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DEVONIAN STYGINID TRILOBITE PARALEJURUS, WITH NEW DATA FROM SPAIN AND MOROCCO, THE

Schraut, Gunnar

ABSTRACT-

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 Rluixeros, 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.

INTRODUCTION

AMONG EARLY Devonian styginids Paralejurus Hawle and Corda, 1847 is distinguished by morphological features that may indicate a different
lifestyle and origin from contemporaneous styginids. Review of all known species of Paralejurus, together with new species and occurrences from
Southern Morocco (Fig. 1) and Spain (Fig. 2) presented herein, constrain its stratigraphical and biogeographical distribution and diversity. In
addition, the importance of Paralejurus in discrimination of crustal blocks and their dynamic relationships is evaluated.

FUNCTIONAL MORPHOLOGY AND LIFB HABITS

A few groups of trilobites have been claimed to have been endobenthic or semiendobenthie due to the gross morphology of their thick-shelled and
strongly vaulted exoskeletons, effaced surface structures, wide axial lobes, and the presence and orientation of terrace ridges. Such trilobites are
found in suitable depositional environments with muddy substrate conditions easily reworked during burrowing activities. The combination of
such criteria is generally not represented in Devonian styginids, with the exception of the highly diversified Paralejurus. Consequently, the
taxonomic position of Paralejurus among contemporaneous styginids is problematic.

Functional morphology.-The axis of the thorax and pygidium in Paralejurus is very wide (tr.) in comparison with the pleurae. This allows the
attachment of powerful muscles that might have provided the coxae with a strong rolling ability (Bergstrom, 1973, p. 44 for lsotelus Dekay,
1824). This morphological structure is unique among the late styginids and suggests a different life habit for Paralejurus. Unlike
contemporaneous styginids with their extended, flattened, often spinose exoskeleton typical for an epibenthic and necto-benthic life activity, the
highly arched morphology of Paralejurus is most similar to that of effaced earlier styginids (i.e., Rhaxeros Lane and Thomas, 1980 and allies) and
illaenids such as Panderia Volborth, 1863 or Illaenus Dalman, 1827, for which Bergström (1973) proposed a semiendobenthie life habit.
Conversely, concave-upward arched exoskeletons that occur frequently in illaenids and have also been observed in Paralejurus were interpreted
by Whittington (1997) as evidence for a life position in shallow depressions on the sea floor. However, this morphological pattern might result
from post-mortem muscle contraction.

Further support for the correlation between vaulted morphology and burrowing life habit comes from hydrodynamic and mechanical
experiments regarding swimming and burrowing in horseshoe crabs. Fisher (1975) suggested greater swimming ability and endurance for the
relatively flat Mesolinndus walchi as a result of its low water resistance and the outstretched appendages, whereas the highly vaulted Limulus
polyphemus has a better ability in burrowing which requires stronger flexed appendages to allow for powerful action against greater resistance.

Terraces.-As in most styginids, terrace ridges arc present in Paralejurus, commonly dorsally but always on the ventral side of the exoskeleton.
The dorsal terraces have their steep slope posteriorly, obliquely, or adaxially directed and are frequently associated with rows of pits paralleling
the base of the steep slopes. The terrace ridges on the ventral doublure invariably form continuous concentric lines that run subparallel to the
margins and have their steep slopes facing outward (anteriorly on rostral plate, abaxially on lateral part of librigenal doublure). The terrace
ridges on the doublure do not exhibit pits along the outwardly directed steep slopes. There are two competing interpretations of the function of
such terrace ridges and both may apply (Fortey, 1986). The current-monitoring system (Miller, 1975) is a plausible model to explain the obvious
relationship between ridges and associated rows of pits interpreted as sensorial organs. The absence of such pits on the ventral side and the
invariably abaxially directed paltern of ventral terraces favors Schmalfuss's (1981) interpretation of a mechanical function to stabilize the
substrate. The forward facing steep slopes on the ventral parts of the anterior cephalon might have provided frictional assistance for burrowing
backward into the substrate (Stitt, 1976; Fortey, 1986).

Geological evidence.-Paralejurus is found mainly in normally oxygenated calcilutites characterized by bioturbation, less commonly in shales and
limey mudsloncs. The Bohemian Pragian species [P. bohemiens najdr, 1960, P. hmngniarli (Barrande, 1846a), P. campanifer (Beyrich, 1845),

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P. richten najdr, I960, and P. zippei (Barrande, 1846a)| occur in the Kon prusy, Vina ice, Lod nice, and Dvorce-Prokop limestones, representing
biodetrital lime-mud substrates away from existing reef cores and therefore suitable for burrowing (Chlupá , 1983, p. 49 and tab. 2). Analogous
Paralejurus-bcaring facies occur in the Steinberger Limestone, Germany (P. richten), the light Crinoidenschuttkalk, Austria (P. cf. s ub
campanifer), the Lower Hamar-Laghdad Limestone, Morocco (P. brongniarti, P. hamlagdadicus G. Alberti, 1983), the Calcaire de la Sarthe,
France (P. brongniarti), the Calcaire à polypiers siliceux, France [P. depressus Feist, 1974, P. rugosus Feist, 1974, P. subcampanifer (Frech,
1887)], the Calcaire de Chaudefonds, France (P. dormitzeri ligeriensis Pillet, 1972), the Calcaire d'Angers, France (P. galloisi Oehlert and
Oehlert, 1890), the Ballersbacher and Ense limestones, Germany [P. dormitzeri applanatus (Novák, 189O)], the Greifenstein Limestone,
Germany (P. rehamnanus G. Alberti, 1970), and other localities containing shales such as the Tentakulitenschiefer of Germany (Paralejurus cf.
dormitzeri Barrande, 1852). In the Dalejan of Bohemia P. dormitzeri dormitzeri (Barrande, 1852) accumulated, after gravitational transport, in
neptunian dyke fillings in the Kon prusy reef (Suchomasty Limestone) and may have been derived from a different environment. In conclusion,
there is, as far as we know, no example of Paralejurus associations in reefs or from hard-rock substrates which would exclude a semiendobenthic
life habit.

STRATIGRAPHY AND GEOGRAPHICAL DISTRIBUTION

Paralejurus has not been recorded with certainty from the Silurian although Chlupá et al. (2000, p. 88) suggested that the genus originated in
the Silurian and continued into the Devonian. Indeed, P. balchashensis Maksimova, 1975 is reported to occur in Silurian-Devonian boundary
beds of Kazakhstan (Maksimova, 1975), but no associated biostratigraphical markers confirm this earliest occurrence. Likewise, the possible
presence of the genus in the Lochkovian is not yet documented with figured material (Pek and Van k, 1989, p. 30). Another report from the
upper Lochkovian of Bohemia (Chlupá et al., 1972, p. 168 and table 1) is not substantiated by any illustration or description. Welldocumented
species of the genus exist no earlier than the Pragian: Bohemia with five species (Beyrich, 1845; Barrande, 1846a, 1846b, 1852; najdr, 1960),
Morocco with five species (G. Alberti, 1983; Schraut, 2000 and herein), Armorican Massif with one species and the Rhenish Slate Mountains
("exotic limestones") with two species (both in open nomenclature). Thus with 12 species Paralejurus experienced a sudden burst of diversification
in the Pragian. In the Lower Emsian six species are documented from Spain, Bohemia, and Montagne Noire (Frech, 1887 and herein; najdr,
1960; Feist, 1974). The youngest documentation of the genus is in the late Emsian where its diversity is reduced to only five species. The fauna is
dominated by different subspecies of Paralejurus dormitzeri Barrande, 1852 and related forms such as P. rehamnanus G. Alberti, 1970 and P.
intumescens A. Roemcr, 1855. In the late Emsian Paralejurus is not only restricted to North Gondwana (Morocco, two species) and the
geodynamically related terranes of Bohemia (one species), the Montagne Noire (one species), and the Armorican Massif (two species), but occurs
for the first time on the Avalonia-Baltica margin (Harz Mts., Rhenish Slate Mts., and Holy Cross Mts.) (Roemer, 1855; Burhenne, 1899;
Osmolska, 1957, among others).

There is some confusion in the literature regarding the occurrence of Paralejurus in the Middle Devonian. However, since the internationally
accepted definition of the Lower-Middle Devonian boundary by the ICS/IUGS is placed at the base of the Polygnathus partitus Biozone (Ziegler and
Klapper, 1985) this level now lies higher than previously accepted. Accordingly, many reports of early Eifelian occurrences are now considered
latest Emsian. The youngest Bohemian representatives of P. dormitzeri range into the partitus Biozone (Chlupá , 1985), thus definitely crossing
the Lower-Middle Devonian boundary. This may also be the case with P. dormitzeri applanatus (Novák, 1890) from the locality "Blauer Bruch"
(Eastern Rhenish Slate Mts.) from where isolated specimens are considered to be Eifelian in age (Basse, 1996). Although the ranges of last
occurrences must currently be checked with index conodonts, we agree with Chlupá (1994, p. 493) that Paralejitrus disappeared within or at the
top of the partitus Biozone, and was probably a victim of the basal Eifelian Chote bioevent.

Annotated species list.-The format of this list follows that of Adrain and Chatterton (1994, p. 311). The list includes all valid species of
Paralejurus, along with several questionable assignments.

balchushensis Maksimova, 1975; uppermost Silurian or lowermost Lochkovian, Central Kazakhstan; two cranidia, one pygidium.
Stratigraphical assignment questionable.

bohemicus najdr, 1960; Pragian, Vina ice Limestone, Bohemia, two pygidia. New material from Issoumour, Maïder, South Morocco, one complete
specimen, one cranidium, one librigena, and four pygidia (herein).

brongniarti (Barrande, 1846a); Pragian to Lower Emsian, kindlei to dehiscens biozones after Chlupá et al. (2000), Bohemia, material see under
subspecies.

brongniarti brongniarti (Barrande, 1846a); Pragian, Dvorce-Prokop Limestone, Bohemia, two complete specimens, two cephaIa, two cranidia,
five pygidia, one rostral plate.

brongniarti menanensis Snajdr, 1960; Pragian, Vina ice Limestone, Bohemia, one pygidium and one cranidium. New material from Foum
Zguid, Dra-valley, South Morocco, three fragmentary cephalo-thoraces, two pygidia (cf. menanensis herein).

brongniarti mixal najdr, 1986; Pragian, Lod nice and Slivenec limestones, Bohemia, four cephala, 60 cranidia and pygidia.

campanifer (Beyrich, 1845); Pragian, upper Kon prusy, and Vina ice limestones, Bohemia, one complete specimen, four cephala, two cranidia,
five pygidia, one free cheek, and one rostral plate. Mentioned from the ?Pragian, Harz Mountains, Germany, one poorly preserved cranidium and
one pygidium (see H. Alberti, 1981); assignment questionable.

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carlxi n. sp.; Lower Emsian, upper Zlíchovian, Valdelinar, Celtiberian chains, Spain, one complete specimen, one cephalon with four thoracic
segments attached and dislocated incomplete pygidium, six cranidia, six pygidia, two free cheeks, and two rostral plates (herein).

depressus Feist, 1974; Lower Emsian, Izarne Formation (middle member), Montagne Noire, France, five cranidia, six pygidia, and one free cheek.

dormitzeri dormitzeri (Barrande, 1852); Upper Emsian, Suchomasty Limestone, Bohemia, two cephala, five cranidia, six pygidia, one
hypostome, and one rostral plate. Specimens referred to this subspecies as cf.-determination are from the Upper Emsian, Holy Cross Mountains,
Poland, one fragmentary craniclium (Osmolska, 1957), from the Upper Emsian, Montagne Noire, France (one cranidium, one free cheek, see
Feist, 1974), and from the ?Upper Emsian, Tentakulitenschiefer, Leun, Rhenish Slate Mountains, Germany (one pygidium, Burhenne, 1899).

dormitzeri ligeriensis Fillet, 1972; Emsian, Calcaire de Chaudefonds, northwestern France, two poorly preserved pygidia; questionable on
subspecies level.

dormitzeri applanatux (Novák, 1890); ?Upper Emsian, Bailersbacher, and Ense Limestones, Bicken, Kellerwald, Germany, two pygidia (holotype
lost) and additional two complete specimens and six pygidia. Stratigraphical attribution questionnable.

elayounensis Schraut, 2000; Pragian, Profil El Ayoun, approx. 16 km southeast of Tata, southeastern Antiatlas, South Morocco, six pygidia.

galloisi Oehlert and Oehlert, 1890; Emsian, Calcaire d'Angers, St.-Barthélemy, northwestern France, one pygidium (see Pillet, 1972).

hamlagdadicus G. Alberti, 1983; Pragian, lower Hamar Laghdad Limestone, five pygidia.

intumescens A. Roemer, 1855; ?Upper Emsian, corbis- to CaIceola members, Harz Mountains, Germany, two cranidia and three pygidia (sec
Basse, 1996).

rehamnanus G. Alberti, 1970; Upper Emsian, Rehamna, Central Morocco, one cephalon, several pygidia (see G. Alberti, 1970), and mentioned
from the Upper Emsian or Lower Eifelian, Greifcnstein Limestone, Greifenstein near Herborn, Hesse, Germany, one cranidium (see Kim, 1997).
New material is referred to this taxon from the Upper Emsian, Oufatène, Maïder, South Morocco, two complete specimens, one cephalon, one
cranidium, jind two pygidia (herein).

richteri najdr, 1960; Pragian to Emsian, Dvorce-Prokop, and ZIíchov limestones, Bohemia, one cephalon, three pygidia. Additional material is
referred to this taxon from the upper Pragian, Steinbcrger Limestone, Giessen, Hesse, Germany, one cranidium (Heinrichs, 1983).

rugosus Feist, 1974; Lower Emsian, Izarne Formation (base of middle member), Montagne Noire, France, two cranidia, five pygidia, one free
cheek, and one hypostome.

spatuliformis n. sp.; Pragian, Oufatène, Maïder, South Morocco, one complete specimen, one cranidium, one librigena, six pygidia (herein).

tenuistriatus n. sp.; Upper Emsian, Oufatène, Ma'ider, South Morocco, one complete specimen, one pygidium (herein).

subcampanifer (Frech, 1887); Lower Emsian, Izarne Formation (middle member), Montagne Noire, France, three cranidia, six pygidia, one
hypostome (see Feist, 1974). Additional material is referred to cf. subcampanifer from the Pragian, Garnie Alps, Austria, one cephalon (see
Ellermann, 1992).

teres Ancigin, 1979; Lower Emsian, gronbergi- to invervai-zones, eastern Urals, Russia, one cranidium, one free cheek, and four pygidia
(Ancigin, 1979, pi. 9, figs. 1-7).

verneuili (Oehlerl and Davoust, 1879); Pragian, Sarthe, France, one cephalon and one incomplete pygidium. Additional material is referred to
cf. vemeuili from the Pragian, Phacops potieri Zone, Sarthe, northwestern France, one pygidium (see Fillet, 1972), questionable.

zippei (Barrande, 1846a); Pragian, Kon prusy Limestone, Bohemia, one eephalon, one cranidium, nine pygidia (see najdr, 1960).

n. sp. A G. Alberti, 1981; Pragian, Lower Hamar Laghdad Limestone, Tafilalet, South Morocco, one fragmentary pygidium.

sp. s. Basse, 1996; Upper Emsian, Harbccke-Formation, Winterberg, Germany, several cranidia, pygidia, free cheeks, not figured.

sp. s. Chlupá , 1983; Lower Emsian, Zlichovian, Bohemia, undocumented.

sp. s. Chlupá et al., 1972; Upper Lochkovian, Bohemia, undocumented.

sp. s. Chlupá and Luke , 1999; Lower Emsian, Guerichina strangulata Zone, Bohemia, undocumented.

sp. s. Fillet, 1972; Emsian, Calcaire de la Sarthe, Sablé, France, single poorly preserved pygidium; questionable.

SYSTEMATIC AND PHYLOGRNETIC ASPECTS

Despite its distinctive morphology the assignment of Paralejurus to the Scutelluinae was hitherto seldom questioned (Harrington et al., 1959;
najdr, 1960, 1986; Erben, 1967; Whittington, 1999, among others) except by Fillet (1972), who erected the Paralejurinae as a distinct
subfamily without giving a diagnostic justification. The evolutionary history and the position of Paralejurus within the styginids have not been

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elucidated in great detail.

Indeed the origin of Paralejurus remains cryptic. Prantl and Pribyl (1947) and Richter and Richter (1956) considered Paralejurus to be derived
from Scutellum Pusch, 1833 at or close to the Silurian-Devonian boundary. However, phylogenetically important features such as the form of
the dorsal and glabellar furrows, the narrow axis, and the undivided median rib in Scutellum argue against its ancestry to Paralejurus.
Maksimova (1968, textfig. 1) derived Paralejurus from Scabriscutellum Richter and Richter, 1956, but typical representatives of this
polyphyletic taxon do not occur earlier than mid-Devonian and are thus much younger than the oldest Paralejurus. P ibyl and Van k (1971)
proposed Scutellum or Decoroscutellum najdr, 1958 as possible ancestors of Paralejurus.

In our view Paralejurus, though related to the Scutelluinae, is distinctive, occupying similar epi- to semiendobenthic habitats as the Illaeninae of
Ordovician and Silurian times. This can be deduced from many morphological "illaenimorph" features such as the convexity of the exoskeleton,
the wide thoracic axis, and trends towards effacement of sculptural surface elements. However, the typical and highly consistent morphology of
the pygidium with seven ribs, a terminal bifurcated median rib, the presence of a short but distinctive axis surrounded by dorsal furrows,
occurrence of 10 thoracic segments, and the presence of occipital and dorsal furrows in the cephalon is quite distinct from the Illaeninae (see for a
detailed comparison Whittington, 1999, p. 417-419). The presence of "illaenimorph" features in Paralejurus is considered to be the result of
adaptive homeomorphy.

Among the characteristic traits that remain unchanged through time within Paralejurus are the relatively high cross vault of the exoskeleton,
the wide axis, the anterior glabellar lobe expanding to border, narrow fixigenae, the hyperbolic course of the posterior axial furrows in the
cranidium, and the faint impressions of glabellar furrows and pygidial axial furrows. Other traits underwent evolutionary changes that allow
Pragian representatives to be distinguished from Emsian species. One such evolutionary trend concerns the occipital region. In the oldest-known
representatives, such as P. brongniarti and the new species P. spatuliformis, the occipital ring remains narrow (sag.), without medial
enlargement, and is only poorly differentiated from the faint occipital furrow. The large occipital muscle scars remain virtually flush with the
glabella. In later forms the occipital ring enlarges medially at the expense of a deeper and narrower occipital furrow. The reduced occipital
muscle scars are placed on a much lower level than both the glabella and the occipital ring. Whereas the Pragian representatives have blunt
librigenal corners, the Emsian species developed spines. In the thorax the fulcral notches situated near to the axis at the beginning tend to
migrate outwards. The pygidium has a transverse section with vertical anterolateral corners in older forms whereas this is bell-shaped with
evenly outstretched anterolateral corners in the younger representatives. The anterolateral lobes (possible remnants of first axial ring?) that
prominently encroach onto the first pleural rib in Pragian species become very tiny in Emsian ones and tend to disappear in the latest Emsian.

Taking into account the portrayed direction of evolutionary trends, we expect that direct ancestors of Paralejurus must have been largely smooth
and highly vaulted, being provided with narrow fixigenae and an outward curved posterior axial furrow in the cranidium and a wide axis and
shorter pleurae in the thorax. In this respect the mid-Late Silurian styginids represented by the Australian Rhaxeros Lane and Thomas (1980)
best meet these criteria. Shared derived characters concern in particular the overall high vault of the exoskeleton, the wide thoracic axis, the
absence of deeply impressed furrows, and prominent sculpture. However these traits are profoundly controlled by environmental conditions and
are therefore subject to convergence leading to homeomorphy.

A possible evolutionary lineage from the Rhaxeros group (including Liolalax Holloway and Lane, 1999) to Paralejurus would be independent of
the other late styginid lines. Yet the inclusion of Paralejurus in late styginids by Whittington (Scutelluinae s. Whittington, 1999) is based on
important diagnostic features such as the pattern of glabellar impressions and the structure of the thoracic pleurae, especially the free terminal
spines lacking articulating facets. It is difficult to argue that these characters were separately acquired in scutelluines and Paralejurus, as would
be the case if the latter were derived from the Rhaxeros group. There is no evidence of intermediates between Paralejurus and the Rhaxeros group
in these characters. Therefore we agree with D. Holloway (personal commun., 2003) that Paralejurus should be retained within the Scutelluinae
(s. Whittington, 1999). However, we consider Paralejurus as a monophyletic group within the latter, evolving with a different lifestyle. We
exclude the possibility that its particular morphology was derived more than once by iterative evolution from various scutelluine ancestors. In
this context, if the Scutelluinae are once again elevated to family rank (as originally proposed by the Richters) we would favor the subfamily
status "Paralejurinae" for this group.

DYNAMIC PALHOGEOGRAPHY

Due to its epi- and semiendobenthic life habit in well-oxygenated shallow neritic biofacies away from the open oceanic realm, Paralejurus is
particularly suited for biogeographic comparisons between continental blocks that are separated by oceanic interspace (Fig. 3).

The earliest known representatives of Paralejurus from the Pragian are exclusively distributed in both the North Gondwana mainland (southern
Morocco, Montagne Noire, Iberian Chain) and Gondwana-derived microblocks, such as the Armorican Terrane Assemblage (including Bohemia
and the Armorican Massif) and the Proto-Alps (including the Garnie Alps) (McKerrow et al., 2000; Tait et al., 2000). The occurrence in
Kazakhstan might also be related to Gondwana. It is noteworthy that morphologically similar styginids such as Rhaxeros and Liolalax from the
mid-Silurian are restricted to Gondwana (Fig. 3a) which might emphasize the importance of vaulted styginids for paleobiogeographic
discrimination at that time.

The Emsian is a crucial period for geodynamic plate movements as many benthic biotas, especially trilobites (G. Alberti, 2000) and ostracodes
(Berdan, 1986) hitherto restricted to North Gondwana, appear for the first time in terranes of the southern Avalonian margin on the Laurussia
supercontinent providing biogeographic constraints on approaching continental margins in Europe. In the context of general compressional

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geodynamics generated by the northward migration of the Gondwana margin and forerunning microplates (Matte, 2001) remnants of
mid-European oceanic pathways should have largely been reduced by the end of the Early Devonian. Indeed, the characteristic late Emsian
species P. dormitzeri occurs on both sides of the previously impassable Rheic ocean. The sudden and contemporaneous disappearance of
Paralejurus altogether in Gondwanan and Avalonian terranes in the earliest Eifelian argues for close geographic relationship between both plates
in which biotas underwent the same extrinsic conditions.

SYSTEMATIC PALEONTOLOGY

The described and figured material is deposited in the collections of the following institutions: SMF (Senckenberg Museum Frankfurt, Germany)
and MPZ (Museo Paleontologico Zaragoza, Spain).

Family STYGINIDAL Vogdes, 1890

Subfamily SCUTELLUINAE Richter and Richter, 1955

Genus PARALEJURUS Hawle and Corda, 1847

Type species.-Bronteus campanifer Beyrich, 1845, from the Kone prusy Limestone (Pragian), Me n any, Bohemia; original designation.

Diagnosis.-Exoskeleton strongly vaulted (sag. and tr.); glabella expanding to anterior border; shallow glabellar furrows; narrow fixigenae;
strongly diverging posterior axial furrows; axial region wider than pleural field; median rib in pygidium bifurcate.

PARALEJURUS BRONGNIARTI cf. MENANENSIS S najdr, 1960

Figure 4.1-4.6

cf. Paralejurus brongniarti menanensis S NAJDR, 1960, p. 196, pi. 25, figs. 10, 11; 1986, p. 129, pi. 3, figs. 1, 2.

Material examined.-Cephala with attached thoracic segments SMF 76646-48; pygidia SMF 76674-75 from Assa Formation (s. Hollard, 1963,
correlated by Jansen, 1998), Early Devonian (Pragian).

Occurrence.-Plain north-northeast of Mdouer-el-Kebir, southeast of Foum Zguid, Southern Morocco.

Discussion.-Among the different subspecies of brongniarti the new material from southern Morocco is nearest to menanensis. In particular the
course of the anterior axial furrow in the cranidium with a straight portion adjacent to the palpebral lobe and its distinctiveness around the
anterolateral corners of the glabella that characterize the Bohemian subspecies is also seen in the African material. Poorly preserved pygidia
seem to have a smooth exoskeleton, unlike the Bohemian material, which has scaly terrace ridges as on the cranidium.

PARALEJURUS BOHEMICUS S najdr, 1960

Figure 4.7-4.13

Paralejurus bohemicus S NAJDR, 1960, p. 261, pi. 25, figs. 4, 5. G. ALBERTI, 1983, p. 25-26; SNAJDR, 1986, p. 128.

Diagnosis.-Cephalon with entire border anteriorly and laterally, glabella unfurrowed, flattened anteriorly and posteriorly, marked occipital
furrow deflected forwards medially, high palpebral lobes, blunt librigenal angles; evenly vaulted pygidium without border, faint triangular axis,
pointed behind; terrace lines on abaxial parts of exoskeleton.

Description.-Cephalon bordered anteriorly and laterally with a continuous, upturned rim extending backwards to genal angle and carrying
strong terrace ridges. Frontal outline of glabella defined by uninterrupted narrow preglabellar furrow. Glabella of moderate vault (tr. and sag.),
axe-shaped, defined anterolaterally by almost straight diverging axial furrows. Frontal lobe of glabella forward projected and pointed in lateral
view. Glabellar furrows inconspicuous. Medial part of glabella in front of occipital ring flattened, laterally merging with large, subtriangular,
and slightly inflated occipital muscle scars. Occipital furrow continuously deep, meeting axial furrows, medially enlarged (sag.) and curved
forward. Occipital ring spindle-shaped, remaining below level of glabella. Palpebral lobes slightly higher than posterior part of glabella, not
protruding further outward than maximum extension of cranidium (tr.). Anterior branches of suture subparallel to axial furrows. Librigenae
vaulted transversely, unfurrowed except at base of eye socle. Genal angle blunt, less than right angle. Entire cephalon densely covered with pits.
Besides pitting, external surface smooth with the exception of frontal glabella lobe, palpebral lobes, posterior part of occipital ring, and marginal
parts of librigenae which carry wavy terrace lines.

Material examined.-Complete specimen SMF 76651; cranidium SMF 76652; librigena SMF 76653; pygidia SMF 76654-57 from Ihandar
Formation, Early Devonian (Pragian).

Occurrence.-Jebel Issoumour, Maider, south-southeast of Alnif, southern Morocco.

Discussion.-Paralejurus bohemicus was hitherto known from its pygidium alone. New material from southern Morocco, including pygidia
indistinguishable from the Bohemian ones and an entire specimen, allows completion of the diagnosis and description of the species with
characteristic features of the cephalic region. Among contemporaneous species, the pygidium of P. elayounensis Schraut, 2000 has a similar

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shape lacking any upturned border region and exhibits the typical triangular backwardly pointed axis as in P. bohemicus. By contrast, the
former has a granular sculpture and a different outline, being narrower in front. However, pygidia of P. bohemicus tend to become more
elongated during postlarval growth. The apparently close relationship between P. bohemicus and P. elayounensis must be reevaluated after the
discovery of cephalic sclerites of the latter.

PARALEJURUS SPATULIFORMIS new species

Figure 5

Diagnosis.-Cranidium strongly vaulted, glabella sculptured with interconnected wrinkles; pygidium tonguelike, moderately vaulted, flat-topped
with steeply declining anterolateral flanks, clearly impressed narrow inter-rib and axial furrows.

Description.-Cephalon highly arched transversely, narrow semielliptic in outline, breadth/height index at base 1.75. Cranidium anteriorly
regularly curved, entire edge framed with narrow upturned rim. Anterior part of glabella strongly expanding laterally, 2.5 times wider than at
base. Glabellar furrows very shallow, inconspicuous. Fused S1 and S2 very shallow, similar to that of P. campanifer (Bcyrich, 1845) (see S najdr,
1960, fig. 58; pi. 23, fig. 2). S3 transversely elongate, connected to axial furrow by shallow depression. L2-L3 not inflated, flush with median
part of glabella. Occipital furrow very shallow, poorly differentiated from front of occipital ring, laterally superseded by protruding triangular
occipital muscle scars. Occipital ring narrow (sag.) and low, uninflated. Palpebral area slightly arched (exsag.), palpebral lobes expanding
beyond projection of maximum extension of anterior cranidium (exsag.). Visual surface of eye lobe vertical behind and less steeply inclined in
front of the palpebral lobe. Librigenae with unfurrowed genal field that steeply slopes from the eye to the lateral margin; lateral edge bordered by
very narrow rim. Genal angle obtuse. Anterior part of glabella with a dense network of irregular wrinkles that progressively vanish to the rear;
fewer isolated wrinkles and short terraces developed on the outer palpebral lobes, occipital ring and periocular portion of the librigenae; posterior
part of glabella in front of occipital ring smooth.

Pygidium elongated in the form of a spatula, strongly vaulted posteriorly and laterally with nearly vertical anterolateral sides (length to width
approx. 0.8-1.1 and length to height approx. 2.0-2.6), flat-topped adaxially. Axis subpentagonal, very low, virtually flush with adaxial region of
the pleural field; median lobe very narrow, U-shaped, not reaching the tip of the axis, with anteriorly strongly diverging longitudinal furrows
that die out in the anterior third of axis well behind articulating furrow; small triangular lateral lobes weak but obvious. Axial furrow and
inter-rib furrows narrow and clearly marked on external surface. Paired ribs straight, median rib narrowest at about one-third length from
anterior, subdivided by shallow longitudinal furrow anteriorly and posteriorly but not in middle part. Border extremely narrow and slightly
upturned, rapidly expanding rearwards being very wide and concave posteriorly. Posterior outline varies ontogenetically from narrow elliptical
to largely semiparabolic. Entire surface of pygidium with densely spaced, backwardly directed short ridges and elongated nodules.

Comparison.-The new species is distinctive in its tonguelike elongated pygidium. This feature is shared with the contemporaneous P. richten from
Bohemia and the younger P. galloisi from the Emsian of northwestern France. The former differs from P. spatuliformis in the longer pygidial axis
with a pronounced triangular outline, and the sculpture that covers the entire glabella, whereas P. galloisi has a regularly and less strongly
vaulted pygidium in lateral view, the inter-rib furrows are nearly effaced, and the subdivision of the unpaired medial rib is not obvious.

Etymology.-After its typical elongated, spatulalike pygidium.

Types.-Holotype, pygidium SMF 76658; Figure 5.6, 5.11, 5.13. Paratypes, cephalon with attached thorax and fragmentary pygidium SMF
76659, coll. Lemke; incomplete cranidium SMF 76660; librigena SMF 76661; pygidia SMF 76662-67 from Ihandar Formation, Early Devonian
(Pragian).

Occurrence.-Jebel Oufatène, Maïder, South Morocco.

PARALEJURUS CARLSI new species

Figure 6

Diagnosis.-Exoskeleton moderately vaulted; cranidium very wide anteriorly, truncated in front with clearly impressed glabellar furrows; short
genal spine; pygidium bell-shaped in posterior view with large prominent axis, outwardly curved ribs and deep, distally expanded inter-rib
furrows.

Description.-Cephalon semicircular in outline, relatively moderately vaulted in lateral view, culminating in height in the posterior part of the
occipital ring; breadth/height index at base approximately three. Glabella axe-shaped, well defined laterally by almost straight, strongly
diverging deep axial furrows, nearly three times broader across the frontal lobe than across occipital furrow. Frontal outline of glabella
truncated, rectilinear medially. Anterior border very narrow, slightly concave adaxially and merging with median part of frontal lobe.
Glabellar impressions relatively well marked. S1 with poorly impressed anterior branch and a deeper posterior one that is fused to S2 by
moderately deep longitudinal furrow; medial elevation of S1 large, inconspicuous; S2 ending blindly anteriorly. S3 elongated, not extending to
dorsal furrow on external surface. L2 + 3 trapezoidal, slightly swollen. Preoccipital part of glabella parabolic in posterior outline. Occipital furrow
wide (sag., exsag.) and shallow medially, even wider laterally where it contains slightly swollen triangular occipital muscle scars merging
posteriorly with narrow, gently convex (exsag.) occipital ring. Posterior slope of fixigenae swollen. Palpebral lobes horizontal, on lower level than
occipital furrow, laterally remaining inside of the line of maximum extension of anterior cranidium (exsag.); anteriorly straight, clearly

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impressed palpebral furrows. Free cheek with broad lateral border and shallow border furrow; genal spine short, broad-based.

Pygidium semielliptic, bell-shaped in transverse section with slightly upturned margins. Prominent axis short (sag.), subpentagonal in outline,
wider anteriorly than pleural field (tr.), with three slightly swollen longitudinal lobes and an additional pair of small anterolateral lobes that
dominate adjacent pleural fields in lateral and posterior view; median lobe slightly higher than lateral ones, defined by shallow anteriorly
diverging furrows that do not reach the broad (sag.) articulating furrow. Outwardly curved pleural ribs separated by deep narrow furrows that
expand and deepen distalIy shortly before vanishing at the upturned border. Median rib twice as broad as neighboring ribs at the tip of the axial
lobe, gently expanding posteriorly and provided in the anterior two thirds of its length with a weak keel, bifurcating in the posteriormost
quarter.

With the exception of the furrows, the exoskeleton is covered with scalelike, short ridges, and tubercles. Peripheral parts, i.e., anterior fixigenae,
librigenae, thoracic pleurae, and anterolateral margins of pygidium, have discontinuous terrace ridges, mostly running exsagittally. Anterior
border of cranidium, and cephalic (including rostral plate) and pygidial doublures with continuous terraces with steep slopes facing outwards.
The entire exoskeleton displays spaced pits that are aligned along the steep slopes of the terraces on the anterior cranidial border and on the
rostral plate. Pygidial doublure seemingly unpitted.

Comparison.-The new species is distinct with respect to the comparatively well-impressed glabellar furrows and deep inter-rib furrows in the
pygidium. It shares shape and outline of the pygidium with contemporaneous taxa such as P. subcampanifer and P. rugosus from southern
France; it has a similar nodular sculpture and the same gradually enlarging median rib as the former but the lateral ribs are outwardly curved
and not straight as in P. subcampanifer. Curved ribs occur likewise in P. rugosus but this species has a less prominent axis and a different
sculpture. Other Emsian species with bell-shaped (in transverse section) pygidia and short genal spines such as P. dormitzeri and P. rehamnanus
are more vaulted transversely, have different sculptures and less prominent pygidial axes.

Etymology.-After Prof. Dr. P. Carls, who found the material of this new species.

Types.-Holotype, pygidium MPZ 01; Figure 6.9-6.11. Paratypes, cephalon with attached four thoracic segments and a dislocated incomplete
pygidium MPZ 02; enrolled entire specimen MPZ 03; cranidia MPZ 04-08; rostral plates MPZ 09-10; librigenae MPZ 11-12; thoracic segment MPZ
13; pygidia MPZ 14-21 from Mariposas Formation, Lower Devonian (Lower Emsian, upper Zlìchovian).

Occurrence.-All from region of Nogueras, Camaras River and Andrea section (Fig. 2), Spain.

PARALEJURUS REHAMNANUS G. Alberti, 1970

Figure 7.1, 7.3-7.5, 7.8, 7.9

Paralejurus dormitzeri rehamnanus G. ALBERTI, 1970, p. 33, pi. 2, figs. 9, 10; KIM, 1997, p. 186, pi. 10, fig. 4; SCHRAUT, 2000, p. 371, pi. 1.
fig. 10. subrectangular.

Diagnosis.-Frontal margin of cranidium with slight medial embaymenl; preglabellar furrow interrupted medially; basal lobe of glabella very
wide transversely; axial furrows diverging in almost straight line anteriorly; stout librigenal spine. Pygidial axis very low and weakly
differentiated from pleural field on external surface, undivided except for median lobe; inter-rib furrows outwardly curved abaxially, median rib
keeled, virtually undivided on external surface.

Description.-Glabella moderately and evenly vaulted (sag. and tr.), axe-shaped, almost straight, diverging axial furrows in front of occipital
furrow, anterolateral corners subrectangular, medially incurved frontal lobe merging with central depression on rostral plate. Glabcllar furrows
not impressed on external surface, with combined S1-S2 as unsculptured flat area. Occipital furrow distinct, narrow sagittally, expanding
(exsag.) abaxially and containing triangular, low, uninflated occipital muscle scars laterally. Occipital ring very broad sagittally, rapidly
narrowing abaxially, moderately vaulted (sag.) and remaining below level of glabella just in front. Palpebral lobes long (exsag.), lower than
glabella, with weak palpebral furrow, protruding beyond maximum extension of anterior cranidium. Anterior branches of suture parallel to
axial furrows. Visual area vertical below posterior half of palpebral lobe, progressively changing inclination forward and reaching 45 degrees in
front. Base of eye surrounded by rather deep and large furrow. Librigenal field unfurrowed, vaulted, framed by a very thin border rim that
vanishes at base of strong, relatively long genal spine. Pygidium semicircular in dorsal outline, rather highly vaulted (tr. and sag.), bell-shaped
in transverse section with broad, flat, and inclined border that is twice as wide posteromedially as anterolaterally. Axis lacking independent
convexity on external surface, being flush with adjacent parts of the pleural field; only median lobe is slightly prominent posteriorly, defined
laterally by faint furrows. Relicts of tiny anterolateral lobes are present. Inter-rib furrows slightly curved outward, vanishing abaxially at inner
edge of border. Unpaired median rib of equal width over most of its length, keeled medially, slightly expanding distally; median longitudinal
furrow not seen on external surface. Entire dorsal exoskeleton densely pitted, pits on pygidial axis enlarged. all parts of exoskeleton are covered
with prominent sharp terrace ridges.

Ontogeny.-Smaller holaspid cranidia (Fig. 7.3, 7.4) have a differently shaped glabella which is less vaulted transversely and relatively more
extended anterolaterally than in larger specimens. In addition glabellar furrows are clearly though slightly impressed on the external surface.
These comprise a very broad (exsag.) S1 with sculptured eccentric median elevation, fused with a short longitudinal S2. S3 forms a rather tiny
isolated groove that remains far from the axial furrow. The occipital muscle scars appear as slightly inflated areas.

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G. Alberti (1970, pi. 2, fig. 10) figured a fragmentary pygidium of Paralejurus rehamnanus of 10 mm length that obviously belongs to a young
holaspis. In this specimen the axis is clearly separated from the pleural field by a continuous axial furrow. In our material a specimen of equal size
that is found in association with advanced holaspidcs exhibits the same configuration with the axial furrow present. We conclude that the axial
furrow in the pygidium shallows and finally disappears during postlarval growth.

Material examined.-Two complete exoskeletons SMF 76668-69, one juvenile exoskeleton with detached cephalon SMF 7667Oa and pygidium
SMF 7667Ob, cranidia SMF 76671-72, pygidium SMF 76673 from Tazoulait Formation, Lower Devonian (Upper Emsian).

Occurrence.-All from Jebel Oufatcnc, Mai'der, South Morocco.

Discussion.-The slight medial embayment of the anterior cranidial margin that characterizes Paralejurus rehamnanus from the central
Moroccan Meseta occurs also in an isolated cephalon from Greifenstein (Southern Rhenish Slate Mountains, Germany) attributed by Kim (1997)
to the same taxon. The new material from Maïdcr (southern Morocco) comprising five specimens representing different growth stages exhibit this
character without exception. In contrast, in the numerous specimens of P. dormitzeri from the Barrandian region as well as in related forms from
Montagne Noire (Feist, 1974, pi. 15, fig. 8 and additional new material), the anterior border, though upturned medially, has an evenly curved
outline and there is a narrow preglabellar furrow running parallel to the border without medial interruption. We think therefore that the slight
cmbayment in the anterior margin is an important feature that defines P. rehamnanus at the species level. Important new material of this taxon
including two entire exoskeletons allows the completion of its diagnosis and description.

G. Alberti (1970, p. 33) did not mention any further differences between P. rehamnanus and P. dormitzeri apart from the medial embayment of
the anterior cephalic margin in the former. His material of P. rehamnanus consisted only of a single cephalon with incomplete librigenae (the
holotype) and a fragment of a juvenile pygidium. The discovery of a poorly preserved cephalon in Greifenstein, Germany (Kim, 1997), was also
not helpful for further comparisons. In addition to the characters mentioned by G. Albcrti and Kim our material reveals the following differences
from P. dormitzeri that emphasize the specific independence of P. rehamnanus: in P. dormitzeri glabellar furrows are visible on the external
surface, the pygidial ribs encroach onto the border, the median rib is subdivided by a short furrow, and terrace lines are less sharp and
prominent.

The occurrence of Paralejurus rehamnanus in the Eifelian reported by G. Alberli (1970, p. 33) after information given by Gigout (1951) is not
dated by index fossils and therefore doubtful. The new material is definitely from the Lower Devonian (upper part of the Upper Emsian, cf. Fig. 1).

PARALKIURUS TKNUISTRIATUS new species

Figure 7.2, 7.6, 7.7, 7.10-7.12

Diagnosis.-Cephalon with continuous border and anterolatcrally extended glabella that is narrow at occipital furrow; strong genal spines;
pygidium bell-shaped in transverse section, moderately vaulted, with short trapezoidal axis.

Description.-Cephalon with broadly parabolic anterior outline. Cranidium widest across lateral extremities of frontal lobe. Glabella strongly
expanding anteriorly, bounded by deep outwardly curved axial furrows and continuous preglabellar furrow. Evenly curved, narrow, slightly
upturned border surrounds entire front of glabella. Anterior glabellar lobe moderately vaulted transversely and sagittally. Glabellar furrows
well marked on external surface. Branches of Sl surround large elongate central elevation; S2 long and straight (exsag.); S3 narrow, crescentic,
remaining far from dorsal furrow. Basal glabellar lobe flat and low, semicircular in posterior outline. Occipital furrow deep and broad (sag.),
containing weakly inflated occipital muscle scars laterally. Occipital ring enlarged medially, evenly vaulted (sag.), of equal height as glabella.
Palpebral lobes short (exsag.), higher than basal glabellar lobe, remaining far inside maximum extension of cranidium, with two pairs of weak
furrows that run parallel to palpebral suture. Visual surface of equal inclination around palpebral lobe. Subocular furrow deep and narrow.
Anterior branch of facial suture running along suturai ridge. Unfurrowcd librigenal field moderately and evenly vaulted (exsag. and tr.),
sloping to shallow marginal depression. Lateral border slightly upturned with thickened edge carrying terrace ridges. Librigenal spine with
broad base. Inner portion of convex thoracic pleurae and first pygidial rib with relicts of faint pleural furrows discernible on adaxial half of
pleurae. Pygidium semicircular in outline, of moderate transverse and longitudinal vault, bell-shaped in posterior view with flat inclined border
that expands posteromedially. Axis very short (sag.), truncated behind, defined by faint continuous axial furrow. Axial lobes low but discernible,
middle lobe slightly inflated, laterally defined by straight depressions that originate in the articulating furrow but vanish shortly before
reaching the dorsal furrow. Inter-rib furrows sigmoidal in curvature, expanding abaxially, and vanishing at base of break in slope before border.
Median rib keeled, enlarged at axis, expanding slightly and uniformly backward, bifurcated at the extreme rear by very short and faint median
longitudinal furrow. The entire exoskeleton is pitted and covered by liny wavy terrace lines and furrows that are continuous on the cephalic
parts but reduced to discontinuous short wrinkles on the pygidium, where they are accompanied by small patches of shallow groovelike
structures.

Etymology.-LaUn, tennis = thin, striatus (lat.) = striated with reference to the sculpture.

Types.-Holotype, complete exoskeleton SMF 76676; Figure 7.2, 7.6, 7.7, 7.10-7.12, Paratype, fragmentary pygidium SMF 76677 from
Tazoulait Formation, Lower Devonian (Upper Emsian).

Occurrence.-All from Jebel Oufatène, Maïder, South Morocco.

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Discussion.-The new species is closely related to the other latest Emsian representatives, such as P. dormitzeri and P. rehamnanus, that all exhibit
genal spines, a pygidium with bell-shaped transverse section, and the same kind of dense sculpture of terrace ridges throughout the entire
carapace. The new taxon is distinguished mainly by the outline of the glabella with its narrow base and its strong anterolateral expansion. In
addition, the short trapezoidal axis in the pygidium is distinctive.

ACKNOWLEDGMENTS

We are very grateful to the editors and the two referees G. Edgecombe and D. Holloway for their critical and useful comments and corrections
that improved our manuscript. We thank both P. Carls and U. Lemke who made available important material used in this study and provided
locality information. R. Lerosey-Aubril and G. Webster made helpful comments on the original manuscript. K. Wcddige is thanked for the loan of
specimens housed in the Senckenberg Museum. The work was supported by a Marie Curie Fellowship by the European Community to GS
(HPMF-CT2000-00591). This is a contribution to IGCP 421 "North Gondwana mid-Palaeozoic bioevent/biogeography patterns in relation to
crustal dynamics." Publication ISEM 2003-059.

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ACCEPTED 3 NOVEMBER 2003

GUNNAR SCHRAUT AND RAIMUND FEIST

Institut des Sciences de l'Evolution, Laboratoire de Paléobotanique et Paléontologie, Université Montpellier II, Place E. Bataillon 34095
Montpellier Cedex 5, France,

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