Relationships of †Codoichthys carnavalii Santos, 1994 (Teleostei, Clupeomorpha, †Ellimmichthyiformes) from the Late Aptian of São Luís-Grajaú Basin, NE Brazil

ABSTRACT

†Codoichthys carnavalii is a clupeomorph fish only found in calcareous concretions of Codó Formation, State of Maranhão. It is known based on three specimens housed in the paleontological collection of the Museu de Ciências da Terra of Departamento Nacional da Produção Mineral, Rio de Janeiro. It was omitted in most of recent cladistic analyses about clupeomorphs. We revisited its anatomy furnishing new data and additional restorations. Furthermore we explored the relationships of †Codoichthys with the computer program TNT based on a matrix with 30 taxa and 60 unordered and unweight characters. Elops was used to root the tree. The strict consensus was obtained from three shortest trees (L=181; CI=0.387; RI=0.632). The monophyly of †Ellimmichthyiformes is supported by a sigmoid cleithrum and an uniquely derived predorsal scute series. Most of subgroups showed low support indices. †Sorbinichthyidae and †Horseshoeichthys appear in the most basal position, and not closely related to †Armigatus or †Diplomystus. A †Diplomystus clade is more advanced than †Armigatus and sister-group of remaining †ellimmichthyiforms. Within †Paraclupeidae, †Codoichthys is sister-group of remaining †paraclupeids (including †thorectichthyines and †paraclupeines). Within †Paraclupeinae, †Triplomystini includes a †Triplomystus clade, a sister group of †Rhombichthys plus †Tycheroichthys, and †Paraclupeini with †S. itapagipensis and all other †ellimmichthyiform taxa.

Key words:
†Codoichthys; São Luis-Grajaú Basin; Lower Cretaceous; systematics; Northeastern Brazil.

INTRODUCTION

Clupeomorpha is a great radiation of teleostean fishes that produced two major lineages: Clupeiformes and †Ellimmichthyiformes (Grande 1985). The former consists of 84 genera (Nelson 2006) and 397 living species (Lavoué et al. 2014) of fishes popularly known as herrings, sardines and anchovies together with many fossil taxa known since the Barremian (Figueiredo 2009). The latter is an archaic group of double-armored herring-like fishes including about 13 genera and at least 35 species found in many marine, estuarine, and freshwater deposits around the world ranging from the Hauterivian-Barremian (Santos and Corrêa 1985) to Middle Eocene (Patterson 1993).

Grande (1982, 1985) was the first to perceive the existence of †Ellimmichthyiformes. He intuitively indicated monophyly of the group on the basis of derived presence of subrectangular predorsal scutes. Unlike ordinary scales, he noted that these unpaired scutes are positioned at the dorsal midline and they are heavily ossified, often covered with cycloid scales. He also pointed out that many taxa of distant lineages of teleosts bear some type of dorsal scutes (e.g., many aulopiforms) and these scutes are commonly preserved in fossil fishes unlike true scales.

At first only †Diplomystus and †Ellimmichthys were placed within †Ellimmichthyiformes (Grande 1982) that also originally contained a single family (i.e.,†Ellimmichthyidae). Afterwards, due to nomenclatural priority, †Paraclupeidae was indicated by Chang and Grande (1997) as replacement name for this family. Since then new diagnoses, definitions and compositions of putative monophyletic subunits were proposed, and new non-clupeiform taxa coming from many fossil localities throughout the world were assigned to †Ellimmichthyiformes. Many phylogenetic analyses were carried out (e.g., Chang and Maisey 2003, Zaragueta-Bagils 2004, Alvarado-Ortega et al. 2008, Murray and Wilson 2013) but until now the monophyly of the group remains uncertain as well as its content.

The following nominal species of †Ellimmichthyiformes are known from Brazilian strata: †Ellimmichthys longicostatus from Hauterivian-Barremian non-marine deposits of Bahia (Longbottom 1988, Silva 1993, Carvalho and Figueiredo Souto 2005), †Ellimmichthys maceioensis from Aptian-Albian shales of Maceio Formation in Sergipe-Alagoas basin (Malabarba et al. 2004), †Ellimma branneri from Hauterivian to Aptian deposits of northeastern Brazil (Maisey 2000, Chang and Maisey 2003), †Ellimma cruzae from Albian-Aptian calcareous concretions of Cabo Formation (Santos 1990), †Codoichthys carnavalli from the Late Aptian of Codó Formation (Santos 1994), and †Scutatuspinosus itapagipensis from Hauterivian-Barremian shales of Marfim Formation of the Reconcavo Basin (Santos and Corrêa 1985). But the number must increase since undescribed material of †Ellimmichthys-like and †Ellimma-like fishes coming from the Lower Cretaceous of Bahia (e.g., Marizal and Candeias formations, Gallo and Figueiredo 2002), Upper Cretaceous (Turonian) of Pelotas Basin (Gallo et al. 2006), and Upper Cretaceous of the Santos Basin (salt beds of so-called Brazilian pre sal) are pending for description.

The fish fauna from Reconcavo Basin is mainly distributed in outcrops of Santo Amaro Group, particularly in the Itaparica, Candeias and Maracangalha formations (Carvalho and Figueiredo Souto 2005). †E. longicostatus was studied by Cope (1886) after material collected by the geologist Joseph Mawson in localities near Salvador and Simões Filho (Carvalho and Figueiredo Souto 2005). Santos (1949) stated that †Ellimmichthys longicostatus was collected together with remains of †Lepidotes, †Cladocyclus, †Calamopleurus and †Mawsonia by the paleontologist Lewellyn Ivor Price and Abel Oliveira, in Ilha de Itaparica. More recently, †E. longicostatus was collected in shales of Maracangalha Formation (Carvalho and Figueiredo Souto 2005).

Chang and Maisey (2003) while revisiting †Ellimma branneri considered only †E. branneri and †E. cruzae (=E. cruzi) as valid species for this genus. They indicated that †Ellimma branneri only partially agree with the diagnosis of †Paraclupeidae because anterior dorsal scutes are longer than broad unlike the posterior ones, broader than long. Thus they suggested a reassessment of characters supporting the monophyly of this family.

At first †E. cruzae (=E. cruzi) was identified as †Ellimmichthys longicostatus (Costa et al. 1979) but Santos (1990) assigned it to †Ellimma and named †Ellimma cruzi in honor of the paleontologist Norma Maria da Costa Cruz, from the staff of DNPM, Rio de Janeiro. We note the ending of the specific epithet is an incorrect original spelling based on an inadvertent mistake (ICZN 1999: art. 32.5.1), and should be corrected to the feminine †E. cruzae.

Murray and Wilson (2013) based on 24 taxa and 62 characters furnished the most recent phylogenetic review on †Ellimmichthyiformes. They divided the group in two major clades: †Armigatoidei (for †Armigatus and †Diplomystus) and †Ellimmichthyoidei (for remaining taxa). They tentatively named many subunits excluding or adding taxa. Thus †Paraclupeidae was divided in five subfamilies: †Scutatuspinosinae, †Thorectichthyinae, †Ellimminae, †Ellimmichthyinae, and †Paraclupeinae. Notwithstanding the relevance to the knowledge of the relationships of †ellimmichthyiform fishes, many problems still remain particularly regarding the phylogenetic status of the Brazilian taxa †Ellimma cruzae, †Scutatuspinosus itapagipensis and †Codoichthys carnavalii.

Certainly †Codoichthys carnavalli is one of the most puzzling clupeomorph taxa described from Brazil. At first Santos, in 1945, considered it a species of †Knightia. Afterwards the same author (Santos 1994), based on comprehensive revisions by Grande (1982, 1985), classified it as Clupeomorpha incertae sedis although recognizing putative affinities with non-clupeiform clupeomorphs as †Armigatus, †Ellimmichthys, and †Diplomystus. Since then no rigorous attempt to decipher its relationships was undertaken.

Our goal in this paper is to describe in detail the morphology of †Codoichthys carnavalii as soon as the material permits and exploring its relationships within Clupeomorpha using cladistic methodology.

MATERIALS AND METHODS

MATERIAL

The specimens of †Codoichthys carnavalli herein studied belong to the paleontological collection of the Museu de Ciências da Terra of the Departamento Nacional de Produção Mineral, Rio de Janeiro, and consists of the type-series (see details in Santos 1994). They are referred with the abbreviation DGM followed by the institutional register number.

According to Santos (1994) the fishes were collected by the late geologists Odorico Albuquerque and Victor Dequech, even belonging to the Divisão de Geologia e Mineralogia of DNPM, in 1945, in the locality of Morro de Umburanas, State of Maranhão, northeastern Brazil (Fig. 1). All specimens examined are poorly preserved and laterally compressed. They are preserved in yellowish calcareous concretions and were previously mechanically prepared with steel needles of different sizes under dissecting microscope by Rubens da Silva Santos. We produced painted silicone peels from the type-series specimens to enhance anatomical features. Camera lucida drawings were made using a Motic-Quimis stereomicroscope with a drawing-tube attachment. Digital photographs of high resolution were obtained with a USB Camera under MIAS(r) Software and photograph camera Nikon D7100. For enhancing anatomical details and reducing bright during photograph sessions fossils were coated with a sublimate of magnesium oxide. Anteriormost vertebrae hidden by opercle were counted as three according to Grande (1985). Measurements and meristic counts follow Forey et al. (2003). Proportions are presented as a percentage of the standard length (SL). A "dagger" (†) preciding taxon indicates that it is known only by fossils.

Figure 1
Location map of Cretaceous fish-bearing beds cropping out in southeastern Maranhão (i.e., Barra do Corda, Codó e Brejo) after Santos (1994).

GEOLOGICAL SETTING

The São Luís - Grajaú Basin is included in the Gurupe Graben System and encompasses an extensive area of about 250,000 km² in the center-north part of the State of Maranhão. It is a typical marginal rift basin, with predominantly Cretaceous filling. Codó Formation is in the lowermost sequence of Cretaceous deposits reaching of about 4,000m thick in the depocenters (Rossetti et al. 2004). This formation was deposited in a pre-rift phase and forms a bedding sequence of about 180 m through an area extending from Araguatins, Tocantins State, to Brejo Municipality, northeastern Maranhão. It is interspersed by Grajaú (below) and Itapecurú (above) formations and constituted mainly of lacustrine dark shales, evaporites and calciferous shales which include calcareous concretions and gypsum lenticular bedding (Santos 1994). There are two facies (Santos and Carvalho 2009): one lacustrine (lower unit), with †Dastilbe elongatus and other fossils also found in the Crato Formation of the Araripe Basin, and other estuarine (upper unit), represented by calcareous concretions from which Santos (1994) pointed out the occurrence of †Codoichthys carnavalii in association to fishes commonly found in Romualdo Member of Araripe Basin, for instance, †Tharrias araripis, †Calamopleurus cylindricus, †Brannerion latum, †Vinctifer comptoni, †Santanichthys diasii, †Rhacolepis buccalis, †Cladocyclus gardneri, and †Araripelepidotes temnurus.

The localities in the State of Maranhão from which Cretaceous fossil fishes were found are plotted in map (Fig. 1). According to Santos (1994) these localities are Morro de Umburanas in Brejo, Codó and Barra do Corda.

CLADISTIC ANALYSIS

We used cladistic principles (e.g., Wiley and Lieberman 2011, Wheeler 2012) and underlying computational technology to explore the phylogenetic relationships of †Codoichthys carnavalii together with other clupeomorphs. The analysis was performed with TNT computer program version 1.1 (Goloboff et al. 2003) to generate the shortest trees based on a polarized data matrix (Table I) composed of 30 taxa and 60 unordered and unweight selected characters taken from foregoing cladistics analyses of the †ellimmichthyiforms (i.e., Chang and Maisey 2003, Forey 2004, Zaragueta-Bagils 2004, Alvarado-Ortega et al. 2008, Murray and Wilson 2013). Only characters suggestive of immediate common ancestry (synapomorphic) are accepted as criteria for the recognition of monophyletic groups(=clades). We excluded some numerical characters of Alvarado-Ortega et al (2008) and Murray and Wilson (2013) because they introduce subjective procedure of cut a continuum when data of new taxa are added to the matrix, always producing new arrangements to accomodate them. Missing characters or unclear states owing to the quality of preservation (including ambiguous data) were coded as "?" in the matrix. ACCTRAN was the optimization choice due to the preservation of primary homology (de Pinna 1991). The tree-building routine was traditional search with random stepwise addition, ten trees held at each iterative step, and TBR (tree bisection and reconnection) branch swapping routine. Branches without support were collapsed.

TABLE I
Data Matrix

Taxon	1-5	6-10	11-15	16-20	21-25	26-30	31-35	36-40	41-45	46-50	51-55	56-60
†Armigatus alticorpus	00010	??00?	1????	??00?	10?1?	??00?	00000	???10	0000?	?1?00	01???	000?0
†Armigatus brevissimus	00010	12001	1001?	1000?	10110	?0001	00000	10010	00000	?1200	01010	000?0
†Armigatus namourensis	00010	12001	11?10	10000	10110	?0001	00000	11010	00001	11200	011?1	000?0
Chirocentrus dorab	01000	01100	21000	01101	01111	10101	10100	0100-	- - - --	-12--	0 -??0	00000
†Codoichthys carnavalii	00110	??00?	2?0??	01?0?	00110	00001	00001	?1010	00000	01100	000?0	01101
Denticeps clupeoides	01000	00110	20001	11201	00210	?1021	01010	0010-	- - - --	-100-	00100	00000
†Diplomystus birdi	00110	?2001	1?0??	???00	11110	0?001	0000?	??111	11000	21?00	011?1	000??
†Diplomystus dentatus	00110	?2001	11?10	11200	11110	10001	00000	?1111	11000	21100	02011	000?1
†Diplomystus dubertreti	00110	?200?	?????	?1?0?	11110	0000?	0?00?	???11	11000	21100	0?1??	000??
†Diplomystus shengliensis	00110	??001	11???	1?10?	10010	00000	00001	10?11	11100	211?0	0?110	000??
†Diplomystus solignaci	101??	??001	?????	1111?	10110	?000?	01000	???11	1?1??	?11??	02???	0?0??
†Ellimma branneri	10011	?0001	20?1?	01110	11110	00000	00000	10010	10111	01110	01110	011??
†Ellimmichthys goodi	1?011	??00?	2????	11111	11110	00000	0000?	1??10	10111	01110	0211?	0?000
†Ellimmichthys longicostatus	10011	?000?	2??1?	11111	11110	00001	00000	10110	10111	?1110	02111	010?0
Elops saurus	00000	00000	00000	10000	00000	00000	00000	1010-	- - - --	-00- -	0-000	00000
†Ezkutuberezi carmenae	10??0	??00?	2????	0?200	1?110	00100	00010	?0010	10111	?1010	011?1	0??0?
†Horseshoeichthys armaserratus	00000	?000?	2?0??	?0?01	10???	?????	??0??	???1?	?0???	?1???	0?0??	?0???
Odaxothrissa vittata	01000	01110	20000	01001	00111	10101	11100	0100-	- - - --	-1000	00010	00000
†Ornategulum sardinoides	00010	?0000	2001?	10000	00010	00000	00000	0100-	- - - --	-00- -	00000	00000
†Paraclupea chetungensis	10011	??001	2??1?	1?11?	10110	10000	01001	00111	01111	11111	02111	00000
†Rhombichthys intoccabilis	100??	??001	2??1?	011?0	10210	00011	00100	0001?	?0111	01121	02111	1000?
†Scutatuspinosus itapagipensis	00010	??00?	20010	01100	11?10	00001	0?000	10010	10111	01112	011?1	10001
†Sorbinichthys africanus	0?00?	?00?1	20???	1010?	01200	0?101	010??	?0010	?110?	21000	00??0	0000?
†Sorbinichthys elusivo	0010?	??001	??110	00100	11200	00000	000??	0?010	?110?	21000	000?0	00001
†Thorectichthys marocensis	10011	??001	2?110	11100	10100	00000	01000	11110	00010	01200	01011	00?0?
†Thorectichthys rhadinus	100?0	??0?1	2??10	11100	10100	00000	01000	1?110	00010	01210	01011	0000?
†Triplomystus applegatei	1?010	??0?1	2????	011??	11?1?	10011	00000	1111?	10111	01211	10010	?000?
†Triplomystus noorae	10010	12001	211??	1110?	10110	10010	00000	10010	10111	01201	12010	00011
†Triplomystus oligoscutatus	10010	12001	211??	?110?	1?110	1001?	00000	1??10	1011?	?1??1	1101?	0001?
†Tycheiroichthys dunveganensis	10000	??001	2?1??	01100	1111?	?00??	0?0?1	00010	0011?	11121	020?0	0011?
Missing data(?), Inapplicable (-)

Selected †Ellimmichthyiformes used in analysis are listed below together with literature review from which anatomical data are better known and discussed. Characters have been taken mainly from literature and checked (if possible), and in the case of changes they are indicated in our list (see ^{Appendix 1}). Additional fossil specimens housed in institutional collections from which we made original observations or confirmed previous data are also indicated.

The fossil material is the following: †Armigatus alticorpus - all data are from Forey et al. (2003); †Armigatus brevissimus - n= 5, uncatalogued Pz. UERJ, data from Patterson (1967) and Grande (1982); †Armigatus namourensis - all data are from Forey et al. (2003); †Diplomystus birdi - all data from Grande (1982); †Diplomystus dentatus - n= 2, uncatalogued Pz.UERJ from Green River Formation, Wyoming; complementary data are taken from Cavender (1966) and Grande (1982); †Diplomystus shengliensis - all data taken from Chang and Maisey (2003); †Diplomystus solignaci - all data taken from Gaudant and Gaudant (1971) and Grande (1982); †Diplomystus dubertreti - data are from Signeux (1951) and Grande (1982); †Ellimma branneri - data are from Chang and Maisey (2003) and Pz.UERJ 77 and 95;†Ellimmichthys longicostatus - data are from Grande (1982), Chang and Grande (1997), Chang and Maisey (2003), and BMNH P. 7109; †Ellimmichthys goodi - data are taken from Eastman (1912) and Chang and Grande (1997); †Ezkutuberezi carmenae - all data are from Poyato-Ariza et al. (2000); †Horseshoeichthys armiserratus - all data are from Newbrey et al. (2010); †Paraclupea chetungensis - all data are from Chang and Grande (1997); †Rhombichthys intoccabilis - all data are from Khalloufi et al. (2010); †Scutatuspinosus itapagipensis: n=72, DGM 1164-P (holotype) to 1176-P, DGM 1207-P to 1264-P, complementary data are from Santos and Corrêa (1985) and Alvarado-Ortega et al. (2008); †Sorbinichthys elusivo - all data taken from Bannikov and Bacchia (2000); †Sorbinichthys africanus - all data taken from Murray and Wilson (2011);†Tycheiroichthys dunveganensis - all data are from Hay et al. (2007); †Thorectichthys marocensis - all data are from Murray and Wilson (2013); †Thorectichthys rhadinus - all data taken Murray and Wilson (2013); †Tryplomystus noorae - all data taken from Forey et al. (2003); †Tryplomystus oligoscutatus (all data taken from Forey et al. (2003); and †Tryplomystus applegatei - all data are from Alvarado-Ortega and Ovalles-Damián (2008).

We have not included in this analysis data of †Ellimma cruzae, †Kwangoclupea dartevellei, and †Ellimmichthys maceioensis because we think that their anatomy demands more study so that they will be reassessed elsewhere (work in progress). We include the living clupeiforms Chirocentrus dorab (data taken of Cavender 1966, Fujita 1990, Di Dario 2009), Denticeps clupeoides (data taken of Greenwood 1968, Di Dario and de Pinna 2006), Odaxothrissa vittata (data taken of Grande 1985), and the primitive clupeomorph †Ornategulum sardinoides (all data from Forey 1973a) as near outgroups together with the elopomorph Elops saurus (D. UERJ 190, n=15, 150-550 mm SL). To maximize applicable states of characters, Elops saurus was choiced as remote outgroup. This species was used for rooting the tree because this taxon is considered a morphologically generalized teleost whose anatomy is relatively well-known (see Ridewood 1904, Vrba 1968, Nybelin 1967, 1968, 1971, Forey 1973b, Taverne 1974).

MORPHOMETRIC ABBREVIATIONS

BD, body depth; HD, head depth; HL, head length; PANL, preanal length; PDL, predorsal length; SL, standard length.

ANATOMICAL ABBREVIATIONS

AA, anguloarticular; abd. sc, abdominal scute; an. PTG, pterygiophores of anal fin; ANT, antorbital; APAL, autopalatine; ASMX, anterior supramaxilla; c.sc; caudal scute; CL, cleithrum; CO, coracoid; D, dentary; EP, epural; EPL, epipleural intermuscular bone; EPM, epimeral intermuscular bones; EPN, epineural intermuscular bone; fac. MX APA, autopalatine facet for articulation with maxillary; f.r, fin ray; FR, frontal; H, hypural; h.sp, hemal spine; IO, infraorbital; io.c, infraorbital sensory canal; LA, lachrymal; lat.pr, lateral process of mesethmoid; LET, lateral ethmoid; md.c, mandibular sensory canal; MES, mesethmoid; MX, maxilla; n. sp, neural spine; n. sp. PU2; neural spine of the second preural centrum; n.a, neural arch; n.a. PU1, neural arch of the first preural centrum; OP, opercle; PA, parietal; pap, parapophysis; PAS, parasphenoid; PH, parhypural; pl.r, pleural rib; PMX, premaxilla; POP, preopercle; PSMX, posterior supramaxilla; PTM, posttemporal; PU1, first preural centrum; RAR, retroarticular; S, symplectic; SCL, supracleithrum; scl.b, sclerotic bone; SOC, supraoccipital; SOP, subopercle; stt.com, commissural supratemporal sensory canal; U1+H2, first ural centrum fused to second hypural; UN, uroneural; VC, vertebral centrum.

INSTITUTIONAL ABBREVIATIONS

BMNH P, formerly British Museum of Natural History, London; DGM, formerly Divisão de Geologia e Mineralogia do Departamento Nacional de Produção Mineral, nowadays Museu de Ciências da Terra; D. UERJ, a collection of cleared and stained fishes in Departamento de Zoologia of Universidade do Estado do Rio de Janeiro; Pz.UERJ, paleozoological collection of the Universidade do Estado do Rio de Janeiro.

SYSTEMATIC PALEONTOLOGY

Subdivision TELEOSTEI Müller, 1845

Cohort CLUPEOCEPHALA Patterson and Rosen, 1977

Subcohort OTOCEPHALA Johnson and Patterson, 1996

Superorder CLUPEOMORPHA Greenwood et al., 1966

Order †ELLIMMICHTHYIFORMES Grande, 1982

Family †PARACLUPEIDAE Chang and Chou, 1977

Diagnosis (amended from Santos 1994). Fusiform fish reaching 62 mm SL showing the following combination of features: HL equals 31% SL; predorsal dorsal profile slightly convex; anamestic antorbital present; medially united parietals; rugose ornamentation on frontal and parietal; well-developed supraoccipital crest; upper jaw and parasphenoid toothless; dentary bearing a patch of minute conical teeth on oral border and prominent coronoid process; two supramaxillae; quadrate-mandibular articulation placed below in the middle of the orbit; dorsal preopercle limb longer than ventral one, two recumbent tubules of preopercular sensory canal on ventral limb of preopercle; opercle equals 33% HL and ornamented with parallel striae on the lower half; L-like cleithrum with expanded posterior lamina; two postcleithra; predorsal scute series complete and composed of at least eight equal-sized keeled elements, all smooth and ovoid; at least seven curved and slender supraneurals; 33 preural vertebrae from which 11 caudal; epineurals fused to abdominal vertebrae; dorsal fin with at least 10 pterygiophores; at least 15 pterygiophores on anal fin; pelvic fin placed in opposition to dorsal fin; 10 prepelvic and nine postpelvic scutes; postpelvic scutes without pungent posterior process; vertebral column blending gradually upwards in the caudal region; three epurals; long neural spine of second preural centrum; parhypural fused to first preural centrum; large leaf-like neural arch of first preural centrum; the first uroneural reaching first preural centrum; six hypurals, the first one slender and with proximal condylar end contacting first ural centrum; second hypural slender and fused to first ural centrum; third hypural large and triangular; caudal diastema present; first ural centra almost equal-sized to preural centra; equal-lobed caudal fin.

TYPE AND ONLY SPECIES: †Codoichthys carnavaliiSantos, 1994.

†Codoichthys carnavaliiSantos, 1994

(Figs. 2- 8)

Figure 2
a. †Codoichthys carnavalliSantos, 1994, DGM 435-P (Holotype), 62 mm SL, from marine deposit of the Late Aptian of Codó Formation, São Luís-Grajaú Basin, Northeastern Brazil. b. Restoration of entire fish based on holotype and complemented with data of DGM 436-P.

Figure 3
a. Cranial bones and pectoral girdle †Codoichthys carnavaliiSantos, 1994, from the Late Aptian of Codó Formation, São Luís-Grajaú basin, State of Maranhão, Northeastern Brazil; DGM 435-P (Holotype); reversed. b. Line drawing of the same and interpretative reconstruction.

Figure 4
a. Ethmoid region and associated structures of †Codoichthys carnavalii, DGM 435-P (Holotype); b. Line drawing and interpretative restoration of ethmoid region.

Figure 5
Photograph of skull roof of †Codoichthys carnavalii, DGM 435-P (Holotype) showing vestiges of ornamentation (indicated with white arrows).

Figure 6
Line drawing of posteriormost abdominal vertebrae, anal fin support, and associated structures of †Codoichthys carnavalii, DGM 435-P (Holotype).

Figure 7
Detail of dorsal scutes and supraneural bones of †Codoichthys carnavalii, DGM 435-P (Holotype). Arrows indicate supraneurals; scutes are numbered.

Figure 8
†Codoichthys carnavalli. a. Caudal endoskeleton of DGM 435-P (holotype); b. Line drawing of caudal endoskeleton of holotype. Arrows indicate position of outer principal fin-rays.

Selected Synonymy:

1945. †Knightia carnavalii Santos [nomen nudum in a draft sent to II Congresso Pan-Americano de Engenharia de Minas e Geologia, Petrópolis].

1994. †Codoichthys carnavalii Santos, An. Acad. Bras. Cienc. 62(3), p. 133, Text-figure 2, Pl. 1, Figs. 1-3 [original description; type-locality: Morro de Umburanas].

2000. †Codoichthys carnavalii Santos: Maisey, Cretaceous Res. 21, p. 295 [only reference].

2000. †Codoichthys carnavalii Santos: Poyato-Ariza, López-Horgue and García-Garmilla: Cretaceous Res. 21, p.581 [only reference].

2009. †Codoichthys carnavalii Santos: Santos and Carvalho, Serv. Geol. Bras., fig. 13.4 [data about taphonomy and paleoenvironment ].

Diagnosis. - as for genus, monotypic genus.

Material examined. - Holotype: DGM 435-P; Paratypes: DGM 436-P; DGM 966-P.

Locality and geological time. - Morro de Umburanas, Brejo Municipality, State of Maranhão, Lower Cretaceous (Aptian) of São Luís - Grajaú Basin (see Rossetti et al. 2004, and particularly, Santos 1994).

DESCRIPTIVE MORPHOLOGY

GENERAL BODY FORM

†Codoichthys carnavalii shows fusiform body, with predorsal profile rather arched (Figs. 2a, b). All specimens are preserved in lateral view suggesting that this fish was laterally compressed in life. HL corresponds to 28% of SL and almost equal to HD. Snout is short and upper jaw not extending behind the orbit. Both dorsal and ventral profile are slightly convex. BD corresponds to 27% SL. PDL is of about half (54%) of SL. Dorsal fin is placed at the middle point of the body, in opposition to the insertion of pelvic fin. PANL is 72% of SL and the base of anal fin length is about 16% of SL. The caudal fin is forked, with deep notch, showing equal-sized dorsal and ventral lobes.

Cranial bones. - Most of skull bones (Fig. 3a, b) are badly or incompletely preserved; hence interpretative restorations are furnished as far as the material permits. In the ethmoid region there is a short and robust mesethmoid (Figs. 3b, 4, MES) with prominent cup-like lateral process for palatine (Fig.4, lat.pr). Its length is almost equal the underlying lachrymal bone. Anteriorly mesethmoid shows a short recess to house the anteriormost end of maxilla.

The lateral ethmoid (Figs. 3b, 4, LET) is represented by two incomplete and imperfect separated portions. One corresponds to a fragment of the main body of left ethmoid lateral which was slightly displaced to occupy the center of a broad gape (a probable condriferous space of nasal pit in life) covered by mesethmoid. Other portion corresponds to a well-developed fan-like shield of perichondral bone associated to a cup-like upper edge (for meeting frontal bone dorsally) of the right ethmoid lateral compounding the anterior limit of the orbit. Ventrally, this bone produces a wing-like outgrowth to touch the shaft of the parasphenoid.

The parasphenoid (Fig. 3b, PAS) is a long, low, and slightly curved bone. Only its orbital portion is visible. It is toothless. Due to state of preservation, we are unable to determine presence of a dermal basipterygoid process.

The frontal (Figs. 3b, 5, FR) is the largest bone of the cranial roof covering completely the orbit and finishing a little behind it. It is narrower at the level of the ethmoid region and broadens considerably at the posterodorsal orbital corner. Close to the contact zone with parietal this bone is ornamented with coarse rugae. Due to preservation it is difficult to determine presence of fontanels or fossae. The supraorbital sensory canal run bone enclosed so that its presence on surface is hardly noted through a faint tubular relief. Pores are not observed.

The meeting between frontal and parietal is through a long transverse suture. There is an ascending profile anterior to the well-developed supraoccipital crest as in the skull of the unnamed †Diplomystus from the Cenomanian of the English Chalk (see Forey 2004) in contrast with †Ellimma branneri and †Paraclupea. Deep grooves for supraorbital sensory canal separated by a medial bony bridge as seen in the unnamed †Diplomystus from the English Chalk and Scutatuspinosus itapagipensis are lacking.

The parietal (Figs. 3b, 5, PA) is a large and subrectangular bone in lateral view. In the skull roof, judged by its placement and position of supraoccipital it meets its partner in the midline as commonly found in well-preserved †ellimmichthyiforms. As for frontal bone, there are rugose ornamentation near lateral border.

The commissural supratemporal sensory canal (Figs. 3b, 5, stt.com) is entirely bone-enclosed, running within a curved tubular relief on parietal. The supraoccipital is placed outside this commissure. Pores are not visible on surface.

Like many clupeomorphs, the otico-occipital region is almost equal in size to orbital and ethmoid regions. Unfortunately, autosphenotic, prootic and pterotic bones are not preserved. There are any evidence of recessus lateralis and temporal fossa. Noteworthy the temporal fossa seems to be lacking in all †ellimmichthyiforms, as for the recessus lateralis, so that dermosphenotic are not reduced and opennings for preopercular and infraorbital branches of sensory canal are distantly positioned in otic neurocranium of certain well-preserved specimens (see Patterson 1970, Grande 1982, Forey 2004).

The epioccipital (Fig. 3b, EPO) is almost trapezoid and placed among parietal, supraoccipital, and an uninformative portion of pterotic. Its position in the skull resembles that found in †Triplomystus noorae (see Forey et al. 2003, p. 271, fig. 41). Posteriorly this bone shows a reduced smooth process to receive the upper arm of the posttemporal bone.

Behind parietals, there is a smooth supraoccipital (Fig. 3b, SOC) bearing a well-developed median crest resembling that of †Diplomystus. This bone does not separate the parietals as usual in clupeiforms. Due to the state of preservation it is not possible to determine presence of pre-epioccipital fenestra and posttemporal fossa. But a so-called parietal excavation as described by Forey (2004) in a three-dimensional skull of †Diplomystus from the Upper Cretaceous of the English Chalk is very probable (Fig. 5). Such excavation is considered by Forey (2004, Fig. 13, Node E) derived featured shared by †Triplomystus, †Sorbinichthys, and †Diplomystus. The same author stated it is absent in †Ellimmichthys longicostatus, †Ellimma branneri and †Paraclupea. Otherwise we also noted its presence in †Scutatuspinosus and †Ellimma cruzae.

The orbit of †C. carnavalii is large. Its diameter is contained of about 3 1⁄2 in HL. The eyeball were supported by two large shields of sclerotic bones from which only one (anterior) is preserved (Figs. 3 and 4, scl. b) in the holotype.

CIRCUMORBITAL BONES

Only anterior elements from this series remain. We interpreted a triangular bone displaced, lying on the anterior border of the lachrymal (=first infraorbital), as an anamestic antorbital (Figs. 3b, 4, ANT). It is very similar in shape to that of †Leptolepides haerteisi, an early euteleostean fish known form the Tithonian of Germany (see Arratia 1997, p. 71, Fig. 47). Besides the absence of antorbital has been claimed as a derived feature for †ellimmichthyiform fishes (Alvarado-Ortega et al. 2008) this bone is seen in †Horseshoeichthys (see Newbrey et al. 2010, fig. 2, identified as the first infraorbital bone, a mistake) in the same position and association of this bone as in Denticeps clupeoides (see Di Dario and de Pinna 2006).

Behind antorbital three infraorbital bones are preserved in a row. The lachrymal (=first infraorbital bone, Figs. 3b, 4, LA) is an elongate and rectangular bone lying on beneath the anterior half of the orbit. It shows an anterior border slightly rounded. It covers only the anteriormost edge of the suspensorium, extending from the facet of articulation to maxilla of autopalatine to the level of the posterior border of ethmoid lateral. The infraorbital sensory canal is consistently present running within the bone close to the dorsal border in a slender bony tube.

The second infraorbital bone (Figs. 3b, 4, IO2) is short and low, almost rectangular. It is placed below the middle point of the orbit. The infraorbital sensory canal pierces the bone at the midline taking into account the presence of a conspicuous tubular relief.

The third infraorbital bone (Fig. 3b, IO3) is the largest of the set. It is subrectangular as in †Diplomystus dentatus (see Grande 1982, p. 10, Fig. 7) and show an anterior flange below the contact zone with second infraorbital. This bone forms most of the ventral rim of orbit extending from the level of the quadrate-mandibular joint to the posterior extremity of the symplectic, but not entirely recovering the cheek. The infraorbital sensory canal is bone-enclosed near the orbital rim. Its membranodermal component is very expanded.

The fourth and fifth infraorbital bones, although not preserved, judging by a great gap above third infraorbital, mighty be tubular flimsy bones as observed in †Diplomystus dentatus (see Grande 1982).

Jaws and suspensorium. - The premaxilla is partially preserved and displaced from the anteriormost region of skull (Fig. 3b, PMX). It is low, triangular and toothless. The maxilla (Figs. 3b, 4, MX) exhibits an elongate anterior process finishing in a dilated articular head. There is a short and blunt autopalatine condyle upon this bone. The main body of the bone produces a slightly convex dentigerous lamina (as in †Diplomystus dentatus) on oral edge and extends backwardly to end at the level of the middle point of orbit. True teeth are lacking.

There are two smooth supramaxillae incompletely preserved. The first one, the anterior supramaxilla (Figs. 3b, 4, ASMX), is an elongate and elliptical bone lying on the anterodorsal border of the maxilla. The posterior supramaxilla (Fig. 3b, PSMX) shows a large and ovoid main body, very fragmented in DGM 435-P, but its anterodorsal process is lost.

The lower jaw is relatively short, deep and well-ossified. Laterally it consists of dentary, anguloarticular and retroarticular (Fig. 3).

Most of the lower jaw is formed by the dentary (Fig. 3b, D) which contributes with at least 80% of length. This bone has a deep symphysis and the oral border ascends abruptly producing a high coronoid process. There is a short row of at least 20 minute conical teeth on oral border. The path of the mandibular sensory canal (Fig. 3b, md.c) is evident on surface through a slight and straight tubular relief. Pores are not visible.

The anguloarticular (Fig. 3b, AA) is a relatively short and deep bone. It contributes to form part of the coronoid process. Its articular facet for quadrate is well-developed. Posteriorly it shows a short and slightly rounded postarticular process. Its median portion is crossed by a extension of the mandibular sensory canal.

As for dentary, the anguloarticular is only a little incurved suggesting the presence of a low meckelian fossa and consequently few volume of adductor muscle in life. The posterior opening for the mandibular sensory canal is not observed laterally so that we interpreted it was placed medially. A small retroarticular (Fig. 3b, RAR) is visible at the posteroventral corner of the lower jaw upon anguloarticular.

The quadrate-mandibular articulation is placed below the middle point of the orbit. The triangular quadrate (Fig. 3b, Q) is well-ossified and slightly curved forwards. Its articular condyle for lower jaw is well-developed and the posteroventral process is short, sharp, and vertically oriented. The anterior border of preopercle is tightly attached to the posterior margin of posteroventral process. The dorsal margin of quadrate seems to be truncate.

A metapterygoid is not preserved but judged by a great gap on suspensorium this was of moderate size. The symplectic (Fig. 3b, S) is a short, narrow, and club-like bone inserted in a notch of posterodorsal margin of quadrate, between the main body of this bone and its posteroventral process.

The shape of the hyomandibula (Fig. 3b, HM) is inferred mainly from the imprint of this bone on the rock. It is vertically oriented in respect to braincase and shows an obliquely positioned articular head for otic neurocranium. The vertical process is elongate and the opercular process is very short and stout.

Anterior to quadrate there are a badly preserved ectopterygoid and an elliptical and a shield-like endopterygoid (Figs. 3b, 4, ENPT) closely associated. The latter is visible in the inner orbit and above the second and third infraorbitals. Both bones are toothless.

From the autopalatine remains only a prominent and robust facet for maxilla (Fig. 3b, APAL) placed anterior to lachrymal bone.

OPERCULAR SERIES

The usual complement of teleostean opercular bones is present in †Codoichthys. The preopercle (Fig. 3b, POP) is a L-shaped bone, with dorsal limb longer than the ventral one. The ventral limb has ventral margin slightly convex. The preopercular sensory canal (Fig. 3b, pop. c) runs into a bony tube in the midline of dorsal limb of the bone. We are unable to determine the presence of tubules in dorsal limb, but in the ventral limb the main canal gives off two conspicuous recumbent tubules (better seen in DGM 436-P) as in †Ellimma branneri (see Chang and Maisey 2003, p. 10, Fig. 5).

The interopercle (Fig. 3b, IOP) is a triangular and elongate bone underlying the entire length of the ventral limb of the preopercle. Its ventral edge is a slightly convex.

The opercle (Fig. 3b, OP) is a well-developed bone, deeper than long, with dorsal margin rounded and an oblique contact zone for subopercle ending in a protruding anteroventral corner. This bone is contained of about three times in the head length. The ventral half shows on surface fading ornamentation under form of parallel striations (better seen in DGM 436-P), a pattern shared with †Ellimma and †Tycheroichthys.

The subopercle (Fig. 3b, SOP) is a large and falcate bone. It shows digit-like anterior ascending process and smooth posteroventral margin. Its major depth is contained of about three times in the opercle depth.

PAIRED FINS AND GIRDLES

The pectoral girdle includes the posttemporal, supracleithrum, cleithrum, postcleithra, scapula, and coracoid. The posttemporal (Fig. 3b, PTM) exhibits a rounded and smooth-bordered main body and an elongate and sharp anterodorsal limb for contacting epioccipital. The ventral limb for intercalar is club-like, very slender and long. An oblique tubular relief for lateral line is seen along the main body of the bone. The supracleithrum (Fig. 3b, SCL) is an elliptical bone lying on the dorsal process of cleithrum. A short tubular relief for lateral line (Fig. 3b, lat.l) is visible on surface.

The cleithrum (Fig. 3b, CL) is a well-ossified L-shaped bone with expanded laminar posterior margin. Therefore it is not sigmoid as in most of †ellimmichthyiform fishes (see Forey 2004). The overall shape is similar to that of the clupeiform †Santanaclupea silvasantosi from the Araripe Basin, Northeastern Brazil (see Maisey 1993). The anteroventral process is elongate and projected forwards along all extension of the margin of subopercle. Regarding this aspect it is very similar to that of †Diplomystus dentatus.

There are two distinct postcleithra crossing over pectoral-fin rays in DGM 436-P. Grande (1985) indicated absence of postcleithra in †Ellimmichthyiformes. But since then postcleithra were registered in †Horseshoeichthys (Newbrey et al. 2010) and †Triplomystus applegatei (Alvarado-Ortega and Ovales-Damian 2008).

Unfortunately endosteal bones of pectoral girdle are badly preserved and uninformative. An imprint of a small and quadrangular scapula is seen in DGM 435-P. The coracoid (Fig. 3b, CO) is large, laminate and L-shaped. We counted at least 15 pectoral fin-rays and the uppermost fin-ray is the thickest of the set.

The pelvic bone is apparently hidden by abdominal scutes so that nothing is visible externally. The pelvic fin is in opposition to dorsal fin and remains in a middle point between insertions of pectoral and anal fins. There are five or six branched pelvic fin-rays.

DORSAL AND ANAL FINS

The dorsal fin origins in a vertical line spanning abdominal vertebrae 10 and 11 (Fig. 2B). There are two rudimentary fin-rays followed of at least 10 fin-rays in association with at least 10 pipe-like proximal pterygiophores. The first pterygiophore shows an anterior lamina with a notch between dorsal and ventral branches, forming an acute angle. It supports the three anteriormost fin-rays. The number of anal-fin rays is inferred of 15 preserved pterygiophores. There are one rudimentary followed of at least 15 fin-rays.

VERTEBRAL COLUMN, PLEURAL RIBS AND INTERMUSCULAR BONES ASSOCIATED

There are, at least, 33 preural vertebrae, 11 in caudal region. All vertebral centra are hour-glass shaped in lateral view. Each vertebral centrum is smooth, except for weak longitudinal ridge separating grooves (Fig. 8B). The precaudal abdominal centra are mostly deeper than long. From the origin of dorsal fin backwards they became gradually longer than deep.

The neural arches (Fig. 6, n.a) are co-ossified to centra and the parapophyses (Fig. 6, pap). Neural spines (Fig. 6, n.sp) are long and touch proximal end of the dorsal pterygiophores. We are unable to determine the presence or not of bifid neural spines in the abdominal region due to the state of preservation. But in the caudal region, only a single neural spine fused to arch is seen.

Hemal spines ( Fig.6, h.sp) are as fine and long as the neural spines. There are any consistent differences in size or thickness among spines contributing to sustain the caudal fin.

The supraneural (=predorsal) bones form a series of eight slender, long, and almost sigmoid elements closely associated to predorsal scutes (Fig. 7). The space among supraneurals are unequal. The first supraneural is very short and positioned far from the other ones. The posteriormost one is the longest and closely associated to anterior lamina of the first pterygiophores of the dorsal fin. Apparently all supraneurals reach the level of the tips of abdominal neural spines.

The epineurals (Fig. 6, EPN) are long, thin, and laterally arched. The anteriormost ones are proximally fused to the base of abdominal neural arches. They are symmetrically aligned along the vertebral column spanning an area corresponding to five to six vertebrae. In the transition between abdominal to caudal regions there are high on the flank detached epineurals forming epimeral hypsiloid intermuscular bones (Fig. 6, EPM), in a framework backwards spanning an area spanning eight vertebrae. They continue backwardly on caudal region in opposition to some detached epipleurals (Fig. 6, EPL).

There are at least 17 pleural ribs. They are long and slightly curved and reach the tip of abdominal scutes (Fig. 8). Each one shows a deep groove along most of its extension. The anteriormost pleural ribs are lodged in a groove of the abdominal vertebra whereas the remaining ones on short parapophyses.

PREDORSAL AND ABDOMINAL SCUTES

There are slight imprints of eight elliptical predorsal scutes bearing a median keel (Fig. 7). All scutes are smooth, subequal-sized and lack posterior spine. They touch the distal end of the supraneurals.

A row of pre-pelvic scutes composed of 10 elements originates closely to pectoral fin at the level of the last fin-ray (Fig. 2). Each scute is ventrally keeled and bears long and sharp triangular lateral processes. The abdominal postpelvic series shows nine scutes. They are triangular, ventrally keeled and show short posterior spine-like process unlike erected and pungent of the most of †ellimmichthyiform fishes. The series finishes at the origin of the anal fin.

CAUDAL ENDOSKELETON AND FIN

The caudal fin is forked and shows equal-sized lobes. It is supported by two preural and two ural centra. As usual in many basal teleosteans, there are 10 principal fin rays in the upper lobe and 9 in the lower lobe. The region of dorsal procurrent rays is very damaged but there are at least four preserved in DGM 435-P. There are at least seven ventral procurrent rays and a probable ventral caudal scute (Fig. 8b, c.sc). The vertebral axis bends gradually upwards as usually found in non-clupeiform clupeomorphs. The neural spine of the second preural centrum is slender and elongate. Most of endoskeletal structures are preserved (Figs. 8a, b).

The proximal end of parhypural (Fig. 8b, PH) is fused to the first preural centrum. The neural arch of first preural centrum is enlarged and leaf-like extending backwardly to cover the dorsal margin of the first ural centrum and to contact laterally the first uroneural. This condition differs from that of †Diplomystus dentatus in which only the neural arch of the first ural centrum is enlarged and backwardly expanded.

The first ural centrum (Fig. 8b, U1) is almost hour-glass shaped and equal-sized to preural centra. is hour-glass shaped and almost equal-sized to preural centra. However the second ural centrum (Fig. 8b, U2) is more reduced, cone-shaped with tubular posterior end.

We counted six hypurals (Fig. 8b). The first one (Fig. 8b, H1) is a large triangular plate, long and enlarged distally. Proximally it attaches the first ural centrum and contributes for the hypural foramen. The second hypural is slender and elongate, and fused to first ural centrum (Fig. 8b, U1+H2). The remaining four hypurals are associated to second ural centrum. The third hypural (Fig. 8b, H3) is triangular and large; its expanded spatulate portion invades ventrally space of lowermost hypurals. Probably it was attached to second ural centrum in life. There is a large diastema between third and second hypurals. This pattern differs from that of †Scutatuspinosus and †Rhombichthys whose third hypural has an upward process constraining the fourth hypural to project forwardly. This condition seems to be shared uniquely for these taxa. The other three hypurals are elongate and gradually decrease in size towards the dorsal margin of the upper lobe.

Our observations are in contrast with the original description. Santos (1994) pointed out the occurrence of four uroneurals. We consistently observed two uroneurals (Fig. 8b), all free from the centra. The existence of a third uppermost uroneural (Fig. 8b, ?UN3) is probable taking into account by a clear space in the region anterior to the tips of invading fin rays and the presence of a strut of bone in this place. The first uroneural (Fig. 8b, UN1) extends forwards for reaching the first preural centrum laterally. It is placed laterally to the neural arch from the first preural centrum. The second uroneural (Fig. 8b, UN2) is short and fusiform, and lies on the dorsal margin of the second ural centrum close to the origin of the second ural centrum.

There are three thin, elongate and curved epurals (Fig. 8b, EP, numbered) occupying a space between neural spine of the second preural centrum and dorsal border of neural arch from the first preural centrum.

RESULTS OF THE PHYLOGENETIC ANALYSIS

The hypothesis generated by the present phylogenetic analysis corresponds to a strict consensus of three equally parsimonious trees (Fig. 9). The majority consensus tree shows the same topology. The tree has 181 evolutionary steps, Consistency Index (CI)=0,387, Retention Index (RI)=0,632.

Figure 9
The strict consensus of the three shortest trees obtained. Uniquely derived characters are indicated with an asterisk symbol (*). Synapomorphies supporting nodes are: Node 1 (†Ellimmichthyiformes): 21[0►1],39*[0►1]; Node 2 (†Sorbinichthyidae): 18[0►1], 22[0►1], 24[1►0], 42[0►1]; Node 3 (unnamed): 4[0►1], 11[2►1], 52[0►1]; Node 4 (unnamed): 18[0►1], 60[0►1]; Node 5 (†Diplomystus clade): 40[0►1], 41[►1], 42[►1]; Node 6 (†Paraclupeidae): 11[1►2], 46[2►0]; Node 7(unnamed): 1[0►1], 44[0►1]; Node 8 (†Thorectichthyinae): 24[1►0], 30[1►], 32[0►1], 38[0►1], 48[1►2]; Node 9 (†Paraclupeinae): 37[1►0], 43[0►1], 45[0►1]; Node 10 (†Triplomystini): 29[0►1], 50[0►1]; Node 11 (unnamed): 36[1►0], 49[01►2]; Node 12 (unnamed): 26[0►1], 48[1►2], 51*[0►1]; Node 13 (†Paraclupeini): 12[1►0], 53[0►1]; Node 14 (unnamed): 30[1►0]; Node 15 (unnamed): 5[0►1], 19[0►1]; Node 16 (unnamed): 16[0►1], 20[0►1], 52 [1►2]; Node 17 (unnamed): 22[1►0], 32[0►1], 40[0►1].

It is not possible to confirm if †Ornategulum belongs with clupeomorphs. In this analysis this taxon is excluded of Clupeiformes and †Ellimmichthyiformes and placed outside as the sister group of all clupeomorphs used in this analysis.

†Sorbinichthyidae appears in a basal polytomy together with the enigmatic †Horseshoeichthys and all other †ellimmichthyiforms. But we think that the position of †Horseshoeichthys is doubtful because it shows many missing data in the matrix and herein confirmed as a "wild-card" (sensuMurray and Wilson 2013). Thus its position among †Ellimmichthyiformes still depends on the collection of well-preserved specimens. If so, excluding †Horseshoeichthys, †Sorbinichthyidae becames the putative sister group of all other †ellimmichthyiforms.

We have found no diagnostic character for †Armigatus so that species of this genus appear in a basal polytomy below more advanced taxa.

Species of †Diplomystus (excluding † "D". solignaci) form a group supported by the second higher value of Bremer index (3). We obtained a different placement for this group so that it is separated of †Armigatus and †Sorbinichthyidae, dismantling †Armigatoidei (†Diplomystus in part plus †Armigatus sensuMurray and Wilson 2013). †Diplomystus is placed in a more advanced position in the tree in comparison with †Sorbinichthys and †Armigatus. †"Diplomystus" solignaci is positioned within an advanced †Paraclupeidae but, in contrast with previous analyses, it appears as sister group of †Paraclupea chetunguensis.

Node 1 (=†Ellimmichthyiformes) is diagnosed by the uniquely derived presence of predorsal scute series (C=39) and S-shaped cleithrum (C= 21), this latter reversed in †Codoichthys and †Horseshoeichthys.

Node 2, linking species of †Sorbinichthyidae, are characterized by homoplastic features of pleural ribs insertion (C=18), subrectangular dorsal arm of posttemporal (C=22), a reversed condition to autogenous hypural (C=24), and spines on predorsal scutes (C=42). The latter is also shared with †Diplomystus.

Node 3, linking species of †Armigatus (in polytomy) and all †ellimmichthyiforms above †Sorbinichthyidae, is characterized by one striking ornamentation of skull roof (C=4), initial presence of teeth in a patch on parasphenoid (C=11), and certain number (22 to 30) of abdominal scutes (C=52).

A Node 4, linking †Diplomystus clade and †Paraclupeidae, is diagnosed by a special relation of pleural ribs and grooves of vertebral centra on abdominal region (C=18) and a conspicuous parietal excavation (C=60).

The Node 5 (Bremer index =3) corresponds to the †Diplomystus clade and is diagnosed by the presence of sub-rectangular scutes in the anterior (C=40) and posterior (C=41) portion of predorsal series, spines on posterior edge of predorsal scutes (C=42, homoplastic feature shared with †Sorbinichthys).

Node 6, †Paraclupeidae new usage is diagnosed by another transformation of the character 11 (state 1 to 2), resulting in loss of teeth on parasphenoid (homoplasy shared with clupeiforms), and a derived number of predorsal scutes (C=46, state 2 to 0). It contains †Codoichthys and all remaining †ellimmichthyiforms.

A Node 7 (unnamed) includes †ellimmichthyiforms more advanced than †Codoichthys. It is divided in two subfamilies †Thorectichthyinae (Node 8, new name) and †Paraclupeinae (Node 9, new use). They share marked angle in front of the insertion of dorsal fin (C=1, secondarily lost is †Codoichthys and †Scutatuspinosus), and the size of scutes increasing backwards (C=44).

A Node 8 (†Thorectichthyinae), linking species of †Thorectichthys is recognized by five apomorphies (C=24, 30, 32, 38, and 48, see Fig. 10) in addition to that proposed by Murray and Wilson (2013). This node shows the highest value of bootstrap (i.e., 75) in our analysis.

For Node 9, †Paraclupeinae, includes two sister tribes: †Triplomystini and †Paraclupeini. The latter includes tribes †Triplomystini (Node 10, new usage) and †Paraclupeini (Node 13, new usage). This node is characterized by reduction of neural spine of first preural centrum (C=37, state 1 to 0 ) and derived presence of ornamentation (C=43) and strong spine (C=45) in predorsal scutes.

†Triplomystini (Node 10) contains all species of †Triplomystus clade (Node 12, diagnosed by the homoplastic features sharp end of first hypural and quantity of predorsal scutes, and an uniquely derived C=51, i.e., postdorsal scutes) that is sister group (Node 11) of †Tycheroichthys plus †Rhombichthys.

This arrangement of Node 11 is new and shows the highest value of decay (i.e., Bremer index equals 4). Node 10 is characterized by two homoplastic features: derived number of uroneurals (C=29) and spatulate abdominal scutes (C=50).

†Paraclupeini (Node 13) is linking †Scutatuspinosus and advanced taxa by sharing homoplastic condition of characters 12 (absence of supraorbital, state 1 to 0) and 53 (loss of postcleithra). We think these characters used to diagnosed both groups demands more study. Noteworthy †Scutatuspinosus shows reversions (characters 1 and 50), and a independent acquisition (character 56).

Within †Paraclupeini, a Node 14 is linking †Ezkutuberei and more advanced taxa (Node 15, †Ellimma, †Ellimmichthys, †Paraclupea and † "Diplomystus" solignaci). Only a reversion of character 30 (state 1 to 0) supported it. Better is the case of Node 15 characterized sculptured skull roof (C=5, state 0 to 1) and extension of epineurals and epipleurals on caudal region (C=19, state 0 to 1). Assuming the condition found in †Rhombichthys and †Triplomystus applegatei not ambiguous (as interpreted by the TNT program), it is a putative uniquely derived feature for the clade.

Alignement between †Ellimmichthys and †Paraclupea was yet detected in early efforts to decipher relationships within †Ellimmichthyiformes (e.g., Chang and Grande 1997, Chang and Maisey 2003, Forey 2004). It is confirmed in Node16 although species of †Ellimmichthys appear in a polytomy reflecting still bad knowledge of their structures. The node also includes the enigmatic †"Diplomystus" solignaci and is supported by three homoplastic features (characters 16, 20, and 52). In this we noted a well-supported clade (Node 17) formed by †Paraclupea and †"Diplomystus"solignaci. This node is the second highest value of bootstrap (i.e., 50) in our analysis although supported by homoplasies (characters 22, 32, and 40). Thus †"Diplomystus"solignaci is confirmed outside that genus.

We made an experimental analysis excluding the so-called 'wild-cards' or "swing taxa" (i.e., †Ornategulum, †Ellimmichthys longicostatus, †Ezkutuberezi carmenae, and †Horseshoeichthys, together with †Ellimmichthys goodi) and using command traditional search, we obtained seven trees (168 steps, CI 0,417, RI 0,611). Besides higher CI and lower steps, applying strict consensus we obtained a paraphyletic group for species of †Sorbinichthys and a great basal polytomy with remaining †Ellimmichthyiformes. In sum, in this step we have found a paradoxical result with Murray and Wilson (2013).

DISCUSSION AND CONCLUSIONS

After Grande (1985), the classification of Ellimmichthyiformes, in a historical perspective, become a continuous effort to add or exclude taxa. It was obviously associated with many attempts of tie the name of the taxon to at least one character that is necessary for membership. In this context Grande's approach was based on drawing-by-hand analysis, with a monophyly accepted a priori with selected characters and omitting homoplasies.

Since then †Ellimmichthyiformes seems to be generally accepted, besides strikingly short of characters. Particularly, we think that the reality of the group is better supported mainly because any member demonstrates close affinities with taxa outside (e.g., clupeiforms, ostariophysans, and elopomorphs).

Chang and Maisey (2003) were the first to use parsimony analysis performed in a computer program (i.e., PAUP version 4.0) to test monophyly of †Ellimmichthyiformes. Their results were obtained on the basis of a data matrix including 30 characters and 11 taxa. The characters used were all taken of Grande (1982, 1985) but they included new taxa too. As a result, they found a clade containing †Armigatus, †Diplomystus, †Ellimma, †Ellimmichthys, and †Paraclupea. This group was subdivided is two subclades, one for †Armigatus plus †Diplomystus and other containing †Ellimma, †Ellimmichthys and †Paraclupea. Noteworthy, these authors were the first to pointed out †Scutatuspinosus as showing some features indicating close affinities with †paraclupeines.

In an approach to known if Ellimmichthyiformes is a clade or a convenience group, Zaragueta-Bagils (2004) used a data matrix including 56 characters and 15 terminal taxa. He indicated that †Armigatus brevissimus, †Diplomystus birdi, and †D. dentatus form a group outside clupeiforms and †ellimmichthyiforms. His results dismantled †Ellimmichthyiformes.

Forey (2004) furnished a drawing-by-hand phylogenetic scheme for †Ellimmichthyiformes diagnosing the clade by a S-shaped cleithrum. Even he recognized two great subclades: a node D (for †Armigatus, †Triplomystus, †Diplomystus) and node C (for †Paraclupea and †Ellimmichthys). He stated Node D diagnosed by enlarged endopterygoid teeth, elongate anal fin, and deep and narrow opercle. His node C was characterized by characters taken of Grande (1982, 1985). Unfortunately, he omitted informative taxa as †Ellimma, †Scutatuspinosus, †Codoichthys, all yet known in that time.

Alvarado-Ortega et al. (2008) using a data matrix of 28 clupemorphs and 58 characters excluded †Armigatus from the †Ellimmichthyiformes. They used mainly characters taken of Grande (1982, 1985) and Zaragueta-Bagils (2004) and divided the group in two clades: †Sorbinichthyidae and †Paraclupeidae. According to them, †Sorbinichthys is a member of a †Diplomystus clade and †Paraclupeidae containing all other †ellimmichthyiforms. But certain taxa were omitted in the analysis of Alvarado-Ortega et al. (2008) as the case of †Codoichthys but other (e.g., †Tycheroichthys, †Rhombichthys, †Torectichthys) were only described in recent years. Other problem detected was about definition of characters and codification of states so that they were critically reassessed by Murray and Wilson (2013).

Murray and Wilson (2013) used 28 taxa and 62 characters in the most recent and complete cladistics analyses. Their results were condensed in a consensus tree of three shorest trees (176 steps, CI=0,44, RI=0,57) . It follows that†Armigatoidei (†Armigatus plus †Diplomystus) was found as sister of all other †ellimmichthyiforms. The strict and majority-role consensus trees were as our results, identical. As in our analysis neither decay (Bremer support) nor bootstrap analyses indicated strong support for any branch except for certain pair of species. †Ellimmichthyoidei contained †Sorbinichthyidae and †Paraclupeidae (including †scutatuspinosines, †thorectichthyines, and †paraclupeines). During their analyses some taxa were excluded because introduce many problems (i.e., many missing data), for instance, †Ornategulum, †Ellimmichthys longicostatus, †Ezkutuberezi, and †Horseshoeichthys. Even, they indicated † Ellimmichthys could be paraphyletic.

Recently some preliminary approaches to decipher relationships of †ellimmichthyiforms were carried out by F.J. de Figueiredo (unpublished data, Ph.D dissertation, Museu Nacional-UFRJ, Rio de Janeiro, 2006), F.J. de Figueiredo and D. Barros (unpublished data, abstracts of the 14th Congresso Latino Americano de Paleontologia de Vertebrados, p. 172, San Juan, Argentina, 2011), D.R.M. Ribeiro and F.J. de Figueiredo (unpublished data, abstracts of the 23th Congresso Brasileiro de Paleontologia, p. 270, Gramado, Rio Grande do Sul, 2013) and A. Nunes and F.J. de Figueiredo (unpublished data, abstracts of the 23th Congresso Brasileiro de Paleontologia, p.255, Gramado, Rio Grande do Sul, 2013).

F.J. de Figueiredo (unpublished data, Ph.D dissertation, Museu Nacional-UFRJ, Rio de Janeiro, 2006) furnished a review of articulated and complete clupeomorphs found in Lower Cretaceous and Tertiary deposits outcropping in many localities from the Northeastern Brazil. At first a data matrix containing 61 characters and 19 taxa were used in that analysis. The anatomy of many taxa were revisited in detail and a phylogenetic scheme was obtained. As a result †Ellimmichthyiformes appears as a weakly supported clade. Only a S-shaped cleithrum was indicated as uniquely derived feature, confirming hypothesis by Forey (2004). In addition, †Paraclupeidae only contained †Ellimmichthys longicostatus, †Ellimma cruzae, †Triplomytus noorae, †Paraclupea and a unnamed †Ellimmichthys-like taxon from the Lower Cretaceous of Reconcavo Basin, State of Bahia. †Codoichthys, †Scutatuspinosus, †Ezkutuberezi, and †Ellimma branneri appeared in a basal polytomy, and †Diplomystus (i.e., †D. shengliensis and †D. dentatus) plus †Armigatus formed a clade. Like Alvarado-Ortega et al. (2008) this analysis become dated because did not include many taxa only described recently.

F.J. de Figueiredo and D. Barros (unpublished data, abstracts of the 14th Congresso Latino Americano de Paleontologia de Vertebrados, p. 172, San Juan, Argentina, 2011) while revisiting the anatomy of †Scutatuspinosus preliminarly proposed a new hypothesis on phylogeny. They obtained eight trees with 128steps based on a data matrix including 15 putative ellimmichthyiforms, two basal clupeiforms (i.e., †Santanaclupea silvasantosi and †Pseudoellimma gallae, see Maisey 1993, Figueiredo 2009). And as for previous analyses, low index (CI=0.5; RI=0.6) indicating many missing data as the main problem. †Ellimmichthyiformes and †Paraclupeidae were confirmed. In addition Figueiredo and Barros suggested the exclusion of †Ellimma cruzae from the genus †Ellimma and put formally †Codoichthys within †Ellimmichthyiformes. But the inclusion of †Santanaclupea silvasantosi and †Pseudoellimma gallae as near outgroup in that analysis introduced many missing data and a doubtful result.

A. Nunes and F.J. de Figueiredo (unpublished data, abstracts of the 23th Congresso Brasileiro de Paleontologia, p.255, Gramado, Rio Grande do Sul, 2013) while describing new data about † Ellimmichthys-like taxon from the Lower Cretaceous of Reconcavo Basin, State of Bahia, once more worked on phylogeny. They used 22 taxa and 39 characters and obtained 102 trees, with 132 steps, CI=0.6; RI=0.79. It follows † Ellimmichthyiformes was diagnosed by a S-shaped cleithrum (one more time agreeing with Forey 2004) and †Paraclupeidae forming a clade including †Rhombichthys, †Tycheroichthys, new taxon and †Paraclupea. But this analysis did not included recently described taxa and many characters of Murray and Wilson (2013).

D.R.M. Ribeiro and F.J. de Figueiredo (unpublished data, abstracts of the 23th Congresso Brasileiro de Paleontologia, p. 270, Gramado, Rio Grande do Sul, 2013) using unchanged data from literature (i.e., Santos 1994, Santos and Correa 1985, Chang and Maisey 2003, Alvarado-Ortega et al. 2008) and taxa recently described (i.e., †Tycheroichthys dunveganensis, †Rhombichthys intoccabilis) performed a preliminary cladistics analysis of † Ellimmichthyiformes. Their analysis was based on a data matrix containing 26 taxa and 57 characters. They obtained 10 shortest trees (L=177 passos, CI=0.463 e RI=0.652).

As a result, †Codoichthys appeared in basal position in a tree as sister group of remaining †Ellimichthyiformes. Once more †Diplomystus is indicated as paraphyletic taxon. †Rhombichthys revealed to be sister group of †Ezkutuberezi and appeared with†Tycheroichthys in a great polytomy together with †Ellimma branneri, †Diplomystus solignaci, †Ellimmichthys, and †Paraclupea plus †Triplomystus. They confirmed †Paraclupeidae with †Scutatuspinosus as sister group of the deep-bodied †Ellimmichthyiformes.

Noteworthy the unpublished results of Ribeiro and Figueiredo and Murray and Wilson (2013) appeared independently in the same year. Both works have not included or discussed data and results from each other.

Unlike results of D.R.M. Ribeiro and F.J. de Figueiredo (unpublished data, abstracts of the 23th Congresso Brasileiro de Paleontologia, p. 270, Gramado, Rio Grande do Sul, 2013) we took into account the corrections furnished by Murray and Wilson (2013), using new insights on characters and states (see ^{Appendix 1}, commented list of characters and states), and including new taxa. Thus we furnished an ultimate character list. Although highlighting character conflict we feel again the urgency of revisiting characters and reassessing taxa. Clearly the situation with regard to relationships of taxa within ellimmichthyiforms is unresolved and most of lineages are supported only by homoplasies. Many problems remain. For instance, although †Horseshoeichthys has an incertae sedis position at the base of the group in polytomy with †Sorbinichthyidae and all other †ellimmichtyiforms, we think that more informative specimens are need to decipher its relationships. Thus the most basal group of †Ellimmichtyiformes is probably †Sorbinichthyidae.

In sum, unlike Zaragueta-Bagils (2004) but agreeing with Alvarado-Ortega at al. (2008) and Murray and Wilson (2013) we confirm the existence of a †Ellimmichthyiformes group, though weakly supported. There are major differences between the topology of trees of Chang and Maisey (2003), Zaragueta-Bagils (2004), Forey (2004), Alvarado-Ortega et al. (2008), and Murray and Wilson (2013) and the hypothesis herein proposed. It confirms the impact of new described taxa in the phylogenetic analysis of a problematic group whose definition, diagnose, and composition wait for better results.

Particularly we noted that the main divergence among most of the analyses is about the position of †Sorbinichthys, †Armigatus and †Diplomystus in relation to the remaining taxa, difficulting linking. They are "swing" groups, sliding from the base of tree to near †Paraclupeidae. This latter includes mostly deep-bodied taxa, but its composition until now remains unresolved.

†Codoichthys is a basal and primitive †Paraclupeidae. In our analysis this taxon appears as sister group of remaining †paraclupeids.

As far as we known most recent phylogenetic analyses were prejudiced by many homoplasies and missing data. All phylogenetic schemes after Chang and Maisey (2003) produced trees with weakly supported branches.Thus, as claimed by Murray and Wilson (2013) we need more complete and informative specimens as well as revisiting old material in a search for new characters.

ACKNOWLEDGMENTS

We are indebted to Drs. Valéria Gallo and Hugo Ricardo S. Santos for reading a draft of this paper and for offering many helpful criticism and suggestions. We also thank Dr. Lionel Cavin and an anonymous reviewer for constructive critique and valuable comments. We are greatful to Maria Regina Salgado de Melo for producing high resolution photographs. Acknowledgement is also made to Fundação de Amparo à Pesquisa do Estado do Rio de Janeiro (FAPERJ) for a grant-in-aid that enabled the junior author to work with fish fossil material in 2012. We thank the staff of the Departamento Nacional de Produção Mineral for hospitality during visits and loan of specimens.

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APPENDIX 1

List of Characters and States Used in the Phylogenetic Analysis

The present cladistic analysis of †Ellimmichthyiformes is based on the characters below. [0] at first represents the assumed plesiomorphic state and [1], and [2], and [3] the apomorphic states. They are present according to anatomical region, in logical flow, from the head to tail, except for four last additional characters from the list. The outgroup used to polarize characters and rooting tree is Elops saurus. Additional taxa used in this analysis are the clupeomorphs †Ornategulum sardinioides, Chirocentrus dorab, Denticeps clupeoides, and Odaxothrissa vittata. Further explanation of the characters and their coding see publications of Chang and Maisey (2003), Zaragueta-Bagils (2004), Alvarado-Ortega et al. (2008), and Murray and Wilson (2013). These publications followed of character number are referred by abbreviations CM#, Z#, AO# and MW# respectively. Additional relevant authors are included in parenthesis. Comments in brackets detail changes we made or add explanation.

(1) Dorsal margin of the body with a marked angle at the insertion of the dorsal fin (modified from AO1, Z56, Chang and Grande 1997, MW1): [0] absent; [1] present [Due to enlargement and grade of ossification of predorsal scutes there is among so-called paraclupeids an almost straight ascending profile in advance of the origin of the dorsal fin. Noteworthy Khalloufi et al. (2010) pointed out this character may be affected by the growth and size of an individual.]

(2) Parietal bones (AO2, CM4, Z1): [0] meeting in the midline; [1] completely separated by supraoccipital. [Early lineages of teleosts have parietals meeting in the midline, as in Elops. It is very difficult to check the states among different taxa due to lateral compressing of most available specimens. But we agree with Chang and Maisey (2003) contra Chang and Grande (1997) about coding primitive state of this character for †Ellimmichthys longicostatus and †E. goodi.]

(3) Supraoccipital crest (AO3, Z2): [0] low and small; [1] high. [A deep crest, with large laminar structure abruptly projected upward, is verified in †Diplomystus shengliensis, †Ezkutuberezi carmenae, and †Codoichthys carnavalii unlike most of †ellimmichthyiforms. It is found in †Diplomystus dentatus, †D. dubertreti and †Rhombichthys. Forey (2004) described it in the well-preserved skull of an unnamed †Diplomystus from the English Chalk. Due to poor definition it has been differently and subjectively interpreted by different researchers. Therefore this character was modified after the matrix by Murray and Wilson (2013) as lateral profile of skull roof, enhancing a distinct angle between anterior and posterior regions.]

(4) Ornamentation of skull roof (Z3, AO4, MW4): [0] Absent; [1] Present. [Murray and Wilson (2013) simplified and joined characters 4 and 52 of Alvarado-Ortega et al. (2008) in a single character, and we used it here.]

(5) Dermal roof sculptured with coarse irregular and radiating ridges or grooves: [0] absent; [1] present. [The grade and topographic site of ornamentation on skull roof varies among taxa and according to age of individuals. Most of non-clupeiform clupeomorphs shows ornamentation under form of radiating ridges and grooves. It is particularly observed in †Spratticeps (Patterson 1970) and †Ornategulum (Forey 1973a). But the skull roof becomes strongly sculptured in †Ellimmichthys, †Paraclupea, and †Ellimma. The state is unknown in †Diplomystus solignaci and †Rhombichthys whereas †Horseshoeichthys exhibits a skull roof smooth. Many advanced clupeiforms commonly show parallel striae on border of the posterior part of frontal. Denticles or spines are homoplastically observed on dermal bones of Denticeps clupeoides and †Sorbinichthys elusivo. Noteworthy this character may be affected by age of the individual inasmuch as young individuals of †Ellimma lack ornamentation on dermal skull bones (Chang and Maisey 2003). Since the variety of ornamentation in frontal and parietal, we simplified the character Z4, AO5, and MW5.]

(6) Posttemporal fossa (AO6, Z6): [0] absent; [1] present. [Forey et al. (2003) described the presence of a posttemporal fossa in †Armigatus namourensis and †Triplomystus noorae. Even, Forey (2004) demonstrated its occurrence in a well-preserved skull of an unnamed †Diplomystus from the English Chalk. Apparently it is present in all other species of †Diplomystus except in the enigmatic †D. solignaci so that we used a question mark in the matrix. Unfortunately a posttemporal fossa is unknown in Brazilian taxa (i.e., †Codoichthys, †Scutatuspinosus, †Ellimma, †Ellimmichthys) and other taxa studied manly due to preservation.]

(7) Cavity or fossa in the temporal region of the skull (MW7, AO7, CM3, Z7, and modified from Forey 2004): [0] absence of cavity or fenestra; [1] pre-epioccipital fossa (i.e., between parietal, epioccipital and pterotic bones); [2] pre-epioccipital fenestra (i.e., between the parietal, epioccipital and supraoccipital bones). [Forey (2004) indicated originally differences between pre-epioccipital fenestra and fossa. It is not clear the state in †Scutatuspinosus, †Codoichthys, †Ellimma and †Ellimmichthys so that we coded as question mark.]

(8) Recessus lateralis (Grande 1985; CM1, Z8, MW8): [0] absent; [1] present. [The presence of a infraorbital canal merging with preopercular canal in the otic region in association with a reduction of dermosphenotic bone as far as we known is only found in clupeiform fishes (Grande 1985).]

(9) Supramaxillae (AO9, CM8, Z10, MW9): [0] two; [1] one or none. [According to Zaragueta-Bagils (2004) †ellimmichthyiform fishes show primitively two supramaxillae; it is the primitive state for teleosts (Arratia 1997, 1999). A reduction in number is assumed as secondary lost.]

(10) Basipterygoid process of parasphenoid (CM 11, AO10, Z11): [0] absent; [1] present.

[The dermal basipterygoid process is present in many †ellimmichthyiform fishes (Chang and Maisey 2003, Zaragueta-Bagils 2004, Forey et al. 2003, Forey 2004, Alvarado-Ortega et al. 2008). But due to poor preservation it is not possible to confirm if †Scutatuspinosus, †Ezkutuberezi, †Ellimmichthys and †Codoichthys show that state, therefore this character is coded as missing data for them.]

(11) Teeth on parasphenoid (modified from AO11, Z12, Grande 1982, 1985; MW11): (0) occupying most of the major extension of the orbital region; (1) only with a posterior osteoglossid-like tooth patch; (2) absent [All clupeiform fishes show toothless parasphenoid so far; the same for †Scutatuspinosus, †Codoichthys, †Ellimma, and †Ellimmichthys. A tooth patch backwardly placed on parasphenoid is only found in species of †Armigatus and †Diplomystus.]

(12) Supraorbital (AO12, Z13, MW12): (0) present; (1) absent. [This anamestic bone is commonly found in Clupeiformes and most of Elopiformes (e.g., Elops and Albula). But its putative presence is very difficult to confirm in many fossils due to flimsy nature and preservation. The situation is worse when few and badly preserved specimens are available. It is certainly found in †Scutatuspinosus itapagipensis (cf. DGM 1255-P and DGM 1253-P), †Ellimma branneri, †Sorbinichthys africanus (Murray and Wilson 2012) but doubtfully in †Rhombichthys. It is coded as missing data (question mark) in †Codoichthys. Furthermore we considered the presence of a supraorbital bone in †Horseshoeichthys uncertain so that it is herein coded ?]

(13) Antorbital (AO13, Z16, MW13): (0) present; (1) absent.[Although assumed as regularly present in many basal lineages of teleostean fishes (Arratia 1999), the presence or not this bone is very difficult to confirm in most of fossil fishes due to the flimsy condition. It commonly becomes easily lost or damaged. It is boomerang-like in †Scutatuspinosus (cf. DGM 1255-P) and quadrangular (and with canal) in †Horseshoeichthys (identified erroneously as a lachrymal by Newbrey et al. 2010) in which is very similar to Denticeps clupeoides (see Di Dario 2004). Gayet (1989) claimed the presence of an anamestic lachrymal as a synapomorphy of Clupeomorpha but as far is known it is not supported by any congruence of characters in foregoing analyses and, at worst, this bone clearly bears canal in Denticeps (see Di Dario and de Pinna 2006) and †Horseshoeichthys (see Newbrey et al. 2010).]

(14) Beryciform foramen on anterior ceratohyal bone (AO14, Z14, CM12, Grande 1985): (0) absent; (1) present. [It occurs in many taxa from early lineages of teleosts, many early euteleosteans, and early acanthomporhs. Otherwise it is absent in osteoglossomorphs, elopomorphs and advanced acanthopterygians. It is found in Scutatuspinosus (cf. DGM 1262-P). We coded this character mostly according to Zaragueta-Bagils (2004) assuming loss of this foramen in clupeiforms.]

(15) Foramen on the posterior ceratohyal (Z15, AO15): (0) absent; (1) present. [Although herein included in analysis, it is very difficult to verify this feature even in well-preserved specimens. It is a result of the enlargement of the groove for the hyoidean afferent artery. Apparently it is found in the hyoid apparatus of the unnamed †Diplomystus of the Cenomanian of English Chalk (Forey 2004). But as far as we known, among living clupeomorphs only Denticeps clupeoides possesses it. We found ceratohyal is widely covered by mandible and opercular bones in available specimens used in analysis. This feature contributes to increase the quantity of missing data in matrix.]

(16) Teeth on endopterygoid (CM9, AO16): (0) absent or minute; (1) present and well-developed. [many times it is only inferred of punctuated surface of this bone in association of isolated teeth.]

(17) Halves of the neural arches of most abdominal vertebrae (AO18, Z17): (0) independent; (1) fused medially. [see Arratia (1997) for explanation and discussion.]

(18) Pleural ribs (Z18, AO19): [0] all articulated with parapophyses along abdominal region; [1] the anteriormost ribs articulate with deep pits on the lateral side of the abdominal centra and those located posteriorly articulate with well-developed parapophyses; [2] all articulated with deep pits on the lateral side of abdominal centra. [Zaragueta-Bagils (2004) indicated states 0 and 1 as relevant and Alvarado-Ortega et al. (2008) modified it, including an additional state 2. We follow basically Murray and Wilson (2013) so that the state 1 occurs in †Ellimmichthys (Grande 1982), †Paraclupea (Chang and Grande 1997), †Triplomystus (Forey et al. 2003), †Ellimma branneri (Chang and Maisey 2003), and †Diplomystus solignaci (Gaudant and Gaudant 1971). †Codoichthys shares this last derived state.]

(19) Epineurals and epipleurals in the caudal region (Z19, AO20): (0) present, (1) absent. [According to Gaudant and Gaudant (1971) †"Diplomystus" solignaci does not show epineurals and epipleurals in the caudal region. We assumed epineurals and epipleurals occur in the caudal region of †Diplomystus birdi, †D. dentatus, †D. dubertreti, †D. shiengliensis, †Sorbinichthys elusivo, †Sorbinichthys africanus, †Triplomystus noorae, †Triplomystus oligoscutatus, Chirocentrus dorab, Elops saurus, Odaxothrissa, Denticeps clupeoides, and †Ornategulum sardinioides. Unlike Alvarado-Ortega et al. (2008), we clearly observed epineurals invading caudal region of †Scutatuspinosus (cf. DGM 1242-P).]

(20) Epicentrals (AO21, Z20, MW21): (0) absent; (1) present.

(21) Shape of cleithrum (modified of AO22, Forey 2004; MW22): (0) L-shaped; (1) sigmoid. [Forey (2004) claimed a sigmoid cleithrum is a synapomorphy of †Ellimmichthyiformes. Afterwards new non-clupeiform taxa were described bearing L-shaped cleithrum (e.g., †Codoichthys and †Horseshoeichthys).]

(22) Dorsal process of posttemporal (Z22, AO23): (0) slender and sharp; (1) sub-rectangular.

(23) Hypurals (AO26, CM20, Z23): (0) seven; (1) six; (2) five. [Alvarado-Ortega et al. (2008) counted six hypurals in †Scutatuspinosus probably assuming a common occurrence in †ellimmichthyiforms. We counted only five in our best preserved specimen examined (cf. DGM 1250-P). Therefore, we used question mark for this taxon.]

(24) Second Hypural (Z24, CM22, AO27): (0) autogenous; (1) fused to first ural centrum.

(25) First hypural and first ural centrum (Z25, AO28): (0) in contact; (1) without contact.

(26) Shape of proximal end of first hypural (Z26, AO29): (0) massive and forming an upward process; (1) sharp and straight.

(27) Shape of second hypural (AO30, Z27): (0) symmetrical to the fourth hypural; (1) very thin and stick-like. [Second and fourth hypurals are almost symmetrical in many clupeomorphs and most of †ellimmichthyiforms (Alvarado-Ortega et al. 2008). Apparently, only in Denticeps clupeoides it is stick-like.]

(28) Size of the first ural centrum (AO32, Z30): (0) roughly the same size as preural centra;

(1) smaller than the first preural centra.

(29) Number of uroneurals (Z29): (0) three; (1) two; (2) one. [Most of clupeomorphs (Grande 1985) and basal elopomorphs (Arratia 1997) exhibit three uroneurals. Unlike Alvarado-Ortega et al. (2008), we note the presence of three uroneurals in †Scutatuspinosus (cf. DGM 1250-P).]

(30) Proximal extremity of the first uroneural (Z31, AO34): (0) contacting second preural centrum; (1) not reaching or only contacting the first preural centrum. [In the basal and primitive teleostean fishes the first uroneural projects forwards anterior to second preural centrum (Patterson and Rosen 1977). In elopomorphs and basal clupeomorphs it reaches only the second preural centrum whereas in more advanced taxa, only first preural centrum is contacted. The coding of this character is based mainly on Zaragueta-Bagils (2004) but agreeing with Alvarado-Ortega et al. (2008) we considered the condition of †Diplomystus birdi and †D. dentatus, whose the first uroneural does not reach the second preural centrum. †D. shengliensis shows a first uroneural projecting forwardly to reach the second preural centrum (see Chang and Maisey 2003, fig. 12). Chang and Grande (1997) in contrast with Grande (1982) and Patterson and Rosen (1977) noted a first uroneural reaching the second preural centrum in †Ellimmichthys longicostatus. According to observations on BMNH P. 7109 we agree with older restorations. In †Scutatuspinosus, the first uroneural reach only the first preural centrum (cf. DGM 1250-P). Although Alvarado-Ortega et al. (2008) claimed an extension of the first uroneural to second preural centrum in †Triplomystus applegatei, it is not seen in the figure and photo of the original description (Alvarado-Ortega and Ovalles-Damian 2008, fig. 5). It is present in †Triplomystus noorae according to Forey et al. (2003).]

(31) Fusion of the first uroneural with the first ural centrum (Z32, AO 35): (0) absent; (1) present. [The pleurostyle is certainly a homoplastic feature shared by clupeiforms and ostariophysans, but absent is all non-clupeiform clupeomorphs.]

(32) Distal end of second uroneural (Z34, AO37): (0) reaching the distal end of the first uroneural (or according Z34 - "ascending part of the hypural 2 as long as the first uroneural"); (1) not reaching the distal end of the first uroneural (According Z34 - "shorter"). [The coding is based mainly on Zaragueta-Bagils (2004) except for modifications presented by Alvarado-Ortega et al. (2008) in which †Sorbinichthys and †Diplomystus solignaci show second uroneural not reaching the distal end of the first uroneural and in †Triplomystus the proximal end of the second uroneural extends as far as that of the first uroneural.]

(33) Parhypural (Z35, AO38): (0) fused with the first preural centrum; (1) autogenous. [Except for isolated and independent cases of secondary fusion, a separation of the parhypural from the first ural centrum is present in most of clupeiform clupeomorphs (Grande 1985). Among species examined, it is fused to preural centrum in †Scutatuspinosus (cf. DGM 1250-P) and †Codoichthys such as in remaining †ellimmichthyiforms.]

(34) Number of epurals (AO40, Z37, CM23): (0) Three; (1) Two. [Three epurals are usually found in living clupeoids and most of non-clupeiform clupeomorphs. †Codoichthys and †Scutatuspinosus have three epurals. Among species examined, only †Ezkutuberezi, and Denticeps exhibit two epural bones. The presence of epurals in †Sorbinichthys is uncertain according to Murray and Wilson (2011).]

(35) Position of epurals (Z38, AO41): (0) Filling space between the neural spines of the first and second preural centra and the first uroneural; (1) placed far from the spine of the second preural centrum, leaving an open space between them. [According to Alvarado-Ortega et al. (2008) we noted epurals filling space between neural arch and first uroneural in †Scutatuspinosus (cf. DGM 1250-P) unlike †Codoichthys. According to Murray and Wilson (2013) the state for †D. birdi is unknown.]

(36) Caudal scutes (Z39, AO42): (0) absent; (1) present. [We coded this character according to Murray and Wilson (2013). But we are unable to determine the occurrence of caudal scutes in †Codoichthys. In caudal skeleton of †Sorbinichthy elusivo (see Bannikov and Bacchia 2000, text-fig. 5) there are about five elements anterior to dorsal procurrent rays of difficult homology. They may be homologous to that two-three of †Paraclupea chetungensis which Chang and Grande (1997) interpreted as dorsal scutes.]

(37) Neural spine of the first preural centrum (modified of Z40, AO43): (0) first preural neural arch plus spine shorter or equal to half of the first uroneural; (1) first preural neural arch plus spine longer than half of the first uroneural [We noted problems with the original definition of this character as large and lanceolate (or even very slender) versus short or sub-rectangular. Poor preservation and deformation can produce subjective interpretation. Murray and Wilson (2011) described a long and fine neural spine in †Sorbinichthys africanus. †Scutatuspinosus (cf. DGM 1250-P) shows lanceolate neural spine and very elongate second neural spine as in †Sorbinichthys. The spine is large, elongate, and slender in †Diplomystus shengliensis, †D. dentatus, †Ellimmichthys longicostatus, and †Triplomystus applegatei (Alvarado-Ortega and Ovalles-Damián 2008, text-fig. 5). In †Triplomystus noorae it is short (see Forey et al. 2003, fig. 44). Also it is clearly short and sub-rectangular in †Ellimma branneri (see Chang and Maisey 2003, fig. 8). According to Cavender (1966, fig. 3A) Chirocentrus dorab shows large neural spine and arch complex disagreeing of codification furnished by Alvarado-Ortega et al. (2008) and Zaragueta-Bagils (2004).]

(38) Neural arch of the first ural centrum (Z42, AO44): (0) absent; (1) present.

(39) Predorsal scutes series (A45, MW 46): (0) absent; (1) present. [We assumed codification of Murray and Wilson (2013) except for †Ellimmichthys longicostatus and †Armigatus brevissimus in which we noted such series present.]

(40) Sub-rectangular scutes in the anterior part of predorsal series (Z47, MW 48): (0) absent; (1) present. [It is common in many non-clupeiform clupeomorphs the presence of sub-rectangular scutes since occiput to the origin of the dorsal fin. In certain advanced deep-bodied †paraclupeids, the anteriormost scutes are elliptical.]

(41) Sub-rectangular scutes in the posterior part of predorsal series (AO 47): (0) absent; (1) present.

(42) Spines on the posterior margin of predorsal scutes (Z48, AO48, MW 50): (0) absent; (1) present. [This feature was originally described as uniquely derived for †Diplomystus (see GRANDE 1982a). After, Santos and Corrêa (1985), claiming close affinities between †Diplomystus and †Scutatuspinosus on the basis of the form of the predorsal scutes pointed out the presence of spines on posterior border of scutes of the holotype of †Scutatuspinosus. However, Alvarado-Ortega et al. (2008) and Barros and Figueiredo (unpublished data 2011) indicated the posterior border all smooth except for the strong median spine. Herein we assumed the scalloped posterior border of †Horseshoeichthys as state 0 contra Murray and Wilson (2013).]

(43) Prominent median strong spine on posteriormost predorsal scutes (AO49, Z49, MW 51): (0) absent; (1) present. [This character is coded mainly based on Zaragueta-Bagils (2004). A median strong spine is present on the posteriormost scutes of many †ellimmichthyiform taxa such as †Ellimmichthys, †Ellimma, †Sorbinichthys, †Paraclupea, †Triplomystus, and † "Diplomystus" solignaci. This feature is also present on †Scutatuspinosus (cf. DGM 1164) but lacking in †Codoichthys. Unlike Murray and Wilson (2013) we coded primitive state for †Diplomystus birdi.]

(44) Size of scutes of predorsal series (Chang and Grande 1997; AO50, Z50; MW 52): (0) all equal-sized; (1) irregular, the size of scutes increases backwards in the series.

(45) Surface of predorsal scutes (CM15; Z51; AO51; MW 53): (0) smooth; (1) ornamented .

[This character is coded based on Alvarado-Ortega et al. (2008) except for †Scutatuspinosus that show fading radiating ornamentation at least in the posteriormost scutes (cf. DGM 1164).]

(46) Number of scutes in predorsal series (Z52, MW 54): [0] 6-14; [1] 16 to 19; [2] 20-41. [We coded according to Murray and Wilson (2013).]

(47) Abdominal scutes series (AO53; Z52; CM16): (0) absent; (1) present. [It is considered a synapomorphy of Clupeomorpha (Arratia 1997) besides certain basal taxa lack it (e.g., †Ornategulum, Chirocentrus).]

(48) Number of predorsal bones (MW 62): [0]10 or more; [1] 7-9; [2] 6 or fewer.

(49) Prepelvic and postpelvic scutes (modified of AO55, Z54, MW 57): (0) similar ; (1) prepelvic normal and postpelvic scutes with strong posterior spine; (2) both prepelvic and post pelvic with strong spines.

(50) Shape of lateral wings of abdominal scutes (modified of MW 59): (0) triangular; (1) spatulate; (2) quadrangular. [Most of clupeomorphs show triangular lateral wing of abdominal scute (Whitehead et al. 1985). In †Paraclupea, †Rhombichthys, †Tycheroichthys, and other deep-bodied †ellimmichthyiforms the scutes are more specialized becoming spatulate.] (51) Postdorsal scutes series (AO 58 ): (0) absent; (1) present. [As far as we known only species of †Tryplomystus show this feature.]

(52) Number of abdominal scutes (MW61): [0] fewer than 20; [1] 22-30; [2] more than 32. [In fact, some taxa will not show a number equivalent to vertebral counts. We coded according to Murray and Wilson (2013).]

(53) Postcleithra: (0) present; (1) absent. [†Triplomystus applegatei, †Horseshoeichthys armaserratus, †Codoichthys carnavalii, †Armigatus namourensis, †Tycheroichthys and †Thorectichthys show postcleithra. According to Murray and Wilson (2013) it is also found in †Diplomystus dentatus. This bone is usually present in living clupeiform fishes, commonly two in number. However, Greenwood (1968) stated occurrence of postcleithra in Denticeps uncertain.]

(54) Basisphenoid: (0) present; (1) absent. [Basisphenoid is found in Elopiformes and Clupeiformes (e.g., Sardinops, Denticeps, Engraulis, Alosa), †Ornategulum, and †Spratticeps (see Patterson 1970) but lack in †Diplomystus, †Triplomystus, †Ellimma, †Ellimmichthys, †Rhombichthys, and †Paraclupea. It is absent in the complete and well-preserved skull of a unnamed †Diplomystus from the English Chalk (Forey 2004). Its presence in †Scutatuspinosus and †Codoichthys is doubtful so that is coded with question mark on the matrix. Apparently its absence is a probable case of secondary loss for †Ellimmichthyiformes.]

(55) Third hypural (Poyato-Ariza et al. 2000): (0) smaller than or almost equal-sized the first hypural; (1) triangular, larger than others hypural bones. [Poyato-Ariza et al. (2000) pointed out that the third hypural is enlarged and triangular in most of non-clupeiform clupeomorphs and it penetrates diastema region projecting downwards unlike the condition found in clupeiforms. Thus the third hypural extends below occupying the diastema region.]

(56) Third hypural with upward process constraining the fourth hypural to project forwardly: (0) absent; (1) present. [It is only found in †Scutatuspinosus (cf. DGM 1250-P) and †Rhombichthys.]

(57) Two long and recumbent tubules of preopercular sensory canal on ventral limb: (0) absent; (1) present. [In †Codoichthys, †Ellimmichthys longicostatus and †Ellimma branneri there are two (rare three) distinct recumbent tubules emerging from the main preopercular sensory canal.]

(58) Striations on ventral part of opercle: (0) absent; (1) present. [It is found in adults of †Ellimma, †Tycheroichthys, †Codoichthys and possibly in †Thorectichthys marocensis (see Murray and Wilson, 2013, fig. 2).]

(59) Head deeper than long: (0) absent; (1) present. [Murray and Wilson (2013) commented (but not included in matrix) that †Tycheiroichthys and †Triplomystus shared a deep head unlike other †ellimmichthyiforms.].

(60) Parietal excavation: (0) absent; (1) present. [Forey (2004) recognized through a drawing-by-hand analysis a clade within †Ellimmichthyiformes composed of †Triplomystus, †Sorbinichthys, and †Diplomystus. One of the derived states of character used was the presence of a parietal excavation. According to him it is absent in †Armigatus, †Paraclupea, and †Ellimmichthys. We found it in †Codoichthys and †Scutatuspinosus (cf. DGM 1279-P). Its presence is very probable in †Ellimma cruzae but not in †E. branneri.]

Publication Dates

• Publication in this collection
  01 Sept 2016
• Date of issue
  Sept 2016

History

• Received
  29 July 2015
• Accepted
  18 Mar 2016