The function of forks: Isotelus‐type hypostomes and trilobite feeding

Hegna, T.A. 2010: The function of forks: Isotelus‐type hypostomes and trilobite feeding. Lethaia, Vol. 43, pp. 411–419. Despite previous investigations, the function of the forked morphology of asaphid trilobite hypostomes is enigmatic. The focus of this study is the large and robust forked hypostome of the largest known genus of trilobite, Isotelus, and the independently‐derived forked hypostome of Hypodicranotus, the longest hypostome relative to body size of any trilobite. Although the trilobite hypostome is analogous to the labrum in other arthropods, forked hypostomes lack an obvious modern functional counterpart. The Isotelus hypostome is distinguished from other trilobite hypostomes by closely‐spaced terrace ridges on a greatly thickened inner surface of the forked posterior margin, with the scarp of the terrace facing antero‐ventrally. This is compatible with a grinding function, suggesting possible limb differentiation to complement this structure. The inner face of the tine (one of the two, prominent, sub‐parallel posterior projections) is also unique in that it has a microstructure which is evident in section, running perpendicular to the surface. Macropredatory and filter‐feeder roles are ruled out, and previous characterizations of the hypostome as knife‐like or serrated are rejected. Its function is incompatible with that of other non‐asaphid trilobites with forked hypostomes, like the remopleuridid Hypodicranotus, which lack similar terrace ridges and thickened inner‐edge cuticle. □Arthropoda, Asaphida, ecology, functional morphology, Trilobita.     

The stability of thoracic segmentation in trilobites: a case study in developmental and ecological constraints

The decline in origination rate of new metazoan body plans following the Cambrian radiation has been suggested to reflect developmental canalization in derived taxa, limiting their ability to evolve forms with radically different morphotypes. Segmentation is a fundamental aspect of arthropod body plan, and here we show that a derived trilobite that secondarily converged on a morphotype characteristic of basal members of the clade also reverted to a pattern of segmental variability common among basal trilobites. Hence a secular trend in loss of variability of the trilobite thorax was not due to the evolution of an inviolable developmental constraint. This result challenges the notion of developmental canalization in phylogenetically derived taxa. Rather, early variability in trilobites may be the result of ecological factors that promoted segment-rich thoracic morphotypes during Cambrian time.     

Wolf-Ernst Reif (1945–2009)

It has been claimed that olenellids, long regarded as trilobites, are more closely related to chelicerates than to the rest of the trilobite clade (Lauterbach 1980). This was based on an interpretation of the homologies of segmental arrangement in the thorax, and of the thoracic axial spine. It is shown that there are more synapomorphies uniting accepted trilobites with the olenellids, than there are uniting olenellids with chelicerates. At least seven characters serve to define Trilobita as a natural, monophyletic group. These include the presence of a pygidium, the unique optical system, the presence of eye ridges, circum‐ocular ecdysial sutures, and the construction of the hypostome. Olenelloids include the most primitive of the trilobites, retaining three (possibly four) primitive characters, including the permarginal suture, flange‐like thoracic articulation and highly expressed segmentation on the larval cranidia. If facial sutures are primitively absent in the Trilobita there is no obvious olenelloid autapomorphy, and they may constitute a paraphyletic group.     

DEVONIAN TRILOBITES FROM THE FALKLAND ISLANDS

Abstract:  New trilobite material is described from the Fox Bay Formation (Lower Devonian; Pragian) of the Falkland Islands (South Atlantic). The fossils were collected from many localities in both East and West Falkland and represent the first significant collection of trilobites from East Falkland. The trilobites belong to three higher taxa: Proetida (the first proetid record in these islands); Homalonotidae, Burmeisteria Salter, 1865; and Calmoniidae, including Bainella Rennie, 1930 and Metacryphaeus Reed, 1907. All of the forms except proetids have now been found in both East and West Falkland.     

Head segmentation of trilobites

Although trilobites have provided research subjects for more than two centuries, their head segmentation has remained unresolved. Four glabellar furrows (SO and S1–S3) marking the segmental boundaries are generally present in the cephalic axis, but there are trilobites with one more pair of furrows, the so‐called S4, in the cephalic axis, causing confusion in understanding trilobite head segmentation. Recent advances in developmental biology and palaeontology have shed light on the arthropod head problem, and thus, trilobite head segmentation can be reviewed in the light of this knowledge. Based on the information from the anatomy of exceptionally preserved trilobites and artiopodans closely related to trilobites, it is inferred that trilobite head contains five segments: the anteriormost ocular segment potentially associated with the hypostome, the antennal segment and the following three segments with walking legs. When present, the S4 furrows are situated where the eye ridges meet the cephalic axis of trilobites, indicating that the furrows are incised ‘within’ the anteriormost segment in trilobites with an anteriorly enlarged frontal lobe. Trilobites of the Order Redlichiida, the most primitive stock, show variable conditions in the frontal glabellar conditions, while in other more derived groups, the condition is rather constant. The frontal glabellar condition, therefore, could provide a clue to elucidate the unresolved Cambrian trilobite phylogeny and the Cambrian roots of the post‐Cambrian trilobites.     

Inferring internal anatomy from the trilobite exoskeleton: the relationship between frontal auxiliary impressions and the digestive system

Lerosey‐Aubril, R., Hegna, T.A. & Olive, S. 2011: Inferring internal anatomy from the trilobite exoskeleton: the relationship between frontal auxiliary impressions and the digestive system. Lethaia, Vol. 44, pp. 166–184. The digestive system of trilobites is rarely preserved. As a result, many aspects of its organization remain unknown. Fortunately, the exoskeleton sometimes preserves evidence of soft‐tissue attachment sites that can be used to infer internal anatomy. Among them are the frontal auxiliary impressions (FAIs), probable soft‐tissue insertion sites located on the fronto‐median glabellar lobe of some trilobites. FAIs are herein described in the Carboniferous trilobite Phillipsia belgica Osmólska 1970 – representing the only known example of such structures in the Proetida and their youngest occurrence. A taphonomic scenario is proposed to explain their variable preservation. Although particularly common in the Phacopina, FAIs or FAI‐like structures are also found in several orders that differ greatly. Comparisons with modern analogues suggest that FAIs might represent attachment sites for extrinsic muscles associated with a differentiated crop within the foregut. A review of purported remains of the trilobite digestive system indicates that it usually consisted of a tube‐like tract flanked by a variable number of metamerically paired diverticulae. Its anterior portion is not particularly individualized, except in a few specimens that might hint at the presence of a crop. This differentiation of a crop might have constituted a secondarily evolution of the foregut in trilobites, occurring independently in different clades. Accompanied by a strengthening of associated extrinsic muscles, this modification of the foregut might explain the presence of more conspicuous muscle insertion sites on the glabella. Study of FAIs might therefore provide new data on the anatomy of the foregut in trilobites and evidence of diverse feeding habits. □Arthropoda, digestive system, ecology, muscle scars, Proetida, Trilobita.     

Rusophycus carleyi (James, 1885), Trace Fossils from the Lower Ordovician of Southern Morocco,and the Trilobites that Made Them

Thin-bedded, pyrite-rich, fine sandstones and mudstones of the Floian-Dapingian Upper Fezouata Formation contain abundant trace fossils Rusophycus carleyi in close association with a species of the asaphid trilobite Asaphellus. The sizes and shapes of this trilobite and the traces match closely. Five specimens have even been found where an articulated specimen of Asaphellus appears to be directly located over a specimen of Rusophycus carleyi within a thin bed of sandstone, suggesting that the trilobite animal may have been trapped on top of a trace that it had just made. Such intimate associations between a putative tracemaker and a trace are rare in the fossil record and particularly rare for Trilobita. The number of coxal impressions that form part of R. carleyi, eleven, matches the number expected for an asaphid trilobite (one for each of eight thoracic segments and one for each of three post-oral cephalic appendages). Impressions of the hypostome, thoracic tip impressions, cephalic margin, and pygidial margin in a few of the traces also match those of this asaphid trilobite. R. carleyi has been found in Ordovician strata of other parts of the world in association with asaphid trilobites.     

Exceptionally preserved Cambrian trilobite digestive system revealed in 3D by synchrotron-radiation X-ray tomographic microscopy

The Cambrian 'Orsten' fauna comprises exceptionally preserved and phosphatised microscopic arthropods. The external morphology of these fossils is well known, but their internal soft-tissue anatomy has remained virtually unknown. Here, we report the first non-biomineralised tissues from a juvenile polymerid trilobite, represented by digestive structures, glands, and connective strands harboured in a hypostome from the Swedish 'Orsten' fauna. Synchrotron-radiation X-ray tomographic microscopy enabled three-dimensional internal recordings at sub-micrometre resolution. The specimen provides the first unambiguous evidence for a J-shaped anterior gut and the presence of a crop with a constricted alimentary tract in the Trilobita. Moreover, the gut is Y-shaped in cross section, probably due to a collapsed lumen of that shape, another feature which has not previously been observed in trilobites. The combination of anatomical features suggests that the trilobite hypostome is functionally analogous to the labrum of euarthropods and that it was a sophisticated element closely integrated with the digestive system. This study also briefly addresses the preservational bias of the 'Orsten' fauna, particularly the near-absence of polymerid trilobites, and the taphonomy of the soft-tissue-harbouring hypostome.     

Trilobite monophyly revisited

Fortey's and Whittington's recent refutation of Lauterbach's hypothesis of a paraphyletic Trilobita is supported. However, much of the character evidence raised by Fortey and Whittington to substantiate the monophyly of the Trilobita (including, inter alia, "Olenellinae”; and Agnostoidea) is ambiguous. Of seven proposed synapomorphies, only one (dorsal cuticle calcification) may be maintained at that node after testing within a cladistic framework. The other six characters are either constrained by calcification or define nodes up or down the cladogram. As positioned by Fortey's and Whittington's characters, Agnostoidea could be regarded either as the most primitive trilobites, or as being outside that clade. Lauterbach's support for an “olenelline"‐chelicerate clade is found to include interdependent characters which are reduced here to two testable derived similarities. Only one of these may conform to general criteria indicative of homology, such as detailed similarity and topology. It is, however, rejected on the basis of parsimony. We emphasize that resolution of the chelicerate‐"olenelline"‐trilobite three‐taxon problem must be based on recognition of homologies among each of these taxa. Nectaspida are excluded from Trilobita as defined by cuticle calcification, but as ingroup “Arachnata”; (sensu Lauterbach) they are important for determining character generality in this clade.     

THE LOWER CAMBRIAN EODISCOID TRILOBITE CALODISCUS LOBATUS FROM SWEDEN: MORPHOLOGY, ONTOGENY AND DISTRIBUTION

Abstract:  Several thousand disarticulated remains together with a few complete enrolled specimens of the lower Cambrian eodiscoid trilobite Calodiscus lobatus ( Hall, 1847 ) have been collected at two outcrop areas in Sweden. The material reveals new details of morphology and morphogenesis during ontogenetic development. Size-frequency analyses show that the material from the Fånån rivulet in Jämtland, central Sweden, represents a natural population dominated by juveniles, whereas the material from Gislövshammar in Scania, southern Sweden, has been sorted during postmortem transport. Three stages of protaspid development can be traced and defined as well as all subsequent ontogenetic stages for the cephalon, hypostome and pygidium. The early meraspid pygidium has a pronounced larval notch, which persists, though becoming progressively less distinct in later meraspides. The number of axial rings in the transitory pygidium increases throughout meraspid development until a third and final thoracic segment is liberated. During ontogeny the articulating half-rings are strongly developed, and both meraspides and holaspides were capable of full sphaeroidal enrollment and outstretched postures. The hypostome undergoes some dramatic modifications; in M0 the anterior margin is axe-shaped, by M1 the area of attachment greatly decreases and the hypostome becomes more elongated and pear-shaped, before attaining its adult form, which has an overall resemblance to that of polymerid trilobites. During ontogeny, the hypostome changes from a conterminant attachment to a natant condition, thereby mirroring hypostomal evolution within trilobites generally. The morphology, ontogeny, enrollment, hypostomal development and the presence of calcified protaspides suggest polymerid rather than agnostoid affinities of the eodiscoids.     

The Phylogeny and Systematics of Blind Cambrian Ptychoparioid Trilobites

The paraphyletic trilobite suborder Ptychopariina includes a large proportion of Cambrian trilobite diversity and is probably ancestral to most groups of post-Cambrian trilobites. Resolution of the phylogenetic relationships within the group is therefore crucial to a better understanding of the initial radiation of trilobites. The recognition of approaches that can successfully resolve the relationships of ptychoparioid taxa is an important first step towards this aim. Cladistic analysis was used to determine relationships within the Cambrian ptychoparioid trilobite family Conocoryphidae, and to test claims that the family is polyphyletic. Ninety-seven characters were coded for 40 conocoryphid species and nine non-conocoryphids. The results indicate that the family consists of four distantly related clades. Three are recognized here as distinct families, including an extensively revised Conocoryphidae, and the families Holocephalidae and Atopidae. The fourth clade is referred to the subfamily Acontheinae (Corynexochida) as the new Tribe Hartshillini. Analysis of the disparity of these four clades shows that they are significantly less morphologically variable than the original polyphyletic taxon, demonstrating the possible effects of taxonomic error on macroevolutionary studies of morphological disparity.     

A FUNCTIONAL STUDY OF THE SILURIAN ODONTOPLEURID TRILOBITE LEONASPIS DEFLEXA (LAKE)

The odontopleurid trilobite Leonaspis deflexa (Lake) was functionally capable of life in two alternative attitudes. In the 'resting' attitude the whole body could lie outstretched on the sea floor, supported on the hindmost anterior denticles of the cephalon and on the genal, thoracic, and pygidial spines. In this attitude the body declined posteriorly.
In the 'active' attitude, however, the cephalon was tilted forwards through 25°, bringing the body up horizontally. The hypostome and presumably the mouth also could then have been brought into close proximity with the sea floor.
Leonaspis, Primaspis, Diacanthaspis , and Anacaenaspis are all functionally comparable. Dudleyaspis, Acidaspis , and other genera were specialised permanently for life in an attitude analogous to the 'active' posture of Leonaspis.
The life attitudes postulated for odontopleurids are compared with those of other trilobites and some comments are given on the evolution of the different patterns.     

Gut evolution in early Cambrian trilobites and the origin of predation on infaunal macroinvertebrates: evidence from muscle scars in Mesolenellus

Trilobites are particularly common Cambrian fossils, but their trophic impact on the rapidly evolving marine ecosystems of that time is difficult to assess, due to uncertainties on how diverse their feeding habits truly were. Gut anatomy might help to constrain inferences on trilobite feeding ecology, but preservation of digestive organs is exceedingly rare. Muscle scars on the glabella, known as ‘frontal auxiliary impressions’ (FAIs), have been interpreted as evidence of the evolution of a pouch‐like organ with powerful extrinsic muscles (i.e. a crop) in some trilobites. Here we describe FAIs in Mesolenellus hyperboreus from Cambrian Stage 4 strata of North Greenland, which represents the oldest example of such structures and their first report in the Suborder Olenellina. Mesolenellus FAIs suggest that the crop in trilobites was clearly differentiated from the rest of the digestive tract, and essentially located under a hypertrophied glabellar frontal lobe. Reviews of the digestive anatomy of trilobite sister‐taxa and the glabellar morphology of the oldest‐known trilobites suggest that the gut of the trilobite ancestor was an essentially simple tract (i.e. no well‐differentiated crop) flanked laterally by numerous midgut glands. A crop first evolved in the Cambrian in groups like olenelloids and (later) paradoxidoids. Using ichnological evidence, we hypothesize that the emergence of olenelloids yields evidence for the evolution of predatory inclinations in a group of arthropods originally dominated by surface‐deposit‐feeders. By allowing the exploitation of a rapidly developing food source, infaunal animals, the diversification of feeding strategies in trilobites might partially explain their unparalleled evolutionary success.     

Phylogenetic analysis of the Malacostraca (Crustacea)

The Malacostraca comprises about 28 000 species with a broad disparity in morphology, anatomy, embryology, behaviour and ecology. The phylogenetic relationships of the major taxa are still under debate. Is the Leptostraca the sister group of the remaining Malacostraca, or is this taxon more closely related to other Crustacea? Does the Stomatopoda or the Bathynellacea represent the most basal taxon within the remaining taxa? Is the Peracarida monophyletic or are some peracarid taxa more closely related to other ‘caridoid’ taxa? Is the Thermosbaenacea part of the Peracarida or its sister group, and how much support is there for a taxon Amphipoda + Isopoda? To answer these questions a phylogenetic analysis of the Malacostraca combining different phylogenetic approaches was undertaken. In a first step, the monophyly of the Malacostraca including the Leptostraca is shown using the ‘Hennigian approach’. A computer cladistic analysis of the Malacostraca was carried out with NONA and PEE ‐WEE , based on 93 characters from morphology, anatomy and embryology. Nineteen higher malacostracan taxa are included in our analysis. Taxa whose representatives are exclusively fossils were not included. The Leptostraca was used as an operational out‐group. The present analysis supports the basal position of the Stomatopoda. Syncarida and Peracarida (including Thermosbaenacea) are supported as monophyletic, the Eucarida is not. Instead a sister‐group relationship is suggested between Euphausiacea and Peracarida (including Thermosbaenacea), with the Syncarida as the sister group to both taxa. Certain embryonic characters are interpreted as support for the monophyly of the Peracarida (without Thermosbaenacea) because convergences or reversals of these characters seem implausible. Within the Peracarida, the Mysidacea (Lophogastrida + Mysida) represents the sister group to the remaining taxa. A sister‐group relationship between Amphipoda and Isopoda is not supported.     

Trilobite tagmosis and body patterning from morphological and developmental perspectives

The Trilobita were characterized by a cephalic region in whichthe biomineralized exoskeleton showed relatively high morphologicaldifferentiation among a taxonomically stable set of well definedsegments, and an ontogenetically and taxonomically dynamic trunkregion in which both exoskeletal segments and ventral appendageswere similar in overall form. Ventral appendages were homonomousbiramous limbs throughout both the cephalon and trunk, exceptfor the most anterior appendage pair that was antenniform, preoral,and uniramous, and a posteriormost pair of antenniform cerci,known only in one species. In some clades trunk exoskeletalsegments were divided into two batches. In some, but not all,of these clades the boundary between batches coincided withthe boundary between the thorax and the adult pygidium. Therepeated differentiation of the trunk into two batches of segmentsfrom the homonomous trunk condition indicates an evolutionarytrend in aspects of body patterning regulation that was achievedindependently in several trilobite clades. The phylogeneticplacement of trilobites and congruence of broad patterns oftagmosis with those seen among extant arthropods suggest thatthe expression domains of trilobite cephalic Hox genes may haveoverlapped in a manner similar to that seen among extant arachnates.This, coupled with the fact that trilobites likely possessedten Hox genes, presents one alternative to a recent model inwhich Hox gene distribution in trilobites was equated to eightputative divisions of the trilobite body plan.     

Trilobite body patterning and the evolution of arthropod tagmosis

Preservation permitting patterns of developmental evolution can be reconstructed within long extinct clades, and the rich fossil record of trilobite ontogeny and phylogeny provides an unparalleled opportunity for doing so. Furthermore, knowledge of Hox gene expression patterns among living arthropods permit inferences about possible Hox gene deployment in trilobites. The trilobite anteroposterior body plan is consistent with recent suggestions that basal euarthropods had a relatively low degree of tagmosis among cephalic limbs, possibly related to overlapping expression domains of cephalic Hox genes. Trilobite trunk segments appeared sequentially at a subterminal generative zone, and were exchanged between regions of fused and freely articulating segments during growth. Homonomous trunk segment shape and gradual size transition were apparently phylogenetically basal conditions and suggest a single trunk tagma. Several derived clades independently evolved functionally distinct tagmata within the trunk, apparently exchanging flexible segment numbers for greater regionally autonomy. The trilobite trunk chronicles how different aspects of arthropod segmentation coevolved as the degree of tagmosis increased.     

Trilobites in Paleozoic carbonate buildups

The trilobite faunas found in carbonate buildups throughout much of the Paleozoic are remarkably similar in both composition and general morphological characteristics. This is primarily the result of the domination of these faunas by a limited number of trilobite families which are long-ranging in the buildup environment. Illaenids are the dominant trilobites in Ordovician-Devonian buildups and are generally accompanied by members of the Lichidae and Cheiruridae. Proetid trilobites are characteristic of Ordovician-Permian buildups but are seldom common. The number of trilobite families found in carbonate buildups is highest in the late Ordovician, with most of the important families persisting into the Devonian. After the Devonian only proetids are found in this environment. The conservative nature of these trilobite faunas suggests that the environmental characteristics of normal-marine Paleozoic carbonate buildups are similar. Trilobita, palaeoecology, faunistics, morphology. carbonate buildups. Palaeozoic .     

Modelling enrolment in Cambrian trilobites

Trilobites were capable of enroling in different ways based on the flexible articulation of thoracic segments and associated interlocking devices; the type of enrolment (spiral or sphaeroidal) is thought to have largely depended on the coaptative devices that each trilobite used to enclose the body. Based on X‐ray microtomography scans of complete enrolled specimens from the Cambrian, we created three‐dimensional (3D) computer models to assess the kinematics needed to achieve both enrolment types. We demonstrate that closely related trilobites with little morphological variation (Bailiaspis?, Conocoryphe and Parabailiella) developed different enrolment types as a result of small variations in the number of thoracic segments and the angle between adjacent segments. Moreover, our models indicate that sphaeroidal enrolment, which is associated with a smaller number of thoracic segments, enabled faster encapsulation. This supports the hypothesis that there was a trend in the evolution of trilobites towards reduction in the number of thoracic segments in phylogenetically derived taxa in order to enhance the efficiency of enrolment.     

A statistical/computer-graphic technique for assessing variation in tectonically deformed fossils and its application to Cambrian trilobites from Kashmir

Hughes, N. C. & Jell, P. A. 1992 07 15: A stdlislical/compuler-graphic technique for assessing variation in tectonically deformed fossils and its application to Cambrian trilobites from Kashmir.
A combined approach using statistics and computer graphics can help resolve patterns of morphological variation in tectonically deformed fossils. Bivariate analyses and Principal Components Analysis can be used to identify a generalized strain vector in populations of deformed fossils, the identification of which permits discrimination of biological and tectonically induced variation. Results can be used to determine the number of taxa and growth relationships within the population. Statistically resolved taxa can be compared with species described from other areas using computer-aided shape restorations. Application of these techniques to a sample of trilobite cranidia from the Cambrian of Kashmir demonstrates that the variation of characters used to diagnose several genera and species are ontogeny- or deformation-controlled. Seven previously described species of Saukia, Prosaukia, Hundwarella and Anomocare are best considered as a single species, Hundwarella personara . These results suggest a Middle Cambrian age for the fauna, which has affinities with other faunas described from India and northern China. Hundwarella personara shows a pattern of developmental flexibility similar to that seen in other Cambrian trilobites. Morphometrics, computer-aided restoration, deformed fossils, allometry, trilobites, Cambrian, Kashmir, developmental flexibility .     

The interrelationships of proseriata (Platyhelminthes: seriata) tested with molecules and morphology

Proseriate flatworms are common members of the interstitial benthic fauna worldwide, predominantly occupying marine environments. As minute animals, having relatively few characters useful for cladistic analysis, they have been difficult to present in a phylogenetic framework using morphology alone. Here we present a new morphological matrix consisting of 16 putatively homologous characters and two molecular data sets to investigate further this major group of free-living members of the Platyhelminthes. Complete 18S rDNA (representing 277 parsimony-informative characters) from 17 ingroup taxa and partial 28S rDNA spanning variable expansion regions D1 to D3 and D1 to D6 (representing 219 and 361 parsimony-informative characters, respectively) from 27 and 14 ingroup taxa, respectively, were determined and aligned as complementary data sets. Morphological and molecular data sets were analyzed separately and together to determine underlying phylogenetic patterns and to resolve conflict between published scenarios based on morphology alone. The monophyly of the Proseriata cannot be confirmed categorically with any of these data sets. However, the constituent taxa are confirmed as basal members of the Neoophora, and a sister group relationship with Tricladida is rejected. Similarly, the monophyly of one of the two subtaxa of the Proseriata, the Lithophora, could not be confirmed with molecules. Concerning intragroup relationships, we could reject one of the two phylogenetic trees formerly proposed, as well as the clade Otoplanidae + Coelogynoporidae. However, a clade Otoplanidae + Archimonocelididae + Monocelididae (to which the Monotoplanidae belong) was supported, and the position of the genus Calviria shifted from the Archimonocelididae to the Coelogynoporidae.