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1 Example of a mating in lamelliform scorpions. Courtship and sperm transfer in Bothriurus burmeisteri: (A) During the courtship the male grasps the female's pedipalps with his own pedipalps. (B) Male pushing of the female during the initial part of the sperm transfer. As a result of this behaviour the lamella of the spermatophore moves down and compresses the trunk (white arrow), unfolding the capsule inside the female's genital atrium. To move his body forward appropriately, the male presses the end of his fi fth metasoma segment against a stone (black arrow) (photos courtesy of Patricia Carrera).
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This note concerns a scorpion species of the genus Buthus recently described in Algeria, Buthus apiatus Lourenço, El Bouhissi & Sadine, 2020. According to the last synthesis of Algerian scorpions, this species is located in the NorthWest of Algeria, in the department of Sidi Bel Abbes and the Tissemsilt. This note provides a new locality of B. apia...
Citations
... Additionally, scorpions offer an interesting model for investigating these topics because, in certain species, we possess substantial knowledge regarding the functional significance of numerous courtship behaviors (Peretti, 2010;Polis & Sissom, 1990). Throughout sexual interactions, individuals engage in signal exchange and various behaviors involve traits to stimulate or appease the female (Carrera et al., 2009;Chantall-Rocha & Japyassú, 2017;Lira et al., 2018;Olivero et al., 2015Olivero et al., , 2019Peretti, 2013;Peretti et al., 2001). ...
... Scorpions present indirect sperm transfer via a sclerotized spermatophore deposited in the substrate (that is regenerated each time the male mates from two chitinous halves-i.e., hemispermatophores produced in internal glandular structures called paraxial organs) (Polis & Sissom, 1990;Proctor, 1998;Weygoldt, 1990). These genital characters are incredibly complex and can be divided into subunits offering interesting opportunities for studying the evolution of genitalia Monod et al., 2017;Peretti, 2003Peretti, , 2010Peretti et al., 2001). In particular, these characters were extensively studied in the family Bothriuridae in the evolutionary framework of sexual selection (Carrera et al., 2009;Olivero et al., 2014Olivero et al., , 2015Olivero et al., , 2019Oviedo-Diego et al., 2020;Peretti, 2003Peretti, , 2010Peretti et al., 2001). ...
... These genital characters are incredibly complex and can be divided into subunits offering interesting opportunities for studying the evolution of genitalia Monod et al., 2017;Peretti, 2003Peretti, , 2010Peretti et al., 2001). In particular, these characters were extensively studied in the family Bothriuridae in the evolutionary framework of sexual selection (Carrera et al., 2009;Olivero et al., 2014Olivero et al., , 2015Olivero et al., , 2019Oviedo-Diego et al., 2020;Peretti, 2003Peretti, , 2010Peretti et al., 2001). ...
Components of the same structure or characters of the same individual might respond differently to natural and sexual selective pressures, showing complex morphological patterns. Besides, studying interactions between species plays a crucial role in understanding the diversification of sex-linked phenotypes. Specifically, when two closely related species coexist and exhibit interspecific sexual interactions (reproductive interference—IR), key traits for mating can diverge in sympatric areas to prevent interbreeding and ensure reproductive isolation (reproductive character displacement—RCD). RCD is primarily
driven by natural selection, although sexual selection pressures can alter the pattern of phenotypic variation. Additionally,
to gain a comprehensive understanding of the patterns of morphological diversification, it is essential to consider changes
related to phenotypic plasticity across environmental gradients. To date, there are no studies evaluating this topic in scorpions,
and two sympatric species (Urophonius brachycentrus and U. achalensis) with RI, provide an ideal model for evaluating
phenotypic variation across environmental gradients and the presence of RCD. In this study, we compared intra-specific
variation, as well as the size and shape of multiple characters involved in courtship and sperm transfer, between individuals
from sympatric and allopatric populations using geometric morphometrics. Our findings revealed an increase in the size of
various characters at lower temperatures (higher altitudes) for U. brachycentrus, making them more similar to heterospecifics
in sympatric areas, resulting in a pattern of morphological convergence between these species. Increased similarity between species combined with a scramble competition mating system could intensify sexual selection pressures on particular characters. Furthermore, we identified asymmetric RCD in the shape of several sexual characters crucial for mating success
(grasping structures) and sperm transfer (genital characters), which could potentially be significant for mechanical isolation
during interspecific interactions. Our results highlight significant morphological variability in the size and shape of somatic
and genital characters in two scorpion species. This variability may reflect different evolutionary responses, driven in part
by natural selection pressures associated with geographic and environmental variations and species recognition mechanisms,
and in part by sexual selection pressures at both the intra- and interspecific levels. This comprehensive study reveals the
complexity of evolving multifunctional traits in an understudied model and offers valuable insights into traits subject to
multiple selective pressures in animal systems experiencing RI.
... Furthermore, during an elaborate courtship, both sexes displayed unique characters with functional roles such as stimulation or increased female receptivity with non-genital contact structures (e.g., the caudal gland in 'rubbing with telson', the sting in 'sexual sting') or grasping characters to overcome female resistance (e.g., apophyses in pedipalps, chelicerae) (Polis & Sissom, 1990;Carrera et al., 2009;Peretti, 2001). In particular, these characters were extensively studied in the family Bothriuridae in the evolutionary framework of sexual selection (Peretti et al., 2001;Carrera et al., 2009;Olivero et al., 2014Olivero et al., , 2019Peretti, 2010). Lastly, scorpions present indirect sperm transfer via a sclerotized spermatophore deposited in the substrate (Weygoldt, 1990;Proctor, 1998). ...
... This spermatophore is regenerated each time the male mates from two chitinous halves (i.e., hemispermatophores) produced in internal glandular structures called paraxial organs (Polis & Sissom, 1990). These genital characters are incredibly complex and can be divided into subunits offering interesting opportunities for studying the evolution of genitalia (Peretti et al., 2001;Peretti, 2003Peretti, , 2010Mattoni et al., 2012;Monod et al., 2017). For example, some characters follow a distinctive pattern of characters under sexual selection pressures (i.e., evolve rapidly and divergently), while others show only minor variations coinciding with what is predicted for characters under natural selection, such as structures with mechanical constraints or with key reproductive functions such as sperm passage (Peretti, 2010;Mattoni et al., 2012). ...
... These genital characters are incredibly complex and can be divided into subunits offering interesting opportunities for studying the evolution of genitalia (Peretti et al., 2001;Peretti, 2003Peretti, , 2010Mattoni et al., 2012;Monod et al., 2017). For example, some characters follow a distinctive pattern of characters under sexual selection pressures (i.e., evolve rapidly and divergently), while others show only minor variations coinciding with what is predicted for characters under natural selection, such as structures with mechanical constraints or with key reproductive functions such as sperm passage (Peretti, 2010;Mattoni et al., 2012). The morphological diversity of sexual characters and spermatophores of scorpions responds to diverse (and not mutually exclusive) evolutionary hypotheses (Peretti, 2010). ...
Reproductive interference (RI) can occur when two related species coexist in sympatry, involving sexual attraction, mating, and even hybridization between heterospecifics. Consequently, reproductive key characters of these species may suffer morphological shifts in sympatry to avoid the success of heterospecific sexual interactions, a phenomenon known as reproductive character displacement (RCD). RCD can be promoted by natural selection, although sexual selection pressures can act synergistically or agonistically so that phenotypic variation can respond in different directions and magnitudes to these forces. In turn, the size and shape of characters may respond differentially (mosaic evolution) to these pressures, so the analysis of multiple dimensions in traits is essential to understand the complexity of their phenotypic variability. To date, there are no studies evaluating this topic in scorpions, and two species ( Urophonius brachycentrus and U. achalensis ) sympatric and synchronous with RI represent an ideal model to evaluate the phenotypic variation and occurrence of RCD. In addition, the populations of these species are found in an altitudinal cline, so environmental factors may also be responsible for explaining their morphological variation. We compared the intra-specific variation, the size and shape of multiple characters involved in courtship, and sperm transfer in individuals from sympatric and allopatric populations using geometric morphometrics. We found asymmetric RCD of several sexual characters for courtship success (grasping structures) and sperm transfer (genital characters). This would evidence the action of natural selection pressures and the existence of a possible mechanism to avoid heterospecific mating success. In addition, we found a pattern of asymmetric morphological variation where one species in the sympatric zone suffered an increase in size in several characters due to environmental factors (pattern of morphological convergence). The convergence of characters combined with RI and a scramble competition mating system could intensify sexual selection pressures on specific characters, which was reflected in their high coefficients of variation. Our results suggest that in this sympatric zone, several selective regimes act differentially on various dimensions of the characters evaluated, which would support a possible mosaic evolution. This comprehensive study illuminates the complexity inherent in the evolution of multi-functional traits in a previously unexplored model, providing novel insights for evaluating traits under multiple selective pressures in animal systems experimenting RI.
... Scorpion spermatozoa consist of an acrosomal vacuole and filament, a long filiform nucleus, an elongated mitochondrial segment, and an axoneme with a typical 9+2 or an abnormal 9+0 [3]. [4]. ...
... In our study, it can be said that it consists of 3 main parts morphologically. When compared with the electron microscope image in Peretti's study [4], the spermatozoa structure matches. In the review by Vignoli et al. [1], they reported that scorpion semen had cytoplasmic droplets in the head and middle part. ...
Scorpions are venomous arthropods, members of the order Arachnida and the order Scorpions. Scorpions and their venom have been used in China, India and Africa for years to treat various diseases. In recent years, there are studies showing that scorpion venoms and toxins can reduce cancerous growth, induce apoptosis and prevent metastasis
... Examples of such structures include the mating plugs typically produced by males, whose main function is to obstruct the female genitalia against new sperm depositions (Garcilazo-Cruz & Alvarez-Padilla, 2015), which could increase their reproductive success by mechanically hindering females from mating with subsequent males (Uhl et al. 2014). Amorphous secretory materials, and male somatic or genital organs are all potential components of mating plugs (Drummond, 1984;Eberhard, 1996;Birkhead & Møller, 1998;Simmons, 2001;Oh & Hankin, 2004;Peretti, 2010;Uhl, Nessler, & Schneider, 2010). In the current study, while examining the specimens of Ketambea falcata sp. ...
An extensive survey of Linyphiidae spiders during 1988–
2006 in the Gaoligong Mountains and the adjacent areas
of Yunnan Province revealed 110 species belonging to 53
genera, including seven new genera and 76 new species:
Absconditus gen. nov., Absconditus acerus sp. nov. (♂♀);
Amfractus gen. nov., A. dentefaberis sp. nov. (♂♀); Genus
Bathyphantes Blackwall, 1859, A. acutala sp. nov. (♂♀); A.
thailandica Barrion & Litsinger, 1995 (♀); Auricula gen.
nov., A. aeda sp. nov. (♂♀), A. rutunda sp. nov. (♀), A.
sanchaheensis sp. nov. (♀), A. triangulara sp. nov. (♂♀);
Genus Bathyphantes Blackwall, 1859, B. ansulis sp. nov.
(♂♀), B. gracilis (Blackwall, 1841) (♂), B. longiscapus sp.
nov. (♂♀), B. magnis sp. nov. (♂♀); Genus Bolyphantes C.
L. Koch, 1837, B. lishadiensis sp. nov. (♀), B. lushuiensis
sp. nov. (♀); Genus Capsulia Saaristo, Tu & Li, 2006, C.
tianmushana (Chen & Song, 1987) (♂♀); Genus Caviphantes
Oi, 1960, C. catomidius sp. nov. (♂♀), C. pseudosaxetorum
Wunderlich, 1979 (♀); Genus Ceratinella Emerton, 1882,
C. acutalum sp. nov. (♂); Genus Cirrosus Zhao & Li, 2014,
C. atrocaudatus Zhao & Li, 2014 (♀); Genus Collinsia O.
Pickard-Cambridge, 1913, C. denticulata sp. nov. (♂♀),
C. inerrans (O. Pickard-Cambridge, 1885) (♂♀); Genus
Conglin Zhao & Li, 2014, C. personatus Zhao and Li, 2014
(♀); Cristatus gen. nov., C. anfractus sp. nov. (♂♀), C.
makuensis sp. nov. (♂♀); Genus Curtimeticus Zhao & Li,
2014, C. rutundus sp. nov. (♀); Genus Erigone Audouin,
1826, E. ansula sp. nov. (♂♀), E. atra Blackwall, 1833
(♂♀), E. prominens Bösenberg & Strand, 1906 (♂♀), E.
sinensis Schenkel, 1936 (♂♀); Genus Frontinellina van
Helsdingen, 1969 (new to China), F. gemalakaensis sp.
nov. (♀); Genus Gnathonarium Karsch, 1881, G. dentatum
(Wider, 1834) (♂♀), G. gibberum Oi, 1960 (♂♀), G.
taczanowskii (O. Pickard-Cambridge, 1873) (♀); Genus
Gongylidiellum Simon, 1884, G. acerosus sp. nov. (♂);
Genus Gongylidioides Oi, 1960, G. laqueus sp. nov. (♀),
G. lingulatus sp. nov. (♀); Genus Gongylidium Menge,
1868, G. bifurcatus sp. nov. (♂♀), G. manibus sp. nov. (♂);
Gracilentus gen. nov., G. denticulatus sp. nov. (♂♀), G.
serratus sp. nov. (♀), G. tenchongensis sp. nov. (♀); Genus
Himalaphantes Tanasevitch, 1992, H. aduncus sp. nov.
(♂♀), H. azumiensis (Oi, 1979) (♂♀), H. auriculus sp. nov.
(♀), H. fugongensis sp. nov. (♀), H. gyratus sp. nov. (♀), H.
pseudoaduncus sp. nov. (♂), H. pulae sp. nov. (♀); Genus
Houshenzinus Tanasevitch, 2006, H. tengchongensis Irfan &
Peng, 2018 (♂♀) (male new to science); Genus Hylyphantes
Simon, 1884, H. graminicola (Sundevall, 1830) (♂♀), H.
spirellus Tu & Li, 2005 (♂); Genus Ketambea Millidge &
Russell-Smith, 1992, K. acuta Tanasevitch, 2017 (♂♀), K.
aseptifera sp. nov. (♀), K. falcata sp. nov. (♀), K. septifera
sp. nov. (♀); Genus Laogone Tanasevitch, 2014, L. lunata
Zhao & Li, 2014 (♂♀); Genus Lepthyphantes Menge, 1866,
L. cordis sp. nov. (♂♀), L. serratus sp. nov. (♂); Genus
Linyphia Latreille, 1804, L. gaoshidongensis sp. nov. (♀), L.
pengdangensis sp. nov. (♀), L. triangularis (Clerck, 1757)
(♀); Lutosus gen. nov., L. projectis sp. nov. (♂); Genus
Megalepthyphantes Wunderlich, 1994, M. gongshanensis
sp. nov. (♂♀), M. sanchaheensis sp. nov. (♀); Genus
Mermessus O. Pickard-Cambridge, 1899, M. datangensis
sp. nov. (♂♀); Genus Microlinyphia Gerhardt, 1928, M.
pusilla (Sundevall, 1830) (♂♀), M. rehaiensis sp. nov.
(♀), M. spirala sp. nov. (♀); Genus Molestia Tu, Saaristo
& Li, 2006, M. ancorarius sp. nov. (♀), M. caudatus sp.
nov. (♂♀), M. hamifer (Simon, 1884) comb. nov. (♂♀),
M. yaojiapingensis sp. nov. (♀); Genus Mughiphantes
Saaristo & Tanasevitch, 1999, M. latus sp. nov. (♂♀); Genus
Nasoona Locket, 1982, N. crucifera (Thorell, 1895) (♀);
Genus Nasoonaria Wunderlich & Song, 1995, N. orthogonia
sp. nov. (♂♀), N. sinensis Wunderlich & Song, 1995 (♂♀),
N. yunnanensis (Xia, Zhang, Gao, Fei & kim, 2001) comb.
nov.; Genus Nematogmus Simon, 1884, N. cikaiensis sp.
nov. (♀), N. lushuiensis sp. nov. (♀), N.membranifer Song
& Li, 2008 (♂♀); Genus Nippononeta Eskov, 1992, N.
coreana (Paik, 1991) (♂♀); Genus Oia Wunderlich, 1973, O.
ceratinum sp. nov. (♂♀); Genus Paikiniana Eskov, 1992, P.
lurida (♂♀); Genus Parameioneta Locket, 1982, P. bilobata
Li & Zhu, 1993 (♂♀), P. tricolorata Zhao & Li, 2014 (♀);
Genus Porrhomma Simon, 1884, P. longjiangense Zhu &
Wang, 1983 (♀); Genus Prosoponoides Millidge & Russell-
Smith, 1992, P. bangbieensis sp. nov. (♀), P. corneus sp.
nov. (♀), P. guanduensis sp. nov. (♀), P. longiprojectus sp.
nov. (♂♀), P. longyangensis sp. nov. (♀), P. minutus sp.
nov. (♂♀), P. pianmaensis sp. nov. (♀), P. sinensis (Chen,
1991) (♂♀), P. yakouensis sp. nov. (♀), P. yani sp. nov.
(♀), P. yapingensis sp. nov. (♂♀), P. yunnanensis sp. nov.
(♀); Genus Ryojius Saito & Ono, 2001, R. furcatus sp. nov.
(♂♀); Sinisterigone gen. nov., S. circularis sp. nov. (♂♀),
S. incurvatis sp. nov. (♀), S. rutundis sp. nov. (♀); Genus
Stemonyphantes Menge, 1866, S. bifurcatus sp. nov. (♂♀);
Genus TaibaishanusTanasevitch, 2006, T. nankangensis sp.
nov. (♂); Genus Tapinopa Westring, 1851, T. longicisterna
sp. nov. (♀); Genus Tchatkalophantes Tanasevitch, 2001,
T. lingulatis sp. nov. (♀); Genus Theoa Saaristo, 1995, T.
hamata Tanasevitch, 2014 (♂) (new to China); Genus
Ummeliata Strand, 1942, U. feminea (Bösenberg & Strand,
1906) (♂♀), U. insecticeps (Bösenberg & Strand, 1906)
(♂♀); Genus Vietnagone Tanasevitch, 2019, V. denticulata
sp. nov. (♂♀); Genus Walckenaeria Blackwall, 1833, W.
circulara sp. nov. (♂♀). Among new species described here,
forty five are known by single sex. The following genus
and species are recorded for the first time in China: Genus
Frontinellina van Helsdingen, 1969 and three species Atypena
thailandica Barrion & Litsinger, 1995 (♀), Ketambea acuta
Tanasevitch, 2017 (♂♀) and Theoa hamata Tanasevitch,
2014 (♂). The taxonomic placements of Lepthyphantes
hamifer Simon, 1884, Lepthyphantes zhangmuensis Hu,
2001 and Walckenaeria yunnanensis Xia, Zhang, Gao, Fei
& kim, 2001 are revised and proposed here as: Molestia
hamifer (Simon, 1884) comb. nov. (♂♀), Nasoonaria
yunnanensis (Xia, Zhang, Gao, Fei & Kim, 2001) comb.
nov. and Himalaphantes zhangmuensis (Hu, 2001) comb.
nov., respectively. Morphological descriptions of all new
species, including photos of body and copulatory organs for
all species, as well as the locality maps are provided.
... The spermatozoa are filiform and flagellate, comprising a head, middle piece, and flagellum (Jespersen and Hartwick, 1973;Alberti, 2000Alberti, , 2005Michalik and Mercati, 2010). Structural differences in the axoneme or the acrosomal complex, between Buthidae C.L. Koch, 1837, and other extant families, suggest variation in the morphology of scorpion sperm may be phylogenetically informative (Vignoli et al., 2008;Michalik and Mercati, 2010;Peretti, 2010). ...
... For the first time, Vrech et al. (2011) analyzed various measurements of individual sperm packages, investigated intraspecific variation, and discussed the utility of sperm packages as characters for scorpion systematics. Based in part on differences in sperm package morphology, Vrech et al. (2011) recommended elevating the subgenus Andibothriurus Maury, 1975, long recognized based on other characters (Mattoni, 2003;Peretti, 2010), to the rank of genus. Vrech et al. (2011Vrech et al. ( , 2016 also described abnormalities in the shape of sperm packages and the number of spermatozoa inside male ejaculates. ...
... Sperm Package Morphology: The general absence of sperm packages in Buthidae is well known (Cruz-Landim and Ferreira, 1973;Peretti and Battán-Horenstein, 2003; Mercati, 2010; Peretti, 2010;Vrech et al., 2011; but see Alberti, 1983). For example, Peretti and Battán-Horenstein (2003) reported the absence of sperm conjugation in Zabius fuscus (Thorell, 1876) and suggested a hypothesis, concerning a difference in sperm transfer between Buthidae and Bothriuridae, which could account for the absence or presence of sperm packages, respectively. ...
The spermatozoa of scorpions are often bundled together, forming a type of sperm conjugation known as a sperm package. Sperm packages may be found inside the testes and seminal vesicles but vanish in the female atrium, leaving free spermatozoa. Previous studies, based on a limited number of taxa, suggested a diversity of sperm package morphology across the order Scorpiones C.L. Koch, 1850. However, the sperm packages of most scorpion taxa remained unknown. The present study provides the first systematic survey of sperm package morphology across the order, covering 89 exemplar species in 66 genera and 19 families representing all suprafamilial ranks, with a more detailed investigation of the family Bothriuridae Simon, 1880. Whereas all exemplar species of scorpions exhibit sperm packages, Buthida Soleglad and Fet, 2003, including Chaerilidae Pocock, 1893, and most Buthidae C.L. Koch, 1837, present unorganized sperm or loosely organized bundles. Although the details vary, three main types of sperm packages may be recognized in all other families: single folded; straight; and multiple folded. Subtypes may be identified according to general shape and folding patterns, mainly among sperm packages that are folded multiple times. Single-folded sperm packages are the most common type observed in the order. Although most sperm packages lack a covering, a conspicuous secretion sheath may be evident, e.g., in some Chactidae Pocock, 1893. Sperm packages vary in length from 112354 m and bent sperm packages are not necessarily longer than straight sperm packages. Four exemplar species of Bothriuridae reveal that variation in sperm count within a single sperm package is consistent with the count derived in spermatogenesis. The diversity of sperm packages suggests a path from free spermatozoa, via bent sperm packages, to other forms. Sperm packages may aid in the transport, cooperation, competition, and survival of spermatozoa. The diverse morphology, function, and evolution of sperm packages merit further investigation.
... This spermatophore is formed by joining two halves (hemispermatophores), generated in the paraxial organs of the male reproductive system (Francke, 1979). The hemispermatophores then join just as the spermatophore emerges from the male gonopore, right before sperm transfer (Hjelle, 1990;Farley, 2001;Peretti, 2010). The production of each hemispermatophore by a different paraxial organ could lead to differences between the structures of the right and left sides of the spermatophore (Peretti, 2000(Peretti, , 2010. ...
... The hemispermatophores then join just as the spermatophore emerges from the male gonopore, right before sperm transfer (Hjelle, 1990;Farley, 2001;Peretti, 2010). The production of each hemispermatophore by a different paraxial organ could lead to differences between the structures of the right and left sides of the spermatophore (Peretti, 2000(Peretti, , 2010. Consequently, the environmental stress could affect these structures causing asymmetry in the spermatophore (Peretti et al., 2001;Olivero et al., 2014). ...
... The male scorpion reproductive system, including the hemispermatophores, is produced during a precise stage of the organism's life, when it reaches adulthood (Francke, 1979;Benton, 2001;Peretti, 2003). Furthermore, in male scorpions, the adult should produce and regenerate both hemispermatophores in a much shorter time period (close to one week) after each mating (Peretti, 2003(Peretti, , 2010Vrech et al., 2019). This means that these types of structures are not exposed to developmental stress during all the organism's life, but for shorter periods of time. ...
Urbanization causes the loss of large amounts of habitat and produces significant changes in environmental conditions with consequences in the individual's behavior, morphology, and physiology in natural populations. The urbanization can impact in the individual's development instability (DI) that may be measured with the fluctuating asymmetry (FA) level making it an environmental quality bioindicator. In addition, the FA of each trait should be interpreted considering the trait functional importance, morphogenesis and history of selection pressures. In this paper, we evaluated the impact of two decades of increasing urbanization levels over a population of the pampean scorpion Bothriurus bonariensis. We analyze the FA level in somatic and genital traits in adult individuals of both sexes, collected every ~ 10 years (1997, 2008, 2018). Interestingly, in this study the somatic traits (pedipalp chelae) would be under directional selection and are shaped throughout the individual's growth. In contrast, the genital traits (male's hemispermatophore) are involved in the sperm transfer process and could be under stabilizing selection. Also, hemispermatophores are affected for less time by environmental stressors since they are formed once the individual has reached sexual maturity. Our results showed a significant increase in the FA levels in somatic traits of males and females, according to the increase of urbanization along years. In contrast, genital traits showed no changes in the level of FA in females, and only a few male genital traits presented an increase in the FA level. This work shows evidence that the measured scorpion traits (under different selective pressures and morphogenesis) might present differences in their susceptibility to environmental stress. We discuss these results in a context of DI of individuals caused by urbanization. We support the idea that the level of FA can be useful as an index to evaluate the effects of environmental stress on invertebrate populations. Furthermore, our paper presents the scorpions as a good model that could be used as an indicator group for studying the impact of the anthropogenic disturbance factors on natural ecosystems.
... During the sperm transfer, sperm is expulsed from the spermatophore passing through the female genital aperture into the genital atrium (covered externally by the genital operculum) (Francke, 1979;Hjelle, 1990;Polis & Sissom, 1990) and finally reach the seminal receptacles or spermathecae (Peretti & Battán-Horenstein, 2003;Volschenk, Mattoni, & Prendini, 2008). Considerable diversity of mating plugs exists in scorpions, many of which have a male origin from the components of the spermatophore, sometimes including glandular secretions or sperm (see Peretti, 2010 for a review). Mating plugs have been described for Urodacidae (Prendini, 2000;Shorthouse & Marples, 1982;Smith, 1966;Stockwell, 1989), Euscorpiidae (Althaus et al., 2010;Angermann, 1957;Jacob et al., 2004), Hemiscorpiidae (Monod, Cauwet, González-Santillán, & Huber, 2017), Chactidae (Monod et al., 2017;Stockwell, 1989), Buthidae (Bucherl, 1956;Probst, 1972;Shulov & Amitai, 1958), Vaejovidae (Ayrey, Jones, & Myers, 2019;Contreras-Garduño et al., 2005;Fox, 1975;Hendrixson, 2001;Monod et al., 2017;Sissom & Stockwell, 1991;Stockwell, 1989), Ischnuridae (Monod et al., 2017) and Bothriuridae (Carrera, 2008;Mattoni & Peretti, 2004;Peretti, 2003Peretti, , 2010Stockwell, 1989). ...
... Considerable diversity of mating plugs exists in scorpions, many of which have a male origin from the components of the spermatophore, sometimes including glandular secretions or sperm (see Peretti, 2010 for a review). Mating plugs have been described for Urodacidae (Prendini, 2000;Shorthouse & Marples, 1982;Smith, 1966;Stockwell, 1989), Euscorpiidae (Althaus et al., 2010;Angermann, 1957;Jacob et al., 2004), Hemiscorpiidae (Monod, Cauwet, González-Santillán, & Huber, 2017), Chactidae (Monod et al., 2017;Stockwell, 1989), Buthidae (Bucherl, 1956;Probst, 1972;Shulov & Amitai, 1958), Vaejovidae (Ayrey, Jones, & Myers, 2019;Contreras-Garduño et al., 2005;Fox, 1975;Hendrixson, 2001;Monod et al., 2017;Sissom & Stockwell, 1991;Stockwell, 1989), Ischnuridae (Monod et al., 2017) and Bothriuridae (Carrera, 2008;Mattoni & Peretti, 2004;Peretti, 2003Peretti, , 2010Stockwell, 1989). Among the principal types of scorpion mating plugs, we find "amorphous" or gel-like plugs formed by sperm or accessory gland substances from male paraxial organs (Althaus et al., 2010;Castelvetri & Peretti, 1999;Peretti, 2003Peretti, , 2010Peretti & Battán-Horenstein, 2003), as well as "sclerotized" plugs derived from one or more cuticular portions that detach from the spermatophore (Contreras-Garduño et al., 2005;Mattoni & Peretti, 2004;Peretti, 2003Peretti, , 2010. ...
... Mating plugs have been described for Urodacidae (Prendini, 2000;Shorthouse & Marples, 1982;Smith, 1966;Stockwell, 1989), Euscorpiidae (Althaus et al., 2010;Angermann, 1957;Jacob et al., 2004), Hemiscorpiidae (Monod, Cauwet, González-Santillán, & Huber, 2017), Chactidae (Monod et al., 2017;Stockwell, 1989), Buthidae (Bucherl, 1956;Probst, 1972;Shulov & Amitai, 1958), Vaejovidae (Ayrey, Jones, & Myers, 2019;Contreras-Garduño et al., 2005;Fox, 1975;Hendrixson, 2001;Monod et al., 2017;Sissom & Stockwell, 1991;Stockwell, 1989), Ischnuridae (Monod et al., 2017) and Bothriuridae (Carrera, 2008;Mattoni & Peretti, 2004;Peretti, 2003Peretti, , 2010Stockwell, 1989). Among the principal types of scorpion mating plugs, we find "amorphous" or gel-like plugs formed by sperm or accessory gland substances from male paraxial organs (Althaus et al., 2010;Castelvetri & Peretti, 1999;Peretti, 2003Peretti, , 2010Peretti & Battán-Horenstein, 2003), as well as "sclerotized" plugs derived from one or more cuticular portions that detach from the spermatophore (Contreras-Garduño et al., 2005;Mattoni & Peretti, 2004;Peretti, 2003Peretti, , 2010. ...
Mating plugs have been proposed as a mechanism that has evolved to avoid sperm competition. Their structure and composition vary across taxa and are related to the effectiveness of its function. This effectiveness could be related to different evolutionary interests of the sexes. Urophonius brachycentrus and Urophonius achalensis (Scorpiones, Bothriuridae) are highly suitable species to study mating plugs because both are monandrous species with specific morphological and physiological responses in the female's genitalia. Here, we analyze (a) the morphology and fine structure of the mating plugs of both species, (b) the site of production in males and the formation process of the mating plug, and (c) the changes that it undergoes over time in the female's reproductive tract. In both species, a complex mating plug obliterates the female's genital aperture and fills the genital atrium. We observed considerable interspecific variation in the mating plug morphology. A mating hemi‐plug was found surrounding the capsular lobes of the hemispermatophore, which could have a mixed composition (involving portions of the hemispermatophore and glandular products). The glandular portion was transferred in a semi‐solid state filling the female's genital atrium and then hardening. Changes that the plug undergoes in the female's genitalia (darkening and increase of the “distal” area of the plug) indicate a participation of the female to the formation of this type of plug. Our study provides new insights into the plugging phenomenon in scorpions, and we discussed the adaptive significance as a post‐copulatory mechanism to avoid sperm competition.
... And the predicted differences in spermatophore complexity in some small arthropods such as pseudoscorpions (Weygeldt 1969) and collembolans (Mayer 1957), in which males never contact females (and species specificity would thus seem especially important) is absent. Their spermatophores are less rather than more complex compared to the spermatophores of others (other pseudoscorpions, amblypygids, scorpions) in which males find and hold females prior to depositing the spermatophore (Weygeldt 1969(Weygeldt , 1999Peretti 2010). ...
Rapid divergence in external genital structures occurs in nearly all animal groups that practice internal insemination; explaining this pattern is a major challenge in evolutionary biology. The hypothesis that species‐specific differences in male genitalia evolved under sexual selection as courtship devices to influence cryptic female choice (CFC), has been slow to be accepted. Doubts may stem from its radical departure from previous ideas, observational difficulties because crucial events occur hidden within the female's body, and alternative hypotheses involving biologically important phenomena such as speciation, sperm competition, and male‐female conflicts of interest. We assess the current status of the CFC hypothesis by reviewing data from two groups in which crucial predictions have been especially well‐tested, Glossina tsetse flies and Roeseliana (formerly Metrioptera) roeselii bushcrickets. Eighteen CFC predictions have been confirmed in Glossina and nineteen in Roeseliana. We found data justifying rejection of alternative hypotheses, but none that contradicted CFC predictions. The number and extent of tests confirming predictions of the CFC hypothesis in these species is greater than that for other generally accepted hypotheses regarding the functions of non‐genital structures. By this criterion, it is reasonable to conclude that some genital structures in both groups likely involved sexual selection by CFC. This article is protected by copyright. All rights reserved
... The male production of mating plugs is a widespread phenomenon in the animal kingdom and has evolved independently many times (Shine, Olsson, & Mason, 2000). Mating plugs are diverse in structure and effectiveness, being generally formed by male substances, such as coagulation of ejaculate, or portions of genitalia (Peretti, 2010;Simmons, 2001;Uhl, Nessler, & Schneider, 2010;Wigby & Chapman, 2004). Sometimes, females may also have a role in mating plug formation (Aisenberg & Eberhard, 2009;Eberhard, 1996). ...
... Studies in several species of scorpions show that they can use chemical signaling to find potential mates and begin courtship (Gaffin & Brownell, 1992Melville, Tallarovic, & Brownell, 2003;Steinmetz, Bost, & Gaffin, 2004;Taylor, Cosper, & Gaffin, 2012). In many scorpion species, males transfer a mating plug after sperm transfer and this plug may be of different types and effectiveness (Peretti, 2010). Although much effort has been put into analyzing pheromonal communication and mating plug morphology, the evaluation of the capability of males to detect the female's mating status, and the relationship between the presence of a mating plug and the female's pheromone emission has been neglected in scorpions. ...
... Urophonius brachycentrus (Thorell, 1876) represents a good model organism for this analysis because the females present a mating plug after insemination (Carrera, 2008;Peretti, 2010). This mating plug is detached from the male spermatophore and cannot be expelled by the female or removed by other males. ...
After mating, females may experience a decline in sexual receptivity and attractive‐
ness that may be associated with changes in the production and emission of sex
pheromones. In some cases, these changes are produced by chemical substances or
structures (e.g., mating plugs) produced by males as a strategy to avoid or reduce
sperm competition. In scorpions, sex pheromones may be involved in finding poten‐
tial mates and starting courtship. Here, we tested the hypothesis that the males of
Urophonius brachycentrus, a species that produces a mating plug, use chemical com‐
munication (sex pheromones) to detect, localize, and discriminate females according
to their mating status (virgin or inseminated), aided by chemical signaling. We also
explored the effect of extracting of the mating plug on chemical communication and
mating acceptance. We used Y‐maze olfactometers with different stimuli to analyze
male choice and exploration time. To evaluate mating acceptance, we measured the
attractiveness and receptivity of females of different mating status. We found that
chemical communication occurs through volatile pheromones, but not contact pher‐
omones. Males equally preferred sites with virgin or inseminated females with re‐
moved mating plug. In turn, females with these mating statuses were more attractive
and receptive for males than inseminated females. This study suggests that the mat‐
ing plug significantly affects female chemical attractiveness with an effect on volatile
pheromones and decreasing sexual mating acceptance of females. The decline in the
female's sexual receptivity is a complex process that may respond to several non‐ex‐
clusive mechanisms imposed by males and strategically modulated by females.
... After fertilization, the viviparous embryos develop within the ovariuterus until the time of parturition [73][74]. In many species of scorpions, females present a genital plug after the sperm transfer [32], although their morphology is very diverse and the function is discussed [31]. In some cases, the formation of the genital plug is almost completely attributed to the male [32,39,43], although the participation of the female has been suggested [31-32, 39, 75-76]. ...
... In many species of scorpions, females present a genital plug after the sperm transfer [32], although their morphology is very diverse and the function is discussed [31]. In some cases, the formation of the genital plug is almost completely attributed to the male [32,39,43], although the participation of the female has been suggested [31-32, 39, 75-76]. In ultrastructural studies of the atrial epithelium of the female, pores and glandular cells and high secretory activity have been described, so the possibility of female participation is strongly expected [38, [76][77]. ...
... In this work, females of three scorpion species are compared. Urophonius brachycentrus (Thorell, 1877) and U. achalensis (Á balos and Hominal, 1974) [31,78] two species of the family Bothriuridae that have genital plugs were compared with Zabius fuscus (Thorell 1877), a buthid species with no genital plug [32]. Urophonius has a 'mixed' plug with a combination of detachable portions of the spermatophore and glandular substances [31]. ...
Immune defense is a key feature in the life history of organisms, expensive to maintain, highly regulated by individuals and exposed to physiological and evolutionary trade-offs. In chelicerates, relatively scarce are the studies that relate postcopulatory mechanisms and immune response parameters. This work makes an approximation to the female’s immunological consequences produced after the placement of a foreign body in the genitalia of three scorpions species, two species that normally receive genital plugs during mating (Urophonius brachycentrus and U. achalensis) and one that does not (Zabius fuscus). Here we performed the first morphological description of the natural plugs of the two Urophonius species. We described complex three zoned structure anchored to the female genital atrium and based on this information we placed implants in the genitalia (for eliciting the local immune response) of virgin females of the three species and measured the immune encapsulation response to this foreign body. We found a greater and heterogeneous response in different zones of the implants in the plug producing species. To corroborate the specificity of this immune response, we compared the local genital reaction with the triggered response at a systemic level by inserting implants into the female body cavity of U. brachycentrus and Zabius fuscus. We found that the systemic response did not differ between species and that only in the plug producing species the local response in the genitalia was higher than the systemic one. We also compared the total hemocyte load before and after the genital implantation to see if this parameter was compromised by the immunological challenge. We confirmed that in Urophonius species the presence of a strange body in the genitalia caused a decrease in the hemocyte load. Besides, we find correlations between the body weight and the immunological parameters, as well as between different immunological parameters with each other. Complementarily, we characterized the hemocytes of the three scorpion species for the first time. This comparative study can help to provide a wider framework of the immunological characteristics of the species, their differences and their relationship with the particular postcopulatory mechanism such as the genital plugs.
