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BULLETIN OF
THE BRITISH MUSEUM
(NATURAL HISTORY)
GEOLOGY
VOL. VII
1962-1963
X R U S T E E S OF
THE BRITISH MUSEUM (NATURAL HISTORY)
LONDON : 1964
DATES OF PUBLICATION OF THE PARTS
No. i. isth May ...... 1962
No. 2. 1 5th May .... . 1962
No. 3. 1 4th August ...... 1962
No. 4. i4th August ...... 1962
No. 5. 1 4th August ...... 1962
No. 6. jth December ..... 1962
No. 7. 7th December ..... 1962
No. 8. 8th February ..... 1963
No. 9. gth July ...... 1963
PRINTED IN GREAT BRITAIN
BY THOMAS DE LA RUE &
COMPANY LIMITED LONDON
2 0
CONTENTS
GEOLOGY VOLUME VII
PAGE
No. i. A new Eocene Primate Genus, Cantius, and a Revision of some allied
European Lemuroids. E. L. SIMONS i
No. 2. The Brachiopod Genus Cyclothyris, E. F. OWEN 37
No. 3. The Trilobites of the Caradoc Series in the Cross Fell Inlier of Northern
England. W. T. DEAN 65
No. 4. Fossil Flora of the Drybrook Sandstone in the Forest of Dean,
Gloucestershire. K. M. LELE & J. WALTON 135
No. 5. Fossil Insects from the Lower Lias of Charmouth, Dorset.
F. E. ZEUNER 153
No. 6. The English Cretaceous Turritellidae and Mathildidae (Gastropoda).
H. L. ABBASS 173
No. 7. The Morphology of Tubicaulis africanus sp. nov. a fossil fern from
Tanganyika. H. S. HOLDEN & W. N. CROFT 197
No. 8. The Ordovician Trilobite Faunas of South Shropshire, III. W. T. DEAN 213
No. 9. The Gastropod Genus Thatcheria and its Relationships. A. J. CHARIG 255
Index to Volume VII 299
7V, /t, V.
A NEW EOCENE PRIMATE
GENUS, CANTIUS, AND A
REVISION OF SOME ALLIED
EUROPEAN LEMUROIDS
E. L. SIMONS
BULLETIN OF
THE BRITISH MUSEUM (NATURAL HISTORY)
GEOLOGY Vo1- 7 No. i
LONDON : 1962
A NEW EOCENE PRIMATE GENUS, CANTIUS,
AND A REVISION OF SOME ALLIED
EUROPEAN LEMUROIDS
BY
ELWYN L. SIMONS
Peabody Museum and Department of Geology, Yale University, New Haven, Conn.
Pp. 1-36 ; Pis. 1-3 ; 4 Text-figures
BULLETIN OF
THE BRITISH MUSEUM (NATURAL HISTORY)
GEOLOGY Vol. 7 No. i
LONDON: 1962
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
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within one calendar year.
This paper is Vol. 7, No. i of the Geological
(Palaeontological) series.
Trustees of the British Museum 1962
PRINTED BY ORDER OF THE TRUSTEES OF
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Issued May, 1962 Price Fifteen Shillings
A NEW EOCENE PRIMATE GENUS, CANTIUS,
AND A REVISION OF SOME ALLIED EUROPEAN
LEMUROIDS
By ELWYN L. SIMONS
CONTENTS
Page
I. A NEW EUROPEAN OMOMYID Cantius ...... 3
Taxonomic Revision ........ 5
II. REVISION OF THE SPECIES OF Protoadapis Lemoine ... 8
III. NOTES ON Pronycticebus gaudryi GRANDIDIER .... 14
IV. THE TAXONOMIC POSITION OF Anchomomys STEHLIN . . .21
V. RELATIONSHIPS OF EARLY TERTIARY LEMUR-LIKE PRIMATES . 23
VI. CLASSIFICATION OF EUROPEAN EOCENE PRIMATES ... 27
VII. CONCLUSIONS .......... 34
VIII. ACKNOWLEDGEMENTS ......... 34
IX. REFERENCES .......... 34
SYNOPSIS
Cantius, a new European early Eocene lemur-like Primate genus, is described and additions
to knowledge of anatomy and affinities for three Eocene lemuroid genera, Protoadapis, Pronyc-
ticebus and Anchomomys are made. The significance of the terms " tarsioid " and " lemuroid "
is discussed as these concepts bear on a consideration of the relationships of early Cenozoic
European and American lemur-like Primates to each other and to contemporary tarsier-like
prosimians. The classification of European Eocene Primates is revised from Simpson (1945)
and the conclusions drawn that some European Eocene lemuroids may relate closely to subse-
quent stocks as well as to American prosimian families contemporary with them.
ABBREVIATIONS
A.M.N.H. — American Museum of Natural History, New York.
B.M.N.H.— British Museum (Natural History).
G.P.I.H. — Geological and Paleontological Institute, University of Halle /Wittenburg.
M.C.Z. — Museum of Comparative Zoology, Harvard.
P.U. — Princeton University.
S.M.G.C. — Sedgwick Museum of Geology, Cambridge.
I. A NEW EUROPEAN OMOMYID CANTIUS
A RECENT examination of fossil mammals from the London Clay at the British
Museum (Natural History) resulted in the discovery of an undescribed upper and two
partial lower dentitions of Protoadapis eppsi Cooper (1932). The specimens belong
to a new genus which can be assigned to the prosimian family Omomyidae.
Gazin (1958 : 47) proposed family status for the Omomyinae, which have previously
been ranked as a subfamily of Anaptomorphidae. The latter taxon is now understood
to be a separate stock primarily characterized by reduced dental formula and much
GEOL. 7, i. i§
4 A NEW EOCENE PRIMATE GENUS, CANTIUS
enlarged third and fourth premolars. There are at least two European primate
species, Cantius eppsi (described below) and Teilhardina belgica, which can best be
assigned to the Omomyidae, although each presents some features of difference from
typical omomyids. One might establish a new family or subfamily for these European
species, but until they are better known such a procedure has little to recommend it.1
Higher categories among early Cenozoic Primates have already been too much
multiplied. This may be due principally to the difficulties of grouping species which
are usually known only from dentitions, on rare occasions associated with fragmen-
tary skeletal materials, and to the emotional element involved in the study of human
relatives and ancestors, deservedly criticized by Simpson (1945 : 181) and elsewhere,
and which, apparently, has even pervaded some discussions of fossil prosimians.
Both Cantius and Teilhardina come from deposits containing Sparnacian (early
Eocene) faunas and consequently represent some of the oldest Old World Primates.
They are the earliest, in fact, that could possibly be near the basal stock or ancestry of
any of the surviving European, Asian, or African members of the order.
Evidence as to whether or not Eocene omomyids were more lemur-like or more
tarsier-like is scanty, for skull parts, other than maxillae, are practically unknown
in this group. Hiirzeler (1948) appears to be correct in stressing dental similarities
between the early Eocene primate Teilhardina, from Belgium, and the much better
known necrolemurines. This view is shared by the writer and is also implied by
Gazin (1958 : 92). The Necrolemurinae are quite definitely tarsier-like, and close
correspondence between premolar and molar cusp patterns in unspecialized necro-
lemurines and omomyids reinforces the idea that Omomys and its allies are nearer to
true tarsioids than to such distinctly lemuriform Primates as Adapis or Pronycticebus.
There is a reasonable probability that necrolemurines, North American omomyids,
and even Old World Anthropoidea were derived from a form like the generalized
prototarsioid Teilhardina, which has a lower dental formula of 2.1.4.3., and which
lacks specializations in known parts that could rule out this possibility. Of course,
Teilhardina itself, of Sparnacian Eocene provincial age, occurs too late in time to have
such a position.
A skull fragment of an omomyid, Hemiacodon, discussed and illustrated by Gazin
(1958 : 55, pi. 4, fig. 4) indicates some expansion of the dorso-frontal area in at least
one member of this family — together with reasonably large orbits. Between the
orbital apertures the rostrum, however, is relatively broader than in the probably
omomyid derived genus Necrolemur, being about as in Microchoerus (S.M.G.C. 9669).
In degree of forward rotation the orbits of Hemiacodon appear to be intermediate
between these latter two necrolemurines.
Simpson (1940 : 190-197) discussed a pelvis, some vertebrae, and some elements
of the hind limb which probably belong to Hemiacodon (A.M.N.H. 12613) and con-
cluded that the " skeleton of Hemiacodon, as far as known, is at least as lemuroid as
tarsioid, probably more ". Such judgements ultimately depend on how these two
1 It also seems possible that Periconodon, Lushius and Hoanghonius represent three other Old World
omomyids. Together with broad similarities in known parts, both of these Primates possess an un-
usual cusp on the antero-internal base of the protocone which has been called a pericone. This struc-
ture often occurs in Omomys but is known in very few other members of the order. For discussion of
Luchius, see Chow (1961).
A NEW EOCENE PRIMATE GENUS, CANTIUS 5
concepts are defined. It seems unlikely that the features of the postcranial skeleton in
any Eocene primate would be closely similar to those in a highly specialized modern
form like Tarsius. The earliest tarsioid postcranial distinctions remain elusive.
Segregating out osteological characters of value in determining earlier radiations of
higher categories, or in indicating phyletic relationships, is always difficult and uncer-
tain. Nevertheless, an attempt to do so can be made for Tarsius. In the class of
late and unusual specializations (of uncertain value in determining tarsioid phylogeny)
may be the following characters : (i) Much enlarged orbits with flaring bony rims.
(Occurring also, in a somewhat less exaggerated form, among other primarily
nocturnal animals, such as Nycticebus, Aotes, owls, etc.) (2) Greatly elongated cal-
caneum and astragalus. (This specialization is seen elsewhere in galagos and in an
analogous way in anurans.) (3) Fused tibia and fibula. (Such fusion occurs in a
variety of hopping tetrapods including many rodents, lagomorphs, in some marsupials
and insectivorans, as well as among Aves. See Barnett & Napier (1953 : 12) for
further discussion of the adaptive significance of this feature.) Should these charac-
ters be relatively recent acquisitions among tarsiines, even the direct Eocene ancestor
of Tarsius, in lacking them, would be difficult to distinguish from lemuroids post-
cranially. In view of this possibility, evidence provided by the postcranial bones of
Hemiacodon appears to be equivocal in relating the animal either to Tarsiiformes or
Lemuriformes. What would help to indicate the affinities of such omomyids would
be knowledge of such features as the size and shape of component parts of the brain
(their size relative to each other), extent of facial shortening, proportions of palate
and skull vault, amount of forward shifting of the foramen magnum, and various
other characters of the basicranium, particularly degree of inflation of the auditory
bullae and mastoid region, and situation or relationships of entocarotid circulation,
of the ectotympanic pterygoid alae, and of cranial foramina. Although for omomyids
these cranial features are not known they can be observed in Necrolemur, which in
most of these areas closely resembles Tarsius, and which dentally (at least) shows
affinity with Omomyidae.
As knowledge of the earliest Primates grows with new finds, taxonomic separation,
at all grades of classification, can be expected to become more arbitrary. In this
respect, some European Primates, particularly Cantius, evidence a closer relationship
between Notharctidae and Omomyidae than has previously been documented ;
although this possibility was briefly considered by Gazin (1958 : 47). Perhaps both
of these families, assuming the two groups deserve separate family status, are not
far removed from a common late Paleocene ancestry.
TAXONOMIC REVISION
Genus CANTIUS1 nov.
(PI. i)
TYPE SPECIES. Protoadapis eppsi Cooper, 1932.
GENERIC CHARACTERS. As for the species.
1 From the Latin for County Kent, England (Cantium, Cantia) in which all specimens of this primate
have been discovered and in analogy with such related forms as Washakius, from the North American
Eocene.
6 A NEW EOCENE PRIMATE GENUS, CANTIUS
DISCUSSION. Size : Cantius is a large omomyid, intermediate in size between
Hemiacodon and Ourayia, but distinctly smaller than any of the species of Protoadapis,
and may have been about the size of the living African Giant Galago, Galago crassi-
caudatus.
The discovery of a right maxilla with P3-M3 of this species (PL la) from the early
Eocene deposits at Abbey Wood, Kent (since Cooper's original description) confirms
that the species belongs to a genus distinct from Protoadapis, a conclusion which
can also be demonstrated by features of the lower dentition of Cantius eppsi (B.M.N.H.,
M-I3773) originally reported on by Cooper (1932), and by two other partial lower
dentitions found subsequently (PI. ib). Indeed, Cooper himself suggested the possi-
bility that the British species belonged to a distinct genus. Re-examination of all
available specimens of Protoadapis indicates that this is so.
Some differences between Cantius and Protoadapis (in the lower dentition) are
that the former is somewhat smaller, has paraconids on P4, Mx, M2, a much longer
and more complex M3 talonid, and a considerably smaller and lower P3. A charac-
teristic feature of Protoadapis, sometimes seen in species of the North American
genus Pelycodus as well, is that the P3 stands noticibly higher than P4. In Proto-
adapis the paraconid decreases in expression posteriorly in the molar series but not
in the same way as in Cantius where the paraconid and metaconid are increasingly
drawn together in the series M-^g, compare Pis. ib and 3/. Throughout the lower
molar series of Protoadapis the paraconid remains widely separated from the meta-
conid and in M3 is represented only by a slight crest — still considerably removed
from the metaconid. In Cantius paraconids are clearly delimited in all three molars
and are nearly as high as metaconids.1
In this progressive alteration of trigonid elements posteriorly Protoadapis resembles
closely Notharctus, while Cantius is much more like some specimens of Omomys in
which, although a distinct paraconid is retained throughout the lower molars, the
paraconid is closer to the metaconid in M^_3 than in Mg. Cantius, in its enlarged and
complex M3 talonid, also agrees well with the elaboration of this element in
omomyids. Yet another distinction in the lower dentition between Cantius and
Protoadapis is that in Cantius a clearly defined paraconid cusp is present in the
P4, a structure which never occurs in Protoadapis (see PL 3/, and Stehlin, 1912 :
1282).
Recognition of the Abbey Wood species as generically distinct from Protoadapis
was delayed, in part, because of the scarcity of comparative material in European
museums. The total number of good specimens belonging to all species of both of
these genera is probably less than ten. Nevertheless, the significance of the facts
that Cantius, an omomyid, and Protoadapis, a notharctid-like adapid, occur in the
European Eocene should be stressed. Knowledge of such ranges gives added import
to the more extensively known North American Primates of the omomyid-notharctid
type because it indicates that forms closely affined to them were present in the early
Tertiary of the Old World. The possiblity is thus strengthened that some North
1 Paraconid and metaconid of the M3 in the holotype of C. eppsi cannot be distinguished, either because
of wear or because they are coalesced. In B.M.N.H., M-i^i^ja, b, however, these M3 cusps are separate
but closely approximated.
A NEW EOCENE PRIMATE GENUS, CANTIUS 7
American species may approximate morphological, or even linear stages in the ances-
try of Old World Primates.
The upper dentition from the Abbey Wood locality (PI. la ; B.M.N.H., M-I5I45)—
quite apart from its occluding well with the type lower dentition of Cantius eppsi
(both are of the right side) — shows definite omomyid affinities to about the same
degree as do the lower teeth. Hence there is little reason to doubt that it can be
assigned to Cantius eppsi. Taken as a whole, this upper dentition is close to that of
Omomys and of Teilhardina but is one-half larger than that of either of these small
Primates. P3 and P4 are rather simple teeth with a constriction between the inner
and outer cusps and are similar in conformation to those of Omomys, Teilhardina
and Hemiacodon. P3 has but a single outer and inner cusp and is smaller relative to
P4 than in some specimens of Omomys. Both P3 and P4 are less crenulate than in
Hemiacodon, but exhibit slight enamel wrinkling, particularly on the median slope
of P4 protocone. P4 carries a small cuspule on the anterior slope of the outer cusp
which occurs also in Hemiacodon and Washakius, but apparently not in typical
Omomys. The anterior, median and posterior protocone crests of the P4 of Cantius
are situated about as in Washakius (see Gazin 1958, pi. 9, No. i).
An interesting similarity of Cantius to necrolemurines (and to Tetonius as well) is
that the posterior protocone crest of M1 and M2 turns down toward the posterolingual
part of the basal protocone cingulum, thus breaking the ridge running toward the
metaconule. This forms a so-called " nannopithex-fold ". There is, however, practic-
ally no indication of an incipient hypocone element, other than a slight thickening of
the posterolingual part of the basal protocone cingulum. As in other omomyids the
para- and metaconules are distinct on all three upper molars. These elements are
usually less well denned in notharctids. Also unlike notharctids Cantius lacks any
suggestion of an upper molar mesostyle. Molar para- and metacones are connected
by straight crests on their facing slopes, much as in Omomys and Washakius. Nearly
continuous lingual and labial basal molar cingula in Cantius are also quite like those
of Omomys. Another resemblance to Omomys, to Teilhardina, to some of the more
generalized late Paleocene Primates particularly Navajovius, and also to Tetonius
is the small size of the M3 compared to M1"2. Para- and metastyle elements are
present, as in most omomyids, except on M3 metacone.
The recent identification of an upper dentition of Protoadapis in the collections of
the Geological Institute of the University of Halle /Wittenburg makes possible further
comparative remarks about it and Cantius. Protoadapis, unlike Cantius, shows no
indication in the upper molars of a nannopithex-fold, and the internal cingulum is
typically pronounced and continuous around the lingual base of the protocone, the
hypocone is large, and there is no indication of a metaconule on any of the upper
molars, see PL 3. In these differences from Cantius, Protoadapis more closely resembles
the notharctid Pelycodus.
In conclusion, Cantius has not been found to exhibit any features of resemblance
to notharctids or adapids not occurring in the North American Omomyidae, and in
view of numerous similarities to Omomys and Hemiacodon it is assigned to the latter
family. The dental formula of Cantius is apparently more reduced than in Teilhardina,
but evidence regarding the exact number of antemolar teeth remains inadequate.
8 A NEW EOCENE PRIMATE GENUS, CANTIUS
Cantius eppsi (Cooper)
(PL i)
1932. Protoadapis eppsi Cooper, p. 461, pi. n, figs. 2, 3.
HOLOTYPE. Right ramus of mandible with P3-M3 and partial alveolar borders of
I3-P2, B.M.N.H., M-I3773.
MATERIAL. B.M.N.H., M-I3773 (Holotype), B.M.N.H., M-I5I45, maxilla with
P3-M3; B.M.N.H., M-i5i47#, mandibular fragment with Mj-Mg ; and B.M.N.H.,
M-I5I47&, mandibular fragment with M2_3.
HORIZON AND LOCALITY. Blackheath Beds (Sparnacian) ; Abbey Wood, one and
three-quarter miles east of Woolwich, Kent.1
DESCRIPTION. Lower dentition: Dental formula: 2(?).i.3(?).3. One small
incisor alveolus, followed by a much larger canine alveolus, narrow anteroposteriorly
and broad buccolingually. If only one premolar was present anterior to P3 it may
have been two-rooted. Two-rooted P3 with crown lower than P4 and lacking distinct
para- and metaconids. Para- and metaconids well developed on P4, connected to
protoconid by crests ; well-defined central crest on talonid, running posteriorly along
mid-line of tooth from protoconid and with small posterior cuspule ; P4 not molar-
ized. Trigonid of Mx large, with three well-separated cusps ; paraconid and metaconid
joined to protoconid by low ridges ; talonid much larger than trigonid with distinct
ento- and hypoconid, small hypoconulid present on slope of posterior crest of hypo-
conid. Outline of M2 (crown view) more nearly circular than that of Mx, as in Ourayia,
Hoanghonius, etc. Mg trigonid broad transversely, narrower anteroposteriorly, with
more closely approximated paraconid and metaconid than Mj. Paraconid and
metaconid of M3 trigonid either coalesced as a single cusp (M-I3773) or closely
approximated (M-i5i47# and 6).
Upper dentition : B.M.N.H., M-I5I45 : P3~4 with slight constriction between
protocone and outer cusp. P4 with small anterior cuspule, as in Hemiacodon. M2,
M1, M3 ; no mesostyle or distinct hypocone on upper molars ; antero-internal base
of molar protocone projecting most mesiad ; nannopithex-fold present on posterior
slope of M1 and M2 ; paraconule and metaconule present on M1"3.
II. REVISION OF THE SPECIES OF PROTOADAPIS LEMOINE
History of Study
Preparation of the foregoing section on the British early Eocene primate Cantius
necessitated investigation of all the available material of Protoadapis. It soon became
clear that species of this genus were much in need of revision.
Unfortunately Stehlin's discussion of members of Protoadapis (1912 : 1284-1286)
intruded some taxonomic confusion, as well as a misspelling of the generic name ;
this was largely corrected by Teilhard (1921 : 66, 67, 88-91) but Stehlin's views, and
additional misconceptions of dating quoted by Osborn (1890 : 55) have continued to
1 Dr. G. G. Simpson of Harvard University has suggested in a personal communication that the
species of Hyracotherium from Abbey Wood indicate an earliest Eocene age for this fauna. This is also
indicated by an M3 of Coryphodon from the same locality figured by Cooper (1932 : 459).
A NEW EOCENE PRIMATE GENUS, CANTIUS
Measurements (in mm.) of Cantius eppsi (Cooper)
(All specimens in the British Museum of Natural History)
Mandible :
Depth of jaw beneath M2
Anteroposterior length of P3-M3
Anteroposterior length of Mx_3
Anteroposterior length of M2~3
Anteroposterior diameter of P3
Transverse diameter
(trigonid)
(talonid)
M2
M,
M2
M3
M!
M2
M,
Maxilla : M-I5I45
Anteroposterior diameter of P3
Anteroposterior diameter of P3
P4
M1
M2
M3
M-I3773
6-7
17-2
11-4
8-1
2-8
3'3
3-6
3-6
4'5
1-8
2-3
2-5
3'3
3'0
2-8
3'i
2-4
15-0
3-6
3'7
8-0
3-6
3'7
4'3
2-3
3-0
2-9
2-8
3'0
2'4
(Transverse
diameter)
3'2
3-8
4'7
5'4
4'3
3-8
4'4
affect the literature on this primate. These errors were largely reproduced by Hill
(1953 : 482-487).
Further complications derive from the observation, made during a recent visit to
the Halle /Wittenburg Geological and Paleontological Institute, that the holotypes
of two Eocene Primates, " Europolemur " klatti and " Megatarsius obeli " described
by Weigelt (1933) can be assigned with confidence to Protoadapis. As it was not
possible for Weigelt (1933) to make direct comparisons with previously described
fossil Primates some specific and generic assignments which might have been recog-
nized for the Geiseltal species were not noted by him. In fact, at that time Weigelt
probably could not have determined his two new genera as upper dentitions and skulls
of Protoadapis, since the associated upper and lower dentition, G.P.I.H. 4310,
discussed below, had not then been discovered.
The assumption that species of this genus occur in the French Paleocene of Cernay
and Rilly dates from an erroneous citation in Lemoine & Aumonier (1880 : 611) as
stated by Teilhard (1921). * However, this misapprehension was repeated in Hill
1 Lemoine & Aumonier list a species of this genus, P. copei, as occurring in the Cernay Paleocene.
It was not figured or described and is now lost, probably because Lemoine later referred the specimen to
a different genus and species, not Protoadapis.
io A NEW EOCENE PRIMATE GENUS, CANTIUS
(1953 : 483). Lemoine later stated (1891 : 28) that this genus is restricted to the
Eocene. D. E. Russell who is currently revising the French Paleocene faunas,
informs me that it does not occur at Cernay. The recognition of several specimens
of Protoadapis from the Middle Eocene Brown Coals of the Geiseltal, Germany,
now in the collections at Halle, serves to date the earlier members of this genus
better than the rather scanty and poorly associated Cuisian fauna with which the
French specimens of Protoadapis curvicuspidens and " Protoadapis recticuspidens "
were recovered.
A further error in the literature on this genus is its recorded occurrence in the
Belgian Eocene deposits (Hill, 1953:483). There is no real evidence for such a
record, but a specimen of Protoadapis from the Quercy phosphorites of south-central
France was, at the time of Teilhard's writing, in a private collection at Louvain and
his reference to it may have led to this confusion of range.
All the materials of this genus discussed by Lemoine, Stehlin, and Teilhard, as
well as the specimens at Halle, belong (after the removal of " Protoadapis " eppsi)
to no more than three species. Protoadapis recticuspidens Lemoine, 1878, is based
(in spite of published remarks to the contrary) on a single specimen in which the
teeth are so worn that only a few characters of M3 can be determined. This tooth,
and the mandible, now in the Paris Museum, are distinctly smaller than are those of
Protoadapis curvicuspidens (here designated as the type species of the genus) and it
may not belong to Protoadapis. I would prefer to regard Protoadapis recticuspidens
as a nomen vanum.
As Teilhard stressed, the specimen called Protiadapis recticuspidens by Stehlin
(1912, fig. 278) is actually P. curvicuspidens, the holotype of which is now in the
Museum National d'Histoire Naturelle in Paris. It is on the basis of this specimen
that the genus Protoadapis was first defined.
TAXONOMIC REVISION
Genus PROTOADAPIS Lemoine
(Text-fig, i)
TYPE SPECIES. Protoadapis curvicuspidens Lemoine (1878). (Unnumbered holo-
type and two other specimens in National Museum of Natural History, Paris.)
AMENDED DIAGNOSIS. Size : somewhat larger than Adapis parisiensis ; lower
dental formula : i?.i.3.3. incisors missing in type but one or more alveoli appear
to be present in other specimens ; canine large (alveolus) ; Pt absent, replaced by
diastema ; P2 short with single pointed cusp, and with two obliquely situated roots,
the anterior located more buccally. P3 distinctly larger, much higher and more pointed
than P2 with nearly vertical anterior margin, no distinct paraconid or metaconid ;
P4 lower than P3 (unlike Cantius) lacking paraconid, but with well-defined metaconid
and talonid possessing central ridge ; molar paraconids reduced, but extending
much more lingually than in Adapis, less so than in Cantius. Mj and M^ with small
hypoconulid on posterior hypoconid crest ; M3 hypoconulid making up a distinct
lobe ; length from P4 to M3 — 22 mm., depth of horizontal ramus beneath M2 from
io to 12 mm.
A NEW EOCENE PRIMATE GENUS, CANTIUS n
DISCUSSION. Two French species of this genus are accepted here as valid, Proto-
adapis curvicuspidens collected in Paris Basin deposits, perhaps of Cuisian age, and
Protoadapis angustidens (Filhol, 1888) from the Quercy phosphorites (see Text-fig, i).
Protoadapis brachyrhynchus Stehlin (1912) is clearly a synonym of the latter species,
as Teilhard (1921 : 97) concluded. A third species, Protoadapis raabi, from the
Middle Eocene Brown Coals of the Geiseltal, Germany, is slightly smaller than
COMPARISONS OF MANDIBLES OF Protoadapis species
A (ALL X 2 APPROX., INTERNAL ASPECT)
Diagram of Protoadapis angustidens from
ilhard (1921, pl.4. fig. 12 reversed)
Protoadapis curvicuspidens from Teilhard
(1921, pl.3, fig.5)
GEOL. 7, I.
Protoadapis curvicuspidens from Stehlin
(1912, fig. 288)
FIG. i. Species of Protoadapis Lemoine from French deposits.
i§§
12 A NEW EOCENE PRIMATE GENUS, CANTIUS
P. curvicuspidens but does not differ greatly from it otherwise, in so far as these
two species can be compared. Better materials may prove that the latter two
" species " intergrade but pending such an eventuality, it is advisable to retain
a separate species for the Geiseltal finds.
SPECIFIC DIAGNOSES. Teilhard (1921) has published full descriptions of the two
species of this genus from the French Eocene and consequently they will not be
separately diagnosed here.1
Protoadapis klatti (Weigelt)
(Pis. 2, 3)
1933. Europolemur klatti Weigelt, p. 123, pi. 4, fig. 5 ; pi. 6, fig. 4.
1933. Megatarsius obeli Weigelt, p. 141, pi. 4, fig. 4 ; pi. 6, figs. 1-3.
MATERIAL. In addition to the holotypes of " Europolemur " klatti and " Mega-
tarsius obeli " (No. 4234) ; numbers 4238, 4258, 4280, 4292 and 4310 of the collections
of the Geological and Paleontological Institute of the University of Halle /Wittenburg
are referable to this species. Some of these specimens were regarded by Heller (1930)
as belonging to Adapis, but this genus does not occur in the Geiseltal fauna.
DIAGNOSIS. A medium-sized primate — apparently about the size of Pronycticebus
2 . 1 . "3 . "3
or the Recent Potto, Perodicticus. Dental formula : .r-1— — '- — ; anterior lower incisor
?l. 1.3.3
not known (if present), I3 procumbent and spatulate ; C long and pointed with
slightly developed posterior shelf — appearance about as in Notharctus, £ followed by
diastema ; P2 two-rooted with simple pointed crown and posterior heel, lacking
metaconid and protoconid ; P3 much higher than P2 and P4 and lacking protoconid ;
P4-M3 as in Protoadapis curvicuspidens but smaller, Mx and M2 with distinct hypo-
conulid on posterior slope of hypocone, M3 hypoconulid and entoconid large.
I2 larger than I3, spatulate with crown anteromedially directed and somewhat
procumbent ; I3 small and simple, may be separated from C by a diastema ; C
much larger and longer than in Adapis, furrowed along its external face by anterior
and posterior vertical grooves and followed by a diastema ; P2 less than half as
large as P3, and with small protocone (two rooted) ; P3 and P4 with single ectoloph
and large protocone ; M1 lacking mesostyle and metaconule, but with distinct para-
conule and prominent internal and external cingula, internal cingulum with large
hypocone, basal cingulum sometimes incomplete across internal face of protocone ;
M2 slightly larger than M1 with more continuous lingual cingulum and larger hypo-
cone ; M3 smaller than JVF-M2, lacking a distinct hypocone, but with pronounced
internal cingulum.
DISCUSSION. " Two " undescribed specimens in the Halle collection form a valuable
basis for the clarification of the dental structure of Protoadapis. One of these G.P.I.H.
4310, from the Leonhardt Coal Mine (the locality of " Europolemur ") includes the
left P4 through M3, the right M3 an associated right lower canine and M1"3 (M1"2
somewhat damaged) see PI. ^e,f. The second specimen, G.P.I.H. 4258 (PI. 2), crushed
facial region with attached lower jaw fragment of the left side with I2 through P3
1 Protoadapis angustidens is larger and occurs later in time than P. curvicuspidens.
A NEW EOCENE PRIMATE GENUS, CANTIUS 13
and I3 through P3 was also collected from Leonhardt Mine. The specimen card of
No. 4310 states that these teeth probably belong to the same individual as No. 4258,
and this association is reinforced by similarity of colour, wear, matrix, and the fact
that the two specimens do not share any overlapping parts, which would rule out
such a possibility. In any event, both are referable to Protoadapis and together
they preserve almost the entire dentition intact. Dental comparisons indicate that
the skull of " Europolemur " klatti belongs to the same species as the foregoing
specimens and is therefore a synonym of Protoadapis. The holotype of " E." klatti
(apparently lost during the last war) has upper molars of the same size and conforma-
tion as those of G.P.I.H. 4310 (judging from Weigelt's published information) and
in so far as these two finds can be compared, there is no basis for a specific distinction.
Teilhard (1921, pi. 3, fig. 6) illustrated an upper left molar which he suggested might
be of P. curvicuspidens ; this is confirmed by the Halle specimens. Because of crush-
ing, measurements on Brown Coal fossil Primates, other than on teeth, are not very
reliable and thus have reduced value when drawing taxonomic distinctions. However,
the length of P4-M3 series of Protoadapis klatti is about 15 mm. in the type and the
length of P4-M3 in G.P.I.H. 4310 is about 17*2 mm. Similar measurements for
Cantius eppsi are 14' 2 and 15*1 mm. respectively.
Regarding the teeth of " Megatarsius obeli " from the Cecilia Coal Mine (locality
Leichenfeld II), as Weigelt noted (1933 : 142) :
" die Beschaffenheit der Kronen 1st
eigenartig stumpf , als wenn die oberste
Schimelzchicht fehlte . . . ".
There is no doubt that this condition is due to post-mortem chemical absorption
of the enamel and some of the dentine of the canine and P4 through M3 of the right
side (all of the dentition that remains in " Megatarsius "). This sort of erosion of
fossil teeth is quite common in early Cenozoic mammals. Neither Heller (1930) J nor
Weigelt (1933) appear to have taken into account the fact that many of the Geiseltal
Primates have lost almost all significant dental characters in this way. Such erosion
also explains the extraordinary appearance of the canine of " Megatarsius " figured
by Weigelt (1933, pi. 6, fig. i). In this tooth, the enamel and much of the dentine of
the crown has been removed and there has also been erosion around the base of the
tooth. The odd basal cingulum and the whole appearance of the tooth is unreal
misleading. In " Megatarsius ", G.P.I.H. 4234, the upper dental formula is the same
as in G.P.I.H. 4258-5310, Protoadapis. Furthermore, the eroded crowns of M1
and M2 show metaconule and mesostyle absent, but paraconule present, and allowing
for the reduction in size caused by enamel erosion, M1"3 of " Megatarsius " are of
appropriate size for Protoadapis klatti. For the writer, no doubt remains that
" Megatarsius " is referable to P. klatti. In any case, G.P.I.H. 4234 is not adequate
as a type specimen, and if not Protoadapis would be a nomen vanum.
Weigelt's discussion of the skulls of " Europolemur " and " Megatarsius " describes
adequately these much damaged specimens. In overall morphology the skull of
Protoadapis is more like that of Pronycticebus than of Adapis. With Pronycticebus,
1 Heller's holotype of Adapis minimus is an indeterminate specimen for the same reason, all of the
enamel having been removed by chemical solution.
i4 A NEW EOCENE PRIMATE GENUS, CANTIUS
it shares a comparatively shorter rostrum, larger orbits, more slender zygoma, P4
with a single outer cusp and two-rooted P2, but differs in having lost P i/i. " Mega-
tarsius " has a broad interorbital septum about i cm. wide and resembles that of
Pronycticebus in the disposition of the medial end of the supraorbital margin. On
the antero-dorsal face of this skull between the temporal crests is a broad and slightly
depressed area. The temporal ridges converge to the mid-line about 2 '25 cm. behind
the posterior end of the nasals. Conformation and extent of the nasals can be deter-
mined in G.P.I. H. 4234. These bones are slightly expanded posteriorly as in Pronycti-
cebus. The orbital region is much crushed and, consequently, the lacrymal foramen
cannot be located, but fragments suggest that the post-orbital bar was more slender
than in Adapis.
Relationships of Protoadapis
Now that both the upper and lower dentitions and something of the skull are
known in Protoadapis, it takes a more significant position in early Primate history.
Placement of this genus as a typical lemuroid primate seems beyond doubt, and in
fact it is the only Old World early Cenozoic form which has the same dental formula
as any of the living lemuroids, although it lacks the specialized tooth comb below
which is typical of the latter taxon. Being older, and yet having a more reduced
dental formula, it cannot be near the lines which gave rise to such Eocene lemuroids
as Adapis, Pronycticebus and Notharctus. Perhaps its greatest similarities are with
Pelycodus and Pronycticebus which, however, retain P I/I. In Pelycodus the manner
of hypocone formation may also be different. Some specimens of Pelycodus, for
instance A.M.N.H. 15022 (see Gregory, 1920, pi. 35), appear to have an incipient
hypocone on the basal cingulum and at the same time another " pseudohypocone "
developing from the nannopithex-fold on the posterior slope of the protocone. In
the line leading to Notharctus the hypocone derived from the basal cingulum is
suppressed, while in Protoadapis the nannopithex-fold is absent. The nearest known
upper molar morphology that could give rise to both these patterns is that of Cantius
but this form is too late to be ancestral to Pelycodus and its dentition too generalized
for classification with the adapids. As suggested by Teilhard (1921), Simpson (1940),
and others, Protoadapis shares with the North American notharctids many of the
features which distinguish the latter group from the Adapidae. The upper dentitions
and the anterior teeth preserved at Halle add somewhat to this conclusion, but
in several respects this primate tends to link adapids and notharctids as Cantius
links the omomyids with the latter families. In view of dental similarities with
Caenopithecus and Pronycticebus, together with its European provenance, Protoadapis
is here referred to the Adapidae.
III. NOTES ON PRONYCTICEBUS GAUDRYI GRANDIDIER
(Text-fig. 2)
The unique skull and mandibular fragment of Pronycticebus was discovered in
1893 and reported, in the original studies of Grandidier (1904, 1905), to be of Bar-
tonian, late Eocene age, from Memerlein-le-Quercy in south-central France. The
Abbe Rene Lavocat has suggested in a personal communication to the writer that
A NEW EOCENE PRIMATE GENUS, CANTIUS 15
judging from his studies of mammals of Oligocene provenance, there is a similarity
in the colour of bone and matrix of this specimen to those of the European Oligocene.
This interesting observation implies, at least, that Pronycticebus may be from the
latest levels of the Quercy caves. If so, it could be of Stampian Oligocene provincial
age.
Pronycticebus has long been a difficult form to assess taxonomically, due in part to
the lack of specialization of the teeth and the presumed absence of a post-orbital bar.
Grandidier originally suggested affinities with the living lorisiform Primate Nycticebus,
hence its generic name. This view has some merit. Subsequently Gregory (1920)
and others held that the species has tarsioid affinities. This conclusion appears to
have been based largely on the brachycephaly of the skull (considerably more pro-
nounced that in Adapis or Notharctus) and which, at the time of Gregory's writing,
was rivalled in degree, among Eocene Primates, only by Necrolemur skulls (also
from Quercy) and by a fragmentary skull, lacking dentition, from the Middle Eocene
of North America described by Granger & Gregory (1917) as " Aphanolemur gibbosus ".
Gazin, in a revision of the Middle and Upper Eocene Primates of North America
(1958), has shown that Aphanolemur is a synonym of Smilodectes. Several specimens
figured by him indicate that expansion of the brain case in the latter primate (appar-
ently closely related to Notharctus) has reached about the same level as in Pronycti-
cebus. Skulls of Smilodectes, and to a lesser extent of Notharctus, demonstrate,
therefore, that the degree of brain expansion which occurred in Pronycticebus was
not unique among Eocene lemuroids. Protoadapis, judging from the crushed skulls
at Halle, also had a rather large brain.
On other grounds, Le Gros Clark's careful study (1934) of the cranial anatomy of
Pronycticebus has already demonstrated that this genus is not a tarsioid, a position
recently re-affirmed by Piveteau (1957 : 55, 56).
Re-examination of the Evidence
Orbital Region
It is, perhaps, an amusing commentary on the nature of appraisal of fossil speci-
mens that the zygomatic arch of the one known skull of Pronycticebus can be said
to have been " evolving " almost continuously since Grandidier 's original studies on
this primate in 1904 and 1905. Apparently some time before the original description,
the zygomatic arch was buried in a thick layer of plaster of Paris, which closely
resembles the colour of the original bone. In 1934 Le Gros Clark remarked that the
zygoma was not quite as broad as is indicated in Grandidier's illustrations and
figured a narrower arch. Piveteau's plate (1957 : 61) shows a still more slender arch.
In the same year, the writer was able to remove most of the remaining plaster which
exposed a yet more gracile arch and the very distinct base of a post-orbital process
(Text-fig. 2). Furthermore, the region of the frontal attachment of the post-orbital
bar shows distinct fractures on both sides of the skull, where the bar has been broken
off. The cross-sectional extent of this area is shown by hatching in Text-fig. 2. One
can clearly distinguish the broken surface, and although this area of attachment is
not very extensive in the vertical dimension, this correlates well the slender zygoma,
16
A NEW EOCENE PRIMATE GENUS, CANTIUS
FIG. 2. Dorsal and lateral views of the skull of Pronycticebus gaudryi (Holotype),
Hatching indicates broken surfaces, stipple missing parts. In dorsal view, right side
restored from left.
A NEW EOCENE PRIMATE GENUS, CANTIUS 17
found under the plaster, and with the much enlarged orbit indicated by the remaining
median part of the orbital margin.1 Living and fossil Primates with comparatively
large orbits often have a slender post-orbital bar and a thin and flaring zygoma, as
in Galago and Necrolemur. In fact, in some Galago species, the temporal base of the
post-orbital bar in cross-sectional area is less extensive (when compared to total
skull size) than is this area in Pronycticebus.
In view of the observed presence in Pronycticebus of both a frontal and a zygomatic
base for the post-orbital bar, the large brain compared to Adapis, the complete
absence of any evidence that other primates of the grade of advance seen in Pronycti-
cebus lack the post-orbital bar, and the suggestion that enlarged orbits do sometimes
correlate with relatively small frontal areas of attachment for the bar, no basis now
remains for thinking that the bar was not continuous. Text-fig. 2 presents a conjec-
tural restoration of its position.
Le Gros Clark (1934 : 20-27) noted most of the observable characters in the skull
of Pronycticebus and discussed occurrences of many of these features in other Primates.
The numerous new fossil primate specimens, which have been discovered or restudied
since that time justify further comparison of some of these structures. Contrary to
the current assumption Pronycticebus gaudryi is not a conservative form. With
existence of a post-orbital bar in this species established, little remains to be seen in
it that is particularly primitive for an Eocene primate. In fact, the expansion of the
brain case, the large, forward-directed eyes and the short rostrum are all features
that have been considered " advanced ".
Location of the lachrymal foramen, together with part of the lachrymal bone,
outside the orbit in Pronycticebus was suggested as a difference from Adapis by Le
Gros Clark (1934). However, a specimen of Adapis magnus, P U. 11481, indicates
that in this species the position of the foramen and forward extension of the lachrymal
may sometimes be about as in Pronycticebus. In Malagasy lemurs the lachrymal
foramen typically lies further outside the orbit than in the foregoing, while in living
lorisines, and the potto in particular, the position of foramen and lachrymal is about
as in Pronycticebus, except for the smaller size of the lachrymal itself. Necrolemur
(M.C.Z. 1179) and Tarsius have also a facial location of this foramen, but in Smilodectes ,
Notharctus and most platyrrhines this foramen clearly lies within the anterior orbital
margin, as in pongids and man. In this character it would seem that Adapis and
Pronycticebus approach more closely the living lemuroids than they do members of
the Notharctidae and Anthropoidea. Forsyth Major (1901 : 151) concluded that the
primitive condition for Primates is to lack a great facial expansion of the lachrymal.
Shape and position of the foramen and lachrymal in Pronycticebus appear to be
equivocal in relating it either to living lemurids or lorisids.
Rostrum
The comparatively foreshortened face of this primate is evident from the parts
preserved. A vertical position of the premaxillo-maxillary suture (as well as the
anterior recurving of the alveolar border around the canines) indicates that the missing
1Relative to skull size Pronycticebus has larger orbits than most, if not all, other early Tertiary Primates.
This may indicate nocturnal habits.
i8 A NEW EOCENE PRIMATE GENUS, CANTIUS
anterior tip of the skull can hardly have been much longer than the conjectural
reconstruction shown in Text-fig. 2. The sharp constriction in transverse diameter
of the rostrum immediately posterior to the canines in Pronycticebus is not as distinct
in living lemurs and is absent in Adapis. However, this feature can be seen in Loris
and Nycticebus, and is one of the reasons why this skull is reminiscent of that of
Nycticebus coucang.
Dorsal Aspect of Skull
Cleaning of the skull has revealed sutures on the top of the brain case more clearly,
and allows further comparison of this aspect of the skull in Pronycticebus with that
of Adapis. Unlike Adapis, in Pronycticebus the posterior margins of the f rentals
extend on to the anterior part of the brain case. A sinus canal, near the posterior
extremity of the parietal, is single in Pronycticebus, but in Adapis parisiensis it is
multiple, as noted by Le Gros Clark (1934 : 22) and indicated by Stehlin's figures
(1912). A large size for this canal has been said to be a primitive feature. After
cleaning, sutures can be determined in Pronycticebus delimiting paired interparietals
and (posterior to the sinus canal) small accessory ossicles (see Text-fig. 2). These
bones have not been reported as occurring in Adapis. Taken as a whole the dorsal
aspect of the brain case of Adapis does not closely resemble that of Pronycticebus.
Such distinctions suggest that by the Middle or Late Eocene (the exact time range
for both forms is uncertain) these two lemuroids had already diverged considerably.
Such an assumption is reinforced by distinctions in other skull components, discussed
above, such as the different shape of the muzzle, divergent character of the dentition,
and difference in position and size of the orbits in these two Primates of the Quercy
phosphorites. At present, it is probably better to retain them in the same family,
as has been done by Hill (1953) and Piveteau (1957). Nevertheless, Pronycticebus
and Adapis are as distinct morphologically as forms from the North American Eocene
currently placed in different families, for example the notharctid, Pelycodus, and
omomyid, Ourayia see Simons (1961^:5).
Apart from the construction of the inside of the auditory bulla the basicranium of
Pronycticebus was fully described by Le Gros Clark (1934). It will not be reconsidered
in detail here. A possible alternative for an identification made by him (1934 : fig. 3)
is that the foramen indicated as the hypoglossal canal may be the inferior petrous
sinus.
On obtaining permission to examine the interior of the auditory bulla it was
determined that the ectotympanic is not tubular, thus completely confirming, in this
regard, Le Gros Clark's thesis that this primate is not a tarsioid. It is noteworthy
that the annulus lies very near the external auditory meatus and, unless it has been
displaced, appears to be fused with the bulla wall in its posterior third, just internal
to the posterior rim of the external meatus. From this point the annulus arches
inward and downward, so that its ascending anterior limb lies some distance from
the antero-lateral wall of the bulla. Near the ventral extremity of the ring (and
external to it) is a shelf on the lateral bulla wall for the support of the annulus
membrane. Compared to that of Adapis this shelf is quite small. Hill (1953 : 113)
figures a dissection of the auditory bulla of Loris tardigradus which indicates only
A NEW EOCENE PRIMATE GENUS, CANTIUS 19
slightly greater reduction of the annulus. The position of the tympanic ring in
Pronycticebus, close to the external meatus, is an additional indication that Grandidier
may have been nearer the truth than has sometimes been thought, when he suggested
an affinity between this species and the modern lorises.
Dentition
It was principally because of the somewhat enlarged brain and generalized charac-
ters of the dentition (resembling teeth of Anchomomys a supposed tarsioid), that
such students as Stehlin (1916 : 1422), Gregory (1922) and Abel (1931 : 186) referred
this primate to the tarsioids. Founding his conclusions principally on characters of
the skull Le Gros Clark demonstrated that this primate cannot be considered a
tarsioid. However, it may not have been sufficiently stressed that the dentition also,
although unspecialized, is not Tarsius-like. Some principal reasons why this dentition
does not indicate tarsioid relationships are as follows :
The cross-sectional area of the upper canine base is considerably greater relative
to adjacent teeth than it is in Tarsius while in necrolemurines the upper canine is
even smaller proportionately than in Tarsius. Tarsiids, with the possible exception
of Nannopithex, lack the P1 present in Pronycticebus. The latter primate, like Adapis,
Protoadapis, Anchomomys and Progalago, has two-rooted second premolars above
and below, while in tarsiines, these teeth are always single-rooted. In configuration
of the remaining teeth Pronycticebus closely resembles Protoadapis, but differs from
it dentally in having a less well-developed lingual cingulum on the upper molar
protocone, and a more reduced M2_3 paraconid. However, these are rather slight
distinctions, known to be variable in other primate species.1 Otherwise, these two
forms agree in such features as the absence of mesostyles, and of a distinct hypocone
on M3 (although it is well developed on M1 and M2). Also, they are closely similar in
proportion and size. It is just possible that better specimens might even show that
species of these two types of Primates are not separable generically. Unlike tarsioids,
Pronycticebus has, as Simpson (1940 : 202) remarks (with reference to Caenopithecus)
" open trigonids and the peculiar mode of paraconid reduction so characteristic of
Adapis and its close allies and unknown among any forms of really probable tarsioid
affinities ".
Cranial Proportions
A logarithm of ratio diagram (Text-fig. 3) indicates graphically the close similarity
in linear proportions between Pronycticebus and Nycticebus and in contrast to those
of an early Cenozoic lemuroid species Adapis magnus? and a Recent Malagasy lemur.2
1 Even in Protoadapis, as is indicated by isolated upper molars from the Geiseltal coals at Halle, the
lingual protocone cingulum of the upper molars is sometimes incomplete, as it also is in Pronycticebus.
2 Measurements indicated in Text-fig. 4 are as follows : (i) Transverse diameter from mid-line of
skull to greatest flare of zygomatic arch. (2) Length of skull, from posterior margin of canine to anterior
margin of foramen magnum. (3) Length of skull from posterior margin of canine to posterior
extremity of skull. (4) Length of dentition from anterior face of canine to posterior edge of M3.
(5) Greatest transverse width of dentition. (6) Greatest transverse diameter of the cranium. (7) Least
post-orbital diameter of skull. (8) Length from posterior margin of the canine to end of hard palate.
(9) Length from anterior margin of canine to point of least post-orbital diameter. (10) Length from
point of least post-orbital diameter to posterior tip of skull, (n) Greatest diameter of orbit from base
of post-orbital bar to opposite rostral orbital margin. (12) Greatest transverse diameter across auditory
bullae. (13) Greatest transverse diameter across canines. (14) Least transverse diameter of muzzle
across upper second premolars.
20 A NEW EOCENE PRIMATE GENUS, CANTIUS
The major differences between Nycticebus and Pronycticebus are in plots 4, 7, and 8.
Measurement number 4, the length of the upper teeth might be expected to be less
in Nycticebus as its dental formula is more reduced, and the same would apply to
measurement number 8, the approximate length ol the hard palate. Measurement
Pronycticebus
Nycticebus
2
3
4
5
6
7
8
9
10
II
12
13
14
L
-.40
Ada pis (standard)
fLemur
\
\
v
*"^"**^w&
\.
I I I I I I I I I I — I — I — I — I — I — I — I
-.35 -.30 -.25 r20 -.15 -,IO-:05 0 +.05 +.10 +.15 +.20 +.25 +.30 +.35 +.40 +.45
FIG. 3. Logarithm of ratio diagram comparing cranial proportions in four prosimians ;
Adapis (standard). Mean, and extremes of range are indicated for a sample of ten
Nycticebus coucang.
number 7, the least post-orbital diameter of the skull (much smaller in Pronycticebus)
is a primitive feature, correlated with a comparatively unexpanded brain, and is
common to all Eocene lemuroids. In the remaining measurements Pronycticebus
falls within (or very close to) the range of size variation of a small sample of the living
lorisid Nycticebus coucang (10 individuals) .
CONCLUSIONS
Perhaps all the above-mentioned similarities are due to parallelism produced by
similar adaptations in Pronycticebus and lorises, but such a view appears to be rather
A NEW EOCENE PRIMATE GENUS, CANTIUS 21
strained. A more likely conclusion is that Pronycticebus does have some relationship
to the radiation which produced extant Lorisiformes. Nevertheless, because of many
primitive structures also shared with the contemporary Adapis and Protoadapis it
seems best not to remove this genus from the Adapidae, at present.
IV. THE TAXONOMIC POSITION OF ANCHOMOMYS STEHLIN (1916)
(Text-fig. 4)
This rather small primate, originally described by Stehlin (1916) occurs in deposits
ranging from early Lutetian to early Ludian age in Europe. Four species have been
proposed, differing little in known parts except for size. Taxonomic placement of this
genus has varied considerably. Abel (1931 : 175) and others considered it a tarsioid,
but Simpson (1940 : 202) quite correctly pointed out that such an assignment was
most improbable. Of European Eocene lemuriforms the closest dental resemblances
to Anchomomys are to be seen in Pronycticebus and Caenopithecus, but because of a
common simplicity in cusp pattern their similarities can, perhaps, be over-emphasized.
In some respects these three genera also resemble Adapis, particularly in type of
paraconid reduction, but all three differ from Adapis in having M3 hypoconulid
much more distinctly set off as a separate lobe from the hypoconid. They also lack
the bicusped ectoloph of P4 and continuous lingual upper molar cingula seen in
Adapis.
For Anchomomys the closest dental resemblances appear to be with a Miocene
lorisiform of Kenya, Progalago. In fact, in spite of the differences in time and place
of occurrence of the species belonging to these two genera, some of the European
Eocene forms, particularly Anchomomys quercyi, can hardly be distinguished generic-
cally, on the basis of parts preserved, from Progalago dome. Species of these genera
also overlap as far as size is concerned and similarities in the lower molars are particu-
larly striking. Both exhibit reduced paraconids, well-developed entoconids, a large
M3 hypoconulid, with remaining molar cusps of closely corresponding shape and
situation relative to one another. Both these prosimians also display in the major
molar cusps a fairly distinctive angular or pyramidal form.
The upper molars of Anchomomys are quite like those of Progalago although in
some, but not all, species of Anchomomys the paraconule and metaconule are more
distinctly developed. In Progalago dorae and Anchomomys quercyi the latter two
cusps are absent, see Text-fig. 4. These two species are alike in having a subquadrate
M1 with large hypocone, more triangular M2 with smaller hypocone and lack a
distinct M3 hypocone, see Text-fig. 4 ; Le Gros Clark (1956, pi. i) and Stehlin (1916,
figs. 327, 328, 332). Moreover, at least one specimen of Progalago dorae (Le Gros
Clark & Thomas, 1952, pi. 3, fig. 9) shows that in this species the P2 was a much
elongated and two-rooted tooth. If the same applies to the maxilla of Anchomomys
quercyi figured by Stehlin, and shape of the anterior alveoli strongly indicates this
possibility, then the upper dental formulae of both species are identical, at least from
the canine posteriorly.
Perhaps the best preserved skull of Anchomomys is that of the unnumbered
holotype of Anchomomys latidens (now in Paris) described by Teilhard (1921 : 13}
22 A NEW EOCENE PRIMATE GENUS, CANTIUS
from the Quercy phosphorites and identified as " Rossignol Collection 1893 : n ".
The specimen is much broken and distorted, but careful preparation could probably
expose more of the cranial anatomy. As in Pronycticebus and Progalago there is a
single large infra-orbital foramen above P4. The anterior root of the zygoma appears
to arise about M2, and through this anterior root opens to the rear a relatively large
posterior infra-orbital canal. On the antero-dorsal surface posteriorly converging
FIG. 4. — Comparison of upper dentition in Anchomomys and Progalago. A. Progalago
sp.. [Diagram after Le Gros Clark, 1956, pi. 4, fig. i.] Right maxilla, reversed,
xy-5 approx. B. Anchomomys quercyi. [After Stehlin, 1916, fig. 332.] XJ'5
approx.
temporal ridges are evident, but the dorsal insertion of the post-orbital bar cannot
be distinguished. Between these temporal ridges, there is a broad depression, appear-
ing rather as does this area in Pronycticebus. An apparently distinctive feature of
Anchomomys latidens is the character of the surface of the temporal bone which is
covered by numerous minute foramina, a condition certainly not to be seen (to
this extent, if at all) in Adapis or Necrolemur. At the back of this skull are well-
developed nuchal crests. These are typical of Eocene Primates and their presence
is consistent with the retention of similar strong ridges in this position in Progalago.
A NEW EOCENE PRIMATE GENUS, CANTIUS 23
Taxonomic Position
Stehlin (1912 : 1426) tended to favour the idea that Pronycticebus and Anchomomys
have about the same relationship to the Galagidae [" Nycticebiden "] as Adapis
has to the lemuriform lemurs, but remarked that until the position of the carotid
foramen and situation of the extotympanic was revealed for one of these Primates
their taxonomic position would remain uncertain. Now that the annulus of Pronycti-
cebus is known to show a possible foreshadowing of the condition seen in lorisiform
Primates, Stehlin's analysis is strengthened. The subsequent placement of these
two genera among the tarsioids, proposed for Anchomomys by Teilhard (1921),,
and for both Pronycticebus and Anchomomys by Gregory (1922) and Abel (1931)
can now be seen to have marked a step backward in determination of their relation-
ship to other prosimians.
Teilhard, although stressing tarsioid relationships for Anchomomys latidens, did
however remark (1921 : 16) : " Tout au plus pourrait-on le rapprocher des Galago
pour le developpment de 1'hypocone et la tendance a la molarisation de P 4 . . . ".
This observation is important taxonomically because the fourth upper premolar in
species of the genera Anchomomys, Progalago and Galago has an interesting similarity
in basic plan, in addition to the other features of resemblance between these genera
mentioned above. The view that these forms were tarsioids was rejected by Simpson
(1940) after Le Gros Clark's study of the cranial anatomy of Pronycticebus, but since
then their position among the non-tarsioid Prosimii has remained uncertain.
The very close dental relationship between Anchomomys and Progalago, and the
similarities which the latter in turn has with living lorisoids reinforce the idea that
this phylum can now be traced back with a fair degree of accuracy as far as the
European Middle Eocene.
V. RELATIONSHIPS OF EARLY TERTIARY LEMUROID PRIMATES
The Concept of " Lemur vid " Primates
One objective of the research reported here and in Simons (1961) has been to
determine whether tarsier-like and lemur-like Primates can be distinguished in the
early Cenozoic. For at least one subfamily of this period, Necrolemurinae, a definite
relationship with Tarsius is indicated. Most remaining Paleocene and Eocene Primates
have (or should have) been termed " lemuroid " in the past. A great many fossil
genera have consistently been classified as tarsioid when there is no basis, in known
parts, for so doing. If one examines the better known lemuroid Primates they seem
to be of four general kinds, as follows :
(i) Species on the border-line between Primates and Insectivora as these orders
are currently understood, of the families :
Amphilemuridae
Microsyopidae
Apatemyidae
24 A NEW EOCENE PRIMATE GENUS, CANTIUS
(2) Specialized Primates of early extinction. Forms too specialized dentally to
be near ancestral lines of later stocks :
Families :
Plesiadapidae
Carpolestidae
Anaptomorphidae [in part]
Paromomyidae [ — Phenacolemuridae, in part]
(3) Relatively generalized groups that could be near the ancestry of some living
families, but which show no convincing evidence of having reached a tarsioid,
or higher, grade of advance.
Families :
Adapidae
Notharctidae
(4) Generalized Primates, possibly close to the stocks which gave rise to living
groups, but which are not well enough known to indicate their grade of
advance.
Omomyidae
perhaps some [paromomyids and anaptomorphids]
Past usage has roughly equated the term lemuroid with at least some members of
each of the four groups of extinct families listed above. Such usage implies, however,
that all these types can be assigned to the Lemuridae, which is not the case. On the
other hand, it is surely valid to assume that the ancestral stocks of all the groups
which have advanced beyond the lemur-like state (represented today by tupaiids
and lemurids) went through such a level of advancement, and consequently their
ancestors of the early Cenozoic can correctly be called lemuroid or lemur-like.
Obviously then, among fossil forms that are distinctly lemuroid may be found
ancestors of the more advanced Primates : tarsioids, and anthropoids. Some members
of the above families do foreshadow tarsioids. For instance, Navajovius of the Tiff an-
ian Paleocene of Colorado and Teilhardina of the Sparnacian Eocene of Belgium,
both exhibit interesting dental resemblances to the European Middle Eocene tarsiid,
Nannopithex, while the only skull with advanced (or tarsioid) features ever discovered
in North America (Tetonius ; A.M.N.H. 4194) also possesses upper molars that,
together with a general similarity of detail, share with those of Nannopithex an
unusual feature, the nannopithex-fold. At present these four genera are assigned to
three different families, viz.
Navajovius and Teilhardina . Omomyidae (Simpson, 1940 : 208)
Nannopithex .... Tarsiidae (Simons, 1961 : 61)
Tetonius . . . . Anaptomorphidae (Gazin, 1958 : 73)
Dental variation among these primates is not as great as their assignment to different
families implies, being in most ways far less than seen in the Malagasy family
Lemuridae. Yet, until better cranial remains are known for some omomyids, paro-
momyids and anaptomorphids it will remain difficult to judge whether or not these
A NEW EOCENE PRIMATE GENUS, CANTIUS 25
groups show the same definite indications of tarsioid relationships in their osteology
than they do in their dental anatomy. All these most interesting Primates fall
within the fourth group listed on p. 24. In regard to the first three groups primatolo-
gists are on safer ground, for they are either so primitive dentally that tarsioid
relationships cannot be considered or they are known from skulls that show no
convincing tarsioid features. In the broadest sense it appears reasonable to call these
forms lemuroid, and it is with them that the following paragraphs will be concerned.
Border-line Primates
A mphilemuridae
This family, proposed by Heller (1935) for the reception of Amphilemur eocaenicus
from the Middle Eocene deposits of the Geiseltal, has recently been assigned to the
Erinaceoidea (Insectivora) by McKenna (1960 : 58). To this subfamily McKenna
also assigns, among others, the following North American Paleocene and Eocene
genera : Entomolestes , Macrocranion and Sespedectes. As far as Primate classification
is concerned, this procedure is not likely to improve the taxonomy of the genera
involved, for the type of Amphilemur cannot be located at Halle, and if lost makes an
unsatisfactory choice as the type genus of a subfamily known otherwise only in the
New World, and to which it may not belong. Both Simpson (1945) and Hill (1953)
refer Amphilemur to the Adapidae, incertae sedis. In its poorly-known state, now
entirely dependent on Heller's text and figures, there is little basis for placing it
elsewhere. In Adapis parisiensis the lower canine is often very small and otherwise
the antemolar teeth exhibit gradual and nearly unbroken size-decreases from back
to front, which is not unlike the size sequence of these lower teeth in Amphilemur
and which shows that the same general kind of tooth proportions seen in Amphilermir
can occur in a primate.
Microsyopidae
McKenna (1960 : 76) has re-defined this family so as to include the following
genera : Microsyops, Craeseops and Cynodontomys. His view, that this family
belongs among the Primates appears to have sound justification. He is of the opinion
that the nearest affinities of the Microsyopidae, as re-defined by him, are with the
more primitive plesiadapids of North America. Moreover, he remarks (1960 : 78, 79)
while discussing Microsyopidae :
" Recently Hiirzeler (1948^, pp. 343-356) has described a new genus and
species, Alsaticopithecus leemanni, from the Lutetian of Alsace, based on
excellent dental material. The animal was placed in the Primates incertae
sedis. Alsaticopithecus appears to me to be a microsyopid ... As Hiirzeler
notes, the genus is indeed primatelike. Thus Alsaticopithecus has a bearing on
the problem of microsyopid origins. The genus is an interesting extension of
the known statigraphic and geographic range of microsyopids."
In view of this assignment of Alsaticopithecus, taken together with what is now
known of microsyopid anatomy, including cranial material, soon to be reported on
26 A NEW EOCENE PRIMATE GENUS, CANTIUS
by McKenna, Hiirzeler's conjecture that this form has significant dental resemblances
to Anthropoidea is no longer tenable.
Apatemyidae
The series of genera now assigned to this family were first clearly distinguished
from plesiadapids by Jepsen (1934). Although they are retained provisionally in
the order Primates by Gazin (1958), research on the basicranium reported by Hiirzeler
(1949 : 485) indicates that this group is almost certainly not to be referred to the
Primates, as the order is now defined. Notwithstanding this, the apatemyids may
have had their origin in the same eutherian stock from which the Primates arose — a
conclusion which is indicated principally by molar-pattern resemblances. They
certainly have no close connection with Paleocene- Recent Primates and will not be
considered further here.
Specialized Lines of Early Extinction
A number of divisions of the order of family and subfamily status, which became
extinct before the end of the Eocene, exhibit such pronounced dental specializations
that it is clear they could not be near the ancestry of any living Primate species. Of
these, the carpolestids are the least well understood, but conformation of the fourth
lower premolar alone (elongate and saw-like) removes them from consideration as
ancestral to surviving lines. The same applies to plesiadapids, and to Phenacolemur1
because of the marked reduction of their anterior dentition. Even so, Plesiadapis
is of interest because of its geographical distribution. Species of this genus are known
from the late Paleocene of both Europe and North America. This occurrence demon-
strates that climatic conditions as well as the requisite land-bridges, allowing not
only other mammals but prosimians to communicate between these two continents
then existed, and helps to explain the presence of omomyids, microsyopids and possi-
bly other primate families, in both continents in the early Eocene.2
Among the plesiadapids, carpolestids, and paromomyids the cranial and post-
cranial anatomy is poorly known except in two species of Plesiadapis. The limb
bones of Plesiadapis studied by Simpson (and greatly amplified by undescribed
specimens recently collected by Russell in the Cernay Paleocene deposits near Rheims
in central France) are primitive, but can be spoken of as lemur-like. Russell (1960)
briefly reported on a remarkably well-preserved skull of Plesiadapis from the Cernay
region. This is the oldest primate skull yet to be described. The skull of Plesiadapis
is very broad, flat, with a small brain-case, while the premaxillaries are enormously
expanded and make up most of the muzzle. Elsewhere, among Primates such
expanded premaxillaries occur only in the Madagascan lemur Daubentonia. However,
a relationship with the latter is excluded by the presence in Plesiadapis, but not
in Daubentonia, of a tubular ectotympanic element. Finally, it is clear that in
Plesiadapis there is no post-orbital bar, nor any indication of post-orbital processes.
1 Here regarded as a paromomyid, following McKenna (1960 : 70).
2 Basically all living prosimians are tropical forest dwellers. Uniformitarian reasoning would suggest
that their early Cenozoic forerunners were also warm-climate animals.
A NEW EOCENE PRIMATE GENUS, CANTIUS 27
From this feature of the Cernay Plesiadapis skull it must be concluded that the basal
stock of the order lacked the post-orbital bar, although it occurs in all other undoubted
Primates (for which this part of the skull is known) with the single exception of
Anagale from the Oligocene of Mongolia.
The families Anaptomorphidae and Paromomyidae are known only from North
America. In terms of present knowledge, they include the most likely candidates
for a relationship with the rise of tarsioids and of surviving higher Primates, possibly
by way of containing forms ancestral to the omomyids. Regrettably members of
these families, with the exception of Tetonius, are known only from dentitions, and
consequently, little can be said as to whether they are lemur-like or tarsier-like.
The same is true for omomyids. Nevertheless, dental analogies suggest that some
of the species referred to these families were at least transitional to the tarsioid grade
and such an assumption is further indicated by the large, forward-directed orbits
and expanded brain of Tetonius.
VI. REVISED CLASSIFICATION OF EUROPEAN EOCENE PRIMATES
Revisions included here, in Simons (1961), and in a number of other works published
since Simpson (1945) now make it possible to assign taxonomically certain European
early Cenozoic Primates formerly listed by Simpson as incertae sedis, as well as to
transfer others for which a better placement can be supported. The most recent
extensive treatment of these forms is by Hill (1953, 1955), but Hill's taxonomy is
clearly synoptic and no revision of fossil prosimians is included in these works. The
following section, therefore, attempts to bring the classification of the fossil Primates
of Europe up to date.
Genera to be considered here are listed in Simpson (1945) as follows :
Plesiadapidae :
Megachiromyoides Weigelt, 1933. M. Eoc.; Germany.
Adapidae incertae sedis :
Caenopithecus Riitimeyer, 1862. M. Eoc.; Switzerland.
(?) PROSIMII of uncertain infraorder or family :
Ceciliolemur Weigelt, 1933. M. Eoc.; Germany.
Europolemur Weigelt, 1933. M. Eoc.; Germany.
Megatarsius Weigelt, 1933. M. Eoc.; Germany.
Microtarsioides Weigelt, 1933. M. Eoc.; Germany.
Two additional Eocene Primates from Europe described by Hiirzeler (1946, 1947)
are also discussed in the following pages.
Gesneropithex peyeri Hiirzeler, 1946. M. Eoc.; Switzerland.
Alsaticopithecus leemanni Hiirzeler, 1947. M. Eoc.; Alsace.
Inasmuch as the European plesiadapids, and a specimen from the Walbeck
Paleocene of Germany, referred to Phenacolemur by Weigelt (1947) but of uncertain
generic and familial assignment are currently under study by others they are not
discussed here in detail. These groups are, however, indicated in Text-chart I, in
28
A NEW EOCENE PRIMATE GENUS, CANTIUS
PALEOCENE
EOCENE
PROVINCIAL AGE
THANETIAN
SPARNACIAN
INCL. EARLY
YPRESIAN
CUISIAN
EQUALS LATE
YPRESIAN
LUTETIAN
BARTONIAN
LUOIAN
EARLY
LATE
EARLY
LATE
_,^-
Necrolemur
Necrolemur
Necrolemur
^"
"--
Microchoerus
Microchoerus
X
,,---'
ftilhordino
,'-
Nannopithex
Nannopithe*
Pseudolor/s
Gesneropithex
Anchomomys
Adapts
Protoadapis
Adopis
Periconodon
> OMOMY
DAE
- Cantius
Anchomomys
— Adapts
Caenopithecus
Anchomomys
Adapts
Protoadapis
Anchomomys
Adapts
Protoadapis
Pronycticebus
\
X
V
Chiromyoides
^•* ^
Protoadapis ^
>
N
*\_
Plesiadopis
Plesiadapis *
• PLESIAC
JAPIOAE
\
\
N
X.
Plalychoerops
^
Phenacolemur
">PAROMO
MYIDAE
\
\
X
***•.
J
Alsoticopithec
us > MICRO
J
SYOPIDAE
CHART I
Temporal occurrence of early Cenozoic European primate genera indicating
approximcte interrelationships. (* Occurrence in this horizon dubious.)
CHART i.
A NEW EOCENE PRIMATE GENUS, CANTIUS 29
which temporal occurrences and approximate interrelationships of all European
early Tertiary Primates are shown.
Megachiromyoides
Since Megachiromyoides is frequently referred to in the literature as a primate, it
should be emphasized that Stehlin & Schaub (1951 : 21) stated that it is a rodent,
assignable to the genus Aeluravus. A recent examination, by the writer, of the holo-
type of " Megachiromyoides " from the Geiseltal Brown Coals (described by Weigelt,
1933) has confirmed the view that this form cannot possibly be a plesiadapid.
Caenopithecus
This interesting primate from Middle Eocene (Lutetian) deposits near Egerkingen,
Switzerland was originally described by Rutimeyer (1862). Stehlin (1916) discussed
the species further and figured a mandible which shows only one incisor alveolus on
either side. He stated that his material indicated the same lower dental formula
as in Tarsius, and noted other advanced features including the loss of the first upper
and lower premolars, and a short rostrum.
Corroboration of Simpson's placement (1940) of this primate among adapidscan
be drawn from its similarities with the Lutetian species Protoadapis klatti. Principally
Caenopithecus differs from Protoadapis in showing more reduced molar paraconids,
in the possession of distinct mesostyles and in having less well-developed upper molar
lingual cingulum. Protoadapis also exhibits a more distinct P3 protocone. In spite of
these generic differences, however, Caenopithecus and Protoadapis are, in upper and
lower dental conformation, in dental formula and in what is known of the skull,
much closer to each other than either is to Adapis, but (in different ways) both
exhibit some features of resemblance to a species of the latter genus.
Stehlin (1916 : 1319) remarked that knowledge then available of Caenopithecus
lemuroides indicated affinities with Protoadapis and Pelycodus and this view has been
strengthened by the better material of Protoadapis klatti now known. Protoadapis
and Caenopithecus agree in possessing the following features, some of which are
unknown among other European Eocene Primates : Long and large canines (above
and below) — not very procumbent ; PI/I absent and replaced by diastema ; P3
higher than P4 ; metaconule of upper molars lacking but distinct paraconule present ;
well-developed hypocones on M1"2 only ; upper molar cingula (labial and lingual)
well defined and in some individuals continuous across lingual base of protocones ;
similar proportions in lower molars.
Common possesssion of pronounced upper molar cingula, absence of a distinct Pa
protocone and near identity in overall proportions of the lower molars between
Caenopithecus and Adapis parisiensis, tend to link species of these two genera as
well. Weigelt noted a slight mesostyle on the M2 of Protoadapis klatti (also in agree-
ment with Caenopithecus} and expressed the view that his Brown Coal species was
closer to Caenopithecus than to Adapis. Moreover, his illustration of the skull of
P. klatti (1933, pi. 6, fig. 4) indicates a vertically deepened anterior base for the
zygomatic arch, as in Caenopithecus lemuroides. One further similarity between
30 A NEW EOCENE PRIMATE GENUS, CANTIUS
these two types of Primates is the possible loss (reported for species of both genera)
of all but one pair of lower incisors. Because of the low angle at which the anterior
alveolar border of the mandible of Caenopithecus (Basel Eh. 733) from which this
was deduced, is broken off, the possibility remains that there were more incisors
below. In some specimens of Adapis parisiensis in the Paris Museum, where this
margin is broken off in a similar way, the incisor alvoli are entirely missing. If these
were the only known specimens of A . parisiensis its lower incisor formula would be
equally uncertain. The exact number of lower incisor pairs in given Eocene primate
species can seldom be stated categorically. Both Caenopithecus and Protoadapis
could have had two pairs of these teeth, but present evidence is not adequate to
prove that they did.
A further note of some interest regarding Caenopithecus , and the nature of true
hypocones and pseudohypocones, is provided by a specimen of this primate at Basel
(Eh. 727) which shows both kinds of hypocone formation in one individual. The
retention of both of these patterns in this dentition suggests that a latency for
producing pseudohypocones, as in the line leading to Notharctus as well as for true
hypocone production in Adapis, probably existed in the ancestral adapid-notharctid
stock.
Periconodon
The genus Periconodon was established by Stehlin (1916) for a species " helveticus "
which had been assigned by Riitimeyer (1891 : 115) to the North American genus
Pelycodus. Like most Caenopithecus materials the type of Periconodon helveticus
came from Middle Eocene deposits near Egerkingen, Switzerland. Apparently no
additional specimens referable to this genus have been discovered since Stehlin's
discussion.1 The individual illustrated by Stehlin (1916, pi. 22 fig. 3), which preserves
P3 and M1"2 shows a distinctive pericone on the antero-internal base of M1"2 proto-
cones. Since a pericone sometimes occurs here in Omonys (see Gazin, 1958, pi. 6,
figs. 3 4) but not elsewhere among early Tertiary Primates, and because omomyids
are known to have been present in the European Eocene fauna, the provisional
reference of Periconodon to this family seems advisable. Furthermore, the small
size of the species and general conformation of the teeth does not suggest adapid
or notharctid affinities.
Ceciliolemur and Microtarsioides
Ceciliolemur is now considered by Matthes (1957), and by others, to be an insecti-
vore, primarily because it seems to have had dermal spines. Recent examination of
the holotypes of Ceciliolemur, G.P.I.H. 4237, and of Microtarsioides, G.P.I.H. 4235,
both from the Geiseltal Brown Coals of Cecilia mine, locality Leichenfeld II, indicates
strongly that they belong to the same species. Cheek teeth do not survive in either
specimen, so dental formula and affinities cannot be determined. Moreover, the
1 Heller (1930) and Weigelt (1933) both mention the presence of isolated upper molars of this Primate
in the Geiseltal Brown Coals. These teeth do not belong to Periconodon, but are those of a larger Primate
from which the enamel has been resorbed. What remains of the cusp bases suggests their reference to
Protoadapis klatti.
A NEW EOCENE PRIMATE GENUS, CANTIUS 31
condition of bones and of eruption of lower teeth indicate that both are very young
individuals, perhaps pre-natal. No basis remains for referring either of these speci-
mens to the Primates. It is a remarkable accident of fossilization that either of these
very delicate specimens were preserved, considering that the body excluding the tail,
in both cases, is only about 4 cm. long. Locality data indicates recovery from the
same place, and possibly they are from the same " litter ".
Europolemur and Megatarsius
As discussed above (pp. 13, 14) these primate genera are junior synonyms of
Protoadapis.
A Isaticopithecus
The species A Isaticopithecus leemanni Hiirzeler (1947) can be assigned to the family
Microsyopidae, as re-defined by McKenna (1960 : 76), see p. 25.
Gesneropithex
Gesneropithex peyeri Hiirzeler (1946) is based on a lower jaw containing P4_M2and
one second upper molar collected in Ludian deposits near Bosgen, Switzerland.
The lower teeth differ from Alsaticopithecus and resemble such adapids as Anchomo-
mys, in that M1_2 lack distinct hypoconulids. Moreover, conformation of the upper
second molar is within the general range of variability of species now assigned to the
Adapidae. M2 is most similar in size and cusp pattern to those of Anchomomys latidens,
but wear obscures some of its features. Anchomomys and Gesneropithex also agree in
one rather unusual feature of M2 ; both have a strong anterior crest running from
the metaconule to the metacone. Presence of an upper molar mesostyle in Gesnero-
pithex is in agreement with Caenopithecus. It seems reasonable to assume that
Gesneropithex peyeri can be referred to the Adapidae (s.s.).
REVISED CLASSIFICATION1
Order PRIMATES
Suborder PROSIMII
Family TARSIIDAE Gray, 1870
Subfamily NECROLEMURINAE Simpson, 1933
MICROCHOERUS Wood, 1844
Synonyms — Microchaerus Forbes, 1894
Necrolemur : Filhol, 1880 (in part)
Microchoerus erinaceus Wood, 1844
Microchoerus edwardsi (Filhol, 1880)
Synonym — Necrolemur edwardsi Filhol, 1880
Microchoerus ornatus Stehlin, 1916
1 Since a revision of the European plesiadapids is currently being undertaken by Russell at the Paris
Museum they have been omitted from this classification.
32 A NEW EOCENE PRIMATE GENUS, CANTIUS
NECROLEMUR Filhol, 1873
Necrolemur antiquus Filhol, 1873
Necrolemur zitteli Schlosser, 1887
NANNOPITHEX Stehlin, 1916
Synonyms — Necrolemur Chantre & Gaillard, 1897
Pseudoloris : Weigelt, 1933
Nannopithex filholi (Chantre & Gaillard, 1897)
Synonyms — Necrolemur filholi Chantre & Gaillard, 1897
Nannopithex pollicaris Stehlin, 1916
Nannopithex raabi (Heller, 1930)
Synonyms — Pseudoloris abderhaldini Weigelt, 1933
Necrolemur raabi Heller, 1930
PSEUDOLORIS Stehlin, 1916
Synonym — Necrolemur : Filhil, 1889
Pseudoloris parvulus (Filhol, 1889)
Synonym — Necrolemur parvulus Filhol, 1889
Family OMOMYIDAE Gazin, 1958
TEILHARDINA Simpson, 1940
Synonym — Omomys Teilhard, 1927
Teilhardina belgica (Teilhard, 1927)
Synonym — Omomys belgicus Teilhard, 1927
CANTIUS nov. gen.
Synonym — Protoadapis : Cooper, 1932
Cantius eppsi (Cooper, 1932)
Synonym — Protoadapis eppsi Cooper, 1932
PERICONODON Stehlin, 1916
Synonym — Pelycodus Riitimeyer, 1891
Periconodon helveticus (Riitimeyer, 1891)
Synonym — Pelycodus helveticus Riitimeyer, 1891
Family ADAPIDAE Trouessart, 1879
ADAPTS Cuvier, 1822
Synonyms — Aphelotherium Gervais, 1848
Leptadapis Gervais, 1852
Palaeolemur Delfortrie, 1873
A NEW EOCENE PRIMATE GENUS, CANTIUS 33
Adapts parisiensis Blainville, I84I1
Adapts magnus Filhol, 1874
A da pis rutimeyeri Stehlin, 1912
Adapis prisons Stehlin, 1916
Adapis sciureus Stehlin, 1916
PROTOADAPIS Lemoine, 1891
Synonyms — Plesiadapis : Lemoine, 1878 (in part)
Protadapis : Stehlin, 1912
Megatarsius Weigelt, 1933
Europolemur Weigelt, 1933
Protoadapis curvicuspidens2 Lemoine, 1878
Synonym — Protadapis recticuspidens : Stehlin 1912 (in part)
Protoadapis angustidens (Filhol, 1888)
Synonyms — Adapis angustidens Filhol, 1888
Protadapis brachyrhynchus Stehlin, 1912
Protoadapis klatti (Weigelt, 1933)
Synonyms — Europolemur klatti Weigelt, 1933
Megatarsius obeli Weigelt, 1933
CAENOPITHECUS Riitimeyer, 1862
Caenopithecus lemur oides Riitimeyer, 1862
PRONYCTICEBUS Grandidier, 1904
Pronycticebus gaudryi Grandidier, 1904
ANCHOMOMYS Stehlin, 1916
Synonym — Caenopithecus : Riitimeyer, 1891
Anchomomys Pygmaeus (Riitimeyer, 1891)
Anchomomys gaillardi Stehlin, 1916
Anchomomys quercyi Stehlin, 1916
Anchomomys latidens Teilhard, 1916
GESNEROPITHEX Hiirzeler, 1946
Gesneropithex peyeri Hiirzeler, 1946
Family MICROSYOPIDAE new sense, McKenna (1960)
ALSATICOPITHECUS Hiirzeler, 1947
Alsaticopithecus leemanni Hiirzeler, 1947
? Primates incertae sedis
AMPHILEMUR Heller, 1935
Amphilemur eocaenicus Heller, 1935
1 For numerous synonyms of the various species of Adapis, see Stehlin (1912, 1916).
2 Here designated as the type species of Protoadapis.
34 A NEW EOCENE PRIMATE GENUS, CANTIUS
VII. CONCLUSIONS
' The occurrence of a new omomyid prosimian genus, Cantius, from the English
Sparnacian is reported on and its distinctions from the Middle Eocene genus Proto-
adapis outlined. Assignment of this form to the Omomyidae reinforces the extension
of the range of this family into the Old World suggested by a few previously
described forms. Although additional resemblances to North American notharctids
are noted for Protoadapis, it is referred to the Adapidae on the basis of its greater
similarities with Caenopithecus and Pronycticebus. Preliminary evidence suggests
that Notharctidae and Adapidae, as denned by Gazin (1958) may not actually be
distinguishable, but solution of this problem requires additional comparative study.
Certain loris-like features of the genus Pronycticebus are pointed out. Just possibly,
these can be interpreted as indicating the differentiation of the lorisiform prosimians
from the general stock of the Adapidae (s.l.) . A high degree of dental similarity between
an undoubted lorisoid prosimian, Progalago, of the early Miocene of Kenya, and the
late Eocene European primate Anchomomys is stressed. These findings are in contrast
to the widespread opinion that no close relationship of early Cenozoic with Recent
prosimian families can be demonstrated, but is in line with the vertical taxonomic
association of necrolemurines and tarsiines proposed by Simons (1961). However,
zoogeographical and morphological considerations alone are sufficient to indicate
that it would be unwise to regard any such relationships direct ancestor-descendant
lineages.
A number of species previously regarded as incertae sedis are here re-assigned or
removed from the order and the taxonomy of European Eocene Primates revised.
VIII. ACKNOWLEDGEMENTS
I should like to thank Dr. K. P. Oakley of the British Museum (Natural History),
Dr. J.-P. Lehman of the National Museum of Natural History, Paris, and Drs. H.-W.
Matthes and G. Krumbiegel of the Geological and Paleontological Institute of the
Martin Luther University of Halle/Wittenburg for the opportunity to study and
report on Eocene Primates in their respective collections. Thanks are also due to
Dr. G. L. Jepsen of Princeton University, Sir Wilfrid Le Gros Clark, and to many
others with whom this subject has been discussed. Figures were prepared by Miss
Christine Court of Oxford University and Mrs. Margaret E. Freeman of New Haven,
and early completion of the work facilitated by a grant from the Wenner-Gren
Foundation for Anthropological Research.
IX. REFERENCES
ABEL, O. 1931. Die Stellung des Menschen im Rahmen der Wirbeltiere. 398 pp. Jena.
BARNETT, C. H. & NAPIER, J. R. 1953. The rotatory mobility of the fibula in eutherian
mammals. /. Anat. Lond., 87 : 11-21.
CHOW, MIN-CHEN. 1961. A New Tarsioid Primate from the Lushi Eocene, Honan. Verte-
brata Palasiatica, Peking, 5 : 1-5, 3 figs.
CLARK, W. E. LE GROS. 1934. On tne skull structure of Pronycticebus gaudryi. Proc. Zool.
Soc. Lond., 1934 : 19-27, 3 figs.
— 1956. A Miocene lemuroid skull from East Africa. Fossil Mammals of Africa, 9, 6 pp.
i pi. British Museum (Nat. Hist.), London.
A NEW EOCENE PRIMATE GENUS, CANTIUS 35
CLARK, W. E. LE GROS & THOMAS, D. P. 1952. The Miocene Lemuroids of East Africa.
Fossil Mammals of Africa, 5, 20 pp., 3 pis. British Museum (Nat. Hist.), London.
COOPER, C. FORSTER. 1932. On some mammalian remains from the Lower Eocene of the
London Clay. Ann. Mag. Nat. Hist., London (10) 9 : 458-467, pis. 11-12.
FILHOL, H. 1888. Description d'une nouvelle espece d'Adapis. Bull. Soc. philom. Paris,
7, 12 : 10-12.
FORSYTH MAJOR, C. I. 1901. On some characters of the skull in the lemurs and monkeys
Proc. Zool. Soc. Land., 1901 : 129-153, pis. 11-13.
GAZIN, C. L. 1958. A review of the Middle and Upper Eocene Primates of North America.
Smithson Misc. Coll., Washington, 126 : 1-112, pis. 1-14.
GRANDIDIER, G. 1904. Un noveau Lemurien fossile de France, le Pronycticebus Gaudryi.
Bull. Mus. Hist. nat. Paris, 10 : 9-13, 3 figs.
1905. Les lemuriens disparus. Nouv. Arch. Mus. Hist. nat. Paris (4) 7 : 1-42.
GREGORY, W. K. 1920. On the structure and relations of Notharctus, an American Eocene
primate. Mem. Amer. Mus. Nat. Hist., New York (n.s.) 3 : 49-243, pis. 23-59.
1922. The origin and evolution of the human dentition, xviii + 548 pp., 14 pis. Baltimore.
GRANGER, W. & GREGORY, W. K. 1917. A revision of the Eocene Primates of the genus
Notharctus. Bull. Amer. Mus. Nat. Hist., New York, 37 : 841-859, pis. 103-107.
HELLER, F. 1930. Die Saugetier fauna der mitteleozanen Braunkohle des Geiseltals bei
Halle. Jb. hallesch. Verb. Erf. mitteldtsch. Bodensch., Halle (n.f.) 9 : 13-41, pis. 1-5.
— 1935. Amphilemur eocaenicus n. g. et n. sp., ein primitiver Primate aus dem Mitteleozan
des Geiseltales bei Halle a. S. Nova Acta Leop. Carol., Halle a. S. (n.s.) 2 : 293-300.
HILL, W. C. O. 1953. Primates. Comparative Anatomy and Taxonomy. I. Strepsirhini.
xxiv + 798 pp., 34 pis. Edinburgh.
1955- Primates. Comparative Anatomy and Taxonomy. II. Haplorhini : Tarsioidea.
347 pp., 14 pis. Edinburgh.
HURZELER, J. 1946. Gesneropithex Peyeri nov. gen. nov. spec, von Gosgen (Solothurn).
Eel. geol. Helv., Lausanne, 39, 2 : 354-361, 4 figs.
— 1947. Alsaticopithecus Leemanni nov. gen. nov. spec., ein neuer Primate aus dem
unteren Lutetien von Buchsweiler im Unterelsass. Eel. geol. Helv., Lausanne, 40 : 343-
356, 12 figs.
— 1948. Zur Stammesgeschichte der Necrolemuriden. Abh. schweiz. palaont. Ges., Zurich,
66 : 1-46, 41 figs.
1950. liber die europaischen Apatemyiden. Eel. geol. Helv., Lausanne, 42, 2 : 485.
JEPSEN, G. L. 1934. A revision of the American Apatemyidae and the description of a new
genus, Sinclair ella, from the White River Oligocene of South Dakota. Proc. Amer. Phil.
Soc., Philadelphia, 74 : 287-305, pis. 1-3.
LEMOINE, V. 1878. Communication sur les ossements fossiles des terrains tertiares inferieurs
des environs de Reims. Bull. Soc. Hist. nat. Reims, 1878 : 90-113, pis. 1-5.
1891. Etude d'ensemble sur les dents des mammiferes fossiles des environs de Reims.
Bull. Soc. geol. Fr., Paris (3) 19 : 263-290, pis. 10, n.
LEMOINE, V. & AUMONIER, M. 1880. Terrains tertiaires des environs de Reims. Ass.
Franc. Avanc. Sci., Paris, 9 : 605-619, pi. 5.
MATTHES, H. W. 1957- Die mitteleozane Saugerfauna des Geiseltales and ihre verwand-
schaftlichen Beziehungen. Wiss. Z. Univ. Halle, Math. Nat., 7 : 37-62, 22 figs.
MCKENNA, M. C. 1960. Fossil mammalia from the early Wasatchian Four Mile fauna,
Eocene of northwest Colorado. Bull. Dep. Geol. Univ. Calif., Berkeley, 37 : 1-130,
64 figs.
OSBORN, H. F. 1891. A review of the Cernaysian Mammalia. Proc. Acad. Nat. Sci. Philad.,
1890 : 51-62, 6 figs.
PIVETEAU, J. 1957. Primates, paleontologie humain. Traite de Paleont., Paris, 7 : 1-675.
RUSSELL, D. E. 1960. Le crane de Plesiadapis. Bull. Soc. geol. Fr., Paris, 7 : 312-314,
pi. 18.
36 A NEW EOCENE PRIMATE GENUS, CANTIUS
RUTIMEYER, L. 1862. Eocane Saugethiere aus dem Gebiete des schweizerischen Jura.
Denkschr. schweiz. Ges. Naturw., Zurich, 19 : 1-248, pis. 1-6.
— 1891. Die Eocane Saugethier-welt von Egerkingen. Abh. schweiz. palaont. Ges., Zurich,
18 : 1-153, pls- T~8-
SIMONS, E. L. 1961. Notes on Eocene Tarsioids and a revision of some Necrolemurinae.
Bull. Brit. Mus. (Nat. Hist.) Geol., London, 5 : 43-69, pis. 12-14.
- 19610. The dentition of Ourayia : — its bearing on relationships of omomyid prosimians.
Postilla, Yale Peabody Mus., New Haven, Conn., 54 : 1-20, 3 figs.
SIMONS, E. L. & RUSSELL, D. E. 1960. Notes on the cranial anatomy of Necrolemur. Breviora
Mus. Comp. Zool. Harv., Cambridge, Mass., 127 : 1-14, 3 figs.
SIMPSON, G. G. 1940. Studies on the earliest primates. Bull. Amer. Mus. Nat. Hist., New
York, 77 : 185-212, 8 figs.
1945- The principles of classification and a classification of mammals. Bull. Amer. Mus.
Nat. Hist., New York, 85 : 1-350.
- 1955. The Phenacolemuridae, new family of early primates. Bull. Amer. Mus. Nat.
Hist., New York, 105 : 415-441, pis. 30-35.
STEHLIN, H. G. 1912. Die Saugetiere des schweizerischen Eocaens. Abh. schweiz. palaont.
Ges., Zurich, 38 : 1165-1298, figs, 244-288.
1916. Die Saugetiere des schweizerischen Eocaens. Abh. schweiz. palaont. Ges., Zurich,
41 : 1299-1552, pis. 21-22.
STEHLIN, H. G. & SCHAUB, S. 1951. Die trigonodontie der simplicidentaten nager. Abh.
schweiz palaont. Ges., Zurich, 67 : 1-385, 620 figs.
TEILHARD DE CHARDIN, P. 1921. Les mammiferes de 1'Eocene inferieur Fran9ais et leurs
gisements. Ann. Paleont., Paris, 10 : 171-176 ; 11 : 9-116, 42 figs.
WEIGELT, J. 1933. Neue Primaten aus der mitteleozanen (oberluteitischen) Braunkohle des
Geiseltals. Nova Acta Leop. Carol., Halle (n.f.) 1 : 97-156, pis. i-u.
PLATE i
Cantius eppsi (Cooper)
A. Right maxilla, B.M.N.H., M 15145, with P3-M2. B. Right mandible, B.M.N.H., M 13773
(Type), with P3-M3. x 6-0 approx.
Bull. B.M. (N.H.) Geol. 7, i
PLATE i
A
\
PLATE 2
Protoadapis klatti (Weigelt)
Anterior dentition, G.P.I. H. 4258. xya approx.
Bull. B.M. (N.H.) Geol. 7, t
PLATE 2
PLATE 3
Protoadapis klatti (Weigelt)
Specimens of Protoadapis at Halle. A. Right M3, G.P.I.H. 7325 ; B. Right M1, G.P.I. H.
4292 ; c. Right M2, cf. Protoadapis, G.P.I.H. 4238 ; D. Left M3, G.P I H 4292 • E Left
M2-3, Left P4_M8, G.P.I.H. 4310. All x 8 -o approx.
(Photographs slightly retouched.)
Bull. B.M. (N.H.) Geol. 7, i
PLATE 3
A
B
PRINTED IN GREAT BRITAIN BY
ADLARD AND SON, LIMITED
BARTHOLOMEW PRESS DORKING
THE BRACHIOPOD GENUS
CYCLOTHYRIS
E. F. OWEN
BULLETIN OF
THE BRITISH MUSEUM (NATURAL HISTORY)
GEOLOGY Vol. 7 No. 2
LONDON: 1962
THE BRACHIOPOD GENUS CYCLOTHYRIS
BY
ELLIS FREDERIC OWEN
Pp- 37-63 ; •?&• 4. 5 ; 10 Text-figures
BULLETIN OF
THE BRITISH MUSEUM (NATURAL HISTORY)
GEOLOGY Vol. 7 No. z
LONDON: 1962
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(NATURAL HISTORY), instituted in 1949, is
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within one calendar year.
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PRINTED BY ORDER OF THE TRUSTEES OF
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THE BRACHIOPOD GENUS CYCLOTHYRIS
By ELLIS FREDERIC OWEN
SYNOPSIS
A revision of M'Coy's genus Cyclothyris from the Upper Aptian is made and serial sections
of the type-species presented for the first time. In addition eight other species of Cyclothyris
are described, two of these are new species. The subfamily Cyclothyrinae of Makridin is
emended and discussed. A comparison is made between Cyclothyris from the Lower Greensand
and the Upper Chalk genus Cretirhynchia. A new generic name is proposed for five Lower
Cretaceous Rhynchonellidae.
INTRODUCTION
THE name Cyclothyris was first used by M'Coy (1844 : 103) to indicate a group of
brachiopods which he called "... those curious species in which the deltidium
completely surrounds the foramen ". No geological age was mentioned nor was
any species named. M'Coy did, however, figure a specimen (p. 150, fig. 29) which is
probably lost.
From the beak characters of Mesozoic and Palaeozoic Rhynchonellidae it seems
likely that M'Coy was referring to an early Cretaceous genus and, from the arrange-
ment of the costae, shell outline, growth-marks and interarea as seen in his fig. 29,
it is probable that M'Coy had selected a specimen of Aptian age.
FIG. i. Copy of M'Coy's (1844) original illustration of fig. 29, p. 150.
Previously, J. de C. Sowerby (1825 : 165, pi. 502, fig. i) had described and figured
a rhynchonellid from the Upper Aptian, Sponge Gravel of Faringdon, Berkshire
under the name Terebratula lata. He subsequently changed this specific name to
T. latissima in the index to the Mineral Conchology published in 1829, having realized
that his father J. Sowerby (1815 : 227) had already used the name T. lata for a
brachiopod from the Drift of Lowestoft, Suffolk.
The outline, arrangement of costae and growth-lines of J. de C. Sowerby's figured
specimen of Terebratula latissima (pi. 502, fig. i) agree with the figure of M'Coy's
genus Cyclothyris. This similarity was certainly noted by Davidson (1855 : 93) who
included Cyclothyris in his synonymy of Fischer's genus Rhynchonella and quoted
C. latissima in brackets after the generic name, probably as a typical species of
Cyclothyris.
GEOL. 7, 2. 2
4o
THE BRACHIOPOD GENUS CYCLOTHYRIS
Buckman (1906 : 326) was the first to designate T. latissima J. de C. Sowerby as
the type species of Cyclothyris and subsequent authors have followed Buckman.
The genus was not further investigated nor adequately defined until Hertlein &
Grant (1944 : 61) published a full synonymy in their work on Caenozoic Brachiopoda
from western North America.
Stratigraphically the genus ranges from the Upper Aptian to the Upper Cenomanian
and was probably developed from such a middle European form as Lepidorhynchia
which Burn (1956 : 689) described from the Neocomian, Lower Barremian, of
Dm
B
SP
FIG. 2. Four drawings illustrating some of the internal characters of Cyclothyris.
A. Pedicle valve showing subquadrate hinge-teeth (Ht) and large pear-shaped diductor
muscle-scars (Dm). B. Subparallel dental lameleae in pedicle valve (Dl). c. Brachial
valve showing ventral surface of the crura (Cr). D. Brachial valve showing the dorsal
concave surface of the crura (Cr) and deep crenulated dental sockets (Ts) and short
septum (Sp).
Switzerland. Cyclothyris appears to have been confined chiefly to the European
continent, although a notable exception to this is C. americana described and figured
by Cooper (1955 : 3, pi. i) from the Aptian of Arizona. Examples of this species
have not yet been examined by the writer but to judge from the figure on Cooper's
pi. i it may well belong to this genus. Other species referred to Cyclothyris by
Hertlein & Grant (1944 : 63) are probably not referable to the genus, but careful
examination including the use of serial sections is necessary before any conclusions
can be drawn. Another species from the American continent referred to this genus
is Cyclothyris ? subtrigonalis Imlay from the Lower Neocomian of Mexico. This is
shown in serial section (Imlay, 1937 : 571) to have a persistent septum in the brachial
THE BRACHIOPOD GENUS CYCLOTHYRIS 41
valve and a large septalium, both characters which do not appear in serial sections
of the type species C. latissima (J. de C. Sowerby).
In the English Aptian the genus is represented by at least four species namely,
C. latissima, C. depressa (J. de C. Sowerby), C. antidichotoma (Buvignier) and C.
lepida sp. n., but there may be others not yet investigated. The rhynchonelloid
species " R." nuciformis described from the Upper Aptian of Faringdon by J. de C.
Sowerby (1825 : 166), and often erroneously referred to Cydothyris, is probably
related to a species described by the writer (Owen, 1960 : 253) as Lamellaerhynchia
larwoodi from the Upper Aptian of Upware, Cambridge.
B
D
FIG. 3. Diagram illustrating different types of anterior commissure found in Cydothyris.
A. Broadly arcuate. B. Trapezoidal, c. Asymmetrical. D. High arcuate.
Some of these Upper Aptian species are recognizable as passage forms in the Lower
Albian deposits at Leighton Buzzard, Bedfordshire. Middlemiss (1959 : 138) regards
the Shenley Limestone fauna as essentially of an Albian-Cenomanian type. While
possibly true of the terebratuloid fauna it is certainly not of the rhynchonelloids.
With the possible exception of two species the fauna is distinctly Albian, having
close affinities with Upper Aptian faunas at Brickhill and Upware in this country
and Lower Albian faunas at Grandpre, Haute-Savoie and Haute-Saone, France.
Two Lower Albian species of Cydothyris are described here from the Leymeriella
tardefurcata Zone of the Shenley Limestone of Leighton Buzzard, Bedfordshire. One
is the widely quoted C. antidichotoma (Buvignier), which also occurs in the Upper
Aptian, and C. levis sp. n.
By far the commonest Cenomanian species of Cydothyris is C. difformis (Valen-
ciennes in Lamarck) which ranges from the Lower to Middle Cenomanian. It has
42 THE BRACHIOPOD GENUS CYCLOTHYRIS
been recorded from Wiltshire, Devon and Dorset in England and from Normandy in
France, Tournai in Belgium, and Essen in North Germany. Examples of this species
with its many varieties are to be found in the general collections of the British Museum
(Natural History).
Apart from C. difformis, other Cenomanian species here described include C. scaldi-
sensis (d'Archiac) from the Tourtia of Belgium, and C. schloenbachi (Davidson),
a common fossil in the Lower Cenomanian of Somerset and Devon.
The generic name Burrirhynchia is proposed for " Rhynchonella " leightonensis
Walker from the Lower Albian, L. tardefurcata Zone of Shenley Hill, Leighton
Buzzard, Bedfordshire. This genus is also represented in the Upper Aptian, P. nut-
fieldensis Zone, of Upware, Cambridge and Brickhill, Buckinghamshire by the
species " R." cantabrigensis Davidson.
An emended description of " R." leightonensis together with serial sections were
published by the writer (Owen, 1956 : 166, 167) and these may be compared with
serial sections of Burrirhynchia cantabrigensis (Davidson) figured here as Text-fig. 10.
TERMINOLOGY
The terms used in the systematic descriptions here are according to Thomson
(1927), Muir-Wood (1934) and Muir-Wood & Cooper (1960).
Cyclothyrid. As applied to the deltidial plates was proposed by the writer (Owen,
1956 : 165) to replace the term " auriculate " which was used by Buckman (1918 : 18)
to describe the encirclement of the foramen by the deltidial plates.
Sulcus. The term sinus has often been used by authors to describe the median
sulcation of a valve. Muir-Wood & Cooper (1960 : 8) pointed out that the term for
this condition should be sulcus. Their definition was in connection with productoids,
but the term is equally applicable to the rhynchonelloids and is quoted as " Sulcus. —
A major depression in either valve usually median in position, which helps channel
the outgoing stream in feeding. A median fold is usually opposite a median sulcus
in the productoids."
The term sulcus, therefore, as defined by Muir-Wood & Cooper is used throughout
the following systematic descriptions.
In the same publication Muir-Wood & Cooper (1960 : 14) redefined the following
terms :
Capillae. Fine radial, elevated lines (with more than 25 in 10 mm.).
Costellae. Radial lines coarser than capillae (about 15-25 in 10 mm.).
In the following systematic descriptions the measurement of 10 mm. is taken to
indicate a position 10 mm. wide at a distance 10 mm. anterior to the umbo on the
brachial valve. These terms have been used as above in the descriptions of C. anti-
dichotoma, C. lepida sp. n. and C. levis sp. n.
Dimensions of Cyclothyris specimens are given in millimetres but are defined
generally as :
Small, measuring up to 15 mm. long, 20 mm. wide.
Medium, 16-28 mm. long, 21-38 mm. wide.
Large, 29-35 mm. long, 39-45 mm. wide.
THE BRACHIOPOD GENUS CYCLOTHYRIS 43
SYSTEMATIC DESCRIPTIONS
Superfamily RHYNCHONELLACEA Schuchert, 1896
Family RHYNCHONELLIDAE Gray, 1848
Rzhonitskaya (1959), in an attempt to reclassify the rhynchonelloids, cited the
subfamily Cyclothyrinae which was proposed by Makridin (1955) for certain rhyn-
chonelloid genera ranging from the Trias to the Cretaceous. Makridin's definition
quoted by Rzhonitskaya is translated by Mrs. G. A. Cooper, Washington, U.S.A. as :
" Ribbed or plicate rhynchonellids without a septalium ; septa may be developed
or be absent, hinge-plate divided."
This very broad definition included such genera as Stolmorhynchia Buckman, 1914
and Lacunosella Wisniewska, 1932 which, unlike most of the other genera included,
are known to possess falcifer crura and are quite distinct from those genera bearing
radulifer crura.
The following emended definition of Makridin's subfamily Cyclothyrinae is there-
fore proposed :
Costate, costellate, capillate or plicate rhynchonelloids without a septalium ; sep-
tum developed or absent, hinge-plates divided, crura radulifer, foramen hypothyrid.
This would include the following genera :
Cyclothyris M'Coy, 1844.
Cretirhynchia Pettitt, 1950.
Sulcirhynchia Burri, 1953.
Lamellaerhynchia Burri, 1953.
Plicarostrum Burri, 1953.
Lepidorhynchia Burri, 1956.
Burrirhynchia nov.
RANGE. Cretaceous.
Subfamily CYCLOTHYRINAE Makridin, 1955 emended
Genus CYCLOTHYRIS M'Coy, 1844
1844 Cyclothyris M'Coy, p. 103, 150, fig. 29.
1852 Cyclothyris Davidson, p. 93.
1877 Cyclothyris Dall, p. 24.
1906 Cyclothyris Buckman, p. 18.
1913 Cyclothyris Schuchert in Zittel, p. 398.
1918 Cyclothyris Buckman, pi. 14, fig. ia.
TYPE-SPECIES (by subsequent designation, Buckman, 1906). Terebmtula latissima
J. de C. Sowerby.
EMENDED DIAGNOSIS. Medium to large biconvex rhynchonellidae, oval to subtri-
angular in outline. Shell multicostate, costae sharp, coarse to fine, subangular to
rounded. Growth-lines lamellar to step-like. Anterior commissure uniplicate.
Beak-ridges usually distinct, interarea well defined. Foramen medium to large,
circular. Deltidial plates conjunct, cyclothyrid.
44 THE BRACHIOPOD GENUS CYCLOTHYRIS
Internal characters. Umbonal cavities oval in cross-section. Denticulae well
developed. Teeth large, deeply inserted. Dental lamellae strong, subparallel.
Hinge-plates broad, slightly arched in transverse section, divided terminally. Crura
anteriorly concave.
DESCRIPTION. Cyclothyris comprises a very variable group of medium to large
rhynchonellidae. With the exception of perhaps one or two species, the outline is
distinctly transversely oval. A fold and sulcus develop late and there is usually
considerable lateral and anterior thickening of the margins in the gerontic stage.
The ornament consists chiefly of strong costae with marked growth-lines sometimes
becoming lamellar. The type and position on the valves of the growth-lines is
regarded as a specific character and can be linked with type of costation when group-
ing the species. The members of the latissima group, for instance, have numerous
subangular or rounded radiating costae with fairly distinct growth-lines at about
one-third the distance from the umbo, and approximately three to five at about
two-thirds the distance from the umbo. They tend to be closer together or more
lamellar and less prominent at the anterior margin. Bifurcation of the costae in
adult forms is rare.
In some species, notably C. antidichotoma (Buvignier), the ornament shows con-
siderable deviation from the typical. Davidson (1851, pi. 14) in the species Capil-
lirhynchia urighti (Davidson) from the Inferior Oolite, and Ager (1958 : 69) in
Furcirhynchia furcata Buckman from the Lias illustrated a similar kind of ornament.
In the genus Cyclothyris this type of ornament is regarded as a specific character.
Marked uniplication of the anterior margin is a strong feature but some species
show a tendency to produce an asymmetrical commissure. These aberrant forms
are noticeable throughout the generic range and particularly in the Cenomanian
species C. difformis.
Some species, for instance C. scaldisensis (d'Archiac), are characterized by their
large circular foramen. This is a common feature of the latissima group.
Internal characters
Pedicle valve. The umbonal cavity is roughly oval in cross-section, a pedicle
collar being developed within the first 4 mm. The cyclothyrid deltidium is seen in
transverse section as two outwardly curving plates, one on either side of the foramen.
The crenulated hinge-teeth are thick, quadrate and deeply inserted, expanding
dorsally. The diductor muscle-scars are large and triangular. The adductor muscle-
scars are too faint for adequate description.
Brachial valve. No cardinal process is developed. The hinge-plates are slender
and gently arched ventrally. Inner and outer socket-ridges are well defined. The
median septum is short and does not support the hinge-plates. Radulifer crura
originate from the distal parts of the hinge-plates and curve ventrally. Each
terminates in a Y-shaped fork which may be slightly deflected dorsally or remains
almost horizontal to the sides of the valve.
DISCUSSION. The chief distinguishing characters of Cyclothyris, apart from its
general outline, are the extensive interarea, absence of bifurcation and intercalation
THE BRACHIOPOD GENUS CYCLOTHYRIS 45
of costae in adult forms, large circular foramen and incipient fold in the brachial
valve. The arched and divided or forked hinge-plates, as seen in transverse section,
distinguish it from other Cretaceous rhynchonellid genera. It is readily distinguished
from Orbirhynchia (Pettitt, 1954) in possessing radulifer crura.
The stratigraphical zones quoted for species of Cyclothyris from the Lower Green-
sand are according to Casey (1961).
Cyclothyris latissima (]. de C. Sowerby)
(PI. 4, figs. 7, 8 ; Text-fig. 4)
1825 Terebratula lata J. de C. Sowerby (non T. lata J. Sowerby), p. 165, pi. 502, fig. i.
1829 Terebratula latissima J. de C. Sowerby, Index.
1852 Rhynchonella latissima (J. de C. Sowerby) : Davidson, p. 82, pi. n, figs. 15—22.
1918 Cyclothyris latissima (J. de C. Sowerby) : Buckman, p. 14, pi. 14, fig. ia.
1950 Cyclothyris latissima (J. de C. Sowerby) : Pettitt, pi. i, figs. 14, 15.
1956 Cyclothyris latissima (J. de C. Sowerby) : Owen, pi. 3, fig. 6.
EMENDED DIAGNOSIS. Medium sized Cyclothyris approximately 21 mm. long,
24 mm. wide and 13 mm. thick. Transversely oval to subtriangular in outline,
lenticular in anterior contour. Brachial valve convex, with broad, flat almost
imperceptible median fold. Pedicle valve less convex, with broad shallow sulcus.
Anterior commissure broadly arcuate. Umbo short, thick, slightly incurved. Fora-
men large. Deltidial plates well exposed. Each valve ornamented by about 60
rounded or subangular costellae (23 per 10 mm.) with approximately 13 on the fold
and a corresponding number in the sulcus.
LECTOTYPE. In the original description of the species J. de C. Sowerby (1825 : 165)
quotes three localities, Faringdon, Sidmouth, and Devizes Canal. The species is
represented in the Sowerby Collection, British Museum (Natural History) by three
specimens from the above localities. Of these, the specimen from Faringdon was
selected and figured as lectotype of the species by Pettitt (1950, pi. i). It is registered
in the British Museum (Natural History) collections as 6.61499. Of the two remain-
ing syntypes, one is recognizable as a young C. difformis (Valenciennes in Lamarck)
from the Cenomanian of Sidmouth, Devon, while the other is crushed and too badly
damaged for identification.
DESCRIPTION. Although C. latissima does not seem to show the same range of
variation as other species of Cyclothyris, a marked variant occurs at Faringdon,
Berkshire and at Brickhill, Buckinghamshire and reaches dimensions considerably
larger than the typical form, measuring approximately 25 mm. long, 34 mm. wide
and 18 mm. thick. The dimensions of the lectotype are 21 mm. long, 28 mm. wide
and 12 mm. thick. A fine series of specimens illustrating the above variant are to
be found in the collections of the Sedgwick Museum, Cambridge numbered SM.
6.25682-6.25708.
The outline of the shell of C. latissima remains oval to subtriangular with a broad
flat fold on the brachial valve. The ornament consists of approximately 60 costellae
on each valve, which are characteristically subangular or rounded and interrupted
by step-like or fairly steep concentric growth-lines at about one-third the distance
GEOL. 7, 2. 2§
46
THE BRACHIOPOD GENUS CYCLOTHYR1S
from the umbo and again at about two-thirds the distance from the umbo. Growth-
lines at the anterior margin tend to be more numerous, usually about 3 to 5 and are
lamellar.
Internal structure. As for genus. Text-fig. 4.
Cyclothyris latissima can be distinguished from other Aptian species of Cydothyris
by its more acutely triangular outline, more extensive interarea, less marked anterior
fold and more prominent growth-lines. It differs from C. depressa (J. de C. Sowerby)
FIG. 4. A series of fifteen transverse serial sections through the umbonal part of a
specimen of Cyclothyris latissima (J. de C. Sowerby) from the type locality, Faringdon,
Berkshire. BM. 66.5482. X2.
in its less produced beak, larger dimensions, less angular costellae and more arcuate
anterior commissure. It is less convex than other Cyclothyris species with the excep-
tion of C. scaldisensis (d'Archiac) from the Tourtia of Belgium which it resembles,
but can be distinguished from this species by its smaller dimensions, more prominent
growth-lines and less well-developed interarea and beak-ridges.
DISTRIBUTION. Apart from the type locality at Faringdon, Berkshire, where it
occurs in the Tropaeum subarcticum subzone of the Parahoplites nutfieldensis Zone,
C. latissima has also been found in the P. nutfieldensis Zone at Upware, Cambridge
and from the junction of the Hythe and Sandgate Beds on the foreshore just west
of Folkestone Harbour, Kent.
THE BRACHIOPOD GENUS CYCLOTHYRIS 47
Cyclothyris antidichotoma (Buvignier)
(PI. 4, figs. 3^6 ; Text-figs. 5, 6)
1842 Terebratula antidichotoma Buvignier, p. 533, pi. 4 fig. 8.
1847 Rhynchonella antidichotoma (Buvignier) d'Orbigny, p. 31, pi. 500, figs. 1-5.
1847 Rhynchonella antidichotoma (Buvignier) : Pictet & Roux, p. 534, pi. 50, figs. $a-g.
1872 Rhynchonella antidichotoma (Buvignier) : Pictet, p. 41, pi. 199, figs. 13-176.
1874 Terebratula antidichotoma Buvignier : Davidson, p. 65, pi. 8, figs. 19-21.
1903 Rhynchonella lineolata var. mirabilis Walker, p. 261, pi. 18, figs. ja-c.
EMENDED DIAGNOSIS. Medium to large biconvex Cyclothyris approximately
31 mm. long, 36 mm. wide and 15 mm. thick. Transversely oval to subcircular in
outline. Brachial valve with broad, faintly developed median fold. Pedicle valve
with wide, shallow sulcus, anteriorly developed. Anterior commissure broadly
arcuate to trapezoidal. Umbo massive to sharp, slightly produced. Foramen large.
Deltidial plates well exposed. Beak-ridges distinct, interarea extensive. Each valve
ornamented with between 15 and 30 fine rounded costellae posteriorly and a few
strong angular costae anteriorly.
TYPE SPECIMEN. The original specimen figured by Buvignier (1842, pi. 4, fig. 8)
cannot be traced and is presumed lost. It is not proposed to erect a neotype of this
species until more material from the type locality has been examined. The only
available specimen said to be from the area of the type locality examined by the
writer is in the d'Orbigny Collection, Museum Nationale d'Histoire Naturelle, Paris
and is registered in the d'Orbigny catalogue as 6014. This specimen, stated to be
from the Albian of Saulce, Ardennes, does not resemble Buvignier's figured specimen
in general outline although the ornament is similar. Instead it resembles one of the
more robust forms commonly found in England at Brickhill and Upware and regarded
as an extreme variant. D'Orbigny (1847, pi. 500) figured two specimens both of
which resemble the one figured by Buvignier (1842, pi. 4, fig. 8). Neither of these
specimens is in the d'Orbigny Collection and their whereabouts cannot be ascer-
tained.
DESCRIPTION. Buvignier's figure shows a large Cyclothyris measuring approximately
32 mm. long and 42 mm. wide, with a sharp beak, large foramen and ornament of
fine costellae and strong costae which are the main distinguishing features. Specimens
possessing similar features have been examined by the writer from Upware, Cam-
bridgeshire, Brickhill, Buckinghamshire and Potton and Leighton Buzzard, Bed-
fordshire. In addition collections of specimens also bearing these features from Albian
localities in France and Switzerland have been studied in the Museum d'Histoire
Naturelle, Geneva.
It is probable that the original specimen described by Buvignier came from the
Gault of Grandpre, although specimens of other species from this locality are usually
preserved as internal casts in phosphatized sandy clay. Buvignier's specimen,
however, has a well-preserved shell and may have come from the beds below the
phosphate workings which are known to be of Upper Aptian age.
In England, the Upper Aptian forms occurring at Brickhill and Upware differ
from Buvignier's figured specimen in their more circular general outline and more
48
THE BRACHIOPOD GENUS CYCLOTHYRIS
convex valves, although variants approaching the typical form have been found.
The Lower Albian forms from Leighton Buzzard, while agreeing in general outline
and proportions with Buvignier's figure, have a less acute apical angle, less produced
beak and smaller interarea and are much nearer a form which occurs at Goudiniere,
Grand St. Bernard and at Mont Saxonet, Savoie, France. This form is represented
o .0
0-2
FIG. 5. Eighteen transverse serial sections through the umbonal part of Cyclothyris
antidichotoma (Buvignier) from the Upper Aptian, Upware, Cambridge. BM. 6.25753.
X 2.
in the Davidson Collection, British Museum (Natural History) by three specimens
from Goudiniere registered as 66.41485-87. Another specimen showing a marked
resemblance to the Leighton Buzzard form comes from the Albian of Vergys, Upper
Savoie and is registered as BM. 6.35284.
Young forms of this species were thought to belong to a new variety by Walker
(1903 : 261) who described them as Rhynchonella lineolata var. mirabilis. They are
THE BRACHIOPOD GENUS CYCLOTHYRIS
49
usually more triangular in outline than the adult forms, with marked anterior sulca-
tion of both valves. The large marginal costae are less numerous at this stage and the
median fold in the brachial valve is not developed. Examples of this juvenile stage
were figured by Pictet & Roux (1847, pi. 50) and by Pictet (1872, pi. 919) as R. anti-
dichotoma (Buvignier) .
The ornament of fine costellae and coarse costae running into one another is
characteristic of the species but, as already stated, authors have described a similar
0-2
FIG. 6. Fourteen transverse serial sections through the umbonal part of Cyclothyris
antidichotoma (Buvignier) from the Lower Albian, Shenley Hill, Leighton Buzzard,
Bedfordshire. BM. 66.17561. x 2.
feature on species in the Jurassic, and less notably, on Upper Cretaceous species such
as Cretirhynchia octoplicata (J. Sowerby) as figured by Davidson (1852, pi. 10, figs,
i-n).
Although there may be sufficient grounds both morphologically and stratigraphic-
ally for separating the Upper Aptian forms from those in the Lower Albian of this
country, the writer proposes to leave this extremely well-known species as it is at
present interpreted until more material from Grandpre, or at least from the French
Ardennes, is available and the limits of variation are possible to define.
DISTRIBUTION. C. antidichotoma as at present interpreted is a common fossil in
50 THE BRACHIOPOD GENUS CYCLOTHYRIS
the Upper Aptian, P. nutfieldensis Zone, at Upware, Cambridge and Brickhill,
Buckinghamshire and occurs, though not commonly, in the Lower Albian, Leymeriella
tardefurcata Zone at Leighton Buzzard, Bedfordshire. It has also been collected
from the Albian of Mont Saxonet and Goudiniere and from the Ardennes, France.
Cyclothyris depressa (J. de C. Sowerby)
(PI. 4, fig- ii)
1825 Terebratula depressa J. de C. Sowerby, p. 165, pi. 502, fig. 3.
1852 Rhynchonella depressa (J. de C. Sowerby) Davidson, p. 89, pi. n, figs. 28-32.
EMENDED DIAGNOSIS. Small Cyclothyris about 9 mm. long, n mm. wide and 8 mm.
thick. Elongate-triangular in outline. Brachial valve convex with moderately
well-developed median fold. Pedicle valve less convex with shallow sulcus. Anterior
commissure uniplicate with trapezoidal linguiform extension. Beak suberect, apical
angle acute. Foramen large. Deltidial plates well exposed. Shell ornamented by
about 25-28 sharp, angular, radiating costae originating from the umbones, with 6
on fold and 7-8 in sulcus.
LECTOTYPE. In the original description of the species J. de C. Sowerby (1825)
figured two specimens as fig. 3 on his pi. 502. Of these the larger of the two specimens
is shown in dorsal and ventral views. This specimen is here selected as lectotype of
the species. It is in the Sowerby Collection, British Museum (Natural History) No.
6.61468, together with the second figured specimen (6.61469) and three other
syntypes (6.61470-72) which are also referable to C. depressa.
DESCRIPTION. Apart from its small dimensions, the distinctive characters of this
species are those of the beak, which is slightly produced, sharp and suberect. The
large circular foramen shows the cyclothyrid deltidial plates to advantage. The pedicle
valve has a marked trapezoidal linguiform extension. The sharp, angular costae
show no signs of bifurcation or intercalation. They are interrupted only by a faint
growth-line which appears at about two-thirds the distance from the umbo to anterior
margin.
Internal structure. As for genus.
REMARKS. C. depressa has been grouped with C. schloenbachi (Davidson) since the
two species have much in common. Their general outline and profile is roughly the
same with a well-defined fold and sulcus. The linguiform extension is trapezoidal
in shape. Both species are relatively small and seem to show the same extremes of
variation with fine and coarse costation. C. depressa, however, is distinguished from
C. schloenbachi by its more acutely triangular outline, slightly produced beak and
shallower sulcus in the pedicle valve and well-developed fold on the brachial valve.
It differs from C. latissima in its general triangular outline, angular costation, pro-
duced beak, smaller dimensions and trapezoidal linguiform extension.
DISTRIBUTION. C. depressa is commonly found in association with C. latissima in
the Upper Aptian (P. nutfieldensis Zone) of Faringdon, Berkshire and also at the same
horizon at Brickhill, Buckinghamshire.
THE BRACHIOPOD GENUS CYCLOTHYRIS 51
Cyclothyris difformis (Valenciennes in Lamarck)
(PI. 5, figs. 1-7 ; Text-figs. 7, 8)
1819 Terebratula difformis Valenciennes in Lamarck, p. 255, fig. indicated Encycl. Meth.,
1789, pi. 242, fig. 5.
Terebratula dimidiata J. Sowerby, p. 138, pi. 277, fig. 5.
Terebratula gallina Brongniart, p. 84, pi. 9, fig. 2.
Terebratula deformis Lamarck : Defrance, p. 160, pi. 5, fig. 3.
Rhynchonella compressa (Lamarck) : d'Orbigny, p. 35, pi. 497, figs. 1-5.
Terebratula difformis Lamarck : Davidson, p. 433, pi. 15, fig. 48.
Terebratula compressa Lamarck : Davidson, p. 80, pi. n, figs. 4, 5.
Terebratula difformis Lamarck : Quenstedt, p. 696, pi. 54, fig. 2.
1900 Rhynchonella difformis (Lamarck) Jukes-Browne, p. 65, figs. 41, 42.
1918 Terebratula difformis Valenciennes in Lamarck : Clerc & Favre, pi. 15, fig. 84.
EMENDED DIAGNOSIS. Medium-sized Cyclothyris approximately 23 mm. long,
31 mm. wide and 17 mm. thick. Shell biconvex, fold low, indistinct, oval to sub-
triangular in outline. Anterior commissure with well-marked uniplication. Umbo
short, massive, slightly incurved. Foramen fairly large. Deltidial plates conjunct,
well exposed. Beak-ridges distinct. Interarea extensive. About 40-45 costae on
either valve (14 per 10 mm.) with 9 on fold and 10-11 in sulcus.
LECTOTYPE. Owing to Lamarck's blindness the brachiopods in his " Animaux
sans Vertebres " (vol. 6, 1819) were described by his pupil A. Valenciennes from
specimens in Lamarck's own collection. In the description of Terebratula difformis
(p. 255) Valenciennes indicated a specimen illustrated in Encyclopedic Methodique
(1789, pi. 242, fig. 5) quoting the localities " near Le Mans, and also at Cap la Heve,
near Le Havre " both Cenomanian localities. No type specimen was named or
indicated and the whereabouts of the figured specimen is not known.
Davidson (1850 : 433) in a description of Lamarck's species states that he bor-
rowed the ten specimens used by Valenciennes in his original description and figured
one of them as T. difformis (pi. 15, fig. 48). This would have served as lectotype of
the species had the specimen not been subsequently lost or mislaid. A further
specimen figured by Clerc & Favre (1918, pi. 15, figs. 8^a-d) has therefore been
selected as lectotype of the species. This specimen is one of eight to be found under
the name Terebratula difformis in the Lamarck Collection at the Museum d'Histoire
Naturelle, Geneva and is registered as No. 48 in the Lamarck catalogue. From the
mode of preservation and adhering matrix it was almost certainly collected from the
Lower Cenomanian of the Normandy coast. The other seven specimens are probably
of Jurassic age. The dimensions of the lectotype are : 23 mm. long, 31 mm. wide and
17 mm. thick.
DESCRIPTION. C. difformis is an extremely variable species ranging in outline from
subcircular to transversely oval to subtriangular. The ornament usually consists of
about 45 coarse to relatively fine costae originating from the umbo of each valve and
becoming more elevated towards the anterolateral margins. These costae are usually
interrupted by one or two step-like growth-lines situated at about one-half to two-
thirds the distance from the umbo to the anterior margin.
The umbo varies from short to massive to slightly produced. An extensive interarea
52 THE BRACHIOPOD GENUS CYCLOTHYRIS
with distinct beak-ridges is seen throughout the range of variation. The median fold
on the brachial valve remains indistinct while the broad, shallow sulcus in the pedicle
valve is late in development. The linguiform extension varies from broadly arcuate
to trapezoidal but asymmetry of the anterior commissure is a notable character.
Variants worthy of special note occur at Wilmington, south Devon and at Cap la
Heve, Normandy. One of the Wilmington varieties departs from the typical in
having more convex valves, shorter umbo, smaller foramen and more acutely arched
FIG. 7. Diagram illustrating thirteen variants of Cyclothyris difformis (Valenciennes in
Lamarck) from Wiltshire, south Devonshire, and Normandy.
anterior commissure. The variety usually found on the Normandy coast, on the
other hand, has a slightly produced beak, less convex valves and commonly exhibits
an asymmetrical anterior commissure.
Another notable variant occurs in the Lower Cenomanian, Tourtia, Belgium. Six
well-preserved specimens in the British Museum (Natural History) are numbered
66.41475-80. The same variety occurs in the Lower Cenomanian deposits of Essen,
North Germany and is represented in the same museum by a single specimen (BB.
4I473)-
The specimen figured and described as Terebratula dimidiata by J. Sowerby (1821 :
138, pi. 277, fig. 5) from Halldown near Chudleigh, south-east Devon, is a large,
almost sphaeroidal rhynchonellid with coarse costae, sharp, produced beak and
THE BRACHIOPOD GENUS CYCLOTHYRIS 53
asymmetrical anterior commissure. This is either an extreme variant of C. difformis
or a pathological form.
A form somewhat resembling the outline of Sowerby's T. dimidiata but with finer
costae, is figured by d'Orbigny (1847, pi- 49$, figs. 6-9) from the Charentes as
Rhynchonella difformis. Some authors have followed d'Orbigny 's interpretation of
Lamarck's species and Mile G. Fage (1935 : 433, pi. n), in a description of some
Upper Cretaceous rhynchonellids from the Charentes, illustrated an evolutionary
development from R. difformis d'Orbigny to a variety R. globata Fage. Her figured
specimen of R. difformis is stated to have been collected from the Coniacian.
A specimen figured by J. de C. Sowerby (1836, pi. 18, fig. 2) as T. dilatata was stated
in the explanation of the plates to have been collected from Blackdown, south Dorset.
FIG. 8. A series of twelve transverse serial sections through the umbonal part of Cyclo-
thyris difformis (Valenciennes in Lamarck) from the lower Cenomanian, Cap la Heve,
near Le Havre, Normandy. BM. 6.35224. x 2.
This figure shows a remarkable similarity to a specimen figured by Schloenbach
(1867, pi. 23, fig. 2) as C. difformis from the Lower Cenomanian of Essen, Germany.
DISTRIBUTION. C. difformis ranges from the Lower to Middle Cenomanian of the
west of England, Normandy and North Germany. In the type area of the Normandy
coast it occurs in the remanie bed at the base of the Cenomanian, is abundant in the
Lower Cenomanian and is less common, but still well represented in the Middle
Cenomanian.
In England the species is almost unrepresented in the Middle Cenomanian though
rare examples have been found in the Middle Cenomanian basement beds of west
Dorset. It is abundant, however, in the Lower Cenomanian, occurring in Wiltshire
(Warminster Greensand) and the Devon coast, as well as in the Isle of Wight. A
well-marked variant is the commonest brachiopod in the sandy Lower Cenomanian
54 THE BRACHIOPOD GENUS CYCLOTHYRIS
of Wilmington, south Devon. It is said to occur in the dispar Zone at Punfield Cove,
south Dorset but specimens of this species are more likely to have been collected
from the bottom Cenomanian beds. Its occurrence below this level is uncertain.
Cyclothyris compressa (Valenciennes in Lamarck)
(PL 5, ng. 8)
1819 Terebratula compressa Valenciennes in Lamarck, p. 256, No. 54.
1850 Terebratula compressa Lamarck : Davidson, p. 455, pi. 15, fig. 54.
1918 Terebratula compressa Valenciennes in Lamarck : Clerc & Favre, pi. 19, figs. 117, 119.
EMENDED DIAGNOSIS. Cyclothyris, about 28 mm. long, 32 mm. wide and 14 mm.
thick. Distinctly subtriangular in general outline ; brachial valve convex with faint
median fold. Pedicle valve posteriorly convex with broad, shallow median sulcus.
Both valves ornamented by about 40 strong, acutely angular, radiating costae,
(8 per 10 mm.) with 8 on fold and a corresponding number in sulcus. Concentric
growth-lines very faint. Beak short, slender, slightly incurved. Beak-ridges distinct,
interarea small. Deltidial plates triangular, foramen small.
LECTOTYPE. The specimen here selected as lectotype of the species is one of four
syntypes in the Lamarck Collection at the Museum d'Histoire Naturelle, Geneva
and is numbered in the Lamarck Catalogue as No. 54. The specimen was figured by
Davidson (1850, pi. 15, fig. 54) and by Clerc & Favre (1918, pi. 19, fig. 117). It was
collected from the Upper Cenomanian of Coulaines near Le Mans in the Sarthe,
south-west of the Paris basin. The dimensions of the lectotype are : 29 mm. long,
35 mm. wide and 14 mm. thick.
DESCRIPTION. C. compressa is one of the most distinctive species of Cyclothyris.
It is characterized by its acutely subtriangular outline and strong angular costae.
It can be distinguished by its low, faintly developed median fold, trapezoidal lingui-
form extension, and marginal thickening of the valves. It is probable that this
species is a direct development of C. difformis. A specimen from the Tourtia of
Belgium, showing characters of both species is in the British Museum (Natural
History) numbered BM. 6.1889. Another specimen figured by Davidson (1852,
pi. n, figs. 4, 5) as C. compressa and figured here on PL 5, fig. 8 as C. difformis shows
the general outline of C. difformis with the anterior and lateral contours and sharp
angular costae of C. compressa but without the marginal thickening of the valves.
DISTRIBUTION. The species, originally described from Coulaines near Le Mans in
the Sarthe, France is confined to the sandy Cenomanian of the south-west Paris
basin. The typical form is characteristic of the Upper Cenomanian, Sables du Perche,
where it is the only rhynchonellid present. Related but not identical forms occur in
the beds above (Marnes a Ostrea biauriculata) and below (Sables et Gres du Mans).
Varieties, stated to belong to this species, were described by Cayeux (1949) from
Le Havre but were not accompanied by any adequate description or illustration,
nor were any type-specimens indicated.
THE BRACHIOPOD GENUS CYCLOTHYRIS 55
Cyclothyris schloenbachi (Davidson)
(PI. 4, figs. 12, 13)
1852 Rhynchonella depressa vars. A and B, Davidson, p. 92, pi. 12, figs. 28-30.
1874 Rhynchonella schloenbachi Davidson, p. 59.
EMENDED DIAGNOSIS. Small Cyclothyris about n mm. long, 15 mm. wide and 13 mm
thick. Shell outline oval, brachial valve convex, slightly inflated, with well-defined
median fold. Pedicle valve less convex with shallow sulcus broadening anteriorly.
Foramen medium sized to small. Beak-ridges distinct, interarea fairly long. Orna-
ment varying from fine to coarse costae.
LECTOTYPE. C. schloenbachi was first described by Davidson (1874 : 59) who
included three specimens which he had previously described and figured (1852 : 92,
pi. 12, figs. 28-30) as varieties A and B of Rhynchonella depressa (]. de C. Sowerby),
an Upper Aptian species. Two of these specimens (figs. 28, 29) were collected from
the Cenomanian of Chardstock, Somerset, BM. 6.8215, and the third (fig. 30) from
the Cenomanian of Shaftesbury, Wiltshire. Of these syntypes, the specimen from
Shaftesbury (BM. 6.8216), figured by Davidson (1852, pi. 12, fig. 30) is here selected
as lectotype of the species.
DESCRIPTION. Cyclothyris schloenbachi is a common fossil is the Lower Cenomanian
beds of south Devon, Somerset, south Wiltshire, and Dorset. As can be seen from
Davidson's original figures (1852, pi. 12, figs. 28-30) the ornament varies from fine,
subangular to strong, coarse costae. Davidson himself (1874 : 59) noted this varia-
tion and stated that intermediate forms were commonly found with the extreme
forms.
The median fold is better developed or more distinct on specimens with finer costae
and is often replaced in the coarser costate forms by a slight sulcation of the brachial
valve.
Bifurcation of the costae is a fairly common feature of the young coarsely costate
forms. Both variants occur with intermediate forms in the same beds at Chardstock
in Somerset, Beer Head, Devon, and Pinhay Cliff, Dorset.
Because of its comparatively small dimensions C. schloenbachi cannot easily be
confused with other known Cyclothyris species. It is distinguished from the somewhat
similar C. depressa. by its oval outline, less produced beak, smaller foramen and better
developed or deeper sulcus in the pedicle valve.
DISTRIBUTION. In England, C. schloenbachi appears to be confined to the Lower
Cenomanian beds of south Devon, Dorset, Somerset and Wiltshire. A slightly larger
form of the same species occurs in the Lower Cenomanian beds of Vivautier in the
department of Orne, France and is represented in the Davidson Collection, British
Museum (Natural History) by fifteen specimens numbered 6.11917. These specimens
show variations identical with those of the English forms.
Cyclothyris lepida sp. n.
(PI. 4, fig. 10)
DIAGNOSIS. Cyclothyris about 22 mm. long, 32 mm. wide and 16 mm. thick. Shell
biconvex, transversely oval in outline. Brachial valve with poorly developed median
56 THE BRACHIOPOD GENUS CYCLOTHYRIS
fold. Pedicle valve with broad shallow sulcus. Both valves ornamented by about
60-65 fine rounded costellae (21 per 10 mm.) with 20 on the fold and about 22 in the
sulcus. Anterior contour lenticular. Linguiform extension trapezoidal, moderately
developed. Beak prominent with large foramen.
HOLOTYPE. SM. 6.25683 from the Upper Aptian, Brickhill, Buckinghamshire in
the collections of the Sedgwick Museum, Cambridge.
Dimensions of holotype. 26 mm. long, 33 mm. wide and 16 mm. thick.
PARATYPES. Twenty-two specimens from the type locality in the Sedgwick Museum
SM. 6.25682, 6.25684-87, 6.25689-702, 6.25706-08. Also 40 specimens from the
same locality and horizon in the 6ritish Museum (Natural History), 6.25546,
6.25547, 5.25549.
DESCRIPTION. C. lepida is a medium-sized Cyclothyris with a characteristic orna-
ment of fine radiating costellae originating from the umbones of each valve. The
absence of any prominent growth-lines gives the species a neat appearance. The
beak is strong and slightly incurved. An extensive interarea is bounded by distinct
beak-ridges. The foramen is large and the deltidial plates well exposed.
Internal characters. As for genus.
REMARKS. This species can be distinguished from others of Cyclothyris mainly
by its distinctly oval outline, fine costellae, regular anterior commissure, moderately
developed but well-marked median fold on the brachial valve, trapezoidal linguiform
extension and absence of marked growth-lines. It occurs with C. antidichotoma
(6uvignier) at 6rickhill but its ornament shows none of the characteristics of this
species. It is probably the species referred to by authors as Rhynchonella lata
d'Orbigny (1847 : 21) from the Neocomian of France.
6ecause if its regular outline, well-marked median fold and absence of prominent
growth-lines, C. lepida is grouped with C. depressa and C. schloenbachi.
DISTRIBUTION. C. lepida appears to be confined to the Upper Aptian of 6rickhill,
6uckinghamshire.
Cyclothyris levis sp. n.
(PI. 4, fig- 9)
DIAGNOSIS. Cyclothyris about 20 mm. long, 24 mm. wide and 12 mm. thick. Shell
biconvex, distinctly subtriangular in outline. No median fold developed on brachial
valve. Shallow, broad sulcus developed anteriorly in pedicle valve. 6oth valves
covered by about 100 very fine capillae (48 per 10 mm.). 6eak short, massive,
foramen small. 6eak-ridges distinct. Interarea small. Deltidial plates not well
exposed.
HOLOTYPE. 66.41493 in 6ritish Museum (Natural History) from the Lower
Albian (L. tardefurcata Zone) of the Shenley Limestone, Leighton 6uzzard, 6edford-
shire. Dimensions of holotype are : 22 mm. long, 27 mm. wide and 14 mm. thick.
PARATYPES. Sixty-one specimens in 6ritish Museum (Natural History) registered
Nos. 6.26541, 6.26542, 6.26595. .
DESCRIPTION. In general outline this species resembles C. latissima but has a
much more inflated brachial valve. A broad, shallow sulcus develops late in the
THE BRACHIOPOD GENUS CYCLOTHYRIS 57
pedicle valve giving rise to a low broadly arcuate anterior commissure. The short,
slightly incurved beak is sharp and massive with a small circular foramen. Distinct
beak-ridges define a small interarea. Faint concentric growth-lines are seen on both
valves and appear more distinct towards the anterolateral margins.
Internal characters. As for genus.
REMARKS. C. levis is distinguished from other Cyclothyris species by its extremely
fine, rounded capillae, short massive beak, absence of median fold in the brachial
valve and poorly developed sulcus in the pedicle valve. It can be distinguished
from C. lepida sp. n. mainly by its general subtriangular outline, fine capillae, absence
of median fold, shorter more massive beak, less extensive interarea, poorly exposed
deltidial plates and broad arcuate anterior commissure. Although resembling
C. latissima in general outline it can be distinguished from this species by its less
extensive interarea, less produced beak, finer ornament of capillae, less marked
growth-lines and broader arcuate anterior commissure.
DISTRIBUTION. Cyclothyris levis is confined to the Lower Albian and is a common
fossil in the limestone lenticles (L. tardefurcata Zone) at Leighton Buzzard, Bedford-
shire.
Cyclothyris scaldisensis (d'Archiac)
(PI. 4 fig. i)
1846 Terebratula scaldisensis d'Archiac, p. 330, pi. 21, fig. n.
EMENDED DIAGNOSIS. Cyclothyris about 24 mm. long, 32 mm. wide and 16 mm.
thick. Shell biconvex, oval to subtriangular in outline, brachial valve with broad
incipient median fold. Broad flattened median sulcus in pedicle valve. Umbo
short, massive, slightly incurved. Foramen large, circular. Deltidial plates well
exposed. Costae fine, rounded, numbering about 68 on each valve (12 per 10 mm.)
with 15-18 on the fold and a corresponding number in the sulcus.
HOLOTYPE. The specimen used by d'Archiac in his description of the species was
stated to have been collected from the Tourtia of Belgium. It is not certain whether
this specimen is still extant and enquiries are still being made. Several well-preserved
specimens from the Tourtia of Belgium are to be found in the Davidson Collection
and in the general collections of the British Museum (Natural History). Two good
examples of this species are from Tournai and are registered 6.1889, 66.41492.
Another specimen, larger than the dimensions given for the typical form, is figured
here on PI. 4, fig. i.
DESCRIPTION. C. scaldisensis is medium sized, characteristically oval to broadly
subtriangular in outline. The brachial valve is convex and slightly inflated posteriorly
with a very faintly developed median fold. The pedicle valve is convex in the umbonal
region but develops a shallow sulcus which broadens anteriorly. The linguiform
extension is moderately developed giving rise to a low broad, arcuate anterior
commissure. Both valves are ornamented by about 65-68 fairly fine, rounded costae
radiating from the umbones. Faint lamellar growth-lines are seen at about half and
again at two-thirds the distance from the beak to the anterior margin. They tend to
become more prominent anteriorly. The short umbo is massive with a large circular
58 THE BRACHIOPOD GENUS CYCLOTHYRIS
foramen and well-exposed deltidial plates. The beak-ridges are distinct and define
a faintly concave extensive interarea.
This is probably the least variable of all species of Cyclothyris. Its characteristically
fine, rounded costae and faint concentric growth-lines make it easily distinguishable
from C. difformis, with which it is often associated. It differs from C. latissima, from
which it was most probably developed, by its less prominent growth-lines, longer or
more extensive interarea and less incurved beak.
FIG. 9. Fourteen transverse serial sections through the umbonal part of Cretirhynchia
norvicensis Pettitt from the Upper Chalk (mucronata Zone), Mousehold Pit, Norwich.
BM. 6.25079. x 2.
DISTRIBUTION. Apart from the area of its original description in the Lower Ceno-
manian, Tourtia, of Belgium, the species is recorded from the Lower Cenomanian
of Honfleur, Cap la Heve and Fecamp, Normandy and from a similar horizon in
the Munster basin at Essen, Germany. It also occurs, though not commonly, in the
Lower Cenomanian beds of Wiltshire, and the south Devon coast.
Genus BURRIRHYNCHIA nov.
DIAGNOSIS. Small to medium biconvex rhynchonellidae. Usually elongate-
triangular in general outline. Umbo massive, suberect ; beak-ridges rounded.
THE BRACHIOPOD GENUS CYCLOTHYRIS
59
Foramen small, circular, hypothyrid. Deltidial plates disjunct, cyclothyrid. Anterior
commissure with broad trapezoidal linguiform extension Faint fold on brachial
valve. Hinge plates narrow, ventrally arched. Median septum strong, persistent.
Ornament of fine, rounded costellae (more than 50 on each valve).
TYPE SPECIES. " Rhynchonella " leightonensis Walker, 1903.
HOLOTYPE. From Lower Albian (L. tardefurcata Zone), Shenley Hill, Leighton
Buzzard, Bedfordshire, No. GSM. 51279 in the Geological Survey Museum.
FIG. 10. A series of thirteen serial sections through the umbonal part of Burrirhynchia
cantabrigensis (Davidson) from the Upper Aptian of Upware, Cambridge. B.M.
B.2574Q. x 2.
REMARKS. The genus differs from Cyclothyris in its more elongate-triangular
outline, strong persistent median septum, small foramen, rounded beak-ridges, and
narrow hinge-plates which are not terminally divided. In transverse outline it
resembles Sulcirhynchia Burri (1953) from the Swiss Neocomian, from which it may
have been developed. It can be distinguished from this genus, however, by its
stronger, more persistent, median septum, narrower, less acutely ventrally deflected
hinge-plates, shorter, less deeply inserted hinge-teeth and more clearly defined inner
and outer socket-ridges.
Burrirhynchia almost certainly gave rise to Cretirhynchia Pettitt, 1950, from the
Upper Chalk but is distinguished from it by its subparallel dental lamellae, disjunct
60 THE BRACHIOPOD GENUS CYCLOTHYRIS
deltidial plates, poorly defined interarea and absence of bifurcation of costellae and
thicker, less clearly defined hinge-plates.
RANGE. Upper Aptian to Lower Albian.
SPECIES ASSIGNED. " R." leightonensis Walker, " R." cantabrigensis Davidson,
" R." gibbsiana (]. de C. Sowerby), " R." bertheloti d'Orbigny. " R." tripartita
Pictet.
MATERIAL AND LOCALITIES. One hundred and sixty specimens of B. leightonensis
(Walker) from the Lower Albian (L. tardefurcata Zone), Shenley Hill, Leighton
Buzzard, Bedfordshire (6.26524-28, 6.26595), 57 specimens of B. cantabrigensis
(Davidson) from the Upper Aptian, of Upware, Cambridge and 85 specimens of the
same species from Brickhill, Buckinghamshire and one specimen of " R" gibbsiana
(J. de C. Sow.) from Folkestone, Kent, all in the British Museum (Natural History).
CONCLUSION
Although, externally, Cyclothyris shows considerable variation both in size, outline
and ornament, internally it remains fairly constant. Within certain limits internal
variation can be confined to the development of the hinge-plates, dental lamellae
and the septum in the brachial valve. In the typical form from the Upper Aptian
the dental lamellae are postero-anteriorly shorter than those of the Albian and
Cenomanian forms and the hinge-plates in the Upper Aptian and Albian forms tend
to be more acutely deflected towards the ventral valve than in the Cenomanian
species, though this may be a variable character.
Some authors, such as Makridin (1955) and Dacque (1934) have suggested that
the genus ranges from the Upper Jurassic to Cretaceous. Their definition of the
genus is most probably based on Central European material which may have con-
tained early ancestral forms.
The Neocomian genus Lepidorhynchia Burri (1956) certainly has both external
and internal characters in common with Cyclothyris. The cyclothyrid deltidial plates
are already evident and the ornament of subangular costae shows a tendency to
marginal dichotomy, a character observed in very young specimens of C. latissima
from Faringdon. Also, in Lepidorhynchia, are seen the ventrally curved and termin-
ally divided hinge-plates which, when further developed in Cyclothyris, form the main
distinguishing characters allowing the genus to be separated from Sulcirhynchia
and Lamellaerhynchia. Both these genera, however, have a great deal in common with
Cyclothyris and have obviously been developed from the same original stock.
It was at first thought that Cyclothyris gave rise to the Upper Cretaceous genus
Cretirhynchia but, in the light of present knowledge, this theory is no longer tenable.
In his description of the genus Cretirhynchia, Pettitt (1950) remarked on the differences
between his genus from the Upper Chalk, and the Lower Cretaceous genus Cyclothyris.
In the same work (p. n, text-fig. 4) he figured a series of serial sections of the type-
species C. plicatilis (J. Sowerby) from Northfleet, Kent showing the strong converging
dental lamellae, short hinge-plates persistent septum and broad radulifer crura.
Recently, the writer, has prepared several serial sections of other species of Cretirhyn-
chia from Norfolk and Kent which compare favourably with Pettitt's illustrations.
These sections show quite clearly that Cretirhynchia is not a direct development of
THE BRACHIOPOD GENUS CYCLOTHYRIS 61
Cyclothyris but is closely related to early Cretaceous forms represented by the species
" Rhynchonella " cantabrigensis Davidson from Upware and Brickhill and " R."
leightonensis Walker from the Shenley Limestone. Both these species are here
referred to Burrirhynchia gen. nov.
A comparison between the serial sections of Burrirhynchia cantabrigensis from
Upware (Text-fig. 10) and Cretirhynchia norvicensis Pettitt from Norfolk (Text-fig. 9)
shows that the two have much in common. Neither species has a pedicle collar ;
the outline in transverse section is similar ; both have narrow thickened hinge-plates.
The median septum in the brachial valve of both forms persists for well over one
half the length of the shell. In addition the broad radulifer crura are given off
dorsally from similarly shaped crural bases. Externally also the two forms are much
alike. Both have a short, massive umbo with small foramen and ornament of fine,
rounded costae. Neither species shows any of the distinguishing characters of
Cyclothyris nor is there any tendency to asymmetry of the anterior margin, a common
feature of some Cyclothyris species.
It is probable, therefore, that Cretirhynchia developed from such early Cretaceous
forms as Burrirhynchia cantabrigensis and B. leightonensis which marked the beginning
or early stages and C. norvicensis the last or later stages of its evolution. Both
B. cantabrigensis and B. leightonensis are closely related to Sulcirhynchia Burri, from
the Neocomian of Switzerland.
Cyclothyris probably died out in the Upper Cenomanian with C. compressa (Valen-
ciennes in Lamarck), though Turonian species such as " Rhynchonella " vespertilio
(Brocchi) which may have been developed from C. compressa and " R." elegans
Hanstein, from the Ciply Chalk, have still to be investigated.
ACKNOWLEDGMENTS
I am greatly indebted to Dr. E. I. White, F.R.S., Keeper of Palaeontology, British
Museum (Natural History) for permission to work on the collections in his department
and to Dr. H. M. Muir-Wood for her kind help and encouragement, and for reading
the manuscript.
My thanks are also due to Dr. F. Burri, Basel ; to Mme. A. Schnorf, Musee Geolo-
gique, Lausanne, Switzerland ; to Mr. A. G. Brighton and Dr. Colin Forbes, Sedgwick
Museum, Cambridge ; to Mr. K. J. Evans, King's College, London and to Dr. VV. T.
Dean and Mr. F. M. Wonnacott, British Museum (Natural History), London.
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PLATE 4
FIGS. ia-c. Cyclothyris scaldisensis (d'Archiac). Tourtia, Gussignies, Belgium. BM.
B.I 1965.
FIGS. za-c. Cyclothyris dimidiata (J. Sowerby) = C. difformis. Cenomanian, Halldown,
south-east Devon. BM. 6.61466.
FIGS. 3«-c. Cyclothyris antidichotoma (Buvignier). Lower Albian, Shenley Hill, Leigh ton
Buzzard, Bedfordshire. BM. 66.41495.
FIGS. ^a-c. Cyclothyris antidichotoma (Buvignier). Lower Albian, Goudiniere, Grand St.
Bernard, Savoie, France. BM. 66.41486.
FIGS. $a-c. Cyclothyris antidichotoma (6uvignier). Upper Aptian, 6rickhill, 6uckingham-
shire. 6M. 66.41490.
FIGS. 6a-c. Cyclothyris antidichotoma (6uvignier). Upper Aptian, Upware, Cambridge.
6M. 6.27264.
FIGS. ja-c. Cyclothyris latissima (J. de C. Sowerby). Upper Aptian, Faringdon, 6erkshire.
8M. 66. 41494.
FIGS. Sa-c. Cyclothyris latissi ma (J. deC. Sowerby). Lectotype. Upper Aptian, Faringdon,
6erkshire. 6M. 6.61499.
FIGS. ga-c. Cyclothyris levis sp. n. Holotype. Lower Albian, Shenley Hill, Leigh ton
6uzzard, 6edfordshire. 6M. 66.41493.
FIGS. ioa-c. Cyclothyris lepida sp. n. Holotype. Upper Aptian, 6rickhill, 6uckingham-
shire. Sedgwick Museum Coll. 6.25683.
FIGS. na-c. Cyclothyris depressa (J. de C. Sowerby). Lectotype. Upper Aptian, Faring-
don, 6erkshire. 6M. 6.61468.
FIGS. I2a-c. Cyclothyris schloenbachi (Davidson). Lectotype. Lower Cenomanian, Shaftes-
bury, Wiltshire. 6M. 6.8216.
FIGS. i$a-c. Cyclothyris schloenbachi (Davidson). Coarsely costate variant from Chard-
stock, Somerset. 6M. 6. 8215.
All figures at natural size unless otherwise stated.
6M. — 6ritish Museum (Natural History).
4 JUN196?
Bull. B.M. (N.H.) Geol. 7, 2
PLATE 4
6a
PLATE 5
FIGS. la-c. Cyclothyris difformis (Valenciennes in Lamarck). Cenomanian, Warminster,
Wiltshire. Typical form. B.M. no. 45336.
FIGS. 2a-c. Cyclothyris difformis (Valenciennes in Lamarck) . Lower Cenomanian, Wilming-
ton, south Devon. BM. 36.41433.
FIGS. 3«-c. Cyclothyris difformis (Valenciennes in Lamarck). Lower Cenomanian, Cap le
Heve, near Le Havre, Normandy BM. 6.82754.
FIGS. 4«-c. Cyclothyris difformis (Valenciennes in Lamarck). Lower Cenomanian, Wilming-
ton, south Devon. BM. 66.15292.
FIGS. 5a— c. Cyclothyris difformis (Valenciennes in Lamarck). Tourtia, Tournai, Belgium.
BM. 66.41476.
FIGS. 6a-c. Cyclothyris difformis (Valenciennes in Lamarck). Cenomanian, Essen, Germany.
Similar variant to fig. 5 from Belgium. 6M. 66.41473.
FIGS. ja-c. Cyclothyris difformis (Valenciennes in Lamarck). Lower Cenomanian, War-
minster, Wiltshire. 6M. 6.8301.
FIGS. Sa-c. Cyclothyris compressa (Valenciennes in Lamarck). Upper Cenomanian, Le
Mans, Sarthe, France. BM. 66.41489.
Bull. B.M. (N.H.) Geol. 7, 2
PLATE 5
la
Ic
2c
2a
8c
THE TRILOBITES OF THE CARADOC
SERIES IN THE CROSS FELL INLIER
OF NORTHERN ENGLAND
W. T. DEAN
BULLETIN OF
THE BRITISH MUSEUM (NATURAL HISTORY)
GEOLOGY Vol. 7 No. 3
LONDON: 1962
THE TRILOBITES OF THE CARADOC SERIES
IN THE CROSS FELL INLIER OF NORTHERN
ENGLAND
BY
WILLIAM THORNTON DEAN
Pp. 65-134 ; Pis. 6-1 8 ; 5 Text-figures
BULLETIN OF
THE BRITISH MUSEUM (NATURAL HISTORY)
GEOLOGY Vol. 7 No. 3
LONDON: 1962
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.
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within one calendar year.
This paper is Vol. 7, No. 3 of the Geological
(Palaeontologicai) series.
Trustees of the British Museum 1962
PRINTED BY ORDER OF THE TRUSTEES OF
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Issued August 1962 Price Forty-two Shillings
THE TRILOBITES OF THE CARADOC SERIES
IN THE CROSS FELL INLIER OF NORTHERN
ENGLAND
By WILLIAM THORNTON DEAN
Page
I. INTRODUCTION AND ACKNOWLEDGMENTS ...... 69
II. STRATIGRAPHICAL SUMMARY ........ 69
III. SYSTEMATIC DESCRIPTIONS ........ 72
Family Raphiophoridae Angelin . . . . . .72
Lonchodomas swindalensis sp. nov. ..... 72
Lonchodomas pennatus (La louche) ..... 78
Family Trinucleidae Hawle & Corda ..... 79
Subfamily Cryptolithinae Angelin ...... 79
Broeggerolithus nicholsoni (Reed) ..... 79
Broeggerolithus melmerbiensis sp. nov. . . . . .81
Broeggerolithus cf. transiens (Bancroft) .... 82
Broeggerolithus sp. . . . . . . .83
Onnia gracilis (Bancroft) ....... 84
Onnia superba (Bancroft) pusgillensis Dean .... 84
Subfamily Tretaspidinae Whittington ..... 85
Tretaspis cf. ceriodes donsi St0rmer ..... 85
Tretaspis convergens Dean . . . . . . .85
Tretaspis kiaeri St0rmer radialis Lament .... 86
Tretaspis kiaeri St0rmer duftonensis Dean . . . .86
Family Cheiruridae Salter ....... 86
Subfamily Cyrtometopinae Opik ...... 86
Pseudosphaerexochus cf. octolobatus (M'Coy) .... 87
Family Encrinuridae Angelin ....... 88
Subfamily Encrinurinae Angelin ...... 88
Encrinurus sp. ........ 88
Subfamily Dindymeninae Pfibyl ...... 89
Dindymene duftonensis sp. nov. ...... 89
Dindymene sp. ......... 90
Subfamily Cybelinae Holliday . . . . . .91
Atractopyge scabra sp. nov. . . . . . .91
Atractopyge? sp. ........ 93
Paracybeloides cf. girvanensis (Reed) ..... 94
Paracybeloides sp. ........ 95
Family Dalmanitidae Reed ....... 95
Subfamily Dalmanitinae Reed ...... 95
Dalmanitina mucronata (Brongniart) matutina subsp. nov. . 95
Subfamily Acastinae Delo . . . . . . -97
Kloucekia (Phacopidina) apiculata (M'Coy) ... 97
Duftonia lacunosa Dean ....... 97
GEOL. 7, 3. 3
68 THE TRILOBITES OF THE CARADOC SERIES
Page
Family Pterygometopidae Reed ...... 98
Subfamily Pterygometopinae Reed ..... 98
Calyptaulax planiformis sp. nov. ..... 98
Estoniops alifrons (M'Coy) ...... 100
Subfamily Chasmopsinae Pillet . . . . . .104
Chasmops cf. extensa (Boeck) . . . . . .104
Chasmops aff. maxima (Schmidt) ..... 105
Chasmops sp. ......... 105
Family Homalonotidae Chapman . ; . . . . .106
Brongniartella minor (Salter) . . . . . .106
Brongniartella ascripta (Reed) . . . . . .106
Brongniartella bisulcata (M'Coy) . ' . . . . . 108
Brongniartella depressa sp. nov. ...... 108
Brongniartella aff. platynota (Dalman) . . . .no
Family Calymenidae Burmeister . . . . . .in
Flexicalymene cf. caractaci (Salter) . . . . .114
Onnicalymene onniensis (Shirley) . . . . .115
Onnicalymene laticeps (Bancroft) . . . . .115
Diacalymene cf. marginata Shirley . . . . .116
Gravicalymene jugifera sp. nov. . . . . . .116
Family Dimeropygidae Hupe . . . . . . .118
Toernquistia aff. reedi Thorslund . . . . .118
Family Illaenidae Hawle & Corda . . . . . .120
Stenopareia? sp. . . . . . . . .120
Family Lichidae Hawle & Corda . . . . . .120
Subfamily Homolichinae Phleger . . . . . .120
Conolichas melmerbiensis (Reed) . . . . .120
Platylichas cf. laxatus (M'Coy) . . . . . .121
Family Odontopleuridae Burmeister . . . . . .122
Primaspis semievoluta (Reed) ...... 122
Odontopleurid gen. et sp. ind. ...... 123
Family Otarionidae R. &. E. Richter . . . . .123
Otarion sp. ......... 123
Family Proetidae Salter . . . . . . . .124
Subfamily Proetidellinae Hupe . . . . . .124
Proetidella? marri sp. nov. ...... 124
Family Remopleurididae Hawle & Corda . . . . .127
Remopleurides spp. ........ 127
IV. STRATIGRAPHICAL DISTRIBUTION AND RELATIONSHIPS OF THE TRILOBITES 128
V. REFERENCES .......... 130
SYNOPSIS
The trilobites known from the Caradoc Series in the Cross Fell Inlier are described and figured,
many of them for the first time. They comprise fifty species and subspecies, assigned to
twenty-eight genera and fifteen families. The relationships of the trilobites to those of other
successions is reviewed. In the Longvillian and Marshbrookian Stages the fauna is of Anglo-
Welsh type with occasional Baltic elements, but in subsequent stages the affinities with corres-
ponding Norwegian and Swedish faunas becomes marked, reaching a maximum in the Pusgillian
Stage.
THE TRILOBITES OF THE CARADOC SERIES 69
I. INTRODUCTION AND ACKNOWLEDGMENTS
THE name Cross Fell Inlier is applied by geologists to the elongated area of Lower
Palaeozoic rocks, a few miles north of Appleby, Westmorland, which extends in a
south-south-easterly direction between the Carboniferous strata of the Pennines
and the Permo-Trias of the Vale of Eden. The geological structure of the area is of
great complexity and has been described by Shotton (1935). The disposition of the
principal outcrops of Caradoc strata is shown in Text-fig, i.
Although the first detailed stratigraphical accounts, accompanied by faunal lists,
of the Caradoc Series in the Inlier were given during the last century by Harkness
& Nicholson (1878), and Nicholson & Marr (1891), it was not until 1907 and 1910
that any figures and descriptions of trilobites were published. In those years Reed
described four new species, Lichas melmerbiensis , Acidaspis semievoluta, Homa-
lonotus ascriptus and Trinudeus nicholsoni, from strata which he called Dufton
Shales at the road-section near Melmerby, as well as listing several other species
from the same locality.
Bancroft (1933) listed a few Shropshire species of trilobites and brachiopods from
the Inlier but it was not until 1936 that another trilobite was illustrated, when
Shirley figured a specimen of Flexicalymene onniensis, a south Shropshire species,
from an unspecified locality and horizon within the Dufton Shales of Pus Gill, near
Dufton. In 1948 Bancroft's manuscript notes on the Cross Fell succession, in-
cluding locality maps of Swindale Beck and part of Pus Gill, were published
posthumously by Lament, who added photographs of certain Shropshire specimens
but figured none from the Inlier.
Since then a new genus, Duftonia, has been described from the Dufton Shales by
the writer (Dean, 1959) who, in a later paper (19590;), discussed the detailed strati-
graphy of the area and gave faunal lists. More recently some of the Trinucleidae
occurring in the upper part of the succession have been described and figured (Dean,
1961). The present work aims at describing and figuring all the trilobites listed
and discussed in the foregoing papers. Extensive collections of trilobites have been
made during field-work financed in part by grants from the Gloyne Fund of the
Geological Society of London, and I am grateful to the Council of the Society for
their assistance. All the material is housed in the British Museum (Natural
History), London. I wish to thank the Council of the Yorkshire Geological Society
for permission to reproduce the sketch-map shown in Text-fig. 4. I am indebted
to Mr. A. G. Brighton for the loan of type-specimens from the Sedgwick Museum,
Cambridge, to my wife for her important share of the collecting, and to Professor
W. F. Whittard who has kindly read and criticized the manuscript.
II. STRATIGRAPHICAL SUMMARY
The following table represents the subdivisions of the Caradoc Series which are
generally recognized today in the main portion of the Inlier, together with the
equivalent faunal zones as established in the type-area of south Shropshire.
The subdivision of the strata into Corona Beds and Dufton Shales is made on
lithological grounds, the former including maroon and grey mudstones and shales
7o
THE TRILOBITES OF THE CARADOC SERIES
SKETCH-MAP
SHOWING THE
DISTRIBUTION
OF CARADOC
OUTCROPS IN THE
CROSS FELL
INLIER [AFTER
SHOTTON. 1935]
ASHGILL SERIES
CARADOC SERIES
A. ALSTON ROAD
B. KNOCK-DUFTON
DISTRICT
C. HILTON BECK
D. ROMAN FELL
D.-
FIG. i.
THE TRILOBITES OF THE CARADOC SERIES
Stage
PUSGILLIAN
ONNIAN
ACTONIAN
MARSHBROOKIAN
U . Broeggerolithus longiceps
LONGVILLIAN
I
Zonal
TrUobite
Onnia superba
Onnia gracilis
Onnia ? cobboldi
Platylichas laxatus
Broeggerolithus transiens
Broeggerolithus globiceps
Zonal
Brachiopod
Onniella broeggeri
Onniella inconstans
Cryptothyris paracyclica
Onniella reuschi
Dalmanella unguis
Dalmanella wattsi
Kjaerina typa
Kjaerina bipartita
Bancroftina typa
Dalmanella indica and D. lepta
Dalmanella horderleyensis ?
Lithological
Subdivision
DUFTON
SHALES
CORONA
BEDS
with occasional limestone bands, whilst the latter comprise essentially a rather
monotonous series of dark-grey mudstones and shales with bands of nodular and
impure limestone which often weather to a rotten-stone or " gingerbread " rock.
The Corona Beds are shown in the table to coincide with the limits of the Lower
Longvillian Substage, but in fact the zone of Dalmanella indica and D. lepta is the
lowest faunal horizon definitely recognized, though the underlying zone may well be
represented. The topmost zone of the Lower Longvillian is represented in the highest
Corona Beds and it seems likely that there is a transition to the overlying Dufton
Shales, but owing to extensive faulting the lowest zone of the Upper Longvillian
has not been detected throughout the greater part of the Inlier. The remainder of
the succeeding Caradoc strata as found in the Shropshire type-area are, for the
most part, represented in the Cross Fell Inlier, with the addition of the Pusgillian
Stage. This subdivision is held to constitute the topmost part of the Caradoc
Series and to pre-date any known Ashgill strata. In Swindale Beck, near Knock,
the Pusgillian is followed in the succession by the Staurocephalus or Swindale Lime-
stone. The junction of the two has often been assumed to be conformable, but the
writer believes there is a marked stratigraphical break above the Pusgillian, and
that the Limestone represents a relatively high Ashgill horizon, as it is followed at
Swindale Beck by Ashgill Shales with Hirnantia sagittifera (M'Coy). South-east of
Dufton the Pusgillian outcrop is separated from that of the Swindale Limestone by
a small thickness of strata, equated with part of the Diacalymene Beds of Cautley,
which are probably overstepped by the Limestone.
The northernmost outcrops of Caradoc strata within the Inlier occur only in the
vicinity of the village of Melmerby, after which they are known as the Melmerby
Beds. The rocks have been divided into an upper and lower series ; the Upper
72 THE TRILOBITES OF THE CARADOC SERIES
Melmerby Beds contain a fauna indicative of the lower zone of the Upper Long-
villian Substage, an horizon which, as stated earlier, is not yet known elsewhere in
the Inlier. In the field, the Lower Melmerby Beds appear to underlie the upper
beds stratigraphically and are considered to be of Lower Longvillian age ; they
contain abundant evidence of a Longvillian age but the zonal brachiopods of the
Shropshire succession have not yet been detected and may be absent owing to the
unsuitability of the environment. The rocks are mainly dark-green or maroon,
blocky mudstones with occasional impure, nodular limestones. Their fauna includes
a conspicuously large number of trinucleid trilobites, accompanied by common
Kloucekia apiculata (M'Coy), features which readily distinguish them from the
equivalent strata farther south in the Inlier, and which are more suggestive of
corresponding North Welsh faunas.
No useful evidence has yet been obtained from the region to elaborate on the cor-
relation between the shelly and graptolitic successions already put forward for south
Shropshire (Dean, 1958 : 226). The Pusgillian, from its position above the Onnian
(believed to represent the topmost Dicranograptus dingani Zone) , is equated approxi-
mately with the Pleurograptus linearis Zone, and the shelly fauna suggests a
correlation with Etage 4ca of southern Norway, an horizon which is itself correlated
with that graptolite zone.
The fossil localities cited in the following descriptions are shown in Text-figs.
2-5. Text-fig 3 covers the well-known section of Swindale Beck, just north-east of
the village of Knock ; Text-fig 2 shows Pus Gill, north-east of Dufton ; Text-fig.
4 covers the large area of east of Dufton, with the important sections of Dufton
Town Sike, Billy's Beck and Harthwaite Sike ; Text-fig. 5 shows the small, isolated
outcrop of the Melmerby Beds north-east of Melmerby, near the northern end of
the Inlier.
III. SYSTEMATIC DESCRIPTIONS
Family RAPHIOPHORIDAE Angelin, 1854
Genus LONCHODOMAS Angelin, 1854
TYPE-SPECIES. Ampyx rostratus Sars, 1835 by subsequent designation of
Bassler (1915 : 41).
Lonchodomas swindalensis sp. nov.
(PL 6, figs. 2, 6, 8)
19590. Lonchodomas aff. rostratus (Sars) Dean, pp. 194, 207.
DIAGNOSIS. Lonchodomas with glabella projecting only short distance in front
of fixigenae. Pygidium semi-elliptical with four pairs of pleural furrows, the first
pair being the most sharply defined.
DESCRIPTION. The material is fragmentary for the most part but includes two
almost whole cranidia and one small, well-preserved pygidium. Excluding the
THE TRILOBITES OF THE CARADOC SERIES
73
74
THE TRILOBITES OF THE CARADOC SERIES
FOSSIL LOCALITIES
IN SWINDALE BECK
NEAR KNOCK
FIG. 3.
THE TRILOBITES OF THE CARADOC SERIES
75
o
£
76
THE TRILOBITES OF THE CARADOC SERIES
FOSSIL LOCALITIES
71
ON
THE
ALSTON ROAD, NEAR XMELMERBY
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1+ + • ^ '* ' * i v- o _±_ _+ + +'+/+ + + -»£
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+ + + + +/* + yf + + + + + + + + + + +r
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—A. FOSSIL LOCALITY
/ FAULTS DELIMITING
CARADOC OUTCROP
FIG. 5.
THE TRILOBITES OF THE CARADOC SERIES 77
frontal spine, the dimensions of the cranidia, numbered respectively In. 49946
and In. 49968, are as follows : length, 5 mm., breadth, 9 mm. ; length, u mm. ;
breadth, 16 mm. Dimensions of the pygidium, In. 49967, are : length, 1-5 mm.,
breadth, 4-5 mm. The cranidium is roughly rhomboidal in plan, produced anteriorly
into a long, frontal spine, the surface of which bears fine, longitudinal striae. The
glabella attains its greatest breadth at its mid-point and is bounded by almost
straight axial furrows containing slot-like hypostomal pits, situated frontally.
One cranidium (PI. 6, fig. 2) shows a faint median ridge extending from near the
occipital furrow to the base of the frontal spine, where it is replaced by a groove ;
similar grooves occur on the sides and undersurface of the frontal spine as in other
species of Lonchodomas , giving rise to the fluted, quadrate cross-section characteristic
of the genus. The posterior margin of the cranidium, and the pleuroccipital
furrows are parallel, transversely straight except medially where, owing to a small
extension of the glabella, they are convex backwards.
The pygidium is semi-elliptical in outline with a strongly-curved, steeply-declined,
posterior border which carries fine, parallel terrace lines, though the latter are not
visible in the photograph (PI. 6, fig. 6). The anterior margin is straight medially
but curves forwards a little anterolaterally. There are four pairs of pleural furrows,
the first pair being the most deeply impressed ; all are deepest anterolaterally
becoming faint at, or near, the shallow axial furrows. No axial rings are visible.
HORIZON AND LOCALITIES. All the known specimens have been collected from
Dufton Shales belonging to the upper half of the Upper Longvillian Substage in
Swindale Beck. There they occur in association with Kjaerina geniculata Bancroft,
Reuschella sp. nov. and Dolerorthis sp. The figured specimens are from localities
B. 5 and B. 6, but the same form has been found also at B. 10 (see Text-fig. 3).
These are the only records of Lonchodomas from the Upper Longvillian of the Anglo-
Welsh area.
HOLOTYPE. In. 49968 (PI. 6, fig. 2).
PARATYPES. In. 49946 (PI. 6, fig. 8) ; In. 49967 (PI. 6, fig. 6).
DISCUSSION. The Swindale Beck species bears an obvious relationship to
Lonchodomas rostratus (Sars) which has been refigured by St0rmer (1940 : 128, pi.
2, figs. 1-4). The latter species has, however, a smaller proportion of the glabella
projecting in front of the fixigenae, and the pygidium of the Norwegian form, the
type-specimens of which were obtained from the Ampyx Limestone (zone of
Nemagraptus gracilis) near Oslo, differs also in having only one pair of well-defined,
distally situated, pleural furrows, which die out before attaining the axial furrows.
Lonchodomas politus (Raymond, 1925 : 39, pi. 2, figs. 8, 9 ; Cooper, 1953 : 18,
pi. 6, figs, i, 2, 5-10), from the " Upper Lenoir Limestone " of Black River age in
Tennessee, has a conspicuously punctate test and a pygidium bearing seven or
eight pairs of furrows, only the first of which is deeply impressed, but otherwise
much resembles the Cross Fell species.
The stratigraphically younger species Lonchodomas pennatus (La Touche),
discussed elsewhere in the present paper, has a relatively larger proportion of the
glabella projecting in front of the fixigenae, whilst the pygidium has a less well-
differentiated axis than that of L. swindalensis , and only one pair of pleural furrows.
78 THE TRILOBITES OF THE CARADOC SERIES
Lonchodomas pennatus (La Touche)
(PI. 6, figs, i, 3-5, 9, 12)
1884. Ampyx pennatus La Touche, p. 57, pi. 3, fig. 56.
1891. Ampyx tetragonus Angelin : Nicholson & Marr, p. 511.
1932. Ampyx (Lonchodomas) pennatus La Touche : Reed, p. 205, pi. n, figs. 5-7.
1949. Lonchodomas pennatus (La Touche) Bancroft, p. 299, pi. 10, fig. 17.
1958. Lonchodomas pennatus (La Touche) : Dean, pp. 213, 224.
19590. Lonchodomas pennatus (La Touche) : Dean, pp. 200, 207.
1960. Lonchodomas pennatus (La Touche) : Dean, p. 82, pi. u, figs. 2, 5, 8-12.
Several cranidia agreeing in all essentials with La Touche's species have been
collected from the Dufton Shales. The often indifferent preservation renders
precise measurement difficult , most of the specimens being compressed vertically,
but the proportions are close to those of similarly preserved Shropshire material,
the length of the cranidium, measured from the base of the frontal spine to the
posterior margin of the occipital ring, being slightly more or less than three-quarters
of the maximum breadth. A feature not previously seen in L. pennatus but now
preserved in one specimen (PL 6, fig. i) is the retrousse form of the prismatic frontal
spine, which curves forwards and gently upwards from the glabella : a similar
curvature is seen also in Ampyx salteri Hicks and Ampyx linleyensis (Whittard,
1955, pi. i, fig. 17 ; pi. 2, fig. 6). A few individuals have traces of a slight median
ridge or carina on the glabella, and this may be either accentuated by lateral com-
pression or absent altogether. A similar ridge occurs in some Shropshire specimens,
but its presence varies with the state of preservation. Two probably immature
pygidia from Cross Fell (PI. 6, fig. 5) are proportionately broader than the forma
typica, their length : breadth radio being i : 3, but in mature pygidia this changes
to about i : 2, that is to say, generally comparable with the Shropshire material.
HORIZON AND LOCALITIES. In the Knock-Dufton district Lonchodomas pennatus
appears for the first time in the Onnian Stage, Onnia gracilis Zone, of Pus Gill, at
localities A. 8, 9, 12 and 14. It has been found less commonly in the overlying
Onnia superba Zone of Pus Gill, at localities A. 5 and 16. The ensuing Pusgillian
Stage has yielded specimens at both Pus Gill, localities A. 6, 18 and 28, and Swindale
Beck, locality B. 25.
DISCUSSION. La Touche's holotype was obtained from the Onnian Stage, Onnia
superba Zone, in the Onny Valley of south Shropshire, in which district Lonchodomas
pennatus ranges upwards from the Actonian Stage (Dean, 1960 : 83). The discoveries
in the Pusgillian Stage of the Cross Fell Inlier increase the known vertical range of
the species, but there is no information yet regarding its relationship to still younger
forms, though there is a general resemblance to certain species of Ashgill age, for
example that figured by Reed (1905 : 97, pi. 4, fig. 2) as Ampyx (Lonchodomas) cf.
rostratus Sars from the Slade Beds of Haverfordwest.
Elsewhere, a small cranidium of Lonchodomas which may be conspecific with L.
pennatus has been figured by Stermer (1945, pi. 2, fig. 10) as Lonchodomas aff.
rostratus (Sars) from the Tretaspis Shales of Hadeland, a horizon approximately
equivalent to the Pusgillian Stage. Lonchodomas carinatus (Cooper, 1953 : 17, pi. 7,
THE TRILOBITES OF THE CARADOC SERIES
79
figs. 12-17, 19-23) from the Black River Stage of Tennessee bears a strong resem-
blance to L. pennatus but possesses an irregularly punctate test, and the glabella
projects less far forwards in front of the fixigenae. The pygidium of L. carinatus
differs in having at least four discernible axial rings.
Family TRINUCLEIDAE Hawle & Corda, 1847
Subfamily CRYPTOLITHINAE Angelin, 1854
Genus BROEGGEROLITHUS Lamont, 1935
TYPE SPECIES. Cryptolithus broeggeri Bancroft, 1929 by original designation.
Broeggerolithus nicholsoni (Reed)
(PL 6, figs. 7, ii?, 14? ; PL 7, figs. 1-12)
1891. Trinucleus goldfussi Barrande ?, Nicholson & Marr, p. 509.
1891. Trinucleus seticornis Hisinger ?, Nicholson & Marr, p. 509.
1910. Tinucleus nicholsoni Reed, p. 212, pi. 16, figs. 1-9.
1912. Trinucleus nicholsoni Reed : Reed, pi. 18, fig. 6 ; pi. 19, figs. 4, 4^.
1914. Trinucleus gibbifrons M'Coy : Reed, p. 356, pi. 29, fig. 6.
1914. Trinucleus nicholsoni Reed : Reed, p. 357.
1927. Cryptolithus nicholsoni (Reed) Stetson, p. 88.
1940. Broeggerolithus nicholsoni (Reed) Whittington, p. 245.
1948. Broeggerolithus nicholsoni (Reed) : Bancroft in Lamont, p. 416.
I959& Broeggerolithus nicholsoni (Reed) : Dean, pp. 212, 214, 220.
1960. Broeggerolithus sp. (? nov.) Dean, p. 119, pi. 17, fig. 12.
1960. Broeggerolithus simplex Dean, p. 120, pi. 17, fig. 14.
DESCRIPTION. The cephalon is sub-semicircular in outline, broader than long,
the maximum breadth of the syntypes ranging from 14 mm. to about 22 mm. The
glabella is of moderate breadth, equal to about two-thirds of the total length, and
often carries a small, apical ocellus just forward of centre. A pair of small, basal,
glabellar furrows with apodemes is located immediately in front of a second pair of
apodemes situated at the distal ends of the occipital furrow. The glabella is
separated from the convex cheek-lobes by moderately-deep, almost straight, axial
furrows. The occipital ring is small, strongly convex transversely, produced
backwards and upwards to form a broadly-based occipital spine. The cephalic
fringe is moderately declined, both frontally and laterally, though this may not
always be apparent owing to compression of the specimen within a shaly matrix.
The upper surface of the fringe is generally smooth and there is no development of
raised interradial ridges such as are found in some earlier species of Broeggerolithus,
for example B. broeggeri (Bancroft). One or two specimens show suggestions of
such ridges, but these are almost certainly due to crushing and are not of constant
form. There are four concentric rows of fringe-pits in front of the glabella, where the
breadth (sag.) of the fringe contracts slightly so as to accommodate the frontal
lobe of the glabella. There is a strong radial arrangement of pits frontally, and this
8o THE TRILOBITES OF THE CARADOC SERIES
persists laterally. The concentric arrangement is also marked, and the variation
and norm of the four outer rows of pits is as follows : E2 = 19-24 (22), Ex = 21-23
(21), Ix = 21-22 (22), I2 = 20-22 (21). Up to three pits of E2 may be missing at
the genal angle, and in certain rare instances an occasional pit of E2 may be absent
laterally or anterolaterally. There is a notable lack of auxiliary E2 pits such as are
found laterally in certain related species of Broeggerolithus. There is little marked
differentiation in the size of pits, but those of Ij and I2 are slightly larger than the
others, especially towards the posterior margin of the fringe. E2 and Ej are sited
close together, but Ex and Ix are separated by a smooth, narrow, concentric " band "
corresponding in position with a conspicuous girder on the underside of the fringe.
A similar band of equal breadth separates I1 and I2, marking the position of a second
less well developed pseudo-girder. There are seven to nine, most commonly eight,
pits along the posterior margin of the fringe. I3 usually comprises fourteen or
fifteen small pits, and is developed from Rn or thereabouts. The posterior margins
of the fringe run slightly backwards, delimiting small genal prolongations, towards
the genal angles which are produced to form librigenal spines ; the latter curve
gently outwards and backwards, and their length is at least equal to that of the
cephalon.
One of the syntypes, A. 29607, retains only five thoracic segments of typical
trinucleid form. The axis of each is narrow (tr.) and bears a pair of apodemes
situated just above the axial furrows. On each pleura, a moderately-deep pleural
furrow, situated just forwards of centre, runs obliquely backwards from the axial
furrow to the pleural point. A complete topotype thorax (PI. 7, fig. 4) exhibits the
customary complement of six thoracic segments.
The syntype pygidium, A. 29615 (PL 7, fig. 2), is preserved as an internal mould
and few details can be seen. The axis has three clearly defined axial rings, and three
or four more are less well defined. The pleural lobes have three, or perhaps four,
segments, separated by faint pleural furrows and widening (exsag.) towards the
lateral margins.
HORIZON AND LOCALITIES. All Reed's syntypes were described by him as having
been collected solely from the Alston Road cutting near Melmerby, but the state of
preservation of most of the specimens indicates that they came from what are known
as the Lower Melmerby Beds, and the probable type-locality is believed to be that
shown in the present paper as locality J (see Text-fig. 5). One syntype, Sedg.
Mus. A. 29614, is preserved differently from the rest and the matrix suggests a
probable origin in the Upper Longvillian strata such as are found at Alston Road,
locality H. All the other localities, from A to G, along the Alston Road have yielded
Broeggerolithus nicholsoni, but the specimens are, in general, smaller than the syn-
types.
Although trinucleid trilobites are so abundant almost everywhere at the Alston
Road outcrop, elsewhere in the Inlier they are almost unknown from strata of
Longvillian age. An uncommon exception is locality E. 3 at Harthwaite Sike where
rare fragments, referred to B. cf. nicholsoni (see PL i, figs, n, 14), occur with an
assemblage indicating the Bancroftina typa Zone of the Lower Longvillian.
LECTOTYPE, here chosen. Sedg. Mus. A. 29613 (PL 7, figs. 6, 9).
THE TRILOBITES OF THE CARADOC SERIES 81
PARATYPES. Sedg. Mus. A. 29607 ; A. 29608 ; A. 29609 (PI. 7, figs, i, 7) ;
A. 29610 (PI. 7, fig. 10) ; A. 29611 (PI. 7, fig. 5) ; A. 29612 (PI. 7, fig. 8) ; A. 29614
(PI. 7, fig. ii) ; A. 29615 (PL 7, fig. 2).
DISCUSSION. In a recent account of trinucleid trilobites in south Shropshire
(Dean, 1960 : 103) the species assigned to the genus Broeggerolithus were divided
into three groups on the basis of small but significant differences in the structure of
the cephalic fringe. B. longiceps (Bancroft) and B. transiens (Bancroft) were placed
together in Group 3, whilst B. simplex Dean was placed doubtfully in Group 2.
The last-named species is here regarded as a synonym of B. nicholsoni and may also
be placed in Group 3. B. simplex was founded on a small cranidium from the Lower
Longvillian of the Onny Valley and was at that time believed to be distinct. The
pit count, however, falls within the limits of variation now established for B.
nicholsoni, and additional material from Shropshire tends to confirm the identity of
the two species
The fringe of Broeggerolithus transiens closely resembles that of the Melmerby
species but its pit-count for the four outermost rows is noticeably smaller, and there
are a few auxiliary pits situated anterolaterally in E2. Broeggerolithus longiceps is
remarkably similar to B. nicholsoni. The former species is, unfortunately, known
from only a comparatively small population sample in south Shropshire, but as far as
can be ascertained the pit-count falls within the limits of that for B. nicholsoni.
Like B. transiens, however, B. longiceps generally possesses a few auxiliary pits in
E2, though situated posterolaterally, and these have not been found in B. nicholsoni.
It is not known whether this is an invariable characteristic of B. longiceps, but if
this should not prove to be the case then there is no other valid reason for regarding
the species as distinct from B. nicholsoni. Certain well-preserved Shropshire speci-
mens of B. longiceps have reticulate cheek-lobes, a feature not seen on any of the
syntypes of B. nicholsoni but found on several specimens of the latter species from
the Upper Melmerby Beds of the Alston Road cutting.
Outside the Cross Fell Inlier Broeggerolithus nicholsoni probably occurs at several
localities in North Wales, associated with faunas of Longvillian age.
Broeggerolithus melmerbiensis sp. nov.
(PI. 6, figs. 10, 13)
1959a- Broeggerolithus aff. nicholsoni (Reed) pars, Dean, p. 214.
DIAGNOSIS. Large Broeggerolithus generally similar to B. nicholsoni but with large
number of fringe-pits, about one hundred, in four outermost concentric rows. Both
Ex and E2 extend to the genal angles.
DESCRIPTION. One well-preserved specimen collected from the Alston Road
cutting appears to resemble Broeggerolithus nicholsoni in most respects, but possesses
a number of fringe-pits much larger than the norm for that species. The specimen
has a closely similar radial and concentric arrangement of pits, but particularly
noticeable is the manner in which rows E2 and Ex are placed close together, whilst
rows E! and Ix are separated from each other by a wide, concentric " band " which
GEOL. 7, 3. 4
82 THE TRILOBITES OF THE CARADOC SERIES
coincides with the girder on the underside of the fringe. Ix and I2 are similarly
separated from each other, and the pits composing these two rows become larger
in size towards the posterior margin of the fringe. A conspicuous feature is the
way in which both Ex and E2 are developed as far as the genal angles. Both
I3 and I4 consist essentially of pits which are comparatively small, though increasing
slightly in size posteriorly, and the two rows are developed respectively from Ri3
and Rig. The pit count for the specimen is as follows : £2 = 24 pits, Ei = 25
pits, Ii = 26 pits, 12 = about 25 pits. There are four continuous concentric rows
of pits in front of the glabella, and seven pits are situated along each posterior margin
of the fringe. The total number of pits in rows E2 to 12 inclusive is about one
hundred, compared with an average of about eighty-five in Broeggerolithus nicholsoni
(s. s.). The single available cranidium possesses four attached thoracic segments
of typical Broeggerolithus aspect, but the total number originally present is unknown,
as is the pygidium.
HORIZON AND LOCALITY. The holotype was collected from a loose nodule of
impure limestone immediately below locality A in the Alston Road cutting, three-
quarters of a mile north-east of Melmerby (see Text-fig. 5). It is almost certain that
the specimen did, in fact, derive from this locality. The horizon is in the Long-
villian Stage, and the strata at this point are believed to be the Upper Melmerby
Beds, of Upper Longvillian age (Dean, 1959*1 : 213). The associated fauna includes
Broeggerolithus nicholsoni, Brongniartella and FlexicaLymene.
HOLOTYPE. BM. In. 52558.
DISCUSSION. Although bearing a general resemblance to certain other species of
Broeggerolithus such as B. nicholsoni, B. longiceps and B. transiens, the new form is
unlikely to be confused with them on account of its much higher pit-count for the
four outer rows, whilst no other species exhibits a comparable development of E2
to the genal angles.
Broeggerolithus cf. transiens (Bancroft)
(PI. 8, figs, i, 3, 4, 6, 8, ii)
1929. Cryptolithus transiens Bancroft, p. 90, pi. 2, fig. 5.
i959«. Broeggerolithus transiens (Bancroft) Dean, pp. 196-8, 207.
1960. Broeggerolithus transiens (Bancroft) : Dean, p. 123, pi. 18, figs. 2, 3, 7, 8, ii, 14. This
reference contains a list of all other synonyms.
Numerous cephala and cranidia have been found, agreeing in all essentials with the
description of the species from the type-area of south Shropshire (Dean, 1960 : 123).
The pit count for the various rows of the cephalic fringe falls within the limits of
variation of the Shropshire specimens, but instead of having one or two pits of Ej
marginal at the genal angle, the Cross Fell specimens commonly have three pits so
situated, though the number has been found to vary from one to four. Generally
there are up to two auxiliary pits in the outermost concentric row, E2, of the Shrop-
shire specimens, and these may sometimes be absent from the Cross Fell examples.
This casts some doubt on the validity of such a feature as a means of separating
closely-related species, but too few complete cranidia are available from either
THE TRILOBITES OF THE CARADOC SERIES 83
Shropshire or Cross Fell to give reliable statistics for the variation in E2. An analogous
situation has already been noted with regard to Broeggerolithus nicholsoni and B.
longiceps. The species is more abundant in the mudstones of the Cross Fell Inlier
than in the corresponding coarser sediments of south Shropshire, and individuals
frequently attain a larger size, the largest-known cephalon having a breadth of about
25 mm.
HORIZON AND LOCALITIES. In the Knock-Dufton district Broeggerolithus cf. tran-
sient is fairly common in Dufton Shales of the Marshbrookian Stage at Swindale
Beck. There it occurs in small numbers in strata thought to belong to the middle
portion of the stage (locality B 12), but becomes more abundant in higher strata
(localities B. n, 14, 16) which contain Kjerulfina cf. polycyma Bancroft and are
considered to be equivalent to the Onniella reuschi Zone of south Shropshire. B. cf.
transiens occurs also in Harthwaite Sike, east-south-east of Dufton (see Text-fig.
4) at localities E. 7, 8, 9?, 10-12, but the fauna there is sparse, composed in the
main of smaller individuals.
Broeggerolithus sp.
(PI. 8, %. 2)
19590. Broeggerolithus sp. (pars.) Dean, p. 207.
Among the numerous trinucleid trilobites found in the Marshbrookian rocks of
Swindale Beck is one individual quite different from all the rest. The specimen
comprises a fragmentary cranidium preserved as a limonitic external mould in a
dark-grey, cleaved mudstone. The estimated length and breadth of the original
cephalon are respectively 12 mm. (approx.) and 26 mm. Its most distinctive feature
is the manner in which the two outermost concentric rows of pits, E2 and Ex, end
abruptly without extending to, or even near, the genal angle as is customary in
normal forms of Broeggerolithus. E2 extends only to the anterolateral angle, leaving
three pits of Ex external to the margin. Ex stops far short of the genal angle, leaving
five pits of Ix external to the margin, though separated from it by a smooth band which
is equal in breadth to one row of pits. Such an arrangement results in the somewhat
unusual, indented form of the cephalic margin in front of the genal angle, and the
obliquely truncated shape of the anterolateral angle. The estimated number of
pits originally present is as follows : E2 + e2 = 12, Ex + et = 15, Ix + ix = 19,
I2 -f i2 = 19, I3 -f i3 = 10. There is a small triangular group of eight pits situated
between I2 and I3 near the posterior margin of the fringe, and seven pits are aligned
parallel to the latter. The pits of Ex and 1^ are of roughly equal size, larger than
those of E2 and I3. The last four pits of I2 nearest the posterior margin are of
particularly large size.
HORIZON AND LOCALITY. Found in mudstones of the Dufton Shales which contain
Broeggerolithus cf. transiens (Bancroft) and Kjerulfina cf. polycyma (Bancroft), and
belong to the topmost part of the Marshbrookian Stage, at locality B. 16 in
Swindale Beck.
DISCUSSION. It is not unusual for specimens of Broeggerolithus to have a de-
ficiency in the number of E2 pits near the genal angle, and in such forms as B.
84 THE TRILOBITES OF THE CARADOC SERIES
soudleyensis (Bancroft) and B. globiceps (Bancroft) some E2 pits may be absent
frontally, but no known species compares in any way with the excessive loss of pits
in the specimen now figured. In view of the large number of specimens of normal
Broeggerolithus cf . tmnsiens associated with the cranidium it seems likely that the
specimen represents a rare mutation or pathological form. Consequently it is not
proposed to erect a new specific name, at least until further material is available.
Genus ONNIA Bancroft, 1933
TYPE SPECIES. Cryptolithus superbus Bancroft, 1929 by original designation of
Bancroft (1933 : 2).
Onnia gracilis (Bancroft)
(PI. 8, figs. 12, 13)
1929. Cryptolithus gracilis Bancroft, p. 94, pi. 2, figs. 8, 9.
JQSQtf- Onnia gracilis (Bancroft) Dean, p. 207.
1960. Onnia gracilis (Bancroft) : Dean, p. 130, pi. 19, figs. 2, 7. This reference contains a
comprehensive synonymy of the species.
This species has recently been redescribed in detail by the writer (Dean, 1960 : 130)
who gave the following figures for the range in variation of the number of pits
present in the outermost four rows of the fringe. E2 + e2 = 31-37 (34), £!+€! =
21-25 (22), Ij + ij = 21-25 (23), I2 + ia = 21-25 (23) The figure in brackets
represents the mode. Insufficient well-preserved material is available from the
Cross Fell Inlier to give authentic figures for the corresponding variation, but the
available material has yielded the following results: E2 + e2 = 32-37, Ej -f- ^ =
20-23, Ix + it = 20-22, I2 + i2 = 21-22. It is therefore apparent that the speci-
mens fall within, or are very close to, the permissible variation for the species in
its type-area.
HORIZON AND LOCALITIES. As in south Shropshire Onnia gracilis characterizes
the middle portion of the Onnian Stage. It has been found in some abundance in
black mudstones of the Dufton Shales at Pus Gill, localities A. 8, 9, u, 12, 13, 14
and 15. Again, as in Shropshire, the species is accompanied by a fauna which includes
Onnicalymene onniensis (Shirley), Lonchodomas pennatus (La Touche), Chonetoidea
sp. and Onniella broeggeri Bancroft.
Onnia superba (Bancroft) pusgillensis Dean
(PL 8, figs. 5, 7, 9, 10)
1961. Onnia superba pusgillensis Dean, p. 120, pi. 7, figs. 1-6. Includes full synonymy.
Onnia superba (Bancroft), the zone fossil of the topmost subdivision of the Onnian
Stage in south Shropshire, has not been found in the Cross Fell Inlier, but is thought
to be represented by a distinct, and possibly local, subspecies 0. superba pusgil-
lensis. Although resembling 0. superba in almost all respects the Cross Fell form
may easily be distinguished by the continuous development of I2 in front of the
THE TRILOBITES OF THE CARADOC SERIES 85
glabella, and by the greater development of I3 and I4, respectively from R$ or R4,
and Ry.
HOLOTYPE. BM. In. 55707.
PARATYPES. BM. In. 50005 (PL 8, fig. 9) ; In. 50008 (PI. 8, fig. 5) ; In. 50049 ;
In. 55704 ; In. 55705 ; In. 55706 ; In. 55708.
HORIZON AND LOCALITIES. Dufton Shales, Onnian Stage, Onnia superba Zone,
Pus Gill, localities A. 5 and A. 16.
Subfamily TRETASPIDINAE Whittington, 1941
Genus TRETASPIS M'Coy, 1849
TYPE SPECIES. Asaphus seticornis Hisinger, 1840 by subsequent designation of
Bassler (1915 : 1285).
Tretaspis cf. ceriodes (Angelin) donsi Stermer
(PL 10, figs. 4, 6, 8)
1945. Tretaspis ceriodes (Angelin) var. donsi Stormer, p. 405, pi. i, fig. 8.
!959<*- Tretaspis cf. ceriodes (Angelin) : Dean, p. 207.
1961. Tretaspis cf. ceriodes donsi St0rmer : Dean, p. 129, pi. 9, figs. 7-9.
In southern Norway, Tretaspis ceriodes donsi has been shown by Stermer (1945 :
404) to be restricted to the upper part of the Upper Chasmops Limestone, 4b#.
The earliest-known occurrence of Tretaspis in the Cross Fell Inlier is at Pus Gill,
locality A. 13, in Dufton Shales belonging to the Onnia gracilis Zone of the Onnian
Stage, and the evidence at this point, though fragmentary, suggests a comparison
with the Norwegian subspecies. Additional, better-preserved material from the Onnia
superba Zone at Pus Gill, locality A. 5, is too incomplete for detailed comparison, but
shows all the principal features of T. ceriodes donsi. The horizon of the Cross
Fell specimens corresponds fairly closely with that of the Norwegian material.
Tretaspis convergens Dean
(PL 10, figs, i, 3, 5)
1961. Tretaspis convergens Dean, p. 127, pi. 9, figs. 1-6.
A detailed description of this species has already been given (Dean, 1961 : 127)
to which nothing need be added here. T. convergens is a distinctive form, its closest
relationships being with the species-group typified by Tretaspis seticornis (Hisinger),
particularly T. seticornis var. anderssoni Stermer (1945 : 402, pi. i, fig. 2). T.
convergens has so far been found at only one locality in the Cross Fell Inlier, namely
B. 25 in Swindale Beck. There it occurs in the highest part of the Dufton Shales,
belonging to the Pusgillian Stage, so that it is generally contemporaneous with
Tretaspis seticornis anderssoni and allied forms as found in Etage 4ca of southern
Norway.
HOLOTYPE. BM. In. 50059^, b (PL 10, figs, i, 3, 5).
Paratypes. In. 50030 ; In. 50037 (PL 10, fig. 2) ; In. 500580, b.
86 THE TRILOBITES OF THE CARADOC SERIES
Tretaspis kiaeri St0rmer radialis Lament
(PL 9, figs. 2-4)
1941. Tretaspis kjaeri mut. radialis Lament, 1941, p. 456, figs. 5, 6.
1961. Tretaspis kiaeri radialis Lament : Dean, p. 122, pi. 7, figs. 7-9, pi. 8, figs, i, 3-5.
Includes full synonymy of the subspecies.
The majority of the abundant specimens of Tretaspis collected from the Dufton
Shales of the Cross Fell Inlier can be referred to this subspecies. T. kiaeri radialis
was founded originally on two fragmentary specimens from the Portrane Limestone
of Eire, but the description has since been supplemented, using material from the
Dufton Shales (Dean, loc. cit.}, and the affinities with other species discussed.
HORIZON AND LOCALITIES. All the known specimens are from Dufton Shales
belonging to the Pusgillian Stage. The exact distribution within the Pusgillian
is not clear, owing to the predominance of successions affected by strike-faulting,
but the subspecies appears to occur throughout most of the stage. Localities include
the following : Pus Gill, A. 6, A. 18, A. 21, A. 24 to A. 30 ; Swindale Beck, B. 18,
B. 22 to B. 26, B. 28, B. 33, B. 34 ; Dufton Town Sike, C. I to C. 6 ; Billy's Beck,
D. i ; also the section at Hurning Lane, one mile north of Dufton.
Tretaspis kiaeri St0rmer duftonensis Dean
(PL 9, figs, i, 5-7)
1961. Tretaspis kiaeri duftonensis Dean, p. 125, pi. 8, figs. 2, 6-8.
The subspecies was described from a single, well-preserved specimen collected
from Dufton Shales belonging to the Pusgillian Stage at Pus Gill, locality A. 27.
Its closest relationship would appear to be with T. kiaeri radialis, from which it
differs in having a conspicuously larger pit-count for Ej.2, a less well-developed
concentric arrangement of I2_4 posterolaterally, as well as a larger, triangular area
of irregularly arranged pits on the genal prolongations of the fringe. T. kiaeri
duftonensis is not known with certainty outside the type-locality, though fragments
of fringe suggest that it may occur elsewhere at Pus Gill, but identification is difficult
in the absence of better-preserved material.
HOLOTYPE. BM. In. 5oo2oa, b.
Family CHEIRURIDAE Salter, 1864
Subfamily CYRTOMETOPINAE Opik, 1937
Genus PSEUDOSPHAEREXOCHUS Schmidt, 1881
TYPE SPECIES. Sphaerexochus hemicranium Kutorga, 1854 by subsequent
designation of Reed (18960 : 119).
THE TRILOBITES OF THE CARADOC SERIES 87
Pseudosphaerexochus cf. octolobatus (M'Coy)
(PI. 10, figs. 7, 9-12)
?i8gi. Youngia trispinosa Nicholson & Etheridge : Nicholson & Marr, p. 511.
I959« Pseudosphaerexochus sp. nov., Dean, pp. 204, 208.
DESCRIPTION. The cephalon is known only from the cranidium. The glabella
is strongly convex both longitudinally and transversely, subovate in plan, its maxi-
mum breadth about, or slightly greater than, three-quarters of the sagittal length.
It is bounded by deep, evenly-curved axial furrows, strongly convex abaxially, with
deep, hypostomal pits. The frontal lobe is small, rather less than one-quarter of
the glabellar length, and there are three pairs of equispaced glabellar furrows. The
first and second pairs are parallel, extending inwards about one-quarter of the
glabellar breadth ; they intersect the axial furrows at right-angles, but curve thence
adaxially backwards, at the same time becoming rapidly shallower. The first and
second pairs of glabellar lobes are identical in size and form, their convexity in line
with the remainder of the glabella. The third glabellar furrows are deeper than,
and parallel to, the first and second pairs, but curve backwards without reaching
the occipital furrow and become shallower proximally. The occipital ring is small,
narrow (tr.), strongly convex transversely, separated from the glabella by a moder-
ately deep occipital furrow. The fixigenae are small, convex, and steeply declined
abaxially. The pleuroccipital segment becomes slightly longer (exsag.) laterally,
ending posterolaterally in a pair of short, broad, blunt fixigenal spines set a short
distance inwards from the lateral margins. The pleuroccipital furrow is transversely
straight for the most part, becoming shallower distally where it curves forwards
towards, though without reaching, the lateral margins. The eyes have not been
found preserved. The palpebral lobes are narrow, moderately convex abaxially,
strongly arched longitudinally, and slightly divergent backwards ; they are defined
proximally by palpebral furrows of moderate depth which die out quickly to both
front and back. The anterior branches of the facial suture extend forwards from
the eyes, parallel to the axial furrows, to cut the anterior border, whilst the
posterior branches curve backwards distally to intersect the lateral margins just in
front of the line of the pleuroccipital furrow. The librigenae have not been found.
The surface of the glabella is covered with fine, evenly distributed granules, whilst
all the furrows are smooth. The surface of the fixigenae is covered with large,
dispersed pits, the intervening spaces, together with the surface of the pleuroc-
cipital segment, being covered with smaller punctae which are apparent only on the
external mould. The internal mould of the glabella is ornamented with closely-
grouped, small, prickly granules, probably representing the infillings of canals in the
original test.
The hypostoma and thorax are not known.
An incomplete pygidium, which must have been more than twice as broad as
long, is the only specimen that has been found (PI. 10, fig. 7). It is made up of four
segments which are produced backwards to form four pairs of strong, tapering
spines ; the latter are separated from one another by well-defined notches, and their
tips are stepped backwards slightly en echelon from first to fourth. Only three
88 THE TRILOBITES OF THE CARADOC SERIES
axial segments are visible, the first two well defined by deep ring furrows, and
the third poorly defined by two shallow transverse notches near the bases of the
fourth pair of pleural spines. The axial furrows comprise broad, shallow grooves,
moderately convergent backwards. Excluding the furrows, the surface of the
pygidium is granulate on the internal mould, but finely pitted on the external mould.
HORIZON AND LOCALITIES. The earliest-known occurrence of the species in the
Cross Fell Inlier is in the Dufton Shales, Onnian Stage, Onnia superba Zone of Pus Gill,
locality A. 5. Most of the available specimens are, however, from Dufton Shales
belonging to the Pusgillian Stage at Swindale Beck, locality B. 25. Other localities,
all in Pusgillian strata, are at Pus Gill, localities A. 6 and A. 30, and at Dufton
Town Sike, locality C. 6, east of Dufton.
DISCUSSION. The specific name Pseudosphaerexochus octolobatus has been used to
cover forms of the genus occurring in British Ordovician strata of various ages.
The holotype, described and figured by M'Coy (1849 : 4°7 » *» Sedgwick & M'Coy,
1851, pi. IG, fig. 10) as Ceraurus octolobatus, is a pygidium, Sedg. Mus. A. 11606,
from the Rhiwlas Limestone, of Ashgill age, near Bala, and the species must be
redescribed in detail before satisfactory comparisons can be made. Specimens from
the Ashgill Series, Drummuck Group, of Girvan were assigned by Reed (1906 : 141,
pi. 18, figs. 8-n) to Cheimrus (Cyrtometopus] octolobatus, and these appear to be
identical with the Cross Fell specimens, at least as far as the cephalon is concerned.
Pseudosphaerexochus has been recorded from the Onnian Stage of the Onny Valley
in south Shropshire (Dean, 19610 : 316), but the material there is too fragmentary
for comparison with that from Cross Fell.
Family ENCRINURIDAE Angelin, 1854
Subfamily ENCRINURINAE Angelin, 1854
Genus ENCRINURUS Emmrich, 1844
TYPE SPECIES. Entomostracites punctatus Wahlenberg, 1821 by original designa-
tion of Emmrich (1844 : 16).
Encrinurus sp.
(PI. n, figs, i, 4)
19590. Encrimtrus sp., Dean, p. 214.
Only one specimen has been collected, a pygidium about 5 mm. long and 4-5 mm.
broad frontally, preserved as an internal mould with part of the corresponding
external mould. Frontally the axis occupies about one-quarter of the maximum
breadth ; it is delimited by deep, smooth axial furrows and tapers backwards to a
narrow tip. There are eight pairs of pleural ribs which terminate laterally in small
free points ; the pleurae of the eighth pair almost coalesce with the terminal piece of the
axis, from which they are separated by a pair of shallow grooves. The axis as far
as the eighth pair of pleurae carries fifteen axial rings, but beyond this position is
apparently smooth. The ring furrows, apart from the first three or four, become
THE TRILOBITES OF THE CARADOC SERIES 89
shallower medially and there are traces of small median nodes on the third, fifth and
eighth axial rings. The remainder of the dorsal surface is smooth.
HORIZON AND LOCALITY. Locality H (see Text-fig. 5) beside the Alston Road,
three-quarters of a mile north-east of Melmerby. The strata, the Upper Melmerby
Beds, contain fossils indicating the lower part of the Upper LongvilHan Substage,
the equivalent of the Kjaerina bipartita Zone or Alternata Limestone in south
Shropshire.
DISCUSSION. Few Caradoc species of Encrinurus are available for comparison.
Encrinurus sp. (? nov.) from the Marshbrookian Stage of Shropshire (Dean, 1961^,
pi. 49, figs. 10, 12) has a similar number of ribs but a larger number of axial rings,
at least seventeen. The Alston Road specimen may well prove to represent a new
species.
Subfamily DINDYMENINAE Pfibyl, 1953
Genus DINDYMENE Hawle & Corda, 1847
TYPE SPECIES. Dindymene fridericiaugusti Hawle & Corde, 1847 by subsequent
designation of Barrande (1852 : 816).
Dindymene duftonensis sp. nov.
(PL n, fig. 13)
19591*. Dindymene sp. nov., Dean, pp. 198, 207.
The holotype is an incomplete cranidium of estimated breadth 4 to 5 mm., ex-
cluding fixigenal spines, the maximum breadth being roughly twice the length.
The glabella is strongly convex, both longitudinally and transversely, narrow
posteriorly but expanding forwards until equal to rather more than one-third of
the cephalic breadth. The axial furrows are deep and narrow posteriorly, but widen
forwards where they curve distally to join the lateral border furrow which skirts the
plump fixigenae. The occipital furrow is shallow and broad (sag.) medially, deepen-
ing laterally where a pair of deep apodemal pits is situated ; the occipital ring is
small, curving forwards abaxially to form a pair of occipital lobes. The pleuroc-
cipital segment is uniformly narrow (exsag.), transversely straight, separated from
the fixigenae by a parallel, moderately-deep, pleuroccipital furrow. The external
mould shows that the narrow (tr.) lateral borders meet the pleuroccipital segment
at the genal angles which are produced to form a pair of fixigenal spines, long,
slender, slightly curved, convex side forwards, and broadly splayed backwards.
The surface of the fixigenal spines, lateral border and pleuroccipital segment is
uniformly and finely granulate, appearing almost smooth. The surface of each
fixigena is covered with fine wrinkles, which contain numerous small pits, and carries
seven large tubercles ; three of the latter are situated anterolaterally in a row parallel
to the lateral border furrow, one is situated medially just in front of the pleuroccipital
furrow, and the remainder are arranged more or less sporadically over the fixigena.
The surface of the glabella, though incomplete, is estimated to have carried about
twenty tubercles of moderate size. Three of these form a median, longitudinal row
9o THE TRILOBITES OF THE CARADOC SERIES
extending half-way from the occipital furrow to the front of the glabella and are
flanked by two pairs of tubercles situated a short distance adaxially from the axial
furrows. The remaining tubercles are grouped more closely together across the
frontal lobe, and the intermediate spaces are finely granulate. These figures apply
to the partially preserved holotype, and the material available is insufficient to
decide whether there is any variation in the disposition of the tubercles.
The remainder of the exoskeleton is unknown.
HORIZON AND LOCALITY. Dufton Shales belonging to the Marshbrookian Stage,
probably the middle or upper third, at locality E. 12 (see Text-fig. 4) in Harthwaite
Sike, east-south-east of Dufton.
HOLOTYPE. BM. In. 546520, b, the latter, an external mould, being figured here
in the form of a latex cast.
DISCUSSION. The known species of Dindymene were reviewed recently by Kielan
(1959 : 146 et seq.). Of them, the species with which D. duftonensis may best be
grouped are D. ornate, Linnarsson, of Ashgill age, and D. plasi (Kielan, 1959 : 151,
pi. 29, figs. 1-3) from the Llanvirn Series of Bohemia. Like them it is equipped with
genal spines directed posterolaterally, but those of the Cross Fell species are notably
longer. Both the extra-British species differ from D. duftonensis in having a particu-
larly large tubercle, almost a cephalic spine, sited medially, one-third of the distance
from the occipital furrow to the front of the glabella ; in this respect they somewhat
resemble the more extreme development of a cephalic spine seen in the recently
described Cornovica, from the Lower Llanvirn of west Shropshire (Whittard, 1960 :
122), a genus which possesses, however, glabellar furrows and eleven thoracic seg-
ments. The fixigenae of D. duftonensis carry a similar number of large tubercles to
those of D. plasi and D. ornata : the glabella, however, has a greater number of
tubercles than that of D, ornata but fewer than that of D. plasi. As with D. ornata
the space between the glabellar tubercles is finely granulate, a feature apparently
not found in D. plasi.
Harper's (1956 : 389) record of Dindymene cf . ornata from Upper Longvillian strata
at Llanystwmdwy, Carnarvonshire, suggests that D. duftonensis may possibly be
represented in the North Welsh faunas.
Dindymene sp.
(PL n, fig. 6)
i959<z. Dindymene sp., Dean, pp. 194, 207.
Two fragmentary cranidia, one of which is figured here, have been collected from
the Upper Longvillian of Swindale Beck. In size and general form they match the
type material of D. duftonensis but, as the dorsal surface is not preserved, precise
identification has not been possible, though there are faint traces of some large
tubercles. The more complete specimen has a narrow (sag.), upturned, anterior
border, apparently separated from the glabella by a narrow (sag.), anterior border
furrow containing the facial suture. The latter follows the outline of the frontal
lobe as far as the axial furrow, beyond which it runs parallel to the margin of the
cephalon and separates a narrow (tr.), smooth border from the fixigena. It has not
THE TRILOBITES OF THE CARADOC SERIES 91
been possible to trace the line of the suture beyond the anterolateral portion of the
glabella.
HORIZON AND LOCALITY. Dufton Shales containing Kjaerina geniculata Bancroft
and forming the upper part, the Kjaerina typa Zone, of the Upper Longvillian
Substage, at locality B. 5 in Swindale Beck.
Subfamily CYBELINAE Holliday, 1942
Genus ATRACTOPYGE Hawle & Corda, 1847
TYPE SPECIES Calymene verrucosa Dalman, 1826 by original designation of
Hawle & Corda (1847 : 9°)-
Atractopyge scabra sp. nov.
(PI. II, figs. 2, 3, 10)
1891. Cybele verrucosa Dalman : Nicholson & Marr, pp. 505, 511.
I959a- Atractopyge aff. aspera (Linnarsson) : Dean, p. 207.
DIAGNOSIS. Glabella clavate, slightly broader than long, with frontal lobe strongly
convex forwards. Three pairs equisized glabellar lobes. Eye-ridges run from
palpebral lobes towards first glabellar furrows. Surface of test, excluding furrows,
covered with coarse tubercles, some of conspicuously large size. Pygidium longer
than broad. Four pairs of ribs ending in free points arranged en echelon. Narrow
axis with four continuous axial rings and a further fifteen or sixteen incomplete
rings.
DESCRIPTION. The length of the cranidium is rather more than half the breadth.
The glabella is moderately convex, clavate in outline, and attains its maximum
breadth, slightly greater than the median length, across the lateral extremities of
the anterior border. There are three pairs of glabellar furrows, represented by deep,
almost slot-like pits situated a short distance adaxially from the axial furrows.
The first pair of glabellar furrows is situated about mid-way between the frontal
margin and the occipital furrow, the second and third pairs being then positioned
at regular intervals so as to give three pairs of almost equisized glabellar lobes, their
long axes strongly divergent forwards. The occipital ring is short (sag.), convex
forwards medially ; laterally it curves forwards slightly to form a pair of occipital
lobes, immediately in front of which a pair of apodemal pits is situated at the extremi-
ties of the broad (sag.), smooth, moderately-deep occipital furrow. The axial furrows
are deep, wide (tr.), smooth, and rounded basally in cross-section ; from the occipital
ring they follow a slightly divergent course forwards as far as the first glabellar
furrows, whence they diverge more sharply to the false preglabellar field. The
frontal lobe of the glabella is strongly convex forwards where it is bounded by a
broad (sag.), smooth furrow, well defined but not deep, which intersects the axial
furrows opposite a pair of deep, hypostomal pits. Beyond this furrow the anterior
border is only slightly raised, but it is rendered more conspicuous by the presence
thereon of a row of well-developed tubercles of both large and medium size. Orna-
mentation of similar type, but including a greater number of large tubercles, covers
92 THE TRILOBITES OF THE CARADOC SERIES
the surface of the remainder of the glabella with the exception of the glabellar lobes.
In some individuals two, or even four, of the largest tubercles on the anterior border
may be arranged symmetrically about the sagittal line, and two or three pairs of
those on the main body of the glabella have been found to behave similarly, but
it has not proved possible to utilize satisfactorily, with the available material, the
notation proposed by Tripp (1957) for the tuberculation of certain species of
Encrinurus. The fixigenae are strongly convex, with faint suggestions of eye ridges
running towards the first glabellar furrows, and are surmounted by prominent,
pedunculate, palpebral lobes. No palpebral furrows have been seen on the latter,
which have a finely granulate, almost smooth surface, contrasting with the coarse
tuberculation of the fixigenae which ends abruptly at the base of the palpebral lobes.
The pleuroccipital furrow is straight for about half its length (tr.) but then turns
markedly backwards and finally dies out just before reaching the lateral margins.
The pleuroccipital segment, which follows the course of the pleuroccipital furrow, is
extremely narrow (exsag.) near the axial furrows but then widens appreciably to-
wards the apparently rounded genal angles. Each half of the pleuroccipital segment
carries about seven or eight tubercles arranged fairly regularly along its length (tr.).
The librigenae, hypostoma and thorax are not known.
Only a few isolated pygidia have been found, the largest of which is just over 12
mm. long. The best-preserved specimen (PI. n, fig. 10) is about 9-5 mm. in length,
with a maximum breadth of 8 mm. It is subovate in plan and only moderately con-
vex transversely. There are four pairs of pleural ribs, separated from each other by
strong rib furrows which become less well developed from first to fourth. The ribs
of the first pair curve abaxially backwards for about half the length of the pygidium
and then, more gradually, adaxially. The remaining ribs are subparallel to the first
pair but become successively less strongly curved until those of the fourth pair are
almost straight, converging backwards subparallel to the axial furrows. The ribs
terminate in small, free points which are " stepped " backwards en echelon from
first to fourth. The axis is in the form of an acute isosceles triangle, the two long
sides of which meet posteriorly at about 20 degrees, and there may be up to nineteen,
or perhaps twenty, axial rings visible. The first four axial rings are well defined,
continuous with the corresponding pairs of ribs. Frontally the axial furrows are
present merely as shallow grooves, but they become sharply defined behind the fourth
axial ring. Sometimes a fifth axial ring may be defined by a transversely continuous
ring furrow, but usually the greater part of the axis bears a smooth median band,
occupying about one-third of its breadth, on either side of which the axial rings are
clearly defined. There is a small, pointed terminal piece, behind which the points of
the fourth pair of ribs extend a little way. The greater part of the surface of the
pygidium is smooth but there are traces of incipient tubercles on occasional axial
rings, whilst up to five larger tubercles may be found along each rib, particularly its
hindmost two-thirds.
HORIZONS AND LOCALITIES. The earliest-known occurrences of Atractopyge
scabra within the Cross Fell Inlier are in Dufton Shales belonging to the Onnian Stage
at Pus Gill, where the species is uncommon in both the Onnia. gracilis Zone, locality
A 14, and the Onnia superba Zone, locality A 5. The new species is more common in
THE TRILOBITES OF THE CARADOC SERIES 93
the Pusgillian Stage, at which horizon it is known from Pus Gill, localities A 25,
27 and 28 ; Swindale Beck, locality B. 25 ; and Dufton Town Sike, localities C. 2,
3 and 5-
HOLOTYPE. BM. In. 50154 (PI. n, fig. 2).
PARATYPES. BM. In. 50147 ; In. 50150 (PI. n, fig. 3) ; In. 50157 (PI. u, fig. 10).
DISCUSSION. The name " Cybele verrucosa Dalman ", a species described originally
from the Red Tretaspis Shales of Vastergotland, Sweden, has often been used in
old faunal lists and collections from the Cross Fell Inlier, and from horizons of
various ages elsewhere. This trilobite, the type species of Atractopyge, has recently
been refigured by Henningsmoen (in Moore, 1959, fig. 349, la, b) whose illustrations
show that it differs markedly from A. scabra in having a glabella which is less
expanded frontally, with a smaller frontal glabellar lobe, as well as a pygidium which
is proportionately shorter, carrying noticeably fewer (15) axial rings, and with the
tips of the pleural ribs ending in line posteriorly.
The holotype of Atractopyge michelli (Reed, 19140 : 42, pi. 7, figs. 7, ja-c] from
the Balclatchie Group of Girvan is generally similar to A. scabra, but the glabella
is less expanded frontally and more coarsely tuberculate, whilst the pygidium is
proportionately shorter with fewer, less well-defined axial rings. Atractopyge
scabra bears a strong resemblance to several forms of the genus said to have been
collected from Ashgill strata in the Anglo- Welsh area, but until these have been
investigated further it is impossible to say whether the species ranges higher than
the Caradoc Series.
The lectotype pygidium of Atractopyge atractopyge (M'Coy in Sedgwick & M'Coy,
1851, pi. IG, fig. 4 only ; Dean, 1961^ : 319), from the Caradoc Series, probably
Longvillian Stage, of North Wales, is proportionately broader than that of A.
scabra and has a less strongly tapered axis, probably with fewer and better-defined
axial rings. M'Coy's illustration shows the pleural ribs ending in-line posteriorly,
but this is not clear on the actual specimen.
Atractopyge? sp.
(PI. ii, figs. 5, 8)
19590. Atractopyge sp. ind., Dean, p. 214.
A single specimen, a fragmentary cranidium, from the Lower Melmerby Beds,
Lower Longvillian Substage, at Alston Road, locality J, figured here in the form of a
latex cast, shows features suggestive of Atractopyge. The glabella is incomplete
but expands forwards and possesses three, almost equisized, pairs of glabellar lobes.
The fixigenae stand as high as the glabella, decline steeply towards the axial
furrows, and are topped by traces of pedunculate palpebral lobes. The surface of
the test is covered with large granules interspersed with larger tubercles, some of
which are arranged in pairs symmetrically about the sagittal line. The cranidium is
too poorly preserved for comparison with known species, but forms an interesting
addition to the trilobites known from the Longvillian Stage of the Anglo-Welsh
area. A further specimen (PL n, fig. 5), in a still worse state of preservation,
from the Corona Beds, Lower Longvillian Substage, Bancroftina typa Zone, at
94 THE TRILOBITES OF THE CARADOC SERIES
Harthwaite Sike, locality A. 3, shows only part of a glabella and occipital ring,
together with part of the corresponding left fixigena and palpebral lobe. Generic
determination is not possible and the specimen is figured here merely as Atractopyge?
sp., but it may prove to be related to the Alston Road trilobite.
Genus PARACYBELOIDES Hupe, 1955
TYPE SPECIES. Cybele loveni Linnarsson var. girvanensis Reed, 1906 by original
designation of Hupe (1955 : 271).
Paracybeloides cf . girvanensis (Reed)
(PI. n, figs. 7, 9, 12, 14, 15)
1891. Cybele Loveni Linnarsson : Nicholson & Marr, p. 511.
1906. Cybele loveni var. girvanensis Reed, p. 126, pi. 17, figs. 1-4.
1959^- Paracybeloides aff. loveni (Linnarsson) : Dean, pp. 207, 208.
Numerous specimens of Paracybeloides have been collected from the Dufton
Shales, and agree in all essential respects with the type material described by Reed
(loc. cit.} from the Ashgill Series of Girvan. According to Reed's original descrip-
tion the pygidial axis carries twenty-two to twenty-eight axial rings, but examina-
tion of several specimens, including syntypes, in the Gray Collection at the British
Museum (Nat. Hist.) suggests that the average number of rings present is about
twenty-three, whilst specimens with twenty or twenty-one rings are not unknown.
The variation in the number of axial rings found on the Cross Fell specimens is from
twenty to twenty-three, the most common number being twenty-one, and this
slight difference is not considered here to be of sufficient importance for specific
differentiation.
The tip of the pygidium of P. girvanensis was inaccurately illustrated by Reed
(loc. cit.}, who has been followed by other workers, for example Hupe (1955 : 272)
and Henningsmoen (in Moore, 1959, fig. 349, 5&). Reed's illustration shows the
terminal piece of the pygidial axis ending in front of the hindmost pair of pleural
points which, according to him, were separated from each other by a small gap.
In fact, the specimens figured by him, including BM. In. 23243 (Reed, 1906, pi.
17, fig. 4), possess a terminal piece which is produced postaxially into a long, flattened,
pointed process, constituting the longest (sag.) portion of the pygidium and united
abaxially with the proximal pair of pleural points so as to form a flattened, trifid,
pygidial termination. One of the Cross Fell specimens, BM. In. 50178, shows that
the terminal process described above was sometimes produced to an even greater
degree, giving a slender, pointed spine ; the total length of this particular pygidium
is approximately 17 mm., of which the axis sensu stricto occupies only about 10 mm.
Reed's illustrations of the cephalon of P. girvanensis show also that it possesses a
smooth occipital ring. This is apparently the case in the syntypes, and also in
other specimens, when preserved as internal moulds, but it is clear from the cor-
responding external moulds that they, together with the Cross Fell specimens,
THE TRILOBITES OF THE CARADOC SERIES 95
have a large median tubercle which may occasionally be produced backwards and
upwards to form a blunt, occipital spine.
HORIZON AND LOCALITIES. Paracybeloides cf. girvanensis appears uncommonly
in the Onnian Stage, Onnia gracilis Zone, of Pus Gill, locality A.5, but has not yet
been recorded from the overlying Onnia superba Zone ; this apparent absence may
be due to the paucity of exposures of the relevant strata. In the Pusgillian Stage
the same species becomes more abundant and has been found at Pus Gill, localities
A. 18, 25, 28-30 ; Swindale Beck, locality B. 25 ; and Dufton Town Sike, locality
C. 2.
Paracybeloides sp.
(PL n, fig. ii)
19590. Paracybeloides sp. ind., Dean, p. 214.
One specimen only has been found in the Longvillian Stage, constituting the sole
record of the genus at this horizon in the Anglo-Welsh area. It comprises a pygidium
which, excluding spines, is about 5 mm. in length with a frontal breadth of about
4 mm. The axis is narrow, less than one-third the frontal breadth, and carries
an estimated eighteen axial rings. Although the specimen is damaged, most of the
axial rings can be seen to be obsolete medially, where a median band extends back-
wards from at least the fourth ring. The main body of the axis is relatively short,
just over two-thirds of the total length of the specimen, but the terminal piece is
long, slender, and produced backwards, though the tip is not visible. The pleural
lobes are imperfectly preserved but appear to be of generally similar form to those of
Paracybeloides girvanensis. The specimen is inadequate for detailed comparison
with other species, but appears to be smaller and more slender than the pygidium
of P. girvanensis, with fewer axial rings. It bears some resemblance to the speci-
men of Paracybeloides figured by Whittington & Williams (1955, pi. 40, fig. no only)
as Atractopyge sp. ind., from the Derfel Limestone of early Caradoc age.
HORIZON AND LOCALITY. Lower Melmerby Beds, Lower Longvillian Substage,
Alston Road, locality J (see Text-fig. 5).
Family DALMANITIDAE Reed, 1905
Subfamily DALMANITINAE Reed, 1905
Genus DALMANITINA Reed, 1905
TYPE SPECIES. Dalmania socialis Barrande, 1852 by original designation of
Reed (19050 : 224).
Dalmanitina mucronata (Brongniart) matutina subsp. nov.
(PL 12, figs, i, 5, 6, 9, 12)
I959«. Dalmanitina sp. nov., Dean, pp. 204, 208.
DIAGNOSIS. Subspecies of Dalmanitina mucronata (Brongniart), characterized
by smaller fixigenal spines ; palpebral lobes developed from opposite mid-points
of first glabellar lobes to corresponding position opposite third glabellar lobes ;
96 THE TRILOBITES OF THE CARADOC SERIES
pygidium, excluding terminal spine, broader than long, with eight axial rings and
seven pleurae.
DESCRIPTION. Dalmanitina mucronata (Brongniart) has been figured and des-
cribed in detail by Temple (1952). The cephalon of the new subspecies resembles
that of D. mucronata, in all save two respects. The more important of these is the
length of the palpebral lobes which, in D. mucronata matutina, extend from just
outside the axial furrows, slightly in front of the mid-points of the first glabellar
lobes, backwards and slightly outwards until almost opposite the mid-points of the
third glabellar lobes. In D. mucronata itself the palpebral lobes extend backwards
from the first glabellar furrows until level with, or slightly behind, the second
glabellar furrows. Only one specimen of the new subspecies has been found with
part of the fixigenal spine preserved, but the latter appears to be smaller than that
of D. mucronata.
The hypostoma and thorax are not known.
The pygidium is subparabolic in outline, broader than long. The most complete
specimen, excluding the articulating half-ring and terminal spine, is 18 mm. broad and
about ii mm. (estimated) long. The axis occupies just over one-quarter of the frontal
breadth, about four-fifths of the median length (excluding terminal spine), and is
bounded by straight axial furrows which converge backwards and die out alongside
the terminal piece. There are eight axial rings, the first three of equal length (sag.),
the remainder becoming successively shorter. The articulating furrow and the
three succeeding ring furrows are moderately broad (sag.) and fairly shallow medially,
but become narrow abaxially and deepen to form apodemal pits. The remaining
ring furrows are of more uniform depth but become shallower from the fourth
furrow onwards, and the seventh and eighth furrows do not quite attain the axial
furrows on the internal mould. The terminal piece is subtriangular in plan and
passes backwards into a well-developed, post-axial ridge which runs, in turn, into a
stout, upturned, terminal spine. The pleural lobes are gently convex dorsally,
ending abaxially in smooth margins where there is a narrow doublure of apparently
uniform breadth. Each lobe carries seven pleurae, the first three of which are
separated by faintly-impressed, interpleural furrows and ornamented by deep,
pleural furrows which curve gently and abaxially backwards. The remaining
pleural and interpleural furrows are of about equal depth but become slightly less
impressed towards the tip of the pygidium. On the internal mould all these furrows
end abruptly at the proximal margin of the doublure, but on the external mould
they may be traced, in a much attenuated condition, to the lateral margin, coincident
with a poorly-defined border. The test of the entire pygidium, including the
terminal spine, is smooth.
Apart from having a smaller number of axial rings than the pygidium of D.
mucronata, eight as compared with eleven, that of the new subspecies differs also
in being relatively broader, almost semicircular in outline, with fewer pleurae, seven
compared with eight or nine, whilst the terminal spine is stouter and perhaps longer.
HORIZON AND LOCALITIES. Most of the type-specimens are from Dufton Shales
near the top of the Pusgillian Stage in Swindale Beck, locality B. 25. One of the
paratypes, however, is from Pus Gill, locality A. 7, where the horizon is low in the
THE TRILOBITES OF THE CARADOC SERIES 97
Pusgillian, not far above its presumed junction with the underlying Onnian Stage.
Another specimen has been collected by Mr. M. Mitchell from Pusgillian strata in
the section at Hurning Lane, 700 yards north-east of St. Cuthbert's Church, Dufton.
HOLOTYPE. BM. In. 49915 (PL 12, fig. 6).
PARATYPES. BM. In. 50112 (PI. 12, fig. i) ; In. 50113 (PL 12, fig. 5) ; In. 49919
(PL 12, fig. 9) ; In. 50114 (PL 12, fig. 12).
Subfamily ACASTINAE Delo, 1935
Genus KLOUCEKIA Delo, 1935
TYPE SPECIES. Phacops phillipsi Barrande, 1846 by original designation of
Delo (1935 : 408).
Subgenus Phacopidina Bancroft, 1949
TYPE SPECIES. Phacopidina harnagensis by original designation of Bancroft
(1949 : 310).
Kloucekia (Phacopidina} apiculata (M'Coy)
(PL 12, fig. n)
1851. Portlockia ? apiculata M'Coy in Sedgwick & M'Coy, p. 162.
1910. Phacops apiculatus Salter : Reed, p. 211.
19610. Kloucekia (Phacopidina} apiculata (M'Coy) Dean, p. 324, pi. 2, figs. 6-9, 12. Includes
full synonymy of the species.
Several specimens of this characteristic middle Caradoc trilobite have been found
in the vicinity of the Alston Road, north-east of Melmerby. The remains are
usually fragmentary but may sometimes include articulated thoracic segments.
The material agrees closely with the descriptions given by Harper (1947 : 169)
and later by Dean (19612 : 324), but the individuals, which occur in ashy mudstones,
are somewhat smaller than typical North Welsh specimens and significantly smaller
than those from south Shropshire, perhaps the result of a less favourable environ-
ment.
HORIZONS AND LOCALITIES. Kloucekia apiculata has been collected from the
Lower Melmerby Beds, Lower Longvillian Substage, of the Alston Road outcrop
(see Text-fig. 5) at localities B, C, E, F, G and J, and from the Upper Melmerby
Beds, Upper Longvillian Substage, of the same section at locality H. The species
has been found nowhere else in the Cross Fell Inlier.
Genus DUFTONIA Dean, 1959
uenus uvf i uiw/\ i^ean, 1959
TYPE SPECIES. Duftonia lacunosa by original designation of Dean (1959 : 143).
Duftonia lacunosa Dean
(PL 12, fig. 3)
1948. Pterygometopus sp., Bancroft in Lamont, p. 468.
1959. Duftonia lacunosa Dean, p. 144, pi. 19, figs. 1-3, 5, 6, 8.
IQ590- Duftonia lacunosa Dean : Dean, pp. 203, 204, 208.
GEOL. 7, 3.
g8 THE TRILOBITES OF THE CARADOC SERIES
The species has already been described in detail. All the type-specimens are
from the uppermost part of the Dufton Shales, Pusgillian Stage, at Swindale Beck,
locality B. 25, but other specimens have been found, though less commonly, at
locality B. 24 ; in Pus Gill, locality A. 22 ; and at Hurning Lane, north of Dufton.
At both the latter localities the horizon is apparently somewhat lower in the Pus-
gillian than are the rocks of the type-locality.
HOLOTYPE. BM. In. 49824 (PL 12, fig. 3).
PARATYPES. BM. In. 49821 ; In. 49826 ; In. 49830 ; In. 49920.
Family PTERYGOMETOPIDAE Reed, 1905
Subfamily PTERYGOMETOPINAE Reed, 1905
Genus CALYPTAULAX Cooper, 1930
TYPE SPECIES. CaLyptaulax glabella by original designation of Cooper (1930 : 387).
Calyptaulax planiformis sp. nov.
(PL 13, figs. 1-5)
1891. Phacops brongniarti Portlock, Nicholson & Marr, p. 505.
?i945- Calyptaulax sp., Stormer, p. 418, pi. 4, fig. 10.
1959^. Calyptaulax aff. norvegicus St0rmer : Dean, pp. 204, 208.
DIAGNOSIS. Glabella slightly longer than broad, pentagonal in outline, bluntly
pointed frontally attaining maximum breadth across frontal globe. Three pairs
of glabellar lobes ; first and second pairs conjunct laterally, third lobes small,
almost separated from remainder of glabella. Eyes long, crescentic in plan, ex-
tending from opposite first glabellar lobes almost to pleuroccipital furrow. Pygidium
subtriangular in plan, broader than long, with large facets, and terminates in blunt
point. Tapering axis with seven to ten axial rings ; pleural lobes with six pleural
furrows.
DESCRIPTION. Entire cephalon unknown. The glabella is only slightly convex,
even in uncrushed specimens, pentagonal in outline, broadly pointed frontally,
the length, excluding occipital ring, being slightly greater than the maximum breadth.
The frontal lobe is alate, extending laterally as far as the anterior ends of the first
glabellar lobes ; there it meets the axial furrows, which diverge forwards at about
30 degrees. The first glabellar furrows are only moderately deep, slightly sigmoidal,
and widely divergent anteriorly ; the first glabellar lobes are long and trapezoidal
in outline. The second glabellar furrows are shallow, curved, convex forwards,
and directed slightly backwards, not reaching the axial furrows, so that the distal
ends of the first glabellar lobes are continuous with the smaller, backwardly-directed,
second glabellar lobes. The third glabellar furrows are deep, with apodemes,
diverge backwards and almost isolate the third glabellar lobes which are thus present
only as semi-detached tubercles at the distal ends of the basal glabellar segment.
The occipital ring is long (sag.), especially medially, moderately broad, and convex
forwards with occipital lobes developed abaxially ; it is separated from the glabella
THE TRILOBITES OF THE CARADOC SERIES 99
by the occipital furrow which is shallow medially but deepens laterally, where it is
indented by the third glabellar lobes. The fixigenae are broad and level with long,
crescentic, palpebral lobes which are separated from the cheeks by conspicuous
palpebral furrows extending from the front of the first glabellar lobes to the pleur-
occipital furrow. The pleuroccipital segment is preserved in only one specimen (PI.
13, fig. 2) ; the pleuroccipital furrow becomes shallower laterally, not reaching the
lateral margin, and the genal angle is rounded. The anterior branches of the facial
suture follow the outline of the glabella and no preglabellar furrow is developed,
whilst the posterior branches curve gently forwards from behind the palpebral lobes
before curving backwards to meet the lateral margins.
The hypostoma and thorax are unknown.
The pygidium is roughly triangular in outline, the length approximately two-
thirds the breadth, with gently convex lateral margins which taper posteriorly to a
blunt, slightly upturned point. The axis tapers backwards fairly sharply and carries
seven to ten axial rings which diminish posteriorly in size and in degree of definition.
Beyond the tip of the axis there is a low, postaxial ridge. The pleural lobes are
only moderately convex and carry six pleural furrows which decrease in depth from
first to sixth, extend just over half-way from the axial furrows to the lateral margin,
and become progressively less divergent backwards. The first pleural furrows are
truncated distally by a pair of large facets. The first rib furrows are deep, extend-
ing rather more than half the breadth of the pleural lobes to cut the lateral margins,
and are directed sharply backwards laterally. Subsequent rib furrows are only
faintly impressed, though about three can usually be traced at the margin.
HORIZONS AND LOCALITIES. In the Cross Fell Inlier Calyptaulax planiformis
appears in small numbers in the Onnia gracilis Zone of the Onnian Stage, has not
yet been found in the Onnia superba Zone, probably as the result of inadequate
exposures, and becomes more common in the Pusgillian. Most of the type specimens
are from the Pusgillian at Swindale Beck, locality B 25, but one paratype is from
the Onnia gracilis Zone at Pus Gill, locality A. 12. Another locality where the
species is known to occur is Pus Gill, locality A. 6, also in the Pusgillian Stage.
HOLOTYPE. BM. In. 50138 (PI. 13, fig. 4).
PARATYPES. BM. In. 49903 (PI. 13, fig. 3) ; In. 49907 (PI. 13, fig. 5) ; In. 49908
(PI. 13, fig. 2) ; In. 49965 (PI. 13, fig. i).
DISCUSSION. Calyptaulax planiformis is clearly related to the approximately
contemporaneous C. norvegicus Stormer (1945 : 417, pi. 4, figs. 2, 3), a species which
may possibly be synonymous with Calyptaulax [Homalops] altumi (Remele) figured
by Wiman (1908, pi. 8, figs. 7-10) from the Baltic region. C. planiformis differs
from C. norvegicus in the following respects : the glabella is proportionately longer
and less rounded frontally ; the axis of the pygidium is longer ; the rib furrows are
definitely, though generally faintly, defined at the pygidial border, contrasting with
the smooth border described by St0rmer (1945 : 418). In the last feature the Cross
Fell species resembles more the pygidium figured by Stermer (1945, pi. 4, fig. 10) as
Calyptaulax sp.
Although Calyptaulax was once thought to be a predominantly North American
genus it is now known from several places in northern Europe and Scandinavia.
ioo THE TRILOBITES OF THE CARADOC SERIES
In addition to the Norwegian and Baltic records mentioned above, Jaanusson
(1953 : 102-103) has recorded Homalops ( = Calyptaulax} cf. altumi Remele from
the Slandrom Limestone, Pleurograptus linearis Zone, of southern Sweden.
Calyptaulax actonensis Dean (ig6ia : 328) from the Actonian and basal Onnian
Stages of south Shropshire strongly resembles the new species, particularly in the
form of the cranidium, but the pygidium is slightly shorter, has a shorter axis with
fewer axial rings, possesses more strongly denned pleural furrows, and terminates in
a better-developed, pointed, caudal spine.
Of the other comparable species of the genus, Calyptaulax compressa (Cooper,
1930, pi. 5, figs. 7, 8) has a shorter glabella and smaller fixigenae than has C. plani-
formis, and the pygidium of C. glabella (Cooper, 1930, pi. 5, figs. 9-11) is both broader
and more pointed posteriorly. Calyptaulax schucherti from the Cape Calhoun
Formation of northern Greenland (Troedsson, 1929, pi. 19, figs. 17-20) may be
distinguished from the new species by its shorter frontal glabellar lobe, and smaller
fixigenae and first glabellar lobes.
Genus ESTONIOPS Mannil, 1957
Estoniops comprises phacopid trilobites generally resembling Pterygometopus
(s.s.) but differing essentially from that genus in their lack of a preglabellar furrow.
The frontal lobe of the glabella is large and broad, and the first glabellar lobes are
relatively large, but the second and third glabellar lobes are much reduced in size.
The genal angles are rounded. The surface of the glabella is covered with coarse
tubercles, but that of the cheeks is pitted. The pygidium is generally similar to
that of Pterygometopus, the tip being bluntly rounded, and has a smooth border.
Estoniops has been found in Britain only in the Upper Longvillian Substage of
the Anglo-Welsh area but the genus is widely distributed geographically and is
known from Scandinavia and the Baltic region. These extra-British occurrences
appear to be roughly contemporaneous with, or somewhat earlier than, the Anglo-
Welsh material.
TYPE SPECIES. A caste exilis Eichwald, 1857 by original designation of Mannil
(1957 : 386).
Estoniops alifrons (M'Coy)
(PI. 12, figs. 2, 4, 7, 8, 10, 13, 14)
1851. Phacops (Phacops) alifrons M'Coy in Sedgwick & M'Coy, p. 159, pi. iG, figs. 12-14.
1852. Phacops (Phacops) alifrons M'Coy : Salter, p. ii.
1853. Phacops jukesi Salter, p. n.
1864. Phacops (Acaste) alifrons M'Coy : Salter, p. 33, pi. i, figs. 31-34.
1864. Phacops (Chasmops ?) jukesii Salter : Salter, p. 36, pi. i, figs. 29, 30.
1873. Phacops (Acaste) alifrons M'Coy : Salter, p. 52.
1891. Phacops (Acaste) alifrons M'Coy : Woods, p. 149.
1922. Pterygometopus jukesii (Salter) Elles, pp. 150, 152, 170.
1923. Phacops (Pterygometopus) jukesii Salter : King, p. 491.
X933- Pterygometopus jukesi (Salter) : Bancroft, table i.
1945. Phacops (Calliops) jukesi Salter : Reed, p. 315.
THE TRILOBITES OF THE CARADOC SERIES 101
1959. Estoniops jukesi (Salter) Dean, p. 146.
i959«. Estoniops jukesi (Salter) : Dean, pp. 194, 207, 220.
ig;jia. Estoniops alifrons (M'Coy) Dean, p. 320.
The species was founded by M'Coy (in Sedgwick & M'Coy, 1851 : 159) on three
syntypes, now in the Sedgwick Museum. One of these, numbered A. 42694
(Sedgwick & M'Coy, 1851, pi. IG, fig. 12), is chosen here as lectotype and is re-
figured (PI. 12, fig. 4). It comprises a damaged, incomplete cephalon which has under-
gone slight lateral compression, so that the glabella appears more elongated than is
normally the case. The specimen was said to be " from the impure limestone of
Capel Gannon " (Denbighshire), a horizon presumed to be of Upper Longvillian
age. The remaining syntypes comprise two pygidia, said to be from the " Lime-
stone of Pont y Glyn, Diffwys " (Denbighshire). One of them, Sedg. Mus. A. 42695
(Sedgwick & M'Coy, 1851, pi. IG, fig. 13), is incomplete and probably slightly crushed,
whilst the other, A. 42696 (Sedgwick & M'Coy, 1851, pi. IG, fig. 14), is a complete
pygidium, preserved as an internal mould which is believed to have undergone slight
longitudinal compression and is therefore thought to be unsuitable for systematic
description owing to the exaggeration of the pleural furrows.
Phacops jukesi Salter (1853 : n), a species regarded here as a synonym of Estoniops
alifrons, was established using two syntype cephala from the Bala district. These
are in the Geological Survey & Museum where they are numbered 19165 and 19166.
Owing to the poor state of preservation of the type specimens of Estoniops alifrons
the following description is founded also on supplementary material from both the
Bala district, north Wales, and the Cross Fell Inlier.
DESCRIPTION. The cephalon is strongly convex, of semi-elliptical form, broader
than long. The glabellar outline is slightly convergent from the posterior border as
far as the outer ends of the second glabellar furrows, but then expands rapidly for-
wards, attaining its maximum breadth a short distance in front of the first glabellar
furrows. The glabella is slightly broader than long and there is no preglabellar
furrow. The frontal lobe is convex, strongly alate, its projected length in an un-
crushed specimen being about two-fifths that of the glabella. The first glabellar
furrows are moderately deep, directed backwards slightly and extending adaxially
about one-quarter to one-third of the glabellar breadth. The second glabellar
furrows are deep, short (tr.), directed forwards slightly so that the first glabellar
lobes are almost subtriangular in outline. The second glabellar lobes are small,
directed backwards, bounded posteriorly by deep third glabellar furrows. The
third glabellar lobes are merely the swollen ends of what may be described as a ring-
like segment, corresponding in position with the occiput or peduncle of the Tri-
nucleidae and separated by a well-defined occipital furrow from the convex occipital
ring, which is slightly wider (tr.) than is the glabella across the basal lobes. The
axial furrows are narrow and deep, becoming shallower opposite the first glabellar
lobes, with hypostomal pits situated opposite the first glabellar furrows. From the
posterior margin as far as the second glabellar lobes, the axial furrows converge
slightly but then diverge markedly until just beyond the first glabellar furrows,
whence they coalesce with the lateral marginal furrows ; the latter run back as far
as the distal ends of the pleuroccipital furrow, towards which they become consider-
102 THE TRILOBITES OF THE CARADOC SERIES
ably shallower. The eyes are prominent, standing as high as, or slightly higher than,
the glabella, and the fixigenae are declined steeply towards the axial furrows. The
eye of one of the Cross Fell specimens contains about 165 lenses arranged in twenty-
four vertical rows, each row containing from four to eight lenses, but Welsh speci-
mens have not yet been found sufficiently well preserved for detailed comparison.
The palpebral lobes are sharply crescentic in form, bounded by deep palpebral
furrows. Each palpebral furrow runs from the axial furrow and turns sharply
through a right angle opposite the mid-point of the eye before attaining the posterior
end of the latter ; from this point a deep furrow runs immediately below the eye,
whilst another furrow curves gently forwards and then laterally backwards, becoming
almost obsolete before intersecting the lateral marginal furrow. The anterior
branches of the facial suture run obliquely forwards from the eyes to cut the axial
furrows just in front of the hypostomal pits ; they then turn sharply inwards to
converge and meet in front of the glabella. The posterior branches curve forwards
gently from behind the eyes and then straighten before cutting the lateral margins ;
they are situated in the furrows already described. The librigenae are fused frontally,
and a median suture is not developed. The pleuroccipital furrows are deep and
transversely straight but die out before reaching the margin. The pleuroccipital
segment widens (exsag.) noticeably towards the smoothly rounded genal angles. The
test of the glabella is covered with coarse tubercles the surface of which, together with
the intervening area, is finely granulate. The surface of the cheeks is pitted, but
that of the palpebral lobes, occipital ring, pleuroccipital segment and cephalic
border is smooth or finely granulate. The cephalic doublure is similarly orna-
mented but there is no sharp demarcation between this and the tuberculate upper
surface of the cephalon at the line of the facial suture.
The hypostoma and thorax are unknown.
No well-preserved topotype pygidium being yet available for examination, the
following description is founded on a pygidium, presumed to belong to the species
(PI. 12, figs. 8, 10), which has been found at Swindale Beck in association with a
cephalon of E. alifrons. The outline is broadly semi-elliptical, the anterior margin
slightly convex forwards, and the tip well rounded. The straight-sided, tapering
axis, bounded by deep axial furrows, has seven, well-defined axial rings, followed
by two further rings which are less distinct medially. The pleural lobes carry six
ribs, separated by deep furrows, with a seventh rib less well defined. Interpleural
furrows are only faintly impressed and neither they nor the pleural furrows attain
the lateral margin, so that a smooth border results.
HORIZON AND LOCALITIES. Although the horizon of M'Coy's syntypes was not
specifically stated, the evidence of all other known occurrences of Estoniops alifrons
in the Anglo-Welsh area suggests that they must have been collected from strata
of Upper Longvillian age. Salter's type material of Phacops jukesi in the Geo-
logical Survey & Museum is labelled merely " Caradoc " of Gelli Grin, Bala, but all
the material of the species from that area contained in the Bancroft Collection at
the British Museum (Nat. Hist.) is described as having been collected from the Upper
Longvillian substage, presumably from what Bancroft (1933, table i) termed the
Pterygometopus jukesi Beds. In her paper on the Bala district Elles (1922 : 150,
THE TRILOBITES OF THE CARADOC SERIES 103
152, 170) recorded Pterygometopus jukesi from the Gelli Grin Calcareous Ash Series
at various localities near Bala. The rocks in which E. alifrons occurs at Cross Fell
belong to the upper half of the Upper Longvillian substage. The species is uncom-
mon there and has, as yet, been found at only two localities, both in Swindale Beck,
B. 7 and B. 10.
Phacops (Pterygometopsis) [sic] cf . jukesi has been recorded from the Pen-y-garnedd
Limestone of the Berwyn Hills, North Wales (Wedd et al., 1929 : 44), a horizon now
known to contain Upper Longvillian fossils (Whittington, 1938 : 436, 451).
LECTOTYPE, here selected. Sedgwick Mus. A. 42694 (PI. 12, fig. 4).
PARATYPES. Sedgwick Mus. A. 42695 ; A. 42696.
DISCUSSION. In redescribing the holotype of the type species of Pterygometopus,
P. sclerops (Dalman), Whittington (1950 : 548) sought to restrict the genus to that
species and P. trigonocephala (Schmidt) on account of their having such features as a
distinct preglabellar furrow, set back slightly from the facial suture in front of the
glabella, and palpebral furrows which continue behind the eye and then, containing
the posterior branches of the facial suture, curve towards the marginal furrows.
Other forms usually assigned to Pterygometopus were said to be distinct because the
preglabellar furrow was not impressed, the glabella and preglabellar field being
continuous, and the genus Estoniops seems to accommodate these phacopids.
Estoniops shares certain characteristics, such as the absence of a preglabellar
furrow and the presence of cheeks with a pitted surface, with the genus Duftonia
(Dean, 1959) but, in addition to other features, the possession of fixigenal spines and
a mucronate pygidium assist in separating the latter generically. Estoniops alifrons
has many features in common with E. exilis Eichwald sp. (Schmidt, 1881 : 86,
pi. i, figs. 18-21 ; pi. 12, fig. 13 ; Wigand, 1888 : 43, pi. 6, figs. 5«, b) from the
Kuckers Shale, Stage C2, of Estonia, a horizon probably slightly earlier in age than
the Upper Longvillian, but the cephalon of the British form differs in having a
more elongate glabella and more divergent axial furrows. A particularly noticeable
feature in both species, and in others assigned to Estoniops, is the curiously angular
form of the palpebral furrow, that of E. exilis having been illustrated by Opik
(1937 : 163, fig. 42). Phacops (Pterygometopus) panderi Schmidt (1881 : 84, pi. i,
figs. 15-17 ; pi. 12, figs. 10-12) from Stage Ci of the Baltic region bears some re-
semblance to Estoniops but differs in having tuberculate cheeks, and it is not known
whether this feature is of more than specific significance.
Two specimens figured by Salter (1864, pi. i, figs. 33, 34) as Phacops (Acaste)
alifrons can probably be assigned to that species. One of them, GSM 19109 (Salter,
1864, pi. i, fig. 35) is an incomplete cephalon labelled, on the tablet, " Tyn-y-cabled,
Pennant " ; the other, GSM 19107, is a somewhat distorted pygidium from " Goetre,
Meifod ". The latter, from its general similarity to the Swindale Beck pygidium
figured here, almost certainly belongs to the same species and is associated with a
cephalon, GSM 19108, which definitely belongs to E. alifrons. None of the localities
above matches those listed by Salter (1864 : 34) for Phacops alifrons.
The specimen recorded in the Oswestry Memoir (Wedd et al., 1929 : 61) as Phacops
cf. alifrons from the Ashgill Series of the Berwyn Hills is, in fact, a crushed Para-
cybeloides sp. ; it is numbered RE 938-9 in the Geological Survey & Museum.
io4 THE TRILOBITES OF THE CARADOC SERIES
The trilobite from the Balclatchie Beds of the Girvan district described by Reed
(1945 : 314, pi. i, fig. 6) as Phacops (Calliops} jukesi var. vicina appears to be a
typical Calliops not unlike C. brongniarti (Portlock) and cannot be considered closely
related to Estoniops alifrons.
Subfamily CHASMOPSINAE Fillet, 1953
Genus CHASMOPS M'Coy, 1849
TYPE SPECIES. Calymene odini Eichwald, 1840 by original designation of M'Coy
(1849:403).
Chasmops cf. extensa (Boeck)
(PL 13, fig. 8)
I959a- Chasmops cf. extensa (Boeck) : Dean, p. 207.
Three fragmentary pygidia belonging to Chasmops of the extensa species-group
have been recovered from Swindale Beck, but only one is sufficiently well preserved
for descriptive purposes. The specimen is about 18 mm. long, and the original
maximum breadth is estimated to have been about 27 mm. The axis is narrow,
with a frontal breadth of about 5 mm., tapers strongly backwards and ends about 3-5
mm. in front of the tip of the pygidium. There is evidence, obscured by crushing,
of a postacial ridge. As far as the state of preservation allows, there appear to be
fifteen axial rings. Each pleural lobe carries fourteen ribs, the pleural furrows
becoming noticeably shallower near the lateral margin. Each rib usually bears a
faintly-impressed interpleural furrow and becomes wider (exsag.) abaxiaUy. The
ribs become smaller towards the tip of the pygidium, at the same time changing
their position relative to the axis until the smallest are almost parallel to the sagittal
line.
HORIZON AND LOCALITY. Dufton Shales, believed to belong to the middle
portion of the Actonian Stage, at Swindale Beck, locality B. 15. The associated
fauna includes Onnicalymene, Remopleurides and Onniella.
DISCUSSION. Chasmops extensa has been redescribed by St0rmer (1940 : 138, pi.
3, figs. 7-11) whose illustrations of the pygidium show that it possesses fifteen axial
rings and fourteen pleural ribs, corresponding with the numbers in the specimen
now figured. The type pygidium is, however, much more elongate, and narrowly
elliptical in outline, but the specimen is uncrushed whereas those from Cross Fell
are flattened in a dark-grey shale. In southern Norway C. extensa is characteristic
of the Upper Chasmops Shale and Limestone, and in south Shropshire members of
the same species are most abundant in the Actonian and Marshbrookian Stages,
though appearing earlier, in the Upper Longvillian, and ranging upwards into the
lowest part of the Onnian (Dean, 19610 : 335).
THE TRILOBITES OF THE CARADOC SERIES 105
Chasmops aff. maxima (Schmidt)
(PL 13, fig- 6)
I959«. Chasmops aff. maxima (Schmidt) : Dean, p. 208.
The material consists only of one damaged cranidium, estimated length about
15 mm. It is moderately convex, both longitudinally and transversely, and the
axial furrows diverge anteriorly at approximately 40 degrees. The frontal lobe is
incomplete but its length is about two-thirds of the estimated breadth. The first
glabellar furrows are almost straight, diverging forwards at no degrees and deepen-
ing towards the axial furrows ; the first glabellar lobes are triangular in outline,
their convexity conforming with the remainder of the glabella. The second glabellar
furrows converge forwards at about 75 degrees and deepen markedly towards the
axial furrows ; the second glabellar lobes are present only as small tubercles. The
third glabellar furrows occur as notches separating the second glabellar lobes from
those of the third pair, which are represented by tubercle-like projections situated
just behind the first glabellar lobes and slightly overlapping the occipital furrow.
The latter is transversely straight, deepening towards the axial furrows, near which
it contains a pair of apodemal pits. Another pair of apodemal pits is situated at the
hind margin of the occipital ring, just inwards from the axial furrows and bordering
two small, anteriorly-projecting, occipital lobes. The pleuroccipital furrow is
moderately deep and the pleuroccipital segment is fairly narrow (exsag.), but the two
are only partly preserved. Even though incompletely preserved, the one surviving
palpebral lobe is prominent, raised above the level of the glabella and carries a
shallow, palpebral furrow. In plan the palpebral lobe extends from a point almost
opposite the middle of the first glabellar lobe to one opposite the third glabellar
lobe. The fixigena declines steeply from the palpebral lobe to the axial furrow.
HORIZON AND LOCALITY. Dufton Shales in, probably, the higher part of the
Pusgillian Stage at Pus Gill, locality A. 28.
DISCUSSION. The species has been discussed in detail by St0rmer (1945 : 420)
who has pointed out that Schmidt (1881 : 112) founded C. maxima on specimens from
both the Jewe and Kegel Stages and figured a variety of forms under the one name.
The specimen from Pus Gill matches best the specimens of C. maxima figured by
St0rmer from the Tretaspis Shales of Hadeland, though a detailed comparison is not
possible with the material available.
Chasmops sp.
(PL 13, fig- 12)
19590. Chasmops sp., Dean, p. 214.
Two specimens of Chasmops have been recovered from the Caradoc Series near
Melmerby. One, a cranidium BM. In. 54655, is too badly damaged for detailed
examination. The other, an almost complete pygidium In. 54656, is slightly
distorted, but the length and breadth are estimated to have been respectively 12
mm. and about 16 mm. The axis possesses eleven axial rings extending almost to
io6 THE TRILOBITES OF THE CARADOC SERIES
the terminal piece which, although damaged, appears to be only partly differentiated
from the postaxial ridge. The pleural lobes have ten, or perhaps eleven, ribs
some of which bear faint traces of interpleural furrows. The ribs are separated
from one another by deep pleural furrows which, as far as can be ascertained, extend
to the lateral margins of the pygidium.
HORIZON AND LOCALITY. Lower Melmerby Beds, probably representing as an
yet undetermined part of the Lower Longvillian Substage, at the Alston Road
section near Melmerby, locality J (see Text-fig. 5).
DISCUSSION. The fragmentary evidence available suggests that the species
may be new. It is certainly distinct from the so-called longicaudate forms of
Chasmops, such as C. extensa (Boeck), which are so abundant higher in the Caradoc
Series of Shropshire and Norway, and its affinities are more likely to lie with the
group typified by Chasmops conicophthalma (Sars & Boeck), a species redescribed by
Stormer (1940 : 137, pi. 3, figs. 1-6) and characteristic of part of the Caradoc Series
in Norway prior to the appearance of C. extensa.
Family HOMALONOTIDAE Chapman, 1890
Genus B RON GN I ART ELL A Reed, 1918
TYPE SPECIES. Homalonotus Usulcatus M'Coy, 1851 by original designation of
Reed (1918 : 322).
Brongniartella minor (Salter)
(PI. 15, fig. 4 ; PI. 16, fig. n)
1852. Homalonotus bisulcatus var. /3 minor Salter, p. v.
1947. Brongniartella parva Harper, p. 165, pi. 6, figs. 7, 8.
19610. Brongniartella minor (Salter) Dean, p. 351, pi. 54, fig. 6 ; pi. 55, fig. n.
A few specimens found at one locality in the Inlier agree in all respects with Salter's
species, the lectotype of which, a pygidium, was originally described from the
Longvillian Stage of the Bala district, North Wales. Although the outer surface of
the test, when preserved, is apparently smooth, internal moulds show a conspicuous
granulation due to the infilling of small canals within the test. A similar feature
is found in Brongniartella ascripta (Reed), described elsewhere in this paper.
HORIZON AND LOCALITY. Upper part of the Corona Beds, Lower Longvillian
Substage, Bancroftina typa Zone, at Pus Gill, locality A. 3 and Harthwaite Sike, £.3.
Brongniartella ascripta (Reed)
(PI. 15, figs, i, 2, 5, 8, ii ; PI- 16, fig. 14?)
1910. Homalonotus ascriptus Reed, p. 216, pi. 17, figs. 4-8.
1918. Homalonotus ascriptus Reed : Reed, p. 272.
I959a. Brongniartella ascripta (Reed) Dean, p. 214.
Reed's description was founded on a single cranidium, which may be taken as
the holotype, supplemented by three additional cranidia and a hypostoma, all
THE TRILOBITES OF THE CARADOC SERIES 107
of which he regarded only as belonging doubtfully to the species. There seems to be
little doubt that these specimens belong to B. ascripta. The holotype cranidium is
somewhat flattened but retains, nevertheless, a longitudinal, median ridge which is
particularly well shown on the anterior half of the glabella. In the same specimen
crushing has produced what appear to be large, basal glabellar lobes, but all the
uncrushed specimens available show no trace of glabellar furrows. Reed described
the test of the cranidium as being covered with closely-set " tubercles ", and con-
sidered this feature sufficient to distinguish the species from Brongniartella bisulcata,
but examination of additional material suggests that the so-called tubercles are, in
fact, infillings of canals within the original test, the outer surface of which was
probably smooth. The straight lateral and frontal margins of the glabella, with
the fairly well-defined anterior border and the median carina, suggest that B.
ascripta may be grouped with forms such as Brongniartella minor (Salter) and B.
minor subcarinata Dean, rather than with B. bisulcata and B. caradociana (Dean,
1961^ : 349). The presumed hypostoma of B. ascripta is quite distinct from that of
B. bisulcata (see Dean, 1961^, pi. 54, fig. 9), and such a difference may eventually
prove to be of subgeneric, or even generic, significance. The thorax is known only
from isolated segments which are not distinguishable from those of B. bisculata.
The presumed type-locality has yielded a few pygidia believed to belong to the species.
The largest and most complete of these, BM. In. 52980, is slightly distorted and has
a median length of about 17 mm. The pleural lobes are well segmented with seven,
and perhaps eight pairs of pleural furrows, the first pair being noticeably deeper
than the rest ; the furrows extend to the lateral margin, near which they become
shallower and almost obsolete. The axis of this specimen is not sufficiently well
preserved for an accurate assessment of the number of axial rings, but another,
incomplete pygidium shows that there are at least seven, and probably more.
HORIZON AND LOCALITIES. It is believed that Reed's original material was
obtained from locality J near the Alston Road, north-east of Melmerby ; the rocks
there are thought to belong to the Lower Melmerby Beds, of probable Lower Long-
villian age. Elsewhere along the Alston Road fragments of the species have been
collected at localities D, E, F and G, from the Lower Melmerby Beds.
In the main portion of the Inlier, specimens tentatively assigned to B. ascripta
have been found at locality E. 3, Harthwaite Sike, near Dufton (see Text-fig. 4),
in the upper part of the Corona Beds, belonging to the Bancroftina typa Zone of the
Lower Longvillian. A fragmentary cranidium from this locality is figured (PL
16, fig. 14), and a large, distorted pygidium, BM. In. 54651, from the same horizon
at locality E. 5 shows traces of eleven axial rings.
In their account of the Cross Fell Inlier Nicholson & Marr (1891 : 510) recorded
" Homalonotus rudis Salter? " from the Corona Beds at Roman Fell, and the speci-
mens were subsequently given the provisional name of Brongniartella cf. rudis
(M'Coy) (Dean, 19590; : 209). More recently it has been pointed out that Homa-
lonotus rudis, described originally by M'Coy (in Sedgwick & M'Coy, 1851 : 168)
from North Wales, is not a recognizable species (Dean, 1961^ : 355). Nicholson &
Marr's specimens from Roman Fell, housed in the Sedgwick Museum and numbered
A. 32906-12, are figured here for the first time (see PL 15, figs. 3, 6, 7, 9, 10). They
io8 THE TRILOBITES OF THE CARADOC SERIES
are too distorted tectonically for certain identification but are undoubtedly generally
similar to B. ascripta. The pygidia show eleven or twelve axial rings, and one
specimen has eight pairs of pleural ribs, though exaggerated by crushing. The
best-preserved cranidium has proportions and a glabellar outline like B. ascripta
but lacks the median carina, though this could well be due to tectonic causes In
the absence of more definite specific features, it has been preferred to figure the
Roman Fell specimens merely as Brongniartella sp. B. ascripta has not yet been
recorded outside the Cross Fell Inlier.
HOLOTYPE. Sedg. Mus. A. 29632 (PL 15, fig. n).
Brongniartella bisulcata (M'Coy)
(PI. 16, fig. 8)
1851. Homalonotus bisulcatus M'Coy in Sedgwick & M'Coy, p. 168, pi. iG, figs. 26, 27.
19613. Brongniartella bisulcata (M'Coy) Dean, p. 346. This reference contains a full synonymy
of the species.
Brongniartella bisulcata has been redescribed in detail from the type-area in south
Shropshire (Dean, 1961^), and the material from the Cross Fell Inlier exhibits no
features not already known. The earliest undoubted occurrence is in the upper half
of the Upper Longvillian Substage at Swindale Beck, localities B. 5, 8 and 9. Frag-
mentary evidence suggests that it may also be found at the same horizon at Harth-
waite Sike, locality E. 6. As in Shropshire, B. bisulcata is fairly common in the
Marshbrookian Stage, at which level it is known from both Swindale Beck, localities
B. n, 14 and 16, and Harthwaite Sike, localities E. 7, 9, 10, n and 12. In the
north of the Inlier, at the Alston Road outcrop, near Melmerby, fragments of
Brongniartella occur in the lower part of the Upper Longvillian, but have not proved
satisfactorily identifiable. They are undoubtedly close to B. bisulcata but may yet
prove to belong to B. ascripta, a species originally described from adjacent and
probably earlier strata.
Brongniartella depressa sp. nov.
(PI. 16, figs, i, 7, 10, 13)
!959«- Brongniartella sp. nov., Dean, p. 208.
DIAGNOSIS. Brongniartella with glabella of low convexity, poorly defined both
laterally and frontally, where outline narrows towards transversely-truncated,
frontal, glabellar lobe. Occipital furrow almost absent from external mould. Axial
furrows of thorax almost obsolete. Pygidial axis broad, poorly defined ; axial
rings and pleural ribs scarcely discernible.
DESCRIPTION. The cephalon is known only from a flattened cranidium, the
median length of which is slightly less than half the maximum breadth. The
glabella is slightly longer than wide, poorly defined laterally by markedly shallow
axial furrows which are shallowest and straight frontally, but which then curve
outwards and back from about their mid-points to the ends of the occipital ring.
The dimensions of the holotype cranidium are as follows : max. length 16 mm.,
THE TRILOBITES OF THE CARADOC SERIES 109
max. breadth 30 mm. (estimated) ; length of glabella 12-5 mm., max. breadth n
mm., min. breadth 7-5 mm. The convexity of the glabella is remarkably low and the
frontal glabellar lobe has a noticeably square appearance, being truncated by a poorly
impressed, transversely straight, preglabellar furrow which meets the axial furrows
almost at right-angles, the anterolateral angles of the frontal lobe being slightly
rounded. The external mould of the holotype shows no trace of glabellar furrows
but a presumably immature specimen (PL 16, fig. 13) carries three pairs of faintly-
marked, almost equispaced, glabellar furrows on the internal mould, whereas the
external mould of the same specimen shows no sign of lobation. The anterior
border is slightly inclined forwards, transversely straight, uniformly broad (sag.),
and is not well differentiated from the frontal glabellar lobe. The occipital furrow
of the holotype is practically obsolete and the dorsal surface of the glabella is almost
continuous with that of the occipital ring ; these two structures are uniformly
broad (tr.) and the occipital ring is not obliquely truncated distally as, for example,
is that of Brongniartella bisulcata (M'Coy). The internal mould of an immature
cranidium, mentioned earlier, has a transversely straight, moderately-deep, occipital
furrow (see PI. 16, fig. 13) but the external mould of this specimen is almost smooth,
comparable with the holotype. On both internal and external mould the pleuroc-
cipital furrow is only moderately impressed, curving gently forwards laterally and
delimiting a pleuroccipital segment of almost uniform breadth (exsag.). The genal
angles are imperfectly known but appear to be rounded. Frontally the fixigenae
are narrow, constricting forwards slightly and almost coalescing with the distal ends
of the anterior border, from which they are separated by only shallow depressions
representing the extension of the preglabellar furrow The posterior portions of
the fixigenae are moderately declined, their breadth (exsag.) between one-quarter
and one-third of the glabellar length. The palpebral lobes are small, short, situated
opposite the mid-point of the glabella, and are moderately declined towards the
axial furrows. The anterior branches of the facial suture are straight, converging
forwards slightly so as to cut the anterior margin just outside the line of the axial
furrows. The posterior branches are transversely straight at first but quickly
turn backwards strongly to cut the genal angles in a normal gonatoparian position.
The eyes, librigenae and hypostoma are not known.
Only one specimen of the thorax attributed to this species is known ; it is in-
complete but there are traces of at least ten thoracic segments. The axis is only
slightly convex, occupies approximately one-third of the total breadth, and is
defined by almost obsolete axial furrows. The pleurae are transversely straight for
the most part, but turn down and backwards distally, ending in bluntly-rounded
tips. Each pleura carries a large anterolateral facet, delimited by a narrow, raised
ridge running from the anterior margin, approximately mid-way between the axial
furrow and the fulcrum, towards the posterolateral corner of the tip. Just outside
the axial furrow, the frontal margin of each pleura is cut by a slit-like pleural furrow
which runs outwards and slightly back so as to intersect the frontal margin mid-way
between the fulcrum and the pleural tip.
One, almost complete pygidium ascribed to B. depressa is known, preserved as an
external mould (PI. 16, fig. 10). It is of depressed form, and broader than long in
no THE TRILOBITES OF THE CARADOC SERIES
the ratio 4 : 3 as far as measurements are possible. The frontal margin is moderately
convex forwards and the posterior margin is broadly subparabolic in plan, with a
rounded tip. The axis is broad, more than one-third the frontal breadth, denned
laterally by faint axial furrows which are moderately convergent posteriorly. There
are six or seven, almost indistinguishable, axial rings, followed by a short terminal
piece which tends to merge into the posterior border, and there is no postaxial ridge.
The pleural lobes are almost smooth, showing only traces of five pairs of pleural
furrows, of which the first pair is the most clearly defined.
Internal moulds show a fine granulation probably resulting from the infilling of
small canals within the test, but the exterior of the exoskeleton, though apparently
smooth, proves, on magnification, to be markedly and finely punctate, the puncta-
tion extending even across the various furrows impressed on the dorsal surface.
HORIZON AND LOCALITY. A thin band of fossiliferous, weathered, impure lime-
stone in the higher beds of the Dufton Shales, Pusgilhan Stage, at Swindale Beck,
locality B. 25.
HOLOTYPE. BM. In. 49882 (PL 16, fig. i).
PARATYPES. BM. In. 49881 (PL 16, fig. 7) ; In. 49884 (PL 16, fig. 10) ; In.
49885 (PL 16, fig. 13).
DISCUSSION. The flattened form and lack of well-defined furrows on the external
surface of the test readily distinguish Brongniartella depressa from any known species
of the genus. StratigraphicaUy earlier species such as B. ascripta and B. minor
subcarinata frequently exhibit a median carination of the glabella, the latter, to-
gether with the thoracic and pygidial axes, being well differentiated by axial furrows,
whilst the pleural lobes of the pygidium are better segmented than is the case for
B. depressa. The glabella of B. bisulcata undergoes a marked constriction at about
its mid-point, and the anterolateral portions of the frontal glabellar lobe are less
angular than those of B. depressa. The glabella of Brongniartella platynota (Dalman,
1828 : 135 ; Kielan, 1959 : 116, pi. 19, figs. 1-3), like that of the Pusgillian form,
shows glabellar furrows only on the internal mould, and the occipital furrow is
shallow externally, but the Swedish species has larger fixigenae, a better differentiated
glabella which is markedly constricted frontally, and both the ring and pleural
furrows of the pygidium are more strongly developed.
Brongniartella aff. platynota (Dalman)
(PL 16, fig. 5)
A single pygidium apparently distinct from coexisting forms in the Pusgillian
Stage has been collected from the Dufton Shales in Swindale Beck, locality B 30.
It is incomplete, preserved as an internal mould, and possesses at least seven axial
rings and five pairs of pleural furrows. The axis is well defined by straight, moderately-
deep, axial furrows which are convergent posteriorly, and the pleural lobes
are moderately declined laterally. The anterior margin is convex forwards and the
pleural ribs are directed backwards, but these features have almost certainly been
exaggerated by crushing. Comparison with described species of Brongniartella is
difficult, but the appearance is generally reminiscent of B. platynota (Dalman,
THE TRILOBITES OF THE CARADOC SERIES in
1828 : 135), a form originally described from the Ashgill Series of Sweden and since
redescribed by Kielan (195 : 116, pi. 19, figs. 1-3). Each pygidium has a strongly
convergent axis and well-defined pleural furrows, and as far as can be judged the
number of axial rings and pleural ribs is generally similar.
Family CALYMENIDAE Burmeister, 1843
The first serious attempt to subdivide the trilobites composing this family was
made by Shirley who, in 1936, erected several new species and genera, and reviewed
previous work on the group. Shirley's genera were later regarded by Richter
(1937) as subgenera of Calymene, but most are now accepted as being of generic
rank. The genus most abundantly represented in the Caradoc Series of the Anglo-
Welsh area is that usually referred to as Flexicalymene Shirley, 1936, but recently
Fisher (1957 : 13) has stated that Orimops Rafinesque, 1832, has precedence over
the former, and it is necessary to investigate this claim further.
Several North American trilobite species were introduced by Green (1832) who,
although he did not figure them, nevertheless gave valid diagnoses and provided
sets of plaster casts of his original specimens. Among them was Calymene callice-
phala (Green, 1832 : 31), said to be from Hampshire, Virginia, a species which
Rafinesque (1832 : 72), later in the same year, considered to be sufficiently distinct
as to warrant the formation of a new subgenus of Calymene which he named
Orimops, though he misspelt the specific name as calicephala. Many of Green's
species are readily recognizable from their plaster casts, but that of Calymene callice-
phala is not one of the best, and the species is obviously in need of redescription.
Consequently Orimops must be regarded as a nomen dubium, and may eventually
prove to be best rejected. Nevertheless, the cast of C. callicephala shows enough
of the structure of the cephalon to suggest that the species is similar in many respects
to the well-known Cincinnatian trilobite Calymene meeki (Foerste, 1910 : 84, pi. 3,
fig. 18 ; 1919, pi. 18, fig. 3), a form usually put in Flexicalymene, and in his original
description of C. callicephala Green (1832 : 31) stated that his species occurred near
Cincinnati, Ohio, though not at Trenton Falls. Both species are characterized by a
markedly triangular glabellar outline, the sides of which are strongly convergent
frontally to a narrow, frontal, glabellar lobe, and by the forward position of the
eyes which, as in Platycalymene, are sited opposite the first pair of glabellar lobes.
These two species, together with a closely similar form in the Ashgill Series of Scot-
land (Shirley, 1936, pi. 29, fig. 8), are regarded here as forming a compact group,
both geographically and stratigraphically distinct. The name Orimops, if it is to
be employed at all, is best used for these species, though the erection of a new name
may prove necessary.
The genus Flexicalymene was founded by Shirley (1936 : 395) on the well-known
south Shropshire trilobite Calymene caractaci Salter, 1865, one of several species
in which the eyes are situated opposite the second glabellar lobes. Fisher's (1957,
pi. 33, fig. 12) use of Orimops to include the Trenton trilobite Calymene senaria
Conrad, 1841, even if the genus were not under suspicion, can hardly be justified
and this species is regarded here as belonging to Flexicalymene (s.s.).
ii2 THE TRILOBITES OF THE CARADOC SERIES
Shirley (1936 : 390-392) regarded the position of the eyes as being constant in
most calymenids, and he supposed Flexicalymene onniensis, with the eyes situated
far back, opposite the third glabellar furrows, to be exceptional. During the present
work it has been noted that F. onniensis is only one of a number of widely distributed
species, again falling within fairly well-defined geographical and stratigraphical
limits, in which the eyes are similarly placed, and it is proposed now to separate
these as a new genus to which the name Onnicaiymene is given, with type species
Flexicalymene onniensis Shirley.
Reacalymene was separated by Shirley (1936 : 395) as a distinct genus and a
similar course has been followed by Whittard (1960 : 152, 158). In practice, how-
ever, whilst the ridging of the preglabellar field which Shirley claimed as a dia-
gnostic feature is generally obvious, though never strongly developed, on the external
mould, when only an internal mould is available all trace of the ridge frequently
disappears and it is then impossible to distinguish the specimen from a typical
Flexicalymene. For this reason it is thought better to follow the course adopted
by Whittington (in Moore, 1959 : 0.452) in regarding Reacalymene as being of sub-
generic status within the genus Flexicalymene. As far as is now known, Reacalymene
has a relatively restricted vertical range in the Anglo-Welsh area, perhaps through
only the lower part of the Caradoc Series. Shirley (1936 : 390) postulated that the
first stages in the thickening of the anterior border of the Calymenidae was to be
found in Reacalymene pusulosa Shirley, from the early Caradoc strata of south
Shropshire. This now appears to be an oversimplified view as Gravicalymene, with
a much more thickened border, is known from the lowest Harnagian Stage, where it
is represented by G. praecox (Bancroft), and may have existed even earlier.
The position of the calymenids normally placed in Flexicalymene may be
summarized as follows :
Flexicalymene Shirley, 1936
DIAGNOSIS. Glabellar outline generally subparabolic, rounded frontally. An-
terior border moderately long, smooth, usually steeply inclined forwards. Pal-
pebral lobes situated opposite, or almost so, second glabellar lobes. Thorax with
thirteen segments (but see later).
DISTRIBUTION. Llandeilo Series to Caradoc Series, probably to highest Marsh-
brookian Stage. Widely distributed geographically, occurring in England, Wales,
Scotland, Ireland, Bohemia and eastern North America.
TYPICAL SPECIES. F. acantha (Bancroft), F. brevicapitata (Porlock), F. cambrensis
(Salter), F. caractaci (Salter), F. declinata (Barrande), -F. forcipata (M'Coy), F.
planimarginata (Reed), F. senaria (Conrad), F. shirleyi Tripp.
Onnicaiymene gen. nov.
DIAGNOSIS. Generally similar to Flexicalymene, but distinguished by anterior
border, which is short and steeply inclined forwards, and by position of palpebral
lobes, opposite third glabellar furrows and frontal portion of third glabellar lobes.
Thorax with thirteen segments.
THE TRILOBITES OF THE CARADOC SERIES 113
DISTRIBUTION. First encountered in the Actonian Stage of the Caradoc Series
in the Anglo- Welsh area, Onnicalymene ranges upwards at least into the Pusgillian
Stage. Its range in Ashgill strata is not known. Found in England, Wales,
Sweden and southern Norway.
TYPICAL SPECIES. 0. jemtlandica (Thorslund), 0. laticeps (Bancroft), 0. onniensis
(Shirley), 0. salteri (Bancroft).
Flexicalymene (Reacalymene) Shirley, 1936
DIAGNOSIS. Generally similar to Flexicalymene (s.s.) but anterior border bears
low, transverse ridge, visible at least on external mould, and glabella is proportion-
ately broader across third glabellar lobes, with sides more convergent forwards,
producing triangular outline.
DISTRIBUTION Only confirmed from lower part of Caradoc Series in Welsh
Borders, but possibly also in Llandeilo Series.
TYPICAL SPECIES. F. (R.) limba (Shirley), F. (R.) pusulosa (Shirley).
British Species of Doubtful Systematic Position
Calymene quadrata (King, 1923 : 504, pi. 26, figs, i, 2), from the Ashgill Series,
is generally referred to Flexicalymene but differs considerably from that genus in
its glabellar outline, and is unique in having only twelve thoracic segments. Its
position and affinities are therefore regarded as being doubtful.
Calymene (Diacalymene) bigener (Reed, 1935 : 47, pi. i, fig. 3), from the Ashgill
Series of Girvan, has been placed tentatively in Reacalymene by Shirley (1936 : 418),
but such a position is doubtful and the species needs to be redescribed.
Calymene (Ptychometopus) grayae (Reed, 1935 : 45, pi. i, figs. 13, 19?), from the
Craighead Limestone, cannot be considered a recognizable species.
Calymene (Colpocoryphe^} aldonensis (Reed, 1935 : 48, pi. i, figs. 14, 15), from the
Didymograptus super stes Shales (= Nemagraptus gracilis Zone) of the Girvan district,
has been placed in Flexicalymene by Shirley (1936 : 418), but the position of the
eyes is reminiscent of that in Onnicalymene, whilst the glabellar outline is unlike
that of any other described species of Flexicalymene.
Diacalymene and Gravicalymene
The typically Upper Ordovician or Silurian genus Diacalymene Kegel, 1927 has
been reported from the Caradoc Series of south Shropshire, supposedly being repre-
sented by D. praecox Bancroft, 1949, but the species in question is considered here
to be better placed in Gravicalymene Shirley, 1936. The two genera bear a consider-
able resemblance to one another in glabellar outline and it is sometimes difficult
to differentiate between the respective anterior borders, which were described by
Shirley (1936 : 395) as being " roll-like " in Gravicalymene and " ridged " in Dia-
calymene. The writer's observations suggest that whereas the type species of
Diacalymene, D. diademata (Barrande), from the Silurian of Bohemia, possesses a
distinctive, ridged, steeply-inclined anterior border, those Ordovician species
GEOL. 7, 3. 6
H4 THE TRILOBITES OF THE CARADOC SERIES
customarily assigned to the genus have an anterior border which is not markedly
different from that of Gravicalymene, a genus which did not undergo much modifica-
tion even as late as the Devonian. A more important criterion for separating the
two genera is the presence or absence of what Shirley called " papillate second
glabellar lobes ", accompanied by " buttresses " on the fixigenae opposite the same
lobes. Stumm & Kaufmann (1958 : 949) have drawn attention to the unsuitability
of this terminology, and it is suggested that one might more appropriately speak of
" conjugate second glabellar lobes and genal buttresses " in Diacalymene. Hupe
(1955 : 245) has attempted to place Diacalymene and Gravicalymene in separate
subfamilies, respectively Calymeninae and Colpocoryphinae, on the basis of their
having unconstricted or constricted axial furrows, but such a course seems obviously
unworkable. Diacalymene diademata differs in some respects from the described
Ordovician species of the genus, the latter in turn resembling Gravicalymene in
glabellar outline. It seems likely that the two genera are closely related, and may
have diverged from a common ancestral group earlier in the Ordovician, but more
evidence is needed. As restricted in this paper, Diacalymene makes its appearance
in the Anglo-Welsh area in the Pusgillian Stage of the Cross Fell Inlier, being un-
known in Shropshire ; it becomes more abundant in the succeeding Ashgill Series
and continues into the lower Silurian. Gravicalymene, though ranging as high as the
Devonian, is an earlier genus, known with certainty from the Harnagian Stage of
the type Caradoc succession but possibly originating still earlier. In Caradoc
strata both genera are subordinate in numbers to the widespread Flexicalymene
and Onnicalymene.
Genus FLEXICALYMENE Shirley, 1936
TYPE SPECIES. Calymene Blumenbachii var. Caractaci Salter, 1865 by original
designation of Shirley (1936 : 395).
Flexicalymene cf. caractaci (Salter)
(PI. 13, fig. 10)
i959«. Flexicalymene cf. caractaci (Salter) : Dean, p. 214.
Calymenid remains have been found at several points in the vicinity of the Alston
Road near Melmerby, and some of the better preserved are generally comparable
with Salter's species. The type-material of F. caractaci came from the Upper
Cheney Longville Flags, Marshbrookian Stage, of south Shropshire and has been
redescribed by Shirley (1931 : 25). In the Cross Fell Inlier only occasional frag-
ments of calymenid trilobites have been collected from the Marshbrookian Stage,
and none has proved identifiable. The strata at Melmerby are probably both Lower
and Upper Longvillian in age, and the relevant localities there comprise C to J
inclusive, and possibly also locality A (see Text-fig. 5). The lectotype of Flexi-
calymene planimarginata (Reed), redescribed by Shirley (1931 : 22) and Harper
(1947 : 167), originated from the Lower Longvillian of North Wales. The species
has not yet been reported from the Cross Fell Inlier at this horizon, though
THE TRILOBITES OF THE CARADOC SERIES 115
Whittard (1960 : 158) has recorded it earlier, from the Soudleyan Stage of the
Shelve Inlier.
Genus ONNICALYMENE nov.
TYPE SPECIES. Flexicalymene onniensis Shirley, 1936. For diagnosis of genus
see p. 112.
Onnicalymene onniensis (Shirley)
(PI. 13, fig. 7 ; PL 14, figs, i, 2, 7, 10)
1936. Flexicalymene onniensis Shirley, p. 405, pi. 29, figs. 5-7.
1949. Flexicalymene onniensis Shirley : Bancroft, p. 308, pi. n, figs. 36, 37.
1958. Flexicalymene onniensis Shirley : Dean, p. 224.
I959&. Flexicalymene onniensis Shirley : Dean, pp. 200, 207.
I959&- Flexicalymene aff. onniensis Shirley : Dean, pp. 202, 208.
This well-known Shropshire species occurs in abundance in the Dufton Shales
of the Knock-Dufton district of the Cross Fell Inlier. The earliest specimens are
from the Onnia gracilis Zone at Pus Gill, localities A. 8, 9, n, 12, 14 and 15, and the
species has been found less commonly in the Onnia superba Zone, also at Pus Gill,
locality A. 5. Specimens are relatively abundant in rocks of the Pusgillian Stage,
and do not appear to be restricted to any particular part of the stage. Localities
are numerous and include : Pus Gill, localities A. i, 6, 18, 21, 24, 25-30 ; Swindale
Beck, localities B. 22, 24-26, 28, 33, 34 ; Dufton Town Sike, localities C. 2-4 ;
and Hurning Lane. Judging from the matrix of the Pus Gill specimen figured by
Shirley (1936, pi. 29, fig. 7) it probably originated from the Pusgillian Stage.
Onnicalymene laticeps (Bancroft)
(PL 14, figs. 5, 6)
1949. Flexicalymene laticeps Bancroft, p. 307, pi. n, figs. 33, 330.
i959«. Flexicalymene cf. laticeps Bancroft : Dean, pp. 199, 207.
Fragments referable to this characteristic south Shropshire form have been
recovered from locality B. 15 in the bed of Swindale Beck, near the eastern bank,
and the best preserved, a cephalon with three attached thoracic segments, is figured
here. In Shropshire 0. laticeps has been found only in the Actonian Stage, particu-
larly the middle third, and the associated fauna at Swindale Beck, comprising
Chasmops cf. extensa (Boeck), Remopleurides sp. (see p. 127), Chonetoidea, Onniella
and Sowerbyella, suggests a similar horizon. Like other members of Onnicalymene,
0. laticeps has the palpebral lobes sited well back, opposite the third glabellar
furrows, but is easily separated specifically by the short glabella with its well-
rounded, parabolic outline. Bancroft's original account of the species did not
describe the pygidium. Two pygidia, numbered BM. In. 50259 and In. 50260,
have been found in association with the figured cephalon at Swindale Beck, and are
of characteristic calymenid form. The axis has five well-defined axial rings, with
n6 THE TRILOBITES OF THE CARADOC SERIES
faint traces of a sixth, followed by a short terminal piece. Each pleural lobe carries
five pleural ribs with faintly impressed rib furrows which deepen distally ; the ribs
are separated from each other by deep pleural furrows running from the axial furrows
to the lateral margins.
Genus DIACALYMENE Kegel, 1927
TYPE SPECIES. Calymene diademata Barrande, 1852 by original designation of
Kegel (1927 : 617).
Diacalymene cf. marginata Shirley
(PI. 13, fig. 13 ; PI. 14, fig. ii)
1936. Diacalymene marginata Shirley, p. 415, pi. 29, figs. 19, 20.
*959a- Diacalymene cf. marginata Shirley : Dean, pp. 204, 208.
Diacalymene marginata was described from the Drummuck Group, Ashgill Series,
of the Girvan district by Shirley, who also recorded the species from the Ashgill
of North Wales and the Cautley district of Yorkshire. The specimens from the
Cross Fell Inlier are closely similar to the holotype in most respects, though the
glabella may be slightly broader frontally and the anterior border situated a little
closer to the frontal glabellar lobe. Such differences may well be due to tectonic
causes. This form is known from the topmost beds of the Pusgillian Stage at
locality B. 25, and perhaps also B. 34, in Swindale Beck. At both places the horizon
is not far below the disconformable base of the Swindale Limestone. Elsewhere in
the Inlier D. cf. marginata has been found at locality A. 27 in Pus Gill ; the horizon
here is also in the Pusgillian Stage, but is probably lower than those at Swindale
Beck, though the relationships of the strata are obscured by strike-faulting.
Genus GRAVICALYMENE Shirley, 1936
TYPE SPECIES. Gravicalymene convolva by original designation of Shirley (1936 :
395).
Gravicalymene jugij era sp. nov.
(PI. 13, figs. 9, ii ; PI. 14, figs. 3, 4, 8, 9)
I959«- Gravicalymene sp. nov., Dean, pp. 204, 208.
DIAGNOSIS. Gravicalymene with well-rounded frontal glabellar lobe, and glabella
which is relatively narrow for genus. Anterior border of cephalon flattened dorsally,
its posterior margin ridged, separated by deep, broad (sag.) furrow from glabella.
Pygidium of characteristic calymenid shape with five axial rings and five pleural
ribs.
DESCRIPTION. The cranidium is moderately convex, both longitudinally and
transversely, and about twice as broad as long. The glabella is a little broader
than long, its sides gently convergent forwards for the most part, but the breadth
increases noticeably at the line of the basal glabellar lobes. The frontal glabellar
THE TRILOBITES OF THE CARADOC SERIES 117
lobe is short, its anterior margin well rounded. There are three pairs of lateral
glabellar lobes, those of the first pair being small, tubercle-like, bounded frontally
by shallow, slit-like, first glabellar furrows and posteriorly by wider (exsag.), deeper,
second glabellar furrows. The second glabellar lobes are larger than those of the
first pair, and subcircular in plan. The third glabellar furrows are deep, at first
directed slightly backwards from the axial furrows but then bifurcating markedly,
the anterior branch so formed being the shorter, curving adaxially forwards around
the second glabellar lobes. The posterior branches are deeper and turn backwards
towards, though they do not attain, the occipital furrow. The third, or basal
glabellar lobes are subangular in plan with gently convex distal margins, bluntly
pointed frontally, and are constricted proximally by the posterior branches of the
third glabellar furrows. The axial furrows are deep, gently sigmoidal in plan, and
of generally uniform breadth except posteriorly, where they narrow slightly as
they flex around the basal glabellar lobes. There are deep hypostomal pits situated
midway between the first glabellar furrows and the line of the front of the glabella.
There is no true preglabellar field or furrow present, and the portion of the cranidium
immediately in front of the glabella is continuous with the anterior border, the whole
being flexed upwards through about a right-angle. The structure so formed carries
a ridge, moderately defined medially but dying out laterally, which marks the
junction of the smooth, broad (sag.), deep furrow immediately in front of the glabella,
and the granulate, flattened, moderately-inclined top of the anterior border. The
occipital ring is transversely convex, broadest (sag.) medially, becoming narrower
laterally towards a pair of poorly-defined occipital lobes. The occipital furrow is
shallow and transversely straight medially, deepening laterally where it curves
backwards slightly to encompass the basal glabellar lobes. The pleuroccipital
furrow is broad (exsag.}, of moderate depth, becoming shallower abaxially ; the
pleuroccipital segment is narrow proximalTy, broadening towards the rounded
genal angles. The anterior parts of the fixigenae are of uniform breadth, converging
forwards slightly, parallel to the axial furrows. The eyes are situated opposite
the second glabellar furrows and the front half of the second glabellar lobes. The
visual surface has not been found preserved, and the palpebral lobes are moderately
declined adaxially, with only traces of palpebral furrows. Poorly-developed eye
ridges run obliquely forwards from the palpebral lobes, ending at the axial furrows
opposite the first glabellar lobes. The librigenae are of characteristic calymenid
type, steeply declined laterally. The hypostoma is unknown.
The thorax is represented by only one crushed example in which the number
of thoracic segments is not visible. In spite of the poor state of preservation the
segments do not appear to differ significantly from those of other calymenids.
In plan the pygidium is broadly kite-shaped, the anterior margin strongly curved
forwards and the lateral margins straight, converging backwards to a blunt point.
The axis is convex dorsally, defined laterally by deep axial furrows extending almost
to the tip. Excluding the articulating half-ring there are four well-developed axial
rings, separated by deep, transversely-straight ring furrows, with a fifth less obvious
axial ring, followed by a relatively long terminal piece extending almost to the tip
of the pygidium, to which it is linked by a suggestion of a low postaxial ridge. The
n8 THE TRILOBITES OF THE CARADOC SERIES
pleural lobes carry four pairs of well-developed pleural ribs which become progres-
sively less divergent backwards, with traces of a fifth pair almost parallel to one
another. The ribs of the first pair have interpleural furrows only on the distal
half of the dorsal surface, but on the remaining pairs the interpleural furrows extend
from just outside the axial furrows to the lateral margins, dividing each rib into two
bands, of which the anterior is the narrower (exsag.) .
Excluding the furrows, which are apparently smooth, the surface of internal
moulds is generally finely granulate, representing the infilling of canals within the
original test. A little of the test remains on the holotype cranidium, and the in-
clined, upper surface of the anterior border is finely granulate whilst that of the
broad (sag.) furrow immediately in front of the glabella is smooth.
HORIZON AND LOCALITIES. All the known occurrences of Gravicalymene jugifera
are in Dufton Shales belonging to the Pusgillian Stage. The holotype is from
Swindale Beck, locality B. 28, but the species is known also from Pus Gill, locality
A. 24 ; Swindale Beck, localities B. 20, B. 30 and, with some doubt, B. 25 ; Dufton
Town Sike, locality C. 6 ; Billy's Beck, locality D. i ; and at Hurning Lane, 700
yards north-east of St. Cuthbert's Church, Dufton.
HOLOTYPE. BM. In. 50263 (PI. 14, figs. 3, 4, 8).
PARATYPES. BM. In. 50245 (PI. 13, figs. 9, n) ; In. 55897.
DISCUSSION. Gravicalymene jugifera. seems to be closely related to the type-
species of the genus, G. convolve, which was described from the Ashgill Series of
Birdshill Quarry, near Llandilo (Shirley, 1936 : 409, pi. 29, figs. 16-18). The new
species may, however, be distinguished by its slightly narrower glabellar outline,
smaller basal glabellar lobes, more dorsally flattened anterior border, and by the
greater breadth (sag.) of the furrow separating the anterior border from the frontal
glabellar lobe. Both species have a glabellar outline remarkably close to that
found in certain Ordovician species of Flexicalymene (s. 1.) described by Stumm &
Kaufmann (1958 : 951, 953, pi. 123, figs. 14, 15 ; pi. 124, figs. 2-16) from Michigan,
but are distinguished by the different structure of the anterior border.
Family DIMEROPYGIDAE Hupe, 1953
Genus TOERNQUISTIA Reed, 1896
TYPE SPECIES. Cyphaspis (Tornquistia) nicholsoni (Reed, 1896 : 433) by mono-
typy.
Toernquistia aff. reedi Thorslund
(PI. 16, figs. 2, 3)
1910. Tornquistia nicholsoni (Reed) Reed, p. 211.
1940. Tornquistia reedi Thorslund, p. 148, pi. 10, figs. 14-19 ; pi. n, fig. 15.
Only a small, incomplete cranidium preserved as an internal mould is available.
The calculated length is estimated to have been about 2 mm. and the breadth across
the palpebral lobes 2-3 mm. The glabella is subparabolic in outline, moderately
convex, bounded laterally by deep, narrow, axial furrows which are continuous
THE TRILOBITES OF THE CARADOC SERIES 119
frontally with the preglabellar furrow. There are no glabellar furrows. The fixi-
genae are narrow, lower than the glabella and continuous frontally with the pre-
glabellar field. The latter is broad (exsag.) distally, but narrows towards the median
line where a deep pit is situated in the preglabellar furrow ; from this pit a slot-
like depression extends forwards almost half-way across the steeply-declined,
preglabellar field. The anterior border is narrow (exsag.), almost flat, separated
from the preglabellar field by a narrow (exsag.), shallow, anterior border furrow.
The surviving left palpebral lobe, though incomplete posteriorly, is long and narrow,
separated from the fixigena by a broad, shallow, palpebral furrow. In front of the
palpebral lobes the anterior branches of the facial suture diverge strongly towards
the margin. Ah1 the furrows of the cranidium are smooth, but the surface of the
anterior border is finely granulate, whilst that of the glabella, palpebral lobes and
preglabellar field is covered with small tubercles which increase in size on the distal
portions of the fixigenae.
HORIZON AND LOCALITY. Probably that denoted here by the letter J (see Text-
fig. 5), east of the Alston Road, north-east of Melmerby. The rocks probably
belong to the Lower Melmerby Beds, Lower Longvillian Substage.
DISCUSSION. In emending the Family Dimeropygidae, Whittington & Evitt
(1954 : 35) placed therein the genera Dimeropyge Opik, Dimeropygiella Ross,
Toernquistia Reed, Mesotaphraspis Whittington & Evitt, and Chomatopyge Whitting-
ton & Evitt. They noted the species described by Thorslund (1940 : 148) as
Tornquistia reedi, but considered its generic position to be uncertain owing to its
possessing a cephalon recalling that of Mesotaphraspis and a pygidium resembling
that of Chomatopyge. Until more material is available the systematic position of
the Alston Road form is debatable. Genetically it appears to be closer to Toern-
quistia than to Mesotaphraspis, whilst bearing a considerable resemblance to Thors-
lund's species, though the latter has faint glabellar furrows and a striate anterior
border. The type material of Toernquistia reedi was obtained from the limestone
facies of the Lower Chasmops Limestone, 4b/?, of Jemtland, Sweden, a horizon
which may not differ appreciably from that of the Alston Road species.
Toernquistia nicholsoni Reed sp. (1896 : 433, pi. 21, figs. 3, 30 ; see also Whitting-
ton, 1950 : 561, pi. 75, figs. 8-16), from the Keisley Limestone of Ashill age in the
Cross Fell Inlier, differs in having palpebral lobes situated farther forwards, and in
being proportionately narrower across the frontal glabellar lobe.
The trilobite originally described by Reed (1904 : 86, pi. 12, figs. 3-7) as Meno-
cephalusl (Tornquistia} cf. nicholsoni, from the Balclatchie Group of Girvan bears a
strong resemblance to that from the Alston Road, but differs from the latter in
having a more transversely straight, anterior cephalic border, as well as a pair of
deep furrows running distally from the mid-points of the axial furrows to just in
front of the palpebral lobes. The Balclatchie species was later renamed Hystricurus
translatus by Reed (1931 : 6), and placed in Toernquistia by Whittington (1950 : 563).
Hupe (1955 : 143, fig. 124, 6a, b) has since recorded " Tornquistia nicholsoni Reed "
from the Llandeilo Series of Scotland, and it must be presumed that his paper
refers to Toernquistia translata, as no genuine record of T. nicholsoni is known from
pre-Ashgill strata.
120 THE TRILOBITES OF THE CARADOC SERIES
Family ILLAENIDAE Hawle & Corda, 1847
Genus STENOPAREIA Holm, 1886
TYPE SPECIES. Illaenus linnarssoni Holm, 1882 by original designation of Holm
(1886 : 152).
Stenopareia? sp.
(PI. 16, fig. 12)
1910. Illaenus Bowmani [sic] Salter (?), Reed, p. 211.
A single pygidium recorded by Reed (1910 : 211) is the only-known representative
of the family in the Caradoc Series of the Inlier. The specimen is relatively long,
the median length only slightly less than the maximum breadth. Stenopareia
has been accepted as a distinct genus by Jaanusson (1954 : 570-572), and he has
been followed by Whittard (1961^ : 216, pi. 30, figs, 10-13) who has described a new
species, 5. camladica, from the Lower Soudleyan Stage of west Shropshire. The
pygidium of S. camladica bears a considerable resemblance to that from the Inlier,
but detailed comparison is not possible owing to lack of material.
HORIZON AND LOCALITY. Probably the Lower Melmerby Beds at locality J
near the Alston Road (see Text-fig. 5).
Family LICHIDAE Hawle & Corda, 1847
Subfamily HOMOLICHINAE Phleger, 1936
Genus CONOLICHAS Dames, 1877
TYPE SPECIES. Lichas aequiloba Steinhardt, 1874 by subsequent designation of
Vogdes (1890 : 23).
Conolichas melmerbiensis (Reed)
(PI. 17, figs. 14, 16 ; PI. 18, figs. 1-4)
1907. Lichas (Homolichas) melmerbiensis Reed, p. 396, pi. 17, figs. 1-7.
1910. Lichas melmerbiensis Reed : Reed, p. 211.
J933- Platylichas melmerbyensis (Reed) Bancroft, table 2.
1937. Tetralichas melmerbiensis (Reed) Phleger, p. 1088.
*939- Conolichas melmerbiensis (Reed) Warburg, p. 73.
1946. Platylichas melmerbiensis (Reed) : Bancroft in Lament, p. 237.
1948. Platylichas melmerbiensis (Reed) : Bancroft in Lament, p. 416.
1958. Conolichas melmerbiensis (Reed) : Tripp, p. 576.
I959#- Conolichas melmerbiensis (Reed) : Dean, pp. 213, 214.
All Reed's type-specimens were described by him merely as having been collected
from what he called the Dufton Shales near Melmerby. The manuscript catalogue
accompanying the specimens, together with their state of preservation, suggests
that the place of origin was locality J, east of the Alston Road, near Melmerby
(see Text-fig. 5), and during the present field-work two more specimens of the species
THE TRILOBITES OF THE CARADOC SERIES 121
have been obtained from this locality. The horizon is in the Longvillian Stage,
and the rocks at locality J are believed to belong to the Lower Melmerby Beds, of
probable Lower Longvillian age. It was claimed by Bancroft (1933, table 2 ; in
Lament, 1948 : 416) that what he called Platylichas melmerbiensis occurred near
Melmerby with Wattsella horderleyensis Whittington, but the latter, a zonal brachio-
pod species in the Lower Longvillian of south Shropshire and North Wales, has not
yet been confirmed in the Cross Fell Inlier. The type-specimens of Conolichas
melmerbiensis are unusual in including several whole, or nearly whole, exoskeletons,
and it may be that they occurred at only one restricted horizon where conditions
were particularly suitable for their preservation.
LECTOTYPE, here selected. Sedg. Mus. A. 29638 (PI. 18, figs. 3, 4).
PARATYPES. Sedg. Mus. A. 29637 (PL 18, fig. i) ; A. 29639 ; A. 29640 ; A.
29641 (PL 17, fig. 16) ; A. 29642 (PL 18, fig. 2) ; A. 29643 (PL 17, fig. 14).
Genus PLATYLICHAS Giirich, 1901
TYPE SPECIES. Lichas margaritifer Nieszkowski, 1857 by original designation of
Giirich (1901 : 522).
Platylichas cf. laxatus (M'Coy)
(PL 17, figs, i, 7)
I959«- Platylichas laxatus (M'Coy) (s.l.) : Dean, pp. 204, 207, 208.
Lichas laxata was described by M'Coy (1846 : 51, pi. 4, fig. 9) from southern
Ireland, and founded by him on an incomplete, compressed cranidium, now in the
National Museum of Ireland, from Ballygarvan Bridge, New Ross. The species
has since been widely quoted by a number of authors from a variety of horizons
within the Caradoc and Ashgill Series, and although it seems likely that more than
one species may be present, comparison with other forms will continue to be difficult
until a detailed redescription is available on the basis of topotype material. A long
description was given by Warburg (1939 : 118) using Scandinavian material of
Caradoc and Ashgill age. The cranidia of these specimens appear indistinguishable
from each other, and the pygidia exhibit only occasional minor differences which
may or may not be of specific significance. The specimens from the Chasmops
Limestone of Norway and Sweden cannot reasonably be separated from those now
under consideration from the Cross Fell Inlier, particularly when allowance is made
for variation within the species.
In the Anglo-Welsh Caradoc Series, Platylichas of the laxatus type has been
found as early as the Derfel Limestone (Whittington & Williams, 1955 : 424, pi.
40, figs. 113-118), whilst the Longvillian strata of North Wales contain remains
identical with Lichas nodulosus (M'Coy in Sedgwick & M'Coy, 1851 : 151, pi. iF,
fig. 16). Tripp (1958 : 579, pi. 85, fig. 5) has refigured the holotype of the latter
species and considers it to be synonymous with Platylichas laxatus, though pointing
out that it may eventually prove to merit subspecific status. The specimen is
122 THE TRILOBITES OF THE CARADOC SERIES
incomplete but exhibits, nevertheless, a remarkably close resemblance to a pygidium
figured here (PL 17, fig. 7).
The earliest-known occurrence of Platylichas in the Cross Fell Inlier is in the Upper
Melmerby Beds of the Alston Road outcrop, locality H (see Text-fig. 5). The sole
specimen found (PL 17, fig. 2) is a fragmentary pygidium, numbered BM. In. 52595,
which belongs unmistakably to the genus but which is insufficient for specific com-
parison and is accordingly named merely Platylichas sp.
The two figured specimens of Platylichas cf. laxatus (M'Coy) are from the highest
beds of the Dufton Shales, Pusgillian Stage at Swindale Beck, locality B. 25. They
are numbered BM. In. 50115 (PL 17, fig. i) and In. 50117 (PL 17, fig. 7). What is
apparently the same form is known earlier, in Dufton Shales of the Onnian Stage,
Onnia gracilis Zone, at Pus Gill, localities A. 12, 13 and 14.
Family ODONTOPLEURIDAE Burmeister, 1843
Genus PRIMASPIS R. & E. Richter, 1917
TYPE SPECIES. Acidaspis primordialis Barrande, 1852 by original designation of
R. & E. Richter (1917 : 466).
Primaspis semievoluta (Reed)
(PL 17, figs, 3, 10, n, 13, 15)
1910. Acidaspis semievoluta Reed, p. 214, pi. 17, figs. 1-3.
Primaspis semievoluta (Reed) Dean, p. 214.
All Reed's syntypes are believed to have been obtained from the Lower Melmerby
Beds, probably Lower Longvillian Substage, at locality J near the Alston Road
(see Text-fig. 5). During the present work one further specimen has been collected
from what is believed to be the same geological horizon at locality F beside the
Alston Road ; this specimen, a distorted cranidium, is in the British Museum, where
it is numbered In. 53000.
Few species of Odontopleuridae are known from the Anglo- Welsh Caradoc Series.
Primaspis harnagensis (Bancroft, 1949 : 301, pi. 10, figs. 21, 22), from the Harnagian
Stage of Shropshire, has a narrower glabellar outline than P. semievoluta, and the
pygidium has seven pairs of marginal spines, compared with five pairs on the Cross
Fell species. One of the paratypes of P. semievoluta (see PL 17, fig. 10) has a rela-
tively narrow glabella not unlike that of P. harnagensis, but this is believed to be the
result of tectonic distortion. The lectotype cranidium of Primaspis semievoluta
bears a noticeable resemblance to that of Primaspis caractaci (Salter, 1857 : 211,
pi. 6, figs. 15-17), a species found most commonly in the Actonian Stage of south
Shropshire. The glabella of the Shropshire form is, however, slightly broader across
the basal glabellar lobes, and a little more pointed frontally. The pygidia are more
distinct, that of P. semievoluta having five pairs of marginal spines, compared with
six pairs in P. caractaci. In each case the third pair of spines is longer and stouter
than the others.
LECTOTYPE, here selected. Sedg. Mus. A. 29951 (PL 17, fig. 3).
THE TRILOBITES OF THE CARADOC SERIES 123
PARATYPES. Sedg. Mus. A. 29952 (PL 17, fig. 10) ; A. 29953 (PI. 17, fig. 15) ;
A. 29954 (PL 17, fig. n).
Odontopleurid gen. et sp. ind.
Apart from the occurrence of Primaspis semievoluta (Reed) in the Alston Road
outcrop, the only other evidence of the family in the Caradoc Series of the Inlier is
an unidentifiable fragment from the Dufton Shales, Pusgillian Stage, at Dufton
Town Sike, locality C. 6 (see Text-fig. 4), collected by Mr. M. Mitchell. The speci-
men, BM. In. 51848, is an external mould showing part of the glabella with two pairs
of glabellar lobes, the right palpebral lobe, situated opposite the basal glabellar lobe,
and a wide (sag.) occipital ring carrying a small but well-defined median tubercle.
There are traces of three thoracic segments and the surface of the test is ornamented
with large closely-set granules. The general aspect is not unlike that of a species of
Primaspis but the specimen is inadequate for even generic determination.
In describing Acidaspis magnospina from the Ashgill Series of the southern Lake
District, Stubblefield (1928 : 427) stated that his new species was probably the same
as that recorded by Marr (1892 : 108) as " Acidaspis n. sp." from the " Sleddale
Group of Pusgill and Applethaite ". Earlier, Nicholson & Marr (1891 : 511), in
listing the fossils then known from the Dufton Shales, noted " Acidaspis n. sp."
from " Pusgill " and the specimen is in the Sedgwick Museum, where it is numbered
A. 32956. The species is apparently Acidaspis magnospina Stubblefield, but the
matrix is not that of the Dufton Shales ; it resembles instead the lithology of the
so-called Staurocephalus or Swindale Limestone, a horizon not known from Pus Gill
but which crops out at Swindale Beck and Billy's Beck, and the specimen is
probably mislabelled.
Family OTARIONIDAE R. & E. Richter, 1926
Genus OTARION Zenker, 1833
TYPE SPECIES. Otarion diffractum Zenker, 1833 by subsequent designation of
R. & E. Richter (1926 : 95).
Otarion sp.
(PL 17, figs. 4, 12)
? 1891. Cyphaspis megalops M'Coy (?) : Nicholson & Marr, p. 509.
i959<z Otarion sp., Dean, p. 214.
Two indifferently-preserved cranidia from the Upper Melmerby Beds, Upper
Longvillian Substage, at locality H by the Alston Road, represent the only specimens
of the genus known from the Cross Fell Inlier. The more complete example, figured
here, bears a general resemblance to the cephalon of Otarion planifrons (Eichwald),
a Baltic species from the Kuckers Stage which has been refigured by Opik (1937,
pi. 2, figs. I, 2, text-fig. 2). The proportions of the glabella, and the size and position
124 THE TRILOBITES OF THE CARADOC SERIES
of the basal glabellar lobes, is similar in both species, but the glabella of 0. planifrons
is the more coarsely tuberculate and possesses a preglabellar field which is slightly
longer. Otarion isoplates Tripp (1954 : 669, pi. 3, figs. 1-4) has a preglabellar field
and anterior border very like those of the Alston Road form but the basal glabellar
lobes are slightly larger and project farther abaxially.
Family PROETIDAE Salter, 1864
Subfamily PROETIDELLINAE Hupe, 1953
Kielan (1959 : 69) has declared that the Subfamily Proetidellinae is synonymous
with that of the Proetinae, on the grounds that Hupe's subdivision is unsatisfactorily
diagnosed, utilizing only the degree of divergence of the anterior branches of the
facial suture. The type genera of these two subfamilies differ considerably in several
respects, including the presence in Proetidella of a definite preglabellar field such as is
not seen in Proetus. Accordingly the writer prefers to follow Richter & Struve (in
Moore, 1959 : 0.395) in retaining Proetidellinae as a distinct Subfamily.
Genus PROETIDELLA Bancroft, 1949
TYPE SPECIES. Proetidella feamsidesi by original designation of Bancroft (1949 :
304)-
Proetidella? rnarri sp. nov.
(PL 16, figs. 4, 6, 9 ; PI. 17, figs. 5, 6, 8, 9)
*959a- Proetidella aff. feamsidesi Bancroft : Dean, p. 206.
DIAGNOSIS. Glabella of approximately equal length and breadth, broadest
posteriorly, narrowing frontally to gently rounded frontal lobe. Preglabellar field
long, flat ; anterior border brim-like in form, steeply inclined. Palpebral lobes
narrow, extending backwards almost to occipital furrow.
DESCRIPTION. The cephalon is known only from discrete cranidia and librigenae.
The glabella is roughly as broad as long, gently convex longitudinally and trans-
versely, its line of greatest breadth situated a little way in front of the occipital
furrow. The glabellar outline constricts frontally, and glabellar furrows are totally
absent. The axial furrows are only moderately deep, converging frontally around
the anterolateral angles of the frontal lobe, the anterior margin of which is gently
convex, to become continuous with the preglabellar furrow, which is of similar
depth. The preglabellar field is flat, fairly long (sag), equal to about one-quarter of
the length of the glabella, arched forwards gently, and differentiated by only a poorly-
defined furrow from the anterior border. The last-named structure forms a con-
spicuous, raised brim to the front of the cranidium and is generally steeply inclined
forwards, though the angle may vary according to the state of preservation. The
occipital ring is slightly convex transversely, less so longitudinally, of greatest
length (sag.) medially but shortening a little laterally where it forms a pair of ill-
THE TRILOBITES OF THE CARADOC SERIES 125
defined occipital lobes. The hindmost parts of the fixigenae have not been found
preserved ; frontally they form only a thin, rim-like continuation of the pre-
glabellar field, and medially are not differentiated from the palpebral lobes. The
latter are narrow, declined proximally, and extend from opposite the mid-point of
the glabella almost to the line of the occipital furrow. The librigenae are known
only from dissociated specimens, and the ocular surface of the eye has not been found
preserved. Beneath each eye is a poorly-defined eye-platform from which the
convex upper surface of the librigena declines steeply to a deep, broad, lateral
border furrow ; beyond the latter is a well-developed lateral border which passes
backwards into a librigenal spine of moderate length. The pleuroccipital furrow is
well defined and of moderate depth, intersected by the lateral border furrow with-
out reaching the margin ; the pleuroccipital segment becomes broader (exsag.)
laterally, where it passes without interruption into the librigenal spine. As far as
can be judged from both cranidia and librigenae, the posterior branches of the facial
suture run laterally back from the posterior end of the eyes to cut the cephalic
margin only a short distance outside the axial furrows ; the anterior branches con-
verge from the eyes almost to the axial furrows but then curve anterolaterally to
cut the anterior border longitudinally in-line with the palpebral lobes
The hypostoma and thorax are not known.
One almost complete pygidium attributed to the species has been collected. The
frontal breadth is between two and three times the median length, and the outline is
generally semi-elliptical, though the frontal margin is gently curved back laterally.
The axis is well defined, the sides converging backwards only gently, and does not
quite reach the tip of the pygidium. It is strongly convex transversely and stands
well above the pleural lobes, which are only a little declined to the entire lateral
margins. It is estimated that there are five axial rings, with a short terminal
piece. The most complete specimen has four pairs of pleural furrows, the first pair
slightly more impressed than the rest (PL 17, fig. 6), and an additional specimen,
In. 55880, shows that each of the pleural ribs is divided by a shallow interpleural
furrow into two unequal bands, of which the anterior is the narrower (exsag.). As
far as can be ascertained, both sets of furrows extend to the lateral margin.
The dorsal surface of the glabella, occipital ring and anterior border is covered
with thin, slightly-raised, anastomosing ridges arranged in a Bertillon pattern, but
the furrows of the cranidium, together with the preglabellar field, are smooth.
The librigenae have not been found sufficiently well preserved to ascertain whether
they are so ornamented, but the doublure can be seen to be covered with terrace
lines subparallel to the lateral margin. The test of the pygidium is not adequately
known.
HORIZON AND LOCALITY. Corona Beds belonging to the Lower Longvillian
Substage, Bancroftina typa Zone, at Harthwaite Sike, locality E. 3, east of Dufton
(see Text-fig. 4).
HOLOTYPE. BM. In. 54644 (PI. 16, fig. 4 ; PL 17, fig. 8).
PARATYPES. BM. In. 54645 (PL 17, fig. 5) ; In. 54646 (PL 16, fig. 9) ; In. 54647
(PL 16, fig. 6) ; In. 54648 (PL 17, fig. 9) ; In. 55880 ; In. 55881 ; In. 55882 (PL 17,
fig. 6).
126 THE TRILOBITES OF THE CARADOC SERIES
DISCUSSION. The holotype of Proetidella fearnsidesi has recently been refigured
in the form of a line drawing by Struve (in Moore, 1959 : O. 396, fig. 301, i) but his
illustration is somewhat misleading. The original specimen is preserved as a flat-
tened external mould in shaly mudstone, and the glabella is distorted by crushing.
The species is to be redescribed by the writer in a future paper, but in the meantime
it may be stated that the presence in Struve's illustration of a bluntly-pointed frontal
glabellar lobe, together with a pair of basal glabellar lobes, is the result of crushing.
In addition, the pygidium of the holotype has well-defined pleural furrows extending
to the lateral margins, as well as traces of interpleural furrows.
In Proetidella? marri the attributed pygidium is shorter than that of the Shrop-
shire species, with fewer pleural furrows and axial rings, numbering four and five
respectively compared with five and seven. The narrowing of the frontal glabellar
lobe of the Cross Fell form, together with its more transverse frontal margin, are
features not seen in P. fearnsidesi. The frontal portion of the cranidium is not easy
to compare in the two species owing to differences in the state of preservation, as
already stated, but the preglabellar field of P.? marri is the longer (sag.), and its
anterior border may be both better differentiated and more steeply inclined.
Of other British Ordovician proetids, " Proetus " girvanensis (Nicholson &
Etheridge, 1878 : 169, pi. 12, figs. 7-10) from the Drummuck Group of Girvan has
a shorter (sag.) preglabellar field and a larger glabella with three pairs of definite,
though poorly-impressed, glabellar furrows. The eyes are also smaller and situated
farther forwards. The trilobite described by Reed (1914 : 27, pi. 4, fig. 8) as
Cyphaspis jamesoni and founded on a single imperfect specimen from the Bal-
clatchie Group, has a cephalic form, including the development of the preglabellar
field, not unlike Proetidella, but comparison of this species must await its further
description.
The genus Decoroproetus Pfibyl, 1946, with type-species Proetus decorus Barrande
from the Silurian of Bohemia, was claimed by Pfibyl (1953 : 60) as a synonym of
Proetidella, but is now held to be generically distinct, separable by means of its
well-defined basal glabellar lobes, and by the pygidium, the pleural furrows of which
are better developed, more numerous, and turn backwards more strongly.
In Ogmocnemis from the Ashgill Series of Poland (Kielan, 1959 : 69-71) there are
only traces, not always visible, of basal glabellar lobes, the anterior branches of the
facial suture are less divergent frontally than in Proetidella, and the eyes are some-
what smaller, ending posteriorly a little farther in front of the pleuroccipital furrow.
The reasons for separating the two generically may not be well founded, and
Ogmocnemis may be a subgenus of Proetidella, if not actually synonymous with it.
Whittard (1961 : 186) has laid stress on the importance of the preglabellar field in
proetid classification, and has used Kielan's genus in describing Ogmocnemis calvus
from the Lower Soudleyan of west Shropshire. It is difficult to distinguish the
incomplete type cranidium figured by him from one of Proetidella fearnsidesi, and the
species may range above the Harnagian Stage. Specimens in the Soudleyan of
south Shropshire are almost identical with 0. calvus, but their apparent differences
from P. fearnsidesi may be merely the expression of a different lithology and preserva-
tion.
THE TRILOBITES OF THE CARADOC SERIES 127
Family REMOPLEURIDIDAE Hawle & Corda, 1847
Genus REMOPLEURIDES Portlock, 1843
TYPE SPECIES. Remopleurides colbii Portlock, 1843 by subsequent designation
of Miller (1889 : 565-566).
Trilobite remains broadly referable to this genus have been found at a number of
different horizons and localities in the Cross Fell Inlier. In no instance has the
hypostoma or pygidium be~en recovered, and the cranidia are usually damaged or
distorted. Consequently it has not proved practical to assign specific names to the
specimens or to make a firm comparison with other species, and in the following
account they are arranged according to their geological horizons.
Remopleurides sp.
(PL 18, fig. 12)
I959a- Remopleurides sp., Dean, p. 214.
A small damaged cranidium, lacking the anterior glabellar tongue, is the only
evidence of the genus in the Lower Melmerby Beds of the Alston Road section, where
it was found at locality E. The specimen is too poor for detailed comparison, but
the main portion of the glabella is generally similar to that of Remopleurides biacu-
leatus, described by Tripp (1954 : 664, pi. 2, figs. 1-12) from the Craighead Mudstones
of the Girvan district.
Remopleurides sp.
(PI. 18, figs. 6, 10)
!959^- Remopleurides sp., Dean, p. 214.
A single cranidium has been found in the Upper Melmerby Beds, Upper Long-
villian Substage, at locality H by the Alston Road, near Melmerby. It is too badly
damaged for comparison with other species, but it is interesting to note that the
genus has been recorded by Harper (1956 : 389) from North Wales, in association
with an Upper Longvillian fauna. The Alston Road specimen is of relatively large
size, the basal breadth of the glabella being approximately 15 mm.
Remopleurides sp.
(PI. 18, figs. 7, 8)
I959«. Remopleurid&s sp., Dean, p. 207.
Two small cranidia have been collected from Dufton Shales believed to belong to
the middle portion of the Actonian Stage at locality B. 15 in Swindale Beck, and the
better preserved is figured here. The species is probably new but its description
must await additional material. Particularly conspicuous is the anterior glabellar
tongue, which is quadrate, parallel sided, and steeply turned down frontally.
128 THE TRILOBITES OF THE CARADOC SERIES
Remopleurides sp.
(PI. 18, fig. 5)
19590. Remopleurides aff. burmeisteri Bancroft : Dean, p. 207.
One flattened, incomplete cranidium from the Dufton Shales, Onnian Stage,
Onnia gracilis Zone, at locality A. 12 in Pus Gill provides the only known record of
the genus from the Onnian of the Cross Fell Inlier. Comparison with other species is
difficult owing to the poor state of preservation, but the glabella lacks the glabellar
furrows of Remopleurides burmeisteri, and the two are not now thought to be related.
In south Shropshire Remopleurides appears rarely in the Onnia gracilis Zone and
becomes more abundant in the succeeding 0. superba Zone.
Remopleurides sp.
(PI. 18, figs. 9, n, 13)
19590. Remopleurides sp. ind., Dean, p. 208.
Several fragments of Remopleurides have been collected from a thin band of
sandy limestone within the highest beds of the Dufton Shales, Pusgillian Stage, at
locality B. 25 in Swindale Beck. They comprise only flattened cranidia, with occa-
sional thoracic segments and librigenae, but it seems unlikely that more than one
species is represented.
The cranidia are too distorted for detailed comparison but bear a general resem-
blance to the cranidium of Remopleurides colbii Portlock (1843 : 254, pi. i, figs.
1-6 ; pi. 24, fig. 10) from the Killey Bridge Beds of County Tyrone, Northern
Ireland, a species redescribed by Whittington (1950 : 540, pi. 70, figs, i, 2, 4, 5).
Each of a group of three conjoined thoracic segments (PI. 13, fig. n) from Swindale
Beck has an axial ring with a serrated posterior margin, a feature found not only in
R. colbii but also in the Swedish species R. validus, described by Thorslund (1940 :
132, pi. 7, fig. 4) from the Lower Chasmops Limestone of the Lockne area. Three
librigenae are known from the Pusgillian, one being figured here (PI. 13, fig. 9). The
librigenal spine is situated forwards from the genal angle, from which it is separated
by what Warburg (1925 : 87) called a subgenal notch, a structure she described in
R. lotus var. kullsbergensis (1925 : 84, text-fig. 15). Subgenal notches are present
in certain other species of Remopleurides, but do not occur in R. colbii.
IV. STRATIGRAPHICAL DISTRIBUTION AND RELATIONSHIPS
OF THE TRILOBITES
Lithologically the Caradoc strata of the Cross Fell Inlier form what is essentially
a succession of shales and mudstones, broken only by thin layers of impure, nodular
limestone. Such a succession is in marked contrast with the, probably, shallower-
water deposits of south Shropshire, and their thick development of arenaceous
rocks. As might be expected, such differences in lithology are accompanied by
variations in the composition of the shelly fauna, though it has generally proved
possible to apply the faunal Stages of the type-succession to the Inlier.
THE TRILOBITES OF THE CARADOC SERIES 129
In south Shropshire the local base of the Caradoc Series is usually formed by the
Costonian Stage and falls within the Nemagraptus gracilis Zone, so that it may be
equated with part of a profound and widespread marine transgression. Subsidence
of the sea-floor probably continued through the whole of the time represented by the
succeeding Diplogmptus multidens Zone, believed to comprise the Harnagian and
Soudleyan Stages, and may have been prolonged still further because the earliest-
known Caradoc strata of the Cross Fell Inlier belong to the Lower Longvillian
Substage and probably he unconformably on the Borrowdale Volcanic Series,
though an unfaulted junction of the two has not been found satisfactorily exposed.
The trilobites of the earlier Longvillian rocks in the Knock-Dufton and Roman
Fell districts are neither abundant nor varied, comprising only a local species of
Proetidellal , with occasional Brongniartella and Broeggerolithus, suggesting affinity
with the northern part of the Inlier. The fauna of the Melmerby district is par-
ticularly characterized by an abundance of Trinucleidae, especially Broeggerolithus
nicholsoni (Reed), accompanied by common Kloucekia apiculata (M'Coy), indicating
an intimate connection with the faunas of the Drygill Shales of the northern Lake
District and parts of North Wales, for example the Llanbedrog Mudstones. These
two trilobites occur also in Shropshire, but in much reduced numbers. The genera
Chasmops, Conolichas, Illaenus and Toernquistia at Melmerby suggest a connection
with corresponding Scandinavian and Baltic faunas, whilst Paracybeloides occurs
commonly in Scottish and North American Ordovician rocks, though it is known from
stratigraphically early horizons in the Caradoc of North Wales.
In the Upper Longvillian of the Knock-Dufton district, Brongniartella bisulcata
(M'Coy) is moderately common, as it is in south Shropshire, but its frequent Shrop-
shire associate Kloucekia apiculata has not yet been found. Conversely, Estoniops
alifrons (M'Coy) represents a group of phacopid trilobites which is widespread in
the Baltic region and Scandinavia, as well as throughout much of North Wales,
though unrecorded in Shropshire. Trinucleid trilobites have not been found in the
Upper Longvillian rocks of the main part of the Inlier, and their absence has not
been explained satisfactorily.
The trilobites of the Marshbrookian Stage are numerically abundant, though
lacking in variety, and are generally comparable with those of Shropshire. Brong-
niartella bisulcata is moderately common and trinucleids, Broeggerolithus cf.
transiens (Bancroft), fairly common. Dindymene, a European genus, is rare and has
not yet been found in Shropshire, though it occurs also in the Upper Longvillian near
Dufton, and in North Wales. A notable absentee from this horizon in the Cross
Fell Inlier is Chasmops, a surprising fact in view of its abundance in Shropshire.
Owing to the lack of outcrops of Actonian strata in the Inlier the trilobites of this
Stage are poorly known, but the assemblage of Chasmops, Onnicalymene and Remo-
pleurides indicates connections with both Shropshire and Scandinavian faunas.
In the rocks of the Onnian Stage the resemblance between the fauna of the Duftou
district and south Shropshire is most marked. Onnia gracilis (Bancroft), a species
found also in North Wales, occurs in the Onny Valley and at Pus Gill, accompanied
at both places by Lonchodomas pennatus (La Touche) and Onnicalymene onniensis
(Shirley). In the succeeding zone the well-known Shropshire species Onnia superba
GEOL. 7, 3. 7
130 THE TRILOBITES OF THE CARADOC SERIES
(Bancroft) is represented by a local subspecies 0. superba pusgillensis. An interest-
ing feature of the Onnian fauna at Cross Fell is the appearance of such genera as
Atractopyge, Calyptaulax, Pseudosphaerexochus and Tretaspis, marking an influx
from the Scandinavian area which assumed greater proportions during the suc-
ceeding Pusgillian Stage.
The Pusgillian has at various times been assigned to the Ashgill Series, but is now
regarded as the topmost subdivision of the Caradoc Series. Trinucleid trilobites
are abundant at this horizon, but the genus Onnia, so characteristic of the Onnian
is absent and the family is represented only by Tretaspis. The latter genus is
represented by several species or subspecies, all of which indicate close affinities
with corresponding faunas in southern Norway and Sweden. Most of, if not all,
the other trilobites in the Pusgillian are members of genera which have generally
been thought to be more characteristic of Ashgill strata. The trilobite faunas of the
Ashgill Series exhibit a uniformity and widespread distribution which are unknown
in those of the Caradoc Series, and several genera and species are common to Scot-
land, the Anglo- Welsh area, Scandinavia and eastern Europe. The Onnian and
Pusgillian strata of the Cross Fell Inlier are of particular interest because they
demonstrate that the Ashgill elements invaded the Anglo- Welsh area at a relatively
early date, after which there was a progressive increase in their numbers.
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STUBBLEFIELD, C' J. 1928. A new trilobite, Acidaspis (Pseudomonaspis) magnospina, from
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134 THE TRILOBITES OF THE CARADOC SERIES
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xiv + 1 80 pp. Cambridge.
PLATES
Most of the type and figured specimens are in the collections of the British Museum
(Natural History), London, and bear numbers with the prefix In. Other numbers
with the prefix A. denote specimens in the Sedgwick Museum, Cambridge, whilst
one specimen, GSM 19165 (PI. 12, fig. 2), is in the Geological Survey & Museum,
London. The locality numbers refer to the sketch-maps elsewhere in this paper.
All the photographs are by the writer, the specimens having been whitened with
ammonium chloride before photographing.
PLATE 6
Lonchodomas pennatus (La Touche) . . . p. 78
Dufton Shales, Pusgillian Stage, Swindale Beck, locality 6.25.
FIG. i. Internal mould of cranidium showing upturned frontal spine. In. 49933. X2.
FIG. 3. Internal mould of incomplete cranidium. In. 49934. X2.
FIG. 4. Internal mould of flattened cranidium showing hypostomal pits. In. 49936. X2.
FIG. 5. Internal moulds of two small pygidia, probably immature. In. 49931. X2-75-
Dufton Shales, Onnian Stage, Onnia gracilis Zone, Pus Gill, locality A. 8.
FIG. 9. Internal mould of cranidium. In. 49979. X2-25.
Horizon as for Fig. 9, Pus Gill, locality A. 14.
FIG. 12. Internal mould of pygidium. In. 49926. X2-5.
Lonchodomas swindalensis sp. nov. . . . p. 72
Dufton Shales, Upper Longvillian Substage, Kjaerina typa Zone, Swindale Beck, locality B.6.
FIG. 2. Cranidium with part of test preserved. Holotype. In. 49968. X2-25.
Horizon as for Fig. 2, Swindale Beck, locality 6.5.
FIG. 6. Latex cast from external mould of pygidium. Paratype. In. 49967. X4'5-
FIG. 8. Latex cast from external mould of cranidium. Paratype. In. 49946. X3-
Broeggerolithus nicholsoni (Reed) . . . p. 79
Probably Upper Melmerby Beds, Upper Longvillian Substage, Alston Road, locality A.
FIG. 7. Internal mould of cranidium. In. 52570. X4-
Broeggerolithus melmerbiensis sp. nov. . . . p. 81
Horizon and locality as for Fig. 7.
FIGS. 10,13. Almost complete cranidium with three attached thoracic segments. Holotype.
In. 52558. Xi -75.
Broeggerolithus cf. nicholsoni (Reed) . . . p. 80
Corona Beds, Lower Longvillian Substage, Bancroftina typa Zone, Harthwaite Sike,
locality £.3.
FIG. ii. Latex cast from external mould of fragmentary cephalic fringe. In. 546406.
X2-5.
FIG. 14. Latex cast from external mould of incomplete pygidium. In. 54643. X3-
Bull. EM. (N.H.) Geol. 7, 3
PLATE 6
LONCHODOMAS, BROEGGEROLITHUS
PLATE 7
Broeggerolithus nicholsoni (Reed) . . . p. 79
Lower Melmerby Beds, Lower Longvillian Substage, probably Alston Road, locality J.
FIGS. 1,7. Almost complete cephalon. Paratype. A. 29609. X2-8.
FIG. 2. Internal mould of pygidium. Paratype. A. 29615. X3-
FIG. 4. Internal mould of almost complete exoskeleton. A. 2994&a. xi-8.
FIG. 5. Fragmentary cranidium. Paratype. A. 29611. X2-8.
FIGS. 6, 9. Almost complete cranidium. Lectotype. A. 29613. X2-5.
FIG. 8. External mould of fragmentary cranidium. Paratype. A. 29612. X2-8.
FIG. 10. Almost complete cranidium. Paratype. A. 29610. X2-8.
Probably Upper Melmerby Beds, Alston Road, locality uncertain.
FIG. ii. Cephalon preserved partly as an internal, partly as an external mould. Paratype.
A. 29614. X2-8.
Probably Upper Melmerby Beds, Upper Longvillian Substage, Alston Road, locality A
FIG. 3. External mould of cephalic fragment showing reticulate surface of cheek-lobe.
In. 52567. X4-
FIG. 12. External mould of underside of fringe showing pits and librigenal spine. In. 52576.
X3-5-
Bull. B.M. (N.H.) Geol. 7, 3
PLATE 7
BROEGGEROLITHUS
PLATE 8
Broeggerolithus cf. transiens (Bancroft) . . . p. 82
Dufton Shales, Marshbrookian Stage, probably topmost third, Swindale Beck, locality B.i6.
FIG. i. Internal mould of cranidium. In. 50216. X2-2.
FIG. 6. Internal mould of underside of fringe. In. 50215. X2.
Horizon as for Fig. i. Swindale Beck, locality B. n.
FIG. 3. Internal mould of fragmentary cephalon showing genal angle. In. 50244. X2-y5.
Horizon as for Fig. i. Swindale Beck, locality B. 14.
FIG. 4. Internal mould of slightly distorted pygidium. In. 49957. X2-5.
FIG. 8. Internal mould of fragmentary cranidium showing development of I3. In. 50190.
X2.
Horizon as for Fig. i. Harthwaite Sike, locality E. 8.
FIG. ii. Internal mould of small cranidium. In. 51853. X3'5-
Broeggerolithus sp. . . . .. . p. 83
Horizon and locality as for Fig. 4.
FIG. 2. Latex cast from external mould of fragmentary cranidium. In. 50222. X2.
Onnia gracilis (Bancroft) . . . . p. 84
Dufton Shales, Onnian Stage, Pus Gill, locality A. 4.
FIG. 12. Internal mould of cranidium showing pitting of fringe. In. 50001. X3-
FIG. 13. Damaged, almost complete specimen showing pygidium and impression of under-
side of fringe. In. 49948. X2-5.
Onnia superba (Bancroft) pusgillensis Dean . . . p. 84
Dufton Shales, Onnian Stage, Onnia superba Zone, Pus Gill, locality A. 5.
FIG. 5. Internal mould of incomplete cranidium. Paratype. In. 50008. xi-5-
FIG 7. Fragmentary cranidium, an internal mould, accompanied by external mould of
Tretaspis. In. 50003. xi*5-
FIG. 9. Latex cast from external mould of two incomplete cranidia. Paratype. In. 50005.
X2.
FIG. 10. Internal mould of pygidium. In. 50013. X2-25.
Bull. B.M. (N.H.) Geol. 1, 3
PLATE 8
12
BROEGGEROLITHUS, ONNIA
PLATE 9
Tretaspis kiaeri (St0rmer) duflonensis Dean . . p. 86
Dufton Shales, Pusgillian Stage, Pus Gill, locality A. 27.
FIG. i. Internal mould of almost complete individual showing median and lateral ocelli,
thorax and pygidium. Holotype. In. 50020^. X3-
FIGS. 5-7. Latex cast from external mould of same specimen, showing surface of thorax
and pygidium, reticulation of glabella, and fringe profile. In. 500206. Figs. 5, 7, X4- Fig. 6,
X3-
Tretaspis kiaeri (St0rmer) radialis Lamont . . . p. 86
Horizon as for Fig. i. Swindale Beck, locality B. 25.
FIGS. 2, 3. Latex cast from external mould of cephalon, illustrating pitting of fringe,
marginal suture, and grooved librigenal spines. In. 500456. X2-5.
FIG. 4. Underside of fringe preserved as internal mould showing development of girder.
In. 50052. X2'5.
Bull. EM. (N.H.J Geol. 7, 3
PLATE 9
TRETASPIS
PLATE 10
Tretaspis convergens Dean . . . . p. 85
Dufton Shales, Pusgillian Stage, Swindale Beck, locality 6.25.
FIG. i. Latex cast from external mould showing reticulation of cephalon. Holotype
In. 500596. X3-
FIGS. 3, 5. Internal mould of same specimen. In. 50059*. X2-5-
FIG. 2. Internal mould of part of cephalic fringe. Paratype. In. 50037.
Tretaspis cf. ceriodes (Angelin) donsi St0rmer . . p. 85
Dufton Shales, Onnian Stage, Onnia superba Zone, Pus Gill, locality A. 5.
FIGS. 4, 8. Latex cast from external mould of cephalon to show fringe and surface of glabella
and cheek-lobes. In. 500186. X3-
Dufton Shales, Onnian Stage, Onnia gracilis Zone, Pus Gill, locality A. 13.
FIG. 6. Latex cast from external mould of fragment of fringe. In. 50096. X3'5.
Pseudosphaerexochus cf. octolobatus (M'Coy) . . . p. 87
Horizon as for Fig. i, Pus Gill, locality A. 6.
FIG. 7. Internal mould of incomplete pygidium. In. 49816. X2-25-
Horizon and locality as for Fig. i.
FIGS. 9,10. Internal mould of cranidium. In. 49803. X3-25.
Horizon as for Fig. i, Pus Gill, locality A. 22.
FIG. ii. Internal mould of incomplete cranidium. In. 4981 ja. xi'75-
FIG. 12. Latex cast from external mould of same specimen. In. 498176. X2-5.
Bull. B.M. (N.H.) Geol. 7, 3
PLATE 10
TRETASPIS, PSEUDOSPHAEREXOCHUS
GEOL. 7, 3.
PLATE ii
Encrinurus sp. . . . . . p. 88
Upper Melmerby Beds, Upper Longvillian Substage, Bancroftina robusta Zone,
Alston Road, locality H.
FIGS, i, 4. Internal mould of pygidium. In. 49820. X5-
Atractopyge scabra sp. nov. . . . . . p. 91
Dufton Shales, Onnian Stage, Onnia, superba Zone, Pus Gill, locality A. 5.
FIG. 2. Internal mould of cranidium. Holotype. In. 50154. X3-
Dufton Shales, Pusgillian Stage, Swindale Beck, locality B. 25.
Fig. 3. Internal mould of cranidium, slightly compressed laterally. In. 50150. X3-
Horizon as for Fig. 3, Pus Gill, locality A.25.
FIG. 10. External mould of pygidium. In. 501576. X4-
Atractopyge ? sp. . . . . . p. 93
Corona Beds, Lower Longillian Substage, Bancroftina type Zone, Harthwaite Sike,
locality £.3.
FIG. 5. Latex cast from external mould of fragmentary cranidium. In. 546386. X5-
Dindymene sp. . . . . . p. 90
Dufton Shales, Upper Longvillian Substage, Kjaerina typa Zone, Swindale Beck,
locality B. 5.
FIG. 6. Incomplete cranidium with test partially preserved. In. 49966. x6.
Paracybeloides cf. girvanensis (Reed) . . . p. 94
Horizon and locality as for Fig. 3.
FIG. 7. Internal mould of pygidium. In. 50177. X3-
FIG. 9. Similar specimen showing development of spines near tip of pygidium. In. 50179.
X3-
FIG. 12. Latex cast from external mould of cranidium. In. 50158. X2-25-
FIG. 14. Internal mould of incomplete librigena. In. 50172. X2-5.
FIG. 15. Internal mould of cranidium. In. 50165. X3-
Atractopyge ? sp. . . . . . . p- 93
Lower Melmerby Beds, Lower Longvillian Substage, Alston Road, locality J.
FIG. 8. Latex cast from exteral mould of cranidium. In. 546546. x 4.
Paracybeloides sp. . . . . p. 95
Horizon and locality as for Fig. 8.
FIG. ii. Latex cast from external mould of pygidium. In. 54657. x6.
Dindymene duftonensis sp. nov. . . . . p. 89
Dufton Shales, Marshbrookian Stage, probably topmost portion, Harthwaite Sike, locality E.I2.
FIG. 13. Latex cast from external mould of incomplete cephalon. Holotype. In. 54652.
X6.
Bull. BM. (N.H.) Geol. 7, 3
PLATE 11
ATRACTOPYGE, DINDYMENE, ENCRINURUS, PARACYBELOIDES
PLATE 12
Dalmanitina mucronata matutina subsp. nov. . . p. 95
Dufton Shales, Pusgillian Stage, Pus Gill, locality A. 7.
FIG. i. Internal mould of incomplete cephalon. Paratype. In. 501 120,. X2-J5.
Horizon as for Fig. i, Swindale Beck, locality 6.25.
FIG. 5. Internal mould of incomplete cranidium showing fixigenal spine. Paratype.
In. 50113. X2-75.
FIG. 6. Internal mould of pygidium. Holotype. In. 49905. X2-75.
FIG. 9. Latex cast from external mould of fragmentary pygidium. Paratype. In. 499*9-
X2-25.
FIG. 12. Internal mould of incomplete cranidium. Paratype. In. 501 140. X2-25.
Estoniops alifrons (M'Coy) . . . . . p. 100
Caradoc Series, probably Upper Longvillian Substage, near Gelli Grin, Bala District.
FIG. 2. Internal mould of cephalon. GSM 19165. X2'5.
Horizon as for Fig. 2, Capel Gannon, Denbighshire.
FIG. 4. Incomplete cephalon with part of test preserved. Lectotype, figured by M'Coy
in Sedgwick & M'Coy, 1851, pi. i.G, fig. 12. Sedgwick Museum A. 42694. x 1-5.
Dufton Shales, Upper Longvillian Substage, Kjaerina typa Zone, Swindale Beck, locality B. 7.
FIGS. 7,13,14. Cephalon with test preserved. In. 49837. X3-
Horizon as for Fig. 7, Swindale Beck, locality B. 10.
FIGS. 8,10. Pygidium with test preserved. In. 49839. X2-5.
Duftonia lacunosa Dean . . . . p. 97
Horizon and locality as for Fig. 5.
FIG. 3. Latex cast from external mould of cranidium. Holotype. In. 49824. X3-
Kloucekia (Phacopidina) apiculata (M'Coy) . . p. 97
Upper Melmerby Beds, Upper Longvillian Substage, Bancroftina robusta Zone, Alston Road,
locality H.
FIG. n. Latex cast from external mould of fragmentary cranidium. In. 52587. X3-
Bull B.M. (N.H.) Geol. 7,3
PLATE 12
14
DALMANITINA, DUFTONIA, ESTONIOPS, KLOUCEKIA
PLATE 13
Calyptaulax planiformis sp. nov. . . . p. 98
Dufton Shales, Pusgillian Stage, Swindale Beck, locality B. 25.
FIG. i. Latex cast from external mould of incomplete cranidium. Paratype. In. 499656.
X2-75.
FIG. 2. Internal mould of distorted cranidium. Paratype. In. 49908. X2.
FIG. 4. Internal mould of pygidium. Holotype. In. 50138. X2.
FIG. 5. Internal mould of incomplete pygidium. Paratype. In. 49907. X3-
Dufton Shales, Onnian Stage, Onnia superba Zone, Pus Gill, locality A. 5.
FIG. 3. Internal mould of cranidium. Paratype. In. 49903. Xi'75-
Chasmops aff. maxima (Schmidt) . . . p. 105
Horizon as for Fig. i, Pus Gill, locality A. 28.
FIG. 6. Internal mould of fragmentary cranidium. In. 50412. X3-
Onnicalymene onniensis (Shirley) . . . . p. 115
Horizon and locality as for Fig. 3.
FIG. 7. Cranidium with part of test preserved. In. 50294. x 3.
Chasmops cf. extensa (Boeck) . . . . p. 104
Dufton Shales, Actonian Stage, Swindale Beck, locality B. 15.
FIG. 8. Internal mould of pygidium. In. 498970. X2-75.
Gravicalymene jugifera sp. nov. . . . p. 116
Horizon as for Fig. i, Swindale Beck, locality B. 30.
FIGS. 9, ii. Internal mould of almost complete individual, slightly compressed laterally.
Paratype. In. 50245. Xi'5-
Flexicalymene cf. caractaci (Salter) . . . p. 114
Lower Melmerby Beds, Lower Longvillian Substage, Alston Road, locality J.
FIG. 10. Internal mould of incomplete cranidium. In. 54658. X2.
Chasmops sp. . . . . . p. 105
Horizon and locality as for Fig. 10.
FIG. 12. Latex cast of incomplete pygidium. In. 546566. X2.
Diacalymene cf. marginata Shirley . . . . p. 116
Horizon and locality as for Fig. i.
FIG. 13. Internal mould of incomplete cranidium, the frontal portion slightly compressed
longitudinally. In. 50256. X4'5-
Bull. B.M. (N.H.) Geol. 7, 3
PLATE 13
L '
CALYPTAULAX, CHASMOPS, DIACALYMENE, FLEXICALYMENE,
GRAVICALYMENE, ONN1CALYMENE
PLATE 14
Onnicalymene onniensis (Shirley) . . . p. 115
Dufton Shales, Pusgillian Stage, Swindale Beck, locality B. 25.
FIG. i. Internal mould of pygidium with attached thoracic segments. In. 50270. X2-25.
FIGS. 2, 10. Internal mould of almost complete individual. In. 50266. xi-75-
Horizon as for Fig. i, Pus Gill, locality A. 28.
FIG. 7. Internal mould of large, incomplete cranidium. In. 50318*. X2-75.
Gravicalymene jugifera sp. nov. . . . p. 116
Horizon as for Fig. i, Swindale Beck, locality B. 28.
FIGS. 3,4,8. Almost complete cranidium with part of test intact. Holotype. In. 50263.
Xi-75-
Horizon as for Fig. i, Swindale Beck, locality B. 30.
FIG. 9. Internal mould showing pygidium and thorax. Paratype. In. 50245. Xi'5-
Onnicalymene laticeps (Bancroft) . . . p. 115
Dufton Shales, Actonian Stage, Swindale Beck, locality B. 15.
FIGS. 5, 6. Internal mould of cephalon with two attached thoracic segments. In. 50258.
X3-
Diacalymene cf. marginata Shirley . . . p. 116
Horizon and locality as for Fig. I.
FIG. ii. Latex cast from external mould showing form of librigena, with conjugate fixigena
and second glabellar lobe. In. 50254. X 1-75.
Bull. B.M. (N.H.) Geol. 7, 3
PLATE 14
II
DIACALYMENE, GRAVICALYMENE, ONNICALYMENE
GEOL. 7, 3.
PLATE 15
Brongniartella ascripta (Reed) . . . . p. 106
Probably Lower Melmerby Beds, Lower Longvillian Substage, Alston Road, locality J.
FIG. i. Incomplete, damaged cranidium. A. 29633. X2.
FIG. 2. Internal mould of small cranidium. A. 29635. X3- Figd. Reed, 1910, pi. 17,
figs. 5, 7.
FIG. 5. Incomplete cranidium showing form of glabella. A. 29634. X3- Figd. Reed,
1910, pi. 17, fig. 6.
FIG. 8. Internal mould of hypostoma. A. 29636^. x6. Figd. Reed, 1910, pi. 17, fig. 8.
FIG. 11. Internal mould of incomplete cranidium showing median ridge. Holotype.
A. 296320. X2'5. Figd. Reed, 1910, pi. 17, fig. 4.
Brongniartella minor (Salter) . . . p. 106
Corona Beds, Lower Longvillian Substage, Bancroftina typa Zone, Pus Gill, locality A. 3.
FIG. 4. Internal mould of pygidium. In. 49844. x 2.
Brongniartella sp. . . . . . p. 108
Corona Beds, probably Lower Longvillian Substage, western flank of Roman Fell,
exact locality unknown.
FIG. 3. Incomplete pygidium with part of thoracic axis. A. 32909. Xi-5-
FIG. 6. Slightly distorted cranidium. A. 32910. xi-5-
FIG. 7. Large pygidium with furrows exaggerated by crushing. A. 32911. xi'4-
FIG. 9. Fragmentary pygidium with part of thorax. A. 32907. x 1-5.
FIG. 10. Incomplete pygidium. A. 32908. xi-8.
Bull. B.M. (N.H.) Geol. 7, 3
PLATE 15
BRONGNIARTELLA
PLATE 16
Brongniartella depressa sp. nov. . . . p. 108
Dufton Shales, Pusgillian Stage, Swindale Beck, locality B. 25.
FIG. i. External mould of cranidium. Holotype. In. 49882. X2.
FIG. 7. Latex cast from external mould of incomplete thorax and pygidium. Paratype.
In. 49881. X2-5.
FIG. 10. External mould of almost complete pygidium. Paratype. In. 49884. X2-5.
FIG. 13. Internal mould of immature cranidium showing traces of glabellar furrows.
Paratype. In. 49885. X2.
Toernquistia aff. reedi Thorslund . . p. 118
Lower Melmerby Beds, Lower Longvillian Substage, Alston Road, probably locality J.
FIGS. 2, 3. Internal mould of incomplete cranidium. A. 29960. x8*5.
Proetidella ? marri sp. nov. . . . . . p. 124
Corona Beds, Lower Longvillian Substage, zone of Bancroftina typa, Harthwaite Sike,
locality £.3.
FIG. 4. Internal mould of cranidium. Holotype. In. 54644. X2-5.
FIG. 6. Internal mould of cranidium. Paratype. In. 54647. X4-
FIG. 9. Latex cast of incomplete cranidium showing raised lines ornamenting surface of
test. Paratype. In. 54646. X2-5.
Brongniartella aff. platynota (Dalman) . . . . p. no
Dufton Shales, Pusgillian Stage, Swindale Beck, locality B. 30.
FIG 5. Internal mould of incomplete pygidium. In. 49878. X2-5.
Brongniartella bisulcata (M'Coy) . . . . p. 108
Dufton Shales, Marshbrookian Stage, probably uppermost third, Swindale Beck,
locality B. 14.
FIG. 8. Incomplete, slightly distorted cranidium. In. 49847. xi'4-
Brongniartella minor (Salter) . . . p. 106
Horizon and locality as for Fig. 4.
FIG. n. Internal mould of cranidium. In. 49840. X2-2.
Stenopareia ? sp. . . . . . p. 120
Horizon and locality as for Fig. 2.
FIG. 12. Internal mould of pygidium. A. 29966. xi'4-
Brongniartella ascripta (Reed) ? .... p. 107
Horizon and locality as for Fig. 4.
FIG. 14. Fragmentary internal mould of cranidium. 1^54639(1. X2-5.
Bull. B.M. (N.H.) Geol. 7, 3
PLATE 16
12
13
14
T A TT T.AF.MT IS. PROFTinFT.T.A ?. TOERNOUISTIA
PLATE 17
Platylichas cf. laxatus (M'Coy) . . . p. 121
Dufton Shales, Pusgillian Stage, Swindale Beck, locality B. 25.
FIG. i. Internal mould of cranidium. In. 50115. X2-5.
FIG. 7. Internal mould of pygidium. In. 50117. X2.
Platylichas sp. . . . . . p. 122
Upper Melmerby Beds, Upper Longvillian Substage, Kjaerina bipartita Zone, Alston Road,
locality H.
FIG. 2. Internal mould of fragmentary pygidium. In. 52595. X2-5.
Primaspis semievoluta (Reed) . . . . p. 122
Lower Melmerby Beds, Lower Longvillian Substage, Alston Road, probably locality J.
FIGS. 3, 13. Internal mould of incomplete cranidium. Lectotype. A. 29951. X3'5-
FIG. 10. Internal mould of small cranidium. Paratype. A. 29952. X4-
FIG. ii. Internal mould of incomplete librigena. Paratype. A. 29954. X3-
FIG. 15. Incomplete pygidium. Paratype. A. 29953. X3'4-
Proetidella ? marri sp. nov. . . . . p. 124
Corona Beds, Lower Longvillian Substage, zone of Bancroftina typa, Harthwaite Sike,
locality £.3.
FIG. 5. Latex cast from external mould of cranidium. Paratype. In. 55882. X2-5.
FIG. 6. Internal mould of pygidium. Paratype. In. 55882. x 3.
FIG. 8. Internal mould of cranidium. Holotype. In. 54644. X2-5.
FIG. 9. Internal mould of librigena. Paratype. In. 54648. X2-5.
Otarion sp. . . . . . p. 123
Horizon and locality as for Fig. 2.
FIG. 4. Internal mould of cranidium. In. 498 iga. xi2.
FIG. 12. Latex cast from external mould of same specimen. In. 498196. x 12.
Conolichas melmerbiensis (Reed) . . . p. 120
Horizon and locality as for Fig. 3.
FIG. 14. Internal mould of hypostoma. Paratype. A. 29643. X3-
FIG. 16. Internal mould of incomplete cranidium. Paratype. A. 29641. Xi-6.
Bull. B.M. (N.H.) Geol 7, 3
PLATE 17
16
CONOLICHAS. OTARION. PLATYLICHAS, PRIMASPIS, PROETIDELLA ?
PLATE 18
Conolichas melmerbiensis (Reed) . . . . p. 120
Lower Melmerby Beds, Lower Longvillian Substage, Alston Road, probably locality J.
FIG. i. Internal mould of almost complete individual. Paratype. A. 29637. xi'7-
FIG. 2. Internal mould of pygidium. Paratype. A. 29642^. xi-y.
FIGS. 3,4. Internal mould of almost complete individual. Lectotype. A. 29638. xi-j.
Remopleurides sp. . . . . p. 128
Dufton Shales, Onnian Stage, Onnia gracilis Zone, Pus Gill, locality A. 4.
FIG. 5. Internal mould of incomplete cranidium. In. 5Oi2ga. X2-8.
Remopleurides sp. ..... p. 127
Upper Melmerby Beds, Upper Longvillian Substage, Kjaerina bipartita Zone, Alston Road,
locality H.
FIGS. 6,10. Internal mould of distorted cranidium. In. 52594. X2-4.
Remopleurides sp. . . . . p. 127
Dufton Shales, Actonian Stage, Swindale Beck, locality B. 15.
FIGS. 7,8. Internal mould of cranidium. In. 50131. X4'5-
Remopleurides sp. . . . . . p. 128
Dufton Shales, Pusgillian Stage, Swindale Beck, locality B. 25.
FIG. 9. Underside of internal mould of left librigena. In. 50140. X2-6.
FIG. ii. Internal mould of three conjoined thoracic segments. In. 501360. X3-
FIG. 13. Internal mould of flattened cranidium. In. 50135. X2-2.
Remopleurides sp. . . . . . p. 127
Lower Melmerby Beds, Lower Longvillian Substage, Alston Road, locality E.
FIG. 12. Small, incomplete cranidium with most of test preserved. 1^52999. X5-
^ ,-
Bull. B.M. (N.H.) Geol. 7, 3
PLATE 18
CONOLICHAS, REMOPLEURIDES
PRINTED IN GREAT BRITAIN BY
ADLARD AND SON, LIMITED
BARTHOLOMEW PRESS, DORKING
FOSSIL FLORA OF THE
DRYBROOK SANDSTONE
IN THE FOREST OF DEAN,
GLOUCESTERSHIRE
K. M. LELE
AND
J. WALTON
BULLETIN OF
THE BRITISH MUSEUM (NATURAL HISTORY)
GEOLOGY Vol. 7 No. 4
LONDON: 1962
FOSSIL FLORA OF THE DRYBROOK
SANDSTONE IN THE FOREST OF DEAN,
GLOUCESTERSHIRE
BY
K. M. LELE
(Birbal Sahni Institute, Lucknow)
AND
J. WALTON
(Professor of Botany in the University of Glasgow)
Pp. 135-152 ; Plates 19-23
BULLETIN OF
THE BRITISH MUSEUM (NATURAL HISTORY)
GEOLOGY Vol. 7 No. 4
LONDON: 1962
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. 7, No. 4 of the Geological
(Palaeontological) series.
Trustees of the British Museum 1962
PRINTED BY ORDER OF THE TRUSTEES OF
THE BRITISH MUSEUM
Issued August 1962 Price Twelve shillings and sixpence
FOSSIL FLORA OF THE DRYBROOK
SANDSTONE IN THE FOREST OF DEAN,
GLOUCESTERSHIRE
By K. M. LELE & J. WALTON
SYNOPSIS
Descriptions of the fossil plants including sporae dispersae found in the Drybrook Sandstone
(Mississippian) at Hazel Hill in the Forest of Dean, Gloucestershire are given. The most
frequently occurring species are Lepidophyllum fimbriatum Kidston, Scutellocladus variabilis
gen. et sp. nov. and Diplopteridium holdeni sp. nov. The spores belong to nineteen genera
including three new species. A close comparison can be made between this flora and the flora
of the Lower Brown Limestones at Diserth in North Wales and the age, judging from the plants,
is that of the Calciferous Sandstone of Scotland. Palaeozoological evidence however supports a
considerably later age.
INTRODUCTION
THE fossil plants described below were collected by Dr. H. S. Holden, Mr. F. M.
Wonnacott, Mr. T. D. West, Mrs. C. J. Croucher and Mr. Keith Allen. Most of the
specimens described are preserved in the British Museum (Nos. ¥.42428-43004,
¥.42048, ¥.42049), a few are in the Geological Museum, University of Birmingham
(B.U. 800-804) while one is in the Hunterian Museum, University of Glasgow (No.
Pb.3452). Preparations of the sporae dispersae isolated from the sandstone are now
in the British Museum (Nos. ¥.43700-43734). In addition we have been kindly
allowed to examine a collection made by members of the staff of the Geological
Department, Kingston Technical College.
The specimens were all collected from a bed of fine sandstone with a grey to pink
coloration in the Hazel Hill Quarry (Grid reference 32/646184) near Puddlebrook in
Gloucestershire. The organic matter of the original plants is in the form of a black
powder and only in exceptional cases (a few lepidophytes) is a cuticular membrane
preserved. The forms of the plants are retained as a fine impression on the sand-
stone which is stained a brownish colour where it was in contact with the plant
material. This helps to show up the shapes of the plants in photographs taken on
orthochromatic plates.
The rock also contains somewhat poorly preserved spores. The sporae dispersae
were isolated by macerating small fragments of the plant-bearing rock in cold
concentrated hydrofluoric acid for about a week. The residue was washed free of
the acid and the organic matter containing the microfossils was further separated
by the swirling method (Funkhouser & Evitt, 1959 : 373). The material thus
concentrated was directly mounted in glycerine jelly. Canada balsam and chloral
hydrate were also found suitable for mounting. In favourable cases, grains were
individually picked out and mounted for study. Staining with Safranin was found
useful especially in case of lightly coloured spores. In view of the rather sparse
GEOL. 7, 4. 10
138 FOSSIL FLORA OF THE DRYBROOK SANDSTONE
spore population several macerations were made and nearly fifty slides were examined
for the identification of spores. The number of better preserved and identifiable
specimens is given in the description of each species. The system of classification
used here is that of Potonie & Kremp (1955).
One of the most frequently occurring fossils, a lepidophyte sporophyll, Lepido-
phyllum fimbriatum Kidston has been described and figured by Allen (1961) and
others under the name of Lepidostrobophyllum fimbriatum (Kidst.) but so far the
generic name Lepidostrobophyllum has no valid basis and it is hoped that this
nomenclatural confusion may be cleared up. The only animal fossil found associated
with the plants is a Conulariid organism almost certainly of marine origin, which
suggests that the sediments containing the plants were deposited near the sea in a
lagoon or estuary.
In the earliest account of the geology of this area and on the one-inch, Old Series,
sheets of the Geological Survey (43 S.E., 43 S.W. and 35) of the Forest of Dean, the
Drybrook Sandstone is stated to be of Millstone Grit age. In the Geological Survey
Memoir Trotter (1942 : 15, 16) divides the Drybrook Sandstone into upper and lower
parts separated by the Drybrook Limestone or its equivalent. In the north the
Coal Measures overlap, unconformably, the Lower Drybrook Sandstone. At Hazel
Hill the sandstone rests on Limestones referred to zones CZS1 of the Avonian zone
sequence and is overlain unconformably by the Coal Measures (Sibly & Reynolds,
1937 : 25, 37). Crookall (1939 : 72) assigns some obscure plant remains in the Dry-
brook Sandstone to ICydostigma, a genus of Devonian plants, but they might well
be the remains of a Carboniferous lycopod.
In the account that follows, descriptions are given of the most complete and
identifiable specimens.
SYSTEMATIC DESCRIPTIONS
Genus SCUTELLOCLADUS nov.
DIAGNOSIS. Stems bearing narrow lanceolate, acuminate, falcate leaves.
Leaves spirally arranged, almost contiguous, rhomboidal in section near their
bases. On the smallest branches the leaves are more widely separated. On small
branches which have lost their leaves the leaf scars are elliptical and level with the
surface of the stem. A punctiform cicatrule is present in the centre of the scar.
There are no persistent leaf bases or cushions and no evidence of a ligule or parichnos.
TYPE SPECIES. Scutellocladus variabilis sp. nov.
Scutellocladus variabilis sp. nov.
(PI. 19, figs. 1-6)
DIAGNOSIS. As for the genus.
SYNTYPES. B.M.N.H. Nos. ¥.42433, ¥.42477, ¥.42538, ¥.42558.
LOCALITY. Hazel Hill Quarry, Puddlebrook, Forest of Dean, Gloucestershire.
HORIZON. Drybrook Sandstone. Mississippian.
DESCRIPTION. There is a number of examples of small branches of a plant with
the appearance of a lycopod. Their stems vary in thickness from i mm. to 8 mm.,
FOSSIL FLORA OF THE DRYBROOK SANDSTONE 139
some having their leaves still attached (PI. 19, figs, i, 2, 3, 5, 6) and some from which
the leaves have fallen (PI. 19, fig. 4). Where branching is seen it is dichotomous.
The leaves vary from about 9 mm. to 5 mm. in length and from 2 mm. to 0-5 mm.
in breadth. The leaves were acuminate, falcate and rhomboidal in section judging
from the shape of their area of attachment to the stem (PI. 19, fig. la). They were
closely set on the stem in a spiral phyllotaxy. On one branch (PI. 19, fig. 2), which
is presumably at a later stage of development, the bases of the leaves which are
still attached exhibit a narrow central vertical groove, are more widely spaced and
are subcircular to elliptical in shape. The groove no doubt represents the vascular
strand in the leaf base. On the smaller branches the leaves are attached by bases
more elongated vertically and the scars are vertically elongated ellipses (1-8 x 1-5
mm.) with a central punctiform cicatrule (PI. 19, fig. 4). The scar is flush with the
surface of the branch and there is no elevated leaf base or cushion. This is evident
when the profile of the branch is studied where it is seen to be almost straight. The
scars measure about I mm. long by 0-7 mm. wide on the very smallest branches
(PI. 19, fig. 6).
There is no evidence of a ligule or parichnos on the leaves or on the defoliated
branches. The complete absence of a persistent leaf cushion is a marked character-
istic and the base of the leaf is not decurrent. There are some lycopod-like stems
described by Radchenko (1956 : 199, pi. 36, figs, i, 2 ; 1957 : 50, pi. 3, figs. 2, 3)
under the name Tomiodendron ostrogianum (Zal.) which at first sight seem to re-
semble our fossils : but on closer examination it is clear that they had projecting
persistent bases, in many cases decurrent, with a definite leaf-scar at the top of the
cushion.
In addition to the compressions of small leafy branches assigned to Scutellocladus
variabilis there are several examples (B.M.N.H., Nos. ¥.42459, ¥.42551, ¥.42557,
¥.42559) in which nothing but the cuticle remains in the form of a perforated lamina
similar to the cuticles of Bothrodendron tenerrimum from the Moscow basin but
lacking any trace of the cuticular linings of the ligular pits characteristic of that
species. These Hazel Hill cuticles are from branches from about 8 mm. to over 13
mm. in diameter and may belong to S. variabilis.
Genus LEPIDOPHYLLUM Brongniart
Lepidophyllum cf. fimbriatum Kidston
(PI. 20, figs. 18-20)
In addition to typical examples of L. fimbriatum (Allen, 1961 : 225) there is a
number of lepidophyte leaves which are much longer, some reaching more than 6-5
cm. and about 6 mm. at the broadest part of the base (PI. 20, fig. 20). The leaves
taper very gradually to an acute apex. A single vein is visible in the middle and,
while several have a smooth margin, two show rather widely spaced lateral hairs
(PI. 20, fig. 18) smaller than those of L. fimbriatum but of the same kind. If a
sporangium had been present it must have been less than 6 mm. in width. A slight
transverse cleft can be seen in one or two (PI. 20, fig. 19) which might indicate the
140 FOSSIL FLORA OF THE DRYBROOK SANDSTONE
presence of a ligule. There is no film of carbonaceous matter at their proximal end
as dense as that found on L. fimbriatum as described by Kidston (1883 : 543).
One can merely speculate as to the nature of these leaves. Perhaps they were
the foliage leaves of the plant, either microsporophylls, sterile bracts of a strobilus,
or intermediates between sporophylls and leaves.
There are some indubitable specimens of L. fimbriatum which measure only 8 mm.
at the widest point and in the region where the sporangium was probably present
are only 5 mm. wide.
Cf. Stigmaria
(PI. 19, fig. 7)
Two or three impressions of which the largest is shown in PI. 19, fig. 7 are in the
collection. They exhibit an extended smooth surface with one or more circular
scars ranging from about 10 mm. to 12 mm. in diameter, each with a small central
elevation. There scars closely resemble those on a Stigmaria. A large part of the
surface shown uppermost in the figure has no such scars. In the Manchester Museum
there is a similar specimen (Manch. Mus. CWM.23) which consists of a smooth
surface, about 5x3 cm., on which there are two scars of the same kind. This
specimen is from the Lower Brown Limestone at Diserth in North Wales. Another
specimen from the same locality consists of a lepidophyte axis (Manch. Mus. LL.ngo)
28 cm. long which tapers from 9-5 cm. in width at the base to 4 cm. at the top ;
on which at the lower end there are similar large stigmarioid scars and on its upper
two-thirds numerous regularly arranged small indentations or scars which are
obviously the marks of leaf traces. As no leaf traces can be seen on the surface
round the large scars it is unlikely that the large scars represent branch scars.
These specimens are possibly parts of the base of a lepidophyte of comparable
size to the " Naples tree " Lepidosigillaria whitei (Krausel & Weyland, 1949 : 148)
which has a slightly bulbous base with stigmarioid scars to which rooting appendages
were attached.
Genus DIPLOPTERIDIUM Walton
Diplopteridium holdeni sp. nov.
(PI. 20, figS. 8-15)
DIAGNOSIS. Fronds tripinnatifid, from 10 cm. to 20 cm. long, ultimate divisions
about 0-5 mm. in width. Main rachis from 2 mm. to 3-5 mm. broad at base ;
forking equally into two main divisions. Pinnae borne on the two divisions of the
frond and on the rachis below the fork. Some fronds have a naked rachis in the
angle of the fork. Its first dichotomy is at right angles to the plane of the frond.
HOLOTYPE. B.M., ¥.42453.
PARATYPES. B.M., ¥.42474, ¥.42487, ¥.43004.
LOCALITY. Hazel Hill Quarry, Puddlebrook, Forest of Dean, Gloucestershire,
England.
HORIZON. Drybrook Sandstone ; Mississippian.
FOSSIL FLORA OF THE DRYBROOK SANDSTONE 141
DESCRIPTION. The most noticeable and one of the most frequently occurring
species found in the Hazel Hill quarry beds is a tripinnatifid frond which bears a
close similarity to Diplopteridium teilianum (Kidston) which was found in beds
assigned to the Upper Black Limestone near Prestatyn in North Wales. The fronds
are, however, much smaller (PI. 20, figs. 8-15) and the ultimate divisions of the
pinnules narrower than those of D. teilianum. As far as one can judge from the
fragments the fronds were probably from 10 cm. to 20 cm. in length and the main
rachis, which in the specimens examined ranges from 2 mm. to 3-5 mm. in width,
forked into two equal divisions in the middle of the frond. Near the proximal end
of the frond (PI. 20, fig. 9) the pinnae are small with few divisions and are opposite
but higher up they increase in size and complexity and are alternate. They appear
in many instances to have had their principal plane facing the rachis (PI. 20, figs.
8, 10) so that as a result of compression the pinna at first sight resembles a Rhacop-
teris pinnule of the R. petiolata type and Rhacopteris geikiei Kidston (19230 : 218,
text-fig, n). The main rachis in one specimen (PI. 20, fig. 9) appears to have been
rugose. The arrangement of the pinnae in relation to the forking of the main
rachis and their degree of subdivision is similar to that found in D. teilianum.
Towards the ends of the two main divisions of the frond the pinnae become smaller
and simpler and in the entire apex seen in PI. 20, fig. n the rachis terminates in a
single spatulate division.
Evidence for the presence of the fertile part of the frond, in the same position as
that in specimens of D. teilianum (Walton, 1926 : 213, pi. 17, figs. 17-19), is afforded
by one specimen (PI. 20, fig. 8) where a bare rachis is seen as a continuation of the
main rachis of the frond in the angle between the two divisions of the pinna-bearing
parts of the frond. Careful examination reveals that the first forking of this pre-
sumably fertile rachis was a dichotomy at right angles to the plane of the frond.
The presence of this structure in the angle of the fork supports the assignment of
this plant to the genus Diplopteridium. Parts of the two divisions of a considerably
larger frond (PI. 20, fig. 13) have, situated between them, portions of a naked branch-
ing system (PL 13, fig. 13%) which probably represents the fertile part of the frond.
Genus SPHENOPTERIS Brongniart
Sphenopteris obfalcata Walton
(PI. 20, figs. 16, 17)
Several fragmentary pieces of this frond from Hazel Hill (PI. 20, figs. 16, 17)
exhibit collectively features which justify identification with the type of this species
from the Upper Black Limestones in N. Wales (Walton, 1931 : 363). The frond
appears to have been at least tripinnatifid with a tendency for the pinnae to be
opposite. The ultimate divisions and the form of the pinnae bear some resemblance
to those of Diplopteridium but the ultimate divisions are more spatulate. There are
several genera such as Sphenopteris, Sphenopteridium, Spathulopteris and Rhodea
which have much divided fronds with fine ultimate segments and it is often difficult
to decide to which genus such specimens as these should be assigned.
142 FOSSIL FLORA OF THE DRYBROOK SANDSTONE
Sphenopteris cuneolata L. & H.
(PI. 21, fig. 21)
Four incomplete specimens of a frond resembling Sphenopteris cuneolata L. & H.
(Lindley & Hutton, 1837, pi. 214) occur in the collections from Hazel Hill. Kidston
(1923 : 156) states that the type specimen is lost. The largest fragment (PI. 21,
fig. 21) probably represents part of a frond below the main fork, for the pinnae
seem to get larger towards its distal end and are opposite. The rachis was evidently
about 2 mm. thick as in Lindley & Button's figure. A smaller fragment (¥.42048)
with a rachis about 1-5 mm. thick has its pinnae alternately attached as in the upper
divisions of the type. In the type and in the specimens from Hazel Hill several of
the pinnae appear to be inaequilateral. This is due to folding of the pinnae as ex-
plained in the description of Diplopteridium holdeni. No venation is visible on the
pinnules which have a finely striated surface. The base of the pinna is shortly
decurrent on the rachis. The chief differences which exist between these specimens
and the drawing of the type lie in the greater breadth of the frond and the wider
spacing of the pinnules in the former. Kidston (1923 : 156) includes in the species
a specimen of a frond from the Oil Shales, Calciferous Sandstone Series in Hailes
Quarry near Edinburgh which differs very obviously from the type as figured by
Lindley & Hutton (1837, pi. 214) in having definitely katadromic venation and
in being irregularly and slightly lobed and unlike the type which, in common with
the Hazel Hill specimens, has pinnately divided pinnae.
In view of the loss of the holotype the specimen described here (PI. 21 fig. 21)
is selected as the lectotype of Sphenopteris cuneolata L. & H.
Genus ARCHAEOPTERIDIUM Kidston
Archaeopteridium tschermaki (Stur)
(PI. 21, fig. 22)
One specimen (B.U. 802) which we assign to this species (PL 21, fig. 22) consists
of a clear impression on the fine sandstone of three pinnae attached to a piece of
rachis. The shape and venation of the pinnules match exactly those of Archaeo-
pteridium tschermaki (Stur) Kidston (1923 : 182) which has records in Britain ranging
from the Limestone Coal Group down to the Oil Shale Group.
A second specimen (¥.42498) which has smaller, more rounded pinnules or lobes
with the same type of venation may also, perhaps, be assigned to this species.
FRUCTIFICATIONS
A considerable number of detached fructifications occur on the surfaces of the
sandstone. Two types are of frequent occurrence and are in most cases on the same
surfaces which have impressions of Diplopteridium fronds ; one must be referred to
Telangium sp., the other seems to consist of bunches of cupules.
FOSSIL FLORA OF THE DRYBROOK SANDSTONE 143
Telangiutn sp.
(PI. 21, figS. 23-26)
There are several examples of fructifications consisting of a number of disc-like
bodies connected in some instances by slender axes and each bearing numerous,
pointed fusiform sporangia (PI. 21, figs. 23-26). The individual sporangia appear
to be about 1-3 mm long. There is a close resemblance in form and size between these
fructifications and the Telangium found in connection with Diplopteridium teilianum
(Kidston) (Walton, 1931, pi. 23, figs. I, 3, 4). The sporangia of the Telangium found
associated with Sphenopteris affinis L. & H. are from 2-5 mm. to 3-5 mm. in length
while those associated with Sphenopteris bifidum are about 2 mm. long (Kidston,
1924:446,454).
Calathiops sp.
(PI. 21, figS. 27-29)
There are a number of examples of fructifications which consist of bunches of
elongated structures which are considerably larger than the sporangia in the
Telangium type. They probably were ovuliferous for in at least one example one
of the bunches distinctly resembles an encupuled seed (PI. 21, fig. 2jc). Similar
fructifications have been described by Kidston (1883 : 539, pi. 31, fig. n) from the
Cementstone Group in Liddesdale where they were found in masses in the shales.
The name Schutzia was later applied by Kidston (1924 : 424) to microsporangiate
fructifications. We have decided, however, to include these fructifications of an
indeterminate nature in the genus Calathiops of Goeppert.
Very probably these Telangium and Calathiops types represent pollen-bearing and
ovuliferous organs of Diplopteridium as they are abundant on the same surface as the
fronds of the latter.
SPORAE DISPERSAE
i. Leiotriletes sphaerotriangulus (Loose) Pot. & Kr.
(PI. 21, figS. 30, 31)
Seven specimens ranging in size from 35 /i to 55 p are closely similar to Leiotriletes
sphaerotriangulus (Loose) Pot. & Kr. (1955 : 41). The trilete rays are distinct and
almost reach the spore margin.
2. Punctatisporites minutus Kos.
(PI. 21, figs. 32-34, ?35~37)
Small forms agreeing in size range with P. minutus Kosanke (1950 : 15) are not
infrequent. The spore figured by Potonie & Kremp (1955, pi. n, fig. 120) although
smaller than the known size range (25-35 /*) seems indistinguishable from our
specimens. The rays, in the present examples, are often eccentric and irregular.
The exine usually shows an infrasculpture.
GEOL. 7, 4. II
144 FOSSIL FLORA OF THE DRYBROOK SANDSTONE
There are several other specimens (size range 18-40 fi) of a similar generalized
pattern but having a thinner laevigate exine and thin or indistinct rays (PI. 21,
ngs- 35-37)- Although these specimens are provisonally placed under P. minutus,
they might possibly belong to Calamospora.
Punctatisporites subobesus sp. nov.
(PI. 21, figS. 38-42)
DIAGNOSIS. Size range 45-80 fi (30 specimens) ; circular-subcircular ; rays
simple, more than | spore radius ; exine commonly split open along the trilete
mark to form a i wide triangular fissure ; folding along fissure occasional ; exine
up to nearly 4 /* thick, ± translucent, smooth to infrasculptured ; folds uncommon.
HOLOTYPE. PI. 21, fig. 40 ; 70 /*. B.M., ¥.43701.
COMPARISON. Spores similar to P. obesus (Loose) Potonie & Kremp (1955 : 43)
in habit but distinguishable by their smaller size and longer trilete rays or fissure.
PI. 21, figs. 39-42 represent different degrees of the split trilete area. P. fissus
Hoffmeister, Staplin & Malloy (1955 : 393) and P. debilis Hacquebard (1957 : 308)
are distinguishable by their smaller size, thinner exine, shorter trilete area and
granular sculpture.
4. Punctatisporites spp.
(PI. 21, fig. 43)
The collection contains several specimens of Punctatisporites which cannot be
specifically identified owing to their bad preservation or insufficient number. These
spores range in size from 45 /i to 90 /£ and may include more than one species. The
spore in PI. 21, fig. 43 has simple rays which reach the margin.
5. Calamospora sp. cf. C. mutabilis (Loose)
(PI. 22, fig. 44)
Size range 82-115 X 30-43 ^ (6 specimens), all folded into boat-shaped form ;
trilete mark seen only in one case (PI. 22, fig. 44), rays thin, simple, longest ray 33 p,
a little more than | spore radius ; exine moderately thin, smooth to faintly infra-
punctate, yellow to brown in colour, with no difference in structure or colour in the
contact area.
The spores, in most of their features, are comparable with Calamospora mutabilis
(Loose) (see Potonie & Kremp, 1955 : 49 and Bhardwaj, 1957 : 82).
6. Granulatisporites tennis sp. nov.
(PI. 22, figs. 45, 46)
DIAGNOSIS. Size range 17-25 fi (10 specimens), triangular to roundly triangular,
rays distinct, occasionally with thin lips, reaching spore margin ; exine crowded
with very minute grana, about 20 grana between two rays on the margin ; ornament
present on rays.
FOSSIL FLORA OF THE DRYBROOK SANDSTONE 145
HOLOTYPE. PI. 22, fig. 45 ; 2O fl. B.M., ¥.43708.
REMARKS. The species is distinguished by its small size, minute grana and long
rays bearing similar ornament.
7. Granulatisporites sp. cf. G. orbiculus (Pot. & Kr.)
(PI. 22, figs. 47, 48)
Size range 24-35 /i (12 specimens), triangular to roundly triangular ; rays extend-
ing almost up to spore margin ; preservation of ornament poor, grana apparently
somewhat irregular and partly coalescent. Nearest comparable species is Granu-
latisporites (Cyclogranisporites) orbiculus (Pot. & Kr) Potonie & Lele (1959).
8. Cyclogranisporites amplus McGregor
(PI. 22, figS. 49-51)
Size range 50-90 [i (16 specimens) ; circular, trilete rays not always seen, f spore
radius or more ; ornament of dense grana of variable size and shape, grana usually
less than or up to i /i in diameter, over 100 grana at the margin, exine thickness
variable, secondary folds common.
The present specimens fall within the size range of C. amplus McGregor and agree
in most respects with it. It may be added that the ornament in C. amplus also
appears to be rather variable as in the present forms (cf. McGregor, 1960, PI. n,
fig. 8).
9. Cyclogranisporites sp.
(PI. 22, fig. 52)
Size range 17-40 p (20 specimens) ; circular, trilete mark weak, often not visible,
when present rays f spore radius or longer, ray ends often indistinct ; exine thin,
with frequent compression folds ; ornamentation of densely set minute grana,
grana i variable in shape and size, often obscure on the equator ; state of preserva-
tion unsatisfactory.
10. Planisporites minimus McGregor
(PI. 22, figs. 53-55)
Size range 25-30 /i (4 specimens), triangular to nearly circular ; rays indistinct,
apparently long ; ornament of minute coni or fine hairy spinules.
The few specimens in the present preparations do not represent the overall size
range. They can, however, be assigned to Planisporites minimus McGregor (1960 :
29, pi. n, fig. 9) with which they agree most. It may also be remarked that the
two species P. minimus McGregor and P. delucidus McGregor (1960 : 30, pi. n, fig.
16) are so similar in characters that they would appear to fall within the size range
of a single species.
146 FOSSIL FLORA OF THE DRYBROOK SANDSTONE
ii. Planisporites sp. cf. P. granifer (Ibr.)
(PI. 22, fig. 56)
Single specimen ; 112 X 100 fi, triangular, corners rounded ; triangular fold in
the centre ; exine with i sparsely set short coni. The specimen is slightly larger
but is comparable with P. granifer (Ibr.) Knox (cf. Potonie & Kremp, 1955 : 71 ;
1960 : 38).
12. Verrucosisporites sp.
(PI. 22, fig. 57)
Specimens referable to Verrucosisporites are too few to justify specific determina-
tions. PI. 22, fig. 57 represents an example somewhat comparable with V '. donarii
Potonie & Kremp (1955 : 67).
13. Camptotriletes sp.
(PI. 22, fig. 58)
Single specimen, 105 X 90 fi, roundly triangular, rays f spore radius, ornament of
verrucae up to 9 /* long and up to 8 /i broad, often connected to form crested ridges.
The ornament of the exine supports the assignment of the specimen to Camptotriletes.
14. Cf. Convolutispora sp.
(PI. 22, fig. 59)
A few specimens, showing somewhat convoluted ridges, are probably referable
to Convolutispora. The specimen in PI. 22, fig. 59 recalls C. tessellata H. S. & M.
of Butterworth & Williams (1958, pi. 2, figs. 17, 18).
15. Microreticulatisporites cf. cribellarius (Horst)
(PI. 22, figS. 60, 6l)
Size range 40-55 fi (10 specimens), roundly triangular, rays distinct, extending up
to the spore margin ; reticulum imperfect, i indistinct ; lumina very small, muri
low and barely evident at the margin.
The specimens are closely comparable with and probably indistinguishable from
Microreticulatisporites cribellarius (Horst) Potonie & Kremp (1955 : 97).
16. Microreticulatisporites spp.
(PI. 22, figS. 62, 63)
(i) PI. 22, fig. 62 :
Size range 65-78 fi (3 specimens) ; triangular to roundly triangular, thick rays
almost reaching the equator, gnarled ; muri 1-2 /* wide, not very high, lumina about
equal to or slightly narrower than the width of muri ; reticulum imperfect, appar-
ently coarse, outline minutely undulating, about 20 curvatures between two rays at
the margin, exine 2 fi thick.
FOSSIL FLORA OF THE DRYBROOK SANDSTONE 147
(ii) PI. 22, fig. 63 :
Size range 20-37 fi (a few specimens) ; triangular to ± circular ; rays indistinct,
about f spore radius ; muri about i JLI thick, close, irregular, anastomosing but no
definite reticulum, margin minutely undulating, about 15-20 curvatures between
two rays at the margin.
17. Dictyotriletes sp.
(PI. 22, fig. 64)
Single specimen, probably comparable with D. falsus Potonie & Kremp (1955 :
109 ; Love, 1960, pi. i, fig. 8).
Remarks. There are a few other ill-preserved specimens with somewhat project-
ing muri which are referable to Reticulatisporites. Besides these, one or two speci-
mens may perhaps belong to Knoxisporites.
18. Lycospora cf. bracteola Butt. & Will.
(PL 22, fig. 65)
Size range 45-55 X 40-48 jn (6 specimens), width of cingulum of a spore often not
uniform ; exine faintly granular, cingulum with much finer granules, visible only
under high magnification.
The specimens are comparable with L. bracteola Butterworth & Williams (1958 :
357)-
19. Lycospora spp.
(PL 22, figS. 66-69)
The genus is, on the whole, very poorly represented in numbers of spores, although
more than one type is present. A few specimens with a narrow cingulum and a
thickened inner zone (crassitude) represents one type (PL 22, fig. 66). Another
type (PL 22, fig. 67) somewhat recalls L. granulata. The form in PL 22, fig. 68 has
a wider cingulum and a narrower inner crassitude. The ornamentation of the
cingulum varies from minute grana to short, narrow bacula. The specimen in
PL 22, fig. 69 has a narrow membraneous cingulum, without a crassitude. Types
shown in PL 22, figs. 67-69 are solitary specimens.
20. Anulatisporites anulatus Pot. & Kr.
(PL 22, figS. 70, 71)
Size range 37-68 X 30-53 fi (12 specimens) ; usually oval, occasionally sub-
triangular ; cingulum ± 3 total spore radius, width often not uniform, outer margin
smooth ; trilete mark indistinct to invisible, exine of central area and cingulum
faintly infrasculptured.
The specimens bear close similarity to A. anulatus (Potonie & Kremp 1956 :
112). A few of them (PL 22, fig. 71), provisionally included under this species,
have a comparatively narrower cingulum.
148 FOSSIL FLORA OF THE DRYBROOK SANDSTONE
21. ? Anulatisporites sp.
(PI. 22, fig. 72)
Single specimen ; 90 x 76 p ; oval ; central area nearly triangular, thin ;
cingulum 25 /* wide, roughly equal to diameter of central area ; a peripheral thicken-
ing (limbus?) about 2 /* wide evident ; inner margin of cingulum also similarly
thickened (about 2 /i) ; exine laevigate to infrasculptured ; trilete mark not visible.
REMARKS. The essentially sculptureless cingulum, its smooth outline and
apparent absence of a trilete mark favour the assignment of this specimen to
Anulatisporites. However, a peripheral limbus-like thickening in Anulatisporites
is hitherto unknown. The dark points seen on the cingulum in the photograph
are due to foreign matter on the spore.
22. Densosporites sp.
(PI. 23, fig. 73)
Single specimen ; 56 X 48 fi ; subcircular ; central area infrapunctate ; cingulum
13 /* wide, roughly equal to radius of central area, rugose in appearance owing to
presence of irregular warts of variable size, specially towards the inner margin of
cingulum where they appear to form a ± continuous, narrow, thickened ill-defined
zone. The cingulum towards the periphery is finely wrinkled ; outline nearly
smooth ; trilete mark invisible.
23. Cf. Cirratriradites sp.
(PI. 23, fig. 74)
Single specimen ; 70 X 48 fi ; triangular ; central body 40 X 30 /*, triangular,
distinct ; equitorial zone thin, about 12 fi wide, outline minutely crenulate ; radial
striations visible ; trilete mark clear, rays extending beyond the central body ;
exine of central body and zone minutely granular-punctate.
The preservation of the spore is poor but its appearance supports an attribution
to Cirratriradites.
24. Endosporites sp.
(PI. 23, fig. 75)
Single specimen, 52-5 fi ; subcircular ; body 36 fi, circular, thin-walled ; bladder
nearly 10 /* wide, ± I radius of body, infrareticulate, with radial folds ; outline
smooth ; trilete mark not seen.
25. Remysporites drybrookensis sp. nov.
(PL 23, figs. 76-81)
Cf. Radforth & McGregor, 1956 : 27, pi. i, fig. 7.
DIAGNOSIS. Size range 70-155 x 70-128 /i (50 specimens) ; usually oval in
outline, occasionally circular ; central body large, 60-115 X 60-98 /*, distinct,
FOSSIL FLORA OF THE DRYBROOK SANDSTONE 149
often denser than the bladder, smooth, with frequent cracks, folds uncommon ;
bladder somewhat thick, apparently attached to the proximal side of body, smooth
to faintly infrapunctate or infragranulate, folds uncommon ; trilete mark often not
evident, rays simple, often eccentric, i half the length of the longer axis of the body.
HOLOTYPE. PL 23, fig. 76 ; 108 x 97 /*. B.M., ¥.43719.
REMARKS. The spores are referable to Remysporites Butt. & Will, sensu Potonie
(1960 : 72). They lack good proximo-distal orientation, with the result that the
bladder may be more or less asymmetric and the trilete mark eccentric. The bladder
width (compensated) ranges between 8-32 fi and is approximately J-^ of the longer
axis of the body. The specimens, however, show all variations between the two
extremes and may, therefore, be placed under the same species. The central body
is variously cracked. There is often a concentric and ± continuous fissure along
the margin of the body which produces a lighter zone between the body and the
bladder (PI. 23, figs. 76-80). In many cases, however, this fissure is absent (PI. 23,
fig. 81) and the crack is evidently not an original feature.
The present spores are distinguished from the known species of Remysporites
(Butterworth & Williams, 1958 ; Staplin, 1960) by their consistently smooth and
relatively thick body and bladder and the lack of secondary folds.
2§. Cf. Remysporites sp.
(PI. 23, fig. 82)
Two or three poorly preserved specimens. A more complete one (PI. 23, fig. 82)
measures 175 /i ; body dense, 100 (JL ; bladder thin, as wide as the radius of body,
folded over the central part ; ornament obscure ; trilete mark not seen. The
spore recalls Remysporites in size and appearance but the bladder is somewhat
broader (cf. Butterworth & Williams, 1958 : 386).
27. SPORAE INSERTAE SEDIS
PI. 23, fig. 83 : Single specimen ; 40 /* ; triangular ; central body convexly
triangular, 25 /* ; apparently enclosed by a rather thick structure (Pcapsula), out-
line of spore irregularly lobed ; surface of spore laevigate but somewhat irregularly
thickened at places to form small sinuous ridges (seen only by proper adjustment
of focus). Spore pale yellow, transparent ; trilete mark distinct, rays almost
reaching equator of central body.
PL 23, fig. 84 : Single specimen ; 53 /* ; roundly triangular ; equitorial ridge
(? cingulum) nearly 2-5 /* wide, about £ radius of central area ; rays distinct, open,
almost reaching margin of central area ; exine of central area with irregular, large
mounts ; outline undulated.
THE AGE OF THE DRYBROOK SANDSTONE
There is a close lithological resemblance between the plant-bearing sandstones at
Hazel Hill and the basement beds of the Lower Brown Limestone at Moel Hiraddug
150 FOSSIL FLORA OF THE DRYBROOK SANDSTONE
at Diserth, North Wales. In both, the sandstone is light coloured and friable and
contains, associated with the plants, Conularia or Conulariid fragments, two from
Hazel Hill (B.M., ¥.42431 and B.U. 8oo«) and numerous examples from Diserth
(Manchester Museum, Cwm 3-28). There is also the occurrence in both places of
specimens of smooth axes with stigmarioid scars which hitherto have not been
observed elsewhere.
At the Diserth locality several examples of the fucoid Chondrites plumosus Kidston
occur in association with abundant remains of Lepidophyllum fimbriatum, Kidston.
These two fossils are also recorded by Kidston from the Cementstones at Glencartholm
in the Borders. Among the specimens from Hazel Hill (e.g. B.M., ¥.42449) there
are examples of a fucoid which consists of filaments 0-3 mm. in diameter and up to
9 cm. in length.
These facts suggest that the Hazel Hill and Diserth sediments were deposited in
lagoons or estuaries near the sea and were of the same age as the lower part of the
Calciferous Sandstone Series in the Borders.
The spores, on the whole, suggest a relatively simple and old type of assemblage
in which Punctatisporites constitutes nearly half of the spore population and the
Azonotriletes forms occur in far greater numbers than the Zonales. Very few exam-
ples of saccate genera except Remysporites are present. Monolete or monocolpate
grains are apparently absent. Evidently, the Drybrook spore florule has little
in common with the well-diversified and rich spore complex described by Butter-
worth & Williams (1958) from the Limestone Coal Group and the Upper Limestone
Group of the Scottish Lower Carboniferous. The spore content of the Oil Shale
Group (Love, 1960) which is comparable with that of Butterworth & Williams is
also considerably varied and apparently younger than the Drybrook spore florule.
Our present knowledge of the microspore assemblages from horizons older than the
Oil Shales is very incomplete. It is, however, interesting to mention that Knox
(1959 : 92) reports the occurrence of 13 genera from the base of the Calciferous
Sandstone Series. They are Punctatisporites, Calamospom, Granulatisporites ,
Cyclogranisporites, Lophotriletes , Apiculatisporis, Planisporites, Microreticulatis-
porites, Cristatisporites, Reticulatisporites , Lycospora, Densosporites and Cirratrira-
dites. Ten out of the above genera are also present in our assemblage. On the
whole the Drybrook assemblage of sporae dispersae and that of the base of the
Calciferous Sandstone Series show a close relationship with one another. We are
inclined to believe that the Drybrook spore florule, like that of Knox, indicates a
horizon fairly low in the Mississippian and supports the macrofossil evidence. On
the other hand from the palaeozoological and stratigraphical evidence several
geologists including Welch & Trotter (1960 : 60) place the Drybrook beds much
higher and consider that they are in the Upper Caninia Zone CgSj. The North Wales
floras from the Lower Brown and Upper Black Limestone like the Drybrook Sand-
stone flora are associated with marine animal fossils and may possibly have been
developed under different ecological or climatic conditions from those prevailing
in other areas during the Lower Carboniferous and this might account for the
difference in the floristic composition between them and floras of the same age in
other parts.
FOSSIL FLORA OF THE DRYBROOK SANDSTONE 151
ACKNOWLEDGEMENTS
We cordially thank those mentioned in the Introduction, Dr. Isles Strachan of
the University of Birmingham, who kindly lent its collection to us, and the staff of
the Geological Department of Kingston Technical College for lending specimens
which they had collected. To the Staff of the Department of Palaeontology,
Natural History Museum, London, we are especially grateful for their help and
encouragement and we thank Mr. W. Anderson for assisting us with the photo-
graphy.
One of us (K. M. L.) thanks the authorities of the Colombo Plan for the award of a
scholarship, and the Director, Birbal Sahni Institute of Palaeobotany, Lucknow,
India, for the grant of study leave.
REFERENCES
ALLEN, K. C. 1961. Lepidostrobophyllum fimbriatum (Kidston, 1883) from the Drybrook
Sandstone (Lower Carboniferous). Geol. Mag. Land., 98 : 225-229, pi. 13.
BHARDWAJ, D. C. 1957. The palynological investigations of the Saar Coals, Pt. i. Morphology
of sporae dispersae. Palaeontographica, Stuttgart, 101, B : 73-125, pis. 22-31.
BUTTERWORTH, M. A. & WILLIAMS, R. W. 1958. The small spore floras of coals in the Lime-
stone Coal Group and Upper Limestone Coal Group and Upper Limestone Group of the
Lower Carboniferous of Scotland. Trans. Roy. Soc. Edinb., 68 : 353-39°* pis. 1-4.
CROOKALL, R. 1939. Lycopodiaceous stems (? Cyclostigma Kiltorkense Haughton) from
Mitcheldean in the Forest of Dean. Bull. Geol. Surv. Gt. Britain, London, 2 : 72-77, pis. 5, 6.
FUNKHOUSER, J. W. & EviTT, W. R. 1959- Preparation techniques for acid-insoluble micro-
fossils. Micropaleont., New York, 5 : 369-375, 2 figs.
HACQUEBARD, P. A. 1957. Plant spores in coal from the Horton Group (Mississippian) of
Nova Scotia. Micropaleont., New York, 3 : 301-324, pis. 1-3.
HOFFMEISTER, W. S., SxAFLiN, F. L. & MALLOY, R. E. 1955- Mississippian plant spores from
the Hardinsburg formation of Illinois and Kentucky. /. Paleont., Chicago, 29 : 372-399,
pis. 36-39.
KIDSTON, R. 1883. Report on fossil plants collected by the Geological Survey of Scotland in
Eskdale and Liddesdale. Trans. Roy. Soc. Edinb., 30 : 531-550, pis. 3°-32-
- 1923. Fossil plants of the Carboniferous rocks of Great Britain. Mem. Geol. Surv. Gt.
Brit., Palaeont., 2, 2 : 111-198, pis. 23-47.
- 19230. Fossil plants of the Carboniferous rocks of Great Britain. Mem. Geol. Surv. Gt.
Brit. Palaeont., 2, 3 : 199-274, pis. 48-68.
1924. Fossil plants of the Carboniferous rocks of Great Britain. Mem. Geol. Surv. Gt.
Brit. Palaeont., 2, 5 : 377-522, pis. 92-122.
KNOX, E. M. 1959. Some aspects of microspore morphology. Trans. Bot. Soc. Edinb., 38 :
89-99, 27 figs.
KOSANKE, R. M. 1950. Pennsylvanian spores of Illinois and their use in correlation. State
Geol. Surv. Illinois, 74 : 1-128, pis. 1-16.
KRAUSEL, R. & WEYLAND, H. 1949. Gilboaphyton und die Protolepidophy tales. Senck-
enbergiana, Frankfurt a.M., 30 : 129-152, pis. 1-7.
LINDLEY, J. & HUTTON, W. 1837. The Fossil Flora of Great Britain, 3, pis. i57-23°- London.
LOVE, L. G. 1960. Assemblages of small spores from the Lower Oil Shale Group of Scotland.
Proc. Roy. Soc. Edinb., 67 : 99-126, pis. i, 2.
MCGREGOR, D. C. 1960. Devonian spores from Melville Island, Canadian Arctic Archi-
pelago. Palaeontology, London, 3 : 26-44, pis. 11-13.
POTONIE, R. 1960. Synopsis der Gattungen der sporae dispersae. Beih. Geol. Jb., Hanover.
39, 3 : 1-189, pis- i-9-
GEOL. 7, 4. I2
152 FOSSIL FLORA OF THE DRYBROOK SANDSTONE
R. & KREMP, G. O. W. 1955. Die sporae dispersae des Ruhrcarbons. Palaeonto-
graphica, Stuttgart, 98, B : 1-136, pis. 1-16.
1956. Die sporae dispersae des Ruhrcarbons. Palaeontographica, Stuttgart, 99,
B : 85-191.
POTONIE, R. & LELE, K. M. 1959. Studies in the Talchir flora of India — Sporae dispersae
from the Talchir Beds of South Rewa Gondwana Basin. Palaeobotanist, Lucknow, 8 :
22-37, pis. 1-3.
RADCHENKO, G. P. 1956. New families and genera of fossil plants, II. All Union Inst. Geol.
Res. (n.s., Palaeont.), 12 : 185-264. (In Russian.)
- 1957- Morphological and anatomical peculiarities of some early Carboniferous vegetative
types from Kouznetsk Province. Kryshtofovich Memorial Volume, pp. 33-54, pis. 1-4.
Moscow & Leningrad.
RADFORTH, N. W. & MCGREGOR, C. 1956. Antiquity of form in Canadian plant micro-
fossils. Trans. Roy. Soc. Canada, Ottawa (3, 5) 50 : 27-33, pis. 1-3.
SIBLY, T. F. & REYNOLDS, S. W. 1937. The Carboniferous Limestone of the Mitcheldean
area, Gloucestershire. Quart. J. Geol. Soc. Lond., 93 : 23-51.
STAPLIN, F. L. 1960. Upper Mississippian plant spores from the Golata Formation, Alberta,
Canada. Palaeontographica, Stuttgart, 107, B : 1-40, pis. 1-8.
TROTTER, F. M. 1942. Geology of the Forest of Dean Coal and Iron Ore Field. Mem. Geol.
Surv.Gt. Britain, London, vi + 95 pp., 5 plates.
WALTON, J. 1926. Contributions to the knowledge of Lower Carboniferous plants, Pts. I, II.
Philos. Trans., London (B) 215 : 201-224, P^s- X6, 17.
— 1931. Contributions to the knowledge of Lower Carboniferous plants, Pt. III. Philos.
Trans., London (B) 219 : 347-379, pis. 23-26.
WELCH, F. B. A. & TROTTER, F. M. 1960. Geology of the Country around Monmouth and
Chepstow. Mem. Geol. Surv. Gt. Britain, London, viii + 164 pp., 2 pis.
PLATE 19
Scutellocladus variabilis gen. et. sp. nov.
FIG. i. Leafy branch with closely set leaf-bases. B.M., V.42433. Nat. size. Syntype.
Fig. i a. Part of the same specimen, x 3.
FIG. 2. Small forking branch with leaves still attached showing the areas of attachment of
the leaves with a longitudinal ridge representing the leaf trace. B.M., V.42538. x 3.
Syntype.
FIG. 3. Dichotomously forking leafy branch. B.U. 800. Nat. size.
FIG. 4. Dichotomously forking branch showing leaf-scars with central punctiform scar.
B.M., ¥.42558. X4.
FIG. 5. Small leafy twig. The apical region of one branch is seen. B.M., V.42477- Nat.
size.
FIG. 6. Smallest size of leafy branch found. U.G. Pb-3452. Nat. size.
Cf. Stigmaria
FIG. 7. Impression of the outer surface of an axis bearing a number of stigmarioid scars.
B.M., V. 42428. Nat. size.
Bull. B.M. (N.H.) Geol. 7, 4
PLATE 19
la
LELE & WALTON
GEOL. 7, .J. 12§
PLATE 20
All figures are natural size
Diplopteridium holdeni sp. nov.
FIG. 8. Specimen showing central part of a frond with remains of fertile rachis. B.M.,
¥.42453. Holotype.
FIG. 9. Base of a frond with small basal pinnae. B.M., ¥.43004. Paratype.
FIG. 10. Part of a frond showing apparently inaequi lateral pinnules on the pinnae. B.M.,
¥.42487. Paratype.
FIG. ii. The two terminal portions of a frond. B.U. 804^.
FIG. 12. Parts of the two divisions of a frond showing clearly the form of the pinnae. B.M.,
¥.42524.
FIG. 13. Part of the largest frond. Portions of its two divisions on the left and right and
parts of presumably its fertile rachis (x) between. B.U. 804.
FIG. 14. Middle part of an apparently sterile frond. B.M., ¥.42474. Paratype.
FIG. 15. Parts of a frond with very slender ultimate segments. B.M., ¥.42457.
Sphenopteris obfalcata (Walton) n. comb.
FIG. 16. PMain rachis of a frond with proximal parts of six subopposite primary pinnae-
B,U. 803.
Fig. 17. Part of a frond to show form of pinnae and their ultimate divisions. B.M., ¥.42488.
Lepidophyllum cf . fimbriatum Kidston
FIG. 18. Distal part of a leaf with a single row of short hairs on each side. B.M., ¥.42516.
FIG. 19. Almost complete leaf showing expanded base. No hairs are evident. B.M.,
¥.43442.
FIG. 20. Complete leaf showing acute apex. B.M., ¥.42531.
Bull. BM. (N.H.) Geol 7, 4
PLATE 20
LELE & WALTON
PLATE 21
Sphenopteris cuneolata L. & H.
FIG. 21. Part of a frond with parts of nine pairs of pinnae. Lectotype ; B.U.Soi. Nat.
size.
Archaeopteridium tschermaki (Stur) Kidston
FIG. 22. Parts of three pinnae. B.U.8O2. X2.
Telangium sp.
FIGS. 23, 24, 25 and 26. Groups of fructifications which appear to consist of microsporangia
attached to discs at the ends of slender rachides. 23, 24 B.U.8O2 and B.U.Soo ; 25, 26 B.M.,
V-42435 and V-43443. x 3.
Calathiops sp.
FIG. 27. Fructification with a cupule c, probably ovuliferous, shown at c. B.M., V. 42432.
X3-
FIG. 28. Fructification probably ovuliferous. B.U.8o2a. X2.
FIG. 29. Cupule with numerous lobes. B.M., V. 42478. X2.
Sporae dispersae
(All figures are X 500. Locality and horizon same as for the macrofossils.)
FIGS. 30, 31. Leiotriletes sphaerotriangulus (Loose). V. 43723, V. 43705.
FIGS. 32-34. Punctatisporites minutus Kos. ¥.43721, ¥.43706, ¥.43707.
FIGS. 35-37. ? Punctatisporites minutus Kos. ¥.43706, ¥.43707, ¥.43713.
FIGS. 38-42. Punctatisporites subobesus sp. nov. (Holotype : Fig. 40). ¥.43700, ¥.43720,
¥.43701, ¥.43722, ¥.43726.
FIG. 43. Punctatisporites sp. ¥.43709.
Bull. B.M. (N.H.) Geol. 7, 4
PLATE 21
38
LELE & WALTON
PLATE 22
A II figures x 500
FIG. 44. Calamospora sp. cf. C. mutabilis (Loose) ¥.43707.
FIG. 45. Granulatisporites tennis sp. nov. Holotype. V-437o8.
FIG. 46. Granulatisporites tennis sp. nov. ¥.43714.
FIGS. 47, 48. Granulatisporites sp. cf. G. orbiculus (Pot. & Kr.). ¥.43702, ¥.43708.
FIGS. 49-51. Cyclogranisporites amplus McGregor. ¥.43704, ¥.43721, ¥.43707.
FIG. 52. Cyclogranisporites sp. ¥.43705.
FIGS. 53-55. — Planisporites minimus McGregor. ¥.43705, ¥.43711, ¥.43712.
FIG. 56. Planisporites sp. cf. granifer (Ibr.). ¥.43724.
FIG. 57. Verrucosisporites sp. ¥.43716.
FIG. 58. Camptotriletes sp. ¥.43701.
FIG. 59. Cf. Convolutispora sp. ¥.43700.
FIGS. 60, 61. Microreticulatisporites cf. cribellarius (Horst). ¥.43710, ¥.43707.
FIGS. 62, 63. Microreticulatisporites spp. ¥.43703, ¥.43708.
FIG. 64. Dictyotriletes sp. ¥.43726.
FIG. 65. Lycospora cf. bracteola Butt. & Will. ¥.43707.
FIGS. 66-69. Lycospora spp. ¥.43710, ¥.43715, ¥.43708.
FIGS. 70, 71. Anulatisporites anulatus Pot. & Kr. ¥.43707.
FIG. 72. ? Anulatisporites sp. ¥.43724.
Bull. B.M. (N.H.) Geol. 7, 4
PLATE 22
70
71
68
LELE & WALTON
PLATE 23
All figures X 500
FIG. 73. Densosporites sp. ¥.43720.
FIG. 74. Cf. Cirratriradites sp. ¥.43700.
FIG. 75. Endosporites sp. ¥.43727.
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LELE & WALTON
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FOSSIL INSECTS FROM
THE LOWER LIAS OF
CHARMOUTH, DORSET
F. E. ZEUNER
BULLETIN OF
THE BRITISH MUSEUM (NATURAL HISTORY)
GEOLOGY Vol. 7 No. 5
LONDON: 1962
FOSSIL INSECTS FROM THE LOWER LIAS
OF CHARMOUTH, DORSET
BY
FREDERICK EVERARD ZEUNER
Pp. I53-I71 ; ^s- 24-27
/ ".'.'<? 4?
BULLETIN OF
THE BRITISH MUSEUM (NATURAL HISTORY)
GEOLOGY Vol. 7 No. 5
LONDON: 1962
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. 7, No. 5 of the Geological
(Palaeontological) series.
Trustees of the British Museum 1962
PRINTED BY ORDER OF THE TRUSTEES OF
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Issued August, 1962 Price Twelve Shillings
FOSSIL INSECTS FROM THE LOWER LIAS
OF CHARMOUTH, DORSET
By FREDERICK EVERARD ZEUNER
SYNOPSIS
The paper discusses the environmental conditions prevailing in the Lower Lias of England
with particular reference to the insect faunas contained in the sediments. It also discusses the
degree of salinity insects can withstand. The environment of the Lias is compared with that
of the Bombay coast of today, with its numerous islands and inlets, and mud sedimentation.
The new fauna from Charmouth, with six dragonflies, one orthopteron and three beetles, is
described. The dragonflies all belong to the almost extinct Anisozygoptera and are among the
largest known. The orthopteron belongs to the almost extinct Prophalangopsidae, being their
most primitive representative and linking them with the Gryllacrididae. Of the beetles, one is
a Cupedid, a member of a family that flourished in the late Palaeozoic ; the two others are of
unknown affinities. The structure and coloration of the elytra are analysed and interpreted
in detail, providing information about the process of fossilization. The taxonomic status of the
two species of Holcoelytrum is discussed and types are designated.
i. GENERAL PALAEOBIOLOGIC AL CONSIDERATIONS
AN important collection of fossil insects has recently been made by Mr. J. F. Jackson
from the " Flatstones " and related deposits near Stonebarrow, Charmouth, on the
Dorset coast. The preliminary classification of the specimens, which have been
acquired by the British Museum (Natural History), has shown that both in the
composition of the fauna and in the preservation they resemble those of the Lower
Lias of Gloucestershire, Warwickshire and Worcestershire, though there are some
significant differences. Conditions of life, death and fossilization were evidently
somewhat similar. In detail, however, several problems arise when the faunal asso-
ciation is studied. Some insects, for instance, including a dragonfly (In. 49573),
rest directly on or are closely associated with ammonites.
According to Mr. Jackson (written communication) the following ammonites
occur in the deposit : Asteroceras obtusum, Xipheroceras dudressieri, Xipheroceras sp.,
Promicroceras planicosta, very abundant. Fishes are present also and, according to
Mr. Jackson's observations, mostly found where plant remains occur. The following
species have been recorded : Dapedium granulatum, Dapedium sp., Pholidophorus
sp. and Chondrosteus sp. (very rare) .
Lamellibranchs are rare, except Inoceramus, and gastropods are absent. Plants,
apart from driftwood, are also rare.
This list of fossils indicates salt-water, with tolerable living conditions in the
water itself, whilst conditions on the sea-floor appear to have been unsuitable for
many groups one might normally expect to be present. The environment would
thus have been essentially marine, and there is a contradiction in the presence of
numerous insects together with a marine fauna. The first possibility requiring
consideration is that sea-water was less saline in Jurassic times than it is at the
present day. An assessment can be made with the aid of Conway's important paper
GEOL. 7, 5. I3§
156 FOSSIL INSECTS FROM THE LOWER LIAS OF CHARMOUTH
(1943). From his data it can be calculated that the salinity of the Jurassic ocean
was of the order of 3-0%, x in other words only slightly less than the ocean of the
present (3'4°/o)-
Since British Liassic insect faunas contain fair numbers of species which pass
their larval lives in water, the resistance of insects to sea-water needs consideration.
Some Apterygota are able to stand temporary submergence and to float on the water
of rock pools which are their ordinary environment. A collembolon, Lipura maritima
L., lives on the shores of Britain ; it is frequent for instance at Selsey in Hampshire
on the quiet surface of pools. Another species of the same order, Actaletes neptuni
Giard, lives on the coast of France where it is temporarily submerged by the tide.
But since Apterygota have not yet been recorded from the British Lias, these forms
merely indicate possibilities of adaptation. Of greater interest are certain species
of Diptera, the larvae of which live on decaying organic matter in jetsam accumu-
lations. They withstand frequent wetting and temporary submergence. A few
Diptera are known from the Lias, but nothing is known about their larval modes
of life.
There are, however, two important groups with aquatic larvae, the adults of which
are well represented in the British Lias, the dragonflies (Odonata) and the caddisflies
(Trichoptera). Both are known to occur in brackish water, and their presence in
the Baltic Sea has received some attention.
Leander (1901) gave a list of the insects occurring in the sea-water west of Helsinki,
Finland. He quoted twenty-one species, comprising one mayfly, nine caddisflies,
two bugs, three mosquitoes (two Chironomus, one Ceratopogon) and six water beetles.
These species were found among the islands and associated with the jelly-fish,
Aurelia aurita, the worm, Nereis diversicolor, the barnacle, Balanus improvisus, the
bivalve, Cardium edule, and others. This mixture of freshwater insects and marine
species exists in water with about 0-5% of salt.
Silfvenius (1905) reported on the caddisflies found at the entrance of the Finnish
Gulf at Tvarminne. The water contains 0-5-0-6% of salt, and no fewer than twenty-
four species of caddisflies live in it as larvae, six of them being abundant. It k note-
worthy that the bladder- wrack, Fucus sp., serves both as food for the larvae and as
building material for their cases.
Ussing (1918) studied the insects of the Randers Fjord in Denmark, in which the
concentration of salt decreases inland. He found the following :
Salt content Insects present
Surface i-7% "1 ^, .
TN j-u n/ X Chironomus larvae only.
Depth 2-4% f
Surface 1-2% "\ One dragonfly, i water bug, 4 water beetles, i leaf
Depth 1-9% J beetle, Chironomus.
Surface 0-5% \ Two caddisflies, i dragonfly, i bug, 4 water beetles,
Depth 1*3% J 2 leaf beetles, Chironomus, i moth.
Surface 0-14%^ T , .., , ,
T\ 2.1, In/ )* Insect life abundant.
Depth o-i6%J
1 Almost the same value is obtained, whether the calculation is based on sodium or on the total of
chlorides.
FOSSIL INSECTS FROM THE LOWER LIAS OF CHARMOUTH 157
This evidence shows that insect life does not altogether shun salt water, although
at concentrations over 0-2% very few species are able to exist, chief among them
dragonflies, caddisflies and Chironomus mosquitoes. Only the caddisflies, however,
have gone some way towards adapting themselves to this environment, as shown
by the use of Fucus as food as well as building material.
The Recent examples given can be supplemented by a Miocene locality (Zeuner,
1938), the Hydrobia Limestone of the Mayence Basin. The brackish character of
the water may here be suspected, though not proved, by the abundance of the
gastropod Hydrobia sp.1. The analysis of the insect fauna presents the following
picture :
Terrestrial and flying individuals . 139
Aquatic individuals . . . More than 8
The aquatic individuals comprise two adult water beetles and larvae of Hydro-
philidae, two larvae of dragonflies, and larvae of Diptera (Stratiomyidae) . Though
the larvae have not all been studied, their total will remain small. This particular
fauna, from Mombach, differs, in the scarcity of aquatic larvae, from neighbouring
localities in which whole beds are composed of the cases of caddisfly larvae. Locally,
therefore, and at certain times, the water was inhabited by enormous numbers of
these insects. Even at Mombach, however, where no caddis cases have been found,
adult caddisflies comprise 50% of the fauna. Adding to these the dragonflies, being
aquatic as larvae, and the larvae of the Hydrophilidae and Stratiomyidae, the total
of insects dependent on water rises to 63%. This composition suggests that at
Mombach the water itself was only partially inhabitable, and this for certain insects
only, whilst at other places in the neighbourhood the water must have harboured
an abundance of insect larvae. It suggests that the salinity of the lake in which
the Hydrobia Limestone formed was near the upper limit of concentration bearable
by insects, and that in the immediate vicinity enough fresh water, possibly from
springs, was available. Such diversity of local biotopes, not evident from the study
of the embedding sediment, is by no means exceptional.
On the shore of the Dead Sea in Jordan, for example, a few freshwater pools are
fed by springs, as at 'Ain Feshka. They support fishes (Cyprinodon sophiae, C. cypris,
C. dispar). Only about 10 ft. of beach gravel separates these fishes from the deadly
brine of the sea, and on this beach the writer found dead water beetles and locusts.
If this combination of species occurred in a fossil context, one would almost certainly
credit the Dead Sea with a fish fauna, with insects living in its water as larvae, and
with other insects living in the neighbourhood. It is this complexity of the local
biotopes that makes the environmental interpretation of fossil insect faunas some-
what difficult. The identification of groups with special environmental requirements
and the relative frequency of individuals in the various environmental groups is,
however, of considerable use as shown by the examples here quoted.
The Liassic deposits of the south-western Midlands have yielded a fauna which
may be regarded as representative of woodlands interrupted by water readily in-
1 The Oligocene of the Rhine Rift is rich in salt deposits, and the partly saline character of the Hydrobia
Limestone was first suggested by Wenz (1921), though he was not using evidence based on insects.
158 FOSSIL INSECTS FROM THE LOWER LIAS OF CHARMOUTH
habitable by insects. Dense vegetation on the water's edge is suggested. The pre-
sence of water of a salinity approaching that of the sea cannot be excluded, but there
must have been plenty of accumulations of water less saline than 0-2%, to enable
the abundant fauna of caddisflies, dragonflies and others to rear their larvae. The
presence of humid ground is indicated by a group of crickets which appears to be
related to Pteronemobius , and which burrows in moist ground near puddles of water
under the surface. Many other Orthoptera Saltatoria are conspicuous in this fauna,
the Prophalangopsidae being an almost exclusively Jurassic family which appears
to have been living among leafy vegetation. The Panorpoid Complex is well repre-
sented, apart from caddisflies, by the Orthophlebiidae, the larval environment of
which is unfortunately not yet known.
Turning now to the insect fauna of the Charmouth Lias, of which 434 specimens
are known, it is noteworthy that no aquatic larvae have so far been discovered.
Adult caddisflies, too, are completely absent. Among the remainder, the beetles
(38-1%) and the Saltatoria (20-8%) dominate. Other groups are represented in
small numbers only. These are the Panorpoid Complex (3-2%), Odonata (2-5),
Rhynchota (2-5%) and Blattodea (1-8%). 31-1% cannot be classified without
further detailed work, and many of these are too poorly preserved to be of interest.
If one compares this list with the brackish faunas quoted above, it becomes clear
that the insects of Charmouth are at least predominantly derived from another
environment, and that their life cycles were not dependent on the water in which
they were embedded and which, presumably, was too saline.
Furthermore, the numerical distribution of insect groups in the Charmouth fauna
is clearly a function of their resistance to mechanical disintegration. This is why
beetles are conspicuous. With them must be ranked the bugs which have hard
bodies and, to a lesser extent, the cockroaches. Wings of dragonflies also are resistant
to decay in water, and, unlike specimens from the Lias of the Midlands, no bodies
have been bound.
The fragile forms belong to the grasshoppers and to the Panorpoid Complex.
The former are often very poorly preserved ; many wings are folded over as if by
a changing water current. But there are a number of legs, and a few specimens
appear to be almost complete with body. This group, therefore, is rather better
preserved than the remainder. Apart from the single specimen of Protohagla langi
sp.n., which is a Prophalangopsid, the great majority of other Saltatoria (if not all)
belong to the Elcanidae. This well-known Jurassic family had enlarged spines on the
hind tibiae with which it was able to swim, the body being held on the water by
surface tension and cutaneous fat, whilst the hind tibiae were dipped into the water,
serving as oars.1 The Elcanidae, which are the ancestors of the Tertiary and Recent
Tridactylidae which live on the edge of water and are likewise able to swim, were
relatively fragile insects. Their abundance implies that the shore was not far away,
and that the surface was not often disturbed by wave action. Salt water would
not have affected them.
The Panorpoid Complex from Charmouth contains several groups, Neuroptera,
Mecoptera (of which Orthophlebia is represented) and Diptera. In addition, there is
1 First noted by Handlirsch (1908), phylogenetic relations discussed by Zeuner (1938).
FOSSIL INSECTS FROM THE LOWER LIAS OF CHARMOUTH 159
a solitary Hymenopteron of the sawfly type. All these are poorly preserved, and their
number may increase as the undetermined remainder is studied. They provide
further evidence that the Charmouth insects had undergone transport and decay
before they were embedded in the sediment.
Whilst this is obvious even from the condition of many Elcanidae, the more
resistant beetles bear it out in an interesting manner. There are many isolated
elytra, though bodies, with or without elytra, are not rare. Legs and other appendages
are lost without exception. The condition of the Coleoptera is in this respect remi-
niscent of those found in present-day jetsam, in which they dry out periodically,
acquire some buoyancy and are thus washed away and re-deposited several times.
Jetsam sedimentation of insects has been described by Trusheim (1929), Schwarz
(1939) and Zeuner (1938 : 151). These authors agree that off-land winds play an
important part in such formations.
In the Charmouth Lias, jetsam is not likely to have been the cause of the presence
of insects. If so, there should be evidence in the form of belts of concentrated frag-
ments of vegetation mixed with marine forms as well as insects. The state of pre-
servation of the present material does not support this idea.
Since the sediment must have been a calcareous mud deposited near land, the
possibility of direct wind transport of insects on to wet mud-flats at low tide has to
be mentioned. It was so in the case of the Solnhofen Limestone of Upper Jurassic
age (Abel, 1929 ; Zeuner, 1939 : 20). Under such conditions, whole insects are often
blown on to the flats, where they remain stuck, unable to rise again. They are thus
not infrequently embedded in positions indicating their struggle to free themselves.
That this interpretation does not apply to the Charmouth Lias is evident.
There remains the possibility of flotsam, of prolonged drifting and slow sinking,
followed by incorporation in soft calcareous and bituminous mud, under either tidal
or non-tidal conditions. Drifting is partly suggested by the fragmentary condition
of most of the insects, though this may be due to other causes such as attack by
predators. There is, however, conclusive evidence for extreme water-logging. Several
of the beetles are compressed dorsoventrally, which is possible only when they were
softened by prolonged wetting and decay. Elytra are often flattened, though by
no means always, and they show wrinkles which were produced as the flattening
proceeded under the weight of the freshly forming sediment. Furthermore, the
wing of Petrophlebia anglicanopsis, a dragonfly to be described later in this paper, has
its margin characteristically frayed as in modern insect wings that have begun
to decompose in water.
These observations favour the interpretation of the insects as flotsam, as indeed
does the composition of the fauna. The same can be applied to the Midlands Lias,
but there the percentage of fragile insects is so much higher, and the abundance of
caddisflies so suggestive of less saline conditions that the environment cannot have
been precisely the same.
In the light of the evidence cited from other localities and of the observations made
on the material from the Midlands Lias and that of Charmouth respectively, the
following deductions may be made concerning the environment. In both areas, mud
sedimentation occurred under protected conditions and proceeded in the virtual
160 FOSSIL INSECTS FROM THE LOWER LIAS OF CHARMOUTH
absence of wave action. The marine fauna may be regarded as implying that tides
were present, and there are plenty of present-day areas of sedimentation that fulfil
these conditions, especially on the mangrove coasts of the tropics. Those known to
the writer are on the coast of Bombay and the Gulf of Cambay. The coast north of
Bombay is particularly suitable for comparison. There are numerous low islands
which, going inland, fuse into strips of land interrupted by numerous water inlets
of varying width. These, in turn, link up with rivers which reduce the salinity of
the water until, some 20 miles inland, it is fresh. Mud sedimentation under quiet
tidal conditions can be observed in many places. Where tested, the mud is both
calcareous and rich in organic matter. Dense vegetation on the islands and the
shores harbours a rich fauna of insects, and marine species penetrate as far inland as
decreasing salinity will permit them to go. In such an area, one would place the
Charmouth Lias among the islands near the open sea, and the Midlands localities
within the inlets.
The complete description of the fauna from Charmouth will require considerable
time, partly because the poor state of preservation imposes a heavy strain on the
eyes of the investigator, and partly because the fragments require large-scale
comparison with more complete material from a great variety of insect orders. In
this first descriptive part, all the remains of Odonata are described and referred to
six species, three of which are new. In addition, the new Prophalangopsid orthopteron
is described because of its phylogenetic interest, together with three species of
beetles.
2. SYSTEMATIC DESCRIPTIONS
Order ODONATA
Suborder ANISOZYGOPTERA Handlirsch
DIAGNOSIS. Nodus placed at great distance from the base of the wing.
DISTRIBUTION. Mainly Jurassic, with two Recent species.
Family LIASSOPHLEBIIDAE Tillyard
EMENDED DIAGNOSIS. Tillyard's definition (1925 : n) has now to be modified in
so far as the discoidal cell of the fore wing is sometimes closed basally.
DISTRIBUTION. So far known from the British Lias only, with two genera con-
taining nine species (including those here described for the first time).
Genus PETROPHLEBIA Tillyard (1925 : u)
TYPE SPECIES. Petrophlebia anglicana Tillyard.
Petrophlebia anglicanopsis sp. n.
(PL 24, figs, i, 2)
DIAGNOSIS. Hind wing with CuA2 less curved than in the type species, and with
indistinct distal portion.
FOSSIL INSECTS FROM THE LOWER LIAS OF CHARMOUTH 161
MATERIAL. The holotype, British Museum (Nat. Hist.) In. 49573, Jackson Coll.,
(PL 24, fig. i) with counterpart ; and a second specimen, In. 59376, from the same
locality, with counterpart (PL 24, fig. 2).
LOCALITY. Lower Lias, Flatstones, Stonebarrow, Dorset.
PARTS KNOWN. Portion of hind wing, and ? apex of wing.
DESCRIPTION. The holotype is a fragment 34-5 mm. long, and 15 mm. wide. It
thus belongs to a large species. From the preserved portions of M4, Cu, and Cu2 a
total length of about 80 mm. can be computed. This is more than in P. anglicana
as estimated by Tillyard. The principal new specimen is part of a hind wing, seen
from the upperside. This has been ascertained from the condition of Cu1} which is
negative (depressed), whilst Cu2 is a positive (raised) vein.
The venation leaves no doubt that this is a Petrophlebia, and this is confirmed by
the pretty colour pattern (not mentioned by Tillyard) produced by a dusky pig-
mentation present along all the cross- veins.
Fragments of R and all branches of M are preserved, but of no particular interest.
The area between M4 and Cut and Cu2 is narrower towards the apex. The long middle
portion of Cu2 is remarkably straight compared with the type species and it disappears
suddenly in the reticulated cross-venation, whilst in P. anglicana it can be traced
as running in a curve down to the hind margin. The hind margin itself is not pre-
served.
REMARKS. This fragment (PL 24, fig. i) shows, oddly enough, about the same
portion of the wing as does the type species, so that a close comparison is possible.
It reveals the differences mentioned ; they make a specific separation necessary.
The new species, P. anglicanopsis, is nevertheless closely related to the type
species.
The holotype of P. anglicanopsis is partly covered by a fragment of an ammonite,
probably Asterocems obtusum (J. Sowerby). The marine context of sedimentation is
thus evident, and the poor condition of the wing, with its corroded edge, suggests
prolonged drifting.
The second specimen is tentatively referred to this species, though it shows some
resemblance to Diastommites liassina (Strickland). It will be necessary to study the
affinities of this species with care, as it may prove to be closely related to Petrophlebia.
Genus LIASSOPHLEBIA Tillyard (1925 : 13)
TYPE SPECIES. Liassophlebia magnifica Tillyard (1925 : 14).
REMARKS. Apart from the type species, Tillyard included in this genus one species
first described by Hagen (1850) as Heterophlebia westwoodi and later (Hagen, 1866)
transferred to Tarsophlebia. This assignation was accepted by Handlirsch ; it
established the relationships of the entire family subsequently erected by Tillyard.
Tillyard further described three new species. Two new species are described in
the present paper, and some additional information concerning the type species is
provided. There is, in addition, a fragment which cannot be assigned to a species
with certainty, though it clearly belongs to the genus.
162 FOSSIL INSECTS FROM THE LOWER LIAS OF CHARMOUTH
Liassophlebia magnified Tillyard
(PI. 27, fig. i)
1925 Liassophlebia magnified Tillyard, p. 15, pi. i, fig. 3 ; pi. 2, fig. 4 ; text-figs. 3, 4.
MATERIAL. British Museum (Nat. Hist.), In. 64000 (PI. 27, fig. i), In. 59106 and
In. 49213, all from The Flatstones, Stonebarrow, Charmouth, Dorset. All with
counterparts.
EMENDED DESCRIPTION. Tillyard's description and illustration of the anal area
of the hind wing (poorly preserved in the holotype) can be improved with the aid
of In. 64000, which shows the details with unusual clarity (PI. 27, fig. i). Most im-
portant, there is an anal angle after all, and as sharp as in most Anisoptera, as well
as a large triangular basal cell, into which a short, blindly-ending cross- vein protrudes
from A. Tillyard was right in suspecting that the subquadrangle contained a group
of three cross-veins, which he dotted in his text-fig. 4, except that the third, which
points downwards, does not join up with another vein, but ends blindly. A similar
" blind end " protrudes into the discoidal cell. The basal vein of the subquadrangle
continues quite straight beyond this structure down to the posterior angle of the
wing.
REMARKS. This is the only dragonfly species common to both the Charmouth
and the Midlands Lias.
Liassophlebia jacksoni sp. n.
(PI- 25)
DIAGNOSIS. Hind wing with 14 postnodals.
HOLOTYPE. British Museum (Nat. Hist.), In. 53999, with counterpart. Jackson
Coll., the only specimen.
LOCALITY. Lower Lias : Flatstones, Stonebarrow, Charmouth, Dorset.
PARTS KNOWN. Hind wing only, portion beyond pterostigma not preserved.
DESCRIPTION. As for Progonophlebia from the Lias of Gloucestershire (Zeuner,
1958), this description and those that follow use exactly the same venational nomen-
clature as Tillyard (1925). It will thus be possible to compare the species and identify
material. It does not, however, imply that I am in agreement with Tillyard regarding
the names given to certain veins. The matter, which raises the problem of the crossing
of tracheae in the immature wing, is too complicated to be discussed here and requires
further studies not only of Recent material but of fossils.
Total length of hind wing about 83 mm. (computed value ; 67 mm. preserved).
This compares with 70 mm. in L. magnified. Greatest width, 28 mm., compared with
21 mm. in L. magnified.
Pterostigma not fully preserved, basal side apparently vertical. Nodus situated
nearer the base than the tip of the wing. Fourteen postnodals (only 10 in L. magnified) .
Subquadrangle traversed by one strong vein, which is concave towards the apex.
M1A weak, reduced to a short length, and joining M2. More distad, the two veins
separate again. This condition is confirmed by the counterpart specimen. Oblique
vein between M2 and Ms present and situated below the seventh postnodal. Between
FOSSIL INSECTS FROM THE LOWER LIAS OF CHARMOUTH 163
M4 and Cux, distad of the triangle, four rows of cells (only two in L. magnified}.
Base of wing with straight vertical vein as in Aeschna. Dark pigmentation at the
base in two distinct patches, one on Sc -}- R, the other on Cu.
REMARKS. This species is much larger than L. magnified Tillyard, described from
Warwickshire, and several of the venational characters are in keeping with this,
as they strengthen the large wing.
The size of the wing has been computed in the following way. If the distance
from the nodus to the pterostigma is used as a basis for the calculation, the total
length would amount to 83-5 mm. On the other hand, if the width is used, assuming
proportions similar to those of L. magnifica, the wing would have been 93 mm. long.
This is improbable, since the preserved length between nodus and pterostigma should
then be longer than it actually is and contain even more than 14 postnodals. Thirdly,
assuming the proportions of both species to be the same and using the distance from
nodus to pterostigma, one arrives at 78 mm. This is too short, for the postnodal
section of the new species is proportionately longer than the antenodal. Thus, the
most likely length is 83-5 mm., and it emerges that the new species is much broader
then L. magnifica. If it had the same length /width ratio as this species, it should
be only 25 mm. broad instead of 28 mm. as measured. It is safe, therefore, to regard
the size given in the description as approximately correct. The insect must have pre-
sented a magnificent sight, with its wing-span of between 7 and 8 inches, when
flying over the estuaries of Liassic Charmouth.
The difference in pigmentation, compared with the type species, confirms that
the Charmouth species cannot be a large individual of the Warwickshire species.
The new species is named after Mr. J. F. Jackson of Charmouth, the indefatigable
collector of Dorset fossils. Liassophlebia jacksoni is the seventh species of the genus
to be described. The genus must have been flourishing in the British Lias, and it is
curious that it has not yet been found elsewhere.
Liassophlebia gigantea sp. n.
(PL 27, fig. 2)
Diagnosis. Very large Liassophlebia with cross- vein linking Cu with arculus in
fore wing so as to form a triangle.
HOLOTYPE. British Museum (Nat. Hist.), In. 51030, with counterpart. Jackson
Coll., the only specimen.
LOCALITY. Lower Lias : Woodstones, Black Ven, Charmouth, Dorset.
PARTS KNOWN. Basal portion of fore wing.
DESCRIPTION. Two very strong antenodals (called " hypertrophied " by Tillyard
in other species). From these and from the width between C and Cu the length of
the wing can be calculated. The former index gives 82 mm., the latter 84 mm. for
the fore wing. The size of this species, therefore, must have been about the same as
that of L. jacksoni.
The arculus is continued by an exceptionally strong vein on the distal side of the
discoidal cell. This vein is approximately at right angles to the long axis of the wing ;
in fact, it is very slightly turned back. In L. magnifica, it is distinctly turned forwards.
164 FOSSIL INSECTS FROM THE LOWER LIAS OF CHARMOUTH
It is also distinguished by a cross vein which connects Cu with the arculus, whilst
in L. magnified the discoidal cell is open at the base.
From L. wither si Tillyard (which is a much smaller species) it is distinguished by
the same characters, except that L. wither si also has a closed discoidal cell. The
closing cross-vein is, however, much nearer to the base of the wing.
REMARKS. Though identical with L. jacksoni in size, this specimen has very much
thicker veins. It is for this reason that it is here regarded as a distinct species.
Liassophlebia sp.
MATERIAL. British Museum (Nat. Hist.), In. 53972, with counterpart. Jackson
CoU.
LOCALITY. Lower Lias : Flatstones, Stonebarrow, Charmouth, Dorset.
DESCRIPTION. A fragment of C, Sc and R of a hind wing as far as the nodus, and
traces of M and Cu. Length from first antenodal to nodus, 23-5 mm.
REMARKS. The two " hypertrophied " antenodals prove that this is a Liasso-
phlebia. The fragment appears to have suffered much from softening in water, for
there are two unusual features difficult to explain as genuine venational elements,
viz., the C is depressed between the two antenodals, and basad of the second ante-
nodal Sc and R seems to be fused. It is inadvisable, therefore, to assign this fragment
to a particular species, though it is likely to belong to one of them.
Family ARCHITEMIDAE Handlirsch
Genus DIASTATOMMITES Handlirsch (1920 : 178)
For status of family and genus, see Tillyard (1925).
Diastatommites liassina (Strickland) ?
(PI. 27, fig. 3)
1840 Aeshna liassina Strickland, p. 301, fig. n.
1856 Diastatomma Hasina (Strickland) Giebel, p. 276.
1906 Diastatomma Hasina (Strickland) : Handlirsch, p. 465.
1925 Diastatommites liassina (Strickland) : Tillyard, p. 23.
MATERIAL. One specimen, British Museum (Nat. Hist.), In. 59375, with counter-
part, from the Lower Lias : Flatstones, Stonebarrow, Charmouth, Dorset. Jackson
Coll.
REMARKS. I refer this small fragment to Diastatommites with considerable hesi-
tation. It appears to be the anal portion of a hind wing. If so, Cu2 is exceptionally
strongly curved. It is to be hoped that more and better material will be discovered
which would help also to clear up the uncertainty regarding the systematic position
of this genus.
Length of the fragment, 22 mm.
Three other tiny fragments of dragonfly wings, In. 49247 (Stonebarrow), In. 53895
FOSSIL INSECTS FROM THE LOWER LIAS OF CHARMOUTH 165
(Black Ven) and In. 59109 (Stonebarrow), may be mentioned to complete the record.
They are unidentifiable.
Order ORTHOPTERA SALTATORIA
Suborder ENSIFERA
Family PROPHALANGOPSIDAE Caudell
Subfamily HAGLINAE Zeuner
Genus PROTOHAGLA nov.
DIAGNOSIS. A Prophalangopsid whose male elytra have almost straight longi-
tudinal veins, except for CuP and lA, which are regularly concave towards the
fore margin.
TYPE SPECIES. Protohagla langi sp.n. (only species).
Protohagla langi sp. n.
(Plate 26)
DIAGNOSIS. Male fore wing broadest at end of basal third. Total length of fore
wing about 67 mm. (61 mm. preserved). Maximum width 26 mm. Pattern of four
dark cross-bands, most pronounced in the anterior portion of wing. Hind wings
appear to exceed fore wings in length.
HOLOTYPE. British Museum (Nat. Hist.), In. 59018, with counterpart. Jackson
Coll., the only specimen.
LOCALITY. Lower Lias : Flatstones, Stonebarrow, Charmouth, Dorset.
PARTS KNOWN. Basal two-thirds of fore wing, distal portion of hind wing.
DESCRIPTION. Fore wing very large, length about 67 mm., of which 61 mm. are
preserved, maximum width 26 mm. In spite of this, the venation is slightly simpler
than in other genera. Four dark cross-bands form a conspicuous colour-pattern of
the fore wing. The tip of the hind wing also appears to have been darkened.
Precostal area well developed and separated from the rest of the fore margin by
a conspicuous long C. The outline of the part resembles that of Tettigonia viridissima
L. Sc extending over about five-sixths of the fore margin ; it has 14 branches.
R separated from Sc by a field of densely-spaced parallel cross-veins, area between
R and Rs similar, but wider, with at least three branches, the first of which is inde-
pendent as in Hagla. Rs with at least four, possibly five, branches.
M separated from R by an area of densely-spaced parallel cross-veins at right
angles to the longitudinal veins, but between Rs and M they are oblique.
M three-branched, as in the female of Hagla, MA being a simple fork, whilst MP
begins with a short oblique stalk, is fused with CuA for some distance, and then free
again. The free portion is almost straight and undivided.
Cu consisting of a forked CuA, an unbranched Cun, and an unbranched CuP, as in
166 FOSSIL INSECTS FROM THE LOWER LIAS OF CHARMOUTH
Hagla. It differs, however, from that of Hagla in that both Cun and CuP branch off
before the junction with MP (in Hagla Cun distad, see Zeuner, 1939, pi. 26, figs. 2, 3).
Cun is slightly concave towards the fore margin, CuP pronouncedly so. The areas
between CuA and Cun, and between Cun and CuP, are thus broadest in the middle.
Of lA, only the basal and distal portions are preserved. It is strongly concave
towards the fore margin near the base and, if a line in the third fifth of the area
between CuP and lA is correctly interpreted as part of the latter vein, slightly bent
towards the fore margin in this zone. This would give it a somewhat wavy run,
though one very much less intense than in Hagla.
The anal area being damaged, 2A, 3A and Ax (if present) cannot be seen.
The hind wing is folded and lies beneath the fore wing. It appears to have exceeded
the latter in length, for its tip is darkened. The traces of veins that can be discerned
call for no comment.
DISCUSSION AND REMARKS. This magnificent fossil is remarkable in several respects.
From the point of view of preservation it is evident that both fore and hind wing
were nipped off simultaneously at the base, and the victim was not then flying.
Considering how detached the movements of the wings of Saltatoria are whilst in
flight, they must have been tightly closed at the time when the insect was being
dismembered. This makes one think of a predator that caught it over the land,
rather than of a dragonfly catching a flying insect over the water. What sort of
predator this may have been is guesswork for the time being ; perhaps it was a
flying reptile.
In general appearance Protohagla must have resembled Tettigonia rather than
Prophalangopsis, or perhaps more still one of the large tropical Phaneropterinae
with irregularly-veined fore wings and long hind wings, such as Zeuneria sp.
The affinities of Protohagla are with Hagla, a well-known genus from the British
Lias. The simple venation of Protohagla might at first sight suggest that the specimen
is a female, but the inverted run of Cun, CuP and lA clearly indicates modifications
in the direction of those observed in the males of Hagla, where the peculiar kink in
these veins foreshadows the way in which the stridulating organ of the later
Prophalangopsidae and Tettigoniidae was to evolve. In the females of Hagla
these veins are unspecialized and convex towards the fore margin.
Protohagla langi is the most primitive Prophalangopsid so far known, though not
the oldest, Notopamphagopsis bolivari Cabrera from Argentina being of Triassic age.
Protohagla is an important phylogenetic link with the Gryllacrididae, of which
Jurassobatea gryllacroides Zeuner from the Upper Jurassic of Solnhofen, Bavaria,
shows several features reminiscent of the fore wing of Protohagla, especially CuP,
i A and 3 A being bent in a similar manner, and the cross-venation being parallel,
not reticulated. The new genus thus connects the Prophalangopsidae more closely
with the Gryllacrididae, the most primitive family of all Saltatoria Ensifera.
The species is dedicated to Dr. W. D. Lang, F.R.S., who first recommended to me
the insects of the British Lias as a subject worthy of study. That he was right in
regarding them as such is borne out by the many descriptions that have appeared
in the last thirty-five years, as well as by Mr. Jackson's discoveries at Charmouth,
with which Dr. Lang is closely connected.
FOSSIL INSECTS FROM THE LOWER LIAS OF CHARMOUTH 167
Order COLEOPTERA
Family CUPEDIDAE
Genus LIASSOCUPES nov.
DIAGNOSIS. Cupedid with rounded pronotum.
TYPE SPECIES. Liassocupes parvus sp. n. (only species).
Liassocupes parvus sp. n.
(PI. 27, fig. 4)
DIAGNOSIS. As for genus.
HOLOTYPE. British Museum (Nat. Hist.), In. 64008. Jackson Coll., the only
specimen.
LOCALITY. Lower Lias : Flatstones, Black Ven, Charmouth, Dorset.
PARTS KNOWN. Elytra, pronotum, head.
DESCRIPTION. The specimen is a well-preserved beetle, but unfortunately the
distal part of the elytra is missing. The elytra must have been about twice as long
as the preserved portion.
Width of beetle across elytra, 2-6 mm., preserved length of elytra, 4-0 mm., pro-
notum, i'Q mm.
The elytra show the dense regular network ol longitudinal ridges with numerous
cross-connections characteristic of the Cupedidae. On the assumption that this
beetle had the same slender shape as Recent Cupes, they should have been long
and parallel-sided. There are eight or nine longitudinal lines in the network, of
which the fourth is distinctly stronger, another Cupedid feature. In life this fossil
must have resembled the Recent Cupes capitatus Fabr. of North America in general
appearance.
The pronotum and head lie directly in front of the elytra, tilted at an angle of
135 degrees to the right, evidence of severe water-logging. The pronotum is almost
circular and shows no spines or other details. It is narrower than the pair of elytra,
and this is again another Cupedid characteristic.
The head is poorly preserved, but its granular surface is discernible, the mandibles
can be recognized and the head is inserted horizontally in front of the pronotum.
No spines or protuberances can be seen, and the antennae are not preserved.
REMARKS. The shape of the pronotum and the exceptionally small size are charac-
teristic. Its sculpture distinguishes this species from the fragment recently described
by Gardiner (1961 : 87) as Metacupes harrisi, from the Rhaetic of Bridgend,
Glamorgan.
Family ?
Genus HOLCOELYTRUM Handlirsch (1906 : 453)
DIAGNOSIS. Beetles with elytra with conspicuous black and white longitudinal
stripes.
TYPE SPECIES. Holcoelytrum giebeli Handlirsch, 1906, by monotypy.
i68 FOSSIL INSECTS FROM THE LOWER LIAS OF CHARMOUTH
DISTRIBUTION. Lower Lias of England.
REMARKS. The two species described by Handlirsch as Holoelytrum giebeli and
Holcoptera schlotheimi are unlikely to belong to different genera. The holotypes of
both are reported to have been incorporated in the British Museum, but only that
of the first species is known. It is unwise, therefore, to use the second as the type
species. The holotype of Holcoelytrum giebeli, though only a fragment of an elytrum,
is recognizable, so that this species defines the genus satisfactorily. It is, incidentally,
by far the more common species. For lack of generic differences, Holcoptera is
therefore here regarded as a synonym of Holcoelytrum, which genus now contains
the two species easily distinguishable as follows :
(1) Elytrum about 12 mm. long, with 5 black stripes . . Holcoelytrum giebeli Hdl.
(2) Elytrum about 5 mm. long, with 4 black stripes . . Holcoelytrum schlotheimi (Hdl.)
Holcoelytrum giebeli Handlirsch
(PI. 27, figs. 6-8)
1845 (Harpalideous Carabidae) Brodie, pp. 101, 124, pi. 10, fig. 2.
1856 Harpalus Schlotheimi Giebel, p. 63. (Partim.)
1906 Holcoelytrum Giebeli Handlirsch, p. 453, pi. 41, fig. 64.
DIAGNOSIS. Elytrum over 10 mm. long, with five black stripes.
DISTRIBUTION. Lower Lias of England.
HOLOTYPE. British Museum (Nat. Hist.), I. 3581. Brodie Coll. Probably from
Apperley, Gloucestershire.
MATERIAL. In addition to the holotype, forty-three specimens from the Flatstones
of the Lower Lias of Charmouth, Jackson Coll., as follows :
(a) Stonebarrow, Charmouth, Dorset
Bodies with both elytra, with counterpart : In. 51002 (PI. 27, figs. 7, 8), 53928,
53937 ; three specimens.
Single elytra, with counterpart : In. 49204, 43981, 49563, 49229, 49570, 49227,
49611, 49228, 53989, 53962, 49239, 53985, 49585, 59129, 59141, 59*34, 59153, 59H7,
64013, 64012, 59145, 49244, 64009 ; twenty-three specimens.
Single elytra without counterpart : In. 64010, 64011, 49149, 53943, 49219, 53974,
49610, 49616 (PI. 27, fig. 6), 59148, 59138 ; ten specimens.
(b) Black Ven, Charmouih, Dorset
Single elytra with counterpart : In. 49209, 59100, 48163, 49621 ; four specimens.
Single elytra without counterpart : In. 49619, 49211, 59393 ; three specimens.
Total, forty-three specimens.
PARTS KNOWN. Elytra, scutellum, tip of abdomen (In. 53928), fragment of leg
(In. 53937).
DESCRIPTION. No complete beetles have so far been found. The three specimens
with pairs of elytra look broader and stouter than they would have appeared in life,
FOSSIL INSECTS FROM THE LOWER LIAS OF CHARMOUTH 169
since they are dorsoventrally compressed, causing the elytra to gape at the distal
end. Size and shape are comparable with a large Tenebrio molitor L., or with Feronia
sp. The scutellum is very small, and the fragments of abdomen and leg present no
features worth mention. In. 51002 shows the underside of the body (on the counter-
part), in addition to the striped elytra (PI. 27, fig. 8).
The elytra are noteworthy for their longitudinal stripes. In the diagnosis they
have been given as black. Alternatively, the unpigmented interstices can be counted.
There are at first sight four of them. The outer (anterior) margin of the elytron
also is represented by a pale line (In. 49616). Counting this as Line i, one finds
that Nos. i and 2 are joined near the apex, continuing as a single line for a short
distance and ending free within the black. Also, Nos. 3 and 5 are similarly joined,
without continuation. They thus enclose No. 4. All are bent towards the shoulder
near the base and towards the apex at the distal end. It is evident that these lines
represent the veins, whilst the black lines are the interstices. The veins represented
are Sc (Line i), R, Rs, M, and Cu, on the interpretation given to the elytral ridges
of the Cupedidae by Zeuner (1933 : 294).
The question arises whether the veins were ridges in the elytra. They could
equally well have been immersed within the elytron. A break between the upper
and the lower cuticles would result in the same structure on the rock surface as
would be produced by ridges.
Some specimens show only three black stripes, the inner and the outer being
absent. In these cases, the white lines are broader than usual. One may be inclined
to regard these as a different species, but In. 53981 demonstrates conclusively that
both variants are merely aspects of the same type of elytron. In the specimen men-
tioned part and counterpart have five and three black stripes respectively. Un-
fortunately, this specimen is much flattened. On the counterpart, the anterior edge
is distinctly bent upwards, as it is in a large number of Recent beetles. It may
thus be presumed that the three-striped aspect is the lower (or underside) of the
elytron. This is confirmed by one of the few specimens that have retained some of
their original curvature. In specimen In. 49611 the surface is distinctly convex,
identifying this as a left elytron. Its colourless lines are very distinct on the coun-
terpart, which is the negative of the upper surface. Since they are wider on the
lower part, they must represent lumina in the elytron. It is highly probable, there-
fore, that these elytra had five ridges corresponding to the major longitudinal veins,
and that the membrane between the veins was pigmented black.
One further detail is supplied by In. 49616, and less clearly by others. There
is a concentration of black along the white lines, whilst the central portions of the
dark stripes are grey. It appears that the latter were thinner than the sides of the
ridges. This is as it should be if the ridges contained tracheal lumina.
Having ascertained the structure of the elytron, it is now possible to describe it
in some detail.
Elytra n-8-13'5 nun. long, and up to 4-0 mm. wide (less originally, since flattened
Post mortem ; 3-5 mm. normal). Black, with four prominent ridges corresponding
to R, Rs, M, Cu. These and the anterior edge (Sc) usually colourless in the fossils
which are split between the upper and lower membranes. Sc and R joined distad,
iyo FOSSIL INSECTS FROM THE LOWER LIAS OF CHARMOUTH
with a short single continuation, Rs and Cu likewise joined distad but not continued.
M inside the area enclosed by Rs and Cu, without touching them. On specimens
showing the veins as white lines, shoulder portion always black. All veins, except
Sc, bent forwards at the base.
REMARKS. Brodie (1845 : 124) regarded this species as " appearing in form
nearest to the Harpalideous Carabidae ". Handlirsch refuted this, and its systematic
position must remain uncertain until a specimen with head and pronotum is found.
Nevertheless, owing to the five " black stripes " separated by four " white lines ",
the species is the most easily recognized among the beetles of the British Lias.
Holcoelytrum schlotheimi (Giebel)
(PL 27, fig. 5)
1845 (Harpalideous Carabidae) Brodie, pp. 101, 124, pi. 6, fig. 28.
1856 Harpalus Schlotheimi Giebel, p. 63.
1906 Holcoptera Schlotheimi Giebel : Handlirsch, p. 453, pi. 41, fig. 63.
DIAGNOSIS. Elytron over 5 mm. long, with four black stripes.
DISTRIBUTION. Lower Lias of England.
HOLOTYPE. British Museum (Nat. Hist.), I. 3582. Brodie Coll., from Binton,
Warwickshire, is supposed to be the holotype. The specimen is labelled as
" Carabidae (Harpalideous), Figd. Brodie, Foss. Ins. pi. 6, f. 28, p. 101, 124, Brodie
Coll." Except in size, however, it does not agree with the figure referred to, being
uniformly black and lacking the stripes. The specimen is marked on the rock itself
" Binton ", and on a label glued to it is written " Carabidae PI. 6, f. 28 " and on
the reverse " Harpalideous Carabidae ". This appears to be in Brodie's own hand-
writing, so that the mistake was made by Brodie himself. The locality given in his
book (p. 101) is Apperley or Brockeridge. Since Apperley is the type locality of
H. giebeli, the type of H. schlotheimi should have come from Brockeridge.
It is clear, therefore, that the specimen marked as the type, I. 3582, is not the
holotype, and that the true holotype was lost long ago, probably in Brodie's time.
For diagnostic characters one has to rely on Brodie's illustration and description,
and a Neotype has been selected from the new material.
NEOTYPE. British Museum (Nat. Hist.), In. 59115, from the Flatstones, Stone-
barrow, Charmouth, Dorset. Jackson Coll. (PI. 27, fig. 5).
OTHER MATERIAL. In. 53990 with counterpart, from the same locality.
PARTS KNOWN. Elytra in pairs, parts of prothorax.
DESCRIPTION. Of the two specimens available, In. 59115 shows the same kind
of black stripes as H. giebeli. There are two in the central part of the elytron, and
one thin one along the hind margin. The latter touches the corresponding stripe of
the other elytron in the position of rest, so that the beetle appears to have five
stripes. The intervening white lines are broader in this species than in H. giebeli,
and all join the pale anterior margin at the apex. Their bases are bent forward as
in the other species. One vein, therefore, is not marked (or atrophied) in H. schlo-
theimi. Most probably R lies close to Sc, the black stripe between them being
missing.
FOSSIL INSECTS FROM THE LOWER LIAS OF CHARMOUTH 171
The dorsal counterpart of the second specimen (In. 53990) is almost uniformly
black, confirming the observations made on the extensive material of H. giebeli ;
but in this specimen the longitudinal ridges are discernible, and there appears to
be a fine punctation present on the surface of the elytron
The scutellum and a portion of the prothorax are preserved in In. 53990, without
providing diagnostic information.
Elytra 5-5 mm. long, 3 mm. wide.
REMARKS. This species is much rarer than H. giebeli. There can be no doubt
that the two specimens described here belong to H. schlotheimi, based on Brodie's
figure (1845, pi. 6, fig. 28).
•
REFERENCES
ABEL, O. 1922. Lebensbilder aus der Tievwelt der Vorzeit. 643 pp., i pi. Jena.
BRODIE, P. B. 1845. A History of the Fossil Insects in the Secondary Rocks of England. 130
pp., ii pis. London.
CONWAY, E. J. 1943. The chemical evolution of the ocean. Proc. R. Irish Acad., London,
48, B : 161-222.
GARDINER, B. G. 1961. New Rhaetic and Liassic Beetles. Palaeontology, London, 4 : 87-
89, 3 figs.
GIEBEL, C. 1856. Fauna der Vorwelt mil steter Beriicksichtigung der lebenden Thieve, 2, i.
511 pp. Leipzig.
HAGEN, H. 1850. In SELYS-LONGCHAMPS, E. de & HAGEN, H. A. Revue des Odonates ou
Libellules d'Europe. Mem. Soc. Sci. Liege, 6 : xxii + 406 pp., n pis.
— 1866. Hemerabiidarum Synopsis synonymica. Entom. Zeit., Stettin, 27:369-462.
HANDLIRSCH, A. 1906-08. Die fossilen Insekten und die Phylogenie der rezenten Formen. 1430
pp., 51 pis. Leipzig.
LEANDER, K. M. 1901. Ubersicht der in der Umgebung von Esbo-Lofo im Meerwasser vorkom-
menden Tiere. Acta Soc. Fauna Flora fenn., Helsingforsiae, 20, 6 : 1-20.
SCHRODER, C. 1920. Handbuch der Entomologie, 3. Jena.
SCHWARZ, A. 1931. Insektenbegrabnis im Meer. Natur u. Mus., Frankfurt, 61 : 453-465.
SILFVENIUS, A. J. 1905. Zur Kenntnis der Trichopterenfauna von Tvarminne. Festschr.
J. A. Palmen, 2, 14 : 1-31. Helsingfors.
STRICKLAND, H. E. 1840. On the Occurrence of a Fossil Dragon-fly in the Lias of Warwick-
shire. Mag. Nat. Hist., London (n.s.) 4 : 301-303, text-figs. 11-13.
TILLYARD, R. J. 1925. The British Liassic Dragonflies (Odonata). 40 pp., 5 pis. British
Museum (Nat. Hist.), London.
TRUSHEIM, F. 1929. Massentod von Insekten. Natur. u. Mus., Frankfurt, 59 : 54-61.
USSING, H. 1918. Insektlivet i og ved Gudenaaens Delta og Randers Fjord. In Johansen,
A. C. Randers Fjords Naturhistorie. K0benhavn.
WENZ, W. 1921. Das Mainzer Becken und seine Randgebiete. 351 pp., 518 figs. Heidelberg.
ZEUNER, F. E. 1933- Die Stammesgeschichte der Kafer. Paldont. Z., Berlin, 15:280-311,
14 figs.
- 1938. Die Insektenfauna des Mainzer Hydrobienkalks. Palaont. Z., Berlin, 20 : 104-159
pis. 13-17.
- 1939- Fossil Orthoptera Ensifera. xiii + 321 pp., 80 pis. British Museum (Nat. Hist.),
London.
— 194°- Saltatoria Ensifera fossilia. Fossilium Catalogus, 1 : Animalia, Pars 90. 108 pp.
The Hague.
— 1958. A new Liassic Dragonfly from Gloucestershire. Palaeontology, London, 1 : 406-
407, pi. 72, fig. 4.
PLATE 24
FIG. i. Petrophlebia anglicanopsis sp. n. Holotype. 111.49573. Xi-85-
FIG. 2. A second specimen doubtfully referred to P. anglicanopsis. In. 59376- X2-2.
Bull. B.M. (TV'.//.) Gcol. 7, 5
PLATE 24
PLATE 25
FIGS, i, 2. Liassophlebia jacksoni sp. n. Holotype (Fig. i) and counterpart (Fig. 2).
In. 53999- X 2.
Bull. B.M. (N.H.) Geol. 7, 5
PLATE 25
-•
PLATE 26
FIGS, i, 2. Protohagla langi gen. et sp. n. Holotype (Fig. i) and counterpart (Fig. 2).
In. 59018. xi -7 and 1-6 respectively.
Bull. B.M. (N.H.) Geol. 7, 5
PLATE 26
PLATE 27
FIG. i. Liassophlebia magnified Tillyaid. In. 64000. X2'4-
FIG. 2. Liassophlebia gigantea sp. n. Holotype. In. 51030. x 2.
FIG. 3. Diastatommites liassina (Strickland) ? In. 59375. X2.
FIG. 4. Liassocupes parvus gen. et sp. n. Holotype. In. 64008. x 10. On the right above
the pair of elytra lie the pronotum and head, cut off in this print.
FIG. 5. Holccelytrum schlotheimi (Giebel). Neotype. In. 59115. X2-g.
FIG. 6. — Holcoelytrum giebeli Handlirsch. In. 49616. X3'i.
FIG. 7. Holcoelytrum giebeli Handlirsch. A pair of elytra. In. 51002. X2-g.
FIG. 8. Holcoelytrum giebeli Handlirsch. Underside of specimen shown in Fig. 7. X2-g.
Bull. B.M. (N.H.) Geol. 7, 5
PLATE 27
2
8
PRINTED IN GREAT BRITAIN BY
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BARTHOLOMEW PRESS, DORKING
THE ENGLISH
CRETACEOUS TURRITELLIDAE
AND MATHILDIDAE (GASTROPODA)
H. L. ABBASS
BULLETIN OF
THE BRITISH MUSEUM (NATURAL HISTORY)
GEOLOGY Vol. 7 No. 6
LONDON : 1962
THE ENGLISH
CRETACEOUS TURRITELLIDAE
AND MATHILDIDAE (GASTROPODA)
BY
HOUSSEIN LOUTFY ABBASS
Department of Geology, Faculty of Science,
Ain Shamo University, Cairo, U.A.R.
Pp. 173-196 ; Plates 28-32 ; 19 Text-figures
BULLETIN OF
THE BRITISH MUSEUM (NATURAL HISTORY)
GEOLOGY Vol. 7 No. 6
LONDON: 1962
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. 7, No. 6 of the Geological
(Palaeontological) series.
Trustees of the British Museum, 1962
PRINTED BY ORDER OF THE TRUSTEES OF
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THE ENGLISH
CRETACEOUS TURRITELLIDAE
AND MATHILDIDAE (GASTROPODA)
By H. L. ABBASS
SYNOPSIS
The paper is a systematic study of the Turritellidae and other Turritella-like gastropods
found in the Cretaceous rocks of England. The number of species recognized is 13, of which
the following are described as new : Turritella (Torquesia) tamra, T. (Torquesia) wagihi, T.
(Torquesia) faizai, T. (Torquesia) hassani, Turritella (s. lat.) ageri, Turritella (s. lat.) sherborni,
Mathilda coxi, M. ahmadi. Formations of Albian age (the Blackdown Greensand and the
Gault) have yielded the greater number of the gastropods described, but some species come
from the Aptian, Cenomanian, and Senonian.
INTRODUCTION
THE work which has led to the present paper was carried out in 1954-6 in the
Department of Geology of the Imperial College of Science, London, in partial fulfil-
ment of the requirements for the degree of Ph.D. of the University of London.
The author has received advice and help from several people, and is especially
grateful to Dr. L. R. Cox, of the British Museum (Natural History), and to Dr.
D. V. Ager, of the Imperial College, in this connection. The material examined
belongs to the collections of the Department of Palaeontology of the British Museum
(Natural History), the Geological Survey of Great Britain, the Sedgwick Museum
(Cambridge), Mr. C. W. Wright, and Dr. J. M. Hancock. The writer tenders his
thanks to these two gentlemen and to the authorities of the institutions mentioned
for the facilities afforded him to carry out the work.
STRATIGRAPHICAL NOTES
Fossil turriculate gastropods occur in several of the English Cretaceous marine
formations. They are, however, known only from a relatively few localities and are
rarely abundant, so that there is no possibility of their being used stratigraphically.
Most of the material described in this paper comes from old museum collections and
in some cases nothing is known of the precise horizons and localities from which the
specimens were collected. There is a probability that in some cases specimens with
similar labels (e.g. " Gault, Folkestone ") may come from a whole range of horizons
within the formation stated.
The marine Neocomian rocks found in Lincolnshire and Yorkshire have yielded no
GEOLOGY 7, 6. 14
I76 ENGLISH CRETACEOUS TURRITELLID AE AND MATHILDIDAE
Cretaceous outcrops are shaded.
FIG. i. Sketch map showing the Cretaceous outcrop in England and Northern Ireland,
with localities marked.
ENGLISH CRETACEOUS TURRITELLID AE AND MATHILDIDAE 177
representatives of the Turritellidae, and the contemporaneous Wealden deposits of
south-eastern England are deltaic and lacustrine sediments with fossil gastropods
belonging to non-marine genera.
Aptian deposition began in the English area with marine transgressions from the
south and north. The isthmus separating the northern and southern basins was
eventually submerged in mid-Aptian times, and throughout the rest of the Creta-
ceous period there was a single basin of deposition in the British area. The Aptian
rocks or Lower Greensand of the Weald and other districts of the English mainland
have not yielded any of the gastropods described in the present study, but the rich
faunas found in the south of the Isle of Wight near Atherfield, where the Lower
Greensand reaches its maximum thickness of 800 ft., include members of both the
Turritellidae and the Mathildidae. The two species here described, Turritella
(Torquesia) tamra sp. nov. and Mathilda coxi sp. nov., both come from the rock
bands known as the Crackers, belonging to the Atherfield Clay series.
The Albian stage is represented in the Wealden district by the upper part of the
Folkestone beds and by the Gault, a stiff clay, the narrow, elliptical outcrop of
which occupies low-lying ground between the Lower Greensand hills and the Chalk
downs. The thickness of the Gault is variable, being about 300 ft. near Eastbourne
and 100 ft. at Folkestone, and at the latter locality its rich fauna includes many
gastropods, most of which, however, are more or less crushed. Two species from
the Gault of Folkestone, Turritella (Torquesia} vibrayeana d'Orbigny and Turritella
(Torquesia} wagihi sp. nov., are described in the present paper, and the first has
also been found in the thin deposit of Gault clay found below the Upper Greensand
at Charmouth, Dorset. The Greensand of Blackdown, Devon, also of Albian age,
is the most prolific source of Cretaceous gastropods in England, and they are silicified
and relatively well preserved. The following species from Blackdown are here
described : Turritella (Torquesia} granulata J. de C. Sowerby, Turritella (Torquesia}
faizai sp. nov., Turritella (s. lat.) ageri sp. nov., Turritella (s. lat.) sherborni sp. nov.,
and Mathilda ahmadi sp. nov. The first occurs in great abundance.
The Cenomanian stage is represented by rocks of greensand facies in south-eastern
Devon and in the Warminster district of Wiltshire. Turritella (Torquesia} granulata
occurs in the Cenomanian of both areas, and a second species, Turritella (Torquesia}
hassani sp. nov., in Devon. Except in these two areas, deposition of calcareous
marl now represented by chalky rocks had begun by the beginning of Cenomanian
times. This change in sedimentation was formerly interpreted as the result of a
gradual sinking of the sea-floor that continued, with slight pauses, for a long period
over almost the whole area that is now the British Isles. The lack of clastic sedi-
ment may, however, have been due to other causes, as the fossils found in the Chalk
do not seem to have been deep-sea forms. The Cenomanian chalky rocks of Kent
and Cambridgeshire have yielded Turritella (Turritella) dibleyi Newton, and those
of Sussex and the Isle of Wight " Turritella " turbinata J. de C. Sowerby, a large
gastropod which belongs most probably to a new genus.
Although rare in the great mass of the post-Cenomanian Chalk, gastropods are
relatively abundant in the rock-bands at the top of the Turonian known at the
Chalk Rock, but no Turritellidae have been found at this horizon. The Senonian
178 ENGLISH CRETACEOUS TURRITELLID AE AND M ATHI LDI D AE
Chalk of Norwich, however, has yielded the species Turritella (Turritelld) unicarinata
(S. Woodward), a form also found in the Upper Chalk of northern Ireland.
TABLE I. — Summary of the Distribution in England of the Species Described
Aptian Albian Cenoman. Turon. Senon.
Turritella (Turritella) dibleyi
,, ,, unicarinata
,, (Torquesia) granulata . . . Devon
,, ,, tamra . . I. o. W. .
,, ,, vibrayeana . . Kent,
Dorset
,, ,, wagihi . . . Kent
,, ,, faizai . . . Devon
hassani
,, (s. lat.) ageri ... . Devon
,, ,, sherborni . . . Devon
Mathilda coxi . . . . . I. o. W. .
,, ahmadi .... . Devon
" Turritella " turbinata
Cenoman.
Kent,
Cambs.
Devon,
Wilts.
Norfolk
Devon
Sussex,
I. o. W.
NOTES ON TAXONOMY AND TERMINOLOGY
Of modern authors who have discussed the morphology of the sheh1 in the Turri-
teUidae and the taxonomic value of its various features, particular mention may
be made of Cossmann (1906, 1912, 1916), Guillaume (1924), Dollfus (1926), Merriam
(1941) and Marwick (1957). Ah1 of these workers have pointed out the significance
of the form of the growth-lines as seen between successive sutures on the spire
whorls, and Marwick has extended the study of the growth-lines to their continua-
tion as seen on the base of the last whorl.
Cossmann, in his review of the family, also took into account the outline and
ornament of the whorls and the degree of acuteness of the spire. He recognized
three genera with numerous subgenera and sections, and he considered Turritella
(Turritella), Turritella (Zaria), Turritella (Haustator), Turritella (Peyrotia), Turritella
(Archimediella) , and Mesalia all to be represented in the Cretaceous, T. (Haustator)
by numerous species. The studies of Guillaume were confined to Tertiary forms.
This author based his classification entirely on the form of the growth-lines as seen
between the sutures, but he did not assign subgeneric names to the groups which
he distinguished. Merriam, in his work on the Upper Cretaceous and Tertiary
Turritellidae of western North America, paid attention, not only to the growth-
lines, but also to the order in which the elements of spiral ornament appear in the
early ontogeny of the shell. He recognized a number of distinct lineages but re-
frained from assigning subgeneric names to them.
Marwick, in his generic revision of the family, considers both the growth-lines and
the ontogeny of the primary elements of spiral ornament to be valuable criteria for
classification, and he also finds that the nature of the protoconch is of help in this
connection. He interprets genera in a more restricted sense than did his pre-
ENGLISH CRETACEOUS TURRITELLID AE AND MATHILDIDAE 179
decessors, recognizing 33 in all, distributed among five subfamilies ; in addition,
he regards nine named taxa as subgenera. The data assembled by Marwick show
that the following genera or subgenera have species of Cretaceous age as their
types : Arcotia Stoliczka (Upper Cretaceous, India), Colposigma Finlay & Marwick
(Danian, New Zealand), Craiginia Stephenson (Cenomanian, North America),
Leptocolpus Finlay & Marwick (Danian, New Zealand), Sechuritella Olsson (Upper
Cretaceous, Peru), and Torquesia Douville (Cretaceous, widespread). The geological
ranges of the various genera are not given in Marwick's paper and it is possible
that he would consider some of those with Tertiary or Recent type-species to be
represented in the Cretaceous.
Owing to the state of preservation of the available material it is seldom possible,
when studying Cretaceous specimens, to observe the protoconch, the development
of the elements of spiral ornament in early ontogeny, or even the growth-lines on the
base of the last whorl. In the present paper, therefore, particular attention is paid
FIGS. 2-6. Terminology of growth-lines in Turritellidae.
Two points of inflection.
2.
3-
4-
5-
6. No point of inflection. Growth-line chord orthocline.
One point of inflection.
No point of inflection. Growth-line chord prosocline.
No point of inflection. Growth-line chord opisthocline.
to the form of the growth-lines between the sutures, and the taxonomic groups of
Turritellidae which are recognized are treated as subgenera of Turritella. The forms
described include the type-species of Torquesia (Turritella granulata ]. de C. Sowerby),
and this subgenus has proved to be suitable for the reception of several other English
species. The other taxa mentioned above as being based on Cretaceous species are
not represented in the material studied.
Two of the Turritella-like species here described have been recognized as repre-
sentatives of the genus Mathilda, the presence of which in the Cretaceous was first
recorded by Cossmann. One of these (M. coxi sp. nov.) is the earliest known
representative of the genus and of the family Mathildidae.
The terminology employed in the present paper is mainly that used by previous
students of the Turritellidae. The less familiar terms are explained by Text-figs.
2-6. The growth-line chord may be defined as a straight line joining the points
where a growth-line meets successive sutures.
Most of the new species are named after geologists and others who have helped
the author during the course of the work.
i8o ENGLISH CRETACEOUS TURRITELLI D AE AND MATHILDIDAE
SYSTEMATIC DESCRIPTIONS
Class GASTROPODA
Subclass PROSOBRANCHIA
Superfamily CERITHIACEA Fleming, 1822
Family TURRITELLIDAE Woodward, 1851
Genus TURRITELLA Lamarck, 1799 : 74
TYPE SPECIES. — Turbo terebra Linnaeus, 1758 (by monotypy).
Subgenus TURRITELLA s. str.
SUBGENERIC CHARACTERS. — Shell medium-sized to large, slender ; whorls evenly
convex, ornamented with spiral cords and threads ; growth-lines between sutures
forming a simple sinus of no great depth.
Turritella (Turritella) dibleyi Newton
(PL 29, figs. 7, 8 ; Text-fig. 7)
1918. Turritella dibleyi Newton, p. 97, pi. 10, figs, i, 2.
HOLOTYPE. — In the British Museum (Natural History), reg. no. G. 29362.
DESCRIPTION. — The shell is moderately large and slender. The apical whorls, up
to a diameter of about 9 mm., are missing in the holotype. The whorl outline is
rather strongly convex, but in the holotype the convexity of the earlier preserved
whorls has apparently been reduced by pressure ; the suture is moderately deep.
About 30 spiral threads are present on the spire whorls. They are crossed by collabral
threads which tend to become prominent at variable intervals, almost forming
varices, and become generally stronger on the later whorls, so that the appearance
of collabral ornament is developed in places. The spiral threads are separated by
interspaces the width of which is almost equal to that of the threads. The base
and aperture are damaged in the holotype. The growth-lines show a broad, shallow
sinus with no points of inflection, and the growth-line chord is prosocline (Text-
fig. 7)-
Measurements of the Holotype :
Height (as now preserved) • • • 75 mm.
Spire angle ...... 13°
Height of penultimate whorl . . . 12 mm.
Diameter of penultimate whorl . . 19 mm.
DISCUSSION. — This species is included in the subgenus Turritella s. str. as its
growth-lines have a simple, shallow sinus and a strong prosocline trend, as in the
type-species of the genus. It differs from T. (T.) unicarinata, described below, in
the presence of a greater number of spirals and of frequent collabral rugae.
OCCURRENCE. The holotype came from the Chalk Marl (Cenomanian) of Margett's
Pit, Burham, Kent. The species has also been reported from the same formation at
ENGLISH CRETACEOUS TURRITELLI D AE AND MATHILDIDAE 181
Folkestone and from the Burwell Rock (Cenomanian) at Burwell and Reach, in
Cambridgeshire.
8
10
FIGS. 7
\— \—
'-19. Diagrams of growth-lines of the species described in this paper.
7. Turritella (Turritella) dibleyi Newton.
7. Turritella (Turritella) dibleyi Newton.
Turritella (Turritella} unicarinata (S. Woodward).
9. Turritella (Torquesia) granulata J. de C. Sowerby.
10. Turritella (Torquesia} tamra sp. nov.
11. Turritella (Torquesia} vibrayeana d'Orbigny.
12. Turritella (Torquesia} wagihi sp. nov.
13. Turritella (Torquesia} faizai sp. nov.
14. Turritella (Torquesia} hassani sp. nov.
15. Turritella ageri sp. nov.
1 6. Turritella sherborni sp. nov.
17. Mathilda coxi sp. nov.
1 8. Mathilda ahmadi sp. nov.
19. " Turritella " turbinata J. de C. Sowerby.
GEOLOGY 7, 6.
i8z ENGLISH CRETACEOUS TURRITELLID AE AND MATHILDIDAE
Turritella (Turritella) unicarinata (S. Woodward)
(PI. 28, figs. 3-5 ; Text-fig. 8)
1833. Cerithium unicarinatum S. Woodward, p. 49, pi. 6, fig. 21.
1854. Nerinaea unicarinata (Woodward) : Morris, p. 264.
1865. Turritella unicarinata (Woodward) : Tate, p. 37, pi. 3, fig. 7.
HOLOTYPE. Not traced.
DESCRIPTION. The shell is moderately large and slender. The earlier whorls,
up to a diameter of 8 mm., are missing in the specimens studied. The whorls which
are preserved are feebly and almost symmetrically convex in outline and the suture
is moderately deep. The later whorls bear about 30 very finely beaded spiral
threads, separated by equal interspaces nearly equal in width to the threads, but
becoming slightly wider than them towards the anterior suture. Each interspace
is usually occupied by one secondary spiral. The base is slightly convex and has
an angular periphery. It is crossed by spirals of the same nature as those on the
whorl-side. The growth-lines have a broad sinus without points of inflection and
the growth-line chord is orthocline (Text-fig. 8). A furrow on the internal mould
appears to be the impression of a depressed median spiral fold on the interior of the
whorls. The aperture is subquadrate.
Measurements of a Typical Specimen (G.S.M. no. 28256) :
Height (as now preserved) . . . . . 51 mm.
Spire angle ........ 13°
Height of penultimate whorl . . . . . 12 mm.
Diameter of penultimate whorl . . . . . 16-5 mm.
DISCUSSION. This species has been referred to such different genera as Cerithium,
Nerinea and Turritella. The absence of an anterior canal or notch excludes it from
the first two genera and it also differs from Nerinea in the absence of internal folds
(apart from the single one mentioned) and of a juxta-sutural sinus band. The
entire aperture and other characters show that it belongs to Turritella, and it is
included in Turritella s. str. on account of the simple arcuate form of its growth
lines.
OCCURRENCE. The holotype was found in the Upper Chalk of Norwich, where
internal moulds are frequent. The species also occurs in the Upper Chalk of Northern
Ireland, where it is common in the White Limestone (Campanian) at Lisburn and
rare in the Spongarian Zone (Santonian) at Woodburn.
Subgenus TORQUESIA Douville, 1929:55
TYPE SPECIES. Turritella granulata J. de C. Sowerby (designated by Inter-
national Commission on Zoological Nomenclature, Opinion 493, 1957).
SUBGENERIC CHARACTERS. Shell medium-sized, more or less slender ; whorls flat
to moderately convex, ornamented with beaded spiral cords ; growth-lines forming
ENGLISH CRETACEOUS TU RRITELLI D AE AND MATHILDIDAE 183
a rather deep sinus with its vertex nearly at mid- whorl and with points of inflection
above and below it ; growth-line chord orthocline to feebly prosocline.
Turritella (Torquesia) granulata J. de C. Sowerby
(PL 30, figs. 10-14 ; Text-fig. 9)
1811. Cerithium turritellatum Parkinson, p. 71 (non Lamarck).
1816. Turritella sp. : Smith, p. 12, " Green Sand " pi., fig. 5.
1827. Turritella granulata ]. de C. Sowerby, p. 125, pi. 565, fig. I.
1840. Turritella granulata Sowerby : Geinitz, p. 44, pi. 15, figs. 7-11.
1842. Turritella granulata Sowerby : d'Orbigny, p. 46, pi. 153, figs. 5-7.
1843. Turritella granulata Sowerby : Geinitz, p. 10, pi. i, fig. 18.
1845. Turritella granulata Sowerby : Geinitz, p. 325, pi. 14, figs. 9, 10.
1849. Turritella granulata Sowerby : Brown, p. 70, pi. 38, fig. 18.
1850. Turritella granulatoides d'Orbigny, p. 190.
1868. Turritella granulata Sowerby : Briart & Cornet, p. 29, pi. 3, figs. 43, 44.
1875. Turritella granulata Sowerby : Geinitz, p. 239, pi. 54, figs. 3, 4.
1920. Turritella (Haustator) granulata Sowerby : Roman & Mazeran, p. 44, pi. 5, fig. 21.
1920. Turritella (Haustator} granulatoides d'Orbigny : Roman & Mazeran, p. 44, pi. 5, figs.
17-20.
HOLOTYPE. In the British Museum (Natural History), reg. no. 43667.
DESCRIPTION. The shell is of medium size and moderately slender. The proto-
conch is not preserved in an uneroded state in the specimens studied. The whorl
outline is feebly convex and the whorls are relatively high. The suture is at first
superficial but becomes progressively more furrowed between the later whorls,
which tend to become disjunct in some specimens. On the earliest whorls of which
the ornament has been observed four beaded spiral cords are present in all variations
of this species, and their interspaces soon become occupied by one or more spiral
threads which are finely beaded. Some of these spiral threads increase in pro-
minence during growth so that on the later whorls they become as strong as the
primary cords, and, as a result, there may be as many as 7-8 principal spirals on the
last whorl. The most posterior spiral cord is stronger than the others and is
separated from them by a relatively broad spiral groove ; this feature is diagnostic
of the species. On the later whorls, especially the last, the beads on the cords are
located along conspicuous growth-lines, and those on the most posterior cord produce
a feeble undulation of the suture. The base is convex and is ornamented with
spiral cords crossed by growth-rugae. The aperture is oval, with a rounded anterior
margin. The thin inner lip is reflected on the columella. The growth-lines have a
deep U-shaped sinus and two points of inflection, one near each suture ; the growth-
line chord is slightly prosocline (Text-fig. 9) .
Measurements of Holotype :
Height ...... 46 mm.
Spire angle ...... 18°
Height of penultimate whorl ... 8 mm.
Diameter of penultimate whorl . . 10 mm.
VARIABILITY. This species is rather variable. One variant is similar to the
184 ENGLISH CRETACEOUS TURRITELLI D AE AND MATHILDIDAE
holotype except that its ornament consists even on the later whorls of only four
principal cords with very conspicuous beads. It would appear that additional
shelly matter was added during growth to the primary spirals instead of to secondary
spirals in their interspaces. (See PI. 30, fig. 14.)
DISCUSSION. The specimens figured by d'Orbigny (1842) as T. granulata had
five spiral cords (not four, as stated by Roman & Mazeran, 1920), and subsequently
d'Orbigny considered them to belong to a different species, T. granulatoides, dis-
tinguished from the true T. granulata by the smaller number of its spiral cords and
by the deeper sinus of its growth-lines. The present writer has found that specimens
agreeing with both forms occur in association at Blackdown, while Roman &
Mazeran mention that they are found together in the Turonian of the Uchaux
Basin (France). It thus seems reasonable to re-unite them as one species.
Cossmann (1912) referred T. granulata to Montfort's subgenus Haustator (type-
species Turritella imbricataria Lamarck, Eocene). He included in this subgenus a
great number of turritellids with growth-lines of the same type as those of the type-
species, namely, with a deep sinus near the middle of the whorl and points of inflec-
tion above and below it. He admitted, however, that forms differing considerably
from the type-species in whorl outline and ornament were thereby brought together.
Roman & Mazeran (1920) also referred T. granulata to Haustator. DouviUe (1929),
however, was of the opinion that Cossmann had interpreted Haustator too widely
and considered that the group of Cretaceous species with rather similar growth-
lines but commonly with a beaded cord near the posterior suture should constitute
a new subgenus Torquesia. He cited as type-species of this new taxon " T. granulosa
de Blackdown ", which the International Commission on Zoological Nomenclature,
following a petition by Dr. J. Marwick, has decided was an unintentional error for
Turritella granulata. Marwick (1957 : 160) rather doubts the taxonomic importance
of beaded spiral cords, but nevertheless considers that Torquesia is a taxon in which
many Cretaceous species may usefully be included. The growth-lines are more
symmetrical and the upper point of inflection is more marked than in the type-
species of Haustator as figured by Marwick (1957 : 146, text-fig. 14), and their general
trend is less pronouncedly prosocline. Torquesia is, therefore, here accepted as a
subgenus in which several English Cretaceous species can be included.
OCCURRENCE. T. granulata is very abundant in the Albian Greensand of its type-
locality, Blackdown, Devon, although specimens retaining the initial whorls and
showing the whole ornament clearly are very rare. Specimens from the Ceno-
manian of Warminster, Wilts., and of Devon (falcatus Zone) have also been examined.
A queried record of the occurrence of the species in the Lower Greensand of Ingolds-
thorpe, Norfolk (Forbes, 1845) is to be rejected.
T. granulata has been recorded from the Cenomanian of various parts of the
European Continent, including the Elbe valley district of Saxony, where it occurs in
both the Lower Quadersandstein and the Lower Planer (Geinitz, 1874), Bohemia
(Reuss, 1845), Bavaria (Sohle, 1896), the Baltic coastlands in glacial erratics
(Noetling, 1885), and Belgium, where it occurs in the so-called meule de Bracquegnies.
D'Orbigny (1842) and Roman & Mazeran (1920) have recorded it from the Turonian
of Uchaux in France.
ENGLISH CRETACEOUS TURRITELLID AE AND MATHILDIDAE 185
Turritella (Torquesia) tamra sp. nov.
(PI. 28, fig. i ; Text-fig. 10)
HOLOTYPE. In the Sedgwick Museum (Cambridge), reg. no. B. 27329.
DESCRIPTION. The shell is small and slender. The actual protoconch is not
preserved in the specimens studied and the earlier whorls are worn. The whorls
are moderately high, with a slightly convex outline, and they are separated by a
well-impressed suture. Four beaded primary cords are present when the whorl
diameter is 2 mm. and on the later whorls the number of principal spirals has in-
creased to about six, one of which is close to the anterior suture. These are separated
by considerably wider interspaces occupied by from one to three spiral threads,
some not very much weaker than the principal spirals. The beads on the spirals
are weak and some are elongated in a spiral direction with the spaces between them
varying in width. The base is convex and is bordered by the primary spiral which
is seen just above the suture on the later spire whorls ; below this are three more
primaries, decreasing in strength inwards, with fine secondary threads between them.
The aperture is not preserved. The growth-lines have a deep sinus the vertex of
which lies between the second and the third primary cord from the posterior suture,
that is, well above the middle of the whorl ; there are two points of inflection. The
growth-line chord is very slightly opisthocline (Text-fig. 10).
Measurements of the Holotype :
Height ...... 26 mm.
Spire angle ...... 14°
Height of penultimate whorl . . . 3-5 mm.
Diameter of penultimate whorl . . 5 mm.
DISCUSSION. Several other specimens of this species are preserved in the small
piece of rock which contains the holotype, but only two (including one which has
been registered separately as B. 27330) show any diagnostic characters.
This species is included in the subgenus Torquesia because of its growth-line
pattern and of its ornament, which bears a general resemblance to that of T. granu-
lata. It differs from that species, of which it is possibly the ancestor, in its smaller
size, more convex whorl outline, weaker spirals, and fewer and weaker beads.
OCCURRENCE. Aptian, Lower Greensand, Atherfield Clay Series (bed known as
the " Crackers "), Atherfield, Isle of Wight.
Turritella (Torquesia} vibrayeana d'Orbigny
(PI. 31, figs. 17-22 ; Text-fig, n)
1842. Turritella vibrayeana d'Orbigny, p. 37, pi. 151, figs. 10-12.
1860. Turritella vibrayeana d'Orbigny : Mackie, p. 324, fig. 34.
1862. Turritella vibrayeana d'Orbigny : Pictet & Campiche, p. 315, pi. 72, figs. 5-7.
1896. Turritella vibrayeana d'Orbigny : Cossmann, p. 255, pi. i, figs. 24-26.
1903. Turritella vibrayeana d'Orbigny : Doncieux, p. 301, pi. i, fig. i.
1954. Turritella cf. vibrayeana d'Orbigny : Gortani, p. 75, pi. 16, figs, za, b.
SYNTYPES. Nine specimens in the d'Orbigny Collection in the Museum National
d'Histoire Naturelle, Paris, reg. no. 5847.
186 ENGLISH CRETACEOUS TURRITELLI D AE AND MATHILDIDAE
DESCRIPTION. The shell is of small to medium size and very slender, with rela-
tively high whorls. The whorl sides are usually flat but may be feebly convex ;
the suture is superficial or slightly furrowed. The initial whorls have a tricostate
ornament. The three primary spiral cords may persist on the later whorls without
the addition of other strong spirals (PL 31, fig. 19), but usually a fourth principal
cord is added near the posterior suture (PI. 31, fig. 20), while a fifth may appear in
the most posterior interspace (PL 31, fig. 18) and a sixth in the most anterior inter-
space (PL 31, fig. 22). The spiral cords are beaded, some rather coarsely, others
faintly. Some of the interspaces are almost equal to the primary cords in width,
but most of them are wider, sometimes as much as five to six times as wide (PL
31, fig. 17). Secondary threads, variable in number, occupy these interspaces, and
usually one or two are more prominent than the rest. The base is slightly convex,
with spiral cords and threads crossed by growth-lines. The aperture is oval and
evenly rounded anteriorly. The growth-lines are of about the same strength as the
secondary spiral threads. They show a deep sinus with its vertex near the middle
of the whorl and two points of inflection, one near each suture. The growth-line
chord is almost orthocline (Text-fig, n).
Measurements of a Typical Specimen (B.M., no. 0.73791) :
Height ......... 40 mm.
Spire angle ........ 10°
Height of penultimate whorl ..... 3-5 mm.
Diameter of penultimate whorl ..... 6 mm.
DISCUSSION. Tunitella vibrayeana has been described by previous authors as
having four primary beaded spiral cords alternating with secondary ones. Actually,
it is a highly variable species. At one extreme of the range of variation are forms
retaining on the later whorls the same number of primary spiral cords as are found
on the initial ones, and at the other extreme are specimens in which as many as
three additional principal cords have been added. There appears to be no justifica-
tion for separating shells with additional spiral cords on the later whorls as distinct
species or subspecies, since all specimens show the same growth-line pattern and
the ornament of the initial whorls is always tricostate. The variation in the number
of principal cords may be due to unimportant environmental or physiological
factors, although its causes may be merely genetic.
The first reference in the literature to specimens belonging to this species was
when Michelin (1838 : 99) referred a Tunitella from the Gault of the Aube Depart-
ment of France to Tunitella rigida J. de C. Sowerby (1833, pi. 38, fig. 19), a species
from Gosau, in Austria. T. rigida has a prominent posterior tuberculate spiral cord
on which the tubercles are markedly elongated and extend to the suture, and this
feature is not found in specimens from the Gault. In view of these differences
d'Orbigny (1842) erected his new species T. vibrayeana for the latter. The figures
of Pictet & Campiche, illustrating specimens from Switzerland, agree with our
description.
Doncieux (1903), when recording specimens from the Gault of the eastern Corbieres,
in the south of France, stated that they differed from the holotype of d'Orbigny in
ENGLISH CRETACEOUS TURRITELLID AE AND MATHILDIDAE 187
the inequality of the spiral cords (the most anterior of these being the strongest)
and also of the secondary spirals. His figures, however, do not show these features.
Cossmann (1912) included T. vibrayeana in the subgenus Haustator. The present
writer agrees that relationship with Turritella granulata is indicated by the growth-
line pattern and by the general character of the ornament. T. vibrayeana is, there-
fore, now included in the subgenus Torquesia. Merriam (1941), in discussing his
" first stock " of T. tolenasensis Merriam, referred to T. vibrayeana as showing
similarities to some American species.
OCCURRENCE. All the specimens examined in the preparation of this paper are
from the Gault of Folkestone, Kent, except one (0.49825) which was collected by
Dr. W. D. Lang from the Lower Gault of Fairy Dell, Stonebarrow, Charmouth,
Dorset. T. vibrayeana has been found in the Albian of France at several localities
in the Aube Department and the eastern Corbieres, and it has also been recorded
from Switzerland, Italy and Russia.
Turritella (Torquesia) wagihi, sp. nov.
(PL 29, fig. 6 ; Text-fig. 12)
HOLOTYPE. In the Geological Survey Museum, reg. no. 93745.
DESCRIPTION. The shell is of medium size and moderately slender. The whorls
are feebly convex, with a weak carina close to the anterior suture, giving them a sub-
imbricate appearance. The protoconch is not preserved. The earliest whorls seen
are tricostate, with primary cords lying at the anterior quarter, the middle, and the
posterior quarter respectively. The number of principal spiral cords remains the
same on the later whorls, but they have moved a little further forward so that the
space behind the most posterior cord is greater than that in front of the most anterior
one. These spirals show a marked difference in strength and in the number of
beads on them. Thus, while the beads are relatively strong and wide apart on the
most posterior cord, those on the most anterior one are more numerous, closer, and
weaker. The primary interspaces are three to four times as wide as the primary
spiral cords, and each is occupied by two secondary spiral cords and four or five
spiral threads. The aperture and base are not preserved in the holotype. The
growth-lines have a broad and rather shallow sinus with its vertex at about the
middle of the whorl and two points of inflection ; the growth-line chord is slightly
opisthocline (Text-fig. 12).
Measurements of the Holotype :
Height ...... 49 mm.
Spire angle ...... 14°
Height of penultimate whorl ... 5 mm.
Diameter of penultimate whorl . . 10 mm.
DISCUSSION. The holotype, the only specimen seen, is crushed and its spire angle
may be greater than was originally the case. This species bears some resemblance
to Turritella vibrayeana and its variations, but can be distinguished by the shape of
the shell, the lower whorls, the more convex whorl outline, the peculiar difference
i88 ENGLISH CRETACEOUS TU RRITELLI D AE AND MATHILDIDAE
in the granulation of the primary spiral cords and the development of the spiral
ornament during the ontogeny of the shell. It differs from T. marticensis Matheron
(1843 : 240, pi. 39, fig. 16) in the presence of the anterior carination and in the
nature of the ornament. The growth-lines are not indicated clearly enough in
Matheron's figure to allow comparison to be made with those of the present species.
OCCURRENCE. Albian, Gault, Folkestone, Kent.
Turritella (Torquesia) faizai sp. nov.
(PI. 30, fig. 15 ; Text-fig. 13)
HOLOTYPE. In the Sedgwick Museum (Cambridge), reg. no. 6.44621.
DESCRIPTION. The shell is of medium size and moderately slender. The earlier
whorls have a slightly convex outline with a feeble anterior carination, but the later
whorls become concave mesially, with anterior and posterior convexities. The
suture is moderately deep. The protoconch is broken off in the holotype. The
earliest preserved whorls are tricostate, the most anterior spiral cord coinciding
with the carination. On later whorls a secondary spiral cord is developed behind
each primary spiral, and these secondaries increase in strength rapidly until they
become as strong as the primaries, so that the total number of principal spiral
cords is increased to six. The cords are finely beaded and are separated by wider
interspaces most of which are occupied by one secondary and two tertiary threads.
The most anterior primary cord is separated from the adjacent suture by an inter-
space wider than that between the posterior primary and the corresponding suture.
A secondary spiral which develops in this latter interspace becomes as strong as the
most posterior primary on the later whorls and fuses with it to form a strong astragal1
which affects the whorl outline. The base is almost flat, with a rounded edge, and
bears concentric cords which alternate in strength. The aperture is broken away
but was evidently subquadrangular. The growth-lines have a deep, broad sinus
and points of inflection coinciding with the anterior and posterior convexities. The
growth-line chord is almost orthocline (Text-fig. 13).
Measurements of the Holotype :
Height 53 mm.
Spire angle ...... 16°
Height of penultimate whorl ... 8 mm.
Diameter of penultimate whorl . . 15 mm.
DISCUSSION. This species is distinguished from Turritella (Torquesia) hassani,
described below, by the relative weakness of the astragal on the posterior part of
the whorls and by the anterior convexity of the whorls. It also has fewer secondary
threads between the principal spiral cords.
OCCURRENCE. Albian of Devon ; the holotype came from the Greensand of
Blackdown and specimens have also been found in the Upper Greensand of Peak
Hill (west of Sidmouth), Seaton and Beer.
1 Astragal. — A moderately broad, parallel-sided elevation with a rounded profile (architectural
term) .
ENGLISH CRETACEOUS TURRITELLID AE AND MATHILDIDAE 189
Turritella (Torquesia) hassani sp. nov.
(PL 28, fig. 2 ; PI. 29, fig. 9 ; Text-fig. 14)
HOLOTYPE. In the British Museum (Natural History), reg. no. 0.49907.
DESCRIPTION. The shell, not known complete, is of medium size, slender, and
nerineiform. The whorl outline is flat or slightly concave and the sutures are
superficial. A rounded posterior astragal, occupying about one-fifth of the height
of the whorl, is developed just below the suture. Eight or nine very faintly beaded
primary spiral cords are distributed equally over the anterior four-fifths of the
whorl ; the interspaces are two to three times as wide as the cords. These inter-
spaces are occupied by four to seven fine spiral threads of which the middle one is
sometimes stronger than the others ; similar spiral threads cover the rounded
posterior astragal. The base and aperture are not preserved in the available speci-
mens. The growth-lines have a deep sinus with its vertex at the middle of the
whorl and two points of inflection. The growth-line chord is slightly prosocline
(Text-fig. 14).
Measurements of the Holotype :
Height (four mid-spire whorls) . . . 40-5 mm.
Spire angle ....... 42^°
Height of last preserved whorl ... 9 mm.
Diameter of last preserved whorl . . . 14 mm.
DISCUSSION. This species is founded on three imperfect specimens now in the
British Museum (Natural History), two collected by Mr. T. F. Grimsdale, the third
by Mr. C. W. Wright. It differs from the other turritellids described in this work
in having concave whorls with a rounded posterior astragal crossed by spiral threads
and in other details of ornament. The growth-line pattern is similar to that of
Turritella {Torquesia} granulata J. de C. Sowerby, and this permits its inclusion in
the same subgenus as that species.
OCCURRENCE. Cenomanian Limestone (Bed B of Jukes-Browne), " near Seaton,
Devon " (type-locality). Same bed, between Havencliff and Dowlands, east of
Seaton, Devon (ex C. W. Wright Collection).
Subgenus novum ?
In the two species described next the growth-lines have a broad sinus with its vertex
at about mid-whorl and an inflection point close to the lower suture ; the growth-
line chord is almost orthocline. The whorls are feebly convex and the ornament is
of beaded spiral cords. Of the turritellid growth-lines figured by Marwick (1957 :
146), those of Pareora (fig. 25) are most like those of the two new species, but the
sinus is deeper and its vertex is higher on the whorl. In Sigmesalia (Marwick's
fig. 31) the growth-lines rather resemble those of these species, but the point of
inflection coincides with the lower suture. In other characters there is little re-
semblance between the forms now described and either Pareora or Sigmesalia.
It is probable that they belong to a new subgenus, but since neither their apertures
igo ENGLISH CRETACEOUS TU RR ITELLI D AE AND MATHILDIDAE
nor their initial whorls can be studied this could not at present be denned satis-
factorily.
Turritella ageri sp. nov.
(PI. 32, fig. 25 ; Text-fig. 15)
HOLOTYPE. In the British Museum (Natural History), reg. no. 0.74106.
DESCRIPTION. The shell is moderately slender and of medium size. The whorl
outline is feebly and symmetrically convex, the suture very slightly impressed.
The initial whorls, up to a diameter of 3 mm., are not preserved in the specimens
studied. The whorls now preserved have 6-8 coarsely beaded spiral cords which
increase in strength during growth. On the last whorl the width of the interspaces
is equal to or slightly exceeds that of the cords. Each interspace is occupied by
three or four spiral threads, the middle one of which may be more prominent than
the others and ultimately become as strong as the primary cords. The beads are
coarse, more or less rounded, and closely spaced. The base is slightly convex, with
a rounded periphery, and is crossed by a number of spiral threads alternating in
strength. The aperture is not complete in the holotype, but it can be inferred
from the outline of the last whorl that it was more or less elliptical. The inner lip
is reflected over the short and thick columella. The growth-lines have a broad,
moderately deep sinus, the vertex of which lies on the third spiral cord from the
posterior suture, that is, at about the posterior two-fifths of the height of the whorl.
They straighten out to a point of inflection on the most anterior spiral cord. The
growth-line chord is almost orthocline (Text-fig. 15).
Measurements of the Holotype :
Height ...... 47 mm.
Spire angle ...... 16°
Height of penultimate whorl ... 8 mm.
Diameter of penultimate whorl . . 14 mm.
DISCUSSION. This new species can be easily distinguished from similar forms,
such as Turritella (Torquesid) granulata J. de C. Sowerby, by the absence of a very
strong posterior spiral cord, by the prominence of the spirals on the last whorl, by
its very coarse and rounded beads, and by the growth-line pattern.
OCCURRENCE. Albian, Greensand, Blackdown, Devon.
Turritella sherborni sp. nov.
(PL 30, fig. 16 ; Text-fig. 16)
HOLOTYPE. In the British Museum (Natural History), reg. no. 0.74107.
DESCRIPTION. The shell is of medium size, with whorls which are subimbricate
owing to the presence of a weak carina which coincides with the second spiral cord
from the anterior suture ; above this carina the whorl surface is almost flat. The
sutures are slightly impressed. The initial whorls, up to a diameter of 4 mm., are
not preserved in the holotype. On the whorls which are still seen there are seven
ENGLISH CRETACEOUS TURRITELLID AE AND MATHILDIDAE 191
equal spiral cords ornamented with small, spirally elongated beads separated by
constrictions which are nearly equal in width to the beads. The interspaces between
the spiral cords are nearly double the width of the latter and are occupied by spiral
threads, one or two of which may be stronger than the rest. The spiral threads
show faint constrictions corresponding to those of the primary spiral cords. The
base is rather obscured by adherent matrix in the holotype, but has a rounded
periphery below which its surface is slightly convex. The aperture has an evenly
rounded anterior margin and its height is about equal to its width. The growth-
lines have a broad and moderately deep sinus the vertex of which lies just above
the middle of the height of the whorl. They straighten out to a point of inflection
just above the lower suture. The growth-line chord is almost orthocline (Text-
fig. 16).
Measurements of the Holotype :
Height ...... 30 mm.
Spire angle . . . . . . 16°
Height of penultimate whorl ... 6 mm.
Diameter of penultimate whorl . . 10 mm.
DISCUSSION. This new species much resembles Turritella (Torquesia] granulata
J. de C. Sowerby, but differs in the equality of its spiral cords, in the presence of
the anterior carination, and in the nature of the growth-lines. It shows a great
similarity to T. cenomanensis d'Orbigny, as figured by Gueranger (1867, pi. 9, figs.
10, n), but differs in having anteriorly carinate whorls, and probably also in the
nature of the spiral cords, which are not clearly shown in Gueranger's figures.
D'Orbigny (1850, p. 148) referred T. granulata Geinitz (1840, pi. 15, figs. 7-11, non
Sowerby) to T. cenomanensis, but Geinitz (1874 : 161) later included pi. 15, fig. 9
in the synonymy of T. multistriata Reuss. T. cenomanensis was included by Coss-
mann in the subgenus Haustator, but its growth-lines are unlike those of the type-
species of that subgenus.
The new species T. sherborni differs from T. ageri, described above, in its anterior
carination and finer granulation.
OCCURRENCE. Albian, Greensand, Blackdown, Devon.
Family MATHILDIDAE Cossmann, 1912
Genus MATHILDA Semper, 1865 : 330
TYPE SPECIES. Turbo quadricarinatus Brocchi, 1814 (by subsequent designation,
de Boury, 1883).
Mathilda coxi sp. nov.
(PI. 32, fig. 23 ; Text-fig. 17)
HOLOTYPE. In the Sedgwick Museum (Cambridge), reg. no. 6.27336.
DESCRIPTION. The shell is of small-medium size and broadly turriculate. Details
of the protoconch have been a little obscured by erosion. Although not conspic-
IQ2 ENGLISH CRETACEOUS TURRITELLID AE AND MATHILDIDAE
uously heterostrophic, it exhibits a distinct discontinuity of coiling ; its first
complete whorl is an open one with a gap in the middle, and the actual nucleus
seems to be partly obscured by this whorl. The first two visible whorls are smooth
and rounded in outline, while the third develops a median carina. On the next
succeeding whorls a second carina makes its appearance just in front of the posterior
suture, but the original carina remains the more prominent. The concavity between
these carinae, which come respectively to occupy positions at about the lower third
and slightly above the upper third of the whorl, is reduced on the later whorls,
which are biangular with an almost flat outer face. The suture is deep. The outer
face of the later whorls is occupied by two to three spiral cords of primary strength,
separated by interspaces which are of the same width or slightly narrower. Each
interspace is occupied by one or by two secondary spirals. The area between each
carina and the adjacent suture is similarly ornamented, but the spirals on the
anterior area are the most prominent. On the last whorl two additional spiral
cords of primary strength and below them a third carina become visible just above
the suture. The periphery of the convex base is formed by a spiral keel continuing
this third carina ; below it is a further carina, and the remainder of the base bears
concentric threads of two orders of strength. Where uneroded, the whole surface
of the shell bears a delicate ornament of closely and regularly spaced collabral
threads. The aperture is oval, with the inner lip slightly reflected. The growth-
lines between the sutures form a broad arch, the chord of which is almost ortho-
cline (Text-fig. 17).
Measurements of the Holotype :
Height . . . . . . 21 mm.
Spire angle ...... 30°
Height of penultimate whorl ... 5 mm.
Diameter of penultimate whorl . . 10 mm.
DISCUSSION. The protoconch of the holotype of this species is not so obviously
heterostrophic as in some of the figures of Tertiary species of Mathilda published
by Semper (1865) and de Boury (1883), showing specimens in which the axis of
coiling of the initial whorls is at right angles to that of the shell axis. In a specimen
from the Miocene of southern France in the Wrigley Collection in the British Museum
(Natural History), identified as belonging to the var. semperi Tournouer of the
type-species of the genus, M. quadricarinata (Brocchi), the protoconch is, however,
very similar to that of the species now described, and in some of the Paris Basin
Eocene shells figured by de Boury (1883, pi. 5, figs. 70, qa, loa, na) it also seems
to be of a comparable type. In its general form and ornament as well as in its
broadly arched growth-lines the present shell seems to be a typical Mathilda.
Cossmann (1912 : 10) recognizes two Cretaceous representatives of the genus,
M. douvillei Cossmann (1912, pi. I, figs. 3-5) and M. faucignyana (Pictet & Roux)
(1849 : 166, pi. 16, fig. i, sub Turritella), both Albian in age. M. douvillei is of
about the same size and proportions as the new species M. coxi, but its ornament,
although of the same general character, differs in detail.
ENGLISH CRETACEOUS TU RRITELLI D AE AND MATHILDIDAE 193
OCCURRENCE. Aptian, Lower Greensand, Atherfield Clay Series (bed known as
the " Crackers "), near Atherfield, Isle of Wight.
Mathilda ahmadi sp. nov.
(PI. 32, fig. 24 ; Text-fig. 18)
HOLOTYPE. In the Sedgwick Museum (Cambridge), reg. no. 6.44649.
DESCRIPTION. The shell is of small-medium size, turriculate, and moderately
broad-spired. The protoconch is missing in the specimens studied, the earliest
preserved whorl being about 1*5 mm. in diameter. All the whorls now present are
bicarinate anteriorly, the posterior carina being slightly the more prominent and
placed weh1 below their median line. The interspace between the carinae is narrower
than the latter and is occupied on the later whorls by one or two secondary spirals.
The part of the whorl posterior to the carinae is occupied by two to five primary
spirals which decrease in strength posteriorly and are separated by interspaces the
width of which is about the same as that of the spirals or slightly greater ; each
interspace is occupied by one secondary spiral. On the later whorls the more
posterior of the main carinae carries spiral threads or (in one specimen) splits up
into two cords. The narrow area anterior to the carinae is occupied by two or
more secondary spirals, and a further carina becomes visible on the last whorl,
continuing the line of the suture. Everywhere very fine and numerous, regularly
spaced collabral threads form a cancellate ornament with the spirals. The base is
convex and the spiral cord which forms its periphery is succeeded inwards by a
number of weaker spirals, alternating in strength. The aperture is rounded and the
inner lip thin and reflected. The growth-lines form a broad, shallow arch with its
vertex at about the middle of the whorl ; the growth-line chord is almost orthocline
(Text-fig. 18).
Measurements of the Holotype :
Height ...... 22 mm.
Spire angle ...... 18°
Height of penultimate whorl ... 4 mm.
Diameter of penultimate whorl . . 8 mm.
DISCUSSION. This species bears a general resemblance to Mathilda coxi, but
differs in the more anterior position of its carinae and the narrower space separating
them, as well as in other details of ornament.
OCCURRENCE. Albian, Greensand, Blackdown, Devon.
INCERTAE SEDIS
" Turritella " turbinata J. de C. Sowerby
(PL 32, figs. 26, 27 ; Text-fig. 19)
1850. Turritella turbinata ]. de C. Sowerby, in Dixon, p. 349, pi. 29, fig. 2.
HOLOTYPE. In the British Museum (Natural History), reg. no. 44520.
DESCRIPTION. The shell is large and phasianelliform. The earlier whorls, up
194 ENGLISH CRETACEOUS TURRITELLI D AE AND MATHILDIDAE
to a diameter of about 15 mm., are not preserved in the specimens studied. The
outline of the later whorls is strongly convex except for a slight concavity near the
posterior suture, the diameter of those forming the spire being almost three times
their height. The whorls are ornamented with about 40 spiral threads which are
finely serrated and separated by interspaces equal to or narrower than their own
width. The threads are of equal strength except for the most posterior six to
eight, which are more prominent than the rest. The base is convex, with a rounded
periphery, and is crossed by spiral threads similar to those on the outer face of the
whorls. The apertural margin is not preserved intact in the available specimens,
but, so far as can be seen, it was evenly rounded anteriorly. The growth-lines
have a broad, shallow sinus, and a point of inflection near each suture ; the growth-
line chord is almost orthocline (Text-fig. 19).
Measurements of Largest Specimen Examined (B.M., no. 98208) :
Height (allowing for missing apex) .... 0.115 mm.
Spire angle ......... 34°
Height of penultimate whorl . . . . . 23 mm.
Diameter of penultimate whorl . . . . . 52 mm.
DISCUSSION. In the holotype the last whorl has been distorted by pressure in a
direction parallel to the axis of the shell, with the result that its height has been
reduced and the convexity of its outline increased. The specimen of which the
measurements are quoted above seems to be undistorted, but the convexity of the
later part of the last whorl at the periphery is stronger than that of the spire whorls.
This species appears to belong to a new genus which probably should be excluded
from the TurriteUidae. Until more perfect specimens are available, however, no
adequate definition of this genus could be given. In its large size and Phasianella-
like shape T. turbinata recalls the long-ranging Jurassic species Bourguetia saemanni
(Oppel) (B. striata (J. Sowerby)), in which, however, the growth-lines are scarcely
arched and slightly opisthocline and the last whorl and aperture are proportionately
higher. Bourguetia is referred to the family Pseudomelaniidae, from which, how-
ever, the growth-lines would appear to exclude T. turbinata.
OCCURRENCE. Cenomanian, Chalk Marl, Sussex (holotype). Same formation,
Ventnor, Isle of Wight.
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ENGLISH CRETACEOUS TURRITELLI D AE AND MATHILDIDAE 195
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Univ. Col. Bull. geol. Sci., 26, i. v -)- 129 pp., 41 pis., map.
MICHELIN, H. 1838. Note sur une argile dependant du Gault observed au Gaty, commune
de G6rodot, Ddpartement de 1'Aube. Mem. Soc. geol. Fr., Paris, 3 : 97-103, pi. 12.
MONTFORT, P. D. DE. 1810. Conchyliologie systematique et classification methodique de coquilles,
2. 676 pp., Paris.
MORRIS, J. 1854. A catalogue of British fossils, 2nd edit, vii + 372 pp. London.
NEWTON, R. B. 1918. Notes on a new species of Turritella and a naticoid shell referred to the
genus Gyrodes from British Cretaceous rocks. Proc. geol. Ass., Lond., 29 : 97-105, pi. 10.
ig6 ENGLISH CRETACEOUS TURRITELLID AE AND MATHILDIDAE
NOETLING, F. 1885. Die Fauna der baltischen Cenoman-geschiebe. Geol. paldont. Abh.,
Berlin, 2 : 199-247 ; pi. 16-23.
ORBIGNY, A. D'. 1842. Paleontologie francaise, terrains cretaces. 2. Gasteropodes. 456 pp.,
pis. 149-236. Paris.
1850. Prodrome de paUontologie stratigraphique universelle des animaux mollusques et
rayonne's, 2. 428 pp. Paris.
PARKINSON, J. 1811. Organic remains of a former world, 3. iv -f- 479 pp., 22 pis. London.
PICTET, F. J. & CAMPICHE, G. 1861-64. Description des fossiles du terrain cretac6 des en-
virons de Sainte-Croix. Materiaux pour la Paleontologie suisse, 3 : 752 pp., pis. 44-98.
Geneve.
PICTET, F. J. & Roux, W. 1849. Description des mollusques fossiles qui se trouvent dans les
gres verts des environs de Geneve. 558 pp., 51 pis. Geneve.
REUSS, A. E. 1845. Die Versteinerungen der bohmischen Kreideformation. Pt. i, iv + 58 pp.,
13 pis. ; Pt. 2, iv + 148 pp., pis. 14-51. Stuttgart.
ROMAN, F. & MAZERAN, P. 1920. Monographic pa!6ontologique de la faune turonien du
Bassin d'Uchaux et de ses dependences. Arch. Mus. Hist, nat., Lyon, 12, M6m. 2. 137
pp., ii pis.
SEMPER, O. 1865. Du genre Mathilda. Additions a la monographic du genre Mathilda.
J. Conchyliol., Paris, 13 : 328-345, pi. 13.
SMITH, W. 1816-19. Strata identified by organized fossils. 32 pp., 18 pis. London. (Pp.
1-16, pis. 1-8, 1816 ; pp. 17-24, pis. 9-13, 1817; pp. 25-32, pis. 14-18, 1819.)
SOHLE, U. 1896. Geologische Aufnahme des Labergebirges bei Oberammergau mit besonderer
Beriicksichtigung des Cenomans in den bayerischen Alpen. Geogn. Jh., Cassel, 9 : 1-66,
pis. 1-8.
SOWERBY, J. DE C. 1827. The mineral conchology of Great Britain, 6. 235 pp., pis. 504-609.
London.
1833. In SEDGWICK, A. & MURCHISON, R. I. A sketch of the structure of the Eastern
Alps with sections through the newer formations on the Northern flanks of the chain and
through the Tertiary deposits of Styria. Trans, geol. Soc. Lond. (2) 3 : 310-420, pis.
35-40-
TATE, R. 1865. On the correlation of the Cretaceous formations of the N.E. of Ireland.
Quart. J. geol. Soc. Lond., 21 : 15-44, pk- 3~5-
WOODWARD, S. P. 1833. Outline of the geology of Norfolk. 54 pp., 6 pis., 5 maps. Norwich.
EXPLANATION OF PLATES
Specimens with numbers prefixed by BM, GSM, or SM are preserved in the British Museum
(Natural History), London, the Geological Survey and Museum, London, and the Sedgwick
Museum, Cambridge, respectively. Where specimens are preserved on a block of matrix, the
vertical line indicates the length of the actual shell.
PLATE 28
FIG. i. Turritella (Torquesia) tamra sp. nov. Aptian, Lower Greensand, Ather-
field Clay Series (bed known as the " Crackers "), near Atherfield, Isle of Wight.
Holotype (bottom right-hand specimen) and paratypes. SM, B. 27329-30. X2 . p. 185
FIG. 2. Turritella (Torquesia) hassani sp. nov. Cenomanian Limestone (bed B
of Jukes-Browne), " near Seaton, Devon ". Holotype. BM, G. 49907. x 2. (See
also PL 29, fig. 9.) . . . . . . . . . . . p. 189
FIG. 3. Turritella (Turritella) unicarinata (S. Woodward). Upper Chalk, Lisburn,
Co. Antrim, Ireland. GSM, 93759. X2. . . . . . -. . p. 182
FIG. 4. Same species, basal view. Same locality. GSM, 93755. X 2.
FIG. 5. Same species. Upper Chalk, White limestone, Northern Ireland. GSM,
28256. X i£.
Bull. B.M. (N.H.) Geol. 7, 6
PLATE 28
TURRITELLA s.str., TURRITELLA (TORQUESIA)
PLATE 29
FIG. 6. Turritella (Torquesia) wagihi sp. nov. Albian, Gault, Folkestone, Kent.
Holotype. GSM, 93745. X2 . . . . . . . . . p. 187
FIGS. 7, 8. Turritella (Turritella) dibleyi Newton. Chalk Marl (Cenomanian),
Margett's Pit, Burham, Kent. Holotype. BM, G. 29362. X i (fig. 7) ; detail,
showing ornament, X3 (fig. 8) . . . . . . . . p. 180
FIG. 9. Turritella (Torquesia) hassani sp. nov. Cenomanian Limestone (bed B of
Jukes-Browne), "near Seaton, Devon". Paratype. BM, GG. 2320. x 2. (See
also PI. 28, fig. 2) ............ p. 189
Bull. B.M. (N.H.) Geol. 7, 6
PLATE 29
TURRITELLA s.str., TURRITELLA (TORQUESIA)
PLATE 30
FIGS. 10, ii. Turritella (Torquesid) granulata J. de C. Sowerby. Albian, Green-
sand, Blackdown, Devon. Holotype. BM, 43667. X 2. Sowerby collection ;
figured J. de C. Sowerby 1827, pi. 565, fig. i. ....... p. 183
FIG. 12. Same species and locality. BM, G. 16149. X2.
FIG. 13. Same species and locality. GSM, 93690. x 2.
FIG. 14. Same species and locality ; variety showing ornament of only four principal
cords on later whorls (see p. 184). BM, G. 74108. x 2.
FIG. 15. Turritella (Torquesia) faizai sp. nov. Albian, Greensand, Blackdown,
Devon. Holotype. SM, B. 44621. X2 p. 188
FIG. 16. Turritella sherborni sp. nov. Albian, Greensand, Blackdown, Devon.
Holotype. BM, G. 74107. X 2 p. 190
Bull. B.M. (N.H.) Geol. 7, 6
PLATE 30
10
12
TURRITELLA (TORQUESIA), TURRITELLA s.lat.
PLATE 31
FIG. 17. Turritella (Torquesia) vibrayeana d'Orbigny. Albian, Gault, Folkestone,
Kent. BM, G. 73790. X2 P- 185
FIG. 18. Same species and locality. BM, G. 73791. X2.
FIG. 19. Same species and locality. BM, G. 73784. x 2.
FIG. 20. Same species and locality. GSM, 93729. X2.
FIG. 21. Same species and locality. GSM, 1703. x 2.
FIG. 22. Same species. Lower Gault, Fairy Dell, Stonebarrow Cliff, Charmouth,
Dorset BM, G. 49825. x i£.
Bull. B.M. (N.H.) Geol. 7, 6
PLATE 31
17
22
TURRITELLA (TORQUESIA)
PLATE 32
FIG. 23. Mathilda coxi sp. nov. Aptian, Lower Greensand, Atherfield Clay
Series (bed known as the " Crackers "), near Atherfield, Isle of Wight. Holotype.
SM, B. 27336. X3 . . p. 191
FIG. 24. Mathilda ahmadi sp. nov. Albian, Greensand, Blackdown, Devon.
Holotype. SM, B. 44649. X 3 . . . . . . . . . p. 193
FIG. 25. Turritella ageri sp. nov. Albian, Greensand, Blackdown, Devon. Holo-
type. BM, G. 74106. X 2 . . . . . . . . . . p. 190
FIG. 26. " Turritella " turbinata J. de C. Sowerby. Cenomanian, Chalk Marl,
Sussex. Holotype. BM, 44520. x i. F. Dixon collection ; figured J. de C.
Sowerby in F. Dixon, 1850, pi. 29, fig. 2 . . . . . . . p. 193
FIG. 27. Same species. Cenomanian, Chalk Marl, Ventnor, Isle of Wight. BM,
98208. x i.
Bull B.M. (N.H.) Geol. 7, 6
PLATE 32
23
26
27
MATHILDA, TURRITELLA s.lat., "TURRITELLA"
PRINTED IN GREAT BRITAIN BY
ADLARD AND SON, LIMITED,
BARTHOLOMEW PRESS, DORKING
•M.
THE MORPHOLOGY OF
TUBICAULIS AFRICANUS
SP. NOV. A FOSSIL FERN
FROM TANGANYIKA
HENRY SMITH HOLDEN
AND
WILLIAM N. CROFT
BULLETIN OF
THE BRITISH MUSEUM (NATURAL HISTORY)
GEOLOGY Vol. 7 No. 7
LONDON: 1962
THE MORPHOLOGY OF
TUBICAULIS AFRICANUS SP. NOV.
A FOSSIL FERN FROM TANGANYIKA
BY
HENRY SMITH HOLDEN and the late WILLIAM N. CROFT
Pp. 197-211 ; Plates 33-36 ; 16 Text-figures
BULLETIN OF
THE BRITISH MUSEUM (NATURAL HISTORY)
GEOLOGY Vol. 7 No. 7
LONDON: 1962
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. 7, No. 7 of the Geological
(Palaeontological) series.
Trustees of the British Museum 1962
Issued December 1962 Price Eleven shillings
THE MORPHOLOGY OF
TUBICAULIS AFRICANUS SP. NOV.
A FOSSIL FERN FROM TANGANYIKA
By HENRY SMITH HOLDEN & WILLIAM NOBLE CROFT
SYNOPSIS
The paper describes the morphology of a new species of fossil fern from the Ruhuhu area of
Tanganyika. This is assigned to the genus Tubicaulis as T. africanus sp. nov. The relation-
ships between the new species and the other fossil ferns assigned to the same genus and to
possibly allied fossils are discussed.
INTRODUCTION
THE stratigraphy of the Ruhuhu area of south-western Tanganyika and its correlation
with the Karroo formations of South Africa have been the subject of detailed study
both by Stockley (1931, 1932, 1947) and Nowack (1937). Stockley's investigations
resulted in the collection of numerous animal and plant fossils among which the
silicified fern stems and their attached roots and leaf bases forming the subject of
this study were included. They were discovered by Charles Kirchstein, who was
at that time Stockley's Field Assistant, lying in a restricted area on the surface slope
of an exposure of Upper Permian age. In this area there is a local disconformity
between the Upper Permian strata and the scarp of the overlying Triassic Kingori
Sandstones and, although it is possible that the specimens could have been dis-
lodged from pockets in the sandstone, Stockley, who was consulted with regard to
this possibility, regarded such derivation as improbable. Inevitably, there must
be some doubt, however, as to whether the specimens are of Upper Permian or Lower
Triassic age. They were sent, with other fossils, to the Department of Palaeonto-
logy of the Natural History Museum and a considerable amount of preliminary work
on them was undertaken by the late W. N. Croft. The pressure of other duties
and the increasing ill-health which ultimately led to his death resulted in their being
put aside and they were not the subject of further study until the latter part of
1959 when Dr. E. I. White, Keeper of the Department, kindly made available both
the specimens and Croft's notes. The specimens, five in number, had been given a
common Locality Number, 8559, and additional distinguishing letters A to E.
They are now registered as ¥.44725-29 in the Department of Palaeontology. Croft
made outline sketches of the specimens indicating their dimensions and these are
reproduced as Text-figures 1-5. It will be seen from these that specimen ¥.44729
was broken into several pieces when received and that a deep crack was present on
one face of specimen ¥.44725. Two of the specimens, ¥.44725 and ¥.44727, are
illustrated in PI. 33, figs, i, 2 and it is clear from a surface examination of these and
the other specimens that this fern possessed an upright stem which was surrounded
GEOLOGY 7, 7. 15
200 THE MORPHOLOGY OF TUBICAULIS AFRICANUS SP. NOV.
FIGS. 1-5. Tubicaulis africanus sp. n. Outline sketches of the five specimens as
originally received indicating their dimensions. Fig. i is the holotype, Figs. 2-5 are
the paratypes. The heavy broken line in each of the figures indicates the position of
the stele. Not drawn to scale.
Fig. i, Holotype, V. 44725, A-B 20-2 cms., C-D 13-3 cms., E-F 13-2 cms. ; Fig. 2,
V. 44726, A-B 9-3 cms., C-D 10*8 cms. ; Fig. 3, ¥.44727, A-B 11-9 cms., C-D 9-7 cms. ;
Fig. 4, ¥.44728, A-B 16-5 cms. ; Fig. 5, ¥.44729, A-B 11-5 cms.
THE MORPHOLOGY OF TUBICAULIS AFRICANUS SP. NOV. 201
by a thick armour of persistent leaf bases. Trial sections indicated that, as so
frequently happens in silicified material, the quality of the preservation of the tissues
varied considerably from one part of any given specimen to another and that the
more delicate tissues of the fossil frequently were replaced, partly or wholly, by
non-structural matrix. This replacement was notably the case in specimen ¥.44727.
The plant tissues, when examined on a cut surface, were mostly light brown in
colour and trial sections indicated that very little carbon had been preserved in
them. Because of this, attempts to obtain " peels " by smoothing and subsequent
etching with hydrofluoric acid often gave very unsatisfactory results, the action of
the acid being uneven and resulting in the production of " ghost peels " in which
either the structural details of the plant tissues were lost or some or all the tissues
were missing completely. During the course of the investigation attempts were
made also to obtain " peels " both of Tubicaulis solenites and Grammatopteris
baldaufi for comparison with the new fern but etching with hydrofluoric acid proved
equally unsuccessful with these also. An exception to the lack of success attending
etching techniques was provided, however, by specimen V. 44729 which yielded a
number of good quality " peels ". The information obtained from these was
amplified by the examination of occasional successful " peels " from some of the
other specimens together with ground sections and selected polished surfaces.
FILICALES
Family COENOPTERIDEAE
Genus TUBICAULIS Cotta, 1832 : 15
Tubicaulis africanus sp. nov.
(Plates 33-36 ; Text-figs, i-n, 15, 16)
DIAGNOSIS. A fossil fern : stem upright surrounded by dense armour of persistent
leaf bases : stem stele cylindrical, protostelic with numerous exarch protoxylem
groups and with medulla of mixed tracheids and parenchyma : pitting of tracheids
multiseriate scalariform or, in narrower tracheids, scalariform : cortex wide with
scattered nests of sclerotic cells : petioles developed in close spiral succession :
petiole traces initially monodesmic, later dividing into three, arched abaxially with
free margins incurved when fully developed : region between stem and persistent
leaf bases filled with long, uniseriate, septate hairs : roots with stout diarch stele
arising at or near free margins of leaf traces.
HOLOTYPE. Specimen ¥.44725 in the Department of Palaeontology, British
Museum, (Natural History).
PARATYPES. Four specimens, ¥.44726-29, in the same Department.
LOCALITY. Ruhuhu area of south-western Tanganyika.
HORIZON. ? Upper Permian.
DESCRIPTION. The Stem. The surface tissues of the stem have not survived
petrifaction and, as a consequence, it has proved impossible to determine its exact
diameter but this ranges from approximately 4-5 cms. to 6-0 cms. Its centre is
occupied by a small cylindrical stele which has a diameter of from 3-5 mm. to 4-0
mm. Its cylindrical character is somewhat obscured by the emission of a close
GEOLOGY, 7, 7. I5§
202 THE MORPHOLOGY OF TUBICAULIS AFRICANUS SP. NOV.
succession of spirally arranged leaf traces showing a complex divergence (PI. 34,
fig. 3 ; PL 36, fig. i). The stelar protoxylem is exarch and consists of a number of
separate groups of small tracheids distributed around the periphery of the stele
(PI. 34, fig3; Pl-35- fig. 3). Themajorityof these possess scalariform pitting although,
occasionally, they show suggestions of close spiral thickening. The metaxylem
tracheids form a continuous hollow cylinder which varies somewhat in depth and
8
FIGS. 6-8. Tubicaulis africanus sp. n. Outline sketches of portions of three longi-
tudinal " peels " from V. 44729. Fig. 6 (Peel n, ¥.44729(13)), Fig. 7 (Peel 13, ¥.44729
(14)), Fig. 8 (Peel 14, ¥.44729(15)), showing the close succession of the petiole traces
and the temporary vascular linkage of some of these. The petiole traces are numbered
in the order of their emission from the stele.
which encloses a " mixed pith " consisting of a parenchymatous ground mass
among which tracheids are scattered singly or in small groups (PI. 34, fig. 3 ; PL
35, figs, i, 2). The medullary parenchyma is not sharply delimited from the main
mass of the metaxylem but extends into it in the form of small groups or radial
plates of cells which may penetrate locally almost to the outer margin of the stele
and even into the bases of the departing leaf traces (PL 35, fig. 4). The type of
pitting of the metaxylem is often difficult to determine owing to imperfect preserva-
tion but, where this is sufficiently good, it is seen to consist of a single row of trans-
THE MORPHOLOGY OF TUBICAULIS AFRICANUS SP. NOV. 203
versely elongated pits in the narrower tracheids whilst the wider ones possess two or
three similar rows. The pitting is thus of the type termed multiseriate scalari-
form by Delevoryas & Morgan (1952). Most of the medullary tracheids are spindle
shaped with pitting similar to that of the rest of the metaxylem although shorter
tracheids with truncated ends are not uncommon, (PI. 35, fig. 2). The phloem and
its associated parenchyma have usually not been preserved and, at the best, are
represented by shreds of pale brown tissue in which no structural features can be
made out.
Scattered through the wide cortex are abundant groups of thick-walled cells
forming sclerotic nests (PI. 34, fig. 3 ; PI. 35, fig. I ; PI. 36, fig. i), the component
cells of which have dark contents. Some of these sclerotic nests have an attached
fringe of parenchyma and are linked occasionally by similar groups of thin-walled
cells. No continuous areas of parenchymatous ground tissue have been preserved
although this does not appear to have led to any great spatial disturbance of the
petiole and root traces traversing the stem cortex. The groups of sclerotic cells
are frequently closely associated with these traces although this may be due, in part,
to the loss of the cortical parenchyma.
The petiole. As already stated, the petiole traces are given off from the stem stele
in close spiral succession and, as they begin to emerge from the outer tissues of the
stem cortex, they produce in this an irregular series of superficial corrugations
(PI. 34, fig. 2). This crowded spiral succession may result also in a temporary
vascular linkage between neighbouring petiole traces near the region of their emer-
gence from the stem and is illustrated in PI. 35, fig. 5 and, in outline, in Text-figs.
6-8. They develop a well-marked abaxial curvature immediately (PI. 35, fig. 3)
with the protoxylem tracheids forming a series of separate groups distributed
along the convex adaxial surface. This distribution of the protoxylem is well
shown not only in transverse sections of the petiole (PI. 35, fig. 3) but also in tangential
sections passing through the petiolar bundle at right angles to the sagittal plane
immediately after it leaves the stem stele (PI. 35, figs. 4, 5). Closely associated
with each petiole trace is an oval mass of sclerotic cells which occupies its abaxial
concavity. A continuation of this particular mass becomes closely applied to the
abaxial face of the petiole trace as it moves outwards and forms a continuous
strip along the whole abaxial face of the trace during its passage through the outer
cortex of the stem. Similar groups of sclerotic cells are distributed also along the
convex adaxial face of the trace and, near its point of origin, often form a more or
less continuous band. Scattered irregularly through the strips of sclerotic cells
which accompany the petiole traces are large ovoid cells which occur singly or in
groups of two or three (PL 36, fig. 4). They are readily distinguished from the sur-
rounding tissues by their larger size and by the absence of dark contents. They
recall the elements described by C. Eg. & P. Bertrand (1911) as occurring in the
inner cortex of Tubicaulis berthieri and which are interpreted by them as glandular
cells. The petiole trace, during its passage outwards, takes a sharp upward course
through the stem cortex. As it enters the base of the petiole, however, its upward
curvature becomes much less pronounced so that, from the time of its inception
to its later stages, the trace has the form of a sigmoid curve. During its transit,
204
THE MORPHOLOGY OF TUBICAULIS AFRICANUS SP. NOV.
the abaxial curvature of the petiolar trace becomes steadily more pronounced with
its free margins strongly recurved so that, when it enters the petiole, the xylem,
in transverse section, has the shape of a slender letter C. (PI. 36, fig. i). The
basal part of the petiolar cortex contains numerous scattered sclerotic nests and
similar tissue, several cells in depth, forms a continuous layer immediately below
the epidermis. This last, if preserved at all, is usually represented by a pale yellowish
layer in which the cellular structure is lost. As the petiole moves further away from
(10)
FIGS. 9-10. Tubicaulis africanus sp. n. Camera lucida sketches of sections of two
petioles. Fig. 9 from V. 44726;' shows, on the left, the complete division of the petiole
trace and the separation of the distal part from the rest by cortical tissue and, on the
right, the thinning of the xylem preparatory to division. Fig. 10 from ¥^44726^ shows
the symmetrical division of the petiole trace into three. Xylem in solid black, cortical
tissues stippled (Both x8).
the stem, the sclerotic nests diminish in number and tend to disappear completely
whilst the cortical tissues become homogeneous and are composed of small, some-
what thick- walled, cells apart from the continous layer of sclerenchyma lying just
below the epidermis. This stage appears to be reached only in the more distal parts
of the persistent leaf bases. At the same level, extramarginal gaps are sometimes
developed in the petiolar xylem. These gaps may occur initially on one side only
or they may arise symmetrically on either flank of the trace (Text-figs. 9, 10). When
the first examples of these gaps were found it was thought they might be artefacts
which had resulted from locally imperfect petrifaction but further investigation
showed that they were a normal feature in which the division of the xylem of the
petiole trace was followed by its separation into three independent strands with the
cortical tissues passing through the gaps between them. Whilst these gaps in the
petiolar vascular supply resemble those resulting from the emission of extramarginal
pinna-traces, it should be pointed out that none of the petioles examined shows any
indication of the existence of such traces, although it is conceivable that they may
develop from the contiguous margins of the xylem gaps or from the central bar at a
higher level. However, no evidence of this has been found. A similar division of
an initially monodesmic leaf trace is not uncommon in existing ferns (Davie, 1914),
THE MORPHOLOGY OF TUBICAULIS AFRICANUS SP. NOV.
205
typical examples being those of Cibotium barometz and Saccoloma elegans. Although
the adaxially curved petiole trace in both these ferns is more elaborate than that of
Tubicaulis africanus both show a transient division into three meristeles, one median
and two lateral, prior to their further sub-division (Bower, 1923 ; Davie, 1914 ;
G wynne- Vaughan, 1903). The persistent petiole bases are embedded in a dense
mass of long uniseriate hairs through which the roots pass to the exterior (PI. 36,
fig. 3). Although the preservation of these hairs is imperfect, there are indications
(13)
FIGS. 11-14. Fig. ii, Tubicaulis africanus sp. n. Portion of the petiolar epidermis
showing the bases of several hairs, ¥.44726;'. (X3oo). Figs. 12, 13, Bases of two
hairs of Cibotium regale showing the thickened basal cell and, in 12, a bifurcation
near the base (x 150). Fig. 14, Bases of two hairs from C. barometz (x 150).
that, like those found in many existing ferns, they were multicellular. Some of
them arise from the basal parts of the petiolar epidermis and appear to develop from
a single basal cell (Text-fig, n) but it seems probable that many were developed
also from the surface tissues of the stem. The existence of such surface hairs as
distinct from flattened scales is widespread among palaeozoic ferns and they occur
also in many existing ferns which are regarded as retaining other primitive characters
(Bower, 1926). Comparatively short uniseriate hairs arising from a single basal
cell form a thick covering to the rhizomes and petiole bases of Matonia pectinata
(Seward, 1899). Those occurring in Tubicaulis africanus, however, are longer and
appear to resemble more closely the long, brownish or golden yellow hairs which
form a dense thatch clothing the stem apices and young petioles of various species
of Cibotium such as C. barometz and C. regale. The hairs in these two species are
206
THE MORPHOLOGY OF TUBICAULIS AFRICANUS SP. NOV.
somewhat coarser than the ones present in Tubicaulis africanus although, like them,
they arise from a single basal cell. (Text-figs. 12-14). In Cibotium regale, however,
several examples have been noted in which the hairs divide into two equal branches
near the base (Text-fig 12). The basal cell in this species has thicker walls than
the remaining cells of which the hair is composed and, when the hair is shed, this
basal cell persists as a large, convex, golden-yellow boss which stands out prominently
from the smaller epidermal cells which surround it.
Roots. The roots arise on or near the free incurved margins of the petiole traces
(PI. 35, fig. 6 ; Text-fig. 16), usually in pairs, one from each margin. No evidence
has been found indicating that they develop from the stem as well. As a rule,
only the comparatively stout diarch stelar xylem of the root is preserved and this
is surrounded by a sheath of dark sclerotic elements with a clear space between this
(15)
FIGS. 15, 16. Tubicaulis africanus sp. n. Fig. 15, T.S. of a root in the stem cortex
showing the stout diarch stele and traces of the superficial tissues. The sclerotic
tissue is stippled. ¥.44729(28). X 350. Fig. 16, The recurved margin of a petiole
trace showing the origin of a root. Note the larger diameter of the tracheids of the
root metaxylem compared with those of the petiole. The sclerotic tissue is stippled.
¥.44729(27). X200.
and the root stele (Text-fig. 15). In this clear space but well separated from the
stele, there are often traces of the other root tissues, these usually taking the form
of a delicate complete or broken cylinder (Text-fig. 15). Occasionally, roots
occur in which the whole of the cortical tissues have been partly petrified but, even
here, the preservation is such that all that can be said is that they are parenchy-
matous. There is some evidence that the roots may branch whilst still within the
stem cortex or among the hairs which enclose the persistent leaf bases. In a few
instances, traces of the roots are found at the surface of the specimens and, where this
is the case, these take the form of short oval rods or papillae representing the root
steles.
DISCUSSION. The seven species at present included in the genus Tubicaulis
comprise ferns showing wide differences in habit, these ranging from the relatively
small epiphytic forms such as T. berthieri and T. scandens to those like T. solenites
THE MORPHOLOGY OF TUBICAULIS AFRICANUS SP. NOV. 207
and T. africanus which possessed stout upright stems enclosed in an armour of
persistent leaf bases. This wide variation in habit is not, in itself, a valid reason
for generic separation since a similarly wide variation in habit occurs in many
existing fern genera. The genus Asplenium, for example, includes, in addition
to both creeping and radially organised terrestrial species, minute epiphytes and
massive specialised ones. They are included in the same genus by taxonomists in
view of the agreement in their soral and sporangial characteristics. In the absence
of the reproductive organs, however, palaeobotanists are compelled to rely solely
on such morphological characters as are available in the specimens they study, this
imposing a limitation which has been stressed already by Mamay (1952) and, more
recently still, by Eggert (1959). A further handicap is often imposed by the small
number of specimens which has been discovered and by their incomplete character,
a handicap well-illustrated by Tubicaulis itself in which five of the seven species
assigned to it have been described from single examples. In spite of these limita-
tions, however, it seems reasonable, in the present state of our knowledge, to assign
them provisionally to the same genus since all possess certain anatomical features
in common, namely :
(a) An essentially terete, protostelic axial vascular system devoid of secondary
thickening with exarch protoxylem groups and in which the protostele may
be wholly tracheidal or possess a core of mixed tracheids and parenchyma ;
(b) petiole traces which depart from the stem stele, either as flat or slightly
curved tangential bands which assume, ultimately, the shape of a letter C
in transverse section and in which the curvature is abaxial, or in which the
abaxial curvature is well-developed immediately on the separation of the
petiole trace from the stem stele.
It will be noted that this definition of the essential generic vascular characteristics
is slightly modified from that proposed by Mamay (1952) in the introduction to his
description of Tubicaulis scandens and has been rendered necessary as a result of
the discovery and description of other species subsequent to the publication of his
paper.
The root systems of the various species provided no indications of common ancestry
apart from the fact that the diarch steles are of the stout osmundaceous type rather
than the slenderer type characteristic of existing leptosporangiate ferns. Like
the roots of the fossil Osmundaceae, they show a wide variation in their source of
origin (Kidston & Gwynne-Vaughan, 1907-14). Thus in T. scandens they arise
solely from the stem (Mamay, 1952), in T. sutcliffii (Stopes, 1906), T. multiscalari-
formis (Delevoryas & Morgan, 1952) and T. stewartii (Eggert, 1959) they arise from
the stems and basal parts of the petioles and in T. africanus from the peti jles only.
Adequate knowledge of the nature of the pinna-traces is lacking also. Stopes
(1906) records the presence of two small strands in the cortex of one of the petioles
of T. sutcliffii which were, presumably, pinna-traces although their origin could not
be determined. Mamay (1952) shows that, in T. scandens, the pinna-traces are
paired and that they are slightly but definitely extramarginal in origin whilst,
in T, solenites in which they are also paired and slightly extramarginal, they bifurcate
208 THE MORPHOLOGY OF TUBICAULIS AFRICANUS SP. NOV.
whilst still within the petiolar cortex, each branch supplying a separate pinna
(Stenzel, 1889, pi. i, figs. 4-9). Morgan & Delevoryas (1954), on the basis of informa-
tion supplied by Nickerson, state that in T. multiscalariformis the pinnae are borne
in two rows, one on either side of the petiole. Grateful acknowledgments are due
to Dr. Nickerson who has kindly supplied " peels " from coal balls in the collection
of the Washington University Department of Botany confirming this.
A point worthy of note is that in both T. solenites and T. scandens the departure of
the pinna-trace does not appear to cause a break in the continuity of the main petiolar
strand. In attempting to indicate affinities within the genus, we are thus left only
with the stelar characters and those of the petiole traces. Judged by these characters,
the seven species assigned to Tubicaulis fall into two groups, T. solenites, T. sutcliffii,
T. berthieri and T. scandens all possessing a homogeneous, wholly tracheidal proto-
stele and a leaf trace which, initially, has the form of a flat or slightly curved tan-
gentially expanded bar, whilst T. multiscalariformis, T. stewartii and T. africanus
are characterized by the development of intrastelar parenchyma, slight in amount in
T. multiscalariformis (Eggert, 1959) but much more abundant in both T. stewartii
and T. africanus. These three species also agree in possessing petiole traces in
which the assumption of an abaxial curvature occurs at a very early stage. Although
there is a close resemblance in vascular structure between T. stewartii and T.
africanus and also in the fact that both possess a surface covering of closely set
uniseriate multicellular hairs, they differ considerably in habit and in numerous
other features and these differences, combined with the wide difference in geological
age, rule out any suggestion that they may have been specifically identical.
The relationship of the seven species now included in the genus Tubicaulis to
other palaeozoic ferns does not appear to be a close one. The genera which resemble
Tubicaulis in possessing an abaxially curved petiole trace are Psalixylon (Holden,
1960), Apotropteris and, possibly, Anachoropteris (Morgan & Delevoryas, 1954).
All these are readily distinguished anatomically from Tubicaulis by their petiolar
vascular characters and the likelihood of any close relationship either with that
genus or with each other seems remote. A recent paper by Hall (1961) has described
for the first time a radially organised plant bearing typical Anachoropteris involuta
petioles. The stem of this plant is protostelic with, probably, exarch protoxylem.
The stelar xylem is not homogeneous but contains plates of parenchyma interspersed
among the tracheids. The trend towards the replacement of tracheids by paren-
chyma in the centre of the stele (i.e. towards medullation) is of wide occurrence in
both the ferns and also in the fossil lycopods and there seems no valid reason why
this plant should not be assigned to the genus Anachoropteris in view of its petiolar
anatomy.
The fact that an abaxially curved leaf trace occurs in ferns along several apparently
unrelated or very distantly related lines of descent suggests that this type of leaf
trace may well have arisen independently from different ancestral stocks.
Of the other palaeozoic fern genera, Grammatopteris has been the one considered
to be nearest Tubicaulis (Scott, 1920) although this view is open to some doubt.
Of the two species of Grammatopteris of which both stem and petiole are known,
G. rigolloti has been studied by Renault (1891, 1896) and Corsin (1937) whilst the
THE MORPHOLOGY OF TUBICAULIS AFRICANUS SP. NOV. 209
most recent investigation of G. baldaufi has been that of Sahni (1932). Both species
possess an exarch protostele and, in both, the petiole trace has the form of a tan-
gentially flattened bar which, in this respect, resembles that of those species of
Tubicaulis in which the abaxial curvature of the petiole trace is, initially, slight or
absent. Renault's account and figures of Grammatopteris rigolloti (Renault, 1891,
1896), although adequate for diagnostic purposes, left a number of features obscure
and Corsin's more detailed study has served both to amplify Renault's account and
to reveal additional points of interest. The most unexpected result of Corsin's
work was that it demonstrated the presence in the holotype of a thin peripheral zone
of secondary xylem surrounding the stem stele. This encloses a solid mass of pri-
mary xylem composed entirely of tracheids of varying diameter in which the pits
apart from the protoxylem were multiseriate.
Renault's co-type was a small, thick preparation which showed no trace of secon-
dary xylem. The tissues forming the core of the stele are imperfectly preserved
but Corsin interpreted them as probably representing the pith, an interpretation
which receives support from the two additional sections prepared from " le culot de
taille " in which the presence of intrastelar parenchyma is undoubted. Corsin con-
sidered that the differences between the type section and the others were due to
their being cut at different levels, the type being from nearer the base than the
co-type and the additional sections (Corsin, 1937 : 52).
The petiole trace of G. rigolloti, although it shows a general resemblance to that of
some species of Tubicaulis (e.g. T. solenites, T. scandens], differs in a number of details.
Briefly, these are (a) that the protoxylem is confined to two slightly adaxial groups
occupying the opposite margins of the trace, (b) that the slight curvature seen in the
more distal parts of the trace is adaxial, and (c) that the pinna-traces are marginal
in origin.
The specimens of Grammatopteris baldaufi do not appear to be so well preserved
as those of G. rigolloti but it seems clear that the plant was also protostelic with a
complete absence of intrastelar parenchyma but with a stellate core, most of which
consisted of normal, elongated tracheids mixed with a small number of short, much
wider ones (the " parenchymatous tracheids" of Sahni). The petiole trace is very
like that of G. rigolloti with the protoxylem forming two groups, one at each end of
the tangentially flattened xylem bar. The trace, in many instances, shows a slight
adaxial curvature in the free petiole although this is possibly an artefact.
From the available data is seems clear that the relationship between Tubicaulis
and Grammatopteris is not a particularly close one and that the genus Tubicaulis,
as at present constituted, must continue to occupy a somewhat isolated position
among the palaeozoic Filicales.
SUMMARY
Tubicaulis africanus is a fern probably of Upper Permian age which was dis-
covered in the Ruhuhu area of Tanganyika. It possesses a stout, upright stem
surrounded by a close armour of persistent leaf bases. The stem has a slender,
cylindrical stele in which the protoxylem is exarch and in which the metaxylem
consists of a peripheral, wholly tracheidal zone enclosing a core of mixed tracheids
210 THE MORPHOLOGY OF TUBICAULIS AFRICANUS SP. NOV.
and parenchyma. The wide cortex is characterized by abundant scattered nests
of sclerotic cells with dark contents among which larger clear ovoid cells occur.
The petioles are given off in close spiral succession. The petiolar vascular supply
is initially monodesmic with a distinct abaxial curvature which becomes steadily
more pronounced distally and in which the free margins ultimately are recurved.
The numerous protoxylem groups are distributed along the whole of the adaxial
surface of the trace. The region between the surface of the stem and the petiole
bases is packed with long uniseriate hairs.
The roots, which arise at and near the margins of the petiole traces, are diarch
with a stout stele resembling that of the zygopterids and Osmundaceae.
ACKNOWLEDGEMENTS
Grateful acknowledgements are due to Dr. E. I. White, Keeper of the Department
of Palaeontology, for placing the specimens of Tubicaulis at my disposal and for
facilities for their study. I am indebted also to Mr. F. M. Wonnacott of the Depart-
ment for several helpful suggestions. I am further indebted to Dr. W. Dean for the
photograph reproduced in Plate 34, fig. i. The expert assistance of the staff of the
Photographic Department has been invaluable and all the remaining figures in
Plates 33-36 are from negatives and prints supplied by them.
REFERENCES
BERTRAND, C.-Eg. & P. 1911. Le Tubicaulis berthieri (sp. nov.). Bull. Soc. Hist. nat. Autun,
24 : 43-92, pis. 1-3.
BOWER, F. O. 1923. The Ferns (Filicales), 1. Analytical Examination of the Criteria of Com-
parison. 350 pp., 307 figs. Cambridge.
— 1926. The Dermal Appendages of the Ferns. Ann. Bot., London, 40 : 479-490.
CORSIN, P. 1937. Contribution a I'etude des Fougeres anciennes du Groupe des Inversicatenales.
247 pp., 42 pis. Lille.
COTTA, C. B. 1832. Die Dendrolithen in Beziehung auf ihren innern Bau. ix + 89 pp., 20
pis. Dresden & Leipzig.
DAVIE, R. C. 1914. The Pinna-Trace in the Ferns. Trans. Roy. Soc. Edinb., 50 : 349-378,
Pis. 33-35-
DELEVORYAS, T. & MORGAN, J. 1952. Tubicaulis multiscalariformis : a new American
coenopterid. Amer. J. Bot., Lancaster, 39 : 160-166, 15 figs.
EGGERT, D. A. 1959. Studies of palaeozoic ferns : Tubicaulis stewartii sp. nov. and evolu-
tionary trends in the genus. Amer. J. Bot., Lancaster, 46 : 594-602, 14 figs.
GWYNNE-VAUGHAN, D. T. 1903. Observations on the Anatomy of Solenostelic Ferns, Pt. 2.
Ann. Bot., London, 17 : 689-742, pis. 33-35.
HALL, J. 1961. Anachoropteris involuta and its attachment to a Tubicaulis type of stem from
the Pennsylvanian of Iowa. Amer. J. Bot., Lancaster, 48 : 731-737, n figs.
HOLDEN, H. S. 1960. The Morphology and Relationships of Rachiopteris cylindrica. Bull.
Brit. Mus. (Nat. Hist.} Geol., London, 4 : 51-69, pis. 8-10.
KIDSTON, R. & GWYNNE-VAUGHAN, D. T. 1907-14. On the Fossil Osmundaceae, Pts. I-V.
Trans. Roy. Soc. Edinb., 45 : 759-780, pis. 1-6 (1907) ; Pt. II, 46 : 213-232, pis. 1-4 (1908) ;
Pt. Ill, 46 : 651-667, pis. 1-8 (1909) ; Pt. IV, 47 : 455-477, pis. 1-4 (1910) ; Pt. V, 50 1469-
479, pis. 41-44 (1914)-
MAM AY, S. H. 1952. An Epiphytic American Species of Tubicaulis. Ann. Bot., London
(n.s.) 16 : 145-162, pi. 9.
THE MORPHOLOGY OF TUBICAULIS AFRICANUS SP. NOV. 211
MORGAN, J. & DELEVORYAS, T. 1954. An anatomical study of a new coenopterid and its
bearing on the morphology of certain coenopterid petioles. Amer. J. Bot., Lancaster,
41 : 198-203, 9 figs.
NOWACK, E. 1937. Zur Kenntnis der Karroo Formation in Ruhuhu Graben (D.O.A.). N.
Jb. Min. Geol. Paldont., Stuttgart, 68, B : 380-412, 7 figs.
RENAULT, B. 1891. Note sur la famille des Botryopte'ridees. Bull. Soc. Hist. nat. Autun,
4: 349-373. pis. 10, n.
1896. Bassin Houiller et permien d'Autun et d'Epinac, IV. Flore Fossile II. 578 pp.,
atlas, pis. 28-89. Paris.
SAHNI, B. 1932. On a Palaeozoic Tree-fern, Grammatopteris baldaufi (Beck) Hirmer, a Link
between the Zygopterideae and Osmundaceae. Ann. Bot., London, 46 : 863-877, pi. 35.
SCOTT, D. H. 1920. Studies in Fossil Botany, I. Pteridophyta. 3rd ed., xxiii + 434 PP-,
190 figs. London.
SEWARD, A. C. 1899. On the Structure and Affinities of Matonia pectinata. Philos. Trans.,
London (B) 191 : 171-209, pis. 1-3.
STENZEL, G. 1889. Die Gattung Tubicaulis Cotta. Bibl. Bot., Cassel, 12 : 1-50, pis. 1-7.
STOCKLEY, G. M. 1931. Report on the Geology of the Ruhuhu Coalfield. Tanganyika
Geol. Surv. Bull., 2 : 1-63, pis. 1-6.
1932. The Geology of the Ruhuhu Coalfield, Tanganyika Territory. Quart. J. Geol. Soc.
Lond., 88 : 610-622, pi. 38.
1947. The Geology and Mineral Resources of Tanganyika Territory. Bull. Imp. Inst.
Lond., 45 : 375-406, pis. 1-4.
STOPES, M. C. 1906. A New Fern from the Coal Measures, Tubicaulis sutcliffii sp. nov. Mem.
Manchr. Lit. Phil. Soc., 50 : 1-34, pis. 1-3.
PLATE 33
Tubicaulis africanus sp. n.
FIG. i. Holotype (V. 44275). x f approx.
FIG. 2. Another specimen, V.4472y, slightly enlarged.
Bull. B.M. (N.H.) Geol. 7, 7
PLATE 33
TUBICAULIS
PLATE 34
Tubicaulis africanus sp. n.
FIG. i. Part of the smoothed and polished surface of a transverse section through ¥.44726^
showing the armour of crowded, persistent leaf bases. About nat. size.
FIG. 2. Part of the smoothed and polished surface of a transverse section through V.44726c
showing the stele and part of the stem cortex and leaf bases, (x i'25).
FIG. 3. T.S. of stele showing the " mixed pith " and five still attached leaf traces, V.44729
(28). (Xi5).
Bull. EM. (N.H.) Geol. 7, 7
PLATE 34
TUBICAULIS
PLATE 35
Tubicaulis africanus sp. n.
FIG. i. L.S. of stele showing the " mixed pith ", bases of leaf traces and patches of cortical
sclerenchyma. ¥.44729(21). Longitudinal series No. 25. (xi5).
FIG. 2. L.S. of part of stele more highly magnified. ¥.44729(24). Longitudinal series
No. 30. (X3o).
FIG. 3. T.S. of a leaf trace at the point of departure from the stele, ¥.44729(25). (X2o).
FIG. 4. Tangential section of a leaf trace showing the peripheral protoxylem groups,
¥.44729(13). (X25).
FIG. 5. Tangential section of two leaf traces showing their temporary vascular linkage,
¥.44729(14). (X2o).
FIG. 6. T.S. of a leaf trace in the stem cortex from which a root trace has just been given off.
¥. 44729(25). (xis).
Bull. B.M. (N.H.) Geol. 7, 7
PLATE 35
4
TUBICAULIS
PLATE 36
Tubicaulis africanus sp. n.
FIG. i . Tangential section through the stem cortex showing the crowded leaf traces and the
scattered areas of sclerenchyma, ¥.44729(13). (X3o).
FIG. 2. T.S. of a fully developed leaf trace, ¥.44729(31). ( x 12).
FIG. 3. Part of the area between two petioles showing the packing of hairs, ¥.447267.
(X3Q).
FIG. 4. An area of petiolar cortical sclerenchyma in L.S. showing the groups of " clear "
cells, ¥.44729(30). (X3o).
Bull. B.M. (N.H.) GeoL 7, 7
PLATE 36
TUBICAULIS
H V-
THE ORDOVICIAN
TRILOBITE FAUNAS OF
SOUTH SHROPSHIRE, III
W. T. DEAN
BULLETIN OF
THE BRITISH MUSEUM (NATURAL HISTORY)
GEOLOGY Vol. 7 No. 8
LONDON: 1963
THE ORDOVICIAN TRILOBITE FAUNAS OF
SOUTH SHROPSHIRE, III
BY
WILLIAM THORNTON DEAN
Pp. 213-254 ; Pis. 37-46
BULLETIN OF
THE BRITISH MUSEUM (NATURAL HISTORY)
GEOLOGY Vol. 7 No. 8
LONDON: 1963
THE BULLETIN OF THE BRITISH MUSEUM
(NATURAL HISTORY), instituted in 1949, is
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within one calendar year.
This paper is Vol. 7, No. 8 of the Geological
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© Trustees of the British Museum 1963
PRINTED BY ORDER OF THE TRUSTEES OF
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Issued February, 1963 Price Thirty Shillings
THE ORDOVICIAN TRILOBITE FAUNAS OF
SOUTH SHROPSHIRE, III
By WILLIAM THORNTON DEAN
SYNOPSIS
This paper is the third of a series of four dealing with the Caradoc trilobites of south Shropshire.
Those described belong to fifteen genera or subgenera, of which one, Remopleurella, is new, and
twenty-six species or subspecies, of which five are new. The trilobites comprise the gonatoparian
family Calymenidae, together with all the known opisthoparian forms, belonging to nine families.
CONTENTS
Page
I. SYSTEMATIC DESCRIPTIONS ........ 216
Family Calymenidae Burmeister . . . . . . .216
Flexicalymene acantha Bancroft . . . . . . .216
Flexicalymene caractaci (Salter) . . . . . .217
Flexicalymene planiwiarginata (Reed) . . . . .217
Flexicalymene cobboldi sp. nov. . . . . . . .218
Flexicalymene (Reacalymene) pusulosa (Shirley) .... 220
Flexicalymene (Reacalymene) cf. limba (Shirley) . . . .221
Flexicalymene (Reacalymene) horderleyensis sp. nov. . . .221
Onnicalymene onniensis (Shirley) . . . . . .223
Onnicalymene laticeps (Bancroft) . . . . . .224
Onnicalymene salteri (Bancroft) . . . . . .224
Gravicalymene praecox (Bancroft) . . . . . .225
Gravicalymene inflata sp. nov. . . . . . . .227
Family Asaphidae Burmeister ... . .229
Subfamily Asaphinae Burmeister . . . . . .229
Parabasilicus powisi (Murchison) ..... 229
Family Illaenidae Hawle & Corda ....... 233
Subfamily Illaeninae Hawle & Corda . . . . 233
Illaenus cf. fallax Holm .... -233
Family Lichidae Hawle & Corda ... -234
Subfamily Lichinae Hawle & Corda ...... 234
Metopolichas ? sp. ........ 234
Subfamily Homolichinae Phleger ... . 235
Platylichas laxatus (M'Coy) . . . 235
Family Odontopleuridae Burmeister . -237
Subfamily Odontopleurinae Burmeister . . . -237
Primaspis harnagensis (Bancroft) . . .237
Primaspis caractaci (Salter) . . . . .239
Family Olenidae Burmeister ... . . 241
Subfamily Triarthrinae Ulrich .... .241
Triarthrus cf. Hnnarssoni Thorslund ... . 241
Family Otarionidae R. & E. Richter 242
Subfamily Otarioninae R. & E. Richter ..... 242
Otarion sp. ..... ... 242
GEOL. 7, 8 1 6
2i6 ORDOVICIAN TRILOBITE FAUNAS OF SOUTH SHROPSHIRE, III
Family Proetidae Salter ........ 243
Subfamily Proetidellinae Hup6 ...... 243
Proetidella fearnsidesi Bancroft ...... 243
Proetidella? sp. ....... 246
Family Remopleurididae Hawle & Corda ...... 246
Subfamily Remopleuridinae Hawle & Corda .... 246
Remopleurides warburgae sp. nov. ..... 246
Remopleurides latus Olin onniensis subsp. nov. . . . 247
Remopleurella burmeisteri (Bancroft) gen. nov. . . . 250
Family Thysanopeltidae Hawle & Corda . . . . . .251
Eobronteus ? sp. . . . . . . . . 251
II. REFERENCES . . . . . . . . . .251
Family CALYMENIDAE Burmeister, 1843
Genus FLEXICALYMENE Shirley, 1936
TYPE SPECIES. Calymene Blumenbachii var. Caractaci Salter, 1865 by original
designation of Shirley (1936 : 395).
Flexicalymene acantha Bancroft
(PI. 37, figs, i, 3-6, 14)
1949. Flexicalymene acantha Bancroft, p. 305, pi. n, figs. 29, 30.
1958. Flexicalymene acantha Bancroft : Dean, p. 219.
Bancroft's original description was founded on two specimens, the holotype
cranidium and a paratype pygidium, both probably immature. A few large topotype
cranidia have been collected and these show slight differences from the holotype.
The anterior border is somewhat shorter proportionately and a little less steeply
upturned, the frontal glabellar lobe is more rounded, and there is a well-defined
break in the glabellar outline, so that the latter narrows sharply, immediately in
front of the second glabellar lobes. Where the external impression of the test is
available it is seen to be practically smooth, with only occasional traces of tubercles,
contrasting markedly with the almost prickly appearance of immature specimens.
One specimen has been found with thorax complete (PL 37, fig. 14). There are
thirteen segments of normal calymenid type and the surface of the axial rings and
pleurae, excluding furrows, is ornamented with small tubercles ; again, the specimen
represents an immature individual. The pygidium figured by Bancroft (1949, pi. II,
fig. 30) has the axis distorted by crushing so that it appears too narrow ; normally
it occupies frontally about one-third of the maximum breadth. There are six axial
rings, of which the sixth is poorly defined, and five pairs of furrowed pleural ribs,
not six as stated by Bancroft (1949 : 305). No well-preserved pygidium of F. acantha
has yet been found in association with the cephalon of the species, therefore some
doubt must exist as to the true identity of the paratype pygidium, as another
calymenid, Gravicalymene praecox (Bancroft), occurs in the same strata.
HORIZON AND LOCALITIES. In the Costonian Stage of the Coston district, near
Aston-on-Clun, and at Brokenstones, near Horderley, there occur numerous caly-
menids which are close to Bancroft's species, and some are figured here as Flexicaly-
ORDOVICIAN TRILOBITE FAUNAS OF SOUTH SHROPSHIRE, III 217
mene cf. acantha (PL 37, figs. 7, 10, 13). Such differences as exist, for example,
degree of inclination of the anterior border, straightening of the normally convex
margins of the glabella, and variation in the shape of the frontal glabellar lobe,
could well be the result of mechanical deformation and it is not proposed to regard
them as distinct from F. acantha. The earliest known, uncommon occurrences are
in the middle Costonian of Coston but specimens are moderately abundant in the
succeeding upper Costonian.
The type locality is the old cartway section near the south end of Smeathen
Wood, Horderley, 250 yards west-south-west of Wood House, where the species
occurs in small numbers in the lowest Smeathen Wood Beds, Harnagian Stage,
Reuscholithus reuschi Zone. Flexicalymene acantha has been found neither at higher
horizons, nor outside the southern part of the Caradoc district. In the northern
part of the district, however, Flexicalymene (s.s.) is believed to occur in the topmost
Costonian Stage, near Cressage, but the available specimens are too fragmentary
to ascertain whether another species is represented ; furthermore the dominant form
in the Cressage strata is Flexicalymene (Reacalymene] pusulosa (Shirley).
HOLOTYPE. BM. In. 42091 (PI. 37, fig. 3).
PARATYPE. BM. In. 49592 (PI. 37, fig. 4).
DISCUSSION. For convenience the discussion of the species of Flexicalymene (s.s.)
has been placed after the description of F. cobboldi.
Flexicalymene planimarginata (Reed)
(PI. 37, fig. 15)
1906. Calymene planimarginata Reed, p. 137, pi. 17, fig. 15.
1960. Flexicalymene planimarginata (Reed), Whittard, p. 158. Includes full synonymy of the
species.
A few pygidia have been collected from the Lower Longvillian Substage south of
the Onny Valley, and the most complete is figured here.
Preserved as an internal mould it shows all the features mentioned by Harper
(1947 : 167) in his re-description of the species, though comparison with the some-
what distorted topotype figured by him (PL 6, fig. 12) is difficult. The Shropshire
specimen now illustrated is from the Dalmanella horderleyensis Zone, that is to say
only slightly earlier than the horizon yielding the type-specimen at Ynys Galed,
Caernarvonshire. Whittard (1960 : 158) has recently found the same species in west
Shropshire, where it occurs earlier, in the Soudleyan Stage.
Flexicalymene caractaci (Salter)
(PL 38, figs. 2, 4-6, 8, n, 12)
i865«. Calymene Blumenbachii var. Caractaci Salter, p. 96, pi. 9, fig. 3.
I931- Calymene caractaci Salter : Shirley, p. 25, pi. 2, figs. 4-6.
1936. Flexicalymene caractaci (Salter) Shirley, p. 395, figs, i, 2.
1949. Flexicalymene trigonoceps Bancroft, p. 306, pi. n, figs. 31, 32.
Specimens of this well-known species are common in the Caradoc Series of south
Shropshire, but records of its existence outside the district are suspect and its true
2i8 ORDOVICIAN TRILOBITE FAUNAS OF SOUTH SHROPSHIRE, III
distribution in the Anglo- Welsh area is by no means well known, though its apparent
absence is likely to result from the lack of the appropriate strata over much of Wales.
Flexicalymene caractaci was re-described by Shirley (1931 : 25) using Salter's
type-material, which comprises a large slab of highly fossiliferous, decalcified sand-
stone. The lithology is typically that of the Upper Cheney Longville Flags, of
Marshbrookian age, and this is confirmed by the fauna which includes, in addition
to F. caractaci, abundant Dalmanella wattsi (Bancroft) and Broeggerolithus transiens
(Bancroft). The specimen illustrated by Shirley (1931, pi. 2, fig. 4) and refigured
here (PI. 2, fig. 6) is now chosen as lectotype as it corresponds best with Salter's
original illustration. Little need be added to Shirley's detailed description, but it
may be noted that the anterior border is somewhat variable in length, and that its
angle of inclination may be dependent on the state of preservation. For this reason
Bancroft's Flexicalymene trigonoceps, said by him to have an anterior border less
steeply inclined than that of F. caractaci, is now considered to fall within the limits
of variation of the latter species. The earliest undoubted occurrences in south
Shropshire are in the lowest Marshbrookian Stage, the Dalmanella wattsi Zone, at
which level it is to be found in abundance at several localities, but fragmentary
evidence from the Kjaerina typa Zone of the Upper Longvillian Substage in the
Onny Valley suggests that it may have appeared there somewhat earlier. F. caractaci
is generally found wherever the strata of the Dalmanella unguis Zone crop out ; in
the topmost third of the Marshbrookian, the Onniella reuschi Zone, the species is
noticeably less common, and as yet it has been found neither in the overlying Actonian
Stage nor co-existing with species of Onnicalymene.
LECTOTYPE. GSM. 47698 (PL 38, fig. 6).
Flexicalymene cobboldi sp. nov.
(PI. 38, figs, i, 3)
DIAGNOSIS. Flexicalymene (s.s.) with relatively short, well-rounded glabella.
Anterior border long, steeply inclined frontally, and convex forwards in plan. Eyes
situated opposite second glabellar lobes and furrows.
DESCRIPTION. The cranidium is moderately convex both longitudinally and
transversely, with the maximum breadth twice the median length. The glabella is
plump and short, about as broad as long, its outline subparabolic. The frontal
glabellar lobe is small and short, only about one-seventh of the glabellar length,
broadly rounded frontally and defined posteriorly by short (tr.), shallow, first
glabellar furrows slightly divergent posteriorly, parallel to the small, ovoid, first
glabellar lobes. The second glabellar lobes are larger, subcircular in plan, about
half the length (exsag.) of the third lobes, which are of the " cat's ear " form described
by Shirley (1931 : 20, 26). The second glabellar furrows are deep, widening (exsag.)
adaxially and showing traces of branching. The third glabellar furrows are some-
what wider, deeper and distinctly bifid, the anterior branches being the shorter,
continuing across the bases of the second lobes, whilst the posterior branches end
opposite the mid-points of the third lobes, though there is a suggestion of their
continuation as shallow grooves across the neck-like structures connecting the lobes
ORDOVICIAN TRILOBITE FAUNAS OF SOUTH SHROPSHIRE, III 219
to the median glabellar lobe. The unfurrowed median lobe of the glabella is broad,
with almost no development of what Shirley (1936 : 386) termed intermediate lobes.
The axial furrows are broad, curved, gently convergent forwards, shallowing slightly
opposite the second glabellar lobes. The anterior border is long, well rounded in
plan, and of scoop-like form, steeply inclined forwards. The palpebral lobes,
incompletely known, are sited in line with the second glabellar lobes and furrows.
The remainder of the cranidium contains no diagnostic features, and is identical
with that of Flexicalymene caractaci, a closely-related species. The librigenae,
hypostoma, thorax and pygidium are not known.
HORIZONS AND LOCALITIES. The holotype is from the Upper Longvillian Substage,
Kjaerina bipartita Zone, by the south-eastern side of the road between Horderley
and Marshbrook, 420 yards north-east of the building known as Crosspipes. At this
point the specimen was collected from grey-green mudstones which are interbedded
with limestone lenses containing Heterorthis alternata (J. de C. Sowerby), the whole
being grouped together as the " Alternata Limestone ". The paratype, the only
other specimen known, is from the Marshbrookian Stage, Dalmanella unguis Zone,
at Marsh Wood Quarry, half-a-mile south of Marshbrook Station. The fossil band
containing this specimen did not yield any other specimens of Flexicalymene, though
F. caractaci has been found in abundance at other levels in the same quarry.
HOLOTYPE. BM. In. 55448 (PI. 38, fig. 3).
PARATYPE. BM. In. 50762 (PL 38, fig. i).
DISCUSSION. The stratigraphically earliest known species of Flexicalymene (s.s.)
in south Shropshire is F. acantha, the glabellar outline of which, narrowing sharply
in front of the second glabellar furrows, is remarkably suggestive of that found in
Flexicalymene cambrensis (Salter, 1865), a Llandeilo species which has been refigured
by Shirley (1931 : 20, pi. I, figs. 11-15), and the two forms may be related. However,
the anterior border of F. acantha is noticeably the shorter of the two, whilst the
palpebral lobes of F. cambrensis are situated farther forwards than in the Harnagian
species.
In the collections of the Geological Survey & Museum there are two specimens
labelled " Calymene fatua ", a manuscript name proposed by Salter but never
published. One of these specimens, numbered 19603, is a typical, though distorted,
cranidium of Flexicalymene acantha from " Horderley ". The other is an unidentifi-
able, damaged cranidium, 19604, apparently a Flexicalymene or Reacalymene,
alleged to be from " Gretton ". This locality is probably incorrect, as only strata of
Marshbrookian or Actonian age crop out in the vicinity of Gretton village, whereas
the matrix of the specimen suggests rather a derivation from the Chatwall Flags, of
Soudleyan age.
Calymene incerta Barrande (1852 : 568, pi. 19, figs. 30, 36) was stated to occur at
a number of localities in Bohemia, and the illustrated specimens show some variation
in the proportions of the glabella, so that more than one form may be present. The
species may be referred to Flexicalymene (s.s.}, and specimens in the Barrande
Collection at the British Museum (Nat. Hist.) from Zahorzan, one of the localities
listed in the original description, show the anterior border to be relatively short,
suggesting a comparison with species such as F, acantha and F. planimarginata.
220 ORDOVICIAN TRILOBITE FAUNAS OF SOUTH SHROPSHIRE, III
rather than with F. caractaci and F. declinata (see later). In general, Flexicalymene
incerta appears to be larger than any of the Anglo- Welsh species, with a glabellar
outline much broader frontally than either F. acantha or F. planimarginata, although
one of the specimens figured by Barrande (1852, pi. 19, fig. 32 only) has a glabella
reminiscent of F. acantha. According to Barrande (1852 : 569), F. incerta is accom-
panied by Trinucleus ornatus (now Onnia ornata Sternberg sp.) and Calymene pulchra
(now Pharostoma pulchrum Beyrich sp.), and is unlikely to be later than Caradoc in
age.
The species to which Flexicalymene caractaci bears, perhaps, the strongest resem-
blance is F. declinata (Hawle & Corda), described originally from the Ordovician of
Beraun (Hawle & Corda, 1847 : 87), but figured first by Barrande (1852 : 570, pi. 43,
figs. 53-58). The glabellar outline is closely similar in both species, but the eyes of
F. declinata are situated farther forwards, opposite the first glabellar lobes, whilst
the outline of the anterior border is less well-rounded than that of F. caractaci.
According to Barrande (1852 : 572), F. declinata is found in company with Remo-
pleurides radians (now Amphitryon radians Barrande sp.) and Trinucleus Bucklandi
(now Tretaspis granulata Wahlenberg sp.), species which suggest an Ashgill age.
Flexicalymene caractaci and F. cobboldi may readily be separated from the earlier
F. acantha and F. planimarginata by their longer anterior border as well as by the
glabellar outline, whilst the glabella of F. caractaci is longer than that of F. cobboldi.
The short glabella of the last-named species is not unlike that of Flexicalymene
brevicapitata Portlock sp. (Shirley, 1931 : 28, pi. 2, figs. 9, 10) from, probably, the
Killey Bridge Beds of Tyrone, but the latter lacks definite first glabellar lobes and
the eyes are situated slightly farther forwards than in the Shropshire trilobite.
Subgenus REACALYMENE Shirley, 1936
TYPE SPECIES. Reacalymene limba by original designation of Shirley (1936 : 395).
Flexicalymene ( Reacalymene) pusulosa (Shirley)
(PL 37, figs. 9, n, 12)
1936. Reacalymene pusulosa Shirley, p. 407, pi. 29, figs. 9, 10.
1958. Reacalymene pusulosa Shirley : Dean, pp. 198, 218.
1962. Flexicalymene (Reacalymene) pusulosa (Shirley) Dean, p. 113.
Numerous specimens corresponding in all details with Shirley's original description
of the species have been found in the topmost subdivision, the Harknessella sub-
quadrata Zone or Limestone, of the Costonian Stage in the Harnage District, where
it occurs at several localities. In the southern part of the Caradoc District, several
miles south of Harnage, the Costonian strata become less calcareous and more
arenaceous than at Harnage and have not yielded material which can satisfactorily
be assigned to F. (R.) pusulosa. The evidence suggests that the species has a some-
what restricted horizontal as well as vertical distribution, and that farther south it
is replaced by Flexicalymene acantha Bancroft. Although Shirley stated that the
pygidium of F. (R.) pusulosa possesses four pleural ribs and six axial rings, further
ORDOVICIAN TRILOBITE FAUNAS OF SOUTH SHROPSHIRE, III 221
collecting shows that some specimens may have five pleural ribs and traces of a
seventh axial ring.
HOLOTYPE. This specimen, a cranidium, was originally part of the E. S. Cobbold
collection, housed in the Shrewsbury Museum, but has since been transferred to the
British Museum (Nat. Hist.) where it is numbered In. 50780 (PL 37, figs. 9, n).
LOCALITIES. The holotype is from the disused quarry near the eastern end of
Black Dick's Coppice, Evenwood, whilst the other figured specimen is from the small
quarry in the north-western part of Round Nursery, south of Harnage Grange.
Flexicalymene (Reacalymene) cf. limba (Shirley)
(PL 37, fig. 16)
1936. Reacalymene limba Shirley, p. 409, pi. 29, figs. 11-13.
1960^. ? Reacalymene pusulosa Shirley : Dean, p. 167.
1961. Reacalymene limba Shirley : Whittard, pi. 22, fig. 5.
An incomplete cranidium preserved as an internal mould resembles the species
described by Shirley in all essentials, as far as can be ascertained. Shirley dis-
tinguished Reacalymene limba from R. pusulosa by the former's possession of a less
convex glabella and longer " preglabellar field " (=anterior border). He also pointed
out that R. limba lacked the so-called " supplementary " furrows on the glabella
just behind the hypostomal pits, and such furrows appear to be absent from the
present specimen. In addition it may be stated that the sides of the glabella are
more convergent forwards in F. (R.) limba than in F. (R.) pusulosa, resulting in a
distinctive, triangular appearance.
HORIZON AND LOCALITY. The so-called " Glyptocrinus Flags ", Soudleyan Stage,
probably Broeggerolithus broeggeri Zone, at the small, disused, road-side quarry
150 yards north-east of Chatwall Farm, Chatwall. Shirley (1936 : 409) believed,
though with reservations, that the holotype of F. (R.) limba originated from the
Rhiwlas Limestone, Ashgill Series, of the Bala District, but recently Whittard
(1960 : 158) has recorded the species from the Shelve Inlier, at the same horizon as
the present specimen. Despite extensive collecting the species has not been found
at any other horizon in south Shropshire, and Shirley's record (1936 : 408) of rare
Reacalymene pusulosa in the Glyptocrinus Flags may refer, in fact, to forms such as
that now figured.
Flexicalymene (Reacalymene) horderleyensis sp. nov.
(PL 37, figs. 2, 8 ; PL 40, figs. 7, 8)
?I947. Flexicalymene sp. nov., Harper, p. 168, pi. 6, fig. 10.
1958. Reacalymene sp. nov., Dean, p. 221.
DIAGNOSIS. Species of Reacalymene characterized by subtriangular glabella with
short frontal lobe, and long, steeply inclined anterior border with low, transverse
ridge.
DESCRIPTION. The species is known, so far, from only three isolated cranidia, two
of them external moulds, the other an internal mould. The glabella is slightly longer
than broad, roughly subtriangular in plan with almost straight lateral margins
222 ORDOVICIAN TRILOBITE FAUNAS OF SOUTH SHROPSHIRE, III
converging forwards at about 35 degrees to a short, broadly rounded, frontal
glabellar lobe. There are three pairs of lateral glabellar lobes, those of the basal
pair the largest, broadly oval in plan, slightly divergent forwards, and bluntly
pointed anterolaterally. The second glabellar lobes are about half the size of the
third pair, oval in plan, broadly divergent forwards and parallel to the lobes of the
first pair, which are short and subrectangular. The first glabellar lobes are separated
from the frontal glabellar lobe by short, shallow, first glabellar furrows which are
parallel to the long axes of the first lobes. The second glabellar furrows run parallel
to those of the first pair, but are deeper, bifurcating slightly so that the posterior
branch is the longer. The third glabellar furrows are more markedly bifid, the
anterior branches short, curving around the second glabellar lobes, the posterior
branches running backwards and ending just short of the occipital furrow ; the two
branches are separated by a small, convex lobe-like swelling. All the glabellar lobes
and furrows end in-line adaxially, leaving a median body which is roughly parallel
sided and occupies a little less than half the breadth of the glabella. The anterior
border is long, convex forwards medially, slightly ridged posteriorly on the external
mould (PL 37, fig. 2), though apparently smooth on the internal mould (PL 40, fig. 8) ;
it is separated from the preglabellar field by a shallow, anterior border furrow. What
may be described as the preglabellar field is short (sag.}, gently inclined forwards,
expanding laterally, truncated by the well-impressed preglabellar furrow which is
continuous with the deep, narrow axial furrows. The occipital furrow is shallow and
transversely straight medially, deepening laterally, at the same time becoming
convex backwards as it circumscribes the third glabellar lobes. Similarly,
the occipital ring is transversely straight and of uniform length (sag.) medially,
shortening laterally where it forms a pair of poorly defined occipital lobes, strongly
divergent forwards, and indistinctly separated from the posterolateral parts of the
fixigenae. The posterior border furrow is broad (exsag.), straight, of uniform and
moderate depth, delimiting a posterior border which is narrow (exsag.) adaxially,
broadening towards the genal angles. The anterior portions of the fixigenae are of
uniform breadth, almost half that of the frontal glabellar lobe, and parallel-sided as
far as the palpebral lobes, whilst the posterior portions are steeply declined. The
palpebral lobes are laterally convex in plan, situated opposite the second glabellar
lobes, with poorly-defined eye-ridges running from the anterior ends of the lobes to
end at the axial furrows opposite the first glabellar lobes. The facial suture is of
characteristic calymenid type, the anterior branches converging gently from the
eyes to cut the anterior margin at points which lie longitudinally between the
palpebral lobes and the axial furrows, whilst the posterior branches curve backwards
from the eyes, at first gently and then more strongly to the genal angles. The surface
of the test is poorly known from only two external moulds, but appears to be finely
granulate.
The librigenae, hypostoma, thorax and pygidium are not known.
HORIZON AND LOCALITIES. The holotype and one paratype are from the Lower
Longvillian Substage, zone of Dalmanella indica and D. lepta, at the outcrop inside
the south-eastern corner of Rookery Wood, south-east of Horderley. The remaining
paratype is from a similar horizon at the small disused quarry just west of the
ORDOVICIAN TRILOBITE FAUNAS OF SOUTH SHROPSHIRE, III 223
western end of Rookery Wood. Fragmentary evidence of what may prove to be
the same species has been obtained from the underlying Dalmanella horderleyensis
Zone, at Long Lane Quarry, three-quarters of a mile north-west of Craven Arms.
HOLOTYPE. BM. In. 50656 (PI. 37, figs. 2, 8).
PARATYPES. BM. In. 49559, BM. In. 52103 (PI. 40, figs. 7, 8).
DISCUSSION. Flexicalymene (Reacalymene] horderleyensis bears a general resem-
blance to F. (R.) limba, from which it is distinguished by its longer, more steeply
upturned anterior border, and shorter frontal glabellar lobe. In the Welsh Borders
the known species of Reacalymene, namely pusulosa, limba and horderleyensis, derive
successively from Costonian, Soudleyan and Lower Longvillian strata, and form a
morphological series, with a progressive increase in the length of the anterior border,
and in the degree of convergence of the axial furrows.
The internal mould of a calymenid cranidium described and figured by Harper
(1947 : 168, pi. 6, fig. 10) as Flexicalymene sp., from the Lower Longvillian of Ynys
Galed, North Wales, has apparently undergone some tectonic deformation, but
nevertheless bears a general resemblance to F. (R.) horderleyensis, particularly in the
glabellar outline and long anterior border (compare PI. 4, fig. 7). The two may be
synonymous, but a final decision must await the collection of additional Welsh
material.
Calymene croneisi Roy, 1941, from the Ordovician, possibly Trenton, of Baffin
Island, was recently redescribed by Whittington (in Miller et al., 1954 : 147, pi. 62,
figs. 13-15) and assigned to Flexicalymene, though he noted that the anterior
border is ridged as in Reacalymene. Flexicalymene (Reacalymene} croneisi bears a
general resemblance to the Anglo-Welsh species, but is particularly distinguished by
the forward position of the palpebral lobes, opposite the second glabellar furrows
and part of the first glabellar lobes. The anterior border of the North American
form, like that of F. (R.) pusulosa, is relatively short and transversely straight,
which feature easily separates it from F. (R.) horderleyensis, whilst the glabellar
outline is narrower than that of F. (R.) pusulosa.
Genus ONNICALYMENE Dean, 1962
TYPE SPECIES. Flexicalymene onniensis Shirley, 1936 by original designation of
Dean (1962 : 112).
Onnicalytnene onniensis (Shirley)
(PL 39, figs. 2, 10, n; PI. 40, fig. 9)
1865. Calymene brevicapitata Portlock, Salter, pi. 9, fig. 7.
1936. Flexicalymene onniensis Shirley, p. 405, pi. 29, figs. 5-7.
1949. Flexicalymene onniensis Shirley : Bancroft, p. 308, pi. u, figs. 36, 37.
1958. Flexicalymene onniensis Shirley : Dean, p. 224.
!959- Flexicalymene onniensis Shirley : Dean, pp. 200, 207.
1959- Flexicalymene aff. onniensis Shirley : Dean, pp. 202, 208.
1962. Onnicalymene onniensis (Shirley) Dean, p. 115, pi. 13, figs. 7; pi. 14, figs, i, 2, 7, 10.
The species was first described by Shirley (1936) from the Onnia gracilis Zone of
the Onnian Stage in the River Onny Valley. The most likely site of the type-locality
224 ORDOVICIAN TRILOBITE FAUNAS OF SOUTH SHROPSHIRE, III
is the prominent, ridge-like outcrop in the river-bed about 100 yards of its junction
with the stream of Batch Gutter, and at this point 0. onniensis occurs abundantly
with the zonal trinucleid. There is ample evidence that the species ranges upwards
into and through the succeeding Onnia superba Zone, but evidence of its earlier
occurrence is tenuous. Possible fragments have been found in the Onnia ? cobboldi
Zone of the Onny Valley, but all the identifiable specimens of Onnicalymene so far
collected from the underlying Actonian Stage have proved to belong to other species.
Outside Shropshire 0. onniensis occurs at Welshpool, in the Onnia gracilis Zone,
and in the Dufton Shales of the Cross Fell Inlier, where it has an extended vertical
range upwards into the Pusgillian Stage (Dean, 1962 : 115).
HOLOTYPE. GSM. RR 1940 (PL 39, figs. 10, n).
Onnicalymene laticeps (Bancroft)
(PL 38, figs. 7, 10, 14, 15)
1949. Flexicalymene laticeps Bancroft, p. 307, pi. n, figs. 33, 33^.
1958. Flexicalymene laticeps Bancroft : Dean, p. 224.
1962. Onnicalymene laticeps (Bancroft) Dean, p. 115, pi. 14, figs. 5, 6.
This small, easily-recognized species of Onnicalymene has been found at several
localities in Shropshire, in both mudstones and sandstones of the Actonian Stage.
0. laticeps has been recorded also in the Cross Fell Inlier (Dean, 1962 : 115), and
may prove to be of stratigraphical value. It is readily separated from other species
of the genus by its short, well-rounded glabella.
HOLOTYPE. BM. In. 42103 (PL 38, fig. 7).
Onnicalymene salteri (Bancroft)
(PL 38, figs. 9, 13)
1949. Flexicalymene salteri Bancroft, p. 306, pi. n, figs. 34, 35.
1958. Flexicalymene salteri Bancroft : Dean, p. 224.
1962. Onnicalymene salteri (Bancroft) Dean, p. 113.
This species was founded on two cranidia, both of Actonian age, one, the holotype,
from the northern end of the small wood west of Rose Villa, Marshbrook, the other
from the locality known as Jack Slither, in the south bank of the River Onny,
45 yards west of its junction with the stream of Batch Gutter. Both specimens
occur in grey, calcareous mudstones, a lithology which contains most of the known
specimens, though a few have been collected from limestone bands of the Actonian
in the vicinity of Jack Slither. Onnicalymene salteri has not yet been found in the
arenaceous facies of the Actonian, such as is found at and near Acton Scott and
Cardington, nor is it known to occur at any horizon other than the Actonian. It is
subordinate in numbers to the contemporaneous Onnicalymene laticeps (Bancroft),
and has not been recorded outside Shropshire.
HOLOTYPE. BM. In. 42100 (PL 38, fig. 9).
PARATYPE. BM. In. 42099.
ORDOVICIAN TRILOBITE FAUNAS OF SOUTH SHROPSHIRE, 111 225
Genus GRAVICALYMENE Shirley, 1936
TYPE SPECIES. Gravicalymene convolva by original designation of Shirley (1936 :
395).
Gravicalymene praecox (Bancroft)
(PL 39, figs, i, 3, 9, 12-14)
1949. Diacalymene praecox Bancroft, p. 308, pi. n, figs. 28, 280,.
1958. Diacalymene ? praecox Bancroft : Dean, p. 219.
1962. Gravicalymene praecox (Bancroft) Dean p. 113.
The holotype of Gravicalymene praecox, like the lectotype of Flexicalymene acantha,
from the same locality and horizon, is a small immature cranidium (PI. 39, figs. 13,
14). The photograph used by Bancroft (1949, pi. n, fig. 28) to illustrate the species
was taken obliquely from the right-hand side of the specimen ; consequently, what
might appear to be a constriction of the right axial furrow is really an illusory effect
produced by the convexity of the right fixigena in line with the axial furrow. In
fact, the axial furrows of the holotype show no lateral constriction, and the species
cannot, therefore, be placed in Diacalymene. Both Diacalymene and Gravicalymene
have recently been discussed (Dean, 1962) and it has been shown that the earlier
usage of the terms " ridged " and " roll-like " with regard to the form of the anterior
border is unsatisfactory as a basis for generic differentiation, but that the form of
the axial furrows is more reliable. On these grounds, therefore, Diacalymene praecox
must be transferred to Gravicalymene. During recent collecting, additional topotype
material has been obtained which apparently represents mature adult specimens of
the species (PI. 39, figs, i, 3, 9, 12). All are internal moulds of incomplete cranidia,
and they show little change from the holotype, though the granulation of the test
found on the latter specimen seems to have disappeared when the adult stage was
attained.
A cranidium from the Actonian Stage at Gretton, GSM. 19588, constituted one
of the syntypes of Salter's (1865^, pi. 9, fig. 4 only) Calymene Blumenbachii var.
Caractaci whilst another cranidium from the same locality was assigned to the
same species by La Touche (1884, pi. 3, fig. 61). These stratigraphically later forms
of Gravicalymene are undoubtedly extremely close to G. praecox, with which species
they are compared here, though identification cannot be certain in the absence of
complete, uncrushed, adult cranidia of the Harnagian species. The thorax, of
characteristic calymenid form, with thirteen thoracic segments, was found in
juxtaposition to both cranidium and pygidium in strata of Onnian age (PL 39,
figs. 4, 5, 8). The pygidium is strongly convex both longitudinally and transversely,
the dorsal surface of the axis being gently convex, steeply declined posteriorly.
The outline of the pygidium, when viewed from above with the axis horizontal, is
broadly semi-elliptical frontally, the posterolateral margins being straight, strongly
divergent forwards as far as the line of maximum breadth, opposite the midpoint
of the pygidium. The axis is broadest frontally, where it is equal to one-third the
maximum pygidial breadth, but narrows a little backwards, and stops short of the
the tip of pygidium. Excluding the articulating half-ring, there are seven well-defined
axial rings, and the axial furrows are straight, slightly convergent backwards where
226 ORDOVICIAN TRILOBITE FAUNAS OF SOUTH SHROPSHIRE, 111
they are continued so as to circumscribe the terminal piece, which is of moderate
size and a little less than one-quarter the length of the axis. The pleural lobes are
strongly arched-down laterally, with six pairs of well-defined, evenly curved ,pleural
ribs, and a seventh, less well-defined pair fused together to form a low, postaxial
ridge. Each pleural rib is divided into two bands of almost equal breadth (exsag.)
by a narrow, shallow, interpleural furrow which cuts the posterior margin of the
rib at its intersection with the axial furrow.
HORIZONS AND LOCALITIES. Gravicalymene praecox was first described from the
Smeathen Wood Beds, Harnagian Stage, Reuscholithus reuschi Zone, exposed in the
old cartway at the southern end of Smeathen Wood, Horderley. What is probably
the same form has been found, rarely, at the corresponding horizon in the north
bank of Coundmoor Brook, about 1,295 yards south-west of Harnage Farm,
Harnage. As noted earlier, specimens referred to Gravicalymene cf. praecox have
been collected from both the Actonian and Onnian Stages. Actonian localities
include the old quarry 210 yards east of Acton Scott church, and the now unavailable
quarry at Quarry Field, Gretton, near Cardington (see PL 39, fig. 7). The only
Onnian locality at which G. cf. praecox has been found is in rocks of the Onnia
gracilis Zone, exposed in the north bank of the River Onny 100 yards east of its
junction with Batch Gutter. These occurrences, at widely separated horizons within
the Caradoc Series, are particularly interesting as no evidence of Gravicalymene has
been found in the intervening strata. It may be noted, however, that in each case
the appearance of the genus is associated with an influx of new f aunal elements, and
this, in turn, may be correlated with a marine transgression which affected much of
the Anglo-Welsh area. The lower of these is the Nemagraptus gracilis Transgression,
which continued into the overlying Diplograptus multidens Zone ; the higher trans-
gression is that occurring high in the Dicranograptus clingani Zone, and perhaps also
in the Pleurograptus linearis Zone, which gave rise to dark shales and mudstones,
including the Nod Glas, in North Wales.
HOLOTYPE. BM. In. 42090 (PL 39, figs. 13, 14).
DISCUSSION. Of the other recorded species of Gravicalymene in the British
Ordovician, G. jugifera Dean (1962, pi. 13, figs. 9, n ; pi. 14, figs. 3, 4, 8, 9) from
the Pusgillian Stage of the Cross Fell Inlier differs from G. praecox in having pro-
portionately smaller basal glabellar lobes, a less quadrate frontal glabellar lobe, and
an anterior border which is separated from the glabella by a broader (sag.), more
conspicuous furrow. Gravicalymene convolva Shirley (1936 : 409, pi. 29, figs. 16-18)
is a relatively large species which may be distinguished from G. praecox by its
broader glabellar outline, with slightly smaller basal glabellar lobes and conspicuously
shorter frontal glabellar lobe.
The trilobite figured by St0rmer (1945, pi. 2, figs. 6-8) as Reacalymene holtedahli
from Stage 4ca of Hadeland, Norway (an horizon approximately equivalent to the
Pusgillian Stage) is probably best placed in Gravicalymene or Diacalymene. It may be
distinguished from G. praecox by its narrower glabellar outline, smaller glabellar lobes,
and shorter frontal glabellar lobe. One of St0rmer's syntypes appears to show the
second glabellar lobes in contact with the fixigenae, but, judging from the distortion
of the specimen, this may well be due to mechanical causes. The pygidium of the Nor-
ORDOVICIAN TRILOBITE FAUNAS OF SOUTH SHROPSHIRE, ill 227
wegian form has fewer axial rings and pleural ribs than that described in the present
paper as Gravicalymene cf. praecox.
It was claimed by Bancroft (1949 : 309) that what he called Diacalymene praecox
was important in anticipating a form of Ashgill age, probably D. marginata Shirley,
judging from his text. Now that D. praecox can be assigned to Gravicalymene, such
a claim is without foundation, and the species probably represents nothing more
than one member of a long-ranging genus. The trilobite figured by Shirley (1931,
pi. 2, fig. n) as Calymene quadrata ? King, from the Llandeilo Flags of South Wales,
is too badly preserved for certain identification, but bears some resemblance to
Gravicalymene, and suggests that the genus may have appeared in the Anglo- Welsh
area before the beginning of Caradoc times. Gravicalymene had a remarkably long
stratigraphical range, and has been recorded as late as the Devonian from New
Zealand (Shirley, 1938).
Gravicalymene inflata sp. nov.
(PI. 39, fig. 6)
1958. Gravicalymene sp. nov., Dean, p. 224.
DIAGNOSIS. Species of Gravicalymene with short (sag.), transversely straight,
anterior border. Frontal glabellar lobe moderately long, markedly quadrate in plan.
Fixigenae strongly convex, almost as anteriorly broad as frontal glabellar lobe.
Palpebral lobes sited opposite second glabellar furrows and anterior half of second
glabellar lobes.
DESCRIPTION. The only available specimen is an incomplete cranidium, lacking
the posterior halves of the fixigenae, but with the test preserved. The median length
is about 15 mm., and the frontal breadth, as measured across the anterior portion
of the fixigenae, is 17 mm.
The glabella is slightly longer than broad, approximately in the ratio 6:5, but
the proportions are somewhat distorted by crushing of the basal glabellar lobes,
which constitute the line of greatest breadth. The frontal glabellar lobe is markedly
rectangular in outline, its breadth more than twice the length. There are three pairs
of glabeUar lobes, those of the third pair being the largest, becoming subangular
anterolaterally. The second glabellar lobes are considerably smaller than those of
the third pair, oval in shape with their long axes gently divergent forwards. The
third lobes are of subcircular form, about half the size of the second lobes ; they
are defined anteriorly by short (tr.), transversely straight, first glabeUar furrows
which widen (exsag.) slightly adaxially. The second glabellar furrows are at first
transversely straight from the axial furrows, but quickly bifurcate, the anterior
branch so formed being short, and the posterior branch longer, extending backwards
until opposite the mid-points of the second glabellar lobes. The third glabellar
furrows extend backwards from the axial furrows for almost the length (tr.) of the
second glabellar lobes before bifurcating, the shorter, anterior branch then running
forwards towards, but not meeting, the posterior branch of the second glabeUar
furrows. The posterior branches of the third glabellar furrows are longer (exsag.),
and appear to end opposite the middle of the third glabellar lobes but, as the latter
are partly crushed, this feature cannot be properly examined. The glabellar lobes
228 ORDOVICIAN TRILOBITE FAUNAS OF SOUTH SHROPSHIRE, III
are thus connected to the median glabellar lobe by constricted, neck-like structures
which, as far as can be seen, are slightly depressed dorsally, though there is no
development of definite furrows. The axial furrows are narrow, uniformly deep,
almost parallel frontally as far as the second glabellar furrows ; they then diverge
slightly as far as the third glabellar furrows, beyond which they diverge more
strongly, curving round the abaxial margins of the basal glabellar lobes to intersect
the occipital furrow. The anterior border is conspicuously short (sag.}, one-seventh
of the median length of the glabella, becoming even shorter abaxially, beyond the
axial furrows ; the anterior margin is transversely straight, as is the posterior margin
opposite the frontal glabellar lobe, though there is a slight lengthening (exsag.)
opposite the axial furrows. The flattened upper surface of the anterior border is
inclined forwards at only a small angle to the dorsal surface of the glabella, that is
to say, it would be approximately horizontal when the cranidium was in its, pre-
sumably, normal position. The anterior border and frontal glabellar lobe are
separated by a deep, narrow (sag.}, transversely straight furrow which curves slightly
forwards abaxially beyond the axial furrows to separate the anterior border from
the fixigenae. The occipital ring is relatively long (sag.}, in length about one-sixth
that of the glabella, parallel-sided medially, but shortening (exsag.} abaxially to
accommodate the projecting posterior margins of the basal glabellar lobes, and
forming a pair of poorly-defined occipital lobes. The occipital furrow is both shallow
and transversely straight medially, but deepens distally, becoming convex back-
wards around the basal glabellar lobes. The pleuroccipital furrow, posterolateral
portions of the fixigenae and the posterior branches of the facial suture, are not
preserved. The palpebral lobes are of moderate size, inclined gently towards the
lateral margins, and extending from opposite the second glabellar furrows to opposite
the third glabellar furrows. The fixigenae are strongly convex both transversely
and longitudinally, standing highest opposite the second glabellar lobes. They are
parallel-sided frontally, the anterior branches of the facial suture running straight
forwards from the eyes, and there is a pair of poorly defined eye-ridges extending
slightly forwards adaxially from the anterior ends of the palpebral lobes and ending
at the axial furrows opposite the first glabellar lobes. The surface of the test,
excluding the furrows, all of which are smooth, is covered with fine, closely-set
granules, generally of uniform size ; slightly larger granules occur sporadically, and
uncommonly, over the surface of the glabella and fixigenae, becoming more common
on the anterior border.
The hypostoma, thorax and pygidium are not known.
HORIZON AND LOCALITY. The only known specimen is from the Onnian Stage,
Onnia gracilis Zone, in the north bank of the River Onny, 100 yards east of its
junction with Batch Gutter.
HOLOTYPE. BM. In. 50653 (PI. 39, fig. 6).
DISCUSSION. This rare form is distinctive, and may easily be separated from all
other known species of the genus, for example G. praecox, by the unusually short
(sag.), straight anterior border, and by the large, inflated fixigenae, which
are much broader than those of other forms. In addition, the palpebral lobes are
situated farther forwards than those of G. praecox.
ORDOVICIAN TRILOBITE FAUNAS OF SOUTH SHROPSHIRE, III 229
Family ASAPHIDAE Burmeister, 1843
Subfamily ASAPHINAE Burmeister, 1843
Recently Parabasilicus was placed by Jaanusson (in Moore, 1959 : 342) in the
subfamily Isotelinae, whilst Basilicus was assigned to the Asaphinae. The affinities
of Parabasilicus appear to lie with Basilicus rather than with the other genera of
the Isotelinae, and accordingly the genus is here transferred to the Asaphinae.
Genus PARABASILICUS Kobayashi, 1934
TYPE SPECIES. Parabasilicus typicalis by original designation of Kobayashi
(1934 •• 475)-
Parabasilicus powisi (Murchison)
(PL 40, figs, i, 3-5 ; PL 41, figs, i, 2 ; PL 42, fig. 9)
1839. Asaphus powisi Murchison, p. 661, pi. 23, fig. gc only.
1851. Isotelus (Basilicus) Powisii (Murchison) M'Coy in Sedgwick & M'Coy, p. 170.
1866. Asaphus (Basilicus) powisi Murchison : Salter, p. 154, pi. 23, figs. 3-7.
1931. Asaphus powisi Murchison : Reed, p. 443.
1934. Parabasilicius powisi (Murchison) Kobayashi, p. 476.
1935. Asaphus (Parabasilicus) powisi Murchison : Reed, p. 13.
1937. Parabasilicus powisi (Murchison) : Kobayashi, p. 503.
1938. Asaphus (Parabasilicus) powisi Murchison : Stubblefield in Pocock et al., pp. 89, 255.
1958. Parabasilicus powisi (Murchison) : Dean p. 220.
1961. Parabasilicus powisi (Murchison) : Dean & Dineley, p. 374, pi. 20, fig. 8.
DESCRIPTION. The species was described first by Murchison (1839 : 661) on the
basis of two syntypes. One of these (Murchison, 1839, pi- 23> n§s- 9a> b) nas been
shown to be a cephalon of Chasmops extensa (Boeck) and the other syntype (Murchi-
son, 1839, pi. 23, fig. gc) has been chosen as lectotype (Dean & Dineley, 1961 : 374).
The latter specimen is a flattened, large pygidium with six attached thoracic seg-
ments, preserved as an external mould and refigured here as a vinyl plastic cast
(PL 41, fig. 2). The pygidium is large, with frontal breadth 85 mm. and median
length 52 mm., though it has been both vertically compressed and slightly sheared
sinistrally. The anterior margin is gently convex forwards, whilst the remainder of
the outline is broadly subparabolic. The axis is triangular in plan, though this has
undoubtedly been exaggerated by crushing, bounded laterally by poorly-defined,
straight, axial furrows which are hardly more than broad, shallow depressions.
Frontally the axis occupies roughly one-third of the total glabellar breadth, and its
anterior third carries traces of three large axial rings : beyond these the axis is
virtually smooth, and ends in a slightly better-defined terminal piece, separated
from the margin by a long (sag.) posterior border. The pleural lobes also have a
broad, gently concave, smooth border, inside which the pleural fields are slightly
convex dorsally with seven pairs of shallow, poorly-defined pleural furrows becoming
progressively fainter posteriorly, the furrowed area extending only for about two-
thirds of the length of the axis. Each pleural lobe has a large, steeply downturned,
anterolateral facet ; on the lectotype, only the left facet is preserved, slightly
distorted by crushing.
GEOL. 7, 8 17
230 ORDOVICIAN TRILOBITE FAUNAS OF SOUTH SHROPSHIRE, III
The thorax of the lectotype retains only six thoracic segments. Each axial ring
is relatively long (sag.), transversely straight as far as the poorly-defined axial
furrows. What appears, superficially, to be another pair of longitudinal furrows is
found on the lectotype (PL 41, fig. 2), extending along the thorax and intersecting
each axial ring at the points where it is curved backwards. This inner pair of
" furrows " is not present on uncrushed specimens assigned to the species, although
it is known from individuals which have undergone distortion, and is believed to
be of tectonic origin. The thoracic pleurae are directed only slightly backwards
distally, and end in blunt tips ; the latter are not preserved on the lectotype, but
on other specimens each can be seen to form a small, spine-like posterolateral process
which, presumably, functioned as a stop during enrollment. There is a well-defined
pleural furrow on each pleura, running from the anterior margin at the axial furrow
towards, but not reaching, the posterolateral part of the pleural tip. Immediately
in front of this furrow is a paraUel ridge which reaches the tip, though in a diminished
form, and beyond the ridge the anterior band of the pleura is turned down antero-
laterally, again a feature which was probably functional in enrollment.
No topotype cephalon is yet available, and the following description is founded
on a cranidium from Pontesford, east Shropshire, which retains the full asaphid
complement of eight thoracic segments, and has one associated librigena (PL 41,
fig. i). This individual shows the cranidium to be of depressed form, only gently
convex both longitudinally and transversely, with straight posterior margin. The
glabella is longer than broad in the ratio 4 : 3 and, although the frontal glabellar
lobe is moderately well defined anteriorly but with no incised preglabellar furrow,
the remainder of the glabella is poorly defined, particularly posteriorly, where it is
continuous with the fixigenae and occipital ring. There are no glabellar furrows and
the axial furrows are practically obsolete, apart from poorly defined depressions
delimiting the lateral margins of the frontal glabellar lobe. The anterior border
forms a brim-like structure, its anterior margin moderately convex forwards and its
dorsal surface smooth, flattened or slightly concave. The palpebral lobes are strongly
curved, placed opposite the middle of the glabella, their dorsal surface flat and
continuous with that of the glabella. The anterior branches of the facial suture
diverge forwards from the eyes for about half their length, but then curve strongly
and evenly inwards to cut the anterior border at an acute angle, longitudinally in-
line with the palpebral lobes. The posterior branches are moderately curved back-
wards from the eyes, and cut the posterior margin of the cephalon midway between
the axial furrows and the lateral margins. One displaced librigena has been found
in association with the cranidium (PL 41, fig. i), though the visual surface of the
eye is too badly damaged for examination. There is a broad, flat platform circum-
scribing the eye, and from this the remaining dorsal surface of the librigena declines
gently to the margin, which is slightly concave, though there is no marginal furrow.
The librigena is produced posterolaterally to form a stout, sharp, librigenal spine
which extended originally as far as the mid-point of the thorax. None of the available
specimens shows any ornamentation of the surface of the test, with the exception of
the distal portions of the thoracic pleurae which carry terrace-lines between the
pleural tips and the fulcrum. There is no trace of a median tubercle on the glabella,
ORDOVICIAN TRILOBITE FAUNAS OF SOUTH SHROPSHIRE, III 231
though the exoskeleton has always been found damaged at the point where such a
tubercle might be expected to occur.
No hypostoma has yet been recorded from the type-locality, nor has one been
found in association with undoubted specimens of Parabasilicus powisi. However,
a specimen, assigned doubtfully to the species, is figured here from the middle
Costonian of the Coston district (PL 40, fig. 2). It is preserved as an internal mould,
lacks the anterior wings, and is longer than broad in the ratio 6:5. The median
body is gently convex both longitudinally and transversely, sub-ovate in plan,
longer than broad, with a pair of well-defined maculae sited posteriorly, a short
distance either side of the axial line. The posterior border is indented by a deep,
parabolic notch which extends forwards as far as the median body and is flanked
posterolaterally by a pair of broad points. From these points the flattened lateral
border runs forwards in a curve, at first expanding and then contracting to meet
the sides of the median body just in front of centre.
LECTOTYPE. GSM. 6841 (PL 41, fig. 2).
HORIZONS AND LOCALITIES. The lectotype of Parabasilicus powisi was obtained
from Trilobite Dingle (= Bron-y-Buckley Wood), Welshpool, and is therefore almost
certainly of Harnagian age ; this tends to be confirmed by the state of preservation,
which resembles that of the so-called Trilobite Dingle Shales. In south Shropshire,
the earliest example of the species is from the Costonian Stage of the Evenwood
district (PL 40, fig. i) ; this is a smaller specimen than the lectotype, but shows
all the specific characters, as far as can be judged, and closely resembles a specimen
figured by Salter (1866, pi. 23, fig. 5) from an unspecified horizon at Waterloo Bridge,
Conway. Asaphid remains are uncommon in the Harnagian strata of south Shrop-
shire, but a fragmentary cranidium (PL 42, fig. 9) and pygidium, B.M. In. 55371,
from the Reuscholithus reuschi Zone of Smeathen Wood, Horderley, can probably
be assigned to P. powisi, as also can a relatively small cranidium and pygidium from
the same horizon at Coundmoor Brook, Harnage (PL 40, figs. 4, 5).
The species Asaphus (Basilicus) marstoni was founded by Salter (1866 : 156, pi. 23,
figs. la, b) on a small damaged dorsal shield preserved as an internal mould, together
with the damaged right librigena of a slightly larger specimen, both from the
"Shales of Horderly ". The syntypes are refigured here (PL 42, figs, i, 6, 8).
Although Reed (1931 : 470) claimed that the species was better referred to the
genus Proetus, there can be little doubt that it is, as Salter believed, an asaphid.
Reed's assertion that the specimen represents an immature individual with less than
the usual proetid quota of thoracic segments seems to be incorrect, as the eight
segments present are typically asaphid in both form and number. The cephalon is
damaged but shows nevertheless a general resemblance to that of Parabasilicus
Powisi, as do the pleural lobes of the thorax, though the thoracic axis is more
convergent posteriorly than that of the latter species. The pygidium resembles that
of P. powisi in outline but the axis is better defined and the pleural fields of the
pleural lobes are more strongly convex, with at least six pairs of moderately defined
pleural furrows, though the number of both these, and the axial rings, present is
obscured by abrasion and crushing. In the absence of additional material it seems
advisable, for the present, to restrict the name Parabasilicus ? marstoni to the type-
232 ORDOVICIAN TRILOBITE FAUNAS OF SOUTH SHROPSHIRE, III
specimens. The preservation of the type-material matches that of the Smeathen
Wood Beds, belonging to the Harnagian Stage, of the Onny Valley near Horderley.
The only other asaphids known from these strata are probably referable to Para-
basilicus powisi, and P.? marstoni may well prove to be a synonym of the former
species. The apparent differences could well be explained by variations in preserva-
tion or in the stage of development, and the more distinct furrowing of the pygidium
of P.? marstoni may have disappeared by the time the adult stage was attained.
Evidence of P. powisi from the Lower Soudleyan, Broeggerolithus broeggeri Zone,
is scanty, but includes a large pygidium and thorax from Glenburrell Farm (PL 40,
fig. 3) as well as the cephalon and thorax already described from Habberley Brook,
Pontesford. In the succeeding Upper Soudleyan and Lower Longvillian strata,
asaphid fragments, especially pygidia, are not uncommon. The pygidium and
hypostoma from " Horderley ", figured by Salter (1866, pi. 23, figs. 4, 6) as Asaphus
(Basilicas) Powisi, are probably from the Horderley Sandstone. The hypostoma,
now refigured (PI. 42, fig. 2), is proportionately narrower than the Costonian specimen
questionably referred here to P. powisi (PL 40, fig. 2), though bearing a general
resemblance to the latter. None of the pygidia from these younger horizons may be
satisfactorily matched with that of P. Powisi, and they (PL 40, fig. 6 ; PL 42, fig. 5),
together with the hypostoma mentioned, are merely referred, with some uncertainty,
to Parabasilicus. In general, the Longvillian pygidia, as well as being of slightly
different outline, tend to exhibit more ring and pleural furrows than P. powisi ; in
this respect they resemble the pygidium of Parabasilicus ? marstoni, already dis-
cussed, but until more satisfactory material becomes available specific identification
is virtually impossible. The only evidence of asaphid trilobites from the Upper
Longvillian of Shropshire is an unidentifiable fragment from the Alternata Lime-
stone, and the family is unknown from all the succeeding strata.
The distribution of Parabasilicus powisi and allied forms in the south of Shropshire
is of particular interest in that the genus is found most commonly elsewhere in the
Chikunsan Beds of South Korea, from which area the type species was described
by Kobayashi (1934 : 475). However, another asaphid genus, Basilicus, though
best known from the Llandeilo Series of Wales, occurs also in Korea (Kobayashi,
1934 : 465). Elsewhere, trilobites assigned to Parabasilicus are recorded from both
Bolivia and Virginia (Kobayashi, 1937 : 503 ; Raymond, 1925 : 85-86), though the
former record has been questioned by Harrington and Leanza (1957 : 36), who
tentatively report the genus from Argentina. Most of the Shropshire occurrences of
P. powisi are in strata correlated with the Diplograptus multidens Zone, though
known from the top of the preceding Nemagraptus gracilis Zone, and extending
upwards at least a little way into the succeeding Dicranograptus clingani Zone.
According to Kobayashi (1934 : 336) the Chikunsan Beds are of Llandeilo age, but
as they contain Diplograptus (Amplexograptus) perexcavatus Lapworth, a species said
to be common in the Llandeilo and early Caradoc Series of Great Britain (Elles &
Wood, 1914 : 521), both they and the Korean species of Parabasilicus may be of an
age comparable to those of Shropshire. The form most resembling P. powisi, as
noted by Kobayashi (1934 : 480) is that described by him as Parabasilicus shirakii ;
the differences between the two are small, the thoracic axis of P. shirakii being the
ORDOVICIAN TRILOBITE FAUNAS OF SOUTH SHROPSHIRE, III 233
narrower, whilst the palpebral lobes are sited farther forwards than in P. powisi.
Although several specific names were erected by Kobayashi for the material of
Parabasilicus from Korea, it seems likely that only one or two species are, in fact,
present, most of the apparent differences being the result of tectonic distortion or
intraspecific variation.
The asaphid trilobite from the Richmond Formation of Ohio described by Foerste
(1919 : 65, pis. 14, I4A, 15, especially 15) as Isotelus brachycephalus is a large form
resembling Parabasilicus in some respects, including the anterior facial suture, the
shape of the librigenae, the thorax, and the pygidial outline. However, the eyes are
situated farther forwards, and the axial and pleural furrows of the pygidium are
even less well defined than in the British species.
Family ILLAENIDAE Hawle & Corda, 1847
Genus ILLAENUS Dalman, 1827
TYPE SPECIES. Entomostracites crassicauda Wahlenberg, 1821 by subsequent
designation of Miller (1889 : 550).
Illaenus cf. fallax Holm
(PL 42, figs. 3, 4, 12)
1958. Illaenus sp., Dean, pp. 211-213, 224-
Several fragments of illaenid trilobites have been recovered from strata of Actonian
and Onnian age in south Shropshire. They are often damaged and distorted, but a
few of the best preserved are now figured. No undamaged cranidium has been
found, and it has not proved possible to make a firm identification of most of those
found (PI. 42, figs. 7, 10, n). Two pygidia, however, are undistorted and show
certain distinctive features (PL 42, figs. 3, 4). The outline is well rounded, subpara-
bolic, the median length about three-quarters of the maximum breadth, attained
just behind the anterior margin. The axis, occupying about one-quarter of the
frontal breadth, is triangular in outline, scarcely defined by faint axial furrows which
converge backwards at 50 degrees to meet just in front of centre of the pygidium,
and continue back as a single faint furrow for about half the distance to the posterior
margin, finally being replaced by an extremely low, thin, median ridge which runs
almost to the margin. The dorsal surface of the axis, which bears traces of four or
five axial rings, is slightly depressed or level, whilst the surface of the pygidium
outside the axial furrows is at first level and then declines fairly steeply towards
the margins. The front of the axis is marked by a moderately deep, transversely
straight, articulating furrow which turns backwards slightly as it crosses the axial
furrows, and then becomes both deeper and markedly broader (exsag.), finally
terminating at about the proximal margin of the doublure. Immediately in front
of this furrow each pleural lobe carries a pronounced, broad (exsag.) ridge, bordered
frontally by a large, steeply downturned, anterolateral facet.
Illaenus fallax was described by Holm (1882 : 82, pi. 2, figs. 11-20, pi. 5, figs. 15-
24) using a number of syntypes stated to be from the " Chasmopskalk " of Sweden ;
234 ORDOVICIAN TRILOBITE FAUNAS OF SOUTH SHROPSHIRE, III
several localities are mentioned in his account, and two or more horizons are almost
certainly involved. One Shropshire cranidium (PL 42, fig. 12) is generally similar
to that of I. fallax, whilst pygidia figured here bear a remarkably close resemblance
to one figured by Holm (1882, PL 2, fig. 17) from Gullerasen, a specimen which
possesses also the characteristic axial outline, followed by a median groove and
ridge. Thorslund (1940 : 140-141) recorded Illaenus fallax from different localities
and horizons in Sweden, but he stated that the species occurs most abundantly in
the Kullsberg Limestone of Dalecarlia, an horizon generally contemporaneous with
those containing the Shropshire specimens.
Although Illaenus fallax has been supposed by Jaanusson (1954 : 574) to belong
to what he calls the " Parillaenus-Gruppe " of Illaenus s.s., some of the specimens
which have been assigned to the species bear a certain resemblance to the genus
Stenopareia Holm, 1886, and Stenopareia camladica has recently been described
from the Soudleyan Stage of the Shelve Inlier by Whittard (1961 : 216, pi. 30,
figs. 10-13).
HORIZONS AND LOCALITIES. Specimens of Actonian age have been found beside
the River Onny, in the vicinity of the junction of the river with Batch Gutter ; at
various exposures in and around the village of Acton Scott ; and at Gretton Quarry
(now filled in), near Cardington. Localities in Onnian strata include the north bank
of the River Onny about 63 yards east of its junction with Batch Gutter (Onnia ?
cobboldi Zone) ; and the river bank 100 yards east of the same junction (Onnia
gracilis Zone).
Family LICHIDAE Hawle & Corda, 1847
Subfamily LICHINAE Hawle & Corda, 1847
Genus METOPOLICHAS Gurich, 1901
TYPE SPECIES. Metopias huebneri Eichwald, 1842 by subsequent designation of
Reed (1902 : 62).
Metopolichas? sp.
(PL 43, figs. 3, 4, 6, 7)
1938. Lichas aff. verrucosa (Eichwald), Stubblefield in Pocock et al., p. 255.
1958. Metopolichas ? aff. verrucosa (Eichwald) Dean, p. 218.
Two fragmentary cranidia from the Costonian Stage may tentatively be referred
to Metopoliches. They are too incomplete to provide a detailed description of the
species, but bear a general resemblance to Metopolichas patriarchus (Wyatt-Edgell),
recently redescribed by Whittard (1961 : 194, pi. 25, figs. 14-17) from west Shrop-
shire. As far as the state of preservation permits, the fragments differ from M.
patriarchus in having a greater convexity, smaller basal lateral lobes, and straighter
longitudinal furrows. Such differences may, however, be nothing more than the
consequence of a different mode of preservation, and a full description of the south
Shropshire form must await additional material. M. patriarchus, although originally
described from the Llanvirn Series of Wales, has been found by Whittard in the
ORDOVICIAN TRILOBITE FAUNAS OF SOUTH SHROPSHIRE, III 235
Llandeilo Series of the Shelve Inlier, an horizon not appreciably earlier than the
Costonian Stage.
HORIZON AND LOCALITIES. Zone of Harknessella subquadrata, Stevenshill Quarry,
by the south-eastern side of the brook, 1,500 yards south-west of Harnage Farm,
Harnage. Zone of Costonia ultima, the old quarry, now filled in, 650 yards west of
Woolston House, Woolston. Both horizons constitute the highest subdivision of
the Costonian Stage in, respectively, the northern and southern parts of the Caradoc
district.
Subfamily HOMOLICHINAE Phleger, 1936
Genus PLATYLICHAS Gurich, 1901
TYPE SPECIES. Lichas margaritifer Nieszkowski, 1857 by original designation of
Gurich (1901 : 522).
Platylichas laxatus (M'Coy)
(PI. 43, figs, i, 2, 5, 8-12)
1846. Lichas laxatus M'Coy, p. 51, pi. 4, fig. 9.
?i846. Calymene (?) forcipata M'Coy, p. 48, pi. 4, fig. 14 (Pygidium only).
1848. Lichas laxatus M'Coy : Salter in Phillips & Salter, p. 340, pi. 8, fig. 6.
1851. Trochurus nodulosus M'Coy in Sedgwick & M'Coy, p. 151, pi. i, F, fig. 16.
1852. Lichas nodulosus (M'Coy) Salter, p. iv.
1854. Lichas sexspinus Angelin, p. 74, pi. 38, figs. j-8a.
1854. Lichas aculeatus Angelin, p. 75, pi. 38, figs, n, na.
1854. Lichas laxatus M'Coy : Murchison, p. 201, fig. 29, 5.
1859. Lichas laxatus M'Coy : Murchison, p. 223, fig. 44, 5.
1866. Lichas laxatus M'Coy : Salter, p. 324, pi. 19, figs. 1—3.
1866. Lichas segmentatus Linnarsson, p. 18, pi. 2, fig. 4.
1867. Lichas laxatus M'Coy : Murchison, p. 204, fig. 46, 5.
1872. Lichas laxatus M'Coy : Murchison, p. 204, fig. 46, 5.
1884. Lichas laxatus M'Coy : La louche, pi. 3, fig. 63.
1885. Lichas laxata M'Coy : Schmidt, p. 125, pi. 6, fig. 24.
1906. Lichas laxatus M'Coy : Olin, p. 53, pi. i, figs. 27, 28.
1908. Lichas laxatus M'Coy : Wiman, p. 133, pi. 8, fig. 23.
1937. Platylichas laxatus (M'Coy)
1938. Platylichas laxatus (M'Coy) pars : Stubblefield, p. 34.
JQ39- Platylichas laxatus (M'Coy)
1945. Platylichas laxatus (M'Coy)
1945. Platylichas laxatus (M'Coy)
1958. Platylichas laxatus (M'Coy)
1958. Platylichas laxatus (M'Coy)
Phleger, p. 1090.
Warburg, p. 118, pi. 12, figs. 1-4, 6, 7, 9-12.
St0rmer, p. 417, pi. 4, fig. 15.
Bancroft, p. 183.
Dean, p. 224.
Tripp, p. 579, pi. 85, figs. 3-5.
This oft-quoted Ordovician trilobite species was founded on a single, incomplete,
slightly compressed cranidium, preserved in a dark-grey, shaly mudstone. The
specimen is in the M'Coy Collection at the National Museum of Ireland, and is
refigured here for the first time since M'Coy's original publication (PL 43, fig. 10).
The outline of the central glabellar lobe is clavate, narrowing posteriorly to a
minimum breadth opposite the posterior portions of the bicomposite lateral lobes,
which themselves converge backwards at roughly 70 degrees. The central glabellar
lobe expands both frontally, constituting the maximum length of the glabella, and
236 ORDOVICIAN TRILOBITE FAUNAS OF SOUTH SHROPSHIRE, III
transversely, so that it is level with the distal margins of the bicomposite lateral
lobes. The short (sag.), flattened, anterior border forms a rim which is separated
from the central lobe by a shallow, narrow (sag.) furrow containing a pair of hypo-
stomal pits opposite the intersection of the axial and longitudinal furrows. The
right palpebral lobe is almost completely preserved ; its length is half that of the
bicomposite lateral lobe, and carries a conspicuous palpebral furrow. The longi-
tudinal furrows are moderately deep, curved, converging backwards for most of
their length, and then diverge to circumscribe the bicomposite lateral lobes. At
this point the holotype is broken, but Tripp (1958, pi. 85, fig. 4) has figured a more
complete topotype cranidium showing most of the essential features.
Platylichas laxatus was described in considerable detail by Warburg (1939 : 118)
on the basis of Norwegian and Swedish material, which she considered to be specifi-
cally identical with that from Ireland. There can be no doubt that, at least as far
as the cranidium of P. laxatus is concerned, there are no significant differences
between Irish, Welsh, English and Scandinavian specimens. The pygidium and
hypostoma of the south Shropshire and Scandinavian forms are, for all practical
purposes, identical but the hypostoma, thorax and pygidium of P. laxatus have not
yet been described from topotype material. Until this has been done, it seems
preferable to follow Warburg's interpretation of the species, though it may eventually
prove necessary to recognize several different species or subspecies.
Under the name Calymene forcipata, M'Coy (1846, pi. 40, fig. 14) figured from the
" Silurian " of " Slieveroe, Rathdrum ", Ireland, a cranidium and pygidium, of
which the latter has been generally supposed to represent Platylichas laxatus. In
order to stabilize the nomenclature of Calymene forcipata, the cranidium figured by
M'Coy, now housed in the National Museum of Ireland, is here chosen as lectotype,
and may be referred to the genus Flexicalymene. The pygidium figured by M'Coy
is refigured in the present paper (PI. 43, fig. 9). The specimen is unusually long,
but this is due in part to mechanical deformation. Excluding the pleural spines, of
which only the third pair is preserved intact, the outline is roughly semi-elliptical.
The axis occupies about one- third of the total breadth, and half the length of the
pygidium ; it is mainly parallel-sided, but posteriorly becomes tapered and
increasingly convex, whilst the terminal piece is poorly defined. Frontally the axial
furrows are well defined and almost parallel, but become convergent near the
terminal piece of the axis ; beyond the latter they diverge slightly and terminate
at the inner margin of the doublure, about halfway to the posterior margin. The
axis has two well-defined axial rings with traces of an additional two. The pleural
lobes carry three pairs of pleurae, each of which has a pair of well-defined pleural
furrows and terminates laterally in a pair of backwardly-directed pleural spines,
though the last-named are only partly preserved. It seems advisable to refer this
specimen merely to Platylichas sp. until further material is available from the
original locality.
HOLOTYPE. M'Coy Collection, National Museum of Ireland (PL 43, fig. 10).
HORIZON AND LOCALITIES. Platichas laxatus has been found at several localities
in south Shropshire, almost all of them comprising strata of the Actonian Stage.
They include the bed and banks of the River Onny immediately east and west of
ORDOVICIAN TRILOBITE FAUNAS OF SOUTH SHROPSHIRE, III 237
its junction with Batch Gutter ; the section in the upper part of Batch Gutter
itself; several points in and around the village of Acton Scott ; exposures near
Plaish, north-east of Church Stretton ; and Gretton Quarry (now filled in), north-
east of Cardington. The species has not been found higher than the Onnia ? cobbolcti
Zone of the Onnian Stage in Shropshire, at which horizon it occurs at the exposure
in the north bank of the Onny, 63 yards east-south-east of its junction with
Batch Gutter. Outside Shropshire, P. cf. laxatus has been recorded from the
Pusgillian Stage and highest Onnian of the Cross Fell Inlier.
Family ODONTOPLEURIDAE Burmeister, 1843
Subfamily ODONTOPLEURINAE Burmeister, 1843
Genus PRIMASPIS R. & E. Richter, 1917
TYPE SPECIES. Acidaspis primordialis Barrande, 1852 by original designation of
R. & E. Richter (1917 : 466).
Primaspis harnagensis (Bancroft)
(PI. 44, figs, i, 4, 6, 8)
1949. Acidaspis harnagensis Bancroft, p. 301, pi. 10, figs. 21, 22.
1958. Primaspis harnagensis (Bancroft) Dean, p. 201.
1962. Primaspis harnagensis (Bancroft) : Dean, p. 122.
DESCRIPTION. The exoskeleton is known only from a few disarticulated cranidia,
together with a single, detached librigena and pygidium. The glabella is subparabolic
in plan, strongly convex, its upper surface declined frontally, with median length
slightly greater than the maximum breadth, measured across the basal glabellar
lobes. The frontal glabellar lobe is small, less than one-fifth of the length of the
glabella, and bluntly pointed. There are three pairs of glabellar lobes, those of the
third pair being the largest, almost oval in plan, slightly divergent forwards, and just
over one-third the length of the glabella. The second glabellar lobes are subcircular
in plan, about half the size of the third lobes. Both pairs of lobes are defined by
deep second and third glabellar furrows which run backwards, and apparently
continue, though shallower, so as to circumscribe the second and third lobes. This
last feature is known only from internal moulds preserved in mudstones or shales,
and may have been exaggerated by crushing. First glabellar lobes are represented
by small, tubercle-like projections, delimited frontally by shallow, straight, first
glabellar furrows directed only slightly backwards. The axial furrows are curved,
strongly convex abaxially, shallowest just behind the line of the second glabellar
furrows and then deepening both anteriorly and posteriorly. The anterior border,
though incompletely known, is apparently of typical odontopleurid type, narrowest
(sag.} medially but broadening abaxially, and delimited posteriorly by a poorly-
defined furrow. The occipital ring is strongly convex transversely, with straight
upper surface ; it is longest (sag.) medially, almost one-quarter the length of the
glabella, but shortens markedly abaxially, at the same time turning sharply forwards
anterolaterally to form a pair of distinct occipital lobes. The glabella and occipital
238 ORDOVICIAN TRILOBITE FAUNAS OF SOUTH SHROPSHIRE, III
ring are separated by a straight, broad (sag.), occipital furrow which deepens
distally. The fixigenae are strongly convex, standing highest posteriorly, where
they are slightly narrower than the basal glabellar lobes, and decline steeply forwards
to end opposite the first glabellar lobes. The palpebral lobes, though poorly known,
are apparently strongly convex abaxially, with well-defined palpebral furrows and
palpebral rims which are continued forwards as eye ridges to join the frontal glabellar
lobe. The ocular surface of the eyes is not known. The anterior branches of the
facial suture are convergent forwards, and apparently cut the cephalic margin
opposite the anterior ends of the axial furrows, though this part of the exoskeleton
has not been found well preserved. As far as can be judged from the available
material, the posterior branches run from the palpebral lobes, a little in front of the
pleuroccipital furrow, backwards to cut the posterior border at the base of the
librigenal spines. The single available librigena (PL 44, fig. 8) is of typical odonto-
pleurid form, steeply declined distally from the eye towards the raised marginal
rim, the latter defined proximally by a deep marginal furrow which ends at the
pleuroccipital furrow. The marginal rim is continued posterolaterally to form a
librigenal spine, the length of which is at least equal to that of the remainder of the
librigena. There are at least thirteen, slender, marginal spines, transversely straight
or directed slightly backwards, and increasing in length (tr.) towards the genal angle.
The hypostoma and thorax are not known.
The pygidium, excluding marginal spines, is transversely semielliptical in plan,
between three and four times as broad (tr.) as long. The axis, occupying frontally one-
quarter of the maximum breadth, narrows backwards slightly at first, but ends in a
small, bluntly pointed terminal piece which reaches the margin. There are three axial
rings, the first well defined, the second and third less so. The axial furrows are
moderately deep except frontally, where the first axial ring is produced postero-
laterally as a pair of strong raised ridges which cross the flat pleural regions and
continue beyond the posterior margin as the fifth and largest of seven pairs of spines
distributed evenly along the margin. The first four pairs of spines increase in size
from small to only slightly shorter than the conspicuous, stout, fifth pair. The sixth
and seventh pairs are slender and subparallel backwards.
The surface of the cephalon, excluding most of the furrows, is covered with closely-
set granules of small, almost uniform, size. Similar granules ornament the entire
dorsal surface of the librigena, including the librigenal spine, but the marginal spines
are smooth. The axis and pleural lobes of the pygidium are covered with slightly
coarser granules, but the furrows and pleural spines are apparently smooth.
LECTOTYPE, here chosen. BM. In. 42086.
PARATYPE. BM. In. 42087 (PI. 44, fig. 6).
HORIZON AND LOCALITIES. The type-locality is the Smeathen Wood Beds,
Harnagian Stage, Reuscholithus reuschi Zone, at the cart-way section near the
southern end of Smeathen Wood, Horderley. A single cranidium has also been
collected from the same horizon in the north bank of Coundmoor Brook, 1,300 yards
south-west of Harnage Farm, Harnage.
DISCUSSION. For convenience the discussion of Primaspis hamagensis is grouped
with that of P, caractaci,
ORDOVICIAN TRILOBITE FAUNAS OF SOUTH SHROPSHIRE, III 239
Primaspis caractaci (Salter)
(PL 44, figs. 3, 7, 9, n, 13, 14)
1853. Acidaspis caractaci Salter, p. 7.
1857. Acidaspis caractaci Salter : Salter, p. 211, pi. 6, figs. 15-17.
1949. Acidaspis caractaci Salter : Bancroft, p. 303.
1958. Primaspis caractaci (Salter) Dean, pp. 211, 223, 224.
1962. Primaspis caractaci (Salter) : Dean, p. 122.
DESCRIPTION. The dorsal exoskeleton, excluding pleural and pygidial spines, is
roughly oval in plan, longer than broad approximately in the ratio 3 : 2. The
cephalon is transversely semielliptical in plan, more than twice as broad as long.
The glabella is steeply declined forwards, almost as broad as long, attaining its
maximum breadth just in front of the occipital furrow, whence it narrows markedly
to the subparabolic frontal glabellar lobe. There are three pairs of glabellar lobes,
those of the basal pair being suboval in plan, moderately divergent forwards, and
of large size, about one-third of the glabellar length. The second glabellar lobes
are just over half the length of the basal lobes and slightly oval in plan, with long
axes parallel to those of the succeeding pair. First glabellar lobes, originally stated
by Salter (1857 : 211) to be obsolete, are poorly represented by a pair of small
tubercles, the sudden diminution in size between the second and first pair of lobes
resulting in a break in the glabellar outline. The second and third glabellar furrows
are parallel to one another, deep, bifurcating, and circumscribe the second and third
glabellar lobes, which are thus separated from the almost paraUel-sided central
glabellar lobe. The axial furrows are narrow and curved, convex abaxially, deepening
both frontally and posteriorly but becoming shallower mediaUy, opposite the third
glabellar furrows. The anterior border is narrow (sag.), moderately inclined forwards,
broadening laterally, and separated from the glabella by a furrow which increases
in depth abaxially. The occipital ring is relatively long (sag.), becoming shorter
laterally where it forms two distinct occipital lobes and extends forwards to join
with the fixigenae ; it is delimited frontally by a broad (sag.), transversely straight,
occipital furrow which deepens laterally. Each librigena is quadrant-shaped, and
its upper surface declines steeply from the eye to a moderately-deep marginal furrow
which ends posterolaterally at its intersection with the pleuroccipital furrow. The
lateral margin is thickened, ridge-like, produced posterolaterally at the genal angle
to form a stout, librigenal spine, the length of which is at least equal to that of the
cephalon. The lateral margin carries thirteen, or occasionally fourteen, short (tr.)
border spines, the hindmost of them almost transversely straight, becoming pro-
gressively less divergent forwards.
The hypostoma is not known.
The thorax comprises ten segments. The axis is moderately convex transversely
and the pleural lobes are flat as far as the bases of the pleural spines. Each pleura
is divided into two bands by a pleural furrow which runs gently backwards distally
from the anterior margin of the pleura at its junction with the axial furrow. The
anterior pleural band is narrow (exsag.), moderately convex, the tip produced
laterally into a short (tr.) pleural spine. The posterior pleural band is broader
(exsag.), more convex, ridge-like, produced posterolaterally to give a posterior
24o ORDOVICIAN TRILOBITE FAUNAS OF SOUTH SHROPSHIRE, III
pleural spine of length almost equal to that (tr.) of the pleura itself. The posterior
pleural spines become progressively more strongly directed backwards from front
to rear of the thorax.
The pygidium, excluding marginal spines, is transversely semielliptical in plan,
with straight anterior margin. The axis occupies about one-quarter of the maximum
breadth of the pygidium, and its anterior portion is strongly convex dorsally ; the
remainder of the axis declines posteriorly and fails to reach the margin, apparently
ending in-line with the inner limit of the doublure. There is evidence of at least two
axial rings. The first of these is produced, in the form of a pair of large raised ridges,
posterolaterally to the margin, beyond which it continues as a pair of stout spines,
slightly divergent backwards. In front of these large spines are situated two pairs
of small, slender, parallel spines, whilst the posterior margin of the pygidium carries
a further three pairs of small spines of similar size and form, parallel to one another
and directed straight backwards.
One of the syntypes of P. caractaci (PI. 44, fig. 13) shows the exoskeleton in what
may have been the position of rest of the animal on the sea-floor. The occipital
ring and pleuroccipital segment are in the same vertical plane, and in this posture
the glabella is steeply declined forwards. At the same time the cephalic margins
rest on the border spines of the librigenae, the progressive increase in length of the
spines posterolaterally assisting in maintaining the position of the cephalon. The
thorax is flexed downwards slightly at its midpoint, rising again towards the pygi-
dium which is, in turn, turned down. A comparable condition is known from other
groups of trilobites, for example, the Trinucleidae and Calymenidae, and may be
posthumous.
The dorsal surface of the glabella, proximal parts of the librigenae, thorax and
pygidium, excluding furrows, is mostly covered with coarse granules, the intervening
space between the latter being ornamented with granules of smaller size. The
glabellar furrows are smooth, but there is a tendency for the smaller granules to
persist across the occipital furrow (PI. 44, fig. 7). Both the lateral border furrow
and the lateral border of the cephalon are finely granulate, the granulation extending
over part of the border spines, as far as can be judged.
HORIZONS AND LOCALITIES. The earliest specimen of Primaspis caractaci known
from south Shropshire is a single librigena from the Onniella reuschi Zone of the
Marshbrookian Stage, just south of Marsh Wood Quarry, Marshbrook. The species
becomes much more abundant, however, in the succeeding Actonian Stage, particu-
larly in the more arenaceous facies, though it is rare in the grey mudstone succession
of the Onny Valley. Localities include the stream section just south of the waterfall
at the western end of Chuney Pool, Acton Scott ; the old quarry 210 yards east
of Acton Scott Church ; the stream section by the east side of the road-bridge about
350 yards west-south-west of Hatton ; and the old quarry, now filled in, at Quarry
Field, Cardington. The last-named locality yielded the type-specimens of P. caractaci,
together with numerous other specimens of the species.
SYNTYPES. GSM. 5214 (PI. 44, fig. 9) ; GSM. 35473 (PI. 44, fig. 13).
DISCUSSION. The cranidium is of generally similar form in both Primaspis
harnagensis and P. caractaci, but the former species may be distinguished by its
ORDOVICIAN TRILOBITE FAUNAS OF SOUTH SHROPSHIRE, III 241
shorter occipital ring, narrower glabella and, perhaps, longer basal glabellar lobes.
The pygidia of these forms may be separated more easily, that of P. harnagensis
having two posterior and four anterior pairs of small marginal spines in addition to
the single large pair, whilst the pygidium of P. caractaci has three posterior and
only two anterior pairs of small spines.
The only other species of Primaspis described from the Caradoc Series of the
Anglo-Welsh area, P. semievoluta (Reed, 1910 : 214, pi. 17, figs. 1-3 ; Dean, 1962)
from the Longvillian Stage of the Cross Fell Inlier, has a pygidium with two posterior
and two anterior pairs of small marginal spines separated by a single pair of large
spines. The pygidium is also slightly longer proportionately than that of the two
Shropshire species.
The cranidium of Primaspis ascitus (Whittington, 1956 : 199, pis. I, 2), from the
Middle Ordovician of Virginia, is relatively broader than that of P. harnagensis and
bears a remarkably close resemblance to that of P. caractaci though the anterior
border of the last-named is slightly longer (sag.). The pygidium of P. ascitus is
more easily distinguished and carries, in addition to the largest pair of margin
spines, a further five pairs of small spines, three pairs situated anteriorly and the
remaining two pairs posteriorly. An unusual feature common to the pygidia of
P. ascitus and P. caractaci is the manner in which the basal part of each of the
large pair of spines is expanded so as to merge with the proximal half of the first
succeeding small, marginal spine (PL 44, fig. 9 ; see also Whittington, 1956, pi. I,
figs. 9, 10).
In a recent paper Whittard (1961 : 203, pi. 27, fig. 12 ; pi. 27, fig. 13) has figured
what he calls Primaspis cf. harnagensis (Bancroft) and P. cf. caractaci (Salter) from
the Spy Wood Grit of the Shelve district, an horizon approximately contemporaneous
with the early Harnagian Stage of south Shropshire. The specimens are too frag-
mentary for certain indentification but appear to be distinct from either of the
south Shropshire forms and may represent a new species.
Family OLENIDAE Burmeister, 1843
Subfamily TRIARTHRINAE Ulrich, 1930
Genus TRIARTHRUS Green, 1832
TYPE SPECIES. Triarthrus becki Green, by monotypy (Green, 1832 : 86-87).
Triarthrus cf. Hnnarssoni Thorslund
(PL 44, figs. 2, 5, 10, 12)
1940. Triarthrus Hnnarssoni Thorslund, p. 128, pi. 12, figs. 4-12.
1945. Triarthrus sp., Bancroft, p. 183.
1949. Triarthrus sp. Bancroft, map on p. 302.
1958. Triarthrus sp., Dean, pp. 213, 225.
1960. Triarthrus cf. Hnnarssoni Thorslund : Dean, p. 86.
Three cranidia in varying degrees of completeness, and a single, small, incomplete
pygidium, have been examined, all from one locality in south Shropshire. They
242 ORDOVICIAN TRILOBITE FAUNAS OF SOUTH SHROPSHIRE, III
match the material described by Thorslund (1940 : 128) in both the shape and
proportions of the cranidium and in the position of the palpebral lobes. Librigenae
of the Shropshire form have not yet been discovered.
The pygidium (PL 44, fig. 2) has a broad axis occupying almost half the total
breadth, though this proportion may have been exaggerated by crushing, and taper-
ing backwards only slightly, its junction with the posterior margin being indistinct.
There are two well-defined axial rings, with a third ring less well defined. The
pleural lobes carry at least two, and possibly three, pleural furrows which extend
from the axial furrows to the margin, the ribs so defined curving backwards strongly
in the same direction. The pygidium of T. linnarssoni has not yet been described,
and comparison is therefore impossible.
HORIZON AND LOCALITY. Onnia Beds, Onnian Stage, Onnia superba Zone, at the
" Cliff Section " in the north bank of the River Onny, 720 yards west-south-west of
Wistanstow Church.
DISCUSSION. Thorslund's type-specimens came from Vastergotland and Jemtland,
Central Sweden, where the species occurs in both the Upper Chasmops beds and
shales of the corresponding Dicranograptus clingani Zone. As has been pointed out
elsewhere (Dean, 1960 : 85-86), the trilobite assemblage of these Swedish strata is
closely comparable with that of the highest Ordovician rocks of the south Shropshire
succession.
Family OTARIONIDAE R. & E. Richter, 1926
Genus OTARION Zenker, 1833
TYPE SPECIES. Otarion diffractum Zenker, 1833 by subsequent designation of
R. & E. Richter (1926 : 95).
Otarion sp.
(PL 45, ng. i)
1958. Otarion sp., Dean, p. 223.
Trilobites which can be assigned broadly to this genus are uncommon in south
Shropshire. A few specimens have been found in the middle and upper Marshbrookian
and in the Actonian, one of the best preserved, an incomplete cranidium about 2-5 mm.
long, being figured here. The glabella is subparabolic in outline, occupies just over
two-thirds of the median length, and is strongly inflated, particularly frontally.
There is one pair of basal glabellar lobes, in length about two-fifths that of the
glabella, from which they are separated by a pair of deep, curved, basal glabellar
furrows. The preglabellar field is convex, steeply declined frontally, separated by a
broad (sag.), well-defined, anterior border furrow from the thick, slightly flattened,
anterior border. Both preglabellar field and anterior border are of equal breadth
(sag.). The fixigenae are only partly preserved but apparently stand highest opposite
the anterior part of the basal glabellar lobes, and are declined frontally, where they
are continuous with the preglabellar field. The occipital ring, delimited anteriorly
by a deep, transversely straight, occipital furrow, is strongly convex transversely,
becoming thicker medially where it is directed upwards and slightly backwards.
ORDOVICIAN TRILOBITE FAUNAS OF SOUTH SHROPSHIRE, III 243
The surface of the glabella, fixigenae and preglabellar field, excluding the furrows,
which are smooth, is covered with conspicuous, coarse tubercles, but that of the
anterior border is finely granulate.
HORIZONS AND LOCALITIES. Specimens belonging, apparently, to the same species
have been found in the Marshbrookian Stage both at and near Marsh Wood Quarry,
Marshbrook ; in the stream-section 550 yards south-south-west of Common Farm,
Wallsbank ; and by the track 2,900 feet north-west of the Methodist Chapel,
Cardington. The highest record is from the middle Actonian at the old quarry
210 yards east of St. Margaret's Church, Acton Scott.
Family PROETIDAE Burmeister, 1843
Subfamily PROETIDELLINAE Hupe, 1953
Genus PROETIDELLA Bancroft, 1949
? Ogmocnemis Kielan, 1959.
TYPE SPECIES. Proetidella fearnsidesi by original designation of Bancroft (1949 :
304).
Proetidella fearnsidesi Bancroft
(PL 45, figs. 3-8, 12, 14)
1949. Proetidella fearnsidesi Bancroft, p. 304, pi. 10, fig. 23.
1953. Decoroproetus fearnsidesi (Bancroft) Pfibyl, p. 60.
1958. Decoroproetus fearnsidesi (Bancroft) : Dean, pp. 201, 219.
?i96i. Ogmocnemis calvus Whittard, p. 186, pi. 24, fig. 15.
1962. Proetidella fearnsidesi Bancroft : Dean, p. 126.
DESCRIPTION. The dorsal exoskeleton of the holotype, the most complete specimen
known, is oval in plan, longer than broad in the ratio 9:7. The roughly semi-
circular cephalon occupies one-third the median length of the holotype, with the
glabella equal to one-third the cephalic breadth. The holotype is a flattened specimen,
preserved in shaly mudstone, and its appearance and proportions are consequently
distorted, but in the case of a specimen preserved in sandy limestones of late
Costonian age (PI. 45, figs. 6, 7) the cephalon is convex both longitudinally and
transversely, with the glabella occupying half the cephalic breadth. The glabella
of the holotype, including occipital ring, is as long as broad ; however, the specimen
is not only compressed but damaged, and its apparently irregular outline, narrowing
forwards to a blunt point, is misleading, though it has been illustrated by Struve
(in Moore, 1959 ; fig. 301, i). The uncrushed glabellar outline is subparabolic, well
rounded frontally, and occasionally the anterior half is slightly constricted. Flat-
tened topotype cranidia tend to appear broader, though with the same general
outline, and there is some variation in the ratio of length to breadth (PI. 45, figs. 12,
14). Most of the known cranidia have no glabellar furrows, though one or two bear
traces of a basal pair ; the conspicuous basal furrows shown in Struve's illustration
(in Moore, 1959, fig. 301, i) are the result of crushing. The glabella is circumscribed
laterally and frontally by a narrow, moderately-deep furrow representing the con-
joined axial and preglabellar furrows. The glabella is preceded by a preglabellar
244 ORDOVICIAN TRILOBITE FAUNAS OF SOUTH SHROPSHIRE, III
field and anterior border of somewhat variable form. The preglabellar field is flat,
varying in length from one-sixth to one-eighth that of the glabella. The line of
demarcation between preglabellar field and anterior border is generally poorly
defined in topotype material, with no distinct anterior border furrow (PI. 45, figs. 3,
12, 14), but specimens in a more arenaceous matrix (see, for example, PL 45, fig. 5)
tend to have a better defined, more steeply upturned, anterior border. The occipital
furrow is well defined and moderately deep, flexing forwards a little both medially
and laterally. The occipital ring follows a similar course, ending laterally in poorly
defined occipital lobes ; its median length is about one-fifth that of the glabella, and
in certain rare instances (PL 45, fig. 14) there is a small median tubercle. The eyes,
the visual surface of which is not known, are elongated, semielliptical in plan, and
slightly convergent forwards, equal in length to just over half that of the glabella.
They are each delimited laterally by a moderately defined eye platform, and extend
forwards from points sited just outside the axial furrows and opposite the occipital
furrow. The combined palpebral lobes and fixigenae are thus small and narrow,
becoming almost obsolete both anteriorly, immediately in front of the eyes, and
posteriorly, where they reach a minimum opposite the pleuroccipital furrow and
then expand suddenly to form part of the pleuroccipital segment (PL 45, fig. 4).
The anterior branches of the facial suture extend only a very short distance forwards
from the eyes, parallel to the axial furrows, before becoming suddenly and markedly
divergent almost as far as the anterolateral margin where they turn adaxially through
a right-angle and cut the margin at points longitudinally in line with the hindmost
parts of the axial furrows. The posterior branches run backwards from the eyes,
just outside and parallel to the axial furrows, as far as the pleuroccipital furrows ;
there they change direction suddenly, running straight outwards and back to cut
the posterior margin of the cephalon at points about mid-way between the axial
furrows and the lateral margins. The librigenae thus formed are of relatively large
size, produced posterolaterally to form long, broad-based librigenal spines, the
points of which end in-line with the mid-point of the thorax, between the fifth and
sixth thoracic segments.
The hypostoma is unknown.
The thorax of the flattened holotype comprises ten segments, and is divided into
three longitudinal lobes of approximately equal breadth by narrow, deep, axial
furrows which are parallel from the first to fifth axial rings, beyond which they
converge backwards until the breadth (tr.) of the tenth axial ring is little more than
half that of the first. The axial rings are of the same general shape as the occipital
ring, though perhaps a little shorter (sag.) . The pleurae are transversely straight for
the most part, but beyond the fulcra become directed backwards and end in short
pleural spines. Each pleura is divided into two bands of unequal breadth (exsag.),
of which the anterior band is the narrower, by a sharply defined pleural furrow ;
each pleural furrow runs slightly backwards distally from the axial furrow, but
turns backwards sharply at the fulcrum towards the pleural spine, at the same time
becoming markedly shallower.
The pygidium is broadly subparabolic in plan with margin entire, the breadth
about two-and-a-half times the median length in the case of the holotype. The
ORDOVICIAN TRILOBITE FAUNAS OF SOUTH SHROPSHIRE, III 245
tapering axis occupies just over three-quarters of the median length, carries six
well-defined axial rings, and ends in a terminal piece of moderate length, with a
small postaxial ridge extending to the margin. The doublure is of even breadth,
extending inwards from the posterior margin as far as the tip of the axis. Each
side-lobe carries four deeply impressed pleural furrows which curve backwards
evenly to intersect the lateral margins ; there are, in addition, one or two further
pairs of only poorly denned pleural furrows. The pleural ribs so formed carry faintly
impressed rib furrows which are not parallel to the pleural furrows, but run back-
wards only slightly for about three-quarters of their length and then turn sharply
back to the margin. Each rib is thus divided into two bands, of which the anterior
is the narrower (exsag.).
Excluding all furrows the entire dorsal surface of the exoskeleton is ornamented
with a series of thin, raised, anastomosing ridges so as to form a Bertillon pattern
which is particularly conspicuous on the glabella.
HORIZONS AND LOCALITIES. The holotype is from shaly mudstones of the basal
Harnagian, Reuscholithus reuschi Zone, in the old cartway near the south end of
Smeathen Wood, Horderley, and a few other specimens have been recovered from
this locality. Proetidella fearnsidesi occurs also, and in small numbers, at the same
horizon as exposed 550 yards north-west of Woolston House, Woolston. In the
northern part of the Ordovician outcrop in south Shropshire the same species may
be found occasionally in the topmost subdivision, the Harknessella subquadrata Beds,
of the Costonian Stage.
HOLOTYPE. BM. In. 42083 (PI. 45, fig. 3).
DISCUSSION. It has recently been suggested by Dean (1962) that Ogmocnemis
calvus, described by Whittard (1961 : 186) from the Lower Soudleyan of the Shelve
Inlier, may prove to be synonymous with P. fearnsidesi. 0. calvus was founded on
a single specimen, so there is no information concerning variation within the species.
The preglabellar field and anterior border are apparently fairly well differentiated,
but this may not be of specific significance, particularly in view of the variation in
Harnagian specimens of P. fearnsidesi, and the two species are regarded here as
being probably synonymous. Specimens of Proetidella are figured in the present
paper from the Upper and, probably, Lower Soudleyan of south Shropshire (PL 45,
figs. 9-11). They show small differences from P. fearnsidesi in glabellar outline and
degree of differentiation of the anterior border, but owing to lack of material it has
been impossible to demonstrate whether or not these are more than an expression
of a different environment and lithology. The specimens, which are closely similar
to the holotype of Ogmocnemis calvus, are therefore only compared with P. fearn-
sidesi.
There are in the old collections of the Geological Survey & Museum certain
specimens, numbered 35614-35616 and 35618-35620, which are labelled Proetus
ovatus, a manuscript name attributed to Salter. The specimens are typical Proetidella
fearnsidesi, and the state of preservation suggests that they originated from the
Smeathen Wood Beds of the Horderley district.
The Norwegian species Proetus asellus (Esmark), from metamorphosed shales of
probable Llanvirn age, has been redescribed and figured by St0rmer (1940 : 122,
GEOL. 7, 8 18
246 ORDOVTCIAN TRILOBITE FAUNAS OF SOUTH SHROPSHIRE, III
pi. I, fig. i). The glabella is of more uniform breadth than that of Proetidella fearn-
sidesi, and the anterior border is apparently better differentiated from the long (sag.)
preglabellar field, though the importance of the latter feature may not be great.
However, the general resemblance of the two forms is enough to suggest that
Proetidella-like trilobites existed earlier than the Caradoc Series, though the group
is inadequately known.
Proetidella? sp.
(PI. 45, fig. 13)
1958. Proetus (s.l.) sp., Dean, p. 223.
A single proetid cranidium has been recovered from the Upper Cheney Longville
Flags, west of the village of Hatton. The glabellar outline is relatively broad,
especially posteriorly, becoming broadly convergent frontally towards the well-
rounded frontal glabellar lobe after constricting noticeably at about its mid-point.
There is a suggestion of a pair of basal glabellar furrows, but as the specimen is
somewhat damaged this feature may have been exaggerated by crushing. The
anterior border is bluntly pointed medially, gently inclined forwards, separated
from the broad (sag.), preglabellar field by a shallow, anterior border furrow. The
glabella is circumscribed by a conspicuous preglabellar furrow, continuous laterally
with deep axial furrows which become shallower medially. Part of the right occipital
lobe remains, showing it to be generally similar to that of Proetidella fearnsidesi,
though perhaps a little broader and situated slightly farther from the axial furrow.
There is insufficient material for the erection of a new species, though the specimen
is probably specifically distinct from P. fearnsidesi.
HORIZON AND LOCALITY. Marshbrookian Stage, Dalmanella unguis Zone, about
350 yards north-west of the road bridge situated 350 yards south-west of Hatton,
near Acton Scott.
Family REMOPLEURIDIDAE Hawle & Corda, 1847
Genus REMOPLEURIDES Portlock, 1843
TYPE SPECIES. Remopleurides colbii Portlock by subsequent designation of Miller
(1889 : 565-566).
Remopleurides warburgae sp. nov.
(PL 46, figs, i, 2)
1958. Remopleurides sp. nov., Dean, p. 224.
DIAGNOSIS. Glabella slightly broader than long, broadest posteriorly, narrowing
frontally. Glabellar tongue strongly convex, steeply downturned, equal to about
half breadth of glabella. Palpebral lobes flattened, narrowing frontally. Three pairs
of equispaced glabellar furrows show through test as black lines. Test mainly smooth.
DESCRIPTION. The glabella is moderately convex longitudinally and transversely,
almost as long as broad, its greatest breadth occurring to the rear of centre of the
main body. The anterior portion of the glabella is convex, including the glabellar
tongue, which is strongly arched downwards. The glabellar tongue is about twice
ORDOVICIAN TRILOBITE FAUNAS OF SOUTH SHROPSHIRE, III 247
as broad as long, its breadth half that of the glabella, and the lateral margins con-
verge slightly towards the almost straight anterior margin. Strong palpebral furrows
delimit the palpebral lobes, which narrow frontally and continue forwards as narrow
rims to the anterolateral angles of the glabellar tongue. Three pairs of equidistant,
parallel, glabellar furrows are present, their position indicated by poorly visible
black lines showing through the test ; the first pair is shorter (tr.) than the second
and third pairs, which are of almost equal length. The glabellar furrows curve
gently backwards from near the palpebral furrows so as to leave an unfurrowed
band extending along the axis of the glabella and occupying about one-fifth of the
maximum breadth. The occipital furrow is deeply incised and transversely straight.
The occipital ring is moderately convex both longitudinally and transversely, but in
the holotype the distal parts are not wholly preserved. The surface of the cranidium
is smooth except for the occipital ring, palpebral lobes and anterior portion of the
glabellar tongue, all of which are ornamented with fine, slightly-raised, anastomosing
ridges, sometimes forming a Bertillon pattern.
The hypostoma, thorax and pygidium are not known (but see later).
HORIZON AND LOCALITIES. The type-material was collected from grey mudstones
in the uppermost third of the Actonian Stage in the north bank of the River Onny,
87 feet east of the junction of the river with Batch Gutter. The same species occurs
also in the Actonian Stage at Acton Scott, and, perhaps, near Hatton.
HOLOTYPE. BM. In. 49751 (PI. 46, figs, i, 2).
DISCUSSION. In plan the glabella of Remopleurides warburgae most resembles
R. dalecarlicus Holm MS., described and figured by Warburg (1925 : 88, pi. i,
figs. 7, 8) from the Upper Leptaena Limestone of Sweden and therefore younger
than the Shropshire species ; the Swedish form also has a smooth test but possesses
a narrower, more convex, glabellar tongue.
Remopleurides latus Olin onniensis subsp. nov.
(PL 46, figs. 4, 5)
DIAGNOSIS. Glabella broader than long, with main body transversely ovate in
plan. Glabellar tongue equal in breadth to half that of glabella, strongly declined
frontally, its anterior margin transversely truncated and indented medially.
Palpebral lobes narrow forwards, forming rim along sides of anterior glabellar
tongue. Three pairs glabellar furrows visible through test as dark lines.
DESCRIPTION. The glabella is moderately convex both longitudinally and trans-
versely, its length (sag.} equal to three-quarters the maximum breadth, which occurs
midway between the occipital furrow and the base of the glabellar tongue. The
latter is longitudinally convex, less so transversely, and strongly arched down
frontally, its breadth half that of the glabella. Apart from a small median indentation
the anterior margin of the glabellar tongue is transversely straight, meeting the
anterolateral margins at right-angles. Three pairs of equidistant glabellar furrows
are present, showing through the test as poorly-discernible dark lines, and not
impressed upon the outer surface of the test. The first pair is shorter (tr.} than the
second and third pairs which extend distally almost to the palpebral furrows ; the
248 ORDOVICIAN TRILOBITE FAUNAS OF SOUTH SHROPSHIRE, III
third pair curves backwards a little more strongly than the first and second pairs,
which are almost parallel to one another. All three pairs of furrows curve gently
backwards and extend adaxially from the lateral margins so as to leave an unmarked
band, running down the centre of the glabella and about one-fifth the breadth of the
latter. The palpebral lobes, which narrow frontally and are separated from the
main body of the glabella by deeply-incised palpebral furrows, continue forwards
in the form of a much attenuated rim as far as the anterolateral angles of the
glabellar tongue. The occipital ring is only slightly convex longitudinally, and
becomes shorter (exsag.) laterally. The surface of the test of the glabellar tongue
and occipital ring is ornamented with fine, anastomosing, raised ridges arranged
parallel to the anterior margin, but the remainder of the test is smooth. The
hypostoma, thorax and pygidium are not known (but see later).
HORIZON AND LOCALITIES. The uppermost third of the Actonian Stage, exposed
in the north bank of the River Onny, 87 feet east of the junction of the river with
Batch Gutter. The type-material was found in a band of shelly debris, about
3 inches thick, containing also Remopleurides warburgae, Tretaspis, Chasmops, Sampo
and Onniella. The species is not known with certainty elsewhere, but fragments
possibly belonging to it have been found in the Actonian Stage at Acton Scott.
HOLOTYPE. BM. In. 49750 (PI. 46, figs. 4, 5).
PARATYPE. BM. In. 49762.
DISCUSSION. The new subspecies is generally similar to Remopleurides lotus (Olin,
1906, pi. 2, figs. 50, b) from the Chasmops Series of Scania, particularly in the form
of the posterior half of the glabella, but the convexity of the glabella is greater and
the glabellar tongue is both longer and broader. R. latus kullsbergensis (Warburg,
1925 : 83, pi. i, figs. 1-6) has more distinct glabellar furrows as well as a smaller,
narrower glabellar tongue than the Shropshire subspecies, though the two are
probably of generally similar age, whilst R. latus var. granensis (St0rmer, 1945 : 408,
pi. 4, fig. 4), from a slightly higher horizon, differs in possessing a narrower glabellar
tongue, and a test which is described as having a completely smooth surface. R.
validus (Thorslund, 1940, pi. 7, figs. 1-9), from the Lower Chasmops Limestone,
shows a glabella of generally similar outline to that of the new subspecies but the
glabellar tongue is both narrower and less well differentiated from the main body of
the glabella. R. wimani (Thorslund, 1940 : 135, pi. 7, fig. 10), though of an age
generally comparable to that of R. latus onniensis, exhibits a much narrower glabellar
tongue and the main body of the glabella is both narrower and more convergent
forwards.
Remopleurides sp.
(PI. 46, figs. 3, 7)
At the type-locality of Remopleurides warburgae and R. latus onniensis, isolated
fragments of Remopleurides have been found in abundance in a fossiliferous band of
shelly mudstone about 3 inches thick. There appears to have been some sorting of
the trilobite remains and although librigenae and thoracic segments are not uncom-
mon, no pygidium has yet been found. Two typical specimens are figured here, but
it has not yet been possible to assign either to a definite species.
ORDOVICIAN TRILOBITE FAUNAS OF SOUTH SHROPSHIRE, III 249
The main part of the librigena is subtriangular in outline (PI. 46, fig. 3), slightly
convex, and arched down laterally. A long, stout spine is developed, the whole of
which is not preserved ; as in the case of Remopleurides latus kullsbergensis (Warburg,
1925 : 84, fig. 15), this is not developed from the true genal angle but arises from a
position just in front of it. The posterior portion of the librigena stands at a some-
what lower level than the remainder, from which it is separated by a strong transverse
furrow. This part of the librigena terminates posterolaterally in a short, blunt,
librigenal spine, separated by a well-defined notch from the long anterior spine,
but the proximal end is marked by a rounded, flange-like notch, the function of
which is to provide articulation with the fulcral process of the first thoracic segment.
The surface of the test is ornamented with several well-developed, anastomosing
ridges, subparallel to the distal margin of the librigena as far as the genal angle, but
transverse in direction near the fulcral notch. The librigena strongly resembles that
of R. latus kullsbergensis and may belong to the allied R. latus onniensis. It also
recalls, but is shorter and broader than, one figured by Thorslund (1940, pi. 7,
fig. 16) and referred by him to R. cf. latus Olin.
In addition to numerous isolated thoracic segments, a group of six articulated
segments has been found (PL 46, fig. 7). The axial rings are gently convex longi-
tudinally and transversely, the breadth at least four times the length (sag.) , excluding
the articulating half-ring. Apart from a small, central portion the posterior margin
of each axial ring is serrated, particularly distally, up to six tooth-like spines being
visible on the lateral half of each ring ; similar structures are developed on the
axial rings of Remopleurides colbii Portlock (Whittington, 1950, pi. 70, figs. I, 4).
The side-lobes are narrow, with flattened pleurae terminating in short (exsag.),
broad, backwardly-directed pleural points. The pleural furrows are curved, deep,
directed distally towards, but not attaining, the pleural points. Just outside the
axial furrow the anterior margin of each pleura is thickened markedly to form a
conspicuous, tooth-like, fulcra! process, the posterior margin being indented to give
a corresponding, flange-like, articulating socket. The test is ornamented with fine
raised ridges distributed more or less parallel to the margins of each segment and
tending to converge at the pleural points.
HORIZON AND LOCALITY. Actonian Stage, north bank of the River Onny, 87 feet
east of its junction with Batch Gutter.
Genus REMOPLEURELLA nov.
DIAGNOSIS. Cranidium of typical remopleuridid outline, the main body only
gently convex, both longitudinally and transversely, with three pairs of deeply-
incised glabellar furrows, those of the first pair noticeably shorter (tr.) than those
of the other two pairs. Surface of test with conspicuous pattern of subconcentric,
thin, raised lines. Librigenae narrow with librigenal spines arising from the genal
angles ; posterior margin entire. Hypostoma as long as, or slightly longer than,
broad, with transversely straight anterior and posterior borders. Median body
suboval, gently convex, with pair of oval areas converging backwards medially ;
small anterior lobe, sometimes produced to form short spine, delimited by broad,
250 ORDOVICIAN TRILOBITE FAUNAS OF SOUTH SHROPSHIRE, III
curved median furrow, convex backwards. Thorax incompletely known. Pygidium
unknown.
Robergiella, type-species R. sagittalis, from the Edinburg Limestone of Virginia
(Whittington, 1959 : 431, pi. 6, figs. 16-33), may be related to Remopleurella, which
it resembles in the depressed glabellar form, with three incised pairs of glabellar
furrows. Robergiella differs, however, in having a larger glabellar tongue which
expands forwards, as well as considerably broader librigenae, each of which carries
a pronounced notch at the base of the librigenal spine.
TYPE SPECIES. Remopleurides burmeisteri Bancroft, 1949.
DISTRIBUTION. Known with certainty from the Onnian Stage of the type Caradoc
Series in south Shropshire. Probably occurs also in the higher Caradoc strata of
southern Norway.
Remopleurella burmeisteri (Bancroft)
(PI. 45, fig. 2 ; PI. 46, figs. 6, 8, 9, n, 12)
1945. Remopleurides (? Caphyra) sp., Lament, p. 118.
1949. Remopleurides burmeisteri Bancroft, p. 300, pi. 10, figs. 19, 20.
1958. Remopleurides burneisteri Bancroft : Dean, pp. 213, 225.
Bancroft's original description covered the cranidium, librigenae and thorax (pars),
and little need be added. Most known cranidia, including one of the syntypes, are
slightly flattened with the glabellar tongue apparently arched down only slightly,
but occasional specimens show that the plane of the tongue is, in fact, steeply
declined frontally, almost at right-angles to the remainder of the glabella. Most of
the known specimens are preserved in decalcified mudstones in which the glabellar
test is not seen, but one specimen, BM. In. 54772, from the unweathered underlying
strata shows the test to have been thin, its dorsal surface, apart from the smooth,
deeply-incised glabellar furrows, ornamented by thin, raised, anastomosing ridges
forming a Bertillon pattern.
A hypostoma presumed to belong to this species occurs in association with the
paratype thorax, and a slightly smaller specimen has recently been collected by
Mr. S. F. Morris. The entire hypostoma is as long as, or slightly longer than, wide.
The gently convex median body is generally ovate in outline with bluntly terminated
anterior and posterior margins, and is circumscribed by a moderately-deep border
furrow. The posterior border is apparently continuous with the narrow (tr.) lateral
border, which is, in turn, continuous with the anterior border. The last-named is
transversely straight produced distally to form a pair of anterior wings, separated
by rounded lateral notches from the lateral borders. The median body is divided
into two unequal parts by a curved median furrow, concave forwards, deepest
medially, and shallowing distally. The anterior lobe is small, convex, and specimen
In. 56800 shows it thickened medially to produce a small spine. The posterior lobe,
occupying about three-quarters of the length of the median body, is traversed
longitudinally by a faint groove, a continuation of the median furrow, which
narrows backwards and ends at the posterior border. The posterior lobe is thus
divided so as to form what Whittington (1950 : 541) has described as two oval
ORDOVICIAN TRILOBITE FAUNAS OF SOUTH SHROPSHIRE, III 251
areas, their long axes convergent backwards. Fragments of the test show it to be
thin, its lower surface covered by subconcentric, thin, raised ridges.
LECTOTYPE, here chosen. BM. In. 42106 (PL 46, fig. n).
PARATYPES. BM. In. 42098 (PL 46, fig. 6) ; BM. In. 49561 (PL 46, figs. 8, 12).
HORIZONS AND LOCALITIES. The earliest known occurrence of the species is a
single cranidium, BM. In. 54772, from the Onnian Stage, Onnia gracilis Zone, by
the north bank of the River Onny about 100 yards east of the junction of the river
with Batch Gutter. All the other known specimens, including the types, are from
the Onnia superba Zone in the north bank of the Onny, 720 yards west-south-west
of Wistanstow Church.
DISCUSSION. Remopleurella burmeisteri appears to belong to a group of remo-
pleuridid trilobites which, although they have received little attention, are not
uncommon in at least the Upper Chasmops Limestone, Stage fod, of southern
Norway, and probably occur also in parts of Sweden. Several such are housed in
the Paleontologisk Institut, Oslo, where some have been labelled as ' Remopleurides
cf. radians ", the type-species of Amphitryon Hawle & Corda, but they cannot be
assigned to the latter genus and probably belong to Remopleurella.
Family THYSANOPELTIDAE Hawle & Corda, 1847
Genus EOBRONTEUS Reed, 1928
TYPE SPECIES. Entomostracites laticauda Wahlenberg, 1818 by original designation
of Reed (1928 : 58, 75).
Eobronteus? sp.
(Plate 46, fig. 10)
1958. Eobronteus ? sp., Dean, p. 224.
Three fragmentary specimens housed in the Sedgwick Museum are the only
representatives of this trilobite family found in south Shropshire, and the best
preserved of these is now figured. The specimen comprises an almost complete
thorax, the full number of thoracic segments not being visible, together with a
fragmentary cephalon, which partly obscures the thorax, and an incomplete pygidium.
The genus Eobronteus has been examined in detail by Sinclair (1949) who has figured
or noted all the known species. None of these agrees wholly with the Shropshire
species, which may be new, but a detailed description must await the collection of
better-preserved material.
HORIZON AND LOCALITY. Onnian Stage, Onnia superba Zone, north bank of the
River Onny, 720 yards west-south-west of Wistanstow Church.
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PLATES
Specimens whitened with ammonium chloride before photographing. Numbers with prefixes
BM. In., GSM., and A. are housed respectively in the British Museum (Natural History), London,
the Geological Survey & Museum, London, and the Sedgwick Museum, Cambridge. Other
figured specimens are in the Shrewsbury Museum and the National Museum of Ireland, Dublin.
PLATE 37
Flexicalymene acantha Bancroft . . . . p. 216
Harnagian Stage, zone of Reuscholithus reeuschi, section in old cartway, near southern end of
Smeathen Wood, near Horderley.
FIG. i. Internal mould of cephalon with incomplete thorax. BM. In. 50635. x 2.
FIG. 3. Lectotype, an immature cranidium preserved as an internal mould. BM. In. 42091.
X3-
FIG. 4. Paratype pygidium, an internal mould. BM. In. 49592. x 3.
FIGS. 5, 6. Latex cast from external mould of adult cranidium. BM. In. 50622. x 2.
FIG. 14. Latex cast from external mould of damaged, almost whole, dorsal exoskeleton.
BM. In. 50623. X2'5.
Flexicalymene cf. acantha Bancroft . . . . p. 217
Costonian Stage, zone of Costonia ultima, old quarry about 500 feet south of Coston Farm, i
mile east of Clunbury.
FIG. 7. Internal mould of cranidium showing glabellar outline. BM. In. 51694. X2.
FIG. 10. Cranidium illustrating variation in glabellar outline. BM. In. 52226. x 2.
FIG. 13. Internal mould of pygidium, with axis broadened by crushing. BM. In. 52227.
X2.
Flexicalymene (Reacalymene} horderleyensis sp. nov. . . p. 221
Lower Longvillian Substage, zone of Dalmanella indica and D. lepta, south-eastern corner of
Rookery Wood, south-east of Horderley.
FIGS. 2, 8. Latex cast of holotype, an external mould. BM. In. 50656. X2.
Flexicalymene planimarginata (Reed) . . . p. 217
Lower Longvillian Substage, zone of Dalmanella horderleyensis, Long Lane Quarry, f mile
north-west of Craven Arms.
FIG. 15. Internal mould of pygidium. BM. In. 52104. x 2*3.
Flexicalymene (Reacalymene} pusulosa (Shirley) . . p. 220
Costonian Stage, probably zone of Harknessella subquadrata, quarry near eastern end of Black
Dick's Coppice, Evenwood.
FIGS. 9, ii. Holotype cranidium with test preserved. BM. In. 50780. X2*5.
Horizon as for Fig. 9. Old quarry in north-western part of Round Nursery, south of Harnage
Grange.
FIG. 12. Pygidium showing granulate surface. BM. In. 54868. x 3.
Flexicalymene (Reacalymene) cf. limba (Shirley) . . p. 221
Soudleyan Stage, probably zone of Broeggerolithus broeggeri, road-side quarry 150 yards
north-east of Chatwall Farm, Chatwall.
FIG. 16. Internal mould of incomplete cranidium. BM. In. 54320. x 2-5.
FIGS. 2, 5, 6, 8 and 14 by Mr. E. W. Seavill, remainder by the writer.
Bull. B.M. (N.H.) Geol. 7, 8
PLATE 3?
14
FLEXICALYMENE. FLEXICALYMENE (REACAI YMENF)
PLATE 38
Flexicalymene cobboldi sp. nov. . . . p. 218
Marshbrookian Stage, zone of Dalmanella unguis, Marshwood Quarry, \ mile south of Marsh-
brook Station.
FIG. i. Internal mould of cranidium, associated with that of Otarion sp. BM. In. 50762.
X2.
Upper Longvillian Substage. zone of Kjaerina bipartita (=Alternata Limestone), south-eastern
side of Horderley-Marshbrook road, 420 yards north-east of Crosspipes, about 2 miles north-east
of Horderley Station.
FIG. 3. Holotype, internal mould of cranidium. BM. In. 55448. x 2*3.
Flexicalymene caractaci (Salter) . . . . p. 217
Marshbrookian Stage, zone of Onniella reuschi, well-sinking at " first house along road south
from The Corner", near Wistanstow.
FIG. 2. Internal mould of incomplete cranidium. BM. In. 52820. xi'Q. Horizon and locality
as for Fig. i.
FIGS. 4, 5. Internal moulds of cranidium, thorax and pygidium, associated on same hand-
specimen. Syntypes of Flexicalymene trigonoceps Bancroft (1949, pi. n, fig. 32). BM.
In. 42102. Fig. 4, X2'4; Fig. 5, x 2.
Marshbrookian Stage, zone of Dalmanella wattsi, " Acton Scott Lodge ", near Acton Scott.
FIG. 6. Lectotype, an internal mould. GSM. 47698. x 1*5. Horizon as for Fig. 2. West side
of track in Marsh Wood, just south of the quarry.
FIGS. 8, ii. Internal mould of cranidium illustrating inclined anterior border. BM. In. 50735.
Fig. 8, X2; Fig. n, X2'5.
Horizon as for Fig. 6, south-eastern side of Horderley road, 520 yards west-south-west of
Marshbrook Station.
FIG. 12. Internal mould of incomplete cranidium. BM. In. 55446. X2.
Onnicalymene laticeps (Bancroft) . . . p. 224
Actonian Stage, north end of wood west of Rose Villa, Marshbrook.
FIG. 7. Holotype cranidium, an internal mould. BM. In. 42103. X2. Uppermost Actonian
Stage, north bank of River Onny, 62 feet east of junction with Batch Gutter.
FIGS. 10, 14, 15. Uncrushed cranidium with test preserved. BM. In. 50723. x 2*5.
Onnicalymene salteri (Bancroft) . . . p. 224
Horizon and locality as for Fig. 7.
FIG. 9. Holotype cranidium, an internal mould. BM. In. 42100. X2. Horizon as for Fig. 7,
south bank of River Onny, 80 yards west of junction with Batch Gutter.
FIG. 13. Incomplete cranidium with test preserved. BM. In. 50736. xi'8.
Fig. i by Mr. E. W. Seavill, remainder by the writer.
Bull. B.M. (N.H.) Geol. 7, 8
PLATE 38
FLEXICALYMENE, ONNICALYMENE
PLATE 39
Gravicalymene praecox (Bancroft) . . . p. 225
Harnagian Stage, zone of Reuscholithus reuschi, section in old cart-way near south end of
Smeathen Wood, near Horderley.
FIG. i. Slightly crushed internal mould of large cranidium. BM. In. 50523. xi'8.
FIGS. 3, 9. Internal mould of large, incomplete cranidium showing glabellar outline and form
of anterior border and axial furrows. BM. In. 50522. x 2.
FIG. 12. Internal mould of incomplete cranidium showing palpebral lobe and undistorted
axial furrows. BM. In. 50657. xi'8.
FIGS. 13, 14. Holotype, the internal mould of an immature cranidium, the glabella bearing
the impression of an ostracod, Tallinella scripta (Harper). BM. In. 42090. X3'5-
Gravicalymene cf. praecox (Bancroft) . . . p. 225
Onnian Stage, zone of Onnia gracilis, north bank of River Onny, 100 yards east of junction
with Batch Gutter.
FIG. 4. Cranidium with test preserved. BM. In. 50654. Xi'5-
FIGS. 5, 8. Thorax and pygidium associated with, and believed to belong, to same individual
as Fig. 4. BM In. 50655. Xi'5-
Actonian Stage, site of old quarry, Quarry Field, Gretton, near Cardington.
FIG. 7. Internal mould of incomplete cranidium. BM. In. 50648. xi'5-
Gravicalymene inflata sp. nov. . . . p. 227
Horizon and locality as for Fig. 4.
FIG. 6. Holotype, an incomplete cranidium with the test preserved. BM. In. 50653. x 1*5.
Onnicalymene onniensis (Shirley) . . . p. 223
Horizon and locality as for Fig. 4.
FIG. 2. Thorax and pygidium with test preserved. BM. In. 52092. x 2. Horizon and locality
probably as for Fig. 4.
FIGS. 10, ii. Holotype cranidium. GSM. RR 1940. X2.
Figs. 4-8 by Mr. E. W. Seavill, remainder by the writer.
Bull. B.M. (N.H.) Geol. 7, 8
PLATE 39
•-.u-*i &%&&m
12
GRAVICAT .YMFMF OMMTPAT YUFMF
GEOL. 7, 8
PLATE 40
Parabasilicus powisi (Murchison) . . . p. 229
Costonian Stage, probably zone of Harknessella, subquadrata, old quarry just west of Round
Nursery, south-east of Harnage Grange.
FIG. i. Internal mould of pygidium and fragment of thorax. BM. In. 52197. x i'3.
Soudleyan Stage, zone of Broeggerolithus broeggeri, exposure behind Glenburrell farmhouse,
Horderley.
FIG. 3. Internal mould of pygidium and thorax. BM. In. 52186. xi'i.
Harnagian Stage, zone of Reuscholithus reuschi, south bank of Coundmoor Brook, 1340 yards
south-west of Harnage Farm, Harnage.
FIG. 4. Internal mould of cranidium with fragments of test, showing course of facial suture.
BM. In. 54237. xi'5.
FIG. 5. Internal mould of slightly distorted pygidium showing anterolateral facets. BM.
In. 54238. X2'2.
Parabasilicus powisi (Murchison) ? . . p. 231
Middle Costonian Stage, Harknessella Beds, north side of cartway 50 yards east of Coston
Farmhouse, i mile east of Clunbury.
FIG. 2. Internal mould of hypostoma lacking anterior wings. BM. In. 50621. xi'6.
Parabasilicus ? sp. . . . . p. 232
Lower Longvillian Substage, zone of Dalmanella indica and D. lepta, top of western end of
Rookery Wood, south-east of Horderley.
FIG. 6. Internal mould of pygidium. BM. In. 50741. X 1*5.
Flexicalymene (Reacalymene) horderleyensis sp. nov. . . p. 221
Horizon as for Fig. 6, old quarry just west of western end of Rookery Wood, south-east of
Horderley.
FIGS. 7, 8. Internal mould of incomplete cranidium. BM. In. 52103. X2.
Onnicalymene onniensis (Shirley) . . . p. 223
Onnian Stage, zone of Onnia superba, north bank of River Onny 720 yards west-south-west of
Wistanstow Church.
FIG. 9. Internal mould of cranidium. BM. In. 50711. X2.
Fig. 9 by Mr. E. W. Seavill, remainder by the writer.
Bull. B.M. (N.H.) Geol. 7, 8
PLATE 40
FLEXICAT.YMFNF mFAHAT YMFNF\ OMMIPAT YUFMF PAR ARA9TT in TQ
PLATE 41
Parabasilious powisi (Murchison) . . . p. 229
Soudleyan Stage, zone of Broeggerolithus broeggeri, eastern bank of Habberley Brook, 500
yards north of The Lyd Hole, near Pontesford, Shropshire.
FIG. i. Internal mould of cranidium, left librigena (displaced), and thorax. BM. In. 54305.
Xi'i.
Probably Harnagian Stage, Bron-y-buckley Wood (=Trilobite Dingle), Welshpool, Mont-
gomeryshire.
FIG. 2. Vinyl plastic cast of lectotype, an external mould of pygidium and incomplete thorax
GSM. 6841. xn.
Fig. i by the writer, Fig. 2 by Mr. J. Brown.
Bull. B.M. (N.H.) GeoL 7, 8
PLATE 41
PARABASILICUS
PLATE 42
Parabasilicus ? marstoni (Salter) . ,. . . p. 231
Horizon probably Smeathen Wood Beds, Harnagian Stage pars, of the Horderley district.
Exact locality unknown.
FIGS, i , 6. Syntype, internal mould of an almost complete dorsal exoskeleton with damaged
cephalon. GSM. 12817. X 4- Fidg. Salter, 1866, pi. 23, figs, i, \a.
FIG. 8. Syntype, internal part external mould of right librigena. GSM. 12818. X2'5. Figd.
Salter, 1865, pi. 23, fig. ib.
Parabasilicus ? sp. . . . . p. 232
Probably from the Horderley Sandstone, Lower Longvillian Substage, of the Horderley
district.
FIG. 2. Internal mould of hypostoma. GSM. 12830. x 2. Figd. Salter, 1866, pi. 23, fig. 6.
Lower Longvillian Substage, zone of Dalmanella horderleyensis, quarry by north side of road,
300 yards south-east of Glenburrell Farm, Horderley.
FIG. 5. Internal mould of pygidium. BM. In. 51459. X2.
Parabasilicus powisi (Murchison) . . . p. 229
Harnagian Stage, zone of Reuscholithus reuschi, section in old cart-way near southern end of
Smeathen Wood, near Horderley.
FIG. 9. Latex cast from external mould of frontal portion of cranidium. BM. In. 51462.
X2.
Illaenus cf. fallax Holm ..... p. 233
Onnian Stage, zone of Onnia ? cobboldi, north bank of River Onny, about 70 yards east of
junction with Batch Gutter.
FIG. 3. Internal mould of pygidium. BM. In. 50760. xi'9- Onnian Stage, zone of Onnia
gracilis, north bank of River Onny, 100 yards east of junction with Batch Gutter.
FIG. 4. Internal mould of pygidium. BM. In. 50515. x 1*9.
FIG. 12. Cranidium with most of test preserved. BM. In. 50759. x i'3-
Illaenus sp. . . . . p. 233
Actonian Stage, site of old quarry, Quarry Field, Gretton, near Cardington.
FIGS. 7, 10. Internal mould of cranidium. BM. In. 50576. x i'6.
FIG. IT. Internal mould of immature cranidium. BM. In. 50755. x 2-3.
Figs. 7, 10 by Mr. E. W. Seavill, remainder by the writer.
Bull. B.M. (N.H.) Geol. 7, 8
PLATE 42
PARABASILICUS, ILLAENUS
PLATE 43
Metopolichas ? sp. . . . . p. 234
Costonian Stage, zone of Harknessella subquadrata, old quarry by south-eastern side of brook,
1500 yards south-west of Harnage Farm, Harnage.
FIGS. 3, 6, 7. Internal mould of incomplete, slightly distorted cranidium. BM. In. 46455.
Figs. 3, 6, xi'4; Fig. 7, X2.
Costonian Stage, zone of Costonia ultima, old quarry 650 yards west of Woolston House,
Woolston.
FIG. 4. Internal mould of fragmentary cranidium. GSM. RR 2140. X2.
Plaiylichas laxatus (M'Coy) . . . p. 235
Actonian Stage, site of old quarry, Quarry Field, Gretton, near Cardington.
FIG. i. Latex cast from external mould of large pygidium. BM. In. 49558. x 1*4.
FIG. 8. Internal mould of incomplete cranidium. BM. In. 50749. Xi'5-
FIG. 12. Internal mould of hypostoma. BM. In. 50524. xi'7-
Horizon as for Fig. i, south bank of River Onny, 80 yards west of junction with Batch Gutter.
FIG. 2. Incomplete cranidium showing surface of test. BM. In. 50747. x 2.
Horizon as for Fig. i, old quarry 1400 feet west-south-west of St. Margaret's Church, Acton
Scott.
FIG. 5. Internal and partly external mould of pygidium, showing impression of terrace-lines
on doublure. BM. In 50754. x 2*4.
Caradoc Series, Slieveroe, Rathdrum, Ireland.
FIG. 10. Holotype, a fragmentary cranidium. National Museum of Ireland Collection, x 2.
Horizon as for Fig. i, locality as for Fig. 2.
FIG. ii. Almost complete cranidium. BM. In. 46440. x 1*4.
Platylichas sp. ..... p. 236
FIG. 9. Internal mould of pygidium, figured by M'Coy, 1846, pi. 4, fig. 14, as Calymene
forcipata. National Museum of Ireland Collection, x i'75-
Caradoc Series, Ballygarvan Bridge, New Ross, Ireland.
Figs. 2,9, 10 by Mr. E. W. Seavill, remainder by the writer.
Bull. B.M. (N.H.) Geol. 7, 8
PLATE 43
LICHAS?. PLATYLICHAS
PLATE 44
Primaspis harnagensis (Bancroft) . . . p. 237
Harnagian Stage, zone of Reuscholithus reuschi, section in old cart-way near southern end of
Smeathen Wood, near Horderley.
FIG. i. Internal mould of cranidium. BM. In. 50802. x 9.
FIG. 6. Paratype pygidium, an internal mould. BM. In. 42087. X2'5.
FIG. 8. Latex cast from external mould of librigena. BM. In. 50803. x 4.
Horizon as for Fig. i. By north bank of Coundmoor Brook, 1300 yards south-west of Harnage
Farm, Harnage.
FIG. 4. Internal mould of cranidium, slightly distorted. BM. In. 52829. x 8.
Primaspis caractaci (Salter) . . . . p. 239
Actonian Stage, site of old quarry, Quarry Field, Gretton, near Cardington.
FIG. 3. Internal mould of cranidium. BM. In 50805. x 4.
FIG. 9. Syn type thorax. GSM. 5214. X5-
FIG. 13. Syntype, internal mould of almost complete dorsal eivoskeleton. GSM. 35473. X4-
Horizon as for Fig. 3. Bank of stream by east side of bridge c. 1000 feet west-south-west of
Hatton.
FIG. 7. Two incomplete cranidia with test preserved. BM. In. 50510. x 3*5. Horizon as for
Fig. 3. Old quarry by road almost opposite Church Farm, Acton Scott.
FIG. ii. Latex cast from external mould of librigena. BM. In. 50801. X4-
Marshbrookian Stage, zone of Onniella reuschi, west side of track just south of quarry in Marsh
Wood, near Marshbrook.
FIG. 14. Poorly-preserved librigena. BM. In. 50799. x 3.
Triarthrus cf. linnarssoni Thorslund . . . p. 241
Onnian Stage, zone of Onnia superba, north bank of River Onny, 720 yards west-south-west of
Wistanstow Church.
FIG. 2. Internal mould of immature pygidium. BM. In. 49320. x 10.
FIG. 5. Incomplete immature cranidium showing position of palpebral lobe. BM. In. 51528.
X5'5-
FIGS. 10,12. Internal mould of cranidium. Shrewsbury Museum 374. X4-
Figs, i, 3, 8, 10-12 by Mr. E. W. Seavill, Fig. 2 by Mr. J. Brown, remainder by the writer.
Bull. B.M. (N.H.) Geol. 7, 8
PLATE 44
PRIMASPIS, TRIARTHRUS
PLATE 45
Otarion sp. . . . . . p. 242
Marshbrookian Stage, zone of Dalmanella unguis, west side of track, 70 yards south of entrance
to quarry, Marsh Wood, near Marshbrook.
FIG. i. Latex cast of cranidium. BM. In. 50618. X4'5.
Remopleurella burmeisteri (Bancroft) . . . p. 250
Onnian Stage, zone of Onnia superba, north bank of River Onny, 720 yards west-south-west of
Wistanstow Church.
FIG. 2. Internal mould of hypostoma. BM. In. 56800^. x 7.
Proetidella fearnsidesi Bancroft . . . p. 243
Harnagian Stage, zone of Reuscholithus reuschi, section in old cartway near southern end of
Smeathen Wood, near Horderley.
FIG. 3. Latex cast from the holotype, an external mould. BM. In. 42083. x 3'4-
FIG. 8. Internal mould of immature thorax. BM. In. 51454. X2-g.
FIGS. 12, 14. Internal moulds of cranidia, showing variation in outline of glabella. Fig. 12,
BM. In. 55449, X 3-4. Fig. 14, BM. In. 51522, x 3-25.
Horizon as for Fig. 3. Exposure in field, 550 yards north-west of Woolston House, Woolston.
FIG. 4. Internal mould of cephalon showing palpebral lobe and facial suture. BM. In. 50616.
X3-
FIG. 5. Internal mould of immature cranidium. BM. In. 51455. X3'5-
Costonian Stage, zone of Harknessella subquadrata, old quarry just west of western end of
Round Nursery, south of Harnage Grange.
FIGS. 6,7. Internal mould of nearly whole dorsal exoskeleton. BM. In. 51152. X3-
Proetidella cf. fearnsidesi Bancroft . . p. 245
Soudleyan Stage, ? zone of Broeggerolithus broeggeri, road-side quarry 150 yards north-east of
Chatwall Farm, Chatwall.
FIG. 9. Internal mould of cranidium. BM. In. 34326. x 4.
FIG. ii. Internal mould of left librigena. BM. In. 34325. X 3'25-
Soudleyan Stage, zone of Broeggerolithus soudlevensis , road-side quarry 100 yards south-east
of Glenburrell Farm, Horderley.
FIG. 10. Internal mould of cranidium. BM. In. 51456. X2.
Proetidella ? sp. . . . . p. 246
Marshbrookian Stage, zone of Dalmanella unguis, stream bank about 350 yards north-west of
road bridge, 350 yards south-west of Hatton, near Acton Scott.
FIG. 13. Internal mould of cranidium. BM. In. 50617. x 4.
Photographs by the writer.
Bull B.M. (N.H.) Geol 7, 8
PLATE 45
OTARION, PROETIDELLA, REMOPLEURELLA
PLATE 46
Remopleurides warburgae sp. nov. . . . p. 246
Highest beds of Actonian Stage, north bank of River Onny, 30 yards east of junction with
Batch Gutter.
FIGS. 1,2. Holotype cranidium with test preserved. BM. In. 49751. X 2-5.
Remopleurides latus Olin onniensis subsp. nov. . . . p. 247
Horizon and locality as for Fig. i.
FIGS. 4, 5. Holotype cranidium. BM. In. 49750. X2'5.
Remopleurides sp. . . . . . p. 248
Horizon and locality as for Fig. i.
FIG. 3. Almost complete left librigena. BM. In. 49752. X2'5_
FIG. 7. Incomplete thorax showing serrated axial rings and shape of pleurae. BM. In. 49753.
X3-
Remopleurella burmeisteri (Bancroft) . . . . p. 250
Onnian Stage, zone of Onnia superba, north bank of River Onny, 720 yards west-south-west
of Wistanstow Church.
FIG. 6. Paratype, internal mould of librigena. BM. In. 42098. x 2-5.
FIGS. 8, 12. Paratype, incomplete, disarticulated exoskeleton preserved as internal mould.
BM. In. 49561. Fig. 8, X2'5. Fig. 12, hypostoma, In. 49561^, x 6.
FIG. 9. Internal mould of cranidium showing palpebral lobes. BM. In. 49748. X5'25.
FIG. ii. Lectotype, internal mould of cranidium. BM. In. 42106. x 275.
Eobronteus ? sp. . . . . p. 251
Horizon and locality as for Fig. 6.
FIG. 10. Internal mould of incomplete dorsal exoskeleton. Sedg. Mus. A. 43833. x 2.
FIGS, i -5, by Mr. E. W. Seavill, Fig. 10 by Mr. J. Brown, remainder by the writer.
Bull. B.M. (N.H.) GeoL 7, 8
PLATE 46
EOBRONTEUS?, REMOPLEURELLA, REMOPLEURIDES
PRINTED IN GREAT BRITAIN BY
ADLARD AND SON, LIMITED,
BARTHOLOMEW PRESS, DORKING
THE GASTROPOD GENUS
THATCHERIA AND ITS
RELATIONSHIPS
A. J. CHARIG
BULLETIN OF
THE BRITISH MUSEUM (NATURAL HISTORY)
GEOLOGY Vol. 7 No. 9
LONDON: 1963
THE GASTROPOD GENUS THATCHERIA
AND ITS RELATIONSHIPS
BY
ALAN JACK CHARIG
Pp. 255-297 ; Plate 47 ; 10 Text-figures
BULLETIN OF
THE BRITISH MUSEUM (NATURAL HISTORY)
GEOLOGY Vol. 7 No. 9
LONDON : 1963
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. 7, No. 9 of the Geological
(Palaeontological] series.
Trustees of the British Museum 1963
PRINTED BY ORDER OF THE TRUSTEES OF
THE BRITISH MUSEUM
Issued July 1963 Price Sixteen Shillings
THE GASTROPOD GENUS THATCHERIA AND
ITS RELATIONSHIPS
By A. J. CHARIG
CONTENTS
Page
I. INTRODUCTION .......... 258
II. HISTORY OF Thatcheria AND OF THE VIEWS ON ITS RELATIONSHIPS 258
III. NOTES ON Thatcheria mirabilis ANGAS ...... 268
Identification of the holotype ....... 268
Additional material in the British Museum (Natural History) . 269
Supplementary description ....... 269
IV. NOTES ON Clinura ......... 271
Material studied . . . . . . . . .271
Posterior sinus ......... 272
Protoconch .......... 274
Transverse sculpture . . . . . . . .274
V. NOTES ON Waitara livatula POWELL . . . . . . 275
Protoconch .......... 275
Transverse sculpture . . . . . . . .276
VI. SYSTEMATICS .......... 276
General principles . . . . . . . . .276
The Clinura-Thatcheria group . . . . . . .278
Division into genera . . . . . . . .281
Interspecific differentiation in Thatcheria . . . . .283
Evolution in the Clinura-Thatcheria group .... 286
Systematic position of the Clinura-Thatcheria group . . . 288
Formal classification . . . . . . . .291
VII. A NEW SPECIES OF Thatcheria FROM THE PLIOCENE OF FIJI . 292
VIII. ACKNOWLEDGMENTS ......... 294
IX. REFERENCES .......... 295
SYNOPSIS
The literature on the subject is reviewed with particular reference to the systematics of the
genera concerned.
A Recent gastropod shell, Brit. Mus. (Nat. Hist.) no. 1960.154, is identified as the holotype
of the type-species of Thatcheria (T. mirabilis from off Japan) . The description is amplified.
The form of the posterior sinus in Clinura has been generally misunderstood. The proto-
conch, hitherto unknown in the genus, is described in the type-species and in one other. The
protoconch of Waitara liratula is re-examined.
The systematics of the genera concerned are reconsidered. Clinura is restricted to a few
species from the Neogene of Europe and of the Western Pacific. Clinura, Waitara and
Thatcheria are closely related to each other but not to Surculites. Fourteen species in those
three genera are reclassified into two genera, Clinura (mainly Miocene) and Thatcheria (Upper
Miocene to Recent, mainly Pliocene) ; Thatcheria is confined to the Western Pacific. The two
genera constitute a sub-family of the Turridae, the Thatcheriinae, which has affinities with
the Daphnellinae.
Thatcheria vitiensis sp. nov. is described and figured from one specimen of probable Lower
Pliocene age from Fiji.
GEOL. 7, 9 20
258 GASTROPOD GENUS THATCHERIA AND ITS RELATIONSHIPS
I. INTRODUCTION
THE discovery of a new fossil gastropod which resembles Thatcheria and Waitara
has led to a re-examination of this little-known group of genera. The conclusions
reached have been based upon all the relevant literature and upon a few shell
characters hitherto unknown or imperfectly understood.
Some of the earlier descriptions and illustrations of the gastropods in question
were inadequate or misleading (especially with reference to the form of the posterior
sinus and of the consequent growth-lines, the taxonomic significance of which had
not then been appreciated). One drawing in particular, upon which some later
workers relied for their knowledge of a type-species, is also quite different in that
respect from the less accessible description and figures of the original author. The
extreme scarcity of actual material obliged most later workers to rely very largely
upon these earlier descriptions and figures, and misunderstandings therefore arose.
Language difficulties appear to have exacerbated the situation. Finally, it is also
evident that certain important works on the Thatcheria-Waitara group of gastropods
were written in ignorance of the existence of some of the others. But this was
sometimes unavoidable ; for example, two of the most important were written and
published at about the same time during the Second World War, one in New Zealand
and the other in the German-occupied Netherlands.
Thus, while the known members of the group are few in number, both as species
and as individuals, their systematics are confused. Although only two named
species (Thatcheria mirabilis and T. gradata) have been referred to the genus That-
cheria itself, those two species have already been placed by various authors in ten
different prosobranch taxa of the genus-group (excluding synonyms). They have
been considered, at least implicitly, as belonging to eight different prosobranch
families, five of which possess alternative names. Indeed, one author alone (Wenz,
1943) has referred the two forms concerned and the closely related genus Waitara
to three separate families. It has also been suggested that T. mirabilis is an opistho-
branch, and elsewhere stated quite dogmatically that it is a " scalariform mon-
strosity ". In all, in connexion with T. mirabilis and T. gradata, sixteen family
names are involved.
However, it is now generally accepted that these molluscs are so closely related
as to form a natural group. The group might be either a sub-family within the
Turridae or, as has been suggested, a separate family with strong affinities therewith.
The status and relationships of the group require careful consideration, and the
systematics within the group are in need of revision.
II. HISTORY OF THATCHERIA AND OF THE VIEWS
ON ITS RELATIONSHIPS
Thatcheria mirabilis gen. et sp. nov. was described and figured by Angas (1877 :
529, pi. 54, figs. la, b}. His material consisted of one fairly large pagodiform shell
believed to be unique ; it had been brought from Japan not long before by Mr.
Charles Thatcher, and its occurrence could be localised no more accurately than
" Seas of Japan ". No indication was given as to where the shell was deposited.
GASTROPOD GENUS THATCHERIA AND ITS RELATIONSHIPS 259
Of its systematic position, Angas wrote : " Without a knowledge of the operculum, its
exact generic position cannot be determined ; but at present I regard it as belonging
to the subfamily Fusinae " (i.e. in the family then called Fusidae, now either called
Fusinidae or regarded as part of the Fasciolariidae).
The next reference to the specimen is by G. B. Sowerby (1880 : 105, pi. 422, figs.
45, 46), who wrote that it " has been thought to exhibit, in the arched sinus of the
outer lip above the angle, a peculiarity of generic value." He nevertheless referred
the species to the genus Pyrula. (The " Thesaurus Conchy liorum " does not place
genera in families, but the genus Pyrula Lamarck 1799, which should be replaced
by the older synonym Ficus Roding 1798, is now referred to its own family Pyrulidae
or Ficidae). Sowerby also considered Angas' type to be immature, and mentioned
that it was " in the collection of Mrs. Deburch ".
Tryon (1881 : 98, 112, pi. 44, figs. 238, 239) retained Angas' genus Thatcheria
and referred it to the sub-family Melongeninae or Melongeniinae of the family Buc-
cinidae ; but his arrangement might indicate that he considered Thatcheria to be
synonymous with the older genus Hemifusus Swainson. Further, he opined that
Angas' specimen did not represent the normal form of the genus, for he wrote " That
this shell is a scalariform monstrosity cannot be doubted ". Later (1883 : 135, pi.
49, fig. 5) he gave an exact repetition of his earlier text.
Fischer (1884 : 623) reported Tryon as considering Thatcheria to be a scalariform
monstrosity of " Semifusus Swainson em. 1840 (Hemifusus}." In fact, Semifusus
Agassiz 1846 is an invalid emendation of Hemifusus Swainson 1840. Fischer placed
this genus in the sub-family Melongeninae of the family Turbinellidae (sometimes
called Xancidae or Vasidae) ; the sub-family in question is now generally regarded
as a separate family, the Melongenidae.
Cossmann was at first (1889 : 162) unable to agree with Tryon on this matter ;
the form of the sinus, as indicated by the growth-lines on the ramp, led him to
believe that Thatcheria could well be a junior synonym of Mayeria Bellardi 1873.
Mayeria, in any case, seemed to be close to " Semifusus " , for Cossmann placed them
both together (in the same family and sub-family as Fischer had done). Later,
however (1901 : 62, 93, 94), Cossmann decided that the canal of Thatcheria dis-
tinguished it from Mayeria, and that Tryon was correct after all in supposing
Thatcheria to be but a monstrosity of " Semifusus ".
Meanwhile Pilsbry (1895 : 28) had reverted to Tryon 's classification and had listed
Thatcheria mirabilis in the Buccinidae.
No further mention of Thatcheria has been discovered earlier than Tomlin's
editorial notes (1919 : 66), in which he stated that the de Burgh Collection was offered
for sale and that it included Thatcheria mirabilis, frequently considered to be a
monstrosity of Fusus. (Presumably Tomlin intended this to refer to the genus
Fusinus Rafinesque 1815 emend, pro Fusus Lamarck 1799, non Helbling 1779).
Yokoyama (1928 : 338, pi. 66, figs. 3, 4) described and figured two specimens from
the Pliocene of Japan as Cochlioconus gradatus gen. et sp. nov. The new genus was
not explicitly referred to any family, but it was described as " C onus-like " and
Yokoyama stated that " The presence of a deep sutural notch reminds us of the
genus Conus." Pilsbry saw this paper and wrote to Yokoyama, drawing his attention
GEOL. 7, 9 20§
260 GASTROPOD GENUS THATCHERIA AND ITS RELATIONSHIPS
to the similarity between Cochlioconus and Thatcheria. Yokoyama then stated in a
subsequent paper (1930) that he regarded Thatcheria (T. mirabilis} and Cochlioconus
(C. gradatus] as congeneric, though not conspecific, the Pliocene species being more
Conus-like in appearance than the Recent form. He did not, however, change his
views on the systematic position of the genus, for he wrote (p. 406) " I think I am
right in placing it near Conus. This opinion seems to be also shared by Mr. Pilsbry
as is evident from his communication." He also stated, probably wrongly, that
Angas' specimen of T. mirabilis " may be the only one now existing [i.e., in a col-
lection] in the whole world."
Thiele (1929 : 320) considered Thatcheria to be a synonym of the sub-genus
(" sectio ") Semifusus sensu stricto, which he placed in the family Galeodidae.
The proper family name is in fact Melongenidae, the name of the type-genus Galeodes
being a junior homonym (Roding 1798 non Olivier 1791). Thiele, who was con-
cerned only with Recent molluscs, did not mention Cochlioconus.
S. Hirase (1934 : 104, pi. 1286, fig. 3) figured a specimen of " Thatcheria mirabilis
Angas (= Semifusus m.?} ", which is appreciably longer and more slender than
Angas' type. The illustration is of interest in that it showed, for the first time, that
more than one individual of the species had been found.
This point was emphasised in 1937 at an Ordinary Meeting of the Malacological
Society of London, at which Tomlin exhibited two shells of the species, both from
Japan. Hopwood wrote in the Proceedings (1937 : 158) that " Tryon's dictum
[that Thatcheria is a scalariform monstrosity] has been confuted recently by the
discovery of several examples, but where it should be placed systematically is as
great a problem as ever. Discussion elicited suggestions that it was a prosobranch
to be placed in Turridae and that it was an opisthobranch allied to Akera." The
suggestion that it was a turrid had not been made before.
The first recorded find of the soft parts of Thatcheria was made at about this time ;
Tomlin had obtained from a Japanese fisherman the anterior portion of a male ani-
mal, which was described and figured by Eales (1938). Unfortunately the specimen
had lost the radula. Eales concluded, however (p. 16), that its characters " show
that the specimen belongs to the Toxoglossa." A comparison of the creature with
various members of the Turridae (e.g., Turricula javana) and of the Conidae (e.g.,
Conus quercinus] showed a very close resemblance to the former and none whatever
to the latter ; and an X-ray photograph of the shell of T. mirabilis, which " shows
a typical columella similar to that of the Turrids, not resorbed as in the Cones ",
confirmed Eales' opinion (p. 17) that " As far as it is possible to judge . . .
Thatcheria mirabilis should be placed with the Turrids."
The next paper to mention Thatcheria was that of Powell (1942), in which he
suggested a close affinity between Thatcheria and the fossil Waitara. At this point
it is necessary to digress a little upon the latter genus.
All the species mentioned immediately below and referred to Waitara are from
New Zealand. Marwick (1926 : 324, pi. 74, fig. 9) had described a specimen from
the Upper Miocene under the new specific name waitaraensis, referring it to the
genus Turricula ; but he also wrote (p. 326) that " the generic location of the Turridae
described above is quite provisional. New genera seem to be required ". Later
GASTROPOD GENUS THATCHERIA AND ITS RELATIONSHIPS 261
(1931 : 149) he erected the new genus Waitara upon this species, and referred thereto
another new Miocene species, W. generosa (pi. 18, fig. 339). Powell (1942 : 168)
considered W. generosa to be of Lower Miocene age. Powell also described two
more species of Waitara in the same work — W. pagodula (p. 168, pi. 14, fig. 7) from
the Upper Miocene or Lower Pliocene and W. liratula (p. 169, pi. 14, fig. 8) from the
Lower Pliocene — and mentioned yet another Lower Pliocene species (p. 168), repre-
sented by a single specimen too poorly preserved for description or illustration but
said to be " somewhat similar [to W. pagodula] but narrower ". The Waitara
liratula material included the protoconch illustrated in his text-fig. 63 (p. 38). The
paper gave a key to the four named species of Waitara (p. 168).
Powell's systematic conclusions were important. He wrote (p. 167) " It is possible
that Waitara may yet prove to be identical with Thatcheria . . . and Cochlioconus
... In any case all three are closely allied and represent a discordant Turrid-like
group that cannot be satisfactorily placed in any of the nine sub-families adopted in
this bulletin. . . . Cochlioconus is almost certainly a synonym [of Thatcheria],
although its author made no reference to Thatcheria, but compared his genus with the
Cones. Anyone acquainted with Thatcheria would scarcely have overlooked the need
for some comparison." Evidently Powell did not know of Yokoyama's second paper
on this subject ; nor, indeed, does he seem to have been aware of Eales' work.
He sought to differentiate Waitara from Thatcheria, at least provisionally, because
its sinus " although very similar to that of Thatcheria, does differ at its point of
origin, in being narrowly concave before swinging forward, whereas the sinus in
Thatcheria swings forward immediately." He also described the form of the sinus
in all these genera as " deep Conid-like." Finally, after discussing the relationship
of the genera in question to certain sub-families of the Turridae (the Daphnellinae
and the Cochlespirinae) and to the Conidae, Powell came to the following conclusions
(p. 168) :
" In order not to prejudice the status of the other Turrid subfamilies by forcing
in such an aberrant group, it seems advisable to consider Thatcheria and its allies
as representative of a new family, closely akin or parallel to the Turridae, for it
seems to have arisen from the Conidae, but probably much later and independent of
the early Conid-like Turrid divergent stock as represented by the Conorbiinae."
He called this new family the Thatcheriidae ; and, in the same work (p. 170), he
wrote " The former [i.e. the Thatcheriidae] in having marked Conid affinity in
respect to its sinus is indicated as a late Tertiary offshoot from the Conidae . . .
Wenz (1943) evidently saw no connexion between Thatcheria, Cochlioconus and
Waitara, for he placed all three in different families. He considered Thatcheria
(p. 1215) to be a synonym of the sub-genus Hemifusus of the genus Pugilina Schu-
macher 1817, family Galeodidae. Cochlioconus (mis-spelt " Cochliconus " in the
text on p. 1470 and in the index on p. 1604, though correct in the legend to fig. 4154)
he regarded as a sub-genus of Conus Linnaeus 1758, family Conidae. And he classi-
fied Waitara (p. 1390, fig. 3929) as a genus of the sub-family Clavatulinae, family
Turridae ; he wrongly cited W. generosa Marwick as the type-species, wrongly
quoted its horizon as Upper Miocene and gave the stratigraphical range of the genus
as Oligocene to Miocene, although no Oligocene species of Waitara has ever been
262 GASTROPOD GENUS THATCHERIA AND ITS RELATIONSHIPS
recorded. (The Pliocene species of Waitara described by Powell (1942) were, of
course, unknown to Wenz because of the Second World War.) The genus Clinura
Bellardi 1875 (see below) was also included by Wenz (p. 1390, fig. 3928) in the
Clavatulinae, as a sub-genus of Surculites Conrad 1865.
Beets (i943# : 296) described as Surculites (Clinura) bituminatus a new species of
gastropod from the asphalt deposits of the island of Buton, off Celebes ; the deposits
were at that time believed to be of Upper Oligocene age. Like Wenz, Beets re-
garded Clinura as a sub-genus of Surculites and considered these forms to be turrids.
In his description of S. (C.) bituminatus he placed Cochlioconus in the synonymy of
Clinura ; but, when doing so, he knew nothing of Thatcheria, for it was not until later
that Yokoyama's second paper (1930) drew his attention towards that genus.
Perhaps the most important work of all on this group of gastropods was a second
article by Beets (19436). He suggested that Thatcheria (including, ywfc Yokoyama,
1930, Cochlioconus}, Clinuropsis Vincent 1913 (non Thiele " 1931 " [should be 1929])
and Nekewis Stewart " 1926 " [should be 1927] should all be regarded as junior
synonyms of the sub-genus Clinura. In this paper Beets discussed a large number
of species — of Clinura, of Clinuropsis, and of other genera such as Cryptoconus,
Surculites, " Pseudotoma ", " Surcula ", " Pleurotoma " and "Turricula " — in order
to ascertain whether they might properly be included in Clinura ; and, since he
considered their great variations in form (especially that of the posterior sinus) to
fall within the range of the "fur die so schwierigen Turridae ' normalen Variabilitat '",
his decision was usually affirmative. He concluded (p. 365) : " Clinura ist jetzt
mit Gewissheit bekannt aus dem Eocan bis Miocan der atlantischen [surely that
should be " pazifischen "?] Kiistenregionen Nordamerikas . . . , aus dem Montien
bis Pliocan Europas und dem Oligocan des ostindischen Archipels ; in Japan tritt sie
erst ins Pliocan auf und lebt dort mit einer einzigen, extremen rezenten Art fort.
Vielleicht (?) ist Clinura auch noch bekannt aus dem Eocan S.W. Afrikas und des
Congos, hochstens mit zwei Arten."
At this point it may be useful to give a short account of Clinura and of its author.
The genus was erected by Aloysius Bellardi (1875 : 20) in a paper written entirely
in Latin ; although "A. Bellardi " and " L. Bellardi " (Luigi) are sometimes listed
separately in bibliographies (e.g., Wenz, 1944 : 1513), there are several reasons
which make it seem certain that " Aloysius " was merely a latinisation of " Luigi "
and not the name of another author. First, both " Aloysius " and Luigi were
Professors of Natural History, living at the same time in the same country and
writing on the same subject ; indeed, not only " Aloysius " but also Luigi published
works concerned with Clinura, including the one mentioned below in which Luigi
quoted verbatim (but without acknowledgment) the generic diagnosis given by
" Aloysius " and then designated a type-species. Secondly, both " Aloysius " and
Luigi published in the Bullettino delta Societa Malacologica Italiana ; and authors
of works in that journal were generally members of the society in question, yet only
Luigi's name is to be found in the list of members for 1875. Thirdly, " Aloysius "
(unlike Luigi) published nothing except this one paper. Fourthly, neither " Aloy-
sius " nor Luigi, when referring to names proposed by a Bellardi or to previous
publications by a Bellardi, was in the habit of distinguishing himself from the other
GASTROPOD GENUS THATCHERIA AND ITS RELATIONSHIPS 263
by the use of an initial. Fifthly, a paper in French by " Louis " Bellardi (1841) is
clearly the work of Luigi ; it is therefore evident that Luigi was in the habit of
altering his Christian name to conform with the style of the language in use.
Finally, an obituary of Bellardi by his fellow malacologist Sacco (1889) gives a list
of the " Publications scientifiques de M. Louis Bellardi " ; this includes all the works
of " Aloysius ", Luigi and " Louis ", and thus provides conclusive proof of their
identity.
The 1875 paper designated no type-species for Clinura but assigned two species to
the new genus, C. calliope (Brocchi, 1814 : 436, pi. 9, figs. 150, b) [Murex] and C.
elegantissima (Foresti, 1868 : 598, pi. 2, figs. 10-13) [Pleurotoma] ; Bellardi placed
Clinura in his new sub-family Pseudotominae, family Pleurotomidae. A far more
complete account of the genus, published a few years later in Italian by Luigi Bellardi
(1877 : 204-209), included five species therein ; Clinura calliope, from the Upper
Miocene of Italy, was clearly designated as the type (p. 204). Other fossil species
were subsequently referred to the genus, at first only from the Neogene of Europe,
e.g., R. Homes & Auinger, 1891 : 362 ; Friedberg, 1912 : 210) ; later, however,
species from rocks as old as the Eocene and as far away as California and Celebes were
also placed in Clinura (e.g., Grant & Gale, 1931 : 494 ; Beets, 1943^, I943&, as
indicated above).
Meanwhile two Recent deep-sea gastropod species had also been referred to
Clinura, C. monochorda Ball (1908 : 292, pi. 13, fig. i) and C. peruviana Dall (1908 :
293, pi. 13, fig. 2). Since these are the only Recent species ever referred to the genus,
they are also the only species mentioned by Thiele (1929 : 371), whose work did not
deal with fossil forms. Thiele wrote of Ball's species, however, that " doch diirften
sie mit dieser [Clinura} kaum verwandt sein ", without giving any reason for this
statement, and he proposed that these two species should constitute a new section
Clinuropsis of the genus Pleurotomella Verrill 1873, with P. (C.} monochorda (Ball
1908) as the type-species. But, by the time he wrote his " Nachtrage und Berichti-
gungen " to the same work, published in 1934, Thiele had learnt that the name
Clinuropsis had already been used by Vincent in 1913 ; he therefore proposed
(p. 1002) the new name Anticlinura to replace it.
Wenz (1943 : 1460) treated the supposed Recent species of Clinura exactly as
Thiele had done. Beets (19436 : 364) also agreed with Thiele (1929) in the matter,
Ball's species being two of the few which he chose to exclude from the genus Clinura ;
unlike Thiele, he gave reasons (such as the presence of lattice-like sculpture on Ball's
species) for so doing. Further, Beets appears to have noted Vincent's preoccupation
of the name Clinuropsis but not Thiele's subsequent correction and proposal of
Anticlinura nom. nov. to replace Clinuropsis Thiele ; for he himself, with the same
intention, proposed Clinuromella nom. nov. Thus Clinuromella Beets 19436 is an
objective junior synonym of Anticlinura Thiele 1934, both being new names for
Clinuropsis Thiele 1929 (non Vincent 1913). In conclusion, it would seem that
Ball's two species should continue to be placed under the sub-generic name Anti-
clinura (of which monochorda is the type-species) in the genus Pleurotomella.
No author has ever disputed the position of Clinura in the Turridae (Pleuroto-
midae). Cossmann (1896) and Friedberg (1912) both regarded the taxon as a sub-
264 GASTROPOD GENUS THATCHERIA AND ITS RELATIONSHIPS
genus of Surcula H. & A. Adams 1853 ; but Grant & Gale (1931) considered it to be
a sub-genus of Surculites, in which opinion, as recorded above, they were followed
by Wenz (1943) and Beets (19430 ; 19436). Wrigley (1939), however, while not
denying the turrid (or at least toxoglossan) nature of Clinura, believed that Surculites
was not a turrid and therefore not related to Clinura.
Thus the connexion of Thatcheria with Clinura was suggested and firmly established
in 1943. The rest of this historical account will deal also with papers referring to
Clinura, even though they make no mention of Thatcheria.
Though complete in most respects, Beets' paper (19436) had not considered
Waitara. Powell's bulletin on the Turridae, published in New Zealand in 1942,
drew attention to the relationship between Waitara and Thatcheria, but had not
been seen by Beets working in that same year in the German-occupied Netherlands ;
nor could he have seen Wenz's treatise (1943) which placed Waitara immediately
next to Clinura. In 1949, however, Beets obtained access to Powell's bulletin ;
and in 1951 he published yet another paper, one part of which was concerned with
" Waitara and its relationships to Surculites-Clinura-Thatcheria." The main object
of this work was to establish Waitara as part of " the species group Clinura . . .
(taken in a broad sense) " ; the three taxa Clinura, Waitara and Thatcheria, hitherto
related only as three different pairs by three different authors, were to be brought
together. Beets recognised, however, that Wenz's conclusions regarding the
taxonomic proximity of Waitara and Clinura had been based entirely on Marwick's
original Miocene species of the former genus, while the affinities of that genus to
Thatcheria depended more on the characters of the younger species described by
Powell (1942).
One noteworthy point from Beets' 1951 paper is that, whereas he had earlier
(19436) considered Thatcheria to be a synonym of the sub-genus Clinura, he now
regarded Clinura, Waitara and Thatcheria as three separate (but closely related)
sub-genera, still in the genus Surculites. He gave no reason for this change. In-
cidentally, this work of Beets contains three minor errors. First, he stated
(p. 14) that Turricula waitaraensis Marwick became the type-species of Waitara
by Powell's subsequent designation in 1942 ; it was, in fact, the type-species
by Marwick's original designation (1931). Secondly, he appeared to claim
(p. 15) that he himself was the first to refer Thatcheria to the Turridae (19426, cited
in the present work as 19436) ; he was, of course, preceded in this by Eales (1938).
Thirdly, he quoted Powell (p. 16) as describing the posterior sinus of Waitara as
" first narrowly convex " ; that should read " concave ".
Beets (1951) also gives a small, semi-schematical figure (pi. i, fig. 6) of " Thatcheria
spec. nov. ; a portion of a specimen from the Pliocene of East-Borneo ". The
fragment was mentioned only briefly in the text (p. 16) and has not yet been des-
cribed. It appears to consist of two whorls from a pagodiform spire which must
indeed have been remarkably like that of T. mirabilis, with a similar type of sinuosity
in the growth-lines on the ramp.
The asphalt deposits of Buton (which, according to Beets (19430 ; 19436), con-
tained two species of Clinura) were supposed to be of Upper Oligocene age. In
1953, however, Beets showed (p. 239) that the age of the moUuscan fauna was most
GASTROPOD GENUS THATCHERIA AND ITS RELATIONSHIPS 265
likely Mio-Pliocene ; there is no evidence to support the suggestion that the molluscs
might be a mixture derived from different stratigraphical horizons, but the possibility
cannot be entirely excluded. He also reported Reinhold as concluding that the
diatoms in the deposits indicate a probable Upper Miocene age.
S. Hirase's posthumous handbook of Japanese shells (1951), which was in effect
his catalogue of 1934 revised and enlarged by Taki, reproduced the figure of Thatcheria
mirabilis which had appeared in the earlier work (pi. 1286, fig. 3, in both). The
later work, however, no longer suggests that Thatcheria might be a synonym of
" Semifusus " ; and the second edition (1954) is provided with a systematic index
to the plates which, on p. 88, classifies Thatcheria as a genus of the Turridae.
Thatcheria was also listed among the Turridae by Kuroda & Habe (1952 : 10).
On p. 90 of the same work they indicated that the geographical range of T. mirabilis
was off the Pacific coast of Japan, between 33° and 35° of latitude.
Hatai & Nisiyama (1952 : 191) listed Yokoyama's two syntypes of Cochlioconus
gradatus with full details of locality and horizon ; the latter was given as Takajo
Formation, believed to be of Lower Pliocene age. It was noted that both specimens
were in the collections of the Geological Institute of the Faculty of Science of Tokyo
University. The specimen illustrated in Yokoyama's fig. 3 (1928, pi. 66) was
designated as lectotype (" holotype ") by these authors, who evidently regarded
C. gradatus as a junior synonym of Thatcheria mirabilis.
Kuroda & Habe (1954 : 80) published important new information on Thatcheria
mirabilis ; this was obtained from three individuals of different ages, each with soft
parts. They were able to describe and figure the radula (text-fig. 2) and the proto-
conch (text-fig. I, showing also the early nepionic whorls) ; the operculum was
reported lost in all the specimens. The peculiar shape of the radula and the obliquely
reticulated sculpture of the protoconch led them to suggest that Thatcheria was a
member of, or had a close affinity with, the Daphnellinae. Unfortunately they had
misinterpreted both Eales' and Powell's papers. First, they thought that Eales'
observation, " No traces of proboscis or radula are present ", indicated the natural
lack of a radula in the species ; but, in fact, the lack of a radula was clearly accidental
in the specimen which constituted the whole of Eales' material, " the body being
torn away immediately posterior to the mouth tube." Secondly, they believed
that Powell had established the family Thatcheriidae to include, not only Thatcheria
mirabilis and some fossil species, but also some other, related Recent groups (but
no others are known) ; and further, that Powell had done this because of the
" nature of its radula quite rudimentary " in Thatcheria (also unknown at that time).
Habe (1955) devoted two short articles to " Thatcheria mirabilis Angas (Turridae) ".
The first is in English ; it consists only of a direct quotation of Angas' description,
reproductions of the figures of the early whorls and the radula (from Kuroda &
Habe 1954), references to the works of Angas, Eales, Powell, and Kuroda & Habe,
and the comment " This strange species seems better to locate under the sub-
family Daphnellinae in having the reticulated protoconch and the dart shaped
radula." The other article is in Japanese but is not the same ; it reproduces the
illustrations to Angas' paper, cites the dimensions of three new specimens of T.
mirabilis and mentions the fossil Cochlioconus.
266 GASTROPOD GENUS THATCHERIA AND ITS RELATIONSHIPS
Kira (1955 : 71, pi. 35, fig. 19) also placed Thatcheria mirabilis in the Turridae.
Rossi Ronchetti (1951-56) included Murex calliope, the type-species of Clinura
(see p. 263), among the species dealt with in her review of Brocchi's types ; she
selected, figured and carefully described a neotype (1955 : 305, fig. 163) because
Brocchi's holotype was no longer preserved. The species appears, however, under
the name Pleurotomella (Clinuropsis) calliope, presumably for the following reasons :
The introduction to Rossi Ronchetti's review stated (1951 : n) that the revision
of the generic names would be based partly upon the works of Wenz (1938-44) and
Schilder (1932), partly upon the works of Thiele (1929-31, 1934-35). Schilder,
of course, dealt only with Cypraeacea. In this particular instance Rossi Ronchetti
could not have used Wenz (or Thiele's " Nachtrage und Berichtigungen ") as the
basis for her classification ; had she done so, she would then have known— as she
clearly did not know — that Thiele had later proposed Anticlinura as a new name to
replace his own Clinuropsis. It therefore appears likely that she based her classifica-
tion only upon Thiele's main work. Noting that Ball's species of Clinura, the only
supposed representatives of that genus mentioned by Thiele, had been transferred
by the latter author to Pleurotomella (sectio Clinuropsis), Rossi Ronchetti did the
same with Clinura calliope — probably without considering the possibility that Ball's
species had no real connexion with Clinura.
Although other authors may not agree that calliope Brocchi and monochorda
Ball are co-sub-generic, or even congeneric, Rossi Ronchetti is nevertheless entitled
to her opinion that they are. In that case she was correct in using for both species
the oldest generic name available — Pleurotomella Verrill 1873. But, for the sub-
genus, she should then have used the oldest generic name previously applied to any
included species — and this was Clinura Bellardi 1875, not Clinuropsis Thiele 1929
(which, in any case, was preoccupied). Moreover, according to Grant & Gale (1931 :
510), Pleurotomella Verrill 1873 (type-species P. packardii Verrill 1873) is a sub-
jective junior synonym of Pleurotomoides Bronn 1831 (type-species Defrancia pagoda
Millet 1826).
With respect to Clinura, Eames (1957 : 51) adopted the classification of Grant &
Gale, Wenz, and Beets, in which Clinura appears as a sub-genus of Surculites in the
Turridae. But he referred the Eocene species Surcula ingens (Mayer-Eymar 1896)
[Pleurotoma] to Clinura, although Beets (1943^ : 363) had clearly expressed the
opinion that it was not related thereto and represented an altogether different group.
The Fiji Geological Survey Bepartment, in its Annual Report for the year 1958
(1959), mentioned on p. 15 the collection of an unusual fossil shell from the island of
Vanua Levu, identified by the present writer as a new species of Thatcheria. The
Report also quoted his observation (in litt.) " ... as far as I am aware, the genus
has never been recorded as a fossil."
Several gastropod species represented in the collections of the Institut Royal
des Sciences Naturelles de Belgique were listed by Glibert (1960 : 25) under the
generic name Clinura. They are all from the Tortonian or Plaisancian of Italy or
the Vienna Basin. Glibert classified Clinura as a genus of the sub-family Cochles-
pirinae (family Turridae).
GASTROPOD GENUS THATCHERIA AND ITS RELATIONSHIPS
267
TABLE I. — SUMMARY OF VIEWS ON SYSTEMATIC POSITION OF THATCHERIA
(i.e., of the only two named species hitherto referred to that genus)
Author
Sp.
Sub-genus Genus Sub-family
Family
Sowerby, 1880
m
. Pyrula (juvenile)
[Pyrulidae
( = Ficus}
(— Ficidae)]
Tryon, 1 88 1, 1883 .
m
Thatcheria, a . Melongeninae, "*
scalariform Melongeniinae
monstrosity listed
^ Buccinidae
under Hemifusus
Pilsbry, 1895
m
. Thatcheria
Thiele, 1929 .
m
Semifusus s.s. . Semifusus
(= Hemifusus s.s.) (= Hemifusus}
Galeodidae
Wenz, 1943 .
m
. Hemifusus . Pugilina
(= Melongen
(Pchelintsev &
m
. Hemifusus
dae)
Korobkov, 1960
Fischer, 1884 \
Cossmann, 1901 j
m
A scalariform
monstrosity of
Semifusus
Melongeninae
Turbinellidae
( —Hemifusus)
Cossmann, 1889
m
. Mayeria J
Angas, 1877 .
m
. Thatcheria . Fusinae "1
(= Fusi-
Tomlin, 1919
m
ninae) J
. A monstrosity of . [Fusinae [
Fusus (= Fusi- '|
[Fusidae
(— Fusinidae
( = Fusinus} ninae)] J
Hopwood, 1937
m
. Thatcheria
? Turridae
Eales, 1938 .
~\
Kuroda&Habe, 1952
\
. Thatcheria
Hirase & Taki, 1954
\
Kira, 1955
>• .
Turridae
Beets, 19430 .
g
. Clinura . Surculites
Beets, 19436 .
m g
. Clinura . Surculites
Beets, 1951
m g
. Thatcheria . Surculites
Kuroda & Habe,
m
. Thatcheria . Daphnellinae .
[Turridae]
1954
Habe, 1955 .
m g
. Thatcheria . Daphnellinae
Turridae
Powell, 1942 . \
MacNeil, 1960 J
m g
. Thatcheria
Thatcheriidae
Yokoyama, 1928
Yokoyama, 1930
g
m g
. Cochlioconus \
. Thatcheria f
[Conidae]
Wenz, 1943 . "1
Pchelintsev & >
g
. Cochliconus . Conus
Conidae
Korobkov, 1960 J
(= Cochlioconus}
Hopwood, 1937
m
. Thatcheria
? [Akeridae]
Charig, 1963 .
me
. Thatcheria . Thatcheriinae
Turridae
Note:
m indicates application to mirabilis Angas.
g indicates application to gradata Yokoyama.
Square brackets [ ] denote implicit references.
Where an author has used a junior synonym or homonym, or family name derived therefrom, the nan
considered here to be correct is also given— in parentheses.
GEOL. 7, 9
20§§
268 GASTROPOD GENUS THATCHERIA AND ITS RELATIONSHIPS
The Osnovy Paleontologii (1960), edited by Pchelintsev & Korobkov, adopts
a very conservative arrangement which seems to be based on that of Wenz ; the
important works of Eales, Powell, Beets, and Kuroda & Habe are all ignored.
Thatcheria (p. 222) is given as a synonym of the genus Hemifusus, family Galeodidae ;
Cochlioconus (p. 241), again mis-spelt " Cochliconus ", appears as a sub-genus of
Conus, family Conidae ; Waitara (p. 240) is cited as a genus incertae sedis in the
Pleurotomidae ; and, in that same family, Clinura (p. 239) is still considered to be
a sub-genus of Surcula.
MacNeil (1960 : 120, pi. 15, figs, u, 12) described two shells from the Neogene of
Okinawa, now in the United States National Museum, as Thatcheria cf. T. gradata
(Yokoyama) ; he retained Powell's family Thatcheriidae for the genus. The
figured specimen was from the Pliocene ; the other was from rocks which might be a
little older, perhaps of either [Lower] Pliocene or [Upper] Miocene age. Both were
compared with T. gradata rather than with T. mirabilis because of their plane to
slightly concave ramp (the ramp in T. mirabilis is " definitely depressed with the
periphery slightly upturned "), the coarser sculpture, the lack of crowded spirals just
below the periphery, and (presumably) the less pagodiform spire. The excellent
preservation of the Okinawan specimen figured by MacNeil, with the sculpture
preserved nearly to the protoconch stage, shows that " The early sculpture is strongly
turrid, the periphery being ornamented with well defined blunt denticulations,
about 9 or 10 visible from an angle, and the denticulations themselves are crossed
by fine spiral lines " ; in this respect comparison with Yokoyama's material is
impossible, for the early whorls are not preserved in either of the Japanese specimens.
Nor, for that matter, are they preserved in MacNeil's other Okinawan (possibly
Miocene) specimen.
MacNeil also expressed some slight doubt as to whether the young shell figured by
Kuroda [& Habe] (1954) was really a Thatcheria, pointing out that the first nepionic
whorl is quite different from that of the Okinawan form. The figured specimen from
Okinawa, however, lacks more of the apex than is represented by the whole of
Kuroda & Habe's figure, and therefore a proper comparison cannot be made.
III. NOTES ON THATCHERIA MIRABILIS ANGAS
Identification of the holotype
It was suspected that the holotype of the species might be a shell in the collection
of the Department of Zoology of the British Museum (Natural History), recently
registered under the number 1960.154. The only available information pertaining
to this specimen is that it formed part of the de Burgh Collection.
As mentioned above (p. 258), Angas gave no indication as to where his unique
shell was deposited. But Sowerby wrote that it was " in the collection of Mrs.
Deburch " ; and a copy of Angas' original paper, now in the possession of Mr. S. P.
Dance, bears the annotation " de Burgh Coll." in Fulton's handwriting. Finally,
as also mentioned above, the de Burgh Collection was offered for sale in 1919 and
GASTROPOD GENUS THATCHERIA AND ITS RELATIONSHIPS 269
included material of T. mirabilis. Although it was not stated explicitly that there
was only one specimen, there is nothing to suggest that there was more than one ;
indeed, it was not until the nineteen-thirties that it became apparent that T.
mirabilis was not just a unique monstrosity.
The British Museum specimen referred to has therefore been compared with
Angas' description and illustrations. Its dimensions and individual peculiarities,
especially the form of the mended break in the outer lip, show that, while the draw-
ings (which bear the indication " G. Sowerby lith.") are not wholly accurate, the
common peculiarities are so many that they cannot be merely fortuitous.
It may therefore be presumed that the B.M. specimen, registered number 1960.
154, is the holotype of Thatcheria mirabilis Angas.
Additional material in the British Museum
(1) The incomplete soft parts described by Eales (1938) (B.M. regd. no. 1937-12-
16 . i) . This specimen was obtained at Kii, Japan, and was presented to the Museum
by Tomlin in 1937. The shell is missing, but it must be presumed that one was
originally associated with these soft parts, for the latter could not otherwise have
been identified ; it may have been one of the two exhibited by Tomlin in 1937 at a
meeting of the Malacological Society of London. (Tomlin's collection, lodged in the
National Museum of Wales, Cardiff, includes two uncatalogued shells of Thatcheria
mirabilis from Kii, and it seems likely that these are the same two.) Dr. Eales has no
information on the subject.
(2) The shell of which an X-ray photograph was given in Eales' (1938) paper.
This photograph was of " a specimen in Mr. Winckworth's collection ". Winck-
worth's entire collection was later bought by the British Museum ; the shell shown
in the photograph may therefore be in the Museum's possession, but it cannot be
found at present.
(3) Another shell (B.M. regd. no. 1937-7-9.40). This specimen is considerably
smaller than the holotype (length 76 mm. as against 87 mm.). It too was obtained
at Kii, Japan, and was bought from H. C. Fulton in 1937. It is certainly not the
X-rayed specimen.
Supplementary description
Angas (1877) described Thatcheria mirabilis as having an " outer lip with a broad
excavated sinus extending from its juncture with the body-whorl to the extremity of
the last keel " ; he gave no more precise indication of its form. His fig. la of the
shell in apical view shows the margin of the posterior sinus (and all the collabral
growth-lines parallel thereto) intersecting the edge of the preceding whorl more or
less at right angles ; as the margin passes outwards from that point it at once
curves smoothly forward and then runs obliquely to meet the carina at an acute angle.
His fig. i b of the same shell in apertural view gives no more information on this
matter.
270 GASTROPOD GENUS THATCHERIA AND ITS RELATIONSHIPS
It is nevertheless obvious that, since each whorl is much wider at the carina than
at the abapical suture, it will hide the inner part of the ramp of the following whorl
when viewed from the direction of the apex ; thus, in Angas' figure, the line which
seems to terminate the margin of the sinus at its inner end is the carina of the pre-
ceding whorl. The margin of the sinus must in fact pass further adaxially beneath
the carina to meet the (hidden) adapical suture. But what is the precise form of
this inner part of the sinus margin?
Except for one drawing by Beets (19436, referred to below), the work of subsequent
authors has done nothing to make known the form of the posterior sinus in Thatcheria
mirabilis ; sometimes, indeed, the form of the sinus has been completely misunder-
stood. The holotype will therefore be briefly re-described, with particular reference
to that feature.
The form of the sinus is clearly shown in Plate 47, fig. 3. The margin of the sinus
does not pass perpendicularly outwards from its point of origin on the adapical suture,
but extends obliquely outwards and backwards for a short distance ; as it does so it
curves right round until it is running obliquely forwards to meet the carina at an acute
angle. The outer limb of this arch is much longer than the inner, so that any given
growth-line meets the carina at a point which is much further forward than its point
of origin, and the vertex or base of the sinus lies much nearer to the suture than to
the carina — at about a quarter of the distance across the ramp. This type of sinus
may be described as deep and juxtasutural, its depth being largely due to the for-
ward extension of the outer lip.
Powell (1942 : 167) was entirely misled by Angas' figure. He wrote that " That-
cheria has ... a very deep sutural sinus " ; but, as mentioned above, he noted
that Thatcheria seemed to differ from Waitara in that its sinus swung forward
immediately from its point of origin.
Beets, on the other hand, seemed to understand the true nature of this sinus. In
his drawing of T. mirabilis in apical view (19436, pi. 36, fig. 2) the position of the
suture, hidden by the projecting carina of the preceding whorl, is marked with a
broken line ; also marked are the whole of the margin of the sinus and of one growth-
line, rightly showing the hidden parts next to the suture passing backwards in an
arc. However, he paradoxically wrote (p. 361) : " Der Sinus der Aussenlippe ist
vollkommen vergleichbar jenem der (anderen) C/wwa-Arten ". Now this is true.
But, as shown below, his conception of the form of the posterior sinus in Clinura
was itself completely wrong ; it was based on the misleading illustrations of other
authors, so that he believed the sinus to be quite shallow. Further, he cited (Beets,
1951 : 1 6) Powell's observations on the Thatcheria sinus, yet at the same time was
puzzled by the apparent fact that the sinuses of Cochlioconus and Waitara, while
similar to each other, differed from that of Thatcheria in that they were first narrowly
concave.
The superficial ornament of the shell is another feature of T. mirabilis that has
not been adequately described. Angas (1877) wrote of his holotype : " above the
keels finely arcuately striate [i.e. with collabral growth-lines parallel to the margin
of the posterior sinus], below irregularly more or less crenately concentrically ridged ".
Beets (19436 : 361) included this description of the superficial ornament of T.
GASTROPOD GENUS THATCHERIA AND ITS RELATIONSHIPS 271
mirabilis : " der Kiel 1st glatt, auch der obere Windungsabschnitt ; vor der Kante
sind die Umgange wieder fein spiralig gefurcht."
There are, in fact, three distinct categories of superficial ornament. These are
spiral ornament, running in the direction of growth of the helicocone ; transverse
ornament, running across (more or less) the direction of growth of the helicocone ;
and collabral ornament (growth-lines), parallel to the outer lip throughout the length
of the latter. Transverse ornament, of course, is usually collabral, but is not
necessarily so (Cox, 1955 : 198).
In Thatcheria mirabilis there are indeed many distinct spiral furrows on the outer
face, often arranged in pairs (PL 47, fig. i). On the lower part of the outer face of
the last whorl, within the inner lip, the furrows are soon obliterated by the forward
growth of the edge of the mantle ; thus each spiral furrow, if traced backwards and
inwards for a few millimetres beyond the inner lip, disappears beneath a shining
white layer of callus. The carina of the shell is smooth. Very weak spiral striations
are present on the ramp, except on its innermost part next to the adapical suture.
Transverse ornament is entirely lacking in T. mirabilis.
As for collabral lines, the form of the outer lip which produces them is simple and
slightly convex between the anterior canal and the carina. On the ramp the outer
lip delimits the posterior sinus, of which the form has already been described.
A final point to mention is that consideration of the length of all the nine measured
shells of T. mirabilis recorded in the literature and in this present work leads to the
conclusion that the holotype is an adult shell of average size. The measured lengths
are 18 mm., 33 mm., 76 mm., 77 mm., 87 mm. (holotype), 89 mm., 94 mm., 97 mm.
and 98-5 mm. Three other shells have been mentioned but without measurements ;
they are the third specimen of Kuroda & Habe (1954), which is presumed not to be
the same as any of the three referred to by Habe (1955), and Tomlin's two shells
in the National Museum of Wales. Thus, in all, twelve shells have been mentioned
hitherto. There is at least one more shell in a private collection in England, there
are probably several in the United States, and it is likely that there are further
specimens in Japan and elsewhere. Indeed, Thatcheria mirabilis, once thought to
be a unique monstrosity, is no longer even a great rarity.
IV. NOTES ON CLINURA
Material studied
The British Museum collections contain two well preserved specimens of Clinura ;
one of these belongs to the type-species C. calliope. The determinations were verified
by detailed comparisons with the descriptions and figures of the original authors.
Particulars of the specimens are as follows :
(1) Clinura calliope (Brocchi 1814) [Murex]. Brit. Mus. (Nat. Hist.) regd. no.
G. 79439-
From the Plaisancian (Lower Pliocene) of Biot, near Antibes, Alpes Maritimes,
France.
(2) Clinura trochlearis (M. Homes 1854) [Pleurotoma]. Brit. Mus. (Nat. Hist.)
regd. no. GG. 2185. (This specimen was formerly in the Wrigley Collection.)
272 GASTROPOD GENUS THATCHERIA AND ITS RELATIONSHIPS
From the Zweite Mediterran-Stufe (Middle or Upper Miocene) of Walbersdorf,
Burgenland, Austria.
This material was studied in order to ascertain the generic characters of Clinura
rather than the specific characters of C. calliope or C. trochlearis. Since the former
is the type-species, the latter was considered only in so far as it afforded additional
evidence of those generic characters.
Posterior sinus
It is evident from Plate 47 that the posterior sinus of Clinura calliope (fig. n) was
exactly like that of Thatcheria mirabilis (fig. 3). But, just as in T. mirabilis, the
form of the posterior sinus is not so readily apparent from the descriptions and has
often been misunderstood — though in an entirely different way. In these fossil
suture
shallow sinus $
outer lip
drawn forwards
carina
outer face
FIGS, i and 2. — Alternative ways of describing the posterior sinus of
Clinura and Thatcheria.
gastropods the outer lip with its sinus is usually broken away, and the form of the
sinus must therefore be inferred from the form of the collabral growth-lines. Some
of the earlier figures (and hence later ones copied from them) either did not show the
growth-lines or else depicted them inaccurately ; and it is unfortunate that later
workers have sometimes ignored descriptions in foreign languages. Finally, the
posterior sinus of Clinura has been variously described as " deep " or " shallow ",
even by the same author ; this point requires explanation.
Confusion has probably arisen over the depth of the posterior sinus for the following
reason. Some workers (including the present writer) define the posterior sinus as
including the whole of the notch between the outer lip and the suture (Text-fig, i).
Others, however, have considered it to end anteriorly (with respect to the direction
of growth of the helicocone) at the level of the junction of its margin with the suture
(Text-fig. 2) ; in which case the sinus of Clinura must be regarded as shallow, but it is
then necessary to mention that there is also a well developed forward extension of
the outer lip.
The original author (Brocchi, 1814, pi. 9) gave two figures of Murex calliope which
GASTROPOD GENUS THATCHERIA AND ITS RELATIONSHIPS 273
were repeated in the later edition of his work (1843). His fig. 150, in apertural view,
shows nothing of the growth-lines on the ramp ; but fig. 156, in abapertural view,
shows them fairly well. All that Brocchi wrote in this connexion (p. 436) was
" ... si osservano alcune finissime rugosita obblique ed arcuate che sono le
tracce dell'antico margine di quella sinuosita, che hanno tutte le pleurotome nel
labbro destro, e ch'e il carattere del genere."
Bellardi (1847, P^ T> fi&- 9) depicted Pleurotoma calliope with strongly curved
growth-lines in which both limbs of the arch appeared to be of about the same length ;
that is, the sinus would have been deep but symmetrical.
The same author (1875 : 20), proposing the new genus Clinura, gave only a generic
diagnosis in Latin. This included two phrases which he later reproduced more
grammatically (Bellardi, 1877 : 204) as " Sinus posticus labri sinistri valde profundus,
arcuatus ; labrum sinistrum antice valde productum, aliforme ". But he also
wrote on p. 204 : "I caratteri principal! di questo genere sono [inter alia] :
3° labbro sinistro sinuoso posteriormente ;
4° seno molto largo e poco profondo, foggiato ad arco, il quale parte direttamente
dalla sutura posteriore ;
5° labbro sinistro protratto a guisa di ala ".
It is not easy to reconcile " sinus . . . valde profundus " with " seno . . . poco
profondo"! Incidentally, it should be noted that the outer lip is the "labbro
destro " of Brocchi but the " labbro sinistro " of Bellardi. In Bellardi's figure
(1877, pi. 7, fig. i) the little that can be seen of the growth-lines makes the sinus
appear shallow.
Cossmann (1896 : 74) described the sinus of Clinura as " largement arrondi en
quart de cercle incompletement ferine et aboutissant presque perpendiculairement
a la suture ". His pi. 5, fig. 19 shows nothing whatever of the growth-lines.
It is evident that Grant & Gale (1931 : 494) based their idea of the form of the
sinus in Clinura upon these works of Bellardi (1877) and Cossmann (1896), for they
referred specifically to the figures of C. calliope therein. Thus they were led to
write of the genus " posterior notch very wide and shallow, rounded ". They
also transferred to Clinura certain American species which, in fact, differ widely
from Clinura in the form of their sinus.
Of modern authors, only Wrigley (1939 : 283) seems to have been aware of the
true form of the posterior sinus in Clinura ; for he wrote of that genus, " the growth
lines sweep back over the rear slope four or five times as far as those of Surculites " .
This character was not illustrated in his paper. Wrigley was wrong, however, in
stating that Nekewis io (Gabb), from the Eocene of California, was much like
Clinura ; for Stewart's figure of that species (1927, pi. 30, fig. n) shows a fairly
shallow, symmetrical sinus.
Beets (19436 : 359) cited the figures of Brocchi, Bellardi and Cossmann mentioned
above, and then wrote : " Wie die Aussenlippe vor der Kante verlauft, ist nicht aus
Bellardi's Abbildung abzulesen, wohl aber aus den anderen zitierten Figuren."
His semi-diagrammatical drawing (19436, pi. 36, fig. 8, pi. 37, fig. 23 ; 1951, pi. i,
fig. 3) is nevertheless based upon Bellardi's figure, to which has been added a growth-
274 GASTROPOD GENUS THATCHERIA AND ITS RELATIONSHIPS
line evidently consequent upon a very shallow sinus. Thus Beets too had been
misled.
Rossi Ronchetti (1955 : 306) was not informative on this point. She wrote of
" Pleurotomella " calliope that " la maggior parte della superficie [della porzione
posteriore ... e] percorsa solo da strie di accrescimento alquanto arcuate."
Her fig. 163 shows but the faintest trace of growth-lines, and then only the part
nearest to the adapical suture.
As for Clinura trochlearis , examination of the material in the British Museum shows
that its posterior sinus is just like that of C. calliope. Indeed, the form of the sinus
in C. trochlearis was depicted very clearly by M. Homes in the figures to his original
description (1854 : 363, pi. 39, figs. 140, b, 150, b) [Pleurotoma] ; it has never been
subject to misunderstanding.
Protoconch
No protoconch has yet been described in any species of Clinura ; it may be that
the apex of the shell was missing in all the material available to those working on
the genus. But the two specimens in the British Museum, of which details have
been given above, both possess a protoconch.
The protoconch of C. calliope (PI. 47, fig. 12 ; Text-fig. 3) is well preserved, though
probably incomplete above ; two whorls remain. It is entirely covered with very
distinct oblique cancellation, each thread being slightly concave towards the anterior
side.
The protoconch of C. trochlearis (Text-fig. 4), of which two and a half whorls
are preserved, is rather abraded. The upper and lower parts of each whorl are
obliquely cancellate, just like the protoconch of the type-species, but the central,
thickest part is worn smooth in places.
Transverse sculpture
The presence of transverse sculpture is well known in Clinura calliope. It consists
of prominent tubercles on the carina ; according to Rossi Ronchetti there are
eighteen per whorl, but in fact the number increases with the size of the helicocone
from not more than ten to at least twenty. Further, on the early whorls of the
teleoconch each tubercle is continued abapically as a short, broad, slightly opistho-
cline rib running down the outer face (see PI. 47, fig. 12).
There is no record of any comparable sculpture in C. trochlearis ; indeed, Homes
(1854 : 364) described his new species as having " nicht eine Spur von Spitzen oder
Dornen an dem Kiele, selbst an den obersten Windungen ". This is certainly true
of all the larger whorls. But the British Museum specimen shows that, just as in
C. calliope, the first two and a half whorls of the teleoconch bear small nodules on
the carina, about twelve on the first whorl and sixteen on the next ; each has a
small rib beneath it. As the helicocone grows these successively diminish in relative
size and eventually disappear.
GASTROPOD GENUS THATCHERIA AND ITS RELATIONSHIPS
5 67
FIGS. 3-7. — Protoconchs :
3. Clinura calliope (Brocchi) ; original drawing by D. L. F. Sealy, based on a
photograph of specimen no. G. 79439 in the British Museum (Natural History).
4. Clinura trochlearis (M. Homes) ; original drawing by D. L. F. Sealy, based on a
photograph of specimen no. GG. 2185 in the British Museum (Natural History).
5. Waitara liratula Powell ; after Powell.
6. Thatcheria mirabilis Angas ; after Kuroda & Habe.
7. Cryptodaphne pseudodrillia Powell ; after Powell.
V. NOTES ON WAITARA LIRATULA POWELL
Protoconch
Powell (1942) was the first to postulate a close affinity between Waitara and
Thatcheria. He mentioned that comparative studies of the protoconchs had yet to
be made ; for, in any of the genera with which this work is concerned, the only
protoconch known at the time was that of W. liratula described by Powell himself.
Since then the protoconch of Thatcheria mirabilis has been described and figured by
Kuroda & Habe (1954) from a beautiful specimen showing very distinct oblique
reticulate ornament, and the remarkably similar protoconchs of two species of
Clinura have now been described in the preceding section of the present work. If
Clinura, Waitara and Thatcheria are indeed closely akin, it would be expected that
the protoconchs also would be alike ; this expectation is rendered even more prob-
able by the Clinura-like tuberculation of the carina on the first few whorls of the
W. liratula teleoconch, noted below. But, according to Powell (1942 : 169), W.
liratula has a " dome-shaped protoconch of two whorls bearing weak protractively
arcuate axial growth threads " ; and his drawing (text-fig. 63 on p. 38 ; Text-fig.
5 in the present work) shows no ornament other than faint, slightly sinuate, slightly
opisthocline striations, with no indication of others running across them. Might not
this appearance have been caused by abrasion?
276 GASTROPOD GENUS THATCHERIA AND ITS RELATIONSHIPS
It was felt that the protoconch of Waitara liratula should be looked at again.
According to Fleming (in litt.), " The paratype figured by Powell (pi. 14, fig. 8) is
apparently the same specimen on which he based his text-fig. 63 ... The
holotype and other paratypes are still more deficient in protoconch. . . . We
have no evidence whether the protoconch was damaged after Powell's figure was
made." The figured paratype (PI. 47, fig. 7) was therefore sent to London and
examined with great care. Unfortunately the specimen does seem to have been
damaged since Powell described it twenty years ago, for, whatever its condition
before, the protoconch is now in such a poor state of preservation that it is difficult
to compare it with Powell's drawing and impossible to see any traces of its original
ornament. Indeed, the protoconch is far too worn and damaged to afford positive
evidence of any significant difference between Waitara and Thatcheria in this taxono-
mically important character. A close relationship between Waitara and Thatcheria,
as originally suggested by Powell himself, is therefore still quite possible.
Transverse sculpture
During the examination of the figured paratype of Waitara liratula it was noticed
that the first two or three whorls of the teleoconch bore unmistakable indications
of nodules on the carina, arranged with an appearance of regularity. The angular
distance between consecutive nodules was estimated at about 20° of arc ; thus,
originally, there would have been about eighteen nodules per whorl. Beneath some
of these there were traces of a short, broad and markedly opisthocline rib running
down the outer face of the whorl. After the first few whorls, however, the carina
became smooth. This character has not previously been recorded in W. liratula.
The Neogene shells from Okinawa which MacNeil (1960 ; see above, p. 268)
described as Thatcheria cf . gradata seem to have a close affinity with W. liratula, even
though it must be admitted that comparisons are difficult when they have to be
made from one illustration of a single incomplete specimen of each species. The
early whorls of the teleoconch of the figured specimen from Okinawa are far better
preserved than in Powell's specimen and bear distinct nodules on the carina, exactly
like those of Clinura calliope ; these gradually become less distinct and eventually
disappear at a point where the transverse diameter of the shell (measured across the
carina) is about 4 mm.
Marwick (1931 : 149) reported that the early whorls of W. gtnerosa bore about
twelve low, rounded axial costae extending downwards from the keel and forming
weak tubercles at their intersection with the shoulder-angle.
VI. SYSTEMATICS
General principles
The classification of the gastropods discussed in this work, except that of the
Recent Thatcheria mirabilis, is necessarily based entirely upon their shell characters.
It has been generally believed that the most fundamental of these are the position
and form of the posterior sinus, indicated in incomplete specimens by the position
GASTROPOD GENUS THATCHERIA AND ITS RELATIONSHIPS 277
and form of the collabral growth-lines ; this feature is considered to be the most
useful of those by which a turrid may be distinguished from other Toxoglossa and
assigned to a sub-family within the Turridae. But the posterior sinus is not in-
fallibly diagnostic in these matters. In the Conidae, which are without doubt
close relatives of the Turridae, the form of the posterior sinus varies not only from
species to species but even within a species, and it is sometimes just like that of
certain Turridae. For example, the collections of the British Museum (Natural
History) include a series of shells of the Recent Conus pennaceus Born 1780, all
characterised by a particular colour pattern. One of these, however, has a posterior
sinus which resembles that figured by Powell (1942, text-fig. F28) as belonging
to the Recent turrid Asperdaphne versivestita (Hedley 1912) [Daphnella} ; another
has the " reversed L " sinus more typical of the Daphnellinae (as found in Daphnella
cancellata Hutton 1878) ; in a third specimen the sinus is similar to that of Clinura
or Thatcheria ; and in a fourth it is virtually absent. In other individuals it is
yet again quite different from any of these. Thus there can be no such thing as a
sinus typical of the Conidae, although Powell (1942 : 167), who mentioned the " deep
Conid-like " sinus of his Thatcheriidae, clearly implied that there was. Again,
Powell (p. 28) wrote of the Turridae that " Position and style of sinus is the best
guide to subfamily grouping." Experience may indeed have shown that there is no
better, but the guidance afforded by the posterior sinus provides no simple key to
this problem ; its form, if not its position, may vary widely within a given sub-
family. In short, the posterior sinus appears to be of systematic value only when
the taxonomic placing indicated thereby is confirmed by other distinctive characters
with which it has no connexion.
It was therefore necessary to bear other characters in mind when considering
whether or not the molluscs in question should be referred to the Turridae and, if so,
to which particular sub-family thereof. Those characters were the nature of the
protoconch and the absence of a tendency towards the resorption of the columella
and early internal walls. Further, when a series of shells was thus referred to a
given grouping, their possession of a common suite of less distinctive characters
(such as a pagodiform spire, a certain distribution of spiral ornament, and a certain
type of columella) offered additional confirmation of their relationship.
Generic and specific separation were effected primarily upon the presence or
absence of transverse sculpture and tubercles, and secondarily upon a number of
other characters — the position of the carina on the whorl, the degree of develop-
ment and precise nature of the spiral ornament, the degree of " pagodiformity ",
the spire angle, and the height of the spire relative to that of the last whorl and
aperture.
In living Toxoglossa the soft anatomy (where known) affords more evidence of
the animal's affinities. The morphology of the radula is important in this connexion.
The presence or absence of an operculum, and, when present, its structure, are also
characters which are generally of taxonomic value only in Recent forms. Fossil
gastropods are rarely associated with opercula, even where they are known to have
possessed them, and the seeming absence of an operculum in a little-known form is
thus without significance.
278 GASTROPOD GENUS THATCHERIA AND ITS RELATIONSHIPS
The Clinura-Thatcheria group
As mentioned above (p. 262), it was suggested by Beets (19436) that Clinuropsis,
Clinura and Thatcheria formed a closely related group. He regarded Cochlioconus
as part of Thatcheria, and did not consider Waitara at all. But Powell (1942) had
already recognised a relationship between Waitara and Thatcheria ; and Wenz
(1943) placed Waitara (at that time known to him only by its Miocene species)
near Clinura, although he thought that Thatcheria and Cochlioconus had no affinities
with either, or even with each other. Beets, therefore, in a later paper (1951), added
Waitara to his group.
This suggested grouping has evoked no mention or comment elsewhere. The
forms in question certainly seemed to possess some characters in common and no
fundamental differences ; in particular, the shell was always more or less pagodiform,
with a juxtasutural posterior sinus (admittedly of variable depth) having its vertex
close to the suture. All the species seemed to be extremely rare, and were probably
abyssal in habit.
More conclusive evidence of this supposed relationship is now available, although
the extent of the group requires limitation. Beets' conception of the taxon Clinura
was a very wide one, even in the more restricted sense in which he used it in 1951.
His misunderstanding of the true form of the posterior sinus in the type-species,
C. calliope (Brocchi), had led him to believe that the depth of the sinus, since it
appeared to vary greatly in species which were otherwise similar, was subject to
" normal " variation within the group ; yet, in the same work (19436 : 364), he stated
that the form of the sinus was " ein Merkmal von hochster Bedeutung in der Sys-
tematik der Turridae". It has now been made quite clear (see p. 272] that the
posterior sinus of the type-species is very deep ; and it is also apparent (see PI. 47,
figs. 3, 6, 8, n) that its exact shape as found in C. calliope is a constant character of
Thatcheria (including Cochlioconus) , of Waitara, and of certain other species referred
to Clinura itself, but not of the remaining species hitherto referred to Clinura and
not of Clinuropsis. It is therefore suggested, in accord with Powell and Beets, that
the highly distinctive type of sinus found so consistently in these genera is indeed
of taxonomic value, notwithstanding what has been written above concerning the
variable position and form of the posterior sinus in the Toxoglossa in general. (This,
of course, would not be true if a sinus like that of Thatcheria could be clearly shown
to have developed as a result of parallel evolution in an entirely different group.)
It is further suggested that any alleged species of Clinura without this deep sinus
have been wrongly placed in that genus. An examination of the original descrip-
tions and figures of most of the species mentioned by Cossmann (1896 : 75) ; Grant
& Gale (1931 : 494) ; Beets (19430 : 296, 297 ; 19436 : 358-64) ; Eames (1957 : 51)
and Glibert (1960 : 25, 26) indicates that this exclusion from Clinura should apply to
all the species from North America (Eocene to Miocene), Africa (Palaeocene and
Eocene), and the Palaeocene and Eocene of Europe, and to most of those from the
Neogene of Europe ; thus it is not possible to agree with Beets that the genera
Clinuropsis Vincent 1913 (non Thiele 1929) and Nekewis Stewart 1927 should be
placed in the synonymy of Clinura. Indeed, the only true species of Clinura (using
GASTROPOD GENUS THATCHERIA AND ITS RELATIONSHIPS 279
the name in Beets' narrower sense, thus also excluding Waitara, Cochlioconus and
Thatcheria) seem to be the type-species and a few others from the Neogene of Europe,
together with the two species from the Neogene of Buton (Celebes).
It is noteworthy that the only species (other than C. calliope) mentioned by
Bellardi when proposing the genus Clinura was C. elegantissima (Foresti 1868)
[Pleurotoma}, which, by the form of its growth-lines, is clearly not a Clinura. Its
complex spiral ornament is also very distinctive. On the other hand, a good example
of a European species accepted here as a Clinura is C. trochlearis (M. Homes 1854)
[Pleurotoma]. In the present work, the decision to retain this species within the
genus Clinura was originally based upon Homes' description and figures, with
particular reference to the form of the posterior sinus ; only later did this decision
receive powerful support from the discovery that an individual of C. trochlearis in
the collections of the British Museum (see p. 274) showed a diagonally cancellate
protoconch, just as in C. calliope, and transverse, slightly opisthocline costae on the
juvenile whorls of the teleoconch. Another species which may be assigned to
Clinura with a high degree of probability is C. controversa (Bellardi 1847) [Pleuro-
toma] ; Bellardi, incidentally, ascribed this new specific name to Jan 1845 " in
litt. et specim." (p. 64).
More doubtful cases, where it would probably be difficult to make a definite
decision on the correct systematic position of the species without handling the actual
type-material, include C. sopronensis (Wolf 1870) [Pleurotoma], C. subtrochlearis
(Friedberg 1912) [Surcula], and C. sabatiorum Bellardi 1878. But it is felt that a
full examination of every species attributed to Clinura by one author or another,
often without justification, is beyond the scope of the present review. The species
excluded would form such a large and varied assemblage, with such a wide geo-
graphical distribution and such a long stratigraphical range, that they would doubt-
less represent several distinct genera. The available genera to which they might
be referred would include those listed by Grant & Gale (1931) and by Beets (19436)
as synonyms of Clinura, other than Thatcheria and Cochlioconus : namely, Clinuropsis
Vincent 1913 (non Thiele 1929) and Nekewis Stewart 1927. It is possible, how-
ever, that not all species might be referred to those or to other existing genera, in
which case new genera would be needed.
The Butonese species of Clinura are C. carinata (Martin 1933) [Cryptoconus] and
C. bituminata (Beets 19430) [Surculites].
If the genus Clinura be limited as suggested above, which to me seems perfectly
reasonable, then the reasons for postulating a close relationship with Waitara and
Thatcheria (including Cochlioconus) are much more convincing. But there are no
grounds for supposing these forms to be related just as closely to the genus
Clinuropsis.
Confirmation of this affinity is given by the protoconchs. Until a few years ago
that of Waitara was the only one described (Powell, 1942). Now, however, the
protoconchs of the type-species of Clinura and Thatcheria have been made known
(see p. 274 above; and Kuroda & Habe, 1954) ; both show the same type of diagonal
reticulation, and this suggests a relationship between those two genera. On the
other hand, the Waitara protoconch described by Powell gives no indication of
280 GASTROPOD GENUS THATCHERIA AND ITS RELATIONSHIPS
diagonal cancellation ; but, as stated above, its condition is so unsatisfactory that
the nature of its original ornament cannot be ascertained. At least it affords no
evidence to refute the suggestion, based on other conchological evidence, that
Waitara is related to Clinura and Thatcheria. In any case, the species in question
(Waitara liratula} is not the type-species of Waitara.
Other common characters of the shells, which might be of little taxonomic value
on their own, together reinforce the evidence already cited for the relationship of the
three genera. They include the pagodiform habit, the distribution of the spiral
ornament on the outer surface of the teleoconch, and the smooth, very slightly
twisted nature of the columella. The distribution of the spiral ornament cannot
always be ascertained very easily from the published descriptions and illustrations ;
but the ornament is typically confined to the outer face of each whorl, except in
that there are comparatively weak striations on the outer edge of the ramp (next to
and parallel with the carina). This type of distribution is certainly found in Clinura
calliope, C. trochlearis, C. bituminata, Waitara pagodula, W. liratula, and in the new
species of Thatcheria from Fiji described below. In the Recent Thatcheria mira-
bilis, of which the material is of course in a far better state of preservation, the
spiral ornament on the ramp extends further towards the adapical suture, but it
is altogether fainter than in the other species and is in no way comparable with the
much stronger striation of the outer face.
The value of Beets' eventual conclusion, that Clinuropsis, Clinura, Waitara and
Thatcheria (including Cochlioconus) form a closely related group, may therefore be
assessed as follows. His suggestion was based at the time on inadequate evidence ;
but a more restricted conception of the genus Clinura, coupled with better under-
standing of the form of the posterior sinus in that genus and with new knowledge
of the form of the protoconchs in Clinura and Thatcheria, now indicates that it is
perfectly reasonable in so far as Clinura, Waitara and Thatcheria are concerned.
Clinuropsis, however, does not form part of this group. Further, it cannot be
agreed that the similarities between the three named taxa in the group are sufficiently
close to justify placing them all in synonymy (senior synonym Clinura, ranked as
a sub-genus of Surculites in Beets, 19436), or even to warrant their being considered
as three separate sub-genera of Surculites (in Beets, 1951). In the following argu-
ment they are regarded initially as three distinct genera. In any case, close affinity
with Surculites is denied in the present work ; the matter is discussed more fully
below (p. 289), where the external relationships of the Clinura-Thatcheria group are
considered in detail.
At present, then, this group appears to include at least the following species :
Clinura calliope Waitara waitaraensis Thatcheria mirabilis
C. trochlearis W. generosa T. gradata
C. controversa W. pagodula Thatcheria sp. nov. Beets
C. bituminata Waitara sp. nov. Powell I95I
C. carinata J942 aff. pagodula
W. liratula
GASTROPOD GENUS THATCHERIA AND ITS RELATIONSHIPS 281
Division into genera
The species referred to Waitara seem, in general, to be intermediate in character
between Clinura and Thatcheria. This division of the group into three genera,
however, appears rather unsatisfactory : like species are separated (C. carinata and
W. pagodula, W. liratula and T. mirabilis} and unlike species placed together (C.
bituminata and C. carinata, W. generosa and W. liratula). A careful consideration
of all the species involved has led to the following conclusions.
All three species listed as Thatcheria are correctly regarded as congeneric with each
other. But several other species listed above (Clinura carinata, Waitara pagodula,
Waitara sp. nov. Powell 1942 aff. pagodula, W. liratula} and the new species from
Fiji described below as Thatcheria vitiensis are all very similar to the three species of
Thatcheria listed ; they resemble them in having the carina generally high on the
larger whorls and only weak spiral ornament, and in being entirely free of transverse
ribbing or tubercles at every stage of development (except, in three cases, for minute
vestigial nodules on the very youngest whorls of the teleoconch), while they differ
from them chiefly in being less pagodiform. Indeed, there is little more than the
variable degree of pagodiformity to distinguish any of these eight species from each
other. It is therefore suggested that they should all be placed in the genus
Thatcheria.
The geographical distribution of the genus as thus enlarged remains entirely
within the area of the Western Pacific. Its stratigraphical range extends from Upper
Miocene to Recent, but it is predominantly Pliocene.
Powell (1942), of course, recognised the affinity of Waitara and Thatcheria (see
pp. 260,261), " probably basing his opinion mainly on the features of two species of
Waitara newly described by him " (Beets, 1951 : 16). But, because he had mis-
interpreted the form of the posterior sinus of T. mirabilis from the growth-lines shown
in Angas' rather misleading figure, he doubtfully retained Waitara as a separate
genus " for New Zealand usage ". Beets (1951 : 16) pointed out that the sinus in
T. gradata [Cochlioconus] " is almost similar to that in Waitara ", but he failed to
realise Powell's mistake in the matter of the T. mirabilis sinus, even though he him-
self had earlier figured it correctly (19436, pi. 36, fig. 2). However, this misinter-
pretation has now been explained (pp. 269, 270) ; and Powell's younger species of
Waitara (though not Marwick's two older species, one of which is the type-species) have
been referred to Thatcheria.
On the other hand, if it should later transpire that the protoconch of T. liratula
and its allies (known in only the one specimen of T. liratula, which seems to have
been subsequently damaged) is, as appears from Powell's description and figure,
essentially different from that of Thatcheria, then it would be necessary to erect a
new genus for those forms.
If the younger species formerly ascribed to Waitara are transferred to Thatcheria,
then the only species remaining in the former genus are the type-species waitaraensis,
from the Upper Miocene of New Zealand, and generosa, from the Lower Miocene
of the same country. Marwick's fairly detailed generic diagnosis of Waitara ap-
pears to be based equally upon waitaraensis and generosa, so that, as mentioned above
(p. 261), Wenz (1943) was misled into believing that generosa was the type-species
282 GASTROPOD GENUS THATCHERIA AND ITS RELATIONSHIPS
of the genus. But, as far as can be judged from Marwick's descriptions of these two
species, only one individual is known of each. The holotype of waitaraensis in
particular is very incomplete and seems to have suffered distortion ; there are
no highly distinctive characters, either positive or negative. It therefore seems that
the genus Waitara, denned objectively by reference to this single specimen, cannot be
denned very clearly. It is nevertheless obvious from Marwick's descriptions and
figures that, unlike W. pagodula and W. liratula, neither W. waitaraensis nor W.
generosa can be referred to Thatcheria. Neither species is much less pagodiform than
is T. liratula or the new Fijian species T. vitiensis — indeed, they are probably more
pagodiform than is T. pagodula — but their whorls have an entirely different aspect.
On the other hand, Beets wrote (1951 : 16) that " there are also relationships
between Waitara in its former sense (i.e., as based upon W. generosa and W. waitara-
ensis) and Clinura " . This appears to be true at least of W. generosa, which has both
the characteristic attributes of a Clinura : the first few whorls of the teleoconch
are ribbed, with tubercles on the carina, and the carina is low on the larger whorls
(that is, the ramp slopes steeply downwards). These show its affinity with the
European species of Clinura (including the type-species, C. calliope] and with the
Butonese C. bituminata. It is therefore proposed that W. generosa be transferred
to the genus Clinura. As for W. waitaraensis, the incomplete and distorted nature
of the specimen prevents its giving any information on these characters ; but, since
Marwick (who handled the material) believed it to be congeneric with W. generosa,
it may also be transferred — albeit provisionally — to Clinura, thus making Waitara
a subjective junior synonym of Clinura. If better material later showed that this
transfer, though correct for W. generosa, was not justified in the case of W. waitara-
ensis, the generic name Waitara would still be available for the latter species.
The genus Clinura, as listed above, has now lost C. carinata to Thatcheria but
has gained the species waitaraensis and generosa from Waitara ; i.e., it includes the
species calliope, trochlearis, controversa, bituminata, generosa, perhaps waitaraensis,
and possibly others from Europe. The geographical distribution of the genus in
this new sense is thus restricted to Europe (the Mediterranean Basin), the East
Indies and New Zealand. The European species are all of Miocene age, although
some (such as C. calliope itself) persist into the Lower Pliocene (Plaisancian) ; the
Butonese species is of Neogene age, very probably Upper Miocene ; and the two
New Zealand species, as mentioned above, are from the Lower and Upper Miocene
respectively. It may therefore be said that the stratigraphical range of Clinura is
from the Lower Miocene to the Lower Pliocene, but that the genus occurs mainly
in the Middle and Upper Miocene.
The stratigraphical range of Clinura contrasts with that of Thatcheria, essentially
Pliocene to Recent. This shows that the most natural break in the Clinura-
Thatcheria group occurs between the earlier and the later species of " Waitara " at
about the level of the Miocene-Pliocene boundary. It seems, however, that there
is a certain amount of overlap on either side of that boundary. In one direction
C. calliope survives into the Plaisancian, as do other possible species of the genus
(see Gilbert, 1960 : 25, 26) ; in the other, rocks which are probably no younger than
GASTROPOD GENUS THATCHERIA AND ITS RELATIONSHIPS 283
Upper Miocene already contain T. carinata, and T. pagodula may likewise be of
Upper Miocene age.
The genus Clinura, even in the restricted sense used here, cannot be denned very
easily ; it is essentially a rather variable assemblage of species, of wide distribution
and of predominantly Miocene age. Thatcheria, on the other hand, even in the
present broad interpretation, is a fairly homogeneous group of species and may be
denned quite clearly. It seems that it originated from Clinura towards the end of
the Miocene, became widespread in the Western Pacific area during the Pliocene,
and survives to the present day as the one rather " extreme " form living in deep
waters off the Japanese coast.
T. mirabilis may in fact occur more widely than is known at present, " off Japan " ;
for a rare abyssal form is far more likely to be recovered in the Kii region, off the south
coasts of Honshu and Shikoku, than anywhere else in the Pacific. Not only is the
Pacific Ocean almost at its deepest off Japan, but in few places are there such deep
waters so near to a large area of land. Indeed, they are nowhere else so close to a
heavily populated shore-line from which intensive fishing is carried out.
Interspecific differentiation in Thatcheria
The eight species referred in this work to the genus Thatcheria seem to differ from
each other in little more than in certain characters of the spire : in its degree of
pagodiformity, in its acuteness, and in its size relative to that of the last whorl
and the aperture. In general, the various species seem to form an evolutionary
series in which the degree of pagodiformity increases with time ; and, as it increases,
the spire appears less acute.
It is very difficult, if not impossible, to define " pagodiformity " objectively.
Several numerical properties of pagodiform shells were examined as possible indices
but rejected for various reasons ; they are shown in Text-fig. 8. Measurement of the
spire angle proved more useful, for this, in a shell of the type under consideration,
can be measured in two different ways (Text-fig. 9). The term " internal spire
angle " may be introduced for the angle between the two lines which, on either side
of the spire, connect the intersections of the suture with the lateral profile. It is
obvious that only an approximate value can be obtained for this, for the lines are
unlikely to be straight ; it is not even likely that smooth continuous curves can be
drawn to pass exactly through all the points, and the best that can be obtained will
nearly always be a slightly concave curve passing close to most of them. The
" external spire angle " is measured in a similar manner but is based on the inter-
sections of the carina with the lateral profile ; it is, of course, larger than the internal
spire angle.
Of the eight species of Thatcheria in which such measurements were possible
(including T. vitiensis sp. nov. described below), seven gave a value for the internal
spire angle which was remarkably constant, lying always between 38° and 46° ;
exact measurements would not be especially significant, for, even in different indi-
viduals of one species (the Recent T. mirabilis), the angle varied between 38° and
44°. Only T. pagodula gave a significantly different value, 57°.
284 GASTROPOD GENUS THATCHERIA AND ITS RELATIONSHIPS
c
FIG. 8. — Plane projection of the spire of an idealised pagodiform shell, to illustrate the
numerical properties examined as possible indices of pagodiformity but rejected for
various reasons. They are :
(a) The projection of the carina, expressed as the ratio of its perpendicular distance
from the shell axis (AB) to the distance of that perpendicular from the apex (BC).
This is the tangent of half the external spire angle (e).
(b) The angle (r) between the profile of the ramp and the shell axis.
(c) The height of the carina on the whorl, expressed as the ratio of its axial distance
from the lower suture (YZ) to the axial distance between the sutures (XZ).
On the other hand, the external spire angle varied greatly, from 50° in the earliest
species T. carinata to 82° in the holotype of the Recent T. mirabilis. It seemed to
show a fairly steady increase with the passage of time ; but T. pagodula was once
again the exception, with an external spire angle of 71°. This higher value, however,
is consistent with the higher value for the internal spire angle. In fact, the ratio
of the external spire angle to the internal, which might be called the " index of
pagodiformity ", is the same in T. pagodula as in the much slimmer T. carinata
(the only other species of possibly Miocene age) — namely, 1.25.
To sum up, then, the situation seems to be that there are two species of Thatcheria
of possibly Miocene age. One of these (T. carinata} has an internal spire angle of
GASTROPOD GENUS THATCHERIA AND ITS RELATIONSHIPS 285
FIG. 9.
Note
-Plane projection of the spire of an idealised pagodiform shell, to illustrate :
i — the internal spire angle,
e — the external spire angle,
ds — the angle of declination of the suture, and
dc — the angle of declination of the carina.
(a) ds is greater than dc in a pagodiform shell, for the suture descends in a tighter
(and therefore steeper) spiral than does the carina.
(£>) Both dc and ds are smaller than they appear in the plane projection. This is
because, in the plane projection, both carina and suture appear to descend from one
side of the shell to the other in a straight line ; whereas, in fact, they do so on the
surface of a cone and therefore less steeply.
tan ds tan e/2
(c) — — should be equal to .
tan dc tan t/2
40°, the other (T. pagodula) has a much broader spire (internal spire angle 57°) ;
but both have exactly the same index of pagodiformity (1-25). All the later species
have an internal spire angle which is much the same as in T. carinata ; but the
index of pagodiformity, like the external spire angle, shows an increase with the
passage of time — to 1-50 in the two supposed Lower Pliocene species (T. liratula,
T, vitiensis), to 1-58-1-62 in the two species described simply as " Pliocene " (T.
286 GASTROPOD GENUS THATCHERIA AND ITS RELATIONSHIPS
gradata, Thatcheria sp. nov. Beets 1951), and to 175-2-05 in the Recent T. mirabilis.
It seems likely that the index of pagodiformity will prove to be more constant and
characteristic for a given species than either the internal or external spire angle ;
for the two spire angles, though they may vary considerably between individuals,
seem to do so together. Use of the index, though liable to considerable error, may
afford a rough indication of the course of evolution.
Of course, the position of a species on the evolutionary scale (and hence its approxi-
mate geological age) should not be assessed on any one character alone. The various
characters probably evolved at different rates in different lineages.
Evolution in the Clinura-Thatcheria group
Clinura calliope, type-species of the genus Clinura, ranged from Upper Miocene to
Lower Pliocene. Clinura, however, was already well diversified at the beginning of
that time ; thus the type-species occurred too late to occupy any central position
in the phylogeny of the genus as defined in the present work. Moreover, in one
prominent feature C. calliope differs from the other species which are considered here
to constitute that genus : the entire carina is furnished with well developed tubercles.
A better claim to a central position may be made on behalf of C. trochlearis (from
the Middle Miocene1 of Europe). Unlike every other species referred to Clinura,
C. trochlearis may be placed in that genus with absolute certainty. Except for the
poorly known, rather peculiar C. generosa in the Lower Miocene, it is probably the
oldest species of Clinura. It is somewhat variable in form, especially with regard
to the acuteness of the spire and the position of the carina on the whorl. And it is
also the most " central " in form ; for it could well be ancestral, by a late Miocene
radiation, to all the Upper Miocene species of Clinura and to Thatcheria.
Of those species in the Upper Miocene, C. waitaraensis is rather like C. bituminata
in the general aspect of its whorls, which, where undamaged, seem to have a steeply
inclined ramp and a low carina ; further, both those species are very similar to
C. trochlearis. C. bituminata and C. waitaraensis could be derived from a common
ancestor which, in turn, had evolved from C. trochlearis with little change in form.
Migration from Europe to the south-western Pacific would have occurred at the
onset of this hypothetical evolution.
Another species in the Upper Miocene of the south-western Pacific which also is
similar to Clinura trochlearis is Thatcheria carinata ; the flatter ramp and higher
carina distinguish it from C. bituminata and C. waitaraensis. Like those two species,
T. carinata may have been derived from a migrating descendant of Clinura
trochlearis. From Thatcheria carinata a line of evolution to the Recent T. mirabilis
may be envisaged ; it has been shown above that, with passage of time, the internal
spire angle remained more or less constant while the external spire angle (and hence
the index of pagodiformity) gradually increased. Only T. pagodula, with its
generally much stouter spire, would necessarily have formed a distinct Upper Mio-
cene offshoot from this series. It also seems that the evolution of Thatcheria from
1 Some authorities now prefer to regard the Zweite Mediterran-Stufe (Suess, 1866) of the Vienna Basin
as Tortonian in age, i.e. Upper Miocene, rather than Helvetian. But C. trochlearis occurs also in the
Middle Miocene of Italy (Bellardi, 1877 : 206).
GASTROPOD GENUS THATCHERIA AND ITS RELATIONSHIPS 287
FIG. 10. — A very tentative evolutionary tree of the Thatcheriinae.
the Miocene onwards has been accompanied by a northward movement of the popula-
tion towards its present home in the north-western Pacific.
The only other species in the Upper Miocene which have definitely been referred to
Clinura are both from the Mediterranean region, C. calliope (the type-species, per-
sisting into the Lower Pliocene) and C. controversa. A list of characters distinguish-
288 GASTROPOD GENUS THATCHERIA AND ITS RELATIONSHIPS
ing these two species (at that time referred to Pleurotoma) was drawn up by Bellardi
(1847 : 64) i in fact» it was largely by means of this comparison with P. calliope
that he described his new species P. controversa. Once again C. trochlearis may be
regarded as a possible ancestor — of C. calliope, C. controversa, or both, — but in this
case there is no need to postulate a distant migration of the stock. The tendency
towards the development of transverse sculpture on the first few whorls of the teleo-
conch, manifested in C. trochlearis and in other early species of the group, might have
led to the evolution of C. calliope by affecting the whole shell ; and, in similar fashion,
C. controversa might have evolved through the development of a notched carina.
As for Clinura generosa in the Lower Miocene, this species has a somewhat different
appearance ; it is difficult to imagine it as a direct ancestor of C. trochlearis and of
all the other species mentioned. But it has the typical posterior sinus of the
Clinura-Thatcheria group, together with the tubercles on the early whorls of the
teleoconch. It must therefore be retained within the group, in the genus Clinura,
and placed on a sterile side-branch of the evolutionary tree.
All these suggestions are summarised in Text-fig. 10.
Systematic position of the Clinura-Thatcheria group
Serious consideration of Thatcheria and its relatives has led most modern authors
to place them in the Turridae (Eales, 1938 ; Beets, 19436, 1951 ; Kuroda & Habe,
1954, implicitly ; Habe, 1955). There have been only two important exceptions.
Yokoyama (1928, 1930), though he did not explicitly refer Thatcheria [Cochlio-
conus] to the Conidae, thought it had affinities therewith ; while Powell (1942)
regarded Waitara, Cochlioconus and Thatcheria as constituting a new family, the
Thatcheriidae, distinct from the Turridae (though parallel to them) and more closely
related to the Conidae.
Preference is given here to the view that this group of gastropods is best placed
in the Turridae. The posterior sinuses of Clinura and Thatcheria may indeed be
compared with those of certain Conidae (e.g., Conus araneosus Solander 1768) ;
although, in general, the outer lip is drawn further forward in Clinura and Thatcheria
before it crosses the line of the carina. On the other hand, their posterior sinuses
also resemble those of certain turrids ; the diagonaUy cancellate ornament on the
protoconchs of Clinura and Thatcheria is a turrid character ; and neither genus is
known to show any manifestation of the tendency, typical of the Conidae, to resorb
the columella and early internal walls. Further, the form of the radula in Thatcheria
mirabilis and other characters of that Recent species confirm the opinion that the
group should be referred to the Turridae, particular significance being attached to
Eales' work upon the soft anatomy. There would seem to be little or no justification
for the erection of a separate family to accommodate Clinura and Thatcheria, even
if that family should be considered as having turrid rather than conid affinities.
Clinura, of course, had been placed in the Turridae (Pleurotomidae) from the
very date of its establishment in 1875, long before Eales (1938) wrote her paper on
Thatcheria and long before it was first suggested that the two genera were related.
A brief history of the more exact classification of Clinura has already been given above
(pp. 263-268) ; sufficient here to note that, of recent years (since 1931), the taxon has
GASTROPOD GENUS THATCHERIA AND ITS RELATIONSHIPS 289
been generally regarded as a sub-genus of Surculites. Grant & Gale wrote (1931 :
494) •'
" Clinura is closely related to Turricula, and might be considered a variational
extreme in the opposite direction from Pleurofusia. It can, however, be distin-
guished generically by its shorter form and broad, shallow notch. It is even more
closely related to Surculites proper, Pseudotoma, and Megasurcula, sharing with them
the broad conical shape and wide shallow notch, so that it is here not separated from
them generically. ... It probably was derived from Turricula, perhaps indirectly,
and later gave rise to, or is a branch of the stock that gave rise to, Pseudotoma and
Megasurcula. The intergradations between it and the last named are evident in
the California and Washington Tertiary. Clinura and typical Surculites are both
well represented in the Eocene of the Pacific coast, . . . Clinura lived on into
the Miocene, where it is represented by [Megasurcula] keepi (Arnold), ... "
It is clear from this, however, that Grant & Gale were basing their arguments largely
upon the characters of the species in the Eocene of North America which they had
referred to Clinura ; and those species, as pointed out above, do not possess the deep
posterior sinus which is considered here to be the most essential character of the
genus. Their opinions, therefore, do not apply to Clinura in its proper sense.
But, in any case, Wrigley (1939 : 282, 283) recognised quite correctly that the
genus Surculites was not a turrid, having no true sinus or notch like that of the typical
members of the family. He concluded that " The sinuous growth lines [of Surculites]
here considered are not an unequivocal mark of the Turridae, they may occur in
other families, linked with and proportionate to whorl-carination. Surculites is long
extinct, we have no guidance from the animal anatomy and, in the light of the
foregoing comparisons, it seems best to place the genus in a not too determinate
position between the Fusinidae and the Buccinidae . . . On the other hand,
he also noted that " Surculites has some resemblance, especially in general form, to
Clinura Bellardi, but in that genus the growth lines sweep back over the rear slope
four or five times as far as those of Surculites, here showing a Turrid character, or
rather, a contour often seen in the Conidae." Thus, if the species properly referred
to Clinura have no affinity with Surculites, then Clinura may be a turrid even though
Surculites is not. Further, this means that Clinura cannot be regarded as a sub-
genus of Surculites (in which classification Grant & Gale (1931) were followed by
Beets (1943^, 19436, 1951)), nor can Thatcheria be considered as part of the latter
genus (as in Beets, 19436, 1951).
The true characters of Clinura and Thatcheria not only link those genera with the
Turridae but with a particular sub-family thereof, the Daphnellinae. Of all the
turrid sub-families listed by Powell (1942 : 29), only the Daphnellinae have (some-
times) a diagonally cancellate protoconch (see Text-fig. 7) ; only the Daphnellinae
possess a juxtasutural posterior sinus ; only the Daphnellinae and the Mangeliinae,
together with a few genera in other sub-families, lack an operculum ; and the radula
of the Daphnellinae, which has marginal teeth only, is very similar to that of
Thatcheria. Indeed, Kuroda & Habe wrote of Thatcheria mirabilis (1954 : 81)
that the protoconch " shows the obliquely reticulated texture of sculpture, suggest-
ing that it is also of a Daphnelloid ", and (p. 80) that the species has a radula, " whose
2QO GASTROPOD GENUS THATCHERIA AND ITS RELATIONSHIPS
peculiar shape shows that it has a close affinity with the Daphnellinae".
But Powell (1942 : 167) had two objections to the location of Thatcheria and its
allies in the Daphnellinae, " the lack of a typical Daphnellid apex . . . and the
actual form of the sinus ". It has already been shown that the only Thatcheria
protoconch known at that time, that of T. liratula [Waitara], was so badly abraded
as to be useless ; but Powell was correct in supposing that the form of the Thatcheria
sinus (either as he wrongly believed it to be in T. mirabilis, having been misled by
Sowerby's figure to Angas' description, or as he actually observed it in T. liratula}
was not quite like that of the Daphnellinae. The other differences between the
Daphnellinae on the one hand and Clinura and Thatcheria on the other (listed
below in Table II) cannot preclude the possibility that the Clinura-Thatcheria
group originated from the Daphnellinae in the Upper Oligocene or Lower Miocene
by the development of a larger, pagodiform shell ; even the protoconchs of Clinura
and Thatcheria, now known from one good specimen of each of three species (C.
calliope, C. trochlearis, T. mirabilis}, agree with such a theory. But, from the taxono-
mic viewpoint, the consistent difference between the groups in the form of the
Form of posterior sinus
(juxtasutural)
Protoconch (see Text-figs.
3-7)
Spire ....
Whole shell
Earliest known occurrence .
Present trend
TABLE II
Daphnellinae
Typically a " reversed L "
Generally diagonally cancel-
late ; but never exactly like
that of Clinura and Thatcheria
Rarely pagodiform, if ever
Generally small
Oligocene, perhaps even Cre-
taceous
Still increasing
Clinura and Thatcheria
Smoothly arcuate, with outer
lip drawn far forward
Diagonally cancellate (where
adequately known)
Always more or less pagodi-
form
Always fairly large
Lower Miocene
Seemingly tending towards
extinction
posterior sinus is more fundamental. All things considered, it seems that, while
Clinura and Thatcheria should be placed in the Turridae and may well have evolved
from the Daphnellinae, they form a group which is so distinctive that it is best not
to regard it as part of the Daphnellinae.
The only logical alternative is to erect for them a special sub-family within the
Turridae, the Thatcheriinae; i.e., it is proposed that the family Thatcheriidae Powell
should be reduced in rank and the name of the taxon amended accordingly. The
Thatcheriinae have a particularly close relationship with the Daphnellinae. The
pagodiform habit, so characteristic of the former sub-family (especially its later mem-
bers), has been evolved in a very similar manner in other sub-families of the Turridae,
e.g. in the Cochlespirinae (compare Leucosyrinx (Aforia} circinata minatoensis
Otuka 1949, pi. 13, fig. n, from the Pliocene of Japan) ; but the Thatcheriinae may
be easily distinguished from such parallel developments by the form of the posterior
sinus.
GASTROPOD GENUS THATCHERIA AND ITS RELATIONSHIPS 291
Formal classification
Family TURRIDAE
Sub-family THATCHERIINAE Powell 1942 nom. transl. herein
Shell generally fairly large. Spire short relative to size of last whorl, very slightly
coeloconoid, more or less pagodiform with deep suture and well developed carina.
Posterior sinus on ramp of last whorl, juxtasutural, rounded, with vertex close to
suture and deepened by forward extension of outer lip ; outer lip simple and slightly
convex between carina and anterior canal ; distinct collabral growth-lines developed
accordingly on ramp and outer face of whole teleoconch. Protoconch diagonally
cancellate. Spiral ornament on outer face of teleoconch, and usually on outer edge
of ramp next to carina ; transverse sculpture may be present on juvenile whorls of
teleoconch (sometimes also on adult whorls), generally leading to formation of
tubercles or notches on carina, but, alternatively, it may be entirely absent.
Columella smooth, very slightly twisted. Operculum unknown and probably
absent ; soft anatomy of the only Recent species typically turrid ; radula of that
species like that of the Daphnellinae, with one pair of marginal teeth and no others.
OCCURRENCE. Lower Miocene to Recent ; Mediterranean Basin and Western
Pacific.
Genus CLINURA Bellardi 1875
?[i93i. Waitara Marwick, p. 149.]
DIAGNOSIS. Spire slightly or moderately pagodiform with carina generally
nearer to abapical suture ; juvenile whorls of teleoconch (sometimes also adult
whorls) bear transverse sculpture, generally leading to formation of tubercles or
notches on carina ; spiral ornament usually well developed.
TYPE-SPECIES. Murex calliope Brocchi 1814 by subsequent designation of
Bellardi, 1877. Upper Miocene to Lower Pliocene ; Italy and Southern France.
RANGE. Lower Miocene to Lower Pliocene (mainly Middle and Upper Miocene) ;
Europe, East Indies, New Zealand.
REFERRED SPECIES. C. generosa (Marwick 1931) [Waitara} ; Lower Miocene, New
Zealand.
C. trochlearis (M. Homes 1854) [Pleurotoma] ; Middle Miocene, Vienna Basin and
Italy.
C. controversa (Bellardi 1847) [Pleurotoma] ; Upper Miocene, Italy.
C. bituminata (Beets 1943) [Surculites] ; Upper Miocene, Buton (Celebes).
SPECIES REFERRED PROVISIONALLY. C. waitaraensis (Marwick 1926) [Turricula] ;
Upper Miocene, New Zealand. (May still represent distinct genus Waitara Marwick
[Clinura may also include :
C. sopronensis (Wolf 1870) [Pleurotoma} ; Upper Miocene, Vienna Basin.
C. subtrochlearis (Friedberg 1912) [Surcula] ; Upper Miocene, Poland,
C. sabatiorum Bellardi 1877 ; Lower Pliocene, Italy.
Other species from the Miocene and Lower Pliocene of Europe.]
292 GASTROPOD GENUS THATCHERIA AND ITS RELATIONSHIPS
Genus THATCHERIA Angas 1877
1928. Cochlioconus Yokoyama, p. 338.
DIAGNOSIS. Spire moderately to extremely pagodiform, with carina generally
nearer to adapical suture ; no transverse ribbing, no tubercles or notches on carina
(except for minute nodules on apical whorls of teleoconch in earlier forms) ; spiral
ornament usually weak.
TYPE-SPECIES. Thatcheria miraUlis Angas 1877 by monotypy. Recent ; off
Japan.
RANGE. Upper Miocene to Recent (mainly Pliocene) ; Western Pacific (New
Zealand, Fiji, East Indies, Japan).
REFERRED SPECIES. T. carinata (Martin 1933) [Ciyptoconus] ; Upper Miocene,
Buton (Celebes).
T. pagodula (Powell 1942) [Waitara] ; Upper Miocene or Lower Pliocene, New
Zealand.
Thatcheria sp. nov. aff. pagodula (Powell 1942) [Waitara] ; Lower Pliocene, New
Zealand.
T. liratula (Powell 1942) [Waitara] ; Lower Pliocene, New Zealand.
T. vitiensis sp. nov. (see below) ; Lower Pliocene, Fiji.
Thatcheria sp. nov. Beets 1951 ; Pliocene, East Borneo.
T. gradata (Yokoyama 1928) [Cochlioconus] ; Pliocene, Japan.
T. cf. gradata (Yokoyama 1928) MacNeil 1960 ; Pliocene (and Upper Miocene?),
Okinawa.
VII. A NEW SPECIES OF THATCHERIA FROM
THE PLIOCENE OF FIJI
Thatcheria vitiensis sp. nov.
PL 47, figs. 4-6
DERIVATION OF NAME. Viti — the Fijian name for Fiji.
DIAGNOSIS. A species of Thatcheria with a fairly thick shell, a relatively long and
slender spire and a moderately projecting carina. The whorls bear dense, distinct
spiral ornament on the outer face and a few weaker spiral striations on the outer part
of the ramp. The first few whorls of the teleoconch bear minute nodules on the
carina.
HOLOTYPE. The unique specimen, Brit. Mus. (Nat. Hist.) Palaeont. Dept. no.
G. 91124 (collector's field number VL . i) ; it lacks the extreme apex of the spire,
the outer lip and the anterior canal.
OCCURRENCE. In tuffaceous marls of the Vanua Levu Formation, of probable
Lower Pliocene age. On the new south coast road near Savusavu, Eastern Vanua
Levu, Fiji ; a large breast-cut 4 miles east of Salt Lake point where the road crosses
higher ground north of Naweni.
DESCRIPTION.
[Note, (i) The protoconch is entirely lacking, and the condition of the early
whorls is poor. (2) Although the whole of the outer lip has been broken off, the
ramp of the last whorl is preserved up to its end. Had it extended further, it
GASTROPOD GENUS THATCHERIA AND ITS RELATIONSHIPS 293
would surely have left some indication of its attachment to the outer face ; and,
in any case, the end of the suture is coincident with the adapical end of the last
striation marking the limit of growth of the inner part of the mantle. (3) The abapi-
cal end of the shell is also missing, but may be reconstructed with a fair degree of
probability by reference to T. mirabilis. (4) The surface of the shell is abraded,
but only very slightly.]
MEASUREMENTS.
Length as preserved ......... 45 mm.
Estimated total length . . . . . . . . .51 mm.
Height of spire as preserved . . . . . . . .22 mm.
Estimated total height of spire . . . . . . -23 mm.
Estimated height of last whorl . . . . . . . -34 mm.
Maximum diameter (from end of carina to point diametrically opposite
on carina of last whorl) . . . . . . . .26 mm.
Internal spire angle . . . . . . . . .38°
External spire angle . . . . . . . . -57°
Angle of declination of suture (from plane perpendicular to shell axis)
(see Text-fig. 9) .......... 14°
Angle of declination of carina (see Text-fig. 9) . . . .8°
This species shows a less extreme development of the characteristic form of
the type-species (T. mirabilis). The shell is rather thick and solid ; this applies
especially to the carina, the outer part of the ramp adjacent thereto, the columella,
and the outer lip in the region of the anterior canal. The internal spire angle is a
little less than in other species of the genus, the external spire angle much less than
in any other species except the Miocene T. carinata ; the index of pagodiformity is
greater than in T. carinata or the much stouter T. pagodula, about the same as in the
slightly stouter T. limtula, and less than in the other species. The height of the spire
is about 80% of the estimated height of the aperture ; the corresponding figure for
T. liratula is about 50% (fide Powell) and for T. mirabilis 57%.
The whorl profile shows that the carina projects rather acutely ; in the younger
whorls both the outer face and the ramp are appreciably concave, the outer face being
directed steeply downwards. On the penultimate and last whorls, however, the
outer face shows a more or less straight profile ; and the ramp of the last whorl has
a very marked angular concavity, the angle lying rather nearer to the suture than to
the carina.
The spiral ornament is very much like that of T. liratula and is clearly shown in
Plate 47, figs. 4-6 ; the striations pass further into the interior of the last whorl,
beyond the inner lip, than they do in T. mirabilis. There are very faint indications
of minute nodules on the carina of the earliest whorls, just as in T. liratula. The
growth-lines are also like those of T. liratula, the parts on the ramp showing that
the characteristic posterior sinus of the Thatcheriinae was present ; the parts on the
outer face, however, are inclined much less obliquely to the shell axis than they are
in the New Zealand species. One peculiarity of the holotype, if not of the species,
is the series of undulating striations on the inner lip which mark the limit of growth
of the inner part of the mantle.
294 GASTROPOD GENUS THATCHERIA AND ITS RELATIONSHIPS
REMARKS. This species is closely related to Thatcheria liratula (Powell), to which
it has an especial resemblance in its degree of pagodiformity, in the type and distri-
bution of the spiral ornament, and in the presence of minute nodules on the first
few whorls of the teleoconch. It may be distinguished from T. liratula, however,
by its more acute spire, which is much longer relative to the aperture ; by the less
oblique growth-lines on the outer face ; and by the fact that its carina is rather more
acute, with the result that both the ramp and the outer face of each whorl generally
appear a little more concave.
T. liratula is from the Opoitian (Lower Pliocene) of New Zealand. The resem-
blance of T. vitiensis to T. liratula rather than to any other species tends to confirm
the suspected Lower Pliocene age of the marls from which T. vitiensis was collected.
T. vitiensis is also like the Okinawan form which MacNeil (1960) described as
T. cf. gradata, and which, as pointed out above, seems to have a close affinity with
T. liratula. The Okinawan shell, however, differs from the Fijian species in certain
details. The carina appears to be rather higher on the whorl and to project further,
producing an almost flat ramp and a more pagodiform spire (just as in T. gradata
itself, from Japan) ; while the nodules on the carina of the early whorls of the teleo-
conch, if not better developed, are at least much better preserved than in T.
vitiensis.
ASSOCIATED FAUNA. Four other shells (three gastropods and one lamellibranch)
were found in the Vanua Levu Formation with Thatcheria vitiensis. They will be
described elsewhere.
VIII. ACKNOWLEDGMENTS
The specimen described above as the holotype of Thatcheria vitiensis sp. nov. was
collected by Mr. R. W. Bartholomew of the Fiji Geological Survey Department, and
was sent to London for examination by permission of Dr. N. J. Guest, Chief Geo-
logist. Their interest and encouragement are gratefully acknowledged.
My especial gratitude is due to Dr. L. R. Cox, F.R.S., for all the help he so patiently
gave me. Among the many others who kindly offered useful information, sugges-
tions and criticism were Dr. C. Beets, Mr. S. P. Dance, Dr. N. B. Eales, Mr. C.
Matheson, Prof. J. E. Morton, Mr. C. P. Nuttall, Dr. A. W. B. PoweU, Mr. E. P.
Smith, Mr. F. M. Wonnacott, and, in particular, Mr. D. L. F. Sealy.
Dr. C. A. Fleming, Chief Palaeontologist to the New Zealand Geological Survey,
allowed me to borrow the figured paratype of Waitara liratula Powell.
Thanks are also due to Dr. G. Archey, Director of the Auckland Institute and
Museum, for permission to reproduce Powell's text-figures of the protoconchs of
Waitara liratula and Cryptodaphne pseudodrillia ; to Drs. T. Kuroda and T. Habe, of
Kyoto University, for permission to reproduce their text-figure of the protoconch of
Thatcheria mirabilis ; to Mr. D. L. F. Sealy, who copied those text-figures and made
the original drawings of the protoconchs of Clinura calliope and C. trochlearis ;
and to Mr. N. P. G. Tanti, who took some of the photographs.
GASTROPOD GENUS THATCHERIA AND ITS RELATIONSHIPS 295
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PLATE 47
Thatcheria mirabilis Angas
Holotype : British Museum (Nat. Hist.), no. 1960.154.
Recent ; off Japan.
FIG. i. — Apertural view, x i.
FIG. 2. — Apical view, x i.
FIG. 3. — Oblique view of ramp to show growth-lines, x 2.
Thatcheria vitiensis sp. nov.
Holotype : British Museum (Nat. Hist.), no. G. 91124.
Lower Pliocene ? (marls of Vanua Levu Formation) ;
near Savusavu, Eastern Vanua Levu, Fiji.
FIG. 4. — Apertural view, x i£.
FIG. 5. — Apical view, x i.
FIG. 6. — Oblique view of ramp to show growth-lines, x 2.
Thatcheria livatula (Powell)
Paratype : New Zealand Geological Survey, no. TM. 2864.
Lower Pliocene (Opoitian) ; N.Z.G.S. loc. 1543, mudstone and argillaceous sandstone beds,
Mangawhero Stream, Taramarama (S.W.) S.D., Wairoa, New Zealand.
FIG. 7. — Lateral view of spire, x 2^.
FIG. 8. — Oblique view of ramp to show growth-lines, x 6.
Clinura calliope (Brocchi)
British Museum (Nat. Hist.), no. G. 79439.
Lower Pliocene (Plaisancian) ; Biot, near Antibes,
Alpes Maritimes, France.
FIG. 9. — Apertural view. X about ij.
FIG. 10. — Abapertural view, x about i£.
FIG. ii. — Oblique view of ramp to show growth-lines. X5-
FIG. 12. — Protoconch and early whorls of teleoconch. x 125.
Bull. B.M. (N.H.) Geol. 7, 9
PLATE 47
12
INDEX TO VOLUME VII
New taxonomic names and the page numbers of the principal references are printed in Bold type.
An asterisk (*) indicates a figure
ABBASS, H. L. 173-196
Abbey Wood, Kent 6, 8
A caste alifrons 100, 103
exilis 100
Acastinae 97
Acidaspis caractaci 239
harnagensis 237
magnospina 123
primordialis 122, 237
semievoluta 69, 122
sp. 123
Actaletes neptuni 156
Adapidae 14, 21, 24, 25, 27, 31, 32, 34
Adapis 4, 10, 12-15, 17-23, 30, 32
angustidens 33
magnus 17, 19, 33
minimus 13
parisiensis 10, 18, 25, 29, 30, 33
priscus 33
rutimeyeri 33
sciwrews 33
Aeluravus 29
Aeschna 163
liassina 164
Akera 260
Akeridae 267
Alsaticopithecus 25, 31, 33
leemanni 25, 27, 31, 33
Alternata limestone 219
Amphilemur 25, 33
eocaenicus 25, 33
Amphilemuridae 23, 25
Amphitryon 251
radians 220
Amplexograptus perexcavatus 232
Ampyx linleyensis 78
pennatus 78
restrains 76, 78
salteri 78
tetragonus 78
Anachoropteris 208
involuta 208
Anagale 27
Anaptomorphidae 3, 24, 27
Anchomomys 3, 19, 21, 23, 31, 33, 34
fi 33
^4 nchomomys — contd.
latidens 21-23, 31, 33
pygmaeus 33
quercyi 21, 22*, 33
Anisozygoptera 155, 160
Anthropoidea 17
Anticlinura 263, 266
Anulatisporites anulatus 147 ; PI. 22, figs. 70, 71
sp. 148; PI. 22, fig. 72
Aotes 5
Apatemyidae 23, 26
Aphanolemur 15
gibbosus 15
Aphelotherium 32
Apiculatisporites 150
Apotropteris 208
Apterygota 156
Archaeopteridium 142
tschermaki 142 ; PI. 21, fig. 22
Archimediella 178
Architemidae 164
Arcotia 179
Asaphidae 229
Asaphinae 229
Asaphus marstoni 231
powisi 229, 232
seticornis 85
Asperdaphne versvestita 277
Asplenium 207
Asteroceras obtusum 155, 161
Atractopyge 91, 93, 130
aspera 91
atractopyge 93
michelli 93
scabra 91, 92, 93 ; PI. n, figs. 2, 3, 10
sp. 93, 94, ; PI. n, figs. 5, 8
Auckland Institute & Museum 294
Aurelia aurita 156
A zonotriletes 150
Balanus improvisus 156
Bancroftina typa 71
Zone 80, 93, 107, 125
Basilicus 229
marstoni 229
powisi 229, 232
3°°
INDEX
Birbal'Sahni Institute 151
Birmingham University 137, 151
Blattodea 158
Bourguetia saemanni 194
striata 194
Bothrodendron tenerrimum 139
Broeggerolithus 79-82, 129
broeggeri 79
Zone 221, 232
globiceps 71, 84
longiceps 71, 81-83
melmerbiensis 81 ; PI. 6, figs. 10, 13
nicholsoni 79, 80-83, I29 .' PI- 6> ngs. 7, n ;
PL 7, figs. 1-12
simplex 79, 8 1
soudleyensis 84
sp. 79, 83 ; PI. 8, fig. 2
transiens 81, 82, 83, 84, 129, 218 ; PI. 8, figs.
i, 3, 4, 6, 8, n
Brongniartella 82, 106, 129
ascripta 106, 107, 108, no ; PI. 15, figs, i, 2,
5, 8, ii ; PL 16, fig. 14
bisulcata 107, 108, 109, no, 129 ; PL 16, fig. 8
depressa 108, 109, no ; PL 16, figs, i, 7,
10, 13
minor 106, 107 ; PL 15, fig. 4, ; PL 16, fig. 11
subcarinata 107, no
parva 106
platynota 110
rwrfis 107
sp. 108
Buccinidae 259, 267, 289
Burrirhynchia 42, 43, 58, 59
cantabrigensis 42, 59*, 60, 61
leightonensis 59, 60, 61
Caenopithecus 14, 19, 21, 27, 29, 30, 31, 33, 34
lemuroides 29, 33
Calamospora 144, 150
mutabilis 144, PL 22, fig. 44
sp. 144 ; PL 22, fig. 44
Calathiops sp. 143 ; PL 21, figs. 27-29
Calliops brongniarti 104
jukesi 100
var. vicina 104
Calymene in
aldonensis 113
bigener 113
blumenbachii var. caractaci 114, 216, 217, 225
brevicapitata 223
callicephala 111
caractaci in, 217
diademata 116
fatua 219
forcipata 235, 236
grayae 113
incerta 219
meeki in
104
planimarginata
Calymene — contd.
pulchra 220
quadrata 113, 227
senaria in
verrucosa 91
Calymenidae 111, 215, 216
Calyptaulax 98, 130
actonensis 100
altumi 99, 100
compressa 100
glabella 98, 100
norvegicus 98, 99
planiformis 98, 99, 100 ; PL 13, figs. 1-5
schucherti 100
sp. 98, 99
Camptotriletes sp. 146 ; PL 22, fig. 58
Cantius 3, 4, 5, 6, 8, 10, 14, 22, 34
e^si 4, 6, 7, 8, 9, 32 ; PL i
Capillae 42
Capillirhynchia wrighti 44
Cardium edule 156
Carpolestidae 24
Ceciliolemur 27, 30
Ceratopogon 156
Ceraurus octolobatus 88
Cerithiacea 180
Cerithium turritellatum 183
unicarinatum 182
CHARIG, A. J. 255-297
Charmouth, Dorset 155
Chasmops 104, 129, 248
conicophthalma 106
extensa 104, 106, 115, 229 ; PL 13, fig. 8
jukesi 100
maxima 105 ; PL 13, fig. 6
sp. 105 ; PL 13, fig. 12
Chasmopsinae 104
Cheiruridae 86
Cheirurus octolobatus 88
Chironomus 156, 157
Chomatopyge 119
Chondrites plumosus 150
Chondrosteus sp. 155
Chonetoidea 115
sp. 84
Cibotium barometz 205
regale 205, 206
Cirratriradites 150
sp. 148 ; PL 23, fig. 74
Clavatulinae 262
Clinura 257, 262-264, 2^6, 268, 270, 271, 272*,
273, 277-283, 286-289, 291
bituminata 262, 279-282, 286, 291
calliope 263, 266, 271-274, 275*, 276, 278-280
282, 286-288, 290, 294
carinata 279-282
controversa 279, 280, 287, 288, 291
elegantissima 263, 279
generosa 286, 288, 291
monochorda 263
INDEX
301
Clinura — contd.
peruviana 263
sabatiorum 279, 291
sopronensis 279, 291
subtrochlearis 291, 279
Thatcheria group 278, 286, 288
trochlearis 271, 272, 274, 275*, 279, 280,
288, 290, 291, 294
waitaraensis 286, 291
Clinuromella 263
Clinuropsis 262, 263, 266, 278-280
calliope 266
monochorda 263
Cochlespirinae 261, 266, 290
Cochlioconus 260-262, 265, 267, 268, 270, 278
288, 292
gradatus 259, 260, 265
Coenopterideae 210
Coleoptera 159, 167
Colpocoryphe aldonensis 113
Colposigma 179
Conidae 261, 267, 268, 277, 288, 289
Conolichas 120, 129
melmerbiensis 120, 121 ; PI. 17, figs. 14,
PI. 1 8, figs. 1-4
Conorbiinae 261
Conularia 150
Conus 259-261, 267, 268
araneosus 288
pennaceus 277
quercinus 260
Convolutispora sp. 146 ; 1*1. 22, fig. 59
tessellata 146
Coryphodon 8
Costellae 42
Costonia ultima Zone 235
Craeseops 25
Craiginia 179
Cretaceous Gastropoda 173-196
Cretirhynchia 39, 43, 59-61
norvicensis 58*, 61
octoplicata 49
plicatilis 60
Cristatisporites 150
CROFT, W. N. 197-211
Cryptoconus 262, 279, 292
Cryptodaphne pseudodrillia 275*, 294
Cryptolithinae 79
Cryptolithus broeggeri 79
gracilis 84
nicholsoni 79
transiens 82
Cryptothyris paracydica 71
Cupedidae 167, 169
Cupes 167
capitatus 167
Cybele loveni 94
var. girvanensis 94
iwrwcosa 91, 93
Cybelinae 91
Cyclogranisporites 150
amplus 145 ; PI. 22, figs. 49-51
orbicularis 145
sp. 145 ; PI. 22. fig. 52
Cy do stigma 138
Cyclothyrid 42
286, Cyclothyrinae 39, 43
Cydothyris 39*, 40*, 41*, 42, 43
americana 40
antidichotoma 41, 42, 44, 47, 48*, 49*, 56 ;
PI. 4. figs. 3-6
compressa 54, 61 ; PI. 5, fig. 8
depressa 41, 50, 56 ; PI. 4, fig. n
difformis 41, 42, 44, 45, 51, 52*, 53*, 54, 58 ;
-280, PI. 5, figs. 1-7
latissima 39, 41, 45, 46*, 50, 56-58, 60 ; PI. 4,
figs. 7, 8
lepida 41, 42, 55, 56, 57 ; PI. 4, fig. 10
levis 41, 42, 56, 57 ; PI. 4, fig. 9
plicatilis 60
scaldisensis 42, 44, 46, 57 ; PI. 4, fig. i
schloenbachi 42, 50, 55, 56 ; PI. 4, figs. 12, 13
subtrigonalis 40
1 6 ; Cynodontomys 25
Cyphaspis jamesoni 126
megalops 123
nicholsoni 118
Cypraeacea 266
Cyrtometopinae 86
Dalmanella horderleyensis 71
Zone 217, 223
indica 71
Zone 222
/e/>ta 71, Zone 222
unguis 71, Zone 218, 219, 246
wattsi 71, Zone 218
Dalmania socialis 95
Dalmanitidae 95
Dalmanitina 95
mucronata 95, 96
matutina 95, 96 ; PI. 12, figs, i, 5, 6, 9, 12
Dalmanitinae 95
Dapedium granulatum 155
sp- 155
Daphnella 277
cancellata 277
Daphnellinae 257, 261, 265, 267, 277, 289-291
Daubentonia 26
DEAN, W. T. 65-134, 213-254
Decoroproetus 126
fearnsidesi 243
Defrancia pagoda 266
Densosporites 150
sp. 148 ; PI. 23, fig. 73
Diacalymene 113, 114, 116
bigener 113
diademata 113, 114
marginata 116, 227 ; PI. 13, fig. 13 ; PI- 14,
fig. ii
302 INDEX
Diacalymene — contd .
praecox 113, 225, 227
Diastatomma liassina 164
Diastatommites 164
liassina 161, 164 ; PI. 27, fig. 3
Dicranograptus clingani Zone 72, 226, 232, 242
Dictyotriletes sp. 147 ; PI. 22, fig. 64
Didymograptus superstes Shales 113
Dimeropyge 119
Dimeropygidae 118, 119
Dindymene 89
duftonensis 89, 90 ; PI. n, fig. 13
fridericiaugusti 89
ornata 90
plasi go
sp. 89, 90 ; PI. ii, fig. 6
Dindymeninae 89
Diplograptus multidens Zone 129, 226, 232
perexcavatus 232
Diplopteridium 140-143
holdeni 137, 140, 142 ; PI. 20, figs. 8-15
teilianum 141, 143
Diptera 156, 158
Dolerorthis sp. 77
Dry brook Sandstone 137, 149
Duftonia 69, 97, 103
lacunosa 97
Elcanidae 158, 159
Encrinuridae 88
Encrinurinae 88
Encrinurus 88, 89
sp. 88 ; PI. n, figs, i, 4
Endosporites sp. 148 ; PL 23, fig. 75
Ensifera 165
Entomolestes 25
Entomostracites crassicauda 233
laticauda 251
punctatus 88
Eobronteus 251
sp. 251 ; PI. 46, fig. 10
Erinaceoidea 25
Estoniops 100, 103
alifrons 100, 101-104, I29 > PI- I2> nSs- 2' 4>
7, 8, 10, 13
exilis 103
jukesi 101
Europolemur 12, 13, 27, 31, 33
£/a#i 9, 12, 13, 33
Ficidae 259, 267
Ficus 259, 267
Fiji Geological Survey 294
Filicales 201
Flexicalymene 82, in, 112, 113, 114, 118, 216,
219, 236
acantha 112, 216, 217, 219, 220, 225 ; PI. 37,
figs, i, 3-6, 14
brevicapitata 112, 220
cambrensis 112, 219
Flexicalymene- — contd.
caractaci 112, 114, 217, 218-220 ; PI. 13, fig.
10 ; PI. 38, figs. 2, 4-6, 8, ii, 12
cobboldi 218, 220 ; PI. 38, figs, i, 3
croneisi 223
declinata 112, 220
forcipata 112
horderleyensis 221, 223 ; PI. 37, figs. 2, 8 ;
PI. 40, figs. 7, 8
incerta 220
laticeps 115, 224
limba 113, 221, 223 ; PI. 37, fig. 16
onniensis 69, 112, 115, 223 ; PI. 13, fig. 7 ;
PI. 14, figs, i, 2, 7, 10
planimarginata 112, 114, 217, 219, 220 ; PI. 37,
ng- 15
pusulosa 113, 217, 220, 221, 223 ; PI. 37,
figs. 9, ii, 12
salteri 224
senaria 112
shirley i 112
sp. 221, 223
trigonoceps 217, 218
Fucus sp. 156
Furcirkynchia furcata 44
Fusidae 267
Fusinae 267
Fusinidae 267, 289
Fusininae 267
Fusinus 259, 267
Fusus 259, 267
Galago 17, 23
crassicaudatus 6
Galeodidae 261, 267, 268
Galigidae 23
Gastropoda 175, 180
Geological Society, London 69
Geological Survey & Museum 59, 102, 103, 187,
224, 225, 240, 245
GesneropithexSl, 33
peyerizi, 31. 33
Glasgow University 137
Glyptocrinus Flags 221
Grammatopteris 208
baldaufi 209, 201
rigolloti 209
Granulatisporites 1 50
orbiculus 145 ; PI. 22, figs. 47, 48
sp. 145 ; PI. 22, figs. 47, 48
tenuis 144 ; PL 22, figs. 45, 46
Gravicalymene 112, 113, 114, 117, 225
convolva 116, 118, 225, 225
inflata 227 ; PL 39, fig. 6
jugifera 116, 118, 226 ; PL 13, figs. 9, ii ;
PL 14, figs. 3, 4, 8, 9
praecox 112, 216, 225, 226-228 ; PL 39, figs, i,
3,9, 12-14
sp. 227
Gryllacrididae 155,166
INDEX
3°3
Hagla 165, 166
Haglinae 165
Halle University 12, 34
Harknessella subquadrata Zone 220, 235, 245
Harpalideous Carabidae 168, 170
Harpalus schlotheimi 168
Haustator 178, 184, 187, 191
granulata 183, 184
granulatoides 183, 184
Hemiacodon 4-8
Hemifusus 259, 261, 267, 268
Heterophlebia westwoodi 161
Heterorthis alternata 219
Hirnantia sagittifera 71
Hoanghonius 4
Holcoelytrum 155, 167
giebeli 167, 168, 170, 171 ; PI. 27, figs. 6-8
schlotheimi 168, 170, 171 ; PI. 27, fig. 5
Holcoptera 168
HOLDEN, H. S. 197-211
Homalonotidae 106
Homalonotus ascriptus 69
bisculatus 106, 108
var. minor 106
rudis 107
Homalops altumi 99, 100
Homolichas melmerbiensis 120
Homolochinae 120, 235
Hydrobia 157
Hydrophilidae 157
Hymenoptera 159
Hyracotherium 8
Illaenidae 120, 233
Illaenus 129, 233
bowmani 120
fallax 233, 234
linnarssoni 120
sp. 233
Imperial College of Science, London 175
Inoceramus 155
Insectivora 25
Ireland National Museum 235, 236
Isotelinea 229
Isotelus brachycephalus 233
Isotelus powisi 229
Jurassobatea gryllacroides 166
King's College, London 61
Kingston Technical College 137, 151
Kjaerina bipartita Zone 71, 89, 219
geniculata 77, 91
typa Zone 71, 91, 218
Kjerulfina polycyma 83
Kloucekia 97
apiculata 72, 97, 129 ; PI. 12, fig. n
Knoxisporites 147
Kyoto University 294
Lacunosella 43
Lamellaerhynchia 43, 60
larwoodi 41
Leiotriletes sphaerotriangulans 143 ; PI. 21, figs.
3°. 31
LELE, K. M. 135-152
Lemuridae 24
Lemuriformes 5
Lemuroid Primates, Concept of 23
Lepidophyllum fimbriatum 137, 138, 139, 140,
150 ; PI. 20, figs. 18-20
Lepidorhynchia 43, 60
Lepidosigillaria whitei 140
Lepidostrobophyllum fimbriatum 138
Leptadapis 32
Leptocolpus 179
Leucosyrinx circinata minatoensis 290
Leymeriella tardefurcata Zone 41, 42, 50, 56, 57,
59,6o
Liassocupes 167
parvus 167 ; PI. 27, fig. 4
Liassophlebia 161
gigantea 163 ; PI. 27, fig. 2
jacksoni 162, 163, 164, ; PI. 25
magnified 161, 162, 163, 164 ; PI. 27, fig. i
sp. 164
wither si 164
Liassophlebiidae 160
Lichas aculeatus 235
aequiloba 120
laxata 121, 235
margaritifer 121, 235
melmerbiensis 69, 120
nodulosus 121, 235
segmentatus 235
sexspinus 235
verrucosa 234
Lichidae 120, 234
Lichinae 234
Lipura maritima 156
Lonchodomus 72, 77, 78
carinatus 78, 79
pennatus 77, 78, 79, 84, 129 ; PI. 6, figs. 1,3-5,
9, 12
politus 77
rostratus 72, 77
swindalensis 72, 77 ; PI. 6, figs. 2, 6, 8
London Clay 3
Lophotriletes 150
Loris 1 8
tardigradus 18
Lushius 4
Lycospora 150
bracteola 147 ; PI. 22, fig. 65
granulata 147
spp. 147 ; PI. 22, figs. 66-69
Macrocranion 25
Manchester Museum 140, 150
INDEX
Mangeliinae 289
Mathilda 179, 191
ahmadi 175, 177, 178, 181*, 193 ; PI. 32,
fig. 24
coxi 175, 177-179, 181*, 191, 192, 193 ; PI.
32, fig. 23
douvillei 192
faucignyana 192
quadricarinata 192
Mathildidae 175, 191
Matonia pectinata 205
Mayeria 259, 267
Mecoptera 158
Megachiromy aides 27, 29
M egasurcula 289
keepi 289
Megatarsius 13, 14, 27, 31, 33
a&eftg, 12, 13,33
Melongenidae 259, 267
Melongeninae 259, 267
Menocephalus nicholsoni 119
Mesalia 178
Mesotaphraspis 119
Metacupes harrisi 167
huebneri 234
M etopolichas 234
patriarchus 234
sp. 234 ; PI. 43, figs. 3, 4, 6, 7
verrucas a 234
Microchoerus 4, 31
edwardsi 3 1
erinaceus 3 1
ornatus 31
Microreticulatisporites 150
cribellarius 146 ; PI. 22, figs. 60, 61
spp. 146 ; PL 22, figs. 62, 63
Microsyopidae 23, 25, 33
Microsyops 25
Microtarsioides 27, 30
Mwre.r 263, 271
calliope 266, 272, 291
Nannopithex 19, 24, 32
filholi 32
pollicaris 32
raa&i 32
Navajovius j, 24
Necrolemur 4, 5, 15, 17, 22, 31, 32
antiquus 32
edwardsi 31
filholi 32
parvulus 32
raabi 32
zitteli 32
Necrolemurinae 4, 23, 31
Nekewis 262, 279
io 273
Nemagraptus gracilis Zone 77, 113, 129, 226, 232
Nereis diversicolor 156
Nerinaea unicarinata 182
Neuroptera 158
New Zealand Geological Survey 294
Notharctidae 5, 17, 24, 34
Notharctus 6, 12, 14, 15, 17, 30
Notopamphagopsis bolivari 166
Nycticebus 5, 15, 18-20
cougang 1 8, 20
Odonata 156, 158, 160
Odontopleuridae 122, 237
Odontopleurinae 237
Ogmocnemis 126, 243
calvus 126, 243, 245
Olenidae 241
Omomyidae 3-5, 7, 24, 32, 34
Omomyinae 3
Omowrys 4, 6, 7, 30, 32
belgicus 32
Onnia 84, 1 30
cobboldi Zone 71, 224, 234, 237
gracilis 84, 129 ; PI. 8, figs. 12, 13
Zone 71, 78, 85, 92, 95, 99, 115, 122, 128,
223, 224, 226, 228, 234
ornata 220
superba 129
pusgillensis 84, 130 ; PI. 8, figs. 5, 7, 9, 10
Zone 71, 78, 84, 85, 88, 92, 95, 99, 115,
128, 224, 242, 251
Onnicalymene 104, 112, 113, 115, 129, 223
jemtlandica 113
laticeps 113, 115, 224 ; PI. 14, figs. 5, 6; PI. 38
figs. 7, 10, 14, 15
salteri 113, 224 ; PI. 38, figs. 9, 13
onniensis 84, 113, 115, 129, 223, 224 ; PI. 39,
figs. 2, 10, ii ; PI. 40, fig. 9
Onniella 104, 115, 248
broeggeri 71, 84
inconstans 71
reuschi Zone 71, 83, 218, 240
Orbirhynchia 45
Ordovician trilobites 215
Orimops 1 1 1
Orthophlebia 158
Orthophlebiidae 158
Orthoptera Saltatoria 158, 165
Osmundaceae 210
Otarion 123, 242
diffractum 123, 242
isoplates 124
planifrons 123, 124
sp. 123, 242 ; PI. 17, figs. 4, 12 ; PI. 45, fig. i
Otarionidae 123, 242
Ourayia 6, 18
OWEN, E. F. 37-63
Palaeolemur 32
Parabasilicus 229
marstoni 231, 232
powisi 229, 230-233 ; PI. 40, figs, i, 3-5 ;
PI. 41, figs, i, 2 ; PI. 42, fig. 9
INDEX
305
Parabasilicus — contd.
shirakii 232
typicalis 229
Paracybeloides 94, 129
girvanensis 94, 95, 96 ; PI. n, figs. 7, 9, 12,
14, 15
sp. 95, 103 ; PI. ii, fig. ii
Parahoplites nutfieldensis Zone 42, 46, 50
Pareora 189
Paromomyidae 24, 27
Pelycodus 6, 7, 14, 18, 29, 30, 32
helveticus 32
Periconodon 4, 30, 32
helveticus 30, 32
Perodicticus 12
Petrophlebia 160, 161
anglicana 160, 161
anglicanopsis 159, 160, 161 ; PI. 24, figs, i, 2
Phacopidina 97
apiculata97 ; PI. 12, fig. ii
harnagensis 97
Phacops alifrons 100,103
apiculatus 97
brongniarti 98
jukesi 100-103
var. vicina 104
panderi 103
phillipsi 97
Phaneropterinae 166
Pharostoma pulchrum 220
Phenacolemur 26, 27
Phenacolemuridae 24
Pholidophorus sp. 155
Planisporites 150
delucidus 145
granifer 146 ; PI. 22, fig. 56
minimus 145 ; PI. 22, figs. 53—55
sp. 146 ; PI. 22, fig. 56
Platycalymene 1 1 1
Platylichas 121, 122, 235
laxatus 71, 121, 122, 235, 236, 237 ; PI. 43,
figs, i, 2, 5, 8-12
melmerbiensis 120, 121
sp. 122
Plesiadapidae 24, 27
Plesiadapis 26, 27, 33
Pleurofusia 289
Pleurograptus linearis Zone 72, 100, 226
Pleurotoma 262, 263, 271, 274, 279, 288, 291
calliope 273, 288
controversa 288
Pleurotomella 263
calliope 266, 274
monochorda 263
packardii 266
Pleurotomidae 263, 268, 288
Pleurotomoides 266
Plicarostrum 43
Portlockia apiculata 97
POWELL, A. W. B. 294
Primaspis 122, 237
ascitus 241
caractaci 122, 238, 239, 240, 241 ; PI. 44, figs.
3. 7-9- ii, 13, M
harnagensis 122, 237, 238, 241 ; PI. 44, figs.
1,4,6,8
semievoluta 122, 123, 241 ; PI. 17, figs. 3, 10,
ii, 13, 15
Princeton University 34
Proetidae 124, 243
Proetidella 124, 126, 129, 243, 245
fearnsidesi 124, 126, 243, 245, 246 ; PI. 45,
figs. 3-8, 12, 14
marri 124, 126 ; PI. 16, figs. 4, 6, 9 ; PL 17,
figs. 5, 6, 8, 9
sp. 246 ; PI. 45, fig. 13
Proetidellinae 124, 243
Proetus 231
asellus 245
decorus 126
girvanensis 126
ovatus 245
sp. 246
Progalago 19, 21-23, 34
dorae 2 1
sp. 22*
Progonophlebia 162
Promi croceras planicosta 155
Pronycticebus 3, 4, 12-15, 17-23, 33, 34
gaudryi 14, 16*, 17, 33
Prophalangopsidae 155, 158, 165, 166
Prophalangopsis 166
Prosimii 27, 31
Prosobranchia 180
Protoadapis 3, 6, 7, 8, 9, 10, 12, 13, 14, 15, 19,
21, 29-34
angustidens ii*, 12, 33
brachyrhynchus ii, 33
copei 9
curvicuspidens 10, ii*, 12, 13, 33
eppsi 3, 5, 8, 10, 32
klatti 12, 13, 29, 30, 33 ; Pis. 2, 3
recticuspidens 10, 33
Protohagla 165, 166
langi 158, 165, 166 ; PI. 26
Psalixylon 208
Pseudoloris 32
abderhaldini 32
parvulus 32
Pseudomelaniidae 194
Pseudospkaerexochus86, 130
octolobatus 87, 88 ; PI. 10, figs. 7, 9-12
Pseudotoma 262, 289
Pseudotominae 263
Pteronemobius 158
Pterygometopidae 98
Pterygometopinae 98
Pterygometopus 100
grayae 113
jukesi 101-103
306 INDEX
Pterygometopus — conid.
pander i 103
sclerops 103
sp. 97
trigonocephala 103
Pugilina 261, 267
Punctatisporites 150
debilis 144
minutus 143, 144 ; PI. 21, figs. 32-37
spp. 144 ; PI. 21, fig. 43
subobesus 144 ; PI. 21, figs 38-42
Pyrula 259, 267
Pyrulidae 259, 267
Raphiophoridae 72
Reacalymene 112, 219, 220
croneisi 223
holtedahli 226
horderleyensis 221, 223 ; PI. 37, figs. 2, 8 ;
PI. 40, figs. 7, 8
limba 113, 220, 221 ; PI. 37, fig. 16
pusulosa 112, 113, 220, 221, 223 ; PI. 37, figs.
9, II, 12
Sp. 221
Remopleurella 215, 249, 250, 251
burmeisteri 250, 251 ; PI. 45, fig. 2 ; PI. 46,
figs. 6, 8, 9, ii, 12
Remopleurides 104, 115, 127, 129, 246
biaculeatus 127
burmeisteri 128
colbii 127, 128, 246, 249
dalecarlicus 246
latus 248
var. granensis 248
kullsbergensis 128, 248, 249
onniensis 247, 248, 249 ; PI. 46, figs. 4, 5
radians 220, 251
spp. 127, 128, 246, 248 ; PL 18, figs. 5-13 ;
PL 46, figs. 3, 7
validus 128, 248
warburgae 246, 247, 248 ; PL 46, figs, i, 2
wimani 248
Remopleurididae 127, 246
Remysporites 149, 150
drybrookensis 148 ; PL 23, figs. 76-81
sp. 149 ; PL 23, fig. 82
Reticulatisporites 147, 150
Reuschella sp. 77
Reuscholithus reuschi Zone 217, 226, 231, 238, 245
Rhacopteris geikiei 141
petiolata 141
Rhodea 141
Rhynchonella 39
antidichotoma 47
bertheloti 60
cantabrigensis 42, 60, 61
compressa 51
depressa 50, 55
difformis 51, 53
elegans 61
Rhynchonella — contd.
gibsiana 60
globata 53
/ate 56
latissima 45
leightonensis 42, 59—61
lineolata var. mirabilis 47, 48
nuciformis 41
schloenbachi 55
tripartita 60
vespertilio 61
Rhynchonellacea 43
Rhynchonellidae 39, 43
Rhynchota 158
Robergiella 250
sagittalis 250
Saccoloma elegans 205
Saltatoria 158
Ensifera 166
Sampo 248
Scutellocladus 138
variabilis 137, 138, 139 ; PL 19, figs. 1-6
Sechuritella 1 79
Sedgwick Museum, Cambridge 45, 56, 61, 69, 80,
81, loi, 107, 121-123, I75> ^5, 188, 191, 193
Semifusus 259, 260, 265, 267
Sespedectes 25
Shrewsbury Museum 221
Sigmesalia 189
SIMONS, E. L. 1-36
Smilodectes 15, 17
SMITH, E. P. 294
Sowerbyella 115
Spathulopteris 141
Sphaerexochus hemicranium 86
Sphenopteridium 141
Sphenopteris 141
affinis 143
bifidum 143
cuneolata 142 ; PL 21, fig. 21
obfalcata 141 ; PL 20, figs. 16, 17
Sporae dispersae 143
Staurocephalus 71, 123
Stenopareia 120, 234
camladica 120, 234
sp. i 20
Stigmaria 140 ; PL 19, fig. 7
Stolmorhynchia 43
Sulcirhynchia 43, 59-61
Sulcus 42
Surcula 262, 264, 268, 279, 291
ingens 266
Surculites 257, 262, 264, 266, 267, 273, 279, 280,
289, 291
biluminatus 262
Tanganyika 199
Tarsiidae 24, 31
Tarsiiformes 5
INDEX
307
Tarsius 5, 17, 19, 23, 29
Tarsophlebia 161
Teilhardina 4, 7, 24, 32
belgica 4, 32
Telangium 143
sp. 142, 143 ; PI. 21, figs. 23-26
Tenebrio molitor 169
Terebratula antidichotoma 47
compressa 54
deformis 51
depressa 50
difformis 5 1
dilatata 53
dimidiata 51-53
gallina 5 1
/a/a 39, 45
latissima 39, 40, 43, 45
scaldisensis 57
Tetonius 7, 24, 27
Tetralichas melmerbiensis 120
Tettigonia 166
viridissima 165
Tettigoniidae 166
Thatcheria 257, 258, 259, 283, 292
carinata 283-286, 292, 293
gradata 258, 268, 276, 280, 281, 292, 294
liratula 282, 285, 290, 292-294
mirabilis 257, 258, 260, 264-267, 268, 269-272
275*, 276, 280, 281, 283, 284, 286, 288-290,
292-294
pagodula 282-286, 292, 293
vitiensis 257, 281-283, 285, 292, 294 ; PI. 47,
figs. 4-6
Thatcheriidae 261, 265, 267, 268, 277, 290
Thatcheriinae 257, 267, 290, 291
Thysanopeltidae 251
Toernquistia 118, 119, 129
nicholsoni 118, 119
reedi 118, 119 ; PI. 16, figs. 2, 3
translata 119
Tomiodendron ostrogianum 139
Torquesia 179, 182, 184, 187
faizai 175, 177, 178, 181*, 188 ; PI. 30, fig. 15
granulata 177-179, 181*, 182, 183, 189-191 ;
PL 30, figs. 10-14
granulosa 184
hassani 175, 177, 178, 181*, 188, 189 ; PI. 28,
fig. 2 ; PI. 29, fig. 9
tamra 175, 177, 178, 181*, 185 ; PI. 28, fig. i
vibrayeana 177, 178, 181*, 185-i87 ; PI. 31,
figs. 17-22
wagihi 175, 177, 178, 181*, 187 ; PI. 29, fig. 6
Tretaspidinae 85
Tretaspis 85, 130, 248
ceriodes 85
donsi 85 ; PI. 10, figs. 4, 6, 8
convergens 85 ; PI. 10, figs, i, 3, 5
granulata 220
kiaeri duftonensis 86 ; PI. 9, figs, i, 5, 7
radialis 86 ; PI. 9, figs. 2-4
Tretaspis — contd
seticornis 85
var. anderssoni 85
Triarthrinae 241
Triarthrus 241
becki 241
linnarssoni 241, 242 ; PI. 44, figs. 2, 5, 10, 12
sp. 241
Trichoptera 156
Tridactylidae 158
Trinucleidae 69, 79, 129
Trinucleus bucklandi 220
gibbifrons 79
goldfussi 79
nicholsoni 69, 79
ornatus 220
seticornis 79
Trochurus nodulosus 235
Tropaeum subarcticum 46
Tubicaulis 201
africanus 199, 200*, 201, 202*, 204*, 205*,
207-209 ; Pis. 33-36
berthieri 203, 206, 208
multiscalanformis 207, 208
scandens 206—209
solenites 201, 206—209
steward 207, 208
sutcliffii 207, 208
Turbinellidae 259
Turbo quadricarinatus 191
terebra 180
Turricula 262, 289, 291
javana 260
waitaraensis 260, 264
Turridae 257, 260, 261, 263-267, 277, 278, 288,
290, 291
Turritella 178, 180
ageri 175, 177, 178, 181*, 190, 191 ; PL 32,
fig- 25
cenomanensis 191
dzWeyi 177, 178, 180, 181* ; PL 29, figs. 7, 8
faizai 175, 177, 178, 181*, 188 ; PL 30, fig. 15
granulata 177—179, 181*, 182, 183, 184, 189-191 ;
PL 30, figs. 10-14
granulatoides 183, 184
hassani 175, 177, 178, 181*, 188, 189 ; PL 28,
fig. 2 ; PL 29, fig. 9
imbricataria 184
multistriata 191
rigida 186
sherborni 175, 177, 178, 181*, 190, 191 ; PL
30, fig. 1 6
tamra 175, 177, 178, 181*, 185 ; PL 28, fig. i
tolenasensis 187
turbinata 177, 178, 181*, 193, 194 ; PL 32,
figs. 26, 27
unicarinata 178, 180, 181*, 182 ; PL 28,
figs- 3-5
vibrayeana 177, 178, 181*, 185, 186, 187 ;
PL 31, figs. 17-22
308 INDEX
Turritella — contd. Wattsellahorderleyensis 121
wagihi 175, 177, 178, 181*, 187 ; PI. 29, fig. 6 WENNER-GREN FOUNDATION 34
Turritellidae 175, 179*, 180
Xancidac 259
asi ae 259 Xipheroceras dudressieri 155
Verrucostspontes sp. 146 ; PI. 22, fig. 57
257, 258, 260-262, 264, 268, 270, 275,
278-281, 288, 291, 292 Yorkshire Geological Society 69
generosa 261, 276, 280-282 Youngia trispinosa 87
liratula 257, 261, 275*, 276, 280-282, 294
pagodula 261, 280-282 Zaria 178
waitaraensis 280-282 ZEUNER, F. E. 153—171
WALTON, J. 135-152 Zeuneriasp. 166
, 7 Zonales 150
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