;oolo,qiml ,Joumnl of the /,innpan So&& (1988), 94: 233-258. \Vith 16 figures
A new camuropiscid arthrodire (Pisces:
Placodermi) from Gogo, Western Australia
JOHN LONG F.L.S.
Geology Department, Uniuersily of Tasmania, P.O. Box 252 C,
Hobart, Tasmania, Australia 7001
Receir'ed J u l p 1987, accepted,fi,r publication December I987
A new camuropiscid arthrodire, Latocamurus coulthardi gen. et sp. nov., is described
from the Upper Devonian Gogo Formation, Western Australia. Latocamurus, known from two
complete specimens. i s recognized as a carnuropiscid by its narrow, spindle-shaped armour, dccp
postnasal plates participating in the orbits, prcorbital plates which meet mesially, cheek unit firmly
sutured to skull roof, posterior cheek plates tightly interconnected and much reduced, and the
robust durophagous dentition. I t i s characlerizcd by its downturned siiout, broad, flat rostral plate,
and narrow, deep parasphenoid. I t is placed phyletically as the plesiomorphic sister taxon to all
other caniuropiscids which are more derived in having, inter a h , a n antrrior lateral plate which
arireriorly contacts the anterior vcritrolateral plate and pointed rostral plates. T h e family
Camuropiscidae Dennis & Miles 1979h i s redefined to incorporate features of the new genus.
Carnuropiscids arid Incisoscutnm are closely related by fcatures of thc postnasal plate and cheek.
KEY \\'ORDS:
anatomy
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Devonian
phylogeny.
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Australia
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camuropiscid
arthrodire
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new species
ethmoid
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CONTEN13
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Introduction .
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Systematic palaeonrology .
Family Carnuropiscidar Dennis & Miles 1979b.
Latocamurus gen. nov . . . . . .
Latocamurus coulthardi sp. nov . . .
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Parasphenoid.
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Ethmoid ossification .
Relationships of Latocamuruj gen. no\
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Ackriowlcdgcmrnts
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IN'I'KODUC'IION
Since the first collections of fossil fishes were made from Gogo in the 1960s a
series of important papers has been published dealing with the placoderms
(Miles, 1971; Miles & Dennis, 1979; Dennis & Miles, 1979a, b, 1980,
1981, 1982, 1983; Dennis-Bryan, 1987; Miles & Young, 1977; Young, 1984;
0024-4082/88/110233
+ 26 $03.00/0
233
1988 T h e Liiineaii Society of L.ondon
231
J . LONG
Fore)- & Gardiner, 1986), lungfishes (Miles, 1977), palaeoniscoids (Gardiner,
1913, 1984; Gardiner & Bartram, 1977) and the crossopterygians (Long, 1985).
Still au.aitirig publication are works dealing with Onychodus (by Dr S . M.
Atidrews), the four or so new species of coccosteids and Bothridepis (>by Dr
Dennis Bryan, Prof. B. Gardiner and Dr R . S. Miles). All of these papers have
t o some extent made an impact on our views of early vertebrate evolution, and
some controversy has arisen over the problem of tetrapod ancestry (Rosen,
Forcy, Gardiner & Patterson, 1981) based on new anatomical data from Gogo
lilrs, 1977). This fauna is regarded as one of the world’s most important
assemblages of‘ Devonian fishes because of the extraordinary preservation of‘
anatomical detail afforded by the acetic acid preparation of the specimens. All
of the above works are the result of the first two collections (1963, 1967), most ot‘
vvhich arc held in the British h4useum (Natural History).
Dennis-Bryan (1987: 2) states that the 1967 tripartite expedition comprised
Dr C. H. C. Brunton (British Museum), Dr W. D I. Rolfe (Hunterian
Museum) and Dr K. S. 1%‘.Campbell (Australian National University), but in
fact Campbell collected material on an independent trip later in the early 1970s,
a s did Gavin Young and Alex Ritchie. M r H . A. Toombs (British Museum) was
the expedition leader and the australian contingent consisted of George
Kendrick, Ken Butler and David Cliff all from the Western Australian
lluseum. It should be noted that Dr Curt Teichert, during his Canning Basin
fictld trips of the late 1930s to early 1940s was the first person to recognize
‘coccostean’ fish remains from the Manticoceras zone in limestone concretions
(Teichert, 1949), and not Dr Basil Balme as stated by Ritchie (1985). ‘Teichcrt
collected Gogo fishes which were reposited in the collections of the University of’
Ct‘estern Australia Geology Museum, and also found the skull of a new genus of’
dinichthyid from a Famennian horizon in the same region (Long, 1987~11.
Denis-Bryan (1987) states that the entire collecting area of the 1967 expeditioli
was within Gogo Station, whereas today many of the important Gogo localitics
arc situated on the adjacent Christmas Creek Station (Paddy’s Valley, Bugle C a p
and Teichert (Hills).Although Gogo Formation as mapped by Playford & Lowry
(1966) is exposed over a large area within Gogo and Christmas Creek Stations.
it is only in restricted regions where weathering has produced a significant
concentration of Gogo concretions that fish can be found in abundance.
This paper represents a new generation of work on the Gogo fauna resulting
from material found on the 1986 and 1987 Gogo expeditions sponsored b y thc
National Geographic Society. T h e new material, all of which shall be reposited i n
the Western Australian Museum, includes the new arthrodire described here a s
well as other new species; a complete skull of the osteolepiform Gogonnsus (Long,
1985); previously undescribed material of the tail and pelvic girdle of Holonemci
utestolli (Miles, 197 1); a complete specimen of the poorly known ptyctodontid
Chnpbellodus decipiens (Miles & Young, 1977); new information on previousl)
described arthrodires (Long, 1987b; 1988, and several skulls and hodirs of
lungfishes, including a new genus of chirodipterid and a new species ol‘
Holod$terus, which are being studied by Prof. K. Campbell and Dr R . Barwick
at the Australian National University.
As the previous Gogo arthrodires have been described in great detail, and thr.
new form does not differ significantly from the other camuropiscids in man).
points of anatomy, I shall leave the illustrations to show the shape and form of
N E b ’ AUSTRALIAN ARlHRODlRE:
235
these components, as adopted by Miles & Dennis (1979) and Dennis & Miles
(1979- 1983). T h e description will follow the numbered character scheme of
Miles & Dennis, but where morphological features are essentially the same as in
the other camuropiscids this will be noted and no further comments made. As
the material is uncrushed and three-dimensionally perfect, there is complete
confidence in the given restorations of the armour, and therefore only unusual
features of the new genus will be described. Measurements (Table 1 ) follow the
scheme adopted by Miles & Dennis (1979) and Dennis-Bryan (1987).
SYS’IEMA’I’IC PALAE:ON‘I’OI.OGY
Order Euarthrodira Gross 1932
Suborder Eubrachythoraci Gross 1932
Family Camuropiscidae Dennis & Miles 197913
Diagnosis
Small eubrachythoracid arthrodires which have the following derived
features: elongated, narrow body shape with large orbits; postnasal plates large,
excluding contact between the suborbital and preorbital plates; rostral plates
either broad and flat or tubular in presumed advanced species; preorbital plates
having mesial contact; suborbital plate firmly sutured to skull roof;
postsuborbital and submarginal reduced and tightly interconnected to each
other; dentition durophagous with thick, rectangular posterior superognathals;
postmarginal sensory-line canal not present, and main lateral line canal having
a ventral course o n the anterior dorsolateral plate.
Remarks
T h e above diagnosis has been amended from Dennis & Miles (197913: 298) to
exclude features of Carnurapiscis, Rolfosteus and Tubonasus which are not seen on
the new form. T h e Camuropiscidae are an endemic group of specialized
arthrodires known at present only from the Gogo Formation.
Referred taxa
Carnurapiscis concinnus, Carnurapiscis laidlawi (Dennis & Miles, 1979a), RolJosleus
canningensis, Tubonasus lennardensis (Dennis & Miles, 1979b), Latocamurus
coulthardi gen. et sp. nov.
Latocamurus gen. nov.
Etyma Logy
From the Latin latus, broad, and camurus, a flat nose, alluding to the shape of
the rostral plate.
Diagnosis
A camuropiscid arthrodire having a headshield with a breadth/length index
of c. 65, and orbits which are 4404 of the postpineal length of the headshield.
Headshield breadth equal to postpineal length of skull roof. T h e snout is
strongly downturned. The rostral plate is broader than long, with a weakly
convex anterior margin and four concave margins for contacting the postnasal
and preorbital plates. T h e suborbital plate has a deep and well rounded
236
,J. LONG
postorbital division, and the postsuborbital is larger than the submarginal which
is firmly sutured to the marginal plate. The anterior and posterior
superognathals have a central row of broad, rounded tubercles; the
irifixrognathal lacks teeth having only broad crushing surfaccs. ' l h e para-
NEW AUS'I'RALIAN AR'I'HRODIRE
237
E'igurc 2. Latocamurus coulthardi gen. et sp. nov. A. armour in latcral vicu (dusted wit11
ammonium chloride]; B, intcrprctation of k a t u r r s iuppcr .jaw toothplatcs addcd). \ V A M 86.9.699.
sphenoid is very narrow and deep in profile, with a crescentic ledge posterior to
the huccohypophysial foramina on the ventral surface. Trunkshield has a
breadth/length index of 46; the median dorsal plate has a sharply pointed
posterior margin, and the spinal plate is long, separating the anterior lateral
from contact with the anterior ventrolateral; anterior median ventral plate with
narrow tapering posterior lobe which has short contact with posterior median
ventral plate.
Remarks
Latocamurus is readily distinguished from the other camuropiscids by its
broader headshield with downturned snout, blunt rostra1 plate and long spinal
plate which excludes the anterior lateral plate from contacting the anterior
ventrolateral plate. It also differs in other minor features such as the shape of
the pineal and postnasal plates and in features of the parasphenoid. T h e
characters it shares with other camuropiscids are discussed in the phylogenetic
section at the end of this paper.
Type species
Latocamurus coulthardi sp. nov.
J . LONG
238
TABLE
1. hleasurcments ( m m ) of Latocamurus coulthardi gen. et sp.
nov. based on the schemr of Miles & Dennis (1979)
H h t ! pc
~
~~~
~~~~~~
~
~~
~~~~~~
'vY.111 86.9.699
.
Lciigtli (11' skull
Brcadtti of skull across postri-ol~itrr~d
aiigl~~s
Rrradtli o f skull across postrromcsial angler
Uqxh ( i f skull
I'rcpiIi(,;iI length
Lerigt ti d o r h i t
I.ellg'h of XI:
Leiigtli of l a t c r d articular fossa
1)cprh of Intrral articular fossa
.\iiglc I)ct~vr.cnaxis of 'trticular fiissa arid
d(irscilatera1 surfacc or skull
l x n g t h i i E r.licck
I.c~tgrhof postortiital di\ iairiii ol'cticck
I.cllgth of If:
1,crigth 01biting dilisioii of Ify
Length of t~.urrksliicld
Hi-cadtti d ' 1 r u n k shield
Dcprh 01' t i n i n k shield
Ro~ri-oc;iudalIcngth of I l m k i i i - i i i o u r
I.cngtl1 of ["'tol-al lcncstr'l
Lclrgttl of S1U
16
31
26
21 4
I3.j
1 15
13 7
I5*
3.7
L.
2
2.6
(.
(.
35*
1%5*
34
I6
88
i. 4 2
23*
17.5
1.8
30
31) ,j
Ib.5
12
63.5
3I
:i:i
17
9T
37
-.
'I() 5
Rrc'ltltll 01 \ll)
lrcllgtll 0"
A n g k l)rr\iecii Sp a n d rostr(icaiidaI
axis of iirmour
21.5
13
19
9
0
1"~Ilgrhol'.\\.l.
I.ciigth of spinal tlivisioir of h1.L
37
0
28
22.5
Ili.7
sp
Latocamurus coulthardi sp. nov.
(Figs 1-15)
I.:1 I'?rlolo<gJ
I n honour of M r ,Jim Coulthard, district rnanager of Christmas Creek and
Gogo Stations, who kindly permitted us to collect specimens.
.llaleTial
Holotype, \VAM 86.9.670 (Figs 4, 5 ) , a complete armour, partly articulated,
prepared out ns two halves by the resin transfer method (Rixon, 1976). t V A M
86.9.699, complete specimen.
Luccilz(v and horizon
Holotype collccted by the author on 17/8/1986, from approxitnatel) 100 m
south-west of 'Stromatoporoid Camp' (close to locality no. 20 of Miles, 1971,
fig. 1). WAM 86.9.699 collected by the author from the north-eastern side of
Paddy's Vdlley in June 1987. Late Devonian (Frasnian) Gogo Formation.
NE\V AUSTRALIAN ARI‘HKODIKE
Figure 3 Latocamurus coulthardi gen. et sp. nov., tiendshield
1 cinbrrd w i t h ammonium chloridc)
111
\entrdl
239
tic\\,
\Z 411 86 9 63’)
Descriplion
‘l’he general shape of the armour (Figs 1, 2), like that of the other
camuropiscids, is elongated and narrow, but not as streamlined as in presumed
advanced members of the family. Proportions of the armour (Table 1) can be
directly compared with the other camuropiscids.
‘The headshield (Figs 1-3) is narrow (breadth/length index 65) with large
orbits which are approximately 4 4 O , of the postpineal skull length in diameter,
similar in size to that of Cumuropiscis concinnus (4.27,).The rostra1 plate is variable
in the two specimens (Figs 1, 7B) but is consistent in its broad shape with a
weakly convex anterior margin and large overlap areas for the postnasal and
preorbital plates. I’he pineal is cruciform in the holotype (Fig. 7A), but what is
preserved of it in WAM 86.9.699 (Fig. 1) is similar to that of Cumuropiscis. T h e
preorbitals have a short area of mesial contact just anterior to the pineal plate,
and a deep descending lamina with large overlap surfaces for the postnasal
plates. They are arched anteriorly. l’he postnasal plates (Figs 3, 4, 7A, 8, 15A)
are very large and deep, and participated in the anterior margin of the orbit.
They have separated contact margins for the suborbital (oa.SO) and preorbital
ioa.PRO: plates (Fig. 81, indicating that these plates did not contact each other
behind the postnasal i n the articulated armour as occurs in Incisosculum (Dennis
8r hiiles, 1981). There is a horizontal lamina of bone projecting at a right angle
to the surface of the postnasal for overlap of the rostra1 plate (oa.R, Fig. 8 ) . In
the articulated armour the postnasals are orientated vertically to enclose a
box-like space between themselves and the skull roof (Figs 3, 1.5). Thc.
postorbital. marginal and paranuchal pIates bear n o special features; their
outlines can be seen from the figures. T h e central plates show clearly the middle
and posterior pit-lines. The postmarginal plate is entirely tucked into thc
subobstantic area of the headshield (soa, Fig. Z), and lacks an) dermal
ornamentation on its lateral surface. T h e ventral aspect of the nuchal platc. has
a iveak nuchal thickening with paired infranuchal pits (pt.u, Fig. 3 i , arid
numerous nutritive foramina in its central region. The triangular dcprcssion
noted on the visceral surface of the marginal plate of Camuropiscis b y Dennis 8
nliles (1979a: fig. 16) is well developed in Latocamurus itri, Figs 3, 1 3 A ) . 'l'hcc
long quadrate (Qd, Fig. 13) does not stretch into this depression as suggested for
hE,\.\’AUS I RALIAX AR I HRODIKt
24 I
Figure
dorsal
Camuropiscis, but bends away below this region, which was probably occupied by
muscles which moved the cheek complex.
T h e dermal ornament is well illustrated (Figs 1 , 2, 6, 7, 9), consisting of
simple low tubercles which may form a concentric disposition on parts of the
ventral thoracic armour. It is not as pronounced as some of the high tubercles
found on the armour of Cumuropiscis (for example see Dennis & Miles, 1979a:
fig. 10B).
T h e following characters are now described with reference to the phylogenetic
scheme of Miles & Dennis (1979) to facilitate comparisons with other Gogo
arthrodires. Points of morphology identical with those on Cumuropiscis are noted
but not discussed further.
1. Great width of skull roof across posterolaterul angles. T h e posterolateral angles
are not seen in dorsal view (Fig. 1). The headshield has an overall broader
shape than Cumuropiscis due to the shorter rostra1 plate, and the postpineal
length of the skull, relative to its breadth is still marginally shorter (postpineal
242
J. LONG
Figurr b. Latocamurus coulfhardi gen. et sp. nov. SESI photographs of rhr drrnial omanirnt
lrom t h r lrli pnstrrioi- dorsolatrral plate. Iiolotypc. A, B, Ohlique virws A. x 120; U, x 60 (:.
Prrpcndicular ti) tuticrclrs x 60'1.
length/breadth skull roof being 100 for Latocnmurus, 78 Camuropisrzs, 87-88 for
Tubonasus and Rolfsteus; 140 for Hanytoomhsia).
2. Two superognathals. As for other arthrodires these are paired anterior and
posterior elements, both with dorsal processes developed (Figs 7A, 8, 10, 13,.
IVhen articulated with the autopalatine and ethmoid bone they appear almo5t
fused together, the fit between them being very close (Fig. IOA,C). ' l h e anterior
superognathals are robust bones having a single row of low rounded teeth
present (the ventral tooth row; Dennis & Miles, 1979a; fig. 12), and lack
symphysial teeth. The posterior superognathals are almost rectangular in form,
N E W AUSTRALIAN ARTHRODIRE
243
C
Figure 7. Latocamurus coulthardi gen. et sp. nov. Holotype WAM 86.9.670. A, Jaw and skull
elements, x2.5 (see also Figs 5, 8). Note the unusual shape of the pineal plate in the top right,
behind the left postnasal plate. B, Ethmoid bone and rostra1 plate, ~ 2 . 2(see also Figs 2, 11).
C, Median dorsal plate in dorsal view, x 1.35 (see also Figs 2, 5).
being very thick in lateral view and with well-developed dorsal processes
(Fig. 1OC). There are three large, rounded teeth situated in a lateral row,
comparable to the multicuspid teeth of Bullerichthys (Dennis & Miles, 1980). The
crushing teeth of Latocarnurus differ from those of Bullerichthys in lacking a dentine
coat and in the absence of scattered isolated teeth. It is clear from close
t'iqutc 8 . Latocamurus coulthardi gen. et sp. nov. Skctcli ol'jaw e h l r I i t 5 . postnasal pLitcs ,tiid
\c11tral surfiicc 0 1 rtllmold kxmc, holoc) pc L V A X l 86.9.670 (ace also Fig. 7'4:. 'Ttic outline ( i t ' t h c
right antcrioi- superogtiatli;d i t i l i k prirition i s shown on thr vcntral surfiice of t h r rthmciid.
iiidic'it(d I)! .in arrow. I n w t slro\vs t l i c p s t e r i o r process of thc left ;interior superognatli;il from t t i ( ,
ctiorr of the right postnasal plate in thc t(Jp Irft of tlic f q u r c \IIOMS t11c
c.\tctisi\ c hcirizotital Iiimitin for the o\ crlap 0 1 ttrc r ( i s r r a l plate.
245
Figurc 9. Latocamurus coulthardi gen. et sp. nov. Ventral asprct of thoracic armour and parts
0 1 thc chcck unit. holot)-pc \VAhl 86.9.670. (dusted with ammonium rhloridc).
comparison between the dentition of these two taxa that Latocamurus was a
durophagous feeder like the other camuropiscids.
3. Ginglymoid dermal neckjoint. T h e dimensions of this feature are stated in the
measurements. It is typical of the neck joint described for other camuropiscids
with the para-articular process and lateral articular fossa well developed, and a
wide nuchal gap present.
4 . Median dorsal plate wilh ventral ridge. Developed as for other Gogo
arthrodires.
5. Occipital cross-commissure passing of the hind margin of the paranuchal. Developed
as for Camuropiscis. Extrascapular plates are not seen on either specimen.
6. Suture lines remaining distinct, sinuous in skull-roof,whose plates have well-developed
overlaps. These characters are seen in the illustrations. Some points regarding
overlap relations are: postnasal plates large, preventing contact behind them
between the suborbital and preorbital plates; inward lamina on the postnasal
plate for overlap of the rostral plate; pineal and rostra1 plates separated by
mesial contact between the preorbital plates; suborbital has a n irregular
J . LONG
246
10. Latocamurus coulthardi gen. et sp. nov. Ethmoid bone with associated
p.ir.i~phciioicl and toothplnrca of the right side. lVAh1 86.9.699. A, Antcrior \iw'; B. oblique. kft
I;itcIal \.ic\v; C. ventral view. A, B? C, x 4.4. D, oblique ventral \icw of anterior of hcad s l i o w i n ~
cthmoicl boric.. parasphrnoid and right supcrognathals in lifc position, x 2.5. (All dustcd w i t h
ammonium chloride I
Fixutc
posterior margin where the postsuborbital plate is firmly sutured; submarginal
plate has large dorsal overlap flange for marginal plate; central platr has a long
suture with postorbital plate, excluding contact between the paranuchal and
preorbital plates; anterior lateral not contacting the anterior ventrolateral due
to long spinal plate; extensive contact margin betwecn the anterior latcral ancl
interolateral plates; posterior lateral has a dorsal overlap surface for thc
posterior dorsolateral plate; anterior median ventral and posterior median
ventral have a short median contact margin.
7. -tuchal plate posteriorb expanded. As for Camuropiscis.
8. Paranuchal plate with postnuchal process. As for Camuropiscis.
9. Xuchal thickening on zlisceral surface o f skull roof. As €or Camuropiscis.
10. Paired pits on uisceral surface of skull roof. As €or Camuropiscis.
11. Pre-endol_vmphatic thickening. As for Camuropiscis.
12. PPcLomIjn long, horizontal base articulating against laterally,facing scapulocorrrroid.
NEM' AUSTRALIAN AR'I'HRODIRE
247
A
Figui-c I I . Latocumurus coulthardi gen. et sp. nov. Parasphrnoid in A, ventral, B, dorsal and
C:. oblique lateral \ icws. Holotypc LVAM 86.9.670.
Presumed to be as for Camuropirczs, except that the spinal plate is slightly longer.
13. ~nferog~ialhal
wilh both blade and biting regiom, which are complele and co-ossified.
The relative lengths of the biting and blade divisions of the inferognathal are
exactly the same as in Camuropiscis. T h e mesial groove is not seen. T h e biting
region is thick with a flat occlusal surface lacking any cusps, but with shallow
depressions opposing the blunt teeth of the posterior superognathals (Fig. 2).
The anterior end of the biting region is narrow with an anteroventral thickening
present. T h e mentomeckelian and articular ossifications are well preserved, the
former being anteriorly narrow but very deep and thin posteriorly; the articular
is essentially as in Camuropiscis with a relatively high articulation surface and a
well formed posterior process.
14. Skull-roof with lateral consolidated region. Visceral features of the skull-roof are
shown in Fig. 3. The left postorbital shows this feature well developed
(Fig. 13A) as a triangular depression exactly similar to that of Camuropiscis
(Dennis & Miles, 1979a: fig. 16). No inframarginal crista (sensu Miles & Westoll,
1968) is developed.
15. Submargznal plate small and closely incorporated into cheek unit. The externally
exposed surface of the submarginal (Fig. 2) is quite small as in the other
camuropiscids, with irregular contact margins for both the postsuborbital and
marginal plates. T h e submarginal has a long anterior lobe, but does not
preclude contact between the suborbital and marginal plates. It meets the
postsuborbital along an irregular suture.
16. Skull-roof with well deueloped supraorbital vault. As for Camuropiscis, including
the delicate ventral post-ocular process developed (pt.o.pr, Fig 3 ) .
17. Suborbital plate with slender suborbital laminae. T h e suborbital has a shorter
suborbital lamina than that of Camuropiscis, with a deeper, more rounded
A
nos. cap
tub
nos cov
br.cov
pr.pos
5 rnm
3
nos cop
postorbital division (Fig. 2 ) . There is an extensive anterodorsal overlap margin
for the postnasal plate and a well marked suborbital crista (cr.so, Figs 3, 13;).
'l'he posterior margin of the suborbital is notched for a firm suture t o the
postsuborbital plate.
18. Medinn dorsal Plnte with carinal process. T h e carinal process jcr.pr, Fig. 2 j is
quite broad, and stout, directly comparable with that of other camuropiscids.
The posterior margin of the median dorsal plate is pointed but not drawn out
N Lit’ AUSTRAL1 AN AKTHRODIKE
’ /
pr det
’
249
I crn
t‘igurr 13. L Q ~ O C U ~ coulthardi
U~SS
gen. et sp. nov. A, A’, Stereo pair showing features of the
\isccral surfacr of the check and skull roof. Notr the long quadrate which hcrrds vrntrally brforc
rcncliing the triangular deprcssion on the postorbital plate. B, Features of the check complex in
\ iscrr;il \.irw. iY.411 86.9.699.
into a spine as in Harrytoombsia. The sides of the median dorsal plate are parallel
for about half the plate length in the holotype (Fig. 7C). T h e anterior margin
comprises two strongly convex sides (Figs 1, 7C). The sub-median dorsal bone is
slender as in Camuropiscis.
19. Prcorbital plate with preorbitaal lamina. As €or Camuropiscis except that the
preorbital plate is arched throughout its anterior half to give the snout its
downturned appearance (Fig. 2). T h e postnasal plates are curved in crosssection having a short dorsal division and a larger lateral lamina posteriorly.
The postnasal plate is quite deep, the depth/length index being 64 (Figs 2, 8).
‘The supraorbital sensory-line canal runs off the postnasal plate at the anterior
free margin which is not in contact with other plates (Fig. 15).
20. Suborbital plale with internal laminae. As for Camuropiscis, described under 17.
21. Palaloquadrate with separate autopalatine and quadrule bones. The autopalatines
are well preserved attached to the lateral faces of the ethmoid bone (Figs
10-13). In both specimens the autopalatine is fused to the ethmoid ossification
to give great rigidity to the attachment surface for the superognathals. The
quadrates are long perichondral ossifications bearing articulatory condyles and
a small elongated detent process (pr.det, Fig. 13) near the ventral end of the
250
J LONG
bone. The anterior end of the quadrate is very narrow, almost slit-like. As in
Camuropurz r the quadrate reaches towards a triangular depression on the
postt"rhit,il bone (Dennis & Miles, 1979a fig. 16), but in Latocamurur i t cur\es
entrally before meeting the triangular depression (Fig. 13A)
22. SuI)a~ognathalJ tedh postpilor procasser. T h e toothplates have ahead) been
described no. 2 ) .
ADDITIOSXI. 1'OlXl.S OF STRUGTIIRE
In this section the parasphenoid and ethmoid ossification of the braincase are
described. The pclvic girdle of WAM 86.9.699 is well-preserved but is identical
l o that of Camuropiscis concinnus and is not described here. Fragments of the
scapulocoracoid and the vertebral skeleton are preserved posterior to the trunk
ximiour in the holotype (vert, Fig. 5) but these do not differ significantly from
those described for other Gogo arthrodires (Dennis & Miles, 1981; DennisBr, ;m, 1987) to ivarrant firrther comment.
T h e parasphenoid
T h i s boric. (Figs 10- 12) is similar in its narrow shape and deep profile to that
ol' Ijruntonichthys (Dennis & Miles, 1980; fig. 4) and Tubonasus (Long, 1988). It is
pc.rf'crtly preserved on WAM 86.9.699, still articulated to the ethmoid
ossification, where it juts out at an angle of c. 45" from the horizontal plant, of'
the autopalatine-ethrnoid suture (Fig. 10B). O n the holotype it sho\vs thc.
complete dorsal and lateral surfaces and part of the Lrentral surface which \\-as
damaged through the plane of splitting when the specimen was fourid (Fig. 1 1 I .
'I'hc ventral surface (Figs IOC, 1 I A ) consists of a smooth, deeply grcto\~d
anterior division with a central median crista (cr.m.), and a relatively flat
posterior division, the two divisions being separated by a prominent curved
ledge (le), just posterior to the depression for the bucohypophysial foramina
il>.hyp). There is a short lateral groove (gr.a.com) for the interrial carotid
arteries as in the other Gogo arthrodires. The ventral division has a narrow
transverse groove for the lateral branchcs of the hypophysial vein (v.1.h) p I .just
anterior to the large central foramen for the posteriorly directcd median
hypophysial vein (v.m.hyp). Numerous foramina (for) are scattered over the
surface of the bone between the small transverse groove and the ledge on the
holotype parasphenoid.
In dorsal view (Fig. 11B) there is a small triangular surface which contains
the paired ovoid depressions, presumably for the lobus hypothalami, in which
the buccohypophysial foramina (b.hyp) are located. There is a deep lateral
groove (t.l.gr) on the dorsal surface just posterior to the median process in1.p~).
The grooves on the dorsal surface run into the centre of the bone from a
dorsolateral direction, rather than horizontally, as do the transverse latera!
grooves on the ventral surface of the parasphenoid. This suggests that the
grooves on the dorsal surface may have been for veins draining downwards into
the median hypophysial vein which then courses posteriorly through the bone
via a small foramen (v.m.hyp). An unusual feature of the dorsal surface of the
parasphenoid of Latocamurus is the presence of a well pronounced median process
m.pr) posterior to the buccohypophysial foramina.
N E W AUSl’RAI.IAN ARTHK0I)IRE
25 1
T h e deep, narrow form of the parasphenoid in Latocamurus and Tubonasus
would suggest that this is a specialized feature of the Camuropiscidae (noting
that the parasphenoid has not been described for Camuropiscis or RolfsteuJ).
Close morphological relationship between Tubonasus and Rolfsteus suggests that
the parasphenoid of Rolfosteus is predicted to be of the narrow, deep type.
The ethmoid ossijication
T h e ethmoid ossification (rhinocapsular bone; Stensio, 1963) has been
described in three other Gogo arthrodires: Rolfosteus (Dennis & Miles, 1979b),
Incisoscutum (Dennis & Miles, 198 1 ) and Eastmanosteus (Dennis-Bryan, 1987).
Stensio (1963) has restored the ethmoid ossification from a grinding series in
Tapinosteus, and described it from actual remains in some other arthrodires (e.g.
Pholzdosteus, Trematosteus), and Goujet (1984) and Miles (1971), amongst others,
provide descriptions of this bone for some primitive arthrodires. T h e ethmoid
ossification of Latocamurus is perfectly preserved in WAM 86.9.699 (Fig. 10) and
fairly complete in the holotype embedded in resin, showing the lateral and
internal surfaces clearly, with the ventral surface preserved in the counterpart
block of resin (Figs 4, 5, 7A,B, 8, 12). As in other arthrodires the ethmoid
ossification has two laminae of perichondral bone enclosing an interperichondral
space which in life was cartilage filled. There is no internasal bone present,
although there is a stout internasal process projecting anteriorly below the
nasal cavities (pr.in, Fig 12D) which would have functioned similarly to an
internasal element. Unlike in Tapinosteus where the internasal process is most
developed in the dorsal part of the anterior surface of the ethmoid bone
(Stensio, 1963: fig. 52), the intcrnasal process in Latocamurus is ventral to the
cavities for the nasal capsules.
The dorsal surface of the ethmoid is not roofed by perichondral bone but was
closed over by the dermal skull roof. In dorsal view the internal surfaces of the
ethmoid ossification are clearly seen (Figs 7B, 12A). It is almost square, having
slightly longer anterior margin than posteriorly, and differs from that of
Incisoscutum in the well defined separate rhinal lobes of the brain cavity (br.cav),
which terminate at the cribrosal ossification. There are several small foramina at
the anterior end of the cavity for the olfactory tract (olf), representing the filia
ofactoria of cranial nerve 1 which run anterovetitrally into the nasal capsule
(nascap). In addition there is a small tubule of perichondral bone running from
the mesial side of the cavity, anterior to nerve 1, into interperichondral space
abovc the nasal capsule, ending at a point in the midline of the capsule on its
dorsal margin. This nerve tube (term?) is seen on both sides of the holotype
specimen and may possibly represent the course of the terminalis nerve,
previously described in only one other placoderm, the acan thothoracid
Brindabellaspis (Young, 1980). I n Brindabellahpis, the terminalis nerve runs from
the brain cavity just anterior of the olfactory nerve into the nasal capsule,
whereas in Latocamurus this nerve ends dorsal to the posterior encapsulated part
of the nasal capsule, presumably entering it in its anterior half. WAM 86.9.699
shows paired perichondral tubules for the terminalis in exactly the same
position. There is a large break in the dorsal rim of the ethmoid where the two
rhinal divisions of the cranial cavity meet, indicating the outwards passage of a
large cranial nerve, the optic nerve (gr.II), by comparison with other
252
J . LOXG
arthrodires (e.g. Tapinosteus, Incisoscutum). Behind the optic nerve foramen ihcrc
tnissing the dorsal region of bone above the opening) the ethmoid terminates at
the basal subocular process, here seen developed exactly as for Incisosculum.
In lateral view the ethmoid ossification (Figs 10B, 12C) shows a curved groove
( 1 .grj running from the base of the subocular process upwards
then down
anteroventrally ,just behind the nasal capsules where the autopalatines (Aup)
are fused to the ethmoid bone. Midway along this suture is a deep excavation
with two smaller pits contained inside its rim: a myodome for extrinsic eye
muscle attachment (my). I t is also noted that the posteriorly facing orbital vault
of the preorbital plate bears a well developed flange (PRO.fl, Fig. 14) which
abuts the ethmoid bone and may have acted as an attachment area for superior
and inferior obliquus muscles. T h e anterodorsal region of the lateral surface
s1mw.s a smooth attachment area to the preorbital plates ( a t t . P R 0 ) which juts
o u t as it short ledge; however, when articulated to the ethmoid bone there is a
small part of' this ledge which faces posteriorly into the orbital chamber
a.or.att) being confluent with the posterior orbital flange of the preorbital
platc. Figure 10D shows the ethmoid bone in life position, with upper jaw
toothplates attached. Ventral to the preorbital plate attachment area is a
smooth depression for attachment of the subocular process of the suborl>ital
p l a t e iatt.SO). Between this region arid the preorbital attachment area is a
Ivell-defined groove (gr.al.eth), possibly for passage of the orbitenasal artery
and vein. O n the left side of the holotype only there is a small circular ring of
bone (pp) dorsal to the suborbital attachment area which may have been for
ligaments from the suborbital, although this structure is not present on
\ Y A M 86.9.699. A large foramen penetrates the posterior wall of' the nasal
capsule and runs anteroventrally into the nasal capsule, via a perichondral tube
ivhich has accessory tubules of the snout sensory plexus branching off it. This
c;tnal is for the profundus nerve (prof). 'Hie course of the profundus is shown in
Fig. 12B, where most of the tubuli of the interperichondral space are
interconnected to this main nerve pathway. Immediately ventral to the large
cmbayment for the optic nerve there is a slit-like opening i n the
autopalatine~ethmoidsuture, possibly for passage of the orbital (or optic) artcr),
! 0rh.a. 1.
'I'he Lrentral surface of the ethmoid ossification is deeply grooved anteriorly for
attachment of the anterior superognathal (att. ASg, Fig. 8), posteriorly it
hecomcs steep sided to leave room for the hypophysial region where the
parasphenoid attaches (Fig. 1OC). Around the anterior margin, just ventral of
the curved embayments for the nasal cavities, there is a small groove (n.buc)
which runs parallel with thr anterior margin of the ethmoid bone, and then
posteriorly to course lateral to where the anterior superognathals were
positioned. This groove was presumably for the buccalis nerve and vessels as
noted for Tapinosteus and other arthrodires by Stensio (1963; fig. 50, gr.mb).
'I'he anterior superognathals attached to the ventral wall of the ethmoid at tbvo
depressed regions (dep 1, dep 2, Fig. 8 ) , and a well developed ridge (ri
posteromesially across the top of the superognathals, terminating at a small
foramen ( f 1 ) . A second foramen (f2)opens laterally to this foramen. Evidence of
another courseway for vascular or nerve supply to the snout and upper lip area
is seen in the form of a very small groove (gr) running antero-posteriorly on the
lateral side of the ethmoid ventral wall.
NEW AUSTRALIAN AKL'HRODIRE
253
PRO
/
I cm
E'igurc 14. Latocamurus coulthardi gen. et sp. nov. Skctch of cthmoid bone in postcrolatcral
vicw situated in orbital cavity t o show relationships brtwren thr ncurocranium, dermal skull roof
and chcck. \\'AM 86.9.699.
In anterior view (Fig. 10A) the nasal cavities are separated from the ventral
lamina by a flat anterior lamina. This surface is interrupted by the internasal
process (pr.in), b u t is otherwise featureless apart from grooves, noted above,
along the ventral edge.
In none of the previously described Gogo arthrodires has the connection
between the ethmoid ossification and the dermal skull roof and cheek been
discussed. WAM 86.9.699 is unique in having a complete ethmoid bone
perfectly preserved which can be easily fitted into life position (Figs 14, 15A)
showing that there were two major pathways for exit of the nerve, artery and
vein supply from the orbital region into the snout. The posterior orbital flange
on the preorbital plate (PRO. fi) fits against the suborbital plate allowing the
grooved anterior end of the subocular crista to run into an open groove plus
foramen (gr.n.buc, Fig. 15A), presumably for the n. buccalis lateralis and
n. maxillaris nerves. Mesial to this opening is a second foramen between the
B
I. rm
_
Figure 15. Latocamurus couZtlurrdi gen. e t sp. nov. A, Anterior view of skull with cthmoid
hone and infrrognathals in life position. B, Attcmptrd restoration of head in anterior view, srnsorvlirirs shown as unbroken lines.
251
J. L O N G
anteromesial edge of the suborbital and the orbital attachment area on thc
ethmoid hone, here termed the anterolateral ethmoid groove (gr.al.cchj . I t
probably enabled the orhito nasal vein and artery to pass into the snout region
as suggested for Tupinos/eu.c by Stensio (1963).
As few arthrodires have been reconstructed in anterior view I take the
opportunity to prrsent such a restoration of Latocamurus. Figure 15A shows the
dermal skeleton with ethmoid bone and inferognathals in life position i n
anterior view, and Fig. 15B shows an attempted restoration of the fish’s fact,
nit11 lateral line canals shown as unbroken lines (in life these were undoubtedly
pwes as in li\ing fishes). ‘I’he notch for the incurrent naris (na.in, Fig. 1514) is
me11 defined between the rostral and postnasal plates. ‘Thc excurrent naris was
probably situated lateral to the internasal process (pr.inj at the ventral limit of
the nasal capsule.
R t:I..4‘I’IONSHIPS O F LA 7.0
The fiimilj. C.hnuropiscidae was defined by Dennis & Miles [ 1979111 t o
include the genera Camuropiscis (two species; Dennis & Miles 1979a), and the
unusual tubular-snouted forms Rolfosleus and Tubonasus (Dennis & Miles, 19791):
Long, 1988). .\I1 three possess a n elongated armour with pointed rostra ;most
dei,eioped i n Rm(fii.cleer.r and Tuhonnsus as tubular rostral plates) as well as other
specializations listed in the original familial diagnosis.
Denison j 1984) suggested that the Camuropiscidae arc the sister group t o
1wi.5o.(cuIuttT i\vhich he placed in the monotypic family Incisoscutidae), a s the!,
share the t‘ollowing synapomorphics:
1 A deep postnasal plate participating in the orbital margin.
(2)The submarginal plate is reduced more so than in typical coccosteids.
13) ‘lhe spinal plate is reduced antcriorly.
1
‘lhese characters are corroborated from my own observations of specimens
although the last character is here modified from Denison’s original wording
“spinal plate reduced anteriorly and posteriorly” because it is difficult t o
demonstrate the posterior reduction of this plate. ‘I’he anterior reduction of the
spinal is confirmed by the more extensive contact area between the iriterolateral
arid anterior lateral plates in these taxa, whereas typical Gogo ‘coccosteids’ show
comparati\dy very short contact between these two plates. One feature Denison
used to separate the Camuropiscidae from Zncisoscutum-the contact between the
anterior lateral and anterior ventrolateral plates--is not found in Zdocamuru.r
arid represents the presumed primitive condition for camuropiscids and othcr
eubrachythoracids. T h e cheek of Incisosculum features reduced submarginal and
postsuborbital elements which arc tightly connected behind the suborbital, a
condition similar to that of camuropiscids but lacking firm suturing of the cheek
u n i t t o the skull roof. Other striking similarities between ~ 7 ~ c ~ . s o , s cand
u~u~~
t,amuropiscids (but not necessarily synapomorphies) are the similar shape of the
i tifi.rognathals in Incisosculum and juvenile C‘umuropiscis, and the same t 1 - x of
scapulocoracoid ii.e. similar shape and same number of foramina in the
articular crest i.
NEW AUSTRALIAN ARTHRODIRE
255
Latocamurus and other camuropiscids are more derived than Incisoscuturn in
having:
(4) A broad rostral plate. Although the exact shape of the rostral plate differs
i n the two specimens, in both it lacks a n extensive posterior median lobe, a
feature present on all T-shaped rostral plates of coccosteomorphs. Incisoscutum
shows some variation in this plate, but the normal condition is to have a
T-shaped rostral plate.
(5) A deep postnasal plate which participates in the orbital margin and
excludes contact between the preorbital and suborbital plates. In Incisoscuturn the
postnasal is comparably broad but does not exclude contact between the
suborbital and preorbital plates.
(6) A suborbital plate which is firmly articulated to the postorbital plate.
( 7 ) The dentition is durophagous.
(8) The submarginal and postsuborbital plates are strongly interconnected,
forming a posterior cheek unit behind the suborbital.
(9) The preorbital plates have short mesial contact with each other.
(10) The postmarginal sensory-line canal is not present.
( 1 1 ) The main lateral-line has a ventral course on the anterior dorsolateral
plate.
Camuropiscis, Rolfosteus and Tubonasus are more derived than Latocarnurus in
having:
(12) A pointed rostral plate.
( 1 3) T h e submarginal lacks an anterior lobe, instead being short and deep.
( 14) Elongated preorbitals having more extensive median contact.
(15) Anterior lateral plates contacting the anterior ventrolaterals along an
extensive margin.
Both Tubonasus and Rolfosteus are more derived than the previous species in
having:
(16) Tubular rostral plates.
T h e recent discovery of a new advanced camuropiscid having a rostral plate
like that of Camuropiscis concinnus but a cheek plate arrangement like Tubonasus
Rolfosteus
Tubonasus
Camuropiscis
Latocamorus
Incsoscutum
Harrytmmbsm
Figurr 16. Cladograrn of suggested relationships betwccn a typical coccostcid iHnrrytoombsia),
carnuropiscids and Zncisosrutum.
J. LONG
256
(in which the postsuborbital sutures with the marginal) suggcsts that the
tubular rostra of Kolfosteus and Tubonasus could be independently derived. This
problem will be considered in detail after the new form is described. 'The abovc
character scheme is summarized in the cladogram (Fig. 16). It is rioted that not
all of the above characters are unique to camuropiscids and Zncisoscutum within the
whole of the Arthrodira, but most represent specializcd conditions within thc
framework of higher eubrachythoracids. As monophyly of camuropiscids is
easily demonstrated (character 5), the acquisition of all the other characters can
be seen as specializations within the family Camuropiscidae, irrespective of thc
fact that certain other arthrodires possess some of these characters.
(:Os(:LUSIONS
A IICW camuropiscid, Lntornrnurus coultfiardi gen. et sp. nov., is described from
the Upper Devonian Gogo Formation, \.l"estern Australia, and thc famil)Camuropiscidae Dennis & Miles, 1979b is redefined to include characters of the
new genus. Lntocamurus is regarded as the primitive sister taxon to a l l othcr
camuropiscids as it lacks contact between the anterior lateral arid antc'rior
ventrolateral plates and does not have a projecting rostra1 plate. The cthnioid
ossification of Lntocamurus shows the possible presence of a terrnirialis ncr\.c
running into interperichondral space above the nasal capsules, and has tlir
autopalatines fused to it. The parasphenoid of Latocnmurus is unusually deep arid
narrow as in Tubonnsus, and is firmly attached to the ethmoid boric..
Carnuropiscids and Zncisosculum are closely related b y shared features of' tlic
postnasal plate and cheek unit, as proposcd by Denison ( 1984).
h(:KN0\VL,ED(;E~~ESlS
The author acknowledges a Xational Research Fellowship-Queen Elizabeth
I1 Award enabling this research to be undertaken at the University of b'estern
;\ustraiia. Fieldwork at Gogo in 1986 1987 was funded through the National
Geographic Society grant Number 3364-86 and the Q E I1 Research Fund. I
offer thanks to the following volunteer workers who assisted me at Gogo: &fr
Chris Nelson, Miss Susan Creagh, Dr Richard Holst and M r Terry Barnes, and
t o hlr Jim Coulthard and M r 'Len Hill, Gogo Station, for permission to collect
specimens. Professor Brian Gardiner (King's College, London) is thanked for his
helpful comments on the work.
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ABBREVIATIONS USED IN T H E FIGURES
ADL
.IL
X\lV
a or
Ar
.%
att
anterior dorsolateral plate
anterior lateral plate
anlerior median ventral plate
anterior orbital attachment surfare
of cthmoid bone
articular
anterior superognathal
atl.
ASg
att.PRO
att.SO
;\up
attachment area for anterior
superognathal
attaclimrnt area for prrorbital
platr
attachment area for suborbital
plate
autopalatine
ri
occipital sensory-line dndl ~ I . < K J \ c
olfactory nerve
ohtected riuchal area
part of thr orbital margin
Ibrarncn for orbital a1-tery:'
pineal plate
posterior dorsolatcrnl platr
prctoral fcnstra
pineal foramen
posterior lateral plate
posterior marginal plate
posterior mcdidn \ rntral pl'itc
postriasal plate
paranuchal platt
p s t r r i o r pit line g r w n c or
posterior process on rthtrioid t i o w
mcdisn posterior process on nuclial
plate
dctrnt process of q u a d r a t e
preurbital platr
prcorbital flange
internasal process
stipcrlieialis prc hndur ri(mc'
postcrior process
posterior wpcrognaihlil boiic
postsuhorhital p l a t e
parasphenoid bone
postorbital plate
\ rntral postocular process
paired pits on visccr,tl surface o f
nuchill plate
posterior vcntrolattral p l a t e
quadrate hone
rostra1 plate
ridge on vrntal surliice of rthmo~tl
Sc
sclrrotii bones
Set.
s11
scapulocoracoid
submargiual plat c
suhorhital plate
subobst;intic area of licadshicld
supraorhital sensor! -line c;iiiiiI
groin e
suborbital crista of suborbital plate
oral pit-linr groovc
spinal plate
supraorbital vault
tcrminalis ncrvc
prc-endolymphatic thickrning
tr;ins\-ersr lateral sr o w c on dursiil
surface of parasphenoid
triangular dcprcasion on p~istwttital
platc
rostra1 tubuli in eihmoid bone
vcrtcbral rlrmrntc
groove on parasphrtioid for l a t e r a l
bi-anches of the hypophysial \rill
1i)ramc.n for rnrdiari hypophysi,rl
:interior \entrolatcral plate
bticcohypophysial for am in:^
Ixairi cavity
OC
< cl1tl;II platc
orb
orb. il
i.iiannel for doi-sal 2isprt.t of
prrort)ital procrss o f nciirocraiiiuni
c Iiiiiincl fool- dorsal aspect of
aulir.i\ q a l process of ~i curocrai i i ut ~~
mibi osal ossilication
nicdian Irist'i of parasphcnoid
posterior crista of mrdian dorsal
Ill a t <,
\uIxiciiLii- crista
rc~i~I:iI
sc~isor)--linccanal groove
opening for the eridol) niplidtic duct
drlm~ssioiison ventral surfice of
rthmoid hoiic h r contact w i t h
aiitcrior superogiiathal
rtlimoid (rhinocapsular) boric
hrarnina i n ventral wall of cthmoid
I10nc
rntial stirficr ol'thc ethninid
hi-amina o t i vciitral surfiicr of
parazpticnoid
groo\ c OII ventral surfice o f
cthmoid bonc
groove on parasphenoid
h r intcrnal carotid arteries
aiitcrdtitcral groovr hctrc..ccn the
cthnioid 2nd suborbital horrrs,
p t ~ o I ) ~ i hfur
l ~ orbitori;isal artery and
\(in
groo\-c and foramen fir i t . buccalis
fbramrn h r the optic nerve
groo\ t' Tor posterior superognathal
groo\ e for submarginal plate
i~ilerog~iatlial
I)oiic
iiifraor1)ital scnsory--liiie c a n a l
groo1.c
in tcrolatcrd pla tr
postpcctoral lamiria of postci-ioi
\ ciitrol,itcral plate
niain late-ral-line canal groin e
~raiisvcrscledge 011 parasphenoid
lcitcrLtlgroovc of cthmoid hone
ni,trgiil,il pliitc
rncdinii doi.saI platr
mcntomrckrlian ossilication
rniddlr pit-line gram e
nirdi;iti process o n dorsal side (if
p;trasphcnoid
possi l i l c myodolnc
~ i o t c l iliir incurrent n,ii-is
\
n'isal capsulc
t i ~ s a lcavity
gt-oovc. fix 11. huccalis and vrssels
nuch'll "late
ovrr1.rp inargin Tor prcorhital p h t c
o\ erlnp lamina for rostal platc
o\-crlap iirra fbr suborbital plat(.
Olf
011
P
P111,
pcc. f
Pi
1'L
PhfG
FhfV
I"
PSC
PP
P.P'
pr.dct
PRO
1'KO.H
pr.in
pr0f
pr.pos
PSI:
PSO
1's p
PTO
pt.o. pr
[Jl.li
PYL
Yd
K
bonr
so
SOa
soc
so.cr
wr
SP
sue.\
term
th.pr
t.l.gr
tri
tub
vrrt
v.l.hyp
\
.m.h)p
\ ('111