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We provide an updated checklist of birds of a unique landscape in northeastern Argentina, which is characterized by three isolated, rocky outcrops and their surrounding agroecosystem. We recorded 188 bird species, including nine that are globally or nationally threatened. We highlight the presence of several grassland-specialist birds of conservation concern. Of the species recorded, 80.6% are residents and 17.7% are migrants. The heterogeneity of the landscape and its structural complexity accounts the existence of a high avian species diversity, which includes both generalists and specialists. This study confirms the conservation value of this ecosystem, due both to the rocky outcrops and grassland matrix. Grasslands are one of the most threatened biomes in the world. Fil: Fernández, Juan Manuel. Universidad Nacional del Nordeste. Facultad de Ciencias Exactas, Naturales y Agrimensura. Departamento de Biología; Argentina. Consejo Nacional de Investigaciones Científicas y Técnicas. Centro Científico Tecnológico Conicet - Nordeste; Argentina Fil: Thomann, Maria Luz. Universidad Nacional del Nordeste. Facultad de Ciencias Exactas, Naturales y Agrimensura. Departamento de Biología; Argentina. Consejo Nacional de Investigaciones Científicas y Técnicas. Centro Científico Tecnológico Conicet - Nordeste; Argentina Fil: Fandiño, Blas. Dirección General de Sistema Provincial de Áreas Naturales Protegidas; Argentina Fil: Cajade, Rodrigo. Universidad Nacional del Nordeste. Facultad de Ciencias Exactas, Naturales y Agrimensura. Departamento de Biología; Argentina. Consejo Nacional de Investigaciones Científicas y Técnicas. Centro Científico Tecnológico Conicet - Nordeste; Argentina Fil: Hernando, Alejandra. Universidad Nacional del Nordeste. Facultad de Ciencias Exactas, Naturales y Agrimensura. Departamento de Biología; Argentina

Isolated rocky outcrops represent biodiversity centers, refuges for endangered species, and favorable scenarios for endemism. Most studies in these ecosystems have been focused mainly on their flora and secondarily on different groups of animals. To highlight the value of rocky outcrops as ecosystems for biodiversity conservation, in the present study, we describe the diversity of herptiles of three isolated rocky outcrops of northeastern Argentina and compare it with that of other natural areas of the region. We conducted fieldwork from September 2010 to March 2017. We calculated the alpha diversity and the number of rare, threatened and endemic species. For comparative diversity analysis, we calculated the importance of each area for amphibians and reptiles and the beta diversity. Were recorded a total of 56 species (23 amphibians and 33 reptiles), representing 35% of the herptiles recorded for Corrientes province. These species included 19 rare species, seven threatened species, and two endemic species. The overall beta diversity showed considerably differences in species composition between the compared areas. The rocky outcrops showed higher importance for amphibians and reptiles than the other areas studied. Our study contributes to the knowledge of rocky outcrops and highlights their importance in biodiversity conservation. Fil: Piñeiro, Jose Miguel. Universidad Nacional del Nordeste. Facultad de Ciencias Exactas, Naturales y Agrimensura. Departamento de Biología. Laboratorio de Herpetología; Argentina. Consejo Nacional de Investigaciones Científicas y Técnicas. Centro Científico Tecnológico Conicet - Nordeste; Argentina Fil: Cajade, Rodrigo. Universidad Nacional del Nordeste. Facultad de Ciencias Exactas, Naturales y Agrimensura. Departamento de Biología. Laboratorio de Herpetología; Argentina. Consejo Nacional de Investigaciones Científicas y Técnicas. Centro Científico Tecnológico Conicet - Nordeste; Argentina Fil: Hernando, Alejandra Beatriz. Universidad Nacional del Nordeste. Facultad de Ciencias Exactas, Naturales y Agrimensura. Departamento de Biología. Laboratorio de Herpetología; Argentina Fil: Courtis, Azul Celeste. Universidad Nacional del Nordeste. Facultad de Ciencias Exactas, Naturales y Agrimensura. Departamento de Biología. Laboratorio de Herpetología; Argentina. Consejo Nacional de Investigaciones Científicas y Técnicas. Centro Científico Tecnológico Conicet - Nordeste; Argentina Fil: Ingaramo, María del Rosario. Universidad Nacional del Nordeste. Facultad de Ciencias Exactas, Naturales y Agrimensura. Departamento de Biología. Laboratorio de Herpetología; Argentina. Consejo Nacional de Investigaciones Científicas y Técnicas. Centro Científico Tecnológico Conicet - Nordeste; Argentina Fil: Marangoni, Federico. Universidad Nacional del Nordeste. Facultad de Ciencias Exactas, Naturales y Agrimensura. Departamento de Biología; Argentina. Consejo Nacional de Investigaciones Científicas y Técnicas. Centro Científico Tecnológico Conicet - Nordeste; Argentina

The composition of a snake assemblage from an orographic island in east-central Argentina, the Sierras de Ventania mountain chain, was analyzed. The aim was to determine the biogeographic resemblance to other snake assemblages from neighboring regions. Species composition of each region was obtained from an exhaustive review of the literature, and both fieldwork and museum records. The higher biogeographic resemblance of the Sierras de Ventania occurred with the Sierras de Tandilia and the Coastal Dunes. These regions formed a well-defined group according to their snake assemblages. On the other hand, the Sierras de Lihué Calel linked to the Sierras de Ventania, and also to the rest of the compared regions, at very low values of biogeographic resemblance. The results obtained in this study contrasted with the classic zoogeographic scheme. Snake assemblages allowed recognizing a more significant division between Central and Pampean domains. In this scheme, the limit between these two regions moved to the southwest of the classical scheme; therefore the Sierras de Ventania was part of the Pampean domain. Also, the recognition of the Subtropical domain was evident, as well as its faunistic link with the Pampean domain. Fil: Di Pietro, Diego Omar. Universidad Nacional de La Plata. Facultad de Ciencias Naturales y Museo. Departamento Científico Zoología Vertebrados; Argentina. Consejo Nacional de Investigaciones Científicas y Técnicas; Argentina Fil: Cabrera, Mario Roberto. Consejo Nacional de Investigaciones Científicas y Técnicas. Centro Científico Tecnológico Conicet - Córdoba. Instituto de Diversidad y Ecología Animal. Universidad Nacional de Córdoba. Facultad de Ciencias Exactas Físicas y Naturales. Instituto de Diversidad y Ecología Animal; Argentina Fil: Williams, Jorge Daniel. Universidad Nacional de La Plata. Facultad de Ciencias Naturales y Museo. Departamento Científico Zoología Vertebrados; Argentina Fil: Alcalde, Leandro. Consejo Nacional de Investigaciones Científicas y Técnicas. Centro Científico Tecnológico Conicet - La Plata. Instituto de Limnología "Dr. Raúl A. Ringuelet". Universidad Nacional de La Plata. Facultad de Ciencias Naturales y Museo. Instituto de Limnología; Argentina Fil: Cajade, Rodrigo. Universidad Nacional del Nordeste. Facultad de Ciencias Exactas, Naturales y Agrimensura. Departamento de Biología. Laboratorio de Herpetología; Argentina. Consejo Nacional de Investigaciones Científicas y Técnicas; Argentina Fil: Kacoliris, Federico Pablo. Universidad Nacional de La Plata. Facultad de Ciencias Naturales y Museo; Argentina. Universidad Nacional de La Plata. Facultad de Ciencias Naturales y Museo. Departamento Científico Zoología Vertebrados; Argentina

Introduction: Rocky outcrops influence both micro and macro habitat for plants and animals by increasing the availability and predictability of food, providing shelter, and unique microclimatic conditions. Objective: We describe the bat assemblage in three isolated rocky outcrops and their surrounding pediments, adding new data about trophic guilds and roosts. Methods: We surveyed the bat fauna of Paraje Tres Cerros, Corrientes Province, Argentina, exploring 13 sites during 26 field trips, using mist nets and search of roosts. We made an accumulation species curve and calculate inventory completeness. The similarity and composition of different habitats was compared by a Non-Metric Multidimensional Scaling (NMDS) and an ANOSIM test. Results: We reported 13 species, with a completeness inventory of the 80 % (Jacknife1), corresponding to the families Vespertilionidae, Molossidae and Phyllostomidae. The insectivorous bats were the best represented guild. We found nine types of roost for eight species in different sites. The analysis expressed low differences in the species composition between both types of habitats in the area. Conclusions: The rocky outcrops and their pediments are important ecosystems for the conservation and diversity of the bats in the region, since their assemblages are regionally unique. Resumen Introducción: Los afloramientos rocosos influyen tanto en el micro como en el macro hábitat de plantas y animales al incrementar la disponibilidad de alimentos, proporcionar refugios y brindar condiciones microclimáticas únicas. Objetivo: Describir el ensamble de murciélagos en tres afloramientos rocosos aislados y sus pendientes circundantes, incorporando nuevos datos sobre gremios tróficos y refugios. Métodos: Muestreamos la fauna de murciélagos del Paraje Tres Cerros, provincia de Corrientes, Argentina, exploramos 13 sitios durante 26 viajes de campo, usando redes de niebla y búsqueda de refugios. Realizamos una curva de acumulación de especies y calculamos la completitud del inventario. La similitud y composición de diferentes hábitats se comparó mediante un escalado multidimensional no métrico (NMDS) y una prueba ANOSIM. Resultados: Reportamos 13 especies, con una completitud del 80 % del inventario (Jacknife1), correspondientes a las familias Vespertilionidae, Molossidae y Phyllostomidae. Los gremios mejor representados fueron los insectívoros. Encontramos nueve tipos de refugios para ocho especies en diferentes sitios. Los análisis expresaron bajas diferencias en la composición de especies entre ambos tipos de hábitats en el área. Conclusión: Los afloramientos rocosos y pendientes son ecosistemas importantes para la conservación y diversidad de los murciélagos en la región, ya que sus comunidades son regionalmente únicas.

Background: Argenteohyla siemersi (red-spotted Argentina frog) is a casque-headed tree frog species belonging to the Hylidae family. This species has a complex combination of anti-predator defense mechanisms that include a highly lethal skin secretion. However, biochemical composition and biological effects of this secretion have not yet been studied. Methods: The A. siemersi skin secretion samples were analyzed by mass spectrometry and chromatographic analysis (MALDI-TOF/MS, RP-HPLC and GC-MS). Proteins were also studied by SDS-PAGE. Among the biological activities evaluated, several enzymatic activities (hemolytic, phospholipase A2 , clotting, proteolytic and amidolytic) were assessed. Furthermore, the cytotoxic activity (cytolysis and fluorescence staining) was evaluated on myoblasts of the C2C12 cell line. Results: The MALDI-TOF/MS analysis identified polypeptides and proteins in the aqueous solution of A. siemersi skin secretion. SDS-PAGE revealed the presence of proteins with molecular masses from 15 to 55 kDa. Steroids, but no alkaloids or peptides (less than 5 kDa), were detected using mass spectrometry. Skin secretion revealed the presence of lipids in methanolic extract, as analyzed by CG-MS. This secretion showed hemolytic and phospholipase A2 activities, but was devoid of amidolytic, proteolytic or clotting activities. Moreover, dose-dependent cytotoxicity in cultured C2C12 myoblasts of the skin secretion was demonstrated. Morphological analysis, quantification of lactate dehydrogenase release and fluorescence staining indicated that the cell death triggered by this secretion involved necrosis. Conclusions: Results presented herein evidence the biochemical composition and biological effects of A. siemersi skin secretion and contribute to the knowledge on the defense mechanisms of casque-headed frogs. Fil: Fusco, Luciano Sebastian. Consejo Nacional de Investigaciones Científicas y Técnicas. Centro Científico Tecnológico Conicet - Nordeste. Instituto de Química Básica y Aplicada del Nordeste Argentino. Universidad Nacional del Nordeste. Facultad de Ciencias Exactas Naturales y Agrimensura. Instituto de Química Básica y Aplicada del Nordeste Argentino; Argentina Fil: Cajade, Rodrigo. Universidad Nacional del Nordeste. Facultad de Ciencias Exactas, Naturales y Agrimensura. Departamento de Biología. Laboratorio de Herpetología; Argentina. Consejo Nacional de Investigaciones Científicas y Técnicas; Argentina Fil: Piñeiro, Jose Miguel. Universidad Nacional del Nordeste. Facultad de Ciencias Exactas, Naturales y Agrimensura. Departamento de Biología. Laboratorio de Herpetología; Argentina. Consejo Nacional de Investigaciones Científicas y Técnicas; Argentina Fil: Torres, Ana M.. Universidad Nacional del Nordeste; Argentina Fil: da Silva, Igor R. F.. Universidade Estadual de Campinas; Brasil Fil: Hyslop, Stephen. Universidade Estadual de Campinas; Brasil Fil: Leiva, Laura Cristina Ana. Consejo Nacional de Investigaciones Científicas y Técnicas. Centro Científico Tecnológico Conicet - Nordeste. Instituto de Química Básica y Aplicada del Nordeste Argentino. Universidad Nacional del Nordeste. Facultad de Ciencias Exactas Naturales y Agrimensura. Instituto de Química Básica y Aplicada del Nordeste Argentino; Argentina Fil: Pimenta, Daniel C.. Governo do Estado de Sao Paulo. Secretaria da Saude. Instituto Butantan; Brasil Fil: Bustillo, Soledad. Consejo Nacional de Investigaciones Científicas y Técnicas. Centro Científico Tecnológico Conicet - Nordeste. Instituto de Química Básica y Aplicada del Nordeste Argentino. Universidad Nacional del Nordeste. Facultad de Ciencias Exactas Naturales y Agrimensura. Instituto de Química Básica y Aplicada del Nordeste Argentino; Argentina

Made available in DSpace on 2020-12-10T19:54:02Z (GMT). No. of bitstreams: 0 Previous issue date: 2020-03-27 Fundação de Amparo à Pesquisa do Estado de São Paulo (FAPESP) Conselho Nacional de Desenvolvimento Científico e Tecnológico (CNPq) Rede Baiana de Pesquisa sobre Anfibios (RBPA) Fundacao O Boticario de Protecao a Natureza Fundacao de Amparo a Pesquisa da Bahia (FAPESB) Coordenação de Aperfeiçoamento de Pessoal de Nível Superior (CAPES) Mohamed bin Zayed Species Conservation Fund Fundação de Amparo à Pesquisa do Estado do Rio de Janeiro (FAPERJ) Smithsonian Institution (USA) Agencia Nacional de Promocion Cientifica y Tecnologica (ANPCyT) The South American and West Indian Casque-headed Treefrogs (Hylidae: Hylinae: Lophyohylini) include 85 species. These are notably diverse in morphology (e.g. disparate levels of cranial hyperossification) and life history (e.g. different reproductive modes, chemical defences), have a wide distribution, and occupy habitats from the tropical rainforests to semiarid scrubland. In this paper, we present a phylogenetic analysis of this hylid tribe based on sequence fragments of up to five mitochondrial (12S, 16S, ND1, COI, Cytb) and six nuclear genes (POMC, RAG-1, RHOD, SIAH, TNS3, TYR). We included most of its species (> 96%), in addition to a number of new species. Our results indicate: (i) the paraphyly of Trachycephalus with respect to Aparasphenodon venezolanus; (ii) the nonmonophyly of Aparasphenodon, with Argenteohyla siemersi, Corythomantis galeata and Nyctimantis rugiceps nested within it, and Ap. venezolanus nested within Trachycephalus; (iii) the polyphyly of Corythomantis; (iv) the nonmonophyly of the recognized species groups of Phyllodytes; and (v) a pervasive low support for the deep relationships among the major clades of Lophyohylini, including C. greeningi and the monotypic genera Itapotihyla and Phytotriades. To remedy the nonmonophyly of Aparasphenodon, Corythomantis, and Trachycephalus, we redefined Nyctimantis to include Aparasphenodon (with the exception of Ap. venezolanus, which we transferred to Trachycephalus), Argenteohyla, and C. galeata. Additionally, our results indicate the need for taxonomic work in the following clades: (i) Trachycephalus dibernardoi and Tr. imitatrix; (ii) Tr. atlas, Tr. mambaiensis and Tr. nigromaculatus; and (iii) Phyllodytes. On the basis of our phylogenetic results, we analyzed the evolution of skull hyperossification and reproductive biology, with emphasis on the multiple independent origins of phytotelm breeding, in the context of Anura. We also analyzed the inter-related aspects of chemical defences, venom delivery, phragmotic behaviour, co-ossification, and prevention of evaporative water loss. Univ Estadual Paulista, Inst Biociencias, Dept Biodiversidade, Av 24A 1515, BR-13506900 Rio Claro, SP, Brazil Univ Estadual Paulista, Inst Biociencias, Ctr Aquicultura, Av 24A 1515, BR-13506900 Rio Claro, SP, Brazil Univ Sao Paulo, Inst Biociencias, Dept Zool, BR-05508090 Sao Paulo, SP, Brazil Univ Fed Rio de Janeiro, Museu Nacl, Dept Vertebrados, Setor Herpetol, BR-20940040 Rio De Janeiro, RJ, Brazil Univ Estadual Santa Cruz, Dept Ciencias Biol, Trop Herpetol Lab, Rodovia Jorge Amado,Km 16, BR-45662900 Ilheus, BA, Brazil Univ Brasilia, Dept Engn Florestal, Lab Fauna & Unidades Conservacao, BR-70910900 Brasilia, DF, Brazil Univ Fed Vicosa, Dept Biol Anim, Museu Zool Joao Moojen, BR-36570900 Vicosa, MG, Brazil Univ West Indies, Zool Museum, Dept Life Sci, St Augustine, Trinidad Tobago Pontificia Univ Catolica Rio Grande do Sul, Lab Sistemat Vertebrados, Av Ipiranga 6681,Predio 40,Sala 110, BR-90619900 Porto Alegre, RS, Brazil Consejo Nacl Invest Cient & Tecn, UNaM, Inst Biol Subtrop, Lab Genet Evolut Claudio Juan Bidau, Felix Azara 1552,CPA N3300LQF Posadas, Misiones, Argentina Univ Estadual Paulista, Inst Biociencias, Lab Herpetol, Campus Litoral Paulista, BR-11330900 Sao Vicente, SP, Brazil Univ Nacl Nordeste, CONICET, Fac Ciencias Exactas & Nat & Agrimensura, Lab Herpetol,Dept Biol, Av Libertad 5470, RA-3400 Corrientes, Argentina Consejo Nacl Invest Cient & Tecn, Fdn Miguel Lillo, Unidad Ejecutora Lillo, Miguel Lillo 251, RA-4000 San Miguel De Tucuman, Tucuman, Argentina Univ Koblenz Landau, Inst Integrated Sci, Dept Biol, Univ Str 1, D-56070 Koblenz, Germany Univ Fed Rural Rio De Janeiro, Inst Biol, Dept Biol Anim, Caixa Postal 74524, BR-23851970 Seropedica, RJ, Brazil Consejo Nacl Invest Cient & Tecn, Museo Argentino Ciencias Nat Bernardino Rivadavia, Div Herpetol, Angel Gallardo 470,C1405DJR, Buenos Aires, DF, Argentina Univ Buenos Aires, Fac Ciencias Exactas & Nat, Dept Biodiversidad & Biol Expt, Buenos Aires, DF, Argentina Univ Estadual Paulista, Inst Biociencias, Dept Biodiversidade, Av 24A 1515, BR-13506900 Rio Claro, SP, Brazil Univ Estadual Paulista, Inst Biociencias, Ctr Aquicultura, Av 24A 1515, BR-13506900 Rio Claro, SP, Brazil Univ Estadual Paulista, Inst Biociencias, Lab Herpetol, Campus Litoral Paulista, BR-11330900 Sao Vicente, SP, Brazil FAPESP: 2013/20423-3 FAPESP: 2017/26162-8 FAPESP: 2003/10335-8 FAPESP: 2011/50146-6 FAPESP: 2018/15425-0 FAPESP: 2012/10000-5 FAPESP: 2013/50741-7 FAPESP: 2007/57067-9 FAPESP: 2012/12500-5 CNPq: 167387/20170 CNPq: 155198/2018-1 CNPq: 476789/2009-5 CNPq: 306823/2017-9 CNPq: 140412/2008-5 CNPq: 150377/2012-6 CNPq: 150122/2014-4 Fundacao O Boticario de Protecao a Natureza: 0991_20132 Fundacao de Amparo a Pesquisa da Bahia (FAPESB): 8951/2014 CAPES: 001 CAPES: 88881.133927/2016-01 CNPq: 140716/2016-5 FAPESP: 2014/50342-8 Agencia Nacional de Promocion Cientifica y Tecnologica (ANPCyT): PICT 2013-404 Agencia Nacional de Promocion Cientifica y Tecnologica (ANPCyT): 2015-820

La quiropterofauna de la provincia de Corrientes es una de las menos conocidas de la Argentina. En este trabajo se reportan ocho nuevas localidades de distribución para ocho especies de murciélagos, pertenecientes a las familias Phyllostomidae, Vespertilionidae y Molossidae. Además, se agregan reportes adicionales de especies nuevas para localidades de registro ya conocidas. Estos registros corresponden a 31 ejemplares procedentes de distintas ecorregiones (Chaco Húmedo, Campos y Malezales y Esteros del Iberá). Finalmente, se incluyen algunos comentarios sobre la historia natural de las especies citadas, ampliando el conocimiento para este grupo de mamíferos en la provincia. The chiropterofauna of Corrientes province is one of the least known of Argentina. In this contribution, eight new localities are reported for eight species, belonging to the families Phyllostomidae, Vespertilionidae and Molossidae. In addition, we added some of those species for localities already known for other species. All these records correspond to 31 specimens from different ecoregions (Chaco Húmedo, Campos y Malezales, and Esteros del Iberá). Finally, we add some comments on the natural history of the mentioned species, expanding the knowledge for this group of mammals in the Province. Fil: Argoitia, Antonella. Universidad Nacional del Nordeste. Facultad de Ciencias Exactas y Naturales y Agrimensura; Argentina Fil: Cajade, Rodrigo. Universidad Nacional del Nordeste. Facultad de Ciencias Exactas y Naturales y Agrimensura; Argentina. Consejo Nacional de Investigaciones Científicas y Técnicas. Centro Científico Tecnológico Conicet - Nordeste; Argentina Fil: Piñeiro, Jose Miguel. Universidad Nacional del Nordeste. Facultad de Ciencias Exactas y Naturales y Agrimensura; Argentina. Consejo Nacional de Investigaciones Científicas y Técnicas. Centro Científico Tecnológico Conicet - Nordeste; Argentina Fil: Teta, Pablo Vicente. Consejo Nacional de Investigaciones Científicas y Técnicas. Oficina de Coordinación Administrativa Parque Centenario. Museo Argentino de Ciencias Naturales "Bernardino Rivadavia"; Argentina

We studied size, age and growth in Chacophrys pierottii, a fossorial amphibian inhabiting the arid region of Gran Chaco, in Northern Argentina. We provide the first detailed demographic data for this cryptic species using skeletochronology. We observed a female-biased sexual size dimorphism, but no differences in age parameters between the two sexes. C. pierottii has a short life span of up to five years, both males and females reaching sexual maturity after one year. Our results suggest that the rapid growth to maturity, followed by a relatively short reproductive lifespan, is a response to the constraints posed by arid environments. The life-history parameters in the studied population of C. pierottii are in line with the observed patterns of other anurans inhabiting xeric habitats. Our study provides conservation managers with the basic data required for the accurate assessment of the species status and the adoption of proper conservation strategies. Fil: Marangoni, Federico. Consejo Nacional de Investigaciones Científicas y Técnicas. Centro Científico Tecnológico Conicet - Nordeste; Argentina. Universidad Nacional del Nordeste; Argentina Fil: Stănescu, Florina. Universitatea Ovidius Din Constanta; Rumania Fil: Courtis, Azul Celeste. Universidad Nacional del Nordeste; Argentina. Consejo Nacional de Investigaciones Científicas y Técnicas. Centro Científico Tecnológico Conicet - Nordeste; Argentina Fil: Piñeiro, Jose Miguel. Consejo Nacional de Investigaciones Científicas y Técnicas. Centro Científico Tecnológico Conicet - Nordeste; Argentina. Fundación Amado Bonpland; Argentina. Universidad Nacional del Nordeste; Argentina Fil: Ingaramo, María del Rosario. Consejo Nacional de Investigaciones Científicas y Técnicas. Centro Científico Tecnológico Conicet - Nordeste; Argentina. Universidad Nacional del Nordeste; Argentina Fil: Cajade, Rodrigo. Consejo Nacional de Investigaciones Científicas y Técnicas. Centro Científico Tecnológico Conicet - Nordeste; Argentina. Universidad Nacional del Nordeste; Argentina. Fundación Amado Bonpland; Argentina Fil: Cogălniceanu, Dan. Universitatea Ovidius Din Constanta; Rumania

La Historia Natural (HN) es una disciplina relevante de las Ciencias Naturales que se encarga de describir a los seres vivos y su ambiente a partir de la observación precisa. Como madre de las ciencias occidentales, sus orígenes se remontan a los escritos de Aristóteles, para atravesar desde entonces la historia de la humanidad brindando conocimientos esenciales para su desarrollo. Luego de su auge entre los siglos XVII y XIX con los viajes exploratorios de los naturalistas europeos a los nuevos continentes, la HN ha perdido valoración en el aporte a la construcción del conocimiento científico. Desde entonces el debate sobre el rol de la HN en las ciencias ha oscilado como un péndulo entre su estimación y desestimación. En esta contribución, presentamos los principales temas de este debate aportando nuestro punto de vista a partir de nuestra experiencia en la revaloración de la historia de la HN, resignificada con la influencia de la obra de Alexander von Humboldt y Amado Bonpland. La especialización de las ciencias, los avances tecnológicos, la pérdida del sentido naturalista y la vinculación con el romanticismo han sido, entre otras, las principales causas que han desacreditado a la HN. Sin embargo, concluimos que la historia nos muestra que la HN ha sido y es fundamental para el crecimiento y desarrollo de la humanidad.

We analyzed the spatial distribution of a snake assemblage from the Sierras de Ventania mountain range, a rich endemism area in east-central Argentina. We used field-collected and museum specimens combined with high-resolution environmental variables to identify common distributional patterns, estimate the suitable habitat for each species, and develop a conservation proposal related to the Protected Natural Areas (PNAs) system. Distributional models diverged into two opposite patterns: suitable habitats for most species at relatively high altitude in the hills, and suitable habitats for Bothrops ammodytoides and Lygophis anomalus at relatively low altitude in the surrounding hill plain. We assessed two proposals based on different conservation algorithms: the additive benefit function (ABF), which favours hilly environments, prioritizing areas with high specific richness, and the core-area zonation (CAZ), which prioritizes the distribution of all species, further indicating areas with low richness in the surrounding plain. Regardless of the algorithm used, our results indicated that the existing PNAs are not effective in protecting the snake assemblage. Protecting at least 5% of the highest species richness areas (ABF) or 5% of the distribution core areas of all species (CAZ), the existing PNAs only represented 2.2% of these priority areas. Our findings are largely consistent with previous work, highlighting that a widespread area needs to be assigned for conservation purposes. Thus, protection of the snake assemblage should take place both inside and outside the PNAs. Fil: Di Pietro, Diego Omar. Universidad Nacional de La Plata. Facultad de Ciencias Naturales y Museo. División Zoología de Vertebrados. Sección Herpetología; Argentina Fil: Cabrera, Mario Roberto. Consejo Nacional de Investigaciones Científicas y Técnicas. Centro Científico Tecnológico Conicet - Córdoba. Instituto de Diversidad y Ecología Animal. Universidad Nacional de Córdoba. Facultad de Ciencias Exactas Físicas y Naturales. Instituto de Diversidad y Ecología Animal; Argentina Fil: Williams, Jorge Daniel. Universidad Nacional de La Plata. Facultad de Ciencias Naturales y Museo. División Zoología de Vertebrados. Sección Herpetología; Argentina Fil: Kacoliris, Federico Pablo. Universidad Nacional de La Plata. Facultad de Ciencias Naturales y Museo. División Zoología de Vertebrados. Sección Herpetología; Argentina Fil: Cajade, Rodrigo. Universidad Nacional del Nordeste. Facultad de Ciencias Exactas, Naturales y Agrimensura. Departamento de Biología. Laboratorio de Herpetología; Argentina Fil: Alcalde, Leandro. Consejo Nacional de Investigaciones Científicas y Técnicas. Centro Científico Tecnológico Conicet - La Plata. Instituto de Limnología "Dr. Raúl A. Ringuelet". Universidad Nacional de La Plata. Facultad de Ciencias Naturales y Museo. Instituto de Limnología; Argentina

Two dimensions of the ecological niche (diet and habitat) of a snake assemblage from an endemic rich area in east-central Argentina, the Sierras de Ventania mountain chain, were analyzed. Field data collection was performed in 15-week study periods between 2010 and 2014. Snakes were hand-captured using transect surveys. Field observations on diet were analyzed together with stomach content data from museum specimens. Our results supported the partitioning of the snake assemblage by both habitat use and diet into at least three functional groups: species restricted to microhabitats under rocks and with a diet composed exclusively of ants (Epictia australis); species found mostly in stream microhabitats and feeding mainly upon anurans (Erythrolamprus poecilogyrus and Lygophis elegantissimus); and species found mostly in grassland microhabitats, with specialized diets of terrestrial prey items (Philodryas patagoniensis and Bothrops alternatus). Consistent with previous work, diet was more important than habitat in explaining ecological niche partitioning of this snake assemblage. Our results showed that high overlap values of microhabitat use were compensated by low overlap values of the trophic niche dimension, thus matching the traditional complementary niches hypothesis., Facultad de Ciencias Naturales y Museo, Instituto de Limnología "Dr. Raúl A. Ringuelet"

Fil: Idoeta, Fabricio Miguel. Universidad Nacional de La Plata Facultad de Ciencias Naturales y Museo; Argentina. Fil: Cajade, Rodrigo. Universidad Nacional del Nordeste. Facultad de Ciencias Exactas y Naturales y Agrimensura; Argentina. Fil: Piñeiro, José Miguel. Nacional del Nordeste. Facultad de Ciencias Exactas y Naturales Y Agrimensura; Argentina. Fil: Acosta, José Luis. Universidad Nacional del Nordeste. Facultad de Ciencias Exactas y Naturales y Agrimensura; Argentina. Fil: Pautasso, Andrés A. Nacional del Nordeste. Facultad de Ciencias Exactas y Naturales y Agrimensura; Argentina. Fil: Pautasso, Andrés A. Museo Provincial de Ciencias Naturales Florentino; Argentina. Se presenta el primer registro de Eumops bonariensis (Chiroptera, Molossidae) para la provincia de Corrientes, Argentina. Se capturó un macho adulto con red de neblina en el cerro “Capará”, ubicado en la Estancia Las Marías, Paraje Tres Cerros, Departamento General San Martín. Con esta adición esta provincia incrementa la riqueza específica de quirópteros a 29 especies. We report the first record of Eumops bonariensis (Chiroptera, Molossidae) for the province of Corrientes, Argentina. The specimen was a male collected at Paraje Tres Cerros, Department General San Martín. This record increases the richness of bats in the province to 29 species.

ResumenLos afloramientos rocosos aislados se describen como paisajes destacados en biodiversidad debido a la gran variedad de microhabitats, presencia de taxones restringidos y al alto recambio de especies con la matriz circundante. Particularmente los afloramientos rocosos aislados del Paraje Tres Cerros (ARPTC), Corrientes, Argentina, representan un paisaje único en la región, biológicamente fueron poco explorados. Se estudió la comunidad de aves de bosques y pastizales de los ARPTC mediante 136 listas de Mackinnon de diez especies durante la temporada reproductiva y no reproductiva. Se evaluó la diversidad, riqueza estimada y completitud de inventarios, abundancia de gremios tróficos y fenología de las especies. Se comparó estos atributos entre los bosques y pastizales. Se registraron 107 especies donde el bosque es 1.3 veces más diverso que el pastizal. La completitud de inventarios varió de 83 a 90 % en el pastizal y de 88 a 99 % en el bosque. Los hábitats se complementaron en un 62 % indicando una elevada ocurrencia de especies exclusivas. Las aves insectívoras fueron abundantes en ambos ambientes, mientras que las frugívoras en bosques y las granívoras en pastizales. Ambos hábitats son significativamente más ricos en especies en la estación reproductiva por la llegada de los migrantes estivales. Los resultados concuerdan con los patrones generales conocidos para las comunidades de aves y con otros específicos para las comunidades de aves de afloramientos rocosos. El presente estudio resalta la importancia de los afloramientos rocosos aislados de Paraje Tres Cerros para la protección de la biodiversidad de aves a nivel local y regional y representa una herramienta de manejo para la conservación de estos ecosistemas. AbstractIsolated rocky outcrops can act as patch habitats that host great biodiversity through the diversity of microhabitats, presence of restricted taxa and the high turnover of species in the patch matrix. The isolated rocky outcrops of Paraje Tres Cerros (ARPTC), Corrientes, Argentina, are a unique landscape in the region and biologically little explored. We conducted a study to describe for the first time the bird community of the forest and grassland of the ARPTC, using 136 lists following the ten species Mackinnon method from the breeding and non-breeding seasons. We evaluated the diversity, trophic guilds and phenology of the species, and compared attributes of grassland and forest communities. The total species richness was 107, with the forest being 1.3 times more diverse than the grassland. Completeness of inventories ranged from 83-90 % in the grassland and 88 to 99 % in the forest. The habitat complementarity was 62 %, indicating the presence of several species exclusive of forest and grassland habitats. The trophic structure of communities showed a good representation of insectivorous birds in both habitats. Frugivorous birds were more frequent in the forest and granivorous birds were more frequent in the grasslands. Due to the arrival of migrant species, both habitats had significantly higher richness in the breeding season than in the non-breeding season. Our results showed similar general aspects and patterns for bird communities and with specific studies for bird communities of rocky outcrops. The present study highlights the importance of the isolated rocky outcrops of Paraje Tres Cerros for the protection of local and regional bird biodiversity and represents a benchmark for the management and the protection of these ecosystems.

Anurans employ a wide variety of anti-predator mechanisms to defend themselves. In casque-headed hylids, defence is thought to be a complex combination of several anti-predator mechanisms. However, the defence traits of only a few species are known; some hypotheses have yet to be addressed, whereas others, already tested in some species, need to be tested in additional taxa. The anti-predator mechanism of the casque-headed frog, Argenteohyla siemersi, is described here. It is a complex mechanism consisting of (1) behavioural and ecological traits, including secretive and semi-phragmotic habits and posture; (2) morphological features, including cryptic and aposematic colourations, a skull covered with bony dermal spines and protuberances that are associated with two types of granular venom glands; and (3) physiological and chemical traits, such as a highly lethal skin secretion. Our results are compared with those of previous studies of defence mechanisms in casque-headed frogs in an effort to understand the mechanisms and evaluate their potential phylogenetic signal in this group of anurans., Instituto de Limnología "Dr. Raúl A. Ringuelet"

Tityus curupi n. sp., belonging to the bolivianus complex, is described from the biogeographically distinct area of Paraje Tres Cerros in north-eastern Argentina. We also present a molecular species delimitation analysis between Tityus curupi n. sp. and its sister species Tityus uruguayensis Borelli 1901 to confirm species integrity. Furthermore, a cytogenetic analysis is presented for these two species which contain different multivalent associations in meiosis, as a consequence of chromosome rearrangements, and the highest chromosome numbers in the genus. Fil: Ojanguren Affilastro, Andres Alejandro. Consejo Nacional de Investigaciones Científicas y Técnicas. Oficina de Coordinación Administrativa Parque Centenario. Museo Argentino de Ciencias Naturales “Bernardino Rivadavia”; Argentina Fil: Adilardi, Renzo Sebastián. Consejo Nacional de Investigaciones Científicas y Técnicas. Oficina de Coordinación Administrativa Ciudad Universitaria. Instituto de Ecología, Genética y Evolución de Buenos Aires. Universidad de Buenos Aires. Facultad de Ciencias Exactas y Naturales. Instituto de Ecología, Genética y Evolución de Buenos Aires; Argentina Fil: Cajade, Rodrigo. Universidad Nacional del Nordeste. Facultad de Ciencias Exactas y Naturales y Agrimensura; Argentina. Consejo Nacional de Investigaciones Científicas y Técnicas; Argentina Fil: Ramirez, Martin Javier. Consejo Nacional de Investigaciones Científicas y Técnicas. Oficina de Coordinación Administrativa Ciudad Universitaria. Instituto de Ecología, Genética y Evolución de Buenos Aires. Universidad de Buenos Aires. Facultad de Ciencias Exactas y Naturales. Instituto de Ecología, Genética y Evolución de Buenos Aires; Argentina Fil: Ceccarelli, Fadia Sara. Consejo Nacional de Investigaciones Científicas y Técnicas. Oficina de Coordinación Administrativa Parque Centenario. Museo Argentino de Ciencias Naturales “Bernardino Rivadavia”; Argentina Fil: Mola, Liliana Maria. Consejo Nacional de Investigaciones Científicas y Técnicas. Oficina de Coordinación Administrativa Ciudad Universitaria. Instituto de Ecología, Genética y Evolución de Buenos Aires. Universidad de Buenos Aires. Facultad de Ciencias Exactas y Naturales. Instituto de Ecología, Genética y Evolución de Buenos Aires; Argentina

In this study we analysed the body size, age structure, age at maturity, longevity and growth pattern of Argenteohyla siemersi pederseni in north-eastern Argentina using skeletochronological methods. Body size was sexually dimorphic; females were significantly larger and heavier than males, regardless of age. As A. s. pederseni is an explosive breeder with only one reproductive episode per year, we assumed that each Line of Arrested Growth (LAG) observed was equivalent to one period of decreasing growth, LAGs reflecting a direct estimation of individual age. The minimum number of LAGs counted was two for males and three for females, but there was no sexual difference in longevity. The von Bertalanffy's growth coefficients (K) were higher in males than in females for both size and body mass. Females had a lower growth rate than males, took longer to reach the minimum maturation size, and were older and larger at reproduction. Fil: Cajade, Rodrigo. Universidad Nacional del Nordeste. Facultad de Ciencias Exactas Naturales y Agrimensura. Departamento de Biología. Laboratorio de Herpetología; Argentina. Consejo Nacional de Investigaciones Científicas y Técnicas; Argentina Fil: Marangoni, Federico. Universidad Nacional de Misiones. Facultad de Ciencias Exactas, Químicas y Naturales; Argentina. Consejo Nacional de Investigaciones Científicas y Técnicas; Argentina Fil: Gangenova, Elena. Universidad Nacional de Misiones. Facultad de Ciencias Exactas, Químicas y Naturales; Argentina. Consejo Nacional de Investigaciones Científicas y Técnicas; Argentina

Natale, G.S., Alcalde, L., Herrera, R., Cajade, R., Schaefer, E.F., Marangoni, F. and Trudeau, V.L. 2011. Underwater acoustic communication in the macrophagic carnivorous larvae of Ceratophrys ornata (Anura: Ceratophryidae). —Acta Zoologica (Stockholm) 92: 46–53. We provide the first evidence for sound production by anuran larvae. In this study, we describe the sounds, their context-specific emission and the structures related to sound production of the carnivorous larvae of Ceratophrys ornata (Amphibia, Anura, Ceratophryidae). Tadpoles emit a brief, clear and very audible metallic-like sound that consists of a short train of notes that occur at all stages of larval development. Tadpoles make sound only when a conspecific tadpole is preying upon it or when touched by an object. Ceratophrys ornata larvae possess the basic required anatomical structures for sound production via expulsion of atmospheric air from the lungs through the open soft-tissue glottis. The glottis is opened and closed via the larval laryngeal muscles (constrictor laryngis and dilatator laryngis). The arytenoid cartilages appear at stage 40 and the cricoid cartilage does at stage 43. Adult laryngeal muscles differentiate from the larval ones at stage 46 together with the vocal sac formation from the adult interhyoideus muscle. We demonstrate (n = 2160 conspecific predator–prey interactions) that larval sounds occur always under predatory attack, probably serving to diminish the chances of cannibalism. These data raise the possibility that other macrophagic carnivorous anuran larvae may produce sound.

The present study describes the reproductive biology of Argenteohyla siemersi pederseni in a modified “quebracho” forest of Schinopsis balansae in northeastern Argentina. Observations were made between January 2007 and March 2009 at El Perichón, Corrientes Province, Argentina. Argenteohyla s. pederseni bred once per year over a 3-day period in a semi-permanent pond during the first spring rains in mid-September. Males called while floating on the water surface. The advertisement call was short and repeated at regular intervals in relatively fast succession, forming groups of multiple calls. Aggressive calls were evoked in two different agonistic interactions. Eggs were black, laid in three-dimensional arrangements interspersed within the submerged vegetation. Egg number per clutch ranged from 2638 to 8987. Tadpole redescription shows many differences and similarities with previous descriptions. The reproductive biology of A. s. pederseni has several similarities with another pond-dwelling casque-headed frog species. Fil: Cajade, Rodrigo. Consejo Nacional de Investigaciones Científicas y Técnicas. Centro Científico Tecnológico Conicet - Nordeste. Centro de Ecología Aplicada del Litoral. Universidad Nacional del Nordeste. Centro de Ecología Aplicada del Litoral; Argentina Fil: Schaefer, Eduardo Federico. Consejo Nacional de Investigaciones Científicas y Técnicas. Centro Científico Tecnológico Conicet - Nordeste. Centro de Ecología Aplicada del Litoral. Universidad Nacional del Nordeste. Centro de Ecología Aplicada del Litoral; Argentina Fil: Duré Pitteri, Marta Inés. Consejo Nacional de Investigaciones Científicas y Técnicas. Centro Científico Tecnológico Conicet - Nordeste. Centro de Ecología Aplicada del Litoral. Universidad Nacional del Nordeste. Centro de Ecología Aplicada del Litoral; Argentina Fil: Kehr, Arturo Ignacio. Consejo Nacional de Investigaciones Científicas y Técnicas. Centro Científico Tecnológico Conicet - Nordeste. Centro de Ecología Aplicada del Litoral. Universidad Nacional del Nordeste. Centro de Ecología Aplicada del Litoral; Argentina Fil: Marangoni, Federico. Universidad Nacional de Posadas, Misiones. Facultad de Ciencias Exactas, Químicas y Naturales. Laboratorio de Genética Evolutiva.; Argentina

The genus Elachistocleis is a neotropical group of small sized microhylid frogs, which is represented by two species in Argentina: Elachistocleis bicolor (Guerin-Meneville, 1838) and E. skotogaster Lavilla, Vaira and Ferrari, 2003. E. skotogaster was only known for the type locality Los Toldos, Santa Victoria department, Salta, 1100 m (Lavilla et al. 2003; Lavilla 2004), however, specimens from Bolivia with spotted bellies that are currently attributed to Elachistocleis ovalis might pertain to E. skotogaster (Lavilla 2004).

The genus Homonota comprises nine South American species of terrestrial and nocturnal lizards. Homonota lizards lack the femoral pores typical of other South American Phyllodactylidae, and their infradigital lamellas are not expanded. We here describe a new species, Homonota taragui sp. nov., exclusively found on a small group of three hills up to 179 meters above sea level in central eastern Corrientes Province, Argentina. The new species differs from other Homonota species by a combination of characters, including: a well-marked dorsal, reticulate, dark pattern contrasting with a lighter colored background; small, star-shaped chromatophores on the abdomen; the post-orbital region of the head covered by granular scales; the dorsal and anterior regions of the thighs covered by keeled scales interspersed with cycloid scales; and the internasal scale in contact with rostral scales. The conservation status of Homonota taragui sp. nov. may be vulnerable, due to its localized endemism with populations on three small hills surrounded by intense agricultural and livestock activity. Two endemic plant species are known from these hills, and this new lizard represents the first endemic animal species.

El 15 Octubre de 2012, a las 19:56 hs. en la Estancia Santo Domingo, Dpto. Paso de los Libres, provincia de Corrientes (29°36?2.10?S, 56°59?23.10?O) se observó un ejemplar hembra de L. annulipes (7,05 cm longitud total) depredando sobre un ejemplar macho de la especie Erythrolamprus poecilogyrus (Wied-Neuwied, 1825) (40,5 cm longitud total). El evento se registró en una charca temporaria de 10 m de largo por 1 m de ancho aproximadamente, con una profundidad de 30 cm. La temperatura del aire y del agua al momento de la captura fue de 21°C y 24°C respectivamente. Al momento de la observación la serpiente se encontraba sin vida. Fil: Piñeiro, Jose Miguel. Consejo Nacional de Investigaciones Científicas y Técnicas; Argentina. Fundación Amado Bonpland; Argentina. Universidad Nacional del Nordeste. Facultad de Ciencias Exactas, Naturales y Agrimensura. Departamento de Biología. Laboratorio de Herpetología; Argentina Fil: Cajade, Rodrigo. Universidad Nacional del Nordeste. Facultad de Ciencias Exactas, Naturales y Agrimensura. Departamento de Biología. Laboratorio de Herpetología; Argentina. Consejo Nacional de Investigaciones Científicas y Técnicas; Argentina. Fundación Amado Bonpland; Argentina Fil: Ingaramo, María del Rosario. Universidad Nacional del Nordeste. Facultad de Ciencias Exactas, Naturales y Agrimensura. Departamento de Biología. Laboratorio de Herpetología; Argentina. Consejo Nacional de Investigaciones Científicas y Técnicas; Argentina Fil: Marangoni, Federico. Consejo Nacional de Investigaciones Científicas y Técnicas. Centro Científico Tecnológico Conicet - Nordeste. Instituto de Biología Subtropical. Instituto de Biología Subtropical - Nodo Posadas | Universidad Nacional de Misiones. Instituto de Biología Subtropical. Instituto de Biología Subtropical - Nodo Posadas; Argentina. Fundación Amado Bonpland; Argentina

Several previous studies have established that the introduction of the American bullfrog, Lithobates catesbeianus, Lithobates catesbeianus, produces negative effects on native communities. Herein, the American bullfrog is reported for the first time in the province of Buenos Aires, Argentina in the locality of 9 de Julio. The stomach contents of the collected specimens were examined and the reproduction of the species was also confirmed at the study area. A total of 85.7% (n = 30) of the analyzed stomachs contained prey items, and eleven of these were identified. The prey items belonged mostly to coleopterans (50%) and vertebrates (11%). The pathogen Batrachochytrium dendrobatidis was not found on skin samples examined. The installation of Lithobates catesbeianusn = 30) of the analyzed stomachs contained prey items, and eleven of these were identified. The prey items belonged mostly to coleopterans (50%) and vertebrates (11%). The pathogen Batrachochytrium dendrobatidis was not found on skin samples examined. The installation of Lithobates catesbeianusBatrachochytrium dendrobatidis was not found on skin samples examined. The installation of Lithobates catesbeianus breeding facilities contributes to the incidental release of this species. A tightening of rules in the regulation of aquaculture activities and the use of control programs for naturalized populations are necessary to address the increasing dispersion of the species in Argentina and potential ecological problems. Fil: Barrasso, Diego Andrés. Universidad Nacional de La Plata. Facultad de Ciencias Exactas. Departamento de Química; Argentina. Consejo Nacional de Investigaciones Científicas y Técnicas; Argentina Fil: Cajade, Rodrigo. Consejo Nacional de Investigaciones Científicas y Técnicas. Centro de Ecología Aplicada del Litoral; Argentina Fil: Nenda, Santiago Javier. Consejo Nacional de Investigaciones Científicas y Técnicas. Oficina de Coordinación Administrativa Parque Centenario. Museo Argentino de Ciencias Naturales ; Argentina Fil: Baloriani, Gabriel. Universidad Nacional de La Plata. Facultad de Ciencias Agrarias y Forestales; Argentina Fil: Herrera, Raúl. Fundación Óga; Argentina

In amphibians from tropical or subtropical regions well-expressed growth marks in bone could be expected because of the existence of strong dry/wet seasons, but little evidence is available. Additionally, temporal patterns of growth-mark formation may be a function of genetic and environmental factors. We used skeletochronology and a laboratory experiment to tease apart both causal sources, establishing direct correspondence between an observed pattern of bone growth and the actual age of Ceratophrys cranwelli and Dermatonotus muelleri from the subtropical Argentinean Chaco. Also, we describe the growth by application of the von Bertalanffy model. Growth rates decreased in both species, tending to reach asymptotic values at age 8¿10 months after metamorphosis. Both species clearly exhibited haematoxylinophilic lines of arrested growth (LAG) after one year growing in captivity. These results reinforce the hypothesis that in subtropical climates the LAG formation is ultimately caused by a general intrinsic (genetic) control.

The genus Homonota comprises nine South American species of terrestrial and nocturnal lizards. Homonota lizards lack the femoral pores typical of other South American Phyllodactylidae, and their infradigital lamellas are not expanded. We here describe a new species, Homonota taragui sp. nov., exclusively found on a small group of three hills up to 179 meters above sea level in central eastern Corrientes Province, Argentina. The new species differs from other Homonota species by a combination of characters, including: a well-marked dorsal, reticulate, dark pattern contrasting with a lighter colored background; small, star-shaped chromatophores on the abdomen; the post-orbital region of the head covered by granular scales; the dorsal and anterior regions of the thighs covered by keeled scales interspersed with cycloid scales; and the internasal scale in contact with rostral scales. The conservation status of Homonota taragui sp. nov. may be vulnerable, due to its localized endemism with populations on three small hills surrounded by intense agricultural and livestock activity. Two endemic plant species are known from these hills, and this new lizard represents the first endemic animal species.