Feedinghabitsintrilobites: by and

FEEDING HABITS IN TRILOBITES
by R. A. FORTEY and R . M. OWENS
ABSTRACT. We briefly review the various types of feeding habits in marine arthropods, and suggest that the trilobites
adopted a range of different feeding strategies. We show that much of the variety of trilobite exoskeletal development,
particularly in the cephalon, can be explained as a response to the adoption of specific feeding modes. We regard the
primitive mode as having been predatory/scavenging, both from morphological grounds and by out-group comparison,
but this habit had a long subsequent history in the group. Predators/scavengers included those trilobites with rigidly
braced and attached conterminant or impendent hypostomes, which often developed posterior forks or rasps used by
the animals for manipulating prey after it had been grasped by the ‘gnathobases’. Advanced predators often acquired
expanded anterior glabellar lobes which are associated with the ingestion of bulky food; concomitantly, the largest
trilobites of all had predatory morphology. Associated trace fossils are of the Rusophycus type in which impressions of
limb bases and rarely the hypostome can be seen. Detritivors were derived from predators by detachment of the
hypostome from the doublure in natant mode; it is significant that the hypostome of such feeders exhibited little change
thereafter. The typical detritivor morphology is of the ‘generalized’ ptychopariid type, common in outer shelf habitats,
with rectangular or tapering glabellas and small to moderate overall size. It is suggested that in some species the
hypostome may have functioned as a ‘scoop’ directly to aid ingestion of sediment. Trace fossils of Cruziana
semiplicata type have been associated with sediment ploughing in this feeding mode. Filter feeders evolved a vaulted
cephalic chamber of trinucleimorph type, and elevated thoraces, often flanked by extended genal spines. Where it is
known, the hypostome is curved up inside the cephalic chamber, within which sediment stirred into suspension by the
limbs was sorted for edible particles. Filter feeding trilobites are typically small, and are uncommon outside muddy
habitats. Bean-like Rusophycus are the associated trace fossils. In trinucleids ingress of the feeding current was
alongside the thorax and out through the fringe pits. The combination of different feeding modes with adaptation for
different prey and/or particle sizes goes some way to account for the variety of trilobites cohabiting in a single site
(alpha diversity). We do not claim that the model accounts for all morphological variation displayed by the group.
TR I L O B I T E S are remarkable for their great variety of exoskeletal shapes. Their morphological complexity
increased from the Cambrian into the Ordovician (Fortey and Owens 1990; Foote 1991), and a
considerable array of forms persisted even into the Carboniferous. Yet comparatively little is known
about how this morphological variety related to life habits. It is common to find reference to certain phases
of trilobite history as ‘radiations’, for example, at the beginning of the Cambrian (Briggs and Fortey 1992),
through much of the Ordovician (Droser et al. 1996) and in the Malvinokaffric Province of the Devonian
(Eldredge 1980), yet although the implication clearly is that radiations were into a variety of niches, it is
unclear what these niches were, nor whether the morphological compass displayed can be related to their
life habits. Broad habitat differentiation is better known. It is clear that the trilobites were able to occupy
normal marine habitats (biofacies) ranging from near inshore, to deep water, even ‘slope’ environments, at
several hundred metres of water depth, from early in the Cambrian until the later Carboniferous (review in
Fortey and Owens 1997). This differentiation accounts for some of the richness in species of trilobites of
any given geological age, but it does not account for the subdivision of each habitat by its constituent
species (alpha diversity). Biofacies can vary in the number of species and genera they include, but it is not
uncommon for a single biofacies to include 20 or more benthic taxa (e.g. Ludvigsen et al. 1989).
Presumably, such co-occurring species found ways of exploiting their total environment which allowed
them to compete adequately with their neighbours.
The kinds of niche subdivisions of habitat which were possible for trilobites have been little investigated
(Bergström 1973). For such an extinct arthropod group, particularly one of which the appendages of so few
species are known, the interpretation of life habits is not an easy matter. However, this should not prevent
attempts to interpret reasonably function from form. This paper presents a model for trilobite feeding
[Palaeontology, Vol. 42, Part 3, 1999, pp. 429–465] q The Palaeontological Association
430 PALAEONTOLOGY, VOLUME 42
habits, with an aim of answering one of the commonest questions advanced about the group: why many
different kinds of trilobites co-exist in the same rock bed. Not all trilobites were benthic; a free swimming
fauna also lived within the water column, at least during the Cambrian and Ordovician (Robison 1972;
Fortey 1985; Fortey and Owens 1987). Robison (1975) explained the co-occurrence of differently sized
planktic agnostids as resource partitioning, whereby adult trilobite size reflected optimal utilization of
different food particle sizes, thus relating taxonomic variety and aspects of morphology to feeding
specialization. However, there are often references to feeding mode in otherwise morphological and
taxonomic accounts, and we have attempted to review the scattered literature of the past 50 years in this
paper. Such a review should be of use to the non-specialist palaeontologist for teaching purposes. Our
prime purpose is to identify a number of morphological designs connected with different feeding habits,
discuss their geological history, and associate them broadly with their appropriate trace fossils.
THE RANGE OF POSSIBILITIES FOR TRILOBITE FEEDING HABITS

Marine arthropods occupy many different niches, and not all of these are capable of being deduced from
fossil evidence. There is no necessary relationship between belonging to a particular high level taxon and
having a specific life habit. Virtually all larger Arachnida, for example, are predatory in habit, while
crustaceans may be filter feeders, sediment feeders, parasites, or predators (Schram 1986). Even within a
single crustacean group (e.g. Cladocera, see Fryer 1968) there may be a variety of feeding habits. Hence
the recognition of the feeding habits of one species is no guarantee that the rest of the group to which it
belongs will have similar habits. For trilobites, we are perforce concerned only with the broadest
categories of feeding strategies, and we cannot approach the refinements known for the living fauna.
To give one example, Laval (1980) describes how hyperiid amphipods have evolved a close relationship
with medusoids and ctenophores, using them as a nursery for young, or in some cases killing them to
construct their own floating accommodation. Such habits will forever elude palaeontological evidence. In
the sections that follow, the attitude and position of the trilobite hypostome will be of importance.
Throughout the rest of this paper we accept the commonly held view that the trilobite’s mouth lay at the
back of the hypostome, and that the oesophagus lay above it. The oesophagus opened into the stomach
situated beneath the glabella, especially its anterior part. The oesophagus was elevated or contracted by
muscles which attached between the hypostome and the oesophagal wall. This is an arrangement similar to
that described for the living notostracan Lepidurus arcticus by Fryer (1988, fig. 63).
The life habits available to arthropods (and conceivably to trilobites) can be summarized briefly as
follows:
1. Predatory. Hunting habits are common among crustaceans (e.g. Stomatopoda) and dominant in
Arachnida. Limulus is capable of breaking clams with its gnathobasic jaws. There is a gradation between
hunting activity and scavenging, and many crustaceans which are capable of hunting, e.g. among the
decapods, are also opportunistic scavengers if the possibility arises. The manipulation of comparatively
bulky food unites these two activities, and, because of the relatively high nutritional value of such food,
larger arthropods tend to have such habits. We cannnot distinguish scavenging from predatory habits here.
2. Particle feeding. Feeding on edible detritus is a common mode of life among crustaceans, including
branchiopods, ostracodes and isopods, among animals ranging from a few millimetres to a few tens of
millimetres long (Brusca and Brusca 1990). Included in this category is feeding using the ventral median
line as a food groove, such as has been described in the diminutive and primitive cephalocarid
Hutchinsoniella by Sanders (1963). Sorting of edible from inedible particles is achieved by the
appendages. A variant on this life mode is scraping or collecting edible material, usually algal, directly
from rocks or sedimentary surfaces.
3. Suspension feeding. Many arthropods, notostracans, conchostracans and phyllocarids among them, stir
up sediment in order to extract edible organic particles, or they exploit algal ‘soup’ in similar fashion. The
notostracan Triops, for example, stirs up clouds of sediment beneath its dorsal carapace, and passes the
edible particles sorted from suspension towards the mouth along the ventral food groove (Fryer 1988). One
might suspect that this mode of life ought to be present among trilobites, especially in view of the
similarity of the vaulted headshield of many species to that of Triops.
FORTEY AND OWENS: TRILOBITE FEEDING HABITS 431
TEXT-FIG. 1. Hypothesis
of relationships of trilobites and their close relatives indicating why predatory habits are likely
to have been primitive for the group as a whole, and specifically for Olenellus. See text for discussion.
4. Filter feeding. In this case the appendages are used to entrap and select out food particles, often
plankton, by forming a net or basket. The appendages are usually modified to perform this function
(Stubbings 1975), as in the cirri of the barnacles (Cirripedia), or the anntenna and antennule of the planktic
copepods. There is no example of a sessile trilobite, all of which had vagrant benthic or nektic habits, and
thus any anology with cirripedes is implausible. However, some trilobites lived as part of the pelagic
community in adult life, and many, perhaps even the majority of trilobites had planktic larval stages, and it
is likely that filter feeding was important at this stage of growth (see below).
5. Parasitic. Several arthropod groups of small size, such as branchiurans, some groups of copepods, and
tantulocarids, have adopted parasitic habits, their hosts including fish and other arthropods (Schram 1986).
In some cases this produces a degenerate adult morphology. The only claim for parasitic habits in trilobites
has been made by Bergström (1973, p. 48) with regard to agnostoids, to which we return below.
PRIMITIVE PREDATORY FEEDING HABITS IN TRILOBITA

The calcified trilobites are a clade descended from a common ancestor during the pre-Atdabanian (early
Cambrian) (Fortey and Whittington 1989; Ramsköld and Edgecombe 1991). It is clearly desirable to
ascertain, if possible, the feeding habits of the most primitive trilobites, so that subsequent adaptive
deployment can be better understood. Cladistic analyses of trilobite relationships are in agreement that
olenelloids (Olenellus and allies) are the sister group of higher trilobites, and these trilobites should be
considered first.
Apart from the antennae, the appendages of olenelloids are not known. Evidence bearing on olenelloid
life habits is perforce indirect, but two lines of evidence point to the same conclusion.
1. Trace fossils. Bergström (1973) and Jensen (1990) have described examples where trilobite tracks have
been seen to terminate ‘worm’ trails, providing direct evidence of predatory activity – although
Bergström’s example has not been universally accepted. In some cases the trilobite dug down into the
layer of sediment through which the ‘worm’ was burrowing, which implies sensory apparatus sophisti-
cated enough to detect subtle vibrations. Jensen (1990) associated the Lower Cambrian hunting trails with
a particular ichnospecies, Rusophycus dispar, made by the hunter. We argue below that certain other kinds
of Rusophycus should be associated with predatory activity.
2. Outgroup comparisons. Another approach considers the sister group of the olenelloids and other
trilobites (Text-fig. 1). If the life habits of the sister group are known then the likelihood is that its closest
TEXT-FIG. 2. Comparative limb anatomy of the Cambrian trilobites, A, Eoredlichia intermedia (Lu), B, Olenoides
serratus (Rominger) and C, Naraoia longicaudata Zhang and Hou (after Ramsköld and Edgecombe 1996), showing
relatively massive basal podomeres associated with predatory habits. Scale bar represents 10 mm.
relatives had the same habits (Coddington 1988). In analyses of the relationships of trilobites to other
arthropods, Naraoia and its allies repeatedly appear as the sister group outboard of olenelloids (e.g. Briggs
and Fortey 1989; Wills et al. 1995). It is possible to regard the uncalcified Naraoia and the trilobites as
derived from a common ancestor, possibly by paedomorphism (Fortey and Theron 1994). The appendage
morphology of Naraoia is well-known (Whittington 1977; see also refinements in Ramsköld and
Edgecombe 1996), and its massive, spiny basal endites (‘gnathobases’) leave little room for doubt that
the habits of some species of Naraoia were predatory: ‘Naraoia appears to have been particularly well-
adapted to capture and to feed on the varied, small, worm-like creatures that abounded in the Burgess
Shale’ (Whittington 1977, p. 438). Chen et al. (1997) have suggested that one Naraoia species from the
Lower Cambrian of China may have rather lived by ingesting sediment, indicating a wider diparity of
feeding habits within naraoiids as a whole. But the evidence derived from the majority of closest-related
sister taxa points to predatory habits for olenelloids also.
In a wider context, Trilobita and naraoiids together comprise a clade within a more inclusive clade of
arachnomorph arthropods, which includes the living Arachnida (chelicerates): for example, limulids,
scorpions and spiders. Virtually all of these animals are also predators and/or scavengers. The major
exception, a number of groups belonging to the mites, are minute and terrestrial. Furthermore, analyses
seeking to place the Burgess Shale arthropods (e.g. Wills et al. 1995) in the context of their general
arthropod relationships show that those arthropods, such as Molaria and Sidneyia, which classify close to
the trilobite clade, are also interpreted as having predominantly predatory habits (Whittington 1985). This
is not likely to be true of other Cambrian possible sister taxa to the arachnomorph clade, such as Marrella
(see Whittington 1971). It is concluded that the habits of the majority of the arachnomorph clade was
predatory from the first, with olenelloid trilobites sharing the same carnivorous predilections. It is now
well-known that by the early Cambrian there was a variety of ‘worms’ available as prey.
PREDATORY TRILOBITES
Trilobites radiated rapidly during the early Cambrian (Briggs and Fortey 1992). However, the Middle
Cambrian trilobite Olenoides serratus, of which the limb anatomy is fully known (see Whittington, 1975,
1980; Text-fig. 2B), shows striking ventral similarity to Naraoia (compare Whittington 1975, fig. 26B and
Whittington 1977, fig. 97; Text-fig. 2C), and the Lower Cambrian naraoiids from China. Its stout spinose
limb bases are equally indicative of a gnathobasic function. Whittington has described how these stout
processes may have functioned as jaws, grasping and shredding worms and other prey and at the same time
passing the food forwards along the ventral medial line to the mouth, which lay at the posterior edge of the
hypostome. Olenoides is of similar dorsal length to Olenellus and may have exploited similar sized prey.
Recently, Shu et al. (1995) described limbs of the primitive redlichioid Eoredlichia from China. Several
details of their reconstruction of limbs of this animal were criticized by Ramsköld and Edgecombe (1996),
who emphasized the similarity of Eoredlichia limbs to those of naraoiids. It, too, is likely to have been a
predator/scavenger. Eoredlichia is particularly significant, because redlichioids of this type are considered
to have been the sister group of a clade comprising all ‘higher’ (non-olenelloid) trilobites (Fortey 1990),
and its life habits may therefore be taken as basal for this group.
Evidence from limb morphology (see Text-fig. 2) thus supports the conclusion that trilobites like
Eoredlichia and Olenoides retained the primitive feeding mode which was likely in Olenellus, and which
was inherited from a still more distant common ancestor of naraoiids and trilobites. Because very few
trilobites have their appendages preserved, however, it is not possible directly to infer life habits in most
taxa. Fortunately, several features of the calcified exoskeleton bear directly upon the life habits, and these
provide a basis for a model of which trilobites retained hunting habits.
TEXT-FIG. 3. Examples of buttressed hypostomes associated with rigid attachment to doublure in the predatory/
scavenging trilobites. Ventral views of A, Cambrian Olenoides serratus (Rominger), B, Ordovician Nileus affinis
Billings and C, Ordovician Niobe morrisi (Billings) (after Whittington 1975 (A), 1965 (B–C)). Such strongly supported
hypostomes are associated with specialized ventral morphology. Scale bar represents 10 mm.
Braced hypostome
In Olenoides, the hypostome is rigidly attached at the cephalic doublure (Whittington 1988a). The mouth
lay at the posteromedian margin of the hypostome, and, not far behind, the anterior pair of post antennal
cephalic limbs were inserted. Coxal movements served to pass food forwards into the mouth. The
hypostome covered the oesophagus, and the space between the frontal lobe of the glabella and the
hypostome provided space for oesphagus and stomach (midgut). In the trilobites, the manipulation of
the food was by means of the basal parts of the limbs, rather than feeding into the mouth by the tips of
the limbs or ‘fingers’, as in the majority of crustaceans. Thus for a trilobite which had to deal with
relatively bulky food, as does a predator or scavenger, it was clearly an advantage to have a buttressed
hypostome, against which the food could be pressed, manipulated and shredded before ingestion. We
regard the very short, terminal spines on the limbs of Olenoides has having been largely sensory.
In trilobites of the order Corynexochida (which includes Olenoides) hypostomal buttressing (Text-
fig. 3) has been taken a stage further by ankylosis of the hypostomal suture. Hypostome and rostral plate
together comprise an even more rigid structure. In Paradoxides and allied genera (see Whittington 1988a)
some species have the rostral plate and hypostome ankylosed, while others do not. Evidence for
anteroventral strengthening of this kind is taken as grounds for suspecting predatory habits.
There is much evidence for hypostome buttressing among post Cambrian trilobites. For example, among
Asaphida both Nileidae (e.g. Fortey 1986; Text-fig. 3B) and Asaphidae (e.g. Whittington 1965; Text-fig.
3C) have extended anterior wings which firmly supported the hypostome against the doublure. Similar
structures have been reported in Scutelluidae, Illaenidae and Phacopida (Whittington 1988b). The
hypostome of trilobites attributed to such groups is often a massive structure carrying an ornament of
strong terrace ridges. Hypostome specializations of advanced trilobites are discussed further below.
Strengthening of anterior cephalic doublure

The cephalic doublure is potentially an area of weakness in the exoskeleton, because it includes the ventral
sutures. Although the sutures were only functional during moulting their traces are seen in entire trilobites
and constitute preferential lines of weakness. The ventral expression of sutures may have become a ‘trade
off’ between the advantages of having a facilitator for moulting, and the disadvantage of a potentially weak
zone. Among predatory trilobites a strong anterior cephalic rim was important. It has been shown that the
hypostomal suture may be lost in corynexochids. However, simplification and strengthening of the
cephalic doublure is typical of other trilobites with buttressed hypostomes. The rostral plate in Asaphida
has been lost, to be replaced by a single, median suture. In some families this process has been taken
further, in that the suture itself is lost in advanced members, which then have the free cheeks yoked
together as a single unit. This is obviously the strongest arrangement of all. There is one example in the
asaphid Isotelus having fused the median suture late in its ontogeny. Fused cheeks combined with wide
cephalic doublure are typical of Nileidae and phacopoids. The changes in such trilobites may be contrasted
with conservatism of ventral sutures in trilobites having unattached (natant) hypostomes, in which a rostral
plate and connective sutures are retained, which also implies a functional explanation connected with
strengthening the border in those trilobites that do change.
Hypostome attachment style

Among Cambrian trilobites with hypostomes rigidly attached to the doublure the attachment style is
invariably of the kind described by Fortey (1990) as conterminant. This means that the anterior furrow
defining the front edge of the glabella coincides ventrally with the front margin of the hypostome. There is
a close match between the profile of the middle body of the hypostome, and the frontal lobe of the glabella.
This morphology implies that the foregut and stomach were confined within a volume circumscribed by
the area inside the glabellar furrows (which connected with the musculature of the cephalic limbs),
anteriorly by the preglabellar furrow, and ventrally by the middle body. It is noted that those trilobites with
strongly buttressed conterminant hypostomes are also often those in which the frontal lobe of the glabella
becomes enlarged – often becoming relatively wide or pyriform. This happens polyphyletically. Typical
examples are Paradoxides, Asaphus, some lichids, and many Encrinuridae. The expansion served to
enlarge the area available for the stomach cavity, and thus modified the constraints imposed by the fixture
of the hypostome at the front of the glabella. More food could be accommodated. It is interesting to
observe that some of the trilobites with fixed hypostome and fowardly expanding glabellas are also among
the largest species known, for example Paradoxides and Uralichas.
Among later and more advanced trilobites the close relationship between glabella and hypostome can be
lost. In the impendent condition (Text-fig. 4A) the glabella has expanded forwards, such that its front edge
now lies well in front of the anterior edge of the hypostome. This conterminant–impendent trend occurred
several times (Fortey 1990), for example, in Phacopida and advanced Proetida, invariably associated with
forwardly expanding glabellas. It is a reasonable interpretation that in these trilobites the volume available
for ingestion of food has increased, perhaps enabling the trilobite to deal with larger prey items, or at least
store more food for subsequent digestion. Note that Fryer (1968, fig. 111) recorded a comparable increase
in stomach size in the one predator among an otherwise mostly herbivorous or detritus-feeding group of
cladocerans. In that impendent morphology is also associated with braced hypostomes it does seem to be
reasonably explained within the context of predatory habits. Typical impendent morphology is present in
Phacopida and Proetida of Devonian, Carboniferous and even Permian (see Owens 1983) age, and it can be
concluded that predatory habits remained an option until the end of the history of the group.
TEXT-FIG. 4. Lateral views of cephala showing hypostomal attachment styles within the body cavity. Heavy shading
shows section through hypostome revealing attitude relative to glabella and cephalic doublure, stipple shows inferred
position of mouth (m) and gut. A, impendent hypostome showing enlarged glabella extending anterior to front edge of
hypostome; B, conterminant hypostome attached to doublure and anterior edge coinciding with that of glabella;
C, natant hypostome, detached from doublure, sited beneath anterior end of glabella; D, conterminant hypostome in
illaenimorph, which is reflexed upwards inside cephalic cavity such that mouth is well above lateral cephalic margin.
Eldredge (1971) described the strong muscle impressions that are developed on the frontal glabellar
lobe, and hypostomal middle body in Phacopina. They were interpreted as insertion areas for musculature
serving to manipulate the stomach and foregut; additionally, some muscles were inserted to offer further
support for the hypostome. Campbell (1975a) did not accept the latter. He showed (1975a, fig. 2) ‘muscles
for raising the hypostome’; however, if Whittington (1988b) was correct in claiming that the hypostome in
Phacopida is fixed rigidly then these muscles must, rather, have served to dilate the foregut. Upon the
interpretation given here, this evidence of strong musculature would combine with the tumid glabella of
phacopids, their impendent hypostomal condition, and the supported, attached hypostome, to suggest
advanced predatory habits. It is conceivable that the enlarged stomach was modified as some kind of
gastric mill which required stronger muscle support for its operation. Both Campbell (1975a) and Eldredge
(1971) inferred the existence of ‘digestive glands’ opening into the stomach. Eldredge (1971), however,
believed that these animals were essentially particle feeders of the kind typified by cephalocarids (see
above). We find this interpretation is not so well-supported in the context of the Trilobita as a whole, as
argued here. Miller (1976) and Stockton and Cowen (1976) also presented arguments that phacopids were
predators, based upon their highly developed sensory fields, and their sophisticated visual system,
respectively. Whether or not the visual system operated stereoscopically as Stockton and Cowen claimed,
it cannot be questioned that phacopid eyes are highly evolved structures (Clarkson and Levi-Setti 1975)
and that high visual acuity is at a premium in predatory animals.
Modifications of the posterior hypostome margin

If buttressed hypostomes are associated with the manipulation of bulky food it might be expected that
trilobites with this kind of hypostome should show appropriate modifications. Because the mouth lies at
TEXT-FIG. 5. Modification of posterior margin of various hypostomes in inferred predator/scavenger trilobites. A, lateral
spines in Paradoxides gracilis (Boeck) (after S̆najdr 1958, pl. 14, figs 1–5); B, postero-median ‘grinder’ in Acidaspis
jessi Perry and Chatterton (after Chatterton and Perry 1983, pl. 23. fig. 12); C, fork in Isotelus gigas DeKay (after
Rudkin and Tripp 1989, text-fig. 2); D, narrow fork in secondarily conterminant Griffithides acanthiceps Woodward
(after Fortey 1990, text-fig. 5b); E, fork in Warburgella rugulosa canadensis Ormiston (after Ormiston 1971, pl. 20,
figs 9–11); F, fork – with rhynchos on middle body – Calymene polgari Siveter and Chatterton (after Siveter and
Chatterton 1996, pl. 3, fig. 2); G, exceedingly long fork extending beneath thorax in Hypodicranotus striatulus
(Walcott) (after Ludvigsen and Chatterton 1991, fig. 2). Various magnifications.
the back end of the hypostome it is the posterior part of the hypostome which will become the focus of any
changes.
Such modifications are extremely common (Text-fig. 5). The change most often seen is the
development of a fork. This is present already in rather weak form in the Mid Cambrian Paradoxides
(Šnajdr 1958, pl. 21, fig. 8), and quite strongly in somewhat younger dameselloideans (Öpik 1967, pl. 51,
fig. 1). The development of the fork is particularly well known in the Asaphidae. Primitively, no fork was
present (e.g. Asaphellus, see Fortey and Owens 1991, fig. 8b); in some Niobinae a shallow notch-like fork
occurs (Whittington 1965, pl. 29, fig.2); in Isotelinae the fork is deep with a broadly U-shaped outline
(Text-fig. 5C). In these trilobites the anterior wing of the hypostome is especially powerfully developed,
indicating robust bracing. In many of these examples the fork is associated with comparatively wide lateral
hypostomal borders. This is particularly the case in Lichida. Shallow forks are developed in Calymenidae
and Homalonotidae. The fork may have acted against the bases of the anterior cephalic limbs as an aid to
processing bulky food. Longer forks may have helped prevent prey escape. The longest fork of all is
present in the remopleuridid Hypodicranotus, in which it extends along almost the entire venter of the
body (Whittington 1952). This genus is interesting in showing how the median ventral line remains
uncovered, presumably to permit the free operation of the limb bases. It also confirms that the appendages
lay above the hypostome, in that it is only in this position that the musculature of the limbs could be
effectively inserted into their axial apodemes.
Fortey (1990) pointed out how conservative the unattached (natant) hypostome is in trilobites from the
Cambrian to the Carboniferous. It is interesting to observe how in those Proetida and Asaphida in which
the hypostome has subsequently docked with the cephalic doublure (secondarily conterminant), forked
posterior margins have developed. A good proetoid example is Carboniferous Griffithides acanthiceps
TEXT-FIG. 6. Rusophycus trace fossils definitely associated with

trilobites assumed to have had predatory/scavenging
habits. A, from Grès Armoricain (Ordovician, Arenig) Camp Français, Laval, Mayenne, France; (150 mm long),
associated with the asaphid Ogygyinus armoricanus. (Photo courtesy G. L. Pillola). B, from the Mithaka Formation
type section, Georgina Basin, Queensland (see Draper 1980) (Ordovician, Llanvirn), associated with large Lycophron
sp. (Asphidae); (340 mm long). Note limb base impressions and doublure.
(Fortey 1990, text-fig. 5B; Text-fig. 5D). There are some interesting species in other Proetoidea in which
the preglabellar field is absent, or has become very short. In these, sutural junction between the front
margin of the hypostome and rostral plate has not, apparently, been taken to completion, because examples
with the hypostome preserved in place against the doublure are unknown. Nonetheless, in some cases their
hypostomes exhibit modifications associated in other trilobites with predator/scavenger morphology.
Warburgellinae, for example, have hypostomal forks (Text-fig. 5E). Cordania shows a posterior median
extension of the doublure which ‘eliminates’ the preglabellar field, and Myoproetus has a triangular,
phacopid-like hypostome (Šnajdr 1980). These proetoideans also tend to have the kind of varied sculpture
seen on hypostomes of predator/scavengers. It seems likely to us that these were, indeed, trilobites with
this life habit, but in which the process of secondary buttressing of hypostome against doublure had not
proceeded to completion. The connection between hypostome and doublure may have been ligamental
(see also Whittington and Campbell 1967, p. 456). Whether these trilobites employed a certain flexibility
along the hypostome/doublure joint as part of the feeding process is something that would repay
investigation.
Further evidence is provided by sculptural modifications to the posterior median region of attached
hypostomes. Chatterton and Perry (1983) described a pair of roughened tuberculate areas in the middle of
the border of the hypostome of odontopleurids. These are burr-like structures situated exactly where they
would be of service in grinding and comminuting food. On the inner surfaces of the fork of the Ordovician
asaphid Isotelus there is a series of fine raised ridges (Tripp and Evitt 1986, fig. 6A; Rudkin and Tripp 1989;
Text-fig. 5). These regular ridges are present nowhere else on the exoskeleton and are obviously of
functional significance. They resemble a rasp, and it is feasible that they had a role in breaking down the
body wall of prey before ingestion into the stomach. Similar structures occur on some lichids.
Trace fossils
There is a variety of possibilities for trace fossils to be produced by trilobites during surface scavenging or
hunting, but only those which involved digging in the sediment are likely to be distinctive enough to be
associated with this life habit in particular. The olenellid hunting trail described by Jensen (1990) has been
mentioned previously. However, if it is correct that prey was manipulated by limb ‘gnathobases’, then we
might anticipate that impressions of limbs should be preserved. Trilobites that habitually caught prey
within the sediment should, in favourable circumstances, be associated with trace fossils showing evidence
of limb bases. For example, Rusophycus traces have been closely associated with large asaphids in the
early mid Ordovician Mithaka Formation of central Australia (Draper 1980; see also Text-fig. 6B), and
indicate that these trilobites excavated shallow burrows in arenaceous sediments. Not only are the limb
bases clearly visible, but the impression of the back end of the hypostome can be seen in some cases.
Because these asaphid trilobites have well-developed hypostomal forks, and buttressed doublure, and are
also very large, they conform closely to the predatory model. Interestingly enough, the Australian
examples are associated with a variety of infaunal ‘worm’ (annelid, polychaete, etc.) burrows, the
occupant of any one of which could have provided a prey item.
Summary. Trilobites with hypostomes rigidly attached to the cephalic doublure include large species
which develop distinctive features of the glabella and hypostome. These features are consistent with
predatory or scavenging habits, and can be interpreted as modifications to cope with greater ingestion
of food or its processing prior to consumption. Some of these predators developed stalked eyes
(homalonotids: Hammann 1983; Whittington 1993; Cybelinae: Ingham 1968), and were capable of a
semi-infaunal life. Although predatory habits were primitive, some advanced trilobites acquired
secondarily the appropriate morphology, which continued even among Permian species. Rusophycus
showing coxal impressions are associated traces. Trilobites of the following groups show morphology
consistent with dominant habits of this kind: Paradoxidoidea, Corynexochida, advanced Asaphida,
Lichida, Odontopleuridae, Calymenoidea, Phacopina and secondarily conterminant Proetida.
PARTICLE FEEDING
Because their source of nourishment was mostly minute and in a finely divided condition, trilobite species
that lived by particle feeding did not require the strong hypostomal support of their predatory
contemporaries. They are potentially very numerous as individuals, both because they occupy a lower
position in the food chain (the classical ‘food pyramid’), and because the main limiting factor on their
abundance is likely to have been the availability of nutrients in the form of detritus in the substrate. In some
environments – such as the outer part of the continental shelf in spring – nutrients are prolific and
populations of particle feeding crustaceans or molluscs can be dense. However, because of the small
particle size and the comparatively unfavourable ratio between energy expended in gathering nutrients, and
the low food value of those nutrients, the large sizes attained by the predator/scavenger are not expected.
The characteristic morphology of a particle feeding trilobite includes the following features.
Natant hypostome
In contrast with those trilobites discussed above in which the hypostome was both attached to the doublure
and rigidly supported, in many trilobites it became detached and lost skeletal support at the doublure. It
was, however, probably still attached ligamentally to the anterior wings (Whittington 1988a). In this natant
condition the hypostome was supported mainly by the ventral membrane. The position of the natant
hypostome in life still corresponded ventrally with the forward part of the glabella. Because of this mode
of support the hypostome easily became detached upon the death of the trilobite, and in situ occurrences of
natant hypostomes are uncommon. However, the natant condition can be inferred directly from cases
where trilobites died ‘on their backs’ (Chatterton and Ludvigsen 1998, figure many examples; see also
Text fig. 9B), or in which the hypostome can be prepared out, and indirectly if the length of the preglabellar
field exceeds that of the anterior cephalic doublure. Fortey (1990) reviewed the distribution of natant
hypostomes in trilobites and identified 30 families in which it may have been present.
TEXT-FIG. 7. The conservative nature of natant hypostomes associated with particle feeding habits. A, Ptychoparia
striata (Emmrich); Middle Cambrian, Czech Republic (from S̆najdr 1958, pl. 39, figs 1, 4; pl. 40, fig. 3).
B, Aulacopleura koninckii (Barrande); Silurian, Wenlock Series, Czech Republic (from Hughes and Chapman 1995,
figs 1G, 3). Scale bars represent 10 mm (A) and 5 mm (B).
The hypostome continued in its rôle as a ventral plate covering foregut and stomach. However, because
the natant hypostome was not rigidly attached, muscles operating on the hypostome were capable of
moving it. The natant hypostome is built very much to a common design (Text-fig. 7): elongate (sag.),
having narrow borders, a rounded posterior margin and, at or behind midlength, a pair of middle furrows.
Like the dorsal glabellar furrows, it is likely that the middle furrows were the sites of muscle attachment.
This being the case, muscular contraction of a pair of muscles to either side of the mouth would have
elevated the hypostome. But, the anterior wings of the hypostome were likely to have been tethered by
ligaments to the anterior pits or fossulae in the axial furrows (Whittington 1988a, 1988b). The presence of
such wings, and their correspondence with the fossulae, is a general character of Trilobita (see Fortey and
Whittington 1989). Accordingly, muscle contraction would not elevate the whole hypostome, but only the
posterior part. This would produce a lifting motion. The foregut and/or stomach would be squeezed in an
upwards direction at the same time. None of these motions would have been possible unless the hypostome
was free-lying, and would assuredly have been impossible for any trilobite with conterminant or
impendent hypostomal attachment. This suggests that the evolution of the natant condition was an
important adaptive innovation.
We envision that the lifting motion of the natant hypostome aided the forward passage of food material
into the digestive system. Although the movement was as much upwards as backwards, constriction of the
posterior part of the oesophagal sac would have impelled the relatively fluid food forwards. This may in
turn have forced the food around the anterior turn in the digestive tract and into the main gut, possibly
aided by peristaltic contraction. How rapidly these movements operated is not known. However, many
natant hypstomes show a short posterior wing (Text-fig. 8) situated a little way behind the middle furrow,
and it is possible that another set of muscles attached to these. If muscles inserted here were opposed
TEXT-FIG. 8. Musculature and operation of natant hypostomes employing a ‘scooping’ motion to collect nutrients.
A, muscles relaxed, with hypostome in ‘down’ position, anchored by interior wings to anterior fossulae; B, muscles
contracted, posterior margin of hypostome elevated to ‘tethered’ anterior wings, inducing pressure of oesophagus
aiding ingestion of food.
(antagonistic) to those on the middle body they might between them have set up a relatively rapid pumping
action. Alternatively, the hypostomal muscles may have contracted from time to time to swallow food
when the oesophagus was full, the hypostome being restored to its ‘down’ position as the muscles relaxed.
Conservative hypostomes
A remarkable feature of the hypostomes associated with the natant condition is their extreme conserva-
tism. Middle Cambrian examples are much like those from the Carboniferous. This conservatism is
consistent with the notion that the hypostome did not play an active part in shredding or manipulating food,
which in turn accords with a particulate diet. As we have seen, this is in contrast with attached hypostomes,
with their array of forks and peripheral modifications. A few hypostomes have delicate posterolateral
spines which are in the right position to have been used as a grooming device for the cephalic appendages
(a gnathobasic toothpick?). Otherwise the pattern of elongate middle body and narrow borders is universal.
There is little evidence of inflation of the middle body, and one must assume that no advantage was to be
gained from storing additional material in the stomach.
Reduced hypostomes?
In some olenids which have died ‘on their backs’ the hypostome has fallen down into the cephalic cavity.
Examples of Porterfieldia puctata (Crosfield and Skeat) from the Arenig of South Wales (see Fortey and
Owens 1978) are illustrated here (Text-fig. 9) to show this mode of preservation. In these specimens it is
extremely likely that the hypostome in question belongs to the trilobite beneath because they came to rest
in an identical position. Henningsmoen (1957, pl. 26, fig. 2) figured a specimen of Peltura scarabaeoides
showing the same. In both of these cases the hypostome is remarkably small, and even the anterior wings
appear to be poorly developed. Certainly, the hypostome does not occupy an area equivalent to that of the
frontal glabellar lobe in such species. Its protective function was therefore probably reduced. Olenids
apparently had thin cuticles (Fortey and Wilmot 1991) and do not seem to have required exoskeletal
protection from predators. This may have been because they lived in specialized, oxygen depleted
palaeoenvironments.
It is clear that in such olenids, even more than in other natant trilobites, the manipulative function of the
hypostome was reduced. Ingestion of food may have been achieved largely by muscular contraction, with
the hypostome acting as little more than a ventral lip. This introduces a further possibility: that the
hypostome may have become completely decalcified in some species. After all, there are many
libristomate trilobites for which no hypostome has been assigned. Whilst this loss is conceivable, many
examples of species known from large fossil populations having particle feeding, natant morphology have
TEXT-FIG. 9. The olenid Porterfieldia punctata (Crosfield and Skeat). A, NHM It26536 dorsal, × 4, B, NHM It26537
ventral, × 3, views, the latter with hypostome fallen into cephalic cavity in approximate life position, showing its
relatively small size in this natant (presumed particle feeding) trilobite. Compare specialized hypostomes (Text-fig. 5).
also eventually yielded hypostomes in place or in close association (see, for example, the type
ptychopariid, Ptychoparia striata (Emmrich): Šnajdr 1958, pl. 39, fig. 7; Text-fig. 7A). This is true even
of advanced olenimorphs such as Aulacopleura (Text-fig. 7). Hence it would not be prudent to assume that
complete decalcification of the hypostome occurred, although it remains a possibility worth investigating.
‘Sluggish’ morphology
Many natant trilobites are olenimorphs (Fortey and Owens 1990), that is, they had a comparatively low,
narrow axial region including a small area for muscle insertion, and often large numbers of narrow (exsag.)
thoracic segments and a small pygidium. They are neither well streamlined, nor vaulted transversly in the
fashion associated with possible infaunal habits (e.g. homalonotids, see Hammann 1983; Whittington
1993). Genal spines, which are usually present, are not particularly strong, and may be needle- or blade-
like. It is considered likely that such trilobites were feeble swimmers, and that they were habitually part of
the vagrant benthos. It is interesting to note in this connection that few trilobites with natant hypostomes
have stalked eyes (Devonian Cyphaspis species are an example). Most of those that do also have attached
hypostomes, such as asaphids (Asaphus kowalewskii), most Cybelinae and many odontopleurids and
homalonotids. All these facts are consistent with rather sluggish, surface feeding for those trilobites with
natant hypostomes. Some of these trilobites, including those olenids that Henningsmoen (1957) termed
‘Peltura-type’, and many Ordovician–Carboniferous proetoideans, are more compact and convex, and
may have been capable of faster swimming (e.g. Fortey and Owens 1991). These tend to be part of more
diverse trilobite faunas, such as those from the Silurian (Wenlock) mentioned below, rather than dominant
members of low diversity faunas. It may be that rapid escape was more important for particle feeders living
in such communities.
High relative abundance

Trilobites with natant hypostomes which are inferred to have been particle feeders are common as
individual fossils and commonly dominate trilobite faunas. In the mid Cambrian Wheeler Shale of Utah
the typical ptychoparioids Elrathia kingi and Asaphiscus wheeleri are abundant on some bedding planes
compared with many other species (see Gunther and Gunther 1981, frontispiece, and inside front covers
for examples). Another natant olenimorph Alokistocare is second in abundance to Elrathia in the Wheeler
Shale according to Robison (1964). Whilst a census which may include moults as well as entire individuals
can be misleading as regards original population structure, it is at least certain that during the deposition of
parts of the Wheeler Shale natant trilobites greatly predominated over corynexochids with attached
hypostomes. This dominance may not be paralleled by species richness; e.g. about half the species from
the Utah Middle Cambrian figured from complete specimens by Gunther and Gunther (1981) probably had
attached hypostomes. The abundance of olenids within the nodules in the late Cambrian Alum Shale of
Scandinavia is remarkable, as Westergård (1922) noted (see the wonderful example shown by Hennings-
moen 1957, pl. 30). Commonly, these concentrations of articulated individuals are mono- or bispecific
(Clarkson and Taylor 1995). Fortey’s (1975a) quantitative data on the Olenid Biofacies in the early
Ordovician of Spitsbergen showed that species of the eponymous family comprised 90 per cent. or more of
the preserved fossils within a given sampling interval, whilst Owens (1979) described a monospecific
accumulation of the proetoidean Cyamella from the Ashgill Boda Limestone of Dalarna, Sweden. All
these species were probably natant. Another example is afforded by the faunas recently described from the
top of the Tremadoc Shineton Shale Formation, Shropshire, by Fortey and Owens (1991). Of more than
500 specimens collected the diminutive Shumardia comprises more than 40 per cent. of the total. Fortey
and Owens figured the hypostome and doublure of this trilobite, which was probably natant. The unusual
orometopid Skljarella accounts for about 20 per cent. of the fauna and may also have been natant. Both
Shumardia and Skljarella can be numerous on single bedding planes (Fortey and Owens 1991, figs 8r,
10q). The agnostid Litagnostus may also have been a particulate feeder (see below) and accounts for 25 per
cent. of the fauna. Conterminant asaphids and remopleuridids are dominant among the rest. Even in strata
younger than Ordovician natant trilobites may be extremely abundant. For example, the late proetoidean
olenimorph Aulacopleura koninckii is the commonest trilobite in the Bohemian Silurian Liten̆ Formation
comprising perhaps 60 per cent. of the articulated material there (Hughes and Chapman 1995). An Upper
Palaeozoic occurrence is a monospecific accumulation of Phillibolina from the Carboniferous of northern
England (Owens 1990).
TEXT-FIG.10. Group of Cyphaspis elachopos Thomas from the Much Wenlock Limestone Formation (Silurian)
Dudley, West Midlands; NHM 46430; × 3. Group of particle feeding trilobites in a fauna in which such species are
generally rare.
However, it is certainly not the case that natant forms always dominate in the total fauna. In some fossil
assemblages with a high proportion of complete specimens, which are likely to reflect naturally co-
occurring taxa, natant forms may comprise quite a small proportion of the total. In collections from the
Much Wenlock Limestone Formation of Dudley, West Midlands, kept in The Natural History Museum,
London, we have estimated that only about 15 per cent. of the articulated specimens (comprising proetids
and aulacopleurids) were natant. This is probably a reasonable sample, because complete Dudley trilobites
were collected exhaustively in the nineteenth century and were conspicuous enough not to have been
missed. But even within this fauna there are certain bedding planes on which natant trilobites appear to
have been particularly common (see Text-fig. 10), such as those with Harpidella. A Dudley collection
(n ¼ 90) at the National Museum of Wales from a single bedding plane in the Nodular Member of the
TEXT-FIG.11. Relative proportions of trilobite species with natant (fine stipple) and conterminant (coarse stipple)
hypostomes in populations from: A, Ordovician, Tremadoc Series, Shineton Shale Formation, Shropshire (sample from
Coundmoor Brook, see Fortey and Owens 1991); B, Silurian, Wenlock Series, Liten̆ Formation, Czech Republic (after
Hughes and Chapman 1995); C, Wenlock Series, Coalbrookdale Formation, Shropshire (based on NMW collections);
D, Wenlock Series, Much Wenlock Limestone Formation, Dudley (based on NHM collections); E, Wenlock Series,
Nodular Beds Member of Much Wenlock Limestone Formation, Dudley (based on NMW collections). These diagrams
demonstrate differences in distribution of major feeding types between different trilobite faunas.
Much Wenlock Limestone Formation made by RMO probably represents a natural population, and
comprises 70 per cent. conterminant species and 30 per cent. natant, proportions of a similar order of
magnitude to estimates from the undifferentiated collections. By contrast, a large collection in the National
Museum of Wales made by M. Rowlands and P. Tarrant from the Coalbrookdale Formation (Silurian,
Wenlock Series) from a locality near Buildwas, comprises 790 individuals of which 98 per cent. were
probably conterminant and only 2 per cent. natant.
It is proposed that the numbers of particle feeding trilobites were limited by the availability of organic
particulate matter in the substrate; when this material was abundant so were the appropriate small- to
medium-sized natant trilobites. In the Cambrian–Ordovician Olenid Biofacies, trilobites of this family
were overwhelmingly dominant. These thin-shelled trilobites were probably protected from predators in
an oxygen-poor (dysaerobic), but nutrient rich environment (Fortey and Wilmot 1991) although they
teetered always on the edge of complete anoxia. Where more than one natant species co-occurs in the
examples mentioned above smaller species appear to outnumber larger ones, but there is always the
possibility of sampling error in favour of the former.
Uncalcified anatomy
There is no satisfactory information on the appendages of those trilobites claimed as particle feeders
(although see remarks on Naraoia above). The closest approach may be the olenid Triarthrus, the limbs of
which were fully redescribed by Whittington and Almond (1987). Triarthrus is the only olenid known to
us in which the hypostome probably became attached to the doublure (i.e. secondarily conterminent), and
it has no preglabellar field, so it cannot be assumed to represent natant trilobites. However, virtually all of
its relatives were probably natant, and it is the best surrogate that we have. There are some observations
that are consistent with particle feeding. Whittington and Almond (1987) showed that the limbs continued
all the way posteriorly to form a rather tight post-cephalic feeding chamber. The horizontal, overlapping
disposition of the lamellar exites shows that they could not have engaged directly with the sediment or the
feeding process, and they are interpreted as having had a respiratory function, although their beating
motion may have assisted ingress of currents. However, it is noticeable that proximal podomeres of the
thoracic legs are equipped with remarkably long, inward-facing spines. The distal joints towards the tip of
the legs are not so equipped (Text-fig. 12). By contrast, the cephalic leg joints are smooth, other than
massive gnathobasic ‘coxae’, which are far more robust than those on the thorax. This apparently odd
pattern becomes explicable if the setose thoracic limbs are regarded as the sorting appendages for food
particles; meanwhile the cephalic gnathobases functioned only to pass the sorted food to the mouth. The
absence of setae on the podomeres of the cephalic limbs does point to a differentiation of function between
thoracic and cephalic limbs, while the prolific spinosity of the former is consistent with a rôle in particle
sorting. It may have been the case that, as in certain cladocerans (Fryer 1968), the cephalic limbs were used
to pull the animal along while the posterior limbs processed the sediment for food.
TEXT-FIG. 12. Limbs of Triarthrus eatoni (Hall); A, from cephalic region with massive gnathobasic coxae for passing
food to the mouth, and B, from thoracic region with long, inward-facing spines for food sorting. Exites omitted. After
Whittington and Almond (1987, fig. 39a–b).
Trilobites with well-developed genal caeca are often also natant (e.g. Papyriaspis). Such caecal patterns
have been attributed to digestive glands (Öpik 1961; Cisne 1975; Chen et al. 1997), and, if so, might be
supposed to have yielded secretions which aided digestion of the particulate organic material sorted by the
trunk limbs. Such a digestive system might account for the comparatively small volume of the stomach in
these trilobites; much of the work would have been done by digestive enzymes. However, Jell (1978)
claimed that caecal patterns related to circulation rather than to digestion; similarities to the arterial pattern
found in Limulus had been pointed out by Bergström (1973) and Fortey (1974). In cases such as Penarosa
(Jell 1977) where the caecal pattern extends all the way to the pygidium it is not likely that the caeca were
secretory organs linked to the digestive system, since they so closely approach the anal region.
Trace fossils
It is probable that those continuously ploughed trace fossils referred to the genus Cruziana might be
associated with particle feeding from organic rich sediments. It is, of course possible that any arthropod
having this mode of feeding could have produced Cruziana, and it is accepted that there were other
trilobitomorphs capable of doing so during the Palaeozoic. We do not wish to repeat the arguments for or
against trilobite Cruzianas here. However, it can be said that trilobites are likely to have made Cruziana of
the semiplicata type (Text-fig. 13), which retain parallel grooves on either side, consistent with the
TEXT-FIG. 13. Feeding traces made by particle feeding trilobites; Cruziana semiplicata Salter; Upper Cambrian.
A, NMW 27.110.G6; Merioneth Series, Marchlyn Formation, Carnedd y Filiast, south of Bethesda, Gwynedd;
× 0·5. B, NHM TF1; Andam Formation, Sultanate of Oman (see Fortey and Seilacher 1997); × 0·5. Note the parallel
grooves to either side of each track, particularly on B.
presence of genal spines. Fortey and Seilacher (1997) have shown that there is good field evidence to
associate C. semiplicata with the late Cambrian trilobite genus Maladioidella. This ‘ptychoparioid’ genus
has a long preglabellar field, relatively narrow cephalic doublure, and was thus very likely to have had a
natant hypostome. Furthermore, the palaeogeographical distribuition of Maladioidella coincides with that
of Cruziana semiplicata in dominantly clastic sediments of what was Palaeozoic Gondwana. Since the
majority of natant trilobites also had genal spines it would be expected to find at least some indication of
them on any Cruziana trail produced during the course of feeding on sediment.
Summary. Trilobites having the hypostome detached from the cephalic doublure in the natant condition
show specializations consistent with a mode of life as organic particle feeders on soft sediments. They are
dominant in some faunas. The typical morphology (‘ptychoparioid’ or ‘olenimorph’) was an enduring one,
having evolved in the early Cambrian and persisting until the Carboniferous. However, this is not to imply
that it was conservative, rather, it was a successful specialization that lasted. They may have been
responsible for excavation of continuous feeding trails showing distinct impression of genal spines,
classified as Cruziana of semiplicata type. Typical trilobites with this kind of morphology are found in
Ptychopariina, Olenina, and primitive members of Asaphida and Proetida.
TEXT-FIG. 14. Some trilobites for which filter-chamber feeding is postulated. Dorsal and lateral views of: A–B, Harpes
macrocephalus Goldfuss (after Whittington, in Moore 1959, fig. 3. 19a–b); C–D, Uromystrum validum (Billings) (after
Whittington, in Moore 1959, fig. 284·2a–b); E–F, Paranorwoodia venusta Rasetti (after Rasetti 1945, pl. 61, figs 24–25
and Ludvigsen et al. 1989, pl. 39, fig. 5); G–H, Cordania falcata Whittington (after Campbell 1977, pl. 5, fig. 3a–b);
I, sagittal section through Uromystrum validum, showing inferred position of hypostoma. See also Text-figure 16,
Cryptolithus tesselatus (Green), which is a further example of this morphology.
FILTER CHAMBER FEEDING

A third major feeding strategy is proposed for trilobites which utilized a cephalic feeding chamber. These
trilobites are considered to have brought sedimentary material into suspension before selecting out edible
particles. For this purpose the cephalic area was modified into a ventral feeding chamber for filtration. In
all of the trilobites discussed previously, the mouth (i.e. the median rear end of the hypostome) was more-
or-less on the same plane as the inner parts of the limbs and the cephalic margin. In the filter feeding group
the cephalic margin, usually developed as a brim, lies at a level well below the mouth, and the space
between brim and mouth is available as a filtration chamber. Hence the typical filter feeding morphology is
where the sagittal convexity of the cephalic shield exceeds that of thorax and pygidium, while the cephalic
shield may be extended into a brim, often with long, broad genal prolongations (Text-fig. 14). A change in
TEXT-FIG.
15. Lateral view of Ampyx nasutus Dalman, lower–middle Ordovician, Sweden, showing position of
hypostome (stippled). If mouth (m) lay at inner end of hypostome then it was elevated inside the cephalic chamber.
Modified from Whittington 1988b, figure 16.
the orientation of the hypostome is known in some species, and increases the available volume for filtration
still further. This feature will be considered first.
Upturned hypostome
In the trilobites so far considered in this paper the hypostome has a more-or-less horizontal orientation, that
is, it might be considered as a ventral extension of the exoskeleton on a plane with the cephalic doublure.
However, there is a number of cases where the hypostome has become sharply reflected upwards inside the
glabella. This orientation was proved in Illaenus by Bruton (1968) and in Symphysurus by Fortey (1986),
both trilobites with particularly convex (sag.) cephalic shields. Here (Text-fig. 15) the same phenomenon
is illustrated in the raphiophorid Ampyx. In this genus, the glabella is convex anteriorly, and much more so
(tr.) than the thorax and pygidium, which were held in an elevated position. The hypostome slopes
upwards steeply on the inside of the glabella close to its anterior face, beneath the frontal median spine. It
is attached to the narrow cephalic doublure in conterminant mode. Note that in this position the posterior
end of the hypostome lies close to a horizontal line produced backwards from the thoracic axial furrows,
and therefore also presumably on the operating plane of the coxal bases of the thoracic limbs. The result of
the configuration of the cephalic shield is a considerable volume beneath genae and glabella available as a
filter chamber.
If the mouth were at the posterior end of the hypostome in its usual position it is difficult to see how
Ampyx could have fed other than by filter feeding. The mouth could not have engaged closely with the
sediment surface, nor is the orientation of the hypostome in a suitable position for the manipulation of
bulky food (cf. predatory morphology). Furthermore, the hypostome comes so close to the dorsal glabellar
surface that the foregut must surely have been shortened, to curve posterodorsally almost immediately
behind the mouth. Presumably the stomach was accommodated in the inflated frontal lobe of the
glabella.
Although the hypostome is not known in place in many genera with trinucliemorph design it is clear
from the morphology of the cephalic doublure that they, too, must have had hypostomes suspended well
above the marginal rim of the cephalic shield. For example, in harpetids (see Whittington 1988b) the lower
lamella curves upwards underneath the brim such that its inner edge lies at the edge of the genal fields, far
above the anterior and lateral edges of the cephalic shield. The hypostome (and mouth) must have been
suspended within the cephalic chamber, even if it were attached more-or-less horizontally at the doublure,
and it may have been inclined even farther upwards in the manner of Ampyx. Certain trinucleids show a
similar arrangement of fringe and lower lamella.
Filter chamber morphotypes

Although the example just given provides the most completely known case, there are many trilobites likely
to belong to the filter feeding group for which the hypostomal position is not yet described. These share a
convex, chamber-like cephalic shield with a long frontal area, and a ‘suspended’ thorax and pygidium
conspicuously less convex than the cephalon (tr.). For this reason, they are most easily recognized in
lateral views of articulated material (cf. Campbell 1975b, fig. 3). The appropriate combination of
morphological features is found in almost all relatives of Ampyx classified in Trinucleina. Those species
(Trinucleidae, Dionididae) with a perforated cephalic fringe were termed trinucleimorphs by Fortey and
Owens (1990). Harpetids show a similar morphology, although the genal spines in many species are more
conspicuously extended posteriorly to form a ‘wall’ alongside the thorax. The filter chamber is thus
enclosed by steep genal extensions. Whittington (1988b) showed that given the wide cephalic doublure in
harpetids and its close reflection against the dorsal surface of the brim, then, as in trinucleoideans, the
hypostome was elevated above the level of the cephalic margin, in the manner typical of filter feeders.
Other families include one or two genera only which exhibit the appropriate morphology. Among
Bathyuridae (Ordovician) Uromystrum (see Whittington 1953; Shaw 1968) differs from other members of
the family in the convexity (sag.) of the cephalic shield, and the genal spines are swept backwards in the
same fashion as described for harpetids. It is also clear that the thorax and pygidium were held in the
‘suspended’ position. Where the genal spines have this enclosing form it is likely that the thorax was also
enclosed into the filter chamber. Whittington (1963) described the genus Harpillaenus, which is an illaenid
convergent upon the same morphology. The only proetoideans which may be close to this design are
members of the Brachymetopidae, for example the Devonian species Cordania falcata (see Campbell
1977, pl. 4, fig. 2). The enigmatic family Ityophoridae may furnish additional examples. It is sufficiently
clear that this characteristic morphology was capable of having evolved more than once and that filter
feeding happened polyphyletically. All the taxa mentioned are likely to have had what Bergström (1973)
described as ‘basket and lid’ enrolment style. It will be interesting to discover whether the hypostome
proves to have the same orientation as in Ampyx in these examples.
In the typical filter feeding design the low vaulting of the thoracic axis and its narrow width compared
with a comparatively massive headshield, coupled with the orientation of the thorax and pygidium, all
point to feeble musculature, with the appendages operating at a mechanical disadvantage. It is likely that
these trilobites were benthic and sluggish (Campbell 1975b). Many raphiophorids carry long median
cephalic spines as well. Eye reduction appears to have been typical of some clades.
In the examples just considered the suspension feeding habit may have evolved from the particulate
feeding one, for two reasons. First, the change from one to the other is a plausible progression which seems
to require no great shift in limb function. Second, where they are known, sister groups of suspension
feeders are natant, and hence likely to have been direct sediment feeders. Even within Trinucleina there are
some taxa which are considered to have had natant hypostomes (Hughes et al. 1975, p. 54), as did relatives
of Uromystrum and Cordania in the order Proetida. The sister goup of harpetids may have been
loganopeltids. which lack a pitted fringe but otherwise already show the vaulted cephalon typical of the
morphotype.
Functional aspects of trinucleimorph design

It is likely that the trinucleimorph design related directly to filter feeding habits: after all, a pitted fringe is
only developed in trilobites which on other criteria are considered to have used a cephalic feeding
chamber. Bergström (1969, 1972) and Cisne (1970) agreed that the morphology of the limbs of
Cryptolithus, studied originally by Raymond (1920), were consistent with ‘a trunk-limb filter feeding
mechanism’ (Cisne 1970, p. 21) such as that employed by many branchiopod and leptostracan crustaceans.
If it is assumed that the mode of operation of the chamber entailed selection of material from suspension
and passage forward to the mouth, then it is likely that water currents were brought into the chamber by
means of metachronal movements of those exites (‘gill branches’) which lie posterior to the mouth. It
makes good functional sense to have ‘clean’ water pass through the breathing apparatus. If this were the
ingress direction, the egress must be separated from it. One possibility is that an exit current moved
ventrally and posteriorly. But it is possible that hydrodynamic stability within the feeding chamber was
maintained by outflow through the pores perforating the fringe. This explanation has the attraction that it
produces a plausible pattern for both water and nutrient movement, as well as an explanation for the
presence of pitting on the flanks of the feeding chamber. Størmer (1930) showed that the pits would also
have a strengthening effect, but in these sluggish and poorly streamlined trilobites it is hard to see how that
alone would have been a high priority, nor would it account for the very particular form of trinucleid pits
(Whittington 1959), in which the central part of the perforation is exceedingly narrow. The explanation
advanced here is opposite to that given by Seilacher (1970) and Cisne (1970) in which the pores in the brim
are supposed to provide a filter to separate ‘finer particles from the sediment to be taken into the feeding
current’ (Cisne 1970, p. 22), that is, the movement is from outside inwards.
If the fringe had been an area of ingress, and the ventral median line was not an exhalent direction, then
the exhalent direction would perforce have had to be sideways between thorax and genal spines – which is
precisely where the respiratory exites are situated. The fine filaments of this appendage (Bergström 1969)
would have rapidly clogged with sediment – especially if Campbell’s (1975b) observation is right that the
length of these filaments had been underestimated by previous authors (they may have been more like
those of Triarthrus). If the logic of this argument is correct it adds plausibility to the opposite circulation
pattern shown in Text-figure 16, because it is the one which prevents fouling.
TEXT-FIG.16. Lateral view of the Ordovician trinucleid Cryptolithus tesselatus (Green) resting in a Rusophycus-type
burrow, showing the circulation pattern entering over the gills and exiting through the pits of the fringe, as postulated
here for a chamber filter feeder.
The feeding habit can be summarized thus: the animal settled on to a new patch of sediment and stirred it
into suspension, in the process bringing in oxygenating currents through the exites; edible particles were
sorted by the limbs within the feeding chamber and passed to the mouth along the food groove, flow of
material being maintained by egress of water through the pores of the fringe. In a short while, the trilobite
would exhaust a given site, move on to a new patch of sediment, and repeat the process. In those
trinucleimorphs lacking a pitted fringe, such as Ampyx, it is possible that current egress was out through the
anterior cephalic arch, which is commonly present in such forms, or perhaps more generally by ‘leakage’
along the broad cephalic margin.
This can scarcely be the whole story, because there are some trinucleimorphs, such as Protolloydolithus
(Whittard 1956 pl. 5, fig. 13), and even some species of Trinucleus itself, in which the major part of the
pitted fringe appears to have been horizontally disposed. In some harpetids there are pits in a concave
anterior part of the fringe, which seems more appropriate to rest on the sediment surface. Nonetheless it
does seem appropriate to attempt to explain the distinctive aspects of these trilobites in terms of feeding
habit, especially as the development of fringes seems to be confined to those trilobites whose morphology
is otherwise that of a filter feeder.
Trace fossils
Campbell (1975b) illustrated almost circular, bilaterally symmetrical trace fossils of Rusophycus
type associated with Cryptolithus. These show (Text fig. 17) that the animal made discrete traces in
one site before moving on to another, rather than ploughing the sediment. There is no coxal trace
TEXT-FIG.17. Rusophycus cryptolithi Osgood, a ‘resting’ burrow produced by a trinucleid trilobite. A, interpretative
drawing. Note the impressions left by the genal spines, and the irregular dark markings in the burrow, interpreted by
Campbell (1975b) as impressions made by the telopodites. He considered the impressions along the front wall of the
burrow to have been produced by the cephalic margin as the animal moved slightly backwards at successive depths
(after Campbell 1975b, fig. 2A and pl. 1, figs 1–3). B, single trace showing median ridge; University of Cincinnati
(UCM) Geology Museum 37587, × 3. C, several successive feeding burrows, produced by a single individual; UCM
37567; × 2. B–C courtesy of Professor K. S. W. Campbell.
on the mid-line, which is consistent with the elevated position of limb bases in the filter feeding
mode. There is a median ridge between the depressions to left and right produced by the activity of
the limbs. This is of interest because if there had been a posteroventral exhalent current it might be
expected to have left a median groove, or at the least to have scoured away any delicate median
ridge. We regard this as providing additional evidence of the exhalent function of the pitted
fringe.
The problem of illaenid morphology

There are some illaenids in which the hypostome also had the orientation described previously, that is,
steeply inclined upwards inside the cephalic cavity. Although such illaenids are large, it is tempting to add
the appropriate members of the family to the filter feeding category. However, we have come to doubt this
because of other aspects of their morphology, and their phylogenetic history. Although axial definition in
illaenids is often poor, the axis is generally wide, and the transverse vaulting is high. Furthermore, genal
spines are almost invariably lacking, so that the typical trinucleimorph design is not possible. Fortey
(1986) described the nileid Symphysurus as having a very illaenid-like morphology. In both illaenids and
Symphysurus the hypostome is strongly buttressed at the cephalic doublure, which carries very strong
terrace ridges.
Neither Illaenus nor Symphysurus have closely related sister taxa which are natant. Symphysurus is a
nileid close to Nileus, and the whole family Nileidae has strongly attached hypostomes showing features
characteristic of the predator/scavenger morphology discussed previously. If Fortey (1990) was correct in
allying illaenids and styginids then this clade also is typified by buttressed hypostomes. Hence the
transition to filter feeding mode seems, on balance, improbable.
Westrop (1983) and Fortey (1986) discussed the body orientation of these trilobites in life, and
concluded that they may have habitually lived partially buried, with the thorax and pygidium in the
sediment and the eyes in a horizontal attitude at the sediment surface (what Fortey termed the ‘bumastoid
stance’). In this position, the hypostome would have lain close to horizontal. This is not at all like the
stance deduced for typical trinucleimorphs. Whittington (1997) did not support the idea that illaenids lived
by partially burying themselves, but argued instead that they were ‘vagrant benthos particularly adapted
for negotiating over irregular surfaces’ based partly on the unusual flexibility of the thorax in a dorsal
direction. Whatever the correct interpretation of illaenimoprh mode of life we consider it unlikely that
these animals were filter feeders of Ampyx type, despite the similarity in the orientation of the hypostome.
Summary. Trilobites which habitually used a filter chamber for feeding had convex cephalic shields (often
swept back into genal spines) with convexity greatly exceeding that of thorax and pygidium, which were
habitually held above the sediment surface. In some forms the brim was perforated, probably as an aid to
current circulation. They were typically small- to medium-sized. These trilobites left a Rusophycus feeding
trace lacking impressions of the ‘coxal’ bases. Trilobites with this kind of morphology are present already
in the Cambrian, for example, among menomoniids and norwoodiids (see Paranorwoodia, Text-fig. 14).
Trinucleids, raphiophorids, harpetids and ityophorids were among Ordovician trilobites showing the
appropriate morphology, and harpetids and some proetoideans exemplify its continuation into the late
Palaeozoic. Although raphiophorids and trinucleids are probably most abundant in outer shelf biofacies, as
in the Ordovician of Wales, niches for filter chamber feeding seem to have been present in all normally
oxygenated biofacies, for example, harpetids can be associated with carbonate ‘mound’ facies (e.g. Lane
1972).
PELAGIC TRILOBITES
Pelagic trilobites typically have the following combination of morphological characters: large, or even
hypertrophied eyes; elongate exoskeleton with pleurae reduced (tr.); powerful thoracic articulation; genal
spines, where present, often directed downwards. Epipelagic trilobites are found in all biofacies, while
mesopelagic trilobites occur in those in proximity to a palaeo-ocean, sometimes in association with blind
TEXT-FIG.18. Pelagic trilobite morphology. A, reconstruction of the ventral side of the cephalon of Degamella evansi
Fortey and Owens showing the transverse hypostome. (Based on Fortey and Owens 1987, figs 38d and 40a–b).
B, Pricyclopyge binodosa (Salter); NHM I 15232; Ordovician, Llanvirn (S̆árka Formation), Bohemia; anterior
view; × 5.
(atheloptic) benthic trilobite assemblages (Fortey and Owens 1987). If these life habits are correctly
understood, pelagic trilobites presumably fed in the water column, and were not in direct competition with
the benthic feeding types considered previously. Even within the water column there may have been
vertical stratification,with different species having optimum feeding at different depths. McCormick and
Fortey (1998) have shown that the optical properties of Carolinites and Pricyclopyge differ in a way which
is consistent with the former being epipelagic, the latter mesopelagic.
It is unfortunate that we do not know the limbs of any non-agnostoid pelagic trilobite. It is unknown, for
example, whether the limbs were modified for swimming or for feeding in the water column. What follows
is inevitably speculative. However, there are some morphological indications on the exoskeleton that
feeding habits must have been modified in some way. Degamella has the most transverse hypostome of
any trilobite (Fortey and Owens 1987, fig. 38d; Text-fig. 18). This feature might become explicable if the
mouth had become comparably wide. Could this increased ‘gape’ have been an adaptation for direct
plankton feeding, in the same fashion as rays and some pelagic fish? Hypostomes of smaller cyclopygids such
as Microparia and Cyclopyge are also transverse, but less so than that of Degamella (Fortey and Owens 1987,
fig. 45b). They are all strongly supported at the doublure, but are never forked. The sister group of
Cyclopygidae is Nileidae, species of which seem to have had scavenger/predator morphologies, if the
arguments presented in this paper are followed. On balance, therefore, rather slim evidence favours the
majority of mesopelagic cyclopygids as actively feeding on zooplankton which also lived in the water column.
Fortey (1985) distinguished two morphological groups of pelagic trilobites (Text-fig. 19): small
(20–40 mm), poorly streamlined forms with bulbous eyes; and larger (up to 100 mm), well-streamlined
forms with extended cephalic shields (sag.) prolonged into ‘noses’. Degamella belongs in the latter
category, Cyclopyge, Telephinidae (Carolinites and allies) and Bohemilla in the former. The smaller and
less streamlined forms are presumably likely to have fed upon smaller and sluggish species of plankton.
We do not know the mode of hypostome attachment in Telephinidae, but it is considered that they are
related to Hystricurinae (Chatterton 1980) which included natant species, and the hypostome of
Carolinites (Fortey 1975b, pl. 38, fig. 2) is like that of typical ptychoparioids. It is therefore possible
that these telephinids fed upon plankton, directly collecting edible material in a way analogous to that of
TEXT-FIG. 19. Subdivision of the pelagic habitat exemplified by large, well-streamlined (A, Degamella, × 0·3) and
smaller, poorly streamlined (B, Cyclopyge, × 1·5; C, Telephina, × 1·5; D, Opipeuter × 2; and E, Bohemilla (Fenniops)
× 2) pelagic trilobites. After Fortey and Owens 1987 (A–B, E), and Fortey 1985 (C–D).
the particle feeders described above, to which they were related phylogenetically. Such a position as a
primary consumer low in the food chain is consistent with the great abundance that telephinids may attain
in favourable circumstances. For example, in one bed 75 m from the base of the Olenidsletta Member of
the Valhallfonna Formation (Ordovician, Arenig, Spitsbergen, see Fortey 1975b) cranidia, cheeks and
pygidia of Carolinites genacinaca genacinaca are often ‘rock formers’ (see Text-fig. 20). In that
Carolinites is found in both shallow water, inner shelf limestones (Ibex area of Utah, see Hintze 1953)
and deep water ‘slope’ graptolitic shales and limestones (Fortey 1975a; Henderson 1983) it seems that it
must have lived in surface waters, whence it spread entirely around the Ordovician palaeoequator.
Individual species of Carolinites were also very widespread, indeed, they probably ’grazed’ the surface
plankton of the whole Ordovician tropical belt.
The larger, streamlined morphologies appear to be confined to deeper water facies. The anomalous
Degamella was mentioned above, and may have been a plankton feeder. Parabarrandia was studied by
Fortey (1985) who showed that its profile was close to that of a hydrofoil, and streamlined by comparison
with a variety of other trilobites; it was designed to be an efficient swimmer. All such larger forms have
rigidly braced hypostomes. At this size, predatory/scavenging habits are more likely, by comparison with
living decapods and stomatopods. Possibly, these trilobites preyed upon the smaller, more sluggish
plankton-feeding species, with which they are rarer associates. However, phyllocarid crustacean carapaces
are common (although neglected) fossils in deeper water Ordovician shales and limestones, and these
would have comprised an abundant source of food.
Summary. Direct evidence of feeding habits in pelagic trilobites is not available, but morphological
evidence and field occurrence suggests that trilobites habitually living in the water column included well-
streamlined predators, and a number of smaller and poorly streamlined plankton feeders which can be
abundant. Different species probably inhabited different parts of the water column.
CHANGES IN FEEDING HABITS DURING ONTOGENY

Larval trilobites, especially protaspides, may also have lived as part of the zooplankton. Like other small
inhabitants of the pelagic realm they were likely to have fed upon phytoplankton and smaller
zooplankton – trochophore and vitellaria larvae, and the like. Speyer and Chatterton (1989) recognized
dorsally rather featureless, globular protaspides (termed asaphoid protaspides) as those most likely to have
been members of the zooplankton, and regarded more discoidal protaspides as being likely to have
manifested direct, progressive development moult-by-moult to a benthic adult, living close to the sea-
floor. Asaphoid protaspides are known from a variety of families of post-Cambrian age (Chatterton and
Speyer 1997), and the typical examples are members of the family Asaphidae. Since most metazoan phyla
primitively have a dispersive, planktic larval phase it is perhaps more likely a priori that planktic habits
TEXT-FIG. 20. Abundance of pelagic trilobites: the Ordovician epipelagic trilobite Carolinites genacinaca Ross,
Valhallfonna Formation (Arenig) Spitsbergen; Paleontological Museum Oslo NF 707; three cranidia include an
immature individual (centre); × 4.
were also primitively present in the trilobites, including those Cambrian forms which have discoidal
protaspides. Such protaspides typify the redlichiids and ptychopariids, for example, and may well have
been general in the early history of the group. The olenellids probably lacked a calcified protaspis
altogether. However, asaphoid protaspides are clearly advanced relative to the discoidal ones, with their
reduction of peripheral spines and overall inflation.
It is likely that there was a change in feeding habit between larva and adult. This is commonly the case in
living arthropods (Fryer 1988). Typical asaphoid protaspides provide some indication of what these
changes might involve. As demonstrated previously, adult asaphids are typical of what we regard as
predator/scavenging morphology – including very large species with braced, forked hypostomes with
well-attributed Rusophycus traces. The larva (Fortey and Chatterton 1988, fig. 10.2) has an attached
hypostome as in the adult, but the spinose hypostome is relatively huge so that it takes up the bulk of the
venter. The space available for the extrusion of appendages is very meagre – essentially small slots
between hypostomal spines – and it it does not seem possible that the animal would have been able to
manipulate large prey items; it must have fed on comparatively minute organic particles or organisms.
Hence at some point in the life cycle the trilobite settled to the sea floor and then or subsequently assumed
the quite different habits of the adult. There must have been something akin to a metamorphosis at that
stage. In fact, smoothed-out asaphoid larvae could grow quite large (those of some nileids are more than
3 mm long) and it is not implausible to assume that the rather sudden change to a meraspis similar to the
adult also coincided with the change to the same feeding habits as the adult.
For other non-asaphoid ontogenies there is less direct evidence of feeding in the protaspis, the
hypostomes of which are often relatively smaller. It is usual for the hypostome to be attached to the
doublure, which means that the kind of particle feeding employed by natant trilobites can be excluded.
Ontogenies of natant trilobites show the hypostome ‘detaching’ from the doublure during growth as the
preglabellar field extends (e.g. Olenus, see Clarkson and Taylor 1995). In most cases the trilobite is only a
few millimetres long when natant condition is attained. However, Clarkson et al. (1997) have shown that
the olenid Parabolina may only have become natant later in ontogeny. It can be reasonably asserted that
the adoption of the natant hypostome marked the beginning of particle-feeding during the life-cycle,
whatever the mode of feeding of the larva. Only in advanced Proetida are there protaspides which have
‘adult-like’ larvae (Scharyia, Chatterton and Speyer 1997), possibly with a natant proto-hypostome in some
cases, and these may have heterochronically ‘displaced’ particle feeding to an earlier stage in ontogeny. In
summary, for the majority of trilobites there is evidence that larval feeding strategy differed from that of the
adult. In some cases this was connected with settling of the immature trilobite to the sea-floor.
PROBLEMATICAL GROUPS
The general strategies for trilobite feeding described above do not pretend to account for the entire array of
TEXT-FIG. 21. A–B, holaspid and protaspid of Isotelus parvirugosus Chatterton and Ludvigsen (after Chatterton and
Ludvigsen 1976, pl. 2, figs 1, 10–12, 18 and Fortey and Chatterton 1988, fig. 10:1B respectively); C–D, meraspis and
holaspis of Olenus wahlenbergi Westergård (after Clarkson and Taylor 1995, figs 8h, l, 14 and 16a). Note the change in
morphology of the hypostome in Isotelus, and the change in its position with respect to the anterior border in Olenus.
Concomitant changes in feeding strategies presumably occurred during ontogeny. See text for discussion. Scale bars
represent 5 mm (A, C) and 0·5 mm (B, D).
exoskeletal designs in this very diverse group, although they may serve as a starting point for future work.
Below we consider a few examples of the kinds of morphology that resist interpretation, or are controversial.
Agnostoid trilobites
The mode of life of agnostoid (i.e. suborders Eodiscina and Agnostina) trilobites has been thoroughly
controversial (Robison 1972; Jago 1973; Pek 1977; Müller and Walossek 1987), and it would be remiss not
to mention the group in this review. Robison (1972, 1975) made a case for the planktic habits of Agnostina,
while Pek (1977) thought that they may have been epiplanktic, attached to floating seaweed, on the basis of
remarkable, aligned articulated specimens. Müller and Walossek (1987) considered that Agnostus
pisiformis lived in a nearly enrolled attitude, with the cephalic shield and pygidium employed almost
like the valves of an ostracod, whilst feeding on organic particles in the flocculent layer at the sediment
surface. Bergström (1973) thought that they might have been parasitic.
Shergold et al. (1990) regarded Agnostina as possibly independently derived from other trilobites, and
not closely related to Eodiscina; Fortey and Whittington (1989), Ramsköld and Edgecombe (1991), and
Fortey and Theron (1994) considered them to be derived trilobites related to Eodiscina within the clade
Agnostida. This argument about affinities surely reflects the remarkable specializations of the group, and is
TEXT-FIG. 22. An agnostoid coquina composed of an undetermined pseudagnostid from Bed 8 (lower Ordovician) of the
Cow Head Peninsula, western Newfoundland; slab showing packed individuals; NHM It16254; × 5. Detail shows that
some individuals are entire and enrolled.
relevant to their mode of life insofar as it relates to the feeding habits of close relatives. There seems to be a
consensus that eodiscoids were trilobites. Eodiscoids, at least the more advanced ones, had a natant
hypostome (Jell 1975; Fortey 1990). By comparison with the trilobites discussed earlier, this suggests that
they were benthic feeders on minute organic particles, including algae, detritus and animalcules. If the
more specialized agnostids, which also had a natant hypostome (Robison 1972), were closely related to the
eodiscoids then it is likely that they, too, had similar habits, at least primitively. The origin of Agnostida
was likely to have been paedomorphic, by progenesis (Briggs and Fortey 1992; Fortey and Theron 1994).
It seems possible that precocious sexual maturity in Eodiscina and Agnostina facilitated feeding on small
particle sizes in cryptic habitats. They may then have occupied the size and feeding niches below that of
ptychoparioids with natant hypostomes. Once achieved, the small-sized developmental strategy of
agnostids could evidently not be reversed; they stayed small.
Bergström’s suggestion about parasitic habits is intriguing. It was based upon the resemblance of highly
caecate agnostids to living parasitic copepods with a comparably well-developed system of digestive
glands. The host was not suggested. Some caecate (scrobiculate) agnostids are extremely abundant and
widespread globally (e.g. Glyptagnostus reticulatus, see Kobayashi 1949). It does seem improbable that
such abundant animals were parasites, especially in view of the absence of any suitable host, which itself
presumably would have had to have been large and abundant. The largest Cambrian animals we know are
the giant predators, anomalocarids, and none of these is associated with agnostids on the body. The
problem is even more marked for such non-scrobiculate species as the late Cambrian Agnostus pisiformis,
which can be so abundant as to be a rock former! The Agnostus pisiformis limestones are widespread
in southern Sweden and are virtually composed of this one species. RAF has estimated that a 10 mm3 of
TEXT-FIG. 23. Differing views of agnostoid life habits. A, Agnostus pisiformis (Wahlenberg) in possible life attitude for
swimming and feeding (after Müller and Walossek 1987, fig. 25B); B, aligned, enrolled specimens of Geragnostella
tullbergi (Novák), possibly attached to an algal thallus (from Pek 1977, pl. 10, fig. 1); C, Corrugatagnostus morea
(Salter), with ‘alimentary apparatus’ (after Öpik 1961, text-figs 5–6). Scale bars represent 0·5 mm (A) and 5 mm (B–C).
this rock holds about 125 specimens, divided equally between headshields and pygidia, assuming an
average length of about 2 mm for each. This is based on measurements taken from a block, It 20108, in The
Natural History Museum collections. Even if this thin limestone were calculated as only 10 mm thick (it is
invariably thicker) it extended across most of southern Sweden and Bornholm and into the Olso region.
One could very conservatively estimate something like 1014 agnostoids within it. Considering the sparse
body-fossil record of parasites as a whole (Boucot 1990) this superabundance does render parasitic habits
most improbable.
The morphology of species belonging to Agnostida is not like that of polymeroid pelagic trilobites, not
only because they are blind, but also because they are isopygous and with a reduced thorax. This does not
necessarily rule out pelagic or planktic habits, although it does indicate adaptations of a fundamentally
different kind. It is certainly the case that abundance of agnostids is generally associated with more
exterior, off-shelf biofacies, or with the transgression of such facies on to the shelf (Palmer 1984).
However, Jago (1973) and Babcock (1994) have noticed a relationship between bottom community
(biofacies) and the occurrence of particular agnostid genera, which is suggestive of benthic habits, as
Müller and Walossek (1987) inferred from a consideration of the limb morphology.
Muller and Walossek’s (1987) suggestion that agnostids fed with the pygidium and cephalon habitually
nearly closed, ostracod-like, is worth discussion. If this were the attitude one would expect them to be
preserved more commonly in the enrolled condition than is the case. The rich Cambrian agnostid faunas
from different continents described by Westergård (1946), Öpik (1979) and Robison (1964, 1984) all
feature a majority of entire specimens in extended position. As an example, a slab of Wheeler Shale
(Middle Cambrian) figured by Gunther and Gunther (1981) shows 17 or 18 entire specimens, none
enrolled. RAF examined similar slabs in the US National Museum. It might be argued that enrolled
specimens might be neglected as less collectable and desirable. The authors made a special attempt to
collect all specimens of Litagnostus meniscus from the Tremadoc Shineton Shale (Fortey and Owens
1991) but found only two enrolled examples among 20 or so entire specimens. Certainly, it seems difficult
to sustain the view that ‘outstretching was impossible’ (Müller and Walossek, 1987, p. 53), even though
careful morphological arguments are adduced to sustain this claim. The only way one could support Müller
and Walossek’s orientation in the face of such examples would be if the cephalon and pygidium were
continuously closed by active muscular contraction during life, which then became quickly released on
death, causing the animal to ‘spring apart’ into an extended position. Clams that do this have the resilium
on the outside of the hinge line, which is scarcely possible in an agnostid. Could the cephalothoracic
aperture have included a resilium-like membrane, which could act as the ‘spring’ to open the valves upon
death? Müller and Walossek (1987, pl. 28, figs 7–9) figured a preserved arthrodial membrane in this
position in two enrolled specimens, but there was no particular reason to suppose that it was under tension.
Although not commonly figured, enrolled specimens of A. pisiformis are known from rock occurrences as
well as örsten (see also Text-fig. 22), but when these occur they are completely enrolled (and they are
obviously adapted for tight enrolment) rather than preserved in their alleged life position with ‘valves’
agape. Hence the evidence from field occurrences does seem to argue against Muller and Walossek’s life
orientation.
The best-preserved, communally enrolled specimens yet discovered were figured by Pek (1977) from
the Ordovician of Bohemia (Text. fig. 23B). These specimens were more or less aligned and Pek (1977, p.
34) concluded that: ‘some agnostids could become adapted to an epifaunal mode of life on free-floating
thalli of algal agglomerations or live as phytals in algal growth of the sublittoral zone’. This adaptation was
likely to have been secondary, and developed after a putatively planktic larval phase. The evidence
presented by these specimens is compelling, and it is plausible that some, at least, later agnostids adopted
this life habit, which is common among living ostracods.
In sum, the evidence remains somewhat equivocal. It seems probable that agnostoids in general fed on
small organic particles of a size range below that of other natant trilobites. It is unlikely that they were
parasitic, and there is some evidence that they did not habitually hold the pygidium and cephalon in a
‘closed’ orientation. Various species may well have been planktic, but facies dependence also indicates
that some forms had benthic habits, but usually in relatively ‘oceanic’ settings. Some Ordovician species
were probably epipelagic. A range of life habits best accounts for the taxonomic variety that these
diminutive arthropods may attain, even within a single horizon, such as the faunas from the Middle
Cambrian of Australia illustrated by Öpik (1979).
Cheiruroidea
We have difficulty in applying the feeding model to Cheiruroidea. Trilobites belonging to the family
Cheiruridae seem to have to the hypostome firmly attached in the conterminant mode, but the orientation is
different from other trilobites of this kind. The hypostome slopes downwards, often at quite a steep angle
below the glabella; for example, Deiphon (Lane 1971, pl. 12, fig. 6b), and Cydonocephalus (Whittington
1963, pl. 27, fig. 7). Pliomeridae are similar (Whittington 1988b). The hypostomes of this group are very
conservative through a long stratigraphical range (Ordovician–Devonian). They invariably have a surface
sculpture comprising granules or small tubercles. Nor are forks developed, nor any other of the specialized
structures which may develop posteromedially on hypostomes of species that we have considered as
predators above. It seems likely that they had a different feeding mode in detail from any considered
previously, but what it was is puzzling. Presumably, the downward directed hypostome allowed for an
even greater expansion of the stomach cavity, and the frontal glabellar lobe of these trilobites is also often
appropriately inflated. The lack of terrace ridges on the venter might be taken as evidence that the doublure
did not interact directly with the sediment, i.e. that these trilobites were active surface feeders. On the other
hand, pliomerids such as Pseudocybele have long, vaulted bodies and stalked eyes, characters that in
homalonotids are associated with burrowing habits (Hammann 1985), and which Ingham (1968) suggested
for the cybeline Paracybeloides. The strong development of glabellar furrow S1 (often hooked backwards)
implies a special function for the appropriate limb pair whose musculature was associated with this region.
Taking this slim evidence together it is possible to speculate that these trilobites were specialized predators
that engulfed prey in an enlarged stomach, processing the food there (gastric mill?) rather than using the
bases of the limbs to macerate and sort material, as was argued for species with specialized hypostomes.
The S1 limbs may have been adapted for a grasping function; in Sphaerexochus this pair of furrows is
developed at the expense of those lying anteriorly, and it is conceivable that more anterior pairs were
comparatively atrophied.
CONCLUSION: TOWARDS EXPLAINING TRILOBITE FAUNAL DIVERSITY

The identification of trilobite feeding habits is of much more than anecdotal interest. An understanding of
feeding history is important in sketching the history of ecosystem utilization through time, and is the basis
of several palaeoecological generalizations. For example, Bambach’s (1983) classic study of changes in
ecospace occupation during the Palaeozoic classified trilobites into a few guilds related to broad
morphological divisions (his table 2). Two of these – ‘blind, short thorax, often pitted fringe’ and
‘agnostid form’ – correspond to feeding types recognized by us. Three others – ‘asymmetry of cephalon to
pygidium’, ‘symmetry of cephalon to pygidium’ and ‘elaborately spinose’ do not bear any relation to the
feeding habits we have identified. Predator/scavenger morphology may occur in trilobites with small
pygidia (Cambrian Paradoxides) or large ones (Ordovician Asaphidae). There is a general trend in trilobite
evolution from the former morphology to the latter (Fortey and Owens 1997) which we do not regard as
related to feeding habits per se. Very spinose morphology occurs in trilobites with both natant (e.g.
Cambrian Pardailhania hispida, see Courtessole 1973) and conterminant (e.g. Devonian Phacopida in
Kowalski 1992) hypostomal attachment. We assert that ecospace utilization as applied to extinct
organisms should be determined through a careful appraisal of their functional morphology before they
are incorporated into large-scale analyses.
If we are correct in identifying predatory/scavenging habits as primitive in trilobites, and typical of the
Lower Cambrian, then this reinforces the conclusion reached from the study of ‘soft bodied’ arthropods
that they included the major predators at this early stage in the radiation of the Metazoa. Chen et al. (1997)
claimed that at least one species of the trilobite sister-taxon Naraoia was a deposit feeder, but the great
majority of early arachnomorphs had predator morphology. Natant, particle- or deposit-feeding ptycho-
parioids had appeared before the end of the Early Cambrian, and were abundant by the Mid Cambrian (e.g.
Zhang et al. 1980). There are plausible filter-chamber feeders by the Late Cambrian (norwoodiids). There
were already trilobites (Centropleura) with possible pelagic morphology by the Mid Cambrian – although
such life habits are not known from examples younger than Ordovician (Fortey and Owens 1990);
agnostoids are known from the Lower Cambrian to uppermost Ordovician. Hence, even within the
Cambrian, there is the full spectrum of feeding possibilites identified in this review, and this may account
for some of the within-habitat diversity (alpha diversity of Sepkoski 1988) exhibited by trilobite faunas
collected from a single locality. With the exception of pelagic feeding, all three major feeding modes can
be identified from the morphology of trilobites which range in age from the Cambrian to the Permian.
Naturally, there is no obligation for a given palaeoenvironment to provide a living for all main feeding
modes: we have already mentioned the Cambro-Ordovician Olenid Biofacies, which is dominated by
sediment feeders, and there are plenty of Ordovician faunas which lack pelagic trilobites. The presence or
absence of a given ecological group depends on extrinsic, physical factors. Nonetheless, feeding mode
provides a way of assessing the morphological spread of common, co-occurring elements in a given fauna.
Different trilobites having the same feeding mode may have co-existed if they divided the available
resources in such a way as to avoid competitive exclusion. For example, predators may have optimized on
different sizes of prey species, with the larger predators taking the larger prey; eagles, buzzards and hawks
are a familiar Recent example of this resource partitioning. The concept of the ecological niche
(Hutchinson 1957) as an optimization of all the ecological factors which might contribute to the fitness
of an organism has been greatly criticized as almost impossible to define among living organisms
(e.g. Schoener 1988) let alone fossils. However, it is reasonable to suppose that an explanation for the
TEXT-FIG. 24. An attempt to sort a co-occurring assemblage of trilobites according to habitat and proposed feeding
strategy (guild). Biota of the late Arenig Pontyfenni Formation of Pont-y-fenni quarry, Whitland, South Wales. This
was a deep water habitat occupied by an atheloptic, benthic assemblage of trilobites, above which swam a series of
probably mesopelagic species. Both groups were probably differentiated by size and feeding strategy.
coexistence of different species with similar feeding strategies is that they partitioned food resources
among themselves, even if there are factors (such as prey specialization and breeding strategies, see Brown
1995) which will probably forever elude us in palaeontology. It is also necessary to remember that the
non-preservable soft-bodied fauna may have included successful competitors as well as prey.
Overall body size may be correlated with the sizes of food particle ingested preferentially. Robison
(1975) and Fortey and Rushton (1980) have invoked this to explain the co-existence of closely-related
species of agnostid and shumardiid trilobites, respectively. Hence it will be instructive to look at body size
in faunas as a whole as a surrogate for feeding specialization. Since death assemblages preserved in the
fossil record are nearly always time-averaged on a scale beyond the customary compass of ecologists,
measurements taken from fossils cannot be claimed to represent living communities, but careful collecting
can at least retrieve an inventory of morphologies present in a single and uniform former habitat.
In the lower Ordovician of South Wales the Pontyfenni Formation yields a deep-water biota including
articulated benthic trilobites, and a range of pelagic trilobites occurring in the same beds (Fortey and
Owens 1987, fig. 13). This thick formation represents a single type of biofacies or palaeocommunity
characterized by its blind (or almost blind) benthic trilobites, and the fauna (Text-fig. 24) from one quarry
at the type locality, Pont-y-fenni, near Whitland, is plausible as a surrogate for a short Ordovician time
period in one place. The authors have collected virtually every specimen recovered from this locality.
Many of the species are rare, and their rôle in the community is uncertain. There are no particle feeders
apart from a tiny species of Shumardia. There are three comparatively common species with predator/
scavenger morphology: Illaenopsis harrisoni, Ormathops nicholsoni and Dindymene saron. Illaenopsis is
large, 53–60 mm long (fragments indicate that the species grew to at least 130 mm long); Ormathops
nicholsoni is medium-sized, about 13–35 mm long; Dindymene saron is small, 5–7 mm long. Of species
with filter chamber morphology, Bergamia rushtoni and Ampyx linleyoides are common; A. linleyoides
(excluding frontal spine) is 8–22 mm long (mean 16 mm) B. rushtoni (with full complement of arcs) is
6–12 mm long. Populations are different at more than the 95 per cent. confidence level. Thus
articulated material from this locality provides some support for the idea that alpha diversity may be
maintained by resource partitioning within a given feeding mode. The next commonest predator/scavenger
is Colpocoryphe taylorum which, at 22 mm long, is in the middle of the range of Ormathops nicholsoni;
however, Colpocoryphinae to which it belongs have been claimed as burrowers (Hammann 1983), and if
so it may not have been in competition with Ormathops. Among pelagic forms, four species are
comparatively common: Degamella evansi, Pricyclopyge binodosa eurycephala, Microparia broeggeri,
and Cyclopyge grandis brevirhachis. Degamella evansi is a well-streamlined ’bullet head’ form with
complete individuals 50–60 mm long; Microparia broeggeri has a similar exoskeletal form but is
12–25 mm long. They thus were able to co-exist while competing for prey within their particular pelagic
habitat. Pricyclopyge binodosa eurycephala is the poorly streamlined type with more globular eyes; it has
a similar size range to Microparia, 12–30 mm long. The other poorly streamlined species, Cyclopyge
grandis, is 9–15 mm long. These, too, were probably able to divide the resources of their habitat in a co-
operative way. Thus, the co-existence of the ten or 11 (including Shumardia) commonest trilobites in a
single fauna can be explained in terms of (1) their feeding deployments (2) their resource partitioning
within a given habitat. This goes some way to answering one of the most frequently asked, but intractable
questions posed by students encountering trilobites in the field for the first time: how did so many trilobites
co-exist? The future lies in quantitative studies of whole faunas to see if such an explanation is of general
application.
Acknowledgements. For comments and discussion on various parts of this paper we are indebted to Nigel Hughes,
Geoff Boxshall and Brian Rosen, although none of them necessarily shares the views of the authors. Lin Norton
produced the line diagrams, and Phil Crabbe was responsible for photography. Ken Campbell and Gian Luigi Pillola
kindly supplied the originals of Text-figures 17B–C and 6A, respectively.
REFERENCES
BABCOCK, L. E. 1994. Biogeography and biofacies patterns of Middle Cambrian polymeroid trilobites from North
Greenland. Bulletin Grønlands Geologiske Undersøgelse, 169, 129–148.
BAMBACH, R. K. 1983. Ecospace utilization and guilds in marine communities through the Phanerozoic. 719–746. In
TEVESZ, M. J. S. and MCCALL, P. L. (eds). Biotic interactions in Recent and fossil benthic communities. Plenum, New
York, 1045 pp.
BERGSTRÖM, J. 1969. Remarks on the appendages of trilobites. Lethaia, 2, 395–414.
—— 1972. Appendage morphology of the trilobite Cryptolithus and its implications. Lethaia, 5, 85–94.
—— 1973. Organisation, life and systematics of trilobites. Fossils and Strata, 2, 1–69.
BOUCOT, A. J. 1990. Evolutionary paleobiology of behavior and coevolution. Elsevier, Amsterdam, xxiii þ 723 pp.
BRIGGS, D. E. G. and FORTEY, R. A. 1989. The early radiation of the Arthropoda. Science, N.Y., 246, 241–243
—— —— 1992. The early Cambrian radiation of arthropods. 335–373. In LIPPS, J. H. and SIGNOR, P. W. (eds). Origin
and early evolution of the Metazoa. Plenum, New York, 570 pp.
BROWN, J. H. 1995. Macroecology. University of Chicago Press, 269 pp.
BRUSCA, R. C. and BRUSCA, G. J. 1990. Invertebrates. Sinauer Associates, Sunderland, Mass., 922 pp.
BRUTON, D. L. 1968. The trilobite genus Panderia from the Ordovician of Scandinavia and the Baltic areas. Norsk
Geologisk Tiddskrift, 48, 1–34.
CAMPBELL, K. S. W. 1975a. The functional anatomy of phacopid trilobites: musculature and eyes. Journal and
Proceedings, Royal Society of New South Wales, 108, 168–188.
—— 1975b. The functional morphology of Cryptolithus. Fossils and Strata, 4, 65–86.
—— 1977. Trilobites of the Haragan, Bois d’Arc and Frisco formations (Early Devonian), Arbuckle Mountains
region, Oklahoma. Bulletin of the Oklahoma Geological Survey, 123, 1–139.
CHATTERTON, B. D. E. 1980. Ontogenetic studies of the Middle Ordovician trilobites of the Esbataottine Formation,
Mackenzie Mountians, Canada. Palaeontographica, Abteilung A, 171, 1–74.
—— and LUDVIGSEN, R. 1976. Silicified Middle Ordovician trilobites from the South Nahanni River area, District of
Mackenzie, Canada. Palaeontographica, Abteilung A, 154, 1–106, pls 1–22.
—— ——— 1998. Upper Steptoan (Upper Cambrian) trilobites from the McKay Group of southeastern British
Columbia, Canada. Memoir of the Paleontological Society, 49, 1–43.
—— and PERRY, D. 1983. Silicified Silurian odontopleurid trilobites from the Mackenzie Mountains. Palaeonto-
graphica Canadiana, 1, 1–55.
—— and SPEYER, S. E. 1997. Ontogeny. 173–247. In WHITTINGTON, H. B., CHATTERTON, B. D. E., SPEYER, S. E., FORTEY, R. A.,
OWENS, R. M., CHANG, W. T., DEAN, W. T., JELL, P. A., LAURIE, J. R., PALMER, A. R., REPINA, L. A., RUSHTON, A. W. A., SHERGOLD,
J. H., CLARKSON, E. N. K., WILMOT, N. V. and KELLY, S. R. A.. Treatise on invertebrate paleontology Part O. Arthropoda 1.
(revised). Geological Society of America and University of Kansas Press, Lawrence, xxiv þ 530 pp.
CHEN, J.-Y., EDGECOMBE, G. D. and RAMSKÖLD, L. 1997. Morphological and ecological disparity in naraoiids (Arthropoda)
from the early Cambrian Chengjiang fauna, China. Records of the Australian Museum, 49, 1–24.
CISNE, J. L. 1970. Beecher’s trilobite bed revisited: ecology of an Ordovician deep water fauna. Postilla, 160, 1–25.
—— 1975. Anatomy of Triarthrus and the relationships of the Trilobita. Fossils and Strata, 4, 45–64.
CLARKSON, E. N. K. and LEVI-SETTI, R. 1975. Trilobite eyes and the optics of Des Cartes and Huygens. Nature, 254 (5502),
663–667.
—— and TAYLOR, C. E. 1995. Ontogeny of the trilobite Olenus wahlenbergi Westergård 1922 from the Upper Cambrian
Alum Shales of Andrarum, Skåne, Sweden. Transactions of the Royal Society of Edinburgh: Earth Sciences, 86, 13–34.
—— —— and AHLBERG, P. 1997. Ontogeny of the trilobite Parabolina spinulosa (Wahlenberg, 1818) from the Upper
Cambrian Alum Shales of Sweden. Transactions of the Royal Society of Edinburgh: Earth Sciences, 88, 69–89.
CODDINGTON, J. A. 1988. Cladistic tests of adaptational hypotheses. Cladistics, 4, 3–22.
COURTESSOLE, R. 1973. Le Cambrien moyen de la Montagne Noire. Université de Toulouse, 241 pp., 27 pls.
DRAPER, J. J. 1980. Rusophycus (Early Ordovician ichnofossil) from the Mithaka Formation, Georgina Basin. Bureau of
Mineral Resources Journal of Australian Geology and Geophysics, 5, 57–61.
DROSER, M. L., FORTEY, R. A. and XING, L. 1996. The Ordovician radiation. American Scientist, 84, 122–131 (and cover).
ELDREDGE, N. 1971. Patterns of cephalic musculature in the Phacopina (Trilobita) and their phylogenetic significance.
Journal of Paleontology, 45, 52–67.
—— 1980. An extravagance of species. Natural History, 89, 45–51.
FOOTE, M. 1991. Morphologic patterns of diversification: examples from trilobites. Palaeontology, 34, 461–485.
FORTEY, R. A. 1974. The Ordovician trilobites of Spitsbergen. I Olenidae. Skrifter Norsk Polarinstitutt, 160, 1–129.
—— 1975a. Early Ordovician trilobite communities. Fossils and Strata, 4, 331–352.
—— 1975b. The Ordovician trilobites of Spitsbergen. II. Asaphidae, Nileidae, Raphiophoridae and Telephinidae of
the Valhallfonna Formation. Skrifter Norsk Polarinstitutt, 162, 1–207.
—— 1985. Pelagic trilobites as an example of deducing the life habits of extinct arthropods. Transactions of the Royal
Society of Edinburgh: Earth Sciences, 76, 219–230.
—— 1986. The type species of the Ordovician trilobite Symphysurus: systematics, functional morphology and terrace
ridges. Paläontologische Zeitschrift, 60, 255–275.
—— 1990. Ontogeny, hypostome attachment and trilobite classification. Palaeontology, 33, 529–576.
—— and CHATTERTON, B. D. E. 1988. Classification of the trilobite suborder Asaphina. Palaeontology, 31, 165–222.
—— and OWENS, R. M. 1978. Early Ordovician (Arenig) stratigraphy and faunas of the Carmarthen District, south-west
Wales. Bulletin of the British Museum (Natural History), Geology Series, 30, 225–294.
—— —— 1987. The Arenig Series in South Wales. Bulletin of the British Museum (Natural History), Geology Series,
41, 69–307.
—— ——1990. Trilobites. 121–142. In MCNAMARA, K. J. (ed.). Evolutionary trends. Belhaven Press, London, 368 pp.
—— —— 1991. A trilobite fauna from the highest Shineton Shales of Shropshire, and the correlation of the latest
Tremadoc. Geological Magazine, 128, 437–464.
—— —— 1997. Evolutionary history. 249–287. In KAESLER, R. L. . (ed.). Treatise on invertebrate paleontology. Part
O. Arthropoda 1. Geological Society of America and University of Kansas Press, Lawrence, xxiv þ 530 pp.
—— and RUSHTON, A. W. A. 1980. Acanthopleurella Groom, 1902: origin and life-habits of a miniature trilobite.
Bulletin of the British Museum (Natural History), Geology Series, 33, 79–89.
——and SEILACHER, A. 1997. The trace fossil Cruziana semiplicata and the trilobite that made it. Lethaia, 30, 105–112.
—— and THERON, J. N. 1994. A new Ordovician arthropod, Soomaspis, and the agnostid problem. Palaeontology, 37,
841–861.
—— and WHITTINGTON, H. B. 1989. The Trilobita as a natural group. Historical Biology, 2, 125–138.
—— and WILMOT, N. V. 1991. Trilobite cuticle thickness in relation to palaeoenvironment. Paläontologische
Zeitschrift, 65, 141–151.
FRYER, G. 1968. Evolution and adaptive radiation in the Chydoridae (Crustacea: Cladocera): a study in comparative
functional morphology and ecology. Philosophical Transactions of the Royal Society of London, Series B, 254,
221–385.
—— 1988. Studies on the functional morphology and biology of the Notostraca (Crustacea: Branchiopoda).
Philosophical Transactions of the Royal Society of London, Series B, 321, 27–124.
GUNTHER, L. F. and GUNTHER, V. G. 1981. Some Middle Cambrian fossils of Utah. Brigham Young University Geology
Studies, 28, 1–87.
HAMMANN, W. 1983. Calymenacea (Trilobita) aus dem Ordovizium von Spanien: ihre Biostratigraphie, Okologie und
Systematik. Abhandlungen von der Senckenbergischen Naturforschenden Gesellschaft, 542, 1–177.
—— 1985 . Life habit and enrolment in Calymenacea (Trilobita) and their significance for classification. Transactions
of the Royal Society of Edinburgh: Earth Sciences, 76, 307–318.
HENDERSON, R. A. 1983. Early Ordovician faunas from the Mount Windsor subprovince, northeastern Queensland.
Memoirs of the Association of Australasian Palaeontologists, 1, 145–175.
HENNINGSMOEN, G. 1957. The trilobite family Olenidae. Skrifter utgitt av det Norske Videnskaps-Akademi I Oslo. 1.
Mathematics and Natural Sciences, 1, 1–303, pls 1–31.
HINTZE, L. F. 1953. Lower Ordovician trilobites from western Utah and eastern Nevada. Bulletin of the Utah Geological
and Mineralogical Survey, 48, 1–249.
HUGHES, C. P., INGHAM, J. K. and ADDISON, R. 1975. The morphology, evolution and classification of the Trinucleidae
(Trilobita). Philosophical Transactions of the Royal Society of London, Series B, 272, 537–607.
HUGHES, N. C. and CHAPMAN, R. E. 1995. Growth and variation in the Silurian proetide trilobite Aulacopleura konincki
and its implication for trilobite palaeobiology. Lethaia, 28, 333–354.
HUTCHINSON, G. E. 1957. Concluding remarks. Cold Spring Harbor Symposia on Quantitative Biology, 22, 415–27.
INGHAM, J. K. 1968. British and Swedish Ordovician species of Cybeloides (Trilobita). Scottish Journal of Geology, 4,
300–316.
JAGO, J. B. 1973. Cambrian agnostid communities in Tasmania. Lethaia, 6, 405–21.
JELL, P. A. 1975. Australian Middle Cambrian eodiscoids with a review of the superfamily. Palaeontographica,
Abteilung A, 150, 1–97.
—— 1977. Penarosa netenta, a new Middle Cambrian trilobite from northwestern Queensland. Memoirs of the
Queensland Museum, 18, 199–123.
—— 1978. Trilobite respiration and genal caeca. Alcheringa, 2, 251–260.
JENSEN, S. 1990. Predation by early Cambrian trilobites on infaunal worms – evidence from the Swedish Mickwitzia
Sandstone. Lethaia, 23, 29–42.
KOBAYASHI, T. 1949. The Glyptagnostus hemera, the oldest world-instant. Japanese Journal of Geology and
Geography, 21, 1–6.
KOWALSKI, H. 1992. Trilobiten Verwandlungskünstler des Paläozoikums. Goldschneck-Verlag, Korb, 157 pp.
LANE, P. D. 1971. British Cheiruridae (Trilobita). Monograph of the Palaeontographical Society, 125 (530), 1–95, pls
1–16.
—— 1972. New trilobites from the Silurian of north-east Greenland, with a note on trilobite faunas in pure limestones.
Palaeontology, 15, 336–364.
LAVAL. P. 1980. Hyperiid amphipods as crustacean parasitoids associated with gelatinous zooplankton. Annual Review
of Oceanography and Marine Biology, 18, 11–56.
LUDVIGSEN, R. and CHATTERTON, B. D. E. 1991. The peculiar Ordovician trilobite Hypodicranotus from the Whittaker
Formation, District of Mackenzie. Canadian Journal of Earth Sciences, 28, 616–622.
—— WESTROP, S. R. and KINDLE, C. H. 1989. Sunwaptan (Upper Cambrian) trilobites of the Cow Head Group, western
Newfoundland, Canada. Palaeontographica Canadiana, 6, 1–175.
MCCORMICK, T. and FORTEY, R. A. 1998. Independent testing of a paleobiological hypothesis: the optical design of two
Ordovician pelagic trilobites reveals their relative paleobathymetry. Paleobiology, 24, 235–253.
MILLER, J. 1976. The sensory fields and life mode of Phacops rana (Green, 1832). Transactions of the Royal Society of
Edinburgh, 69, 337–67.
MOORE, R. C. (ed.) 1959. Treatise on invertebrate paleontology. Part O. Arthropoda 1. Geological Society of America
and University of Kansas, Boulder, Colorado, and Lawrence, Kansas, xix þ 560 pp.
MÜLLER, K. J. and WALOSSEK, D. 1987. Morphology, ontogeny and life habit of Agnostus pisiformis from the Upper
Cambrian of Sweden. Fossils and Strata, 19, 1–123.
ÖPIK, A. A. 1961. Alimentary caeca of agnostids and other trilobites. Palaeontology, 3, 410–438.
—— 1967. The Mindyallan fauna of northwestern Queensland. Bureau of Mineral Resources, Geology and
Geophysics, Australia, 74, 1–404, pls 1–51 [2 vols].
—— 1979. Middle Cambrian agnostoids: systematics and biostratigraphy. Bulletin of the Bureau of Mineral
Resources, Geology and Geophysics, Australia, 172, 1–188, pls 1–67.
ORMISTON, A. R. 1971. Silicified specimens of the Gedinnian trilobite, Warburgella rugulosa canadensis Ormiston,
from the Northwest Territories, Canada. Paläontologische Zeitschrift, 45, 173–180, pls 19–21.
OWENS, R. M. 1979. The trilobite genera Panarchaeogonus Öpik, Isbergia Warburg and Cyamops gen. nov. from the
Ordovician of Balto-Scandia and the British Isles. Norsk Geologisk Tidsskrift, 58, 199–219.
—— 1983. A review of Permian trilobite genera. Special Papers in Palaeontology, 30, 15–42.
—— 1990. Carboniferous trilobites: the beginning of the end. Geology Today, 6, 96–100.
PALMER, A. R. 1984. The biomere problem: evolution of an idea. Journal of Paleontology, 58, 599–611.
PEK, I. 1977. Agnostid trilobites of the central Bohemian Ordovician. Sbornik Geologickych Ved Praha, 1977 (19),
7–44.
RAMSKÖLD, L. and EDGECOMBE, G. D. 1991. Trilobite monophyly revisited. Historical Biology, 4, 267–283.
—— —— 1996. Trilobite appendage structure of Eoredlichia reconsidered. Alcheringa, 20, 269–276.
RASETTI, F. 1945. New Upper Cambrian trilobites from the Lévis Conglomerate. Journal of Paleontology, 19, 462–478,
pls 60–62.
RAYMOND, P. E. 1920. The appendages, anatomy and relationships of trilobites. Memoirs of the Connecticut Academy of
Arts and Sciences, 7, 1–169.
ROBISON, R. A. 1964. Late Middle Cambrian trilobites from western Utah. Journal of Paleontology, 38, 510–566.
—— 1972. Mode of life of agnostid trilobites. 24th International Geological Congress, Montreal, Session 7, 33–40.
—— 1975. Species diversity among agnostoid trilobites. Fossils and Strata, 4, 219–226.
—— 1984. Cambrian agnostids of North America and Greenland. Part 1. Ptychagnostidae. Paleontological
Contributions of the University of Kansas, 109, 1–59.
RUDKIN, D. and TRIPP, R. P. 1989. The type species of the Ordovician trilobite genus Isotelus: I. gigas Dekay, 1824. Royal
Ontario Museum Life Sciences Contributions, 152, 1–19.
SANDERS, H. L. 1963. The Cephalocarida: functional morphology, larval development, comparative external anatomy.
Memoirs of the Connecticut Academy of Arts and Sciences, 15, 1–80.
SCHOENER, T. W. 1988. The ecological niche. 79–113. In CHERRETT, J. M. (ed.). Ecological concepts. Blackwell
Scientific, Oxford, 503 pp.
SCHRAM, F. R. 1986. Crustacea. Oxford University Press, Oxford, 606 pp.
SEILACHER, A. 1970. Cruziana stratigraphy of ‘non-fossiliferous’ Palaeozoic sandstones. 447–476, In CRIMES, T. P. and
HARPER, J. C. (eds). Trace fossils. Special Issue of the Geological Journal, 3. Seal House Press, Liverpool, 547 pp.
SEPKOSKI, J. J. JR 1988. Alpha, beta, or gamma: where does all the diversity go? Paleobiology, 14, 221–234.
SHAW, F. C. 1968. Middle Ordovician Chazy trilobites of New York. Memoir of the New York State Museum of Natural
History, 17, 1–114.
SHERGOLD, J. H., JAURIE, J. R. and SUN XIAO-WEN 1990. Classification and review of the trilobite order Agnostida. Report
of the Bureau of Mineral Resources, Geology and Geophysics, Australia, 296, 1–70.
SHU, D. G., GEYER, G., CHEN, L. and ZHANG X. L. 1995. Redlichiacean trilobites with soft-parts from the Lower Cambrian
Chengjiang Fauna (South China), Beringeria Special Issue, 2, 203–241.
SIVETER, D. J. and CHATTERTON, B. D. E. 1996. Silicified calymenid trilobites from the Mackenzie Mountains, northwest
Canada. Palaeontographica, Abteilung A, 239, 43–60, pls 1–5.
ŠNAJDR, M. 1958. Trilobiti c̆eského str̆ednı́ho kambria. Rozpravy Ústr̆ednı́ho ústavu geologického, 24, 1–279, pls 1–46.
—— 1980. Bohemian Silurian and Devonian Proetidae (Trilobita). Rozpravy Ústr̆ednı́ho ústavu geologického, 45,
1–323, pls 1–64.
SPEYER, S. E. and CHATTERTON, B. D. E. 1989. Trilobite larvae and larval ecology. Historical Biology, 3, 27–60.
STOCKTON, W. L. and COWEN, R. 1976. Stereoscopic vision in one eye: paleophysiology of the schizochroal eye of
trilobites. Paleobiology, 2, 304–315.
STØRMER, L. 1930. Scandinavian Trinucleidae with special reference to Norwegian species and varieties. Skrifter Utgitt
av der Norsk Videnskaps-Akademi i Oslo, 1930 (4), 1–111.
STUBBINGS, H. G. 1975. Balanus balanoides. Memoir of the Liverpool Marine Biology Committee on typical British
marine plants and animals, 37, 1–23.
TRIPP, R. P. and EVITT, W. R. 1986. Silicified trilobites of the family Asaphidae from the Middle Ordovician of Virginia.
Palaeontology, 29, 705–724.
WESTERGÅRD, A. H. 1922. Sveriges Olenidskiffer. Sveriges Geologiske Undersogelse, Series Ca, 18, 1–205, pls 1–16.
—— 1946. Agnostidea of the Middle Cambrian of Sweden. Sveriges Geologiske Undersogelse, Series C, 477, 1–141.
WESTROP, S. R. 1983. The life habits of the Ordovician illaenine trilobite Bumastoides. Lethaia, 16, 15–24.
WHITTARD, W. F. 1956. The Ordovician trilobites of the Shelve Inlier, west Shropshire. Part 2. Monograph of the
Palaeontographical Society, 110 (473), 41–70, pls 5–9.
WHITTINGTON, H. B. 1952. A unique remopleuridid trilobite. Breviora, 1952 (4), 1–9.
—— 1953. North American Bathyuridae and Leiostegiidae (Trilobita). Journal of Paleontology, 27, 647–678.
—— 1959. Silicified Middle Ordovician trilobites: Remopleurididae, Trinucleidae, Raphiophoridae, Endymioniidae.
Bulletin of the Museum of Comparative Zoology, Harvard University, 121, 371–496.
—— 1963. Middle Ordovician trilobites from Lower Head, western Newfoundland. Bulletin of the Museum of
Comparative Zoology, Harvard University, 129, 1–118.
—— 1965. Trilobites of the Ordovician Table Head Formation, western Newfoundland. Bulletin of the Museum of
Comparative Zoology, Harvard University, 132, 275–442.
—— 1971. Redescription of Marrella splendens (Trilobitoidea) from the Burgess Shale, Middle Cambrian, British
Columbia. Bulletin of the Geological Survey of Canada, 209, 1–24.
—— 1975. Trilobites with appendages from the Middle Cambrian, Burgess Shale, British Columbia. Fossils and
Strata, 4, 97–136.
—— 1977. The Middle Cambrian trilobite Naraoia, Burgess Shale, British Columbia. Philosophical Transactions of
the Royal Society of London, Series B, 280, 409–443.
—— 1980. Exoskeleton, moult stage, appendage morphology and habits of the Middle Cambrian trilobite Olenoides
serratus. Palaeontology, 23, 171–204.
—— 1985. The Burgess Shale. Yale University Press, New Haven, 151 pp.
—— 1988a. Hypostomes and ventral cephalic sutures in Cambrian trilobites. Palaeontology, 31, 577–610.
—— 1988b. Hypostomes of post-Cambrian trilobites. Memoirs of the New Mexico Bureau of Mines and Mineral
Resources, 44, 321–339.
—— 1993. Morphology, anatomy and habits of the Silurian homalonotid trilobite Trimerus. Memoirs of the
Association of Australasian Palaeontologists, 15, 69–83.
—— 1997. Illaenidae (Trilobita): morphology of thorax, classification, and mode of life. Journal of Paleontology, 71,
878–895.
—— and ALMOND, J. E. 1987. Appendages and habits of the Upper Ordovician trilobite Triarthrus eatoni.
Philosophical Transactions of the Royal Society of London, Series B, 317, 1–46.
—— and CAMPBELL, K. S. W. 1967. Silicified Silurian trilobites from Maine. Bulletin of the Museum of Comparative
Zoology, 135, 447–483, pls 1–19.
WILLS, M. A., BRIGGS, D. E. G., FORTEY, R. A. and WILKINSON, M. 1995. The significance of fossils in understanding arthropod
evolution. Verhandlungen der Deutschen Zoologischen Gesellschaft, 88, 203–215.
ZHANG, W.-T., ZHANG, S.-G., LIN, H.-L., ZHU, Z.-L., QIAN, Y.-Y., ZHOU, Z.-Y., YUAN, J.-L. and LU, Y.-H. 1980. Cambrian trilobite
faunas of southwestern China. Palaeontologica Sinica, New Series B, 16, 1–497, pls 1–134.
R. A. FORTEY
The Natural History Museum

Cromwell Road
London SW7 5BD, UK
R. M. OWENS
Typescript received 29 June 1998 National Museum of Wales

Revised typescript received 12 December 1998 Cardiff CF1 3NP, UK

Feedinghabitsintrilobites: by and

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FEEDING HABITS IN TRILOBITES

by R. A. FORTEY and R . M. OWENS

THE RANGE OF POSSIBILITIES FOR TRILOBITE FEEDING HABITS

PRIMITIVE PREDATORY FEEDING HABITS IN TRILOBITA

Strengthening of anterior cephalic doublure

Hypostome attachment style

Modifications of the posterior hypostome margin

TEXT-FIG. 6. Rusophycus trace fossils definitely associated with

High relative abundance

FILTER CHAMBER FEEDING

Filter chamber morphotypes

Functional aspects of trinucleimorph design

The problem of illaenid morphology

CHANGES IN FEEDING HABITS DURING ONTOGENY

CONCLUSION: TOWARDS EXPLAINING TRILOBITE FAUNAL DIVERSITY

The Natural History Museum

Typescript received 29 June 1998 National Museum of Wales

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