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Vi'Nv’i I i W
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Q) The Palaeontological Association, 1975
Cover: Marrolitlnis favus (Salter).
Reconstruction of Ordovician trinucicid trilobitc, prepared by Dr. J. K. Ingham as the symbol for the Symposium on
the Ordovician System, Birmingham, lt)74. Based on silicified material collected by Dr. R. Addison from limestones
of Upper Llandcilo age from Wales.
PALAEOECOLOGY OF A BITUMINOUS SHALE-
THE LOWER OXFORD CLAY OF
CENTRAL ENGLAND
by K. L. DUFF
Abstract. Quantitative palaeoecological studies, using triangular plots, rarefaction curves, trophic nuclei, trophic
group composition, and Diversity Index, have allowed the definition of ten different biofacies within the Lower
Oxford Clay (Upper Jurassic, Middle Callovian) of central England. Analysis of the distribution of these biofacies
and seven lithofacies groups, has led to the recognition of the Lower Oxford Clay as a deepening-water sequence,
in which two distinct environmental cycles are present. By comparison with other Mesozoic shale facies, the Lower
Oxford Clay appears different in having its fauna dominated by infaunal deposit-feeders and by high-level (‘pendent’)
epifaunal suspension-feeders; only the Upper Lias is comparable. Evolutionary changes are considered between
Palaeozoic and Mesozoic deposit-feeder dominated assemblages, with siphonate bivalves occupying most of the
niches previously held by articulate brachiopods.
Palaeoecological studies on clay sequences are of great importance in building
up an understanding of Mesozoic environmental conditions. Most thick clay sequences
seem homogeneous, but close inspection reveals many lithological and faunal varia-
tions capable of analysis. This is especially true of bituminous shale sequences, which
until the work of Hallam (1960, p. 10), were usually thought of as homogeneous.
Hallam showed that in the Blue Lias several lithologies, each with a characteristic
fauna, could be recognized; only some lacked benthonic fossils, which had previously
been considered a typical feature of bituminous shales. Since then other studies have
been made on similar rock sequences, and it is the purpose of this paper to show the
high variability, both in lithology and fauna, that exists within the Lower Oxford
Clay.
STRATIGRAPHY
The Callovian deposits of England represent a transgressive marine phase after the
lagoonal and estuarine conditions of the Bathonian, the base of the transgression
being marked by the Upper Cornbrash, which passes upwards into the Kellaways
Clay and Kellaways Rock. After the deposition of the Kellaways Rock, conditions
appear to have stabilized, with deposition of a thick argillaceous sequence of bitu-
minous shales, shaly clays, and more calcareous clays. The Yorkshire succession
(text-fig. 1) differs in that it is developed in a more marginal facies, and has been well
described by Wright (1968, p. 367): in this paper only the clay facies will be con-
sidered in detail. The Lower Oxford Clay has been defined by Callomon (1968,
p. 265), and occupies the whole of the Middle Callovian together with the top subzone
of the Lower Callovian and the lower part of the Upper Callovian (text-fig. 2). It
eonsists of about 16-25 m of grey bituminous shaly clays with other minor lithologies
developed within them (text-fig. 3). The biostratigraphy is further described else-
where (Duff, unpublished Ph.D. thesis, Leicester University, 1974). The Lower
Oxford Clay is of great importance for brickmaking, and is extensively quarried
[Palaeontology, Vol. 18, Part 3, 1975, pp. 443-482.]
444
PALAEONTOLOGY, VOLUME 18
TEXT-FIG. 1 . Comparison of the lower part of the Upper Jurassic of Yorkshire
and southern England, showing the development of a marginal facies in
Yorkshire during Callovian times. After Wright (1968). All zones drawn to
equal thickness, therefore the lithological sections are not to scale.
between Peterborough and Aylesbury. Higher parts of the Oxford Clay are more
plastic, less bituminous, and not so easily used for brickmaking; consequently
exposures are rarer.
The zonal divisions (text-fig. 2) have been considered by Callomon (1955, p. 254,
1964, 1968, p. 265), and are now widely accepted. Previous workers on the Oxford
Clay have dealt mainly with the general stratigraphy of the formation, and paid
scant attention to the Lower Oxford Clay itself (Woodward 1 895, p. 5 ; Morley Davies
1916; Neaverson 1925; Arkell 1933, p. 341), although Brinkmann (1929, p. 28) and
Callomon (1955, 1968) have dealt with that part of the formation in considerable
detail. The only palaeoecological studies have been by Rutten (1956) and Hudson
and Palframan (1969).
DUFF: OXFORD CLAY PALAEOECOLOG Y
445
STAGES
ZONES
SUBZONE
DIVISION
C. cordatum
z
<
Cardioceras
cordatum
C. costicardia
UPPER
o
QC
CC
tu
C. bukowskii
OXFORD
O
u.
O
u
Quenstedtoceras
C. praecordatum
CLAY
X
o
mariae
C. scarburgense
oc
Quenstedtoceras
lamberti
MIDDLE
Q.
a
D
Peitoceras
athleta
Upper
Middle
OXFORD
CLAY
Lower
K. (Zugokosmoceras)
Erymnoceras
grossouvrei
coronatum
K. (Zugokosmoceras)
LOWER
LU
obductum
z
O
o
K. (Gulielmites)
OXFORD
<
s
Kosmoceras
jason
>
jason
K. (Gulielmites)
CLAY
o
_I
medea
S. (Catasigaloceras)
o
enodatum
GC
Sigaloceras
calloviense
Sigaloceras
calloviense
KELLAWAYS
LU
$
Proplanulites
ROCK
o
koenigi
Macrocephalites
M. (Kamptokephalites)
kamptus
KELLAWAYS
CLAY
macrocephalus
M. (Macrocephalites)
UPPER
macrocephalus
CORNBRASH
TEXT-FIG. 2. The zonal sequence of the Callovian and Lower Oxfordian stages
of NW. Europe. After Callomon (1968).
NATURE OE THE FAUNA
The composition of the Lower Oxford Clay fauna has been considered elsewhere
(Callomon 1968, p. 269); molluscs are very dominant, mostly cephalopods and
bivalves. Other common macrofauna include gastropods, scaphopods, brachiopods,
Crustacea, annelids, and occasional echinoderms ; a diverse and well-preserved verte-
brate fauna has been described by several authors (Arkell 1933, p. 357). A faunal list
of the macro-invertebrates is as follows:
POLYCHAETA CEPHALOPODA
Genicularia vertebralis (J. de C. Sowerby) "Binatisphinctes' comptoni (Pratt)
Serpula sp. 'B.' fluctuosus (Pratt)
446
PALAEONTOLOGY, VOLUME 18
CEPHALOPODA {COnt.)
‘fi.’ spp.
Chojfatia spp.
Erymnoceras spp.
Hecticoceras spp.
Kosmoceras (Gulielmiceras) gulielmi (J. Sowerby)
K. (Kosmoceras) baylei Tintant
K. (K.) grossouvrei Douville
K. (K.) nodosum Callomon
K. (Spinikosmoceras) aculeatum (Eichwald)
K. (Sp.) acutistriatum Buckman
K. (Sp.) castor (Reinecke)
K. (Sp.) pollux (Reinecke)
K. (Zugokosmoceras) enodatum (Nikitin)
K. (Z.) Jason (Reinecke)
K. (Z.) medea Callomon
K. (Z.) obductum (Buckman)
K. (Z.) zugium (Buckman)
Pseudocadoceras spp.
Reineckeia spp.
Sigaloceras calloviense (J. Sowerby)
Belemnopsis sulcata (Miller)
Belemnoteuthis antiquus Pearce
Cylindroteuthis puzosianus (d’Orbigny)
BIVALVIA
Anisocardia (Anisocardia) tenera (J. de C. Sowerby)
Bositra buchii (Roemer)
"Entolium' sp. nov.
Camptonectes (Camptonectes) auritus (Schlotheim)
Chlamys (Cblamys) sp. nov.
C. (Radulopecten) fibrosa (J. Sowerby)
C. (R.) scarburgensis (Young and Bird)
Corbulomima macneillii (Morris)
Discomiltha lirata (Phillips)
Entolium (Entolium) corneolum (Young and Bird)
Grammatodon (Grammatodon) clathrata (Leckenby)
G. (G.) concinna (Phillips)
G. (G.) minima (Leckenby)
G. (G.) montaneyensis (de Loriol)
Gryphaea (Bilobissa) sp. nov.
Isocyprina (Isocyprina) roederi Arkell
MeleagrineUa braamburiensis (Phillips)
Mesosaccella morrisi (Desha yes)
Modiolus (Modiolus) bipartitus J. Sowerby
Myophorella (Myophorella) irregularis (Seebach)
Nanogyra nana (J. Sowerby)
Neocrassina (Neocrassina) sp. nov.
N. (N.) ungulata (Lycett)
Nuculoma sp. nov.
N. pollux (Raspail ex d’Orbigny)
Oxytoma (Oxytoma) inequivalvis (J. Sowerby)
Palaeonucula cottaldi (de Loriol)
Palaeonucula sp. nov.
Parainoceramus subtilis (Lahusen)
Pinna (Pinna) mitis Phillips
Plicatula (Plicatula) cf. fistulosa Morris and Lycett
Pleuromya alduini (Brongniart)
P. uniformis (J. Sowerby)
Protocardia (Protocardia) striatulum (J. de C.
Sowerby)
Protocardia sp.
Pteroperna? pygmaea (Dunker)
Rollierella minima (J. Sowerby)
Solemya woodwardiana Leckenby
Thracia (Thracia) depressa (J. de C. Sowerby)
Trautscholdia phillis (d’Orbigny)
GASTROPODA
Dicroloma bispinosa (Phillips)
D. trifida (Phillips)
Pleurotomaria reticulata (J. Sowerby)
' Procerithium' damonis (Lycett)
Spinigera spinosa d’Orbigny
SCAPHOPODA
Prodentalium calvertensis Palmer
BRACHIOPODA
Lingula craneae Davidson
'Orbiculoidea' sp.
' Rhynchonella' sp.
CRUSTACEA
Mecocheirus pearcei M’Coy
ECHINODERMATA
Unidentified ophiuroids
The microfauna is more restricted than that of the Middle and Upper Oxford Clays,
the foraminifera having been studied by Cordey (1962, 1963) and Barnard (1952,
1953), the ostracodes by Whatley (1970), and the coccoliths by Rood, Hay and
Barnard (1971) and Rood and Barnard (1972). Apparently, the darker, more organic
rich shaly clays of the Middle Callovian were less conducive to the development of
a diverse benthonic microfauna than were the more calcareous clays of the Upper
Callovian-Lower Oxfordian.
This paper is based on detailed studies made on the Lower Oxford Clay in 1970-
DUFF: OXFORD CLAY PALAEOECOLOG Y
447
1971 at four brickpits in central England (text-fig. 4), collections coming from beds
of Calloviense-Coronatum Zone age. The beds are well exposed in continuously
accessible profiles, and are clearly marked off at the base by the sandy Kellaways
Rock, and at the top by a concretionary limestone bed, the Acutistriatum Band (text-
fig. 3); this marker horizon was shown by Callomon (1968, p. 272) to be the basal bed
of the Athleta Zone. The palaeoautecology of the bivalves is summarized in Table 1.
TABLE 1. Life habits of the bivalve genera recognized in the Lower Oxford Clay of southern England.
Epifaunal Infaunal
Genera and feeding
Taxonomic
groups
*u
CL)
■c
•D
X _
o
o
Vi
c
c
(J
Maximum s
length (mm
Swimming
Free-living >
cemented
Vi
Vi
CQ
‘Pendent’
o
-c:
.9r
'55
C
O
Z
o
O
s:
c/5
o
.9"
C
o
h-J
-D
2
CO
72
U
s
Vi
Vi
ffl
Mobile
Deposit-feeders
Superfamily
Palaeonucula
18-4
X
NUCULACEA
Mesosaccella
17-6
X
NUCULANACEA
Suspension-feeders
Solemya
380
X
SOLEMYACEA
Grammatodon
28-0
X
X
ARCACEA
Modiolus
700
X
MYTILACEA
Pinna
82-4
X
PINNACEA
Pteroperna
15-3
X
PTERIACEA
Parainoceramus
72-5
X
PTERIACEA
Bositra
15-0
X
PECTINACEA
Oxytoma
39-8
X
PECTINACEA
Meleagrinella
33'7
X
PECTINACEA
Entolium
310
X
PECTINACEA
'Entolium' gen. nov.
12-3
X
PECTINACEA
Camptonectes
580
X
PECTINACEA
Chlamvs
9-7
X
PECTINACEA
Radulopecten
760
X
PECTINACEA
Plicatula
26-8
X
PECTINACEA
Gryphaea
800
X
OSTREACEA
Nanogyra
11-2
X
OSTREACEA
Myophorella
880
X
X
TRIGONACEA
Discomiltha
47-0
X
X
LUCINACEA
Neocrassina
21-2
X
X
ASTARTACEA
Trautscholdia
12-5
X
X
ASTARTACEA
Protocardia
30-3
X
X
CARDIACEA
Anisocardia
25-0
X
X
ARCTICACEA
Isocyprina
20-0
X
X
ARCTICACEA
Rollierella
24-0
X
X
ARCTICACEA
Corbulomima
6-8
X
X
MYACEA
Pleuromya
71-0
X
PHOLADOMYACEA
Thracia
650
X
X
PANDORACEA
448
PALAEONTOLOGY, VOLUME 18
•3
I/I
(T LiJ
> Z
8S
to CD
^ 3
O to
cc
o
tri to
o
O N
to CD
< Z|
Q IS] I
Ltl ffl ,
2 3I
(/) I
TEXT-FIG. 3. Lithological sections measured at the four quarries examined in central England.
DUFF: OXFORD CLAY PALAEOECOLOG Y
449
The life habits of the benthonic invertebrates other than bivalves which occur in
the Lower Oxford Clay are as follows :
Suspension-feeders : Genicularia vertehralis (epifaunal), Serpula sp. (epifaunal), "Orbiculoidea sp. (epifaunal),
'Rhynchonella sp. (epifaunal), Lingula craneae (infaunal).
Deposit-feeders: Procerithium damonis (epifaunal), Dicroloma bispinosa (infaunal), Dicroloma trifida
(infaunal), Spinigera spinosa (infaunal), Prodentalium calvertensis (infaunal).
Scavengers: Mecocheirus pearcei, ophiuroids.
Browsing herbivores: Pleurotomaria reticulata.
The methods of analysis used here are a combination of those introduced by both
zoologists and palaeontologists, and have not previously been applied to Mesozoic
clay deposits. Thus there is a lack of comparative data, and the Lower Oxford Clay
has been compared only qualitatively with other Mesozoic sediments, especially clays.
TEXT-FIG. 4. The outcrop of the Oxford Clay in Britain, showing the location of the major sections
examined.
Preservation. The Lower Oxford Clay is notable for the preservation of original shell
aragonite in the shales, especially in the bituminous shales; a feature caused by the
impervious nature of the sediment (Hudson and Palframan 1969, p. 398). However,
most of the material is crushed.
450
PALAEONTOLOGY, VOLUME 18
In some of the more porous lithologies, notably the shell beds, there has been
replacement of the original aragonite by secondary calcite, precipitated from cal-
careous fluids moving through the rock, or by recrystallization of shell material in
situ. A more notable post-depositional preservational change has been the growth
of pyrite in many of the shell beds, and in local pockets within the shales. It appears
that the porous shell beds have acted as ‘aquifers’ along which sulphide-rich fluids
moved, and that when the pyrite was precipitated it became concentrated in the
central parts of the shell beds. This is particularly noticeable in many of the Nuculacean
shell beds, where the central part of the shell bed is strongly pyritized, with pyritiza-
tion decreasing towards the margins. Many pyritized shells have had the shell material
replaced by pyrite, rather than having had pyrite grow outward from the shell surface.
Within the bituminous shales, many shells have developed pyrite overgrowths. The
pyrite is usually rather patchily developed and often seems to be concentrated on
aragonitic shells such as Thracia, Pinna, and Palaeonucula where it occurs as small
patches on the shell surfaces. Hudson and Palframan (1969, p. 404) describe a com-
parable situation in the Middle and Upper Oxford Clay of Woodham, Bucks.,
where pyrite is patchily developed on the surfaces of bivalves preserved as clay moulds.
They attribute the pyrite formation to local sulphate reduction by bacteria acting on
the organic matrix of the dissolving shell. It is possible that the patchy pyrite developed
on aragonitic shells in the Lower Oxford Clay formed in a similar manner, although
the aragonite has not totally disappeared.
Another characteristic feature of the Lower Oxford Clay is the presence of con-
cretionary limestones at certain levels (text-fig. 3). Dr. J. D. Hudson informs me of
the existence of two phases of concretion development, one pre-compaction and the
other post-compaction, each with distinctive carbon and oxygen isotopic com-
positions. The early pre-compaction concretions are septarian, and are found within
the various Lower Oxford Clay bituminous shales and shell beds (text-fig. 3); they
contain uncrushed fossils preserved in partially dissolved aragonite or secondary
calcite. The later, post-compaction concretions occur as lenticular limestones within
the Acutistriatum Band (text-fig. 3), and contain crushed fossils, usually preserved
in secondary calcite. Both limestones are of diagenetic origin, and contain the same
fauna as the enveloping shales.
Methods of analysis. At the four pits studied (text-fig. 4), the Lower Oxford Clay is
up to 18 m thick, and worked in large open-cast pits by means of draglines, giving
sloping faces for collections on which a continuous profile may also be measured.
As the sections were measured, detailed counts were made of all the fossils found in
each bed, collections being made over a horizontal distance of up to 2 m, and con-
tinuing until no new species appeared in the sample. In practice, collection ceased
after about 2000 specimens had been counted, and when all the dominant species
had appeared. In the case of beds over 50 cm thick, each 50 cm was then considered
as a separate sample; this gave a method for evaluating the faunal similarity of dif-
ferent parts of the thicker units. In addition to these field counts (usually conducted
on up to 2000 specimens), blocks from each sampled bed were taken back to the
laboratory and broken up under more controlled conditions to check the accuracy
of the field counts. While this sometimes revealed the presence of one or two
DUFF: OXFORD CLAY PALAEOECOLOGY 451
TEXT-FIG. 5. Distribution of the ten biofacies at each of the four Midlands quarries.
452
PALAEONTOLOGY, VOLUME 18
additional species (in small quantities), in most cases it merely confirmed the field
counts, and so the analysis presented here is based on the field data only.
Analyses of the organic carbon and insoluble residue percentages of the samples
are shown in Tables 2 and 3. The organic carbon content was determined volumetric-
ally, the clay samples being treated with a solution of potassium chromate in phos-
phoric acid, and gives a measure of the amount of detrital organic matter available
TABLE 2. Organic carbon contents measured in the various Lower Oxford Clay biofacies.
Biofacies
N
Max
Min
Mean
Silts and silty clays
1
10
10
10
Deposit-feeder bituminous shales
10
3-5
2-1
2-9
Gramniatodon-rich bituminous shales
2
61
3-4
4-8
Foram-rich bituminous shales
3
4-9
3-5
41
Nuculacean shell beds
1
1-8
1-8
1-8
Gryphaea shell beds
1
L7
1-7
L7
Meleagrinella shell beds
4
3-8
2-3
2-9
Calcareous clays
1
M
11
11
N = number of samples; Max = maximum observed value; Min = minimum observed value,
for collection by feeding organisms. The insoluble residue determinations give
a measure of the amount of lime present in the sediment, either as cement or as
original particles of shell or other carbonate. Organic carbon contents of over 3%
in Recent mud deposits have been shown by Bader (1954, p. 40) to cause a diminution
in bivalve diversity, with infaunal deposit-feeding protobranchs becoming dominant.
The organic carbon contents of many of the Lower Oxford Clay samples (Table 2)
show that they belong to this type of lithology, with an impoverished benthonic fauna.
The degree of correlation between the organic carbon contents of Recent muds and
fossil shales is uncertain, and it is not possible to tell whether or not the values
obtained from compacted rocks are true reflections of the primary organic carbon
content. Comparison of Bader’s values with those from the Lower Oxford Clay
suggests that in some cases, they are.
Most of the analysis was carried out on the benthonic fauna only, with the nektonic
elements (such as the cephalopods) removed from consideration. However, before
the nektonic elements were deducted, the relative percentages of nektonic predators
and scavengers were calculated; in the bituminous shales they constitute 10-15% of
the fauna. Having removed the nektonic elements, the data for the remaining benthos
were recalculated to give percentages of epifaunal suspension, infaunal suspension,
and deposit-feeders only. The reasons for using feeding groups were discussed by
Rhoads ct a/. ( 1 972, p. 1 1 00), who suggested that it is sedimentary and hydrographical
conditions which most closely control the distribution of bivalves, with sediment
grain-size and texture, bottom turbidity, and food availability all being of importance
in determining the spatial distribution of suspension and deposit-feeding bivalves.
The status and mode of life of bivalve feeding groups in general have been studied by
Stanley (1970). The ecological positions of the various bivalve genera are shown in
Table 1.
The data for the bivalves was then further subdivided, because of the high pro-
portion of epifaunal suspension-feeders such as Bositra, Oxytoma, Meleagrinello,
Parainoceranms, and Pteroperna in some beds. It seems very likely that these genera
DUFF: OXFORD CLAY PALAEOECOLOG Y
453
were not strictly benthonic, but lived byssally attached to organic matter at some
distance above the sea floor, as there appears to be a lack of suitable benthonic attach-
ment areas, and the genera show a tendency to occur in clusters. It is suggested that
they were attached to algal fronds, probably not cemented to the sea floor, and which
could be moved by currents; attachment to floating driftwood is also likely, as this
material is characteristic of the bituminous shales, and is often associated with
clusters of Parainoceramus and Meleagrinella. There is also the possibility that
Bositra was pseudo-planktonic (Jefiferies and Minton 1965). This group of genera is
grouped together as ‘pendent’ epifaunal suspension-feeders, and as they tend to be
rather abundant in the bituminous shales, thereby obscuring the relative importance
of the more strictly benthonic elements, the bivalve percentages were recalculated to
omit them. Consideration of the whole bivalve assemblage then shows the over-all
faunal composition of a bed, the relative abundance of the strictly benthonic species
being seen after the removal of the pendent species. The abundance of driftwood,
frequently in large pieces, suggests that the bituminous shales were probably laid
down fairly near shore, in a quiet-water environment, and where a large amount of
suspended organic particles provided a rich food source for high-level suspension-
feeders.
BIOFACIES ANALYSIS
Ten biofacies have been recognized within the Lower Oxford Clay of the Midlands,
the major lithofacies groupings being subdivided by faunal content; the data are
summarized in Table 3, while text-fig. 5 shows the distribution of each of these facies
at the major pits. The data were then analysed in five ways to give a synthesis of the
palaeoecology, the plots used being (a) triangular plots, (h) rarefaction curves,
(c) trophic nuclei, (d) trophic group composition, (e) Diversity Index.
The triangular diagrams are based on bivalve feeding groups, the corners of the
triangles representing 100% epifaunal suspension-feeders (ES), 100% infaunal
suspension-feeders (IS), and 100% infaunal deposit-feeders (ID). Each sample has
both the total bivalve fauna and the over-all benthonic fauna (excluding pendent
genera) divided into these three groups, and may then be represented on the diagram
by a single point. It can be seen from text-figs. 6 and 7 that each biofacies yields
a group of points, all falling within a certain field of the triangle, with varying degrees
of overlap.
The rarefaction curve method was conceived by Sanders (1968) as a means of
comparing the diversities of different samples of benthonic organisms. He showed
that most diversity measurements were affected by sample size, as individuals are
added to a population at an arithmetic rate, while species are added at a decreasing
logarithmic rate. The rarefaction method depends on the shape of the species
abundance curve rather than the absolute number of specimens in a sample, and has
the advantage that each sample generates a curve. The method of calculating and
plotting rarefaction curves is described by Sanders (1968, p. 245). Each aquatic
environment was shown by Sanders to have its own characteristic rate of species
increment, with its rarefaction curves lying in a particular field. The curves generated
by the various Lower Oxford Clay biofacies (text-fig. 8) agree closely with those
454
PALAEONTOLOGY, VOLUME 18
TEXT-FIG. 6. Triangular plots of bivalve feeding
groups (ID, infaunal deposit; ES, epifaunal sus-
pension; IS, infaunal suspension) for three of the
biofacies, a, silts and silty clays, all bivalves; b, silts
and silty clays, pendent bivalves removed ; c, deposit-
feeder bituminous shales, all bivalves; d, deposit-
feeder bituminous shales, pendent bivalves removed ;
e, Grammatodon-nch bituminous shales, all bivalves ;
/, Grammatodon-r'ich bituminous shales, pendent
bivalves removed.
TEXT-FIG. 7. Triangular plots of bivalve feeding
groups for the remaining biofacies, a, foraminifera-
rich bituminous shales, all bivalves; /), Meleagrinella
shell beds, all bivalves (a); blocky claystone, all
bivalves (b); c, shell beds (apart from Meleagrinella
shell bed), all bivalves; Gryphaea shell beds (a),
Nuculacean shell beds (b), Grammatodon-nch. shell
beds (c); d, shell beds (apart from Meleagrinella
bed), pendent bivalves removed; Gryphaea shell
beds (a), Nuculacean shell beds (b), Grammatodon
shell beds (c); e, calcareous clays, all bivalves;
/, calcareous clays, pendent bivalves removed.
generated by Recent Boreal shallow-water samples, and Hallam (1969, p. 1 1) placed
England in his Boreal Province during Callovian times. However, the Boreal Province
of the Jurassic is not necessarily equivalent to the Recent Boreal area.
The trophic nucleus of an assemblage or community is defined as the numerically
dominant species which make up 80% of the fauna (Neyman 1967, p. 151). Analysis
of the trophic nucleus helps clarify the relationships between the various members of
the assemblage, notably in the relative abundance of the species, and the importance
of deposit-feeders. The trophic nucleus of most communities consists of up to five
species, although in some tropical marine environments high specific diversity
DUFF: OXFORD CLAY PAL AEOECOLOGY
455
TABLE 3. A summary of the main faunal and lithological characteristics of the ten Lower Oxford Clay
biofacies.
Biofacies Lithology Dominant faunal elements Organic Insoluble
carbon
7o
residue
%
Silts and silty clays
Silts and silty clays
Cephalopods, Pinna, Protocardia,
Trautscholdia, Corbulomima,
Meleagrinella
10
93
Deposit-feeder bituminous
shales
Dark olive-green shaly
clays
Pendent epifaunal suspension-
feeders (Bositra, Meleagrinella,
and Oxytoma), together with
Palaeonucula and Mesosaccella
2-9
90
Grammatodon-rkh bituminous
shales
Dark olive-green shaly
clays
Bositra, Oxytoma, Meleagrinella,
Palaeonucula, Mesosaccella,
together with infaunal
suspension-feeders such as
Grammatodon, Thracia, and
Isocyprina
4-8
90
Foram-rich bituminous shales
Light green, rather
fissile, shaly clays
Bositra, Parainoceramus,
Mesosaccella, Corbulomima,
Prodentalium, foraminifers
(Brotzenia)
41
87
Nuculacean shell beds
Shell concentrate
Palaeonucula, Mesosaccella
1-8
79
Grammatodon-rkh shell beds
Shell concentrate in
clay matrix
Grammatodon, Isocyprina, oysters,
Oxytoma, Discomiltha,
Protocardia, Trautscholdia,
Neocrassina, Myophorella
80
Gryphaea shell beds
Shell concentrate
Gryphaeate oysters, bone
fragments, cephalopods
1-7
82
Meleagrinella shell beds
Shell concentrate
Meleagrinella
2-9
69
Blocky claystone
Light grey plastic clay
with dark streaks
Bositra, Meleagrinella,
Palaeonucula, Mesosaccella,
Procerithium, Lingula, Solemya
96
Calcareous clays
Light grey or grey-
green, rather plastic,
clay
Palaeonucula, Mesosaccella,
Discomiltha, Isocyprina,
Myophorella, ^ Entolium' ,
Genicularia, oysters
11
74
greatly increases its size. Table 4 shows the over-all trophic nucleus of the whole
benthonic fauna, whilst Table 5 shows the trophic nucleus discounting the pendent
bivalves. Columns 7-9 of each table show the percentage of the various kinds of
deposit-feeders within each biofacies.
As well as determining the size of the trophic nucleus, it is useful to examine the
trophic group composition of each biofacies, a technique introduced, and later
refined, by Turpaeva (1948). This method of determining the trophic relationships
of all the benthonic invertebrates in an assemblage was shown by Walker (1972,
p. 83) to be a useful method of ecological analysis. Turpaeva’s work on the Recent
faunas of the Barents Sea revealed several generalizations which apply to most
Recent communities; Walker also showed that they could be applied to many Lower
Palaeozoic communities, and the evidence from the Lower Oxford Clay suggests
that they are also applicable to Mesozoic assemblages. Turpaeva chiefly showed that
(a) each community is dominated by a single trophic group, (b) each of the dominant
species in the trophic nucleus belongs to a different trophic group, (c) one species
456
PALAEONTOLOGY, VOLUME 18
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TEXT-FIG. 14. Trophic group composition of the Nuculacean
shell bed biofacies. All benthos included.
triangular plots (text-fig. 7c, d) show the relationship between this and the preceding
biofacies, indicating that there is some overlap. However, the two biofacies are easily
distinguished by their faunal content and the surrounding lithology. This biofacies is
also likely to be caused by slower sedimentation, and slight increase in current
activity.
Gryphaea shell beds, like the two preceding types, are restricted to a particular
lithology, the transition beds between the Kellaways Rock and the Oxford Clay.
They, too, have a low dominance diversity, with Gryphaea 62-2% of the fauna (text-
fig. 16), the remaining 37-8% being fairly evenly distributed between twelve species
of suspension-feeder (28-7%) and three species of deposit-feeder (8-5%). Amongst
the dominant species, there is evidence of niche separation, although in general the
biofacies is characterized by suspension-feeders. The high oyster content places the
field of this biofacies close to the epifaunal suspension-feeder corner of the triangle
(text-fig. Id, e). Again, the fauna is similar to that of the beds in which the shell beds
occur, and the sediment seems to be another shell concentrate formed more or less
in situ. There is, however, evidence to suggest that the fossils found in this biofacies
may have been transported, as bivalves occur as disarticulated shells rather than
articulated shells, many of the valves are fragmented, and there are disproportionate
amounts of left and right valves. In addition, broken cephalopod fragments, reptile
bones, and fish teeth are fairly common, and it seems likely that the Gryphaea shell
beds represent a transported fossil assemblage. Facies associations (their occurrence
in the silts and silty clays), and consideration of the mode of life of the various faunal
DUFF: OXFORD CLAY PALAEOECOLOGY
469
Percent
0 10 20 30 40
1 I I I i_
50 60
-I 1
Mesosaccella
morrisi
Palaeonucula
sp nov
3 a
Procerithium
damonis
Bositra buchii
Grammatodon
minima
Meleagrinella
braamburiensis
Corbulomima
macneillii
Flat oyster
Trautscholdia
phillis
□ LF
Entolium
corneolum
r 50 mm
- 25 mm
•- 0
Thracia
TEXT-FIG. 15. Trophic group composition of the Grammatodon
shell bed biofacies. All benthos included.
elements, particularly the oysters, suggest that these were deposited in shallower
water than the bulk of the Lower Oxford Clay, subject to wave-scouring at times.
Meleagrinella shell beds are largely confined to the upper part of the Grossouvrei
Subzone, where they are interbedded with calcareous clays, and consist over-
whelmingly of a concentration of broken and unbroken specimens of Meleagrinella
braamburiensis. Accurate counts are difficult, and consequently the data in text-
fig. 17 is less accurate than that of other biofacies. Over 70% of the fauna is Melea-
grinella, the remainder consists of well niche-partitioned species occupying several
habitats. Most of the shell beds are bounded above or below by burrowed surfaces,
where fragments of the overlying bed are piped down into the bed beneath. Burrowed
horizons are very rare elsewhere in the Lower Oxford Clay (except in the silts), and
these may represent phases of slow or nil-deposition, with increased current activity.
470
PALAEONTOLOGY, VOLUME 18
Percent
0 10 20 30 40 50 60
1 1 1 I I I I
LF
Meleagrinella
braamburiensis
HF
Procerithium
damonis
Corbulomima
macneillii
© EZD LF
Grammatodon
concinna
Palaeonucula
sp nov
Discomiltha
1 1 rata
r 50 mm
- 25 mm
L 0
Anomalodesmatan LF
sp. A
TEXT-FIG. 16. Trophic group composition of the Gryphaea shell
bed biofacies. All benthos included.
Others
The overwhelming abundance of Meleagrinella is probably original, as the shells are
too fragile to have withstood much post-mortem transport, and if the shell bed were
current-concentrated, there would be more larger, heavier shells. To explain the
numerous Meleagrinella, bearing in mind the inferred pendent mode of life of the
genus, it is probable that there was a large amount of floating organic matter to which
Meleagrinella may have been attached; the resulting environmental reconstruction
may be comparable with the Recent S'^rga^^wm-dominated environments. However,
this comparison is tentative, as most of the epifauna of Sargassum is soft bodied
(Friedrich 1965, p. 198), and would leave no traces, and there is no direct evidence
for large amounts of floating organic material in the Lower Oxford Clay, and its
existence is deduced from the abundant thin-shelled byssally attached Pteriacea and
Pectinacea in sediments such as bituminous shales, where the substrate would have
been too soft to allow such animals to live on the sea floor.
DUFF: OXFORD CLAY PALAEOECOLOGY
471
Percent
0 10 20 30 40 50 60 70
■ ■ ■ I I 1 I 1
Others [ 1 TEXT-FIG. 17. Trophic group composition of the Me/eagr/«e//a
LJ shell bed biofacies. All benthos included.
Blocky clay stone. This lithology is known only from the Jason Subzone (Bed 3B)
of Calvert. As with the deposit-feeder bituminous shales (within which it occurs), the
first two dominance positions are occupied by pendent bivalves (58-4%), whilst the
remainder of the fauna is dominated by an alternation of infaunal deposit and
suspension-feeders, Lingula and Solemya being the characteristic members of the
latter group (text-fig. 18). The total number of species found is small (only eleven),
but there is a relatively high dominance diversity, with only two species less than 1%
of the fauna. The lithology is a distinctive light grey, rather plastic, non-fissile clay,
with many black organic fragments spread throughout it. Many of the Lingula are
preserved upright in life position, unknown elsewhere in the Oxford Clay, which
suggests that there was no deep scouring of the sea floor. The over-all trophic nucleus
(Table 4) is similar to the normal bituminous shales, due mainly to the large content
of pendent species, and it is not until these are removed (Table 5) that the unusual
nature of the fauna becomes apparent.
Calcareous clays. This facies is restricted to the upper Grossouvrei Subzone, where it
forms a regular alternation with the Meleagrinella shell beds with the junctions
between the two always burrowed. The main characteristic of the calcareous clays is
their high dominance diversity, with five species each more than 10% of the fauna
(text-fig. 19), and a relatively large trophic nucleus (Table 4). The most abundant
species is an infaunal deposit-feeder (Mesosaccella), the next two species being
pendent bivalves {'Entoliuni and Meleagrinella), which together make up 60-8% of
the fauna. 'Entolium' (a new genus to be described elsewhere) is the most charac-
teristic faunal element, known only rarely from other facies; the polychaet worm
Genicular ia is also typical. Further analysis reveals that there is an approximately
equal distribution of deposit and suspension-feeders (Table 4), with most of the
suspension-feeders being pendent; infaunal suspension-feeders are not abundant
(text-fig. 19), and the degree of aeration within the sediment was probably not great,
in the same way as in the more calcareous Middle and Upper Oxford Clays. The
low content of infaunal suspension-feeders is also apparent from the triangular plots
472
Meleagrinella
braamburiensis
Bositra buchii
Palaeonucula
sp nov
Lingula craneae
Mesosaccella
morrisi
Solemya
woodwardiana
Dicroloma
trifida
Oxytoma
inequivalvis
Procerithium
damonis
PALAEONTOLOGY, VOLUME 18
Percent
0 10 20 30
1 I I I
40
50
I
60
HF
Others
- 50 mm
- 25 mm
- 0
TEXT-FIG. 18. Trophic group composition of the blocky
claystone biofacies. All benthos included.
(text-fig. le-f). The calcareous clays are rather rich in carbonate, and consequently
have a low insoluble residue content (74%), together with a low organic carbon
content (M%).
Relations between the hiofaeies. The over-all succession in the Lower Oxford Clay
at the major pits examined in the Midlands (text-fig. 5) is generally similar, although
there are local variations, especially at Stewartby and Calvert. Superimposed on these
local variations in the details of the succession is a marked southward increase in
thickness, the Enodatum-Grossouvrei Subzone succession being 12 m thick at
Peterborough, and gradually increasing through 14 m at Stewartby, 17 m at Bletchley,
to 18 m at Calvert. These thickness changes probably indicate greater distance from
the shoreline or a submarine swell, with consequent decrease in the number and thick-
ness of shell beds. This is particularly noticeable in the case of the shell bed which
occurs at or just above the base of the Obductum Subzone at Peterborough and
Stewartby; the shell bed is missing from the Bletchley section. Most of the thickness
variation occurs within the Jason and Calloviense Zones (text-fig. 6), which thicken
from 0-5 m at Peterborough to over 3 0 m at Calvert, suggesting that during the
deposition of the initial subzones of the Oxford Clay, conditions were more variable,
and controlled by local configurations of the sea floor. By the time of the Coronatum
DUFF: OXFORD CLAY PALAEOECOLOGY
473
Percent
10
I
20
I
30
—I
40
_l_
50
1
60
_j
Mesosaccella mornsi
5w
Entolium sp.nov
Meleagrinella
braambunensis
Procerithium
damonis
1 HP
HP
.
Co.
Bositra buchii
Palaeonucula
sp nov.
]hf
iiiliiiii Sw.
Corbulomima
macneillii
Genicularia
vertebralis
© CZI LP
■ 50 mm
- 25 mm
TEXT-FIG. 19. Trophic group composition of the calcareous
clay biofacies. All benthos included.
Zone, conditions appear to have stabilized over the whole of the Midland area,
giving a much more uniform thickness of Lower Oxford Clay.
The transition beds between the Kellaways Rock and the Oxford Clay are best
developed at Peterborough and Bletchley, where they consist of an alternation of
silts and silty clays, Gryphaea shell beds, and deposit-feeder bituminous shales. At
Peterborough the transition beds occupy the whole of the Enodatum and Medea
Subzones (0-5 m), but at Bletchley are restricted to the Medea Subzone (2 0 m),
Kellaways Rock deposition having continued until the top of the Enodatum Subzone.
At Stewartby the transition beds appear to be absent, a bipartite shell bed, the upper
part a Grammatodon shell bed and the lower part a Gryphaea shell bed, rests directly
on the silts of the Kellaways Rock, which are dated as Enodatum Subzone; deposit-
feeder bituminous shales of the Medea Subzone follow these shell beds directly.
Thus Kellaways Rock deposition ended earlier in the northern parts of the Midlands,
and Oxford Clay did not reach the south Midlands until the end of Enodatum Sub-
zone times; this suggests that the shoreline lay to the south during this time. There
then appears to have been rather variable current activity, phases of shallowing and
increased current activity giving Gryphaea shell beds, whilst in the intermittent quiet
phases silts or bituminous shales were laid down. The bituminous shales indicate that
474
PALAEONTOLOGY, VOLUME 18
the influence of the olfshore clay facies was greater at that time, and that distance
from shore, and probably water depth, was gradually increasing. The transition beds
are not exposed at Calvert, the pit ending in the Jason Subzone, although Callomon
(1968, p. 287) records 10 ft of combined Medea and Enodatum Subzones in a bore-
hole there.
Gradual recession of the shoreline during deposition of the transition beds eventu-
ally allowed the establishment of quiet water conditions in which bituminous shales
were laid down. At Peterborough, Bletchley, and Calvert, this phase seems to have
begun more or less at the start of Jason Subzone times, while at Stewartby similar
conditions became established slightly earlier. At all the pits the lowermost few
centimetres of the shales are markedly fossiliferous and contain the same species as
the underlying shell beds, indicating the gradual dying out of the fauna of the pre-
ceding bed. There then followed thick dominantly deposit-feeder bituminous shales,
which occupy the whole of Jason and Obductum Subzone time at Peterborough,
Bletchley, and Calvert, with the exception of the 1-m band of blocky claystone near
the base at Calvert. During this time bottom conditions were quiet, and water
circulation probably poor, producing an impoverished benthonic fauna dominated
by deposit-feeding bivalves and gastropods, with rare benthonic suspension-feeders.
Living above the bottom and attached to postulated organic material were large
numbers of pendent epifaunal suspension-feeding bivalves, chiefly Bositra, Melea-
grinella, and Oxytoma. The suspension-feeder dominated fauna is unusual, and was
probably caused by a superabundance of the suspended food source, as discussed
above.
The general sequence seen in the Lower Oxford Clay of the Midlands, Kellaways
Rock through transition beds to bituminous shales and then to more fossiliferous
shales, adds substance to the suggestion of Hallam (1967a, p. 489) that bituminous
shales are often laid down in relatively shallow water. Hallam suggested that near the
base of transgressive sequences widespread bituminous shale deposition was charac-
teristic, and that it was followed by more fossiliferous, deeper-water clays and shales,
laid down as water circulation improved, and sediment oxygenation increased. The
Kellaways Beds-Upper Oxford Clay sequence agrees with this model, the bulk of
the Lower Oxford Clay representing the bituminous shale part of the cycle.
The presence of the 1-m band of blocky claystone near the base of the Jason Sub-
zone at Calvert indicates that conditions were slightly different there during much of
the Jason Subzone times, as the Lingula-hc\\ blocky claystone fauna continues for
most of the subzone. Both Lingula and Solemya have been considered by previous
authors to be genera tolerant of poorly aerated water, and this, together with the
low organic carbon content, the increased thickness, and reduced benthonic fauna,
suggests that in the south Midlands area this part of the sequence was laid down more
rapidly than the deposit-feeder bituminous shales and possibly in slightly deeper
water.
At Stewartby the Obductum Subzone sequence is different to that seen at the other
pits, a thin development of deposit-feeder bituminous shale, containing two very
well-developed Nuculacean shell beds, being followed by a very thick Meleagrinella
shell bed, and a development of foraminifera-rich bituminous shale. The combina-
tion of the two Nuculacean shell beds, being followed by a very thick Meleagrinella
DUFF: OXFORD CLAY PALAEOECOLOGY
475
shell bed, suggests that local current activity must have been increased at this time,
probably on a local swell. The foraminifera-rich bituminous shale above the Melea-
grinella shell bed shows a return to quieter and deeper water, with well-developed
niche partitioning within the benthonic fauna.
Throughout the Midlands the end of the Obductum Subzone coincides with a wide-
spread phase of slow deposition, marked by Nuculacean or Grammatodon shell beds.
Brinkmann (1929, p. 81) showed that at Peterborough the shell bed at the top of the
Obductum Subzone represents a significant pause in sedimentation, and it seems
likely that a similar situation prevailed over much of the Midlands. This phase of
condensed deposition is followed at all the pits by a similar succession in the
Grossouvrei Subzone. The well-aerated conditions of the Nuculacean and Gram-
matodon shell beds persisted into the next phase of bituminous-shale deposition,
giving a sequence of Grammatodon-nch bituminous shales, rich in infaunal suspension-
feeders such as Grammatodon, Isocyprina, and Thracia. The start of the Grossouvrei
Subzone sees the first appearance in the Oxford Clay of Grammatodon minima, and
also marks the arrival of abundant Isocyprina. This event can be recognized at the
same level in Dorset, and is useful for defining the base of the Grossouvrei Subzone.
Phases of increased current activity during this time are marked by Grammatodon
shell beds. As subsidence continued, and oxygenation of the water gradually decreased,
the Grammatodon-rich bituminous shales were replaced by foraminifera-rich
bituminous shales in which deposit-feeders became more abundant and infaunal
suspension-feeders fewer, although the conditions which resulted in the deposit-
feeder bituminous shales never became re-established.
Later, in Grossouvrei Subzone times, there was a renewed phase of shallowing,
producing the characteristic alternation of calcareous clays and Meleagrinella shell
beds, bringing the Middle Callovian to a close in southern England. As suggested
above, this sequence must have been characterized by periodic explosions in coloniza-
tion by organic matter, allowing dense Meleagrinella shell beds to accumulate, and
producing many small non-sequences. Aeration in the calcareous clays must have been
relatively good as they supported a diverse fauna of suspension-feeders, and there is
a sudden increase in the abundance of tube-building annelid worms. Grossouvrei
Subzone deposition was concluded over the whole area by a thick Nuculacean shell
bed— the Comptoni Bed— which again represents a phase of increased current
activity, and a pause in sedimentation. This part of the sequence is usually capped by
a eoncretionary diagenetic limestone, the Acutistriatum Band, which is developed
within a band of very bituminous foraminifera-rich shaly clay, and which represents
the basal bed of the Athleta Zone.
Thus within a relatively thin sequence of Lower Oxford Clay (12-18 m), occupying
just over two ammonite zones, there are two cycles of environmental conditions.
Firstly, there is the deepening sequence from the Kellaways Rock through the transi-
tion beds into the deposit-feeder bituminous shales of the Jason-Obductum Subzones,
with indications of shallowing towards the top, and secondly, the more balanced
cycle of the Grossouvrei Subzone, which shows distinct deepening and shallowing
phases, ending with a pronounced non-sequence.
c
476
PALAEONTOLOGY, VOLUME 18
COMPARISONS WITH OTHER FACIES
In view of the lack of comparable quantitative data from similar environments,
comparisons must be limited to more qualitative observations, largely gleaned from
the literature. In particular, the work of Melville (1956), Hallam (1960, 1967), Palmer
(1966, 1966fl, 1973), and Sellwood (1972) on the Lias, Hudson and Palframan (1969)
on the Middle and Upper Oxford Clay, Price (1879) on the Gault, and Scott (1970)
on the Lower Cretaceous of the United States, has been used for comparison of clay
faunas, and shows the Lower Oxford Clay to be unusual because of its very high
content of pendent epifaunal suspension-feeders and infaunal deposit-feeders. With
the exception of Scott, no quantitative data was given, fossils merely being recorded,
or being said to be rare, common, or occur, and thus direct comparison is difficult.
There is also the assessment of the relative importance of evolutionary and environ-
mental changes when the palaeoecology of different ages is being compared. However,
in this consideration all the rocks are of Jurassic or Lower Cretaceous age, and the
importance of evolutionary changes appears minor; there is little change in the over-
all faunal composition, although there is much variation in the relative importance
of species within it. Protobranch bivalves are known to be a slowly evolving group,
so evolutionary effects in this protobranch-dominated assemblage are likely to have
been small. In general terms it appears to be environmental conditions which have
exercised the greater control over benthonic assemblages during Jurassic and Lower
Cretaceous times.
The shales of the Oxford Clay have frequently been compared with the Lias, but
this study shows many differences between the two deposits. Hallam (1960, p. 12)
described the bituminous shales of the Blue Lias of Dorset and Glamorgan, and showed
them to have a very high organic carbon content (3-9-8-0%), and a fauna consisting
almost entirely of ammonites, fish scales, and bivalve spat, indicating that bottom
conditions must have been anaerobic. Most of the fossils are preserved in the marls
and limestone bands, which Hallam showed to be essentially of primary origin,
although he later (1964) amended his views, but similarities with the Lower Oxford
Clay are negligible, as in the Lias limestones there is a rich and varied fauna of normal
infaunal and epifaunal suspension-feeders, while deposit-feeders and pendent bivalves
are rare. It thus seems very likely that the limestone part of the Blue Lias rhythm was
much better aerated than the Lower Oxford Clay, as it contains a much more varied
fauna, including gastropods, brachiopods, and echinoderms.
Sellwood (1972) gives similar results on the Sinemurian-Pliensbachian Lias to
those of Hallam (1960, p. 10), in so far as the fauna is dominated by infaunal and
epifaunal suspension-feeders, with few deposit-feeders. Rocks of this age over most
of Britain are clearly less bituminous and more well aerated than the bituminous
shales of the Blue Lias, and contain many genera which are also characteristic of the
Lower Oxford Clay, but again there is a lack of abundant pendent bivalves and
deposit-feeders. The same is true of the Middle and Lower Lias described by Melville
(1956, p. 74) from the Stowell Park borehole in Gloucestershire, pendent bivalves
and deposit-feeders again not being abundant, the fauna being dominated by infaunal
and epifaunal suspension-feeders. Palmer’s (1973, p. 252) work on the upper parts of
the Lower Lias in Gloucestershire shows a faunal list rather similar to that of the
DUFF: OXFORD CLAY PALAEOECOLOGY
477
Lower Oxford Clay, but again pendent bivalves and deposit-feeders are neither
abundant nor widespread. The Middle Lias (Palmer 1966, 1966a; Hallam 1967)
shows similar conditions to have prevailed during deposition of the more sandy
shales.
The Upper Lias (Melville 1956; Hallam 1967) of Britain is probably the most
similar deposit to the Lower Oxford Clay, consisting of dark shales and shaly clays
with a sparse benthonic bivalve fauna, dominated by deposit-feeding Nuculaceans
such as Nuculana and 'Nucula\ often with a pendent bivalve fauna of Bositra radiata
and Inoceramus dubius. These shales, which are also rich in cephalopods and Pro-
cerithium, often contain local concentrations of comminuted fish debris, insect
remains, and Crustacea, emphasizing the similarity with the bituminous shales of the
Lower Oxford Clay. Quantitative work on the fauna of the Upper Lias shales would
be of interest for detailed comparisons with the Oxford Clay.
Hudson and Palframan (1969) have described the palaeoecology of part of the
Middle-Upper Oxford Clay of the Midlands, and shown that there are clear dif-
ferences between the fauna of this part of the Oxford Clay and the Lower Oxford
Clay. The dark grey, well-laminated bituminous shales have been replaced by light
grey, rather calcareous clays, often rich in fossils preserved as pyritic internal moulds,
with no preserved aragonite. The Spinosum Clays (Athleta-Lamberti Zones) have
a sparse benthonic fauna, dominated by shallow infaunal species (mostly deposit-
feeders), with the epifauna characteristically rich in Chlamys and Gryphaea; other
Pectinacea are locally common. Near the top, Gryphaea beds appear, alternating with
the normal clay facies, and having, as well as common Gryphaea lituola, suspension-
feeders dominating over deposit-feeders. However, these Gryphaea beds are not
equivalent to those of the Lower Oxford Clay; they merely consist of a concentration
of oysters (estimated at four per square foot) in slightly harder and more calcareous
clay, and do not represent phases of non-deposition, although there must have been
some slowing of sedimentation. The rest of the Spinosum Clays make up a sequence
of quiet water muds similar to, but not equivalent to, the bituminous shales of the
Lower Oxford Clay. The faunal density is also less than that of the Lower Oxford
Clay, and there are none of the abundant Bositra, Meleagrinella, or Oxytoma so
typical of the Middle Callovian. The abundance of Astarte s.l. in the Spinosum Clays
suggests that the bottom sediments must have been fairly well aerated.
The Mariae Clays (Mariae Zone) are darker and more organic-rich than the
Spinosum Clays, and have a different faunal composition. The benthonic fauna is
reduced in variety, and is mostly infaunal, with Dicroloma, Procerithium, and
Nuculacea abundant, and Pinna the only common suspension-feeder. This part of
the Upper Oxford Clay is much more similar to the Lower Oxford Clay in faunal
content, although the presence of pyritic ammonites is a notable difference and the
shales are not well laminated. It seems likely, however, that during at least part of the
Mariae Zone conditions were somewhat similar to those of the Lower Oxford Clay.
The fauna of the Gault Clay (Cretaceous, Albian) has been summarized by Casey
(1966, p. 102), but a more comprehensive faunal list was given by Price (1879, p. 60),
who charted the distribution of the bivalve fauna. Price recognized eighty-six bivalve
species in the English Gault, fourteen of which are deposit-feeders, almost all species of
"Nucula\ In spite of this apparently high diversity of deposit-feeding protobranchs.
478
PALAEONTOLOGY, VOLUME 18
they are not as abundant as they are in the Lower Oxford Clay, nor are there so
many siphonate forms. There are, however, large swarms of Inoceramus through-
out the Gault, and so in this respect there are close similarities with the Lower Oxford
Clay. The main difference is in the greater diversity of infaunal and epifaunal
suspension-feeders, especially deep burrowers. The Gault has long been divided into
two lithological parts, the Upper Gault, consisting of light-coloured rather calcareous
clay, while the Lower Gault is much darker, less calcareous, and is generally more
similar to the Lower Oxford Clay, although it is not bituminous. Casey (1966,
p. 105) records Inoceramus and Nuculacea (Nucula spp., Acila, and Mesosaccella)
as the most abundant bivalves of the Lower Gault, with infaunal suspension-feeders
becoming more abundant in the Upper Gault. Most of the molluscs preserved in the
Lower Gault retain the original unaltered shell aragonite, although the cephalopods
in particular, as well as some of the bivalves and gastropods, are usually pyritized.
As in the Lower Oxford Clay, preservation of aragonite is related to the very impervious
nature of the sediment.
Scott (1970) has described the palaeontology and palaeoecology of the Kiowa
Formation (Lower Cretaceous, Aptian-Cenomanian) of Kansas, and recognized
six lithofacies groupings, of which one, the dark-grey shale lithofacies, is comparable
with the Lower Oxford Clay. It is a dark grey, fissile, well-laminated shale, with
a general lack of small-scale sedimentary structures, and Scott believes the fossil
assemblages to represent ‘disturbed neighbourhood’ and mixed-fossil assemblages.
This lithofacies is characterized by the Nuciilana association, dominated by Nuculana,
Yoldia, Nucula (nuculaceans), Breviarca (Arcacea), Pholadomya, Turritella, Drepano-
cheilus, and Lingula, which constitute 18-84% of the fauna; most of the remainder
of the fauna is composed of a corbulid.
The other comparable lithofacies recognized by Scott is his shell conglomerate
facies, which corresponds closely to the Gryphaea shell beds of the Lower Oxford
Clay. Similarities include the high content of Gryphaea (51-100%), the common
occurrence of calcitic shells, and the laterally discontinuous nature of the shell beds.
There are, however, some sedimentary structures which suggest that the Kiowa shell
conglomerates were deposited in very shallow water, possibly by storm-generated
currents, and there is no direct evidence that this is the case for the Lower Oxford
Clay Gryphaea shell beds.
Evolutionary changes — comparison with Palaeozoic and Recent assemblages. Deposit-
feeder dominated assemblages occur in many argillaceous deposits, from the
Ordovician to the present. In general trophic composition the assemblages are
similar, but marked evolutionary changes have altered the structure of the younger
ones, showing the importance of evolutionary changes over a long period. The main
changes are in the composition of the suspension-feeding part of the fauna since the
Lower Palaeozoic; bivalves having taken the place of the articulate brachiopods, as
a result of the development of siphon formation (Stanley 1 968, p. 224). The suspension-
feeder groups present in the Lower Palaeozoic have also been replaced by more
highly evolved superfamilies, leaving only the slowly evolving deposit-feeding
Nuculoida as a conservative stock.
The most similar assemblages to those of the Lower Oxford Clay are the various
DUFF: OXFORD CLAY PAL AEOECOLOG Y
479
Lingula ‘communities’ described from the British and American Palaeozoic (Bretsky
1970, p. 61 ; Ziegler et al. 1968, p. 5; Craig 1955, p. 114). The data for these com-
munities have been replotted by Walker (1972, pp. 87, 88, 90) to show the trophic
structure of the assemblages, and it is clearly apparent that both the Ordovician and
the Lower Carboniferous Lingula assemblages (described by Bretsky and Craig
respectively), are dominated by deposit-feeders, and show well-developed niche
partitioning. As in parts of the deposit-feeder bituminous shale biofacies, Lingula is
an important constituent of the fauna, and occupies second place in the assemblage ;
the dominant species in both these Lower Palaeozoic assemblages is an infaunal
nuculoid. In the Ordovician Lingula community described by Bretsky, third and
fifth places are occupied by archaeogastropods, which functioned as epifaunal
browsing herbivores. These elements are absent from the equivalent Mesozoic
assemblages.
Ziegler et al. (1968) have described several ‘communities’ from the Silurian of the
Welsh borderlands, the Lingula community being particularly relevant here. The
commonest species is an epifaunal suspension-feeding brachiopod (Camarotoeclua),
with Lingula and Palaeoneilo (a nuculanid) oceupying the next two positions; the
epifaunal pterioid Pteronitella is also characteristic. This assemblage has a low
diversity (Diversity Index 6-2), but shows a wide range of feeding types, although it
differs from the other Palaeozoic Lingula associations in having a high content of
filter-feeders; Walker (1972, p. 91) has suggested that perhaps this is not a true
example of the Lingula assemblage. This Lingula community is usually developed in
a more marginal facies, consisting mainly of sandstones, and so epifaunal suspension-
feeders, such as Camarotoechia, are most abundant. Thus Lingula communities in
general may be of varying type, and developed in several lithologies; only shale
occurrences are of relevance here. The Diversity Index value for Ziegler’s Lingula
community (6-2) agrees well with the values from the Lower Oxford Clay (5-9-
6-7 in the bituminous shales), in contrast to the other brachiopod-dominated Silurian
communities, which have much higher diversities (D.I. 7-8-1 1-8) due to their low
dominance diversity.
In general terms, there are close similarities between Palaeozoic and Mesozoic
deposit-feeder dominated assemblages, notably in the abundance of infaunal deposit-
feeding bivalves and infaunal suspension-feeding Lingula. In the Mesozoic, many
of the niches originally occupied by brachiopods have been taken over by bivalves,
many of the new superfamilies still being extant. One of the major differences between
the Lower Oxford Clay and the Palaeozoic assemblages, is the lack of a rich fauna
of pendent epifaunal suspension-feeding bivalves in the Palaeozoic, although they
oecur sporadically in some of the later assemblages (i.e. the Posidonia Band of Craig
1955, p. 112).
Recent offshore soft-mud eommunities have been described by many authors, and
are broadly comparable with the Lower Oxford Clay assemblages, although there are
differences. The silt-clay facies occupying the central axis of Buzzards Bay, Massa-
ehusetts (Sanders 1960; Rhoads and Young 1970; and Rhoads 1973) is dominated
by deposit-feeders, both infaunal and epifaunal. Sanders identified the fauna as
belonging to the Nucula proxirna-Nephthys incisa community, with these two species
(the latter a polychaet) making up 76% by number of the specimens collected. The
480
PALAEONTOLOGY, VOLUME 18
trophic nucleus consists of three deposit-feeders, Nucula proxima being the most
abundant, and suspension-feeders are of minor importance; no pendent epifaunal
suspension-feeding bivalves are known. The same is true of other offshore mud com-
munities (Jones 1950, p. 308) which are usually dominated by deposit-feeding proto-
branchs and polychaets, usually with a conspicuous associated fauna of infaunal
suspension-feeding bivalves. In this respect, the similarities with the various Lower
Oxford Clay biofacies are many when the strictly benthonic fauna alone is considered,
but, again, there is a noticeable lack of pendent suspension-feeders. The only known
Recent assemblage with a high content of pendent epifaunal species is that found on
Sargassum weed (Friedrich 1965, p. 198), but molluscs are not of great importance,
only five species having been described from this habitat. Stanley (1972, p. 189) has
recorded that many Recent species of Pteria attach preferentially to alcyonarian sea-
whips, a method of obtaining stable fixation in an agitated environment. Recent
parallels for the mode of life postulated for the Oxford Clay pendent bivalves are not
known, and this may be accounted for by evolutionary effects. Alcyonarians did not
appear until the Jurassic, and gorgonaceans until the Cretaceous (Stanley 1972,
p. 190), so it is possible that the lack of abundant rooted organic material during the
Jurassic led to the colonization of floating or rooted organic material (including algae)
by species that needed to live above, rather than on, a soft-mud substrate. Thus the
Lower Oxford Clay bituminous shale assemblages were both of different structure
and occupied slightly different environments to Palaeozoic and Recent offshore mud
assemblages, a consequence of evolutionary, rather than environmental changes^
CONCLUSIONS
Hallam ( 1967a, p. 489) suggested that bituminous shales were relatively shallow-
water deposits laid down in quiet, but not invariably stagnant water below wave
base, in contrast to the deep-water ‘barred basin’ model postulated by earlier authors.
The evidence deduced from the Oxford Clay appears to support this hypothesis, as
there is a deepening sequence from the sands and silts of the Kellaways Rock, through
the laminated bituminous shales of the Lower Oxford Clay, into the more massive
calcareous clays of the Middle-Upper Oxford Clay. The many small-scale alterna-
tions of lithology within the Lower Oxford Clay indicate relatively shallow-water
deposition, where a slight change in water depth could have a marked effect on hydro-
graphic conditions.
Faunally, bituminous-shale sequences show variability through time, with
Palaeozoic black shales either lacking benthonic elements (the graptolitic shales),
or with a benthonic fauna dominated by deposit-feeding nuculoids and suspension-
feeding linguloids. At various times pendent or benthonic byssally attached bivalves
were fairly common, but were never as important as in the Mesozoic. In the European
Jurassic, bituminous shales are particularly important in the Lower Hettangian,
Lower Toarcian, and the Middle Callovian, and all tend to show a fauna consisting
mainly of nuculoids and pendent bivalves. Recent organic-rich mud communities
have rather more infaunal suspension-feeders and no pendent bivalves, but, again,
nuculoids are numerically dominant. The role of the infaunal deposit-feeding proto-
branchs seems to have persisted more or less unchanged since the Lower Palaeozoic,
DUFF. OXFORD CLAY PALAEOECOLOG Y
481
their mode of life (inhabiting quiet water muds in areas of environmental stability)
needing little adaptive change. The replacement of brachiopods by bivalves as the
dominant members of the epifauna after the Palaeozoic followed siphon formation
in the Bivalvia, and marks the main change in the composition of the faunas of
organic-rich shales since the Palaeozoic.
Acknowledgements. I would like to thank Dr. J. D. Hudson for help and advice given during the period of
this research, and Mrs. A. M. Bowden, also of Leicester University, who carried out the organic carbon
determinations. The work was largely financed by an N.E.R.C. research studentship.
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ROOD, A. p. and Barnard, t. 1972. On Jurassic coccoliths: Stephanolithion, Diadozygus and related genera.
Eclog. geol. Helv. 65, 327-342, pis. 1, 2.
HAY, w. w. and Barnard, t. 1971. Electron microscope studies of Oxford Clay coccoliths. Ibid. 64,
245-271, pis. 1-5.
rutten, m. g. 1956. Depositional environment of Oxford Clay at Woodham clay pit. Geologic Mijnb.
N.S. 18, 344-346.
SANDERS, H. L. 1960. Benthic studies in Buzzards Bay. III. The structure of the soft-bottom community.
Limnol. Oceanogr. 5, 138-153.
1968. Marine benthic diversity; a comparative study. Am. Nat. 102, 243-282.
SCHAFER, w. 1972. Ecology and Palaeoecology of Marine Environments. Ed. CRAIG, G. Y. Oliver & Boyd,
Edinburgh.
SCOTT, R. w. 1970. Paleoecology and paleontology of the Lower Cretaceous Kiowa Formation, Kansas.
Paleont. Contr. Univ. Kansas, Art. 52, 94 pp., 7 pis.
SELLWOOD, B. w. 1972. Regional environmental changes across a Lower Jurassic stage-boundary in Britain.
Palaeontology, 15, 125-157, pis. 28, 29.
STANLEY, s. M. 1968. Post-Paleozoic adaptive radiation of infaunal bivalve molluscs— a consequence of
mantle fusion and siphon formation. J. Paleont. 42, 214-229.
— 1970. Relation of shell-form to life habits in the Bivalvia (Mollusca). Mem. geol. Soc. Am. 125.
1972. Functional morphology and evolution of byssally attached bivalve mollusks. J. Paleont. 46,
165-212.
TURPAEVA, E. p. 1948. The feeding of some benthic invertebrates of the Barents Sea. Zool. Zh. Ukr. 27.
WALKER, K. R. 1972. Trophic analysis : a method for studying the function of ancient communities. J. Paleont.
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WHATLEY, R. c. 1970. Scottish Callovian and Oxfordian Ostracoda. Bull. Br. Mus. nat. Hist. Geol. 19,
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WOODWARD, H. B. 1895. The Jurassic rocks of Britain. V. The Middle and Upper Oolitic rocks of England.
Mem. geol. Surv. G.B.
WRIGHT, J. K. 1968. The stratigraphy of the Callovian rocks between Newtondale and the Scarborough
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K. L. DUFF
The Nature Conservancy Council
Foxhold House, Thornford Road
Crookham Common
Newbury, Berkshire, RG15 8EL
Original typescript submitted 1 October 1974
Revised typescript submitted 4 December 1974
MEGASPORES AND MASSULAE OF AZOLLA
PRISCA FROM THE OLIGOCENE OF THE
ISLE OF WIGHT
by K. FOWLER
Abstract. Scanning and transmission electron microscopy, ultra-thin sectioning and light microscopy are employed
in this investigation of Azolla prisca, which is placed in Section Trisepta sect. nov. of the genus. The columella and
structural modifications of the proximal megaspore wall of A. prisca are compared with other species, both fossil
and modern, and the phylogenetic interrelationship of these structures are discussed. The complex megaspore wall
reveals an exine and two-layered perine, the outer perine layer being further subdivided into four zones. The inner
perine layer thickens to form the proximal wall and associated labra. The wall structure of most Azolla species appears
to be of this same basic pattern. Massulae, found dispersed and attached to megaspore apparatuses, reveal funnel-
shaped cavities connecting the microspores to the exterior. Although previously unrecorded in Azolla such structure
is present in both fossil and modern species, and is thought to form an escape mechanism for spermatozoids. A list
of the better-known pre-Miocene Azolla species is presented, which includes stratigraphic range, and characteristics
expressed as a formula. The evolutionary trends in Azolla are briefly reviewed.
The genus Azolla belongs to the Salviniaceae, a family of heterosporous ferns.
Plants are free-floating, occurring in freshwater habitats mainly in warm temperate
to tropical zones. Some forty-eight species are recorded, six of which are extant. The
fossil record of Azolla, dating back to the late Cretaceous of North America (Hall
\969b), is based mainly on its highly distinctive reproductive structures, the mega-
spore apparatus, and massula. In modern species, sporocarps are borne in pairs
which arise from the ventral lobe of the first leaf of a branch. These paired sporocarps
may be male and female, termed microsporocarp and megasporocarp respectively,
or they may be of the same sex. The microsporocarp contains microsporangia hold-
ing a number of massulae; the megasporocarp contains a megasporangium within
which is a single megaspore apparatus. The latter consists of a megaspore with
an elaborate wall, and a unique complex swimming apparatus. The term
‘Schwimmaparrat’, first applied by Strasburger (1873), is a misnomer in that the
structure does not endow buoyancy, mature megaspores sinking following liberation
(Sculthorpe 1967). The massula is a frothy pseudocellular structure in which micro-
spores are embedded. The six modern species, together with their present geo-
graphical distribution are as follows (data from Mahabale 1963; Sculthorpe 1967;
and Svenson 1944);
A. filiculoides Lam. From Alaska to Guatamala in North America, and in Andean and southern South
America. Introduced into eastern U.S.A., Hawaii, and Europe. A. filiculoides var. ruhra(K. Br.) Strasburger,
originally described from Australia, has since been found scattered throughout America.
A. caroliniana Willd. Essentially warm temperate. In eastern U.S.A. from Massachusetts to Florida,
extending to the West Indies and Brazil. Introduced in western, central, and southern Europe.
A. pinnata R. Br. Australasia, Indomalaya, and Africa, including Madagascar. Introduced into southern
Europe (Sculthorpe 1967).
A. microphylla Kaulfuss. Mainly South America, especially lowlands of Brazil and British Guiana.
[Palaeontology, Vol. 18, Part 3, 1975, pp. 483-507, pis. 59-61.]
484 PALAEONTOLOGY, VOLUME 18
Scattered distribution in western South America and northward to central America, extending into California
and the West Indies.
A. mexicana Presl. Mainly Mexico, with scattered occurrences northward through the Pacific States to
British Columbia, and eastward to Lake Michigan. Extending southward in lowlands to French Guiana
and Bolivia.
A. nilotica Dene. African species with a less extensive latitudinal span than A. pinnata, occurring from
Senegal and Ethiopia south to northern Transvaal.
As early as 1847 Mettenius reported seven species of Azolla, together with an
already confused synonymy, and interpretation of the development of reproductive
structures had commenced (Meyen 1836; Griffith 1844). The most significant work
on morphology and development of reproductive structures in Azolla concentrates
on A. filiculoides (Strasburger 1873, 1889; Campbell 1893; Hannig 1911; Duncan
1940; and Bonnet 1957). A. caroliniana was studied by Berggren (1880) and Pfeiffer
(1907), A. pinnata by Rao (1935), and A. nilotica by Demalsy (1954). Initial develop-
mental stages of megasporangium and microsporangium are identical, until the
production of thirty-two spores embedded in a multinucleate mass derived from
a periplasmodial tapetum. Subsequent megasporangial development involves abor-
tion of all but a single spore, the degenerating nuclei becoming aggregated into three
large vacuoles which later form floats of the swimming apparatus above the develop-
ing megaspore. Thickening of the megaspore wall occurs with deposition of an
elaborate perine derived from surrounding cytoplasm. The hairs which then develop
on the perine surface are considered homologous with glochidia which form on the
massula. Glochidia serve as a means of attaching the massula to the megaspore
apparatus. During microsporangial development, sixty-four spores are produced,
eight to twelve of which become distributed toward the periphery of each of five to
eight vacuoles. Each vacuole develops into the pseudocellular massula which bears
glochidia at the surface. A float is regarded as the homologue of a massula.
Although the fundamental development and organization of reproductive strue-
tures in Azolla have been understood since 1889, little detailed comparative work of
taxonomic importance has been attempted on modern species. This seems especially
surprising in that probably no other group of embryophyte plants possess such
complex spores.
The genus ranges from the late Cretaceous to the present. Taxonomic diversity
within the genus, eoupled with short stratigraphic ranges of individual species,
renders Azolla a potentially valuable stratigraphic indicator. The stratigraphic ranges
and characteristic features of the more important, and best-known, Azolla species
from the late Cretaceous and Palaeogene are presented in Table 1. This list excludes
certain species considered synonymous with others, and those established on such
limited information as (i) only vegetative remains recorded, (ii) only massulae known,
and (hi) megaspore apparatus inadequately described. Certain species established
mainly on the basis of megaspore wall structure are included, but should be regarded
as temporary, awaiting further information.
Taxonomically, Azolla species are placed in six sections of the genus, based on
features of the megaspore apparatus and massulae. The essential characteristics of
each section are given below, together with a list of suggested members which includes
extant species, and those from the late Cretaceous and Palaeogene.
TABLE 1. Stratigraphic range and characteristic features of the reproductive structures of late Cretaceous
and Palaeogene species of Azolla.
Important literature related to each species is listed in the second column. Characteristics are expressed
as a formula in the third column. The formula is in three sections, referring to swimming apparatus, proximal
pole of megaspore, and massula. Swimming apparatus: prefixed by C for columella, considered the basic
component. Followed by columellar type (d dome— or cone-shaped; t triseptate), nature of floats
(F = smooth, pseudovacuolate ; f ^ hairy, or poorly known float-like structures), float number (N =
numerous) and number of float tiers (in brackets). Proximal pole: prefixed by P; c = collar; 1 labra, or
similar structures. Massula: prefixed by M ( + M = massulae found attached to megaspores). In brackets,
g = glochidia recorded; -g = eglochidiate; a =- anchor-shaped glochidia; d = anchor-shaped with distal
dilation ; s = simple hair-like or coiled glochidia. In some instances the presence of certain features is
indicated only as a possibility.
CRETACEOUS
PALAEOCENE
EOCENE
OLIGOCENE
MIOCENE
A. simplex
Hall (1969b)
Cd I/P / M (gad)
A. barbata
Snead (1969)
Cd FN(2)/P / +M (gs)
Hall & Bergad (1971)
A. extinota
Jain (1971)
Cd fN /P (?!)/ +M (-g)
A. geneseana
Hills & Weiner (1965)
Cdl /P / M (ga)
A. lauta
Snead (1969)
Cd fN /P / M
A. distincta
Snead (1969)
Cd FN(3)/ P(?l)/ +M (ga)
Hall & Bergad (1971)
A. sahapfi
Dijkstra (1961)
Cd fl8(3)/P (1)/ M (ga)
Snead (1969)
A, montana
Hall k Swanson (196R)
Cd f 15-20/P (1)/ +M (ga)
Jain & Hall (1969)
A. bulbosa
Snead (1969)
Cd fl8(3)/P / M
A. fragilis
Jain & Hall (1969)
Cd FN/P (1) +M
A. Stanley i
Jain & Hall (1969)
Cd FI5+/P /+M (ga)
A . ve lus
Jain & Hall (1969)
Cd FN/P (1) / +M (ga)
A. tesohiana
Florschutz (1945)
Cd F24(3)/P / +M (ga)
Dijkstra (1961)
A. intertrappea
Sahni & Rao (1943)
C? F3/P(c + 1)/ +M (gad)
Hall (1969a)
Trivedi & Verma (1971)
A. indica
Trivedi & Verma (1971)
C? F3/P(c + 1)/ M (gad)
A. primoBVa
Arnold (1955a)
Cd FI/P /M (gad)
Hills & Weiner (1965)
Hills S Gopal (1967)
Hall (1969a)
A. antiqua
Dorofeev (1959)
Ct F6-9(2)/P (c+1)/ M
A. prisoa
Reid & Chandler (1926)
Ct F9 (2)/P (c+1)/ +M (gad)
■
A. nana
Dorofeev (1959)
Ct F9 (2)/P (c+1)/ M (-g)
A. turgaioa
Dorofeev (1959)
Ct F9 (2)/P (c+1)/ M (-g)
A. sibirioa
Dorofeev (1959)
Ct F9 (2)/P (c+1)/ M
A. ventrioosa
Nikitin (1955)
Ct F9 (21/P (c+1)/ M f-a)
Dorofeev (1959)
A, YiiM.'itvYi'i'i
Dorofeev (1955)
Ct F9 (2)/P (c+1) / M
A. aspera
Dorofeev (1963)
Ct F9 (2)/P (c+1) / M
•
486
PALAEONTOLOGY, VOLUME 18
Section simplicispora Hall, 1970. Float-like columella or single float; anchor-shaped glochidia. Fossil
species A. geneseana, A. primaeva, A. simplex.
Section kremastospora Jain and Hall, 1969. Megaspore apparatus with more than nine floats; anchor-
shaped glochidia. Fossil species A. distincia, A. montana, A. schopfi, A. stanleyi, A. tescliiana, A. velus.
Section filifera Hall, 1 968. Megaspore apparatus with more than nine floats ; hair-like or coiled glochidia.
Fossil species A. harbata.
Section antiqua Dorofeev, 1959. Megaspore apparatus with six to nine floats in two tiers; massulae
undescribed. Fossil species A. antiqua, A. aspera, A. nikitinii, A. sibirica.
Section rhizosperma Meyen, 1836. Megaspore apparatus with nine floats in two tiers; glochidia absent,
or simple, straight or branched structures. Fossil species A. nana, A. turgaica, A. ventricosa. Modern
species A. nilotica, A. pinnata.
Section azolla Meyen, 1836. Megaspore apparatus with three large floats; glochidia simple, hooked
or anchor-shaped. Fossil species A. indica, A. intertrappea. Modern species A. caroliniana, A. filiculoides,
A. mexicana, A. microphylla.
Important contributions to our knowledge of the morphology of the megaspore
apparatus and massula of fossil species of Azolla and their phylogenetic significance,
have been made by Hills and Gopal (1967), Hall and Swanson (1968), Jain and Hall
(1969), and Jain (1971). In some instances lack of suitable material has caused mis-
interpretation of structure. This has resulted in some terminological confusion,
especially with regard to the application of the term columella. Apart from variation
in form of the glochidia, little has been written concerning the structure of the massula.
It now seems clear that the multifloated swimming apparatus is more primitive than
both the nine-floated and three-floated type. Massulae with anchor-shaped glochidia
appear to be more primitive than those with hair-like glochidia, or those in which the
glochidia are absent. Although the megaspore wall structure of several fossil species
has now been examined, comparatively few have received detailed attention. The
most important of such contributions is that of Kempf (1969a and h) who examined,
by means of light and transmission electron microscopy, the megaspore wall of
A. tescliiana, A. nana, A. cf. aspera, A. tomentosa, and A. tegeliensis. Structural
details of the megaspore wall of modern species of Azolla are equally lacking, with
the exception of A. filiculoides (Bonnet 1957), A. nilotica (Demalsy 1954), and
A. pinnata (Kao 1935 ; Sweet and Hills 1971), and surprisingly little progress has been
made in tracing the developmental history of the wall layers. Some confusion has
resulted from the number of different terms which have been applied to these wall
layers in both fossil and modern species. Structural variation of the megaspore wall
between fossil Azolla species has been demonstrated, and attention drawn to this
potentially useful method of identifying wall fragments (Kempf 1969a; Snead 1969,
1970; Hall and Bergad 1971).
A. prisca has the distinction of being the earliest fossil species of Azolla in which
both vegetative and fertile remains were described (Reid and Chandler 1926). The
only record prior to this was that of Azollophyllum prirnaeviim Penhallow based on
vegetative material (Dawson 1890), later to be known as Azolla primaeva. Increasing
interest shown by many workers in Upper Cretaceous and Palaeogene species of
Azolla stimulated this reappraisal of A. prisca, using ultra-thin sectioning, and both
scanning and transmission electron microscopy. A. prisca is consistently described
in the literature as an enigmatic species which cannot be placed in any existing
FOWLER: OLIGOCENE AZOLLA
487
section of the genus due to the presence of anchor-shaped glochidia (Arnold 1955;
Hills and Gopal 1967; Trivedi and Verma 1971). In this present work, the establish-
ment of a new section to accommodate this species is considered both desirable and
necessary. Azolla species with nine floats in the swimming apparatus are particularly
characteristic of the Oligocene (see Table 1), a number of species having been recorded
from Britain and the U.S.S.R. (Reid and Chandler 1926; Dorofeev 1959). Its occur-
rence in the lowest Oligocene makes A. prisca one of the oldest nine-floated species
to be recorded, this type of swimming apparatus being rare in pre-Oligocene rocks.
This species affords the opportunity of investigating what may possibly be an
important evolutionary link between the Eocene and Oligocene representatives.
Critical study, coupled with adequate description and illustration, has never previously
been attempted for any Oligocene species of Azolla.
LOCALITY AND STRATIGRAPHY
A. prisca was described from the Insect Limestone of the Isle of Wight as part of the
rich Bembridge flora (Reid and Chandler 1926). The Insect Limestone is a fine-
grained blue-grey argillaceous limestone, varying in thickness up to 0-3 m and
located just above the base of the Bembridge Marls which reaches a maximum thick-
ness of some 33 m. The Bembridge Marls rest on the freshwater Bembridge Lime-
stone. Much of the Insect Limestone is barren and, according to Reid and Chandler
(1926), the plant collection was made from small pockets over a twenty-five year
period. Numerous insect remains are reported from this horizon (Woodward 1879).
The Insect Limestone is seen in the cliff-section of Gurnard Bay (SZ 467 943), but
to the west in Thorness Bay it reaches shore level.
During the Palaeogene, Hampshire occupied a marginal position between sea to
the east and land to the west. The Bembridge Marls accumulated during a regressive
phase when non-marine conditions were re-established after an initial transgressive
phase of short duration. According to Daley (1973), the depositional environment of
the Insect Limestone is not well understood.
Most workers consider the Bembridge Marls to be of Oligocene age, though an
Upper Eocene age has been suggested (Blondeau, Cavelier, Leugueur and Pomerol,
1965). Machin (1971), using palynological evidence, placed the Eocene-Oligocene
boundary at the base of the Lower Hamstead Beds, at the same time suggesting that
the base of the Bembridge Marls, lower in the succession, might be considered
a possible alternative. Preliminary palynological investigation of the Insect Lime-
stone by the author suggests that a lowermost Oligocene age might be appropriate
for the base of the Bembridge Marls.
Rock specimens used in this investigation had well-preserved massulae and mega-
spore apparatuses scattered over the surface (Specimens V. 17729 and counterpart,
British Museum (Natural History)). This material was originally collected by
J. E. E. A’Court Smith during the latter half of the nineteenth century. Repeated
attempts by the author to find suitable material of A. prisca from the type locality
were unsuccessful.
488
PALAEONTOLOGY, VOLUME 18
METHODS
Megaspore apparatuses and massulae were excavated from the rock surface by means
of fine needles, cleaned of adhering particles of matrix in 40% hydrofluoric acid,
then washed thoroughly in distilled water. Megaspore apparatuses were examined
and photographed, in dry condition and in water, using a Wild-Heerbrugg M7
Stereomicroscope and Photoautomat camera. Massulae were examined and photo-
graphed by means of a Wild-Heerbrugg M20 light microscope with Photoautomat.
Air-dried specimens for scanning electron microscopy, using a Cambridge Instru-
ments Company Stereoscan, were attached to double-sided sellotape on stubs, and
coated with gold using a Polaron E5000 Spatter Coater. Ultra-thin sections of the
megaspore apparatus were cut with a LKB Ultratome and glass-knife, after embedding
in Taab resin. Sections of thickness 1-5 /xm stained with basic fuchsin and methylene
blue were used for light microscopical examination, and of thickness 600 A stained
with uranyl acetate and lead citrate for examination with the transmission electron
microscope (Philips EM 300).
SYSTEMATIC DESCRIPTION
Order salviniales
Family salviniaceae
Genus azolla Lamarck, 1783
Section trisepta sect. nov.
Azolla prisca Reid and Chandler, 1926
The new section Trisepta is erected to include species of Azolla with the following
characteristics: megaspore apparatus clearly differentiated into megaspore and
swimming apparatus; swimming apparatus consisting of a hairy peltate columella
forming three compartments, each separated by a septum of columellar material;
attached to the columella are nine floats arranged in two tiers, three triangular floats
above and six oval floats beneath ; each compartment of the columella accommodates
one triangular and two oval floats; the massula bears non-septate anchor-shaped
glochidia each with a distal dilation immediately beneath the anchor-shaped tip.
The name Trisepta refers to the three septa of the columella which divide the
swimming apparatus into compartments housing the floats.
Details of vegetative features, structure of the sporocarp and sporangial wall of
A. prisca, are not included in this investigation which is concerned solely with the
dispersed megaspore apparatus and massula. According to Reid and Chandler
(1926) the megasporocarps and microsporocarps occur together in pairs with at least
twelve massulae in the microsporangium.
TEXT-FIG. 1. Azolla prisca Reid and Chandler, a and b, swimming apparatus, x 160. a, intact, b, floats
removed showing columella and labra on proximal surface of megaspore, c and d, section of megaspore
wall, c, with tubercle, and showing stratification, x 1750. d, at proximal pole, showing structural modifica-
tion and labra, x700. b base of striated layer; c columellate zone; ca ^ tomentose cap; co -= collar;
d dense zone; E = exine of megaspore proper; G granular layer, much thickened at proximal pole;
h hairy zone; H homogeneous layer; 1 labrum;o oval float; p = proximal surface of megaspore;
p perine;s tomentose septum; sp = spongy zone; S = striated layer; t ~ triangular float; tr position
of triradiate suture in exine.
FOWLER: OLIGOCENE AZOLLA
489
D
P
I ^
''"^1 >
t3::i5Mji:ra
E P
490
PALAEONTOLOGY, VOLUME 18
DESCRIPTION OF MEGASPORE APPARATUS
The megaspore is rounded, thick-walled, and bears several large rounded to vermi-
culate tubercles toward the distal surface. Above the proximal surface of the mega-
spore lies the swimming apparatus, the basic component of which, the columella, is
composed of long, unbranched and intertwined hair-like filaments. This columella
is peltate, with a small dome-shaped cap at the apex, and a hollow central strand with
wing-like longitudinal extensions dividing the swimming apparatus into three com-
partments. Attached to the columella are nine floats, an upper tier of three triangular
floats and a lower tier of six smaller, oval floats (text-fig. 1a, b; PI. 59, figs. 1, 2).
Most of the hairs forming the columella originate from a thickened area of mega-
spore wall which forms a prominent collar around the megaspore delimiting the
periphery of the proximal face. The columellar structure of A. prisca may best be
understood if it is regarded as resulting from the invagination of a dome-shaped
tomentose columella to form three compartments, each compartment being lined by
hairs. As a result, adjoining compartments are separated by a thick double-layer of
tomentose material which forms the septum. The thinner tomentose layer covering
the base of each compartment is supplemented by hairs from the proximal surface of
the megaspore. Each compartment accommodates a single upper triangular float and
two oval floats beneath. Hair-like filaments of the columella are continuous with
similar structures on the megaspore wall which, though scattered over the surface,
are particularly abundant near tubercles. The cap covering the uppermost area of the
triangular floats (PI. 59, figs. 1-3) has an inner layer of closely interwoven hairs,
continuous with the columella, and an outer darkly pigmented membraneous layer.
Length of megaspore apparatus 446 ixm to 475 ixm, average 455 /um; width of mega-
spore apparatus 237 ju,m to 270 ^um, average 255 |um (twelve specimens measured).
The floats are vacuolate pseudocellular masses, loosely attached to the columella
by relatively few hairs limited to the inner faces of the float (PI. 60, fig. 1). The central
region of the float is occupied by pseudocellular cavities approaching 25 |U,m in
diameter, whilst a well-delimited narrow zone of smaller cavities, average diameter
2-5 ju.m, occurs at the surface (PI. 60, fig. 3). Hairs on the floats are tubular extensions
of the peripheral cells (PI. 60, fig. 2) with a uniform diameter of approximately 2 jum,
often exceeding 75 |um in length and with blunt ends. A few closely spaced septa may
occasionally be found toward the hair base. The knot-like structure described by
various authors (Strasburger 1873; Campbell 1893; Hannig 1911; Kempf 1969^) can
often be seen within the float occupying a more or less central position (PI. 61, fig. 1).
EXPLANATION OF PLATE 59
Figs. 1 -5. Scanning electron micrographs of Azolla prisca Reid and Chandler from the Insect Limestone
(Bembridge Marls, Oligocene), Gurnard Bay, Isle of Wight. 1-2, intact megaspore apparatuses in dif-
ferent view. 1, swimming apparatus showing triangular float and two oval floats above hairy collar,
and apical cap. Distal surface of megaspore shows tubercles, x 175. 2, tomentose septum between two
smooth oval floats, x 175. 3, megaspore apparatus dissected revealing megaspore exine within perine.
Vacuolate floats and apical cap clearly seen, x 170. 4-5, perine surface. 4, tubercles formed of anastomos-
ing cylindrical elements. The spherical bodies are fungal spore contaminants, x 750. 5, surface regulate
verrucate, with foveae in which hair bases lie, X 1900.
PLATE 59
FOWLER, Azolla prisca
492
PALAEONTOLOGY, VOLUME 18
The substance forming this structure is regarded as rudimentary exine by Hannig
(1911) and Kempf (19696).
Removal of the columella shows a triradiate wall which attains a height of some
75 (um at the centre of the proximal surface of the megaspore, decreasing in height
toward the periphery (text-fig. 1b). In median longitudinal section the triradiate
wall is seen to be formed by the close association of two vertically orientated labra
bordering a suture (text-fig. Id; PI. 61, fig. 1). Pronounced sculptural and struc-
tural differences occur between the wall of the proximal face and that of the rest of
the megaspore. This was initially indicated by the darker coloration of the latter,
later to be confirmed by examination of the wall in thin section. The three compart-
ments of the columella lie directly above the inter-labral areas, the septa thus coinciding
with the labra.
The megaspore wall is composed of two principal parts, the exine of the megaspore
proper to the inside, and perine to the outside. The perine surface is rugulate-verrucate,
with foveae to a depth of some 2 ^um. Hairs scattered over the surface are 0-5-1 i^Lin
in diameter, their bases appearing to originate in the depressions of the foveae
(PI. 59, figs. 4, 5).
Dissection of the megaspore apparatus, via the proximal surface, reveals the
detached exine of the megaspore proper with the trilete mark uppermost (PI. 59,
fig. 3). This simply bordered trilete mark is small, with laesurae extending little more
than a third of the way to the equator; its position is coincident with the trilete suture
on the proximal face of the megaspore. The diameter of the megaspore proper is
approximately 190 |U.m; the surface is finely pitted to reticulate (PI. 60, figs. 4-7).
The megaspore wall is composed of three main layers, an exine and two-layered
perine, structural details of which can be seen in text-fig. Ic and Plate 61, figs. 3, 4.
In optical and thin section the exine is seen to have a thin basal zone above which it
is striated, the total thickness approaching 3 (PI. 60, fig. 5; PI. 61, fig. 3). As seen
with the transmission electron microscope, the exine has an essentially granular
structure, with the basal zone formed by fusion of elements (PI. 61, fig. 4). Above the
basement zone the elements, though fused to form a spongy network, are orientated
in such a way as to provide numerous narrow, radially arranged sinuous cavities of
varying length. Lateral fusion of these radially arranged elements in the uppermost
part of this layer provides a relatively smooth exinous surface. Numerous small
granules on this outer surface allow for a certain degree of interlock with the layer
above, yet rendering the exine readily detachable. The radially orientated elements
EXPLANATION OF PLATE 60
Figs. 1-12. Azolla prisca, megaspore apparatus and massula. 1-3, float structure. 1, oval float, biconvex in
side view, with hairs near apex of inner face, x375. 2, hair base, x 1550. 3, surface showing small
pseudocellular cavities, with larger cavities beneath, x 480. 4-7, megaspore proper. 4, proximal surface
showing trilete mark, X 180. 5, exine in optical section, x 1875. 6, exine surface at high, and 7, low
level of focus, x 1875. 8-T 1, massula. 8, massulae attached to megaspore apparatus, just below collar,
x75. 9, massula with anchor-shaped glochidia, x250. 10, anchor-tip showing two recurved prongs and
dilation beneath, X 1250. II, massula with microspores and associated funnel-shaped cavities, x940.
12, microspore-containing cavity and associated funnel-shaped cavity opening by a pore to the exterior,
X 940.
PLATE 60
FOWLER, AzoUa prisca
494
PALAEONTOLOGY, VOLUME 18
give the characteristic striated structure at lower magnification, and is consistent with
the sculpture of the exine.
The perine is divisible into an inner granular and outer homogeneous layer, the
latter stratified into four zones. Its thickness, though variable due to protuberances
and elaboration at the proximal surface, is approximately 10-5 ixm, excluding the
outer hairy zone. The granular layer, which is approximately 3 i^m thick, has a smooth
base and undulating upper surface (PI. 61, fig. 3). Transmission electron microscopy
shows that the structural elements within the striated and granular layers are similar,
though more haphazardly arranged in the latter (PI. 6 1 , fig. 4). The homogeneous layer
is supported above the granular layer by small solid columellae, 0-5-1 ;u,m in height,
forming the columellate zone. This passes up into a spongy zone, approximately
4-5 /xm thick, composed of an irregular network of large elements which fuse to form
a solid outer covering about 2-5 fxm in thickness. The surface of this outer dense zone
is dissected by grooves and pierced by foveae to give the sculpture already mentioned.
The outermost zone of the perine is composed of hair-like filaments originating from
the dense or spongy zones at the bases of the foveae, or from the reduced spongy
zone within the tubercles (text-fig. Ic; PI. 61, fig. 3). The wall of the tubercle is com-
posed of an anastomosing network of cylindrical elements developed from the dense
zone (PI. 59, fig. 4).
At the proximal pole of the megaspore, the perine structure is much modified
from that described above, this transformation taking place in the collar region
(text-fig. Id; PI. 61, figs. 1, 2). Here, the wall approaches 50 fxm in thickness (exclud-
ing the hairy zone), most of this being composed of the greatly thickened and vacuo-
lated granular layer. Correspondingly, the homogeneous layer is much reduced in
this region, being represented only by the dense and hairy zones. The proximal wall,
with an average thickness of 25 nm increasing to 100 |um in the formation of labra,
is composed almost entirely of the granular layer. On this proximal surface of the
megaspore the homogeneous layer almost completely disappears, remaining only as
scattered granules from which hairs may arise. The structure of the proximal wall,
with pseudocellular cavities as large as 21 jj.m in diameter, is reminiscent of that of
both float and massula. A single layer of small cavities, approximately 4 jxm in
diameter, form the outer surface of the proximal wall.
EXPLANATION OF PLATE 61
Figs. 1-4. Azolla prisca, megaspore apparatus in thin section. 1, median section through megaspore
apparatus, showing two triangular floats covered by cap, oval float beneath with ‘knot’ at centre, partially
detached exine, thick perine with tubercles, and vacuolate proximal wall with labra. Megaspore wall has
fractured in the collar region, x 200. 2, wall in collar region, showing thickening and vacuolation of
granular layer and reduced homogeneous layer represented only by dense zone. Hairs occur on dense
zone. Detached exine at bottom of figure, x 1400. 3, wall and tubercle. Outside the detached striated
exine, the granular layer supports the homogeneous layer divisible into columellate, spongy and dense
zones. Tubercle composed of anastomosing cylindrical elements formed by the dense zone. Within the
tubercle the spongy zone appears reduced, giving rise to hair-like structures, x950. 4, transmission
electron micrograph showing partially detached exine, granular layer, spongy and dense zones of homo-
geneous layer, x 4000.
PLATE 61
FOWLER, Azolla prisca
496
PALAEONTOLOGY, VOLUME 18
DESCRIPTION OF MASSULA
Numerous massulae of the same type were found associated with, and attached to,
megaspore apparatuses of A. prisca. There seems little reason to doubt that both
types of reproductive structure belong to the same parent species. Invariably, the
massulae are found attached by their anchor-shaped glochidia at, or near, the hairy
collar region of the megaspore apparatus (PI. 60, fig. 8). The massulae are vacuolate
pseudocellular discoid bodies ranging in maximum diameter, excluding glochidia,
from 98-8 to 167-2 [j.m, averaging 136-7 ij.m (fifty specimens) (PI. 60, fig. 9). As in
floats, the pseudocellular cavities of the massula range in diameter from approximately
5 /xm at the periphery, to about 25 fxm toward the centre. Occasionally, in both floats
and massulae, large cavities may extend to the periphery. Glochidia are rarely seen
projecting from the massula, being mainly restricted, and closely adpressed, to the
flattened surfaces. The nonseptate glochidia have an average length of 70 ;u,m; the
stalk, approximately 5-5 p.m wide in the median region, tapers to about 2-3 ^m at
the proximal and distal ends. A slight dilation of the stalk near its distal extremity
becomes abruptly constricted again at the Junction with the anchor-shaped tip.
Average width of the anchor-shaped tip is 8-4 ^m, and the two prongs of the anchor
are recurved (PI. 60, fig. 10). An inverted V-shaped diaphragm, often seen 5-6 fxm
beneath the distal dilation, separates what appears to be a tubular stalk from
a solid tip.
Microspores, averaging 20-8 ;u,m in diameter (range 15-25 |U,m, 150 specimens
measured), occupy large pseudocellular cavities within the massula. Six microspores
per massula appears usual, though the number ranges from three to nine. Micro-
spore walls are laevigate, with a thickness less than 1 jj.m. Each microspore-containing
cavity is closely associated with the bulbous base of a funnel-shaped cavity, the neck
of which extends to the periphery, opening by a pore to the exterior (PI. 60, figs. 11, 12).
The diameter of this pore approximates to that of the small cavities to the outside of
the massula. The germinal area of the microspores, represented by both closed and
open triradiate sutures, consistently occurs in a position adjacent to the base of the
funnel-shaped cavity (PI. 60, fig. 11). An incomplete partition appears to connect
these two adjoining cavities.
DISCUSSION
Most Azolla species with nine floats in two tiers in the swimming apparatus can be
included in sections Rhizosperma or Antiqua. Section Antiqua is established on the
basis of A. antiqua, a fossil species from the late Eocene and early Oligocene of the
U.S.S.R. in which the massulae are unknown (Dorofeev 1959). The nine-floated
species A. nana, A. turgaica, and A. ventricosa, described by Dorofeev (1959), and
in which the massulae are reported as eglochidiate (Hall and Swanson 1968; Trivedi
and Verma 1971), can be included in the Rhizosperma, together with the modern
species A. nilotica and A.pinnata. It would seem expedient to include the fossil species
A. aspera, A. nikitinii, and A. sihirica, together with yl. antiqua, in the section Antiqua
until further information is available on the massulae. Since A. prisca cannot be
included in either of these sections of the genus, due to its anchor-shaped glochidia.
FOWLER: OLIGOCENE AZOLLA
497
it seems appropriate to establish a new section, Section Trisepta, to accommodate
this, and similar species.
The columella in fossil and modern Azolla. Most pre-Eocene Azolla species were
described between 1969 and 1971, resulting in some synonymy, and confusion in the
descriptive morphology of the reproductive structures. The columella, as first recog-
nized by Meyen (1836), is well defined by the time Campbell (1893) describes it as
a short stalk from which microsporangia develop laterally. Since then, the term has
taken on dual usage, being also applied to that part of the swimming apparatus bear-
ing floats. Misinterpretation of the structure of the megaspore apparatus in both
fossil and modern species has resulted in a confused and inaccurate definition of the
term columella. One of the first applications of the term to the megaspore apparatus
is made by Eames (1936) in an account of the reproductive structures of modern
Azolla species. Here it is regarded as a conical pad of tissue situated, as the name
suggests, in a central position between the floats. Hall and Swanson (1968) illustrate
this interpretation of the columella with reference to the vacuolate pseudocellular
peg-like structure between the floats of the modern A. mexicana. However, in the
same work the authors describe the columella of the fossil species A. montana as
a hairy, hollow, thimble-shaped structure with attached floats. Further complica-
tions arise with the definition by Jain and Hall (1969), given as ‘a peg-like or cone-
shaped structure, distally continuous with the perispore and commonly hairy or
highly vacuolate’. Even when applied to the megaspore apparatus, the term columella
is apparently being used to describe two different structures, features of both having
become incorporated within the definition. This study of A. prisca indicates that the
vacuolate peg-like structure is best regarded as an elaboration of the proximal wall
of the megaspore, and not as part of the hairy superstructure primarily concerned
with holding the floats. On the bases of priority and aptness, the term columella
should be applied to the peg-like structure. However, as alternative terms may be
found for modification of the proximal wall of the megaspore, and since the term
has now been widely adopted for the tomentose superstructure in fossil species, it is
proposed to retain the term for that particular purpose. A reappraisal of the structure
of the columella would seem pertinent, especially as it is the major component of the
swimming apparatus in Palaeogene species of Azolla.
The columella is essentially a hairy superstructure over the proximal surface of the
megaspore, formed from hair-like filaments of the megaspore wall. The swimming
apparatus can be composed solely of this one component, forming a structure called
a columellate float, as in A. simplex (Hall \969b). Commonly, a second component,
the float, is developed. It is generally accepted that true floats are distinguished from
the columella by their vacuolate pseudocellular structure and lack of surface hairs.
Such distinction should be preserved, to the extent that a term such as columellate
float should be abandoned, this type of structure simply being regarded as an undif-
ferentiated columella. Similarly, in species such as A. montana, where structures
described as floats are hardly distinguishable from the columella, it would seem more
appropriate to refer to the structure as a segmented columella. The form of the
columella varies in different species, such variation depending on the volume of the
swimming apparatus given over to float production, and the size, shape, and number
498 PALAEONTOLOGY, VOLUME 18
of floats. At its simplest, the columella appears to be dome- or cone-shaped, as in
A. simplex and many multi-floated species. In A. prisca, and possibly all nine- and
three-floated species, modification of the dome-shaped type of columella by float
production in three sectors has restricted its development to a thin layer in the form
of a triseptate structure. A transverse section of this type of columella, as seen in the
Lower Pleistocene species A. tegeliensis, is well illustrated by Kempf (19696, fig. 8).
Here, two layers of tomentose material are seen to form each septum dividing float
compartments. The suggestion that the basic form of the columella is dome-shaped
is an oversimplification. In some species there is a tendency for only the central part
of the columella to be hollow, often with a pore-like opening at the apex. A. rnoniana
shows such a pore (Jain and Hall 1969), and Hall (19696), commenting on A. simplex,
states ‘in many specimens there is a canal in the columellate float, extending from the
apex of the megaspore body to the tip of the swimming apparatus’. The canal, though
narrow, is present in A. prisca, passing up through the central strand of the columella,
the position of the pore being marked at the apex by a small indentation in the apical
cap (text-fig. 1a; PI. 59, fig. 3).
There is some doubt as to whether the apical cap, so characteristic of A. prisca and
other nine-floated species in the Oligocene, is truly part of the columella. According to
Rao (1935), describing a similar structure in the modern nine-floated species A . pinnata,
it is a remnant of the inner part of the megasporangial wall. However, as can be seen
in A. prisca, both the columella and the outer membraneous megasporangial wall
would appear to be represented (PI. 59, fig. 1). Scanning electron microscopical
examination of megaspore apparatuses of modern nine- and three-floated species,
after removal of the enveloping megasporocarp and megasporangial walls, shows
that the columella forms a tomentose peltate structure lining the inside of the mega-
sporangial wall at the apex of the swimming apparatus. The columella within the
swimming apparatus of the modern A. filiculoides, A. pinnata, and A. nilotica has
been described by various authors (Bonnet 1957; Campbell 1893; Demalsy 1954;
Rao 1935) but its apparent insignificance precluded a term being applied. In these
species, representing both the three-floated and nine-floated condition, the peltate
part of the columella, which becomes inverted on removal of the megasporocarp and
megasporangial wall, is described simply as an abundance of hairs at the apex of the
swimming apparatus. The present work indicates that the columella of all nine- and
three-floated species, fossil and modern, is both peltate and triseptate. Furthermore,
the small tomentose columellar cap seen in fossil species such as A. prisca, was
probably more extensively developed in life, having been lost after release from the
parent plant. The tomentose cap is not recorded with any degree of certainty before
the Oligocene, though Hall (1969u) suggests that this feature may be present in
A. intcrtrappea, an Eocene species from India. The author is not aware of a similar
structure having been recorded for any megaspore apparatus other than the nine-
and three-floated type.
There is variation in the extent to which the swimming apparatus, and hence the
columella, covers the megaspore in fossil species. Jain and Hall (1969), suggests that
the swimming apparatus completely envelops the megaspore in ancestral types
similar to AzoUopsis, becoming progressively more restricted to the proximal pole
of the megaspore in the course of evolution.
FOWLER; OLIGOCENE AZOLLA
499
A. prisca appears to be the first species, chronologically and stratigraphically, in
which the triseptate columella is considered of structural and evolutionary significance,
hence the use of this feature in naming the section Trisepta.
Wall structure. Modification of the proximal pole of the megaspore can be seen in
fossil and modern species of Azolla. The earliest example of such modification can
be found in the late Cretaceous to Palaeocene species A. schopfi, described by Dijkstra
(1961) as having triradiate ridges, 60 pm in height, which are clearly visible when the
swimming apparatus is absent. A. distincta, with a similar stratigraphic range, is
described as having columella and floats with the ‘same foamy texture’ (Hall and
Bergad 1971). Possibly this foamy columella in A. distincta is really the proximal face
of the megaspore with vacuolate pseudocellular structure like that of the floats, as
in A. prisca. Study of the literature suggests that apart from the above-mentioned
species similar modification may occur in extincta, A. montana, A. fragilis, A. velus,
A. intertrappea, and indica, and may be a common feature of pre-Oligocene species.
In A. prisca structural modification of the proximal surface takes the form of lips
bordering the triradiate suture. A transition zone of this sort between sutures and the
remainder of the proximal surface, due to increased wall thickness, sculptural
modification, or both, is termed a labrum (Couper and Grebe 1961). The term gula,
applied by Kempf (19696) to a similar structure in A. tegeliensis, is best retained for
a more marked extension of the labra than that seen in A. prisca.
Wall stratification in Azolla megaspores is complex and a number of terminologies
has been applied. Details of terminologies used, together with suggestions as to
various authors’ interpretations of the megaspore wall structure, is not within the
scope of this work which is not primarily concerned with developmental history of
the wall layers. However, it would be appropriate simply to outline the main termino-
logies used for both modern and fossil species. Campbell (1893) recognized two
principal layers in the megaspore wall of A. filicidoides, an inner exospore, and an
outer epispore which could be sub-divided into two zones. Later workers adopted
the same terminology in describing the wall of modern Azolla, though Demalsy
(1954) and Bonnet (1957) delimited an innermost endospore from the exospore.
By this time, perispore was being used as an equivalent term to epispore. Kempf
(1969fl and h) uses the terms exine, instead of endospore and exospore, and perine,
instead of epispore and perispore, for both fossil and modern species. In earlier work
on fossil species the exine was termed endospore and most of the perine was called
exospore, the term perispore being reserved only for the outer hairy zone of the
perine (Hall and Swanson 1968; Jain and Hall 1969). Jain (1971), though using
the same basic delimitation of layers, termed the hairy zone as perine, and applied the
terms endexine and ectexine to the exine and remainder of perine respectively.
The terminology adopted for A. prisca, and for fossil and modern species mentioned
in this work, is that used by Kempf (1969a and b). Sweet and Hills (1971), describing
the megaspore wall of A. pirmata, indirectly make use of the term perine by intro-
ducing the terms inperine and experine. Such introduction of new terms is, in the
author’s opinion, both unjustifiable and unnecessary.
Wall structure of modern species has received little detailed attention, and the
development of the wall layers is imperfectly understood. Such information available
500
PALAEONTOLOGY, VOLUME 18
indicates that, as in prisca, the exine is composed of a basement zone and radially
striated zone. The inner layer of the perine (equivalent to the granular layer of
A. prisca) is described as granulate, reticulate, or foamy, in A. pinnata, A. nilotica,
and A. filiculoides respectively, such differences possibly reflecting the degree of
vacuolation of this layer in each species. In A. filiculoides it is called the ‘couche
ecumeuse’, is characteristically vacuolated and extensively developed, forming
eruptions toward the surface (Bonnet 1957). Demalsy (1954), describing structural
variation in the megaspore wall of A. nilotica, points out that this inner layer of the
perine appears thicker and heterogeneous in some megaspores. However, similar
structural differences may be observed in oblique sections of the wall of A. prisca.
The outer homogeneous layer of the perine is composed primarily of radially elongated
elements in A.pinnata and A. nilotica, that of A. filiculoides appearing to have a denser
structure. As in A. prisca, the outermost zone of the homogeneous layer is composed
of hairs. Hannig (1911) suggests that in A. filiculoides hairs originate from eruptions
into the dense zone of the vacuolate granular layer, a suggestion later accepted by
Bonnet (1957). Photographs of the ultrastructure of the megaspore wall of A. fili-
culoides, loaned to the author (Pettitt 1974, pers. comm.), would appear to sub-
stantiate this. However, it is possible that a very thin layer of homogeneous material
is deposited over the granular layer, at the site of the eruption, before hairs form.
Hairs certainly appear to originate directly from the surface of the dense zone in the
intervening areas, as illustrated by Bonnet (1957, fig. 35). In A. prisca, hairs appear
to be composed of the same material as that of the homogeneous layer, and they can
be seen to arise from the spongy zone within the tubercle and from the dense zone in
the collar region. Hairs which originate in foveae, away from the tubercle, appear to
originate from the dense zone, though a connection between the hair base and spongy
zone beneath is not discounted.
Details of wall structure are now known for several Palaeogene species, with
A. prisca furnishing the best example from the Oligocene. Although there is distinct
variation in thickness and structure of individual layers or zones, a similar pattern
to that seen in modern species is shared by the majority of fossil types. This basic
pattern in fossil species can be summarized as follows: An exine, 3-4 /j,m thick, is
covered by a two-layered perine. The inner layer of the perine is smooth, granular,
or laminated, and between 3-8 |U.m in thickness. It is the outer homogeneous layer
which shows most interspecific variation, and it is divided into at least three zones by
most authors. There is often a columellate zone supporting a reticulate or clavate
zone, the elements of which become fused toward the surface from which hairs arise.
As Pettitt (1966) points out, the term perine, and equivalent terms, are indis-
criminately used for a variety of exinous and extra-exinous layers of the spore or
pollen wall. Erdtman (1952) defines the perine as an outer extra-exinous layer formed
by the activity of a tapetal plasmodium. It is acknowledged that use of such terms as
exine and perine for A. prisca has developmental implications. However, structural
similarities between the vacuolated granular layer, massulae, and floats of A. prisca
cannot be disregarded, and suggest that these structures may be homologous. This
similarity is even more marked in modern T . /7//cu/o/V/(?5' where the vacuolated granular
layer is extensively developed in the megaspore wall, not merely limited to the
proximal pole as in A. prisca. As massulae and floats are usually accepted as perinous.
FOWLER: OLIGOCENE AZOLLA
501
there seems reasonable justification in regarding the granular and homogeneous
layers of A.priscaas perine. Only intensive study of the development of the megaspore
wall of modern Azolla will reveal whether a particular layer is exinous or perinous.
Structural resemblances between the wall of modern and fossil species suggest the
possibility that conclusions reached from study of modern species may also be applied
to fossil species.
Kempf (1969(3 and b, 1973), equates the perine of megaspore walls with the ektexine
of angiosperm pollen grains, emphasizing the presence of foot layer, columellae, and
tectum in both. At the same time the spore exine is equated with the endexine of the
pollen grain. This would seem unacceptable, as the term perine can only correctly be
applied to plants with a periplasmodial tapetum, and which is known to contribute
sporopollenin to the pollen grain wall. Though Kempf ’s (1969fl) terminology is
used, the author does not hold the view, expressed by Kempf (1973), that all spore
walls have perine to the outside, the exine beneath playing no part in surface orna-
mentation. The terms adopted here are intended only for species of Azolla.
Snead’s (1969) contention that wall structure is sufficiently variable in fossil
Azolla to allow species identification from wall fragments would seem to be correct,
though it should be applied with caution. More intensive study using ultra-thin
sectioning would seem necessary before individual species can be identified with any
degree of confidence. The extent of variation within a single megaspore wall, and
between spores of the same species must be evaluated. Interpretation of the wall of
fossil megaspores found attached to the parent plant may result in description of
immature structure, since megaspores become detached at maturity.
Development of labra, or similar structures, appears to be linked with a thickening
of the megaspore wall to form a collar around the proximal face of the megaspore.
The Eocene species A. intertrappea not only provides the earliest example of this
association, but is also the earliest species to demonstrate the significant part played
by the granular layer in the formation of the proximal wall and labra (Sahni 1941).
Such development of the proximal pole is readily seen in A. prisca, where the collar
and labra are prominent features of the megaspore. Here, the collar region and
proximal wall with labra are composed almost entirely of the thickened vacuolated
granular layer, the homogeneous layer becoming reduced to a thin covering of
scattered granules which may be associated with hair development. This association
of labra and collar, with a prominent granular layer in the proximal wall, probably
occurs in other nine-floate(i Oligocene species. After the Oligocene, it can again be
seen in the Lower Pleistocene species A. tegeliensis (Kempf 19696) and A. pyrenaica
(Florschiitz and Menendez Amor 1960), representing the nine-floated and three-
floated condition respectively. The apparent size difference between the gula of
A. nana, as illustrated by Kempf (1969(3, pi. 13, fig. 8), and the labrum of A. prisca,
may possibly be accounted for by differences in preparation technique. In A. nana,
contraction of the megaspore wall in the collar region, with associated infolding of
the exine, has also resulted in the extrusion of the vacuolate proximal wall upward
between the floats, so giving an exaggerated size to the structure which has been
termed a gula. It is conceivable that the apparent size of the gula is partly responsible
for the importance attached to this structure by Kempf (19696) with regard to anchor-
age of floats, the tomentose columella being considered of minor significance.
502
PALAEONTOLOGY, VOLUME 18
Maximum development of the proximal pole of the megaspore occurs in modern
species. Concerning the role of the granular layer, Rao (1935) states that this layer
forms the spongy conical structure at the top of the megaspore in A. piimata. The
author’s own observations, together with published description and illustration,
indicate that modern species have a much enlarged collar, a central conical structure
trisected by sutures, and a similar role for the granular layer as that seen in A. prisca
(Rao 1935, pi. 19, fig. 45; Demalsy 1954, pi. 10, figs. 201-202; Bonnet 1957, text-
fig. 3; Hall and Swanson 1968, fig. 14).
Relevance of funnel-shaped cavities. Funnel-shaped cavities occurring in association
with microspore-containing cavities is a consistent feature of all massulae of A. prisca
examined. The germinal area of the microspore, represented by a triradiate mark,
consistently occurs in a position adjacent to the base of the funnel-shaped cavity,
which often appears as a thin incomplete partition. The exterior pore and the open-
ing in the base of the funnel-shaped cavity are found associated with microspores
having both closed and open sutures. The occurrence of uniform funnel-shaped
cavities associated with closed sutures, together with the intact nature of the funnel,
not appearing to have formed by breakdown of pseudocellular material, would seem
to support the suggestion that these structures are features of the primary develop-
ment of the massula, and not formed as a result of post-germination prothallial
activity. Such organization could conceivably have formed an escape mechanism
for spermatozoids, and to the author’s knowledge, no directly comparable structure
has previously been recorded for Azolla species.
Fames (1936), describing microspore germination in modern Azolla, states that
a papilla protrudes through the opened sutures, then differentiates to form a small
prothallus on which develops an antheridium producing eight spermatozoids. The
gametophyte remains embedded in the massula, spermatozoids being freed by the
eventual breakdown of the outer part of the massula. Most accounts of microspore
germination in Azolla are similar, there being no mention of funnel-shaped cavities.
It seems unlikely that such organization, which would seem to impart distinct bio-
logical advantage, was no longer a feature of the massulae of modern species. Pre-
liminary examination of massulae of modern A. filiculoides by the author reveals
a similar organization to that described for A. prisca. It seems possible, therefore,
that such organization may occur in the massulae of all Azolla species, fossil and
modern.
Evolutionary trends. Pioneering work by Hills and Gopal (1967) led to the con-
clusion that the three-floated condition in the swimming apparatus of Azolla is
ancestral to the nine-floated condition. This conclusion was based largely on the
discovery of A. geneseana, a late Cretaceous species purported to possess three
floats from study of a few poorly preserved specimens (Hills and Weiner 1965).
Other workers have since failed to establish the presence of three-floated specimens
at the type locality (Snead 1969; Jain 1971). Together with A. simplex, the oldest
species yet described (Hall 19696), A. geneseana is best considered as possessing
a tomentose columella without float diflerentiation. A. primaeva, an Eocene species
once erroneously reported as having three floats (Hills and Weiner 1965; Hills and
Gopal 1967), is now believed to have a single large vacuolate pseudocellular float-
FOWLER: OLIGOCENE AZOLLA
503
like structure on which surface hairs may be present (Hall 1969n). Apart from species
just mentioned, the majority of late Cretaceous and Palaeocene species are multi-
floated, bearing up to twenty-four floats which may, or may not, have structure
different from that of the columella. Readily distinguishable floats probably made
their appearance before the close of the Cretaceous, as in A. barbata (Snead 1969;
Hall and Bergad 1971). Many multifloated species, however, have little more than
a segmented columella, as in A. montana, in which the so-called floats are hardly
differentiated from the hairy columella. The float-like structures described in
A. distincta, A. fragilis, A. stanleyi, Sind A. veins may represent an evolutionary
advancement in float development, in that they lack hairs and are readily removed
from the dome-shaped columella (Hall and Bergad 1971; Jain and Hall 1969).
Megaspores in which the floats, proximal wall, and collar region are distinctly vacuo-
late make their first appearance in the Eocene of India, with A. intertrappea and
A. indica. These two species are the earliest recorded as showing the three-floated
condition, though this is still a matter for speculation. Hall (1969<7), in a revision of
A. intertrappea, accepts the presence of three floats, and describes this species as
having a well-developed peltate columella bearing small floats approximately half
the size of those of extant three-floated species. Megaspores with three floats, similar
to those of extant species do not appear until the Oligo-Miocene (Lancucka-
Srodoniowa 1958). Sahni (1941 ) points out that the exact number of floats in A. inter-
trappea is difficult to ascertain in longitudinal section, due mainly to the thickness of
the sectioned material, the floats being seen at different levels of focus. From such
material, interpretation of A. intertrappea as having one or nine floats would appear
equally possible, and on stratigraphic evidence, would seem more plausible. A. indica,
described by Trivedi and Verma (1971 ) as probably having three floats, furnishes no
better evidence for the existence of the three-floated condition in the Eocene. Until
further information is available, A. intertrappea and A. indica are accepted as having
three floats in the swimming apparatus. These two species are very similar, and may
eventually be regarded as con-specific. Main differences reported are number of
microsporangia, size of microsporangia and massulae, and structure of the glochidia,
which is described as septate in A. indicia and non-septate in A. intertrappea (Trivedi
and Verma 1971). Presence or absence of a septum provides the only difference of
possible value, though it seems likely that the diaphragm separating the solid glochidial
head from the stalk, as seen in A. prisca, may have been interpreted as a septum.
Evolutionary development of the swimming apparatus of Azolla seems reasonably
clear, and may be summarized as follows. The multifloated condition appears more
ancient than the nine-floated and three-floated condition, but due to the presence of
A. simplex and A. geneseana in the Cretaceous, it is not known if the most ancient
types were multifloated or were without floats. Considering the tomentose columella
as the basic component of the swimming apparatus, it seems likely that the dome-
shaped columella without floats is the most primitive type. From this type of struc-
ture the columella becomes progressively organized into separate areas to form
float-like structures of the multifloated condition. At first the columella becomes
segmented, as in A. montana, the segments hardly distinguishable from the hairy
columella. Some species, like A. distincta, are reported as having foamy floats, imply-
ing that true floats appear before the close of the Cretaceous. A number of Cretaceous
504
PALAEONTOLOGY, VOLUME 18
species probably possess floats which, though distinguishable from the columella,
do not have the vacuolate structure of later species. Numerous vacuolate float-like
structures, readily distinguished and easily removed from the columella, is a feature of
some Palaeocene species. Eocene records are scanty and not particularly reliable,
though it is evident that the number of floats is reduced, the columella becomes less
extensive and floats take up a correspondingly larger volume of the swimming
apparatus. Eocene species are known to possess true floats showing smooth surface
with vacuolate pseudocellular structure, and the proximal wall of the megaspore is
modified to form a collar region and labra, as in /4. inter irappea. In this species, the
columella is still a significant feature of the swimming apparatus, the floats are much
smaller than those of extant species and undifferentiated columella remains in the
form of a dome at the apex. In Oligocene species such as A. prisca, perhaps we see
maximum elaboration in the form of the triseptate columella, resulting from float
development in three sectors of the swimming apparatus. The columella, though much
reduced in volume, is still a significant structure supporting nine floats. The three
floats of the upper tier are held together by the cap-like development of the columella
at the apex, whilst those of the lower tier are supported in position by the contours
of the modified proximal pole and held together by the triseptate columella.
Species having three large floats like those of living species first occur, with certainty,
in the late Oligocene or Miocene. The structure and role of the columella appears to
have remained largely unaltered since the close of the Eocene, despite the develop-
ment of the three-floated condition. However, with the development of three large
floats, the significance of the columella has possibly been slightly diminished as
a result of increased elaboration of the proximal pole, more support for the floats
being provided by the development of the collar, labra, gulae, etc. How a species such
as primaeva, with a single pseudocellular float, could fit into this basic evolutionary
pattern is not known. Its late appearance, in the Eocene, suggests that it might have
developed from an ancestor similar to A. simplex, as a result of vacuolation.
Evolutionary tendencies are not clear regarding the massulae. High numbers of
microspores per massula is a feature of multifloated species, with fifty microspores
recorded in the massula of A. extincta, considered by Jain (1971) to represent the
contents of a single microsporangium. Less than eight microspores per massula occur
in the Eocene species A. indica and the Oligocene species A. prisca, this low number
being retained as a feature of post-Oligocene species, both fossil and modern.
Mahabale (1963) proposed an evolutionary scheme for Azolla based on structure of
glochidia, on the assumption that section Azolla, with septate anchor-shaped
glochidia, is primitive. He suggested that these primitive glochidia became reduced,
losing both septa and anchor-tip, to become filamentous as in modern A. pinnata
and eventually eglochidiate as in modern A. nilotica. Section Azolla is no longer
considered primitive, but anchor-shaped glochidia, though found occurring with
every known type of megaspore apparatus throughout the stratigraphical range of
Azolla, are consistently associated with multifloated megaspores (Jain and Hall
1 969). According to Hills and Gopal ( 1 967), septation is a recently acquired character,
not an ancestral one, the earliest record of septate glochidia occurring in the Pleisto-
cene. As previously mentioned, there is some doubt concerning the reported presence
of septa in the glochidia of the Eocene species A. indica.
FOWLER: OLIGOCENE AZOLLA
505
It was Kempf (1969a and b) who provided the basis of our understanding of the
structural morphology of the megaspore apparatus of AzoUa species, with particular
emphasis being placed on the ultrastructure of the megaspore wall. This present
investigation of A. prisca, using modern techniques, further extends this knowledge.
Ultra-thin sectioning and transmission electron microscopy of the megaspore wall
supports Kempf ’s (1969a) suggestion that wall structure is sufficiently variable as
to provide a useful means of taxonomic separation and identification of both modern
and fossil types, and provides further information on the origin of the hair-like
structures. Furthermore, the significance of the innermost layer of the perine in the
formation of the proximal wall of the megaspore and associated labra, gulae, etc.,
is given more attention than in previous work. Scanning electron microscopy employed
in the study of the megaspore apparatus of A. prisca and some modern species, has
led to a better understanding of the structure and significance of the columella, the
sculpture of the megaspore wall, and the nature of the apical cap. Critical study of the
massulae of A. prisca and modern species, has revealed structural organization
previously unrecorded. As a result of this investigation on the reproductive structures
of A. prisca, this species becomes one of the best known of all species of Azolla, both
fossil and modern.
Our knowledge concerning fossil Salviniaceae has expanded considerably in
recent years, indicating great diversity in reproductive morphology and megaspore
wall structure. At one time, the Salviniaceae was regarded as having two living genera,
Salvinia and AzoUa, the latter divided into two sections, Azolla and Rhizosperma.
We are now aware of the importance of the genus AzoUa in the Upper Cretaceous and
Lower Tertiary, from which over thirty species have been recorded and placed in
seven sections of the genus. In addition, two new genera, AzoUopsis (Hall 1968) and
Parazolla (Hall 1969^), have been established within the Salviniaceae.
Acknowledgements. The author is indebted to the Keeper of Palaeontology, British Museum (Natural
History) for material of Azolla prisca. My gratitude to Dr. J. M. Pettitt, British Museum (Nat. Hist.) and
Dr. K. R. Sporne, Department of Botany, University of Cambridge, for their helpful advice on certain
aspects. However, responsibility for opinions expressed, and any errors, rest with the author. Sincere
thanks are due to Mrs. P. Palmer for sectioning material, to Diane Irwin and Steve Furtado for assistance
with scanning electron microscopy, to Helen Harris for typing the manuscript and to the photography
section of the Portsmouth department. Lastly, to Dr. L. V. Hills, University of Calgary, who stimulated my
interest in A. prisca during an unsuccessful collecting trip to the type locality in the summer of 1972.
REFERENCES
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37-45.
BERGGREN, M. s. 1880. Om Azollas prothallium och embryo. Lunds. Univ. Arsskrift, 16, 1 1 1.
BLONDEAU, A., CAVELIER, c., FEUGUEUR, L. and POMCROL, c. 1965. Stratigraphie du Paleogene du Bassin de
Paris en relation avec les bassins avoisinants. Bull. Soc. geol. Fr. 1, 200-221.
BONNET, A. L.-M. 1957. Contribution a I’etude des Hydropteridees 3. Recherches sur Azolla filiculoides
Lamk. Rev. Cytol. Biol. Veg. 28, 1-88.
CAMPBELL, D. H. 1893. On the development of Azolla filieuloides Lam. Ann. Bot. 1, 155-187.
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Rendu, 3 {Reunion Comm. Internal. Microflore Paleozoique, Rept. Group 16), 15 pp.
DALEY, B. 1973. The palaeoenvironment of the Bembridge Marls (Oligocene) of the Isle of Wight, Hamp-
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PALAEONTOLOGY, VOLUME 18
DAWSON, J. w. 1890. On fossil plants from the Smilkameen Valley and other places in the southern interior
of British Columbia. Trans. R. Soc. Can. 8, 75-91.
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DUNCAN, R. E. 1940. The cytology of sporangium development in Azolla filiculoides. Bull. Torrev bot. Club,
67,391-412.
EAMES, A. J. 1936. Morphology of vascular plants. Lower groups. McGraw-Hill Book Co., New York, 433 pp.
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FLORSCHUTZ, F. 1945. Azolla uit het Nederlandsche Palaeoceen en Pleistoceen. Verb. Geol. Mijnb. Gen.
Nederl. KoL, Geol. 14, 191-197.
and MENENDEZ AMOR, J. 1960. Une Azolla fossile dans les Pyrenees Orientales. Pollen et Spores, 2,
285-292.
GRIFFITH, w. 1844. On Azolla and Salvinia. Calcutta J. Nat. Hist. 5, 227-273.
HALL, J. w. 1968. A new genus of Salviniaceae and a new species of Azolla from the late Cretaceous. Am.
Fern. J. 58, 77-88.
1969fl. A reappraisal of the megaspores of two Eocene species of Azolla. J. Paleont. 43, 528-531.
19696. Studies on fossil Azolla: Primitive types of megaspores and massulae from the Cretaceous.
Am. J. Bot. 56, 1173-1180.
1970. A new section of Azolla. Taxon, 19, 302-303.
and BERGAD, R. D. 1971. A critical study of three Cretaceous salviniaceous megaspores. Micro-
paleontology, 17, 345-356.
and SWANSON, n. p. 1968. Studies on fossil Azolla: Azolla montana, a Cretaceous megaspore with
many small floats. Am. J. Bot. 55, 1055-1061.
HANNIG, E. 1911. liber die Bedeutung der Periplasmodien. 2. Die Bildung der Massulae von Azolla. Flora
Jena, 102, 243-278.
HILLS, L. V. andGOPAL, B. 1967. Azolla primaeva and its phylogenetic significance. Can. J. Bot. 45, 1179-1 191.
and WEINER, N. 1965. Azolla geneseana, n. sp., and revision of Azolla primaeva. Micropaleontology,
11, 255-261.
JAIN, R. K. 1971. Pre-Tertiary records of Salviniaceae. Am. J. Bot. 58, 487-496.
and HALL, J. w. 1969. A contribution to the early Tertiary fossil record of the Salviniaceae. Ibid. 56,
527-539.
KEMPF, E. K. 1969u. Elektronenmikroskopieder Sporodermis von kanozoischen Megasporen der Wasserfarn-
Gattung Azolla. Paldont. Z. 43, 95-108.
19696. Elektronenmikroskopie der Megasporen von Azolla tegeliensis aus dem Altpleistozan der
Niederlande. Palaeontographica, B 128, 167-179.
1973. Transmission electron microscopy of fossil spores. Palaeontology, 16, 787-797.
LANCUCKA-SRODONIOWA, M. 1958. Salvinia i Azolla w miocenie Polski. Acta Biol. Cracov. Bot. 1, 15-23.
MACHiN, J. 1971. Plant microfossils from Tertiary deposits of the Isle of Wight. New. Phytol. 70, 851-872.
MAHABALE, T. s. 1963. Evolutionary tendencies in the genus Azolla. J. Indian Bot. Soc. Mem. 4, 51-54.
METTENius, G. 1847. Uber Azolla. Linnaea, 20, 259-282.
MEYEN, F. J. F. 1836. Beitrage zur Kenntnis der Azollen. Nova Acta Acad. Caesar. Leap. Carol. 18, 505-524.
PETTiTT, J. M. 1966. Exine structure in some fossil and recent spores and pollen as revealed by light and
electron microscopy. Bull. Br. Mus. nat. Hist. (Geol.), 13, 221-257.
PFEIFFER, w. M. 1907. Differentiation of sporocarps in Azolla. Bot. Gaz. 44, 445-454.
RAO, ti. s. 1935. The structure and life history of Azolla pinnata R. Brown with remarks on the fossil history
of the Hydropteridae. Proc. Indian Acad. Sci. 2, 175-200.
REID, E. M. and CHANDLER, M. E. J. 1926. Catalogue of Cainozoic plants in the Department of Geology, I. The
Bemhridge Flora. Brit. Mus. (Nat. Hist.), London, 206 pp.
.SAHNi, B. 1941. Indian silicilied plants. 1. Azolla intertrappea Sah. & H. S. Rao. Proc. Indian Acad. Sci.
B 14, 489-501.
- and RAO, H. s. 1943. A silicilied flora from the Intertrappean cherts round Saugar in the Deccan.
Proc. natn. Acad. Sci. India, 13, 36-75.
SCULFHORPE, c. D. 1967. The biology of aquatic vascular plants. Edward Arnold, London, 610 pp.
FOWLER; OLIGOCENE AZOLLA
507
SNEAD, R. G. 1969. Microfloral diagnosis of the Cretaceous-Tertiary boundary, central Alberta. Bull.
Alberta Res. Council, 25, 1-148.
1970. A new approach to the classification of Azolla megaspore species (abstract). Geoscience and
Man, 1, p. 135.
STRASBURGER, E. 1873. Uber Azolla. Jena.
1889. Histologische Beitrdge. 2. Uber das Wachsthum vegetabUischer Zellhaute. Jena.
SVENSON, H. K. 1944. The New World species of Azolla. Am. Fern. J. 34, 69-84.
SWEET, A. and hills, l. v. 1971. A study of Azolla pinnata R. Brown. Ibid. 71, 1-13.
TRiVEDi, B. s. and VERMA, c. L. 1971. Contributions to the knowledge of Azolla indica sp. nov. from the Deccan
Intertrappean Series M.P., India. Palaeontographica, B 136, 71-82.
WOODWARD, H. 1879. On the occurrence of Branehipus (or Chirocephalus) in a fossil state, associated with
Eosphaeroma and with numerous insect remains, in the Eocene Fresh Water (Bembridge) Limestone of
Gurnet Bay, Isle of Wight. Q. J! geol. Soc. Land. 35, 342-350.
Typescript received 12 August 1974
Revised typescript received 2 December 1974
K. FOWLER
Department of Biological Sciences
Portsmouth Polytechnic
King Henry I Street
Portsmouth POl 2DY
E
LUDLOW BENTHONIC ASSEMBLAGES
by J. D. LAWSON
Abstract. The communities recently described by Calef and Hancock are considered to provide an inadequate
picture of Ludlow faunas and their palaeoecological significance. Alternative assemblages, including the important
non-brachiopod benthos, have been compiled from the evidence of published faunal lists. It is here maintained that
these four assemblages reflect more accurately than those of Calef and Hancock the faunal distribution within the
Ludlow rocks but no special significance is claimed for them; each contains subdivisions which may be more readily
explained in palaeoecological terms. It is suggested that the recent emphasis on depth-communities has led to neglect
of other very important environmental controls, particularly the nature of the substrate. The concept of continuous
regression through the Ludlow is considered untenable in the light of sedimentological evidence. The degree of
diachronism of the shelly faunas is assessed. It is concluded that the picture drawn by Calef and Hancock is an over-
simplification resulting, perhaps, from the attempt to impose a relatively straightforward Llandovery pattern on to
the more complex Ludlow rocks.
A RECENT paper in Palaeontology by Calef and Hancock (1974) describes five major
marine benthonic communities occurring in clastic (i.e. terrigenous) sediments laid
down in areas of increasing depth of water ‘from the shoreline to deep areas in Wales
and the Welsh Borderland during Wenlock and Ludlow times’. Only four of these
communities are well developed in the Ludlow rocks, mainly on the stable eastern
margin of the Welsh basin. They are named after characteristic brachiopod genera
as follows; (1) Salopina, (2) Sphaerirhynchia, (3) Isorthis, and (4) Dicoelosia com-
munities. They are considered as approximate equivalents of the upper Llandovery
depth-communities; (1) Eocoelia, (2) Pentamerus, (3) Stricklandia, and (4) Clorinda,
the last-named being the deepest of the four. A fifth, deeper Visbyella community
has only been recognized at one locality in the Ludlow rocks.
The purpose of this paper is to critically examine some of Calef and Hancock’s
conclusions in the light of the very detailed published evidence on Ludlow faunas
and sediments. Text-fig. 1 shows the localities mentioned in this paper.
RECOGNITION OF COMMUNITIES
The statistical description of their communities by Calef and Hancock is most
welcome but it is not clear how collections were assigned to a particular community
in the first place. Perhaps the allocation was made on the basis of survivors from
Llandovery communities but these form a small proportion of the faunas and some
genera apparently change their modal communities (Calef and Hancock 1974,
Table 12). It is stated that the communities completely intergrade and that the com-
munity boundaries are arbitrary lines through a continuum (Calef and Hancock 1974,
p. 779). The number of divisions is therefore a matter of practical convenience and
ten communities could have been postulated instead of five. The recognition of five
communities has proved to be of value in the upper Llandovery and this fact has
presumably influenced the choice of five for the Wenlock and Ludlow.
Although it is at first reassuring to have the presence percentage and frequency
[Palaeontology, Vol. 18, Part 3, 1975, pp. 509-525.]
510
PALAEONTOLOGY, VOLUME 18
TEXT-FIG. 1, Map of south-east Wales and the Welsh Borderland showing the outcrops of Ludlow rocks
(stippled) and localities mentioned in the text.
presence recorded for each fossil in each community it is not clear how a geologist |
attempting to use these tables is expected to allocate a particular fauna to one of these
communities. For instance, in the Wenlock Salopina community (Calef and Hancock
1974, Table 2) five of the seven prevalent fossils are present in less than one-third of
the seventeen localities examined—evidently not prevailing very successfully.
ASSEMBLAGES AS LIFE-ASSEMBLAGES
Calef and Hancock follow the practice of Ziegler, Cocks and Bambach (1968, p. 3) t
in accepting the shell assemblages as reasonably representative of the preservable *
LAWSON: LUDLOW BENTHONIC ASSEMBLAGES
511
elements of the communities. This seems well justified in the Llandovery for the
following reasons:
1. Analysis of shells suggests only limited post-mortem transport.
2. The faunal associations are so frequently repeated.
3. Life-assemblages of similar composition to the transported assemblages have
been recorded.
4. The community belts are about ten miles wide and only extensive shell transport
could confuse the basic pattern.
Points (1) and (2) apply also to the Wenlock and Ludlow rocks (Calef and Hancock
1974, p. 781) but so far no depth-patterns have been established or life-assemblages
described. Although it seems likely that Calef and Hancock’s thesis does apply to
most of the shelf Ludlow faunas, it should be accepted with caution until broad dis-
tribution belts have been established, life-assemblages recognized, and further
analysis of shell wear and distribution carried out.
COMPOSITION OF ASSEMBLAGES
If, however, the above contention is broadly acceptable it means that previously
described Ludlow assemblages may now merit consideration as life-assemblages and
deserve close comparison with the communities listed by Calef and Hancock. Straw
(1937, p. 411) and Lawson (1960) both described Ludlow faunas but much new
information has been published since those papers and it is now possible, using
detailed faunal lists, to draw up tables of four major Ludlow benthonic assemblages
occupying the main shelf area of the Welsh Borderlands and the English Midlands.
They characterize the four Ludlow stages (Eltonian, Bringewoodian, Leintwardinian,
and Whitcliffian) on the shelf and, therefore, succeed each other vertically in any
particular area. Unlike the intergrading communities postulated by Calef and
Hancock three of these four assemblages suflFer abrupt vertical changes in faunal
content. This can be most clearly seen in the range charts included in the papers on
Usk (Walmsley 1959, p. 490), Woolhope (Squirrell and Tucker 1960, p. 144), Malvern
(Phipps and Reeve 1967, p. 354), Wenlock Edge (Shergold and Shirley 1968, p. 135),
and in text-fig. 2 of this paper. This apparent distinction between the faunas is prob-
ably due to breaks in deposition or slow deposition between the main stratigraphical
units. There is a widespread conglomerate at the junction of the Bringewood and
Leintwardine Beds, a frequently developed phosphatized fragment-bed at the
Leintwardine-Whitcliffe junction and the Ludlow Bone-Bed where the Whitcliffe
Beds join the Downton Castle Sandstone.
The term ‘assemblage’ is here used in its most general sense to denote those fossils
which tend to be found together in the rocks, without drawing any conclusions about
the life-assemblages from which they might have been derived or implying any
statistically proved separation from an adjacent assemblage. The scope of the word
‘together’ is also important. If only four assemblages are to be recognized in a shelf
thickness of over 360 m of Ludlow rocks then the average thickness per assemblage
is at least 90 m. If the thickness examined is limited to about 20 m then the collection
of fossils occurring ‘together’ has a different composition ; these are here called ‘minor
512
PALAEONTOLOGY, VOLUME 18
assemblages’. If the thickness is restricted to one or two metres the fossil composition
is again different and much more limited ; these are here called faunal units and are,
perhaps, the associations of Calef and Hancock (1974, p. 796). The signihcance of
these distinctions is discussed after the assemblage lists.
It must be made clear that no particular significance is claimed in this paper for the
four major assemblages described. Calef and Hancock, however, claim that there are
four successive intergrading communities related to depth. It is here agreed that four
successive assemblages are present and this contention is supported by records from
previous papers but it is maintained that there are substantial differences in com-
position from the community lists of Calef and Hancock. It is further maintained
that only one pair of assemblages intergrade and that their ecological significance is
more complex than Calef and Hancock realize. It is agreed that the highest assemblage
almost certainly represents much shallower water than the lowest and earliest
assemblage but the two intermediate assemblages are more complex and contain
subdivisions of considerable palaeoecological significance. The changes in the major
assemblages may be due to some major environmental factors, such as changes in
late Silurian palaeogeography causing restriction of seas and closing and opening
of connections to other regions.
The major assemblages listed below have been named after two characteristic
genera— not necessarily the most abundant. Although generic names change, they
have been preferred to species names which are more likely to be duplicated (e.g.
ludloviensis or lewisii). Ideally, both generic and specific names should be used as
different species of the same genus may have different ecological preferences. This
procedure would make the titles of the assemblages cumbersome and has not been
adopted. However, species names have been given in the assemblage lists. The lists
have been compiled from faunal lists from the following eight areas on the shelf:
May Hill (Lawson 1955), Usk (Walmsley 1959), Woolhope (Squirrell and Tucker
1960), Malverns (Phipps and Reeve 1967; Penn 1969), Aymestrey (Lawson 1973),
Ludlow (Holland, Lawson and Walmsley 1963), Leintwardine (Whitaker 1962), and
Wenlock Edge (Shergold and Shirley 1968). Three points have been allocated to
a fossil recorded as common, two for fairly common, and one for present, giving a pos-
sible top score of twenty-four for ‘commonness’. Because the major Ludlow divisions
each contain two or three subdivisions, usually with separate recordings of species
abundance, average values have had to be taken resulting in non-integers. The
figure after the stroke (maximum eight) indicates the number of areas of occurrence. I
Only benthonic forms are listed, i.e. graptolites and cephalopods are omitted. These
lists differ in intent from those of Calef and Hancock in that non-brachiopod benthos
is listed (indicated by an asterisk), and often seems of greater importance than they
allow. Fossils collected from the limestones as well as the terrigenous sediments are
included and are considered to be essential if an over-all picture of Ludlow palaeo-
ecology is required. The brachiopod contents of these assemblages are sufficiently
similar to those of the communities of Calef and Hancock to invite closer com- |
parison. A more detailed examination, however, reveals some important disparities, ,
which are briefly discussed. I
The author’s name is provided only at the first mention of a species but, in the I
interests of clarity, generic names are repeated in subsequent lists. Text-fig. 2 provides !•
LAWSON: LUDLOW BENTHONIC ASSEMBLAGES
513
a graphic presentation of these faunal changes. Most of the important fossils are
figured in Holland, Lawson and Walmsley (1963, pis. 3-7), and in Calef and Hancoek
(1974, pi. 106).
BENTHONIC ASSEMBLAGES
A. Dicoelosia-Skenidioides assemblage
1.
Aegiria grayi (Davidson)
14-3/8
*2.
Dalmanites myops (Konig)
13-6/8
Hemsiella maccoyana (Jones)
12-2/7
4.
isorthid
10-7/7
5.
Atrypa reticularis (Linnaeus)
9-7/8
6.
Craniops implicata (J. de C. Sowerby)
9-4/8
7.
Shagamella ludloviensis Boucot and Harper
8-0/8
8.
Howellella elegans (Muir-Wood)
8-0/6
9.
Mesopholidostrophia sp.
6-9/6
10.
Dicoelosia biloba (Linnaeus)
6-3I6
11.
Protochonetes minimus (J. de C. Sowerby)
6-3/5
*12.
Calymene sp.
6-1/8
13.
Leptaena depressa (J. de C. Sowerby)
5-9/6
14. 1
1 Strophonella euglypha (Hisinger)
5-8/6
15.J
1 Amphistrophia funiculata (M’Coy)
5-8/6
*16.
Leonaspis sp.
5-5/7
17.
Skenidioides lewisii (Davidson)
5-2/6
18.1
' Leptostroplna filosa (J. de C. Sowerby)
5-0/5
19. J
1 Eospirifer spp.
5-0/5
20.
Sphaerirhynchia wilsoni (J. Sowerby)
A-116
*21.
proetid
4-6/6
22.
Dalejina cf. hybrida (J. de C. Sowerby)
4-5/4
23.
Orbiculoidea rugata (J. de C. Sowerby)
4-4/5
24.
Gypidula cf. galeata (Dalman)
4-3/6
25.
Glassia sp.
4-1/5
26.
Coolinia pecten (Linnaeus)
4-0/5
This list compares well with the Dicoelosia eommunity of Calef and Hancock but
contains four trilobites and one ostracod. As they indicate, it is a high-diversity, low-
density assemblage of predominantly small brachiopods. Dalejina, Skenidioides,
Nucleospira, Cyrtia, and Leangella are more important in their list, perhaps beeause
seven (maybe eight) of their nine Dicoelosia localities are in the Lower Elton Beds
where these forms are commoner. The Eltonian succession has, therefore, been
sampled very unevenly by Calef and Hancock.
B. Strophonella-Gypidula assemblage
1.
Atrypa reticularis
20-5/8
2.
Strophonella euglypha
19-8/8
3.
Leptaena depressa
19-3/8
4.
Sphaerirhynchia wilsoni
15-5/8
5.1
1 Gypidula lata Alexander
15-0/8
6.J
Leptostroplna filosa
15-0/8
7.
Shagamella ludloviensis
14-0/8
8.
Isorthis orbicularis (J. de C. Sowerby)
13-8/8
*9
solitary trochoid coral
13-5/8
514
PALAEONTOLOGY, VOLUME 18
*10.'
Poleumita globosa (Schlotheim)
11-3/8
♦11. J
Dalmanites myops
11-3/8
12.
Howellella elegans
11-0/8
13.
Amphistrophia funiculata
10-3/8
14.
Mesopholidostrophia sp.
10-3/7
♦15.
Hemsiella maccoyana
10-0/7
16.
Camarotoechia micula (J. de C. Sowerby)
9-5/8
17.
Craniops implicata
9-5/8
♦18.
Favosites spp.
9-3/8
19.
Shaleria sp. nov.
8-5/7
20.
Coolinia pecten
l-Sj!
♦21.
Cypricardinia spp.
7-5/7
♦22.
PtUodictya spp.
1-516
23.
Kirkidium knightii (J. de C. Sowerby)
6-5/5
24.
Eospirifer spp.
6-3/8
♦25.
Protochonetes ludloviensis Muir-Wood
6-3/7
26.
Encrinmus sp.
6-3/6
♦27.
Calymene sp.
6-0/8
28.
Aegiria grayi
6-0/6
29.
Dayia navicula (J. de C. Sowerby)
5-5/6
30. 1
Protochonetes minimus
5-0/6
♦31.
Halysites sp.
5-0/6
♦32.
Rhabdocyclus porpitoides (Lang and Smith)
5-0/4
This is a diverse, high-density assemblage dominated by brachiopods, particularly
strophomenids, but with quite a significant variety of other groups, i.e. trilobites,
corals, bivalve, gastropod, ostracod, and bryozoan. It most closely resembles the
Isorthis community of Calef and Hancock but the order of abundance is very different.
At least two of their samples are from Elton Beds which may explain the records of
Dalejina, Homeospira, Glassia, and Skenidioides.
C. Dayia-Isorthis assemblage
1.
Camarotoechia nucula
21-5/8
2.
Dayia navicula
17-8/8
3.
Protochonetes ludloviensis
16-8/8
4.
Sphaerirhynchia wilsoni
14-8/8
5.
Isorthis orbicularis
14-6/8
6.
A trypa reticularis
14-2/8
1.]
1 Shaleria ornatella (Davidson)
13-3/8
8.J
I Shagamella ludloviensis
13-3/8
9.
Howellella elegans
13-0/7
10.
Salopina lunata (J. de C. Sowerby)
11-7/7
11.
Leptaena depressa
10-5/8
12.
Whitfieldella canalis (J. de C. Sowerby)
10-3/7
13.
Craniops implicata
10-0/8
14.
Orbiculoidea rugata (J. de C. Sowerby)
9-5/8
♦15.
Bythocypris siliqua (Jones)
9-2/6
♦16.
Sedgwickia [Fuchsella] amygdalina (J. de C. Sowerby)
8-9/8
♦17.
Calymene neointermedia (R. & E. Richter)
8-4/8
18.
Lep tostrophia filosa
8-3/7
19. i
1 Aegiria grayi
7-7/7
♦20. J
1 Pteronitella retrqflexa (Wahlenberg)
7-7/7
♦21.
Goniophora cymbaeformis (J. de C. Sowerby)
7-6/7
♦22.
Serpulites longissimus (J. de C. Sowerby)
7-5/8
LAWSON: LUDLOW BENTHONIC ASSEMBLAGES
515
*23. Hemsiella maccoyana 7-2/6
*24. Neobeyrichia torosa (Jones) 6-7/7
*25. proetid 6-2/8
*26. //ov<7;7 (J. de C. Sowerby) 6-1/7
*27. Cyclonema corallii 5-2/6
28. Lingula lewisii 5-0/6
This is a diverse, high-density assemblage, still with brachiopods dominant but the
non-brachiopod benthos (ostracods, trilobites, bivalves, gastropods, worm) becoming
important in the ‘second division’. The brachiopod component compares most
closely with the Sphaerirhynchia community of Calef and Hancock. Their list omits
Shaleria and Aegiria, presumably because the Upper Leintwardine Beds were not
sampled at all. It includes, however, Mesopholidostrophia, Gypidula, Strophonella,
and Amphistrophia^-genera. which do not, in fact, occur with an abundance of Dayia,
Protochonetes, and Salopina.
D. Protochonetes-Salopina assemblage
1.
Protochonetes ludloviensis
22-5/8
2.
Camarotoechia nucula
21-5/8
3.
Salopina lunata
20-0/8
*4
Sedgwickia [Fuchsella] amygdalina
16-7/8
*5.
Serpulites longissimus
15-0/7
*6.
Neobeyrichia torosa (Jones)
12-3/8
*7.
Pteronitella retroflexa
10-8/6
*8.
Goniophora cymbaeformis
10-3/6
Cornulites serpularius Schlotheim
9-7/8
10.
Orbiculoidea rugata
9-5/6
*11.
Nuculites spp.
9-0/6
12.
Craniops implicata
8-0/6
13.
Howellella elegans
7-3/7
*14.)
Loxonema spp.
5-8/6
*15.j
Tentaculites tenuis (J. de C. Sowerby)
5-8/6
16.
Dayia navicula
5-7/5
*17.
Bythocypris siliqua
5-7/4
This is a low diversity, high-density assemblage with three brachiopods dominant
but ten of the seventeen commonest fossils are not brachiopods. Bivalves are parti-
cularly important. This fauna corresponds reasonably closely with the Salopina
community of Calef and Hancock in its brachiopod content but they report Sphaeri-
rhynchia and Dayia as being prevalent. This may be due partly to their four samples
from the atypical Llandeilo-Llandovery area and partly to their three Leintwardinian
localities out of a total of twelve localities. It is difficult to comprehend why these
seven samples should be grouped with the five from the Whitcliffian in the first place.
Text-fig. 2 presents the same data in graphic form but with more detailed informa-
tion on the variation in abundance of the various taxa with time. The following points
should be noted :
1. The four assemblage-zones (or, perhaps, concurrent-range zones) correspond
with the four main divisions of the Ludlow Series into Elton, Bringewood, Leint-
wardine, and Whitcliffe Beds.
2. The chart is confined to benthonic forms so that the graptolites and the cephalo-
pods are important absentees. Because of the more refined presentation of the vertical
ASSEMBLAGE
FOSSILS
DIcoelosia -
Skenidioides
Strophonella •
Gypidc'la
Doyia -
Isorthis
Protochonetes -
Salopina
Dicoelosio bilobo
Leonaspis sp.
isorthid
Dalejina cf. hybrida
Skenidioides lewisi i
Glassio sp.
Gypidgla cf. galeata
Profochonefes minimus
Coolinia pecfen
Eospirifef spp.
Halysites sp.
Sholerio sp. nov.
Gypidula lato
Kirkidium knight ij
Dalmanites myops
Strophonella euglypha
Amphistrophio funiculata
Mesopholidostrophia
Poleumito globoso
Favosites spp.
solitary trochoid corals
Rhabdocyclus porpitoides
Cypricardinia spp.
Ptilodictya spp.
Hemsiella maccoyana
Leptostrophia filosa
Sphaerirhynchia wilsoni
Whitfieldella canalis
Neobeyrichia lauensis
Aegiria grayi
Atrypo reticularis
Leptaena depressa
Isorthis orbicularis
proetids
Encrinurus spp.
Shagamella ludloviensis
Bembexia Iloydii
Shaleria ornatella
Calymene neointermedia
Doyio navicula
Lingula lata
C rani ops implicata
Howellella elegans
Calymene spp.
Orbiculoidea rugata
Lingula lewisii
Camarotoechia nucula
Pteronitella retroflexa
Sedgwickia amygdalina
Goniophoro cymbaeformis
Nuculites spp.
Cornulites serpularius
Tentaculites spp.
Loxonemo spp.
By thocypris si 1 iqua
Protochonetes ludloviensis
Cyclonema corallji
Salopina I una ta
Neobeyr ichio torosa
Serpulites longissimus
TEXT-FIG. 2. Range chart of benthonic fossils in the Ludlow rocks of the Welsh Borderland shelf facies:
based on records from May Hill, Usk, Woolhope, Malvern, Aymestrey, Ludlow, Leintwardine, and
Wenlock Edge (see p. 515 for discussion).
LAWSON: LUDLOW BENTHONIC ASSEMBLAGES
517
variation, the stratigraphically important fossils Neobeyrichia lauensis and Lingula
lata become eligible for inclusion on this chart, although not in the assemblage lists.
3. Most of the species and almost all the genera have longer ranges and different
abundance patterns outside this region ; in other words, these are mostly local ranges
and local acmes due to ecological controls.
4. The common constituents of these assemblages do not necessarily enter and
depart, or wax and wane, together, e.g. Howellella elegans is prevalent in four
assemblages, Atrypa reticularis in three, Salopina lunata in two, Shaleria ornatella
in one, whilst Kirkidium kniglitii and Dicoelosia biloba are dominant only for part of
an assemblage. This suggests that these species are not all reacting to one ecological
factor, such as depth of water— as is implied in Calef and Hancock’s account. It also
seems inappropriate to group species of varying tolerances in the same community;
the components of a community should come and go together.
5. There is quite a degree of lateral faunal variation not evident from this chart
of vertical abundance; the main contrast is between the inner and outer shelf areas.
There are obviously important differences between these two versions of the four
major Ludlow benthonic assemblages— even if the comparison is restricted to the
brachiopod content. At first sight, it might be thought that the conclusions of Calef
and Hancock have the greater objective validity as they have described the com-
munities statistically, counting all macrofossils in collections of 100-200 specimens.
Their records are therefore more objective and precise than the familiar categories
of ‘common’, ‘fairly common’, and ‘present’. On the other hand, presumably because
of the time involved in rock-splitting and counting, only 53 localities were collected —
usually represented by single beds or up to 20 cm of rock. This means a possible
maximum thickness sampled of 10-6 m in a succession at least 360 m thick, i.e. a per-
centage of only 3-4 in a series of rocks characterized by many vertical and lateral
facies and faunal changes. The four major communities are based on only 44 localities,
an average of 1 1 per community. Of these 44, 14 are from the Sawdde Gorge which is
just one of the four main sections in the Llandovery-Llandeilo area, where the shelf
facies is atypical.
In contrast, in the eight areas from which the alternative lists have been compiled,
a total of 2600 Ludlovian localities has been examined— an average of 325 per area.
Although it is improbable that any single bed was collected and studied as thoroughly
as those examined by Calef and Hancock it is certain that a large percentage of these
2600 localities were studied bed by bed in order to establish the faunal succession
and subdivisions and to delimit accurately the boundaries between them.
A close look at the locality list in their Appendix (p. 810) reveals some important
differences in faunal records between Calef and Hancock and previous authors.
For instance, they record three examples of Isorthis communities from the Leint-
wardine Beds of Ludlow but Holland, Lawson and Walmsley (1963) did not record
any occurrences of the supposedly prevalent Isorthis community fossils Meso-
pholidostrophia spp., Dalejina, and Amphistrophia in their Leintwardine Beds. The
collections from these three localities have now been examined by the present author
at the British Museum (Natural History), by courtesy of Dr. L. R. M. Cocks. Locality
Lud 2 evidently yielded abundant Kirkidium knightii and is undoubtedly in Upper
518
PALAEONTOLOGY, VOLUME 18
Bringewood Beds and not Leintwardine Beds; it is hardly a very good example of
an Isorthis community in that 7 out of the 10 prevalent fossils are missing, the most
notable absence being Isorthis itself with its statistically assessed presence percentage
of 100. Lud 7 and Lud 8 are, however, correctly assigned to the Leintwardine Beds
but are again not very convincing representatives of the Isorthis community. Indeed,
Lud 7 yielded only 4 of the 10 prevalent forms of the /sorz/jw community but contained
8 of the 10 prevalent fossils of the Sphaerirhynchia community. This locality is Sunny-
hill Quarry, which has recently been studied in detail by Miss Lesley Cherns of
Glasgow University. She reports that there are about 15 m of Leintwardine Beds at
this exposure of which only 20 cm were sampled by Calef and Hancock (i.e. 1-33%).
At this level in the quarry successive bands are dominated by different fossils, e.g.
Isorthis, Sphaerirhynchia, Dayia, and Shagamella. Calef and Hancock evidently
struck a band rich in Isorthis but if they had collected a metre above or a metre
below they might well have hit a Sphaerirhynchia band, and allocated their col-
lection to that community. Indeed, Miss Cherns has studied another locality in the
Leintwardine Beds of the Ludlow area (4619 7360) where, in a thickness of 3 m,
the four community index fossils Lingula, Salopina, Sphaerirhynchia, and Isorthis
are all very common, taking it in turn to dominate different bands. Such a faunal
pattern might be expected to instil some doubt in the mind of even the strictest devotee
of the depth-community religion.
Equally anomalous is the record of a Sphaerirhynchia community from the Lower
Perton Beds of Woolhope (i.e. Lower Whitcliffe Beds). The prevalent fossils of this
community include Sphaerirhynchia wilsoni (with a presence percentage of 100),
Whitfieldella and Leptostrophia filosa, none of which are recorded from their Lower
Perton Beds by Squirrell and Tucker. This collection has also been examined by the
present author and a list of fossils, with numbers present, was submitted to Dr. E. V.
Tucker for his expert opinion. He places the fauna in his lowest Lower Bodenham
Beds (Lower Leintwardine Beds). He also points out that the locality map reference
given by Calef and Hancock appears to indicate a collection from the southern face
of the extended Perton Quarry where Perton Beds do not occur at all.
The above lists are supported by less complete evidence in the following publica-
tions on the areas of Usk (Squirrell and Downing et al. 1 969), Church Stretton (Greig
et al. 1968), Tites Point and Newnham (Cave and White 1971), and Gorsley (Lawson
1954).
The Llandovery-Llandeilo district has been excluded from this analysis as it repre-
sents an unusual and ‘sandy variety of the shelf facies’ (Potter and Price 1965, p. 396)
and the faunas display variations which may relate to the sandy, shallow-water facies.
Calef and Hancock, however, include fourteen localities from this area in their com-
munity analysis based on fifty-three Ludlow localities and this may explain some of
the peculiarities in the associations recorded by them. Presumably because of their
commitment to brachiopod communities they fail to recognize what is probably the
most significant assemblage palaeogeographically in this area. This is a strong
molluscan fauna which Potter and Price (1965, p. 390) considered to be ‘well adapted
for sandy, shallower and possibly less saline conditions’. It occurs in sandstones of
middle Ludlow age and includes the bivalve genera Grammysia and Modiolopsis and
the gastropods Loxonema, Platyschisma, and Bucanopsis. Lingula and Orbiculoidea
LAWSON: LUDLOW BENTHONIC ASSEMBLAGES
519
also occur in association with this fauna, which resembles that of the Downton Castle
Sandstone and also the persistent Palaeozoic linguloid-molluscan community
described by Bretsky (1969) as characterizing near-shore sandy and silty environ-
ments.
Although Calef and Hancock (1974, p. 779) refer to shelly faunas occurring in the
basin they record collections only from Builth Wells (four localities) and Denbighshire
(one), and none from Clun Forest, Knighton, Long Mountain, and Radnor Forest.
They would certainly not have found a simple succession of their four shallowing
benthonic communities. In many places, as at Bishop’s Castle, the Dicoelosia-
Skenidioides assemblage is well developed at the base of the Ludlow succeeded by
Diversograptus nilssoni shales. Above follows a Dayia-Isorthis fauna with some
elements of the Strophonella-Gypidula fauna in the more calcareous siltstones (e.g.
Gypidula, Poleumita, and Favosites at Builth). Then succeeds a normal Dayia-
Isorthis fauna followed by shales with Lingula lata and Saetograptus leintwardinensis.
This latter association is very interesting when it is recalled that in the upper Llandovery
the Lingula community is in the shallowest belt and the graptolitic beds in the deepest
belt. Above this fauna comes the distinctive Aegiria grayi-Neobeyrichia lauensis
assemblage, followed by a Dayia-Protochonetes-Fuchsella assemblage and then
a typical Protochonetes-Salopina assemblage as on the shelf. This is only a very
generalized pattern covering a large area but it serves to demonstrate the limitations
of a palaeoecological interpretation based only on brachiopod communities and also
includes some distinctive associations not recognized by Calef and Hancock.
ASSEMBLAGES AS COMMUNITIES
If four major benthonic assemblages are worthy of recognition in the shelf Ludlow
it is here maintained that the above lists are a more accurate record of the associations
of the fossils than are the lists of Calef and Hancock, even allowing for their restric-
tion to brachiopods. The palaeoecological significance of these major assemblages
must, however, be questioned. Are they, for instance, communities?
In its normal, non-biological usage the term ‘community’ suggests that the com-
ponents have something in common, some interdependence, which results in
a nucleated gathering. In palaeontology the term has come to be used quite commonly,
as in Calef and Hancock’s paper for a completely intergrading life-assemblage. It
consists of a number of species inhabiting the same area at the same time. In the case
of the major shelf assemblages listed above it should be realized that the forms
recorded occur over an area of 12 000 sq km and each assemblage spans at least
30 m of strata, i.e. at least one million years of time. The same applies to the com-
munities of Calef and Hancock. They might claim that this is not a serious objection
if one is concerned with the regional palaeogeographical picture. There are, however,
lateral and vertical faunal variations within these major assemblages which may have
greater palaeoecological and palaeogeographical significance than the differences
between the major faunal units. An example of significant lateral variation occurs in
the Dayia-Isorthis assemblage of Leintwardinian age; the eastern shelf is charac-
terized by the common occurrence of Protochonetes ludloviensis and Salopina lunata
in the siltier near-shore shallow-water facies (Holland and Lawson 1963, p. 287)
520
PALAEONTOLOGY, VOLUME 18
whereas the shelf-edge area, muddier and perhaps deeper, shows a reduced abundance
of these species and the increased importance of Dayia navicula and Shagamella
ludloviensis. Vertical variation is well illustrated in the Bringewoodian of the western
shelf where the Strophonella-Gypidula assemblage can be divided into a lower
Amphistrophia funiculata fauna and an upper Kirkidium-Favosites fauna. Are these
minor assemblages communities in the sense that the constituent species inhabited
the same place at the same time -in the same depth of water? It is doubtful, for
Newall (1966) has subdivided the Kirkidium-Favosites fauna into three units of
palaeoecological significance viz. ;
1. Atrypa-Strophonella units formed in conditions of least turbulence.
2. Coral units of tabulate coral colonies formed in the shallow photic zone in
conditions of fairly high turbulence.
3. Kirkidium units formed in a high-energy environment and possibly within the
breaker zone.
Such faunal units, of depth significance, are completely masked by being lumped
together in major assemblages or communities. Even the units mentioned above,
usually several feet thick, may benefit from refinement. Contrary to the statement by
Calef and Hancock (1974, p. 780), Ludlow fossils commonly occur in bands, often
dominated by particular fossils. Studies of these bedding-plane assemblages would
probably repay study; there may even be more than one community on one bedding
plane ! In the Leintwardinian of the eastern shelf successive bedding planes are often
dominated by Isorthis, Sphaerirhynchia, and Protochonetes with Salopina in turn; it is
surely too much to postulate depth changes every few centimetres through the
succession to explain the repetition of Calef and Hancock’s communities.
The ultimate degree of refinement is to investigate the palaeoecology of the
individual species, paying particular attention to its relationship to the sediment
and to the possible functional significance of some of its morphological characters.
Mr. John Hurst, of Oxford University, has already derived significant results from
some such studies on Silurian brachiopods (Fiirsich and Hurst 1974).
BRACHIOPOD COMMUNITIES
The community tables published by Calef and Hancock ( 1 974, p. 783) are based solely
on the brachiopod fraction of the fauna for two reasons: (1) brachiopods generally
make up at least 90% of the total fauna, (2) the taxonomic uncertainty is less with
brachiopods than with most other groups. This second reason is particularly uncon-
vincing as the trilobites and ostracods have been quite well studied and even the
negleeted groups, such as corals, bivalves, gastropods, worms, bryozoa, can often
yield information of palaeoecological significance in spite of their nomenclatorial
impreeision. The first point, on the dominance of brachiopods, can be seen to be well
justified from the lists published here. Nevertheless, non-brachiopods are evidently
not unimportant. Indeed, in the Protochonetes-Salopina fauna here listed, ten out
of the seventeen fossils are not brachiopods, bivalves being particularly important.
It has already been pointed out that there are important coral units on the main shelf
LAWSON: LUDLOW BENTHONIC ASSEMBLAGES
521
and a bivalve assemblage in the Llandeilo-Llandovery area during Bringewoodian
times.
It must, however, be appreciated that the study of brachiopod communities has
yielded important ideas on Silurian palaeogeography in recent years. It would be
interesting to see to what extent the study of the non-brachiopods will confirm,
refine, or contradict these ideas. Corals, stromatoporoids, and algae should certainly
be helpful as depth-indicators particularly in the Wenlock and in the carbonate
developments. It should be emphasized in this respect that Calef and Hancock’s
study is restricted to the clastic rocks.
DEPTH COMMUNITIES
Calef and Hancock wisely refer to depth-rc/fl/ct/ communities rather than depth-
controlled communities. It is difficult to understand how depth can directly control
the distribution of organisms in the sea. Nevertheless, most of the controlling factors
normally vary with depth— some directly, such as pressure, light, and temperature
and some less inevitably such as substrate, sedimentation, turbulence, salinity, and
food supply. Muddy substrate and still water are commonest at greater depth but
are not uncommon in shallow water; hence the need for caution.
The depth-patterns plotted for the upper Llandovery (Ziegler 1965) nevertheless
seem convincing proof of the depth-relationship of the communities. Even here
there is need for some caution as a progression from onshore to offshore does not
always correlate with increasing depth. Indeed, in the case of the middle Ludlow,
Alexander (1963, pp. 111-112) adduced evidence that the shell-banks of Kirkidium
accumulated on a shelf-edge ridge, i.e. in very shallow water even though far off
shore.
Calef and Hancock do not, however, produce such depth-pattern maps for the
Ludlovian, to demonstrate their communities succeeding each other laterally and
basinwards at particular times. The main reason for this (Hancock, pers. comm.) is
their uncertainty about precise time-correlations in the Ludlow rocks of the Welsh
Borderlands. Presumably, they require lineage zones such as have been established
for the upper Llandovery based on the evolution of Eocoelia, etc. These zones did
not, however, prove the established graptolite zones to be inadequate or diachronous
and it is therefore not clear why the widespread graptolite zones of Diversograptus
nilssoni and Saetograptus leintwardinensis are not acceptable in the Ludlovian. If the
correlation by Holland, Lawson and Walmsley (1963, p. 150, Table 2) is followed,
Calef and Hancock’s communities can be plotted for each of the stages of the Ludlow.
No clear patterns emerge, partly because more data are needed and partly because
single communities tend to spread over most of the shelf, perhaps because the slope
was much more gentle than in the Llandovery. There are also some puzzling anomalies.
In the lower Eltonian the south-eastern inkers of Usk, May Hill, and Woolhope
display a Dicoelosia community whereas the further offshore area of Wenlock Edge
has a ‘shallower’ Isorthis community. In the Bringewoodian the Isorthis community
occurs at Ludlow and Wenlock Edge but the ‘deeper-water’ Dicoelosia community
is reported from May Hill, which is well on to the shelf. The Leintwardinian plots
show an equal mixture of Sphaerirhynchia and Salop ina communities at Usk and
522
PALAEONTOLOGY, VOLUME 18
May Hill— apparently completely, not merely marginally, overlapping. The Isorthis
community is reported from Ludlow, which is indeed further offshore in the tradi-
tional interpretation. In the Whitcliffian, the Salopina community is widespread,
occurring at Usk and May Hill on the inner shelf, at Ludlow on the outer shelf, and
at Builth in the basin. At Woolhope, however, on the inner shelf a Sphaerirhynchia
community is recorded. The direct interpretation of these communities in terms of
depths therefore results in inconsistent and confusing patterns.
Calef and Hancock state (1974, p. 797) that ‘no good correlation has been seen
between sediment type and community within the clastic facies covered by this paper’.
This is contrary to the experience of previous workers who have felt compelled to refer
informally to the "Dicoelosia mudstones’ (actually fine siltstones), the ‘strophomenid
siltstones’, the "Dayia shales’, and the 'Chonetes flags’. It would be interesting to
know whether the Dicoelosia community of Calef and Hancock has ever been found
other than in fine olive siltstones with irregular bedding.
Nevertheless, the suggestion that the Salopina community normally inhabited
shallower water than the Dicoelosia community is not disputed. Also, Calef and
Hancock’s use of density and diversity indices to interpret depths is a welcome new
approach, to be used with caution.
CONTINUOUS REGRESSION
Calef and Hancock contend that the upward Ludlow succession represents a single
regression and (1974, p. 800) ‘have found no evidence of widespread cyclic trans-
gressions and regressions such as those postulated by Phipps and Reeve (1967, fig. 6)
for the Malvern Hills area’. It is here maintained that there is adequate evidence from
both the sediments and the fauna that the pattern figured by Phipps and Reeve is the
regional picture for the shelf area. The Main Outcrop (Wenlock Edge to Aymestrey)
confirms this. The Dicoelosia mudstones of the Lower Elton Beds obviously accumu-
lated in still water with a muddy substrate; the high faunal diversity and low density
lead Calef and Hancock to the conclusion that the water was relatively deep. This
seems quite acceptable.
The succeeding Middle Elton Beds are characterized by graptolites and orthocones
with a very small benthonic fauna. It has usually been considered that these deposits
probably represent a further deepening of the sea. Calef and Hancock record
a Visbyella community from the Middle Elton Beds of Ludlow and presumably
agree on this continued deepening (perhaps to 1000 or 1500 m according to Hancock,
Hurst and Fursich 1974) rather than a regression. The graptolitic Upper Elton Beds
contain slumps and few benthonic forms. They pass up into the richly benthonic
Strophonella-Gypidula calcareous siltstones of the Lower Bringewood Beds which are
succeeded by the Kirkidium-Favosites limestones of the Upper Bringewood Beds.
Newall (1966) has concluded, from detailed palaeoecological studies, that the
tabulate corals lived in moderately turbulent water and that the Kirkidium banks
were probably within the breaker zone. Cross-bedding is fairly common (Whitaker
1962, p. 339) and is indicative of current action. Lawson has found algal remains in
these beds at Aymestrey (Elliott 1971) suggesting water no deeper than 30 m. These
indications of extreme shallowing are confirmed by the widespread occurrence of
LAWSON: LUDLOW BENTHONIC ASSEMBLAGES
523
a limestone conglomerate at the base of the succeeding Lower Leintwardine Beds,
suggesting actual emergence of most of the shelf area. Ooliths have been found at
this level on Wenlock Edge (Shergold and Shirley 1968, pi. 126) and the occurrence
there of the large ostracod Leperditia might be taken to indicate extreme shallowing
as Berdan (1968) suggests that these ostracods were adapted to temporary subaerial
exposure. The dark shales of the Lower Leintwardine Beds at Aymestrey containing
a Dayia navicula-Shagamella ludloviensis sub-fauna plus Saetograptus leintwardinensis
must therefore represent some degree of deepening as postulated by Phipps and
Reeve (1967) not continued regression. This period of emergence in the middle
Ludlow is even more convincingly demonstrated by Potter and Price (1965, p. 398)
in the Llandovery-Llandeilo area where the Old Red Sandstone facies in the Bringe-
woodian Trichrug Beds is succeeded by the fully marine Dayia-Isorthis assemblage
(the Sphaerirhynchia community of Calef and Hancock) of the Leintwardinian.
There is then general agreement on progressive shallowing up through the Whitcliffe
Beds into the Downton Castle Sandstone, with its Lingula-moWusc assemblage.
The regional pattern for the shelf is therefore of two periods of maximum trans-
gression (Middle Elton Beds and Lower Leintwardine Beds) and two periods of
maximum regression (tops of the Bringewood Beds and Whitcliffe Beds), approxi-
mately as depicted by Phipps and Reeve (1967, fig. 6). The recognition of this pattern
raises serious problems for the believers in depth communities. It means that the
same depth of water probably obtained three, or even four, times in the Ludlovian
period and yet there is no repetition of Calef and Hancock’s depth-communities.
The Dicoelosia mudstones, the strophomenid siltstones, and the Dayia shales may
all have been deposited at similar depths and it may have been the difference in sub-
strate (or some other factors) which resulted in the differences in faunal assemblage.
DIACHRONOUS COMMUNITIES
Although Calef and Hancock postulate a succession of regressive benthonic com-
munities they do not explicitly suggest that these are diachronous in the way that the
upper Llandovery communities are. This reticence may be due to their uncertainty
about the precise time correlation of the Ludlow rocks. It has for long been recog-
nized that the shelly faunas are largely facies dependent and the present Ludlow
correlation from basin to shelf has therefore been based on the occurrences of
graptolites, trilobites, and ostracods rather than brachiopods. The internationally
recognizable graptolite zones of Diver sograptus nilssoni and Saetograptus leint-
wardinensis, although best developed in the basin facies, spread well on to the shelf
and interdigitate with the shelly divisions, particularly along Wenlock Edge. Further-
more, at the top of the Leintwardinian there occur not only the highest specimens of
Saetograptus leintwardinensis but also the short-range species Neobeyrichia lauensis
and Calymene neointermedia which occur together at a similar level on the Baltic
island of Gotland.
If this correlation is accepted some of the brachiopod assemblages are seen to be
diachronous. The brachiopods characteristic of the Strophonella-Gypidula assemblage
appear in the Eltonian of the basin but in the Bringewoodian of the shelf. The
Protochonetes-Salopina assemblage is strongly developed in the Leintwardinian of
F
524
PALAEONTOLOGY, VOLUME 18
the southern and eastern shelf but does not reach the basin areas of Kerry and
Knighton until Whitcliffian times. Within the main shelf area diachronism of the
shelly divisions is less easily demonstrated, perhaps because of fairly uniform condi-
tions, including depth, over most of the area.
CONCLUSIONS
The four successive benthonic assemblages here listed for the Ludlow are considered
to give a fuller and more accurate picture of the shelf faunas than the communities
listed by Calef and Hancock, which seem to be based on inadequate sampling and
are inevitably limited by restriction to brachiopods in clastic sediments. The palaeo-
ecological significance of these four major assemblages is not clear. The minor
assemblages, characterizing smaller thicknesses of rock, are likely to be closer to the
life assemblages. The study of the functional morphology and facies preference of
particular species is also a promising approach.
The recent emphasis on depth-communities has led to a neglect of other important,
and more direct, environmental controls, particularly the nature of the substrate.
A consideration of sedimentary evidence demonstrates that Calef and Hancock’s
postulation of continuous regression throughout the Ludlow is unacceptable.
The present correlation of the Ludlow rocks, based mainly on graptolites, trilobites,
and ostracods, is thought to be reasonably sound. Some of the shelly assemblages
are, however, markedly diachronous from shelf to basin but not noticeably so on the
main shelf.
It is concluded that the picture drawn by Calef and Hancock is an over-simplification
resulting, perhaps, from an attempt to impose a relatively straightforward Llandovery
pattern on to the more complex Ludlow rocks.
Acknowledgements. I thank Mr. Nigel Hancock and other members of the Oxford ‘community school’
for the ready access to their writings and for many stimulating and amicable discussions— in spite of the
divergence of our scientific views. Professors T. Neville George and C. H. Holland, and Miss Lesley Cherns
kindly read and criticized the manuscript.
REFERENCES
ALEXANDER, F. E. s. 1936. The AymesUy Limestone of the Main Outcrop. Q. Jlgeol. Soc. Land. 92, 103-1 15.
BERDAN, J. 1968. Possible paleoecological significance of Leperditiid ostracodes. Geol. Soc. Amer. Prog.
Annu. Mtng. North-eastern Sect. Washington D.C., p. 17 (Abstr.).
BRETSKY, p. w. 1969. Evolution of Paleozoic benthic marine invertebrate communities. Palaeogeogr.,
Palaeoclimat., Palaeoecol. 6, 45-59.
CALEF, c. E. and Hancock, n. j. 1974. Wenlock and Ludlow marine communities in Wales and the Welsh
Borderland. Palaeontology, 17, 779-810.
CAVE, R. and white, d. e. 1971. The exposures of Ludlow rocks and associated beds at Tites Point and near
Newnham, Gloucestershire. Geol. J. 7, 239-254.
ELLIOTT, G. F. 1971. A new fossil alga from the English Silurian. Palaeontology, 14, 637-641.
FURSiCH, F. T. and HURST, J. M. 1974. Environmental factors determining the distribution of brachiopods.
Ibid. 17, 879-900.
GREiG, D. c., WRIGHT, J. E., HAiNS, B. A., MITCHELL, G. H., et al. 1968. Geology of the country around Church
Stretton, Craven Arms, Wenlock Edge and Brown Clee. Mem. geol. Surv. U.K. 1-379.
HANCOCK, N. J., HURST, J. M. and FURSICH, F. T. 1974. The depths inhabited by Silurian brachiopod com-
munities. Jlgeol. Soc. bond. 130, 151-156.
LAWSON: LUDLOW BENTHONIC ASSEMBLAGES
525
HOLLAND, c. H. and LAWSON, J. D. 1963. Facies patterns in the Ludlovian of Wales and the Welsh Borderland.
Lpool Manchr geol. J. 3, 269-288.
and WALMSLEY, V. G. 1962. Ludlovian Classification— A reply. Geol. Mag. 99, 393-398.
1963. The Silurian rocks of the Ludlow district, Shropshire. Bull. Br. Mas. nat. Hist.
(Geol.), 8, 95-171.
LAWSON, J. D. 1954. The Silurian succession at Gorsley (Herefordshire). Geol. Mag. 91, 227-237.
1955. The geology of the May Hill inlier. Q. J I geol. Soc. Load. Ill, 85-1 16.
1960. The succession of shelly faunas in the British Ludlovian. C.R. Intern. Geol. Congr. 21st Session,
Nor den, 7, 114-125.
1973. Facies and faunal changes in the Ludlovian rocks of Aymestrey, Herefordshire. Geol. J. 8,
247-278.
NEWALL, G. 1966. A faunal and sedimentary study of the Aymestry Limestone and adjacent beds m parts
of Herefordshire and Shropshire. Ph.D. thesis, Univ. Manchester.
PENN, J. s. w. 1969. The Silurian rocks to the west of the Malvern Hills from Clenchers Mill to Knightsford
Bridge. Ph.D. thesis, Univ. London.
PHIPPS, c. B. and REEVE, F. A. E. 1967. Stratigraphy and geological history of the Malvern, Abberley and
Ledbury Hills. Geol. J. 5, 339-368.
POTTER, J. F. and PRICE, J. H. 1965. Comparative sections through rocks of Ludlovian-Downtonian age in
the Llandovery and Llandeilo districts. Proc. Geol. ^455. Lond. 76, 379-402.
SHERGOLD, J. H. and SHIRLEY, J. 1968. The faunal stratigraphy of the Ludlovian rocks between Craven Arms
and Bourton, near Much Wenlock, Shropshire. Geol. J. 6, 1 19-138.
SQUiRRELL, H. c., DOWNING, R. A. et al. 1969. Geology of the South Wales Coalfield pt. 1. The country
around Newport (Mon.). Mem. geol. Surv. U.K. 1-333.
and TUCKER, e. v. 1960. The geology of the Woolhope inlier (Herefordshire). Q. Jl geol. Soc. Lond.
116, 139-185.
STRAW, s. H. 1937. The higher Ludlovian rocks of the Builth district. Ibid. 93, 406-456.
WALMSLEY, v. G. 1959. The geology of the Usk inlier (Monmouthshire). Ibid. 114, 483-521.
WHITAKER, J. H. MCD. 1962. The geology of the area around Leintwardine. Ibid. 118, 319-351.
ZIEGLER, A. M. 1965. Silurian marine communities and their environmental significance. Nature, Lond.
207, 270-272.
COCKS, L. R. M. and bambach, r. k. 1968. The composition and structure of Lower Silurian marine
communities. Lethaia, 1, 1-27.
J. D. LAWSON
Department of Geology
Typescript received 23 May 1974 University
Revised typescript received 22 November 1974 Glasgow G12 8QQ
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THE TRILOBITE LEJOPYGE HAWLE AND
CORDA AND THE MIDDLE-UPPER
CAMBRIAN BOUNDARY
by B. DAILY and J. B. jago
Abstract. The species and subspecies of the late middle Cambrian agnostid trilobite Lejopyge are reviewed. Lejopyge
cos Opik is shown to be a junior synonym of Lejopyge laevigata armata. In Sweden the middle-upper Cambrian
boundary is placed at the boundary between the Lejopyge laevigata and Agnostus pisiformis Zones. The reassignment
of L. cos to L. 1. armata and other criteria suggest that this boundary in Australia should be drawn within the
Mindyallan Cyclagnostus quasivespa Zone between the L. cos and Blackwelderia sahulosa faunas.
It is suggested that the middle-upper Cambrian boundary in North America be placed well up into the Cedaria
Zone ; in China it is at some as yet undefined position within the Blackwelderia sinensis Zone ; on the Siberian Platform
it should be placed between the zones of Lejopyge laevigata armata- Lomsucaspis alta and Agnostus pisiformis-
' Homagnostus fecundus’ and in north-west Siberia between the zones of Maiaspis spinosa-Oidalagnostus trispinifer
and Acrocephalella granulosa- Koldiniella prolixa.
Various species and subspecies of Lejopyge are important index fossils of the late
middle Cambrian of Sweden (Westergard 1946), Utah (Robison 1964^, b), Queens-
land (Opik 1961a, 1967), Siberia (Demokidov 1968), and Alaska (Palmer 1968).
This paper reviews the status of the species and subspecies of Lejopyge and discusses
the intercontinental correlations arising out of this work. The availability of large
numbers of latex moulds and silicone-rubber casts of trilobites (especially those
illustrated by A. H. Westergard from Sweden), allowed many conclusions to be
drawn which otherwise could not have been made from the published literature.
Order miomera Jaekel, 1909
Suborder agnostina Salter, 1864
Superfamily AGNOSTACEA M’Coy, 1849
Family agnostidae M’Coy, 1849
Subfamily ptychagnostinae Kobayashi, 1939
Genus lejopyge Hawle and Corda, 1847
Synonymy. Hawle and Corda, 1847,p. 51 ; Kobayashi 1937, pp. 437-447; 1939, p. 131 ; Lermontova
1940, p. 130; Westergard 1946, p. 87; Hupe 1953, p. 61; Pokrovskaya 1958, p. 72; 1960, p. 60; Howell
1959, p. 178; Opik 1961a, p. 85; 1967, p. 93; Robison 1964a, p. 521 ; Palmer 1968, p. 27. Miagnostus iaekel,
1909, p. 401.
Type species. Battus laevigatas Dalman, 1828, p. 136.
Discussion. Westergard (1946, p. 87) and Opik (1961a, pp. 76, 85) have discussed
Lejopyge, its species and subspecies, and its relationships with other genera, especially
Ptychagnostus Jaekel. Westergard (1946, p. 75) suggested, and Opik (1961a, p. 85)
agreed, that Ptychagnostus (Triplagnostus) elegans (Tullberg), P. elegans laevissimus
Westergard (PI. 63, figs. 12, 13), andL. /acv/ga/a (Dalman) ‘constitute an evolutionary
series with very small intervals’.
[Palaeontology, Vol. 18, Part 3, 1975, pp. 527-550, pis. 62-63.]
528
PALAEONTOLOGY, VOLUME 18
The following species and subspecies have been included in Lejopyge: L. calva
Robison, L. cos Opik, L. empozadensis Rusconi, L. exilis Whitehouse, L. laevigata
(Dalman), L. laevigata armata (Linnarsson), L. 1. forfex (Brogger), L. 1. perrugata
Westergard, L. 1. rugifera Westergard, L. 1. similis (Brogger), and L. obsoletus
(Kobayashi).
Opik (1961fl, p. 86) suggested that the holotype cephalon of L. exilis belongs in
either L. laevigata or L. 1. armata and that the pygidium of L. exilis figured by White-
house (1936, pi. 9, fig. 12) belongs in either Phalacromal dubium Whitehouse or
Hypagnostus hippalus Opik. This pygidium is very poorly preserved (PI. 63, fig. 11)
and cannot be assigned to any species or genus with certainty. In our opinion the
border is far too wide to include the specimen in L. laevigata. Westergard (1946,
p. 88) suggested that L. 1. similis belongs in Cotalagnostus confusus (Westergard),
and that L. 1. forfex resembles the pygidium figured as L. 1. armata by Westergard
(1946, pi. 13, fig. 31). The pygidium described by Kobayashi (1935) as Agnostus
(Lejopygel) obsoletus was reassigned by him (Kobayashi 1962, p. 30) to Phoida-
gnostus limbatus. L.l controversa Kryskov {in Borovikov and Kryskov 1963) belongs
in Peratagnostus Opik (1967, p. 35). L. ? sugandensis Kryskov was described in
Borovikov and Kryskov (1963, p. 275, pi. 1, fig. 9). However, a footnote (p. 274)
indicates reassignment of sugandensis to Phaldagnostus Ivshin.
Rusconi (1953, p. 5) described a single pygidium as L. empozadensis. He later
redescribed and figured the same specimen (1954, p. 33, pi. 2, fig. 10) as L. empozadense.
As far as can be determined from the figure, this species has a much wider border
than any described species of Lejopyge. The specimen described and figured by
Rusconi (1951, p. 8, fig. 9) as Spinagnostus pedrensis was later assigned by him to
L. pedrensis ( Rusconi 1 953). However, the figure given by Rusconi ( 1 95 1 ) is inadequate
for either generic or specific identification.
Robison (1964a) described L. calva from Utah and Nevada where it is the nominate
species of the youngest of the three subzones of his late middle Cambrian Bolaspidella
Assemblage Zone. Palmer (1968) described L. calva from Alaska. L. calva is more
effaced (PI. 63, fig. 10) than L. laevigata and its subspecies.
Robison (1964a, p. 522) reported the occurrence of an unnamed subspecies of
L. calva from U.S. Geological Survey Collection 2523-CO from Schell Creek Range,
Nevada, characterized by postero-lateral border spines on the cephalon, but not on
the pygidium. A pygidium is figured (PI. 63, fig. 9), but none of the available associated
cephala show undoubted cephalic spines. Cephala and pygidia from a Lejopyge-
coquina from Patterson Pass, Snake Range, East Nevada, are almost entirely effaced
and are figured as Lejopyge sp. (PI. 63, figs. 7 and 8) but may well be representatives
of L. calva.
L. cos was described by Opik (1967, p. 93) from the lower two zones {Erediaspis
eretes and Cyclagnostus quasivespa Zones) of the Mindyallan Stage of north-west
Queensland, which were placed in the upper Cambrian, thus making L. cos the
youngest species of Lejopyge. All other described and authenticated species of
Lejopyge come from late middle Cambrian horizons. As concluded below, we
believe that L. cos is a junior synonym to L. 1. armata and that it is of late middle
Cambrian age.
L. laevigata and L. 1. armata are differentiated on the basis of the latter having
DAILY AND JAGO: LEJOPYGE
529
cephalic and pygidial spines. However, there are small postero-lateral spines on the
pygidium of L. laevigata (Westergard 1946, pi. 13, fig. 25; PI. 62, fig. 10). There are
also short spines on the cephalon of L. laevigata (Westergard 1946, pi. 13, fig. 24;
PI. 62, fig. 2). L. 1. perrugata and L. 1. rugifera were erected by Westergard (1946) for
forms with cephala showing a greater degree of scrobiculation than in either L. laevi-
gata or L. 1. armata. However, some of the cephala of L. laevigata and L. 1. armata
illustrated by Westergard (1946, pi. 13, figs. 22, 35) are scfobiculate to varying degrees
(PI. 62, fig. 3). L. 1. rugifera was differentiated from L. 1. perrugata by Westergard
on the basis of the latter having short cephalic spines with no mention of cephalic
spines in the diagnosis of L. 1. rugifera. The cephalic spines of the holotype of perrugata
are quite large (PI. 63, fig. 1) and the holotype of rugifera also has cephalic spines
albeit short (PI. 63, fig. 6). The pygidia (PI. 62, figs. 12, 13 ; PI. 63, figs. 2-4) associated
with the holotype cephala of rugifera and perrugata are indistinguishable from pygidia
of L. laevigata and L. 1. armata.
Westergard noted the great morphological variation within L. laevigata and also
the presence of intermediate forms between L. laevigata and L. 1. armata, L. 1. per-
rugata and L. 1. rugifera, and between the subspecies (see text-fig. 1). This variation
and the presence of intermediate forms indicate that we are dealing with a species
complex with the subspecies armata, perrugata, and rugifera representing extreme
forms of L. laevigata.
Lejopyge laevigata rugifera
Lejopyge laevigata armata < > Lejopyge laevigata perrugata
TEXT-FIG. 1. Summary of gradations between the species and subspecies of Lejopyge from Sweden. The
arrows indicate the presence of gradational characteristics, which include the over-all shape of the cephalon
and pygidium, the degree of effacement, the width of the pygidial axis, the presence or absence of cephalic
and pygidial spines, the length of spines, and the degree of cephalic scrobiculation.
Opik (1967, p. 93) diagnosed L. cos as follows:
Leiopyge cos sp. nov. is distinguished by well developed posterior section of the cephalic axial furrows
and rather distinct but relatively small basal lobes, short pygidial marginal spines, and two median nodes
on the pygidial axial lobe; the additional node is placed on the anterior axial annulation.
Opik’s differential diagnosis of L. cos is as follows:
The marginal pygidial spines of L. cos are shared by Leiopyge laevigata armata (Linnarsson) but armata
has only one node, on the second axial annulation; furthermore, the cephalic spines of armata are long
(short in cos, as observed on specimens not illustrated).
The specimens figured by Westergard (1946, pi. 13, figs. 28, 29, 30, 31) as L. 1.
armata (Linnarsson) fit the diagnosis of L. cos perfectly. (The anterior of the two
nodes cannot be seen in Westergard’s figures.) The pygidia of armata (Westergard
530
PALAEONTOLOGY, VOLUME 18
1946, pi. 13, figs. 30, 31 ; PI. 62, figs. 15, 16) have nodes on both the first and second
pygidial axial segments in identical positions to the two nodes illustrated on L. cos
by Opik (1967, fig. 20). Close examination of the holotype pygidium of L. cos reveals
the presence of a faint, but distinct, third node placed at about the centre of the third
axial segment (PI. 62, fig. 18). A third node in a similar position is also present on
L. laevigata, L. 1. armata, and on pygidia associated with the holotype cephala of
L. /. perrugata and L. 1. rugifera and the unnamed subspecies of L. calva of Robison
(1964a). Palmer (1968, p. 26) noted that Lejopyge has ‘the posterior axial node on
the axial lobe and not at its terminus, comparable to the position in Ptychagnostus' .
The presence or absence of pygidial nodes and spines on the various species of
Lejopyge is shown in Table 1. Not all pygidia possess a third node; where it is present
it is usually small and faint and is not always visible in the photographs. However,
in some specimens the node is reasonably prominent (e.g. PI. 62, figs. 7, 8; PI. 63,
figs. 3, 9). At least one pygidium of L. laevigata (Westergard 1946, pi. 13, fig. 23;
PI. 62, fig. 7) has an anterior axial node as do some of the pygidia associated with the
holotype cephalon of L. 1. perrugata. In most pygidia not possessing a definite anterior
axial node there is a slight general swelling in the expected position of the node. Thus
the presence or absence of the first or third nodes cannot be used to differentiate
L. cos, L. laevigata, and L. 1. armata. The pygidium of Ptychagnostus elegans
laevissimus (Westergard 1946, pi. 10, fig. 22; PI. 63, fig. 13), the supposed ancestor
of L. laevigata, shows no sign of either a first or a third pygidial node.
L. eos is also similar to L. 1. armata in its pygidial spine characteristics. In this
discussion of spine characters the line diagram of Opik (1967, fig. 20) is referred to
rather than his photograph of the holotype of L. cos (Opik 1967, pi. 57, fig. 5; PI. 62,
fig. 18), because the border is poorly preserved on the holotype and Opik had access
to other unfigured pygidia of L. cos.
Opik (1961a, p. 87; 1967, p. 93) maintained that L. 1. armata has long cephalic
and pygidial postero-lateral spines. However, Westergard (1946, pi. 13, figs. 28-36)
allows great variations in the length of these spines— they vary from quite small
to very long. Westergard (1946, p. 89) also notes, when discussing armata that:
Forms with shorter spines and more or less distinctly furrowed cheeks connect this long-spined
and smooth form on the one hand with the typical laevigata and on the other hand with the subspecies
perrugata.
This is borne out by a cephalon with short spines (PI. 62, fig. 14) which occurs on
the same slab as the pygidia figured as L. 1. armata in Westergard (1946, pi. 13, figs. 30,
31) (see also PI. 62, figs. 15, 16). Further, a cephalon figured as L. /acv/gata (Westergard
1946, pi. 13, fig. 24; PI. 62, fig. 2) has short cephalic spines. The pygidia of L. 1. armata
(Westergard 1946, pi. 13, figs. 30, 31; PI. 62, figs. 15, 16) have quite small spines
which in fact are smaller than those of L. cos (Opik 1967, p. 93, fig. 20). Thus, as far
as cephalic and pygidial spines and pygidial nodes are concerned, L. cos and L. 1. armata
are indistinguishable. The over-all shape of the holotype pygidium of L. cos (PI. 62,
fig. 18) is similar to the shape of many of the pygidia of L. laevigata and L. 1. armata
figured by Westergard (1946). Unfortunately, the only cephalon of L. cos figured by
Opik (1967, pi. 57, fig. 6) (see also PI. 62, fig. 17) is a poorly preserved collapsed
specimen in which the border has not been preserved. Opik’s diagnosis of L. cos
notes the well-developed posterior section of the cephalic axial furrows and the small
DAILY AND JAGO: LEJOPYGE
TABLE 1 . Pygidial characteristics of the species and subspecies of Lejopyge.
531
Designated name
Figuring
Figuring in
Spine
Axial nodes
or association
herein
previous works
characteristics
1
2
3
Other remarks
Lejopyge laevigata
PI. 62, fig. 7
Westergard (1946,
pi. 13, fig. 23)
Absent
7
P
P
5 or 6 pairs of muscle
scars. Associated with
Dalman's syntype
L. laevigata
PI. 62, fig. 4
WestergSrd (1946,
pi. 13, fig. 26)
Absent
A
P
A
At least four pairs of
muscle scars
L. laevigata
PI. 62, fig. 10
Westergard (1946,
pl. 13, fig. 25)
Very small
7
P
P
L. laevigata
PI. 62, fig. 8
Unfigured pygidium
on same slab as
cephalon figured
in Westergdrd
(1946, pl. 13,
fig. 24)
Absent
P
P
P
Faint trace of post-
axial median furrow
L. laevigata
PI. 62, fig. 6
Unfigured pygidium
on same slab as
above specimen
Minute
A
P
P
Faint trace of post-
axial median furrow
L. laevigata
PI. 62, fig. 9
Westergard (1946,
pl. 13, fig. 20)
Absent
A
P
P
L. laevigata
PI. 62, fig. 5
Unfigured specimen
on same slab as
above specimen
Absent
P
P
P
Trace of post-axial
median furrow
L. laevigata
PI. 62, fig, 1
Westergard (1946,
pl. 16, fig. 9)
Absent
A
P
7
Complete specimen
L. 1. armata
PI. 62, fig. 15
Westergard (1946,
pl. 13. fig. 30)
Very small
P
P
7
L. 1. armata
PI. 62, fig. 16
Westergard (1946,
pl. 13, fig. 31)
Very small
P
P
P
Pygidium associated
with holotype
cephalon of L. 1.
perrugata
PI. 63, fig. 3
Unfigured
Present
A
P
A
Broad low ridge
posterior to node 2.
Spines broken —
length indeterminate
Largest pygidium
PI. 63, fig. 2
Unfigured
Present
7
P
P
Very large spine base
associated with
holotype cephalon of
L. 1. perrugata
Pygidium associated
PI. 62, fig. 13
Unfigured
Large spines
P
P
P
with holotype
cephalon of
L. 1. perrugata
Pygidium associated
PI. 62, fig. 12
Unfigured
Present
P
P
P
with holotype
cephalon of
L. 1. perrugata
Pygidium associated
PI. 63, fig. 4
Unfigured
Absent
A
P
P
with holotype
cephalon of
L. 1. rugifera
Holotype pygidium of
L. cos
PI. 62, fig. 18
Opik (1967, pl. 57,
fig- 5)
Present
P
P
P
L. calva
Unfigured
Robison (1964,
pl. 83, fig. 3)
Absent
A
P
A
Strikingly effaced in
both cephalon and
pygidium
U.S.G.S. Collection
2523-CO (unnamed
subspecies of L. calva,
see Robison 1964a,
PI. 63, fig. 9
Unfigured
Absent
7
P
P
Wide border. Not all
pygidia on this
specimen have the
third node
p. 522)
Lejopyge sp. (probably
PI. 63, fig. 8
Unfigured
Absent
A
P
A
All pygidia in these
L. calva) from
coquina, Patterson
Pass, Snake Range.
Nevada
Plychagnostus elegans
laevissimus
PI. 63, fig. 13 Westerg^rd (1946,
pi. 10, fig. 22)
Absent
A P A
specimens are
strikingly effaced
Wide axis
A ^ absent. P present. ? = indeterminate.
532
PALAEONTOLOGY, VOLUME 18
but distinct basal lobes. However, the basal lobes of all species of Lejopyge are small.
The rear part of the cephalic axial furrows of almost all the specimens of L. laevigata
and its subspecies figured in Westergard (1946) and herein also have well-developed
posterior axial furrows.
The facts noted above indicate that L. cos is a junior synonym of L. /. armata.
Another point is that the pygidia of L. 1. armata, as illustrated by Opik (1961a,
pi. 22, figs. 2, 3, 4) presumably have no node on the anterior axial annulation. Whether
this is so or not cannot be clearly determined from the illustrations given by Opik.
If, in fact, there is no anterior node on the Queensland middle Cambrian and Passage
Zone forms, then this may indicate a difference due to geographical variation.
A further point of difference between the Swedish specimens of L. I. armata and
those from Queensland illustrated by Opik (1961a, pi. 22, figs. 2-4) is that in the
Queensland forms the pygidial spines are posterior to those figured by Westergard
(1946).
LEJOPYGE THE MIDDLE-UPPER CAMBRIAN BOUNDARY
Scandinavia
Within the Acado-Baltic province (type province of the Cambrian System) in
both the Oslo region and adjacent parts of Sweden, the Cambrian occurs as very
markedly condensed platform sequences. Although sections of these seemingly
shallow-water deposits contain breaks, the painstaking collection and documentation
of the fossils, mainly trilobites, has allowed a reliable and very fine zonation of the
System, especially for the middle and upper Cambrian. Westergard (1922; 1946,
p. 19; 1947, pp. 20-21) has shown that the most complete sections for the middle and
upper Cambrian in Sweden are in Scania. However, even there breaks of varying
magnitude are evident.
In Scandinavia the middle Cambrian-upper Cambrian boundary is drawn at the
top of the Lejopyge laevigata Zone (see Tables 2 and 3). However, when discussing
the biostratigraphy of the Swedish middle Cambrian, Westergard (1946, p. 7) pointed
out that ‘The boundary is not very well defined, the zone of Lejopyge laevigata
merging into that of Agnostus pisiformis'. There are several reasons why this appears
to be so :
1. In contrast to the rich and varied fauna of the L. laevigata Zone, only eight
trilobite species or subspecies (even one of these questionably) are known in the
A. pisiformis Zone in Sweden; three in Norway, where Olenus alpha Henningsmoen
constitutes a further species. However, the rare O. alpha is unknown outside of the
Ringsaker area (Henningsmoen 1957; 1958).
2. A. pisiformis (Linnaeus) is the only common trilobite in the A. pisiformis
Zone, all other species being generally rare or absent in collections from most
localities where the zone is recognized. The fossils which occur in black bituminous
shales (alum shales) and dark bituminous limestone (stinkstone) were probably
specialized planktonic forms that were able to avoid the poisonous bottom habitat
( Bergstrom c/ a/. 1972).
3. Of the trilobites found in the A. pisiformis Zone only A. pisiformis ranges down
into the L. laevigata Zone (Table 2). Originally Westergard (1947, p. 22) showed
DAILY AND JAGO: LEJOPYGE
533
TABLE 2. Trilobite zonation for the late middle Cambrian-early upper Cambrian of Sweden. The ranges of
the majority of Scandinavian trilobites mentioned in the text are presented to facilitate discussion. The
thicknesses given for the various divisions are taken from Westergard (1944a, p. 29). The unbracketed
figures are for the Andrarum No. 1 borehole; those bracketed are for the Sodra Sandby borehole about
40 km west of Andrarum, Scania. Note that the thickness for the L. laevigata Zone includes the unfossili-
ferous interval 10 m (3 0) immediately below the designated A. pisiformis Zone.
(P
m
g
(p
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m
CP
Acrocephalites stenometopus
Acrocephalites stenometopus agnostorum
Acrocephalites stenometopus olenorum
Agnostus pisiformis
Agnostus pisiformis subsulcatus
Clavagnostus sulcatus
Diplagnostus planicauda vestgothicus
Drepanura eremita
Glyptagnostus reticulatus
Glyptagnostus reticulatus nodulosus
Homagnostus obesus
Hypagnostus sulcifer
Lejopyge laevigata
Lejopyge laevigata armata
Oidalagnostus trispinifer
Olenus alpha
Peronopsis insignis
Phalacroma glandiforme
Phalagnostus bituberculatus
Proceratopyge conifrons
Proceratopyge nathorsti
Ptychagnostus (Goniagnostus) spiniger
Ptychagnostus (Ptychagnostus) aculeatus
Schmalenseeia amphioneura
Solenopleura brachymetopa
MIDDLE CAMBRIAN
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534
PALAEONTOLOGY, VOLUME 18
Acrocephalites stenometopus (Angelin) in the Agnostus pisiformis and Olenus Zones,
but he later (1948) referred forms from each of these zones to the subspecies A. steno-
metopus agnostorum and A. s. olenorum respectively (Table 2). Moreover, he regarded
the middle Cambrian A. stenometopus and its two upper Cambrian subspecies as con-
stituting an evolutionary series which spanned the middle-upper Cambrian boundary.
Thus in Scandinavia and elsewhere, rocks with L. laevigata signify the middle
Cambrian.
4. Where unfossiliferous intervals occur between rocks containing the L. laevigata
and Agnostus pisiformis faunas, there must be an interval of uncertainty concerning
the zonal and series boundaries. In practice, the boundary has been drawn either
immediately above the barren interval (Westergard 1944a, h) or immediately below
it (Westergard 1922, p. 18).
Should a convenient reference section for the middle-upper Cambrian boundary
be required, the section described by Westergard (1922, fig. 33, pp. 67-68) from
Odegarden, Falbygden district in Vastergotland would be suitable, for at that
locality the L. laevigata and A. pisiformis Zones are in contact and the ranges of the
two nominate zonal species overlap. This unbroken section provides an unambiguous
solution to the boundary problem.
The Cambrian world exclusive of Scandinavia
Australia. Since Lejopyge cos Opik is a synonym of L. laevigata armata (Linnarsson),
it is evident that L. 1. armata ranges as high as the Mindyallan Zone of Cyclagnostus
quasivespa (see Opik 1967, Table 4, p. 41). Providing the upper limits of the ranges
of this subspecies are the same in Queensland and Sweden then part of the C. quasivespa
Zone and the top part of the Swedish L. laevigata Zone are correlatives (Table 3).
The described specimens of L. cos came from the Mungerebar Limestone at
locality G 131 in the Zone of C. quasivespa. In the Mungerebar-Mindyalla area dips
are low and outcrops are small and discontinuous so that Opik’s stratigraphic suc-
cession was pieced together on faunal evidence rather than on superposition. This
has led to uncertainties, for example, Opik (1967, vol. 2, p. 9) commented that the
collection from locality G 131 was ‘apparently below G 130’ which among other
species contained Blackwelderia sabulosa Opik. As indicated on the collection ;
locality map (Opik 1967, fig. 3, p. 12), the G 131 site is not far removed from the \
lower boundary of the zonal limits. An analysis of faunal lists from collecting sites |
within the C. quasivespa Zone suggests a clear separation of the G 131 fauna (and its *
presumed equivalent the G 10 fauna, see Opik 1967, vol. 2, p. 6) from those contain- {
ing B. sabulosa (G 124-G 127 ; G 1 30), which as he suggested are presumably younger.
Thus for the Australian region it is advocated that the middle-upper Cambrian |
boundary be drawn within the C. quasivespa Zone between the L. cos{ = L. 1. armata)
and B. sabulosa faunas (Table 3). In passing, we note that in Australia Blackwelderia
was already present in the late middle Cambrian for at locality G 1 19, 5. cf. sabulosa r.i
is found in the zone of E. eretes. Moreover, Blackwelderia succeeds Damesella in i
Australia as in China. In Australia Damesella first appears in the D. torosa-A. janitrix
Zone and D. torosa itself ranges into the E. eretes Zone (Opik 1967, p. 307) where
Blackwelderia is present. '
TABLE 3. Correlation chart of late middle and early upper Cambrian trilobite faunas of Scandinavia and other areas discussed in the text. It should
be noted that the top and bottom lines of the correlation chart have no temporal significance, e.g. it does not indicate that the top of the North
American Aphelaspis Zone corresponds to the top of the Glyptagnostus reticulatus Zone from Australia or that the base of the North American
Bolaspidella Zone is equivalent to the base of the Swedish Solenopleura brachymetopa Zone. Chu (1959) uses the term Damesella paronai Zone
rather than the D. blackwelderi Zone.
DAILY AND JAGO: LEJOPYGE
535
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SIBEFUAN PLATFORM NORTH-WEST SIBERIA
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Notes: 1 and 2. The age of the basal part of the Manceiter Grits and Shales is uncertain and may range within the limits shown. 3 and 4. The
relative positions of two important fossils in the Merevale No. 3 Borehole.
Notes 5 and 6, The lower boundary of the Maiaspis spinosa-Oidalagnosius irispinifer Zone is shown extending to position 5, well into the
Solenopleura brachymetopa Zone. Table 4 which is based mainly on Datsenko et a/. (1968, ‘Atlas’, pp. 28-31) shows that the ranges of Phalacroma
glandi/orme and Oidalagnostus irispinifer overlap. Consequently, if it is conceded that P. glandiforme ranges into the Swedish L. laevigata Zone,
then this boundary will need to be shifted to approximately position 6,
536
PALAEONTOLOGY, VOLUME 18
The vast majority of all the other trilobites listed in Opik’s Table 4 are endemic
species and so have little value for refined intercontinental correlations. However,
a check of the non-endemic forms listed suggests that the correlation proposed
above is correct. The following species of agnostids listed by Opik deserve comment
(reference to Tables 2 and 3 will assist the reader);
1. In Sweden Ptychagnostus {Goniagnostus) spiniger (Westergard) occurs in the
‘Zone of Lejopyge laevigata, basal layer’ (Westergard 1946, p. 82). Opik (1967,
p. 90) reported this species from limestone in the Northern Territory (locality T 87)
and from the Steamboat Sandstone in Queensland (localities G 106 and D 96).
In the discussion of the Australian material Opik (1967, p. 90) stated that P. (G.)
spiniger occurs ‘in the upper part of the L. laevigata II and in the laevigata III Zones’.
Now the L. laevigata III Zone is shown as the uppermost middle Cambrian Zone
in the biostratigraphic chart given by Opik (1961a, %. 15, p. 34). However, from the
Devoncourt Limestone (locality D 18, which is a direct correlative of, or at the most
one zone older than the T 87 fauna cited above) and the older Roaring Siltstone
(locality D 7/15) in Queensland, Opik (1961a, p. 44) reported Ptychagnostus {Ptycha-
gnostus) aculeatus (Angelin), a species which in Sweden is confined to the Solenopleura
brachymetopa Zone. Thus, the positioning of the D 18 fauna on Opik’s chart (Opik
1961a, fig. 15, p. 34) appears to be too high in terms of the Swedish zonal scale
and in the writers’ opinion the Australian L. laevigata II Zone is not younger
than the upper half of the Swedish S. brachymetopa Zone. The occurrence of the
Swedish Diplagnostus planicauda vestgothicus (Wallerius) in the D 18 fauna also
tends to support the correlation of the Australian L. laevigata II Zone with the Zone
of S. brachymetopa although in Sweden this form also occurs in the overlying
L. laevigata Zone. Thus it appears likely that the L. laevigata III Zone will correlate
approximately with the basal part of the Swedish Zone of L. laevigata. In terms of
the Swedish Scale we suggest that P. (G.) spiniger in Australia spans the boundary
separating the S. brachymetopa- L. laevigata Zones.
2. Opik (1967) showed that Oidalagnostus trispinifer Westergard ranged from the
late middle Cambrian L. laevigata III Zone (localities G 121 and G 133) to the Zone
of C. quasivespa (locality G 131) where it is associated with L. cos. Further, Opik
(1967, p. 1 34) stated that O. trispinifer occurs in the superjacent zone of Glyptagnostus
stolidotus in Tasmania. However, the only species of Oidalagnostus from Tasmania
known to the writers is indeterminate. Its age is probably the Erediaspis eretes Zone
or the C. quasivespa Zone.
In Sweden the very rare O. trispinifer has been found only in the upper part of the
L. laevigata Zone (Westergard 1946, p. 67). (Dr. Lars Karis, Geological Survey of
Sweden (pers. comm.), has found O. trispinifer in a limestone concretion containing
faunal elements of the Zone of S. brachymetopa in the Tasjo area, central Swedish
Caledonides. Thus the stratigraphic range of this species is more extensive than that
shown on Table 2. Consequently, the lower boundary of the Siberian Zone of
Maiaspis spinosa-0. trispinifer can now be confidently drawn at position 5 on
Table 3.) Thus it seems likely that the lower portions of the C. quasivespa Zone and
the top part of the Swedish L. laevigata Zone are correlatives and that the species
may in fact cover the full range of the Swedish L. laevigata Zone. This latter suggestion
DAILY AND JAGO: LEJOPYGE
537
TABLE 4. Stratigraphic distribution of trilobites important for the correlation of the north-west Siberian
middle and upper Cambrian rocks. The relative sizes of the zones and the ranges of the fossils were
calculated from Datsenko et al. (1968, ‘Atlas’, pp. 28-31) and Rosova (1968).
TRILOBITES
Agraulos punctatus
Maiaspis spinosa-
Oidalagnostus trispinifer
Acrocephalella granulosa —
Koldiniella prolixa
Pedinocephalina—
Toxotisc?)
Phalacroma glandiforme
Maiaspis spinosa
Oidalagnostus trispinifer
"Homagnostus fecundus’
X
Grdnwallia decora
X
Koldiniella convexa
-
Nganasella nganasanensis
-
Acidaspidella limita
Pseudagnostus nganasanicus
—
■peronopsis insignis”
“Clavagnostus sulcatus”
X
is supported by the common occurrence of O. trispinifer in north-western Siberia in
the middle Cambrian Mayanian Stage where according to Datsenko et al. (1968, in
‘Atlas of stratigraphic schemes’, pp. 28-29) it is found in all but the basal part of the
Zone of Maiaspis spinosa-Oidalagnostus trispinifer which, in our opinion (see below),
marks the top of the middle Cambrian (Table 4).
3. An agnostid cephalon from the Mungerebar Limestone (locality G 119, Zone
of E. eretes) figured (Opik 1967, pi. 58, fig. 1) as Agnostus‘1 sp. aff. Agnostus pisiformis
subsulcatus Westergard, may belong in our opinion to Westergard’s subspecies
which was described by him from the Paradoxides forchammeri beds, although on his
range chart (Westergard 1946, p. 102), he indicated that the species occurred only
in the L. laevigata Zone. Apart from minor taxonomic differences it would seem that
the uncertainty of Opik’s assignment was partly influenced by the belief that the
E. eretes Zone was younger than the L. laevigata Zone of Sweden.
4. According to Opik (1967, pp. 131-132) the Proagnostusl sp. from Woodstock,
Alabama, U.S.A. (see Palmer 1962), is Connagnostus venerabilis Opik, a species
which in Australia is confined to the Glyptagnostus stolidotus Zone. It is one of the few
new species of Australian agnostids described by Opik common to both continents.
Of even greater significance is its occurrence in Alabama, in the Conasauga Formation,
in association with G. stolidotus Opik (Palmer 1962, fig. 4) the nominate zone fossil
for the uppermost zone of the Australian Mindyallan Stage. Thus the intercontinental
538
PALAEONTOLOGY, VOLUME 18
correlation of the G. stolidotus Zone with the lower levels of the Crepicephalus Zone
in North America and probably an undefined part of the subjacent Cedaria Zone
seems assured (Table 3). Further, in both Australia and North America G. stolidotus
is succeeded by Glyptagnostus reticulatus (Angelin) (Opik \96\b, 1963; Palmer 1962,
Table I, p. 7). G. reticulatus is also present in Sweden where it occurs in the two oldest
subzones of the Olenus Zone and its subspecies G. r. nodulosus Westergard passes
into the overlying subzone.
Therefore it seems that providing the lower part of the C. quasivespa Zone marks
the top of the Swedish L. laevigata Zone as indicated above, then the upper part of
the C. quasivespa Zone (from the base of the B. sabulosa fauna) together with the
overlying G. stolidotus Zone must equate with the Scandinavian A. pisiformis Zone.
Thus in Australia the middle-upper Cambrian boundary would occur within the
Mindyallan Stage and within the C. quasivespa Zone as shown in Table 3.
Great Britain. Until recently the L. laevigata and A. pisiformis Zones were unknown
with certainty in Britain but they have now been positively identified from fossils
obtained from the Merevale No. 3 Borehole, Warwickshire (Rushton in Taylor and
Rushton 1972; Cowie et al. 1972). However, L. laevigata has not yet been found in
British rocks. The L. laevigata Zone is present within the Mancetter Grits and Shales.
The oldest identifiable fossil within this formation is the bradoriid crustacean Svealuta
primordialis (Linnarsson). It was found one-third of the way through the forma-
tion but fragments assigned to this species occur almost to its base. In Sweden the
species occurs in the L. laevigata Zone (Westergard 1944a, p. 33) and it is ‘abundant in
the Zone with Solenopleura brachymetopa' (Opik 1961a, p. 175). Hence it seems
EXPLANATION OF PLATE 62
All figures are rubber casts whitened with magnesium oxide prior to photography. All figures are untouched.
Figs. 1-10. Lejopyge laevigata (Dalman). 1, complete specimen (Westergard 1946, pi. 16, fig. 9) from
Ullavi (boulder), Narke, x7-3. 2, cephalon (Westergard 1946, pi. 13,fig. 24)fromDjupadalen, Vastergot-
land, x8-4. 3, cephalon showing scrobiculation (Westergard 1946, pi. 13, fig. 22) from Honsater,
Kinnekulle, Vastergotland, X 11-2 (the black hole is a hole in the cast). 4, pygidium (Westergard 1946,
pi. 13, fig. 26) from Gudhem, Vastergotland, x 8. 5, small pygidium showing post-axial median furrow
(associated with specimen figured PI. 62, fig. 9), Andrarum, Scania, x 1 2-4. 6, minutely spinose pygidium
(associated with cephalon figured PI. 62, fig. 2), Djupadalen, Vastergotland, xlO. 7, pygidium
(Westergard 1946, pi. 13, fig. 23) from Honsater, Kinnekulle, Vastergotland, x8. Note the very faint
third pygidial node and the several pairs of muscle scars on the third pygidial lobe. 8, pygidium (associated
with cephalon figured PI. 62, fig. 2), Djupadalen, Vastergotland, x 10. 9, pygidium (Westergard 1946,
pi. 13, fig. 20) from Andrarum, Scania, x9-7. 10, pygidium (Westergard 1946, pi. 13, fig. 25) from
Djupadalen, Vastergotland, x 8.
Figs. 11-18. Lejopyge laevigata armata (Linnarsson). Figs. 11, 12, 13 are of specimens associated with the
cephalon (Westergard 1946, pi. 14, fig. 2) figured herein (PI. 63, fig. 1) as the holotype of Lejopyge laevigata
perrugata from Karlfors, Billingen, Vastergotland. 11, cephalon with long spines, x8-4. 12, pygidium
with long spines, x7-9. 13, small pygidium, x 10. 14, spinose cephalon associated with pygidia of
Lejopyge laevigata armata (see PI. 62, figs. 15, 16) from Gudhem, Vastergotland, x 13. 15, pygidium
with small spines (Westergard 1946, pi. 13, fig. 30), x7-6. 16, pygidium with small spines (Westergard
1946, pi. 13, fig. 31), x7-5. 17, crushed cephalon (Opik 1967, pi. 57, fig. 6 as Lejopyge cos) from
Mungerebar Limestone, Queensland at Lat. 22° 15-5' S., Long. 139° 01' E., x 13-5. 18, pygidium figured
(Opik 1967, pi. 57, fig. 5) as holotype of Lejopyge cos, Mungerebar Limestone, Queensland, at Lat.
22° 15-5' S., Long. 139° OF E., x9-4.
PLATE 62
DAILY and JAGO, Lejopyge
540
PALAEONTOLOGY, VOLUME 18
likely that the lower third of the Mancetter Grits and Shales could conceivably
incorporate part of the S. brachymetopa Zone, rather than all of it belonging to the
L. laevigata Zone as suggested by Rushton. Such an uncertainty is expressed in
Table 3. Irrespective of its age, the basal part of the formation is a conglomerate
(see also Illing 1916, p. 395; Stubblefield 1956, p. 31) which may reflect an erosional
event comparable with that of the Exporrecta conglomerate of Sweden. The youngest
fossil which can be assigned confidently to the L. laevigata Zone is Hypagnostus
sulcifer (Wallerius), found near the top of the formation. Westergard (1946, p. 52)
reports this species only from the upper part of the Swedish L. laevigata Zone.
The A. pisiformis Zone is contained with certainty in the lower part of the over-
lying Outwoods Shales. A. pisiformis and Schmalenseeia cf. amphionura occur
together at or near the base of the zone, a 10-m interval below this level remaining
unassigned due to lack of diagnostic fossils.
An important find about three-fifths of the way through the Mancetter Grits and
Shales was Ptychagnostus (Goniagnostus) fumicola Opik (Rushton in Taylor and
Rushton 1972, p. 9). However, on the bore log record (ibid., pi. 4) the identification
appears to be less certain for there it is given as Ptychagnostus cf. fumicola. Through
the kind efforts of Dr. A. Rushton we have examined latex casts of this material and
believe that the assignment of P. (G.) fumicola Opik is correct. Now in the Mungerebar
area in Queensland, P. (G.) fumicola occurs with Oidalagnostus trispinifer in rocks
(locality G 121) referred by Opik (1967, p. 91) to the L. laevigata III Zone. It is
also found in the succeeding zone with Damesella torosa and Ascionepa janitrix which
Opik called the middle-upper Cambrian zone of passage. However, as pointed out
above, O. trispinifer in Queensland is known to range upwards into the C. quasivespa
EXPLANATION OF PLATE 63
All figures are photographs of rubber casts, except figs. 7 and 8 which are of the actual specimens. All
were whitened with magnesium oxide prior to photography. All figures are untouched. Catalogue numbers
are those of the palaeontology collections, South Australian Museum, Adelaide, South Australia.
Fig. 1. Holotype cephalon of Lejopyge laevigata perrugata (Westergard 1946, pi. 14, fig. 2) from Karlfors,
Billingen, Vastergotland, x 9.
Figs. 2, 3. Pygidia associated with the holotype cephalon of Lejopyge laevigata perrugata. 2, pygidium with
very large spine base, x 10-8. 3, pygidium with broad low ridge posterior to the second axial node, x 1 1.
Figs. 4, 5. Pygidium and rugose cephalon associated with the holotype cephalon of Lejopyge laevigata
rugifera from Sjogestad, Ostergdtland. 4, pygidium, x 7-7. 5, cephalon, x 7-4.
Fig. 6. Holotype cephalon of Lejopyge laevigata rugifera (Westergard 1946, pi. 14, fig. 3), x 8-4.
Figs. 7, 8. Lejopyge sp. (probably Lejopyge calva) from coquina at Patterson Pass, Snake Range, East
Nevada. 7, P. 14545, cephalon, x8-8. 8, P. 14546, pygidium, x 10-4.
Fig. 9. Pygidium of unnamed subspecies of Lejopyge calva (see Robison 1964a, p. 522) from U.S. Geo-
logical Survey Collection 2523-CO, Schell Creek Range, Nevada, x8-5. Note the third pygidial node.
Fig. 10. Lejopyge calva Robison, holotype cephalon (Robison 1964a, pi. 83, fig. 1) from 1336 ft above
base of the Marjum Formation, Wheeler Amphitheater, House Range, Western Utah, x 10-3.
Fig. 1 1. Lejopyge exilis pygidium (Whitehouse 1936, pi. 9, fig. 12) from 8 miles north-east of Duchess,
Queensland, x7T.
Figs. 12, 13. Ptychagnostus elegans laevissismus Westerg&rd, from Gislovshammer (boulder 18), Scania.
12, holotype cephalon (Westergard 1946, pi. 10, fig. 21), x9-7. 13, pygidium (Westergard 1946, pi. 10,
fig. 22), x81.
Figs. 14, 15. Drepanura eremita Westergard . 14, cranidium (Westergdrd 1947, pi. 3, fig. 9), locality unknown,
x3-5. 15, holotype pygidium (Westergard 1947, pi. 3, fig. 11) from Djupadalen, Vastergotland, x2.
PLATE 63
DAILY and JAGO, Lejopyge, Ptychagnostiis, and Drepanura
542
PALAEONTOLOGY, VOLUME 18
Zone where, at locality G 131, it is associated with L. cos Opik ( = L. /. armata) and
Svealuta cf. primordialis. The range of the British P. (G.) fumicola is unknown, being
found only in one thin bed, just above the mid-point of the interval allotted by
Rushton to the L. laevigata Zone. Unfortunately, without further fossil control on
the upper and particularly the lower limits of the zone, the value of P. {G.) fumicola
for refined intercontinental correlation remains untested. Nevertheless, as the species
occurs well below the occurrence of H. sulcifer, which in Sweden seems to have the
same range as O. trispinifer, it appears likely that P. (G.) fumicola may be confined
to the interval represented by the central portion of the Swedish L. laevigata Zone.
North America. In North America L. calva Robison occurs in the uppermost subzone
of the late middle Cambrian Bolaspidella Assemblage Zone. Lu (1960, p. 213) and
Robison (1964/?) independently proposed that the middle-upper Cambrian boundary
in North America be placed at the top of the Bolaspidella Zone. In reaching his con-
clusion Robison {\96Ab) assumed that the range of L. calva was contained within the
time interval occupied by the Swedish Zone of L. laevigata. However, Palmer (1968,
p. 10) has shown that in Canada L. calva is associated with Phalagnostus bituberculatus
(Angelin) and Ptychagnostus {P.) aculeatus (Angelin) both of which in Sweden are
confined to the S. brachymetopa Zone (Table 2). Palmer (1968, p. 10) also reported
L. laevigata from the Hillard Peak area in Alaska within a mile or so of the Canadian
occurrence of L. calva. Unfortunately, both species are unknown in the same section
in Alaska (or elsewhere), and thus all that can be said presently with any degree of
certainty is that L. calva, based on the Canadian occurrence, covers only the lower
part of the range of L. laevigata. Thus in North America the middle-upper Cambrian
boundary may well lie within the Cedaria Zone rather than at its base as suggested
by Robison (1964Z), c). Using the generic range of trilobites. Palmer (1962, fig. 9) was
the first to suggest that the Series boundary lay somewhere within the Cedaria Zone.
This conclusion is in harmony with our views (Table 3), which, however, are based on
more recent information at the species level. Indeed, it is the writers’ view that
correlations based on species have the best chances of being correct, for the accuracy
of correlation using genera or higher taxa is of a much lower order and should be
viewed as such. For example, of the many polymerid species listed by Opik (1967,
Table 4), only Corynexochus plumula Whitehouse and Stephanocare richthofeni
Monke presently allow for intercontinental correlation. Corynexochus has in the
past been regarded as a middle Cambrian genus. However, the anachronistic C.
plumula, which succeeds G. reticulatus in all its known occurrences in Australia and
elsewhere (Opik 1963; Palmer 1968), is clearly upper Cambrian in age.
China. Recently, Kobayashi (1967, p. 476, and fig. 5, p. 477) has discussed and
shown the areal distribution of three distinct Cambrian faunas in eastern Asia. Two
of these, namely the Hwangho Fauna and the Chiangnan Fauna are of interest here.
The Hwangho Fauna is a shallow-sea fauna which contains mainly endemic elements
with rare cosmopolitan elements. In contrast, the Chiangnan Fauna is interpreted as
a pelagic or offshore fauna preserved in mainly black carbonaceous shales; its facies
is similar to the dark-coloured Scandinavian alumshale and stinkstone facies.
S. richthofeni, an important member of the Hwangho Fauna, provides a firm
correlation of the Australian C. quasivespa Zone with part of the Kushan Formation
DAILY AND JAGO: LEJOPYGE
543
sensu stricto, of northern China. There S. richthofeni is confined to the Blackwelderia
paronai Zone (elsewhere in the text and Table 3 the term B. sinensis Zone is used in
preference to the term B. paronai Zone) and the lower part of the succeeding Drepanura
premesnili Zone (Chu 1959). Sun (1948), on the basis of the occurrence of D. eremita
Westergard in the Swedish A. pisiformis Zone, argued for an early upper Cambrian
age for the Kushan Formation sensu stricto. Opik ( 1 967) assigned both D. eremita and
D. ketteleri Monke (note D. ketteleri is confined apparently to the D. premesnili Zone)
to Palaeadotes Opik, which in Australia occurs in both the C. quasivespa and G. stoli-
dotus Zones. Palaeadotes Opik is, however, a synonym of Bergeronites Sun whose
genotype is Drepanura ketteleri Monke (see Kuo 1965, p. 637). We have re-examined
D. eremita and believe that although it is close to Bergeronites it should be reassigned
to a new genus. For example, its anterior facial sutures are distinctly divergent and
not convergent as in Bergeronites and its pygidium has a well-defined border (see
PI. 63, figs. 14, 15). Thus, less importance should be accorded this species for inter-
continental correlation than has been in the past.
In contrast to the paucity of agnostids in the Hwangho Fauna, there is a relative
abundance of cosmopolitan agnostids in the Chiangnan Fauna. This fauna occurs in
a broad north-easterly trending belt of rocks across south-eastern China and embraces
parts of South Korea. Within this belt, on the Hunan-Kueichow border in southern
China, Egorova et al. (1963) have reported Drepanura in the Para-Kushan Fauna
in association with Proceratopyge conifrons Wallerius, a species confined to the
upper part of the Swedish L. laevigata Zone (Table 2). At another locality Drepanura
was found with "Glyptagnostus fossus' Pokrovskaya { = G. stolidotus Opik) and
G. ret/cM/flto (Kobayashi 1971, Table 13, p. 177). Hence, in terms of the Scandinavian
scale, and providing the determinations of the fauna are correct (we have not seen
Egorova et al. 1963), Drepanura (or Drepanurinae if the determinations are not
precise) would range from the upper part of the L. laevigata Zone to at least the base
of the Olenus Zone where G. reticulatus is present in its lower part. Note that in
Australia Opik (1961Z), p. 430) reports that G. stolidotus and G. reticulatus ‘overlap
for a short interval (represented by a few feet of sediment only)’. This range for the
Drepanurinae, therefore, is comparable to that cited above for Queensland. However,
in northern China, Chu (1959) has shown that Drepanura and Bergeronites are pre-
sumably restricted to the D. premesnili Zone whereas S. richthofeni ranges downwards
into the lower levels of the B. sinensis Zone. As Bergeronites aff. dissidens occurs in
Queensland in the C. quasivespa Zone with L. laevigata armata [= L. cos] (Locality
G131) and with S. richthofeni (Locality G153) it would seem that the B. sinensis and
C. quasivespa Zones are correlatives either fully or at least in part and that the
D. premesnili Zone must in turn be correlated with the Australian G. stolidotus Zone
and the upper part of the Swedish A. pisiformis Zone (Table 3). This agrees with con-
clusions cited above. Likewise the Stephanocare Zone below the Drepanura Zone in
South Korea will correlate to the C. quasivespa Zone as S. richthofeni is confined to
the Stephanocare Zone in that region.
Within the Chiangnan faunal belt in China, Kobayashi (1967, pp. 459-461) has
reported the occurrence of Lejopyge in the Yanglioukang limestone in west Chekiang
and south Anhwei provinces. In west Chekiang L. 1. armata occurs in the upper part
of the formation (Kobayashi 1971, p. 176) and below Glyptagnostus beds above. In
544
PALAEONTOLOGY, VOLUME 18
south Anhwei Lejopyge occurs below rocks with Drepanura, Blackwelderia, and Pro-
ceratopyge and many other genera, but further pertinent details are unavailable to us.
Kobayashi (1967, p. 501) also reports Lejopyge from the dark- and light-grey bedded
limestones and shales of the Mehuershan Series in the Eastern Tienshan. Glyptag-
nostus occurs in the 25-m thick basal member of the overlying Torsuqtagh Series.
In presenting a list of the middle and upper Cambrian trilobites from the Chiangnan
faunal belt of central and south China, Kobayashi (1967, p. 462) reported L. 1. armata
in the middle Cambrian sequence of the Kueichow-Hunan border region. Its occur-
rence is listed together with the Swedish agnostids Ptychagnostus aculeatus (Angelin),
P. atavus (Tullberg), and Diplagnostus planicauda bilobatus Kobayashi. We have
been unable to check either the original locality data (presumably this is in Egorova
et al. 1963) to see if further stratigraphic refinement is possible, or to check the fossil
identifications. However, in the Handbook of standard fossils of south China (Chinese
Academy of Science, 1964) some of the named species are figured but without
accompanying locality and stratigraphic data. We believe the squashed specimen on
plate 3, fig. 10 therein is correctly referred to L. 1. armata although we have some
reservations about the identity of their P. atavus (pi. 2, figs. 8, 9). The material figured
as P. aculeatus (pi. 2, figs. 10, 11) is not Chinese but Swedish material figured by
Westergard (1946, pi. 12, figs. 9, 8). Judging the data presented by Kobayashi (1971,
pp. 175-177) it seems likely that the listed L. 1. armata is from the west Chekiang
occurrence cited above and that it has been inadvertently placed in the list of material
from the Kweichow-Hunan border. Until more concrete facts are known concerning
the occurrence of L. 1. armata and its relationship to immediately overlying faunas
in this part of China, a final decision concerning the Series boundary cannot be given.
However, the present evidence seems to favour the drawing of the boundary at some
point within the Blackwelderia sinensis Zone rather than at its base as has so often
been suggested. This conclusion pertains only to the Hwangho faunal facies belt.
Lejopyge is yet unknown in this facies and is seemingly restricted to the Chiangnan
Fauna. It is critical that further studies be conducted to find areas of intertongueing
of the two faunal belts to prove or negate the above conclusion.
U.S.S.R. Three zones, namely the Agnostus pisiformis-^ Homagnostus fecundus',
G. stolidotus, and G. reticulatus Zones constitute the early upper Cambrian Tuorski
or Tuorian Stage, Siberian Platform (Table 3). Its stratotype occurs in the foothills
of the Tuora-Sis Ridge, 6 km below Chekurovka village on the River Lena (Lazarenko
1966; Ivshin and Pokrovskaya 1968). In northern Siberia Demokidov (1968) has
referred to the interval covered by the two lower zones as the Sukhanski Horizon.
The middle-upper Cambrian boundary is drawn between the Mayanian ( = Maisky)
Stage and the overlying Tuorian Stage (Table 3). The uppermost zone of the Mayanian
Stage is the Zone of L. armata- Lomsucaspis alta (Table 3). In Lazarenko’s zonal
scheme the same zone is called the Zone of ‘L. armata-M. mirabilis'. Presumably,
L. 1. armata is not necessarily present, as in the accompanying faunal list 'Lejopyge
ex gr. laevigata" is cited. However, elsewhere in Siberia L. 1. armata has been recorded
from many sections, for example in northern Siberia (Demokidov 1968) and in the
north-western portion of the Siberian Platform, within the upper levels of the
Mayanian Stage, in the Gremyakinskaya Anticline and on the River Mokoutey at
DAILY AND JAGO: LEJOPYGE
545
the Rylninskii Ledge (Datsenko et al. 1968). Note also that only A. cf. pisiformis has
been recorded from the Altay-Sayan fold belt (Romanenko 1972), so it seems invalid
to use it as one of the nominate species in a zonal scheme. "Homagnostus fecundus',
however, is not yet described and is a nomen nudum (Lazarenko, pers. comm. 1974).
The faunal lists for the two oldest zones of the Tuorian Stage stratotype given by
Lazarenko (1966, chart opposite p. 34) and by Ivshin and Pokrovskaya (1968,
pp. 98-99) are significantly different. It is difficult to make a judgement without
figures of the listed species and one might assume that the later of the two lists has
updated the earlier one and includes taxa from more recent collections. With this in
mind the following comments are offered. Four of the species listed for the
A. pisiformis-" H . fecundus' Zone occur outside the limits of the U.S.S.R. In Sweden
Damesella{l) eremita { = Drepanura eremita Westergard) and Proceratopyge nathorsti
Westergard are known only from the A. pisiformis Zone whereas Acrocephalites
stenometopus (Angelin) is confined to the L. laevigata Zone (Westergard 1952 and
Table 2 herein). However, in her determination of fossils from the G. stolidotus Zone,
Lazarenko (1966) identified A. stenometopus agnostorum Westergard and if this is
correct, then the Swedish yI./?/5z/brw A Zone is indicated (Westergard 1948). Lazarenko
(pers. comm. 1974) has not only reaffirmed the identification but has pointed out that
the subspecies is now known from the A. pisiformis-" H . fecundus' Zone as well as the
lower G. stolidotus Zone. We presume that Acrocephalites stenometopus recorded in
Ivshin and Pokrovskaya (1968) is in reality the subspecies A. s. agnostorum in which
case the base of the A. pisiformis-" H. fecundus' Zone will coincide with the middle-
upper Cambrian boundary. If, however, Acrocephalites stenometopus is really present
below A. s. agnostorum, then the middle-upper Cambrian boundary would need to
be drawn within the zone and not at its base as indicated in Table 3 herein. The fourth
species Pseudagnostina contracta was described by Palmer ( 1 962) from the G. stolidotus
beds in Alabama, U.S.A., where it is unknown outside that zone. In the Tuorian Stage
stratotype P. contracta and Proceratopyge nathorsti pass from the A. pisiformis-
"H. fecundus' Zone into the overlying interval referred to as the G. stolidotus Zone
thus suggesting that the upper levels of the A. pisiformis-" H . fecundus' Zone may
correlate with the lowest parts of the G. stolidotus Zone elsewhere. Such an idea is
expressed in Table 3. It should also be emphasized that Ivshin and Pokrovskaya
(1968, p. 98) recorded G. reticulatus angelini Resser and Homagnostus obesus (Belt)
in the G. stolidotus Zone in addition to the nominate species. In Sweden H. obesus is
confined to the Olenus Zone. Thus it appears that the upper part of the Siberian
G. stolidotus Zone in the Tuorian Stage stratotype already includes rocks that can be
correlated with the lower levels of the Swedish G. reticulatus Zone and consequently
the upper boundary of the Siberian G. stolidotus Zone is drawn a little higher than
the base of the Swedish G. reticulatus Zone (Table 3).
In the middle section of the River Kulyumbe, a tributary of the River Yenisey in
north-western Siberia, the listed Swedish agnostids given in Datsenko et al. (1968,
‘Atlas’, Table 3, pp. 6-7) suggests that the Mayanian Stage, as recognized in that
region, is represented by the time interval equivalent to that covering the Swedish
Zone of Ptychagnostus punctuosus to the top of the L. laevigata Zone (but see below).
Its two uppermost zones are the Zone of Maiaspis spinosa-Oidalagnostus trispinifer
below and the Zone of Acrocephalella granulosa- Koldiniella prolixa above. All the
546
PALAEONTOLOGY, VOLUME 18
species in the latter zone are endemic to the U.S.S.R. except for Peronopsis insignis
(Wallerius) which in Sweden is confined to the upper part of the L. laevigata Zone
(Westergard 1946, p. 43). Rosova (1964, fig. 2) has indicated that P. insignis is
restricted to the lower and midsections of the Sakhaiski Horizon, the upper-
most division of the Middle Cambrian in her stratigraphic scheme. As well,
Datsenko et al. (1968, p. 7) included P. insignis in their list of fossils contained in the
Acrocephalella granulosa-Koldiniella prolixa Zone which together with the upper
levels of the underlying Maiaspis spinosa-Oidalagnostus trispinifer Zone they equated
with the Sakhaiski Horizon. However, on their charts Datsenko et al. (1968,
fig. 31, p. 31) and Lazarenko and Nikiforov (1968, chart opposite p. 20) have also
shown the occurrence of P. insignis in the very basal part of the overlying
Pedinoeephalina-Toxotis(l) Zone (Table 4). This seems to support the observation
by Lazarenko and Datsenko (1967, chart opposite p. 16) of the presence of P. insignis
in both the A. granulosa- K. prolixa and Pedinocephalina-Toxotis{l) Zones. Like
Westergard (1946) we regard P. insignis as indicative of a late middle Cambrian
age. However, in our opinion the agnostid figured as P. insignis by Lazarenko and
Nikiforov (1968, pi. 1, figs. 1-5) is incorrectly assigned because the pygidial axes of
the two forms are different and the glabella of the Swedish form is shorter than the
Siberian form; likewise for the pygidium figured by Rosova (1964, pi. 13, fig. 16).
Also Lazarenko and Nikiforov (1968) charted Clavagnostus suleatus Westergard
(known in Sweden only from the upper part of the L. laevigata Zone) as occurring
above the form they called P. insignis (Table 4). The pygidia figured as C. suleatus
(Lazarenko and Nikiforov, pi. 3, figs. 13, 14) may be incorrectly assigned (Jago and
Daily 1974, p. 99). Thus neitW of these two agnostids are important for the boundary
problem. However, their ranges are shown herein on Table 4 for comparison with
those of other trilobites mentioned in the text.
Many of the species recorded in the A. granulosa- K. prolixa Zone range up from
the underlying zone. Among the new forms is ' Homagnostus fecundus' Pokrovskaya,
the nominate zone fossil in the Siberian A. pisiformis- H . fecundus' Zone of the type
Tuorian Stage. Datsenko et al. (1968) have indicated on their stratigraphic tables
(p. 7 and Table 13, p. 41) that the A. granulosa-K. prolixa Zone at the top of their
Mayanian Stage is middle Cambrian in age. However, as the Swedish agnostid
O. trispinifer ranges only to the top of the M. spinosa-0. trispinifer Zone (Table 4),
we suggest that the middle-upper Cambrian boundary should be placed at the top
of this zone (Table 3) rather than at the top of the succeeding A. granulosa-K. prolixa
Zone as suggested by most Soviet workers. Also because of the spot occurrence of
‘//. fecundus' in the latter zone (Table 4) the present authors suggest that this zone
would better equate with the A. pisiformis-'H. feeundus' Zone of the type Tuorian
Stage, in which case it is upper Cambrian in age (Table 3).
The lower levels of the overlying Zone of Pedinocephalina-Toxotisfl) can be
correlated with the lower Nganasanski Horizon at the bottom of the Kulyumbeiski
Superhorizon or Substage of Rosova (1963, 1964, 1968, 1970) by means of the short-
ranging Nganasanella nganasanensis Rosova, Koldiniella convexa Lazarenko, and
Groenwallina decora Rosova (Tables 3 and 4). Pseudagnostus nganasanicus Rosova
occurs in the same horizon (Rosova 1964, fig. 2). Also of importance for correlation
is the reported occurrence of the very distinctive Acidaspidella limita Rosova, the
DAILY AND JAGO: LEJOPYGE
547
lower range of which according to Rosova (1964, 1968, 1970) is near the base of the
Nganasanski Horizon, although Datsenko et al. (1968, ‘Atlas’, fig. 31, p. 31) and
Lazarenko and Nikiforov (1968, chart opposite p. 20) record its first appearance
above the upper range of N. nganasanensis. Rosova’s observations for the species’
range are accepted herein (Table 4) particularly as Rosova (1970) has re-emphasized
its occurrence near the base of the Nganasanski Horizon. P. nganasanicus and
A. limita appear to be endemic to the U.S.S.R. Their occurrence also in the G. stoli-
dotus Zone of the Tuorian Stage stratotype (Ivshin and Pokrovskaya 1968) permit
reference of both the lower Nganasanski Horizon and the lower part of the
Pedinocephalina-Toxotis{l) Zone to the G. stolidotus Zone. Such a conclusion
reinforces the view suggested above that the A. granulosa- K. prolixa Zone is to be
correlated with the A. pisiformis-" H . fecundus' Zone of the Tuorian Stage stratotype
and with the lower part of the Swedish A. pisiformis Zone (Table 3).
CONCLUSIONS
The present revision of the taxonomic status of L. cos Opik has led to the conclusion
that it is a junior synonym of the morphologically variable L. 1. armata Westergard.
All known species of Lejopyge are of late middle Cambrian age.
In Sweden L. laevigata and its subspecies range through the Solenopleura brachy-
metopa Zone and throughout the succeeding Zone of L. laevigata, the top of which
marks the middle-upper Cambrian boundary.
For Australia, it is advocated that because L. cos Opik is synonymous with L. 1.
armata Westergard, the middle-upper Cambrian boundary should be drawn within
the Mindyallan Stage and at a level within the Cyclagnostus quasivespa Zone between
the L. cos and Blackwelderia sabulosa faunas. Previously the boundary has been
drawn at the base of the Mindyallan Stage.
L. laevigata is presently unknown from British rocks. In England recent finds of
agnostids and other fossils in the Merevale No. 3 Borehole show that the middle-
upper Cambrian boundary lies within an unfossiliferous interval between the occur-
rence of Hypagnostus sulcifer (Wallerius), found near the top of the Mancetter Grits
and Shales, and below the occurrence of Agnostus pisiformis (Linnaeus) and
Schmalenseeia cf. amphioneura, found towards the base of the overlying Outwoods
Shales (Table 3).
In North America the top of the Bolaspidella Assemblage Zone, which contains
L. calva, has been regarded as the uppermost zone of the middle Cambrian. However,
present evidence from Alaska where both L. calva and L. laevigata are found, sug-
gests that the middle-upper Cambrian boundary for North America is more likely
to occur within the overlying Cedaria Zone (Table 3).
In China L. laevigata is apparently absent within the shallow-water shelf facies of
the Hwangho Faunal Facies belt. Existing evidence favours the positioning of the
middle-upper Cambrian boundary at some undefined level within the Blackwelderia
sinensis Zone rather than at its base. However, elsewhere in China and within the
Chiangnan Faunal Facies belt, the occurrence of L. 1. armata and other cosmopolitan
agnostids should permit a reliable positioning of the Series boundary.
On the Siberian Platform, in the foothills of the Tuora-Sis Ridge, the middle-upper
548
PALAEONTOLOGY, VOLUME 18
Cambrian boundary appears to be correctly drawn between the L. 1. armata-
Lomsucaspis alta Zone below, and the A. pisiformis-'' Homagnostus fecundus' Zone
above. However, in north-west Siberia evidence presented above suggests that the
middle-upper Cambrian boundary should be drawn at the top of the Maiaspis
spinosa-Oidalagnostus trispinifer Zone (Table 3) rather than at the top of the succeed-
ing Acrocephalella granulosa-Koldiniella prolixa Zone as is presently done by Soviet
authors.
Acknowledgements. We are indebted to Professor A. R. Palmer, Drs. V. Jaanusson and H. Mutvei, the late
Professor F. Brotzen, Professor D. Hill, and Dr. J. Shergold for kindly allowing us to obtain rubber moulds
from specimens in their care and to Dr. A. W. A. Rushton who sent us rubber moulds of fossils from the
Merevale No. 3 Borehole, Warwickshire. Dr. V. A. Gostin translated some of the Russian literature.
One of us (J. B. J.) was supported by a grant from the Australian Research Grants Committee.
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PALAEONTOLOGY, VOLUME 18
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B. DAILY
Department of Geology and Mineralogy
University of Adelaide
G.P.O. Box 498B
Adelaide, South Australia 5001
J. B. JAGO
School of Applied Geology
South Australian Institute of Technology
Typescript received 23 April 1974 North Terrace
Revised typescript received 29 November 1974 Adelaide, South Australia 5000
THE OSTRACOD PARAPARCHITES MINAX
IVANOV, SP. NOV. FROM THE PERMIAN
OF THE U.S.S.R., AND ITS MUSCLE-SCAR
FIELD
by M. N. GRAMM and v. k. ivanov
Abstract. The ostracod Paraparchites minax sp. nov., from the early Permian of the Pre-Donetz Depression of
the Rostov area of the Soviet Union, is described and figured. Particular attention is paid to the muscle scars, to
mandibular and frontal scars and especially to the adductor muscle scars, which are in the form of a cluster of up
to 190 spots. An outline of the ontogenetic development of the scars is given. The systematic position of the Para-
parchitacea is discussed in the light of outline, inner lamella, dimorphism, and central muscle-scar pattern, with the
conclusion that the superfamily is related neither to the Platycopa, nor the Kloedenellidae, and in consequence,
a new suborder of the Podocopida, the Paraparchitocopa, is proposed.
In the early Permian strata of the Donetz, amongst the commonest ostracods are
members of the Paraparchitacea, a preliminary account of which has been given by
Ivanov (1964). Particularly well-preserved specimens, including large numbers of
the form Paraparchites minax sp. nov., were recovered from a depth of 474-475 m
in Asselian stage beds in drillings in the Rostov region (Tatzin district, Skosyr area).
Such was the preservation that some thirty specimens showed details of adductor,
mandibular, and frontal scars, improving our hitherto scanty knowledge of the
muscle-scar patterns of Palaeozoic ostracods. Thus, the main purpose of this paper
is to describe and analyse these structures and to discuss the systematic position of
the Paraparchitacea. In most earlier works the central muscle-scar field has been
studied from internal moulds, or from the inner surface of valves. In the specimens
described here the details have been obtained by treating translucent or semi-
transparent carapaces with castor oil or sugar solutions and photographing the
specimens in reflected light.
All specimens referred to in the text under No. 146 have been deposited in the Ukrainian Scientific
Research Institute for natural gas (UkrNIIGas), Kharkov.
SYSTEMATIC DESCRIPTION
Order podocopida Muller, 1894
Suborder paraparchitocopa Gramm, n. suborder
.Diagnosis. Dorsal margin straight, ventral margin generally convex. Surface smooth,
one or two postero-dorsal spines may be present. Calcified inner lamella narrow.
Adductor muscle scar in the form of a cluster, which may contain a large number
of spots. Mandibular scar elongate; frontal scar complex. Dimorphism of non-
kloedenellid type. One superfamily— Paraparchitacea Scott, 1959. Range: Devonian
to Permian.
[Paleontology, Vol. 18, Part 3, 1975, pp. 551-561, pi. 64.]
552
PALAEONTOLOGY, VOLUME 18
Superfamily paraparchitacea Scott, 1959
Family paraparchitidae Scott, 1959
Genus paraparchites Ulrich and Bassler, 1906
Paraparchites minax Ivanov, sp. nov.
Plate 64, figs. 1-9
1964 Paraparchites humerosus Ulrich and Bassler, 1906, morpha magna Ivanov, 1964, p. 110,
pi. 2, fig. \a-c.
Derivation of name. ‘Minax’ = prominent (Latin).
Holotype. Complete carapace, 146/1.
Paratypes. Thirteen complete carapaces (146/2, 3, 4, 5, 146/9-3, 146/10-1, 146/10-2, 146/11-1, 146/11-2,
146/11-4, 146/12-3, 146/13, and one right valve, 146/6). All types are from Borehole 2323, from the Asselian
Stage, at 474/475 m, Tatzin district, Skosyr area, Rostov.
Material. Eighty carapaces, and over 100 valves.
Diagnosis. Carapace large, up to 2800 ij.m, elongate and sub-oval; left valve slightly
overlaps right valve along the entire free margin, with reversal of overlap along the
hinge margin.
In lateral view, anterior and posterior margins evenly rounded, although the former
is more fully curved ; dorsal margin short, straight, and somewhat inclined posteriorly ;
cardinal angles weakly developed; ventral margin convex, merging smoothly with
anterior and posterior margins. Shell surface smooth, with a few scattered pits
corresponding to normal pore canals. Parallel to the free margin, and close to it,
thin elevated ridges sometimes observed. No internal features other than the central
muscle-scar field are known, these consisting of the adductor field located centrally
within the valve and made up of up to 190 spots, an elongate mandibular scar, and
a frontal scar.
Dimensions. Details of type specimens given in Table 1.
Ontogeny. The smallest specimens, 725 ptm long, possibly Instar III, differ little
morphologically from the holotype (an adult carapace). Changes during growth
follow a pattern of regular increase in all basic dimensions relating to shape. There
is size increase in the adductor scar field, as well as an increase in number and size of
EXPLANATION OF PLATE 64
Figs. 1-5. Paraparchites niina.xlnvdno\,sp. nov. 1, 2, carapace, holotype, no. 146/1. 1, right view; 2, dorsal
view. 3, 4, carapace, no. 146/3; 3, right view. 4, dorsal view. 5, carapace, no. 146/4, right view. Rostov
region, Skosyr area; Lower Permian. All x 15.
Figs. 6-9. Central muscle-scar field of Paraparchites minax Ivanov. 6, 7, larval stages. 6, right side of
carapace, VI? instar, L 1400 /xm, no. 146/10-1 ; adductor muscle spots, mandibular spot, and frontal
spot are seen. 7, right side of carapace, VI? instar, L=- 1500 ;u,m, no. 146/9-3; adductor muscle spots,
mandibular spot, and frontal spot are seen. 8, 9, adults. 8, right side of carapace, L = 2525 jum, holotype,
no. 146/1 ; adductor muscle spots and mandibular spot are seen. 9, left side of carapace, L = 2550 jum,
no. 146/4; adductor muscle spots, mandibular spot, and frontal spot are seen. Photographs were taken
from the outer side in reflected light. L— length of carapace. Rostov region, Skosyr area; Lower Permian.
All X 150.
Fig. 10. Paraparchites sp., right valve, no. 1 1 16/76-1, internal view in transmitted light; the inner lamella
is seen. Leningrad region; Lower Carboniferous, x 30.
PLATE 64
GRAMM and IVANOV, Paraparchites
554
PALAEONTOLOGY, VOLUME 18
the spots. There is limited variation in the adult stage, rare specimens showing greater
inflation, or concave ventral margins. Other than the inflation mentioned above, no
clearly dimorphic features have been observed.
Remarks. The new species differs from Paraparchites scotoburdigalensis (Hibbert)
from the British Carboniferous in its greater dimensions, and length : height ratio.
Some of our specimens are morphologically close to those figured as P. humerosus
Ulrich and Bassler by Scott (1959), but these are of much smaller dimensions (length
2000 |Ltm).
Ecology. The early Permian paraparchitaceans from the Donetz area appear to
have lived in conditions of varied salinity, leading Ivanov (1964) to conclude that
they were euryhaline. A similar conclusion was reached by Robinson (1969) and
Sohn (1971) who both thought that, although essentially a marine genus, Paraparchites
may have tolerated brackish and hypersaline conditions at times. In the Donetz
region P. minax sp. nov. occurs in grey and dark-grey argillaceous limestones,
accompanied by an abundance of darwinulaceans and carbonitids (Darwinula sp.
and Carbonita sp.). Other fauna includes micro-gastropods and bivalves, calcareous
worm tubes, stick bryozoan fragments, fish scales, and denticles. Particularly the
abundance of darwinulids, and the paucity of marine invertebrates, suggest abnormal
salinity conditions, verging upon fresh water.
THE MORPHOLOGY OF THE CENTRAL MUSCLE-SCAR FIELD
OF PARAPARCHITES MINAX
Adductor scar field. On the surface of adult carapaces, 2450-2800 /xm long, the
adductor scar field is sometimes evident as a shallow, circular depression located in
the centre of the valve. In the adult, the adductor scar field is a circular to elliptical
cluster of small spots, the long axis of the cluster aligned dorso-ventrally. The cluster
can be 270 /xm in length and 300 ju,m in height. The number of spots within the cluster
varies from 128 to 190, and may differ in the two valves of a single carapace. As can
be seen from Table 1, there is no close correlation between spot number and size of
carapace, indeed, in the right valve of one of the largest specimens examined (2800
length), one of the lowest spot counts, 128, was recorded. The shape of the spots
varies from circular or oval to angular, the packing being usually close-set. Any kind
of consistent pattern of spots within the scar is difiicult to detect. While details of the
ontogenetic development of the scar is scanty, the present material suggests a general
increase both in size and number of spots with growth. Thus, in specimens c. 1100 jj.m
long, spot counts range from 25 to 35; for specimens c. 1400 /xm long, the count is
40-60; for carapaces greater than 1500 |xm, the count is 46-plus.
Mandibular scar. Antero-ventral to the adductor-scar field, there lies an elongate
scar which is best interpreted as a mandibular scar. Sometimes visible on the outer
surface of the valves, the scar may be horizontal but sometimes slightly bowed.
Although the scar might suggest the coalescence of spots, there is no evidence to
support this idea. There is a gradual increase in size through ontogeny.
Frontal scar. Dorsal to the adductor-scar field, there is an oval frontal scar, 75-90 /xin
high in carapaces 1400-1500 /xm long, increasing to 100 in adults.
GRAMM AND IVANOV: THE OSTRACOD PARAPARCHITES MINAX IVANOV 555
TABLE 1. Dimensions of Paraparchites minax sp. nov. and details of the central muscle-scar field.
Mandibular Frontal
Adductor muscle scar
scar
scar
Length
Number
Length
Height
Length
Height
Collection no.
(^m)
of spots
(ium)
{^rn)
(ium)
(;um)
146/12-3
C left view
1100
16
146/11-1
C right view
1200
25
60
146/11-2
C right view
1400
35-40
146/10-1 <
\ C right view
[ C left view
1400
60
125
125
140
140
175
170
75
90
146/9-2
C left view
1500
46
146/9-3 <
f C right view
[ C left view
1500
40
40
125
140
150
160
200
200
80
146/10-2 j
1 C right view
1650
170
170
210
50?
/ C left view
150
160
200
146/10-3
C right view
2050
200
146/7
C right view
2250
250
250
250
146/6
RV
2450
151
225
240
325
146/1 j
C right view
^ C left view
2525
190
184
250
250
290
300
250
146/4 j
1 C right view
^ C left view
2550
132
138
225
225
250
250
250
250
100
146/3 j
1 C right view
2650
131
250
250
300
[ C left view
270
300
325
146/5 ^
\ C right view
2800
128
225
250
350
[ C left view
250
250
335
The central muscle field of Paraparchites minax can be homologized with this
structure in bairdids, cyprids, and cytherids, the scar representing the points of
attachment of muscle and chitin elements of the soft-part anatomy. Its mandibular
scar was presumably the attachment point of the chitinous rods springing from the
dorsal apex of the basal podomere of the mandible protopodite (Triebel 1960). The
presence of two mandibular scars reported by Ivanov (1964) and Robinson (1969),
and to be seen in Sohn’s plates (1971), may prompt the idea that these have become
fused to form the single scar of P. minax. As Smith (1971) has demonstrated that the
dorsal anterior scar in Recent cytherids and cyprids has no direct relationship to the
antennae, the term frontal scar is employed for the scars here described. The relative
disposition of the scars described by Ivanov (1964), Robinson (1969), and Sohn
(1971), together with the present evidence from P. minax, removes any doubt as to
the orientation of Paraparchites. Orientation in fact, is as described by Scott (1959).
The Paraparchitacean central muscle-scar field. Data as to the central muscle-scar
field of Paraparchites is limited, and usually refers to a smooth muscle scar in the
centre of the valve (Tschigova 1960), or a central muscle scar with faint marks
(Kummerow 1953). The first detailed description appears to be that of Ivanov (1964,
p. 110, pi. 2, fig. 4) for P. humerosus morpha oblima. Later, in 1967, Bless described
and illustrated fifty discrete spots as the muscle pattern for P. cantelii Bless, 1967,
from the Upper Carboniferous of Spain. Robinson (1969) noted that the central
muscle-scar field of Paraparchites is essentially the same as that for Bernix, a large
H
556
PALAEONTOLOGY, VOLUME 18
patch area covered with clusters of small pits, with one or two linear scars obliquely
below. Such scars were figured for Paraparchites sp. from Tournaisian, and for
Paraparchites cf. inornalus (M’Coy) from the Visean (Robinson 1969, pi. 3, figs. 3
and 4).
The fullest documentation of paraparchitid muscle-scar patterns is to be found in
the monographs of Sohn (1971, 1972), in which he specifically mentions the presence
of a ‘cyprid adductor muscle scar pattern in some of the genera’ (Sohn 1971, Al,
Abstract). According to Sohn’s schematic illustration, the most complete cypridid
pattern is that of Shishaella marathonensis (Hamilton, 1942) in which there are some
six elongate obliquely arranged adductor scars, and two closely adjoined mandibular
scars (Sohn 1971, A5, fig. 2). At the same time the scar pattern in the genus Chami-
shaella Sohn, 1971 is described as follows, ‘The subcentral adductor scar consists
of a circle of small individual scars’ (Sohn 1971, All).
Available data indicate three types of paraparchitid adductor muscle-scar patterns :
1. The pattern of P. minax, characterized by a circular cluster of many spots (up to 190). Close to this
type are the patterns of P. sp. and P. cf. inornatus from the Lower Carboniferous (Robinson 1969) and of
Cliamishaella (Sohn 1971). P. cantelii Bless, 1967 also has this type of adductor muscle scar, as does Bernix \
and Robinson (1969) has argued persuasively that Bernix belongs to the Family Paraparchitidae. The
presence of one or two mandibular scars is also typical, but a frontal scar is, at present, known in P. minax
only.
2. The pattern of P. humerosus morpha oblima, consisting of a circular group of a few scars (up to ten?)
associated with two mandibular scars (Ivanov 1964).
3. The pattern of Shishaella marathonensis, with six large scars associated with two elongate mandibular
scars. This pattern was regarded as being of cyprid type by Sohn (1971).
It is difficult to envisage three such strongly dissimilar adductor muscle-scar patterns
forming a morphological series within the paraparchitid group. At the moment, the
available data, especially for the second and third adductor muscle-scar types, are
very limited and any final assessment of the taxonomic significance of the second and
third types mentioned above must await further information.
(Latest observations on some well-preserved Visean paraparchitids from Novgorod
region revealed that in some old individuals on the adductor muscle-scar area an
intense calcification took place, due to which the structure acquired a form of a coarse,
uneven elevation. May this be the cause of scars which give the impression of a cyprid-
like adductor muscle-scar pattern?)
THE SYSTEMATIC POSITION OF THE PAR APARCHITACE A
In the past, three general views have been widely held:
1. Assigning the genus Paraparchites to the Family Kloedenellidae Ulrich and Bassler, 1923, which in
turn would place it within the Order Palaeocopa (Henningsmoen 1953; Mertens 1958), or alternatively
within the Platycopa, Podocopida (Pokorny 1958).
2. That of the 1961 Treatise oj Invertebrate Paleontology, placing the Superfamily Paraparchitacea
Scott, 1959, within the Suborder Kloedenellocopina Scott, 1961, which in turn belongs to the Order
Palaeocopida (Scott, 1961).
3. Amalgamating the Paraparchitacea with the Kloedenellacea and the Cytherellacea within a Suborder
Platycopina (opinion of Schallreuter 1968).
Other views to record are those expressed in Ostwvy, placing Paraparchites within
GRAMM AND IVANOV: THE OSTRACOD P A RA P A RC HIT ES MINAX IVANOV 557
the Family Aparchitidae Jones, 1910 (Orlov 1960) and more recently, Sohn’s defini-
tion of the Paraparchitacea as Podocopida incertae subordinis (Sohn 1971). In all
these opinions, there appear to have been judgements based upon the following
criteria. First, the presence of a form of kloedenellid dimorphism. Second, carapace
outline. Third, the presence of what is judged a calcified inner lamella. Fourth, the
type of central muscle-scar pattern.
Taking these in turn, a presumed kloedenellid dimorphism in Paraparchites has
been taken as evidence of affinity to the Kloedenellidae (Pokorny 1958 and Schall-
reuter 1968). On the other hand, evidence of dimorphism was regarded as inconclusive
by Scott (1961, p. Q86), and of limited value by Griindel (1967, p. 323). Because their
possible dimorphic features are so weak, Kniipfer has rejected any relationship of
Paraparchitacea to the Platycopina, preferring to regard them as a discrete branch
of the Podocopida, equal in status with the Platycopina and Metacopina (Kniipfer
1968). A kloedenellid-type dimorphism in paraparchitids has been completely
rejected by some, including Tschigova (1967). The same author has noted a ventral
inflation in possible female carapaces (Tschigova 1960; Buschmina 1968), a view
repeated by Robinson for Paraparchites and Bernix (1969) and by Sohn (1971,
p. A5). In P. minax some forms are ‘inflated’ with obtuse extremities, whereas others
are ‘thin or uninflated’ with acute extremities, but no traces of kloedenellid dimorphism
have been revealed. All this leads to the conclusion that any sexual dimorphism in
paraparchitids would seem to be of non-kloedenellid type, and no basis for allocation
of the group within either the Kloedenellidae, or the Platycopa.
Carapace outlines do not provide a reliable basis for placing the paraparchitids
within the Kloedenellacea or the Platycopa, groups which normally possess a recti-
linear or slightly concave ventral margin in contrast to the strongly convex venter of
Paraparchites. In P. minax the ventral margin is convex with the exception of a few
rare specimens with obvious concave ventral margins.
Published information concerning the calcified inner lamella is scanty, and even
contradictory. Scott notes that a duplicature is generally absent in the Kloedenelli-
copina, but present in the Geisinidae (Scott 1 96 1 , p. Q90 ; Sohn in Scott 1 96 1 , p. Q 1 82).
Such observations have been extended more recently by Pollard (1966) and Kniipfer
(1968) to include the genera Glyptopleura Girty, 1910, Electia Tschigova, 1960,
Hypotetragona Morey, 1935, Knoxites Egorov, 1950, Mennerella Egorov, 1950,
IMarginia Polenova, 1952, and others. Data are scarce for the Paraparchitecea. Scott
has written of a ‘vestibule’ in P. humerosus Ulrich and Bassler, 1959. Once again
Sohn (1971) is our main source of information, recording a narrow inner lamella in
the genera Shivaella Sohn, Shamishaella Sohn, Shishaella Sohn, and Shemonaella
Sohn. Working with the complete carapaces of P. minax, it has been impossible to
confirm such structures, but in well-preserved single valves of Paraparchites from the
Lower Carboniferous of the Leningrad region, a clearly visible inner lamella has
been found (PI. 64, fig. 10). Thus it can be said that the possession of a calcified inner
lamella is a characteristic of paraparchitaceans as well as of some kloedenellaceans,
separating both from Platycopa sensu stricto, the latter possessing only rudimentary
traces at best (Van Morkhoven 1962).
Our total knowledge of the central muscle-scar field of P. minax confirms the
opinion of Sohn that, ‘the lateral outline, hingement, calcified inner lamella and
558
PALAEONTOLOGY, VOLUME 18
adductor muscle scar pattern negates this assignment’ (of the Paraparchitacea to the
Platycopina: Sohn 1971, p. A5). For the Platycopa, the pattern and its evolution
could be said to be well established, changing from the multiserial scar of the
Cavellinidae (six rows of from 7 to 10 spots, totalling between 40 and 56 spots,
Triebel 1941 and Scott 1944), to the biserial scar of the Cytherellidae (Gramm
1972). In contrast, relatively little is known of the adductor muscle scar of the
Kloedenellidae. Nyhamnella from the Lower Silurian, has an oval group of spots
(23) somewhat drawn out in a dorsal direction (Adamczak 1966, fig. 1). The Lower
Carboniferous genera Geisina and Kloedenellitina have biserially arranged adductor
scars with up to 11 spots (Knupfer 1968, also Pollard 1966). With so little evidence,
it is impossible to discuss any morphological evolution of the kloedenellid scar,
except to observe that the scar type differs considerably from that of the Platycopa,
that of the paraparchitids described by Sohn (1971), and that described herein for
P. minax. Table 2 summarizes our knowledge of muscle-scar patterns for Ostracoda,
TABLE 2. Central muscle-field elements of various ostracod groups.
-I- known, — unknown.
From data published by the following authors: Sars 1922-1928; Triebel 1941, 1960; Scott 1944, 1951;
Schweyer 1949; Swartz 1949; Schneider 1956; Kashevarova 1958; Abushik 1960; authors in Osnovy
Paleontologii, 1960; authors in Treatise on Invertebrate Paleontology, Pt. Q, 1961 ; Morkhoven 1962, 1963;
Sandberg 1964; Darby 1965; Smith 1965, 1971; Gramm 1970; Gramm et al. 1972; Gramm and Posner
1972; Hartmann 1966; Adamczak 1966, 1968; Benson 1967; McKenzie 1967; Knupfer 1968; Maddocks
1969; Grundel 1970; Bolz 1971 ; Malz 1971 ; Ishizaki 1973; Shornikov and Gramm 1974.
Central muscle field
Adductor
Mandibular
Frontal
muscle scar
scar
scar
Leperditiida
4
-
-
Palaeocopida-Beyrichicopa :
Scrobicida (possibly Podocopida)
4
—
?
Placidea
+
—
—
Sulcicuneus, Svislinella, Kielciella
+
—
—
Puncia, Manawa
+
—
—
Kloedenellacea :
Nyhamnella
+
—
—
Geisina
7-
—
. —
Kloedenellitina
4-
—
—
Myodocopida :
Myodocopa
-f
—
—
Cladocopa
-4
—
—
Podocopida Platycopa :
Cavellinidae
+
—
—
Cytherellidae
f
—
—
Metacopa :
Healdiidae
f
+
+
Podocopa :
Darwinulacea
L
+
—
Bairdiacea
■f
+
+
Cypridacea
i
+
+
Cytheracea
*
+
+
Sigilliidae
t
—
+
GRAMM AND IVANOV: THE OSTRACOD PARAPARCHITES MINAX IVANOV 559
requiring it to be said that data relating to several important Palaeozoic groups are
very limited.
CONCLUSIONS
On the basis of the absence of kloedenellid dimorphism, aspects of outline, the nature
of the central muscle-scar field and its pattern, it is apparent that the Paraparchitacea
cannot be united with either the Kloedenellacea or the Platycopa. The presence of
a calcified inner lamella moves the superfamily still further from a relationship to the
Platycopa, while the development of the same structure in some kloedenellids may
be regarded as instances of evolution in parallel. On the possible criteria for a more
refined taxonomic judgement upon the Paraparchitacea, that which appeals most is
consideration of the central muscle-scar field. Such structures are, we believe,
important, because the scars are intimately associated with the soft-body anatomy
of the Ostracod, and in fossil carapaces provide as Smith said, ‘one of the common
meeting grounds between the palaeontologic and zoologic systems of classification’
(Smith 1965, p. 1). Of course, it is necessary to take other criteria into consideration,
but many internal structures in Palaeozoic ostracods are very poorly known and
ideas and opinions are frequently based on insufficient evidence. As a result, the
importance attached to certain features for taxonomic purposes varies, and the same
features may have varying significance in different groups’ ability to recognize
homologous structures of independent origin, which is crucial for phylogenetic
systematics. As our discussion has shown, the central muscle-scar field of the Para-
parchitacea can best be compared with that of the Podocopa— a view strengthened
by the record of the elongate mandibular scar. Thus in taxonomy, serious attention
should be given to the close relationship with the Podocopida postulated by Sohn
(1971). As, however, aspects of shape and outline, the absence of radial pore canals
coupled with the rudimentary nature of the duplicature, and special features of the
scar pattern, do not allow the assignment of the Paraparchitacea to any of the recog-
nized Suborders of the Podocopida, we feel that it is necessary and appropriate to
propose a new Suborder Paraparchitocopa to accommodate this group.
Acknowledgements. Thanks are due to many colleagues cited in this paper for sending literature, and to
Dr. Alan Lord, Professor P. C. Sylvester-Bradley and especially Dr. Eric Robinson for help with the manu-
script. Mrs. O. G. Gein kindly typed the paper.
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560
PALAEONTOLOGY, VOLUME 18
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GRUNDEL, j. 1967. Zur Grossgliegerung der Ordnung Podocopida G. W. Muller, 1894 (Ostracoda). N. Jb.
Geol. Paldont. Mh. 6, 321-332.
1970. Die Ausbildung der Muskelnarben an liassischen Vertretern der Healdiidae (Ostrac.). Freib.
Forsch.-H. C256, 47-63.
HARTMANN, G. 1966. Ostracoda. In Bronns Klassen des Tierreich, Bd. V, Abt. I, 2 Buch, Leipzig, 1-216.
HENNINGSMOEN, G. 1953. Classification of Paleozoic straight-hinged ostracods. Norsk Geol. Tidsskr. 31,
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republik Polen. Beiheft z. Zeitschrift Geologic, Berlin, 7, 1-75.
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MCKENZIE, K. G. 1967. Saipanellidae I a new family of Podocopid ostracoda. Crustaceana, 13, 103-113.
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derzeitigen Stand der Systematik. Geol. Jb. Hannover, 75, 311-318.
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and ecology of Recent Ostracoda. Edinburgh, Oliver and Boyd, 14-20.
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SCHALLREUTER, R. 1968. Ordovizischc Ostracoden mit geradem Schlossrand und konkaven Ventral-
rand. Ernst Moritz- Arndt-Universitdt Greifswald, Wiss. Zeitschr. 17, Math.-naturwiss. Reihe 1-2,
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SCHNEIDER, G. F. 1956. Family Placideidae Schneider fam. nov. In kiparissova, l. d., markovsky, b. p.,
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Moscow, 12, 95-96. [In Russian.]
GRAMM AND IVANOV: THE OSTRACOD PARAPARCHITES MINAX IVANOV 561
SCHORNIKOV, E. I. and GRAMM, M. N. 1974. Saipunetta McKenzie (Ostracoda) from the northern Pacific
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SCHWEYER, A. V. 1949. Principles of morphology and systematics of Pliocene and Post-Pliocene Ostracodes.
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1951. Instars and shell morphology of Eoleperditia fabulites. Ibid. 25, 321-326.
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neuer Gattungen und Arten. Senckenbergiana, 23, 294-400.
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TSCHiGOVA, V. A. 1960. Age relationship of Rakovka and Lower Malinovka deposits of Kama-Kinel
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Moscow, 30, 169-233. [In Russian.]
1967. Ostracodes in boundary beds of Devonian and Carboniferous deposits of Russian Platform.
Ibid. 49, 256 pp. [In Russian.]
M. N. GRAMM
Institute of Biology and Pedology
Far East Scientific Centre
USSR Academy of Sciences
690022 Vladivostok, U.S.S.R.
Typescript received 3 April 1974
Revised typescript received 25 October 1974
V. K. IVANOV
Ukrainian Scientific-Research
Institute for Natural Gases
Kharkov, U.S.S.R.
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THE BRADYCNEMIDAE, A NEW FAMILY
OF OWLS FROM THE UPPER CRETACEOUS OF
ROMANIA
by c. J. o. HARRISON and c. a. walker
Abstract. The only bird hitherto known from the upper Cretaceous (supposed Maestrichtian) of Transylvania is
the pelecaniform Elopteryx nopscai Andrews (1913), based on the proximal half of a femur; referred material includes
the distal ends of three tibiotarsi from the same beds. Re-examination of these tibiotarsi, however, shows that they
belong to the owls (Strigiformes); they represent the oldest owls known and are described and named as two species
in new genera, Bradycneme and Heptasteornis. The posterior side of the condylar region in both genera differs so
much from that of the tibiotarsi of Recent owls as to warrant the separation of these Cretaceous forms into the
new family Bradycnemidae.
Andrews(1913) described a large bird Elopteryx nopscai from the upper Cretaceous
of Transylvania, Romania, the holotype being the proximal half of a femur (A 1234).
He placed the new genus in the Pelecaniformes, an assignment which is confirmed by
our re-examination of the specimen. Andrews also referred to his new species, albeit
tentatively, the distal end of a tibiotarsus from the same locality for which he found
no parallel among other birds; indeed he may have thought it to belong to the same
individual, for he gave it the same register number, though it is now re-registered
A 4359— all material is in the British Museum (Natural History). When Lambrecht
(1929) discussed bird fossils from this region, and subsequently (1933) included
them in his handbook, he referred to this species a further proximal end of a femur
(A 1235) and also the distal ends of two more tibiotarsi (A 1528 and A 1588). Of the
three distal ends of tibiotarsi A 1588 is wide distally, with a large intercondylar
hollow. A 4359, described and figured by Andrews (1913), has condyles of similar
diameter but is much narrower. Since it has a median fracture it was first thought
that the middle region might have been lost through lateral pressure, but the third
specimen, A 1528, which is worn and has some small accretions of hard matrix,
does not show similar signs of damage and confirms the general size and shape of
the second specimen. Since the differences are not of a kind which could be attributed
to sexual dimorphism it would appear that two related forms are involved, one
represented by A 1588 and the other by A 4359 and A 1528.
The characters shown by these two forms, however, cannot be reconciled with those
of known pelecaniform birds and it is evident that there are no grounds for referring
them to the genus Elopteryx which is therefore represented only by the femora.
A search was therefore made for other avian taxa with which they might show
affinities. The broader specimen, A 1588, showed some general similarities to falconi-
form species and a particular similarity to Ealco rusticolus (Falconidae) in the shape
and position of the condyles and in the development of the external ligamental
prominence (text-fig. 1). The Falconidae, however, have a complex tendinal bridge
over the tendinal fossa which is absent from the fossils; the resemblance of the
[Palaeontology, Vol. 18, Part 3, 1975, pp. 563-570, pis. 65-66.]
564
PALAEONTOLOGY, VOLUME 18
fossils to Falco is therefore more likely to be the result of convergence than evidence
of affinity. In features such as the absence of a tendinal bridge, the size and shape of
the tendinal fossa, the size of the peroneus groove, the proximally placed prominence
for muscle attachment on the internal side of the anterior face of the shaft, and the
pit just proximal to the anterior face of the internal condyle, these distal ends of
tibiotarsi resemble those of owls, Strigiformes; this particular combination of
characters is not found in any other order of birds. These fragments appear to repre-
sent the earliest known owls; they are earlier than the genera Eostrix and Protostrix,
family Protostigidae (Eocene of North America), and the earliest species of the family
Strigidae, a family which is still extant, is from the upper Eocene or Oligocene of
France. They differ from Recent owls, however, in lacking any marked posterior
projection of the condyles. In modern forms the condyles project posteriorly to
a greater distance; the intercondylar groove is deep and extends back to a hollow just
proximal to the condyles on the posterior surface of the shaft, exaggerating the degree
to which the condyles appear to project. The absence of posteriorly projecting
condyles might be either primitive or secondary. The accompanying text-fig. 1 shows
the condition in a Recent owl (u), in the two fossils {b, c) and in two Recent falconids
{d, e); the distal views show very clearly that only in (a) do the condyles project
a
K*
TEXT-FIG. 1 . Anterior and distal views of right tibiotarsi : a, Strix varia (Strigidae) ; b, Heptasteornis andrewsi
gen. et sp. nov. (left tibiotarsus reversed); c, Bradycneme draculae gen. et sp. nov. ; d, Falco msticolus
(Falconidae); e, Polyborus plancus (Falconidae). Various magnifications.
posteriorly. The fact that these fossils differ consistently in this respect from Recent
owls, more so than the tibiotarsi of the two Recent owl families differ from each
other, appears to justify their segregation into a new family; and the structural dif-
ferences between the two fossil forms suggest that they represent separate genera.
Order strigiformes
Family bradycnemidae nov.
Type genus. Bradycneme nov.
Diagnosis. Large owls much larger than any described species. Distal end of tibio-
tarsus flattened. Condyles not projecting posteriorly; external condyle projecting
distally beyond internal condyle; intercondylar groove shallow. Anterior tendinal
HARRISON AND WALKER: CRETACEOUS OWLS FROM ROMANIA 565
fossa well defined, broad, fairly deep, its distal margin partly undercutting inter-
condylar region and external condyle.
BRADYCNEME gen. nov.
Type species. B. draculae sp. nov.
Diagnosis. Distal end of tibiotarsus broad and antero-posteriorly flattened. Distal
projection of external condyle beyond internal condyle very marked. Internal and
external condyles projecting anteriorly to same extent ; projection of internal condyle
also directed internally to some extent. Anterior intercondylar fossa transverse, deep.
On external side well-developed groove terminating at large external ligamental
prominence, proximally situated on external surface of external condyle.
Bradycneme draculae sp. nov.
Plate 65, figs. 1-5
Etymology. The generic name is formed from the Greek bradys ( heavy or massive), cneme (= leg) and
is feminine. The specific name is derived from the Romanian word dracul meaning evil one.
Material. Holotype only. The distal end of a right tibiotarsus, A 1588. Collected and presented by Lady
Smith- Wood ward, 1923.
Occurrence. Szentpeterfalva, Hatszeg, Transylvania, Romania. Beds attributed previously to the Danian
(now of the Palaeocene period) but according to Jeletzky (1962) and other modern authors the reptilian
forms associated with the deposits indicate a Maestrichtian upper Cretaceous age. The dating problem
is now under review by Finnigan.
Description. The specimen is a distal end of a right tibiotarsus with a short portion of shaft. It is in fairly
good condition, but damaged in places along the proximal external edge, and with small areas of crushing
elsewhere. The proximal part of the shaft shows a ridged and roughened surface. The shaft and head are
antero-posteriorly flattened, the shaft being thickest along the internal side. The external condyle projects
further distally than does the internal condyle, and is distally tilted towards the internal side. The internal
condyle projects internally, its distal edge and the distal edge of the intercondylar groove forming a level
transverse surface. Posteriorly the bone is flattened, with smooth curving edges, the central portion of its
surface merging smoothly at its distal end with the intercondylar region. Distally and posteriorly the inner
edges of the condyles are little in evidence, the intercondylar groove curving smoothly upwards laterally to
the outer edges of the condyles. The latter are only slightly prominent posteriorly. Distally the condyles
are widely spaced and most prominent at their outer edges, the intercondylar groove being deeper where it
borders the external condyle. Anteriorly the condyles have prominent rounded surfaces projecting beyond
the line of the anterior surface of the shaft.
On the anterior surface of the intercondylar groove there is the deep transverse groove of the anterior
intercondylar fossa ; the part of the intercondylar region between this and the tendinal fossa of the shaft
forming a narrow lip. The anterior part of the outer surface of the external condyle is hollowed, but there
is a large, projecting, external ligamental prominence towards the proximal edge of the condyle. A deep
groove extends along the anterior side of the external edge of the shaft, ending abruptly at the external
ligamental prominence, at a level with the distal edge of the tendinal fossa. Posterior to this groove a narrow
ridge extends along the middle of the internal surface of the shaft. The anterior surface of the shaft is rela-
tively smooth, a large rounded tendinal fossa occupying most of the distal end of the shaft. This fossa is
nearer the external side and leaves a thick ridge on the internal side extending to the proximal base of the
internal condyle where it bears a small elongated pit. The fossa tapers a little towards the proximal end
where it is shallowest and distally deepens to a point just proximal to the condyles. Small areas within the
fossa appear to have been crushed, and detail is more satisfactory on the other species, but there is a small
566
PALAEONTOLOGY, VOLUME 18
rounded hollow at the distal internal corner, slightly undercutting the intercondylar region bordering the
internal condyle, and another broader and more shallow just undercutting the internal side of the proximal
edge of the external condyle.
The edge of the fossa is rounded on the internal side but a narrow ridge borders it on the external side,
slanting externally towards the external ligamental prominence and separating the fossa from the peroneus
groove. From comparison with the other specimen it appears that this ridge should form a thick structure,
its upper surface slanting internally, just proximal to the external condyle, but on the present specimen
crushing has produced a double ridge with a hollow between. There is also a small area of crushing towards
the proximal external end of the main narrow ridge. The internal side of the shaft is rounded but much
thicker than the external side, and this thickness increases proximally to a point where, on the tibiotarsi
of strigiform species, a prominence is present on the anterior internal edge. The shaft of the present specimen
appears to have been broken at the point where this prominence occurs.
Measurements. Length from proximal end to distal tip of external condyle 68-7, to internal condyle 60-5.
Width across posterior edges of condyles 33-5, across distal ends 35-2, across anterior edges 37-8. Distal/
proximal depth of internal condyle 16-9, of external condyle 16-4. Anterior/posterior thickness of internal
condyle 20-9, of external condyle 2 TO. Length of anterior intercondylar fossa 121. Distal end of shaft,
internal to external edges 21-7, thickness on internal side 12-6, on external side 8-7, greatest length of
tendinal fossa 24 0, greatest width 21-4. Height of external ligamental prominence 4-5 mm.
HEPTASTEORNis gen. nov.
Type species. H. andrewsi sp. nov.
Diagnosis. Distal end of tibiotarsus less broad and less flattened antero-posteriorly
than in Bradycnemis. Distal projection of external condyle beyond internal condyle
very small. Internal condyle projecting much further anteriorly than external condyle.
Anterior intercondylar fossa less marked than in Bradycnemis.
Heptasteornis andrewsi sp. nov.
Plate 65, figs. 6, 7; Plate 66, figs. 1-7
Etymology. The generic name is formed from the Greek hepta{ = seven), asty- ( = town), and ornis ( = a bird)
in reference to the name of the area of origin, and is feminine. It is named after C. W. Andrews.
Diagnosis. The only known species of its genus.
Material. Holotype: distal end of a left tibiotarsus A 4359, presented by Baron von Nopcsa, 1913. Paratype:
another distal end of a left tibiotarsus, A 1528. Presented by Baron von Nopcsa, 1922.
Occurrence. Szentpeterfalva, Hatszeg, Transylvania, Romania. Maestrichtian (upper Cretaceous). (See
eomments under previous species.)
Description. The holotype is a distal end of a left tibiotarsus, broken off before the proximal end of the
tendinal fossa. The surfaces are in good condition but the external edge is broken away to the external
ligamental prominence, and the specimen had been irregularly fractured along the median axis. The surface
EXPLANATION OF PLATE 65
Bradycneme draculae gen. et sp. nov. Holotype: distal end of right tibiotarsus (A 1588). Stereopairs, x^.
Fig. I, anterior; Fig. 2, external; Fig. 3, posterior; Fig. 4, internal; Fig. 5, distal.
Heptasteornis andrewsi gen. et sp. nov. Paratype: distal end of left tibiotarsus (A 1528). Stereopairs, xf.
Fig. 6, anterior; Fig. 7, internal.
PLATE 65
HARRISON and WALKER, Cretaceous owls
568
PALAEONTOLOGY, VOLUME 18
of the bone shows some very fine ridging and irregular texturing. The posterior surface is flat and smooth,
rounded at the edges. The posterior edges of the condyles show only a very slight prominence along the
outer edges, and the posterior condylar surfaces continue smoothly from the posterior side over the distal
end, with only a shallow intercondylar groove between them. Distally the external condyle projects slightly
beyond the internal condyle, but shows the converse condition to that of Bradycneme in that the distal
surface of the external condyle is shorter and more rounded distally. Anteriorly both condyles are pro-
minent, with a deeper groove between them. The internal condyle projects further anteriorly than does the
external condyle, and the anterior ends of both show some internal deflection. The internal side is smooth
and flat, slightly rounded at its edges; and the internal side of the condyle shows a hollow towards the
anterior edge. On the external side there is some evidence of a projecting ridge which has broken away.
There is a portion of a deep, tapering groove. There is a worn area where a ligamental prominence might
have been, and a hollow on the anterior side of the outer condylar surface. On the anterior intercondylar
fossa, but proximal to it the surface of the groove forms a narrow prominent lip above the distal end of the
anterior tendinal fossa which undercuts it. The broad ridge formed by the anterior surface along the internal
side of the tendinal fossa terminates at the proximal base of the internal condyle. It bears an elongated pit
which appears to have been enlarged by erosion. On the external side of the anterior shaft surface the floor
of the tendinal fossa forms a similar ridge with an inward-slanting surface, the floor of the fossa deepening
markedly in the central part of the shaft. The distal end of the fossa shows a small rounded hollow in the
proximal side of the intercondylar region adjacent to the inner side of the internal condyle, and a similar
hollow proximally undercutting the inner side of the external condyle.
The referred second specimen is worn and eroded over much of the surface and elsewhere shows small
accretions of matrix which obscure detail. Apart from confirming the general configuration of the holotype
it adds little except that, having a longer portion of shaft, it shows the outline of the tendinal fossa, tapering
to a point proximally near the external edge of the shaft.
Measurements. Holotype. Length on internal side 30T, on external side 35-7. Greatest width at distal
end 32-5. Distal/proximal depth of internal condyle 13-9, of external condyle 15-7. Anterior/posterior
thickness of internal condyle 19 0, of external condyle 17-8, of external side of shaft 11-3, of internal side
12-6 mm.
Paratype. Length on internal side 5 10, on external 54-9, greatest width at distal end 33-8, width of
proximal end of shaft 17-2. Distal/proximal depth of internal condyle 18 0, of external condyle 16-3.
Anterior/posterior thickness of internal condyle 19-7, of external condyle 181, of internal side of shaft
1 I T, of external side of shaft lOT. Greatest length of tendinal fossa 27-9 mm.
Discussion. Apart from the flattened condition of the posterior condylar region, the
specimens resemble the tibiotarsi of Recent owls such as Strix. The existence of
giant forms at a period when the other known fauna consists of a huge aquatic bird
and a number of large reptiles would not be surprising. In general, in Recent diurnal
raptors, the broader, more flattened distal end of the tibiotarsus is associated with
species which are relatively sedentary and rely on a rapid flight and swift seizure with
the feet to capture their prey, the narrower bone being usually associated with species
which walk or run more frequently. The difference in shape of the specimens discussed
here might be the result of similar selective pressures.
EXPLANATION OF PLATE 66
Heptasteornis andrewsi gen. et sp. nov. Paratype: distal end of left tibiotarsus (A 1528). Stereopairs, xf.
Fig. 1, posterior; Fig. 2, external; Fig. 3, distal.
Heptasteornis andrewsi gen. et sp. nov. Holotype: distal end of left tibiotarsus (A 4359). Stereopairs, x^.
Fig. 4, anterior; Fig. 5, internal; Fig. 6, posterior; Fig. 7, external; Fig. 8, distal.
PLATE 66
HARRISON and WALKER, Cretaceous owls
570
PALAEONTOLOGY, VOLUME 18
REFERENCES
ANDREWS, c. w. 1913. On some bird remains from the Upper Cretaceous of Transylvania. Geol. Mag.
(5) 10, 193-196.
JELETZKY, j. A. 1962. The allegedly Danian dinosaur bearing rocks of the globe and the problem of the
Mesozoic-Cenozoic. J. Palaeont. 36, 1005-1018.
LAMBRECHT, K. 1929. Mesozoische und tertiare Vogelreste aus Siebenbiirgen. C.R. 10th Congr. Int. Zool.
Budapest 1927. Sect. 8, 1262-1275.
1933. Handbuch der palaeornithologie. xix^ 1024 pp., 209 figs. Berlin.
C. J. O. HARRISON
Subdepartment of Ornithology
British Museum (Natural Elistory)
Tring, Herts.
C. A. WALKER
Department of Palaeontology
Original typescript submitted 24 April 1974 British Museum (Natural History)
Revised typescript submitted 20 November 1974 London SW7 5BD
A NEW 7BRYOZOAN FROM THE
CARBONIFEROUS OF EASTERN AUSTRALIA
by BRIAN A. ENGEL
Abstract. Revision of Australian Carboniferous cryptostome fenestrate bryozoans has resulted in the recognition
of a new genus, Septatopora, which has been defined on the basis of nine species, four of which, 5. flemingi,
S. gloucesterensis, S. nodosa, and 5.(?) williamsensis, are new, with the remaining five species having been previously
assigned to Fenestella Lonsdale or Polypora M’Coy.
The existence of eight apertural septa and an additional orifice on the branch surface proximal to each aperture,
place the affinities of the genus in doubt. Grouping with either bryozoans or octocorals is suggested, with the con-
clusion being drawn that greatest affinities lie with the contemporary genera of fenestrate bryozoans. A new family,
doubtfully positioned close to the Family Fenestellidae King, 1850, is erected to contain the new genus.
A BiosTRATiGRAPHiCAL, taxonomic, and evolutionary study of Australian
Carboniferous fenestrate bryozoans, has led to the recognition of a new, morpho-
logically distinct group of species, previously described members of which have been
distributed generically between Fenestella Lonsdale and Polypora M’Coy.
Division of Australian species between these two taxa, based largely upon the
number of rows of zooecial apertures per branch, has been found to be impractical.
There exists a distinct evolutionary trend throughout the Carboniferous Period for
all initially two-rowed fenestrate species to develop a third row of apertures at an
increasing distance proximal to each branch bifurcation. One result of this is that
it is no longer possible to decide if some Mid to Upper Carboniferous species are
basically two- or three-rowed forms. This problem has already been raised in the
case of Fenestella{l) altinodosa Campbell (Campbell 1961) where that author sug-
gests his species could equally well be placed in Polypora M’Coy.
As a result of an extensive statistical survey in the present study of numerous
Australian Carboniferous fenestrate specimens it became apparent that there were
variations in apertural form, in conjunction with several other features, which pro-
vided a more satisfactory grouping of the Australian material. In particular, three
basic types of aperture were recorded, namely :
1. Fenestellid type— a simple, weakly exserted, circular aperture with a narrow
peristomal rim. Mean apertural diameter lies between 0 08 and 0T5 mm.
2. Polyporid type— a larger, simple aperture with a broad, low, peristomal collar
which may become horseshoe-shaped in some species. Mean apertural diameter is
usually about OT 4-0-23 mm.
3. Septate type— a circular, strongly exserted aperture with a thin, high peristome
within which there are eight radially disposed septa surrounding a very small central
orifice. Mean apertural diameter ranges between 0 07 and 013 mm.
This last group was also found to share several additional morphological features
which together define the new genus described here as Septatopora gen. nov.
[Palaeontology, Vol. 18, Part 3, 1975, pp. 571-605, pis. 67-70.]
I
572
PALAEONTOLOGY, VOLUME 18
The geological range of this new genus commences in strata which can be cor-
related, on the basis of other fauna, with the Tournaisian-Visean boundary. It
extends up through the remainder of the Australian Carboniferous sequence but
has not yet been recorded from the overlying Permian strata. For purposes of brevity
the following morphological discussion will refer to low, mid, and high zonal dis-
tribution, each of which correlates approximately with Lower Visean, Upper Visean-
Namurian, and Westphalian-Stephanian respectively. More specific stratigraphic
data are given with the systematic descriptions.
A total of nine species, two of which are of dubious relationship, are here assigned
to the new genus :
Septatopora pustulosa (Crockford) 1949
Septatopora flemingi sp. nov.
Septatopora isaacsensis (Campbell) 1961
Septatopora stellaris (Campbell) 1961
Septatopora(l) sulcifera (Crockford) 1947
Septatopora gloucesterensis sp. nov.
Septatopora acarinata (Crockford) 1947
Septatopora nodosa sp. nov.
Septatopora(l) williamsensis sp. nov.
[= Polypora pustulosa]
[= Polypora isaacsensis]
Fenestella stellaris]
[= Polypora sulcifera]
[= Fenestella acarinata]
The stratigraphic distribution of these species is illustrated in text-fig. 2.
DIAGNOSTIC MORPHOLOGY OF SEPTATOPORA
Apart from the fact that all species have a standard cryptostome fenestrate mesh-
work with a normal zooecial chamber/vestibule arrangement, the following are the
four major, additional generically-distinguishing morphological features:
Septation. All apertures are strongly exserted and contain eight apertural septa
which commence on the sides of the vestibule from where they taper upwards and
inwards towards the axis to leave only a small circular opening in the centre of the
external aperture. Each aperture also bears a narrow, elevated peristomal collar
which gives it a cup-like form very similar to the calice of some solitary corals.
Auxiliary tube. In low zonal species the proximal side of the exserted aperture has
a small opening or gap on to the obverse branch surface. This detail is quite difficult
to observe in the very fine meshwork of these older species.
Upper zonal species have an obvious small, conical or slit-like depression situated
some distance proximal to each aperture on the branch surface. This depression bears
the surface ornament of the branch and is connected by a narrow auxiliary tube to
the proximal region of the elongated vestibule just anterior to the hemiseptum.
Position and orientation of the auxiliary tube vary according to the form of the
zooecial chamber.
Ovicellular structures. Most species have additional large, irregularly spaced, hemi-
spherical depressions on the branch surface. When present, they are situated adjacent
to the proximal rim of an aperture where they obliterate the smaller conical depres-
sion. The surface of these larger depressions is smooth and they are also connected
ENGEL: CARBONIFEROUS SEPTATOPORA
573
TEXT -FIG. 1. A, B, side-sectional diagrams along one row of zooecial apertures in a branch showing zooecial
chambers, hemisepta, septate vestibules, auxiliary tubes, and surface ovicellular depressions, (a, Septatopora
flemingi, x 75; B, Septatopora acarinata, x 60.) c, D, E, reverse views of the method of packing of zooecial
chambers immediately beneath the back wall of the branch, c illustrates a low zonal species with only one
additional aperture appearing at bifurcation, d is a mid-zonal species with a third row of apertures appear-
ing some distance before bifurcation, e demonstrates the change in zooecial packing in the late Carboni-
ferous forms, (c, Septatopora acarinata, x 25; D, Septatopora ghucesterensis, x 25; E, Septatopora flemingi,
x20.)
to the lower vestibule by the auxiliary tube. They may be the sites of former external
ovicellular chambers.
Ornamentation. Most species lack carina and are ornamented with fine, pustulose,
sinuous, longitudinal striations of distinctive appearance.
The above features define a morphologically compact species group dilferent from
other fenestrate taxa. Several other variable features have proved also to be of con-
siderable stratigraphic value. They are detailed in the comparative discussion which
follows the description of each species.
BRYOZOAN OR OCTOCORAL?
The classification of Septatopora gen. nov. presents numerous difficulties which can-
not be resolved on the basis of evidence at present available. The novel occurrence of
eight septa in the vestibule of a form with a fenestrate bryozoan habit combines
aspects of both bryozoan and possibly octocorallian affinities, a final decision between
which must await further detailed thin section study.
Unfortunately, with rare exceptions, Eastern Australian Carboniferous fenestrate
species are preserved in fine clastic sediments as either internal or external moulds,
the original calcareous skeleton having been leached away or perhaps replaced by
structureless secondary mineral deposition (calcite or hydrous silica). Consequently,
almost no information is available on skeletal microstructure. Despite this serious
deficiency, it is possible to reconstruct from the moulds many of the important struc-
tural details, some of which would be quite difficult to observe on complete specimens.
574
PALAEONTOLOGY, VOLUME 18
Amongst the rare material suitable for thin section work, all sections made so far
have revealed a standard fenestellid microstructural arrangement (Tavener-Smith
1969; Tavener-Smith and Williams 1972). Unfortunately most of these sections have
not been identifiable generically, and hence it is not possible to be certain that
specimens of Septatopora have been included, although by their frequency, this is
thought to be quite probable. Some internal moulds of Septatopora exhibit short
skeletal rods which extend from the base of the zooecial chambers out to the side and
reverse walls of the branch. These rods are almost certainly a replacement of the
skeletal rods which occur normally in the laminated wall tissue of all fenestellids,
lending further support to the supposition that the microstructure of Septatopora
is of the fenestellid type.
Despite the major problems which will arise consequent upon the decision, the
writer is of the opinion that Septatopora must be classified with its contemporary
fenestellids. In recognition of its distinct morphology, the genus has been placed
herein in a separate family and, with slight reservation, grouped most closely with the
Family Fenestellidae pending the resolution of the generic microstructural details
of the new genus. Some of the reasons for this decision are given below.
Growth habit. Where known, species of Septatopora have a broadly funnel-shaped
or flared zoarium structurally identical with contemporary fenestellids. Apertures
are arranged in regular rows along the branches on the inner surface of the cone.
Exact equivalence of so many structural aspects is so great that an explanation of
similarity based upon convergence from separate phyla is regarded as being highly
improbable. Both FenesteUajPolypora and Septatopora also exhibit the same evolu-
tionary trends throughout the Carboniferous in the development of their zoaria,
some details of which are discussed later.
In Lower Carboniferous species septation is the major visible distinguishing
feature, and in the case of poor preservation of this aspect, it is not possible to make
a generic decision between Septatopora and Fenestella. It is only in the much larger,
late Carboniferous species that the septation and auxiliary tube become readily
evident, but even there the similarity of form is still very clear.
Growth habit in the octocorals is of extremely wide variation and a fenestrate form
is known in a number of groups (e.g. gorgonids). No examples have been observed
of the regular funnel-shaped zoarial form, and, although of limited significance, size
differences between this group and Septatopora are of quite major proportions.
Septation. The existence of eight apertural septa is considered to be the main argu-
ment against a bryozoan origin for Septatopora. Modern ideas of the lophophore and
gut of a bryozoan would appear to be incompatible with septation. It is not possible
to argue this matter without further details of the skeletal microstructure of the
vestibular region.
It should be pointed out that the septation is generally much shorter (longitudinally)
than the vestibule in which it is housed and there appears to be no difficulty with the
protrusion of the tentacles between the septa. In their fully extended mode, the
tentacles would fill the calice-like external aperture and raise the mouth to a position
beneath the central orifice in the base of the calice. To do this requires some slight
invagination of the tentacle ring which in turn would provide a suitable secreting I
ENGEL: CARBONIFEROUS SEPTATOPORA
575
surface for the septal development. Such possible modifications to the lophophore
require further investigation.
An additional aspect of septation concerns the type species of Polypora M’Coy
{P. dendr aides M’Coy) which has been redescribed by Miller ( 1 963 ) as having apertures
with ‘fifteen or sixteen short thin internal projections resembling the septa of corals’.
Although of only slight form, their presence and number is highly suggestive, and
lends some possible support to the argument that Septatopora should be grouped
with these fenestrates.
Internal form. In spite of a lack of thin-section detail it is possible to establish that,
internally, Septatopora is quite different from most octocorals of comparable
arborescent form.
All species of Septatopora have a calcified skeleton in which the body chambers
are regularly packed in contact with each other in rows adjacent to the thin reverse
wall in a fashion identical with that of the fenestellids. These body chambers almost
fill the branch having some variable skeletal thickening surrounding them.
In the space available in each branch it is quite impossible to develop an inner
coenenchymal (medullar or axial) zone with an outer layer in which the chambers
are shallowly embedded, as is a common condition in the gorgonid octocorals. It
would appear that there is a variety of similar basic structural differences between
Septatopora and most arborescent groups of living and fossil octocorals which would
make their combination improbable.
Finally, all zooecia in Septatopora are distinctly subdivided into a body chamber
and a vestibule separated by a marked hemiseptum. This dual chamber arrangement
appears to have no modern analogue in the octocorals but is a well-established
bryozoan feature.
Septation remains the most difficult aspect of the new genus to encompass within
modern ideas on bryozoans. Despite this problem, the case has been argued above
that Septatopora is basically inseparable from its contemporary fenestellids with
which it closely approximates in both structure and form. It would seem that if
Septatopora is unacceptable within the Phylum Bryozoa, then further close investiga-
tion must be made of the systematic position of the Family Fenestellidae.
FUNCTIONAL MORPHOLOGY
Whilst lacking any clear understanding of the reasons behind the development of
apertural septation in all species of Septatopora, it is readily evident that the polypide
was greatly restricted in its ability to extrude out of the zooecial cavity. In the fully
extended mode, the tentacles would have been placed between the septal partitions
and the mouth must have been located beneath the small central opening in the base
of the calice-like depression. Of necessity, the tentacle ring or lophophore was thus
contained within the vestibule. Assuming the genus was a normal ectoproct, this
means that the anus, being outside the lophophore, would also have been enclosed
within the vestibule. To overcome this major problem, the development of a separate
anal opening would seem to have been an essential requirement.
In low zonal species with their globular zooecial chamber close to the obverse
surface it is postulated that this was initially achieved by the simple development of
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PALAEONTOLOGY, VOLUME 18
a breach in the side of the exserted vestibule, or by the construction of a short, narrow,
horizontal connection from the base of the vestibule to the branch surface on the
proximal side of the aperture. In high zonal species the chamber became elongate
oval in form and located quite close to the reverse branch surface. This required the
elongation of both the vestibule and the auxiliary ‘anal’ tube, each of which then
developed as quite separate structures situated perpendicular to the branch surface.
From the simple expedient solution of a lateral breach in the side of the exserted
aperture, changes in chamber shape and position would thus have inevitably resulted
in the need for the elaborate auxiliary tube as can be observed in the late Carboni-
ferous representatives of the genus. It is perhaps not surprising that such a complex
arrangement apparently did not survive beyond the Carboniferous Period.
It also appears reasonable to postulate that the reproductive system would have
used this auxiliary tube for the release of fertilized ova which were then stored in
enlarged spherical chambers on the branch surface, prior to final release. This would
explain the coincidence of the auxiliary tube opening with the frequent, large hemi-
spherical depressions observed on the branch surface adjacent to the proximal rim
of selected fertile polypides in most zoaria.
The assignment of such an alimentary/reproductive role to the auxiliary tube
presents several major difficulties. It is customary to extrapolate backwards from
modern functional morphology to fossil morphology and unfortunately there appears
to be no modern equivalent which can lead to the above interpretation. Modern
calcified bryozoans have a budding pattern in which the anus is distally placed in
the tentacular crown and given this information it is quite difficult with the budding
pattern in Septatopora to postulate a proximal anus. Study of all species of Septatopora
makes it quite evident that the auxiliary tube connects to the base of the vestibule
adjacent to the hemiseptum making it obligatory to propose a proximal anus if the
above proposed theory is to have any substance. In addition, modern species have
their coelomic pores for egg extrusion placed distally and the transfer of eggs to the
distally positioned brooding cavities requires a great deal of movement and manipula-
tion on the part of the tentacle crown, a process clearly not possible from the base
of the vestibule. Finally, external proximal ovicells are unknown in modern forms
although little doubt is held that this is the only likely interpretation of these large,
spherical, proximally situated structures in the fossil species.
From the above discussion it is apparent that an alimentary/reproductive role for
the auxiliary tube requires a major reversal of the polypide construction from fossil
to modern species. Although the proposition remains attractive, the absence of
skeletal detail makes it impossible to arrive at a positive conclusion.
A second suggestion that the auxiliary tube served an hydrostatic function, with
the surface opening being analogous to an ascopore is possible. This structure is
defined by Bassler (1953) as a ‘median small opening in the frontal wall of some
cheilostomes leading to the compensatrix, located proximally with reference to the
aperture’. There is a close similarity with this cheilostome feature and the opening
found in Septatopora making it necessary to propose an hydrostatic function as an
alternative explanation. However, since the polypide was unable to move very far
because of the septation, it is unlikely to have developed the need for any great degree
of compensation. The term ‘ascopore’ has been used for several non-homologous
ENGEL: CARBONIFEROUS SEPTATOPORA
577
systems in bryozoans and it is possible that other hydrostatic functions than that of
compensation may have been the role of the auxiliary tube.
Although of major significance to the interpretation of the new genus, a lack of
thin-section data and modern analogous structures means that a satisfactory explana-
tion has yet to be achieved for the biological function of the apertural septation and
its closely associated, proximally situated auxiliary tube.
Abbreviations. The following abbreviations are used in the statistical treatment of fenestrate mesh. FL =
fenestrule length (centre to centre of dissepiments) ; FW = fenestrule width (centre to centre of branches) ;
BW = branch width ; DW = dissepiment width ; ZD = zooecial diameter ; Z-Z = zooecial aperture spacing
(centre to centre of apertures); N-N = nodal spacing; F/10 = number of fenestrules in 10 mm; B/10 =
number of branches in 10 mm; Z/5 = number of zooecial apertures in 5 mm; N/5 = number of nodes in
5 mm ; Z/F = number of zooecial apertures per fenestrule ; m = arithmetic mean value of dimension ;
s = standard deviation; OR = observed range of dimension; N = number of measurements recorded.
Repositories. All specimens have had their catalogue number prefixed by the letter ‘F’, preceded by the
following Museum coding: QU = Queensland University; QGS = Queensland Geological Survey;
NEU = University of New England; NU = University of Newcastle; SU = Sydney University.
Most specimens recorded by Crockford (1947, 1949) in the University of Queensland Catalogue have
had new numbers allocated since their original publication.
Fossil localities. Localities are recorded, wherever possible, with the prefix NUL, followed by a number,
all of which refers to the University of Newcastle Fossil Locality Index. Localities not present in that index
are given in descriptive detail in the text. NUL9— 3 km east of Booral, N.S.W. (Campbell 1961); NUL39—
Cameron’s Bridge, Rouchel Brook, N.S.W. (Crockford 1947); NUL258— Barrington, N.S.W. (Cvancara
1958); NUL361— Glen William, north of Clarencetown, N.S.W. (Crockford 1947); NUL372— Hilldale,
N.S.W. (Crockford 1947); NUL390-Oaky Creek, N.S.W. (Campbell 1962); NUL4 14- Barrington Guest
House, N.S.W. (Crockford 1947); NUL448— Raglan property, east of Dungog, N.S.W.; NUL454—
Isaacs Road, Dungog 1 mile Military Map (Grid Reference 019840), N.S.W. (Campbell 1961); NUL472 —
Ridgelands 1 mile Military Map (Grid Reference 194827), Queensland (Fleming 1969).
Photographic methods. Because most specimens used in this study occur as external moulds, it was generally
necessary for photographic purposes to prepare latex casts which were firstly painted with a uniform black
coating over which a thin grey-white layer of ammonium chloride was deposited. Depending upon the size
of the specimen and the magnification required, photographic negatives were produced with a stand-
mounted, close-up camera, or with a camera attached to a stereobinocular microscope.
SYSTEMATIC DESCRIPTIONS
Phylum ?BRYOZOA
Order ?cryptostomata Shrubsole & Vine, 1882
Family septatoporidae fam. nov.
Type genus. Septatopora gen. nov.
Family diagnosis. ?Cryptostomata with zoaria of a broadly flared funnel shape with
obverse surface on inner side of cone; composed of sub-parallel to radiating non-
carinate branches connected by non-celluliferous dissepiments into a regular fenestrate
meshwork; branches with two to four rows of small to medium-sized, strongly
exserted calice-like apertures; apertures bear eight septa which taper upwards and
inwards from the sides of the vestibule, converging in the base of the external aperture
on a small central orifice ; branch surface with fine pustulose, striate ornament ; nodes
present or absent, but when developed, irregularly placed on the distal rim of some
578
PALAEONTOLOGY, VOLUME 18
apertures; hemiseptum distinct; base of vestibule connected to branch surface where
a second opening is located on the proximal side of each aperture.
Geological range. Upper Tournaisian to Stephanian.
Remarks. This family has been erected solely for the reception of the new genus
Septatopora which cannot readily be combined with any other existing taxon. For
reasons outlined in the previous discussion, the new family is regarded as belonging
to the Order Cryptostomata Shrubsole & Vine, 1882, and is doubtfully grouped
closest to the Family Fenestellidae King, 1850, with which it would appear to share
the greatest number of common features.
Genus septatopora gen. nov.
Type species. Septatopora pustulosa (Crockford) (= Polypora pustulosa Crockford, 1949).
Generic diagnosis. See family diagnosis given above.
Generic description. Septatoporid with zoaria of a flared funnel shape (where known)
with obverse surface on inner side of cone; fine to medium-sized fenestrate mesh
composed of narrow to wide, finely striated, pustulose branches, each having
a rounded, non-carinate cross-section; fenestrules oval to sub-rectangular being
moderately to strongly indented by zooecial apertures; apertures small to medium-
sized, circular, septate, strongly exserted, being surrounded by a thin, high, complete
peristome of calice-like form; apertures in two to four rows per branch with increase
in number before, and decrease after, each branch bifurcation; nodes absent or
irregularly developed adjacent to the distal or disto-central rim of some apertures in
any zooecial row ; arrangement may appear more regular in branches with only two
rows of zooecia.
Zooecial chambers globular to elongate oval in form, being joined on their distal
margin to curved or L-shaped vestibules respectively; distinct hemiseptum at base
of vestibule ; each vestibule with eight short, radially disposed septa extending upwards
and inwards from the sides of the vestibule to converge on a narrow axial opening in
the centre of the external aperture, which thus assumes a rosette pattern on the base
and sides of the calice-like depression.
Additional small, funnel-shaped depressions located on the branch surface,
proximal to each aperture; a narrow tube connects this orifice to the base of the
vestibule just anterior to the hemiseptum; irregularly disposed, larger, smooth,
hemispherical depressions may occur on the branch surface where they replace the
funnel-shaped depressions and abut against the proximal rim of an aperture.
Reverse surface of rounded, pustulose branches joined by narrower, near level,
sometimes inclined dissepiments; some reverse branches may also bear numerous,
irregularly arranged spines. Zooecial bases irregularly pentagonal in lateral rows and
rhomboidal in central rows; with increase in the number of zooecial rows they become
elongate oval with little overlap between rows.
Geological range. Upper Tournaisian to Stephanian.
Generic comparisons. The reasons for excluding Septatopora from either Fenestella
ENGEL: CARBONIEEROUS SEPTATOPORA
579
Lonsdale or Polypora M’Coy have been discussed previously. No other genus of
comparable form has been observed in the literature.
It is possible that the species discussed by Miller (1963) as Polypora(l) verrucosa
(M’Coy) could be assigned to this genus. In his revision of Polypora M’Coy, Miller
excluded this species on the basis of its stalk-like apertures, but was not prepared at
that stage to designate a new generic category until similar morphology had been
observed on further material. In its apertural form, the species is quite similar to
Septatopora, but until further examination of the original material is made it is not
possible to confirm its inclusion.
Septatopora piistulosa (Crockford), 1949
Plate 67, figs. 1-9
1949 Polypora pustulosa Crockford, p. 426, text-fig. 9.
1949 Polypora tenuirama Crockford, p. 428, text -fig. 1 1.
1961 Polypora septata Campbell, pp. 462-463, pi. 58, figs. 1-2.
1962 Polypora cf. septata Campbell, p. 47, pi. 13, fig. 8a-c.
1972 Polypora pustulosa Crockford; Fleming, pp. 6-7, pi. 3, fig. 8; pi. 4, figs. 1-6; text-figs. 1-2.
Revised diagnosis. Septatopora with wide pustulose branches ; mesh open, medium-
sized, with sub-oval to sub-rectangular fenestrules; apertures septate, strongly
exserted, distantly spaced, with frequent, proximally associated, auxiliary pits and
hemispherical depressions; zooecia in three rows per branch; carina and nodes
lacking; reverse branch profile rounded; zooecial bases elongate oval.
Revised description. Zoarium : gently radiating to sub-parallel branches of maximum
radius 90 mm; orientation unknown. Obverse surface, {a) Branches. Wide, normally
with three rows of zooecia (m.BW 0-49 mm) ; two-rowed branches medium to narrow
(~0-3 mm) and four- to five-rowed branches very wide (~0-8 mm); branch cross-
section rounded, becoming oval at bifurcations; ornament of distinctive pustules
arranged along slightly wavy, faint, longitudinal ribbing, {b) Dissepiments. Medium
to wide (m.DW 0-21 mm); expanded gently outwards from centre to branch junction
in a broad curve; level with or slightly below branches; most dissepiments inclined
towards base of colony; ornament as on branches, (c) Fenestrules. Sub-oval to sub-
rectangular; medium-sized mesh moderately regular but varied by regions of wide
pre-bifurcation, and narrow post-bifurcation branches; fenestrules wider than
branches, resulting in an open-mesh appearance; fenestrules medium length (m.FL
1-68 mm), medium to wide (m.FW 1-03 mm), {d) Carina. Absent, {e) Nodes. Absent.
if) Zooecial apertures. Medium size, circular (m.ZD 0- 1 1 mm) ; strongly exserted
being surrounded by high peristome; apertures bear eight radially disposed septa
surrounding a narrow axial opening which widens downwards towards the zooecial
chamber; erect or laterally inclined apertures arranged in straight rows with alter-
nating positions in adjoining rows; marginal apertures strongly indent fenestrules
and are not stabilized with respect to the dissepiments ; apertures in each row medium
to distantly spaced (m.Z-Z 0-43 mm) with from three to five zooecia per fenestrule
(m.Z/F 3-9); usually three rows per branch with increase to four (rarely five) rows
up to 2 mm before, and decrease to two rows up to 2 mm after bifurcation, (g) Addi-
tional features. Proximal to each aperture there is a small, funnel-shaped pit which
580
PALAEONTOLOGY, VOLUME 18
bears normal, slightly deflected branch ornament; a narrow tube connects the bottom
of this pit to the base of the vestibule; some specimens also bear irregularly dis-
tributed, smooth, hemispherical depressions (diam. 0-25-0-35 mm) on the obverse
branch surface, situated so that the distal rim of the depression is in contact with the
proximal margin of an aperture; an opening at the base of the depression may be
visible, being the site of the tubular connection.
Reverse surface, {a) Form. Broadly rounded branches joined by medium width, level
dissepiments; both branches and dissepiments bear fine, pustulose striations.
{b) Zooecial bases. Elongate oval with little or no overlap between adjoining rows of
zooecia.
Material. Holotype QUF24980; topotype QGSF10910 (For. 2v. Ph. Stan well);
others QGSF11920, QGSF10936 (Neerkol Creek); QGSF10905, QGSF10988-
10989, QGSF10889 (Malchi Creek); QUF24954-24955 (Malchi Creek);
NEUF4708D, NEUF4715C/D, NEUF4734C/D (NUL9); NEUF5632-5634
(NUL390). Further material from various localities in New South Wales and Queens-
land has been placed in the University of Newcastle Collection.
Remarks. Fleming (1972) first proposed that the three species P. pustulosa Crockford,
P. tenuirama Crockford, and P. septata Campbell should be combined into the one
species. Concurrent and subsequent detailed mesh studies by the present writer
strongly support this decision. However, the work of Miller (1963) clearly indicates
that the Australian material cannot be assigned to the genus Polypora M’Coy, and
hence a new genus is proposed here for its reception.
Based upon Queensland specimens, Crockford (1949) erected both her species in
the one paper, but made no comparative remarks. Distinction appears to have been
based upon absolute differences in mesh size and zooecial spacing. No mention was
made of apertural septation in P. pustulosa, no doubt because of the very poor state
of preservation of the type material.
Campbell (1961) erected the third species P. septata upon specimens from New
South Wales, and hesitantly distinguished it from P. tenuirama because of small dif-
ferences in mesh dimensions and apertural septation. Further specimens from another
New South Wales locality (Campbell 1962) were referred to P. cf. septata because of
their weak development of apertural septation, a reduced branching frequency, an
EXPLANATION OF PLATE 67
All figures of latex casts except fig. 7.
Figs. 1-9. Septatopora pustulosa (Crockford). 1-4, obverse surface of holotype of Polypora septata Campbell,
NEUF4708D, x 20, x 30, x 30, x 50 respectively. Note auxiliary openings proximal to each aperture
on figs. 1 and 2 and strongly exserted, septate form of apertures together with the distinctive pustulose
ornament on branch surfaces in figs. 3 and 4. 5, 6, obverse surface of holotype of Polypora tenuirama
Crockford, QUF24955, x20, x 10 respectively. Note frequent occurrence of proximal ovicellular pits.
Weaker occurrence of surface ornament is due to preservation. 7, reverse view of zooecial chamber
infillings of P. tenuirama Crockford showing skeletal rods extending between chamber and walls,
QUF24955, x5. 8, obverse surface of QGSF10988, x5. 9, obverse surface of QGSF11920, x 5.
(Figs. 1-4 from locality NUL9, Booral, N.S.W.; figs. 5-9 from Malchi Creek, Queensland.)
PLATE 67
ENGEL, Septatopora
582
PALAEONTOLOGY, VOLUME 18
increase to five, rather than four zooecial rows before bifurcation, and the develop-
ment of surface hemispherical depressions not previously observed on P. septata.
Detailed measurement by the present writer on a wide variety of specimens from
both states, including all type material, has established that all three species belong
to the one continuously expanding mesh series, and that there are no grounds for
subdivision upon this basis. Furthermore, all material bears the diagnostic obverse
features of pustulose ornament, apertural septation, and separate auxiliary orifices,
and is therefore considered to be conspecific.
All authors have recorded the occurrence of surface hemispherical depressions
(?ovicell sites) but Fleming (1972) also noted a few of the smaller openings on one
specimen (holotype of P. tenuirama) and described them as incompletely formed
ovicellular structures. Close examination of most specimens reveals this structure to
be associated with all apertures as a basic functional feature.
Septatopora pustulosa (Crockford) has been chosen as the type species of the new
genus because of its widespread, common occurrence in Eastern Australia. Further,
it is generally of sufficiently coarse mesh to enable easy recognition of the diagnostic
features of the genus.
Stratigraphy. S. pustulosa (Crockford) has a wide geographic and stratigraphic range
through the Levipustula levis Zone in Australian Upper Carboniferous sediments.
It is found intermittently through the whole thickness of the Neerkol Formation
(2100 m) near Rockhampton, and in the Poperima/Rands Formations (Maxwell
1964) in the Yarrol Syncline of Queensland. Occurrences in New South Wales are
more restricted but are found at various uncorrelated levels in the Booral Formation
(2000 m) and in the Kullatine Formation as recorded by Campbell (1962).
The age of the L. levis Zone is generally considered to be Westphalian but no
evidence exists for more precise correlation.
Septatopora flemingi sp. nov.
Plate 68, figs. 4-8; text-fig. 1a, e
Diagnosis. Septatopora with very wide, weakly pustulose branches; mesh closed.
EXPLANATION OF PLATE 68
All figures of latex casts except figs. 6-8.
Figs. 1-3. Septatopora{l) williamsensis sp. nov. 1, 2, obverse surface of holotype showing lateral position
of partly exserted apertures and the wide expanse in centre of branch without carina or nodes, NUF2421a,
locality NUL414, x 10, x30 respectively. 3, reverse surface of holotype, NUF2421b, x 10.
Figs. 4-8. Septatopora flemingi sp. nov. 4, obverse surface of holotype showing weakly exserted apertures
together with their prominent adjacent auxiliary openings which form a longitudinal furrow between
apertures in each row, NUF2357, locality NUL472, x 10. 5, obverse surface of NUF2362a, locality
NUL472, X 5. 6, 7, oblique reverse view of an eroded zoarium with reverse wall removed. Visible is
a complete internal mould of one chamber. In fig. 7 the upper vestibule is partly obscured by fenestrule
infilling and only the curved and lower horizontal portion can be seen leading back to the distinct hemi-
septum. Immediately below the hemiseptum, the vertical auxiliary tube (shaded) connects the lower
vestibule to the obverse surface below. The body chamber extends to the right of the vestibule where its
termination tends to merge with adjacent infillings, NUF2358, locality NUL472, x 20, X 40 respectively.
8, vertical view of same showing one complete chamber infilling from the reverse with adjacent infillings
being broken off to leave the septate moulds of the upper vestibules, NUF2358, x40.
PLATE 68
ENGEL, Septatopora
584
PALAEONTOLOGY, VOLUME 18
medium-sized with sub-oval to sub-rectangular fenestrules; apertures septate,
moderately to weakly exserted, distantly spaced with very strong development of
proximal auxiliary pits; hemispherical surface depressions rare; zooecia in three or
four rows per branch ; carina and nodes absent, but with pseudo-carinal relief between
zooecial rows; reverse branch profile rounded; zooecial bases elongate oval.
Description. Zoarium: moderately radiating to sub-parallel branches of maximum
radius 40 mm; zoarial margins crenulate; orientation unknown. Obverse surface,
{a) Branches. Wide to very wide (m.BW 0-54 mm), straight, usually with three to
four rows of zooecia; branch cross-section round to oval, well-preserved branches
bear sinuous, faintly pustulose, longitudinal ribbing of moderate elevation; large
auxiliary orifices in the form of strongly depressed elongated pits extend between
adjacent apertures ; each apertural row appears to be located in a linear furrow and
separated from adjoining rows by a weak carinal rise. This relief effect is more
apparent on deflated branch surfaces, {b) Dissepiments. Medium to wide (m.DW
0-3 mm) with gradual expansion to branch junction in a gentle curve; level with or
below branches, they are inclined with the obverse face being directed proximally;
ornament as on branches, (c) Fenestrules. Sub-oval to sub-rectangular ; mesh medium-
sized, generally regular except in regions where several adjacent branches bifurcate;
fenestrule openings narrower than branches resulting in a closed-mesh appearance;
fenestrules of medium length (m.FL 1-65 mm) and medium width (m.FW 0-91 mm).
{d) Carina. Absent, (e) Nodes. Absent. (/) Zooecial apertures. Medium size, circular
(m.ZD 0T3 mm) moderately exserted being surrounded by an entire, low peristome;
each aperture bears eight septa which taper down the sides of the vestibule ; apertures
are erect or laterally inclined, and arranged in three to four rows per branch with
increase to five (rarely six) pre-bifurcation, and decrease to three (rarely two) post-
bifurcation ; apertures alternate in adjacent rows only slightly indenting the fenestrule
margin and are not stabilized with respect to the dissepiments; spacing between
apertures medium to distant (m.Z-Z 0-38 mm) with from three to five zooecia per
fenestrule (m.Z/F 4-3). (g) Additional features. Elongate, proximally directed oval-
shaped zooecial chambers are located close to the reverse surface wall. The long
vestibule is geniculate in form with a short horizontal section being joined by a longer
vertical section. Septa commence above the geniculation, and taper upwards and
inwards towards the axis. The auxiliary tube extends vertically from the horizontal
section of the vestibule immediately anterior to the strong hemiseptum, to join the
branch in a deeply depressed slit-like surface pit of dimensions comparable with those
of the adjacent aperture. Only rare hemispherical surface pits have been observed.
Reverse surface, (a) Form. Branches broadly rounded joined by medium width, near
level or slightly depressed dissepiments both of which generally lack much ornament;
some specimens bear strong, ribbed attachment spines, {b) Zooecial bases. Narrow,
elongate oval in form, with little or no overlap between adjacent rows.
Material. Holotype NUF2357 (NUL472); paratypes NUF2358, 2360a/b, 2361,
2362a/b, 2365, 2366 (NUL472); others NUF2359a/b, 2363, 2364 (NUL472).
Remarks. Although quite similar to S. pustulosa in most mesh dimensions these
ENGEL: CARBONIFEROUS SEPTATOPORA
585
specimens display sufficient morphological differences to justify the erection of a new
species.
Major differences between the two species are that S.flemingi has (a) an extra row
of zooecial apertures in only slightly wider branches. Zooecia are more closely packed
and apertures can be located low on the sides of the branches; (b) apertures which
are somewhat larger but more closely spaced, and less exserted, resulting in reduced
fenestrule indentation; (c) auxiliary tube openings very strongly developed being
sub-equal to the apertures in size. This results in a crowded branch surface of distinctly
different aspect to S. pustulosa where the auxiliary tube openings are still quite small.
The strong depression of the auxiliary tube openings between apertures in S.flemingi
results in linear furrows along the zooecial rows not seen in the other species. The
opposed relief effect of apparent carinae between zooecial rows is another feature
not observed on S. pustulosa.
Stratigraphy. S. flemingi sp. nov. is known only from the top 300 m of the Neerkol
Formation as recorded by Fleming (1969) in association with the Cancrinella levis
Zone. Because of lack of stratigraphic continuity it is not possible to be certain of the
exact age of this late Carboniferous zone. However, the occurrence of an associated
brachiopod-bivalve fauna of late Carboniferous-early Permian aspect would sug-
gest that the C. levis Zone is at least of late Westphalian to Stephanian age.
The zone is known only from localities in the Stan well- Ridgelands and Yarrol
districts of Queensland.
Septatopora isaacsensis (Campbell), 1961
Plate 69, figs. 7-8
1961 Polypora isaacsensis Campbell, pp. 463-464, pi. 63, fig. la-e.
Revised diagnosis. Septatopora with wide, weakly pustulose branches; mesh closed,
medium to fine with oval to sub-oval fenestrules; apertures septate, moderately
exserted, medium spaced with frequent, proximally associated, hemispherical depres-
sions obliterating most auxiliary pits ; zooecia in three rows per branch ; carina absent ;
blunt nodes distantly spaced; reverse branch profile tapered, with ornament of
numerous irregular spines; zooecial bases oval.
Revised description. Zoarium : large zoarium of gently radiating, strongly crenulate
branches; maximum radius of specimen 50 mm; obverse surface faces upwards on
the interior of a broadly flattened cone-like zoarium. Obverse surface, {a) Branches.
Wide (m.BW 0-47 mm), straight, broadly rounded to flattened with weak pustulose
ornament, fb) Dissepiments. Medium to wide (m.DW 0-26 mm); growth expands
from centre to branch junction in a semi-circular curve ; level with, or just below,
branches; no ornament observed, (c) Fenestrules. Oval to sub-oval; medium to small,
moderately regular mesh with small fenestrule openings resulting in a closed
appearance; short to medium length, medium width fenestrules (m.FL 0-91 mm;
m.FW 0-75 mm), {d) Carina. Absent, (c) Nodes. Medium size, circular, blunt nodes
very poorly developed due to large number of hemispherical depressions present ;
irregularly spaced when present (m.N-N 0-94 mm); usually located on distal rim of
an aperture. (/) Zooecial apertures. Medium size, circular to slightly oval (m.ZD
586
PALAEONTOLOGY, VOLUME 18
012 mm); exserted with distinct peristome; eight septa in each aperture with a rela-
tively large central opening; apertures erect, with slight fenestrule indentation, not
stabilized with respect to dissepiments; apertures in each row of medium spacing
(m.Z-Z 0-32 mm) with two to three zooecia per fenestrule (m.Z/F 2-8); three rows
per branch increasing up to five pre-bifurcation, (g) Additional features. Obverse
surface largely covered with many hemispherical depressions (diam. 0-2-0-25 mm)
although smaller auxiliary tube pits can be observed at a few locations.
Reverse surface, (a) Form. Branches tapered becoming narrowly rounded and sub-
equal to near level dissepiments; ornament of strong, variable-size spines arranged
irregularly near the centre line of the branch, {b) Zooecial bases. Form not clear but
have a well-rounded to oval form near the basal plate.
Material. Holotype NEUF4744A/C (NUL454).
Remarks. This species is known only by the holotype. No further material from the
type locality has been found.
5. isaacsensis (Campbell) is the smallest Upper Carboniferous member of the new
genus and can be grouped with S. stellaris (Campbell) and S', gloucesterensis sp. nov.
on the basis of their common development of strong obverse nodes.
Stratigraphy. S. isaacsensis (Campbell) occurs only at one locality in the Isaacs
Formation as described by Campbell (1961). By its association with rare specimens
of Levipustula levis it is considered to be situated high in that zone, but unfortunately
there are no overlying marine faunas at this locality which can be used to fix its
position with any degree of accuracy.
The brachiopod fauna, with which S. isaacsensis is associated, is located about
1500 m above the profuse but vertically restricted development of the L. levis fauna
near the base of the Booral Formation. On rather tenuous grounds therefore, the
fenestrate species is considered to be of late Westphalian age.
Septatopora stellaris (Campbell), 1961
Plate 69, figs. 2-4
1961 Fenestella stellaris Campbell, pp. 456-457, pi. 58, fig. 4a-d.
Revised diagnosis. Septatopora with medium width, finely pustulose branches; mesh
uniform, medium to fine, with oval to sub-oval fenestrules ; apertures septate, strongly
EXPLANATION OF PLATE 69
All figures of latex casts except fig. 1.
Fig. 1. Septatopora{l) sulcifera (Crockford). Obverse surface of holotype, QUF14909, locality For. 21/22,
Ph. Malmoe, Queensland, x 10.
Figs. 2-4. Septatopora stellaris (Campbell). 2, obverse surface of holotype, NEUF4716A, locality NUL9,
X 10. 3, 4, reverse surface of holotype showing tapered branches with equal width to that of the dissepi-
ments in a polygonal mesh. Note reverse surface spines, x 10, x 10 respectively.
Figs. 5-6. Septatopora gloucesterensis sp. nov. 5,6, obverse surface of holotype, NUF2398, locality NUL258,
x20, X 10 respectively.
Figs. 7-8. Septatopora isaacsensis (Campbell). 7, 8, obverse and reverse of holotype, NEUF4744A/B,
locality NUL9, x 30, x 10 respectively.
PLATE 69
: »
%
%
%
ENGEL. Septatopora
588
PALAEONTOLOGY, VOLUME 18
exserted, medium spaced, being associated with profuse, proximal, hemispherical
depressions; zooecia in two to three rows per branch; carina absent; nodes large,
irregular, distantly spaced ; reverse branch profile tapered, with ornament of irregular
spines; zooecial bases unknown.
Revised deseription. Zoarium'. flattened, funnel-shaped expansion with a very small
cone of attachment; maximum radius 55 mm; obverse surface faces upwards or to
the interior of the funnel. Obverse surface, (a) Branches. Medium width (m.BW
0-36 mm), straight, broadly rounded with some deflation; ornament of fine ribbing
and faint pustules, fb) Dissepiments. Medium width (m.DW 0-14 mm); growth
expands from centre to branch junction in a semi-circular curve; level with branches;
ornament of fine ribbing, (c) Fenestrules. Sub-oval to oval; medium to fine mesh;
regular distally but variable proximally ; fenestrule openings of about branch width
resulting in a uniform appearance; short to medium length, medium width fenestrules
(m.FL 0-95 mm; m.FW 0-63 mm), {d) Carina. Absent, (e) Nodes. Large, circular
(diam. 0-15 mm), erect or slightly inclined distally; spacing irregular (m.N-N
0-95 mm) where present, but large areas nodeless; situated distal to an aperture
resulting in zigzag placement on two-rowed branches but centrally placed in three-
rowed branches. (/) Zooecial apertures. Medium size, circular (m.ZD 01 3 mm);
strongly exserted with high peristome; each aperture with eight septa and strong
axial tube; erect apertures moderately indent the fenestrules and are not stabilized
with respect to the dissepiments; medium spacing (m.Z-Z 0-33 mm) with from two
to three zooecia per fenestrule (m.Z/F 2-9); two rows of zooecia per branch with
three rows developing up to half-way between successive bifurcations, but normally
only for about one-third of this distance, (g) Additional features. Obverse surface
covered by profuse hemispherical depressions (diam. 0-2-0-28 mm) situated between
apertures and projecting into the fenestrule margin.
Reverse surface, {a) Form. Branches taper in width to become acutely rounded and
equal in dimension to the near level dissepiments. Reduced branch width is accom-
panied by a wavy to zigzag branch outline which results in an irregular mesh of
rectangular-polygonal fenestrules with no thickening at branch-dissepiment junc-
tions; branch and dissepiment width about 0-2 mm; ornament of fine, longitudinal
ribbing with a pustulose overgrowth; numerous irregularly disposed spines occur
with large variation in size, position, and attitude; most are distally inclined.
{b) Zooecial bases. Poorly preserved in type material, being of oval form some little
distance above the basal plate.
Material. Holotype NEUF4716A/B (NUL9); paratype NEUF4717 (NUL9). Addi-
tional material in the University of Newcastle Collection.
Remarks. Campbell (1961) noted the very different morphology of this species as
compared with other Carboniferous fenestellids, but was guided by the widespread
development of two rows of apertures into placing it in the genus Fenestella Lonsdale.
It is removed here from that genus on the basis of its distinctive apertural features.
There is a considerable degree of similarity between S. isaacsensis (Campbell) and
S. stellaris (Campbell), but they can be readily distinguished by the number of rows
ENGEL: CARBONIFEROUS SEPTATOPORA
589
of zooecial apertures per branch at which taxonomic level this feature is given its
greatest significance.
Stratigraphy. This species comes from low in the Booral Formation, where it is
associated with the principal occurrence of Levipustula levis. To date it has only been
found at localities of similar age in the Stroud-Gloucester Syncline, N.S.W. On this
basis it can only be assigned a probable early Westphalian age.
Septatoporaip.) sulcifera (Crockford), 1947
Plate 69, fig. 1
1947 Polypora sulcifera Crockford, pp. 15-16, pi. 1, fig. 2; text-fig. 13.
Revised diagnosis. Septatopora-\ike species with medium to wide, pustulose branches ;
mesh open, medium sized, with oval to sub-rectangular fenestrules; apertures small,
strongly exserted, closely spaced, without visible septation; carina and nodes absent;
reverse branch profile rounded; zooecial bases unknown.
Revised description. Zoarium : small fragment of radiating branches from base of the
zoarium; orientation unknown; maximum radius 6-5 mm. Obverse surface:
{a) Branches. Medium to wide (m.BW 0-41 mm), straight, slightly flattened (depth
0- 36 mm); ornament of fine, pustulose ribbing, (b) Dissepiments. Small to medium
width (m.DW 0-2 mm); centrally straight with slight expansion at branch junction;
slightly depressed below branch level; pustulose ornament, (c) Fenestrules. Elongate
oval proximally becoming sub-rectangular distally; proximal mesh, medium size,
irregular with an open appearance; fenestrules of medium length and width (m.FL
1- 36 mm; m.FW 0-74 mm), {d) Carina. Absent, (e) Nodes. Absent. (/) Zooecial
apertures. Small (m.ZD 0 07 mm); circular; strongly exserted; no internal structure
preserved; narrow peristome; apertures closely spaced (m.Z-Z 0-29 mm) with from
three to six zooecia per fenestrule (m.Z/F 4-7) but distal portions indicate a dis-
tribution of five to six per fenestrule ; three rows per branch with two or three post-
bifurcation and three to five pre-bifurcation.
Reverse surface, {a) Form. Rounded branches joined by narrow, near level
dissepiments, both of which bear pustulose ornament, {b) Zooecial bases. Unknown.
Material. Holotype QUF14908 (formerly QUF5768c) Riverleigh Limestone For.
21/22, Ph. Malmoe, 8 km NW. of Mundubbera, Queensland.
Remarks. The only known specimen is the holotype which comprises two small,
silicified fragments dissolved from the Riverleigh Limestone near Mundubbera,
Queensland. Both fragments are very close to the base of the zoarium indicating
little significance for the accompanying dimensions. Despite further solution of
limestone no other samples have been recovered from the type locality.
Since morphological details of apertural structures, zooecial bases, and distal
zoarial form are unknown, generic assignment must be conjectural. Based upon the
occurrence of small, strongly exserted apertures on nodeless branches which bear
only a fine ornament of pustulose striations, this species has been provisionally
grouped with Septatopora. This assignment is subject to confirmation by the recovery
of distally located, better-preserved specimens.
590
PALAEONTOLOGY, VOLUME 18
Stratigraphy. The Riverleigh Limestone, located in an isolated fault block, has been
previously correlated with beds which lie just below the Rhipidomella fortimuscula
Zone (Hill 1934; Driscoll 1960). More recent studies by McKellar (1967) and Jull
(1968, 1969) indicate an older age which would correlate either with the Delepinea
aspinosa Zone, or perhaps with the Orthotetes australis Zone. Experience in New
South Wales would indicate this latter age is too old, since extensive sampling has
failed to reveal any multi-rowed fenestrates in beds belonging to that zone. For the
present, a D. aspinosa age is preferred, but no conclusive evidence for this age exists.
Septatopora gloucesterensis sp. nov.
Plate 69. figs. 5-6; text-fig. Id
Diagnosis. Septatopora with medium width, weakly pustulose branches; mesh open,
fine, with sub-oval to sub-rectangular fenestrules; apertures small, septate, strongly
exserted, associated with numerous proximal, hemispherical depressions; zooecia
in two to three rows per branch; carina absent; nodes small, irregularly placed,
moderately spaced, situated on disto-central rim of some apertures; reverse branch
profile rounded ; zooecial bases irregularly pentagonal and rhomboidal.
Description. Zoarium: shallow, cone-shaped zoarium of radiating branches; obverse
surface on interior of cone; maximum radius 20 mm. Obverse surface: {a) Branches.
Straight, narrow to medium width (m.BW 0-3 mm); obverse acutely rounded with
prominent apertures ; ornament of fine pustules and very weak longitudinal ribbing.
(b) Dissepiments. Medium to wide (m.DW 0T8 mm); centrally straight with moderate
expansion to branch junction; profile tapers obversely so as to appear slightly
carinate, (c) Fenestrules. Fine, slightly irregular, open mesh of sub-oval to sub-
rectangular, medium to short length, narrow fenestrules (m.FL 1T2 mm; m.FW
0-58 mm), {d) Carina. Absent, (c) Nodes. Small, erect, irregularly developed,
moderately spaced nodes (m.N-N 0-51 mm) situated adjacent to an aperture on its
distal or central rim; most nodes placed near to centre of branch when present.
(/) Zooecial apertures. Small, circular (m.ZD OT mm); strongly exserted; narrow,
entire peristome ; apertures bear eight septa which radiate from a central perforation ;
erect apertures arranged in straight rows with moderate fenestrule indentation;
mostly stabilized with respect to the dissepiments, on to which they commonly
encroach; zooecia closely spaced (m.Z-Z 0-27 mm) with from four to five zooecia
per fenestrule (m.Z/F 4T); usually two to three rows per branch with increase to four
rows up to 3 mm before bifurcation, (g) Additional features. Hemispherical depres-
sions developed on branch surface between apertures and projecting into fenestrules;
auxiliary tube connection to vestibule visible at base of depression; pronounced
hemiseptum developed in most zooecial chambers.
Reverse surface, (a) Form. Rounded branches joined by slightly narrower dissepi-
ments near branch level; ornament of fine pustules over longitudinal striations.
(b) Zooecial bases. Elongate, irregularly pentagonal in two-rowed branches, becoming
shorter where three or four rows occur; central rows rhomboidal in shape.
Material. Holotype NUF2398 (NUL258); paratypes NUF2383 (NUL258), NUF2425
(NUL414).
ENGEL: CARBONIFEROUS SEPTATOPORA
591
Remarks. This new species has the distinctive apertural features of the genus Septato-
pora. Further, it has considerable, more or less equal, areas of either two or three
rows of apertures per branch, making it transitional from the older two-rowed
species to the younger three-rowed forms. The existence of occasional nodes in
a near central row is a further characteristic of this species.
Stratigraphy. S. gloucesterensis occurs in the Delepinea aspinosa and Rhipidomella
fortimuscula Zones in New South Wales. As such it is the first occurrence of a multi-
rowed fenestrate in the Australian Carboniferous sequence, and it is joined in the
R. fortimuscula Zone by the first representative of the genus Polypora M’Coy. The
incoming of these multi-rowed fenestrates is thus considerably younger than their
development in other parts of the world, but their appearance in the Australian record
is of maximum zonal value.
Septatopora acarinata (Crockford), 1947
Plate 70, figs. 6-8; text-fig. 1b, c
1947 Fenestrellim acarinata Crockford, p. 36, pi. 4, fig. 3; text-fig. 45.
1968 Levifenestella acarinata (Crockford) Wass, p. 87.
Revised diagnosis. Septatopora with narrow, straight to zigzag, pustulose branches;
mesh open, fine, with sub-oval to sub-rectangular fenestrules; apertures small,
septate, closely spaced, strongly exserted with frequent, proximally associated hemi-
spherical depressions; zooecia in two rows per branch; carina and nodes absent;
reverse branch profile rounded; zooecial bases irregularly pentagonal.
Revised description. Zoarium: shallow, cone-shaped zoarium of radiating branches;
obverse surface on interior of cone; maximum radius 20 mm (holotype 6 mm).
Obverse surface, {a) Branches. Narrow, straight or slightly zigzag; rounded profile
without carina; ornament of very fine, pustulose, longitudinal striations (m.BW
0-21 mm), {b) Dissepiments. Medium width centrally with gradual expansion to
branch junction in an expanding curve; some dissepiments inclined from vertical
plane; level with or slightly below branches; ornament of fine striations (m.DW
Oil mm), (c) Fenestrules. Fine, irregular mesh of sub-oval to sub-rectangular short,
narrow fenestrules (m.FL 0-75 mm; m.FW 0-48 mm), {d) Carina. Absent, (c) Nodes.
Absent. (/) Zooecial apertures. Small, circular (m.ZD 0 08 mm); strongly exserted;
narrow, raised, entire peristome; apertures bear eight septa radiating from a minute
axial tube ; mostly erect with some lateral apertures moderately indenting the fenestrule
margin ; apertures generally stabilized with respect to dissepiments on to which they
commonly encroach; zooecia closely spaced (m.Z-Z 0-25 mm) in each row, with
from three to four zooecia per fenestrule (m.Z/F 3 0); two rows per branch with
a third row appearing at or immediately prior to branch bifurcation, (g) Additional
features. Large, hemispherical branch depressions located adjacent to the proximal
rim of some apertures; horizontal lateral connection between exserted portion of
vestibule and proximal branch surface occurs with all apertures; distinct hemiseptum
visible on casts of most zooecial chambers.
Reverse surface, (a) Form. Branches and dissepiments rounded with the latter depressed
592
PALAEONTOLOGY, VOLUME 18
slightly below branch level ; reverse wall very thin ; ornament of longitudinal striations.
(b) Zooecial bases. Irregularly pentagonal.
Material. Holotype SUF7402 (NUL372); paratypes SUF7406 (NUL372); SUF6438
(NUL36 1 ) ; others NUF2478, NUF2483-2485 (NUL448) ; NUF25 1 8-252 1 (NUL39) ;
NUF2402-2403 (NUL414).
Remarks. S. acarinata (Crockford) has long been known as an aberrant species of
Fenestella in that it lacks the generically diagnostic, nodose carina and has septate
apertures. Wass (1968) placed the species in Levifenestella Miller, but this seems
inappropriate, in view of the importance of the nodeless carina of that genus.
As at present defined, S. acarinata extends over a wide range covering much of the
Visean interval. It should be noted that the specimens from the lowest zone have
a much narrower branch profile which appears to zigzag between apertures which
thus dominate the obverse surface. By contrast, specimens from later zones have
a more robust, rounded branch profile on which the apertures appear to have been
superimposed. This variation in branch width has not been given taxonomic status
at this time, although it results in specimens with quite dissimilar appearance.
Because S. acarinata is a very fine-meshed species, it is quite difficult to observe
the auxiliary opening in the side of the exserted vestibule on most specimens. Certainly
the type material is too poorly preserved to enable the description of such features.
Stratigraphy. The species first appears in the Schellwienella cf. burlingtonensis Zone
at Raglan (NUL448) and other stratigraphically similar localities. It is of common
occurrence in the Pustula gracilis Zone at Rouchel Brook (NUL39) where the thin,
zigzag branch form is most common. The species is plentiful in the Orthotetes
australis Zone at Glen William (NUL361) and the type material comes from Hilldale
(NUL372) which is situated low in the Delepinea aspinosa Zone. The final occurrence
of the species is in the Barrington Guest House fauna (NUL414), a little higher in the
above zone, where its range overlaps with the first occurrence of the two- to three-
rowed species, S. gloucesterensis sp. nov.
EXPLANATION OF PLATE 70
All figures of latex casts except fig. 2.
Figs. 1-5. Septatopora nodosa sp. nov. 1, obverse surface of holotype exhibiting an ill-defined central carina,
NUF2386, locality NUL258, x 10. 2, obverse mould of the base of a fan-shaped zoarium showing early
branches bending away from the mesh to anchor the colony to the surface of a brachiopod, NUF2431a,
locality NUL258. x 5. 3, reverse surface with spherical ovicells attached to the sides of the branches
and projecting above the reverse surface level, NUF2525, locality NUL39, x 5. 4, obverse surface of
NUF2524 illustrating strong development of a central carina in a low zonal form, locality NUL39, x 10.
5, obverse surface of NUF2523 showing multiple lateral branches developed from the side of a marginal
branch in the positions normally occupied by dissepiments, locality NUL39, X 5.
Figs. 6-8. Septatopora acarinata (Crockford). 6, obverse surface of NUF2519 exhibiting large spherical
ovicells positioned adjacent to the proximal rim of some apertures. Note also the zigzag obverse appear-
ance of this low zonal form, locality NUL39, x 10. 7, obverse surface of a funnel-shaped zoarium,
NUF2519, locality NUL39, x 5. 8, enlarged obverse surface of NUF2519 showing strong exsertion of
septate apertures and the absence of a central carina and nodes, locality NUL39, x20.
PLATE 70
ENGEL, Septatopora
594
PALAEONTOLOGY, VOLUME 18
Septatopora nodosa sp. nov.
Plate 70, figs. 1-5
Diagnosis. Septatopora with narrow, straight to zigzag, pustulose branches; mesh
open, fine with sub-rectangular to rectangular fenestrules; apertures small, septate,
closely spaced, strongly to moderately exserted with proximally associated hemi-
spherical depressions and/or spherical bodies which project into fenestrules and on
to reverse surface; zooecia in two rows per branch; carina weakly developed or
absent; nodes small, closely spaced in a central row; reverse branch profile rounded,
with spiny ornament ; zooecial bases broadly triangular to pentagonal.
Description. Zoarium : radiating fan-shaped, gently undulose to laminate fragments
of unknown orientation; some zoaria grow rapidly outwards in a fan shape from the
base of the colony, in which specimens several of the outside branches droop away
from the mesh to become recumbent sterile spine-like supporting anchors ; maximum
radius 50 mm. Obverse surface, (a) Branches. Straight or broadly curved, narrow
(m.BW 0-25 mm); rounded with no carina or with low, indistinct median ridge
between closely set nodes; profile strongly modified by apertural exsertion; orna-
ment of fine, pustulose striations. (b) Dissepiments. Narrow (m.DW 009 mm);
centrally straight with gradual expansion close to branch junction; slightly depressed
below branch level; ornament of fine, pustulose striations. (c) Fenestrules. Sub-
rectangular to rectangular; mesh fine, moderately regular, of even appearance;
fenestrules short and narrow (m.FL 0-92 mm; m.FW 0-5 mm), {d) Carina. Lacking
or represented by ill-defined ridge produced by narrowing of the branch profile;
development variable within each zoarium. (e) Nodes. Small, erect, pointed nodes
with rounded or slightly elongated bases; closely spaced (m.N-N 0-28 mm) with
linear distribution in a central row; each node associated with an aperture on its
disto-central rim. (/) Zooecial apertures. Small, circular (m.ZD 0-1 mm), strongly
to moderately exserted; peristome narrow, raised, entire; apertures with eight septa
surrounding a fine axial perforation; septa extend for only a very short distance
down into the vestibule; erect apertures situated on branch shoulder and showing
some slight lateral inclination; apertures indent fenestrule margin and may be
stabilized with respect to the dissepiments on to which they frequently encroach;
closely spaced (m.Z-Z 0-26 mm) with from three to five zooecia per fenestrule
(m.Z/F 3-6); zooecia in two rows with a third appearing only in the fork at each
bifurcation, (g) Additional features. A short horizontal tube proximally connects the
upper exserted vestibule to the branch surface; ovicellular structures uncommon but
when present they are usually located on the branch side in the fenestrule where they
may also project around above the level of the reverse surface. Some specimens bear
strong, ribbed, spine-like projections (diam. c. 0-2 mm) standing erect on the obverse
surface at distant intervals.
Reverse surface, {a) Form. Branches and dissepiments rounded to slightly tapered
with dissepiments at or below branch level; surface of branches and dissepiments
longitudinally striate; high zonal specimens also bear numerous large pustules or
small spines irregularly developed along the striations. {b) Zooecial bases. Broadly
triangular; irregularly pentagonal on wider branches.
ENGEL: CARBONIFEROUS SEPTATOPORA
595
Material. Holotype NUF2386 (NUL258); paratypes NUF2388a/b, NUF2431a/b,
NUF2445 (NUL258); NUF2523-2525 (NUL39); others NUF2432a/b, NUF2436-
2438, NUF2385, NUF2387, NUF2389a/b, NUF2390-2391 (NUL258); NUF2404,
NUF2424 (NUL414); NUF2488 (NUL448); NUF2522, NUF2526 (NUL39);
NUF2412 (NUL361).
Remarks. The most distinctively different aspect of this species is its regular develop-
ment of nodes. All other species of Septatopora either lack nodes or have them placed
adjacent to some apertures in a generally irregular pattern. In S. nodosa they are
associated with apertures but, because of the narrow branch width, they have assumed
a linear, or slightly zigzag arrangement. In total appearance the species is not a good
representative of the genus to which it has been attached with some misgivings. Were
it not for the apertural septation, this species could be grouped readily with Fenestella
Lonsdale.
The distinctively septate apertures and the proximo-lateral branch connection
with an auxiliary tube form the basis of its assignment here to Septatopora. However,
because of the short vestibular septa, mould infillings of the apertures do not always
appear obviously septate and very close study is needed for correct generic assign-
ment.
As in S. acarinata (Crockford), which shares a similar low zonal range with the
present species, the oldest specimens have a very narrow branch profile which appears
to zigzag between the exserted apertures. Subsequent material assumes a wider branch
profile which eliminates this effect. Coupled with this, there is a reduction in the height
of the low Carina which becomes scarcely apparent, if at all.
A few specimens of S. nodosa have been found attached to the surface of brachiopods
in the Rhipidomella fortimuseula Zone, where they use recumbent basal branches as
an additional attachment device.
Comparisons. S. nodosa is the only species assigned to the new genus which can
be reasonably compared with existing species of Fenestella Lonsdale. Generally,
the distinction is readily based upon the development of apertural septation in
5. nodosa.
Fenestella wilsoni Roberts occurs within a part of the range of S. nodosa and they
share an identical mesh configuration. Because of a crystalline silica coating on all the
type specimens of F. wilsoni it is difficult to observe apertural details, but it does not
appear to be septate, and hence can probably be distinguished on this basis. Should
better-preserved material of F. wilsoni be found, it may be possible to show that these
two species are identical, but for the present they have been retained as separate
taxa.
Other Lower Carboniferous species of Fenestella in Australia are readily dis-
tinguished on mesh grounds alone, quite apart from the apertural details.
Stratigraphy. S. nodosa first appears rarely in the Schellwienella cf. burlingtonensis
Zone at Raglan (NUL448). It is of common occurrence in all subsequent zones up
to and including the Rhipidomella fortimuseula Zone, thus spanning a range com-
parable with much of the Visean interval.
596
PALAEONTOLOGY, VOLUME 18
Septatopora{l) williamsensis sp. nov.
Plate 68, figs. 1-3
Diagnosis. Septatopora-\i\.Q species with straight, narrow, pustulose branches ; mesh
open, medium to fine, with sub-rectangular to rectangular fenestrules; apertures
small, weakly septate, medium spaced being arranged on the extreme margin of the
branch where only the fenestrular rim is weakly exserted; zooecia in two rows per
branch; carina and nodes absent; reverse branch profile rounded; zooecial bases
quadrate to pentagonal.
Description. Zoarium: small, radiating fragments of unknown orientation ; maximum
radius 20 mm. Obverse surface, {a) Branches. Straight to broadly curved, narrow
(m.BW 0-22 mm); narrowly rounded without carina; ornament of fine, pustulose,
longitudinal striations. {b) Dissepiments. Very narrow (m.DW 0-08 mm), centrally
straight with moderate expansion at branch junction; situated well below branch
level; ornament of longitudinal striations. (c) Fenestrules. Sub-rectangular to
rectangular; regular mesh of medium size and open appearance; fenestrules of
medium length, narrow width (m.FL 1-29 mm; m.FW 0-57 mm), {d) Carina. Absent.
(e) Nodes. Absent. (/) Zooecial apertures. Small, circular (m.ZD 0-09 mm); weakly
exserted with indistinct, low peristome developed only on the fenestrule margin of
the aperture; apertures bear radiating septal plates which extend only half-way
towards the axis and which do not extend far down into the vestibule; apertures
situated low on the sides of the branch where they project weakly into the fenestrules
with their low peristomal margin; inner apertural margins depressed into side of
branch resulting in apertures which have a slight proximal and lateral inclination
towards the fenestrule; moderately stabilized with respect to the dissepiments;
zooecia medium spaced (m.Z-Z 0-32 mm) with from three to five zooecia per fenestrule
(m.Z/F 4-0); apertures in two rows with a third row developing in the fork at each
bifurcation.
Reverse surface, {a) Form. Rounded branches joined by narrower level dissepiments;
ornament of fine, pustulose striations, identical to those of the obverse surface.
{b) Zooecial bases. Quadrate to pentagonal in form being arranged in weakly over-
lapping rows.
Material. Holotype NUF2423 (NUL414); paratypes NUF2421-2422 (NUL414).
Remarks. The generic status of this species is somewhat doubtful. On the basis of
its partly septate apertures and distinctive branch ornament it has been placed here
with Septatopora. However, the lack of strongly exserted apertures and the absence
of any auxiliary tube means that it lacks some of the essential diagnostic features of
that genus.
As a relative of Septatopora, it would appear that the recessed apertures and short
radial septa of this species produced only a partial constriction of the vestibule. This
would mean it was still possible to extend both tentacles and lophophore perhaps
making the development of an auxiliary tube unnecessary. The stratigraphic occur-
rence of 5.(?) williamsensis, which correlates approximately with Mid-Visean pre-
cludes any suggestion of an evolving sequence, since other species of Septatopora with
ENGEL: CARBONIFEROUS SEPTATOPORA
597
Strongly developed apertural exsertion and septation are known from late Tournaisian
onwards.
As it is outside the morphological limits of Septatopora, this new species really
deserves its own generic category. However, until much more material of com-
parable form is found, it is not considered appropriate to propose such a taxon.
Another notable morphological aspect of S'.C?) williamsensis is the wide expanse
of central, obverse branch surface which lacks carina, nodes, and apertures. Indeed,
both reverse and obverse surfaces are so similar that careful inspection of the apertural
form is necessary to determine which surface is being examined.
No existing species of Septatopora or Fenestella in the Australian Carboniferous
resemble this distinctive species.
Stratigraphy. S.p.) williamsensis has been found only at the Barrington Guest House
locality (NUL414) where it is associated with S. nodosa, S. acarinata, and S. glou-
cesterensis. On the basis of the whole brachiopod-bryozoan fauna, this horizon has
been placed in the lower parts of the Delepinea aspinosa Zone which correlates
approximately with a Mid-Visean age.
COMPARISON OF SPECIES OE SEPTATOPORA
Descriptive and statistical aspects of all nine species of Septatopora have been given
in Tables 1 and 2.
From a study of information so displayed it is possible to indicate the general
nature of variation present within the genus. Features which show very little varia-
tion include : ( 1 ) A standard zoarial form with only slight changes in fenestrule outline
(oval to sub-rectangular). (2) A lack of any central carina on the branches (excluding
S. nodosa sp. nov.). (3) Development of surface ornament in the form of distinctive
pustulose striations. (4) Apertural septation. (5) Strong apertural exsertion with a high
peristome forming a calice-like depression on each aperture. (6) Common surface
development of hemispherical depressions proximal to some apertures in each
zoarium.
By contrast the following features display considerable variation: (1) Increase in
mesh size. (2) Increase in the number of zooecial rows per branch. (3) Increase in
branch width and changes in branch profile. (4) Irregular nodal development.
(5) Increase in apertural size and spacing. (6) Change in zooecial chamber form and
auxiliary tube. (7) Variations in reverse surface ornament. Each of these aspects is
detailed below with reference to the appropriate species and to the interval over
which they have maximum zonal potential.
1 . Increase in mesh size. Mesh dimensions reveal a progressive increase from small-
to medium-sized species. Low zonal forms {S. acarinata, S. nodosa) are rather
delicate compared with the larger species found in the high zones. Mesh variation is
indicated by the large changes in fenestrule dimensions, shown in Table 2.
2. Zooecial rows per branch. A general trend throughout the Carboniferous in
Australia is for all species of Septatopora to increase the number of zooecial rows per
branch from two to four.
Low zonal species {S. acarinata, S. nodosa) are basically composed of two rows
598
PALAEONTOLOGY, VOLUME 18
TABLE 1 . A descriptive comparison of the important morphological features of the fenestrate mesh of all
species of Septatopora.
Mesh
Branch Width
Form
Prof i le
Fenestrules
Nodes
Apertural
Size &
Spacing
Zooecial Rows
Zooecia
per
Fenestrule
Zooecial
Bases
Branch
Reverse
Post Norm Pre
bif. -al bif.
S. acarinata
fine,
open
narrow;
straight or
zig-zag;
narrowly
rounded
sub-oval to
sub-rectangular;
short ;
narrow
small;
close
2/2/3
3-4
irregularly
pentagonal
rounded
S nodosa
fine;
open
narrow;
straight
to zig-zag;
rounded
sub-rectangular
to rectangular;
short ;
na r row
close;
central
row
small ;
close
2/2/3
3 - 4
broadly
triangular
to
pentagonal
rounded;
spiny
S.l?) williamsensis
medium;
open
narrow;
straight;
narrowly
rounded
sub-rectangular
to rectangular;
medium length;
narrow
-
small;
medium
2/2/3
3 - 5
quadrate to
pentagonal
rounded
S. gloucesterensis
fine;
open
narrow to
medium ;
straight;
rounded
sub-oval to
sub-rectangular;
medium to short
narrow
medium
spacing
small;
close
2 /2-3/4
4 - 5
irregularly
pentagonal
and
rhomboidal
rounded
S.C’l sulcifera
medium;
open
medium to
wi de:
straight;
oval
oval to
sub-rectangular;
medium length;
medium width
small,
close
2-3 / 3 / 3-5
13)5-6
7
rounded
S. stellaris
medium
to fine;
even
medium ;
straight ;
rounded
oval to
sub-oval ;
medium to short;
medium width
distant
spacing
medium;
medium
2 / 2-3 / 3
2 -3
(?) oval
tapered;
spiny
S. Isaacsensis
medium
to fine;
closed
wide ;
straight;
oval
oval to
sub-oval;
medium to short;
medium width
distant
spacing
medium ;
medium
3 / 3 / 4-5
2 -3
oval ;
no overlap
between rows
tapered ;
spiny
S pustulosa
medium;
open
wide,
straight ;
round -oval
sub-oval to
sub- rectangular;
medium length,
medium width
medium;
distant
2 / 3/4-5
3-5
elongate oval
no overlap
between rows
rounded
S Uemingi
medium;
closed
wide to
very wide;
straight,
oval
sub-oval to
sub-rectangular;
medium length;
medium width
“
medium ;
distant
2-3/3-4Z5-6
3-5
elongate oval;
no overlap
between rows
rounded
per branch with additional apertures appearing only at bifurcation. S. gloucesterensis
in the Delepinea aspinosa and Rhipidomella fortimuscula Zones has about equal
development of two and three rows of zooecia between successive bifurcations. In
the Levipustula levis Zone, S. stellaris continues in the trend of S. gloucesterensis but
it is associated with S. pustulosa and S. isaacsensis which are dominantly three-rowed
ENGEL; CARBONIFEROUS SEPTATOPORA
599
TABLE 2. A statistical summary of the principal mesh dimensions of all species of Septatopora. Species
known by only one specimen have only the mean and observed range recorded. Explanation of the abbrevia-
tions are given in the text.
FL
FW
BW
DW
ZD
z-z
N-N
N9
F/10
B/10
Z/5
Z/F
S. acarinata
m
s
OR
mm
0-748
0-123
0-44-1-08
mm
0-484
0-068
0-34-0-64
mm
0-213
0-026
0-16-0-28
mm
0-106
0-037
0-06-0-18
mm
0-078
0-011
0-06-0-10
0-246
0-021.
0-20-0-32
mm
140
13-8
20-7
20-4
3-0
S. nodosa
m
s
OR
0-912
0-150
0-56-1-46
0-504
0-073
0-34-0-70
0-244
O-OU
0-14-0-36
0-092
0-019
0-04-0-14
0-095
0-011
0-06-0-12
0-258
0-02i.
0-20-0-34
0-281
0-040
0-18-0-38
360
11-1
20-0
19-4
3-5
S.(?l williamsensis
m
s
OR
1-289
0-23i
0-86-1-84
0-574
0-079
0-40-0-72
0-223
0-017
0-18-0-26
0-075
O-OU.
0-04-0-10
0-087
0-010
0-08-0-10
0-319
0-020
0-28-0-38
-
60
7-8
17-4
15-7
4-0
S gloucesterensis
m
s
OR
1-116
0-080
0-96-1-28
0-581
0-106
0-44-0-86
0-297
0-030
0-24-0-36
0-183
0-027
0-14-0-26
0-098
0-009
0-08-0-12
0-272
0-019
0-22-0-30
0-512
0-089
0-34-0-64
40
9-0
17-3
18-4
4-1
S.(?) sulcifera
m
OR
1-358
0-76-2-24
0-742
0-70-0-76
0-411
0-35-0-50
0-200
0-14-0-22
0-069
0-04-0-08
0-288
0-24-0-32
-
20
7-4
13-5
17-4
4-7
S. stellaris
m
s
OR
0-953
0-079
0-70-1-10
0-633
0-092
0-50-0-86
0-361
0-053
0-28-0-46
0-140
0-029
0-10-0-24
0-128
0-013
0-10-0-16
0-333
0-031
0-28-0-42
0-951
0-150
0-60-1-22
40
10-5
15-8
15-0
2-9
S. isaacsensis
m
OR
0-914
0-82-1-00
0-747
0-60-0-96
0-470
0-34-0-60
0-263
0-18-0-40
0-120
0-10-0-16
0-324
0-26-0-38
0-937
0-70-1-20
20
10-9
13-4
15-4
2-8
S. pustulosa
m
s
OR
1-684
0-305
1-00-2-60
1-030
0-192
0-60-1-50
0-491
0-092
0-32-0-74
0-212
0-039
0-14-0-36
0-106
0-012
0-08-0-14
0-433
0-055
0-30-0-60
-
200
6-1
9-8
11-6
3-9
S. flemingi
m
s
OR
1-652
0-U.8
1-20-2-12
0-905
0-158
0-60-1-40
0-541
0-097
0-36-0-86
0-300
0-065
0-16-0-46
0-128
0-015
0-10-0-16
0-381
O-Oi.3
0-28-0-50
-
140
6-1
11-0
13-1
4-3
species. Finally, in the Cancrinella levis Zone S. pustulosa is joined by S. flemingi
which is dominantly four-rowed with pre-bifurcation increase up to five (rarely
six) rows.
Both increase in mesh size and number of rows of apertures are universal trends
which affect all Australian Carboniferous fenestrates. Increase is progressive in
both cases and is not readily divided into arbitrary sub-groups.
3. Branch width and profile. Extra rows of zooecia, up to three rows, are accommodated
in increasingly wider branches. Increase to four rows as in S. flemingi is, however,
achieved by change in chamber arrangement which enables the extra row to be con-
tained in branches which have not greatly increased in width.
Increase in branch width is also accompanied by a reduction in relative size of the
fenestrule openings resulting in open-meshed forms changing to a closed-mesh
appearance.
600
PALAEONTOLOGY, VOLUME 18
Variation in branch profile is considerable. As noted previously zigzag profiles
of the low zonal species {S. acarinata, S. nodosa) change to a rounded profile in the
Orthotetes australis Zone beyond which the profile gradually becomes more oval
in form.
4. Nodal development. Some species of Septatopora lack any obverse nodes whereas
others have irregular nodes adjacent to the distal or disto-central rim of some
apertures. There appears to be no stratigraphic control over this feature which has
resulted in three species groups.
(fl) Nodeless: S. acarinata, S'.(?) williamsensis, S.{1) sulcifera, S. pustulosa,
S. flemingi.
{b) Irregular nodes: S. gloucesterensis, S. stellaris, S. isaacsensis.
(c) Regular nodes : S. nodosa.
No major importance above the species level can be given to the position and
occurrence of these nodes. The absence of thin-section detail makes it unlikely that
further functional significance will become known, and without this it is not possible
to assess the weighting which should be given to this morphological feature. Even
within node-bearing species some branches can be found which lack nodal
development.
It is of some interest to note that most Australian species referrable to Polypora
M’Coy also bear similar, randomly distributed nodes which differ in position from
those of Septatopora, being placed adjacent to the proximal or proximo-central rim
of an aperture rather than in the equivalent distal position.
5. Increase in apertural size and spacing. Low zonal species have slightly smaller
apertures (mean range 0-07-0T mm) compared with the medium-sized apertures
(mean range 011-0T3 mm) in the higher zones. In parallel with this variation the
apertures change from closely spaced (mean range 0-25-0-29 mm) to distantly spaced
(mean range 0-33-0-44 mm) over the same stratigraphic interval.
6. Zooecial chambers and associated structures. Chambers in two- or two to three-
rowed species are globular in form and almost entirely fill the thickness of the branch.
This form results in a very short vestibule beginning quite close to the obverse surface
and extending up into the exserted aperture. In this case the auxiliary tube is only
a breach in the side of the vestibule or a very short horizontal connection to the
branch surface on the proximal side of the aperture. As such it is quite difficult to
observe. With zigzag packing of globular chambers, the zooecial base form is irregu-
larly pentagonal with considerable overlap between rows. Central rows of apertures
appear rhomboidal in shape between the lateral rows.
With increase in the number of rows of apertures in the high zonal species
(iS. pustulosa, S. isaacsensis, S. flemingi), the chambers are necessarily packed much
closer. They therefore assume an elongate, compressed-oval form with little or no
overlap between rows. In addition, the ffatter chambers are now located close to the
reverse surface of the branch and this requires considerable changes in the shape of
the vestibule. It is lengthened into an L-shaped or geniculate form with the auxiliary
tube being connected to the short, posterior, horizontal section and extending up to
the obverse surface in parallel with the longer anterior section of the vestibule. As
ENGEL: CARBONIFEROUS SEPTATOPORA
601
two quite separated structures they are now clearly visible on the obverse surface in
their respective positions.
Primarily as a result of changes in branch width, the larger hemispherical depres-
sions can also vary their position. On wide branches in the higher zones the branch
width and zooecial spacing are sufficient to allow full development on the obverse
branch surface. However, in the low zonal species the branch width tends to be very
narrow and the apertures quite close together so that the spherical (?)ovicells are
attached more to the side of the branch where they protrude into the fenestrules, or
they can even continue around on to the reverse side of the zoarium where they may
extend above the reverse surface level.
7. Reverse ornament. All except two species of Septatopora have a broadly rounded
branch profile, joined by narrower dissepiments on the reverse surface. Normal
ornament consists of distinctive pustulose striations similar to that developed on the
obverse surface.
By contrast, two closely related high zonal species, S. stellaris (Campbell) and
S. isaacsensis (Campbell) have a narrow, tapered reverse branch profile. Branches are
joined by equal width, level dissepiments in a distinctive regular or proximally-
polygonal mesh. The branches also bear a near central, very irregular array of
numerous spines which are inclined distally.
Some features of reverse ornament are probably the product of exceptionally good
preservation or are of some ecological significance. For example, all fenestrate genera
from one locality (NUL258, Barrington, N.S.W.) have the same network of very
fine spines or large thorny pustules over their reverse surface. This particular form
of ornament has not been observed at any other locality.
AUSTRALIAN STRATIGRAPHIC DISTRIBUTION OF SEPTATOPORA
From the foregoing discussion, it is apparent that variation in the species of Septa-
topora makes it a most useful local zonal indicator. Text-fig. 2 sets out the range of all
nine species in terms of the brachiopod zones of Campbell and McKellar (1969),
Roberts and Oversby (1972), and Jones, Campbell and Roberts (1973) with which
there is a strong parallel.
Tournaisian-earliest Visean
Tulcumbella tenuistriata and Spirifer sol Zones have not yielded species referrable
to the new genus. However, outcrops are few, and as other fenestrates are known from
this level, it is quite possible that representatives will be found.
Schellwienella cf. burlingtonensis and Pustula gracilis Zones contain the first repre-
sentatives of the new genus, namely Septatopora acarinata and S. nodosa. At this
stratigraphic level, examples of both species have very narrow branches which zigzag
between alternating apertures in a most distinctive pattern.
Visean
Orthotetes australis Zone contains the same two species, but at this higher level both
have developed wider, straight branches in which the apertures no longer are the
dominating element.
602
PALAEONTOLOGY, VOLUME 18
EASTERN AUSTRALIAN
FAUNAL ZONES
STRATIGRAPHIC DISTRIBUTION
OF SEPTATOPORA
EUROPEAN
STAGES /ZONES
Cancrinella levis
of
c
r S
to ^
c
STEPHANIAN
Levipustula levis
O
to
o
« 5
51 a
QiU
\ "
Vll
Sif i/j
ll
t/)
WESTPHALIAN
NAMURIAN
Oriocrassatella compressa
to
Marginirugus barringtonensis
-a s
10 -Si
Cu IIip-8
CunioA
VISEAN
Rhipidomella fortimuscula
ifera
amser
ouces
Delepinea aspinosa
ILfl 3
Orthotetes australis
D
o
c
Q
nodosa
t4 uo
Cull S
]
[DI
r
Pus tula gracilis
O
O
t^
cuim
Schellwienella cf.
burlingtonensis
t/i
CuIIoc
TOURNAISIAN
Spirifer sol
Tulcumbella tenuistriato
Cl
I
TEXT-FIG. 2. Stratigraphic distribution of the various species of Septaiopora in terms of the Eastern Australian
Carboniferous zones. Tentative correlation with equivalent European zonation is also included.
Delepinea aspinosa Zone contains representatives of the two previous species plus
5.(?) williamsensis, and the first of the two- to three-rowed forms S. gloucesterensis
and 5.(?) sulcifera.
Rhipidomella fortimuscula Zone contains S. gloucesterensis and S. nodosa with all
other prior species having disappeared.
Late Visean-Narnurian-earliest Westphalian
Marginirugus barringtonensis Zone is dominated lithologically by coarse detrital
sedimentary units within which no bryozoan remains have been preserved. Most
brachiopods occur as reworked detrital shell deposits indicating a medium quite
unsuited to delicate bryozoan preservation. The same observations are true for the
Oriocrassatella compressa Zone (new) where large banks of heavy bivalve shells
dominate the fossil record.
Westphalian -Stephanian
Levipustula levis Zone includes no species of Septatopora from lower stratigraphic
ENGEL: CARBONIFEROUS SEPTATOPORA
603
levels, although some generalized species of Fenestella (e.g. F. osbornei Crockford)
do appear to have continued across the coarse sedimentary interval noted above.
The fauna in this zone consists of S. stellaris (two to three rows), S. pustulosa (three
rows), and S. isaacsensis (three rows) of which the three-rowed species are by far the
most abundant forms.
Cancrinella levis Zone (new) includes the final species S.flemingi (three to four rows)
together with S. pustulosa. The new species S.flemingi is the most common bryozoan
at this level in a zone which is known only from regions of Carboniferous outcrop in
east-central Queensland. The type locality for this newly defined zone is placed in the
Stanwell-Ridgelands district where Fleming (1969) has published a geological map
together with a description of some of the other elements of this late Carboniferous
fauna.
It appears that Septatopora did not continue into overlying strata, for a literature
survey of Australian Permian species referred to Fenestella Lonsdale and Polypora
M’Coy has failed to reveal any probable members of the new genus. It is possible,
however, that apertural septation has not been observed because of the generally
poor state of preservation, in very coarse sandstones, of many Permian species. It
should be noted also that the degree of apertural exsertion is slightly reduced in the
final Carboniferous species, indicating that this may not be an obvious aspect in any
Permian successors. Finally, it is of some value to indicate that most Australian
Permian species of Polypora M’Coy have five or more rows of apertures, thus con-
tinuing the general trend noted in all Carboniferous fenestrates.
Correlation of the above assemblage zones with the European succession is diffi-
cult. Controls used by the previously mentioned authors in the definition of their
Eastern Australian assemblage zones depend not upon the nominate, long-ranging
brachiopod species, but upon the associated goniatites, a rare and often confusing
component of the local faunas.
Initial correlation (Campbell and McKellar 1969) assumed that the standard
German goniatite sequence was continuous. With the discovery by various European
conodont workers including Rhodes et al. (1969, 1971) and Matthews (1969a, b,
1970a, b) that the Lower Carboniferous European goniatite sequence contains
several major time gaps, the need arose to recorrelate the Eastern Australian zones.
A definitive statement of the revised position is given by Jones et al. (1973) who give
detailed arguments for their currently adopted position.
Assessment of the magnitude of the time breaks in the European sequence continues
to present difficulties, making even the most recent efforts only very tentative.
Conodont studies presently being undertaken by Dr. H. Jenkins and colleagues
(University of Sydney) would suggest that the correlations adopted in text-fig. 2
must be revised so that the Rhipidomella fortimuscula Zone is of an early rather than
late Visean age, with the position of the Tournaisian-Visean boundary remaining
unaltered within the Schellwienellaci. bur lingtonensis Zone. Changes of this magnitude
in the Lower Carboniferous will obviously also vary the Upper Carboniferous
correlations and hence it is not possible to offer confident comparisons at this
stage. A study of the brachiopod content of the Cancrinella levis Zone reveals,
however, a fauna of distinctively Permian affinities and it is with some confidence
L
604
PALAEONTOLOGY, VOLUME 18
that the range of Septatopora is extended up to the level of the late Westphalian or
Stephanian age.
It is therefore concluded on present evidence that Septatopora ranges from late
Tournaisian to Stephanian, at which point it appears to have become extinct.
GEOGRAPHICAL DISTRIBUTION
In Australia the pre-Carboniferous record is generally poor in fenestrates, so that no
suitable ancestral material has yet been described.
With the possible exception of Polypora(l) verrucosa (M’Coy) no other described
fenestrate has been observed in the literature, at present available to the author,
which could be placed in Septatopora.
The only other record of the genus outside Australia would appear to be in the
Upper Carboniferous of Argentina, where Amos and Sabattini (1969) and Mrs. N.
Sabattini (pers. comm.) have listed the occurrence of several fenestrate species,
originally described by Campbell (1961), some of which have been transferred here
to the genus Septatopora.
Dependent upon further published descriptive details by Mrs. N. Sabattini, it
would appear that Septatopora is another of the many genera shared in common by
these two continents during Upper Carboniferous times.
Acknowledgements. I am indebted to Professor B. Nashar of the University of Newcastle and Dr. K. S. W.
Campbell of the Australian National University for their continued guidance and support during the
course of this project. Thanks are also due to Emeritus Professor D. Hill, University of Queensland,
Mr. G. Fleming, Queensland Geological Survey, Mr. G. Foldvary, University of Sydney, Dr. B. Runnegar
and Mr. G. Brown, University of New England, and Dr. A. Ritchie, Australian Museum, for their generous
provision of access to, or loan of, type material from their respective collections.
The writer is especially grateful for the constructive comments given freely by Dr. R. S. Boardman,
Smithsonian Institution, Emeritus Professor D. Hill, University of Queensland, and Dr. P. L. Cook and
Dr. B. R. Rosen of the British Museum (Natural History) upon the problems of classification and func-
tional morphology of the new genus. Provisional assignment to the Phylum Bryozoa, however, remains the
responsibility of the author and does not necessarily reflect the opinion of these distinguished contributors.
Finally, thanks are recorded for the support given by Mrs. N. Morris and a number of honours students
at the University of Newcastle, whose field studies have allowed me to gather together a very large quantity
of bryozoan material in the University of Newcastle collections.
REFERENCES
AMOS, A. j. and sabattini, n. 1969. Upper Palaeozoic faunal similitude between Argentina and Australia.
Gondwana Stratigraphy, I.U.G.S. Symposium in Buenos Aires, 1-15 Oct. 1967, UNESCO Earth Sciences
Publication, 2, 235-248, 1 table.
BASSLER, R. s. 1953. In MOORE, R. c. (ed.). Treatise on invertebrate paleontology. Part G, Bryozoa. Geol. Soc.
Amer. and University of Kansas Press. G1-G253.
CAMPBELL, K. s. w. 1961. Carboniferous fossils from the Kuttung rocks of New South Wales. Palaeontology,
4, 428-474, pis. 53-63.
— 1962. Marine fossils from the Carboniferous glacial rocks of New South Wales. J. Paleont. 36, 38-52,
pis. 11-13, 4 figs.
- - and MCKELLAR, R. G. 1969. Eastern Australian Carboniferous invertebrates: sequence and affinities.
In CAMPBELL, K. s. w. (ed.). Stratigraphy and Palaeontology. Aust. Nat. University Press, Canberra,
77-119.
ENGEL: CARBONIFEROUS SEPTATOPORA
605
CROCKFORD, J. M. 1947. Bryozoa from the Lower Carboniferous of New South Wales and Queensland.
Proc. Linn. Soc. N.S.W. 72, 1-48, pis. 1-6.
1949. Bryozoa from the Upper Carboniferous of Queensland and New South Wales. Ibid. 73, 419-
429, 4 figs.
CVANCARA, A. M. 1958. Invertebrate fossils from the Lower Carboniferous of New South Wales. J. Paleont.
32, 846-887, pis. 109-113.
DRISCOLL, E. G. 1960. Geology of the Mundubbera District. Pap. Dept. Geol. Univ. Qd, 5, 5-27, figs. 1-3.
FLEMING, p. J. G. 1969. Fossils from the Neerkol Formation of Central Queensland. In Campbell, k. s. w.
(ed.). Stratigraphy and Palaeontology. Aust. Nat. University Press, Canberra, 264-275, pis. 16-17.
1972. Redescription of fenestellid species from the Upper Carboniferous of Eastern Australia. Pubis.
geol. Surv. Qd, 354, Palaeont. Pap. 29, 1-8, pis. 1-4.
HILL, D. 1934. The Lower Carboniferous corals of Australia. Proc. R. Soc. Qd, 45, 63-1 15.
JONES, p. J., CAMPBELL, K. s. w. and ROBERTS, J. 1973. Correlation chart for the Carboniferous System of
Australia. Aust. Bur. Miner. Resour. Rec. 69, 1-80.
JULL, R. K. 1968. The Lower Carboniferous limestones in the Monto-Old Cannindah district; a brief
description and a proposed name. Qd Gov. Min. J. 69, 199-201.
1969. The Lower Carboniferous corals of Eastern Australia, a review. In Campbell, k. s. w. (ed.).
Stratigraphy and Palaeontology. Aust. Nat. University Press, Canberra, 120-139, pis. 9-10.
MCKELLAR, R. G. 1967. The Geology of the Cannindah Creek area, Monto District, Queensland. Pubis,
geol. Surv. Qd, 331, 1-38, figs. 1-8.
MATTHEWS, s. c. 1969a. A Lower Carboniferous conodont fauna from East Cornwall. Palaeontology, 12,
262-275, pis. 46-50.
19696. Two conodont faunas from the Lower Carboniferous of Chudleigh, South Devon. Ibid.
276-280, pi. 51.
1970a. A new cephalopod fauna from the Lower Carboniferous of East Cornwall. Ibid. 13, 112-131.
19706. Comments on palaeontological standards for the Dinantian. Congr. Avanc. Etud. Stratigr.
Garb., 6th, Sheffield, 1967, 3, 1159-1163.
MAXWELL, w. G. H. 1964. The geology of the Yarrol Region, Part 1, Biostratigraphy. Pap. Dept. Geol. Univ.
Qd. 5, 1-65, pis. 1-14.
MILLER, T. G. 1963. The bryozoan genus Polypora M’Coy. Palaeontology, 6, 166-171, pis. 23-24.
RHODES, F. H. T. and AUSTIN, R. L. 1971. Carboniferous conodont faunas of Europe. Mem. geol. Soc. Amer.
Ml, 317-352.
and DRUCE, e. c. 1969. British Avonian (Carboniferous) conodont faunas and their value in
local and intercontinental correlation. Bull. Br. Mus. nat. Hist. Geol. Supp. 5, 1-313, pis. 1-31.
ROBERTS, J. and oversby, b. s. 1972. The Lower Carboniferous geology of the Rouchel District, New South
Wales. Aust. Bur. Miner. Resour. Rec. 119, 1-59, pis. 1-19, figs. 1-16.
TAVENER-SMiTH, R. 1969. Skeletal structure and growth in the Fenestellidae (Bryozoa). Palaeontology, 12,
281-309, pis. 52-56.
and williams, a. 1972. The secretion and structure of the skeleton of living and fossil bryozoa. Philos.
Trans. R. Soc. Lond. 264 (859), 97-159, pis. 1-32.
WASS, R. E. 1968. Permian polyzoa from the Bowen Basin. Bull. Aust. Bur. Miner. Resour. 90, 1-135, pis. 1-18.
BRIAN A. ENGEL
Department of Geology
University of Newcastle
Typescript received 20 May 1974 Newcastle, N.S.W. 2308
Revised typescript received 10 October 1974 Australia
Addendum : The author has very recently been shown fenestrate specimens from the Middle Permian of the
Bowen Basin, Queensland, which have strongly exserted apertures bearing eight apertural septa. This
discovery thus extends the probable range of Septatopora gen. nov. into Permian strata in Eastern Australia.
As an oversight during manuscript preparation, reference has not been made to Polypora stenostoma
Tavener-Smith (1971) which shares some aspects of morphological similarity with the new genus.
TAVENER-SMITH, R. 1971. Polypora stenostoma: a Carboniferous bryozoan with cheilostomatous features.
Palaeontology, 14, 178-187, pi. 25.
1.. •'4^
iW
■ . :. *«.
■ 'f r • J5i“* '■ ' -
f. -' .,--v =-’' ^'' .
■?. -
'*^'4
r^r..
>s V
• :,. ■ ■'■'
' , -■■■ -■ 'ui
. , i*. ^ , ' ('‘i.yj,'' I*. , (
i , i • V.. 'V ,^ •
-. ♦" . .^''.4* '
N.,
K- K«*.H -.• ■
■' ■ .f "rtHtWf* ’■
. o^-
■ ‘f.-v '
• ^
• ■.-, ; •.•. ..>,» — ■ '•, '’ .
- •^d) to typical rhodocrinitid types (text-
fig. 3c) where the basals are large, in lateral contact, and support the radials and inter-
brachials. Transitional crinoids are also known. Note that small basals (relative to
size of the infrabasals and radials) are associated with the zygodiplobathrid condition
and large basals with the normal rhodocrinitid or eudiplobathrid base. Ubaghs (1950,
p. 120) notes that two of the radials of Paulocrinus biturbinatus Springer (1926/?,
p. 21, pi. 4, figs. 5, 5a-c) lie on truncated infrabasals whereas the other three rays
have the normal rhodocrinitid condition. As in Rhipidocrinus crenatus, the relatively
large infrabasals and small basals are associated with the zygodiplobathrid type base.
These examples clearly support Ubaghs (1950) and Breimer (1960) in their contention
that zygodiplobathrids and normal dicyclic camerates are both variants of the same
basic organization. It also implies that one was probably ancestral to the other.
These variations within rhodocrinitids are believed to indicate genetic instability
with respect to this character, at least in certain species. On the other hand, the
proximal structure of the cup seems to be stabilized in zygodiplobathrid taxa. Although
these crinoids are rare and do not seem to have been successful, moderately large
642
PALAEONTOLOGY, VOLUME 18
TEXT-FIG. 3. Origin of Dimerocrinites pentlandicus sp. nov. a, lateral view of holotype ; note shape of primi-
brachs, structure of interbrachials, and zygodiplobathrid base, x3-4. b-d, sketches showing probable
ontogeny of rhodocrinitids with eudiplobathrid base and zygodiplobathrids. b, ‘larva’ of dicyclic camerate
crinoid with interbrachials separating the radials; note that the plates are still separated by tissues; during
later ontogeny, these join to form a rigid mosaic of plates; reconstructed by analogy with living crinoids,
based on the growth sequence of plates and statistical data of Brower (1973, pp. 292-328); height of larval
calyx is about 0-5 mm. c, D, bases of mature crinoids simulated by growth of the ‘larva’, c, eudiplobathrid
base of rhodocrinitid, growth rates of height and width of the plates relative to total size of the calyx:
infrabasals 014, basals 0-43, radials 0-43. d, zygodiplobathrid base, relative growth rates of height and width
of the plates: infrabasals 0-62, basals 013, radials 0-25. e-g, sketches showing postulated ontogeny of
zygodiplobathrid base seen in D. pentlandicus sp. nov. and normal dimerocrinitids. e, ‘larva’ of dicyclic
camerate crinoid in which the proximal interbrachials are at the level of the primibrachs, reconstructed as
before, f, g, bases of mature dimerocrinitids simulated by growth of the ‘larva’, f, eudiplobathrid base of
normal dimerocrinitid, relative growth rates for plates: infrabasals 014, basals 0-43, radials 0-43.
G, zygodiplobathrid base of D. pentlandicus sp. nov., relative growth rates for plates: infrabasals 0-62,
basals 013, radials 0-25. All relative growth rates of plates are approximate averages for both height and
width. Symbols: radials black, interbrachials stippled.
BROWER; SILURIAN CRINOIDS FROM SCOTLAND
643
samples are available in Cleiocrinus laevis Springer (191 1, p. 44). Examination of five
to ten calyx bases discloses no variation.
The oldest zygodiplobathrid is the bizarre Cleiocrinus from middle Ordovician
rocks of North America (Springer 1905; Ubaghs 1950, pp. 116-120; 1953, p. 691).
In Cleiocrinus, the combined radial-basal circlet overlaps and hangs down over the
infrabasals. Except for the CD interray, there are no interbrachials and the adjacent
ray plates join one another above the radials and basals. The calyx is large because of
the numerous fixed-brachials and all calyx plates are pierced by complex sutural pores
which are interpreted as respiratory devices connected to body coeloms. The free
arms are composed of uniserial pinnulate brachials. The evolutionary history of
Cleiocrinus is unknown because there are no known connecting links between the
genus and any known Ordovician archaeocrinids or rhodocrinitids. Nevertheless,
Cleiocrinus is believed to have been derived from an archaeocrinid or rhodocrinitid
stock.
The Devonian zygodiplobathrid Spyridiocrinus is less obscure (see detailed dis-
cussion in Ubaghs 1950). Aside from the large basal concavity and a reduced number
of interbrachials, intersecundibrachs, etc., the over-all crown habit of Spyridiocrinus
is ‘typical rhodocrinitid’. Comparison of Cleiocrinus and Spyridiocrinus shows only
one similarity— the zygodiplobathrid base. Consequently the two crinoids are not
considered closely related. Obviously, the bizarre Cleiocrinus was not ancestral to
Spyridiocrinus. The large time gap between the two taxa is certainly consistent with
this belief. The total morphological similarities between Spyridiocrinus and Silurian-
Devonian Rhodocrinitidae seem much greater than those between Spyridiocrinus and
Cleiocrinus. The most similar rhodocrinitid genera are : Anthemocrinus Wachsmuth
and Springer (1881, p. 208 (382)), Wenlock. Paulocrinus Springer (1926^, p. 22),
middle or upper Silurian. Condylocrinus Eichwald (1860, p. 612), Devonian. Pre-
sumably the ancestral stock of Spyridiocrinus is within one of these genera. At any
rate, rhodocrinitid ancestry is postulated for Spyridiocrinus with little doubt. If these
considerations are correct, then the two zygodiplobathrid genera had independent
evolutionary histories regardless of the origin of Cleiocrinus. Thus zygodiplobathrids
were probably polyphyletic. If the ancestry of Cleiocrinus can be clarified, it seems
advisable to drop the suborder Zygodiplobathrina and group the two genera within
the Eudiplobathrina along with the most closely related families.
As previously mentioned, the zygodiplobathrid base is associated with relatively
small basals and large infrabasals whereas the reverse characterizes eudiplobathrids.
This suggests that one type can be derived from the other by means of ‘mutations’
which affected the growth of the youngest crinoids; these ‘mutations’ would increase
or decrease the growth rates of the height and width of the basals relative to those of
the surrounding plates. [The word ‘mutations’ is used in a highly general sense, namely
to include gene changes, chromosome additions, translocations, etc. ; a detailed dis-
cussion of this concept is given by Brower (1973, p. 328 et seq.).] In order to test
this hypothesis, a series of hypothetical crinoids was drawn based on statistical data
(text-fig. 3b-g). A simulated rhodocrinitid larva is pictured in text-fig. 3b; this was
reconstructed based on analogy with living crinoids, using the statistical data and
plate-growth sequences for camerate crinoids (Brower 1973, pp. 292-328). The
subsequent growth of this ‘larva’ was simulated in two ways. In Case I the growth
■ 644
PALAEONTOLOGY, VOLUME 18
rates of the basals are large compared to those of the infrabasals and radials. In
Case II the basals are characterized by relatively small rates of growth. Note that the
growth rates are only approximate averages for height and width and that the growth
rates are listed as proportions. The data are ;
Case Approximate values of subsequent growth Type of base
rates for height and width of the listed plates produced
relative to total height of the larval calyx
Infrabasals
Basals
Radials
014
0-43
0-43
Rhodocrinitid-
eudiplobathrid
(text-fig. 3c)
0-62
013
0-25
Zygodiplobathrid
(text-fig. ’id)
Text-fig. l>b-d indicates that the simulation is geometrically feasible and thus
zygodiplobathrids can be derived from eudiplobathrids by a decrease in the growth
rates of the basals relative to those of the surrounding plates.
Comparison. Assignment to the Dimerocrinitidae seems probable because the
Pentland crinoid is basically a dicyclic camerate crinoid with many fixed-brachials
and radials in contact within the lateral interrays. For reasons discussed above, the
presence of a zygodiplobathrid base is not thought to be a fundamental character.
Considering the affinities of the Pentland species within the Dimerocrinitidae, the
most closely allied crinoids should have; (1) four arms per ray with axillary secundi-
brach 2; (2) primitive structure of the fixed-brachials. In D. pentlandicus primibrach 1
has eight sides and the primaxil eight or nine sides. The most closely related dimero-
crinitid should exhibit a similar structure, probably consisting of hexagonal primi-
brach 1 and septagonal primaxil. (3) Slender calyx. (4) Relatively high infrabasals.
Only a few Ordovician and Silurian dimerocrinitids fit these specifications. These
include Ptychocrinus parvus (Flail) (Wachsmuth and Springer 1897, p. 199) and
several species of Dimerocrinites with four arms in each ray; i.e. specimens from the
Wenlock of Gotland and Dudley, England, labelled D. quinquangularis (Angelin),
D. ornatus (Angelin), and D. speciosus (Angelin). These crinoids were described by
Angelin (1878), but unfortunately the original figures are not reliable because many
are probably composites of several crinoids which are not always conspecific; hence,
emphasis is placed on specimens seen by the writer. These crinoids are easily separated
from D. pentlandicus by the shapes of the primibrachs, the nature of the adjacent
interbrachials, and the type of cup base present. The Pentland crinoid has primibrachs
with eight or nine sides whereas those of the other taxa bear from four to nine sides.
Due to the structure of the primibrachs, D. pentlandicus has four ranges of inter-
brachials below the proximal secundibrachs but only one, two, or three ranges are
found in the other forms. As mentioned above, the Pentland species has a
zygodiplobathrid-type base whereas the other crinoids have eudiplobathrid bases, and
Ptychocrinus parvus has more prominent median-ray ridges than in D. pentlandicus.
Phylogeny. The most likely ancestor for D. pentlandicus is an Ordovician or lower
Llandovery dimerocrinitid or ptychocrinid with four arms in each ray. Two main
evolutionary changes were involved. First is the development of a zygodiplobathrid
BROWER: SILURIAN CRINOIDS FROM SCOTLAND
645
from a eudiplobathrid base. This probably occurred in roughly the same way that
Spyridiocrinus was derived from the Rhodocrinitidae, by means of a growth ‘mutation’
which reduced the growth rates of height and width of the basals relative to those of
the sub- and superjacent infrabasals and radials. Variation of this sort is unknown
within the Dimerocrinitidae, but precedent for such evolution is shown by variation
within various rhodocrinitids such as Rhipidocrinus crenatus and the Carboniferous
inadunate Woodocrinus gravis Wright (1950-1954, pi. 25, cf. figs. 2, 3, 5, 6, 9). Where
the basals are small, they are not in lateral contact and the radials rest on the truncated
infrabasals. If the basals are large, they are in lateral contact and the radials are fully
separated from the infrabasals. Small basals are rectangular like those of
D. pentlandicus but the large basals are hexagonal like those of normal dimero-
crinitids. The simulated crinoids in text-fig. 3e-g show that such a change is at least
geometrically plausible.
The second change is the divergence in the structure of the primibrachs and inter-
brachials. Increase in the number of sides of the primibrachs probably began when
the plates formed early during ontogeny. The shape changes were achieved through
adjustments of the various growth rates of widths of the primibrachs (see Brower
1973, pp. 401-407 for outline of similar evolution in patelliocrinids). An increase in
the supply rate of interbrachials in conjunction with the shape changes of the primi-
brachs mentioned above would be sufficient to develop the interbrachial areas of
D. pentlandicus from the ancestral type.
Subclass INADUNATA Wachsmuth and Springer, 1885
Order disparida Moore and Laudon, 1943
Superfamily homocrinicae Ubaghs, 1953
Family pisocrinidae Angelin, 1878
PISOCRINUS de Koninck, 1858
Type species. P. pilula de Koninck, 1858.
Pisocrinus campana Miller
Plate 74, figs. 1, 2, 4; text-fig. 4
1891 Pisocrinus campana Miller, p. 32, pi. 11, figs. 4, 5.
1892 Pisocrinus campana Miller, p. 642, pi. 11, figs. 4, 5.
1897 Pisocrinus sp., Wachsmuth and Springer, pi. 8, fig. 10.
1915 Pisocrinus campana Miller; Bassler, p. 980.
1926fi Pisocrinus campana Miller; Springer, p. 76, pi. 24, figs. 6-27.
1943 Pisocrinus campana Miller; Bassler and Moodey, p. 612.
1952 Pisocrinus cf. campana Miller; Lament, p. 29.
Scottish material. A crown and a cup (RSM 1970.42.3, 4) occur on a small slab. Unfortunately, the original
has been lost and the two crinoids are only represented by latex casts. Two other crowns, RSM 1970.42.2,
1970.43. Part and counterpart of a cup, Hunterian Museum (HM) 3173a, b.
Type locality. Upper Llandovery or Wenlock; Salamonie Dolomite; Wabash, Indiana, U.S.A.
Other American localities. Upper Llandovery; Osgood Formation; St. Paul and adjacent areas in southern
Indiana. Lower Wenlock; Laurel Limestone; St. Paul, Indiana. Lower Ludlow; Brownsport Formation;
various localities in Wayne, Perry, and Decatur Counties, Tennessee.
646 PALAEONTOLOGY, VOLUME 18
Scottish locality. Plectodonta Mudstones, lower part of River North Esk, about 220 yd north-east of the
North Esk Reservoir.
Diagnosis. A species of Pisocrinus with moderately high cup which shows wide varia-
tions in shape, height/width ranges from 0-75 to 1-2; walls of cup slightly rounded;
basals high relative to radials regardless of cup shape; radial processes weakly
developed ; plates of cup smooth. Arms long and slender, arm length/height of cup
ranges from 4 0 to 8 0, arms of mature crinoid consist of about five brachials. Dorsal
sides of brachials rounded or distinctly triangular. Stem round, composed of only
one order of plates; distal columnals nodose.
Description of Scottish specimens. Cup moderately high; height/width ranges from 0-85 to 10; sides of cup
slightly rounded; basals high relative to cup height; height of basals/cup height equals about 0-41 ; surfaces
of cup smooth, may be faintly rugose in one specimen. Basals five, three pentagonal and two rectangular
(A ray, BC interray); pentagonal basals larger than rectangular ones, height/width of pentagonal basal is
0-8 to 1 -0 ; same for rectangular basal equals 0-5. Large inferradial occurs under B and C ray radials, septa-
gonal, height/width about 10. B and C ray superradials basically pentagonal, height/width 0-7. A and D
ray radials largest plates in cup, pentagonal to septagonal, height/width ranges from 0-8 to 1 0. E ray radial
not seen. Radial facets wide, about two-thirds width of radials, faintly curved with small radial processes.
Tegmen unknown. Arms large, massive, blade-like, consist of uniserial, non-pinnulate brachials; each
arm has from three to eight brachials ; most of arm tapers gently ; at distal arm tips, the taper angle increases
and the last brachial is bullet-shaped; arm length/cup height varies from 4 0 to 5-5. Primibrach 1 small,
rectangular, much wider than high, partially set inside of radial processes; higher primibrachs much larger
and more massive, sutures obscure; height/width variable, ranges from 10 to 16; dorsal sides of brachials
are more or less strongly triangular. Large part of column observed, round with small round axial canal;
entire column consists of only one order of columnals; column tapers distally from below calyx to mid-
distal region of stem; distal-most stem plates become wider than those of mid-distal region. Proximal
columnals wide relative to height, shaped like thin discs with slightly nodose edges.
Comparison. Despite the recent monographic treatment of pisocrinids by Bouska
(1956), the American species remain in need of revision. For example, P. campana
Miller (see Springer 1926Zt, p. 76) and P. benedicti Miller (1891, p. 29; Springer
1926/t, p. 77) commonly occur together; the former is separated by a cup with
straight or slightly curved walls which is high relative to width whereas the latter is
lower and more globose. Springer noted intergradations between the two species
(1926Z), p. 76): 'As stated, with the expanding and bell shaped forms the identifica-
tion is easy, but those with a lower calyx, ovoid to globose, are confusing; if they have
EXPLANATION OF PLATE 74
Figs. 1, 2, 4. Pisocrinus campana Miller, note blade-like arms, relatively high cup with straight walls and
high basals, figured specimens, Plectodonta Mudstones, lower part of River North Esk, about 220 yd
north-east of the North Esk Reservoir. 1, crown with rugose markings on plates of cup and relatively
short arms, /I, B, and Cray view of RSM 1970.42.1, x6-4. 2, poorly preserved crown with tumid plates
and cigar-like arms, lateral view of RSM 1970.43, x7 0. 4, left, relatively wide and globose cup with
smooth plates, A and B ray view of RSM 1970.42.3; right, crown with comparatively high cup with
smooth plates and long arms, lateral view of RSM 1970.42.4, x 5-3.
Fig. 3. Ptychocrinus longibrachialis sp. nov., note long slender arms composed of elongate brachials,
C ray view of holotype. Grant Institute of Geology 134, mudstone layer, just above top plantation on
Gutterford Burn, x2-8.
Fig. 5. Dimerocrinites pentlandicus sp. nov., note small basals and zygodiplobathrid-type base of cup,
shape of primibrachs and structure of interbrachials, lateral view of holotype RSM 1885.26.78h, ‘Starfish
Bed’, Gutterford Burn, x3-8.
PLATE 74
BROWER, Scottish Silurian crinoids
648
PALAEONTOLOGY, VOLUME
TEXT-FIG. 4. Figured specimens of Pisoaimis campana. Note slender calyx with
high basals and long blade-like arms, a, lateral view of crown, RSM 1970.42.4,
X 5-3. B, A and B ray view of cup with a partial arm, RSM 1970.42.3, x 5-3.
c, A, B, and C ray view of partially disarticulated crown, RSM 1970.42.1,
X 5 0. D, lateral view of RSM 1970.43, x 5-3, the plate structure of the cup
is conjectural. Symbols: radials and superradials black, inferradials are ruled
horizontally.
fairly high basals, we may call them campana, while those with basals but little visible
will have to go into benedicti. Thus there will be an intermediate zone in which the
distinction is shadowy.’ This is shown by measurements made on the specimens in
the Springer Collection, United States National Museum, in which the height/width
ratios of the cup of specimens assigned to P. campana by Springer ranges from 0-75
to 1 -2 while that of P. benedicti varies from 0-60 to 1 -2. The figures overlap and further
data are required to either recombine or fully define the two species. Pending restudy,
the specimens with the higher cups having high basals and straight or nearly straight
walls are assigned to P. campana while those with lower and more globose cups with
shorter basals are placed in P. benedicti. Thus P. campana is a highly variable species
which ranges from the upper Llandovery to the lower part of the Ludlow.
The specimens from the Pentland Hills differ from typical North American
individuals in several respects. Most American crinoids have well-developed radial
processes although these grade into individuals with faint radial processes (Springer
\92bb, pi. 24). The Scottish specimens are characterized by shallow radial processes
like some of the end-member crinoids from America. The Pentland specimens range
much smaller. The largest crown is about 13 mm high whereas Springer (1926/?,
pi. 24, figs. 7, 8) illustrated crowns about 65 mm high. Both the Pentland and the
BROWER. SILURIAN CRINOIDS FROM SCOTLAND
649
American crinoids possess comparable numbers of brachials in each arm. The fact
that crinoids develop new brachials at the arm tips throughout life indicates the
Pentland crowns were probably not juveniles. These are believed to be adults which
exhibited reduced growth rates of size with respect to time compared with typical
American individuals. Nevertheless, the high cup with slightly rounded walls and
large basals in conjunction with the long blade-like arms and nodose columnals in the
stem indicate that the Scottish crinoids should be referred to P. campana.
P. campana is also closely related to P. pilula de Koninck (see Bather 1893, p. 27 ;
Bouska 1956, pp. 13, 62, 104) of the Wenlock and Ludlow of England, Gotland, and
Bohemia. Both species are long ranging and widespread and both vary in the height/
width ratio of the cup and the nature of the radial processes. P. campana can be
separated from P. pilula by several characteristics. Better-developed radial processes
are seen in P. pilula. The distal columnals of P. pilula exhibit smooth sides whereas
the columnals of P. campana are nodose. Also the proximal stem of P. pilula appears
to have been secondarily thickened, a feature which is not known in P. campana.
The basals of P. pilula are always low while those of P. campana are much higher rela-
tive to the height of the cup. The cup walls of P. campana are commonly rounded but
those of P. pilula are generally straight. P.pocillum Angelin (1878, p. 21 ; Bather 1893,
p. 33; Springer 19266, p. 80) from the Silurian of Gotland, P. ubaglisi Bouska 1956
and P. mofinensis Bouska 1956 both from the Ludlow of Bohemia, have higher cups
(height/width ranges from 1-3 to 1-7) with straight and angular sides in contrast to
the lower cup with straight or slightly rounded walls of P. campana.
Superfamily iocrinicae Ubaghs, 1953
Family myelodactylidae Miller, 1883
HERPETOCRiNUS Salter, 1873
Type species. H. ftetcheri SaAiQX 1873.
Herpetocrinus parvispinifer sp. nov.
Plate 73, figs. 3, 5
Holotype. RSM 1897.32.285, a terminal coil with part of the straight portion of the stem, in which the crown
is only represented by the distal parts of the arms. The lack of the cup and most of the crown does not
preclude definite generic and specific placement because all Myelodactylidae can be classified on stems
alone: the crowns are only known in three of the five genera assigned to the family by Moore (1962,
pp. 40-44).
Paratypes. Straight stem segments with cirri; RSM 1885.26.78e (part and counterpart), 1897.32.286 (part
and counterpart). A partial terminal coil and straight stem segment: RSM 1897.32.287 (part and counter-
part). A partial terminal coil: RSM 1897.32.288 (part and counterpart).
Derivation of name. From the short spines on the distal margin of each cirral.
Type locality. Gutterford Burn Flagstones, ‘Starfish Bed’, Gutterford Burn.
Diagnosis. A species of Herpetocrinus with characteristic cirral ornamentation;
distal end of cirral expanding outward to form small angular rim-like process;
typically the rim bears two to six short spines; rarely, the rim is absent or weakly
developed but such cirrals always show traces of spines. Column with crescent-
shaped cross-section, concave part of crescent faces the outside of the coil of the stem,
convex side of crescent located on the inside of the coil.
650
PALAEONTOLOGY, VOLUME 18
Description. Proximal part of stem coiled in an S-shaped bend which is followed by another half-circle of
coil, distal part of stem nearly straight. Proximal portion of stem round, about 14 mm long, diameter
increases from 1 -2 to 1 -6 mm in distal direction, lacking cirri ; longitudinal sutures well developed in middle
of stem. Columnals nodose, with parallel sides in straight parts of column; where the column bends, the
columnals become slightly wedge-shaped, height/average width of proximal columnals about 0T8. Next
part of stem with crescentic cross-section, convex part of crescent occurs on the inside of the coil, concave
part of crescent on outside of the coil, diameter of this stem segment increases from 1-8 to 2-8 mm distally,
cirri lacking, longitudinal sutures obscure. Columnals slightly nodose, somewhat wedge-shaped, ratio of
average height/width ranges from 0-5 to 0-36 with distal columnals having the lowest values. Third region
of stem similar to previous part, having crescentic cross-section, diameter of stem segment decreases
distally from 3-2 to 2-4 mm, several cirri present in distal part of stem segment, adjacent nodals separated
by one or two columnals, longitudinal sutures are poorly developed. Columnals not nodose, proximal
columnals slightly wedge-shaped but distal ones have parallel sides, average height/width of columnals
ranges from 0-25 to 0-35, internodals lacking cirri; nodals with cirri, portion of nodal with cirrus scar
expands, this constricts the adjacent part of the internodal; articular facet for cirrus round, somewhat
protuberant. Distal portion of stem almost straight, with crescentic cross-section, width varies from 2-0
to 3 0 mm, cirri generally present, adjacent nodals usually separated by one columnal, longitudinal sutures
obscure. Columnals not nodose, like those in previous part of stem except that the sides are parallel and
the columnals are not wedge-shaped, average height/width of columnals ranges from 0-35 to 0-30.
Cirri long and slender, longest known cirrus is incomplete, observed length about 30 mm, consisting of
forty-one plates. Cirrals with round cross-section, sides expanding distally to form rim-like process which
bears two to six short spines, some cirrals lack rims but some spines are always present. Crown only known
from distal brachials; brachials uniserial, lacking pinnules, ranging from equidimensional to higher than
wide. Distal part of column not observed.
Comparison. The Pentland species is most closely related to H.fletcheri Salter (1873,
p. 118; Bather 1893, p. 46; Springer 19266, p. 86; 1926a, p. 10; Moore 1962, p. 42)
from the Wenlock of Great Britain and Gotland. In H.fletcheri the cirrals are evenly
nodose whereas those of H. parvispinifer generally possess small angular distal rims
which commonly bear two to six short spines or spine bases. The Pentland individuals
exhibit some variability in the nature of the cirrals while the distal rims range from
prominent to weakly developed or absent, but these are normally seen. The number
of spines varies in increments of two, either two, four, or six. When four or six are
present, the two spines lying within the plane of stem coil are generally the best
developed. H. parvispinifer is distinguished from the Gotland H.flabellicirrus Bather,
1 893 by the shape of the individual cirrals, these being nodose in the Gotland form
and rim and spine bearing in the Pentland animal. Also, the cirri of H. flabellicirrus
are ponderous and expand distally whereas those of H. parvispinifer are more slender
and taper evenly distally. The Pentland species differs from Myelodactylus in the
cirri, which in myleodactylids are elongate and unornamented, but the cirrals of the
Pentland herpetocrinid are wider and either nodose or rim and spine bearing. Also,
the longitudinal sutures are well developed throughout the stem of H. parvispinifer
whereas they tend to disappear on the distal end of the column of Myleodactyhis.
Order cladida Moore and Laudon, 1943
Suborder dendrocrinina Bather, 1899
Family dendrocrinidae Miller, 1899
DENDROCRINUS Hall, 1852
Type .species. D. longukiclylus HM, 1852.
BROWER: SILURIAN CRINOIDS FROM SCOTLAND
651
Dendrocrinm extensidiscus sp. nov.
Plate 75, figs. 1-3; text-fig. 5
Holotype. A well-preserved crown of a young specimen with an attached stem segment on RSM 1897.32.289.
Paratypes. Young crowns: RSM 1897.32.290, 291. Long stem segment with partial cup of a young crinoid
on RSM 1897.32.289. Cup of mature specimen with a long stem segment : RSM 1885.26.78g. Stem segment
with attached cirrus roots: RSM 1897.32.301.
Derivation of name. In allusion to the relatively high columnals.
Type locality. Gutterford Burn Flagstones, ‘Starfish Bed’, Gutterford Burn.
Diagnosis. A species of Dendrocrinus with relatively slender cup, brachials high with
respect to width; column round; distal columnals have relatively large ratios of
height/width compared to most dendrocrinids.
Description. Cup conical with straight walls, height/width ranges from 0-8 to 10. Plates of cup convex
with depressed sutures, otherwise smooth. Infrabasals high, pentagonal, higher than wide, infrabasal
circlet occupies from 17 to 24% of the cup height. Basals large, lateral interray basals hexagonal, higher
than wide, height of basal circlet ranges about 50% of the cup height. CD interray basal septagonal, largest
plate in cup, distally truncated for reception of anal X. Radials basically pentagonal, equidimensional or
wider than high, radial circlet generally represents about 30% of the cup height. Radial facets smooth,
narrow, horseshoe-shaped, sloping outward, width of facet varies from 45 to 60% of the width of the radial.
Radianal poorly known, large, lying under C ray radial. C ray radial pentagonal with wide radial facet,
located above radianal and between anal X and B ray radial. Anal X five-sided, occurring between C ray
radial, radianal, and D ray radial and above truncated CD interray basal. Arms only known in young
crinoid, slender, branching isotomously, once or twice; usually six primibrachs present, rarely five or seven
plates occur, distal primibrach is axillary; number of secundibrachs uncertain, twelve plates present in
unbranched arm segment, branched arm segment probably has roughly the same number of secundibrachs ;
from one to three tertibrachs present. Brachials uniserial, nonpinnulate, smooth, slender, with round or
sharp backs. Nonaxillary primibrachs rectangular; primibrach 1 is shortest primibrach, height/width
ranges from 0-7 to 1-2; other nonaxillary primibrachs much higher, height/width varies from 1-7 to 4 0;
axillary primibrach pentagonal, spear-shaped, height/width ranges from 1-6 to 2-3. Secundibrachs higher
than wide, height/width equals from 2 0 to 3-3. Column round, composed of smooth columnals that are
high relative to width compared to other species of Dendrocrinus, column lacking cirri, only one order of
columnals can be differentiated. Proximal columnals which are immediately below the calyx disc-shaped,
much wider than high, height/width varies from 0-2 to 0-5. Distal columnals of young specimens much
higher than wide, height/width ranges from 1-7 to 4 0. Distal columnals of adult have height/width ranging
from 0-25 to 0-8. Columnals near rooting device have strongly crenulate sutures, height/width of columnals
is about 0-5. Rooting device partially known, consisting of at least two heavy cirri, each of which branches
several times.
Remarks. This species is represented by cups which fall into two height intervals.
The largest crinoid has a cup height of 8 0 mm. The smaller crowns range from I T
to 2 0 mm in cup height. The large specimen is considered conspecific with the
smaller ones because of similarities in outlines of the cup and its component plates
and because all specimens have similar convex plates with depressed sutures. The
main differences between the young and mature specimens are in the column. Distal
columnals of the smaller specimens are much higher than wide and the height/width
ratios of these plates range from 1 -7 to 4 0. The equivalent ratios for the adult crinoid
vary from 0-25 to 0-8 indicating columnals that are wider with respect to height. The
columnals of the mature crinoid are higher than in the young specimens. The dif-
ference in shape of the columnals between the young and adult specimen is attributed
o
652
PALAEONTOLOGY, VOLUME 18
TEXT-FIG. 5. Dendrocrinus extensidiscus sp. nov. A, B, D and B ray views of holotype, counterpart and part,
respectively; note relatively long and slender arms with elongate brachials; the cup is crushed so that the
plates appear wider than in uncrushed specimens ; if the crinoid is interpreted correctly, the C ray radial is
located above the radianal and well above the other radials, this is higher than in normal specimens and the
holotype is considered abnormal in this respect, x 4-9. c, d, lateral ray view of external mould and CD
interray view of internal mould, paratype RSM 1885.26.78g, X 3. The radials are black.
to progressive growth. In most crinoids the height/width ratios of columnals decrease
as the columnals become older and larger (Brower 1973, pp. 298, 299). The arms are
not known in the larger crinoid and these cannot be compared with those of younger
specimens.
Comparison. D. extensidiscus is only remotely related to the Wenlock age species
from North America. These include: (1) Z). longidactylus Hall (1852, p. 193) which
has a cup with slightly rounded walls and more numerous arm branches; (2) D. celsus
Ringueberg (1888, p. 132) which shows eleven primibrachs (compared to about six
in this form), more arm branches, and a column which expands near the calyx; and
EXPLANATION OF PLATE 75
Figs. 1-3. Dendrocrinus extensidiscus sp. nov., note relatively high columnals and slender arms consisting
of elongate brachials, ‘Starfish Bed’, Gutterford Burn. 1, D ray view of holotype RSM 1897.32.289
(counterpart, a well-preserved young crown with long stem segment) and paratype RSM 1897.32.289
(crushed cup of young specimen with long stem segment showing well-preserved columnals, located on
right side of photograph), x4-2. 2, C ray view of holotype RSM 1897.32.289 (part), x4-2. 3, lateral
view of a mature specimen, paratype RSM 1885.26.78g, x 1-4.
Figs. 4, 5. Macrostylocrinus silurocirrifer sp. nov. 4, stem segment with well-preserved cirri, paratype
RSM 1897.32.293, ‘Starfish Bed’, Gutterford Burn, x2-4. 5, lateral view of immature crinoid, paratype
GSE 12791, Deerhope Burn Flagstones, River North Esk, at bend 935 yd N. 30° W. of North Esk
Cottage, x4-8.
PLATE 75
BROWER, Scottish Silurian crinoids
654
PALAEONTOLOGY, VOLUME 18
(3) the peculiar Z).? nodobrachiatus Ringueberg (1890, p. 303) which is probably not
referable to Dendrocrinus because either ramules or pinnules are present according
to Ringueberg : also the cup is wider and there are only two main arms per ray with
primibrach 3 forming the axillary. In addition, D. extensidiscus differs from all other
Silurian species by the relatively long and slender columnals and brachials.
Only two other species of Dendrocrinus, D. rugocyathus Ramsbottom (1961, p. 16)
and D. granditubus Ramsbottom (1961, p. 15), are known from Britain; both are
upper Ordovician. They are characterized by stellate plates in the cup and a pentalo-
bate stem. In D. extensidiscus smooth calyx plates and a round stem are observed.
The two British Ordovician species are closely related to the American upper
Ordovician D. casei Meek (1871, p. 295; 1873, p. 28). This American form has very
similar calyx ornament and shape, stem type, and general stem and crown habit to the
British Ordovician crinoids.
The most similar crinoids consist of a series of middle and upper Ordovician
forms from North America. In general, these and the Pentland animal resemble each
other in having round stems which are non-nodose, similar calyx shapes with smooth
plates and slender arms which branch two to four times. Middle Ordovician forms
in this category are: (1) D. acutidactylus Billings (1857, p. 266; 1859, p. 37);
(2) D. gregarius Billings (1857, p. 265; 1859, p. 36); and (3) D. gracilis (Hall) (1847,
p. 84). Upper Ordovician species are: (1) D. navigiolum Miller (1880, p. 235) and
(2) /).? sp. nov. aff. D.l navigiolum Brower (1973, p. 457). Of all the above species,
D. acutidactylus is judged the closest with respect to over-all morphology. D. extensi-
discus is separated from all the above species by the higher, relative to width, columnals
and brachials. Therefore the affinities of the Pentland crinoid lie closer to Ordovician
forms than to Silurian ones.
Acknowledgements. I cordially thank the following for loan of specimens: Dr. C. D. Waterston (Royal
Scottish Museum, Edinburgh), Dr. W. D. I. Rolfe, Dr. J. K. Ingham, and Miss Sylvia Jackson (Hunterian
Museum, Glasgow), Dr. R. Wilson and Mr. P. Brand (Institute of Geological Sciences, Edinburgh),
Professor G. Y. Craig and Miss Helen Nisbet (Grant Institute of Geology, Edinburgh), and Dr. P. M. Kier
(United States National Museum). Dr. A. Lamont generously donated several important specimens to
the Royal Scottish Museum. Most of this work was completed on academic leave from Syracuse University,
at the Royal Scottish Museum during 1969-1970, where I was kindly helped by Dr. C. D. Waterston and
his staff. The specimens were developed and cast by Mr. Robert Rieke of the Museum. Problems of strati-
graphy, correlation, and palaeoecology were discussed with Dr. Waterston, Dr. Rolfe, Dr. P. Toghill,
and Dr. L. R. M. Cocks.
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J. C. BROWER
Department of Geology
Heroy Geological Laboratory
Syracuse University
Syracuse, New York, U.S. A.
Original typescript received 2 September 1974
Revised typescript received 6 January 1975
A NEW CARINATE PH YLLOCERATID
AMMONITE FROM THE EARLY ALBIAN
(CRETACEOUS) OF ZULULAND, SOUTH AFRICA
by H. C. KLINGER, J. WIEDMANN and W. J. KENNEDY
Abstract. The early Albian sediments of Zululand yield abundant specimens of a keeled phylloceratid, Carino-
phylloceras collignoni gen. et sp. nov., superfically homeomorphous with the desmoceratid Damesites. Investigation
of the suture line confirms the phylloceratid affinities of the genus, which is an independent Cretaceous relative of the
P. ( Hypophyllocems) velledae (d’Orbigny) group, and unrelated to the keeled Jurassic phylloceratids HarpophyUoceras
Spath, 1927 and Menegheniceras Hyatt, 1900.
The early Albian deposits of Zululand (Kennedy and Klinger 1972, 1974) yield rich
ammonite faunas consisting of abundant douvilleiceratids, Lyelliceras lyelli
(d’Orbigny), L. pseudolyelli (Parona and Bonarelli), Neosilesites, Phylloceras {Hypo-
phylloceras) ^ Beaudanticeras\ 'Cleoniceras\ and ^ Sonneratm species, Rosallites,
Ammonoceratites, abundant Anagaudryceras sacya (Forbes), Eubrancoceras aff.
aegoceratoides (Steinmann), and Oxytropidoceras species. Accompanying this
assemblage are abundant specimens of a keeled oxyconic ammonite resembling the
desmoceratid genus Damesites Matsumoto, 1942, and referred to as such in a previous
publication (Kennedy and Klinger 1975).
Subsequent investigation of the external and internal suture of this form revealed it
to be more appropriately referable to the ammonite subfamily Phylloceratinae, as
a new genus and species, Carinophylloceras collignoni.
Location of specimens. The following abbreviations indicate repositories of specimens :
SAS— South African Geological Survey Collections, Pretoria,
UPE --University of Pretoria (Boschoff Collection).
BMNH— British Museum (Natural History).
Full details of locality numbers cited are given by Kennedy and Klinger (1975).
SYSTEMATIC DESCRIPTION
Subclass AMMONOIDEA Zittcl, 1884
Order phylloceratida Arkell, 1950
Superfamily phyllocerataceae Zittel, 1884
Family phylloceratidae Zittel, 1884
Subfamily phylloceratinae Zittel, 1884
Genus carinophylloceras gen. nov.
Type species. Carinophylloceras collignoni gen. et sp. nov.
Diagnosis. Phylloceratid ammonites with fastigate to distinctly keeled venters.
Whorl section ovoid, higher than wide, with maximum width at umbilical margin;
[Palaeontology, Vol. 18, Part 3, 1975, pp. 657-664, pis. 76-77.]
658
PALAEONTOLOGY, VOLUME 18
narrowly umbilicated. Ornament typically phylloceratid, consisting of biconcave
striae. Suture phylloid, with lituid 7, trifid L, saddles EjL asymmetrically diphyllic,
Lj U2 asymmetrically tetraphyllic. Saddles in C/3 asymmetrically diphyllic.
Carinophylloceras collignoni sp. nov.
Plate 76, fig. [a-h\ Plate 77, figs. 1-3; text-figs. 1-3
Derivation of name. The species is named for General Maurice Collignon.
Holotype. SAS A1577 from the Mzinene Formation, Stream Cliff section along the Mzinene River 1200 m
NE. of the Farm Amatis, north of Hluhluwe, Zululand, South Africa, 27° 58' 03" S., 32° 18' 34" E. Locality
35 of Kennedy and Klinger (1974).
Paratypes. Thirty-nine specimens; SAS UMS/2, SAS A1133, and BMNH C78639, C78644, C78647-
C78648, C78767, C78769, C78770 from Locality 35, on the Mzinene River; BMNH C78640-C78643,
C78645-C78646, C78651, C78768 from Locality 36, also on the Mzinene River. SAS H 93D/1, SAS H 93/1,
SAS H 93/2, SAS H 93/3, SAS H 93/5 from Locality 142, Nxala Estate, southern part of Mkuze Game Reserve,
Zululand. SAS EM 91, SAS EM 92, SAS EM 77 from the Msunduzi Pan at 26° 57' 25" S., 32° 12' 40" E. ;
UPE B 33 from the same area at 26° 57' 10" S., 32° 12' 45" E. SAS EM 245a, b, c, SAS EM 93, SAS EM 244,
SAS EM 1 14 from the Ndumu region, northern Zululand at 26° 55' 55" S., 32° 12' 55" E. SAS LJE 134A,
UPE B 463, UPE B 464, UPE B 41 1, and BMNH C78649-C78650 from Locality 174; BMNH C78766 and
C78771 from Locality 171, Mlambongwenya Spruit, northern Zululand. UPE B 23 from Aloe Flats Estate,
northern Zululand at 26° 59' 50" S., 32° 11' 50" E.
All specimens are from the Mzinene Eormation of late early Albian age, Albian III of Kennedy and
Klinger (1975).
Dimensions. All measurements are in millimetres; figures in parentheses are percentages of total diameter.
D = diameter, Wb = whorl breadth, Wh ^ whorl height, U = umbilical diameter.
Specimen
D
Wb
Wh
Wb/Wh
U
Holotype
SAS A1577
149
60-5(41)
88-5(59)
0-68
8-5(6)
Paratypes
SAS EM24c
123-5
44-5(36)
68-0(55)
0-65
80(6-5)
SAS H98/1
77-5
32-5(41)
44-5(58)
0-73
6-0(8)
SAS H93/3
108
40-5(37)
63-5(58)
0-64
7-5(7)
SAS UMS/2
132-5
47-0(36)
77-5(58)
0-60
9-0(6-8)
Description. Coiling is moderately involute with a narrow funnel-shaped umbilicus
(6-8% of diameter). Whorl section is subtrigonal with a fastigate to distinctly keeled
venter. Maximum width is at the umbilical edge. In juvenile stages the venter is
fastigate, but in the adult a distinct keel is developed. The keel is of the floored type,
and, depending on the mode of preservation, may either be present or absent on
internal moulds.
Ornament consists of pronounced biconcave striae which arise at the umbilical
wall, are bent forwards at first, then sweep gently backward near the middle of the
flanks, finally being strongly projected on the outer part of the flanks. They are
bundled at their origin, and much stronger on the outer part of the whorls and venter,
producing a chevron-like ventro-lateral and ventral ornament. On internal moulds
the ornamentation is still present, though very much subdued.
Suture line as for genus. Auxiliary saddles in are triphyllic.
EXPLANATION OF PLATE 76
Fig. \a~h. Carinophylloceras collignoni gen. et sp. nov. Holotype, SAS A 1577.
PLATE 76
KLINGER et al., Carinophylloceras
660
PALAEONTOLOGY, VOLUME 18
E
t
TEXT-FIG. 1 , Sutures of Carinophylloceras collignoni gen. et sp. nov. a, external suture of UPE B464,
X 2. b, external suture of UPE B33, x 2.
Discussion. In the description of the stratigraphy of Natal and Zululand (Kennedy
and Klinger 1975), the present specimens were referred to the desmoceratid genus
Damesites because of the presence of a ventral keel. Homeomorphy with Damesites
is, indeed, very close. Not only the whorl section and the presence and shape of the
keel, but also the degree of shell involution, the course of the ornamentation, and even
the external suture line show such similarities that the genera can scarcely be dis-
tinguished. Examination of the suture line, especially the internal part, reveals that
EXPLANATION OF PLATE 77
Figs. 1-3. Carinophylloceras collignoni gen. et sp. nov. \a-h, paratype BMNH C78644, showing details of
ornament and keel and deep lituid internal lobe in section (arrowed). 2, paratype SAS EM 1 14, showing
lituid internal lobe, x2. 3a-/), paratype BMNH C78768, showing juvenile ornament and fastigate
venter.
PLATE 77
KLINGER et al., Carinophylloceras
662
PALAEONTOLOGY, VOLUME 18
TEXT-FIG. 2. Carinophylloceras coUignoni gen. et sp. nov. External suture of UPE B33, x 2.
TEXT-FIG. 3. Carinophylloceras col-
lignoni gen. et sp. nov. Internal
sutures of SAS EMI 14, showing
overlapping lituid lobes, x 12-5.
KLINGER ET AL.: A NEW CARINATE PH YLLOCERATID
663
it is typically phylloid with a lituid internal lobe, characteristic of all phylloceratinids
(Wiedmann 1968, p. 115; Kullmann and Wiedmann 1970, pp. 11-14). The internal
lobe of Dame sites as figured by Matsumoto (1954, fig. 11, reproduced here as text-
fig. Ab), is intensively frilled, and of desmoceratid type; indeed, no desmoceratids
known possess a lituid internal lobe. Text-fig. 4 shows Damesites sutures for com-
parative purposes.
TEXT-FIG. 4. a, external suture lines of Damesites damesi (Jimbo), after Matsumoto
1954, fig. 10, x4. b, external and internal sutures of a juvenile D. damesi at
D = 8-5 mm. After Matsumoto 1954, fig. 1 1.
Keeled phylloceratids occur in the Jurassic, i.e. Harpophylloceras Spath, 1927 and
Menegheniceras Hyatt, 1900. There are, however, no Cretaceous taxa referable to
these genera and affinities of Carinophylloceras with these forms may be ruled out.
The suture line of Carinophylloceras, with an asymmetrical diphyllic saddle EjL
and asymmetrical tetraphyllic L/t/2, the ornamentation, degree of evolution, and
to a lesser extent whorl section point to affinities with the Albian/Cenomanian
Phylloceras (Hypophylloceras) velledae {sensu Wiedmann 1964), and to the Albian/
Aptian Ph. {H.) cy pris cyprisY dWoi and Termier (Wiedmann 1964, fig. 50, pi. 13,
fig. 3, etc.). Apart from the keel, the whorl section is intermediate between Ph. (H.)
velledae velledae and Ph. (//.) velledae morelianum. The presence of a keel, however,
clearly separates Carinophylloceras collignoni from these forms.
It is interesting to note that within the Tetragonitaceae an analogous development
of a keel occurs in Carinites Wiedmann, 1973, thus also mimicking a desmoceratid
exterior to a certain extent.
Carinophylloceras provides a further example of homeomorphy within the
Ammonoidea, and demonstrates how consideration of the sutural formula can
clarify relationships which are obscure when only external features are taken into
account.
664
PALAEONTOLOGY, VOLUME 18
Acknowledgements. We are grateful to Professor J. Visser (Pretoria University) for placing material from
the Boschoff Collection at our disposal and to Mr. D. Phillips and Dr. M. K. Howarth of the British Museum
for assistance and discussion. The paper is published by permission of the Director of the South African
Geological Survey, Pretoria.
REFERENCES
ARKELL, w. J. 1950. A classification of the Jurassic ammonites. J. Palaeont. 24, 354-364.
HYATT, A. 1900. Cephalopoda, pp. 502-604. In zittel, k. a. von, 1896-1900. Textbook of Palaeontology.
Eastman & Co., London.
KENNEDY, w. J. and KLINGER, H. c. 1972. Hiatus concretions and hardground horizons in the Cretaceous of
Zululand. Palaeontology, 15, 539-549, pis. 106-108.
1975. Cretaceous faunas from Zululand and Natal, South Africa. Introduction, Stratigraphy.
Bull. Br. Mus. nat. Hist. (Geol.), 25, 266-312.
KULLMANN, J. and wiEDMANN, J. 1970. Significance of sutures in phytogeny of Ammonoidea. Univ. Kansas
Palaeont. Contrib. 47, 1-32.
MATSUMOTO, T. 1954. Selected leading Cretaceous ammonites in Hokkaido and Sakhalin, pp. 243-313,
pis. 17-36. In MATSUMOTO, T. (ed.). The Cretaceous System in the Japanese Islands. Jap. Soc. Prom.
Sci., Tokyo.
SPATH, L. E. 1927-1933. Revision of the Jurassic cephalopod faunas of Kach (Cutch). India Geol. Survey
Mem., Palaeont. Indica, N.s. 9, mem. 2, pts. 1-6, 945 pp., 130 pis.
WIEDMANN, J. 1964. Unterkreide-Ammoniten von Mallorca. 2. Lief. Phylloceratina. Abh. Akad. Wiss.
Lit. Mainz. Math.-naturw. KL, 1963, nr. 4, 157-264, 64 figs., 21 pis.
1968. Neue Vorstellungen uber Stammesgeschichte und System der Kreideammoniten. Proceed.
IPU, XXIII d' International Geol. Congress, 93-120, 1 pi.
1973. The Albian and Cenomanian Tetragonitidae (Cretaceous Ammonoidea) with special reference
to the circum-indic species. Eclogae geol. Helv. 66, 586-616, 8 pis.
ZITTEL, K. A. VON. 1884. Handbuch der Paleontologie. 1. Abt. II, Lief. Ill, Cephalopoda, pp. 329-522.
Munchen & Leipzig.
H. C. KLINGER
Geological Survey of South Africa
Private Bag XI 12
Pretoria 0001
South Africa
J. WIEDMANN
Geologisch-Palaontologisches Institut
Universitat Tubingen
Sigwartstrasse 10
D 74 Tubingen
Germany
W. J. KENNEDY
Department of Geology and Mineralogy
Parks Road
Oxford, 0X1 3PR
Typescript received 7 October 1974
Final typescript received 28 November 1974
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Palaeontology
VOLUME 18 ■ PART 3
CONTENTS
Palaeoecology of a bituminous shale— the Lower
Oxford Clay of central England
K. L. DUFF 443
Megaspores and massulae of Azolla prisca from
the Oligocene of the Isle of Wight
K. fowLer 483
Ludlow benthonic assemblages
J. D. LAWSON 509
The trilobite Lejopyge Hawle and Corda and
the middle-upper Cambrian boundary
B. DAILY and J. B. JAGO 527
The ostracod Paraparchites minax Ivanov,
sp. nov. from the Permian of the U.S.S.R., and
its muscle-scar field
M. N. GRAMM and V. K. IVANOV 551
The Bradycnemidae, a new family of owls from
the upper Cretaceous of Romania
C. J. O. HARRISON and C. A. WALKER 563
A new ?bryozoan from the Carboniferous of
eastern Australia
B. A. ENGEL 571
The Hauterivian ammonite genus Lyticoceras
Hyatt, 1900 and its synonym Endemoceras
Thiermann, 1963
C. W. WRIGHT 607
Two Triassic fish from South Africa and
Australia, with comments on the evolution of
the Chondrostei
P. HUTCHINSON 613
Silurian crinoids from the Pentland Hills,
Scotland
J. C. BROWER 631
A new carinate phylloceratid ammonite from the
early Albian (Cretaceous) of Zululand, South
Africa
H. C. KLINGER, J. WIEDMANN and
W. J. KENNEDY 657
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Cover : Marrolithus favus (Salter).
Reconstruction of Ordovician trinuclcid trilobitc, prepared by Or. I. K. Ingham as the symbol lor the Symposium on
the Ordovician System, Birmingham. 1974. Based on silicihed material collected by Dr. R. Addison Irom limestones
of Upper Llandeilo age from Wales.
KIRKLANDIA TEXANA CASTER-
CRETACEOUS HYDROZOAN MEDUSOID
OR TRACE FOSSIL CHIMAERA?
by F. T. FURSiCH and w. j. Kennedy
Abstract. A re-examination of Kirklandia texana Caster, 1945 described originally as a medusoid hydrozoan,
revealed stratinomic, preservational, and morphological features incompatible with interpretations as a body fossil.
An alternative interpretation, with the ‘bell’ of Kirklandia as a feeding trace of Gyrophyllites type and the ‘arms’
as fecal-pellet-lined burrows comparable with Granularia, satisfactorily explains these anomalous features. The genus
Kirklandia and the family Kirklandidae should be removed from the Coelenterata and the medusoid hydrozoans
(order Trachylinida) thus have no unequivocal fossil representatives.
The last hundred years has seen the description of a variety of actual or supposed
fossil hydrozoan medusae. These range from mere stellate impressions on the top
surfaces or soles of sandstones, through lobate concretions and composite moulds
to material retaining details of internal organs, tentacles, and umbrellar ornament
(Riiger and Riiger-Haas 1925; Kuhn 1937; Riiger 1933; Kiderlen 1935; Lorcher
1931; Huene 1901; Caster 1945; Kolb 1951; Sprigg 1947, 1949; Harrington and
Moore 1956; Glaessner 1961, 1962, 1966; Glaessner and Wade 1966; Wade 1968).
These fossils have in turn been used to draw conclusions on topics as distant as
coelenterate phylogeny (e.g. Ruger 1933; Caster 1945) and intertidal exposure
(Ruger 1933).
In the Treatise (Harrington and Moore 1956) some eight fossil genera are tentatively
classed as hydrozoan medusae. Of these, Beltanella Sprigg, 1947 and Ediacaria
Sprigg, 1947 from the late Precambrian Ediacara fauna of South Australia are
undoubted medusoids, but cannot be referred with confidence to any of the coelen-
terate classes (Glaessner and Wade 1966). Acalepha Beyrich, 1849, Acraspedites
Haeckel, 1869, and Hydrocraspedota Kolb, 1951 are doubtfully classed as Trachylinid
medusae, whilst Atollites Maas, 1902 and Palaeosemaeostoma Riiger, 1933 are
definitely trace fossils (Seilacher 1955, 1962; Vialov 1968; Hantzschel 1970 with
references; Grubic 1970 with references). Thus only the genus and species Kirklandia
texana Caster, 1945 remains as a hitherto undisputed fossil record of the medusoid
hydrozoans, the Trachylinida, and the sole member of the Family Kirklandidae
Caster, 1945. This species is known from scores of individuals from the Albian Paw
Paw Formation of Texas, and there is an additional doubtful record of the genus
from the German Dogger (Lorcher 1931). The Cretaceous material occurs typically
as sharp sandstone external moulds preserved in full relief. Caster recognized a
remarkable degree of structure interpreted as a lobate body typically divided by eight
adradial sulci, petaloid stomach pouches, genital sacs with paired gonads, a quad-
rate, functional mouth, and eight apparently rod-like tentacles covered in pustules,
interpreted as nettling structures.
During the summer of 1974 we had the opportunity of studying the holotype and
[Palaeontology, Vol. 18, Part 4, pp. 665-679, pis. 78-80.]
A
666
PALAEONTOLOGY, VOLUME 18
paratype material preserved in the Smithsonian Institution, Washington, D.C., and
a large number of additional specimens housed in that institution and the Texas
Memorial Museum, Austin, Texas. With the paratype material preserved in the
University of Cincinnati Museum, Princeton University Museum, and other collec-
tions (Caster 1945, p. 186) over a hundred individuals are available for study. They
suggest that, on the basis of preservation and morphology, Kirklandia is not a medu-
soid, but rather a chimaera— a chance association of a feeding burrow of Gyrophyllites
type and a fecal-pellet-lined or stuffed hmro'w-Granularia.
THE MATERIAL
Preservation. The Kirklandia material studied here consists predominantly of
depressions— described as natural moulds by Caster (1945)— on the top surfaces of
thin-bedded, ripple cross-laminated fine sandstones with a calcareous cement. The
sandstone slabs are generally less than 10 cm in thickness, and laminations are
generally well preserved, although showing some biogenic disturbance. These are
cylindrical burrows both normal and sub-parallel to bedding, some being empty
(although originally clay infilled), others with meniscus-like back fills. Escape struc-
tures are frequent. Bottom surfaces bear common sole markings; some of these are
of inorganic origin, while others are meandering and branching burrows preserved
in positive hyporelief. Top surfaces are often covered in diverse burrows in addition
to Kirklandia (Caster 1945, p. 186, pi. 4, fig. 6). There are also three supposed natural
casts of Kirklandia preserved as ellipsoidal sideritic and pyritic concretions (Caster
1945, pi. 5, figs. 1-5).
Occurrence. The bulk of the Kirklandia material originates from the area around
Roanoke in Denton County, Texas. The supposed natural casts are from Gaines-
ville, Texas. The former region coincides with what Sellards et al. (1966) describe
as their second facies of the Paw Paw, a blackish lustrous clay with ironstone and
jasper-like concretions and occasional sandstone ledges. The facies around Gaines-
ville is somewhat similar (Hill 1901). The sandstones occur interbedded with clays
which yield an extensive normal marine fauna, including diverse ammonites (Adkins
1920, 1928 ; Clark 1965). The whole is interpreted as representing a relatively offshore
environment.
Morphology. Caster (1945) provides a lengthy description and extensive illustrations
of Kirklandia and only an outline is therefore needed here. Kirklandia consist of
scalloped depressions, generally from 40 to 100 mm in diameter and up to 35 mm
depth. There is, in our view, no consistent symmetry other than radial. Up to three
EXPLANATION OF PLATE 78
Fig. 1 a-h. The holotype of Kirklandia texana Caster, la, the negative epirelief, from the Paw Paw Formation
(Albian), 2 miles west of Roanoke, Denton County, Texas, USNM 136131a. \h, silicone mould of same,
OUM KT 8/P.
Fig. 2. Silicone mould of a specimen from the Paw Paw Formation (Albian) at USGS Mesozoic Locality
22258. Blue Mound, 5 miles south of Haslet, Tarrant County, Texas. OUM KT 9/P.
Fig. 3. A further specimen, preserved as a negative epirelief, from the same locality.
PLATE 78
FURSICH and KENNEDY, Kirklandia
668
PALAEONTOLOGY, VOLUME 18
cycles, each with from five to sixteen lobes, are present within the structure (Plate 78,
figs. 1-3; Plate 79, figs. 1-3; Plate 80, fig. 3; text-fig. 1), six or seven being the com-
monest lobe number. Lobes vary enormously in relative development within cycles,
and may be equal or highly unequal. The divisions between lobes, preserved as
tapering walls of sandstone (Caster’s radial sulci), are likewise variable in develop-
ment, whilst the depressions between vary from deep and narrowly rounded (Plate
78, fig. \a-b) to the shallowest of scoops (Plate 78, fig. 2; Plate 79, fig. 2; text-fig. 2).
One of the most critical features of these lobes is that many are associated with
definite overhangs— partial roofs of sandstone (text-fig. 2).
The outermost cycle of lobes is in general the shallowest and most poorly dilfer-
entiated. The intermediate, generally deeper, cycle consists of two lobe types. There
are elongate petaloid structures extending to the centre of the disc (Caster’s insert
lobes) and those which are reduced, triangular, and peripheral (Caster’s exsert lobes ;
see text-fig. 1). The lobes of the innermost cycle are deep inflated structures corre-
sponding approximately with the position of the longer petaloid ‘insert’ lobes of the
middle cycle. Lobe surfaces commonly show distinct, subparallel concretic striations
(Plate 79, figs. 2-3). Caster regarded these as wrinkles resulting either from desicca-
tion shrinkage prior to burial, or rigor mortis contraction (Caster 1945, pp. 176, 180).
In the centre of the disc there may be depressed areas with a conical protuberance
and an axial and tubular pit (Plate 78, fig. \a-b\ Plate 79, figs. 2-3). The ‘arms’ of
Kirklandia described by Caster are tubular, sometimes branching cavity systems
extending through the sandstone slabs (Plate 80, fig. la-b), or mere depressions on
top surfaces (Plate 80, fig. 6). The tubes are about 10 mm diameter, and their surfaces
are covered in randomly orientated ellipsoidal depressions up to 1-5 mm in length
(Plate 79, fig. 5 and Plate 80, figs. 6 and 7 show silicone rubber moulds of these).
THE MEDUSOID INTERPRETATION
The original view. Text-fig. \a-b shows Caster’s original interpretation of Kirklandia
as natural moulds of the oral or subumbrellar surfaces of trachylinid medusae. The
outer cycle of lobes are interpreted as the peripheral zone of the umbrella, the central
cycle as the gastric lobes, and the inner cycle as gastrogenital sacs. Obscure structures
on some specimens are interpreted as paired gonads within genital sacs. The central
area, conical protuberance, and central pit are interpreted as the mouth and associated
EXPLANATION OF PLATE 79
Fig. \a-b. A paratype of Kirklandia texana Caster, la, the negative epirelief, USNM 136131b, from the
Paw Paw Formation (Albian), 2 miles west of Roanoke, Denton County, Texas, lb, silicone mould of
same, OUM KT 11/P.
Fig. 2. A silicone mould of Kirklandia texana, from the Paw Paw Formation of USGS Mesozoic Locality
22258, Blue Mound, 5 miles south of Haslet, Tarrant County, Texas. OUM KT 13/P.
Fig. 3. Same locality as fig. 2, preserved as a negative epirelief.
Fig. 4. Surface details of a specimen of the fecal-pellet-iined burrows Granularia from the Atherfield Clay
Series (Lower Aptian), Atherfield, Isle of Wight, Hampshire. BMNH A6208 (Stinton Collection), x2.
Fig. 5. Surface details of a silicone mould of the ‘arms’ of Kirklandia texana from the same locality as
fig. 2; compare with the Granularia shown in fig. 4, OUM KT 12/P, x 2.
PLATE 79
FURSICH and KENNEDY, Kirklandia, Granularia
670
PALAEONTOLOGY, VOLUME 18
organs, the mouth being quadrate (text-fig. \a). The granules on the ‘arms’ are
interpreted as stinging cells.
Objections. 1. Preservation potential of medusoids. There are three chief situations
in which medusoids can become fossilized. (1) In the intertidal zone, (2) associated
with very fine grained sediments in restricted environments and burial by special
mechanisms, as is the case of the Solnhofen Limestone and Burgess Shale occurrences,
and (3) as a result of rapid burial by clastic influx in exceptional conditions, as at
Ediacara.
Observations on the preservational potential of medusoids, albeit brief, are wide-
spread. The early studies and experiments of Walcott (1898) are now classic; there
are more recent observations by Trusheim (1937), Wagner (1932), Schafer (1941),
Lincke (1956), Muller (1970), experiments by Hertweck (1966), and extensive bio-
stratinomic discussion by Schafer (1962, pp. 212-216; available in English translation
1972, pp. 157-190, pis. 31b-39a). Medusoids can be preserved only as moulds, and
exposure and desiccation are a prerequisite; they cannot occur as body fossils. The
mould is produced by body and organs within a few hours of stranding, and the
precise organs identifiable on moulds depend on sediment grain size, water content,
and rate of decomposition amongst other factors. As the buried bell decays and
subsides the overlying sediment collapses, and all that remains is a composite mould,
perhaps picked out by a thin clay veneer, above which is a region of disturbed and
crumpled lamination. In some cases the gastrovascular cavity and genital pouches
can fill with sediment, either passively, or as a result of inadvertent ingestion during
pumping motions as the stranded organism tries to escape. These ‘stomach stones’
have preservation potential, and indeed Walther (e.g. 1910) and others have described
such objects.
The second category of fossilization again results in the preservation of the medu-
soid as a composite mould, or mere film of organic material. The best-known examples
are the Solnhofen Limestone, Germany (Barthel 1964, 1970; Van Straaten 1971)
and Burgess Shale, Canada (Whittington 1971; Piper 1972a, b). In both cases the
medusoids (and indeed other fauna) were buried in very fine-grained material,
apparently by turbiditic mud clouds (microturbidites) in local euxinic basins.
The Ediacara occurrences of Australia (Wade 1968; Goldring and Curnow 1967)
are in relatively coarse-grained sediment. Medusoids occur as moulds in two situa-
tions. The commonest is in positive relief on the bottom of sandstone laminae; after
rapid burial tissues decayed, and sediment collapsed into the void to produce a species
of composite mould. If a clay lamina was present between quartzitic laminae, a
counterpart mould may also occur on the subadjacent lamina (Wade 1968, figs. 7,
9, 11). In all cases the mould has an extremely low relief.
2. Comparisons with Kirklandia. The Kirklandia material shows none of the features
of preservation and preservational environment seen in undoubted fossil medusoids.
The fauna, sedimentology, and palaeogeographic setting of the Paw Paw in the area
yielding the specimens studied are offshore, normal marine, with no evidence of inter-
tidal exposure, nor of restricted bottom conditions nor burial in fine-grained sedi-
ment. The material occurs with full three-dimensional relief on the top surfaces of
FURSICH AND KENNEDY: KIRKLANDIA
671
Rr
TEXT-FIG. 1. The medusoid interpretation of Kirklandia. a, oral or sub-
umbrellar view showing the basic four-part symmetry. According to
Caster, the mutability of the species is attained by asymmetrical centri-
petal insertion of exsert lobes and perhaps by splitting of the canaliculate
radii. B, axial section through the restored disc of Kirklandia. Known
features are shown in solid outlines; all others inferred from similarities
to the trachyline hydrozoa. Ar, adradius; Cd, central disc; Cm, delicate
circular corrugations or rugae of the peripheral zone (possibly ring-muscles
or velar muscles); Ex, exsert lobes; G, inferred internal gonads; Ggs,
gastrogenital sacs on the radial canals ; Gu, implied gelatinous umbrella ;
Gw, gastric wall or shrunken residue of gelatinous umbrella; In, insert
lobe; Ir, interradius; Mg, low carina between ovoid depressions on the
swollen protuberances; Pd, paired depressions on the swollen pro-
tuberances (perhaps indications of the gonads within the gastro-genital
pouches); Pf, peripheral field (subumbrella or velum); Pr, perradius;
Uc, umbral concavity of aboral surface (modified from Caster 1945,
fig. 1 and fig. 4).
672
PALAEONTOLOGY, VOLUME 18
sandstones and therefore cannot be compared with the Ediacara material. Further-
more, sections show the Kirklandia cutting across laminations, rather than distorting
laminae, which suggests a post-depositional emplacement. Perhaps the most serious
objection to a medusoid origin is the presence of overhangs in many sections (text-
fig. 2). These could survive only if cementation occurred prior to decomposition of
TEXT-FIG. 2. Cross-sections of Kirklandia lobes (after Caster 1945, fig. 5). Note overhanging rims and partial
roofs of sediment to lobes in many specimens.
the coelenterate tissue. In the environment indicated for the Paw Paw, such de-
composition would take only hours or days; there is none of the petrographic
evidence of early cements reviewed by Bathurst (1971), whilst the distortion by
compaction of some burrows and the obvious cross-cutting relations of others
suggest a relatively late date for cementation. Finally, if these indeed are medusoids,
the absence of material preserved on bottom surfaces is curious. The supposed
EXPLANATION OF PLATE 80
Figs. 1-2. "Caulerpa carruthersi' —Gyrophyllites preserved in three dimensions. Fig. 1 is BMNH 25 (Damon
Collection) a section normal to bedding; fig. 2 is BMNH V 2546 (Damon Collection) a section parallel
to bedding. These specimens have been preserved by early diagenetic cementation of the clay matrix
they were excavated in (compare text-fig. 3a). Kimmeridge Clay (Kimmeridgian) of Sandsfoot, Dorset,
x2.
Fig. 3. Kirklandia texana Caster. Specimen preserved as a negative epirelief on the upper surface of a sand-
stone slab from the Paw Paw formation (Albian) of USGS Mesozoic Locality 22258, Blue Mound,
5 miles south of Haslet, Tarrant County, Texas.
Fig. 4. Granularia sp. BMNH A789 (Wethrell Collection) from the London Clay (Yprisian) of Chalk Farm,
London.
Fig. 5. Granularia sp. BMNH A6154, a club-shaped specimen from the Atherfield Clay Series (Lower
Aptian) of the Lower Greensand, Atherfield, Isle of Wight, Hampshire. Compare with fig. 6 of this plate.
Figs. 6, la~b. Moulds (6, Ih) and silicone impression (7a; OUM KT 10/P) of Kirklandia arms from the
Paw Paw formation (Albian) of USGS Mesozoic Locality 22258, Blue Mound, 5 miles south of Haslet,
Tarrant County, Texas.
PLATE 80
FURSICH and KENNEDY, Kirklandia, Granularia
674
PALAEONTOLOGY, VOLUME 18
‘natural casts’ of Kirklandia are equally unlikely to be medusoids, since all other
workers have concluded that body fossils of medusoids cannot be preserved.
We stress that the depositional environment of the Paw Paw, where Kirklandia
is found, is neither one of fine-grained substrate and rapid burial under restricted
conditions, nor is it intertidal. In addition there is no stratinomic or petrographic
evidence to suggest that these were body fossils buried in sediment and preserved
in three-dimensions by rapid cementation ; they are a post-deposition phenomenon
and their matrix was cemented at a relatively late date.
The biological problems of accepting Kirklandia as a medusoid were discussed by
Caster (1945, pp. 187 et seq.). Thus his genus not only mingles features of the Narco-
medusidae and Trachymedusidae, but also includes many unique traits, the most
striking of which is the enormous variation in the number of body lobes and sym-
metry.
KIRKLANDIA AS TRACE FOSSILS
What we believe to be the true nature of Kirklandia is suggested by Caster’s (1945,
p. 184) comment, where he notes two specimens which ‘show either arms or worm
burrows emanating from the central area of the mould. In one the burrow-like
structure enters the rock, and its termination is unknown. In the illustrated specimen,
the two “burrows” terminate in large fusiform expansions, unlike anything seen on
any of the worm spoor interlacing the matrix of most slabs.’
We have sectioned a number of specimens, and some of these clearly show an open,
or sediment filled, vertical cylindrical burrow extending down into the sandstone slab
from the centre of the ‘mouth’ of Kirklandia. This feature recalls the relationship
demonstrated by Hantzschel (1970, p. 207, pi. 2) in very similar stellate depressions
on the top surfaces of Lower Jurassic sandstones. Hantzschel interpreted these
structures as a surface trace, produced by the surface grazing of an animal dwelling
in the central tube. The difficulty in applying this interpretation to Kirklandia is that
it does not explain the overhangs associated with many of the deeper lobes, the
presence of several cycles of lobes, or the observation that a central burrow is not
always present. We would therefore suggest that Kirklandia in fact represents the
preservation of the distal parts of a much more extensive feeding burrow of Gyro-
phyllites type. Gyrophyllites consists of a vertical shaft from which arise rosettes of
short, simple, tear-shaped lobes, interpreted by Seilacher (1955) and Hantzschel
( 1 962) as feeding structures. Some typical ‘three-dimensional’ Gyrophyllites are shown
in Plate 80, figs. 1-2 and schematically in text-fig. 3.
The preservation potential of Gyrophyllites {text-fig. 3). The preservation of a burrow
in the sedimentary record depends mainly on the following factors; (a) the nature
of the infilling, (b) the sedimentary interfaces through which the trace fossils cut,
and (c) the subsequent diagenetic history of the sediment.
The Gyrophyllites animal seems to have preferred fine-grained substrates, pre-
sumably because of their high organic content. A large percentage of the known
occurrences of Gyrophyllites are therefore in mudstones, silts, or even clays. In these
cases the burrow fill usually does not differ a great deal from the matrix, and after
diagenesis the burrows will be difficult to pick out, especially as compaction will
FURSICH AND KENNEDY: KIRKLANDIA
675
endorelief
TEXT-FIG. 3. The preservation potential of Gywphyllites. a, reconstruction of the complete burrow; b, c,
three-dimensional preservation of the burrow or parts of it by early diagenetic mineralization. The forma-
tion of concretions can be confined either to the burrow fill (6) or to the surrounding sediment (c) ; d, preser-
vation as compacted endoreliefs in fine-grained sediments as in most Flysch and Molasse occurrences;
e, preservation as negative epirelief at clay/sandstone interfaces.
largely destroy their three-dimensional nature. If the fill differs from the matrix, thin
impressions of rosettes can be found on the upper or lower surfaces of slabs— the
common preservation of Flysch and Molasse specimens (text-fig. 3d). To guarantee
a three-dimensional preservation of Gyrophyllites differential diagenesis must take
place. This can either be an early cementation of the infilling, resulting in a concretion
whilst the surrounding sediment remains uncemented (text-fig. 3b). Such seems to
have been the case in, for instance, the three-dimensional specimen of Medusina liasica
of Ruger and Riiger-Haas (1925). Alternatively, the matrix can have been cemented
early in diagenesis, while the burrow infillings remained soft. Damon’s (1888)
specimens of Caulerpa carruthersi from the Kimmeridge Clay of Dorset illustrate
the latter case (text-fig. 3c; Plate 80, figs. 1-2). Here, calcareous/sideritic mudstone
concretions formed around the burrows which are filled with soft clay. Finally, parts
of the Gyrophyllites system can be preserved at sedimentary interfaces, usually at
clay/sand junctions. This is the mode of preservation of the bulk of the ^ Kirklandia’
and many other fossil ‘medusoids’. In these cases, parts of the burrow system.
676
PALAEONTOLOGY, VOLUME 18
especially the more or less horizontal rosettes of lobes, are found as negative epireliefs
on sandstone surfaces (text-fig. 3c), which usually seem to have set the lower limit of
sediment penetration.
Kirklandia as Gyrophyllites. As a deposit feeder, the producer of Gyrophyllites
mined the sediment for food, probably inhabiting a vertical shaft from which the
sediment was explored in a radial fashion. The preservation of Gyrophyllites at a
sedimentary interface suggests the burrows were excavated primarily in clays. These
were exploited for food by the animals shifting their burrows downwards when a
rosette was completed to start a further series of radial excavations at a lower level
(text-fig. 4). When a clay/sand interface was reached, mining generally terminated,
for the sands were low in nutrients due to their larger grain size. In some cases a
probing shaft was extended down into the sand (text-fig. 4c), but always abandoned;
the ‘bell’ of Kirklandia thus represents the lowest rosette or rosettes of tunnels pro-
duced immediately before abandoning the excavation.
Text-fig. 4 illustrates how the great variety of Kirklandia can be explained by
a combination of slight variations in burrowing behaviour and by preservational
phenomena associated with the position of the rosette/rosettes of feeding lobes
relative to the clay/sand interface. The size of the central ‘disc’ depends on the position
of the axial tube of the burrow system (text-fig. 4), from where the radial feeding
lobes originate, relative to the buried sedimentary interface. When well above the
interface (text-fig. 4a), only the distal parts of the feeding lobes reach the sand, and
the result is a large central ‘disc’. As the distance decreases (text-fig. Aa-b) the central
A
TEXT -FIG. 4. Generalized features of Kirklandia interpreted as a feeding burrow of the Gyrophyllites type.
Vertical sections are reconstructions; only the negative epireliefs are preserved. Horizontal shading is
clay, stippling is sand.
FURSICH AND KENNEDY: KIRKLANDIA
677
disc becomes smaller, and the lobes deeper and their more proximal parts are also
preserved (text-fig. Ab). If the animal extended an exploratory shaft down into the
sandstone, then the specimen will bear a central sandstone plug and associated
features— Caster’s (1945) manubrial apparatus (text-fig. Ac).
The presence of two or three rosettes suggests that lobes were in some cases closely
stacked ; this is clearly an efficient means of exploiting the sediment (text-fig. Ad-e),
whilst crowding may also have been produced by final intense exploitation of the
clay above the top of the sand layer before the burrow system was abandoned. The
variation in relative development of rosettes can be explained by the position of the
termination of the axial tunnel to clay/sand interface (compare text-figs. Aa and 3e),
whilst shape and depth of lobes depend on the angle between the axis of feeding lobes
and the central shaft. With an angle of 90°, the lobes will be very elongate (text-fig.
Aa-b) ; as the angle decreases the lobes become deeper, increasingly circular in section,
and, in general, will lack their proximal portions (text-fig. Ad-e).
By combining these variables, it is thus possible to generate the variety of structures
described in the original account of the ‘bell’ of Kirklandia.
Kirklandia "arms' as Granularia. The nature of the ‘arms’ of Kirklandia is equally
explicable in trace-fossil terms; they are simple or branched burrows which were
lined or stuffed with clay pellets, referable to the ichnogenus Granularia Pomel,
1849. The ‘utricating structures’ of Caster (1945) are no more than individual pellets.
The association with Gyrophyllites is thus no more than chance; we do not regard
them as necessarily products of the same animal. Specimens of Granularia from the
English Eocene and Cretaceous are figured for comparison in Plate 79, fig. 4 and
Plate 80, figs. 4, 5a-b. The remaining Kirklandia material can also be interpreted
as trace fossils. The supposed natural casts (Caster 1945, pi. 5) represent no more
than concretions developed around Gyrophyllites systems (text-fig. 5b).
Apart from Kirklandia, it is clear that the supposed medusoid Palaeosemaeostoma
geryonides (von Huene) is also part of a Gyrophyllites, preserved at a sedimentary
interface. The overhanging rim of the lobes in the type species (Kiderlen 1935,
fig. 3) suggests that an origin at a sediment /water interface is unlikely. Lorcher’s
(1931, pi. 1, figs. 1-3) Medusina, from the German Dogger a, referred to Kirklandia
in the Treatise (Harrington and Moore 1956, p. 870) is again part of a Gyrophyllites,
preserved as a negative epirelief at a clay/sand junction. Identical "Kirklandia' stellate
traces in the same preservation in interbedded clay/sandstone successions of Car-
boniferous age from central Texas have been shown to us by Professor J. E. Warme
of Rice University.
CONCLUSIONS
On the basis of stratinomic considerations and morphological criteria, we therefore
regard the interpretation of Kirklandia texana Caster, 1945 as a fossil medusoid as
untenable. We prefer to interpret it as a trace-fossil chimaera, the ‘bell’ being the
distal parts of a feeding burrow of Gyrophyllites type, while the ‘arms’ are fecal-pellet-
lined burrows of Granularia type. This interpretation satisfactorily explains preserva-
tional and morphological aspects of the structures described by Caster which are
inconsistent with a medusoid origin.
678
PALAEONTOLOGY, VOLUME 18
The genus Kirklandia Caster, 1945 and the Family Kirklandidae Caster, 1945
should be removed from the Coelenterata ; Kirklandia should be tentatively classed
as a synonym of Gyrophyllites Heer, 1841 (non Wiedmann 1962, Cretaceous Ammo-
noidea). There is therefore no known fossil record of the medusoid hydrozoa (Order
Trachylinida Haeckel, 1877).
Acknowledgements. We thank Dr. E. G. Kauffman and Mr. F. Collier of the Smithsonian Institution,
Washington, D.C. and Dr. C. Duerden of the Texas Memorial Museum, Austin, Texas for allowing us to
study collections in their care. Professor A. Seilacher, Tubingen, provided useful discussion. Financial
support from the Lindemann Trust (W. J. K.) and the Sonderforschungsbereich 53 ‘Palokologie’, Tubingen
(F. T. F.) is gratefully acknowledged. This is contribution No. 43 of the research programme Konstruktions-
morphologie of the Sonderforschungsbereich 53 ‘Palokologie’, Tubingen.
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F. T. FURSICH
Geol. Palaontologisches Institut der Universitat
Sigwartstrasse 10
74 Tubingen, West Germany
W. J. KENNEDY
Original typescript received 3 March 1975 Department of Geology and Mineralogy
Revised typescript received 20 April 1975 Parks Road, Oxford
V
' • 'I
I
*
i k'j »
*iSJ
AN OUTLINE HISTORY OF SEAGRASS
COMMUNITIES
by M. D. BRASIER
Abstract. The primary ecological role played by seagrasses results from their ability to modify the physical environ-
ment. Trapping and binding of nutrient-enriched sediments encourages deposit and suspension-feeding invertebrates.
Leaves provide a substrate and shelter for flourishing populations of bacteria, algae, protozoans, coelenterates,
molluscs, bryozoans, and echinoderms which in turn contribute CaC03 to the sediment, forming strata with a good
preservation potential. The seagrass community is best developed in tropical and subtropical regions, especially
where alternative nutrient sources are limited.
The geological history of seagrass communities is traced with the aid of foraminifera. Gradual encroachment of
seagrasses into the sublittoral of the late Cretaceous or early Caenozoic was followed in the Miocene by a rapid
dispersal of Thalassia and associated biota, arriving for the first time in the Caribbean and mid Pacific.
Few palaeoecologists would doubt that marine vegetation has played a significant
role in the ecostystems of the past. Unfortunately, this role must remain largely
enigmatic because of the general lack of non-calcified plant material in the fossil
record. The basis for this paper was laid in 1970 when the writer examined recent
seagrass associated biota (especially foraminifera) around the Caribbean. It was
evident from these studies (e.g. Brasier 1973, 1975a, 1975/?, 1975c) and from the con-
siderable work of others, that seagrasses exert great influence over both sedimentation
and ecology in shallow-water habitats. Their first appearance in the sublittoral might
therefore have been marked by a significant change in community structure, and
hence of biofacies. The intention of this paper is to examine briefly the present-day
ecological role of seagrasses and then to trace their geographic dispersal through
time, leading to an assessment of their probable palaeoecological and evolutionary
significance.
SEAGRASS ECOLOGY
Adaptive features. Seagrasses (‘eel’, ‘turtle’, ‘widgeon’, or ‘manatee’ grass) are the
only group of angiosperms known to have successfully invaded the sea. Their means
of attachment to the substrate, propagation, and nutrient absorption {s.l.) differ
considerably from those of marine algae. It is for these reasons that the group has
a more marked effect upon water movement, sedimentation, fauna, and flora. Sea-
grasses somewhat resemble true grasses in mode of growth but are more closely allied
to the freshwater monocotyledons. Den Hartog (1970) has reviewed the main features
of the group, noting that it is not yet certain whether marine forms evolved from
freshwater forms or vice versa. However, genera adapted to either habitat are known
from within the several families so that seagrasses form an ecological rather than
a taxonomic group.
Most seagrass genera have adapted to the aquatic medium by the development of
hydrophilous pollination (Den Hartog 1970). Certain seagrass fruits also float, thus
[Palaeontology, Vol. 18, Part 4, 1975, pp. 681-702.]
B
682
PALAEONTOLOGY, VOLUME 18
aiding plant dispersal (Opurt and Boral 1964). The principal method of increase is,
however, by rhizomatous growth, leaves sprouting at regular intervals along the
rhizome. The size and shape of these rhizomes and leaves varies greatly between
genera and is well reflected in their ecology. Forms with large strap-like leaves and
extensive rhizomatous growth such as Thalassia, Cymodocea, and Posidonia are the
ones with most interest for the palaeoecologist for these have the greatest effect on
the environment and biota.
Habitat conditions. Generalizations concerning the physical factors controlling sea-
grass distribution are difficult to make. Most are found below mean low water and
above 12 m depth. Some of the larger forms (e.g. Thalassia, Cymodocea, Posidonia)
are tolerant of hypersaline conditions (see Brasier 1975a; Logan and Cebulski 1970).
However, differing tolerances of dessication, turbidity, current agitation, sediment
thickness, grain size, and humic content are amongst the factors which cause ecological
zonations of seagrasses in the sublittoral (e.g. Scoffin 1970; Davies 1970; Thomassin
1971 ; Pichon 1971). Furthermore, light intensity and periodicity, as well as tempera-
ture, may affect the latitudinal distribution of species (see Marmelstein et al. 1968).
Influence of seagrass. Seagrass is notable for its ability to influence the character of
the sediment substrate. This results from four more or less independent factors. In
the first place it supplies biogenic CaC03 to the substrate in the form of epibionts and
shells from invertebrates and calcareous algae. Secondly, the dense plant growth
encourages the sedimentation of suspended particles by reducing current velocities
(‘baffling’) and/or trapping the material on the blades. Thirdly, none of these would
be significant were it not for the rhizomes which stabilize the accumulated sediment
and bind it together. Fourthly, it may modify the chemical environment. Seagrass
photosynthesis and respiration are thought to cause variation in the O2 and CO2
content of seawater, which in turn may influence the rate of fixation of CaCOj by
marine organisms (Davies 1970). Seagrasses can also exert a direct control on the
interstitial environment by the reducing action of exudates from plant cells (which
probably arise from bacterial action). The role of these factors in sedimentation has
already been discussed in some detail by Davies (1970). Their significance is a function
of the density of plant growth, the size of the plants, and the width of the leaves,
being greatest where all three are maximal.
Seagrasses are, indirectly, important producers of biogenic CaCOj because their
epibionts are often very dense. Foraminifera and coralline algae may contribute as
much as 5100 gm/m^ per year (see Land 1970; Patriquin 1972; Brasier 1975a),
productivity of both being higher on shallow, turbulent shores. Furthermore, cal-
careous green algae (e.g. Penicillus) which thrive between the blades are copious
producers of aragonitic lime mud (Perkins et al. 1972). In many areas the net result
of these processes is the formation of carbonate banks (see Moulinier and Picard
1952; Scoffin 1970; Davies 1970; Farrow 1971).
SEAGRASS COMMUNITIES
A simplified model of community energetics for tropical seagrass communities is
outlined in text-fig. 1 , compiled from the extensive literature and the author’s observa-
BRASIER: SEAGRASS COMMUNITIES
683
TEXT-FIG. 1 . Generalized diagram of the energy flow in a seagrass community. B = bacteria ; f = foraminifera
and other microherbivores ; a = microscopic algae.
tions. A review or synthesis of the ecology of seagrass communities is beyond the
scope of this study. The comments below are of necessity generalized, with emphasis
on forms likely to be preserved in ancient strata.
Primary producers. Most workers on seagrass communities have commented on the
associated algae (e.g. Scoffin 1970; Taylor and Lewis 1970; Davies 1970) which pro-
vide shelter for many animals. Calcareous algae are common and include benthic
codiaceans and corallines (e.g. Penicillus, Halimeda, Goniolithon, Lithothamnion)
and encrusting epiphytic corallines (e.g. Leptoporolithon = " Melobesia' of many
workers). Abundant non calcified algae (e.g. Laurencia) may also cause entrapment
of sediment, adding to the growth of a seagrass bank.
The primary food source for much of the food web is not seagrass or macro-algae
but populations of benthic and epiphytic unicellular algae and bacteria (see Lee et al.
1966; Lipps and Valentine 1970) or the accumulated detritus (Taylor and Lewis 1970).
Epizoans. Feeding upon these epiphytic ‘blooms’ and detritus are a diverse population
of epizoans, most significant of which (in terms of numbers and diversity) are the
foraminifera. Brasier (1975a) has divided these into primary weed dwellers and
secondary weed dwellers, the latter being facultative forms from sediment substrates.
Primary weed dwellers include many forms of discoidal shape and sessile habit (e.g.
Sorites, Amphisorus, Marginopora, Cyclogyra, Planorbulina) some of which are
684
PALAEONTOLOGY, VOLUME 18
extremely large in size (5 mm + ). Morphological criteria for recognizing other kinds
of weed-dwelling foraminifera have been discussed by Brasier (1975Z?). Micro-
carnivores may include the paradoxostomatid ostracods (McKenzie 1971 ; Maddocks
1966). Organisms using the leaves as a firm substrate for suspension feeding include
annelids (e.g. Spirorbis), bivalves (e.g. Pteria, Barbatia), coelenterates, bryozoans,
and sponges. Detritus feeders (e.g. Cerithium) also flourish on the leaves.
Substrate fauna. The lime mud and decaying organic material trapped and bound by
dense plant growth are an encouragement to bacteria and these together with micro-
algae and detritus may form the staple diet of benthic foraminifera (Lee et al. 1966;
Taylor and Lewis 1970; Lipps and Valentine 1970). Many of these comprise thin-
shelled, elongate miliolids of the genus Cycloforina (see Brasier \915b). Foraminifera,
algae, and bacteria are in turn a major dietary component of suspension and deposit
feeders (Newell 1965; Lipps and Valentine 1970). Gastropods are abundant and
diverse (e.g. Cerithium, Strombus, Cypraea, Olivella, Bulla) and an abundance of
small gastropods may be one criterion for the detection of the former presence of
seagrass (Moulinier and Picard 1952; Davies 1970). Scleractinian corals may also
abound, especially forms tolerant offluctuating salinity and pH (e.g. Porites, Manicina,
Siderastrea).
Dead coral skeletons comprise much of the framework of some seagrass mounds,
affording a hard substrate for colonization by epiphytes and epizoans. Echinoids
(e.g. Diadema, Tripneustes, Lyteclmus, Toxaneustes, Clypeaster) have adapted to
grazing on seagrass leaves and may also be used as palaeoecological indicators (Kier
and Grant 1965).
Infauna. Infaunal deposit and suspension-feeding bivalves of seagrass communities
are often tolerant of low pH and low oxygen supply (Taylor and Lewis 1970; Taylor
1971) and may increase in diversity with increasing organic content (Thomassin
1971). They include lucinoids, tellinids, and pinnids. The deposit-feeding holo-
thurians, polychaetes, and sipunculids thrive in the nutrient-enriched sediments
around seagrass, whilst suspension-feeding crustaceans (e.g. Neaxius and Calianassa)
utilize the organic rich seston (see Farrow 1971 ; Aller and Dodge 1974). The carni-
vorous gastropod Conus is also found in the seagrass community, attracted by the
plentiful food supply and protective canopy (Taylor 1971). Jackson (1972) and
Levinton and Bambach (1975) have discussed the molluscan ecology of tropical and
temperate seagrass communities.
Diversity. Various studies (e.g. Logan and Cebulski 1970; Taylor 1971; Brasier
1975fl, 1975c) have shown that diversity, biomass, standing crop, and productivity
are strikingly greater in seagrass communities than in those of surrounding waters.
This is essentially because of the wide variety of habitats afforded to the fauna and
flora by seagrass. For example, the variety of foraminiferal tests found around Alli-
gator Reef, south of Jamaica, is significantly higher in the vicinity of seagrass (text-
fig. 2). A similar temporal example has been described from Abu Dhabi Lagoon
(Murray 1 970) in which recent colonization by seagrass greatly improved foraminiferal
diversity and standing crop. Conversely, annihilation of the backreef seagrass stands
of Buccoo Reef, Tobago, by Hurricane Flora in 1963, resulted in a towering of
foraminiferal diversity and standing crop (Dr. S. Radford, pers. comm. 1972). The
BRASIER; SEAGRASS COMMUNITIES
685
Textulorio ogglutinons
T. candeiana
Bigenerina irregularis
Valvulino oviedoiona
Vertebralina cassis
Spiroloculina antillarum
Cornuspiramio ontillarum
Quinqueloculino agglutinans
Q bicostata
Q. bidentota
0 candeiana
Q cuvieriana
Q lomarckiana
Q. poeyana
0 polygona
0 quadriiateralis
0 steliigera
Q subpoeyona
Q triconnota
Triloculma carinota
T linneiana
T oblonga
T rotundo
T trigonula
Pyrgo subsphoerica
Sigmoilina orenata
Schlumbergerina alveoliniformis
Miliolinella circularis
Houerina bradyi
H ornatissima
Articulina mexicana
— •=10-20’/. □ =>20’/.
A mucronata
Archoios angulatus
Sorites marginalis
Amphisorus hemprichii
Peneroplis bradyii
P pertusus
P profeus
Monalysidium pohtum
Borelis pulchrus
Bulimina marginota
Botivina sp.
Neoconorbina orbicularis
Oiscorbis spp indet.
0. bertheloti
0 tloridana
D volvulata
Volvulinerio candeiana
Astengerina carinota
Siphonina pulchra
Ammonia beccarii var tepida
Rotorbinetla mira
R rosea
Elphidium discoidale
E poeyonum
Amphistegina gibboso
Cibicides tobatulus
C pseudoungerianus
Ranorbulino acervalis
Cymbaloporetta squammosa
Florilus grateloupi
DIVERSITV ( V volue )
(58)
ED @ i
[m
17 I 20 I 4 ' 38
^
TEXT-FIG. 2. Distribution of recent foraminiferid tests in samples from Alligator Reef,
Jamaica; 1 52 = unvegetated backreef calcarenite (depth 1 m); 153 = backreef muddy
calcarenite from Thalassia meadow (1 m); 155 = muddy calcarenite from Thalassia-
colonized inter-reef channel (2 m); 1 57 = unvegetated inter-reef Halimeda sand (3m);
159 = carbonate-rich terrigenous muds from Thalassia meadow on open shelf (c. 12 m).
Sample numbers refer to H.M.S. Fox/U.C.L. Geology Dept, programme, CICAR 1970.
686
PALAEONTOLOGY, VOLUME 18
epidemic infection of the temperate seagrass Zoster a in the 1930s was also accom-
panied by a lowering of community diversity (see the review by Johnson 1964).
Ancient seagrass assemblages might therefore be expected to show increases in
diversity compared with neighbouring biofacies.
Community dispersal. Before discussing the distribution of the seagrass community
in space and time it is worthwhile to consider the available dispersal mechanisms.
Here the concern is primarily the way in which benthonic organisms have crossed
wide oceanic barriers such as the tropical Atlantic and the mid Pacific. Some of the
possible mechanisms are illustrated in text-fig. 3, none of which can be ruled out on
the basis of chance because of the extremely long time period involved.
TEXT-FIG. 3. Diagram of dispersal mechanisms available to tropical shallow-water benthos for crossing
ocean barriers in the northern hemisphere. Oceanic islands serve as staging posts. The equatorial surface
current (east to west) and undercurrent (west to east) are important. Although dispersal by atmospheric
phenomena (e.g. hurricanes) or by birds and marine vertebrates need not be insignificant.
For certain organisms planktonic larvae are the major means of dispersal. Scheltma
(1968) has shown that such larvae make it possible for some marine species to breach
faunal barriers and colonize new regions. He found that stenothermal tropical larvae
are distributed not only throughout the westerly travelling Equatorial Current but
also throughout the easterly travelling Equatorial Undercurrent, which runs around
the equator. Scheltma concluded that both currents can account for the amphi-
Atlantic distributions of much of the tropical shallow-water benthos between West
Africa and South America.
Although these various currents are of value to invertebrates with lengthy plank-
tonic larval stages, those with short ones, such as the foraminifera, stand little chance
of reaching their destination. Hence Vaughan (1933) suggested that foraminifera
were dispersed by rafting as individuals or small colonies upon seaweed, and Bock
(1969) specifically mentioned Thalassia as the means of their dispersal throughout
BRASIER: SEAGRASS COMMUNITIES
687
the Caribbean region. But whilst many types of foraminifera can live on anchored
seagrass, only those firmly adherent and encrusting forms such as Planorbulina,
Sorites, Amphisorus, Marginopora, discorbids, cibicidids, and phytal miliolids were
found alive and abundant by Brasier (1975c) on grass blades floating on the sea’s
surface. Forms that are better adapted to a sediment-dwelling niche, such as Archaias,
were not only rare or absent on floating seagrass but also on populations of floating
Sargassum and filamentous algae. From this one may expect that rafting on weed,
including seagrass, is an unlikely mechanism of dispersal for forms ill-adapted for
attachment, although not impossible.
That adherent foraminifera are more widely dispersed is evident from studies of
recent forms. Stenothermal ‘phytal’ foraminifera have a more or less pantropical
distribution today, whilst relatively eurythermal phytal forms are almost pandemic
(see Brasier 1975a, \915b). Conversely, typically sediment-dwelling foraminifera of
the tropics, such as Archaias and the alveolinids, are endemic to certain provinces.
Very isolated islands such as Midway Atoll in the mid North Pacific have been
colonized by the firmly adherent Sorites and Marginopora rather than by the loosely
adherent or free-lying Baculogypsina and Calcarina, genera which have not yet made
the journey successfully (see Cole 1969).
The pelagic and pseudopelagic dispersal of these organisms is greatly dependent
on the current velocity, the length of the journey, and the ecological stresses
encountered en route. Because of the nature of the oceanic surface currents, migration
is also more difficult from west to east in the tropics and from east to west in high
latitudes. However, these constraints may be overcome with the aid of very mobile
vertebrates (especially turtles, birds, and man) or special atmospheric and oceanic
phenomena (e.g. hurricanes). All such journeys may have their chances of success
improved by ‘staging posts’, volcanic and coral islands for example. The role of some
of these factors will be considered in special cases below.
DISTRIBUTION IN TIME AND SPACE
Several lines of evidence can be pursued to indicate the origin and evolution of sea-
grass and its associated biota. The plant remains themselves are naturally of prime
importance but are usually rare and indifferently preserved (see Den Hartog 1970).
Unless the reproductive parts have been fossilized, the remains are not easily dis-
tinguished from other monocotyledons, causing dissent as to how meaningful many
of the so-called fossil seagrasses are. Therefore only those recognized by Den Hartog
(1970) are considered here. Unfortunately, the pollen of seagrass lacks exine and is
therefore not preserved. Hence less direct methods must be considered.
It is possible to use elements of the rest of the seagrass community such as fora-
minifera, molluscs, echinoids, and crustaceans as indices of seagrass in earlier seas.
Furthermore, Farrow (1971) and others have shown that the sediments which
accumulate around seagrass communities are distinctive and, if they escape channelling
and other forms of erosion associated with the biotope, they stand a first-class chance
of preservation.
A final and most valuable clue to the history of seagrass and its community lies
in studies of present-day biogeography, especially that of seagrass, which Den
688
PALAEONTOLOGY, VOLUME 18
Hartog (1970) has recently discussed. Examination of those data reveals that sea-
grass distributions fall generally into three associations: the Zostera association
(which may be taken to include Heterozostera, Phyllospadix, Amphibolis, and Posi-
donia) is predominantly of temperate water forms with a more or less bipolar distribu-
tion (text-figs. 4 and 5) ; the Cymodocea association (including also Thalassodendron
and Enhalus) is mostly tropical but is notably absent from the Neotropics and tropical
West Africa (text-fig. 5); the third or Thalassia association (including Halophila,
TEXT-FIG. 4. Recent distribution of Zostera (Zostera) (shown as stipples) and Heterozostera (shown in
black). Modified from Den Hartog (1970) with ocean surface currents after Sverdrup et al. (1942).
Syringodium, and Halodide) is also tropical but differs basically in being present in
the Neotropics but absent from most of the Mediterranean (text-fig. 6). It should be
noted here that seagrasses are, significantly, absent from the coasts of South America
excepting the tropical Atlantic region and single records from Chile and Argentina
(Den Hartog 1970).
These interesting distributions are remarkably similar to the faunal realms of
shallow-water benthic foraminifera. This need be no surprise for they have most
probably evolved side by side throughout most of the Cainozoic. The ecological
requirements of certain foraminifera have become almost dependent upon seagrass,
especially those of the larger soritids (‘peneroplids’). Hence Peneroplis planatus
parallels the Cymodocea association in its distribution (text-fig. 4) and is known to
live attached to that genus (Blanc-Vernet 1969; Davies 1970). Other living larger
foraminifera such as Alveolinella, Calcar ina, Baculogypsina, and Operculina are
further restricted to the Indo-West Pacific. Conversely, Amphisorus hemprichii and
Sorites marginalis appear to parallel the Thalassia association in their distribution
BRASIER: SEAGRASS COMMUNITIES
689
(text-fig. 6) and are known to live as epifaunas on Thalassia (Bock 1969; Brasier
1975a, 1975c). Phytal microfaunas on Zostera are not welt known but appear to be
dominated by the smaller hyaline forms Rosalina, Discorbis, Cibicides, and Planor-
bulina (J. Scott, pers. comm. 1970). These are equally common on other plants and
hard surfaces both in and out of the tropics and therefore do not constitute a specially
adapted seagrass faunule. Soritids and other larger tropical foraminifera are usually
absent from temperate Zostera communities.
TEXT-FIG. 5. Recent distribution of Posidonia (dashed lines), the 'Cymodocea association’ (stippled), and
records of the foraminiferid Peneroplis planaius (black circles). Data from Den Hartog (1970) and others.
Whilst Other invertebrate groups are similarly associated, the advantages of fora-
minifera as palaeoecological indices are well known and numerous (e.g. Brasier
\915b). Nevertheless they can only, at best, be an indication of the presence of sea-
grass and further finds of plants remains themselves or other corroborative evidence
should be looked for.
Cretaceous
The oldest seagrass-like fossils are protozosteroids and cymodoceoids (Den
Hartog 1970). These have been found as imprints and silicified remains in the upper
Cretaceous rocks of Japan and northern Europe (Koriba and Miki 1931, 1960;
Oishi 1931 ; Voigt and Domke 1955). Posidonioids are also known but Den Hartog
considered all these Cretaceous forms to be but poorly adapted to marine conditions.
Furthermore, the numerous other truly terrestrial angiosperms and cycads found
in the Japanese beds must question the validity of referring to the examples as ‘sea-
grasses’. Hence they may have been of little significance to the marine biota of that
690
PALAEONTOLOGY, VOLUME 18
time. However, contemporaneous foraminiferal faunas contain a few questionably
seagrass-adapted forms such as the peneroplinid Vandenbroeckia and meandrop-
sinids such as Broeckina, Edomia, Qataria, and Pseudedomia. These all have a dis-
tinctive Tethyan (i.e. Mediterranean to Indo-West Pacific) distribution (text-fig. 7),
similar to that of the Cretaceous ‘alveolinids’, but the latter are more likely to have
flourished in unvegetated backreef-type sands, as do their descendants. These
alveolinids, peneroplinids, and meandropsinids are absent from neotropical strata,
and suggest that the Americas were effectively isolated from Africa and Europe in
the late Cretaceous (Dilley 1971, 1973).
The present bipolar distribution of the Zostera association nevertheless suggests
that seagrasses were evolving in Cretaceous times and have since become isolated
by continental drift and by the expansion of a more specialized tropical seagrass
flora (Den Hartog 1970). The occurrence of the primitive Heterozostera both around
South Australia and at an isolated locality in Chile (Van Steenis 1962) suggests, like
the marsupial evidence (Cox 1973), an archipelagic link between South America and
Antarctica in the late Cretaceous. The genus is unrecorded from New Zealand
although Zostera occurs there.
Kennedy and Juignet (1974) have recently described possible seagrass bioherms
from the upper Cretaceous of Normandy, although the inferred water depth would
seem to be greater than tolerated by seagrasses today.
Palaeocene- Eocene
Fossil seagrasses of the Cymodocea association are first known from the lower
Eocene of the Paris Basin where Cymodocea occurred with Posidonia (Den Hartog
1970). Significantly their appearance is accompanied more or less contemporaneously
by that of the large foraminifer Orbitolites, whose living, close relatives Marginopora
and Sorites are unquestionable seagrass dwellers. In the Ypresian limestones of
Corbieres, southern France, the analogy with recent seagrass sediments is supported
by foraminifera (especially Orbitolites, Valvulina, and thin-shelled miliolids) together
with burrowing crustaceans and lithological evidence (Plaziat and Secretan 1971).
These fossil seagrass beds even pass laterally into sand blanket calcarenites with the
foraminifer Alveolina and the crustacean Calianassa, much as they do today in the
tropics. This Orbitolites-Alveolina-Calianassa assemblage has also been described
from the Eocene of Somalia (Silvestri 1939).
It is clear from biofacies and lithofacies that some of the later Calcaire Grossier
accumulated under similar conditions. Wright and Murray (1972) have further
deduced from foraminiferal evidence that seagrass stands (presumably of Cymodocea
and Posidonia) were widespread in the middle and upper Eocene of the English
Channel and this is supported by finds of Posidonia in the Bracklesham Beds of
Selsey Bill, associated with an unusually diverse phytal fauna of miliolids, bryozoa,
and molluscs (Curry 1965).
As mentioned above, the alveolinid sand facies of the Eocene occurred also in the
late Cretaceous tropical regions, excepting those of the Americas. This provincialism
was maintained, but the niche of Alveolina may have been occupied in the Eocene of
the American region by Archaias (as it is today) and also by the extinct Yaberinella.
It must also be emphasized that Orbitolites never reached the Neotropics and that
TEXT-FIG. 6. Recent distribution of the 'Thalassia association’ (stippled) with records of the foraminiferids
Sorites, Amphisorus, and Marginopora (black circles). Data mainly from Den Hartog (1970) and Wright
and Murray (1972).
TEXT-FIG. 7. Land, seas, and oceans in late Cretaceous times, with records of fossil cymodoceoids (c) and
protozosteroids (z). Black circles show records of possible seagrass-dwelling foraminiferids (peneroplinids,
meandropsinids, Praeosorites). Stipples represent inferred distribution of ‘tropical’ seagrasses. Palaeo-
geography and palaeocurrents modified from Cox et al. (1973) and Gordon (1973).
692
PALAEONTOLOGY, VOLUME 18
the Cymodocea association is not in evidence there at present, both facts strongly
suggesting that seagrass did not succeed in colonizing this region during the Eocene
(see text-fig. 8). None the less, seagrasses are likely to have had a complete Tethyan
distribution in the early Eocene because Orbitolites occurred at that time as far east
as West Pakistan (Nutall 1925) and has been reported from Tibet and Hyderabad
(Davies and Pinfold 1937).
Other foraminifers which might have been seagrass dwellers are Pseudorbitolites
and 'Taberina daviesi from the Palaeocene of the Middle East (Henson 1950;
Morley-Davies 1971). Rhipidionina, a middle Eocene soritid from Istria and the
Middle East, and Saudia, a dicyclinid from the Palaeocene to middle Eocene of
northern Iraq and Arabia, could also be included here.
TEXT-FIG. 8. Land, seas, and oceans in Eocene times, with records of fossil Posidonia (p) and Cymodocea (c).
Symbols show records of possible seagrass-dwelling foraminiferids: black circles = Orbitolites\ stars =
Saudia-, crosses = Rhipidionina. Stipples represent inferred distribution of ‘tropical’ seagrasses. Palaeo-
geography and palaeocurrents modified from Cox et al. (1973), Gordon (1973), and Ramsay (1973).
It might therefore be concluded that the present-day Mediterranean-Indo-West
Pacific distribution of the Cymodocea association was initiated at least by early
Eocene times and probably during the Palaeocene. The late Cretaceous soritids
mentioned above might have heralded the advent of this association of tropical sea-
grasses for their distribution is remarkably similar, but there are no plant remains
to support this speculation.
BRASIER: SEAGRASS COMMUNITIES
693
Oligocene
The extinction of Orbitolites and similar forms before the end of the Eocene and
their lack of replacement is not easily explained. It was concurrent with the extinction
of many other Palaeogene larger foraminifera (Adams 1973) and may perhaps be
connected with a contraction of the tropical belt in the Oligocene as shown by
palaeoclimatology (Haq 1973). This would have resulted in a scarcity of suitable
habitats. Furthermore, Orbitolites may have had a life cycle extending over three
or four years, as do recent Marginopora (Ross 1972) and the consequent lack of
adaptability may have rendered it vulnerable to fluctuations of trophic source,
habitat, and climate.
It is therefore more difficult to suggest what the distribution of seagrass com-
munities may have been during the Oligocene. Cymodocea has been recorded from
the Oligocene of Bembridge, Isle of Wight (Chesters et al. 1967) which may indicate
that this genus maintained its distribution after the demise of its specialized epifauna.
No details of stratum, locality of preservation are given, however. The miliolid and
rotaliid foraminifera recorded from the Headon Beds of that area were thought by
Bhatia (1957) to indicate the presence of vegetation and do in many respects resemble
those from recent lagoonal seagrass beds in the tropics (Brasier 1975a). The present
distribution of Cymodocea suggests hardiness compared with the other tropical
genera, which concurs with its presence in the English Oligocene.
Backreef sediments of the tropical Americas during middle and late Oligocene
times, contained flourishing ‘microfaunas’ of Miogypsina and Miogypsinoides. These
large, lenticular, complexly structured foraminifera spread rapidly to other tropical
regions, migrating (like the deeper-water Lepidocyclina) to the Indo-West Pacific via
West Africa and the Mediterranean sea (Adams 1967, 1973). Migration in the opposite
direction was apparently difficult at this time. There is, therefore, little to suggest
that seagrasses or their biota were able to reach the Neotropics during the Oligocene.
None the less, Chesters et al. (1967) mention macrofossil records of Cymodocea
from the Oligocene of Florissant, U.S.A., but unfortunately give no citation. Until
there is better evidence this unfigured occurrence may be treated with caution because
Cymodocea is otherwise unknown around the Americas. The fossil (if a seagrass)
would more likely have been of Thalassia, Halophila, or Zostera (?).
Miocene
At the end of the Oligocene and in the early Miocene, the seas began to withdraw
from the Near and Middle East (Savage 1967). This effectively isolated the Mediter-
ranean from the Indo-Pacific province, an event which was further accentuated by
the junction of the Betic peninsula of Europe with Africa in the middle Miocene
(Berggren 1972).
Somewhat paradoxically it is at the same time that a pantropical expansion of
certain shallow-water foraminifera took place, and especially it would appear, of
seagrass-dwelling forms. These included Sorites, Amphisorus, Marginopora, and
Peneroplis, some of which are very similar in appearance to Orbitolites (see text-fig. 9).
They occur in the lower Miocene limestones of the Caribbean region and appeared
at about the same time in the Mediterranean and Indo-West Pacific Region (Adams
1967) and Mid Pacific (Cole 1969) where they still occur today. Equally remarkable
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PALAEONTOLOGY, VOLUME 18
is the fact that the sediment-dwelling Borelis became, at the same time, the first
alveolinid to reach the Neotropics. Spiroclypeus also made a unique appearance
outside of the Indo-Pacific realm at this time (Adams 1973) and McKenzie (1967)
records that the paradoxostomatid ostracods, which are phytal in habit, appeared
and spread very widely in the Miocene. Furthermore, certain coralline algae, other-
wise unknown from the Neotropics, arrived in the Caribbean from the Indo-Pacific
in middle Miocene times (Brasier and Mather 1975).
This new or improved connection between the tropical Americas and the Indo-
West Pacific province in the early Neogene is likely in part to be correlated with the
invasion of the Neotropical area by the Thalassia association. Although largely
TEXT-FIG. 9. Land, seas, and oceans in Miocene times, with the distribution of fossil Cymodocea (c). Black
circles show the wide distribution of the seagrass-dwelling foraminiferids Sorites, Amphisorus, and Margino-
pora. Stipples represent the inferred, almost pantropical, distribution of the Thalassia association. Palaeo-
geography modified from Cox et al. (1973), palaeocurrents conjectural.
absent from the Mediterranean it is unique in being present at isolated localities
around the west coast of Africa (text-fig. 6). This is consistent with a colonization
via the south cape of Africa at a time when the Suez isthmus was closed. Geological
evidence can substantiate this. According to Haq (1973) a climatic amelioration of
5-8 °C occurred in the higher latitudes of the southern hemisphere during early and
middle Miocene times. The probable net northward movement of the African
continent during the Tertiary (Newell 1971) may in combination have made it
possible for Indo-West Pacific shallow-water biotas to ‘round the cape’ and establish
along the west coast of Africa. Closure of the Suez isthmus and Persian Gulf
BRASIER: SEAGRASS COMMUNITIES
695
could also have diverted warm Indian Ocean water down the east coast of Africa
rather than through the Mediterranean as before, rendering the southern cape more
tropical.
That the temperatures were unusually suitable in the early and middle Miocene is
supported by otherwise unprecedented records of Lepidocyclina and Miogypsina
from Angola (Lemoine and Douville 1904), Miogypsina from Brazil (Closs 1966),
and Miogypsinoides from Nigeria (Adams 1973). The present-day occurrence of the
Indo-West Pacific colonist Borelis, around the Atol das Rocas, off Brazil (Tinoco
1965) and also around the Cape Verde and Ascension Islands (Adams 1967) suggests
that these mid-Atlantic islands served as ‘staging-posts’ on the westward route.
Briggs (1974, pp. 90-11 1) has pointed out that the (probably) Miocene island of St.
Helena, Ascension Islands, has a mixed American Indo-West Pacific fish fauna. The
latter, he suggests, arrived on surface currents via the Cape of Good Hope. One may
add that one of the two living species of seagrass shared by the Caribbean with the rest
of the tropics {Halodule wrightii) is the only species extant along the coast of tropical
West Africa but is absent from the Pacific and Mediterranean (Den Hartog 1970).
Other than this, the Caribbean and Indo-West Pacific share ‘twin-species’ of Thalassia,
Syringodium, and Halodule (ibid.).
It is reasonable, therefore, to infer that the Thalassia association of seagrass,
together with other flora and fauna, progressed around the south cape and west coast
of Africa during the warmer spell in the early and middle Miocene, to be the first to
arrive in the hitherto isolated Neotropical waters. After the onset of cooler late
Miocene conditions this southern migration route may have become impassible.
Tropical and subtropical faunas certainly show evidence of climatic restriction at
this later date (Bandy 1968; Newell 1971).
As shown in text-fig. 6, the Thalassia association is widely dispersed at present in
the North Pacific, Halophila reaching as far as Hawaii and Tuamotu (D. R. Stoddart,
written comm. 1974). The seagrass-dwelling foraminifer Marginopora first appeared
in this region during the Miocene and others have arrived since (Cole 1969). This
suggests that this spread of the Thalassia association (especially Halophila) may
also have occurred primarily at that time.
Plio-Pleistocene
Climatic deterioration probably prevented the relatively stenothermal Thalassia
association from colonizing the higher latitudes of North and South America so
that these may have been, as yet, unvegetated by seagrass. Den Hartog (1970) has
suggested that colonization of the north Atlantic shores by Zostera (Zosterella) took
place in the Plio-Pleistocene from the north Pacific (see text-fig. 10). This was pre-
sumably achieved via the ‘arctic’ shores of Canada, including Hudson Bay where
a relict flora remains. The late Pliocene molluscs of England, Iceland, and New
England also share strong north Pacific affinities, which suggests that a two-way
migration of biotas took place across North America before the onset of the Pleisto-
cene glaciations (Ekman 1953, p. 122).
Also in the late Pliocene, the Panama isthmus became closed, isolating the Pacific
and Atlantic communities. This limited the flow of warm water along the west coasts
696
PALAEONTOLOGY, VOLUME 18
TEXT-FIG. 10. Recent distribution of the seagrass Zostera (Zosterella) and a Pleistocene record (z).
of the tropical Americas which once had flourishing Neotropical faunas. The
presently restricted distribution of the Thalassia association (text-fig. 6) on the west
side of Panama attests to this.
DISCUSSION
Perhaps the major paradox to arise from the foregoing is the suggestion that tropical
seagrass communities were expanding their geographic range in the Miocene, when
it is generally held that coral-reef communities became more isolated and impoverished
at about that time (see Newell 1971). Both changes may largely have been brought
about by Alpine earth movements which reached a climax in the Miocene, uplifting
many of the Tethyan continental margins. This must have resulted in a concomitant
increase in the input of terrestrial material, causing in places greater turbidity and
more highly fluctuating levels of nutrients (especially nitrates and phosphates) than
occurred before. According to Tappan (1971) and Valentine (1971) such changes
would have ‘rejuvenated’ the coral community ecosystem, causing a reduction in
diversity. As the seagrass community is apparently more tolerant of great fluctuations
in nutrient levels, acidity, and oxidation, it need not have been adversely affected by
the increased ‘continentality’.
The east to west dispersal of the Thalassia association across the Atlantic in the
Miocene could well help to explain the unusual migratory behaviour of Chelonia
mydas, the seagrass-eating green turtle. Carr and Coleman (1974) recently accounted
for its yearly migration from Brazil to the Ascension Islands as a product of seafloor
spreading. They have suggested that the breeding grounds on the Mid-Atlantic
BRASIER: SEAGRASS COMMUNITIES
697
Ridge gradually spread further and further away from the South American feeding
grounds, starting at least in the early Caenozoic. As a result C. mydas now swims
WNW.-ESE. against the prevailing equatorial current for nearly eight weeks. It
breeds around the Ascension Islands, lays eggs from which juveniles hatch and drift
back with the current to Brazil. For this behavioural pattern to develop they imply
that the home breeding grounds of South America became inexplicably unsuitable
at some stage. Whilst remaining good for feeding, only the mid-ocean volcanic
islands provided suitable beaches for egg laying.
Bearing in mind the extremely conservative breeding behaviour of past and present
amphibians and primitive reptiles, one wonders how Chelonia evolved the above-
mentioned exploratory breeding behaviour whilst retaining conservative feeding
behaviour. The reverse seems to be more in keeping with present evidence. Firstly,
Chelonia is not recorded before the Miocene (Carr and Coleman 1974). Secondly,
other evidence indicates that seagrass (the food source) did not reach the Americas
from the Indo-West Pacific until the Miocene. Thirdly, no such behaviour is known
in green turtles from the Indian Ocean (which would have been inherited if they had
American origins). It is therefore more likely that Chelonia arrived in the Atlantic like
many other organisms, via the Cape of Good Hope during Miocene times. It would
have found the new American feeding grounds by passively drifting across the
equatorial Atlantic, colonizing the Ascension Islands on route. At breeding time
instinctive behaviour still leads them eastwards to the Ascension Islands, their
original base (Brasier 1974).
One should now consider why it was not possible for the Eocene Cymodocea com-
munity to cross the Atlantic from east to west in a similar way. Paradoxically, the
Atlantic is thought to have been narrower in the Eocene (Smith et al. 1973). The
answer may therefore lie in the complex and conflicting evidence for palaeocurrents,
especially that for the Atlantic’s North Equatorial Current which is the one which
could have acted as the requisite agent. Interestingly, this is thought to have been
weaker than the Equatorial Undercurrent and Counter Current in the Eocene
(Ramsay 1973). A warm current from the Indian Ocean into the Mediterranean may
also have pertained at that time, produced by north-east trade winds which then had
a more northerly sphere of influence (Schwarzbach 1963). The deflection of this
current by closure of the eastern end of the Mediterranean may then have helped
to extend the tropical zone to the southern cape of Africa, as previously discussed.
The closure of the western end of the Mediterranean in the middle Miocene could
also have improved the return of the north Atlantic water down the west coast of
Europe and North Africa to form a cooler branch of the North Equatorial Current.
This cooler water effectively isolated the Mediterranean from the Neotropics and
West Africa, as indicated by fossil faunas (Ekman 1953; Berggren 1972).
A final point arises out of the foregoing observations: that the nature of ocean
surface currents and the availability of migration routes can be more important than
geographical proximity in controlling the similarity of shallow-water benthos.
Reconstruction of past continental configurations by analysis of faunal and floral
similarity should therefore be viewed with some caution (see Jell 1974).
c
698
PALAEONTOLOGY, VOLUME 18
SOME PALAEOECOLOGICAL IMPLICATIONS
The gradual encroachment of seagrasses into the sublittoral in late Cretaceous and
Tertiary times may be surmised from recent studies to have been a significant ecological
event. The resultant eutrophication of sediment substrates would have encouraged
the development of deposit-feeding organisms, especially prosobranch gastropods
and miliolid foraminifera. These two groups have certainly diversified greatly since
the Mesozoic. Seagrasses also provide shelter for coralline algae, which likewise
have diversified in the Caenozoic (Wray 1971).
One may further suggest that those organisms feeding on herbivores and detritus
feeders, such as teleosts and rays, have benefited from the innovation. The archaic
turtles could have survived their marine reptile contemporaries after the Mesozoic
because of their gastropod and seagrass diet. Mosasaurs were probably used to
a cephalopod diet— hence were ill-adapted to exploit such a trophic innovation (see
Nicol 1961).
The more rapid encroachment of seagrasses into Neotropical waters in the Miocene
must also have been a significant event. At a time when land barriers were making
biotic exchange with the rest of the Tethyan tropics increasingly difficult, seagrasses
could have aided the dispersal of sessile organisms across the Atlantic and probably
to islands in the mid North Pacific. Such relatively sudden innovations are thought
to cause rejuvenation of the ecosystem, threatening specialized tropical organisms
with extinction (Tappan 1971). The forms most likely to suffer in the Miocene of
the Neotropics were those endemics adapted to lower nutrient levels or to coarser
and less stable substrates. Hence it may be no coincidence that the specialized, sedi-
ment (?) dwelling Neotropical foraminifera Miogypsina and Lepidocyclina became
extinct in their own province during the middle Miocene and not long after in other
regions. Their extremely large size, structure, and palaeoecology suggests that they
cultured symbiotic algae, which often indicates a relatively low but steady external
food supply. If the food supply was increased and trophic stability was upset by the
rejuvenation of continental margins and oceanic islands, together with the encroach-
ment of seagrasses, these foraminifera would have had difficulty in adapting, especially
if they had long life cycles (see Tappan 1971 ; Valentine 1971). Other endemics, such
as scleractinian corals, seem to have suffered in a like manner (see Newell 1971).
Unfortunately, it is not yet possible to say to what extent the above speculations
are justified but they point to some interesting lines for future research.
CONCLUSIONS
The distributions of Recent and fossil seagrasses are similar to the distributions of
Recent and fossil seagrass-dwelling foraminifera. The latter may therefore be used
as indices (only) of the geographical dispersal of seagrass communities through time.
The suggested dispersal patterns are biased towards tropical seagrass communities
because tropical foraminifera indices are more distinctive than temperate ones and
are therefore more reliable.
Seagrass communities were probably present in the shallow sublittoral waters of
the Tethys in late Cretaceous times and almost certainly in Eocene times.
BRASIER: SEAGRASS COMMUNITIES
699
The gradual encroachment of seagrasses into the sublittoral in late Cretaceous
to early Tertiary times can be surmised from recent studies to have resulted in
three significant modifications of the ecosystem: (i) increase in habitat diversity;
(ii) eutrophication of sediment substrates; (iii) additional means of dispersal for
sessile organisms (i.e. rafting).
The more or less contemporaneous radiation of deposit feeding and epiphytic
gastropods and miliolid foraminifera (including soritids) in the late Cretaceous and
Tertiary could be attributed in part to these factors, particularly in tropical waters.
A more rapid colonization of the relatively isolated Neotropical and mid-Pacific
sublittoral waters by seagrass communities seems to have occurred during the Mio-
cene. Seagrasses, epibionts, and associated faunas (including the green turtle Chelonia)
were probably dispersed by oceanic surface currents from the Indian Ocean via the
Cape of Good Hope and the islands of the Mid-Atlantic Ridge. Earth movements
may also have aided dispersal at this time by the provision of broader shelves and
new islands.
The Miocene earth movements and seagrass dispersal could likewise have con-
tributed to the extinction of specialized endemic faunas such as Lepidocyclina and
Miogypsina.
The ecological effects were and are most marked in tropical carbonate environ-
ments where the influence of land masses (‘continentality’) is low and alternative
sources of organic and inorganic nutrients are few.
The similarity of fossil assemblages is not always a reliable indication of their
former (or present) geographical proximity. Caution should therefore be exercized
in applying techniques of similarity analysis to continental reconstructions.
Acknowledgements. The author acknowledges the valuable assistance of many who took part in CICAR
1970 (Co-operative Investigation of the Caribbean and Adjacent Regions), notably Professor T. Barnard
and Dr. A. J. Smith who organized the U.C.L. Geology Department contribution, Lt.-Cdr. D. Scott of
the R.N. Hydrography Dept., the officers and crew of H.M.S. Fox and H.M.S. Fawn, and colleagues
Dr. P. Dolan, Dr. P. Wigley, Dr. S. Radford, Dr. J. Scott, and Mr. J. White. Dr. D. R. Stoddart, Dr.
J. W. Murray, and Dr. R. Goldring are thanked for their constructive criticisms of the manuscript. Messrs.
A. Cross and J. Watkins of Reading Geology Department prepared the illustrations.
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M. D. BRASIER
Typescript received 20 June 1974
Revised typescript received 1 3 December 1 974
Geology Department
The University
Cottingham Road
Kingston upon Hull, HU6 7RX
THE BIOSTRATIGRAPHY OF
THE UPPER ORDOVICIAN AND
LOWER SILURIAN OF SOUTH-WEST DYFED,
WITH COMMENTS ON THE HIRNANTIA FAUNA
by L. R. M. COCKS and d. price
Abstract. Late Ordovician and early Silurian beds around Haverfordwest, South-west Dyfed (Pembrokeshire),
Wales, are remapped and their faunas reviewed. Five formations are defined, representing a fairly continuous
succession from mid Ashgill (Cautleyan) to late Llandovery (Telychian). New faunal evidence places the local
Ordovician-Silurian boundary higher than has been previously suggested, within the Haverford Mudstone Forma-
tion. The world-wide late Ashgill Hirnantia fauna is reviewed, and the conclusion reached that it represents
a diachronous animal life assemblage and that it need not necessarily represent the latest local Ordovician fauna.
Rock successions spanning the Ordovician-Silurian boundary are not common and
with the current world-wide reassessment of late Ordovician and early Silurian rocks,
their correlation, faunas, and ecology, it is desirable to review and evaluate the more
important sections available.
One such section occurs in South-west Wales. Here a series of large thrust-faults
separate blocks of various Palaeozoic ages and the upper Ordovician and lower
Silurian are exposed together within one of these blocks around the town of Haver-
fordwest. Our field-work there, together with work on museum collections, provides
new faunal and stratigraphical evidence in the light of which earlier accounts of late
Ordovician and early Silurian strata in Pembrokeshire need revision. We attempt
also to rationalize the stratigraphical terminology and we review the position of the
Ordovician-Silurian boundary within the area.
PREVIOUS WORK
Marr and Roberts (1885) first recognized and determined a detailed succession within
Ordovician and Silurian rocks in the neighbourhood of Haverfordwest. They
regarded the ‘Slade Calcareous Shales’ (the highest unit of the " Trinucleus seticornis
Beds’ of their succession) as forming the summit of the Ordovician System. These
shales were succeeded by their ‘Conglomerate Series’, represented at Haverfordwest
by the unit to be subsequently designated the Cethings Sandstone, and elsewhere by
conglomerate or grit or both. The relationship of the Conglomerate Series to higher
horizons was not entirely clear to them, but in the Cethings railway cutting at Haver-
fordwest the sandstone mentioned above was succeeded by strata which graded
upwards into the fossiliferous horizons near Haverfordwest gasworks, which they
regarded as of definite lower Llandovery age. Outcrop patterns elsewhere supported
this relationship. Provided the position of the Conglomerate Series was as this
evidence suggested, they argued (1885, p. 489) that it formed ‘a satisfactory base to
[Palaeontology, Vol. 18, Part 4, 1975, pp. 703-724, pis. 81-84.]
704
PALAEONTOLOGY, VOLUME 18
the Silurian rocks of this area’ and that it should be included within the lower Llan-
dovery. The horizon was lithologically compared with the Mulloch Hill Con-
glomerate (now Lady Burn Conglomerate, Cocks and Toghill 1973) of the Girvan
district, Scotland (Lapworth 1882).
The Geological Survey (Strahan et al. 1914) accepted this conclusion and followed
Marr and Roberts in regarding the Slade (and Redhill) Beds as the youngest of the
Ordovician formations. The succeeding strata the Survey termed the ‘Basement
Beds’, comprising at Haverfordwest the Cethings Sandstone (named for the first
time) with shale units above and below it. Elsewhere conglomerates were variably
developed within or at the base of the shales beneath the sandstone.
For Strahan and his colleagues the problem of delimiting the local Ordovician-
Silurian boundary was primarily one of mapping the junction between the Basement
Beds and the Slade and Redhill Beds, a task in which the Cethings Sandstone played
an important part as a marker horizon. It was on this basis that the rich faunas
collected by V. M. Turnbull from localities near St. Martin’s Cemetery, Haverford-
west, were placed within the Silurian (Reed 1906), even though Reed (1905, pp. 98,
103) had earlier regarded them as of ‘Upper Bala’ age. There was some palaeonto-
logical evidence for the lower Silurian age, mainly from the misidentifications
of graptolites found in the succession (see p. 714), but it is clear, particularly from the
accounts written by Cantrill (1907 and in Strahan et al. 1914, p. 101), that the chief
reason for reconsidering the age of the St. Martin’s Cemetery faunas was their
occurrence above the Cethings Sandstone marker horizon. Strahan et al. (1914) also
comprehensively described the succeeding Silurian horizons.
Since the publication of the Survey Memoir, and also following Jones (1925,
p. 354), the horizon yielding the St. Martin’s Cemetery fauna has been widely regarded
as of lowermost Llandovery age. Recently, however, Ingham and Wright (1970,
p. 240) recognized elements of the Hirnantia fauna amongst the brachiopods and
trilobites recorded from this horizon and suggested for it a Hirnantian age.
STRATIGRAPHY AND FAUNAS
Due to various palaeontological and stratigraphical misconceptions, outlined below,
the previous rock terminology is in several parts unsatisfactory and we have found
it necessary to present a revised succession for the Haverfordwest area (Table 1).
Geological maps of the area are shown in text-figs. 1 and 2. Each rock unit and its
fauna is now reviewed in turn.
Slade and Redhill Mudstone Formation
Lithostratigraphy. The ‘Redhill Shales’ and ‘Slade Calcareous Shales’ of Marr and
Roberts (1885) were grouped by the Geological Survey into a single formation, their
‘Slade and Redhill Beds’, since in certain developments of their ‘northern type’
(Strahan et al. 1914, pp. 55-56) they had been unable to map the two divisions
separately. Price (1973) has considered these beds in part and also found that dis-
tinctions between various levels within them are not consistent over any area and
that even where broad lithological distinction between an upper and a lower level
can be made, the boundary region is very vague. We therefore propose that these
COCKS AND PRICE: ORDOVICIAN AND SILURIAN BIOSTRATIGRAPHY 705
TABLE 1. Stratigraphical terminology within the Haverfordwest area.
Standard Scale
IDWIAN
a
>-
YOUNGER
q:
LU
>
C5
Q
<
_l
1
RHUDDANIAN
HIRNANTIAN
tP
X
CO
<
RAWTHEYAN
a
OLDER
Strahanetat, 1914
This paper
Fbsitions of selected localities and faunas
UZMASTON
BEDS
( Lower part of )
GASWORKS
SANDSTONE
GASWORKS
MUDSTONES
CARTLETT
BEDS
MILLIN MUDSTONE
FORMATION
(Lowest part of)
GASWORKS SANDSTONE
FORMATION
(85 metres)
HAVERFORD
MUDSTONE
FORMATION
(370 metres)
basement CethingsSondstone
BEDS
PORTFIELD
'Cuckoo Shale Member
Cethings SarxJst. Member
FORMATION
(65metres) (Scotchwel! Shale MemI
iber
SLADE a REDHILL
BEDS
(Highest part of)
SLADE a REDHILL
MUDSTONE FORMATION
(Highest part of)
• M Clonnda undata,
Eocoelia curtisi,
Eoplecfodonta penkiHensis,
Leptaena purpurea Sc.
Rich shelly Rhudtjanian
faunas with Catymene,
Dalmanites, Ctorinda,
Stncktandia Sc.
R Stricktandia
Q
— CHmacograptus cf norma! is
Faunas with Mucronaspis
mucronato, Anisopieureiio
^ gracilis, Eospirigerina,
Eopiectodonta, Leptaena ac
— ■P—'! CHmacograptus norma iis
St Marlin’s Cemetery horizon,
localities A,D,EandN;
Hirnantia fauna
— ? OnnieUa
F Tretaspis %^^Eochonetes,
Plectothyrella
Tretaspis, Stenopareia, Diacaiymene
I Chasmops ^Z-,PhoHdostrophia
horizons should be regarded as a single formation formally designated the Slade
and Redhill Mudstone Formation. The type development should be taken as the
outcrop strip immediately north-west of Haverfordwest, which includes both Marr
and Roberts original localities along the B4330 road and also the outcrops along
the A487 road between St. Martin’s Cemetery and the school near Pelcomb Cross,
which are much more continuous and include exposures near both base (Grid Ref.
SM 9140 1820, loc. 1 of Price 1973) and summit (Grid Ref. SM 9463 1575, see
text-fig. 2) of the formation.
Fauna. Price (1973) has dated the lower parts of the formation, which is diachronous
within the Cautleyan Stage at its base, and reviewed the trilobite fauna. In this paper
we consider only the youngest part of the formation. The stratigraphically highest
diagnostic fauna from the Slade and Redhill Mudstones known so far comes from
locality I on text-fig. 2, a disused quarry on the west bank of the Western Cleddau
(Grid Ref. SM 953 162). As well as brachiopods (pp. 706, 709) and other shelly
fossils, the following trilobites occur:
Calyptaulax sp. (PI. 81, fig. 6), Chasmopsci. >7rarri (Reed, 1894) (PI. 81, fig. 4), "Diacaiymene' cf. marginata
Shirley, 1936 (PI. 81, figs. 1-3) (for discussion of the genus see Temple 1975, pp. 146-149), Remopleurides
sp. (PI. 81, fig. 5), Stenopareia bowmanni Salter, 1848 (PI. 81, fig. 7), Tretaspis cf. hadelandica St0rmer, 1945
brachystichus Ingham, 1970 (PI. 81, figs. 8, 9).
706
PALAEONTOLOGY, VOLUME 18
Tretaspis hadelandica brachystichus ranges up to Rawtheyan Zone 6 in the north
of England (Ingham 1970) and its probable descendant, T. latilimbus (Linnarsson)
distichus Ingham, appears in Zone 7. When histograms of fringe characters are
plotted separately for forms from Zones 5 and 6 (Ingham 1970, text-fig. 10) there
are differences indicating a gradual change towards T. 1. distichus. The fringe characters
of the Slade and Redhill Mudstone specimens are like those of the Cautley forms
from Zone 5 and also like those of specimens from the highest part of the underlying
Sholeshook Limestone (probably Cautleyan Zone 3), but it is not known whether
the changes in Welsh populations exactly paralleled those seen in the north of
England. All that can yet be said is that the Slade and Redhill specimens are earlier
than Zone 7 and most likely to be low to mid Rawtheyan, an age consistent with the
rest of the fauna.
Cocks (1968, p. 304) reported a specimen of Tretaspis from a temporary exposure
near St. Martin’s Cemetery (locality F on text-fig. 1, Grid Ref. SM 9434 1570), in
an area marked on the Survey map as early Silurian. The specimen (BM It. 13243)
is certainly a Tretaspis, although it is too incomplete to allow specific determination,
but our recent remapping, with the aid of many new exposures in the foundations
of a housing estate, shows the locality to be unequivocally within the highest Slade
and Redhill Mudstones. Thus, as far as is known, there is still no authenticated record
of Tretaspis from rocks of Silurian age.
The brachiopod faunas of the Slade and Redhill Mudstone Formation are much
in need of exhaustive collecting and systematic revision. The assemblages differ much
from place to place. The temporary exposure mentioned above (locality F) just west
of St. Martin’s Cemetery, was dominated by Eochonetes aff. advena Reed, 1917
(PI. 83, figs. 7, 9, 12), a genus hitherto unknown outside its type area at Girvan,
Scotland, which was in late Ordovician time on the further side of the lapetus Ocean.
Eochonetes made up of 84% of the collection (n = 287), with Plectothyrella at 8% the
next most common element. (The method of calculating percentages follows Ziegler
et al. 1968, p. 4.) Topographically close, but over 50 m lower in the formation, a
temporary exposure at SM 942 158 (locality G on text-fig. 2) yielded a quite different
assemblage dominated by Eostropheodonta (43%, n ^ 181), amongst thirteen different
EXPLANATION OF PLATE 81
Figs. 1-3. "Diacalymene' cf. marginata Shirley, x 3. 1, SM A53047a, internal mould of enrolled exo-
skeleton, dorsal view of anterior part. 2, SM A85288a, internal mould of distorted enrolled specimen,
cranidium in dorsal view. 3, SM A3 1207, internal mould of cranidium in dorsal view.
Fig. 4. Chasmops cf. marri (Reed). SM A30961, internal mould of incomplete pygidium, dorsal view, x 2.
Fig. 5. Remopleurides sp. SM A30966, internal mould of pygidium, dorsal view, x 8.
Fig. 6. Calyptaulax sp. SM A30960b, cast from external mould of right free cheek, dorso-lateral view, x 3.
Fig. 7. Stenopareia howmcumi (Salter). SM A30943, internal mould of incomplete cranidium, dorsal view,
X 2.
Figs. 8, 9. Tretaspis cf. hadelandica St0rmer brachystichus Ingham. SM A53049a, b, internal mould and
cast from external mould of cephalon, dorsal views, x 5.
All specimens from the high Slade and Redhill Mudstones of locality I on text-fig. 2. Originals of figs. 1,
8, and 9 collected by Mrs. M. R. Cave, originals of figs. 2 and 4-6 collected by V. M. Turnbull, original
of fig. 3 collected by D. P., original of fig. 7 collected by J. E. Marr.
PLATE 81
COCKS and PRICE, Ordovician trilobites
708
PALAEONTOLOGY, VOLUME 18
TEXT-FIG. 1. Geological map of the area between Little Cuckoo and St. Martin’s Cemetery, west of Haverfordwest. Lettered localities are those
referred to in the text.
COCKS AND PRICE; ORDOVICIAN AND SILURIAN BIOSTRATIGRAPHY 709
brachiopods and nine species of other groups. The Sowerbyella present here at 3%
is of interest in that it appears morphologically indistinguishable from the Eochonetes
mentioned above from higher strata, apart from the lack of perforations along the
hinge-line, and suggests an ancestor for the aberrant Eochonetes stock. Another
assemblage, from the old quarry near the Western Cleddau (locality I), yielded
a fauna in which Pholidostrophia (Eopholidostrophia) matutinum (Lamont, 1935),
a species previously recorded only from the Cautleyan of the Girvan area and of
Cautley itself, occurs commonly (BM BB 69611-69614). A further exposure in the
formation south of Little Clerkenhill, 9 km west of Haverfordwest (Grid Ref. SN
045 150) also yielded Eoehonetes (26%, n= 173) and Pleetothyrella (13%), but the
most common brachiopod (at 39%) was an enteletacean probably related to Resserella.
In no part of the Slade and Redhill Formation, however, do we consider a Hirnantia
fauna to be present, although several of its constituents occur at different localities
and horizons within the formation.
Portfield Formation
Lithostratigraphy. The name ‘Portfield Formation’, after the region of Haverford-
west known as Portfield (see Portfield House on text-fig. 1), is proposed for the unit
designated ‘Basement Beds’ by Strahan et al. (1914)— an unsatisfactory term because
of its erroneous assumption about the position of the unit in relation to the Ordovician-
Silurian boundary, as well as its lack of a local name. The Portfield Formation is
divided into three members, two of shale with an intervening sandstone.
The type section is in the railway cutting at Cethings (see text-fig. 2; Grid Ref.
SM 966 161). At the north-east end of this cutting is exposed the unit which we
propose to designate the Cethings Sandstone Member; here a thickness of about
8-5 m of well-sorted, fine-grained grey to buff sandstone dipping at 62° to the south.
Below the sandstone, dark sooty shales of the Scotchwell Shale Member (named
after Scotchwell, see text-fig. 2) are poorly exposed in a shallow ditch on the east
side of the railway line. Their thickness was estimated by Strahan et al. (1914, p. 89)
as at least 46 m. Shales of similar aspect form the Cuckoo Shale Member (named
after Little Cuckoo, text-fig. 1), exposed to a thickness of about 9 m above the sand-
stone and grading up over a distance of about 1 m in bioturbated beds into the greener
mudstones of the overlying Haverford Mudstone Formation. From regional mapping
the shales of the lowest member appear to be conformable with the underlying Slade
and Redhill Mudstone Formation, but the junction is nowhere exposed.
The sequence in the Cethings railway cutting applies over a region from about
7 km west of Haverfordwest to about 4 km to the east. Further to the east con-
glomerates are locally and variably developed at or near the base of the formation,
one of the most prominent forming the hill at Robeston Wathen (Grid Ref. SN
081 158); further details are given in Strahan et al. 1914, pp. 83-101. The Cethings
Sandstone Member, although in places attenuated and locally containing some
intercalated shale bands, is persistent in the sequence at least as far east as Narberth
and Lampeter Velfrey and forms a valuable mapping horizon.
Fauna. The Portfield Formation is largely barren of fossils. We have found none and
the only known specimen (SM A32094) was collected by V. M. Turnbull from the
710
PALAEONTOLOGY, VOLUME 18
Scotchwell Shale Member in the Cethings railway cutting; it comprises the con-
joined valves of an enteletacean brachiopod, perhaps referable to Onniella.
The St. Martin’s Cemetery Horizon
Litbostratigrapby. V. M. Turnbull made painstaking collections, now in the Sedg-
wick Museum, from two fossiliferous localities (D and E on text-fig. 1) at the roadside
west of St. Martin’s Cemetery. Because the fossils were described (Reed 1905, 1906,
1907) before the complete survey of the area (Strahan et al. 1914), a stratigraphical
name, variously the St. Martin’s Mudstone or the St. Martin’s Cemetery Beds or
Formation has been applied to the rocks from which Turnbull’s fossils came.
The original localities, though no longer exposed, can be accurately located and
lie at the junction of the Portfield Formation and the Haverford Mudstone Forma-
tion; the fossiliferous horizon involved was almost certainly that also seen in the
Cethings railway cutting. There a fossiliferous band about one metre in thickness
occurs at the same junction (locality N on text-fig. 2), fossils from its lowest part
having a dark shaley matrix and those from the upper part a light, more silty one.
Most specimens from the St. Martin’s Cemetery localities have either a light siltstone
or olive-green mudstone matrix which, as at Cethings, is often bioturbated. Elements
of the same fauna, in a matrix similar to that from the St. Martin’s Cemetery localities,
are known from Little Cuckoo (locality A, text-fig. 1), suggesting that there is a
constant band from Little Cuckoo, through St. Martin’s Cemetery to Cethings railway
cutting, a distance of approximately 4 km.
This fossiliferous band, because of its thinness and because its dominant lithology
is more similar to the rocks above rather than below it, we include within the basal
part of the Haverford Mudstone Formation. We refer to the band informally as the
St. Martin’s Cemetery horizon and recommend that the designation ‘St. Martin’s
Cemetery’ should not again be used as a separate formal stratigraphical name for
a formation or member.
EXPLANATION OF PLATE 82
Figs. 1, 2. Lichas cf. laciniatus (Wahlenberg), x2. 1, SM A32014, internal mould of cranidium, dorsal
view. 2, SM A4650, internal mould of pygidium, dorsal view.
Figs. 3-8. Otarion cf. megalops (M’Coy). 3, SM A85575, internal mould of left free cheek, dorsal view, x 6.
4, SM A4648, internal mould of cranidium, dorsal view, x 8. 5, 6, SM A32068, internal mould of
cranidium, right-lateral and dorsal views, x 8. 7, SM A85576a, internal mould of cranidium, dorsal
view, X 8. 8, SM A4647b, internal mould of cranidium, dorsal view, x 8.
Fig. 9. Brongniartella sp. GSM Pg 17, cast from external mould of incomplete pygidium, dorsal view, x 2.
Figs. 10-13. Diacanthaspis stadensis (Reed), X 10. 10, SM A4646b, lectotype (selected Temple 1969,
p. 203), cast from external mould of cranidium, dorsal view. 11, 12, SM A4644a, b, internal mould and
cast from external mould of cranidium, dorsal views. 13, SM A32010, internal mould of small pygidium,
dorsal view.
Figs. 14, 15. cf. Leonaspis girvanensis (Reed), X 10. 14, SM A88536, internal mould of pygidium, dorsal
view. 15, SM A32021, internal mould of partial pygidium, dorsal view.
All specimens from St. Martin’s Cemetery horizon (basal Haverford Mudstone Formation); collected by
V. M. Turnbull. Figs. 1, 2, 4, 6-15 from either of localities D and E on text-fig. 1. Fig. 3 from the Cethings
railway cutting, locality N. Fig. 4 from Little Cuckoo, locality A.
PLATE 82
COCKS and PRICE, Ordovician trilobites
712
PALAEONTOLOGY, VOLUME 18
Fauna. The fauna in Turnbull’s collections was listed by Reed (1906, p. 537). We have
reidentified the trilobites and brachiopods in these collections in addition to those
collected subsequently from all localities at this restricted horizon (an asterisk denotes
species originally described from the horizon) :
Trilobites. Brongniartella sp. (PI. 82, fig. 9), * Diacanthaspis sladensis (Reed, 1905) (PI. 82, figs. 10-13),
cf. Leonaspis girvanensis (Reed, 1914) (PI. 82, figs. 14, 15), Lichasci. laciniatus (Wahlenberg, 1818) (PI. 82,
figs. 1, 2), Mucronaspis mucronata (Brongniart, 1822) (PI. 83, figs. 1-4), Otarion cf. megalops (M’Coy,
1846) (PI. 82, figs. 3-8).
Brachiopods. Lingula sp., Orbiculoidea concentrica (Wahlenberg, 1821) (PI. 83, fig. 5), Philhedra sp.,
Craniops sp., Skenidioides sp., *Giraldiella giraldi Bancroft, 1949, * Dalmanellal biconvexa 'WilWams, 1951,
Dalmanella aff. testudinaria (Dalman, 1828) (PI. 83, fig. 1 1), Hirnantia sagittifera (M’Coy, 1851) (PI. 83, figs.
13, 14) of which *Orthis porcata sladensis Reed, 1905 is a junior synonym, Chonetoidea cf. papillosa (Reed,
1905), *Leptaena martinensis Cocks, 1968 (PI. 83, fig. 6), * Eostropheodonta whittingtoni Bancroft, 1949
(PI. 83, figs. 6, 8) of which * Eostropheodonta hirnantensis delicatula Bancroft, 1949 is a junior synonym,
*Cliftonia lamellosa Williams, 1951 which may be a junior synonym of Cliftonia psittacina (Wahlenberg,
1821) (see Bergstrom, 1968, p. 11), Cryptothyrella crassa (J. de C. Sowerby, 1839) incipiens (Williams, 1951)
(PI. 84, fig. 3).
Diacanthaspis sladensis, together with forms very similar to Otarion cf. megalops
and Lichas cf. laciniatus, is known from limestone bands immediately overlying the
Keisley Limestone of Westmorland (Temple 1969), an horizon which Temple con-
sidered to be lowest Silurian in age, but which Ingham and Wright (in Williams et al.
1972, p. 47) more recently considered to be Hirnantian. The type material of Leonaspis
girvanensis is from the upper Drummuck Group of Girvan which is late Rawtheyan
in age (Ingham 1966, p. 495). In addition, a pygidium (SM A43243) very similar to
that figured here as Plate 82, fig. 14 is known from the 'Mucronatus Beds’ west of
Trout beck in the Lake District, another late Rawtheyan horizon. In what can be
seen of its ornamentation, the Welsh pygidium shows more affinity with these two
forms than with the species from the basal Silurian of Watley Gill, Cautley, mentioned
EXPLANATION OF PLATE 83
Figs. 1-4. Mucronaspis mucronata {Brongniart). 1,2, BM It 13246a, b, internal mould and cast from external
mould of pygidium, dorsal views, x6. 3, BM It 13247, internal mould of cephalon, dorsal view, x4.
4, SM A32020, internal mould of cephalon, dorsal view, x4.
Fig. 5. Orbiculoidea concentrica (Wahlenberg). SM A3 1850a, brachial valve, x2.
Figs. 6, 8. Eostropheodonta whittingtoni Bancroft. 6, SM A30039a, pedicle valve with brachial valve of
Leptaena martinensis Cocks, X 1-4. 8, SM A32035, pedicle valve, x 1-5.
Figs. 7, 9, 12. Eochonetes a&. advena Reed. 7, BM BB 31678-31679, internal moulds of pedicle valves, note
perforated hinge-lines, especially on top left-hand specimen, x 1-3. 9, BM BB 32230, brachial valve,
X 1-5. 12, BM BB 31683, brachial valve, x3.
Fig. 10. Anisopleurella gracilis (Jones). GSM 37555, internal mould of brachial valve, x5.
Fig. 11. Dalmanella aff. testudinaria (Dalman). SM A3 1 899, internal mould of large brachial valve showing
musculature, x 2.
Figs. 13, 14. Hirnantia sagittifera (WPCoy). 13, SM A31908, pedicle valve, X 1-4. 14, SM A31912, brachial
valve, X 2.
Figs. 1-3 and 10 from lower Haverford Mudstones, locality K; 1-3 collected by L. R. M. C., 10 collected
by O. T. Jones. Figs. 4-6, 8, 11, 13, and 14 from St. Martin’s Cemetery horizon (basal Haverford Mud-
stones), 4-6, 1 1, 13, and 14 from either of localities D and E, 8 from locality N; all collected by V. M.
Turnbull. Figs. 7, 9, and 12 from upper Slade and Redhill Mudstones, locality F; collected by L. R. M. C.
PLATE 83
COCKS and PRICE, Ordovician trilobites and brachiopods
714
PALAEONTOLOGY, VOLUME 18
by Temple (1975, p. 158). Similarly, the Welsh material of Mucronaspis mucronata
does not appear to belong to the subspecies M. m. brevispina which at Watley Gill
also occurs in the basal Silurian (Temple 1952). Brongniartella is known in the fauna
only from two pygidia, the better of which is figured here (PI. 81, fig. 1), and one
poor, very small (?holaspid) cranidium (SM A85611). Although what features can
be seen from this material are similar to the upper Ordovician B. platynota (Dalman,
1828), adequate comparison is not really possible.
There are only two graptolite specimens known from the St. Martin’s Cemetery
horizon, one collected by Turnbull from the Cemetery localities and one by Pringle
from the Cethings cutting. Both were identified at Cambridge, the first by Miss Elies
as Diplograptus cf. modestus Lapworth (Reed 1907, p. 537) and the second by her
colleague Mrs. Shakespeare (Miss Wood) as Diplograptus modestusl Lapworth
(Strahan et al. 1914, p. 101). Unfortunately the Turnbull specimen is now lost, but
the Pringle specimen (GSM Pg. 54) has been kindly re-examined for us by Dr. R. B.
Rickards who states that, while the specimen superficially resembles D. modestus,
it is definitely not that form and represents an undescribed diplograptid species. It
seems reasonable to assume that the lost Turnbull specimen was probably of the
same form (particularly in view of its ‘cf.’ identification) so that these graptolites are
at present of little use in any determination of the detailed age of the St. Martin’s
Cemetery horizon. This fact is important since the original "D. modestus' determina-
tion was one of the main palaeontological factors in the decision by Reed and the
Survey Officers to assign the horizon to the Silurian.
The shelly fauna from the St. Martin’s Cemetery horizon we interpret as a Hirnantia
fauna of late Ashgill age; it is further discussed at the end of this paper.
The Haverford Mudstone Formation
Lithostratigraphy. Although in their vertical sections (e.g. Strahan et al. 1914, fig. 12)
the Survey workers distinguished between their ‘Cartlett Beds’ and ‘Gasworks Mud-
stones’, they made no attempt to map these as separate divisions. This is consistent
with the experience of the present authors who have reached the conclusion that it
is only the presence of distinctive shelly faunas at certain localities that enables them
to be classed as developments of Gasworks Mudstones rather than Cartlett Beds;
lithologically there is no clear distinction. Although there is a gradual increase in
the number of thin sandstone beds upwards and also an increase in faunal content,
there is no sharp change that could be used to separate two formations. We therefore
group the Survey’s Cartlett Beds and Gasworks Mudstones together as a single
formation for which we propose the name Haverford Mudstone Formation— taking
the name used by the Survey workers for the lowest stage (what would today be
termed a group) of their Silurian succession.
There is no one section through which the entire formation may be followed.
Many excellent localities, however, have become temporarily available for periods
over the last ten years, including those due to the building of the Haverfordwest
bypass, improvements along the A40 road to the east of the town, and the con-
struction of a series of housing estates to the west of the town. The most important
permanent sections in the formation are the Cethings railway cutting (text-fig. 2),
where the contact with the underlying Portfield Formation is seen (locality N), the
COCKS AND PRICE: ORDOVICIAN AND SILURIAN BIOSTRATIGRAPHY
715
TEXT-FIG. 2. Map of the solid geology of the Haverfordwest area. Lettered localities are those referred to in the text. The Carboniferous outcrop
boundary is taken from the Geological Survey six-inch map (Pembrokeshire 27 NE.) and some of the fossil localities shown are Geological Survey
localities. There are more strike-faults and minor local folds than are shown.
716
PALAEONTOLOGY, VOLUME 18
sections in the railway cuttings south-west of Haverfordwest Station (locality K),
and the classic section running from New Road to opposite the entrance of Haver-
fordwest gasworks (Strahan et al. 1914, pp. 89-91) (locality L), where the contact
with the overlying Gasworks Sandstone Formation is exposed.
The structure of the outcrop area (text-figs. 1 and 2) is complex and the exposures
restricted; there are certainly more strike faults than we have shown on our maps,
as well as many minor local folds.
Fauna. The Haverford Mudstone Formation is estimated to be between 350 and
390 m in thickness and is divided faunally into three parts. The basal metre includes
the St. Martin’s Cemetery horizon which carries a Hirnantia fauna, the next 210 to
250 m carries a sparse fauna discussed further below, and the uppermost 140 m con-
tains a rich shelly fauna of typical lower Llandovery (Rhuddanian) aspect, best
known through the classic exposures opposite the gasworks at Haverfordwest.
The fauna of the St. Martin’s Cemetery horizon has been discussed above. The next
division of the formation is largely barren of macrofauna but it has yielded fossils in
four areas :
(i) 9 m above the base of the formation in the Cethings railway cutting a single
graptolite was collected by Strahan et al. (1914, p. 89) and identified by them as
IDiplograptus modestus parvulus (H. Lapworth). Dr. R. B. Rickards reidentifies the
specimen as IClimacograptus normalis Lapworth, although he reports a hint of
apertural spines recalling the C. innotatus group. In either case the stratigraphical
significance is doubtful.
(ii) A series of exposures in the railway cutting south of Haverfordwest (locality K,
Grid Ref. SM 955 146), where dark-green micaceous mudstones have yielded a shelly
fauna dominated (80% in one collection, n= 103) by the small brachiopod Aniso-
pleurella gracilis (Jones, 1925) (PI. 83, fig. 10; Cocks 1970, pi. 16, figs. 1-9), with
subsidiary Mucronaspis mucronata{V\. 83, figs. 1-4), Leptaena, Eoplectodonta, Lingula,
EXPLANATION OF PLATE 84
Figs. 1, 2, 5, 6. Hirnantia sagittifera (M’Coy). 1,2, BM BB 68722 and BM BB 68709, internal moulds of large
brachial valves, x 2-5. 5, BM BB 38699, pedicle valve, x 2. 6, BM BB 68720, internal mould of small
brachial valve, x 2.
Fig. 3. CryptothyreUa crassa incipiens (Williams). SM A3 1930, internal mould of conjoined valves, posterior
view, X 2.
Figs. 4, 7. Eostropheodonta whittingtoni Bancroft. 4, SM A30041a, brachial valve, x 1-4. 7, SM A31884,
pedicle valve, x 1-5.
Figs. 8, 10. (M’Coy). 8, BM BB 38634, brachial valve, x 1-5. 10, BM BB 38660,
pedicle valve together with Hirnantia sagittifera brachial valve, x 1 -7.
Figs. 9, 13. Stricklandia lens lens (i . deC. Sowerby), brachial valves. 9, BM BB 69607, X 2. 13, BM BB 69608,
X 1-5.
Figs. 11,12. Katastrophomenaai{.scotica(^'dncroi\). 1 1, BM BB69610, pedicle valve, X 2. 12, BM BB 69609,
brachial valve, x 1 -6.
Figs. 1, 2, 5, 6, 8, and 10 from Flirnant Quarry, south-east of Bala, Merionethshire (Gwynedd), Grid Ref.
SH 945 285; collected by L. R. M. C. Figs. 3, 4, 7 from St. Martin’s Cemetery horizon (basal Haverford
Mudstones), localities D and E; collected by V. M. Turnbull. Figs. 9, 13 from upper Haverford Mud-
stones, locality J; collected by L. R. M. C. Figs. 11,12 from upper Haverford Mudstones, locality H;
collected by L. R. M. C.
PLATE 84
COCKS and PRICE, Ordovician and Silurian brachiopods
718
PALAEONTOLOGY, VOLUME 18
Eospirigerina, and two undetermined species of enteletacean brachiopod. Un-
fortunately the exact stratigraphical horizon of this locality within the formation
is difficult to determine, since it lies on the southern limb of a syncline when com-
pared with other localities (text-fig. 2).
(iii) Several temporary exposures in the foundations of new houses in the area
around Portfield House (text-fig. 1). At one locality (locality B, Grid Ref. SM
9375 1550) Anisopleurella gracilis occurred in some quantity, and 60 m north of this
(SM 9374 1556, locality C) an indeterminate graptolite fragment. The former locality
is an estimated 45 m from the base of the formation, the latter about 20 m.
(iv) A series of exposures, mainly temporary, along the main A40 road just north
of Bethany Farm (text-fig. 2). At locality P (Grid Ref. SM 9653 1590) A. gracilis
occurred as many specimens on a single bedding plane and Orbiculoidea, Eospiri-
gerina, Leptaena, Eoplectodonta, Skenidioides, and Resserella (BM BB 70622-70631)
as scattered single specimens. This is near a locality (O on text-fig. 2) termed EA 23 by
the Survey, which yielded to them, as well as A. gracilis, a graptolite (GSM TCC
1876/7) which Dr. Rickards identifies as Climacograptus cf. normalis Lapworth.
From a stratigraphically slightly higher horizon (locality Q, Grid Ref. SM 9661 1589),
no longer exposed, the Survey (their locality EA 24) recovered single specimens of
Cryptothyrella, Leangella scissa (Davidson, 1871), Eospirigerina, IClorinda, Eostro-
pheodonta, and "I Anisopleurella and three specimens of Resserella, a fauna which,
although not fully diagnostic, suggests for the first time in the formation a positive
Llandovery, rather than an Ashgill, age. Within 1 5 m above this (locality R, Survey loc.
EA 28, Grid Ref. SM 9666 1591) a typical Llandovery fauna, including Stricklandia,
occurs.
Thus the age of this central division of the Haverford Mudstone Formation remains
equivocal. Below it there is a definite Hirnantian horizon and above it a definite
Rhuddanian horizon. Where within this division the boundary limit between these
Stages, and hence between the Ordovician and Silurian, should be drawn is uncertain.
There seems a good case for putting the highest 15 m of the division (locality Q and
above) into the Rhuddanian, but the underlying 235 m with its sparse Anisopleurella
fauna and Mucronaspis mucronata could just as well be Hirnantian; we do not regard
the presence of Mucronaspis as being necessarily diagnostic in this respect. It is hoped
that more graptolites may be recovered from these horizons in the future, although
even graptolite faunas near the Ordovician-Silurian boundary still await a defini-
tive study.
The fauna of the uppermost 140 m of the formation is one of the richest in the
Rhuddanian of Britain. Temple (1975) has reviewed the trilobites and redescribes
Calymene crassa (Shirley, 1936), Calymene sp. A, Brongniartella sp., Hadromeros
elongatus (Reed, 1931), Acernaspis sp., Dalmanites sp., Stenopareia sp., and an
indeterminable odontopleurine. The faunas, however, are numerically dominated
by brachiopods, of which only the strophomenides have yet been redescribed (Cocks
1968, 1970), although many of the forms described by Temple (1970) from Meifod
also occur in the Haverfordwest area. The chief localities are;
(i) The lane leading from New Road to the gasworks, Haverfordwest (locality L,
Grid Ref. SM 9622 1500 to SM 9582 1537; Strahan et al. 1914, pp. 90-91).
COCKS AND PRICE: ORDOVICIAN AND SILURIAN BIOSTRATIGRAPHY 719
(ii) Along the path known as Fortune’s Frolic (text-fig. 2) on the east bank of the
Western Cleddau (Grid Ref. SM 9622 1500 to SM 9647 1461; Strahan et al. 1914,
pp. 92-96), where the succession is repeated by local folding and faulting.
(iii) The area around Priory Mill (Grid Ref. SM 959 149), including some large
temporary exposures made during the construction of the southern bypass in 1974,
from locality J (SM 9540 1478) eastwards.
(iv) The area around Merlin’s Bridge (text-fig. 2) where there are several isolated
exposures. Jones {in Strahan et al. 1914, pp. 96-99) was concerned that the faunal
aspect of these ' Pentamerus undatus beds’, as he called them, was not the same as
that of those opposite the gasworks entrance. We have examined the material
collected by the Survey and made new collections from several localities, in par-
ticular an old quarry west of Merlin’s Bridge (locality H, Grid Ref. SM 946 146),
and there is no doubt that the exposures fall within the uppermost part of our Haver-
ford Mudstone Formation.
There are three different main brachiopod assemblages present in the upper
division :
(i) Faunas dominated by Clorinda undata (J. de C. Sowerby) and Eoplectodonta
duplicata (J. de C. Sowerby), and typical of most of the exposures in the Merlin’s
Bridge area. A collection from locality H yielded Clorinda undata (23%), Eoplecto-
donta duplicata (18%), Resserella llandoveriana Williams (10%), Skenidioides wood-
landiense (Davidson) (9%), nine other species of brachiopod, and eight species of
other groups (n = 122).
(ii) Faunas dominated by Stricklandia lens, very often in nearly monospecific
assemblages covering single bedding planes, with some specimens even in position
of growth (Ziegler et al. 1966). The subspecies present is chiefly S. lens lens (J. de C.
Sowerby), but some populations and individuals are morphologically closer to S. 1.
prima Williams, whose type horizon and locality (Williams 1951) is A2_3 Beds at
Llandovery itself, and which is the earliest Silurian stricklandiid form. These Strick-
/a«(i/a-dominated assemblages are widespread, extending well outside the area to
such localities as that north of Woodford Cottage, south of Robeston Wathen (Grid
Ref. SN 0848 1504).
(iii) Diverse faunas without common pentamerids, the best examples coming from
the exposures opposite the entrance to the gasworks at Haverfordwest (locality L).
In a large collection from a band approximately 14 m below the top of the formation,
brachiopods dominated the assemblage, with twenty different species, including
Eoplectodonta duplicata (18%), Resserella llandoveriana Williams (9%), Leangella
scissa (8%), Eopholidostrophia sefinensis ellisae Hurst (5%), Eostrophonella eothen
Bancroft (3%), and Schizonema sowerbyiana Davidson (2%) (n = 658), but the
collection also contained at least seventeen species of other phyla, including a thick
stick bryozoan, possibly Hallopora (15%), Tentaculites (10%), the dasyclad alga
Mastopora fava Salter (5%), a thin stick bryozoan (4%), and a compound bryozoan
(3%), one species of orthoceratid, three species of gastropod, one of bivalve, two
species of coral, much crinoid debris, and four different trilobites {Calymene crassa,
an odontopleurine— probably Leonaspis, a phacopid, and an encrinurid). Another,
much smaller, collection from the railway cutting 40 m NE. of the railway bridge
720
PALAEONTOLOGY, VOLUME 18
(Grid Ref. SM9579 1551) was from a single thin band and was very much less diverse,
being dominated by Katastrophomena scotica (Bancroft) (59%), with the next most
common taxa each at 7% (Resserella Uandoveriana and a bryozoan). A notable
feature of this faunal group is the variety of assemblages encountered, with particular
abundances of groups other than brachiopods, and an unusual absence of penta-
merids.
The interpretation of the first two of these upper division assemblages is that they
can be identified with the Clorinda and Stricklandia Communities defined in the
upper Llandovery of the Welsh Borderland (Ziegler et al. 1968). However, the
interpretation of the third group, which is both varied and diverse, presents more of
a problem. Perhaps the most likely solution is that assemblages are represented which
ecologically parallel a typical Clorinda Community, but from an environment locally
unsuited to Clorinda itself or other pentamerids; the abundance of bryozoa and
Mastopora are also unusual. Whether or not the deposition was unusual is uncertain.
Sanzen-Baker (1972, p. 1 52) suggests that the upper parts of the Haverford Formation
were partly deposited from turbidity currents, and the jumbled nature of many
of the third assemblage occurrences would be explained by this. On the other hand,
the fact that many of the brachiopods are found with conjoined valves, and that
the fragile calcareous algae are found at all, implies that the amount of pre-depositional
disruption was not great. In any case a Rhuddanian age for the upper part of the
Haverford Mudstone Formation is quite certain.
Gasworks Sandstone and Millin Mudstone Formations
Lithostratigraphy. Detailed discussion of these later formations is outside the scope
of this paper. Sanzen-Baker (1972) has described the turbidites of the Gasworks
Sandstone Formation, which succeed the Haverford Mudstone Formation apparently
conformably. Above the Gasworks Sandstone there are thick mudstones which
Strahan et al. (1914) divided into lower Uzmaston Beds and upper Canaston Beds.
The Survey workers were not able to map these two divisions separately, and the
present authors treat them as one formation, the Millin Mudstone Formation, whose
name is taken from the Survey’s ‘Series’ name.
Fauna. Fossils, chiefly brachiopods, occur sporadically within the Gasworks Sand-
stone Formation, but their age is not diagnostic. The Millin Mudstone Formation
is more fossiliferous, yielding late Llandovery faunas chiefly representing the Clorinda
Community. These beds may be dated by stricklandiids and Eocoelia to a variety of
ages within the Fronian and Telychian stages; a representative locality occurring
on our map (locality M, Survey loc. FA 25, Grid Ref. SM 9620 1525) included
Anthirhynchonella linguifera (J. de C. Sowerby), Eoplectodonta penkillensis (Reed),
Clorinda undata (J. de C. Sowerby), Leptaena purpurea Cocks, Atrypa reticularis
(Linnaeus), and Eocoelia curtisi Ziegler, indicating a Telychian age at approximately
90 m above the base of the Millin Mudstone Formation.
THE HIRNANTIA FAUNA
For many years a fauna has been known, typified by the association of the enteletacean
Hirnantia sagittifera (M’Coy) and the strophomenide Eostropheodonta hirnantensis
COCKS AND PRICE: ORDOVICIAN AND SILURIAN BIOSTRATIGRAPHY 721
(M’Coy), from near the Ordovician-Silurian boundary. The type area is at Aber
Hirnant, south-east of Bala, North Wales, where the beds were often regarded as
post-Ashgill (Elies 1922; Bancroft 1933, p. 4), although more recently these beds at
Hirnant were revised to form the type Hirnantian, the highest stage within the Ashgill
Series (Ingham and Wright 1970). It is perhaps unfortunate that the name of the
fauna and the name of the stratigraphical time division should be so similar ; although
this misfortune has since been compounded into error, for example by Lesperance
(1974), who even explicitly equates the two terms, resulting in an erroneous hybrid
‘Hirnantian fauna’, which Lesperance identifies, as if by definition, with the Hirnantian
time horizon (Stage).
The beds around Aber Hirnant await modern redescription, but a collection from
Hirnant Quarry yielded the following seven brachiopods: E. hirnantensis (30%,
n=153), H. sagittifera (21%), Plectothyrella crassicosta (Dalman, 1828) (10%),
Kinnella kielanae (Temple, 1965) (9%), Dalmanella sp. (9%), Bancroftinal bouceki
(Havlicek, 1950) (8%), and Skenidioides sp. (1%). The only representatives of other
phyla were a few crinoid ossicles, some compound bryozoa, and some borings,
presumably of sponges, into the brachiopod shells. These data are comparable with
the percentages (of brachiopods only) from the same locality recorded by Temple
(1965, p. 419), and consist of the same species, with the addition only of the uncommon
Skenidioides. The lithology of Hirnant Quarry is variable, but includes a pisolitic
oolite not seen elsewhere.
This Hirnantia fauna is very comparable in composition and diversity with others
from near Llangollen, North Wales, and from Hoi Beck (Temple 1965, p. 418) and
Cautley (Wright 1968, p. 361), both in northern England. These Hirnantia faunas,
however, are less diverse than others which possess the same basic elements but with
the addition of other taxa. One such is from the St. Martin’s Cemetery horizon,
described above. Another, from Stawy, Poland, contains three inarticulate and seven
articulate species of brachiopod as well as ‘trilobites, ostracods, bryozoans, worm
tubes, a hyolithid and crinoid and graptolite fragments’ (Temple 1965, p. 380).
A further fauna, from the Kildare Limestone, Ireland (Wright 1968) consisted of
Cyptothyrella crassa incipiens (35%), Cliftonia oxoplecioides (28%), Plectothyrella
platystrophoides (19%), dalmanellids (including Dalmanella) (12%), cf. Leptaenopoma
(4%), H. sagittifera (3%), and Eostropheodonta sp. (less than 1%). Rare trilobites
included Dalmanitina sp. and an odontopleurid. It is of great interest that Wright
records that these Kildare Beds, which include the Hirnantia fauna elements, are
actually interbedded with reef limestones with a rich and diverse fauna including
Cliftonia, Streptis, Triplesia, Anisopleurella, Leptaena, and Christiania, as well as
Sphaeroxochus, illanenids and other trilobites, ostracodes, byrozoans, and algae;
and it is also interesting to note the records of fragmentary trinucleids from beds
above those with the Hirnantia fauna.
Bergstrom (1968) gives valuable data on the brachiopod fauna of the Dalmanitina
Beds in Vastergotland, Sweden. Of the eighteen localities from which he records
brachiopods, seven (his fig. 4 locality nos. 1, 3, 6-7, 8, 10, 14, and 28) are typical,
fairly restricted, Hirnantia assemblages, four (his localities 18 and 22-24) are not
Hirnantia assemblages, and the rest are diverse assemblages which include Hirnantia
fauna elements to greater or lesser extents. His most prolific exposure (his locality 5)
722
PALAEONTOLOGY, VOLUME 18
yielded an interesting assemblage consisting of (percentages calculated from his
fig. 4 of brachiopods only) ; Coolinia dalmani (43%) (n = 625), H. sagittifera (9%),
K. kielanae (9%), E. hirnantensis (7%), P. crassicosta (6%), Cliftonia psittacina (5%^
Horderleyella fragilis (4%), Aphanomena schmalenseei (4%), Leptaenopoma trifidum
(3%), Leptaena rugosa (3%), Orbiculoidea concentrica (2%), Drabovia westrogothica
(1%), Giraldiella bella (1%), Dalmanella testudinaria (1%), Draborthis caelebs (1%),
Titanomena grandis (1%), and rare Dalmanella pectinoides and Petrocrania aperta,
a total of eighteen brachiopod species, which is quite diverse. Bergstrom also records
(1968, p. 5) Hirnantia fauna elements from Jamtland, northern Sweden, occurring
in the same beds as Dalmanitina mucronata, Brongniartella platynota, and also
Tr etas pis.
Marek and Havlicek (1967) described comparable assemblages from the Kosov
Formation of Bohemia in which Hirnantia fauna elements— Kinnella,
Dalmanella testudinaria, Cliftonia, E. hirnanentensis, Leptaena rugosa, Cryptothyrella,
and Plectothyrella occur with other {oxm?,— Giraldiella subsilurica, Comatopoma
sororia, Drabovia agnata, Draborthis caelebs, Onniella rava, Aegiromena ultima,
Rafinesquina urbicola, R. ultrix, Leptaenopoma trifidum, Bracteoleptaena polonica,
Eardenia comes, and Zygospira fallax. Although a few of these latter species have
been recorded with Hirnantia fauna elements elsewhere (e.g. Bergstrom 1968), most
of them are not found in the strict Hirnantia fauna at all.
Havlicek (1971) also described a comparable fauna from the Deuxieme Bani of the
Anti-Atlas, Morocco. Here, in addition to Hirnantia, Eostropheodonta, and Plecto-
thyrella, he described another ten brachiopod species not so far recognized elsewhere.
Hirnantia alf. sagittifera and endemic species of Plectothyrella have also been
described from the Memouniat Formation of Libya (Havlicek and Massa 1973).
Lesperance (1974, tables 1 and 2) reports a fauna from Perce, Canada, consisting
of Brongniartella sp., M. mucronata, M. olini, Philipsinella parabola, two other
endemic trilobites, Dalmanellal, E. hirnantensis, Hirnantia^}, Kinnella kielanae, and
Plectothyrella, but he does not give relative abundances, or state whether the fauna
was all collected from the same horizon. From the Portage River area, 17 km away,
Lesperance records a fauna (without brachiopods) in which the trilobites M. mucro-
nata, M. olini, Brongniartella, Portaginus, and Cryptolithus occur together with what
he records as Climacograptus rectangularismedius, the latter suggesting an early
Silurian age. However, Dr. Rickards (pers. comm.) believes that these graptolites
probably represent an earlier climacograptid stock, perhaps evolved from C. normalis,
and that their age, although uncertain, is more probably late Ordovician.
The situation in Kazakhstan, U.S.S.R., requires further clarification: Nikitin
(1971, p. 339) records M. mucronata and M. olini from the upper Tolen Beds in
association with Glyptograptus persculptus, but a Hirnantia fauna as such does not
seem to be present, the only brachiopod recorded being "Conchidium’’ munsteri.
Thus the Hirnantia fauna assemblages are widely variable; sometimes restricted
to a mere two genera, at other times consisting of the basic half-dozen Hirnantia
fauna elements with up to twelve associated cosmopolitan or endemic brachiopods.
Sometimes no trilobites are present; at other times one or more representatives of
the ‘'Dalmanitina' fauna with or without other cosmopolitan or endemic forms. We
believe that the Hirnantia fauna is best interpreted as representing an animal com-
COCKS AND PRICE: ORDOVICIAN AND SILURIAN BIOSTRATIGRAPHY 723
munity comparable with those described by Ziegler et al. (1968), rather than as a
single time assemblage zone, and that its occurrence is therefore a priori as likely
to be diachronous as synchronous. As Wright (1968, p. 365) suggested, the Hirnantia
Community, at least in its restricted form, suggests original deposition under relatively
shallow water; but perhaps the more diverse Hirnantia assemblages could reflect
deposition under slightly deeper water.
Spectacular evidence for a late Ordovician-early Silurian glacial event has been
accumulating from many parts of the world for some years, and this event is probably
connected with the faunal changes seen across the Ordovician/Silurian boundary.
Obviously near the then poles this glacial event would have been more prolonged
than in equatorial regions, but to what extent the Hirnantia Community is a direct
reflection of cold-water conditions is as yet uncertain. In the same way the degree
to which this glaciation coincides with the extent of the Hirnantian Stage remains
unknown.
The age range of the Hirnantia Community also remains uncertain. Eostropheo-
donta ranges from Cautleyan up to the Wenlock, Hirnantia itself is known from the
early Ashgill to late Llandovery (Walmsley et al. 1969, p. 515), Cryptothyrella from
the early Ashgill to the Wenlock, Dalmanella s.s. from the Caradoc to the Llandovery,
and Plectothyrella from the Ashgill to the Llandovery. Of the typical Hirnantia
Community forms only Kinnella appears to be confined to the late Ashgill, and that
enteletacean has only been recognized comparatively recently (Bergstrom 1968) as
a separate genus; its range is not definitively known. Sometimes the Hirnantia Com-
munity occurs below tretaspid trilobites, at other times in the same beds, and at yet
other times apparently later than the last local tretaspid fauna. In a similar fashion
the ranges of the trilobites that make up the " Dalmanitina" fauna also vary and their
occurrences should be assessed separately from those of the whole Hirnantia Com-
munity. Very often the two occur together in a single assemblage, but at other times
each is found separately. Thus to conclude that the Hirnantia Community occurrences
are all of the same age appears to us to be a dangerous assumption.
Acknowledgements. We are grateful to Dr. R. B. Rickards for identifying our graptolites and for re-examining
old specimens, and to Dr. J. T. Temple for discussion and for access to unpublished material. We are also
most grateful to Dr. A. W. A. Rushton for access to old Geological Survey collections and data.
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PALAEONTOLOGY, VOLUME 18
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Early Old Red Sandstone of Pembrokeshire. Proc. Geol. 83, 139-164.
STRAHAN, A., CANTRiLL, T. c., DIXON, E. E. L., THOMAS, H. H. and JONES, o. T. 1914. The geology of the South
Wales Coalfield, Part XL The country around Haverfordwest. Mem. geol. Surv. U.K. 228, 1-262.
TEMPLE, J. T. 1952. A revision of the trilobite Dalmanitina mucronata (Brongniart) and related species.
Lunds. Univ. Arssk. N.s. (2), 48, 1-33, pis. 1-4.
1965. Upper Ordovician brachiopods from Poland and Britain. Acta palaeont. Pol. 10, 379-450,
pis. 1-21.
1969. Lower Llandovery (Silurian) trilobites from Keisley, Westmorland. Bull. Br. Mus. nat. Hist.
(Geol), 18, 199-230, pis. 1-6.
1970. The Lower Llandovery brachiopods and trilobites from Ffridd Mathrafal, near Meifod,
Montgomeryshire. Palaeontogr. Soc. [Monogr.], 1-76, pis. 1-19.
1975. Early Llandovery trilobites from Wales with notes on British Llandovery calymenids. Palaeon-
tology, 18, 137-159, pis. 15-21 .
WALMSLEY, V. G., BOUCOT, A. J. and JOHNSON, J. G. 1969. Silurian and lower Devonian salopinid brachiopods.
J. Paleont. 43, 492-516, pis. 71-80.
WILLIAMS, A. 1951. Llandovery brachiopods from Wales with special reference to the Llandovery District.
Q. Jl geol. Soc. Lond. 107, 85-136, pis. 3-8.
STRACHAN, L, BASSETT, D. A., DEAN, W. T., INGHAM, J. K., WRIGHT, A. D. and WHITTINGTON, H. B. 1972.
A correlation of Ordovician rocks in the British Isles. Spec. Kept. Geol. Soc. Lond. 3, 1 -74.
WRIGHT, A. D. 1968. A westward extension of the Upper Ashgillian Hirnantia Eauna. Lethaia, 1, 352-367.
ZIEGLER, A. M., BOUCOT, A. J. and SHELDON, R. p. 1966. Silurian pentameroid brachiopods preserved in
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Typescript received 21 January 1975
Revised typescript received 23 April 1975
L. R. M. COCKS
Department of Palaeontology
British Museum (Natural History)
London, SW7 5BD
D. PRICE
Department of Geology
Sedgwick Museum
Downing Street
Cambridge, CB2 3EQ
COMPARATIVE ANALYSIS OF FOSSIL AND
RECENT ECHINOID BIOEROSION
by R. G. BROMLEY
Abstract. One of the most abundant forms of bioerosion sculpture on Mesozoic and Cainozoic shells and other
hard substrates has a pentaradiate symmetry based on a regular, stellate module consisting of five radiating grooves.
Regular echinoids today, browsing on encrusting and boring organisms on hard substrates, produce identical sculp-
ture to the trace fossil, and a common origin is suggested. The tooth scratches lose their pentaradiate orientation and
become subparallel where the echinoid gnaws along edges of shells and flat pebbles; a corresponding sculpture is
also encountered in the trace fossil. The pentaradiate trace fossil is designated as Gnathichnus pentax, ichnogen. et
ichnosp. nov.
A TYPE of hard substrate trace fossil that, despite its abundance, has received scant
attention from geologists, is the sculpture produced by browsing and foraging
animals. Such sculpture, commonly termed ‘bioerosion’ (Neumann 1966), occurs
on substrate surfaces that have been exposed at the sea floor for a period of time.
Indeed, in some post-Palaeozoic sediments of shallow marine origin it is difficult to
find fossils unaffected by superficial bioerosion.
One of the most common types of bioerosion sculpture has a basic pentaradiate
symmetry and comparison with the traces produced by the teeth of living regular
echinoids strongly suggests a common origin.
DESCRIPTION OF FOSSIL MATERIAL
The surface sculpture of bioeroded shells and other substrates may consist of random
or grouped scratches of various depths and sizes. Examples have been described by
Abel (1935), Boekschoten (1966, 1967), Riegraf (1973), Bishop (1975), and many
others. Among these different types of scratches there is a single group that can be
treated separately owing to its distinctive pentaradiate organization and regular
morphology. The aspect presented by this type depends on (1) the spacing between
the scratches, (2) the curvature of the substrate, and (3) the presence of borings in
the substrate.
More or less flat surfaces
In its simplest form the trace fossil consists of a regular stellate arrangement of
five radiating grooves making an angle of approximately 72° with each other. Such
a ‘star’ is rarely seen in isolation but may be regarded as the ‘modular unit’ (Heinberg
1973) of the ultimate bioerosion sculpture. The ‘rays’ of this stellate module have
a uniform length and depth. The diameter of the module rarely exceeds 5 mm and
in most cases is less than 2 mm.
Considerably more common than single stars are compound traces built up of
a series of overlapping identical stars in which equivalent rays are repeated parallel
[Palaeontology, Vol. 18, Part 4, 1975, pp. 725-739, pis. 85-89.]
726
PALAEONTOLOGY, VOLUME 18
to one another (PI. 85, fig. 1). In those cases where repeated rays lie closer together
than they do to other rays of the same star, the closely spaced group of parallel grooves
representing the repetition of a single ray may be erroneously interpreted as the
module (as e.g. McKerrow et al. 1969, pi. 12, fig. 3) rather than the pentaradiate star
itself.
Compound stellate bioerosion sculptures may become increasingly complex by
repetition of overlapping stars until the individual stars can no longer be distinguished
and the substrate surface is completely covered with grooves of similar depth and
length (PI. 86, figs. 4-5). Nevertheless, plotting of the orientation of the grooves
over selected areas of the surface reveals the underlying stellate module. Indeed, the
pentaradiate distribution of the grooves is so pronounced in most cases that it can
be discerned with the naked eye, and accentuated by oblique illumination.
Sharply curved and irregular surfaces
The full development of the trace fossil as described above is seen only on flat
substrates (e.g. smooth mollusc shells). Where the surface is irregular, the stellate
module is commonly reduced to fewer rays, but the 72° angle between rays remains
constant (PI. 85, fig. 3; PI. 89, fig. 1). In cases where the substrate describes a sudden
and extensive convex curve, as at the edge of a flat pebble, the basic module changes
so that the grooves become densely packed and more or less parallel (PI. 86, fig. 2).
Where perforations occur in the substrate which exceed the diameter of the basic
module, these are commonly surrounded by a series of closely packed grooves in
this case aligned more or less perpendicular or normal to the edge of the hole. The
dimensions of the individual grooves are precisely similar to those of the pentaradiate
sculptures on related flat surfaces.
Presence of sessile organisms within and upon the substrate
Concentration of the grooves also occurs around borings and encrusting skeletons
that are smaller than the diameter of the module. In contrast to the subparallel
orientation of the grooves around large-scale perforations in the substrate, the
grooves around smaller sites exhibit the characteristic regular pentaradiate arrange-
ment. This relationship is particularly noticeable in the case of borings of organisms
in the substrate, where the grooves are generally deep and associated with breakage
of the rim of the boring. Well-developed stars are seen around acrothoracic barnacle
EXPLANATION OF PLATE 85
Fig. 1. Gnathichnus pentax on a belemnite from Reutlingen, near Tubingen, West Germany. Upper Lias
delta. Printed from a peel. x9.
Fig. 2. Bite traces of a Recent echinoid on gastropod shell, Charonia tritonis. Locality unknown. MMH
13387, Mineralogisk Museum, Copenhagen. x7.
Fig. 3. Gnathichnus pentax (holotype) on an oyster shell from lowermost Pleistocene at Kritika, Rhodes.
MMH 13386. x8.
Fig. 4. Bite traces of Sphaerechinus granularis on a flat, algal-coated limestone pebble. The traces were
made in an aquarium at the Institut Rudjer Boskovic-Zagreb, Centre for Marine Research at Rovinj,
Yugoslavia. The specimen is housed in the North Adriatic Collection of the Geologisch-Palaontolo-
gisches Institut, Gottingen University. x9.
PLATE 85
BROMLEY, echinoid bite traces
728
PALAEONTOLOGY, VOLUME 18
borings (PI. 88, figs. 1-3), borings of small worms (PI. 86, fig. 3; PI. 87, fig. 6), and
the papillar orifices to clionid sponge borings. A particularly close association is
found in the last case, and commonly a large proportion of the papillar orifices are
surrounded by multiple stellate modules (PI. 87, figs. 1-5). Other comparable
inhomogeneities of the substrate can also be surrounded by stellate groove patterns,
such as the ambulacral pores of echinoids (PI. 88, figs. 4-5).
GNAWING TRACES PRODUCED BY PRESENT-DAY MARINE ANIMALS
Several groups of animals cause bioerosion through their eating activities. In many
of these groups the process is chiefly one of ‘biting’ and ‘crushing’ and is caused by
biramous tools, i.e. a pair of opposable jaws or claws, as in, for example, parrot fish
and grapsid crabs. These animals work over hard substrates, breaking up the surface,
especially around borings, in search of epilithic and endolithic organisms or eroding
living substrate such as coral (Bakus 1964, 1966). The result of such foraging is
generally a series of highly irregular scratches, pits, and broken protuberances that
in no way resembles the uniform, pentaradiate traces discussed here (PI. 89, fig. 3)
(Abel 1935, p. 325). In certain circumstances, however, fish can produce groups of
subparallel scratches at breakage cavities (Kier and Grant 1965, p. 55).
A more homogeneous form of bioerosion is produced by the methodical grazing
of gastropods and polyplacophores. Boekschoten (1966, fig. 11) illustrated typical
chiton rasping traces, while those of grazing gastropods have received more attention
(see particularly Abel 1935, fig. 338 ; Ankel 1929, 1936, 1937). In these cases, extensive
areas of substrate are eroded, but the rather even scratches have a subparallel distribu-
tion within arcuate groups that reflect the swinging movement of the head as the
gastropod progresses slowly over the substrate, and scratches of the individual teeth
of the radula may be preserved (Ankel 1936, fig. 8, 1937, figs. 2 and 11). Predatory
boring gastropods may scratch the shell surface of their prey with their radula before
boring, but these scratches have random orientation and very local distribution
(Carriker 1969, figs. 11-15). This radular erosion does not produce a sculpture of
pentaradiate module.
EXPLANATION OF PLATE 86
Fig. 1 . Gnawing traces of Sphaerechinus granularis on the rounded edge of an algal-coated limestone pebble.
Details as for Plate 85, fig. 4. x 3.
Fig. 2. Gnawing traces on the edge of a shell fragment (Inoceramus sp.) from Lower Maastrichtian white
chalk, Dronningestolen, M^ns Klint, Denmark. MMH 13388. x6.
Fig. 3. Gnalhichnus pentax along a worm boring, the roof of which has been largely broken away. Upper
Campanian, Bluffport Marl Member, Demopolis chalk, north of Parker, Alabama, U.S.A. MMH
13389. X 6. See also Plate 87, fig. 6.
Fig. 4. Surface of an oyster {Arctostrea dilmiana) entirely sculptured with Gnalhichnus pentax. Uppermost
Lower Campanian calcarenite, Ivo Klack, Ivb, Scania, Sweden. MMH 13390. x6.
Fig. 5. Enlargement of part of fig. 4. x 20.
PLATE 86
BROMLEY, echinoid bite traces
730
PALAEONTOLOGY, VOLUME 18
Echinoid-gnawing traces
The work of one particular group of organisms, however, is highly distinctive and
provides an excellent model for the pentaradiate trace fossils, namely the browsing
traces of regular echinoids. A considerable part of the body of these animals is taken
up by the jaw apparatus, the so-called ‘Aristotle’s Lantern’, the five teeth of which
can be brought to bear on the substrate with considerable power through the muscula-
ture of the lantern and the concerted effort of the numerous tube feet. Regular
echinoids are highly efficient bioeroders and cause considerable rock destruction in
their quest for food. Umbgrove (1939) described intertidal notch erosion in coral
rock chiefly by Echinometra mathaei in the East Indies (text-fig. 1) and Neumann
(1966) attributed extensive notch erosion in limestone immediately below spring
low tide in Bermuda chiefly to the work of Lytechinus variegatus. Neumann (1965)
also emphasized the production of quantities of fine rock powder by the bioerosive
TEXT-FIG. 1. A ‘toadstool island’ from the north coast of
Batoe Daka, Togian Islands, Celebes, sketched after Umb-
grove (1939, fig. 21). The intertidal notch is chiefly eroded
by Echinometra mathaei.
TEXT-FIG. 2. Echinus esculentus clears a
browsing path about as wide as its own
test by meandering over the substrate.
Sketched from a photograph of an animal
in the natural environment on the sea
floor at Heligoland, West Germany. Inset
photograph of gnawing traces of this
species in an aquarium. Both from W. E.
Krumbein, pers. comm. Meandering line
from Krumbein and Van der Pers (1974,
fig. 6a).
BROMLEY: ECHINOID BIOEROSION
731
activities of Lytechinus variegatus. In many rocky shores echinoids use their teeth
to bore deep protective cavities, commonly with narrow entrances, in granite, lime-
stone, and artificial substrates (Market and Maier 1967 ; references in Bromley 1970).
The prime function of the great jaw apparatus is for the exploitation of organisms
encrusting and boring into hard substrates. A study of the feeding habits of regular
echinoids reveals many features that render these animals likely candidates for the
originators of the pentaradiate trace fossils.
The mode of employment of the echinoid tooth involves a powerful scraping action,
producing a single groove that has, on an even substrate, a characteristic and uniform
width and depth. The simultaneous action of all five teeth produces a stellate pattern
of grooves identical in form to the module of the trace fossil (PI. 85, fig. 2). This trace
has been described by many workers.
Krumbach (1914) stated that the scratch produced by Sphaerechinus granularis on
limestone in the Adriatic Sea was up to 0-5 mm deep (PI. 85, fig. 4; see also Abel 1935,
fig. 310). Krumbach observed that each bite in this species took 30-35 seconds at
a temperature of 10°C— a little faster at higher temperatures— and comparable
speeds have been recorded for other species (e.g. Milligan 1916). Between individual
bites, i.e. during the time required to reopen the jaws, the echinoid will have travelled
a certain distance over the substrate, so that at the next bite the five teeth cut fresh
substrate adjacent to the first bite. In this way successive stars are scratched side by
side. Krumbein and Van der Pers (1974) observed that as the browsing echinoid pro-
gresses over the substrate it follows a regularly meandering course and covers a strip
of substrate approximately the same width as the animal’s test (text-fig. 2). Owing
to the non-cephalization of regular echinoids, the orientation of the body and its
tooth apparatus remains more or less unaltered as the animal wanders, and the stellate
grooves of the browsed area consequently show a corresponding constant alignment.
This constancy of orientation is also characteristic of the trace fossil (PI. 85, figs.
3 and 4).
There has been much speculation over the food preferences of regular echinoids
and conflicting evidence has been recorded in the literature. Most species, however,
appear to take advantage of a variety of types of organic matter. Shallow-water
species feed predominantly on algal films on hard substrates, and some appear to be
exclusively algal browsers. Encrusting animals, particularly bryozoans, are also
scraped off shell and rock surfaces. Milligan (1916) recorded Psammechinus miliar is
from British waters eating empty mollusc shell and serpulid tubes, and also the perio-
stracum of the shells of living molluscs. Jensen (1969) regarded bryozoans as of
vital importance as food for P. miliaris in Danish and Norwegian waters, whereas
Krumbein and Van der Pers (1974) recorded a preference for the boring worm
Polydora ciliata in this species and Echinus esculentus at Heligoland.
Ormond and Campbell (1971) emphasized the browsing efficiency of Diadema
setosum and Echinothrix diadema in the Red Sea off Sudan. These echinoids emerged
from their borings and crevices at night to browse freely on the surrounding coral
rock surfaces, which were kept largely clean from encrusting organisms. Only locally,
where both echinoids were absent, were rich developments of encrusting algae to
be found.
The methodical exploitation of encrusting organisms for food invariably causes
732
PALAEONTOLOGY, VOLUME 18
scratching of the underlying substrate. The resulting stellate pattern of overlapping
superimposed grooves (Krumbach 1914, fig. 1; Abel 1935, fig. 310; Neumann 1966,
fig. 7 ; Krumbein and Van der Pers 1974, fig. 1 1 ; PI. 85, figs. 2 and 4) precisely dupli-
cates the trace fossil.
Furthermore, when an echinoid, browsing over a flat, algal-coated pebble, arrives
at a sharply curved edge of the pebble, the mode of employment of the teeth changes
to accommodate the different topography. The hitherto stellate orientation of the
grooves is replaced around this edge by a subparallel orientation perpendicular to
the boundary of the pebble (PI. 86, fig. 1), since on the curved surface only two
or three teeth can operate at a time. The pattern again duplicates the trace fossil
(PI. 86, fig. 2).
The occurrence of the trace fossil finds a parallel in the feeding predelection of
present-day regular echinoids for boring animals in general and clionid sponges
in particular. Hancock (1957) reported that Psammechinus miliaris kept under
experimental conditions attacked only those oyster shells (dead or alive) that were
infested by the sponge Cliona celata or the polychaete worm Polydora ciliata, and
that these boring organisms were the sole reason for the attack. The echinoid caused
severe erosion of the shell in order to expose and eat the enclosed worms and sponges.
In the natural environment of the North Sea, Krumbein and Van der Pers (1974)
also recorded active erosion of limestone by Echinus esculentus feeding on Polydora
ciliata.
EXPLANATION OF PLATE 87
Fig. 1. Gnathichnus pentax around a pore of a sponge-boring (Entobia megastoma (Fischer)) in a belemnite
(Belemnitella mucronata) from Upper Campanian white chalk, Keswick, Norfolk, England. 85.964(4),
Norwich Castle Museum. x20.
Fig. 2. Gnathichnus pentax around a pore of Entobia megastoma in Belemnitella mucronata from Upper
Campanian white chalk, Norwich, England. 2127(1), Norwich Castle Museum. X 20.
Fig. 3. Gnathichnus pentax around pores of Entobia cretacea Portlock (sponge-boring) in Inoceramus
digitatusi. de C. Sowerby (non Schliiter). Probably Coniacian, white chalk, south-east England, locality
unknown. Paratype. GSM 115027. x5.
Fig. 4. Gnathichnus pentax around broken open boring of a sponge in Belemnitella aff. lanceolata. Degree
of destruction of the substrate lies between those in figs. 2 and 8. Lower Maastrichtian white chalk,
Kongsted, Denmark. MMH 13391. x6.
Fig. 5. An unusually clearly pentaradiate Gnathichnus pentax around a clionid sponge papillar boring in
Belemnitella sp. Lower Maastrichtian, Zeltberg/Liineburg, West Germany. Collection of Nieder-
sachsischen Landesamtes fiir Bodenforschung, Hannover, catalogue kma 12. Photo E. Voigt. X 12.
Fig. 6. As Plate 86, fig. 3. x4.
Fig. 7. Typical location of Gnathichnus pentax, around the muscle attachment area of oysters. Loss of the
aragonite myostracum leaves only that part of the trace that extended on to the surrounding calcite shell
surface. Lower Campanian Burditt Marl Member, Austin chalk from Little Walnut Creek, east of
Austin, Texas, U.S.A. MMH 13392. x3-5.
Fig. 8. ‘Wreck’ of a Belemnitella sp. broken open by foraging animals eating the boring sponges that pro-
duced the internal cavities. Scratches around the holes indicate the work of regular echinoids. Uppermost
Campanian white chalk, 1 m below horizon 595, Saturn Quarry, Kronsmoor, West Germany. MMH
13393. x 3.
PLATE 87
BROMLEY, echinoid bioerosion
734
PALAEONTOLOGY, VOLUME 18
DISCUSSION
The close similarity of the fossil material with the work of many species of browsing
and foraging regular echinoids renders it probable beyond reasonable doubt that the
trace fossil is also the work of echinoids. The geological occurrence lends further
support to this interpretation. The bioerosion sculpture has so far been found only
in deposits that appear to have been laid down in fully marine conditions. The trace
fossil is particularly abundant in sediments representing shallow-water well-
oxygenated environments.
So far the pentaradiate trace fossil, common in Jurassic and younger strata, has
not been found in Triassic or Palaeozoic rocks. This may be correlated with evo-
lutionary changes in the structural development of the jaw apparatus in regular
echinoids. The first perignathic girdles appeared in the Permian and progressive
evolutionary changes of both the girdle and the lantern occurred during the Mesozoic.
Kier (1974, pp. 53-56 and 62) interpreted these changes in terms of promoting the
biting power and mobility of the lantern. The appearance of the strengthened tooth
with T-shaped cross-section in the stirodont lantern, and change over from apophyses
to auricles in the girdle occurred in late Triassic to early Jurassic times. It is significant,
therefore, that the earliest pentaradiate bite known is of early Jurassic age (PI. 85,
fig. 1). The fully developed camarodont lantern appeared in the Maastrichtian, having
extremely large muscles capable of moving the pyramids and their teeth with great
force against the bottom (Kier 1974, p. 55). It may therefore also be significant that
the earliest known occurrences of extensive areas browsed uniformly by five teeth
date from only shortly before the Maastrichtian (PI. 86, figs. 4-5; PI. 88, fig. 7).
From Upper Cretaceous rocks there is some evidence that the sculpture may prove
a useful palaeoenvironmental indicator. Substrates from localities representing
shallow-water environments have been extensively browsed (PI. 86, fig. 4) while in
deeper-water deposits of comparable age the trace fossil is concentrated locally
around borings (PI. 87, figs. 1-4). This difference may be a reflection of the presence
and absence, respectively, of algal films in these environments.
In most of the rocks in which these trace fossils have been found, one or more
EXPLANATION OF PLATE 88
Figs. 1-3. Gnathichnus pentax around acrothoracic borings (Rogerella mathieui Saint-Seine) in Echimcorys
sp., showing different stages in the destruction of the borings. x6. 1, Santonian or Coniacian, white
chalk from Guston near Dover, England. Institute of Geological Sciences, London, CJW 806. 2, Cam-
panian white chalk from West Harnham, SW. of Salisbury, England. I.G.S., London, Zn 1903. 3, Lower
Maastrichtian white chalk of Gr§ryg, M0ns Klint, Denmark. MMH 13394.
Fig. 4. Gnathichnus pentax around the aperture of a small boring in Echinocorys sp. Bed S, Lower Maas-
trichtian white chalk at Sidestrand, England. X 10.
Fig. 5. Gnathichnus pentax at site of attack of four tubefeet of Echinocorys sp. Santonian white chalk of the
coast between Kingsgate and Foreness, Thanet, Kent, England. I.G.S., London, GSM 88258. x6.
Fig. 6. Gnathichnus pentax around a broken open boring in Inoceramus digitatus. Same specimen as Plate
87, fig. 3. X 6.
Fig. 7. Paratype of Gnathichnus pentax. Extensive but light bioerosion of the external surface of an Ino-
ceramus sp. of "cuvierC group. Turonian? White chalk of SE. England, locality unknown. GSM 1 15029.
X 1-5.
PLATE 88
7
BROMLEY, Gnathicimus pentax
736
PALAEONTOLOGY, VOLUME 18
regular echinoids are preserved that may represent the trace maker(s) (e.g. Voigt
1972, p. 119). In the Lower Campanian chalk {Gonioteuthis quadrata Zone) of
Hampshire, England, the trace fossil occurs with Stereocidaris sp., Salenia granulosa,
and Phymosoma sp. In the Upper Campanian chalk of England they are accompanied
by Stereocidaris 'serrifera', Salenia heberti, and Phymosoma regularis, while in the
Maastrichtian chalk of England, Germany, and Denmark the sculpture is accom-
panied by Salenia pygmaea, Stereocidaris spp., Phymosoma sp., and (Denmark only)
Tylocidaris baltica (C. J. Wood, pers. comm. 1975). On the other hand, the extensive
but otherwise rather similar grooves on shells in the Campanian littoral deposits of
Sweden are again accompanied by species of Salenia, Tylocidaris, and Stereocidaris
but the species are different from those of the correlative deeper-water chalk facies.
In those cases where several regular echinoid species accompany the trace fossil
it is doubtful whether the traces produced by the different species can be distinguished.
The large Echinus esculentus and the small Psammechinus miliaris, feeding together
in aquaria, each produce large or small traces according to local changes of feeding
habits (W. Krumbein, pers. comm. 1974). Stars of a particular echinoid individual
also tend to be smaller on hard substrates than on softer rocks (Krumbein and Van der
Pers 1974, p. 12). Krumbach (1914), however, noted that Sphaerechinus granularis
consistently produced more widely separated stars than Strongylocentrotus lividus in
the same environment. The scratches of S. lividus were so close together that they
overlapped so that the entire substrate surface was eroded clean of algae, but the
grazing was restricted to limited territories. Arbacia pustulosa also fed in the same
environment, but this species restricted its activities to narrow clefts in the substrate
and to the underside of stones and overhangs, so that its feeding traces were readily
separable from those of the other two species.
NAMING THE TRACE FOSSIL
For future reference it is necessary to name the distinctive groove pattern as a trace
fossil. The name is applied to the basic stellate unit, multiplication of which produces
the characteristic bioerosion sculpture. This is analogous to soft sediment trace
fossils in which the isolated burrow bears the name, but its repetition produces a
EXPLANATION OF PLATE 89
Fig. 1. Phosphatic pebble from Cenomanian greensand at Miilheim/Ruhr, near Essen, West Germany.
Grooves scattered over the entire surface resemble the work of browsing regular echinoids. Private
collection of H. Klaumann. x4.
Fig. 2. External surface of a shell of Arctica islandica (L.) bioeroded by browsing organisms (Bromley and
Tendal 1973, pi. Id). A remnant island of original shell surface at left, carrying a patch of black perio-
stracum, shows the surface to have been lowered generally by about 1 mm. The double ridge is a ghost
of an encrusting serpulid tube that originally protected the shell surface. Four orifices of a sponge-boring
(papillae of Cliona celata are visible within) are surrounded by radiating grooves typical of the work of
browsing echinoids (probably Psammechinus miliaris). The remainder of the surface bears fine striae
from the radulae of gastropods. Dredged in southern Kattegat, Denmark. Housed in the Zoological
Museum, Copenhagen, x 13.
Fig. 3. Rodent gnawing traces on terrapin bone (plastron). Nacogdoches, Texas, U.S.A. x 3.
PLATE 89
BROMLEY, echinoid bioerosion
738
PALAEONTOLOGY, VOLUME 18
bioturbation fabric. The definition of the name is based on morphological characters
alone, with the understanding that the structures have a biogenic origin. The generic
name is available for trace fossils of other types produced by rasping, biting, and
gnawing animals.
Ichnogenus gnathichnus nov.
Type ichnospecies. Gnathichnus pentax nov.
Diagnosis. Biogenic sculpture consisting of grooves, pits, and scratches on hard
substrates.
Ichnospecies Gnathichnus pentax nov.
Plate 85, figs. 1-3; Plate 86, figs. 3-5; Plate 87, figs. 1-7; Plate 88, figs. 1-7.
Type material. The variable aspects of the trace fossil cannot be represented by a single specimen. The
holotype is an example showing particularly good preservation of the grooves. The paratypes illustrate
two other typical modes of occurrence.
Holotype. MMH 13386, housed in the Mineralogisk Museum, Copenhagen, Denmark (PI. 85, fig. 3).
Paratypes. GSM 1 1 5029 (PI. 88, fig. 7) and GSM 1 1 5027 (PI. 87, fig. 3 ; PI. 88, fig. 6), housed in the Institute
of Geological Sciences, London.
Locus typicus. Kritika, Rhodes, Greece.
Stratum typicum. Sgourou Formation, lowermost Pleistocene.
Diagnosis. Gnathichnus consisting of a regular stellate grouping of five similar grooves
radiating at c. 12°.
Description. The trace fossil almost invariably occurs in multiples of several over-
lapping stars and can cover considerable areas of substrate with grooves of similar
dimensions intersecting at more or less 72° (and 144°). Commonly concentrated
around skeletons of encrusting organisms and apertures to borings.
Interpretation. Browsing and foraging traces attributed to dental erosion by regular
echinoids. '
Range. Lower Jurassic to Recent.
Acknowledgements. This paper has benefited considerably from the help and advice of C. J. Wood (London)
and R. Goldring (Reading) who read earlier drafts; W. E. Krumbein (Oldenburg) provided photographs
and facts (text-fig. 2); J. Schneider (Gottingen) lent recent material (PI. 85, fig. 4; PI. 86, fig. 1); W. Riegraf
(Tubingen) supplied the peel for Plate 85, fig. 1 and E. Voigt (Hamburg) the photograph for Plate 87,
fig. 5. The specimen in Plate 89, fig. 1 was lent by H. Klaumann (Mulheim/Ruhr) and W. Pockrandt
(Hannover) also loaned comparative fossil material.
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ABEL, o. 1935. Vorzeitliche Lehensspuren. Fischer, Jena. 644 pp.
ANKEL, w. E. 1929. Frassspuren einer Meeresschnecke. Natur Museum, 59, 95-99.
1936. Die Frassspuren von Helcion und Littorina und die Funktion der Radula. Zool. Anzeiger, 9
(Suppl.), 174-182.
1937. Wie frisst Littorinal Senckenhergiana, 19, 317-333.
BROMLEY: ECHINOID BIOEROSION
739
BAKUS, G. J. 1964. The effects of fish-grazing on invertebrate evolution in shallow tropical waters. Occ. Pap.
Allan Hancock Fdn, no. 27.
1966. Some relationships of fishes to benthic organisms on coral reefs. Nature, London, 210, 280.
BISHOP, G. 1975. Fossil evidence of predation and paleopredation. In frey, r. w. (ed.). The study of trace
fossils. Springer, New York.
BOEKSCHOTEN, G. J. 1966. Shell borings of sessile epibiontic organisms as palaeoecological guides (with
examples from the Dutch coast). Palaeogeogr., PalaeoclimatoL, Palaeoecol. 2, 333-379.
1967. Palaeoecology of some Mollusca from the Tielrode Sands (Pliocene, Belgium). Ibid. 3, 311-362.
BROMLEY, R. G. 1970. Borings as trace fossils and Entobia cretacea Portlock, as an example. In crimes, t. p.
and HARPER, J. c. (eds.). Trace fossils. Geol. J. special Issues, 3, 49-90.
and TENDAL, o. s. 1973. Example of substrate competition and phobotropism between two clionid
sponges. J. ZooL, Land. 169, 151-155.
CARRiKER, M. R. 1969. Excavation of boreholes by the gastropod, Urosalpin.x: an analysis by light and
scanning electron microscopy. Am. Zoologist, 9, 917-933.
HANCOCK, D. A. 1957. The feeding behaviour of the sea urchin Psammechinus miliaris (Gmelin) in the
laboratory. Proc. zool. Soc. Lond. 129, 255-262.
HEiNBERG, c. 1973. The internal structure of the trace fossils Gyrochorte and Curvolithus. Letliaia, 6,
227-238.
JENSEN, M. 1969. Breeding and growth of Psammechinus miliaris (Gmelin). Ophelia, 7, 65-78.
KiER, p. M. 1974. Evolutionary trends and their functional significance in the post-Paleozoic echinoids.
Paleont. Soc., Mem. 5, 95 pp.
and GRANT, R. E. 1965. Echinoid distribution and habits, Key Largo Coral Reef Reserve, Florida.
Smithsonian misc. Coll. 149, 68 pp.
KRUMBACH, T. 1914. Mitteilungcn fiber die Nahrung felsenbewohnender Seeigel der nordlichen Adria.
Notizen fiber die Fauna der Adria bei Rovigno. Zool. Anzeiger, 44, 440-451.
KRUMBEiN, w. E. and VAN DER PERS, J. N. c. 1974. Diving investigations on biodeterioration by sea-urchins
in the rocky sublittoral of Helgoland. Helgoldnder wiss. Meeresunters. 26, 1-17.
MARKED, K. and MAiER, R. 1967. Beobachtungen an lochbewohnenden Seeigeln. Natur Museum, 97, 233-243.
MCKERROW, w. s., JOHNSON, R. T. and JAKOBSON, M. E. 1969. Palaeoecological studies in the Great Oolite
at Kirtlington, Oxfordshire. Palaeontology, 12, 56-83.
MILLIGAN, H. N. 1916. Observations on the feeding habits of the purple-tipped sea-urchin. Zoologist (4),
20, 81 99.
NEUMANN, A. c. 1965. Processes of recent carbonate sedimentation in Harrington Sound, Bermuda. Bull,
mar. Sci. 15, 987-1035.
1966. Observations on coastal erosion in Bermuda and measurements of the boring rate of the sponge,
Cliona lampa. Limnol. Oceanogr. 11, 92-108.
ORMOND, R. F. G. and CAMPBELL, A. c. 1971. Observations on Acanthaster planci and other coral reef echino-
derms in the Sudanese Red Sea. Symp. zool. Soc. Lond. no. 28, 433-454.
RiEGRAF, w. 1973. Bissspuren auf jurassischen Belemnitenrostren. N. Jb. Geol. Paldont., Mh., 1973, 494-
500.
UMBGROVE, J. H. F. 1939. De atollen en barriere-riffen der Togian-Eilanden. Leidsche geol. Mededeel. 11,
VOIGT, E. 1972. liber Talpina ramosa v. Hagenow 1840, ein wahrscheinlich zu den Phoronidea gehoriger
Bohrorganismus aus der Oberen Kreide, nebst Bemerkungen zu den fibrigen bisher beschreibenen
kretazischen ‘Talpina’-Arten. Nachrichten Akad. ^Viss. Gottingen 2, math.-phys. Kl. 1972, 93-126.
132-187.
Typescript received 10 March 1975
Revised typescript received 18 April 1975
R. G. BROMLEY
Institute of Historical Geology and Palaeontology
University of Copenhagen
DK-1350 Copenhagen
Denmark
ENGLISH HYPSILOPHODONTID
DINOSAURS (REPTILIA: ORNITHISCHI A)
by PETER M. GALTON
Abstract. A premaxillary tooth from the Stonesfield Basin (Bathonian) of Stonesfield may represent the oldest
hypsilophodontid described to date. However, small bones from the Lias (Jurassic) of Charmouth are not hypsilopho-
dontid and were correctly referred to the primitive ankylosaur Scelidosaurus harrisoni. A femur from the Oxford Clay
(Callovian) of Peterborough is regarded as an iguanodontid (Camptosaunis (?) leedsi Lydekker). A dentary tooth from
the Kimmeridge Clays of Weymouth represents the oldest undoubted hypsilophodontid described to date from
England. In addition to the well-known Hypsilophodon fo.xii Huxley, a new Wealden species is tentatively referred to
the genus Dryosaurus. Some Wealden specimens previously referred to Iguanodon are hypsilophodontid and represent
individuals with a length of up to 4-2 m.
The Hypsilophodontidae are a family of conservative bipedal ornithischian dinosaurs
(ornithopods) with inset cheek teeth, and neither a rostral bone nor any marked
thickening of the skull roof. They were herbivorous and fast-running or cursorial with
an elongate hind limb with the tibia longer than the femur (Galton 1972, 1973,
1974fl, b). My concept of the family Hypsilophodontidae is more restricted than that
ofThulborn(1970, 1971, 1972); I refer only the following to this family: Nanosaurus^l)
rex, specimens referred to Laosaurus, Dryosaurus altus (all upper Jurassic, North
America; Galton and Jensen 1973a, Gilmore 1925; Marsh 1896; Dysalotosaurus
lettow-vorbecki (upper Jurassic, Tanzania; Janensch 1955); Hypsilophodon foxii
(lower Cretaceous, England; see Galton 1974a); and 'Laosaurus' minimus, Parkso-
saurus warreni (upper Cretaceous, North America; see Galton 1973; Gilmore 1924a;
Parks 1926). Dryosaurus Marsh, 1894 and Dysalotosaurus Pompeckj, 1920 are
extremely similar and, as will be detailed elsewhere, these two genera are probably
synonymous. Consequently, the reported record of hypsilophodontids is very limited
with the English records representing the whole of Eurasia. Previously described and
new hypsilophodontid material from England is reviewed in stratigraphical sequence
starting with the oldest record. The abbreviations used for measurements are explained
in the caption to Table 1 and institution names have been abbreviated as follows;
AM, American Museum of Natural History, New York; BM, British Museum
(Natural History), London; CM, Carnegie Museum, Pittsburgh, Penn., U.S.A.;
UCMP, University of California Museum of Paleontology, Berkeley, U.S.A.; US,
United States National Museum, Washington D.C.; YPM, Peabody Museum, Yale
University, New Haven, Conn., U.S.A.
JURASSIC
Sinemurian. Owen ( 1861 ) described several specimens of the ornithischian dinosaur Scelidosaurus harrisoni
Owen from the Lower Lias of Charmouth, Dorset. Amongst this material are the bones which Owen
(1861) regarded as representing a juvenile of Scelidosaurus harrisoni (dorsal centrum, phalanges, partial
right hind limb, text-fig. 1a-d, casts as BM 5909, originals in Charmouth Museum; see Owen 1861, pi. 2,
[Palaeontology, Vol. 18, Part 4, 1975, pp. 741-752.]
742
PALAEONTOLOGY, VOLUME 18
TABLE 1 . Measurements of femora in millimeters. FT, minimum distance from proximal end to distal edge of fourth
trochanter; L, maximum length; LA, estimated total length of body, for hypsilophodontids calculated on a pro-
portional basis from BM R196. Camptosaurus based on Gilmore (1909); Wd, greatest width of distal end; Wm,
minimum width of shaft; Wp, greatest width of proximal end.
LA
L
Wp
Wd
Wm
FT
FT/L
m
ft
Hypsilophodon foxii
BM R5830
101
27
25
11
43
0-43
0-91
3-0
BM R196
150
—
—
—
65
0-43
1-36
4-5
BM R5829
202
52
26
—
87
0-43
1-82
60
Dryosauriis (?) canaliculatus^
140
3
33
14
58
0-41
1-27
4-2
Dryosaurus alius
AM 834
222
59
55
22
96
0-43
2-03
6-7
YPM 1876
362
84
98
39
168^
0-44
3-30
10-9
CM 1949
470
131
—
—
212
0-45
4-27
141
Camptosaurus (?) leedsP
280
65
73
122^
0-48
3-33
11-0
Camptosaurus ampins
US 2210
258
72
71
137
0-53
3-03
100
YPM 1877
585
208
192
■
300
0-51
5-15
17-0
' BM R185. ^ BM R1993. ^ Estimated.
distal half of humerus figured as proximal part of tibia, metatarsal 3 figured as a partial fibula). Recently,
Newman (1968) considered that these bones probably belong to the genus Hypsilophodon or some allied
form. However, the femur (text-fig. 1a, b) differs from those of hypsilophodontids (text-figs. 2g-l, 3) in
several respects; the shaft is straight rather than bowed antero-posteriorly, the apex of the lesser trochanter
is well below that of the greater trochanter rather than at the same level, and the non-pendant fourth
trochanter is at mid-length as against a pendant fourth trochanter more proximally placed. In these features
the femur (text-fig. 1a, b) appears to agree with that of the complete skeleton of S. harrisoni (BM R1 1 1 1).
The femur of the juvenile is almost identical to the femur (Charig 1972, pi. 6A) of the partial skeleton
(BM R6704, see Rixon 1968, fig. 103) of a small individual that is referred to as Cf. 'Scelidosaurus harrisoni
Owen’ by Charig ( 1 972, p. 138). The length of the femur (text-fig. 1 a) is about 1 33 mm and that of metatarsal
3 about 60 mm to give a femur to metatarsal 3 ratio of about 0-45. In Hypsilophodon foxii this ratio is 0-62
in BM R5830 (femoral length 101 mm) and 0-56 in BM R196 (femoral length 151 mm). On the basis of this
ratio the bones from Charmouth should not be referred to the family Hypsilophodontidae which, as noted
by Gabon (1972, 1974a, b), is restricted to genera which were cursorial. The value of 0-45 is about correct if
the remains are regarded as a juvenile individual of S. harrisoni because the corresponding value for BM
R1 1 1 1 is about 0-34 (from Owen 1863, pi. 10).
Comparisons of BM R5909 and R6704 with the much larger BM Rllll (size ratio about 1:4) show
that corresponding bones are almost identical, so all specimens are referable to Scelidosaurus harrisoni
Owen rather than to two separate ornithopod families (BM R5909, R6704 to Fabrosauridae; BM Rllll
to Scelidosauridae) as suggested by Thulborn (1974). The combined lengths of the femur, tibia, and meta-
tarsal 3 of BM Rllll is short relative to the trunk (combined length of centre of dorsal vertebrae) so
the hind limb to trunk ratio at 0-85 is comparable to that of fully quadrupedal ornithischians (stegosaurs,
0-86-0-90; Cretaceous ankylosaurs, 0-69; ceratopsians, 0-90-1 08; see Gabon 1970, table 2) but much less
than that of the facultitively or fully bipedal ornithopods (hadrosaurs, 122-144, see Gabon, 1970, table 1 ;
iguanodontids I-08-1-24; hypsilophodontids, 1-44 1-53; psittacosaurids 1-3; see Gabon, 19716, table 1).
1 disagree with the referral of Scelidosaurus, an obligatory quadruped while it walked or ran, to the Orni-
thopoda, an order whose members were characterized by bipedality. Scelidosaurus is probably a very primi-
tive ankylosaur but any discussion of its affinities must await the detailed anatomical study of Scelidosaurus
being prepared by Dr. A. J. Charig.
Balhonian. The tooth YPM 7367 (text-fig. 4a-c) was collected prior to 1870 by G. J. Chesler from the
Stonesfield Slate (Oracilisphinctes progracilis Zone) of the Great Oolite Series at Stonesfield, Oxfordshire.
The form of the tooth is similar to that of the premaxillary teeth of Hypsilophodon (Gabon 1974a) but it
differs in the absence of small denticles on the anterior and posterior edges of the crown (text-fig. 4a, b)
and the presence of a large concave wear surface (text-fig. 4b, c). However, isolated premaxillary teeth of
the late Cretaceous iguanodontid Thescelosaurus (Gabon 19746, pi. 1, figs. 7-11) are almost identical to
GALTON: H YPSILOPHODONTID DINOSAURS
743
A
B
C
E
F
G
TEXT-FIG. 1. A-D, SceMosaurus harrisoni, juvenile after Owen (1861), bones of right hind limb, xO-5;
A, femur in anterior view; b, femur in posterior view; c, metatarsal 3 in anterior view; d, metatarsal 4 in
anterior view; e-m, Camptosaurus nanus, left femur, xO-25, after Gilmore (1909); F, G, hypsilophodontid
right tibia, BM 36506 in f, medial view, G, anterior view; h, hypsilophodontid left ischium, BM 2183, lateral
view of proximal end, xO-33; i, hypsilophodontid left pubis, BM R720, lateral view with cross-section of
postpubic rod, xO-25; J, k, hypsilophodontid right pubes in lateral view with cross-section of anterior
process, xO-33; J, BM R169, k, BM 36538; l, m, hypsilophodontid left femoral shaft, BM R8669, xO-25
with L, proximal cross-section, m, medial view, a, acetabulum; ap, anterior or prepubic process; c, cnemial
crest ; d, depression, area of insertion of M. caudi-femoralis longus ; il, surface for ilium ; 1, lesser trochanter ;
o, obturator foramen; op, obturator process; p, posterior process or postpubic rod; pu, surface for pubis;
4, fourth trochanter. Scale line represents 10 cm.
744
PALAEONTOLOGY, VOLUME 18
E
TEXT-FIG. 2. Upper Jurassic ornithopod femora, a-f, Camptosaurns{l) leedsi Lydekker, holotype
left femur, BM R1993, xO-3; g-l, Dryosaurus altus (Marsh), left femur of holotype, YPM
1876, xO-23. Views: a, g, lateral; b, h, posterior; c, i, medial; d, j, anterior; E, k, proximal;
F, L, distal. Fourth trochanter indices (minimum distance from proximal surface of head to distal
edge of fourth trochanter) in a: a, 0-44; h, 0-46; c, 0-48; d, 0-50; for identification of structures
see text-fig. 3.
GALTON: H YPSILOPHODONTI D DINOSAURS
745
YPM 7367. Wear facets are also reported on the medial surface of teeth preserved in situ in a premaxilla
of the Triassic ornithopod described as Lycorhimis by Thulborn 1970, fig. 2. The wear on these ornithopod
premaxillary teeth was presumably caused by contact with the horny predentary sheath. YPM 7367 (text-
fig. 4a-c) is tentatively identified as a left premaxillary tooth of an ornithopod dinosaur and it may represent
the oldest hypsilophodontid yet described.
Oxfordian. Lydekker (1889) described a left femur (BM R1993, text-fig. 2a-f) from the Oxford Clay near
Peterborough, Northamptonshire, as a new species of Camptosaurus, C. leedsi. Gilmore (1909) pointed
out that the fourth trochanter extends on to the distal half of the shaft in all described species of Campto-
saurus and noted that, if C. leedsi is referable to an American genus, then its closest affinities are with
Dryosaurus. ‘Camptosaurus' leedsi is shown as being closely related to the hypsilophodontids Dryosaiirus
and Dysalotosaurus in the phyletic charts given by Galton (1972, 1973, 1974a, b).
The lesser trochanter of all hypsilophodontids is relatively slender (text-figs. 2g, i, 3a, c, g, i) and it is
not expanded antero-posteriorly as it is in Camptosaurus (text -fig. 1e) and BM R1993 (text-fig. 2a, c, e).
On the basis of the figures given by Lydekker (1889, 1890), the fourth trochanter index of BM R1993 is
about 0-45 (text-fig. 2a), a value comparable to that of hypsilophodontids (Table 1 ; Dysalotosaurus 0-45,
Nanosaurus (?) rex 0-43, Galton and Jensen 19736). However, the distal surface of the fourth trochanter
as given by Lydekker (1889, 1890) is based on a broken surface (text-fig. 2a, c). The exact value of the
fourth trochanter index is not known but it was probably close to 0-48 (see text-fig. 2a). In Camptosaurus
(text-fig. 1e) the fourth trochanter index is 0-51-0-53 (Table 1). In Camptosaurus (text-fig. 1e) and BM
RI993 (text-fig. 2c) the depression for the M. caudi-femoralis longus (Galton 1969) is shallow and close
to the fourth trochanter. In Dryosaurus (text-fig. 2i) and Dysalotosaurus (Janensch 1955, pi. 14, figs, lb, 2)
it is deep and situated more anteriorly on the shaft but this position is unique for hypsilophodontids. The
difference in depth is probably not significant because in Hypsilopliodon foxii this depression is shallow
in some femora and deep in others (Galton 1969, 1974a). The distal ends of the femora of Camptosaurus
(Gilmore 1909, YPM 1877) and Dryosaurus (text-fig. 2g-j, l) are very similar with a well-developed anterior
intercondylar groove (text-fig. 2l) where as that of BM R1993 is quite shallow (text-fig. 2f).
The femur BM R1993 differs from those of Camptosaurus in only a couple of respects; the fourth tro-
chanter is more proximally placed (text-figs. 1e, 2a) and the anterior intercondylar groove is more shallow
(text-fig. 2f). On the basis of the femur, BM R1993 from the Oxfordian is an ideal ancestor for the American
species of Camptosaurus which are of Kimmeridgian or possibly Portlandian age. Unfortunately no other
parts of the anatomy of the English form are known. I now consider that BM R1993 is best assigned to
the family Iguanodontidae as Camptosaurus (?) leedsi Lydekker rather than as a hypsilophodontid related
to Dryosaurus as suggested by Gilmore (1909) and Galton (1972, 1973, 1974a, b).
Kimmeridgian. The history of the dentary tooth UCMP 4961 1 (text-fig. 4d-f) is unknown but it came from
the Kimmeridge Clays of Weymouth, Dorset. It was collected along with three theropod teeth (UCMP
49612; two complete crowns, height 15 mm, one tip from a larger tooth) tentatively identified as Megalo-
saurus sp. The more thickly enamelled surface of the crown (text-fig. 4d) has a strong central ridge, the
size of which is not obvious because of the other longitudinal ridges on either side (text-fig. 4d, e). The
longitudinal ridges on both sides of the crown (text-fig. 4d, f) are more numerous and more prominent
than those on the teeth of Hypsilopliodon (Galton 1974a), Laosaurus (Marsh 1896, pi. 55, fig. 1), Dryosaurus
(Marsh 1878 as Laosaurus, Galton and Jensen 1973a), and Dysalotosaurus (]ar\m&c\\ 1955). The longitudinal
ridges are even more prominent on the teeth of Thescelosaurus with those of the thickly enamelled surface
forming two converging cresentic patterns (Galton 19746; Sternberg 1940). Undescribed teeth (YPM,
unnumbered) from the upper Jurassic of North America are very similar to UCMP 4961 1. UCMP 49611
undoubtedly represents a hypsilophodontid dinosaur but discussion of its affinities must await revision of
the American Jurassic hypsilophodontids.
CRETACEOUS
Hypsilopliodon foxii. The holotype of H. foxii Huxley, 1869 is a skull and the centrum of a dorsal vertebra
(BM R197, Huxley 1870) from the Wealden Beds (pre-Aptian and probably Barremian) exposed near
F
A
B
C
D
E
TEXT-FIG. 3. Lower Cretaceous hypsilophodontid femora, a-f, Dryosaums (?) canaliculatus n. sp., left femur,
BM R185 (with some details from right femur, BM R184), xO-45; G-l, Hypsilophodon foxii Huxley, left
femur, BM R5830, x 0-75 with indication of lines of actions of muscles associated with trochanters, modified
from Galton (1969). Views as in text-fig. 2. AG, anterior intercondylar groove; C-FB, M. caudi-femoralis
brevis ; C-FL, M . caudi-femoralis longus ; FT, fourth trochanter ; GT, greater trochanter ; IC, inner condyle ;
IF, M. ilio-femoralis; IT, M. ilio-trochantericus; LT, lesser trochanter; OC, outer condyle; PIFI, pubo-
ischio-femoralis internus, dorsal part.
GALTON. HYPSILOPHODONTID DINOSAURS
747
TEXT-FIG. 4. Jurassic hypsilophodontid teeth, x 3. a-c, left premaxillary tooth YPM 7367 in a, lateral
(labial); b, posterior and c, medial (lingual) views; d-f, right dentary tooth UCMP 49611 in d, medial
(lingual) ; e, posterior and f, lateral (labial) views.
Cowleaze Chine on the south-western shore of the Isle of Wight. H.foxii (text-fig. 5) is the best-represented
hypsilophodontid from England. Its diagnosis (Galton 1974a) is as follows:
Five premaxillary teeth separated by step from maxillary row with 10 or 11 teeth, 13 or 14 on dentary;
enamelled medial surface of a dentary tooth has a strong central ridge that is absent on the lateral surface
of a maxillary tooth. Narial openings completely separated by anterior process of premaxillae; large
antorbital recess or depression plus row of large foramina in maxilla; jugal does not contact quadrate;
large fenestrated quadratojugal borders lower temporal opening. Five or six sacral ribs, the additional one
borne on the anterior part of the first sacral vertebra. Scapula same length as humerus; obturator process
on middle of ischium. Femur with following combination of characters: fourth trochanter on proximal
half, lesser trochanter triangular in cross-section with a shallow cleft separating it from the greater tro-
chanter, practically no anterior condylar groove and posteriorly outer condyle almost as large as inner.
The holotype of Camptosaurus valdensis Lydekker, 1889, a large left femur (BM R167), represents a large
individual (body length about 2-27 m or 7-5 ft) of H. foxii (see Galton 1974a, pp. 102-103, pi. 2, fig. 4).
Hypsilophodon is usually considered to have been arboreal but, as discussed by Galton ( 1971a, 6, 1974a),
Hypsdophodon was a ground-living and cursorial dinosaur.
Dryosaurusl canaliculatus sp. nov.
Derivation of name. From Latin caniculatus, a channel or conduit, with reference to the deep anterior
intercondylar groove.
Diagnosis. Femur with pendant fourth trochanter well on proximal half of shaft,
rod-like lesser trochanter separated by deep cleft from greater trochanter, distally
a deep anterior intercondylar groove.
Lydekker (1888) listed under Hypsilophodon foxii the associated right and left
femora (BM R184, R185) from the Wealden of the Isle of Wight. He noted that
a small tibia (BM R186) was apparently associated with these femora but subse-
quently (1891) he referred the tibia to the coelurosaur Calamosaurus. BM R185 (text-
fig. 3a-f) resembles the femur of H.foxii (text-fig. 3g-l) in the proximal position of
the fourth trochanter. In both femora the lesser trochanter is triangular in cross-
section (text-fig. 3e, k) but the cleft separating it from the greater trochanter is deep
748
PALAEONTOLOGY, VOLUME 18
in BM R185 (text-fig. 3c, d) and shallow in Hypsilophodon (text-fig. 3i, j). At the
distal end the anterior intercondylar groove is deep in BM R185 (text-fig. 3d, f) and
practically non-existent in Hypsilophodon (text-fig. 3j, l). Posteriorly the outer con-
dyle of BM R185 (text-fig. 3b, f) is sheet-like while that of Hypsilophodon (text-fig.
3h, l) is more massive so that it is almost as large as the inner condyle. The inner
condyle of BM R185 (text-fig. 3c, f) is much squarer than that of Hypsilophodon
(text-fig. 3i, l) and the rugose area of origin of the medial head of the M. gastrocnemius
is much larger in R 185 (text-fig. 3c), extending on to the shaft and delimited anteriorly
by a sharp edge. The form of the ends of BM R185 (text-fig. 3a-f) differs from those
of Hypsilophodon (text-fig. 3g-l) in several other minor respects as can be seen by
comparing equivalent views.
These differences between BM R185 and Hypsilophodon are too great to make it
likely that BM R185 belongs to that genus. (Even though there is individual variation
in some features of Hypsilophodon foxii (Galton 1974a), this does not affect the
femur.) Another possibility is that BM R185 might belong to another Wealden
ornithopod. Two others are known: Vectisaurus?Lnd Yavelandia. Vectisaurus valdensis
Hulke, 1879 is based on an ilium and a few vertebrae but, from an examination of
the holotype (BM R2494) and of another specimen (BM R5849) that I refer to this
genus, I conclude that Vectisaurus is an iguanodontid (Galton in press). It is very
unlikely that BM R185 is referable to Vectisaurus because in iguanodontids the
fourth trochanter is on the distal half of the femur. The primitive pachycephalosaurid
Yaverlandia bitholus Galton (1971c) is based on a partial skull cap. The fourth tro-
chanter is on the proximal half of the femur of the pachycephalosaurids Stegoceras
(Gilmore 19246, pers. obs.), Homalocephale, and Prenocephale (Maryahska and
Osmolska 1974) but the lesser trochanter is separated by a shallow cleft from the
greater trochanter (Stegoceras, Prenocephale) and the anterior intercondylar groove
is shallow (Stegoceras, Homalocephale). BM R185 might be a femur of Y. bitholus
but this is considered very unlikely.
The closest approach to the femur BM R185 (text-fig. 3a-f) are those of Dryo-
saurus altus (text-fig. 2g-l) and Dysalotosaurus lettow-vorbecki (Janensch 1955,
fig. 40; pi. 14, figs. 1, 2). BM R185 can be distinguished from femora of both taxa by
the relative slenderness of the lesser trochanter (text-fig. 3a, c), the more proximal
position of the fourth trochanter (text-fig. 3a), and distally by the greater depth of
the anterior intercondylar groove (text-fig. 3d, f). The apparent slenderness of BM
R185 in comparison with that of Dryosaurus (text-fig. 2g-j) is size related because
smaller femora of Dryosaurus (AM 834) are comparably slender to BM R185. The
femora of Nanosaurus(l) rex are similar to BM R185 except that distally there is
practically no anterior intercondylar groove (Galton and Jensen 19736).
The femora (BM R184, R185) may represent a new genus but, because of the limited
nature of the available material, these specimens are made the holotype of a new
species of hypsilophodontid tentatively referred to Dryosaurus Marsh. Dryosaurus (?)
canaliculatus is shown on the phyletic charts in Galton (1972, 1973, 1974a, 6) as the
Wealden hypsilophodontid.
Additional material. BM 2459. Proximal parts of a large pair of femora ( Wm 46 mm, FT 1 52 mm, LA about
3 m or 9-84 ft), Wealden (Blue Clay), Heathfield, Sussex.
BM 28697. Distal end left femur (Wd 45 mm). Isle of Wight.
GALTON: H YPSILOPHODONTID DINOSAURS
749
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PALAEONTOLOGY, VOLUME 18
BM 36509. Distal end of small right femur (Wd 27 mm) catalogued by Lydekker (1888) as Hypsilophodon
foxii', Cuckfield, Sussex. The fossil record of H. foxii is now restricted to the Isle of Wight because this
was the only specimen from elsewhere that was referred to H. foxii (Lydekker 1888; Swinton 1936).
BM R8420, R8421. Distal ends of two femora (Wd 40 mm, 34 mm) previously catalogued by Lydekker
(1888) under BM R 170 as Iguanodon, Isle of Wight.
BM R8670. Distal end of right femur (Wd 50 mm), Bone Bed between high and low water, Clinton
Chine, Isle of Wight. The partial left ischium (BM 2183, text-fig. 1h) from the Wealden of Cuckfield, Sussex
is similar to those of Dryosaums and Dysalotosaurus (Janensch 1955). This ischium is probably also referable
to Dryosaurus (?) canaliculatus rather than to Iguanodon as listed by Lydekker (1888, p. 235).
Larger Wealden hypsilophodontid material. A search through the Wealden Iguano-
don material resulted in the identification of several hypsilophodontid specimens
representing larger individuals. The anterior or prepubic process of the pubis of
hypsilophodontids is bar-shaped where as that of iguanodontids is a deep and
laterally flattened plate. The following pubes are identified as hypsilophodontid :
BM 36538. Fragmentary right pubis (text-fig. Ik), Cuckfield, Sussex; figured by Mantell (1827, pi. 16,
fig. 3) as part of a scapula, listed by Lydekker (1888, p. 235) as Iguanodon pubis. The lips of the obturator
process are separated by an obliquely inclined gap of 1 mm which in life was probably filled with cartilage.
BM R169. Fragmentary right pubis (text-fig. It), Isle of Wight, listed by Lydekker (1888, p. 235) as
Iguanodon.
BM R720. Left pubis incomplete anteriorly (text-fig. Ill), Horsham, Sussex; listed by Lydekker (1888,
p. 223) as I. mantelli. The range of variation of these pubes is comparable to that within Hypsilophodon
foxii (Galton 1974a, figs. 46, 48, 49) and all may be referable to this taxon. The length of the postpubic
rod of BM 720 at 340 mm is almost twice that of BM R 1 96 so BM 720 is from an animal with a total length
of about 2-74 m or 9 ft. The section of femoral shaft (BM R8669, text-fig. 1l, m) from Compton Bay, Isle
of Wight has a maximum width of 57 mm immediately below the fourth trochanter (originally pendant).
On the basis of comparisons with the femora of Camptosaums and Dryosaurus the original length of this
femur was about 470 mm so LA about 4-2 m or 14 ft. Three tibiae listed by Lydekker (1888, p. 237) as
Iguanodon are much too slender to be iguanodontid and are regarded as hypsilophodontid.
BM 36506. Right tibia (text-fig. If, g), Cuckfield, Sussex, L 332 mm, Wp 72 mm, Wd 65 mm, LA about
2-51 m or 8-3 ft.
BM 36508. Left tibia. Isle of Wight, L 168 mm.
BM R124. Right tibia. Isle of Wight, L 280 mm, Wp 67 mm, Wd 58 mm, LA about 212 m or 7 0 ft.
It should be noted that the following specimens cited in the literature as H. foxii should be regarded as
hypsilophodontid, generically and specifically indeterminate: BM Nos. R170, R183, R186, R199, R200,
R202a, R752, R2481, R8422 (for details of specimens and citations see Galton 1974a, pp. 7-12) as are the
following uncited specimens as H. foxii: BM Nos. R198, R201, R2479, R2480, R2482-2486, R2489-2493,
R519LR6373.
Acknowledgements. I thank the following for access to collections, A. J. Charig and C. A. Walker, British
Museum (Natural History); J. H. Ostrom, Peabody Museum, Yale University; and S. P. Welles and
R. Long of the University of California Museum of Paleontology at Berkeley. R. C. Fox kindly provided
a cast of Stegoceros. P. Olsen drew the teeth and Miss P. Rubino typed the manuscript. I acknowledge
a N.E.E.C. grant while at the Department of Zoology, King’s College, London University, and a Faculty
Research Grant from the University of Bridgeport.
REFERENCES
CHARIG, A. J. 1972. The evolution of the archosaur pelvis and hind-limb: an explanation in functional terms,
pp. 121-155. In JOYSEY, K. A. and kemp, t. s. (eds.). Studies in vertebrate evolution. Edinburgh, 284 pp.
GALTON, p. M. 1969. The pelvic musculature of the dinosaur Hypsilophodon (Reptilia: Ornithischia).
Post ilia, 131, I 64.
GALTON: H YPSILOPHODONTID DINOSAURS
751
GALTON, p. M. 1971a. Hypsilophodon, the cursorial non-arboreal dinosaur. Nature, Loud. 231, 159-161.
19716. The mode of life of Hvpsilophodon, the supposedly arboreal ornithopod dinosaur. Lethaia,
4, 453-465.
1971c. A primitive dome-headed dinosaur (Ornithischia : Pachycephalosauridae) from the Lower
Cretaceous of England and the function of the dome of pachycephalosaurids. J. Paleont. 45, 40-47.
1972. Classification and evolution of ornithopod dinosaurs. Nature, Land. 239, 464-466.
1973. Redescription of the skull and mandible of Parksosaurus from the Late Cretaceous with
comments on the family Hypsilophodontidae (Ornithischia). Contr. Life Sci. Div. R. Out. Mus. 89,
1-21.
1974a. The ornithischian dinosaur Hypsilophodon from the Wealden of the Isle of Wight. Bull. Br.
Mus. (nat. Hist.) Geol. 25, 1-1 52c.
19746. Notes on Thescelosaurus, a conservative ornithopod from the Upper Cretaceous of North
America, with comments on ornithopod classification. J. Paleont. 48, 1048-1067.
(in press). The dinosaur Vectisaurus valdensis (Ornithischia: Iguanodontidae) from the Lower
Cretaceous of England. Ibid.
and JENSEN, J. A. 1973a. Small bones of the hypsilophodontid dinosaur Dryosaurus altus from the
Upper Jurassic of Colorado. Gt Basin Nat. 33, 129-132.
19736. Skeleton of a hypsilophodontid dinosaur (Nannosaurus (?) rex) from the Upper Jurassic
of Utah. Brigham Young Univ. Geol. Stud. 20, 135-157.
GILMORE, c. w. 1909. Osteology of the Jurassic reptile Camptosaurus with a revision of the species of the
genus, and descriptions of two new species. Proc. U.S. natn. Mus. 36, 197-332.
1924a. A new species of Laosaurus, an ornithischian dinosaur from the Cretaceous of Alberta. Trans.
Roy. Soc. Can. (3) 18, 4, 1-6.
19246. On Troddon validus an orthopodous dinosaur from the Belly River Cretaceous of Alberta,
Canada. Bull. Dept. Geol. Univ. Alberta, 1, 1-43.
1925. Osteology of ornithopodous dinosaurs from the Dinosaur National Monument, Utah. Mem.
Carnegie Mus. 10, 385-410.
HULKE, J. w. 1879. Vectisaurus valdensis, a new Wealden dinosaur. Q. Jl geol. Soc., Lond. 35, 421-424.
HUXLEY, T. H. 1869. On Hypsilophodon, a new genus of Dinosauria. Abstr. Proc. geol. Soc. Lond. 204, 3-4.
1870. On Hypsilophodon foxii, a new dinosaurian from the Wealden of the Isle of Wight. Q. Jl geol.
Soc. Lond. 26, 3-12.
JANENSCH, w. 1955. Der Ornithopoda Dysalotosaurus der Tendaguru-Schichten. Palaeontographica,
Suppl. 7, E.R. 3, 105-176.
LYDEKKER, R. 1888. Catalogue of the fossil Reptilia and Amphibia in the British Museum {Natural History),
Part 1 . London, 309 pp.
1889. On the remains and affinities of five genera of Mesozoic reptiles. Q. Jlgeol. Soc. Lond. 45, 41-59.
1890. Catalogue of the fossil Reptilia and Amphibia in the British Museum {Natural History). Part 4,
London, 295 pp.
1891. On certain ornithosaurian and dinosaurian remains. Ibid. 47, 41-44.
MANTELL, G. A. 1827. Illustrations of the geology of Sussex, with figures and descriptions of the fossils of
Tilgate Forest. London, 92 pp.
MARSH, o. c. 1878. Principal characters of American Jurassic dinosaurs. Am. J. Sci. 16, 411-416.
1894. The typical Ornithopoda of the American Jurassic. Ibid. 48, 86-90.
1896. The dinosaurs of North America. Rep. U.S. geol. Surv. 16, 133-244.
MARYANSKA, T. and OSMOLSKA, H. 1974. Pachycephalosauria, a new suborder of ornithischian dinosaurs.
Palaeont. pol. 30, 45-102.
NEWMAN, B. H. 1968. The Jurassic dinosaur Scelidosaurus harrisoni, Owen. Palaeontology, 11, 40-43.
OWEN, R. 1861. The fossil Reptilia of the Liassic Formations. Part 1. Palaeontogr. Soc. {Monogr.), 1-14.
1863. The fossil Reptilia of the Liassic Formations. Part 2. Ibid. 1-26.
PARKS, w. A. 1926. Thescelosaurus warreni a new species of orthopodous dinosaur from the Edmonton
Formation of Alberta. Univ. Toronto Stud. geol. Ser. 21, 1-42.
POMPECKJ, J. F. 1920. Das angebliche Vorkommen and Wandern des Parietalforamens bei Dinosauriern.
Sber. Ges. naturf. Freunde Berl. 1920, 109-129.
RixoN, A. E. 1968. The development of the remains of a small Scelidosaurus from a Lias nodule. Museums J.
67, 315-327.
752
PALAEONTOLOGY, VOLUME 18
STERNBERG, c. M. 1940. Thescelosaurus edmontensis n. sp., and the classification of the Hypsilophodontidae.
J. Paleont. 4, 481-494.
swiNTON, w. E. 1936. The dinosaurs of the Isle of Wight. Proc. Geol. Ass. 47, 204-220.
THULBORN, R. A. 1970. The systematic position of the Triassic ornithischian dinosaur Lycorhinus angustidens.
Zool. J. Linn. Soc. 49, 235-245.
1971. Origins and evolution of ornithischian dinosaurs. Nature, Land. 234, 75-78.
1972. The post-cranial skeleton of the Triassic ornithischian dinosaur Fabrosaurus australis. Palaeon-
tology, 15, 29-60.
1974. A new heterodontosaurid dinosaur (Reptilia; Ornithischia) from the Upper Triassic Red Beds
of Lesotho. Zool. J. Linn. Soc. 55, 151-175.
Original typescript received 18 October 1974
Revised typescript received 16 January 1975
P. M. GALTON
Department of Biology
University of Bridgeport
Bridgeport, Conn. 06602 U.S.A.
BIBLIOGRAPHY AND INDEX OF CATALOGUES
OF TYPE, FIGURED, AND CITED FOSSILS IN
MUSEUMS IN BRITAIN
by MICHAEL G. BASSETT
Abstract. Published (and some unpublished) information on the distribution of type, figured, and cited fossils in
museums in Great Britain and Ireland is collated in a bibliography as an initial aid in tracing type collections and
individual specimens. The catalogues are indexed taxonomically, stratigraphically, and by museums. A supplementary
reference list draws attention to some further publications which may be useful in locating old collections.
. . . Of what advantage was it to science that, when Dr Otto Jaekel was writing his admirable memoir on the Devonian
crinoids of Germany, all the type specimens described by Schultze in his "Echinodermen des Eifler Kalkes’ were locked
up in dusty boxes in a store room at Harvard? . . .
F. A. BATHER. 1897. Science, New Ser. 5, 695.
. . . The value of all types and figured specimens, and the necessity for their safe preservation are now duly recognised.
The recognition has come none too soon. Specialists in particular have to regret the disappearance of many of the types
figured by older authors. And the more doubtful the identification of a species, the more is the disappearance of the type
to be regretted, and the greater would be its value if it could be recovered. . . .
S. S. BUCKMAN. 1899. Proc. Cotteswold Nat. Eld Club, 13, 133.
At the fifty-ninth annual meeting of the British Association for the Advancement of
Science held at Newcastle upon Tyne in September 1889, a Committee was appointed
‘To consider the best methods for the registration of all Type Specimens of Fossils
in the British Isles, and to report on the same’ {Rep. Br. Ass. Advmt Sci. 1 890, p. Ixxxiv).
The following year the Committee gave details of a recording form which they
recommended should be sent to the curators of all museums and owners of private
collections, and at the meeting for 1891 they were able to report that ‘several valuable
lists have already been received’. Progress in the gathering of this information was
reported briefly and intermittently at subsequent meetings of the Association, up to
that of 1903, after which the Committee appears to have become defunct, although
there is no record of it being formally disbanded. Unfortunately the data accumulated
as a result of this exercise were never collated, and a great deal of information on the
whereabouts of many type specimens remained unpublished, notably those in private
collections.
However, in response to the stimulus generated by the British Association survey,
and partly as a result of the direct influence of some members of the Committee,
a number of museums did publish their own catalogues of type and figured specimens.
In some cases the inventories have subsequently been revised and/or expanded from
time to time, and other institutions have since also produced catalogues of all, or
specialized parts of their collections. Together with a few earlier, nineteenth-century
publications, which include information on type specimens, these catalogues form
the main basis of this bibliography.
In 1967 a similar compilation on a world-wide scale was attempted by the
[Palaeontology, Vol. 18, Part 4, pp. 753-773.]
754
PALAEONTOLOGY, VOLUME 18
International Council of Museums (I.C.O.M.), to cover both zoological and
palaeontological collections. This resulted in the publication in 1968 of A preliminary
list of catalogues of type specimens in zoology and palaeontology (30 pp., compiled by
A. W. F. Banfield, published by the State Committee of Culture and Art on the
occasion of the 20th anniversary of the I.C.O.M., Bucharest, Romania, in French
and English). This list contains only thirty-seven references to palaeontological
collections for the whole of the world, with just eight from Britain, and has a limited
index; it thus provides little guidance to the distribution of type-fossil specimens in
British museums, a factor which partly prompted the present compilation.
In modern systematic palaeontological literature it is standard practice to quote
details of the repositories and registration numbers of type, figured, and individually
cited specimens; indeed, most journals rightly insist that this information should be
included as standard, in accordance with recommendations made by the International
Commissions on Zoological and Botanical Nomenclature. Such practice ensures
that specimens will be readily traceable in the future, but it is a relatively recent
innovation and a vast bulk of past publications conspicuously lacks these basic data.
It is frequently difficult or impossible, therefore, to determine from the primary
literature the present whereabouts of old type or figured specimens which may be
essential for revisionary studies of some fossil groups, or important for comparative
purposes, especially where those specimens are not housed in major museums, and
it is all too easy to regard old material as ‘lost’. Yet the published catalogues of type
specimens contain a great deal of information on individual fossils and collections
described in the past, which have fortunately found their way into museums ; a number
refer to small institutions or are published in local journals which may be unfamiliar
to many individuals. The time-consuming effort of tracking down old collections can
often be solved simply by referring to these catalogues, and the primary aim of this
bibliography is to draw the attention of palaeontologists to the published lists as an
initial aid in such a search.
Of course many old type and figured specimens are genuinely lost, but it seems
certain too that many others exist unknowingly in public and private collections. The
responsibility for tracing old type material in any systematic study rests very much
with the individual, but there are limits to the extent that anyone can go in ensuring
beyond all doubt that particular specimens no longer exist. These limits would be
reduced significantly if all institutions were to accept their share of responsibility in
checking collections for type material, and to ensure that details of such material
are widely publicized ; this institutional responsibility is best summarized by Recom-
mendation 72D of the International Code of Zoological Nomenclature, which
states that :
Every institution in which types are deposited should :
( 1 ) ensure that all are clearly marked so that they will be unmistakably recognized ;
(2) take all necessary steps for their safe preservation;
(3) make them accessible for study;
(4) publish lists of type-material in its possession or custody; and
(5) so far as possible, communicate information concerning types when requested
by zoologists.
BASSETT: TYPES OF FOSSILS
755
Unfortunately it is a sad fact that many institutions and individuals are unaware
of, or neglect these recommendations with the result that some type material can still
become mislaid or lost. Any such museum should carefully heed the following advice
of D. E. Owen concerning the care of type specimens (1964, Mus. J. 63, 288-291).
It is a prerequisite that such a museum must have a suitably qualified member of the
staff always in charge of the types. For instance a museum with fossil types must have
a geologist on the staff who will be replaced by another geologist if he leaves. This is
even more important with perishable specimens which require regular technical treat-
ment. The small museum that may be under the care of a geologist for a few years, an
archaeologist next, and then an art expert, had much better place its types in more
permanent hands. The University department with types but no permanent curator, had
much better place these types in an institution whose staff are appointed primarily to
care for the specimens.
Owen also stresses that The publishing of a list of type and figured specimens in the
collections must be the aim of every museum holding such specimens, and efforts should
be made to keep this up to date. The specialist, studying a group, usually has great
difficulty locating types, and such lists are invaluable.
Strict attention to all these comments would ensure that essential specimens are
both housed properly and brought to the attention of palaeontologists as a whole.
One of the aims of the recently constituted Geological Curators Group is to trace
type fossil specimens in museums in Britain, particularly those which have no
permanent geological staff to uphold the responsibilities outlined above. Where
necessary the Group will publish further catalogues of type material in its Newsletter,
to add to those cited here.
BIBLIOGRAPHY
The format and content of the catalogues listed here varies considerably. Ideally they
are published inventories of individual fossil specimens, with information on the
repository, museum registration numbers, type data (where applicable), and details
of page, plate, and figure numbers in a previous publication referring to those
individual specimens; in the comparatively few cases where all these details are not
included, the information that is given will generally allow an individual specimen
to be identified. The bibliography specifically excludes many museum ‘Catalogues’
which are published as systematic monographs of particular fossil groups in the
collections. The best known of these are the many monographic Catalogues published
by the British Museum (Natural History), which will be familiar to specialists working
on a particular fossil group. However, where such catalogues do give references to
type or figured specimens in addition to those described systematically, they are listed
here. Also excluded are the many Guides to displays of fossils in museum galleries,
together with straightforward inventories of collections which contain no specific
data on type, figured, or cited specimens. Unpublished, manuscript lists are included
only where they have been widely distributed by their authors, or are available in the
institutions to which they refer.
Information in square brackets after some of the references draws attention to
changes in the names or locations of some institutions, and to cases where specimens
756
PALAEONTOLOGY, VOLUME 18
are known to have been transferred to different institutions. In this supplementary
information The Geological Museum of the Institute of Geological Sciences is referred
to as IGS, London, the regional offices of the Institute are referred to as IGS, Leeds
and Edinburgh, and the British Museum (Natural History) as BM(NH).
ALLEN, H. A. 1900. Catalogue of types and figured specimens from the Eocene and Oligocene Series pre-
served in the Museum of Practical Geology. Summ. Progr. geol. Surv. Land, for 1899, 195-208. [Speci-
mens now in IGS, London.]
1901u. Catalogue of types and figured specimens from British Pliocene and Pleistocene strata pre-
served in the Museum of Practical Geology, London. Ibid, for 1900, 182-195. [Specimens now in IGS,
London.]
19016. Catalogue of types and figured specimens from British Devonian strata preserved in the
Museum of Practical Geology, London. Ibid. 196-216. [Specimens now in IGS, London.]
1902a. Catalogue of types and figured specimens of British fossil Phyllocarida preserved in the Museum
of Practical Geology, London. Ibid, for 1901, Appendix A, 200-203. [Most specimens now in IGS,
London; Carboniferous specimens in IGS, Leeds.]
19026. Catalogue of types and figured specimens of British Palaeozoic Echinodermata preserved in
the Museum of Practical Geology, London. Ibid. Appendix B, 204-211. [Most specimens now in IGS,
London; Carboniferous specimens in IGS, Leeds.]
1903. Catalogue of types and figured specimens of British Gasteropoda and Scaphopoda from the
Rhaetic beds. Lias and Inferior Oolite, preserved in the Museum of Practical Geology, London. Ibid,
for 1902, 217-228. [Specimens now in IGS, London.]
1904. Catalogue of the types and figured specimens of British Gasteropoda and Scaphopoda from the
Lower, Middle and Upper Oolites, preserved in the Museum of Practical Geology, London. Ibid, for
1903, 175-187. [Specimens now in IGS, London.]
1905. Catalogue of types and figured specimens of British Lamellibranchiata from the Rhaetic beds
and Lias, preserved in the Museum of Practical Geology, London. Ibid, for 1904, 172-177. [Specimens
now in IGS, London.]
1906. Catalogue of types and figured specimens of British Lamellibranchiata from the Lower, Middle
and Upper Oolites, preserved in the Museum of Practical Geology. Ibid, for 1905, 175-195. [Specimens
now in IGS, London.]
1915. Catalogue of types and figured specimens of British Cretaceous Lamellibranchiata preserved
in the Museum of Practical Geology, London. Ibid, for 1914, 66-79. [Specimens now in IGS, London.]
1916. Catalogue of types and figured specimens of British Cretaceous Gasteropoda preserved in the
Museum of Practical Geology, London. Ibid, for 1915, 47-51. [Specimens now in IGS, London.]
ANDERSON, E. M. 1936. Catalogue of types and figured specimens of fossils in the Geological Survey collections
now exhibited in The Royal Scottish Museum, Edinburgh. 1-77, H.M.S.O., London. [Specimens now in
IGS, Edinburgh.]
ANON. 1896. Museum Sub-Committee. Report for 1894-1895. In Rep. Brighton publ. Mus. for 1894-1895,
3-7. [Includes note on type specimens added to the collections.]
1957. Index to collection of sections and preparations of fossil plants. John Walton collection. [Typed
MS., 25 pp. (numbered 1-12 only); specimens formerly in Department of Botany, University of Glasgow,
now in Department of Geology, Hunterian Museum, from where copies of the catalogue are available;
the manuscript is undated, but is here referred for convenience to 1957 since the latest paper quoted
is 1956.]
APPLEBY, R. M. 1958. A Catalogue of the Ophthalmosauridae in the collections of the Leicester and Peterborough
Museums. 1-47, pis. 1-7, Leicester Museums and Art Gallery, Department of Geology.
BASSETT, M. G. 1972. Catalogue of type, figured and cited fossils in the National Museum of Wales. 1-113,
The National Museum of Wales, Cardiff.
BATHER, F. A. 1899. The genera and species of Blastoidea, with a list of the specimens in the British Museum
{Natural History), i-x, 1-70, British Museum (Natural History), London.
BELL, A. 1917. A list of type and figured specimens in the Geological Gallery, Ipswich Museum. J. Ipswich
Distr. nat. Hist. Soc. 5 [for 1916], 41-49. [Also reprinted verbatim (1917) by the Ipswich Museum, with
emended pagination, 1-11.]
BASSETT: TYPES OF FOSSILS
757
BLAKE, J. F. 1902. List of the types and figured specimens recognised by C. D. Sherborn, F.G.S., in the col-
lection of the Geological Society of London. Verified and arranged, with additions, by J. F. Blake, M.A.,
F.G.S. {with an appendix). 1-100, i-xxxii. Geological Society, London. [British specimens now in IGS,
London and Leeds, foreign specimens in BM(NH), to which institutions they were transferred in
1911.]
BOLTON, H. 1892. A catalogue of types and figured specimens contained in the Geological Department of
the Manchester Museum, Owens College. Rep. Proc. Mus. Ass. 96-129.
1894. Supplementary list of type and figured specimens in the Geological Department, Manchester
Museum, Owens College. Ibid. 250-254.
[BRIGHTON, A. G.j [1954]. List of ammonites in Sedgwick Museum, fig’d by S. Buckman 1886-1907. Mon.
Pal. Soc. Amm. Inf. Oolite. [Typed MS., 7 pp., undated but approximately 1954 (fide H. S. Torrens).]
BUCKMAN, s. s. 1899. List of types and figured specimens of Brachiopoda. Proc. Cotteswold Nat. Fid Club,
13 (2), 133-141.
[1929]. [Catalogue of the S. S. Buckman collection.] [Handwritten MS. list compiled by Buckman
between about 1 880 and 1 928, now in the BM(NH), bound in a single ledger ; undated, but for convenience
referred here to 1929 since that was the date that part of the collection, together with the catalogue, was
sold to the BM(NH). Collection now broken up and housed in a number of museums both in Britain and
abroad, of which those in Britain are known to include at least the BM(NH), IGS, London, Sedgwick
Museum, The Manchester Museum, Oxford University Museum; some specimens acquired by the City
Museum, Bristol were destroyed in November 1940.]
CALDER, M. G. 1959. Catalogue of the Kidston collection of sections of fossil plants in the Department of Botany
of the University of Glasgow. [Typed MS., 1 14 pp., based on an unpublished catalogue compiled between
1933 and 1936; copies available from the Hunterian Museum, University of Glasgow, where all the
specimens are now stored in the Department of Geology.]
CANTRiLL, T. c., DIXON, E. E. L., THOMAS, H. H. and JONES, o. T. 1916. A list of types and figured specimens
from Sheet 227 in the Survey and Museum Collections. Appendix III, P- 176, in The geology of the South
Wales coal-field. Part 12, The country around Milford. Mem. geol. Surv. U.K. i-vii, 1-185, pis. 1-7.
[Most specimens now in IGS, London; Carboniferous specimens in IGS, Leeds.]
CARRECK, J. N. 1955. The Quaternary vertebrates of Dorset, fossil and subfossil. Proc. Dorset nat. Hist,
archaeol. Soc. 75 [for 1953], 164-188.
COLE, w. w. see Enniskillen, earl of.
cox, L. R. and ARKELL, w. J. 1948-1950. A survey of the Mollusca of the British Great Oolite Series: primarily
a nomenclatorial revision of the monographs by Morris & Lycett (1851-55), Lycett (1863) and Blake
(1905-07). Palaeontogr. Soc. [Monogr.], i-xxiv, 1-105. [Text and revised plate explanations give data
on previously figured specimens.]
CRANE, E. 1892. Catalogue of types and figured specimens now in the Brighton Museum. In Rep. Brighton
publ. Mus. for 1891-1892, Appendix B, 9-20.
1893. Museum Sub-committee. Report for 1892-3. In Ibid, for 1892-1893, 5-8. [Contains note on
type specimens added to the collections.]
CRICK, G. c. 1 898. List of the types and figured specimens of fossil Cephalopoda in the British Museum (Natural
History). 1-103, British Museum (Natural History), London.
1922. Notes on specimens of Cephalopoda figured in Tate and Blake’s ‘Yorkshire Lias’, 1876.
Naturalist, Aug.-Sept., 273-288. [Specimens in BM(NH).]
CURRIE, E. D. and george, t. n. 1963. Catalogue of described and figured specimens in the Begg Collection
in the Hunterian Museum of the University of Glasgow. Palaeontology, 6, 378-396.
CURTIS, M. L. K. 1956. Type and figured specimens from the Tortworth Inlier, Gloucestershire. Proc. Bristol
Nat. Soc. 29, 147-154.
[1970]. [Bristol City Museum list of specimens figured in I909-1930''Type Ammonites' by S. S. Buckman.]
[Handwritten MS., 3 pp., provided by M. L. K. Curtis for H. S. Torrens; undated but approximately
1970 (fide H. S. Torrens).]
[CUTBILL, J. L.] 1973. Sedgwick Museum catalogue, H Section [Scn'ei] Devonian. [Computer printed catalogue
dated 31 January 1973, comprising: H Series in numerical order (4 vols., 1-930), locality index (2 vols.,
1-307), taxonomic index (2 vols., 1-182), stratigraphic index (1 vol., iv -1-1-145), personal names index
(1 vol., 1-38), bibliographic index (1 vol., 1-36). Locality index subsequently revised slightly (pp. 1-306)
and both vols. reissued on 27 July 1973, but in original covers.]
758 PALAEONTOLOGY, VOLUME 18
DAVIES, w. 1871a. Alphabetical catalogue of type specimens of fossil fishes in the British Museum. Geol.
Mag. 8, 208-216.
18716. Supplementary list of type specimens of fossil fishes in the British Museum. Ibid. 334-
335.
DELAIR, j. B. 1966a. Fossil footprints from Dumfriesshire, with descriptions of new forms from Annandale.
Trans. J. Proc. Dumfries. Galloway, nat. Hist. Antiq. Soc. 43, 14-30.
19666. Catalogue of the fossil vertebrates in the Museum and Art Gallery, Paisley. [Typed MS., 23 pp.,
dated by author February 1966.]
[delair, j. b.] 1966c. A catalogue of the vertebrate fossils in Kilmarnock Museum. [Typed MS., 22 pp., dated
by author November 1966.]
DONOVAN, d. t. 1954. Synoptic supplement to T. Wright’s ‘Monograph on the Lias ammonites of the British
Islands’ ( 1 878-86). Palaeontogr. Soc. [Monogr.], 1-54. [Revised plate explanations give data on previously
figured specimens.]
DOUGHTY, p. 1974. Collections or information currently sought: 7. Captain R. B. Bennett. Newsletter of
the Geological Curators Group, No. 2, 68-69. [Includes note on a figured Carboniferous bivalve in Ulster
Museum.]
EDMONDS, J. M. 1949. Types and figured specimens of Lower Palaeozoic Trilobites in the University Museum,
Oxford. Geol. Mag. 86, 57-66.
EGERTON, P. G. 1836. Catalogue of fossil fish, in the collections of Lord Cote and Sir Philip Grey Egerton,
arranged alphabetically, with references to the localities, strata, and published description of the species.
13 pp., J. Seacombe, Chester. [Specimens now in BM(NH).]
[egerton, p. m. g.] 1869. Alphabetical catalogue of type specimens of fossil fishes in the collection of Sir
Philip de Malpas Grey Egerton, Bart., M.P., at Oulton Park. Geol. Mag. 6, 408-413. [Also published
separately (1869) with emended pagination, 1-10. Specimens now in BM(NH).]
ENNISKILLEN, EARL OE [w. w. COLE]. 1869. Alphabetical catalogue of the type specimens of fossil fishes in
the collection of the Earl of Enniskillen, at Florence Court. Ibid. 556-561. [Also published separately
(1869) with emended pagination, 1-9. Specimens now in BM(NH).]
[GREGORY, J. w.] 1928. University of Glasgow, Hunterian Museum Geological Department, 1-12. [Pamphlet
giving brief history of the collections, including some type and figured specimens.]
HALLAM, A. D. 1937. Report on the geological collections in the Somerset County Museum. Proc. Somerset
archaeol. nat. Hist. Soc. 82 [for 1936], 62-66.
HENRICHSEN, I. G. c. 1970. A Catalogue of fossil vertebrates in the Royal Scottish Museum, Edinburgh.
Part One. Actinopterygii. Royal Scottish Museum Information Series. Geology, 1, i-x, 1-102.
1971 . A catalogue of fossil vertebrates in the Royal Scottish Museum, Edinburgh. Part Two Agnatha.
Ibid. 2, i-vi, 1-38.
1972. A catalogue of fossil vertebrates in the Royal Scottish Museum, Edinburgh. Part Three/
Actinistia and Dipnoi. Ibid. 3, i-vi, 1-26.
HOPPING, c. A. 1957. Catalogue of fossil plants in the Hunterian Museum of the University of Glasgow
[with a foreword by J. Walton]. [Typed MS., 390 pp., copies available from the Hunterian
Museum.]
HOWARTH, M. K. 1962. The Yorkshire type ammonites and nautiloids of Young and Bird, Phillips, and
Martin Simpson. Palaeontology, 5, 93-136, pis. 13-19.
HOWSE, R. 1888. Contributions towards a catalogue of the flora of the Carboniferous System of Northumber-
land and Durham. Part 1.— Fossil Plants from the Hutton Collection. Catalogue of those specimens of
the Hutton Collection of fossil plants that have been presented to the Natural History Society by the
Council of the Mining Institute, and are now exhibited in the Geological Room of the Museum, at Barras
Bridge, Newcastle-upon-Tyne. Nat. Hist. Trans. Northumb. 10, 19-151, pis. 1-6. [Also reprinted verbatim
(1888) with emended pagination, 1-135. Specimens now in The Hancock Museum, Newcastle upon
Tyne.]
JACKSON, J. w. 1952. Catalogue of types and figured specimens in the geological department of the Man-
chester Museum. Manchester Museum Publication, No. 6, i-vii, 1-170.
JONES, T. R. 1882. Catalogue of the fossil Foraminifera in the collection of the British Museum {Natural
History), Cromwell Road, S.W. i-xxiv, 1-100. British Museum (Natural History), London.
JUKES-BROWN, A. J. and ELSE, w. J. 1907. A list of the type fossils and figured specimens in the Museum of
the Torquay Natural History Society. Rep. Trans. Devon. Advmt Sci. 39, 399^09.
BASSETT: TYPES OF FOSSILS
759
LANG, w. D. 1947. James Harrison of Charmouth, geologist (1819-1864). Proc. Dorset not. Hist, archaeol.
Soc. 68 [for 1946], 103-118. [Includes list of specimens from Harrison’s collection purchased by BM(NH)
in 1865.]
LEBOUR, G. A. 1878. Catalogue of the Hutton Collection of fossil plants, including a synoptical list of the
chief Carboniferous species not in the Collection. Drawn up by order of the Council of the North of England
Institute of Mining and Mechanical Engineers, i-xii, 1-132, Newcastle upon Tyne. [Specimens now in
The Hancock Museum, Newcastle upon Tyne.]
LENEY, F. 1902. A list of the ‘Type’, figured and described fossils in the Norwich Castle Museum. Geol. Mag.
Dec. 4, 9, 166-171,220-231.
LYDEKKER, R. 1885-1887. Catalogue of the fossil Mammalia in the British Museum (Natural History),
Cromwell Road, S.fV. 5 vols.; Part 1 (1885), i-xxx, 1-268; Part 2 (1885), i-xxii, 1-324; Part 3 (1886),
i-xvi, 1-186; Part 4(1886), i-xxiv, 1-233; Part 5 (1887), i-xxxv, 1-345. British Museum (Natural History),
London.
1888-1890. Catalogue of the fossil Reptilia and Amphibia in the British Museum (Natural History),
Cromwell Road, S.W. 4 vols.; Part 1 (1888), i-xxviii, 1-309; Part 2 (1889), i-xxi, 1-307; Part 3 (1889),
i-xviii, 1-239; Part 4 (1890), i-xxiii, 1-296. British Museum (Natural History), London.
1891. Catalogue of the fossil birds in the British Museum (Natural History), Cromwell Road, S.W.
i-xxvii, 1-368. British Museum (Natural History), London.
MCHENRY, A. and WATTS, w. w. 1898. Guide to the collections of rocks and fossils belonging to the Geological
Survey of Ireland, arranged in the curved gallery of the Museum of Science and Art, Dublin. 1-155, H.M.S.O.,
Dublin. [Section 3, pp. 120-127, figured and type specimens of fossils.]
MELMORE, s. 1945-1946. Catalogue of types and figured specimens in the Geological Department of the
Yorkshire Museum. N-West Nat. [in three parts], 207-221 [1945], 72-91, 234-245 [1946]. [Also reissued,
verbatim in one volume with original pagination, by The Yorkshire Museum.]
MITCHELL, M. and WHITE, D. E. 1966. Catalogue of figured, described and cited Carboniferous corals in
the collections of the Geological Survey and Museum, London. Bull. geol. Surv. Gt Br. 24, 19-56.
[Specimens now in IGS, Leeds.]
MORELLET, L. and MORELLET, J. 1939. Tertiary Siphoneous Algae in the W. K. Parker Collection with descrip-
tions of some Eocene Siphoneae from England, i-xi, 1-55, Pis. 1-6, British Museum (Natural History),
London.
[morris, j. and owen, r.] 1856. Descriptive catalogue of the fossil organic remains of Invertebrata contained
in the Museum of the Royal College of Surgeons of England, i-vi, 1-260, Taylor and Francis, London.
NEA VERSON, E. 1950. The foundation of the University geological collection. Proc. Lpool geol. Soc. 20,
149-157. [Includes notes on specimens in Liverpool University, many of which have since been trans-
ferred to BM(NH).]
NEWTON, R. B. 1891. Systematic list of the Frederick E. Edwards collection of British Oligocene and Eocene
Mollusca in the British Museum (Natural History), with references to the type-specimens from similar
horizons contained in other collections belonging to the Geological Department of the Museum, i-xxviii,
1-365, British Museum (Natural History), London.
1902. List of Thomas Say’s types of Maryland (U.S.) Tertiary Mollusca in the British Museum. Geol.
Mag. Dec. 4, 9, 303-305.
NORTH, F. J. 1928. Type and figured fossils in the National Museum of Wales. Ibid. 65, 193-210. [Reprinted
verbatim (1928) by the National Museum of Wales with emended pagination, 1-20.]
OWEN, R. 1845. Descriptive and illustrated catalogue of the fossil organic remains of Mammalia and Aves
contained in the Museum of the Royal College of Surgeons of England, i-vii, 1-391, pis. 110. Taylor and
Francis, London.
PATON, R. L. 1975. A catalogue of fossil vertebrates in the Royal Scottish Museum, Edinburgh. Part Four,
Amphibia & Reptilia. Royal Scottish Museum Information Series. Geology. 5, i-ix, 1-38.
PLATNAUER, H. M. 1891. List of figured specimens in York Museum. Rep. Yorks, phil. Soc. [for
1890], 56-89.
1894. Appendix to the list of figured specimens in the Museum of the Yorkshire Philosophical Society.
Ibid, [for 1893], 45-56.
PYRAH, B. J. In Press. Catalogue of type and figured material in the geological collections of the Yorkshire
Museum. Part 1. Porifera, Coelenterata, Bryozoa, Annelida, ‘Unknown’, Brachiopoda, Crustacea,
Insecta. Proc. Yorks, geol. Soc.
760
PALAEONTOLOGY, VOLUME 18
RILEY, T. H. 1974. Type specimens in the palaeontological collections of Sheffield City Museums, England.
Newsletter of the Geological Curators Group, No. 2, 36-31.
ROWE, F. w. E. 1974. Catalogue of the Sladen Collection in the Royal Albert Memorial Museum, Exeter,
Devon. Biol. J. Linn. Soc. 6, 179-243, pis. 1-3. [Deals mainly with one of the most important collections
of living and fossil echinoderms ever made, of which only the type and figured fossils are indexed here;
there is also brief mention of the W. B. Carpenter Foraminifer Collection, which includes type fossil
material.]
SALTER, J. w. 1873. A catalogue of the collection of Cambrian and Silurian fossils contained in the geological
museum of the University of Cambridge. With a preface by The Rev. Adam Sedgwick, Ll.D., F.R.S.
Woodwardian Professor of Geology in the University of Cambridge, and a table of genera and index added
by Professor Morris, F.G.S. i-xlviii, 1-204, University Press, Cambridge.
[SAMUEL, E. M.] [1970]. Type specimens from the Jurassic [Dorset County Museum]. [Typed MS., 2 pp.,
undated but approximately 1970 {fide H. S. Torrens).]
SANFORD, w. A. 1869. Catalogue of the feline fossils in the Taunton Museum. Proc. Somerset archaeol. nat.
Hist. Soc. 14 [for 1867], 103-160. [Also published separately by the Society (? 1869) in large format,
together with 26 plates.]
SEELEY, H. G. 1869. Index to the fossil remains of Aves, Ornithosauria, and Reptilia, from the Secondary
System of strata arranged in the Woodwardian Museum of the University of Cambridge. With a prefatory
notice by the Rev. Adam Sedgwick, LL.D., F.R.S. Woodwardian Professor and Senior Fellow of Trinity
College, i-xxiii, 1-143, Deighton, Bell and Co. Cambridge. [Now Sedgwick Museum.]
SEWARD, A. c. 1894. Notes on the Bunbury Collection of fossil plants with a list of type specimens in the
Cambridge Botanical Museum. Proc. Cambridge Phil. Soc. 3, 187-198.
1900. Notes on some Jurassic plants in the Manchester Museum. Mem. Proc. Manchr lit. phil. Soc.
44, 1-28, pis. 1-4. [Also issued separately as Notes from the Manchester Museum, No. 6; includes short
list of type and figured specimens.]
siME, I. F. 1972. A catalogue of Carboniferous corals in the Royal Scottish Museum, Edinburgh. Royal
Scottish Museum Information Series. Geology, 4, i-x, 1-72.
SIZER, c. A. 1962. A catalogue of the figured and cited specimens in the Department of Geology. 1-46, 1 pi.,
Leicester Museums and Art Gallery, Department of Geology.
STRACHAN, I. 1971. A synoptic supplement to ‘A monograph of British graptolites by Miss G. L. Elies and
Miss E. M. R. Wood’. Palaeontogr. Soc. [Monogr.], 130 pp. [Revised plate explanations give data on
previously figured specimens.]
STRAHAN, A., CANTRiLL, T. c., DIXON, E. E. L., THOMAS, H. H. and JONES, o. T. 1914. A list of type and figured
specimens from Sheet 228 in the Survey and Museum collections. Appendix III, pp. 248-249. In The
geology of the South Wales eoalfield. Part 11, The country around Haverfordwest. Mem. geol. Surv.
U.K. i-viii, 1-249. [Specimens now in IGS, London.]
STUBBLEFIELD, c. J. 1936. Notes on the types and figured specimens acquired from the late S. S. Buckman
by the Geological Survey of Great Britain. Summ. Progr. geol. Surv. Lond. for 1934, Pt. 2. 52-59.
[Specimens now in IGS, London.]
1938. The types and figured specimens in Phillips and Salter’s Palaeontological Appendix to John
Phillips’ Memoir on ‘The Malvern Hills compared with the Palaeozoic districts of Abberley, etc.’ (Mem.
Geol. Surv. Volume II, Part 1, June 1848.) Summ. Progr. geol. Surv. Lond. for 1936, Pt. 2, 27-51. [Speci-
mens now in IGS, London.]
THOMPSON, B. 1929. Obituary. Mr. Thomas Jesson, B.A., F.G.S. J. Northampt. nat. Hist. Soc. 25,
49-50. [Includes reference to publications based on Jurassic ammonites and fishes in Northampton
Museum.]
and GEORGE, T. J. A catalogue of the geological collection in the Northampton Museum.
Part 1. The Silurian System. Ibid. 3, 39-46, 1 pi. [Includes note (p. 36) referring to one
figured crinoid.]
TORRENS, H. s. 1974. Collections and information: lost and found: A. Collections previously sought:
5. Wyville Thomson. Newsletter of the Geological Curators Group, No. 2, 67-68. [Figured Ordovician
and Silurian trilobites in Oxford University Museum.]
In press. Type, figured and cited fossils formerly in the Sherborne School Museum. BM(NH) = British
Museum (Natural History) collection. Proc. Dorset nat. Hist, archaeol. Soc. 96. [Most specimens now
in BM(NH), one ammonite in Oxford University Museum.]
BASSETT: TYPES OF FOSSILS
761
TUTCHER, J. w. [1937], Lists of the types and figured specimens of British fossil organisms in the collection of
J. W. Tutcher, Bristol. [Handwritten MS., 35 pp., original in BM(NH); undated, but for convenience
referred here to 1937 since that was the date that the collection, together with the catalogue, was sold to
the BM(NH).]
WATERSTON, c. D. 1954. Catalogue of type and figured specimens of fossil fishes and amphibians in the
Royal Scottish Museum, Edinburgh. Trans. Edinb. geol. Soc. 16, i-x, 1-91.
[ ] 1968a. A list of specimens in the Forres Museum: Altyre and other collections. [Typed MS., 2 pp.,
dated 1968. All type and figured specimens since purchased by Royal Scottish Museum, Edinburgh.]
[ ] 19686. Specimens of fossil fishes described and figured in Elgin Museum. [Typed MS., 2 pp., dated
1968.]
1968c. List of specimens in the Elgin Museum. [Typed MS., 2 pp., dated 1968 ]
WILSON, E. 1890. Fossil types in the Bristol Museum. Geol. Mag. Dec. 3, 7, 363-372, 411-416.
[wiNWOOD, H. H. and wilson, e.] 1892. Charles Moore, F.G.S. and his work; with a list of the fossil types
and described specimens in the Bath Museum. Proc. Bath nat. Hist, antic/. Eld Club, 7, 232-292.
WOODS, H. 1891. Catalogue of the type fossils in the Woodwardian Museum, Cambridge, i-xvi, 1-180, Uni-
versity Press, Cambridge. [Now Sedgwick Museum.]
1893. Additions to the type fossils in the Woodwardian Museum. Geol. Mag. Dec. 3, 10, 111-118.
[Now Sedgwick Museum.]
WOODWARD, A. s. 1 889- 1901. Catalogue of the fossil fishes in the British Museum {Natural History), Cromwell
Road, S.W. 4 vols.; Part 1 (1889), i-xlvii, 1-474, pis. 1-17; Part 2 (1891), i-xliv, 1-567, pis. 1-16; Part 3
(1895), i-xlii, 1-544, pis. 1-18; Part 4 (1901), i-xxxviii, 1-636, pis. 1-19. British Museum (Natural
History), London.
and SHERBORN, c. D. 1890. Catalogue of British fossil Vertebrata. i-xxxv, 1-396, Dulau & Co., London.
WRIGHT, c. w. and wright, e. v. 1951. A survey of the fossil Cephalopoda of the Chalk of Great Britain:
primarily a nomenclatorial revision of Daniel Sharpe’s 'Description of the fossil remains of Mollusca
found in the Chalk of England. Part 1, Cephalopoda’ (1853-1857). Palaeontogr. Soc. [Monogr.], 40 pp.
[Revised plate explanations give data on previously figured specimens.]
WYATT, A. 1974. Geological collections at U.C.W. Aberystwyth. Newsletter of the Geological Curators
Group, No. 2, 65. [University College of Wales, Aberystwyth.]
INDEX
The index is arranged in three sections ;
1 . A taxonomic index in which genera and species listed in individual catalogues
are grouped together in major taxonomic divisions, each of which is then broken
down stratigraphically and cross-referenced to authors. To save repetition and space
the date of a publication is not given after authors’ names in cases where those authors
have only one publication under their name, since those references can be located
immediately in the bibliography; in all other cases both authors’ names and dates
of publications are given. The major taxonomic groupings are generally those which
are employed as headings in most of the catalogues and correspond with classificatory
divisions which will be immediately familiar to palaeontologists. For the inverte-
brates the groupings are initially at the Phylum level, with each Phylum being further
subdivided, generally at the Class or Sub-Class level, wherever those lower categories
are commonly studied as fossil groups. The vertebrates are listed under familiar
Class groups, with the exception that agnathans and gnathostomes are loosely
included together under the single heading Fishes. Plants are listed simply as Plantae
or Algae.
Where authors of catalogues have not themselves separated their specimens into
the groupings adopted here, their genera and species are included as undifferentiated
G
762
PALAEONTOLOGY, VOLUME 18
members of the highest appropriate division listed. The stratigraphical breakdown
within the taxonomic index is as discussed below.
2. A stratigraphical index in which individual specimens listed in catalogues are
grouped stratigraphically, with each stratigraphical division then broken down into
major taxonomic groups corresponding with those outlined above. No reference is
made here to authors since this information can be obtained simply by cross-reference
back to the taxonomic index.
In most cases the stratigraphical horizons given for specimens in the catalogues
have been grouped together in the index within the geological Systems. Exceptions
are made for specimens recorded as coming from undifferentiated Tertiary beds,
which are listed here as such, and for specimens from the Tremadoc, Old Red Sand-
stone, and Rhaetian which are listed separately in order to avoid any confusion which
might arise from assigning the material to one or more Systems.
3. A museums index in which all the museums and institutions recorded in the
catalogues as including type, figured, or cited fossil specimens are listed and cross-
referenced to authors. As in the taxonomic index, dates of publications are given only
in cases in which there is more than one reference by any one particular author.
TAXONOMIC INDEX
INVERTEBRATA
Actinozoa: see Coelenterata.
Ammonoidea ; see Mollusca.
Annelida :
CAMBRIAN, Anderson; Blake; Salter.
ORDOVICIAN, Salter; Woods 1891.
SILURIAN, Blake ; Curtis 1956 ; Salter ; Stubble-
field 1938; Woods 1891.
DEVONIAN, Blake.
CARBONIFEROUS, Anderson; Lebour; Sizer.
RHAETIAN, Sizer.
JURASSIC, Anderson; Blake; Melmore;
Pyrah; Sizer.
CRETACEOUS, Blake; Melmore; Woods 1891.
TERTIARY, Blake.
PLEISTOCENE, Leney; Sizer.
Anthozoa: see Coelenterata.
Arachnida : see Arthropoda.
Arthropoda (undifferentiated);
CAMBRIAN, Anderson; Bolton 1892; North;
Salter.
ORDOVICIAN, Jackson.
SILURIAN, Anderson; Jackson; McHenry
and Watts.
OLD RED SANDSTONE, Andcrson.
DEVONIAN, [Cutbill].
CARBONIFEROUS, Anderson ; Jackson.
JURASSIC, Jackson.
Arachnida :
DEVONIAN, [Cutbill].
CARBONIFEROUS, Bassett.
Crustacea ;
CAMBRIAN, Allen 1902a; Bassett; Bolton
1892; North; Salter; Woods 1891, 1893.
TREMADOC, Bassett.
ORDOVICIAN, Allen 1902a; Bassett; Blake;
North; Woods 1891, 1893.
SILURIAN, Allen 1902a; Bassett; Blake;
Bolton 1892, 1894; North; Salter; Woods
1891, 1893.
DEVONIAN, Allen 19026; [Cutbill]; Jukes-
Browne and Else; Woods 1891.
CARBONIFEROUS, Allen 1 902a ; Bassett ; Blake ;
Bolton 1892; Sizer.
PERMIAN, Blake.
TRiASSic, Blake; Sizer.
RHAETIAN, Blake; Sizer.
JURASSIC, Blake; Lang, Melmore; Platnauer
1891; Pyrah; Sizer; [Winwood and
Wilson]; Woods 1891.
CRETACEOUS, Blake; Crane 1892; Melmore;
Pyrah; Platnauer 1891.
EOCENE, Blake.
TERTIARY, Blake; Pyrah.
PLEISTOCENE, Bell; Leney, Sizer.
BASSETT: TYPES OF FOSSILS
763
Insecta :
CARBONIFEROUS, Bassctt; North.
RHAETIAN, Pyrah; Sizer.
JURASSIC, Blake; Sizer.
TERTIARY, Blake.
Isopoda :
CRETACEOUS, Woods 1891.
Merostomata :
SILURIAN, Bolton 1892; Woods 1891.
CARBONIFEROUS, Bassett.
Trilobita:
CAMBRIAN, Bassett; Blake; Bolton 1892;
Edmonds; North; Salter; Stubblefield
1938; Woods 1891, 1893.
TREMADOC, Bassett; Curtis 1956; Salter.
ORDOVICIAN, Anderson; Bassett; Blake;
Currie and George; Edmonds; Gregory;
Neaverson; Salter; Strachan el al.\
Stubblefield 1938; Torrens 1974; Woods
1891.
SILURIAN, Bassett; Blake; Cantrill et al.;
Currie and George; Curtis 1956;
Edmonds; McHenry and Watts; Salter;
Stubblefield 1938; Torrens 1974; Woods
1891.
DEVONIAN, Blake; Bolton 1892; Jukes-
Browne and Else; McHenry and Watts;
Woods 1891.
CARBONIFEROUS, Currie and George; Riley.
Asteroidea: see Echinodermata.
Belemnoidea : see Mollusca.
Bivalvia: see Mollusca.
Blastoidea : see Echinodermata.
Brachiopoda :
CAMBRIAN, Anderson; Salter; Woods 1891.
TREMADOC, Curtis 1956.
ORDOVICIAN, Bassett; Blake; Cantrill et al.-,
Currie and George; Jackson; Strachan
et al. -, Stubblefield 1938; Woods 1891.
SILURIAN, Bassett; Blake; Bolton 1892;
Cantrill et al. -, Currie and George; Curtis
1956; Jackson; McHenry and Watts;
Salter; Stubblefield 1938; Woods 1891.
OLD RED SANDSTONE, Blake.
DEVONIAN, Allen 1901^; Blake; [Cutbill];
[Gregory]; Jukes-Browne and Else;
Woods 1891.
CARBONIFEROUS, Anderson ; Bassett; Blake;
Currie and George; [Gregory]; Jackson;
McHenry and Watts; Melmore; North;
Platnauer 1891 ; Pyrah; Woods 1891.
PERMIAN, Jackson; Woods 1891.
TRiASSic, [Gregory] ; Sizer.
JURASSIC, Anderson; Bassett; Blake; Buck-
man 1899, [1929]; North; Pyrah; Sizer;
Tutcher; [Winwood and Wilson]; Woods
1891, 1893.
CRETACEOUS, Blake; Jackson; Melmore;
Platnauer 1891; Pyrah; Woods 1891,
1893.
TERTIARY, Blake; Pyrah.
PLEISTOCENE, Leney; Melmore; Platnauer
1891.
Bryozoa :
CAMBRIAN, Salter.
ORDOVICIAN, Bassett; North; Woods 1891.
SILURIAN, Bassett; Blake; Woods 1891.
DEVONIAN, Allen 19016; [Cutbill]; Jukes-
Browne and Else.
CARBONIFEROUS, Anderson ; Jackson ; Woods
1891.
JURASSIC, Melmore; Platnauer 1891; Pyrah;
[Winwood and Wilson]; Woods 1891.
CRETACEOUS, Blake; Woods 1891.
PLIOCENE, Allen 1901u.
TERTIARY, Blake; Melmore.
PLEISTOCENE, Leney; Pyrah.
Cephalopoda : see Mollusca.
Coelenterata (undifferentiated):
CAMBRIAN, Jackson ; Salter.
SILURIAN, Curtis 1956; Jackson; McHenry
and Watts; Salter; Stubblefield 1938;
Woods 1891.
DEVONIAN, Allen 19016; [Cutbill].
CARBONIFEROUS, Jackson; McHenry and
Watts; Melmore; Pyrah.
PERMIAN, Jackson; Woods 1891.
JURASSIC, Jackson; Sizer.
CRETACEOUS, Melmore; Pyrah.
OLIGOCENE, Allen 1900.
TERTIARY, Pyrah.
PLEISTOCENE, Melmore; Pyrah.
Anthozoa :
ORDOVICIAN, Woods 1891.
SILURIAN, Blake; Cantrill et al.-. Woods
1891.
DEVONIAN, Blake; [Cutbill]; Jukes-Browne
and Else.
CARBONIFEROUS, Anderson; Bassett; Blake;
Mitchell and White; Neaverson; North;
Platnauer 1891 ; Sime; Woods 1891.
PERMIAN, Blake.
JURASSIC, Blake; [Gregory]; Platnauer 1891 ;
Woods 1891.
CRETACEOUS, Blake; [Gregory]; Platnauer
1891.
EOCENE, Blake; [Gregory]; Woods 1891.
MIOCENE, Blake.
TERTIARY, Blake.
PLEISTOCENE, Bell.
764
PALAEONTOLOGY, VOLUME 18
Coelenterata (undifferentiated) (cont.) :
Conulata :
TREMADOC, Salter.
ORDOVICIAN, Bassett; Currie and George.
SILURIAN, Blake.
DEVONIAN, Blake.
CARBONIFEROUS, Bassett ; Mitchell and White.
Hydrozoa :
CAMBRIAN, Salter.
DEVONIAN, Salter.
CARBONIFEROUS, Anderson.
PLIOCENE, Allen 1901a.
Chitinozoa:
ORDOVICIAN, Wyatt.
Conodonts: see Miscellanea.
Conulata: see Coelenterata.
Crinoidea: see Echinodermata.
Crustacea : see Arthropoda.
Cystoidea : see Echinodermata.
Decapoda : see Mollusca.
Derived fossils: see Miscellanea.
Echinodermata (undifferentiated):
CAMBRIAN, Bolton 1892; Salter.
ORDOVICIAN, Jackson.
SILURIAN, Jackson; Melmore; Salter.
DEVONIAN, Allen 1901Z); [Cutbill]: Jukes-
Browne and Else.
CARBONIFEROUS, Anderson; Jackson; Mel-
more; Platnauer 1891.
JURASSIC, Anderson; Blake; Bolton 1892;
Jackson; Melmore; Platnauer 1891 ; Sizer.
CRETACEOUS, Blake; Crane 1892; Jackson.
EOCENE, Allen 1900.
PLIOCENE, Allen 1901a; Platnauer 1891.
PLEISTOCENE, Leney : Melmore ; Platnauer 1891.
Asteroidea:
ORDOVICIAN, Allen 19026; Woods 1891.
SILURIAN, Allen 19026; Woods 1891.
DEVONIAN, Allen 19026.
JURASSIC, Woods 1891.
Blastoidea:
SILURIAN, Bather.
DEVONIAN, Bather; [Cutbill]; Rowe.
CARBONIFEROUS, Bather; Rowe.
Crinoidea:
ORDOVICIAN, Allen 19026; Bassett; Currie
and George; Woods 1891.
SILURIAN, Allen 19026; Neaverson; Thomp-
son and George; Woods 1891, 1893.
DEVONIAN, Allen 19026; Blake; [Cutbill].
CARBONIFEROUS, Allen 19026; Currie and
George; McHenry and Watts; Sizer;
Woods 1891.
JURASSIC, Rowe; Woods 1891.
CRETACEOUS, Woods 1891.
Cystoidea :
ORDOVICIAN, Allen 19026; Currie and
George.
SILURIAN, Allen 19026; Salter.
Echinoidea :
ORDOVICIAN, Currie and George; Strachan
et al.
SILURIAN, Stubblefield 1938.
DEVONIAN, [Cutbill].
CARBONIFEROUS, McHenry and Watts;
Woods 1891.
RHAETIAN, Sizer.
JURASSIC, [Gregory]; Tutcher; Woods
1891.
CRETACEOUS, Bassett; Blake; [Gregory];
Woods 1891.
MIOCENE, Blake.
TERTIARY, Blake.
PLEISTOCENE, Bell.
Edrioasteroidea :
ORDOVICIAN, Allen 19026.
CARBONIFEROUS, Allen 19026; Rowe.
Ophiuroidea :
ORDOVICIAN, Woods 1891.
DEVONIAN, Allen 19026.
CARBONIFEROUS, Allen 19026.
JURASSIC, Blake.
CRETACEOUS, Blake.
Stelleroidea :
CARBONIFEROUS, McHenry and Watts.
Echinoidea : see Echinodermata.
Edrioasteroidea : see Echinodermata.
Foraminifera : see Protozoa.
Gastropoda : see Mollusca.
Graptolithina:
TREMADOC, Strachan.
ORDOVICIAN, Anderson; Blake; Strachan;
Strahan et al.; Woods 1891.
SILURIAN, Anderson; Blake; Bassett;
McHenry and Watts; Strachan; Wyatt.
DEVONIAN, [Cutbill].
Hydrozoa: see Coelenterata.
Hyolitha : see Miscellanea.
Insecta: see Arthropoda.
Isopoda: see Arthropoda.
Lamellibranchia : see Bivalvia.
Merostomata: see Arthropoda.
Miscellanea:
Conodonts:
DEVONIAN, [Cutbill].
Derived fossils :
in CARBONIFEROUS, Wyatt.
in TRiASSic, Bassett.
Hyolitha:
ORDOVICIAN, Salter.
BASSETT: TYPES OF FOSSILS
765
Problematica :
PRECAMBRIAN, SizCf.
SILURIAN, Blake.
TRiASSic, Blake.
JURASSIC, Pyrah.
EOCENE, Blake.
Tentaculitida :
SILURIAN, Blake.
Trace fossils:
SILURIAN, Blake; McHenry and Watts.
CARBONIFEROUS, Bassett; McHenry and
Watts.
TRIASSIC, Neaverson; Sizer.
‘Unknown’: see Problematica.
Mollusca (undifferentiated):
CAMBRIAN, Bolton 1892.
SILURIAN, Bolton 1892.
DEVONIAN, [Gregory].
CARBONIFEROUS, Bolton 1892; [Gregory];
Lebour.
TRIASSIC, [Gregory].
JURASSIC, [Gregory]; Torrens, in press.
CRETACEOUS, [Gregory].
TERTIARY, Newton 1902; Bell.
Ammonoidea :
DEVONIAN, [Cutbill].
CARBONIFEROUS, Blake ; McHenry and Watts.
JURASSIC, Blake; [Brighton]; Buckman
[1929]; Cox and Arkell; Crick 1922;
Curtis [1970]; Donovan; Lang;
Neaverson; Sizer; Stubblefield 1939;
Thompson; Torrens, in press.
CRETACEOUS, Blake; Wright and Wright.
Belemnoidea :
JURASSIC, Blake; Cox and Arkell; Crick
1922; Lang; Sizer.
CRETACEOUS, Blake; Wright and Wright.
TERTIARY, Blake.
Bivalvia :
CAMBRIAN, Salter.
ORDOVICIAN, Blake; Currie and George;
Jackson; Stubblefield 1938; Woods 1891.
SILURIAN, Blake; Currie and George;
McHenry and Watts; Salter; Stubble-
field 1938; Wilson; Woods 1891.
OLD RED SANDSTONE, Blake; McHenry and
Watts.
DEVONIAN, Allen 19016; Blake; [Cutbill];
Jukes-Browne and Else; Woods 1891,
1893.
CARBONIFEROUS, Anderson; Bassett; Blake;
Bolton 1894; Doughty; Jackson;
McHenry and Watts; Melmore; North;
Platnauer 1891; Sizer; Wilson; Woods
1891.
PERMIAN, Platnauer 1891.
TRIASSIC, Blake.
RHAETIAN, Allen 1904; Sizer; Tutcher.
JURASSIC, Allen 1904, 1906; Anderson;
Blake; Cox and Arkell; Jackson; Mel-
more; Platnauer 1891; Sizer; Torrens,
in press; Tutcher; [Winwood and Wilson];
Woods 1891.
CRETACEOUS, Allen 1915; Blake; Hallam;
Jackson; Melmore; Platnauer 1891; Wil-
son; [Winwood and Wilson]; Woods 1891 .
EOCENE, Allen 1900; Jackson; Newton 1891.
OLiGOCENE, Allen 1900; Newton 1891.
MIOCENE, Blake.
PLIOCENE, Allen 1910a; Jackson; Platnauer
1891; Woods 1891.
TERTIARY, Blake; Jackson; Melmore.
PLEISTOCENE, Bell; Leney; Platnauer 1891;
Sizer.
Cephalopoda (undifferentiated):
CAMBRIAN, Anderson; Jackson.
TREMADOC, Salter.
ORDOVICIAN, Anderson; Blake; Crick 1898;
Woods 1891.
SILURIAN, Anderson; Bassett; Blake; Crick
1898; Curtis 1956; Jackson; Salter;
Stubblefield 1938; Woods 1891.
DEVONIAN, Allen 19016; Crick 1898; [Cut-
bill] ; Jukes-Browne and Else ; Woods 1891.
CARBONIFEROUS, Anderson; Bassett; Crick
1898; Jackson; McHenry and Watts;
Melmore; Neaverson; North; Platnauer
1891 ; Sizer; Tutcher; Woods 1891.
TRIASSIC, Crick 1898.
JURASSIC, Anderson; Bassett; Crick 1898;
Howarth ; Jackson ; Melmore ; Neaverson ;
Platnauer 1898; Tutcher; Wilson; Woods
1891, 1893.
CRETACEOUS, Blake; Crane 1892; Crick
1898; Howarth; Jackson; Melmore; Plat-
nauer 1891; Tutcher; Wilson; Woods
1891.
EOCENE, Crick 1898.
Decapoda :
CRETACEOUS, Woods 1891.
Gastropoda :
CAMBRIAN, Anderson; Salter; Woods 1891.
TREMADOC, Curtis 1956; Salter.
ORDOVICIAN, Anderson; Blake; Currie and
George; Jackson; Stubblefield 1938;
Woods 1891.
SILURIAN, Anderson; Bassett; Blake; Curtis
1956; McHenry and Watts; Jackson;
North; Salter; Stubblefield 1938; Wilson;
Woods 1891.
766
PALAEONTOLOGY, VOLUME 18
Gastropoda (cont.)\
OLD RED SANDSTONE, Blake.
DEVONIAN, Allen 1901ft; Blake; [Cutbill];
Jukes-Browne and Else; Woods 1891,
1893.
CARBONIFEROUS, Anderson; Blake; Bolton
1894; Currie and George; Jackson;
McHenry and Watts ; North ; Sizer ; Woods
1891.
TRiASSic, [Winwood and Wilson].
RHAETiAN, Allen 1903.
JURASSIC, Allen 1903, 1904; Anderson;
Blake; Cox and Arkell: Jackson; Plat-
nauer 1891; Sizer; Tutcher; [Winwood
and Wilson]; Woods 1891, 1893.
CRETACEOUS, Allen 1916; Blake; Crane
1892; Platnauer 1891; Wilson; Woods
1891.
EOCENE, Allen 1900.
OLiGOCENE, Allen 1900.
MIOCENE, Blake; Jackson.
PLIOCENE, Allen 1901a; Platnauer 1891.
TERTIARY, Blake.
PLEISTOCENE, Allen 1901a; Bell; Blake;
Leney; Platnauer 1891 ; Sizer.
Lamellibranchia : see Bivalvia.
Nautiloidea:
ORDOVICIAN, Blake.
SILURIAN, Blake.
OLD RED SANDSTONE, Blake.
CARBONIFEROUS, Blake.
JURASSIC, Blake; Cox and Arkell; Howarth;
Neaverson; Sizer.
CRETACEOUS, Blake; Wright and Wright.
TERTIARY, Blake.
Scaphopoda :
DEVONIAN, [Cutbill].
CARBONIFEROUS, Anderson.
PERMIAN, Riley.
RHAETIAN, Allen 1903.
JURASSIC, Allen 1903, 1904; Cox and Arkell;
[Winwood and Wilson].
CRETACEOUS, Woods 1891.
PLEISTOCENE, Melmore.
Nautiloidea : see Mollusca.
Ophiuroidea : see Echinodermata.
Polyzoa: see Bryozoa.
Porifera :
CAMBRIAN, Salter; Woods 1891.
ORDOVICIAN, Salter; Woods 1891 ; Wyatt.
SILURIAN, Salter; Anderson; Woods 1891;
Wyatt.
DEVONIAN, [Cutbill].
JURASSIC, Blake; Platnauer 1891 ; Pyrah.
CRETACEOUS, Melmore; Platnauer 1891;
Pyrah; Woods 1891.
PLEISTOCENE, Leney; Melmore.
Problematica : see Miscellanea.
Protozoa (undifferentiated) ;
CARBONIFEROUS, Bolton 1892; Jackson.
CRETACEOUS, Blake.
EOCENE, Allen 1900.
Foraminifera :
SILURIAN, Cantrill et al. ; Jones.
DEVONIAN, Jones.
PERMIAN, Jones.
JURASSIC, Blake; Jones; [Winwood and Wil-
son].
CRETACEOUS, Jones.
EOCENE, Jones.
TERTIARY, Blake.
Radiolaria :
ORDOVICIAN, Anderson.
Radiolaria: see Protozoa.
Scaphopoda : see Mollusca.
Stelleroidea : see Arthropoda.
Stromatoporoidea :
SILURIAN, Salter.
DEVONIAN, [Gregory].
Tentaculitida: see Miscellanea.
Trace fossils: see Miscellanea.
Trilobita: see Arthropoda.
‘Unknown’: see Miscellanea.
Vermes: see Annelida.
VERTEBRATA
Vertebrata (undifferentiated):
OLD RED SANDSTONE, Anderson.
EOCENE, Allen 1900.
OLIGOCENE, Allen 1900.
PLEISTOCENE, Bell.
Amphibia:
CARBONIFEROUS, Blake; Lydekker 1889;
McHenry and Watts; Paton; Waterston
1954; Woods 1891 ; Woodward and Sher-
born.
PERMIAN, Lydekker 1889; Paton; Waterston
1954; Woodward and Sherborn.
TRIASSIC, Blake; Lydekker 1889; Paton;
Woodward and Sherborn.
RHAETIAN, Sizer; Woodward and Sherborn.
CRETACEOUS, Lydekker 1889.
MIOCENE, Lydekker 1889.
PLEISTOCENE, Lydekker 1889; Woodward
and Sherborn.
HOLOCENE, Carreck.
BASSETT: TYPES OF FOSSILS
767
Aves:
CRETACEOUS, Melmore ; Seeley ; Woods 1891;
Woodward and Sherborn.
EOCENE, Lydekker 1891 ; Woods 1891 ; Wood-
ward and Sherborn.
OLiGOCENE, Woods 1891; Woodward and
Sherborn.
PLIOCENE, Lydekker 1891.
TERTIARY, Blake.
PLEISTOCENE, Allen 1901a; Bell; Leney;
Lydekker 1891; Melmore; Woodward
and Sherborn.
HOLOCENE, Carreck.
Fishes:
ORDOVICIAN, Henrichsen 1971.
SILURIAN, Anderson; Bassett; Bolton 1892;
Henrichsen 1971 ; Salter; Waterston 1954;
Woodward and Sherborn.
OLD RED SANDSTONE, Blake; Bolton 1892;
McHenry and Watts; Waterston 1954,
1968a, 19686, 1968c; Woods 1891 ; Wood-
ward 1891 ; Woodward and Sherborn.
DEVONIAN, Blake; [Cutbill]; Davies 1871a;
Egerton 1869; Enniskillen; Henrichsen
1970, 1971, 1972; Waterston 1954; Wood-
ward and Sherborn.
CARBONIFEROUS, Anderson; Bassett; Blake;
Bolton 1892, 1894; Davies 1871a; [Delair]
1966c; Egerton 1836, 1869; Enniskillen;
Henrichsen 1970, 1972; Lebour; McHenry
and Watts; North; Platnauer 1891 ; Sizer;
Waterston 1954; Wilson; Woods 1891,
1893; Woodward 1891; Woodward and
Sherborn.
PERMIAN, Davies 1871a; Egerton 1836, 1869;
Enniskillen; Henrichsen 1970, 1972;
Waterston 1954; Woodward 1891 ; Wood-
ward and Sherborn.
TRiASSic, Bassett; Blake; Egerton 1869;
Henrichsen 1970, 1972; Sizer; Waterston
1954; Woodward and Sherborn.
RHAETiAN, Blake; Bolton 1894; Egerton
1869; Wilson; Woodward and Sherborn.
JURASSIC, Blake; Davies 1871a; Egerton
1836, 1869; Enniskillen; Hallam; Hen-
richsen 1970, 1972; Lang; Platnauer 1891 ;
Sizer; Thompson; Torrens, in press; Wil-
son; Winwood and Wilson; Woods 1891 ;
Woodward 1889, 1891; Woodward and
Sherborn.
CRETACEOUS, Anon 1896; Blake; Bolton
1894; Crane 1892, 1893; Davies 1871a;
Egerton 1836, 1869; Enniskillen; Hen-
richsen 1970, 1971 ; Platnauer 1891 ; Sizer;
Waterston 1954; Woods 1891 ; Woodward
1889, 1891, 1895, 1901; Woodward and
Sherborn.
EOCENE, Bolton 1894; Davies 1871a; Egerton
1869; Enniskillen; Henrichsen 1970;
Woodward 1889, 1891, 1901; Woodward
and Sherborn.
OLIGOCENE, Egerton 1 869 ; Enniskillen ; Hen-
richsen 1970; Woodward 1901; Wood-
ward and Sherborn.
MIOCENE, Davies 1871a; Egerton 1869;
Enniskillen; Woods 1891.
PLIOCENE, Allen 1901a; Bell; Gregory; Hen-
richsen 1970; Woodward 1891.
TERTIARY, Egerton 1836, 1869; Enniskillen;
[Gregory]; Henrichsen 1970; Leney; Plat-
nauer 1891 ; Woodward and Sherborn.
Mammalia ;
TRIASSIC, Winwood and Wilson.
RHAETIAN, Woodward and Sherborn.
JURASSIC, Melmore; Woodward and Sher-
born.
CRETACEOUS, Woodward and Sherborn.
EOCENE, Blake; Lydekker 1885, 1887; Mel-
more; Platnauer 1891; Woods 1891;
Woodward and Sherborn.
OLIGOCENE, Blake; Lydekker 1889; Woods
1891 ; Woodward and Sherborn.
MIOCENE, Blake; Lydekker 1885a, 18856,
1886a, 1887; Woods 1891.
PLIOCENE, Allen 1901a; Lydekker 1885a,
18856, 18866, 1887.
TERTIARY, Blake; Owen.
PLEISTOCENE, Allen 1901a; Blake; Bolton
1892; Carreck; [Gregory]; Leney;
Lydekker 1885a, 18856, 1886a, 18866,
1887; Melmore; Owen; Platnauer 1891;
Sanford; Sizer; Woods 1891 ; Woodward
and Sherborn.
HOLOCENE, Carreck.
Reptilia :
CARBONIFEROUS, Blake; Baton.
PERMIAN, Blake; Baton.
TRIASSIC, Blake; Bassett; Lydekker 1889;
North; Baton; Seeley; Sizer; Wilson;
Woodward and Sherborn.
RHAETIAN, Paton ; Sizer; Woodward and
Sherborn.
JURASSIC, Appleby; Blake; [Gregory]; Lang;
Lydekker 1889; Melmore; Paton; Plat-
nauer 1891; Seeley; Sizer; Torrens, in
press; Woods 1891 ; Woodward and Sher-
born.
CRETACEOUS, Crane 1892; Lydekker 1889;
Paton; Platnauer 1891; Seeley; Woods
1891 ; Woodward and Sherborn.
768
PALAEONTOLOGY, VOLUME 18
Reptilla (cont.):
EOCENE, Lydekker 1889; Woodward and
Sherborn.
OLiGOCENE, Woodward and Sherborn.
PLIOCENE, [Gregory]; Lydekker 1889.
TERTIARY, Blake.
PLEISTOCENE, Leney; Lydekker 1889; Wood-
ward and Sherborn.
Vertebrate footprints:
PERMIAN, Delair 1966a, 19666; Paton.
TRiASSic, Bassett; Neaverson; Paton.
JURASSIC, Paton.
PLANTAE
Plantae (undifferentiated) :
CAMBRIAN, Salter.
ORDOVICIAN, Woods 1891.
SILURIAN, Bassett; Blake; Jackson; Salter.
OLD RED SANDSTONE, Andcrson; Jackson;
McHenry and Watts.
DEVONIAN, Bassett; Blake; Calder; Hopping.
CARBONIFEROUS, Andcrson; Anon 1957;
Bassett; Blake; Bolton 1892, 1894; Calder;
Hopping; Jackson; Lebour; McHenry
and Watts; Neaverson; Platnauer 1891;
Riley; Seward 1894; Sizer; Winwood and
Wilson; Woods 1891, 1893.
PERMIAN, Calder; Hopping; Jackson; Sizer.
TRIASSIC, Blake; Jackson; Lebour; Seward
1894; Sizer.
RHAETIAN, Sizcr.
JURASSIC, Anderson; Blake; Bolton 1892;
Calder; Howse; Jackson; Lebour; Mel-
more ; Seward 1 894, 1 900 ; Sizer.
CRETACEOUS, Blake; Calder; Jackson; Plat-
nauer 1891; Woods 1891.
EOCENE, Allen 1900; McHenry and Watts;
Woods 1891.
OLIGOCENE, Allen 1900.
PLIOCENE, Allen 1901a.
TERTIARY, Calder; [Gregory].
PLEISTOCENE, Allen 1901a; Sizer.
Algae (including stromatolites) :
ORDOVICIAN, Salter.
SILURIAN, Blake; Curtis 1956; Salter; Wyatt.
DEVONIAN, [Cutbill].
EOCENE, Morellet and Morellet.
STRATIGRAPHICAL INDEX
PRECAMBRiAN : See Problematica.
CAMBRIAN: see Annelida, Arthropoda (undiffer-
entiated), Bivalvia, Brachiopoda, Bryozoa,
Cephalopoda, Coelenterata (undifferentiated),
Crustacea, Echinodermata (undifferentiated).
Gastropoda, Hydrozoa, Mollusca (undiffer-
entiated), Plantae (undifferentiated), Porifera.
TREMADOC: see Brachiopoda, Crustacea, Gastro-
poda, Graptolithina, Trilobita.
ORDOVICIAN: see Actinozoa, Algae, Annelida,
Asteroidea, Bivalvia, Brachiopoda, Bryozoa,
Cephalopoda, Chitinozoa, Conulata, Crinoidea,
Crustacea, Cystoidea, Echinodermata (undiffer-
entiated), Echinoidea, Edrioasteroidea, Fishes,
Gastropoda, Graptolithina, Lamellibranchia,
Nautiloidea, Plantae (undifferentiated), Polyzoa,
Porifera, Radiolaria, Trilobita.
SILURIAN: see Algae, Annelida, Anthozoa, Arthro-
poda (undifferentiated), Asteroidea, Bivalvia,
Blastoidea, Brachiopoda, Bryozoa, Cephalopoda,
Coelenterata (undifferentiated), Conulata,
Crinoidea, Crustacea, Cystoidea, Echinodermata
(undifferentiated), Echinoidea, Fishes, Foramini-
fera. Gastropoda, Graptolithina, Hydrozoa,
Lamellibranchia, Merostomata, Mollusca (un-
differentiated), Nautiloidea, Plantae (undiffer-
entiated), Polyzoa, Porifera, Problematica, Stro-
matolites, Stromatoporoidea, Tentaculitida, Tri-
lobita.
OLD RED sandstone: See Arthropoda (undiffer-
entiated), Bivalvia, Brachiopoda, Fishes, Nau-
tiloidea, Plantae (undifferentiated), Vertebrata
(undifferentiated).
DEVONIAN : see Actinozoa, Algae, Annelida, Antho-
zoa, Arachnida, Asteroidea, Bivalvia, Brachio-
poda, Blastoidea, Bryozoa, Cephalopoda,
Coelenterata (undifferentiated), Conodonts,
Conulata, Crinoidea, Crustacea, Echinodermata
(undifferentiated), Echinoidea, Fishes, Foramini-
fera. Gastropoda, Graptolithina, Lamellibran-
chia, Mollusca (undifferentiated), Ophiuroidea,
Plantae (undifferentiated), Porifera, Protozoa,
Scaphopoda, Trilobita.
CARBONIFEROUS : See Actinozoa, Ammonoidea, Am-
phibia, Annelida, Anthozoa, Arachnida, Arthro-
poda (undifferentiated), Bivalvia, Blastoidea,
Brachiopoda, Bryozoa, Cephalopoda, Coelen-
terata (undifferentiated), Conulata, Crinoidea,
Crustacea, Derived fossils, Echinodermata
(undifferentiated), Edrioasteroidea, Fishes,
Gastropoda, Hydrozoa, Insecta, Merostomata,
Mollusca (undifferentiated), Nautiloidea, Ophiu-
BASSETT: TYPES OF FOSSILS
769
roidea, Plantae (undifferentiated), Polyzoa,
Protozoa (undifferentiated), Reptilia, Scapho-
poda, Stelferoidea, Trace fossils, Trilobita,
PERMIAN: see Amphibia, Anthozoa, Bivalvia,
Brachiopoda, Crustacea, Fishes, Foraminifera,
Plantae (undifferentiated), Reptilia, Scaphopoda,
Vertebrate footprints.
TRiASSic: see Amphibia, Bivalvia, Brachiopoda,
Cephalopoda, Crustacea, Derived fossils. Fishes,
Gastropoda, Mammalia, Mollusca (undiffer-
entiated), Plantae (undifferentiated), Proble-
matica, Reptilia, Trace fossils. Vertebrate foot-
prints.
rhaetian: see Amphibia, Annelida, Bivalvia,
Brachiopoda, Crustacea, Echinoidea, Fishes,
Gastropoda, Insecta, Lamellibranchia, Mam-
malia, Plantae (undifferentiated), Reptilia,
Scaphopoda.
JURASSIC; see Actinozoa, Ammonoidea, Annelida,
Anthozoa, Arthropoda (undifferentiated), Aste-
roidea, Belemnoidea, Bivalvia, Brachiopoda,
Bryozoa, Cephalopoda, Coelenterata (undiffer-
entiated), Crinoidea, Crustacea, Echinodermata
(undifferentiated), Echinoidea, Fishes, Foramini-
fera, Gastropoda, Insecta, Lamellibranchia,
Mammalia, Mollusca (undifferentiated), Nauti-
loidea, Ophiuroidea, Plantae (undifferentiated),
Polyzoa, Porifera, Problematica, Reptilia,
Scaphopoda, Vertebrate footprints.
cretaceous: see Ammonoidea, Amphibia, Antho-
zoa, Annelida, Aves, Belemnoides, Bivalvia,
Brachiopoda, Bryozoa, Cephalopoda, Coelen-
terata (undifferentiated), Crinoidea, Crustacea,
Decapoda, Echinodermata (undifferentiated),
Echinoidea, Fishes, Foraminifera, Gastropoda,
Isopoda, Lamellibranchia, Mollusca (undiffer-
entiated), Nautiloidea, Ophiuroidea, Plantae (un-
differentiated), Polyzoa, Porifera, Protozoa (un-
differentiated), Reptilia, Scaphopoda.
eocene: see Algae, Anthozoa, Aves, Bivalvia,
Cephalopoda, Crustacea, Echinodermata (un-
differentiated), Fishes, Foraminifera, Gastro-
poda, Lamellibranchia, Mammalia, Plantae
(undifferentiated), Problematica, Protozoa (un-
differentiated), Reptilia, Vertebrata (undiffer-
entiated).
oligocene: see Aves, Bivalvia, Coelenterata (un-
differentiated), Fishes, Gastropoda, Lamelli-
branchia, Mammalia, Plantae (undifferentiated),
Vertebrata (undifferentiated).
MIOCENE: see Amphibia, Anthozoa, Bivalvia, Echi-
noidea, Fishes, Gastropoda, Mammalia.
PLIOCENE; see Aves, Bivalvia, Bryozoa, Echino-
dermata (undifferentiated). Fishes, Gastropoda,
Hydrozoa, Lamellibranchia, Mammalia, Plantae
(undifferentiated), Reptilia.
TERTIARY (UNDIFFERENTIATED): see Annelida,
Anthozoa, Aves, Belemnoidea, Bivalvia, Brachio-
poda, Bryozoa, Coelenterata, Crustacea, Echi-
noidea, Foraminifera, Fishes, Gastropoda,
Insecta, Lamellibranchia, Mammalia, Mollusca
( undifferentiated), Nautiloidea, Plantae ( undiffer-
entiated), Polyzoa, Reptilia, Vertebrata (undiffer-
entated).
PLEISTOCENE : See Amphibia, Annelida, Aves,
Bivalvia, Brachiopoda, Bryozoa, Coelenterata
(undifferentiated), Crustacea, Echinodermata
(undifferentiated). Fishes, Gastropoda, Lamelli-
branchia, Mammalia, Plantae (undifferentiated),
Polyzoa, Porifera, Reptilia, Scaphoda, Verte-
brata (undifferentiated).
QUATERNARY : see PLEISTOCENE.
HOLOCENE: see Amphibia, Aves, Mammalia.
MUSEUMS INDEX
ABERYSTWYTH; See University College of Wales,
Aberystwyth.
AYLESBURY : See Buckinghamshire County Museum.
BATH : see Victoria Art Gallery, Bath.
BEDFORD museum: Woodward and Sherborn.
BELFAST: see Ulster Museum.
BIRMINGHAM: See University of Birmingham Geo-
logical Department.
BOTANY DEPARTMENT, UNIVERSITY OF GLASGOW ; See
Hunterian Museum.
BRADFORD: See City Art Gallery and Museum,
Bradford.
BRIGHTON: see Natural History Museum, Brighton.
BRISTOL: see City Museum, Bristol, and University
of Bristol.
BRITISH MUSEUM (NATURAL HISTORY) : Bather ; Blake ;
Buckman [1929]; Carreck; Cox and Arkell;
Crick 1898, 1922; Curtis 1956; Davies 1871a,
18716; Delair 1966a; Donovan; Egerton 1836,
1869; Enniskillen; Howarth; Jones; Lang;
Lydekker 1885-1887, 1891 ; Morellet and Morel-
let; Newton 1891, 1902; Strachan; Torrens in
press; Tutcher; Woodward 1889-1901; Wood-
ward and Sherborn; Wright and Wright.
BUCKINGHAMSHIRE COUNTY MUSEUM; Woodward
and Sherborn.
BURGH MUSEUM, DUMFRIES: Delair 1966a.
CAMBRIDGE; see Sedgwick Museum, antf University
of Cambridge, Botanical Museum.
CARDIFF: see National Museum of Wales.
770
PALAEONTOLOGY, VOLUME 18
CASTLE MUSEUM, NORWICH: Lcncy; Woodward and
Sherborn; Wright and Wright.
CENTRAL LIBRARY, MUSEUM AND ART GALLERY, HULL :
Woodward and Sherborn.
CENTRAL MUSEUM AND ART GALLERY, NORTHAMPTON :
Cox and Arkell; Thompson; Thompson and
George; Woodward and Sherborn.
CITY ART GALLERY AND MUSEUM, BRADFORD : Wood-
ward and Sherborn.
CITY MUSEUM, BRISTOL: Buckman [1929]; Curtis
1970; Wilson; Woodward and Sherborn.
CITY MUSEUM, LEEDS: Woodward and Sherborn.
CITY MUSEUM AND ART GALLERY, PETERBOROUGH:
Appleby.
CITY MUSEUM AND ART GALLERY, WORCESTER : Wood-
ward and Sherborn.
COUNTY MUSEUM, WARWICK : Woodward and Sher-
born.
DORCHESTER : see Dorset County Museum.
DORSET COUNTY MUSEUM : Carreck ; [Samuel] ; Wood-
ward and Sherborn.
DUBLIN : see Geological Survey of Ireland, National
Museum of Ireland, unJ Trinity College, Dublin.
DUBLIN UNIVERSITY MUSEUM: See Trinity College,
Dublin.
DUMFRIES: see Burgh Museum, Dumfries.
EDINBURGH: See Institute of Geological Sciences,
Edinburgh, and Royal Scottish Museum.
ELGIN museum: Watcrston 1968h, 1968c; Wood-
ward and Sherborn.
EXETER : see Royal Albert Memorial Museum,
Exeter.
FARNHAM : See Pitt Rivers Museum, Farnham.
FORRES museum: Waterston 1968a; Woodward and
Sherborn, see also Royal Scottish Museum,
Edinburgh.
GEOLOGICAL SURVEY OF IRELAND: McHcury and
Watts; Woodward and Sherborn.
GEOLOGICAL SURVEY MUSEUM: see Institute of
Geological Sciences, Leeds, London, and Edin-
burgh.
GEOLOGY MUSEUM, UNIVERSITY OF BRISTOL: see
University of Bristol.
GLASGOW: see Hunterian Museum, University of
Glasgow, and Royal College of Science and
Technology, Glasgow.
HALIFAX : see Museums and Art Galleries, Halifax.
HANCOCK MUSEUM, NEWCASTLE UPON TYNE: HOWSC;
Lebour; Woodward and Sherborn.
hull: see Central Library, Museum and Art
Gallery, Hull, and University of Hull Geology
Department.
HUNTERIAN MUSEUM, UNIVERSITY OF GLASGOW:
Anon 1957; Calder; Currie and George;
[Gregory]; Hopping; Woodward and Sherborn.
INSTITUTE OF GEOLOGICAL SCIENCES, EDINBURGH:
Anderson; Strachan; Woodward and Sherborn.
INSTITUTE OF GEOLOGICAL SCIENCES, LEEDS: Allen
1902a, 19026; Blake; Cantrill et al. ; Mitchell and
White; Strachan.
INSTITUTE OF GEOLOGICAL SCIENCES, LONDON : Allen
1900, 1901a, 19016, 1902a, 19026, 1903, 1904.
1905, 1906, 1915, 1916; Blake; Buckman [1929];
Cantrill et al. ', Carreck; Cox and Arkell; Curtis
1956; Donovan; Howarth; Strachan; Strachan
et al.', Stubblefield 1936, 1938; Woodward and
Sherborn; Wright and Wright.
IPSWICH museum: Bell; Woodward and Sherborn.
KILMARNOCK: See Public Library, Museum and
Art Gallery, Kilmarnock.
LEEDS: see City Museum, Leeds, and Institute of
Geological Sciences, Leeds.
LEICESTERSHIRE MUSEUM, ART GALLERIES AND
RECORDS service: Appleby; Sizer; Woodward
and Sherborn.
LUDLOW museum: Woodward and Sherborn.
MALTON MUSEUM : Woodward and Sherborn.
MANCHESTER MUSEUM: Bolton 1892, 1894; Buckman
[1929]; Cox and Arkell; Seward 1900; Wood-
ward and Sherborn.
MUSEUM AND ART GALLERY, PAISLEY: Delair 19666.
MUSEUM OF NATURAL HISTORY, SCARBOROUGH:
Howarth; Woodward and Sherborn.
MUSEUM OF PRACTICAL GEOLOGY, LONDON : See
Institute of Geological Sciences, London.
MUSEUM OF SCIENCE AND ART, DUBLIN: see National
Museum of Ireland.
MUSEUMS AND ART GALLERIES, HALIFAX: Woodward
and Sherborn.
NATIONAL MUSEUM OF IRELAND, DUBLIN : McHenry
and Watts; Woodward and Sherborn.
NATIONAL MUSEUM OF WALES, CARDIFF: BaSSett ;
North; Strachan.
NATURAL HISTORY AND ANTIQUARIAN MUSEUM,
PENZANCE: Woodward and Sherborn.
NATURAL HISTORY MUSEUM, BRIGHTON : Anon 1 896 ;
Crane 1892, 1893; Woodward and Sherborn.
NEWCASTLE UPON TYNE: See Hancock Museum,
Newcastle upon Tyne.
NORTHAMPTON: See Central Museum and Art
Gallery, Northampton.
NORWICH: see Castle Museum, Norwich.
OWENS COLLEGE, MANCHESTER: See Manchester
Museum.
oxford: see University Museum, Oxford.
paisley: see Museum and Art Gallery, Paisley.
PENZANCE: see Natural History and Antiquarian
Museum, Penzance.
PETERBOROUGH: See City Museum and Art Gallery,
Peterborough.
BASSETT: TYPES OE EOSSILS
771
PITT RIVERS MUSEUM, FARNHAM : Carreck.
PUBLIC LIBRARY, MUSEUM AND ART GALLERY, KIL-
MARNOCK: Delair 1966c.
PUBLIC MUSEUM AND ART GALLERY, SUNDERLAND:
Woodward and Sherborn.
ROYAL ALBERT MEMORIAL MUSEUM, EXETER : Rowe.
ROYAL COLLEGE OF SCIENCE AND TECHNOLOGY,
GLASGOW: Howarth.
ROYAL COLLEGE OF SURGEONS, HUNTERIAN MUSEUM :
[Morris and Owen]; Owen; Woodward and
Sherborn.
ROYAL SCOTTISH MUSEUM, EDINBURGH: Anderson ;
Delair 1966a; Hennchsen 1970, 1971, 1972;
Paton; Sime; Waterston 1954, 1968a; Wood-
ward and Sherborn.
ST. ANDREWS MUSEUM : Woodward and Sherborn.
SALFORD : see Science Museum, Salford.
SALISBURY AND SOUTH WILTSHIRE MUSEUM: Wood-
ward and Sherborn.
SCARBOROUGH: see Museum of Natural History,
Scarborough.
SCIENCE MUSEUM, SALFORD: Woodward and Sher-
born.
SEDGWICK MUSEUM, CAMBRIDGE UNIVERSITY:
[Brighton]; Buckman [1929]; Cox and Arkell;
Curtis 1956; [Cutbill]; Donovan; Howarth;
Salter; Seeley; Strachan; Woods 1891, 1893;
Woodward and Sherborn; Wright and Wright.
SHEFFIELD CITY MUSEUMS : Riley.
SHERBORNE SCHOOL MUSEUM: Torrens, in press.
SHREWSBURY MUSEUM: Woodward and Sher-
born.
SOMERSET COUNTY MUSEUM, TAUNTON CASTLE :
Carreck; Hallam; Sanford; Woodward and
Sherborn.
SUNDERLAND: See Public Museum and Art Gallery,
Sunderland.
TAUNTON : see Somerset County Museum, Taunton
Castle.
TORQUAY NATURAL HISTORY SOCIETY MUSEUM : JukeS-
Browne and Else.
TRINITY COLLEGE, DUBLIN: Strachan; Woodward
and Sherborn.
ULSTER museum: Donovan; Doughty.
UNIVERSITY COLLEGE OF WALES, ABERYSTWYTH :
Wyatt.
UNIVERSITY MUSEUM, OXFORD: Buckman [1929];
Delair 1966a; Edmonds; Torrens 1974, in press;
Woodward and Sherborn.
UNIVERSITY OF BIRMINGHAM GEOLOGY DEPARTMENT :
Strachan.
UNIVERSITY OF BRISTOL, GEOLOGY MUSEUM: Curtis
1956.
UNIVERSITY OF CAMBRIDGE, BOTANICAL MUSEUM:
Seward 1894.
UNIVERSITY OF HULL GEOLOGY DEPARTMENT :
Strachan.
VICTORIA ART GALLERY, BATH: [Winwood and
Wilson]; Woodward and Sherborn.
wales: see National Museum of Wales.
WARWICK: see County Museum, Warwick.
WHITBY museum: Howarth; Woodward and Sher-
born. '
WOODWARDIAN MUSEUM, CAMBRIDGE : See Sedgwick
Museum, Cambridge University.
WORCESTER: see City Museum and Art Gallery,
Worcester.
YORKSHIRE MUSEUM, YORK: Cox and Arkell;
Howarth; Melmore; Platnauer 1891, 1894;
Pyrah; Woodward and Sherborn.
SUPPLEMENTARY REFERENCES
As noted earlier the whereabouts of a great many type, figured, and cited fossils
may remain unknown if there is no published information on them, and there can
be no guarantee that some museums are even aware that they house such material.
Apart from catalogues of types, however, there are numerous other publications
which contain information on old collections, and which may give some guidance
in a search for a particular specimen. Most museums produce Annual Reports which
list major accessions during any one particular year, and many also publish guides
to the collections and galleries, often including notes on particular items of interest.
Biographical and/or obituary notices of known collectors may include details of the
whereabouts of their collections, with many journals of local natural history societies
giving a great deal of information for a particular area. The new Newsletter of the
Geological Curators Group plans to collate and publish these kinds of data and to
provide general guidance on the location of collections of fossils in Britain. It is
clearly not possible to cite the whole range of these publications here, but the following
list is intended to draw attention to the variety of sources of information and to some
772
PALAEONTOLOGY, VOLUME 18
Standard references on museums, collections, and biographies of collectors; it makes
no claim to be either complete or comprehensive in its coverage. Also included are
a few useful references giving guidance to the maintenance and storage of type fossil
collections.
CHALMERS-HUNT, J. M. In prcss. Natural History auctions 1700-1972: a register of sales in the British Isles.
Sotheby & Co., London. [Information on the disposal of collections at auctions.]
COOPER, J. A. Geological collections and collectors of note; 2. Northampton Central Museum. Newsletter
of the Geological Curators Group, No. 2, 40-45. [With an appendix (pp. 46-51) by H. S. Torrens on
collectors represented at Northampton ; both the paper and appendix note important collections which
have been discovered recently, and there is a note that a type catalogue is in preparation jointly by Cooper
and Torrens.]
cox, L. R. 1956. Fossil invertebrate collections from India and Pakistan in the British Museum (Natural
History). J. Palaeont. Soc. India, 1, 94-98. [Valuable notes on collections and collectors, with the publica-
tions in which the specimens are described.]
CURTIS, M. L. K. 1962. [Note on location of type specimens of Silurian Bivalvia and Gastropoda.] Ludlow
Research Group Bulletin, No. 10, p. 4. [City Museum, Bristol, and IGS, London.]
DANCE, s. p. 1967. Report on the Linnaean shell collection. Proc. Linn. Soc. Lond. 178, 1-24, pis. 1-10.
[Mainly conchological, but also with details of collectors who provided Linnaeus with fossils.]
HUXLEY, T. H. and ETHERIDGE, R. 1865. A catalogue of the collections of fossils in the Museum of Practical
Geology, with an explanatory introduction, lxxix + 381 pp., H.M.S.O., London. [Valuable information
in footnotes referring to donors of collections in IGS.]
LAMBRECHT, K., QUENSTEDT, w. and QUENSTEDT, A. 1938. FossUium catalogus I. Animalia. Pars 72: Palaeon-
tologi. Catalogus bio-hibliographicus. xxii + 495 pp., W. Junk, Gravenhage. [A major, though often
neglected, source of biographical details of palaeontologists; in German but with abundant data relevant
to Britain.]
LEBOUR, G. A. 1886. Materials for a palaeontology of Northumberland. Chapter 14, pp. 108-1 13. In Outlines
of the geology of Northumberland and Durham. viii+ 156 pp., 5 pis., Lambert and Co., Newcastle upon
Tyne. [An example of data on regional collections; many of those listed are now in the Hancock Museum.]
LEEDS, E. T. Edited with notes and additions by w. e. swinton, 1956. The Leeds collection of fossil reptiles
from the Oxford Clay of Peterborough. xii4 104 pp., 6 pis., B. H. Blackwell, Oxford. [History of the
collection and disposal of one of the most important collections of British vertebrates.]
MURRAY, D. 1904. Museums: their history and their use: with a bibliography and list of museums in the United
Kingdom. Vol. 1, xvi-( 339 pp.; Vol. 2, 363 pp.; Vol. 3, 341 pp., James MacLehose and Sons, Glasgow.
[Contains a great deal of information on early collectors and collections, and an extensive bibliography
of museum publications, including fossil catalogues.]
MURRAY, J. w. 1971. The W. B. Carpenter Collection. Micropalaeontology, 17, 105-106. [Notes on Car-
penter’s collection of foraminifers in the P. F. Sladen Collection at the Royal Albert Memorial Museum,
Exeter.]
OWEN, D. E. 1964. Care of type specimens. Mus. J. 63, 288-291.
PYRAH, B. J. 1974. Geological collections and collectors of note: 3. Yorkshire Museum. Newsletter of the
Geological Curators Group, No. 2, 52-55. [With an appendix (pp. 56-58) by H. S. Torrens of notes on
some Yorkshire Museum collectors.]
SARJEANT, w. A. s. 1974. A history and bibliography of the study of fossil vertebrate footprints in the British
Isles. Palaeogeography, Palaeoclimatology, Palaeoecology, 16 (Special issue), 165-378. [Contains a great
deal of useful information on collectors and collections, including present repositories.]
SHERBORN, c. D. 1940. Where is the Collection? An account of the various Natural History Collections
which have come under the notice of the compiler Charles Davies Sherborn D.Sc. Oxon. between 1880 and
1939. 148 pp.. University Press, Cambridge. [The standard primary source of information on the location
of natural history collections, with a bias towards geology; the information is being updated under the
editorship of R. Cleevely of the BM(NH) to be incorporated in a 2nd edition of the book, which will
be published by the BM(NH) and the Society for the Bibliography of Natural History.]
lORRENS, H. s. I974u. Geological collections and collectors of note: 1. Lichheld Museums (pre 1850).
Newsletter of the Geological Curators Group, No. 1, 5-10.
BASSETT: TYPES OF FOSSILS
773
TORRENS, H. s. \91Ab. Geological collections and collectors of note: 1. Lichfield Museums (pre 1850) post-
script. Ibid. No. 2, 38-39.
1974c. Locating and identifying collections of palaeontological material. Ibid. No. 1, 12-17. [Includes
a useful list of published sources of biographies of geologists in addition to those listed here.]
WOODWARD, A. s. 1904. The department of geology, pp. 197-340. In The history of the collections contained
in the Natural History departments of the British Museum. Vol. 1, 442 pp., British Museum (Natural
History), London. [Abundant data on important early collectors and collections, with information on
publications in which specimens are described.]
YOCHELSON, E. 1969. Fossils— the how and why of collecting and storing. In cohen, d. m. and cressey, r. f.
(eds.). Symposium on Natural History collections, past, present, future. Proc. biol. Soc. Wash. 82,
585-601.
Acknowledgements. I am particularly grateful to Dr. H. S. Torrens (University of Keele) for providing me
with a great deal of information in the preparation of this bibliography. Dr. W. D. I. Rolfe (Hunterian
Museum, Glasgow University), Dr. W. H. C. Ramsbottom (IGS, Leeds), and Dr. C. D. Waterston (Royal
Scottish Museum) also gave me valuable information and together with Dr. Torrens and Dr. D. A. Bassett
(National Museum of Wales) kindly read a first draft of the manuscript. Dr. R. L. Paton (Royal Scottish
Museum) and Miss B. J. Pyrah (The Yorkshire Museum) allowed me to study their unpublished manu-
script catalogues, and Dr. M. K. Howarth (British Museum, Natural History) gave me access to manuscripts
in his care. I also thank the Library staffs at the National Museum of Wales, Geological Society of London,
and Department of Palaeontology at the British Museum (Natural History) for their help in obtaining and
checking numerous references. Miss G. Newton, Mrs. S. Thackray, and Mr. S. R. Howe helped to compile
and check the index.
MICHAEL G. BASSETT
Department of Geology
The National Museum of Wales
Typescript received 14 February 1975 Cardiff, CFl 3NP
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MID-CRETACEOUS ANGIOSPERM POLLEN
FROM SOUTHERN ENGLAND
AND NORTHERN FRANCE
by J. F. LAING
ABSTRACT. Angiosperm pollen grains are described from the Upper Albian to Middle Cenomanian strata of several
localities in southern England and northern France. Twenty-two species are described, of which the following
thirteen are new: Psilatricolpites rectilatibus, Retitricolpites amplifissus, R. crassitransennus, R. exiguiexemplum,
R. meumendum, R. promiscuus, R. sarthensis, R. subtilimaculatus, Psilatricolporites complanatius, Retitricolporites
ecommoyensis, R. insolitimorus, Triporopollenites curtisi, and T. worbarrowensis. A sequence of angiosperm pollen
assemblages is suggested and an attempt is made to relate this to the ammonite zonation.
The description of angiosperm pollen from the Albian and Cenomanian strata of
southern Britain and northern France is important since this is an area which includes
the stratotypes of both stages, and where the stratigraphy is understood in detail.
Previous studies on early angiosperm pollen assemblages (e.g. Doyle 1969; Pacltova
1971) have generally been concerned with areas where the age of the strata is impre-
cisely known. This is because correlation with the standard European sequence is
impossible, owing to a lack of any suitable marine fossils.
An exception to this is, perhaps, Dettmann’s (1973) recent study on the angiosperm
pollen from the eastern Australian Albian to Turonian, but here the sequence appears
to be rather different from that which occurs in contemporaneous European and
North American strata.
The present work represents an attempt to relate the sequence of angiosperm pollen
assemblages to the ammonite zonation of the Albian and Cenomanian of north-west
Europe, in the hope that it will help other workers to define better the stratigraphy in
areas where the mid-Cretaceous sequence occurs in non-marine strata.
LOCATION AND STRATIGRAPHY OF SAMPLES
Samples have been examined from six localities: Lulworth Cove, Worbarrow Bay,
Punfield Cove, and Compton Bay on the south coast of England; Saint-Jouin on the
coast of Normandy; and Ecommoy near Le Mans. All the samples collected are of
marine rocks, except those from Ecommoy which are of a non-marine or brackish
facies. The position of each locality is shown on text-fig. 1, and details of each section
are given in text-figs. 2-9.
PREPARATION TECHNIQUE
Samples were prepared as follows: crushing; removal of CaCOj with cold HCl;
removal of silicates with cold 60% HE ; removal of excess fluorides with warm
HCl; removal of remaining minerals by flotation in ZnBr2 solution (SG 2-5); short
[Palaeontology, Vol. 18, Part 4, 1975, pp. 775-808, pis. 90-94.]
776
PALAEONTOLOGY, VOLUME 18
TEXT-FIG. 1. Map showing localities sampled.
centrifuging; up to 30-minutes oxidation with cold concentrated HNO3; removal
of oxidation products with cold 5% NH4OH for 5 minutes.
Residues were mounted on eover slips using ‘Clearcol’, the cover slips then being
mounted on slides using ‘DePeX’. Exeess residues were stored in glycerine (plus
a few drops of phenol) in elosed plastic vials.
SYSTEMATIC PALYNOLOGY
A major problem with early angiosperm palynology is the small size of the pollen
grains and of their ornamentation. This difficulty seems to have deterred many
authors from producing descriptions of new species which are at all adequate for
comparative purposes, with the result that it becomes necessary to create many
new species.
Angiosperm pollen is rare in the strata which I have studied, especially so in the
marine roeks, and so it has not been possible to base my new species on as many
specimens as I would have wished. A further problem is that many of the features of
these grains are too small to be measured directly by light microscopy and for this
reason I have often had to estimate sizes. Thus where fractions of a micron have been
given these are only estimates.
Where I have attributed my specimens to a previously published species, I have used the scheme of
graded comparisons proposed by Hughes and Moody-Stuart (1967). When I have given the occurrences
of the specimens that I have found, I have done so in terms of the zonal schemes of Spath (1923-1943)
for the Albian, and of Kennedy (1969) for the Cenomanian, both summarized on Table 2. When I have
referred to the occurrences of other specimens, and to the known ranges of species, the stratigraphic informa-
tion is generally as stated by the original author; in areas lacking suitable zone fossils, this information may
be only approximately correct.
LAING: MID-CRETACEOUS ANGIOSPERM POLLEN
111
TEXT-FIG. 2. Section at Punfield Cove, Dorset text-fig. 3. Section at north end of Swanage Bay,
(SZ 045813). (Zonation after Hancock 1969; and Dorset (SZ 043812). Continuation of section at
Drummond 1970.) Punfield Cove. (Zonation after Wright, in Arkell
1947.)
TEXT-FIG. 4. Section c. 200 m SSW. of steps from text-fig. 5. Section c. 500 m NNE. of steps from
Valleuse Boucherot, Saint-Jouin, Seine-Maritime. Valleuse Boucherot, Saint-Jouin, Seine-Maritime.
(Zonation after Juignet 1972.) (Zonation after Juignet 1972.)
H
778
PALAEONTOLOGY, VOLUME 18
tompling
hptdon
UPPER
GREENSAND
I
I (■•How 9'sen,
rhplo/nogenie MIDDLE
Zona CENOMANIAN
photphole
nodulai
Text-fig. 6.
GlAUCONITtC
MARL
UPPER
GREENSAI'Jd
torcilonensit monleUi LOWER
Horiton Zone CENOMANIAN
phoipKaie nodulet
Text-fig. 7.
Fine, dork
yellowish
oronge sond
with greyish
clovey lentils
ona bonds
Medium light
S.grey cloy
and silt
Qfiofr
conglomeratic
boulders
rho tomogense MIDDLE
^ Zone CENOMANIAN
f— 2 Om
— l-5m
— 1-Om
“0 5m
Text-fig. 9.
TEXT-FIG. 6. Section at Lulworth Cove, Dorset
(SY 825800). (Zonation after Hancock 1969.)
TEXT-FIG. 7. Section at Compton Bay, Isle of Wight
(SY 366853). (Zonation after Kennedy 1969.)
TEXT-FIG. 8. Section at Worbarrow Bay, Dorset
(SY 862804). (Zonation after Wright, in Arkell
1947; and Kennedy, pers. comm.)
middle
CENOMANIAN
UPPER
ALBIAN
Text-fig. 8.
TEXT-FIG. 9. Section at Carriere de Bezonnais,
Ecommoy, Sarthe. (Zonation after Juignet and
Medus 1972.) Palynological comparisons with the
marine sequence suggest the Argiles noires to be
of approximately lower Middle Cenomanian age
(Laing 1973.)
LAING: MID-CRETACEOUS ANGIOSPERM POLLEN
779
The following abbreviations are used in the text ;
D
depth
NT
nexine thickness
DC
depth of colpus (colpi)
OA
oblique aspect
E
equatorial aspect
P
polar aspect
ED
equatorial diameter
PD
polar diameter
ET
exine thickness
s
standard deviation
L
length
ST
sexine thickness
LC
length of colpus (colpi)
W
width
LD
MW
diameter of lumina of microreticulum
width of muri of microreticulum
WC
width of colpus (colpi)
The occurrence of each species in each horizon and locality is summarized on Table 1.
All specimens are deposited in the Sedgwick Museum, Cambridge, where a catalogue of specimens may
also be found on ‘Taxon Register’ forms (form 6109). Co-ordinates given are for Leitz Dialux microscope
no. 469843, also in the Sedgwick Museum. Specimens shown in scanning electron micrographs are located
according to the co-ordinate reference system described by Laing (1974).
Anteturma pollenites Potonie, 1931
Turma plicates Naumova emend. Potonie, 1960
Subturma monocolpates Iversen and Troels-Smith, 1950
Genus asteropollis Hedlund and Norris, 1968
1968 Asteropollis Hedlund and Norris, p. 152.
Type species. Asteropollis asteroides Hedlund and Norris, 1968, p. 153, pi. 6, figs. 18-20; pi. 7, figs. 1-5.
Comments. Hedlund and Norris originally placed this genus in the subturma Polyp-
tyches. However, I consider that it is better regarded as being a monosulcate form
and I have accordingly transferred it to the subturma Monocolpates.
CfB. Asteropollis asteroides Hedlund and Norris, 1968
Plate 94, figs. 12-14
1968 Asteropollis asteroides Hedlund and Norris, p. 153, pi. 6, figs. 18-20; pi. 7, figs. 1-5.
Description of five specimens from samples EC03 and ECO 5. Sub-circular to circular in polar view, semi-
circular in equatorial view. Tri-, tetra-, penta-, or hexachotomosulcate; sulcus margins poorly defined and
rather ragged. Clear exine stratification into unstructured nexine and microreticulate sexine; sexine usually
thicker than nexine (occasionally nexine thicker than sexine); sexine shows no tendency to detach from
nexine. Microreticulum usually slightly imperfect (occasionally perfect) with some discrete clavae present ;
lumina all of about the same size; sexine absent in patches over surface of sulcus.
Dimensions. ED 21-26 /^tm (5), PD 13 ^^m (1), ET 1-2-1-5 jum (5), NT 0-5-0-8 (um (5), ST 0-5-10 /^m (5),
ST/NT 0-6-2-0 (5), LD 0-2 ^.im (5), MW OT-0-2 pm (5), sulcus diameter 15-20 pm (5), sulcus diameter/ED
0-7-0-8 (5), L rays of sulcus/sulcus diameter 0-2-0-4 (5), W rays of sulcus/sulcus diameter 0-2-0-4 (5).
Orientation. P 80 0%, E 20-0%.
Occurrence. This study: inflatum Zone, Upper Albian to costatus Horizon, rhotomagense Zone, Middle
Cenomanian. Other occurrences: Hedlund and Norris (1968), Middle Albian, Oklahoma; Dettmann
(1973), Cenomanian and Turonian, eastern Australia. Known range: Middle Albian to Turonian.
Comments. My attribution of these specimens to this species strictly requires an
emendation of the genus, which should only include tetra- and pentachotomosulcate
forms according to Hedlund and Norris’s diagnosis. However, I am reluctant to
780
PALAEONTOLOGY, VOLUME 18
make any formal emendation based on the few specimens which I have seen. The !
form described by Hedlund and Norris also differs in having the nexine slightly j
thicker than the sexine. j
\
Genus LiLiACiDiTES Couper, 1953 "
1953 Liliacidites Couper, p. 56. i,
1958 Clavatipollenites Couper, p. 159.
1961 Retimonocolpites Pierce, p. 47.
Type species. Liliacidites kaitangataensis Couper, 1953, p. 56, pi. 7, fig. 97.
Comments. Couper (1953) proposed the genus Liliacidites for monosulcate pollen
grains with reticulate exines, adding that the genus be for the reception of pollen
of liliaceous affinity which cannot be more accurately placed. The same author (1958)
proposed the genus Clavatipollenites for monosulcate grains with a sexine of clavate
projections which tend to expand and fuse at their tips to form a tectate exine. Couper
made no suggestion as to how this genus was to be distinguished from Liliacidites.
Kemp (1968) noted the morphological similarity of the two genera but implied that
Liliacidites be used for grains of unquestionable angiospermous origin, whereas
Clavatipollenites should be retained as a separate genus for forms of a more question-
able affinity. In my opinion, such a distinction, based on the supposed affinity of
a dispersed grain, has no place in palaeopalynology and any distinction should be
made on morphological grounds.
Dettmann (1973) attempted such a morphological distinction. She proposed that
Liliacidites should include forms with a differentially thickened exine and a sulcus
clearly formed in both the nexine and sexine. Clavatipollenites was proposed to
include forms with an exine of uniform thickness in which the sulcus is invariably
developed in the nexine but only occasionally in the sexine. I do not think that the
question of the presence or absence of differential exine thickening is sufficiently
important a character to justify the distinction of two genera. As for the question of
the development of the sulcus in the sexine, the holotype of the type species of
Clavatipollenites, C. liughesii, clearly has the sulcus developed in both the nexine
and sexine. This is also the case with C. rotundas Kemp, which Dettmann seems pre-
pared to leave in the genus Clavatipollenites. Thus, I do not think that the degree of
development of the sulcus in the sexine is a valid feature for the distinguishing of these |
two genera. I am thus of the opinion that no useful purpose is served by the retention j
of Clavatipollenites as a genus separate from Liliacidites.
CfA. Liliacidites peroreticulatus (Brenner) Singh, 1971 j
Plate 93. figs. 2-5
1963 Peromonoliles peroreticulatus Brenner, p. 94, pi. 41, figs. 1-2. ^ '
1971 Liliacidites peroreticulatus (Brenner) Singh, p. 188, pi. 28, figs. 6-1 1.
De.scription of twenty-one speeimens from samples ECO 2, ECO 4, and ECO 5. Oval in polar view; semi- |
circular to oval, sub-circular or circular in equatorial view. Monosulcate; sulcus parallel-sided or gaping
with pointed ends; sexinal margins usually entire (occasionally slightly ragged); nexinal margins entire,
often slightly infolded. Clear exine stratification into unstructured nexine and microreticulate sexine, which
are often partially or completely separated from each other by a cavity ; where separation has not occurred, :
LAING: MID-CRETACEOUS ANGIOSPERM POLLEN
781
the sexine is attached to the nexine by bacula, the muri being swollen over the point of attachment. Micro-
reticulum perfect, lumina of varying size and of irregularly polygonal shape.
Dimensions. L (whole grain) 17 (20-4) 26 jum (15), L (nexinal inner body) 14 (17-4) 21 ,um (15), W (whole
grain) 13 (18-8) 23 fxm (17), W (nexinal inner body) 10 (15-2) 21 ;um (17), D (whole grain) 12-19 jj.m (6),
D (nexinal inner body) 10-20 ixm (6), L/W (whole gram) 10-1-6 (6), L/W (nexinal inner body) 10-1-9 (6),
ET (where nexine and sexine are still in contact) 10 (1-6) 2 0 jj.m (21), NT 0-5 (0-7) 1 0 ;um (21), ST 0-5 (0-8)
10 ^^m (21), ST/NT 0-5 (1-2) 2-0 (21), LD (least) 0-2 (0-8) 1-5 fem to (greatest) 1-5 (2-5) 4 0 /xm (21), MW 0-2
(0-4) 10 /xm (21), sulcus L 12-18 /xm (8), sulcus L/L (whole grain) 0-7-0-9 (8), sulcus W 0-5 (2-8) 1 10 /ixm
(19), sulcus W/W (whole grain) <01 (0-2) 0-5 (15), sulcus D 0-5 (21) 5-0 /xm (14), sulcus D/D (whole
grain) < OT-0-3 (6).
Orientation. P 381%, equatorial transverse aspect 28-6%, equatorial longitudinal aspect 0 0%, OA 33-3%.
Occurrence. This study: inflatum Zone, Upper Albian to costatus Horizon, rhotomagense Zone, Middle
Cenomanian. Other occurrences: Brenner (1963), Upper Barremian to Albian, Maryland; Norris (1967),
Late Albian to ?Cenomanian, Alberta ; Brenner ( 1 968), Albian, Peru ; Agasie ( 1 969), Cenomanian, Arizona ;
Singh (1971), Middle to Upper Albian, Alberta; Azema, Durand and Medus (1972), Middle Cenomanian,
France. Known range: Upper Barremian to Cenomanian.
TABLE 1. Presence ( + ) or absence ( — ) of individual species at
each locality and horizon. A, inflatum Zone, Punfield Cove;
B, inflatum Zone, Saint-Jouin; C, dispar Zone, Saint-Jouin;
D, carcitanensis Horizon, Saint-Jouin; E, carcitanensis
Horizon, Compton Bay; F, costatus Horizon, Punfield Cove;
GJukes-brownei Horizon, Worbarrow Bay; HJukes-brownei
Horizon, Lulworth Cove; I,
Argiles
noires
of Ecommoy.
Locality
A
B
C
D
E
F
G
H
I
A. asteroides
—
-b
—
—
—
+
—
—
+
L. peroreticulatus
+
+
—
+
+
+
—
—
+
L. rotundus
+
-b
-b
+
+
4-
—
—
-b
P. erugatus
—
—
—
—
-b
—
—
—
P. rectilatibus
—
—
-f
—
—
—
+
+
+
R. amplifissus
—
+
+
+
+
—
+
+
+
R. crassitransennus
+
+
-b
R. exiguiexemplum
-
+
+
—
—
+
—
—
+
R. georgensis
—
-b
—
+
-f
-b
+
+
+
R. meumendum
—
+
+
—
—
—
—
—
+
R. nemejci
—
—
+
—
—
—
—
—
+
R. promiscuus
—
-f
+
+
—
+
+
—
+
R. sarthensis
—
+
—
+
—
+
-b
—
+
R. subtilimaculatus
—
+
—
—
+
—
—
—
+
Retitricolpites sp 1
-I-
+
—
—
+
+
—
—
+
S. sarstedtensis
—
+
—
—
+
—
—
+
P. complanatius
-
-
-
-
-b
-b
—
—
+
R. ecommoyensis
—
+
—
—
+
+
+
—
-b
R. insolitimorus
-b
—
+
+
' +
-b
—
+
-b
C. subtilis
+
—
T. curtisi
—
—
—
—
—
—
+
—
—
T. worbarrowensis
—
—
—
—
—
—
-b
—
—
Comments. Distinction from L. rotundus (Kemp) is sometimes a little difficult.
Generally, however, L. rotundus has a finer-meshed microreticulum with narrower
muri, and has the muri composed of bacula or clavae rather than being supported
by bacula.
782
PALAEONTOLOGY, VOLUME 18
Brenner (1963) described two similar species from the Potomac Group, Pero-
monolites reticulatus and P. peroreticulatus, stating that they were distinguishable
on the basis of grain size and lumen diameter. However, the specimens found in the
present study overlap the ranges for these characters of both of Brenner’s species.
I have examined some material from the Patapsco Formation of the Potomac Group
and have found that these species differ as follows : P. peroreticulatus has more or
less polygonal lumina and muri with a beaded surface, whereas P. reticulatus has
more sinuous muri, such that the lumina are not so polygonal, the muri having
smooth surfaces.
CfB. Liliacidites rotundus (Kemp 1968) comb. nov.
Plate 90, figs. 1 -6
1963 Liliacidites dividuus (Pierce) Brenner, p. 93, pi. 40, figs. 7-10.
1968 Clavatipollenites rotundus Kemp, p. 424, pi. 79, figs. 1-19; pi. 80, figs. 1-8; text-fig. 2.
Description of seven specimens from samples JOU 3 and JOU 4. Oval to sub-circular in polar view, oval in
equatorial longitudinal view, sub-circular or oval in equatorial transverse view. Monosulcate, sulcus slit-
like or gaping, margins entire. Clear exine stratification into unstructured nexine and microreticulate
sexine composed of clavae or bacula, sexine occasionally tending to detach from nexine. Microreticulum
perfect, lumina generally of varying size (very rarely of more uniform size) and irregularly polygonal shape.
Grain outlines generally finely indented (sometimes almost smooth).
Dmensions. L 18-29 /urn (4), W 16-22 fxm (4), D 14-22 /uin (4), L/W M-L3 (2), D/W 0-7-1 -2 (2), L/D
M ( 1 ), ET 1 -5-2-5 ;nm (7), NT 0-5- 1 -5 fxm (7), ST 0-9- 1 -0 pm (7), ST/NT 0-7-2-0 (7), LD (least) 0- 1 -0-3 /xm
to (greatest) 0-3-1-5 pm (7), MW 0-2-0-3 pm (7), sulcus L 13-22 pm (4), sulcus L/L 0-7-0-8 (4), sulcus
W 2-10 (um (4), sulcus W/W 01-0-5 (4), sulcus D 1-9 pm (4), sulcus D/D 01-0-6 (4).
Orientation. P 28-6%, equatorial transverse aspect 28-6%, equatorial longitudinal aspect, 14-3%, OA 28-6%.
Occurrence. This study: inflatum Zone, Upper Albian to costatus Horizon, rhotomagense Zone, Middle
Cenomanian. Other occurrences: Brenner (1963), Albian, Maryland; Hedlund (1966), Cenomanian,
Oklahoma; Brenner (1967), Cenomanian, New Jersey; Hedlund and Norris (1968), Middle Albian,
Oklahoma; Kemp (1968, 1970), Lower Albian (Itardefurcata Zone) to Upper Albian, England; Habib
(1969), Middle Cretaceous, sea bed near the Bahamas; Playford (1971), Albian, Saskatchewan and Mani-
toba; Singh (1971), Albian to Cenomanian, Alberta; Azema, Durand and Medus (1972), Middle Ceno-
manian, France. Known range: Itardefurcata Zone, Lower Albian to Cenomanian.
EXPLANATION OF PLATE 90
All figures x 2000.
Figs. 1-6. CfB. Liliacidites rotundus (Kemp). 1-2, oblique aspect; JOU 4, JL 169.1 ; 26.9, 106.9; 1, median
focus; 2, low focus. 3-4, specimen with sexine partially detached from nexine; distal polar aspect;
JOU 12, JL 177.1; 25.6, 104.6; 3, high focus; 4, median focus. 5-6, specimen with darkened sulcus
borders; proximal polar aspect; ECO 6, JL 188.2; 24.1, 098.9; 5, median focus; 6, low focus.
Figs. 7-8. CfC. PsHatricolpites erugatus {\\Qd\ux\d). Near equatorial aspect; ECO 5, JL 187.3; 34.1, 108.3;
7, high focus; 8, median focus.
Figs. 9-12. Psilatricolpites rectilatihus sp. nov. 9- 10, holotype; polar aspect: ECO 5, JL 187.2; 59.0, 1 10.8;
9, high focus; 10, median focus. 1 1-12, oblique aspect; ECO 5, JL 209.3; 35.6, 101.2; 1 1, median focus;
12, high focus.
Figs. 13-16. Retitricolpites amplifissus sp. nov. 13-14, holotype; oblique aspect; ECO 5, JL 209.2; 40.8,
096.6; 13, high focus to show ornament; 14, median focus. 15- 16, small specimen; polar aspect; ECO 5,
JL 187.3; 41.4, 097.0; 15, high focus to show ornament; 16, median focus.
PLATE 90
LAING, angiosperm pollen
784
PALAEONTOLOGY, VOLUME 18
Comments. See comments for L. peroreticulatus for distinction from this species.
Kemp (1968) attempted to distinguish this species from Liliacidites (al. Clavati-
pollenites) hughesii (Couper 1958, p. 159, pi. 31, figs. 19-22 emend. Kemp 1968,
p. 426, pi. 80, figs. 9-19) comb. nov. on the basis of size-range and shape. However,
I have found that the two species seem to show too great an overlap in the ranges
of these characters to allow a reliable distinction to be made on this basis. Kemp
further suggested that the nature of the sulcus could be used to differentiate the
species, the sulcus having entire margins with darkened borders in L. rotundus, and
being indistinct or with ragged margins and no darkened borders in L. hughesii.
Although I have found a few specimens of L. rotundus with darkened sulcus borders
(e.g. PI. 90, figs. 5-6), it is the exception in the specimens that I have observed ; further-
more, the holotype of L. hughesii has a distinct sulcus with entire margins. Thus
I do not consider that the nature of the sulcus should be used as a distinguishing
character. In my opinion the most useful distinguishing character is (as Kemp also
stated) the degree of development of the microreticulum, it being perfect in L. rotundus
and imperfect in L. hughesii.
Retimonocolpites dividuus Pierce, 1961, was poorly described and figured. Pierce
did, however, state that the aperture (i.e. sulcus) almost encircles the grain, dividing
it into two hemispheres; on the basis of this feature, Pierce’s species seems to be
distinct. Brenner (1963) and later authors (Hedlund 1966; Hedlund and Norris 1968;
Brenner 1967; Habib 1969; and Singh 1971) described or figured forms identified as
Pierce’s species, which seem to more closely resemble L. rotundus.
Subturma triptyches Naumova emend. Potonie 1960
Several genera have been used by various authors for early tricolpate pollen. I find
the genera of van der Hammen (1956) the most convenient, at least for light micro-
scope studies. For S.E.M. studies, different genera may be preferred based on the
greater structural variation which can be observed, and for practical purposes it may
become necessary to have separate taxonomies for each mode of observation (in
which case the S.E.M. taxonomy will probably be the correct ‘formal’ one, and the
light microscope taxonomy will be rather more informal).
Genus psilatricolpites van der Hammen ex van der Hammen and Wymstra, 1964
1956 Psilatricolpites van der Hammen, p. 88.
1964 Psilatricolpites van der Hammen ex van der Hammen and Wymstra, p. 234.
Type species. Psilatricolpites clarissimus (van der Hammen) emend, van der Hammen and Wymstra, 1964,
p. 235, pi. 2, fig. 2.
CfC. Psilatricolpites erugatus (Hedlund 1966) comb. nov.
Plate 90, figs. 7-8
1966 Tricolpites erugatus Hedlund, p. 30, pi. 9, fig. 2a-b.
1967 Psilatricolpites parvulus (Groot and Penny) Norris, p. 107, pi. 17, figs. 5-6.
Description of thirty-two specimens from samples ECO 1, ECO 3, ECO 4, ECO 5, and ECO 7. Perprolate
to prolate (very occasionally prolate spheroidal); elliptical in equatorial view. Three distinct (occasionally
LAING: MID-CRETACEOUS ANGIOSPERM POLLEN
785
indistinct) slit-like (occasionally more gaping) colpi with entire (occasionally more ragged) margins. No
clear exine stratification, Psilate.
Dimensions: PD 10 (12-2) 17 fxm (27) s2-0 ;um, ED 5 (7-6) 1 1 /xm (32) sl-4 /Ltm, PD/ED 11 (1-6) 2-3 (27),
ET 0-2 (0-7) M ^m (32), EC 7 (91) 13 /^m (27), LC/PD 0-5 (0-7) 0-9 (27).
Orientation. P 0 0%, E 844%, OA 15-6%.
Occurrence. This study: carcitanensis Horizon, mantelli Zone, Lower Cenomanian to rhotomagense Zone
0-costatus Horizon), Middle Cenomanian. Other occurrences: Hedlund (1966), Cenomanian, Oklahoma;
Norris (1967), Late Albian, Alberta; Singh (1971), Upper Albian and Cenomanian, Alberta; Azema,
Durand and Medus (1972), Middle Cenomanian, France. Known range: Upper Albian and Cenomanian.
Comments. It is difficult to decide whether this form is more closely referable to
Tricolpopollenites parvulus Groot and Penny, 1960 or Tricolpites erugatus Hedlund,
1966. Groot and Penny’s description of T. parvulus stated ‘grains about isodiametric,
usually seen in polar view’, which seems to suggest that this species should have a
spheroidal shape (although one of Groot and Penny’s figures, pi. 42, fig. 9, shows
a clearly prolate form). Norris (1967) did not describe this species but, of the two
specimens which he figured, one is prolate and the other more or less spheroidal. The
forms described as Psilatricolpites parvulus by Singh (1971) are ‘sub-prolate to almost
isodiametric’. I have placed the specimens that I have found in Hedlund’s species
since this definitely includes prolate forms. However, Hedlund included both smooth
and microgranulose specimens in this species, and the latter condition has not been
observed in the specimens found in the present study.
P. rectilatibus sp. nov. differs by generally having a greater equatorial diameter
and a more spheroidal or oblate shape. It also seems to be more deeply trilobate.
Psilatricolpites rectilatibus sp. nov.
Plate 90, figs. 9-12
Description of fourteen specimens from samples ECO 5 and ECO 7. Spheroidal to oblate; deeply trilobate,
more or less straight-sided triangular outline in polar view. Three distinct long slit-like or gaping colpi
generally with entire margins, angul-aperturate. No clear exine stratification. Psilate.
Holotype. Sample ECO 5, slide JL 187.2; 59.0, 110.8. Plate 90, figs. 9-12. Argiles noires, Ecommoy;
rhotomagense Zone, Middle Cenomanian.
Dimensions. ED 10 (12T) 15 /xm (14), ET 0-2 (0-7) 10 /xm (14), WC 0-2 (2-6) 5-0 ;ixm (4), WC/ED <01
(0-2) 0 4(4), DC 01 (l-6)3 0;ixm (4), DC/ED <01 (01) 0-2 (4).
Orientation. P 28-6%, E 0-0%, OA 714%.
Occurrence. This study: dispar Zone, Upper Albian to jukes-hrownei Horizon, rhotomagense Zone, Middle
Cenomanian.
Comments. Tricolpopollenites parvulus Groot and Penny, 1960 was not described
adequately enough to allow a close comparison ; it does seem, however, to differ from
this species in having short colpi. Tricolpites pachyexinus Couper, 1953 differs by
being larger, by having a much thicker exine and by occasionally being tetracolpate.
T. gillii Cookson, 1957 differs by being larger and by having a finely granular tectate
exine. Tricolporopollenites triangulus Groot, Penny and Groot, 1961 differs by having
pores. Psilatricolpites tetradus Brenner, 1968 differs by having a more circular amb,
exine stratification, and by most commonly occurring in tetrads. See comments for
P. erugatus for distinction from this species.
786
PALAEONTOLOGY, VOLUME 18
Genus retitricolpites van der Hammen ex van der Hammen and Wymstra, 1964
1956 Retitricolpites van der Hammen, p. 90.
1964 Retitricolpites van der Hammen ex van der Hammen and Wymstra, p. 234.
Type species. Retitricolpites ovalis van der Hammen and Wymstra, 1964, p. 234, pi. 1, figs. 5-6.
Retitricolpites amplifissm sp. nov.
Plate 90, figs. 13-16; Plate 91, figs. 1-2
1968 Tricolpites sp. 2 Kemp, p. 432, pi. 81, figs. 23-24.
Description of sixteen specimens from samples ECO 3, ECO 4, ECO 5, and ECO 7. Oblate, perhaps to
spheroidal; circular to sub-triangular, deeply trilobate outline in polar view. Three distinct, quite short
colpi which gape at the equator, margins entire to ragged. Clear exine stratification into unstructured
nexine and microreticulate sexine; sexine of equal thickness to, or more commonly thicker than, nexine.
Microreticulum most commonly perfect, but sometimes imperfect; lumina generally of varying size (but
sometimes of fairly constant size) and irregularly polygonal in shape. Some specimens with a smooth inner
central body (see PI. 91, figs. 1-2). Grain outlines finely indented.
Holotype. Sample ECO 5, slide JL 209.2; 40.8, 096.6. Plate 90, figs. 13-14. Argiles noires, Ecommoy;
rhotomagense Zone, Middle Cenomanian.
Dimensions. Type material: ED 11 (19-9) 27 fim (16), ET 10 (T3) 2-0 /xm (16), NT 0-2 (0-5) 10 [xm (16),
ST 0-5 (0-9) 10 fxm (16), ST/NT 0-9 (2-1) 5-0 (16), LD (least) 0-1 (0-2) 0-8 ,xm to (greatest) 01 (0-9) 1-5 /xm
(16), MW 01 (0-2) 0-3 /xm (16), WC 2 0 (4-9) 8-0 /xm (8), WC/ED 01 (0-2) 0-3 (8), DC 2-0 (4-9) 9-0 /xm (8),
DC/ED OT (0-2) 0-3 (8). Other material: Sample JOU 4, ED 15-23 /xm (6); Sample JOU 15, ED 16-
22 /xm (5).
Orientation. Type material; P 50 0%, OA 50-0%.
Occurrence. This study; inflatum Zone, Upper Albian to jukes-brownei Horizon, rhotomagense Zone,
Middle Cenomanian. Other occurrences: Kemp (1968, 1970), Lower Albian (Itardefurcata Zone), England.
Known range : Itardefurcata Zone, Lower Albian to jukes-brownei Horizon, rhotomagense Zone, Middle
Cenomanian.
EXPLANATION OF PLATE 91
All figures x 2000.
Figs. 1-2. Retitricolpites amplifissus sp. nov. Large specimen with inner central body; polar aspect; ECO 4,
JL 1 84. 1 ; 3 1 . 1 , 111.6; 1 , low focus to show ornament ; 2, median focus to show inner central body.
Figs. 3-8. Retitricolpites crassitransennus sp. nov. 3-4, holotype; polar aspect; ECO 5, JL 209.1; 40.1,
101.0; 3, high focus to show ornament; 4, median focus. 5-6, oblique aspect; ECO 5, JL 209.1; 41.3,
108.8; 5, median focus; 6, low focus. 7-8, equatorial aspect; ECO 5, JL 209.3; 32.8, 103.5; 7, high focus;
8, median focus.
Figs. 9-10. Retitricolpites exiguiexemplum sp. nov. Holotype; oblique aspect; ECO 5, JL 209.1; 47.4,
099.1 ; 9, high focus; 10, median focus.
Figs. 11-12. CfB. Retitricolpites georgensis Brenner. Equatorial aspect; ECO 7, JL 191.1; 29.5, 097.8;
11, median focus; 12, low focus to show ornament.
Figs. 13-14. Retitricolpites meumendum sp. nov. Holotype; equatorial aspect; ECO 5, JL 209.3; 42.1,
108.8; 13, median focus; 14, low focus to show ornament.
Figs. 15-16. CfC. Retitricolpites nemejci (Pacitova). Equatorial aspect; ECO 5, JL 209.1; 44.4, 100.6;
15, median focus; 16, low focus.
Figs. 17-18. Retitricolpites promiscuus sp. nov. Holotype; specimen with lumina of varying size; equatorial
aspect; ECO 5, JL 187.2; 26.4, 101.4; 17, median focus; 18, low focus to show ornament.
r/
PLATE 91
LAING, angiosperm pollen
788
PALAEONTOLOGY, VOLUME 18
Comments. The size distribution of the type material suggests that two species could
be present, one with an average equatorial diameter of about 17 /xm, the other of
about 25 /xm, but insufficient specimens have been found to show this conclusively.
Retitricolpites crassitransennus differs by having slightly longer colpi, a generally
finer-meshed microreticulum, wider muri, and undulating grain outlines. R. pro-
miscuus has a generally more prolate shape and longer, narrower, more slit-like colpi.
R. vulgaris Pierce, 1961 and R. oblatoides Pierce, 1961 were both inadequately
described; either could be conspecific with this species. Tricolpopollenites platyreti-
culatus Groot, Penny and Groot, 1961 is a more coarsely reticulate form. Tricolpites
sp. 2 of Agasie (1969) is, on average, a rather larger form, with a more straight-
sided triangular outline and a microreticulum which becomes finer over the poles
and at the colpus margins. Tricolpites heusseri Kimyai, 1966 seems to differ by having
narrower colpi. T. sagax Norris, 1967 differs by sometimes being sub-prolate, by
sometimes having a sub-granular exine, and by generally having narrower colpi.
Tricolpites sp. A of Pacltova (1971) seems to differ in having a sexine of closely spaced
(presumably discrete) pila rather than a microreticulum. R. maximus Singh, 1971 is
larger and has thicker, more vermiculate muri and a more prolate shape. T. reticulata
Cookson, 1947 is a generally rather larger form. T. cooksonae Dettmann, 1973 has
a generally slightly coarser-meshed microreticulum with wider muri.
Retitricolpites crassitransennus sp. nov.
Plate 91, figs. 3-8
Description of sixteen specimens from samples ECO 1, ECO 5, and ECO 6. Oblate to prolate; circular to
sub-triangular, quite deeply trilobate outline in polar view, elliptical to sub-circular in equatorial view.
Three distinct colpi which gape at the equator, margins entire to slightly ragged. Clear exine stratification
into unstructured nexine and microreticulate sexine; sexine of equal thickness to, or more commonly,
thicker than nexine (very rarely sexine thinner than nexine). Microreticulum perfect; lumina usually of
uniform size (rarely of varying size), irregularly shaped; muri sometimes of varying width, usually wider
than lumina. Grain outlines undulating.
Holotype. Sample ECO 5, slide JL 209.1 ; 40.1, 101.0. Plate 91, figs. 3-4. Argiles noires, Ecommoy; rhoto-
magense Zone, Middle Cenomanian.
Dimensions. PD 17-22 ;ixm (3), ED 13 (19-3) 27 ;xm(16), PD/ED 10-L4(3), ET 10 (1-5) 21 ^xm(16), NT
0-2 (0-6) M ^m (16), ST 0-5 (0-9) 1-2 p.m (16), ST/NT 0-5 (2-2) 6 0 (16), LD 0-2 (0-4) 10 pm (16), MW
0-2 (0-6) 10 /xm (16), LC 10-18 ^m (3), LC/PD 0-7-0-8 (3), WC 2 0-6 0 pm (4), WC/ED 01-0-3 (4), DC
2-0-5-0 ;um (4), DC/ED 01 -0-2 (4).
Orientation. P 25-0%, E 18-8%, OA 56-3%.
Occurrence. This study : rhotoniagense Zone, Icostatus Horizon to jukes-brownei Horizon, Middle Ceno-
manian.
Comments. See comments for Retitricolpites amplifissus and CfC. R. nemejci for
distinction from these species. Tricolpopollenites virgeus Groot, Penny and Groot,
1961 has a coarser-meshed reticulum and is a little larger. R. fragosus Hedlund and
Norris, 1968 is a little smaller and also differs in having the lumina reduced in size
on the mesocolpia. Tricolpites harrandei Pacltova, 1971 is smaller and perhaps has
a rather coarser-meshed microreticulum. Foveotricolpites concinnus Singh, 1971
differs by always being prolate, by having wider muri, and a coarser-meshed reticulum,
and by being rather larger.
LAING; MID-CRETACEOUS ANGIOSPERM POLLEN
789
Retitricolpites exiguiexemplum sp. nov.
Plate 9 1 , figs. 9-10; text-figs. 10-11
Description of nine specimens from samples ECO 3, ECO 5, and ECO 7. Spheroidal. Three distinct long
deep colpi, most commonly narrow and slit-like, less commonly wider and gaping at the equator, margins
entire. Clear exine stratification into unstructured nexine and microreticulate sexine; nexine and sexine
usually of equal thickness (occasionally sexine thicker than nexine). Microreticulum usually perfect
(occasionally imperfect with some free bacula) ; lumina of approximately uniform size and equidimensional
shape; some specimens psilate over the poles. Grain outlines smooth to slightly indented.
Holotype. Sample ECO 5, slide JL 209.1; 47.4, 099.1. Plate 91, figs. 9-10. Argiles noires, Ecommoy;
rhotomagense Zone, Middle Cenomanian.
Dimensions. ED 7-12 fxm (9), ET 0-8-1 -8 /xm (9), NT 0-2-0-9 ;um (9), ST 0-4-0-9 /xm (9), ST/NT 1-0-4-0 (9),
ED < 01-0-2 fxm (9), MW < 01-0-1 (9).
Orientation. OA 100-0%.
Occurrence. This study: inflatum Zone, Upper Albian to costatus Horizon, rhotomagense Zone, Middle
Cenomanian.
jiiimininirm
immmiin.
TEXT-FIG. 10. Cross-sections of sexine of
Retitricolpites subtilimaculatus sp. nov. (left)
and R. exiguiexemplum sp. nov. (right).
LOBE
TEXT-FIG. 1 1 . Holotypes of Tricolpites albiensis
Kemp (left) and Retitricolpites exiguiexemplum
sp. nov. (right), x 1000.
Comments. Retitricolpites meumendum is rather more prolate and has shallower
colpi. Distinction from R. subtilimaculatus is sometimes difficult. The main difference
seems to lie in the nature of the sexine ; in the case of R. subtilimaculatus the sculptural
elements stand up separately from each other to give a more indented outline (see
text-fig. 10), whereas in R. exiguiexemplum the sculptural elements appear more
fused, so that the outline is less indented. Tricolpites albiensis Kemp, 1968 is a similar
form. It differs from this species in having less rounded ‘lobes’ (i.e. the area between
the colpi— see text-fig. 1 1 ). Also, in the case of T. albiensis the sexine shows a tendency
to thicken as it passes over the centre of the lobe (this feature is well shown in Kemp’s
(1968) pi. 81, fig. 7); such a tendency is less apparent in R. exiguiexemplum. Judging
by Kemp’s description, a further difference is that T. albiensis has a coarser-meshed
microreticulum (LD 0-3-0-4 pm) which is imperfect: however, the holotype appears
to have a perfect microreticulum of LD 0-2 ,um. Further differences are that T. albi-
ensis includes quite strongly prolate forms and is on average a little larger than
R. exiguiexemplum. Tricolpopollenites micromunus Groot and Penny, 1960 was
inadequately described; it might possibly be conspecific with this species. Tricolpites
reticulominutus Jardine and Magloire, 1965 differs from this species by being some-
what larger.
790
PALAEONTOLOGY, VOLUME 18
CfB. Retitricolpites georgensis BrtnnQV, 1963
Plate 91, figs. 1 1-12; text-fig. 12
1963 Retitricolpites georgensis Brenner, p. 91, pi. 38, figs. 6-7.
1973 Rousea georgensis (Brenner) Dettmann, p. 14, pi. 2, figs. 16-17.
Description of eight specimens from samples ECO 1, ECO 3, ECO 5, and ECO 7. Sub-prolate to prolate;
elliptical in equatorial view. Three distinct, narrow slit-like colpi, margins entire. Clear exine stratification
into unstructured nexine and microreticulate sexine; sexine of equal thickness to, or thicker than, nexine
(very rarely sexine thinner than nexine). Microreticulum perfect; lumina of varying size, usually becoming
smaller towards the poles, irregularly shaped; muri sometimes of varying width, usually narrower than
lumina. Grain outlines notched or coarsely indented at equator, becoming smoother towards the poles.
Dimensions. PD 15-26 (5), ED 10-16 /um (8), PD/ED M-1-6 (5), ET 10- 1-8 /xm (8), NT 0-5- TO /xm (8),
ST 0'5-lT /xm (8), ST/NT 0-5-2-8 (8), LD (least) 0-2-0-5 ju.m to (greatest) 10-2-5 pm (8), MW 0-2-0-8 /xm
(8), LC 9 (13-3) 20 ^m (5), LC/PD 0-6 (0-7) 0-8 (5).
Orientation. P 0 0%, E 62-5%, OA 37-5%.
Occurrence. This study: inflatum Zone, Upper Albian to jukes-brownei Horizon, rhotomagense Zone,
Middle Cenomanian. Other occurrences: Brenner (1963), Albian, Maryland; Norris (1967), Late Albian,
Alberta; Habib (1969), Middle Cretaceous, sea bed near the Bahamas; Paden Phillips and Felix (1971),
Cenomanian, Louisiana; Playford (1971), Middle and ?Late Albian, Saskatchewan and Manitoba; Singh
(1971), Middle Albian to Cenomanian, Alberta; Azema, Durand and Medus (1972), Middle Cenomanian,
France; Dettmann (1973), Late Albian and Cenomanian, eastern Australia. Known range: Albian and
Cenomanian.
Comments. The specimens described by Brenner (1963) are quite similar, but are, on
average, larger (PD 18 (26) 36 ;ixm, ED 17 (22) 28 ju.m) and show a smaller range of
variation in the lumina diameter (0-5- 1-5 /xm). See comments for Retitricolpites
crassitransennus and Retitricolpites sp. 1 for distinction from these species. R. sar-
thensis differs by having a finer-meshed microreticulum, a thinner exine either with
unclear stratification or a thinner nexine relative to the sexine, and finely indented
grain outlines. R. promiscuus has a rather finer-meshed microreticulum and more
finely indented grain outlines. CfC. R. nemejci differs by having generally rather
longer colpi, a finer-meshed more regular microreticulum, a generally thinner exine,
a thinner nexine as compared to the sexine, and more finely indented grain outlines.
Retitricolpites meumendum sp. nov.
Plate 91, figs. 13-14
Description of twelve specimens from sample ECO 5. Prolate to sub-prolate (? perhaps to prolate spheroidal) ;
elliptical or sub-circular in equatorial view. Three generally distinct narrow slit-like shallow colpi, margins
entire (rarely ragged). Exine usually clearly stratified into unstructured nexine and a sexine which is usually
psilate at the poles and microreticulate elsewhere, exine stratification unclear in some specimens; sexine
usually thicker than nexine. Microreticulum perfect but not strongly developed; lumina of approximately
uniform size and equidimensional shape. Grain outlines smooth at poles, elsewhere very finely indented.
Holotype. Sample ECO 5, slide JL 209.3; 42.1, 108.8. Plate 91, figs. 11-12. Argiles noires, Ecommoy;
rhotomagense Zone, Middle Cenomanian.
Dimensions. PD 10-15 /xm (8), ED 7 (9-3) 14 /un (12), PD/ED T3-T9 (8), ET 0-4 (0-6) 11 ,-m (12), NT
0- 1 -0-2 /xm (8), ST 0-3-0-9 /xm (8), ST/NT 2 0-5 0 (8), LD < 0- 1 (0-2) 0-4 /xm ( 1 2), MW < 0- 1 (0- 1 ) 0-2 /xm
(12), LC 8 (9-5) 13 /xm (8), LC/PD 0-6 (0-8) 0-9 (8).
Orientation. P 0 0%, E 66-7%, OA 33-3%.
LAING: MID-CRETACEOUS ANGIOSPERM POLLEN
791
Occurrence. This study : inflatum Zone, Upper Albian to rhotomagense Zone (? costatus Horizon), Middle
Cenomanian.
Comments. See comments for Retitricolpites exiguiexemplum for distinction from
this species. CfC. R. nemejci is generally rather larger, is less commonly psilate at the
poles, and has a more distinct microreticulum with generally wider muri. R. sub-
tilimaculatus has rather deeper colpi, the microreticulum developed all over the
grain, and a thicker exine.
CfC. Retitricolpites nemejci (Pacltova 1971) comb. nov.
Plate 91, figs. 15-16
1971 Tricolpites nemejci Pacltova, p. 113, pi. 4, figs. 1-5; pi. 5, figs. 1-12; pi. 6, figs. 1-12.
Description of five specimens from sample ECO 5. Sub-prolate to prolate; elliptical in equatorial view. Three
distinct parallel-sided colpi, margins entire. Clear exine stratification into unstructured nexine and micro-
reticulate sexine (sexine very occasionally psilate at the poles) ; sexine thicker than nexine. Microreticulum
perfect; lumina either of approximately uniform or of slightly varying size, irregularly polygonal in shape;
muri often of varying width. Grain outlines finely indented (very occasionally smooth at the poles).
Dimensions. PD 16-24 /urn (5), ED 9-16 /nm (5), PD/ED 1-3- 1-8 (5), ET 0-6- 10 /iim (5), NT OT-0-2 jum (5),
ST 0-5-0-8 fxm (5), ST/NT 4-0-7-0 (5), ED 01-0-4 ^.m (5), MW 0-2-L0 ,xm (5), EC 12 (17-1) 22 ixm (5),
LC/PD 0-8 (0-8) 0-9 (5).
Orientation. E 100 0%.
Occurrence. This study: dispar Zone, Upper Albian to rhotomagense Zone (? costatus Horizon), Middle
Cenomanian. Other occurrence: Pacltova (1971), Cenomanian, Bohemia. Known range: dispar Zone,
Upper Albian and Cenomanian.
Comments. Some of the specimens figured by Pacltova (1971) are similar to the forms
described here. Pacltova described the shape as being oblate to sub-prolate although
some of her figured specimens are clearly prolate. Pacltova’s specimens are, on
average, rather larger (PD 23 (25) 28 p.m) and have slightly coarser microreticula than
the specimens found in this study. See comments for CfB. Retitricolpites georgensis
and R. meumendum for distinction from these species. R. crassitransennus differs by
being more spheroidal and by having slightly shorter colpi, a generally slightly
coarser-meshed microreticulum, a generally thicker exine, and undulating grain
outlines. R. promiscuus has a coarser-meshed microreticulum, a generally thicker
exine, nexine and sexine typically of about equal thickness, and possibly has narrower
colpi. Foveotricolpites sphaeroides Pierce, 1961 and Querco'idites sp. of Azema and
Ters (1971) seem to be similar forms but both were inadequately described for com-
parative purposes. Retitricolpites sp. B of Hedlund and Norris (1968) also seems to
be similar, but it was undescribed.
Retitricolpites promiscuus sp. nov.
Plate 91, figs. 17-18; Plate 92, figs. 1-2
Description of fifty-four specimens from samples ECO 1, ECO 2, ECO 5, and ECO 6. Spheroidal to prolate;
sub-circular trilobate outline in polar view, elliptical in equatorial view. Three, usually distinct, slit-like
(occasionally more gaping) colpi, margins entire or slightly ragged. Clear exine stratification into un-
structured nexine and microreticula te sexine ; nexine and sexine usually of about equal thickness (occasionally
792
PALAEONTOLOGY, VOLUME 18
sexine a little thicker than nexine). Microreticulum perfect; lumina of varying size in about two-thirds of
the specimens examined, and of more uniform size in about one-third of the specimens, irregularly polygonal
in shape. Grain outlines finely indented.
Holotype. Sample ECO 5, slide JL 187.2; 26.4, 101.4. Plate 91, figs. 17-18. Argiles noires, Ecommoy;
rhotomagense Zone, Middle Cenomanian.
Dimensions. Type material: PD 12 (17-4) 23 ;um (27) s3-4 /^m, ED 8 (13-7) 21 ;um (54) s3-5 ;um, PD/ED 10
(1-4) 1-8 (27), ET 10 (1-4) 2-0 ^m (54), NT 0-3 (0-6) l O^m (54), ST 0-5 (0-7) 10/xm(54), ST/NT 0-8 (1-3)
3-3 (54), LD (least) OT (0-2) 0-5 /xm to (greatest) OT (0-6) 2-0 fxm (54), MW OT (0-2) 0-3 jum (54), LC 8(11-6)
1 8 ,xm (26), LC/PD 0-5 (0-7) 0-9 (26), WC 0-5-2-0 /xm (2), WC/ED < 0- 1 -0- 1 (2), DC 1 -0- 1 -5 /xm (2), DC/ED
01 (2). Other material: Samples JOU 1, JOU 2, and JOU 4, PD 12 (14-6) 20 jum (13), ED 8 (101) 14 ;um
(17); Samples JOU 8, JOU 16, and JOU 17, PD 13-20 (6), ED 10-17 f.nn (8).
Orientation. Type material: P 3-7%, E 50 0%, OA 46-3%.
Occurrence. This study: inflatum Zone, Upper Albian to jukes-brownei Horizon, rhotomagense Zone,
Middle Cenomanian.
Comments. See comments for Retitricolpites ampUfissus, CfB R. georgensis, and
CfC. R. nemejci for distinction from these species. R. sarthensis differs by having a
generally thinner exine, either with unclear exine stratification or with the nexine
rather thinner than the sexine.
Retitricolporites insolitimorus is superficially similar but it is sometimes tetrahedral
in shape. Further, it does not always have three colpi, but has pores, and tends to
have a thicker nexine than sexine, and shorter colpi.
The specimens described as TricolpopoUenites retiformis Pflug and Thomson by
Groot, Penny and Groot (1961) might be the same species, but they are inadequately
described for a close comparison (Thomson and Pflug’s (1953) original description
also is inadequate for comparative purposes). T. haraldii Manum, 1962 is a larger
form with a thinner nexine relative to the sexine. Retitricolpites prosimilis Norris,
1967 differs by having a decrease in lumina diameter towards the poles and a generally
thinner exine. Tricolpites variabilis Burger, 1970 differs in having a generally slightly
EXPLANATION OF PLATE 92
All figures x 2000.
Figs. 1-2. Retitricolpites promiscuus sp. nov. Specimen with lumina of approximately uniform size; equa-
torial aspect; ECO 5, JL 209.1 ; 33.9, 097.7; 1, high focus to show ornament; 2, median focus.
Figs. 3-6. Retitricolpites sarthensis sp. nov. 3-4, holotype; equatorial aspect; ECO 5, JL 187.3; 53.6, 105.9;
3, high focus; 4, median focus. 5-6, oblique aspects; PUN 1, JL 106.3; 40.4, 097.9; 5, median focus;
6, low focus.
Figs. 7-10. Retitricolpites subtilimaculatus sp. nov. 7-8, holotype; equatorial aspect; ECO 5, JL 187.2;
32.1, 103.3; 7, high focus; 8, median focus. 9-10, oblique aspect; ECO 5, JL 209.2; 35.1, 105.7; 9, high
focus; 10, median focus.
Figs. II 14. Retitricolpites sp. 1. 11-12, equatorial aspect; ECO 5, JL 209.2; 52.5, 105.1; 11, median
focus; 12, high focus to show ornament. 13-14, oblique aspect; ECO 5, JL 187.2; 54.5, 111.6; 13, high
focus to show ornament; 14, median focus.
Figs. 15-20. CfC. Striatopollis sarstedtensis Krutzsch. 15-16, oblique aspect; ECO 5, JL 187.2; 51.5,
099.8; 15, high focus; 16, median focus. 17-18, polar aspect; ECO 5, JL 187.2; 25.0, 100.5; 17, median
focus; 18, high focus to show ornament, note the development of the polar microreticulum. 19-20,
equatorial aspect; ECO 5, JL 209.2; 28.1, 1 1 1.8; 19, high focus to show ornament, note the cross pieces
linking the muri; 20, median focus.
PLATE 92
LAING, angiosperm pollen
794
PALAEONTOLOGY, VOLUME 18
thinner exine, the nexine thinner than the sexine, and by having narrow costae
bordering the colpi. T. brnicensis Pacltova, 1971 differs by sometimes being oblate,
by being generally larger, and by always having more or less uniform-sized lumina.
Retitricolpites sarthensis sp. nov.
Plate 92, figs. 3-6
1971 Tricolpites vulgaris (Pierce) Pacltova, p. 113, pi. 3, figs. 6-13.
Description of forty-four specimens from samples ECO 2, ECO 5, ECO 6, and ECO 7. Prolate spheroidal
to prolate; elliptical to circular in equatorial view. Three narrow slit-like (occasionally more gaping) colpi,
margins entire (occasionally slightly ragged). Exine thin, microreticulate, stratification often unclear
(specimens with thicker exines show stratification into unstructured nexine and thicker microreticulate
sexine). Microreticulum perfect; lumina of varying size, irregularly polygonal in shape. Grain outlines
finely indented.
Holotype. Sample ECO 5, slide JL 187.3; 53.6, 105.9. Plate 92, figs. 3-4. Argiles noires, Ecommoy; rhoto-
magense Zone, Middle Cenomanian.
Dimensions. Type material: PD 13 (14-6) 18 /xm (26) sl-7 /xm, ED 7 (11-8) 19 /xm (44) s3-4 ;ixm, PD/ED
IT (1-4) 1-9 (26), ET 0-4 (0-8) 1-2 /xm (44), NT (where determinable) OT (0-2) 0-2 /xm (21), ST (where
determinable) 0-8 (0-9) 10 /xm (21), ST/NT (where determinable) 4-0 (5-8) 9 0 (21), LD (least) OT (0-3)
0-5 /xm to (greatest) 0-4 (TO) 2 0 /xm (44), MW OT (0-2) 0-5 /xm (44), LC 8 (11-2) 16 /xm (26), LC/PD 0-6
(0-8) 0-9 (26). Other material; Sample PUN 1, PD 16 (19-9) 24 /xm (11), ED 10 (14-7) 19 /xm (16); Sample
WOR 1, PD 17-23 /xm (6), ED 9-16 /xm (6).
Orientation. Type material: P 0-0%, E 61-4%, OA 38-6%.
Occurrence. This study: inflatum Zone, Upper Albian to jukes-brownei Horizon, rhotomagense Zone,
Middle Cenomanian. Other occurrence: Pacltova (1971), Cenomanian, Bohemia. Known range', inflatum
Zone, Upper Albian to Cenomanian.
Comments. See comments for CfB. Retitricolpites georgensis, R. promiscuus, and
Retitricolpites sp. 1 for distinction from these species. The form described by Pacltova
(1971) as Tricolpites vulgaris (Pierce) is very similar, but is a little larger (PD 20-
28 /um, ED 18-24 /u.m). R. vulgaris Pierce, 1961 was too briefly described for a close
comparison, but it does seem to be a larger form, perhaps with a thicker exine and
wider colpi. R. minutus Pierce, 1961 also seems to be a similar form. It also was
inadequately described for a close comparison; however, it does seem to differ from
this species by having wider colpi with slightly thickened margins. R. prosimilis
Norris, 1967 differs by usually having a reduced ornament on the apocolpia. Norris
noted, however, that some of his specimens lacked this feature and that these might
represent a separate species; perhaps the latter belong to R. sarthensis. Tricol-
popollenites microreticulatus Takahashi, 1961 is superficially similar; unfortunately,
it was inadequately described for a close comparison.
Retitricolpites subtilimaculatus sp. nov.
Plate 92, figs. 7-10; text-fig. 10
Description of seven specimens from sample ECO 5. Spheroidal to prolate; elliptical to circular in equatorial
view. Three colpi, narrow and slit-like or wide and gaping at the equator, quite deep, margins entire
(occasionally slightly ragged), sometimes rather indistinct. Clear exine stratification into unstructured
nexine and microreticulate sexine; sexine thicker than nexine. Microreticulum perfect; lumina of about
uniform size and equidimensionally shaped. Grain outlines finely indented.
LAING. MID-CRETACEOUS ANGIOSPERM POLLEN
795
Holotype. Sample ECO 5, slide JL 187.2; 32.1, 103.3. Plate 92, figs. 7-8. Argiles noires, Ecommoy; rhoto-
magense Zone, Middle Cenomanian.
Dimensions. PD 11-12 jiim (2), ED 8-12 fxm (7), PD/ED 10-1-4 (2), ET 10-1-7 nm (7), NT 0-2-0-8 (um (7),
ST 0-7-0-9 (ixm (7), ST/NT M-4-0 (7), LD 0-1 -0-2 /urn (7), MW 0-1 ^^m (7), LC 7-10 /^m (2), LC/PD 0-6-
0-8(2).
Orientation. P 0-0%, E 28-6%, OA 71-4%.
Occurrence. This study: inflatum Zone, Upper Albian to rliotomagense Zone (? costatus Horizon), Middle
Cenomanian.
Comments. See comments for Retitricolpites exiguiexemplum and R. meumendum for
distinction from these species. Tricolpopollenites mimitus Brenner, 1963 is a very
similar form which perhaps differs in having a generally slightly thinner exine and
possibly also in having smoother or less indented grain outlines. Tricolpites albiensis
Kemp, 1968 differs by being somewhat larger and by having much more closely-
spaced sexinal sculptural elements, such that the grain outlines are more or less
smooth.
Retitricolpites sp. 1
Plate 92, figs. 11-14; text-fig. 12
Description of seven specimens from sample ECO 5. Prolate spheroidal to prolate; sub-circular to elliptical
in equatorial view. Three colpi, usually narrow and slit-like but occasionally wide and gaping at the equator,
margins more commonly entire than ragged and sometimes a little thickened, sometimes indistinct. Clear
exine stratification into unstructured nexine and a sexine composed of bacula which support a micro-
reticulum; nexine usually thinner than sexine (very rarely nexine and sexine of about equal thickness).
Microreticulum usually imperfect (often with some discrete bacula) but sometimes perfect; lumina of
varying size, irregularly polygonal in shape. Grain outlines indented.
Dimensions. Sample ECO 5: PD 15-17 ;um (3), ED 11-20 ^^m (7), PD/ED M-1-4 (3), ET 10-1-5 i^m (7),
NT 0-2-0-5 /xm (7), ST 0-5-10 jum (7), ST/NT 1-0-4-5 (7), LD (least) 0-2-0-5 /xm to (greatest) T5-2-5 /xm
(7), MW 01-0-2 ;txm (7), LC 7-13 ^xrn (3), LC/PD 0-5-0-8 (3). Samples JOU 1 and JOU 4; PD 16-23 ^tm
(4), ED 12-17 nxm (6).
Orientation. Sample ECO 5: P 0-0%, E 42-9%, OA 57-1%.
Occurrence. This study: inflatum Zone, Upper Albian to costatus Horizon, rhotomagense Zone, Middle
Cenomanian.
Comments. Since only a few specimens of this type have been found from any one
locality, and of these some are not well preserved, I have not attempted to erect a new
species to accommodate them. Retitricolpites sarthensis differs by having a constantly
perfect, rather finer-meshed microreticulum. CfB. R. georgensis differs by having
a microreticulum composed of solid muri which are in continuous contact with the
nexine thus giving a different LO pattern (see text-fig. 12). The microreticulum of this
species is also constantly perfect. Tricolpopollenites platyreticulatus Groot, Penny
and Groot, 1961 seems to differ by having a constantly perfect microreticulum which
(judging from the figures) continues down to the nexine rather than being supported
on bacula. T. macroreticulatus Groot and Groot, 1962 seems to differ by being rather
larger, by having the colpi bordered by a margo, and an apparently constantly perfect
microreticulum which is in continuous contact with the nexine. T. virgeus Groot,
Penny and Groot, 1961 seems to differ in having the microreticulum made up of.
796
PALAEONTOLOGY, VOLUME 18
LO PATTERN
CROSS SECTION OF
EQUATORIAL EXINE
TEXT-FIG. 12. LO patterns and cross-sections of
equatorial exine of Retitricolpites sp. 1 {left) and
CfB. R. georgensis Brenner {right).
rather than supported by, bacula. Tricolpites sp. 1 of Kemp (1968) differs by having
a perfect microreticulum which tends to detach from the nexine. R. marginatus
van Hoeken-Klinkenberg, 1966 differs by being larger, by having a rather coarser-
meshed microreticulum which is reduced along the colpus margins, and by having
a thicker exine.
Genus striatopollis Krutzsch, 1959
1959 Striatopollis Krutzsch, p. 142.
1962 Striopollis Rouse, p. 212.
Type species. Striatopollis sarstedtensis Krutzsch, 1959, p. 143, pi. 34, figs. 1-24; text-fig. 12.
CfC. Striatopollis sarstedtensis K.r\xX.z^ch., 1959
Plate 92, figs. 15-20; Plate 93, fig. 1
1959 Striatopollis sarstedtensis Krutzsch, p. 143, pi. 34, figs. 1-24; text-fig. 12.
EXPLANATION OF PLATE 93
Fig. 1. CfC. Striatopollis sarstedtensis Krutzsch. Scanning electron micrograph; near equatorial aspect;
ECO 5, stub JL 8; 335714; x 5000.
Figs. 2-5. CfA. Liliacidites peroreticulatus {Brenner). 2-3, distal polar aspect; ECO 4, JL 184. 1 ; 32.0, 1 10.6;
2, high focus to show ornament ; 3, median focus ; x 2000. 4-5, scanning electron micrographs ; oblique
aspect; ECO 5, stub JL 8; 224803; 4, x2000; 5, x 5000.
Figs. 6-7. Psilatricolporites complanatius sp. nov. 6, holotype; syncolpate specimen; polar aspect; ECO 5,
JL 209.1 ; 30.2, 101.6; median focus; x2000. 7, specimen with small apocolpia; polar aspect; ECO 5,
JL 209.2; 38.9, 103.1 ; median focus; x2000.
Eigs. 8-13. Retitricolporites insolitimorus sp. nov. 8-9, holotype; ellipsoidal specimen with two colpi (one
apparently with no pore) and two pores (one apparently not associated with a colpus); equatorial aspect;
ECO 5, JL 209.2; 23.7, 105.0; 8, high focus to show ornament and pore; 9, median focus; x2000.
10-1 1, ellipsoidal specimen with three pores and three colpi; polar aspect; PUN 1,JL 106.3;39.3, 103.0;
10, high focus to show ornament; 11, median focus; x2000. 12-13, tetrahedral specimen with three
colpi and three pores; ECO 1,JL 179.1; 35.2; 110.4; 12, median focus; 13, low focus; x2000.
Figs. 14-15. Retitricolporites ecommoyensis sp. nov. Holotype; equatorial aspect; ECO 5, JL 209.2; 32.4,
106.9; 14, high focus; 15, median focus; x2000.
PLATE 93
LAING, angiosperm pollen
798
PALAEONTOLOGY, VOLUME 18
Description of five specimens from sample ECO 5. Spheroidal to prolate; triangular trilobate outline
in polar view, sub-circular to elliptical in equatorial view. Three slit-like colpi, margins entire, angul-
aperturate, sometimes syncolpate. Clear exine stratification into unstructured nexine and striate or
striato-microreticulate sexine; sexine thicker than nexine. Muri and striae tend to parallel the polar
axis, sometimes anastomosing and occasionally (especially near the poles) linked by cross pieces to form
a microreticulum; S.E.M. observation shows the muri to bear small cones.
Dimensions. PD 15-24 ;^m (3), ED 10-23 nm (5), PD/ED 1-4-T6 (3), ET lT-1-6 /xm (5), NT 0-2-0-6 /xm
(5), ST 0-7-1 0 /xm (5), ST/NT 1 -4-4-5 (5), LD 0-3 /xm (1), MW 0-2-0-5 /xm (5), W striae 0-2-0-5 /xm (5),
number of muri between each pair of colpi 7-19 (4), LC 11-17 /xm (2), LC/PD 0-7-0-8 (2), WC 0-2 /xm (1),
WC/ED<0-1 (1), DC l-0/xm(l), DC/ED<0-1 (1).
Orientation. P 20-0%, E 60-0%, OA 20-0%.
Occurrence. This study: inflatum Zone, Upper Albian to costatus Horizon, rhotomagense Zone, Middle
Cenomanian. Other occurrences: Krutzsch (1959), Lower Paleocene, G.D.R.; Groot and Groot (1962),
Cenomanian, Portugal. Known range', inflatum Zone, Upper Albian to Lower Paleocene.
Comments. None of the specimens described by Krutzsch (1959) seems to show the
development of a polar microreticulum.
StriatopoUis cf. paraneus (Norris) in Dettmann (1973) differs by having ‘ropy’
muri (under S.E.M. observation).
Subturma ptychotriporines Naumova emend. Potonie, 1960
As with the tricolpates, the genera of van der Hammen (1956) are used as they are
the most convenient for light microscope studies.
Genus psilatricolporites van der Hammen ex van der Hammen and Wymstra, 1964
1956 Psilatricolporites van der Hammen, p. 91.
1964 Psilatricolporites van der Hammen ex van der Hammen and Wymstra, p. 236.
Type species. Psilatricolporites operculatus van der Hammen and Wymstra, 1964, p. 236, pi. 1, fig. 13.
Psilatricolporites complanatius sp. nov.
Plate 93, figs. 6-7
Description of nine specimens from samples ECO 5 and ECO 7. Strongly oblate; triangular or rounded
triangular, usually quite shallowly trilobate outline in polar view. Three distinct long gaping (less commonly
more slit-like) colpi, margins entire but usually faint, angul-aperturate, sometimes syncolpate; single
equatorial pore in each colpus, often rather unclear. No clear exine stratification. Psilate.
Holotype. Sample ECO 5, slide JL 209.1; 30.2, 101.6. Plate 93, fig. 6. Argiles noires, Ecommoy; rhoto-
magense Zone, Middle Cenomanian.
Dimensions. PD c. 1 - 5 /xm (1), ED 13-15 /xm (9), PD/ED c. 0-1 (1), ET 0-4-0-8 /xm (9), WC 1-5 (2-7) 6-0 /xm
(9), WC/ED 0-1 (0-2) 0-4 (9), DC 0-0 (0-6) 1-5 /xm (9), DC/ED 0-0 (< 0-1) 0-1 (9), pore diameter 1-0 (2-4)
4-5 /xm (8).
Orientation. P 100-0%.
Occurrence. This study: carcitanensis Horizon, mantelli Zone, Lower Cenomanian to costatus Horizon,
rhotomagense Zone, Middle Cenomanian.
Comments. Tricolporopollenites orhiculatiisGroo\.,Ptnv\y dndGxooi, 1961 differs from
this species by being slightly prolate. Tricolporopollenites sp. S. Cl. 215 of Jardine
LAING: MID-CRETACEOUS ANGIOSPERM POLLEN
799
and Magloire (1965) has a more sub-spherical shape. TricoIporopoUenites sp. S. Cl.
141 of Jardine and Magloire (1965) is larger, has pores with annuli and perhaps has
shorter colpi. T. aliquantulus Hedlund, 1966 has a more prolate shape. TricoIporo-
poUenites sp. B of Brenner (1968) has much shorter colpi. Nyssapollenites albertensis
Singh, 1971 differs by having rim-like thickenings around the pores, and by having
much shorter colpi.
Genus retitricolporites van der Hammen ex van der Hammen and Wymstra, 1964
1956 Retitricolporites van der Hammen, p. 93.
1964 Retitricolporites van der Hammen ex van der Hammen and Wymstra, p. 235.
Type species. Retitricolporites guianaensis van der Hammen and Wymstra, 1964, p. 235, pi. 3, figs. 1-2.
Retitricolporites ecommoyensis sp. nov.
Plate 93, figs. 14-15
Description of nine specimens from sample ECO 5. Prolate spheroidal to prolate; sub-circular to elliptical
in equatorial view. Three distinct narrow slit-like colpi, margins entire and with nexinal thickening; single
distinct equatorial pore in each colpus; many specimens have the colpi buckled out at the equator (i.e. they
are tricolporoidate in the sense of Doyle 1969). Clear exine stratification into unstructured nexine and
microreticulate sexine (sexine occasionally psilate at the poles); sexine generally thicker than nexine (except
at colpus margins and sometimes at the poles), occasionally exine (particularly nexine) thickens at the poles.
Microreticulum perfect; lumina of about uniform size and equidimensionally shaped. Grain outlines finely
indented to smooth.
Holotype. Sample ECO 5, slide JL 209.2; 32.4, 106.9. Plate 93, figs. 14-15. Argiles noires, Ecommoy;
rliotomagense Zone, Middle Cenomanian.
Dimensions. PD 10-16 /xm (8), ED 6-13 /um (9), PD/ED lT-1-7 (8), equatorial ET 0-5- 1-2 ;um (9), equatorial
NT OT-0-4 nm (9), equatorial ST 0-4-0-8 /xm (9), equatorial ST/NT 1-5-4 0 (9), LD <0T-0-3 jum (9),
MW <0T-0-2 ij.m (9), LC 7 (9-4) 13 ;um (8), LC/PD 0-6 (0-7) 0-8 (8), thickness of colpus margins 0-5-
10 jixm (9), pore diameter 0-2 (0-7) 1-2 jum (9).
Orientation. P 0-0%, E 88-9%, OA 1 11%.
Occurrence. This study: inflatum Zone, Upper Albian to jukes-brownei Horizon, rliotomagense Zone,
Middle Cenomanian.
Comments. TricoIporopoUenites inaequalis Groot, Penny and Groot, 1961 is a larger
form which seems to have unthickened colpus margins. TricoIporopoUenites sp. S.
Cl. 428 of Jardine and Magloire (1965) differs by its larger size and by the fact that
it often occurs in tetrads. Tricolporoidites minimus Pacltova, 1971 has a rather thicker
exine and a granulate nexine. T. subtilis Pacltova, 1971 differs in having a chagrenate
exine.
Retitricolporites insolitimorus sp. nov.
Plate 93, figs. 8-13
Description of thirty-nine specimens from samples ECO 1, ECO 2, ECO 4, ECO 5, ECO 6, and ECO 7.
Variable in shape, either ellipsoidal (oblate spheroidal to sub-prolate) or a more tetrahedral shape ; ellipsoidal
grains with triangular quite deeply trilobate outline in polar view, elliptical in equatorial view; tetrahedral
grains with a more triangular outline. Tricolporate development often aberrant with up to three colpi which
may or may not have pores, or up to three pores, some of which may not be associated with colpi (the
800
PALAEONTOLOGY, VOLUME 18
apparent lack of a full complement of pores and/or colpi in some specimens may simply be caused by the
aspect of these specimens). Ellipsoidal grains fossaperturate. Pores sometimes elongate parallel to the polar
axis but more commonly approximately circular; colpi relatively short and deep. Clear exine stratification
into unstructured nexine and microreticulate sexine; nexine most commonly thicker than sexine, less
commonly nexine and sexine of about equal thickness, only rarely sexine thicker than nexine. Micro-
reticulum perfect; lumina either of about uniform size and regularly polygonal shape, or of more varying
size and irregularly polygonal shape. Grain outlines finely indented to almost smooth.
Holotype. Sample ECO 5, slide JL 209.2; 23.7, 105.0. Plate 93, figs. 8-9. Argiles noires, Ecommoy; rhoto-
magense Zone, Middle Cenomanian.
Dimensions. Type material: PD 13 (17-8) 22 /im (15), ED 9 (15-7) 22 ;um (39) s3-7 fj.m, PD/ED 0-8 (IT)
1 -2 ( 1 5), ET 1 -0 ( 1 -5) 2-5 ^m (39), NT 04 (0-9) 1 -5 ^^m (39), ST 0 4 (0-6) 1 0 ,xm (39), ST/NT 0-3 (0-7) 2-3 (39),
ED (when uniform) OT (0-2) 04 fxm (26), LD (when varying) (least) OT (0-2) 0-2 ^m to (greatest) 0-3 (0-9)
2-0 ^m (13), MW OT (0-2) 0-3 i^m (39), LC 3 (10-0) 15 ;um (15), LC/PD 0-2 (0-5) 0-8 (15), WC 2-0-5 0 fxm
(2), WC/ED 01-0-3 (2), DC 1-5-3 0 /xm (2), DC/ED 01-0-2 (2), pore diameter (including W of elongate
pores) 0-8 (2-3) 60 fxm (39), L of elongate pores 30-7-5 jum (6). Other material: Sample JOU 15, PD 15-
19 ^m (5), ED 12-17 ^m (5).
Orientation. Type material: P 5-1%, E 410%, OA 53-8%.
Occurrence. This study: inflatiim Zone, Upper Albian to jukes-brownei Horizon, rhotomagense Zone,
Middle Cenomanian.
Comments. See comments for Retitricolpites promiscuus for distinction from this
species.
Tricolporopollenites microreticulatus Thomson and Pflug, 1953 differs by being
on average rather larger and in not showing an aberrant tricolporate development
and tetrahedral condition. T. inaequalis Groot, Penny and Groot, 1961 differs by
having the nexine thinner than the sexine and in not showing an aberrant tricolporate
development and tetrahedral condition. T. subobscurus Groot and Penny, 1960,
Retitricolporites crassicostatus van der Hammen and Wymstra, 1964,. and Tricol-
poroidites bohemicus Pacltova, 1971 all differ in not showing an aberrant tricolporate
development and tetrahedral condition.
Turma poroses Naumova emend. Potonie, 1960
Subturma triporines Naumova emend. Potonie 1960
Genus complexiopollis Krutzsch emend. Goczan, Groot, Krutzsch
and Pacltova, 1967
1959 Cowp/ex/opo//;5 Krutzsch, p. 134.
1967 Complexiopollis Krutzsch emend. Goczan et al., p. 453.
Type species. Complexiopollis praeatumescens Krutzsch, 1959, p. 135, pi. 31, figs. 39-54; text-fig. 6.
CfB. Complexiopollis subtilis (Krutzsch) Goczan, Groot, Krutzsch and
Pacltova, 1967
Plate 94, figs. 1-3; text-fig. 13
1959 LatipoUis subtilis Krutzsch, p. 129, pi. 31, figs. 1-13; text-fig. 1.
1967 Complexiopollis subtilis (Krutzsch) Goczan et a!., p. 445.
Description of three specimens from sample WOR 1. Sub-oblate; elliptical in equatorial view, strongly
LAING: MID-CRETACEOUS ANGIOSPERM POLLEN
801
TEXT-FIG. 13. Pore structure of CfB. text-fig. 14. Pore structure of Triporopollenites
Complexiopollis subtilis (Krutzsch). curtisi sp. nov.
three-rayed shape in polar view. Three pores, often unclear, seemingly with faint vestibula (see text-fig. 13),
angul-aperturate. Exine stratified into unstructured nexine and a sexine with a rather rough surface; nexine
and sexine of about equal thickness. Grain outlines rather irregular.
Dimensions. PD 13-14 (2), ED 15-17 /xm (3), PD/ED 0-8-0-9 (2), ET 10- 1-2 fxm (3), NT 0-5-0-6 fivn
(3), ST 0-5-0-6 /xm (3), ST/NT 10 (3), exopore diameter 0-5- 1-2 /xm (3), exopore diameter/ED 0 03-0 07 (3),
endopore diameter 0-5- 1-2 /xm (3), endopore diameter/ED 0 03-0 07 (3), vestibulum diameter L5-3 0 /xm
(3), vestibulum diameter/ED 0T0-0T7 (3), D of pores 2 0-2T /xm (3), D of pores/ED 0T17-0T33 (3).
Orientation. P 0-0%, E 66-7%, OA 33-3%.
Occurrence. This study : jukes-brownei Horizon, rhotomagense Zone, Middle Cenomanian. Other occurrence :
Krutzsch ( 1959), ?Upper Cenomanian and Turonian, Central Europe. Known range: jukes-brownei Horizon,
rhotomagense Zone, Middle Cenomanian to Turonian.
Comments. Though the pore structure of the specimens described above is not clear,
it does not appear to be as complicated as that described by Krutzsch (1959).
Genus triporopollenites Thomson and Pflug, 1953
1953 Triporopollenites Thomson and Pflug, p. 82.
Type species. Triporopollenites coryloides Thomson and Pflug, 1953, p. 84, pi. 9, figs. 20-24.
Triporopollenites curtisi sp. nov.
Plate 94, figs. 4-7; text-fig. 14
Description of six speeimens from sample WOR 1. Oblate: three-rayed or concave-sided triangular outline
in polar view. Three, usually distinct, simple pores; exine sometimes a little thickened around the pores,
and often a little constricted behind the endopore such that an arrow-head shaped or triangular cavity
appears to be present behind the endopore (see text-fig. 14), angul-aperturate. Exine stratified into un-
structured nexine and a sexine with a rather rough surface; nexine most commonly thinner than sexine,
occasionally nexine and sexine of about equal thickness; exine stratification often rather unclear. Grain
outlines rather irregular.
Holotype. Sample WOR 1, slide JL 236.1; 41.8, 100.5. Plate 94, figs. 4-5. Glauconitic Marl, Worbarrow
Bay; probable jukes-brownei Horizon, rhotomagense Zone, Middle Cenomanian.
Dimensions. ED 14-20 /xm (6), ET 1 0- 1 -2 /xm (6), NT 0-2-0-6 /xm (6), ST 0-6-0-8 /xm (6), ST/NT 1 0- 1 -4 (6),
exopore diameter 0-2 (0-7) IT /xm (6), exopore diameter/ED 0 01 (0 03) 0 07 (6), endopore diameter 0-2
(0-7) 10 /xm (6), endopore diameter/ED 0 01 (0-03) 0-07 (6), D of pore 10 (1-6) 2 0 /xm (6), D of pore/ED
0 052 (0 084) OTll (6).
802
PALAEONTOLOGY, VOLUME 18
Orientation. P 50 0%, OA 50 0%.
Occurrence. This study: jukes-brownei Horizon, rhotomagense Zone, Middle Cenomanian.
Comments. ComplexiopoUis praeatumescens Krutzsch, 1959 differs in having more
complex pores, more exine layers, a finely punctate to reticulate sculpture, and by
being larger. Latipollis vulgaris Groot and Groot, 1962 has a more circular cavity
behind the endopore (see text-fig. 2 of Groot and Groot). TuronipoUis helmigii van
Amerom, 1965 differs in having small atria and a granulate to reticulate sculpture.
Conclavipollis densilatus Kimyai, 1 966 is a larger form with a thicker exine and perhaps
wider pores. Triporopollenites pseudocanalis Paden Phillips and Felix, 1971 is a larger
form which has annuli around the pores and a more convex- (or at least not so strongly
concave-) sided shape with protruding apices.
Triporopollenites worbarrowensis sp. nov.
Plate 94, figs. 8-1 1
Description of twelve specimens from sample WOR 1. Oblate; straight or slightly convex-sided triangular
outline in polar view, equatorial apices slightly protruding. Three, usually distinct, simple pores; when
damaged the pores occasionally have the appearance of short slightly gaping colpi with a V-shaped cross-
section, angul-aperturate. Exine stratified into unstructured nexine and a sexine with a smooth or slightly
rough surface; sexine slightly thicker than, or of equal thickness to, nexine (occasionally nexine slightly
thicker than sexine) ; exine stratification occasionally unclear. Grain outlines smooth or slightly irregular.
Holotype. Sample WOR 1, slide JL 236.1 ; 26.0, 096.4. Plate 94, figs. 8-9. Glauconitic Marl, Worbarrow
Bay; probable jukes-brownei Horizon, rhotomagense Zone, Middle Cenomanian.
Dimensions. ED 16(18-4) 23 p.m (12), ET 0-7 (1-0) 1-3 ^m (12), NT 0-3 (0-4) 0-6 (12), ST 0-4 (0-6) 1-0 ^lm
(12), ST/NT 0-7 (1-5) 3-3 (12), exopore diameter 0-5 (1-2) 2-2 ;um (11), exopore diameter/ED 0-02 (0-05)
0-10 (11), endopore diameter 0-5 (10) 2 0 /um (11), endopore diameter/ED 0 02 (0 04) 0 08 (11), D of pore
1 -0 ( 1 -3) 2 0 ^m ( 1 1 ), D of pore/ED 0-052 (0-068) 0-111 (11).
Orientation. P 50-0%, OA 50 0%.
Occurrence. This study: jukes-brownei Horizon, rhotomagense Zone, Middle Cenomanian.
Comments. Triorites africanensis Jardine and Magloire, 1965 differs in being rather
larger.
EXPLANATION OF PLATE 94
All figures x 2000.
Figs. 1-3. CfB. ComplexiopoUis subtilis (Krutzsch). 1-2, equatorial aspect; WOR 1, JL 236.1 ; 54.0, 095.9;
1, median focus; 2, low focus. 3, oblique aspect; WOR 1, JL 236.2; 33.1, 098.2; median focus.
Figs. 4-7. Triporopollenites curtisi sp. nov. 4-5, holotype; polar aspect; WOR 1, JL 236.1; 41.8; 100.5;
4, high focus; 5, median focus. 6-7, oblique aspect; WOR 1, JL 236.2; 51.6, 106.5; 6, high focus; 7, median
focus.
Figs. 8-11. Triporopollenites worbarrowensis sp. nov. 8-9, holotype; polar aspect; WOR 1, JL 236.1 ; 26.0,
096.4; 8, high focus; 9, median focus. 10-1 1, oblique aspect; WOR 1, JL 238.1 ; 27.2, 1 1 1.2; 10, median
focus; 1 1, low focus.
Figs. 12-14. CfB. Asteropollis asteroides Hedlund and Norris. 12-13, proximal polar aspect; ECO 5, JL
209.3; 32.6, 103.1; 12, high focus to show ornament; 13, median focus to show sulcus. 14, equatorial
aspect; ECO 5, JL 209.3; 32.6, 101.9; high focus to show ornament.
PLATE 94
LAING, angiosperm pollen
804
PALAEONTOLOGY, VOLUME 18
SEQUENCE OF POLLEN ASSEMBLAGES
Using the results of this study and the results of earlier workers (Couper 1958 ; Hughes
1958; Kemp 1968, 1970), it is possible to suggest a sequence of angiosperm pollen
assemblages. Unfortunately, the data is as yet insufficient for the establishment of
a zonal scheme, but I hope that this sequence will help others to make ‘rule-of-thumb’
stratigraphic assessments on material coming from this area. In another paper
(Laing 1975), I have described a scheme which is, I hope, applicable over a wider
area (perhaps over Europe and North America).
The approximate level of the earliest occurrence of each assemblage has been
referred to the ammonite zonations of Spath (1923-1943) for the Albian and of
Kennedy (1969) for the Cenomanian. The sequence of pollen assemblages and their
approximate correlation with the ammonite zones are outlined on Table 2.
Liliacidites hughesii Assemblage.
Base. Within the Upper Barremian.
Diagnostic features. This assemblage is characterized by having reticulate and/or
clavate monosulcate forms as the only angiospermous pollen present. In Britain
and France there is only one angiospermous species at present known from this
assemblage and that is Liliacidites (al. ClavatipoUenites) hughesii (Couper 1958
emend. Kemp 1968) comb. nov. The base of the occurrence of this assemblage is
thus defined by the first appearance of this species which occurs in the Upper Bar-
remian (Hughes 1958).
Liliacidites rotundas- Retitricolpites amplifissus Assemblage
Base. Within the Leymeriella tar defur cat a Zone (Lower Albian).
Diagnostic features. This assemblage is characterized by the presence of monosulcate
and tricolpate forms; tricolporate forms may also be present but triporate forms are
absent. Although Retitricolpites sarthensis may be present it is never the dominant
angiosperm species (neither, for that matter, is it present in any abundance). In this
area, the base of the occurrence of this assemblage may be defined by the first appear-
ance of Liliacidites rotundas and R. amplifissus, which according to Kemp (1968,
1970) is in her sample F270. This is probably from the tardefurcata Zone of the Lower
Albian. At this level, the only angiospermous species present are these two species
and L. hughesii.
I have only studied the Upper Albian and Lower Cenomanian part of the time
interval represented by this assemblage, and the evidence both from my work and
that of Kemp (1968) is that L. rotundas, R. amplifissus, R. promiscuus, Retitricolpites
sp. 1 and/or Tricolpites albiensisK.Qmp, 1968 are the dominant angiospermous species
in the Upper Albian, and that Retitricolporites insolitimorus and/or Retitricolpites
amplifissus may be the most abundant angiosperm species in the Lower Cenomanian.
The time interval occupied by this assemblage is important in that it is the period
during which reticulate, striate and psilate tricolpate, reticulate tri- to hexachotomo-
sulcate (e.g. Asteropollis), and reticulate and psilate tricolporate forms first appear
(for a fuller discussion of this see Laing 1975).
LAING: MID-CRETACEOUS ANGIOSPERM POLLEN
805
TABLE 2. Sequence of angiosperm pollen assemblages and their approximate
correlation with the ammonite zonation of the mid-Cretaceous of southern
Britain and northern France. (Ammonite zonation after Spath 1923-1943;
and Kennedy 1969.)
STAGE
AMMONITE
ZONE
ASSEMBLAGE OR
FAUNAL HORIZON
POLLEN
ASSEMBLAGE
Acanthoceras
jukes-brownei
T riporopolleni tes
worbarrowensis
MIDDLE
CENOMANIAN
Acanthoceras
rhotomagense
Turrilites
acutus
9
Retitricolpites
sarthensis
Turrilites
costatus
(uncharacterized)
9
LOWER
CENOMANIAN
Mantelliceras
mantelli
Mantelliceras
gr. dixoni
Mantelliceras
saxbii
Hypoturrilites
carcitanensis
UPPER
Stoliczkaia
dispar
Liliacidites
ALBIAN
Mortoniceras
inflatum
rotundus-
Retitricolpites
amplifissus
MIDDLE
Euhoplites
lautus
ALBIAN
Hoplites
dentatus
LOWER
Douvilleiceras
mammillatum
ALBIAN
Leymeriella
tardefurcata
9
APTIAN
Liliacidites
hughesii
BARREMIAN
9
806
PALAEONTOLOGY, VOLUME 18
Retitricolpites sarthensis Assemblage
Base. Between the Hypoturrilites carcitanensis Assemblage horizon of the Man-
telliceras mantelli Zone (Lower Cenomanian) and the Turrilites costatus Assemblage
horizon of the Acanthoceras rhotomagense Zone (Middle Cenomanian).
Diagnostic features. This assemblage is characterized by the presence of either
Retitricolpites sarthensis or R. promiscuus as the most abundant angiosperm species.
When R. promiscuus is the most abundant species, then R. sarthensis is the next most
abundant. R. sarthensis is always more abundant than R. amplifissus. Triporate
grains are absent. R. crassitransennus makes its first appearance during the time
interval represented by this assemblage and, apart from the possibility of the presence
of this species, and the relative abundance of R. sarthensis, this assemblage is quite
like the later (i.e. Lower Cenomanian) part of the L. rotundus-R. amplifissus
Assemblage.
This assemblage occurs both in the base of the Glauconitic Marl at Punfield Cove
and in the Argiles noires of Ecommoy.
Triporopollenites worbarrowensis Assemblage
Base. Between the Turrilites costatus and Acanthoceras jukes-brownei Assemblage
horizons of the A. rhotomagense Zone (Middle Cenomanian).
Diagnostic features. The base of the occurrence of this assemblage is defined by the
first appearance of triporate pollen. Three triporate species occur in the material
which I have examined, Triporopollenites worbarrowensis, T. curtisi, and Com-
plexiopollis subtilis, the first mentioned being the most abundant angiosperm species
present.
This assemblage occurs in the base of the Glauconitic Marl at Worbarrow Bay.
Although it would also be expected to occur in the base of the Glauconitic Marl at
Lulworth Cove, I have found no triporate grains in this material. Indeed, this material
is peculiar in being generally depleted in angiosperm pollen, such pollen being about
six times as abundant (by comparison with the total spore/pollen content) in the
material from Worbarrow Bay as it is in the material from Lulworth Cove.
Acknowledgements. This research was carried out whilst I was in receipt of a N.E.R.C. training award.
I wish to thank Mr. N. F. Hughes for his encouragement and advice, Messrs. D. Marriage and D. Bursill
for their assistance with the photographic work, and Mr. R. S. Curtis for his assistance in the preparation
of certain samples.
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Original typescript received 18 January 1975
Revised typescript received 28 April 1975
J. F. LAING
Department of Geology
Sedgwick Museum
Downing Street
Cambridge
RADNORIA, A NEW SILURIAN
PROETACEAN TRILOBITE, AND THE ORIGINS
OF THE BRACHYMETOPIDAE
by R. M. OWENS and a. t. thomas
Abstract. The new trilobite genus Radnoria is proposed to include the type, R. syrphetodes sp. nov., and two other
species, R. triquetra sp. nov. and R. humUlima (Barrande, 1852), from the Silurian of Britain and Czechoslovakia.
Its morphology includes features typical of both the Brachymetopidae and Warburgellinae, suggesting a phyletic
link between the two groups. The composition of the Brachymetopidae is discussed, and new family and subfamily
diagnoses are given.
Recent collecting from the Dolyhir Limestone (Silurian, Wenlock Series) in the
Old Radnor district, Powys (Radnorshire), the Much Wenlock Limestone Formation
of Wren’s Nest Hill, Dudley, West Midlands, and from a limestone of Wenlock age
near Llandeilo, Dyfed (Carmarthenshire) has furnished large numbers of dissociated
exoskeletal parts of two undescribed proetacean trilobite species, here placed in
a new genus. Cyphaspis humillima Barrande, 1 852 from the late Wenlock of the Prague
district, Czechoslovakia also belongs to the same genus. The significance of this new
genus lies in its similarities both to members of the proetid subfamily Warburgellinae
and to the family Brachymetopidae, and in the consequent implications for the
origins of the latter.
Terminology. Terms used in the descriptions are those dehned by Harrington et al. (in Moore 1959,
pp. 0117-0126) and Owens (1973, pp. 3, 5; text-fig. 1a, p. 4).
Repositories. The following abbreviations are used herein: GSM— The Geological Museum, Institute of
Geological Sciences, London; NMW— National Museum of Wales, Cardiff; NMP— National Museum,
Prague.
SYSTEMATIC PALAEONTOLOGY
Family brachymetopidae Prantl and Pfibyl, 1951
Diagnosis. The following diagnosis is based on that of Whittington (1960, p. 407),
modified to include the Warburgellinae and other slight amendments. Cephalon with
preglabellar field; tropidium or tropidial ridges may be present; glabella narrows
forwards, commonly with well defined Ip lobe; 2p and 3p furrows may be present;
palpebral lobe far back and close to glabella; anterior branches of facial sutures
divergent; connective sutures diverge backwards; thorax of 8-10 segments; no
preannulus; pygidium relatively large; axis with 76-14 rings, pleural ribs with flat-
topped profile, or with posterior band elevated above anterior; pygidial margin
entire or with short spines; external surface smooth, granular, tuberculate, rugulose,
pitted, or spinose, or combination of these.
[Palaeontology, Vol. 18. Part 4, 1975, pp. 809-822, pis. 95-96.]
K
810
PALAEONTOLOGY, VOLUME 18
Stratigraphical range. Silurian (Llandovery) to uppermost Carboniferous, possibly
to Permian.
Subfamily warburgellinae Owens, 1973
[= V^arburgellinae Yolkin, 1974]
Diagnosis. Ip furrow deep; tropidium or tropidial ridges may be present; occipital
ring with lateral lobes; thorax of 8-10 segments; pygidium with narrow axis with
?6-14 rings; pleural areas with 15-1 pairs of ribs with flat-topped profile; pygidial
border may be present ; sculpture granular or rugulose, or exoskeleton smooth.
Genera and subgenera. Warburgella {Warburgella) Reed, 1931 ; W. (Tetinia) Chlupac,
1971 ; Prantlia Pfibyl, 1946; Tropidocare Chlupac, 1971 ; IKoneprusites Pfibyl, 1964.
Stratigraphical range. Silurian (Llandovery) to Devonian (Gedinnian), possibly to
Middle Devonian.
Yolkin (1974, p. 64) included Warburgella, AstroproetusBegg, 1939, Tetinia,
Cyphoproetus Kegel, 1927, and Paleodechenella Maximova, 1970 in the Warburgel-
linae. Of these genera, Owens (1973, p. 8) placed Cyphoproetus in the Proetinae and
Astroproetus in the Tropidocoryphinae (ibid., p. 40), and reasons for doing so are
discussed therein. Without having seen original material of Paleodechenella, we cannot
comment on its subfamilial affinities.
Yolkin (1974, p. 64) referred the Warburgellinae to the Dechenellidae. Members
of this family (which we consider to be a proetid subfamily) do show a broad
resemblance to warburgellines, but possess a preannulus, and the pygidial pleural
rib structure is like that of the Proetinae. On the basis of the thoracic and pygidial
differences, and also because dechenellines are almost certainly phyletically linked
with proetines (Owens 1973, p. 84), we consider that their general similarity to
warburgellines is due to homoeomorphy.
Subfamily brachymetopinae Prantl and Pfibyl, 1951
Diagnosis. Preglabellar field broad, concave or weakly convex in longitudinal section ;
Ip lobe isolated except in Brachymetopus', 2p and 3p commonly absent or ill defined;
anterior branches of facial sutures widely divergent (ankylosed in Brachymetopus) ',
tropidium absent; occipital ring without lateral lobes; rostral plate large, may
extend as far back as genal angle; thorax of nine segments; pygidial axis with 10-13
axial rings; pleural ribs commonly with posterior band elevated above anterior, but
EXPLANATION OF PLATE 95
Figs. 1-6. Radnoria syrphetodes gen. et sp. nov. \a-e. Limestone of probable Wenlock age, old quarry
opposite Ty-newydd Farm, 1-3 km at 127° from Llanarthney church, Dyfed (SN 5442 1951): GSM
103744, cranidium, anterolateral oblique, dorsal, posterolateral oblique, anterior, and left lateral views,
X 8. 2-6, Wenlock Series, Dolyhir Limestone, disused quarry 475 m W. of Dolyhir Bridge, Old Radnor,
Powys (SO 2409 5812): 2a-h, NMW 74.30G.12a, left free cheek, dorsal and left lateral views, x8.
3, NMW 74.30G.62b, right free cheek, ventral view, x8. 4, NMW 74.30G.15, left free cheek, oblique
dorsal view, x 8. Note course of connective suture. 5, NMW 72.18G.177, holotype cranidium, dorsal
view, x8. 6a~h, NMW 74.30G.23, pygidium and thoracic segment, left lateral and dorsal views, x8.
PLATE 95
OWENS and THOMAS, Radnoria
812
PALAEONTOLOGY, VOLUME 18
rarely with flat-topped profile; pygidial margin entire or with short spines; external
surface tuberculate, pitted, spinose, or smooth.
Stratigraphical range. Silurian (Wenlock Series) to Carboniferous, possibly to
Permian.
Genera. Australosutura Campbell and Goldring, 1960; Brachymetopus M’Coy, 1847;
Cordania Clarke, 1892; Mystrocephala Whittington, 1960; Proetides Walter, 1924;
Radnoria gen. nov. ; Tscliernyshewiella Toll, 1899; ICheiropyge Diener, 1897.
Discussion. Brachymetopus, Cordania, Mystrocephala and Australosutura are well
known and there is no doubt as to their membership of the family. We agree with
Hessler (1962, p. 812) that Proetides is a brachymetopid. No good illustrations are
available for Tscliernyshewiella, but we accept Whittington’s (1960, p. 407) observa-
tions on its close similarity to Cordania, and include it in the family. We consider that
Piltonia Goldring, 1955 should be excluded; both Hahn (1964, p. 362) and Osmolska
(1970, pp. 12, 13) considered it to be closely allied to ‘phillipsiid’ genera, in particular
to Eocyphinium Reed, 1942, a view supported by Owens’s unpublished work. Cheiro-
pyge, from the Permian of the Himalayas, was excluded from the family by Whitting-
ton (1960, p. 408). His reasons for doing so were not discussed, except to state that
the pygidium on which the genus is based did not, in his view, resemble those of other
brachymetopids. Schmidt {in Moore 1959, p. 0408) questionably included Cheiropyge
in the Brachymetopidae, and although existing illustrations (Diener 1897, pi. 1,
fig. 2a-c; Moore 1959, fig. 310.4, p. 0407) are poor, they do show that the pygidium
resembles spinose Brachymetopus pygidia. Until this genus can be revised, we follow
Schmidt, and provisionally assign it to the Brachymetopidae. Namuropyge R. and
E. Richter, 1939 was included with question in the Brachymetopidae by Schmidt {in
Moore 1959, p. 0408), but we agree with Whittington (1960, p. 408) that it should
be excluded; we also agree with Schmidt {in Moore 1959, p. 0408, footnote) and
Whittington (1960, p. 408) that Panarchaeogonus Opik, 1937 is not a brachymetopid,
and Owens (1974, p. 687) and Fortey and Owens (1975, p. 231) considered it to be
an otarionid.
Genus radnoria gen. nov.
Derivation of name. From Old Radnor, Powys, from where most of the material of the type species originates.
Type species. Radnoria syrphetodes sp. nov.
Other species. R. triquetra sp. nov., R. humiltima (Barrande, 1852).
Diagnosis. Preglabellar field concave or flat in longitudinal section; shallow depres-
sion traverses its posterior part and inner part of fixed and free cheeks, running
parallel to margin; anterior branch of facial suture diverges at 60-70° from an
exsagittal line through y; pygidial axis with 10-13 rings, pleural areas with 6-7 pairs
of ribs; latter with flat-topped profile or with posterior pleural band elevated above
anterior; dorsal surface smooth or with fine pits and sporadic granules.
OWENS AND THOMAS: RADNORIA
813
Radnor ia syrphetodes sp. nov.
Plate 95, figs. 1-6; Plate 96, figs. 1,2; text-fig. 1
V. 1972 Warburgella cf. stokesii (Murchison); Bassett, p. 31.
V. 1973 Warburgella (Warburgella) stokesii (Murchison); Owens, p. 67 pars, [reference only to
single specimen from Ty-newydd Farm].
V. 1974 IPrantlia sp.; Bassett, p. 759.
TEXT-FIG. 1. Reconstruction of Radnoria syrphetodes gen. et sp.
nov. Topmost illustration represents the ventral aspect of the
anterior part of the cephalon to show the inferred shape of the
rostral plate (blank area). All x 1 1 approx.
814 PALAEONTOLOGY, VOLUME 18
Derivation of name. Greek syrphetodes, ]\imb\ed together; reference to the mosaic of characters of various
genera seen in Radnoria.
Holotype. Cranidium NMW 72.18G.177, from Wenlock Series, Dolyhir Limestone (lower Wenlock);
disused quarry 475 m W. of Dolyhir Bridge, Old Radnor, Powys (SO 2409 5812) (Quarry ‘D’ of Garwood
and Goodyear 1918, pi. 7).
Paratvpes. Cranidia NMW 72.18G.178, 74.30G.4a-b, c, lOb-c, 13c-d, 18-19, 68, 73a, free cheeks NMW
74.30G.8-9, 10a, 11-13, 15-17, 62b, pygidia NMW 72.18G.179-180, 74.30G.3e-f, lOd, 14, 20-61, 62c-d,
67, 69-72, 73b, 74 from the type locality; pygidium NMW 74.30G.77 from shale band in Dolyhir Lime-
stone, type locality; cranidia GSM 103744, NMW 74.30G.93a-b, pygidia GSM DEX2927, NMW
74. 30G. 94-98 from limestone of probable lower-middle Wenlock age, old quarry opposite Ty-newydd
Farm, 1-3 km at 127° from Llanarthney church, Dyfed (SN 5442 1951). An ill-preserved cranidium, NMW
74.30G.99 from the Much Wenlock Limestone Formation (lundgreni Zone) of Wren’s Nest Hill, Dudley,
may also belong to this species.
Diagnosis. Glabella with weak forward taper; Ip furrows broad and shallow, Ip
lobes reduced; pygidium with 10-13 axial rings and six pairs of pleural ribs with
flat-topped profile.
Description. Cranidium moderately vaulted, palpebral width about two-thirds sagittal length. Glabella
about as wide as long, defined laterally by deep axial furrows; these shallow and narrow at Ip lobes, and
at anterolateral corner of glabella run into shallower preglabellar furrow, which is shallowest at sagittal
line. From its posterolateral corners, glabella tapers gently forwards to bluntly rounded frontal lobe. In
lateral profile it is gently convex, with the posterior end elevated well above the occipital ring (PI. 95,
fig. le). In transverse section it is strongly convex (PI. 95, fig. Id). Ip furrow runs into axial furrow opposite
anterior part of palpebral lobe, and is directed inwards and backwards at about 25° to an exsagittal line,
with steep adaxial slope and shallow abaxial slope. Ip furrow defines partially isolated, reduced, roughly
triangular Ip lobe, which is fused with remainder of glabella at its inner end. 2p furrow a smooth area,
not impressed, meeting axial furrow about two-thirds of way along glabella from its posterior end, directed
backwards at about same angle as Ip. 3p small, not reaching axial furrow, placed a short distance in front
of 2p, directed backwards at about 40°.
Occipital furrow rather broad and shallow in its median section, deepening markedly behind Ip lobes,
and arched forwards very weakly sagittally. Occipital ring about as wide (trans.) as base of glabella, its
sagittal length about one-third that of preglabellar area. No lateral lobes; presence or absence of occipital
node unknown. Preglabellar field about half sagittal length of glabella, weakly concave in longitudinal
section, anteriorly merges almost imperceptibly into gently upturned, weakly convex anterior border,
which is between one-third and one-half the sagittal length of preglabellar field. Preglabellar field traversed
on its posterior portion by shallow depression running parallel with margin, and which continues on to
EXPLANATION OF PLATE 96
Figs. 1-2. Radnoria syrphetodes gen. et sp. nov. Wenlock Series, Dolyhir Limestone, disused quarry 475 m
W. of Dolyhir Bridge, Old Radnor, Powys (SO 2409 5812). la-c, NMW 74.30G.33a, pygidium, dorsal,
posterior, and left lateral views, x8. 2, NMW 72.18G.179, pygidium, dorsal view, x8.
Figs. 3-5. Radnoria triquetra gen. et sp. nov. Wenlock Series, Much Wenlock Limestone Formation,
Nodular Beds, large bedding plane exposure on west side of Wren’s Nest Hill, Dudley, West Midlands
(SO 9350 9210). 3a-c, NMW 71.6G.239, holotype, internal mould of cranidium, dorsal, left lateral,
and anterior views, x 10. 4, NMW 71.6G.260, internal mould of cranidium, dorsal view, x8. 5a-d,
NMW 71.6G.240, pygidium, dorsal, right lateral, oblique posterodorsal, and posterior views, x 12.
Fig. 6. Radnoria humiUima (Barrande, 1852). Wenlock Series, Liten Formation, Lodenice, near Beroun,
Czechoslovakia. NMP 215/68, latex cast of cranidium, oblique anterolateral, right lateral, and dorsal
views, X 12. Original of Horny, Prantl and Vanek 1958, pi. 2, fig. 6.
PLATE 96
OWENS and THOMAS, Radnoria
816 PALAEONTOLOGY, VOLUME 18
field of free cheek. This depression appears to correspond roughly with the inner edge of the cephalic
doublure (PI. 95, figs. 2a, 3).
Anterior branch of facial suture diverges at 60-70° from an exsagittal line through y. y a broad curve,
a little way out from axial furrow. Palpebral lobe backwardly placed, subsemicircular and about one-third
sagittal length of glabella, e and | a single angle, a short distance in front of posterior border furrow.
Section e+^ to co short, nearly straight, defining minute, triangular posterior portion of fixed cheek. Visual
surface unknown. Distinct eye socle, the lower margin of which is not incised and which diverges strongly
from the upper margin at either end; median section directed almost exsagittally. Field of free cheek broad,
lateral border more upturned and a little narrower than anterior. Posterior border furrow shallow, com-
parable to lateral. Long genal spine with deep median groove, which dies out before reaching posterior
end. Cephalic doublure broad (PI. 95, fig. 3); connective sutures of rostral plate backwardly divergent
(PI. 95, fig. 4). Hypostome unknown. Thorax known only from one incomplete segment (PI. 95, fig. 6u-6).
Pygidium rather weakly vaulted, about two-thirds as long as wide. Axis about one-quarter greatest
transverse pygidial width, tapering gently backwards, not reaching posterior margin ; axial rings defined
by shallow ring furrows which become progressively shallower posteriorly. No postaxial ridge. Pleural
areas broad, weakly convex with six pairs of pleural ribs of flat-topped profile whose anterior and posterior
pleural bands are of approximately equal width (exsag.). Pleural furrows rather shallow, of constant depth
along their length, much deeper than ill-defined interpleural furrows, which become deeper at their abaxial
ends. Both pleural and interpleural furrows reach close to pygidial margin. No border, but marginal area
of pygidium flattened. Sculpture of fine pits (on preglabellar field, cheeks and Ip lobe) and granules (on
glabella, anterior border, and palpebral lobe) seen on some cranidia (PI. 95, fig. 1), and some pygidia have
very fine granulation (PI. 96, fig. 1) or pits (e.g. GSM DEX2927). The apparent absence of these sculptural
elements on some specimens (PI. 95, figs. 5, 6) may be a product of preservation or of variation. The similarity
of other features of all the material included in this species suggest that the presence or absence of fine
sculptural details is not of taxonomic significance.
Radnor ia triquetra sp. nov.
Plate 96, figs. 3-5
Derivation of name. Latin triquetrus, triangular; with reference to the shape of the glabella.
Holotype. Cranidium NMW 71.6G.239, from Wenlock Series, Much Wenlock Limestone Formation,
Nodular Beds (lundgreni Zone), large bedding plane exposure on west side of Wren’s Nest Hill, 200 m SW.
of ‘Caves’ public house, Dudley, West Midlands (SO 9350 9210).
Paratypes. Cranidium NMW 71.6G.260 and pygidia NMW 71.6G.240, 72.18G.181; horizon and locality
of holotype.
Diagnosis. Glabella triangular with distinct Ip lobes; pygidium with thirteen axial
rings, each with a posteriorly placed median node, and seven pairs of pleural ribs in
which posterior pleural band is elevated above anterior.
Description. Cranidium weakly vaulted, with palpebral width three-quarters of sagittal length. Glabella
as wide posteriorly as long, defined by deep axial furrows which merge anteriorly with shallow preglabellar
furrow. In lateral profile glabella gently convex, more strongly so in transverse section, with posterior end
elevated above occipital ring. Ip furrow runs from axial furrow backwards and inwards at 60° from an
exsagittal line, curved adaxially, into the occipital furrow, widening and shallowing at its posterior end.
Ip lobe isolated, semi-oval, about one-third glabellar length. 2p and 3p furrows not seen on available
material. Occipital furrow broad and shallow, arched very weakly forwards sagittally, deepening laterally
behind Ip lobes. Occipital ring nearly one-third length (sag.) of preglabellar field, and marginally wider
(trans.) than widest part of glabella. No lateral lobes. Preglabellar field approximately two-thirds sagittal
length of glabella, nearly flat in longitudinal section, traversed on its posterior portion by a shallow depres-
sion. Anterior border furrow weak, anterior border about half sagittal length of preglabellar field. Anterior
branches of facial sutures strongly divergent, each branch diverging at about 70° from an exsagittal line
through y, which is some distance out from axial furrow. Palpebral lobe crescentic, about half sagittal
OWENS AND THOMAS: RADNORIA
817
length of glabella. Posterior section of facial suture unknown. Hypostome, rostral plate, free cheek, and
thorax unknown.
Pygidium about two-thirds as long (sag.) as wide (trans.). Axis anteriorly one-quarter greatest pygidial
width, tapering gently backwards, not reaching posterior margin and with thirteen rings defined by
moderately distinct ring furrows which become progressively shallower towards posterior. Median node
on posterior edge of each ring. No postaxial ridge. Pleural areas broad, adaxial part horizontal in trans-
verse section, abaxial part rather steeply declined. Seven pairs of pleural ribs, with the posterior pleural
bands elevated above the anterior. Pleural furrows deeper and wider than the interpleural adaxially, but
abaxial ends of latter are deeper than corresponding sections of pleural furrows. Interpleural furrows reach
pygidial margin, pleural do not. Anterior and posterior pleural bands of approximately equal width (exsag.),
abaxial ends of latter distinctly elevated and crest-like. No border. Section of pygidial doublure seen
(PI. 96, fig. 5fl, c) shows that it has fine, parallel, terrace lines. Dorsal exoskeleton smooth.
Radnoria hiimillima (Barrande, 1852)
Plate 96, fig. 6a-c
*1852 Cyphaspis hiimillima Barrande, p. 492, pi. 18, figs. 57-58.
1868 Cyphaspis humillimus Barrande; Bigsby, p. 47.
1951 Otarion (?) humillimum (Barrande); Prantl and Pfibyl, pi. 1, figs. 27, 28.
V. 1958 Otarion! Immillinmm (Barrande); Horny, Prantl and Vanek, pi. 2, fig. 6.
V. 1970 Otarion! humillimum (Barrande); Horny and Bastl, p. 169.
Type specimens. Barrande (1852, p. 492) states that he had several specimens (all cranidia) at hand when
he erected this species. Therefore the statement by Horny and Bastl (1970, p. 169) that specimen NMP
IT309 is the holotype by monotypy is incorrect, and it is here designated lectotype. It is from high Liten
Formation (late Wenlock), Listice, near Beroun, Czechoslovakia. Barrande’s other syntypes are also from
this locality.
Other material. One cranidium NMP 215/68, Liten Formation, Lodenice, near Beroun. Figured Horny,
Prantl and Vanek 1958, pi. 2, fig. 6.
We have only had the opportunity to examine the specimen figured by Horny,
Prantl, and Vanek. This is poorly preserved and no preparation has been possible.
A full description and comparison must await revision of Barrande’s material, but
a latex cast is figured for comparison with the British species. Although R. humillimum
shows certain similarities to R. syrphetodes, it is distinguished by the relatively
longer, narrower, and more ovate glabella, while the anterior border is relatively
narrower (sag. and exsag.).
RELATIONSHIPS OF THE BR ACH YMETOPIDAE
As previously conceived, the Brachymetopidae comprised a number of Upper
Palaeozoic genera, the earliest being from the Lower Devonian. Different authors
have classified these trilobites in different ways; Prantl and Pfibyl (1951, p. 439) pro-
posed the Brachymetopinae as a subfamily of the Otarionidae; Hupe (1953, p. 220;
1955, p. 210) elevated them to family status, and believed that they were allied to
proetids rather than to otarionids; Maximova (1957, pp. 60-61) and Whittington
0960, p. 407) considered their morphology to suggest relationship with the phillip-
siids; Whittington and Campbell (1967, pp. 450-451) and Fortey and Owens (1975,
p. 23 1 ) surmised that they could have evolved from otarionids ; the latter also suggested
(1975, p. 231 ) a possible origin in the proetid subfamily Warburgellinae. So far, how-
ever, no convincing evidence has been advanced in support of either a proetid or an
otarionid ancestry.
818
PALAEONTOLOGY, VOLUME 18
In an attempt to trace brachymetopid relationships and ancestry, we have con-
sidered the following morphological characters: rostral plate, cephalic doublure,
lateral occipital lobes, preannulus, outline of pygidium, and structure of pygidial
pleural ribs. We have selected these features since they show the greatest variation in
the groups considered, and we believe that, taken together, they are of phylogenetic
significance.
Rostral plate. The Warburgellinae is the only proetid (sensu Owens 1973, p. 6) sub-
family in which the connective sutures diverge backwards. The connective sutures
also diverge backwards in Mystrocephala (Whittington 1960, pi. 54, fig. 3), Brachyme-
topus (Hahn 1964, pi. 32, fig. 3), and Australosutura (Amos, Campbell and Goldring
1960, pi. 39, figs. 10, 11; pi. 40, figs. 1, 5, 6) (although in the last two genera the
posterolateral corners of the greatly expanded rostral plate extend to the base of the
genal spine). The only species of Cordania in which any part of the rostral plate is
known is C.falcata Whittington (1960, p. 41 1, pi. 51, fig. 16), one specimen showing
the anterior part of the left-hand connective suture. Whittington points out that
the connective sutures converge backwards. He also figured free cheeks of C. macro-
bins (Billings, 1869) and one ofthese (ibid., pi. 53, fig. 10) shows a very broad doublure.
If the doublure of C. falcata is similar, only part of the connective suture is seen on
the specimen mentioned above. It may therefore be that the connective sutures of
C.falcata (and presumably other Cordania species) do diverge, after initially converg-
ing. As all other known brachymetopids have backwardly diverging connective
sutures it would be surprising if the same condition did not obtain in Cordania.
R. syrphetodes has backwardly diverging connective sutures (PI. 95, fig. 4).
In otarionids, and typical proetids, the rostral plate is small and triangular or
i tr ,
y
7
TEXT-FIG. 2. Schematic sections through doublures of free cheeks of: 1, Cor-
dania macrobius (Billings, 1869) (based on Whittington 1960, pi. 53, figs. 1,
10); 2, Australosutura gar dneriC&m^hcW and Goldring, 1960 (based on Amos,
Campbell and Goldring 1960, pi. 39, figs. 1, 10); 3, Proetides msignis iyJmcite.W,
1863) (after Hessler 1962, fig. 1a, p. 812); 4, Radnoria syrphetodes sp. nov.
(based on PI. 95, figs. 3, 4); 5, Proetus pluteus Whittington and Campbell,
1967 (based on Whittington and Campbell 1967, pi. 1, fig. 11; pi. 2, fig. 2);
6, Otarion plautum Whittington and Campbell, 1967 (based on Whittington
and Campbell 1967, pi. 7, figs. 1, 6); 7, Warburgella rugulosa canadensis
Ormiston, 1967 (based onOrmiston 1971, pi. 21, figs. 4, 5); 8, Prantliagrindrodi
Owens, 1973 (based on Owens 1973, pi. 15, figs. 3, 5). Arrows indicate position
of lateral border furrow, ‘tr’ the tropidium.
OWENS AND THOMAS: RADNORIA
819
trapezoidal with backwardly converging connective sutures (see Whittington and
Campbell 1967, pi. 6, figs. 2, 9; pi. 7, fig. 6; pi. 10, fig. 15).
Cephalic doublure. In many Proetacea (see Owens 1973; Whittington and Campbell,
1967; Ormiston 1971) the cephalic border and doublure form a ‘tube’ (Fortey
and Owens 1975, p. 236) (see text-fig. 2), where the inner margin of the doublure
coincides with the border furrow, which in all these cases tends to be well defined.
In some genera, as Hessler (1962, p. 811) has observed, the doublure extends well
inside the border furrow (which in these cases tends to be ill-defined), and its inner
section runs more or less parallel with the dorsal surface of the corresponding part
of the cephalon (see text-fig. 2). In such cases a nearly flat ‘trough’ is commonly
developed which crosses the cheeks and preglabellar field parallel to the cephalic
margins. The inner edge of this ‘trough’ corresponds with the inner edge of the
doublure, and is represented by an abrupt change in slope. This structure is found in
R. syrphetodes {P\. 95, fig. 2a), Prantlia grindrodi Owens (1973, pi. 15, fig. 3), Proetides
insignis (Winchell, 1863) and P. colemani Hessler, 1962 (see Hessler 1962, text-fig. 1,
p. 812), Cordaniamacrobius (QWWngs, 1869) (see Whittington 1960, pi. 53, figs. 10-12),
Mystrocephala pulchra (Cooper and Cloud, 1938) (see Whittington 1960, pi. 53,
fig. 15), and Australosutura gardneri (Mitchell, 1922) (see Amos, Campbell and
Goldring 1960, pi. 39, figs. 1, 6, 10, 11).
Lateral occipital lobes. These are of common occurrence in the Proetacea, but are
lacking in the Otarionidae, Brachymetopidae, and many Tropidocoryphinae. Lateral
occipital lobes are well developed in Silurian Proetinae (but absent in later members
of the subfamily), present in most Warburgellinae (but suppressed in some species
of Warburgella), and have evidently been repeatedly acquired and lost at various
times in different proetacean lineages.
Preannulus. This feature is found in the Proetinae and their derivatives— e.g. Dechenel-
linae and ‘phillipsiids’, but is lacking in Tropidocoryphinae, Warburgellinae, Brachy-
metopinae, and Otarionidae.
Structure and shape of the pygidium. Owens (1973, pp. 5-6; text-fig. 2, p. 5) has
recognized three types of pygidial pleural rib structure in Lower Palaeozoic Proetidae,
with different kinds characterizing the Tropidocoryphinae, Warburgellinae, and the
Proetinae and their derivatives. R. syrphetodes has pleural ribs like those found in
Warburgellinae. R. triquetra and other Brachymetopinae, however, have a different
type, in which the posterior pleural bands are elevated above the anterior; this type
of structure appears to be a modification of that typical of Warburgellinae. In
Otarionidae the structure is similar to that found in Proetinae.
Proetinae, Tropidocoryphinae, Warburgellinae, and Brachymetopinae normally
have a pygidium of subparabolic outline with five (or commonly many more) axial
rings. Within any one lineage there is commonly an increase in number of rings in
successively younger genera; less commonly there is a decrease. Otarionidae differ
from all the above groups in that the pygidium is short (sag.), with its width (trans.)
commonly over twice its sagittal length, and the number of axial rings never exceeds
seven, and is most commonly in the range three to four.
820
PALAEONTOLOGY, VOLUME 18
Inferred relationships. Brachymetopinae and Prantlia have in common the shape of
the rostral plate, type of cephalic doublure, lack of preannulus, pygidial outline, and
large number of pygidial axial rings. Other Warburgellinae share these characters,
but have a different kind of cephalic doublure structure. Proetinae and Otarionidae
have less in common with the above groups and, in particular, are distinguished by
the type of rostral plate and pygidium, and the possession of the preannulus in the
former. The Proetinae and their ‘phillipsiid’ derivatives, therefore, do not seem to
be closely related to the Brachymetopinae. Otarionidae bear a close general re-
semblance to certain Brachymetopinae, especially to Cordania species, in general
cephalic morphology. There are marked contrasts, however, in the structure of the
cephalic doublure, and probably also in the rostral plate. The similarity between
otarionids and Cordania is therefore considered to be due to homoeomorphy.
It would seem to be too great a coincidence for so many common features to be
independently acquired in brachymetopines and warburgellines and we consider
Radnoria and later brachymetopines to be derived from warburgellines along the
paths outlined below. The earliest known warburgellines are Warburgella species
from the mid-Llandovery (Owens 1973, p. 72). Warburgella has a backwardly
widening rostral plate, a tropidium or tropidial ridges, lateral occipital lobes, and the
pygidial pleural ribs are flat-topped in profile; the cephalic border and doublure
together form a ‘tube’. The earliest known Prantlia species is P. grindrodi Owens,
1973 from the highest Llandovery and Wenlock. The stratigraphical occurrence and
morphology of Prantlia suggest that it is derived from Warburgella through secondary
loss of the tropidium and by modification and widening of the cephalic doublure.
W. scutterdinensis Owens, 1973 from the early Wenlock is more similar to P. grindrodi
than is any other Warburgella species. In particular, it has a flat-bottomed ‘trough’
running parallel to the margin, similar to that typical of P. grindrodi. The presence
of this structure suggests that the doublure may be widened, but it is still unknown in
this species.
The pygidium of R. syrphetodes is similar to those of Prantlia species, particularly
that of P. longula (Hawle and Corda, 1847) (see Chlupac 1971, pi. 20, fig. 10). The
cephalon is also similar in its broad ‘trough’ and lack of tropidium but lateral occi-
pital lobes are absent, the Ip lobes reduced, and the anterior branches of the facial
sutures much more strongly divergent. R. triquetra is additionally distinguished by
the structure of the pygidial pleural ribs. The range of characters of Radnoria species
include some found in Prantlia (see above) and others— especially the strongly
divergent anterior branches of the facial sutures, lack of lateral occipital lobes, and
the pygidial pleural rib structure (of R. triquetra)^Io\xnd in Cordania. Radnoria thus
has a mosaic of Prantlia and Cordania characters, implying a close relationship
between the three genera.
Systematic position of the Warburgellinae q/ Radnoria. Hitherto, warburgellines
and brachymetopines have been classified in different families but, because of their
inferred relationships, we consider such a division to be artificial. Because war-
burgellines have more characters in common with brachymetopines than their
presumed proetid ancestors, the tropidocoryphines, we classify them with the former.
All three Radnoria species possess highly divergent anterior branches of the facial
OWENS AND THOMAS: RADNORIA
821
sutures and lack occipital lobes— features typical of brachymetopines. The pygidial
pleural rib structure of R. triquetra is also similar to members of this subfamily,
although that of R. syrphetodes is more like that of warburgellines. We consider that
brachymetopine characters outweigh warburgelline ones, and place Radnoria in the
Brachymetopinae.
Acknowledgements. We are indebted to H. B. Whittington, M. G. Bassett, and R. A. Fortey for valuable
discussion and criticism of previous drafts. D. E. White (IGS) and V. Zazvorka and F. Bastl (NMP) kindly
gave access to specimens in their care. We also thank Miss L. Cherns and P. D. Lane for assistance in the
field. A. T. T. acknowledges a Research Studentship from the N.E.R.C.; R. M. O. thanks the National
Museum of Wales for financial support for the fieldwork.
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R. M. OWENS
A. T. THOMAS
Department of Geology
National Museum of Wales
Cardiff CFl 3NP
Original manuscript received 31 December 1974
Revised manuscript received 16 February 1975
Department of Geology
Sedgwick Museum
Downing Street
Cambridge CB2 3EQ
A NEW CRAB, COSTACOPLUMA CONCAVA
FROM THE UPPER CRETACEOUS OF NIGERIA
by j. s. H. COLLINS and s. f. morris
Abstract. A new genus and species of retroplumid crab is described from the upper Cretaceous of Nigeria and
comparisons are made with Retropluma and the closely allied genus, Archaeopus. Archaeopus senegalensis Remy is
transferred to the new genus.
Included in a collection of fossils from the upper Cretaceous of Nigeria, deposited
in the Department of Palaeontology, British Museum (Natural History) by Professor
R. A. Reyment (formerly of the Nigerian Geological Survey), are two crabs from the
Coniacian of Abakaliki Province of East Central Region, three from the ?Maastrich-
tian of Shendam, Plateau Province of Benue-Plateau Region, and ten from the
upper Campanian of Enugu Province, East Central Region.
The transverse ridges on the dorsal surface of the carapace, the narrow front,
together with the structure of the orbital and antennular cavities clearly place these
crabs in the Retroplumidae as defined by Glaessner (1969, R531). The new material,
however, possesses features sufficiently distinct from Retropluma to allow a new
genus, Costacopluma, to be described.
SYSTEMATICS
Section brachyrhyncha Borradaile, 1907
Superfamily ocypodoidea Rafinesque, 1815
Family retroplumidae Gill, 1894
(= Ptenoplacidae Alcock, 1900)
Genus costacopluma gen. nov.
Type species. Costacopluma concava sp. nov.
Derivation of name. Referring to the strong transverse ridges and the familial root.
Diagnosis. Carapace transversely suboval with three transverse arched ridges, the
foremost extending across the protogastric lobes to unite with the mesogastric lobe;
the areas between the ridges are concave ; the lateral edges are thinly raised from the
front to the posterior ridge and the urocardiac depression is distinct. The 5th coxae
are subdorsal.
Costacopluma concava sp. nov.
Plate 97, figs. 1 -9
Derivation of name. The trivial name refers to the concave areas between the transverse ridges on the
carapace.
Diagnosis. Costacopluma with anterolateral notch, anterior transverse ridge reaches
[Palaeontology, Vol. 18, Vol. 4, 1975, pp. 823-829, pi. 97.]
824
PALAEONTOLOGY, VOLUME 18
the lateral margin, areas between ridges deeply concave. Flat triangular rostrum not
strongly produced.
Material. Fifteen more or less complete carapaces from three horizons in South-east Nigeria : Holotype,
In. 44642 (PI. 97, figs. 1-3) and paratypes, In. 44643-In. 44648, In. 44650-In. 44652, upper Campanian,
Zone of B. polyplocum, horizon of Libycoceras afikpoense, Anofia, on River Cross c. 8 miles (c. 12 km)
south of Afikpo Government Station, Enugu Province, East Central Region. Additional paratypes are
In. 46496-In. 46498, from ?Maastrichtian, Shendam, Plateau Province and In. 46499-In. 46500, Coniacian,
Awgu Limestone, Awgu, Abakaliki Province, East Central Region.
Description. The carapace is suboval in outline, the length being about two-thirds the
greatest width; it is gently arched longitudinally and transversely nearly flat. The
lateral margins are inclined almost at right angles to the dorsal surface and the lateral
edges from the front to the ridge on the metabranchial lobe, are drawn up slightly
into a thin rounded ridge. The anterolateral margin is short and the broadly rounded
lateral angle is about two-thirds distant from the front. The posterolateral angles are
sharp and lead by shallow incisions for the 5th coxae into the posterior margin which
is nearly straight, bounded by a low ridge and as wide as the orbitofrontal margin.
The orbitofrontal margin is nearly straight and occupies about three-quarters of the
greatest width of the carapace. The sharply triangular rostrum is steeply downturned
and thinly grooved round the apex of fused frontal lobes reaching nearly to its tip;
the frontal lobes become more elongate and less prominent as growth advances.
The orbits are subovate and directed forwards. The upper orbital margin, divided
by a blunt spine into two nearly equal parts, is thickened to form a low granulated
ridge and terminates externally in a rounded spine barely projecting beyond the
front. The short, similarly thickened, and granulated lower orbital margin terminates
at the buccal margin in a tuberant spine projecting somewhat beyond the upper
orbital margin. A thin septum separates the basal antennular segments which occupy
about a third of the antennular-orbital cavity; the segments are not well preserved,
but appear to be triangular with the angles much rounded. The orbital peduncle is
long, slightly contracted medially, and almost circular in transverse section. The
corneal surface is oblique and, where it is laying in position on the type, directed
downwards.
From a shallow marginal notch, the cervical groove curves inconspicuously to
a pit level with the narrow part of the mesogastric lobe, then commencing with another
pit the cervical groove deepens considerably and passes across the mid-line of the
carapace some two-thirds distant from the front. The dorsal surface is divided trans-
versely by three ridges; the foremost curves broadly forward from the lateral margin
across the protogastric lobes to unite with the tip of the mesogastric lobe; the crest
of the second ridge crosses the fused epi- and mesobranchial lobes and extends
EXPLANATION OF PLATE 97
Costacopluma concava gen. et sp. nov.
Figs. 1-5, 7-9 from the upper Campanian of Enugu Province, Nigeria. 1-4, holotype BM. In. 44642.
1, dorsal view. 2, right lateral view. 3, anterior view showing orbital peduncle. 4, ventral view. 5, 9, para-
type BM. In. 44644. 5, dorsal view. 9, anterior view. 7, 8, paratype BM. In. 44647. 7, dorsal view.
8, ventral view. All x2.
Fig. 6, from the ?Maastrichtian of Plateau Province, Nigeria. Paratype BM. In. 46497. Dorsal view, x 3.
PLATE 97
COLLINS and MORRIS, Costacopluma
826
PALAEONTOLOGY, VOLUME 18
downwards from the lateral margin towards the base of the mesogastric lobe. The
third ridge, on the metabranchial lobe, occurs about midway from the second ridge
to the posterior margin ; it is directed upwards in a broadly sinuous curve and although
interrupted by deep epimeral muscle scars, it continues across the carapace by a row
of four tubercles on the anterior of the cardiac region. The cardiac region is sub-
pentagonal in outline and has another small tubercle at its base. The mesogastric
lobe forms an elongated oval ; at its base are three forwardly directed pits set in an
inverted triangle, the lateral ones being the most prominent. On each metabranchial
lobe is an elongated tubercle close to the posterior margin. There is a tendency for
the ridges to become sharper as growth advances. While the subsurface shell layer of
the ridge tops exposed on the holotype is seen to be pitted, they are normally smooth
or lined with granules. The areas between the ridges are markedly concave and finely
pitted ; several larger pits are scattered along the outer course of the cervical groove in
addition to those already mentioned.
The triangular pterygostomian process is inflected almost at right-angles to the
carapace margin; a faint groove extending from the cervical groove curves towards
the buccal margin, and the sternal border is bounded by a strong granulated ridge
(In. 44647, PI. 97, fig. 7). The buccal cavity is about as broad as long and the margins
are straight. The ischiognath of the 3rd maxilliped is about twice as long as wide and
a shallow longitudinal depression reaches a little over one-half the width from the
convex inner margin; the outer margin is nearly straight. The merognath is subovate
in outline and almost as long as the ischiognath, a low ridge extends to the articulating
facet and the outer margin is thickened ; the three segments of the palp are of equal
length. The exognath tapers distally and reaches to about the middle of the merognath,
its width is a third of its length and there is a depression along the outer margin.
The abdominal sternites are very wide; the lst-3rd are separated by transverse
grooves and are divided by a median cleft widening posteriorly ; the groove separating
the 3rd from the 4th sternites runs back a short way from the margin before turning
sharply inwards. The 5th-7th sternites are drawn up into strong oblique ridges, while
the 8th is much reduced and subdorsal.
Male abdomina only are preserved and none is complete. The 4th-6th somites are
of about equal length and rapidly decrease in width so that the width of the posterior
margin of the 6th is one-half that of the anterior margin of the 4th; they are divided
from each other by sutures and each has a median transverse ridge. The telson is
about twice as long as broad, it widens slightly coincident with the 4th/5th sternal
groove before tapering to a broadly rounded apex. The abdominal trough is deep
and extends almost the entire length of the 4th sternite.
The specimens range in size from 6-9 mm to 26 mm across the carapace ; those from
the Coniacian and ?Maastrichtian localities are smaller than those from the Cam-
panian, but this may represent only a collecting bias, since fewer specimens were
collected from the first two localities.
Discussion. Hitherto, the earliest known member of the Retroplumidae has been
Retropluma eocenica Via. It differs from Costacopluma concava in having much
straighter anterior and posterior transverse ridges and in the posterior one (across
the metabranchial and cardiac lobes) being more entire; the anterolateral margin in
COLLINS AND MORRIS: CRETACEOUS CRAB FROM NIGERIA 827
C. concava is continuous with the general marginal curvature towards the front, but
in R. eocenica it is slightly hollowed.
The family Retroplumidae was erected by Gill (1894) to contain Retropluma Gill,
1894 ( = Archaeoplax Alcock and Anderson, 1894 non Stimpson, 1863) which is
represented in the Lutetian of Spain by R. eocenica Via, in the Pliocene of Italy by
R. craverii (Crema), and by four Recent species inhabiting the Indo-Pacific region.
Archaeopus senegalensis Remy (19606, p. 316) from the Palaeocene of Senegal is very
close to C. concava and must be included in Costacopluma. It differs from C. concava
by the absence of an anterolateral notch and the anterior ridge does not reach the
lateral margin ; the mesogastric lobe of C. concava is smaller and the transverse ridges
are narrower with steeper slopes and the areas between the ridges are more concave.
The median transverse ridge of C. senegalensis is more continuous, i.e. with shorter
gaps between the segments of the ridge. The triangular rostrum of C. senegalensis is
more strongly produced, with its median depressed and the margins elevated, also
the orbits are wider. C. senegalensis would appear to be a direct descendant of
C. concava.
TEXT-FIG. 1. Costacopluma senegalensis (Remy). Holotype,
X 2. Reproduced from Bull. Soc. geol. Fr. (7) 1, pi, 19a, fig. 1,
with the kind permission of the Societe geologique de France.
Beurlen (1930, p. 352) included the Cretaceous Archaeopus Rathbun, 1908 from
North America in the Retroplumidae, and whilst Glaessner (1969, R532) tentatively
placed it in the Palicidae, Via (1969, p. 339) again drew attention to the affinities of
this genus to Retropluma', also it appears to be closely related to Costacopluma.
Archaeopus is known by two species, A. antennatus Rathbun and A. vancouverensis
(Woodward) both from the upper Cretaceous (probably Campanian) of western
North America. Both species have well-developed ridges across the carapace, but the
gastric one (less well defined in A. antennatus) is more or less straight, not arched as
in Costacopluma, in which the branchial ridge also differs by curving down towards
the posterolateral angles. The cervical groove follows much the same course in both
828
PALAEONTOLOGY, VOLUME 18
genera, but does not weaken laterally in Archaeopus. The lateral margin of Costaco-
pluma is entire whereas the margin of Archaeopus is dentate. Via (1957, p. 554; 1969,
p. 339 et seq.) postulated that the origin of the Retroplumidae (in which he includes
Archaeopus, Re tr op luma, as well as Ophthalmoplax) lay in the Americas during the
Cretaceous. He suggested that a primitive, more robust, stock stayed in America—
Arehaeopus in North America and Ophthalmoplax in the region of the Gulf of Mexico
and northern South America— and another migrated eastward, adapting as it did
so, to deeper waters. It is difficult to accept entirely Via’s suggestion as to the origin
of the family since the Coniacian specimens from Nigeria are older than all the
American species, except for Ophthalmoplax comancheensis Rathbun (1935, p. 54)
which was based only on fingers from the Comanche Series (Albian) of Texas. It
would appear that Costacopluma started to adapt to deeper water within the North
African area. There is no evidence for any deep-water deposits in the Cretaceous of
Nigeria but by Palaeocene times Tessier (1952, p. 414) thought that the total Palaeo-
cene fauna of Senegal indicated a well-aerated, moderately deep sea of a maximum
of 50 m depth. Via (1969, p. 325) summarizes the details of catches of Recent Retro-
pluma spp. The shallowest species he records is R. denticulata Rathbun which is
caught off the coasts of Japan in the depth range 80-125 m. The presence of the
ocypodid Goniocypoda tessieri Remy in the Maastrichtian of Senegal suggests that
it is possible that Africa might have been the centre of ocypodid evolution. An east-
ward trend from Africa, possibly through southern Europe towards south-east Asia
had certainly been established by Miocene times for both the Retroplumidae and
Macrophthalminae.
Previous knowledge of fossil crabs from Nigeria has been limited to descriptions
by Withers (1924) of the xanthid, Holcocarcinus suleatus and a xanthid cheliped, both
from the middle Eocene (Lutetian) of Ameki, southern Nigeria. From nearby Senegal
and Ivory Coast, Tessier (1952) and Remy (1954; 1960a, b) have recorded: from the
Maastrichtian — Zanthopsis africana (Remy), Goniocypoda tessieri Remy; from the
Palaeocene — Necroearcinus simplex Remy, Raninella ornata Remy, Laeviranina sp.,
Pleolohites erinaceus Remy, Menippe frescoensis Remy, Zanthopsis multispinosa
Remy, Zanthopsis sp., Branchioplax ballingi Remy, Glyphithyreus wetherelli (Bell);
from the Ypresian— Glyphithyreus wetherelli', and from the Guitinm—Colneptunus
hungaricus lutetianus Remy, Colneptunus sp., Palaeocarpilius straeleni Remy, Micro-
maia simplex Remy, Atelecyclus gorodiskii Remy, Zanthopsis africana, and Branchio-
plax bcdlingi. Joleaud and Hsu (1935) described Necroearcinus multituberculatus
(Joleaud and Hsu) and a new genus of the family Potamidae from the upper Cretaceous
of the Niger Territory.
REFERENCES
ALCOCK, A. 1900. Materials for a Carcinological Fauna of India. No. 6. The Brachyura Catometopa, or
Grapsoidea. J. Asiat. Soc. Beng. 69, 279-456.
and ANDERSON, A. R. 1894. Natural History Notes from H.M. Indian Survey Steamer Investigator,
Commander C. F. Oldham, R.N. Commanding. Series II, No. 14. An Account of Recent Collections of
Deep Sea Crustacea from the Bay of Bengal and Laccadive Sea. Ibid. 63 (3), 141-185, 9 pis.
BEURLEN, K. 1930. Vcrgleichende Stammesgeschichte Grundlagen, Methoden, Probleme unter besonderer
Beriicksichtigung der hoheren Krebse. Fortschr. Geol. Palaeont. 8, 317-586.
COLLINS AND MORRIS: CRETACEOUS CRAB FROM NIGERIA
829
BORRADAiLE, L. A. 1907. On the classification of the Decapoda. Atm. Mag. not. Hist. (7) 19, 457-486.
CREMA, c. 1895. Sopra alcuni decapod! terziarii del Piemonte. Atti Accad. Sci. Torino, 30, 664-681.
GILL, T. 1894. A New Bassalian Type of Crabs. Am. Nat. 28, 1043-1045.
GLAESSNER, M. F. 1969. Decapoda: R399-533, 626-628. In moore, r. c. (ed.). Treatise on Invertebrate
Paleontology, Part R, Arthropoda 4 (2), Geol. Soc. America and Univ. Kansas Press.
JOLEAUD, L. and hsu, t.-y. 1935. Crustaces decapodes du Cretace de Tanout (Damergou Niger frangais).
Archs Mus. natn. Hist. nat. Paris, (6) 13, 99-1 10, 11 figs.
RAFINESQUE-SCHMALTZ, c. s. (rafinesque). 1815. Analyse de la nature, ou tableau de Tunivers et des corps
organises. 224 pp. Palermo.
RATHBUN, M. j. 1908. Descriptions of fossil Crabs from California. Proc. U.S. natn. Mus. 35, 341-349,
pis. 45-49.
1926. The fossil stalk-eyed Crustacea of the Pacific slope of North America. Bull. U.S. natn. Mus.
138, 156 pp., 39 pis., 6 text -figs.
1935. Fossil Crustacea of the Atlantic and Gulf Coastal Plain. Spec. Pap. geol. Soc. Am. 2, 160 pp.,
26 pis., 2 text-figs.
REMY, J.-M. 1954. In REMY, J.-M. and TESSiER, F. Decapodes nouveaux de la partie ouest du Senegal. Bull.
Soc. geol. Fr. (6)4, 185-191, pi. 11.
1960a. Etudes paleontologiques et geologiques sur les falaises de Fresco (Cote d’Ivoire). 2 Crustaces.
Annls Fac. Sci. Dakar, 5, 55-64, 1 pi.
19606. In GORODiSKi, a. and remy, j.-m. Sur les Decapodes eocenes du Senegal occidental. Bidl. Soc.
geol. Fr. (7) 1, 315-319, pi. 19a, fig. 1.
REYMENT, R. A. 1956. On the stratigraphy and palaeontology of the Cretaceous of Nigeria and the Cameroons,
British West Africa. Geol. For. Stockh. Fork. 78, 17-96.
TESSIER, F. 1952. Contribution a la stratigraphic et a la paleontologie de la partie ouest du Senegal (Cretace
et Tertiaire). Bull. Dir. Mines Geol. Afr. Occid.fr. 14, 1-465.
VIA, L. 1957. Contribution a I’etude paleontologique du Ocypodoida, Beurlen. C.r. hebd. Seanc. Acad. Sci.
Paris, 245, 553-554.
1969. Crustaceos Decapodos del Eoceno espanol. Pirineos, 91-94, 479 pp., 39 pis., 41 figs.
WITHERS, T. H. 1924. Eoccnc Brachyurous Decapod Crustaceans from Nigeria. Ann. Mag. nat. Hist. (9) 13,
94-97, pi. 5.
WOODWARD, H. 1896. On some Podophthalmatous Crustacea from the Cretaceous Formation of Vancouver
and Queen Charlotte Islands. Q. Jl geol. Soc. Land. 52, 221-228.
J. S. H. COLLINS
63 Oakhurst Grove
London, S.E.22
Typescript received 14 November 1974
Revised typescript received 10 April 1975
S. F. MORRIS
Department of Palaeontology
British Museum (Natural History)
London, SW7 5BD
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A LOWER PERMIAN TEMNOSPOND YLOUS
AMPHIBIAN FROM THE ENGLISH MIDLANDS
by ROBERTA L. PATON
Abstract. The holotype skull of Dasyceps bucklandi (Lloyd, 1850) from the Lower Permian (Autunian) Kenilworth
Breccia is redescribed and its relationships with other members of the family Zatrachydidae are discussed. The two
species of the American genus Zatrachys, the type species Z. serratus Cope, 1878 and Z. microphthalmus Cope, 1896
are considered. Z. microphthalmus is transferred to the genus Dasyceps as a new species D. microphthalmus, leaving
Z. serratus as the only species of the genus Zatrachys. It is shown that Acanthostomatops Kuhn, 1961 is not the
larval form of Dasyceps, but may instead be the larva of Z. serratus. It is suggested that Dasyceps was a terrestrial
labyrinthodont, and a possible function for its enormous median nasal vacuity is put forward.
The unique skull (Warwick County Museum, no. Gz 42) is the holotype of the species
Labyrinthodon bucklandi Lloyd, 1850. It was more fully described by Huxley (1859)
who recognized it as a form distinct from other known labyrinthodonts and separated
it as a new genus Dasyceps. Von Huene (1910) studied it further, giving the most
complete description to date. Case (1911) was the first to note its close relationship
to the aberrant family of labyrinthodonts, the Zatrachydidae. The other generally
recognized members of this family are Stegops Moodie, 1909, a primitive form from
the Westphalian D of Linton, Ohio (Romer 1930); Acanthostomatops Kuhn, 1961
(previously known as Acanthostoma Credner, 1883, but Kuhn pointed out that this
name was preoccupied by a polychaete), a small form found in the lower Permian
Niederhasslich deposits of Saxony (Steen 1937); and Zatrachys Cope, 1878 from the
lower Permian of Texas and New Mexico. The poorly known genus Platyhystrix
Williston, 1911, from the lower Permian of New Mexico, was thought to belong in
this family at one time, but this was because Williston (1911, 1916) figured what is
almost certainly a Z. microphthalmus skull (Langston, 1953) in artificial association
with long-spined Platyhystrix vertebrae, believing the two to belong to one genus.
This has given rise to the erroneous belief that some zatrachydids possessed ‘sails’
of pelycosaurian type on their backs. The affinities of Platyhystrix, which did have
a ‘sail’, are uncertain, and are irrelevant to the present topic; for a good discussion
of them see Langston (1953).
The Zatrachydidae have always been considered as rhachitomous labyrinthodonts,
and Romer (1947, 1966) placed them in the superfamily Eryopoidea. The dilferences
used by Case (1911) to distinguish Dasyceps from Zatrachys will be discussed later.
Dasyceps has been brieffy discussed by Romer (1930, 1939, 1945), but a detailed study
of it has not been attempted since von Huene (1910) examined it. Some authors
(e.g. Romer 1947, p. 172) have suggested that Acanthostomatops, which appears to
be a juvenile form, is in fact the larva of Dasyceps. At the same time Romer suggested
that Dasyceps and Zatrachys be synonymous, a view also held by Broom (1913).
These points will be discussed later.
The specimen, an almost complete skull, is in two pieces, part and counterpart.
[Palaeontology, Vol. 18. Part 4, 1975, pp. 831-845, pis. 98-99.]
832
PALAEONTOLOGY, VOLUME 18
Much of the skull roof has adhered to one portion, so exposing the ventral surface
of the bones with some impressions of their dorsal surfaces. The other portion shows
some of the skull roof, but anteriorly a considerable part of the dorsal surface of the
palate is exposed. The occiput has at some time been destroyed. Further preparation
of the specimen would appear to be impossible owing to its extremely fragile nature,
but the thick coat of dark shellac which covered both parts of the specimen, obscuring
all detail, has been removed. The skull has been crushed dorso-ventrally but it must
in life have been shallow anteriorly and of a moderate depth posteriorly.
The skull was found in a quarry close to Kenilworth itself (grid ref. SP 290720) ;
the quarry has long been disused and its exact location cannot now be determined.
The matrix is a coarse, red, loosely cemented sandstone containing pellets of red clay.
The horizon of the specimen is probably Autunian (Paton \91Ab).
DESCRIPTION
Von Huene (1910) described and figured the specimen fairly accurately, although his
plates show that it was then covered by the above-mentioned shellac. The skull is
much larger than those of other known zatrachydids, being twice as large as the
largest known specimen of Zatrachys; its maximum length is 298 mm and the maxi-
mum width is 230 mm. Its general shape, and the arrangement of the bones, is typical
of a rhachitomous temnospondyl, as can be seen from text-figs. 1-4 and Plates 98
and 99. Several features of interest are apparent.
TEXT-FIG. I. Specimen Gz 42, Dasyceps huckkmdi
(Lloyd), showing the ventral surface of the skull
roof. Stippled areas indicate where bone is missing.
(For abbreviations see p. 837.)
TEXT-FIG. 2. Reconstruction of skull roof of Dasyceps
bucklancii (Lloyd). (For abbreviations see p. 837.)
PATON: PERMIAN AMPHIBIAN FROM ENGLAND
833
The triangular skull is constricted slightly at the level of the maxillary /quadratojugal
suture, so that it appears to be swollen across the maxillae. Posterior to this con-
striction, the quadratojugals flare outwards in a lateral flange which shows signs of
having carried a bony frill at the edge. The quadratojugals also have posterior
extensions, which extend further back than the tabular horns. The latter rise to a level
somewhat above the rest of the skull surface. The postparietals are unusual in having
small posterior extensions; they are not, however, as prominent as was shown in
previous reconstructions. This feature has also been seen in ^ Platyhystrix' (Williston
1916), Zatrachys (Broom 1913), and, to a much lesser extent, in Stegops (Romer
1930). The orbits are very small and are situated in the posterior third of the skull.
They are elevated above the general skull surface and lie at the apices of two quite
sharply defined bony prominences, connected by a transverse ridge 16 mm in front
of the pineal foramen. Four other ridges radiate out from each prominence (these
are best seen as depressions on the ventral surface of the skull roof). The largest of
these extends forward antero-medial to the orbit to the level of the posterior edge
of the median nasal vacuity. The other three are of approximately equal size, one
extending laterally from the orbit to about the centre of the jugal, another extending
posteriorly from the orbit, and the third postero-medially to the middle of the parietal.
The height of these prominences and ridges is exaggerated antero-lateral and postero-
lateral to the orbit by two depressions: a particularly deep one on the jugal and
lachrymal, and a shallower one on the squamosal and jugal, the two being separated
by the bony ridges on the jugal. Case (1911) noted similarly situated preorbital
depressions in Zatrachys, although they were not at that time known to be present
in Dasyceps. Identically positioned ridges and depressions can be seen on the skulls
of the terrestrial labyrinthodonts Eryops (Cope 1877) and Peltobatrachus (Panchen
1959). Sawin (1941) has suggested that they were occupied by the trabecular cartilages
in life.
The nares are small, oval, with their long axes transverse, and lie far back, at the
junction of the premaxillae, maxillae, and nasals. Their posterior position is caused
by the very great enlargement of the premaxillae, which occupy approximately one-
third of the total skull length. The main zones of intensive growth (Bystrow 1935),
which determine the adult skull shape, are confined to (i) the premaxillae and maxillae ;
(ii) the jugal and quadratojugal; (iii) the tabular horns. There is a lesser zone across
the nasals and maxillae just posterior to the nares. There are no signs of these zones
posterior to the orbits. The pineal foramen, which is small, is thus situated very close
behind the level of the posterior edges of the orbits.
Probably the most dominant feature of the skull is the very large median vacuity
situated between the premaxillae and the nasals. It is drop-shaped, with the blunt
end facing anteriorly, and is 86 mm long and 38 mm wide. The edge of this median
nasal vacuity, where preserved, is smooth. The anterior part of the broken palate
shows the posterior edge of the median anterior palatal vacuity, which corresponds
in position to the dorsal vacuity, although the palatal vacuity is considerably smaller.
Only a small part of the dorsal surface of the palate is visible (text-fig. 3), but it
shows that the longitudinal elongation of the premaxillae also affected the palate.
In addition, the vomers are very large, extending into the posterior half of the skull.
This means that the interpterygoid vacuities, although not visible, must be relatively
834
PALAEONTOLOGY, VOLUME 18
TEXT-FIG. 3. Specimen Gz 42, Dasyceps bucklandi text-fig. 4. Reconstruction of palate of Dasyceps
(Lloyd), showing parts of the palate and skull roof. bucklandi (Lloyd). (For abbreviations see p. 837.)
Stippled areas indicate where bone is missing; rest
of cranial surface damaged except where ornament
is shown. (For abbreviations see p. 837.)
very small. Portions of the palatine and ectopterygoid bones can be seen, as well as
the posterior palatal extension of the maxilla, which extends back to about the level
of the squamosal/quadratojugal suture. The bases of three vomerine tusks can be
seen, one on the left side, and two on the right side ; one palatine tusk on each side
can be seen just posterior to each choana. The choanae are, in contrast to the external
nares, very large— they are oval in shape with the long axis antero-posterior and
38 mm in length, and the transverse axis 24 mm long. Because of the elongation of the
anterior part of the snout, they are situated about half-way along the skull.
Only a few marginal teeth are preserved, on the left premaxilla and maxilla. The
exact number is very difficult to determine because of the fragmentary nature of the
specimen in this region, but remains of eighteen teeth and seven sockets are thought
to be present in a maxillary length of 149 mm. Most of the teeth are broken off close
to the jaw, but two are almost complete. They are situated at the level of the choanae
and are 5 mm long, curving slightly inwards while being directed slightly outwards.
For their position and the over-all skull size, the teeth are small. The palatal tusks are
also relatively small.
The dorsal surface of the cranial bones is very poorly preserved, and only a small
EXPLANATION OF PLATE 98
Dasyceps bucklandi (Lloyd). Kenilworth. Ventral surface of skull roof, x^. Warwick County Museum,
Gz 42.
PLATE 98
PATON, Permian amphibian
836
PALAEONTOLOGY, VOLUME 18
part of the ornamentation is intact. It apparently consisted of very small, fairly
shallow pits separated by wide ridges which form blunt, upwardly directed points at
junctions of two or more ridges, thus giving the bone surface a pustular appearance.
A similar slightly pustular type of ornament can be seen in Stegops, "Platyhystrix',
and Zatrachys.
No trace of any lateral-line system can be seen, and it seems likely that von Huene
(1910) was incorrect in figuring parts of it in his reconstruction. The only other
member of the family in which this system has been figured is Stegops, and even here
there is some doubt about its presence.
No other material of Dasyceps bucklandi is known, although part of a rib in the
Institute of Geological Sciences, GSM 90490, from the lower Permian of Kenilworth,
is labelled as ‘? Dasyceps'. It appears to be a labyrinthodont rib, but is so poorly
preserved that further identification is impossible.
DISCUSSION
Case (191 1) noted the similarities between Dasyceps and Zatrachys, but distinguished
between them because of the apparent lack of the deep preorbital pits in Dasyceps,
and the apparent absence of the median nasal vacuity in Zatrachys. These features
were in fact later found in both genera, and Broom (1913) concluded that the two
were congeneric. This conclusion does not appear to have been widely accepted and
Romer (1966) still recognized both genera.
Langston (1953) has discussed the genus Zatrachys in considerable detail, based
upon a study of a large number of specimens of Z. serratus. A cast of one of the best
specimens used by Langston was available to the author. Case (1911) suggested that
Z. serratus and Z. microphthalmus were conspecific. Langston (1953), however, gives
fairly conclusive evidence that they are in fact distinct species. He cites many dif-
ferences in the skulls of the two species of Zatrachys, and goes on to list various skull
characters in these, and in Dasyceps bucklandi, Acanthostoniatops, and Stegops.
A modified version of this table, omitting Stegops and bringing the information on
Dasyceps up to date, is given here (Table 1). Some of the points Langston included
are left out, as they are considered irrelevant to the discussion on Dasyceps', these
are the basal articulation, ossifications in palatal roof, occiput, sclerotic plates, and
scutes.
This table again emphasizes the differences between Z. serratus and Z. micro-
phthalmus. Of the 17 characters listed here, they differ in 9 and agree in 5, while 3 are
unknown in one or the other. When Z. microphthalmus is compared with D. bucklandi,
there are only 2 differences, 4 unknown characters (the cultriform process in Dasyceps
is hypothetical and therefore considered here as an unknown character), and 1 1
similarities. Some of the characters are dependent upon the over-all size of the speci-
men, e.g. orbital size, depth of orbital pit, quadratojugal spikes, otic notch depth,
size of interpterygoid vacuities. The two features in which D. bucklandi differs from
Z. microphthalmus (the narrower otic notch and narrower internasal vacuity) are
here considered to be growth factors probably affected by the much larger size of
D. bucklandi. That Z. microphthalmus resembles D. bucklandi much more than it
PATON: PERMIAN AMPHIBIAN FROM ENGLAND
837
TABLE 1. Comparative osteology of the Zatrachydidae. Modified after Langston (1953).
Dasyceps bucklandi
Zatrachys
microphthalmus
Zatrachys serratus
A can thostomatops
vorax
Dorsal outline
Acute U-shape, slight
Acute U-shape, max.
Broad U-shape, no
Broad U-shape, no
max. exp.
exp.
max. exp.
max. exp.
Jaw articln.
Post, to occiput
Post, to occiput
In line with occiput
In line with occiput
Dermal bone
Pits + ridges, low
Similar to D. buck-
Pits-bridges, pro-
Similar to Z. serratus.
ornament
bosses on ridges
landi
minent spikes +
bosses
but less complex
Preorb. pit
Deep
Deeper
Deep
Shallower
L.l.g.
Not known
Not known
Not known
Not known
Cornua
Broad T and PP
cornua
Broad T and PP
cornua
Slender T and PP
cornua
Weak T cornua only
QJ spikes
Lateral QJ flange and
Like D. bucklandi but
Many long spikes on
3 spikes on QJ, small
postartic. proc.
less pronounced
QJ and postartic.
proc.
postartic. proc.
Orbits
Small, rims elevated
Small, rims elevated
Slightly larger, rims
less elevated
Large, rims slightly
elevated
Nares
Small, far posterior
Small, posterior
Large, posterior
Large, more anterior
Otic notch
Narrow, deep
Broad, deep
Broad, deep
Broad, shallow
Median nasal
Long, fairly narrow.
Shorter, wider, partly
Very large, separates
Small, hardly separates
vacuity
partly separates Ns.
separates Ns.
Ns., partly divides
Fs.
Same size as nares,
post, to nares
Ns.
Choanae
Larger than nares,
post, to nares
Not known
Same size as nares,
posterior to nares
Interpt.
vacuities
Widely triangular,
very short (?)
Widely triangular
Widely triangular
Widely triangular,
relatively large
Cultr, proc.
? short, broad
Broad, ? longer
Short, broad
Short, narrow
Ant. pal. vac.
Large
Not known
Not known
Large
Mandible
Not known
Not known
ANG. bSA. ridged,
low bosses on ANG.
5 long spines on ANG.
Dentition
Marginal teeth small.
Marginal teeth small.
Marginal teeth small.
Marginal teeth small.
3 tusk pairs
tusk pairs not
known
3 tusk pairs
3 tusk pairs
Abbreviations
used in text-figures and in Table 1
ANG
angular
PO
postorbital
ant. pal. vac.
anterior palatal vacuity
postartic. proc.
postarticular process
ch
choana
PP
postparietal
cultr. proc.
cultriform process
preorb. pit
preorbital pit
ECT
ectopterygoid
PRF
prefrontal
F
frontal
PSP
parasphenoid
interpt. vac.
interpterygoid vacuity
PT
pterygoid
J
jugal
p.t.
palatal tusk
L
lachrymal
Q
quadrate
l.l.g.
lateral line groove
QJ
quadratojugal
max. exp.
maxillary expansion
r.o.
ridges around orbit
MX
maxilla
SA
surangular
N
nasal
SM
septomaxilla
P
parietal
SQ
squamosal
p.a.p.v.
position of anterior palatal vacuity
ST
supratemporal
PF
postfrontal
T
tabular
PL
palatine
V
vomer
PMX
premaxilla
838
PALAEONTOLOGY, VOLUME 18
Zatrachys serratus Cope, after Langston (1953). /, ‘Platyhystrix' (probably Dasyceps microphthalmus),
after Williston (1911). g, ^Zatrachys' microphthalmus Cope, after Broom (1913). h, Dasyceps bucklandi
(Lloyd).
resembles Z. serratus can also be seen quite clearly from text-fig. 5. The positions
of the various sutures in Z. microphthalmus and D. bucklandi are very similar (although
Broom’s 1913 figure of the former shows a large lachrymal and small jugal, the
suture between these bones is dotted in, so presumably there is some doubt about its
position). The tabular meets the squamosal, excluding the supratemporal from the
otic notch, only in Z. microphthalmus and D. bucklandi.
Since the differences between Z. serratus and Z. microphthalmus are of a much
greater order than those between Z. microphthalmus and D. bucklandi, it is felt that
this confirms Broom’s (1913) decision that the latter two species belong to the same
genus. The name of Dasyceps Huxley, 1859 antedates that of Zatrachys Cope, 1878
and Z. microphthalmus must therefore become a species of Dasyceps, D. micro-
phthalmus. It was thought advisable to retain this as a distinct species, separate from
D. bucklandi, because of the poor illustrations of D. microphthalmus, the imper-
fections of the specimen of D. bucklandi, and the difference in size between the
specimens.
Langston (1953) considered that it would serve no useful purpose to synonymize
Zatrachys and Dasyceps. This is not the case, if only from a stratigraphical point of
view. The presence of two very closely related, possibly even conspecific, forms, one
in North America and the other in England, provides yet more evidence for the lower
Permian age of the Kenilworth Breccia and for the proximity of the two countries
at this time.
EXPLANATION OF PLATE 99
Dasyceps bucklandi (Lloyd). Kenilworth. Part of dorsal surface of skull roof and palate, xf. Warwick
County Museum, Gz 42.
PLATE 99
PATON, Permian amphibian
840
PALAEONTOLOGY, VOLUME 18
Z. serratus shows so many differences from the two species of Dasyceps that it is
considered necessary to retain it in a separate genus.
The other point put forward by previous authors is the possibility of Acantho-
stomatops being the juvenile form of Dasyceps. Probably the most complete growth
series known in fossil Amphibia is that shown by the skull of Benthosuchus sushkini
(Bystrow and Efremov 1940; Westoll 1950) in which most stages between skull
lengths of 30 mm and 600 mm are known. The development of the fairly elongated
skull can be seen easily when specimens of different sizes are reduced to a standard
width (Westoll 1950, fig. 26). A similar diagram has been produced for some growth
stages of Acanthostomatops, and for the skulls of D. bucklandi, D. microphthalmus,
and Z. serratus (text-fig. 5). From this diagram it can be seen that 'Platyhystrix'
(almost certainly a badly preserved skull of D. microphthalmus) and D. micro-
phthalmus are only slightly larger than the largest known specimen of Acantho-
stomatops and that there are substantial differences between the two forms in the
skull shape and in the positions of bones. Other dilferences are apparent from Table 1 .
The skulls of D. microphthalmus and D. bucklandi, however, although very different
in size, are very similar in shape and bone arrangement. Thus it seems most unlikely
that Acanthostomatops can be the larva of either species of Dasyceps.
However, a different picture emerges when Acanthostomatops and Z. serratus are
compared. Even the largest specimens of Acanthostomatops are believed to be
juvenile, and no true adult of this genus is known (Romer 1947; Langston 1953).
The converse is true of Z. serratus; many specimens are known, all are of similar size
and all are undoubtedly adults. Langston (1953, p. 396) states that ‘this suggests that
near adulthood was attained elsewhere, but nothing is known of the habitat’. The
skull width of the largest known Acanthostomatops specimen is 92 mm measured
across the quadratojugals, while that of the smallest Z. serratus is approximately
117 mm. It can be seen from Table 1 that, in a comparison between these two forms,
there are 1 1 similarities, 5 differences, and 1 unknown character. The only significant
difference between them is the apparent absence of the anterior palatal vacuity in
Z. serratus. Other differences (i.e. no postparietal cornua, small internasal vacuity,
short premaxillae and therefore more anteriorly placed nares, shallower otic notch)
are features which are likely to be associated with growth. Since juveniles of Z. serratus
are unknown, it is obvious that its larval stages grew and metamorphosed elsewhere—
perhaps this was a mechanism to prevent possible cannibalism by the adults. Equally,
no adults of Acanthostomatops are known, therefore metamorphosis occurred else-
where and the adult form inhabited a different environment. Because of this and the
many similarities between Acanthostomatops and Z. serratus, it is suggested that the
former is the juvenile form of Z. serratus. It would appear that, just before meta-
morphosis occurred, the juvenile migrated and only moved to the adult habitat after
it had reached a definite size. A possible reason for the large and relatively rapid
increase in size of the median nasal vacuity at metamorphosis is given later (p. 844).
It is not suggested that the juvenile Z. serratus migrated the considerable distance
which would separate Niederhasslich and New Mexico even when the effects of
continental drift are taken into account and the two placed on a single Laurasian
continent. It is merely thought to indicate that Z. .serratus was fairly widespread over
the whole of Laurasia (see later, p. 843), but that larva and adult, inhabiting different
PATON: PERMIAN AMPHIBIAN FROM ENGLAND
841
environments, would have been preserved under differing conditions, and therefore
would not have been preserved together.
Z. serratus Cope, 1878 has priority over A. vorax (Credner, 1883) so the species
remains Z. serratus. The apparent absence of the anterior palatal vacuity in the adults
of Z. serratus is most surprising in view of its large development in other members
of the family. Langston (1953) has indicated that a very small vacuity might perhaps
be present in the adult Z. serratus. He states, however, that the vomer is not usually
preserved and is in no case intact, so there is possibly some doubt about this point.
The author (1974a) has suggested that minor differences in the normal labyrintho-
dont ornamentation pattern of pits and ridges may be of taxonomic value, and it is
therefore interesting to note that the patterns found in the four groups discussed
above fall into two distinct types. That in Dasyceps bucklandi and D. microphthalmus
consists of very small, shallow pits separated by wide ridges which have low bosses
on them at their junctions. Langston (1953) describes that of the adult Z. serratus
as having prominent spikes and bosses superimposed on the ridges, which also appear
to be wide, separating small, shallow pits. He states that the ornament of ‘‘Acantho-
stomatops' is very similar to that of Z. serratus but is less complex. This is what would
be expected, as Bystrow and Efremov (1940) have shown that the juvenile pattern of
dermal bone ornamentation is less complex than that in the adult. This division of the
ornament into distinct groups confirms the conclusions reached above that D. buek-
landi and D. microphthalmus are closely related, as are Z. serratus and 'Acantho-
stomatops\ the two groups being separate.
Normal rhachitomous vertebrae are found in all the genera of Zatrachydidae (the
vertebral structure of D. bucklandi is unknown, but there can be little doubt that it
too was rhachitomous). Stegops is the earliest zatrachydid known but is considered
to be too aberrant to be ancestral to other members of the family, and Milner (pers.
comm.) suggests that it constitutes a separate family of specialized early dissorophoids.
The ancestry of the Zatrachydidae is unknown, but it seems probable that it was
derived from an early edopoid. It was included by Romer (1966) in the superfamily
Eryopoidea and it is not proposed to remove it from this superfamily, which contains
many widely divergent families and which is probably a polyphyletic assemblage of
advanced rhachitomes.
SYSTEMATIC PALAEONTOLOGY OF THE ZATRACHYDIDAE
Order temnospondyli
Superfamily eryopoidea
Family zatrachydidae
for family diagnosis see Langston (1953)
Genus dasyceps Huxley, 1859
Zatrachydids with acutely U-shaped skulls showing slight expansion across maxillae;
large median nasal vacuity; premaxillae much expanded antero-posteriorly; nares
far posterior; orbits small, in posterior third of skull, with rims much elevated; pro-
nounced lateral and posterior flanges on quadratojugal ; tabulars broad; small
M
842
PALAEONTOLOGY, VOLUME 18
postparietal process; supratemporal excluded from otic notch; ornament of pits and
ridges but with low bosses on the ridges; jaw articulation posterior to occiput.
Dasyceps bucklandi (Lloyd)
1850 Labyrinthodon bucklandi lAoyd, p. 56.
Very large zatrachydid with drop-shaped median nasal vacuity; orbits and nares
relatively very small ; laehrymal excluded from both orbit and naris.
Dasyceps microphthalmus (Cope)
1896 Zatrachys microphthalmus Cope, p. 436.
Small zatrachydid with oval median nasal vacuity; lachrymal excluded from naris
but not from orbit; quadratojugal flanges less pronounced than in D. bucklandi.
Genus zatrachys Cope, 1878
1878 Zatrachys serratus Cope, p. 523.
Zatrachys serratus Cope
Small zatrachydids with broad U-shaped skulls; no maxillary expansion; very large
median nasal vacuity in adult ; premaxillae with moderate antero-posterior expansion ;
nares slightly posterior and large; orbits small, in posterior half of skull, with slightly
elevated rims; lachrymal forms part of orbital border; quadratojugal flanges orna-
mented with long spikes; tabulars narrow; small postparietal process present in
adult; supratemporal forms part of edge of otic notch; ornament of pits and ridges
with prominent spikes and bosses on ridges; jaw articulation in line with occiput.
MODE OF LIFE
Dasyeeps and Zatrachys have always been considered as aquatic forms. The reason
for this is not clear and it seems possible that at least one species of Dasyceps was
terrestrial. Lateral-line grooves seem to be unknown in Dasyceps and Zatrachys, and
the skull appears to have been protected by an outstanding bony frill on the postero-
lateral edges; this was probably present in all members of the family but is not often
preserved complete. Such a bony frill would seem to be a distinct disadvantage to
a purely aquatic animal, as it would impede its progress through the water. In addition,
the skull shape, shallow anteriorly but moderately deep posteriorly and with ridges
radiating outwards from the orbits, is very similar to that found in known terrestrial
labyrinthodonts such as Eryops and Peltobatrachus. Advanced procolophonids and
the pareiasaurs, particularly Elginia, which were certainly terrestrial, also had spiny
edges on the posterior margins of the skull. Similar spines are found in present-day
lizards, e.g. Moloch horridus and Phrynosoma, where their function may be camouflage
as they break up the skull outline (Walker, pers. comm. ; Langston 1953). Thus from
an anatomical point of view it seems likely that Dasyceps and Zatrachys were terrestrial
forms. This view is confirmed by the nature of the sediments in which D. bucklandi
PATON: PERMIAN AMPHIBIAN FROM ENGLAND
843
occurs. The Kenilworth Sandstone or Breccia is a coarse, red, terrestrial deposit of
lower Permian (Autunian) age (Hains and Horton 1969; Paton 1974^). These deposits
are almost barren, the only other fossils known being three genera of pelycosaurs
(Paton 1974^?), a species of the conifer Lebachia (Walchia) (Hains and Horton 1969),
and some reptilian and amphibian footprints (Haubold 1970, 1971, 1972). Thus
D. bucklandi occurs in deposits of terrestrial origin and in association with a com-
pletely terrestrial fauna and flora. It therefore seems most unlikely that the species
was aquatic. Anatomical similarities in the skulls of D. bucklandi, D. microphthalmus,
and the adults of Z. serratus indicate that all three were probably terrestrial. Milner
and Panchen (1973) suggest that terrestrial animals were able to move freely over
the single continent of Laurasia during the lower Permian while a partial barrier seems
to have separated the aquatic tetrapods of the eastern and western parts of the super-
continent. If, as suggested here, D. bucklandi and D. microphthalmus, and Z. serratus
and ' Acanthostomatops' form two genera, the fact that the members of these two genera
are found widely apart on what was the Laurasian continent is added evidence for
their being considered terrestrial.
This leads to a consideration of the function of the relatively enormous median
nasal vacuity found in all zatrachydids except Stegops. The occurrence of a very small
interpremaxillary foramen is widespread among labyrinthodonts, and it is generally
accepted that this foramen was connected to a mucus-producing gland which had
another opening into the mouth in the anterior palatal vacuity. The mucus pre-
sumably lubricated the edges and inside of the mouth, and it has also been suggested
that it may have functioned either to attract prey or to repel predators (see Langston
1953). In all cases the interpremaxillary foramen is situated wholly between the pre-
maxillae and is in the vertical overhang of these bones above the mouth— a position
whence gravity would aid the mucus to run downwards to the mouth.
This is not the case in Dasyceps. The vacuity is situated between the premaxillae
and nasals, is very large, and lies horizontally on the dorsal surface of the snout. The
anterior palatal vacuity is positioned directly below it but is considerably smaller,
its position relative to the median nasal vacuity is shown in text-fig. 2. The appearance
of the two vacuities suggests that they were in fact confluent in life. Confluent fora-
mina between snout and palate are known, for example in Mastodonsaurus where
they accommodate the relatively enormous symphysial tusks of the lower jaws. This
is obviously not their function in Dasyceps. While it is possible that a small part of
the vacuity may still be glandular in function, it seems most unlikely that such an
enormous area could be given over entirely to mucus production. Assuming that
Dasyceps was terrestrial, sueh vast quantities of mucus spilling on to the skull surface
would cause considerable evaporation and consequent heat loss, and thus might be
disadvantageous.
D. bueklandi probably inhabited a fairly hostile environment. Its teeth are not large
and suggest a diet of small, fairly inactive animals. But it in turn may have been
hunted by predators including the large carnivorous pelycosaur Sphenacodon
britannicus which occurs at the same locality near Kenilworth. In common with other
large terrestrial labyrinthodonts, it was probably a ponderous animal which would
have to rely upon forms of protection other than a speedy retreat. No postcranial
material is associated with D. bucklandi but evidence (admittedly poor) from the
844
PALAEONTOLOGY, VOLUME 18
Other species of zatrachydids suggests that little armour was present on the body.
The skull, however, possesses a prominent bony frill around the posterior edge which
may have helped to camouflage the animal but which, in view of its size, also suggests
that the head could be used as a means of defending the whole animal. For this reason
it is very tentatively suggested that the large median nasal vacuity which is thought to
connect directly with the mouth may have been the site of an expandable sac which
could be inflated as an aggressive defence mechanism. The anterior palatal vacuity,
situated directly below and approximately in the centre of the median nasal vacuity,
but being much smaller, may have been the site of a valve which could cut off the
inflated sac from the mouth, thus enabling it to remain inflated independent of
respiration. This suggestion may appear unlikely at first sight, but such forms of
defence mechanisms occur in modern amphibians and reptiles and are known to deter
relatively large predators very effectively (see references in Cott 1940). It is also
possible that the throat could be inflated as in some modern frogs and the resultant
apparent increase in size combined with the visual effect of the bony frill round the
skull would probably be quite effective as a psychological deterrent.
The bony frill and internasal vacuity are relatively undeveloped in the juvenile
Zatrachys serratus although their development can be traced in the larger specimens.
If the functions for these structures suggested above are correct, they would of course
be unnecessary in the aquatic larval form, although the median nasal vacuity would
probably be glandular at this time.
Acknowledgements. I wish to thank Dr. A. D. Walker most sincerely for his continued help throughout this
work, and for his constructive criticisms of the manuscript. I am also grateful to Dr. S. M. Andrews for
reading the manuscript and for helpful suggestions, and Dr. A. R. Milner for useful discussions. I wish
to thank Miss J. Morris, Dr. W. Allen, and the Trustees of Warwick County Museum for their help and
permission to borrow and study Dasyceps bucklandi, and Mr. D. E. Butler of the Institute of Geological
Sciences, London, for the loan of specimens.
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R. Soc. B 242, 207-281.
PATON, R. L. 1974u. Capitosauroid labyrinthodonts from the Trias of England. Palaeontology, 17, 253-290.
19746. Lower Permian pelycosaurs from the English Midlands. Ibid. 541-552.
ROMER, A. s. 1930. The Pennsylvanian tetrapods of Linton, Ohio. Bull. Am. Mus. nat. Hist. 59, 77-147.
1939. Notes on branchiosaurs. Am. J. Sci. 237, 748-761.
1945. The late Carboniferous vertebrate fauna of Kounova (Bohemia) compared with that of the
Texas Red Beds. Ibid. 243, 417-442.
1947. Review of the Labyrinthodontia. Bull. Mus. comp. Zool. Harv. 99, 1-352.
1966. Vertebrate Paleontology. (3rd edition) Chicago, University Press. 468 pp.
SAWIN, H. J. 1941 . The cranial anatomy of Eryops megacephalus. Bull. Mus. comp. Zool. Harv. 86, 407-463.
STEEN, M. c. 1937. On Acanthostoma vorax Credner. Proc. zool. Soc. Lond. B 107, 491-500.
WESTOLL, T. s. 1950. In ‘A discussion on the measurement of growth and form.’ Proc. R. Soc. Lond. B 137,
490-509.
WILLISTON, s. w. 1911. American Permian Vertebrates. Chicago, University Press. 145 pp.
1916. Synopsis of the American Permo-Carboniferous Tetrapoda. Contr. Walker Mus. 1, 193-236.
ROBERTA L. PATON
Department of Geology
Royal Scottish Museum
Chambers Street
Edinburgh, EH 1 IJE
Original typescript received 22 November 1974
Revised manuscript received 17 February 1975
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A REVISION OF THE TRIASSIC TO LOWEST
JURASSIC DINOFLAGELLATE
RHAETOGONYA ULAX
by R. HARLAND, S. J. MORBEY and W. A. S. SARJEANT
Abstract. A re-evaluation of the dinoflagellate cyst genus Rhaetogonyaulax shows it to have a complex tabulation
of 4'-?6', 5a-?6a, lav, 7", 7c, 7'", Ip, Ipv, 3s, 3'"'. The archaeopyle is shown to be developed by loss of the epitract
anterior to the precingular plate areas. In consequence of recognition of the high degree of intraspecific variability,
all specimens examined are attributed to the single species R. rhaetica', the former species R. chaloneri is recognized
as an extreme in the morphological range and reduced to varietal status.
One of the authors (S. J. M.), in his research on the Rhaetic Formation (Morbey
and Neves 1974, p. 161) of a borehole at Bunny Hill, Nottinghamshire, and the
Rhaetian Stage of the Kendelbachgraben, Austria, encountered dinoflagellate cysts
attributable to the genus Rhaetogonyaulax Sarjeant, 1966. Since the holotypes of
the two species of this genus (including the type species) were lodged with H.M.
Geological Survey, now the Institute of Geological Sciences, a request was made
to I.G.S. for permission to study the type material, which was located, remounted,
and re-examined. In addition, it was decided to supplement the types by processing
topotype material from a sub-sample of the original rock specimen. Normal palyno-
logical processing techniques were used and a total of thirty-eight single grain mounts
were made, together with several strew mounts. The type material and topotype
material, together with other specimens from other localities under investigation by
S. J. M., are used here in a reconsideration of the genus.
The two described species of Rhaetogonyaulax have a particular significance in
dinoflagellate studies, since they are among the oldest undoubted dinoflagellate cysts
known. They have in consequence figured in many discussions of dinoflagellate
evolution (see Wall and Dale (1968, p. 288), Sarjeant in Erdtman (1969, p. 179),
Evitt in Tschudy and Scott (1969, p. 462), and Evitt (1970, p. 31)).
HISTORICAL BACKGROUND
In 1963 W. A. S. S. described two new species of dinoflagellates that had been dis-
covered, initially by Professor W. G. Chaloner, during a palynological study of H.M.
Geological Survey Stowell Park Borehole. The specimens were encountered at
a depth of 2059 ft 2 in (627-63 m) in Rhaetic (Upper Triassic) strata. Their tabulation,
which is poorly developed, was interpreted as being of a relatively simple character,
according with that of the living motile genus Gonyaulax\ in consequence, though
they were from the outset recognized to be cysts, they were named G. rhaetica and
G. chaloneri. The holotypes were deposited by W. A. S. S. in the collections of H.M.
Geological Survey. Archaeopyle formation was said to have taken place ‘by breakage
immediately anterior to, and not along, the transverse furrow’ (Sarjeant 1963, p. 353).
[Palaeontology, Vol. 18, Part 4, 1975, pp. 847-864, pis. 100-104.]
PALAEONTOLOGY, VOLUME 18
Sarjeant, in Davey et al. (1966), reviewing cysts with a Gonyaulax-iy^Q tabulation,
erected the new genus Rhaetogonyaulax to accommodate the species R. rhaetica and
R. chaloneri, arguing that, although they possessed a gonyaulacacean tabulation,
they were spindle-shaped and had an epitractal archaeopyle. This is in marked con-
trast to Gonyaulacysta (Deflandre) Sarjeant, the genus to which most fossil species
formerly attributed to Gonyaulax had by then been referred; this latter genus had
been diagnosed as being spheroidal, ovoidal, or polyhedral with a single-plate
precingular archaeopyle. Unfortunately, the transference of the two species to the
genus Rhaetogonyaulax did not conform to Article 33 of the International Code of
Botanical Nomenclature (Lanjouw et al. 1966), since their basionyms were not
clearly indicated; Loeblich and Loeblich (1968) rectified the position for Gonyaulax
rhaetica and subsequently Sarjeant, in Davey etal. (1969), did likewise for G. chaloneri.
The original studies of these cysts were made with a monocular petrological micro-
scope, with a maximum attainable magnification of X 800 and relatively low intensity
of illumination. The new studies here reported have taken full advantage of the
improvements in microscope technology since 1963. The work was done using
a Vickers microscope, Gillet and Sibert and Leitz photomicroscopes equipped for
phase-contrast work, and a Zeiss photomicroscope equipped for both phase-contrast
and Nomarski-interference contrast work. In addition, specimens were mounted for
scanning-electron photomicrography.
The morphology of this genus is one of especial complexity. It was early recognized
that the original interpretation of the tabulation was greatly oversimplified and that
archaeopyle formation was by schism between plates, not across plates as had
originally been believed. Moreover, difficulty was encountered in assigning speci-
mens to the two species originally proposed by Sarjeant. The results of our restudy
of the genus are presented below.
GENERIC RECONSIDERATION
Introduction
It has become increasingly apparent in dinoflagellate taxonomy that one of the
principal criteria in any systematic scheme is the form and method of archaeopyle
formation (Wall and Dale 1969, p. 287). Evitt (1967) has suggested, for instance, that
EXPLANATION OF PLATE 100
All figures at a magnification of x750 and in plain light unless otherwise stated.
Figs. 1-6. Rhaetogonyaulax rhaetica (Sarjeant) Loeblich and Loeblich emend. 1, dorsal view of specimen
MPK 805. On the epitract plates 4", 5", and 4a are most readily seen; on the hypotract, plate 5"'.
Cingular plate 4c is also well shown ; all other reflected plates are not in clear focus. Nomarski interference
contrast xc. 1200. 2, ventral view of MPK 805. Only the hypotract is in focus showing plates 6"',
T", 2'”', and 3"", together with 7c and the median sulcal plate. Nomarski interference contrast X c. 1 200.
3-6, holotype, specimen 1, PF 1983. 3, dorsal view. 4, ventral view. 5, dorsal view showing the antapical
horn, the initial separation of one of the intercalary plates (?5a). Phase contrast. 6, ventral view showing
some of the intercalary plates and the suture between those and the apical plates. Nomarski interference
contrast.
Figs. 7-8. Holotype of Rhaetogonyaulax rhaetica (Sarjeant) var. chaloneri stat. nov, specimen 2, PF 1983.
7, oblique dorsal view showing the reticulate ornamentation and sutures between precingular and inter-
calary plates. 8, oblique ventral view showing antapical horn and some precingular plates.
PLATE 100
HARLAND et a!., Rhaetogonyaulax
850
PALAEONTOLOGY, VOLUME 18
archaeopyle type or form should be used in circumscribing taxa at the generic level,
though it should be noted that Wall, Dale and Harada (1973) have demonstrated
some variation in archaeopyle form in Lingulodinium, a Cainozoic genus. Wall and
Dale (1968, table 2) included Rhaetogonyaulax with genera possessing an apical
archaeopyle, whereas Evitt (1967) regarded the genus as having an AP combination
archaeopyle, i.e. one in which all plates of the apical and precingular series were lost.
Archaeopyle formation and reflected tabulation
Detailed examination of cysts with archaeopyles shows that the margins left, after
the operculum has been shed, are regular in form and show a consistent ‘scalloping’
(PI. 101, figs. 5, 6 and PI. 102, figs. 2, 3). This does not accord with the cross-plate
schism originally visualized by Sarjeant (1963), which is in any case without parallels
in other genera; instead, it indicates that separation has taken place along the
boundary between two reflected-plate series. There is a distinct sulcal notch which
enables ready orientation of such specimens, and seven precingular plates, of very
meagre dimension, are shown to be present between cingulum and archaeopyle
margin. Some specimens appear to show only six precingular plates together with
a doubtful area around the sulcus (PI. 101, figs. 5, 6). We believe that the stereoscan
photomicrographs demonstrate the presence of seven precingular plates, the sulcal
area lying immediately to the right of reflected plate \ " (see especially PL 102, fig. 3).
A particular feature of this plate series is the prominent \" plate. Text-fig. 1a, b
TEXT-FIG. 1 . A, B, a semidiagrammatic sketch of Rhaetogonyaulax rhaetica (Sarjeant) Loeblich and Loeblich
emend, showing complete archaeopyle development, particularly the prominent 1" reflected plate and the
scalloped suture on the dorsal surface. A~ventral surface, b— dorsal surface, c, an interpretative semi-
diagrammatic drawing of the ventral surface of the holotype of Rhaetogonyaulax rhaetica (Sarjeant)
Loeblich and Loeblich emend, showing the tabulation where discernible. The dorsal surface of the specimen
is not shown, as no clear tabulation could be seen on that surface. (For abbreviations, see legend to text-
fig. 2.)
HARLAND ET AL.: DINOFL AGELL ATE RH AETOGONYAU LAX 851
gives a diagrammatic representation of a fully ruptured cyst of Rhaetogonyaulax
showing the precingular plate series.
In certain of the specimens studied (e.g. PI. 103, figs. 1-3), some plates anterior to
the precingular series are still attached to the abandoned cyst. This might result from
chance damage only; but we consider it much more probable that this is evidence
that the archaeopyle is developed by successive loss of opercular pieces, not by the
complete apical third of the cyst being simultaneously thrown off. The recovery of
complete apices with four definite (possibly six) apical plates still present, quite
separate from the anterior intercalary plates (PI. 103, fig. 4, and PI. 104, fig. 9),
supports this concept.
The plates posterior to the apical series and anterior to the precingular series, five
or six in number, appear to form a complete ring surrounding the epitract. However,
we do not consider them truly analogous to the anterior circle plates of Egmonto-
dinium (see Gitmez and Sarjeant 1972), since their position is rather on the flanks
of the cyst than on its anterior surface. One of the authors (Morbey 1975) has pro-
posed the term ‘postapical plates’ for a similarly positioned plate series in another
genus. For the moment, however, we have preferred to follow Wiggins (1973) by
designating these plates as ‘intercalary plates’.
We believe that the archaeopyle begins to develop by initial splitting along the
margins of some of the reflected intercalary plates (PI. 103, fig. 13; PI. 104, figs. 1-3).
One or more intercalary plate-areas separate from each other and from the cyst,
leaving the cyst otherwise intact. Further splitting results in the progressive loss of
the remaining intercalary plate-areas, the apex as a unit, the anterior ventral and
anterior sulcal plates ; the order of these events has not yet been definitely determined,
but it seems likely that the apex and sulcal plates are the last to detach. The holotype
of R. rhaetica shows initial separation of one of the intercalary plates (PI. 100, fig. 5;
text-fig. Ic), and other specimens have been observed in which the only discernible
rupture is between the apical and intercalary plates. Although several detached
apices have been found (see above), none show any surviving attachment to the
anterior ventral and anterior sulcal plates; the latter, if they separate (as seems
probable), would be so small as to be difficult to identify as such.
The reflected tabulation of Rhaetogonyaulax is extremely difficult to observe on
complete specimens, including the holotypes, whose orientation is consequently
often indeterminate. Indeed, the original photographs of the holotypes of R. rhaetica
and R. chaloneri are inverted, as are the specimens illustrated by Orbell (1973) and
Fisher (1972^). Some suture lines are occasionally evident, and in some cases the
plate boundaries are defined by partially or incompletely developed rows of ‘orna-
ment’. Plate areas are most readily to be observed after complete or partial rupture,
especially on the epitract. The hypotractal tabulation is especially difficult to decipher
clearly, as evidenced in the stereoscan photomicrographs. The interpretation here
presented was arrived at only after some weeks of study by one of us in particular
(W. A. S. S.), of specimens by Nomarski-interference contrast, in which it proved
possible laboriously to trace the plate boundaries (PI. 100, figs. 1, 2; PI. 101, figs.
1, 2). Even after this study, some doubt concerning the tabulation exists. In particular,
a rupture line has been observed on some specimens, perhaps separating off a poly-
gonal platelet at the apical end of plate \" (see text-fig. 1a). It is also possible that
852
PALAEONTOLOGY, VOLUME 18
additional posterior intercalary plates may be developed (see text-fig. 2d). Some
degree of variation in plate shape is also apparent, as seen especially in the most
prominent and easily recognizable plate, I" (see text-figs. 1 and 2; PI. 101, figs. 6, 12;
PI. 102, fig. 2; and PI. 103, figs. 7, 9).
The tabulation inferred is of great complexity, according neither with the gonyaula-
cacean lineages nor the peridiniacean lineages of Wall and Dale (1968). There is,
however, close accord with the tabulation of the Upper Triassic dinoflagellate
SImblikodinium Wiggins, 1973, which likewise has 4-6 apical plates lost together
with a complete series of ‘intercalary’ plates in archaeopyle formation, and has a
hypotract with three antapical plates. The two genera, though differing in symmetry
and in other tabulation details, are quite evidently closely related and may represent
an ancestral stock from which the two named lineages diverged. In fact, any residual
differences in tabulation are probably a result of two subjective interpretations. The
long-ranging genus Pareodinia Deflandre, whose tabulation has not yet been fully
determined but is known to be complex, may be a persistent representative of this
ancestral stock.
Variability
It is apparent that there is a great deal of variability in many aspects of the mor-
phology of Rhaetogonyaulax. The cysts are typically spindle-shaped but vary in
outline from slender and elongate to broad and squat (see Pis. 103 and 104); speci-
mens have been seen which appear to show the presence of an incipient second
antapical horn (see PI. 104, figs. 2, 5). Although this may be of some significance, the
authors wish to reserve further comment at this time. In addition, specimens have
been observed, typically from the Swabian Facies and the Limestone Li tliodendron
(Group VI), Kendelbachgraben, Austria, which show only rudimentary antapical
horn development (PI. 103, figs. 1, 2).
The nature of the cyst ‘ornamentation’ is such that specimens may carry variously
EXPLANATION OF PLATE 101
All figures at a magnification of x 750 and by Nomarski interference contrast unless otherwise stated.
Figs. 1-2, 5-12. Rhaetogonyaulax rhaetica (Sarjeant) Loeblich and Loeblich emend. 1, dorsal view of
MPK 806 showing a prominent cingulum, plates 2", 3", and 4" on the epitract and plates 2'" and 3"'
and the edge of plates \"' and Ip and the antapical plates 2"" and 3""; compare with text-fig. 3a.
xc. 1200. 2, ventral view of MPK 807, showing plates ?5a, ?6a, lav, la, and 2a on the epitract and
plates L", Ip, and Ipv on the hypotract, compare with text-fig. 3c. Plain light xc. 1200. 5, specimen
MPK 804, in dorsal view showing the ‘scalloping' and precingular plates 4", 5", and 6". 6, specimen
MPK 804, in ventral view showing the sulcus and prominent 1". 7, specimen MPK 803, ventral view
showing some intercalary and postcingular plates together with a broad sulcus and plate bounding
granular ornamentation on hypotractal plate 2"'. 8, specimen MPK 802, oblique ventral view showing
apical and intercalary plates. Plain light. 9, specimen MPK 802, oblique ventral view. 10, specimen MPK
802. oblique dorsal view showing the antapical horn. 1 1, specimen MPK 809, dorsal view showing broad
cingulum. Plain light. 12, specimen MPK 809, ventral view showing prominent 1 ", as and 7" plates.
Compare with fig. 6 for variation in plate shapes. Plain light.
Figs. 3-4. Rhaetogonyaulax rhaetica (Sarjeant) var. ehaloneri stat. nov. 3, holotype in oblique ventral view
showing the ornamentation. Phase contrast. 4, holotype in oblique dorsal view showing lack of orna-
mentation on the cingulum.
PLATE 101
HARLAND et ai, Rhaetogonyaulax
12
854
PALAEONTOLOGY, VOLUME 18
developed elements, i.e. spinelets, granules, and/or bacules. The elements of the
‘ornamentation’ (i.e. granules-spinelets) possess tapered or expanded stems and
truncate, furcate, or rounded terminations; a reticulation and a microgranulation
may also be developed. Any one specimen may carry several types of such abbreviate
‘processes’, in addition to being rough- or smooth- walled. (The holotype of R. chaloneri
(PI. 100, figs. 7, 8, and PI. 101, figs. 3, 4) carries both a reticulation and granules and
spinelets.) Occasionally, forms may occur which are more or less completely smooth-
walled and virtually devoid of ‘ornament’ (PI. 103, fig. 12). The ‘ornamentation’ is
so disposed that the cingulum may, for instance, be devoid of ‘ornament’ (PI. 102,
fig. 2) or may not differ in ornamentation from that on the epitract or hypotract
(PI. 102, fig. 1). The ‘ornamentation’ occurs in the form of random or orientated
intratabular or plate-bounding processes on the epitract and hypotract, being
distinctly plate-bounding between the cingulum and precingular and postcingular
plates, and between the sulcus and adjacent hypotractal plates. The continuation of
the sulcus on the epitract is defined either by plate-bounding processes or by an
apparent difference in texture between the wall structure of the sulcus and the
adjacent plate areas (PI. 103, fig. 2).
Variation in reflected plate morphology has also been noted and is commented
upon in the section on archaeopyle formation and reflected tabulation.
A biometrical study was undertaken on the topotype material. Sixty-five complete
specimens were used to construct the scatter diagram and the histograms seen in
text-figs. 3-5. The parameters used were the length (distance between the apex and
antapex), width, and the maximum cingulum width of the cysts and it was hoped to
learn something of the variation within the topotype population of Rhaetogonyaulax.
The resulting scatter diagram of cyst length/cyst width v. cyst length/width of cingu-
lum (text-fig. 3) produced a distribution of points indicating a relationship between
EXPLANATION OF PLATE 102
All figures at a magnification of xc. 1500 unless otherwise stated.
Figs. 1-9. Rhaetogonyaulax rhaetica (Sarjeant) Loeblich and Loeblich emend. Scanning electron photo-
micrographs of specimens showing archaeopyle formation and the variability of the cyst surface ‘orna-
mentation’. 1, a reticulate form in oblique ventral view that carries granules and bacules; the ornamenta-
tion is also present on the cingulum and sulcus. Plates ps, pv, Ip, T", and 2"' are possibly discernible.
MPK 820. 2, a form in lateral view showing complete archaeopyle formation, the seven precingular
reflected plate areas, and the over-all smooth cyst surface with some intratabular ‘ornamentation’ elements
and a cluster of intratabular elements in the centre of a postcingular plate area. MPK 815. 3, a specimen
in lateral view showing complete archaeopyle formation and a slightly rougher ‘ornament’ than fig. 2.
MPK 813. 4, as form in oblique ventral view showing a rough cyst surface with plate-bounding and
intratabular granules and bacules. Compare with text-fig. 2b. M PK 819. 5, a specimen in dorsal view show-
ing complete archaeopyle formation, rough ‘ornamentation’ and two antapical plates 1"" and 2"",
also Ipv, ps, and the inbulge of ms, a single intercalary plate, at least three precingular plates, three
postcingular plates, and two antapical plates. MPK 814. 6, form with well-developed granules and
bacules in oblique ventral view, plates ps, pv may also be seen. MPK 824. 7, detail of MPK 8 1 3 showing
the shape of the precingular plates 4" and 5" and a cingular plate boundary and their ornament, x c. 3000.
8, detail of MPK 8 1 4 showing the plates and ornament of the antapex, x c. 3000. 9, detail of MPK 820
to show the nature of the reticulate ornamentation in the sulcus, x c. 4500.
PLATE 102
HARLAND et al., Rhaetogonyaulax
856
PALAEONTOLOGY, VOLUME 18
the parameters, that the more slender the cyst the wider the cingulum width in relation
to the length. The data gives a correlation coefficient of 0-3706. It would appear that
only one population is involved here, with respect to the parameters measured, as
there is no suggestion of clumping or separation into two or more groups, and the
histograms of the frequency of cyst length/cyst width (text-fig. 4) and of cyst length/
cingulum width (text-fig. 5), are unimodal and are skewed to the right. The bio-
metrical study does not and cannot, however, in itself prove or disprove the existence
of one, two, or many species; but it is additional evidence for treating the topotype
specimens as belonging to a single population.
In the light of the present study, the authors feel that an emendation, not only of
the genus Rhaetogonyaulax but also of the species R. rhaetica, is necessary. Sarjeant’s
(1963) division into two species, rhaetica and chaloneri, based largely upon ornamenta-
tion, is no longer felt to be justified. In particular, we believe that Rhaetogonyaulax
EXPLANATION OF PLATE 103
All figures are phase contrast photomicrographs and at a magnification of x750 unless otherwise stated.
Slide co-ordinates given here and subsequently refer to microscope No. 158226 housed in the Depart-
ment of Geology, University of Sheffield.
Figs. 1-14. Rhaetogonyaulax rhaetica (Sarjeant) Loeblich and Loeblich emend. 1-9, cysts in various stages
of rupture, archaeopyle formation, and preservation. 1, specimen slide ref 23//lq/l/458/1254; Lime-
stone (Group VI), Kendelbachgraben, Austria; Rhaetian (sensu lato). Antapical horn
rudimentary in development and partial archaeopyle formation. 2, specimen slide ref 22>l /lq/1/347/1214;
as above, following incomplete rupture, a single intercalary, seven precingular reflected plates clearly
visible. Antapical horn rudimentary in development. Ornamentation well developed. Processes clearly
intratabular on anterior intercalary and precingular reflected plates. 3, specimen slide ref 5/ /2j/2/300/
1254; Swabian Facies (Lower), Kendelbachgraben, Austria; Rhaetian (.?. /.). Antapical horn showing
greater development and also showing precingular plates, one intercalary, and some apical plates.
Specimens not possessing clear antapical horns are morphologically very similar to some specimens of
Shublikodinium and possibly bridge the gap between the two genera. 4, specimen slide ref 26d/246/1305;
Westbury Member, Rhaetic Formation, Bunny Flill Borehole, Notts., England; Rhaetian. Detached
apical horn comprising at least four reflected apical plates. 5, specimen slide ref 52c/420/1254; Gotham
Member, Rhaetic Formation, Bunny Hill Borehole, Notts., England; Hettangian Stage, apical com-
pression of completely ruptured cyst. 6, specimen slide ref 44/ /x 13/beta/400/1276; x875. Equatorial
compression of completely ruptured cyst. Archaeopyle clearly evident. Precingular reflected plates
splitting apart. 7, specimen slide ref 44//x 13/beta/363/1291 ; x875. 6 and 7, in Salzburg Facies,
Kendelbachgraben, Austria; cyst illustrating archaeopyle formation. 8, specimen slide ref 14//lz/5/
252/1284; x 875; cyst illustrating archaeopyle formation and also the boundary between 2'"' and 3"".
9, specimen slide ref. 14//lz/4/332/1285; x 875. 8 and 9, in Swabian Facies, Kendelbachgraben, Austria;
cysts illustrating archaeopyle formation. 10-14, cysts illustrating various outline shapes and degrees of
development of body ornamentation. 10, specimen slide ref 52c/420/1254; Gotham Member, Rhaetic
Formation, Bunny Hill Borehole, Notts., England; Hettangian Stage; randomly ornamented with
processes. Orientation indeterminate. Gyst complete. 11, specimen slide ref. 29//lj/2/586/1276; Gar-
pathian Facies, Kendelbachgraben, Austria; Rhaetian (5. /.); x 875; sulcus well developed, ornamenta-
tion randomly developed. Sulcus tending to be broader in hypotract. 12, specimen slide ref 16//lx/l/
375/1294; Swabian Facies (Upper), Kendelbachgraben, Austria; Rhaetian {s. /.); x875; cyst un-
ornamented, wall smooth. Orientation indeterminate. 13, specimen slide ref 16//lx/3/269/1289; x875;
Swabian Facies (Upper), Kendelbachgraben, Austria. Orientation indeterminate. 14, specimen slide
ref 33b/418/1219; Westbury Member, Rhaetic Formation, Bunny Hill Borehole, Notts., England;
Rhaetian. Gyst inequifusiform in outline, wall reticulation superimposed by ‘ornament’.
PLATE 103
HARLAND et al., Rhaetogonyaulax
858
PALAEONTOLOGY, VOLUME 18
TEXT-FIG. 2. An interpretative reconstruction of the tabulation of
Rhaetogonyaulax rhaetica (Sarjeant) Loeblich and Loeblich emend.
A-B, specimen MPK 806. a, in left lateral view, slightly oblique;
B, in right lateral view, slightly oblique, c, specimen MPK 807
in ventral view, d-e, specimen MPK 805. d, in left lateral view;
E, in right lateral view. Abbreviations: l'-6', apical plates; lav,
anterior ventral plate; la-6a, anterior intercalary plates; as, anterior
sulcal plate; ms, median sulcal plate; l"-7", precingular plates;
lc-7c, cingular plates; l'"-7"', postcingular plates; Ipv, posterior
ventral plate; Ip, posterior intercalary plate; antapical
plates.
is a genus that has a demonstrably large variation of surface ornamentation, such
that there is no justification in treating a reticulate form as a separate species. The fact
that the topotype material, based upon certain measurable parameters, acts as a
unified population, is additional evidence for our view.
HARLAND ET AL.\ DINOFLAGELLATE RH AETOGONY AU LAX
859
30-
1-0-
-1 1 ^ r-
7 0 9 0 no 130
TEXT-FIG. 3. Scatter diagram of the topotype material based upon sixty-five complete speci-
mens. h'— holotype of Rhaetogonyaulax rhaetica; h”— holotype of R. chaloneri', m— mean.
The diagram has a correlation coefficient of 0-3706.
h
TEXT-FIG. 4. Histogram showing the frequency of text-fig. 5. Histogram showing the frequency of
the L/W parameter in the topotype material h', h", the L/Wc parameter in the topotype material h', h",
and m as in text-fig. 3. F calculated as percentages of and m as in text-fig. 3. F calculated as percentages of
the total. Standard deviation of L/W is 0-36. the total. Standard deviation of L/Wc is 1 -72.
860
PALAEONTOLOGY, VOLUME 18
SYSTEMATIC PALAEONTOLOGY
Division pyrrhophyta Pascher
Class DiNOPHYCEAE Pascher
Order peridiniales Lindemann
Genus Rhaetogonyaulax Sarjeant, 1966, emend.
1966 Rhaetogonyaulax 152.
1967 [Rhaetogonyaulax Sarjeant]; Evitt, p. 46.
1968 Rhaetogonyaulax Sarjeant; Loeblich Jun. and Loeblich III, p. 212.
1968 Rhaetogonyaulax Sarjeant; Wall and Dale, table 2.
1973 Rhaetogonyaulax Sarjeant; Lentin and Williams, p. 119.
Type species. Rhaetogonyaulax rhaetica (Sarjeant 1963) Loeblich Jun. and Loeblich III, 1968 O.D. emend.
Upper Triassic (Rhaetic), England.
Emended diagnosis. Cyst proximate, elongate-biconical to spindle-shaped, with rudimentary or pronounced
apical and antapical horns. Wall apparently single-layered, smooth, rough; punctate, granulate, or reticu-
late. Cingulum helicoid, laevorotatory, moderately indented; cingulum and sulcus generally defined by
ridges. Tabulation 4'-?6', 5-?6a, lav, 1", 7c, 1"', Ip, Ipv, 3s, 3""; boundaries of plate-areas marked by
raised lines, lines of short processes, or rupture. Processes, where developed, may be sutural or intratabular,
simple or furcate. Archaeopyle development by progressive loss of all plates anterior to the precingular
series.
Remarks. Wiggins (1973, p. 4) commented that ‘it is remarkable that Shublikodinium
superficially resembles Rhaetogonyaulax ... in both cyst and archaeopyle outline,
when their sutural tabulation series is so different’. Our restudy emphasizes the close
comparability of the two genera ; residual differences are in over-all shape, Shubliko-
dinium being ovoidal with two antapical horns, one of which may be reduced, whereas
EXPLANATION OF PLATE 104
All figures are phase contrast photomicrographs and at a magnification of x 875.
Figs. 1-12. Rhaetogonyaulax rhaetica (Sarjeant) Loeblich and Loeblich emend. 1-5, cyst specimens
illustrating partial rupture of some anterior intercalary plates, variation in over-all outline with
development of antapical horn, and broad sulcus on hypotract. 1, specimen slide ref. 44/ /x 13/beta/
400/1276; Swabian Facies, Kendelbachgraben, Austria; Rhaetian (5. /.). Cyst equifusiform in outline,
broad sulcus on hypotract. 2, specimen slide ref. 33a/274/1248; Westbury Member, Rhaetic Formation,
Bunny Hill Borehole, Notts., England; Rhaetian. On epitract, apical, anterior intercalary, and pre-
cingular reflected plates clearly defined, broad sulcus on hypotract, rudimentary secondary (?) horn
developing (? process) in addition to prominent antapical horn. 3, specimen slide ref. 44/ / x 13/gamma/
397/1310, Salzburg Facies, Kendelbachgraben, Austria; Rhaetian {s. /.). 4, specimen slide ref. 54/ /x4/
5/589/1290; Pre-planorbis Beds, Kendelbachgraben, Austria; Rhaetian (.9. /.). 5, specimen slide ref
5//2j/beta/380/1295; Swabian Facies (Lower), Kendelbachgraben, Austria; Rhaetian. Rudimentary
antapical horn (? process), coarsely ornamented. 6-8, 10-12, cyst specimens showing variation in over-all
outline, preservation, and development of ornamentation. 6, specimen slide ref 55// x 3/gamma/612/1200;
Pre-planorbisBeds, Kendelbachgraben, Austria; Rhaetian. 7, specimen slide ref 45/ / x 12/beta/305/1283; 1
Salzburg Facies, Kendelbachgraben, Austria; Rhaetian. 8, specimen slide ref 44/ /x 13/beta/400/1276; j
Salzburg Facies, Kendelbachgraben, Austria; Rhaetian (s. /.). 9, specimen slide ref 56/ / x 3/1/691/1230; j
Pre-planorbis Beds, Rhaetian. Kendelbachgraben, Austria; apical horn comprising at least four pro- 1
minent reflected apical plates. 10, specimen slide ref 44//x 13/gamma/588/1303, Salzburg Facies, 1
Kendelbachgraben, Austria; Rhaetian (s. /.). 11, specimen slide ref 29//lj/2/586/1276; Carpathian 1
Facies, Kendelbachgraben, Austria; Rhaetian. Coarse body ornament. Antapical horn developed 1
(preservation ?). 12, specimen slide ref 41d/352/1228; Cotham Member, Bunny Hill Borehole, Notts., !
England, Rhaetic Formation; Hettangian. Oblique ventral view, with plates I"', 2'" visible also Ip
and much of 7'", 1'"', and 3"".
PLATE 104
HARLAND et at., Rhaetogonyaulax
862
PALAEONTOLOGY, VOLUME 18
Rhaetogonyaulax is basically spindle-shaped, rarely ovoidal, and rarely exhibiting
two antapical horns. Rhaetogonyaulax is also much more sharply attenuated than
Shublikodinium. The two genera also differ in tabulation details, especially on the
hypotract, as Rhaetogonyaulax possesses postcingular, posterior intercalary, sulcal
and antapical plates as opposed to the postcingular and antapical plates of Shubliko-
dinium. Since the plate designated V by Wiggins (1973, text-fig. 2) does not in fact
form a part of the apex, we have preferred to refer to it as an anterior ventral plate
(lav) in reference to its position. The plate which Wiggins (1973) referred to as the
‘apical closing plate’ is here designated V .
Rhaetogonyaulax rhaetica (Sarjeant 1963) Loeblich and Loeblich 1968 emend.
Plates 100-104; text-figs. 1-2
V* 1963 Gonyaulax rhaetica Sarjeant, p. 353, figs. 1, 2 left.
1963 Gonyaulax chaloneri Sarjeant, p. 354, figs. 2 right, 3.
1964 Gonyaulax rhaetica Sarjeant; Downie and Sarjeant, p. 115.
1966 Gonyaulax chaloneri Sarjeant; Downie and Sarjeant, p. 114.
1966 Rhaetogonyaulax rhaetica Sarjeant nom. nud.\ Sarjeant, p. 153.
1966 Rhaetogonyaulax chaloneri Sarjeant nom. nud.; Sarjeant, p. 153.
1968 Rhaetogonyaulax rhaetica (Sarjeant) Loeblich and Loeblich, p. 212.
1969 c/ta/o/ten (Sarjeant); Sarjeant, p. 15.
\912b Rhaetogonyaulax sp.; Fisher, p. 105, pi. 2, fig. 15.
1973 Rhaetogonyaulax rhaetica (Sarjeant) Loeblich and Loeblich; Orbell, pi. 2, fig. 1.
1973 Rhaetogonyaulax rhaetica (Sarjeant) Loeblich and Loeblich; Lentin and Williams, p. 120.
1973 Rhaetogonyaulax chaloneri (Sarjeant) Sarjeant; Lentin and Williams, p. 1 19.
1974 Rhaetogonyaulax rhaetica (Sarjeant) Loeblich and Loeblich; Morbey and Neves, p. 168,
pi. Ill, figs. 3, 4.
1975 Rhaetogonyaulax rhaetica (Sarjeant) Davey, Downie, Sarjeant and Williams; Felix, pi. II,
fig. 2.
1975 Rhaetogonyaulax rhaetica (Sarjeant) Loeblich and Loeblich; Morbey, pi. 14, fig. 17, pi. 15,
figs. 1-4.
Emended diagnosis. Cyst proximate, typically spindle-shaped, unornamented or ornamented. Antapical
horn may be rudimentary or well developed. Wall apparently single-layered, thin, smooth, rough, punctate,
reticulate or granulate. Processes small, variable in development and distribution; random or orientated,
intratabular or plate-bounding, simple and furcate. Cingulum helicoid laevorotatory, unornamented or
ornamented, forming a moderately indented furrow; displacement may be as much as twice the cingulum
width. Seven cingular plates are developed. Sulcus ornamented or unornamented, generally expanded on
the hypotract. Cingulum and hypotractal sulcus clearly defined by plate-bounding processes or ridges; the
epitractal sulcus defined by plate-bounding processes or by textural differences between the epitractal plate
wall and the sulcal wall. Plate sutures most readily evident where partial or complete detachment of opercular
plates has occurred; hypotractal tabulation especially difficult to define. Reflected tabulation 4'-?6', lav,
5a- ?6a, 7”, 7c, 1"', 1 p, 1 pv, 3a, 3"" ; an additional plate-area may be present in the anterior extension of plate
la as here delineated. Archaeopyle formed initially by detachment of one or more intercalary plates, there-
after by loss of the remaining intercalary plates, the apex as a unit, and the anterior ventral and anterior sulcal
plates; cysts with archaeopyles thus show a ‘scalloped’ edge formed by the anterior margins of the pre-
cingular plates and a deep sulcal notch resulting from the loss of the anterior sulcal plate. The median
sulcal plate is often deeply indented and probably corresponds to the position of origin of the two flagella.
Typification. Holotype specimen 1, slide PF 1983 (not specimen 98, slide PS 1983, as quoted by Sarjeant
(1963)).
Type locality. Rock specimen Bj 6011, Stowell Park Borehole (N.G.R. SP 0835 1 173), Northleach, Glou-
cestershire. Depth 2059 ft 2 in, i.e. 627-63 m.
Repository. The holotype and all figured specimens from the topotype material are held in the Palynological
HARLAND ET AL.: DINOFLAGELLATE RH AETOGON Y AU LAX
863
Collections of the Institute of Geological Sciences, Leeds, England, and are registered in the PF and MPK
collections. Comparative material from the Bunny Hill Borehole, Nottinghamshire, and the Kendel-
bachgraben, Austria, is held in the collections of the Department of Geology, University of Sheffield,
Dimensions. Holotype: cyst length 63-75 /urn, width 37-5 jum, cingulum width 8-75 /nm.
Topotype material; length 47-5 (64-72) ll-S fj.m, width 17-5 (35-72) 48-75 jum; cingulum width 5-0 (7-73)
1 1 -25 /Lim, based upon sixty-five complete specimens.
Bunny Hill Borehole and Kendelbachgraben material: cyst length 50-0 (69-0) 92-0 /um; width 34-0 (43-0)
54-0 /rm ; cingulum width 7-0- 13-0 /^rm, based upon thirty-one complete specimens. (The data given are the
minimum, mean, and maximum measurements.)
Rhaetogonyaulax rhaetica (Sarjeant) var. chaloneri, stat. nov.
Plate 100, figs. 7, 8; Plate 101, figs. 3, 4
v*1963 Gonyaulax chaloneri Sarjeant, p. 354, figs. 2 right, 3.
1964 Gonyaulax chaloneri Sarjeant; Downie and Sarjeant, p. 1 14.
1966 Rhaetogonyaulax chaloneri (Sarjeant), nom. nud. ; Sarjeant, p. 153.
1969 Rhaetogonyaulax chaloneri (Sarjeant) ; Sarjeant, p. 15.
1973 Rhaetogonyaulax chaloneri (Sarjeant); Lentin and Williams, p. 1 19.
Typification. Holotype specimen 2, slide PF 1983.
Remarks. Rhaetogonyaulax chaloneri is here redesignated as a variety of R. rhaetica
as no justification can be found in maintaining this form as a distinct species in an
obviously variable morphological group.
Geological and geographical range o/ Rhaetogonyaulax rhaetica
Cotham Beds [Rhaetic], Stowell Park Borehole, Gloucestershire, England (Sarjeant 1963); Cotham Beds
[Rhaetic] and Pre-planorbis Beds [Lias], Barnstone Railway Cutting, Nottinghamshire, England (Fisher
1972a); lower ‘Cotham Beds’ [lower Rhaetian and basal upper Rhaetian], Bristol Channel region, England
(Fisher 19726); Grey Marls, Westbury Beds, Cotham Beds, White Lias, lowermost Watchet Beds [Rhaetic],
Lavernock Point, Glamorgan, Wales. Westbury Beds, Cotham Beds [Rhaetic], Owthorpe, Nottingham-
shire, England. Westbury Beds, Cotham Beds [Rhaetic], Upton Borehole, Oxfordshire, England (Orbell
1973); Tea-green Marl Member, Parva Formation and Westbury Member, Cotham Member, Rhaetic
Formation [Rhaetian sensu lato and Hettangian], Bunny Hill Borehole, Nottinghamshire, England.
Swabian Facies, Limestone ±Lithodendron (Group VI), Carpathian Facies, Kossen Facies, Salzburg
Facies, Lre-planorbis Beds [Rhaetian sensu lato and Hettangian], Kendelbachgraben, Austria (Morbey
and Neves 1974); Highest Grey Marls [Keuper Marl], Westbury Beds, lower Cotham Beds [Rhaetic],
Watchet, Somerset, England (Warrington 1974); Westbury Beds, Cotham Beds [Rhaetic], Larne Borehole,
Antrim, Northern Ireland (Warrington and Harland in press). Westbury Beds, Cotham Beds [Rhaetic],
Steeple Aston and other boreholes near Chipping Norton, Oxfordshire, England. Westbury Beds, Cotham
Beds [Rhaetic], borehole near Chipping Sodbury, Gloucestershire, England (G. Warrington, pers. comm.).
Flatsalen Formation [?Rhaetian], Hopen, Svalbard, Norway (D. Smith, pers. comm.) and recorded as
cf. B R. rhaetica (Smith, Harland and Hughes 1975).
Acknowledgements. The authors would like to thank Dr. G. Warrington for his assistance in reprocessing
the topotype material, for his discussions, and for taking certain of the scanning electron photomicrographs.
Mrs. Brenda Coleman has also assisted with the scanning electron microscope work; her help is gratefully
acknowledged. S. J. Morbey is most grateful to the Natural Environment Research Council for financial
assistance toward his research and to Professor Moore, Dr. R. Neves, and the Department of Geology,
University of Sheffield, for the facilities provided. One of us (R. H.) publishes with the permission of the
Director, Institute of Geological Sciences, London. The work of W. A. S. Sarjeant was supported by
Research Grant A-8393 of the National Research Council of Canada. Helpful discussions in correspondence
with Dr. David Wall (Woods Hole Oceanographic Institution) and the provision of specimens by Mr.
Michael J. Fisher (Robertson Research, Calgary, Alberta) are also gratefully acknowledged.
864
PALAEONTOLOGY, VOLUME 18
REFERENCES
DOWNiE, c. and sarjeant, w. a. s. 1964. Bibliography and index of fossil dinoflagellates and acritarchs.
Mem. Geol. Soc. Amer. 94, 1-180.
EviTT, w. R. 1967. Dinoflagellate studies IE The archeopyle. Stanford Univ. Publ. Geol. Sci. 10, No. 3, 1-82.
1969. Dinoflagellates and other organisms in palynological preparations. In tchudy, r. h. and
SCOTT, r. a. (eds.). Aspects of Palynology. Wiley-Interscience, New York, 439-479.
1970. Dinoflagellates— a selective review. Geoscience and Man, 1, 29-45.
FELIX, c. J. 1975. Palynological evidence for Triassic sediments in Ellet Ringnes Island, Arctic Canada.
Rev. Palaeobot. Palynol. 20, 109-117.
FISHER, M. J. 1972fl. Rhaeto-Liassic palynomorphs from the Barnstone railway cutting, Nottinghamshire.
Mercian Geologist, 4, 101-106.
19726. The Triassic palynofloral succession in England. Geoscience and Man, 4, 101-109.
GITMEZ, G. u. and sarjeant, w. a. s. 1972. Dinoflagellate cysts and acritarchs from Kimmeridgian (Upper
Jurassic) of England, Scotland and France. Bull. Br. Mas. nat. Hist. (Geol.), 21, 171-257.
LANJOUW, J. et al. 1966. International Code of Botanical Nomenclature. International Bureau for Plant
Taxonomy and Nomenclature, Utrecht, 1-402.
LENTIN, J. K. and Vv^iLLiAMS, G. L. 1973. Fossil dinoflagellates; index to genera and species. Geol. Surv.
Canada, Paper 73-42, 1-176.
LOEBLiCH, A. R., Jun. and loeblich, a. r. III. 1968. Index to the genera, subgenera, and sections of the
Pyrrhophyta, II. J. Paleont. 42, 210-213.
morbey, s. j. 1975. The palynostratigraphy of the Rhaetian Stage, Upper Triassic in the Kendelbach-
graben, Austria. Palaeontographica, B, 152, 1-75.
and NEVES, R. 1974. A scheme of palynologically defined Concurrent-range Zones and Subzones for
the Triassic Rhaetian Stage (sensu lato). Rev. Palaeobot. Palynol. 17, 161-173.
ORBELL, G. 1973. Palynology of the British Rhaeto-Liassic. Bull. Geol. Surv. G.B. 44, 1-44.
SARJEANT, w. A. s. 1963. Fossil dinoflagellates from Upper Triassic sediments. Nature, Land. 199, 353-354.
1966. Dinoflagellate cysts with Gonyaulax-iype, tabulation. In davey, r. j. et al. Studies on Mesozoic
and Cainozoic dinoflagellate cysts. Bull. Br. Mus. nat. Hist. (Geol.), Suppl. 3, 107-156.
1969. Taxonomic changes proposed by W. A. S. Sarjeant. In davey, r. j. et al. Appendix to 'Studies
on Mesozoic and Cainozoic dinoflagellate cysts’. Ibid. Appendix to Suppl. 3, 7-15.
1969. Microfossils other than pollen and spores in palynological preparations. In erdtman, g. Hand-
book of Palynology. Hafner Publishing Co., New York, 165-208.
SMITH, D. G., HARLAND, w. B. and HUGHES, N. F. 1975. Geology of Hopen, Svalbard. Geol. Mag. 112, 1-23.
WALL, D. and DALE, B. 1 968. Modern dinoflagellate cysts and evolution of the Peridiniales. Micropaleontology,
14, 265-304.
and HARADA, K. 1973. Descriptions of new fossil dinoflagellates from the late Quaternary of the
Black Sea. Ibid. 19, 18-31.
WARRINGTON, G. 1974. Studies in the palynological biostratigraphy of the British Trias. I. Reference sections
in west Lancashire and north Somerset. Rev. Palaeobot. Palynol. 17, 133-147.
and HARLAND, R. (in press). Palynology of the Trias and Lower Lias of the Larne Borehole. In
MANNING, p. I. and WILSON, H. E. The stratigraphy of the Larne Borehole, Co. Antrim. Bull. Geol. Surv.
WIGGINS, V. D. 1973. Upper Triassic dinoflagellates from arctic Alaska. Micropaleontology, 19, 1-17.
G.B. 50.
REX HARLAND
Institute of Geological Sciences
Ring Road, Halton
Leeds, LSI 5 8TQ
S. JACK MORBEY
Robertson Research International Ltd.
‘Ty’n-y-Coed’
Llanrhos
Llandudno, North Wales
Original typescript received 14 October 1974
Revised typescript received 16 December 1974
WILLIAM A. S. SARJEANT
Department of Geological Sciences
University of Saskatchewan
Saskatoon, Saskatchewan, Canada
THE PRODUCTION OF FAUNAL LISTS
BY AUTOMATIC METHODS
by IAN E. PENN
Abstract. Computer programs have been developed in the Institute of Geological Sciences which, after eliminating
any unwanted data from original determinations, generate correctly type-set and punctuated faunal lists. These are
suitable for direct publication or for easy incorporation into accounts dealing with wider geological topics.
Lists of species found are a simple and most fundamental means of recording
faunal (or floral) distribution and, since the days of William Smith and his recognition
of strata distinguished by means of fossil content, they have occupied a place of special
importance in geology. After the palaeontologist has recorded his observations, he
may still spend a considerable amount of time in non-palaeontological work when he
communicates his discoveries, even by a means apparently as simple as a faunal list.
Thus the fossil names must be written, typed, and checked (often in several copies).
This process may be repeated more than once when, in a large organization, the
palaeontologist’s report is incorporated in a larger work such as a geological account
or memoir written by a colleague, which itself must be typed and checked (often in
several copies). Finally, the lists must again be checked at least once on return from
the printers, prior to eventual publication. When for any reason some part of the
data has to be published additionally or even separately, the whole process may need
to be repeated.
Within the Institute of Geological Sciences, analogous problems had been
encountered in the production of the relatively more sophisticated stratigraphical
range-diagram, and a package of computer programs was written to eliminate non-
palaeontological activity as far as possible (Penn 1974; Farmer and Johnson 1975,
in press). It was then decided to generate fossil lists suitable for direct publication in
the same manner as the range-diagrams and to incorporate a program which would
simultaneously eliminate unwanted data. The main features of these programs are
here outlined (text-fig. 1), while full program listings may be obtained on request.
The programs are housed on the Institute’s IBM 1130 computer configuration and
also on the Edinburgh Regional Computing Centre’s PDP- 11/45 configuration.
They may be installed and used to produce ‘crude’ line-printer output by anyone
having access to such machines. Production of type-set lists requires access, however,
to a more specialized instrument such as the Institute’s Addressograph-Multigraph
AM -747. The species dictionaries contain the names used by individual palaeon-
tologists within the Institute, and it is intended that these will be subsequently
integrated, as will the data generated by them. It is envisaged that users outside I.G.S.
could construct similar dictionaries and data files which may subsequently be brought
together. The Institute’s facilities are, however, available at the discretion of the
Director.
[Palaeontology, Vol. 18, Part 4, 1975, pp. 865-869.]
866
PALAEONTOLOGY, VOLUME 18
PROGRAM INPUT, CONTENTS AND OUTPUT
The fossil data (F DATA in text-fig. 1) are presented on punch cards in stratigraphical
order such that a preliminary card lists the number of batches in the stratigraphic
section being described. Then the first card in each batch states the number of species
found as well as the measured stratal range (e.g. depth range in a borehole of the
sampled horizon), while each subsequent card records the species code number
(corresponding to the full fossil name stored in a companion dictionary) and a code
denoting species abundance at that horizon (see Penn 1974). At the present embryonic
stage of this data bank, each such stratigraphical section is identified manually.
Program CUF 10 reads these data and stores them on magnetic discs ready for
accession by remaining programs. But program RCVET is firstly presented with
a list of the code numbers of those species which it is desired to eliminate (or retain,
as the case may be) from the main body of the data. Only these selected species, with
their abundance codes, are read and stored by program RCVET. Indication is given
if such selection results in the elimination of all species from the data. Thus in the
example shown (text-fig. 2), the Bivalves have first been selected from the total data
and listed, followed by listing of the remaining species.
Preliminary inspection of the data may be made by printing a list of the encoded
TEXT-FIG. 1. Flow chart of the various programs. F DATA represents the input of fossil data.
The various identifiers within rectangles represent the various program decks. PREPT produces
paper-tape and line-printer output.
PENN: AUTOMATIC FAUNAL LISTS
867
data, and a list of all the species found in the total data, by using programs CUF 1 1
and CUZ 88 respectively (Penn 1974). Program PREPT, however, translates the
encoded data and, after consulting the species dictionary, punches out a paper-tape
of the full fossil name preceded by an indication of abundance for every determination
made. This tape, which contains type-setting instructions obtained from the species
dictionary, is fed into a phototypesetter to produce a correctly type-set and punctuated
list (text-fig. 2) for each sampling horizon. Provision has been made for an addition
and multiplication factor to be incorporated into the depth-range values, so as to
SPECIMEN DATA. ALL MACROFOSSILS SPECIMEN DATA. BIVALVES
SPECIMEN DATA. NON BIVALVES
SAMPLING HORIZON I 35,55 to 36,23
p wood [frag.]
fc RhynchoneUoidella smiihi (Davidson)
p Anisocardia bathemis Cox
fc bivalve [indet.]
p Caiinula cf. ampulla (d’Archiac)
p Eniolium sp.
p Grammatodon bathonicus Cox & Arkell
p burrow [horizontal and straight]
SAMPLING HORIZON 2 36.23 to 37.19
p bivalve [indet ]
p Modiolus sp.
p belemnite [indet.]
SAMPLING HORIZON 3 37.37to 38.12
p serpulid [indet.]
Ic rhynchonellacean [indet ]
fc RhynchoneUoidella smithi (Davidson)
p RhynchoneUoidella waitonensis Muir-Wood
fc RhynchoneUoidella sp.
fc arcacean [indet ]
c bivalve [indet ]
fc Caiinula cf. ampulla (d’Archiac)
p Chlamys ( Radulopecten ) sp.
p Gervillella sp.
p pectinoid [mdet ]
SAMPLING HORIZON 4 38.12 to 38 39
p RhynchoneUoidella sp.
p bivalve [indet ]
fc Eniolium sp.
p Gervillella sp.
SAMPLING HORIZON 5 38,39 to 38.60
SAMPLING HORIZON 1 35.55 to 36.23
p Anisocardia baihensis Cox
fc bivalve [indet.]
p Cannula d. ampulla {6' fiixchizc)
p Eniolium sp.
p Grammatodon bathonicus Cox & Arkell
SAMPLING HORIZON 2 36.23to 37 19
p bivalve [indet.]
p Modiolus sp.
SAMPLING HORIZON 3 37.37 to 38.12
fc arcacean [indet.]
c bivalve [indet.]
fc Caiinula cf. ampulla (d’Archiac)
p Chlamys (Radulopecten) sp.
p Gervillella sp.
p pectinoid [indet ]
SAMPLING HORIZON 4 38.l2to 38.39
p bivalve [indet ]
fc Eniolium sp.
p Gervillella sp.
SAMPLING HORIZON 5 38.39 to 38.60
fc bivalve [indet ]
? Campionectes sp.
fc Eniolium sp.
p Inoperna plicata (J . Sowerby)
p Liostrea sp.
fc Modiolus anatinus Wm. Smith
p Modiolus sp.
p Vaugonia sp.
SAMPLING HORIZON 1 35.55 to 36.23
p wood [frag ]
fc RhynchoneUoidella smithi ( Davidson)
p burrow [horizontal and straight]
SAMPLING HORIZON 2 36,23 to 37 19
p belemnite [indet ]
SAMPLING HORIZON 3 37.37 to 38 12
p serpulid [indet ]
fc rhynchonellacean [indet ]
fc RhynchoneUoidella smithi (Davidson)
p RhynchoneUoidella waitonensis Muir-Wood
fc RhynchoneUoidella sp.
SAMPLING HORIZON 4 38.l2to 38,39
p RhynchoneUoidella sp.
SAMPLING HORIZON 5 38,39 to 38.60
p Sarcinella socialis (Goldfuss)
fc serpulid [indet ]
fc RhynchoneUoidella sp.
p Ornithella balhonica (RoWier)
p Proceriihium sp.
p Sarcinella socialis (Goldfuss)
fc serpulid [indet.]
fc RhynchoneUoidella sp.
p Ornithella (Rollier)
p Proceriihium sp.
fc bivalve [indet ]
? Campionectes sp.
fc Eniolium sp.
p Inoperna plicata (J . Sowerby)
p Liostrea sp.
fc Modiolus anatinus Smith
p Modiolus sp.
p Vaugonia sp.
TEXT-FIG. 2. Specimen output from a Middle Jurassic borehole near Bath. ? = possibly occurring; p =
present; fc = fairly common; c = common. The total data on the left of the diagram has been split into the
Bivalve and non-Bivalve sections shown on the right-hand side.
868
PALAEONTOLOGY, VOLUME 18
eliminate the punching of unnecessary digits where, for example, closely spaced
samples have been taken from considerable depth in a borehole. Such factors may
also be used as a security ‘link’ on the depth range of the sample in the case of con-
fidential material.
PROGRAM PERFORMANCE
The size limits for the data are at present 100 (species) x 100 (horizons) and the
number of species at any one horizon must be limited to 50 as in the range-chart
program (Penn 1974). Data preparation (including the noting of the preliminary raw
observations) and checking for the computer takes about the same time as preparing
a first draft of a clean-copy manuscript for typing. Typing the first copy, however,
takes about as long as drafting the manuscript, and checking the first and each
subsequent typescript is conservatively estimated at 50% of the time taken to prepare
the first manuscript. Typing of subsequent copies is slightly quicker (perhaps by
about 25%). Thus, after the first draft of the manuscript, the time spent by the
palaeontologist in non-palaeontological work increases by 50% of his original time,
and similarly that of the typist by 75% for each successive ‘round’ of typing. The
final ‘round’ is that done by the printer and is estimated to be 120% of the time taken
by an ordinary typist. Thus a manuscript taking 1 hr for the preparation of the first
draft of a clean-copy manuscript would take (assuming two further drafts) 2 hr 45 min
to submission and 1 hr 20 min work by the printer as against 1 hr for punching and
checking and 4 min computer time. Program RCVET, which selects data during the
loading process, performs basically the same functions as the normal loading pro-
gram and, indeed by setting it so that no selection is made, may be used as a substitute
for program CUF 10. The operational time taken by program PREPT is dominated
by the number of species determinations in the data. Since most of the activity is in
punching paper-tape, the time taken to produce the output is almost entirely depen-
dent on the size of machine, the speed, and the arrangement of output devices. Thus
the IBM 1 130 configuration, on which the program was established, does not con-
veniently allow separate operation of the central processor and paper-tape punch.
The speed of the operation was therefore determined by the speed of the punch
which, at 14-8 characters per sec, is very slow. Operation on a larger computer
(a PDP- 11/45) with a faster paper-tape improved the run times, in a conservative
estimate, by a factor of five times. In practice this means a job of around 250 deter-
minations is run in around 5 min. In fact, in a multi-user environment, output devices
would be operated simultaneously with entirely different operations, meaning that
the time spent producing the fossil list would be almost negligible. The advantage is
even more marked if the palaeontologist needs an accompanying range-chart, since
the same data input is used, and producing the fossil list simply involves a small
amount of extra computer time.
It is thus possible to free the palaeontologist from a very considerable proportion
of non-palaeontological work once he has initially recorded and checked his data.
In addition, his data can be stored in computer-processable form ready for the per-
formance of other analytical techniques and, in the long term, ready for incorporation
into a computerized data bank.
PENN. AUTOMATIC FAUNAL LISTS
869
Acknowledgements. The programs were written by Dr. D. G. Farmer (Computer Unit, I.G.S.) and Dr.
T. J. Dhonau (Editorial and Publication Section, I.G.S.) advised on the use of the phototypesetter. The
paper is published by permission of the Director of the Institute of Geological Sciences.
REFERENCES
FARMER, D. G. and JOHNSON, L. 1975. Rep. Inst. geol. Sci. (In press.)
PENN, I. E. 1974. The production of stratigraphical range-diagrams by automatic methods. Palaeontology,
17, 553-563.
Typescript received 23 December 1974
Revised typescript received 24 Eebruary 1975
IAN E. PENN
Department of Palaeontology
Institute of Geological Sciences
Exhibition Road
London, SW7 2DE
THE LECTOTYPE OF THE AMMONITE
CADOMITES PSILACANTHUS (WERMBTER)
by G. E. G. WESTERMANN and M. RIOULT
Abstract. The inflated "Am. humphriesianus’’ d’Orbigny (1845, pi. 134, figs. 1-4), non J. Sowerby (1825), has sub-
sequently been named three times; the oldest objective synonym is Stephanoceras psilacanthus Wermbter (1891).
The lectotype has been found in the British Museum (Natural History) and is redescribed. It probably came from the
boundary Humphriesianum-Subfurcatum Zones at Sully, near Bayeux, Normandy. Specimens from the Parkinsoni
Zone described under Cadomites arkelli Sturani (1964), the youngest objective synonym, are C. psilacanthus sturanii
subsp. nov.
The type specimen of 'Ammonites Humphriesianus, Sow.’ figured by d’Orbigny
(1845) on plate 134, figs. 1-2 {non Am. humphriesianus Sowerby, 1825) has now been
found in the Tesson Collection of the British Museum (Natural History). Tesson,
mentioned by d’Orbigny (1845, p. 400) as one of the collectors of the ‘species’, was
a teacher at the Lycee of Caen, a naturalist and friend of Professor J. A. Eudes-
Deslongchamps. After his death, his fossil collections from all regions of Normandy
were purchased by the British Museum (1857). Separate examinations by us in
1972-1973 of specimen No. 37309 labelled 'Ammonites (Stephanoeeras) Brodioei,
J. Sow., Inferior oolite, Bajocian, Bayeux. Figd. as Amm. Humphriesianus, by
A. d’Orbigny, Pal. Fr. Terr, jurass. Vol. I, p. 398 (1846), pi. 134, figs. 1-2’ confirmed
beyond doubt the identity with d’Orbigny’s illustration as labelled.
Both of us had searched independently for this type specimen in the Museum
National d’Histoire Naturelle, Paris, where A. d’Orbigny apparently stored most of
his collections. Following the conviction of most authorities that in d’Orbigny’s
figures, characters of several specimens were united into one, the inability to match
specimens with illustrations seemed to confirm the artistic licence of the engraver,
rather than their absence from the collections. However, several specimens from the
Tesson Collection have been shown beyond reasonable doubt to be the originals to
A. d’Orbigny’s illustrations (e.g. B.M. 37325, lectotype of Wagnericeras wagneri
(Oppel) = Ammonites planula d’Orbigny, pi. 144, non Hehl in Zieten 1830, illustrated
by Arkell 1958, text-fig. 65), and we hope that this new find of a complete specimen
closely matching d’Orbigny’s figures will lead to a thorough search for other type
specimens.
Moreover, this is also an example of the nomenclatural confusion arising from the
designation of old illustrations in old monographs as the holotype or even syntypes
of new species, rather than specimens ; particularly if this is done repeatedly for the
same illustrations.
[Palaeontology, Vol. 18, Part 4, 1975, pp. 871-877, pi. 105.]
872
PALAEONTOLOGY, VOLUME 18
Family stephanoceratidae Neumayr, 1875
Subfamily cadomitinae Westermann, 1956
Genus cadomites Munier-Chalmas, 1892
Cadomites psilacanthus (Wermbter, 1891)
Plate 105
vl845 Ammonites Humphriesianus, Sowerby; d’Orbigny, pp. 398-400, pi. 134, figs. 1-2 [lectotype],
?3-4 [nucleus or juv.]. 1852, d’Orbigny, 11(2), p. 489, fig. 430 [reproduction of d’Orbigny
1845, pi. 134, xO-5].
*1891 Stephanoceras psilacanthus Behr. ms. (= Humphriesianum d’Orb., p.p., pi. 134, non 133);
Wermbter, pp. 270-271 [citation of types only, not his specimens].
1895 Coeloceras cosmopoliticum = Coeloceras Humphriesianum d’Orb. (non Sow.); Moricke,
pp. 20, 21 [for d’Orbigny, pi. 134, figs. 1-2]. 71923, Cadomites cosmopoliticum Moericke;
Fallot and Blanchet, p. 151, pi. XI, fig. 10. Non 1937, Gillet, p. 82, pi. 5, fig. 8 [fide Sturani
1966]. 71939, Roche, pp. 200-201.
1964 Cadomites n. sp. (=' Ammonites humphriesianus' (non Sow.) d’Orbigny 1842-1849, t. 134,
ff. 1-2; = Cadomites cosmopoliticus (non Moricke) Fallot and Blanchet 1923, t. 11, f. 10);
Sturani (1964u), p. 37, pi. 6, fig. 6 [ = C. p. sturanii subsp. nov.].
1964 Cadomites arkelli n. sp. ; Sturani (1964(t), p. 20 [text-fig. 20, pi. 2, fig. 5 = C. p. sturanii subsp.
nov., holotype].
1966 Cadomites psilacanthus (Wermbter); Sturani, p. 27 [correction for Sturani 19647); text-
fig. 2 = 1C. p. sturanii subsp. nov.].
71974 Cadomites (Cadomites) psilacanthus (Wermbter, 1891); Kopik, p. 13, pi. 1, fig. la, b.
History
A. d’Orbigny (1845, pp. 398-400, pis. 133-135) described and illustrated several
specimens under Ammonites Humphriesianus Sow. none of which belongs to the true
species of J. de C. Sowerby (1825, pi. 500, fig. 1 centre; S. S. Buckman 1908, pi. VII,
fig. \a, b). In the text, d’Orbigny mixed characters now attributed to several genera
and/or subgenera; according to text and plates {excl. pathological specimen, pi. 135,
fig. 2), he distinguished (1) a serpenticone variety with subcircular whorl section
(pi. 133) which became the holotype of Stephanoceras (Skirroceras) bayleanum
(Oppel 1856-1858, p. 377), and (2) an involute inflated variety with depressed whorl
section (pi. 134) which Oppel (1856-1858, p. 376) regarded as the true Am. hum-
phriesianus [= Stephanoceras s.s.], but Wermbter (1891) separated as S. psilacanthus.
No details on d’Orbigny’s locality are available and Wermbter did not explain why
he distinguished his species; in fact, Wermbter’s faunal lists and field experience in
the Weser Mountains, Lower Saxony, of one of us (G. E. G. W.), permit the con-
clusion that the German specimens were Stephanoceras s.s. or S. (Stemmatoceras)
EXPLANATION OF PLATE 105
Cadomites psilacanthus (Wermbter), lectotype; type specimen to 'Ammonites Humphriesianus, Sowerby’
of d’Orbigny (1845, plate 134, figs. 1-2). Boundary of Humphriesianum and Subfurcatum Zones, near
Bayeux, Normandy. Tesson Collection No. 37309, British Museum (Natural History). 1, left side with
complete peristome and damaged shell wall; 2, apertural view; 3, right side with damaged peristome,
complete shell wall, and relatively retarded uncoiling of the body chamber; 4, ventral view. All figs, at
xO-9.
PLATE 105
WESTERMANN and RIOULT, Cadomites psilacanthus
874
PALAEONTOLOGY, VOLUME 18
from the Humphriesianum and possibly also from the Sauzei Zone, i.e. generically
distinct from the type specimens. However, the designated plate 134 of d’Orbigny
illustrates two specimens, one (figs. 1-2) large and complete, the other (figs. 3-4) very
small and septate; they are thus syntypes, although all authors quite obviously had
the large complete specimen in mind which was finally designated as the lectotype
by Sturani (1966).
Apparently without knowledge of Wermbter’s work, W. Moricke (1895) desig-
nated the same large specimen of d’Orbigny’s plate 134 as the type of Coeloceras
cosmopoliticum (in heading p. 20 and text p. 21), this time in conjunction with Chilean
stephanoceratids ; even if the specimen figured by Steinmann (1881) referred to in
the synonymy is taxonomically different— he did not himself describe any specimens
—the case is similar to that of C. psilacanthus, so that Moricke’s name is a junior
objective synonym (cf. Sturani 1966). Because of the nomenclatural procedure,
Sturani (1964u, b) considered Moricke’s name a nomen nudum and, not knowing of
Wermbter’s previous naming, introduced the new (third) name, Cadomites arkelli,
for the same figure in d’Orbigny. In correcting his error, Sturani (1966) formally
designated the complete large specimen of d’Orbigny’s figs. 1-2 as the lectotype. His
own material, mainly from the Parkinson! Zone of the Venetian Alps, however,
differs from the designated holotype and is here distinguished at the subspecific level.
The lectotype
Specimen No. 37309 of the Tesson Collection, British Museum (Natural History),
is a complete macroconch with 1 10 mm diameter and most of the test and aperture
preserved, from the ‘Inferior oolite, Bajocian, of Bayeux’ (according to label).
The outer phragmocone whorls terminating at 75 mm diameter, are moderately
involute and depressed subovate in section with steep umbilical wall, convex flanks
(no lateral/umbilical shoulder), and a more gently convex venter. The ornament
consists of sharp fiexuous ribbing, with moderately spaced primaries and dense
secondaries, and mid-lateral pointed to slightly elongate tubercles; the subradiate
primaries are adaperturally concave and very prominent ; the much finer secondaries
are almost four times as numerous and cross the venter with slight convexity. The
umbilical seam runs along the line of tubercles. The test is complete on the phragmo-
cone so that the septal suture is not exposed (d’Orbigny’s plate 135, fig. 1 is not of this
specimen). The body chamber, three-fifths whorls (225°) in length, becomes rapidly
more evolute with the overlap decreasing from about two-fifths to one-third, resulting
in moderate ‘elliptical’ coiling, and slightly contracted towards the end; there is
marked negative allometry for both whorl height and breadth (see measurements).
The ornament becomes increasingly prominent, particularly on the venter. The con-
cave primaries lengthen at the end of the phragmocone and at the beginning of the
body chamber, so that the tubercles move from mid-lateral outward to approximately
three-fifths whorl height ; but at the end of the body chamber, tuberculation is again
mid-lateral. Parallel growth lines appear between the distant primaries. Slight
asymmetry is present at the beginning of the body chamber with the egression of
the umbilical seam being relatively retarded on the right side. Noteworthy is the strong
reduction of relief on the internal mould of the left flank of the body chamber where
the sharp ribbing of the shell becomes blunt.
WESTERMANN AND RIOULT: CADOMITES PSILACANTHUS 875
The peristome begins with a prominent slightly prorsiradiate collar parallel to the
ribs and growth lines, and continues with a smooth sinuous margin covered by growth
lines which indicate a broad umbo-lateral sinus and a straight ventral lip.
Measurements in mm on ribs/tubercles
Whorl
Diameter height
Peristome 110-3 40
End body chamber 105-5 35-5 (0-33)
Beginning body 81-5 33 (0-40)
chamber
Locality and age
The sediment in the body chamber is a grey micritic limestone with small shiny
ferruginous ooids. This lithofacies is well exposed in the ancient quarry of Sully,
a few kilometres north of Bayeux, at the base of the ‘Oolithe ferrugineuse de Bayeux’ ;
i.e. the boundary of the condensed Humphriesianum and Subfurcatum Zones (lower
or middle/upper Bajocian) (Rioult 1964, layers 3a-3b). The lectotype originated
either here or in any of several other quarries of the immediate vicinity which have
long been filled in.
D'Orbigny's illustration
Figs. 1 and 2 on plate 134 of d’Orbigny (1845) show good likeness to the actual
specimen and are thus not synthesograms. It appears, however, that the lithographer,
J. Delarue, engraved the more complete right side (marked by relatively retarded
egression of the umbilical seam) and reversed the sides in the printing process; while
the aperture is from the left side. D’Orbigny’s figures differ from the original also as
follows: (1) 10% reduction; (2) narrower and less depressed whorl section, parti-
cularly at the beginning of the body chamber (B/H = 1-34 v. 1-48) which thus appears
to grow isometrically rather than negatively allometric; (3) section of aperture too
narrowly curved laterally; (4) less flexuous ribbing particularly at the base of the
secondaries; (5) more distant primaries on the inner whorls; and (6) most tubercles
pointed rather than somewhat elongate.
The paralectotypes
The original to figs. 3 and 4 of d’Orbigny’s plate 134 is neither in the Tesson nor in
the d’Orbigny Collections. The illustrations, said to be natural size, show a small
septate specimen of 22 mm diameter which closely resembles the nucleus of the lecto-
type (for which the primaries were illustrated too widely spaced) and appears to be
a juvenile, nucleus, or microconchiate phragmocone of the same or closely allied
species. The septum is shown to have two (paired) subequal saddle axes indicating
two saddles of the internal suture, typical for Cadomites (see Westermann 1956). The
third original collection referred to by d’Orbigny (1845, p. 400) was made by Deslong-
champs and kept at the University of Caen; it was destroyed by fire and bombing in
1944.
Umbilical
Primaries/
Secondaries/
Breadth
diameter
whorl
whorl
56
44
33
127
50-6 (0-47)
41 (0-38)
32
48-7 (0-59)
28 (0-34)
876
PALAEONTOLOGY, VOLUME 18
Taxonomic remarks and comparison
In spite of appreciable efforts, no topotypes of C. psilacanthus have been discovered.
It appears that in general the genus Cadomites becomes more abundant only in the
uppermost Bajocian, Parkinsoni Zone.
C. psilacanthus is morphologically transitional between Stephanoceras and Cado-
mites combining the relatively short primaries of the former with the body chamber
coiling and sharpness and density of ribbing of the latter; the primaries, however,
are still widely spaced as in Stephanoceras and the body chamber does not contract
significantly as in Cadomites. However, the septum of the paralectotype (above) and
the stratigraphic range to the top of the Bajocian indicate strong affinity to Cadomites.
The specimens, mainly from the Parkinsoni Zone of the Alps described under
C. psilacanthus or ‘C. arkelli by Sturani (1964«, b, 1966) are here distinguished as
a new subspecies of C. psilacanthus.
The Andean 'Coeloceras cosmopoliticum" Moricke (1895) from northern Chile has
been illustrated only in a single specimen by Steinmann (1881, p. 268, pi. XII, fig. 7).
It has the curved primaries and round tubercles of C. psilacanthus but differs in the
circular whorl section and the more distant primaries on the inner whorls. Current
reinvestigation by one of us (G. E. G. W.) of the Caracoles fauna suggests closer
affiliation to Stephanoceras s.s. than to Cadomites. There is close resemblance to
C. deslongchampsi (Defrance in d’Orbigny, nom. eorr.) from the Parkinsoni and
Zigzag Zones and the closely allied C. homalogaster Buckman (1925, pi. 543) from
the ‘Leptosphinctes hemera’ (= Subfurcatum Zone), except for the shorter primaries
and the absence of a lateral shoulder in C. psilaeanthus. The somewhat similar
probable Cadomites described by Roche (1939, pis. 2 and 3) from the Humphriesi-
anum-Subfurcatum Zones of eastern France (C. humphriesiformis, C. perplicatus,
C. ? lissajousi) are all much more compressed than C. psilacanthus.
Cadomites psilaeanthus sturanii subsp. nov.
(Synonymy see under C. psilacanthus.)
Diagnosis. More rounded (narrower) whorls and less flexed ribs than in C. psila-
eanthus s.s.
Holotype. Cadomites arkelli Sturani, \%Ah, p. 20, text-fig. 20, pi. 2, fig. 5, from the Parkinsoni Zone of
Cava Magnavacca, Venetian Alps (for nomenclature of C. arkelli see above).
Age. This subspecies occurs at a higher level in the upper Bajocian (Parkinsoni Zone) than C. psilacanthus s.s.
Acknowledgements. We thank Dr. M. K. Howarth and Mr. D. Philips of the British Museum (Natural
History) for their kind co-operation during our visits and for furnishing some of the photographs.
REFERENCES
ARKELL, w. j. 1951-1958. Monograph of English Bathonian ammonites, parts 1-8. Palaeonlogr. Soc.
[Monogr.]. 1-264, 33 pis.
BUCKMAN, s. s. 1908. Illustrations of type specimens of the Inferior Oolite ammonites in the Sowerby
collection. Ibid. pis. I-VII with explanations.
1909-1930. (Yorkshire) Type ammonites, parts 1-7. London, text and 790 pis.
FALLOT, p. and BLANCHET, F. 1923. Observations sur la faune des terrains jurassiques dans la region de
Cardo et de Tortosa (Province de Tarragone). Treh. Inst. Catal. Hist. nat. 1921-1922, 71-260, 13 pis.
WESTERMANN AND RIOULT: CADOMITES PSILACANTHUS
877
GILLET, s. 1937. Les ammonites du Bajocien d’Alsace et de Lorraine. Serv. Carte geol. Alsace-Lorraine,
Mem. 5, 130 pp., 5 pis.
KOPiK, J. 1974. Genus Cadomites Munier-Chalmas, 1892 (Ammonitina) in the Upper Bajocian and
Bathonian of the Cracow-Wielun Jurassic Range and the Gory Swietokrzyskie Mountains (southern
Poland). Inst. Geol., Bull. 276, 7-53, pis. I-XI.
MORiCKE, w. 1895. Beitrage zur Geologic und Palaeontologie von Stidamerika. II, Die Versteinerungen des
Lias und Unteroolith von Chile. Neues Jb. Miner. Geol. Paldont. Beil.-Bd. 9, 1-100, pis. I-VI.
MUNIER-CHALMAS, E. c. p. A. 1892. Sur la possibilitc d’admettre un dimorphism sexuel chez les Ammoni-
tides. C.r. Soc. geol. Fr. ser. 3, 20, 170-174.
NEUMAYR, M. 1875. Die Ammoniten der Kreide und die Systematik der Ammonitiden. Z. dt. geol. Ges. 27,
854-892.
OPPEL, A. 1856-1858. Die Juraformation Englands, Frankreichs und des sudwestlichen Deutschlands.
Jh. Ver. vaterl. Naturk. Wiirtt. XII, 221-556; XIII, 141-396; XIV, 129-291.
ORBiGNY, A. d’. 1842-1849. Paleontologie Frangaise, Terrains jurassiques, Cepbalopodes. Paris (Masson),
text 1-642, Atlas pis. 1-234.
1850-1852. Cours elementaire de paleontologie et de geologie stratigraphique. V. Masson ed., Paris,
1-847, 628 figs.
RIOULT, M. 1964. Le stratotype du Bajocien. In CoU. Jurass. Luxembourg 1962, Compt. Rend. Mem. Inst.
grand-ducal, sect. sci. nat. phys. math. Luxembourg, 239-263.
ROCHE, p. 1939. Aalenien et Bajocien du Maconnais et de quelques regions voisines. Trav. Lab. Geol. Univ.
Lyon, 35, Mem. 29, 1-355, pis. I-XIII.
SOWERBY, J. DE c. 1822-1846. Mineral Conchology. Pis. 338-648.
STEiNMANN, G. 1881. Zur Kenntniss der Jura- und Kreideformation von Caracoles (Bolivia). Neues Jh.
Miner. Geol. Paldont. Beil.-Bd. 1, 239-301, pis. I-XVII.
STURANi, c. 1964fl. La successione delle faune ad ammoniti nelle formazioni mediogiurassiche delle Prealpi
Veneto occidentale (region! tra il Lago di Garda e la valle del Brenta). 1st. Geol. Miner. Univ. Padova,
Mem. 24, 1-64, pis. I-VI.
19646. Ammoniti mediogiurassiche del Veneto, faune del Baiociano terminale (Zone a Garantiana
e a Parkinsoni). Ibid. 1-43, pis. I-IV.
1966. Ammonites and stratigraphy of the Bathonian in the Digne-Barreme area. Boll. Soc. paleont.
ital. 5, 3-57, pis. 1-24.
WERMBTER, H. 1891. Der Gebirgsbau des Leinethales zwischen Greene und Banteln. Neues Jh. Miner. Geol.
Paldont. Beil.-Bd. 7, 246-294, pis. IV-V.
WESTERMANN, G. E. G. 1956. Phylogenie der Stephanocerataceae und Perisphinctaceae des Dogger. Neues.
Jb. Geol. Paldont. Abh. 103, 233-279.
ziETEN, c. H. V. 1830-1834. Die Versteinerungen Wiirttemhergs, parts 1-12, 102 pp., 72 pis. Stuttgart.
G. E. G. WESTERMANN
Department of Geology
McMaster University
Hamilton, Ontario L8S 4M1
Canada
M. RIOULT
Departement de Geologie
Universite de Caen
Esplanade de la Paix
14000-Caen, France
Typescript received 4 September 1974
Revised typescript received 6 February 1975
SHORT COMMUNICATIONS
TEREBRATULIDE AFFINITY OFTHE
BRACHIOPOD SPIRIFERA MINIMA MOORE
by P. G. BAKER and c. J. t. copp
Abstract. Investigation of seventy-three recently rediscovered specimens of Spiriferinal minima (Moore) enables
resolution of the problem of possible synonymy with Nannirhynchia longirostra Baker, 1971. Comparison of the
cardinal areas and delthyria of the two species enables the distinction between S. ? minima and N. longirostra to be
clearly demonstrated. Further, the characters of the shell show that 5.? minima (Moore) is a juvenile terebratulidine
assignable to Terehratula and that N. longirostra Baker is validly designated.
The micromorphic Spiriferinal minima has periodically attracted the attention of
palaeontologists (Davidson 1876; Buckman 1918; Ager 1967; Baker 1971) since the
first record (Moore 1861) of its occurrence in the Inferior Oolite of Dundry Hill near
Bristol. Unfortunately, the precise location from which Moore obtained his material
is not known. All investigation of the species has been hampered by the absence of
the holotype and the apparent lack of any syntypes or topotypes. It is particularly
gratifying, therefore, that a part of the Charles Moore Collection, recently rediscovered
in the Somerset County Museum, Taunton Castle, should include a box containing
seventy-three specimens labelled, ‘4462. Spirifera minima, Dundry’. No precise
horizon is given but the adherent matrix is identical with that of the accompanying
thecidellinids of undisputed Bajocian age. The material was presented to the museum
in 1905 by the Revd. H. H. Winwood, who had been in charge of the Moore Collec-
tion at Bath following Moore’s death in 1881. The part of the collection presented
to the Taunton Museum apparently consisted of specimens which were kept at
Moore’s house and, therefore, not sold with the main collection housed in the Bath
Literary and Scientific Institution. It is probable that Moore’s widow kept them and
later gave them to Winwood. This would account for the absence of S.l minima from
the Bath Museum when earlier workers wished to refer to it.
Davidson (1876, p. 103) tentatively proposed the name Spiriferinal for Spirifera
Moore, 1861, but did not formally designate the genus, observing ‘I know so little
of this minute fossil I cannot venture to express any opinion with respect to the genus
to which it belongs’. As S.l minima is certainly a juvenile terebratulidine and as there
is, at present, no way of assigning the species to an adult genus, the authors would
prefer formal designation to remain in abeyance. However, in view of the con-
siderable time which has elapsed since Moore’s description, and in view of the recurring
interest in S.l minima, it is considered that Moore’s original diagnosis merits repro-
duction and emendation.
[Palaeontology, Vol. 18, Part 4, 1975, pp. 879-882.]
880
PALAEONTOLOGY, VOLUME 18
"Terehratula' minima (Moore)
Text-fig. Ia-d; text-fig. 2
1861 Spirifera minima Moore, p. 190, pi. ii, figs. 19, 20.
1876 Spiriferinal minima (Moore) Davidson, p. 103, pi. XI, fig. 17.
1918 Nannirhynchial (Spiriferinal) minima (Moore) Buckman, p. 68.
1967 Nannirhynchial minima (y[ooxQ), Kgtr,p. 137.
Original diagnosis. Shell microscopic, often one sided or asymmetrical, slightly
rugose; valves moderately convex; deltidium triangular; area broad and flattened;
hinge line broad ; front of shell rounded. In some specimens the shell presents a uni-
formly flattened surface, whilst in the majority the outer surface of the smaller valve
possesses mesial folds and in the larger valve a central sinus (Moore).
Emended diagnosis. Minute, asymmetric " Ter ebr alula' \ planoconvex to ventribi-
convex, slightly longer than wide, characterized by low, poorly defined mesial folds
which fail to deflect the commissure. Apex of the delthyrium closed by a rudimentary
pedicle collar. Shell endopunctate.
Lectotype. There is very little evidence of a type specimen or specimens having been
used by Moore in his original description of the species in 1861. It is likely that,
adopting the procedure of many palaeontologists of the time, he established the
species on knowledge obtained from several examples which he considered to be
typical or characteristic forms. This belief is borne out by the comments ‘In some
specimens’ and ‘whilst in the majority’ in Moore’s original diagnosis. In addition,
figs. 19 and 20 (Moore 1861, pi. ii) are quite clearly prepared from different specimens.
His locality and stratigraphical details were vague.
Since the only known specimens are those in the Somerset County Museum,
No. 4462, the specimen figured in this paper (text-fig. 1a-d) is here proposed as a
lectotype.
TEXT-FIG. 1. A-D. Stereoscaii photomicrographs of the lectotype (No. 4462A) of 'Terehratula' minima
(Moore), showing the general morphology. Specimen coated with evaporated aluminium before photo-
graphy. A, brachial view showing the characteristic lateral deflection of the beak, x 35. b, lateral view, x 35.
c, anterior view, x 50. Specimen tilted forwards slightly, to show the poorly defined mesial folds, d, enlarged
view of the umbonal region, showing the rudimentary pedicle collar closing the apex of the delthyrium, X 85.
BAKER AND COPP: 'TEREBRATULA' MINIMA (MOORE) 881
Dimensions of lectotype. Length 1-2 mm, width 10 mm, thickness 0-4 mm.
Distribution. Uncertain. Moore (1861, p. 190) states that the species is not uncommon
in the Inferior Oolite of Dundry. The Somerset County Museum specimens are
simply labelled Dundry and it is presumed that they are topotypes. They are associated
with moorellinids of Murchisonae Zone age.
Description. External characters. A juvenile terebratulidine up to about T8 mm long,
1-6 mm wide, and 0-4 mm thick, often symmetrical but more commonly with a
marked lateral deflection of the beak. Typically planoconvex or ventribiconvex but
biconvex forms are seen. A characteristic feature is the flattening of the cardinal area
to form an interarea sensu lato, bounded by very sharp beak ridges. Small disjunct
deltidial plates are present, with their inner edges elevated above the plane of the
cardinal area. Specimens, in which the structure is not obscured by matrix, show
a rudimentary pedicle collar closing the apex of the delthyrium, undoubtedly repre-
senting the triagular ‘deltidium’ noted by both Moore (1861) and Davidson (1876).
Internal characters. Apart from immature hinge teeth and sockets, no other internal
characters have been noted.
Discussion. As noted earlier, attempts to study the species have been hampered by
the unavailability of a holotype. An important feature overlooked by Moore but
presumably noted by Davidson (hence Spiriferinal) is the endopunctate shell.
Although even Davidson (1876, p. 103) had to rely on drawing ‘one of Moore’s
specimens’. It appears that by 1918 even the topotypes had been mislaid or Buckman
(1918, p. 68) would surely have been able to differentiate between 'Terehratula
minima and Nannirhynchia subpygmaea Buckman (ex Walker MS.) particularly as
weathered specimens of ‘T.’ minima are so obviously endopunctate. In the absence
of actual specimens for study, subsequent workers (Ager 1967; Baker 1971) have
also been misled by the superficial resemblance between ‘T.’ minima and Nanni-
rhynchia Buckman. Of particular interest was the possibility of synonymy of ‘T.’
minima with N. longirostra Baker, 1971. The recently discovered specimens show
that ‘T.’ minima is much more dorso-ventrally compressed than N. longirostra.
Diagnostic differences are seen in the cardinal areas and delthyria of the two species
and in the observation that ‘T.’ minima is endopunctate whereas N. longirostra is
impunctate. The flat cardinal area and sharp beak ridges (text-fig. 2a, b) of ‘T.’ minima
are in sharp contrast with the rounded beak ridges and well-defined palintropes
(text-fig. 2c) of N. longirostra. A pedicle collar is characteristic of both species but
in N. longirostra this is an almost sessile structure (Baker 1971, pi. 135, fig. 10;
pi. 136, fig. 1) not usually visible externally. A further minor difference is that in the
brachial valve of ‘T.’ minima, a shallow sulcus develops on either side of the umbonal
region but in N. longirostra the brachial valve is regularly convex in this region.
Demonstration of the distinction between ‘T.’ minima and N. longirostra still leaves
the affinity of ‘T.’ minima for consideration. ‘T.’ minima displays all the characters
which are typical of juvenile cancellothyridids. If the cardinal area and delthyrial
characters are compared with an early juvenile cancellothyridid aflf. Plectothyris
(text-fig. 2d) from a different locality (Baker 1 97 1 , p. 696) they are found to correspond
in almost every detail, even to the rudimentary pedicle collar. The closeness of the
882
PALAEONTOLOGY, VOLUME 18
TEXT-FIG. 2. Detail of the morphology of the posterior portion of the shells of four specimens investigated.
All x25 magnification, a, b, small, A, and larger, b, specimens oVTerebratula' minima (Moore), c, Nanni-
rhynchia longirostra, holotype (Brit. Mus. BB.45820). D, early juvenile terebratulidine afif. Plectothyris.
resemblance must be genetic rather than coincidental and the conclusion drawn from
the study of the newly available material must be, therefore, that S.l minima (Moore)
is a juvenile of an undetermined terebratulidine brachiopod assignable to 'Tere-
bratula\
Acknowledgements. The authors wish to thank Dr. H. Torrens, Department of Geology, University of
Keele, for information leading to the rediscovery of the material and Dr. J. D. Hudson, Department
of Geology, University of Leicester, for comments on a previous version of the manuscript.
REFERENCES
AGER, D. v. 1967. The British Liassic Rhynchonellidae, Part IV. Palaeontogr. Soc. (Monogr.), London,
121, 137-172.
BAKER, p. G. 1971. A new micromorphic rhynchonellide brachiopod from the Middle Jurassic of England.
Palaeontology, 14, 696-703.
BUCKMAN, s. s. i 9 1 7 ( 1 9 1 8). The Brachiopoda of the Namyau Beds, Northern Shan States, Burma. Palaeont.
Indica, new ser. 3, 299 pp.
DAVIDSON, T. 1876. Supplement to the British Jurassic and Triassic Brachiopoda, British Fossil Brachio-
poda, pt. 4, 103, Suppl. PI. 11. Palaeontogr. Soc. (Monogr.), London.
MOORE, c. 1861. On new Brachiopoda and the development of the loop in Terebratella. Geologist, 4, 190-194.
P. G. BAKER
Division of Geology
Derby College of Art and Technology
Kedleston Road
Derby, DE3 1GB
C. J. T. COPP
Department of Geology
University of Keele
Keele, STS 5BG
Typescript received 12 March 1975
Revised typescript received 18 April 1975
NEW DATA ON TREMADOC GRAPTOLITES
FROM YUKON, CANADA
by D. E. JACKSON
Abstract. The Tremadocian subzones of Staurograptus tenuis, Anisograptus richardsoni, Clonograptus aureus, and
Adelograptus antiquus are given full zonal status in northern Richardson Mountains. Bryograptus ramosus Br0gger
and a species of Dictyonema are described from the upper Tremadoc. The occurrence of B. ramosus in the C. aureus
Zone supports a correlation with shales of 3ajS age in Norway.
In 1974 Jackson presented findings on the sequence of graptolites found in Trema-
docian shales on Peel River, Yukon Territory. In addition to simplifying zonal
terminology the paper proposed that the Adelograptus Zone and the Staurograptus
Zone each be divided into two subzones. This short paper offers important new data
from two river sections in the Richardson Mountains. The Rock River section which
lies 100 km north of Peel River demonstrates the usefulness of the Peel River zonal
scheme, and the Canyon Creek section 40 km north-west of the Upper Canyon on
Peel River provides supplementary data on the faunal content of the upper Tremadoc.
TEXT-FIG. la, b, d, Bryograptus ramosus Br0gger; a, d, GSC hypotypes 27000
and 27001 respectively, x 3; ft, GSC hypotype 27002, x 5; c, Dictyonema sp.
GSC hypotype 27003 (not described); e, Dictyonema cf. percancellatum
Ruedemann GSC hypotype 27004, x3',f, 1 Dendrograptus sp. GSC hypotype
27005 (not described), x 3. All figures are camera lucida drawings.
[Palaeontology, Vol. 18, Part 4, 1975, pp. 883-887.]
884
PALAEONTOLOGY, VOLUME 18
TABLE 1. Rock River Section (66° 48' N.; 136° 07' W.) measured
by geologists of Chevron Standard Ltd., in 1968; field designation ZB- 19.
“d
o
o
^ o n 4^ X n "s cd b n "s
P
(T
O
C) Oq
oq
II
D T3
S 5
C)
O5 Oq
t3 -T3
P 2 ::::
^ cd
I
si
§■ s
ts|
^ ^
III
S s 5
S-T3 ?
f“ -S' ®
2. ^ ~. •
^ K a ^
S P
== H H S'
• • C/5
^ c/3 g
X K
P CO
II
5 2
a a
a 2
a. a
a
a- 05
T3
2 2 S 2
H
on
X
p
ZONES
8490
8310
8250
8240
8145
8125
8025
7945
7855
7805
X X
9
X
? X X
X
X X
T. approximatus
■ A. antiquus
7765
7760
7750
7690
7685
C. aureus
A. richardsoni
S. tenuis
TABLE 2. Canyon Creek Section (66° 10' N.; 136° 05' W.)
measured by Dr. B. S. Norford, Geological Survey, Canada, 1963.
O
o o
S o’
cr .05
0? o'
■< fc
g JS
3 3
0“ <
a n,
n
p
D
P
CL
O
O
P
cr
o
<
rp
cr
p
S ^ 5^
a I
11-
2 ■§ 5
c« a
Ti
b n
S' S'
'§ a
a 3?
a a
2 i
» a
. O 55
^ a .'o t/5
■a
Xi
NJ "J /a O
S ^
CD
OQ
OQ
P
O
?*r
c/3
£ o
c
bIQQ
-=—33
.2 2 a a
35 c^ 05
•S a .2
^ “a as
a a a a
5' Qd
a. ; .
53034
53032
53031
1054-7
862
503-504
X X
XXX
XXX
JACKSON: TREMADOC GRAPTOLITES
885
Remarks. The four graptolite subzones proposed by Jackson (1974) are recognized
at the following levels in the Rock River section: Adelograptus antiquus 7805-8310 ft;
Clonograptus aureus 7760-7765 ft; Anisograptus richardsoni 7690-7750(7) ft; and
Staurograptus tenuis 7685 ft.
In the Canyon Creek section, a limestone breccia at 948-967 ft is correlative with
a conglomerate marker horizon in the Upper Canyon on Peel River (Jackson 1974,
p. 57). The assemblage from GSC loc. 53031 is probably from the C. aureus Zone
because adelograptids and Anisograptus are absent. The two younger collections
probably represent the Adelograptus antiquus Zone on the basis of the clonograptid
composition.
In conclusion, the occurrence of Jackson’s (1974) proposed subzones on Rock
River, 100 km north of Peel River, demonstrates that these biostratigraphic units
are widely distributed along the Richardson Trough and for this reason are raised
to zonal status. The discovery of Bryograptus ramosus in the basal upper Tremadoc
tends to support the correlation of the Clonograptus aureus Zone with shales of
3a/3 age in Norway.
SYSTEMATIC DESCRIPTIONS
Class GRAPTOLiTHiNA Bronn, 1849
Order dendroidea Nicholson, 1872
Family dendrograptidae Roemer in Freeh, 1897
Genus dictyonema Hall, 1851
Dictyonema cf. percancellatum Ruedemann
Text-fig. \e
cf. Dictyonema percancellatum n. sp. Ruedemann 1947, p. 172, pi. 4, figs. 13, 14.
Material. One compressed and fragmented rhabdosome GSC (Geological Survey of Canada) hypotype
27004 from GSC loc. 53032, Canyon Creek, collected by B. S. Norford, 1963.
Description. Rhabdosome fragmented 11 mm long and 10 mm wide. Stipes are
0-3-0-4 mm wide dorsally and number fourteen per cm. Details of thecae not seen.
Dissepiments are less robust, 0-4 mm long, number eighteen per cm, and tend to have
a common alignment across the entire rhabdosome. The frequency of dissepimental
spacing suggests that dissepiments are produced at the level of every autotheca (or
bitheca) or at alternate autothecae and bithecae. Intra rhabdosomal spaces tend
to be square.
Remarks. The close spacing of stipes and dissepiments makes this a distinctive
dendroid. The nearest comparison that I have been able to make is with D. per-
cancellatum from St. Pauls Inlet, Newfoundland. Ruedemann’s original description
merely dated the species as Ordovician. However, published accounts by Kindle and
Whittington (1958, p. 327) show that Tremadocian rocks do exist in the area.
886
PALAEONTOLOGY, VOLUME 18
Family anisograptidae Bulman, 1950
Genus bryograptus Lapworth, 1880
Bryograptus ramosus Br0gger, 1882
Text-fig. la, b, d
1882 Bryograptus ramosus Br^gger, p. 37, pi. XII, fig. 21.
non 1894 Bryograptus ramosus Br^gger; Marr, p. 125, figs. 1-5.
1925 Bryograptus ramosus Br0gger; Monsen, p. 160, pi. 1, fig. 9; text-fig. 3.
1954 Bryograptus cf. ramosus Br0gger; Bulman, p. 34, pi. 4, fig. 9.
1963 Bryograptus ramosus Br0gger; Spjeldnaes, p. 122, pi. XVII, figs. 7-9; text-fig. 1.
1965 Bryograptus ramosus Br0gger; Erdtmann, p. 105, pi. 2, fig. 5.
1966 Bryograptus ramosus (Br0gger); Szymanski, pp. 50, 59, pi. vi, fig. 9.
1971 Bryograptus ramosus Br0gger; Bulman, pp. 365-366, figs. le,f, 2c.
Material. Five compressed specimens GSC hypotypes 27000, 27001, and 27002 from GSC loc. 53031,
Canyon Creek, collected by B. S. Norford, 1963.
Description. Rhabdosome 21 mm long and 20 mm across distally. Fourteen terminal
stipes are produced by dichotomous and ? lateral branching from three primary stipes.
Two zones of branching occurs at 3 0-3-6 mm and 6-6-9 0 mm from sicula. Sicula
1-3 mm long furnished with a fine nema. Two primary stipes have three thecae and
the third stipe has one theca, these stipes diverge from sicula at about 60-80° then
curve rapidly inwards to become sub-parallel. Stipes have a maximum dorso-ventral
width of 0-6-0-7 mm across thecal aperture and 04-0-5 mm just above the aperture;
free ventral wall of thecae are concave and inclined at 30-40° near aperture. Auto-
thecae number 16-20 per cm, bithecae not seen.
Remarks. Bryograptus ramosus differs from B. kjerulfi in having more widely spaced
zones of dichotomy and closer spacing of autothecae.
It is distinct from B. broeggeri Monsen which has more widely dispersed zones of
branching and a characteristically long and slender sicula.
In Scandinavia, this species is characteristic of the lower part of 3a^ (Monsen
1925) and according to Erdtmann (1965) ranges upwards into 3ay beds. Similarly,
Szymanski (1966) recorded it from upper Tremadoc of Bialowieza, Poland. The
assemblage from GSC loc. 53032 possibly should be assigned to the C. aureus Zone
on account of its position relative to the conglomerate marker and because of the
lack of adelograptids.
REFERENCES
BROGGER, w. c. 1882. Die silurischen Etagen 2 und 3 im Kristianiagebiet und auf Eker. Krisfiania (Oslo).
376 pp.
BRONN, H. G. 1849. Index Palaeontologicus B. Enumerator. Stuttgart. 980 pp.
BULMAN, o. M. B. 1950. Graptolites from the Dictyonema Shales of Quebec. Q. Jlgeol. Soc. Land. 106, 63-99.
1954. The graptolite fauna of the Dictyonema Shales of the Oslo region. Norsk geol. Tidsskr. 33, 1-40.
1971. Some species of Bryograptus and Pseudohryograptus from Northwest Europe. Geol. Mag.
108,361-371.
ERDTMANN, B.-D. 1965. Fine spat-Tremadocische Graptolithen fauna in Oslo. Norsk geol. Tidsskr. 45,
97-112.
FRECH, F. 1897. Letliaea Geognostica, 1, Th. Leth. pal. 1. 11, Graptolithen. Stuttgart.
HALL, J. 1851. New genera of fossil corals etc. Am. J. Sci. (2), 11, 398-401.
JACKSON: TREMADOC GRAPTOLITES
887
JACKSON, D. E. 1974. Tremadoc Graptolites from Yukon Territory Canada. In rickards, r. b., jackson,
D. E. and HUGHES, c. p. (eds.). Graptolite Studies in Honour of O. M. B. Bulman. Spec. Pap. Palaeont.
13, 35-58.
KINDLE, c. H. and WHITTINGTON, H. B. 1958. Stratigraphy of the Cow Head Region, Western Newfoundland.
Geol. Soc. Am. Bull. 69, 315-342.
LAPWORTH, c. 1880. On New British Graptolites. Ann. Mag. nat. Hist. (5), 5, 149-177.
MARR, J. E. 1894. Notes on the Skiddaw Slates. Geol. Mag. 31, 122-130.
MONSEN, A. 1925. liber eine neue Ordovicishe Graptolithen fauna. Norsk geol. Tidsskr. 8, 147-187.
NICHOLSON, H. A. 1872. Monograph of British Graptolites. Edinburgh and London.
NORFORD, B. s. 1964. Reconnaissance of the Ordovician and Silurian rocks of Northern Yukon Territory.
Geol. Surv. Canada Paper 63-39, 1 39 pp.
RUEDEMANN, R. 1947. Graptolites of North America. Mem. Geol. Soc. Am. 19, x-|-652 pp., 92 pis.
SPJELDNAES, N. 1963. Some Upper Tremadocian graptolites from Norway. Palaeontology, 6, 121-131.
SZYMANSKI, B. 1966. Dictyonema Shales of the Krzyze Beds Region of Bialowieza. Kwart. geol. 10, 44-62.
Typescript received 16 October 1974
Revised typescript received 11 December 1974
D. E. JACKSON
Department of Earth Sciences
The Open University
Milton Keynes, MK7 6AA
THE PALAEONTOLOGICAL ASSOCIATION
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(Vice-President), Dr. J. M. Hancock (Treasurer), Dr. C. T. Scrutton (Secretary), Dr. D. D. Bayliss, Dr.
M. C. Boulter, Dr. C. H. C. Brunton, and Dr. A. W. A. Rushton. The Committee looked first at the future
utilization of the Association’s financial resources and detailed proposals are currently before Council for
consideration. The procedure for election of Council was also given early attention. This has resulted in
a proposal for changes in the Constitution to allow a postal ballot when nominations exceed vacancies;
THE PALAEONTOLOGICAL ASSOCIATION
891
guide-lines for the conduct of a ballot have now been agreed by Council. At the same time it is proposed
to write into the Constitution the terms of office for Officers agreed by Council last year. An informal
meeting has been held between representatives of the Association and the Geological Society of London
to discuss matters of common interest. The meeting resulted in a useful exchange of views and ideas covering
among other things publications, promotional activities, arrangements for meetings, the Geological
Society’s Library, and geological conservation. As well as arranging the Association’s established annual
programme of events, Council continues to review suggestions for additional indoor and outdoor meetings.
Following the success of the International Symposium on the Ordovician System, Council is actively
exploring the organization of future symposia on similar lines. Dr. C. A. Fleming retired from his post as
Overseas Representative for New Zealand at the end of 1974. Dr. G. R. Stevens (New Zealand Geological
Survey) has agreed to serve in his place. Thanks are due to Dr. Fleming for his long service to the Association.
P2
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