IN HONOROF~. P (PHIL) P~STER, CONSERVATIQNIST

4 l

I i

I

r

1 MA. DE LOURDES LOZANO-VILANO AN D ARMANDO J. CONTRERAS-BALDERAS EDITORS -C José Antonio González-Treviño Rector-

Jesús Áncer-~odrí~uez Secretario General

Rogelio Villnrrral-Elizondo Serretarzo de Extenszón y C~lltura

José Santos García-Alvnrado Dztector de la Facctltad de Crrnczo~Bzológzii~(

Celso José Garza-Aciiña Direccikn de Pub1lcarione.s

First editiori, November 2006

O Ma. de Lourdes Lozano-Vilano and ,kmando J. Contreras-Balderas O Universidad Autónoma de Nuevo LeOn

ISBN: 970-694-336-6

Pririted and made iii Monterrq, México CONSERVATION OF MEXICAN NATIVE TROUT AND THE DISCOVERY, STATUS, PROTECTION AND RECOVERY OF THE CONCHOS TROUT, THE FIRST NATIVE

Conservación de truchas natives mexicanas y su descubrimiento, estatus, protección y redescubrimiento de la trucha del Conchos, la primera especie nativa. i DEAN A. HENDRICKSON1, DAVID A. NEELY2, RICHARD L. MAYDEN2, KARL 1 ANDERSON2, JAMES E. BROOKS3, FAUS1-INO CAMARENA-ROSALES4, RALPH CLITTER5, LISA CUTTER5, ANNA BELLIA DE LOS SANTOS6, GUY W. ERNSTING7, HECTOR ESPINOSA-PEREZ~,LLOYD T. FINDLEY9, FRANCISCO J. GARCIA-DE LEÓN~, ANNA L. GEORGEIO, JOHN HATCH", BERNARD R. KUHAJDA12, KYLE E. MAYDEN13, KRIS1-INA MCNYSET14, JENNIFER L. NIELSENI5, FRAIVK W. PFEIFER16, DAVID L. PROPST17, GORGONIO RUIZ-CAMPOS4, ERlC ST. CLAIR18, JOSEPH R. TOMELLER119 AND ALEJANDRO VARELA-ROMER020

'. University of Texas, Texas Natural Science Center, Texas Natural History Collections, PRC 176 / R4000, 10100 Burnet Road, Austin, Texas 78758 USA. '. Departrnent of Biology, 3507 Laclede Ave., Saint Louis University, St. Louis, Missouri 631 03 USA. 3. U. S. Fish and Wildlife Services, New Mexico Fishery Resources Office, 3800 Cornrnons N.E., Albuquerque, New Mexico 871 09 USA. 4. Facultad de Ciencias, Universidad Autónoma de Baja California, Ensenada, Baia California Norte, México. '. California School of Flyfishing, 141 40 Sunrock Rd., Nevada City, California 95959 USA6. Centro de Investigaciones Biológicas del Noroeste, S.C. Mar Berrneio 195, Col. Playa Palo Santa Rito, C.P 23090, PO. Box 128. La Paz, México. 7. Rt. 2, Box 153, Ellinwood, Kansas 67526 USA R. Instituto de Biología, Universidad Nacional Autónoma de México, México, D.F., México. CIAD-Unidad Guayrnas, Lote 18 Manzana 1, Calle Banamichi, Fracc. Lomas de Cortes, Guaymas Sonora 85450 México 'O. Tennessee Aquarium, One Broad St., P O. Box 1 1048, Chattanooga, Tennessee 37401 USA. ". Domicilio conocido, Colonia Juarez Chihuahua, México. 12. Department of Biological Sciences, Scientific Collections, Box 870345, The University of Alabama, Tuscaloosa, Alabarna 35487-0345 USA. 13. John F. Kennedy Catholic High School, 500 Woods Mil1 Rd,

FACIiLTAD DE C LhCIAS B OL~GCAS, UAhL DEAN A. HENDRICKSOh, EJAL.

Manchester, Missouri 6301 1 USA I4.U. S. Environnlental Protection Agency, 200 SW 35th St., Corvallis, Oregon 97333 USA. 15. USGS, Alaska Science Center, Biological Science Office, 101 1 E. Tudor Rd., MS 701, Anchorage, Alaska 99503 USA. 16. Pfeiffer USA. 17. Conservation Services Division, New Mexico Departrnent of Garne and Fish, PO. Box 25 1 12, Santa Fe, New Mexico 87504 USA. la.Alabarna State Oil and Gas Board, 420 Hackberry Lane, Tuscaloosa, Alabama 35486 USA 19. Cimarron Trading, 8436 Meadow Lane, Leawood, Kansas 66206 USA. 'O. Universidad de Sonora; DICTUS - Departamento de Investigaciones Científicas y Tecnológicas, Herrnosillo, Sonoro, Mé[email protected]

ABSTRACT. The Northwestern Sierra Madre Occidental areas on the biodiversity. In addition to human-indiiced of Mexico is a rugged mountain range covering portions of changes to these diverse ecosystems impacts of invasive Sonora, Chihuahua, Durango and Sinaloa, and is drained aquatic species are becoming more and more apparent, and by multiple Pacific Slope Rivers to the west and the Casas the potential for severely reducing population sizes of spe- Grandes, Conchos, and Nazas to the east. The overall area cies or their extirpation or extinction is real. While severa1 is topographically, climatically and biotically diverse, rang- invasive or exotic species are identified as potentially de- ing from endorheic basins (Casas Grandes) to mountainous structive to these communities, the exotic Rainbow Trout areas elevations up to 3348 m, average mean temperatures (Oncorhynchus mykiss) that is derived from hatcheries or from 10-20°C and precipitation from 250- 1100 mmlyr. The culture facilities within the region represents one ofthe most regions is also geological complex that, combined with these critica1 threats to the aquatic and semi-aquatic biodiversity. other variables, provides a great diversity of both aquatic The native trout of mainland México represent the south- and terrestrial habitats conducive to both biotic endemicity ern-inost salmonids, and are at imminent risk of introgres- and diversity. The over al1 diversity of the region has con- sion andlor replacement by feral Rainbow Trout, tribiited to the recent listing of this region by Conservation Oncorhynchus mykiss. Our recent survey efforts have ex- International as one of six new high-priority biodiversity panded the known diversity and ranges of each of several hotspots. Our understanding of the aquatic and terrestrial distinct forms, which we feel represent valid species. We biodiversity of this region, however, is poorly understudied discuss our discovery of multiple new species from the Si- and in urgent need of rapid investigation by collaborative erra Madre Occidental and focus on a new species of trout communities. The combination of an incredibly rugged land- restricted to the upper Conchos drainage, the fírst native scape, drug and bandit activities and indigenous peoples that species of Oncorhynchus known to occupy the Atlantic Slope have not always been welcoming to visitors has resulted in in Mexico. Many of these taxa are restricted to small areas a general paucity of roads in the region, and the lack of ac- of intact habitat in headwater areas of high-elevation streams, cess has limited inventory studies. Recently, however, ac- and are at risk from a suite of human-associated perturba- cess to the region has changed dramatically and many of the tions, emerging diseases, and introduced species. These areas are now accessible enough for logging, mining, and fishes occupy unique habitats, and represent a diverse por- agriculture practices, al1 exerting extreme pressures in some tion of the Méxican montane ichthyofauna. The habitats on

STUDIES OF NORTh AMERICAN DESERT FiSrlES CONSERVATION OF ME%ICA)\IhATlVE TROUT AND frtE DSCOVERY, STATUS. PROT'ECTOh AND RECOVERY OF ThE COhCHOS TROUT, ThE FIRSF NATIVE

which they depend support a wide range of other aquatic development throughout the region that incorporates a likely organisms, most of which are grossly understudied. The dis- highly successful ecotourism system for the region. covery of the Conchos Trout, derived primarily from the use of GARP niche modeling of a subset of localities of Key words: Sierra Madre Occidental, Trout, Oncorhynchus, previously sampled undescribed native Mexican trout, pro- Aquatic Biodiversity, lnvasive Species. vides only a snapshot of the biodiversity awaiting to be dis- covered in this region. The rugged landscape of the Sierra RESUMEN. El Noroeste de la Sierra Madre Occidental de Madre Occidental simply precludes routine sampling at eas- México es una montaña escarpada con grandes pendientes ily accesible locations of streams and most sampling loca- que abarca una porción de Sonora, Chihuahua, Durango y tions require time-intensive access. In an effort to rapidly Sinaloa, y drena por múltiples ríos de la vertiente del Pacifico assess the biodiversity of rivers of this region we employed hacia el oeste y el Casas Grandes, Conchos, y Nazas hacia I this method to aid in predicting the most suitable and highly el este. El área total es topográfica, climatológica y I probable Mexican trout niches in the region. This method biológicamente diverso, desde una cuenca endorreica (Casas I offered highly efficient and powerful results that not only Grandes) a áreas montañosas con elevaciones arriba de 3348 Ii predicted the occurrence of a previously unknown trout in m, promediando una temperatura entre 10-20" C y una 1 the upper Rio Conchos but also provided excellent predic- precipitación que varía de 250- 1 100 mmlaño. La región es & ; . 9 : tions of available habitats in drainages where previously además geológicamente compleja, que, combinada con &!&~d , unknown trout have been discovered by the Truchas algunas otras variables, provee una gran diversidad de am- Mexicanas team in the last nine years. Multiple threats exist bos hábitats: acuático y terrestre que conduce a endemismos to the biodiversity of the northern Sierra Madre Occidental, bióticos y diversidad. El total de la diversidad de la región including uncontrolled introductions of exotic and invasive ha contribuido a la aparición en la reciente lista de esta región species, emerging diseases such as whirling disease, por Conservation International como una de seis nuevos I Myxobolus cerebralis, infectious pancreatic necrosis (IPN), puntos importantes de alta prioridad por su diversidad. iridioviruses and pathogenic water mold, Saprolegnia ferax, Nosotros entendemos de la diversidad acuática y terrestre land-use practices leading to habitat degradation via over- de esta región, generalmente es pobremente estudiada y es grazing, logging, deforestation and road construction, in- necesaria y urgente, una rápida investigación por creasing human population growth, over-fishing or over- colaboración de las comunidades. La combinación de un harvesting of aquatic resources and global climate cliange increíble paisaje accidentado, drogas y actividades ilegales, reducing surface and ground water in the area and creating y las poblaciones indígenas que no siempre les dan environments more conducive to the spread of invasive spe- bienvenida a los visitantes ha causado una falta general de cies, congregated and dense human populations, and emerg- caminos en la región y la carencia de accesos, ha limitado ing diseases. Immediate actions need to be developed to los estudios. Recientemente, sin embargo, el acceso a la aid in public education as to the threats to these ecosystems, región tiene cambios dramáticos y muchas de las áreas son protection of areas, assessment of diversity, and sustainable ahora accesibles suficientemente para talar y para trabajos

FACULTAD DE ClENC AS B~OLÓGICAS,UAhL CONSERVATION OF MEXLCAN hATWE TAOUT AND TriE D SCOVERY, STATUS, PROTECT Oh AND RECOVERY OF ThE COlvCHOS TROVT, ThE FlRST NATIVE

whicli they depend support a wide range of other aquatic development throughout the region that incorporates a likely organisms, most of which are grossly understudied. The dis- highly successful ecotourism system for the region. covery of the Conchos Trout, derived primarily from the use of GARP niche modeling of a subset of localities of Key words: Sierra Madre Occidental, Trout, Oncorhynchus, previously sampled undescribed native Mexican trout, pro- Aquatic Biodiversity, lnvasive Species. vides only a snapshot of the biodiversity awaiting to be dis- covered in this region. The rugged landscape of the Sierra RESUMEN. El Noroeste de la Sierra Madre Occidental de Madre Occidental simply precludes routine sampling at eas- México es una montaña escarpada con grandes pendientes ily accessible locations of streams and most sampling loca- que abarca una porción de Sonora, Chihuahua, Durango y tions require time-intensive access. In an effort to rapidly Sinaloa, y drena por múltiples ríos de la vertiente del Pacifico assess the biodiversity of rivers of this region we employed hacia el oeste y el Casas Grandes, Conchos, y Nazas hacia l I this method to aid in predicting the most suitable and highly el este. El área total es topográfica, climatológica y / probable Mexican trout niches in the region. This method biológicamente diverso, desde una cuenca endorreica (Casas 1 offered higlily efficient and powerful results tliat not only Grandes) a áreas montañosas con elevaciones arriba de 3348 1 predicted the occurrence of a previously unknown trout in m, promediando una temperatura entre 10-20" C y una

1,q,,,,, ;s;sr;e* the upper Rio Conchos but also provided excellent predic- precipitación que varía de 250- 1 100 mmlaño. La región es gt:r, 4 abs3qedlg tions of available habitats in drainages where previously además geológicamente compleja, que, combinada con ...E&&@ ...E&&@ ~~s8?3FsF ..I":EQ-I ..I":EQ-I unknown trout have been discovered by the Truchas algunas otras variables, provee una gran diversidad de atn- Mexicanas team in the last nine years. Multiple threats exist bos hábitats: acuático y terrestre que conduce a endemismos to the biodiversity of the nortliern Sierra Madre Occidental, bióticos y diversidad. El total de la diversidad de la región including uncontrolled introductions of exotic and invasive ha contribuido a la aparición en la reciente lista de esta región species, emerging diseases such as whirling disease, por Conservation International como una de seis nuevos Myxobolus cerebralis, infectious pancreatic necrosis (IPN), puntos importantes de alta prioridad por su diversidad. iridioviruses and pathogenic water mold, Saprolegnia ferax, Nosotros entendemos de la diversidad acuática y terrestre land-use practices leading to habitat degradation via over- de esta región, generalmente es pobremente estudiada y es grazing, logging, deforestation and road construction, in- necesaria y urgente, una rápida investigación por creasing human population growth, over-fishing or over- colaboración de las comunidades. La combinación de un harvesting of aquatic resources and global climate change increíble paisaje accidentado, drogas y actividades ilegales, reducing surface and ground water in the area and creating y las poblaciones indígenas que no siempre les dan environments more conducive to the spread of invasive spe- bienvenida a los visitantes ha causado una falta general de cies, congregated and dense human populations, and emerg- caminos en la región y la carencia de accesos, ha limitado ing diseases. Immediate actions need to be developed to los estudios. Recientemente, sin embargo, el acceso a la aid in public education as to the threats to these ecosystems, región tiene cambios dramáticos y muchas de las áreas son protection of areas, assessment of diversity, and sustainable ahora accesibles suficientemente para talar y para trabajos de minería, y prácticas agrícolas, todas ellas causando ex- deriva primeramente del uso del modelo de nicho GARP, de trema presión en algunas áreas y su diversidad. En suma, un subconjunto de localidades de muestras previas de truchas los cambios inducidos por el hombre a estos ecosistemas mexicanas indescritas, proveyendo solo una chasquido de impactan con especies acuáticas invasivas se hacen mas y la biodiversidad aguardando para ser descubierta en esta mas aparentes, y el potencial para reducir severamente el región. El paisaje accidentado de la Sierra Madre Occiden- tamaño de las poblaciones de especies o su extirpación o su tal, simplemente excluye la rutina de muestre0 en las real extinción. Mientras las especies invasivas o especies locaciones fácilmente accesibles de corrientes y las exótica son identificadas como potencialmente destructivas locaciones que requieren un intenso tiempo de acceso. En a estas comunidades, la exótica Trucha arcoiris un esfuerzo de rápido acceso a la biodiversidad de ríos de (0ncorhynchu.s rnykiss) que es derivada de granjas o cultivos, esta región, nosotros empleamos este método para ayudar a facilitan que para la región represente una de las mas criticas predecir la probabilidad mas alta y conveniente para la trucha amenazas para la biodiversidad acuática y semi-acuática. mexicana. Este método ofreció resultados sumamente La trucha nativa de las tierras principales de México, eficientes y poderosos que no sólo predijeron la presencia representa la distribución mas sureña de los salmónidos, y de una trucha antes desconocida en la parte superior del Río presenta un inminente riesgo de introgresion y10 reemplazo Conchos, pero también las excelentes predicciones por individuos ferales de la Trucha arcoiris, Oncorhynchus proporcionadas de los hábitats disponibles en los drenajes rnykiss. Nuestros recientes viajes y esfuerzos han expandido donde la trucha ha sido descubierto por el equipo Truchas el conocimiento de la diversidad y la distribución de cada Mexicanas en los últimos nueve años. Múltiples amenazas una de estas distintas formas, que nosotros sentimos, existen a la diversidad biológica del norte de la Sierra Madre 1 representan especies validas. Nosotros discutinlos nuestros Occidental, incluyendo las introducciones incontroladas de 1 descubrimientos de múltiples especies nuevas de la Sierra las especies exóticas e invasoras, surgiendo enfermedades Madre Occidental y nos enfocamos a una nueva especie de como la enfermedad del remolino, Myxobolus cerebrales, trucha restringida al alto Río Conchos, la primera especie infección de necrosis pancreática (INP), iridovirus y moho nativa conocida de 0ncorhyrichu.s conocida, que ocupa la de agua patógena, Saprolegnia ferax, las practicas del uso Vertiente del Atlántico en México. Muchas de estas son de suelo llevan a la degradación de los hábitat por medio restringidas a pequeñas áreas de hábitat intactos en áreas de del sobrepastoreo, tala, deforestación y construcción de cabecera con arroyos de gran elevación, y están en riesgo, carreteras, incrementando en el crecimiento demográfico, asociadas por las actividades humanas perturbadoras sobrepesca o sobre explotación de los recursos acuáticos y enfermedades emergentes, y especies introducidas. Estos cambios climaticos globales, reduciendo las aguas peces ocupan hábitat únicos, y representan una diversa superficiales y de los mantos acuíferos en el área y creando porción de la ictiofauna de montaña de México. El hábitat medios ambientes que llevan a la dispersión de las especies del que ellos dependen, dan soporte a una amplia gama de invasivas, congregadas y densas poblaciones humana y otros organisnlos acuáticos, la mayoría no han sido apariciones de enfermedades. Se necesitan acciones estudiados. El descubrimiento de la Tnicha del Río Conchos, inmediatas en el desarrollo en programas de educación

STUDIES OF NORTh AMERICAN DESERT F SrlES CONSWAT ONQF MEXICAN hATlVE TROUT AhD THE DISCOVERY, 5TATUS, PROTECTION AND RECOVERY OF THE CONChOS TROUT ThE FIRST'NATNE pública en cuanto a las amenazas a estos ecosistemas, 1983, Miller et al. 2005), the aquatic fauna of the Sierra protección de áreas, evaluación de diversidad, y desarrollo Madre Occidental remains poorly known, especially in ar- sustentable a través de la región que incorpora un sistema eas above 1500 m where trout species and many other cool- probable y sumamente acertado de ecoturismo para la región. water adapted aquatic life forms occur. Through more detailed investigations on the distribu- Palabras clave: Sierra Madre Occidental, Tr~ichas, tions, taxonomy and systematics of the aquatic faunas of Ot~corhynchus,Biodiversidad acuática, Especies invasivas. this region we predict that the Northern Sierra Madre Occi- dental will also be recognized by Conservation International ZN TROD UCTZON as a priority biodiversity hotspot for aquatic species. Un- The Northern Sierra Madre Occidental is a mountain range fortunately, these diverse, yet poorly known, aquatic eco- covering portions of four Mexican States (Sonora, Chihua- systems are under an imminent threat of extirpation and hua, Durango and Sinaloa) (Fig. 1). The range is topographi- extinction. Threats include 1) uncontrolled introductions cally, climatically and biotically diverse (Brown 1994), with of exotic and invasive species including hatchery-reared elevations up to 3348 m, average mean temperatures from Rainbow Trout (Oncorhynchus mykiss), the chytrid, 1 O-20°C and precipitation from 250- 1 100 mmlyr. Its geo- Batrachochytridium dendrobatidis and New Zeland logical complexity (Ferrusquia 1993) further increases habi- mudsnail (Potamopyrgus antipodurum), 2) emerging dis- tat diversity that contributed additional justification for its eases such as whirling disease, Myxobolus cerebralis, in- recent listing by Conservation lnternational (Mittermeier et fectious pancreatic necrosis (IPN), iridioviruses and patho- al. 2004) as one of six new high-priority biodiversity getiic water mold, Saprolegt~ia,ferax,3) land-use practices hotspots. The combination of an incredibly rugged land- leading to habitat degradation via overgrazing, logging, de- scape, drug activities and indigenous peoples that have not forestation and road construction, 4) increasing human popu- always been welcoming to visitors has resulted in a general lation growth, 5) over-fishing or over-harvesting of aquatic paucity of roads in the region. As late as 1936, large por- resources and 6) global climate change that is not only modi- tions of the Río Yaqui basin had not been substantially modi- fying habitats to be more conducive to the invasion of non- fied from a pre-European contact state (Leopold 1949). native species but is leading to decreased rainfall and low- Despite recent construction of several new roads that facili- ering of the water table. Al1 of these activities and their tate access, travel within most the range is still extremely synergistic interactions will ~iltimatelylead to the loss of, or difficult, sometimes requiring several days and travel by foot reductions in, the native biodiversity of this ~iniqueregion. or mule to access a single locality. Thus, despite both its Our collaborative international alliance of investigators proximity to the US, and interest in both the terrestrial (e.g. called fi-uchus Mexicanas (est. 1997) (Camarena et al. 200 1, Escalante-Pliego et al. 1993, Ramamoorty et al. 1993, Hendrickson et al. 2003, Mayden 2002, Mayden 2004) is Navarro and Benítez 1993, Ceballos et al. 1998, Navarro devoted to understanding the diversity of native aquatic spe- and Peterson 2004) and aquatic biodiversity of México (e.g., cies of the desert southwest, particularly trout species of Hulbert and Villalobos-Figueroa 1982, Villalobos-Figueroa México. The mission of T,.~uchasMexicanas is: A binational

FACJCTAD DE CIENCIAS BIOL~)GICAS, JANL DEAh A. riENDR CKSOb., €7 AL. organization, est. 1997, founded by scientists and concerned on conducting extensive tieldwork in aquatic ecosystems citizens of México and United States, dedicated to proac- of the Sierra Madre Occidental for samples of native and tive efforts conducive to better understanding the exotic fishes for taxonomic, systematic, biogeographic, eco- biodiversity and conservation of aquatic species of high- logical, genetic and conservation studies. Samples of di- elevation aquatic systenis of the Sierra Madre Occidental, versity from these expeditions have been deposited in natu- especially the genus Oncorhynchus, and the ecosystems ral history collections in México and US and are available upon which they depend, via collaboration with indigenous for research and conservation purposes. To date, our efforts peoples, local communities, and government and private have focused prinlarily on native trout diversity and sam- conservation groups, to preserve these natural habitats, spe- pling for morphological and molecular variability in these cies diversity and ecosystems through partnership, educa- tishes because of the urgency for knowledge of this diver- tion, sustainable development and acceptable models of sity that is under siege by the region-wide threat posed by ecotourism. The efforts of Truchas Mexicanas focus on re- the exotic and invasive hatchery-reared Rainbow Trout vealing the distribution of trout and other aquatic diversity (0ncorhynchu.s mykiss). and habitat needs, evaluating threats to their continued ex- istence, assessing comnlunity structure, niodeling niches of STUDY AREA vertebrate and invertebrate species and communities and Sierra Madre Occidental working closely with Mexican citizens (including iiidigenous The specific foca1 region of Truchas Mexicanos centers on Rarámuri) and private groups and citizen education for the high-elevation (> 1500 m) aquatic habitats throughout the lasting conservation of these ecosystems through protection, northern Sierra Madre Occidental (Figs. 1,2) a north-south I public appreciation and education and ecotourism. The trending volcanic formation ranging Erom Cretaceous to I United States has been a terrible steward of similar high- Miocene in age (de Cserna 1989), with an average eleva- ' elevation aquatic communities of its western and southwest- tion of 2000 m (max. 3348 m). It is a natural biogeographic ern rivers through poor management of once pristine and and physiographic entity (Brown 1994, Debano et al. 1994) diverse ecosystems, introductions of exotic hatchery-reared covering about 300,000 km2. This area is extremely poorly trout into basins witli native trout species, a history of relo- sampled for aquatic organisms; Miller et al. (2005), for ex- cations of trout into different basins, non-existent or poor ample, presented data for 14 Río Conchos localities, only water and land management practices and a lack of fore- four above 2000 111, and almost al1 taken at road crossings. sight for sustainable development, agriculture aiid ranch- This is despite this drainage being an economically extremely ing. Our overarching goal is to prevent this mistake from important and very large, major tributary of the Río Grande happening in the Sierra Madre Occidental south of the US- (Rio Bravo), with relatively good access over much of its México boarder and to educate the citizens and government total area. Our searches of al1 North American fish collec- for the lasting survival of tliese ecosystems and the species tions databases revealed few collections from our study re- they contain. gion and very sparse and uneven spatial sampling across it. The diverse efforts of Truchas Mexicanas have focused While the Yaqui River basin was relatively thoroughly sur-

STUDIES OF NORTH AMERfCAN DESERT FICHES CONSERVATION OF MEXlCAh HATIVE TROUT AND l'Hf DISCOVERY, STATU$ PROTECTIDN' AhD RECOVERY DF ThE CONCHO5 TROUT, TrlE F RST NATIVE veyed for fishes in 1978 (Hendrickson 1980), few recent levels of stocking and culture facilities for Rainbow Trout collections exist, and collections from more remote areas and other activities. In light of the rich interdisciplinary (deep canyons and highest elevations) remain virtually ab- knowledge-base on species and subspecies of this trout clade sent. More southern basins like the ríos Sinaloa, Culiacán, residing north of the US-México border, the near total lack San Lorenzo and Baluarte, prior to our work each had only of knowledge of Mexican taxa is remarkable and unfortu- two to five fish collections above 2000 m. nate. The collective knowledge on non-Mexican taxa, how- ever, provides an exceptionally rare scientific opportunity Taxonomic Groups Included in Surveys to reap disproportionate rewards from detailed inventory and Our taxonomic focus has been on fishes but future efforts analyses of the new, or at least nearly unstudied, Mexican will also include amphibians, riparian reptiles and aquatic taxa. While morphology-based taxonomic and systematic invertebrates. Our preliminary data from morphological and research can be difficult with trout due to phenotypic plas- molecular analyses of trout samples suggest that the Sierra ticity, our incorporation of a strong molecular component Madre Occidental in México supports roughly one-fifth of benefited immediately and immensely from the extensive the planet's trout species, almost al1 of which are currently genetic databases existing on the rest of the clade including unrecognized as distinct or unique. Our investigations have Oncorhynchz~smykiss, 0. chrysogaster and their US and included analyses ofthe observations on the impact of hatch- Canadian relatives. Furthermore, much of the notion of plas- ery-reared or cultured Rainbow Trout on native trout, am- ticity of morphology that is argued for non-Mexican trout phibian and inveitebrate communities and our surveys of has not been observed to the same degree in our Mexican other fishes provided unpublished morphological and mo- samples. Our observations indicate that the argument that lecular sequence data that support the existence of signifi- morphology is of limited value in taxon delimitation is in- cant undescribed diversity in those groups as well. Given trinsically related to a "polytypic" species concept or Bio- the'patterns oE diversity in the historically poorly studied logical Species Concept superimposed on many years oftrout fishes, we predict that collections of aquatic invertebrates species being considered composites of many subspecies or and amphibians will similarly include high numbers of races. Our investigations of Sierra Madre trout employ the undescribed taxa in these even more poorly sampled groups. Evolutionary Species Concept with the Phylogenetic Spe- cies Concept as an operational surrogate (see Mayden 1997, Trout 1999). When non-Mexican trout taxa are evaluated within Although existence of native trout in the Sierra Madre Oc- this context and characters are evaluated in a phylogenetic cidental was first reported in the 1880s, inaccessibility of context, arguments for polytypic species supported by a the region has hindered further detailed studies. The his- confusing mosaic of morphological variation are found to toric focus of Truchu.~Mexicanus on trout is largely a prod- be largely unfounded and support recognition of O. mykiss uct of the general expertise of most participants and the need ssp. north of México as multiple species or lineages (Mayden to rectify a striking lack of knowledge on this extremely 2004, Mayden and Wood 1995, Mayden 1997, 1999). The impoitant clade before it faces demise due to heightened polytypic concept for 0. myki,s,s is unquestionably falsified DEAN A. XENDRICCSON. ET AL.

by the historic occurrence of subspecies in sympatry. We by our studies, note the loss andlor modification of regions find it more profitable to discover and describe diversity in of the mtDNA control region (CR) in the restricted Mexi- Sierra Madre Occidental trout before it is lost or contami- can trout examined relative to Oncorhynchus in more north- nated by introductions because of misguided transplants and erti latitudes. While at presetit the functional significatice of stockings of "subspecies." this difference is uiiknown, the CR is known to contain regu- Our combined morphological and molecular investiga- latory elements and Coskun et al. (2003) hypothesize vari- tions itidicate that in addition to two species masquerading ants of CR are related to longevity, climatic adaptation, or within the Mexican Golden Trout (0. chrysognster) (Ruiz- enhancing the immune system via slowing the turnover of Campos et al. 2003), the trout of the Sierra Madre Occiden- memory T-cells, aild may function in this manner, as well as tal may consist of as many as 1 O new species within what others visa-via binding to tiDNA-encoded regulatory fac- was once thought to be a single subspecies of the Rainbow tors. lntroductions of Oncorhynchus are docuinented over Trout lineage, the Coastal Rainbow Trout (O. mykiss irideus). and over to lead to introgressive hybridization with native

These new taxa inhabit Pacific Slope rivers from the Río species a process that will compromise adaptations of these I Yaqui south to the ríos Acaponeta and Baluarte, and the upper native trout to their more southern latitude environments i Río Conchos, the latter being the first record of a tiative and jeopardize these populations. Such loss of this diver- i trout of the Rainbow Trout clade from the Atlantic Slope. sity (species, genes, proteins) or their contamination via in- i Historically, researchers have thought that the most south- trogressive hybridization with cultured Rainbow Trout is ern populations of Rainbow-like trout of México were the tragic and irresponsible. products of introductions and not native elements of the Native trout are iinportant sources of protein for human fauna (Needham and Gard 1959, Behnke 1992, Miller et al. populations, constit~itinga major dietary component for in- 2005). Wowever, our analyses of varied character suites digenous peoples especially in some regions of the Sierra indicate that these trout are not only native but represetit Madre Occidental. Historical accounts (e.g., Needham and highly distinctive lineages worthy of taxonomic recogni- Gard 1959) discuss use of dynamite to gather trout, several tion and protection. Only in areas near culture facilities for guides demonstrated remarkable prowess with handlines, hatchery Rainbow Trout have we found coloration differ- and ratichers told us how they collect trout by hand, with ences and microsatellite and DNA sequence-based evidence rocks, levers, knives, or using powdered gypsum to stun of feral hatchery trout. fishes. Others showed us severa1 varieties of native plants The native trout of the Sierra Madre Occidental are the that contain natural fish toxins used to collect trout. Native southern-most living native salmonids and are adapted to trout fisheries clearly represent an ~indevelopedeconomic the unicl~iethermal and hydrological regimes of these eco- resource to these regions. The rugged landscape and spec- systems. We have only recently started to discover some of tacular scenery already is driving a burgeoning tourism in- the unique qualities of these species, members of a clade dustry in portions of the Sierra Madre Occidental, most no- typified as cold-water adapted to streams in low north lati- tably in the famous Barranca del Cobre, or Copper Canyon. tudes. As an example, Nielsen et al. (2001). and confirmed Understanding and promoting native trout species diversity

STJDiE2 OF hORTt4 AMERlCAN DESERT F SHES CONSERVATlOh OF MEXlW NATIVE TROUT AhD THE DISCOVERY. STATUS, PROTECTION AI\ID RECOVEKY OF ThE CONCHOS TKO!JT, THE F RST NATIVE

in México provides an incentive to retain native salmonids, gion, Catostomus plebeius, is clearly a complex of severa1 and trout represent an un-tapped incentive for additional undescribed species (Ferris et al. 1982, Crabtree and Buth tourism based on sport fishing and ecotourism. Trout an- 1987, pers. obs.), however, none of these groups have been glers make a substantial contribution to the economy of the rigorously analyzed in a phylogenetic framework . Efforts neighboring American Southwest; in Arizona and New toward describing the additional fish diversity within the Mexico during 200 1, 733,000 anglers spent 5 12 million US Sierra Madre Occidental have barely scratched the surface dollars oii trips and equipment (USFWS 2002). With pub- of the diversity remaining within Catostomus, Gila, Codoma licity, similar expenditures by anglers visiting the Sierra and Canipostoma. Madre Occidental are a reasonable expectation, thus pro- viding a tremendously large new infusion of capital into this Amphibians economically depressed region. Efforis by groups such as Our biotic surveys have also included observations of a va- Trout Unlimited, Conservation International and World riety of amphibiaiis, another group that is poorly invento- Wildlife Fund, in collaboration with developing governmen- ried and studied in this region (J. Campbell, D. Frost, pers. tal aiid private ecotourism initiatives could make this and com.) aiid that will very likely be heavily impacted by in- I 1 sustainable development a reality. troduced Rainbow Trout as well as possible chytrid aiid iridovirus (Ranuvirus; Iridoviridae) infections. The Sierra Other Fishes Madre Occidental hosts a suite of endemic stream-associ- In addition to trout most of the high elevation Sierra Madre ated amphibians that are likely to be complexes of species, I Occidental diversity in other fish groups is in four includiiig Ambystoniu mexicanum, A. rosaceum, ; Cypriniform genera: Cumpostonzcz, Codoma aiid GiZa (Cyp- Pseudeurycra helli sierraoccidentalis, Lithobcrtes rinidae), and Ccrtmtomus (Catostomidae). The first two gen- tarahumara und L. pustulosa. Iiiventories and taxonomic 1 era have only two described species in our study area revisions oithese and other amphibians in the regions are (Codoma ornata and Campostomu ornatum), but our ex- critically important at this early stage before the exotic Rain- I1 amination of live and preserved specimens over the past nine bow Trout becomes more widely established. years indicates that each represents polytypic complexes (some also noted in Contreras-Balderas 1975, Burr 1976, lnvertebrates Miller et al. 2005), and preliminary molecular data reinforces To date, our inventories of invertebrate faunas of high-el- those diagnoses. A few new species have recently been de- evation streams of the Sierra Madre Occidental have been scribed from the area in the other two genera (Cato.stomu.s highly cursory, but have included species of Odonata, leopoldi and C. cahita by Siebert and Minckley 1986; Gila Trichoptera, Ephemeroptera, Plecoptera, Diptera hrevicauda by Norris et al. 2003), but within Catostomus, (Chironomidae, Simuliidae, Tabanidae), Coleoptera species relationships in the region are not well-resolved and (Elmidae, Gyrinidae, Psephenidae). A dietaiy study by A. several undescribed species are knowii (Miller et al. 2005; Reyes Valdez (2005), however, of gut contents o€the differ- pers. obs.). The most widely-distributed species in the re- ent trout from drainages across the Sierra Madre Occidental DEAN 4 riE'uDR CKSON, ETAL.

(summarized for al1 taxa in Table 1) revealed a tretnendous from within and outside of the area. Itidigenous villagers diversity of both aquatic and terrestrial invertebrates. lt is refer to two types of trout, the native "aparí" or "aparique" highly likely that a thorough inventory of the regio11 will and an introduced "arco iris" (Rainbow Trout), atid te11 us reveal unknown biodiversity from these poorly inventoried that the "arco iris drives out the aparí." Througliout much streams. Such inventories are needed as soon as possible of the Sierra Madre Occidental, federal, state and municipal because as exotic Rainbow Trout expand predation pres- governments promote Rainbow Trout culture or grow-out sure will be heightened and biodiversity could be lost. facilities, much like the US government in western drain- ages. Eggs andior larvae are readily available for transport Diversity, Sustainability and Socioeconon~ics to sometiines extremely reinote and minimal facilities where Based on our preliminary data from morphology, nDNA and they are reared for sale. Though promoted as a new source mtDNA, Mexican trout populations and other fish species of food and income for local people, the short-term mon- of these poorly inventoried streams are morphologically and etary gain to be inade by promoting Rainbow Trout-based genetically distinctive and offer important insights into the aquaculture in the mountains of the Sierra Madre Occiden- zoogeography of the region. These species and the amphib- tal is not justiiiable when compared to the long-term worth ians and invertebrates are also important indicator species and value of intact native faunas. Without concerted action of habitat quality and environmental integrity in the region. and development of regional socioeconomic cures for largely Our discovery of severa1 new species of cyprinid and non-sustainable resource management practices, not only catostomid fishes suggests additional still-undocumented will tiative Mexican trout soon be in grave danger of extinc- diversity in fishes, aquatic invertebrates and amphibians in tion, but the ecosystems of the region and associated 1 the region's poorly sampled and studied stream systems. biodiversity stand to be severely negatively impacted by While the future of native trout and other aquatic invasive species and land management issues. Our work with biodiversity of the Sierra Madre Occidental appears bleak Trout Unlimited, World Wildlife Fz~nd,Conservation Inter- and heading down the same path that led to the high leve1 of national, México-Norte and others is critica1 for advancing 1 imperilment ofdiversity of numerous western US taxa, there alternative means of subsistence and economic alternatives remains room for cautious optimism. On visiting the town that focus on carefully managed use of native trout to re- of Las Truchas, Durango, we observed that the municipal place Rainbow Trout. Tourism based on native trout, and basketball court had a leaping trout painted on the backboard. perhaps one day even limited culture of them, is much more lndigenous peoples in the region are very aware of their likely to produce sustainable income in this beautiful rug- native trout, atid in some cases fiercely proud. These people ged country with inultiple other tourist attractions. are also aware of the introduced trout. Most living in the Maintenance of this rich biodiversity and the outstand- countryside where trout occur can easily distinguish between ing array of ecosystems in the Sierra Madre Occidental calls the native and non-native trout when tested with unlabeled for developmetit of an organized effort by indigenoiis peoples color identiiication sheets of the different trout and Gila of the region, supported through Mexican governmental (sometimes also called trout or truchus in some areas) known agencies, to enact legislation for the sustainable develop-

STUDIES OF NORTh AMERICAN DESERT F SHES CONSERVATION OF MEXICAN hATNE TROClT AND.1HE OISCOVERY,STATUS, .. . .PROTfCnQN Ah3 XECOVEW OF TME CONC~OSTROLT, TriE PlRST hATlVE ment of the region before it is severely impacted. Strong such areas (e.g. Parmesan and Galbraith 2004, O'Neal2002). economic incentives exist for maintaining this biodiversity Climate change altering the amount of rainfall in the region if a holistic plan can be developed in concert with partners may also provide environmental conditions more conducive sharing a common goal to maintain the native diversity and to spread of invasive andlor infectious exotic pathogenic its habitat that will serve as underpinnings for the plan. species. The native trout are additionally imminently threat- Scientists and managers of' Puchas Mexicanas as well as ened by introgressive hybridization and competition with representatives from Tmut Unlimited, Wovld Wildlife Fund, introduced Rainbow Trout. Protection of the region's na- Consevvation Intevnational, México-Norte and our other tive biodiversity and educational efforts with indigenous collaborators are willing participants to provide any form people are critica1 and integral efforts that tllust accompany of assistance in developing long-term plans in collabora- the inventory and discovery process. Such efforts must be tion with the indigenous peoples of this important biotic developed with the aid of NGOs, public and private agen- region. cies and governmental bodies through input and direction provided by scientists and sociologists and with a sensitiv- Iinminent Threats to the Aquatic Biodiversity of Sierra ity for indigenous cultures. Only with the efforts and coop- Madre Occidental eration of the indigenous peoples can these sensitive eco- The aquatic biodiversity of the Sierra Madre Occidental, systei-ils continue to flourish. known and to be discovered, is at imminent risk of loss, either by extirpation or extinction, from a whole suite of Habitat Degradation adverse impacts from habitat degradation via logging, road Logging is currently the principal form of legal resource building, overgrazing, infectious disease introduced via ex- extraction in the region, and even the most remote areas otic Rainbow Trout, human population growth and land- visited in expeditions exhibited signs of overgrazing and use practices without education towards sustainable devel- logging. Many stream channels display evidence of abnor- opment. Furthermore, as is now well documented throughout mally high maximum flows and extreme low flow periods, North America, Europe and elsewhere for many taxa characteristic of streams with disturbed watersheds. Loss of (Parmesan et al. 1999; Parmesan and Yohe 2003; Paimesan riparian ground cover through overgrazing andlor logging and Galbraith 2004), distributions of organisms are being can dramatically increase both the amplitude of stormwater forced to shifi upward in both elevation and latitude as a runoff and maximum temperature at baseflow conditions, result of global climate change. Native Mexican trout, oc- and result in reduced survival of adapted biodiversity (Propst curring as they do at the most extreme altitudes on et al. 1992, Li et al. 1994, Knapp and Matthews 1996). Both mountaintops, and constrained to their headwater streams, overgrazing and logging also lead to increased erosion and are severely threatened by and exceedingly susceptible to accumulation of siltation in aquatic habitats. This, in itself, climate change. It is only through greater understanding of is perhaps one of the most significant threats to freshwater their distributions, ecology and evolutionary history that ecosystems worldwide. The negative impacts of siltation humans might be able to help preserve the biodiversity of on locally adapted aquatic cominunities is well documented

FACULTAD DE CIENCIAS BIO-~G~CAS,UAhL DEAN A. hFNDRICKSON, €TAL. to clog the gills of iishes, smothers eggs of fishes, ainphib- To date, while reservoir construction has mostly been at ians and invertebrates, lead to a generally silent elimination lower elevations, there is increasing intcrest in hydropower of microhabitats for al1 species and significatitly diminishes development of rivers of the Sierra Madre Occidental, and the filtration capacity of the hyporheic zone (Williams et al. many smaller reservoirs are being constructcd or proposed 1993, Lydeard and Mayden 1995, Carpentcr et al. 1998, in higher elevation locations. Such reservoirs will lead lo Wi!cove et al. 1998, Boulton et 31. 2003). Trout are espe- an unequivocal loss of valuable habitat for the native aquatic cially susceptible to siltation because successSul egg incu- biodiversity, and along with them will come additional set- bation can otily occur in clean gravels with large amounts tlers, as well as introductions of exotic ilshes, amphibians, of interstitial spaces that provide water tlow sufficient to invertebrates and likcly new diseases that will negatively keep eggs and larvae well oxygenated. We have visited inany impact the nativc biodiversity. Sierra Madre streams that might supporl trout were it not for severe siltation. Furthermore, soil particles washed into lnvasive Raitibow Trout streams serve as vehicles for the transport of toxins like pes- Rainbow Trout have been reared in hatcheries across 1Yorth ticides, herbicides and Sertilizers that can become attachcd America since the U.S. Fish Commission established a hatch- and leach out into aquatic systems (Goldman et al. 1986, ery on the McCloud River, California in 1872. Over the sub-

Carpenter et al. 1998, Osterkamp et al. 1998). Coilstruction sequent -1 30 years, Rainbow Trout have been broadly in- v;laepoa8gz40i ofroads to support logging increases access Sor scttlers and troduced around the globe (Fuller et a1 1999). with kiiii impacts associated with higher human density. Sawmill-re- considerable impacts on local faunas. The first hatcheries in lated contaminants such as sawdust, oils, diesel fue1 and tan- México were founded in 1888 with a shipment OS-25,000 1 i nin leachate have local point-sourcc etkcts on aquatic sys- McCloud River Rainbow Trout eggs toa hatchery in México l I tems. City, but the extent to which progeny from these fish were Few of the many villages of the Sierra Madre Occiden- distributed across the Sierra Madre Occidental is unknown. 1 tal have any form of sewage treatment and contamination However, Rainbow Trout production is now a significant ; of streams by human waste is nearly pandemic. Nutrient business in México, with the greatest production (5,000 tonsl enrichment from livestock and human wastes can drive in- year) in the state of México. Productioti in Chihuahua and creased microbial production, depleting dissolved oxygen Durango is exceeded by 5 or 6 central and southern states and causing catastrophic mortality of native aquatic organ- (far from currently known native trout ranges), but the num- isms (Fleischner 1994, Strand and Merritt 1999). Likewise, ber of new facilities is increasing very quickly throughout increased human populations with limited availability of the country (Gómez Lepe and Sarmiento Frader 1999). food resources has exerted "unnatural" pressure on the tia- Though Chihuahua and Durango are far from the big- tive fish populations. lntense fishing pressure on some gest Rainbow Trout producen in the country (or were in streams has driven some of the diversity to extirpation within 1999 - Gómez Lepe and Sarmiento Frader 1999), one of the the period of tiine that we have worked in the Sierra Madre inost striking field observation from our expeditions was Occidental. that Rainbow Trout culture operations are abundant, and we

STUOIES OF NORTh AMERLCAN DESERT F SrlES CONSERVATlOk OF ME%iCAN hATlVE TROUT AND THE OISCOVERY, STATUS, PROTECMON AhD RECOVERY OF THE COMChOS TROUT, THE FiR5T bATlVE have witnessed an apparent rapid increase in them through- Einum and Fleming 2001) and displace them from the best out the range of Mexican native trout. We have collected habitat. We have documented feral Rainbow Trout in Mexi- Rainbow Trout at numerous sites in ihe Sierra Madre Occi- can rivers and have evidence from microsatellite markers dental, both at areas in close proximity to hatcheries or cul- for gene exchange in some areas; thus demonstrating that ture facilities as well as in relatively remote headwater they are surviving in these drainages and non-native genes streams. Some local entrepreneurs have also developed basic are introgressitig into native trout populations. This may be grow-out Facilities by diverting portions of streain channels the inost disastrous long-term impact, as even low levels of into side areas dammed by cetnent block, cobble and ce- hybridization with introduced individuals of low fitness can inent or even galvanized sheet metal roofing; these make- lead to breakdown of locally-adapted gene complexes and shift facilities are Frequently destroyed by floods releasing reductions in fitness of the introgressed population their captive non-native trout. lntroduction of Rainbow Trout (McGinnity et al. 2003). lntrogression with introduced Rain- in Sierra Madre Occidental streams will result in a situation bow Trout is one of the most severe threats to cutthroat closely paralleling that of cutthroat trout in the U.S., where (Weigel et al. 2003), Gila (Rinne 1990), Apache (Rinne pure native populations are largely restricted to headwater 1990) and California golden trout (in Rainbow Trout liti- streams. Populations in such habitats are more susceptible eage) (Cordes et al. 2006), and likely represents the greatest to both demographic effects of low population size, as well threat to native Mexican trout as well. as externa1 threats as climate change, droughts, fire, human The potential for introgression is exacerbated by behav- utilization, etc. This suite of threats has led to a recent reas- ioral and life-history attributes of salmonids; as hatchery sessment of the long-tenn viability of conservation strate- fish are usually larger at the same age as native tish, and gies based on isolation (Kr~iseet al. 200 1, Hilderbrand and body size plays a large role in reproductive success in saltno- Kirschner 2000, Novinger atid Rahel 2003). Our genetic nid populations (Foote and Larkin 1988, Dowling and Childs assessments of soine populations of native trout reveal simi- 1992, Rosenfield et al. 2000, Ostberg et al. 2004). Larger lar evidence for genetic bottlenecks. males are preferred by females (Foote 1989) and may ex- Abundant literature clearly demonstrates that hybridiza- clude smaller males from access to females (Quinn and Foote tion between Oncovhynchus spp. is a major obstacle to ef- 1994). Larger females occupy higher quality nest sites (van fective management of native stocks throughout the range den Berghe and Gross 1989, Foote 1990) and dig deeper of the genus in North Atnerica. Recent studies in Montana nests (Steen and Quinn 1999) resulting in lower mortality (Hitt et al. 2003) suggest that even areas previously thought of eggs and higher larval survivorship. safe from introgression are at risk. While hatchery-reared The negative impacts of Rainbow Trout are not limited Rainbow Trout, whether intentionally stocked or accidental to native Mexican trout. Introduced Rainbow Trout have escapees, may not be very successful due to lack of adapta- been strongly implicated in declines of both tion to local environmental conditions (Leider et al. 1990, macroinvertebrate and amphibian communities in western Flemitig and Petersson 200 1, McLean et al. 2004) they may North America (Taylor et al. 1984, Dunham et al. 2004). still competitively interact with native trout (Griffith 1988, Trout impact composition and function of benthic commu-

FACJLTAD DE CIENCIAS B~OL~GICAS.JAN? DEAN A. HENDRICKSON, ETAL.

nities by eithcr al~cringbcliavior of invertebrates (Peckarsky or exchange of water in culture I'acilities (Bartholoinew and et al. 2002), or throiigh dircct predation (Simon and Wilson 7002). Thc rapid spread of Mj:uoholus cer-ehrrrli.~, Townsend 2003). Trout tcnd to select larger tbod iterns, cs- the causnl agcnt of v~hirlingdisease, is iinplicated in cats- pecially grazing caddisllies (Flecker and Townsend 1994. strophic declines in lrout populations in the wcstern US Mclntosh cí al. 1996) aiid predatoiy stoneflies (Hcrbst cí al. (Bergcrscn and Anderson 1997). Siinilarly. icccnt discov- 2003), and tlie effects of prcdation can includc incrcases in ery of inl'cc~iouspancreatic necrosis (IPN) in trout-based both algal biomass and abundante of chironomid midges aqliaculture facilities iii México (Ortega et al. 7002) is of (1-ierbst et al. 2003). Wiíli [he incieascd algal biomass thei-e grave coiicern. IPN is a highly \,irulent virus causing sub- can be toxic algal blooins (Cai-pentcrct al. 1998) and streains ctantial inortality in salmonids (Reno 1999). Ifwild popula- exposed to increased suiilight through clear-ciitting oi' Sor- tions nrc cxposed they are potentially compromised, and oiir ests will increase in teiiiperatuie and lead to anoxic coidi- icscarch indicates that IPN-exposed hatcliery stocks were tions that will be intolerable to natix species. shipped to culturc tiicilities in Cliihuahua witliin the range Populations of several ampliibians. particularly ranid of nativc trout (Cierardo Zamora, perc. cornin.). frogs of tlie geniis Lithohurc.~(foriiierly Rana) arid We llave no insights as to how native troiit will responci nmbystoinatid salainaiiders appcar particularly susccptiblc to these ncw pathogens that have now almost surely been to iinpacts Sroni introduccd trout (Drost 1996, Tyler et al. cariied into streains in somc areas ofthe Sierra Madre Occi- 1998. Matthe~vsct al. 7001, Vredcnburg 2004, Knapp 2005, dental. Clcarly the longer we wait for preventive measures, Welsh 2006). as both groups lack chemical defcnses to lish tlie inore pervasive the impacts of these complex issues will (Kats et al. 1988) and use perinancnt bodies of water for become, furthcr coiil'ounding their rcsolution and exacer- reproductioii. Introduced salmonids prey on larvae of both bating biodiversity loss. The huge part of the diversity of taxa, and can have ma.jor impacts on local populatioii si7e OncorhU~/ich~lsresiding in México remains more vulnerable (Welsh et al. 2006). Interestingly, in the past sevcral years than ever io extirpationiextinction via hybridization. com- of'our expeditions, only sites without fishes or with native peíition. predation, diseases and habitat alteration. The same salinonids have been observed to maintain populations of factors will. OS course, have repercussions throughout the Arnhj.~tol?iaor Lithohatw. As this portion of the Sierra Madre aquatic ecosystems, exlending obviously far beyond fiches. Occidental hosts a suite of endemic stream-associated am- phibians, including Amhystoma mL";icanum, A. rosaceum, Other Emerging Diseases Pseudezrr.j.cerr helli sierruoccidentalis, Lithohutes The decline of aniphibian populations, both in the Sierra ta~-uli~~muruand L. piisti~lo~sa,the impacts of Rainbow Trout Madre Occidental (e.g. Clarkson and Rorabaugh 1989, Hale introductions could be devastating. et al. 7005) and at a worldwide scale (Blaustein and Wake 1990, Wake 1991. Wake and Morowitz 1991, Houlahan Fish Diseases and Hatchery Rainbow Trout 2000), is a source of considerable concern. While causal Aquaculture of Rainbow Trout poses a high risk of infect- factors ranging from cliniate change (Carey and Alevander ing native trout with exotic diseases, either through escape 2003, Pounds et al. 2006), UV radiation (Kiesecker and

STUDIES QF NORTH AME RlCAN DESERT FISHES CONSERVATION OF WiCAN NATlVE TROW AND TiiE DISCUVERY, STATUS, PROTECTION ANO RECOVERY OF THE CONCHO5 TROUT, THE FLRST NATM

Blaustein 1995, Blaustein and Kiesecker 1997), air pollu- suggest complete mortality of some species of infected tion (Clarkson and Rorabaugh 1989, Beebee et al. 19901, postmetamorphic frogs (Briggs et al. 2005), the presence of pesticides (Davidson et al. 2002, Relyea 2004), endocrine B. dendvobatidis in wild populations is not automatically a disruptors (Reeder et al. 2004), diseases from fish culture death knell for a population (Berger et al. 1998, Davidson (Kiesecker et al. 2001), emerging diseases (Laurance et al. et al. 2003, Daszak et al. 2005), and some species (Xenopus 1996, Berger et al. 1998), introduced bullfrogs (Hayes and laevis, Lithob0te.s catesbiana) may serve as vectors of the Jennings 1986. Kiesecker and Blaustein 1998) and fishes fungus into new areas (Briggs et al. 2005, Blaustein and (Bradford 1989, Kiesecker and Blaustein 1998, Tyler et al. Dobson 2006). However, enough studies have implicated 1998, Matthews et al. 2001) have al1 been implicated at lo- chytrids as a principal cause of amphibian declines that iden- cal scales, considerable disagreement over the relative role tification of the current range of the chytrid would allow of each of these factors exists (Collins and Storfer 2003). additional attention to be placed on infected populations. However, emergent diseases appear to play a large role Climate change may also exacerbate effects of these patho- (Berger et al. 1998, Daszak et al. 2003, Ron et al. 2003, gens (Alford and Richards 1999, Berger et al. 2004, Retallick et al. 2004). Populations of Chiricahua and Piotrowski et al. 2004, Ron 2005) and changes in the climatic Tarahumara leopard frogs (Litlzobates chir.icahuensis and characteristicsofthe Sierra Madre Occidental with global warm-

'r$';+FPg.. L. tavahumat.a, respectively) in Arizona crashed during the ing can provide habitats and conditions more conducive to their .. . :,.a.,ar.;@&a&$, late 1970s and early 1980s (Clarkson and Rorabaugh 1989). invasion or spread throughout the aquatic systems. ,:,~,.,2p.~.. - z~;.S,TS~S One is now extirpated from the US. Their decline has been tied to a chytrid, Batvachocl7jtt.idium dt7t7dt.obatidis (Hale Other Exotics et al. 2005). Other evidence suggests that an iridovirus may The New Zealand mudsnail (Potamopyrgus antipodavum) be spreading across much of North America (Jancovitch et is a small (<5mm), invasive hydrobiid snail that is rapidly al. 2005), with similar effects on amphibian populations. spreading across coldwater streams in western North Severa1 studies have documented transmission of America. lt is currently known to occur in 10 U.S. states, iridoviruses (Mao et al. 1999) and the pathogenic water mold including Arizona: but has not to date been reported from Sap~ole~yni~l,fevax (Kiesecker et al. 2001 ) from introduced México. lt reaches densities of up to 800,000/m2 (Dorgelo fishes to native amphibians. All of these were recently re- 1987), carpeting substrates and causing dramatic reductions in viewed in a scientific symposium involving experts (2006 other benthic invertebrates. lt is not generally used as food by American Society of Ichthyologists and Herpetologists An- fishes (M. Vinson, unpubl. studies suggest -45% of mudsnails nual Meeting, New Orleans, LA, Symposium). Amphibian ingested by trout pass through the digestive tract alive), and populations in high-elevation habitats of the Sierra Madre has the potential to seriously impact streams of the area. Occidental have not been thoroughly surveyed, and the in- cidence of Batvachochytvidiirm dendvohatidis and Climate Change and Sierra Madre Occidental Biodiversity iridoviruses in the region will provide valuable data for fo- Global warning under a predicted 2xC0, model is expected cused management efforts. Despite laboratory studies that to negatively affect coldwater fishes and other biotas of these

FACULTAD DE CIENCIAS BIO~ÓGICA~,UANL DEAN A. HENDRICKSON. ETAL cornrnunities overall, although the individual effects of cli- as we pass by them enroute to higher elevations. Though mate change may bc dificult or irnpossible to predict (Hauer rnany of our collectioiis have been niade at locations acces- et al. 1997). Longitudinal displacernent of trout upstrearn as sible via 4-wheel drive vehicles, many more have required a response to carrning (Meisner 1990, Rahel et al. 1996, sornetirnes long hikes. Though rnany of our collections have Eaton arid Scheller 1996, Jager et al. 1999) is unlikely in the been rnade well within areas determined by the U.S. Drug Sierra Madre Occidental since they are already found only Enforcernent Agcncy to have the highest densities of Can- in the uppermost headwater strearns, generally with dry riabis and Opiurn cultivation in the regiori (U.S Drug En- reaches for some distance above, and sonletimes below, es- forcenient Agency 2001 a, b), occasionally local guides have pecially during droughts. Results obtained by Jager et al. advised against collecting in certain areas that are known to (1 999), who docurnented an irnprovernent of headwater habi- have high levels of drug-related crirne, thus further affect- tat for coldwater fish in high-elevation strearns in Colorado ing distribution of our sarnpling. under global warming scenarios, are not applicable to our Fiches Iiave been collected using a variety of gears, pri- study arca. In that study, persistence and population dynarn- marily D.C. electrofishing using Smith-Root battery and ics are controlled by cold winter ternperatures and forma- gasoline-powered backpack units from Coffelt adjiisted to tion of anchor and frazil ice, both of which are unknown in function efficiently at each location without causing undue the Sierra Madre Occidental. Persistence of trout and otlier harrn to individuals and populations (Snyder 2003). We have fishes under typical conditions in the Sierra Madre is often also used a diversity of nets, rnostly sniall seines of various dependent on availability and spatial distribution of deep rneshes, gil1 nets, and dip nets, and have deployed sport fish- pool habitats, which serve as refugia during droughts. Thus, ing equipment (rod and reel). Occasionally we purchased the aquatic faunas of the Sierra Madre Occidental will likely specirnens, especially Rainbow Trout frorn culture facili- be highly negatively irnpacted by global warming, especially ties. frorn loca1 residents. Our observations on arnphibians, if logging in riparian areas continiies and exposes stream reptiles and invertebrates have been rnade coincident with areas to more sunlight, and hurnan population growth and fish collection activities, though we routinely dedicate sorne agriculture activities continue to increase demands on sur- time at each field sites for targeted observations of these face and ground waters. groups, including surveying invertebrates by turning rocks and using dip nets. ME THODS All of our collections have been located using GPS re- Field Collections ceivers while in the field. Though details habitat conditions Our collections have been done primarily in winter and have not generally been rigoroiisly qiiantifíed, general de- spring months, and the localities sarnpled have been prefer- scriptions of al1 habitats have been recorded in standard ich- entially those that we judged most likely to harbor trout, thyological fieldnotes. based on elevation (generally > 1500 m) and general habi- Specimens for morphology have been preserved in 10% tat conditions, advice of local residents, and GARP niodel- forrnalin after fin clips are taken and preserved in absolute ing. Occasionally we have collected lower elevation strearns ethanol. After transport to the lab. specirnens are moved to

STUDlES OF NQRTH AMERICAN DESERT FISHES CQNSERVATION OF MU(1CXN MAflVE TROUT AND TnE DISCOVERV, STATUS, PROTECTION AND RECOERY OF THE CONCHO5 TRDUT, THE FIRST NATWE

70% ethanol for long-term storage. Occasionally entire Lorenzo). These analyses were conducted by Anna Bellia smaller specimens have been preserved in the field directly De Los Santos Camarilla and Dr. F. J. Garcia de Leon at in absolute ethanol. Centro de Investigaciones Biologicas del Noroeste (CIBNOR), La Paz, Baja California Sur, México. Genetic and Phylogenetic Analyses Tests of linkage disequilibriuni, HWE and genotypic Tissues from 246 specimens representing 20 wild popula- differentiation were performed using GENEPOP (Raymond tions of Mexican Oncorh~ynchuscollected between 1997 and and Russet 1997). Genetix was used for Factorial Corre- 2002 were air-dried or preserved in 95% ethanol. An addi- spondence Analysis. Analysis of molecular variation tional 3 1 specimens were obtained from three hatcheries in (AMOVA) was performed using Arlequin v2.0 (Schneider México. Total genomic DNA was extracted with QIAGEN et al. 2000). DNEasy kits following QIAGEN protocols. Twelve DNA sequences (mtDNA & nDNA) were deri~edfor microsatellite loci (Omy2, Omy27, Omy77, Omy207, 27 1 native Mexican trout and hatchery or cultured Rainbow Oniy325, Onem8, Onenill, Otsl, Sfo8, Ssa14, Ssa85 and Trout individuals for Control Region, ATPase 618, GHlC Ssa289) were amplified for the preliininary study using stan- and GH2C. Sequences for outgroup taxa and the data set of dard PCR protocols described in Nielsen and Sage (2001); Bagley and Gall (1998) were downloaded from GenBank al1 niicrosatellite data generation Por these saniples was con- and included in al1 analyses. Outgroup taxa included ducted in Dr. Jennifer Nielsen's laboratory. Anchorage, 017corhynchlrs clarkii, O. masou and 0. keta. Phylogenetic Alaska. Microsatellites were visualized on a LI-COR Long analyses were examined under the maximum parsimony Reader 4200 automatic sequencer with Gene ImagIR (v3.0) (MP) criterion using PAUP* (Swofford 2000) and with software. Mitochondrial D-loop sequence data was gener- Bayesian analysis (Huelsenbeck and Ronquist 2001). Heu- ated from a subset of 45 individuals and amplified using ristic searches were conducted with characters weighted primers DLI F and DL4R (Bagley and Gall 1998). ATPase 6 equally and unordered. Gaps were treated as missing data. and 8, and GH 1 C and GH2C were amplified from a further Starting trees were obtained by stepwise addition, with 1000 subset of 20 individuals using primers from Kontula et al. random addition sequence replicates and TBR branch swap- (2003) and Phillips et al. (2004). QIAGEN gel purification ping. Support for nodes was assessed using nonparametric kits were used to purifi PCR products prior to sequencing jackknife resampling (37% deletion, 1000 pseudoreplicates; on an ABI3 100 Genetic Analyzer. Conlplete sequences were Farris et al. 1996). Jackknife scores above 85 were consid- assenibled from individual electropherogranis, edited and ered well supported (Zander 2004). Bayesian analyses uti- aligned with BioEdit v6.0.7 (Hall 2001). lized MrBayes v3.0.4 (Huelsenbeck and Ronquist 2001). A Additionally, two microsatellite loci (Ssa289, Onel 1) four-chain heated MCMC analysis was run for 2,000,000 have been analyzed for a total of 545 individual trout from generations, with sampling every 1000 generations. expeditions up to 2004. This included 257 wild individuals Stationarity was reached by 100,000 generations; to be con- and 48 hatchery in'dividuals (1 8 Yaqui trout from G. Zamora senative, the first million generations were discarded as and 30 0.n~.vkiss from Granja Truticola on A. la Sidra, San burn-in. Posterior probabilities above 90 were considered

FACULTAD DE CIENCIAS BIOLÓGICAS, UANl DEAN A. HENDRICKSON, ETAL. to be well supported (Zander 2004). respectively). The training data are resampled to 1250 train- ing data points, and 1250 points are randomly sampled from Morphological Variation al1 cells where training data do not occur (as a pseudo-ab- Native Mexican trout are being evaluated for variation in sence data set). These 2500 points are used to develop the standard characters used routinely in studies of other spe- rules and as intrinsic testing data (for evaluation of rule cies of the genus, including coloration patterns of breeding predictivity). Spatial predictions of presence and ahsence and non-breeding adiilts and juveniles, disposition and color can hold two types of error: omission (areas of known pres- of the lateral stripe, arrangement, size and shape of the smali ente predicted absent) and commission (areas of known melanophore spotting, number of parr marks, number, shape absence predicted present), which can be summarized in a and size of auxiliary parr marks. vertebral counts, fin-ray measure of predictive accuracy as the percentage of points counts and scale counts. correctly predicted as present or absent (the correct classifi- cation rate of Fielding and Bell (1 997)). Changes in predic- GIS and GARP modeling tive accuracy from one iteration to the next are used to evalu- We used the Genetic Algorithm for Rule-set Prediction ate whether particular rules should be incorporated into the (GARP; Stockwell and Peters, 1999; Anderson et al., 2003; model or not, and the algorithm runs either 1000 iterations Peterson and Robins 2003) to estimate species' ecological or until convergence (Stockwell and Peters, 1999). The fi- niches and predict their distributions throughout our study nal ecological niche model rule-set is then projected onto area. GARP is a machine-learning approach for detecting the digital maps that are the environmental data sets input associations between known occurrences of species and into the algorithm to identify areas fitting the model param- landscape characteristics, particularly when the ecological eters, a hypothesis of the potential geographic distribution landscape is complex and relationships are not straightfor- of the species. Predictive accuracy of the resulting model is ward (Stockwell and Noble, 1992; Stockwell and Peters, evaluated based on the extrinsic testing data. Accuracy 1999). lnspired by models of genetic evolution, GARP metrics based on these data are used to select the 10 best- models are composed of sets of rules that "evolve" through subset models from al1 models generated. an iterative process of rule selection, evaluation, testing and We georeferenced Mexican trout collections extracted incorporation or rejection (Holland, 1975; Anderson et al., from museum specimen records and personal collections, 2002). GARP searches for non-random associations between scientific literature and our own field surveys (Hendrickson occurrences of species (georeferenced localities in geo- et al. 2003 and additional unpublished data from Truchas graphic coordinates) and environmental variables (¡.e., digi- Mexicanas team surveys). Environmental data sets used in tal maps of relevant ecological parameters). GARP divides our GARP analyses included 13 digital maps ("coverages") occurrence data randomly into data input into the genetic summarizing aspects of topography (elevation, topographic algorithm for model development ("training") and an inde- index, slope, aspect and flow accumulation, from the US pendent data set ("extrinsic testing data") for evaluation of Geological Survey's Hydro-1K data set, native resolution model quality at user-determined proportions (80% and 20%, 1 x 1 km), climate (as absolute and average minimum and

STUDIES OF NORTH AMERICAN DESERT FlSHES CONSERYATJCX-4 OF MMICAN NATrVE TROUT AND THE DISCOVERY, STATUS, PRQTFCilON PJJD RECQVERY Of THE CQNCHOS fROUT, THE FlRST NATIVE maximum temperature (1 : 1.000,000)), precipitation rios Acaponeta and Baluarte. Prior to these efforts sporadic (1 : 1,000,000), evapotranspiration (1 :4,000,000) and soils samples of Mexican Golden Trout were known from the (1:250,000 polygon coverage). Climate and soil coverages rios Fuerte, Sinaloa and Culiacan systems and reports of a are from the CONABIO digital map library, available at: Rainbow Trout-like trout were known from the upper Río I~ttp:llconabioweb.conabio.gob.mxhetacartolmetadatos.pl. Yaqui and the endorheic Casas Grandes basin. Our collec- All coverage data were resampled at 0.01 x 0.01 decimal tions made since the publication of the Hendrickson et al. degree resolution and clipped into 2 datasets; one consist- (2003) review have added 20 additional localities from ing of al1 of México and one to a buffered distance of 100 throughout our study area, expanding the known distribu- km of the locality data. Models are built on the buffered set tion of native trout in México. and then projected onto al1 of México to facilitate analysis. Once the model was developed for Pacific Slope Mexi- Genetics can trout species, using the same topographic and remotely Microsatellite Variation sensed physical data sets for the Río Conchos drainage, we No significant linkage disequilibrium was found among any asked Desktop GARP to predict the distribution in that drain- of the 12 loci examined. All but two populations were in age of any species with similar ecological parameters. As- Hardy-Weinberg equilibrium; the two populations deviat- suming that the Conchos Trout would have habitat require- ing from Hardy-Weinberg equilibrium were those from the ments similar to those oftrout native to the Pacific drainages, Granja Truticola hatchery and the wild population immedi- and most likely rnost similar to the geographically closest ately below the hatchery on Arroyo la Sidra. Factorial cor- other native trout, those from ihe the Yaqui basin, the GARP respondence analysis recovered three groups among the trout model based on input Yaqui trout data, indicated consider- populations examined, one from the northem drainages, ríos able potential trout habitat in the Río Conchos and elsewhere Yaqui and Casas Grandes, one containing the Mexican throughout the Sierra Madre Occidental region (Figure 3). golden trout, and one from the southern drainages (ríos Presidio and San Lorenzo) and al1 hatchery individuals. RESUL TS Variation explained by these groups was low (16.74%), but Distribution and Diversity of Trout in México this was possibly due to smaller sample sizes in the south- As reviewed in Hendrickson et al. (2003) relatively few em drainages adding "noise" to the analysis. AMOVA con- samples of trout from México predated the fonnation of the ducted for these three groups revealed a higher variance binational research and educational team of Truchas between groups than between populations in each group Mexicanas. Through our concerted efforts of intensive re- (29% versus 19%). Within population variance comprised view of the literature and private historic correspondence, 52% of the total variance in the dataset. The high F,, value reconnaissance of maps and areas throughout the northem of 0.4722 indicates significant population substructure. Sierra Madre Occidental, interviews with indigenous people Additional AMOVAs conducted for each of the groups and nine years of surveys we have documented the occur- yielded two groups within the Mexican Golden Trout (O. rence of native trout from the Río Yaqui system south to the clit?~sogaster),one composed of trout from the Río Fuerie

FACULTAD DE CIENCIAS 810~6~1~~5,UANL

CONSERVATIOh OF MEXICAN hAT VE TROUT AND THE OISCOVERY, STATUS, PROTECTlOh AND RECOVERY OF THE CONCHOS TROUT, THE F.RST NATM hatcheries, they are largely uninformative as to the question formative. Mexican Golden Trout (0. chrysogaster) were of more widespread introgression in these systems. We have always embedded within a larger, well-supported clade of collected fish that were morphologically "good" Rainbow samples of the undescribed Mexican trout species. Analysis Trout in streanis in close proximity to both hatcheries, and of partial sequences from O. apache and 0. gil~lealso re- in synipatry with native trout. We have also observed early sulted in those taxa being embedded within these spawning among native populations of trout. Specimens undescribed taxa. Relationships among al1 forms within the collected from the rios Acaponeta, Baluarte, Yaqui and Casas clade inclusive of the Rainbow Trout (0. rnykiss) were not Grandes in early March were in spawning condition; males resolved in either MP or Bayesian analyses, however, some released milt freely upon removal from the water and fe- redband, steelhead and rainbow samples from Bagley and males expelled eggs on light pressure. Size structure of these Gall(1999) were consistently resolved with undescribed taxa populations indicates that reproductive maturity is reached from the Sierra Madre Occidental in al1 analyses. Some dif- earlier and at a smaller size than is typical of most popula- ferentiation among Mexican populations is evident, espe- tions of native fluvial Oncorhj~nchusrnykiss in the United cially among populations in the ríos AcaponetalBaluartel States. These factors may reduce the impact of hybridiza- Piaxtla and San LorenzolPresidio drainages. This suggests tion or introgression by reducing temporal overlap in re- that these populations are native. and have been isolated productive seasonality between Mexican Oncor-hynchus spp. from northern populations of native (not hatchery) 0. rnykiss and exotic 0. rnykiss. This may also produce a skewed di- for a considerable time. The ATPase 618 sequences also re- rectionality of hybridization, resulting from size-assortative vealed relatively low variation; with only 59 parsimony in- pairing, as has been observed in both Rainbow Trout and fonnative characters out of 858 bp. Gila Trout and Apache other species of Oncorhynchus (Foote and Larkin 1988, Trout were recovered as sister to the clade containing na- Dowling and Childs 1992, Rosenfield et al. 2000, Ostberg tive (not hatchery) Rainbow Trout. We did recover similar et al. 7004). While our preliminary data suggest both local groupings of the southern drainages as found in the control hybridization and reductions of variation from bottlenecks, region phylogeny. The nuclear genes CiH 1 C and GH2C were with our sample sizes. additional data are clearly required not informative for variation within clade including Rain- to accurately identify the long-term impact of 0. rnykiss- bow Trout. However, relationships between Gila, Apache, based hatcheries in México. Cutthroat and Rainbow trout were different from those re- covered with mitochondrial genes, especially in the tree re- DNA Sequence Variation covered using GH 1C sequence variation. The genetic varia- While our analysis did yield limited variation in control re- tion of these genes across the native trout of the clade gion sequences from across niuch of the natural range of inclusive of native Rainbow Trout and Gila, Apache, Cut- Oncorh~vnchusrnykiss, this marker is very useful for phylo- throat trout is very promising for ultimately resolving rela- genetic inferences and evaluation of species boundaries. In tionships and species boundaries within these clades. Addi- our preliniinary molecular analysis involving only the con- tional genes (ND2, COI, COII, Cyt-b) and n~icrosatellite trol region. 91 out of 1008 characters were parsimony-in- markers for these and other samples will provide further

FACULTAD DE CIENCIAS BIOL~GICAS,UANL DEAN A. HENDRICKSON, ET AL

resolution of the genetic diversity in these river systems. can trout. Obviously, putting names on these species is of critica1 importance for their conservation given the largely Morphological Variation species-oriented legal protection mechanisms currently in Among the samples examined thus far differences between place in Mexico and elsewhere. thc different proposed undescribed species exist in al1 of the standard morphological features (see inethods) that are di- GIS and GARP Modeling agnostic for up to 10 different species. Results ofthese analy- Because we had evidence from Iiistoric correspondence (re- ses will be presented in a separate publication. viewed in Hendrickson et al. 2003) of native trout in the Rio Conchos basin, and our review of drainage patterns, Taxonomy faunal distributions and habitats ir, the upper Río Conchos Descriptions of what we believe to be distinct native trout indicated that trout could very well exist in these headwa- from the Sierra Madre Occidental of mainland México will ters, we used GARP analysis to predict what streains were be presented elsewhere. However, historically some of these most likely to harbor trout in this system. have formerly (and currently) been grouped under the catch- We input environmental data derived from remotely al1 term of coastal rainbow trout (O. n2,vkiss iridrus). Based sensed imagery coverage of the Sierra Madre Occidental, on molecular analyses, however, they are distinct from what and our known trout localities from the Río Yaqui basin. As has been traditionally referred to as "Rainbow Trout," there is a stochastic element to the GARP process, there is Oncorhynch~ismykis.~, and are more closely related to other no unique solution. Consequently, a number of models are trout species. These taxa are diagnosable on the basis of generated (300 in this case) and a best-subset is chosen. By morphology and molecular traits and such conflation ofthem intersecting the 10 best-subset model convergence map with with the "Rainbow Trout" is an inaccurate depiction of both a stream coverage, streams could be assigned respective the morphological and genetic diversity in México and does model convergence values. These values can then be ranked not accurately depict either their origins or their ongoing to reflect likelihood of native trout populations - based upon evolutionary divergence. Further, we believe that "lump- information about the known localities of native trout in the ing" these trout with O. mykiss will promote the spread of Río Yaqui. Using this algorithm we generated a map of likely hatchery or cultured Rainbow Trout throughout the Sierra locations for trout throughout the Sierra Madre Occidental, Madre Occidental, as was historically done the in US under including the Rio Conchos (Figure 3). The model correctly the polytypic species concept, and consequently endanger identified areas known to contain trout in the remaining parts the gene pools of these native trout. of Pacific-slope drainages, and predicted trout occurrences Therefore, we anticipate describing multiple new taxa in tributaries of the upper Rio Conchos. (of trout as well as minnows and suckers) in the near future Because the terrain of the Sierra Madre Occidental is and providing a revised taxonomy that recognizes and ac- very rugged and travel to collection localities is very time curately reflects the diversity discovered, and the consuming in most areas, we found the GARP predictions phyolgenetic relationships among natural groups of Mexi- to be a useful new tool that helped us more quickly sample LWSERVAT1ON OF MUQG4N NAT1YE TRON AND MEDISCO@RY, STATUS, PR5TECTIOT.IAND RECJOVERY OF THE CONCHQC TROUT, ME FiST NATiVE habitats throughout our study area that were likely to har- the diversity first seen by Lupton. bor native tro~it.Discovery of the Conchos Trout at a local- Our first expedition to the Río Conchos in 2002 (funded ity predicted by GARP to have a high likelihood of harbor- by a grant from the National Science Foundation) focused ing trout (below), together with the high success of the primarily on southern and eastern tributaries but failed to method reciprocally predicting previously sampled locali- produce any native trout populations, despite some appro- ties harboring either undescribed trout or Mexican Golden priate tro~ithabitat. Despite these results and still suspect- Trout based on only a subsample of our known localities ing the existence of an undescribed Conchos Trout, we in- (Fig. xx), exemplifies the power and utility of GARP. put our recent native trout collection localities from other basins into Desktop GARP. Discovery of the Conchos Trout Using the GARP model predictions Truchas Mexiconas Although native Mexican trout have been known to biolo- rapidly focused surveys in 2005 in the streams of the Río gists for well over a century, scientists, as well as the gen- Conchos basin with the highest probabilities of trout occur- eral public, have remained surprisingly ignorant of them. rence. Our extensive efforts thus far have only revealed The tírst Mexican trout collection was made in the 1880's two populations of a trout in the Río Conchos. When the by Nathaniel T. Lupton, Professor of Chemistry at Vanderbilt first population was discovered in 2005 it appeared srnall University. Lupton gave the specimens to Edward Drinker (< 100 individuals) and restricted to a short stream segment Cope, who wrote a brief paragraph about them in the Ameri- (

"aparique," and that most locals. especially Rarámuri living headwaters and a small set of voucher specimens were re- in the Conchos basin, clearly distinguish them from simi- tained for scientific study. Adjacent tributaries also drain- lar-looking introduced, non-native Rainbow Trout (locally ing into the sanle mainstem stream did not maintain any "trucha arco iris"). We have spoken with many local, older populations of trout. Using standard sampling methods for residents who quickly and correctly identified Conchos Trout rapid population evaluation we estimated the population size from a large set of illustrations displayed as an array of simi- in this stream as < 300 adults. While sampling fishes and lar trout and indicated that the species occurred in nearby invertebrates from this stream team members identified three streams (ones we had identified in GARP analyses as high new small dwellings being constructed by Rarámuri, small- probability trout habitats). Unfortunately, after confirming scale timber removal activities in the watershed and evi- that they had seen and captured this species locally, resi- dence of sonle pollution in the stream basin. However, dur- dents almost invariably proceeded to explain that they had ing the sampling period no evidence of local peoples not seen "apariques" for 3 to 10 years, and our sampling of harvesting trout from the stream was encountered through- those streams substantiated their observations. Thorough out the basin. The combination of our own collection ef- reconnaissance and sampling of a wide area of the upper forts and observations by locals has convinced us that not Río Conchos by multiple teams of fiirchas M~J-icanas,and only is this undescribed endemic Mexican trout fading from discussions with local Mexican and Rarámuri revealed mul- local memory, but is also critically endangered and rapidly tiple stream reaches where the "aparí" or "aparique" had disappearing throughout its small remaining range. As can once existed, habitats that were evaluated by our team as be seen from discussions below, our data regarding this dis- excellent for trout, or stream reaches that were dry and had mal conservation situation is not just anecdotal. been dry according to local residents for many years due to Thus, GARP modeling of niches of species in our ef- reduced rainfall and increased pressure for water use. Where forts to sample remote and difficult terrain has been ex- streams were intact, sampling yielded native species of tremely successful and highly valuable for effective and ef- Ccrtostomus. Gila and Codomcr, but no trout. The extirpa- ficient inventory work. The validity of GARP modeling for tion of the Conchos Trout from streams in these areas is Mexican trout is bolstered by the fact that modeling based most likely due to depletion of water in the streams. cli- on subsets of our data from Pacific drainages accurately matic changes in northwestern Mexico with reduced rain- predicted stream reaches containing known samples that fall, overexploitation of these populations as a food source, included the omitted subsets of known trout localities. Va- or a combination of these factors. lidity of extending our model to the Río Conchos basin is During the March 2006 expedition Truchas Mexicanas also bolstered by the fact that the GARP model predicted team members did, however, discover another previously that both streams where we have found the Conchos Trout unknown locality for the undescribed Conchos Trout, in a to have high probabilites of harboring trout. Furthermore. stream reach predicted by GARP analysis to have a high through blind testing by picking from multiple possible na- probability of habitat appropriate for Sierra Madre Occi- tive and cultured trout drawings by Joseph R. Tomelleri ,in- dental trout. This stream was also surveyed from mouth to digenous peoples contirmed existence of the Conchos Trout CONSEHATIUN OF WC-AN NATIVE TROUT AND TtlE MSCQWRY, 5TATUS, PROTECTlON AldD ECQWRWF THE CONCHOS TROUT, THE F1RST MATIVE

in years past in other streams that GARP suggested were disastrous impacts on the population. They did indicate that likely to harbor them.. We thus consider the GARP model they are aware of the presence of apariques and many other predictions to be a reasonable representation of the maxi- regional residents explained to us that locals traditionally mal, theoretical distribution of trout in the Río Conchos ba- tish apariques at Easter. sin. However, various lines of evidence inake it clear to us Following discovery of this population we focused on that human impacts, variables that are near iinpossible to better documenting local habitat conditions via quick pre- model in GARP, have dramatically restricted the area na- litninary studies where the trout lived and in adjacent ar- tive trout now occupy and that protective measures, as well royos that locals and GARP analyses also indicated high as educational efforts are now essential to the continued probabilities of containing trout. We determined that the existence of this and other native trout in the Sierra Madre newly discovered population consists of perhaps 300 indi- Occidental. vidual native trout >lo0 mm total length distributed over approximately 2.5 km of high-gradient stream. Based on Conservation Status of the Conchos Trout the release of eggs by at least one female handled, spawn- This troiit is clearly one of the most imperiled in North ing was occurring at the time of our visit, and our collection America with its current known existence being from only included at least 4 size classes, indicating reasonable and a single stream less than 3 km in length and a population recent recruitment for at least several years. All evidence, size estimated to be less than 300 individuals. Given the ranging from reports by local residents and our own evalu- inferences from sampling multiple locations for trout through ation, indicates that a somewhat larger stream nearby re- the Sierras and the state of other natural trout populations, it cently supported the native aparique. Extensive tish surveys was not surprising that the recently discovered population there during 2005 and 2006, however, failed to produce of the Río Conchos Trout, in a very small stream, was found specimens. We attribute the near absence of apariques in in the least impacted arroyo sampled. This population is the other stream to heavy fishing pressure, as evidenced by constrained to an area near the headwaters in a narrow, high- levers wedged under many boulders in the stream. This tech- gradient canyon with almost no available tillable land. The nique appears to be in widespread use by local residents stream's discharge is exceptionally small (on March 22,2006 throughout the region, is highly effective and capable of from 0.20 - 0.27 liters per second (Lis) near top of trout rapidly decimating small populations already impacted by population to 28 Lis below it), thus limiting irrigation pos- myriad other factors. Cursory examination of invertebrate sibilities. Human population density in the canyon is low; samples from both arroyos with and without apariques sug- only last year was the first house built, iininediately below gest that toxins might be responsible for paucity of apariques the trout population in this arroyo. The new residents of the in the tributary lacking them, an observation that merits fur- canyon are grazing goats and cutting trees to make bowls ther study. Despite having flows that we believed sufficient and other saleable items. While we found no evidence that for supporting trout, no apariques were found in a small they are currently capturing trout. it would be extremely easy tributary immediately below the new house. Seepage of tox- for them to do so in this very small stream, with potentially ins from an abandoned upstream mine or from soils con-

FACULTAD DE CIENCIAS DEAN A. HENDRICKSON, ET AL taminated by niining might explain their absence from tlie Trout is intensive management. We propose declaration of tributary, as well as their absence below the mouth of this a small reserve that includes the entire drainage of the stream tributary. described above. The total area is approxiniately 100 km2. We were told that another small tributary between the The only humans directly affected by such a reserve would two streanis nientioned above never did harbor apariques. be those occupying the new homestead on the stream that and we did not sample it for fishes, but we did visit a small liarbors the population, immediately below the lowermost impoundment ("presita") on it about 200 nl above its mouth. collection of apariques. At a minimum, this faniily should This impoundment, apparently built recently with support be made fully aware of al1 ways in which they currently obtained by the local municipality, had been stocked in Janu- (grazing, tree-cutting), or potentially might (fishing, plow- ary 2006 with 500 fingerling cultured Rainbow Trout. We inglplanting fields, washing clothes or other discharges of have documented many areas in other drainages where Rain- contaminants) inipact the species and should be compen- bow Trout are interacting with natives (via competition and sated for protecting it by changing the ways in which they hybridization) to the detriment of the natives, and these in- use resources. The best solution would probably be to com- troduced trout could easily leave this presita and invade pensate this faniily to move elsewhere, but we realize this native trout habitats. We therefore recommend renloval of might not be possible. Al1 land that we propose to be de- rainbows from this presa as well as the removal of the dan1 clared a reserve is (as far as we can determine) property of so it cannot be re-stocked, and erection of a barrier on the an ejido, and presumably, the family living on the stream lower reaches of the stream harboring apariques to prevent harboring the aparique population is a member ofthat ejido. upstreani niovenient of any non-native fishes. Sonie habi- Following our sampling efforts in this area in March, we tat modifications to this stream to aid the Conchos Trout in revisited the area in ApriI only to find additional impacts on reproduction success, larval development and evade human the native habitat of the trout and new tishing pressures of and non-human predators are also higlily recommended for this snlall population. Empty chlorine bottles were found the continued survival of the species. along the streanl bank and multiple large levers were dis- covered beneath large boulders used by trout for habitat and DISCUSSION concealment from predators. Where these levers were How to Conserve and Protect the Río Conchos Trout present no trout were found in the pools. It is likely that the Members of Truchns Mexicanas are fully committed to con- chlorine was used to kill the trout prior to their removal servation of Conchos Trout. We not only conclude that it is froni pools using the levers to displace boulders. endangered throughout the basin and one known popula- To further justify its protection and managernent, it ap- tion was driven to extinction by local fishing pressure and pears to us that the intrinsic value of this newly discovered habitat rnodifications, but the only viable extant population Conchos Trout stream and the nearby streams we propose we know is critically endangered. protecting goes well beyond having the only known, man- The niost obvious solution for preservation of the only ageable population of Conchos Trout. During our inventory known potentially viable population of the new Conchos efforts we also collected speciniens of a unique, and previ-

5TUDIES OF NQRTH AI4ER1CAN DESERT FlSt-lfS CONSERVATION OF MENCAN NATNE TROUT AND THE DISCOVERY, STATUS, PROTECT Oh AhD RECOVERY OF THE COhCHOS TROLT, ThE FlRST NATNE

ously unknown and undescribed species of sucker (appar- tially could still attract economically significant tourism and ently a relative of Catostomus plebeius) from the stream such a business might one day be possible in the other stream that also harbors the apariques and are in the process of de- where apariques apparently were once more abundant, and scribing this taxon. In addition to these undescribed spe- likely elsewhere, provided that the small surviving aparique cies, the same stream also supports Mexican Stoneroller, population is properly managed and conserved. In the mean- Campostoma ornatum, Ornate Shiner, Codoma 01-nata and time, an ecotourism facility provides convenient housing Conchos Chub, Gila pulchra. Our study of other inverte- for researchers and conservation workers. brates and other organisms was cursory, but al1 of the ar- A barrier on the lower reaches of the stream that harbors royos described above are unusual in the upper Río Conchos the aparique population to protect them from invasions of because they flow to the north, are mostly uninhabited, and Rainbow Trout and the highly likely deleterious impacts on consequently likely harbor a unique assemblage of trees, the new species through introgressive hybridization, con]- forbs and other organisms otherwise uncommon in the Río petition and food depletion is crucial. Truchas Me.~icanas Conchos basin. The adjacent stream that apparently once members committed not only personal contributions of funds harbored apariques has caves with petroglyphs from pre- to establish a base for such an effort, but also committed historic inhabitants along its banks. provision of our experience and expertise toward accom- We view a nearby ecotourism facility as a serendipitously plishing construction of such a barrier. Although other, larger fortunate occurrence for protection and recovery of Conchos efforts are required, especially outreach and education of Trout. The manager, a Spanish-speaking Rarámuri who grew the local community, we believe constniction of a barrier, up in the area and is a member of the ejido that owns the local drainage management efforts with residents, and some streams mentioned above, guided us throughout the area. habitat rehabilitations provide not only tangible protection He is exceptionally knowledgeable regarding local natural that can be quickly accomplished, and which are both sym- history, and interested in the apariques. He introduced us to bolic and educationally useful. These modifications would the owner of a ranch on the lowermost reaches of the stream utilize local labor, thus demonstrating that conservation of that harbors apariques (well below the aparique population), resources, such as aparique, can have tangible and immedi- who explained he has never eaten aparique and is not inter- ate benefits to the local community. ested in them for food. He would th~isseem likely to be The Truchas Mexiconas team is sincerely interested in unaffected by any recovery actions for the trout, but poten- conservation of the Conchos Trout and other trout species tially a beneficiary, together with the owners of an in Mexico. We also believe immediate action to conserve ecotourism business and the ejido (which we understand the small population of the Conchos Trout we discovered is receives part of its income from the tourism business). a critica1 first step toward this end. However, we realize that Ecotourism could be easily extended to include the aparique as biologists, al1 foreign to the local area (and in niany cases and other aspects of the area's natural history as an addi- the country) we are highly ignorant of, and mostly unpre- tional attraction. Though the aparique population is currently pared to deal with the complexities of the local political and too small to allow catch and release fly fishing, it poten- cultural systems through which any actions would have to DEAN A. HENDRICKSON, ET AL.

be taken. We therefore offer whatever assistance we can bow Trout have been introduced into habitats it likely once provide with our diverse and highly relevant backgrounds, occupied, but at least in this case they appear not to have but depend on the participation of others more qualified to established. An incredible diversity of impacts now threaten interact with the appropriate local authorities and who can the tiny remaining population, which, as far as we know help us generate and administer the required funds. may be the last of the species. If not the last population, our We cannot emphasize enough the urgency of taking im- extensive fieldwork leave us confident that not many more mediate steps to save this exceedingly rare and threatened will be found. trout, as well as to forge ahead as quickly as possible to It is clear to us that n~uchremains to be done to assure locate other, similarly imperiled taxa in the Sierra Madre persistence of native Mexican trout and their habitats for Occidental. The impending extinction of this species and future generations. Not only are broad-scale management the decline of the other native trout of the region will affect actions urgently needed, ranging from changing land use Rarámuri tradition as well as other biodiversity. We were practices to reconsidering promotion of Rainbow Trout cul- fortunate to havc located the population we describe, but ture to implementing sweeping socio-economic changes, but the situation requires immediate implementation of recov- we also need additional basic scientific research. System- ery actions and proactive planning for sustainable develop- atic studies are required to understand the evolutionary re- ment efforts and education throughout the region, with a lationships of populations and to allow them to be named "--% vision towards ecotourism capturing a replacement for eco- with appropriate taxonomy Naming these species is essen- nomic growth in the area without the eventual destruction tial for integration of this unique fauna into socio-political of habitats that serve as the primary revenue of the region. conservation frameworks. Additional genetic research is Members of the Truchas Mexicanas organization are pre- required not only for evolutionary studies, but to provide pared to do whatever we personally can to help conserve insights into population biology of the species, and basic this new species, but we clearly need international assis- biological and ecological studies are required so we might tance from citizens, conservation organizations, groups in- understand the complexities of the ecosystems in which this terested in fishing and fishes and their conservation, and biota resides so as to become better informed managers govemment agencies. A Call for International Proactive Efforts The Future of al1 Native Mexican Trout Most native Mexican trout populations are not yet in the The Conchos Trout story provides many lessons. It not only dire condition in which we find the Conchos Trout, but they exemplifies the extreme habitat degradation that is perva- are clearly headed that way. We thus invite al1 those inter- sive and expanding throughout much of our study area, but ested to join the Truchas hfe.siccrncrs team in our efforts to also exemplifies the possible future of many other popula- learn more about these valuable and fascinating fishes, and tions of Mexican trout. As many have told us, the Conchos to join us in helping to assure that the remaining popula- Trout was once much more widely distributed by now finds tions of al1 Mexican trout species persist in their natural refuge only in the uppeimost and tiniest headwaters. Rain- habitats in perpetuity for the benefit of future generations. CONSERVAllDN OF hi(EWS^AN NATIVE TROUT AND THE DISCOERY, STAWS, PROTECTION ANO RECOWRY Df ME CONCHO5 TROUT, THE FIRST NATiVE

lnterested parties and financial contributors wishing to this paper (GRC) on the native trout of Baja California promote the protection and sustainable conservation of the (Oncorl~~vnchusmykiss nelsoni) helped cal1 attention to its Conchos Trout and other new trout species of the Sierra plight arid confer it appropriate legal protection. As a result, Madre Occidental, as well as educational sociological ef- it is today one of the least endangered native Mexican trout. forts with the indigenous Rarámuri (Tarahumara) and Mexi- Finally, Phil's work with the Desert Fishes Council and long- can peoples for existence of their societies in a more envi- standing support of its work in México is testimony to his ronmentally sustainable manner with the local biodiversity commitment to conservation without borders. While we have are encouraged to contact members of the Truchas long been inviting Phil to accompany us on our fieldwork, Mexicanas group andlor authors of this publication, or email unfortunately it has not yet been possible for him to join us, [email protected]~.More information about the but his spirit has always clearly been with us as we wander Truchas Mexicanas project and regarding how others can through the Sierra Madre Occidental searching for Mexi- assist in its efforts and contribute financially to the survival can trout. of these unique ecosystenls can also be found at http:ll It is therefore not only highly appropriate, but also a dis- tinct pleasure, to dedicate this paper to Phil. Hopefully, by following his model. one day the Conchos Trout and al1 other Dedication native Mexican trout will enjoy levels of protection and This paper is dedicated to Edwin "Phil" Pister, long-time conservation security equal to what he has helped attain for leader of the Desert Fishes Council and supporter of con- the species mentioned above. servation efforts for native fishes throughout the arid west of North America. Phil was not only a leader in the famous ACKNO WLEDGMENTS battle for protection of the Devils Hole pupfish (Cyprinodon We thank the Mexican Government for permits to sample diabolis), the population size of which was once similar to the aquatic biodiversity for the past nine years throughout that of the Conchos Trout that this paper highlights, but he the Sierra Madre Occidental (permito of Pesca de Fomento also has extensive experience with trout conservation. His DGOPA 2 10205/0765,25040611606 and 03050611 677). We career-long involvement in management of California's also tharik many of the indigenous peoples of the Sierra golden trout (Oncorhynchus aguabonita .s.spp.)has achieved Madre Occidental, especially those who kindly served as significant legal protection for both them and their habitat, invaluable guides. for their assistance during our expedi- while recovering most of populations from hybridization tions. We thank Jack Williams, Charles Gauvin and Chris with introduced non-native rainbows and the impacts of Woods of Trout Unlirnited for their kind assistance with habitat degradation. These same issues confront conserva- conservation efforts for Conchos Trout, Trout Unlirnited, The tion of al1 native Mexican trout. and it is therefore perhaps Ncrtiiv Fish Conservancy and Mexico Norte for financia1 not surprising that Phil's work extended into that country support of Conchos Trout conservation, and Héctor Arias long before the Truchu.~Mexicrrnas team that authored this and Citlali Cortés Montaño, Oficial de Programa Sierra paper was born. Phil's work with one of the co-authors of Tarahumara, WWF Programa Desierto Chihuahuense, for DEAN A HENDRICKSON, ETAL. their support with community-leve1 conservation actions for spread of Myxobolus cpr-ehralis in the United States. Fisher- the Conchos Trout. ies 22(8): 6-7. Blaustein, A.R. and A. Dobson. 2006. A message from the frogs. Nature 439: 143-144. LITERATURE CITED Blaustein, A.R. and J.M. Kiesecker. 1997. The significance of ul- traviolet-B radiation to amphibian population declines. Rev. Alford, R.A. and S.J. Richards. 1999. Global amphibian declines: Toxicology 1 : 309-327. a problem in applied ecology. Annu. Rev. Ecol. Syst. 30: Blaustein, A.R. and D.B. Wake. 1990. Declining amphibian popu- 133-165. lations - a global phenomenon. Trends Ecol. Evol. 5: 203- Anderson, R.P., D. Lew and A.T. Peterson. 2003. Evaluating pre- 204. dictive models of species' distributions: criteria for selecting Boulton, A. J., W. F. Humphreys and S. M. Eberhard. 2003. Im- optimal models. Ecol. Modeling, 162: 21 1-232. periled Subsurface Waters in Australia: Biodiversity, Threat- Bagley. M.J. and G.A.E. Gall. 1998. Mitochondrial and nuclear ening Processes and C'onservation. Aquatic Ecosystem DNA sequence variability among populatioiis of rainbow Health & Managemeiit 6( 1):4 1-54 trout (0ncor.hjnchus mykiss). Molecular Ecology 7: 945-96 1. Bradford, D.F. 1989. Allotopic distribution of native frogs and Bartholomew, J.L. and J.C. Wilson, eds. 2002. Whirling disease: introduced fishes in high Sierra Nevada lakes of California: reviews and current topics. American Fisheries Society, implication of the negative effect of tish introductions. Copeia Bethesda, Maryland. 1989(3): 775-778. Beebee, T.J.C., R.J. Flower, A.C. Stevenson. S.T. Patrick, P.C. Bradford, D.F., F. Tabatabai and D.M. Graber. 1993. lsolation of Appleby, C. Fletcher, C. Marsh, J. Natkanski, B. Rippey and remaining populations of the native frog, Rana muscosa, by R.W. Battarbee. 1990. Decline of the natterjack toad Bufo introduced fishes in Sequoia and Kings Canyon National calamita in Britain: paleoecological, documentary and ex- Parks, C'alifornia. Cons. Biol. 7: 882-888. perimental evidence for breeding site acidification. Biol. Brach. A.R. and H. Song 2005. ActKey: a Web-based interactive Conserv. 53(1): 1-20. identification key program. Taxon 54(4): 1031-1046 Behnke, R.J. 1992. Native trout of westem North America. Ameri- Brach,A.R. and H. Song. 2006. e-Floras: New direction for online can Fisheries Society Monograph 6. 275 pp. floras exemplified by the Flora of China Project. Taxon 55(1): Behnke. R.J. and J.R. Tomelleri. 2002. Trout and Salmon of North 188-192 America. The Free Press, New York, New York. 384 pp. Briggs, C.J., V.T. Vredenburg, R.A. Knapp and L.J. Rachowiczi. Berger, L., R. Speare, P. Daszak, D.E. Green, A.A. Cunningharn. 2005. lnvestigating the population-leve1 effects of C.L. Goggin, R. Slocombe, M.A. Ragan, A.D. Hyatt, K.R. chytridiomycosis: an emerging infectious disease of amphib- McDonald, H.B. Hines, K.R. Lips, G. Maraiitelli and H. ians. Ecology 86(12): 3 149-3 159. Parkes. 1998. Chytridiomycosis causes amphibian mortality Bronmark, C. and P. Edenhainn. 1994. Does the presence of fish associated with population declines in the rain forests of affect the distribution of tree frogs (Hilla arborea)? Cons. Australia and Central America. PNAS 95: 903 1-9036. Biol. 8: 841-845. Berger, L., R. Speare, H.B. Hines, G, Marantelli,A.D. Hyatt, K.R. Brown, D.E. (Ed.). 1994. Biotic Communities Southwestem United McDonald, L.F. Skerratt. V. Olsen, J.M. Clarke, G. Gillespie, States and Northwestem Mexico. Salt Lake City: University M. Mahony, N. Sheppard, C. Willianis and M.J. Tyle. 2004. of Utah Press. Effect of season and temperature on rnortality in amphibians Bull, E.L. and D.B. Mars. 2002. lnfluence of fish and habitat on due to chytridiomycosis. Aust. Vet. J. 82: 434-439 amphibian coniniunities in high-elevation lakes in iiortheast- Bergersen, E.P. and D.E. Anderson. 1997. The distribution and erii Oregon. Northwest Sci. 76:240-248.

STUDIES OF NORTH AMERICAN COhSERVATION OF MWCAh hATlVE TROLJT ANO THE DISCOVERY, STATUS, PROTECTION AND RECOVERY OF THE CONCHOS TROUT, THE FIRST NATNE

Burr, B.M. 1976. A review ofthe Mexican stoneroller, Campostonza of California Golden Trout based on molecular markers. ornatum Girard (Pisces: Cyprinidae). Trans. San Diego Soc. Trans. Am. Fish. Soc. 135: 110-128. Nat. Hist. 18: 127.144. Coskun, P.E., E. Ruiz-Pesini and D.C. Wallace. 2003. Coritrol re- Burr, B. M. and R. L. Mayden. 1999. A new species of Cyclepflrs gion mtDNA variants: Longevity, climatic adaptation, and a (Cypriniformes: Catostomidae) from Gulf Slope drainages foreiisic conundrum. PNAS 1 OO(5): 2 174-2 176. of Alabama, Mississippi, and Louisiana, with a review of Crabtree, C.B. and D.G. Buth. 1987. Biochemical systematics of distribution, biology and conservation status of the genus. the catostomid genus Catostomus: assessment of C. clarki, Bulletin Alabama Museum of Natural History 20: 19-57. C. plebeius, and C. discobolus including the Zuni Sucker, C. Camarena-Rosales, F. G. Ruiz-Campos, J. De La Rosa-Vélez, R. d. yarrowi. Copeia 1987: 843-854. L. Mayden, D. A. Hendrickson and A. Varela-Romero. 2007. Daszak, P., A.A. Cunningham and A.D. Hyatt. 2003. Infectious Variatioii in mitochondrial haplotypes of wild trout popula- disease and amphibian population declines. Divers. Distrib. tions (Teleostei: Salmonidae) from northwestern Mexico. 9: 141-150. Rev. Fish Biol. Fish. 17(2007):000-000. Davidson, C., H.B. Shaffer and M.R. Jennings. 2002. Spatial tests Carey, C. and M.A. Alexander. 2003. Climate change and am- of the pesticide drift, habitat destruction, UV-B, and climate- phibian declines: is there a link? Divers. Distrib 9(2): 11 1- change hypotheses for California amphibian declines. Coris. 121. Biol. 16(6): 1588- Carpenter. S. R., N. F. Caraco, D. L. Correll, R. W. Howarth, A. Davidson, E. W., M. Parris, J.P. Collins, J.E. Longcore, A.P. Pessier N. Sharpley, V. H. Smith. 1998. Nonpoint Pollution of Sur- and J. Bruner. 2003. Pathogenicity and transmission of

Im=8r"iX""."." ..,, ..rirmz face Waters with Phosphoms aiid Nitrogen. Ecological Ap- chytridiomycosis in tiger salamanders (Ambystoma tigrinum). 81 2&. '*i.i gp$# plications 8(3):559-568. Copeia 2003(3): 601-607. #p&e Ceballos, G., P. Rodriguez and R.A. Medellín. 1998. Assessing Debano. L.F., G.J. Gottfried, R.H. Hamre. C.B. Edminster, P.F. conservation priorities in megadiverse Mexico: mammalian Ffolliott and A. Ortega-Rubio (Eds.). 1994. The Sky Islands diversity, endemicity. and endangerment. Ecol. Appl. 8(1): of southwestern United States and northwestern Mexico. 8-17. USDA Forest Service: General Technical Report RM-GTR- Chang-Sheng Kuoh and H. Song. 2005. lnteractive Key to Tai- 264. wan Grasses Using Characters of LeafAnatomy- The ActKey de Cserna, Z., 1989. An outline of the geology of Mexico. Pp. Approach. Taiwania 50(4): 261-271 233-264 ln: Bally, A.W. and A.R. Palmer (eds.), The Geol- Clarkson, R.W. and J.C. Rorabaugh. 1989. Status of leopard frogs ogy of North Arnerica - An overview. Boulder. Colorado. (Rana pipiens complex: Ranidae) in Arizona and southeast- Geological Society of America. ern California. Southwest. Nat. 34: 531-538. Dorgelo. J. 1987. Density fluctuations in populations (1982- 1986) Collins, J.P. and A. Storfer. 2003. Global anlphibian declines: Sort- and biological observations of Potumopyrgus jenkinsi in two ing the hypotheses. Divers. Distrib. 9: 89-98. trophically differing lakes. Hydrobiol. Bull. 21: 95-110. Contreras-Balderas S. 1975. Zoogeography and evolution of Dowling, T.E. and M.R. Childs. 1992. Impact of hybridization on iVotr.opis lutrensis and Notropis or.r~atlrsin the Rio Grande a threatened trout of Southwesterri United States. Cons. Biol. basin and range. Mexico and United States (Pisces: Cyprin- 6: 355-364. idae) Ph. D. dissertation, Tulane University, New Orleans, Drost, C.A. and G.M. Fellers. 1996. Collapse of a regional frog LA. 146 p. fauna in the Yosemite area of the California Sierra Nevada, Cope, E.D. 1886. The most southern salmon. Am. Nat. 20:735. USA. Cons. Biol. 10: 414-425. Cordes, J.F., M.R. Stephens, M.A. Blumberg and B. May. 2006. Dunham, J.B., D.S. Pilliod and M.K. Young. 2004. Assessing the ldentifying introgressive hybridization in native popiilations consequences of rionnative trout in headwater ecosystems in

FACULTAü DE CIENCIAS BIOL~GICAS,UANL westem North America. Fisheries 29(6): 18-26. Behaviour 106:43-62. Eaton, J.G. and R.M. Scheller. 1996. Effects of climate warming Fuller, P.L., L.G. Nico, J.D. Williams. 1999. Nonindigenous Fishes on fish thermal habitat in streams of the Uriited States. lntroduced into lnland Waters of the United States. Ameri- Limnol. Oceaiiogr. 4 l(5): 1 109- 1 1 15. can Fisheries Society, Bethesda, MD. Einum, S. and 1.A. Flemiiig. 2001. Implicatioris of stockirig: eco- Goldman, S. J., K. Jackson and T. A. Bursztynsky. 1986. Erosioii logical iiiteractions between wild and released salmonids. and sediment control handbook. McGraw-Hill, New York. Nordic J. Freshwater Res. 75: 56-70. Gómez Lepe, C. and M. Sarmiento Frader. De truchas y trucheros. Escalante Pliego, P.. A.G. Navarro S., A. G. and A.T. Peterson. México Desconocido 272, 1999 (available Aug 12, 2006 at 1993. A geographic, historical, and ecological analysis of http://www.mexicodesconocido.com.mx/espanol/ avian diversity iii Mexico. Pp. 281 307 In: The biological culturapypsociedadlactividades~econornicas/ diversity ot' Mexico: origins and distribution (T. P. detalle.~fm?idcat=3&idsec=17&idsub=84&idpag=3 159#). Ramamoorthy, R. Bye, A. Lot and J. Fa, Eds.). Osford Uni- Griffith, J.S., Jr. 1988. Review of competition between cutthroat versity Press, New York. trout and other salmonids. In: R.E. Gresswell (Ed.). Proceed- Ferris, S.D., D.G. Buth and G.S. Whitt. 1982. Substantial gerietic ings of the American Fisheries Society Symposium of the dit'ferentiation among populations of Catostomzrs plebeius. Status and Management of Interior Stocks of Cutthroat Trout, Copeia 1982: 444-449. vol. 4. Bethesda, MD, pp. 134-140. Ferrusquia, 1. 1993. Geology of Mexico: A synopsis. Pp. 3-108 Hale, S.F., P.C. Rosen, J.L. Jarchow and G.A. Bradley. 2005. Ef- In: T. P. Ramamoorthy, R. Bye, A. Lot and J. Fa. (Eds.), fects of the chytrid fungus on the Tarahumara frog (Raria

Biological Diversity of Mexico: Origins and Distribution. tarahumarae) in Arizona and Sonora, Mexico. Pages 407- ~~~~:+~J~~. Oxford: Oxford University Press. 411 IN: G.J. Gottfried, B.S. Gebow, L.G. Eskew and C.B. YA'."'. Fielding, A.H. and Bell, J.F., 1997. A review of methods for the Edminster, editors. Connecting mountain islands and desert ggB assessment of prediction errors in conservation presencel seas: biodiversity and management of the Madrean Archi- absence models. Environ. Conserv. 24: 38-49. pelago 11. Proc. RMRS-P-36. Fort Collins, CO: U.S. Depart- Flecker, A.S. and C.R. Townsend. 1994. Community-wide conse- meiit of Agriculture, Forest Service, Rocky Mountain Re- quences of trout introduction in New Zealand streams. Ecol. search Station. 63 1 pp. Appl. 4: 798-807. Hall. T. 2001. BioEdit v5.0.9. North Carolina State University, Fleischner, T.L. 1994. Ecological costs of livestock grazing in Raleigh, NC. western North America. Cons. Biol. 8(3): 629-644. Hauer. F.R., J.S. Baron. D.H. Carripbell, K.D. Fausch, S.W. Fleming, I.A. and E. Petersson. 2001. The ability ofreleased, hatch- Hostetler, G.H. Leavesley. P.R. Leavitt, D.M. McKnight and ery salmonids to breed and contribute to the natural produc- J.A. Stanford. 1997. Assessment ofclimate change and fresh- tivity of wild populations. Nordic J. Freshwater Res. 75: 7 1 - water ecosystems of the Rocky Mountains, USA and Canada. 98. Hydrological Processes 11 : 903-934. Foote, C.J. 1989. Female mate preference in Pacific salmon. Ani- Hayes, M.P. and M.R. Jennings. 1986. Decline of ranid frog spe- mal Behaviour 38: 721 -722. cies in western North America: are bullfrogs (Raria Foote, C.J. 1990. An experimental comparison of male and fe- catesbeiaria) responsible? J. Herpetol. 20(4):490-509. iiiale spawning territoriality in a Pacific salmon. Behaviour Hendrickson, D.A., W.L, Minckley, R.R. Miller, D.J. Siebert and 1 15: 283-3 14. P.L. Haddock. 1980. Fishes of the Río Yaqui, México and Foote. C.J. and P.A. Larkin. 1988. the role of male choice in as- the United States. J. Ariz. Nev. Acad. Sci. 15: 65-106. sortative mating of sockeye salmon arid kokariee, the anadro- Hendrickson D.A., H. Espinosa-Pérez, L.T. Findley, W. Forbes, moiis aiid non-anadromous fortiis of Or~corllyrichusrierka. J.R. TomeIleri, R.L. Mayden, J.L.Nielsen, B. Jeiisen, G. Ruiz-

STUDfES OF NORTH AMERíCAN DESERT FISHEC( CQNSERVATON OF MBICAN HATIVETRQUT AND THE DlSCQVfRY, 5STAIU5, PROTECTfQPs AND ECOVERY OF THE CONCHO5 TROUT, THE FIRST NATlVE

Campos, A. Varela-Romero, A.M.Van Der Heiden, F. Chinchar, J.P. Collins, B.L. Jacobs and A. Storfer. 2005. Camarena and F.J. García de León. 2003. Mexican native Evidence for emergence of an amphibian iridoviral disease trouts: a review of tlieir history and current systematic and because of human- enhanced spread. Mol. Ecol. 14(1): 2 13- conservation status. Rev. Fish Biol. Fisli. 12(2002)(4): 273- 224 316. Jarvie, J. K. & Stevens. P. F. 1998. ínteractive keys. inventory, Herbst, D.B., E.L. Silldorff and S.D. Cooper. 2003. The influence and conservation. Cons. Biol. 12: 222-224 of introduced trout on native aquatic invertebrate communi- Justice, C.O., E. Vermote, J.R.G. Townshend, R. Defries, D.P. Roy, ties in a paired watershed study of High Sierran streams. D.K. Hall, V.V. Salomonson, J.L. Privette, G. Riggs, A. University of California Water Resources Center. Technical Straliler, W. Lucht, R.B. Myneni. Y. Knyazikhin, Y., Run- Completion Reports. http://repositories.cdlib.org/wrc/tcr/ ning. S.W., Nemani, R.R., N'an, Z., Huete, A.R.. van lierbst Leeuwen, W., Wolfe, R.E., Giglio, L., Muller, J.-P., Lewis, Hilderbrand, R.H. and J.L. Kershner. 2000. Conserving inland P. and Barnsley, M.J., 1998. The moderate resolution imag- cutthroat trout in small streams: how mucli stream is enough? ing Spectroradiometer (MODIS): Land Remote Serising for North Am. J. Fish. Mgmt. 20: 51 3-520. Global Change Research. IEEE Trans. Geosci. Remote Sens., Hitt, N.P., C.A. Frissell, C.C. Muhlfeld and F.W. Allendorf. 2003. 36: 1228-1 249. Spread of hybridization between native westslope cutthroat Kats, L.B., J.W. Petranka and A. Sih. 1988. Antipredator defenses trout, Oncovhynchus clnvki lewisi, and nonnative rainbow and the persistence of amphibian larvae with fiches. Ecol- trout, Oncovh.vnchtis mykiss. Can. J. Fish. Aq. Sci. 60: 1440- ogy 69: 1865- 1 870. 1451. Kiesecker, J.M., A.R. Blaustein and C.L. Miller. 2001. Transfer Holland, J.H. 1975. Adaptation in natural and artificial systems. ol'a pathogen from fish to arriphibians. Cons. Biol. 15(4): The University of Michigan Press. Anii Arbor, MI. 1064-1 070. Houlahan, J.E., C.S. Findlay, B.R. Schmidt, A.H. Meyer and S.L. Kiesecker, J.M., A.R. Blaustein and L.K. Belden. 2001. Complex Kuzmin. 2000. Quantitative evidence for global amphibian causes of amphibian population declines. Nature 4 10: 68 1- population declines. Nature 404(6779): 752-755. 684. Huelsenbeck, J. P. and F. Ronquist. 200 1. MR BAYES: Bayesian Kiesecker, J.M.. A.R. Blaustein. 1995. Synergisrri between UV-B inference of phylogeny. Bioinformatics 17: 754- 755. radiatioii and a pathogen magiiifies amphibian einbryo mor- Huete, A., C. Justice and W. van Leeuwen. 1999. Modis Vegeta- tality in nature. PNAS 92: 1 1049-1 1052. tion lndex (MOD13). Algorithm theoretical basis document. Kiesecker. J.M., A.R. Blaustein. 1998. Effects of introduced bull- http://tbrs.arizona.edu/project/MOD1S/index.php frogs and smallmouth bass ori rriicrohabitat use, growtli and Huete, A., K. Didan, T. Miura, E.P. Rodriguez, X. Gao and L.G. survival of native red-legged frogs (Rann ntivova). Cons. Biol. Ferreira. 2002. Overview of the radiometric and biophysical 12: 776-787. perforniarice of the MODIS vegetatiori iridices. Remote Sens. Knapp, R.A. 2005. Effects of nonnative fish and habitat charac- Eiiviron. 83: 195-2 1 3. teristics on lentic herpetofauna in Yoserriite National Park, Hulbert, S.H. and A. Villalobos-Figueroa (Eds.). 1982. Aquatic USA. Biol. Cons. 121(2): 265-279. Biota of México, Central America and the West Indies. San Knapp, R.A. and K.R. Matthews. 1996. Livestock grazing, golde11 Diego State University, San Diego, California. 529 pp trout, and streams in the Golden Trout Wilderness, Califor- Jager, H.I., W. Van Winkle and B.D. Holcomb. 1999. Would hy- nia: Irripacts and managemerit irnplications. N. Am. J. Fish. drologic climate changes in Sierra Nevada streams influence Manage. 16: 805-820. trout persistence? Trans. Am. Fish. Soc. 128: 222-240 Kontula, T., S. Kirilchik and R. Vainola. 2003. Endemic diversifi- Jancovich. J.K., E.W. Davidson, N. Pararrieswaran, J. Mao, V.G. cation of the monophyletic cottoid fish species flock in Lake Baikal explored with mtDNA sequencing. Molecular concerns, and management reconirriendations. Pages 269-282 Phylogenetics and Evolution 27: 143-155. In: de Lourdes Lozano Vilano, M. and A.J. Contreras Kruse, C.G., W.A. Hubert and F.J. Rahel. 2001. As assessment of Balderas, eds. 2004. Homenaje al Doctor Andrés Reséndez headwater isolation as a conservation strategy for cutthroat Medina, un ictiólogico mexicano. Universidad Autónoma de trout in the Absaroka Mountains of Wyoming. Northwest Sci. Nuevo León. Monterrey, Mexico. 316 pp. 75:l-11. Mayden, R.L. and R.M. Wood. 1995. Systematics, species con- Laurance, W. F., K.R. McDonald and R. Speare. 1996. Epidemic cepts, and the evolutionarily signficant unit in biodiversity disease and the catastrophic decline of Australian rain forest and conservation hiology. Special Publication No. 17. Am. frogs. Cons. Biol. 10: 406-413. Fish. Soc., Bethesda, Maryland. Pp. 58-1 13. Leider, S.A.. P.L. Hulett, J.J. Loch and M.W. Chilcote. 1990. Elec- McGinnity, P., P. Prodohl, A. Ferguson, R. Hynes, N.O. trophoretic comparison of tlie reproductive success of natu- Maoileidigh, N. Baker, D. Cotter, B. O'Hea, D. Cooke, G. rally spawning transplanted and wild steelhead trout through Rogan, J. Taggart and T. Cross. 2003 Fitness reduction and the returning adult stage. Aquaculture 88: 239-252. potential extinction of wild populations of Atlantic salmon, Leopold, A. 1949. A Sand County Almanac. Oxford University Solrno salar as a result of interactions with escaped farrn Press, New York. salmon. Proc. Biol. Sci. 270(1532): 2443-2445 Li, H.W., J.L. Li and J.C. Buckhouse. 1994. Cumulative effects of Mclntosh, A.R. and C.R. Townsend. 1996. lnteractions between riparian disturbances along high desert trout streams of the fish, grazing invertehrates and algae in a New Zealand strearn: John Day Basin, Oregon. Trans Am. Fish. Soc. 123: 627- a trophic cascade mediated by fish-induced clianges to grazer 640. behaviour. Oecologia 108: 174-1 81. Lydeard, C. and R. L. Mayden. 1995. A diverse and endangered McLean, J.E., P. Bentzen and T.P. Quinn. 2004. Differential re- aquatic ecosystem of the southeast United States. Cons. Biol. productive success of sympatric, naturally-spawning hatch- 9(4):800-805. ery and wild steelhead trout, Oncorhynchus mykiss. Environ. Mao, J., D.E. Green, G. Fellers and V.G. Chinchar. 1999. Molecu- Biol. Fish. 69: 359-369. lar characterization of iridoviruses isolated from sympatric Meisner, J.D. 1990. Effect of clirnatic wamling on the southern amphihians and fish. Virus Res. 63: 45-52. margins of the native range of brook trout, Salvelinus Matthews, K.R., K.L. Pope, H.K. Pressler and R.A. Knapp. 2001. ,fonrii~alis.Can. J. Fish. Aq. Sci. 47(6): 1065-1070 Effects of nonnative trout on Pacific treefrogs (Hyla regilla) Miller, R.R.. W.L. Minckley and S.M. Norris. 2005. Freshwater in the Sierra Nevada. Copeia 200 l(4): 1130-1 137. Fishes of México. University of Chicago Press. 652 pp. Mayden, R.L. 1997. A hierarchy of species concepts: tlie denoue- Mittermeier, R.A.. da Fonseca, G.A.B., Hoffman. M., Pilgrim, J., ment in the saga of the species problem, p. 381-424. 111: Spe- Brooks. T.. Robles Gil, P., Mittermeier, C.G. and Lamoreux, cies: The Units of Biodiversity. Claridge, M.F., Dawah, H.A. J. 2004. tlotspots Revisited: Earth's Biologically Richest and and Wilson, M.R. (eds). Chapman and Hall Ltd., London. Most Endangered Terrestrial Ecoregions. CEMEX, Conser- Mayden. R.L. 1999. Consilience and a hierarchy of species con- vation International, Agrupación Sierra Madre, University cepts: Advances towards closure on the species puzzle. J. of Virginia, Washington, D.C. Nematol. 3 1:95-116. Navarro, A. G. and Benitez-Diaz. H. 1993. Patrones deriqueza y Mayden, R.L. 2002. On biological species, species concepts, and endemismo de las aves. Ciencias 7: 45-54. individuation in the natural world. Fish and Fisheries. 2: 1- Navarro-Sigüenzal, A.G. and A.T. Peterson. 2004. An alternative 26. species taxonomy of the birds of Mexico. Biota Neotropica Mayden, R.L. 2004. Biodiversity of Mexican trout (Teleostei: 4(2): 1-32, Salmonidae: OIICO~~.IIC~LI.S):recent findings. conservation Needham, P.R. and Gard, R. 1959. Rainbow trout in Mexico and

STUDIES OF NORTH AMERfCAN DESERT FISHES CONSERVATION OF MEXiCAN hATlM TROUT AND THE DISCOVERY, STATUS, PROTECTlOh AND RECOVERY OF THE CONCHOS TROUT, THE FlRST NATNE

Califomia with notes on ttie cutttiroat series. Univ. Calif. Publ. Parrriesan, C. and Galbraith, H. 2004. Obsewed lmpacts of Glo- Zool. 67(1): 1-124. bal Climate Change in the U.S., Peu Center on Global Cli- Nielsen, J.L., M.C. Fountain, J.C. Favela, K. Cobble and B.L. mate Change, Arlington, Virginia, 67 pp. Jensen. 200 1. Oncorhynchus at the southem extent of their Peckarsky, B.L., A.R. Mclntosh, B.W. Taylor and J. Dahl. 2002. range: a study of mtDNA control-region sequence with spe- Predator chemicals induce changes in mayfly life history cial reference to an undescribed subspecies of 0.mykis.5 from traits: a whole-stream manipulation. Ecology 83: 612-618. Mexico. Environ. Biol. Fish. 51(1): 7-23. Peterson, A.T. and Robins, C.R. 2003. Using ecological-niche Nielsen, J.L. and G.K. Sage. 2001. Microsatellite analysis of the modeling to predict barred owl invasions with implications trout of northwest Mexico. Genetica 11 1 : 269-278. for spotted owl conservation. Cons. Biol., 17: 1161-1 165. Norris, S.M., J.M. Fischer and and W.L. Minckley. 2003. Gila Phillips R.B., M.P. Matsuoka, N.R. Konkol and S. McKay. 2004. brevicauda (Teleostei: Cyprinidae), a new species of fish from Molecular systematics and evolution of the growth hormone the Sierra Madre Occidental of México. Ichthyol. Explor. introns in the Salmoninae. Environmental Biology of Fishes Freshwaters 14(1): 19-30 69: 433-440. Novinger, D.C. and F.J. Raliel. 2003. Isolation management with Piotrowski, J.S., S.L. Annis and J.E. Longcore. 2004. Physiology artificial barriers as a conservation strategy for cutthroat trout of Batrachochytrium dendroba~idis,a cliytrid pathogen of in headwater streams. Cons. Biol. 17: 772-781. amphibians. Mycologia 96: 9-15 O'Neal. K. (2002) Effects of Global Warming on Trout and Salmon Pounds, J.A., M.R. Bustaniante, L.A. Coloma, J.A. Consuegra, in U.S. Streams. Defenders of Wildlife and National Re- M.P.L. Fogden, P.N. Foster, E. La Marca, K.L. Masters. A. . .""".. ".,."m,... m"ms.s.... sources Defense Council, Washington, D.C., 46 pp. Merino-Viteri, R. Puschendorf, S.R. Ron, C.A. Sánchez- g, 8 ., 'S, Ortega-S. C. Montes De Oca RM, Groman D, Yason C, Nicholson Azofeifa, C.J. Still and B.E. Young. 2006. Widespread am- B. Blake S. 2002. Case Report: viral infectious pancreatic phibian extinctions from epidemic disease driven by global necrosis in farmed Rainbow Trout frorri Mexico. J. Aqua. warming. Nature 439(7073): 161-167. Anim. Health 14(4): 305-1 0. Propst, D.L., J.A. Stefferud and P.R. Turner. 1992. Consewation Ostberg, C.O., S.L. Slatton and R.J. Rodriguez. 2004. Spatial par- and status of Gila trout, Oncorh).nch~rsgilae. Southwest. Nat. titioning and asymmetric Iiybridization among sympatric 37(2): 117-125. coastal steelhead trout (Oncorhynchus rnykiss irideus), coastal Quinn, T.P. and C.J. Foote. 1994. The effects of body size and cutthroat trout (O. clar-ki clarki) and interspecific hybrids. sexual dimorphism on the reproductive behaviour of sock- Mol. Ecol. 13(9): 2773-2788. eye salmon, Oncorhynchus nerka. Anirrial Behaviour 48: 75 1- Osterkamp, W. R., P. Heilman and L. J. Lane. 1998. Economic 761. considerations of a continental sediment-monitoring program. Rahel, F.J.: C.J. Keleher and J.L. Anderson. 1996. Potential habi- Inter. J. Sed. Res. 13(4):12-24. tat loss and population fragmentation for cold water fish in Parmesan. C., N. Ryrholm, C. Stefanescu, J. K. Hill. C. D. Tho- the North Platte River drainage of the Rocky Mountains: mas, H. Descimon, B. Huntley, L. Kaila, J. Kullberg, T. response to climate warming on the properties of boreal lakes Tarrirriani, W. J. Tennent, J. A. Thomas, and M. Warren. 1999. and streams at tlie Experimental; Lakes Area, Ontario. Poleward shifts in geographical ranges of butterfly species Limnol. Oceanogr. 4 1 : 1 1 16- 1 123. associated with regional warming. flature 399 (6736):579- Ramamoorthy, T.P.. R. Bye, A. Lot, and J. Fa. 1998. Diversidad 583. Biológica de México, México, D.F.: Instituto de Biología, Parmesan, C. and G. Yohe. 2003. A globally coherent fingerprint Universidad Nacional Autónoma de México. of climate change impacts across natural systems. Nature Raymond, M. and F. Rousset. 1995. Genepop (version 1.2), popu- 421 (691 8):37-42. lation genetics software for exact tests and ecumenicism. J.

FACULT4D DE CIENCIA! BIOLOGICAC,UANL Hered. 86: 248-249. software for population genetics data analysis. http:l/ Reeder, A.L., M.O. Ruiz, A. Pessier, L.E. Brown, J.M. Levengood. anthro.unige.ch1arlequin. C.A. Phillips, M.B. Wheeler, R.E. Warner and V.R. Beasley. Siebert, D.J. and W.L. Minckley. 1986. Two new catostomid fishes 2004. lntersexuality and the cricket frog decline: historic and (Cypriniformes) from the northern Sierra Madre Occidental geographic trends. Env. Health Perspect. 1 13(3): 26 1-265. of Mexico. Am. Mus. Novit. 2849: 1 - 17 Relyea, R.A. 2004. The lethal impacts of Roundup and predatory Simon, K.S. and C.R. Townsend. 2003. The impacts of freshwater stress on six species of North American tadpoles. Archiv. invaders at different levels of ecological organisation, with Environ. Cont. Tox. 48(3): 351-357. emphasis on salmonids and ecosystem consequences. Fresh- Reno P. 1999. Infectious pancreatic necrosis and associated aquatic water Biol. 48: 982-994. birnaviruses. In: Fish Diseases and Disorders, Vol. 3 (ed. by Snyder, D.E. 2003. Electrofishing and Its Harmful Effects on Fish, P.T.K. Woo & D.W. Bruno), pp. 1-55. CAB lnternational Information and Technology Report USGSlBRDlITR-2003- Ltd, Wallingford. 0002, Denver. Co1orado:U.S. Government Printing Office. Retallick, R.W.R., H. McCallum and R. Speare. 2004. Endemic 149 pages. infection of the amphibian chytrid fungus in a frog commu- Steen, R.P. and T.P. Quinn. 1999. Egg burial depth by sockeye nity post-decline. PLoS Biol. 2: 1966-1 97 1. salmon: implications for survival of embryos and natural Reyes Valdez. C.A. 2005. Análisis Alimentario Comparativo de selection on female body size. Can. J. Zool. 77: 836-841. Truchas Nativas del Género Oncorhylichus (Teleostei: Stockwell, D.R.B. and Noble, 1.R. 1992. Induction of sets of rules Salmonidae), de la Sierra Madre Occidental, en el Noroeste from animal distribution data: a robust and informative de México. Unpubl. M.S. Thesis, Universidad Autonoma method of analysis. Math. Comp. Simul., 33: 385-390. de Baja California. Ensenada, Baja California, México. Stockwell, D. and Peters, D. 1999. The GARP modelling system: Rinne, J.N. 1990. Status, distribution, and conservation of two problems and solutions to automated spatial prediction. Int. rare southwesterti (U.S.A.) salmonids, the Apache trout, J. Geogr. Inf. Sci.. 13: 143-158. O~irorhyr~ch~rstrptrche Miller and the Gila trout, 0. gilae Strand, M. and R.W. Merritt. 1999. Impacts of livestock grazing Miller. J. Fish Biol. 37: 189-191. activities on streatri insect cotritriutiities and the riverine en- Ron, S.R. 2005. Predicting the distributioti ofthe amphibian patho- vironment. Ani. Entomol. 45(1): 13-29. gen Batrachochytrium dendrobatidis in the New World. Swofford, D.L. 2000. PAUP. Phylogenetic analysis using parsi- Biotropica 37: 209-22 1. mony (and other methods). Version 4.0b8. Sinauer Associ- Ron, S.R., W.E. Duellman, L.A. Coloma and M. Bustamante. 2003. ates, Sunderland, MA. Population decline of the Jambato toad Ate1opu.s igncscc~ls Taylor, J.N., W.R. Courtenay and J.A. McCann. 1984. Known (Anura: Bufonidae) in the Andes of Ecuador. J. Herpetol. impacts of exotic fishes in the continental United States. Pp 37: 116-126. 322-373 1N: W.R. Courtenay and J.R. Stauffer, Jr., Eds. Dis- Rosenfield, J.A., T. Todd and R. Greil. 2000. Asymmetric hybrid- tribution, biology, and management of exotic fishes. John ization and introgression between pink salmon and Chinook Hopkins Press, Baltimore, Maryland. salmon in the Laurentian Great Lakes. Trans. Am. Fish. Soc. Tyler, T., W.J. Liss, L.M. Ganio, G.L. Larson, R. Hoffman, E. 129: 670-679. Deimling and G. Lomnicky. 1998. Interactions between in- Ruiz-Campos G, Camarena-Rosales F, Varela-Romero A, Sancliez- troduced trout and larval salamanders (Amhy.stoma Gonzales S, de la Rosa-Velez J. 2003. Morphonietric varia- macrodactylum) in high-elevation lakes. Cons. Biol. 12: 94- tion of wild trout populations from northwesterti Mexico 105. (Pisces: Salmonidae). Rev. Fish Biol. Fish. 13(1): 91 -1 10. U.S. Drug Enforcement Agency. 200 1. Cannabis Cultivation Den- Schneider, S., D. Roessli and L. Excoftier. 2000. Arlequin v2.0. A sity from Mexico - Countrj Brief, DEA-01002 (available

STUDlES OF NORTH AMERICAN DESERT FISHES CONSEWAJION OF MEXILAN NATM TROUT AND THf DISCOYERY, STATUS, PRO'FEUiOM AND RECavERY O? THE CONCHO5 TRDUT, THE flRST NATIVE

Aug. 12. 2006 at http://wwu.lib.utexas.eduhnaps/americas/ 860 me~ico~cannabis-densitym.giQ Wake. D.B. and J.J. Morowitz. 1991. Declining amphibian popu- U.S. Drug Enforcement Agency. 2001. Opium Poppy Cultivation lations a global phenomenon? Findings and recommenda- Density from Mexico - Country Brief, DEA-01002 (avail- tions. Alytes 9: 33-42. able Aug. 12, 2006 at http://www.Iib.utexas.edu/n1aps1 Weigel, D.A., J.T. Peterson and P. Spruell. 2003. Introgressive americas/mexicogoppy_cultivation_200 I .giQ hybridization between native cutthroat trout and introduced U.S. Fish and Wildlife Service. 2002. 2001 National Suwey of rainbow trout. Ecol. Appl. 13: 38-50. Fishing, Hunting, and Wildlife-Associated Recreation. Wash- Welsh, H.H., K.L. Pope and D. Boiano. 2006. Sub-alpine amphib- ington, D.C. 170pp. http://www.census.goviprod/2002pubc/ ian distributions related to species palatability to non-native FHWOI .pdf salinonids in the Klamath mountains of northern California. van den Berghe, E.P. and M.R. Gross. 1989. Natural selection re- Divers. Distrib. 12(3): 298-309. sulting from female breeding competition in a Pacific salmon Wilcove, D. S., D. Rothstein, J. Dubow, A. Phillips, E. Losos. (Oncorhynchus kisutch). Evolution 43: 125- 140. 1998. Quantifying Threats to Imperiled Species in the United Villalobos-Figueroa, A. 1983. Crayfishes of México (Crustacea: States. BioScience 48 (8):607-615 Decapoda). Smithsonian lnstitution Librarles and the Na- Williams, J. D., M. L. Warren Jr., K. S. Cumrriings, J. L. tiarris tional Science Foundation. Amerind Publishing, New Delhi. and R. J. Neves. 1993. Conservation Status of Freshwater 276 pp. Mussels of the United States and Canada 1993. Fisheries Vredenburg, V.T. 2004. Reversing introduced species effects: ex- 18(9):6-22 >7.v8r"e,. -.->.. perimental reinoval of introduced tish leads to rapid recov- Zander, R.H. 2004. Minimal values for reliability of bootstrap and "=>-=iP"<".".&;1.!; $~+; $~+; "a, *il;.: ery of a declining frog. PNAS lOl(20): 7646-7650. jackknife proportions. decay index, and Bayesian posterior & Wake, D.B. 1991. Declining amphibian populations. Science 253: probability. Phylolnformatics 2: 1-13.

FACULTAD DE C1ENCiAS BIOLÓGICAS,UANL DEAN A I-IINDRICKSON, f7 Ai

Fig~irel. General distributions ofnalive Mexican troot (Oncoi*l?~iiichusspp.) with irnages of diffcrent distinct foi-rns. Trout from top to bottorn include forins from the rios Casas Grandes, Yaqui, Mayo, Conchos, O chr-ysogtrsler, San Lorenzo, Piaxla, Baluarte and Acaponeta. Fish images copyrighted by Joseph R. Tornelleri; base map adapted from Behnke and Tomelleri (2002).

STUDIES OF NORTH AMERICAN DESFRr FISHES ONSERVATIQN~OF-MWCAN NATIVE TROUT AND THE DISCOVERY, STATUS,. PROTECTION AND RECOVERY OF THE CONCHOS TRQUT,.THE FIRST NATIVE ......

Solid Red Areas Are Elevation Areas 1,828 m (6,000 ft.) in Sierra Madre Occidental

Figure 2. Distribution of high-elevation areas in Sonora, Chihuahua and Durango, Mexico, showing areas likely to contain isolated populations of native troot. Figure 3. CARP rnap of the Sierra Madre Occidental with streain seginents scoring a perfect fit with the model higlilighted in green. Black circles represent the known Yaq~iiTrolit sites used for rnodel development.