Cretaceous Research (1998) 19, 25±42

Taxonomic recognition of species of

Neogastroplites (Ammonoidea, Cenomanian) by
geometric morphometric methods

*R. A. Reyment and {W. J. Kennedy

* Institute of Earth Sciences, University of Uppsala, S 75236, Uppsala, Sweden
{ Oxford University Museum of Natural History, Parks Road, Oxford, OX1 3PW, UK

Revised manuscript accepted 12 August 1997

Species of Neogastroplites display a wide range of morphological variability such that without a priori
knowledge of the stratigraphical location of some individuals, it is not possible to make an
unequivocal speci®c determination on familiar traditional grounds. Using new techniques of
geometric morphometric analysis it is possible to narrow down the margin for error in determining
isolated specimens. We have also demonstrated that there are slight though signi®cant differences in
average lateral shape of the three species studied in this note: N. americanus, N. muelleri and N.
cornutus. If counts of rib-frequencies are admitted into the analysis, the discriminatory effect is
heightened. Unexpectedly, the addition of maximum whorl-breadth, a standard distance measure, to
the analysis has a relatively slight positive effect on the results. A canonical analysis of the full data-set
provides evidence of a gradual shift in shape and ornament over time.
# 1998 Academic Press Limited

KEY WORDS: geometric morphometrics; shape-differentiation; Neogastroplites; evolution; Cenomanian;

North America.

Introduction
The analysis of shape-variation in ammonites by quantitative methods might seem
to be a quite straightforward procedure and one that may be expected to follow
the same course as applications to other organisms. However, the special con-
straint on growth and shape imposed by the logarithmic spiral introduces meth-
odological complications of a statistical nature. The most tangible of these is that
the outline of the shell in lateral aspect is ®xed. In other words it constitutes a
trait that is not free to vary within limits such as is the case for bones of ver-
tebrates and the shells of most invertebrates. Hence, measurements made on the
lateral dimensions of the conch will only yield information on size-variation when
subjected to a multivariate statistical analysis. This situation is further aggravated
when, as is not infrequently done, lateral measurements are standardized by divid-
ing through with the largest measure, usually the diameter of the shell. This
added element inserts the factor of closure whereby the use of standard statistical
methods is ruled out (Reyment & JoÈreskog, 1993).
There is, however, a measurable component of lateral shape variability that can
be brought to the surface by appropriate morphometric methods, particularly
those that are based on geometrical considerations. Bookstein (1989, 1991) has
introduced a well integrated theory of geometric morphometrics which we have
here applied to our data. In very general terms, geometric morphometrics is de®ned
as a class of methods that preserve complete information about the relative spatial
arrangement of the data throughout an analysis. These methods permit the visual-
0195 ± 6671/98/010025 + 18 $25.00/0/cr970094 # 1998 Academic Press Limited
26 R. A. Reyment and W. J. Kennedy

ization of group and individual differences, sample variation, and other results in
the space of the original specimens.
The problem taken up in the present note concerns the following points in
relation to Cenomanian (Cretaceous) material from the Mowry Shale of the Wes-
tern Interior USA and Canada:
(1) The speci®c identity of three Neogastroplites assemblages, separated in time
(and hence presumed not to overlap).
(2) Whether evolutionary changes in the morphology of shape can be recog-
nized.
The three species display a parallel pattern of morphological variability, to wit,
a range of variation from highly compressed, more or less feebly ornate shells,
through to relatively in¯ated, coarsely ribbed, nodate individuals. Without prior
knowledge of the stratigraphical provenance of a particular specimen it is not an
easy matter to identify it speci®cally just by inspection.

2. Material
Reeside & Cobban (1960), in an impressively well illustrated monograph,
described a series of faunal assemblages from early diagenetic calcareous nodules
in beds that form the Mowry, Colorado and Aspen Shales of the Western Interior
USA as well as adjacent, e.g., Alberta, Canada. The Mowry and contempora-
neous formations have yielded freshwater plant and animal fossils, as well as
marine invertebrates in concretions. At a loss to ®nd an explanation of these
concretions, and noting that the surrounding sediment was far from being rich
in fossils, Reeside & Cobban (1960) suggested that they represented faecal
accumulations of an unidenti®ed carnivore.
The ammonite associations exhibit an impressively wide range of variation in
shape, as viewed aperturally, as well as in ornament. The ®ve species recognized
by Reeside & Cobban (1960) were judged to represent ®ve faunal zones currently
regarded as of late Albian and early Cenomanian date. For a discussion of the
Albian-Cenomanian boundary within the Western Interior Neogastroplites-
sequence, see Hancock et al. (1993, p. 461, note 7), and Cobban & Kennedy
(1989). The three Cenomanian species studied here are, in ascending biostratigra-
phical order: Neogastroplites cornutus (Whiteaves) Neogastroplites muelleri Reeside &
Cobban, and Neogastroplites americanus (Reeside & Weymouth).
The variation in form displayed by each of these species ranges over com-
pressed costate, stout nodose, and subglobose spinose. Reeside & Cobban (1960)
observed that the subglobose spinose morphology dominates in the oldest assem-
blages, the stout, nodose morphology predominates in the second, and the com-
pressed costate type tends to be more common in the upper levels. They tried to
categorize subordinate morphological units by assigning them letters, thereby
introducing a semi-quantitative element into their presentation. Reeside & Cob-
ban (1960) broke with traditional ammonite taxonomical dogma by recognizing
that the multitude of forms confronting them did not represent a great number of
distinct species, but were rather the outcome of a kind of variation now known to
be widespread among ammonites. Just a few examples are the variability of hopli-
tids, vascoceratids and dactylioceratids. They referred especially to the morpho-
logical variability in Triassic Trachyceras associations of the USA, monographed
by Silberling (1956), and with speci®c reference to Protrachyceras shastense Smith,
the variational pattern of which approximates that of Neogastroplites.
 Taxonomic recognition of Neogastroplites 27

Extreme variability in morphology is a variety of polymorphism, the mode of

triggering of which still remains uncertain. Continuous variability in morphology
has been best studied for Ostracoda (Ducasse, 1983; Ducasse & Rousselle, 1979;
Reyment, 1995; Reyment & Abe, 1995, for example).
Preservation. The material available to us for study consists of plaster casts of
the ®gured specimens, kept in the University Museum, Oxford University; the
originals are in the US National Museum of Natural History (USNM) in
Washington, DC. Reeside & Cobban (1960) noted that few of the shells in the
vast collections at their disposal (many thousands of specimens) preserve the body
chamber intact. The faecal accumulation hypothesis rests on the absence of sort-
ing effects in the concretions and the number of damaged shells. Against this they
forward the observation that the concretions also contain plant material, twigs
and branch fragments. Concretions of a similar nature are, however, not uncom-
mon in the geological record and can be reasonably interpreted in terms of bio-
chemical reactions. An example that springs readily to mind is that of the
orstensbollar of the Swedish Cambrian alum shale. These concretions are (not
invariably) rich in exceptionally well preserved arthropods and brachiopods,
whereas the host-sediment is nigh on barren. This topic is not germane to our
study and we leave it here.

3. Short Diagnoses
Neogastroplites cornutus (Whiteaves) (Figures 1a-l; 2a,b,e,f)
The most diagnostic features of this species were said to be its relatively coarse
ornament, weak near the umbilicus, and gently convex venter. Costate individuals
soon lose their ornament to become smooth. The stouter forms were reported to
possess lateral, ventrolateral and median ventral nodes. Subglobose forms seem
to be rare (Reeside & Cobban, 1960, p. 67).
Neogastroplites muelleri Reeside & Cobban (Figures 3a-l; 4a-l)
The most distinctive features of this species were stated as being its relatively
coarse ornament, tending to be weaker in the middle of the ¯ank, and the gently
convex venter with weak forwardly directed convex ribs. The compressed costate
variants are inclined to retain vestiges of the primary ribs and median ventral
elevations. On some stout, nodose individuals lateral and ventral ornament may
become weak. In subglobose variants, the lateral nodes displace outward to create
a ¯attening of the ventral region (Reeside & Cobban, 1960, p. 81).
Neogastroplites americanus (Reeside & Weymouth) (Figures 2c,d,g±j; 5a-p)
This species is set apart by its strong and relatively ®ne ornament, ventral rib-
bing strong in early ontogeny, and a tabulate to feebly arched venter. Ornament is
quickly lost in compressed individuals, whereas in moderately compressed var-
iants, a weak median ventral node develops. In stout variants, the lateral and
median ventral nodes become strong. Subglobose specimens are marked by an
outward migration of the lateral nodes to produce an arched ventral region, still
feebly noded (Reeside & Cobban, 1960, p. 98).
The subdivisions recognized by Reeside & Cobban are not without an element
of arbitrariness and notwithstanding the fact that it is possible to pick out speci-
mens that accord with their criteria, there are many that defy assignation, unless
one resorts to a priori knowledge of their provenances.
28 R. A. Reyment and W. J. Kennedy

Figure 1. a-l, Neogastroplites cornutus (Whiteaves). Plaster casts of: a, b, USNM 129320c; c, d,
USNM 129320e; e, f, USNM 129320b; g, h, USNM 129320f; i, j, USNM 129320 g; k, l,
USNM 129320j. Originals are from a single concretion in the Mowry Shale, hillside 15 m south
of side road along southwest ¯ank of Rattlesnake-Cedar Mountain anticline, 14.9 km (9 mi.)
southwest of Cody in SE1/4 Sec. 13, T. 52N, R. 103W, Park County, Wyoming. All ®gures are
1.5.
 Taxonomic recognition of Neogastroplites 29

Figure 2. a, b, e, f, Neogastroplites cornutus (Whiteaves). Plaster casts of: a, b, USNM 129320i; e, f,

USNM 129320m, from a single concretion in the Mowry Shale, hillside 15 m south of side road
along southwest ¯ank of Rattlesnake-Cedar Mountain anticline, 14.9 km (9 mi.) southwest of
Cody in SE1/4 Sec. 13, T. 52N, R. 103W, Park County, Wyoming. c, d, g±j, Neogastroplites
americanus (Reeside & Weymouth). Plaster casts of: c, d, USNM 129528j; g, h, USNM
129528k; i, j, USNM 129528l, all from a single concretion in the Mowry Shale on the east side
of Timber Coulee on the Winnecook Ranch, 14 km (8.5 mi.) southwest of Harlowtown on north
¯ank of middle dome of Shawmut anticline, centre of SW1/4 Sec. 14, T. 7N, R. 16E, Wheatland
County, Montana. All ®gures are 1.5.
30 R. A. Reyment and W. J. Kennedy

Figure 3. a-l, Neogastroplites muelleri Reeside & Cobban. Plaster casts of: a, b, USNM 129416a; c, d,
USNM 129416d; e, f, USNM 129416c; g, h, USNM 129416b; i, j, USNM 129416e; k, l,
USNM 129416 g. Originals are from a single concretion in the Mowry Shale, about 4.13 km
(2.5 mi.) southeast of Teigen in the centre of Sec. 4, T. 14N, R. 25E, Petroleum County,
Montana. All ®gures are 1.5.
 Taxonomic recognition of Neogastroplites 31

Figure 4. a-l, Neogastroplites muelleri Reeside & Cobban. Plaster casts of: a, b, USNM 129416h; c,
d, USNM 129416i; e, f, USNM 129416j; g, h, USNM 129416o; i, j, USNM 129416m; k, l,
USNM 129416p. Originals are from a single concretion in the Mowry Shale, about 4.13 km
(2.5 mi.) southeast of Teigen in the centre of Sec. 4, T. 14N, R. 25E, Petroleum County,
Montana. All ®gures are 1.5.
32 R. A. Reyment and W. J. Kennedy
 Taxonomic recognition of Neogastroplites 33

4. The analysis of shape in relation to ornament

The approach used here is to extract morphometric measures of shape (i.e., true
shape, free of size-effects) by the geometric method of Relative Warps (Bookstein,
1991) for each of the species, separately, and for all three groups considered as a
whole. The morphometric relationship between shape-measures and costation is
then assessed by standard methods of multiple regression and a generalization of
that technique, canonical correlation. Multiple regression is a reasonably widely
known technique, adequate descriptions of which can be found in any standard
textbook of statistical analysis. The method of relative warps, on the other hand,
is a new development, yet to ®nd its way into the generally available reference
literature. For this reason, we attempt here to give a verbal summary of the salient
features of the new geometric morphometrics in the full realization that this can
hardly do justice to the theory, but in a modest way may contribute to its
appreciation. Reyment (1996) discusses examples of size and shape analysis in
palaeontology, illustrated by geometric morphometric analyses.

5. Landmarks and coordinates

The practice of geometric morphometrics rests ®rmly on the concept of landmarks
identi®ed in a Cartesian coordinate system. In the usual course of events, measure-
ments on organisms are made on distances between chosen points. Few palaeon-
tologists lack familiarity with diameters, widths, breadths, lengths, whatever.
Without decrying the practical usefulness of traditional measures, the fact remains
that they suffer from a serious and irremediable de®ciency, namely, that once the
ruler has been removed from the specimen, the spatial relationships between traits
cannot be restored. Arguments can be (and have been) advanced, forsooth, to
attempt to gainsay this statement - for example, by recording angles between all
lines of measurement. Such artifacts are no more than just that. The data cannot
be used for any kind of useful statistical analysis and, far more importantly,
information on size and shape is just as inextricably jumbled as in the case of the
raw distances. Distance measures have their place in many analyses, particularly
those leading to ordinations, and it is not suggested that landmark methods
supplant them; however, each has its own appropriate sphere of application.
A landmark is de®ned as a speci®c point on a biological form or image. Land-
marks with the same epithet are required to correspond over the forms of a data-
set. A constellation of landmarks (the term is a borrowing from topographical
surveying via craniometrics) in two dimensions are related to a ®xed base by any
two of the landmarks. All x- and y-coordinates are rigidly translated, rotated and
rescaled to have one end of the baseline at (0,0) and the other at (1,0) by dividing
the rotated and translated coordinates by the length of the baseline. This
manipulation results in the other n-2 landmarks forming the apices of triangles,

Figure 5. a-p, Neogastroplites americanus (Reeside & Weymouth). Plaster casts of: a, b, USNM
129528b; c, d, USNM 129528f; e, f, USNM 129528g; g, h, USNM 129528a; i, j, USNM
129528e; k, l, USNM 129528d; m, n, USNM 129528m; o, p, USNM 129528i. Originals are all
from a single concretion in the Mowry Shale on the east side of Timber Coulee on the Winne-
cook Ranch, 14 km (8.5 mi.) southwest of Harlowtown on north ¯ank of middle dome of Shaw-
mut anticline, centre of SW1/4 Sec. 14, T. 7N, R. 16E, Wheatland County, Montana. All ®gures
are 1.5.
34 R. A. Reyment and W. J. Kennedy

Figure 6. The locations of the ®ve landmarks observed on the specimens.

each with a base of length 1. The outcome of this procedure is that the data now
consist of n-2 x- and y-coordinate pairs referred to as two-point shape-
coordinates, or Booksteinian coordinates. Clearly, should distance measures be
needed in any connexion, they may be easily computed from landmark data by
elementary geometry. The general idea of shape-coordinates was ®rst put forward
by Francis Galton, but it took a hundred years for a rigorous mathematical
treatment to appear in the hands of Fred L. Bookstein (see, for example,
Bookstein, 1989, 1991).
Change in shape (divorced of mere differences in size) is a compound of regular
change and irregular change. The new morphometrics makes it possible to decom-
pose overall shape-differences among shape-coordinates for a set of organisms into
af®ne (regular) and non-af®ne (irregular) components. The method of two-point
shape-coordinates is actually a kind of superimposition analysis. The fanciful
name is Procrustean analysis, derived from the antics of the bandit Procrustes (or
Damastes) of Greek mythology, who used to lay unwary travellers on a bed and if
they were too long, cut short their limbs; if, however, the bed was longer, stretched
their limbs to make a good ®t. He was slain by Theseus (Harvey, 1966). Superim-
position methods overlay a set of specimens so that corresponding landmarks
match as closely as possible. Differences in shape are recorded as residuals from a
reference shape (which could be a type specimen or, more usually, a sample aver-
age). A recent account of methods of superimposition is that of Slice (1996).

6. Selection of landmarks
The cephalopod shell poses a particular problem with respect to the choice of
meaningful landmarks. Owing to the fact that the most obvious ornamental fea-
tures, ribbing tubercles and nodes, are not ®xed in number, size or location, they
are disquali®ed - it is not possible to equate them on a 1:1 basis from specimen to
specimen. This leaves us with intersections. The points selected for the present
study are illustrated in Figure 6. Using the maximum diameter of the shell as
baseline, the coordinates at the intersection of that line across the lateral surface
at two points on the umbilical margin were recorded plus the intersection of the
penultimate whorl with the last preserved chamber. This gives a measure of lateral
 Taxonomic recognition of Neogastroplites 35

shape that involves mainly the coiling of the shell. It was not possible to locate
landmarks usefully on the cross-sections of whorls, although the maximum
breadth could be determined on most specimens.

7. Relative warp analysis

The operative mathematical tool for doing geometric morphometrics is known as
the Thin-Plate Spline, a method for visualizing a tangent space for the statistical
analysis of variation in shape (Bookstein, 1989). The mathematical concept of a
spline derives from the specialized use of an East Anglian dialect word for a ¯ex-
ible wooden or rubber strip used for drawing large, smooth curves in railway
engineering. The thin-plate spline is a smooth function that maps all points in the
physical space of the reference con®guration onto corresponding points in the
space of the i-th specimen. It provides a convenient mechanism for depicting
shape-change as a deformation and was adapted from the realm of continuum
mechanics where it is used to model the form taken by a thin metal plate that is
constrained at some combination of points and lines, but is otherwise free to
adopt the form that minimizes bending energy. The extrapolation to
morphometrics via the well known construction of D'Arcy Thompson follows
readily. Here, the same interpolation is applied once for each Cartesian
coordinate to provide a unique solution to Thompson's (1917) deformational
grids for data in the form of two landmark con®gurations. Rohlf (1996, p. 120)
summarizes the salient mathematical features of the method.
Outgoing from the thin-plate spline transformation, an analysis by relative
warps maps a reference con®guration of landmarks, usually the mean of a sample,
onto each specimen. In general terms, relative warps are the latent roots and
vectors (cf. Reyment & JoÈreskog, 1993) of the covariances of shape-coordinates
with respect to, for example, Procrustean distance. The parameters of these thin-
plate spline transformations can be used as new variables in multivariate statistical
analysis, which is what we do in this paper. In particular, the methods of principal
components and canonical variates can be conveniently invoked to analyze the
shape geometry of a sample, respectively, several samples, compared and
contrasted. We note that the new variables are linear combinations of the
residuals from the reference con®guration for the specimens.
There is a key parameter in relative warp analysis, a, the details of which cannot
be entered into here (see Bookstein, 1991; Rohlf, 1996) and we restrict ourselves
to outlining its fundamental roÃle in the study of shape. This parameter `adjusts'
the weights for the linear combinations such that for a value of a = 1, the principal
warps of the reference are scaled so as to give large-scale changes greater
in¯uence. At the other end of the scale, a = 0 causes all warps to be equally
weighted. The former choice is to be recommended if interest resides in studying
patterns of growth, whereas the latter is likely to be more appropriate in
exploratory systematic work. In the present analysis, we have used a = 0 in all
computations involving relative warps in recognition of the exploratory systematic
nature of the problem under consideration.

8. Multivariate analysis of the shape data

The method of canonical variate analysis, with af®liated procedures, can be
applied to the new data-array produced by the relative warp extraction. The main
36 R. A. Reyment and W. J. Kennedy

interest in doing such an analysis is that it permits one to display as an ordination

the group centroids (means) and scatter within groups. The group centroids can
then be compared in a one-way multivariate analysis of variance (acronym MAN-
OVA), often a useful diagnostic step in many taxonomically oriented studies. The
generalized Mahalanobis statistical distances between centroids is a byproduct of
the computations as are also a set of linear discriminant functions. For reasons of
stability (Campbell, 1980), it can be advisable to carry out the calculations in the
space of the correlations rather than in the usual space of the covariances.
It is of interest to enquire whether the shape of the shell is related to the
ornament. Eventual associations between shape parameters and ornamental
features (ribbing, tuberculation) can be examined by multiple regression and, in a
broader connexion, by canonical correlation analysis, a method which is designed
for seeking signi®cant relationships between sets of traits.

9. Findings
The relative warp ordination
The pooled material. The total sample of 28 specimens was subjected to relative
warp analysis, using the current version of the program TPSRELW, prepared by
F. J. Rohlf for the Morphometrics Work-Group (Rohlf, 1996).
The main feature of interest for present purposes concerns the result of the
ordination produced by the scores for the ®rst and second relative warps. This is
displayed in Figure 7. Inspection of this graph makes it clear that these scores
(remembering that they are, in effect, principal component scores) do not disclose
any differentiation at all in the data. Thus, if one were to terminate the analysis at
this point, the conclusion would be that on the basis of lateral shape alone, it is
not possible to hold the three forms apart. The corresponding plot of the scores
for the second and third relative warps gives virtually the same outcome. How-

Relative warp scores

cornutus

muelleri
Second relative warp

First relative warp

americanus

americanus
cornutus
muelleri

Figure 7. Ordination produced by the plot of the scores for the ®rst and second relative warps for
all species considered simultaneously. Convex hulls enclose the points for each species.
 Taxonomic recognition of Neogastroplites 37

ever, practical experience shows that a pooled principal component analysis may
not always offer the most diagnostic approach to multiple group discrimination
and there are methods available that better serve that purpose. Such a method is
canonical variate analysis.

Canonical variate analysis

1. The four shape functions. The shape functions for the three samples of Neogastro-
plites were submitted to a canonical variate analysis in the correlation space and in
the covariance space, for comparative purposes (Campbell, 1980). It is instructive
to carry out a one-way univariate analysis of variance (acronym = ANOVA) as a
preliminary step; in the present case it is highly informative in that it indicates
that the means for the ®rst and third scores for partial warps differ highly signi®-
cantly. In the case of the ®rst `new variable', P = 0.008 and, for the third `new
variable', P = 0.015.
The generalized statistical distances are also informative. For the comparison
between samples 1 and 2, D2 = 11.08, for samples 2 and 3, D2 = 12.83, both
highly signi®cant values, whereas the comparison between samples 1 and 3 yields
a value of just 0.57, which shows the two species to be similar with respect to our
shape criteria. Table 1 summarizes the main results of the canonical variate
analysis for the shape-functions.
The ordination on the ®rst two canonical variate scores (not shown) indicates
that none of the species is well separated from the other two and that there is
almost complete overlap between all of them. The Mahalanobis generalized
distances between sample means support this observation. It is important to keep
in mind that we are here dealing solely with lateral shape (free of size), without
any information from ornamental features.
2. The shape-functions combined with two extraneous traits. The next step in the anal-
ysis is to add information from the lateral ornament and the thickness of the spe-
cimens. This can be done conveniently by appending counts on rib frequencies
and observations on whorl-breadths. This introduces a special feature, the effects
of which can be undesirable, granted that we are now mixing three quite different
kinds of measures. There is also the added risk that the variability in rib-counts
could swamp the other variables. This speaks in favour of a canonical variate
solution in the space of the correlations, in accordance with the recommendations

Table 1. Canonical variate analysis of four relative warps

in the space of the covariances

Canonical Vectors
I II

RW1 ÿ3.865 ÿ1.466

RW2 ÿ2.274 ÿ0.713
RW3 6.336 ÿ2.433
RW4 ÿ1.971 8.616
Canonical roots 1.949 0.114

There are three samples which means that there can only be
two canonical vectors.
Note that almost all variability is bound to the ®rst canonical
root and, in fact, there is only one signi®cant root on the
Wilk's test criterion.
MANOVA result, P ˆ 0.001. This indicates the centroids to
differ signi®cantly. RW denotes Relative Warp.
38 R. A. Reyment and W. J. Kennedy

Table 2. Canonical variate analysis of the four shape-

criteria and whorl-breadth and rib-frequencies in the space
of the correlations.

Canonical Vectors
I II

RW1 0.05 0.65

RW2 0.16 ÿ0.29
RW3 0.08 0.50
RW4 ÿ0.15 ÿ0.35
Breadth 0.68 ÿ0.07
Ribs 0.72 ÿ0.35
Canonical roots 2.150 0.546

Both of these canonical roots are statistically signi®cant.

MANOVA result, P > 0.001. This con®rms the previous
indication (Table 1) that there is a shift in the multivariate
means over time.

Generalized distances:
Comparison D2

cornutus/muelleri 3.476
cornutus/americanus 11.338
muelleri/americanus 7.850

(RW denotes Relative Warp)

of Campbell (1980) and Seber (1984). The main details of the results for the
canonical variate analysis are summarized in Table 2.
The plot of the canonical variate scores in Figure 8 shows that none of
the species is well separated and all distributions overlap. There is, however, an
evolutionarily interesting trend in the dispersions such that there is a successive
displacement of the convex hulls, a feature that does not emerge from the plot of
the ®rst two relative warps. Moreover, the dispersions are `expanding', thus
manifesting increasing variability. The generalized statistical distances re¯ect this
relationship in that the distances from the oldest form to the youngest species
increase regularly by increments of roughly 4. The two canonical vectors listed in
Table 2 are, in effect, discriminant functions that can be called into play for
assigning a new specimen to its appropriate category. The ®rst function is
connected to 79.7% of the total variability in the material. Bearing in mind the
disparity in the variables, and their variability, it is probably advisable to apply the
second discriminant function to problems involving the assignation of specimens.
These relationships can be more closely investigated by examining the results of
the canonical correlations and multiple regressions for the three species.

10. Canonical correlation and multiple regression analysis

1. Neogastroplites, all species

The partial warp scores (i.e., the new data-matrix) obtained from the relative
warp computations (the independent variables) and rib-counts were examined by
multiple regression. The multiple correlation coef®cient of R = 0.54 yields a
variance ratio of F = 3.38 which for 4 and 33 degrees of freedom is signi®cant
(P >0.05). All correlations between the four shape indicators and rib-density are
 Taxonomic recognition of Neogastroplites 39

Canonical variate scores

cornutus

Second canoical axis

muelleri

americanus

americanus
cornutus
muelleri

First canonical axis

Figure 8. Ordination produced by the canonical variate scores in the space of the correlations,
using the four relative warps and rib-densities. Convex hulls enclose the points for each species.
There is a gradual displacement of points (centroids and clusters) from older to younger. There
is also a sequential widening of the variability. This was also apparent in Figure 7. The X-axis is
scaled at 2/3 of the Y-axis for ease of presentation.

relatively low, although still statistically signi®cant. The details of the regression
equations for this result, as well as the others, are summarized in Table 3.
The same analysis for the whorl-breadth as independent variable gives an
analogous result with R = 0.56 and F433 = 3.73; P >0.05.

2. Neogastroplites muelleri
The correlation pattern evinced by this species indicates a remarkable contrast
with the foregoing one in that RW2, RW3 and RW4 are relatively strongly
negatively correlated with rib-density, all of them being around r = ÿ 0.5 (NB,
here, and elsewhere, RW denotes Relative Warp). As regards breadth, there is
one moderately strong correlation, namely r35 = 0.59. The multiple correlation
coef®cient is R = 0.62 which as F =1.39, is not signi®cant.

3. Neogastroplites americanus
When this species is viewed on its own, we note that there are two signi®cant
correlations, namely a negative one of 0.56 between RW2 and rib-density and a
positive one of 0.67 between RW4 and rib-density. The multiple correlation
40 R. A. Reyment and W. J. Kennedy

Table 3. Summary of the details for the multiple regressions.

Species RW1 RW2 RW3 RW4 Intercept

1. Ribbing as the independent variable

All species 27.96 ÿ2.90 ÿ2.35 0.06 21.95
cornutus 0.86 5.08 ÿ9.45 ÿ0.98 19.55
muelleri 2.86 ÿ5.41 ÿ2.87 ÿ5.09 20.64
americanus 22.48 ÿ16.98 ÿ0.49 ÿ0.45 25.38
2. Maximum whorl-breadth as the independent variable
All species ÿ14.86 2.20 8.18 ÿ4.78 15.07
cornutus 0.75 ÿ8.12 8.33 2.25 13.77
muelleri ÿ3.62 2.28 7.10 1.60 14.11
americanus ÿ7.66 9.78 0.39 ÿ4.04 17.22

coef®cient of R = 0.82 is associated with a variance ratio of F = 4.17 which for 4

and 8 degrees of freedom is signi®cant on the 5% level. This points to the
likelihood of a genuine association between lateral shape and the ribbing pattern
in this species. Examination of the residuals from regression shows one very poor
®t to occur.
The result for maximum whorl-breadth as dependent variable yields two strong
correlations, to wit, a positive one of 0.71 with RW2 and a negative one of 0.79
with RW4. This almost mirrors the result obtained for the density of the ribbing
and leads one to suspect an inverse relationship to exist between ribbing and
thickness, not manifested in the two older forms. The multiple correlation coef®-
cient is R = 0.86 which as F = 5.65 on 4 and 8 degrees of freedom is signi®cant
on the 0.05% level.

11. Correlation between sets

The ®ndings yielded by the multiple regressions are interesting enough to warrant
a more detailed probing. This can be done by examining the relationships
between, on the one hand, the four relative warps, constituting one set, and, on
the other, the two extraneous variables, breadth and ribbing density, forming the
second set. The standard method of canonical correlation analysis is an appropri-
ate vehicle for realizing this aim. Canonical correlation provides a means of
seeking linear relationships between sets that are highly correlated. For an intro-
duction to palaeobiological applications, see Reyment (1991).

The analysis
The pooled set of observations (N = 38, variables = 6). There is only one signi®-
cant canonical root on Wilk's criterion, which yields a chi-square of 20.78 on 8
degrees of freedom. The associated canonical variables are listed in Table 4.
In summary, there is evidence for an association between ribbing, whorl thick-
ness the relative-warp shape parameters at the level of the species and with respect
to all three viewed as a whole. Moreover, these relationships could well be
species-speci®c, although more extensive material would be required before a
more con®dent assertion can be ventured. The pronounced differences in the
regression equations for americanus and muelleri, as well as cornutus, is interesting
supportive evidence in favour of a speci®cally relevant relationship between lateral
shape and ornament. Whether this apparent differentiation sums up to a real
speci®c difference is a moot point. The canonical correlation analysis concretizes
 Taxonomic recognition of Neogastroplites 41

Table 4. Correlation structure for ®rst canonical

correlation

Canonical correlation ˆ 0.63

Relative warp combination
ÿ0.56RW1 ‡ 0.27RW2 ‡ 0.30RW3 ÿ0.09RW4
Extraneous variables combination
0.54 Breadth -0.51 ribs

The ®rst linear combination depends largely on the

contribution from RW1. The second linear combina-
tion re¯ects an equally balanced negative relationship
between the two extraneous variables.

the results of the regressions regarding an equally balanced inverse relationship

between ribbing density and whorl-thickness.

12. Discussion
It is now appropriate to introduce the stratigraphical aspect into the analysis. The
overwhelming weight of evidence from all sources shows N. cornutus to be separ-
able from the two offshoots in a gradual relationship. The general impression con-
veyed by the statistical analysis is that the three species are located on an
evolutionary gradient with respect to shape and ornament.
The table of stratigraphical distributions of Neogastroplites (Reeside & Cobban,
1960, table 2, p. 27) seems to be reasonably ®rmly founded as regards the bio-
stratigraphical location of N. cornutus (keeping in mind that some of the crucial
support for that table is inferred). Likewise, N. americanus seems to be located in
time, at the top of the sequence, on satisfactory criteria. The evidence for
N. muelleri appears to be more circumstantial, but the evidence yielded by the pre-
sent study is in favour of the ordering advanced by Reeside & Cobban (1960).
Cobban & Kennedy (1989) suggested that the muelleri-americanus sequence
should be reversed on indirect evidence. Our analysis supports the original
sequence proposed by Reeside & Cobban (1960). Dr Cobban has, in a recent per-
sonal communication, now told us that he believes the original sequence to be the
correct one. Our conclusions, based on morphometric analysis, were reached
independently of Cobban's reappraisal of the succession. The question arises as
to whether the latter two species are suf®ciently distinct to warrant separation at
the speci®c level, at least on the basis of lateral shape and ornament as rep-
resented by ribbing. In summary, there seem to be suf®ciently many differences in
covariance structure between them to justify their recognition as distinct species.
Assuming that the material available to us gives a reasonably accurate represen-
tation of the morphometric relationships in the three species, it may be concluded
that evolution in Neogastroplites proceeded by the accumulation of gradual
changes. The detailed statistical analysis provides no evidence for abrupt
transitions.

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