The eyes of trilobites: The oldest preserved visual system

The oldest preserved visual systems are to be found in the extinct trilobites, marine euarthropods which existed between about 520 and 250 million years ago. Because they possessed a calcified cuticle, they have a good fossil record, and commonly the lens-bearing surfaces of their paired compound eyes are well preserved. The sublensar structures, however, remain unknown. Three kinds of eyes have been distinguished. Holochroal eyes, apomorphic for trilobites, typically have many contiguous small lenses, set on a kidney-shaped visual surface. Lens optics, angular range of vision, and ontogeny have been established for many compound eyes. Some pelagic trilobites have enormous eyes, subtending a panoramic field of view. Schizochroal eyes are found only in one group, the phacopids (Ordovician to Devonian). These have large lenses, separated from each other by cuticular material, and the lenses have a complex doublet or triplet internal structure, which could focus light sharply. The optics of phacopid eyes are becoming increasingly well known despite the fact that there are no direct counterparts in any living arthropods today. Schizochroal eyes are apomorphic for phacopids and were derived by paedomorphosis from a holochroal precursor. Abathochroal eyes are confined to a short-lived Cambrian group, the eodiscids (of which most representatives were blind). Less is known about them than other trilobite eyes and their origins remain obscure. Some trilobite groups had no eyes, but had other kinds of sensory organs. In Upper Devonian times several groups of trilobites independently underwent progressive eye-reduction leading to blindness, related to prevailing environmental conditions of the time. The last trilobites (of Carboniferous and Permian age), however, had normal holochroal eyes, which persisted until the final extinction of trilobites at the end of the Permian.     

Abnormalities in early Paleozoic trilobites from central and eastern China

Malformations are common in trilobites, but the majority of described specimens are from Europe and North America. Only a few abnormal trilobites have been reported from China. Ten abnormal trilobites from Cambrian, Ordovician and Silurian strata in central and eastern China are documented. The abnormal Ordovician trilobites are found for the first time in China. All malformations occur in the thoraxes and pygidia, and were caused by a sub-lethal predatory attack, genetic or embryological malfunction, or injury sustained during molting. It is difficult to identify the predators of the six injured trilobites, but potential predators include Cambrian non-trilobite arthropods, Ordovician cephalopods, and Silurian eurypterids, chondrichthyes or cephalopods, even cannibalistic trilobites. Abnormal specimens caused by sub-lethal predatory attacks mainly occur in Cambrian strata in China and other areas in the world, and are rare in post-Cambrian strata. This pattern may reflect the rise of predators and increased predation in the post-Cambrian, which led to an increased trilobite fatality rate, thus reducing the probability that injured specimens would become fossilized.     

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.     

A tiny eye indicating a planktonic trilobite

Abstract: Cambrian trilobites mainly lived on the sea floor, and up till now few, if any, unequivocally planktonic trilobites have been reported from earlier than the Ordovician. The late Cambrian (Furongian) to late Ordovician olenids are a distinctive group of benthic (sea‐floor dwelling) or nekto‐benthic trilobites. Here we show, however, that one recently described, miniaturized and very spiny olenid species, Ctenopyge ceciliae must have been planktonic (passively drifting or feebly swimming in the upper waters of the sea). This interpretation is based not only upon body form but also on the analysis of its visual system and may be one of the earliest records of the planktonic realm being invaded by trilobites.     

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.     

Trilobite cuticle thickness in relation to palaeoenvironment

100 thickness measurements from thin sections of cephala or pygidia of early Ordovician trilobites occurring across an onshore to offshore environmental gradient show that progressively greater maximum cuticle thickness was characteristic of increasingly inshore sites. There is a 40-fold difference between the thinnest and thickest cuticles, and exclusively thin cuticles are confined to the offshore Olenid Biofacies. Variability in cuticle thickness increases offshore to onshore. Environmental control is shown to be more influential on cuticle thickness than is the overall length of the trilobite: some comparatively large trilobites having thin cuticles and small trilobites thick cuticles. The environmental factors which might be responsible for the pattern are briefly discussed. The thin cuticles dominating the offshore Olenid Biofacies were probably appropriate for dysaerobic conditions. Thick cuticles in the most inshore biofacies may have offered protection against predators and turbulence, but the additional presence there of trilobites with thinner cuticles is considered to reflect the greater heterogeneity of the epeiric habitat.     

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.     

Asymmetry of Early Paleozoic trilobites

Asymmetry in fossils can arise through a variety of biological and geological mechanisms. If geological sources of asymmetry can be minimized or factored out, it might be possible to assess biological sources of asymmetry. Fluctuating asymmetry (FA), a general measure of developmental precision, is documented for nine species of lower Paleozoic trilobites. Taphonomic analyses suggest that the populations studied for each taxon span relatively short time intervals that are approximately equal in duration. Tectonic deformation may have affected the specimens studied, since deviations from normal distributions are common. Several measures of FA were applied to 3–5 homologous measures in each taxon. Measurement error was assessed by the analysis of variance (ANOVA) for repeated measurements of individual specimens and by analysis of the statistical moments of the distributions of asymmetry measures. Measurement error was significantly smaller than the difference between measures taken on each side of a specimen. However, the distribution of differences between sides often deviated from a mean of zero, or was skewed or kurtosic. Regression of levels of FA against geologic age revealed no statistically significant changes in levels of asymmetry through time. Geological and taphonomic effects make it difficult to identify asymmetry due to biological factors. Although fluctuating asymmetry is a function of both intrinsic and extrinsic factors, the results suggest that early Cambrian trilobites possessed genetic or developmental mechanisms used to maintain developmental stability comparable to those of younger trilobites. Although the measures are biased by time averaging and deviations from the normal distribution, these data do not lend strong support to 'genomic' hypotheses that have been suggested to control the tempo of the Cambrian radiation.     

Hypostoma of agnostid trilobites

The hypostoma, which is a ventral sclerite that covers the oral region, has recently been discovered in four species of agnostid trilobites. The discovery is significant because the suborder Agnostina is the only major trilobite taxon for which the hypostoma has not been described. The agnostid hypostoma is distinctly different in shape from that of non-agnostid trilobites. Minus its wings, it is widest posteriorly, lacks border furrows, has unusually long wings, and in some species is fenestrate. It is unattached to other sclerites, and presumably was held in place beneath the anterior part of the glabella by muscle tissue. Hypostomal differences probably indicate basic differences in food or feeding habits of agnostid and non-agnostid trilobites. This, along with other evidence, suggests a major divergence in mode of life of these two trilobite groups, and should be given strong consideration in future attempts at trilobite classification.     

Clustered trilobite assemblages in the Middle Devonian Hamilton Group

Clusters of well-preserved trilobites occur throughout lower and middle Paleozoic strata. This phenomenon, however, remains poorly understood. Several thin, regionally persistent beds in the Middle Devonian (Givetian) Hamilton Group of New York State display taphonomic and sedimentary features which indicate rapid burial by storm-generated clouds of fine-grained sediment. Furthermore, components in these units display little or no post-mortem disturbance and, therefore, reflect in faunal associations. These beds contain two types of clustered assemblages of the trilobites Phacops (Green), Greenops boothi (Green) and Dechenella rowi (Green). Body clusters comprise dense assemblages of three or more complete trilobite corpses. Moult clusters , in contrast, are composed of three more moult ensembles; i.e., articulated thoracopygidia in close proximity to corresponding cephala. Clusters are species-segregated and contain only a small portion of the total size range exhibited by trilobite species. Moreover, body clusters and moult clusters are morphometrically undifferentiated indicating that these two cluster modes represent related phenomena. Analogy with modern marine arthropods suggests that trilobites assembled into monospecific, age-segregated clusters and moulted prior to en masse copulation.     

Traces of predation in the Cambrian

Series two marks a revolution in Cambrian predation when new predators and new predation methods appeared, which led to general increase in predation intensities and in the diversity of prey groups. The number of bored taxa and taxa with the predation scars is similar in the Cambrian. Most of the borings are associated with brachiopods and most of the scars with trilobites. Brachiopods, arthropods, molluscs, cnidarians and echinoderms were the most common prey in the Cambrian. The Cambrian record of predation is dominated by damage inflicted on brachiopods and trilobites. The fossils with predation signs are known from a majority of paleocontinents and all the Cambrian series.     

The Cambrian radiation of bilaterians: Evolutionary origins and palaeontological emergence; earth history change and biotic factors

Evidence from a variety of research areas, including phylogenetic palaeobiogeographic studies of trilobites, indicates that there may be a fuse to the Cambrian radiation, with a duration on the order of 20–70 myr. Evolution in trilobites appears to have been powerfully influenced by the tectonic changes occurring at the end of the Neoproterozoic: especially the breakup of Pannotia. This continental fragmentation may have also elevated opportunities for vicariance and speciation in trilobites, and other metazoans, given that speciation rates at this time period were high, though not phenomenally so. This provides clear evidence that abiotic factors played an important role in motivating evolution during this key episode in the history of life; biotic factors probably also played a role. The evidence for the role of biotic factors is considered in light of information from some problematic Cambrian taxa. These may show affinities with modern problematic pseudocoelomate phyla, although Cambrian and modern exponents differ dramatically in body size.     

Aligned trace fossils from the Utica Shale: implications for mode of life and feeding in the trilobite Triarthrus beckii

The olenid trilobite Triarthrus commonly occurs in nearly monospecific assemblages within otherwise relatively barren black shales. As such, it has been proposed that these trilobites preferred dysoxic or even anoxic habitats and suggested feeding habits range from predation and particle feeding to chemoautotrophism. A unique bedding‐plane assemblage of aligned traces that grade from Rusophycus to Cruziana, with associated Triarthrus beckii carcasses, are described from three localities in the Indian Castle Formation (upper Utica Shale) just below a K‐bentonite bed. Although few body fossils are preserved, it is clear that the 15‐cm‐thick, laterally extensive Thruway K‐bentonite created a unique taphonomic window that preserved the activities of numerous olenid trilobites. Rusophycus and Cruziana, consistently observed in densities above 100 trackways/m², were excavated by the trilobites into the upper surface of a micro‐graded bed, likely a distal turbidite deposit. Sedimentological and trace metal data (Mo, Mn, V and U) support dysoxic but not persistently anoxic conditions through this interval. Measurements from over 500 individual trackways have mean orientations of 259.1° (WSW) at the Myers Rd. locality and 224.59° (SW) at the Dolgeville Dam locality. These orientations are closely similar to independent sedimentological indicators of current transport direction at these sites, which indicates that the trilobites preferentially faced into the prevailing current. The trackways provide compelling evidence that Triarthrus beckii individuals were engaged in feeding, probably assisted by current transport of particles. These results do not support previous suggestions that Triarthrus may have relied upon a chemoautotrophic life habit, but are consistent with suggestions of predation or scavenging from the sediment.     

Lower Cambrian trace fossil evidence for predation on trilobites

Predation upon trilobites previously has been inferred from large coprolites containing trilobite fragments, and from specimens of trilobites with healed wounds. The discovery of large burrows ( Dolopichnus gulosus , n. ichnogen., n. ichnosp.) in micritic quartz arenite of the Lower Cambrian Poleta Formation in Esmeralda County, Nevada, suggests that sea anemones preyed upon trilobites. Dolopichnus n. ichnogen., vertical cylindrical burrows with a central cylindrical core, is interpreted as dwelling burrows of sea anemones. In the specimens studied, the core contains coarser-grained material, and in one series of burrows, is composed of trilobitc fragments and micrite pellets. cemented with sparite. The central cylinder is interpreted to be a cast of the sea anemone's coelenteron, which in some specimens contains stomach contents.     

Deep Endichnial Cruziana from the Lower-Middle Ordovician of Spain — A Unique Trace Fossil Record of Trilobitomorph Deep Burrowing Behavior

Full reliefs of Cruziana furcifera from the Lower-Middle Ordovician quartzite sandstone beds (Pochico Formation, southern Spain) points to deep, infaunal burrowing of trilobites. Some specimens show an unusual vertical extension with a wider lower part and a narrower upper part in cross section. They are referred to trilobites, which burrowed deeply in the sediment and were oriented obliquely head down and tail up. Deep burrowing seems to be common for other members of the Cruziana rugosa group, foremost C. rugosa and C. furcifera, less for C. goldfussi. The deep burrowing recorded in the discussed trace fossils can be referred to the earliest common infaunalization caused by trilobites and other arthropods during the Ordovician, probably in a response to a food competition on the sea floor, which promoted a behavioral plasticity within the same taxon or closely related taxa of trilobites.     

A Curious Rusophycus (Arthropod Ichnofossil) Assemblage from the Upper Ordovician of Ontario,Canada

The ichnogenus Rusophycus includes a wide range of short bilobate excavations generally attributed to variable feeding behaviors of arthropods, especially trilobites. An unusual Rusophycus assemblage from Upper Ordovician Georgian Bay Formation in Ontario departs radically from previously described examples and presents new challenges for understanding the behavior represented by these traces. This specimen is unique in the arrangement of multiple Rusophycus burrows in a circular, lens-shaped array (as opposed to a linear or random arrangement typical of other Rusophycus assemblages). The size and shape of the individual Rusophycus components are consistent with traces attributed to the coeval trilobite Flexicalymene. Multiple Rusophycus assemblages likely reflect aggregations of trilobites in response to a local concentration of food. The topology of this particular Rusophycus assemblage suggests that the trilobites opportunistically exploited a rich and narrowly restricted food source, perhaps the decaying remains of a buried organism.     

Two middle Cambrian diceratocephalid trilobites, Cyclolorenzella convexa and Diceratocephalus cornutus, from Korea: development and functional morphology

Park, T.-Y. & Choi, D.K. 2010: Two middle Cambrian diceratocephalid trilobites, Cyclolorenzella convexa and Diceratocephalus cornutus , from Korea: development and functional morphology. Lethaia , Vol. 43, pp. 73–87.
Silicified sclerites of the latest middle Cambrian trilobites, Cyclolorenzella convexa and Diceratocephalus cornutus , have been recovered from the Sesong Formation, Korea. Their morphological similarity and stratigraphic occurrences suggest that D. cornutus is a descendant of C. convexa . The ontogenies of both trilobites demonstrate that a pair of long frontal horns in the cephalon of D. cornutus is an evolutionarily novel structure. It is inferred that redeployment of some pre-existing regulatory gene played a significant role in constructing the frontal horns of D. cornutus . The frontal horns may have been a defensive structure to deter predators. The facial suture of D. cornutus , which extends onto the frontal horns and splits them into the dorsal and ventral halves, was a solution to enable easier forward egression during ecdysis. □ Functional morphology, Korea, Middle Cambrian, ontogeny, trilobites .     

Cambrian agnostid communities in Tasmania

Jago, J. B.: Cambrian agnostid communities in Tasmania.
Two or possibly three different agnostid trilobite assemblages can be distinguished in the late Middle and early Upper Cambrian sequences of northern and western Tasmania. This is significant because in recent years agnostid trilobites have been widely used in local and international correlations of Middle and Upper Cambrian rocks. The three assemblages recognized are (1) an agnostid assemblage in which polymerid trilobites are abscnt, rare or present as thanatocoenotic fossils, (2) a ptychagnostid-non-nepeid assemblage, and (3) a nepeid-clavagnostid-peronopsid assemblage in which non-agnostid trilobites are abundant but ptychag-nostids are absent. It is proposed that assemblage (1) represents an open sea fauna, with assemblages (2) and (3) occurring in progressively shallower water.     

中国寒武纪三叶虫生物地理趋议

寒武纪三叶虫除北疆和藏南尚无可靠报道外,在中国各地均有广泛分布。三叶虫证据表明,除兴安区的额尔古纳一兴安地体和早寒武世中天山一北山地体显示了与西伯利亚和劳伦陆块在动物群方面的密切相关外,中国寒武纪的所有板块和大部分地体（包括中及晚寒武世的中天山～北山地体）均系当时东泛冈瓦纳的重要组成部分,在生物地理上彼此密切关联。对这些属于东泛冈瓦纳的中国区块的生物地理格局的进一步探讨,主要侧重于它们台地相浅水三叶虫动物群之间的对比,特别是立足于来自动物群纪录比较完整的华南、塔里木和华北板块的证据。综合分析研究表明这些板块和有关的地体在中及晚寒武世应归属同一生物地理区,而在早寒武世则可划分为两个生物地理亚区：一个由华南、塔里木板块和安南或印支地体组成,另一仅以华北板块为代表。     

Primary and diagenetic microstructures in trilobites

McAllister, John E. & Brand, Uwe 1989 01 15: Primary and diagenetic microstructures in trilobites. Lethaia , Vol. 22, pp. 101–111. Oslo. ISSN 0024–1164.
Ordovician and Devonian trilobites were studied by scanning electron microscopy (SEM) to define primary and diagenetic microstructures. Primary structures such as Osmólska cavities and hexagonal surface openings are present in samples obtained from both shale and limestone lenses in shales. Generally, samples with primary microstructures are also geochemically well preserved. Diagenetic microstructures were found in samples from shales and limestones, but those from limestones are the most geochemically and microstructurally altered. 'Dendritic' patterns and fused matrices were found in the altered cuticles of trilobites of different species and ages and are therefore considered a normal feature of the diagenetic alteration of cuticular calcite. Primary and diagenetic microstructures of trilobite cuticles are independent of size, species and age of the specimens, but are largely controlled by lithology and thus diagenesis. * Trilobita, primary-, diagenetic microstructures, fossil diagenesis .