Changes in the palaeoenvironment are investigated in two representative Frasnian sections of the NW Algerian Sahara, integrating sedimentology and magnetic susceptibility (MS). The Ben Zireg section is characterized by condensed and ferruginous calcareous deposits; in the South Marhouma section the sedimentation rate is high, dominated by muddy nodular limestones with several hypoxic shale intervals. In both sections, sediments were mostly emplaced on pelagic outer ramps below the limit of storm wave-base, evolving through time from proximal to distal setting. Investigations of the temporal evolution of facies and MS data permit a first estimate of the local sea-level trends in NW Algeria. These trends match the overall long-term rise of sea level recognized worldwide from Frasnian Zone 5 upwards. Noteable positive excursions of the sea-level curve related to the semichatovae transgression, as well as to the late Frasnian transgression prior to the late Kellwasser event, can be established in this area. Although the sharp regression of sea level at the upper Kellwasser level can be confirmed from our data, no particular trend is depicted at the transition of conodont zones (Frasnian Zones 12–13) where the presence of the lower Kellwasser level has not yet been clearly recognized.

The recovery of well-preserved silicified larvae from the early Tournaisian of Montagne Noire, southern France, allows the first description of the early ontogeny of proetoid trilobites, survivors of the end-Devonian Hangenberg extinction event. The fauna comprises various taxa that can only partly and tentatively be assigned to the genera Liobolina, Diacoryphe, and Pedinocoryphe without specific attribution. The close resemblance of these larvae to previously described proetoid larvae from the Ordovician and Devonian emphasizes the ontogenetic homogeneity of this superfamily. Moreover, it supports the view that the Proetida is composed of two clades: the Aulacopleuroidea/Bathyuroidea group and the Proetoidea. The Carboniferous anaprotaspides exhibit an unusually wide size-range that may provide evidence of the acquisition of an extended planktonic period in the early ontogeny of at least some Carboniferous proetoids. This change in the ontogenetic strategy may have enabled proetoid trilobites to survive during the end-Devonian biocrises.

A review of all known species of the scutelluine trilobite Paralejurus corroborates its distinction from contemporaneous representatives of the subfamily. The overall morphology together with observed habitat conditions favor the view of its having had a semiendobenthic life habit analogous to that of many earlier illaenids. Particular features that are shared with the Silurian Rhaxeros, such as the pronounced convexity of the exoskeleton and the wide axis, are considered as homeomorphies related to a burrowing lifestyle rather than the expression of phyletic relationships. The data set based on conodont biostratigraphy reveals earliest occurrences of Paralejurus in the Pragian and the disappearance of last representatives at the base of the Middle Devonian. In the lower Early Devonian Paralejurus was restricted to the North Gondwana margin and related microplates. Later and in consequence of geodynamic plate movements that led to the reduction of oceanic barriers, the Paralejurus dormitzeri group succeeded in spreading to the epicontinental margin of Laurussia in Late Emsian times. New material from Southern Morocco, including Paralejurus spatuliformis n. sp. and Paralejurus tenuistriatus n. sp., and the first representative of the genus from Spain, Paralejurus carlsi n. sp., are described. Paralejurus dormitzeri rehamnanus Alberti, 1970 is considered an independent species emphasized by the discovery of complete exoskeletons.

Two new species of scutelluine styginid trilobites are described from Frasnian strata in the Virgin Hills Formation in the Canning Basin of Western Australia. They are placed in the genus Telopeltis n. gen., reflecting the fact that their final occurrence in the late Frasnian, up to the latest Frasnian Kellwasser extinction event, is the last known record of scutelluine trilobites. As such, it also represents the youngest record of the order Corynexochida. The two species, Telopeltis woodwardi n. sp. and Telopeltis microphthalmus n. sp., are unlike most other scutelluines in possessing extremely vaulted pygidia and showing trends to eye reduction. Such eye reduction is a common feature of late Frasnian trilobites. The characteristic morphological features of this small genus are indicative of evolution of this last scutelluine by paedomorphosis.

A detailed morphometric approach based on size and on outline analyses has been used on an exceptionally well-preserved assemblage of silicified trilobite exuvia, recovered from a Late Devonian limestone from southeastern Morocco. The material comprises a series of late larval to postlarval growth stages belonging to a single phacopine species, Trimerocephalus lelievrei Crônier and Feist, 1997.

Plurimodality of size distribution has allowed us to discriminate postlarval instars. Distinct dimensional classes of isolated parts are obtained using the intertooth distances on the posterior pygidial margin and the internotch distances in the cephalic vincular furrow, which are functionally linked during trilobite enrollment. Morphometric analysis of development permitted demonstration of progressive shape change in agreement with ontogenetic ordination and a comparison of the timing of size and shape changes. The main shape changes appear to occur early in development, and once the “adult” morphology is obtained, size increases significantly. The growth rate during ontogeny is estimated by analogy with extant deep-sea crustaceans. Exponential size increase resulting from constant duration of intermolt periods may be regarded as a life history strategy to compete in a nutrient-impoverished offshore environment. The particular phacopine mode of molting, which involves the opening of the neck joint after ankylosis of the facial sutures, occurred in Trimerocephalus lelievrei between the first two postlarval instars, later than in its ancestor. Trimerocephalus lelievrei occupies an intermediate position within the phacopine paedomorphocline as indicated by the delayed onset of ankylosis.

Distributional patterns of eye lens variation in different morphs of the phacopine trilobite Acuticryphops acuticeps (Kayser 1889) are investigated. The specimens were collected from the latest Frasnian preceding the Upper Kellwasser (UKW) global extinction event in the Frasnian-Famennian (Late Devonian) stratotype section at Coumiac, southern France. In six successive populations a gradual reduction in the mean number of lenses occurs within the short time span of a single conodont Zone. This morphological change cannot be imputed either to the size of individual specimens or to variation in cephalic morphology. Thus morphs with different numbers of eye lenses are considered intraspecific. However, the intrapopulational percentage relation between morphs does not remain constant, as the coefficient of variation in lens number continuously increases from one population to the next. Cases of individuals with asymmetric eyes appear in the two latest assemblages prior to the UKW level. The fact that the total variation of the shape of the cephalon is not affected in these assemblages suggests that the morphological changes observed in the visual complex may account for the relaxation of selective pressures on this trait. Such a change in the regime of selection would have been accompanied by a lessening of the processes that control the development of this trait. As the phenomenon of eye reduction is not restrained by local conditions at Coumiac (Montagne Noire) but occurs contemporaneously to various extents in other crustal blocks such as Rhenish Slate Mountains (Avalonia), Thuringia (Armorica), and Morocco (Northern Gondwana), it is considered as constituting an adaptation to global eustatic deepening that occurred in the terminal Frasnian just before the global extinction event.

Twenty-seven graptolite species are reported from the lowermost Silurian (Rhuddanian) ascensus-acuminatus and vesiculosus biozones of Montagne Noire in southwestern France. Four species are left in open nomenclature, four are new: Normalograptus crassus, Neodiplograptus lautus, Dimorphograptus compactus and Huttagraptus solidus. The joint occurrence of Akidograptus ascensus, Parakidograptus acuminatus and Cystograptus ancestralis, recorded in the Gabian-1 section, suggests correlation of the whole succession at this site with the middle part of a combined ascensus-acuminatus Biozone. Revision of museum collections revealed that other elements that are characteristic of the ascensus-acuminatus Biozone assemblage in peri-Gondwanan Europe, such as Normalograptus trifilis, N. longifilis and Neodiplograptus lanceolatus, have been recovered from several other localities in Montagne Noire that are no longer accessible. The graptolite assemblage of the Gabian-2 section, with common Cystograptus vesiculosus, Dimorphograptus extenuatus, D. elongatus and Atavograptus atavus, belongs to the lower part of the vesiculosus Biozone, whilst the appearance of D. swanstoni and D.? physophora in the upper part of the section represents a higher, although not the uppermost, part of the biozone. This is the first record of a diverse dimorphograptid fauna within the vesiculosus Biozone in the Gondwanan part of Europe.

An account of the systematics, biostratigraphy, ecological setting and the biogeographic implications of rugose corals from olistoliths of the Cabrières area is given. Corals of the lower member of the Izarne Formation are dated by conodonts as gronbergi Zone age, and include Lythophyllum sp. indet., Calceola sandalina (Linné), Tryplasma sp. A, Breviphrentis roharti Pedder new species, B. exigua Pedder new species, and Aqishaphyllum sp. A. Corals from the middle member of the formation include Frechocystis pertinax Pedder new genus and species, Calceola sp. undet., Rhizophyllum sp. aff. R. ukalundense Hill and Jell, Tryplasma enorme Pedder new species, Tryplasma sp. A, Breviphrentis sp. A, Platysmatophyllum halleri Pedder new genus and species, Pseudochonophyllum sentum Pedder new species, and Izarneophyllum barroisi (Frech) new genus. No age significant conodont has been recovered from the middle member. However, scutelluid trilobites, which, together with other trilobites, evidently used the underside of Izarne corals for shelter during molting, provide correlation with conodont sequences in the nappe domain to the north and southwest of Cabrières. From this line of evidence, the middle Izarne coral fauna is deduced to be nothoperbonus Zone age. The association of a variety of benthic trilobites, all with large eyes, provides evidence of a photic zone environment for the middle Izarne corals.

Rugose corals from the Izarne Formation belong to the Old World Realm and have nothing in common with similar age Rugosa of the Eastern Americas Realm. This implies that the dissolution of the boundary between these realms, which occurred in the Middle Devonian, did not begin before latest Emsian time.

One of the most severe restrictions in taxonomic diversity of trilobites coincides with the Late Devonian “Hangenberg” Event, which is characterized by a pronounced eustatic low-stand. One of the survivors of this event is Pudoproetus, which, being adapted to shallow water conditions, experienced an opportunistic worldwide spread during this time. The oldest representatives of this genus are reported from the pre-Hangenberg Late Famennian of the Urals and Kazakhstan. Other known occurrences of this genus, midcontinent U.S.A., Utah, southern China, and southern France, occur at the Devonian-Carboniferous boundary, shortly after the Hangenberg event, and in the earliest Carboniferous. This pattern of distribution suggests that neither climatic nor geographic migration barriers existed around the Old Red landmass at that time. On the other hand, the Hangenberg crisis severely affected outer shelf benthic habitats, whereas nearshore trilobites were largely unaffected by the event.

A proetine origin of Pudoproetus, as originally proposed by Hessler (1963), is emphasized on the basis of paleoecologic, stratigraphic, and morphological arguments. In particular, the presence of ancestral features, such as the possession of 10 thoracic segments and lateral occipital lobes still occurring in early Pudoproetus species, points to a phylogenetic link with proetines rather than with phillipsiids. Within this group two new species are named: P. priscus and P. mediterraneus.

The Ben Zireg anticline NW of Bechar yields the currently most-important Frasnian succession on the northern margin of the Algerian Sahara. It represents 26.5 m of calcilutites which are attributed to the middle–late Frasnian. As the early Frasnian is not represented, the succession rests conformably on undifferentiated, probably late Givetian, substrate. Fine-scaled conodont biostratigraphy reveals a continuous sequence of Montagne Noire Zones 5–13, superseded by the earliest Famennian Lower triangularis Zone. The deposits are organized into 15 sedimentary cycles that are initially condensed and become dilated upwards. Conodont biofacies indicate an overall deepening towards the top of the section with an episode of slight shallowing at the MN11/12 transition marked by an increase of ancyrodellids, which is also seen in the Marhouma section of the Ougarta region (SW Algeria). Homogeneous calcareous microfacies with only a few shaly intercalations determine a predominantly oxygenated depositional environment on an outer platform, submarine rise or ramp setting. This is also indicated by relatively low sedimentation rates which are similar in selected sections from the Tafilalt platform, and contrast with those from the Marhouma trough and the Maider basin. Unlike other Frasnian successions south of the Atlas Fault, but similar to the Moroccan Meseta, the Upper Kellwasser horizon is clearly discriminated at Ben Zireg by an outstanding occurrence of black shales on top of the oxygenated latest Frasnian deposits.

We present detailed biostratigraphy based on conodonts and palaeoenvironmental trends deduced from microfacies and conodont abundance through the Famennian (Late Devonian) at Col des Tribes (Montagne Noire, France). The succession is characterized by micritic limestones deposited in settings oscillating between mid to outer ramp. Facies contain poor fauna, widely dominated by nektonic organisms. This section is complete and one of the most conodont-rich for the Famennian of the north Gondwana-related area. The Upper Kellwasser event (Frasnian–Famennian boundary) and the Hangenberg (Devonian–Carboniferous boundary) have been lithologically identified. They are characterized by decimetre-thick black dysoxic to anoxic argillaceous sediments. The Condroz and annulata events, although not materialized by lithological changes, have been positioned due to the precise stratigraphy. The first event occurred during the deposition of condensed ferruginous facies (griotte limestones) and the second event during the deposition of micrites barren of benthic fauna. The combination of information from both facies and conodont biofacies changes allows a general sea-level curve through the entire Famennian for north Gondwana to be proposed for the first time. At Col des Tribes, the general trend is a slight deepening upwards from triangularis to trachytera zones, then a pronounced shallowing-upwards trend from upper trachytera to praesulcata zones. This curve correlates with the well-known reference curve from Euramerica concerning the late Famennian (trachytera to praesulcata Zones). There are some discrepancies in minor cycles which can be explained by tectonical phenomena at the onset of the edification of the Variscan belt in Europe.

A diverse fauna of phacopid trilobites is described from the Late Devonian (middle Frasnian to early Famennian) of the northern Canning Basin, Western Australia. One new genus and four species in two genera are described from zones 11, 13a and 13b of the middle and late Frasnian: Trimerocephaloides sinevisus gen. nov. and sp. nov., T. ? linguiformis sp. nov., Acuticryphops acuticeps (Kayser, 1889) and A. klapperi sp. nov. Late Frasnian phacopines are either blind, as shown for the first time in Trimerocephaloides sinevisus, or show trends to decreasing eye size up to the Frasnian–Famennian ‘Kellwasser’ mass extinction event. This evolutionary trend in Acuticryphops is demonstrated to have been global at this time. One new genus and six species of early Famennian phacopids are described, from the Upper triangularis, crepida and rhomboidea zones: Houseops gen. nov. with the new taxa H. canningensis sp. nov., H. beckeri sp. nov. and H. sp. A, Babinops planiventer Feist & Becker, 1997, B. minor sp. nov., Trimerocephalus tardispinosus Feist & Becker, 1997 and T. mimbi sp. nov. In contrast to European sections where exclusively blind phacopids are known in earliest Famennian sites, initial recovery following the mass extinction event in Canning peri-reefal environments is characterized by oculated forms. These trilobites must have evolved from conservative ancestors with normal eyes that had succeeded in surviving the Kellwasser biocrises in reef-related shallow water niches. Thus the origin of post-event phacopids from shallow water environments is demonstrated for the first time. Descendant lineages show increasing eye size, increased cephalic vaulting and effacement during the early Famennian.

A new otarionine trilobite Otarionella rastrum sp. nov., from the late Emsian Mont Peyroux Formation (Montagne Noire, France), is described. Silicified remains, recovered from the underlying Bissounel Formation (early to late Emsian), are also attributed to this new species. These isolated silicified sclerites represent metaprotaspid to young holaspid growth stages, which enables the almost complete ontogeny of an otarionine trilobite with a spinose adult morphology to be described for the first time. Comparison with associated larval and juvenile growth stages of Cyphaspis reveal that the pattern of juvenile cranidial spine distribution in Otarionella rastrum sp. nov. differs from all patterns described so far in the Otarioninae, in particular that characterizing the tribe Otarionini. A second species, Otarionella lkomalii sp. nov., known only from a complete articulated specimen discovered in the early Eifelian of southern Morocco, is also described. Like the middle Eifelian Otarionella chamaeleo (Basse, 1997), this new species has only ten thoracic segments, with the fourth and the sixth segments each bearing a long axial spine. In the light of the new elements provided by the ontogenetic sequence of O. rastrum sp. nov. and the adult specimens of this species and O. lkomalii sp. nov., the putative synonymy of Otarionella and Otarion is rejected and a restricted concept of the genus Otarionella is defined.

Biostratigraphical ranges and palaeogeographical distribution of mid-Givetian to end-Frasnian odontopleurids are investigated. The discovery of Leonaspis rhenohercynica sp. nov. in mid-Givetian strata extends this genus unexpectedly up to the late Middle Devonian. New material of Radiaspis radiata (Goldfuss, 1843) and the first koneprusiine in Britain, Koneprusia? sp., are described from the famous Lummaton shell-bed, Torquay, Devon. New taxa of Koneprusia, K. serrensis, K. aboussalamae, K. brevispina, and K. sp. A and K. sp. B are defined. Ceratocephala (Leonaspis) harborti Richter & Richter, 1926, is revised and reassigned to Gondwanaspis Feist, 2002. Two new species of Gondwanaspis, G. dracula and G. spinosa, plus three others left in open nomenclature, are described from the late Frasnian of Western Australia. A further species of Gondwanaspis, G. prisca, is described from the early Frasnian of Montagne Noire. Species of Gondwanaspis are shown to possess a number of paedomorphic features. A functional analysis suggests that, unlike other odontopleurids, Gondwanaspis actively fed and rested with the same cephalic orientation. The sole odontopleurid survivors of the severe terminal mid-Givetian biocrisis (‘Taghanic Event’) belong to the koneprusiine Koneprusia in the late Givetian and Frasnian, and, of cryptogenic origin, the acidaspidine Gondwanaspis in the Frasnian. Whereas the former became extinct in the late Frasnian at the Lower Kellwasser Event, the latter disappeared, and with it the entire Odontopleuroidea, at the terminal Frasnian Upper Kellwasser global biocrisis.