PRELIMINARY ANALYSIS OF THE INTERRELATIONSHIPS OF SOME

PALEOZOIC ACTINOPTERYGH

RICHARD LUND, CECILE P O P L I N & KELLY McCARTHY

LUND R., POPLIN C. & McCARTHY K. 1995. Preliminary analysis of the interrelationships of some paleozoic actinopterygii. [Analyse pr~liminaire des relations phylog~n~tiques de quelques Actinopt~rygiens pal~ozo'fques]. GEOBIOS, M.S. n ° 19 : 215-220.

ABSTRACT

A cladistic analysis of 20 actinopterygian species, including several new taxa from the Lower Carboniferous Bear Gulch Limestone of Montana, USA, is presented. This analysis is based upon 74 dermal cranial and shoulder girdle characteristics. The results strongly support several classical groupings such as that of Paleoniscimorpha. On the basis of these cranial data, there is some relationships between Rhadinichthys canobiensis and two of the Bear Gulch actinopterygians, all characterized by the lack of premaxillae. This analysis will later be expanded with other characters such as postcranial ones.

KEY-WORDS : CLADISTICS, ACTINOPTERYGII, DERMAL SKULL, SHOULDER GIRDLE.

RI~SUMI~

Une analyse cladistique de 20 esp~ces d'actinopt~rygiens, dont plusieurs in~dites du Carbonil~re inf~rieur du calcaire de Bear Gulch, Montana, USA, est pr~sent~e. Elle est fond~e sur 74 caract~res dermiques du cr~_rle et de la ceinture pectorale. Les r~sultats confirment des groupements classiques dont celui des Paleoniscimorpha. Ces donn4es indiquent des relations phyl~tiques entre Rhadinichthys canobiensis et deux actinopt~rygiens de Bear Gulch, tous caractSris~s par l'absence de pr~maxillaire. Cette analyse sera ult6rieurement pr~cis~e par l'adjonction d'autres caract~res en particulier postcr~niens.

MOTS-CLt~S : ANALYSE CLADISTIQUE, ACTINOPTERYGII, CR~NE DERMIQUE, CEINTURE PECTORALE.

A cladistic analysis of 20 species of actinoptery- gians (18 fossil ones including several unpublis- hed taxa from the Lower Carboniferous Bear Gulch Limestone of Montana, USA ; and two re- cent) is presented. This analysis is based upon 74 cranial and shoulder girdle characteristics (Tabl. 1, 2). The theoretical outgroup was coded with all characteristics primitive (0). The cladistic pro- grams used were Hennig 86 and Clados. When certain characteristics were considered additive- ly, the analysis resulted in one tree, with tree length 286, consistency index 50 and retention index 55 (Fig. 1). When all characteristics were considered non additively, two trees resulted, with tree length 256, consistency index 53 and retention index 55. One of the latter trees is identical in structure to Figure 1, while the other differs only in the connection of Pteronisculus di- rectly to the preceding node.

The data strongly support several hypotheses. The taxa "Guilday" and "Discos" are sister taxa within the Cladistia. There is an apparently mo- nophyletic taxon, the Paleoniscimorpha, as sug- gested by Kazantseva (1971). On the basis of these cranial data, there is some relationships between Rhadinichthys canobiensis and two of the Bear Gulch actinopterygians.

The taxa involved in the analysis and sources of information are as follows : Polypterus bichir, Allis 1922 ; Pehrson 1947; and specimens. "Guilday" and "Discos", Bear Gulch Limestone, Lurid 1995, ms. Caturus driani, Saint Seine 1949. Amia calva, Allis 1897 ; Jollie 1969. Enneles audax, Da Silva Santos 1960. Paratarrasius hibbardi, L u n d & Melton 1982.

216

*0 Premaxi[[ary teeth position

1 Prema.xillm 3' teeth assoc-iatioQ

*2 Prem~x~Jlm3' line condition

*3 Ro.~ral bone

4 Thoothed rostral

"5 Ethmoide eOmrni~5-Lll~

6 Rostra,a] hones number

"7 Pos'trostral bone

*8 Snore

9 Roatral notch

10 Nasal bones

11 Rear of nasal (s)

12 Tectals

13 Supraorbital bones

"14 Supraorbital canal

"15 Antorbital bone

16 Amorbital bone

"17 Supraorbital series

18 Frontal bones

19 Parietal/frontal size

20 Pineal foramen

"21 Otic canal bones

22 Supraorbital canal

23 Infraorbital canal

*24 Dermosphenotic position

25 Dermosphenofic/nasal contact

*26 Dermosphenotic bone size

27 Supraoceipit,al bone

28 Occipital scale row

*29 Supratemporal commissure

*30 Ex'u'ascapular bones

31 Profondus canal

*32 Posteroventral infraorbital is #

*33 Posterodorsal m£raorbital is ;~

34 Posterodorsal inf.raorbital shape

35 Po~erodorsal infraorbital

36 Suborbital bone arrangment

37 Suborbital bone numbers g

*38 Cheek fit

39 Suborbital bone thickness

40 Suborbital-maxil]a relationship

*41 Preopcrcular canal

*42 Prec~eronhml/sqamosa! shape

pleslomorphie state 0

in midlin e

on prem~_xillary bone

sutured

separate

absent

inAm der rostral

median

several median

momh subterminal

absent

many, mosaic

preorbital

present x~th lateral llne canal

Series

mesial to nasal openings

recumbent

indistinct/absent

single series

many

large

in p ustrosta-al area

2

to profondtm

to demaosphenotic

at postorbital comer

absent

subequal to dermopterotlc

absent

absent

across midliue

3

present

2

3

slmp]e

on cheek

none

none

fight, abutting

thick, ganoine

not betwee~ maxilla & infraorbital

three bones

La[[. narrow

apomorphic state l

not in midllne

oa rostral

loose

fused to postrostral

present

in/under premaxilla

paired

single median

mouth terminal

vertical

serial

midorbital

present

s-hale

between nasais

inverted L

with infraorbital canal only

double series

two pairs

small

in frontal

3

to p arietals

to dermopterotlc

anterior to postorb, process

contacts nasal

reduced

only on braincase

present

not across midline

I pair

absent

l

2

T-shaped

on postorbltal process

1 row

I-3, rounded

thin. overlapping

thin, no ganoine

baween maxilla & infi-aorb~al

2 bones

"hatchet-shape"

apomorphic state 2

absenl

on antorbital

fused

fused to premaLxilla

apomorphic state 3

one infraorbital

(+ state 4 other)

absent

absent

multiple absent

median & paired paired

~aom sharply protrude

keyhole-shaped

single bone absent

absent

absent

absent

both

vertical

with three canals

absent

1 pair

2 pairs

absent

1

other

other

posterior absent

expanded, elongate

on skull roof

loop to skull roof

2 pairs

>2

4

p ost erinrl.v expanded

rows

3-6. elongate

thick, peripheral gaps

absent

] bone

trapezoidal

canal post. to nasals

absent/fused

>2 pairs

3-5 or more

many

scale-like

nude

many bones

217

"43 Imeropercular bone(s) present, series shagle bone absent

• 44 Precrper~lar canal to posteroventral comer orbit-jugal to mid-otJc region to posto~ital process

45 Quzdratojugal line on quadratojugal & maxilla on preoperonlum" reduced

46 .lugal line to infraorbital reduced absent

47 Quadratojugal bone present reduced absent

48 Anteoperculum (dermohval) absent single series

• 49 Posterior anteoperculars absent 1/3 do~xaa operculum length of operculum

50 Maxillary mobility absent present

51 Maxilla length posterior to orbit orbkal preorbital

52 Maxilla posterior end narrow moderate plate high plate

53 MaxRlar 3' fossa of mandi'bule absent rear of mandible most of mandible

• 54 Ma_x.il]a-preopercular ganoine circumferential/horizon ridges vertical ridges tuberculafions

• 55 Marginal tooth orientation vertical forward posterior

56 Ma~&qfle rimes o eoronoid process no eoronoid process peudocoronoid process

• 57 Dentition single row m£xed large & small setifosm

58 Supraopercular bones absent present

• 59 Operculum tall rectangula~ short reduced

• 60 Suboperculum shorter than operc'alar taller than opercular obscure/absent

61 Anterior branchiostegals numerous few not below mandible

62 Posterior branchiostegals tmiform short behind rnmada~ole modified

63 Ex't.ralateral gular absent present

• 64 Gulars median median median row absent

• 65 Gulars lateral short extended absent

66 Clavicles present absent

• 67 Clavicle contact absent sho~ elongate

• 68 Clehhrum broad laminae arrow top spike

69 Anocleithrum present with canal present no canal absent

70 Scleroties many 4 2

71 Parasphenoid short length of braincase

72 Nasal caps-ale ventrolateral dorsolateral

73 Suspensorium autodiastylic limited methyostyly ex'tensive methyostyly

both 0 & 1

> one rolA '

~Sth supramaxilla

absent

absent

Table 1 - List of characters used in the phylogenetic analysis, (0) plesiomorphic state, (1, 2, 3, 4) apomorpkic states, (?) missing data. * indicates characters traited as non additive. Liste des caract~res utilisgs pour l'analyse phyloggndtique, (0) gtat pHsiornor- phe, (1, 2, 3, 4) dtats ddrivds, (?) donnges manquantes. * indique les caract~res qui sont traitgs cornrne non additifs.

Birgeria groenlandica, Nielsen 1949. "Tubesnt", Bear Gulch Limestone, Lowney 1980 and specimens. Cheirolepis trailli, Pearson 1982. Moythomasia nitida, Pearson 1982. Pteronisculus magna, Nielsen 1942. "Aesopich", Bear Gulch Limestone, Lund & Po- plin 1995, ms. Canobius ramsayi, Moy-Thomas & Bradley Dyne 1938. Paleoniscus freieslebeni, Aldinger 1937.

"berger", Bear Gulch Limestone, Lund & Poplin 1995, ms. Rhadinichthys canobiensis, Moy-Thomas & Bra- dley Dyne 1938 ; Lowney 1980 ; and specimens. "Rdicks", Bear Gulch Limestone, Lund & Poplin 1995, ms.

R E F E R E N C E S

ALDINGER H. 1937 - P e r m i s c h e s G a n o i d f i s h e aus Os t - g r e n l a n d . Meddelelser orn Gronland, 102 : 1-392.

2 1 8

NO- Outgroup 0 0 0 0 0 0 0 0 0 0 0 Cheirolepls 0 0 2 0 0 0 2 0 1 0 1 Hoythomasla 0 0 O I 0 2 0 I 1 0 2 Paratacrasius 0 0 0 0 0 0 0 1 0 0 1 Guildayic 0 0 1 0 0 0 0 2 I 0 I Rdlcks 2 4 3 1 0 0 0 I 0 2 2 berqer ~ 2 4 3 l 0 0 0 i 2 1 2 Aesop]ch Discos Polypterus Rhcanoblen Tubesnt Birgerla Cramsay Boreo rleronis r f r e i e s A m i a C . d r l a n l E a u d a x

NO. Outgroup

0 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 0 0 0 0 0 0 0 0 0 0 0 I 1 I 0 I 2 2 0 0 1 2 I I 0 I 2 2 0 1 0 1 0 1 0 2 I 1 1 2 I 2 i ? 0 I I I I 2 I 2 2 I 1 2 2 2 I 2 1 2 2 1 0 2 2 2 1 2

1 0 1 1 0 0 0 1 0 1 2 1 2 2 1 2 2 2 2 1 2 0 0 1 0 0 0 0 2 1 0 1 0 2 1 ? 0 1 1 1 0 2 0 0 2 0 0 0 0 3 1 0 1. I 2 2 0 0 1 2 2 0 2 2 4 3 1 0 0 0 1 0 2 2 I 2 2 1 1 2 2 2 I 2 0 0 0 O 0 0 0 I I 0 2 I 2 0 1 0 2 0 2 I 1 0 0 0 2 0 1 0 1 1 0 2 1 2 0 2 0 2 1 2 I 2 0 0 0 1 @ 0 0 1 1 0 2 1 2 2 1 0 2 2 2 1 2 2 4 0 .1 " 0 0 0 1 0 0 2 1 2 0 1 0 2 2 2 1 2 0 0 0 0 0 0 1 i 0 0 2 1 2 2 1 0 2 2 2 1 2 2 4 3 ] 0 0 0 1 0 0 2 0 2 0 1 0 2 0 2 1 2 0 0 0 0 0 0 0 3 1 0 3 2 2 2 2 0 2 2 2 1 2 0 0 0 0 0 0 0 2 1 0 3 2 2 0 0 0 0 1 2 1 2 0 0 0 0 0 0 0 4 1 0 2 0 2 0 1 0 3 1 2 1 2

21 22 23 24 25 26 27 28 29 30 31 32 33 34 35 36 37 38 39 40 0 0 0 0 0 0 0 0 0 0 0 0 0 0 O 0 0 0 0 0

C h e i r o l e p i s 0 1 0 0 0 0 0 0 0 1 1 0 0 1 0 0 0 0 0 0 H o y t h o m a s i a 0 1 0 0 0 0 0 0 ? 2 1 0 0 1 0 1 1 0 0 0 ~ r a t a r r a s i u s 0 2 0 2 0 0 I ] 0 1 1 0 0 1 0 3 ,3 0 0 0 G u i l d a y i c Rdicks berger Aesopich Discos Po]ypterus Rhcanobien Tubesnt Di[geria Ctamsay eoreo Pteronis Pfreies ~ . i a C d ~ i a n i Eaudax

1 2 0 0 1 2 2 0 0 1 0 2 2 0 1 3 3 0 0 I 0 1 0 0 1 0 0 0 1 2 1 0 0 1 0 1 1 1 1 I 0 1 0 0 I 0 0 0 I 2 1 0 0 1 0 I 1 1 0 I 0 1 1 1 1 1 0 0 0 2 1 1 0 1 1 1 1 1 1 0 1 1 0 0 0 0 2 0 2 2 1 2 2 0 0 3 3 0 0 I 2 0 2 0 0 3 0 0 0 2 0 1 1 0 0 1 1 2 O 0 1 1 0 0 0 0 0 0 0 2 1 O 0 1 0 1 1 1 1 1 0 1 0 0 0 0 0 0 0 0 1 0 3 0 0 1 2 0 O 0 0 1 0 0 0 0 0 0 0 1 1 0 3 1 0 3 3 0 0 1 2 1 1 3 0 3 0 0 0 1 1 1 0 1 0 1 1 0 0 0 2 1 O 1 1 0 0 0 0 2 1 0 2 1 0 1 2 0 0 0 0 1 0 l l 2 0 0 ] 2 1 0 3 1 0 ] 1 0 0 1 0 1 0 0 0 1 0 0 0 2 1 0 3 0 0 1 1 0 0 1 0 1 0 0 0 1 0 0 0 1 1 2 3 2 0 0 0 2 3 0 0 1 0 0 " 0 1 0 0 0 1 1 0 0 1 0 1 1 1 ? 0 2 2 1 3 0 3 0 0 0 1 1 2 3 0 0 0 0 2 3 0

NO. 41 42 43 44 45 46 47 48 49 50 51 52 53 54 55 56 57 58 59 60 Outgroup 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 Cheirolepls 2 1 2 1 0 1 l 2 0 0 0 1 1 ? 0 l 1 0 0 0 Hoythomasia 2 1 2 1 0 1 2 2 0 0 0 1 1 ? 0 1 1 0 0 0 Pa[atarraslus 2 1 2 i 0 1 2 2 i 0 0 1 1 0 0 ] 0 0 1 0 Guildayic Rdicks berger Aesoplch Discos Polypterus Rhcanoblen Tubesnt Birgerla Cramsay Doreo Pteronis P f r e t e s A m l a C d r i a n i Eaudax

3 0 0 1 0 1 2 0 0 0 1 0 2 0 0 0 0 1 0 2 2 1 2 1 0 1 2 1 0 0 0 1 1 3 I I 0 0 0 0 2 1 2 1 0 i 2 1 0 0 0 ] 1 2 1 I 0 0 0 0 2 2 2 2 0 1 2 1 0 0 0 2 2 1 0 1 0 0 0 0 3 , , 0 0 1 0 1 2 0 0 1 2 0 2 3 0 0 2 1 0 2 2 2 2 1 0 1 2 1 0 0 0 0 0 2 0 0 0 1 0 0 2 1 2 1 0 1 2 1 0 0 0 I 1 ? 0 1 0 0 0 0 2 1 2 1 0 1 2 1 0 0 0 1 1 0 0 1 1 0 0 0 2 1 2 1 0 1 2 0 0 0 0 1 1 0 0 2 1 0 1 0 2 2 2 1 0 I 2 I 0 0 0 2 1 0 0 1 0 0 1 1 2 0 2 1 0 1 2 1 0 0 0 1 1 0 0 1 0 0 0 0 2 1 2 1 0 I 2 1 0 0 0 1 1 ? 0 1 . 0 0 0 0 2 1 2 1 0 1 2 0 0 0 0 1 1 ? 0 1 0 0 1 0 2 0 1 1 0 2 2 0 0 1 0 3 0 3 0 0 0 0 0 0 2 0 1 1 0 2 2 0 " 0 1 0 3 0 3 0 0 0 0 0 0 2 0 1 1 0 2 2 0 0 1 0 3 0 3 0 0 0 0 0 0

NO. 61 62 63 64 65 66 67 68 69 70 71 72 73 Outgroup 0 0 0 0 0 0 0 0 0 0 0 0 0 C h e l r o l e p i s 0 0 0 0 0 0 ? 0 1 i 0 1 2 Hoythomasla 0 0 0 0 0 0 I 0 1 1 0 I 2 Paratar~asius 2 0 0 0 0 0 1 0 0 0 0 I 2 Guildayic Rdlcks berger Aesoplch Discos Polypterus Rhcanobien Tubesnt Dirgerla Cramsay Doreo P t e r o n i s P f r e l e s Amla C d r l a n l Eaudax

2 0 0 0 1 I 0 0 1 0 1 I 1 0 1 1 0 0 0 2 0 1 1 0 1 2 0 0 1 0 0 O 2 0 1 1 0 1 2 2 1 ? ? I ? ? 0 1 1 1 1 2 2 2 0 0 1 1 0 0 1 0 1 1 2 2 3 0 2 1 1 0 0 0 ? 1 1 ] 0 0 ? 0 0 0 ? 0 1 1 O 1 2 0 1 0 0 0 0 2 0 1 1 0 I 2 0 0 0 0 -0 0 2 0 2 1 1 1 2 I 0 0 0 0 ? ? 0 ? 1 0 I 2 i 0 0 1 0 0 ? 0 2 1 0 1 2 1 0 0 0 0 ? ? 0 1 1 0 i 2 1 0 0 0 0 ? ? 0 2 ? 0 1 2 O 0 0 O 2 1 0 2 0 1 1 1 2 0 0 0 0 2 ? ? 2 0 1 I 1 2 0 0 0 0 2 ? ? 2 0 1 I ). 2

Table 2 - Da ta ma t r ix for 20 taxa and 74 characters . Matrice de don- ndes pour 20 taxons et 74 caract~- res.

219

4

2

5

7

outgroup Polypterus bichir Guilday Discos

t j,

I j !

I i

Caturus driani Amia calva Enneles audax Paratarrasius hibbardi Birgeria groenlandica

Tubesnt Cheirolepis trailli Moythomasia nitida

Pteronisculus magna Aesopich Canobius ramsayi Boreosomus piveteaui

L L ~ Paleoniscus freieslebeni Berger Rhadinichthys canobiensis Rdicks

'~ CLADISTIA

I HOLOSTEI

PALEONISCIMORPHA

Figure 1 - Der ived charac te rs a t pr incipal nodes. N O D E 1 Actinopterygii : 8, snout, 1 - mou th t e rmina l ; 10, nasal bones, 1 - serial ; 11, r e a r of nasal(s), 1 - midorbi ta l ; 12, tectals , 2 - absen t ; 16, antorbi ta l bone, 1 - w i th infraorbi ta l canal only ; 17, supraorb i ta l ser ies , 1 - double ser ies ; 18 frontal bones, 2 - 1 pai r ; 20, pineal foramen, 2 - ab sen t ; 36, suborbi ta l bone a r range- ment , 1 - l r o w ; 37, suborbi ta l numbers , 1 - 1-3, rounded ; 41, preopercular canal, 1 - to mid-otic region ; 46, juga l line, 1 - reduced ; 47, quadra to jugal bone, 2 - absen t ; 72, nasal capsule, 1 - dorsola tera l ; 73, suspensor ium, 1 - l imited methyostyly . N O D E 2 Cladis t ia : 13, supraorb i ta l bones, 1 - s ingle ; 30, ex t rascapula r bones, 2 - 2 pairs ; 58, supraopercu la r bones, 1 - present ; 61, an te r io r branchios tegals , 2 - not below mandib le ; 65, gulars la teral , 1 - ex tended ; 66, clavicles, 1 - ab sen t ; 71, pa rasphenoid , 1 - l eng th of braincase . N O D E 4 Act inopter i : 10, nasa l bones, 2 - s ingle bone ; 19, par ie ta l / f ronta l size, 1-small ; 22, supraorb i ta l canal , 1 - to par ie ta ls ; 30, ex t rascapu la r bones, 1 - 1 pai r ; 31, profondus canal, 1 - absen t ; 34, posterodorsal infraorbi ta l shape , 1 - T-shaped ; 43, in te ropercu la r bone(s), 2 - absen t ; 52, maxil la poster ior end, 1 - modera te p la te ; 70, sclerotics, 1- 4 ; 73, suspensor ium, 2 - extensive methyostyly . N O D E 5 "Holostei" : 26, de rmosphenot ic bone size, 1 - reduced ; 43, in te ropercu la r bone(s), 1 - s ingle bone ; 46, jugal line, 2- absen t ; 50, maxi l lary mobility, 1 - p r e sen t ; 52, maxi l la poster ior end, 3 - wi th supramax i l l a ; 54, maxi l la -preopercular ganoine, 3 - absen t ; 65, la tera l gulars , 2 - absen t ; 66, clavicles, 1 - absen t ; 68, c le i thrum, 1 - na r row ; 71, pa rasphenoid , 1 - l eng th of braincase. N O D E 7 Paleonisc imorpha : 7, post ros t ra l bones, 1 - s ingle med ian ; 14, supraorb i ta l canal, 1 - be tween nasa l s ; 16, antorbi ta l bone, 2 - wi th 3 canals ; 42, p reopercu lum/squamosa l shape, 1 - "ha tche t shaped" ; 53, maxi l la ry fossa of mandible , 1 - no coronoid process, ; 67, clavicle contact , 2 - elongate. Caract~res dgrivds aux principaux noeuds.

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NIELSEN E. 1942 - S t u d i e s o n t h e T r i a s s i c f i s h e s f r o m E a s t G r e e n l a n d . I . Glaucolepis a n d Boreosomus. Pa- laeozoologica Groenlandica, 1 : 1-403.

220

NIELSEN E. 1949 - Studies on the Triassic fishes from East Greenland'. II Australosomus and Birgeria. Pa- laeozoologica Groenlandica, 3 : 1-309.

PEARSON D.M. 1982 - Pr imi t ive bony fishes, with espe- cial reference to Cheirolepis and paleonisciform ac- t inopterygians. Zoological Journal of the Linnean Society of London, 74 : 35-67.

PEHRSON T. 1947 ~ Some new interpre ta t ions of the skull of Polypterus. Acta Zoologica, 28 : 400-455.

SAINT-SEINE P. de 1949 - Les poissons des calcaires ]i thographiques de Cerin. Nouvelles archives du Musge d'Histoire Naturelle ~d e Lyon, 2 : 357 p.

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R. L U N D & K. M c C A R T H Y Department of Environmental Studies

Adelphi University Garden City NY 11530, U.S.A.

C. POPLIN Museum national d'Histoire naturelle

Laboratoire de Pal~ontologie et URA 12 du CNRS 8 rue Buffon

F-75005 Paris