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Litter Size in Micro-Buthoid Scorpions (Chelicerata, Scorpiones)

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Litter Size in Micro-Buthoid Scorpions (Chelicerata, Scorpiones) Boletín Sociedad Entomológica Aragonesa, n1 40 (2007) : 473− 477. LITTER SIZE IN MICRO-BUTHOID SCORPIONS (CHELICERATA, SCORPIONES) Wilson R. Lourenço Département de Systématique et Evolution, USM 0602, Section Arthropodes (Arachnologie), Muséum national d’Histoire natu- relle, CP 053, 61 rue Buffon, 75005 Paris, France − [email protected] Abstract: All available information about litter size in buthoid scorpions is presented in tabular form. Selected examples con- cerning species of micro-buthoid scorpions are given. The possible factors responsible for litter-size variation in these species are discussed. It is concluded that (i) reduction of litter size in these species is independent of ecological factors, (ii) litter size is directly proportional to the size of the female and inversely proportional to the size of the young, (iii) large body size of pro- juveniles at birth seems to be associated with a more ‘complete’ embryonic development, (iv) post-embryonic development is achieved with a smaller number of instars, (v) morphometric growth values are higher than those observed for other buthoids. Key words: Scorpiones, micro-buthoids, litter size, evolutionary factors. El tamaño de la camada en los escorpiones microbutoides (Chelicerata, Scorpiones) Resumen: Se presenta, tabulada, toda la información disponible sobre el tamaño de la camada en los escorpiones butoides. Se incluyen ejemplos escogidos de especies de escorpiones microbutoides. Se discuten los posibles factores que determinan el tamaño de las camadas, y se concluye que (i) la reducción en el tamaño de la camada de estas especies es independiente de factores ecológicos, (ii) el tamaño de la camada es directamente proporcional al tamaño de la hembra e inversamente pro- porcional al tamaño de las crías, (iii) el nacimiento de pro-juveniles de gran tamaño parece ir asociado a un desarrollo embrio- nario más “completo”, (iv) el desarrollo postembrionario se lleva a cabo en un menor número de estadios, (v) los valores del crecimiento morfométrico son más altos que los que se observan en otros butoides. Palabras clave: Scorpiones, micro-butoides, tamaño de las camadas, factores evolutivos. Introduction Scorpions are unusual among terrestrial arthropods in se- several species of buthoid scorpions have become available. veral traits of their life-history. Because of this, many as- These are summarized in Table I, with special attention pects of the reproductive biology of scorpions were poorly being given to micro-buthoid species (sensu Lourenço, understood by early authors (see Polis & Sissom, 1990 for 2000), including members the families Buthidae Koch and details). However, some precise studies on scorpion embry- Microcharmidae Lourenço. ology had already been carried out by the end of the 19th Many, if not most, of these small species of buthids century (Laurie, 1890, 1891, 1896a,b) and these were fol- and microcharmids show strongly reduced ‘average litter lowed by the publications of Pavlovsky (1924, 1925) and size’, in most cases with numbers less than 10 and in others Pflugfelder (1930). less than 7 juveniles. It is important to note that small Since the mid-1970s, there has been a renewal of in- broods have also been observed for large species of buthids, terest in the reproductive biology of scorpions and, in par- and that these numbers are frequently reported as ‘standard’ ticular, their post-embryonic development. Research on this numbers by several authors (Polis & Sissom, 1990). In fact, subject was increased during the 1980s and continued this kind of situation can be observed for females of any throughout the 1990s. Little attention, however, was paid to species which arrive to the end of their life cycle, when the the aspect of litter size in scorpions and only a few isolated number of follicles starts to decline (Lourenço, 1979). publications have provided additional information. A pio- These numbers should not be retained as ‘average’ values. neering work was that of Francke (1981), who attempted to Here I will focus on selected examples for which pre- explain the factors involved in litter size of scorpions of the cise life history data, or at least life history predictions, are family Diplocentridae. According to this author, litter size available. in diplocentrids is directly proportional to the size of the female and inversely proportional to the size of the young. Selected examples Thus, the size of the mother and the size of the young were found to account for 81 percent of the variation in litter size Microtityus Kjellesvig-Waering between the species of this family. Polis & Sissom (1990) Armas (1974) and González-Sponga (1984) provided some briefly re-discussed this matter, but referred to Francke’s biological data about Microtityus species, respectively (1981) results as being intuitive. The syntheses proposed by showing that the litter sizes of Microtityus fundorai Armas Polis & Sissom (1990) and Lourenço (1991, 2002a) only and Microtityus joseantonioi González-Sponga were par- resume the known data about litter size in scorpions, with- ticularly small, comprising only 6 or 7 juveniles on average. out further discussion about the factors that might be re- The observations carried out on M. consuelo and M. sponsible for its variability. rickyi (Lourenço et al., 1999) confirm this biological aspect. Subsequent to the publications by Polis & Sissom In both species, the broods are composed of very small (1990) and Lourenço (2002a), new data on litter size of numbers: 3 to 5 for M. consuelo and 4 to 6 for M. rickyi 473 (Fig. 1). In contrast, the size of pro-juveniles at birth is of Tityus. Only a few species of the subgenus Archaeotityus large. For example, in M. rickyi the female body size (pro- have similarly low numbers. The initial body size of pro- soma + mesosoma) averages 7 mm in length while pro- juveniles at birth is slightly larger (in relation to the juveniles body size averages 2.5 mm in length. mother’s body size) than that observed in other species of The duration of embryonic development ranged from Tityus. This more complete embryonic development may be 3 to 4 months, while the acquisition of adulthood took place correlated with the smaller number of post-embryonic instars. at an average of 229 days for M. consuelo and 249 days for The average growth rates observed for different instars M. rickyi. These developmental periods are not very diffe- are similar to those observed for several species of Tityus. rent from those in other buthids, but the growth rate values For males and females of T. (C.) elii, the growth rates were: observed between different instars are globally higher in carapace length 1.22 and 1.28; metasomal segment V length comparison with those of other species of Buthidae studied. 1.28 and 1.29; movable finger length 1.24 and 1.26. Morphometric growth values of the different instars for both the species of Microtityus studied are higher on Microbuthus fagei Vachon average than those of other studied species of the family Observations carried out on M. fagei showed that broods are Buthidae. This aspect can probably be explained by the composed of very small numbers, 3-4 (Lourenço, 2002b). In following factors: the number of molts necessary to reach contrast, the size of pro-juveniles at birth is very large. adulthood is only 4, as opposed to the 5 or 6 observed for Female body length averages 8.4 mm, while pro-juveniles other buthid species (Lourenço, 2002a); the initial body size body length averages 4.2 mm. of Microtityus pro-juveniles at birth is much larger (in rela- No precise data are available on the embryonic and tion to mother’s body size), than those observed for species postembryonic development of the species. However, using belonging to other buthoid genera (Lourenço, 2002a). I indirect methods and the morphometric values of juveniles suggest at present that the large body size of pro-juveniles at collected in nature, the number of molts necessary to reach birth could be associated with a more ‘complete’ embryonic adulthood can be estimated to be 4. development. This factor ‘may’ facilitate a strategy of post- embryonic development with a smaller number of instars. Orthochirus Karsch The theoretical morphometric growth value (constant) Observations carried out on both Orthochirus negebensis defined by Dyar (1890) and Przibram & Megusar (1912) for (Shulov & Amitai, 1960) and an undescribed species of the development of arthropods is 1:26. The values observed Orthochirus from Africa showed that broods are composed for several species of buthoids vary from 1:22 to 1:33, de- of very small numbers, 5-9 (Fig. 3). In contrast, the size of pending on the parameter (segment) measured (Lourenço, pro-juveniles at birth is very large. In both species, female 1979, 2002a). For both Microtityus consuelo and Microti- body length averages 11.5 mm, while pro-juveniles body tyus rickyi the observed growth values were: carapace length averages 4.4 mm. length 1:31 and 1:28; metasomal segment V length 1:55 and No precise data are available on the embryonic and 1:50; movable finger length 1:28 and 1:31. postembryonic development of Orthochirus species. How- ever, using indirect methods and the morphometric values Tityus (Caribetityus) elii (Armas & Marcano Fondeur) of juveniles collected in nature, the number of molts neces- With a total length of 30-35 mm, this species should not sary to reach adulthood can be estimated to be 4. really be ranged among micro-buthoid scorpions. However, some of its biological traits show similarities with micro- Ananteris coineaui Lourenço buthoids and are therefore presented here. Although many different species of the genus Ananteris The broods obtained for Tityus (Caribetityus) elii were Thorell have been described in recent years, almost nothing composed of 10 to 12 individuals. These numbers are lower is known about the life cycles of members of this genus.
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