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http://dx.doi.org/10.18268/BSGM2019v71n3a12 P. 831‒839 VOL. 71NO. 3 BOL. SOC.GEOL.MEX.2019 Coyoacán, CDMX,04510,México. Investigación S/N, CiudadUniversitaria, Autónoma deMéxico, Circuito dela Instituto de Geología, Universidad Nacional Jesús Alvarado-Ortega [email protected] Kleyton M.Cantalice Kleyton M. Mexico southeastern Chiapas, Ixtapa Formation(Miocene), the in Teleostei) The firstoccurrence of a freshwater percomorph (: Short note Manuscript May accepted: 11,2018. Corrected manuscript received: May 5,2018. Manuscript received: 9,2018. April Cantalice , Jesús Alvarado-Ortega apas, Mexico. water, Cenozoic, Ixtapa, Chi Keywords: Percomorph, Fresh through time. freshwater diversityfish inthisregion the ant complement to understand ent discovery constitutes an import Miocene–Pliocene. Hence, the pres Mexico the since endof the southern freshwaterprimary in percomorphs this fossilreveals the occurrence of mination at species level; however, imen impedes a taxonomical deter this spec imperfect preservation of the ural centrum 2.The fin lacks and itscaudal unsegmented spines andanalhavefins because its dorsal the Subdivision Percomorphaceae of from Ixtapaconsideredis amember proboscideans. fossil fish first This including horses, rhinoceros, and pods, aswell as terrestrial mammals charophytes, andaquatic gastro leaves,includes abundant angiosperm previouslyrecovered inthisformation Formation. fossilassemblage The the Ixtapa continental sediments of which belong to the Upper Miocene conditions,under fluvio-lacustrine ashfall sedimentsdeposited contains Mexico. Ixtapa site southeastern The near Tuxtla Gutiérrez City, Chiapas, located at theIxtapa municipality, lected in the Ixtapa fossiliferoussite, incomplete fossil fish specimencol In thispaper we and describe a single ABSTRACT Boletín de la Sociedad Geológica Mexicana Geológica Sociedad la de Boletín ------en elMunicipio del mismo nombre, cerca de tres del sitio fosilífero localizado de Ixtapa, fluvio-lacusfósil colectado en los depósitos Aquí se describe un único e incompleto pez RESUMEN pa, Chiapas, México. Dulceacuícola, Cenozoico, Ixta Palabras clave: Percomorfo, del tiempo. a través peces dulceacuícolas en esta región importante para comprender ladiversidad de descubrimiento constituye uncomplemento Mioceno-Plioceno. Por lo tanto, el presente dulceacuícolas en elsur de Méxicodesde el lapresenciadesvela depeces percomorfos nómica más precisa, sin embargo, este fósil taxo unadeterminación vación no permite el esqueletocaudal. La imperfecta preser además de la ausencia de centro ural 2en en las aletas dorsalno-segmentadas y anal, Percomorphaceae por la presencia de espinas derado como un miembro dela Subdivisión pez y esconsi fósil la localidad en Ixtapa proboscidios. deun Este es elprimer registro cómo, por ejemplo, caballos, rinocerontes y demamíferostambién registros terrestres algas carofitas, gasterópodos acuáticos y amplia diversidad de hojas de angiospermas, incluyerecuperada esta en formación una La asociación fósil a laFormación Ixtapa. cenizas del Mioceno superior pertenecientes el sureste de México. En este sitio se exponen la ciudad de Tuxtla Chiapas, Gutiérrez, en /2019 831 - - - - -

Freshwater percomorph fish of Ixtapa ABSTRACT INTRODUCTION / MATERIAL AND Freshwater percomorph fish of Ixtapa METHODS 832 832 1. Introduction Road 195), next to the bridge that goes through the Soyaló-Ixtapa highway (State the east side of from Tuxtla Gutiérrez City.exposedsite is at This dinates 16°48ʹ42.31ʺN, 92°54ʹ33.58ʺW, 28 km east Ixtapa fossiliferousThe site is located at the coor i fthe present paper is to describe this fossil aim of The unexpected. was unit this in fish a of finding Formation are carbonized leaves; therefore, the the Ixtapa mostabundantThe fossilsat the baseof es-Rocha, 2001;Hernández-Villalba cideans ( horses, rhinoceros, and probos such asteeth of remains, mammal and wood silicified of remains above, whereas dark-gray conglomerates preserve carbonized as well as gastropods and charophytes oogonia; bear angiosperms, leaves andstems of remains of tuff–shales well-stratified the thick and thin the formation, the base of (1992)reported that, atMartínez-Hernández glomerates are sporadicallypresent. and clays, where and calcareous crystalline con conglomerates, sandstones, interbedded layersof part of forming theformation, towardsthe baseof frequent more are tuffs. Tuffs and gravels pebbly pyroclastic materials interbedded with calcite of franca (1996),the Ixtapa Formationa sequence is varado (1965). According to Ferrusquía-Villa unit which wasoriginally named by González-Al Ixtapa Formation, anUpper Miocenegeological to thevolcano-sedimentarysequence known the as the Ixtapa site belong fossil-bearingThe rocksof Hernández-Villalba lafranca, 1996; Martínez-Amador Punta de Llano Formation ( the the Pliocene–Pleistocene volcanic of deposits Formation. Ixtapa Formation The iscovered by theElBosque the Eocenemarinelimestonesof the Coyolar Formation and tal of sandstones conditions, over the Middle Miocene continen fluvio-lacustrine low-energy in deposited is and Ixtapa FormationThe is Late Miocene in age town Ixtapa(Figure 1). the RíoHondo, one-kilometer North fromdown / Boletín de la Sociedad Geológica Mexicana Geológica Sociedad la de Boletín e.g. , DailyandDurham,1966; Menez et al. , 2013). e.g. , Ferrusquía-Vil et al. et al. , 2013). , 2004; ------/2019 2. Materialandmethods (Universidad Nacional Autónoma de México). Mexico Autonomous National University of (Colección NacionaldePaleontología), at the Mexico Paleontology of National Collection of specimendescribed here in the The isdeposited water fishknown percomorph inMexico. fossil alsorepresentsThis fresh the oldest primary North and Central America. the extreme south of in percomorph the of patterns geographic the paleobio be an important clue to understand fish, which despite not being well preserved, could (IGM 4716); cuevasae and include the Paleocene species the species here described same institution of All the taxa used for comparison belong to the al. rusquía-Villafranca, 1996; Hernández-Villalba thearea ( of previouspaleontological and geological studies the Ixtapa Formation follows Lower Pliocene of frominterpretation the Upper Miocene to the Stiassny, 1986; Johnson and Patterson, 1993). The ( the worksclassic osteology on acanthomorph bone nomenclaturethe specimen. The follows of andmeristic data compose the morphometric 11anatomic measures and10counts A total of the specimenwascoated withmagnesiumsmoke. ter contrast between the bones andthe sediment, brushstrokesacid. Toand 5%sulfamic obtainbet needles, andchemically removed withpunctual mechanically removed withthinexcavatorsand thebodywere rockmatrix over manyparts of of patches Thesmall skeleton. the cover rocks, tuff olive byThe sediments, gray formed volcanic caudal fins. and thepelvic dorsal, anal,fins, and and the tip of thevertebral the pectoral column, rior portionof bones onthe head. Furthermore, it lacks the ante isolated undefined only contains and counterpart part and both partially preserves fish fossil This , 2013). Cantalice andAlvarado-Ortega, 2016 i.e. Kelemejtubus castroi , Martínez-Hernández, 1992;Fer, Martínez-Hernández, Cantalice and Eekaulostomus e.g. et - - - - - , 3.1. SYSTEMATIC PALEONTOLOGY 3. Results circles). Modified ofHernández-Villalba Figure 1 Alvarado-Ortega, 2015 (IGM4864-67, 4908-09); Alvarado-Ortega, al. at Tepexi de Rodriguez, Puebla (Espinosa-Pérez Ahuehuetes, Oligocene outcrops of below)sion of (IGM7967;seediscus cyprinodontiform dubious Rosa locality, at Amatitlán, Jalisco State; and the de Santa Barranca 7966), from the Pliocene of dentalis IHNFG 6876),from ChiapasState; andAlaniz-Galvan,tega, 2018(IGM9469-9477; Paleoserranus lakamhae Mapof theStateofChiapas,showing theIxtapa fossiliferous site(blackandcircle) and whitenearby municipalities(black , 1991). Class ACTINOPTERYGII Woodward, 1891 Subsection ACANTHOMORPHATA Subdivision PERCOMORPHACEAE Infraclass TELEOSTEIMüller, 1845 Álvarez 1972 (IGM andArriola-Longoria, ( sensu Wiley andJohnson, 2010) Cantalice, Alvarado-Or et al. (2013). Tapatia occi Boletín de la Sociedad Geológica Mexicana Geológica Sociedad la de Boletín et - - - here describedPerco belongstothe Subdivision species the acanthomorphs, Inside fins. anal and unsegmented beforespines softraysboth dorsal on true because ithas acanthomorphs) (or only Acanthomorphata Subsection the to belongs fish taxonomic characters. Morphological sources of Municipality, Chiapas, Mexico. Formation (Ferrusquía-Villafranca, 1996), Ixtapa Ixtapa the of base the at sediments tuff Miocene Type locality and horizon. anal, andcaudalfinisalsolost. the rays that the dorsal, composes extremity of body, are not preserved (Figure posterior 2). The the of portion antero-ventral the as well as fins), third (includingthehead,pectoral, and pelvic anterior preserved The part inandcounterpart. specimen. Referred (= PERCOMORPHACEA, Gen. andsp. undetermined Wiley andJohnson, 2010) Family IGM 7968, incomplete IGM specimen INCERTAE SEDIS Ixtapa locality, Upper /2019 sensu This fossilThis 833 833 -

MATERIAL AND METHODS / Freshwater percomorph fish of Ixtapa RESULTS Freshwater percomorph fish of Ixtapa RESULTS 834 834 indicate5 mm. supracleithrum. Thescales Scl, ribs; Rb, neuralNs,spine; spine; haemal finHs, spines;dorsal Ds, fin rays; dorsal pterygiophore;Dr, based onbothsides.Abbreviations:Ap,Analfin pterygiophore; Ar,analfinrays;As, analfin spine;Cl,cleithrum; Dp,dorsalfin Figure 2 General features. Description I+8—7+I. pattern the with rays, principal seventeen with fin caudal hypural plates onthe caudalskeleton; andthe the second preural centrum (PU2); five autogenous to havemorphaceae autogenous haemal spine on height without the dorsal andanalfins. height without the dorsal the body relatively deep, occupying about 45%of is peduncle caudal The fin. caudal the of end the to parallel almost is fin anal the of beginning The notched. slightly be to seems dorsal and continuous is fin The preserved. not are fins pectoral pelvic and and jaws, head, The 1). (Table mm 31.2 about is length standard estimated Its flattened. not but low, relatively body the with fish, drical to observe that the specimen isasmallandcylin / Boletín de la Sociedad Geológica Mexicana Geológica Sociedad la de Boletín IGM 7968,apercomorph fishfrom theMioceneofdrawing Ixtapa,side; C)schematic Chiapas,Mexico: A)rightside;B)left Although not complete, it ispossible - /2019 ilar to allneural spines visible, are thin andlong fin vertebrae are stout and short.The others, sim the anal the two centra anterior the originof of smaller andmore haemal spines elongated. The the body. vertebral The centra are progressively areposteriors curved to the posterior portion of anterior ribsare partially preserved; while the The ribs. thin and long of pairs 13 includes skeleton the axial (Table antero-ventral 2).The portion of caudal 15 and more) (or abdominals 13 tebrae, thatsuggest the axial skeleton has at least 28 ver ribs andanterior neural spines, it ispossible to or apophysis isobserved. Bythe presence of skeleton isnot preserved. Noprezygapophysis Axial skeleton. fthe body.of structures, curved towards the posterior portion Teatro oto fthe axial anterior portion of The - - based ontheminimum measure found. Formation (in millimeters). The plus sign indicates anestimation Table 1.Measurements of thespecimenIGM7968of theIxtapa adlpdnl egt 5.03 +3.11 Caudal peduncleheight 5.88 Estimated caudallength 8.77 Post anallength 5.90 Anal finheight Anal finlength 5.13 5.34 Post dorsallength +13.85 Dorsal finheight Estimated dorsallength 15.65 Body height +31.22 Estimated standardlength Measurements oyhih ihu is 11.06 Body heightwithoutfins shows the preural centra (Pu) 3 and 2, urostyle, on the lower lobe, are bone complex lost. This the caudalfin rays, andsome procurrent rays well preserved; onlythe posterior portion of Caudal skeleton. spine, aswell inthesoftrays. expanded pterygiophore supportsthe longest third spines. first Posteriorly,two ventrally and thin one the series andsupportsthe bone of the is biggest pterygiophore first The (Table2). pterygiophores followed by seven soft rays;all supported by nine thebody. There areportion of three stout spines, fin. Anal spine.dorsal be continuous, withaslightnotch before the last to seems It fin. dorsal the compose rygiophores, 22pte and 15softrays, supportedbya total of spines 10 least At lost. are fin dorsal the on spines fin. Dorsal curved posteriorly, with its ventral tip pointed. sal tip rounded; while the last seems to be strongly dor its has and cylindrical is first The preserved. the supracleithrum andthe cleithrum bones are Pectoral girdle. parhypural, five autogenous hypural plates, one The anal fin is on the third posterior the third on is fin anal The The supraneural bones and the first first the and bones supraneural The This portion is very This deteriorated. Only The caudal skeleton isrelatively The IGM 7968 Boletín de la Sociedad Geológica Mexicana Geológica Sociedad la de Boletín - - the minimalvaluefound. Formation. Theplussigninboldindicate anestimationbased on Ixtapa the of 7968 specimen IGM the of Counts 2. Table rnrl 1 2 III+7 Hypural platesformulaH1/H2/H3/H4/H5 I+8-7+I 9 Uroneral 13 Epurals Caudal finformula 28+(+13+15) +22 Anal finpterygiophores Anal fin Dorsal finpterygiophores Vertebrae Ribs Counts Dorsal fin oe yol n eiso spines. posed by onlyone series of com and the ctenii portionissmall,seemingly scales isalmost quadrangular last kind of This the body.on the anterior and middle portion of cycloid; however, there are some ctenoid scales Scales. Pu3 andthe stegural. of gated bones placed between the neural spine the epurals. two are The epurals thin andelon is thin andplaced between the hypural 5and hypural plates 4and5.Posteriorly, the stegural wide andcovers partially the urostyle andthe stegural isanteriorlyand the uralcentra 1. The resents a fusion between the preural centra 1 structure found inmany rep acanthomorphs, hypural 5isextremely reduced. urostyle, The a the same size, while hypural 4iswide, andthe urostyle. The hypural plates 1 to 3 have almost ral plates are triangularandnot fused tothe commonly found onpercomorphs. hypu The not possible to observe the hypurapophysis, tion is not well preserved and, hence, it is parhypuralThe isspatulate; its anterior por reduced (Figure 3). Pu2 isvery autogenous, andthe neural spine of both The Pu3 haemal ad spines and of Pu2 are uroneural (stegural), andtwo (Table epurals 2). Tesae r i n oto them are scales are The bigandmost of IGM 7968 /2019 +X+15 835 835 - - - - -

Freshwater percomorph fish of Ixtapa RESULTS Freshwater percomorph fish of Ixtapa DISCUSSION AND CONCLUSIONS 836 836 4. DiscussionandConclusions Us, urostyle. Thescaleindicates 5mm. spine; Ns,neuralspine; Pu2-3,preuralstegural;haemal St, Hs, Ep, central; hypural ray; epural;H1-5, plates; Dpr, dorsal principal ray; Figure 3 In the Paleogene, there is also a small bony fish fish bony small a also is there Paleogene, the In Ixtapa it is at least 28. but of in the percomorph vertebrae in number of Furthermore, theanal finspineisthe biggest. this last species are stouter, andthe second of Paleoserranus lakamhae fossil herein described resembles the serranid the of fin anal the on threespines of presencethe 2017; Cantalice tega and (Alvarado-Or percomorphs some undetermined sea basses, fishes, flute-mouth known are 2015). Inside the Paleogene strata, currently, Alvarado-Ortega 2013; peninsula (González-RodríguezCalifornia PaleogeneChiapas or of to the the Neogene Baja of represent marine environments andbelong to the the fossil fossil beds with percomorph records of spiny-rayed fishes are scarce inMexico. Most Cenozoic freshwater fishlocalities containing / Boletín de la Sociedad Geológica Mexicana Geológica Sociedad la de Boletín Schematicdrawing of thecaudal fin skeleton ofIGM 7968.Abbreviations: Dbr,dorsal branched ray; Dpcr,dorsal procurrent et al. , 2015; Cantalice and Alvarado-Ortega, 2015; , et al. , 2018). The bodyshape and , 2018).The ; however, the analspines et al. P. lakamhae , 2015;Guzmán, is only 24, is et al. - /2019 , body height and caudalpeduncle slightly deeper. andanalrays dorsal preserved, andthe number of greater a fin, anal and dorsal both on spines true species herein described iseasily distinguished with sedis thus, considered best fishes; as is it Teleostei bony of subgroup any in classification proper the this fossil specimen, for the moment, does not allow by González-Rodríguez mentioned first as such Cypriniformes, Order the of ostariophysians the with affinities some indicate ceratobranchial 5 withankylosed teeth) that could features ( morphological the material andthe specimen seems to have some environments inMexico. Nevertheless, we revised recordoldest percomorph restricted to freshwater Guzmán, 2015),which couldbe considered the (Espinosa-Pérez Cyprinodontiformes the Order considered member of a percomorph Ahuehuetes locality and was originally strata of continental Oligocene in the recovered was fish fossil found inTepexi de Rodriguez, Puebla. This until amore detailed revision. However, the et al. i.e. (2013). Unfortunately, , the well-developed et al. , 1991; , 1991; incertae incertae eidi on h igs iest fextinct Period isfound diversity the biggest of species impossible. Finally, inthe Quaternary making the anatomical comparisonbetween these Ixtapa, are not preserved of inthe percomorph elements (Near andKoppelmann, 2009), which jaw only preserves species Miller, This and 1975. Centrarchidae the family member of reported the percomorph the Lake ChapalaBasin,Jalisco, Mexico, is its of From the Late Pliocene–Early Pleistocene depos Guzmán, 2015:663). (see fin anal the on gonopodium of absence the from distinguish Ixtapa from fish et al. Jalisco (Guzmán de Santa Rosa, Barranca Amatitlán, locality of the familyGoodeidae from the Pliocene species of record is comorph Neogene. In this Period, the oldest Mexican per freshwater perch-like fish in Mexico started in the endemic derived of diversification early the that fossil fish inmorph the Paleogene, we can suggest perco primary After refuting the presence of fMexico. of endemic species inthe central region lishment of estab before the country the portion of southwestern this group already occupied lakes and rivers inthe pri freshwatermary inMexico, percomorph andthat of diversification early the for Ma 6 about of an increase thus, suggesting, Mexico, in fish aperch-like record of oldest andsouthernmost Therefore, the specimen here described isthe from the Pliocene to (Guzmán,2015). the familyGoodeidae, recovered tifom species of freshwater fossil are percomorphs cyprinodon in continental Mexico. To date,waters of the known restricted group the of diversification the and the localization provide new insights about division Percomorphaceae, its Miocene Epoch allow a more detailed identification than the Sub Although the specimen studied herein does not considerably from the Ixtapaspecies. for the species are (except non-percomorph of most however, Mexico; of fauna fish freshwater Micropterus relictus Micropterus , 2013, Guzmán, 2015). The percomorph percomorph The 2015). Guzmán, 2013, , Micropterus relictus Micropterus et al. Tapatia occidentalis , mentioned above),mentioned differing , , 1998;González-Rodríguez Smith, Cavender T. occidentalis , anextinct Boletín de la Sociedad Geológica Mexicana Geológica Sociedad la de Boletín by by ------Acknowledgments References uigtepoeso publication. Finally,during the process of thanks K. Lüneberg, andS. Ramos-Amézquita for help the to manuscript. F. Thanks Vega-Vera,sion of ver final the to improvementsfor reviewer mous Martínez-Melo,A. D. Mayrinck,and ananony to and specimen; the of identification proper the oros, andC. McMahan for about the discussion graphs. We are grateful to P.M. Brito, W.A. Matam alsoto J.Thanks Miguel Contreras for the photo site. Ixtapa the on collected specimen fish unique the UNAM paleo-botanical team for providing the and Ferriz Cevallos S. thank to like would We Cantalice, K.M.,Alvarado-Ortega, J., 2017, Cantalice, K.M.,Alvarado-Ortega, J., 2016, Álvarez, J., J., Arriola-Longoria, 1972,Primer Alvarado-Ortega, J., Cuevas-García, M.,del port toK.M.Cantalice. sup financial providing for UNAM (DGAPA)of to the Dirección General de Académicos Asuntos percomorph (Teleostei,percomorph Actinopterygii) Kelemejtubus castroi 1–24. https://doi.org/10.26879/682 18(3), Electronica, Palaeontologia Mexico: Belisario Domínguez, Chiapas, southeastern fromDanian (Paleocene) of marine deposits ancient armored trumpetfish (Aulostomoidea) Eekaulostomus cuevasae 6–15. Sociedad de CienciasNaturales deJalisco, 6, Jaliscense (Pisces, Teleostomi): Boletín de la goodeido fósil procedente del Pliocene 1–22. https://doi.org/10.26879/536 Mexico: Palaeontologia Electronica, 18(2), southeastern Chiapas, Palenque, from fishes G., Than-Marchese, B.A., 2015,Paleocene Cantalice, K.M., Alaniz-Galvan, A.,Solano-Templos, M., Melgarejo-Damián, Pilar gen. et sp. nov., an ancient gen. and sp. nov., an /2019 837 837 - - - - -

DISCUSSION AND CONCLUSIONS / Freshwater percomorph fish of Ixtapa ACKNOWLEDGMENTS Freshwater percomorph fish of Ixtapa ACKNOWLEDGMENTS / REFERENCES 838 838 Cantalice, K.M.,Alvarado-Ortega, J., Alaniz- Guzmán, A.F., Stinnesbeck, W.,A.F.,Robles-Camacho,Stinnesbeck, Guzmán, González-Rodríguez, K.A.,Espinosa- Arrubarrena, González-Alvarado, I.,1965,Geología del área Ferrusquía-Villafranca, I., 1996, Contribución Espinosa-Pérez, H.,González-Rodríguez, K., Miocene 1966, J.W., Durham F.K., Daily / Boletín de la Sociedad Geológica Mexicana Geológica Sociedad la de Boletín Galvan, A.,2018, 34.2017.1383265 https://doi.org/10.1080/027246 e1383265. 12 37, Paleontology, Vertebrate of Journal Palenque,Mexico: Chiapas, southeastern from the Paleocene near marine deposits 8(2), 127–134. 8(2), 127–134. de laSociedad Mexicana de Paleontología, occidentalis paleontología de lalocalidadtipo de Jalisco: sedimentología y estratigrafía, J., Polaco, O., 1998,ElPaleolago de Amatitán, and Evolution: Germany, Verlag Diversity Dr. Friedrich Pfeil, 9–34. –Global 5 fishes Mesozoic V.H. (eds.) Wilson, H-P., Schultze, record fish fossil Mexican the L., González-Barba, G., 2013, An overview of geológico 507. Petróleos mexicanos, ZonaSur, Informe Ixtapa-San Cristóbal de lasCasas, Chiapas: 109, 130. Universidad Nacional Autónoma de México, área Ixtapa-Soyaló: Instituto de Geología, al conocimiento geológica de Chiapas- el book, 33. de LosGarza,Nuevo León, México, Abstract Nacionalde Ictiología, San Nicolás Congreso Peces Dulceacuícolas Fósiles en México, Applegate, S.P., 1991,Nuevos Registros de Palentology,Journal 40(5),1191–1199. of charophytes from Ixtapa, Chiapas, Mexico: org/10.1016/j. jsames.2018.01.010. https://doi. 137–146, 83, sciences, earth of southeastern Mexico: South American journal fromSerranidae) Palenque, Chiapas, nov.,sp. et a Paleocene seabass (Perciformes: (: Goodeidae): Revista Paleoserranus lakamhae in Arratia G., Tapatia gen. in II II /2019 Martínez-Amador, H.,Rosendo-Brito, B., Fitz- Percomorph 1993, C., Patterson, D.G., Johnson, Hernández-Villalba, D.V., Calvillo-Canadell, L., Guzmán, A.F., 2015,Elregistro fósilde lospeces Near, T.J., Koppelman, J.B., 2009, Species Müller, J., 1845, Untersuchungen über die Menezes-Rocha, J.J., 2001,Tectonic evolution of E.,1992,Caracterización Martínez-Hernández, 52(1), 554–626. Marine Science, a new proposal: Bulletin of and acanthomorphs phylogeny: asurvey of 63, 48–65. Mexicana, Paleontología fósiles: los de foliar fisionomía la Chiapas,utilizando Ixtapa paleoclima de laFormación miocénica en del Inferencia 2013, S.R.S., Cevallos-Ferriz org/10.1016/j.rmb.2015.05.003 de Biodiversidad, 86(3), 661–673. https://doi. mexicanos de aguadulce: Revista Mexicana Science, 1–38. and Conservation: London, Blackwell (eds.), Centrarchid Fishes: Diversity, Biology, Centrarchidae diversity, phylogeny, andphylogeography of Berlin, 109-170, 5láms. Königlichen Akademie der Wissenschaften zu Eingeweide der Fische: Abhandlungen der 75, 183–216. basins, andpetroleum systems: AAPG Memoir Mexico Basin:Tectonics, sedimentary of Gulf C.; Buffler, R.T., Cantú, A. (eds.), The western for regional petroleum systems, i Mexico: Implications basin insoutheastern astrike-slip anexample of the Ixtapa graben, de México, 10,54–64. Geología, Universidad NacionalAutónoma de Instituto del Revista palinoestratigráfico: Ixtapa, Estado de Chiapas—un enfoque ambiental del Terciario de laregión de México: Consejo de Recursos Minerales, 172. Oaxaca, escala 1:250000.Pachuca, Hidalgo, Tuxtla Gutiérrez E15-11, estados de Chiapasy Bravo, C.,2004,Carta Geológico–Minera in Cooke, S.J., Philipp, D.P. n Bartolini, Bartolini, Stiassny, limits andrelationships M.L.,1986,The Cavender,Smith, M.L., T.M.,Miller, R.R.,1975, Michigan 12,29–38. Paleontology University of the Museum of (Jalisco, Mexico): Papers onPaleontology Pliocene-Early the Lake Chapala Basin Pleistocene of Late the from fauna fish a of significance biogeographic and Climatic org/10.1111/j.1096-3642.1986.tb00644.x Zoology, 1(2), 411–460. https://doi. of . Journal the acanthomorph of Boletín de la Sociedad Geológica Mexicana Geológica Sociedad la de Boletín Woodward, A.S., 1891, Catalogue of fossil fishes fossil of Catalogue 1891, A.S.,Woodward, teleost A 2010, G.D., Johnson, E.O., Wiley, (Natural 567. History), the British Museum II: London,Trustees of in the British Museum(Natural Part History), Verlag Dr. Friedrich Pfeil,123–182. Teleosts: Germany,Interrelationshipsof M.V.H.(eds.), OriginandPhylogenetic in groups, monophyletic on based classification esn JS, cute H-. Wilson, H.-P., Schultze, J.S., Nelson, /2019 839 839

Freshwater percomorph fish of Ixtapa REFERENCES