Mesozoic Fishes 5 – Global Diversity and Evolution, G. Arratia, H.-P. Schultze & M. V. H. Wilson (eds.): pp. 73-86, 6 figs., 1 tab. © 2013 by Verlag Dr. Friedrich Pfeil, München, Germany – ISBN 978-3-89937-159-8 Amiiforms from the Iberian Peninsula: historic review and research prospects Hugo MARTÍN-ABAD and Francisco José POYATO-ARIZA Abstract The fossil record of amiiform fishes from the Iberian Peninsula is known from 12 different sites. It extends along the three periods of the Mesozoic era, being much more abundant during the Early Cretaceous. The majority of these sites has yielded isolated remains, mostly teeth; complete and articulated specimens are known from three Konservat-Lagerstätten only: Montral-Alcover, El Montsec, and Las Hoyas. Generic and specific level assessments are possible in these three localities only. The historical taxonomical problems of the cited amiiform taxa are commented. Special mention deserves the case of the genus Urocles (= Megalurus), traditionally cited from El Montsec and Las Hoyas. This genus is invalid since 1998, but its taxonomic history goes back to the 1830’s. The amiiform fishes from Las Hoyas have traditionally been assigned to the same taxa as those coming from El Montsec, but they have not been studied in detail yet, so their taxonomical assessment is in need of confirma- tion. These specimens, characterized by a great quality of preservation and an abundant record, may provide significant information concerning the ontogenetic development of the fishes of this order. Introduction The amiiform fishes: Amiiformes HAY (1929) is an order of neopterygian fishes with an only extant species of the type genus, Amia calva; their peak of diversity occurred during the Mesozoic. Since there are very few living non-teleosts neotperygians, this order is considered to be as a very relevant group from a systematic and paleobiogeographic point of view, especially when dealing with its phylogenetic relationships with the Teleostei. In the last thirty years diverse hypotheses about the relationship be- tween amiiforms and teleosts have been offered, in an attempt to solve the debate on whether they are sister-groups or not. GRANDE & BEMIS (1998) provided a cladistic analysis resolving a clade containing both teleosts and amiiforms as sister-groups; this clade has traditionally been called Halecostomi. They redescribed the order Amiiformes, which they diagnosed by the following three evolutionary novelties: phylogenetic reduction in the number of ossified ural neural arches to two or less (character secondarily lost within Caturidae); phylogenetic loss of the opisthotic bone (character apparently convergent with lepisosteids and more derived teleosts); and phylogenetic loss of the pterotic bone. GRANDE & BEMIS (1998) also divided the amiiforms into two superfamilies, Amioidea and Caturoidea, each of which is in turn divided into two families, respectively: Amiidae and Sinamiidae, and Caturidae and Liodesmidae. Although it is considered as a principally Mesozoic order, Amiiformes present a relatively long chrono- stratigraphical record, which extends at least from the Jurassic, or even the Triassic, if the identifications made by BELTAN (1972, 1984) and LIU et al. (2002) are correct, to the Recent Amia calva, that inhabits freshwaters systems of the eastern North America. The amiiform fossil record also presents a wide-ranged geographical distribution that could be considered almost cosmopolite; however, the vast majority of the known fossil material has been discovered in sites from the North Hemisphere, probably as a result of bias in paleontological field work. This is partly evidenced by their distribution reaching very high latitudes (ESTES & HUTCHINSON 1980, LEHMAN 1951). 73 MESOZOIC The Spanish record: A number of nominal PICKS genera have been described within the Amii- PERIOD EPOCH AGE (Ma) 12. Cacém-Alcântara 65.5 formes, mainly in the family Amiidae. The MAASTRICHTIAN 70.6 present paper is included in a revision project CAMPANIAN 11. Las Hoyas on the amiiforms from Las Hoyas fossil site LATE SANTONIAN 83.5 85.8 CONIACIAN (Cuenca, Spain), currently in progress. In this 89.3 TURONIAN 93.5 12. Buenache de la Sierra sense, the main objective of the present paper CENOMANIAN 99.6 is to offer the most relevant references on ALBIAN 9. Alcaine amiiform fishes from the Iberian Peninsula, in 112 order to provide a biogeographical and chro- APTIAN EARLY 125 noestratigraphical framework for the amiiforms BARREMIAN 8. Galve CRETACEOUS 130 HAUTERIVIAN of Las Hoyas. Some of these references appear 136 VALANGINIAN 140 in the few compilations of the Spanish material BERRIASIAN 145.5 7. Vega de Pas I TITHONIAN published in the second half of the past century, 151 LATE KIMMERIDGIAN which are faunal lists mainly focused on the 156 OXFORDIAN 161 CALLOVIAN 6. El Montsec Cretaceous period (BATALLER 1960, POYATO- 165 BATHONIAN MIDDLE 168 BAJOCIAN ARIZA & WENZ 1990, POYATO-ARIZA et al. 172 AALENIAN 176 5. Cuenca de Cameros 1999). Those concerning only amiids that were TOARCIAN 183 published before 1998 were already included in EARLY PLIENSBACHIAN JURASSIC 190 SINEMURIAN GRANDE & BEMIS (1998); the posterior refer- 197 4. Tereñes & Lastres HETTANGIAN 201.6 ences to this family in the Iberian Peninsula RHAETIAN 204 are compiled in the present paper for the first 3. Guimarota time, together with all available references to NORIAN LATE the other families of this order. 228 2. Sot de Chera & Loriguilla Chronostratigraphicly, the Iberian record of CARNIAN 235 LADINIAN amiiform fishes extends from the Oxfordian- MIDDLE 241 TRIASSIC ANISIAN 245 1. Montral-Alcover Kimmeridgian (Late Jurassic) to the middle- OLENEKIAN EARLY 250 INDUAN 251.0 to-late Cenomanian (Upper Cretaceous), being Fig. 1. thus exclusively Mesozoic (Fig. 1). The first Chronostratigraphic distribution of the Iberian sites that appearance of the order could be traced back have yielded remains of amiiform fishes. The assessment of to the late Ladinian (Middle Triassic) accord- the material from Montral-Alcover to the order Amiiformes ing to BELTAN (1972, 1984) and VÍA-BOADA is doubtful. et al. (1977), but other authors (CARTANYÀ Table 1. Distribution of the taxonomic assessments of the amiiform fishes cited from Mesozoic localities of the Iberian Peninsula. Montral-Alcover Sot de Chera & Loriguilla Guimarota Tereñes & Lastres Cuenca de Cameros El Montsec Vega de Pas I Galve Alcaine Buenache de la Sierra Las Hoyas Cacém-Alcântara Amiiformes indet. %% %% Caturidae indet. %% Caturus sp. ? %%% Caturus tarraconensis % Amiidae indet. %% %% % Amiopsinae indet. % Amiopsis woodwardi %% Vidalamia catalunica %% 74 4 7 5 6 1 8 10 9 11 3 2 12 Cretaceous Jurassic Triassic Fig. 2. Mesozoic Iberian sites that have yielded remains of amiiform fishes. Numeration of localities corresponds to that from Figure 1. 1995, 1999) considered their assessment doubtful and in need of a revision. Geographically, the sites that have yielded remains of amiiform fishes are mainly concentrated in the northern and oriental regions of Spain and in the occidental coast of Portugal. To date, no insular outcrops have been reported from Spain or Portugal. Figure 2 shows the Iberian localities from Triassic, Jurassic, and Cretaceous beds. In general terms, the Iberian record could be divided into two sets. On one hand there are several sites whose remains appear to be more or less isolated and disarticulated, thereby creating problems with taxonomy. This set includes remains that have been reported as Actinopterygii indet., Neopterygii indet., Halecostomi indet., Halecomorphi indet., or Holostei indet., but they will not be considered in the present paper, since their assessment to the order Amiiformes would require a detailed revision that lies outside of the scope of the present work. On the other hand, there are a few Konservat-Lagerstätten which present an extensive and excellently-preserved fish record that usually allows a much more precise taxonomic assignment. Among these last ones, there are two localities that are especially relevant: El Montsec and Las Hoyas. In some cases, the material was originally described under names that have been changed in the course of history. In other cases, the remains do not really belong to the amiiform taxa they were initially assigned to or the corresponding taxa are not included in the Amiiformes anymore. Each particular case will be presented and discussed in detail below, following a systematic order (Table 1), with special reference to the genus Urocles (= Megalurus), of which the taxonomic history is quite complex. Vernacular suffixes and institutional abbreviations: The vernacular suffixes are used in this paper following GRANDE & BEMIS (1999). Institutional abbreviations used here are as follow: MCCM, Museo de las Ciencias de Castilla-La Mancha, Cuenca, Spain (LH, Las Hoyas Collection; MSE, El Montsec Collection); MMGB, Museo Municipal de Geología de Barcelona, Barcelona, Spain; MNHN, Muséum National d’Histoire Naturelle (Institut de Paléontologie), Paris, France. 75 The amiiform record from the Iberian Peninsula Taxa no longer assigned to the Amiiformes: Two different genera have been traditionally included within the Amiiformes, but subsequently were transferred to other orders. These are Eoeugnathus (i. e. BELTAN 1984, CARTANYÀ 1995) and Ophiopsis (i. e. CARTANYÀ 1999). Amiiformes indet.: In many localities the remains of amiiform fishes have been cited, of which the quality and/or integrity of preservation only allows an identification