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NEW STYGINIDS FROM THE LATE DEVONIAN OF WESTERN AUSTRALIA-THE LAST CORYNEXOCHID TRILOBITES

McNamara, Kenneth J

ABSTRACT-

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.

INTRODUCTION

WITHIN THE trilobite family Styginidae, the Scutelluinae is a highly diverse subfamily, ranging in age from the Early Ordovician to the Late
Devonian. Hitherto, the last recorded scutelluines were considered to occur in the early and middle Frasnian. They are all assigned to Scutelliim
Pusch. 1833, a long-lived conservative genus that originated in the Late Silurian (Maksimova, 1968). Most of the Frasnian specimens have been
assigned to its type species S. costatum Pusch, 1833, widely distributed in perireefal limestones along the Avalonian margin of the Old Red
Continent and in its type area, the Polish Holy Cross Mountains (Richter and Richter, 1926). It is also present in Gondwana-related terraines,
such as the Montagne Noire of southern France (Feist, 1974), the Kazakhstanian Mugodjar, the Southern Urals (Maksimova. 1955) and the
Kuznetsk Basin (Tchernysheva, 1951). In the latter regions other species related to S. costatum occur. In continental North America a single
species, S. thomasi (Walter, 1924), is known from northern Iowa. In contrast, no Late Devonian scutelluine has previously been described from
the main Gondwanan region, including South America, Africa, eastern Asia, and Australia-Antarctica. As all known last occurrences of
Scutelliim are from the latest Mid-Frasnian conodont Zone 12 (Klapper, 1988). the Styginidae were thought to have become extinct prior to, or
at, the Lower Kellwasser horizon (boundary Zone 12/13). being victims of the early pulse of the Kellwasser crises (Feist. 1991; Feist and
Schindler, 1994; Fortey and Owens, 1997, p. 287). However, as part of our study of an undescribed Middle and Late Frasnian trilobite fauna
from the Canning Basin, Western Australia, we here record and describe the youngest known representatives of the Scutelluinae. These Frasnian
representatives of the subfamily display some unusual morphological characteristics, including a pronounced vaulting of the pygidium and
marked eye reduction. Their occurrence through the last three conodont zones of the Frasnian (i.e., zones 11-13 a and b sensu Klapper et al.,
2004), up to and including the basal part of the final conodont zone of the Frasnian, the linguiformis Zone (=Zone 13b) (see Girard et al, in press),
confirms that the family extended close to the end of the Frasnian (but not reaching the Frasnian/Famennian boundary), becoming extinct at a
level that is time-equivalent to the base of the Upper Kellwasser Horizon (the latter is not developed in Canning sequences). As such the Canning
Basin species represent the last known members of the order Corynexochida.

GEOLOGICAL SETTING, STRATIGRAPHY, AND AGE

The Late Devonian sedimentary rocks that outcrop along the northern margin of the Canning Basin of Western Australia form one of the
best-preserved Paleozoic reef complexes in the world (Playford and Lowry. 1966: Playford. 1980, 1981, 1984; Becker et al., 1989, 1991).
Extending for about 350 km along the Lennard Shelf along the northern margin of the Canning Basin from the Napier Range in the northwest to
the Lawford Range in the southeast (Fig. 1), this reef complex contains rich invertebrate and vertebrate faunas that range in age from ?Late
Givetian to Famennian.

The first Devonian trilobites recorded from the Canning Basin were specimens collected by Harry P. Woodward, the Western Australian
Government Geologist, while on a visit to the Kimberley region of Western Australia in 1906. These were sent to the British Museum, London,
where they were identified by his father, Henry B. Woodward, as "tail of a Trilobite genus-Proetus, a new species" and "head of a Trilobite (part of
same specimen) (?) and counterpart of tail" (Glauert, 1910). These specimens were collected from Barker Gorge in the Napier Range, probably
from the Virgin Hills Formation. Only the "Proetus" specimen is still in existence (WAM 2051). These, and a few other specimens of brachiopods
and corals, were significant in enabling Devonian strata to be unequivocally recognized in Western Australia for the first time, by H. B.
Woodward.

The pioneering studies by Teichert (1943, 1949) on the reef complexes and inter-reefal basins provided further indications of the existence of a
rich Late Devonian trilobite fauna. From the Frasnian part of the Virgin Hills Formation, a forereef facies, Teichert recorded Harpes Goldfuss,
1839, Scutellum, and Pteroparia Richter, 1913, along with proetids that he assigned to Chaunoproetus Richter and Richter. 1919,
Drevermannia Richter, 1913, and Cyrtosymbole Richter, 1913. From the Famennian "Cheiloceras Zone" (= Lower Nehdenian, Late Devonian II-A
to F) he recorded Cyrtosymbole sp. From the "Sporadoceras Zone" (=Upper H = Nehdenian to Lower Hembergian, UD II-G-1II-C) he recognized a
form which he assigned to "Perliproetus sp. nov." The material which Teichert collected has since disappeared and was never formally described.

Some of the Famennian trilobites from this area, however, have been recently described: the phacopines Babinops Feist and Becker, 1997 and
Trimerocephalus McCoy. 1849, and the proetid Cyrtosymbole. In this present paper some of the Frasnian elements of the trilobite fauna are

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formally described for the first time. The remaining trilobite fauna! elements, including representatives of the Proetidae, Tropidocoryphidae,
Odontopleuridae, Harpetidae. and Phacopidae, will be described in subsequent papers.

The material described in this paper came mainly from the western and eastern Hanks of McWhae Ridge, a faulted reef spine located at the
southern end of the Lawford Range in the Kimberley District of Western Australia (Fig. 1). Here, Frasnian reef-building stromatoporoid and coral
limestones of the Pillara Formation are overlain by the steep forereef facies of the Frasnian Sadler Limestone, and the Frasnian-Famennian Virgin
Hills Formation (Playford. 1980). All the trilobites occur within the Virgin Hills Formation. This consists of very fine-grained calcarenites and
calcilutites that are thinly bedded, gently dipping (10°-20°), and commonly hematite-rich. They contain a diverse fossil assemblage, dominated
by goniatites (Becker et al., 1991), with 103 species/subspecies having been recognized (Becker, 2000). Other elements of the fauna, in addition
to trilobites and goniatites, include: orthoceratid nautiloids, corals, sponges, bivalves, gastropods, crinoids. ostracodes, conodonts, and localized
horizons of stromatolites (Playford and Lowry, 1966).

A composite stratigraphie section that includes both Frasnian and Famennian strata (36 m) was measured from the base of the Virgin Hills
Formation on the east side of McWhae Ridge (1: 100,000, Bohemia Sheet 4160, Grid Reference 924258-924263), where some of the specimens
used in this study were collected (Fig. 2). The Frasnian-Famennian boundary occurs at a height of about 30.5 m. The Virgin Hills Formation
rests disconformably on the Sadler Limestone, as shown by the presence of a thin brecciated layer at the base. The fossiliferous beds from the
Virgin Hills Formation that onlap the ridge close to the contact between the Sadler and Pillara Formations have been interpreted as phases of reef
retreat with steady bioerosion and fine detritus supply (Becker et al., 1991).

The section on the eastern side of McWhae Ridge consists of a sequence of grey, yellow, and red, thin-bedded alternating calcilutites and
calcarenites. Fossiliferous beds containing trilobites within the Frasnian section are restricted to five discrete horizons and are generally
separated by sparsely fossiliferous. yellowish grey calcareous limestones. For convenience, beds containing rich macrofossil assemblages have
been designated informal names. From the base these are: Esko's Gully (7.0-9.0 m), Berigora Gully (11.0-12.5 m). Scutelluid Bed (12.5-13.5 m),
Phacopid Gully (19.5-27.0 m), and Calyx Corner (28.5-29.5 m) (Fig. 2). These fossil beds are characterized by their sedimentological nature and
faunal assemblages. They form prominent marker horizons that can be traced throughout the area and can be correlated with sections on the
western side of the ridge (see Becker et al., 1991). The fossiliferous limestones in Berigora Gully and at Calyx Corner comprise dark red and grey
micritic cephalopod and crinoidal limestones that are similar in many respects to coeval beds from late Frasnian localities in Europe (Becker et
al., 1989; Feist, 1991).

Trilobites were also collected from a section on the western side of McWhae Ridge (1:100,000, Bohemia Sheet 4160, Grid Reference 920260). This
section has received much more attention from paleontologists than the section on the east side of the ridge, and is discussed in detail by Becker et
al. (1991). Trilobites are abundant in two transgressive marker beds, both of which contain abundant specimens of goniatites. The lower of these
two beds contains many scutelluines, and equates with the Scutelluid Bed on the eastern side of the ridge. The other bed is in the lower part of
Becker et al.'s (1989) Upper Belocerax Bed, and contains a relatively rich trilobite fauna. This level is probably equivalent to the lower part of the
Phacopid Gully beds on the eastern side of the ridge, which contain scutelluines.

Zonation of trilobite-bearing sediments in the studied section is based on ammonoid and conodont biostratigraphy. Becker et al. (1991) and
Becker (2000) have provided the most comprehensive accounts of age diagnostic goniatite faunas in Frasnian sediments of the Virgin Hills
Formation, on the basis of more than 100 taxa. The goniatite zones of the Frasnian at McWhae Ridge range from the Miiternoceras retorcjuatum
Zone (J1) to the disappearance of this genus [end of regional zone (L2) (Becker et al., 1991)].

A finer scale zonation of the Late Devonian has been obtained using conodonts (Klapper, 1988; Ziegler and Sandberg, 1990). Frasnian-Famennian
conodonts from the Canning Basin have been documented by Glenister and Klapper (1966), Seddon (1970), Druce (1976), and Nicoll (1984).
Ziegler and Sandberg (1990) subdivided the Frasnian into 10 conodont zones, three of which are represented at McWhae Ridge: the early
rhenana Zone, late rhenana Zone, and linguiformis Zone (Fig. 2). These three zones correspond to the upper part of Zone 12, very latest Zone 12
and Zone 13a, and Zone 13b, respectively, of Klapper (1988) and Girard et al. (2005). Scutelluines were collected from the sections on the eastern
side of McWhae Ridge, from Berigora Gully, Scutelluid Bed, Phacopid Gully, and Calyx Corner. Conodont evidence places the first two horizons in
Zone 12 and the latter two in zones 13a and b, respectively. Specimens were also collected from the western side of the ridge from beds that are
equivalent to the Scutelluid Bed and Phacopid Gully horizons.

Scutelluines also occur to the north of McWhae Ridge in Bugle Gap, 10 km north-northwest of McWhae Ridge. The poorly preserved specimens
were derived from a bioclastic limestone a few meters above conglomeratic beds taken as the base of the Virgin Hills Formation (Seddon, 1970).
Conodont data indicate that these beds are older than those at McWhae Ridge, occurring within Zone 11. The third section where scutelluines
were collected was Horse Spring, in the Horse Spring Range, 60 km north of McWhae Ridge (1:100,000, Elma Sheet 41601 Grid Reference
858854). This 35 m thick Frasnian section extends through conodont zones 6-13c (G. Klapper personal commun.). Specimens were found in bed
23, about 5.5-5.7 m above the base of the section. From the overlying bed 24, Becker et al. (1991, p. 186) recorded proetid trilobites and
"Scutellum." These beds occur within the 12 ammonoid zone (Playfordites tripartus) which correlates with conodont Zone 11. Another recently
discovered nearby locality, 1.5 km to the south of Horse Spring, and just north of Siphon Spring, has also yielded scutelluine remains, and
similarly correlates with conodont Zone 11.

At McWhae Ridge the Frasnian-Famennian boundary is situated about a meter above the Calyx Corner horizon, these beds occurring at the lower
part of the linguiforinis Zone (=Zone 13b). The scutelluines present in the Calyx Corner beds therefore represent not only the last known record of

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the family Styginidae, but indeed of the order Corynexochida.

SYSTEMATIC PALEONTOLOGY

Material examined.-In 1989 collections were made in the Virgin Hills Formation at McWhae Ridge by G. Rokylle and E. Routasuo, who discovered
many of the localities subsequently collected by the authors, assisted by J. Long, D. Friend, C. McGeachie, and D. Haig. Further material was
collected there and from the locality in Bugle Gap by one of us (RF), T. Becker, M. House, P. Jell, and P. Playford. Specimens studied from Horse
Spring and Siphon Spring were collected by the authors and R. Lerosey-Aubril. Material is housed in the collections of the Western Australian
Museum, Perth (WAM). the Geological Survey of Western Australia, Perth (GSWA). and Museum Victoria, Melbourne (MV).

Terminology follows Whittington and Kelly (1997). For the cephalic morphological features we follow Whittington (1999), which is the most
recent account of these features in scutelluines.

Family STYGINIDAE Vogdes. 1890

Subfamily SCUTELLUINAE Richter and Richter. 1955

Genus TELOPELTIS new genus

Type species.-Telopeltis woodwardi n. sp. from the late Middle Frasnian (zones 11 and 12) of the Canning Basin, northwest Australia.

Assigned species.-Telopeltis woodwardi and Telopeltis microphthalmus n. sp.

Occurrence.-Late Middle through Late Frasnian (zones 11-13b). Canning Basin. Australia.

Diagnosis.-Small scutelluine in which anterior margin of frontal lobe in contact with convex anterior border; glabellar furrows, especially S2 and
S3, isolated from each other and very weakly developed; occipital ring very long, two-fifths glabellar length, with short occipital spine in small
holaspids; eye lobe small to very small; no genal spine. Pygidium extremely vaulted, height up to 60% of length: bell-shaped in posterior view;
short, up to 1.4-1.9 times wider than long: axis prominent, with small anterolateral lobes, inconspicuous medial lobe; well-incised, broad axial
furrow; fulcrum approximately midway between axial furrow and lateral margin; median rib not divided, broad adaxially; interrib furrows
narrow and deep; smooth, flat border well developed with entire margin, making angular junction with very convex pleural field.

Etymology.-From 'telos,' Greek, meaning 'last,' and the Greek 'peltis,' meaning 'shield.'

Discuxxion.-Scutelluines are generally large trilobites characterized by their relatively large and nearly flat pygidia. The only exceptions to this
are Telopeltis and the Early Devonian Paralejurus Hawle and Corda, 1847 [see Schraut and Feist (2004) for recent revision], which both have
highly vaulted exoskeletons. Despite this similarity, it is unlikely that Telopeltis is phylogenetically related to Paralejurus because of the many
other morphological differences between the two taxa. Telopeltis differs from Paralejurus not only in being much smaller, but in having a much
shorter pygidium with an undivided median rib, deeper furrows, and possession of a relatively broad, smooth border. Although both genera have
very reduced glabellar furrows, Telopeltis differs in having a glabella that is much narrower posteriorly, a deeper S1 partly enclosing a
well-defined node, S2 isolated from S1, and much smaller eyes.

Another scutelluine that, like Telopeltis, is characterized by small eyes, is the Emsian-Givetian Tenuipeltis Liitke. 1965. Although present in the
upper Givetian, there is no evidence that this genus persisted into the Frasnian, as currently recognized on the basis of conodont zonation.
Telopeltis can be distinguished from Tenuipeltis by having palpebral lobes set closer to the glabella, resulting in a narrower (tr.) fixigena.
Moreover, the glabellar furrows are weaker and not united by a longitudinal furrow, the frontal lobe is longer (sag.), and the cephalon is more
strongly vaulted. The pygidium of Telopeltis is very different from Tenuipeltis: it is far more vaulted; it is much wider; it has a median rib that
does not bifurcate distally; and it has broad pleural ribs, with relatively narrow interrib furrows and a longer axis.

The only other scutelluines that have been recorded from the Frasnian are the long-ranging genus Scutellum and Frasnielluin Basse in Basse and
Müller, 2004. The type species of Scutelluin, S. costatum. has its type locality in the Frasnian of the Holy Cross Mountains (Richter and Richter,
1926; Chlupac, 1992). A number of authors (Wright and Chatterton. 1988; Holloway, 1996) have pointed out that the material from the type
locality is not particularly well known. However, the neotype, a pygidium. illustrated by Archinal (1994, pl. 1, fig. 4) and topotype material
illustrated by Chlupac (1992, fig. 4). are sufficiently well preserved to highlight the differences between this form and Telopeltis. Unlike
Telopeltis, S. costatum has larger eyes and a better-defined S2 and S3. The most distinctive differences lie in the pygidium, which is much longer
and flatter than in Telopeltis and possesses a median rib that attenuates strongly adaxially. Like other scutelluines, S. costatum is much larger
than Telopeltis, with Middle Devonian forms from southwest England having a pygidium more than twice the length of the largest known
specimen of Telopeltis (Selwood, 1966).

Frasnielluin. from the mid-Frasnian of Germany (Basse and Müller, 2004), can be distinguished from Telopeltis in a number of ways. The frontal
lobe of the glabella is longer (sag.): the distance between S3 and the front of the glabella is equivalent to that between S3 and occipital furrow
(this distance being twice in Telopeltis). The front of the glabella is set far distant from the anterior border (sag.) in Fraxniellum, unlike
Telopeltis, in which it is in contact with the anterior border. The frontal lobe of the glabella slopes gently anteriorly in Frasnielluin, and merges
with a conspiciously large (sag.), flat area that is framed anteriorly by a thin, sharply edged, uninflated, upturned border. In Frasniellum S3 is a
rather short (tr.), transverse, relatively deep, narrow impression, unlike the fainter S3 of Telopeltis. The anterior sutures of the two genera

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follow different courses. In Frasnielluin it extends anteriorly to ?, initially running parallel to the axial furrow, then from opposite S3 it runs
exsagittally to meet the anterior border at ß at a right angle. In Frasniellum swollen eye ridges run from the anterior of the palpebral lobe
obliquely forwards toward anterior of L2, before dying out and before reaching the axial furrow.

Pygidia tentatively assigned to Frasnielluin (Basse and Müller, 2004, figs. 142-145) are quite different from pygidia of Telopeltis. The axis is
rather small, pointed posteriorly, with axial furrows sustaining an angle of less than 90° (much less than in Telopeltis). The outline of the
pygidium is nearly semicircular (unlike the semielliptical shape in Telopeltis). In lateral view, the pygidium of Frasniellum has a sigmoidal
profile, lacking the swollen form of Telopeltis, with a wide, upturned, sharp-edged border. The median pleural rib tapers more strongly anteriorly
in Frasniellum, becoming much narrower behind the axis than in Telopeltis.

TELOPELTIS WOODWARDI new species

Figures 3, 4

Diagnosis.-Glabella tumid with relatively long frontal lobe that projects anteriorly in large individuals: palpebral lobe 20% cranidial length,
situated opposite occipital furrow and posterior branch of S1 ; posterior branch of facial suture very short and transverse abaxially. Pygidium
very strongly vaulted (sag. and tr.), height more than half pygidial length. Exoskeleton with coarse tuberculation that on frontal lobe
diminishes and devolves into transverse terrace ridges; sparse covering on pygidial axis, where arranged in two exsagittal rows.

Description.-Cephalon reaching a maximum known length of 13 mm; strongly vaulted in lateral view, being highest at posterior of Ll and
posterior of occipital ring. Frontal outline of glabella gently arched forward in larger specimens; glabella narrowest at occipital furrow, defined
laterally by well-incised, narrow, and sinuous axial furrows that diverge forwards slightly from occipital furrow to anterior branch of Sl. then
diverge more strongly at about 100°, the frontal lobe reaching two and a half times the width of the glabella at the occipital furrow. Anterior
border very narrow, with up to five transverse terrace ridges; preglabellar furrow absent medially where frontal glabella is in contact with
upturned border. Glabellar furrows poorly developed: anterior and posterior branches of Sl broad and shallow, partly enclosing a well-defined,
abaxially protruding median node. S2 and S3 very faint and do not reach axial furrow; S2 isolated impression, broadly oval, very shallow, short;
S3 oval, fainter, and narrower than S2. Occipital furrow long (sag., exsag.) and moderately impressed, deepening anteriorly; weakly developed
lateral occipital lobes present. Occipital ring very long (sag., exsag.) and wide (tr.), being one and a half times wider than glabella at occipital
furrow; flat, inclined slightly anteriorly; short occipital spine present; combined length of occipital ring and furrow nearly 30% cephalic length.
Close to axial furrow faint fixigenal impression present on vaulted fixigenae. which do not reach as high as sagittal part of glabella; decline
steeply anteriorly and very steeply posteriorly. Palpebral lobe nearly 20% cranidial length, strongly curved, and situated high on fixigenae;
posteriorly positioned opposite occipital furrow and Ll; laterally anterior of lobe slightly adaxial of distal lateral extremity of frontal lobe (exsag.);
palpebral furrow shallow. Eye with large visual surface; moderately convex, consisting of curving rows containing up to 25 holochroal lenses.
Visual range extends through about 150°, allowing lateral and dorsolateral vision; no furrow ventral to eye surface. Posterior branch of facial
suture very short, extending exsagittally for very short distance before running transversely, then recurving strongly close to ill-defined
posterior border. Anterior branch moderately divergent close to eye, recurving anteriorly to be slightly convergent close to anterior border.
Posterior border with prominent fulcral process behind posterior tip of palpebral lobe. Pygidium semicircular, short; width up to 1.6 times
sagittal length. Very strongly vaulted, but gently concave marginally; height up to 60% pygidial length; distally pleural field dips at about 60°
to horizontal. When viewed in transverse profile, fulcrum positioned closer to axial furrow than to lateral margin. Small facet present lateral to
prominent fulcral process, bordered anteriorly by a thin, raised rim that bears terrace ridges. Axis broad, almost transverse in outline
posteriorly; narrower than pleural field; strongly vaulted semicircular posterior section behind relatively deep, transverse articulating furrow,
anterior to which is a prominent articulating half ring; weak anterolateral axial lobes present. Ill-defined medial axial lobe, anteriorly narrower
than lateral lobes, tapering rearwards when becoming flush with lateral lobes in its posterior part, remaining narrower (tr.) than proximal end
of median pleural rib. Pleural ribs moderately convex in profile, flattening adaxially; median rib widest, gradually widening posteriorly to
become slightly more than one and one half times anterior width; median furrow absent. In some specimens median rib enlarges shortly before
reaching axial furrow. Ribs separated by narrow, well-defined, anteriorly gently outwards curving furrows that broaden and deepen distally
before terminating prior to margin; anterior pair deepest and confluent with axial furrow; most other furrows reach axial furrow. Sharp, angled
break between pleural field and flat border that is inclined at about 45° to horizontal: slightly narrower than distal pleural rib width.

Fixigenae and occipital ring sparsely tuberculated. Glabellaalso with relatively sparsely distributed tubercles; smallest posteriorly and
anteriorly. Although present anterolaterally, tubercles absent anteromedially, where replaced by transverse wavy terrace ridges, steep scarp
facing posteriorly. Ridges crenulate and bear incipient tubercles anteriorly. Posteriorly these tiny irregularities become progressively larger and
develop into tubercles whose long axis is aligned posteriorly. As tubercles enlarge, terrace lines diminish in intensity then disappear altogether on
posterior one-third of glabella. Tubercles on pygidial axis posterior to articulating furrow arranged in a pair of gently curving rows close to
sagittal line that are slightly convergent posteriorly and some extremely small, posteriorly directed, barblike tubercles. Pleural ribs with
relatively dense covering of tubercles; single rows of tubercles adaxially, increasing to three or four across the ribs distally. Articulating half ring
covered by fine terrace ridges; these absent on facet. Border lacking tubercles.

Ontogeny.-Telopeltis woodwardi displays a certain amount of variation in pygidial characters, largely resulting from ontogenetic changes
during holaspid development. As the pygidium doubles in size, from 10 to 20 mm in length, the degree of vaulting reduces. Thus, when viewed in
lateral profile, the pleural region in smaller forms extends horizontally for about one-third of the pygidial length from the axial furrow, before
arching very strongly, reaching a steep angle posteriorly of up to about 60°. In larger forms the pleural field is inclined at about 10° as it extends

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out from the axial furrow, then progressively increases to about 45°. In the smaller, more strongly vaulted forms the flat border is inclined at an
angle of about 45°, the angle reducing to about 30° in larger, less vaulted pygidia. The angle between the border and pleural field is quite acute
in the smaller, more highly vaulted pygidia. There is also a tendency for the border to broaden during holaspid ontogeny, the width being less
than the distal lateral pleural width in small pygidia, but equal to that in the larger, less vaulted forms. The pleural furrows are a little more
incised in the more strongly vaulted forms, principally because the pleurae themselves are more convex (tr.). A consequence of the shallowing of
the pleural furrows is that some diminish to such an extent that they fail to meet the axial furrow. As the pygidia increase in size the pleural
furrows become more flexed; consequently the median rib, which steadily increases in width posteriorly in smaller forms, initially slightly
constricts posteriorly, before broadening distally. The axial furrow shallows through holaspid ontogeny, resulting in the axis being less defined in
larger forms.

Etymology.-Named in honor of Harry Page Woodward, who found the first Devonian trilobites in the Canning Basin.

Types.-Holotype, pygidium, WAM 04.304: paratypes WAM 91.306, 04.298-04.303, McWhae Ridge, Lawford Range, Canning Basin, Western
Australia: Virgin Hills Formation, Frasnian Zone 12.

Other material examined.-Many hundreds of specimens, mainly pygidia, from the "Scutelluid Bed," McWhae Ridge, Lawford Range, Canning
Basin, Western Australia; Virgin Hills Formation, Frasnian Zone 12. Four pygidia and IO incomplete cranidia from Bugle Gap. 10 km
north-northwest of McWhae Ridge, Lawford Range, Western Australia; Virgin Hills Formation, Zone 11; two pygidia (WAM 05.165, 05.166), one
cranidium (WAM 05.163). and one librigena (WAM 05.164) from Horse Spring and Siphon Spring, Horse Spring Range. Western Australia;
Virgin Hills Formation. Frasnian Zone 1 1 (Fig. 4).

Occurrence.-Late Middle Frasnian, zones 11 and 12.

Discussion.-One of the more distinctive aspects of Telopeltis woodward! is the sculpture on the glabella. As noted above, the anterior of the frontal
lobe is covered by transverse, wavy terrace ridges, in which the steeper scarp faces posteriorly. These are replaced by tubercles on the central
and posterior parts of the glabella. However, there is an area of overlap of the terrace ridges and tubercles, and this clearly shows that the
tubercles arise directly from irregularities on the ridges. The terrace ridges are not of even height, but are crenulate and the tubercles develop
from these crenulations. They start as slight irregularities on the crest of the ridge, and become more pronounced towards the back of the
glabella. In longitudinal profile the axes of tubercles are not perpendicular to the surface of the exoskeleton, but are angled posteriorly. Archinal
(1994. fig. 11) figured part of the frontal lobe of a specimen of Scutellum flahelliferum (Goldfuss. 1843), which shows a similar, though less
well-developed, effect. A combination of terrace ridges and tubercles on the anterior of the glabella has also been described in S. sudorum
Holloway, 1996.

Among scutelluines described from the Devonian of Australia there are a number of species of Scutellum. As this genus persists into the Late
Devonian, comparison is made with these species. S. hollandi Wright and Chatterton, 1988 from the Emsian Jesse Limestone in New South Wales
differs from T. woodward! in possessing a shallow preglabellar furrow; a short genal spine; pronounced occipital spine; larger eye and narrow (tr.)
fixigena; far more distinct S3; much larger, flatter pygidium which is concave distally: and a relatively shorter pygidial axis.

Scntelliim droseron Holloway and Neil, 1982 from the early Lochkovian Mount Ida Formation in Victoria differs from T. woadwardi in having
much coarser tuberculation; far more well-defined glabellar furrows; a preglabellar furrow; a frontal lobe which is much more expanded
laterally: and a much shorter occipital ring. The pygidium is poorly known, but unlike T. woodwardi it is flat and long.

Scutellum sudorwn Holloway, 1996 from the Emsian Murrindal Limestone in Victoria is much closer to T. woodwardi. It differs in possessing a
more forwardly expanding glabella; relatively well-defined S3 that meets the axial furrow; distinct preglabellar furrow; more variable
tuberculation; much flatter pygidium; and median rib that is much narrower adaxially.

Scutellum calniin Chatterton, 1971 from the Emsian Taemas Formation is similar to T. woodwardi in possessing relatively small, posteriorly
positioned eyes and also a glabellar ornamentation of anterior terrace ridges that transform into tubercles posteriorly. However, it has a glabella
that is much broader posteriorly; more well-defined glabellar furrows; shorter occipital ring; weak lateral occipital lobes; and flatter, longer
pygidium with a more heavily tuberculated axis and less well-defined border.

Scutellum lenuistriatum Feist and Talent, 2000. from the late Eifelian to possibly earliest Givetian of the Broken River region of Queensland, is
only known from the pygidium. It is possibly conspecific with "Brontem" tenuistriatus Tchernysheva, 1951 from the Middle Devonian of the
Kuznetsk Basin, from which only the cranidium is known. Like other species of Scutellum it differs from Telopeltix n. gen. in being much flatter
and longer, although in common with larger specimens of Telopeltix it possesses the characteristic dared median rib, which initally narrows
distal to the axial furrow before expanding towards the posterior border. Feist and Talent (2000) have noted how Archinal (1994) used the
presence of an adaxially broadening median rib to delineate the separate subgenus Scutellum (Calycosculellum) Archinal, 1994. However, as
they noted, this feature changes ontogenetically and is also present in Telopeltis, making its use as a diagnostic subgeneric trait questionable.

TELOPELTIS MICROPHTHALMUS new species

Figure 5

Diagnosis.-Small species with gently convex glabella: frontal lobe transverse anteriorly to slightly anteriorly projecting; node enclosed by Sl very

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small: anteriorly positioned palpebral lobe short, only 10% cranidial length, situated opposite S1; posterior branch of facial suture straight,
diverging, relatively long, extending posterolaterally at about 45°. Pygidium strongly vaulted (sag. and tr.), height a little less than half
pygidial length; fulcrum midway between axial furrow and lateral border. Exoskeleton with dense tuberculation that on frontal lobe extends to
anterior border; dense covering on pygidial axis.

Description.-Cephalon nearly semicircular in outline, reaching a maximum known length of 8.5 mm: moderately vaulted in lateral view, being
highest at posterior of glabella. Frontal outline of glabella truncated and rectilinear in smaller holaspids, slightly arched forward in largest
holaspid: glabella narrowest posteriorly, defined laterally by well-incised, narrow axial furrows that diverge slightly to anterior branch of S1,
then diverge more strongly at about 90°, resulting in the frontal lobe reaching nearly two and a half times the posterior width of the glabella.
Anterior border narrow, with up to five transverse terrace ridges; preglabellar furrow absent. Glabellar furrows very shallow; anterior and
posterior branches of Sl broad, demarcating a faint median node that laterally causes slight deflection in course of axial furrow. S2 and S3 do not
reach dorsal furrow; S2 broadly oval, short, diverging at about 150° and isolated from S1: S3 oval, fainter, and shorter than S2, being little more
than interruptions in the glabellar tuberculation. Occipital furrow wide (sag., exsag.) and shallow, deepening a little anteriorly; sharply
delineated by slightly posteriorly curved basal glabellar lobe; very faint lateral occipital lobes present. Occipital ring very long (sag., exsag.) and
wide, being half as wide again as posterior of glabella, posteriorly acuminate; flat, inclined slightly anteriorly and merging with occipital furrow,
protruding centrally at the expense of the occipital furrow; short occipital spine present in small holaspids; combined length of occipital ring and
furrow nearly 30% cephalic length. Close to axial furrow fixigenal impression present on posterior of vaulted fixigenae, which are as high as the
sagittal part of the glabella; decline steeply anteriorly and posteriorly. Palpebral lobe short, only 10% cranidial length, horizontal. and situated
high on vaulted fixigenae: anteriorly positioned opposite S1; laterally in line (exsag.) with distal lateral extremity of frontal lobe: shallow
sigmoidal palpebral furrow present. Small eye surface consisting of irregular curving rows containing up to 10 holochroal lenses; gently convex,
allowing lateral and limited dorsolateral vision; bounded ventrally by narrow, shallow furrow. Posterior branches of facial suture divergent,
relatively long, extending posterolaterally at about 45°, before recurving strongly close to ill-defined posterior border. Anterior branches
moderately divergent close to eye, recurving anteriorly to be slightly convergent close to anterior border. Posterior border with prominent
fulcrul processs behind posterior tip of palpebral lobe. Lateral border a thin raised rim demarcated by broad, shallow furrow. No genal spine.

Pygiclium semicircular, short, with breadth 1.4 times sagittal length in large holaspids. hut up to 1.9 in small ones. Strongly vaulted; height
40%-50% pygidial length. Fulcrum positioned hallway between axial furrow and steeply inclined lateral margin; well-developed facet present
lateral to fulcrum. Axis rhombicshaped: well defined by deep axial furrows; narrower than pleural field; strongly vaulted triangular posterior
section behind transverse articulating furrow, anterior to which is a prominent articulating half ring; small anterolateral lobes present; axial
lobes illdefined by absence of subdividing longitudinal furrows. Pleural ribs moderately convex in profile; median rib widest, being relatively
wide anteriorly and widening posteriorly to become about twice anterior width; lacking median furrow. Ribs separated by narrow, well-defined,
adaxially gently out-curving furrows that broaden distally before terminating prior to margin; anterior pair deepest and meet axial furrow;
other furrows may or may not reach axial furrow. Sharp-angled break between pleural field and flat, gently inclined border, which is similar in
width to distal pleural rib width.

Exoskeleton densely tuberculated; extensive covering on glabella and genal areas. On lateral cephalic border tubercles absent, although a few
short terrace ridges occur near the genal angle, running almost perpendicular to the border. Relatively large tubercles on eephalon set in a
groundmass of very fine tubercles. Tubercles on glabella with axes aligned at very low angle to glabellar surface. This angle increases posteriorly,
such that on posterior of glabella tubercle axes aligned close to perpendicular to surface. The consequence of this orientation is that in life position
the long axes of the tubercles would always be aligned vertically. On pygidium axis posterior to articulating furrow with relatively dense cover of
tubercles, although they are absent on small anterolateral lobes. Close to the sagittal line form a pair of gently curving exsagittal rows of
tubercles. Pleural ribs with relatively dense covering of turbercles; rows of single tubercles adaxially, increasing to five to six across the ribs
distally. Articulating half ring and facet both covered by fine terrace ridges. On facet die out laterally on anterolateral border. Rest of border
lacks tubercles.

Etymology.-From 'microphthalmus.' (Greek): provided with small eyes.

Types.-Holotype: WAM 96.479, cephalon, Calyx Corner, McWhae Ridge, Lawford Range, Canning Basin, Western Australia; Virgin Hills
Formation, Frasnian Zone 13a and b. Paratypes: WAM 96.474, 04.305-04.310.

Other material examined.-Twenty incomplete pygidia and librigenae from Phacopid Gully and Calyx Corner. McWhae Ridge, Lawford Range,
Canning Basin, Western Australia; Virgin Hills Formation, Frasnian Zone 13a and b.

Occurrence.-Late Frasnian, Zone 13a and b, late rhenana and early linguiformis zones.

Discussion.-Telopeltis microphthalmus differs from T. woodwardi n. sp. in its smaller size, the largest known pygidium being only slightly more
than half the size of the largest in T. woodwanli. The largest cephalon is 8.5 mm, compared with 13 mm in T. woodwardi. The frontal lobe is more
transverse anteriorly in T. microphthalmus, less tumid, and extends slightly more anterolaterally. The only difference in the glabellar furrows
of the two species is in S1, which is a little more incised in T. woodwardi and encloses a better-defined median tubercle. The glabellar
ornamentation is quite different in the two species, the dense concentration of tubercles across the entire frontal lobe in T. microphthalmus
contrasting with the transition from terrace ridges through to tubercles posteriorly across the glabella in T. woodwardi. Moreover, the very fine
secondary tubercles present in T. microphtluilmus are not evident in T. woodwardi. The main cephalic feature that distinguishes the two species

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is the palpebral lobe. Not only is it appreciably smaller in T. microphthalinus. with a correspondingly smaller eye surface consisting of irregular,
curving rows containing up to IO holochroal lenses, hut it is also much more anteriorly positioned. Consequently, the course of the posterior
branch of the facial suture is quite different between the two species. In T. woodward/, the suture extends transversely abaxially, before finally
recurving to the border, whereas in T.

microphthalmus the sutures are divergent posteriorly, each branch extending posterolaterally abaxially at about 45°, before recurving
adaxially close to the posterior border. Librigena of T. microphthalmux with lateral border with a thin, raised rim demarcated by broad, shallow
furrow. No genal spine is present.

The pygidia of the two species are similar, but that of T. woodward i is even more strongly vaulted (height 40%-5()% pygidial length in T.
microphthalmns, 60% in T. woodwurdi). Fulcrum in T. woodward! positioned slightly closer to the axial furrrow when viewed in transverse
profile. Well-developed terrace ridges are present on the facet in T. microphthulmits but not in T. woodwardi. As with the cephalon, the
tuberculation on the pygidium is different in the two species. Tubercles are more densely distributed on the pleural ribs in T. microphthalmiix.
and on the axis they are not only confined to a pair of exsagittal rows (although there are more in T. microphtlwlmus), but extend over the entire
surface of the axis, apart from the two lateral lobes. In both species some pleural furrows tail to meet the axial furrow in some specimens.
Moreover, in one specimen of T. microphthalmus the median rib is coalesced with the adjoining pleural rib adaxially (Fig. 5.14), a teratological
feature that has been previously recorded in scutelluines (Holloway, 1996. fig. 3.23).

Regarding the particular features of the reduced visual complex and related course of the posterior sutures, it is tempting to assign T.
microphthalmus to an independent new genus. However, paedomorphic evolution leading to eye reduction is a general phenomenon in
contemporaneous latest Frasnian trilobite lineages [i.e., Palpehralia lineage, Aculicryphops lineage (Feist, 1991; Crônier and Feist, 2004)],
whereas all other main diagnostic features (i.e., the particular shape of the pygidium in the case of Telopeltis n. gen.) remain unchanged.
Awaiting new evidence on diversity patterns of these last reduced eyed scutelluines we prefer to consider microphthalmux as a direct descendant
of woodwardi, both being congeneric.

PHYLOGENETlC AFFINITIES

Any assessment of the phylogenetic affinities of Telopeltis n. gen. must, of necessity, involve an appraisal of its likely derivation from Scutellum.
Given the highly conservative nature of this genus over a long time period (Selwood. 1966; Wright and Chatterton, 1988). the markedly
different morphological characteristics possessed by holaspids of Telopeltis compared with holaspids of Scutellum would seem to argue against a
close phylogenetic link between them: Scutellum is a typically flat scutelluine with larger eyes and long pygidium, whereas Telopeltis is small,
highly vaulted with a trend to eye reduction, and has a short pygidium.

In order to interpret the derivation from a seemingly unlikely ancestor, it is necessary to compare the morphological features of the holaspids of
Telopeltis (meraspids are unknown) with the ontogenetic development of other Devonian scutelluines. Fortunately, one of the best documented
scutelluine ontogenies is of a species of Scutellum, S. calruin. Another is of Dentaloxcutelluni hudxoni Chatterton, 1971. Both are from the Upper
Emsian of New South Wales. Australia. A third example is Meridioscutellum Feist, 1970 from the Lower Emsian of southern France. Chatterton
(1971) has noted that there are a number of similarities in the patterns of ontogenetic development of these genera, indicating that broad
ontogenetic trends in morphological change may have been common among scutelluines. When the holaspids of the two species of Telopeltix are
compared with early developmental stages of these scutelluines. a number of similarities are apparent.

One of the characteristic trends in scutelluine ontogenetic development is for the prolaspid and early meraspids to be highly vaulted, but for this
vaulting to diminish markedly through to, and during, the holaspid stage. Other trends include a posterior migration of the eye in early
development: the appearance and subsequent deepening of the glabellar furrows-these are hardly present at all in early to midmeraspids; a
reduction in length of the occipital spine in meraspids, such that it is only represented by an occipital tubercle in holaspids; and a pronounced
elongation and flattening of the pygidium during development. Even during holaspid ontogenetic development of T. micmphthalmus n. sp., an
appreciable relative lengthening of the pygidium still occurs.

All of the major features that characterize the holaspids of Telopeltix are those which are present in the meraspid stages of Scutellum.
Dentaloxcntelhim Chatterton, 1971 and Meridio.icutelhwt. Given that Telopeltix postdates these genera, this indicates that many of its
diagnostic features arose by paedomorphosis. Six morphological features present in the holaspids of T. wiwdwardi n. sp. are paedomorphic, while
in the last scutelluine, T. microphthalmux, there are nine. The paedomorphic traits present in both species of Telopeltix are: a moderately
vaulted cephalon; ill-defined S2 and S3 furrows; very weak lateral occipital lobes; the retention of an occipital spine (in a 6 mm long holaspid this
is of equivalent length to that of a late stage meraspid of Scutellum c(ilvnm); and a very short, broad, and strongly vaulted pygidium. While the
holaspid pygidium of S. calvitm is almost flat, it is highly vaulted in the meraspid stage (Chatterton. 1971, pi. 5. figs. 3, 4, 8); the pygidial
length/breadth dimensions of early meraspids are also very close to those of holaspids of Telopeltix.

Other paedomorphic traits only possessed by T. micmphthalmus are an almost transverse frontal lobe (compare with Chatterton, 1971. fig. 6: pi.
5. figs. 11, 14, 18); a very poorly defined Sl; and an anteriorly positioned eye lobe (compare with Chatterton, 1971, fig. 6, pi. 5, figs. 2. II).

The evolution of paedomorphic features in Telopeltis and the increase in number of paedomorphic traits in the later species of Telopeltis suggest
that the environmental conditions prior to each of the Kellwasser events were selecting for paedomorphic morphologies. Paedomorphosis has been
recorded in other late Frasnian taxa, where a trend to paedomorphic eye reduction occurred independently in a number of families in association

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with the Kellwasser crises (Feist, 2002). While it can be dangerous to speculate on growth rates in extinct organisms, it could be argued that the
combination of paedomorphic features and small body size may indicate that the paedomorphic process was progenesis. As this results from early
sexual maturation, it is often a feature of selection in unstable, stressed environments, where selection targets early maturation and rapid
reproduction. Such may have been the case prior to the Kellwasser events, when environmental stress was caused by periods of oxygen depletion
on the shelf.

ACKNOWLEDGMENTS

We wish to thank G. Rokylle and E. Routasuo for initially finding the trilobite-bearing horizons, assisting one of us (KJM) to get to the localities,
and for their help collecting material. We wish to thank others who helped to collect material used in this study: J. Long, D. Friend, C. McGeachie,
D. Haig, T. Becker, M. House. R Jell. R Playford, and R. Lerosey-Aubril. A. George and K. Trinjastic are thanked for assistance in the field. We are
grateful to D. Holloway and B. Chatterton for their constructive comments, which improved the manuscript. U. Lemke and M. Basse are
acknowledged for kindly providing photos and a latex cast of the holotype of Frasniellum. G. Klapper kindly provided conodont data for
biostratigraphy. A. and J. Henwood are thanked for providing access to Fossil Downs Station. KJM is grateful for funding from the French
Government (EGIDE) that allowed him to undertake this research in Montpellier. Funding for RF to visit the field was provided by the French
National Centre for Scientific Research (CNRS). This is a contribution to ISEM, UMR 5554 CNRS (2005-016).

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ACCEPTED 5 JUNE 2005

KENNETH J. McNAMARA1 AND RAIMUND FEIST2

1 Department of Earth & Planetary Sciences. Western Australian Museum, Francis Street, Perth, Western Australia 6000, Australia, and 2
Institut des Sciences de l'Evolution. Laboratoire de Paléontologie, Université Montpellier II, Place E. Bataillon, 34095 Montpellier Cedex 5, France.

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