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Cytogenetic Characterization of Three Species of Sigara from Argentina and the Plausible Mechanisms of Karyotype Evolution Within Nepomorpha

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Cytogenetic Characterization of Three Species of Sigara from Argentina and the Plausible Mechanisms of Karyotype Evolution Within Nepomorpha ISSN 0373-5680 Rev. Soc. Entomol. Argent. 66 (3-4): 81-89, 2007 81 New contributions to the study of Corixoidea: cytogenetic characterization of three species of Sigara from Argentina and the plausible mechanisms of karyotype evolution within Nepomorpha BRESSA, María José and Alba Graciela PAPESCHI Laboratorio de Citogenética y Evolución, Departamento de Ecología, Genética y Evolución, Facultad de Ciencias Exactas y Naturales, Universidad de Buenos Aires, Ciudad Universitaria. Pabellón 2, C1428EGA, Ciudad Autónoma de Buenos Aires, Argentina; e-mail: [email protected] Nuevas contribuciones al estudio de Corixoidea: caracterización citogenética de tres especies de Sigara de Argentina y los posibles mecanismos de evolución del cariotipo en Nepomorpha RESUMEN. Los estudios citogenéticos en Heteroptera contribuyen al análisis de las tendencias evolutivas en el taxón. Los Heteroptera se caracterizan por poseer cromosomas holocinéticos, diferentes sistemas de cromosomas sexuales y un par de cromosomas m en algunas especies. En este trabajo describimos el cariotipo y la meiosis masculina de Sigara denseconscripta (Breddin), S. chrostowskii Jaczewski y S. rubyae (Hungerford). Las tres especies tienen un número diploide de 24, con un par de cromosomas m y un sistema de cromosomas sexuales XY/XX. Con estos resultados son 30 las especies de Corixoidea estudiadas citogenéticamente y el cariotipo modal de la superfamilia es 2n= 20+2m+XY en machos. La información citogenética disponible hasta el presente en Heteroptera nos permite sugerir que la presencia de cromosomas m y cromosomas sexuales XY/XX, serían caracteres plesiomórficos para Nepomorpha. La ausencia de cromosomas m en especies de Nepoidea y Ochteroidea, y los sistemas de cromosomas sexuales X0 y Xn0 (en machos) en especies de Corixoidea, Naucoroidea y Nepoidea, serían caracteres derivados que habrían surgido evolutivamente más tarde. PALABRAS CLAVE. Citogenética. Heteroptera. Cromosomas holocinéticos. Meiosis. Cromosomas m. ABSTRACT. Cytogenetic studies in Heteroptera contribute to the analysis of evolutionary trends within the group. Heteroptera are characterized by the possession of holokinetic chromosomes, different sex chromosome mechanisms and a pair of m chromosomes in some species. In the present work, the male karyotype and meiosis in Sigara denseconscripta (Breddin), S. chrostowskii Jaczewski, and S. rubyae (Hungerford) are described. The three species share a diploid chromosome number of 2n= 24 with a pair of m chromosomes and an XY/XX sex Recibido: 22-V-2007; aceptado: 10-VII-2007 82 Rev. Soc. Entomol. Argent. 66 (3-4): 81-89, 2007 chromosome system. With this study the chromosome number of 30 species of Corixoidea are known and the modal karyotype is 2n= 20+2m+XY in males. The available cytogenetic information in Heteroptera led us to suggest that the presence of a pair of m chromosomes and an XY/XX sex chromosome system could be considered as plesiomorphic for Nepomorpha. The absence of m chromosomes in species of Ochteroidea and Nepoidea, and the sex chromosome systems X0 and Xn0 (male) in species of Corixoidea, Naucoroidea, and Nepoidea should be considered as derived characters, which arose later in evolution. KEY WORDS. Cytogenetics. Heteroptera. Holokinetic chromosomes. Meiosis. m chromosomes. INTRODUCTION superfamily and 26 genera, with the large and widespread genus Sigara Fabricius being Corixoidea is the largest nepomorphan divided into many subgenera (Bachmann, group and their members are commonly 1981; Schuh & Slater, 1995). The called water boatmen. They occur worldwide cosmopolitan genus Sigara comprises in various types of stable and temporary, approximately 70 species in America, continental and insular, fresh and saline distributed from Canada to southern waters (Bachmann, 1981; Schaefer & Panizzi, Argentina in Santa Cruz province and the 2000). Corixids have a high dispersal Malvinas Islands (Bachmann, 1981; Morrone potential, which allows them to utilize et al., 2004). Micronectidae includes two various available habitats (Jannson, 1986). genera: Micronecta Kirkaldy distributed in the Most of them fly very well, and they are the Old World and Australia, and Tenagobia typical insect invaders of water bodies, Bergroth distributed from Mexico to central including newly developed ones (Bachmann, Argentina in Buenos Aires province 1981; Schaefer & Panizzi, 2000). Corixids (Bachmann, 1981; Morrone et al., 2004). represent one of the most important True bugs, the Heteroptera, have many predators; they can feed on algal cells, cytogenetic characteristics that make them filamentous blue-greens, diatoms, unique among most insect groups: the microscopic protozoans and rotifers, small possession of chromosomes without a invertebrates, fish eggs and detritus primary constriction, the centromere, namely (Bachmann, 1981; Schaefer & Panizzi, 2000). holokinetic chromosomes; a pair of «m Various corixid species are incidental or chromosomes» in 16 families, belonging to obligatory predators on mosquito larvae in four infraorders; a different meiotic behaviour various regions of the world (Reynolds, 1975; for autosomes and sex chromosomes; and a Reynolds & Scudder, 1987a, b) and may be mean chiasma frequency of only one chiasma important control agents of mosquitoes. They per bivalent (Ueshima, 1979; Nokkala, 1986; can even be of significant importance for the Papeschi & Bressa, 2006). As a rule, development of some young stages of fish, autosomal bivalents are chiasmatic while the as food in small water bodies (Rask, 1983). sex chromosomes and m chromosomes are However, Corixids can be pests of fish achiasmatic. The m chromosomes are culture; some species are facultative predators generally of small size and show allocycly of fish eggs and larvae (Schaefer & Panizzi, with respect to both the autosomes and the 2000). sex chromosomes during male meiosis; they The superfamily Corixoidea comprises are usually unpaired and thus achiasmatic about 400 species, which are included in during early meiotic prophase (Ueshima, Corixidae and Micronectidae. The former 1979; Papeschi & Bressa, 2006). contains the majority of known species of the The pattern of meiosis in the Heteroptera BRESSA, M. J. and A. G. PAPESCHI. Karyotype evolution in Corixoidea 83 Table I. Diploid chromosome number in species of Corixoidea (previous data cited in Ituarte & Papeschi, 2003, 2004 are included for comparisons) Taxa 2n n Corixidae Cymatinae Cymatia borsdorffi (Sahlberg) 26 (XY) - Corixinae Arctocorisa carinata (Sahlberg) 24 (XY) 10A+m+XY A. germari (Fieber) 24 (XY) - Callicorixa concinna (Fieber) 24 (XY) - C. praeusta (Fieber) 24 (XY) 10A+m+XY C. wollastoni (Douglas & Scott) 24 (XY) 10A+m+XY Corixa dentipes (Thomson) 24 (XY) 10A+m+XY C. punctata (Illiger) 24 (XY) - Glaenocorisa cavifrons (Thomson) 24 (XY) - Hesperocorixa castanea (Thomson) 24 (XY) - H. linnaei (Fieber) 24 (XY) 10A+m+XY H. sahlbergi (Fieber) 24 (XY) 10A+m+XY Krizousacorixa femorata (Guérin) 24 (XY) 10A+m+XY Sigara denseconscripta (Breddin)* 24 (XY) 10A+m+XY S. chrowstoskii Jaczewski* 24 (XY) 10A+m+XY S. distincta (Fieber) 24 (XY) 10A+m+XY S. dorsalis (Leach) 24 (XY) - S. falleni (Fieber) 24 (XY) 10A+m+XY S. fossarum (Leach) 24 (XY) 10A+m+XY S. lateralis (Leach) 24 (XY) - S. nigrolineata (Fieber) 24 (XY) 10A+m+XY S. platensis Bachmann 24 (XY) 10A+m+XY S. rubyae (Hungerford)* 24 (XY) 10A+m+XY S. scotti (Douglas & Scott) 24 (XY) - S. semistriata (Fieber) 24 (XY) 10A+m+XY S. striata (Linnaeus) 24 (XY) 10A+m+XY S. sp 24 (XY) - Trichocorixa verticalis (Fieber) 24 (XY) - Micronectidae Micronecta poweri (Douglas & Scott) 24 (XY) - Tenagobia fuscata (Stål) 30 (XY) 14+XY * present contribution 84 Rev. Soc. Entomol. Argent. 66 (3-4): 81-89, 2007 varies between species, particularly for the & Scott) and Tenagobia fuscata (Stål), have behaviour of the sex chromosomes and the been cytogenetically analyzed (Table I) m chromosomes. During the early prophase, (Southwood & Leston, 1959; Ituarte & the sex chromosomes X and Y are positively Papeschi, 2004). While M. poweri has the heteropycnotic and remain in this condition modal chromosome number of Corixidae until diakinesis. By late diakinesis, the X and (2n= 24), T. fuscata presents many interesting Y chromosomes are separated from each cytogenetic features, such as achiasmatic other, become isopycnotic and each is male meiosis. Besides, both species are composed of two sister chromatids (Ueshima, characterized by the absence of an m 1979; Papeschi & Bressa, 2006). chromosome pair. Within Corixidae, nine The autosomal bivalents segregate genera and 25 species belonging to the reductionally during the first meiotic division subfamilies Cymatinae (Cymatia borsdorfii and equationally during the second division. (Sahlberg)) and Corixinae (24 species) have The achiasmatic m chromosomes associate been cytogenetically analyzed (Table I) at late diakinesis end-to-end, through the so- (Ituarte & Papeschi, 2004). All species of called «touch-and-go pairing»; they form a Corixinae show a diploid chromosome pseudobivalent, which segregates number of 24, with a pair of m chromosomes reductionally at anaphase I, and divides and an XY/XX sex chromosome system (male/ equationally at the second meiotic division. female), while C. borsdorfii has 2n= 26= On the other hand, sex chromosomes behave 24+XY (Ueshima, 1979). Sigara platensis as univalents in male meiosis I; they divide Bachmann was the only cytogenetically equationally at anaphase I, associate at described species inhabiting Argentina. It
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