A quantitative comparison of the ontogeny of two closely-related Upper Devonian phacopid trilobites

The best insight into the development of Devonian phacopids has been obtained from Trimerocephalus lelievrei Crônier & Feist, 1997, a Famennian phacopine from Morocco, where changes in size and shape have been quantified. In this study, a morphometric approach has been used: (1) to retrodeform and then establish patterns of morphological variation in a well preserved but tectonically deformed assemblage belonging to another phacopine species Weyerites ensae (Richter & Richter, 1926), a Famennian phacopine from Thuringia, and (2) to establish patterns of developmental and evolutionary changes within two closely related species: Weyerites ensae and Trimerocephalus lelievrei. The method of retrodeformation using a set of discrete points presumed to be homologous on all studied individuals, has demonstrated that the next analyses are possible on the retrodeformed material as compared to the undeformed material. Morphometric analysis based on outline analysis has permitted demonstration of progressive shape change in agreement with ontogenetic ordination and a comparison of changes in size and shape in Weyerites ensae. The main changes in shape appear to occur in the meraspid period, whereas increase in size takes place mainly in the holaspid period. This pattern, already reported for Trimerocephalus lelievrei, can be generalized for phacopine trilobites from the Late Devonian. Moreover, the comparison of the two ontogenetic trajectories has shown that most of the differences are related to 'structural' changes, probably linked to a relative pre- post-displacement. The results suggest that ecological adaptation may be studied by examining the changes in development that occur within species through time and space.     

ONTOGENY AND HETEROCHRONY IN THE EARLY CAMBRIAN ORYCTOCEPHALID TRILOBITES CHANGASPIS, DUYUNASPIS AND BALANGIA FROM CHINA

Abstract:  Ontogenies are described for the first time for three species of Early Cambrian oryctocephalid trilobites: the oryctocephaline Changaspis elongata Lee, in Chien and the oryctocarines Duyunaspis duyunensis Chang and Chien, in Zhou et al . and Balangia balangensis Chien from the Balang Formation in eastern Ghuizhou Province, south-west China. The complete protaspid to holaspid ontogeny for Duyunaspis duyunensis and incomplete meraspid to holaspid ontogenies for Changaspis elongata and Balangia balangensis are described. The relative degree of ontogenetic variation within the lineage Duyunaspis duyunensis – Balangia balangensis is analysed and indicates that Balangia may have evolved from Duyunaspis by paedomorphosis. Decrease in thoracic segment number and concomitant increase in pygidial segment number in the later genus mirrors the pattern seen in the evolution of another oryctocephalid genus, Arthricocephalus . In both cases, selection was probably targeting an increase in cephalic convexity. This resulted in the evolution of a larger pygidium due to increased elevation of the axis above the substrate. The ontogenetic development of Changaspis provides evidence supporting the view that the Oryctocarinae may be considered as paedomorphic descendants of the Oryctocephalinae.     

Decoupling of local and regional dominance in trilobite assemblages from northwestern Argentina: new insights into Cambro‐Ordovician ecological changes

Components of biodiversity are strongly scale dependent, but the relative importance of the patterns that operate at different scales and the links between them have been overlooked. To disentangle the ecological structure of Cambro‐Ordovician trilobite assemblages from the Argentine Cordillera Oriental at different scales, we explore patterns of abundance, dominance and occupancy across the onshore–offshore profile, and through three time intervals: Furongian, earliest Late Tremadocian (Tr2), latest Middle Floian–earliest Late Floian (Fl2–Fl3). At the regional scale, single taxa are overwhelming dominant in the Furongian (Parabolina) and in the earliest Late Tremadocian (Leptoplastides). Several dominants occur in the Floian, but just one (Famatinolithus) attains high occupancy and, rarely, high dominance. In contrast, only the Furongian records highly dominated local assemblages, whereas dominance distinctly decreases among Tr2 and Fl2–Fl3 ones. Thus, when both scales of analysis are combined, an unexpected scenario becomes evident: Tr2 assemblages resemble those of the Furongian at the regional scale, but mirror those of the Floian at the local scale. These results highlight a decoupling in local versus regional structures triggered by an earlier switch in dominance in local communities and a delayed change at the regional scale. Interestingly, this decrease in local dominance matches previous analyses accounting for a coeval step‐up in local evenness, suggesting that the Tr2 appears as a pivotal interval in the reorganization of communities in the Cordillera Oriental. This scenario emphasizes that biogeographical regions witnessed different regional‐scale processes, and suggests that scaling local and regional patterns provides new insights to unravel the history of biodiversity among benthic communities.     

Punctuated equilibrium excused: the original examples fail to support it

The four original stipulations of the definition of punctuated equilibrium (PE; Eldredge & Gould, 1972) are shown to be unsupported, and even contradicted, by the two evidential examples these authors supplied: the trilobite Phacops rana of the Middle Devonian, and the land snail Poecilozonites bermudensis of Late Pleistocene Bermuda.
The additional, much-discussed example of Bellamya unicolor and the accompanying suite of molluscan species of Pliocene-Recent African Lake Turkana, studied by Williamson, also fails to exemplify PE. In particular, the data produced for these cases appear to represent counterexamples to Mayr's paradigm of peripatric speciation , embraced by Eldredge and Gould as the central effective mechanism of PE. The data actually illustrate instead a prevailingly centrifugal pattern of speciation that will accommodate and explain episodic aspects of microevolution in a more realistic way.     

The North China Steptoean positive carbon isotope excursion and its global correlation with the base of the Paibian Stage (early Furongian Series), Cambrian

The use of carbon isotope excursion in Cambrian stratigraphical correlation is a standard practice at both the intercontinental and intracontinental scales. The Steptoean positive carbon isotope excursion (SPICE) is one of the prime examples in this regard in correlating the base of the Paibian Stage and Furongian Series. A lack of definite SPICE evidence in the North China craton has been a challenge in precision correlation between North China and other palaeo‐continents. This study provides new carbonates carbon isotope data from the type locality of the Changshan Formation in Hebei Province, North China. Our new δ13C_carb data provide new objective evidence for the presence of the SPICE in North China. The sampling section is relatively condensed, and the interval of the SPICE curve is less than one and half‐metres after analysing 64 samples (the sampling interval within the SPICE is less than 10 cm). The onset of the SPICE curve in Tangshan, Hebei, occurs in the barren interval between the Neodrepanura and Chuangia trilobite zones. Based on this study and previous work, this could imply the middle part of the Prochuangia‐Paracoosia trilobite Zone in North China and can be correlated with the base of the Paibian Stage and Furongian Series.     

Silicified Middle Cambrian trilobites from Kyrgyzstan

Abstract:  A medium diversity silicified Middle Cambrian faunal assemblage occurs in an olistolith of bioclastic limestone within Silurian siliciclastic turbidites on the slopes of Arpatektyr Mountain in the eastern Alai Range, Kyrgyzstan. The fauna includes the trilobites Raragnostus cf. mirus , Altikolia kurshabica sp. nov., Corynexochina weberi , Dorypyge richthofeniformis , aff. Jincella sp., Olenoides comptus , Skreiaspis aff. spinosus and Suluktella mambetovi sp. nov. Three of these taxa ( C .  weberi , O .  comptus and R . cf.  mirus ) are considered locally to be characteristic of the Pseudanomocarina Beds, although Pseudanomocarina itself is not present. Altikolia and Suluktella are endemic to early Palaeozoic terranes of Central Asia, Corynexochina also occurs in South China and Skreiaspis is known elsewhere exclusively from the Middle Cambrian of 'West' Gondwana. Because of preservation in full relief, the silicified trilobite fauna allows a detailed understanding of some morphological structures, related especially to trilobite enrolment, otherwise inadequately known for many Mid-Cambrian polymerid trilobites.     

Cambrian high-resolution biostratigraphy and carbon isotope chemostratigraphy in Scania, Sweden: first record of the SPICE and DICE excursions in Scandinavia

A core drilling (Andrarum‐3), from the classical locality at Andrarum, Scania, southernmost Sweden, penetrated a 28.90‐m‐thick Cambrian succession. The core comprises dark grey to black, finely laminated mudstones and shales with early concretionary carbonate lenses (stinkstones or orsten) and a few primary carbonate beds. The middle Cambrian (provisional Series 3) part of the core comprises 17.35 m, whereas the Furongian Series (upper Cambrian) part covers the remaining 11.55 m. Nineteen trilobite and two phosphatocopine genera are present in the middle Cambrian, whereas the less diverse Furongian interval yielded four trilobite and three phosphatocopine genera. Other, less frequent, faunal elements include conodonts (s. l.), brachiopods, sponge spicules, bradoriids, and coprolites. Trilobites and phosphatocopines were used to subdivide the core into seven biozones ranging from the Ptychagnostus atavus Zone to the Parabolina spinulosa Zone (P. spinulosa Subzone). Carbon isotopic analyses (δ¹³C_org) through the core show two important excursions, the negative DrumIan Carbon isotope Excursion (DICE) in the Pt. atavus Zone, and the Steptoean Positive Carbon Isotope Excursion (SPICE) beginning near the first appearance of Glyptagnostus reticulatus and extending upward into the Olenus and Agnostus (Homagnostus) obesus Zone. The DICE displays a peak value, in the samples at hand, of –30.45‰δ¹³C_org in the lower part of the P. atavus Zone. The δ¹³C_org values increase through the overlying L. laevigata and A. pisiformis zones and display peak values of c. –28.00‰δ¹³C_org in the lowermost Furongian Olenus wahlenbergi and O. attenuatus subzones. Thereafter the values decrease significantly through the O. scanicus Subzone. Both isotopic excursions have been documented from several palaeocontinents, but never before from Baltica. Moreover, for the first time these excursions are recorded from organic matter in an alum shale setting. The recorded shift of +1.50–2.00‰δ¹³C_org is approximately half the magnitude of the SPICE documented from other regions. This discrepancy may be related to temporal variations in the type, origin, or diagenesis of the organic fraction analysed.     

Biomechanical analyses of Cambrian euarthropod limbs reveal their effectiveness in mastication and durophagy

Durophagy arose in the Cambrian and greatly influenced the diversification of biomineralized defensive structures throughout the Phanerozoic. Spinose gnathobases on protopodites of Cambrian euarthropod limbs are considered key innovations for shell-crushing, yet few studies have demonstrated their effectiveness with biomechanical models. Here we present finite-element analysis models of two Cambrian trilobites with prominent gnathobases—Redlichia rex and Olenoides serratus—and compare these to the protopodites of the Cambrian euarthropod Sidneyia inexpectans and the modern American horseshoe crab, Limulus polyphemus. Results show that L. polyphemus, S. inexpectans and R. rex have broadly similar microstrain patterns, reflecting effective durophagous abilities. Conversely, low microstrain values across the O. serratus protopodite suggest that the elongate gnathobasic spines transferred minimal strain, implying that this species was less well-adapted to masticate hard prey. These results confirm that Cambrian euarthropods with transversely elongate protopodites bearing short, robust gnathobasic spines were likely durophages. Comparatively, taxa with shorter protopodites armed with long spines, such as O. serratus, were more likely restricted to a soft food diet. The prevalence of Cambrian gnathobase-bearing euarthropods and their various feeding specializations may have accelerated the development of complex trophic relationships within early animal ecosystems, especially the ‘arms race'' between predators and biomineralized prey.     

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.     

Ecological disruption close to the Ordovician‐Silurian boundary

Following a long period of stability there was substantial ecological disruption in the late Ordovician, accompanied by widespread extinction. The ecological upheaval was spread over a period of time (1 My?) and varied in intensity between different groups.

The greatest change in brachiopod faunas occurred in the tropics, where there were widespread extinctions. In temperate regions the community structure, although disrupted, was not destroyed. The deep shelf associations (Benthic Assemblages 4–5) of the Rawtheyan persisted in an attenuated form into the Hirnantian when they were supplemented by genera of the ”Hirnantia fauna” and then became strongly re‐established in the Llandovery.

Trilobites, in contrast to brachiopods, suffered a worldwide extinction in the late Ashgill. Approximately 40% of trilobite genera became extinct at, or near, the Rawtheyan‐Hirnantian boundary.

Analysis of the distribution of Rawtheyan trilobite and brachiopod genera supports a correlation between geographical distribution and survival. Global range charts for brachiopod and trilobite genera, incorporating ecological and palaeogeographical data, have been compiled for this study.