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The monophyly of the Caturidae (Pisces; Actinopterygii) and the phylogeny of the Halecomorphi
THE MONOPHYLY OF THE CATURIDAE (PISCES, ACTINOPTERYGII) AND THE PHYLOGENY OF THE HAI,ECOMORPHI
PAUL H. LAMBERS LAMBERS P.H. 1995. The monophyly of the Caturidae (Pisces, Actinopterygii) and the phylogeny of the Halecomorphi. [La monophylie des Caturidae (Pisces, Actinopterygii) et la phylog~nie des Halecomorphi]. GEOBIOS, M.S. n ° 19 : 201-203. ABSTRACT The monophyly of the halecomorph fishes Caturidae and their position among the Halecomorphi is tested. Using Watsonulus as outgroup, two monophyletic groups, Furo as the sister-group of the Ophiopsidae and Amblysemius + Caturus as the sister-group of the the Amiiformes, are recognized. The Ionoscopidae are the most primitive taxon above Watsonulus. KEY-WORDS : AMIIFORMES,CATURIDAE,HALECOMORPHI,IONOSCOPIDAE, OPHIOPSIDAE, PHYLOGENY. RESUME La monophylyie des poissons hal~comorphes Caturidae et leur position parmi les Hal~comorphes ont 5t~ test~es. Avec Watsonulus comme "outgroup", deux groupes monophyl~tiques sont identifi~s : Furo comme groupe-fr~re des Ophiopsidae et Arnblysemius + Caturus comme groupe-fr~re des Amiiformes. Les Ionoscopidae sont le taxon le plus primitif apr~s Watsonulus. MOTS-CLt~S : AMIIFORMES,CATURIDAE,HALECOMORPHI,IONOSCOPIDAE, OPHIOPSIDAE, PHYLOGI~NIE.
INTRODUCTION The Caturidae Owen 1860 (Eugnathidae, Furidae) are a family of Mesozoic halecomorph fishes t h a t originally contained Caturus, Furo (= Eugnathus) and several pachycormiform genera. Patterson (1973) listed Allolepidotes, Furo, Caturus, Hetero-
lepidotes, Lophiostomus, Macrepistius, Neorhombolepis, Osteorachis, Otomitla and Sinoeugnathus in the family. Callopterus, Paraliodesmus and Plesiofuro are also classified as caturids by some authors. Patterson (1973), B a r t r a m (1975) and Schaeffer & Patterson (1984) considered the family to be a grade group, of which some of the better known forms might constitute a monophyletic group, some might be related to the Ophiopsidae and others to the Parasemionotidae. A revision of the caturid fishes from the Tithonian lithographic limestone of Solnhofen showed t h a t the old genus Amblysemius, closely related to Caturus, can be recognized (Lambers 1994).
The Caturidae, or Caturus, are regarded as the primitive sister-group of the Amiiformes (Patterson 1973 ; Schultze & Wiley 1984). It is however still unclear which genera constitute the Caturidae and which characters define this family. In this paper the results will be presented of a phylogenetic analysis including several supposed caturids, ophiopsids, some ionoscopids and some amiiforms. The monophyly of the family Caturidae and its position among the Halecomorphi will be tested. An extensive description of the genera Caturus and Amblysemius, discussion of the characters and the results of the analysis will be published elsewhere.
ANALYSIS The fishes included in the analysis are considered to be halecomorphs, because the symplectic is incorporated in the lower j a w joint (Patterson
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A F i g u r e 1 - P h y l o g e n y of h a l e c o m o r p h s , r e s u l t i n g from an analysis using P A U P 3.1.1 ; including 57 c h a r a c t e r s a n d 17 taxa, Watsonulus as outgroup. E x p l a n a t i o n of the nodes (* = reversals). A : 4, 5, 6, 9, 10, 15, 18, 19, 21, 26, 27, 35, 41, 42, 43, 44, 46, 50, 5 6 ; B : 4 6 , 4 8 , 5 5 ; C : 13, 37, 4 7 ; D : 17* ; E :12,16, 47 ; F : 37*, 57: G: 20 ; H: 21* ; I : 3, 28, 29, 34, 56 ; J : 1, 1 8 , 2 7 " , 3 0 , 31, 46, 55 ; K: 8, 28*, 29", 36, 39* ; L: 47 ; M : 45 ; N : 14, 35*, 53* ; O : 5", 22, 33, 38, 49, 51, 52, 54. E n d n o t e s : O s h u n i a : 10 ; T e o i c h t h y s : 7 ; O p h i o p s i s : 10", 25 ; O s t e o r a c h i s : 35* ; M a c r e p i s t i u s : 3, 7 ; H e t e r o l e p i d o t e s : 10, 29 ; C a l a m o p l e u r u s : 7, 9 *, 10" ; V i d a l a m i a : 7 ; S i n a m l a : 2, 15", 21", 40, 47*, 56* ; A m i a : 7; C a t u r u s : 7, 8, 19, 44* ; A r n b l y s e m i u s : 11, 21", 23, 24, 32. Relations phylogdndtiques des Haldcormophes, rdsultant d'une analyse informatisde (PAUP 3.1.1) comprenant 57 caract~res et 17 taxons avec W a t s o n u l u s utilisg comrne extragroupe. Caract~res apparaissant aux diffgrents noeuds (* = rdversion). A : 4, 5, 6, 9, 10, 15, 18, 19, 21, 26, 27, 35, 41, 42, 43, 44, 46~ 50, 56 ; B : 46, 48, 55 ; C : 13, 37, 47 ; D : 17" ; E : 12, 16, 47 ; F : 37", 57 G: 20 ; H : 21 * ; I : 3, 28, 29, 34, 56 ; J : 1, 18, 2 7 , 30, 31, 46, 55 ; K : 8, 2 8 " , 2 9 " , 36, 39" ; L : 47 ; M : 45 ; N : 14, 3 5 " , 5 3 " ; 0 5", 22, 33, 38, 49, 51, 52, 54. Carat~res prdsents chez certains taxons terminaux : O s h u n i a : 10 ; T e o i c h t h y s : 7; O p h i o p s i s 1 0 " 25 ; O s t e o r a v h i s : 35* ; M a c r e p i s t i u s : 3, 7 ; H e t e r o l e p i d o t e s : 10, 29 ; C a l a m o p l e u r u s : 7, 9 * 10" ; V i d a l a m i a : 7 S i n a m i a : 2, 1 5 , 2 1 , 40, 4 7 * 5 6 * A m i a : 7 ; C a t u r u s : 7, 8, 19, 44*; A m b l y s e m i u s : 11, 2 1 " 23, 24, 32.
1973). The halecomorph symplectic and j a w joint differs from the joint known from more primitive forms such as Pteronisculus, including a short nodular symplectic, and does not represent a primitive character-state (contra Olsen 1984 ; V6ran 1988 ; Olsen & McCune 1991). Whether the similar symplectic arrangement now known from Vinctifer (Brito 1988), pycnodonts (Nursall & Maisey 1991) and some other teleost-like forms (Arratia & Lambers in prep.) is homologous remains to be determined. Watsonulus is the primitive outgroup in the analysis, as it is regarded as the most primitive halecomorph (Patterson 1973 ; Schultze & Wiley 1984). Information not known
on the postcranial skeleton of Watsonulus is taken from Patterson's description (1973) of a Triassic parasemionotid axial skeleton. Two analyses were carried out. One included the following halecomorph genera (see textfigure) : Am-
blysemius, Amia, Amiopsis, Calamopleurus, Caturus, Furo, Heterolepidotes, Ionoscopus, Macrepistius, Ophiopsis, Oshunia, Osteorachis, Pachyamia, Sinamia, Teoichthys and Vidalamia. A second analysis included Callopterus and Neorhombolepis as well. Other genera were not included in the analyses, because they are poorly known.
203 The phylogenetic, analysis using PAUP 3.1.1, including 57 characters, results in one tree, CI=0.699 (see Fig 1). W i t h i n t h e Halecomorphi two m a i n monophyletic groups can be recognized. In one group Amblysemius a n d Caturus form a monophyletic group, being the primitive sistergroup to t h e monophyletic Amiiformes. In the other group Furo is the primitive sister-group of a monophyletic Ophiopsidae (see B a r t r a m 1975) ; the Ionoscopidae t u r n out to be the m o s t primitive t a x o n above Watsonulus. An analysis also including Callopterus a n d Neorhombolepis, results in t h r e e e q u a l l y parsimonious trees. Callopterus is the primitive sister-group to all halecomorphs above t h e Ionoscopidae, a n d Neorhombolepis eit h e r forms a t r i c h o t o m y w i t h Furo and the Ophiopsidae ; or is the sister-group of Furo + Ophiopsidae ; or Furo is the sister-group to Neorhombolepis + Ophiopsidae. List of t h e characters occurring at different nodes of the cladogram (Fig. 1) : 1, large parietals ; 2, m e d i a n p a r i e t a l s ; 3, descending l a m i n a of dermopterotic p r e s e n t ; 4, small n a s a l s ; 5, n a s a l fen e s t r a ; 6, rostral t r i a n g u l a r ; 7 (3 states) supraorbitals in two rows, more t h a n two rows, absent; 8, dermosphenotic small, plate-like ; 9, dermosphenotic in skull roof ; 10, dorsal l a m i n a on infraorbital 1 absent ; 10, sphenotic obscurred ; 11, dorsal l a m i n a on infraorbital 1 ; 12, ethmoid exposed b e t w e e n antorbital and orbit ; 13, large infraorbital in t h e corner of the orbit ; 14, last infraorbital e x p a n d e d ; 15, infraorbital I infraorbital 2/3 ; 16, deep infraorbitals ; 17, posterior border of t h e last infraorbital inclined backwards; 18, suborbitals present, region occupied by infraorbitals ; 19, lower p a r t preoperculum e x p a n d e d ; 20, canal in maxilla ; 21, posterior notch in maxilla ; 22, bar-like maxilla ; 23, low n u m b e r of m a x i l l a r y t e e t h ; 24, posterior p a r t of maxilla inclined v e n t r a l l y ; 25, v e n t r a l hook anterior on maxilla ; 26, long p r e m a x i l l a r y process ; 27, two coronoids ; 28, platform of t e e t h on the prearticular ; 29, cluster of t e e t h on the coronoid ; 30, m e n t o m e c k e l i a n obscurred ; 31, s e p a r a t e articular bones in the lower j a w ; 32, low n u m b e r of d e n t a r y t e e t h ; 33, d e r m o p a l a t i n e t e e t h ; 34, m e m b r a n e o u s outgrowths on the symplectic ; 35, i n t e r c a l a r in contact w i t h the p a r a s p h e n o i d ; 36, pterotic p r e s e n t ; 37, opisthotic p r e s e n t ; 38, par a s p h e n o i d e x t e n d i n g beyond basioccipital ; 39, vomerine t e e t h in a single row ; 40, more t h a n two s u p r a t e m p o r a l s ; 41, broad mesocoracoid ; 42, scapula r e d u c e d ; 43, coracoid anteriorly developped ; 44, s e r r a t e d appendages ; 45, dorsal fin elongate ; 46, (three states) centra including hem i c h o r d a c e n t r a , a m i d ring centra, ionoscopid cent r a ; 47, a l t e r n a t e diplospondyly ; 48, v e r t e b r a e
w i t h m e d i a n ridge ; 49, block-like ural n e u r a l arches ; 50, slender n e u r a l spines ; 51, epineural processes p r e s e n t ; 52, broad h a e m a l spines ; 53, h y p u r a l s fused to arches ; 54, ural n e u r a l spines and h a e m a l spines strongly inclined to the horizontal ; 55, caudal fin r o u n d ; 56, cycloid scales of amiid type ; 57, lateral line extending into the tail. REFERENCES
BARTRAM A.W.H. 1975 - The holostean fish genus Ophiopsis Agassiz. Zoological Journal of the Linnean Society, London, 56 : 183-205. BRITO P.M. 1988 - La structure du suspensorium de Vinctifer, poisson actinopt~rygien m~sozoique : remarques sur les implications phylog~n~tiques. Geobios, 21 : 819-823. LAMBERSP.H. 1994 - The halecomorph fishes Caturus and Amblysemius in the lithographic limestone of Solnhofen (Tithonian), Bavaria. Geobios, M.S. 16 : 91-99. NURSALL R. & MAISEYJ.G. 1991 -Neoproscinetes FIGUEIREDO • SILVA SANTOS, 1987. In MAISEY J.G., (ed.) : Santana fossils, an illustrated atlas. 125-137. T.F.H. Publications, Neptune City. OLSEN P.E. 1984 - The skull and pectoral girdle of the parasemionotid fish Watsonulus eugnathoides from the Early Triassic Sakamena Group of Madagascar, with comments on the relationships of the holostean fishes. Journal of Vertebrate Paleontology, 4 : 481-499. OLSEN P.E. & MCCUNE A.R. 1991 - Morphology of the Semionotus elegans species group from the Early Jurassic part of the Newark supergroup of Eastern North America with comments on the family Semionotidae (Neopterygii). Journal of Vertebrate Paleontology, 11 : 269-292. PATTERSON C. 1973 - Interrelationships of Holosteans. In GREENWOODP.H., MILES R.S., PATTERSON C. & COLIN (eds) : Interrelationships of fishes. Zoological Journal of the Linnean Society London 53, Supplement 1 : 233-305. SCHAEFFER B. & PATTERSON C. 1 9 8 4 - Jurassic fishes from the Western USA, with comments on Jurassic fish distribution. American Museum Novitates, 2 7 9 6 : 86 p.
SCHULTZE H.-P. & WILEY E.O. 1984 - The neopterygian Amia as a living fossil. In ELDREDGE N. 8~ STANLEY S.M. (eds.) : Living fossils. Springer Verlag, New York : 153-159. VI~RAN M. 1988 - Les ~Mments accessoires de l'arc hyo~dien des poissons t61~ostomes (Acanthodiens et Osteichthyens) fossiles et actuels. Mdmoires du Musdum national d'Histoire naturelle, sdrie C, Sciences de la Terre, 54 : 113 p. i
P . H . LAMBERS Museum fiir Naturkunde der Humbold-Universit~it Institut fiir Pal~iontologie Invalidenstrasse 43 10115 Berlin, Allemagne
PAUL H. LAMBERS LAMBERS P.H. 1995. The monophyly of the Caturidae (Pisces, Actinopterygii) and the phylogeny of the Halecomorphi. [La monophylie des Caturidae (Pisces, Actinopterygii) et la phylog~nie des Halecomorphi]. GEOBIOS, M.S. n ° 19 : 201-203. ABSTRACT The monophyly of the halecomorph fishes Caturidae and their position among the Halecomorphi is tested. Using Watsonulus as outgroup, two monophyletic groups, Furo as the sister-group of the Ophiopsidae and Amblysemius + Caturus as the sister-group of the the Amiiformes, are recognized. The Ionoscopidae are the most primitive taxon above Watsonulus. KEY-WORDS : AMIIFORMES,CATURIDAE,HALECOMORPHI,IONOSCOPIDAE, OPHIOPSIDAE, PHYLOGENY. RESUME La monophylyie des poissons hal~comorphes Caturidae et leur position parmi les Hal~comorphes ont 5t~ test~es. Avec Watsonulus comme "outgroup", deux groupes monophyl~tiques sont identifi~s : Furo comme groupe-fr~re des Ophiopsidae et Arnblysemius + Caturus comme groupe-fr~re des Amiiformes. Les Ionoscopidae sont le taxon le plus primitif apr~s Watsonulus. MOTS-CLt~S : AMIIFORMES,CATURIDAE,HALECOMORPHI,IONOSCOPIDAE, OPHIOPSIDAE, PHYLOGI~NIE.
INTRODUCTION The Caturidae Owen 1860 (Eugnathidae, Furidae) are a family of Mesozoic halecomorph fishes t h a t originally contained Caturus, Furo (= Eugnathus) and several pachycormiform genera. Patterson (1973) listed Allolepidotes, Furo, Caturus, Hetero-
lepidotes, Lophiostomus, Macrepistius, Neorhombolepis, Osteorachis, Otomitla and Sinoeugnathus in the family. Callopterus, Paraliodesmus and Plesiofuro are also classified as caturids by some authors. Patterson (1973), B a r t r a m (1975) and Schaeffer & Patterson (1984) considered the family to be a grade group, of which some of the better known forms might constitute a monophyletic group, some might be related to the Ophiopsidae and others to the Parasemionotidae. A revision of the caturid fishes from the Tithonian lithographic limestone of Solnhofen showed t h a t the old genus Amblysemius, closely related to Caturus, can be recognized (Lambers 1994).
The Caturidae, or Caturus, are regarded as the primitive sister-group of the Amiiformes (Patterson 1973 ; Schultze & Wiley 1984). It is however still unclear which genera constitute the Caturidae and which characters define this family. In this paper the results will be presented of a phylogenetic analysis including several supposed caturids, ophiopsids, some ionoscopids and some amiiforms. The monophyly of the family Caturidae and its position among the Halecomorphi will be tested. An extensive description of the genera Caturus and Amblysemius, discussion of the characters and the results of the analysis will be published elsewhere.
ANALYSIS The fishes included in the analysis are considered to be halecomorphs, because the symplectic is incorporated in the lower j a w joint (Patterson
202
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A F i g u r e 1 - P h y l o g e n y of h a l e c o m o r p h s , r e s u l t i n g from an analysis using P A U P 3.1.1 ; including 57 c h a r a c t e r s a n d 17 taxa, Watsonulus as outgroup. E x p l a n a t i o n of the nodes (* = reversals). A : 4, 5, 6, 9, 10, 15, 18, 19, 21, 26, 27, 35, 41, 42, 43, 44, 46, 50, 5 6 ; B : 4 6 , 4 8 , 5 5 ; C : 13, 37, 4 7 ; D : 17* ; E :12,16, 47 ; F : 37*, 57: G: 20 ; H: 21* ; I : 3, 28, 29, 34, 56 ; J : 1, 1 8 , 2 7 " , 3 0 , 31, 46, 55 ; K: 8, 28*, 29", 36, 39* ; L: 47 ; M : 45 ; N : 14, 35*, 53* ; O : 5", 22, 33, 38, 49, 51, 52, 54. E n d n o t e s : O s h u n i a : 10 ; T e o i c h t h y s : 7 ; O p h i o p s i s : 10", 25 ; O s t e o r a c h i s : 35* ; M a c r e p i s t i u s : 3, 7 ; H e t e r o l e p i d o t e s : 10, 29 ; C a l a m o p l e u r u s : 7, 9 *, 10" ; V i d a l a m i a : 7 ; S i n a m l a : 2, 15", 21", 40, 47*, 56* ; A m i a : 7; C a t u r u s : 7, 8, 19, 44* ; A r n b l y s e m i u s : 11, 21", 23, 24, 32. Relations phylogdndtiques des Haldcormophes, rdsultant d'une analyse informatisde (PAUP 3.1.1) comprenant 57 caract~res et 17 taxons avec W a t s o n u l u s utilisg comrne extragroupe. Caract~res apparaissant aux diffgrents noeuds (* = rdversion). A : 4, 5, 6, 9, 10, 15, 18, 19, 21, 26, 27, 35, 41, 42, 43, 44, 46~ 50, 56 ; B : 46, 48, 55 ; C : 13, 37, 47 ; D : 17" ; E : 12, 16, 47 ; F : 37", 57 G: 20 ; H : 21 * ; I : 3, 28, 29, 34, 56 ; J : 1, 18, 2 7 , 30, 31, 46, 55 ; K : 8, 2 8 " , 2 9 " , 36, 39" ; L : 47 ; M : 45 ; N : 14, 3 5 " , 5 3 " ; 0 5", 22, 33, 38, 49, 51, 52, 54. Carat~res prdsents chez certains taxons terminaux : O s h u n i a : 10 ; T e o i c h t h y s : 7; O p h i o p s i s 1 0 " 25 ; O s t e o r a v h i s : 35* ; M a c r e p i s t i u s : 3, 7 ; H e t e r o l e p i d o t e s : 10, 29 ; C a l a m o p l e u r u s : 7, 9 * 10" ; V i d a l a m i a : 7 S i n a m i a : 2, 1 5 , 2 1 , 40, 4 7 * 5 6 * A m i a : 7 ; C a t u r u s : 7, 8, 19, 44*; A m b l y s e m i u s : 11, 2 1 " 23, 24, 32.
1973). The halecomorph symplectic and j a w joint differs from the joint known from more primitive forms such as Pteronisculus, including a short nodular symplectic, and does not represent a primitive character-state (contra Olsen 1984 ; V6ran 1988 ; Olsen & McCune 1991). Whether the similar symplectic arrangement now known from Vinctifer (Brito 1988), pycnodonts (Nursall & Maisey 1991) and some other teleost-like forms (Arratia & Lambers in prep.) is homologous remains to be determined. Watsonulus is the primitive outgroup in the analysis, as it is regarded as the most primitive halecomorph (Patterson 1973 ; Schultze & Wiley 1984). Information not known
on the postcranial skeleton of Watsonulus is taken from Patterson's description (1973) of a Triassic parasemionotid axial skeleton. Two analyses were carried out. One included the following halecomorph genera (see textfigure) : Am-
blysemius, Amia, Amiopsis, Calamopleurus, Caturus, Furo, Heterolepidotes, Ionoscopus, Macrepistius, Ophiopsis, Oshunia, Osteorachis, Pachyamia, Sinamia, Teoichthys and Vidalamia. A second analysis included Callopterus and Neorhombolepis as well. Other genera were not included in the analyses, because they are poorly known.
203 The phylogenetic, analysis using PAUP 3.1.1, including 57 characters, results in one tree, CI=0.699 (see Fig 1). W i t h i n t h e Halecomorphi two m a i n monophyletic groups can be recognized. In one group Amblysemius a n d Caturus form a monophyletic group, being the primitive sistergroup to t h e monophyletic Amiiformes. In the other group Furo is the primitive sister-group of a monophyletic Ophiopsidae (see B a r t r a m 1975) ; the Ionoscopidae t u r n out to be the m o s t primitive t a x o n above Watsonulus. An analysis also including Callopterus a n d Neorhombolepis, results in t h r e e e q u a l l y parsimonious trees. Callopterus is the primitive sister-group to all halecomorphs above t h e Ionoscopidae, a n d Neorhombolepis eit h e r forms a t r i c h o t o m y w i t h Furo and the Ophiopsidae ; or is the sister-group of Furo + Ophiopsidae ; or Furo is the sister-group to Neorhombolepis + Ophiopsidae. List of t h e characters occurring at different nodes of the cladogram (Fig. 1) : 1, large parietals ; 2, m e d i a n p a r i e t a l s ; 3, descending l a m i n a of dermopterotic p r e s e n t ; 4, small n a s a l s ; 5, n a s a l fen e s t r a ; 6, rostral t r i a n g u l a r ; 7 (3 states) supraorbitals in two rows, more t h a n two rows, absent; 8, dermosphenotic small, plate-like ; 9, dermosphenotic in skull roof ; 10, dorsal l a m i n a on infraorbital 1 absent ; 10, sphenotic obscurred ; 11, dorsal l a m i n a on infraorbital 1 ; 12, ethmoid exposed b e t w e e n antorbital and orbit ; 13, large infraorbital in t h e corner of the orbit ; 14, last infraorbital e x p a n d e d ; 15, infraorbital I infraorbital 2/3 ; 16, deep infraorbitals ; 17, posterior border of t h e last infraorbital inclined backwards; 18, suborbitals present, region occupied by infraorbitals ; 19, lower p a r t preoperculum e x p a n d e d ; 20, canal in maxilla ; 21, posterior notch in maxilla ; 22, bar-like maxilla ; 23, low n u m b e r of m a x i l l a r y t e e t h ; 24, posterior p a r t of maxilla inclined v e n t r a l l y ; 25, v e n t r a l hook anterior on maxilla ; 26, long p r e m a x i l l a r y process ; 27, two coronoids ; 28, platform of t e e t h on the prearticular ; 29, cluster of t e e t h on the coronoid ; 30, m e n t o m e c k e l i a n obscurred ; 31, s e p a r a t e articular bones in the lower j a w ; 32, low n u m b e r of d e n t a r y t e e t h ; 33, d e r m o p a l a t i n e t e e t h ; 34, m e m b r a n e o u s outgrowths on the symplectic ; 35, i n t e r c a l a r in contact w i t h the p a r a s p h e n o i d ; 36, pterotic p r e s e n t ; 37, opisthotic p r e s e n t ; 38, par a s p h e n o i d e x t e n d i n g beyond basioccipital ; 39, vomerine t e e t h in a single row ; 40, more t h a n two s u p r a t e m p o r a l s ; 41, broad mesocoracoid ; 42, scapula r e d u c e d ; 43, coracoid anteriorly developped ; 44, s e r r a t e d appendages ; 45, dorsal fin elongate ; 46, (three states) centra including hem i c h o r d a c e n t r a , a m i d ring centra, ionoscopid cent r a ; 47, a l t e r n a t e diplospondyly ; 48, v e r t e b r a e
w i t h m e d i a n ridge ; 49, block-like ural n e u r a l arches ; 50, slender n e u r a l spines ; 51, epineural processes p r e s e n t ; 52, broad h a e m a l spines ; 53, h y p u r a l s fused to arches ; 54, ural n e u r a l spines and h a e m a l spines strongly inclined to the horizontal ; 55, caudal fin r o u n d ; 56, cycloid scales of amiid type ; 57, lateral line extending into the tail. REFERENCES
BARTRAM A.W.H. 1975 - The holostean fish genus Ophiopsis Agassiz. Zoological Journal of the Linnean Society, London, 56 : 183-205. BRITO P.M. 1988 - La structure du suspensorium de Vinctifer, poisson actinopt~rygien m~sozoique : remarques sur les implications phylog~n~tiques. Geobios, 21 : 819-823. LAMBERSP.H. 1994 - The halecomorph fishes Caturus and Amblysemius in the lithographic limestone of Solnhofen (Tithonian), Bavaria. Geobios, M.S. 16 : 91-99. NURSALL R. & MAISEYJ.G. 1991 -Neoproscinetes FIGUEIREDO • SILVA SANTOS, 1987. In MAISEY J.G., (ed.) : Santana fossils, an illustrated atlas. 125-137. T.F.H. Publications, Neptune City. OLSEN P.E. 1984 - The skull and pectoral girdle of the parasemionotid fish Watsonulus eugnathoides from the Early Triassic Sakamena Group of Madagascar, with comments on the relationships of the holostean fishes. Journal of Vertebrate Paleontology, 4 : 481-499. OLSEN P.E. & MCCUNE A.R. 1991 - Morphology of the Semionotus elegans species group from the Early Jurassic part of the Newark supergroup of Eastern North America with comments on the family Semionotidae (Neopterygii). Journal of Vertebrate Paleontology, 11 : 269-292. PATTERSON C. 1973 - Interrelationships of Holosteans. In GREENWOODP.H., MILES R.S., PATTERSON C. & COLIN (eds) : Interrelationships of fishes. Zoological Journal of the Linnean Society London 53, Supplement 1 : 233-305. SCHAEFFER B. & PATTERSON C. 1 9 8 4 - Jurassic fishes from the Western USA, with comments on Jurassic fish distribution. American Museum Novitates, 2 7 9 6 : 86 p.
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