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UNIVERSITÀ DEGLI STUDI DI URBINO “CARLO BO” DIPARTIMENTO DI SCIENZE PURE E APPLICATE CORSO DI DOTTORATO DI RICERCA IN “SCIENZE AMBIENTALI” CICLO XXVIII Tesi in cotutela: Università degli Studi di Urbino “Carlo Bo” e “Universidad de Alicante” Effects of different pastures management on dung beetle communities in a sub-mountainous landscape of central Italy: a multicomponent biodiversity and ecological process analysis Settore Scientifico Disciplinare: BIO/05 DOTTORANDO: Dott. Mattia Tonelli RELATORI: Prof. Mario Zunino Prof. José Ramón Verdú Faraco Dipartimento di Scienze Pure e Centro Iberoamericano de la Applicate Biodiversidad Università degli Studi di Urbino “Carlo CIBIO (Universidad de Alicante) Bo” ANNO ACCADEMICO 2015/2016 M. Tonelli Effects of different grazing management on dung beetle biodiversity and ecological process M. Tonelli Effects of different grazing management on dung beetle biodiversity and ecological process A te vengo, balena che tutto distruggi ma non vinci: fino all’ultimo lotto con te; dal cuore dell’inferno ti trafiggo; in nome dell’odio, vomito a te l’ultimo mio respiro (Herman Melville) …nulla si può tentare se non stabilire l’inizio e la direzione di una strada infinitamente lunga. La pretesa di qualsiasi completezza sistematica e definitiva sarebbe, se non altro, un’illusione. Qui il singolo ricercatore può ottenere la perfezione solo nel senso soggettivo che egli comunichi tutto ciò che è riuscito a vedere. (Georg Simmel) M. Tonelli Effects of different grazing management on dung beetle biodiversity and ecological process M. Tonelli Effects of different grazing management on dung beetle biodiversity and ecological process Al coraggio di un pettirosso da combattimento M. Tonelli Effects of different grazing management on dung beetle biodiversity and ecological process M. Tonelli Effects of different grazing management on dung beetle biodiversity and ecological process INDEX Resumen…………………………………………………………………………………....1 Chapter 1: General introduction and objectives……………………………………...........7 Chapter 2: Effects of grazing intensity and the use of veterinary medical products on dung beetle biodiversity in the sub-mountainous landscape of Central Italy………….....……..29 Chapter 3: Effects of progressive grazing abandonment on dung beetle biodiversity in a central Italian province: nesting behavior and body size matters………………………....75 Chapter 4: The effects of grazing intensity and historical veterinary medical products use on dung beetle functional diversity and ecological process………………………...........123 Chapter 5: The effects of progressive grazing abandonment on dung beetle functional diversity and ecological process………………………………………………………….159 Chapter 6: General discussion and conclusions…………………………………………193 Annex 1: Catalog, biology and phenology of the species………………………………..211 Annex 2: Selected and measured functional traits……………………………………….351 Acknowledgments……………………………………………………………………….377 M. Tonelli Effects of different grazing management on dung beetle biodiversity and ecological process M. Tonelli Effects of different grazing management on dung beetle biodiversity and ecological process RESUMEN M. Tonelli Effects of different grazing management on dung beetle biodiversity and ecological process M. Tonelli Effects of different grazing management on dung beetle biodiversity and ecological process RESUMEN Los escarabajos coprófagos (Coleoptera: Scarabaeoidea) son un grupo de insectos muy importante en términos de diversidad y biomasa en los pastizales de la cuenca del Mediterráneo. A través del uso de estiércol como recurso trófico y para nidificar, estos organismos están involucrados en numerosos procesos ecológicos relacionados con la degradación del estiércol, tales como en el ciclo de nutrientes, en la dispersión secundaria de semillas, contribuyen mejorando la estructura del suelo, en la reducción de emisión de gases de efecto invernadero, así como en el control de endo-ecto parásitos del ganado. A pesar de su importancia ecológica, los escarabajos coprófagos están amenazados por varios factores, entre los cuales se destacan los cambios en el manejo de la ganadería tradicional. En las últimas décadas, la ganadería tradicional ha sufrido cambios radicales debidos principalmente a dos procesos: el abandono progresivo de la ganadería y su intensificación. Por un lado, el abandono progresivo de la ganadería tradicional es un factor clave en la conservación de los escarabajos coprófagos. Debido a la dependencia que este grupo de insectos tiene con el estiércol, este factor llevaría a una disminución del recurso trófico con repercusiones negativas para la comunidad coprófaga. Históricamente, los escarabajos coprófagos han dependido del uso de los excrementos de la megafauna salvaje, y desde su disminución, el estiércol de la ganadería doméstica ha sido su principal recurso trófico. A pesar de ello, en los últimos 50 años, la ganadería tradicional ha comenzado un proceso de abandono progresivo, sobre todo en las áreas marginales como las de montaña. Por otra parte, el uso/abuso de productos médico veterinarios para el control de los endoparásitos y ectoparásitos del ganado es otro factor importante en tema de conservación de la fauna coprófaga y está relacionado con la 1 M. Tonelli Effects of different grazing management on dung beetle biodiversity and ecological process intensificación de la ganadería. Estos compuestos médico veterinarios, entre los cuales la ivermectina destaca por su frecuancia en el uso y el efecto negativo en la fauna coprófaga, son eliminados progresivamente en las heces de los organismos. Estudios recientes han demostrado que los residuos de algunos de estos compuestos tienen efectos sub-letales y letales sobre la fauna coprófaga y sobre todo en los escarabajos. A pesar de la importancia que estos dos factores pueden tener en la conservación de los escarabajos coprófagos, pocos estudios comparativos han investigado el impacto sobre las comunidades de este grupo y como consecuencia sobre el papel ecológico en los cuales están involucrados. Debido a este vacío de conocimiento, el objetivo general de esta tesis fue evaluar los efectos del abandono de la ganadería, su extensificación y uso histórico de productos médico veterinarios sobre las comunidades de escarabajos coprófagos usando medidas de biodiversidad y cuantificando la capacidad de remoción de estiércol en un paisaje sub-montano de la provincia de Pesaro-Urbino (Italia). Después de una introducción general (Capítulo 1) en los capítulos 2 y 3 de esta tesis se abordan los efectos de los diferentes métodos de manejo de la ganadería y se analiza el impacto de estos factores sobre la diversidad alfa y beta, abundancia y biomasa, especies indicadoras, grupos funcionales y clases de tallas de las especies. En los capítulos 4 y 5 se evalúan los efectos de estos factores desde una perspectiva funcional, analizando la diversidad funcional de las comunidades y el desempeño que cada comunidad tiene en la remoción del estiércol. Así mismo, se analizaron las relaciones entre varias medidas de diversidad funcional y de diversidad clásicas (número de especies y exponencial de Shannon) y la relación entre la diversidad funcional y el proceso ecológico. 2 M. Tonelli Effects of different grazing management on dung beetle biodiversity and ecological process En este trabajo se colectaron un total de 156.936 individuos, pertenecientes a 58 especies de escarabajos coprófagos (3 Geotrupinae; 16 Scarabaeinae; 39 Aphodiinae) y por primera vez se cita Calamosternus mayeri (Pilleri, 1953) en la región Marche. Los resultados de este trabajo sugieren que el uso histórico de productos médico veterinarios tiene un impacto negativo y ubiquitario sobre todas las especies de la comunidad de estudio, llevando a un empobrecimiento de los ensambles y a una pérdida de abundancia y biomasa. No obstante se ha registrado el mantenimiento de la estructura de la comunidad. Estos resultados son corroborados igualmente cuando nos enfocamos en la diversidad funcional. En efecto, el uso histórico de productos médico veterinarios lleva a una pérdida en el número de roles funcionales dentro de la comunidad, sin embargo las abundancias se encuentran repartidas equitativamente dentro de cada nicho funcional. Todos estos efectos sobre la comunidad de escarabajos coprófagos debido al uso de productos médico veterinarios, al final quedan reflejados en una pérdida del 70% en la capacidad de remoción de estiércol, comparados con áreas donde estos productos no son aplicados. La extensificación de la ganadería está representada por áreas que tienen una intensidad de pastoreo muy baja, las cuales pueden ser consideradas como la primera etapa hacia el abandono completo. En este sentido, los resultados de este trabajo demuestran como una simple reducción de la carga ganadera lleva a efectos negativos sobre las comunidades de escarabajos coprófagos, registrando una disminución en la diversidad alfa y en el número de especies indicadoras. Esta disminución en la cantidad de recurso trófico tiene implicaciones sobre la composición de la comunidad, favoreciendo especies oportunistas cuya biología le permite evitar la competencia, como por ejemplo; especies cleptoparásitas o especies cuyas larvas tienen un comportamiento 3 M. Tonelli Effects of different grazing management on dung beetle biodiversity and ecological process saprófago. Este “efecto filtro” (filter effect) debido a la disminución de la cantidad trófica, está confirmado por los resultados de diversidad funcional, en los cuales
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  • Quick Guide for the Identification Of

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    Quick Guide for the Identification of Maryland Scarabaeoidea Mallory Hagadorn Dr. Dana L. Price Department of Biological Sciences Salisbury University This document is a pictorial reference of Maryland Scarabaeoidea genera (and sometimes species) that was created to expedite the identification of Maryland Scarabs. Our current understanding of Maryland Scarabs comes from “An Annotated Checklist of the Scarabaeoidea (Coleoptera) of Maryland” (Staines 1984). Staines reported 266 species and subspecies using literature and review of several Maryland Museums. Dr. Price and her research students are currently conducting a bioinventory of Maryland Scarabs that will be used to create a “Taxonomic Guide to the Scarabaeoidea of Maryland”. This will include dichotomous keys to family and species based on historical reports and collections from all 23 counties in Maryland. This document should be cited as: Hagadorn, M.A. and D.L. Price. 2012. Quick Guide for the Identification of Maryland Scarabaeoidea. Salisbury University. Pp. 54. Questions regarding this document should be sent to: Dr. Dana L. Price - [email protected] **All pictures within are linked to their copyright holder. Table of Contents Families of Scarabaeoidea of Maryland……………………………………... 6 Geotrupidae……………………………………………………………………. 7 Subfamily Bolboceratinae……………………………………………… 7 Genus Bolbocerosoma………………………………………… 7 Genus Eucanthus………………………………………………. 7 Subfamily Geotrupinae………………………………………………… 8 Genus Geotrupes………………………………………………. 8 Genus Odonteus...……………………………………………… 9 Glaphyridae..............................................................................................
  • Phylogenetic Analysis of Geotrupidae (Coleoptera, Scarabaeoidea) Based on Larvae

    Phylogenetic Analysis of Geotrupidae (Coleoptera, Scarabaeoidea) Based on Larvae

    Systematic Entomology (2004) 29, 509–523 Phylogenetic analysis of Geotrupidae (Coleoptera, Scarabaeoidea) based on larvae JOSE´ R. VERDU´ 1 , EDUARDO GALANTE1 , JEAN-PIERRE LUMARET2 andFRANCISCO J. CABRERO-SAN˜ UDO3 1Centro Iberoamericano de la Biodiversidad (CIBIO), Universidad de Alicante, Spain; 2CEFE, UMR 5175, De´ partement Ecologie des Arthropodes, Universite´ Paul Vale´ ry, Montpellier, France; and 3Departamento Biodiversidad y Biologı´ a Evolutiva, Museo Nacional de Ciencias Naturales (CSIC), Madrid, Spain Abstract. Thirty-eight characters derived from the larvae of Geotrupidae (Scarabaeoidea, Coleoptera) were analysed using parsimony and Bayesian infer- ence. Trees were rooted with two Trogidae species and one species of Pleocomidae as outgroups. The monophyly of Geotrupidae (including Bolboceratinae) is supported by four autapomorphies: abdominal segments 3–7 with two dorsal annulets, chaetoparia and acanthoparia of the epipharynx not prominent, glossa and hypopharynx fused and without sclerome, trochanter and femur without fossorial setae. Bolboceratinae showed notable differences with Pleocomidae, being more related to Geotrupinae than to other groups. Odonteus species (Bolboceratinae s.str.) appear to constitute the closest sister group to Geotrupi- nae. Polyphyly of Bolboceratinae is implied by the following apomorphic char- acters observed in the ‘Odonteus lineage’: anterior and posterior epitormae of epipharynx developed, tormae of epipharynx fused, oncyli of hypopharynx devel- oped, tarsal claws reduced or absent, plectrum and pars stridens of legs well developed and apex of antennal segment 2 with a unique sensorium. A ‘Bolbelas- mus lineage’ is supported by the autapomorphic presence of various sensoria on the apex of the antennal segment, and the subtriangular labrum (except Eucanthus). This group constituted by Bolbelasmus, Bolbocerosoma and Eucanthus is the first evidence for a close relationship among genera, but more characters should be analysed to test the support for the clade.