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Geographic Variation of the Brachypterous Grasshopper Parapodisma Setouchiensis Group in Western Honshu, with Its Taxonomic Revision

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Geographic Variation of the Brachypterous Grasshopper Parapodisma Setouchiensis Group in Western Honshu, with Its Taxonomic Revision Species Diversity. 1999, 4. 43 61 Geographic Variation of the Brachypterous Grasshopper Parapodisma setouchiensis Group in Western Honshu, with Its Taxonomic Revision Yasushi Kawakami' Department ofBiology, Faculty ofEducation, Tottori University, Tottori, 680-0945 Japan (Received 10 March 1997; Accepted 6 July 1998) The Parapodisma setouchiensis group (Catantopidae) is revised based on geographic variation in external morphology. This group appears to constitute a single polytypic species, P. setouchiensis Inoue, consisting of four geographic forms (Basic, Mt. Hyonosen, Tanba, and Yamato). Four taxa, Parapodisma tanba, P. yamato, P. hyonosenensis hyonosenensis, and P. hyonosenensis kibi, are treated as junior synonyms of P. setouchiensis. The polarity of character states in the P. setouchiensis group was inferred from the distribution pattern of the geographic forms. All pairs of adjacent forms intergrade through relatively narrow zones that may have originated from secondary contact. In two genital characters (triangular marking and cercus morphology), there are clines with different ranges across the boundary between the Tanba and Mt. Hyonosen forms. Key Words: geographic variation, brachypterous grasshopper, Parapodisma setouchiensis group, morphology, taxonomic revision, transitional zone, hybrid zone. Introduction The Parapodisma setouchiensis group is a series of brachypterous grasshoppers which is indigenous to Japan and adjacent islands. The group is mainly distributed in the Japanese Archipelago from Aomori Prefecture to Yakushima Island, and also in Cheju Island of the Republic of Korea. Of the species or species-groups belonging to the genus Parapodisma, this group has the widest distribution. Tominaga et al. (1996) mentioned that the P. setouchiensis group comprises five taxa all having mutually allopatric or parapatric distributional ranges: Parapodisma setouchiensis Inoue, 1979; P. tanbaensis Tominaga and Kano, 1989; P. yamato Tominaga and Storozhenko, 1996; P. hyonosenensis hyonosenensis Tominaga and Kano, 1996; and P. hyonosenensis kibi Tominaga and Kano, 1996. However, classification of the group remained unclear, since its geographical variation in external characters is consider able and had not yet been examined in detail. The purpose of the present paper is first to determine the pattern of geographic variation of the P. setouchiensis group mainly around the eastern part of the Chugoku mountains, western Honshu, where the group exhibits its highest morphological diversity. The second purpose is to revise this polymorphic and taxonomically intractable group. In this paper, I will treat this group as a single species comprising four geographic forms, and, from the patterns of geographic variation, infer the origins of the transitional zones for various characters found in the eastern Chugoku "Present address: Tottori West Technical High School, Kita 3-250, Koyama-cho, Tottori-shi, Tottori, 680-0941 Japan 44 Yasushi Kawakami mountains. Materials and Methods A total of 409 specimens was collected from 46 localities (Fig. 3) in western Honshu from July to September in 1994 and 1995. The specimens were preserved in 80% ethanol and used for the measurement of external characters. All the specimens examined and localities where they were collected are listed in the Appendix. Each sample is designated by the code number given in the Appendix. Males: Males were collected from all 46 localities. The average sample size for males per locality was 6.9. Eight morphological characters shown in Figure 1 were measured for each male specimen. Body length, head width, pronotum length, fore wing length, and hind femur length were measured by using a vernier caliper. The lengths of the subgenital plate and triangular marking (a dark mark on the dorsal subgenital plate) were measured with a micrometer installed on a stereo-microscope. The left cercus in lateral view was accurately sketched with a drawing apparatus, and the angle between its distal protuberance and basal part (Fig. 1) was then measured. The eight characters were compared in detail among populations. Since the fore wing length is supposedly correlated with the size of the thorax, the character state of fore wing length was expressed as the ratio of the fore wing length to the pronotum length. In comparing cerci, the shape and the angle between the distal process and the base of the cercus were used (Fig. 1). The data for 1994 and 1995 were pooled, since no significant between-year difference was detected in any of the eight characters in samples from locality 13 where the largest number of males was obtained each year (13 and 16 males, respectively; Kolmogorov-Smirnov test, P>0.20). Females: Females were obtained from 34 localities. Five characters (body length, head width, pronotum length, fore wing length, and hind femur length) were measured for each female using the same procedures for males (Fig. 1). Morphological Variation 1. Body size Of the eight characters measured, body length (BL), head width (HW), pronotum length (PL), and hind femur length (HFL), all of which are correlated with body size, showed no particular geographical trend along the same latitudes in either males or females. The body size of univoltine insects often varies with altitude or latitude (Masaki 1996). Hence the relation of the four characters to the altitudes of the localities was analyzed in the univoltine P. setouchiensis group. The influence of latitude was ignored because all the localities were at almost the same latitude, from 34° 50' to 35° 30' north. All the characters except for BL of females showed a negative correlation with altitude [BL: male r2= 0.48, P<0.01; female r2= 0.10, P>0.05. HW: male (Fig. 2) r2= 0.69, P<0.01; female r2= 0.50, P<0.01. PL: male r2= 0.54, P<0.01; female r2= 0.21, P<0.01. HFL: male 1^ = 0.54, P<0.01; female r2= 0.23, P<0.01]. The reason for the absence of correlation between female BL and altitude may be the Geographic variation of a grasshopper 45 ca tml;;[ Fig. 1. Eight characters of the P. setouchiensis group measured for the analyses of geographic variation. Abbreviations: BL = body length, CA = cercus angle, HFL = hind femur length, HW = head width, PL = pronotum length, SPL = subgenital plate length, TML = triangular marking length, WL = fore wing length. 46 Yasushi Kawakami large variation in female BL that results from the different degree of ovarian development among individuals. Of all the characters, HW showed the strongest correlation with altitude in both males and females (Fig. 2). 2. Triangular marking The triangular marking (TM) of males varied geographically (Figs 4-6). It was distinct in western and eastern populations but was obscure or absent in the middle part of the area surveyed. Figure 5 displays the geographic pattern of variation in TML. Populations 1-8 and 43-46 consisted of individuals with a prominent TM (Fig. 6A, C), about 0.4 mm long, while populations 22-39 consisted almost exclusively of individuals without a TM (Fig. 6B). In the two zones comprising populations 9-21 and 40-42, TML was highly variable (Figs 4A, 7). In each of these two zones a steep cline was found in the development of TM (Figs 4, 5). One transitional zone, comprised of populations 9-21, extends from Mt. Jubo (Shikano-cho, Tottori) south to the northern part of Okayama city. The other zone, including populations 40-42, occupies a small area around a watershed between the Yura and Kako Rivers. 3. Cercus Figures 4B and 8 show the geographic variation in the shape and the angle (lateral view) of the cerci in males. The tip of the cercus became slenderer and the angle became smaller eastward, but the pattern of geographic variation was some what different between the two characters. The tip was broad in western populations 1-9 whereas it was relatively slender 4.8 y=-7.3xl0"4x+4.7 r2=0.69(r=-0.83) 4.7 P < 0.01 ^o *°«° o 4.6 ^ o o Oo> o 4.5 o ° O ° o O o 5 4.4 8^ z 4.3 oo OlSsQo o 4.2 4.1 o 4 100 200 300 400 500 600 700 800 Altitude (m) Fig. 2. Relationship of male head width to the altitude of localities in the P. setouchiensis group. Each sample is represented by the population mean (open circle). All 46 samples listed in the Appendix are included. o used for the morphological analyses. Each sample is shown by the code number given Fig. 3. Provenance of samples of the P. setouchiensis group in the Appendix. 48 Yasushi Kawakami A. Triangular marking length uii ''iiJHjj..*...*...** il i • • • • i 160 t 150 j T B. Cercus angle 60 lil:"!"""!!..!.,!.,,..!.o» 120 . - 110 I ± *I < 100 w. ° 90 rt 80 Jlii.! 70 I ••' • I 1.2 C. Subgenital plate length ? 1.1 6 1 _j .9 a. 0) .8 .7 Xk0 .6 i > ••• i i • • • • i 1.6 D. Fore wing length / Pronotum length S 1.5 6 1.4 6 1.3 6 >-' 1.2 _j Q. 1.1 "^ -1 1 £ .9 i'!.jrt^p"" " 1 3 5 7 9 11 13 15 17 19 21 23 25 27 29 31 33 35 37 39 41 43 45 2 4 6 8 10 12 14 16 18 20 22 24 26 28 30 32 34 36 38 40 42 44 46 Locality numbers Fig. 4. Geographic variation of four characters in males of the P. setouchiensis group. Code numbers for localities correspond to those in Fig. 3 and localities are approximately arranged from west (left) to east (right). Eachsample is represented by the mean (circle) and the range of 1 SD (bar). a OKAYAMA Fig. 5. Relative frequencies of three grades of length of male triangular marking in the P. setouchiensis group. The three grades are given in the left box. Circle size shows the number of examined individuals, as illustrated in the right box.
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