Tergal and Sexual Anomalies in Bothriurid Scorpions (Scorpiones, Bothriuridae)
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2005. The Journal of Arachnology 33:622±628 TERGAL AND SEXUAL ANOMALIES IN BOTHRIURID SCORPIONS (SCORPIONES, BOTHRIURIDAE) Camilo I. Mattoni: Division of Invertebrate Zoology, American Museum of Natural History, Central Park West at 79th Street, New York, NY 10024-5192, USA. E-mail: [email protected] ABSTRACT. New data concerning developmental anomalies observed among species of the family Bothriuridae (Scorpiones) are presented. Tergal malformations in Bothriurus coriaceus, Brachistosternus roigalsinai and Bothriurus noa are described and illustrated. Two new cases of intersexuality in scorpions, in specimens of Brachistosternus pentheri and Bothriurus araguayae, are reported and discussed. Keywords: Developmental anomalies, tergites, intersexuality, Scorpiones, Bothriuridae There are numerous reports of developmen- Expression of both states in one specimen is tal anomalies in scorpions (Table 1). Most re- clearly an abnormality and the existence of ports relate to the duplication of posterior rare abnormalities does not necessitate the body segments (Vachon 1952; Hjelle 1990; need to abandon these character systems. Fur- Sissom & Shelley 1995, see the latter for thermore, many systematists have observed overview); reference to other types of devel- this kind of variation on the chelicerae of sev- opmental anomalies in scorpions is scarce. eral species, including cases where both che- One work includes information about anom- licerae are different from the usual morphol- alies of the legs and pedipalps (Armas 1977) ogy of the species (Mattoni 2003; Prendini and another describes a tergal and two cara- pers. comm.). They would not consider these pacial malformations (Armas 1976). Most re- represent any obstacle for identifying taxa in cently, Teruel (2004) presented a list and brief question, because these occurrences are rare description of tergal (see below) and pedipal- in scorpion populations, and there are many pal anomalies; however, only Armas (1977) additional characters that can assist with an illustrated the anomalies described, a prereq- identi®cation. uisite for understanding the anomalies report- The only references to tergal anomalies in ed. scorpions are the works of Armas (1976), who Teruel (2004: 237) references a ``cheliceral described a specimen of Didymocentrus trin- anomaly'' on one specimen of a buthid, Ly- itarius Franganillo 1930 (Diplocentridae) with chas obsti Kraepelin 1913. This specimen has fusion of the carapace and the ®rst tergite, and two teeth on the ventral surface of the ®xed Teruel (2004) who described anomalies in one ®nger of one chelicera and one on the other, male of Microtityus jaumei Armas 1974 the latter expression being the typical condi- (Buthidae), that possessed a double anomaly, tion for Lychas (Vachon 1963; KovarÏõÂk 1997). with tergite V completely divided on the pos- Teruel (2004) notes that this is interesting terior half, and tergite VII fused dorsally to from a taxonomic viewpoint, because, tradi- metasomal segment I. Teruel (2004) also re- tionally, the number of ventral teeth on the ported two female diplocentrids (Cazierius ®xed ®nger has been used as a strong char- parvus Armas 1984 and C. gundlachii (Karsch acter in the generic differentiation of the scor- 1880) and one euscorpiid (Euscorpius ¯avi- pions of the family Buthidae (Kraepelin 1899; caudis (DeGeer 1778)), possessing totally di- Sissom 1990). Teruel (2004) suggested that vided tergites. using this character to identify buthids from The references to sexual malformations are Northwest Africa could present problems, be- restricted to 5 reports, involving hermaphro- cause it could cause erroneous identi®cations. ditism (with male and female genitalia), gyn- The two states clearly represent normal vari- andromorphism (with both sexes discretely ation in morphology, and are not anomalous. combined) and intersexualism (where the en- 622 MATTONIÐANOMALIES IN SCORPIONS 623 Figures 1±3.ÐCarapace and tergites of malformed scorpions. 1±2. Females of Bothriurus coriaceus;1. specimen from 4 km N Los Vilos; 2. specimen from Cuesta de Chacabuco. 3. female of Brachistosternus roigalsinai, carapace and tergites. Scale 5 5 mm. tire body is intermediate between sexes). Mat- aly reported by Teruel (2004) on Centromach- thiesen (1968) described a hermaphrodite etes pocockii (Kraepelin 1894) and Urophon- specimen of the buthid Tityus bahiensis (Perty ius granulatus Pocock 1898, are the only 1833). Cokendolpher & Sissom (1988) de- references to malformations among Bothriur- scribed two gynandromorphic diplocentrids (a idae. Cazierius gundlachii (Karsch 1880) and Bio- The main goal of this contribution is to de- culus comondae Stahnke 1968). Armas (1990) scribe and illustrate the tergal anomalies that reported a case of one hermaphrodite Alayo- were found in specimens of three Bothriuridae tityus juraguaensis Armas 1973 and a gyn- species, and to report two more cases of in- andromorphic specimen of Tityopsis inae- tersexuality in scorpions. qualis (Armas 1974) (Buthidae). Maury The specimens studied are preserved in 80 % (1983) described an adult hermaphrodite of ethanol and belong to the following collec- Brachistosternus pentheri Mello-LeitaÅo 1931 tions: AMNH 5 American Museum of Nat- (Bothriuridae) showing intersexual and gyn- ural History (New York, USA); MACN-Ar 5 andromorphic characteristics, and with both Museo Argentino de Ciencias Naturales ``Ber- embryos and hemispermatophores. Another nardino Rivadavia'' (Buenos Aires, Argenti- interesting malformation was reported in two na) and CDA 5 CaÂtedra de Diversidad Ani- males of the bothriurid Bothriurus bonariensis mal I, Universidad Nacional de CoÂrdoba (C.L. Koch 1842), found mating with females (CoÂrdoba, Argentina). Illustrations were pro- in the ®eld, yet presenting only one hemi- duced using a Leica MS5 stereomicroscope spermatophore, the right paraxial organ (that equipped with a camera lucida. Photographs produces the hemispermatophore) being ab- were taken with an Olympus Stylus 400 dig- sent in both specimens (Peretti 2000). The last ital camera under long-wave ultraviolet light. two reports, together with the pedipalps anom- Specimens examined.ÐBothriurus cori- 624 THE JOURNAL OF ARACHNOLOGY aceus Pocock 1893. CHILE: Santiago Region, mation, except for the specimen of B. cori- Chacabuco Province: 1 /, Cuesta de Chaca- aceus from Chacabuco that has a slightly buco, S side, elev. 3900 ft, dry mountainside abnormal telson vesicle, with the left ventral (328599 S, 708449 W), 14 I 1985, N. Platnick, side a little depressed. O.F. Francke, AMNH; Coquimbo Region, The pigmentation pattern of the malformed Choapa Province: 1 /, 4 km N Los Vilos tergites is normal on the Bothriurus coriaceus (318729 S, 718319 W), 5 I 1985, N. Platnick, and B. noa specimens, and apparently in the O. F. Francke, AMNH. Bothriurus noa Maury. Brachistostenus (L.) roigalsinai female as ARGENTINA: TucumaÂn Province: 1 / (par- well. However, in the latter case the specimen atype), TafõÂ del Valle (268529 S, 658519 W), is not well ®xed, and not all the pigmentation 1970 m, 16 I 1981, E. Maury, MACN-Ar has been preserved. 7571. Brachistosternus (Leptosternus) roigal- Tergal malformations include division of sinai Ojanguren-Af®lastro 2002. CHILE: At- tergites and fusion of tergal parts to one an- acama Region, Huasco Province: 1 /, Llanos other. Armas (1976) described a fusion of ter- de Challe National Park, (28809939.80 S, gite I to carapace, and Teruel (2004) observed 71803920.00 W), 205 m, XII 1997, J. Cepeda- division of tergites and fusion to metasomal Pizarro, CDA. Brachistosternus (L.) pentheri. segment I. All the specimens with anomalous ARGENTINA: Mendoza Province: 1 ? (?), tergites examined here were adult females. Te- Reserva de la BioÂsfera NÄ acunÄaÂn (348029 S, ruel (2004) observed the same pattern but 678549 W), 540 m, 20 XI 2003, C. Mattoni, with four females and one male, and the spec- L. Prendini, J. Ochoa, CDA. Bothriurus ara- imen referred by Armas (1974) is an adult guayae Vellard 1934. BRAZIL: Sao Paulo male. I have examined 226 specimens of B. State: 1 ? (?), EstacËao EcoloÂgica de Itirapina, coriaceus (57 females, 62 males and 107 ju- Municipio de Itirapina (228159 S, 478499 W), veniles), and found this kind of anomaly only pitfall, 27 VIII 1999, G. Machado. on the two females referred to here (Mattoni Two of the females (both B. coriaceus) 2003). Despite the few cases reported in scor- were pregnant when preserved. pions, the presence of tergal malformations Tergal malformations.ÐThe B. coriaceus only on adults suggests that they arise during specimen from Cuesta de Chacabuco (Fig. 1) last molt. shows completely longitudinally divided ter- The causes of these tergal malformations gites IV and V. Both parts of each tergite rep- are completely unknown, but they do not resent almost exactly in shape and size the seem to affect the life of the scorpion or its corresponding half of the tergite; only a small mating. The pigmentation pattern in the ter- central portion is lacking from the posterior gites of all the specimens appears not to be edge. The B. noa female shows the same kind altered. of anomaly but only on tergite III. Sexual malformations.ÐBrachistosternus The specimen of B. coriaceus from 4 km N (L.) pentheri: The specimen has intermediate Los Vilos (Fig. 2) also presents some divided sexual characteristics: the small size and num- tergites, with a different arrangement: tergite ber of pectinal teeth (26/30, left/right) suggest IV is completely divided but tergite