The Micropaleontology Project, Inc.

A Study of the Morphology and Variation of Some Upper Lias Foraminifera

Author(s): G. C. Adams
Reviewed work(s):
Source: Micropaleontology, Vol. 3, No. 3 (Jul., 1957), pp. 205-226
Published by: The Micropaleontology Project, Inc.
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ABSTRACT: This paper gives an accountof the morphologic variationand dimorphismshown by certainspeciesof
foraminiferabelonging to thefamily Lagenidae in the UpperLias of northernLincolnshire, England.The variation
of the is
generation described,
megalospheric and the relative
importanceof diferentmorphologicalfeatures
commonlyusedin
the diagnosisof speciesis discussed.The clearmorphologic and
separationof microspheric megalospheric generationsis
established
in onecase,butno supportfor the is
theoryof trimorphism givenby this investigation.

A study of the morphology and variation of

some Upper Lias foraminifera

G. C. ADAMS
King's College
Universityof London

INTRODUCTION

This communication forms part of a general study have been examined, and all the membersof a grad-
of the Upper Lias foraminiferaof northern Lincoln- ing series are regarded as belonging to a single spe-
shire, England, the results of which are contained cies where gradation between these characters has
in an unpublished Ph.D. thesis written at the Uni- been established. Exceptions are made for those
versity of Nottingham in 1955. A paper incorporat- forms where gradations are merely between extreme
ing a systematic description of the whole foram- variants of two species groups, and where other
iniferal fauna is at present being prepared. The charactersof definitive importance have been found.
original work showed that many Liassic species and Such exceptions will be described in a subsequent
genera of the family Lagenidae exhibit great mor- paper.
phologic variation, a fact recognized by most previ-
ous workers, and the present paper seeks to analyse In many early studies of the family Lagenidae,
this variation in a number of cases. species have been erected on solitary or rare megalo-
In the author's opinion, studies of the morphologic spheric individuals, an understandable error when
variation shown by members of a community are of the alterationof generationswas unknown. However,
more importance in modern paleontology than the in much of the literature, including some of that
more recently published, new species are described
compilation of species lists, which may not reflect with no regard to the variation shown by the species
the true interrelationships of the fauna. A multi-
plicity of specific names often tends to obscure the group as a whole. When microsphericspecimens are
close affinity of individuals which may be poten- lacking, identification is necessarily based on their
tially interbreedingmembersof a community. A bet- megalospheric counterparts, even though the latter
ter understanding of the evolution of a group and may be highly variable. Nevertheless, even in an
of the problems involved in species differentiation assemblage comprised of two or more species, the
will be reached if each assemblage of individuals megalospheric forms of which intergrade, measure-
ments of the definitive biocharacters, e.g., number
showing gradation in their morphology is treated of ribs, degree of coiling, overlap of chambers, etc.,
as an evolving plexus. An attempt has thus been
made to analyse the variation shown by some should reveal the presence of more than one species
when plotted on distribution diagrams. The modal
lagenid plexuses, rather than to list a number of
arbitrarily defined species. position of each species should be distinctive,
although some overlapping will occur. It is, of
Where an assemblage of lagenids appears to include course, possible that end members of a plexus that
groups of two or more closely related species, the have previously been regarded as distinct species
biocharacters on which those species were founded may merit specific rank at other horizons or in

vol. 3, no. 3, pp. 205-226,text-figs.1-26,july,1957

micropaleontology, 205
 ADAMS

different localities. An incidental result of such It has been the experience of the present author that
studies is that the tendency to regard recurrent these characters are not always constant even
homeomorphs as long-ranging species is avoided. within the limits of a single species. The genus Den-
talina serves to illustrate this. Dentalinatenuistriata
In addition to the foregoing, an attempt has been Terquem is a commonly occurring Lias species, and
made to determine whether or not any species ex- in the large amount of material possessed by the
hibitstrimorphism.The relationships,if any, between author is always markedly costate, that is to say, it
dimorphism and variation have also been investi- possessesnumerous fine ribs which on no specimens
gated. show any sign of reduction. Dentalina primaeva
The present paper is intended to show how some d'Orbigny, on the other hand, shows all stages of
costation from a well-ribbed form at one extreme to
genera and species varied within a small area a completely smooth form at the other. In the first
throughout a relatively short period of time. case the presence of ribs is a diagnostic feature of the
Although the conclusions reached may be valid
species, whilst in the second it is not. Similar ex-
only for this area, a number of such studies in other amples could be cited from other lagenid genera,
areas will, it is hoped, lead to a morecomplete under- such as Frondicularia,Lingulina, and Astacolus,to
standing of the variation and evolution of the family mention but a few.
Lagenidae.
Nor is Brotzen on safe ground when he deals with
The author is indebted to Professor W. D. Gill, the cross-sectional shape of the test. The genera
Trinity College, Dublin, and to Dr. J. E. Prentice, Lenticulinaand Astacolusgrade into one another as
King's College, London, for reading the typescript do Astacolusand Planularia;similarly, Marginulina,
and for offering much helpful advice. Thanks are Dentalina, and Vaginulinaare variable, and any
also due to the University of Nottingham for the attempt to separate them on the cross-sectional
award of a Revis Studentship, during the tenure of shape of the test alone, save in the most arbitrary
which this work was undertaken. fashion, seems impossible. It may perhaps be said
that if the problem of the nomenclature of the
Mesozoic lagenids is ever satisfactorilysolved it will
HISTORICAL REVIEW be by means of variation studies of the earlier plex-
uses and not by the application of a terminology
The only recent work on the Liassic Lagenidae of based on their later, perhaps more stable, descend-
Britain has been carried out by Macfadyen (1941) ants.
and by Barnard (1950a, 1950b, 1956). Most of this
work concerns type and homoeotype material only, PREPARATION OF MATERIAL AND WORKING METHODS
but the latter author has studied variation in some The foraminifera used in this study all came from
genera, notably Planulariaand Pseudoglandulina. His a cored bore-hole situated at Risby-Warren near
comprehensivework on the Lagenidae of the Lias is Scunthorpe, Lincolnshire. The core included
awaited with interest. The microfaunaof the Lias of 57 feet of Upper Lias shale, representingthe Dactyli-
northern Lincolnshirehas not been studied hitherto. ocerastenuicostatum and Harpocerasfalcifer zones. The
whole of the Upper Lias in this area is known to
Many authors have commented on the difficulties
encountered when trying to classify members of the belong to these divisions, and zonal ammonites were
obtained from this particular core. Identifications
Lagenidae, owing to their extremevariability (Cush- of ammonites were carried out by the Geological
man, 1940; Glaessner, 1945; Bartenstein, 1948a, b,c,
Survey of Great Britain, and the specimens remain
etc.), and Bartenstein (1948a, b, c) has attempted to in their keeping. The types and figured specimens
resolve some of the nomenclatural confusion that
of the foraminiferaare deposited in the collections of
exists. Brotzen (1953) has suggested a new method
the British Museum (Natural History) (B.M.N.H.),
of classifying the Lagenidae based on the supposed
"constantcharacters"of TriassicandJurassicgenera. London.
This method would involve the introduction of a The cored material was divided into one-foot
large number of new generic names, at least fifty-six lengths and its lithology noted. The only variations
generic names being required for the uniserial recti- appeared to be in hardnessand in mica content, the
linear types alone. The two basic characters which lowest 6 feet being very soft and micaceous. In view
Brotzen believes to be constant are (a) the shape of of the thickness of shale comprising each sample
the test in cross-section; and (b) the presence or (one foot), it cannot be claimed that the contained
absence of costae. foraminifera represent a single community, but,

206
 UPPERLIASFORAMINIFERA

owing to the condition of the core, it was impossible An important feature of every assemblage is the
to deal with smaller divisions. It is therefore ac- marked lack of microspheric forms. At some levels
knowledged that the variation seen amongst speci- these are entirely absent, a phenomenon which it is
mens in a given sample did not necessarily exist difficult to explain. The large amount of variation
amongst a single living community, although even which occurs is therefore virtually limited to mega-
that is not impossible, as only a minute area is con- lospheric forms, a factor which probably accounts
sidered; nevertheless, it all must have occurred for the large number of specific names which have
within a relatively short span of geologic time. in the past been assigned to this group.
Each sample comprised about one and a half pounds From time to time authors have recorded different
of shale, which was disaggregated by the usual morphological features, or combinations of features,
methods, the residue being collected on four sieves as being of diagnostic importance in the identifi-
of different sizes, the finest being 200 mesh. Speci- cation of species of Planulariaand Falsopalmula.An
mens were carefully collected by hand, nothing attempt has been made to assess the value of these
characters when applied to the present material.
being ignored except for the very immature stages
of large species, such as Marginulina Over 1000 specimens were examined.
primad'Orbigny,
which collected on the 200-mesh sieve. All subjective The following characters have, at various times,
selection was thereby avoided. been considered of diagnostic value in delimiting
species of Planularia:Size of test; degree of initial
Only species showing considerable variation and coiling; angle made by later sutures with the periph-
representedby numerous individuals are dealt with eral wall; presence of ribs or striations; shape of
here. Thus, out of some seventy species of Lagenidae individual chambers; shape and size of proloculum;
identified, only six fulfilled these conditions. In and amount of lateral compression of the test.
some cases, e.g., Planularia,where direct comparison
of specimens was necessary, it was found useful to Size of test: At first sight, variation in the size of the
photograph all the specimens on a Vickers projec- test might appear to constitute a good criterion of
tion microscope and to work with the resulting species,and indeedit may sometimesdo so. Obviously,
prints. there is an optimum size to which the individuals
of a species will grow in a particular environment
MORPHOLOGY AND VARIATION
when the conditionstherein remainconstant. Equally
clearly, if conditions in the environment change
The most variable and readily studied species were slightly, the maximum size attainable by the indi-
those belonging to the genera PlanulariaDefrance, viduals of the same species might be increased or
FalsopalmulaBartenstein, Marginulina d'Orbigny, decreased. Such environmental changes may or may
FrondiculariaDefrance, and Lingulina d'Orbigny. not be reflected in slight lithologic changes within
With the exception of Frondicularia
these are all dis- the sediments, and if such changes do occur, then
cussed below. changes in test size are not surprising.
The appearance of the material used in this in-
SYSTEMATIC DESCRIPTIONS
vestigation seems to suggest that test size is affected
Order FORAMINIFERA by environment, but as similar specimens have not
been compared from different localities, nothing can
Family LAGENIDAE yet be proved. The observed changes in size prima-
Genus PlnnulariaDefrance, 1824 rily affect genera other than Planularia,e.g., Marginu-
lina, where a change in the lithology of the sediments
Representatives of this genus occur abundantly seems to be reflected in the abundance and size of
throughout the core. They are present at all levels the tests present. This lithologic change is slight,
but are especially numerous in the upper part. A merely an alteration from soft micaceous shale to
number of immature specimens were found in each somewhat harder non-micaceous shale. The pres-
sample, but it was impossible to decide whether ence or absence of the mica itself has no direct
these should be referredto Planulariaor Falsopalmula, connection with the condition of the fauna, for mica-
as the young stages of these genera are identical and rich shales are known with both rich and impover-
the two occur together. The greatest degree of vari- ished faunas. However, in this case the sudden
ation is seen, as might be expected, where the spec- change to non-micaceous shale may reflect some
imens are most abundant, and a careful study has local fluctuation in conditions, other aspects of
been made of the population at those levels. which could account for the changes.

207
 ADAMS

On the whole, not much information was obtained These two series illustrate the variations that may
with regard to variation in test size in the case of occur when one feature of the morphology is re-
Planularia.It can only be stated that individuals garded as constant. However, in practice it has not
tended to be of a larger size toward the bottom of been possible to find any constant features, so that
the section sampled (i.e., in the micaceous shale) the resulting possibilities of variation in shape
than they were at the top. Unfortunately, only a within any assemblage are enormous.
small number of specimens occurred in the lower The remaining morphological features are dealt
part of the section. with under Falsopalmula,a genus closely related to
Planularia.
Degree of initial coiling: A characteristic feature of
the genus is that all its members either possess an Planularia pseudocrepidula Adams, new species
initial coil or show an initial tendency to coil.
Text-figures 1-5
Microspheric forms are, without exception, coiled. Cristellariacrepidula(Fichtel and Moll). - BLAKE, 1876, York-
Measurements of the amount of coiling have been shire Lias, p. 466, pl. 17, fig. 25; pl. 19, figs. 12, 14, 15. -
made by counting the number of chambers that are FRANKE, 1936,Preuss.Geol. Landesanst.,Abh., new ser.,
in contact with the proloculum, or that overlap the no. 169, p. 95, pl. 9, figs. 22, 23a-b. -BARTENSTEINAND
1937, Senckenb.Naturf. Ges., Abh., no. 439,
proloculum and touch the second chamber. This
BRAND,

method has a disadvantage in that it reflects only p. 169, pl. 3, fig. 48; pl. 4, fig. 76; pl. 5, fig. 74; pl. 9,
fig. 45a-c.
the degree of curvature of the inner side of each
crepidula(Fichtel and Moll). - BRADY, 1867, Somer-
?Cristellaria
chamber. The outer side may be equally curved or setshireArchaeol.Nat. Hist. Soc., Proc., vol. 13, pt. 2,
completely evolute. Whilst there is no theoretical p. 228, pl. 3, fig. 39. - ISSLER,1908, Palaeontographica,
vol. 55, p. 79, pl. 5, figs. 244-248.
upper limit to the number of chambers in a coil Not NautiluscrepidulaFICHTELAND MOLL, 1798, Test. Micr.,
(save that it cannot exceed the total number of p. 107, pl. 19, figs.g-i.
chambers in the test), the lower limit is, of course,
two. Text-figure la shows a series of forms that Diagnosis: Test highly compressed, thin and glassy in
appearance,consistingof up to twelve chambers,of
grade from an initially close-coiledindividual (no. 1) which the first few normally form an initial coil; later
to a virtually uncoiled specimen (no. 5). The graph chambers become evolute on the peripheral side, but
below indicates the number of specimens of each on the ventral side tend to reach downward toward
type present in the sample, and it is clear that the the coil. Sutures narrow, flush or slightly deepened at the
close-coiled and uncoiled individuals are end mem- surface. Proloculum variable in shape, commonly either
bersof a continuouslygradingseries,typical members subspherical or oval. Aperture terminal, situated at the
of which are centrally situated on the graph. peripheral edge of the last chamber.
Text-figure lb indicates variations in the width of Dimensions: Maximum length 0.71 mm.; maximum
the test at the same level amongst specimens that width 0.23 mm.
are equally coiled. In this case three "species"could Variation:This species is extremely variable, especially
possibly be recognized: Planularia pauperata (Jones in size, shape, and degree of coiling (detailsare given
and Parker) (nos. 1 and 2), Planulariapseudocrepidula below). It appears to grade into and be inseparable
Adams, n. sp. (= Planularia crepidula (Fichtel and from Planulariapauperata(Jones and Parker), Planularia
Moll) of most workerson the Lias) (nos. 3 and 4), and protracta(Bornemann), and Planulariafilosa (Terquem),
Planularia protracta (Bornemann) (nos. 5 and 7). althoughat least the firsttwo of theseseemto be readily

TEXT-FIGURES 1-4
1a (1-5), Planulariapseudocrepidula Adams, n. sp., series showing variation in degree of coiling at one level, all x 60;
the graph below shows the number of individuals in the sample which have the same degree of coiling; the numbers
on the graph correspond with those of the specimens above; all the specimens are megalospheric. 1b (1-7), Planularia
pseudocrepidula Adams, n. sp., a grading series of individuals, all of which have the same amount of initial coil, all x 40;
in this case the width varies; all the specimens are megalospheric and are from the same level as those in text-figure 1a.
2, Planulariapseudocrepidula Adams var. robustaAdams, n. var., six typical specimens, all megalospheric, all x 47.
3, PlanulariapseudocrepidulaAdams, n. sp.; a, variation at 26-27 feet (all megalospheric, all x 50); the specimens resemble
the following "species": 1-4, Planulariacrepidula(Fichtel and Moll) of various authors; 5-7, Planulariaprotracta(Borne-
mann) of various authors; 8-10, Planulariadorsoarcuata (Wisniowski) of various authors; 12-13, Planulariaanceps(Ter-
quem) of various authors; b, variation at 22-23 feet (all x 50): 6-9, Planulariadorsoarcuata (Wisniowski) or Planularia
filosa (Terquem) of various authors; c, variation at 16} to 17- feet (all X ca. 47) (see also text-fig. 5a): 1-5, Planularia
pseudocrepidula Adams var. robusta Adams, n. var. 4, graphs showing the differences in the number of chambers devel-
oped by Planulariapseudocrepidula and Planulariapseudocrepidula var. robusta.

208
 UPPER LIAS FORAMINIFERA

34 5

. ,

;;?-tl~ ~ ~ ~
~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~F*I1.~~~~~~~~~~~~~~~~~~~~~~~~
1

1 2 3 4 5 6
2 3 4 5

30-
No. of
20
specimens
10-

2 5
4 7 8 9 10 II
Degree of coiling

TEXT-FIGURE la

12 13 14
TEXT-FIGURE 3a
2 3 4 5 6 7

TEXT-FIGURE 1b

"
I
;?

?urC't t'
r? ??.?

s;s? t??

I 3 4

'I'm'
I?w
--- 9
TEXT-FIGURE 2

5 6 7 8 9
TEXT-FIGURE 3b

I I 1- -A
4 5 6 7 8 9 10 II 12
No. of chombers
Plonuloria pseudocrepidula sp. nov.----
11 11 vor. robusta- I 2 3 4 5
TEXT-FIGURE 4 TEXT-FIGURE 3c

209
 ADAMS

I 2 3 4 5 6
TEXT-FIGURE Sb
I 2 3 4 5

1 2 3 4 5
TEXT-FIGURE 5c
6 7 8 9 10

II 12 13 1 2 3 4 5 6
TEXT-FIGURE Sa TEXT-FIGURE 5d

TEXT-FIGURE 5
Planulariapseudocrepidula Adams, n. sp., all x ca. 46; a, variation at 161 to 17| feet (continued): 12-13, typical
uncoiled forms of a type that has often been referredto Vaginulina;b, variation at 14 toi 15i feet; of these specimens,
no. 6 could be referred to the "species" Planulariaanceps(Terquem); c, variation at 101 to 111 feet; these are some
of the smaller forms at this level; see text-figures la and lb for more typical individuals; d, variation at 9-10 feet: 1-3,
typical specimens of Planulariapseudocrepidula.

a b c d e

TEXT-FIGURE 6
Five small specimens of Planularia(or Falsopalmula?)from the upper part of the section. Similar specimens have been
gracilisKiibler and Zwingli, Cristellaria
recorded as Cristellaria Kiibler and Zwingli, Cristellaria
argovensis parkinsoniKiibler
and Zwingli, CristellariamitraKiibler and Zwingli, and Cristellariasemi-involutina
Terquem var. plana Deecke; they are,
however, clearly young stages of either Planulariaor Falsopalmula.It is significant that all such specimens are mega-
lospheric.

210
 UPPER LIAS FORAMINIFERA

identifiablein some districts.Its young stages are in- The general tendency here is for individuals to be
distinguishablefrom those of Falsopalmula deslongchampsi longer and narrower than the typical Planularia
(Terquem). pseudocrepidula.
Remarks: Planularia is a very commonLias
pseudocrepidula The usual series in coiling is present at 22-23 feet.
speciesover the whole of westernEurope.In northern
Lincolnshireit is found throughoutthe Upper Lias. Text-figure 3 b shows the variation. Two main vari-
Previous authors have referredits representativesto ants are present, nos. 1-3 being a larger, slightly
Cristellariacrepidula(Fichtel and Moll), which was, more robust form, and nos. 6-9 an uncoiled elongate
however, originally described from Recent material, form, Planulariadorsoarcuata (Wisniowski) or Planu-
and which, in any case, is the type speciesof Astacolus laria filosa (Terquem). Decrease in width and in
Montfort, 1808. It is much longer, more robust, and coiling occurs simultaneously. The form protractaof
less compressedthan the Lias form here under con- Bornemann is less noticeable and not so well de-
sideration.The name "pseudocrepidula" has been chosen veloped.
because it immediatelyrecalls the older name which
has so oftenbeen usedin error. Two principal variants occur at 164 to 17? feet.
The first is a short, broad form which has a very
Types:Holotype, B.M.N.H. no. P.43374. Paratypes
(fourspecimens),B.M.N.H. no. P.43375. large proloculum and a relatively small number of
chambers succeeding it, usually seven or eight. The
Material:Approximately400 specimensfrom the Upper
Lias of northernLincolnshire. coiling series can again be distinguished. Nearly all
the specimens are megalospheric, the largest pro-
Variationin successivecommunities in the UpperLias of loculum measuring 0.11 x 0.04 mm.; the shape of
northernLincolnshire: The variation was studied in these forms varies from oval to subspherical. The
every sample that yielded sufficient specimens. The second variant is a rather well-developed triangular
numbers given below refer to the depth of the form of Planulariapseudocrepidula. This variant forms
sample below the top of the Upper Lias, a horizon an uncoiling series, at the end of which are a small
which was itself 80 feet below ground level. It is number of specimens that resemble Vaginulina rather
clear that the range of variation is not uniform at than Planularia;they are, however, no more than
all horizons, the principal variants tending to differ end members (all megalospheric) of the latter.
in number and in type from one horizon to another.
In the lower part of the successionthe specimens are The first variant (text-fig. 3c, nos. 1-5) occurs in
neither numerous nor very variable. All the indivi- large numbers and is so strikinglydifferent from the
duals begin with an initial coil. A few specimens typical form of the species that it merits a distinct
varietal name, Planulariapseudocrepidula robusta.Text-
occur which closely resemble Planularia pauperata
figure 4 shows graphs of the number of chambers
(Jones and Parker) in shape, but as they are outside
the normal range of variation exhibited by Planu- possessed by Planularia pseudocrepidula Planularia
and
laria pseudocrepidulaelsewhere in the section, they are pseudocrepidula robustaAdams, n. var., at this horizon.
best referred to the former species.
The difference between the two can be seen even
with this simple measurement. The genus Planularia
Above 29 feet, specimens become more numerous attains its maximum abundance at the 16- to 171
and are especially abundant at some levels, where foot level.
the variation has been studied in detail. At inter-
mediate horizons, specimens are less numerous and A return to a smaller size is evident at 144 to 154
always occur together with Falsopalmula, so that feet (text-fig. 5b). The larger, more robust varieties
most of them are probably young stages of a species are absent, their places being taken by small (usu-
of that genus. ally less than 0.4 mm. in length), narrow individuals
At 26-27 feet, microspheric forms are very rare. The
with highly oblique sutures. These could be referred
range of morphology is illustrated in text-figure 3 a.
to as Planulariadorsoarcuata (Wisniowski), Planularia
The initial coil is usually rather weak, and quite a filosa (Terquem), and Planularia anceps(Terquem).
Many may be young stages of Falsopalmula, of which
large number of specimens are almost completely individuals are
uncoiled. All the specimens illustrated are megalo- comparable present.
spheric and might have been ascribed by earlier At 10 to 11 feet, the specimens again attain a
workers to the following species: larger size, this time tending to become broadly
Nos. 1-4: Planularia crepidula(Fichtel and Moll); triangular in outline. The variation in coiling and
Nos. 5-7: Planulariaprotracta(Bornemann); in width of test has already been referred to above
Nos. 8-10: Planularia dorsoarcuata(Wisniowski); and indicated in text-figure 1. Other variants pres-
Nos. 12-13: Planularia anceps(Terquem). ent at this horizon are referable to Planulariapro-

211
 ADAMS

20- I1-2' 20- 2-3

4
c .I c
15- I
' I
I | 15-
E I
I
I
I I
I
0. o I
10- S-10- I
I

II
o0 0 I
I
I
6 5- 6 5- t
I
z z I

No3 o f6 cha1 9 6 ' ' 12

No. of chambers No. of chambers

20- 20-
14- 15\ 16e- 17'
c: c
E 15- 15-
C?

0
- 10- /
,,
'

C.

6 5- / '7o5-
z
/

5 I t I I I i I. . -. .

3 6 9 12 N3 6 9 1'2
No. of chambers No. of chambers

20- 26-27
A Number of Planularian
d 15-
I,,
chambers
i /
E
I I

.-10-
I
.- - Number of chambers
taking part in coil
0 /
5-
z
I
t ? I I , i I I

3 6 9 12
No. of chambers
TEXT-FIGURE 7

Falsopalmula;frequency polygons showing the number of planularian chambers and the number of chambers in a
coil at five different levels.

212
 UPPER LIAS FORAMINIFERA

tracta (Bornemann) and Planulariafilosa (Terquem) specimen that differed in shape from its fellows. Yet
(text-fig. 5c). There does not appear to be any way this difference in appearance is largely a feature of
of distinguishing between them, and both are numer- the megalosphericgeneration, and most of the vari-
ically of subsidiary importance to more typical ex- ants that occur, although common at a given level,
amples of Planulariapseudocrepidula. are impersistentin time. The fact that similar forms,
At 9 to 10 feet, variation is very similar to that at e.g., Planulariadorsoarcuata (Wisniowski) appeared
several times in the Jurassic does not necessarily
10? to 11? feet. The triangular form of Planularia
imply any direct genetic continuity between them.
pseudocrepidulapersists, and specimens referable to On the contrary, it is far more likely that such
"protracta"and "filosa" are still present, although in
reduced numbers. The absence of the long, narrow, morphologic "species" are homeomorphs of one
another.
uncoiled "dorsoarcuata"type is notable.
The author therefore suggests that, when dealing
From this point on, Planularia is gradually replaced with communities of Jurassic members of the genus
in importance and numbers by Falsopalmula, and it
Planularia,different names should not be allotted to
is impossible to separate the young stages of the two what at first sight seem to be radically different
genera. All the specimens of Planularia tend to be species unless (a) enough individuals have been
small, although they continue to exhibit various examined to show that there is no gradation between
stages of coiling and uncoiling. Text-figure 6 shows them, and (b) microspheric forms matching each
some of this variation. Many specimens similar to
type of megalospheric form can be found. In the
these have, in the past, received specific names, but
present investigation the strikinglydifferent megalo-
it is evident that they are merely young stages of
spheric forms could not be matched by different
Planularia and Falsopalmula. Specimens similar to
microsphericforms, and the former graded into one
those figured have been recorded as Cristellariagra- another. Specific differences based on the amount
cilis Kubler and Zwingli, CristellariaargovensisKtibler of coiling of the megalosphericform are seen to have
and Zwingli, Cristellaria parkinsoni Kubler and no validity in the present case, as all gradations
Zwingli, Cristellaria mitra Kubler and Zwingli, between coiled and uncoiled forms exist.
and Cristellariasemi-involutina
Terquem var. plana
Deecke. It is significant that all these "species" were PlanulariapseudocrepidulaAdams var. robusta
recorded as rare by the original authors, and it is to
be noted that they are all megalospheric.
Adams, new variety
Text-figures 2-4
Conclusions:In this succession, the genus Planularia
is represented by Planularia pseudocrepidulaAdams,
Diagnosis:Test short and broad, composedof from six
to elevenchambers,usuallyaboutseven.In the megalo-
n. sp., a species that exhibits considerable morpho- sphericform the proloculumis large, generallyoval in
logic variation. Of common occurrence at certain shape, and is usuallyfollowed by chamberswhich in-
horizons are specimens referable to Planularia pro- crease rapidly in width, the first few tending to form
tracta (Bornemann), Planularia dorsoarcuata(Wis- part of the initial coil.
niowski), etc. Isolated specimens appear to be quite Dimensions:Maximum length 0.64 mm.; maximum
distinct forms, but when large numbers are examined width 0.25 mm.
they are not clearly separable from Planularia Variation:The testsvary greatlyin shape, but the aper-
pseudocrepidula.It is not intended to suggest that turaledge is usuallyfairlystronglyarcuate,especiallyin
such forms can never be present without grading the early stages. The number of chambers making
(this may occur in some areas), but only that in the up the coil varies from two to seven, with threeto five
present case theycannot be satisfactorilydistinguished. being the usualnumber.
There occur, as end members of a series graded Remarks: This variety differsfrom typical membersof
in coiling, a number of specimens similar to those the speciesin the largersize and smallernumberof the
which Franke (1936) designated as Vaginulina(Vagi- chambers.No definitemicrosphericformswereobserved,
nulina anceps Terquem). Such specimens do not, in but the specimenswith the smallestprolocula always
fact, belong to the genus Vaginulinad'Orbigny, but possessa fairly tight initial coil. This form was found
to Planularia Defrance; they differ from typical mem- at only one level.
bers of the latter genus only in being less coiled. The Types:Holotype, B.M.N.H. no. P.43376. Paratypes,
reason for the confused nomenclature associated B.M.N.H. no. P.43377.
with these species appears to be that the early work- Material:Ninety-onespecimensfrom the Upper Lias of
ers, knowing little or nothing about reproductive northernLincolnshire,at 161 to 171 feet fromthe top
dimorphism, gave a new specific name to every of the section.

213
 .

ADAMS

a b
TEXT-FIGURE 9

I 2 3
TEXT-FIGURE 8

.. . :. ..
es
?

9
'
":'?
?,

0 2 3 4 S 6
TEXT-FIGURE 10 TEXT-FIGURE 11

Width in mm. Width in mm.

0 0n 0 0n O n o 0 n 0 0
N N n * * n n N N 0
00 0000 0 o0 o0 0 9 99. lb 9 .

*100 .100
*095 *095
'090 -090
'085 *085
*080* * '080
E -075 E *075
E '070 ** *:: oo * E *070 *00 * **-
c 065 . . c *065 ':
*060 * 00 *060 ?
-c
*
'055 *055 .
* .0 c *050 0 *? ...
*?050 0_
_J
- 045 ?045 0 0

-040
*040 * . * 0 -040 ** 0 . 000
*035 .035 0

*030 .030
-025 .025
'020 *020

A 16 -17' B 26'-27'
TEXT-FIGURE 12

214
 UPPER LIAS FORAMINIFERA

Genus FalsopalmulaBartenstein, 1948 with characters of specific significance. Bearing in

Text-figures 7-12 mind the difficulties experienced by previous work-
This recently erected genus comprisesmany Jurassic ers, an attempt has been made to assess the value
of the features hitherto deemed to be of specific im-
species previously classed as Flabellina,a generic portance, when these are applied to the present
name which Bartenstein (1948b) replaced by JNeo- material. To this end, some 400 specimens have
flabellinaon the ground that the former name was been examined.
preoccupied by that of a mollusk. In the same year,
Bartenstein (1948a) further stated that Neoflabellina The following characters have, from time to time,
was not known to occur before the Cretaceous. been considered as being of diagnostic value in the
Falsopalmula also includes species previously assigned identification of species of Falsopalmula(Flabellinaof
by many American workers to the genus Palmula older authors): Number of "cristellarian" (planu-
Lea. The latter name had often been used as a syno- larian) chambers; number of chevron-shapedcham-
nym of Flabellina,but Bartenstein(1948 b) established bers; amount of overlap of the chevrons onto the
it as a valid genus, the earliest recorded appear- earlier chambers; degree of initial coiling; angle
ance of which is in the upper part of the Lower made by later sutures of the planularian part and
Cretaceous. the peripheral wall; presence of ribs or striations;
shape of individual chambers; shape and size of
Falsopalmulaappeared first in the Lower Lias, proloculum; amount of lateral compression of the
although not immediately in great numbers. In test; and depth of sutures.
Upper Lias time, however, it became abundant and Thenumber chambers:Theminimum num-
was of universal distribution. One of the most strik- ofplanularian
ber recorded was three (including the proloculum).
ing features of the genus is the wide variety of form
displayed by the individuals included in certain of Only one such specimen was found, the usual mini-
its species, and there can be little doubt that this mum at any level being four. Individuals of this
wide degree of individual variation has led to the type are few in number and invariably representthe
multitude of specific names that have been erected. megalospheric generation. The maximum number
recorded was twelve, and again only one such in-
Falsopalmulais present in almost all of the samples dividual was found, the usual maximum being ten
from northern Lincolnshire, being least numerous or eleven according to the horizon; these forms are
in the lowest 25 feet, where the majority of other always microspheric. At every level where Falsopal-
genera are most strongly represented, and most mulawas found, there was always a difference of six
abundant in the upper part, where many others to eight chambersbetween the one with the least and
are lacking. In this respect it parallels almost ex- the one with the greatest number. Text-figure 7
actly the distribution of Planularia. shows several graphs which indicate the number of
"planularian" chambers possessed by specimens at
The typicalform: A typical Lias example of Falsopal- different levels. Only one species is represented.
mula is a small, free-living, calcareous perforate (Superimposed on these diagrams are graphs show-
form consisting of an initial coiled series of planu- ing the number of chambers taking part in the ini-
larian chambers followed by a number of chevron- tial coil.) The main peak varies from the five- to the
shaped chambers. The aperture is terminal and seven-chambered stage at different horizons, and
situated at the peripheral edge of the last chamber the microspheric forms may show a small peak at
in the young stage, and at the angle of the chevroned nine or more. Despite the relatively wide variation
chambers in the adult. It appears to be radiate. in number of planularianchambers (three to nine in
Recognition of the generic charactersis, therefore, a the megalospheric generation), there is clearly no
relatively simple matter, but this is often not the case indication that more than one species is present.

TEXT-FIGURES 8-12
8, Falsopalmula,showing variation in the amount of overlap of the first equitant chamber on the dorsal wall of the test,
from sampleat 49-50 feet. 9, Falsopalmula,
two specimensillustratingdifferentmethodsof formationof the equitant
chambers;gradationsoccur between the two. 10, Falsopalmula cf. deslongchampsi
(Terquem),showingvariationin
coiling amongst megalospheric forms, all x ca. 39. 11, Falsopalmulacf. deslongchampsi (Terquem), drawings showing
some of the variation that occurs amongst small individuals toward the top of the section; shape of proloculum and of
early chambers, and mode of formation and degree of overlap of chevron chambers are particularly variable. 12,
Falsopalmulacf. deslongchampsi(Terquem), showing variation in the size of the proloculum at two levels; each point
represents one specimen; the rarity of microspheric individuals is notable.

215
 ADAMS

Number of chevron-shaped chambers: This varies ferred to above (text-fig. 10) at least two "species"
between one and seven in the present material. can be recognized. Specimen 3 could be assigned to
Brief consideration reveals that this is an unsatis- Falsopalmuladeslongchampsi (Terquem), and speci-
factory distinguishing feature because it obviously men 6 to Falsopalmula semicristellaria(Schwager).
reflects the growth stages of the individuals. As it
is difficult to know whether a particularspecimen is Completely uncoiled specimens, such as number 6
fully developed or not, and as changes in the en- in text-figure 10, are fairly common in the Upper
vironment may possibly limit growth, the matter Lias, occurring as end members of a gradational
becomes complex. However, counting the number series in coiling. Cretaceous and later specimens ex-
of equitant chambers and plotting them against the hibiting a similar morphology, i.e., an initial
number of planularian chambers developed pro- "vaginuline" stage followed by equitant chambers,
duced a normal scatter-diagramwhich gave no indi- are placed in a separate genus, FlabellinellaSchu-
cation (by separate concentrations of points) that bert. Individuals resembling Flabellinella in the
more than one species was present. Upper Lias are certainly not generically distinct,
The equitant chambers although they may represent the beginning of a
Amountof overlapof chevrons: new stock. Macfadyen (1941, p. 58) notes in his de-
may overlap the earlier chambers either on the
on the side of the test, scription of Flabellina centro-gyrataTerquem that
apertural or non-apertural "Flabellina"semi-cristellaria Schwager, found in the
and sometimes on the two together. In the present
Dogger, differs from the former only in its degree
instance, overlap onto more than one "planularian" of coiling. He did not, therefore, include it in his
chamber is common only on the apertural side.
synonymy, but accepted the difference as being of
Inspection of a group of forms from any level specific importance. Similar considerations have
generally revealsthat the amount of overlap is highly affected other authors; Loeblich and Tappan (1950,
variable, as shown in text-figure 8. Here the first p. 58) state, regarding Falsopalmulaantrorsa,that it
equitant chamber overlaps from one to three plan- "... differsfrom FlabellinacentralisTerquem in hav-
ularian chambers. At one horizon, overlap over
ing a tighter coil. It is about one-third smaller than
four planularian chambers was seen. Flabellinacentro-gyrata Terquem, has more oblique
The equitant chambers may form in two ways. The sutures and a less well-developed coil. Flabellinaob-
most common method involves a sudden change in liqua Terquem is somewhat similar, but has a more
growth pattern with resultant overlap of one arm enrolled early stage...." These authors lay em-
of the chevron down the apertural side of the test phasis on the condition of the coil, but they do not
(text-fig. 9a). The second type is formed by the say how many specimens were found or how much
variation existed amongst them. The description
gradual inward curvature of the apertural wall so
that there is no abrupt alteration in the mode of and figure given would fit a number of the speci-
growth (text-fig. 9b). A specimen of the latter was mens from Risby-Warren. It is therefore evident
figured by Terquem (1864, pl. 10, fig. 13) as Fla- that the amount of coiling shown by an individual,
bellinadeslongchampsi Terquem. if considered alone, may be of no value in determin-
ing its species.
The time of onset of growth of the chevron cham-
bers is, as stated earlier, highly variable. It may Thesutureangle: The angle at which the dividing
occur after three ordinary chambers or after as sutures cross the test has sometimes been thought to
many as twelve. It may follow a tightly coiled or be of diagnostic value (e.g., Macfadyen, 1941) in
completely uncoiled initial part, or it may appear differentiating between Falsopalmulaobliqua and
after any intermediate stage between the two. Falsopalmulacentro-gyrata. Examination of numerous
specimens has convinced the author that in the
Degreeof initial coiling:In the structure of its initial case it is a feature of little value. In other
does not differ from Planularia,and present
coil, Falsopalmula individual variation is lesspronounced,
the remarks made under that genus apply equally regions,where
as Barnard (1950a, p. 350) states is the case in
to the present one.
the Lower Lias of the Dorset area, then the obliquity
Text-figure 10 indicates the variation in coiling of the sutures may be a feature of more significance.
that exists at one level, and the number of specimens It was unfortunately not found possible to analyse
with the same amount of coiling at that level can this variation quantitatively because of the difficulty
be seen in text-figure 7. The degree of coiling has involved in measuring the angles accurately in
been used many times as a means of delimiting small specimens and also because the shape of in-
species, and it thus happens that in the series re- dividual chambers is variable.

216
 UPPER LIAS FORAMINIFERA

Ornament: Ornamented examples of Falsopalmula are Lateralcompression of the test: This character raises
not known to occur in the Upper Lias of England. fundamental questions concerning the status of
Such species as Falsopalmula tenuistriata
(Franke) and the genera Falsopalmula and Planularia.Regarding the
"Flabellinainsignis"Terquem and Berthelin appear former, this investigation yielded a few forms, the
to be characteristic of the continental Lias but to initial coiled part of which is arched in cross-section
be lacking in Britain. It is interesting to note that and lenticuline in appearance. These were assigned
striate specimens of Planularia are also virtually to the genus Lenticulina,as they were evidently
absent. variants of one of its species. Barnard (1950b) noted
a similar phenomenon among specimens from the
Shapeof thechambers: The shape of individual cham- Upper Lias of Byfield.
bers often varies considerably within a single speci- In the case of Planularia,increase in the arching of
men and cannot be consideredof much significance. the walls leads to the production of forms which can
Modification during growth, due to a response to be referred only to the genus Astacolus.Such speci-
an external stimulus, may be responsible for many mens have been found. Brotzen (1953, p. 36) states
individual peculiarities. Particularly noticeable is that "... all lagenid genera, as early as the Triassic
the tendency for a small chamber to be inserted and Jurassic, were divided into a number of very
occasionally between larger ones. The chambers of different groups, which had very constant charac-
megalospheric forms appear to be more variable in ters at that time. One of these constant characters
shape than those of microspheric forms, but few of is the shape of the cross-section, which may be
the latter are ever available for examination. Text-
rounded, oval, triangular or rectangular." To the
figure 11 shows some of the variation that occurs at present author this statement is unacceptable.
one level high in the succession.
Taking but a single instance, that of Planulariaand
Astacolus,it is evident that if the cross-sectionof the
Shapeand size of proloculum:In the past it has been former always revealed parallel sides, and that of
the practice of authors to state the shape of the the latter biconvex sides, no difficulty would be
proloculum when describing a new species; terms experienced in distinguishing between them. In
such as oval, elongate, globular, elliptical, egg-
point of fact, as Bartenstein pointed out (1948c,
shaped, etc., are in common use. However, except p. 52), transitions between the two do occur, and
in the case of a spherical proloculum, the shape
they are often difficult to separate.
appears to vary according to the angle from which
it is observed, and this is usually determined by the Depthof the sutures:This feature is probably not of
arrangement of the following chambers. Examina- very much significance. Slight variation does occur
tion has shown that, although a great variety of between individuals, and Franke (1936) used the
difference in depth of the sutures to distinguish
shapes are present, most individuals have spherical
or oval prolocula with various modifications usually between Flabellinadeslongchampsi Terquem and Fla-
caused by a flattening of part of the wall (see text- bellinajurensisFranke. Later workers have not ac-
fig. 11). cepted these differences as being of diagnostic im-
portance. There is a general tendency amongstmany
In the present case, proloculum size was found to be lagenid genera and species, including Falsopalmula,
of more significance than shape, though here again for the sutures over the later part of the test to
a continuousgradation can usually be found between become slightly deepened relative to the earlier
the smallest and the largest in any group. Generally sutures.
the diameter varies from 0.01 to 0.10 mm. Text- Remarks:The foregoingobservationshave been made
figure 12 shows the variation in proloculum size at on a very variable suite of specimens which, despite
two levels. The absence of the microspheric form their variation, appear to belong to a single species
is evident in the second diagram. of the genus Falsopalmula.A very constant and re-
markable feature of the genus is its association with
A relationship between the size of the proloculum Planularia.This has been noted previouslyby Barten-
and the degree of coiling is evident after inspection stein (1948b), who observed the similarity between
of a few specimens. All of the microsphericforms are Planulariacordiformis (Terquem) and the initial part
tightly coiled (eight or more chambers taking part of Falsopalmuladeslongchampsi (Terquem), and be-
in the coil), whilst the megalospheric forms vary tween Planulariacrepidula(Fichtel and Moll) and
greatly in their degree of enrolling. A feature of Falsopalmulaobliqua (Terquem). He deduced that
these assemblagesis the great rarity of microspheric in each case the species of Falsopalmulawas derived
individuals. from the corresponding species of Planularia. In

217
 ADAMS

180- *4

160-
.4

+9
140-

5 .7 .8
120-

?b-b

.4
100- .3 +5
PROLOCULUM
L.XD. +4 .4 5 .5 .5 .6 .7 .9 +8
*7 *7
80- +5 .5 ,5 .6
+4 *6
.9
.5 4S .S
. ^+ .7 +9
60- 6
*4 *4 +5 +5 +b +.6 +7 *7
*6 +7 *8
08
*9 +-8
40- .9

20-

,lI 013 .i6

0O +17

I I
0o3 0;4 o;5 o06 0:7 0:8 0:9 1:o 1:1 1:2 1: 1-4

LENGTH OF TEST IN NM. TEXT-FIGURE 13a

+4 +5
180-

160- 9
5 4-
.7

140-
+b +8
+s
120- .6

+5 .7 +7
+ .5
+. .b
100- ?10
*8
PROLOCULUM
+7 ,7. 1.8 .10
L.X D. .9
+6
80- + .5

b +7 .8.7 .10
6 ,.7 e7 .8
.7 48 *8 +8
60- 46 *8 48.8 .9
b +7 +7 .9 .9 .II
08 e8 +8
46.6
0+ 6 8 4*7 .14
.9
o8 4?8 ?
40- . .10 .11
9
*10

20-

0- .13 .144L34J4 .j3 .J 5.4 .I6

0o3 0;4
I o:5 C) 6 0'7 0o8 0;9 I'o Tl' 1'2 1'3 1-4 I:'
LENGTH OF TEST IN MM.
Marginulinaprima * TEXT-FIGURE 13b
Marginulinaprima var. spnata +

218
 UPPERLIASFORAMINIFERA

180

.7 *7

160

.4 ,7
140-

+5 .5 +$

.7
120-

100- +4 .5
PROLOCULUM
+5 .6 .7 +7 .9
L. XD.
*6 ,7
80- +5.5 4+6 + .7 +7 45
*5 b5 .+6 46 .7 .10
.5 15 .5.6 .6 4 .7

.5 .5 +6 .7 .9
60-
.55 .6 6 7+>t.7. 7+
+9
*9

+5 +7
40-
*10 .10

20-

+10 ,
4,11 J4)
1 412 43 *15
0-

. . w . . w
w w w w . .

0:3 0:4 0o5 0o6 0-7 08 09 10 11 12 1?3 1:4

LENGTH OF TEST IN MM.

Marginulina primao
Morginullno prima var spinoto + TEXT-FIGURE 13c

TEXT-FIGURE 13
Marginulina primad'Orbigny,distributiondiagramsshowing dimorphism.The length of the test is plotted against
the horizontalscale,the productof the length x breadthof the proloculum(x 1000) againstthe verticalscale. The
numberbesideeach point indicatesthe numberof chambersin the test. Microsphericand megalosphericformsare
clearly separated,the latter being highly variable:a, at 51-52 feet; b, at 53-54 feet; c, at 55-561 feet.

northern Lincolnshire, Falsopalmulais associated champsidoes not show an initial stage resembling a
with Planularia pseudocrepidula Adams, n. sp., a typical Planulariacordiformis(Terquem) as Barten-
species already stated to be very variable, and it is stein indicates, so that the nomenclature remains
interesting to see that the variation in coiling seen very involved.)
in Planulariapseudocrepidula can be exactly matched Conclusions: No single character has been discovered
by that of Falsopalmula. It would seem from this that which, in itself, is sufficiently stable to be of great
the specimens of Falsopalmulafrom the Upper Lias value in delimiting species. The same general
of northern Lincolnshire should be referred to the remarks concerning the futility of erecting species
species Falsopalmulaobliqua(Terquem), yet they do on too few specimens apply here as in the case
not exactly match those described under this name of Planulariadiscussedearlier. The variation seen in
by Bartenstein.Actually, they more closely resemble Falsopalmulaparallels that of Planulariaso closely as
Terquem's original figure of Falsopalmuladeslong- to suggest that, bearing in mind Bartenstein'searlier
champsi,and their range of variation is very great. evidence, almost any species of Planularia,at least
These differences, however, may perhaps be ac- in the Upper Lias, may produce equitant chambers
counted for by minor differencesin the environment and thus become a Falsopalmula.The similarity in
affecting the growth of the tests. (It should be noted coiling between the two genera is very significant
that Terquem's original figure of Flabellinadeslong- in this connexion.

219
 ADAMS
C2D

40- ss'-s

u
30- 5
w

e0
40 A 5354' 20-

/NTEXT-FIGURE \ 15a TEXT-FIGURE 15b

z /

S 6 7 B 9 10 II
m - X /

/ -
50 3/- Y \ 30-
z u
w / 0a.
u Lo

u 30- U
0. 10- c I

U. 20- 1 /
O z

N
mr II- 0'7 0.9 1'I 1'3 I'5 1'7 1'9 2.1 2.3
I:0-

Z O SLENGTH BREADTH RATIO 1: 0-7 1:23

LINGULINA LONGISCATA

5 6 7 8 9 0 II VAR

NUMBER OF RIBS TEXT-FIGURE 16

50-

40- 1-52f 5
number idctshttettwsbonadt
nubrocabrucrann)ir ifo saprt7
40.
~~~~Z~~~~~~~~~~~~~ 7
w n 1
30-x
U 30- .4 ,7
0 .5
t .
20- z 6 4
LL ' 20- .9
-- \ 6 77.76 .7 ..*
I5:5 S
.8 16 I
: .. . e:7
a: '0- -6-6 * 79
wu lo- .6 , 7

0 -

II . *
5 6 7 a 9 10 II 0: 215 0 :2 025 0:3 035 04 04 s

NUMBER OF RIBS LENGTH OF TEST IN MM.

TEXT-FIGURE 14 TEXT-FIGURE 17

TEXT-FIGURES 14-17
14, Marginulinaprima d'Orbigny, frequency polygons showing variation in number of ribs at three different levels.
15, Lingulinalongiscata(Terquem): a, external appearance, x 70; b, interior of chambers showing siphons viewed by
transmitted light after oil immersion. 16, graphs showing the difference in length: breadth ratio of Lingulinalongiscata
(Terquem) and Lingulinalongiscata(Terquem) var. alpha Adams, n. var.; these graphs include the full number of
specimens from all levels. 17, LingulinateneraBornemann, distribution diagram showing variation in the megalospheric
form at 51-52 feet; the number beside each point indicates the number of chambers in the test; the absence of a
number indicates that the test was broken and the number of chambers uncertain; no microspheric forms appear to
be present.

2o0
 UPPER LIAS FORAMINIFERA

d'Orbigny, emend.
Genus MARGINULINA initial coil in which four or five chambers take part. The
Marie, 1941 test then increases rapidly in size, the adult stage being
similar to that of the megalospheric form.
Margimilinaprima d'Orbigny The microspheric and megalospheric individuals are
Text-figures 13-14, 18-20 quite distinct, no specimens with prolocula of inter-
Marginulinaprima D'ORBIGNY, 1850, Prodrome Pal., vol. 1,
mediate size having been discovered. Text-figure 13a-c
p. 242, no. 262. - MACFADYEN, 1936, Roy. Micr. Soc. shows quite clearly the separation of the two forms.
London,Jour., ser. 3, vol. 56, p. 151, pl. 1, fig. 262a. - Variation:This species exhibits considerable variation
FRANKE, 1936, Preuss.Geol. Landesanst.,Abh., new ser.,
no. 169,pl. 8, figs. 1-7 (includingthe formsrugosaBorne- both vertically through the succession and among
mann, gibbosa,recta,acuta,and ornataTerquem, and prae- specimens obtained from a single sample. The distri-
longa and gibberulaTerquem and Berthelin). - BARTEN- bution diagrams (text-fig. 13a-c) indicate the range of
STEIN AND BRAND, 1937, Senckenb. Naturf. Ges., Abh., variation which occurs and also show that it is the
no. 439, p. 161, pl. 2B, fig. 26; pl. 3, figs. 39-40; pl. 4, megalospheric form which is responsible for the greater
fig. 60a-b; pl. 5, fig. 46a-b. - MACFADYEN,1941, Roy. part. In spite of the range of size shown by the megalo-
Soc.London,Philos.Trans.,ser.B, vol. 231, no. 576,p.38, sphere, there is no indication of trimorphism.
pl. 2, fig. 32a-b. - BARNARD, 1950, Geol. Soc. London,
Quart.Jour., vol. 105,pt. 3, p. 372, text-fig.5a-b, g. At 53-54 feet there is a tendency for very elongate forms
Marginulinaraphanus(Linn6). - BRADY, 1867, Somersetshire
to develop; these are fairly robust but slender. At 51-52
Archaeol.Nat. Hist. Soc., Proc., vol. 13, pt. 2, p. 225, feet a shorter, broader form is more in evidence. This
pl. 2, fig. 21. - CRICKAND SHERBORN,1891, Northampton- particular variant increases rather rapidly in width and
shireNat. Hist. Soc., Jour., vol. 6, p. 211, pl. 1, fig. 18. appears conical when viewed from the side. At higher
MarginulinaburgundiaeTerquem. - ISSLER, 1908, Palaeonto- horizons, individuals are reduced in number, probably
graphica, vol. 55, p. 67, pl. 4, figs. 171-174. - FRANKE, owing to an environmental control. At 51 feet there is
1936,Preuss.Geol. Landesanst.,Abh., new ser., no. 169, a change in lithology, grey micaceous shale giving way
p. 78, pl. 8, fig. 8. to a slightly harder non-micaceous shale, whereupon
MarginulinaincisaFRANKE,1936, ibid., p. 78, pl. 8, figs. 11-12. the number of individuals is immediately reduced.
Description:The megalospherictest consistsof a recti- A certain amount of variation occurs in the number of
linearseriesof chambersup to fourteenin number.In- ribs present on each specimen. In general, the number
dividualspossessingsix to nine chambersare the most is dependent on two factors:
common. The proloculum is subspherical,generally
a) The age of the specimen, i. e., the number of cham-
apiculate,and is followedby chamberswhich increase bers it has formed. Later chambers tend to be larger
fairly regularlyin size, the rate of increasedepending and to possess more ribs.
on the size of the proloculum.Specimenswith large
proloculatend to increasebut little in width with age, b) The size of the proloculum. Megalospheric forms
many remaining almost parallel-sided.The chamber generally possess more ribs than microspheric
walls are often very thick; the septa are usuallytrans- forms because at a given chamber the megalospheric
individual is usually the larger. Text-figure 14 shows
verse, but in some cases slope obliquely toward the
the variation in the number of ribs per specimen at
non-aperturalside. In such forms the test is often three levels where material was abundant.
slightly arcuate, the aperturalside being convex. The
apertureis terminal,radiate,raisedon a shortneckand Remarks:The large number of names under which this
offset to one side of the test. If the extremetip of the species has been recorded reflects the wide range of
apertureis missing,the inside of the neck is seen to be variation which it shows. Franke (1936) recognised
circular. seven forms of Marginulinaprima (Marginulinaprima
Ornamentationconsistsof a numberof strongribswhich formae rugosa,gibbosa,praelonga,recta,acuta,gibberula,and
traversethe test longitudinally.These crossthe sutures ornata), these names having been used earlier as
withoutinterruptionand continuedown to the proloc- specific names. The enumeration of such forms appears
ulum. The ribs vary from five to ten in number, but to be quite pointless, especially in the present instance,
when all occur together within a very few feet of strata.
usually there are about seven, fresh ones being added Franke apparently applied different "form" names to
as the test increasesin circumference.Each rib may
remainindependentover its whole length or may fuse megalospheric and microspheric individuals. Macfadyen
with its neighbourson the upper surface of the last (1941) included many previously recognised "species"
chamber. In most cases the suturesare flush, but in under the name Marginulinaprima.
some adult specimensthereis a tendencyfor the cham- This is believed to be the first record of the species from
berjunctionsto become incised, thus causingthe later the Upper Lias of England. Bartenstein and Brand did
chambersto appearmoreglobular. not record it from any strata higher than Lias 8 in
The microsphericform consists,when fully grown, of Germany (= spinatumzone in Britain).
from eleven to sixteen chambers. Usually there are Types:Hypotypes, B.M.N.H. no. P.43378.
about thirteenor fourteen.A minute proloculum(ap- Material:Approximately 200 specimens from the Upper
proximatediameter0.01 x 0.01 mm.) is followedby an Lias of northern Lincolnshire.

221
 ADAMS

Marginilina prima d'Orbigny var. spinata Terquem does not invariably occur, but, on the other hand, no
Text-figure 21 specimens having the sutures flush with the surface have
been observed to possess interrupted ribs.
MarginulinaspinataTERQUEM,1858, Acad. Imp. Metz, MWm.,
vol. 39, p. 615 (55), pl. 3, fig. 8. - MACFADYEN,
1941, Types:Hypotypes, B.M.N.H. no. P.43379.
Roy. Soc. London, Philos. Trans., ser. B, vol. 231, no. Material:One hundred sixty specimens from the Upper
576, p. 39, pl. 2, fig. 33a-b. Lias of northern Lincolnshire.
Marginulina interruptaTerquem forma spinata Terquem. -
FRANKE, 1936,Preuss.Geol. Landesanst.,
Abh., new ser., Genus Lingulina d'Orbigny, 1826
no. 169,p. 79, pl. 8, fig. 10.
Marginulina spinata spinata Terquem. - BARTENSTEIN
AND Lingulina is widespread throughout the Upper Lias
BRAND, 1937, Senckenb.Naturf. Ges., Abh., no. 439, of Lincolnshire, and at least seven described species
p. 161, pl. 4, fig. 61; pl. 5, fig. 47. have been recognized during the present investiga-
Description:The test is similar in general appearance to tion. Confusion has always existed concerning the
that of Marginulinaprima, but differs from the latter in diagnosis of the genus, the trouble arising from a
that the ribs are interrupted at the sutures and tend very poor original description by d'Orbigny (1826).
to project downward as little spines. In some specimens A full discussion of the nomenclatural problem is in
it is only the lower part of the test which develops this the course of preparation, and it is sufficient to state
spinate type of ribbing, whereas in the upper portion here that the author has included in the genus Lin-
the ribs are complete.
gulina all lagenids possessing a rectilinear series of
Remarks: It has previously been the custom of all chambers, often strongly compressed laterally, which
authors to separate Marginulina spinata from Marginulina have the septa arranged at right angles to the long
prima as a distinct species, the difference in the character axis of the test or which are only weakly chevron-
of the ornament having been considered a diagnostic
shaped, and in which the aperture is central, ter-
feature. Careful examination of the present material has
minal, elongate, slit-like or oval. Although all the
led the author to conclude that such a separation is not
characters must be considered when determining
warranted on this occasion. Apart from the ornament,
there are no features by which the "species" may be the genus, the slit-like nature of the aperture seems
to be the most important. Of the species investigated,
distinguished. Measurements of the size of the proloc-
ulum and length of test, when plotted together with the Lingulina longiscata (Terquem) proved to be one of
number of chambers (see text-fig. 13a-c), do not reveal the most interesting.
any differences, nor is the total number of ribs any differ- Lingulina longiscata (Terquem)
ent (text-fig. 14). Many individuals are found in which
only the lower part of the test bears incised, spinous ribs, Text-figures 15-16, 23-24
and in others the ribs are not completely incised but Lingulinacarinatad'Orbigny. -JONEsANDPARKER, 1860 (part),
are saw-shaped. Franke (1936) described the ribs of Geol. Soc. London,Quart.Jour., vol. 16, p. 453,.pl. 19,
Marginulinainterruptaforma spinata as saw-shaped, the fig. 15 only.
rib section increasing in height from the upper to Frondicularia
striatulaReuss. - JONESANDPARKER,1860, ibid.,
the lower suture. Macfadyen (1941), in describing the p. 453, pl. 19, figs. 17-18.
Frondicularia
longiscataTERQUEM, 1870, Acad. Imp. Metz, MWm.,
species Marginulinaspinata, had only two specimens to vol. 51, p. 318, pl. 22, figs. 23-24.
which to refer. These possessed fifteen and thirteen FrondicularianodosariaTERQUEM,
1870 (part), ibid., p. 319,
medium to small costae, respectively. The number of pl. 22, fig. 25 only.
ribs possessed by the present specimens never exceeds lanceolata(Hausler). - ISSLER,1908 (part), Palae-
Frondicularia
ten, and more usually is seven or eight. In his original ontographica,vol. 55, p. 54, pl. 2, fig. 103 only.
mesoliassicaBrand. - BARTENSTEIN
?Frondicularia
description, Terquem stated that seven weak ribs were AND BRAND,
1937(part),Senckenb.Naturf.Ges.,Abh.,no. 439,p. 158,
present.
pl. 4, fig. 66 (not p. 159, text-fig.16).
In view of the extreme difficulty experienced in distin-
Description:A very fragile test formed of up to twelve
guishing between many of the specimens, especially chambers in a rectilinear series. Although laterally
when they are found with the outer edges of the ribs
strongly compressed, in cross-section the sides are
broken, and because both types have the same range, rounded. Growth, whilst fairly regular in some speci-
it is not considered that the spinate forms merit more
than varietal status. The proportion of Marginulina mens, is sometimes irregular, alternately large and
small chambers tending to develop. The later chambers
prima to Marginulina prima var. spinata was found to vary often tend to decrease slightly in width, giving the test
between different levels, but as they are greatly reduced a lanceolate appearance. The proloculum is either
in numbers above 50 feet, little signifance can be
attached to this fact. elongate or subspherical, and its junction with the
second chamber is perfectly straight. The aperture is
It has been noted that there is a tendency for specimens central, terminal, oval or slit-like, and flush with the
to have discontinuous ribs whenever the chamber surface. Usually it is very difficult to see. In rare cases,
junctions become moderately or strongly incised. This when the end chamber is particularly pointed in outline,

222
 UPPER LIAS FORAMINIFERA

21

18

19
. .

22
20

:fv

. '.

; ... t.,

: i ..
.. . .
.E .. . r

)? ,.~
*
.. ^ '.^
.,
tr-
.s ' -
25','i .r
-:
.

-jI

?
r-I * t2J

23 24 ..J L26

TEXT-FIGURES 18-26
18-20, Marginulinaprima d'Orbigny, x ca. 76:18, microspheric form; 19-20, megalospheric forms. 21, Marginulina
primad'Orbigny var. spinataTerquem, x ca. 76. 22, LingulinateneraBornemann, x ca. 105. 23-24, Lingulinalongiscata
(Terquem), x ca. 105. 25-26, Lingulinalongiscata(Terquem) var. alpha Adams, n. var., X ca. 105: 25, an excep-
tionally broad individual.

223
 ADAMS

the aperture approaches a round condition. The sutures Material:One hundred thirty-seven specimens from the
are chevron-shaped, except in the early stages of some Upper Lias of northern Lincolnshire.
specimens, where they are straight.
When mounted in a clarifier and viewed with trans- Lingulina longiscata (Terquem) var. alpha Adams,
new variety
mitted light, a short "siphon" can be seen projecting
down into the lumen of each chamber from the aper- Text-figures 16, 25-26
tural margin (text-fig. 15). In longitudinal sections, this cf. Lingulina (Terquem). - BARNARD, 1950, Geol. Soc.
nodosaria
structure resembles a tiny funnel with its walls concave London,Quart.Jour., vol. 106, pt. 1, p. 29, text-fig. 16.
inward. The aperture itself is often blocked by a calcite
Diagnosis: Test fairly small, formed of up to twelve
plug. The author knows of no other record of this small chambers, similar in many respects to Lingulinalongis-
structural feature in Liassic species of Lingulina,but this
cata,but differing as follows: The numerous striae which
may be due to other workers having examined the tests ornament the test tend to diverge downward. This is
in the dry condition. Buchner (1942) has described caused by the greater width of the test and is not a
somewhat similar structures in living representatives fundamental difference. The test increases in width
of the genus. He calls the downward projections
fairly rapidly with age, so that the chambers in adult
"siphons," and in his figured specimens they are often life are considerably broader than long. Internal
very long and quite straight. "siphons" are present in some specimens but are very
The test is ornamented externally by numerous very small; in others they appear to be lacking. In fully
fine striae running longitudinally. In some cases they grown specimens, the later chambers tend to be reduced
are absent over the upper part of each chamber, a con- in width. The length: width ratio of the adult chambers
dition observed by Bartenstein and Brand (1937) in varies from 1:1.4 to 1:2.9. In text-figure 16, these
Frondiculariamesoliassica,which "species" is, in part, ratios are compared with those of Lingulinalongiscata.
very similar if not identical to this. Dimensions:Length up to 0.63 mm.; width up to 0.19 mm.
Measurements of the length and breadth of the cham- Variation:Irregularities of growth are common. A few
bers in the adult stage reveal that the length:breadth increase regularly in size; in others the increase is rapid
ratio always lies within the range 1:0.8 to 1:1.4. This at first, little further change occurring later. Small
may be compared with the ratios calculated for Lingulina chambers are sometimes intercalated amongst the
longiscatavar. alpha (text-fig. 16). larger.
Dimensions: Length up to 0.63 mm.; width up to Remarks:Barnard (1950b, p. 29) described and illustrated
0.14 mm. a form from Byfield which he called Lingulinanodo-
Variation:In a few specimens, apparently microspheric saria (Terquem). The present author feels that this form
should probably be included under Lingulinalongiscata
forms, the ornament is absent from the first two or three
chambers. These first-formed chambers, which are var. alpha.
longer than broad, are hardly, if at all, chevron-shaped. These specimens are regarded as a distinct variety of
In view of this, and because of the elongate nature of Lingulina longiscata because of the difference in the
the aperture, it is considered that these forms are best length:width ratios of the chambers, and because one
assigned to the genus Lingulinarather than to Frondicu- replaces the other in the succession. There is an interval
laria. of 28 feet in which only rare specimens of either type are
Remarks:Terquem (1870, p. 318) recorded this species present. The author cannot explain the bimodal appear-
as Frondicularialongiscata,from the Middle Jurassic of ance of the graph (text-fig. 16) for this variety. It is not
France. He stated that it was "rather rare." In the caused by the presence of megalospheric and micro-
same work, he figured specimens of Frondicularianodo- spheric forms.
saria Terquem, the first of which (pl. 22, fig. 25a-b) Types: Holotype, B.M.N.H. no. P.43380. Paratypes,
appears to be another example of Lingulinalongiscata. B.M.N.H. no. P.43381.
Jones and Parker (1860) illustrated two specimens which
probably belong here, and Issler (1908, pl. 2, fig. 103 Material: Fifty specimens from the Upper Lias of
northern Lincolnshire.
only) figured a specimen which appears identical to
these. This he called Frondicularialanceolata(Hausler),
the original of which is a smooth form. The striae are Lingulina tenera Bornemann
in many cases difficult to see, and may have escaped Text-figures 17, 22
Issler's attention. Bartenstein and Brand (1937, pl. 4,
Lingulinatenera Bornemann,1854,Liasform.Gottingen,p. 38,
fig. 66) show a broken specimen, which they call Fron- pl. 3, fig. 24a-c.
diculariamesoliassicaBrand. In the description of this
"species," no mention is made of the aperture, and the This species and its allied forms will be treated fully in a
text-figure (p. 159, text-fig. 6) looks decidedly different separate paper. It is introduced here because it exhibits
from the specimen on the plate. considerable morphological variation, one aspect of

224
 UPPER LIAS FORAMINIFERA

which,namely,theshapeofthetest,is largelydetermined of Marginulina,very clear dimorphism could be

by the size of the proloculum. Microsphericindivid- seen in every sample. It may perhaps be suspected
uals have the lower part of the test pointed in outline, from these results that the arrival of the megalo-
whereasmegalosphericindividualshave the test more spheric generation is followed by the production of
parallel-sided. a number of similar generations, the later individu-
Measurementsmade of proloculumsize and length of als of which tend to increase the size of the pro-
test at differentlevels of the same core in the Middle loculum. Forms having very large prolocula are,
Lias seem to indicate that the two generationsdo not however, few in number.
gradeinto one another.The distributiondiagram(text-
It may be that in some genera and species there is
fig. 17) is typical of the conditionfound at all levels in
the Upper Lias, and is notablemainlyfor the complete no clear-cut distinction between the proloculum
absenceof microsphericindividuals. size of megalospheric and microsphericindividuals.
This is suspected in the case of certain species of
Frondiculariaand Nodosaria,but further work is re-
SUMMARY AND CONCLUSIONS
quired before this can be confirmed.
The detailed examination of some Upper Lia
BIBLIOGRAPHY
genera and species of the Lagenidae illustrates the
variation which occurs in their morphology. Appli- BARNARD,T.
cation of several different criteria which have been 1950a Foraminifera from the Lower Lias of the Dorset coast.
Geol. Soc. London, Quart. Jour., vol. 105, pt. 3,
used for the establishment of species, as in the case pp. 347-391, text-figs. 1-11.
of Falsopalmula,has shown that wherever a large 1950b Foraminifera from the UpperLias of Byfield,JNorthamp-
number of specimens are investigated, grading tonshire.Geol. Soc. London, Quart. Jour., vol. 106,
series in different morphological characters can be pt. 1, pp. 1-36, pls. 1-3, text-figs. 1-17.
1956 - SomeLingulinaefrom the Lias of England. Micropale-
established. This leads to the conclusion that, in
ontology, vol. 2, no. 3, pp. 271-282, pls. 1-3.
defining a species, care must be taken to indicate BARTENSTEIN, H.
the complete range of variation exhibited by its 1948a Untersuchungen zur Systematik und Stratigraphieder
component individuals, otherwise an unnecessary Flabellina-gestaltigenForaminiferen.Erdol und Kohle,
vol. 1, no. 6, pp. 180-184, text-figs. 1-14.
multiplication of specific names will result. 1948b TaxonomischeAbgrenzungder Foraminiferen-Gattungen
Statistical investigation, involving the measurement Palmula Lea, Flabellina d'Orbigny und Falsopalmula
of proloculum and test size in thousands of individ- n. g., gleichzeitigeine RevisionderJura-Artenvon "Fla-
bellina."Senckenbergiana, vol. 28, no. 4-6, pp. 119-
uals, indicates that the megalospheric form is re-
137, pls. 1-2, text-figs. 1-5.
sponsible for by far the greater part of the variation, 1948c Entwicklungder GattungLenticulina(Lenticulina)La-
and an examination of the older literature reveals marck1804 mit ihrenUnter-Gattungen (For.). Sencken-
that the majority of species were erected on megalo- bergiana, vol. 29, no. 1-6, pp. 41-65, pls. 1-6,
11 text-figs.
spheric forms alone. Not only this, but the number 1949 - Neoflabellina,nomennovumpour Flabellina d'Orbigny
of specimens was very often small, numerous cases 1839 (foraminiferes).Soc. Geol. France, C. R. Somm.,
of "rare," "very rare," or "single specimens" oc- no. 9, pp. 164-166.
curring in the literature. Nevertheless, as it is some- BARTENSTEIN, H., AND BRAND, E.
times impossible to find microspheric individuals, 1937 - Mikro-paldontologische zur Stratigraphie
Untersuchungen
it becomes all the more important that the complete des nordwest-deutschen Lias und Doggers. Senckenb.
Naturf. Ges., Abh., no. 439, pp. 1-224, pls. 1-20,
range of variation of the megalospheric generation
should be given. (It is not suggested that the micro- text-figs. 1-20, 5 tables.
BLAKE,J. F.
spheric forms do not vary, but in most cases their 1876 - Class Rhizopoda. In: TATE, R., AND BLAKE, J. F.,
rarity makes it impossible to study their variation.) The rorkshireLias. London: J. van Voorst, pp. 449-
It has been averred by Hofker (1951) that trimor- 473, pls. 17-19.
J. G.
BORNEMANN,
phism occurs in the early stages of a family's history. 1854 - Oberdie Liasformationin derUmgegendvonGottingenund
The Lias deposits represent an early period in the ihre organischenEinschliisse.Berlin: A. W. Schade,
evolutionaryhistoryof the familyLagenidae (Trias- pp. 1-77, pls. 1-4.
Recent), and many of its species at that time were BRADY, H. B.
1867 - Foraminifera. In: MOORE, C., On the Middle and Upper
highly variable. Measurements have been made of Lias of the south west of England. Somersetshire
proloculum and test size in species belonging to the Archaeol. Nat. Hist. Soc., Proc., vol. 13 (1865-
genera Marginulina,Lingulina,Frondicularia,Planu- 1866), pt. 2, pp. 220-236, pls. 1-3.
laria and Falsopalmulaand, although only the first BROTZEN, F.
of these is fully discussedin this paper, in no case was 1953 - Problemsin the nomenclature of theforaminifera.Micro-
any sign of trimorphism observed. In the case paleontologist, vol. 7, no. 2, pp. 35-38.

225
 ADAMS

BUCHNER, P. LOEBLICH, A. R., AND TAPPAN, H.

1942 - Die Lingulinendes Golfes von Neapel und der marinen 1950 - North AmericanJurassic foraminifera; I - The type
Ablagerungenauf Ischia. K. Leop.-Carol. Deutsch. Redwatershale (Oxfordian)of SouthDakota.Jour. Pal.,
Akad. Naturf., Abh. (Nova Acta), new ser., vol. 11, vol. 24, no. 1, pp. 39-60, pls. 11-16.
no. 75, pp. 103-145, text-figs. 1-18. MACFADYEN, W. A.
CRICK, W. D., AND SHERBORN,C. D. 1936 - D'Orbigny'sLiasforaminifera. Roy. Micr. Soc. London,
1891 - On some Liassic foraminiferafrom Northamptonshire. Jour., ser. 3, vol. 56, pp. 147-154, pl. 1.
Northamptonshire Nat. Hist. Soc., Jour., vol. 6, 1941 - Foraminifera from the GreenAmmonitebeds,LowerLias,
pp. 208-214, 1 pl. of Dorset. Roy. Soc. London, Philos. Trans., ser. B,
CUSHMAN, J. A. vol. 231, no. 576, pp. 1-73, pls. 1-4, text-figs. 1-6.
1940 - Foraminifera,theirclassification
and economicuse. Ed. 3.
MARIE, P.
Cambridge, Mass.: Harvard University Press, pp. 1941 - Foraminiferesde la Craie:Lesforaminiferesde la Craiea
1-535, pls. 1-31, text-figs., tables, charts; Key, Belemnitellamucronata du Bassin de Paris. Paris, Mus.
pls. 1-48. National Hist. Nat., MWm.,new ser., vol. 12, fasc. 1,
DEFRANCE, M. J. L. pp. 1-296, pls. 1-37, text-figs. 1-7.
1826 - Miniralogie et geologie. In: Dictionnairedes Sciences
ORBIGNY,A. D'
haturelles.Paris: F. G. Levrault, vol. 41 (pin - plo), 1826 - Tableaum6thodique de la classedes Ciphalopodes.Ann.
pp. 1-558; Atlas, Zoologie: Vers et zoophytes; Sci. Nat., ser. 1, vol. 7, pp. 96-169, 245-314, pls.
Conchyliologie et malacologie. 10-17.
FICHTEL,L. VON, AND MOLL, J. P. C. VON
1798 - Testaceamicroscopica 1850 - Prodromede paleontologiestratigraphique universelledes
aliaqueminutaex generibusArgo- animauxmollusques et rayonnis.Paris: V. Masson, vol. 1,
nautaetNautilusadnaturamdelineataet descripta.
Vienna:
Camesina (1803 reprint), pp. 1-124, pls. 1-24. pp. 1-392.
FRANKE,A. PAALZOW,R.
1936 - Die Foraminiferendes deutschenLias. Preuss. Geol. 1922 - Die ForaminiferenderParkinsoni-Mergel
vonHeidenheim
Landesanst., Abh., new ser., no. 169, pp. 1-138, am Hahnenkamm. Naturh. Ges. Nurnberg, Abh., vol.
pls. 1-12. 22, pp. 1-35, pls. 1-4.
GLAESSNER, M. F. TERQUEM, 0.
1945 - Principles of micropalaeontology.
Carlton, Victoria: 1858 - Mlmoiresur lesforaminiferesdu Lias dudepartement de la
Melbourne University Press, pp. 3-296, pls. 1-14, Moselle; Partie I Acad. Imp. Metz, MWm.,vol. 39,
text-figs. 1-64. pp. 563-654 (1-94), pls. 1-4.
HOFKER, J. 1864- Troisiememimoiresur les foraminiffresdu Lias des d6-
1951 - The toothplate-foraminifera.
ArchivesNeerl. Zool., vol. 8 partementsde la Moselle, de la Cote-d'Or,du Rhone,de
(1947-1951), pt. 4, pp. 353-373, text-figs. 1-30. la Vienneet du Calvados.Acad. Imp. Metz, M^m.,
ISSLER,A. vol. 44 (ser. 2, vol. 11) (1862-1863), pp. 361-438,
1908 - Beitrdgezur Stratigraphieund Mikrofaunades Lias in pls. 7-10.
Schwaben.Palaeontographica, vol. 55, no. 1, pp. 1- 1870 - Troisiemememoire surlesforaminifiresduSystemeOolithi-
103, pls. 1-7. que, comprenant les genresFrondicularia,Flabellina,No-
JONES, T. R., AND PARKER, W. K. dosaria,Dentalina,etc. de la zonea Ammonitesparkinsoni
1860 - On somefossil foraminiferafrom Chellastonnear Derby. deFontoy(Moselle). Acad. Imp. Metz, MWm.,vol. 51,
Geol. Soc. London, Quart. Jour., vol. 16, pp. 452- pp. 299-380 (197-278), pls. 22-29.
458, pls. 19-20. T.
WISNIOWSKI,
KUBLER,J., AND ZWINGLI,H. 1890 - Mikrofaunailow ornatowych okolicyKrakowa;Czesi I -
1866 - MikroskopischeBilder aus der Urwelt der Schweiz; Otwornice
grnego Kellowayuw Grojcu.Akad. Umiej. w
Heft II Winterthur, Burgersbibl., Neujahrsbl., Krakowie, Wydz. Mat.-Przyr., Pamietnik, vol. 17,
pp. 1-28, pls. 1-3. pp. 181-242, pls. 8-10 (1-3).

226