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We have a strong focus on the wellbeing of our guests and course participants, a good training environment, air quality and modern equipment. Ihr PDF ist nun fertig. Weitere Informationen Verstanden und fortfahren Einstellungen. Campus Studenten aller weiteren Programme Mitarbeiter Alumni. Another tubercle lateral of L1 in the holotype of C. Posterior tegimen: many species have a densely granulose area that covers a large part of the pygidium and continues onto the lateral extremities of the thoracic pleurae. This area is often demarcated anteriorly by larger, rounded e. We term this feature posterior tegimen Lat.

Subocular patches: first described by Basse , pp. The feature is also present on internal moulds see Basse, , pl. A posterior subocular patch can be present as well e. Subocular patches can also be recognised in other proetides, as for example in species of Diademaproetus. Thoracic dorsal spine: a long posteriorly directed spine centrally on the sixth thoracic axial ring is characteristic of most species.

In these cases the axial ring has grown significantly beyond the size of the adjacent rings. Other species can have a rudimentary spine e. Secondary loss of the dorsal spine is particularly widespread in members of the hamidi group. Reversals i. Additional taxa: Cyphaspis cf. Members of the ceratophthalmus group occur in middle to upper Eifelian strata in the Ardenno-Rhenish Mountains and probably coeval sections in Morocco.

All have a strongly inflated to somewhat cone-shaped, anteriorly overhanging median glabellar lobe; interocular fixigena is strongly inflated tr. Distinctive of at least three members C. This feature has been demonstrated to vary intraspecifically in C. Cyphaspis ceratophthalmus has two granules on the posterior cranidial border and serially homologous ones on each thoracic posterior pleural band; C. All members except the type species and an undescribed species from Morocco show a fairly stable inclination of the glabella lateral view between S0 and the dorsoventrally highest point, which is anterior of the eyes.

The brownish colour is common among trilobites from this locality. Complete specimen NHMM , in dorsal G and lateral H views the right axial furrow of the cranidium is filled with glue. Scale bars are 2 mm. The holotype is presumed lost; to replace this, a complete specimen IRSNB a; see Figs 3, 4 is here designated neotype. Articulated specimens of C.

Savelsberg and the late H. Prescher in yet surprisingly few photographs of whitened specimens have been published cf. The morphology of this species was briefly described by Richter and Basse ; those accounts generally are to the point except that Richter reported 12 thoracic segments. We agree with Basse that this must have been a typographical error in view of the fact that we have never counted more than 11 segments however, see discussion under hamidi group below. As a preliminary note we wish to mention the variable thickness and initial direction of the thoracic dorsal spine; a large median tubercle may be developed on the occipital ring or may be absent altogether; median glabellar lobe is distinctly cone-shaped and anteriorly overhanging or comparatively weakly vaulted and smoothly rounded sagittal section ; ornament on median glabellar lobe varies from granules alternating with tubercles adaxially, to exclusively tubercles; G2 and G3 can be very small similar to other tubercles in this area , developed as long cones, or as thick, short, yet very thick cones section of a cone can equal sagittal length of occipital ring ; G2 can be much larger than G3; G2 or G3 tubercles are occasionally misaligned i.

These variations also have a bearing on C. Erben established this species on comparatively poorly preserved cephalic remains only some of which are actually strato- and topotypical see Basse, , p. Regardless of the specific affiliations of the paratypes, the morphology of the holotype of C. This raises the question of whether C. A reassessment of C.

Most trilobites were collected at the type locality with limited stratigraphic control, or none at all, and a description of the successions has never been published. Van Viersen et al. The morphology of the holotype is known exclusively from circumstantial evidence. Goldfuss , pl. Its whereabouts remain unknown despite efforts by us and other workers to trace it pers. Basse, G. Heumann and B. Schoenemann in , and the late H.

Prescher in However, we believe that the variation among putative specimens of C. The only tangible connection between the holotype and specimens assigned to C. The combination of these features is a fairly unique one and known to us exclusively in some of the members assigned to the ceratophthalmus group herein. The arbitrary assignments of other morphotypes to C. Derived from eximius Lat. IRSNB a, complete specimen. Cephalon is exceedingly strongly vaulted sag.

Low occipital ring with granules along its posterior margin; median glabellar lobe of ovoid outline, widest tr. Cyphaspis eximia is in many ways similar to C. Among shared features are the deep, pit-like posterior border furrow of librigena; comparatively few tubercles on the librigenal and preglabellar fields; weakly impressed pygidial axial and pleural forrws.

The German species demonstrates a more smoothly rounded outline lateral view of the median glabellar lobe; G2 and G3 are never spiny; genal spines are proximally dorsally and distally inward curved; a generally less vaulted cephalon sag. Named after Rainer Bluhm. Small sized member of the ceratophthalmus group with comparatively large eyes; comparatively weakly vaulted tr.

Low occipital ring bearing randomly scattered tubercles; median glabellar lobe of ovoid outline, widest tr. Cyphaspis bluhmi and C. An age indication of the species from Zireg cannot be precisely established at present but the type material of C. Bonino and A. Both species bear all of the characteristic features of basal members of the group while at the same time they attest to a trend of eye size increase and decreased glabellar inflation.

Additionally, both species are very small. The G2 and G3 tubercles are retained in C. Scale bar is 1 mm. Named after Enrico Bonino who donated the holotype for study. IRSNB a, pygidium. Resteigne locality, Belgium; lower part of Hanonet Formation, uppermost Eifelian. Member of the ceratophthalmus group with exceedingly small L1; median glabellar lobe lower than the eyes; librigenal field bearing several tubercles adjacent to the eye; broad tr.

Low occipital ring bearing numerous tubercles; median glabellar lobe of elongated ovoid outline, widest tr. Pygidium with four axial rings; each ring bears two tubercles pairs close to the axial furrow, each consecutive ring has an increasing amount of tubercles between these pairs; four pairs of pleurae, the posterior pleural bands of which bear a row of granules; border furrow is not developed; border widens slightly backwards; postaxial field is depressed medially.

Cyphaspis boninoi sp. The anterior pleural bands of the paratype are damaged just interior of the change in inclination of the pygidial pleural field. This is precisely the area where exceedingly large tubercles may be found in other Cyphaspis species exceptionally so in C. The localised damage in the paratype pygidium leads us to speculate that it carried large tubercles here. Additional taxa: Otarion aff. I herein , Givetian?

Along with the description of C. Our hamidi group is a much extended concept of Chatterton et al. Stratigraphically low members have a weakly vaulted cephalon covered with ubiquitous, closely spaced and uniformly sized ornament; exceedingly narrow interocular fixigenae; weakly inflated median glabellar lobe as opposed to the peramorphic, anteriorly overhanging median glabellar lobe of most other Cyphaspis species ; large, spherical eyes; long sag. The dorsal spine on the sixth thoracic axial ring varies between species from long to short, or absent, but the oldest species of which the thorax is known C.

Basse in We report the discovery of a unique specimen of an undescribed hamidi group member at our Foum Zguid locality which has 12 thorax segments. This amount is striking for it has been generally assumed that post-cephalic segment number variability was constrained in phylogenetically derived trilobites subsequent to the Cambrian radiation.

We assume that likewise, some Cyphaspis species may have been able to loosen constraints on thoracic segmentation and return to the plesiomorphic 12 segments. Future efforts by us will be focused on the recovery of additional specimens from the particular level at Foum Zguid. The incursions of hamidi group members were often associated with one other, non- hamidi group Cyphaspis species.

Cyphaspis lerougei is the oldest member from Morocco known to date. The exceedingly similar C. Another early representative of the hamidi group, C. Vidal et al. These last three units are of early late Emsian age and comparable to the lower and middle parts of the Marrettes Formation Vidal et al. The main trend of these is involved with the original concept of Chatterton et al. Developments that occurred in the uppermost Emsian to Eifelian include, typically, increased L1 size tr.

An increase in eye size peaked in the middle Eifelian whereas the earlier L1 size increase, reversed in some species e. The origin of C. Both are considered to be basal taxa of the hamidi group in bearing most of the primitive characters. On the other hand, they already differed markedly from coeval members of the main trend in having several conspicuous features in common: very robust genal spines; scant cephalic ornament; short, rounded median glabellar lobe that slightly overhangs anteriorly.

These dissimilarities suggest that C. If this is true then the hamidi group is at least diphyletic. However, as already explained above, our Cyphaspis groups are based on overall similarities between species and thus not necessarily monophyletic. Figures as far as they are not taken from the present note: C. At the time of this writing there exists an undescribed species from the middle upper Emsian in Germany which could be ancestral to Eifelian hamidi group members in the Ardenno-Rhenish Mountains M. Basse, pers. The cephalic ornament is ubiquitous but can be sparse and of increased size on and near the anterior to lateral cephalic borders; median glabellar lobe is comparatively strongly inflated, but very rarely overhanging anteriorly, and posterolaterally distinctly incised by large, teardrop-shaped L1; small pits on the librigenal field near the lateral border furrow are sometimes seen; eyes are comparatively small for members of the hamidi group; the numbers of pygidial axial rings six and pygidial pleurae five are clearly increased.

Species belonging to this trend are rarely known from complete specimens. Basse , pl. This species is very similar to C. All specimens are on a single rock slab. Named after Frederik Lerouge. Member of the hamidi group with dorsally flattened anterior cephalic border; subcircular median glabellar lobe; genal spine is very robust proximally, blunted distally and subsequently extended as a thin spine; cephalic border, librigenal, fixigenal and preglabellar fields bear few, weak granules concentrated near the eye; pygidium with two well developed axial rings; a third pygidial axial ring is weakly indicated only medially.

Cephalon is scantily ornamented; Fx1 and Fx4 are retained in large holaspides; preglabellar field bears a row of granules; librigena bears few, randomly scattered granules; occipital ring bears a median tubercle with a few weak granules nearby; eyes are somewhat outward directed, overhanging the librigenal field. Anterior two thoracic axial rings bear a single median granule; each consecutive ring bears more granules; sixth axial ring bears a median tubercle; posterior pleural bands each bear two granules.

Small pygidium that is almost entirely covered by posterior tegimen consisting of exceedingly finely grained and densely spaced granules that become slightly larger anteriorly; posterior border is upturned medially. One of the paratypes Fig. It is not clear at this point whether these differences should be attributed to intraspecific variation. This specimen is similar to C.

We did not position it in the diagram because we do not know its exact origin other than the indication that it comes from Jbel Issoumour not necessarily equivalent to any of our Issoumour localities. An Emsian age may be inferred from its morphological resemblances to the other two mentioned species. Named after Michael Kipping. IRSNB a, a complete specimen of uncertain origin; in all probability from the type locality and horizon.

Member of the hamidi group with a weakly inflated anterior cephalic border; proximally robust and distally acuminate, sword-shaped genal spines with dense granulation; small dorsal spine on the sixth thoracic axial ring; thorax is devoid of prosopon except for the posterior tegimen abaxially on segments 9 to Cyphaspis kippingi has been positioned in the diagram as an intermediary form between Otarion aff. Some of the features of basal members of the hamidi group are retained such as the long genal spines that are covered with granules; weakly inflated, weakly dorsally arched anterior cephalic border.

At the same time C. These last two differences may be explained by the slightly poorer preservation of the thorax and pygidium of the holotype. Named after its geographic occurrence. Member of the hamidi group with a comparatively weakly vaulted tr. This specimen is an indeterminate member of the hamidi group and likely represents a new species.

Of note is that it has three tubercle pairs on the preglabellar field which occur sporadically in species of Otarionini cf. Holotype complete specimen IRSNB a, in dorsal A , oblique anterolateral B , close-up of first nine thorax segments C , anterior D , close-up of thorax from sixth segment and including pygidium E , lateral F views. D; Alberti, pl.

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Member of the hamidi group with small, inflated anterior and posterior subocular patches; preglabellar, fixigenal and librigenal fields as well as frontal part of median glabellar lobe covered by moderately closely spaced granules; straight, fairly long genal spine with inward curved tip; small, narrow median glabellar lobe bearing tightly packed tubercles dorsally; median tubercle on occipital ring and serially homologous ones on thoracic axial rings 1, 2, 4, ; from third thoracic axial ring backwards, each consecutive ring bears more tubercles.

Cyphaspis juergenhollandi is much more similar to stratigraphically lower species than to roughly coeval C. When positioning C. However, the teardrop-shaped median glabellar lobe and genal spines of C. His specimen is generally similar to the holotype of C. After Zwier Smeenk, in recognition of his work on Devonian trilobites. Member of the hamidi group with an exceedingly scantily ornamented cephalon; median glabellar lobe bearing widely spaced granules; genal spine entirely straight, gently tapered; cephalic outline subtrapezoidal; dorsoventrally high anterior to lateral cephalic borders of subrectangular cross-section; shallow anterior to lateral cephalic border furrows.

Cyphaspis smeenki displays an unusual combination of features that does not fit any specific trend of the hamidi group. These include a very long thoracic spine; sparse granules on the median glabellar lobe and absence of ornament elsewhere on the cephalon. We have linked C. The dorsoventrally high anterior to lateral cephalic borders are also seen in C. Strictly taken i. We do not consider such an anteriorly overhanging glabella to be consistent with the weak inflation and overall cephalic morphology of this species. Cephalon with attached thorax segments IRSNB a, in anterior A , lateral B , oblique dorsal C , anterolateral D , oblique anterolateral E , close-up of right librigenal border furrow with row of pits F , posterior G views.

Notice the anomalous right genal spine.

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A single specimen of Cyphaspis sp. I is recorded which is of Eifelian or Givetian provenance at Jbel Issoumour pers. DH with local workers in There are only two similar forms to C. I that we know. One is a paratype cephalon of Cyphaspis balanops see Basse, , pl. The other is a large morphotype of C. The Moroccan taxon only differs significantly from the latter in having broad, very well-developed anterior to lateral cephalic border furrows; longer genal spines; slightly less pronounced cephalic ornament. Cyphaspis sp.


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I has a continuous row of small pits just interior to the anterior to lateral cephalic border furrow. This is generally a rare feature among Cyphaspis species yet again it is known in C.

Translation — EGO

A more straightforward example is a librigena of coeval and morphologically similar C. Based on these analogies to the Ardenno-Rhenish species we believe that our specimen is of Eifelian rather than Givetian age. A of Chatterton et al. The agayuara group represents the main rootstock of Cyphaspis including the oldest known species from the Silurian. Its members were widely palaeogeographically distributed in the Lower Devonian whereas only few representatives remained in the Middle Devonian.

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Here we describe two Pragian species, C. Members of the group are conservative in that they all have many typical i. The exclusion of phylogenetically derived taxa could render the agayuara group as a paraphyletic collection of species with a primitive morphology. It is possible that a future cladistic analysis leads to the conclusion that Cyphaspis should be subdivided but until that time it remains the oldest available name to accommodate species.

It must be recollected that in the very hypothetical scenario that each of our Cyphaspis groups should be warranted generic rank, the genus Cyphaspis would encompass the ceratophthalmus group because the latter contains the type species. Nevertheless he concluded that detailed comparisons were not feasible because the Spanish taxon is not very well known. Cyphaspis frankeorum belongs to the agayuara rather than the hamidi group in having smaller eyes; shorter sag. Other perspectives than dorsal view were not illustrated by Basse but an internal mould of a cephalon figured and identified as C.

Member of the agayuara group with moderately short, proximally well developed and distally acuminate genal spines bearing several tubercles; preglabellar field overhangs anteriorly, accentuated by a medial indention near border furrow; posterior pleural bands are increasingly strongly inflated towards the posterior end of the thorax and the pygidium.

Conspicuous yet damaged bulges are located on the thoracic and pygidial posterior pleural bands. These structures run obliquely across the posterior thoracic pleurae and onto the pygidium; their location suggests that they represent the anterior margin of the posterior tegimen. They are, however, too large to qualify as regular tubercles. The posteriorly lying area of the posterior tegimen consists of densely spaced granules whereas the pygidial pleurae are flattened; this creates a distinctive contrast with the otherwise strongly inflated thoracic and pygidial posterior pleural bands.

This species is exceedingly similar to stratigraphically lower C. Holotype complete specimen IRSNB a, in lateral A , dorsal B , oblique lateral C , close-up of left genal spine and anterior five thoracic pleurae D , oblique posterolateral E , anterior F , close-up of pygidium G views. Scale bar is 5 mm. Named after Horst Heising. Member of the agayuara group with high, collar-shaped occipital ring bearing a large median tubercle; large L1; comparatively large eyes for a member of the group ; furrowed genal spines; tubercles on preglabellar field verge to forming three consecutive, border parallel rows; anterior margin of posterior tegimen consists of verrucous tubercles and is extended onto the pleural tips of all thoracic segments.

Short, pear-shaped glabella, somewhat indented by elongate L1; palpebral lobe bears a row of granules on its margin; L1, fixigena, and median glabellar lobe bear closely spaced tubercles; dorsally flattened anterior cephalic border; anterior to lateral cephalic borders bear faint granules on outer half; trench-like anterior and lateral border furrow; genal spines are proximally posterolaterally directed and subsequently slightly convergent, yet they retain a posterolateral direction.

The thorax is broader across the middle segments than the ones anterior and posterior to these. This feature is also in line with the varying shapes of the pleural tips: in anterior segments the posterior tegimen seems to fortify and extend the posterior pleural band; in posterior segments the posterior tegimen is developed at the cost tr.

Cephalon with seven tergites attached IRSNB a, in anterior A , anterolateral B , dorsal C , oblique anterolateral D , oblique posterolateral E , close-up of occipital ring and anterior six thoracic axial rings F , lateral G views. Our material is almost indistinguishable from the single holotype specimen of C. We hope that the photos of a second representative of this species are helpful.

We regard them as an undescribed new species of the agayuara group. These workers noticed many similarities to C. Our specimen comes from a third locality in the same area and is coeval to C. A from which it only clearly differs in having much smaller L1. Our specimen is furthermore different from C. All three taxa are obviously closely allied and so any descriptions of new species are probably best postponed until more material becomes available for comparison. IRSNB a, cephalon. Member of the agayuara group with occipital ring bearing a median complex of granules instead of a single tubercle; median glabellar lobe, preglabellar field and anterior half of librigenal field covered with numerous granules; palpebral lobes and anterior to lateral cephalic borders covered with exceedingly closely spaced, fine granules; small eyes.

Like the stratigraphically higher C. Yet the rather distinctive morphology including the fine and densely spaced ornament, much more inflated median glabellar lobe, and tiny eyes warrant the description of C.


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A single specimen from Issoumour1 was obtained from workers but the stratigraphic origin is not known exactly. Its cephalon can only be discriminated from C. A tentative Emsian age is inferred from the morphological similarities between these two taxa. IRSNB a, incomplete cephalon. Tadachacht locality, Morocco; Khebchia Formation, basalmost upper Emsian. Member of the agayuara group with cephalon bearing large, cone-shaped tubercles on median glabellar lobe, L1 and fixigenae; large L1 and Fx4; tubercles are ubiquitous on preglabellar and librigenal fields.

Cyphaspis tadachachtensis is a very rare species that co-occurs in Tadachacht with C. It is similar to C. We are hopeful that future collections at the type locality will yield more material.