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R. D. K. Thomas
Paleobiology, Vol. 4, No. 2. (Spring, 1978), pp. 181-194.
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Paleobiology, 4 ( 2) , 1978, pp. 181-194
Shell form and the ecological range
of living and extinct Arcoida
R. D. K. Thomas
Abstract.-Arcoid bivalves occupy an intermediate position, in terms both of morphology and
of adaptive range between the Pterioida and the Veneroida. The range and limits of arcoid
adaptations are related to the growth patterns of their shells. Both the arcoid hinge and ligament
grow by the serial repetition of simple structures, in contrast with the development of more
specialized, complex structures in other groups. These simple growth patterns place signifi-
cant mechanical constraints on the range of possible shell forms. Most arcoids live in moderately
unstable environments, where they are liable to be excavated or detached from their substrates.
Many employ recovery strategies, being adapted to regain their life positions. However, a
variety of specialized forms, convergent on other groups of bivalves, have become adapted to
avoid being dislodged in the first place. Thus, intrinsic growth patterns and substrate relation-
ships have been the major factors in the evolution of the Arcoida.
R. D. K. Thomas. Department of Geology, Franklin and Marshall College; Lancastcr, Pennsyl-
vania 17604
Accepted: December 29, 1977
Introduction
The patterns of organic evolution are not
fully explained as consequences of natural se-
lection acting on random variation, due to
chance genetic configurations and accidents of
geographic distribution. Selection is the effi-
cient, creative cause of evolutionary change,
but it acts within limits set both by function
and by growth processes. On the other hand,
the course of evolution has surely not been
inevitable; the diverse adaptations of living
and extinct organisms argue strongly against
even a mechanistic version of the principle of
plenitude. The particular course and patterns
of evolution on Earth are the results of chance,
historical events that have occurred in the
context of deterministic physical laws, at every
taxonomic level. The purpose of this paper is
to attempt to explain the evolution of the
arcoid bivalves in these terms.
Arcoid Characteristics and Relationships
The arcoids are, in general, distinctly un-
specialized bivalves, intermediate in form and
mode of life between typical members of the
@ 1978 The Paleontological Society. All rights reserved.
Pterioida and the Veneroida. Living and ex-
-
tinct arcoids are or were epifaunal, endobys-
sate, or shallow burrowers. Like the burrow-
ing veneroids, most arcoids are equivalve and
nearly isomyarian. The sturdy arcoid shell,
composed only of aragonite, with outer
crossed-lamellar and inner complex crossed-
lamellar layers, is similar in structure and
composition to the shells of many veneroids
and quite unlike those of the pterioids (Tay-
lor, Kennedy and Hall 1969, 1973). On the
other hand, the arcoids have filibranch gills, a
level of integration which they share with most
pterioids, whereas veneroid gills are eulamelli-
branch. The chevron or duplivincular liga-
ment, which occurs today only among the
arcoids, is also characteristic of the Paleozoic
pterioids, their later, more specialized liga-
ments having been derived from it (Newell
1937; Trueman 1969; Pojeta 1971).
I t is well established that the taxodont hinge
does not reflect a close relationship between
the arcoids and the protobranch Nuculoida
(Cox 1959; Owen 1959). The hinge teeth in
the two groups are based in different shell
layers and have quite different microstructures
(Taylor, Kennedy and Hall 1969; Flajs 1972).
182 THOMAS
FIGURE 1. Hinge plates and ligament attachnlent areas of typical arcoids. ( a ) Ct~ctcllaea uulgaris Morton, Up-
per Cretaceous, Ripley Formation, Coon Creek, Tennessee. ( b ) Anadara antiqtcata (Li nnk), Lumbo, Mozam-
hiclue. ( c ) Litnopsis multistriata ( Fbrskal), Shimoni, Kenya. ( d ) Striarca lactea ( Linnh), Bay of Naples. ( e )
Arca none Linni., Bay of Naples. ( f ) Barbntia cancellaria (Lamarck), Soldier Key, Florida. ( g ) Glycymeris
glycymeris (Linni.), Port Erin, Isle of Rlan. All except the first are speciluens fro111 living populations.
The taxodont pattern has evolved indepen-
dently in these and other bivalves (Pojeta and
Runnegar 1974), as well as the rugose coral
Calceola.
Apart from the shell, the anatomy of living
arcoids exhibits a very small range of variation
(see Heath 1941; Purchon 1957; Lim 1966),
indicating extreme evolutionary conservatism.
I have argued elsewhere (Thomas 1975) that
the anatonly of Glycymeris is unspecialized
and that its respiratory and feeding mecha-
nisms are physiologically inefficient compared
with those of many bivalves; these arguments
apply with equal force to the arcoids in gen-
eral.
It is generally accepted that the later arcoids
evolved from the early Paleozoic Cyrtodon-
tacea (Douvill6 1913; Newell 1954; Pojeta
1971) . The robust, suborbicular or posteriorly
extended shells of the cyrtodontids, together
with paleoecological data, suggest that they
were endobysate or shallow burrowers in un-
stable substrates, in common with many living
arcoids (Pojeta 1971; Stanley 1972). A good
case has been made for the derivation of the
typical taxodont dentition of later arcoids from
the cyrtodontid hinge (Newell 1954; Neves-
skaya et al. 1971). The "duplivincular" liga-
ment of the cyrtodo~ltids differs from those of
most of the later arcoids in that it lies entirely
behind the strongly prosogyrate umbones, with
the fine grooves on its attachment areas run-
ning parallel rather than oblique to the hinge
axis (Fig. 3b below). However, the hornology
of these ligaments is reaffirmed by the occur-
rence of similar and intermediate types among
the Myalinidae.
The later Paleozoic arcoids have all been as-
signed to the single genus Parallelodon, follow-
ing Arkell (1930; Newel1 1969). This group
appears to include both epibyssate and endo-
byssate forms (Stanley 1972); i t is evidently
more diverse than our current understanding
of it might suggest. Nevertheless, it is during
the Mesozoic that the major diversification of
the arcoids took place, giving rise to all the
major elements of the modern fauna (Fig. 1).
The broad adaptive trends of this radiation
ARCOID SHELL FORM AND EVOLUTION 183
are reasonably clear (Arkell 1930; Stanley
1972) but much remains to be learned about
direct phylogenetic relationships, even at the
fanlily level. The follo~ving analysis of the
functional and morphogenetic factors which
have determined shell form is intended to pro-
vide a basis for the reconstruction of arcoid
phylogeny. Shell form is then related to di-
vergent strategies employed by the arcoids in
adaptation to physically unstable environ-
ments. These different strategies are reflected
by patterns of evolution observed in the
Arcoida.
The Arcoid Shell: Functional and
Constructional Morphology
The shapes and structures of organisms are
usually interpreted and related to one another
in terms of analogy and homology, of adaptive
function and historical, phylogenetic relation-
ship. Seilacher (1970) has pointed out that a
third factor, the constructional, growth process
is itself a significant determinant of form.
Thus, the various arcoid shells are products of
particular adaptive, phylogenetic and morph-
ogenetic processes. Put another way, arcoid
diversity has evolved in consequence of oppor-
tunities presented by, and within the con-
straints of particular adaptive, phylogenetic
and morphogenetic patterns. This conceptual
framework, in which all three factors are
involved in the genesis of any form or struc-
ture, is here applied to the analysis of char-
acteristic features of arcoid shell form.
Spiral coiling and shell shape.-The pattern
of spiral, accretionary growth of the exoskele-
ton constitutes a fundamental constraint on
the range of possible arcoid shells. A continu-
ous, exoskeletal structure can increase in size
without changing in shape only by logarith-
mic spiral growth. Furthermore, Raup ( 1966)
has shown that, in a broad array of computer-
generated spiral cones, only a narrow range of
forms is geometrically suited to be articulated
in pairs about bilaterally symmetrical animals.
These morphogenetic constraints on form,
common to nearly all bivalves (Kauffman
1969), constitute part of the arcoid phy-
logenetic heritage.
On the other hand, the particular propor-
tions of arcoid shells are closely related to their
modes of life. Stanley's (1970) observation
that in free-living arcoids the shell length/
height ratio is less than 1.35, while byssally-
attached forms are more elongated, is corrobo-
rated by my data (Fig. 2) . Circular shells,
such as that of Glycymeris glycymeris are
adapted for anterior-posterior rocking locomo-
tion, by means of which the animals plough
through the superficial sediment (Thomas
1975). Posterior truncation of shells like those
of Cuculluea and many species of Anadara
minimizes excavation of shallow burrowers by
scour on unstable substrates (Stanley 1975b).
Shells which are elongated and laterally com-
pressed are suited to nestling in crevices,
among stones and in branching coral, as in
Barbatia. In contrast, Arca is elongated and
dorso-ventrally compressed, the shell being
held closely against rocky surfaces by a mas-
sive byssus (Kauffman 1969; cf. Stanley 1970
on mytilids ) . This dorso-ventral flattening is
achieved primarily by rapid growth of the
shell between the umbones.
Both Stasek (1963) and Raup (1966) have
observed that inter-umbonal growth is es-
sential in equivalve shells, since without it the
umbones of simple spiral cones would inter-
sect. This morphogenetic necessity may
largely explain interumbonal growth in some
species of Glycymeris and Anadara, where
mutual abrasion frequently produces small
facets on the umbones, but in many other
arcoids the umbones are set far enough apart
that no normal ventral gape of the shell would
bring them into opposition. Growth between
the umbones and along relatively tight spirals
at the ventral margin produces strongly in-
flated shells, such as those typical of Ana&ra,
Noetia and Cucullaea. The adaptive signifi-
cance of this inflated form is not clear, since it
makes burrowing much more difficult (Stan-
ley 1970). At least, it provides a large space
for the soft-parts and mantle cavity. In combi-
nation with a thick, heavy shell it also pro-
vides inertial stability to species such as Noetia
ponderosa and Seniliz senilis, which live in
unstable substrates.
The varied shapes of arcoid shells deveIop
by differential rates of growth around the
commissure. They are not strongly affected by
any tendency for the plane of the commissure
to twist anteriorly or posteriorly. Cuculhea
and other orthogyrate arcoids, prosogyrate
forms such as Anadara and Senilia, and the
variably opisthogyrate species of Noetia share
broadly similar shell forms. However, the de-
THOMAS
""I
10 20 30 40 50 60 70 80 90 mm
SHELL LENGTH
2.
arcoids which burrow freely or are byssally attached in unconsolidated sediment from those that nestle among
stones or in coral, are truly epifaunal, or are borers. Key to genera represented: 1, Arca. 2, Litharca. 3-6,
Baibatia. 7, Sanzacar. 8, Bentharca. 9, Trisidos. 10-11, Anadara. 12, Senilia. 13, Scapht~la.14, Poltelius. 15,
Cucullaea. 16, Noetia. 17, Sheldonella. 18-19, Glycynzeris. 20-21, Limopsis. 22, E~npleconia.
FIGURE Shapes of adult arcoid shells. Dashed line is length/height ratio of 1.35 (Stanley 1970), separating
gree and direction of twisting of the shell
away from the umbones does bring about sig-
nificant changes in the growth patterns and
functions of the ligament and hinge plate, as
will be shown in the next sections.
Growth and function of the ligament.-The
duplivincular ligament of typical arcoids con-
sists of alternating tanned lamellar and par-
tially calcified fibrous layers of proteinaceous
material (Figs. 1, 3). The lamellar ligament,
which is elastic under tensional stress (Newel1
1937), is stretched between the diverging liga-
mental areas as the shell grows. It thereby ex-
erts a continuous force which tends to open
the shell. The fibrous ligament is elastic only
under compressional stress, simply acting as
the hinge axis about which the valves rotate.
I have argued in detail elsewhere (Thomas
1976) that this ligament is inefficiently de-
signed for its purpose, compared with those of
many more advanced bivalves, in that the
layers of lamellar and fibrous ligament are not
set apart in positions where they could best
perform their respective functions.
The duplivincular ligament evidently
evolved by serial repetition of the inner and
outer layers of the archetypal bivalve ligament,
homologous (Owen, Truenian and Yonge
ARCOID SHELL FORM AND EVOLUTION
FIGURE3. Evolution of arcoid ligaments. Lamellar layers shown in black. (a) Hypothetical primary ligament.
( b ) Fine, horizontal sheets: cyrtodontids. ( c ) Typical chevrons: parallelodontids, cucullaeids, glycymerids.
( d ) Outer chevrons only: some anadarinids. ( e ) Central resilium, limopsids. ( f ) Vertical strips: noetiids. (g,
h ) Anterior sheets reduced or lost: many arcids, Anadara campechensis (Gmelin).
1953), but no longer contiguous, with the inner
and outer layers of the calcareous shell. From
this common phylogenetic heritage, a variety
of related ligament types has arisen among the
arcoids (Fig. 3) . Previous authors (Newell
1937; Trueman 1969) seem to have assumed
that the typical chevron ligament evolved
directly from a postulated primitive ligament
like that recently inferred to have been present
in the Cambrian bivalve Fordilla (Pojeta et al.
1973). However, among the cyrtodontids and
Ordovician members of the Ambonychiacea,
the grooves on the ligamental areas run paral-
lel to the hinge axis and lie entirely posterior
of the umbones (Fig. 3b; Pojeta 1971). Ap-
parently, the mantle isthmus in these bivalves
secreted lamellar and fibrous ligament alter-
nately along its length.
A true chevron ligament, with simultaneous
secretion of oblique sheets of both materials
and the addition of new sheets at the center,
is present in the questionable Devonian arcoid
Ptychodesma (Newell 1937). This basic pat-
tern is characteristic of parallelodontids, Meso-
zoic cucullaeids, glycymerids and arcids.
However, in the living Cucullaea (Figs. 3d, 6f)
and some species of Anadara, the later, central
lamellar sheets are omitted, leaving only the
anterior and posterior sheets. Alternatively,
the anterior or posterior limbs of the chevrons
have been lost, where the shell is strongly
prosogyrate, as in Barbatia, or opisthogyrate,
as in Litharca (Figs. 3g, h, 6a, c, e ) . In the
Limopsidae, the ligament is largely confined to
a resilium, consisting of a fibrous wedge
bordered anteriorly and posteriorly by lamellar
material (Fig. l c ) . As Trueman (1969) has
pointed out, this ligament is neotenous, retain-
ing the form of the juvenile arcoid ligament
elegantly described by Bernard ( 1896). Fi-
nally, in the Noetiidae, the chevron sheets have
been replaced by narrow vertical strips of
lamellar ligament, set in the fibrous material
(Fig. I d) . In this case, new strips are added
anteriorly and/or posteriorly, instead of at the
center. The origin of this growth pattern is un-
certain, but it can be generated by the re-
peated bifurcation and reorientation of the
distal lamellar sheets, as in some species of
Limopsis.
In most arcoids, the ligament and its attach-
ment areas grow proportionately more rapidly
than the rest of the shell. I have previously
shown that the need for this allometry is di-
rectly related to the growth pattern of the liga-
ment (Thomas 1976). As the umbones di-
verge during ontogeny, the ligament breaks
dorsally. Without allometric growth, the
strength of the unbroken portion of the liga-
ment would increase much less rapidly than
the weight of the animal. The allometric
growth has undesirable side effects: ventral
displacement of the hinge axis leads to inter-
ference with the function of the hinge teeth
and a reduction in the closing moment that can
be exerted by the adductor muscles. Further-
more, the rapid growth of the ligament itself
involves more divergence of the umbones, dor-
sal breakage of the ligament and a need for
yet more allometry.
In consequence of its dorsal breakage and
lack of segregation of lamellar and fibrous ma-
1.0 1 1 1 1 I I I 1 l l 1 i l 1 1 1 ' 1 1 1 ' l " l 1
0.1 1 0 10 100
SHELL "VOLUME" in cm3
FIGURE 4. Strengths of arcoid ligaments in relation
to shell "volume" (height x width x convexity). A
key to the species and parameters of the reduced
major axis regressions plotted here are given in Table
1. Slopes of regressions close to 1.0 show strength and
weight increasing in linear proportion. Both Striarca
and Noetia show substantial relative increase in liga-
ment strength with size; apparent decrease in strength
in Anadara caml?echensis must be confirmed by addi-
tional data.
terials, the arcoid ligament can never be very
strong, even where considerable allometric
growth occurs. This has been confirmed ex-
perimentally in several species of living arcoids
(Trueman 1964; Thomas 1976). Hence, the
growth pattern of the ligament has played a
major part in directing and limiting the evo-
lutionary radiation of the Arcoida. The ex-
ploitation of fully infaunal habitats has been
limited by the fact that the weak ligament is
unable to brace the shell firmly against the
substrate; arcoids are slow, inefficient burrow-
ers (Ansell and Trueman 1967). Specialization
for an epifaunal mode of life is usually accom-
panied by substantial reduction in the anterior
part of the shell (Yonge 1953) ; the arcoid liga-
ment is too weak to accommodate the tan-
gential stress developed by anterior and poste-
rior adductors exerting substantially different
closing moments about the hinge axis (Thomas
1976). Moreover, the retention of the large,
muscular foot, needed by epifaunal arcoids
such as Arca and presumably some species of
Parallelodon to aid in opening the shell, has
facilitated the repeated reversion of arcoids to
shallow burrowing modes of life, documented
by Stanley (1972).
Among the arcoids, ligament growth pat-
terns and strengths are closely related to shell
form and mode of life (Fig. 4, Table 1).
Active burrowers such as Glycymeris and
crawlers such as Barbatia have the most com-
pressed shells; their ligaments are conse-
quently less deeply split and relatively strong.
In contrast, the ligament of the dorso-ventrally
flattened Arca is functional only very close to
the hinge axis and extremely weak. In shells
with prosogyrate umbones, the ligament be-
comes largely or entirely opisthodetic, allow-
ing an adequate gape with less interumbonal
growth and less consequent dorsal splitting
of the ligament, as in Mytilus and most
heterodonts. This has been carried to an
extreme in Anadara campechensis (Figs. 3h,
6e), but here the lamellar ligament, which is
often reduced to two sheets, splits anteriorly as
TABLE1. Strengths of arcoid ligaments: data on which Fig. 4 is based. Correlation coefficients and reduced
major axis regressions of the form y =ar%alculated for logarithms of ligament strength ( y in g, mm) versus
shell "volume" ( x in mm". Strengths are compared at standardized sizes in the last two columns. For
explanation of measure of strength (g. mm) see Thomas ( 1976).
Key to
graph
Strength Strength
Sample Correlation Regression Regression g, mm g.mm
size coefficient b a at 50 c m" t 0.5 cm" Species
Glycymeris glycymeris
Glycymeris violacescens
Glycy meris bimaculata
Anadara campechensis
Noetia ponderosa
Striarca lactea
Bavbatia barbata
Arca noae
187
ARCOID SHELL FORM AND EVOLUTION
FIGURE 5. Evolution of arcoid hinge plates. ( a ) Hypothetical actinodont ancestor. ( b ) Cyrtodontid. ( c )
Parallelodontid. (d, e ) Arcids. ( f ) Cucullaeid. ( g) Glycymerid. ( h )Liinopsid.
it does in Glossus (Owen 1953), and is very
weak. The adaptive value of the noetiid liga-
ment of vertical strips is uncertain. Limited
data suggest that Striarca luctea has a strong
ligament, for its small size, but determinations
made on Noetia ponderosa show its ligament
to be extremely weak, consistent with the wide
separation of its umbones. However, the sharp
opisthogyrate curvature of the umbones in
Noetia tends to reduce the angle between the
two ligamental areas, anteriorly. This is clearly
adaptive, in that the ligament splits less rapidly
and is strongest anteriorly, where the animal is
typically partially buried in the substrate. In
conclusion, it appears that the distribution of
prosogyrate, orthogyrate and opisthogyrate
shells among the arcoids largely depends on
the growth patterns and functions of their liga-
ments, except in Anadara campechmsis, where
the prosogyrate form probably aids directly
in burrowing, as described by Stanley (1975a).
Shape and clistribution of hinge teeth.-The
posterior teeth of Cyrtodonta, of Ordovician
age, suggest that the arcoid hinge teeth origi-
nated as series of similar ridges and grooves,
oriented at a low, oblique angle to the arch of
the hinge plate and shell margin (Fig. 5;
Newell 1954; Pojeta 1971). These gave rise,
by reorientation and differentiation, to the
characteristic long, horizontal posterior and
short, oblique anterior teeth of the Parallelo-
dontidae (Newell 1954). During the Meso-
zoic, the full range of oblique, chevron-shaped
and vertical hinge teeth seen in living arcoids
evolved from these ancestors (Arkell 1930).
The evolution of the arcoid hinge consists of
innumerable variations on a single theme: the
development of interlocking hinge plates bear-
ing anterior and posterior series of simple,
relatively undifferentiated teeth.
The shapes of arcoid hinge teeth are closely
linked to the shapes of the hinge plates on
which they are borne (Fig. 5) . Chevron-
shaped teeth, like those of many cucullaeids
and glycymerids, and teeth sub-parallel to the
shell margin, such as those of Cyrtodonta and
Pseudocucullaea, can only be born on the
broad, heavy hinge plates of relatively thick-
shelled species. At the other extreme, hinge
plates as light and narrow as that of Arca will
only accommodate delicate, vertical teeth.
Shell microstructure certainly played a sig-
nificant part in making the evolution of the
arcoid hinge possible. Well developed bivalve
hinge teeth are almost invariably composed of
nacreous, crossed-lamellar or homogeneous
aragonite (data of Taylor, Kennedy and Hall
1969, 1973). The hinge teeth of extant arcoids
are all composed of crossed-lamellar aragonite
of the outer shell layer; it seems likely that the
cyrtodontid hinge, and perhaps even that of an
actinodontid ancestor common to the arcoids
and veneroids, were similarly constituted. The
shapes of the first arcoid hinge teeth may have
been related to the shape of the hinge plate;
alternatively, they may have been determined
by the distribution of lenticular crossed-lamel-
lae, which run parallel to the arch of the hinge
plate and shell margin in modern arcoids.
The function of bivalve hinge teeth has been
well explained by Dall (1895). They must
guide the valves to precise apposition during
closure and effectively resist shear in the plane
of the commissure once the shell is shut.
THOMAS
Cyrtodontid hinge teeth illustrated by Pojeta
(1971) bear distinct parallel ridges and
grooves, identical with those along which
the hinge teeth of many later arcoids slide past
one another. The widespread occurrence
among the arcoids of teeth which parallel the
hinge axis, or of chevrons with long horizontal
elements (Figs. 5c, f; 6f), is explained by the
need to align the valves correctly. In conse-
quence of the growth pattern of the ligament,
the position of the hinge axis is not precisely
defined; it is simply near the base of the liga-
ment ( Newell 1937; Thomas 1976). Further-
more, since the dorsal portion of the ligament
breaks irregularly, the position of the hinge axis
may vary slightly during ontogeny As a result,
well developed horizontal guides are needed
to ensure the apposition of the valves. This is
corroborated by the fact that in those arcoids
where the hinge teeth are reduced to vertical
elements alone, such as Arca and some anadari-
nids, the hinge axis is well defined, as only
the most ventral part of the ligament is func-
tional at all,
Chevron-shaped teeth are reduced to verti-
cal elements along the central portions of
many arcoid hinges, as they are over-iidden
during ontogeny by the ventrally expanding
ligament. This is an inevitable consequence of
the growth patterns of the arcoid hinge and
ligament (Thomas 1975). Reduction of all the
teeth to vertical elements has occurred where
adaptation dictated that the height, thickness
or curvature of the hinge plate should be re-
duced, as in Arca. The advantage of chevron-
shaped teeth over any single set of parallel
teeth in resisting variably oriented shear
stresses in the plane of the commissure is ob-
vious (Thomas 1975). It is therefore not sur-
prising to observe that strong chevron-shaped
teeth occur in active burrowers, such as gly-
cymerids, and especially in burrowers such
as cucullaeids which lack interlocking margi-
nal crenulations around the commissure.
Arcoids which have vertical tooth elements
alone are either epifaunal or they have well-
developed marginal crenulations.
Adcluctor function and attachment scars.-
Most arcoids have retained the subequal ad-
ductor muscles characteristic of the cyrtodon-
tids and the earliest bivalves (Pojeta and
Runnegar 1974). These muscles are attached
to scars composed of prismatic aragonite, con-
tinuous with the pallial myostracum; this is
consistent with the idea that the adductors
originated by cross-fusion of anterior and
posterior pallial muscle ( Yonge 1953). The
adductors close the shell by exerting a moment,
proportional to their cross-sectional areas and
distances from the hinge axis, about the hinge
axis (Thomas 1976). In isometrically growing
shells, this moment increases at the same
rate as animal weight or volume. In conse-
quence, the adductor muscle scars of most
arcoids change little in proportion or position
during post-dissoconch ontogeny.
In many arcoids, the adductor myostracum
forms a low buttress, on which the muscle is
attached. Such buttresses may serve to rein-
force the shell against stresses developed by
the adductors ( Kauffman 1969). However,
buttressed adductor scars are also more nearly
perpendicular to the lengths of the adductor
muscles than scars flush with the inner surface
of the shell. This must improve the effective
adhesion of the muscles to the shell, by reduc-
ing the shear component of stress acting on
their attachments. Among the cucullaeids,
where the posterior margin of the shell is
sharply truncated, the posterior adductor scar
is commonly held up by a flange, extending
out from the steep shell wall (Fig. 6f). These
flanges are remarkably thin, providing suit-
ably oriented but not massive support for the
attachment of the adductor.
At small body sizes, anterior reduction of
the shell is not limited by the strength of the
ligament. In endobysate species of Limopsis,
the anterior adductor is considerably smaller
and closer to the hinge axis than the posterior
adductor. The epibyssate philobryids, con-
firmed to be tiny, neotenous arcoids by Tevesz
( 1977), have gone even further, substantially
reducing the anterior adductor or losing it
altogether. They have evolved in the same di-
rection, for similar adaptive reasons, as the bet-
ter known anisomyarians described by Yonge
(1953).
External shell sczllptzrre.-Although the
cyrtodontids appear to have had smooth shells,
ornamented only by concentric growth lines,
later arcoids have acquired varied suiface
reliefs related to their modes of life (Kauff-
man 1969). Shell sculpture and phylogenetic
relationship coincide anlong the arcoids only
to the extent that taxa occupy characteristic
adaptive zones. There are no obvious mor-
phogenetic constraints on arcoid shell sculp-
ARCOID SHELL FORM AND EVOLUTION 189
ture, aside from those which govern all bivalve
shells. Simple, radial ribs form the dominant
pattern on all arcoid shells with strong surface
relief. Divaricate patterns, spines, and other
specialized structures are absent but not as a
consequence of the pattern of shell growth or
microstructure.
Shallow burrowing arcoids and those that
are endobyssate in sandy substrates either have
smooth shells or shells with strong, broad ribs.
Prominent ribs may assist or hinder in cutting
into the sediment (Stanley 1970), but they
surely aid the weak arcoid ligament in bracing
the shell against the sediment, during the prob-
ing phase of the digging cycle (Thomas 1975).
Ribs are also believed to serve shallow bur-
rowers by increasing stability and resisting
scour (Kauffman 1969; Stanley 1970). Together
with the thick shell, they inhibit predation by
carnivorous gastropods, which normally drill
bivalves with smooth shells (Carriker and
Yochelson 1968).
Nestling arcoids are characterized by finer
ribs which increase friction between the shell
and the crevice it occupies. Kauffman (1969)
has suggested that the ribs and posterior ridge
of Arca help to direct the inhalent and exhalent
currents. It is notable that Arca, which must
hold its own weight to the rock suiface, has
an extremely thin shell, for an arcoid.
Finer surface patterns are related to the at-
tachment of the periostracum in some arcoids.
Radial grooves, born on flat ribs, bear rows
of recurved periostracal barbs, which aid in
burrowing, in some species of Gl ycymr i s
(Thomas 1975). Indentations perpendicular
to the ribs, which yield cancellate ornaments,
anchor irregular flaps of periostracum which
screen the shell margin in Arca and Barbatia
( Kauffman 1969). From observations such as
these, the nature of the periostracum in fossil
arcoids can be inferred, but its structure and
functions among the living arcoids have been
as yet little studied.
The Adaptive Range of Living Arcoids
Morphologically and physiologically un-
specialized, most arcoids live in periodically
unstable environments, although the infaunal
arcoids are excluded from constantly mobile
substrates by their weak ligaments and lack of
siphons, which enable them to burrow neither
rapidly nor deeply. The diversification of the
Arcoida primarily reflects adaptation to dif-
ferent substrates, notably shifting sand and
gravel or current swept rock and coral. There
are two kinds of adaptive response to a limit-
ing stress: it can either be accomodated or its
effects can be neutralized, circumvented.
Typically, arcoids accommodate the instability
of their substrates by employing a "recovery
strategy." Glycymerids, some limopsids and
some anadarinids are frequently washed out of
their shallow burrows, but they are adapted,
by their thick, inflated shells, to withstand
rolling about and then plough through the
surface sediment in search of new burrowing
sites (Thomas 1975). Many epifaunal arcoids,
such as Barbatia and Striarca, live among
stones or in the branches of Goniolithon and
Porites, whence they are easily liable to be
detached during storms. These animals crawl
easily over hard surfaces to regain suitable at-
tachments. The large, muscular foot, adapted
by its longitudinal ventral groove to act as an
anchor in burrowing or to form a sole on a
flat surface, is critical to both of these recovery
strategies.
In contrast, Arca resists stresses that tend to
dislodge it from its exposed attachments, em-
ploying a "stronghold strategy." Hypertrophied
pedal muscles, attached to large scars beneath
the ligamental areas, hold the animal securely
down against its massive, flaring byssal attach-
ment.
While typical arcoids accommodate the in-
stability of their environments, a variety of
more specialized forms have evolved adapta-
tions, convergent on those of other bivalves,
which reduce or enable them to avoid the
risk of disturbance. However, the constraints
of arcoid shell growth and form, outlined
above, have limited the extent of these con-
vergences.
The 'mytilid' arcoid, Barbatia obliquata, has
acquired many of the characteristics associated
by Yonge (1962) and Stanley (1972) with
epibyssal attachment (Figs. 6a, b) . The an-
terior perimeter of the shell, the anterior ad-
ductor and its attachment scars are greatly
reduced. The anterior pedal retractor is also
reduced, while the posterior pedal retractor
is much enlarged, suited thereby to hold the
mytiliform shell tightly to the mid-ventral
byssus. The anterior arc of hinge teeth is re-
duced and the individual teeth are somewhat
modified. Most striking, the largely opistho-
190 THOMAS
detic ligament is very massive, exhibiting
strong positive allometry in its growth. This
strong ligament is needed to withstand the
tangential stress resulting from the different
closing moments exerted by the anterior and
posterior adductors (see p. 186). However, it
requires the secretion of a great deal of liga-
mental material and impinges more and more
on the hinge and internal volume of the shell
as it grows. Clearly the arcoid ligament,
much less well designed for this purpose than
that of the mytilids, limits the evolution of the
arcoids in this direction.
A more modest reduction of the anterior
shell and adductor is seen in two 'modiolid'
arcoids from the Cenozoic of western Europe,
Barbatia moclioliformis (Deshayes) and B.
mytiloides ( Brocchi) . The overall shapes and
smooth, thin shells of these species indicate
an endobyssate mode of life, like that of
Modiolus modiolus (LinnB) (Stanley 1970),
with which they are convergent. A much more
bizarre endobyssate adaptation is adopted by
the 'myalinid' arcoid, Trisidos, which twists
through 90" along its anterior-posterior axis.
In the ultimate compromise between infaunal
and epifaunal habits, the anterior portion of
the shell is all but that of an equivalve bur-
rower, while the postero-ventral margin lies
flat, at the sediment-water interface. In its
adoption of an inequivalve shell to increase
both stability in the sediment and the length
of its exposed postero-ventral margin, Trisidos
parallels the adaptation of intermediate mem-
bers of late Paleozoic myalinid lineages de-
scribed by Newell (1937) and Stanley (1972).
Although both Trisidos and the 'modiolid'
arcoids have evolved shell forms particularly
suited to endobyssate attachment within the
sediment, they retain the muscular foot which
enables them to reestablish themselves after
disturbance, like typical nestling species of
Barbatia.
The unique 'lithophagid' arcoid, Litharca
litlzodomus is remarkable for its convergence
with the boring mytilids (Figs. 6c, d ) . The
boring mechanism of this species has not been
described, but its straight, anteriorly tapering
shell, weak ligament and much reduced an-
terior pedal retractor muscles are clearly not
designed for either rocking or rotation in the
burrow. Rather, the hypertrophy of the poste-
rior pedal retractor scars indicates that the
animal pulls itself forward in its burrow
against an antero-ventral byssal attachment,
while the heavily corroded exterior of the shell
reflects boring by chemical solution (Thomas
1976). 111 both these respects the boring of
Litharca compares with that of Lithophaga, as
described by Yonge ( 1955). Nevertheless,
Litharca appears to be a much less efficient
borer than Lithophaga. Its acid secretion cor-
rodes its own thicker shell, whereas the thin,
more easily maneuvered, shell of Lithophaga
is protected by a continuous, resistant perio-
stracum. Its ligament is weak and associated
with interumbonal growth which requires ad-
ditional, dead space in the burrow. The weak
ligament has necessitated the retention of a
large anterior adductor, so the anterior portion
of the shell cannot be reduced, as in Litho-
phaga; Litharca has the form of an elongated
blade or wedge. Adoption of the boring habit
by Litharca was made possible by a strong
byssus; its efficiency as a borer is now most
limited by its weak ligament and widely di-
verging umbones. All these characteristics are
derived from its evident direct ancestor, Arca.
Many shallowly burrowing arcoids have
ovate or posteriorly truncated, inflated shells,
some also with coarse radial ribs. As in simi-
larly adapted veneroid bivalves, these char-
acteristics help to reduce scour around the
shell in unstable substrates (Stanley 1975b).
However, the ultimate 'cardiid' arcoid is cer-
tainly Anadara campechensis ( = A, pexata =
Lunarca ovalis auct. ) . In its oval shape, simple
ribs and marginal crenulations this species is
entirely comparable with Cardium edule. Its
shell is strongly prosogyrate, the ligament
lying entirely behind the umbones, but very
much weaker than the similarly placed
C-spring ligament of C. eclule (data above and
in Trueman 1964). In consequence of this
spiral shell growth pattern, the hinge plate
has been substantially modified, such that this
arcoid is truly heterodont (Fig. 6e). A small,
anterior group of irregular, deep teeth and
sockets, is analogous in forill and function with
the cardinal teeth of the veneroids. Posteriorly,
an arc of simple taxodont teeth function as a
posterior lateral. Remarkable as these parallels
are, in the irregular development of its hinge
teeth and especially in its weak ligament, A.
campeclzensis is much less well designed as a
burrower than the cardiids themselves. Stan-
ley (1970) confirms that this species is a slow,
sporadic burrower, whereas the cardiids tend
ARCOID SHELL FORM AND EVOLUTION
FIGURE 6. Arcoids convergent on other bivalve taxa. (a) Barbatia obliquata (Wood), Trincoinalee, Sri Lanka.
( b ) ,Mytilus edulis LinnB, St. Ives, Cornwall, England. ( c ) Litharca lithodomus (Sowerby), Manta, Ecuador;
syntype, BMNH 1969230. ( d) Lithophaga antillarum d'orbigny, Bonefish Key, Florida Bay. ( e ) Anadara
campechensis (Ginelin), Wachapieague, Virginia. ( f ) Cucullaea labiata (Solander), Awaji shinla, Japan. For
discussion see text.
to burrow rapidly. Once again, it is the liga-
ment that has primarily limited specialization,
this time with respect to a burrowing, shallow
infaunal mode of life.
Convergence is also common among the
arcoids themselves. The individual families
and subfamilies recognized by Newell (1969)
occupy characteristic adaptive zones, but with
considerable overlap at the generic and even
specific levels. For example, some limopsids
and cucullaeids converge with glycymerids,
while noetiids are convergent with anadari-
nids, with Barbatia, and with Arca. Over geo-
logical time, such parallels are seen between
groups that have replaced one another, eco-
logically. The Ordovician cyrtodontids antici-
pate several adaptive types of later arcoids,
as shown by Pojeta (1971). Parallelodontids
have been replaced by arcids and cucullaeids
occupying the same niches. Cenozoic anadari-
nids and glycymerids have largely taken over
the adaptive range of the Mesozoic cucul-
laeids. As Stanley (1972) has pointed out, this
has involved evolutionary reversals from bur-
rowing to byssate and back to free-burrowing
modes of adult life.
The occurrence of these reversals is a direct
consequence of the historical and morphoge-
netic limitations to arcoid specialization that I
have here described. The weak ligament has
prevented the arcoids froin becoming special-
ized as shallow burrowers; the lack of fused
mantle siphons further precludes deeper bur-
rowing. The weak ligament has also restrained
192 THOMAS
epibyssal specialization, both by limiting an-
terior reduction of the shell and by requiring
the retention of the foot to help in opening
the valves. From a bivalve's point of view,
coarse-grained shifting substrates are inter-
mediate betn,een rock and soft bottoms.
Among large grains, the necessarily epifaunal
mode of life of the post-larval bivalve (Stanley
1972) is prolonged. In shifting sediments, no
burrow or attachment is permanently secure,
necessitating adult as well as juvenile mobility.
It is in these unstable habitats that the gener-
alized arcoids have characteristically had, and
continue to have, their day.
Conclusions
The variety and range of arcoid morpholo-
gies is a consequence of adaptation to exploit
ecological opportunities, to the extent per-
mitted by fundamental morphogenetic pat-
terns, transmitted in particular phylogenetic
lineages (see Seilacher 1970; Vermeij 1970;
Thomas in press). Both deterministic laws
and chance configurations have played their
parts in this evolutionary radiation.
In the branching of evolving lineages, at any
given taxonomic level, some pathways are
taken and others neglected, in consequence of
chance, immediate circumstances. However,
once the "bauplan" of a group is established,
its evolution is significantly determined by its
preadaptation of form and function for life
in certain environments and not others (Stan-
ley 197513). As inolluscs, the arcoids have in-
herited a spirally coiled, calcareous exoskele-
ton, gills adaptable to respiration and feeding,
and a muscular, hydrostatic foot, suitable for
either crawling or burrowing. As bivalves,
they have inherited bilateral symmetry, an
elastic ligament linking the two valves, an-
terior and posterior adductor muscles, and a
byssus adaptable for both juvenile and adult
attachment. Finally, to their common ancestor
the arcoids owe a shell material structurally
suited to form a sturdy, interlocking hinge; a
weak, duplivincular ligament; and a notably
unspecialized soft-part anatomy.
The evolutionary consequences of this in-
heritance, discussed in detail above, suggest
three conclusions. 1) The lack of specializa-
tion of the arcoids serves them well in the
physically rigorous environments, requiring
adaptations for stabilization and mobility, that
they characteristically occupy. It has also
facilitated the evolutionary reversals noted by
Stanley (1972). 2) The origin of the elastic
ligament was at least as important as bilat-
erally symmetrical, prosogyrate shell form and
the origin of the byssus in the adaptive radia-
tion of these and other shallow burrowing and
endobyssate bivalves. 3) At their inception,
the arcoids opportunistically adopted the sim-
plest means of increasing shell size and com-
plexity. Both the taxodont hinge and the
duplivincular ligament are based on the serial
repetition of simple structures. I t seems highly
probable that such elaboration would have
required less genetic change in a generalized
ancestor than the evolution of more specialized
structures. This early opportunism made pos-
sible the Middle Ordovician radiation of the
diverse cyrtodontids (Pojeta 1971) and largely
determined the evolutionary course of the
later arcoids.
The range and limits of arcoid diversity are
direct consequences of their growth patterns,
as I have shown. The spiral coiling of the
accretionary, bivalved exoskeleton and the
serial repetition of hinge and ligament ele-
ments, selected from a small number of im-
manent, possible patterns, set constructional
limits on form. In the course of adaptation,
biologically useful structures have evolved
from among those forms that can be generated
by these patterns. As a result of the functional
correlation of anatomical parts, the constraints
of growth patterns extend beyond the struc-
tures they affect directly. So, the growth pat-
tern of the ligament and its consequent weak-
ness has required the retention of the anterior
adductor and massive foot in all larger arcoids.
Within the constraints set by growth pattern
and inheritance, diverse arcoids have evolved,
adapted to varied marine environments. Par-
ticular adaptations have arisen at random, re-
peatedly in some cases, in the larger context of
geological time and geographical distribution.
At the same time, these adaptations have been
closely determined by immediate environ-
mental circumstances. Modification of the
shell in response to the requirements of life on
different substrates is by far the predominant
feature of arcoid adaptation; remarkably little
divergence of soft-part anatomy has occurred.
Two adaptive modes are discerned in the radi-
ation of the arcoids. 1) The more typical
arcoids are adapted to survive on unstable
193 ARCOID SHELL FORM AND EVOLUTION
substrates, intermediate between truly infaunal
and epifaunal habitats, by the employment of
a recovery strategy. Frequently dislodged,
they are able to reburrow or crawl and regain
their attachments. The genus Arca is excep-
tional among the arcoid generalists in its se-
cure attachment to exposed rockv surfaces.
L
These arcoids characteristically have wide geo-
graphical distributions and long stratigraphic
ranges. 2) More specialized arcoids have
acquired a variety of adaptations, convergent
with those of other groups of bivalves, which
serve to increase their stability in or on the
substrate. Such specializations, some of which
involve substantial changes in overall shell
u
form, are largely based on simple changes in
growth pattern, which need only have involved
very minor changes in structural or regulatory
genes. These arcoids, individual species or
genera with few species, have more restricted
geographic and temporal ranges. Their local
opportunism has evidently been uninhibited
by competition with the more specialized bi-
valves on which they converge, although the
arcoids have been constrained to adopt less
efficient solutions to some of the common
adaptive problems they face.
Acknowledgments
I thank Sir Maurice Yonge for inviting me to
present some of the ideas expanded upon here
at a Royal Society discussion meeting in 1977.
Among many others, I am grateful to E. G.
Kauffman, J. Pojeta, W. E. Reif, A. Seilacher,
S. M. Stanley, J. D. Taylor, R. D. Turner, and
S. Whybrow for advice, information, and ac-
cess to collections, in the course of this study.
Special thanks are due to J. J. McDermott for
his help in collecting and maintaining live
arcoids; to L. Meszoly for Figures 1and 6; and
to M. K. Hull for her patience with the manu-
script. This paper is based on research gen-
erously supported, at different times, by SFB
53 "Palokologie," Franklin and Marshall Col-
lege, and NSF grant DEB77-01570.
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