Palaeontology - 2003 - Rouget - Embryonic Ammonoid Shell Features Intraspecific Variation Revisited

EMBRYONIC AMMONOID SHELL FEATURES:
INTRASPECIFIC VARIATION REVISITED
by ISABELLE ROUGET and P A S C A L NEIGE
ABSTRACT. Two samples of ammonoids belonging to the Oppeliidae, Sublunuloceras virguloides and Hecticoceras
(Brightii) canaliculatum, are analyzed to estimate the intraspeci®c variability of embryonic shell features. The study of
embryonic shell characters reveals two main shapes of protoconch, ¯attened and round. Prosiphons may be straight or
slightly curved. New parameters for area are added to the linear parameters commonly found in the literature.
Prosiphon length and caecum area vary greatly whereas protoconch and ammonitella diameter vary only slightly, and
the ammonitella angle is almost constant. The protoconch-to-ammonitella size ratio behaves differently in each
species, suggesting different patterns of embryonic growth. We compare our results with published data and discuss
their signi®cance for species determination and ontogenetic interpretation. The main ®nding is that intraspeci®c
embryonic variation is greater than was previously believed.
KEY WORDS: ammonoid, embryonic shell, intraspeci®c variation, Callovian, Hecticoceratinae.
M A N Y palaeontologists have investigated the ontogenetic development of ammonites through the

microstructure of the shell and the morphology of the early whorls (e.g. Druschits and Khiami 1970;
Kulicki 1974; Dauphin 1978; Tanabe et al. 1979; Landman 1987). The embryonic shell (or ammonitella)
consists of a spheroidal or ellipsoidal initial chamber (or protoconch), followed by a planispiral whorl
(Text-®g. 1). At approximately one whorl, a characteristic constriction of the tube (primary constriction)
and associated thickening of the shell outer wall (primary varix) mark the end of the ammonitella phase.
The siphuncle originates as a bulbous cñcum attached to the inner wall of the protoconch by prosiphonal
sheets (prosiphon).
Earlier investigations (Birkelund and Hansen 1968; Druschits and Khiami 1970; Kulicki 1974, 1979;
Druschits et al. 1977; Bandel and Boletzky 1979; Tanabe et al. 1980; Druschits and Doguzhaeva 1981;
Bandel et al. 1982; Landman 1982; Tanabe 1989) and analogy with recent cephalopods (Boletzky 1974)
support the hypothesis that ammonoids developed directly without a larval stage. Most workers divide
ammonite development into embryonic (secretion of the ammonitella) and post-embryonic (secretion of
the remaining shell) stages. The transition point between the two stages is marked by sudden changes in
surface ornament, the appearance of a nacreous layer, and acceleration of growth (e.g. Erben et al. 1969;
Birkelund and Hansen 1974; Kulicki 1974, 1979; Dauphin 1978; Landman 1987; Tanabe et al. 1993;
Neige 1997). This transition point corresponds to the ammonite hatching event.
All these ®ndings are summarized in the review paper (Landman et al. 1996a) of the latest ammonoid
palaeobiology `red' book (Landman et al. 1996b). It mainly re¯ects published data and concentrates on
(1) variations in the shape and size of the protoconch and ammonitella (mostly data from before 1985)
and (2) the effect of such variation on early life history (mainly Tanabe and Ohtsuka 1985, but see also
Kulicki 1974; Druschits et al. 1977). This gives the impression that the diversity of ammonoid
embryonic features has been well described from whole ammonoids (bauplan disparity) to the operational
evolutionary unit (population or species). In fact most studies of the early ontogeny of ammonoids have
concentrated on interspeci®c variation in the size of the protoconch and ammonitella (Druschits et al.
1977; Tanabe et al. 1979; Tanabe and Ohtsuka 1985; Ohtsuka 1986; Tanabe et al. 1994), rather than on
intraspeci®c variability: intraspeci®c data relate to small and sometimes unde®ned numbers of speci-
mens (Text-®g. 2). This may be because of the dif®culties of analysis and of obtaining well-preserved
material.
[Palaeontology, Vol. 44, Part 1, 2001, pp. 53±64, 2 pls] q The Palaeontological Association
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54 PALAEONTOLOGY, VOLUME 44
TEXT-FIG. 1. Basic morphology and terminology of the early internal shell structure of an ammonoid (in median section).
The shortage of intraspeci®c data raises two related questions: (1) can embryonic features be used for
species identi®cation, and (2) can the diversity of embryonic features at species level tell us more about
ammonoid development?
In this paper we attempt to characterize intraspeci®c variability of the embryonic shell (ammonitella and
protoconch size and shape), as well as of the cñcum and prosiphon. Two species in the Oppeliidae are
studied: Sublunuloceras virguloides (Spath, 1928) and Hecticoceras (Brightia) canaliculatum (Quenstedt,
1846). Although these two sympatric species belong to two different genera, they are closely related and
both have a hecticoceratoid shape (Elmi 1967). The results for these two species are compared ®rst and
then further compared with published data for other Ammonitina. We conclude by discussing the use of
embryonic data as clues to ammonoid development and as an aid to species identi®cation.
MATERIAL AND METHODS

Sample collection
The observation of ammonoid embryonic structures and the study of their intraspeci®c variability require a
large, well-preserved sample. The two species under review are from the Callovian Stage [Middle Jurassic of
TEXT-FIG.2. Synthesis of the number of specimens measured for each species studied in previous embryonic ammonoid
studies (for example embryonic measurements for 136 species are known from a single specimen). It is based on all
Ammonoidea, and covers 350 species and more than 1800 specimens. The total number of specimens could not be
calculated for 129 of the 350 species cited in the literature because it was not possible to ®nd the relevant information
in the publications.
ROUGET AND NEIGE: EMBRYONIC AMMONITE SHELLS 55
Burgundy (Athleta Zone, Collotiformis Subzone; see Bonnot et al. 1999)]. The specimens were all collected
from a 10-cm-thick limestone bed. The specimens are well preserved (intact lappets, absence of intra-thalamic
epizoans) and many of the associated fauna (gastropods, bivalves, scaphopods, brachiopods) exhibit
fragile structures such as spines or intricate ornamentation. The good state of preservation is indicative of
rapid burial, suggesting that the assemblage is an accurate census of the former living community.
In the sample examined, specimens of S. virguloides are rather small. The shell is discoidal, oxycone,
more or less involute, and ogival in section. The ¯anks are slightly convex. The venter is narrow and
unicarinate. The inner whorls are smooth at ®rst and then proverse primary ribs appear. These ribs arise at
the umbilical edge and bifurcate near the venter. Secondary and intercalated ribs arise near the middle of
the ¯ank. The occurrence of two different adult morphologies (presence of lappet on small specimens),
exhibiting similar juvenile stages suggests sexual dimorphism (Bonnot et al. 1999).
H. (B.) canaliculatum is thinner than S. virguloides. It also has a marked lateral sulcus, and ®ner
ornamentation. Its ¯anks are almost parallel and the ventral area is thin with a sharp keel. The inner whorls
are smooth and unicarinate. After several whorls, tubercles and ®ne ribbing appear below and above the
sulcus respectively. The ribs are initially retroverse and then clearly proverse towards the venter. No
dimorphic pattern is assumed, although one was suggested by Bonnot et al. (1999).
The material used in this study is housed at the Centre des Sciences de la Terre, UniversiteÂ de
Bourgogne (number pre®x GD).
Sample preparation
The material was prepared in median polished section to expose the ®rst whorls. A number of thin sections
were prepared. Embryonic shells could be studied in 52 specimens (34 S. virguloides, 17 macroconchs and
17 microconchs, and 18 H. (B.) canaliculatum) of the 104 polished sections.
Method of measuring
The following linear (mm) and surface area (mm2) measurements were made on the median sections
(Text-®g. 3): maximum and minimum protoconch diameter (PD1 and PD2 respectively); protoconch area
(PS); caecum area (CS); prosiphon length (PL); ammonitella diameter (AD); ammonitella angle in degrees
(AA). Measuring the diameters of the protoconch and ammonitella involved locating the centre of the
protoconch. This was taken here as the centre of the circle inscribed in the basal part of the protoconch
(Grandjean 1910; see Landman 1987 and Neige 1997 for a discussion of this choice).
TEXT-FIG. 3. Measurement of the early internal shell structure of ammonoids in median cross-section through the
ammonitella.
TABLE 1. Measurement data of selected internal shell characters in the two species studied. N, number of specimens;
SD, standard deviation; CV, coef®cient of variation.
N Mean SD Range CV
PD1 S. virguloides 34 363 25´4 303±413 0´07

PD1 H. (B.) canaliculatum 18 390 28´0 355±452 0´07
PD2 S. virguloides 34 284 22´4 237±333 0´08
PD2 H. (B.) canaliculatum 18 302 21´5 266±348 0´07
PS S. virguloides 33 0´082 0´012 0´054±0´102 0´14
PS H. (B.) canaliculatum 18 0´093 0´011 0´078±0´123 0´12
CS S. virguloides 33 0´0034 0´001 0´001±0´008 0´37
CS H. (B.) canaliculatum 18 0´0043 0´001 0´002±0´006 0´29
PL S. virguloides 24 166 0´046 88±269 0´28
PL H. (B.) canaliculatum 12 188 0´059 81±302 0´31
AD S. virguloides 34 600 43´1 491±706 0´07

AD H. (B.) canaliculatum 18 664 54´3 589±738 0´08
AA S. virguloides 34 259 14´1 225±298 0´05
AA H. (B.) canaliculatum 18 257 14´5 234±288 0´05
RESULTS
Intraspeci®c comparisons
Within the range of variation for S. virguloides (Table 1), there are no signi®cant differences in the size of
the protoconch (PD1) between the dimorphs (303±400 mm for the microconch and 303±413 mm for the
macroconch). For H. (B.) canaliculatum, PD1 ranges from 355 to 452 (Table 1). Protoconch maximum and
minimum diameters are signi®cantly different at the 1 per cent level for both species (Table 2). This
re¯ects the elliptical shape of the protoconch as is consistently described for all Ammonitina (Landman
et al. 1996a). Two main types of protoconch shape can be distinguished: (1) an irregular protoconch, with
a rounded basal part and ¯attened upper part (Pl. 1, ®gs 1±2); and (2) a subspherical protoconch, with
regularly curved basal and upper parts (Pl. 1, ®gs 3±4). These two shapes occur both in the microconchs
and in the macroconchs, and for large or small protoconchs. The mean area of the protoconch (Table 1) is
0´082 mm2 for S. virguloides and 0´093 mm2 for H. (B.) canaliculatum. The larger protoconchs may be
twice the size of the smaller ones, implying very variable volume of the initial chamber within the same
species.
All the observed cñcum look elliptical. The mean area (CS) is 0´0034 mm2 for S. virguloides and
0´0043 mm2 for H. (B.) canaliculatum and the coef®cient of variation indicates wide variability in both
species (Table 1). Cñcum area accounts for around 5 per cent of the protoconch surface, but there is no link
between the two (r close to 0´5 for both species, Tables 3±4).
TABLE 2. Student's t test of the mean of maximum and minimum protoconch diameters within species. ****, signi®cant
at 1 per cent level.
DDL t p
S. virguloides (PD1, PD2) 63 ± 13´008 ****

H. (B.) canaliculatum (PD1, PD2) 34 ± 10´547 ****
TABLE 3. Matrix of correlation of the selected internal shell characters in S. virguloides. Signi®cant values are
underlined.
PD1 PD2 AD AA PL PS CS
PD1 1´00 0´87 0´87 0´10 0´39 0´95 0´49

PD2 1´00 0´80 0´12 0´38 0´93 0´46
AD 1´00 0´26 0´23 0´87 0´28
AA 1´00 0´05 0´12 0´33
PL 1´00 0´30 0´04
PS 1´00 0´51
CS 1´00
Most prosiphons observed in S. virguloides and H. (B.) canaliculatum are straight (Pl. 2, ®gs 1±2).
Several specimens, however, have slightly curved prosiphons (Pl. 2, ®gs 3±4). The coef®cient of variation
(Table 1) indicates wide variation in prosiphon length within species (Pl. 2, ®gs 5±6). Such intraspeci®c
variation in prosiphon size and shape is not reported in previous studies.
A Student's t test indicates no signi®cant difference between the two dimorphs for ammonitella
diameters of S. virguloides (491±693 mm in the microconch and 524±706 mm in the macroconch).
Ammonitella angle ranges from 225 to 280 degrees for the microconch and 240 to 298 degrees in the
macroconch with a mean value of 259 degrees (Table 1). No correlation was found between ammonitella
diameter and ammonitella angle for S. virguloides and H. (B.) canaliculatum (Tables 3±4). However,
ammonitella diameter and protoconch diameter are closely correlated for the two species (Tables 3±4).
This result is consistent with other published ®ndings at different taxonomic levels (Tanabe et al. 1979;
Tanabe and Ohtsuka 1985; Landman 1987; Tanabe et al. 1994).
Interspeci®c comparisons
A Student's t test of (1) protoconch diameter, (2) protoconch area, (3) caecum area, and (4) ammonitella
diameter indicates that H. (B.) canaliculatum is signi®cantly larger than S. virguloides (Table 5). The same
test indicates the length of the prosiphon and the angle of the ammonitella are not signi®cantly different
between the two species (Table 5).
The linear relationship between protoconch and ammonitella diameter seems to be different between
both species (Text-®g. 4). A Z test to de®ne isometry and allometry statistically (Hayami and Matsukuma
1970) reveals that for S. virguloides the AD:PD1 ratio remains constant as PD1 increases, whereas for
H. (B.) canaliculatum the AD:PD1 ratio decreases with increasing protoconch diameter. This also means
that for H. (B.) canaliculatum embryonic size variability was proportionally lower at hatching than at the
start of embryonic growth. These results suggest (1) that differences in embryonic size between species
TABLE 4. Matrix of correlation of the selected internal shell characters in H. (B.) canaliculatum. Signi®cant values are
underlined.
PD1 PD2 AD AA PL PS CS
PD1 1´00 0´87 0´60 0´08 0´68 0´92 0´45

PD2 1´00 0´47 0´07 0´66 0´93 0´56
AD 1´00 0´36 0´54 0´59 0´27
AA 1´00 0´19 0´09 0´14
PL 1´00 0´70 0´46
PS 1´00 0´50
CS 1´00
TABLE 5. Student's t test of means of embryonic shell measurement (PD, AD, AA, PS, CS, PL) in S. virguloides and
H. (B.) canaliculatum. ****, signi®cant at 1 per cent level; ***, signi®cant at 5 per cent level; NS, not signi®cant.
DDL t p
PD1 50 3?578 ****

AD 50 4?890 ****
AA 50 ± 0?086 NS
PS 49 3?139 ***
CS 49 2?834 ***
PL 34 1?230 NS
TEXT-FIG. 4. Plot of the diameter of the ammonitella versus the diameter of the initial chamber in S. virguloides and
H. (B.) canaliculatum. Regression lines show intraspeci®c linear relationships.
were greater in the early growth stages than at hatching and (2) that the embryonic pattern of growth was
probably not the same for each species.
DISCUSSION
Reconstructing the early life history of ammonoids remains dif®cult. Previous authors have used evidence
from specimens preserved at early ontogenetic stages or from the early whorls of larger specimens. As
EXPLANATION OF PLATE 1
Figs 1±4. Sublunuloceras virguloides (Spath 1928). 1±2, GD1730-H18 (macroconch), ´ 120 and GD1729-H24
(microconch), ´ 110, embryonic shell in median section with a basal part of the protoconch round and the upper part
¯attened. 3±4, GD1727-H17 (microconch) and GD1728-H9 (macroconch), subspherical protoconch; ´ 120. All
specimens from the Athleta Zone (Callovian, Jurassic) Burgundy.
PLATE 1
ROUGET and NEIGE, Sublunuloceras

claimed by Landman et al. (1996a), data from comparative anatomy, microstructural study and
morphometry are complementary in determining the sequence of embryonic development.
Broad taxonomic surveys suggest differences among higher taxa (e.g. ammonitella angle, ammonitella
diameter, prosiphon length and caecum shape) and probably assume only slight intraspeci®c variation
(Tanabe and Ohtsuka 1985; Ohtsuka 1986; Tanabe et al. 1994). However, our ®ndings show high
intraspeci®c variation for some features.
Among Ammonoidea as a whole, protoconch diameter varies from 250 mm to 1´6 mm (Landman et al.
1996a). Except for the Agoniatitidae, the mean protoconch diameter does not vary between suborders
(Landman et al. 1996a). Ammonitina protoconch size ranges from 250 to 700 mm. Protoconch size as
reported for eight species of Oppeliidae (Palframan 1966, 1967, 1969; Druschits and Khiami 1970;
Druschits and Doguzhaeva 1981; Neige 1997) is small (250±450 mm) compared with other Ammonitina
(Text-®g. 5). However, intraspeci®c variability of the two species studied here is almost as great as that
found for the Oppeliidae as a whole (Text-®g. 5). This suggests that protoconch size is not as stable a
character at species level as was previously thought.
Earlier authors have assumed that the protoconch provided buoyancy. It was believed to be ®lled
originally with cameral liquid which was discharged and replaced by gas just before or after hatching.
Thereafter the newly hatched, neutrally buoyant, ammonoid supposedly led a planktic mode of life
(Kulicki 1974, 1979; Druschits et al. 1977; Tanabe et al. 1980; Bandel 1982; Landman 1985; Tanabe and
Ohtsuka 1985). The volumetric relationship between protoconch and ammonitella, indicating an almost
constant volume ratio in many ammonoids, supports the neutral buoyancy hypothesis (Shigeta 1993). The
large variation in protoconch area reported in this paper could mean that the initial chamber volume may
have varied widely between and within species. Assuming the initial chamber was completely full of
cameral liquid at the beginning of development, our ®ndings suggest that the volume of cameral liquid
varied greatly within species. This could have been directly related to the regulating factors of a planktic
mode of life such as the rate of cameral liquid discharge (Shigeta 1993). Consequently, it would be
worthwhile comparing protoconch and siphuncular areas in further studies.
Cñcum and prosiphon. Published data about the cñcum and prosiphon are rare. However, in the summary
paper by Landman et al. (1996a), the cñcum of several Ammonitina are described as elliptical. Available
bibliographic data distinguish two main types of prosiphon (Landman et al. 1996a): (1) a long, nearly
straight prosiphon in most Ammonitina except for the Amaltheidae, Collignoceratidae and Placenti-
ceratidae; and (2) a short, curved prosiphon in Ancyloceratina, Bacritina, Goniatitina, Lytoceratina,
Phylloceratina and Prolecanitida. For S. virguloides and H. (B.) canaliculatum the cñcum observed are all
elliptical and shape seems to be constant within and between species. In contrast, the prosiphon may be
straight or slightly curved and length varies greatly: prosiphon length and cñcum size are the most variable
quantitative characters of all embryonic features.
Ammonitella. Previous authors have suggested that most ammonoids hatched with a relatively small
embryonic shell. Variation in ammonitella diameter was reported among (1) the different suborders of
Ammonoidea and (2) species of the same suborder (e.g. Goniatitina). It has also been shown that the size of
the ammonitella varies little within species (Hirano 1975; Tanabe and Ohtsuka 1985). In the suborder
Ammonitina, the diameter of the ammonitella ranges from 520 mm to 1´3 mm (Text-®g. 5). In the Oppeliidae,
EXPLANATION OF PLATE 2
Figs 1±6. Sublunuloceras virguloides (Spath 1928). 1±2, GD1733-H30 (microconch) and GD1730-H18 (macro-
conch), prosiphon long and straight; ´ 120. 3, GD1731-H1 (macroconch), prosiphon long and slightly curved; ´ 200.
4, GD1732-H2 (macroconch), prosiphon slightly curved; ´ 120. 5, GD1729-H24 (microconch), small protoconch;
the prosiphon is long and straight, but curves near the point of attachment to the wall; ´ 120. 6, GD1734-H8
(microconch), prosiphon short and straight; ´ 120. All specimens from the Athleta Zone (Callovian, Jurassic)
Burgundy.
PLATE 2
ROUGET and NEIGE, Sublunuloceras

TEXT-FIG. 5. Comparisons of the variation in the diameter of the protoconch and ammonitella at different taxonomic levels:
Ammonitina, Oppeliidae (bibliographical data excluding the present study), and S. virguloides and H. (B.) canaliculatum.
shells are small at hatching (520±900 mm) compared with the Ammonitina. Our results show that intra-
speci®c variation of S. virguloides and H. (B.) canaliculatum partly overlaps with that of the Oppeliidae
(Text-®g. 5). We also report that (1) intraspeci®c variability was not as stable as assumed by earlier workers
(as for protoconchs), and (2) size at hatching may have differed between phyletically close species.
The angle of the ammonitella is the most variable embryonic shell character at the scale of the suborder
(Druschits and Khiami 1970; Tanabe and Ohtsuka 1985; Tanabe et al. 1994; Landman et al. 1996a).
However, at lower taxonomic levels, this variation is largely reduced (Tanabe et al. 1979; Landman 1987).
Results obtained for S. virguloides and H. (B.) canaliculatum are consistent with this view. Coef®cients of
variation are smaller for ammonitella angle than for protoconch and ammonitella diameters and the mean
ammonitella angle is similar in both species.
Therefore, as demonstrated here, size and even shape cannot be used for identifying characters at
species level. For example, individuals of the same species may have either long, straight or short, curved
prosiphons. Moreover, size ranges (e.g. diameter of protoconch and ammonitella) within the families as
generally reported in the literature are similar to size ranges at the species level. This suggests that data
about ammonoid families re¯ect the state-of-the-art rather than reality. In contrast to the large variation
in size and shape at species level, anatomical characters are extremely stable in terms of position and
occurrence. We consider that within-species size and shape data are not a sound basis for tracing
embryonic development. As summarized by Landman et al. (1996a), shell-wall microstructure, orna-
mentation and general anatomy are the three main strands of evidence used by authors to reconstruct
embryonic development sequences. Quantitative measurements have yielded information solely about a
sudden change in mode of life at the embryonic/post-embryonic event (change in rate of shell growth: see
Landman 1987; Neige 1997).
This analysis shows the need to extend our knowledge of intraspeci®c embryonic variation. Studies on
the relationship between embryonic and post-embryonic development (e.g. the effect of size and coiling
variation of the former stage on the rest of development) are essential for a complete understanding of
ammonoid growth.
Acknowledgements. We thank D. Marchand for providing some of the material studied, N. H. Landman and N. Monks
for their reviews of the manuscript, and C. Sutcliffe for assistance with the English. This paper is a contribution to the
theme `Signal Morphologique de l'Evolution' of the UMR±CNRS 5561 `BiogeÂosciences'.
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ISABELLE ROUGET
PASCAL NEIGE
Centre des Sciences de la Terre

UMR CNRS 5561
UniversiteÂ de Bourgogne
6, boulevard Gabriel
F-21000 Dijon, France
Typescript received 31 January 2000 e-mail Isabelle.Rouget@u-bourgogne.fr
Revised typescript received 28 June 2000 Pascal.Neige@u-bourgogne.fr

Palaeontology - 2003 - Rouget - Embryonic Ammonoid Shell Features Intraspecific Variation Revisited

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EMBRYONIC AMMONOID SHELL FEATURES:

INTRASPECIFIC VARIATION REVISITED

by ISABELLE ROUGET and P A S C A L NEIGE

KEY WORDS: ammonoid, embryonic shell, intraspeci®c variation, Callovian, Hecticoceratinae.

M A N Y palaeontologists have investigated the ontogenetic development of ammonites through the

MATERIAL AND METHODS

PD1 S. virguloides 34 363 25´4 303±413 0´07

AD S. virguloides 34 600 43´1 491±706 0´07

S. virguloides (PD1, PD2) 63 ± 13´008 ****

PD1 1´00 0´87 0´87 0´10 0´39 0´95 0´49

PD1 1´00 0´87 0´60 0´08 0´68 0´92 0´45

PD1 50 3?578 ****

ROUGET and NEIGE, Sublunuloceras

ROUGET and NEIGE, Sublunuloceras

Centre des Sciences de la Terre

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