Shape deformation during fossilization can prevent accurate reconstruction of an organism's form during life, hampering areas of paleontology ranging from functional morphology to systematics. Retrodeformation attempts to restore the original shape of deformed fossil specimens and requires an adequate knowledge of the deformation process. Although tectonic processes and retrodeformation are relatively well understood, research on quantifying the effect of compressive deformation on fossil morphology is scant. Here we investigate the factors that can cause changes in the shape of fossil specimens during compressive deformation. Three-dimensional (3D) models of trilobite cranidia/cephala are subjected to simulated deposition and compaction using rigid body simulation and scaling features of the open-source 3D software Blender. The variation in pitch and roll angle is lowest on flat surfaces, intermediate on tilted surfaces, and highest on irregular surfaces. These trends are reflected in the morphological differences captured by principal component scores in geometric morphometric analyses using landmarks. In addition, the different shapes of trilobite cranidia/cephala according to their systematic affinity influence the degree of angular variation, which in turn affects their posture—normal or inverted. Inverted cranidia/cephala show greater morphological variability than those with normal postures.

Walcottaspis vanhornei (Walcott, 1914) is a large, late Cambrian trilobite with a unique pygidial morphology known only from a narrow outcrop belt of the St. Lawrence Formation in the Upper Mississippi Valley. Found in carbonate-rich layers within heterolithic facies that represent the toesets of a prograding shoreface, it is restricted to a single or small number of parasequences. Only four specimens of any of its sclerites have been illustrated previously. Here holaspid examples of all its biomineralized sclerites are described and illustrated, along with a morphometric analysis of cranidial landmarks and landmarks plus semilandmarks for the pygidium. Ontogenetic allometry accounts for 29% of the variance among holaspid cranidia and includes a relative shortening and narrowing of the palpebral lobe and a reduction in the relative length and width of the frontal area. Notable pygidial phenotypic variation occurs in the extent of the postaxial region and in the proportion of the structure occupied by the axis. Phylogenetic analysis suggests that W. vanhornei is sister taxon to Dikelocephalus minnesotensis Owen, 1852, which also occurs in the St. Lawrence Formation and has a broadly similar cephalon but distinctive trunk. The holaspid pygidium of W. vanhornei is uniquely characterized by the interpleural furrows of the first two segments becoming abruptly obsolete on approaching the axial furrow. Some pygidia show narrow, shallow, flat-bottomed grooves etched into the internal sides of the dorsal surface or doublure that apparently represent infestation of the live trilobite.

Two specimens of Metaconularia manni (Roy, 1935) from the lower Middle Silurian Scotch Grove Formation (eastern Iowa) exhibit well-defined, relict soft parts replicated in silica. One of these specimens bears phosphatic periderm, whereas the other specimen is a mold. Present within the erect, undistorted apical region of the specimen preserving periderm, on opposite sides of the peridermal cavity, are two small, elongate masses of silica located near the midlines of two of the four faces. Present in the central portion of the other specimen, at a somewhat greater distance from the apex, are five pairs of hollow, elongate, keeled pouch-like bodies (hereafter pouches), the long axes of which converge on the center of the fossil. Each pair of pouches is associated with a short, narrow, gently curved or broadly U-shaped tube, also composed of silica. Additionally, two of the pouch/tube combinations are associated with a pair of rectilinear furrows that correspond to the paired internal carinae that straddled the conulariid's facial midlines. We interpret the paired pouches and short tubes in the moldic specimen as relic conulariid soft parts homologous, respectively, to the interradial gonads and retractor muscles of extant, stauromedusan and polypoid scyphozoan cnidarians. Unlike most conulariids, which exhibit four faces, this individual had five faces, an aberrant morphology known in one other conulariid. The two small masses in the other specimen are more difficult to interpret, but they, too, could be relic gonads or longitudinal muscles. These interpretations suggest that, as in certain extant scyphozoans, at least one conulariid lost the free-living, sexual medusoid life phase.

Efficient extraction of oxygen from ambient waters played a critical role in the development of early arthropods. Maximizing gill surface area enhanced oxygen uptake ability but, with gills necessarily exposed to the external environment, also presented the issue of gill contamination. Here we document setae inserted on the dorsal surface of walking legs of the benthic-dwelling middle Cambrian Olenoides serratus and on the gill shaft of the Late Ordovician Triarthrus eatoni. Based on their physical positions relative to gill filaments, we interpret these setae to have been used to groom the gills, removing particles trapped among the filaments. The coordination between setae and gill filaments is comparable to that seen among modern crustaceans, which use a diverse set of setae-bearing appendages to penetrate between gill filaments when grooming. Grooming is known relatively early in trilobite evolutionary history and would have enhanced gill efficiency by maximizing the surface area for oxygen uptake.

Acritarch biostratigraphic and δ13C chemostratigraphic data from the Krol A Formation in the Solan area (Lesser Himalaya, northern India) are integrated to aid inter-basinal correlation of early–middle Ediacaran strata. We identified a prominent negative δ13C excursion (likely equivalent to EN2 in the lower Doushantuo Formation in the Yangtze Gorges area of South China), over a dozen species of acanthomorphs (including two new species—Cavaspina tiwariae Xiao n. sp., Dictyotidium grazhdankinii Xiao n. sp.), and numerous other microfossils from an interval in the Krol A Formation. Most microfossil taxa from the Krol A and the underlying Infra-Krol formations are also present in the Doushantuo Formation. Infra-Krol acanthomorphs support a correlation with the earliest Doushantuo biozone: the Appendisphaera grandis-Weissiella grandistella-Tianzhushania spinosa Assemblage Zone. Krol A microfossils indicate a correlation with the second or (more likely, when δ13C data are considered) the third biozone in the lower Doushantuo Formation (i.e., the Tanarium tuberosum-Schizofusa zangwenlongii or Tanarium conoideum-Cavaspina basiconica Assemblage Zone). The association of acanthomorphs with EN2 in the Krol Formation fills a critical gap in South China where chert nodules, and thus acanthomorphs, are rare in the EN2 interval. Like many other Ediacaran acanthomorphs assemblages, Krol A and Doushantuo acanthomorphs are distributed in low paleolatitudes, and they may represent a distinct paleobiogeographic province in east Gondwana. The Indian data affirm the stratigraphic significance of acanthomorphs and δ13C, clarify key issues of lower Ediacaran bio- and chemostratigraphic correlation, and strengthen the basis for the study of Ediacaran eukaryote evolution and paleobiogeography.

UUID: http://zoobank.org/5289fdb2-0e49-4b3b-880f-f5b21acab371.

A Glyptagnostus reticulatus (Angelin, 1851)–bearing trilobite assemblage has been found from an unnamed Cambrian formation in the Northern Qilian Mountains area, of which geographical placement in the Cambrian is contested. Glyptagnostus reticulatus is a biostratigraphic indicator of the Furongian Series and Paibian Stage, and three agnostoid and six polymerid taxa from the Changgou section, Daliang area are described herein, along with conspecific forms from the nearby Chuancigou section. This well-preserved assemblage shows strong taxonomic affinity with northwestern Queensland, Australia, and western Hunan–eastern Guizhou, China, and likely comes from deep outer-shelf to slope setting associated with the Northern Qilian arc. It is consistent with other arguments that during the Cambrian, the Northern Qilian arc, along with the Hexi Corridor of the Alxa terrane, were more closely allied to South China than to North China.

Later Cambrian and earliest Ordovician trilobites and brachiopods spanning eight horizons from five localities within the Sông Mã, Hàm Rồng and Đông Sơn formations of the Thanh Hóa province of Việt Nam, constrain the age and faunal affinities of rocks within the Sông Đà terrane, one of several suture/fault-bounded units situated between South China to the north and Indochina to the south. ‘Ghost-like’ preservation in dolomite coupled with tectonic deformation leaves many of the fossils poorly preserved, and poor exposure precludes collecting within continuously exposed stratigraphic successions. Cambrian carbonate facies pass conformably into Lower Ordovician carbonate-rich strata that also include minor siliciclastic facies, and the recovered fauna spans several uppermost Cambrian and Lower Ordovician biozones. The fauna is of equatorial Gondwanan affinity, and comparable to that from South China, North China, Sibumasu and Australia. A new species of Miaolingian ‘ptychopariid’ trilobite, Kaotaia xuanensis, is described. Detrital zircon samples from Cambrian–Ordovician rocks of the North Việt Nam and Sông Đà terranes, and from Palaeozoic samples from the Trường Sơn sector of Indochina immediately to the south, contain a predominance of ages spanning the Neoproterozoic period and have a typical equatorial Gondwanan signature. We associate the Cambrian and Tremadocian of the Sông Đà terrane with areas immediately to the north of it, including the North Việt Nam terrane and the southern parts of Yunnan and Guangxi provinces of China.

Abundant articulated specimens of the oryctocarine trilobite Oryctocarella duyunensis from the lower Cambrian (Stage 4, Series 2) Balang Formation at the Bulin section in western Hunan Province, South China, permit the description of all meraspid degrees. The maximum number of thoracic segments observed in this collection is 11. Meraspid growth was accompanied by progressive and gradual change in overall form, and this animal showed an homonymously segmented trunk with variation in the number of pygidial segments during ontogeny. Such variation permits a variety of plausible explanations, but a model of successive instars defined by the number of thoracic segments, and in suborder by the number of pygidial segments, is highly unlikely to explain the growth pattern because it would result in the loss of trunk segments between some instars. Degree-based ontogenetic staging is compatible with the variation observed.

A short stratigraphic interval near Bulin in western Hunan (China) yields multiple specimens of the ~514-Myr-old oryctocarine trilobite Oryctocarella duyunensis. Size data obtained from these specimens indicate that, from meraspid degree 1 onward, degrees represent successive instars. Meraspid growth persisted until a terminal stage was reached, providing the first example of determinate growth in trilobites and, notably, in an early Cambrian species. The sample contains three varieties of such terminal stages, recognized as holaspids, with 9, 10, or 11 thoracic segments, respectively. During the meraspid phase, growth rates were not constant in this species. The pattern of growth seen in the Bulin assemblage differs modestly from that reported in the same species from two other localities, attesting to microevolutionary variation in developmental patterns among these collections.

In order to maximize the utility of future studies of trilobite ontogeny, we propose a set of standard practices that relate to the collection, nomenclature, description, depiction, and interpretation of ontogenetic series inferred from articulated specimens belonging to individual species. In some cases, these suggestions may also apply to ontogenetic studies of other fossilized taxa.

The Ao Mo Lae Formation of the Tarutao Group crops out on Thailand's Tarutao Island and contains a diverse assemblage of late Furongian trilobite taxa, including several endemic forms. This study presents a new genus and species, Satunarcus molaensis, discovered at two locations on the island. A cladistic analysis of the kaolishaniid subfamily Mansuyiinae in light of Satunarcus and similar genera known from across upper Cambrian equatorial Gondwanan rocks suggests that the subfamily is polyphyletic in its current definition, and thus is not a natural group. Separating Mansuyia Sun, 1924 from the other taxa conventionally placed in Mansuyiinae permits recognition of a previously unrecognized monophyletic subfamily Ceronocarinae new subfamily. As established herein, this kaolishaniid subfamily contains Satunarcus n. gen. and all genera previously recognized as Mansuyiinae. with the exception of Mansuyia itself. Ceronocarinae n. subfam. occur in middle Jiangshanian to middle Cambrian Stage 10 sedimentary rocks from Australia, South China, North China, and Sibumasu, with most genera endemic to Australia.

UUID: http://zoobank.org/618c5136-73f0-4912-a7d3-e56559d2a76c

Articulated meraspid and holaspid exoskeletons of Guangxiaspis guangxiensis from the Guole Township, Jingxi County, Guangxi Province, China, are preserved in mudstone deposited during an obrution event. The species has a short dorsal pre-cranidial median suture that splits ventrally into a pair of posteriorly divergent connective sutures. The rostral plate of G. guangxiensis is thus triangular in outline, as in the co-occurrent Shergoldia laevigata, which also bore a conterminant hypostome. These two taxa appear to be closely related. The cephalic venter of Shergoldia laevigata has recently been interpreted to suggest a diphyletic origin of the median suture within the order Asaphida, but Guangxiaspis guangxiensis, Shergoldia laevigata and other tsinaniid trilobites display several characters reminiscent of members of the non-asaphide suborder Leiostegiina. These include swellings adjacent to the margins of the L1 glabellar lobe, the shape and furrows of the glabella, a semi-circular pygidium with a long and thin axis, and macrospinous first opisthopleurae of the holaspid pygidium. Based on these characters and on other new information on the early ontogeny of other tsinaniids, all these taxa likely belong within Leiostegiina. This suggests that the median suture arose independently in corynexochide and asaphide trilobites. The degree of convergence between S. laevigata and members of the derived asaphide family Asaphidae was remarkable. Guangxiaspis guangxiensis shows marked morphological change during both meraspid and holaspid ontogeny and might include more than a single morphotype.

New collections of trilobites from the type section of the Parahio Formation in the Parahio Valley, Spiti, and from the Parahio, Karsha, and Kurgiakh formations in the Zanskar Valley, permit biozonation based on material precisely located within measured stratigraphic sections. Specimens preserved in limestone with mild tectonic deformation clarify the features of several Himalayan taxa known previously only from severely deformed specimens preserved in shale. A total of 75 trilobite taxa from the Cambrian of Spiti and Zanskar can be referred, questionably at least, at the generic level or below, and 61 of these are present in our new collections. This new material is assigned with confidence to 29 existing species, and to 12 new species. Three new genera, Haydenaspis, Bhargavia, and Himalisania, are established; new species include Haydenaspis parvatya, Prozacanthoides lahiri, Probowmania bhatti, Xingrenaspis parthiva, X. shyamalae, Bhargavia prakritika, Kaotaia prachina, Gunnia smithi, Sudanamonocarina sinindica, Proasaphiscus simoni, Koldinia odelli, and Torifera jelli. Ten additional Himalayan forms are assigned at the generic level only, and another 11 are questionably assigned to genera or species. The zonation proposed includes 6 zones and 3 levels, including the Haydenaspis parvatya level, the Oryctocephalus indicus level, the Kaotaia prachina Zone, the Paramecephalus defossus Zone, the Oryctocephalus salteri Zone, the Iranoleesia butes level, the Sudanomocarina sinindica Zone, the Lejopyge acantha Zone, and the Proagnostus bulbus Zone. The sections span from the upper part of the informal Stage 4, Series 2 of the Cambrian System, about 511 Ma old, to the Proagnostus bulbus zone of the Guzhangian Stage near the top of Series 3, dated at about 501 Ma. This time interval is represented by about 2000 m of section, which is thick compared to similar intervals elsewhere and is consistent with high rates of sedimentation along the Himalayan margin at the time. The fauna resembles others from equatorial peri-Gondwanaland, with closest similarity to that of South China. It also bears strong affinity to the North China fauna. Juvenile trilobites are described for the first time from India. A new Chinese species, Monanocephalus liquani, is also described.

New discoveries of the early Cambrian yiliangellinine trilobite Zhangshania typica Li and Zhang in Kunming preserve almost all instars from early postembryonic (protaspid) to mature (holaspid) phases in articulated state, in addition to mature specimens with antennae bearing paired spines on the basal articles. The ontogenetic series shows protarthrous development with some, but likely not all, early holaspid instars expressing additional pygidial segments, gradual rearward migration of the location of the longest pleural spines on the trunk segments, and striking positive allometry of the genal spines. It also reveals Parazhangshania sichuanensis Li and Zhang, 1990 to be the holaspid stage 3 of Z. typica, and therefore its junior synonym. This new find in the Hongjingshao Formation provides species-based regional correlation across the South China block and Z. typica may provide an important biostratigraphic marker for the base of the traditional Tsanglangpuan Stage.

Cambrian biostratigraphy of the Indian subcontinent is best documented from the Parahio Formation of the Tethyan Himalaya. Recently established trilobite biostratigraphy shows that the formation encompasses the latest part of unnamed Stage 4 and much of unnamed Stage 5. A variety of small shelly fossils have been recovered via acid digestion of carbonate beds and include tetract and pentact hexactinellid sponge spicules, chancelloriid spicules belonging to Chancelloria sp. and a new species, Archiasterella dhiraji, shells of an helcionelloid comparable to Igorella maidipingensis, a meraspid ptychopariid trilobite, the tubular Cupitheca sp., a poorly preserved hyolith, and an assortment of spinose microfossils of uncertain affinity. These newly recovered microfossils are consistent with the trilobite-based lower and middle Cambrian age determination and do not support a late Cambrian age for the top of the Parahio Formation advocated in some recent literature. The microfossils reported herein significantly expand the known diversity of such fossils from Cambrian strata in the Himalayan region, and allow for comparison of this fauna with others from Gondwanaland and elsewhere. Integration with trilobite data indicate that the stratigraphic ranges of many small shelly fossils described in this study are greater than previously recognized.

Several localities within the heterolithic facies of the St. Lawrence Formation (Upper Cambrian) of Wisconsin and Minnesota yield specimens with phosphatic exoskeletons, quadrate cross sections composed of four equidimensional faces each bearing a midline, and possible holdfast attachment during life. These specimens are here referred to the order Conulariida, class Scyphozoa. Their fine, tuberculate surface ornament and serially invaginated midline structure serve to define a new genus, Baccaconularia, to which two new species, B. robinsoni and B. meyeri, are assigned. Conularia cambria Walcott 1890, also from the Cambrian of the northern Mississippi Valley and long dismissed as a misidentified trilobite fragment, is illustrated photographically for the first time. This species occurs in rocks stratigraphically beneath the St. Lawrence Formation. Specimens assigned to this species by Walcott are conulariids, but lack features now considered diagnostic of either Conularia or Baccaconularia. Walcott's material is insufficient to permit detailed taxonomic evaluation, and we isolate this name to this material, pending the collection of additional, better preserved specimens. Together, Baccaconularia and Conularia cambria contain the oldest large conulariids, and these narrow a stratigraphic gap between other large conulariids known from the Lower Ordovician onwards, and smaller fossils with conulariid affinities known only from Lower Cambrian rocks.

A new, low-density trilobite body cluster from an Upper Ordovician Kope Formation mudstone in the Cincinnati region is characterized by well-preserved, articulated specimens. This cluster is polytaxic, size-segregated, and shows an unusual variety of flexure postures in the most common taxon, Flexicalymene aff. granulosa. Specimens show internal pyritization and associated pyritefilled tubular burrows, and were preserved by rapid deposition of mud from a sediment-laden flow. Size-segregation among species, but size-consistency within species, coupled with sedimentological data and apparently haphazard body orientation data, suggest that the trilobites represent a biological association, rather than the product of mechanical sorting. While the majority of specimens are inverted, many are dorsal-up, and some are inclined to bedding. Postures include prone (i.e., nonflexed), simple dorsal or ventral flexure, and complex stances that combine both dorsal and ventral flexure. Flexure pattern appears largely unrelated to sediment compaction. We interpret the cluster as an association of live trilobites that were entombed in situ. Specimens may have inverted prior to burial as a behavioral posture, or as a response to onset of burial. The animals were likely preserved in the process of escaping, as they tried to free themselves from being buried. This is in contrast to the tightly enrolled stance in some trilobites, especially other Flexicalymene, commonly associated with rapid burial events, and indicates a new behavioral aspect of trilobite paleobiology. The distribution of associated burrows suggests that the rotting carcasses attracted scavengers.

It has been proposed previously that the northern Mississippi Valley Sunwaptan trilobite genus Dikelocephalus comprises 26 species. Morphometric analyses demonstrate that many of the criteria that had been used to define species of Dikelocephalus are invalid and additional analysis of biostratigraphically and biogeographically constrained collections is necessary before the taxonomic status of Dikelocephalus can be fully resolved. Our results indicate that infrageneric morphological variation in Dikelocephalus is continuous and lacks gaps that could be the basis for establishing multiple species. Many of the characters shown to be taxonomically insignificant in Dikelocephalus are also widely used in the definition of other trilobite taxa. This suggests that the species-level taxonomy of many trilobites may be substantially oversplit. Recognition of widespread oversplitting will have important consequences for biostratigraphic zonations, paleogeographic distributions, and estimates of taxonomic diversity.

Rare specimens of the Late Cambrian primitive libristomate trilobite “Cedaria” woosteri from the Eau Claire Formation of north-central Wisconsin preserve an unusual and new ocular suture, and the impressions of lens facets. The suture is circumocular, but loops abaxially onto the ocular platform. The portion of the ocular platform that is enclosed by this suture and is fused with the visual surface is here termed the “ocular plate”. The size and arrangement of the lens facets show that “C.” woosteri possessed a holochroal eye, typical of that known in other Cambrian libristomate trilobites. This nearshore assemblage, while low in diversity, contains a disparate array of trilobite morphotypes.