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Confirmation of Parthenogenesis in the Medically Significant, Synanthropic Scorpion Tityus Stigmurus (Thorell, 1876) (Scorpiones: Buthidae)

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Confirmation of Parthenogenesis in the Medically Significant, Synanthropic Scorpion Tityus Stigmurus (Thorell, 1876) (Scorpiones: Buthidae) NOTA BREVE: Confirmation of parthenogenesis in the medically significant, synanthropic scorpion Tityus stigmurus (Thorell, 1876) (Scorpiones: Buthidae) Lucian K. Ross NOTA BREVE: Abstract: Parthenogenesis (asexuality) or reproduction of viable offspring without fertiliza- Confirmation of parthenogenesis tion by a male gamete is confirmed for the medically significant, synanthropic in the medically significant, synan- scorpion Tityus (Tityus) stigmurus (Thorell, 1876) (Buthidae), based on the litters thropic scorpion Tityus stigmurus of four virgin females (62.3–64.6 mm) reared in isolation in the laboratory since (Thorell, 1876) (Scorpiones: Buthi- birth. Mature females were capable of producing initial litters of 10–21 thely- dae) tokous offspring each; 93–117 days post-maturation. While Tityus stigmurus has been historically considered a parthenogenetic species in the pertinent literature, Lucian K. Ross the present contribution is the first to demonstrate and confirm thelytokous 6303 Tarnow parthenogenesis in this species. Detroit, Michigan 48210-1558 U.S.A. Keywords: Buthidae, Tityus stigmurus, parthenogenesis, reproduction, thelytoky. Phone/Fax: +1 (313) 285-9336 Mobile Phone: +1 (313) 898-1615 E-mail: [email protected] Confirmación de la partenogénesis en el escorpión sinantrópico y de relevan- cia médica Tityus stigmurus (Thorell, 1876) (Scorpiones: Buthidae) Resumen: Se confirma la partenogénesis (asexualidad) o reproducción con progenie viable Revista Ibérica de Aracnología sin fertilización mediante gametos masculinos en el escorpión sinantrópico y de ISSN: 1576 - 9518. relevancia médica Tityus stigmurus (Thorell, 1876) (Scorpiones: Buthidae). Cua- Dep. Legal: Z-2656-2000. tro hembras vírgenes (Mn 62.3-64.6) se criaron en el laboratorio desde su naci- Vol. 18 miento. Al alcanzar el estadio adulto tuvieron una descendencia telitoca inicial de Sección: Artículos y Notas. 10 - 21 crías cada una después de 93-117 días de alcanzar su madurez. Mien- Pp: 115-121 tras Tityus stigmurus ha sido considerado históricamente una especie parteno- Fecha publicación: 30-Junio-2010 genética, la presente contribución es la primera en demostrar y confirmar la partenogénesis telitoca de esta especie. Palabras clave: Buthidae, Tityus stigmurus, partenogénesis, reproducción, thelitoquia. Edita: Grupo Ibérico de Aracnología (GIA) Grupo de trabajo en Aracnología de la Sociedad Entomológica Aragonesa (SEA) Avda. Radio Juventud, 37 Introduction 50012 Zaragoza (ESPAÑA) Tef. 976 324415 Parthenogenesis (asexuality) or production of genetically identical off- Fax. 976 535697 C-elect.: [email protected] spring from unfertilized ova has been reported to be a relatively rare phe- nomenon in nature (Lynch 1984; Palmer & Norton 1991; Francke 2007; Director: Carles Ribera Lourenço 2008a), with 95% of all living organisms reproducing sexually C-elect.: [email protected] (Lourenço & Cuellar 1995). However, despite its seeming rarity in nature, asexual reproduction occurs in plants and in at least 2000 animal species Indice, resúmenes, abstracts (Rao 2010). Arrhenotoky (production of all-male progeny), deutherotoky vols. publicados: (production of male and female progeny), and thelytoky (production of all- http:/www.sea-entomologia.org female progeny) have been reported to occur in scorpions (Lourenço & Cuellar 1999; Lourenço 2008a). Thelytoky is the most widespread form of Página web GIA: http://gia.sea-entomologia.org parthenogenesis in scorpions and other organisms (Carson 1967; Lynch 1984; Francke 2007; Lourenço 2002a, 2008a; Lourenço et al. 2000). In the Arachnida, parthenogenetic reproduction has been reported to occur in mites Página web SEA: (Acari), spiders (Araneae), harvestmen (Opiliones), and scorpions (Scor- http:/www.sea-entomologia.org piones) (references in, Francke 2007; Lourenço 2008a). Parthenogenetic re- production in scorpions was first reported for the Brazilian yellow scorpion, 116 Lucian K. Ross Tityus (Tityus) serrulatus Lutz & Mello, 1922 by Mat- 2009). It is primarily of synanthropic habitats in the thiesen (1962). Subsequently, parthenogenesis has been states of Alagoas, Bahia, Ceará, Paraíba, Pernambuco, reported to occur in an additional 15 scorpion species, Piauí, Rio Grande do Norte, Sergipe, and the island of belonging to four families, and nine genera (Francke Fernando de Noronha (Pernambuco), northeastern Brazil 2007; Lourenço 2008a,b). The majority of parthenoge- (Dias et al. 2006; Freitas & Vasconcelos, 2008; Souza et netic scorpion species belong to the cosmotropical fam- al. 2009). This species is also considered to be one of ily Buthidae C. L. Koch, 1837, and the exclusively the three most toxic species and leading causal agents of Neotropical buthid genus Tityus C. L. Koch, 1836. Par- scorpionism in Brazil, where T. stigmurus is responsible thenogenesis has been reported to occur in seven Tityus for hundreds of envenomations each year (Eickstedt species: Tityus (Archaeotityus) columbianus (Thorell, 1983/84; Lira-Da-Silva et al. 2000; Barbosa et al. 2003; 1876), Tityus (Atreus) metuendus Pocock, 1897, Tityus Batista et al. 2007; Freitas & Vasconcelos 2008; Albu- (Tityus) serrulatus Lutz & Mello, 1922, Tityus (Tityus) querque et al. 2009a,b). The state of Pernambuco has the stigmurus (Thorell, 1876), Tityus (Tityus) trivittatus highest rate of T. stigmurus envenomations in Brazil, Kraepelin, 1898, Tityus (Tityus) uruguayensis Borelli, with 43.5 incidents per 100,000 inhabitants (Albuquer- 1901, and Tityus (Atreus) neblina Lourenço, 2008 que et al. 2009a). In urban areas, this species is abundant (Francke 2007; Lourenço 2008a,b). However, some of and commonly found in homes and other manmade those reports have been controversial and others have structures; among debris in backyards and construction many shortcomings and/or factual inaccuracies (Francke sites, in waste material, sewage ditches, cess pits, stored 2007; Lourenço 2008a). Francke (2007) considered par- building materials, and garbage (Aguiar et al. 2008; Al- thenogenesis unsubstantiated and published reports as buquerque et al. 2009a). The majority of breeding sites invalid for the medically significant, synanthropic scor- used by T. stigmurus are in homes or their vicinity (Al- pion species T. stigmurus. Historically, T. stigmurus was buquerque et al. 2009a). Knowledge of the biology and considered a sexual species until unpublished field ob- development of T. stigmurus is important due to the servations of parthenogenesis were reported by medical significance of its venom, synanthropic habits, Lourenço & Cloudsley-Thompson (1999). While synan- high rates of envenomations in humans, and the ability thropic populations of T. stigmurus are considered to be of this species to rapidly proliferate and colonize human parthenogenetic, a geographically limited sexual popula- occupied areas. tion of T. stigmurus has been reported from an undis- Francke (2007) briefly reviewed the history of claims of turbed region of Exu in the state of Pernambuco, north- parthenogenesis in T. stigmurus. In that paper he con- eastern Brazil (Lourenço & Cloudsley-Thompson 1999). cluded that while several claims have alluded to the abil- As parthenogenesis indicates the ability of an unmated ity of females of T. stigmurus to reproduce parthenoge- sexually mature female to produce offspring without netically, no evidence was provided by any of the au- insemination by a male, then the most accurate and di- thors to support this claim (Lourenço & Cloudsley- rect means to determine the ability of a female to pro- Thompson 1999; Lourenço et al. 2000; Lourenço duce offspring parthenogenetically is by independent 2002a), and thereby, considered parthenogenetic repro- rearing of immature specimens to maturity without the duction as unsubstantiated for T. stigmurus. Since the presence of males (Toscano-Gadea 2004; Yamazaki & publication of Lourenço (2002a), subsequent reviews Makioka 2004; Francke 2007). Sexually immature fe- and studies of parthenogenesis and various life history males (pre-adult or younger) are as a general rule both aspects of T. stigmurus have been published without unreceptive and unattractive to sexually mature males providing evidence to substantiate claims of partheno- (Francke 2007). Thus, parturition in captivity by a fe- genesis occurring in this species (Aguiar et al. 2008; male which was caught as an immature, and which Lourenço 2008a; Outeda-Jorge et al. 2009; Souza et al. molted (once or more) to attain sexual maturity while in 2009). Freitas & Vasconcelos (2008) indicated that par- captivity, can be construed as indisputable evidence of thenogenesis as a form of reproduction has yet to be parthenogenesis (Toscano-Gadea 2004; Yamazaki & confirmed for T. stigmurus. Recently, Scholte et al. Makioka 2004; Francke 2007). However, Toscano- (2009) listed Tityus (A.) metuendus Pocock, 1897, T. (T.) Gadea (2001) reported a single case of a post-maturation serrulatus, T. (T.) trivittatus Kraepelin, 1898, and T. (T.) molt occurring in a female Tityus (Tityus) uruguayensis uruguayensis Borelli, 1901 as the only parthenogenetic Borelli, 1901. Prior to and since the report of Toscano- scorpion species to occur in Brazil, lending further sup- Gadea (2001), no additional mention of post-maturation port to the earlier assessment of Francke (2007). molts in scorpions has been reported. Additional studies are needed to confirm the occurrence and extent of this Methods phenomenon in scorpions. Tityus stigmurus is a medium size (Total Length (TL) = On May 2008, four
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