Geobios 35 (2002) 111–125

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Hoplolichoides, Allolichas, Autoloxolichas and Akantharges,

and the classification of lichid trilobites
Hoplolichoides, Allolichas, Autoloxolichas et Akantharges,
et la classification des trilobites Lichidae
David J. Holloway a,*, Alan T. Thomas b
a
Invertebrate Palaeontology, Museum Victoria, PO Box 666E, Melbourne, Victoria 3001, Australia
b
School of Earth Sciences, University of Birmingham, Edgbaston, Birmingham, B15 2TT, United Kingdom
Received 9 November 2000; accepted 24 July 2001

Abstract

Hoplolichoides PHLEGER, 1936 and Autoloxolichas PHLEGER, 1936 are revised based on redescription of their type species,
H. conicotuberculatus (NIESZKOWSKI, 1859) and A. sanctamathiae (SCHMIDT, 1885) respectively. Hoplolichoides is considered to be
most closely related to Hoplolichas DAMES, 1877, and both are included in a revised Homolichinae. Autoloxolichas is restricted to its type
species, and other species previously assigned to the genus are placed in Allolichas KRUEGER, 1992, the concept of which is expanded.
Autoloxolichas and Allolichas are included together with three other genera and subgenera in the Subfamily Platylichinae, previously
considered to be a junior synonym of Homolichinae. The hypostomal morphology previously regarded as important in uniting homolichine
and platylichine genera is now regarded as primitive for the Lichidae. Metopolichas GÜRICH, 1901, previously included tentatively in the
Homolichinae on the basis of its hypostomal morphology, is reassigned tentatively to the Lichinae. The trochurine ‘Acanthopyge’ erbeni
MEISCHNER, 1965, from the Givetian of Germany, is assigned to Akantharges PHLEGER, 1936, permitting clarification of the pygidial
morphology of the genus. Akantharges and Ceratarges GÜRICH, 1901 are unique amongst lichids in having a curved, transverse ridge on
the posterolateral cranidial lobe, suggesting that these genera may have been derived from a common ancestor. © 2002 Éditions
scientifiques et médicales Elsevier SAS. All rights reserved.

Résumé

Les genres Hoplolichoides PHLEGER, 1936 et Autoloxolichas PHLEGER, 1936 sont révisés sur la base d’une redescription de leurs
espèces types respectives: H. conicotuberculatus (NIESZKOWSKI, 1859) et A. sanctamathiae (SCHMIDT, 1885). Hoplolichoides est
considéré comme le genre le plus proche de Hoplolichas DAMES, 1877. Les deux genres sont inclus dans les Homolichinae également
révisés. Autoloxolichas ne contient que son espèce-type et d’autres espèces rapportées aurapavant à Autoloxolichas sont placées dans
Allolichas KRUEGER, 1992 dont la définition est ainsi étendue. Autoloxolichas et Allolichas sont inclus avec trois autres genres et
sous-genres dans la sous-famille des Platylichinae que l’on considérait avant comme un synonyme des Homolichinae. La morphologie de
l’hypostome considérée autrefois comme importante (unissant les genres homolichines et platylichines) est à présent considérée comme
primitive pour les Lichinae. Metopolichas GÜRICH, 1901 qui avait été placé dans les Homolichinae sur la base de sa morphologie
hypostomale est placé dans les Lichinae. Le trochuriné ‘Acanthopyge’ erbeni MEISCHNER, 1965 du Givétien de l’Allemagne est attribué
à Akantharges PHLEGER, 1936 permettant une classification de la morphologie pygidiale du genre. Akantharges et Ceratarges GÜRICH,

* Corresponding author.
E-mail addresses: dhollow@museum.vic.gov.au (D.J. Holloway), a.t.thomas@bham.ac.uk (A.T. Thomas).

© 2002 Éditions scientifiques et médicales Elsevier SAS. All rights reserved.

PII: S 0 0 1 6 - 6 9 9 5 ( 0 2 ) 0 0 0 1 4 - 1
112 D.J. Holloway, A.T. Thomas / Geobios 35 (2002) 111–125
1901 sont uniques chez les lichidés par leur crête transversale courbe sur le lobe cranidial postéro-latéral, suggérant que ces deux genres
dérivent d’un ancêtre commun. © 2002 Éditions scientifiques et médicales Elsevier SAS. Tous droits réservés.
Keywords: Trilobites; Lichidae; Systematics; Ordovician; Devonian
Mots clés: Trilobites; Lichidae; Systématique; Ordovicien; Dévonien
1. Introduction
The trilobite genera Hoplolichoides, Autoloxolichas and
Akantharges, all erected by Phleger (1936), were reviewed
by Thomas and Holloway (1988) in their revision of the
Lichida. Additional information on these genera has become
available subsequently, providing the opportunity to clarify
their taxonomic concepts as well as to assess their likely
relationships. In the light of our new data, we assign
Hoplolichoides to a revised Homolichinae. The Subfamily
Platylichinae is resurrected to accommodate Allolichas and
Autoloxolichas, together with other genera.
Hoplolichoides has not generally been accepted as a
distinct genus, most authors (Warburg, 1939; Hupé, 1953;
Tripp, 1957, 1959; Balashova, 1960) considering it to be a
junior subjective synonym of Hoplolichas DAMES, 1877.
Thomas and Holloway (1988) recognised Hoplolichoides
but expressed reservations about its utility as an indepen-
dent genus, stating that the pygidial morphology was
insufficiently known for a full comparison with other
genera. In the present paper, we redescribe cephala and
pygidia of the type species, H. conicotuberculatus (NIESZ-
KOWSKI, 1859), and compare it with other species con-
sidered to be congeneric, showing that Hoplolichoides
warrants recognition as a separate genus.
Autoloxolichas was based on Lichas sanctamathiae
SCHMIDT, 1885, and that was the only species listed by
Phleger (1937) as belonging to the genus. The genus was
briefly diagnosed by Phleger (1936) as differing from
Platylichas in lacking the continuation of the longitudinal
furrow behind the bullar lobe, but Öpik (1937) dismissed
this supposed difference as being due to a misunderstanding
and placed sanctamathiae in Platylichas. Tripp (1957,
1959) and Dean (1974) also considered Autoloxolichas to be
a junior synonym of Platylichas. Thomas and Holloway
(1988) recognised Autoloxolichas as a distinct genus to
which they assigned, in addition to sanctamathiae, a num-
ber of species resembling ‘Lichas’ laxatus MCCOY, 1846.
We now believe, based on the study of plaster replicas of the
type material of sanctamathiae, that Autoloxolichas should
be restricted to that species, and that the other species
included in the genus by Thomas and Holloway (1988)
should be placed in Allolichas KRUEGER, 1992, the
concept of which should be expanded.
Akantharges was until relatively recently known only
from the type species, Lichas gourdoni BARROIS, 1886,
the material of which is strongly deformed and difficult to
interpret. A well preserved cranidium of an undescribed
species of Akantharges from the Devonian of Morocco was
illustrated by Thomas and Holloway (1988, pl. 14, figs 299,
200, 303), who based their concept of the genus largely on
that specimen. We have subsequently been able to examine
the material of ‘Acanthopyge’ erbeni MEISCHNER, 1965
from the Givetian of Germany, and recognised that this
species belongs to Akantharges. The specimens of erbeni
are relatively well preserved and clarify some of the
characters of the genus, particularly pygidial characters.
2. Systematic palaeontology
Morphological terminology used here is that of Thomas
and Holloway (1988). Stratigraphical nomenclature follows
the revised Ordovician series proposed by Fortey et al.
(1995, 2000).
Several of the species dealt with herein were documented
by Schmidt (1885). Until recently, the specimens from this
and other parts of Schmidt’s series of trilobite monographs
were not available for study. Photographs of most of
Schmidt’s specimens are now available on the World Wide
Web at the site entitled ‘Catalogue of the trilobites figured in
Friedrich Schmidt’s ‘Revision der ostbaltischen silurischen
Trilobiten’ (1881–1907)’ by D.L. Bruton, O.A. Hoel,
L.T. Beyene and A. Yu. Ivantsov (internet address
http://www.toyen.uio.no/palmus/schmidt/index.html). The
specimens are also listed but not illustrated in Bruton et al.
(1997).
Repositories of specimens are denoted by the following
abbreviations: CNIGR, Central Scientific-Research Geo-
logical Exploration Museum named after F.N. Chernyshev,
St Petersburg; GIE, Institute of Geology, Tallinn Technical
University; Gö, Geologisch-Paläontologishes Institut und
Museum, Georg-August-Universität, Göttingen; MB, Mu-
seum für Naturkunde, Humboldt Universität, Berlin; RM,
Naturhistoriska Riksmuseet, Stockholm.
Family: LICHIDAE Hawle & Corda, 1847
Subfamily: HOMOLICHINAE Phleger, 1936
Diagnosis – S1 effaced or almost effaced, so that bullar
lobe is more or less completely fused with L1b; resulting
composite lobe circumscribed by furrows, except in Leioli-
chas which has longitudinal furrow effaced externally.
Palpebral lobe narrow, of almost uniform width throughout,
moderately curved in outline. Hypostome with middle body
as wide across posterior lobe as across anterior lobe, or
wider; shoulder situated level with posterior border furrow;
border greatly expanded and flap-like at and behind
 D.J. Holloway, A.T. Thomas / Geobios 35 (2002) 111–125 113

shoulder. Pygidium with two to three pairs of pleural

furrows and six to eight marginal spines (spines absent in
Leiolichas); anterior and posterior pleural bands flattened.
Genera included – Homolichas SCHMIDT, 1885;
Conolichas DAMES, 1877; Hoplolichas DAMES, 1877;
Hoplolichoides PHLEGER, 1936; Leiolichas SCHMIDT,
1885; Otarozoum THOMAS & HOLLOWAY, 1988.
Remarks – S1 is effaced, or almost so, in Conolichas,
Hoplolichas, Hoplolichoides and Otarozoum, the bullar lobe
almost completely fusing with L1b. The resulting composite
lobe is circumscribed by the longitudinal and axial furrows,
and by the transverse furrow in front of L1a (Fig. 1.1–4).
The longitudinal furrow is effaced externally in Leiolichas
but is present on internal moulds, though very shallow
posteriorly (see Warburg, 1939, pl. 3, figs 7, 8). On some
internal moulds of Leiolichas cranidia, S1 is also present but
very weak. It meets the axial furrow just behind the
intersection of the latter with the palpebral furrow, and is
directed strongly obliquely backwards towards the point at
which the longitudinal furrow abruptly shallows (Fig. 1.5).
This glabellar structure is different from that in genera such
as Platylichas, Autoloxolichas and Metalichas, which were
previously included in the Homolichinae (Tripp, 1957,
1959; Thomas and Holloway, 1988), but which we now Fig. 1. Cranidial morphology in Homolichinae (1-5) and Platylichinae
place in the Platylichinae (see below). In these genera, the (6-8). 1. Conolichas (after Thomas and Holloway 1988, pl. 7, fig. 141). 2.
Hoplolichas (after Thomas and Holloway 1988, pl. 8, fig. 155). 3.
axial furrow is effaced alongside L1b, and the bullar lobe is Hoplolichoides (after Fig. 2.6). 4. Otarozoum (after Thomas and Holloway
circumscribed by the longitudinal furrow, S1, and the short 1988, pl. 8, fig. 161). 5. Leiolichas (after Thomas and Holloway 1988, pl.
section of axial furrow between the anterior end of the 7, fig. 147). 6. Platylichas (after Warburg 1939, pl. 11, fig. 1c). 7.
longitudinal furrow and the outer end of S1 (Fig. 1.6–7). Autoloxolichas (after Fig. 3.1 and Thomas and Holloway 1988, pl. 10, fig.
213). 8. Allolichas (after Thomas and Holloway 1988, pl. 10, fig. 188 and
Metopolichas GÜRICH, 1901 was tentatively included in
Krueger 1992, pl. 1, fig. 1b). Dashed lines indicate furrows that are very
the Homolichinae by Thomas and Holloway (1988) by weakly developed, or present only on internal moulds; see text for details.
weighting its hypostomal morphology, despite the fact that Non-standard abbreviations: af, axial furrow; b, bullar lobe; cl, composite
it resembles Lichas and some other Lichinae in glabellar lobe; lf, longitudinal furrow; of, occipital furrow.
structure (longitudinal furrow not generally extending be- Morphologie du cranidium chez les Homolichinae et les Platylichinae. 1.
Conolichas (d’après Thomas et Holloway 1988, pl. 7, fig. 141). 2.
hind S1, with which it is united in a curve; S1 commonly
Hoplolichas (d’après Thomas et Holloway 1988, pl. 8, fig. 155). 3.
incomplete abaxially). The hypostome of Metopolichas Hoplolichoides (d’après Fig. 2.6). 4. Otarozoum (d’après Thomas et
resembles those of Homolichinae and some Platylichinae, Holloway 1988, pl. 8, fig. 161). 5. Leiolichas (d’après Thomas et Holloway
rather than Lichinae, in being wider than long and having a 1988, pl. 7, fig. 147). 6. Platylichas (d’après Warburg 1939, pl. 11, fig. 1c).
deep posterior border furrow (see Thomas and Holloway, 7. Autoloxolichas (d’après Fig. 3.1 et Thomas et Holloway 1988, pl. 10, fig.
213). 8. Allolichas (d’après Thomas et Holloway 1988, pl. 10, fig. 188 et
1988, pl. 11, figs 227, 231, 235). However, hypostomes of
Krueger 1992, pl. 1, fig. 1b). Les hachures indiquent des sillons très
Tetralichinae (e.g. Thomas and Holloway, 1988, pl. 12, figs faiblement développés ou présents seulement sur les moules internes.
250, 267) and Trochurinae (e.g. Chatterton and Ludvigsen, Abbréviations non standards: af, sillon axial; b, lobe bulleux; cl, lobe
1976, pl. 19, figs 17–20) also share these character states, composé; lf, sillon longitudinal; of, sillon occipital.
which we now consider primitive for lichids as a whole and
thus not useful for inferring relationships within the family. or wider, instead of narrower; the border is more expanded
The interpretation of these character states as primitive is in width at the shoulder, where it has a flap-like appearance;
supported by their presence in the oldest known lichid, and the shoulder is situated somewhat farther back, opposite
Metopolichas? klouceki (RŮŽIČKA, 1926) from the lower the posterior border furrow rather than opposite the middle
Tremadoc of the Czech Republic (see Thomas and Hollo- furrow or the posterior lobe of the middle body (compare
way, 1988, pl. 11, fig. 236). In view of the similarities in Thomas and Holloway, 1988, pl. 10, fig. 196 and Fig. 3.4
glabellar structure between Metopolichas and some Lichi- herein).
nae we now assign the genus to that subfamily. In comparing the thoracic and pygidial structure of
Notwithstanding their similarity in the primitive charac- lichines and Platylichas, Whittington (in press) noted that in
ter states mentioned above, hypostomes of Homolichinae both the inner parts of the thoracic pleurae are wide and
differ from those of Platylichinae in that the middle body is evenly inflated, and the pleural furrows extend almost to the
as wide across the posterior lobe as across the anterior lobe, pleural tips. The pygidial pleural bands in these taxa are of
114 D.J. Holloway, A.T. Thomas / Geobios 35 (2002) 111–125
similarly low convexity, are divided by the pleural furrow,
and are extended distally into the marginal spines. This kind
of thoracic and pygidial structure occurs in homolichines
also (e.g. Otarozoum, see Thomas and Holloway, 1988, pl.
8, figs 167–170) and, by comparison with other trilobites,
probably represents the primitive condition for the Lichidae.
The pygidial pleural bands of Hoplolichoides are more
inflated than is usual for a member of these subfamilies, and
the genus is derived in that regard.
Genus Hoplolichoides PHLEGER, 1936
Type species – Lichas conico-tuberculata NIESZ-
KOWSKI, 1859, from the Kukruse Stage (C2, lower Cara-
doc) of Estonia; original designation.
Other species – H. curvifrons (WARBURG, 1939); H.
furcifer (SCHMIDT, 1885).
Stratigraphical and geographical range – Middle
Llanvirn–lower Caradoc; Russia (St Petersburg district),
Estonia, Sweden, Norway.
Diagnosis – Homolichine having glabella with L1b in-
distinctly differentiated from bullar lobe by slight backward
deflection and/or shallowing of longitudinal furrow, and
very weak S1. Frontomedian and bullar lobes moderately
convex (sag., exsag.), frontal lobe slightly overhanging
medial part of anterior border in palpebral view; bullar lobe
with long axis subparallel to sagittal line; minimum width of
median lobe less than width of bullar lobe measured across
same transverse line. Occipital ring strongly elevated medi-
ally, with large paired spines. Cranidial tuberculation very
coarse. Pygidium transverse, approximately 160% as wide
as long (sag.), with three pairs of marginal spines and small
posteromedian spine; third pair of spines very short and
broad, subtriangular or lobate, situated very close to second
pair of spines. Axis approximately 30% maximum pygidial
width anteriorly and 50% sagittal length. Pleurae with
prominent border that is transverse between second pair of
spines; first two pleural furrows and furrow defining post-
axial band terminating at inner edge of border; third pleural
furrow not defined; anterior and posterior pleural bands well
rounded (exsag.).
Remarks – Large paired occipital spines are preserved
only in the type species, the occipital ring being incomplete
posteriorly in known specimens of the other two species
assigned to the genus. We would not exclude a species from
Hoplolichoides solely on the absence of these spines.
Hoplolichoides shows the greatest similarities with
Hoplolichas and Otarozoum. The similarities with Hoploli-
chas include the presence of large occipital spines (unpaired
in Hoplolichas), the weak differentiation of L1b by a
backward deflection of the longitudinal furrow and a faint
S1, the very coarse cranidial tuberculation, the absence of
the third pygidial pleural furrow, and the well developed
border on the posterior part of the pygidium. Pygidia of
some species of Hoplolichas also resemble Hoplolichoides
pygidia in having a very short, broad marginal spine situated
immediately behind the second spine, and/or an unpaired
posteromedian spine (e.g. Warburg, 1939, pl. 8, fig. 5;
Neben and Krueger, 1971, pl. 35, figs 15, 17). Of these
similarities, we do not attribute great taxonomic importance
to the presence or absence of occipital spines, while the
weak differentiation of L1b may be a primitive character
state, compared with the condition in Conolichas and
Otarozoum (in which L1b is indistinguishable), and thus not
indicative of close relationship. On the basis of the pygidial
similarities, however, we consider Hoplolichas to be most
closely related to Hoplolichoides. Hoplolichas differs from
Hoplolichoides in that the glabella is more convex (sag.,
exsag.), the frontal lobe being more inflated and markedly
overhanging the anterior border in palpebral view; the
minimum width of the median lobe is greater (rather than
less) than the width of the bullar lobe measured across the
same transverse line; and the pygidium is not as transverse.
In addition, pygidia of Hoplolichas species typically have
much longer marginal spines, there is no spine situated
immediately behind the second spine, and the posterior pair
of spines are situated very close together, commonly being
fused proximally (e.g. Thomas and Holloway, 1988, pl. 8,
fig. 162).
Otarozoum resembles Hoplolichoides in the moderate
cranidial convexity and the proportions of the glabellar
lobes, but differs in the absence of large occipital spines;
L1b is not differentiated at all from the bullar lobe; the
cranidial tuberculation is much finer; there is no pygidial
border; the third pygidial pleural furrow is impressed; the
third pair of pygidial marginal spines is situated very close
together; and the pygidium has no posteromedian spine. The
pygidium of Otarozoum is very similar to that of Conoli-
chas, suggesting that these genera are most closely related
to each other.
Apart from conicotuberculatus and furcifer, Phleger
(1937) also included in Hoplolichoides the species Lichas
(Hoplolichas) plautini SCHMIDT, 1885, L. (H.) longispina
[sic] SCHMIDT, 1885 and Lichas media [sic] POMPECKI,
1890. The last two belong to Hoplolichas, as indicated by
the proportions of the glabellar lobes, the strongly over-
hanging frontal lobe and the large, unpaired occipital spine
in the illustrated cranidium of medius (Pompecki 1890, pl.
2, fig. 25, 25a), and the three pairs of long marginal spines,
the posteriormost pair situated fairly close together, in the
holotype pygidium of longispinus (Schmidt 1885, pl. 2, fig.
25). In plautini, Schmidt (1885, pl. 2, figs 17–24) associated
cranidia and an incomplete cephalon of Hoplolichas type
with pygidia having the morphology of Hoplolichoides. We
consider these pygidia to belong to Hoplolichoides furcifer,
a species based on cranidia from the same locality and
horizon as plautini. Schmidt chose to assign the pygidia to
plautini rather than to furcifer because one of the pygidia is
preserved beneath an incomplete cephalon of plautini.
However, examination of a plaster replica of this specimen
(CNIGR 36/11101) revealed that the pygidium is not in
enrolled position beneath the cephalon, and consequently
there is no reason to conclude that both cephalon and
pygidium belong to the same individual as believed by
 D.J. Holloway, A.T. Thomas / Geobios 35 (2002) 111–125
Schmidt. In addition, the pygidium is so incomplete (more
so than shown by Schmidt 1885, pl. 2, fig. 17b) that it is not
possible to determine whether it belongs to Hoplolichoides
or Hoplolichas. The fragmentary pygidium later assigned to
furcifer by Schmidt (1907, pl. 2, fig. 9) resembles the
pygidia previously assigned erroneously to plautini.
Hoplolichoides conicotuberculatus (NIESZKOWSKI,
1859)
Fig. 2.1–19
1859 Lichas conico-tuberculata NIESZKOWSKI, p.
365, pl. 1, figs 7–10.
1877 Lichas (Hoplolichas) conico-tuberculata NIESZ-
KOWSKI - Dames, p. 802, pl. 14, figs 2–6.
1885 Lichas (Hoplolichas) conicotuberculata NIESZ-
KOWSKI - Schmidt, p. 82, pl. 3, figs 13–25.
1901 Hoplolichas conicotuberculatus (NIESZKOWSKI)
- Gürich, pl. 20, fig. 14.
1928 Lichas (Hoplolichas) conicotuberculatus NIESZ-
KOWSKI - Kummerow, p. 36, pl. 2, fig. 5.
1936 Hoplolichoides conicotuberculatus
(NIEZSKOWSKI) - Phleger, p. 602, fig. 23.
1937 Hoplolichoides conicotuberculatus (NIESZ-
KOWSKI) - Phleger, p. 1087.
1939 Hoplolichas conicotuberculatus (NIESZKOWSKI)
- Warburg, p. 97, pl. 7, fig. 2.
1957 Hoplolichas conicotuberculatus (NIESZKOWSKI)
- Tripp, p. 108, text-fig. 4O; p, 110, text-fig. 5C.
1958 Hoplolichas conicotuberculatus (NIEZSKOWSKI)
- Tripp, p. 576.
1971 Hoplolichas conicotuberculatus (NIESZKOWSKI)
- Neben and Krueger, pl. 34, figs 7–9.
1988 Hoplolichas conicotuberculatus (NIESZKOWSKI)
- Thomas and Holloway, pl. 10, figs 193–197.
Lectotype – Selected by Warburg (1939); cranidium
figured by Nieszkowski (1859, pl. 1, figs 7, 8), from the
Kukruse Stage (C2, lower Caradoc) at Erras (= Erra) or
Wannamois (= Vanamõisa), Estonia. The records of the
Institute of Geology, Tallinn Technical University, question-
ably identify the lectotype as specimen no. Tr2099 (figured
Thomas and Holloway, 1988, pl. 10, fig. 195). That appears
unlikely as the specimen does not match Nieszkowki’s
illustration (e.g. the posterior borders and occipital ring,
excluding the occipital spines, are shown in the illustration
but are mostly covered by matrix in the specimen).
Material examined Five cranidia (MB T1809–T1811,
T1814, T1815), two librigenae (MB T1821, T1822) and
four pygidia (MB T1812, T1813, T1819, T1820), all from
Kukruse near Jõhvi, Estonia.
Diagnosis – Minimum width of median glabellar lobe
situated level with intersection of axial and palpebral
furrows; L1a transverse, ovate. Pygidium with first two
marginal spines steeply inclined across their width, con-
verging backwards; third spine subtriangular.
Description – Glabella approximately equal in width
across occipital ring and L1a, expanding weakly forwards
from front of L1a to front of palpebral lobe, thereafter
115
narrowing at increasing rate to front of bullar lobe; width of
glabella at front of palpebral lobe 140% width across frontal
lobe and approximately equal to sagittal length (excluding
occipital ring) in palpebral view (with palpebral suture
horizontal). Occipital ring increasing strongly in length
(exsag.) and height adaxially to base of occipital spines;
median part of ring steep and slightly concave (sag.) in front
of spines (Fig. 2.1). Occipital spines stout, diverging back-
wards at about 60° proximally, gently curved backwards in
lateral and dorsal profiles. Large median occipital tubercle
present in front of spines, and a pair of slightly smaller
tubercles close to posterior edge of ring behind L1a. Median
part of occipital furrow shallow, transverse medially; lateral
part deeply incised and deflected backwards behind L1a.
Glabella in front of occipital ring more convex sagittally
than transversely, in palpebral view descending forwards in
front of δ-δ and overhanging medial part of anterior border;
longitudinal furrow deepest anteriorly, meeting occipital
furrow posteriorly in line (exsag.) with adaxial extremity of
L1a. L1a with two prominent tubercles situated slightly
anteromedially and posterolaterally of centre; furrow in
front of L1a collinear with median part of occipital furrow
but shorter (exsag.) and deeper. L1b indistinctly defined by
slight backward deflection of longitudinal furrow and by
faint transverse depression separating it from bullar lobe
(Fig. 2.6); L1b with large tubercle adaxially. Bullar lobe
widest at midlength in palpebral view, in front of midlength
partially subdivided adaxially (most distinctly on internal
moulds) by very narrow (tr.), shallow, transverse S2; posi-
tion of S2 commonly indicated on exterior of exoskeleton
only by weak abaxial deflection and shallowing of longitu-
dinal furrow (Fig. 2.10). Posterior half of median lobe with
fairly regular arrangement of coarse, paired tubercles in
three rows; posterior two rows separated on internal mould
by weak transverse depression (S1) in line (exsag.) with
front of L1b (Fig. 2.6).
Axial furrow deep, narrow; preglabellar furrow beco-
ming more sharply impressed adaxially. Anterior border
flattened (sag., exsag.), decreasing in length adaxially, its
margin gently concave in outline abaxially. Fixigenal field
steeply declined in front of and behind palpebral lobe (Fig.
2.1), overhanging posterior border adaxially in palpebral
view. Posterior border increasing in height and length
(exsag.) abaxially, its posterior edge curving gently back-
wards; posterior border furrow deep, curving slightly for-
wards abaxially. Palpebral lobe narrow (tr.), tapering more
strongly posteriorly than anteriorly, posterior edge situated
opposite front of L1b in palpebral view (Fig. 2.6, 10) and
slightly farther from sagittal line than anterior edge; palpe-
bral furrow strongly curved, shallow at midlength (exsag.);
palpebral area inflated above level of palpebral lobe (Fig.
2.1). Anterior branch of facial suture initially diverging
slightly in front of palpebral lobe, thereafter converging
strongly to lateral border furrow, diverging less strongly
forwards across anterior border; width α-α almost equal to
maximum width of glabella. Posterior branch of suture
116 D.J. Holloway, A.T. Thomas / Geobios 35 (2002) 111–125

Fig. 2. Hoplolichoides conicotuberculatus (NIESZKOWSKI, 1859), Kukruse Stage (C2, lower Caradoc), Kukruse, Estonia. 1, 5-6 MB T1810, cranidium,
figured Dames (1877, pl. 14, fig. 3, 3a); lateral, oblique and palpebral views, × 1.75. 2-3 MB T1809, cranidium, figured Dames (1877, pl. 14, fig. 2); plan
and oblique views, × 2. 4, 8, 12-13 MB T1812, pygidium, figured Dames (1877, pl. 14, fig. 5); lateral, oblique, posterodorsal and dorsal views, × 2. 7. MB
T1821, librigena, collected Dames 1877; dorsal view, × 2. 9, 14. MB T1822, librigena, collected Dames; dorsal and ventral views, × 2. 10-11 MB T1814,
cranidium, collected Dames 1877; palpebral and oblique views, × 1.75. 15-16 MB T1820, pygidium, collected Dames 1876; posterodorsal and dorsal views,
× 2. 17. MB T1815, fragmentary cranidium, collected Dames 1876; plan view, × 2. 18-19 MB T1813, pygidium, figured Dames (1877, pl. 14, fig. 6); dorsal
view, × 2; marginal spines in dorsolateral view, × 3.
Hoplolichoides conicotuberculatus (NIESZKOWSKI, 1859), Kukruse Stage (C2, Caradoc inférieur), Kukruse, Estonia. 1, 5-6 MB T1810, cranidium, Dames
(1877, pl. 14, fig. 3, 3a); vues latérale, oblique et palpébrale, × 1.75. 2-3 MB T1809, cranidium, Dames (1877, pl. 14, fig. 2); vues plane et oblique, × 2.
4, 8,12-13 MB T1812, pygidium, Dames (1877, pl. 14, fig. 5); vues latérale, oblique, postérodorsale et dorsale, × 2. 7. MB T1821, librigena, collecté par
Dames 1877; vue dorsale, × 2. 9, 14. MB T1822, librigena, collecté par Dames; vues dorsale et ventrale, × 2. 10-11 MB T1814, cranidium, collecté par Dames
1877; vues palpébrale et oblique, × 1.75. 15-16 MB T1820, pygidium, collecté par Dames 1876; vues postérodorsale et dorsale, × 2. 17. MB T1815, fragment
de cranidium, collecté par Dames 1876; vue plane, × 2. 18-19 MB T1813, pygidium, figuré par Dames (1877, pl. 14, fig. 6); vue dorsale, × 2; vue dorsolatérale
et épine marginale, × 3.
 D.J. Holloway, A.T. Thomas / Geobios 35 (2002) 111–125
curving strongly backwards abaxially, converging back-
wards slightly across posterior half of posterior border.
Librigena with gently convex borders merging with base
of long, broad, flattened, sickle-shaped genal spine; lateral
border expands greatly in width posteriorly, outer margin
with shallow antennal notch anteromedially. Borders sepa-
rated from more convex librigenal field by shallow border
furrows that unite in a uniform curve (Schmidt, 1885, pl. 3,
fig. 13b, 19); border furrows preserved in MB T1822 but
most of field missing (Fig. 2.9). Numerous small spines
present along adaxial edge of genal spine. Librigenal
doublure broad, its inner edge corresponding approximately
in position with border furrows.
Pygidium with maximum width across base of first
marginal spine and level with pygidial midlength. Axis
strongly convex (tr.), tapering gently backwards, poorly
defined posteriorly; transverse row of prominent tubercles
situated at change of slope between flattened (sag.) crest of
axis and steeply inclined postaxial region (Fig. 2.4). Articu-
lating half ring short (sag., exsag.), of uniform length over
most of axial width. First axial ring only slightly longer
sagittally than articulating half ring, posterior edge with
weak embayment medially to accommodate poorly defined
pseudo-articulating half ring on second axial ring; second
ring longer sagittally than first ring, decreasing slightly in
length laterally; second ring furrow very shallow medially.
First and second rings each with a transverse row of six
tubercles; third ring defined only by alignment of tubercles.
Furrow alongside postaxial region collinear with axial
furrow, becoming broad, shallow and poorly defined poste-
riorly at intersection with border furrow. Pleural field
convex (tr.), steeply downturned abaxially; posterior pleural
bands more prominent than anterior bands, extending onto
border distally. Each posterior band bearing a row of four
prominent tubercles; second tubercle (from adaxial end)
very high and spinose, situated about half way between
axial and border furrows; most distal tubercle situated on
border. Pleural and interpleural furrows deep on pleural
field, shallower on border where pleural furrows are de-
flected laterally to reach margin between spines. Posterior
part of pleural field with two or three prominent tubercles
arranged in a line directed slightly more strongly backwards
than second interpleural furrow. First marginal spine termi-
nating just behind transverse line through tip of notch
between second and third spines; second spine broader and
converging more strongly backwards than first, terminating
level with tip of posteromedian spine; third spine terminat-
ing in front of transverse line through tips of second and
posteromedian spines; posteromedian spine becoming
rounded in cross section distally. Border wider between
third spines than in front of third spines; border furrow
shallow behind postaxial region, even shallower where it
crosses posterior pleural bands on first two segments.
Doublure convex.
Dorsal surface of cranidium and pygidium with dense
sculpture of coarse tubercles together with superimposed
117
and interspersed granules; tubercles on occipital spines
point towards distal ends of spines. Coarse tubercles less
abundant on librigena than on cranidium. Most of librigenal
doublure with two sets of terrace ridges: a low, widely
spaced set running subparallel to inner margin, and an
intervening, very fine, anastomosing set; librigenal margin
and outer part of doublure with coarser, closely spaced,
subparallel terrace ridges. Coarse terrace ridges on pygidial
doublure extend onto dorsal surface along (mainly abaxial)
edges of first two spines, where they run oblique to margin
and bear a row of fine granules (Fig. 2.19).
Discussion – Schmidt (1885) recorded conicotubercula-
tus in Estonia from the Kukruse Stage (C2) at Kuckers
(= Kukruse), Erras (= Erra), Wannamois near Tolks (= Van-
amõisa near Kohala), and Kawast (Kavastu near Rakvere),
and from the uppermost part of the underlying Echino-
sphaerites Limestone (Uhaku Stage, C1c) at Reval
(= Tallinn). From the last locality he illustrated an incom-
plete cephalon with several attached thoracic segments
(Schmidt 1885, pl. 3, fig. 13a–c), the cephalon being
depicted with a single median occipital spine rather than
paired spines. However, the photograph of this specimen
(Palaeontological Institute, Moscow, no. 4248/121) pro-
vided by Bruton et al. in their internet catalogue of
Schmidt’s trilobites (see above), shows that the appearance
of a single occipital spine is apparently caused by breakage,
and that paired spines were probably originally present.
Warburg (1939) considered that this specimen may belong
to her species Hoplolichoides curvifrons, based on speci-
mens from an erratic of early Caradoc age from Sweden, but
the specimen seems to agree in most features with other
specimens of conicotuberculatus from Estonia. Outside
Estonia, conicotuberculatus has been recorded from Swe-
den (island of Björkön, Uppland), southern Norway, and
from glacial erratics in northern Germany (Kummerow,
1928; Warburg, 1939; Neben and Krueger, 1971).
The oldest species of the genus is Hoplolichoides furcifer
(SCHMIDT, 1885), from the Aseri-Uhaku Stages (C1,
middle Llanvirn-lowermost Caradoc) of Estonia and Russia
(St Petersburg district). It differs from conicotuberculatus in
having a broader bullar lobe; the minimum width of the
median lobe is situated farther back, behind a transverse line
through the intersection of the axial and palpebral furrows;
the longitudinal furrow is more strongly curved and is
shallower posteriorly behind S1; and L1a is subtriangular
rather than ovate. In the pygidium of furcifer, incorrectly
assigned to Hoplolichas plautini by Schmidt (1885) (see
above), the first two marginal spines are broader than in
conicotuberculatus, are less inclined across their width, and
do not converge backwards. Hoplolichoides curvifrons
(WARBURG, 1939) is most readily distinguishable from
conicotuberculatus by the third pair of pygidial spines that
are rounded and lobate rather than subtriangular.
Subfamily PLATYLICHINAE Phleger, 1936
Diagnosis – Axial furrow effaced adjacent to L1b, which
merges abaxially with fixigenal field. Bullar lobe circum
118 D.J. Holloway, A.T. Thomas / Geobios 35 (2002) 111–125
scribed, bounded posterolaterally by S1; in some genera,
bullar lobe expanded backwards to reach or almost reach
L1a, displacing adaxial part of L1b; longitudinal furrow
invariably extends to occipital furrow or L1a. Hypostome
with middle body narrower across posterior lobe than across
anterior lobe; shoulder situated opposite middle furrow or
posterior lobe of middle body. Pygidium with 2–3 pairs of
pleural furrows and three pairs of flattened marginal spines;
anterior and posterior bands usually flattened.
Genera and subgenera included – Platylichas (Platyli-
chas) GÜRICH, 1901; P. (Rontrippia) THOMAS & HOL-
LOWAY, 1988; Allolichas KRUEGER, 1992; Autoloxoli-
chas PHLEGER, 1936; Metalichas REED, 1902.
Remarks – Thomas and Holloway (1988, fig. 362, p.
249) recognized the close affinities between Platylichas and
Autoloxolichas, a pair of genera to which Allolichas is also
related. It is appropriate to use the Subfamily Platylichinae
to reflect this grouping, to which we also assign the poorly
known Metalichas (see Thomas and Holloway, 1988). This
subfamily was recognised by Hupé (1953) and Balashova
(1960), but Tripp (1957, 1959) and Thomas and Holloway
(1988) implicitly placed it in synonymy with the Homoli-
chinae by including Platylichas in the latter subfamily.
Members of the Platylichinae differ from the Homolichinae
in having the axial furrow effaced adjacent to L1b, which
consequently is fused with the fixigenal field instead of with
the bullar lobe to form a composite lobe. The strongly
oblique furrow bounding the bullar lobe posterolaterally in
the Platylichinae is not the axial furrow but S1, which meets
the longitudinal furrow posteriorly and joins the axial
furrow anteriorly near the front of the palpebral lobe (Fig.
1.6–8). The homology of the glabellar lobes and furrows is
fairly easily determined in Platylichas and Metalichas,
which have the bullar lobe relatively widely separated from
L1a. In Allolichas and Autoloxolichas, however, the bullar
lobe has expanded backwards to reach or almost reach L1a,
thus assuming more of the appearance of the composite
lateral lobe of the Homolichinae. In some specimens of
Allolichas and Autoloxolichas the axial furrow is weakly
discernible adjacent to L1b, running almost parallel with the
sagittal axis, showing that the enlarged bullar lobe in these
genera has displaced L1b, which is very small and subtri-
angular in outline. Given the similarities in gross glabellar
morphology shared by Platylichas and the lichakephalid
Lichakephalus, the Platylichinae appear less derived in their
glabellar structure than the Homolichinae.
Thomas and Holloway (1988) regarded the hypostomal
characters as important in characterizing their concept of a
united Homolichinae + Platylichinae, but those shared simi-
larities are now regarded as primitive for lichids as a whole
(see remarks on Homolichinae). Not all platylichine hypos-
tomes retain the primitive condition of being longer than
wide, some being more elongated (e.g. Allolichas, see
below; some species of Platylichas, see Thomas and Hol-
loway, 1988, pp. 184, 213, pl. 9, fig. 173).
Genus Allolichas KRUEGER, 1992
Type species – Allolichas longispinus KRUEGER, 1992,
p. 271, from glacial erratics of the Backsteinkalk (upper
Kukruse Stage, C2β, lower Caradoc) of north-eastern Ger-
many; original designation.
Other species – A. crescenticus (REED, 1935) (= Lichas
(Platylichas) vicinus REED, 1935; see Morris 1988, p.
179); A. glenos (WHITTINGTON, 1962); A. halli (FOER-
STE, 1888); A. laxatus (MCCOY, 1846); A. micus (ANTSY-
GIN, 1973); A. noctua (PRICE, 1980); A. nodulosus (MC-
COY, 1851); A. thraivensis (REED, 1935); A.? gracile
(KUMMEROW, 1928); A.? inconsuetus (RAYMOND,
1925); A.? miseneri (FOERSTE, 1920).
Stratigraphical and geographical range – Arenig,
Urals; Caradoc-Ashgill, North America (Indiana,
Ohio, ?New York), Ireland, England, Scotland, Wales, Nor-
way, Sweden, northern Germany (erratics), Estonia, Arctic
Russia (Vaigach Island and Pai-Khoi).
Diagnosis – Platylichine with gently convex (sag., tr.)
cranidium, glabella not overhanging anteriorly; anterior
border flattened, commonly expanding weakly adaxially,
comprising about 10% sagittal cranidial length. Longitudi-
nal furrow meeting inner end of S1 posteriorly in acute
angle at adaxial extremity of L1a. Long axis of bullar lobe
almost parallel to sagittal line; minimum width of median
lobe situated level with midlength (exsag.) of bullar lobe,
less than width of bullar lobe measured across same
transverse line. Hypostome approximately as wide across
shoulders as long (sag.), with subcircular middle body.
Pygidium with axis comprising half sagittal length or more;
three or four axial rings, last ring furrow commonly
incomplete medially; postaxial band narrow. Pleurae with
three pairs of pleural furrows and three pairs of long and
slender marginal spines, first two pleural furrows running
distally onto bases of marginal spines; gently convex border
present posteriorly, bounded by distinct border furrow
joining furrow alongside postaxial band with last pleural
and interpleural furrows.
Remarks – Krueger (1992) included in Allolichas only
the type species, A. longispinus, the long median occipital
spine of which he considered to be the most important
diagnostic character of the genus. In other characters, such
as the intersection of the longitudinal furrow and S1 in an
acute angle at the adaxial extremity of L1a, the narrowness
of the median lobe between the bullar lobes, the weakly
adaxially expanding anterior border, the long (sag.) pygidial
axis and narrow postaxial band, the long and slender
pygidial marginal spines, and the gently convex border on
the posterior part of the pygidium, longispinus resembles
species such as ‘Lichas’ laxatus MCCOY, 1846 (see Dean,
1963, p. 235, pl. 43, figs 1–2, 5, 8–12), ‘Trochurus’
nodulosus MCCOY, 1851 (see Thomas and Holloway,
1988, pl. 9, figs 185, 188–192), and ‘Platylichas’ noctua
PRICE, 1980. Such species, which were included in Au-
toloxolichas by Thomas and Holloway (1988), lack the long
occipital spine of longispinus, though a small median spine
or tubercle may be present (e.g. Warburg, 1939, pl. 12, fig.
 D.J. Holloway, A.T. Thomas / Geobios 35 (2002) 111–125
8a; Thomas and Holloway, 1988, pl. 9, figs 188–190).
However, we do not consider the presence of cranidial
spines to be especially important taxonomically in Lichida
(Thomas and Holloway, 1988). We therefore expand the
concept of Allolichas to include all the species that we
previously assigned to Autoloxolichas, except for the type
species of the latter. See the discussion of Autoloxolichas for
comparison of that genus with Allolichas.
The hypostome of Allolichas is distinctive in its rather
elongated outline and subcircular middle body (see Whit-
tington, 1962, pl. 7, figs 9, 10; Dean, 1963, pl. 43, fig. 12;
Price, 1980, pl. 113, fig. 2; Krueger, 1992, pl. 1, fig. 3).
Genus Autoloxolichas PHLEGER, 1936
Type species –Lichas St. Mathiae SCHMIDT, 1885, p.
115, from the Jõhvi (D1) and Keila (D2) stages (middle–up-
per Caradoc) of Estonia; original designation.
Other species – None known.
Diagnosis – Platylichine with gently convex (sag., ex-
sag., tr.) cranidium; anterior border short (less than half
sagittal length of occipital ring), upper surface sloping
forward at almost same angle as front of glabella, anterior
surface vertical. Bullar lobe ovate in outline, circumscribed
by axial and longitudinal furrows and S1, separated from
L1a posteriorly by very short (exsag.), slightly depressed
region; long axis of bullar lobe oriented at about 30° to
sagittal line Minimum width of median lobe situated well
behind midlength (exsag.) of bullar lobe, less than width of
bullar lobe measured across same transverse line.
Cranidium with sculpture of dense granules grading into
terrace ridges on front of glabella.
Remarks – Öpik (1937) considered that the pygidia
assigned to sanctamathiae by Schmidt are not conspecific
with the cranidia. Sculptural differences (granules grading
anteriorly into undulating ridges on the cranidia; zigzag
scaly ridges on the pygidium) lend some support to this
view. Because of the uncertainty in the assignment of the
pygidia, the generic diagnosis is based on cranidial charac-
ters alone.
Autoloxolichas is here restricted to the type species, the
other species included in the genus by Thomas and Hollo-
way (1988) being placed in Allolichas. Allolichas is very
similar to Autoloxolichas in the development of the
cranidial lobes and furrows but differs in having the
longitudinal furrow extending farther posteriorly, reaching
the adaxial extremity of L1a instead of uniting with the
inner end of S1 a short distance in front of L1a; the bullar
lobe is acute posteriorly rather than rounded, and is less
obliquely oriented, its long axis lying approximately parallel
to the sagittal line; the minimum width of the median lobe
is situated farther forwards, level with the midlength (ex-
sag.) of the bullar lobe; and the anterior border is horizontal
rather than sloping forwards, and commonly lengthens
slightly medially. Pygidia of Allolichas are not similar to
those assigned to Autoloxolichas sanctamathiae by Schmidt
(1885).
Autoloxolichas sanctamathiae (SCHMIDT, 1885)
119
Fig. 3.1–10
1885 Lichas St. Mathiae SCHMIDT, p. 115, pl. 5, figs
11–16.
1936 Autoloxolichas st.-mathiae (SCHMIDT) - Phleger,
fig. 78.
1937 Platylichas st. mathiae FR. SCHMIDT - Öpik, p.
57, pl. 22, fig. 2, text-fig. 15.
1957 Platylichas sanctaemathiasae (SCHMIDT) - Tripp,
p. 116.
1958 Platylichas sanctaemathiasae (SCHMIDT, 1885) -
Tripp, p. 576.
1988 Autoloxolichas sanctamathiae (SCHMIDT, 1885) -
Thomas and Holloway, p. 206, pl. 10, figs 212, 213.
in press Autoloxolichas sanctamathiae (SCHMIDT,
1885) - Whittington, fig. 4.1–4.5.
Type material – Lectotype designated herein, cranidium
CNIGR 92/11101, figured Schmidt (1885, pl. 5, fig. 13),
Fig. 3.6 herein; from the Keila (D2) stage (middle–upper
Caradoc) of Kegel (= Keila), Estonia. This designation is
made in order to clarify the application of the species name,
as the pygidia associated with the cranidia by Schmidt
(1885) may not be conspecific. Paralectotypes include
cranidia CNIGR 90/11101 (figured Schmidt, 1885, pl. 5, fig.
11; Fig. 3.5 herein) and CNIGR 91/11101 (Schmidt, 1885,
pl. 5, fig. 12; Fig. 3.8 herein); hypostome CNIGR 93/11101
(Schmidt, 1885, pl. 5, fig. 14; Fig. 3.4 herein); and pygidia
CNIGR 94/11101 (Schmidt, 1885, pl. 5, fig. 15a, b; Fig. 3.9
herein) and CNIGR 95/11101 (Schmidt 1885, pl. 5, fig. 16;
Fig. 3.7, 10 herein). Additional syntypes mentioned by
Schmidt (1885) but not figured by him have not been traced.
Other material – Cranidia GIE Tr2254 (figured Öpik
1937, pl. 22, fig. 2; Thomas and Holloway, 1988, pl. 10, fig.
212), GIE Tr19546 (figured Thomas and Holloway, 1988,
pl. 10, fig. 213), and RM Ar54838a–b (Fig. 3.1–3).
Description – Cranidium approximately twice as wide at
posterior margin as long (sag.); glabella widest across bullar
lobes (level with intersection of axial and palpebral furrows)
and occipital ring, slightly narrower across frontal lobe and
L1a, narrowest across L1b (level with posterior end of
bullar lobe; see below). Occipital ring with small, weak
median swelling close to posterior margin. L1a, bullar and
frontomedian lobes with slight independent convexity. L1a
lenticular, almost as wide (tr.) as median part of occipital
furrow. L1b generally not defined abaxially, but in some
specimens (e.g. Thomas and Holloway, 1988, pl. 10, fig.
213, left side) axial furrow faintly visible in front of L1a,
running almost parallel to sagittal line from a point level
with 33% distance between abaxial and adaxial extremities
of L1a, meeting outer end of S1 just behind intersection of
axial and palpebral furrows. S1 directed posteromedially at
about 40° to sagittal line, not clearly differentiated in depth
or course from axial furrow immediately in front of S1,
shallowing posteriorly towards junction with longitudinal
furrow. Bullar lobe ovoid, more pointed posteriorly than
anteriorly. Longitudinal furrow meeting axial furrow at
about 25% glabellar length (sag.), directed slightly forwards
120 D.J. Holloway, A.T. Thomas / Geobios 35 (2002) 111–125

Fig. 3. Autoloxolichas sanctamathiae (SCHMIDT, 1885), Keila Stage (D2, middle-upper Caradoc), Estonia. 1-3. RM Ar54838a, exfoliated cranidium,
collected V. Jaanusson 1941; dorsal, oblique and lateral views, × 2.25; Pääsküla. 4 CNIGR 93/11101, mostly exfoliated hypostome, figured Schmidt (1885,
pl. 5, fig. 14); ventral view, × 2; St Mathias (= Madise). 5. CNIGR 90/11101, partly exfoliated cranidium, figured Schmidt (1885, pl. 5, fig. 11); dorsal view,
× 1.75; Madise. 6. CNIGR 92/11101, cranidium, lectotype, figured Schmidt (1885, pl. 5, fig. 13a, b); dorsal view, × 3; Keila. 7, 10. CNIGR 95/11101,
exfoliated pygidium, figured Schmidt (1885, pl. 5, fig. 16); posterior and dorsal views, × 1.75; Haljal (= Haljala) or Friedrichshof (= Saue). 8. CNIGR
91/11101, cranidium, figured Schmidt (1885, pl. 5, fig. 12); dorsal view, × 4; Spitham (= Põõsaspea). 9. CNIGR 94/11101, partly exfoliated pygidium, figured
Schmidt (1885, pl. 5, fig. 15a, b); dorsal view, × 3.5; Madise. All photographs except Fig. 6 are of plaster replicas.
Autoloxolichas sanctamathiae (SCHMIDT, 1885), Keila Stage (D2, Caradoc moyen-supérieur), Estonia. 1-3. RM Ar54838a, cranidium esfolié, collecté par
V. Jaanusson 1941; vues dorsale, oblique et latérale, × 2.25; Pääsküla. 4 CNIGR 93/11101, hypostome presque entièrement exfolié, figuré par Schmidt (1885,
pl. 5, fig. 14); vue ventrale, × 2; St Mathias (= Madise). 5. CNIGR 90/11101, cranidium partiellement exfolié, figuré par Schmidt (1885, pl. 5, fig. 11); vue
dorsale, × 1.75; Madise. 6. CNIGR 92/11101, cranidium, lectotype, figuré par Schmidt (1885, pl. 5, fig. 13a, b); vue dorsale, × 3; Keila. 7, 10. CNIGR
95/11101, pygidium exfolié, figuré par Schmidt (1885, pl. 5, fig. 16); vues postérieure et dorsale, x 1.75; Haljal (= Haljala) ou Friedrichshof (= Saue). 8.
CNIGR 91/11101, cranidium, figuré par Schmidt (1885, pl. 5, fig. 12); vue dorsale, × 4; Spitham (= Põõsaspea). 9. CNIGR 94/11101 pygidium partiellement
exfolié, figuré par Schmidt (1885, pl. 5, fig. 15a, b); vue dorsale, × 3.5; Madise.

before curving backwards in a broad arc to join S1;
narrowest part of median lobe situated at about 60%
glabellar length (sag.) from anterior. Palpebral lobe situated
less than its own length from posterior cranidial margin,
sloping abaxially almost at same angle as fixigena, lying
farther from sagittal line posteriorly than anteriorly, poste-
rior edge opposite posterior edge of bullar lobe; palpebral
furrow much more weakly curved than palpebral margin,
meeting axial furrow level with cranidial midlength (sag.).
Posterior branch of facial suture sigmoidal, diverging
strongly backwards; anterior branch running parallel to
axial and preglabellar furrows. Posterior border weakly
convex (exsag.), expanding strongly abaxially; posterior
border furrow directed obliquely forwards abaxially. Ante-
rior border of uniform length (sag., exsag.). approximately
50% sagittal length of occipital ring; anterior border furrow
much weaker on external surface than on internal mould.
Cranidium with sculpture of dense granules of two sizes, on
larger specimens merging into undulating transverse ridges
on frontal lobe of glabella.
Hypostome with sagittal length 80% width across shoul-
ders, latter situated just behind midlength (sag.). Middle
 D.J. Holloway, A.T. Thomas / Geobios 35 (2002) 111–125
body gently convex transversely and weakly convex sagit-
tally; anterior lobe comprising about 75% sagittal length of
middle body, subelliptical in outline, widest at midlength
(sag.); middle furrow situated opposite shoulder, deep,
transverse, abruptly dying out about half way between
lateral border furrow and sagittal line; posterior lobe 70%
maximum width of anterior lobe, with elongated maculae
extending posteromedially from middle furrow. Lateral
border convex (tr.) behind shoulder, outer edge with slight
embayment opposite posterior border furrow; lateral border
furrow shallower and broader opposite posterior lobe of
middle body than opposite anterior lobe, even shallower
behind intersection with posterior border furrow, where it is
deflected outwards slightly before curving inwards and
dying out. Posterior border slightly convex (sag., exsag.),
highest anteriorly, posterior margin with broad, rounded
median embayment; posterior border furrow deep. Adhering
fragment of exoskeleton with terrace ridges on anterior
wing and around margin of shoulder, and with weak pits on
border opposite shoulder; internal mould with weak
wrinkles on outer part of lateral border posteriorly, replaced
by shallow dimples (swellings on interior of exoskeleton)
adaxially on lateral and posterior borders.
Pygidium (specimens possibly incorrectly assigned to
this species) with maximum width situated well in front of
midlength (sag.) and level with posterior end of axis. Axis
conical, approximately 40% maximum pygidial width ante-
riorly, poorly defined posteriorly; in transverse profile, axis
strongly convex with maximum curvature along sagittal
line, in lateral profile gradually decreasing in height poste-
riorly over most of its length but subsiding more abruptly
behind transverse line through adaxial ends of second
interpleural furrows. First axial ring with slight relief,
longest (sag., exsag.) distally and midway between axial
furrow and sagittal line, defined by continuous ring furrow;
second ring much weaker, ring furrow dying out medially;
very faint trace of third ring furrow present abaxially on
larger pygidium. Pleurae weakly convex (tr.), sloping gently
abaxially; lateral margin curving strongly adaxially from
region of maximum pygidial width to tip of first marginal
spine. First spine very short, broad, separated from second
spine by narrow incision; shape of second and third (if
present) spines unknown. First two pleural furrows not
running onto marginal spines distally but meeting outer end
of succeeding interpleural furrow at pygidial margin be-
tween spines; first pleural furrow more strongly curved than
second; third pleural furrow not impressed. First interpleural
furrow weakly concave backwards, almost straight; second
convex backward. Postaxial band funnel-shaped, narrowing
backwards more strongly in front of transverse line through
distal ends of first pleural and interpleural furrows than
behind this line. Doublure very wide, extending forwards in
sagittal line to posterior end of axis, having shallow radial
furrows abaxially beneath pleural furrows. Dorsal surface of
pygidium with scaly sculpture, on pleurae having appea-
rance of discontinuous, serrated terrace ridges aligned more
121
or les stransversely; doublure with widely spaced, continu-
ous terrace ridges concentric with margins.
Remarks – The smallest cranidium, CNIGR 91/11101
(Fig. 3.8), differs from the larger ones in lacking any trace
of transverse ridges on the front of the glabella, all pre-
served areas of exoskeleton (which include part, but not the
anteriormost part, of the frontal lobe) having a dense
sculpture of granules of two sizes. On the large cranidium
RM Ar54838a-b the exoskeleton adheres to the external
mould so that the exterior surface is not visible, but the
internal mould shows traces of the undulating, transverse
ridges on the frontal lobe (Fig. 3.1–3) The illustration of the
largest cranidium, CNIGR 90/11101, given by Schmidt
(1885, pl. 5, fig. 11) shows transverse ridges on the fragment
of exoskeleton adhering to the frontal lobe, but no trace of
sculpture now remains on the specimen itself, the surface of
the exoskeletal fragment having been obliterated by scra-
ping. This specimen is also less complete on the left side
than shown in Schmidt’s illustration, most of the fixigena,
the anterior border, and the lateral part of the occipital ring
being missing. These differences may be due to restoration
in the illustration or to later damage to the specimen.
Subfamily TROCHURINAE Phleger, 1936
Genus Akantharges PHLEGER, 1936
Type species – Lichas Gourdoni BARROIS, 1886, p.
126, from the Eifelian of the central Pyrenees, France;
original designation.
Other species – A. erbeni (MEISCHNER, 1965); A. sp.
of Thomas and Holloway (1988).
Stratigraphical and geographical range – Middle De-
vonian (Eifelian–Givetian); France (Pyrenees), Germany
(Rheinisches Schiefergebirge), Morocco.
Diagnosis – Trochurine with strongly convex (sag., ex-
sag.) glabella not overhanging anterior border; longitudinal
furrow very shallow anteriorly and behind S1; L1 depressed
medially; L1a not defined. Palpebral lobe small, situated
low on cheek rather far from glabella; anterior and posterior
sections of facial suture collinear. Posterolateral cranidial
lobe bearing long (exsag.) ridge curving forwards and
backwards abaxially, and bounded posteriorly by distinct
furrow; ridge may interrupt posterior border furrow distally.
Subgenal notch may be well developed; genal spine long
and blade-like. Pygidium with three pairs of long, slender
marginal spines that are rounded in cross-section. Axis
comprising 66% sagittal length of pygidium or more, with
one or two prominent axial rings and at least eight addi-
tional, poorly defined rings; postaxial ridge indistinct or
absent. Anterior pleural bands slightly inflated on segments
1–3, on third segment running subparallel with sagittal axis
anteriorly and converging posteriorly; pleural and interpleu-
ral furrows shallow; border absent.
Remarks – A distinctive feature of Akantharges is the
rounded (exsag.) ridge curving forwards and backwards
122 D.J. Holloway, A.T. Thomas / Geobios 35 (2002) 111–125
abaxially across the posterolateral cranidial lobe. This ridge,
which bears a number of large tubercles, is bounded
posteriorly by a rather sharp furrow originating near the
outer end of the occipital furrow and curving forwards
initially and then backwards abaxially towards the posterior
border furrow. The ridge and furrow are well developed in
the type species, A. gourdoni, and in the unnamed species
from Morocco (see Thomas and Holloway, 1988, pl. 14, figs
300, 303, 305), but they are weaker in A. erbeni (see Basse,
1998, pl. 15, fig. 26a–b; Fig. 4.6, 8, 10–11 herein). A similar
ridge and furrow are present also in Ceratarges, as noted by
Thomas and Holloway (1988), but are not known in other
lichids. In Ceratarges the ridge carries the eye (situated at
the end of a long stalk) and, lateral to the base of the eye
stalk, the posterior branch of the facial suture, but in
Akantharges the eye and the posterior branch of the facial
suture have migrated laterally so that they are no longer
situated on the ridge.
The relationships of many genus-group taxa within the
Trochurinae are unclear. This is particularly so for the
Devonian forms, which are commonly both distinctive and
imperfectly known. Based on glabellar and pygidial fea-
tures, and on the supposed absence from both of a subgenal
notch, Thomas and Holloway (1988, p. 252, fig. 365)
tentatively suggested that Akantharges might have been
derived from Richterarges. It is now known that a subgenal
notch may be present in Akantharges. Thomas and Hollo-
way also suggested a possible derivation of Ceratarges, and
other genera with deep subgenal notches and medially
shortened L1, from Acanthopyge (Lobopyge).
Adrain (1994) adopted a restricted definition of the
related Richterarges, assigning some species formerly ac-
commodated there to his Borealarges. He (p. 1084) reco-
gnised a Richterarges/Borealarges/Terranovia clade within
the Trochurinae but, in discussion, identified no particular
points of similarity suggesting a close relationship between
Ceratarges and this clade. The common possession of the
curved ridge and furrow on the posterolateral cranidial lobe
may indicate that Akantharges and Ceratarges are closely
related, and were derived from a common ancestor.
The pygidium of A. gourdoni is not well known due to
deformation of the specimens, and consequently Thomas
and Holloway (1988) did not include pygidial characters in
their generic diagnosis. Pygidial characters in the revised
diagnosis given above are based mainly on the somewhat
better preserved specimens of A. erbeni. In overall propor-
tions, and in the three pairs of long, slender marginal spines
that are rounded in cross section, the pygidium of
Akantharges resembles that of Acanthopyge (Acanthopyge),
but the latter is distinguished mainly by the presence of a
distinct postaxial band bordered laterally by furrows. If the
above suggestion of a common ancestry for Akantharges
and Ceratarges is correct, then A. (Acanthopyge) is less
closely related to Akantharges.
Akantharges erbeni (MEISCHNER, 1965)
Fig. 4.1–14
1965 Acanthopyge erbeni MEISCHNER, p. 124, pl. 1,
figs 1–7, text–figs 1–6.
1998 Akantharges? cf. erbeni (MEISCHNER, 1965) -
Basse, pl. 15, fig. 26a–b.
?1998 Akantharges? gr. erbeni (MEISCHNER, 1965) -
Basse, p. 78, pl. 11, fig. 8.
Holotype - Internal and external moulds of cranidium Gö
548–1; figured Meischner (1965, pl. 1, fig. 1a–c), Fig.
4.1–2, 5 herein; from Givetian phyllites near Gellershausen,
Kellerwald, Germany.
Diagnosis – L1 shorter (sag.) than occipital ring; median
and bullar lobes without paired arrangement of coarse
tubercles; curved ridge on posterolateral cranidial lobe
relatively weak, not interrupting posterior border furrow
distally; subgenal notch well developed.
Remarks – Meischner (1965) gave an exhaustive de-
scription of A. erbeni, as well as providing reconstructions
of the cephalon and pygidium. The specimens have all been
flattened tectonically to some extent, and some have also
been sheared, affecting the convexity of the exoskeleton to
varying degrees. From examination of a range of specimens
it is apparent that Meischner’s (1965, fig. 2e) depiction of
the bullar and posterolateral cranidial lobes as subconical,
posterolaterally curving projections represents an artefact of
deformation rather than the original appearance of these
lobes. These lobes were probably originally evenly rounded
(exsag.).
Comparison with A. gourdoni is hindered by the strong
deformation of the material of that species, but obvious
differences from A. erbeni include: L1 is longer (sagittal
length greater than that of the occipital ring); the curved
ridge on the posterolateral cranidial lobe is more distinct
and interrupts the posterior border furrow distally; the
subgenal notch is much shallower; and the pygidial axis is
longer (about 85% of the sagittal length of the pygidium).
The well preserved cranidium of Akantharges sp. from
Morocco (Thomas and Holloway, 1988, pl. 14, figs
299–300, 303) differs from A. erbeni in that L1 is longer
(sag.); the curved ridge on the posterolateral cranidial lobe
is more distinct and interrupts the posterior border furrow
distally; the subgenal notch is non-existent; there are promi-
nent paired tubercles on the median glabellar and bullar
lobes; and there are three very coarse tubercles in a row on
the curved ridge on the posterolateral cranidial lobe.
The cranidium from Usseln, Germany, figured by Basse
(1998, pl. 15, fig. 26a–b) as Akantharges? cf. erbeni agrees
with the types in all observable characters and is undoub-
tedly conspecific, but worthwhile comparisons cannot be
made with the incomplete pygidium figured by Basse (1998,
pl. 11, fig. 8) as Akantharges? gr. erbeni.
 D.J. Holloway, A.T. Thomas / Geobios 35 (2002) 111–125 123

Fig. 4. Akantharges erbeni (MEISCHNER, 1965), Givetian phyllites near Gellershausen, Kellerwald, Germany. 1-2, 5. Gö548-1 cranidium, holotype, figured
Meischner (1965, pl. 1, fig. 1a–c); latex cast of external mould, dorsal view, and internal mould, dorsal and oblique views, × 3.25. 3, 11. Gö548-69 cranidium;
latex cast of external mould and internal mould, dorsal views × 3.25. 4. Gö548-7 librigena; latex cast of external mould, ventral view, × 3.5. 6. Gö548-137
cranidium; latex cast of external mould, dorsal view, × 3.25. 7. Gö548-237 hypostome; latex cast of external mould, ventral view, × 4.5. 8. Gö548-76
cranidium; latex cast of external mould, × 3. 9. Gö548-134 cranidium; latex cast of external mould, dorsal view, × 3. 10. Gö548-24 cranidium; latex cast
of external mould, dorsal view, × 3.25. 12. Gö548-12 pygidium, figured Meischner (1965, pl. 1, fig. 6; left and right reversed in photograph); natural mould,
dorsal view, × 2. 13. Gö548-74, pygidium; latex cast of external mould, dorsal view, × 1.75. 14. Gö548-278 pygidium; latex cast of natural mould, ventral
view, × 2.
Akantharges erbeni (MEISCHNER 1965), phyllites du Givétien près de Gellershausen, Kellerwald, Germany. 1-2, 5. Gö548-1 cranidium, holotype, figuré
par Meischner(1965, pl. 1, fig. 1a–c); moulage en latex d’un moule externe, vue dorsale et moule interne, vues dorsale et oblique, × 3.25. 3, 11. Gö548-69
cranidium; moulage en latex d’un moule externe et d’un moule interne, vue dorsale, × 3.25. 4. Gö548-7 librigena; moulage en latex d’un moule externe, vue
dorsale, × 3.5. 6. Gö548-137 cranidium; moulage en latex d’un moule externe, vue ventrale, × 3.25. 7. Gö548-237 hypostome; moulage en latex d’un moule
externe, vue ventrale, × 4.5. 8. Gö548-76 cranidium; moulage en latex d’un moule externe, × 3. 9. Gö548-134 cranidium; moulage en latex d’un moule
externe, vue dorsale, × 3. 10. Gö548-24 cranidium; moulage en latex d’un moule externe, vue dorsale, × 3.25. 12. Gö548-12 pygidium, figuré par
Meischner(1965, pl. 1, fig. 6; photographies gauche et droite inversées); moule naturel, vue dorsale, × 2. 13. Gö548-74 pygidium; moulage en latex d’un
moule externe, vue dorsale, × 1.75. 14. Gö548-278 pygidium; moulage en latex d’un moule naturel, vue ventrale, × 2.
124 D.J. Holloway, A.T. Thomas / Geobios 35 (2002) 111–125
. Biblio non appelée
Tripp, 1958
Acknowledgements
We are grateful to H. Jahnke (Georg-August-Universität,
Göttingen) and E. Pietrzeniuk (Museum für Naturkunde der
Humboldt Universität, Berlin) for granting access to collec-
tions and arranging the loan of material; J. Bergström
(Naturhistoriska Riksmuseet, Stockholm) and I.M.
Kolobova (formerly of VSEGEI, St Petersburg) for provid-
ing latex moulds of specimens; and D.L. Bruton (Paleon-
tologisk Museum, Oslo) for providing the image of the
lectotype of Autoloxolichas sanctamathiae in Fig. 3.6. DJH
gratefully acknowledges a grant from the Deutscher Aka-
demischer Austauschdienst, enabling him to travel to Ger-
many in 1993, and thanks Hans-Hartmut and Ingrid Krueger
for their kind hospitality in Berlin. We thank E.N.K.
Clarkson and R. Feist for their constructive reviews of the
manuscript.
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