Conodont Affinity and Chordate Phylogeny

Biol. Rev. (2000), 75, pp.
191–251 Printed in the United Kingdom # Cambridge Philosophical Society 191
Conodont affinity and chordate phylogeny

PHILIP C. J. DONOGHUE", PETER L. FOREY#
and RICHARD J. ALDRIDGE$
" School of Earth Sciences, University of Birmingham, Edgbaston, Birmingham B15 2TT, UK (p.c.j.donoghue!bham.ac.uk)
# The Natural History Museum, South Kensington, London SW7 5BD, UK (plf!nhm.ac.uk)
$ Department of Geology, University of Leicester, University Road, Leicester LE1 7RH, UK (ra12!le.ac.uk)
(Received 4 June 1999 ; revised 29 November 1999 ; accepted 29 November 1999)
ABSTRACT
Current information on the conodonts Clydagnathus windsorensis (Globensky) and Promissum pulchrum Kova! cs–
Endro$ dy, together with the latest interpretations of conodont hard tissues, are reviewed and it is concluded
that sufficient evidence exists to justify interpretation of the conodonts on a chordate model. A new
phylogenetic analysis is undertaken, consisting of 17 chordate taxa and 103 morphological, physiological and
biochemical characters ; conodonts are included as a primary taxon. Various experiments with character
coding, taxon deletion and the use of constraint trees are carried out. We conclude that conodonts are
cladistically more derived than either hagfishes or lampreys because they possess a mineralised dermal
skeleton and that they are the most plesiomorphic member of the total group Gnathostomata. We discuss
the evolution of the nervous and sensory systems and the skeleton in the context of our optimal phylogenetic
tree. There appears to be no simple evolution of free to canal-enclosed neuromasts ; organised neuromasts
within canals appear to have arisen at least three times from free neuromasts or neuromasts arranged within
grooves. The mineralised vertebrate skeleton first appeared as odontodes of dentine or dentine plus enamel
in the paraconodont\euconodont feeding apparatus. Bone appeared later, co-ordinate with the development
of a dermal skeleton, and it appears to have been primitively acellular. Atubular dentine is more primitive
than tubular dentine. However, the subsequent distribution of the different types of dentine (e.g.
mesodentine, orthodentine), suggests that these tissue types are homoplastic. The topology of relationships
and known stratigraphic ranges of taxa in our phylogeny predict the existence of myxinoids and
petromyzontids in the Cambrian.
Key words : Agnatha, Gnathostomata, cladistics, conodont, chordate, craniate, evolution, phylogeny,
vertebrate, skeleton.
CONTENTS
I. Introduction ............................................................................................................................ 192

(1) Poly-, para- and monophyly of the Conodonta ................................................................ 194
II. Soft tissue anatomy.................................................................................................................. 195
(1) The head........................................................................................................................... 197
(2) The trunk.......................................................................................................................... 199
(3) The tail ............................................................................................................................. 199
III. Hard tissues ............................................................................................................................. 199
(1) The histological debate ..................................................................................................... 201
(2) Tissue types ....................................................................................................................... 201
(a) Lamellar crown tissue ................................................................................................. 201
(b) White matter .............................................................................................................. 202
(c) Basal tissue.................................................................................................................. 202
(3) Relative growth of the tissues ........................................................................................... 202
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192 P. C. J. Donoghue, P. L. Forey and R. J. Aldridge
(4) Hard tissue homologies ..................................................................................................... 202

(a) Lamellar crown tissue ................................................................................................. 202
(b) White matter .............................................................................................................. 203
(c) Basal tissue.................................................................................................................. 203
(5) Conodont element growth................................................................................................. 203
(6) Histochemical studies ........................................................................................................ 204
IV. Phylogenetic analysis ............................................................................................................... 204
(1) Taxon sampling ................................................................................................................ 204
(2) Character coding............................................................................................................... 207
(3) Character matrix............................................................................................................... 208
(a) Brain, sensory and nervous systems ............................................................................ 208
(b) Mouth and branchial system ...................................................................................... 211
(c) Circulatory system ...................................................................................................... 211
(d) Fins and fin folds ........................................................................................................ 212
(e) Skeletal ....................................................................................................................... 213
(f) Physiological ............................................................................................................... 215
(g) Miscellaneous.............................................................................................................. 215
(4) Results............................................................................................................................... 215
(a) Experimental analysis of the data set ......................................................................... 219
(b) Cyclostome monophyly............................................................................................... 220
(c) Alternative hypotheses of chordate relationships ........................................................ 220
(d) The effects of alternative interpretations of conodont anatomy and histology ........... 221
(e) Testing alternative hypotheses of conodont affinity.................................................... 223
(f) Summary of conclusions drawn from experimental analyses of our data set.............. 225
(5) Character changes............................................................................................................. 225
(a) General ....................................................................................................................... 225
(b) Nervous and sensory systems ...................................................................................... 225
(c) Skeleton ...................................................................................................................... 233
V. Conodont affinity..................................................................................................................... 236
VI. Directions for future research .................................................................................................. 238
(1) General.............................................................................................................................. 238
(2) The origin of the Conodonta ............................................................................................ 238
(3) Euconodont phylogeny...................................................................................................... 239
(4) Histology ........................................................................................................................... 239
(5) Microevolution.................................................................................................................. 240
(6) Conodont element function............................................................................................... 240
VII. Revised classification ............................................................................................................... 240
VIII. Conclusions .............................................................................................................................. 240
IX. Acknowledgements .................................................................................................................. 241
X. References................................................................................................................................ 241
XI. Appendix ................................................................................................................................. 249
(1) Character diagnostics ........................................................................................................ 249
I. INTRODUCTION Klussendorf, 1985 a, b ; Aldridge et al., 1993 ; Gabbott,

Aldridge & Theron, 1995) has resulted in a
‘ The problem of the zoological affinities of this group revolution in our understanding ; we now know that
remains … one of the most fascinating and perplexing conodonts are a group of eel-shaped animals with
problems of palaeozoology ’ (Rhodes, 1954, p. 419). the phosphatic elements constituting a complex
feeding apparatus. Although the systematic position
When Frank Rhodes wrote his article on conodont of conodonts is still contentious, the diversity of
relationships for Biological Reviews in 1954, conodonts opinion has narrowed greatly. Just a year before the
were known exclusively in the form of phosphatic first conodont fossil with preserved soft tissues was
tooth-like microfossils of unknown or uncertain found, Mu$ ller (1981) compiled a list of groups to
affinity. The perception of conodonts is now very which conodonts had been attributed ; his list
different. The discovery of soft tissue remains (Briggs, includes at least three kingdoms and almost every
Clarkson & Aldridge, 1983 ; Mikulic, Briggs & major animal phylum. The debate is now much
Conodont affinity and chordate phylogeny 193
more tightly constrained (see Aldridge, 1987 ; a bilaterally disposed grasping array in what is
Aldridge et al., 1993) and almost all authorities agree interpreted as the cephalic region. A longitudinal
that conodonts are chordates, although a few gut, possible ovaries and a laterally disposed ray-
workers still advocate a chaetognath affinity (e.g. supported caudal fin are also preserved. The overall
Kasatkina & Buryi, 1996 a, b, 1997). Among those anatomy of T. rokycanensis is distinctly chaetognath-
authors who accept a chordate assignment, however, like and quite dissimilar to preserved remains of
there is still controversy regarding the precise place conodonts.
of the conodonts within the chordate clade. The interpretation of early Cambrian conodonti-
To conduct a cladistic analysis involving both form elements known as protoconodonts as the
chordates and chaetognaths is currently not feasible, grasping spines of chaetognaths (Szaniawski, 1982)
as the affinity of chaetognaths themselves is highly provides another possible line of evidence, if the
contentious. Some authors suggest that they are best evolutionary sequence from protoconodonts,
placed as a sister-group to the vertebrates through paraconodonts to euconodonts (the true
(Christoffersen & Arau! jo-de-Almeida, 1994), conodonts) proposed by Bengtson (1976 ; Fig. 1)
whereas others doubt even their deuterostome were sustained. This linkage was further emphasised
affinity (Halanych, 1996 ; Nielsen, 1995 ; Nielsen, by Bengtson (1983 a, b) and Szaniawski (1987), who
Scharff & Eibye-Jacobsen, 1996). A relationship noted that the hypothesis that chaetognaths and
between conodonts and chaetognaths was first (eu)conodonts shared a common ancestor did not
suggested by Rietschel (1973) and was resurrected necessarily conflict with conodonts being closely
by Briggs et al. (1983) as one of the possible related to the chordates. Andres (1988) has also
interpretations consistent with the features of the suggested a reconciliation between the chordate and
first known conodont with preserved soft tissues chaetognath hypotheses of affinity, proposing chaeto-
(IGSE [British Geological Survey, Keyworth] gnaths as ancestors to euconodonts, which are in
13822). This interpretation was more strongly turn ancestral to the vertebrates. This suggestion is
advocated by Bengtson (1983 b), and is supported by consistent with a close relationship between chaeto-
Kasatkina & Buryi (1996 a, b, 1997). However, the gnaths and chordates (Christoffersen & Arau! jo-de-
resemblances between conodont and chaetognath Almeida, 1994 ; Cavalier-Smith, 1998 ; Hall, 1998),
anatomy are vague ; both are small and bilaterally but at odds with recent molecular and morphological
symmetrical, and both bear ray-supported fins, but work which supports the view that chaetognath
the caudal fin of conodonts is dorso-ventrally rather affinity lies amongst the protostomes (Telford &
than laterally disposed as in chaetognaths (contra Holland, 1993 ; Philippe, Chenuil & Adoutte, 1994 ;
Kasatkina & Buryi, 1996 a). Furthermore, the V- Wada & Satoh, 1994 ; Nielsen, 1995 ; Nielsen et al.,
shaped structures apparent in the trunk of conodonts 1996 ; Halanych, 1996 ; Zrzavy! et al., 1998 ; current
have no parallel in the musculature of chaetognaths, evidence suggests that chaetognaths are closely
although there is some resemblance to the ovaries of related to the nematodes and gnathostomulids,
some taxa [e.g. Pterosagitta draco (Krohn, 1853)]. possibly forming a monophyletic group, see
The possibility remains that the taphonomic state Littlewood et al., 1998). Until the affinities of
of specimens preserving conodont anatomy masks chaetognaths are resolved, the relative position of
similarities between the two groups, but, as no conodonts and chaetognaths remains untestable,
taphonomic studies of chaetognaths have been and an apparent morphological gulf between proto-
undertaken, this remains difficult to assess. Com- conodonts and paraconodonts limits the applica-
parison of fossil conodonts with fossil chaetognaths is bility of the evolutionary model forwarded by
limited by the rarity of fossil chaetognath specimens. Bengtson (1976 ; Fig. 1).
Amiskwia sagittiformis Walcott from the Middle Aldridge & Purnell (1996), in a review of the
Cambrian Burgess Shale was originally described as debate, considered six competing hypotheses put
a chaetognath, but has been reinterpreted as a forward in the recent literature for the systematic
possible nemertine (Conway Morris, 1977). position of conodonts amongst the living chordates
Paucijaculum samamithion Schram from the mid- (Dzik, 1995 ; Kemp & Nicoll, 1995 a, 1996 ; Peterson,
Pennsylvanian Francis Creek Shale of Illinois has 1994 ; Aldridge et al., 1986, 1993 ; Gabbott et al.,
been similarly reassessed (Richardson 1982). More 1995 ; Janvier, 1995). Although most of these
recently, Kraft & Mergl (1989) have described a hypotheses have been couched in cladistic termin-
more convincing candidate, Titerina rokycanensis from ology or expressed in the form of cladograms, not
the Lower Ordovician of Bohemia, which preserves one is the result of a numerical cladistic analysis.
A B C
Fig. 1. Proto- (A), para- (B) and eucondont (C) grades of organisation (after Bengtson, 1976, and Szaniawski &
Bengtson, 1993). (A) Different shadings distinguish three layers which have no homologues in B and\or C. (B, C)
Dark shading represents basal tissue of euconodonts and its putative homologue in paraconodonts ; light shading
represents lamellar crown tissue of euconodonts.
Instead, each hypothesis represents the result of divided into three main groups on histological
placing conodonts in a pre-existing cladogram using criteria by Mu$ ller & Nogami (1971, 1972) and
pre-established synapomorphies, or classifies them Bengtson (1976 ; Fig. 1). The stratigraphically oldest
according to unsubstantiated a priori assumptions of group, the protoconodonts, are represented by
character polarity in chordate phylogeny, or places dominantly organic spine-like fossils which are
them within a phylogeny based on a consideration of multilayered and grew by adding new layers to the
limited parts of the anatomy. This is unfortunate, as concave side of the elements only (Fig. 1 A).
conodonts clearly have an important contribution to Protoconodont elements bear a strong histological
make to our understanding of chordate evolution. resemblance to chaetognath grasping spines
Their significance centres around the feeding (Szaniawski, 1982) and all available evidence indi-
elements, which represent the only biomineralised cates a probable link between these two groups
component of the conodont skeleton. Depending on (Szaniawski, 1983, 1987). Paraconodont elements
whether conodonts are acraniate or craniate are also relatively poorly mineralised ; in growth,
chordates, their elements are either an interesting each successive phosphatic layer accreted onto the
but esoteric feature (in the former case), or an margins and base, but not the tip of its predecessor
important clue to the development of the mineralised (Szaniawski & Bengtson, 1993 ; Fig. 1 B). These
skeleton in craniate phylogeny (in the latter case). elements bear a strong resemblance to the basal
The main aim of this contribution is to determine body of euconodont elements (Lindstro$ m, 1964 ;
the systematic position of the Conodonta through Bengtson, 1976). Euconodont elements are com-
formal cladistic analysis. To this end, we briefly posed of two structures : the crown, which is very
review interpretations of the anatomy and histology heavily mineralised and relatively coarsely crys-
of conodonts in order to explain our coding of talline, and the basal body which is finely crystalline
characters in the data matrix on which our analysis and contains more organic material (Fig. 1 C). Both
is based. First, however, it is pertinent to consider the components grew by external apposition of new
monophyly of the Conodonta and to establish layers of mineral on the outer surface, which were
precisely what constitutes a conodont. either added synchronously (Mu$ ller & Nogami,
1971) or at least in step (Lindstro$ m & Ziegler, 1971).
The lack of demonstrated links between proto-
conodonts and paraconodonts and the evidence that
(1) Poly-, para- and monophyly of the
the former are related to chaetognaths means that it
Conodonta
is likely that protoconodonts belong to a clade of
Microscopic phosphatic cones, or ‘ conodontiform ’ animals distinct from euconodonts (see also Peterson,
fossils, first occur in the uppermost Precambrian and 1994). An evolutionary link between paraconodont
become diverse in Cambrian strata. They were and euconodont elements has been more firmly
Paraconodontida
Proconodontida
Panderodontida
unnamed group
unnamed group
Ozarkodinida
Prioniodinida
Teridontus
Pr
ot
op
a
tid
an
on
de
od
ro
do
ni
io
nt
Pr
id
a
? ? ?
Fig. 2. Attempted reconstruction of the relationships of the major groups of euconodont-grade taxa, based on the
suprageneric scheme and proposed relationships of Sweet (1988). This is not a cladogram but our interpretation of
the views expressed by Sweet (1988) in the form of a phylogenetic tree. The three orders of complex conodonts are
generally given equal taxonomic rank. However, it is implicitly accepted that the Ozarkodinida and Prioniodinida
are derived from taxa classified within the Prioniodontida. Therefore, we have chosen to express this relationship
explicitly in the figure and in the text.
established, primarily on the similarity in growth unclear. The Teridontus lineage includes the over-
pattern. Szaniawski & Bengtson (1993) have whelming majority of conodonts and, although the
attempted to demonstrate some transitional forms in maximum diversity of this clade was also attained in
which layers of crown tissue were added to the basal the Ordovician, it is known to have survived through
body only in late ontogeny. More recently, Mu$ ller & to the end of the Triassic Period. The three taxa
Hinz-Schallreuter (1998) have documented the known with preserved soft tissue features all belong
histology of a number of Cambrian paraconodonts to the Teridontus lineage, as do the vast majority of
which possess microstructural features comparable taxa that have been studied histologically. Given the
with the basal tissue of euconodonts. The link uncertain and probable independent origins of
between paraconodonts and euconodonts points to Proconodontus and Teridontus, we restrict our phylo-
an origin for the whole group in the Early or Mid genetic analyses of euconodonts to the monophyletic
Cambrian and renders the aptly named Para- lineage first represented by Teridontus.
conodontida a paraphyletic assemblage unless it is
expanded to include the Euconodonta. However,
monophyly of the Euconodonta is far from certain
and not all taxa currently regarded as euconodonts II. SOFT TISSUE ANATOMY
necessarily fall within an inclusive Paraconodontida.
The early evolutionary history of the Euco- The soft tissue anatomy of conodonts is known from
nodonta, or true conodonts, has been most clearly at least three euconodont taxa : Clydagnathus
documented by Miller (1969, 1980, 1984) who windsorensis (Globensky) from the Lower Carbon-
distinguished two distinct lineages that are first iferous Granton Shrimp Bed of Edinburgh, Scotland
represented in the stratigraphic record by Procono- (Aldridge et al., 1993) ; Panderodus unicostatus (Branson
dontus and Teridontus (Fig. 2). Both lineages are of & Mehl) from the Lower Silurian of Waukesha,
uncertain ancestry and both made their first ap- Wisconsin (Mikulic et al., 1985 a, b ; Smith, Briggs &
pearance in the late Cambrian. The Proconodontus Aldridge, 1987), and Promissum pulchrum Kova! cs-
lineage is known to have flourished in the Early Endro$ dy from the Upper Ordovician Soom Shale of
Ordovician but its subsequent evolution remains South Africa (Aldridge & Theron, 1993 ; Gabbott et
Fig. 3. For legend see opposite.
al., 1995). There may be additional taxa represented to distinguish between otic and optic capsules (see
in the specimens from the Granton Shrimp Bed. Donoghue, Purnell & Aldridge, 1998), and in
specimen IGSE 13821\2 from Granton, the lobes lie
anterior to a smaller pair of round structures which
(1) The head
have themselves been interpreted as otic capsules
The most frequently preserved soft tissues are a (Aldridge et al., 1993 ; Aldridge & Donoghue, 1998 ;
single pair of lobate structures, which occur com- Donoghue et al., 1998). There is, furthermore, no
monly in association with natural assemblages of evidence that otic capsules are more likely to be
conodont elements in the Soom Shale and are also preserved than optic capsules ; indeed, remnants of
evident in two of the specimens from Granton. The eyes have been reported in all fossil representatives of
lobes occur as carbonised remains that are round to soft-bodied jawless vertebrates. In contrast, the
oval in outline with a thickened or darker rim, and remains of unmineralised otic capsules are rare and
have been reconstructed to three dimensions as a normally limited to moldic preservation (e.g.
pair of outwardly expanding cups (Aldridge et al., Myxinikela ; Bardack, 1991).
1993 ; Aldridge & Theron, 1993). From the Granton The presence of extrinsic eye musculature in
specimens it is evident that these cups were located specimen GSSA C721 is supported by evidence from
slightly in front of and above the feeding apparatus additional, recently discovered specimens of
(Fig. 3 A–C). Aldridge & Theron (1993) drew Promissum pulchrum that also display paired lobes
comparison with similar structures in the head of the composed of fibrous white clay minerals. Although
fossil jawless vertebrate Jamoytius kerwoodii White, no other soft tissue remains are preserved in these
where they have been interpreted as eye capsules specimens, the lobate patches occur in consistent
(Ritchie 1968). Similar structures, interpreted as topological association with natural assemblages of
traces of retinal pigments, have also been recorded in elements, indicative of an original position above
a number of entirely soft-bodied fossil jawless and in front of the feeding apparatus. In a few
vertebrates including Myxinikela (Bardack 1991, specimens, the patches are preserved only on the
1998), Mayomyzon (Bardack & Zangerl, 1968, 1971) part, but are matched by lobate traces of organic
and Hardistiella (Janvier & Lund, 1983 ; Lund & film on the counterpart that are identical with the
Janvier, 1986). The most completely preserved structures interpreted as eye capsules on other
specimen of Promissum pulchrum (GSSA [Geological material. In extant lampreys, the extrinsic eye
Survey of South Africa, Pretoria] C721) includes muscles also match the eye capsules in their
oval patches of white tissue in a similar position distribution (see e.g. Fig. 3.6.I in Janvier, 1996 a).
relative to the feeding apparatus to that occupied by Scanning electron micrography of the white patches
the carbonised lobes in other specimens. This white reveals a fibrous microstructure similar to the
tissue has a fibrous texture comparable with preserved trunk musculature of specimen GSSA
preserved muscle tissue in the trunk of the same C721 (Fig. 4 F–H).
specimen, and was interpreted as extrinsic eye In our codings for conodonts we have accepted the
musculature by Gabbott et al. (1995). growing evidence for the presence of extrinsic eye
The interpretation of these structures as eye musculature and we have taken the presence of
capsules and eye musculature has been criticised by paired sensory organs (eyes and, perhaps, otic
Pridmore, Barwick & Nicoll (1997), who argued capsules) to indicate the presence of a brain.
that the paired lobes more probably represent otic Specimen IGSE 13821\2 also shows faint traces of
capsules. Pridmore et al. (1997) based their argu- approximately five paired rectangular box-like
ments on the size, shape and position of the lobes and structures posterior to the paired capsules (Fig. 3 A
on an assertion that otic capsules are more likely to in Briggs et al., 1983 ; Fig. 3 C). Each of these
be preserved in the fossil record than optic capsules. structures is oriented with its long axis transverse to
However, size and shape are inadequate characters the body axis. These compare well in shape and
Fig. 3. (A–E) Clydagnathus windsorensis (Globensky). (A, B) Part and counterpart of whole animal (respectively IGSE
13821 & 13822) ; frame widths 16 mm and 18 mm respectively. (C) Head region of part (IGSE 13821) ; frame width
25 mm. (D) Bilobed tail of IGSE 13821 ; frame width 7 mm. (E) Representative portion of the trunk (RMS GY
1992.41.1) ; frame width 10.5 mm. (A, C, D from Briggs et al. 1983 ; B, E from Aldridge et al. 1993). A, C, D are
reproduced from Briggs et al. (1983) with the permission of the Lethaia Foundation.
Fig. 4. (A–H) Promissum pulchrum Kova' cs-Endro$ dy. (A, B) Part and counterpart of only known specimen preserving
trunk musculature (GSSA C721b, a respectively) ; frame widths 106 mm and 112 mm respectively. (C) Head region
including feeding apparatus and preserved extrinsic eye musculature (GSSA C721a) ; frame width 16 mm. (D, E)
Scanning electron micrograph of trunk musculature (GSSA C721a) ; D frame width 74 µm ; E frame width 22 µm.
(F–H) Extrinsic eye musculature under progressively higher magnification (GSSA C1320) ; F frame width 2900 µm ;
G frame width 930 µm ; H frame width 360 µm. B, D, E reproduced from Gabbott et al. (1995), with the permission
of Macmillan Magazines Ltd.
position to gill pouches in the fossil lamprey the most extensive development of this fin was on the
Mayomyzon (Aldridge & Donoghue, 1998), and in dorsal or the ventral margin of the animal, but our
our coding we follow Aldridge et al. (1993) in their re-examination of IGSE 13822 indicates that the
tentative interpretation of these structures as gill bilobed portion occurs on the dorsal margin (Fig.
pouches. 3 D). The second specimen (RMS [Royal Museum
of Scotland, Edinburgh] GY 1992.41.3) preserves an
(2) The trunk unequally developed ray-supported fin that is longer
on the dorsal margin. The shape of the bilobed fin
Trunk muscle tissues occur as distinct anteriorly
compares to that of lampreys, where it is traditionally
directed chevrons in the Granton specimens ; their
interpreted as hypocercal. Janvier (1998) has re-
discrete nature is presumably the result of post mortem
cently compared the tail of RMS GY 1992.41.3 with
decay and shrinkage, a process recognised in
that of Myxine glutinosa, which he interprets as
decomposition experiments on Branchiostoma (Briggs
cryptically hypocercal. Here, we regard the tail of
& Kear, 1994). The architecture of the muscle
conodonts to be hypocercal because the bilobed fin is
blocks appears to be V-shaped, although the possi-
restricted to the dorsal margin of the notochord, but
bility remains that the myomeres were originally W-
we do not accept Janvier’s (1998) argument as there
shaped with only the central V preserved (Donoghue
is no evidence of a ventral bend of the conodont
et al., 1998). In modern ammocoetes the myomeres
notochord at the caudal termination.
are W-shaped, but the dorsal limbs of the muscle
blocks are extremely short (P. C. J. Donoghue, pers.
III. HARD TISSUES
obs.). Although there is some evidence for individual
fibre preservation in one of the Granton specimens
The mineralised skeleton of conodonts is limited to
(RMS GY 1992.41.3 ; Aldridge et al., 1993) muscle
the phosphatic elements that are generally inter-
ultrastructure is best preserved in P. pulchrum (GSSA
preted to have comprised a feeding apparatus. The
C721), in which each myomere is demonstrably
composition and architecture of this apparatus is
composed of fibril bundles (Fig. 4 E), together with
only fully understood for a few taxa, almost all of
possible sarcolemmic membranes and collagenous
which are highly derived, within the context of
connective tissues (Gabbott et al., 1995). This
conodont phylogeny as it is understood currently
specimen also includes a black organic patch within
(Fig. 2). We know very little of the apparatuses of
the mid-trunk region that is probably the remains of
the earliest conodonts, with the simplest fully
a visceral organ. Paired axial lines interpreted as the
reconstructed apparatus being that of Panderodus
margins of a notochord (Aldridge et al., 1993), run
which has cone-like elements of similar morphology
the length of the trunk in all Granton specimens.
to those of primitive conodonts. The apparatus of
This interpretation has been corroborated by ex-
Panderodus was bilaterally arranged with eight pairs
perimental decay of Branchiostoma, in which the
of opposed elements and one symmetrical
notochord decomposes to leave the notochordal
‘ symphysial ’ element lying on the axis of symmetry
sheath, which then collapses to a pair of linear
(Smith et al., 1987 ; Sansom, Armstrong & Smith,
thickened margins (Briggs & Kear, 1994). Remains
1994). These elements are considered to have
of a notochord have not been found in association
performed a grasping function (Smith et al., 1987 ;
with Promissum pulchrum, but its position has been
Sansom et al., 1994 a), although this hypothesis
considered to be represented by a 2 mm gap between
remains to be tested.
the dorsal and ventral limbs of the myomeres
More derived conodont groups typically possessed
(Gabbott et al., 1995). A possible nerve cord has been
elements of more complex morphology and their
identified stopping short of the paired lobes in two of
apparatuses were composed from morphologically
the Granton specimens (Aldridge et al., 1993).
distinct groups of elements. Almost all ‘ complex
conodonts ’ belong to the Prioniodontida and many
(3) The tail
of these fall within the Ozarkodinida, the most
The tail is preserved in only two of the 10 specimens derived of all conodont groups (Fig. 2). The
from Granton, one of which (IGSE 13822) displays architecture of non-ozarkodinid prioniodontids has
an asymmetrical bilobed ray-supported fin. The fin only been fully resolved for Promissum pulchrum,
rays are closely set, without apparent branching, which possessed nineteen elements, divided into four
and there is no evidence for supporting musculature. pairs of robust ‘ P ’ elements which lay behind a pair
Briggs et al. (1983) were unable to determine whether of ‘ M ’ elements, and above an array of ‘ S ’ elements
Fig. 5. For legend see opposite.
comprising four pairs of asymmetrical elements and hard tissues of conodonts and vertebrates as homo-
a single, axial symmetrical element (Aldridge et al., logues (Barskov, Moskalenko & Starostina, 1982 ;
1995). The function of this apparatus is not well Dzik, 1986 ; Burnett & Hall, 1992 ; Sansom et al.,
understood but S and M elements are interpreted as 1992 ; Sansom, Smith & Smith, 1994 ; Sansom, 1996 ;
having grasped prey (Aldridge et al., 1987) that was Smith, Sansom & Smith, 1996 ; Donoghue, 1998),
sheared and crushed by the P elements prior to and those who seek to refute this hypothesis (Kemp
digestion (Aldridge et al., 1995). & Nicoll, 1995 a, b, 1996 ; Schultze, 1996).
The apparatus of the most derived conodont
group, the Ozarkodinida, is known from several taxa
(see Purnell & Donoghue, 1998, for a review). The (2) Tissue types
ozarkodinids had sets of M, S and P elements, but, in The crown of euconodont elements is either com-
contrast to P. pulchrum, these units were arranged posed entirely from lamellar crown tissue, or includes
linearly from rostral to caudal (Aldridge et al., 1987 ; a core of opaque cancellous tissue known as white
Purnell & Donoghue, 1997, 1998 ; Purnell, matter. The basal body has also been interpreted as
Donoghue & Aldridge, 2000). Functional hypotheses a two-component structure (Gross, 1957, 1960), but
suggest that the S and M elements were used for is currently regarded as a single tissue, termed basal
grasping, while a shearing and crushing function for tissue by Donoghue (1998). Biomineralised basal
the P elements has been corroborated by studies of tissue is not preserved in the majority of post-
microwear and functional morphology (Purnell, Devonian taxa, although pathological features in
1995 ; Donoghue & Purnell, 1999 b). some elements suggest that a basal body was present
in life (Donoghue, 1998).
(1) The histological debate
(a) Lamellar crown tissue
For the first 127 years of conodont research element
morphology and histology were the only clues to Within an element of any one taxon, lamellar crown
affinity (for a review see Donoghue, 1998). Without tissue exhibits a variable microstructure that is,
independent constraint over the phylogenetic pos- nevertheless, consistent between homologous
ition of conodonts, however, attempts at compara- portions of homologous elements (Donoghue, 1998).
tive histology proved futile. The discovery and The crystallites that build the lamellae are typically
interpretation of soft tissue remains has now provided a few microns long but range from less than 1 µm to
a context within which conodont hard tissue his- more than 30 µm in length ; these layers of crystallites
tology can be assessed. Although there is a range of are bounded at each end by incremental growth
opinion among those who have published histo- lines (Fig. 5 B, C). In some of the taxa referred to the
logical results, recent contributors to the debate fall Order Prioniodinida by Sweet (1988) the crystallites
clearly into two schools : those who interpret the are fibre-like in appearance, typically reaching 30
Fig. 5. Histology of conodont hard tissues. (A) Element of Coryssognathus dubius (Link & Druce) exhibiting the division
into crown and basal body ; (BU [Birmingham University, Lapworth Museum of Geology] 2616) ; frame width 547
µm. B,C. Lamellar crown tissue. (B) Horizontal section through a Pa element of Ozarkodina confluens ; (BU 2621) ; frame
width 78 µm. (C) Transverse section through a Pa element of Idiognathodus sp. ; (ROM [Royal Ontario Museum,
Toronto] 53261) ; frame width 124 µm. (D, G, H) Recurrent patterns of surface microwear compared to internal
discontinuities. (D) (ROM 49777) ; frame width 224 µm, inset element 698 µm in maximum length. (G) (ROM
49780) ; frame width 344 µm, inset element 1675 µm in maximum length. (H) (ROM 53445) ; frame width 352 µm.
(E, F) Basal tissue. (E) Tubular microstructure comparable to mesodentine in the Middle Ordovician Neocoleodus ;
(BU 2257) ; frame width 270 µm. (F) Globular and lamellar microstructure in Drepanodus from the Arenig of Estonia :
left frame in Nomarski differential interference, right frame in cross-polarised light ; (BU 2694) ; total frame width
250µm. (I, J) White matter. (I) Scanning electron micrograph of an etched section through a Pa element of Ozarkodina
confluens from the Upper Silurian of Gotland, Sweden ; (BU 2615) ; frame width 27 µm. (J) Pa element of O. confluens
immersed in optical oil demonstrating the relationship between white matter (the cancellar denticle core) and the
surrounding lamellar crown tissue ; (BU 2627) ; frame width 134 µm. (K, L) A natural pair of Idiognathodus Pa elements
dissected from the articulated skeletal remains of an individual conodont, articulated in the opposing extremes of their
occlusal cycle and viewed from the cauda ; (BU 2683) ; maximum length of dextral element 1182 µm. (M) Sc element
of Carniodus carnulus from the Lower Silurian of Estonia, immersed in optical oil and demonstrating the composite
nature of its struture ; (BU 2628) ; frame width 414 µm, maximum length of inset 890 µm.
µm or more in length with growth increments that quently demonstrated that white matter and la-
are only weakly discernible (Donoghue, 1998) ; the mellar crown tissue share a common microstructural
arrangement of these fibres varies throughout in- fabric which, together with evidence of synchronous
dividual elements. growth suggests that both tissues are products of a
common developmental process.
(b) White matter
(4) Hard tissue homologies
White matter is more finely crystalline than lamellar
crown tissue and is also distinguished by its albid Given the evidence for conodont affinities based on
appearance in reflected light. The opacity of the soft tissue anatomy, the hard tissues can now be
tissue results from the inclusion of small spaces (Fig. assessed within a chordate framework. Ascidiacean
5 I), ranging in size from less than 1 µm to more than and soberacean tunicates are capable of phosphatic
30 µm in maximum dimension. White matter is also biomineralisation, but their spicules are homogenous
more resistant to etching agents than the enveloping and composed of amorphous deposits or, in some
lamellar crown tissue (Stauffer & Plummer, 1932) cases, dahllite. Extant myxinoids and lampreys
and as a result the distinction between the two tissues biomineralise phosphate in the form of statoliths
becomes more obvious in etched sections. Despite (Carlstro$ m, 1963) which are composed of an
these differences, some specimens clearly demon- amorphous (polyhydroxyl) form of calcium phos-
strate that these two tissues grew synchronously phate (Donoghue, 1998) ; these structures, again, do
(Donoghue, 1998 ; Fig. 5 J). The boundary between not compare histologically with conodont elements.
the two tissues appears transitional in transmitted There is some evidence that lampreys are capable of
light, but there is a sharp distinction between them endoskeletal biomineralisation (Bardack & Zangerl,
in etched section. 1971 ; Langille & Hall, 1993). In myxinoids, how-
ever, only the keratinous cap of the lingual teeth is
mineralised, and this mineralisation is limited to
(c) Basal tissue
isolated crystals embedded in a keratin matrix
Basal tissue is the most variable of conodont hard (Dieckwisch & Vahadi, 1997). Although this may
tissues. It almost always incorporates incremental prove to be of fundamental importance to our
growth lines, but may be internally globular (Fig. understanding of the origin of teeth and of the
5 F), or tubular (Fig. 5 E), or globular and tubular, vertebrate dermal skeleton as a whole, it is not
or lacking in both globules and tubules (Donoghue, possible to suggest primary homologies between the
1998). mineralised hagfish teeth and the hard tissues of
conodonts.
(3) Relative growth of the tissues
(a) Lamellar crown tissue
Crown tissue is known to have grown by outer
apposition because many elements exhibit evidence The relatively coarse crystalline microstructure of
of damage and subsequent repair (Furnish, 1938 ; lamellar crown tissue shares no similarity with the
Hass, 1941 ; Donoghue, 1998 ; Donoghue & Purnell, hard tissues of invertebrate chordates, even if we
1999 a ; Fig. 5 D, G, H). Incremental growth lines in consider the tissue in isolation (Donoghue, 1998).
the lamellar crown and basal tissue meet at the basal The tissue is, however, closely comparable with
body-crown junction indicating that the two tissues enamel and enameloid, the hypermineralised tissues
grew in synchrony (Mu$ ller & Nogami, 1971), and of vertebrate teeth and scales. The lamellar crown
hence basal tissue also grew by outer apposition. The tissue of the vast majority of conodont elements
innermost core of any element therefore represents compares most closely to enamel in its coarsely
the earliest growth stage, and the outermost the crystalline microstructure and punctuating incre-
latest. The two tissues grew in opposing directions mental growth lines (e.g. Fig. 5 B). It has been
relative to the crown-basal body junction. suggested that the degree of microstructural vari-
Donoghue (1998) used the architecture of the ation apparent both within and between conodont
putative cell and cell-process spaces in white matter taxa, and even within an individual element, is
to argue that this tissue and lamellar crown tissue incompatible with an interpretation of this tissue as
had grown in synchrony, but in opposing directions. an enamel homologue (Forey & Janvier, 1993). This
However, Donoghue & Chauffe (1999) have subse- variation, however, coincides precisely with the
requirements of element function and relates to the a homology with bone is unlikely. A homology with
different biomechanical forces that were imposed dentine is more sustainable, as the full range of
upon the elements during feeding (e.g. Donoghue & conodont basal tissues from lamellar atubular, to
Purnell, 1999 b). Similar controls can be recognised tubular, to calcospheritic can be encompassed within
in the enamel microstructure of derived mammals various dentine types (Donoghue, 1998). A plausible
(v. Koenigswald, 1997), indicating that the enamel argument has been presented for the presence of
of both mammals and conodonts was influenced by globular calcified cartilage in the basal body of
analogous selective pressures (Donoghue & Purnell, Cordylodus angulatus (a member of the Proconodontus
1999 b). Although by strict definition conodonts were lineage according to Miller, 1984) by Sansom et al.
not teeth (Donoghue, 1998) they functioned in a (1992), through comparison with the endoskeletal
manner which is directly analogous (Purnell, 1995 ; cartilage found in association with the dermal
Donoghue & Purnell, 1999 a, b). The general lack of skeleton of the Ordovician vertebrate Eriptychius
complex enamels in lower vertebrates probably (Denison, 1967). The microstructure of the basal
reflects the fact that their teeth were not performing tissue in C. angulatus, however, falls well within the
complex occlusal functions. Some exceptions occur, range exhibited by dentine (e.g. Plate 3 and Fig. 3 in
such as Uromastyx (the agamid lizard ; Cooper & Sansom et al., 1997 ; see Donoghue, 1998). There is a
Poole, 1973) which has independently evolved problem in resolving between the microstructures of
prismatic enamel in parallel with a complex occlusal cartilage and dentine in fossils as small as conodont
dentition. Similarly, sharks have independently elements, as features that might distinguish cartilage,
evolved a differentiated enameloid microstructure in such as prismatic structure or an approximation of
the dentitions of at least two distinct lineages, in it, occur in other taxa on a much larger scale.
response to dental stresses (Reif, 1973, 1979, 1981 ; However, examination of the calcospheritic structure
Preuschoft, Reif & Mu$ ller, 1974). in the basal tissue of additional specimens of C.
angulatus and of Drepanodus arcuatus reveals that the
calcospheres often appear to have grown inde-
(b) White matter
pendently and are rarely enveloped within the
White matter has been the most controversial of all lamellae in the manner displayed by calcospheritic
the component hard tissues of conodont elements. dentines. The weight of evidence at present, there-
Sansom et al. (1992) interpreted it as cellular dermal fore, indicates that one group of Lower Ordovician
bone, but this is inconsistent with the growth inter- conodonts (of the Proconodontus lineage) bore
relationship now recognised between lamellar crown elements partially composed from mineralised car-
tissue and white matter (Donoghue, 1998). White tilage. Most basal tissues, however, are purely
matter was secreted by the same cell population that lamellar and atubular, and resemble lamelin
produced lamellar crown tissue (Donoghue & (Donoghue, 1998), a form of dentine first described
Chauffe, 1999) and thus it can be neither a dentine from a Silurian chondrichthyan (Karatajute-
nor an enameloid tissue. It is more likely that white Talimaa et al., 1990). In our coding, we recognise
matter was secreted by a slightly modified set of the that the major lineages of euconodonts have a basal
cells which secreted lamellar crown tissue. This body composed of dentine, and regard the presence
modification could have been effected by differen- of dentine as plesiomorphic for euconodont elements.
tiation of the cells into a new type, by changing the
proximity of the secreting cells to the mineralising
(5) Conodont element growth
front, or by the timing of secretion relative to other
members of the cell population. In any event, it is Conodont elements grew in a manner directly
now clear that white matter is autapomorphic to comparable to odontodes, the building blocks of the
some conodonts although developmentally hom- vertebrate dermal skeleton (Donoghue, 1998). Each
ologous to enamel. element is reducible to a number of distinct
morphogenetic units, each composed from an in-
dividual crown and basal body (Donoghue, 1998).
(c) Basal tissue
These morphogenetic units were usually added
The basal tissue of various conodont taxa has been sequentially to the pre-existing, pre-formed struc-
compared to various dentines, types of bone and ture. Internal discontinuities can be identified as
calcified cartilage. However, no convincing evidence resulting from episodes of function alternating with
has been presented for the presence of cell spaces and episodes of growth (Donoghue, 1998 ; Donoghue &
Purnell, 1999 a). Conodont elements performed a biochemical preservation is negligible. Furthermore,
tooth function while continuing to grow by marginal the instability of collagen is such that it can only be
addition of successive odontode generations in the expected to survive biochemically for up to 1 million
same manner as the dentigerous jaw bones of years (see Aldridge & Purnell, 1996, and references
acanthodians (Ørvig, 1973) and tooth plates of therein). Further cause for caution is raised by the
lungfish (Kemp, 1977). More derived groups, such preparation of materials for histochemical analysis.
as the ozarkodinids, must have undergone phases of Picrosirius Red and comparable histochemical stains
dormancy as successive odontodes were added for collagen are intended for use on fully decalcified
circumferentially, facilitating element repair. This sections ; while Kemp & Nicoll (1995 a, b, 1996) used
exaptation allowed the evolution of complex dental decalcified sections of lungfish toothplates as their
mechanisms (Donoghue & Purnell, 1999 b). control, the conodont elements were surface-etched.
The position of the elements within the mouth\ An effect of the mineral salt in attracting the staining
pharynx has led us to code pharyngeal odontodes as molecule cannot be excluded in the interpretation of
present in conodonts, and conodont elements are their results.
considered to be homologous to the pharyngeal
odontodes of both thelodonts and sharks.
IV. PHYLOGENETIC ANALYSIS
(6) Histochemical studies
Many commentators now accept the evidence for a
Histochemical tests were first applied to conodont chordate affinity for conodonts, but the exact
elements by Fa/ hraeus & Fa/ hraeus-van Ree (1987, position of the Conodonta within the Chordata
1993), who used stains to investigate the insoluble remains debatable (Aldridge & Purnell, 1996). To
organic residues of decalcified specimens. The results date, only one attempt has been made to resolve the
from these early studies were interesting but equivo- systematic position of conodonts by formal cladistic
cal ; simple stained mounts of the organic films analysis (Janvier, 1996 b). This resulted in 79 equally
revealed the presence of structures resembling parsimonious trees, the strict consensus of which
collagen fibres, although the source of the tissues placed conodonts in an unresolved position, but
within the host elements was unknown. The ap- more derived than either group of acraniate
proach has been refined by Kemp & Nicoll chordates. Further runs of a modified matrix placed
(1995 a, b,1996) who extended the range of histo- conodonts as a sister-group of lampreys.
chemical stains used and applied them directly to the The phylogenetic analysis presented here con-
mineralised tissues. This technique produced positive siders representatives of a variety of lower craniate
results for collagen in lamellar crown tissue, leading taxa (Fig. 6), with the inclusion of Urochordata and
Kemp & Nicoll (1995a, b, 1996) to reject the Cephalochordata as outgroups. Our data matrix is a
hypothesis that this tissue is an homologue of enamel, derivative of one compiled by P. Janvier & P. L.
an entirely epithelial product lacking collagen. Forey (in preparation) in which all characters were
White matter failed to stain for collagen and the entered in binary fashion (either a structure is there
hypothesis that white matter is dermal bone was also or not), whereas we have opted to express some
rejected because bone always contains collagen. characters as multistate entries ; the reasons for this
Kemp & Nicoll (1995 a, b, 1996) concluded that are explained below.
conodont and vertebrate hard tissues are not
comparable. Their results also indicate the absence
(1) Taxon sampling
of keratin, and the presence of cartilage ; they also
recorded a positive stain result for DNA in a Middle The results of any phylogenetic analysis are subject
Ordovician conodont element. to the sampling of included taxa. Although this is a
Attempts to repeat these results with modern and particular problem with molecular data, where
fossil vertebrate hard tissues have failed (M.M. single species are taken to represent large groups
Smith personal communication 1996 in Donoghue, (e.g. Xenopus laevis is usually taken as the rep-
1998) and Kemp & Nicoll have also failed to resentative of the Amphibia), it is also a concern
demonstrate the effectiveness of these tests on with morphological data. Here, two of the terminal
undoubted fossil vertebrate material. Towe (1980) taxa cause us difficulty. For the Conodonta our
has demonstrated that, although tissues like collagen knowledge of soft tissue anatomy is limited to just
may be preserved physically with high fidelity, two species (Clydagnathus windsorensis and Promissum
Fig. 6. Diagrammatic sketches of the terminal taxa used in our analyses. (A) Eptatretus stoutii (Lockington), a modern
hagfish (600 mm). (B) Petromyzon marinus Linneaus, a modern lamprey (800 mm). (C) Jamoytius kerwoodi White, a
naked anaspid-like fish from the Lower Silurian of Scotland (130 mm) ; K. A. Freedman (in press) has suggested that
there is absence of evidence for a dorsal and caudal fin in J. kerwoodi. (D) Clydagnathus windsorensis (Globensky), a
euconodont from the Lower Carboniferous of Scotland (60 mm). (E) Pharyngolepis oblongus Kiaer, a scaled anaspid
from the Upper Silurian of Norway (150 mm). (F) Errivaspis wayensis (White), a heterostracan from the Lower Devo-
nian of England (150 mm). (G) Sacabambaspis janvieri Gagnier, an arandaspid from the Upper Ordovician of Bolivia
(after Gagnier 1989 ; 300 mm) – details of the tail are uncertain. (H) Furcacauda heintzae (Dineley & Loeffler), a deep-
bodied thelodont from the Lower Devonian of Canada (35 mm). (I) Hemicyclaspis murchisoni Egerton, an osteostracan
from the Lower Devonian of England (150 mm). (J) Loganellia scotica (Traquair), a thelodont from the Upper Silurian
of Scotland (120 mm). (K) Geraspis rara Pan & Chen, a galeaspid from the Middle Silurian of China – the tail is not
known in detail (150 mm). (L) Pituriaspis doylei Young, a pituriaspid from the Middle Devonian of Australia (head-
shield 45 mm in length).
pulchrum) and it is possible that these are not a similarity in preserved soft anatomy that suggests
representative of all conodonts (Panderodus unicostatus that they can be taken as representative of the
from the Lower Silurian of Wisconsin, U.S.A., also monophyletic group Conodonta. This group is
exhibits evidence of soft tissue remains although characterised by phosphatic conodontiform elements
these are too poorly preserved for adequate in- exhibiting a distinct crown composed of enamel and
terpretation ; see Smith et al., 1987, and Conway a basal body composed of dentine ; as far as is known,
Morris 1989 a). However, C. windsorensis and P. the body is laterally compressed with a short rostral
pulchrum scale a great taxonomic disparity but display head portion in which the elements are located.
There is also evidence of a pair of rostro-lateral eyes development of an oral sucker equipped with
with extrinsic eye musculature, a pair of otic capsules denticles and the presence of a pharynx which is a
located caudo-medial to the eyes and, possibly, at blind-ended diverticulum from the main tract of the
least four pairs of box-like gill pouches. The trunk is gut. Heterostracans are a large group of approxi-
elongate relative to its width, and constructed from mately 300 species of Lower Silurian to Upper
repeated symmetrical chevron-shaped muscle blocks Devonian fishes, restricted to North America,
which are convex in a rostral direction ; the caudal Europe and Siberia. They are characterised by a
portion of the animal is dominated by an asym- single branchial opening covered by a single bran-
metrically developed median fin supported by rays. chial plate associated with a large dorsal and ventral
The fin is more extensive on the dorsal margin where shield. The most comprehensive and recent review of
it is divided into two lobes. the entire heterostracan group has been published
Thelodonts are more problematic because of a by Blieck (1984). In the present review, we assume
lack of agreement over their monophyly (e.g. an internal phylogeny of heterostracans in which
Turner, 1991, versus Janvier, 1981, for discussion). forms such as traquairaspids represent the most
Thelodonts are mostly represented in the fossil record plesiomorphic condition. We have adjusted our
by isolated scales although some complete, articu- coding accordingly. For instance, we code the
lated fossils are known. Thelodonts were charac- presence of oak-leaf shaped tubercles as represen-
terised by Novitskaya & Turner (1998 : p. 533) as tative of heterostracans but we acknowledge that
having ‘ discrete dentinal scales with a base of such ornament is only found in our presumed
acellular bone-like tissue (aspidin) which is capable plesiomorphic representatives.
of growth, and the production of simple to complex The Astraspida, Arandaspida and Eriptychiida
anchoring devices ’. Most known articulated thelo- have traditionally been associated with the hetero-
donts are dorsoventrally flattened with asymmetrical stracans on the basis of comparative histology.
tails and with lateral flaps which lie above the gill However, since there is no clear consensus as to
openings. However, the Furcacaudiformes are lat- which type of histology is plesiomorphic or
erally compressed, possess a symmetrical tail and apomorphic and since discussion of histology is
lack lateral flaps ; they are nevertheless covered by a central to this review we consider each of these taxa
micromeric dermal skeleton of thelodontiform scales separately.
(Wilson & Caldwell, 1993, 1998). If thelodonts are a Astraspis has been recovered from the Upper
monophyletic group, as advocated by Turner Ordovician of North America, and exhibits an
(1991), it remains unclear which of the body shapes armour constructed from small polygonal plates
is plesiomorphic. A phylogenetic analysis carried out composed of aspidin, orthodentine and enameloid.
by Wilson & Caldwell (1998) placed dorso-ventrally There are separate branchial openings, each
flattened thelodont genera in a polytomy with associated with a small plate. Astraspis has been most
gnathostomes, while Furcacaudiformes were recently and most completely described by Sansom
regarded as the monophyletic sister-group to this et al. (1997).
combined group. If scale morphology is considered The Arandaspida include the earliest-known fully
as a more general character (e.g. if it is present in armoured craniates, Arandaspis from the Upper
galeaspids as suggested by Janvier, 1996b), then we Ordovician of Australia (Ritchie & Gilbert
should either include both types of animal in our Tomlinson, 1977), and Sacabambaspis from the Upper
cladistic analysis or simply select a single species\ Ordovician of Bolivia (Gagnier, 1993 a, b). The
genus with the proviso that this may not be arandaspids are characterised by large dorsal and
representative of all thelodonts. We have chosen to ventral shields, forward-looking eyes and a long
adopt the latter course (selecting Loganellia as our slanting series of rectangular branchial plates along
thelodont taxon) as we have not had the opportunity the flanks. The most comprehensive review of the
to study for ourselves representatives of the Furca- group was provided by Gagnier (1993 a, b).
caudiformes. Eriptychius is an enigmatic genus usually placed in
To justify the monophyly of other taxa included in its own higher taxon (Eriptychiida), primarily
this analysis we have used the following criteria, because its affinities are so poorly constrained. It is
although these features are by no means exhaustive. known exclusively from fragments of dermal armour
Hagfishes (Myxiniformes) show posterior displace- which include tubercles composed of dentine and a
ment of the gill-pouch series and the presence of bony base underlain by globular calcified cartilage
slime glands. Lampreys (Petromyzontiformes) show (Denison, 1967). It has proven difficult to determine
the precise systematic position of this taxon because deal of new information is now available both to
of the paucity of data available (see below). clarify ambiguities and to refute some aspects of
The Osteostraci is the best-known group of previous interpretations.
armoured jawless craniates ; approximately 300 The Pituriaspida is represented by two species
species have been found in the Silurian and from the Lower Devonian of Australia (Young,
Devonian of North America, Europe, Siberia and 1991) ; these are known only from natural moulds.
Central Asia. Osteostracans possess a large semi- Although poorly known, they appear to have a solid
circular dorsal head shield pierced by lateral and head shield perforated by large fenestrae immedi-
median sensory fields, which are areas of small ately behind the eyes. We include them here because
tesserae connected to the labyrinth by means of bony the casts show some internal anatomy implying the
canals. The most comprehensive revision of the presence of bone and pectoral fins.
group is that of Janvier (1985). Here, we accept The Gnathostomata has always been recognised
Janvier’s (1985) internal phylogeny for the group as a monophyletic group whose Recent members
which suggests that forms such as Ateleaspis and show many synapomorphies (Maisey, 1986), a few of
Aceraspis, which possess pectoral fins, are plesio- which may be listed as : primary upper (palato-
morphic, and that the tremataspids, which lack quadrate) and lower (meckelian cartilage) jaws
paired fins, are the most derived of osteostracans. which carry teeth erupting from a dental lamina ; a
The Galeaspida (approximately 70 species supporting hyoid arch ; segmented branchial arches
recovered from the Lower Silurian to the Upper lying internally to the blood and nerve supply and
Devonian of China and North Vietnam) is a group the gill lamellae ; separate endoskeletal pectoral and
of craniates that superficially resemble osteostracans. pelvic girdles and fin skeletons ; basals and radials
The dorsal surface of the head is covered by a single supporting dorsal and anal fins ; horizontal septum
broad and flattened head shield which is usually dividing epaxial from hypaxial musculature ; hori-
semicircular in outline, but can be extended into zontal semicircular canal ; myelinated nerve fibres ;
long cornuae and\or a long rostra in some taxa. genitial ducts (Wolffian and Mullerian ducts)
Galeaspid synapomorphies include a large, median developing from mesonephros ; renal portal system
anterodorsal opening, interpreted as a nasohypo- and subcardinal veins. Some fossil gnathostomes
physial opening, and a unique scalloped pattern of may lack some of these synapomorphies : for instance,
the main lateral-line canal upon the cephalic shield. it is doubtful if placoderms had a dental lamina since
This group was most comprehensively reviewed by their dentition is non-replaceable.
Janvier (1996 a). For some of our characters gnathostomes are
The Anaspida (approximately 25 species from the polymorphic and thus an unambiguous coding may
Silurian to Upper Devonian of Europe and North depend upon an assumed phylogeny for basal
America) is a group of small fusiform fishes charac- gnathostomes. There is little dispute that the
terised by postbranchial scales or rods and a series of Chondrichthyes are the most plesiomorphic extant
elaborate mid-dorsal scales extending the length of gnathostomes and therefore we have opted to code
the body instead of a dorsal fin. for the character states in this taxon. However, we
Jamoytius, Endeiolepis, Euphanerops and Legendrelepis recognise that some workers accept that Placodermi
have traditionally been classified with anaspids but is the plesiomorphic taxon (e.g. Goujet & Young,
lack both the modified postbranchial scales and mid- 1995) and it is therefore possible that some codings
dorsal scales typical of the Anaspida. Some authors should be different. For instance, character 78 in our
have considered one or more of these to be more data matrix refers to the condition of the dermal
closely related to lampreys than to anaspids (e.g. head covering, which is micromeric in chondri-
Forey & Gardiner, 1981, Arsenault & Janvier, chthyans and macromeric in most placoderms. For
1991). Only the better known of these genera, each character where there is a potential dispute
Jamoytius (Lower Silurian) and Euphanerops (Upper over its coding in gnathostomes, we have discussed
Devonian) have been considered for the purposes of and justified our scoring below.
the present phylogenetic analysis. Jamoytius has been
subject to very different interpretations (White,
(2) Character coding
1946 ; Ritchie, 1968 ; Forey & Gardiner, 1981), in
part because the preservation at the single known As mentioned above we have used presence\absence
locality is unusual. Recently, Jamoytius has been coding (Pleijel, 1995) in the majority of characters
restudied by K. A. Freedman (in press) and a great but we have also used some multistate characters.
There is currently some debate about the usefulness coding for those taxa which lack the structure.
and theoretical underpinning of each of these Computationally, the ‘ not applicable ’ is coded as a
strategies (Pimental & Riggins, 1987 ; Pleijel, 1995 ; question mark. But this leads to further compu-
Wilkinson, 1995 ; Hawkins, Hughes & Scotland, tational difficulties (Platnick, Griswold &
1997 ; Forey & Kitching, in press) which may lead to Coddington, 1991) such as the generation of multiple
the establishment of different relationships. equally parsimonious cladograms, many of which
Difficulty usually arises where some taxa do not contain spurious nodes.
show a structure that others show in one or more At present there is no ‘ right ’ or ‘ wrong ’ way to
conditions. For instance, character number 18 relates code for characters. We have chosen options that, to
to the presence or absence of olfactory organs as well us, make the most biological sense. But, as an
as to the state of such organs, which may be paired experiment we have recoded our data by translating
or unpaired. There are at least eight ways of coding all the multistate characters to presence\absence
such observations (see Forey & Kitching, in press, codings, and to contingent codings, and we have
for discussion). Extreme presence\absence coding of used Sankoff coding which weights certain trans-
this character would code this as two columns of formations (see below), to determine the effects on
data : 1. paired olfactory organs present or absent ; 2. the final topology.
unpaired olfactory organs present or absent. This
means that taxa having no olfactory organs need to
(3) Character matrix
be coded ‘ 0 ’ for both characters and the analysis
runs the risk of grouping taxa on the zero codings In this section, we list the characters, their possible
(plesiomorphic long branch attraction). Also, it is an states and, where necessary, provide discussion of
assumption for the purposes of cladistic analysis used how particular codes have been determined for
here that each column of data provides potentially certain taxa.
independent evidence of relationship ; that is, the
columns are not linked biologically or logically. But
(a) Brain, sensory and nervous systems
this is patently not so for the character under
discussion since the codings in one column pre- 1. Neural crest. Absent l 0, present l 1. Inferred
determine the codings in adjacent columns. The from the presence of known neural crest derivatives
advantages of presence\absence coding are that it (e.g. dermal skeleton, gill arches, pigment cells). The
maximises character congruence, reveals potential recording of neural crest in conodonts is based on the
homoplasy within the character and is the most presence of dentine and extrinsic eye musculature.
exacting test of homology. Smith, Graveson & Hall (1994) have proposed that
Linking the observations into a single column as a neural crest and epidermal placodes are derived
multistate character assumes that there is trans- from a common phylogenetic precursor, while
formation between one observation and another and Northcutt (1996 b) has subsequently provided pre-
denies the possibility that the evolution of unpaired liminary evidence to suggest that neural crest and
olfactory organs is independent of the origin of epidermal neurogenic placodes have a common
paired olfactory organs. In this particular example, developmental origin. It is possible, therefore, that
it is possible that there may be no argument, since it the presence of placode-derivatives may also be
is known through ontogenetic studies of the lamprey taken for the presence of neural crest. As such, this
that the seemingly unpaired olfactory organ is a character partially overlaps with character 2 (brain)
transformation of the paired condition. But in linking which relies on the presence of paired sensory organs
the nature of the dermal head covering (character for coding in many fossil taxa. Baker & Bronner-
78 : absent, micromeric, macromeric) there is no Fraser (1997) have recently argued for the presence
such ontogenetic evidence of transformation either of neural crest precursors or putative homologues in
way and there may be a case for suggesting that acraniate chordates based on : the presence of
micromeric and macromeric are two independently AmphiDll- (the amphioxus homologue of Distal-less)
acquired expressions of a dermal skeleton. expressing migratory epidermis in amphioxus (Hol-
An intermediate way of coding such characters is land et al., 1996) which exhibits parallels to neural
the contingent coding method where one column of crest in vertebrates ; the presence of dorsal sensory
data denotes the presence or absence of a structure neurons in the central nervous system of amphioxus
and the second column expresses the various con- which may share a common origin with neural-crest-
ditions of that character with a ‘ not applicable ’ related sensory neurons in the vertebrate CNS (e.g.
mesencephalic nucleus of the trigeminal nerve ; interpreted by Aldridge et al., 1993) appears to stop
Narayanan & Narayanan, 1978) ; the presence of short of the anterior end of the body (cf. Branchio-
putatively neurogenic migratory cells in tunicates stoma) and this may also indicate the existence of a
(Muske, 1993 ; Bollner, Beesley & Thorndyke, 1993, brain. It may also be pointed out that sensory
1997 ; Bollner et al., 1995 ; Mackie, 1995) ; the capsules etc. are also associated with a neurocranium
presence of mechanoreceptors in ascidian tadpoles but in many of our chosen taxa there is no direct
(Torrence & Cloney, 1982 ; Crowther & Whittaker, evidence of this.
1994), thaliaceans (Bone & Ryan, 1978) and 3. Olfactory peduncles. Absent l 0, present l
amphioxus (Bone & Best, 1978 ; Baatrup, 1981). 1. These are assumed to be present in heterostracans,
Baker & Bronner-Fraser (1997) even identify poss- on the basis of the ridge leading from the pineal
ible neural crest and placode precursors in non- recess to the nasal sac impressions, which corresponds
chordate deuterostomes. However, neither pre- to the position of the olfactory peduncles observed in
cursors nor putative homologues can substitute for galeaspids.
the presence of neural crest and\or epidermal 4. Pineal organ. Absent l 0, present and covered
placodes, and so we score this character as absent for l 1, present and uncovered l 2. In fossil forms, this
cephalochordates and tunicates. is usually indicated by a foramen or a depression on
2. Brain : absent l 0, present l 1. A brain here the inner surface of the dermal skeleton. In Astraspis,
is interpreted as an anterior enlargement of the Sansom et al. (1997) have described a slightly raised
dorsal nerve chord which is morphologically differen- area of tesserae in the presumed position of the
tiated and associated with one or more complex pineal organ and this has been interpreted as
sense organs (nasal sacs, paired and\or median eyes, evidence of the presence of a pineal organ. A paired
labyrinth) and cranial nerves with dorsal somato- pineal organ has been reported in the arandaspids
motor roots and ventral visceromotor and viscero- Arandaspis (Ritchie & Gilbert-Tomlinson, 1977) and
sensory roots. Garcia-Ferna' ndez & Holland (1994 ; Sacabambaspis (Gagnier, 1993 a) and thus is coded
see also Holland & Garcia-Ferna' ndez, 1996) have present here. However, it should be pointed out that
demonstrated that a portion of the cerebral vesicle these paired openings lie well behind the eyes which
and dorsal nerve chord of Branchiostoma is hom- is an unusual position for such structures, and there
ologous to the fore- and hindbrain of vertebrates. remains the possibility that these are paired endo-
More recently, Wada et al. (1998) have identified a lymphatic openings. In hagfishes, there is no pineal
tripartite organisation to the neural tube of an organ even though a habenular swelling and the
ascidian based on comparison between expression habenular ganglion are present (Wicht, 1996). As a
patterns of Hroth, HrHox1 and HrPax-258 and their result, the presence of a habenular swelling in the
homologues in vertebrates (Otx, Hox and Pax-2, -5, cranial endocast of pituriaspids (Fig. 6 in Young,
-8, respectively). However, these are ‘ field ’ homo- 1991) does not constitute sufficient evidence for the
logues rather than primary homologues and in presence of a pineal organ. On structural evidence,
neither of the above cases do the fields give rise to Lacalli, Holland & West (1994) and Lacalli (1996)
any clearly differentiated sensory organs [although identified homology between the frontal eye and
Wada et al. (1998) suggest that later expression of lamellar body of Branchiostoma floridae and the paired
HrPax-258 is associated with epidermal thickenings eyes and pineal organ (respectively) of vertebrates.
(possible neurogenic placodes) which give rise to Again, this only demonstrates that the structures in
what the authors suggest to be a homologue of the amphioxus and vertebrates are field homologues (cf.
vertebrate ear, and Sharman, Shimeld & Holland Wicht, 1996). The expression of a Pax-6 homologue
(1999) describe the presence of placodes or placode- (AmphiPax-6) in precursor cells of the frontal eye and
homologues in Branchiostoma]. There are no extant lamellar body in amphioxus (Glardon et al., 1998)
craniates which show sensory organs and no brain. demonstrates only that a gene known to have a role
Thus, it is assumed that evidence of any of the in the development of photoreceptors in a broad
sensory organs signifies the presence of a brain. range of animals (Callaerts, Halder & Gehring,
Direct evidence of the brain in fossil forms is seen as 1997) is also implicated in photoreceptor devel-
endocasts of the brain cavity and is known in opment in amphioxus. Nevertheless, because of the
osteostracans, galeaspids, and pituriaspids (Young, correlation between these two reports, we record ?
1991). In conodonts, the evidence for the brain is for this character in cephalochordates.
indirect ; in addition to the presence of paired 5. Pituitary divided to adenohypophysis and
sensory organs, the notochord of conodonts (as neurohypophysis. Absent l 0, present l 1.
6. Adenohypophysis. Absent l 0, simple l 1, organ (see Fig. 25 in Goodrich, 1909). This is taken
segmented and compartmentalised l 2. to indicate a paired ontogenetic origin.
7. Optic tectum. Absent l 0, present l 1. However, it needs to be pointed out that
Ronan & Northcutt (1998) have recently described Gorbmann and Tamarin (1985) demonstrated that
an optic tectum in the brain of the Pacific hagfish in Petromyzon marinus the olfactory placode giving rise
Eptatretus stouti. to the olfactory epithelium is unpaired from the
8. Cerebellum. Absent l 0, present l 1. The beginning. It is therefore possible that lampreys are
presence of a cerebellum in lampreys is usually polymorphic for this character. We have opted to
recorded. However, recent work (C. Weigle, per- code lampreys as having paired olfactory organs on
sonal communication ; Weigle & Northcutt, 1998) the basis that other cranial sensory structures are
suggests that the structure in lampreys cannot be paired and hence, the plesiomorphic condition for
considered as a cerebellum since it lacks the auricles olfactory capsules is likely to be paired.
which contain the emenentia granularis of 19. Extrinsic eye musculature. Absent l 0, pres-
gnathostomes. The coding in fossil taxa showing ent l 1. This is inferred in fossils from the presence
evidence of brain contours is based on the assumption of myodomes and oculomotor nerve canals ; in
that the prominent paired swellings of the brain conodonts it is recorded on the basis of muscle fibres
cavity seen in heterostracans, osteostracans and preserved in intimate association with the organic
galeaspids (and possibly pituriaspids) are for the films interpreted by Aldridge & Theron (1993) as
cerebellum and not the optic lobes (see discussion in eye capsules (e.g. Gabbott et al., 1995).
Janvier, 1985). 20. Presence\absence and number of semicircular
9. Pretrematic branches in branchial nerves. canals in labyrinth. None l 0, one l 1, two l 2,
Absent l 0, present l 1. three l 3. In heterostracans, there are clearly at
10. Flattened spinal chord. Absent l 0, present least two semicircular canals which appear to
l 1. correspond to the vertical semicircular canals of
11. Ventral and dorsal spinal nerve roots united. gnathostomes, but since nothing is preserved other
Absent l 0, present l 1. Here, the hagfishes and than shield impressions it remains a possibility that
gnathostomes are coded the same ; that is, they both an horizontal semicircular canal was also present.
have dorsal and ventral roots united. However, it However, we note that the angle between the two
needs to be pointed out that in gnathostomes the impressions for the vertical semicircular canals
union lies immediately outside the dorsal nerve cord, appears to be marked by the impression for a gill
whereas in hagfishes the union lies deep within the pouch (e.g. see Fig. 4.5A1 in Janvier 1996 a), which
body musculature away from the nerve cord. would preclude the existence of a horizontal canal.
12. Mauthner fibres in central nervous system. 21. Vertical semicircular canals forming loops,
Absent l 0, present l 1. well separate from the vestibular division of the
13. Synaptic ribbons in retinal receptors. Absent labyrinth. Absent l 0, present l 1.
l 0, present l 1. 22. Externally open endolymphatic ducts. Absent
14. Number of nasal openings. None l 0, paired l 0, present l 1. For galeaspids, this is clearly
l 1, single median l 2. The coding for hetero- observed only in the genus Xiushuiaspis (Wang,
stracans depends upon reconstruction of the grooves 1991), but since this form is one of the most
that mark the undersurface of the snout as being plesiomorphic galeaspids (Janvier, 1996 a), the
evidence of paired prenasal sinuses. character is assumed general for the entire group.
15. Nasohypophyseal opening serving respiration We have already commented upon the possibility of
(nasopharyngeal duct). Absent l0, present l 1. endolymphatic ducts in arandaspids (see discussion
16. Single nasohypophyseal opening. Absent l of character 4) but code here following the des-
0, present l 1. criptions of the original authors.
17. Position of nasohypophyseal opening. None 23. Sensory-line system with neuromasts. Absent
l 0, terminal l 1, dorsal l 2. l 0, present l 1. The presence of neuromasts is
18. Olfactory organ. Absent l 0, paired l 1, assumed in fossils when sensory-line canals or long
unpaired l 2. Where an olfactory organ is present continuous grooves are observed. Simple short or
the presence of a septum between the two halves of irregular grooves are inconclusive, since such grooves
the olfactory organ denotes a paired structure. Thus, devoid of neuromasts exist in hagfishes. That the
in Petromyzon fluviatilis there is a prominent median grooves in hagfishes represent homologues of the
septum dividing an outwardly median olfactory lateral line system of other craniates is supported by
their development (Wicht & Northcutt, 1995), The evidence for pouch-like gills in Eriptychius is
innervation (Braun & Northcutt, 1997, 1998) and based on Ørvig’s (1958 : pl. 2, figs 4, 5) observation
central projections (Kishida et al., 1987). However, of a branchial plate (incorrectly identified as a
the phylogenetic polarity of this condition (i.e. branchio-cornual plate) with pore-like openings.
primitive versus degenerate) is a moot point 28. Gills alternate l 0, symmetrical l 1. In hag-
(Fernholm, 1985 ; Wicht & Northcutt, 1995 ; Braun, fishes, the gill pouches of either side alternate with a
1996 ; Braun & Northcutt, 1997) ; as the sensory hair corresponding alternation of the blood supply from
cells are unoriented and lack cupulae, they are not the ventral aorta. This may be an adaptation of an
true neuromasts. eel-shaped body. However, a similar alternation of
24. Electroreceptive cells. Absent l 0, present l 1. the gills occurs in Branchiostoma and the condition
25. Sensory-line grooves or canals. Absent l 0, may, therefore, be plesiomorphic for craniates ; a
present on head only l 1, present on head plus body symmetrical distribution of the gill pouches on either
l 2. The coding for Astraspida is based on the side of the midline would then be the derived
recent description by Sansom et al. (1997), who condition.
recorded the presence of paired short sensory grooves 29. Elongate branchial series. More than 10 gill
confined to the pineal region of the head shield. It pouches\slits l 0, fewer than 10 l 1. In Branchio-
has traditionally been observed that the lateral lines stoma, there are numerous gill slits which extend well
of lampreys are restricted to the head. However, down the body and this is accepted here as the
Johnston (1905) documented the presence of lateral plesiomorphic craniate condition. The few but more
lines continuing from head to trunk in Petromyzon, complexly developed gill pouches of most craniates
and R. G. Northcutt (personal communication, are regarded as derived. Some galeaspids, however,
September 1999) records the presence of trunk have up to 40 branchial fossae. This is regarded here
lateral lines in all genera of lampreys that he has as a derived state within this group, since all of the
studied. most generalized galeaspids have fewer than 10 gill
26. Sensory-line. Absent l 0, enclosed in grooves openings or branchial fossae.
l 1, enclosed in canals l 2. In some terminal taxa 30. Gill openings lateral and arranged in slanting
used here sensory canals may be in grooves or canals row. Absent l 0, present l 1.
(e.g. osteostracans and galeaspids). Here, we record 31. Position of gill openings. Gills opening lat-
the taxa as showing canals if any members have this erally l 0, ventrally l 1.
condition. Linking the condition of the lateral line 32. Opercular flaps associated with gill openings.
into a multistate character is justified by the Absent l 0, present l 1. This refers to a small
heterochronic relationship between the two states as opercular flap, covered with minute dermal plates,
demonstrated for Ambystoma mexicanum by Northcutt, which partly covers the gill openings (e.g. in
Catania & Criley (1994). osteostracans, thelodonts). These are presumed
absent when the gill openings are small, rounded,
and surrounded by solid dermal elements (e.g. in
(b) Mouth and branchial system
arandaspids, astraspids, anaspids).
27. Pouch-shaped gills. Absent l 0, present l 1. 33. Endodermal gill lamellae. Absent l 0, pres-
The presence of pouch-like gills signifies : the shape ent l 1.
of the openings as being pore-like ; that the gill 34. Gill lamellae with filaments. Absent l 0,
lamellae are inclined towards the centre of the gill present l 1. This is inferred in fossils when typical
chamber in frontal section ; and the skeleton is gill lamellae impressions are observed.
external to the lamellae. The pouch-like gill may be 35. Mouth terminal l 0 or ventral l 1. Our
round (lampreys and hagfishes) or transversely coding for Jamoytius is taken from K. A. Freedman
elongated (some thelodonts, galeaspids and osteo- (in press).
stracans). There is no known instance where the 36. Velum. Absent l 0, present l 1.
opening is pore-like but the internal anatomy is like
the slit-like gills of gnathostomes (i.e. lamellae
inclined towards the opening and the skeleton medial
(c) Circulatory system
to the lamellae). Thus, the presence of pore-like
openings even in the absence of other knowledge (as 37. Relative position of atrium and ventricle of
in arandaspids and anaspids) is assumed to indicate heart. Well separated l 0, close to each other l 1.
a gill structure like that of lampreys and hagfishes. In fossils, the structure of the heart is only known in
osteostracans from the morphology of the pericardial is a paired lateral fin fold developed at the base of the
cavity (Janvier, 1985). tail in some taxa, such as Hemicyclaspis (Heintz,
38. Closed pericardium. Absent l 0, present l 1. 1967).
39. Open blood system. Absent l 0, present l 1. 46. Unpaired fin ray supports closely set. Absent
40. Paired dorsal aortae. Absent l 0, present l l 0, present l 1. In several fossil taxa (Hetero-
1. This is inferred in some fossils from the shape of the straci, Anaspida, Osteostracans and Thelodonti),
aortic groove. The condition within gnathostomes the ray supports are not observed directly, but their
does vary but here the presumed plesiomorphic position is inferred from the zonation of the
paired condition as represented in chondrichthyans squamation.
is coded. 47. Paired lateral fin folds. Absent l 0, present
41. Large lateral head vein. Absent l 0, present l 1. This refers to any lateral or ventrolateral fin-
l 1. like fold, irrespective of whether they contain radials
42. True lymphocytes. Absent l 0, present l 1. and musculature. Hagfishes are coded here as
43. Subaponeurotic vascular plexus. Absent l 0, lacking such folds but it needs to be mentioned that
present l 1. This character is inferred to be present Jarvik (1980) has recorded the presence of lateral fin
in heterostracans, on the basis of the numerous folds in Neomyxine plicata. These structures are not
vascular grooves in Torpedaspis (Broad & Dineley, present in any other hagfish and N. plicata is not the
1973), although most other heterostracans have a most plesiomorphic hagfish (Fernholm, 1998), there-
smooth internal surface of the exoskeleton and show fore we regard these fin folds as non-homologous
no clear evidence of such a network. The subdermal with those developed in anapsids.
vascular canals of Eriptychius (Denison, 1967) are 48. Constricted pectorals. Absent l 0, present l
assumed here to belong to this network. Sansom et al. 1. This character refers to the gnathostome-like
(1997) documented the presence of a canalised postbranchial pectorals which have a constricted
network deep within the dermal armour of Astraspis. base with an endoskeleton (cartilage or bone) and
We interpret this as a subaponeurotic vascular associated musculature. No fossil jawless vertebrate
plexus based on its similarity to the condition met shows a clearly recognisable endoskeleton (although
with in Eriptychius. some indication may be present in the Upper
Devonian Escuminaspis – Belles-Isles, 1989) but the
occurrence of musculature and possibly an endo-
(d) Fins and fin-folds
skeleton is inferred in osteostracans and pituriaspids,
44. Dorsal fin. Separate dorsal fin absent l 0, based on the scars and foramina inferred to be for
present l 1. In cephalochordates and vertebrate nerve\blood vessels left within the pectoral fenestra
embryos, there is a continuous fin fold which extends of some species. To score osteostracans for this
around the dorsal and ventral half of the body and character requires an internal phylogeny of osteo-
primitively continues ventrally in front of the cloaca stracans which proposes that Ateleaspis and Hemi-
as the preanal fin fold. This initial fold is not cyclaspis, which have well-developed pectoral fins,
supported by any fin rays. In most adult vertebrates, are plesiomorphic osteostracans (see Janvier, 1985).
this fold becomes supported by radials and is These deductions from the phylogenetic tree pro-
differentiated into separate dorsal, caudal and duced by Janvier (1985) echo earlier suggestions of
sometimes anal fins : such differentiation is here Stensio$ (1927) that tremataspids had lost paired fins,
regarded as derived. In scaled anaspids such as and those of Heintz (1939) who considered that
Birkenia and Pterygolepis etc. the dorsal margin of the Ateleaspis and Aceraspis are the most primitive
trunk is covered with modified and enlarged scales osteostracans.
which may be modifications of a dorsal fin, but since 49. Tail shape. Isocercal l 0, hypocercal l 1,
there are no obvious signs of fin rays this group is epicercal l 2. It is almost certain that the isocercal
coded as lacking a dorsal fin. Our coding for tail is the plesiomorphic condition and evidence for
Jamoytius as having no dorsal fin is based on the this may be seen in embryos of tail-bearing
evidence provided by K. A. Freedman (in press). vertebrates (ontogenetic criterion of character po-
45. Anal fin separate. Absent l 0, present l 1. larity), and in cephalochordates (outgroup
The primitive condition is taken to be a continuous criterion). Both other conditions are here regarded
fin fold, with differentiation of a separate anal fin as equally derived and transformation between any
being the derived condition. Here, there is assumed of the three is regarded as equally likely. Although
to be no anal fin in osteostracans even though there the position of the notochord is unknown in
heterostracans, we opt here for an isocercal tail for lampreys, the neurocranium extends back to enclose
this group following descriptions by Denison (1971), the labyrinth within the otic capsule. In
Soehn & Wilson (1990) and Wilson & Soehn (1990). gnathostomes, osteostracans, galeaspids and prob-
For Jamoytius, we have followed K. A. Freedman (in ably pituriaspids, it extends more posteriorly to
press) who stated that the tail shape cannot be enclose both the glossopharyngeal and vagus nerves.
reliably inferred from currently available specimens. 61. Annular cartilage. Absent l 0, present l 1.
The tail of conodonts is scored as hypocercal for This is best developed in the modern lamprey where
reasons discussed above. it supports the sucker. A comparable structure can
50. Preanal median fold. Absent l 0, present l be seen in some fossil jawless vertebrates but there is
1. This character is extended to include the presence no direct evidence that a sucker was similarly
of preanal scutes or crest in fossils possessing a developed, nor that the preserved structure was
mineralised exoskeleton. necessarily cartilaginous ; presence is scored for all
taxa with annular structures surrounding the mouth.
62. Trunk dermal skeleton. Absent l 0, present
(e) Skeletal
l 1.
51. Ability to synthesise creatine phosphatase. Ab- 63. Perichondral bone. Absent l 0, present l 1.
sent l 0, present l 1. This function is assumed to Although pectoral fins are not always present in
be present in all fossils having a skeleton made up of association with perichondral bone (e.g. galeaspids),
calcium phosphate. the converse is true. Thus, we have been able to score
52. Visceral arches fused to the neurocranium. perichondral bone as present in pituriaspids even
Absent l 0, present l 1. though there is no record of histology. Further
53. Horny teeth. Absent l 0, present l 1. evidence in support of this inference is supplied by
Ritchie (1960) noted denticles within the mouth preservation of brain endocasts in pituriapids ; such
region of Jamoytius but did not describe their features could not be preserved in the absence of an
composition. Re-examination of the material has originally mineralised brain case.
revealed that the structures in question are associated 64. Calcified cartilage. Absent l 0, present l 1.
with the naso-hypophysial opening and not the 65. Calcified dermal skeleton. Absent l 0, pres-
mouth. Furthermore, identification of these struc- ent l 1. The coding for conodonts is based on the
tures as denticles is unlikely. presence of calcified hard tissues forming the
54. Trematic rings. Absent l 0, present l 1. elements. There is some dispute in the literature
The series of ring-like impressions observed in concerning the presence of hard tissues in Jamoytius
Jamoytius and Euphanerops are tentatively homo- (Ritchie, 1984 versus Forey & Gardiner, 1981).
logised with the trematic rings. Taphonomic studies by K. A. Freedman (in press)
55. Arcualia. Absent l 0, present l 1. The pres- indicate that Jamoytius possessed mineralised scales,
ence of arcualia (neural arches and spines) is inferred so we code the calcified dermal skeleton as present.
in osteostracans and heterostracans, which show 66. Spongy aspidin. Absent l 0, present l 1.
impressions of these structures on the undersurface of Although originally coined for a very specific tissue
the head shields (Janvier & Blieck, 1979), and in type shown by pteraspids and psammosteids (Gross,
galeaspids, which possess an elongated endoskeletal 1930), the term aspidin now represents an unnatural
occipital region which encloses the vagus nerve and assemblage of histological types that are grouped
hence is inferred to include arcual elements. only by the presence of profuse fine fibre spaces.
56. Cartilaginous copula associated with tongue While it is certain that not all these tissue types are
protractor and retractor muscles. Absent l 0, pres- homologous, further classification requires a sys-
ent l 1. tematic study. We suggest that in future, the term
57. Chondroitin 6-sulphate in cartilage. Absent ‘ aspidin ’ should be restricted to a tissue type that
l 0, present l 1. incorporated aspidones. For this study we have
58. Braincase with lateral walls. Absent l 0, chosen to distinguish two types of ‘ aspidin ’ which
present l 1. may not be mutually exclusive. Spongy aspidin
59. Neurocranium entirely closed dorsally and refers to tissue types which are alamellar and
covering the brain. Absent l 0, present l 1. incorporate aspidones.
60. Occiput enclosing vagus and glosso- 67. Lamellar aspidin. Absent l 0, present l 1.
pharyngeal. Enclosure of cranial nerves IX and X, See notes for character 66 above. Lamellar aspidin
absent l 0, present l 1. In both hagfishes and refers to tissue types that have traditionally been
interpreted as aspidin, but which lack aspidones and arandaspids this is known to be three-layered in at
are entirely lamellar. least Sacabambaspis (Gagnier, 1993 a ; Young, 1997).
68. Cellular bone. Absent l 0, present l 1. In 72. Cancellar layer in exoskeleton, with honeycomb-
conodonts, we record cellular bone as absent for shaped cavities. Absent l 0, present l 1.
reasons given in the text above. 73. Composition of the scales\denticles\teeth.
69. Dentine absent l 0, mesodentine l 1, ortho- Absent l 0, made up by a single odontode l 1,
dentine l 2. Here, we classify dentine according to made up by several odontodes l 2. Gnathostomes
the scheme of Ørvig (1967) where mesodentine refers are polymorphic for this character. Here, we accept
to a dentine in which the odontoblasts are enclosed that the polyodontic scales of Palaeozoic sharks
in the mineralised matrix and cell processes are represent the plesiomorphic state for gnathostomes.
unpolarised. Orthodentine is typified by no cell 74. Scale shape. Scale absent l 0, diamond-
inclusion and parallel to sub-parallel, polarised cell shaped l 1, rod-shaped l 2. New observations of
processes that are enclosed in the mineralised matrix. Jamoytius suggest that there are rod-shaped scales
The grouping of mesodentine and orthodentine into over the dorsal flank (K. A. Freedman, in press). The
a multistate character is justified by the evolutionary primitive condition for the isolated scales in osteo-
relationship between these tissues, first proposed by stracans is typified by Ateleaspis in which they are
Ørvig (1967) and subsequently followed by Smith & diamond-shaped.
Hall (1990, 1993). While dentine is present in 75. Oak-leaf-shaped tubercles. Absent l 0, pres-
conodonts (see above), it is neither mesodentine nor ent l 1. These are highly characteristic styles of
orthodentine plesiomorphically ; we chose not to ornament which consist of elongated tubercles with
introduce a further multistate for ‘ atubular dentine ’ scalloped edges.
because no other group in our analysis is known to 76. Oral plates. Absent l 0, present l 1. These
possess atubular dentine plesiomorphically and so are found as either small parallel plates or larger
there is no potential homology. We have therefore plates within the lower lip of some fossil forms. It is
scored conodonts as ‘ ? ’ for this character. thought that they were linked by soft tissue, so that
70. Enamel\oid absent l 0, (monotypic) enamel they could be extended as a scoop-shaped fan.
l 1, enameloid (bitypic enamel) l 2. Here, we 77. Denticles in pharynx. Absent l 0, present l
follow Smith (1989) in our distinction of enamel 1. The coding for conodonts makes the assumption
(monotypic enamel) and enameloid (bitypic en- that the conodont elements are homologous with
amel). Grouping of the two tissues into a multistate pharyngeal denticles in thelodonts and sharks.
character is justified by their heterochronic re- However, it should be noted that the position of
lationship, as elucidated by Shellis & Miles (1974) conodont elements relative to the alimentary canal
and subsequently followed by Smith (1989, 1992, remains unresolved and our contention that the
1995). Although the superficial glassy layer of feeding apparatus occupied an oro\pharyngeal
thyestidian dermal armour has traditionally been position represents equivocation over whether it was
interpreted as enameloid, the clear absence of oral, pharyngeal or oropharyngeal in position.
evidence for the presence of a basal lamina, and thus 78. Dermal head covering in adult state. Absent
an enameloid-dentine junction, precludes unequivo- l 0, small micromeric l 1, large (macromeric)
cal interpretation of this tissue layer as such. Indeed, dermal plates or a shield l 2. The coding for
it is equally likely that the superficial glassy layer in gnathostomes is debatable since they are polymor-
thyestidian osteostracans is a hypermineralised den- phic for this character. The plesiomorphic
tine in which no tubules are present, resulting from gnathostome condition clearly depends on an
retraction of the cell processes prior to mineralisation internal phylogeny and, more specifically the
or from secondary backfilling of tubules. Further- relationships between placoderms (mostly macro-
more, although this condition has traditionally been meric) and chondrichthyans (micromeric) as well as
extrapolated to typify the dermal histology of all an internal phylogeny of placoderms. Recent litera-
osteostracans, thyestidian histology is atypical and ture (reviewed in Janvier, 1996 a) has settled on two
derived. competing theories of relationships of placoderms
71. Three-layered exoskeleton consisting of a and other gnathostomes. Either they are the sister-
basal lamella, middle spongy (or cancellar) layer group of all other gnathostomes or they are the
and a superficial (often ornamented) layer. Absent sister-group of chondrichthyans. These theories lead
l 0, present l 1. Although it is not possible to to ambiguity in polarising this character. Con-
establish the exact nature of the dermal skeleton of sideration of an internal phylogeny of placoderms
similarly leads to uncertainty. More traditional system where the walls of the blood vessels are
theories such as those of Denison (1978) suggest that thickened, equipped with valves, and function as
micromeric forms such as stensioellids are the most accessory hearts.
plesiomorphic placoderms, in which case macromery 88. Hyperosmoregulation. Absent l 0, present
may have developed independently within the group l 1.
and, irrespective of the relationships as a group, 89. High proportion of serine and theronine
micromery would be plesiomorphic for gnatho- collagen. Absent l 0, present l 1.
stomes. However, more modern theories include the 90. Presence of lactate dehydrogenase 5. Absent
suggestions that stensioellids cannot be shown to be l 0, present l 1.
placoderms (Janvier, 1996 a) and that other micro- 91. Pituitary control of melanophores. Absent l
meric placoderms (radotinids) are not the most 0, present l 1.
plesiomorphic (Goujet & Young, 1995). Thus, we 92. Pituitary control of gametogenesis. Absent l
are left with doubt. Because of this we disregard the 0, present l 1.
macromery of placoderms and accept that, since the 93. High metabolic rate. Absent l 0, present l
relationships of chondrichthyans to other living 1. See Hardisty (1979, 1982) for discussion.
gnathostomes is rarely in dispute, we should code the 94. Ion transport in gills. Absent l 0, present l 1.
plesiomorphic gnathostome condition as micro-
meric.
(g) Miscellaneous
79. Large unpaired ventral and dorsal dermal
plates on head. Absent l 0, present l1. 95. Typhlosole in intestine. Absent l 0, present
80. Massive endoskeletal head shield covering the l 1.
gills dorsally. Absent l 0, present l 1. This refers 96. Spleen. Absent l 0, present l 1.
to the endoskeletal shield of galeaspids, osteostracans 97. Collecting tubules in kidneys. Absent l 0,
and pituriaspids (inferred from the complex internal present l 1.
casts). 98. Condensed and discrete pancreas. Absent l
81. Sclerotic ossicles. Absent l 0, present l 1. 0, present l 1.
These dermal elements which surround the eye are 99. A islet cells in the endocrine pancreas. Absent
present in a wide variety of gnathostomes but only l 0, present l 1.
rarely seen in jawless vertebrates. Their presence in 100. Male gametes shed directly through the
arandaspids as recorded by Gagnier (1993 a) is coelom. Absent l 0, presentl 1.
accepted here. 101. Forward migration of postotic myomeres.
82. Ossified endoskeletal sclera encapsulating the Absent l 0, present l 1.
eye. Absent l 0, present l 1. Gagnier (1993 a) 102. Sexual dimorphism. Absent l 0, present l 1.
describes a calcification around the eyeball of 103. Larval phase. Absent l 0, present l 1.
arandaspids. Although there is some doubt about its Gnathostomes are assumed to have no larval phase
nature, this interpretation is accepted here. as the plesiomorphic condition.
( f ) Physiological
(4) Results
83. Blood volume. More than 10 % of body
volume l 0, less than 10 % of body volume l 1. ‘ to some palaeontologists, fossils preserve information so
84. Haemoglobins with low O affinity and different from that derivable from living organisms that the
#
significant Bohr effect. Absent l 0, present l 1. two cannot be classified in the same way. Surely the real
85. Nervous regulation of heart. Absent l 0, question is ‘‘ How much of this additional information is
present l 1. essential to investigate a particular idea ?’’ ’ (Donovan &
86. Heart response to catecholamines. Absent l Paul, 1998).
0, present l 1.
87. High blood pressure. Absent l 0, present l Parsimony analysis of the data presented in Table 1
1. Lampreys and gnathostomes show a considerably resulted in three equally parsimonious trees, the
higher basal metabolic rate and higher blood strict consensus of which is shown in Fig. 7 A. The
pressures than hagfishes (Hardisty, 1979, 1982). dataset was analysed using both PAUP 3.1.1
Indeed, the blood pressure of the hagfish is so low (Swofford, 1993) and Hennig86 (Farris, 1988),
that there are several regions of the circulatory which both produced the same topological results.
Tunicata
A Tunicata
B Cephalochordata
Myxinoidea
Cephalochordata Petromyzontida
Conodonta
Myxinoidea Astraspis
Pituriaspida
Galeaspida
10 Petromyzontida Osteostraci
100
Jawed vertebrates
Conodonta Eriptychius
6
99 Loganellia
Astraspis
Euphanerops
2 Jamoytius
53 Heterostraci
Anaspida
Arandaspida
Arandaspida Heterostraci
2 Anaspida
58 C Tunicata
Cephalochordata
Jamoytius Myxinoidea
Petromyzontida
Euphanerops Pituriaspida
Galeaspida
Loganellia Osteostraci
Jawed vertebrates
Eriptychius Eriptychius
Loganellia
Jawed vertebrates Euphanerops
Jamoytius
Osteostraci Anaspida
Arandaspida
Pituriaspida Heterostraci
Astraspis
Galeaspida Conodonta
Fig. 7. (A) Strict consensus of the three equally most parsimonious trees ; 180 steps, CIe 0.654, RI 0.699, RC 0.458 ;
annotated nodes signify the Bremer Support (Clade Decay) value (top) and bootstrap value (bottom). Values are
only given for nodes with greater than one-step Bremer Support or greater than 50 % bootstrap support. (B) Strict
consensus of the 55 trees which are one step longer than the most parsimonious ; CIe 0.650, RI 0.694, RC 0.453. (C)
Strict consensus of the 459 trees which are two steps longer than the most parsimonious ; CIe 0.646, RI 0.689, RC
0.447.
The PAUP results are presented here (Fig. 7 A). (see Fig. 14) and two (under ACCTRAN opti-
Multistate characters were left unordered and the misation) or four nodes (under the DELTRAN
tree-building routine used was ie* implicit enu- optimisation) are supported entirely by homoplasy.
meration (Hennig86) or branch and bound (PAUP). Furthermore, two frequently used measures of the
Tunicates and cephalochordates were used as a robustness of the tree – the bootstrap and Bremer
paraphyletic outgroup but it should be stressed that support (or clade decay) methods – imply weak
the topological results for the interrelationships of support for this topology (Fig. 7 A and Appendix).
craniates and conodonts are the same if either The phylogenetic positions of two taxa, Eriptychius
tunicates or cephalochordates are used alone as the and Pituriaspida, invite particular comment. Both
outgroup. The same results were also obtained if a are very incompletely known (see Table 1) and the
consensus or monophyletic outgroup was used, and high number of question marks introduces con-
in no instance did the results change after im- siderable uncertainty to the analysis.
plementation of a posteriori reweighting (we used Pituriaspids are here resolved in a trichotomy with
retention index values). We also tried an all zero osteostracans and galeaspids. With ACCTRAN
ancestor as an outgroup taxon to explore the optimisation applied to the character changes on the
possibility that conodonts might be resolved as the three most-parsimonious trees this trichotomy is
sister-group to tunicates, but this also produced an supported by seven characters. There are five
identical topology. homoplasies (involving characters 35, 45, 76–78)
A number of comments are necessary about this among which pituriaspids are scored as ‘ ? ’ for three
most-parsimonious solution. Most of the nodes on (45, 76, 77). There are two synapomorphies, 31 and
this tree are supported by very few synapomorphies 80. Character 31 is coded as a question mark for
Table 1. Data matrix of 17 chordate taxa. For descriptions of characters and state assignments see section IV(3).
Numbers to the right of the data matrix are percentage values of missing data for each of our operational taxonomic units.
‘ ? ’ against Recent taxa are due to non-applicable codings or codings where comparisons are of questionable primary
homology. Note that conodonts are not the most incompletely known taxa.
pituriaspids. Character 64 records the presence of a analysis as the sister-group of jawed vertebrates may,
massive endoskeletal head shield which covers the therefore, appear radical. However, global par-
gills dorsally, and our coding of ‘ 1 ’ for pituriaspids simony resolves both attributes of aspidin as homo-
is an inference from the complexity of the brain plastic. The sister-group relationship between
endocast. Thus, in this analysis the only apparently Eriptychius and jawed vertebrates in our most
unambiguous support for the relationship of parsimonious trees is supported by nine character
pituriaspids with osteostracans and galeaspids is changes ; eight of these are homoplastic, six of which
based on inference. are scored as ‘ ? ’ for Eriptychius (4, 14, 16, 17, 40, 52).
Of the three possible resolutions to this trichotomy The other two are : character 69, which relates to the
the sister pairing of osteostracans and galeaspids acquisition of orthodentine (also seen in Astraspis and
receives no support ; that is, it is a zero length branch heterostracans) and character 70, the acquisition of
under the ACCTRAN optimisation. It is retained (monotypic) enamel (also seen in conodonts). The
because DELTRAN optimisation places characters single synapomorphy linking Eriptychius and
31 and 76 (both of which are ‘ ? ’ for pituriaspids) as gnathostomes (character 20) is coded as ‘ ? ’ for
characters linking osteostracans and galeaspids. The Eriptychius. Similarly, the association of Eriptychius
theory that pituriaspids and galeaspids are sister with the clade containing gnathostomes, osteo-
groups is supported by changes in six characters (15, stracans and galeaspids is based on eight characters,
32, 69, 71, 73, 82) all of which are homoplasies and including a single synapomorphy which is unknown
‘ ? ’ in pituriaspids. Finally, the grouping Osteostraci in Eriptychius, and a further six homoplastic character
jPituriaspida is supported by three homoplastic changes unknown for Eriptychius.
character changes in states coded ‘ ? ’ for pituriaspids. We cannot claim that the evidence for the
Therefore, the grouping of pituriaspids with osteo- association of Eriptychius with gnathostomes in
stracans and galeaspids is at best questionable and at particular or with the larger clade containing
worst disruptive in disguising synapomorphies be- gnathostomes, osteostracans and galeaspids in gen-
tween osteostracans and galeaspids which might eral is well supported. However, we have retained
otherwise resolve a sister-group relationship between Eriptychius within the tree listing character changes
these taxa. We have, therefore, undertaken another because (a) its phylogenetic position is constant
analysis with pituriaspids excluded, and we have among the three most parsimonious trees, (b) it does
chosen to depict the character changes on the single not destroy the association between gnathostomes,
resulting tree (see Figs 14 A, B, 15, 16). osteostracans and galeaspids and (c) histological
Eriptychius has traditionally been placed close to details of Eriptychius have traditionally played a
Astraspis, arandaspids and heterostracans, primarily significant part in attempts to understand both the
on the basis of common possession of spongy and interrelationships of ‘ agnathans ’ and the evolution
lamellar aspidin (the latter is also found in Loganellia of the vertebrate skeleton.
and galeaspids) ; its phylogenetic position in this The Bremer support values on our most-
A B
Tunicata Tunicata
Cephalochordata Cephalochordata
Myxinoidea Myxinoidea
Petromyzontida Petromyzontida
Conodonta Conodonta
Anaspida Astraspis
Jamoytius Anaspida
Euphanerops Jamoytius
Arandaspida Euphanerops
Galeaspida Loganellia
Pituriaspida Eriptychius
Osteostraci Jawed vertebrates
Heterostraci Pituriaspida
Astraspis Osteostraci
Loganellia Galeaspida
Eriptychius Heterostraci
Jawed vertebrates Arandaspida
C D
Tunicata Tunicata
Petromyzontida Pituriaspida
Conodonta Galeaspida
Euphanerops Osteostraci
Jamoytius Jawed vertebrates
Anaspida Eriptychius
Eriptychius Loganellia
Astraspis Euphanerops
Heterostraci Jamoytius
Arandaspida Anaspida
Loganellia Arandaspida
Galeaspida Heterostraci
Jawed vertebrates Astraspis
Pituriaspida Conodonta
Osteostraci Petromyzontida
Fig. 8. Effects of different ways of coding characters upon resolution and topology. In each case the strict consensus
tree is given with the numbers of fundamental trees, lengths, CIe – consistency indices (excluding uniformative char-
acters), RI – retention indices, and RC – rescaled consistency indices. (A) presence\absence coding ; 9 trees, 199 steps,
CIe 0.585, RI 0.646, RC 0.386. (B) contingent coding ; 22 trees, 181 steps, CIe 0.638, RI 0.700, RC 0.456. (C) Sankoff
coding ; 10 trees, 185 steps, CIe 0.662, RI 0.713, RC 0.474. (D) Effect of deletion of all hard tissue characters ; 13007
trees, 125 steps, CIe 0.718, RI 0.735 and RC 0.529.
parsimonious trees indicate how many extra steps have 89 binary characters, most of which are
must be added before a particular node collapses. presence\absence codings. There are 14 multistate
It can be seen (Fig. 7 A) that with the exception characters, all but one of which are three-state
of the nodes subtending hagfishes and lampreys characters which usually describe the absence of a
with derived sister-groups most nodes are poorly feature plus two alternative states of that feature
supported using this index. We also note that there (e.g. character 78, no head covering l 0, head
are many suboptimal trees at one step (Fig. 7 B) and covering micromeric l 1, head covering macro-
two steps (Fig. 7 C) longer ; therefore, the optimal meric l 2). This coding implies that there is a
result is not strongly supported. However, the strict transformational homology between the condition of
consensus of the trees one step and two steps longer the head covering and does not allow for the
show a range of included tree topologies funda- possibility they may be independent characters. One
mentally consistent with our most parsimonious radical alternative is extreme presence\absence
solution. More particularly, with respect to coding (Pleijel, 1995) where each separate obser-
conodonts, none of these trees is incongruent with vation is coded as a discrete character (character A,
the classification (hagfishes (lampreys (gnathostomes micromeric head covering absent l 0, present l 1 ;
j conodonts))). character B, macromeric head covering absent l 0,
present l 1). Recoding the data matrix in this way
results in 119 characters and nine trees, the strict
(a) Experimental analysis of the data set
consensus of which is shown in Fig. 8 A ; the position
Previous attempts at classifying agnathan fishes of conodonts relative to the extant chordates is
(Forey & Janvier, 1993 ; Forey, 1995 ; Gagnier common to our original optimal trees. In this
1993 b, 1995) have all yielded results that are fragile particular analysis, a heuristic search was used,
in the sense that individual nodes are poorly employing 100 replicates of a random addition
supported. The possibility then remains that slight sequence. It should be noted that major dis-
changes in coding, taxon deletion\addition or choice advantages of presence\absence coding are that
of outgroup may have severe consequences on the analyses of such matrices tend to overemphasise the
phylogenetic hypothesis. To a very large extent the plesiomorphic attributes (the ‘ 0 ’ coding) and that
fragility results from the difficulty expressed in our such coding fails to discriminate between primary
epigram. Basically, modern agnathans are known absence and secondary loss.
from only soft parts while the fossils are known only Another possible approach is to use contingent
from hard parts leading to the inclusion of many coding (Hawkins et al., 1997), in which one character
question marks in the data matrix (see legend to expresses the presence\absence of the feature (e.g.
Table 1). The presence of large numbers of question head covering), and another expresses the nature of
marks can generate many equally parsimonious that feature (e.g. micromeric l 0, macromeric l
trees, including some that are spurious (that is, some 1), with those taxa lacking the feature scored as ‘ ? ’
alternative nodes cannot be supported by any real (meaning not-applicable). Contingent recoding of
values which may be substituted for a question the multistate characters in our matrix results in 118
mark). While we can lessen the effect of large characters, analysis of which gives 22 equally
numbers of question marks we cannot eliminate the parsimonious trees (Fig. 8 B). Although many of the
basic problem, which resides with the fossils and not taxa are placed in an unresolved polytomy, the
with any particular method of analysis. position of conodonts remains more derived than
We have submitted our data set to a series of hagfishes and lampreys but less derived than
experiments in order to test the robustness of the armoured jawless vertebrates.
data, to determine the effect that coding data in A third approach is to apply a priori weights to
different ways might have, to evaluate the effects of certain character changes through the device of
the inclusion or exclusion of characters about which Sankoff coding (Sankoff & Rousseau, 1975 ; Forey &
the fossils are mute, and to address specific criticisms Kitching, in press). This type of coding applies a
regarding the inclusion of conodonts within a ‘ cost matrix ’ to the inference of character changes
chordate classification (e.g. Schultze, 1996). that take place as the computer program is searching
Different ways of coding data embody different for the most parsimonious tree(s). For example, we
theories of homology and may lead to different may specify that to transform from no head covering
phylogenetic outcomes (Pimental & Riggins, 1987 ; to micromeric costs one step, from no head covering
Forey & Kitching, in press). In our data matrix, we to macromeric costs two, from micromeric to
macromeric costs one and from either state of the these results are equivocal : for example, although
head covering to no head covering costs one. the analysis of Lipscomb et al. (1998) of small subunit
Applying these types of step matrices to the eight (SSU) ribosomal RNA indicated cyclostome
multistate characters for which they make potential monophyly when entire SSU ribosomal sequences
biological sense (characters 4, 17, 18, 25, 26, 69, 70, were analysed, when transversional changes of SSU
78), analysis results in 10 equally parsimonious trees ribosomal sequences were analysed, the data
with the strict consensus tree shown in Fig. 8 C. The supported cyclostome paraphyly (see also Philippe et
relationships remain largely compatible with our al., 1994).
preferred solution, with the position of conodonts A large number of characters used in our analysis
retained. can be identified a posteriori as synapomorphies
We also analysed the data set with all the uniting lampreys and jawed vertebrates (and thus
characters dealing with hard tissues deleted separating hagfish and lampreys), but otherwise
(characters 46–67). 13007 equally most-parsimoni- contribute nothing to the resolution of relationships.
ous trees were found with the strict consensus tree It could be considered, therefore, that our analysis is
shown in Figure 8 D ; the lack of resolution underlines an unfair test of cyclostome monophyly. To assess
the importance of hard tissue characters. this we conducted an analysis in which all but five of
As a final experiment, we eliminated all of those these characters (6, 12, 13, 19, 20) were excluded
characters in which all fossils, including conodonts, (characters 17, 23–25, 28–30, 34, 37–40, 42, 49, 50,
could only be scored as question marks (6, 9–13, 24, 55, 57, 58, 83–99, 102 were excluded). The result of
33, 36, 38, 39, 42, 56, 57, 83–103). This tests for the a branch-and-bound analysis was three trees (strict
possible tendency of soft anatomical characters to consensus shown in Fig. 9 A) which support cyclo-
place lampreys as more derived than hagfishes. stome monophyly. Addition of a further synapo-
However, the resulting topology was exactly the morphy (character 23) retained these three trees and
same as our preferred result (Fig. 7 A) and we added a further three in which there is no support for
conclude, therefore, that unavailability of soft tissue cyclostome monophyly ; the strict consensus of all six
characters in fossils is not affecting the ability of a trees is reproduced in Fig. 9 B. Addition of another
phylogenetic signal to come through (whether it is synapomorphy (character 24) resulted in the three
correct is a different matter). trees from our original analysis, exhibiting no
support for cyclostome monophyly.
(b) Cyclostome monophyly
Dume! ril (1806) first placed hagfish and lampreys
(c) Alternative hypotheses of chordate relationships
into a monophyletic group Cyclostomi (‘ rounded
mouth’) on the basis of two characters : a large Other authors have proposed very different patterns
notochord and horny teeth. Through the work of of relationship from that which we derive here, and
Cope (1889) the fundamental significance of the we have selected four of these to act as backbone
presence and absence of jaws was first recognised constraint trees. The mutual relationships of the taxa
and the concept of a monophyletic Cyclostomata considered by the original authors were maintained
persisted until the early 1970s. Subsequently, in each case while the remaining taxa were allowed
Løvtrup (1977), Hardisty (1979, 1982) and a to establish their own relationships within this
number of other anatomists and palaeontologists framework (strictly, we used simple constraint trees
recognised that lampreys shared more derived to represent previous hypotheses of relationships,
characters with jawed vertebrates than with hag- and taxa not included in previous analyses were
fishes, and the Cyclostomata came to be regarded as placed in an unresolved polytomy at the base of the
a paraphyletic group. However, molecular studies starting tree ; attempts at using conventional
have rekindled the concept of a monophyletic backbone-constraint trees, in which only taxa con-
Cyclostomata (Stock & Whitt, 1992 ; Lanfranchi et sidered by a previous author are included in the
al., 1994 ; Turberville, Schultz & Raff, 1994 ; Suzuki constraint tree, failed due to the effect of numerous
et al., 1995 ; Mallatt & Sullivan, 1998 ; Lipscomb et synapomorphies stacked on the nodes separating
al., 1998), and Yalden (1985) and Mallatt (1997 a, b) cephalochordates, hagfishes and lampreys). This
have both argued for cyclostome monophyly on kind of experiment is in no way a test or criticism of
the basis of morphological characters (usually the the original theories : two of the four trees (Fig. 10 A,
feeding apparatus and characters therein). Some of B) were not based on cladistic methods, all use only
Fig. 9. (A) Cyclostome constraint tree ; 3 trees, 128 steps, CIe 0.614, RI 0.667, RC 0.411. (B) Cyclostome constraint
tree with synapomorphies removed ; 6 trees, 130 steps, CIe 0.612, RI 0.664, RC 0.409 ; (see text for discussion).
a subset of the taxa used in our analysis, none were probably have little effect on our understanding of
based on our data set, and three (Fig. 10 A–C) were the phylogenetic significance of conodonts. However,
based on a much smaller and eclectic set of in the absence of such strong phylogenetic signals we
characters. Thus, very little should be read into the conclude that the relationships (and hence the
differing tree-lengths and other statistics. What is of significance) of conodonts cannot be established
interest is that none of these analyses places independently of a theory of relationships of
conodonts in our favoured position ; within the chordates including conodonts. Furthermore, the
constrained analyses conodonts are resolved as either effect of exclusion of conodonts from our analysis
stem-craniates or stem-vertebrates. We interpret this (Fig. 11 A) demonstrates that the unique suite of
to suggest that the phylogenetic position of cono- chordate characters expressed by conodonts has
donts is dependent upon the phylogenetic relation- great significance for our understanding of chordate
ships of other chordates and so, if we assume different phylogeny.
hypotheses of relationship amongst non-conodont
chordates, the phylogenetic position of conodonts is
very differently resolved. Thus, to place conodonts
(d ) The effects of alternative interpretations of conodont
into an existing chordate classification as a means of
anatomy and histology
explaining character evolution and the phylogenetic
significance of conodonts, it is first necessary to Although we contend that homology between
justify the choice of chordate phylogeny. All previous conodont and vertebrate hard tissues is well-
studies (e.g. those cited in Fig. 4 of Aldridge & founded, several authors have expressed doubts (e.g.
Purnell, 1996), bar that of Janvier (1996 b), have Forey & Janvier, 1993 ; Kemp & Nicoll, 1995 a, b,
failed to do this. 1996 ; Schultze, 1996 ; Pridmore et al., 1997). For this
If there were several unambiguous synapo- reason we have undertaken analyses with all the
morphies which linked conodonts with one or histological characters set to question marks against
another terminal taxon (e.g. cephalochordates), the conodonts. As a separate experiment, we have also
differing theories of chordate relationships would set the presence of eye muscles to a question mark, to
A B
Tunicata Tunicata
Heterostraci Conodonta
Myxinoidea Arandaspida
Conodonta Astraspis
Jamoytius Galeaspida
Arandaspida Eriptychius
Osteostraci Jawed vertebrates
Petromyzontida Jamoytius
Anaspida Euphanerops
Pituriaspida Loganellia
Loganellia Myxinoidea
Jawed vertebrates Anaspida
Astraspis Petromyzontida
Eriptychius Osteostraci
C D
Tunicata Tunicata
Myxinoidea Conodonta
Conodonta Pituriaspida
Pituriaspida Eriptychius
Arandaspida Euphanerops
Euphanerops Jamoytius
Eriptychius Myxinoidea
Jawed vertebrates Petromyzontida
Loganellia Osteostraci
Heterostraci Galeaspida
Astraspis Arandaspida
Osteostraci Astraspis
Petromyzontida Anaspida
Anaspida Jawed vertebrates
Jamoytius Loganellia
Fig. 10. Result of incorporating backbone constraint trees (bold lines) based upon existing hypotheses of relationships.
(A) Stensio$ (1968) constraint tree ; 1 tree, 215 steps, CIe 0.547, RI 0.527, RC 0.290. B. Moy-Thomas & Miles (1971)
constraint tree ; 2 trees, 248 steps, CIe 0.474, RI 0.369, RC 0.176. C. Halstead (1982) constraint tree ; 1 tree, 206
steps, CIe 0.571, RI 0.573, RC 0.328. D. Gagnier (1995) constraint tree ; 9 trees, 209 steps, CIe 0.562, RI 0.558, RC
0.315.
test the contentions that conodonts are resolved as sidered Tunicates and Cephalochordates to be more
vertebrates through assumed homologies of the hard closely related to each other than either group is to
tissues and eye muscles (Kemp & Nicoll, 1995 a, b, conodonts, hagfish or lampreys. We conducted a
1996 ; Schultze, 1996 ; Pridmore et al., 1997). These further analysis incorporating this constraint ; the
experiments produced identical results (Fig. 11 B), result is 42 most-parsimonious trees at 193 steps with
with conodonts again resolved as cladistically more a topology meeting the requirements of the backbone
derived than lampreys. Setting the character codings constraint tree (Fig. 12 B). In both instances, the
for the same characters to ‘ 0 ’ instead of ‘ ? ’ for topology of the strict consensus tree is the same (Fig.
conodonts produces an identical consensus tree. 12 A).
Thus, even if we claim that the histological structures Kemp & Nicoll (1995 a, 1996) proposed that
in conodonts and the conodont eye muscles are conodonts are more closely related to the cephalo-
definitely not those of vertebrates, conodonts are still chordates than to any other group of chordates.
resolved as more derived than lampreys. The Analysis of the data set while enforcing a backbone
synapomorphy supporting this position in this case is constraint tree compatible with this hypothesis
the presence of a mineralised dermal skeleton produces 42 equally most-parsimonious trees at 190
(character 65). Conodont hard tissues have been steps (Fig. 12 C).
recognised as dermal at least since the work of Gross Both Peterson (1994) and Pridmore et al. (1997)
(1957, 1960), an hypothesis that has not been suggested that conodonts might best be positioned as
disputed. Nevertheless, if we experiment with this a sister-group to the Craniata. 45 trees at 186 steps
coding by scoring ‘ ? ’ for the presence of a dermal are identified as the shortest compatible with their
skeleton, ‘ ? ’ for the presence of specific hard tissues, hypothesis (Fig. 12 D). Krejsa, Bringas & Slavkin
and ‘ ? ’ for the presence of extrinsic eye musculature (1990 a, b) argued that conodonts and hagfish are
in conodonts, parsimony analysis results in 48 more closely related to each other than either is to
optimal trees in which the relationship between any other group based largely upon putative
conodonts and lampreys is unresolved (Fig. 11 C). If homology between conodont elements and hagfish
these codings are all changed to ‘ 0 ’ for conodonts, toothlets. A branch-and-bound search found 45 trees
parsimony analysis identifies 42 optimal trees, the at 186 steps as the shortest compatible with this
strict consensus of which places conodonts below hypothesis ; a strict consensus is shown in Fig. 13 A.
lampreys and above hagfishes (Fig. 11 D). The hypothesis of conodont affinity commonly
preferred by workers on non-skeletal remains places
conodonts as a sister group to the lampreysjjawed
(e) Testing alternative hypotheses of conodont affinity
vertebrates (Aldridge et al., 1986, 1993 ; Aldridge &
Despite the limited data, conodonts have been Briggs, 1986 ; Aldridge, 1987 ; Aldridge & Purnell,
interleaved in almost every possible permutation 1996 ; Aldridge & Donoghue, 1998 ; Donoghue et al.,
with the major groups of living chordates. To 1998). 45 trees at 182 steps are identified as the
compare the results of our analysis with the various shortest compatible with this hypothesis ; a strict
previous hypotheses we have re-analysed our data consensus of all appears in Fig. 13 B. Alternatively,
set enforcing topological backbone constraint trees Janvier (1996 b) has proposed that conodonts and
representing each hypothesis. lampreys are sister-taxa ; we found three trees at 182
In discussing the affinity of Yunnanozoon, Dzik steps as equally most-parsimonious compatible with
(1995) proposed that conodonts are more primitive this constraint topology (Fig. 13 C). The topology of
than all the living chordates. Assessing this hy- the strict consensus tree is otherwise common to the
pothesis is problematic as it necessitates moving strict consensus of the three optimal trees generated
Tunicata and Cephalochordata to the in-group ; for without a backbone constraint (Fig. 7 A). Janvier
the purposes of this experiment we have adopted an (1995) has also proposed that conodonts might be
all-zero outgroup. Branch-and-bound analysis more closely related to a group including the
identified 42 most-parsimonious trees at 192 steps Osteostraci, Pituriaspida and crown-group gnatho-
with a topology meeting the requirements of the stomes, based upon a possible homology between
backbone constraint tree (Fig. 12 A), in contrast conodont white matter and mesodentine\cellular
with the 180 steps in our optimal tree (Fig. 7 A). dermal bone. Branch-and-bound search identified
However, this is not necessarily an adequate test of six trees at 187 steps as the shortest compatible with
how Dzik’s (1995) hypothesis of conodont affinity the enforced backbone constraint tree ; the topology
compares to our preferred hypothesis, as he con- of a strict consensus is shown in Fig. 13 D).
A B
Tunicata Tunicata
Anaspida Conodonta
Astraspis
Jamoytius
Heterostraci
Euphanerops
Arandaspida
Loganellia
Anaspida
Eriptychius
Jamoytius
Jawed vertebrates
Euphanerops
Pituriaspida
Loganellia
Osteostraci
Eriptychius
Galeaspida Jawed vertebrates
Astraspis Osteostraci
Heterostraci Pituriaspida
Arandaspida Galeaspida
C D
Tunicata Tunicata
Conodonta Conodonta
Anaspida Anaspida
Jamoytius Jamoytius
Euphanerops Euphanerops
Loganellia Loganellia
Jawed vertebrates Jawed vertebrates
Pituriaspida Pituriaspida
Osteostraci Osteostraci
Galeaspida Galeaspida
Astraspis Astraspis
Heterostraci Heterostraci
Arandaspida Arandaspida
Fig. 11. (A) Effect of exclusion of conodonts from the analysis ; 42 trees, 178 steps, CIe 0.661, RI 0.703, RC 0.466.
(B) Effect of changing conodont histological characters and the eye muscle character to question marks (a number of
separate analyses all resulted in the same topology, but with slightly different statistics) ; 3 trees, 178 steps, CIe 0.661,
RI 0.706, RC 0.468. C. Effect of changing all histological characters, extrinsic eye musculature and ‘ dermal skeleton ’
in conodonts to ‘ ?’ ; 48 trees, 178 steps, CIe 0.661, RI 0.706, RC 0.468. D. Effect of changing all histological char-
acters, extrinsic eye musculature and ‘ dermal skeleton ’ in conodonts to ‘ 0’ ; 42 trees, 178 steps, CIe 0.661, RI 0.710,
RC 0.471.
( f ) Summary of conclusions drawn from experimental according to the scala naturae (Northcutt, 1981).
analysis of our data set However, both hagfishes and lampreys are com-
monly thought to be wholly unrepresentative of the
The results of our phylogenetic experiments show ancestral vertebrate state (Northcutt, 1985), due to
that conodonts are consistently placed within verte- their highly specialised modes of life (Conway
brates and that, with the exception of one type of Morris, 1989b ; Northcutt, 1996 b) which are
coding (substituting ‘ 0 ’ for eye muscle and all reflected in an assumed derived morphology.
histological characters in conodonts) the results are Northcutt (1981) and Wicht & Northcutt (1992)
consistent with the proposition that conodonts are have, therefore, adopted a cladistic approach to
cladistically more derived than lampreys, that is : resolving the condition of the hypothetical common
they are basal members of the Gnathostomata. In ancestor, or morphotype, at the respective nodes.
commentary on the epigram to this section we would The topological relationship between hagfishes,
suggest that the presence of ‘ V ’-shaped myotomes lampreys and jawed vertebrates that results from our
and paired sensory organs place conodonts within phylogenetic analysis concurs with many previous
the Chordata but that the extra information required studies and so reveals nothing more regarding the
to place them more accurately centres on interpret- evolution of the brain and sensory systems than
ation of conodont elements as part of a mineralised covered in the excellent reviews by Braun (1996),
dermal skeleton. Northcutt (1996 a), and Wicht (1996). However, a
number of subsequent studies have yielded data
(5) Character changes pertinent to the early evolution of the brain and
sensory systems of vertebrates ; these were incor-
(a) General porated into our analysis and the implications of the
The topology of our most-parsimonious tree (Fig. character changes which arise from our work are
7 A) is very similar to the results of other recent discussed here.
phylogenetic analyses of primitive vertebrates. As Weigle & Northcutt (1998) have provided pre-
such, the implications of character changes for the liminary results of a restudy of the lamprey cer-
phylogeny of many organ systems differ little from ebellum which indicate that a cerebellum is not
these publications, and in these instances we direct present in lampreys. The optimisation used
the reader to the work of Forey & Janvier (1993), (ACCTRAN ; Figs 14 A and 15) suggests that a true
Forey (1995) and Janvier (1996 a, b). Below we cerebellum first evolved in conodonts. However, this
discuss the implications for phylogeny where the conclusion may be an artefact of optimising the
results of our analysis differ, and where new data coding of this character (number 8) as a question
have provided a better understanding of character mark in Conodonta, Jamoytius, Euphanerops,
evolution. Detailed paths of character changes are Anaspida, Eriptychius and Astraspis. Evidence for a
shown optimised onto the most-parsimonious tree cerebellum occurs in heterostracans, galeaspids,
after the deletion of pituriaspids (see above for jawed vertebrates and osteostracans (and possibly
justification of this) ; Fig. 14 A shows ACCTRAN pituriaspids), and so a conservative estimate for the
optimisation while Fig. 14 B shows DELTRAN origin of the cerebellum would be at the node
optimisation. If there are equally parsimonious ways representing the common ancestor of heterostracans
of optimising a homoplastic character change (e.g. if and galeaspids (thus including Astraspis ; see Fig.
the alternatives mean that either the character has 14 B, DELTRAN optimisation).
been gained and independently lost, or that the A sensory line system is first seen in Eptatretus, the
character is gained twice) then ACCTRAN favours Pacific hagfish ; no other hagfish possesses a sensory
gain and loss while DELTRAN favours parallel line system and the sensory hair cells of the
gains. A choice between these alternatives can only mechanoreceptors which comprise the sensory line
be made by invoking ad hoc assumptions. system are unpolarised and lack cupulae, both of
which are characteristic of true neuromasts. It is not
clear whether this condition is representative of the
(b) Nervous and sensory systems
plesiomorphic condition for vertebrates or results
For over a century, hagfishes and lampreys have from secondary reduction and eventual loss in other
been taken as proxies for the most primitive of hagfishes (Braun, 1996 ; Braun & Northcutt, 1997,
vertebrates and the condition of their organ systems 1998). Braun & Northcutt (1997, 1998) described
has been taken as the plesiomorphic condition the presence of three groups of sensory grooves :
A B
All zero All zero
Conodonta Conodonta
Tunicata Cephalochordata
Cephalochordata Tunicata
Anaspida Osteostraci
Jamoytius Galeaspida
Loganellia Eriptychius
jawed vertebrates jawed vertebrates
Pituriaspida Euphanerops
Galeaspida Jamoytius
Astraspis Astraspis
Arandaspida Heterostraci
Heterostraci Arandaspida
C D
Tunicata Tunicata
Conodonta Cephalochordata
Cephalochordata Conodonta
Anaspida Galeaspida
Jamoytius Pituriaspida
Euphanerops Osteostraci
Loganellia jawed vertebrates
jawed vertebrates Eriptychius
Pituriaspida Jamoytius
Galeaspida Euphanerops
Astraspis Astraspis
Fig. 12. (A–D) Constraint trees based on previous hypotheses of conodont affinity amongst the Chordata (see text for
discussion). (A, B) Dzik (1995) constraint trees ; (A) 42 trees, 192 steps, CIe 0.613, RI 0.694, RC 0.307 ; (B) 42 trees,
193 steps, CIe 0.609, 0.690, 0.422. (C) Kemp & Nicoll (1995b) constraint tree ; 42 trees, 190 steps, CIe 0.619, RI
0.650, RC 0.404. (D) Peterson (1994) and Pridmore et al. (1997) constraint tree ; 45 trees, 186 steps, CIe 0.632, RI
0.670, RC 0.425. Dotted lines represent taxa not considered in the original analysis.
A B
Tunicata Tunicata
Conodonta Myxinoidea
Myxinoidea Conodonta
Galeaspida Galeaspida
Pituriaspida Pituriaspida
Osteostraci Osteostraci
jawed vertebrates jawed vertebrates
Loganellia Loganellia
Jamoytius Jamoytius
Euphanerops Euphanerops
Anaspida Anaspida
Astraspis Astraspis
C D
Tunicata Tunicata
Conodonta Petromyzontida
Petromyzontida Jamoytius
Astraspis Euphanerops
Arandaspida Anaspida
Heterostraci Eriptychius
Anaspida Astraspis
Jamoytius Arandaspida
Euphanerops Heterostraci
Loganellia Galeaspida
Eriptychius Conodonta
jawed vertebrates Loganellia
Osteostraci jawed vertebrates
Pituriaspida Osteostraci
Galeaspida Pituriaspida
Fig. 13. (A–D) Constraint trees based on previous hypotheses of conodont affinity amongst the Chordata (see text for
discussion). (A) Krejsa et al. (1990 a, b) constraint tree ; 45 trees, 186 steps, CIe 0.632, RI 0.670, RC 0.425. (B)
Aldridge et al. (1986) constraint tree ; 45 trees, 182 steps, CIe 0.646, RI 0.689, RC 0.447. (C) Janvier (1996 b) con-
straint tree ; 3 trees, 182 steps, CIe 0.646, RI 0.689, RC 0.447. (D) Janvier (1995, hypothesis b) constraint tree ; 6
trees, 187 steps, CIe 0.629, RI 0.665, RC 0.420. Dotted lines represent taxa not considered in the original analysis.
228
(a)
Tunicata
0
Cephalochordata
1
0 2 1 1 1 1 1 1 0 0
0 1 1 1 2 0
Myxinoidea Petromyzontida
9 4 3 3 4 1 1 1 6 6 4 4 4 1
5 0 6 3 8 5 0 4 1 5 4 3 1
0 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 2 1 1 1 1 1 1 1 3
1 1 5 5 5 5 5 4 4 2 2 2 2 1 1 1 1 1 1 7 6 5 2 1 1 1
0 0 6 4 3 2 1 6 3 7 6 5 0 8 7 6 4 1 0
3 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 0 1 1 0 1 1 1 1 1 1 1 2 1 1 2 1 2 1 1 2 Conodonta
7 7
7 0
1 9 9 9 9 9 9 9 9 9 9 8 8 8 8 8 8 8 5 5 5 5 4 4 4 3 3 3 3 3 2 2 2 2 2 2 1 1 1 1 6 2 1 1
0 9 8 7 6 5 4 3 2 1 0 9 8 7 6 5 4 3 8 7 5 0 9 2 0 9 8 7 4 0 9 8 5 4 3 0 9 7 3 2
2 Astraspis
1 1 1 2 1 0 1 7 2 4 1 1 2 0 0
0 5
Arandaspida
7 7 7 6 6 4 1 1 2 1 8 8 7 6 2
0 0 2 1 1 0 0 1 0 0 1 0 1 1 1 6 5 1 9 6 9 4 2 1 4 9 9
7 7 7 1 0 2
1 1 7 6 6 5 5 4 3 3 2 1 9 8 3 9 8 2
0 0 3 5 0 6 3 1 6 3 1 0 Heterostraci
1 0 5 3 2
0 0 6
1
1 1 1 1 1 Anaspida
2 1 0 7
7 7 6 6 5 6 1
8 4 7 2 9
7 6 2 0 1 Jamoytius
4 1 9 3
5
7 1
8 7 Euphanerops
1
1 1 1 Loganellia
7
3
4 4 2
7 5 2
1 1 1 0 1 1 2 1
1 2 0 1 3 0 0 1 1
Eriptychius
8 7 6 5 4 3 2 6
1 7 9 4 4 2 6 6
7 6 5 4 2 1 1 1 4 1 0 0 0
0 9 2 0 0 7 6 4 Jawed
6 6 4 2 vertebrates
1 1 1 1 2 1 0 0 8 7 3 7
8 7 6 6 4 4 4 3
2 1 4 3 9 8 7 0 0 1 0 0 0 0 1
1 2 0 1 0 1 1
Galeaspida
8 7 7 6 4 3 1
2 3 1 9 8 2 5
8 7 7 7 4 3 3 1 0 0
0 8 7 6 5 5 1
Osteostraci
6 6 3
8 7
Fig. 14 a. For legend see opposite.
P. C. J. Donoghue, P. L. Forey and R. J. Aldridge
Conodont affinity and chordate phylogeny

(b)
Tunicata
0 0 1
Cephalochordata
1 9 4
0 5 0 0 1 0 1 1 1 1 1 1 2 1 1 1 1 1 1 1 1 1 1 1 1 1
1 1 0
Myxinoidea Petromyzontida
3 3 1 1 9 5 5 4 4 2 2 1 1 1 1 1 6 1 6 6 5 5 4 4 1
6 3 0 0 5 6 3 3 0 5 0 8 7 5 1 0 0 4 1 6 3 5 4 0
1 1 1 1 1 1 1 2 1 1 1 1 3 1 1
5 5 5 4 2 2 1 1 7 5 2 1 1 1
4 2 1 6 7 6 6 4
1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 0 1 1 1 1 1 1 1 1 1 2 1 1 2 1 1 2 1 1 2 2 Conodonta
7 7
7 0
1 9 9 9 9 9 9 9 9 9 8 8 8 8 8 8 8 5 5 5 5 4 4 3 3 3 3 3 2 2 2 2 2 2 1 1 1 1 1 6 4 2 2 1 1
0 9 8 7 6 4 3 2 1 0 9 8 7 6 5 4 3 8 7 5 0 9 2 9 8 7 4 0 9 8 5 4 3 0 9 8 7 3 2
2 Astraspis
1 1 1 7 6 2 4 1 1 2 0
0 9 5
Arandaspida
7 7 6 1 2 1 1 8 8 7 2
2 1 5 1 6 2 1 4 9
7 7 7 7 2 1 0 0 2 1
7 6 9 8 6 2
3 5 Heterostraci
6 5 4 3 2 1
9 0 9 0 6 4
1 1 1 1 1 1 1 1
1
7 7 6 6 4 2 8 3
8 4 7 2 3 1 Anaspida
2 1 0 7
6 1 1
7 4 2
4 7 9
0 1 Jamoytius
3 1
5 7
7 6
8 1 Euphanerops
1
1 1 1 1
4
5 Loganellia
7 7 6 4
7 3 9 7
1 1 2 1
1 1 2 6
Eriptychius
4 3 2
4 2 6 6
7 7 6 0 0 1 1 1 0 0 0 1 0 1 0 0 0 3 0 0 1 1 1 1
1 0 9 Jawed
1 1 8 7 6 6 5 5 4 4 4 3 3 2 2 1 1 1 1 9 4 vertebrates
1 1 1 1 1 2 1 0 1 0 0 2 7 8 7 4 2 8 3 0 6 3 7 0 7 6 4 1
3 0
8 6 6 6 5 4 4 3 2
1 4 3 0 9 9 1 0 2 1 1 1 1 0 1 0 0
1 2 1 1 1
Osteostraci
8 7 6 6 6 4 4 3
2 1 9 8 7 8 5
8 7 7 3 3 1 0 1
0 8 6 5 1
Galeaspida
7 3 1
3 2 5
Fig. 14. Character change trees ; each character change is denoted by a box below which appears the character number, and above
which appears the character state. Boxes and arrowheads represent the type of change : a solid black cell represents a synapomorphy,
a right-facing arrowhead represents a forward homoplasy, and a left-facing arrowhead represents a homoplasy reversal. (A) Under
ACCTRAN (accelerated transformation) optimisation. (B) Under DELTRAN (delayed transformation) optimisation. Diagrams
should be read in landscape.
229
Jawed vertebrates
Cephalochordata
Petromyzontida
Euphanerops
Arandaspida
Heterostraci
Myxinoidea
Galeaspida
Osteostraci
Conodonta
Eriptychius
Loganellia
Jamoytius
Anaspida
Astraspis
Tunicata
26 a GAIN: neural crest, brain, divided pituitary,

25 Adenohypophysis (simple), optic tectum, flattened
24 spinal chord, united d+v spinal nerve roots, single
23 median nasal opening, single nasohypophysial
22 opening (terminal), paired olfactory, single scc,
21 cranial lateral line grooves
20 b GAIN: segmented and compartmentalised
19 adenohypophysis, two types giant Mauthner cells,
18 synaptic ribbons in retinal receptors, dorsal
nasohypophysial opening, extrinsic eye musculature,
17 two scc, sensory-line neuromasts, electroreceptive cells,
16 sensory-lines on head and body
15 c GAIN: olfactory tract, cerebellum, pretrematic
14 branches of branchial nerves, flattened spinal
13 chord, vertical scc
12 d GAIN: paired nasal openings
11 e GAIN: externally open endolymphatic ducts
10 f GAIN: terminal nasohypophysial opening
9 g GAIN: sensory-line system enclosed in canals
8 h GAIN: pineal present but uncovered, paired nasal
7 openings, 3 semicircular canals
6 LOSS: single nasohypophysial opening
5
4
3
2
1
f h
d g
c
b
a
Fig. 15. Nervous character distribution under ACCTRAN optimisation. Symbols represent different character states.
preoptic, dorsal postoptic, and ventral postoptic. neuromasts arranged into a number of lines on both
The preoptic grooves are innervated by a single pair the head and trunk (Johnston, 1905), many of which
of ganglionated cranial nerves and the two postoptic have been homologised with counterparts in jawed
groove series are innervated by a pair of cranial vertebrates (Northcutt 1985, 1989).
nerves which each possess two ganglia, possibly Sansom et al. (1997) have recently described the
resulting from the fusion of two pairs of ganglionated presence of an asymmetrical pair of lateral line
nerves. grooves caudal to the pineal region of Astraspis,
Many characteristics of the sensory line system of possibly representing postpineal lines (they
lampreys are common to the lateral line system of questioned whether these structures are the same as
jawed vertebrates. The mechanoreceptors lack described by Stensio$ , 1964, p. 178). Heterostracans
cupulae and are composed of a polarised arrange- possess a well-developed lateral line system enclosed
ment of sensory hair cells and occur as free in canals which run through the middle layer of
dermal skeleton ; possible homologies of gnathostome gnathostomes, and possibly also within other groups.
lateral lines have been identified by Northcutt (1985 ; Thus, there is no evidence of a simple phylogenetic
1989) but it must be stressed that the pattern of pattern of free to canal-enclosed neuromasts. An
lateral line distribution is highly variable (Blieck, explanation may be afforded by consideration of
1984). Sacabambaspis bears evidence of a lateral line generalised lateral line development in gnathostomes
system where neuromasts were located in shallow (Northcutt et al., 1994). If this model is applicable to
grooves in the dermal exoskeleton (Gagnier, 1993a), all lower vertebrates it is possible that the super-
possibly representing homologues of the supra- ficially random pattern of lateral-line evolution is
orbital, infraorbital, mediodorsal (fragmented), due solely to the effect of heterochronic pattern-
lateroventral longitudinal, and both dorsal and producing processes upon a compartmentalised
ventral trunk lines, many of which are directly developmental sequence (e.g. Northcutt et al., 1994 ;
comparable to the condition in heterostracans. Braun & Northcutt, 1997 ; Northcutt, 1997). How-
Smith (1957) described a number of shallow grooves ever, following Raff (1996), we recognise hetero-
in the dermal skeleton of Pharyngolepis (an anaspid) chrony only in its explanatory powers for recognising
which he, and subsequently Northcutt (1989), were and describing pattern ; heterochrony is not a
able to identify as homologous with specific lines of process.
lampreys and gnathostomes ; evidence of a lateral The lack of definitive morphological criteria on
line system is not apparent in specimens of Jamoytius which the presence of electroreceptive capability can
or Euphanerops studied by us. The lateral line system be recognised in fossils leads to difficulty in inter-
of thelodonts is generally poorly known although preting the phylogeny of electroreception in verte-
Ma$ rss (1979) has described fully the condition for brates. Hagfishes possess no such capability (Braun,
Phlebolepis. Evidence from isolated scales indicates 1996) but lampreys are electroreceptive (Bodznick
that a lateral line system enclosed within specialised & Northcutt, 1981 ; Bullock, Bodznick & Northcutt,
canal scales occurred in Loganellia (P. C. J. 1983). Of the remaining jawless gnathostomes, only
Donoghue, personal observations). The lateral line the heterostracans, galeaspids and osteostracans
system of osteostracans has been described and have been interpreted as having possessed electro-
discussed by Janvier (1974) and Northcutt (1985, reception. In heterostracans and osteostracans, it has
1989) although little is known of the distribution of been suggested that the enigmatic pore canal system
lines on the ventral surface of the head shield, or on is an extension of the lateral line system and that the
the trunk. The lateral line system of galeaspids is as flask-shaped cavities within this system are the sites
variable as in heterostracans, and the neuromasts of electroreceptors (Denison, 1951, 1964 ; Thomson,
were situated within deep grooves which were 1977 ; Northcutt & Gans, 1983 ; Northcutt, 1985).
arranged in a scalloped pattern reminiscent of More recently, however, evidence has been pre-
anchipteraspids amongst heterostracans, but gen- sented to cast doubt on the validity of this linkage.
erally considered an autapomorphy of the The dermal skeleton of Palaeozoic lungfishes is
Galeaspida (e.g. Janvier, 1996a). In some poly- composed of a tissue combination known as cosmine
branchiaspiforms, the lateral line system is enclosed which is permeated by a pore canal system directly
in canals which penetrate the dermal skeleton and comparable with the pore-canal systems of hetero-
open at their extremity into pits which may represent stracans and osteostracans. Bemis & Northcutt
the sites of ampullary organs. (1992) demonstrated a comparable system of canals
There is no clear pattern of evolution in the in Recent lungfishes, which lack a mineralised
condition of the lateral line system during this early skeleton ; this canal-system is vascular and Bemis &
phase of vertebrate phylogeny. The mechano- Northcutt (1992) suggested that the pore-canal
receptive hair cells of eptatretid hagfishes are located system in Palaeozoic lungfishes was similarly vas-
in shallow grooves, as presumably were the neuro- cular, linked to the secretion and resorption of the
masts in lateral lines of Astraspis, arandaspids and mineralised skeleton itself. Clearly, we can no longer
most galeaspids. In lampreys, neuromasts appear as accept the presence of a pore-canal system as
free organs arranged in lines (Kleerekoper, 1972), undoubted evidence for electroreception ; more re-
and osteostracans are polymorphic, as the lateral search is required.
lines include both shallow grooves and sites of Further evidence exists for electroreception in
individual free neuromasts (cf. Wangsjo$ , 1952). osteostracans. Stensio$ (1927) suggested that the
Lateral lines arranged in canals have been attained cephalic sensory fields were the site of electro-
independently in heterostracans, thelodonts and receptors, although both Bohlin (1941) and Wangsjo$
Jawed vertebrates
Cephalochordata
Petromyzontida
Euphanerops
Arandaspida
Heterostraci
Myxinoidea
Galeaspida
Osteostraci
Conodonta
Eriptychius
Loganellia
Jamoytius
Anaspida
Astraspis
Tunicata
86
85
84
83
82
81
80 a GAIN: neural crest, creatine phosphatase,
79 horny teeth, fin radials, visceral arches fused to
78 neurocranium
77 b GAIN: calcified dermal skeleton, polyodontida
76 LOSS: horny teeth
75 c GAIN: trunk dermal skeleton, lamellar aspidin,
74 diamond shaped-scales, dermal head covering
73 d GAIN: spongy aspidin, orthodentine, three-layered
72 exoskeleton, oak-leaf tubercles, oral plates
71 e GAIN: cancellar layer, macromery, unpaired
70 headplates
69 f GAIN: annular cartilage, rod-shaped scales,
68 g LOSS: dermal head scales
67 h GAIN: mesodentine, pharyngeal denticles,
66 scleroctic ossicles
65 i GAIN: perichondral bone, calcified cartilage, three
64 layered exoskeleton,
63 jGAIN: orthodentine, enamel
62 LOSS: visceral arches fused to neurocranium
61 k GAIN: oral plates, macromeric dermal head
60 scales, massive endoskeletal head shield
59 LOSS: pharyngeal denticles
58
57
56
55
54
53
52
51
e g j k
i
d f
h
c
b
a
Fig. 16. Hard tissue character distribution under ACCTRAN optimisation. Symbols represent different character
states.
(1952) reinterpreted the fields as sites of mechano- Carboniferous (a subsequent gap of approximately
receptors. Northcutt (1985) also suggested that these 300 million years).
structures housed mechanoreceptors, representing Our phylogenetic tree (Fig. 16 & see Fig. 17)
extensive evaginations of the membranous labyrinth. suggests that a mineralised skeleton evolved after the
Notwithstanding the function of the pore-canal origin of the hagfish and lamprey lineages, and is
system or cephalic sensory fields of osteostracans, first expressed in conodonts in the form of a
Northcutt (1985) suggested that the presence of a mineralised feeding apparatus composed from
ramus lateralis communicans interconnecting the odontodes of enamel and dentine. The next most-
ganglia of the anterior and posterior lateral line derived group of vertebrates are the pteraspido-
nerves deduced to have been present in Procephalaspis morphs (sensu Gagnier, 1995, uniting Astraspis,
(Janvier, 1974) indicated that osteostracans were heterostracans and arandaspids), all of which possess
electroreceptive, because this is a diagnostic feature an extensively developed dermal exoskeleton com-
of electroreceptive species that possess trunk electro- posed predominantly from aspidin, with a sub-
receptors innervated by the lateral line nerve (Ronan ordinate component of orthodentine. This sequence
& Northcutt, 1987). Future research into the of character changes indicates that odontodes are the
phylogeny of electroreception, particularly amongst primitive patterning unit of the vertebrate dermal
the extinct groups of jawless gnathostomes might be skeleton. As originally defined (Ørvig, 1967),
directed towards consideration of the relative ar- odontodes ‘ consist of dentine or dentinous tissue and
rangement of lateral line neuromasts and potential in most cases of superficial enameloid substance,
evidence for the presence of electroreceptors ; and...arise ontogenetically, each of them, in a single,
Northcutt, Bra$ ndle & Fritsch (1995) have demon- undivided dental papilla adjoined peripherally by
strated that both arise from common placodes in the an epithelial dental organ of epidermis cells ’ (Ørvig,
axolotl. Thus, evidence of a common association of 1967 : p. 389). Reif (1982) later modified the
lateral lines and electroreceptors would provide odontode concept to an isolated superficial structure
better evidence (e.g. Ørvig, 1971, for brachythoracid of the dermal skeleton which consists of a dentinous
arthrodires) than evidence for connection between tissue, a hypermineralised cap of enamel or
lateral line canals and putative electroreceptive enameloid (which may or may not be present), and
complexes (e.g. Denison, 1951). a bony base (either cellular or acellular) which
functions as an attachment tissue ; the odontode
forms within a single, undivided dental papilla of
mesenchyme bounded at its outer surface by an
(c) Skeleton
epithelial dental organ ; vascular supply through the
Although both hagfishes and lampreys are capable basal and\or neck canals ; attachment is enabled by
of biomineralisation (Carlstro$ m, 1963) and lampreys anchoring fibres which originate in the bony base, or
may be capable of calcifying their endoskeleton ankylosis to underlying bone. Thus far, no histo-
(Bardack & Zangerl, 1971 ; Langille & Hall, 1993), logically investigated conodont exhibits evidence of
neither group expresses the ability to produce a bone of attachment and it must be assumed that
mineralised exoskeleton and both are assumed to be primitively, odontodes were composed solely from
primitively naked. The fossil record of hagfishes enamel and dentine ; bone of attachment must have
extends back to the Upper Carboniferous (approxi- evolved after the divergence of conodonts. Evidence
mately 300 Ma ; Bardack, 1991), and the oldest fossil for attachment tissues has been presented by Sansom
lampreys are of Lower Carboniferous age (approxi- (1996), who described the presence of extinction
mately 320 Ma ; Lund & Janvier, 1986). However, patterns in the dentine base of Pseudooneotodus
because our analysis suggests that conodonts are elements, which he interpreted as either backfilled
more derived than hagfishes and lampreys, and dentine tubules or attachment fibres ; comparison
because the fossil record of conodonts extends at least with extinction patterns in the bone of attachment in
as far back as the Late Cambrian (Aldridge & more derived vertebrates suggests that the latter
Smith, 1993), our phylogeny indicates that the interpretation is most likely. The subsequent evol-
hagfish and lamprey lineages are at least a further utionary history of bone and attachment tissues
220 and 200 million years older (respectively) than depends critically upon the interpretation of aspidin
their fossil record would suggest. This is not which comprises the basal layer of the heterostracan
surprising given that neither hagfishes nor lampreys exoskeleton.
have been recovered from strata younger than Upper Aspidin exists in many forms, some of which may
0
Quaternary
1.8
Neogene
Tertiary
PLI
Conodonta
UMI
Cephalochordata
Tunicata
Petromyzontida
Myxinoidea
Jawed vertebrates
LIA
206
u
Triassic
SCY
248
ZEC
Permian
ROT
290
GZE
Pennsylvanian
KAS
MOS
Carboniferous
BSK
Heterostraci
SPK
Mississippian
Euphanerops
Osteostraci
Galeaspida
VIS
Pituriaspida
TOU
Loganellia
354
u
Devonian
m
Jamoytius
Anaspida
PRI 417
LUD
Silurian
WEN
Arandaspida
Eriptychius
LLY
Astraspis
443
ASH
Bala
Ordovician
CRD
Dyfed
LLO
LLN
ARG
TRE
490
MER
Vendian Cambrian
STD
CRF Fig. 17. Preferred phylogeny (grey) plotted against stratigraphic

543
EDI ranges (black). Numbers are in millions of years before present.
have different phylogenetic histories ; we have attributed to jawed vertebrates (Skiichthys ; Smith &
attempted an initial first-fold division into lamellar Sansom, 1997). Contrary to previous studies, our
and spongy forms as part of this analysis. Aspidin is analysis suggests that cellular bone evolved in-
traditionally interpreted as a form of acellular bone, dependently in jawed gnathostomes and osteo-
secondarily derived from cellular bone (Ørvig, 1951, stracans.
1965, 1989). However, Smith & Hall (1990) con- Maisey (1986) hypothesised that enamel tissues
curred with Denison’s (1967) interpretation of the were the first vertebrate hard tissues to arise, and the
middle layer in Astraspis as trabecular dentine, but distribution of characters in our analysis is super-
also sided with Halstead Tarlo’s (1964) view that ficially consistent with this proposal. However, the
early in vertebrate evolution aspidin and dentine distribution of enamel and enamel-like tissues
were hardly distinguishable. Denison (1967) also throughout the cladogram of relationships does not
interpreted the middle and basal layers of Eriptychius, appear to corroborate the hypothesis that enamel is
and the basal layer of Astraspis, as aspidin, and he a synapomorphy of vertebrates (Smith, 1995).
was able to distinguish dentine from aspidin on the Tissues which possess the structural and devel-
presence of odontoblast cell-processes incorporated opmental characteristics of true (monotypic) enamel
into the matrix of dentine (the middle layer) in are found in conodonts, Eriptychius and jawed
Astraspis ; Denison (1967) made no attempt to draw gnathostomes (sarcopterygians and actino-
homology between aspidin and any other vertebrate pterygians ; Smith, 1989, 1992). Enameloid has been
hard tissue, although he rejected interpretations of described from Astraspis (Denison, 1967 ; Reif, 1979 ;
bone and dentine. Smith & Hall (1990 : p.302) cited Dzik, 1986 ; Smith et al., 1995) but its microstructure
‘ the view that dentine is primitive and dentine- cannot be compared to the enameloid of sharks and
related tissues (aspidin\cementum) formed in close actinopterygians (cf. Smith, Sansom & Smith, 1995).
sequence, both in development, topographically and Other records of enameloid in the dermal skeleton of
in time, and in evolution ’. This view is based upon jawless vertebrates (e.g. osteostracans Stensio$ , 1927 ;
the work of Palmer & Lumsden (1987) who, amongst Belles-Isles, 1989 ; galeaspids Janvier, 1990 ; Zhu &
others (Ten Cate & Mills, 1972 ; Ten Cate, 1975 ; Janvier, 1998) mistake the distinction between
Osborn, 1984 ; Osborn & Price, 1988), distinguished hypermineralised dentine and true enameloid ; there
skeletogenic and odontogenic components of the is no evidence for a basal lamina between the
vertebrate dermal skeleton, differentiating from the putative enameloid and the underlying ectomesen-
dental papilla early in odontogenesis. It is not clear chymally derived tissue.
how Smith & Hall (1990) interpreted the middle Conodont elements also incorporate (usually) an
layer in Astraspis or Eriptychius, but they returned to atubular dentine (this is not evident from the distri-
Ørvig’s (1951, 1965, 1989) interpretation of the bution of characters on Fig. 14 A because atubular
basal layer of aspidin in various jawless vertebrate dentine is not found in any of the other vertebrates in-
groups as a homologue of bone of attachment, albeit cluded in the analysis, and so could not help resolve
acellular, and an odontogenic derivative. Accepting relationships). The systematic position of conodonts
this interpretation of lamellar aspidin (alone), our and the distribution of dentine types over the
phylogenetic analysis suggests that acellular bone is cladogram would appear to indicate that atubular
more primitive than cellular bone, concurring with dentine is more primitive than tubular dentines.
Maisey (1988) but contrary to Ørvig (1951, 1965, However, Huysseune & Sire (1998) have suggested
1989), Smith & Hall (1990), Smith (1991) and that atubular dentines result from spatially confined
Sansom et al. (1992). In the absence of a phylogenetic development. Considering the small size of conodont
framework, Ørvig (1951, 1965, 1989) argued that elements, and the presence of tubular dentines in
cellular bone was primitive for vertebrates ; in all only the largest of elements (mesodentine and,
instances of acellular bone amongst jawed possibly, orthodentine ; Sansom et al., 1994 b), our
vertebrates it could be demonstrated unequivocally evidence would support this conclusion. Never-
that cellular bone was primitive (Schaeffer, 1977 ; theless, the presence of dentine in conodont elements
Parenti, 1986). Smith & Hall (1990) and Smith indicates that dentine is at least as primitive as
(1991) argued that cellular and acellular bone are of enamel, and the presence of a dentine in the absence
equal antiquity because they are both found at the of bone or a bone-like tissue indicates that dentine
same stratigraphic level. However, stratigraphy is no appeared before bone. Progressing from the root of
basis for establishing polarity, and the oldest skeletal the cladogram, orthodentine is first found in
remains to include cellular bone have since been pteraspidomorphs. Mesodentine unites Loganellia
with its sister-group, although Eriptychius plus jawed canalised and thus was open to a great deal of
vertebrates are distinguished by the possession of experimentation through natural selection. This
orthodentine. The random pattern of dentine types equates well with the view of phylogenetic reduction
apparent from the topology of relationships in our in histological diversity in, for example, conodonts,
most-parsimonious tree suggests that the structural heterostracans (Halstead, 1987) and vertebrates in
divisions between dentine types may be artificial, general (Sansom et al., 1994b ; Smith et al., 1996).
resulting from epigenetic factors which are incon- Obviously, it is not possible to test these hypotheses
gruent with phylogeny. directly because the animals in which the devel-
The random distribution of dentine structural opmental system was established have long been
types brings into focus the random distribution of extinct and there are no closely related living models.
hard tissues in general ; this is not a new observation. Nevertheless, the skeleton is the one organ system in
The random distribution of histological characters which a record of development is potentially pre-
has generally been accounted for by the plasticity of served in the fossil record and careful examination of
the odontode (Schaeffer, 1977). Conversely, the pattern of development of the exoskeleton
Halstead (1982) saw no such pattern, placing all his topologically and across different taxonomic levels,
faith in the a priori assumed importance of his- may yet provide a test of such hypotheses. Fur-
tological (plus other) characters to reconstruct thermore, a shift in focus from the study of
phylogeny. Schaeffer (1977) and Smith & Hall odontogenesis from traditional model animals with
(1990,1993) established the now prevalent view that their inherent problems (Bolker, 1995) to taxa which
the odontode is ‘ a single, modifiable, morphogenetic are phylogenetically more critical to this problem
system ’ such that any of the component tissues (e.g. Polypterus ; to some extent this shift in focus
(enamel, enameloid, dentine, bone, cartilage) could has begun – see Huysseune & Sire, 1998) might
be absent or present in combination with any other, provide the basis for elegant experiments to test the
and Smith & Hall (1993 : p. 395) have suggested coherence of the odontode model.
that the random pattern may result from the effect of
heterochrony upon the odontode differentiation pro-
gram. The role of heterochronic processes in the V. CONODONT AFFINITY
development of enamel versus enameloid was estab-
lished by Shellis & Miles (1974 ; but see also Smith, ‘ It is suggested that the present state of knowledge does not
1989, 1992, 1995). Evidence for loss and subsequent justify a final conclusion as to the affinities of the conodonts,
reappearance of other odontogenic tissues is based although they appear to represent an extinct group of either
upon heterotypic tissue recombinations in which a worm-like creatures or primitive vertebrates ’ (Rhodes,
taxon which is presumed, through lack of expression, 1954 : p. 450).
to have lost the ability to produce a tissue during
phylogeny can be demonstrated experimentally to Although Rhodes (1954) vacillated about the
have retained the potential. The obvious example is affinities of conodonts it is clear that he favoured a
the experiment conducted by Kollar & Fisher (1980) relationship to the annelids. He expressed reser-
in which avian mandibular epithelium was demon- vations about an affinity with vertebrates, which
strated to retain a capability for ameloblast differen- appeared ‘ to rest upon the answer to the question
tiation and enamel synthesis, when recombined with ‘‘ Must the chemical composition of the conodonts
murine dental papilla ectomesenchyme (see Kollar, (calcium phosphate) be interpreted as evidence of a
1998, for a discussion on the state of the debate over vertebrate origin, or is it possible for worms to secrete
this experiment). However, understanding of the such a substance internally ?’’ ’ (Rhodes, 1954 : pp.
possible mechanisms is based on a highly generalised 448–449). We now know that annelids are capable of
summary of experiments based on very divergent biomineralisation using calcium phosphate (e.g.
taxa. Moss (1964) argued that ‘ the earliest ossified Lowenstam, 1981), but no one any longer supports
vertebrates possessed the intrinsic capacity to pro- the hypothesis that conodonts are annelids.
duce the entire spectrum of vertebrate skeletal Our phylogenetic analysis provides unequivocal
tissues’ ; in effect that the developmental program of support for the hypothesis that conodonts are
odonto\skeletogenesis (now perceived to be two vertebrates, and not only in the widely used sense of
exoskeletal subsystems which arise from a common vertebrates l craniates (e.g. Donoghue et al., 1998),
cell population ; Palmer & Lumsden, 1987 ; Hall, but also in the sense used by Janvier (1981, 1993,
1992 ; Smith & Hall, 1990, 1993) had not been 1996 a, b). Furthermore, our analysis suggests that
A crown-group Homology of conodont and vertebrate hard tissues

total-group would appear to be supported by congruence.
TØ
Within the constraint that conodonts are vertebrates
the conclusion that conodont hard tissues are an
independent experiment, acquired in parallel with
the common ancestor of the Pteraspidomorpha and
all other vertebrates, appears unlikely. Thus, hy-
potheses of conodont affinity which argue that
conodont hard tissues are autapomorphic, e.g. Dzik
(1995), Kemp & Nicoll (1995 a) and Pridmore et al.
stem-group (1997 ; see also Peterson 1994), are (respectively) 13,
10 and six steps longer than the optimal phylogenetic
crown-group position for conodonts, based upon our data set.
Gnathostomata Nevertheless, the acquisition of a mineralised skel-
eton concomitant with a highly specialised feeding
B total-group apparatus does appear to occur very suddenly.
Gnathostomata However, if the ancestors of conodonts (in our sense)
lie amongst the paraconodonts, there may be
stem-group evidence that this group of characters did not appear
Gnathostomata all at once ; paraconodonts may prove to be sclerites
composed solely from dentine (cf. Bengtson, 1976)
and the origin of (eu)conodonts may be signalled by
the origin of enamel which, according to our
preferred phylogeny must have appeared more than
once (Fig. 14). However, patterning in the growth of
paraconodont elements appears to suggest that they
too grew appositionally, although an epithelial
Fig. 18. Stem, crown and total group classification. (A) component is not present, or at least not preserved.
Theory. (B) Practice. Providing the homology between paraconodont
elements and euconodont basal bodies is correct, the
conodonts are more derived than lampreys, and in pattern of growth requires an epithelial contribution
accordance with Jefferies ’ (1979) criteria for recog- to have been present, a possible candidate being
nising stem and crown groups (Fig. 18 A, B), keratin, which has a very low preservation potential
conodonts are stem-Gnathostomata. Although (Davis & Briggs, 1995 ; Davis, in press).
crown and total groups can be given separate names Following Schmidt & Mu$ ller (1964) and Priddle
(de Queiroz & Gauthier, 1992), this approach results (1974), Krejsa et al. (1990 a, b) have argued that
in an unnecessarily expanded classification scheme hagfish toothlets and conodont elements are hom-
and in one of the groups (stem-group) being ologous, although their hypothesis has here been
paraphyletic (unless that group is represented by one rejected because the primary observations are clearly
species only, in which case the need for a higher inconsistent with the pattern of morphogenesis
group name is unnecessary). Generally, only the exhibited by conodont elements (Szaniawski &
total group is recognised by formal Linnean rank Bengtson, 1993 ; Smith et al., 1996 ; Donoghue,
(Patterson, 1993 ; Smith, 1994). Thus, conodonts 1998). However, recent reappraisal of reports on the
belong to the Gnathostomata ; they are gnatho- presence of enamel-like antigens in the pokal cell
stomes, albeit without jaws (Fig. 18 B). cone of hagfish toothlets (Slavkin et al., 1983 ; see
The results of this analysis indicate that conodonts Slavkin & Dieckwisch, 1996, 1997), together with
lie in a position between the living jawless verte- analysis of toothlet microstructure (Dieckwisch &
brates, which lack a mineralised exoskeleton, and Vahadi, 1997), require that we look at this hy-
the extinct groups of jawless vertebrates which pothesis anew. Slavkin & Dieckwisch (1996, 1997)
possessed a mineralised exoskeleton in the form of a have demonstrated the presence of amelogenins not
dermal armour. Recodings in the experimental only in the enigmatic pokal cells, but also within the
section of this paper demonstrate that the affinity of keratin itself, with particular concentrations in the
conodonts does not rest upon histological characters. near-surface layers of the toothlets (but see Girondot,
Delgado & Laurin, 1998). Dieckwisch & Vahadi distinct in a multielement context ’. These values
(1997) also determined that the toothlets have a high should be taken as a highly conservative estimate
calcium to phosphate ratio and even found evidence (Sweet, 1988 : p. 131) ; even so, conodonts comprise
of keratin mineralisation. Krejsa et al. (1990 b) almost 65 % of all known jawless vertebrates, and
speculated that, were keratin heavily mineralised, it 2.8 % of total vertebrate diversity, living and extinct
would resemble the lamellar crown tissue of cono- (by comparison with figures in Nelson, 1994).
dont elements (although see Slavkin & Dieckwisch, Despite this, and despite the importance of cono-
1996). However, we now know of many instances of donts in interpretations of early vertebrate evolution,
mineralised keratin (T. G. H. Dieckwisch, personal we remain ignorant of all but the most fundamental
communication, 1998) and these are not comparable aspects of conodont palaeobiology and evolution.
histologically to conodont hard tissues. Nevertheless, Below, we outline some of the major unanswered
the possibility that hagfishes (and possibly lampreys) questions and outstanding problems.
express the plesiomorphic condition for the ver-
tebrate dermal skeleton must be considered more
(2) The origin of the Conodonta
seriously.
Smith & Hall (1990) argued that conodont Debate over the affinity of the group has reached a
elements and hagfish toothlets were independent state of maturity and will progress further only with
experiments with skeletal biomineralisation and discovery of unexpected new anatomical data.
Young, Karatajute-Talimaa & Smith (1996) con- However, as we signalled with a caveat over the
sidered the conodont skeleton to be an independent monophyly of the Conodonta at the beginning of this
specialisation which diverged from other vertebrates paper, we remain unaware of the direct ancestry of
soon after the establishment of the vertebrate the group. Little is known regarding the apparatus
skeleton. Indeed, Smith & Coates (1998) have architecture of the early Teridontus and Proconodontus
argued that the conodont skeleton may be an entirely lineages ; the overall similarity in growth patterns
distinct experiment in oral skeletal structures. How- and histology shown by these two putative clades
ever, available evidence suggests that the conodont suggests that they are closely related, but the
skeleton is the earliest expression of skeletal bio- question remains, how closely ? Resolution of this is
mineralisation amongst vertebrates, and that an most likely to rest with the unravelling of the
oropharyngeal skeleton\dentition is plesiomorphic relationships between these lineages and the para-
for the clade. Whether conodont elements are conodonts (following Bengtson, 1976). However, the
homologous to true teeth is a moot point. They paucity of diagnostic features in these groups means
clearly do not occur topologically within a jaw, but that determination of these relationships will be very
they do exhibit evidence of replacement (Donoghue, difficult. The paraphyletic paraconodonts are dis-
1998 ; Donoghue & Purnell, 1999 a) ; they also grew tinguished from euconodonts by the absence of a
in a manner directly comparable to teeth (Schmidt character, i.e. a crown composed from a discrete
& Mu$ ller, 1964 ; Mu$ ller & Nogami, 1971 ; Sansom, tissue type. On current knowledge, the only potential
1996 ; Donoghue, 1998), functioned like teeth synapomorphy of para- and (primitive) euconodonts
(Purnell, 1995 ; Donoghue & Purnell, 1999 b), and is the possession of phosphatic cone-shaped sclerites
even look like teeth (Pander, 1856 ; Jeppsson, 1979). which grow by episodic secretion of successive layers
that only partially encapsulate preceding layers.
Sclerites from many other groups also exhibit a
VI. DIRECTIONS FOR FUTURE RESEARCH
grade of organisation which parallels this, and so we
need to assess whether there is any evidence that
(1) General
para- and euconodonts should be linked at all.
Recent years have seen a dramatic increase in our Cambrian strata yield a wide variety of micro-
understanding of many aspects of conodont palaeo- scopic phosphatic sclerites, several of which possess
biology, leading to the results presented in this histological attributes that might suggest comparison
paper. However, the apparent magnitude of these with microvertebrate remains. For example, sclerites
advances simply reflects how little was previously referred to the genera Hadimopanella and Milaculum
known, and there is still much to do. Sweet (1988 : p. have in the past been allied to the jawless vertebrates
189) estimated the number of named conodont on the basis of tissues that resemble enamel and
species at ‘ nearly 5000 ’, although he considered only dentine (see e.g. Dzik, 1986). However, Hinz et al.
1446 species and 246 genera to be ‘ reasonably (1990) and Mu$ ller & Hinz-Schallreuter (1993) have
demonstrated unequivocally that these micro- progress towards a truly hierarchical suprageneric
remains belong to a group of hitherto poorly known system. The most recent classifications (Sweet, 1988 ;
Palaeozoic annelids, the Palaeoscolecida. This dis- Dzik, 1991) are dominated by groupings that are
covery highlights the problems of using histological unashamedly para- and polyphyletic, as recognised
data alone to determine relationships, without the by Aldridge & Smith (1993), who nevertheless
benefit of a broader anatomical context. adopted a modified version of the Sweet (1988)
One way to demonstrate a link between para- scheme. It is now appropriate to undertake more
conodont and euconodont elements would be to rigorous analyses of relationships within the cono-
document a series of intermediate forms. This donts to produce a classification based on mono-
approach was adopted by Szaniawski & Bengtson phyletic groups ; the implementation of cladistic
(1993), who described specimens that resemble methodology should be aided by the fact that current
paraconodonts in early ontogeny but appear to conodont taxonomy is largely based on topological
acquire a crown in late ontogeny. Histogenetic homology. There is some urgency here, because until
investigation of such specimens should show that conodont phylogeny has been unravelled, we cannot
mature sclerites display no correlation between advance our understanding of the evolution of
incremental layers in the base and crown in the element histology, growth and function.
portion of the growth record representing early
ontogeny ; unfortunately, this is not clearly demon-
(4) Histology
strated in any of their material.
More recently, Mu$ ller & Hinz-Schallreuter The vertebrate skeleton is the system of choice for
(1998) have described the histology of Wester- studying organ development in biological and
gaardodina sp., clearly a paraconodont in its grade of biomedical research (Hall, 1992). Its evolutionary
histological organisation, which displays a calco- history is also more easily understood than that of
spheritic microstructure directly comparable with any other organ system, because it is mineralised and
the basal body of the euconodont Cordylodus (e.g. therefore readily fossilised. The results of our
Mu$ ller & Nogami, 1971 ; Sansom et al., 1992 ; phylogenetic analysis are congruent with the hy-
Aldridge & Donoghue, 1998). An even more pothesis that the vertebrate dermal skeletal system
persuasive link is evident in the histology of was established in the common ancestor of conodonts
Serratocambria minuta Mu$ ller & Hinz, also described and more-derived vertebrates after separation of the
by Mu$ ller & Hinz-Schallreuter (1998), in which the lamprey lineage. Conodonts are therefore an ideal
pattern of growth is comparable with the basal body model group in which to study the evolution of the
of much younger, highly derived euconodont taxa skeletal developmental system : the conodont clade
(e.g. Carniodus in Donoghue, 1998). persisted for some 300 million years, showing low
It remains an open question whether acquisition disparity and high diversity, and the elements exhibit
of a phosphatic crown unites the Teridontus and a very wide range of hard tissue and morphogenetic
Proconodontus lineages, or whether the crown evolved diversity.
more than once. This can only be resolved by The existing histological database shows that the
determining relationships between the taxa of para- odontogenic system in conodonts underwent con-
conodonts and primitive euconodonts and thereby siderable change during the phylogeny of the group
testing the traditional grade-based groupings. (Smith et al., 1996 ; Donoghue, 1998). However, only
a small percentage of conodont taxa have been
investigated histologically. Of the seven orders of
(3) Euconodont phylogeny
conodonts proposed by Sweet (1988), only repre-
The adoption of a wholly artificial form-taxonomy sentatives of the Ozarkodinida, the most derived
for elements during the first century of conodont group, have been studied in any detail (Donoghue,
research meant that it was impossible to construct a 1998), and the results are probably not represen-
meaningful scheme of high level classification. With tative of the conodont clade as a whole. Published
the rise of multielement taxonomy (e. g. Huckreide, histological data from the remaining six orders
1958 ; Walliser, 1964 ; Bergstro$ m & Sweet, 1966 ; (Proconodontida, Belodellida, Protopandero-
Webers, 1966), however, the basis was provided for dontida, Panderodontida, Prioniodontida, Prionio-
the development of a homology-based scheme dinida) are largely restricted to abstract accounts
(Sweet & Scho$ nlaub, 1975 ; Sweet, 1981, 1988 ; (summarised in Donoghue, 1998). Limited research
Purnell et al., 2000), although there has been little on the Panderodontida, an equally derived but
divergent group of conodonts, has revealed an to little more than informed conjecture. Detailed
entirely different mode of growth from that shown investigations are now required of diverse, well-
by ozarkodinids (Barnes, Sass & Poplawski, 1973), preserved taxa, set in a clear phylogenetic context.
and preliminary studies of the Prioniodontida In this way, it should become possible to establish
(Donoghue & Chauffe, 1999) have demonstrated the functional adaptations and constraints driving
the presence of a unique suite of hard tissue types. the morphological evolution of the euconodont
Most critically, histological data on primitive cono- clade(s).
donts are extremely limited.
VII. REVISED CLASSIFICATION
(5) Microevolution
As has been pointed out in other reviews of In our Linnean rank-free classification below we use
conodonts, the elements hold clear potential for a total group concept (see Fig. 18) in which taxa
microevolutionary studies. Although the useable forming a stem-lineage series leading to a crown
record of conodonts is limited exclusively to their group are classified as members of the total group.
teeth, these show clear sequential changes and often This is the most unambiguous way of specifying
occur in great numbers. There have been several phylogenetic relationships. We also use the
morphometric studies of conodont elements over the sequencing convention of Nelson (1972), the plesion
years (e.g. Rhodes, Williams & Robinson, 1973 ; rank as introduced by Patterson & Rosen (1977) and
Barnett, 1971 ; Croll, Aldridge & Harvey, 1982 ; the sedis mutabilis suffix introduced by Wiley (1979).
Croll & Aldridge, 1982 ; Murphy & Matti, 1982 ;
Murphy & Cebecioglu, 1986, 1987 ; Klapper & Craniata Linnaeus, 1758
Foster, 1986, 1993), but to date there have been few Myxiniformes Berg, 1940
serious attempts to use the conodont record as a Vertebrata Linnaeus, 1758
testing ground for hypotheses of evolutionary Petromyzontiformes Berg, 1940
patterns and processes. Gnathostomata Cope, 1889
plesion Conodonta Eichenberg, 1930
(6) Conodont element function plesion Pteraspidomorphi Goodrich, 1909
Astraspis
The documentation of microwear features on the
Heterostraci Lankester, 1868
surfaces of a variety of conodont elements (Purnell,
Arandaspidiformes Ritchie & Gilbert-
1995) has provided clear evidence that they per-
Tomlinson, 1977
formed a tooth function. Investigation of the
plesion unnamed group A
functional morphology of molarised elements of the
plesion Anaspida Traquair, 1899
Pennsylvanian conodont Idiognathodus (Donoghue &
Jamoytius
Purnell, 1999 b) has also shown that these occluded
Euphanerops
so precisely that no room would have been available
plesion Loganellia
for any intervening soft tissue cover. There is a need
Unnamed group B
to reconcile the appositional growth of these elements
Eriptychius
with their occlusive function, and this may be
Jawed vertebrates
explained by evidence of periodic soft tissue cover
plesion unnamed group C
(Donoghue & Purnell, 1999 a).
Osteostraci Lankester, 1868 sedis mutabilis
Although the functional morphology of some
Pituriaspida Young, 1991 sedis mutabilis
elements of a few taxa has now been interpreted in
Galeaspida Halstead, 1982 sedis mutabilis
some detail, very little is still known regarding
element and apparatus function in general. Coni-
form taxa are particularly poorly understood, the VIII. CONCLUSIONS
only current direct evidence arising from surface
scratches on an element of Drepanoistodus figured by (1) The presence of myomeres and a notochord in
Purnell (1995). In the more complex apparatuses, body fossils of euconodonts justifies interpretation of
studies have been largely directed at the posterior- the group in the context of a chordate bauplan.
most P elements, with limited data also available on (2) The significance of conodonts to our under-
"
the function of the P elements. Our knowledge of standing of chordate evolution cannot be assessed by
#
the functions of M and S elements currently amounts placing the group within an existing hypothesis of
phylogenetic relationships, based upon a priori permission of the Lethaia Foundation ; Figures 4 b, d, e
determined synapomorphy distribution. are reproduced with the permission of Macmillan
(3) Cladistic analyses show that conodonts are Magazines Ltd ; Figures 5 d, g, h are reproduced with the
vertebrates and that they are more derived than permission of the Geological Society of America ; Figures
either hagfishes or lampreys because they possess a 5 k, l are reproduced with the permission of the Paleonto-
logical Society. Donoghue was funded by an 1851
calcified dermal skeleton ; this phylogenetic position
Research Fellowship from The Royal Commission for the
does not rely on the nature of the mineralised tissues. Exhibition of 1851 and a NERC post-doctoral research
(4) The Conodonta are stem-group Gnatho- fellowship (GT5\99\ES\2).
stomata ; paraconodonts and euconodonts are the
most primitive members of the total-group Gnatho-
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XI. APPENDIX
(1) Character diagnostics
Character statistics for the tree presented in Fig. (7A). Guide to abbreviations : CI, consistency index, HI,
homoplasy index, RI, retention index, RC, rescaled consistency index.
Appendix.
Minimum Tree Maximum

Character Steps Steps Steps CI HI RI RC
1. 1 1 2 1.000 0.000 1.000 1.000

2. 1 1 2 1.000 0.000 1.000 1.000
3. 1 2 3 0.500 0.500 0.500 0.250
4. 2 3 3 0.667 0.333 0.000 0.000
5. 1 1 2 1.000 0.000 1.000 1.000
6. 2 2 3 1.000 0.000 1.000 1.000
Appendix. (cont.)

7. 1 1 2 1.000 0.000 1.000 1.000

8. 1 1 4 1.000 0.000 1.000 1.000
9. 1 1 1 1.000 0.000 0\0 0\0
10. 1 2 2 0.500 0.500 0.000 0.000
11. 1 2 2 0.500 0.500 0.000 0.000
12. 1 1 2 1.000 0.000 1.000 1.000
13. 1 1 2 1.000 0.000 1.000 1.000
14. 2 3 4 0.667 0.333 0.500 0.333
15. 1 2 2 0.500 0.500 0.000 0.000
16. 1 2 3 0.500 0.500 0.500 0.250
17. 2 4 5 0.500 0.500 0.333 0.167
18. 2 2 3 1.000 0.000 1.000 1.000
19. 1 1 3 1.000 0.000 1.000 1.000
20. 3 3 4 1.000 0.000 1.000 1.000
21. 1 1 4 1.000 0.000 1.000 1.000
22. 1 1 3 1.000 0.000 1.000 1.000
23. 1 1 3 1.000 0.000 1.000 1.000
24. 1 1 2 1.000 0.000 1.000 1.000
25. 2 3 4 0.667 0.333 0.500 0.333
26. 2 3 7 0.667 0.333 0.800 0.533
27. 1 2 3 0.500 0.500 0.500 0.250
28. 1 1 2 1.000 0.000 1.000 1.000
29. 1 3 7 0.333 0.667 0.667 0.222
30. 1 3 6 0.333 0.667 0.600 0.200
31. 1 1 2 1.000 0.000 1.000 1.000
32. 1 2 3 0.500 0.500 0.500 0.250
33. 1 2 2 0.500 0.500 0.000 0.000
34. 1 1 3 1.000 0.000 1.000 1.000
35. 1 2 3 0.500 0.500 0.500 0.250
36. 1 2 2 0.500 0.500 0.000 0.000
37. 1 1 3 1.000 0.000 1.000 1.000
38. 1 1 3 1.000 0.000 1.000 1.000
39. 1 1 2 1.000 0.000 1.000 1.000
40. 1 3 3 0.333 0.667 0.000 0.000
41. 1 1 3 1.000 0.000 1.000 1.000
42. 1 1 2 1.000 0.000 1.000 1.000
43. 1 3 4 0.333 0.667 0.333 0.111
44. 1 2 4 0.500 0.500 0.667 0.333
45. 1 3 5 0.333 0.667 0.500 0.167
46. 1 1 2 1.000 0.000 1.000 1.000
47. 1 2 3 0.500 0.500 0.500 0.250
48. 1 2 2 0.500 0.500 0.000 0.000
49. 2 3 6 0.667 0.333 0.750 0.500
50. 1 2 3 0.500 0.500 0.500 0.250
51. 1 1 2 1.000 0.000 1.000 1.000
52. 1 2 3 0.500 0.500 0.500 0.250
53. 1 2 2 0.500 0.500 0.000 0.000
54. 1 2 3 0.500 0.500 0.500 0.250
55. 1 1 3 1.000 0.000 1.000 1.000
56. 1 2 2 0.500 0.500 0.000 0.000
57. 1 1 2 1.000 0.000 1.000 1.000
58. 1 1 3 1.000 0.000 1.000 1.000
Appendix. (cont.)

59. 1 1 3 1.000 0.000 1.000 1.000

60. 1 1 3 1.000 0.000 1.000 1.000
61. 1 2 3 0.500 0.500 0.500 0.250
62. 1 1 5 1.000 0.000 1.000 1.000
63. 1 1 3 1.000 0.000 1.000 1.000
64. 1 2 5 0.500 0.500 0.750 0.375
65. 1 1 3 1.000 0.000 1.000 1.000
66. 1 2 3 0.500 0.500 0.500 0.250
67. 1 3 6 0.333 0.667 0.600 0.200
68. 1 2 2 0.500 0.500 0.000 0.000
69. 2 5 6 0.400 0.600 0.250 0.100
70. 2 3 4 0.667 0.333 0.500 0.333
71. 1 3 6 0.333 0.667 0.600 0.200
72. 1 1 2 1.000 0.000 1.000 1.000
73. 2 3 6 0.667 0.333 0.750 0.500
74. 2 3 9 0.667 0.333 0.857 0.571
75. 1 1 3 1.000 0.000 1.000 1.000
76. 1 3 5 0.333 0.667 0.500 0.167
77. 1 3 3 0.333 0.667 0.000 0.000
78. 2 4 9 0.500 0.500 0.714 0.357
79. 1 1 2 1.000 0.000 1.000 1.000
80. 1 1 2 1.000 0.000 1.000 1.000
81. 1 2 4 0.500 0.500 0.667 0.333
82. 1 3 3 0.333 0.667 0.000 0.000
83. 1 1 2 1.000 0.000 1.000 1.000
84. 1 1 2 1.000 0.000 1.000 1.000
85. 1 1 2 1.000 0.000 1.000 1.000
86. 1 1 2 1.000 0.000 1.000 1.000
87. 1 1 2 1.000 0.000 1.000 1.000
88. 1 1 2 1.000 0.000 1.000 1.000
89. 1 1 2 1.000 0.000 1.000 1.000
90. 1 1 2 1.000 0.000 1.000 1.000
91. 1 1 2 1.000 0.000 1.000 1.000
92. 1 1 2 1.000 0.000 1.000 1.000
93. 1 1 2 1.000 0.000 1.000 1.000
94. 1 1 2 1.000 0.000 1.000 1.000
95. 1 2 2 0.500 0.500 0.000 0.000
96. 1 1 2 1.000 0.000 1.000 1.000
97. 1 1 2 1.000 0.000 1.000 1.000
98. 1 1 2 1.000 0.000 1.000 1.000
99. 1 1 2 1.000 0.000 1.000 1.000
100. 1 2 2 0.500 0.500 0.000 0.000
101. 1 2 2 0.500 0.500 0.000 0.000
102. 1 1 2 1.000 0.000 1.000 1.000
103. 1 2 2 0.500 0.500 0.000 0.000

Conodont Affinity and Chordate Phylogeny

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Biol. Rev. (2000), 75, pp.

191–251 Printed in the United Kingdom # Cambridge Philosophical Society 191

Conodont affinity and chordate phylogeny

(Received 4 June 1999 ; revised 29 November 1999 ; accepted 29 November 1999)

I. Introduction ............................................................................................................................ 192

(4) Hard tissue homologies ..................................................................................................... 202

I. INTRODUCTION Klussendorf, 1985 a, b ; Aldridge et al., 1993 ; Gabbott,

Fig. 3. For legend see opposite.

Fig. 5. For legend see opposite.

Fig. 14 a. For legend see opposite.

Conodont affinity and chordate phylogeny

26 a GAIN: neural crest, brain, divided pituitary,

CRF Fig. 17. Preferred phylogeny (grey) plotted against stratigraphic

A crown-group Homology of conodont and vertebrate hard tissues

Minimum Tree Maximum

1. 1 1 2 1.000 0.000 1.000 1.000

Minimum Tree Maximum

7. 1 1 2 1.000 0.000 1.000 1.000

Minimum Tree Maximum

59. 1 1 3 1.000 0.000 1.000 1.000

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