Variation of the Poison Duct in Chilopoda Centipedes from Taiwan
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Norw. J. Entomol. 53, 139-151. 3 Nov. 2006 Variation of the poison duct in Chilopoda centipedes from Taiwan Jui-Lung Chao & Hsueh-Wen Chang Chao, J. L. & Chang, H. W. 2006. Variation of the poison duct in Chilopoda centipedes from Taiwan. Norw. J. Entomol. 53, 139-151. We present a new technique to observe the structure the poison ducts of the forcipules of chilopod centipedes under light microscope. We examined twenty one species in fourteen genera of chilopods from Taiwan, Thereuopoda, Lithobius, Bothropolys, Esastigmatobius, Scolopendra, Rhysida, Otostigmus, Scolopocryptops, Cryptops, Scolioplanes, Stigmatogaster, Mecistocephalus, Prolamnonyx, and Taiwanella. Morphology and structure of poison ducts of chilopoda are consistent in the higher taxa genera and families. Morphological characters of poison duct support a close relationship between Scolopocryptops and Scolopendridae, suggesting that the current separation of Cryptopidae from Scolopocryptopidae is correct. We also revised Takakuwa’s description of the poison duct of Scolopocryptops rubiginosus, and consider Taiwanella yanagiharai is a valid species in the genus Taiwanella. Key words: poison calyx, glycerin, transparent specimens, taxonomy. Jui-Lung Chao and Hsueh-Wen Chang (corresponding author), Department of Biological Sciences, National Sun Yat-Sen University, 70 Lien-Hai Rd., 804 Kaohsiung, Taiwan, R.O.C. E-mail: [email protected] INTRODUCTION Foddai et al. 2003). Takakuwa (1940b) drew and described a short poison duct in the tarsungulum The forcipules (maxillipedes) are one of the of the forcipules of Otocryptops rubiginosus (L. common characters of Chilopod centipedes. Koch, 1878) (now Scolopocryptops rubiginosus Each forcipule of Chilopoda comprises four (L. Koch, 1878)) (Figure 1c). When studying articles: trochanteroprefemur, femur, tibia, and this species with new material, we found his tarsungulum (Lewis et al. 2005, Figure 1). Within description and fi gure might have some mistakes. each forcipule is a poison duct opening to the tip With a new and simple method developed in our of the tarsungulum and ending in a poriferous lab, we considered it was necessary to re-examine region, poison calyx, which connects to the poison the poison ducts of Chilopoda. With the variation glands by many short tubular canals (Figure 2). of the structure and length of the poison ducts in Because it is an internal structure and small in centipedes, we considered that the characters of the size, it is diffi cult to observe the poison ducts poison ducts may suggest some information about under dissecting microscope and light microscope. the relationships among members of Chilopoda. Therefore, the structure of the poison duct had rarely been studied and described in Chilopoda centipedes, and only few morphological fi gures MATERIALS AND METHODS of the forcipules with their poison ducts have been drawn (Takakuwa 1940a, b, 1941, Verhoeff We examined the poison ducts of twenty one 1940, Lewis 1996, 1999, 2002, Bonato et al. 2002, species of Chilopoda (Table 1), including one 139 Chao & Chang: Variation of poison duct in Chilopoda from Taiwan Figure 1. The forcipules of Chilopoda are composed of four parts: tarsungulum (Ta), tibia (Ti), femur (Fe), and trochanteroprefemur (Tr). A. Mecistocephalus takakuwai. B. Bothropolys asperatus. C. Scolopocryptops rubiginosus. D. Thereuonema hilgendorfi . (after Takakuwa1940). species of Scutigeromorpha, Thereuopoda Rhysida immarginata (Porat, 1876), Otostigmus clunifera (Wood, 1863); four species of scaber Porat, 1876, Otostigmus aculeatus Lithobiomorpha, Esastigmatobius longicornis Haase, 1887, Scolopocryptops rubiginosus (L. Takakuwa, 1936, Lithobius ongi Takakuwa, Koch, 1878), Scolopocryptops capillipedatus 1941, Bothropolys asperatus Koch, 1878, , and (Takakuwa, 1938), Cryptops japonicus Takakuwa, Bothropolys richthofeni Verhoeff, 1938; ten species 1934, and Cryptops nigropictus Takakuwa, 1936; of Scolopendromorpha, Scolopendra subspinipes six species of Geophilomorpha, Taiwanella Leach, 1815, Scolopendra morsitans Linnaeus, yanagiharai Takakuwa, 1936, Mecistocephalus 1758, Rhysida longipes longipes (Newport, 1845), smithi Pocock, 1895, Mecistocephalus mikado 140 Norw. J. Entomol. 53, 139-151. 3 Nov. 2006 A B Figure 2. Micrographs of the poison calyx of Scolopendra subspinipes. A. The poison gland is around the poison calyx. B. The cylindrical poison calyx has numerous short tubular canals connec- ting the poison glandular cells (GC). B A Figure 3. Micrographs of the poison duct of Thereuopoda clunifera. A. The cylindrical poison calyx is located at the base of the tarsungulum, but the poison gland extends across tibia and femur, reaching the anterior part of trochanteroprefemur (arrow). B. The poison gland cells (GC) are around the poison calyx. Scale is 0.4 mm. A B Figure 4. Micrographs of the poison duct in Esastigmatobius longicornis. A. The poison duct is loca- ted within tarsungulum (Ta). Scale is 0.3 mm. B. The poison calyx is cylindrical with about 100 pores. 141 Chao & Chang: Variation of poison duct in Chilopoda from Taiwan Table 1. The situations of poison calyxes of Chilopoda. Ta: tarsungulum; Ti: tibia; Fe: femur; Tr: trochanteroprefemur; Co: coxosternum. +: Situated; -: Absent. Situation of poison calyx Species Ta Ti Fe Tr Co Scutigeromorpha Scutigeridae Thereuopoda clunifera (Wood, 1863) + - - - - Lithobiomorpha Henicopidae Esastigmatobius longicornis Takakuwa, 1936 + - - - - Lithobiidae Lithobius ongi Takakuwa, 1941 + + - - - Bothropolys asperatus Koch, 1878 + + - - - Bothropolys richthofeni Verhoeff, 1938 + + - - - Scolopendromorpha Scolopendridae Scolopendra subspinipes Leach, 1815 + + + + - Scolopendra morsitans Linnaeus, 1758 + + + + - Rhysida longipes longipes (Newport, 1845) + + + + - Rhysida immarginata (Porat, 1876) + + + + - Otostigmus scaber Porat, 1876 + + + + - Otostigmus aculeatus Haase, 1887 + + + + - Scolopocryptopidae Scolopocryptops rubiginosus (L. Koch, 1878) + + + + - Scolopocryptops capillipedatus (Takakuwa, 1938) + + + + - Cryptopidae Cryptops japonicus Takakuwa, 1934 - - - + - Cryptops nigropictus Takakuwa, 1936 - - - + - Geophilomorpha Mecistocephalidae: Mecistocephalinae Taiwanella yanagiharai Takakuwa, 1936 + + + - - Mecistocephalus smithi Pocock, 1895 + + + - - Mecistocephalus mikado Attems, 1928 + + + - - Mecistocephalidae: Aruppinae Prolamnonyx holstii (Pocock, 1919) + + + + + Himantariidae Stigmatogaster japonica Takakuwa, 1935 - + - - - Geophilidae Scolioplanes maritimus japonicus Verhoeff, 1935 - - + + - 142 Norw. J. Entomol. 53, 139-151. 3 Nov. 2006 A B Figure 5. Micrographs of the poison duct in Bothropolys richthofeni. A. The poison duct reaches the tibia (Ti). Scale is 0.8 mm. B. The poison calyx is bell-shaped at bottom with numerous small pores. A B Figure 6. Micrographs of the poison duct in Bothropolys asperatus. A. The poison duct reaches the tibia (Ti). Scale is 0.5 mm. B. The cylindrical poison calyx is thick at base and with many small pores. Attems, 1928, Prolamnonyx holstii (Pocock, the specimens transparent, so that the poison ducts 1919), Stigmatogaster japonica Takakuwa, 1935, can be observed without dissection. and Scolioplanes maritimus japonicus Verhoeff, 1935. Centipedes were anaesthetized and sandwiched between two glass slides, then fi xed RESULTS and immersed in glycerin (99.5%) over one week. The specimens were then observed under light We found the shape and length of poison ducts microscope. Long immersion in glycerin makes varied among species. The poison duct may 143 Chao & Chang: Variation of poison duct in Chilopoda from Taiwan Figure 7. Micrograph of the poison duct in Lit- hobius ongi. The bended poison calyx reaches the anterior part of tibia (Ti) with about 200 small pores. Scale is 0.2 mm. Figure 8. Micrograph of the poison duct in Cryp- tops japonicus. The anterior non-porous duct reaches trochanteroprefemur, and the poison calyx situates the anterior part of trochanteropre- femur. Scale is 0.1 mm. reach tarsungulum, intermediate articles (tibia distribute across tibia and femur, reaching to the and femur), trochanteroprefemur or coxosternum anterior part of trochanteroprefemur. The poison (Table 2). The shapes of poison calyxes may be calyx of poison duct is a short elongated cylinder short cylindrical, long elongated cylindrical, sub- with numerous small pores, and the number of cordate, round or bell-bottomed. pores increases towards the posterior part. In Scutigeromorpha, the poison duct of In Lithobiomorpha, the poison duct of Thereuopoda clunifera is located entirely within Esastigmatobius longicornis is also located the tarsungulum (Figure 3). The poison gland cells within the tarsungulum (Figure 4), and the poison 144 Norw. J. Entomol. 53, 139-151. 3 Nov. 2006 C A B Figure 9. Micrographs of the poison ducts in Scolopendridae. A. Scolopendra subspinipes (scale is 1.2 mm). B. Otostigmus aculeatus (scale is 0.5 mm). C. Rhysida immarginata (scale is 0.5 mm). Their cylindrical poison calyxes (arrow), reaching the base of trochanteroprefemur, are arched with numerous pores. A C B Figure 10. The poison ducts of Scolopocryptops are cylindrical and longest among all specimens examined, being serpentine in the trochanteroprefemur (arrow). A. Scolopocryptops rubiginosus (scale is 1.0 mm). B. Scolopocryptops capillipedatus (scale is 0.5 mm). C. The end of poison calyx reaches the edge between trochanteroprefemur and coxosternum (circle). A trachea passes the end of poison calyx ventrally. It is easily to treat the trachea as the extension of poison