INTERRELATIONSHIPS OF

PLACODERMS REVISITED

DANIEL G O U J E T & GAVIN C. YOUNG

GOUJET D. & YOUNG G.C. 1995. Interrelationships of Placoderms revisited. [Les relations de parent,s des
Placodermes revisit~es]. GEOBIOS, M.S. n 19 : 89-95.
ABSTRACT
From a set of new characters, the phylogeny of placoderms taken as a whole has been investigated using the
computer programs PAUP and Mac Clade. The result corroborate the hypothesis of an inclusion of the Phylolepida
within the Arthrodira and the basal position of the Acanthothoraci among Placodermata.
KEY-WORDS : PLACODERMS,PARSIMONYANALYSIS,PHYLOGENY.
RI~SUMI~
A partir d'un ensemble de nouveaux caract~res, la phylog~nie des Placodermes pris dans leur ensemble a 5t~
estim~e en utilisant les logiciels PAUP et Mac Clade. Le r~sultat conforte l~ypoth~se de l'inclusion des Phyllolepida au sein des Arthrodira et la position basale parmi les Placodermes des Acanthothoraci.
MOTS-CLI~S : PLACODERMES,ANALYSEDE PARCIMONIE,PHYLOGt~NIE.

INTRODUCTION
Since the publication of the Handbook of Paleoichthyology (Denison 1978), the group Placodermi has been subject to increasing interest
among students of paleozoic fishes. Its phylogenetic relationships has been investigated by several
authors either at a general level, in trying to determine its position within gnathostomes (Goujet
1982 ; Gardiner 1984 ; Young 1986 ; Forey &
Gardiner 1986), or to determine the internal relationships among placoderm orders or even within
these orders (Denison 1983, 1984 ; Long 1984 ;
Leli~vre 1991 ; Young 1984, 1988).
During these last t w e n t y years or so our knowledge of the group has rapidly increased through
the description of numerous new taxa ( see Long
1988 ; Dennis & Miles 1979a, 1979b, 1980, 1981,
1982 ; Dennis-Bryant & Miles 1983 ; Goujet
1984) but also with a better understanding of the
fine anatomical features exhibited by better preserved specimens on which acid preparation has
been applicable. The discovery of such localities

like Gogo and other australian localities, South

China (Liu 1991 ; Wang 1991), Morocco (Leli~vre
1984, 1991) and the reinvestigation of the Cleveland Shale fauna (Carr 1991) played a major role
in this revival of Placoderm studies.
Interrelationships of Placodermi have been investigated by several authors, most of t h e m influenced by the evolutionary views implicitly favored
by Denison (1978) : most researchers supposed
the group m a y have started from short-armoured
forms similar to all osteichthyans (with a short
shoulder girdle) and gradually developed a long
body armour leading to the box-like carapace enclosing most of the anterior part of the body like
those present in Antiarcha and Arthrodira. This
view has lead to propose a close relationships between Antiarcha and Arthrodira even if both
groups show m a n y striking differences both in
the skull-roof p a t t e r n and in the composition of
their body armour. This quasi-general agreement
was also due to our f r a g m e n t a r y knowledge of

90
the other placoderm orders (Acanthothoraci, Petalichthyida, Rhenanida, Ptyctodontida, Phyllolepida, Pseudopetalichthyida) most of them being
represented by isolated elements or by a few
complete specimens always used as models for
the entire order in spite of their individual peculiarities.
Recently the effort has been focusing on the Arthrodira, the best known order among Placodermi
but new insight on Phyllolepidida and Acanthothoraci allow us to reinvestigate the relationships
problem within Placoderms and to propose a new
scheme after a parsimony analysis using both
PAUP (Swofford 1991) and Mac Clade (Maddison
& Maddison 1992) computer programs.
Until now, for several placoderm interrelationships schemes which have been proposed
none or very few of t h e m were performed using a
computerised methodology even if these methods
have been applied for Arthrodira, or for Antiarchi
t a k e n as a group.
The provisionary scheme proposed here is a first
approach. This a t t e m p t is biased by a major
question : in order to polarize the character states, an outgroup is needed. In our case which
choice do we have for an outgroup? We are quite
restricted. The phylogenetic position of placoderms within gnathostomes has been subject to
different options. The group has been considered
either as the sister group of Chondrichthyes (cartilaginous fishes : elasmobranchs and holocephalans) or of Osteichthyes (see Goujet 1982 ; Gardiner 1984) the t ~ r d option is to consider them as
the sister-group to all other gnathostomes (Young
1986). In t h a t case the out-group should be searched among the agnathans stock. The Osteostraci, supposed to be the sister-group to all gnathostomes (Janvier 1993) could be the best candidate.
However, this doesn't help much in our problem
since there is very little similarities between placoderms and Osteostraci, preventing us from a
sound comparative study through homologous
features. Our knowledge of both Osteostraci and
Placodermi is quite reasonable but both groups
have been presumably diverging separately a
long way from what could be a common basic design, blurring the comparison. As a temporary solution, processing our data with a hypothetical
ancestor has been choosen, an option included in
the computer programs. By definition, this hypothetical ancestor, which helps in rooting the cladogram, is exhibiting all characters in a plesiomorphous condition.

In most previous hypotheses of placoderm interrelationships, the primitive state of the placoderm dermal skeleton was supposed to be composed of a short shoulder girdle composing a single
row of plates encircling competely, through a median dorsal plate, the anterior part of the body
j u s t behind the skull to which it was connected
by means of a dermal neck-joint. The skull roof
was supposed to exhibit a variable p a t t e r n of dermal components but always including three posterior plates : a nuchal plate flanked by paired
paranuchals bearing the skull component(s) of
the dermal articulation (see Stensi5 1963 ; Goujet
1984).
In all previous phylogenetic schemes, less one
(Goujet 1984), a grouping of Antiarcha and Arthrodira (sometimes called Dolichothoracomorpha, Young 1980) was proposed based on a common long body armour. One of us (Goujet 1984)
questioned this grouping and proposed a separation of the placoderms in two major ensembles
based on several independant (or so supposed)
characters, mainly the position of the nostrils
and the presence of a premedian plate in front of
the snout but also the histological n a t u r e of the
dermal cover and the presence of tesserae - a
character in need of a serious revision to avoid
confusion. This provisional solution was emphasizing the important position of the Acanthothoraci
in the early diversification of placoderms. Until
that, t h e y were more or less considered as oddities among them. The weakness of this previous
analysis is linked to our poor information on the
Acanthothoraci and it resulted in splitting t h a t
group into several units placed as sister groups
to different major groups. Some placed as sistergroups of (Petalichthyida + Ptyctodontida) + (Arthrodires + Phyllolepida) on one side. On the
o t h e r side, some were closer to Antiarcha, and a
third ensemble placed as sister-group to Rhenanida. The Acanthothoraci were considered as a paraphyletic "stem group" within placoderms. This
scheme doesn't find support in the present analysis because the character used then have not
been included : we have restricted ourselves to
the most complete data ; those used in the analysis j u s t cited, taken from fragmentary or uncompletely known specimens, have been revealed unsufficient for the present purpose. For the same
reasons, two groups commonly included in placoderms (i.e. pseudopetalichthyid and Stensioellida)
are not taken into account for the present analysis. They are represented by complete individual
but our information on their dermal skeletton is
two unperfect to lead to meaningful comparisons.

91
,. ~ r,_

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length

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103

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=
o
_.
" ~ "g , ~

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length

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103

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-[
l
I
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3,

length

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--~

103

Tree

4,

_., ~

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Tree

length

103

1
5,

length

103

Figure 1 - The six most

parsimonious placederm
cladograms obtained from
the annexed data matrix.
L e s s i x c l a d o g r a m r n e s les
plus parcimonieux obtenus
p a r t r a i t e r n e n t de la m a trice d e d o n n g e s en annexe.

~.

~i ~

,~ =

Tree

6,

length

103

92
THE NEW ANALYSIS
The present analysis is taking into account a
large set of characters (49) from both cephalic
(endocranial and dermal cover) and body skeleton
(Table 1). The sample of taxa has been selected
to avoid polymorphic states within generally accepted monophyletic taxa (mainly arthrodires and
antiarchs) : even if MAC CLADE and PAUP are
able to accept polymorphic states, in their treatment polymorphism is considered more or less as
an indetermination which m a y have a strange
impact on the final result. Consequently the matrix includes taxa of different levels from orders
to genera. For example, the actinolepid arthrodires are represented by several different genera
exhibiting various skull roof patterns ; for the antiarchs two groups have been used (yunnanolepids, represented by Yunnanolepis and the euantiarchs). This heterogeneity has no impact on the
analysis any longer each of the terminal taxa included is supposed monophyletic. Another possibility would have been to use as basic components only ordinal or subordinal taxa and then to
define for every one a hypothetical ancestral
state for each character. Such a solution has not
been retained since the "ancestral" state should
be a result a posteriori of the analysis and not
an a priori statement.
As already r e m a r k e d elsewhere (Gauthier et al.
1988 ; Lecointre et al. 1993), the taxon sampling
has an important impact on the final result : in a
previous attempt, using the antiarchs as a single
taxon within the analysis, their position was
highly variable due to problems related to the
possible homologies between various plates
When replaced by two components, the antiarchs
always form a monophyletic group with a stable
position inside the global cladogram.
To run PAUP or MAC CLADE the same options
have been chosen and give the same results. All
multistate characters have been treated as unordered when we did'nt have any clue about their
possible transformation. For the binary characters, the reversible option has been retained
which means t h a t we would accept multiple losses and multiple appearance of supposed homologous features. In fact very few if none of the
character states are reverting. The analysis resuits into six equally parsimonious cladograms
(Fig. 1) which shows a high stability in most nodes. In all six, acanthothoracids and Brindabellaspis are grouped together and branch out near
the hypothetical ancestor defined as the placoderm plesiomorphous for all its character states.
Next are the rhenanids which share with their

sister group the presence of one single pair ef

centrals. Then are the antiarchs which share
with their sister group [composed of two ensembles : Petalichthyiformes (Petalichthyida + Ptyctodontida) and Arthrodira] at least six synapomorphies : four (14, 15, 16, 17) relative to the
head and lower jaw ; two relating to the body armour (31 and 35). In all cases the Petalichthyida
are grouped with Ptyctodontida which are characterized by numerous plate losses or bone regression.
The six different cladograms are due to an instability among the arthrodira, but, in all cases they
include the Phyllolepida. This solution confirms
the phylogenetic position of phyllolepids within
the arthrodira promoted by Long (1984).
This most parsimonious solution (treelength 103)
is characterized by reasonably good consistency
(CI = 0.689) and retention (RI = 0.752) indexes
and a low homoplasy index (0.311).
Some important questions remain, among which
those relative to the original position of the nasal
capsules and the nostrils. In the resulting cladogram, dorsal nostrils and the presence of a premedian plate could be either a primitive condition or an anatomical feature appearing very low
in the cladogram j u s t after the initial common
ancestor and lost at the node corresponding to
the common ancestry of ((Petalichthyida + Ptyctodontida) + (Phyllolepida + Arthrodira)). This illustrates the importance of the actual outgroup
to polarize the character transformation series.
In our case, the absence of such an outgroup results in a sheer ambiguity for such important
characters which m a y be interpreted either as
primitive or derived. As a general result, it seems
the initial composition of the body armour plays
a decisive role in shaping the final cladogram. Given the new information collected on the Acanthothoraci body armour, some plates (e.g., the
posterior ventro-lateral plates) were intitially
present in the ancestral body armour pattern.
The discussion about the justification of the character retained in the annexed matrix will be discussed elsewhere but, to keep open the general
discussion, the list of characters and character
states is annexed to the matrix.
As a conclusion, even if this cladogram does not
solve all problems, it gives some strong clues on
the position of some groups traditionally considered as of uncertain affinities : Ptyctodontida and
Phyllolepida can be clearly interpreted as regressive relative to theirs sister-groups respectively
the Petalichthyida and Arthrodira. The Antiarcha

93

.=

!1 t'
]

Consensus

length

103

Figure 2 - The consensus cladogram obtained from the six

preceeding topologies. Multiple branching within the group
[Arthrodira + Phyllolepida] result from the unsufficiency of
the data used here which doesn't help in solving this multiple
branching. Le c[adogramme consensus des six topologies prdc@dentes. Les branchernents multiples au sein de t'ensemble [Arthrodires + Phyllolepides] traduisent l'insuffisance des caract@res retenus ici pour rdsoudre cette rnultichotomie.

t a k e a p l a c e i n b e t w e e n t h e t e s s e r a t e f o r m s tow a r d s t h e t r e e r o o t a n d t h e m o r e " n o r m a l " placoderms.

LIST OF
STATES

CHARACTERS

AND

CHARACTER

1) P a r a n u c h a l p l a t e s , 0 : s e v e r a l , 1 : o n e ;
2) C e n t r a l p l a t e s ( n u m b e r a n d fusion), 0 : sev e r a l , 1 : o n e only, 2 : no , 3 : c e n t r o n u c h a l c o m pound ;
3) C e n t r a l p l a t e s (contacts), 0 : s e p a r a t e , 1 : in
contact ;
4) P r e m e d i a n p l a t e , 0 : a b s e n t , 1 : p r e s e n t ;
5) P o s t m a r g i n a l p l a t e s , 0 : a b s e n t , 1 : p r e s e n t ;
6) S u b o r b i t a l p l a t e , 0 : p r e s e n t , 1 : a b s e n t ;
7) R o s t r a l p l a t e , 0 : p r e s e n t , 1 : a b s e n t ;
8) E n d o l y m p h a t i c pore, 0 : v e r t i c a l , 1 : l o n g oblique tube ;

9) O p e n i n g of n e r v e V I I H m / e p i h y a l facet, 0 :
anterior, 1 : posterior ;
10) A n t e r i o r p o s t o r b i t a l p r o c e s s , 0 : b r o a d , 1 :
narrow ;
11) P o s t e r i o r p o s t o r b i t a l p r o c e s s , 0 : double, 1 :
simple ;
12) S u b p i t u i t a r y f e n e s t r a , 0 : a b s e n t , 1 : p r e s e n t ;
13) L a t e r a l groove or n o t c h on P a r a s p e n o i d , 0 :
absent, 1 : present ;
14) P o s t v a g a l m u s c u l a r d e p r e s s i o n , 0 : deep, I :
e x t e n s i v e , 2 : shallow, 3 : a b s e n t ;
15) I G on m e c k e l i a n c a r t i l a g e , 0 : a b s e n t , 1 : present ;
16) D i f f e r e n c i a t e d b i t i n g a r e a on IG, 0 : a b s e n t , 1:
present ;
17) M e c k e l ' s c a r t i l a g e , 0 : e n t i r e l y p e r i c h o n d r a l l y
ossified, 1 : p o s t e r i o r e l e m e n t only, 2 : t w o s e p a rate elements ;
18) A S G on e n d o c r a n i u m , 0 : a b s e n t , 1 : p r e s e n t ;
19) P S G on P a l a t o q u a d r a t e , 0 : a b s e n t , i : present ;
20) P a l a t o q u a d r a t e , 0 : o n e s i n g l e u n i t , 1 : two
units, 2 : three units ;
21) M e d i a n d o r s a l p l a t e s , 0 : one, 1 : t w o ;
22) M e d i a n d o r s a l s h a p e , 0 : b r o a d a n d s h o r t , 1:
elongate ;
23) V e n t r a l r i d g e or k e e l on m e d i a n d o r s a l , 0 :
absent, 1 : present, 2 : keel with cupula ;
24) P D L p l a t e , 0 : p r e s e n t , 1 : a b s e n t ;
25) P L p l a t e , 0 : p r e s e n t , 1 : a b s e n t ;
26) I L p l a t e , 0 a b s e n t , 1 p r e s e n t , 2 : f u s e d to
AVL;
27) A V p l a t e s , 0 : a b s e n t , 1 : p r e s e n t ;
28) A L p l a t e , 0 p r e s e n t , 1 a b s e n t or f u s e d to
AVL;
29) S p i n a l p l a t e , 0 : p r e s e n t , 1 : a b s e n t ;
30) A M V p l a t e , 0 : p r e s e n t , 1 : a b s e n t ;
31) P M V p l a t e , 0 a b s e n t , 1 p r e s e n t (large), 2
small ;
32) A M V - P M V c o n t a c t , 0 : a b s e n t , 1 : p r e s e n t ;
33) C o n t a c t b e t w e e n A V L ' s , 0 : no, 1 : s h o r t , 2 :
long ;
34) P V L ' s , 0 : a b s e n t , 1 : no c o n t a c t , 2 : l o n g contact ;
35) D e r m a l p e c t o r a l f e n e s t r a , 0 ' o p e n p o s t e r i o r l y
I : closed ;
36) C o n t o u r o f p e c t o r a l f e n e s t r a , 0 " closed b y
PL+PVL, 1 : PL excluded, 2 AVL only (antiarcha);
37) F o r m of p e c t o r a l a r t i c u l a r f a c e t , 0 r o u n d , 1
elongate short, 2 : elongate long ;
38) N u m b e r of p o t e n t i a l r a d i a l s , 0 " t h r e e , 1 "
two, 2 : m o r e t h a n t h r e e , 3 : o n e ;
39) P e l v i c girdle, 0 ' d e r m a l ~: e n d o s k e l , 1 : endoskel, o n l y ;
40) S e n s o r y lines, 0 : g r o o v e s o n l y , 1 : g r o o v e s +
tubes, 2 : tubes with pores ;
41) P o s t e r i o r p i t - l i n e (pp) a n d o c c i p i t a l l i n e (occ),
0 : on s e p a r a t e p l a t e s , I : o n t h e s a m e ;

94

Hyp. ancestor
Yunnanolepis
Euantiarcha
Petalichthyida
Rhenanida
Acanthothoraci
Ptyctodontida
Actinolepids
Aethaspis
Phlyctaeniida
Brachy~horaci
Phyllotepididae
Wuttagoonaspis
i Bri, Mabellaspis

I
0
I
1
0
?
0
1
1
1
1
1
I
1
0

Hyp. ancestor
Yunnanoletois
Euantiarcha
Petalichthyida
Rhenanida
Acanthothoraci
Ptyctodontida
Actinolepids
AethasDis
Phlvctaeniida
Brachythoraci
Phvllolepididae
Wuttagoonaspis

Actinolepids
Aethaspis
Phlyctaeniida
Brachythoraci

18
0
?
0
?
0
1
l
1
1
I
1
0
?

19
0
?
0
?
0
?
1
1
1
1
1
0
?

20
0
?
0
?
0
0
2
?
?
0
I
?
?

36

35
0
1
1
1
0
0
0
1,
1
1
1
i,

Hyp. ancestor
Yunnanolepis
Euantiarcha
Petatichthyida
Rhenanida
Acanthothoraci
Ptyctodontida

0
2
2
1
1
0
I
I
1
I
I
3
3
0

BHndabellaspis

0
?
0
0

0
1
1
0
?
0
0
I
1
I
1
l
1
0

0
0
0
1
0
0
1
0
0
0
0
0
0
0

0
0
0
0
0
0
?
0
I
0
0
1
0
0

0
0
0
0
0
0
?
1
1
1
1
?
1
0

0
?
?
0
?
0
?
0
0
1
1
?
?
0

10
0
?
?
0
0
0
?
0
0
I
1
0
0
0

22
0

23
0

25
0
0
0
0
1
0
1
0
0
0
0
1
0

26
0
1
1
0
I
0
2
0
0
0
0
0
0

27
0
0
0
0
?
0
I
0
1
I
0
0
I

28
0
I
1
0
0
0
0
0
0
0
0
0
0

29
0
1
1
0
0
0
0
0
0
0
0
0
0

30

40
41
42
0
0
0
0
0
?
0
0
0
2
0
1
0
?
0
1
0
0
2
0
1
0
1
2
0
1
2
0
1
2
0
1
2
ii me
i

43
0
0
0
0
0
0
1
0
?
I
1

44
0
0
2
I
0
0
2
1
1
I
1

45
46
0
0
2
1
2
1
1
3
2
?
2
0
?
3
1
2
1
2
1
4
1
4
i n

47
0
0
0
1
2
0
0
1
1
1
I

0
1
1
0
1
1
0
0
0
0
0
0.,
0
I

0
1
0
0
1
0
I
1

1
1
0

2t

0
0
0
0
0
0
1
1
0
0

2
0
0
2
1
?
0
I
?
?

24
0
0
0
0
1
0
0
0
0
0
0
1
0

37
0
0
0
?
0
0
0
1
1
1
2
~

38
0
3
3
?
3
?
1
0
?
0
2
i

39
0
?
1
0
0
0
0
0
?
1
1

0
1
1
0
0
0
0
0
0
0
0
0
0

11
0
?
?
0
?
0
?
0
0
0
1
0
?
0

12
0
?
?
0
0
1
?
1
?
1
I
?
?
1

I l l i l l i

42) C o n v e r g i n g s e n s o r y lines, 0 : n o n c o n v e r g e n t ,
1 : soc and pp, 2 : soc, cc a n d pp o n c e n t r a l plate,
3 : soc cc et pp on n u c h a l p l a t e ;
43) B o d y scale, 0 : large, 1 : n o ;
44) D e r m a l o r n a m e n t , 0 : s t e l l a t e or f l a t t u b e r c u les, 1 : d e n t i n e t u b e r c u l e s , 2 : b o n y r i d g e s or tubercules ;
45) Nostrils, 0 : t e r m i n a l , 1 : v e n t r a l , 2 : d o r s a l ;
46) D e r m a l cervical joint, 0 : l i m i t e d overlap, 1 :
transverse condyle on head, 2 : simple transverse
overlap, 3 : l o n g i t u d i n a l joint, 4 : c o m p l e x glenoid;
47) S c a p u l a r c o m p l e x , 0 : p r e s e n t , I : a b s e n t (diss o c i a t e d plates), 2 : o t h e r p a t t e r n ;

13
0
?
?
?
0
0
?
I
?
I
I
1
?
?

14
0
3
3
1
0
0
?
2
2
3
3
?
2
0

15
16
17
0
0
0
?
?
?
1
1
1
?
?
?
0
0
0
? , ? , ? ,
1
1
1
I
0
?
I
1
?
1
?
?
1
1
2
1
0
?
?
?
?
?
?
?

31
0
1
1
1
0
0

32

1
I
I
0
1
I
0
0
0
0
0
0
0

I
I
1
1
1
1

0
0
0

1
1
1
1
0
1
0

33
0
2
2
1
2
0
0
0
0
0
0
2
0

34
?
2
2
2
1
2
0
2
2
2
2
2
?

Table
1
ters-taxa

Placoderm
characmatrix.
Matrice
de
donndes
charact&es-taxons
pour les P l a c o d e r r n e s .

48
49
0
0
?
0
?
0
0
0
0
1
0
1
0
0
1
0
?
0
1
0
1
0
,i ~]

I l l i i J

48) P a l a t o q u a d r a t e a n t e r i o r a r t i c u l a t i o n , 0 : lateral, 1 : v e n t r a l o n e c t e t h m o i d p r o c e s s o f n e u r o cranium ;

49) T e s s e r a e , 0 : a b s e n t , 1 : p r e s e n t .

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D. G O U J E T
Mus6um national d'Histoire naturelle
Laboratoire de Pal6ontologie et LIRA 12 du CNRS
8 rue Buffon
F-75005 Paris
G.C. YOUNG
Australian Geological Survey Organisation
P.O. Box 378
Camberra ACT 2601, Australia