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Evidencia De Cortejo Copulatorio En El Orden Scorpiones (Arachnida), Con Un Análisis En Zabius Fuscus (Buthidae)

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Rev. Soco Entomol. Argent. 56 (1-4): 21-30,1997 Evidencia de cortejo copulatorio en el orden Scorpiones (Arachnida), con un análisis en Zabius fuscus (Buthidae) PERETTI, ALFREDO V. Cátedra de Diversidad Animal J, Facultad de Ciencia s Exactas, Físicas y Naturales, Universidad Nacional de Córdoba, Avda. Vélez Sársfield 299, 5000 Córdoba, Argentina. o ABSTRACT. Evidence ofcopulatory courtship inthe orderScorpiones (Arachnida), with an analysis in Zabiusfuscus (Buthidae). The presence of courtship behavior during and after sperm transfer of scorpions is studied. The whole literature on mating behavior of scorpions was reviewed. The re­ view included six families, 22 genera, and 37 species. In laboratory condi­ tions, the mating and sperm transfer on Zabius fuscus (Buthidae) were re­ corded. The relationship between copulatory courtship and sperm uptake was studied, 11 % of the species show copulatory courtship. The behavioral patterns of copulatory courtship in scorpions are the rubbings with cheliceres (RQ) and legs (RP). The occurrence of these patterns may be restricted by mechanism offemale subjection during sperm transfer. The possible underes­ timate of the frequency of copulatory courtship in scorpions is discussed. O INTRODUCCiÓN para la misma finalidad. Éstas se utilizarían para excitar a la hembra, actuando a modo de "cortejo A partir del enunciado de Thornhill (1983) interno" (Eberhard, 1985) dentro de sus vías sobre la existencia de elección femenina encu­ genitales o para asociarse de forma más precisa bierta ("female cryptic choice"), actualmente se está con los genitalia de ella. La presencia de cortejo produciendo un proceso de revisión general del copulatorio y/o excitación por medio de los geni­ cortejo de las especies con fertilización interna. talia sería un valioso indicativo de la existencia de Por medio de este tipo de elección, la hembra elección femenina encubierta, estando ella bajo puede en cierta forma promover, influir positiva o una considerable presión de selección sexual negativamente en procesos post-inseminatorios (Eberhard, 1990). (transporte de esperma, utilización en fertilización Si bien esta teoría ha sido analizada en ciertos de huevos, etc.). grupos de insectos y a rañas (Eberhard, 1991, Eberhard (1991) ha postulado que en las es­ 1992a, b, 1993a, b, e, 1994; Rodríguez & Eber­ pecies en las que esto se presenta, la selección fa­ hard, 1994), actualmente está en etapa de com­ vorecería en el macho la aparición de estrategias probación. Principalmente es necesario confirmar de conducta o de estructuras genitales que induz­ ciertos supuestos, tales como si la realización de can a la hembra a realizar los procesos de post­ cortejo copulatorio realmente incrementa el éxito intromisión que incrementen su oportunidad de de la inseminación y/o fecundación. Además, una fertilización. También se incluyen estructuras no tarea importante es examinar otros grupos para genitales (patas, alas, etc.) que intervienen en el determinar si el fenómeno está realmente generali­ cortejo y componentes químicos del esperma. La zado, o siestá limitado a ciertos gnJpos taxonómicos. aparición de conductas típicas de cortejo durante El presente trabajo tiene como objetivo deter­ o después de la cópula, cumpliría esa función, minar si existe cortejo copulatorio en el orden comportamiento denominado "cortejo copulato­ Scorpiones, considerando en el análisis todas las rio" (Eberhard, 1991). Si se considera que un ma­ especies estudiadas hasta el momento. Si bien en cho efectúa el cortejo para inducir a que la hembra este grupo por el tipo de mecanismo de insemi­ ferti lice sus huevos con sus espermatozoides, su nación -con espermatóforo (Thomas & Zeh, 1984; realización durante o después de la inseminación Polis & Sissom, 1990)- está más generalizado el promovería que ella luego realice los procesos que término "transferencia espermática" que "CÓp U­ incrementen sus posibilidades de paternidad. En la", también conviene utilizar este último para fa­ cuanto a los genitalia, Eberhard (1985, 1993a, b, cilitar el análisis. Además, desde un punto de vis­ e) sostiene que ciertas estructuras, tales como espi­ ta funcional, en los escorpiones la hembra real­ nas, tubérculos y otras ornamentaciones, servirían mente "copula" con el espermátoforo, proceso Rev. Soco Entomol. Argent. 56 (1-4), 1997 que el macho guía y dirige (Polis & Sissorn, 1990; o no esperma durante la cópula. Por tal motivo, Peretti, 1992; Benton, 1993). Recientemente, en este trabajo incluye como complemento un estu­ muchas especies de Bothriuridae se han estable­ dio preliminar sobre la relación que existe entre el cido las secuencias de los patrones de conducta cortejo copulatorio y la transferencia espermática. que intervienen en el cortejo "pre-copulatorio" (Peretti, 1991, 1993, 1995a, 1996). Esto se suma a lo que se conoce en el resto del orden sobre las MATERIAL Y MÉTODOS posibles funciones de los distintos patrones ge­ nerales de cortejo ya identificados (Tabla 1), he­ Estudio comparado del orden Scorpiones. Se cho que permite comenzar a abordar este tema. revisaron los trabajos publicados sobre el compor­ Debido a que los escorpiones transfieren su tamiento de apareamiento en escorpiones, más da­ esperma por medio de un espermatóforo, es po­ tos inéditos de Timogenes dorbignyi (Guérin) sible determinar si aquél pasa o no al interiorde la (Bothriuridae). Esto incluye un total de seis familias, hembra. Cabe señalar que esto representa una 22 géneros y 37 especies (Tabla 11). Sólo se han ventaja metodológica para el investigador, com­ considerado los trabajos originales, ya que ulterio­ parado con los animales que poseen pene u otros res publicaciones comparativas pueden omitir la órganos copuladores, en los cuales puede ser más cita de conductas asociadas al cortejo copulatorio. difícil determinar a simple vista si se ha transferido Con los datos registrados se confeccionaron tablas Tabla I. Conductas de cortejo del macho para la excitación sexual e inhibición de la agresión en la hembra. Entre paréntesis figuran las denominaciones que han utilizado otros autores. Para cada conducta se especi­ fica su función y se citan los trabajos que analizaron (*) o propusieron dicha función. (1) Conductas presentes en Zabius fuscus . ------------------ Tipos generales de conductas Función Referencias "Arqueo conjunto de metasomas Inhibe la agresión de la hembra Polis & Sissom, 1990; Peretti, (AMC)" (Arbre droit ! clubbing) (1) 1991, 1993 "Clavada de aguijón (CLA)" (piqfues ídem Polis & Sissom, 1990; Peretti, sexuelles, sexual mating, pungere) 1993 (*) "Estremecimientos pre y post agarre Reconocimiento sexual, secundaria­ Alexander, 1959; Polis & Sissom, de hembra (EPAjEPOA)", "vibración mente excitación sexual 1990; Gaffin & Brownell, 1992 (*)¡ (VIB)" (variantes de juddering) (1) Peretti, 1995a (*) "Frote con pinza (FP)" Inhibe la agresión de la hembra Peretti, 1993 "Jaladura y aproximación (JA)" Excitación sexual Peretti, 1993 "Roce con quelíceros (RQ)" (kiss, kiss­ Excitación sexual, inhibidor de la Polis & Farley, 1979; Polis & ing, embrassades, chericeral massage) agresión de la hembra Sissom, 1990; Peretti, 1991, (1) 1993 (*), en prensa "Roce con patas (RP)" (trilling, thrills, Excitación sexual Peretti, 1991, 1993 (*) tickling, trilles, trillerphase) (1) ''Roce con glándulas del metasoma (RP)" Excitación sexual, inhibidor de la Peretti, 1993 (*), 1995a agresión "Sacudida del metasoma (SAC)" ídem Peretti, 1993 Total: 9 en Z. fuscus: 4 de presencia-ausencia de conductas para deterrni­ senta si fue descripto al menos en una especie. nar las frecuencias del cortejo copulatorio en las Para la identificación y descripción de los pa.. especies, géneros y familias, así como qué tipos de trones de conducta se siguieron las pautas ya esta­ patrones intervienen. De acuerdo con el criterio de blecidas en trabajos anteriores (Peretti, 1991, 1996). Eberhard (1991) se considera que una especie tiene Para definir cuándo una conducta podía ser con­ cortejo copulatorio si al menos un trabajo lo descri­ siderada de "cortejo", se tuvieron en cuenta los be. Se considera que una familia o género lo pre- conceptos ya establecidos por Alexander (1957, 22 PERETfI, A. Y., cortejo copulatorio en escorpiones Tabla II. Lista de las 37 especies y subespecies de escorpiones en las que se analiza si presentan cortejo copulatorio o no. Sólo se han considerado los trabajos que incluyen la secuencia completa de al menos una transferencia espermática. * Especies estudiadas con mayor detalle. Taxón Tipo de País Referencias Cortejo espermatóforo copulatorio BUTHIDAE Androctonus australis (Linné) flageliforme Argelia Auber-Thomay, 1974 no Buthus occitanus Amoreux Francia Auber,1963 no Hottentota judaica (Simon) Israel Shulov & Amitai, 1958 no Leiurus quinquestriatus (Hemprich Israel Shulov & Amitai, 1958; Abu- si & Erenberg)* shama, 1968; Benton, 1992 Parabuthus planicauda Pocock" Sudáfrica Alexander, 1959 no Mesobuthus tamulus tamulus (Fa- India Yadav & Kamble, 1989 no bricius) Centruroides vittatus (Say) USA McAlister, 1965 no C. guanensis cubensis Armas Cuba Armas, 1980 no C. gracilis Armas Cuba Armas, 1980 no C. armadai Armas Cuba Armas, ]980 no Rophalurus garridoi Armas Cuba Armas, ]986 no Tityus trinitatis Pocock" Trinidad Alexander, 1959 no T. bahiensis (Perty) Brasil Bücherl, ]956; Matthiesen, ] 960 no T. trivittatus Kraepelin Brasil Bücherl, 1956 no T. fasciolatus Pessóa Brasil Lourenco, 1979 no Zabius juscus (Thorell)* Argentina Peretti, 1991 si lsometrus maculatus (De Geer) Tanzania
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    Os escorpioes UNIVERSIDADE FEDERAL DA BAHIA Reitora Dora Leal Rosa Vice Reitor Luiz Rogério Bastos Leal EDITORA DA UNIVERSIDADE FEDERAL DA BAHIA Diretora Flávia Goulart Mota Garcia Rosa CONSELHO EDITORIAL Titulares Ângelo Szaniecki Perret Serpa Caiuby Alves da Costa Charbel Ninõ El-Hani Dante Eustachio Lucchesi Ramacciotti José Teixeira Cavalcante Filho Alberto Brum Novaes Suplentes Evelina de Carvalho Sá Hoisel Cleise Furtado Mendes Maria Vidal de Negreiros Camargo Tania Kobler Brazil Tiago Jordão Porto Os escorpioes EDUFBA Salvador 2010 ©2010 by autores (org.) Direitos para esta edição cedidos à Edufba. Feito o depósito legal. Projeto gráfico, capa e programação visual Rodrigo Oyarzábal Schlabitz Revisão Magel Castilho Normalização Susane Barros Sistema de Bibliotecas - UFBA Brazil, Tania Kobler. Os escorpiões / Tania Kobler Brazil, Tiago Jordão Porto ; prefácio Sylvia Marlene Lucas ; apresentação Tania Kobler Brazil. - Salvador : EDUFBA, 2010. 84 p. ISBN 978-85-232-0729-8 1. Escorpiões - Brasil. I. Porto, Tiago Jordão. II. Lucas, Sylvia Marlene. III.Título. CDD - 595.460981 Associação Brasileira de Editoras Universitárias EDUFBA Rua Barão de Jeremoabo, s/n, Campus de Ondina, 40170-115, Salvador-BA, Brasil Tel/fax: (71) 3283-6164 www.edufba.ufba.br | [email protected] AGRADECIMENTOS Parte deste livro é produto das revisões bibliográficas e dos resultados de pesquisas conduzidas durante a graduação de Tiago Jordão Porto. Por isso agradecemos ao Conselho Nacional de Desen- volvimento Científico e Tecnológico (CNPq), que financiou duas bolsas de Iniciação Científica no Programa Institucional de Bolsas de Iniciação Científica da Universidade Federal da Bahia (PIBIC- UFBA). À Fundação de Amparo à Pesquisa do Estado da Bahia (FAPESB), que financiou outras duas bolsas de Iniciação Científicas de Tiago vinculadas a projetos de pesquisa coordenados pela professo- ra Rejâne Maria Lira da Silva, a quem também agradecemos.
  • Meiosis in the Scorpion Tityus Silvestris: New Insights Into

    Meiosis in the Scorpion Tityus Silvestris: New Insights Into

    © 2019. Published by The Company of Biologists Ltd | Biology Open (2019) 8, bio040352. doi:10.1242/bio.040352 RESEARCH ARTICLE Meiosis in the scorpion Tityus silvestris: new insights into achiasmatic chromosomes Bruno Rafael Ribeiro de Almeida, Renata Coelho Rodrigues Noronha, Marlyson Jeremias Rodrigues da Costa, Cleusa Yoshiko Nagamachi and Julio Cesar Pieczarka* ABSTRACT 1950a,b; Cunha and Pavan, 1954; Adilardi et al., 2016; Ojanguren- Achiasmatic male meiosis in scorpions is characterized by a high Affilastro et al., 2017). The presence of holocentric chromosomes frequency of gaps, asynaptic regions and multivalent associations. favors fixation of these rearrangements in several natural Here, we performed an immunocytogenetic analysis to investigate populations (Mattos et al., 2013). Direct consequences of such recombination, and synapsis and chromatin-remodeling events during changes are detected during prophase I events, as evidenced by meiosis of the scorpion Tityus silvestris. Our results demonstrate synaptic anomalies, such as gaps, asynapsed regions, loop-like that the synaptonemal complex (SC) begins its organization in the structures and a high frequency of heterosynapsis, in several zygotene stage and persists until metaphase I. The advancement of members of this genus (Schneider et al., 2009b). the synaptic process is related to the epigenetic modification histone Another interesting aspect of Tityus meiosis is the absence of H3 lysine 27 trimethylation (H3K27m3). The distribution and dynamics chiasma in males (Schneider et al., 2009a). Achiasmatic meiosis is patterns of variant γH2AX and recombinase Rad51 during achiasmatic not unique to this genus, occurring in all Scorpiones families, as meiosis suggests formation and repair of DNA double-strand well as in other organisms, such as plants, flatworms and insects breaks (DSBs) during early stages of prophase I.
  • Insights Into the Origin of the High Variability of Multivalent-Meiotic Associations in Holocentric Chromosomes of Tityus (Archaeotityus) Scorpions

    Insights Into the Origin of the High Variability of Multivalent-Meiotic Associations in Holocentric Chromosomes of Tityus (Archaeotityus) Scorpions

    RESEARCH ARTICLE Insights into the origin of the high variability of multivalent-meiotic associations in holocentric chromosomes of Tityus (Archaeotityus) scorpions Viviane Fagundes Mattos1, Leonardo Sousa Carvalho2¤, Marcos Andre Carvalho3, Marielle Cristina Schneider4* 1 Universidade Estadual Paulista ªJuÂlio de Mesquita Filhoº, UNESP, Departamento de Biologia, Rio Claro, São Paulo, Brazil, 2 Universidade Federal do PiauõÂ, UFPI, Campus AlmõÂcar Ferreira Sobral Floriano, PiauõÂ, Brazil, 3 Universidade Federal de Mato Grosso, UFMT, Departamento de Biologia e Zoologia, CuiabaÂ, Mato Grosso, Brazil, 4 Universidade Federal de São Paulo, UNIFESP, Departamento de Ecologia e Biologia a1111111111 Evolutiva, Diadema, São Paulo, Brazil a1111111111 a1111111111 ¤ Current address: PoÂs-GraduacËão em Zoologia, Universidade Federal de Minas Gerais, UFMG, Belo Horizonte, Minas Gerais, Brazil a1111111111 * [email protected] a1111111111 Abstract Scorpions represent an intriguing group of animals characterized by a high incidence of het- OPEN ACCESS erozygous chromosomal rearrangements. In this work, we examined six species of Tityus Citation: Mattos VF, Carvalho LS, Carvalho MA, Schneider MC (2018) Insights into the origin of the (Archaeotityus) from Brazilian fauna with a particular focus on elucidating the rearrange- high variability of multivalent-meiotic associations ments responsible for the intraspecific variability of diploid number and the presence of long in holocentric chromosomes of Tityus chromosomal chains in meiosis. To access any interpopulation diversity, we also studied (Archaeotityus) scorpions. PLoS ONE 13(2): individuals from four species representing distinct localities. Most species demonstrated e0192070. https://doi.org/10.1371/journal. pone.0192070 intraspecific polymorphism in diploid number (2n = 19 and 2n = 20 in T. clathratus, T. matto- grossensis, and T. pusillus, 2n = 16, 2n = 17 and 2n = 18 in T.