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Teleostean phylogeny based on osteological and myological characters/Filogenia de teleosteos basada en caracteristicas osteologicas y miologicas.

Clade C40: [158:1 [right arrow] 0]

This clade is supported by a bootstrap value of 78% and by an unambiguous synapomorphy: the presence of supramaxillae (158: 1 [right arrow] 0, among the ostariophysans examined, the four fossil taxa included in this clade are the only ones with supramaxillae; some characiforms do have supramaxillae, but this is seemingly a derived feature for the order: e.g. Fink & Fink, 1981, 1996). Whether these and other 'fossil Otophysi' (sensu Grande & De Pinna) are placed in a monophyletic group or not, their placement outside the clade including the four extant otophysan orders (Fig. 2) has important phylogenetic and evolutionary implications. For instance, this indicates that the characteristic Weberian apparatus of the members of these four extant otophysan orders was acquired only once, thus supporting the view of authors such as Fink & Fink (1981, 1996), Fink et al., (1984) and Patterson (1975) and contradicting those of authors such as Gayet (1981, 1985, 1986). We plan to discuss this subject and the respective evolutionary implications of the phylogenetic results obtained in further detail in a future work.

[dagger] Clupavus maroccanus: No unambiguous features; [dagger] Santanichthys diasii: [55:0 [right arrow] 1]

Clade 41: [80:0 [right arrow] 1]

Although there is only a single unambiguous synapomorphy uniting [dagger] Chanoides macropoma and [dagger] Lusitanichthys characiformis (centrum of third free vertebra markedly shorter than other surrounding centra), it is interesting to notice that this feature is homoplasy free within the numerous fishes included in the cladistic analysis of the present work, and also that this clade C41 is supported by a bootstrap value of 62%.

[FIGURE 29 OMITTED]

[dagger] Chanoides macropoma: [28:0 [right arrow] 1], [65:1 [right arrow] 0], [206:1 [right arrow] 0];

[dagger] Lusitanichthys characiformis: [43:0 [right arrow] 1], [75:1 [right arrow] 0], [102:0a1]

Clade 42: [69:1 [right arrow] 2], [72:1 [right arrow] 2], [76:1 [right arrow] 2], [78:1 [right arrow] 2]

This clade is supported by a bootstrap value of 75% and by four unambiguous synapomorphies: presence of characteristic scaphium (69: 1 [right arrow] 2), intercalarium (72: 1 [right arrow] 2), tripus (76: 1 [right arrow] 2) and os suspensorium (78: 1 [right arrow] 2). Apart these features, two other features with ambiguous distributions may be interpreted as synapomorphies of this clade if a 'fast optimization' is chosen (95: 0 [right arrow] 1, 'ligament between posttemporal and posterior margin of neurocranium' not ossified) or, alternatively, if a 'slow optimization' is chosen (83: 0 [right arrow] 1, perilymph system of inner ear peculiarly extended posteriorly, constituting sinus impar; coded as '?' in the five fossil taxa included in the cladistic analysis). See comments above for 'Clade C40'.

Clade 43: [22:1 [right arrow] 0], [28:0 [right arrow] 1], [32:1 [right arrow] 0], [49:0 [right arrow] 1], [62:0 [right arrow] 1], [88:0 [right arrow] 1], [150:0 [right arrow] [right arrow] 1], [186:0 [right arrow] 1], [187:0 [right arrow] 1]

The grouping of the cypriniform fishes examined is expected (see Fig. 1) and supported by a bootstrap value of 91%. Contrary to the other synapomorphies listed above, the arrector dorsalis not subdivided into different sections (22: 1 [right arrow] 0) and the dorsomesial limb of posttemporal (or posttemporo-supracleithrum) not markedly thin and mesially extended (88: 0 [right arrow] 1) were not proposed as potential cypriniform synapomorphies in previous works.

Clade 44: [133:0 [right arrow] 1]

The assembly of the two taxa examined belonging to the Cobitoidea is expected (see e.g. Siebert, 1987; Liu et al., 2002; Liu, 2004).

Catostomus: [93:0 [right arrow] 1], [140:0 [right arrow] 1], [145:0 [right arrow] 1], [216:0 [right arrow] 1]; Cobitis: [31:0 [right arrow] 1], [34:0 [right arrow] 1], [61:0 [right arrow] 1], [135:0 [right arrow] 1], [136:0 [right arrow] 1], [191:0 [right arrow] 1], [192:0 [right arrow] 1]

Clade 45: [150:1 [right arrow] 2], [182:0 [right arrow] 1]

The assembly of the three taxa examined belonging to the Cyprinoidea is expected (see e.g. Siebert, 1987; Liu et al., 2002; Liu, 2004). This clade 45 is supported by a bootstrap value of 56% (Fig. 2).

Opsariichthys: [137:0 [right arrow] 1], [140:0 [right arrow] 1], [142:0 [right arrow] 1], [191:0 [right arrow] 1]

Clade 46: [79:0 [right arrow] 1]

This clade 46 is supported by a bootstrap value of 77% (Fig. 2).

Danio: [223:1 [right arrow] 0], [253:0 [right arrow] 1]; Barbus: No unambiguous features

Clade 47: [16:1 [right arrow] 0], [33:0 [right arrow] 1], [70:0 [right arrow] 1], [93:0 [right arrow] 1], [131:1 [right arrow] 0], [160:1 [right arrow] 0], [174:1 [right arrow] 0], [206:1 [right arrow] 0], [242:1 [right arrow] 0]

The close relationship between the characiform, gymnotiform and siluriform fishes examined is expected (see Fig. 1) and is supported by a bootstrap value of 77% (Fig. 2). It should be noted that among the nine features characterizing this clade, six concern reversions to the plesiomorphic condition. Although this distribution appears to be the most parsimonious one by the strict application of the principle of parsimony, in such a discussion one should also analyze carefully each feature and discuss it in a critical way. That the first fishes of this clade C47 suffered a truly 'explosive morphological reversion', with two thirds of the characters diagnosing this clade being reversions, seems, at least at first sight, rather unsound. However, this does not mean necessarily that there is something wrong with this clade. In fact, the grouping of characiforms, gymnotiforms and siluriforms in a monophyletic unit has been strongly and repeatedly supported by numerous morphological (e.g. Fink, 1981, 1996; Lauder & Liem; Arratia, 1992; this study) and molecular (e.g. Dimmick & Larson, 1996; Saitoh et al.; Lavoue et al.) phylogenetic studies in the last years. What we think might be happening here is that by strictly applying the principle of parsimony, the absence of premaxillary (174, state 1) and mandibular (242, state 1) teeth in gonorynchiforms, cypriniforms and the fossil taxa examined is interpreted as the plesiomorphic condition for ostariophysans, later reverted in this clade 47. However, this could be an example of one of those cases in which evolution might not compulsorily work by following strict parsimony. That is, one can suppose, for instance, that these teeth were independently lost in gonorynchiforms, in cypriniforms, and in the fossil groups included in the cladistic analysis, instead of being completely lost in an evolutionary line of fishes of which part later completely reacquired them. The principle of parsimony may well be the common rule, and we do think that this is very likely the case (thence our commitment to phylogenetic analyses following the cladistic methodology), but there is no prove, so far, that exceptions to this rule are completely impossible in evolution. The example concerning the loss of teeth referred just above might be one of those exceptions.

[FIGURE 30 OMITTED]

Clade 48: [54:0 [right arrow] 1], [55:0 [right arrow] 1], [73:0 [right arrow] 1], [95:01 [right arrow] 0], [145:0 [right arrow] 1], [167:0 [right arrow] 1], [191:0 [right arrow] 1], [205:0 [right arrow] 1], [265:0 [right arrow] [right arrow] 1]

The grouping of the characiform fishes examined is expected (see Fig. 1) and is supported by a bootstrap value of 99% (Fig. 2). The present cladistic analysis did not support, but did neither contradict, the grouping of the distichodontid genera Distichodus and Xenocharax, nor the close relationship between these distichodontid taxa and the citharinid Citarhinus, as would be expected according to the works of authors such as Vari (1979), Orti & Meyer (1997) and Buckup (1998).

[FIGURE 31 OMITTED]

Xenocharax: [140:0 [right arrow] 1], [155:1 [right arrow] 0], [193:1 [right arrow] 0]; Distichodus: [1:1 [right arrow] 0], [128:0 [right arrow] 1], [133:0 [right arrow] 1], [212:0 [right arrow] 1], [243:0 [right arrow] 1]; Citharinus: [49:0 [right arrow] 1], [67:0 [right arrow] 1], [111:1 [right arrow] 0], [136:0 [right arrow] 1], [165:0 [right arrow] 1], [216:0 [right arrow] 1]; Brycon: [43:0 [right arrow] 1], [88:0 [right arrow] 1], [111:1 [right arrow] 0], [128:0 [right arrow] 1], [140:0 [right arrow] 1], [155:1 [right arrow] 0], [165:0 [right arrow] 1], [205:1 [right arrow] 0], [206:0 [right arrow] 1]

Clade 49: [34:0 [right arrow] 1], [61:0 [right arrow] 1], [79:0 [right arrow] 1], [94:0 [right arrow] 1], [190:0 [right arrow] 1], [225:0 [right arrow] 1]

As can be seen in Figure 1, most researchers now agree that gymnotiforms and siluriforms are sister-groups. However, this sister-group relationship has been mostly supported by morphological evidence (e.g. Fink & Fink 1981, 1996; Lauder & Liem; this study). Some molecular cladistic analyses published in the last years support, instead, a sister-group relationship between gymnotiforms and characiforms (e.g. Dimmick & Larson; Saitoh et al.; Peng et al., 2006) or possibly between characiforms and siluriforms (e.g. Lavoue et al.). The present cladistic analysis does provide strong evidence for a clade including siluriforms and gymnotiforms (Fig. 2). This is because it did not only corroborated many of the synapomorphies provided by Fink & Fink (1981, 1996) to support such a clade (34: 0 [right arrow] 1; 61: 0 [right arrow] 1; 79: 0 [right arrow] 1; 94: 0 [right arrow] 1), but also provided additional synapomorphies to support this clade, e.g. the ossification of the ligament connecting the suspensorium to the ethmoid region (190: 0 [right arrow] 1) and the interhyal (ossified or not) being connected by ligaments to both the hyoid arch and the suspensorium (225: 0 [right arrow] 1) (see Appendix 2). In fact, this clade C49 is supported by a bootstrap value of 93% (Fig. 2).

Clade 50: [39:0 [right arrow] 1], [130:0 [right arrow] 1], [140:0 [right arrow] 1], [180:0 [right arrow] 1], [216:0 [right arrow] 1]

This clade is expected (see Fig. 1) and is supported by a bootstrap value of 95%. The second and third synapomorphies listed above (130: 0 [right arrow] 1, levator arcus palatini markedly lateral to all bundles of adductor mandibulae; 140: 0 [right arrow] 1, insertion of a significant part of adductor arcus palatini on lateral surface of suspensorium) concern muscular features that were not proposed as potential gymnotiform synapomorphies in previous works.

Brachyhypopomus: [1:1 [right arrow] 0], [174:0 [right arrow] 1], [242:0 [right arrow] 1]

Clade 51: [13:0 [right arrow] 1], [151:0 [right arrow] 1]

[FIGURE 32 OMITTED]

[FIGURE 33 OMITTED]

[FIGURE 34 OMITTED]

[FIGURE 35 OMITTED]

Authors such as De La Hoz (1974), Albert & Camposda-Paz (1998) and Albert (2001) defended that the sternopygid gymnotiforms are more closely related to hypopomids than to gymnotids. Authors such as Triques (1993), Gayet et al., (1994) and Alves-Gomez et al., (1995) have, instead, argued that hypopomids are more closely related to gymnotids than to sternopygids. Curiously, the two synapomorphies listed above, which concern muscular features that are homoplasy free within the fishes examined, support a closer relationship between the sternopygid Sternopygus and the gymnotid Gymnotus than between any of these taxa and the hypopomid Brachyhypopomus. This clade C51 is supported by a bootstrap value of 57% (Fig. 2).

Sternopygus: [16:0 [right arrow] 1], [193:1 [right arrow] 0], [254:0 [right arrow] 1]; Gymnotus: [94:1 [right arrow] 0]

Clade 52: [22:1 [right arrow] 0], [48:0 [right arrow] 1], [49:0 [right arrow] 1], [59:0 [right arrow] 1], [74:0 [right arrow] 1], [86:0 [right arrow] 1], [88:0 [right arrow] 1], [92:0 [right arrow] 1], [99:0 [right arrow] 1], [100:0 [right arrow] 1], [103:0 [right arrow] 1], [111:1 [right arrow] 0], [136:0 [right arrow] 1], [201:0 [right arrow] 1], [208:0 [right arrow] 1], [247:0 [right arrow] 1], [256:0 [right arrow] 1], [266:0 [right arrow] 1]

The assembly of the siluriform fishes examined is expected (see Fig. 1) and is supported by a bootstrap value of 100%. Some of the features listed above have been listed in previous phylogenetic works as potential synapomorphies of the Siluriformes (e.g. 49: 0 [right arrow] 1; 59: 0 [right arrow] 1; 74: 0 [right arrow] 1; 86: 0 [right arrow] 1; 88: 0 [right arrow] 1; 99: 0 [right arrow] 1; 100: 0 [right arrow] 1; 136: 0 [right arrow] 1; 247: 0 [right arrow] 1) 201: 0 [right arrow] 1; 208: 0 [right arrow] 1; 256. 0 [right arrow] 1; 266: 0 [right arrow] 1; see e.g. Regan, 1911; Chardon, 1968; Roberts, 1973; Lundberg, 1975; Howes, 1983ab; Fink & Fink, 1981, 1996; Arratia, 1987; Schaefer, 1990; Mo, 1991; Arratia, 1992; De Pinna, 1993, 1998; Diogo, 2004) but others seemingly constitute additional potential synapomorphies to diagnose the order, such as: arrector dorsalis not subdivided into different sections (22: 1 [right arrow] 0); frontal and autopterotic not contacting in dorsal view (48: 0 [right arrow] 1); absence of 'ligament between posttemporal and posterior margin of neurocranium' (92: 0 [right arrow] 1); presence of coracoid bridge (103: 0 [right arrow] 1); adductor mandibulae attaching exclusively on mandible and/or primordial ligament, near its mandibular insertion (111: 1 [right arrow] 0). Within this clade C52, the phylogenetic scenario shown in Fig. 2 is essentially similar to that of Diogo's recent work. As Diogo has already provided a detailed discussion of the siluriform clades obtained in the present work, as well as a comparison with previous works on catfish phylogeny, we will only list the synapomorphies supporting these clades C53, C54, C55, C56, C57, C58 and C59, which are supported by bootstrap values of 51%, 73%, 63%, 53%, 66%, 70% and 54%, respectively (Fig. 2).

Diplomystes: [137:1 [right arrow] 0], [155:0 [right arrow] 1]; Clade 53: [16:0 [right arrow] 1], [132:1 [right arrow] 0], [138:1 [right arrow] 0]; Clade 54: [128:0 [right arrow] 1], [133:0 [right arrow] 1]; Callichthys: [12:0 [right arrow] 1], [91:0 [right arrow] 1], [101:0 [right arrow] 2], [174:0 [right arrow] 1], [256:1 [right arrow] 0], [257:0 [right arrow] 1]; Clade 55: [207:0 [right arrow] 1], [216:0 [right arrow] 1], [220:0 [right arrow] 1], [252:0 [right arrow] 1]; Nematogenys: [103:1 [right arrow] 0], [138:2 [right arrow] 1], [140:0 [right arrow] 1], [149:0 [right arrow] 1], [230:0 [right arrow] 1], [231:0 [right arrow] 1], [267:0 [right arrow] 1]; Trichomycterus: [145:0 [right arrow] 1], [147:0 [right arrow] 1]; Clade 56: [5:0 [right arrow] 1], [25:0 [right arrow] 1], [267:0 [right arrow] 1]; Cetopsis: [51:0 [right arrow] 1], [99:1 [right arrow] 0], [122:0 [right arrow] 1], [141:0 [right arrow] 1], [144:0 [right arrow] 1], [214:1 [right arrow] 0]; Clade 57: [26:0 [right arrow] 1]; Silurus: [38:0 [right arrow] 1], [48:1 [right arrow] 0], [256:1 [right arrow] 0]; Clade 58: [16:1 [right arrow] 0], [101:0 [right arrow] 2]; Chrysichthys: [193:1 [right arrow] 0]; Clade 59: [189:0 [right arrow] 1]; Bagrus: [102:0 [right arrow] 1]; Pimelodus: [42:0 [right arrow] 1], [48:1 [right arrow] 0], [152:0 [right arrow] 1], [205:0 [right arrow] 1]

CONCLUSIONS

The elopomorphs appear grouped in a monophyletic clade, which is the sister-group of the clade including all the other teleosts included in the cladistic analysis. The osteoglossomorphs examined are also included in a monophyletic clade, which appears as the sister-group of the remaining non-elopomorph teleostean fishes examined. The clupeomorph and ostariophysan fishes are grouped together, thus contradicting the results of some recent molecular cladistic analyses placing the Alepocephaloidea inside the Otocephala. In fact, the monophyly of the Argentiniformes (Alepocephaloidea + Argentinoidea) is well supported in the cladistic analysis of the present work. This cladistic analysis also provides strong for the monophyly of the Alepocephaloidea, of the Argentinoidea, of the Galaxioidea + Osmeroidea, and of the Esociformes. However, it does not provide strong evidence to resolve the relationships between the Argentiniformes, Salmoniformes, Esociformes, Osmeriformes and Neoteleostei, although it does indicate that the salmoniforms might be closely related to the Neoteleostei and that the Esociformes and the Osmeriformes might constitute a monophyletic unit. The monophyly of the Cypriniformes + Characiformes + Gymnotiformes + Siluriformes, of the Characiformes + Gymnotiformes + Siluriformes and of the Gymnotiformes + Siluriformes is well supported.

ACKNOWLEDGEMENTS

We specially thank J. Snoeks, E. Vreven, and the late G.G. Teugels (Musee Royal de l'Afrique Centrale), P. Laleye (Universite Nationale du Benin), R. Vari, J. Williams and S. Jewett (National Museum of Natural History), T. Grande (Field Museum of Natural History), D. Catania (California Academy of Sciences), M. Stiassny (American Museum of Natural History), Mark Sabad and J. Lundberg (Academy of Natural Sciences of Philadelphia), L. Page and M. Retzer (Illinois Natural History Survey) and P. Pruvost and G. Duhamel (Museum National d'Histoire Naturelle) for kindly providing a large part of the specimens analyzed for this study. We are particularly grateful to L. Taverne and Michel Chardon, who undertook a revision of a previous version of the paper and contributed to it with valuable comments and suggestions.

Appendix 1: Material Examined

A list of the specimens examined of the extant genera included in the cladistic analysis is given below; the trypsine-cleared and alizarine-stained (cands) or alcohol fixed (alc) condition of the studied fishes is given in parentheses following the number of specimens dissected (AMNH: American Museum of Natural History; ANSP: Academy of Natural Sciences of Philadelphia; CAS: California Academy of Sciences; FMNH: Field Museum of Natural History; INHS: Illinois Natural History Survey; LFEM: Laboratory of Functional and Evolutionary Morphology of the University of Liege; MNCN: Museo Nacional de Ciencias Naturales; MNHN: Museum National d'Histoire Naturelle; MRAC: Musee Royal de l'Afrique Centrale; UNB: Universite Nationale du Benin; USNM: National Museum of Natural History):

Outgroup: Amia calva: MNCN 35961, 2 (alc), 1 (cands). Lepisosteus osseus: ANSP 107961, 2 (alc); ANSP 172630, 1 (alc); MNCN 246557, 1 (cands). Lepisosteus platyrhincus: AMNH 74789, 2 (alc).

Osteoglossomorpha: Hiodon tergisus: MNCN 36019, 3 (alc). Mormyrus niloticus: LFEM, 1 (alc). Mormyrus tapirus: MNCN 80593, 3 (alc); MNCN 85283, 1 (alc). Pantodon buchholzi: MNCN 73493, 4 (alc). Xenomystus nigri: MNCN 227824, 25 (alc).

Elopomorpha: Albula vulpes: MNCN 52124, 2 (alc). Anguilla anguilla: MNCN 41049, 3 (alc). Elops lacerta: LFEM, 2 (alc). Elops saurus: MNCN 48752, 2 (alc). Conger conger: MNCN 1530, 5 (alc). Eurypharynx pelecanoides: AMNH 44315, 1 (alc); AMNH 44344, 1 (alc). Megalops cyprinoides: MNCN 48858, 3 (alc). Notacanthus bonaparte: MNCN 107324, 3 (alc).

Clupeomorpha: Denticeps clupeoides: MRAC 76-032-P-1, 2 (alc). Engraulis encrasicolus: MNCN 68048, 2 (alc); MNCN 65097, 8 (alc); MNCN 1099, 3 (alc). Engraulis sp: MNCN 48896, 3 (alc). Ethmalosa fimbriata: MNCN 48865, 3 (alc). Ilisha fuerthii: MNCN 49338, 8 (alc). Thryssa setirostris: MNCN 49294, 2 (alc).

Ostariophysi: Bagrus bajad: LFEM, 1 (alc), 1 (cands). Bagrus docmak: MRAC 86-07-P-512, 1 (alc). Barbus barbus: LFEM, 1 (cands). Barbus guiraonis: MNCN 245730, 3 (alc). Brachyhypopomus brevirostris: LFEM, 2 (alc). Brachyhypopomus sp: INHS 89761, 2 (alc). Brycon guatemalensis: MNCN 180536, 3 (alc). Brycon henni: CAS 39499, 1 (alc). Callichthys callichthys: USNM 226210, 2 (alc). Catostomus commersonii: MNCN 36124, 10 (alc). Citharinus sp.: 86-016-P-72, 3 (alc). Cetopsis coecutiens: USNM 265628, 2 (alc). Chanos chanos: USNM 347536, 1 (alc), LFEM, 1 (alc). Chrysichthys auratus: UNB, 2 (alc). Chrysichthys nigrodigitatus: LFEM, 1 (cands). Cobitis paludica: MNCN 248076, 7 (alc). Cromeria nilotica: MRAC P.141098, 2 (alc). Danio rerio: LFEM, 5 (alc). Diplomystes chilensis: LFEM, 3 (alc). Distichodus notospilus: MRAC A0-048-P-2630, 3 (alc). Gonorynchus gonorynchus: LFEM, 2 (alc). Gonorynchus greyi: FMNH 103977, 1 (alc). Grasseichthys gabonensis: MRAC 73-002-P-264, 3 (alc). Gymnotus carapo: ILNS 35493, 2 (alc); MNCN 115675, 2 (alc).

Kneria wittei: MRAC P-33512, 2 (alc). Nematogenys inermis: USNM 084346, 2 (alc). Opsariichthys uncirostris: MNCN 56668, 3 (alc). Parakneria abbreviata: MRAC 99-090-P-703, 3 (alc). Phractolaemus ansorgii: MRAC P.137982, 3 (alc). Pimelodus blochii: LFEM, 2 (alc), 1 (cands). Silurus aristotelis: LFEM, 2 (alc). Silurus glanis: LFEM, 2 (alc). Sternopygus macrurus: CAS 48241, 1 (alc); INHS 62059, 2 (alc). Trichomycterus areolatus: LFEM, 2 (alc). Xenocharax spilurus: MRAC A0-048-P-2539, 3 (alc).

Euteleostei: Alepocephalus rostratus: MNCN 108199, 2 (alc). Argentina brucei: USNM 239005, 2 (alc). Argentina sphyraena: MNCN 001134, 12 (alc); MNCN 78530, 5 (alc). Astronesthes niger: MNCN 1102, 1 (alc). Aulopus filamentosus: MNCN 1170, 6 (alc). Bathylagus euryops: MNCN 124597, 1 (alc). Bathylagus longirostris: USNM 384823, 2 (alc). Bathylagus tenuis: MNHN 2005-1978, 2 (alc). Chlorophthalmus agassizi: MNCN 1193, 3 (alc); MNCN 1182, 5 (alc). Coregonus lavaretus: MNCN 75424, 1 (alc). Coregonus tugun: MNCN 75422, 2 (alc). Esox lucius: MNCN 197706, 5 (alc). Galaxias maculatus: USNM 344889, 2 (alc). Osmerus eperlanus: MNCN 193795, 11 (alc). Osmerus mordax: USNM 32565, 2 (alc). Plecoglossus altivelis: MNCN 192036, 1 (alc). Retropinna retropinna: AMNH 30890, 1 (alc). Salmo trutta: MNCN 136179, 2 (alc), 1 (cands); MNCN 16373, 2 (alc); MNCN 40685, 2 (alc). Salmo sp: MNCN 48863, 2 (alc). Searsia koefoedi: USNM 206896, 2 (alc). Stokellia anisodon: AMNH 31037, 1 (alc). Stomias boa: MNCN 74444, 8 (alc); MNCN 74456, 4 (alc). Thymallus thymallus: MNCN 115147, 1 (alc); MNCN 114992, 1 (alc). Umbra limi: MNCN 35672, 2 (alc); 36072, 2 (alc). Umbra krameri: MNCN 36659, 3 (alc). Xenodermichthys copei: MNCN 78950, 2 (alc); MNCN 1584, 2 (alc); USNM 215527, 2 (alc).

Appendix 2: List of characters included in the cladistic analysis.

A list of the characters included in the cladistic analysis is given below. Unless stated otherwise, the nomenclature of the morphological structures mentioned in this list follows that of Diogo. Due to limitations of size, this list of characters will be kept as short and simple as possible. Also, it is obviously not possible to include, in the present paper, anatomical drawings to illustrate all the numerous morphological features mentioned in this list. However, apart the numerous anatomical drawings provided in the present paper, an effort was made in order to supply, for most of the characters, references to figures provided in previous works in which the configuration corresponding to the derived states of these characters has been illustrated. In this way, we also want to pay a tribute to previous works done on teleostean comparative anatomy and phylogeny. It is important to note that, unless explicitly stated otherwise, all the morphological features mentioned in the list below refer exclusively to the adult configuration. It should also be noted that in the matrix shown in Table 1, inapplicable and missing character states for a certain taxon are indicated with '-' and with '?', respectively. Unless otherwise stated, inapplicable characters are used in cases in which for example a certain character refers to the shape of a bone that is not found in a certain taxon; missing character states are used in cases in which it was not possible to appropriately discern the respective state in a certain taxon (e.g. due to the poor preservation of the fossils or of the extant specimens dissected) (for more details see Diogo). As stressed by authors such as Hilton & Bemis (1999), there are documented cases, within actinopterygian fishes, of remarkable morphological variation within a single genus, a single species, and even within a single population of the same species. As will be mentioned throughout the list of 271 characters given below, among these characters there are effectively cases in which different wild-type, adult members (examined by us and/or previously described in the literature) of a certain terminal taxon do seemingly exhibit different character states of a same character. Since in those cases one cannot assign the wild-type, adult members of the respective terminal taxon in which such a variation occurs to a single character state, the taxon is coded as '?' (see below).

Ventral cephalic musculature.

1. Two sections of intermandibularis (anterior and posterior intermandibularis). [0] Absent. [1] Present (e.g. Fig. 3; Greenwood, 1971: fig. 10). As shown in Edgeworth (1935), figure 271, in the members of the genus Albula dissected in the present work the intermandibularis is not subdivided into these two sections.

2. Posterior intermandibularis and interhyoideus (ordered multistate character). [0] Not associated to each other. [1] Posterior intermandibularis deeply associated with interhyoideus, forming the protractor hyoidei (e.g. Fig. 3), but also deeply associated with anterior intermandibularis. [2] Posterior intermandibularis deeply associated with interhyoideus, forming the protractor hyoidei, and not associated with anterior intermandibularis (e.g. Figs. 4, 5, 6; Greenwood, 1971: fig. 7). Greenwood (1971) stated that in osteoglossomorph notopterids, including the members of the genus Xenomystus, the posterior intermandibularis and the interhyoideus do not form a protractor hyoidei. However, in the Xenomystus specimens analyzed in the present work, it is not completely clear that this is effectively the case. This because the muscle named 'posterior intermandibularis' in Greenwood's (1971) page 21 does appear to have a myocommata dividing its anterior and posterior portions, which is very similar to the myocommata dividing the anterior and posterior portions (i.e. posterior intermandibularis and interhyoideus portions) of the protractor hyoidei in many other teleosts. Also, in the Xenomystus specimens examined, the muscle named 'interhyoideus' in Greenwood's (1971) page 21 is very similar to the 'hyohyoideus inferioris' (sensu Winterbottom's 1974 and the present works) of many other teleosts. Contrary to the interhyoideus of teleosts such as Albula and Mormyrus, the muscle named 'interhyoideus' in Greenwood's (1971) page 21 does not attach anteriorly to the lower jaw: instead, it fuses anteroventromesially with its counterpart and inserts onto the hypohyals through a short tendon, as is precisely the case with the hyohyoideus inferior of many other teleosts. Until more detailed data is available, Xenomystus will be prudently coded as '?'.

3. Interhyoideus. [0] Present (either as an independent element or included in the protractor hyoidei) (e.g. Fig. 3). [1] Missing.

4. Anterior intermandibularis. [0] Attaching partially or exclusively on dentary bones (e.g. Figs. 3, 4, 5, 6). [1] Not attaching on dentary bones (e.g. Howes, 1985: figs. 20, 21).

5. Peculiar differentiation of protractor hyoidei into pars dorsalis, pars ventralis and pars lateralis. [0] No differentiation (e.g. Figs. 3, 4, 5, 6). [1] Differentiation (e.g. Diogo, 2004: fig. 3-41).

6. Protractor hyoidei. [0] Not inserted high on mandibular coronoid process. [1] Inserted high on mandibular coronoid process (e.g. Howes, 1985: fig. 5).

7. Hyohyoideus abductor. [0] Present (e.g. Figs. 3, 4, 5, 6). [1] Missing.

8. Hyohyoideus inferior and hyohyoideus abductor. [0] Blended to each other (e.g. Fig. 3). [1] Not blended to each other (e.g. Fig. 4; Diogo, 2004: fig. 3-44).

9. Presence of hyohyoidei adductores. [0] Not missing (e.g. Figs. 3, 4, 5, 6). [1] Missing.

10. Hyohyoideus abductor and hyohyoidei adductores. [0] Not highly modified (e.g. Figs. 3, 4, 5, 6). [1] Highly modified, being hypertrophied and deeply blended to each other (e.g. Greenwood, 1971: figs. 10, 12).

11. Hyohyoidei adductores. [0] Not covering a significant part of the lateral margin of the cranium. [1] Covering a significant part of the lateral margin of the cranium (e.g. Edgeworth: fig. 279).

12. Significant part of hyohyoideus abductor and/or hyohyoidei adductores. [0] Not attaching to pectoral girdle. [1] Attaching to pectoral girdle.

13. Hyohyoideus ventralis. [0] Absent (e.g. Figs. 3, 4, 5, 6). [1] Present (e.g. De la Hoz & Chardon, 1984: fig. 16).

14. Branchiomandibularis. [0] Present (e.g. Lauder: fig. 3B). [1] Absent (e.g. Figs. 3, 4, 5, 6).

Musculature associated with pectoral girdle and fins.

15. Presence of sternohyoideus. [0] Present (e.g. Figs. 4, 5, 6). [1] Absent.

16. Anteroventromesial portion of hypoaxialis. [0] Not continuous with posteroventromesial portion of sternohyoideus (e.g. Lauder: figs. 2B, 3B). [1] Continuous with posteroventromesial portion of sternohyoideus (e.g. Diogo: fig. 3-113).

17. Sternohyoideus. [0] Not consolidated into a single median muscle (e.g. Lauder: figs. 2B, 3B). [1] Consolidated into a single median muscle (e.g. Figs. 4, 5, 6; Diogo, 2004: fig. 3-121).

18. Distinct muscle 'arrector 3'. [0] Absent (e.g. Figs. 7, 8A). [1] Present (e.g. Figs. 9, 10, 11, 12). This muscle was curiously not described in Winterbottom's 1974 work. One possible explanation is that this muscle is only found in some teleostean taxa examined, being absent in fishes such as elopomorphs and osteoglossomorphs, as well as in non-teleosts such as Amia and Lepisosteus (e.g. Figs. 7, 8). The names of the pectoral girdle muscles used in works on catfishes such as Diogo et al. (2000) and Diogo do not fully correspond to the nomenclature proposed by Winterbottom. One of the main reasons for this is precisely due to the presence, in catfishes, of the muscle 'arrector 3', which was not described by Winterbottom and was tentatively named arrector ventralis by Diogo et al. (2001a) and Diogo. In the present work the nomenclature of the pectoral muscles essentially follows that used by Winterbottom. Thus, in order to facilitate comparisons with Diogo et al.' (2000) and Diogo' previous works on catfishes, it should be stressed that the "arrector ventralis" of those works corresponds to the 'arrector 3' of the present work; the "arrector dorsalis", "abductor superficialis 1", "abductor superficialis 2", "adductor superficialis 1", "adductor superficialis 2" and "abductor profundus" of those works correspond respectively to the arrector ventralis, abductor superficialis, abductor profundus, adductor superficialis, adductor profundus and arrector dorsalis of Winterbottom and of the present work.

19. Distinct muscle coracoradialis [0] Absent (e.g. Figs. 8B, 13, 14, 15, 16, 17, 18). [1] Present (e.g. Fig. 19; Winterbottom: fig. 38).

20. Distinct muscle arrector ventralis. [0] Absent (e.g. Fig. 7). [1] Present (e.g. Figs. 8A, 9, 10, 11, 12, 13, 14; Winterbottom: fig. 32).

21. Abductor superficialis and/or abductor profundus. [0] Not hypertrophied (e.g. Fig. 18). [1] Hypertrophied, a significant part also originating on mesial surface of pectoral girdle. (e.g. Greenwood & Thompson, 1960: figs. 6, 8).

22. Arrector dorsalis (ordered multistate character). [0] Not subdivided (e.g. Fig. 16). [1] Subdivided into two well-developed sections (e.g. Fig. 8B). [2] Subdivided into three well-developed sections (e.g. Fig. 19).

23. Attachment of arrector dorsalis. [0] Not attaching on both the first and second pectoral rays (e.g. Fig. 16). [1] Attaching on both the first and second pectoral rays (e.g. Fig. 8B).

24. Protractor pectoralis. [0] Present: according to Greenwood & Lauder the presence of a recognizable protractor pectoralis is seemingly the plesiomorphic condition for actinopterygians, being present in at least some extant members of e.g. the Acipenseriformes, Polypteriformes and Amiiformes (e.g. Fig. 11; Greenwood & Lauder: fig. 2). [1] Absent as an independent element. As noted by these authors, in some members of the genus Galaxias this muscle is missing, while in others it is present as an independent element; this genus is thus coded as '?'.

25. Significant part of mesial portion of arrector ventralis. [0] Not passing through coracoid-cleithrum foramen. [1] Passing through coracoid-cleithrum foramen (e.g. Diogo: fig. 3-56).

26. Arrector ventralis divided into two distinct, well-developed bundles widely separated by large horizontal lamina of coracoid (or scapulo-coracoid). [0] Not divided (e.g. Fig. 8B). [1] Divided (e.g. Diogo: fig. 3-56).

27. Arrector ventralis peculiarly divided into well-developed posterodorsal and anteroventral bundles, both originating on ventrolateral surface of pectoral girdle. [0] Not divided (e.g. Fig. 8A). [1] Divided.

Neurocranium, anterior vertebrae and related structures.

28. 'Kinethmoid' bone. [0] Absent (e.g. Figs. 20, 21, 22, 23, 24). [1] Present (e.g. Fig. 10; Fink & Fink, 1981: fig. 2A).

29. Rostrodermethmoids. [0] Not ossified and/or fused with median supraethmoid. [1] Ossified and not fused with median supraethmoid (e.g. Sanford 2000: figs. 10, 11, 12). According to authors such as Jollie (1975) and Wilson & Veilleux (1982) the adults of genera as e.g. Esox and Umbra have paired rostrodermethmoids (sensu Sanford, 2000) fused with paired endoskeletal structures. However, as stressed by Sanford (2000), this does not invalidate the coding of Exox and Umbra as CS-1, since the specimens of these two genera examined in the present work do fulfill the two requirements needed to be coded as CS-1: their rostrodermethmoids are ossified and are not fused with a median supraethmoid (if the rostrodermethmoids were completely fused with a median, unpaired supraethmoid, the compound structure resulting from this fusion would not be constituted by two lateral, paired portions that never completely fuse in the midline, as is the case in the specimens examined of these two genera). According to authors such as Patterson (1973) the ossified 'pre-ethmoids' of Amia seemingly correspond to ventral ethmoid ossifications, and not to the rostrodermethmoids found in certain teleosts (see character below); therefore Amia is coded here as CS-0.

30. Ventral ethmoids. [0] Not ossified and/or fused with prevomer (e.g. Fig. 21). [1] Ossified and not fused with prevomer (e.g. Sanford, 2000: figs. 10, 11, 12).

31. Mesethmoid. [0] If present, not fused with prevomer (e.g. Fig. 21). [1] Fused with prevomer (e.g. Ramaswami, 1953: fig. 1).

32. Anteroventral margin of prevomer. [0] Does not situate well posteriorly to anteroventral margin of mesethmoid (or of supraethmoid and/or rostrodermethmoids: see above) (e.g. Fig. 21). [1] Situates well posteriorly to anteroventral margin of mesethmoid (or of supraethmoid and/or rostrodermethmoids) (e.g. Taverne, 1974: figs. 2, 4; Fink & Fink, 1981: figs. 2C,D, 4A).

33. Anterolateral processes of mesethmoid (or of supraethmoid and/ or rostrodermethmoids) supporting and/or articulating with premaxillae. [0] Not present (e.g. Fig. 21). [1] Present (e.g. Fig. 24; Fink & Fink, 1981: fig. 3C,D, E, F). As pointed out by Gayet (1985: 109), it is difficult to discern this character appropriately in [dagger] Lusitanichthys characiformis. However, the illustrations provided in Gayet (1981, 1985) suggest that in the members of this species the anterolateral margins of the mesethmoid are mainly associated with the proximal surfaces of the maxillae, and not with the premaxillae ("anterolaterally, the mesethmoid presents a profound depression that receives the articular process of the maxilla": Gayet, 1981, p. 175). Thus [dagger] Lusitanichthys characiformis does not seem to exhibit CS-1.

34. Posterodorsal portion of mesethmoid (or of supraethmoid). [0] Not appearing markedly compressed transversally when seen in dorsal view. [1] Appearing markedly compressed transversally when seen in dorsal view (e.g. Fink & Fink, 1981: fig. 3E, F).

35. 'Ethmoid endoskeleton'. [0] Not markedly reduced ossification of 'ethmoid endoskeleton' (sensu Patterson & Johnson, 1996: p. 254) (e.g. Fig. 23). [1] Markedly reduced ossification of the 'ethmoid endoskeleton' (e.g. Fig. 24; Sanford, 2000: fig. 12). The Astronesthes specimens dissected exhibit a condition similar to that described for CS-1, while certain members of this genus appear to exhibit a condition similar to that described for CS-0 (see e.g. Weitzman, 1967a, b); this genus is thus coded as '?'.

36. Mesethmoid. [0] If present, not peculiarly shaped, markedly compressed anteroposteriorly and expanded transversally (e.g. Fig. 21). [1] Peculiarly shaped, being markedly compressed anteroposteriorly and expanded transversally (e.g. Thys van den Audenaerde, 1961: fig. 13).

37. Lateral ethmoids. [0] Not exhibiting remarkably large, peculiar lateral extensions (e.g. Fig. 22). [1] Exhibiting remarkably large, peculiar lateral extensions (e.g. Greenwood et al., 1966: fig. 6; Grande & Poyato-Ariza: fig. 4B).

38. Thin, elongated lateral laminar projection of lateral ethmoid contacting autosphenotic. [0] Not present (e.g. Fig. 22). [1] Present (e.g. Bornbusch, 1995: fig. 5B).

39. Peculiar anteroventrolateral, anteroventrally pointed process of lateral ethmoid. [0] Not present (e.g. Fig. 22). [1] Present (e.g. De la Hoz & Chardon, 1984: figs. 1, 4).

40. Prevomer. [0] Paired (e.g. Mayhew, 1924: fig. 3). [1] Unpaired (e.g. Taverne, 1974: fig. 4). As stressed by authors such as Patterson (1975) and De Pinna, some specimens of the genera Hiodon and Osmerus may exhibit CS-O, while others exhibit CS-1; these two genera are thus coded as '?'.

41. Large, strong 'pseudocartilaginous ligament' between prevomer, autopalatines and/or mandibles. [0] Not present. [1] Present (e.g. Thys van den Audenaerde, 1961: fig. 19).

42. Prevomerine teeth. [0] Present. [1] Absent (e.g. Fig. 21; Fink & Fink, 1981: fig. 2). In Sanford's (2000) table II, this author states that the members of the genus Alepocephalus have prevomerine teeth. However, this seems to be an error, because all the Alepocephalus specimens examined in the present work and in the works of authors such as Gegenbaur (1878) and Gosline (1969) do not have prevomerine teeth.

43. Rhinosphenoid. [0] Absent (e.g. Fig. 21). [1] Present (e.g. Fig. 23; Fink & Fink, 1981: fig. 2C). The presence/absence of a rhinosphenoid in [dagger] Lusitanichthys characiformis has been subject of controversy (see e.g. Gayet, 1985). In our opinion, the position of Gayet's (1981, 1985) 'rhinosphenoid' seems to be somewhat similar to that of the rhinosphenoid of some characiforms. Moreover, Cavin (1999) described a new species of [dagger] Lusitanichthys, [dagger] L. Africans, which, according to this author, also appears to have a 'rhinosphenoid'. We therefore consider that we cannot simply discard a priori the hypothesis that the 'rhinosphenoid' of these authors might be homologous to the rhinosphenoid of certain characiforms. These structures will thus be tentatively coded here as primary homologues; this primary homology will be tested against the results of the cladistic analysis including all the available characters. Cavin (p. 692) stated that he "observed a trace of a subrectangular bone behind the lateral ethmoid on several specimens of [dagger] Clupavus maroccanus that could be interpreted as a rhinosphenoid'. Since in Taverne's (1977a, 1995) descriptions of [dagger] Clupavus maroccanus this author does not make any reference to the possible presence of a rhinosphenoid, we prefer to prudently code this latter species as '?'.

44. Independent orbitosphenoid. [0] Present (e.g. Fig. 21). [1] Absent (e.g. Fig. 22; Smith, 1989a: fig. 4). Authors such as Jollie (1975) and Arratia (1997, 1999) have described/coded the orbitosphenoid as present in Esox and/or Thymallus. However, the 'orbitosphenoid' of Jollie's (1975) figure 10 clearly does not seem to correspond to the orbitosphenoid of the present work; in the Esox and Thymallus specimens examined in the present work, as well as in works of other authors such as Norden (1961), Johnson & Patterson (1996) and Sanford (2000), the orbitosphenoid is absent as an independent ossification.

45. Independent basisphenoid. [0] Present (e.g. Fig. 9). [1] Absent (e.g. Diogo: 3-66). Sanford (2000) coded Esox as not having a basisphenoid. However, all Esox specimens examined in the present work and in the works of other authors such as Jollie (1975) and Johnson & Patterson do exhibit an independent basisphenoid.

46. Frontals. [0] Not markedly separated from each other along dorsal midline. [1] Markedly separated from each other along dorsal midline (e.g. Grande & Poyato-Ariza: fig. 4B).

47. Autopterotic and dermopterotic. [0] At least one of these bones is not present as an independent ossification (e.g. Fig. 21). [1] Both these bones are present as independent, distinct ossifications (e.g. Gosline, 1969: fig. 6; Greenwood & Rosen: figs. 22, 24). As explained by authors such as Patterson (1973) and Grande & Bemis (1998), it is difficult to discern if both the autopterotic and dermopterotic are or not present as independent ossifications in Lepisosteus; this genus is thus coded as '?'.

48. Frontal and autopterotic (and/or dermospterotic). [0] Contacting in dorsal view (e.g. Fig. 20). [1] Not contacting in dorsal view (e.g. Diogo: fig. 3-67).

49. Vertical, complete laminar bony connection mesially to the eye between frontal, dorsally, and parasphenoid, ventrally. [0] Not present (e.g. Fig. 20). [1] Present (e.g. Fig. 10; Diogo: fig. 3-112). Some specimens of the genera Pantodon and Amia exhibit CS-0 (e.g. Taverne, 1978: fig. 31; Jollie, 1984a: fig. 13B), while others exhibit CS-1 (e.g. Kershaw, 1970: fig. 3; Patterson, 1973: fig. 9B); these genera are thus coded as '?'.

50. Frontals. [0] Not completely fused along midline. [1] Completely fused along midline (e.g. Smith, 1989b: fig. 505E). Forey et al. coded the elopomorph saccopharyngiforms as having the frontals fused along the midline. However, as explained by authors such as Regan (1912) and Tchernavin (1947a, b), this is not the case in Eurypharynx.

51. Peculiar anterodorsal process of pterosphenoid contacting posterodorsal process of lateral ethmoid. [0] Not present (e.g. Fig. 10). [1] Present (e.g. Diogo: fig. 3-46).

52. Prootic and/or pterotic bulla lodging diverticulum of swimbladder. [0] Not present (e.g. Fig. 25). [1] Present (e.g. Grande, 1986: fig. 31).

53. Sacculi and lagenae. [0] Not lying in a posterior position and/ or not lying near the midline. [1] Lying in a posterior position and near the midline (e.g. Rosen & Greenwood, 1970: fig. 15B,C). Filleul & Maisey reported a 'lagenar capsule' (see below) on the exoccipital and also the basioccipital of [dagger] Santanichthys diasii; this seems to indicate that the members of this taxon exhibited CS-1. 54. Peculiar, large 'auditory foramen', which is usually an ovoid opening on the ventral face of the prootic through which the utricular otolith is visible. [0] Not present. [1] Present (e.g. Weitzman, 1962: fig. 4).

55. Markedly large, globular 'lagenar capsule'. [0] Not present (e.g. Fig. 26). [1] Present (e.g. Fig. 25; Weitzman, 1962: fig. 4). Gayet (1985) considered that it was not possible to discern the presence/ absence of this feature in the specimens of [dagger] Lusitanichthys characiformis that she analyzed. However, in Gayet's (1985) figure 23 the posterior portion of the basioccipital is illustrated, and a markedly large, globular 'lagenar capsule' such as that present in the taxa listed under CS-1 does not seem to be present.

56. Peculiar 'recessus lateralis'. [0] Absent (e.g. Fig. 20). [1] Present (e.g. Fig. 9; Forey, 1973a, b; Grande, 1986: figs. 27, 31; Di Dario, 2004: figs. 3, 4).

57. Exoccipitals. [0] Not markedly expanded posterolaterally (e.g. Fig. 26). [1] Markedly expanded posterolaterally, extending well over neural arch of first free vertebra (e.g. Poyato-Ariza, 1996; Grande & Poyato-Ariza).

58. Exoccipitals and basioccipital. [0] Nor completely fused (e.g. Fig. 25). [1] Completely fused.

59. Parietals (or parieto-extrascapulars). [0] Not fused with supraoccipital (e.g. Fig. 21). [1] Fused with supraoccipital (e.g. Fink & Fink, 1981: fig. 5D).

60. Main bodies of parietals (or of parieto-extrascapulars). [0] Not widely separated from each other in dorsal view (e.g. Fig. 21). [1] Widely separated from each other in dorsal view (e.g. Fig. 24; Sanford, 2000: fig. 27). Some members of the genera Esox, Thymallus and Coregonus exhibit CS-0, while others exhibit CS-1 (this study; see also descriptions of e.g. Norden, 1961; Shaposhnikova, 1967; Sanford, 2000; Grande et al., 2004); these genera are thus coded as '?'.

61. Intercalar. [0] According to authors such as Jollie (1986) the intercalar is primitively present as an independent ossification is neopterygians (e.g. Fig. 25). [1] Intercalar absent as independent ossification (e.g. Fig. 26; Patterson, 1973: fig. 11A).

62. Markedly enlarged, ventral 'pharyngeal process' of basioccipital (sensu Winterbottom, 1974). [0] Not present (e.g. Fig. 25). [1] Present (e.g. Vandewalle, 1975; Fink & Fink, 1981: fig. 5D).

63. Markedly elongated, posteriorly pointed posterior process of parasphenoid. [0] Not present (e.g. Fig. 25). [1] Present (e.g. Ridewood, 1904a: fig. 122).

64. Teeth on parasphenoid. [0] Present. [1] Absent (e.g. Albert, 2001: fig. 2).

65. Basipterygoid process. [0] According to authors such as Patterson (1973: p. 254), the "loss of the basipterygoid process, as in Amia and most teleosts" is a "derived condition"; this view is supported by the recent work of Moritz & Britz (2005). [1] Absence of 'basipterygoid process' (e.g. Taverne, 1974: fig. 4). Taverne (1977a) stated that a 'basipterygoid process' is missing in [dagger] Clupavus maroccanus, but Taverne (1995) stated that such a processus was in fact perhaps present in this taxon; until more data is available, we prefer to prudently code this taxon as '?'.

66. Supraoccipital. [0] Not ossified. [1] Ossified (e.g. Fig. 21; Patterson, 1973: fig. 3A,B).

67. 'Highly ossified triangular pars sustentaculum complex'. [0] Not present. [1] Present (e.g. Vari: fig. 32).

68. Anteroposterior elongation of anterior neural arches. In order to properly define this character, it should be mentioned that contrary to taxa of CS-0 [0] (e.g. Tchernavin, 1947a: fig-text 10) in the specimens examined of taxa of CS-1 [1] the length of at least part of at least one of the most anterior neural arches is almost equal, or even greater, than that of the centrum of the free vertebra to which it is associated (e.g. Fig. 27; Grande, 1994: figs. 6, 7).

69. Scaphium (ordered multistate character). [0] Not present (e.g. Fig. 27). As pointed out by authors such as Patterson (1984) and Fink & Fink (1996), in the fishes of CS-1 [1] (e.g. Patterson, 1984: figs. 13-16) the scaphium does not seem to be as derived from the plesiomorphic condition found in fishes of CS-0 as it is in the fishes of CS-2 [2] (e.g. Fink & Fink, 1981: figs. 14-18).

70. Anterior margin of neural arch of third free vertebra. [0] Not closely approaching posterior border of neurocranium (e.g. Fig. 27). [1] Closely approaching posterior border of neurocranium (e.g. Fink & Fink, 1981: figs. 15-18).

71. Claustrum. [0] Not present (e.g. Fig. 27). [1] Present (e.g. Fink & Fink, 1981: figs. 14-17). According to authors such as De Pinna & Grande (2003) and Grande & De Pinna the claustrum of the fishes coded under CS-1 is homologous with the 'accessory neural arch' of some other teleosts. If this is effectively the case, the modification of this 'accessory neural arch' in the characteristic claustrum of the fishes of CS-1 will constitute a derived feature anyway.

72. Intercalarium (ordered multistate character) [0] Not present (e.g. Fig. 27). As pointed out by authors such as Patterson (1984) and Fink & Fink (1996), in the fishes of CS-1 [1] (e.g. Patterson, 1984: figs. 13-16) the intercalarium does not seems to be as derived from the plesiomorphic condition found in those fishes coded under CS-0 as it is in the fishes coded under CS-2 [2] (e.g. Fink & Fink, 1981: figs. 14-18).

73. Peculiar, prominent anterodorsal process of neural arch of third free vertebra. [0] Not present (e.g. Fig. 27). [1] Present (e.g. Fink & Fink, 1981: fig. 15) [State 1].

74. Parapophyses of second free vertebra [0] Not markedly reduced in size or missing (nor completely fused with ribs) (e.g. Fig. 27). [1] Markedly reduced in size or missing (or occasionally completely fused with ribs) (e.g. Fink & Fink, 1981: fig. 17) [State 1].

75. Parapophyses of two first free vertebrae. [0] Not fused to centra (e.g. Fig. 27). [1] At least some parapophyses of two first free vertebrae fused to centra. (e.g. Fink & Fink, 1981: fig. 14).

76. Tripus (ordered multistate character). [0] Not present (e.g. Fig. 27). As pointed out by authors such as Patterson (1984) and Fink & Fink (1996), in the fishes coded under CS-1 [1] (e.g. Patterson, 1984: figs. 13-16) the tripus does not seems to be as derived from the plesiomorphic condition found in the fishes coded under CS-0 as it is in the fishes coded under CS-2 [2] (e.g. Fink & Fink, 1981: figs. 14-18).

77. Ribs/parapophyses of third free vertebra. [0] Not highly modified in relation to, and/or broader than, the ribs/parapophyses of the following free vertebrae. [1] Highly modified in relation to, and/or broader than, ribs/parapophyses of the following free vertebrae (e.g. Grande & Poyato-Ariza: fig. 12). According to authors such as Rosen & Greenwood, Fink & Fink (1981, 1996), Patterson (1984) and Grande & De Pinna the fishes exhibiting a tripus (see above) should be coded under CS-1 in a character such as this, because the tripus is seemingly at least partially constituted by enlarged ribs/parapophyses of the third free vertebra. Fink & Fink (1996) suggested that Kneria and Parakneria might have a configuration such as that described for CS-1, although they recognized that in the members of these two genera the ribs/ parapophyses of the third free vertebral column are only slightly broader than those of the following free vertebrae. In the Kneria and Parakneria specimens examined by us, as well as in those specimens illustrated by Lenglet (1974: figs. 17-19), the ribs/ parapophyses of the third free vertebra are not considerably larger than those of the following free vertebrae. Also, Grande & Poyato-Ariza seem to suggest that Grasseichthys and Cromeria should be coded as CS-1, but the illustrations of Grande (1994: figs. 6, 9) seem to indicate that the ribs/parapophyses of the third free vertebra are not considerably broader than those of the following free vertebrae. Until more data is available, these four genera will be prudently coded here as '?'.

78.'Os suspensorium' (ordered multistate character). [0] Not present (e.g. Fig. 27). As pointed out by authors such as Patterson (1984) and Fink & Fink (1996), in the fishes coded as CS-1 [1] (e.g. Patterson, 1984: figs. 13, 14, 16) the 'os suspensorium' does not seem to be as derived from the plesiomorphic condition found in the fishes coded as CS-0 as it is in the fishes coded as CS-2 [2] (e.g. Fink & Fink, 1981: figs. 14-18).

79. Highly modified, ovoid peculiar anterolateral face of 'transverse process' of fourth free vertebra. [0] Not present (e.g. Fig. 27). [1] Present (e.g. Fink & Fink, 1981: fig. 17, 18). Fink & Fink (1981, 1996) did not mention the presence of this feature in cypriniforms, but the Danio and Barbus specimens examined by us do exhibit a highly modified, ovoid anterolateral face of the 'transverse process' of the fourth free vertebra that is similar to that found in many gymnotiforms and siluriforms.

80. Centrum of third free vertebra. [0] Not markedly shorter than surrounding centra (e.g. Fig. 27). [1] Markedly shorter than surrounding centra (e.g. Patterson, 1984: figs. 13, 14; Gayet, 1985: figs. 22-25).

81. Peculiar silvery peritoneal tunic of swimbladder covering at least part of its anterior portion. [0] Not present. [1] Present (e.g. Rosen & Greenwood: fig. 4). As noted by these authors, the swimbladder is missing in Gonorynchus specimens, but these specimens do have remains of a silvery peritoneal tunic associated to the structures of the anterior free vertebrae.

82. Swimbladder. [0] Not markedly divided into peculiar anterior and posterior chambers. [1] Markedly divided into peculiar anterior and posterior chambers (e.g. Rosen & Greenwood: fig. 4).

83. Perilymph system of inner ear. [0] Not peculiarly extended posteriorly. [1] Peculiarly extended posteriorly, constituting sinus impar (e.g. Chardon et al., 2003: fig. 3.4).

84. Peculiarly large, distinct 'precervical gap' filled mainly with connective tissue between first free vertebra and neurocranium. [0] Not present (e.g. Fig. 26). [1] Present (e.g. Rosen, 1985: figs. 14, 15). This character has been the subject of controversy. For example, authors such as Rosen (1985) stated that members of non-neoteleostean taxa as e.g. Osmerus might also have a large 'precervical gap'. This statement was however contradicted by authors such as Johnson & Patterson (p. 278), which stated that "in osmeroids the articulation between the occipital condyle and V1 is normally close". Within the non-aulopiform and non-stomiiform taxa examined in the present work there were effectively some specimens that appeared to have a 'precervical gap', although this gap was not as peculiarly large as that found in the specimens examined of the genera Aulopus, Chlorophthalmus, Astronesthes and Stomias.

Pectoral girdle and fins.

85. Posttemporal. [0] Present (e.g. Fig. 28, 29). [1] Absent (e.g. Smith, 1989a: fig. 9).

86. Supracleithrum. [0] Present as independent ossification (e.g. Fig. 28). [1] Absent as independent ossification (e.g. Diogo, 346).

87. Main body of posttemporal (or posttemporo-supracleithrum). [0] Not lying considerably far from neurocranium (e.g. Fig. 23). [1] Lying considerably far from neurocranium, with almost no contact between these two structures. The association between the posttemporal (or posttemporo-supracleithrum) and the neurocranium is thus a rather feeble one made essentially through the 'ligament between the posttemporal and the posterior margin of the neurocranium' described below and/or occasionally through thin/small extrascapulars (e.g. Fig. 24; Monod: fig. 45).

88. Dorsomesial limb of posttemporal (or posttemporosupracleithrum). [0] Markedly thin and mesially extended (e.g. Fig. 21). [1] Not markedly thin and mesially extended (e.g. Figs. 9, 10, 23).

89. Scapula. [0] Ossified (e.g. Fig. 8). [1] Not ossified (e.g. Fig. 15; Jollie, 1984a: fig. 16).

90. Coracoid. [0] Ossified (e.g. Fig. 8). [1] Not ossified (e.g. Fig. 15; Jollie, 1984b: fig. 19).

91. Baudelot's ligament. [0] Present (e.g. Fig. 25). [1] Absent. Authors such as Patterson & Johnson stated that gonorynchiforms do not have a Baudelot's ligament. However, most gonorynchiforms examined by us exhibit a well-developed, paired ossification that is usually named 'cephalic rib' but that is strikingly similar to the ossified Baudelot's ligament of teleosts such as catfishes, connecting the cleithrum and/or supracleithrum to the posteromesial surface of the neurocranium (usually exoccipital and/or basioccipital) (e.g. Fig. 26). A potential homology between such gonorynchiform peculiar 'cephalic rib' (e.g. Fig. 26) and the 'Baudelot's ligament' of other teleostean fishes (e.g. Fig. 25) has actually been already proposed by authors such as Ridewood (1905b). Until more detailed data is available on this subject, we prefer to code all the gonorynchiform fishes examined by us as '?', with exception to Grasseichthys, in which such peculiar 'cephalic ribs' are completely missing; this latter genus is thus coded as CS-0. Patterson & Johnson stated that in the elopomorph Notacanthus the Baudelot's ligament is missing. However, the specimens of this genus examined by us do have a Baudelot's ligament, which is effectively peculiarly shaped, but is similar to that found in other elopomorph fishes such as Anguilla and Conger: it is markedly thin transversally and markedly broad anteroposteriorly, attaching to various free vertebrae.

92. 'Ligament between posttemporal and posterior region of neurocranium (usually intercalar)'. [0] Present (e.g. Figs. 21, 25, 28, 29). [1] Absent. This ligament corresponds to the 'posttemporal-intercalar ligament' of authors such as Taverne (1974), but because in some cases (e.g. when the intercalar is missing) it may attach to other bones such as the autopterotic, we prefer to use the less restrictive name 'ligament between posttemporal and posterior margin of neurocranium'.

93. Baudelot's ligament. [0] Contacting mesially with anterior free vertebrae, being exclusively attached to neurocranium. [1] Not contacting mesially with anterior free vertebrae, being exclusively attached to neurocranium (e.g. Fig. 25; Fink & Fink, 1981: fig. 19B,C,D).

94. Ossification of Baudelot's ligament. [0] Not ossified (e.g. Fig. 25). [1] At least partially ossified (e.g. Fink & Fink, 1981: fig. 19C,D). Some Hiodon specimens exhibit CS-0 while others exhibit CS-1 (see e.g. Taverne, 1977b); this genus is thus coded as '?'.

95. 'Ligament between posttemporal and posterior region of neurocranium'. [0] At least partially ossified (e.g. Fig. 25). [1] Not ossified. Taverne (1972, 1977b, 1978) stated that in the osteoglossomorphs Hiodon, Xenomystus and Mormyrus there is no "process of the posttemporal for the intercalar" (that is, there is no ossification of the ligament mentioned in the present character). However, in the specimens examined of these three genera the ligament is, in fact, ossified (e.g. Fig. 29; see also Figs. 21, 28). The presence of an ossified ligament in the specimens of these three genera can be effectively overlooked, for example in analyses that do not include the observation of soft structures. This because these specimens display a peculiar configuration in which the anteroventrolateral surface of the posttemporal is bifurcated anteriorly into a shorter, lateral arm that is essentially a tubular structure carrying a sensorial canal, and a longer, mesial arm that extends well anteriorly to the lateral one and that is attached by a thick ligament to the intercalar (or occasionally the autopterotic in Mormyrus, in which the interhyal is missing) (e.g. Fig. 29; see also Fig. 21). This latter arm thus clearly seems to be homologous to the ossified 'ligament between the posttemporal and the posterior region of neurocranium' found in many other teleosts. In the Pantodon specimens examined by us the ligament is very thick, but is not ossified.

96. Posttemporal. [0] As explained above, although many other taxa may have an ossified 'ligament between the posttemporal and the posterior margin of the neurocranium', their posttemporal is not peculiarly bifurcated anteroventrolaterally in a shorter, lateral arm carrying a sensorial canal and a longer, mesial arm that corresponds to the ossified 'ligament between posttemporal and posterior margin of neurocranium' of the present work. [1] Such a peculiar configuration of the posttemporal is only found in the specimens examined of the osteoglossomorph genera Hiodon, Xenomystus and Mormyrus (see e.g. Figs. 21, 29).

97. Cleithrum. [0] Present as independent ossification (e.g. Fig. 20). [1] Absent as independent ossification (e.g. Tchernavin, 1947a: text-fig. 14).

98. Deep, long, curved fossa on lateral surface of cleithrum (e.g. Fig. 20). [0] Not present. [1] Present.

99. Bifurcation of cleithrum. [0] Not markedly bifurcated dorsally into well-developed anterodorsal and posterodorsal arms for articulation with supracleithrum (or posttemporo-supracleithrum) (e.g. Fig. 28). [1] Markedly bifurcated dorsally into well-developed anterodorsal and posterodorsal arms for articulation with supracleithrum (or posttemporo-supracleithrum) (e.g. Taverne, 1972: fig. 1; 1978: fig. 44; Diogo, fig. 3-58).

100. Compound bone scapulocoracoid. [0] Not present (e.g. Fig. 28). [1] Present (e.g. Gosline, 1977; Diogo et al., 2001a: fig. 12).

101. Mesial limb of coracoids (or scapulo-coracoids) (ordered multistate character). [0] Not broad and anteroposteriorly elongated (e.g. Fig. 8B). [1] Broad and anteroposteriorly elongated, but does not meet its counterpart in a strong median interdigitation (e.g. Brosseau, 1978a: fig. 1). [2] Broad and anteroposteriorly elongated, meeting its counterpart in a strong median interdigitation (e.g. Fig. 28; Diogo, fig. 3-56). Some specimens of the genus Galaxias exhibit CS-0, while others seemingly exhibit CS-1 (see e.g. Swinnerton, 1903; McDowall, 1969); this genus is thus coded as '?'.

102. Prominent posteroventral process on ventral surface of coracoid. [0] Not present (e.g. Fig. 28). [1] Present (e.g. Fig. 17; Diogo: fig. 3-40A) (such a process, found in a few teleosts, should not be confused with the posterior process that is present on the posterolateral surface of the coracoid (or scapulo-coracoid) of numerous teleostean fishes: see e.g. Brosseau, 1987b: fig. 8).

103. 'Coracoid bridge'. [0] Not present (e.g. Fig. 28). [1] Present (e.g. Diogo, fig. 3-56).

104. Mesocoracoid arch. [0] Present (ossified or not) (e.g. Fig. 28). [1] Either undifferentiated or completely fused anteriorly with the posterior margin of the coracoid and/or scapula (e.g. Fig. 19; Smith, 1989a: fig. 9).

105. Ossification of mesocoracoid arch. [0] Among those fishes examined with a mesocoracoid arch (see above), in fishes such as Lepisosteus and Amia this structure is not ossified (e.g. Fig. 16); this is also the case in other basal Actinopteri such as many members of the extant genera Acipenser, Psephurus and Polyodon (e.g. Jollie, 1980; Mabee & Noordsy, 2004; Hilton, pers. comm.). [1] Mesocoracoid arch ossified (e.g. Fig. 28; Taverne, 1974: 23).

106. Mesocoracoid arch and coracoid and/or scapula. [0] Among those fishes examined by us exhibiting an ossified mesocoracoid arch, the plesiomorphic condition is seemingly that in which this arch is not firmly and rigidly attached, through suture or complete fusion, to the coracoid and/or scapula. This is the case in e.g. elopomorph fishes such as Elops, Megalops and Albula and osteoglossomorph fishes such as Hiodon and Mormyrus, in which the mesocoracoid arch articulates ventrally with the coracoid and/ or scapula and, thus, in which this arch has some mobility in relation to these latter bones (e.g. Fig. 8B; 28; Taverne, 1974: fig. 23). This is also the case in e.g. some basal Actinopteri such as some members of the acipenseriform genus Acipenser with an ossified mesocoracoid arch (Hilton, pers. comm.). [1] The taxa coded as CS-1 have an ossified mesocoracoid arch that is firmly and rigidly associated, often through suture or complete fusion, to the coracoid and/or scapula (or scapulo-coracoid) (e.g. Fig. 18; Diogo et al., 2001a: fig. 12).

107. Length of mesocoracoid arch. [0] Not markedly elongated dorsoventrally (e.g. Fig. 28). [1] Markedly elongated dorsoventrally (e.g. Taverne, 1978: figs. 30, 44).

108. Shape of mesocoracoid arch. [0] Not markedly enlarged transversally (e.g. Fig. 28). [1] Markedly enlarged transversally (e.g. Taverne, 1978: figs. 30, 44). Some specimens of Mormyrus exhibit CS-0, while others display CS-1 (Taverne, 1972; this study); this genus is thus coded as '?'.

109. 'Pectoral splints'. [0] Present (e.g. Figs. 7, 8). [1] Absent (e.g. Fig. 11; Gosline, 1980: fig. 2).

110. First pectoral ray. [0] Not articulating directly with scapula and/or occasionally with coracoid. [1] Articulating directly with scapula and/or occasionally with coracoid (e.g. Fig. 8B; Jessen, 1872; Gosline, 1980: fig. 3).

Lateral cephalic musculature.

111. Adductor mandibulae. [0] Attaching only on mandible and/ or primordial ligament, near its mandibular insertion (e.g. Fig. 9). [1] Attaching not only on mandible and/or primordial ligament, near its mandibular insertion, but also on other structures (e.g. maxilla, lacrimal and/or other bones) (e.g. Fig. 10; Bishai, 1967: fig. 2; Kershaw, 1976: fig.23).

112. Adductor mandibulae insertion on bones other than the mandible. [0] When most lateral bundles of the adductor mandibulae attach also, or exclusively, on bony structures other than the mandible, the non-mandibular insertions include only the maxilla (e.g. Fig. 10). [1] When most lateral bundles of adductor mandibulae attach also, or exclusively, on bony structures other than the mandible, the non-mandibular insertions include bones other than maxilla, such as those of the infraorbital series and/or the dermopalatine/autopalatine (e.g. Greenwood, 1977: figs. 10, 11, 21).

113. Adductor mandibulae A1-OST. [0] Not present (e.g. Fig. 9, 24, 30, 31, 32, 33). [0] Present (e.g. Figs. 10, 22, 23, 34; Diogo & Chardon, 2000: figs. 1-11; Diogo & Vandewalle, fig. 2.3).

114. Well-developed, dorsolateral bundle A1 of the adductor mandibulae. [0] Not present (e.g. Fig. 9). [1] Presence of well-developed, dorsolateral bundle A1 of the adductor mandibulae (sensu Diogo & Chardon; not to be confused with the adductor mandibulae A1-OST of these authors: see above) (e.g. Fig. 33; Diogo & Chardon, fig. 1).

115. Distinct adductor mandibulae A3-MAX. [0] Not present (e.g. Fig. 9). [1] The name adductor mandibulae A3-MAX as not been used so far, but the use of this new name for this adductor mandibulae section seems to be the best option, because there has been much confusion concerning its nomenclature and homologies in the literature. The adductor mandibulae A3-MAX corresponds to the 'A1-b' of authors such as Greenwood (1977: e.g. figs. 3, 10, 11, 12), Winterbottom and Wu & Shen (e.g. Fig. 2). However, as stressed by these authors, many other names have been used to designate this adductor mandibulae section in the literature, as e.g. 'pterygo-maxillaire', 'pterygo-maxillaris', or 'levator maxillae superior'. The position and origin of this section is actually somewhat similar to that of the A3'/A3'' of other fishes examined in the present work (see below), the main difference being that the A3-MAX attaches on the maxilla and not on the mandible. That is why, in order to differentiate this section from the A3'/A3'' of other teleosts, we use the name A3-MAX.

116. Adductor mandibulae sections 'palatomandibularis minor' and 'palatomandibularis major'. [0] Not present (e.g. Fig. 9). [0] Present (e.g. Fig. 30; Lauder fig. 2A). In Lauder's table II it is suggested that the adductor mandibulae sections 'palatomandibularis minor' and 'palatomandibularis major' of Lepisosteus are likely homologous to the 'levator maxillae superioris 3 and 4' of Amia, because all these structures represent an 'anterior division' of the adductor mandibulae. However, the overall shape, position and attachments of the 'palatomandibularis minor' and 'palatomandibularis major' of Lepisosteus are markedly different from those of the 'levator maxillae superioris 3 and 4' of Amia. Just to give an example, the 'palatomandibularis minor' and 'palatomandibularis major' of Lepisosteus originate dorsally on the ectopterygoid/entopterygoid and insert ventrally on the mandible, while the section 3 of the 'levator maxillae superioris 3 and 4' of Amia originates dorsally on the neurocranium and orbital bones and inserts ventrally mainly on the autopalatine (e.g. Lauder, fig. 3A).

117. Adductor mandibulae sections 'levator maxillae superioris 3 and 4'. [0] Not present (e.g. Fig. 9). [0] Present (e.g. Lauder, fig. 3A) (see character above).

118. 'Abductor mandibulae'. [0] Not present (e.g. Fig. 9). [1] In Eurypharynx a part of the adductor mandibulae has differentiated into an 'abductor mandibulae' (e.g. Tchernavin, 1947a: figs. 4, 5), which inserts on the posterior end of the mandible behind the mandibulo-quadrate articulation, its contraction thus helping to open, and not to close, the mouth.

119. Peculiar bundle of adductor mandibulae markedly extended anteriorly in order to attach to the anterodorsal surface of mandible. [0] Not present (e.g. Fig. 30). [1] Present (e.g. Bishai, 1967: figs. 1-3).

120. Quite peculiar configuration of A1-OST, its anterior portion being almost perpendicular to its posterior portion. [0] Not present (e.g. Fig. 22). [0] Present (e.g. Howes, 1985: figs. 18, 19).

121. Several small, peculiar tendons branching off from A2. [0] Not present (e.g. Fig. 9). [1] Present (e.g. Howes, 1985: figs. 18, 19).

122. Origin of adductor mandibulae A1-OST. [0] Not extending to neurocranium (e.g. Fig. 22). [1] Extending to neurocranium (e.g. Diogo, 2004: fig. 3-43).

123. Peculiar, well-differentiated bundle A1-OST-m running from anteroventral surface of quadrate to maxilla. [0] Not present (e.g. Fig. 22). [1] Present ('a1i' of Howes's 1985 figs. 11, 12, 16).

124. Attachment of mainly undivided A2 on mesial surface of mandible. [0] Not accomplished by means of two well-distinguished, thick tendons (e.g. Fig. 30). [1] Accomplished by means of two well-distinguished, thick tendons, the most lateral one usually attaching to the coronomeckelian and the most mesial one usually attaching posteriorly to the Aw (e.g. Fig. 33). It should be noted that we consider 'Inapplicable' not only those cases in which there is no 'mainly undivided A2' but also in which there is a 'mainly undivided A2 not attaching on the mesial surface of the mandible by means of two well-distinguished tendons' but not an A3'. This is because the A3 may well be the result of the differentiation of a dorsal part of the A2 plus the incorporation of one of those two ventral tendons attaching on the mesial surface of the mandible in fishes coded as CS-1.

125. Insertion of adductor mandibulae A2. [0] Not directly inserted on anteromesial surface of dentary bone (e.g. Fig. 30). [1] Directly inserted on anteromesial surface of dentary bone s (e.g. Sanford, 2000: figs. 95, 96).

126. Recognizable dilatator operculi. [0] Not present (e.g. Fig. 22). [1] Present (e.g. Tchernavin, 1947: text-figs. 4, 5).

127. Recognizable adductor operculi. [0] Not present (e.g. Fig. 22). [1] Present (e.g. Tchernavin, 1947: text-figs. 4, 5).

128. Dilatator operculi. [0] Not markedly lateral to A2 (e.g. Fig. 22). [1] Markedly lateral to A2 (e.g. Fig. 23; Diogo, fig. 3-88).

129. Adductor mandibulae A2. [0] Not exhibiting strong tendon that is perpendicular to its main body and that connects this bundle to the anteroventral surface of quadrate (e.g. Fig. 22). [1] Exhibiting strong tendon that is perpendicular to its main body and that connects this bundle to the anteroventral surface of quadrate (e.g. Howes, 1985: fig. 16).

130. Levator arcus palatini. [0] Not markedly lateral to all bundles of adductor mandibulae (e.g. Fig. 22). [1] Markedly lateral to all bundles of adductor mandibulae (e.g. De la Hoz & Chardon: fig. 13).

131. Well-differentiated section A3' of adductor mandibulae. [0] As explained by Lauder (1980: table II), the plesiomorphic condition for actinopterygians is seemingly that in which there are two 'mesial adductor mandibulae divisions', as is the case in the Amia and Lepisosteus specimens examined by us (e.g. Lauder: figs. 2A, 3A). The two mesial adductor mandibulae divisions found in Amia and Lepisosteus correspond to the A3' and A3'' of Diogo & Chardon and of the present work (see Figs. 30, 31), and thus the presence of A3' and A3'' is accordingly coded here as CS-0. [1] No well-differentiated section A3' (e.g. Fig. 32). Sanford (fig. 94) reported an A3' in members of Galaxias. However, in the Galaxias specimens examined by us some fibers of the A2 lie mesial to the levator arcus palatini but ventrally these fibers meet, and deeply blend with, the remaining fibers of the A2. Galaxias is thus coded here as '?'. It should be noted that we consider that one of the adductor mandibulae bundles attaching on the coronomeckelian bone in the members of the genus Albula corresponds to an A3', and, thus, this genus is coded here as CS-0.

132. Well-differentiated, separated section A3'' of adductor mandibulae. [0] Present (e.g. Fig. 30, 31). [1] Absent (e.g. Fig. 23) (see character above).

133. Adductor mandibulae Aw. [0] Present (e.g. Fig. 31). [1] Absent (e.g. Fig. 30; Howes 1985: fig. 16). The bundle named 'Aw' in Vari's (1979) figure 42 of Distichodus corresponds to part of the A1-OST of Diogo & Chardon and not to an Aw.

134. Division of Aw. [0] Not divided into well-developed, distinct Aw-D and Aw-V bundles (e.g. Fig. 31). [1] Divided into well-developed, distinct Aw-D and Aw-V bundles, with the Aw-V attaching anteriorly to the suspensorium and/or opercular series (e.g. Fig. 33). Gosline (1986, 1989) described the adductor mandibulae section Aw of Aulopus japonicus as an undivided section that does not attach to the suspensorium and/or the opercular series. The Aulopus specimens examined by us do have an Aw divided into a well-developed dorsal Aw-D bundle and a well-developed Aw-V bundle (e.g. Fig. 33) attaching posteriorly on the opercular series. This is also the case of the specimens examined of the other aulopiform genus analyzed, Chlorophthalmus, as well as of several other aulopiform and non-aulopiform eurypterygians (see Fig. 1) described in the literature, which exhibit a configuration strikingly similar to that found in the Aulopus specimens examined by us (see e.g. Winterbottom, 1974; Gosline, 1986; Sato & Nakabo, 2002; Wu & Shen). We thus consider that it is very likely that Gosline (1986, 1989) failed to recognize the Aw-V in the Aulopus specimens examined by him. In fact, in our first observations of the Aulopus specimens we also failed to recognize the Aw-V. Not because this bundle was small or really absent, but because when one separates the mandible from the other cephalic structures, this bundle tends to remain attached to the opercular series. Therefore, when one then analyses the separated mandible and the structures attached to it, one can effectively easily fail to recognize the Aw-V. 135. Small bundle of adductor mandibulae attaching to lateral ethmoid by means of a thin, long tendon. [0] Not present (e.g. Fig. 22). [1] Present.

136. Adductor arcus palatini and autopalatine and/or dermopalatine. [0] Adductor arcus palatini, or muscle differentiated from it, not inserting on autopalatine and/or dermopalatine (e.g. Fig. 22). [1] Adductor arcus palatini, or muscle differentiated from it, inserting on autopalatine and/or dermopalatine (e.g. Takahasi, 1925: figs. 8, 9, 12).

137. Adductor arcus palatini. [0] Part of adductor arcus palatini and/or of muscle differentiated from it not deeply blended with ligaments connecting the anterior region of the suspensorium and the ethmoid region. [1] Part of adductor arcus palatini and/or of muscle differentiated from it deeply blended with ligaments connecting the anterior region of the suspensorium and the ethmoid region (e.g. Diogo, fig. 3-73).

138. Extensor tentaculi (ordered multistate character). [0] Whiting those fishes in which there an insertion of the adductor arcus palatini or of a part differentiated from it on the autopalatine and/or dermopalatine (see above), the plesiomorphic condition is to lack an extensor tentaculi. [1] Some fibers of the extensor tentaculi are blended with those of the adductor arcus palatini (e.g. Diogo, fig. 3-89). [2] Extensor tentaculi completely separated from adductor arcus palatini (e.g. Diogo et al., 2003: fig. 7-7, 7-8, 7.9).

139. Adductor arcus palatini and preopercle. [0] No significant attachment of adductor arcus palatini to preopercle (e.g. Fig. 22). [1] Significant part of adductor arcus palatini inserting to preopercle (see e.g. Howes, 1985).

140. Adductor arcus palatini and lateral surface of suspensorium. [0] No significant attachment of adductor arcus palatini to lateral surface of suspensorium (e.g. Fig. 24). [1] Significant part of adductor arcus palatini inserting on lateral surface of suspensorium (e.g. Fig. 23; Diogo, fig. 3-20).

141. Anterior and posterior sections of levator arcus palatini. [0] Not peculiarly differentiated into well-developed anterior and posterior sections (e.g. Fig. 22). [1] Peculiarly differentiated into well-developed anterior and posterior sections (e.g. Greenwood, 1977: fig. 11).

142. Dorsolateral and ventromesial sections of levator arcus palatini. [0] Not differentiated into a well-developed dorsolateral and a well-developed ventromesial sections (e.g. Fig. 22). [1] Differentiated into well-developed dorsolateral and well-developed ventromesial sections (e.g. Fig. 24; Kirchhoff, 1958: figs. 38, 39; Greenwood, 1968: fig. 34).

143. Levator arcus palatini and metapterygoid. [0] The plesiomorphic condition seems to be that in which the levator adductor arcus palatini inserts not only on the hyomandibula but also on the metapterygoid, as is the case for example in Amia, Lepisosteus and many teleosts examined (e.g. Fig. 24; Lauder, fig. 2A). [1] Does not insert on metapterygoid.

144. Origin of levator arcus palatini. [0] No attachment of significant part of levator arcus palatini to dorsal surface of cranial roof (e.g. Fig. 22). [1] Significant part of levator arcus palatini originated on dorsal surface of cranial roof (see e.g. Diogo).

145. Origin of dilatator operculi. [0] No attachment of significant part of dilatator operculi to dorsal surface of cranial roof (e.g. Fig. 22). [1] Significant part of dilatator operculi originated on dorsal surface of cranial roof (see e.g. Diogo).

146. Dilatator operculi [0] Not almost completely covered in lateral view by dorsal surface of preopercle. (e.g. Fig. 22). [1] Almost completely covered in lateral view by dorsal surface of preopercle. Such a peculiar configuration of the dilatator operculi is found in Denticeps (e.g. Fig. 9) and has led some authors such as Greenwood (1968) to incorrectly state that this muscle is absent in the members of this genus.

147. Differentiation of dilatator operculi. [0] Not differentiated into two well-developed, distinct divisions. [1] Differentiated into two well-developed, distinct divisions (see e.g. Diogo).

148. Levator operculi. [0] As stated by authors such as Schaeffer & Rosen and Lauder the plesiomorphic condition for actinopterygians is seemingly that found in most basal actinopterygians and in fishes such as Lepisosteus, in which there is no recognizable levator operculi. [1] A recognizable levator operculi is found in those fishes coded as CS-1 (e.g. Fig. 22; Lauder, fig. 3A).

149. Levator operculi and lateral surface of opercle. [0] No insertion of significant part of levator operculi on lateral surface of opercle. [1] Significant part of levator operculi inserted on lateral surface of opercle (e.g. Diogo: fig. 3-88).

150. Levator arcus branchialis V (ordered multistate character). [0] Not hypertrophied. [1] Hypertrophied. [2] Still more enlarged and voluminous than in CS-1 (e.g. Matthes, 1963: plate 9c).

151. Division of levator operculi. [0] Not peculiarly divided into an anterior, mesial bundle and a posterior, lateral bundle. [1] Peculiarly divided into an anterior, mesial bundle and a posterior, lateral bundle (e.g. De la Hoz & Chardon, fig. 13a; Aguilera, 1986: fig. 1).

152. Drumming muscle of swimbladder. [0] Not present. [1] Present (e.g. Ladich, 2001; Diogo, fig. 3-100).

153. Fibers of hypaxialis and/or epaxialis. [0] Not peculiarly covering great part of neurocranial floor. [1] Peculiarly covering great part of neurocranial floor (e.g. Fig. 24; Gunther & Deckert, 1959: figs. 11, 12; Gosline, 1969: fig. 8; Fink & Fink, 1986).

Splanchnocranium.

154. Maxilla and infraorbitals. [0] Not forming peculiar, long compound structure (e.g. Fig. 20). [1] Maxilla and infraorbitals fused, forming peculiar, long compound structure (e.g. Arratia & Schultze, 1991: fig. 19).

155. Maxillary teeth. [0] Present (e.g. Fig. 20). [1] Absent (e.g. Fig. 22; Greenwood, 1977: fig. 21). We agree with the interpretation of authors such as Belouze in that the upper jaw of Eurypharynx is at least partially constituted by the maxilla. The presence of a well-defined, proximal head of the toothed element constituting the upper jaw in the Eurypharynx specimens examined, the presence of a strong ligament between the distal surface of this element and the mandible, and its overall position and shape, combined with the developmental data of Orton (1963), strongly support this interpretation. Some members of the genus Alepocephalus have maxillary teeth (e.g. Gosline, 1969; Sanford) while others not (e.g. Fig. 24; see also Begle, 1991, 1992); this genus is thus coded as '?'. As stressed by authors such as Greenwood (1968), the Denticeps specimens dissected by us have numerous odontodes in various bones of the skull, and is effectively difficult to discern, in certain specimens, if the 'teeth' present in some of these bones are or not exclusively odontodes. Until more detailed, conclusive data is available, we prudently code this genus as '?'.

156. Peculiar, deep lateral fossa on distal margin of maxilla, in which attaches significant part of adductor mandibulae. [0] Not present (e.g. Fig. 22). [1] Present (e.g. Howes, 1985: figs. 18, 19)

157. Premaxillae. [0] Present as independent ossifications (e.g. Fig. 20). [1] Not present as independent ossifications (e.g. Belouze, fig. 2A).

158. Supramaxillae. [0] Present as independent ossifications (e.g. Fig. 20). [1] Not present as independent ossifications (e.g. Fig. 22; Patterson, 1973: fig. 5A). Some members of the genus Astronesthes have supramaxillary bones, while others not (see e.g. Weitzman 1967b); this genus is thud coded as '?'.

159. Maxillae. [0] Markedly ankylosed with neurocranium. [1] Not markedly ankylosed with neurocranium (e.g. Fig. 20; Patterson, 1973: fig. 5A).

160. Distinct, strong ligaments connecting the anterior surface/anterior cartilage of autopalatines and/or dermopalatines and maxilla and/or premaxillae. [0] Not present. [1] Present (e.g. Fig. 10; Vrba, 1968: fig. 3).

161.Well-developed 'rostral' cartilaginous or cartilaginous-like structures associated with posterior surface of well-developed premaxillary dorsomedial processes attached to/articulating with ethmoid region. [0] Not present. [1] Present. The presence/absence and homologies of these structures have been the subject of controversy in the literature. For example, authors such as Fink and Weitzman (1982) and Fink (1984) stated that some fishes of the orders Stomiiformes and Aulopiformes have 'rostral' premaxillary cartilages similar to those present in members of more derived neoteleostean groups. However, authors such as Hartel & Stiassny (1986) and Stiassny (1986, 1996) questioned such an interpretation. Stiassny (1996: p. 455) called the attention to the fact that some of the structures that are often considered to be 'rostral' cartilages are in fact "not composed of hyaline cartilage"; they are, instead, "of an essentially fibrous composition of minimal matrix secretion". We thus prefer to define CS-1 in a way that it includes all the cases in which we have found "well-developed rostral cartilaginous or cartilaginous-like structures associated with the posterior surface of well-developed premaxillary dorsomesial processes attached to/articulating with the ethmoid region". In fact, whether these structures are paired or not, or are completely cartilaginous or not, among all the specimens examined we have only seen such well-developed structures in those few specimens coded as CS-1 (see e.g. Rosen, 1985: figs. 40A,C, 41A,B,C).

162. Premaxillae. [0] Not peculiarly fused in a single median structure (e.g. Fig. 20). [1] Peculiarly fused in a single median structure (e.g. Taverne, 1972: fig. 6; 1978: fig. 36).

163. Premaxillae and neurocranium. [0] Marked ankylosis (but not complete fusion) between these structures. [1] No marked ankylosis (nor complete fusion) between these structures (e.g. Fig. 20; Vrba, figs. 2, 3; Patterson, 1973: fig. 5C).

164. Maxilla. [0] Not exhibiting peculiar, somewhat /\-shaped overall configuration (e.g. Fig. 20). [1] Exhibiting peculiar, somewhat /\-shaped overall configuration (e.g. D'Aubenton, 1961: fig. 8; Lenglet, figs. 12, 13)

165. Peculiar mesial interdigitations between premaxillae. [0] Not present (e.g. Fig. 20). [1] Present (e.g. Weitzman, 1962: figs. 2, 4).

166. Prominent, well-defined, roundish anterior process of maxilla for articulation with posterior/mesial surface of premaxilla. [0] Not present (e.g. Fig. 10). [1] Present (e.g. Fig. 22; Sulak, 1977: figs. 3A, 7A).

167. Primordial ligament. [0] As the primordial ligament of Amia (e.g. Lauder, fig. 3A) and many teleosts examined connects the maxilla to the dorsal surface of the coronoid process, we are tentatively coding such a configuration as CS-0 for the fishes included in this cladistic analysis. [1] In the specimens examined of the genera coded as CS-1 the primordial ligament connects the maxilla to the posterolateral surface of the mandible, somewhat near its articulation with the quadrate (e.g. Fig. 23; Vrba, fig. 3).

168. Strong, well-defined ligament between premaxilla and proximal surface of maxilla. [0] Not present. [1] Present (e.g. Diogo, fig. 3-55). Some specimens of the genus Albula do not seem to have such a ligament (e.g. those examined in the present work) while others seemingly do (e.g. those described by Greenwood, 1977); this genus is thus coded as '?'

169. Peculiar ventromesial, roughly circular process of each premaxilla to articulate with its counterpart. [0] Present. [1] Not present (e.g. Monod, fig. 11, 'bouton articulaire median interpremaxillaires').

170. Prominent, roughly triangular anterolateral processes of premaxillae. [0] Not present (e.g. Fig. 21). [1] Present (e.g. Greenwood & Rosen, fig. 24).

171. Premaxillae. [0] Not syndesmotically attached to proximal head of maxillae in adults (e.g. Fig. 21). [1] Syndesmotically attached to proximal head of maxillae in adults (e.g. Howes & Sanford 1987: fig. 2).

172. Peculiar lateral excavation of upper and lower jaws, in which are anchored numerous outer, epithelially implanted comb-teeth. [0] Not present (e.g. Fig. 21). [1] Present (e.g. Howes & Sanford, fig. 2).

173. Large, peculiar tooth-bearing interpremaxillary pad between premaxillae in adults. [0] Not present (e.g. Fig. 21). [1] Present (e.g. Howes & Sanford, fig. 2).

174. Premaxillary teeth. [0] Present (e.g. Fig. 21). [1] Absent (e.g. Fig. 22). Contrary to Eurypharynx, which is coded here as '?' because it is difficult to discern if the premaxillae are present (e.g. fused with mesethmoid and/or prevomer) or not (in this case, this character would be inapplicable), there is a good amount of evidence suggesting that in the members of Anguilla and Conger the premaxillae are fused to the mesethmoid and prevomer and that these fishes do have premaxillary teeth (see e.g. Belouze, fig. 2D). 175. Long, strong ligament between premaxilla and anteromesial surface of mandible. [0] Not present. [1] Present (e.g. Thys van den Audenaerde, fig. 18).

176. 'Rictal cartilages' between upper jaws and between lower jaws. [0] Not present. [1] Present (e.g. Thys van den Audenaerde, fig. 18).

177. 'Gingival teeth' on upper jaw. [0] Not present. [1] As explained by authors such as Grande & Poyato-Ariza (p. 210) the 'gingival teeth' reported by Monod (1993) in Gonorynchus are non-osseous "fringes on the soft issue of the premaxillae".

178. Toothed dermopalatine. [0] Present (e.g. Fig. 21), occasionally fused to other structures (e.g. ectopterygoids: see below). [1] Absent (e.g. Figs. 22, 35; Lauder & Liem, fig. 38B; Arratia, 1992: fig. 8B). We agree with Johnson & Patterson in that in members of the genera Retropinna and Stokellia the toothed structure situated ventrally and posteroventrally to the autopalatine is very likely the result of the fusion of the dermopalatine with the ectopterygoid; these genera are thus coded here as CS-0, and as CS-1 in the character below.

179. Compound, dermopalatine-ectopterygoid toothed structure. [0] Not present (e.g. Fig. 21). [1] Present (e.g. Johnson & Patterson, fig. 4A). Authors such as Ridewood (1905a), Kershaw (1970), Taverne (1978), Hilton and Moritz & Britz (2005) stated that Xenomystus and Pantodon exhibit a compound dermopalatine-ectopterygoid toothed structure. The observations of the present work support such a statement. However Arratia & Schultze,'s figure 20A seems to suggest that in certain Pantodon specimens these two bones are not fused; this latter genus is thus coded as '?'. 180. Autopalatine. [0] Well ossified (e.g. Fig. 35). [1] Very poorly ossified or completely unossified (e.g. Fig. 21; De la Hoz & Chardon, 1984: fig. 2).

181. Articulation, either direct or indirect, between autopalatine/ dermopalatine and maxilla. [0] Not present. [1] Present. (e.g. Fig. 10; Vrba, 1968: fig. 3). It should be noted that, despite being poorly developed, there is a maxillary articulatory facet for the autopalatine/ dermopalatine in the specimens examined of the osteoglossomorph genera Hiodon, Pantodon and Xenomystus (it was however not possible to discern if this is also the case, or not, in the Mormyrus specimens dissected).

182. Peculiar, large, deep anterodorsal concavity of entopterygoid for articulation with ventral surface of lateral ethmoid. [0] Not present (e.g. Fig. 21). [1] Present (e.g. Vandewalle, 1975: fig. 51). 183. Pars autopalatina. [0] Not peculiarly separated from pars pterygoquadrata (e.g. Fig. 35). [1] Peculiarly separated from pars pterygoquadrata (e.g. Arratia, 1992: figs. 16, 17; Diogo et al., 2001b: fig. 1).

184. Large anteroventral expansion of laminar bone of autopalatine. [0] Not present (e.g. Fig. 35). [1] Present (e.g. Monod, fig. 30; Pasleau, 1974: fig. 23).

185. Anterior portion and/or anterior cartilage of autopalatine. [0] Not forming peculiar 'broad hook' that covers a great portion of the proximal portion of maxilla in lateral view (e.g. Fig. 24). [1] As noted by authors such as Sanford, although in a few fishes coded as CS-0 the anterior portion and/or anterior cartilage of the autopalatine may be significantly elongated anteroposteriorly and/or occasionally form a 'small hook', such a peculiar 'broad wook' covering a great portion of the proximal portion of the maxilla in lateral view is only found, within the specimens examined, in those fishes coded as CS-2 (e.g. Stiassny, 1986: fig. 5A; Sanford, fig. 32).

186. Well-developed, peculiar 'processus dorsomedialis' of autopalatine. [0] Not present (e.g. Fig. 35). [1] Present (e.g. Fig. 10; Fink & Fink, 1981: figs. 3B, 9).

187. Semimovable articulation between entopterygoid and posterior portion and/or posterior cartilage of autopalatine. [0] Not present (e.g. Fig. 35). [1] As explained by authors such as Fink & Fink (1981, 1996), although some fishes of CS-0 could occasionally exhibit a somewhat similar articulation between the anterior pterygoid bones and the autopalatine (e.g. some gonorynchiforms), only in those fishes of CS-1 such an articulation is made between a characteristic, concave facet of the entopterygoid and the posterior portion and/or posterior cartilage of the autopalatine.

188. Peculiar dorsoventral enlargement of posterior portion of autopalatine. [0] Not present (e.g. Fig. 35). [1] Present (e.g. Fig. 24; Gosline, 1969: figs. 4A, 5).

189. Strong, long ligament connecting anterior surface of pterygoid bones to maxilla. [0] Not present (e.g. Fig. 24). [1] Present (e.g. Diogo, fig. 3-39C).

190. Ossification of ligament connecting anterior margin of suspensorium to ethmoid region. [0] No ossification (e.g. Fig. 24). [1] Partial or complete ossification (e.g. Chapmam, 1942: fig. 3; De la Hoz & Chardon, 1984: fig. 8; Diogo, fig. 3-73).

191. Large metapterygoid-quadrate fenestra. [0] Not present (e.g. Fig. 35). [1] Present (e.g. Fink & Fink, 1981: figs. 9, 10).

192. Peculiar configuration of anterior portion of suspensorium, the main bodies of the entopterygoid and ectopterygoid being widely separated by the quadrate, with almost no contact, or no contact at all, between the entopterygoid and ectopterygoid. [0] Absence of such peculiar configuration (e.g. Fig. 35). [1] Presence of such peculiar configuration (e.g. Lekander, 1949: fig. 67).

193. Teeth on pterygoid bones. [0] Present. [1] Absent (e.g. Fig. 35). Some Alepocephalus specimens dissected by us have teeth on the pterygoid region while others not. Also, the Distichodus specimens examined by us have no teeth on this region, but Buckup stated that in the specimens examined by him there were teeth on both the entopterygoid and the ectopterygoid. These two genera are thus coded as '?'.

194. Peculiar, stout posteromesial process of ectopterygoid for articulation with ethmoid region. [0] Not present (e.g. Fig. 21). [1] Present (e.g. Thys van den Audenaerde, fig. 17).

195. Prominent dorsal process of ectopterygoid abutting in infraorbitals. [0] Not present (e.g. Fig. 21). [1] Present (e.g. Ridewood, 1904b; Forey et al., 1996).

196. Peculiar, prominent hyomandibular lateral spur at or below the level of the opercular process, projecting caudally to contact the preopercle. [0] Not present (e.g. Fig. 35). [1] Present (e.g. Johnson & Patterson, 1996: fig. 4B). In Johnson & Patterson's (p. 326) appendix 1, these authors coded this character as an unordered multistate character: for example, Plecoglossus was coded as CS-1 ("short vertical crest fitting against preopercular"), Osmerus as CS2 ("triangular spur"), and Galaxias as CS-3 ("obliquely oriented spurlike crest"). We usually agree with Johnson & Patterson's criticism of Begle's (1992) coding. However, in this case we prefer to follow Begle's (1992) coding for this character, because we do not think that the 'peculiar lateral hyomandibular spur at or below the level of the opercular process projecting caudally to contact the preopercle' found in the fishes of CS-1 being e.g. little bit more (as e.g. in Plecoglossus) or a little bit less (as e.g. in Galaxias) vertical is enough to exclude, a priori, the possibility that this feature constitutes a primary homology between these fishes.

197. Prominent, long, thin and posteroventrally directed posterior process of hyomandibula. [0] Not present (e.g. Fig. 35). [1] Present (e.g. Monod, figs. 36, 37).

198. Prominent anteroventromesial process of quadrate. [0] Not present (e.g. Fig. 35). [1] Present (e.g. Lenglet, fig. 12).

199. Prominent, thin, anterodorsally directed anteroventrolateral process of quadrate. [0] Not present (e.g. Fig. 35). [1] Present (e.g. Thys van den Audenaerde, fig. 17).

200. Metapterygoid. [0] Present as independent ossification (e.g. Fig. 35). [1] Absent as independent ossification (e.g. Smith, 1989a: fig. 6). In some specimens of Bathylagus the metapterygoid, although small, is present, while in others it is missing; this genus is thus coded as '?'.

201. Symplectic. [0] Present as independent ossification (e.g. Fig. 35). [1] Not present as independent ossification (e.g. Smith, 1989a: fig. 6).

202. Quadratojugal as independent ossification. [0] Present as independent ossification. [1] Not present as independent ossification (e.g. Fig. 35; Arratia & Schultze, fig. 23).

203. Two articulatory points between ventral portion of suspensorium and mandible. [0] The ventral portion of the suspensorium and the mandible do not articulate by means of two distinct articulatory points (e.g. Fig. 30). [1] The ventral portion of the suspensorium and the mandible articulate by means of two distinct articulatory points (e.g. Fig. 31; Patterson, 1973: fig. 6B).

204. Toothed 'dermometapterygoids'. [0] Not present (e.g. Fig. 35). [1] Present (e.g. Arratia & Schultze, fig. 42B).

205. Broad, deep, roughly circular fossa on anteroventromesial surface of quadrate. [0] Not present. [1] Present (e.g. Diogo et al., 1999: fig. 4).

206. Two articulatory heads of hyomandibula for neurocranium. [0] In Lepisosteus and Amia, as well as in most teleosts examined by us, the hyomandibula exhibits a single, continuous dorsal articulatory head for the neurocranium (e.g. Arratia & Schultze, fig. 23) (note: although in some taxa coded as CS-0, e.g. Megalops, it might seem that there two hyomandibular cartilaginous heads for the neurocranium, these cartilaginous structures are in fact continuous). [1] Two articulatory heads for neurocranium (e.g. Arratia & Schultze, fig. 23).

207. Prominent posterodorsal projection of hyomandibula (or hyomandibulo-metapterygoid) firmly attached to neurocranium by strong, short connective tissue. [0] Not present. [1] Present (e.g. Diogo).

208. Subopercle. [0] Present as independent ossification (e.g. Fig. 35). [1] Absent as independent ossification (e.g. Taverne, 1978: figs. 41, 104).

209. Subopercle and hyoid arch. [0] Subopercle not articulating directly with hyoid arch by a peculiar, prominent anterior spine. [1] Subopercle articulating directly with hyoid arch by a peculiar, prominent anterior spine (e.g. Ridewood, 1904a: fig. 135C).

210. Opercle. [0] Present as independent ossification (e.g. Fig. 35). [1] Not present as independent ossification (e.g. Tchernavin, 1947a: text-figs. 2, 3).

211. Peculiar, dorsoventrally elongated mesial crest of opercle for articulation with subopercle. [0] Not present. [1] Present (e.g. Monod, fig. 16).

212. Large fenestra on anterodorsal surface of opercle. [0] Not present (e.g. Fig. 35). [1] Present (e.g. Vari, fig. 19B).

213. Preopercle. [0] Present as independent ossification (e.g. Fig. 35). [1] Not present as independent ossification (e.g. Tchernavin, 1947a: text-figs. 2, 3).

214. Opercle. [0] Not with characteristic triangular shape (e.g. Fig. 35). [1] With characteristic triangular shape (e.g. Fink & Fink, 1981: figs. 11, 12).

215. Several peculiar thin spiny structures on posterior and/or posteroventral portions of opercle. [0] Not present (e.g. Fig. 35). [1] Present (e.g. Fig. 24). Some specimens of the genera Elops, Megalops, Galaxias and Esox have such peculiar spiny structures (e.g. Fig. 20; Taverne, 1974), while others not (e.g. Ridewood, 1904; Vrba; Winterbottom, 1974); these genera are thus coded as '?'.

216. Prominent, thin, dorsally/anterodorsally oriented anterodorsal projection of opercle. [0] Not present (e.g. Fig. 20). [1] Present (e.g. Fig. 35; De la Hoz & Chardon, 1984: fig. 13C; Diogo, fig. 389).

217. Prominent, broad posteroventral spine-like process of preopercle. [0] Not present (e.g. Fig. 35). [1] Present (e.g. Fig. 9; Greenwood, 1968: fig. 7).

218. Preopercles. [0] Not markedly expanded ventrally (e.g. Fig. 4). [1] Markedly expanded ventrally, one preopercle overlapping the other along ventral midline (e.g. This van den Audenaerde, fig. 14).

219. Large anterolateral articulatory facet of interopercle for articulation with quadrate. [0] Not present (e.g. Fig. 35). [1] Present.

220. Well-defined, long, strong ligament running from anterodorsal surface of interopercle to posterodorsal surface of preopercle. [0] Not present. [1] Present (e.g. Diogo).

221. Interopercle. [0] Not a markedly thin and long structure (e.g. Fig. 35). [1] A markedly thin and long structure (e.g. Thys van den Audenaerde, fig. 20).

222. Opercle/subopercle. [0] Not widely separated from interopercle (e.g. Fig. 35). [1] Widely separated from interopercle (e.g. Gosline, 1969: fig. 3A).

223. Interhyal. [0] As noted by authors such as Patterson (1977, 1982) and Jollie (1980, 1984ab, 1986), in Amia and Lepisosteus, as well as in other basal Actinopteri such as Acipenser and Polyodon and in many teleosts, the interhyal (sensu these authors) is not ossified (see e.g. Fig. 4). [1] Presence of ossified interhyal (e.g. Fig. 5; Arratia & Schultze, fig. 2D). Some members of the genera Chanos, Gonorynchus, Phractolaemus, Nematogenys and Catostomus do not have an ossified interhyal (e.g. Fig. 6), while others have (see e.g. Weisel, 1960; Arratia, 1992; Grande & Poyato-Ariza; Diogo). These genera are thus coded as '?'.

224. Length of interhyal. [0] Interhyal (ossified or not) not markedly elongated dorsoventrally (e.g. Fig. 4). [1] Interhyal (ossified or not) markedly elongated dorsoventrally, such a peculiar elongation being seemingly related to a rather peculiar mechanism of mouth opening/ closure (e.g. Tchernavin, 1953; Gunther & Deckert, figs. 26, 33).

225. Interhyal, hyoid arch and suspensorium. [0] Interhyal (ossified or not) not connected by ligaments to both hyoid arch and suspensorium (e.g. Fig. 5). [1] Interhyal (ossified or not) connected by ligaments to both hyoid arch and suspensorium and thus not articulating directly with these structures. Arratia (1992) described this derived feature for catfishes. However the gymnotiform of the genera Sternopygus, Gymnotus and Brachyhypopomus examined by us and by authors such as De la Hoz & Chardon (e.g. fig. 8) also have ligaments between the interhyal and both the hyoid arch and the suspensorium.

226. Shape of interhyal. [0] Interhyal (ossified or not) not with peculiar, somewhat dumbbell shape (e.g. Fig. 4). [1] Interhyal (ossified or not) with peculiar, somewhat dumbbell shape. The original definition of this character by Begle was 'interhyal short, dumbbell-shaped'. However, as stressed by Johnson & Patterson, in specimens such as those of the genus Bathylagus the interhyal is not significantly shorter than in certain other fishes coded as CS-0. Therefore, we prefer to use the less restricting terms 'interhyal with peculiar, somewhat dumbbell shape': such a description does effectively apply to the condition found all in the specimens examined of those taxa coded as CS-1.

227. Distinct mandibulo-hyoid ligament. Present (e.g. Fig. 24; Lauder, fig. 3A, 18). [1] Not present.

228. Mandibulo-hyoid and mandibulo-interopercular ligaments. [0] Within those fishes having mandibulo-hyoid and mandibulo-interopercular ligaments, plesiomorphically these ligaments are well-separated from each other (e.g. Vrba, fig. 5; Lauder, fig. 3A). [1] In the specimens examined of the taxa coded as CS-1 one of these three conditions, or a combination of them, occurs: the two ligaments are deeply mixed anteriorly, giving the appearance that there is a single ligament attaching anteriorly on the mandible (e.g. Fig. 33); a significant portion of the mandibulo-interopercular ligament attaches also on the posterior ceratohyal (e.g. Figs. 5, 24); a significant portion of the mandibulo-hyoid ligament attaches also on the interopercle. It should be noted that some authors referred to a 'shift in insertion of the mandibulo-hyoid ligament to the interopercle' as a synapomorphy of the Eurypterygii (e.g. Lauder & Liem; Johnson, 1992) or of the Neoteleostei (e.g. Rosen, 1985). Thus, according to Rosen (1985) fishes such as the basal Neoteleostei stomiiforms, as well as basal Eurypterygii such as the aulopiforms (see Fig. 1), seemingly exhibit such a feature, while according to Lauder & Liem and Johnson (1992) the stomiiforms lack this feature. However, Stiassny (1996) and Sato & Nakabo stressed that such a feature is not present in stomiiforms nor in numerous aulopiforms. The observations of the present study support the statement of these latter authors. The stomiiform and aulopiform fishes examined by us exhibit a condition similar to that found in many non-neoteleostean taxa of CS-1: the mandibulo-hyoid and mandibulo-interopercular ligaments are both present, but are deeply blended anteriorly, giving the appearance that there is a single ligament attaching anteriorly on the mandible (e.g. Fig. 33).

229. Well-developed, posterodorsally pointed dorsal process of posterior ceratohyal. [0] Not present (e.g. Fig. 5). [1] Present (e.g. Thys van den Audenaerde, fig. 17).

230. Well-developed ventrolateral laminar expansion of anterior ceratohyal. [0] Not present (e.g. Figs. 5, 6). [1] Present (e.g. Diogo).

231. Prominent, broad anteroventral lamina of anterior ceratohyal. [0] Not present (e.g. Figs. 5, 6). [1] Present (e.g. Diogo).

232. Peculiar tooth plates associated with anterior and/or posterior ceratohyals. [0] Not present (e.g. Figs. 5, 6). [1] Present (e.g. Arratia & Schultze, fig. 2C,D). Apart Elops and Amia, the Megalops specimens examined by us also exhibit such tooth plates (Megalops was not included in the list of fishes analyzed in Arratia & Schultze's work).

233.Peculiar articulation between prominent anteromesial process of anterior ceratohyal and broad, deep, circular concavity formed by the lateral margins of both urohyal and basihyal. [0] Not present (e.g. Fig. 6). [1] Present (e.g. Taverne, 1972: figs. 9, 10).

234. Broad, deep, roughly circular concavity in each of the anterodorsolateral margins of the urohyal for lodging the anteroventral surface of anterior ceratohyals. [0] Not present (e.g. Fig. 6). [1] Present (e.g. Belouze, fig. 37).

235. Branchiostegal rays. [0] Present (e.g. Fig. 6). [1] Absent (e.g. Tchernavin, 1947a: text-figs. 2, 3; McAllister, 1968).

236. Ossified urohyal/parurohyal. [0] Not present. [1] Present (e.g. Fig. 6; Arratia & Schultze, fig. 13).

237. Peculiar long and thin cartilages of branchiostegal rays for articulation with hyoid arch. [0] Not present (e.g. Fig. 6). [1] Present.

238. Branchiostegal photophores. [0] Absence of several peculiar branchiostegal photophores. [1] Presence several peculiar branchiostegal photophores (e.g. Fink, 1985; Harold & Weitzman, 1996).

239. Ossification of ventral and dorsal hypohyals. [0] At least one ossified hypohyal present (e.g. Fig. 6). [1] Absence of ossified hypohyals (e.g. Smith, 1989a: fig. 7; Baldwin & Johnson, 1996: fig. 12B). As explained by authors such as Taverne (1972) and Hilton, Mormyrus apparently has hypohyals, although they are very small.

240. Branchiostegal rays and hypohyals. [0] Not articulating with hypohyals (e.g. Fig. 6). [1] Some branchiostegal rays articulating with hypohyals (e.g. Weitzman, 1967b: fig. 11).

241. Ossified gular plate. [0] Present (e.g. Fig. 20). [1] Absent (e.g. Fig. 4; Gosline 1963: fig. 26B; Jessen 1968: fig. 1). Forey et al. (1996) coded Albula as having an ossified gular plate, but the specimens of this genus examined by us do not seem to have such an ossified gular plate; this genus is thus prudently coded as '?'.

242. Mandibular teeth. [0] Present (e.g. Fig. 21). [1] Absent (e.g. Fig. 10; Vandewalle figs. 1, 2). In some members of the genus Coregonus there are small mandibular teeth while in others such teeth are apparently missing (e.g. Shaposhnikova; Nelson, 1973; Sanford); this genus is thus coded as '?'. Mandibular teeth were drawn in Gayet's (1981) figure 4 of [dagger] Lusitanichthys characiformis but were not drawn in Gayet's (1985) figure 18 of the same species. Cavin described a new species of [dagger] Lusitanichthys, [dagger] L. africanus, which, according to the latter author, does not have mandibular teeth, suggesting that these teeth might effectively be missing in [dagger] L. characiformis. However, until more conclusive information is available, we prefer to prudently code [dagger] L. characiformis as '?'.

243. Peculiar articulation between dentary bone and posterior portion of mandible. [0] Not present (e.g. Fig. 34). [1] Present (e.g. Vari, fig. 42).

244. Coronoid bones. [0] Present as independent ossifications (e.g. Fig. 31). [1] Absent as independent ossifications (e.g. Fig. 32; Patterson, 1973: fig. C).

245. Prearticulars. [0] Present as independent ossifications (e.g. Fig. 31). [1] Absent as independent ossifications (e.g. Fig. 32; Patterson, 1973: fig. C).

246. Articulars. [0] Not mainly fused with angulars (and/or retroarticulars) (e.g. Fig. 30). [1] Mainly fused with angulars (and/ or retroarticulars) (e.g. Fig. 32; Nelson 1973: fig. 6H). According to the descriptions of authors such as Nelson (1973), Taverne (1974) and Hilton the articulars of the members of the genera Megalops and Elops are not fused with the angulo-retroarticulars (see e.g. Fig. 20). However, Arratia (1999: pp. 275, 289) coded Megalops as having articulars "fused with angular and retroarticular bones". Megalops is thus coded here as '?'. Concerning Elops, Arratia (1999: pp. 275, 289) coded this genus as having articulars "partially fused with anguloretroarticulars late in ontogeny". Elops is coded here under CS-0, since, even if in some adult specimens of this genus the articulars are occasionally 'partially' fused with the angulo-retroarticulars, as stated Arratia (1999), they are not 'mainly' fused with these latter structures, as described for CS-1.

247. Retroarticulars. [0] Present as independent ossifications (e.g. Fig. 31). [1] Absent as independent ossifications (e.g. Fig. 20; Nelson, 1973: fig. 3K). According to authors such as Nelson (1973) and Hilton in Hiodon the retroarticulars are fused with the angulars, but this is seemingly not always the case, since the Hiodon specimens illustrated by Arratia (1997: fig. 85B) and Taverne (1977b: fig. 15) appear to have independent retroarticulars (see e.g. Fig. 21). This genus is thus coded as '?'.

248. Retroarticular and quadrato-mandibular joint. [0] Retroarticular not excluded from quadrato-mandibular joint (e.g. Fig. 30). [1] Retroarticular excluded from quadrato-mandibular joint (e.g. Nelson, 1973: fig. 6D) [State 1]. Arratia (1992) stated that adult gymnotiforms display a configuration such as CS-0. This is effectively the case for the Gymnotus and Brachyhypopomus specimens that we have dissected. However, this is not the case for the Sternopygus specimens examined in the present work and in works such as De la Hoz & Chardon (figs. 2, 8). Nelson (1973: 340) stated that in the notopterid osteoglossomorphs (which include Xenomystus), as well as in the osteoglossomorph Pantodon, the retroarticular is not included in the quadrato-mandibular joint. But Hilton questioned this statement because, according to him, the retroarticular of Pantodon is included in quadrato-mandibular joint. Concerning Xenomystus, Hilton stated that in members of this genus the retroarticular lies close to the quadrato-mandibular joint. In the Xenomystus and Pantodon specimens analyzed by us the retroarticular is included in the joint. So, there is seemingly a variation of this character within the members of these two latter genera, and they are thus coded as '?' (it should be noted that the fishes in which the retroarticular is not present as an independent ossification--see above--were coded as 'Inapplicable').

249. Lower jaw. [0] Not highly modified, i.e. dentary bone not roughly perpendicular to remaining of mandible (e.g. Fig. 32). [1] Highly modified, with dentary bone roughly perpendicular to remaining of mandible (e.g. Howes, figs. 6, 19C).

250. Various peculiar, prominent lateral, mesial and dorsal processes of mandible. [0] Not present (e.g. Fig. 32). [1] Present (e.g. Howes, fig. 13).

251. Right and left halves of lower jaw. [0] Firmly attached to each other at midline (e.g. Fig. 4). [1] Not firmly attached to each other at midline (e.g. Howes & Sanford).

252. Dorsal tip of coronoid process. [0] Not markedly curved mesially (e.g. Fig. 34). [1] Markedly curved mesially (e.g. Fig. 30; Diogo, fig. 3-91).

253. Prominent, posterodorsally pointed anterolateral spine of dentary bone in adults. [0] Not present (e.g. Fig. 9). [1] Present (e.g. Fig. 10).

254. Prominent, stout, dorsoventrally elongated posterolateral and anterodorsally pointed process of angulo-articular. [0] Not present (e.g. Fig. 9). [1] Present (e.g. De la Hoz & Chardon, fig. 2).

255. 'Interdentary'. [0] Not present. [1] The Plecoglossus specimens examined by us exhibit a 'mandibular postsymphysial element with ossified tips', which was named 'interdentary' by Howes & Sanford (1987b: fig. 3). Howes & Sanford and Johnson & Patterson found, in a specimen of Osmerus mordax, a spherical, partially ossified body lying posterior to the mandibular symphysis that, according to Howes & Sanford, appears to be somewhat similar to the 'interdentary' of Plecoglossus. Such a structure was not found in the Osmerus specimens examined by us; this genus is thus coded as '?'.

256. Characteristic 'ascending portion' of Meckel's cartilage. [0] Not present (e.g. Fig. 32). [1] Present (e.g. Mo, 1991: fig. 37; Arratia, 1992; Diogo, fig. 3.36).

257. Ossified coronomeckelian bone. [0] Not present (e.g. Nelson 1973: fig. 1). [1] Present (e.g. Fig. 32; Nelson, 1973: fig. 2).

258. Dorsal margin of coronomeckelian bone. [0] Not situating significantly dorsal to upper margin of other mandibular bones (e.g. Fig. 32). [1] Situating significantly dorsal to upper margin of other mandibular bones (e.g. Howes, fig. 13).

259. Prominent, roughly circular articulatory surface on dorsomesial margin of angulo-articular for articulation with quadrate and/or ectopterygoid. [0] Not present (e.g. Fig. 32). [1] Present.

Miscellaneous

260. 'Saclike bioluminescent organ' at posterior margin of pectoral fin. [0] Not present. [1] As noted by authors such as Parr (1951, 1960), Matsui & Rosenblatt (1987), Begle and Johnson & Patterson, the fishes coded as CS-1 exhibit a 'saclike bioluminescent organ' at the posterior margin of the pectoral fin.

261. 'Tongue-bite mechanism' with dorsal teeth on parasphenoid and occasionally on other bones. [0] Not present. [1] Present (e.g. Sanford & Lauder, fig. 2). As explained by authors such as Sanford, a few other teleosts, e.g. some salmoniforms, may exhibit a mechanism roughly similar to that of fishes of CS-1. But there are significant differences between the two mechanism and only the mechanism of fishes of CS-1 involves the presence of dorsal teeth on the parasphenoid.

262. Peculiar muscle retractor dorsalis. [0] Not present. [1] Present (e.g. Winterbottom, 1971: fig. 19, 23). As noted by authors such as Johnson (1992: p. 11), "although musculature between the free vertebral column and dorsal gill arch elements occurs in a few lower teleosts (e.g. Pantodon, some muraenid, some cyprinids, siluriforms)", this muscle has been interpreted by most authors as "nonhomologous with the retractor dorsalis of neoteleosts". According to this author, the peculiar neoteleostean muscle retractor dorsalis is associated with an also peculiar "modification of the dorsal gill arch muscles" of fishes such as stomiiforms and aulopiforms, namely the "insertion of the third internal levator on the fifth upper pharyngeal toothplate". As also stressed by Johnson (p. 11), "Fink (1984) noted the presence of a 'retractor dorsalis' in Lepidogalaxias", a genus not included in the present study that is nowadays usually included in the Galaxioidea. According to Johnson the configuration found in the members of the genus Lepidogalaxias is unlike that found in fishes such as stomiiforms and aulopiforms, since the former "retain insertion of the third internal levator on the fourth pharyngobranchial cartilage", this "lack of the associated neoteleostean modification" being "at least consistent with an independent origin of the 'retractor dorsalis' of Lepidogalaxias". 263. Pair of well-developed 'nasal tubes' in anterolateral surfaces of head. [0] Not present. [1] Present (e.g. Thys van den Audenaerde, 1961: fig. 1).

264. Peculiar 'accessory cartilage of the fifth ceratobranchial'. [0] Not present. [1] As explained by authors such as Nelson (1967, 1969, 1970), the fishes coded as CS-1 have a peculiar 'accessory cartilage of the fifth ceratobranchial', which forms part of the 'crumeral organ' of these fishes (e.g. Greenwood & Rosen, figs. 1A, 2A, 4A, 6B).

265. Characteristic multicuspid teeth. [0] Not present (e.g. Fig. 21). [1] Present (e.g. Fig. 23; Weitzman, 1962: fig. 10, 1964) [State 1].

266. Characteristic maxillary barbels moved by palatine-maxillary system. [0] Not present. [1] Present (e.g. Alexander, 1965: fig. 8; Gosline, 1975: fig. 1; Diogo et al., 2000, 2003; Diogo & Chardon)

267. Paired 'mandibular barbels' associated with peculiar basal cartilages. [0] Not present. [1] Present (e.g. Diogo et al., 2003: fig. 7.3A,B).

268. 'Leptocephalus larva'. [0] Not present. [1] Present (e.g. Inoue et al., 2004: figs. 1, 5). Authors such as Gosline (1973) have stated that a 'leptocephalus larva' is also found in fishes other than those usually included in the clade Elopomorpha (see Fig. 1), and that the character state described as CS-1 might be a plesiomorphic feature for teleosts. However, authors such as Forey (1973b) and Forey et al. (1996) pointed out that the 'leptocephalus larva' of fishes included in the 'Elopomorpha' exhibits, in fact, several peculiar, derived features in relation to the larvae of other lower teleosts (see e.g. Forey, 1973b, for more details on these features). In the present work the use of the term 'leptocephalus larva' follows the more restrictive definition given by authors such as Forey (1973b) and Forey et al. (1996).

269. 'Symphysial barbel'. [0] Not present. [1] Present (e.g. Howes, fig. 7).

270. 'Luminous chin barbel'. [0] Not present. [1] Present (e.g. Weitzman, 1967b: fig. 1).

271. One or more 'abdominal scutes', each of a single element which crosses the ventral midline of the fish. [0] Not present. [1] As explained by authors such as Grande (1985) the fishes coded as CS-1 present such peculiar 'abdominal scutes'. As noted by Hilton (p. 68), some osteoglossomorphs, e.g. Xenomystus, have somewhat similar 'abdominal scutes', but "the abdominal scutes found in clupeoids are formed as single median elements, whereas those of notopterids are paired, and therefore do not pass the test of similarity in the establishment of (primary) homology". Xenomystus is thus coded as CS-0.

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*,**,*** Rui Diogo; ** Ignacio Doadrio & *** Pierre Vandewalle

* Department of Anthropology, George Washington University, Washington DC, USA.

** Museo Nacional de Ciencias Naturales, Madrid, Spain.

*** Laboratory of Functional and Evolutionary Morphology, University of Liege, Liege, Belgium.

Correspondence to:

Dr. Rui Diogo

Department of Anthropology,

The George Washington University,

2110 G St. NW, Washington,

DC 20052, USA

Phone: 202-248-1440

Email: ruidiogo@gwmail.gwu.edu

Received: 29-03-2008

Accepted: 07-07-2008
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Title Annotation:Part 2
Author:Diogo, Rui; Doadrio, Ignacio; Vandewalle, Pierre
Publication:International Journal of Morphology
Date:Sep 1, 2008
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