Localities bearing abundant enrolled Cambrian trilobites are extremely rare, but here we report a new and exceptional such assemblage with a high diversity of enrolled specimens from the middle Cambrian of Spain. This novel fi nd demonstrates that encapsulated enrollment was a behavioral response available to many of the early benthic trilobite morphotypes. This result provides a new perspective for reviewing the evolution of trilobite enrollment through the Paleozoic. We fi nd a signifi cant and consistent increase in the proportion of enrolled trilobites within assemblages bearing abundant articulated trilobites. Similarly, the mean number of coaptative devices also increased progressively during the Paleozoic. Such devices likely provided later enrolled trilobites with greater structural integrity, and may support an escalatory model of trilobite defense evolution. Likewise, environmental circumstances favoring the preservation of enrolled trilobites may have become more common during post-Cambrian time.on June 4, 2015 geology.gsapubs.org Downloaded from

Laurentia, the Oryctocephalus indicus Zone in the Indian Himalaya and North Greenland, near the base of the Delamaran Stage in Australia, and within the Eccaparadocides sdzuyi Zone in Iberia and the Ornamentaspis frequens Zone in Morocco.

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.

From an epidemiological and public health perspective there is an interest in quantifying differences in incidence and mortality between either time points, geographical areas or males and females. We propose a method for splitting such a difference in the number of cases/deaths into three components: (1) those due to risk; (2) those due to population structure (i.e. age distribution); and (3) those due to population size. We also propose graphical methods for presenting the results. Three examples are used to illustrate our methodology.

Oryctocephalus indicus is a widespread trilobite species described from the lower–middle Cambrian (provisional Cambrian Stage 5) of south China, North Korea, India, USA and possibly North Greenland. Principal component analysis (PCA) and thin‐plate splines (TPSs) based on a pooled sample of 359 specimens from four sections in the Guizhou Province, south China, demonstrated that the shape of this species is very constant, and only a small difference is detected in specimens from deeper part of the basin. Most of the variation is taphonomically controlled, and only the specimens from Sanwan Section show a modest natural difference due to a slightly different depositional environment. A comparison between the Chinese specimens and Oryctocephalus ‘reticulatus’ from the Molodo Section, Siberia, has been carried out. PCA and TPS show that O. ‘reticulatus occupies the same morphospace of O. indicus. However, the canonical variates analysis and the analysis of similarities detect small differences between the Siberian and Chinese populations, indicating a minor geographical variation caused by different environmental settings. The ontogenic variation is also described from early meraspids to holaspids. The most substantial changes take place during the first period of the early meraspid development, in which the glabella changes from a bell‐like shape to a cylindrical shape. The ocular ridge changes from poorly defined in early meraspid to well developed in late merapids. The results indicate that O. ‘reticulatus’ is a junior synonym of O. indicus, suggesting that the base O. indicus of Siberia can be correlated with the O. indicus Zone of south China.

Fifty specimens belonging to species of Solenopleuropsis and Pardailhania from Spain and France demonstrate sphaeroidal enrolment in Cambrian trilobites for the first time. These solenopleuropsines show novel coaptative structures in different regions of the exoskeleton: in the cephalon there are vincular furrows and notches; in the thorax an articulating facet is developed at the pleural margins, with a ball and socket connection on the adaxial most portion, and an articulating half-ring axially; the pygidium possesses an articulating facet. The interaction of these coaptative structures resulted in a sphaeroidal enrolment that was a progressive act from the first articulation between the occipital ring and the first segment to the pygidial articulating facet. A similar type of sphaeroidal enrolment is observed in the Devonian trilobite Phacops. Both Cambrian and Devonian trilobites developed a vincular furrow in the ventral surface of the cephalon to close their bodies tightly. In both cases, this is probably a convergent adaptation to protect against predators and obrution. Indeed, the enrolled trilobites are very common in obrution deposits restricted to shallow and soft muddy substrates.

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Morphology permits the extracting of information to study patterns of disparity and diversity of a particular group of animals through time. Enrolment is a characteristic behaviour of trilobites, having been first recorded in the Cambrian and continued until their demise at the end of Permian mass extinction. Morphological trends related to enrolment strategies have been documented previously but are yet to have been analysed in a morphospace framework. Here we analyse the cephalic shape of a wide range of trilobite taxa using geometric morphometrics. These methodologies were implemented to explore whether the relationship between the shape of the head and the evolution of different enrolment styles correspond with an increase in morphospace or disparity during the Palaeozoic. Our results show distinct patterns of diversity and disparity in the functional morphology of trilobites during the Palaeozoic. Furthermore, these analyses show that enrolment types in trilobites without interlocking devices (e.g. cylindrical) permit more flexibility in the head shape, promoting higher rates of disparity, possibly leading to an enhanced ability to occupy a broad range of ecological niches. On the contrary, lower disparity cephalic shape constrains the way to enclose the body and consequently places limitations on the ability to adapt to new niches. Our results indicate that the higher rates of morpho-functional diversity and disparity coincide with the increase of morphospace in radiations in the early Palaeozoic (Cambrian and Ordovician) but also immediately after the Carboniferous-Permian biodiversification event. Low disparity rates constrain the success of trilobites with a specific enrolment type. Finally, the results show that sphaeroidal enrolment style was the most successful type that evolved independently in all trilobite orders. □ Arthropods, ecology, ecomorphology, evolution, functional morphology.