Marys Medicine



Origin and
of Teleosts
Honoring Gloria Arratia
Joseph S. Nelson, Hans-Peter Schultze & Mark V. H. Wilson (editors) More advanced teleosts
Verlag Dr. Friedrich Pfeil • München Acknowledgments . Gloria Arratia's contribution to our understanding of lower teleostean phylogeny and classifi cation – Joseph S. Nelson . The case for pycnodont fi shes as the fossil sister-group of teleosts – J. Ralph Nursall . Phylogeny of teleosts based on mitochondrial genome sequences – Richard E. Broughton . Occipito-vertebral fusion in actinopterygians: conjecture, myth and reality. Part 1: Non-teleosts – Ralf Britz and G. David Johnson . Occipito-vertebral fusion in actinopterygians: conjecture, myth and reality. Part 2: Teleosts – G. David Johnson and Ralf Britz . The Late Jurassic ray-fi nned fi sh peak of diversity: biological radiation or preservational bias? – A teleost classifi cation based on monophyletic groups – E. O. Wiley and G. David Johnson . Structure and relationships of †Brannerion (Albuloidei), an Early Cretaceous teleost from Brazil – Peter L. Forey and John G. Maisey . The caudal skeleton of osteoglossomorph fi shes, revisited: comparisons, homologies, and characters – Eric J. Hilton and Ralf Britz . Validity of the osteoglossomorph genus †Asiatolepis and a revision of †Asiatolepis muroii (†Lycoptera muroii) – Zhang Jiang-yong . The branchial arches of the primitive clupeomorph fi sh, Denticeps clupeoides, and their phylogenetic implication (Clupeiformes, Denticipitidae) – Mário de Pinna and Fábio Di Dario . General overview of fossil and Recent Gonorynchiformes (Teleostei, Ostariophysi) – Francisco José Poyato-Ariza, Terry Grande and Rui Diogo . Cypriniformes: systematics and paleontology – Kevin W. Conway, M. Vincent Hirt, Lei Yang, Richard L. Mayden and Andrew M. Simons. Biogeography of Characiformes: an evaluation of the available information of fossil and extant taxa. – Maria Claudia Malabarba and Luiz R. Malabarba . Evolutionary morphology of trichomycterid catfi shes: about hanging on and digging in – Dominique Adriaens, Jonathan N. Baskin and Hendrik Coppens . Systematics of ictalurid catfi shes: a review of the evidence – Jacob J. D. Egge . Salmoniform fi shes: key fossils, supertree, and possible morphological synapomorphies – Mark V. H. Wilson and Robert R. G. Williams . Morphological development of the axial skeletons of Esox lucius and Esox masquinongy (Euteleostei: Esociformes), with comparisons in developmental and mineralization rates – Amanda Burdi and Terry Grande . Evolutionary relationships of the Aulopiformes (Euteleostei: Cyclosquamata): a molecular and total evidence approach – Matthew P. Davis . Karyological and morphological analysis of divergence among species of the killifi sh genus Orestias (Teleostei: Cyprinodontidae) from the southern Altiplano – Irma Vila, Sergio Scott, Natalia Lam, Patricia Iturra and Marco A. Méndez . Origin and Phylogenetic Interrelationships of Teleosts J. S. Nelson, H.-P. Schultze & M. V. H. Wilson (eds.): pp. 471-480, 5 fi gs., 4 tabs., 1 app.
2010 by Verlag Dr. Friedrich Pfeil, München, Germany – ISBN 978-3-89937-107-9 Karyological and morphological analysis
of divergence among species
of the killifi sh genus Orestias
(Teleostei: Cyprinodontidae)
from the southern Altiplano
Irma Vila, Sergio Scott, Natalia Lam, Patricia Iturra and Marco A. Méndez
Orestias Valenciennes, 1839, a genus of killifi sh classifi ed in the family Cyprinodontidae, is endemic to the Andean high plains (Altiplano) aquatic systems. Species have been classifi ed in four species complexes: O. cuvieri, mulleri, gilsoni, and agassizii. Previous taxonomic studies on the agassizii complex have been based mainly on external morphology. The present study describes the species of the agassizii complex of the southern Altiplano using chromosomal, meristic, and morphometric characters. Species show differences in number and morphology of chromosomes. Meristic data based on examination of large numbers of juveniles and adults show that the characters evaluated were in agreement with the original systematic descriptions; however, we detected a high degree of overlap among species. Multivariate analyses showed that O. ascotanensis, O. agassizii, O. laucaensis, and O. chungarensis could be morphologically distinguished from the other species; only O. parinacotensis and O. piacotensis were not different from each other. Our results show that the chromosomal and morphological char- acters are informative traits in the study of the systematics of the species of the Orestias agassizii complex.
The genus Orestias Valenciennes, 1839 (Cyprinodontidae: subfamily Orestiinae) comprises 44 valid spe- cies, more than half of them inhabiting the Lago Titicaca basin (Lauzanne 1982, Parenti 1984a, Vila 2006). Orestias is a diversifi ed assemblage of fi sh species endemic to high-altitude lacustrine and lotic systems that are geographically restricted to the endorrheic inter-Andean basins. These systems presently lack interconnections and differ markedly in salt composition. The genus ranges from Lago Lacsha (9° S) in central Perú to the Salar de Ascotán (22° S) in northern Chile. It is diagnosed by absence of vomer and pelvic fi ns and reduced body squamation and tooth cusp number. Based on systematic reviews and phylogenetic studies, Parenti (1981, 1984a) and Costa (1997, 1998, 2003) recognized four groups or com- plexes distinguished by body shape, squamation, neuromast pattern, and meristics: the O. cuvieri complex (4 species), mulleri complex (5 species), gilsoni complex (10 species), and the agassizii complex (24 species). The agassizii complex (recognized with 25 species given Vila 2006) shows external morphological varia- tion at the intra and interspecifi c levels, which has been the basis for their taxonomy. Nevertheless, the systematics of this group has been limited by a strong typological orientation. This has motivated our studies of the systematics of the group using both classical morphological and molecular approaches. The species of the southwestern Andes in Chile or southern Altiplano (17° S - 22° S), form part of the "agassizii" complex. This region has been one of the most arid regions on earth since the middle Miocene. The species inhabiting this area occur in lacustrine and lotic systems that belong to different hydro- graphic basins. During the Quaternary, the Altiplano alternated between cold, wet glacial periods and dry interglacial periods. During wetter periods, the Altiplano lakes were deeper than at present. Due to the reiterative expansions and contractions of the lakes, the aquatic organisms have been subjected to drastic environmental variations, particularly those related to changes of water temperature and salinity. These conditions, together with the high altitude, resulted in separate ecosystems with a unique biodiversity. Orestias, therefore, is an interesting model for assessing the role of geographic isolation and abiotic vari- ables in the genetic differentiation and speciation in the southern Altiplano. A fi rst step toward having a better understanding of these processes is an evaluation of the taxonomy of the group. In this paper we evaluate the characters used historically, and we add new characters that assist in distinguishing the species of the agassizii complex in the region.
In the southern Altiplano (Fig. 1), six species of Orestias have been described as members of the O. agassizii complex (Arratia 1982, Parenti 1984a, Vila & Pinto 1986, Vila 2006). These species and their areas of occurrence are as follows: O. parinacotensis Arratia, 1982, Bofedal de Parinacota marsh (18°10' S, 69°20' W); O. laucaensis Arratia, 1982, Río Lauca (18°05' S, 69°15' W); O. chungarensis Vila & Pinto, 1986, Lago Chungará (18°15' S, 69°07' W); O. ascotanensis Parenti, 1984a, springs feeding the Salar de Ascotán (21°29' S, 68°19' W); O. piacotensis Vila, 2006, Laguna Piacota (18°11' S, 69°15' W); O. agassizii, the species with the widest distribution of the complex, from Lago Titicaca (14°09' S, 68°03' W) to the Salar de Uyuni (16°22.56' S, 67°30.85' W) in Bolivia, the Salar de Huasco (20°05' S, 68°15' W), the Río Isluga (19°15' S, 68°10' W) and the Río Collacahua (20°45' S, 69°10' W) in Chile.
The validity of the species described for the agassizii complex has been debated. Villwock & Sienknecht (1995, 1996) postulated that the various species represent ontogenetic stages (juveniles or adults) of the same species, based on the experimental production of fi rst generation hybrids from some species. These authors proposed that some species of Orestias from the southern Altiplano are geographic variants of one species. The taxonomic disagreements could be due to the fact that the original descriptions were often based on relatively few individuals. On the other hand, Parenti (1984a,b), Costa (1998, 2003), and Dyer (2000) maintained that the species of the agassizii complex are taxonomically valid. More recently, Lüssen et al. (2003), using mitochondrial DNA (D-loop), recovered two clades: a northern clade between 17° S - 20° S and a southern clade distributed between 21° S - 22° S. These authors found no major differ- ences among the described southern Orestias species; they reported that O. parinacotensis, O. laucaensis, and O. chungarensis share haplotypes independent of geographic origin. In this chapter we will follow Arratia (1982) who was the fi rst scientist in Chile to describe and generate the discussion on the Altiplano fi shes, and on the speciation of Orestias belonging to the agassizii complex. We add new information for the species of Orestias from the southern Altiplano, based on morphometric and meristic measures on a large number of individuals of different sizes (ontogenetic stages) of each species. We also report new
chromosomal characterizations for the species of this group. Materials and Methods
Species were collected in the localities shown in Figure 1. The specimens examined are deposited at Museo Nacional de Historia Natural, Santiago, Chile (Appendix 1).
Morphological analyses. The following morphometric measurements (standardized by standard length)
were made on 30 individuals of different sizes per species: predorsal length, preanal length, length of head, depth of head, maximum depth, length of caudal peduncle, depth of caudal peduncle, diameter of orbit, mouth width, and preorbital distance. For meristic analyses we also used 30 individuals per species and counted number of lateral line scales and number of rays in dorsal, anal, and pectoral fi ns. In all analyses except for number of vertebrae, we used juvenile and adult specimens. We made vertebral counts from Digital X-ray plates (35 kV, 100 mA, 0.01 sec) from three individuals per species. The data analyses included: (a) Univariate analysis (ANOVA) and posterior Tukey tests, for which the data were log-transformed to satisfy the requirements of ANOVA, and (b) Principal Component Analysis (PCA) on variance-covariance matrices, using MVSP 3.1 (Kovach 1993-1998). Subsequently, a linear discriminant analysis (LDA) was carried out using SYSTAT 11 program (Systat Software, Inc) on the eigenvalues of the fi ve primary PCA axes, with each species as a classifying variable. The discriminate function was evaluated in a classifi cation matrix using the jackknife option. Orestias par Orestias par Orestias agassizi Orestias agassizii Huasco Distribution of species of the agassizii complex of the ge- nus Orestias in the southern Karyotype analyses. Chromosomal characterization was performed on fi sh of both sexes obtained from
the following fi ve locations: O. chungarensis from Lago Chungará, O. piacotensis from Laguna de Piacota, O. parinacotensis from Bofedal de Parinacota, O. agassizii from Salar de Huasco, and O. ascotanensis from Salar de Ascotán (Fig. 1). The fi sh were transported to the laboratory and maintained alive in aquaria until processed. Mitotic chromosomes were obtained by the squash method from branchial epithelia in specimens previously injected with 0.2 % colchicine for two hours. The branchial tissue was subjected to hypotonic conditions in distilled water for 90 minutes, followed by fi xation in 50 % acetic acid. The met- aphase plates were stained with 4 % Giemsa (pH 7.2: Na2HPO4 0.6 M and K2HPO4 0.6 M). Chromosome counts were made using a Nikon Optiphot light microscope; the best metaphase plates were photographed using a digital camera. The short and long chromosome arms were measured using MicroMeasure version 3.2 (Reeves & Tear 1999). The ratio of the length of the short arm of the chromosome to that of the total chromosome expressed as a percentage was calculated (Centromeric Index: CI) to establish chromosome morphology according to Levan et al. (1964). The number of chromosomal arms (fundamental number = NF) was determined following Elder et al. (1993), with metacentric and submetacentric chromosomes treated as biarmed chromosomes, and subtelocentric and telocentric as uniarmed. All fi sh were euthanized using 100 mg/L tricaine methanosulfonate. Table 1 shows the meristic data for the ontogenetic series analyzed. The values obtained for fi n ray counts, scales on the lateral line, and number of vertebrae did not differ between juveniles and adults in any of the species studied; we also detected a high degree of interspecies overlap in meristics. It is important to note, however, that the meristic characters were in agreement with the original species descriptions. The analysis of variance for morphometric characters showed signifi cant among-species differences for all characters (Table 2). Tukey a posteriori tests indicated that O. ascotanensis differed from the other species in head length, depth of head, and maximum depth. No signifi cant differences were found between O. ascotanensis and O. agassizii in orbit diameter, preanal length, length of caudal peduncle, or preorbital distance. Signifi cant differences among species were found for head length, head depth, length of caudal peduncle, and width of mouth among O. laucaensis, O. parinacotensis, O. chungarensis, and O. piacotensis. In this last group, only length of caudal peduncle and preorbital distance varied signifi cantly among The fi rst two primary axes (PC1 and PC2) of the Principal Components Analysis (PCA) accounted for 58.1 % of the variance (Fig. 2). Predorsal length, height of head, preanal length, maximum depth, and length of head loaded most heavily on PC1. Scores on PC1 and PC2 showed some divergence among species, but with broadly overlapping values among various species (Fig. 2). Linear Discriminant Func- tion Analysis (LDA) based on scores on the fi rst fi ve PCA axes (accounting for 90.2 % of the variance) showed signifi cant differences among the species (Wilks' lambda = 0.028; F (25, 428) = 27.7; p = 0.00001), with high rates of correct classifi cation for O. ascotanensis (75 %), O. agassizii (91 %), O. laucaensis (79 %), and O. chungarensis (100 %), whereas < 70 % of the O. parinacotensis and O. piacotensis were correctly clas- sifi ed (Table 3). Table 4 shows the results of karyotyping 36 specimens from fi ve species of Orestias. The diploid numbers ranged from 2n = 48 to 2n = 55. Two groups are clearly present (Figs. 3, 4), one with 2n = 48 (O. ascotanensis, O. parinacotensis, and O. agassizii of Salar de Huasco) and two with higher numbers (O. pia cotensis, 2n = 52; O. chungarensis, 2n = 55). Analysis of variance on morphometric characters in The karyotypes consist mainly of subtelocentric species of the Orestias agassizii group. (centromeric index = CI = 12.5-25 %) and telocen- tric (CI = 1-12.5 %) chromosomes with a gradual variation in size. The morphology of chromosomes among species with 48 chromosomes indicates Predorsal length that the different karyotypes have a uniarmed chromosome formula characteristic of each spe- cies (Table 4, Fig. 3). The karyotype of O. piacotensis and O. chun- garensis (Fig. 4) is characterized by having very Length of caudal peduncle small chromosomes, as well as punctiform chro- Height of caudal peduncle mosomes corresponding to less than 2 % of the haploid complement. We classify the latter as micro-chromosomes following Arratia (1982). The Preorbital distance Summary of meristic characters in species of the Orestias agassizii group. O. agassizii O. parinacotensis O. laucaensis O. chungarensis O. ascotanensis O. piacotensis O. chungarensis O. parinacotensis O. piacotensis O. ascotanensis O. agassizii (Isluga) O. laucaensis PC 2 (21.20 %) –0.001 – Fig. 2.
Principal Component Analysis of 11 morphometric characters measured in species of the agassizii complex
of the southern Altiplano. Values in "( )" represent the percentage of variance accounted for.
Table 3.
Discriminate analysis. Number of individuals of Orestias species correctly classifi ed based on their morphology.
Percentages of correct classifi cations were obtained using a jackknife correction (see text).
ascotanensis agassizii laucaensis parinacotensis chungarensis O. ascotanensis O. agassizii O. laucaensis O. parinacotensis O. chungarensis O. piacotensis Table 4.
Diploid number and chromosomal formula for fi ve species of the Orestias agassizii complex from the southern
Altiplano. Abbreviations: nd, sex undetermined; M, metacentric; SM, submetacentric; ST, subtelocentric;
T, telocentric; NF, chromosome arm number; *, microchromosomes excluded.
Number and sex Number of metaphases 2n NF O. agassizii O. ascotanensis O. parinacotensis O. piacotensis O. chungarensis

Fig. 3.
Karyotypes. A, Orestias agassizii; B, O. ascotanensis; C, O. parinacotensis.
chromosome numbers and the chromosome formulae are different between O. piacotensis and O. chunga- rensis (Table 4). No variations in chromosome numbers were observed among metaphase plates obtained from various individuals of the same species, nor did we fi nd differences between males and females. Our meristic analyses carried out on ontogenetic series showed agreement with original descriptions of the species (Arratia 1982, Parenti 1984b, Vila & Pinto 1986, Vila 2006). There were no observed meristic differences between juveniles and adults. Our results showed a high degree of among-species overlap in meristic characters, making it diffi cult to discriminate species based on both univariate meristic values and multivariate functions of those values. Only O. agassizii and O. ascotanensis could be distinguished at the univariate level. Four species, O. ascotanensis, O. agassizii, O. laucaensis, and O. chungarensis, had high rates of correct classifi cation in the linear discriminate function analysis, whereas O. parinacotensis and O. piacotensis did not. The species located at 18°S, particularly O. piacotensis and O. parinacotensis, show weak morphological divergence, possibly as a result of recent geographic isolation. On the other hand, the karyotype evidence shows that each of the fi ve species of Orestias studied have

Fig. 4.
Karyotypes: A, Orestias piacotensis; B, O. chungarensis. mc, microchromosomes.
a characteristic karyotype, with differences in chromosome number or chromosome morphology, or both. Karyological data on the Orestias agassizii complex are scarce. Lueken (1962) described the presence of 24 meiotic bivalents in O. agassizii from Lago Titicaca (Puno). Arratia (1982) reported the karyotypes of O. laucaensis from Río Lauca as 2n = 51-52 and O. parinacotensis from the Bofedal de Parinacota as 2n = 48. Our results confi rm 2n = 48 for O. parinacotensis and O. agassizii from the Salar de Huasco to the southern extreme of the distribution of this species. Our results for O. ascotanensis extend the 2n = 48 chromosome number of this species to the southern Altiplano. Although O. agassizii, O. ascotanensis, and O. parinaco- tensis had 2n = 48 chromosomes, they differ in the proportion of subtelocentric and telocentric chromo- somes, resulting in different chromosome formulas. On the other hand, O. piacotensis, O. chungarensis, and O. lauca ensis (Arratia, 1982) form a group of species with high diploid numbers (2n = 51 and 2n = 55) and with microchromosomes. Two species, O. piacotensis (2n = 52) and O. parinacotensis (2n = 48), have divergent karyotypes, but showed no morphological divergence, even in the multivariate analysis. The variation we observed in numbers of uniarmed chromosomes of Orestias might be explained by pericentric inversions and addition or loss of constitutive heterochromatin. The number of chromosome arms is conserved (NF = 54), even among species with different diploid numbers (Table 4). This suggests that Robertsonian rearrangements may have played a role in the chromosome evolution of these species. Karyotypic information on the family Cyprinodontidae is limited, with Cyprinodon being the best studied group. In that genus both the diploid number (48) and chromosomal morphology are conserved (Nirchio et al. 2003). Chromosomal rearrangements such as those proposed herein for Orestias of the southern Altiplano appear to have been important in chromosomal evolution in other groups of killifi sh such as Cynolebias (García et al. 2001, García 2006) and Chromaphyosemion (Völker et al. 2005). The presence of microchromosomes in the karyotype of three species of the agassizii complex appears to be unique among cyprinodontids. The microchromosomes may correspond to supernumerary chromo- somes, since B or supernumerary chromosomes (macro or micro) have been reported in a broad range of fi sh species. Venere et al. (1999) suggested that many of the fi sh species which have B chromosomes are distributed neotropically, particularly the perciforms and siluriforms. In contrast with our results, the supernumerary chromosomes in many of the cases occurred in only a few individuals of a popula- tion (Feldberg et al. 2004), or in one population along the extent of the distribution (Moreira-Filho et al.

2004), and even then in only one of the sexes (Ueno et al. 2001). In the present study, microchromosomes were present in all examined individuals of O. piacotensis and O. chungarensis. We are conducting further cytogenetic studies to rule out the possibility of supernumerary or B chromosomes for these microchro- mosomes. This study is a contribution to the karyotype data for the species of the agassizii complex that should be extended to other Orestias species, using conventional and molecular cytogenetic techniques, in order to identify which types of chromosomal rearrangements are involved in the karyotype variations observed in Orestias. Finally, ongoing molecular studies (Scott, Vila and Mendez, unpublished data) suggest that a recent divergence of O. laucaensis, O. chungarensis, O. parinacotensis, and O. piacotensis occurred after the disappear- ance of the Tauca paleolake (18.1-14.1 ka) (Placzek et al. 2006). This information, along with the moderate morphological differentiation in these species of this area, suggests speciation via chromosomal and/or ecological factors. The mechanisms involved in driving the differentiation in this group of species will be an interesting topic of future studies. The authors thank Rodrigo Pardo, Hernán Thielemann, Manuel Contreras, Gonzalo Collado, and Claudio Correa, all from Santiago, for assistance in the fi eld. The present study was partially fi nanced through Multidisciplinary Project MULT 05/04-2 DI and DOMEYKO, Iniciativa transversal 1 of the Universidad de Chile and FONDECYT Projects 1061256; 1080390. Sergio Scott was supported by a MeceSup UCO-214 scholarship. This research was authorized by the Subsecretaría de Pesca, Chile, Resolución Exenta # 1042. Arratia, G. (1982): Peces del altiplano de Chile. – In: Veloso, A. & Bustos-Obregon, E. (eds.). El Ambiente Natural y las Poblaciónes Humanas de los Andes del Norte Grande de Chile (Arica, Lat. 18°28'S) UNESCO, MAB-6, vol. 1, La Vegetación y los Vertebrados Inferiores de los Pisos Altitudinales entre Arica y Lago Chungará:
93-133; Montevideo, Uruguay (Ofi cina Regional de Ciencia y Tecnología de la Unesco para América Latina Costa, W. (1997): Phylogeny and classifi cation of the Cyprinodontidae revisited (Teleostei: Cyprinodontiformes): Are Andean and Anatolian killifi shes sister taxa? – J. Comp. Biol. 2 (1): 1-17.
– (1998): Phylogeny and classifi cation of the Cyprinodontiformes (Euteleostei: Atherinomorpha): a reap- praisal. – In: Malabarba, L. R., Reis, R. E., Vari, R. P., Lucena, Z. M. S. & Lucena, C. A. S. (eds.). Phylogeny and Classifi cation of Neotropical Fishes: 537-560; Porto Alegre (Edipucrs). – (2003): Family Cyprinodontidae (Pupfi shes). – In: Reis, R. E., Kullander, S. O. & Ferraris, K. J. (eds.). Check List of the Freshwater Fishes of South and Central America: 549-554; Porto Alegre (Edipucrs).
Dyer, B. (2000): Systematic review and biogeography of the freshwater fi shes of Chile. – Est. Oceanol. 19: 77-
Elder, J. F., Turner, B. J., Thomerson, J. E. & Taphorn, D. C. (1993): Karyotypes of nine Venezuelan annual killifi shes (Cyprinodontidae), with comments on karyotype differentiation in annual killifi shes. – Ichthyol. Explor. Freshwater 4 (3): 261-268.
Feldberg, E., Porto, J. I. R., Alves-Brinn, M. N., Mendonça, M. N. C. & Benzaquem, D. C. (2004): B chromosomes in Amazonian cichlid species. – Cytogenet. Genome Res. 106: 195-198.
García, G. (2006): Multiple simultaneous speciation in killifi shes of the Cynolebias adloffi species complex (Cyprinodontiformes, Rivulidae) from phylogeography and chromosome data. – J. Zool. System. Evol. Res. 44: 75-87.
García, G., Lalanne, A. I., Aguirre, G. & Cappetta, M. (2001): Chromosome evolution in annual killifi sh genus Cynolebias and mitochondrial phylogenetic analysis. – Chrom. Res. 9: 93-100.
Kovach, W. L. (1993-1998): MVSP 3.1. Multivariate statistical package, version 3.1. – Pentraeth, Anglesey, North Wales (Kovach Computing Services). Lauzanne, L. (1982): Les Orestias (Pisces, Cyprinodontidae) du Petit Lac Titicaca. – Rev. Hydrobiol. Trop. 15 (1):
Levan, A., Fredga, K. & Sandberg, A. (1964): Nomenclature for centromeric position on chromosomes. – Heredi- tas 52: 201-220.
Lüssen A., Falk, T. M. &Villwock, W. (2003): Phylogenetic patterns in populations of Chilean species of the genus Orestias (Teleostei: Cyprinodontidae): results of mitochondrial DNA analysis. – Molec. Phylogenet. Evol. 29 (1): 151-160.
Lueken, W. (1962): Chromosomenzahlen bei Orestias (Pisces, Cyprinodontidae). – Mitt. Hamburg. Zool. Mus. Inst. 60: 195-198.
Moreira-Filho, O., Galetti Jr. P. M. & Bertollo, L. A. C. (2004): B chromosomes in the fi sh Astyanax scabripin- nis (Characidae, Tetragonopterinae): An overview in natural populations. – Cytogenet Genome Res. 106:
Nirchio, M., Cequea, H. & Turner, B. J. (2003): Karyotypic characterization and nucleolus organizer regions in Cyprinodon dearborni (Meek, 1909) from Venezuela. – Interciencia 28 (6): 352-354.
Parenti, L. (1981): Phylogenetic and biogeographic analysis of cyprinodontiform fi shes (Teleostei, Atherinomor- pha). – Bull. Amer. Mus. Natur. Hist. 168: 335-557.
– (1984a): A taxonomic revision of the Andean killifi sh genus Orestias (Cyprinodontiformes, Cyprinodonti- dae). – Bull. Amer. Mus. Natur. Hist. 178: 107-214.
– (1984b): Biogeography of the Andean killifi sh genus Orestias with comments on the species fl ock concept. – In: Echelle, A.A. & Kornfi eld, I. (eds.) Evolution of fi sh species fl ocks: 85-92; Orono (Univ. of Maine Press).
Placzek, C., Quade, J. & Patchett, P. J. (2006): Geochronology and stratigraphy of late Pleistocene lake cycles on the southern Bolivian Altiplano: Implications for causes of tropical climate change. – Geol. Soc. Amer. Bull. 118 (5/6): 515-532.
Reeves, A. & Tear, J. (1999): MicroMeasure for Windows, version 3.3. Free program distributed by the authors over the Internet: Systat Software, Inc., Point Richmond, California ( Ueno, K., Ota, K. & Kobayashi, T. (2001): Heteromorphic sex chromosomes of lizardfi sh (Synodontidae): focus on the ZZ-ZW1W2 system in Trachinocephalus myops. – Genetica 111 (1-3): 133-142.
Venere, P., Suetoshi, C. & Galetti P. Jr. (1999): New cases of supernumerary chromosomes in characiform fi shes. – Genet. Molec. Biol. 22 (3): 345-349.
Vila, I. (2006): A new species of killifi sh (Teleostei; Cyprinodontiformes) from the southern Altiplano, Chile. – Copeia 2006 (3):471-476.
Vila, I. & Pinto, M. (1986): A new species of killifi sh (Pisces, Cyprinodontidae) from the Chilean Altiplano. – Rev. Hydrobiol. Tropicale 19: 233-239.
Villwock, W. & Sienknecht, U. (1995) Intraspezifi sche Variabilität im Genus Orestias Valenciennes 1839 (Teleostei: Cyprinodontidae) und zum Problem der Artidentität. – Mitt. Hamburg. Zool. Mus. Inst. 92: 381-398.
– (1996): Contribución al conocimiento e historia de los peces chilenos. Los Cyprinodóntidos del género Orestias Val. 1839 (Teleostei: Cyprinodontidae) del altiplano chileno. – Medio Ambiente 13:119-126.
Völker, M., Ráb, P. & Kullmann, H. 2005: Karyotype differentiation in Chromaphyosemion killifi shes (Cyprino- dontiformes, Nothobranchiidae). I: Chromosome banding patterns of C. alpha, C. kouamense and C. lugens. – Genetica 125: 33-41.
Appendix 1
List of specimens of the genus Orestias deposited in the Museo Nacional de Historia Natural de Chile, Santiago, Orestias piacotensis Vila, 2006 Orestias agassizii Valenciennes, 1846 Orestias agassizii Valenciennes, 1847 Orestias parinacotensis Arratia, 1982 Bofedal de Parinacota Orestias parinacotensis Arratia, 1982 Bofedal de Parinacota Orestias piacotensis Vila, 2006 Orestias chungarensis Vila & Pinto, 1986 Orestias chungarensis Vila & Pinto, 1986 Orestias ascotanensis Parenti, 1984a Salar Ascotan, Vertiente 2 Orestias ascotanensis Parenti, 1984a Salar Ascotan, Vertiente 2 Authors' addresses: Irma Vila and Sergio Scott, Laboratorio de Limnología, Facultad de Ciencias, Universidad de Chile, Santiago, Chile. E-mail: [email protected], [email protected] Natalia Lam2 and Patricia Iturra, ICBM, Facultad de Medicina, Universidad de Chile, Santiago, Chile.
Marco A. Méndez, Laboratorio de Genética y Evolución, Departamento de Ciencias Ecológicas, Facultad de Ciencias, Universidad de Chile; Las Palmeras 3425, Ñuñoa, Santiago, Chile; Casilla 653. E-mail: [email protected] The origin and the phylogenetic interrelationships of teleosts have been contro-versial subjects ever since Greenwood, P. H., Rosen, D. E., Weitzman, S. H. and Myers, G. S. in 1966 presented a revision of teleost phylogeny. Different taxa (Amia, Lepisosteus, Amia + Lepisosteus, †Pycnodontiformes, †Dapedium, †Pachycormi-formes, and others) have been proposed as the sister group of teleosts. Tremendous advances have occurred in our knowledge of Neopterygii, basal to teleosts, and in their major component the teleosts over the past 40 years. Many new key fossils have been studied, and many extant teleost clades have been traced back to the Jurassic in detailed studies by Gloria Arratia in 1987, 1996, and 2000. In addition to new fossils, a large number of new morphological and molecular characters have been incorporated in recent phylo genetic analyses, adding to our arsenal of approaches. This book gives a modern view of these approaches. It includes a compilation of synapomorphies of numerous teleostean taxa with a new proposal of their classifi cation, a proposal that pycnodonts are the fossil sister group of tele osts, a phylogeny based on mitochondrial genome sequences, separate analyses of basal teleostean taxa (Osteoglossomorpha, Clupeiformes, Gonorynchiformes, Cypriniformes, Characiformes, Siluriformes, Salmoniformes, Esociformes) and the euteleostean Aulopiformes, karyological studies of Cyprinodontidae, and morpho-logical analyses of the posterior part of the neurocranium. A biography of Gloria Arratia is also presented. The book represents contributions to the symposium "Origin and phylogenetic interrelationships of teleosts" sponsored by the American Society of Ichthyologists and Herpetologists (ASIH) and organized by the three editors of this volume and held at the Society's annual meeting in St. Louis, Missouri, on 14 July 2007. At the same meeting, Gloria Arratia was honored with the Robert H. Gibbs, Jr. Memorial Award, 2007, for her outstanding contributions to systematic ichthyology. The volume presents the current state of phylogenetic knowledge of the origin of teleosts and the interrelationships of teleost groups, both key issues in fi sh systematics, based on both morphological (of extant and fossil taxa) and molecular evidence. The many contributors to the volume present and evaluate progress in studying both characters and taxa and in establishing databases (morphological and molecular) that will be of use in future.
ISBN 978-3-89937-107-9


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TOXICOLOGICAL SCIENCES 67, 219 –231 (2002)Copyright © 2002 by the Society of Toxicology Gene Expression Analysis Reveals Chemical-Specific Profiles Hisham K. Hamadeh,* Pierre R. Bushel,* Supriya Jayadev,† Karla Martin,* Olimpia DiSorbo,† Stella Sieber,* Lee Bennett,* Raymond Tennant*,1 Raymond Stoll,† J. Carl Barrett,* Kerry Blanchard,† Richard S. Paules,* and Cynthia A. Afshari*,1