phylogeny of the trilobite subgenus acanthopyge (lobopyge)

Cladistics 17, 1–10 (2001)

doi:10.1006/clad.1999.0145, available online at http://www.idealibrary.com on

Phylogeny of the Trilobite SubgenusAcanthopyge (Lobopyge)

Malte C. Ebach*,†,‡,§ and Shane T. Ahyong‡,¶

*Department of Earth and Planetary Sciences, Western Australian Museum, Francis Street, Perth,Western Australia 6000, Australia; †School of Botany, University of Melbourne, Parkville, Victoria 3052, Australia;¶Department of Marine Invertebrates, ‡Australian Museum, 6 College Street, Sydney, New South Wales2010, Australia; and §School of Geosciences, University of Sydney, Sydney, New South Wales 2006, Australia

related to A. (Lobopyge) with varying degrees of confi-

dence. This study aims to determine the affinities of

Accepted August 1, 2000; published online February 7, 2001

Cladistic analysis of the trilobite subgenus Acanthopyge(Lobopyge) has not been previously attempted, apart froman inferred phylogeny of the Lichida by Thomas andHolloway (1988, Philos. Trans. R. Soc. London B Biol. Sci.321, 179–262). Results of two separate analyses withvariable taxonomic sampling show a possible cosmopoli-
tan affinity for Australian Devonian species of A. (Lobo- pyge) and corroborate the placement of A. (Lobopyge)rohri in A. (Lobopyge). Benelopyge is a junior subjectivesynonym of A. (Lobopyge). q 2001 The Willi Hennig Society
INTRODUCTION

A recent phylogeny of the Lichida (Thomas and Hol-

loway, 1988) modeled Acanthopyge (Lobopyge) Pribyl

and Erben, 1952 and Richterarges Phleger, 1936, as being

derived from the predominantly Ordovician HemiargesGurich, 1901. Thomas and Holloway’s (1988, p. 252,

Fig. 365) phylogeny assumed that Acanthopyge (Acan-thopyge) Hawle and Corda, 1847, A. (Jasperia) Thomas

and Holloway, 1988, and A. (Perunaspis) Pribyl, 1949,

stem from the A. (Lobopyge) lineage. The genera

Ceratarges Gurich, 1901, Eifliarges Richter and Richter,

1917, Mephiarges Richter and Richter, 1930, Terranovia

0748-3007/01 $35.00 1Copyright q 2001 by The Willi Hennig Society

All rights of reproduction in any form reserved

Maksimova, 1977, and species later assigned to Bore-alarges Adrain, 1994, were also thought to be closely

Australian species of A. (Lobopyge) using cladistic

analysis.

SYSTEMATIC ANALYSIS OF Acanthopyge(Lobopyge)

Of the 38 named A. (Lobopyge) species, 33 have been

coded (Fig. 1). Taxa with more than half their character

states coded as not known will be removed, thus

avoiding an increase in ambiguity.

Outgroup Selection

Thomas and Holloway (1988) considered Hemiargesto display the closest relationship to A. (Lobopyge) (Fig.

2). H. wesenbergensis, the type of Hemiarges, was chosen

as outgroup for this analysis. H. wesenbergensis is plesi-

omorphic for most character states assigned to A. (Lobo-pyge) (Fig. 1) and can be scored for most characters

used herein.

ge
FIG. 1. Character matrix for Hemiarges wesenbergensis and Acanthopy
Some recent analyses have rooted trees using hypo-

thetical ancestors (Ramskold and Werdelin, 1991) even

when suitable outgroups were available. The use of

hypothetical ancestors is justifiable only when it is “nei-

ther feasible nor convenient to select an outgroup”

(Nixon and Carpenter, 1993, p. 421, Bryant, 1997). The

use of a hypothetical ancestor and not an actual taxon

is likely to result in a different ingroup topology re-

gardless of whether or not the data set is complete

(Maddison et al., 1984), when compared to results pro-

duced when using explicit outgroups (Nixon and Car-

penter, 1993, Bryant, 1997).

Characters

Descriptive terminology for cranidial furrows and

lobation in A. (Lobopyge) follows Thomas and Hol-

loway (1988, Fig. 2j), following ontogenetic evidence

of Chatterton (1971). Morphological characters of A.(Lobopyge) are shown in Fig. 3 and all character states

are listed in Fig. 1.

1. Posterior extent of longitudinal glabellar furrow.

Outgroup. In H. wesenbergensis the longitudinal gla-

bellar furrows efface at S1.

Copyright q 2001 by The Willi Hennig Society


(Lobopyge).

States. 0: longitudinal furrow effaced at S1, not con-

tinuous with S0; 1: longitudinal furrow continuous

with S0.

2. Course of S1 medially.

Within A. (Lobopyge), the two character states are

only associated with (1:1).

Outgroup. The character is inapplicable to Hemiargesand cannot be coded and hence is assigned as missing.

States. 0: not merging with S0 medially; 1: merges

with S0 medially.

3. Tuberculation on preoccipital glabellar lobe(s).

The preoccipital region is either nontuberculate or

has two distinct tubercles on its lateral margins in Dev-

onian A. (Lobopyge).Outgroup. In H. wesenbergensis the area is always tu-

berculate.

States. 0: numerous scattered tubercles, 1: nontuber-

culate; 2: two distinct nodes present.

4. Width of median glabellar lobe.

Variation in the width of median glabellar lobe is

measured as the ratio of its width at its anterior limit

to its width at juncture of S1 and the longitudinal fur-

row. In all known A. (Lobopyge) species, the anterior

2 Ebach and Ahyong

width is equal or greater than the posterior width.

Outgroup. Hemiarges has a large, broad median lobe.

FIG. 2. Line drawings of (A) Hemiarges and (B) Acanthopyge (Lobo-
pyge), both from the Wenlock Limestone of Dudley, United Kingdom.
Modified from Tripp (1957, p. 107, text Fig 3.)

States. 0: width anteriorly twice the width posteri-

orly; 1: width anteriorly greater than but less than twice

the width posteriorly; 2: equal width anteriorly and

posteriorly.

5. Median glabellar ornamentation.

Species show either a single pair of large nodes or

two rows of tubercles either as matching pairs or scat-

tered.

Outgroup. Hemiarges lacks spines or distinct nodes,

having tubercles evenly scattered over the glabella.

States. 0: evenly scattered tubercles; 1: a distinct sin-

gle pair or row of paired larger tubercles.

6. Distal shortening of L0.

The occipital ring in A. (Lobopyge) is longest at the

midline (sag.) and shorter distally but the position at

which this commences varies within the ingroup. In

Hemiarges, the occipital ring is of subequal length for

a considerable extent and tapers at the abaxial edge.

In many Devonian species the occipital ring begins

tapering out near the midline (sag).

FIG. 3. Schematic illustration of a Trochurinid lichid trilobite (the left



3

States. 0: tapering commences near lateral margin

(exsag.); 1: tapering commences near midline (sag.).

7. Pitting on middle body of hypostome.

Ornament lacking pitting is usually strongly tubercu-

late. Pitting and tuberculation can occur together; how-

ever, tuberculation is confined mainly to the anterior

area of the middle body.

Outgroup. Large deep pitting (as in Hemiarges) is con-

sidered primitive.

States. 0: middle body pitted, 1: middle body not

pitted.

8. Hypostome tuberculation.

Tuberculation varies on the hypostome, either oc-

curring as sparsely scattered tubercles over the whole

hypostome, concentrated on the lateral border, densely

concentrated near the anterior border of the middle

body or as coarse tuberculation over the whole mid-

dle body.

States. 0: hypostome tuberculate; 1: tuberculation on

middle body only; 2: tuberculation absent.

9. Pygidial posterior border development.

The border is absent in Devonian A. (Lobopyge); how-

ever, vestigial structures, such as a raised margin and

border furrow exist in some Silurian species (A. (L.)orientalis and A. (Lobopyge) rohri).

Outgroup. Border distinct in Hemiarges, state 0 consid-

ered primitive.

States. 0: border distinct; 1: border absent.

10. Postaxial ridge.

The posteromedian region of the pygidium may fea-

Phylogeny of Acanthopyge (Lobopyge)

ture a fusion of the posterior border with a postaxialOutgroup. Hemiarges possesses an occipital ring that

tapers out near the lateral margin (exsag). ridge. A raised margin fused with the postaxial ridge

half of a lichid pygidium and the right half of a lichid cephalon).

States. 0: wide, more than half width of pygidial axis;

1: half width of pygidial axis; 2: narrow, less than half

A includes all 33 coded taxa and Analysis B excludes

8 taxa in which 45% or more of their characters could

4

is present in Silurian A. (Lobopyge), whereas Devonian

species lack a border or raised margin in connection

with a distinct postaxial ridge.

Outgroup. Hemiarges has a postaxial ridge.

States. 0: postaxial ridge present; 1: postaxial ridge

absent.

11. Second axial ring furrow.

A. (Lobopyge) has one distinct axial ring (in line with

the first pygidial spine pair) and up to 4 or more (as in

A. (L.) balliviani) discontinuous rings. A distinct second

axial ring is here defined as continuous medially. The

distinctness of the second ring furrow (sag.) becomes

less from Silurian to Devonian A. (Lobopyge) species.

Outgroup. Hemiarges has two distinct ring furrows,

with an indistinct third. A distinct second ring furrow

is considered primitive.

States. 0: ring furrow continuous; 1: ring furrow im-

pressed across more than half of axis; 2: distinct across

less than half of axis; 3: ring furrow absent.

12. Tuberculation on second pygidial rib.

A pair of large tubercles can be present on the second

pygidial pair in holaspides.

Outgroup: Large tubercule pair absent in Hemiarges.States. 0: absent; 1: present.

13. Direction of second pygidial spine pair.

A. (Lobopyge) has two pairs of major border spines,

representing the first and second pleural ribs. The sec-

ond spine varies in direction. Silurian species have an

inwardly directed spine set, while Devonian species

possess a straight spine parallel to the axis (as in A.(L.) sinuata) or an outwardly directed spine.

Outgroup. A. (L.) rohri has distinct, inwardly directed

spines, considered primitive in A. (Lobopyge).States. 0: inwardly directed; 1: posteriorly directed

(running exsagitally); 2: outwardly directed.

14. Presence of a secondary marginal spine pair(s).

All A. (Lobopyge) posses a posterior marginal spine

pair(s) behind the second pair of pleural spines. Some

A. (Lobopyge) species exhibit an additional pair of mar-

ginal spines or nodes between these two spine pairs.

The secondary spine pair is present as a node, a

spine(s), or as a set of three to four small spines.

Outgroup. H. wesenbergensis has a second minor spine.

This is considered as a primitive state.

States. 0: spine(s) distinct; 1: subdued spine or node;

2: absent; 3: three small spines.

15. Distance between posterior marginal spine pair.

This is measured as the distance between the primary



Ebach and Ahyong

spines (on the midpoint of the spine) relative to the

pygidial axial width (tr.).

Outgroup. H. wesenbergensis codes as wide. The width

varies in A. (Lobopyge).

width of pygidial axis.

CLADISTIC ANALYSIS

Phylogenetic analysis of A. (Lobopyge) was conducted

using PAUP* 4.0b4 (Swofford, 1998) on an IBM Pen-

tium. All analyses were conducted using a random

heuristic search (with 1000 replicates), using the TBR

algorithm and ACCTRAN optimization. All multistate

characters are treated as unordered. Node strength was

subjected to a Decay analysis (Bremer, 1994). Analysis

not be coded.

RESULTS

Analysis A used all taxa coded in Fig. 1, including

taxa with a high proportion of missing data.

FIG. 4. Acanthopyge (Lobopyge) run with taxa with less than 45%

of character state information missing. Strict consensus of 18 trees.

of the 61 (in this case tree 1 (Fig. 5), a summary rather

than a series of taxonomic statements can be made


The analysis retrieved 50,600 trees of 71 steps, with

a consistency index (CI) of 0.33 and a retention index

(RI) of 0.633. The strict consensus tree was totally

unresolved.

Because of the unresolved results in Analysis A, only

species of Acanthopyge (Lobopyge) with less than 50%

missing data were used in Analysis B, including the

outgroup H. wesenbergensis (see Discussion below).

Analysis B resulted in 18 optimal trees (length 61 steps,

CI 0.377, RI 0.596) (Figs. 4 and 5). Bremer support

analysis showed that all nodes collapsed with a single

step increase in tree length.

Missing Values

The deletion of taxa with more than 45% missing data

(denoted by “?”) yielded fewer equally parsimonious

trees than Analysis A. Similar results, observed by

Kitching et al. (1998) using hypothetical data, led to

the conclusion that missing data may not only increase

the number of equally parsimonious trees, but also

cause some cladistic computer programs to yield spuri-

ous cladograms.

The exclusion of terminals with a high proportion

of missing data as outlined in Kitching et al. (1998) is

trees. Node 1 [1:0], [12:0], [14:1]; Node 2 [2:1], [3:2], [13:2]; Node 3 [4:1]

8 [6:1]; Node 9 [15:1]; Node 10 [7:1]; Node 11 [6:1]; Node 12 [5:0], [11:1]

[15:2]; Node 17 [5:0]; Node 18 [10:1]; Node 19 [6:1], [13:0]; Node 20 [9:0



5

majority of missing data concerns either the cranial or

the pygidial regions (approximately 45% of the data).

Thus, the majority of deleted taxa are based on either

cranidial or pygidial data only.The exhaustive study of A. (Lobopyge) has yielded

fewer characters than known species. Acquiring suffi-

cient characters for a full resolution of species in cladis-

tic analysis is frequently impossible in palaeontology.

However, by discussing a single randomly chosen tree

regarding A. (Lobopyge).

DISCUSSION

Analysis B highlights one distinct clade in the strict

consensus (Fig. 4). This includes A. (L.) docekali (Vanek,

1959), A. (L.) sp. Holloway and Neil, 1982, from Victo-

ria, A. (L.) hirsuta (Fletcher, 1850), A. (L.) trinodis (Eller-

mann, 1992). A. (L.) orientalis (Wu, 1977), and A. (L.)parapleura Curtis and Lane, 1998.

The first tree of Analysis B yields two additional

clades, an Australian clade consisting of A. (L.) sinuata
herein justified (Fig. 4). In the case of A. (Lobopyge), the (Ratte, 1886), A. (L.) australis (McCoy, 1876), A. (L.)
FIG. 5. Acanthopyge (Lobopyge) run with taxa with less than 45% of character state information missing (CI50.377, RI50.596). First of 61

; Node 4 [6:1]; Node 5 [9:1]; Node 6 [14:1]; Node 7 [4:0] [13:1]; Node

; Node 13 [8:1], [12:0]; Node 14 [15:0]; Node15 [13:1]; Node 16 [11:1],

]; Node 21 [3:1]; Node 22 [2:0] and Node 23 [7:0].

6

australiformis Chatterton et al. 1979, and A. (L.) campbelliChatterton and Wright, 1986; and a second consisting

of A. (L.) balliviani (Kozlowski, 1923), A. (L.) richteri(Vanek, 1959), and A. (L.) uralensis (Maksimova, 1979).

Two smaller clades consist of A. (L.) brevis (Maksimova,

1968), A. (L.) limbata (Maksimova, 1968), and A. (L.)parva (Barrande, 1846) and A. (L.) pragensis (Boucek,

1933).

The analyses also consistently support the generic

placement of A. (L.) rohri, a species initially described

in Hemiarges. Thomas and Holloway (1988) considered

Perry and Chatterton’s (1977) A. (L.) rohri to be a con-

signment of parts from other species because the crani-

dium resembles Richterarges, and the pygidium resem-

bles A. (Lobopyge). Thomas and Holloway’s (1988)

placement of A. (L.) rohri in A. (Lobopyge) is supported

by the present results. The pygidium of A. (L.) rohrishows A. (Lobopyge) characteristics (13:0 and 14:2)

(shared with five A. (Lobopyge) species). The cranidium

also displays characteristics within (Lobopyge) (1:1 and

2:0) that are shared with four other A. (Lobopyge)

the two subspecies are indistinguishable and must be

considered synonymous. L. (B.) balliviani baldsi is con-

species.

SYSTEMATIC PALAEONTOLOGY

Superfamily Lichoidea sensu Fortey, 1997

Family Lichidae Hawle and Corda, 1847

Subfamily Trochurinae Phleger, 1936

Genus Acanthopyge Hawle and Corda, 1847

Subgenus Acanthopyge (Lobopyge) Pribyl and Erben,

1952

Type Species

Lichas Branikensis Barrande, 1872. Lower Devonian

(Pragian) in age, from the Dvorce-Prokop Limestone

of the Czech Republic.

Acanthopyge (Lobopyge) balliviani (Kozlowski, 1923)

1923 Lichas balliviani Kozlowski

1967 Acanthopyge balliviani Baldis, plate l, Figs. 5

and 6

1977 Acanthopyge balliviani Baldis and Longobucco,

p. 158, plate 2, Figs. 1–7.

1988 Acanthopyge (Lobopyge) balliviani Thomas and

Holloway, p. 224.



Ebach and Ahyong

1991 Lobopyge (Belenopyge) balliviani baldisi and Lo-bopyge (Belenopyge) balliviani balliviani Pek and Vanek,

1991, p. 85, plate 4, Fig. 2–7, plate 5, Fig. 1 and 2.

Discussion

Chatterton et al. (1979) regarded Lobopyge as a subge-

nus of Acanthopyge, a decision supported by Thomas

and Holloway (1988, p. 225, Discussion). The subgenus

L. (Belenopyge) shares all characteristics with A. (Lobo-pyge), such as a short postaxial ridge (present in some

A. (Lobopyge)), axis not inflated posteriorly, broad py-

gidial pleurae (exsag.), and granules often arranged in

rows along the axis. Hence, L. (Belenopyge) herein is

considered a junior synonym of A. (Lobopyge). Pek and

Vanek (1991) justified the erection of two subspecies,

L. (B.) balliviani balliviani and L. (B.) balliviani baldisi,based on significant variation in the length (sag.) and

width of the pygidal axis, and distinctive postaxial area

on the pygidium of the latter. However, restudy of

the type material shows that on the pygidium of both

nominal subspecies, the postaxial area (resembling an

agglomeration of five tubercles on the posterior of the

terminal piece) is present in both subspecies (plate 5,

Fig. 2, Pek and Vanek, 1991 and plate 2, p. 153 Baldis

and Longobucco, 1977). There is neither a distinct dif-

ference in size or length of spines on the posterior

margin of the pygidium, nor any distinct variation in

the width and length (sag.) of the pygidial axis. Hence,

sidered a junior synonym of A. (L.) balliviani.

APPENDIX 1

List of Acanthopyge Subgenera and Species

The list includes all known species and subgenera

of Acanthopyge. Several of the listed species are junior

synonyms of other listed species. The species name

is followed by author(s); date; language of original

description; original assignment; age; area; revisions;

present assignment; and whether coded from figures

(photography) (f), illustrations (i) or specimen (s). The

format is adopted from Ramskold and Chatterton

(1991).

Perunaspis; Eifelian; Czech Republic; described and


altirhachis Chernysheva, 1951; Russian; Lichas (Eu-arges); Devonian; Kuznetsk Basin, Russia; to Acantho-pyge (Lobopyge) by Thomas and Holloway (1988); A.(Lobopyge); not coded.

australiformis Chatterton, Johnston, and Campbell,

1979; English: A. (Lobopyge); Lochovian–Pragian; New

South Wales, Australia; A. (Lobopyge). (f).

australis McCoy, 1876; English; Lichas; Lower Dev-

onian; Victoria, Australia; described by Gill (1939) and

discussed as A. (Lobopyge) by Chatterton and Wright

(1986), A. (Lobopyge); (f).

baldisi Pek and Vanek, 1991; English; Lobopyge (Belen-opyge); Middle Devonian (Givetian–Frasnian bound-

ary); San Juan Province, Argentina; figured by Baldis

and Longobucco (1977) as A. balliviani; herein A. (L.)balliviani; (f).

balliviani Kozlowski, 1923; French; Lichas; Bolivia;

illustrated by Ahlfeld and Branisa (1960) as Lobopyge(Acanthopyge) and A. balliviani, respectively, and fig-

ured by Pek and Vanek, 1991 as L. (Belenopyge) ballivani;herein Acanthopyge (Lobopyge); (f), (s).

bifida Edgell, 1955; English; A. (Mephiarges); Emsian;

New South Wales, Australia; described and figured

by Chatterton (1971), to A. (Jasperia) by Thomas and

Holloway (1988); A. (Jasperia); (f).

brankiensis Barrande, 1872; French; Lichas; Lower

Devonian; Czech Republic; figured by Pribyl and Erben

(1952) as L. (Lobopyge), refigured and illustrated by

Vanek (1959) as Lobopyge; figured by Thomas and Hol-

loway (1988) as A. (Lobopyge); A. (Lobopyge); (i), (f).

brevis Maksimova, 1968; Russian; Lobopyge; Lower

Devonian; Kazakhstan; to A. (Lobopyge) by Thomas and

Holloway (1988); A. (Lobopyge); (f).

campbelli Chatterton and Wright, 1986; English; A.(Lobopyge); Zlichkovian; New South Wales, Australia;

A. (Lobopyge); (f).

consanguinea Clarke, 1894; English; Arges; Lower

Devonian; New York, United States, figured and de-

scribed by Whittington (1956) as Acanthopyge to A. (Lo-bopyge) by Thomas and Holloway (1988); A. (Lobo-pyge): (f).

contusa Hall and Clarke, 1888; English; Lichas (Arges);Lower Devonian; New York, United States: to A. (Lobo-pyge) by Thomas and Holloway (1988); A. (Lobopyge);(i).

decheni Holzapfel, 1895; German; Lichas; Lower Dev-

onian (?); Martenberg, Germany; briefly discussed as

Acanthopyge and A. cf. decheni by Basse (1998), to A.



7

(Lobopyge) by Thomas and Holloway (1988); A (Lobo-pyge); (i).

docekali Vanek, 1959; Czech with English translation;

Lobopyge; Upper Lower Devonian: Czech Republic: to

A. (Lobopyge) by Thomas and Holloway (1988); A. (Lobo-pyge); (i), (f).

duplicispinata Kaneko, 1984; English; A. (Acantho-pyge); Middle Devonian; Kitakami Mountains, North-

east Japan; A. (Acanthopyge); not coded.

edgecombei; Ebach, 2001; Lochkovian; English; A. (Lo-bopyge) New South Wales, Australia; herein A. (Lobo-pyge); (f), (s).

erinacea Haas, 1968; German; Lobopyge; Pragian–

Lower Emsian; Ankara, Turkey; to A (Lobopyge) by

Thomas and Holloway (1988); A (Lobopyge); (f).

granulosa Roemer, 1852; German; Cheirurus; Middle

Givetium; Harz, Ketterwald, and North Sauerland,

Germany; to Lichas by Holzapfel, 1895, to Acanthopygeby Basse, 1998; Herein A. (Acanthopyge); not coded.

haueri Barrande, 1846; French; Lichas: Eifelian; Bohe-

mia, Czech Republic; to Euarges Gurich (1901), A.(Acanthopyge) leuchtenbergii considered subjective syn-

onym of A. haueri by Vanek (1959); figured by Thomas

and Holloway (1988) as A. (Acanthopyge): A. (Acantho-pyge); (f), (s).

helga Pribyl et al. 1986; English; Pernuaspis; Lower

Devonian; Prague, Czech Republic: to A. (Perniaspis)Thomas and Holloway (1988); A. (Perniaspis); not

coded.

hexapteryx Pribyl and Erben, 1952; German; Lichassexlobatus Herrmann (1912); Middle Devonian; Bohe-

mia, Czech Republic; redescribed by Pribyl and Erben

(1952) as a new species of Lobopyge, to A. (Lobopyge)by Thomas and Holloway (1988); A. (Lobopyge): (f).

hirsuta Fletchter, 1850; English; Lichas; Wenlock; Dud-

ley, England; to Acanthopyge (also figured) by Thomas

(1981), to A. (Lobopyge) by Thomas and Holloway

(1988); A. (Lobopyge); (f), (i).

limbata Maksimova, 1968; Russian; Lobopyge; Dev-

onian; Kazakhstan; to A. (Lobopyge) by Thomas and


longiaxis Maksimova 1968; Russian; Lobopyge; Dev-

onian?; Kazakhstan; to A. (Lobopyge) by Thomas and


longispina Pribyl, 1949; Czech with English summary;

figured by Vanek (1959), Lobopyge (Nitidulopyge) dev-onica synonymized with P. longispinus by Vanek (1959),

8

figured by Thomas and Holloway (1988) as A. (Pernu-aspis); A. (Pernuaspis); not coded.

mediosulcatus Kaneko, 1984; Nipponarges; Middle

Devonian; Kitakami Mountains, northeast Japan; to A.(Acanthopyge) by Thomas and Holloway (1988): A.(Acanthopyge); not coded.

minuta Barrande, 1846; French; Cheirurus; Lower

Devonian; Bohemia, Czech Republic; to A. (Pernuaspis)by Thomas and Holloway (1988) A. (Pernuaspis); not

coded.

niobe Basse, 1998; German, with English abstract; Lo-bopyge; Upper Emsian; Sauerland, Germany; herein A.(Lobopyge); not coded.

orientalis Wu, 1977; Chinese, with English abstract;

Acanthopyge; Upper Silurian; Southwest China; figured

by Fang et al. (1985) as Acanthopyge, to A. (Lobopyge)by Thomas and Holloway (1988); A. (Lobopyge); (f).

parapleura Curtis and Lane, 1998; English; A. (Lobo-pyge); Upper Silurian (Llandovery); England; ? A.(Lobopyge), possibly Hemiarges; (f).

parva Barrande, 1846; French; Lichas; Lower Dev-

onian; Czech Republic; figured by Vanek (1959) and

Ellermann (1992) as A. parva parva, to A. (Lobopyge) by

Thomas and Holloway (1988) and herein, respectively;

A. (Lobopyge); (f).

parvulus Novak, 1890; Russian; Lichas; Devonian?;

Russia; figured by Chernysheva (1951) as a new sub-

species Lichas (Eurages) parvulus convexa Chernysheva,

1951, from the Kuznetsk Basin, Russia; A. (Lobopyge)herein; not figured.

permarginata Pribyl and Erben, 1952; German; Lobo-pyge; Lower Devonian; Lower Harz Region, Germany;

herein A. (Lobopyge); not coded.

pervasta Pribyl et al. 1986; English; Acanthopyge; Ei-

felian; Bohemia, Czech Republic, to A. (Lobopyge) by

Thomas and Holloway (1988); A. (Lobopyge); (f).

pragensis Boucek, 1933; Czech?; Lichas; Ludlow;

Prague, Czech Republic; refigured by Pribyl and Erben

(1952) as Lobopyge (Lobopyge), to A. (Lobopyge) by

Thomas and Holloway (1988); A. (Lobopyge); (f).

pulex Haas, 1968; German; Lobopyge; Upper Emsian;

Pendik Burnu, Turkey; herein A. (Lobopyge); not coded.

pusilla Angelin, 1854; Latin; Lichas; Gotland, Sweden;

to A. (Lobopyge) by Thomas and Holloway (1988); A.
(Lobopyge); (i).
pustulosa Morzadec, 1983; French, with German and

English abstract; Lobopyge; Emsian; Amorica, France;



Ebach and Ahyong

to A. (Lobopyge) by Thomas and Holloway (1988); A.(Lobopyge); (f).

richteri Vanek, 1959; Czech with English translation;

Lobopyge; Upper Ludlovian; Czech Republic; to A. (Lo-bopyge) by Thomas and Holloway (1988); A. (Lobopyge);(i), (f)

rohri Perry and Chatterton, 1977; English; Hemiarges;Wenlock; Ballie–Hamilton Island, Canadian Arctic Ar-

chipeligo: to A. (Lobopyge) by Thomas and Holloway

(1988), who consider the cranidium to belonging to

Richterarges; herein A. (Lobopyge); (f).

sexlobata Roemer, 1855; German; Lichas; Lower Dev-

onian; Lower Harz Region, Germany; to Lobopyge (Niti-dulopyge) by Pribyl and Erben (1952), synonymized by

Thomas and Holloway (1988) with A. (Perniaspis): A.(Perniaspis); not coded.

sibirica Chernysheva, 1951; Russian; Lobopyge (?); Si-

lurian–Devonian (?); Russia; A. (Lobopyge); not coded.

sinuata Ratte, 1886; English; Lichas;: late Lochkovian–

middle Pragian; New South Wales, Australia; to A.(Lobopyge) by Chatterton et al. (1979); A. (Lobopyge);(f), (s).

trinodis Ellermann, 1992; German; Lobopyge; Pragian;

Karnian Alps, Austria; A. (Lobopyge) herein: (f)

uralensis Maksimova, 1979; Russian; A. haueri ura-lensis; Devonian; Russia; to A. (Lobopyge) by Thomas

and Holloway (1988); A. (Lobopyge); (f).

sp. Alps Ellermann, 1992; German; Lobopyge; Karnian

Alps, Austria; Herein; A. (Lobopyge) (f).

sp. Sauerland Basse, 1998; German; Lobopyge?; Givet-

ium; Sauerland, Germany; Herein Odontopleurid in-

dent. sp.; not coded.

sp. 1 Harz Pribyl and Erben, 1952; German; Lobopyge;Lower Devonian; Lower Harz Region, Germany;


sp. 2 Harz Pribyl and Erben, 1952; German; Lobopyge;Lower Devonian; Lower Harz Region, Germany;


sp. 1. Russia Maksimova, 1968; Russian; Lobopyge;Devonian ?; Kazakhstan; A. (Lobopyge); not coded.

sp 2. Russia Chernysheva, 1951; Russian; Lichas (Eu-arges); Devonian?; Kuznetsk Basin, Russian; to A. (Lobo-pyge) by Thomas and Holloway (1988); not coded.

sp. Victoria Holloway and Neil, 1982; English; A.
(Lobopyge); Late Lochkovian; Victoria, Australia; A. (Lo-bopyge); (f).
sp. 1 New South Wales Chatteron, Johnston and


Campbell, 1979; English; A. (Lobopyge); Emsian; New

South Wales, Australia; A. (Lobopyge). (f).

sp. 2 New South Wales Jones, Hall, Wright and Carr,

1986; English; ?A. (Lobopyge); Lochkovian–Pragian;

New South Wales, Australia; A. (Lobopyge) herein; (f),

(s).

ACKNOWLEDGMENTS

We are grateful for the help and assistance of Dr. Gregory D.

Edgecombe. We also thank an anonymous reviewer for helpful com-

ments and suggestions. M.C.E. is grateful to the University of Sydney

for granting the D. B. Helby Scholarship in 1997.

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