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.     

Palaeoecology and evolutionary implications of enrolled trilobites from the Kushan Formation,Guzhangian of North China

Damesella paronai is the earliest enrolled odontopleurid trilobite to be found in the fossil record. Whereas its interlocking devices were very poorly developed, its trunk shows advanced articulations, which only lack an articulating facet, though an anterior notch is present in the outer part of each pleura. Its body pattern and structures did not allow it an encapsulated enrolment style. On the other hand, it is coeval with the early asaphid Monkaspis daulis, found in the same beds and at the same level. This co-inhabiting trilobite had a more advanced structure that enabled it to achieve a fully enrolled, encapsulated style. These superior structures enhance the preservation of enrolled specimens of M. daulis, and it seems to be generally the case that more elegant structures actually improve preservation. The evolutionary trends of the lineages of these two trilobites show that while odontopleurids were very conservative, using the same enrolment style throughout their history, the asaphids developed different enrolment styles, which was one factor in giving them a capacity to occupy different ecological niches and a greater range of environments.     

Trilobite larvae and larval ecology

Silicified trilobite faunas yield a well preserved and abundant record of early development in many taxa. Although their morphologies are well known, protaspid larvae are poorly understood in terms of trilobite life history and developmental biology. Two distinct morphologic types are recognized among the great diversity of protaspides studied; these are termed adult‐like and nonadult‐tike according to gross similarity to later developmental stages. Transitions between nonadult‐like and adult‐like morphologies are abrupt, occurring between successsive instars, and, thereby, constitute metamorphoses.

By analogy with developmental patterns among modern marine arthropod taxa, metamorphoses during early trilobite ontogeny correspond to radical modifications in life mode and ecology. Adult‐like trilobite protaspides possess a dorso‐ventrally flattened tergum which displays a coarsely featured prosopon and surface‐parallel spines; these are interpreted as benthic larvae. In contrast, nonadult‐like protaspid larvae display an ovoid to spheroidal shape with spine pairs that project at right angles to one another; these are considered to have been planktonic. Protaspid morphologies are compared and discussed in light of these inferred modes of life.     

Ontogeny of the Upper Cambrian (Furongian) Olenid trilobite Protopeltura aciculate (Angelin, 1854) from Skåne and Västergötland,Sweden

Abstract: The ontogeny of the trilobite Protopeltura aciculata (Angelin, 1854) is described on the basis of material from the upper Cambrian (Furongian) of Andrarum (Skåne) and Hjelmsäter (Västergötland), Sweden. P. aciculata is present in the Parabolina brevispina and Parabolina spinulosa zones. Protopeltura aciculata is represented by all stages of growth, from early protaspides to holaspides, although most of the specimens are disarticulated and precise degrees are unknown. The cranidia have therefore been allocated to five morphological groups. Cuticular sculpture of the cranidia changes throughout ontogeny. Large tubercles are present in earlier stages, disappear gradually in middle meraspid stages and are replaced with a very faint granulation. The transitory pygidium, relatively large and shield‐shaped with upwardly and backwardly directed marginal spines in early meraspides, later becomes very small, triangular‐shaped and lacking spines as a late meraspid and holaspid. The development of hypostomes and librigenae is also described. Protopeltura aciculata shows major intraspecific variations throughout development, especially regarding the pygidium where variation is much less constrained than in many other olenids. This high developmental plasticity may be a survival strategy for a trilobite living in a stressed environment. Protopeltura inhabited a dysoxic environment, possibly unusually prone to localised spreading of anoxic or toxic water. Some morphs may have been less vulnerable than others to such stresses, surviving by chance and thus enabling the species to continue.     

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.     

Phylogenetic support for the monophyly of proetide trilobites

The monophyly of the order Proetida, the only trilobite group to survive the end‐Devonian mass extinction, has been regularly questioned since its erection almost three decades ago. Through analysis of a novel phylogenetic data set comprising 114 characters coded for 55 taxa, including both traditional members of the Proetida along with a number of other trilobite groups, the monophyly of proetide trilobites is rigorously tested for the first time. Proetida is shown to be monophyletic, united by the initial compound eye formation in early protaspids occurring at the lateral margin rather than the anterior margin, and the form of the protaspid glabella being tapering with a pre‐glabellar field. A number of adult characters, including the possession of a quadrate or shield‐shaped hypostome with angular posterior margins, the hypostome median body being divided by a deep groove that entirely traverses the median body, the presence of an enlarged thoracic spine on the sixth tergite and a tergite count of between 7 and 10, also define the basal node. Hystricurid and dimeropygoid trilobites are shown to resolve at the base of the group, while the remaining proetide taxa are divided between large proetoid and aulacopleuroid clades. Some taxa previously allied with Aulacopleuroidea, such as rorringtoniids and scharyiids, are retrieved as basal members of the Proetoidea.     

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.     

Silicified Upper Ordovician trilobites from Pai‐Khoi,Arctic Russia

Abstract: A collection of silicified trilobites extracted from samples obtained by F. Nansen from the Upper Ordovician of Khabarovo, Pai Khoi, Russian Arctic is dominated by two new species of the proetide Lasarchopyge, a genus known hitherto only from the Argentine Precordillera. Comparison of Lasarchopyge with Scharyia reveals a range of common characters that support inclusion of both genera in the same family. The rest of the fauna comprises a metagnostid (Trinodus elspethi), an asaphid (Isotelus sp.), two remopleuridids (Remopleurides cf. caelatus and R. sp. nov. A), an aulacopleurid (Harpidella triloba), a lichid and an odontopleurid (Apianurus cf. barbatus), the last two being represented by very fragmentary material. T. elspethi, H. triloba, R. cf. caelatus and A. cf. barbatus are common to or closely resemble taxa from the Edinburg Formation, Virginia. Closely related taxa are present also in the Esbataottine Formation, NW Territories, Canada, and in the Albany Group, Girvan district, Scotland. All of these occurrences are of early Caradoc (Sandbian) age, and most lay at palaeolatitudes of 20°–30° south; all are interpreted as having been deposited on the outer shelf, which is consistent with their distribution.     

Thoracic structure and enrolment style in middle Cambrian Eccaparadoxides pradoanus presages caudalization of the derived trilobite trunk

Abstract: The ability to enrol effectively evolved several times among trilobites. Here, we show that, unlike most redlichiid trilobites that could not enrol, both morphotypes of Eccaparadoxides pradoanus from the middle Cambrian of Spain enrolled so as to enclose most of the ventral surface beneath the exoskeleton and possessed specialized articulating devices that facilitated this behaviour. The holaspid thorax of all E. pradoanus was divided into two principal regions. The boundary between these marked a transition from anterior segments with short pleural spines, fulcra and ridge‐and‐groove inner pleural regions to posterior segments with longer, acuminate pleural spines that lack fulcra and inner pleural regions. Devices that aid articulation, such as fulcra with short articulating pleural surfaces, the petaloid articulating facet and long articulating half rings, are concentrated in the anterior region. These features, and the large number of specimens preserved in various degrees of enrolment, suggest an enrolment procedure in which the rear part of the trunk, containing both the posterior thorax and the pygidium, rotated as a single unit without significant internal flexure. As these posterior trunk articulations were apparently not required to permit enrolment, concentrating flexure in the anterior may have presaged the caudalized condition seen in many derived trilobite groups that encapsulated, in which a larger proportion of the trunk segments were allocated to the mature pygidium, and therefore unable to articulate.     

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.     

Egg capsules of the dusky catshark Bythaelurus canescens (Carcharhiniformes,Scyliorhinidae) from the south‐eastern Pacific Ocean

The external morphology of the egg capsule of Bythaelurus canescens and its fixation to the substratum are described. Bythaelurus canescens egg capsules are typically vase‐shaped, dorso‐ventrally flattened, pale yellow in colour when fresh and covered by 12–15 longitudinal ridges. The anterior border of the capsule is straight, whereas the posterior border is semicircular. Two horns bearing long, coiled tendrils arise from the anterior and posterior ends of the capsule. The presence of longitudinal ridges and long coiled tendrils at both anterior and posterior ends of the capsule readily distinguish these egg capsules from those of other chondrichthyans occurring in the south‐east Pacific Ocean.     

The biology and functional morphology of Arca noae (Bivalvia: Arcidae) from the Adriatic Sea, Croatia, with a discussion on the evolution of the bivalve mantle margin

In the Croatian Adriatic, Arca noae occurs from the low intertidal to a depth of 60 m; it can live for > 15 years and is either solitary or forms byssally attached clumps with Modiolus barbatus. The shell is anteriorly foreshortened and posteriorly elongate. The major inhalant flow is from the posterior although a remnant anterior stream is retained. There are no anterior but huge posterior byssal retractor muscles and both anterior and posterior pedal retractors. The ctenidia are of Type B(1a) and the ctenidial–labial palp junction is Category 3. The ctenidia collect, filter and undertake the primary sorting of potential food in the inhalant water. The labial palps are small with simple re‐sorting tracks on the ridges of their inner surfaces. The ciliary currents of the mantle cavity appear largely concerned with the rejection of particulate material. The mantle margin comprises an outer and an (either) inner or middle fold. The outer fold is divided into outer and inner components that secrete the shell and are photo‐sensory, respectively. The latter bears a large number of pallial eyes, especially posteriorly. The inner/middle mantle fold of A. noae, possibly representative of simpler, more primitive conditions, may have differentiated into distinct folds in other recent representatives of the Bivalvia.     

Unusual trilobite biofacies from the Lower Ordovician of the Argentine Cordillera Oriental: new insights into olenid palaeoecology

Balseiro, D., Waisfeld, B.G. & Buatois, L.A. 2010: Unusual trilobite biofacies from the Lower Ordovician of the Argentine Cordillera Oriental: new insights into olenid palaeoecology. Lethaia, Vol. 44, pp. 58–75. The study of biofacies has proven to be relevant in the understanding of trilobite palaeoecology, palaeobiogeography and macroevolution. The widespread Olenid biofacies is one of the best known, and is usually interpreted as occuring in dysoxic environments. Tremadocian successions of the Argentinian Cordillera Oriental bear a diverse and long‐studied olenid‐dominated fauna. Based on cluster analysis, five distinct biofacies are defined for the middle Tremadocian (Tr2 stage slice), distributed from shelf (below storm wave base) to lower‐shoreface settings (above fair‐weather wave base). Ordination shows biofacies along two gradients, a bathymetrical one and another related to oxygen content. All of them are dominated both taxonomically and ecologically by olenids. This detailed quantitative palaeoecological study challenges current views suggesting instead that the Olenidae dominated a broad range of environments, from oxygenated shallow‐marine to dysoxic deep‐marine. Comparisons with largely coeval trilobite records from geodynamically and palaeobiogeographically disparate sites suggest that siliciclastic sedimentation appears as the most influential controlling environmental factor upon olenid distribution and dominance. Further comparisons across different climatic belts show that siliciclastic input controlled trilobite diversity gradients, even more than latitude. From an autoecological viewpoint distribution of traditional olenid morphotypes shows no relation to depth or to oxygen content, and at least some members of the group appear to have had the possibility of coping with low oxygen content, rather than being restricted to oxygen‐deficient environments. The analysis performed herein, together with recent research on the group, demonstrate that factors controlling olenid distribution are more complex than previously envisaged. □Biofacies, diversity, Olenidae, palaeoecology, Tremadocian, trilobite.     

Identifizierung vonAgnostus calkeri Kruizinga 1918 als larvales Asaphiden-Hypostom (Trilobita)

New findings in a geschiebe (glacial erratic boulder) of the Red Orthoceras Limestone from the Münsterland (Westphalia) ofAgnostus calkeri originally described 1918 byKruizinga from a Dutch geschiebe of the same kind proved to be an larval asaphid hypostome.     

Facies of a Lower Ordovician carbonate shelf (Mungok Formation: Taebaeksan Basin,Korea)

Summary The lithologic associations within the Lower Ordovician Mungok Formation in Korea define four depositional facies that formed across a continental margin fringing the Sino-Korean block: these facies represent lagoonal/restricted marine, shoal, inner shelf, and outer shelf environments. The stacking pattern of these facies reveals two systems tracts composed of five depositional sequences. The lower highstand systems tract consists of the lagoonal/restricted marine and shoal facies, whereas the upper lowstand systems tract comprises, in ascending order, inner shelf, outer shelf, and inner shelf facies. Three trilobite biofacies are recognized in the Mungok Formation: i.e.,Yosimuraspis, Kainella, andShumardia biofacies in ascending order. TheYosimuraspis Biofacies is dominated byYosimuraspis but also containsJujuyaspis andElkanaspis. The predominance of the endemic eponymous taxon suggests a lagoonal/restricted marine environment. The nearly monotaxicKainella Biofacies, which comprises pandemic genera such asKainella and occasionallyLeiostegium, may represent a less restricted environment than theYosimuraspis Biofacies. TheShumardia Biofacies occurs in the marlstone/shale lithofacies through relatively thick stratigraphic interval and is dominated by cosmopolitan trilobite taxa with some endemic species. The lithofacies and cosmopolitan trilobite assemblage of theShumardia Biofacies indicate that it occupied an outer shelf environment. The vertical succession of lithofacies and trilobite biofacies in the Mungok Formation records in general a shift from a restricted, shallow water environment to deeper-water environment.     

Biostratigraphy of the Peri-Gondwana Cambrian trilobite fauna (northern Kerman,Iran) and correlation with other countries

Polymeroid trilobite fauna from two stratigraphic sections (Godbondar and Kuhbanan) of the Kuhbanan Formation in northern Kerman (central Iran) were studied and subjected to biostratigraphic analysis. Eleven genera and species are recognised from the latest Early Cambrian and Middle Cambrian Peri-Gondwanian successions of the study sections. The recognised fauna includes Afghanocare lategenatum, Blountia blountia, Iranoleesia sp., Iranoleesia pisiformis, Kermanella kuhbananensis, Kermanella lata lata, Kermanella lata minuta, Kermanella sp., Redlichia chinensis, Redlichia noetlingi and Redlichia sp. Based on trilobite distribution, three trilobite biozones were recognised in the study sections, namely Redlichia noetlingi biozone, Kermanella kuhbananensis biozone and Iranoleesia pisiformis biozone. The age of the study sections is late Early Cambrian to late Middle Cambrian based on the recognised trilobite biozones. The recognised late Early Cambrian trilobite assemblages (especially Redlichia and Kermanella) from northern Kerman are similar to those found from some other parts of Gondwana or Peri-Gondwana terrains (north India, Pakistan, northwestern Kashmir, Tajikistan, South Australia, South China and Afghanistan) and show affinities with fauna found in some other parts of Iran (Alborz, northern Iran; Tabas, eastern Iran; southeast Karman).     

The fowl tick, <Emphasis Type="Italic">Argas (Persicargas) persicus</Emphasis> (Ixodoidea: Argasidae): Description of the egg and redescription of the larva by Scanning Electron Microscopy

This scanning electron microscopy study revealed that the egg of Argas persicus was covered with chorion which appeared as a wrinkled layer containing regions of three textures. The first had elevated parts of slightly rough surface. The second had irregular smooth elevations; each carried numerous parallel horizontal foldings with vertical ridges. The last region had rough surface with irregularly shaped projections. Following the removal of the chorion, shell was observed to have one polar micropyle and numerous slit like openings. Length (L), width (W) and L/W ratio of the egg were measured. Investigation of larvae revealed extensively folded integument of idiosoma and spherical or elongated tubercles on dorsal plate. Mouth enclosed between ventral hypostome and two dorsal chelicerae. Hypostome carried four longitudinal rows of conical denticles. Each chelicera was made up of two segments; the basal one appeared as a pocket for the distal one. Haller’s organ consisted of an anterior pit containing seven sensilla and a posterior capsule with four apertures. Distribution of chemo- and mechano-sensilla on the body was examined. Measurements of whole body, idiosoma, dorsal plate, capitulum, hypostome, palp and different types of sensilla both on the body and Haller’s organ are also presented.     

A rare non‐trilobite artiopodan from the Guzhangian (Cambrian Series 3) Weeks Formation Konservat‐Lagerstätte in Utah,USA

We describe a weakly biomineralized non‐trilobite artiopodan arthropod from the Guzhangian Weeks Formation of Utah. Falcatamacaris bellua gen. et sp. nov. is typified by a thin calcitic cuticle, broad cephalon without eyes or dorsal ecdysial sutures, an elongate trunk with distinctively sickle‐shaped pleural spines and a long tailspine with a bifurcate termination. The precise affinities of Falcatamacaris gen. nov. are problematic due to the presence of unique features within Artiopoda, such as the peculiar morphology of the pleural and posterior regions of the trunk. Possible affinities with aglaspidid‐like arthropods and concilitergans are discussed based on the possession of 11 trunk tergites, edge‐to‐edge articulations and overall body spinosity. The new taxon highlights the importance of the Weeks Formation Konservat‐Lagerstätte for further understanding the diversity of extinct arthropod groups in the upper Cambrian.     

Mansuyia Sun,and Tsinania Walcott,from the Furongian of North China and the evolution of the trilobite family Tsinaniidae

The Furongian trilobite family Tsinaniidae is characterized by a highly effaced surface and forms an important constituent of the Furongian trilobite faunas of east Gondwana. However, the origin of the characteristic morphology of this family has remained unclear. Only recently has the tsinaniid trilobite Lonchopygella megaspina been suggested to represent an intermediate stage in the evolutionary transition to other tsinaniids on the basis of the trunk segmentation. Here, we report successive occurrences of four species of the kaolishaniid genus Mansuyia and a tsinaniid trilobite Tsinania canens from the Furongian (late Cambrian) Chaomidian Formation in Shandong Province, China. A cladistic analysis including these taxa reveals that the four species of Mansuyia constitute stem‐group taxa to the family Tsinaniidae, rendering Mansuyia and the Kaolishaniidae paraphyletic. The youngest species of Mansuyia, M. taianfuensis, turns out to be the immediate sister taxon of the Tsinaniidae, displaying a closely similar morphology to the tsinaniid trilobite, Shergoldia laevigata. The generic and familial boundary therefore situated between M. taianfuensis and S. laevigata.     

Tentacle morphogenesis in hydra

Summary Hydra oligactis exposed to 3 g/ml actinomycin D for 24 hours regenerated only the first pair of tentacles (the mid-laterals). If left uncut, actinomycin D treated animals underwent a reduction of the normal number of tentacles to two or less.Inductive activity was retained in the 2-tentacled hypostomes. However, the tentacles present exhibited reduced capacities to capture and manipulate prey.Histological studies showed that the tentacles of actinomycin D treated hydra were morphologically identical to those of the controls. The interstitial cell (I-cell) population of the treated animals, however, became depleted. Replacing the hypostome of an actinomycin D treated hydra with a normal hypostome reversed the cellular effects of actinomycin D treatment.The modifications in tentacle morphogenesis occurring after actinomycin D treatment are consistent with an impairment of hypostomal function in the animal. It is suggested that the morphological site of this malfunction may be in the nervous system.Research supported by American Cancer Society Institutional Grant No. 342-9157, USPHS Institutional Grant No. 342-9241 and by a grant from Research Corporation.Part of this work was completed while L.H. was an undergraduate research student supported by the National Science Foundation.