Parafontaria tonominea, (Attems, 1899)

Tanabe, T., 2002, Revision of the millipede genus Parafontaria VerhoeV, 1936 (Diplopoda, Xystodesmidae), Journal of Natural History 36 (18), pp. 2139-2183 : 2165-2176

Parafontari a tonominea (Attems, 1899)

(gures 1, 5, 8, 9, 12, 14, 15K–N, 17–19) Fontaria tonominea Attems, 1899: 260 , pl. 13, gure 310, female holotype (Zoologisches

Institute und Zoologisches Museum , Hamburg, examined), type locality: Tounomine ,

Sakurai-shi, Nara Prefecture.

2014 February 13 09: 23 at] Bath of University [by Downloaded

1 mm

Fontaria coarctata circula Attems, 1901: 97 , pl. 1, gures 5–7, one female syntype from Hozu, Kameoka-shi, Kyoto Prefecture (NMW, examined), other syntypes may have been deposited in Zoologisches Institute und Zoologisches Museum , Hamburg, type localities: Nara-shi , Nara Prefecture; Hozu , Kameoka-shi , Kyoto Prefecture. NEW SYNONYMY.

Fontaria coarctata acutidens Attems, 1909: 30 , two male syntypes from Ikaho-machi, Kitagunma-gun, Gunma Prefecture (ZSM), and two male syntypes (preparations of gonopods and their adjacent legs; these preparations made by previous worker) from Mangaeshi, Gotenba-shi, Shizuoka Prefecture (NMW, examined); other syntypes from the following localities perhaps deposited elsewhere: Kusatsu-machi , Azuma-gun , Gunma Prefecture; ‘ Mettan (unidenti ed locality name)’. NEW SYNONYMY.

Fontaria (Japonaria) spiraligera VerhoeV, 1937: 315, gures 3–6, four male syntypes ( ZSM, examined), type locality: Ishiyama, Ôtsu-shi , Shiga Prefecture. NEW SYNONYMY .

Fontaria (Japonaria) marmorata VerhoeV, 1937: 316, gure 7, male holotype (gonopods and their adjacent legs prepared by previous worker, ZSM, examined), type locality: Mt Nachi-san, Nachikatsuura-cho, Higashimuro-gun , Wakayama Prefecture. NEW SYNONYMY .

Apheloria tonominea: Attems, 1938: 171 .

Japonaria (Japonaria) acutidens: Attems, 1938: 176 , gure 193.

Japonaria (Japonaria) circula: Attems, 1938: 177 , gures 195, 196.

Japonaria (Japonaria) spiraligera: Attems, 1938: 178 .

Japonaria (Japonaria) marmorata: Attems, 1938: 179 .

Japonaria spiraligera: Takakuwa, 1942: 41 , gure 5; 1954: 81, gures 88, 89; Miyosi, 1959: 85, pl. 4, gures 60–62.

Japonaria longispinosa longispinosa Miyosi, 1951: 266 , gure 2a, types (YM, not examined) destroyed in re in 1945 by air raid on Matsuyama-shi, type locality: Yawatahama-shi, Ehime Prefecture; Miyosi , 1959: 89, pl. 6, gure 92. NEW SYNONYMY.

Japonaria longispinosa falcata Miyosi, 1951: 266 , gure 2b, types (YM, not examined) destroyed in re in 1945 by air raid on Matsuyama-shi, type locality: Seizui, Hiromi-cho , Kitauwa-gun , Ehime Prefecture; Miyosi , 1959: 90, pl. 6, gure 93. NEW SYNONYMY.

Japonaria circula: Takakuwa, 1954: 76 , gure 80; Miyosi, 1959: 89, pl. 6, gures 90, 91.

Japonaria acutidens: Takakuwa, 1954: 82 , gures 90, 91; Miyosi, 1959: 86, pl. 5, gures 71–73.

Japonaria marmorata: Takakuwa, 1954: 83 , gure 92; Miyosi, 1959: 85, pl. 4, gures 65, 66.

Japonaria acutidens takashimai Shinohara, 1957 , in Takashima and Shinohara, 1957: 406, gure 2, male holotype and male paratype (NSMT, examined), type locality: Mt Kumotori-san, 200 m alt., Chichibu-gun, Saitama Prefecture, Honshu. Another male paratype from Mt Bukô-san , Chichibu-gun, Saitama Prefecture (deposited in private collection of K. Shinohara, not examined). NEW SYNONYMY.

Japonaria egregia Haga, 1968: 6 , pl. 5, gure 11, male holotype and one male paratype (NSMT-My 160, examined), type locality: Mt Sarakura-san, Yahatahigashi-ku, Kitakyushu-shi , Fukuoka Prefecture. NEW SYNONYMY.

Parafontaria tonominea: HoVman, 1978: 219 , gures 4, 5.

Parafontaria acutidens: Murakami, 1993: 99 .

Parafontaria circula: Murakami, 1993: 99 .

Parafontaria egregia: Murakami, 1993: 99 .

Parafontaria longispinos a longispinos a: Murakami, 1993: 99.

Parafontaria longispinosa falcata: Murakami, 1993: 99 .

Parafontaria marmorata: Murakami, 1993: 99 .

Parafontaria spiraligera: Murakami, 1993: 100 .

Diagnosis. Separable from all the other congeners except P. takakuwai in having the tibiotarsal process at mid-length of the tibiotarsus of the gonopods (gures 8,

9). Separable from P. takakuwai by the details of the distal part of the gonopods

(gures 11H, 12).

Male (based on male from Harimichi, Sakurai-shi, Nara Prefecture, 2.5 km

distant from type locality). Gonopodal characters as follows (gures 8A, 12A).

Telopodite forming one and a half loops. Femur with femoral bend and mesal tooth, 2014 without tibiotarsus lateral; tibiotarsal tooth. process Tibiotarsus long bent and dorsally accurate, near located border at mid-length between femur, directed and February in small mesally mesal process, 45 view ° angle.; apical to process main branch spatulate of tibiotarsus; mesal process in ventral slender view; solenomerite, completely forming visible 13 Colour in life (based on same male as above). Tergites pink, with brown stripes 09 along paranotal and posterior margins of metatergites and both anterior and 23: posterior margins of collum. See Variation for geographic variation of colour. at Female holotype. Coxae and sternite of segment 3 modi ed as in gure 14A. Bath] on Cyphopodal posterior surface bellows (tube gure somewhat 15K). sclerotized in proximal region, with at region of Variation. Geographic variation in the gonopods is shown in gures 8, 9, 12. University (The portion gure Males lateral 12 of B in, the E tooth some). apical of populations process the gonopods of have the gonopods a is mesal present bifurcate is in bent many mesally projection populations in some on. the populations The coxae apical of [by the sixth legs (gure 17). Downloaded (in stripes 1) The Geographic the gure. In tergites tergal 14 this. That form coloration variation bluish-greenish of, the intrapopulational cyphopod of also the shows grey coxae is or shown and geographic variation greenish sternite in grey gure sometimes variation of segment or 15. brown. exists There 3, of without: females the are tergal two metatergal is forms colour shown:

varies continuously from brown to bluish-greenish grey or greenish grey within a population. (2) The tergites pink or orange, with brown stripes along the posterior margins of the metatergites and both the anterior and posterior margins of the collum. Some populations seem to show intermediate coloration between these two forms.

Measurements (in mm, mean (range)). Males (N =240): HW 4.37 (3.2–5.2),

MTW 7.62 (5.7–9.6). Females (N =159) HW 4.73 (3.7–5.6), MTW 8.03 (5.8–9.8).

Karyotype. 2 n (male)=12 ( Tanabe, 1992); the number is the smallest in the

Diplopoda; the karyotype was observed for P. circula and Parafontaria sp. , which are treated as P. tonominea in this study.

Distribution (gure 5). Honshu, Shikoku, Kyushu and Sadogashima Is.

Remarks. A distinct species group is recognized in the Kansai district, central

Japan. The species group is composed of P. circula , P. spiraligera , P. marmorata ,

P. acutidens acutidens and P. longispinosa longispinosa . In the species group, there 2014 February 13 09: 23 at] Bath of University [by Downloaded are some sympatric occurrences of two diVerent forms, which diVer mainly in the genital morphology and body size. In each sympatric pair, the two forms are thought to be reproductively isolated from each other. The two forms are designated as forms a and b in each pair (see Specimens examined). This does not mean that forms a and b in one sympatric pair are the same as forms a and b in another sympatric pair. Thus, the labels a and b are used only for distinguishing the two forms in each sympatric pair. Based on morphometrics for 74 populations of the species group, Tanabe et al. (2001) demonstrated that the degree of diVerence in the genital morphology and body size may not correspond with the establishment of reproductive isolation for the species group. The species group seems to be an aggregation of very closely related forms comprising diverse evolutionary states ranging from mere geographic variation to full reproductive units. Thus, the taxonomy of the species group remains unresolved. Although biochemical or molecular data may provide useful information for the recognition of species (reproductive units) within the species group, classi cation based on non-morphological information is not practical for identi cation. One possible solution to this problem is to treat the species group operationally as a single species. It is hence proposed that P. circula , P. spiraligera , P. marmorata , P. acutidens acutidens and P. longispinosa longispinosa are synonymized with P. tonominea . Accordingly, the newly de ned P. tonominea has some sympatric pairs of two reproductively isolated forms in the Kansai district.

Parafontaria tonominea was described based only on the female holotype; the male of this species has never been described. Materials from Harimichi, Sakuraishi, Nara Prefecture, which is near the type locality of P. tonominea , were examined. The females from the locality are recognized as P. tonominea based mainly on the female genital structure. On the other hand, the males collected from the same locality are identi ed as P. marmorata based on the gonopodal structure. These facts show that P. marmorata is a synonym of P. tonominea .

According to the original description, Parafontaria longispinosa falcata diVers from P. longispinosa longispinosa in having an additional inner process on the tibiotarsus of the gonopods. However, this diVerence is recognized as minor intraspeci c geographic variation. It is hence proposed that P. longispinosa falcata is synonymized with P. longispinosa longispinosa (= P. tonominea ).

According to the original description, Parafontaria egregia diVers from P. longispinosa in the following gonopodal characters: the femur has a large projection (gure 9D), the mesal tooth is longer and the distal part is narrower. However, these diVerences are recognized as intraspeci c geographic variation. It is hence proposed that P. egregia is synonymized with P. longispinosa (= P. tonominea ).

According to the original description, P. acutidens takashimai (= Japonaria acutidens takashimai ) diVers from P. acutidens acutidens in having the larger body size, the hook-like tip on the gonopods, and the prefemur with setae in its edge and without setae in its central area in the ventral surface. However, these diVerences are recognized as intraspeci c geographic variation. It is hence proposed that P. acutidens takashimai is synonymized with P. acutidens acutidens (= P. tonominea ).

Specimens examined (two sympatric forms are indicated as form a, form b, see Remarks). Yamagata Prefecture: CJ, along NR 286, 480 m alt., Niiyama, Yamagata-shi, 10 October 1988 ( TT), 10 males , one 7S male, 12 females, three 7S females. Fukushima Prefecture: Southern lakeside of Lake Akimoto-ko, Inawashiromachi , Yama-gun , 24 May 1998 ( SY), one male . Tochigi Prefecture: Mashiko ,

Mashiko-machi, 15 May 1998 ( SY), one male, one female. Gunma Prefecture: Ikaho-machi , Kitagunma-gun, date and collector unknown, two male syntypes of Fontaria coarctata acutidens Attems ( ZSM, see synonymy). CJ, Shimotazawa, Kurohone-mura, Seta-gun , 9 September 1986 ( TT), one male . Locality and collector as above, 30 May 1987, one male . Locality and collector as above, 2 August 1987, three 7S males, one 7S female. Shima, Nakanojo-machi, Agatsuma-gun , 15 July 1975 ( KS), two males . CJ, near Haruna-jinja Shrine, Gunma-gun , 28 May 1987 ( TT), one female . Kirizumi Highlands, Matsuida-cho, Usui-gun , 3 July 1966 ( KS), four males, four females . CJ, Matsuida—Kamigo, Matsuida-machi, Usui-gun , 31 March 1991 ( TT), two males, one female . Saitama Prefecture: types of Japonaria acutidens takashimai (see synonymy), 3 August 1953 ( KS) . CJ, Kamisa, Shiroku, Arakawa-mura , Chichibu-gun , 2 June 1991 ( TT), one male, two females . Chiba Prefecture: Yachimata-shi , 1 June 1981 (E. Ishitani), one female . Locality and

2014 above collector, 21 as June above 1981, 8, two June males 1981., Locality one male and, one collector female as. Locality above, 22 and June collector 1981, one as

February 10 female collector October. Locality as 1968 above ( KS and, 31), collector one May male 1982, as one, two above female males, 29. Gongen-mori, two June females 1981,,. one Nagara-machi Toke-machi male. Locality, Chiba-shi, Chousei- and ,

13 gun, 29–30 May 1991 ( TT), four males, two females. Rokujizo, Nagara-machi ,

09 Chousei-gun, 5 May 1969 (K. Ishii), three males. Kyodai, Mt Kiyosumi-san,

23

:

Amatsukominato-machi, Awa-gun, 20 May 1979, collector unknown, one male. Mt

at Tomi-san, Tomiyama-machi, Awa-gun, 20 August 1969 (K. Ishii), ve males, two

Bath

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one females male. Tokyo. Mt Honnita-san Prefecture: Renkô-ji, Okutama-machi Temple , Tama-shi , Nishitama-gun , 23 August , 3 1994 May (K 1977. Niijima ( KS)) ,,

of three males. Mt Mutsuishi-san , Okutama-machi, Nishitama-gun, 10 June 1979

University

(12 four Nishitanzawa H. September Okamura males, one,) 1986, Yamakita-cho one female ( TT male).,. one CJ Kanagawa ,, male Miyanoshita Ashigarakami-gun. Niigata Prefecture, Prefecture Hakone-machi,: 8 Masugoya-zawa August: Mt , Kinpoku-san Ashigarashimo-gun 1979, Houkizawa (M,. Kanai- Ôrui) ,,,

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by machi, Sado-gun, 1 August 1969 (K. Ishii), one male. Kotaki, 690 m alt., Itoigawa-

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shi one Uenohara-cho Higashiyamanashi-gun, 27 male July. Yamanashi 1983, Kitatsuru-gun (Y. Ôsumi Prefecture, 2 August), one, 16: CJ male 1991 May, Shiobira. Kotaki ( TT 1986 ) , — two, (Itoigawa-shi K Ikedori. males Niijima ,, 1000 ve),, 5 m four females June alt. males, 1998 Makioka-cho. Yuzurihara. (T Misaka-. Sota ),,, sanchi, Minamitsuru-gun , 23 June 1970 ( KS) , one male. Mt Ashiwada-san, Ashiwada-mura, Minamitsuru-gun , 23 July 1969 ( KS) , one male, one female. Near Kômori-ana Cave, Narusawa-mura, Minamitsuru-gun , 22 July 1969 ( KS) , two males, one female. Fuji-midori-no-kyukamura, Narusawa-mura, Minamitsuru-gun , 21 September 1995 ( NT) , one female. Nagano Prefecture: Shiga Highlands, Yamanouchi-machi, Shimotakai-gun , 30 July 1971 ( KS) , one male, two 7S males. Minamiaiki-mura, Minamisaku-gun , 24 July 1977 ( KS) , one male. Shizuoka Prefecture: Mangaeshi, Gotenba-shi, two male syntypes of Fontaria coarctata acutidens (see synonymy), date and collector unknown ( NMH) . CJ, 50 m alt., near Ôkunijinja Shrine, Mori-machi, Shyuchi-gun , 6 August 1987 ( TT) , seven males, seven females, two 7S males. CJ Kunôzantôshôgû Shrine, Shizuoka-shi , 5 August 1987 ( TT) , ve males, 17 females, two 7S males, two 6S males. Locality as above, October 1995 ( SY) , two females. Kotonomamahachimang û Shrine , along NR 1 , Yasaka, Kakegawa-shi , 6 August 1987 ( TT) , ve males, four females, one 7S male, one 6S female. Locality and collector as above, 6 October 1991 collected—cultured —

2014 February 13 09: 23 at] Bath of University [by Downloaded 20 October 1991 xed, one male, three females. CJ, along NR 152 , Funakira— Ôkawa , Tenryu-shi , 25 March 1991 ( TT), one male, one 7S male. Ishikawa Prefecture: CJ, along NR 157, 530 m alt., Dônomori, Shiramine-mura, Ishikawagun, 24 July 1993 ( TT), eight males. CJ, Mt Rokuman-san , Shiramine-mura, Ishikawa-gun, 29 July 1991 (H. Tokumoto), one male. Fukui Prefecture: Toyohara, 160 m alt., Maruoka-cho, Sakai-gun, 1 October 2000 (Y. Nishikawa), one female. CJ, along NR 8 , Okuno, Tsuruga-shi, 22 May 1989 ( TT), three males, three females. Rainbow Line, Mihama-cho, Mikata-gun, 11 May 1990 (K. Ishii), one male. CJ, Kihi-jinja Shrine, 160 m alt., Toune, Tsuruga-shi, 2 November 1990 ( TT), one 7S male, one 7S female. Gifu Prefecture: Shinhodaka, Yoshiki-gun, 15 August 1986 ( TT), one male, two females. Bijo-tôge Pass, Takayama-shi, 16 August 1986 ( TT), one male. Deciduous forest , Utsue, 800 m alt., Kokufu-cho, Yoshiki-gun, 10 June 1991 (Y. Takai), one female. Ôshirakawa, Shirakawa-mura, Ôno-gun, 3 August 1986 (Y. Takai), one male. CJ, Shimoyoshida, Kawabe-cho, Kamo-gun, 14 August 1986 ( TT), two males, three females. CJ, 60 m alt., Shimokawa, Mino-shi, 10 August 1987 ( TT), one male, two females. Near JR Suhara station, Suhara, along NR 156 , Mino-shi, 10 August 1987 ( TT), one male. Kôga-jinja Shrine, Horado-mura, Mugigun, 10 August 1987 ( TT and Y. Takai), two males, four females. CJ, near tomb of Yoshitaka Ôtani, Yamanaka, Sekigahara-cho, Fuwa-gun, 21 August 1991 ( NT), 14 males, eight females. Tara-kyo Ravine, 120 m alt., Kamiishizu-cho, Yourou-gun, 21 August 1991 ( NT), one female. Uchiage, Kamiishizu-cho, Yourou-gun, 21 August 1991 ( NT), one female. Aichi Prefecture: CJ, along NR 301 , near Yamamura, Shinshiro-shi, 13 September 1986 ( TT), one male, two females, one 7S male, one 7S female. CJ, Ôtaga-keikoku Ravine, 650 m alt., Asuke-cho, Higashikamo-gun, 7 October 1994 ( NT), one male, three females. Tokai-shizenhodô, 220 m alt., Utsutsu-tôge Pass, Utsutsu-cho, Kasugai-shi, 17 September 2000 (Y. Nishikawa), two males, one female. Mie Prefecture: CJ, Yamaguchi, foot of Mt Fujiwara-dake , 210 m alt., Fujiwara-cho, Inabe-gun, 21 August 1991 ( NT), form a, one male, one female; form b, two males. CJ, Asake-keikoku Ravine, Komono-cho, Mie-gun, 22 August 1991 ( NT), one male, one female. CJ, Yunoyama, Komono-cho, Miegun, 19 August 1986 ( TT), two males. Locality as above, 14 September 1986 ( TT and T. Morino), three males, two females. CJ, along NR 25 , Kabutokitazaike, Sekicho, Suzuka-gun, 1 September 1986 ( TT), one male. Locality as above, 14 May 1988 ( TT), two females. CJ, Sono, Miyagawa-mura, Taki-gun, 19 August 1986 ( TT), seven males, three females. Locality and collector as above, 15 May 1988, two males, two females. Shiga Prefecture: Iiura, Kinomoto-cho, Ika-gun, 9 July 2000 (H. Nishi), one male. Okishima Is., Oumihachiman-shi, 25 June 1997 (K. Yahiro), one male, one female. CJ, near Mizusaka Tunnel, along NR 303, 900 m west of junction with NR 367 , Imazu-cho, Takashima-gun, 18 May 1993 ( TT), two males, seven females. CJ, Tochyuudani, Imazu-cho, Takashima-gun, 14 June 1989 ( TT), three females. Utodani, Kutsuki-mura, Takashima-gun, 1 October 1997 (K. Yahiro), one female. Nakamaki, Kutsuki-mura, Takashima-gun, 10 June 2000 (H. Nishi), one male. CJ, along NR 367 , Nakamura , near border between Nakamura and Kidouchi, Ôtsu-shi, 17 May 1993 ( TT), two females. Ishiyama, Ôtsu-shi, date and collector unknown, two male syntypes of Fontaria (Japonaria) spiraligera (see synonymy). CJ, 90 m alt., Sotohata, Ôtsu-shi, 18 May 1993 ( TT), two males, one female. CJ, along NR 422 , 3.6 km south-east of Shikatobi-bashi Bridge, Ôishitomikawa, Ôtsu-shi, 18 May 1993 ( TT), ve males, four females. CJ, along NR 422 , Miyajiri, 3.2 km north of junction with NR 307 , Shigaraki-cho, Kôga-gun,

18 May 1993 ( TT), six males, six females. CJ, along NR 307 , 19 km east of junction with NR 24, 350 m alt., Urajiro-tôge Pass, Shigaraki-cho, Kôga-gun, 17 September

1986 ( TT), two males, one female. Locality and collector as above, 12 August 1987,

ve females. Kyoto Prefecture: CJ, 370 m alt., Ashiu— Sugo, Miyama-cho ,

Kitakuwada-gun , 31 July 1991 ( NT), one male, one female . Ashiu experimental plantation of Kyoto University, Ashiu , Miyama-cho , Kitakuwada-gun , 14 June 2000

(H. Nishi), two males. CJ, 300 m alt., Ewa—Tauta, Miyama-cho, Kitakuwada-gun ,

31 July 1991 ( NT), one male, one female. Fukami Tunnel, Miyama-cho ,

Kitakuwada-gun , 31 July 1991 ( NT), ve males, one female . CJ, 340 m alt., north of Mt Chitose-san , 270 m alt., Yagi-cho, Funai-gun, 31 July 1991 ( NT), two males ,

two females. CJ, 180 m alt., Atagotanigawa woodland path, Hozu, Kameoka-shi ,

31 July 1991 ( NT), seven males, four females. Mt Hozu-san, Kameoka-shi, date and collector unknown, one female syntype of Fontaria coarctata circula (see syn- 2014 onymy Kamigamo). Kumogahata, Kyoto-shi,, Kyoto-shi 26 May, 2000 25 June (T. 2000 Sota) (, H one. Nishi male),. ve Experimental males, two females Station,. February cho Konzô-ji Kamigamo, Kyoto-shi Temple, Kyoto-shi, 7, October Kyoto-shi, 26 1999 May, 29 (T July 2000. Sota 1991 (N),. pitfall Osawa ( NT) trap,) two,, one one males male female.. Shugakuinsenmanda CJ. Locality, 440 m as alt above., near -, 13 7 November 1999 (T. Sota), pitfall trap, one female. Iwakura, Kyoto-shi, 09 20 September 2000 (Y. Kuwahara), two males. CJ, 220 m alt., Hosotani, Wachi- 23: cho, Funai-gun, 1 August 1991 ( NT), one male, one female. CJ, Kashihara, Miyamaat cho, Kitakuwada-gun, 10 May 1992 ( TT), one male. CJ, about 300 m alt., Nagase,] near Ôno Dam, Wachi-cho, 9 May 1992 ( TT), one male, six females. CJ, along Bath NR 9, 1.35 km west of junction with NR 173, Ijiri, Mizuho-cho, Funai-gun, 13 May of 1988 ( TT), form a, ve males, six females; form b, one male, one female. CJ, along University 13 NR and 1992 August 173 collector ( TT, Itasaka-tôge) 1986, one as (above TT male),, form Pass, two 13, May females a near, two 1988 Itasaka females. CJ, form, Ruri-kei Tunnel; form a, one, b Mizuho-cho, Ravine four male males;, form Sonobe-cho,, one Funai-gun b, two female, males Funai-gun., Locality 10, three May, [by females. CJ, along NR 27, near Itsubo-ôhashi Bridge, Tonase, Ayabe-shi, 9 May Downloaded 9 gun 2000 1992 May, (1 ( TT Y 1992 August.), Kawakami two ( TT 1991 males), one. () NT, CJ male one,) 360,. one Cryptomeria female m alt female.., Enoki-tôge CJ., Mt japonica Sai-jinja Ôe-san Pass and Shrine,, Ôe-cho Shizushi Chamaecyparis , Mikata-cho,, Kasa-gun Mizuho-cho obtusa ,, Ayabe-shi 2, August Funaimixed,

forest, 150 m alt., Mt Karasuga-dake, Fukuchiyama-shi, 1 August 1991 ( NT),

three males. CJ, 200 m alt., Shingu, Fukuchiyama-shi, 3 August 1992 ( NT) , one male, two females. CJ, 180 m alt., Nakayama, Yasaka-cho, Takeno-gun, 4 August

1992 ( NT), form a, one male; form b, one male. Osaka Prefecture: CJ, Mt

Myojinga-dake, Takatsuki-shi, 29 July 1991 ( NT), form a, one male, one female ;

form b, two males, one female. Mt Mikusa-san , Nose-cho , Toyono-gun, 25 July

2000 (Y. Kawakami), one male. Kôdaiji Temple, Toyono-cho, Toyono-gun, mesh code: 5235–23–95, 26 October 2000 (T. Wada), ve males, three females. CJ, 350 m

alt., Katsuô-ji Temple, Minoo, Minoo-shi, 30 July 1991 ( NT), one female. Zenihara ,

440 m alt., Ibaraki-shi, 23–26 October 1976 (Y. Nishikawa), one male, sugar trap.

Fukui, Ibaraki-shi , 5 October 2000 (T. Wada), two males, two females. CJ, Izuhara ,

380 m alt., Ibaraki-shi, 21 May 2000 (Y. Nishikawa), one male, under stone. Mt

Inunari-san, Izumisano-shi, 30 July 2000 (T. Wada and K. Yamanaka), two females.

Western slope of Mt Ikoma-san, Higashiosaka-shi, 22 September 2000 (T. Wada),

form a, one female. Locality as above, 24 September 2000 (T. Wada), form a, one 2014 February 13 09: 23 at] Bath of University [by Downloaded female. Locality as above, 24 September 2000 (Y. Kawakami), form a, two females, one 7S female; form b, one male. Locality as above, 24 September 2000 (S. Kitaguchi), form a, three males, two females; form b, one 7S male, one 7S female. Sobura, mesh code: 5135 –43–34, Kaizuka-shi, 27 September 2000 (T. Wada), one male. Mt Izumikatsuragi-san, Sobura, Kaizuka-shi, 29 October 2000 (T. Wada), one male. Hyogo Prefecture: CJ, along NR 178 , Kawanashi-tôge Pass, Toyookashi, 3 June 1990 ( NT), one female. CJ, along NR 173 , Kodanaka, Sasayama-shi, 10 May 1992 ( TT), one male, one female. CJ, along NR 173 , Fukuzumi, Sasayamashi, 13 May 1988 ( TT), two males. CJ Hioki, Sasayama-shi, 10 May 1992 ( TT), one female. CJ, Furusaka-tôge Pass, near Furusaka Tunnel, along prefectural road, Sasayama-shi, 10 May 1992 ( TT), three males, two females. CJ, Furuichi, along prefectural road, Sasayama-shi, 10 May 1992 ( TT), two males, ve females. CJ, 360 m alt., Kuroishi Dam, Sasayama-shi, 2 August 1991 ( NT), one male. CJ, 360 m alt., Amadani-tôge Pass, Tantou-cho, Izushi-gun, 3 August 1992 ( NT), one female. Itoi-keikoku Ravine, 400 m alt., Wadayama-cho, Asago-gun, 20 August 1990 ( NT), one female. CJ, along NR 312 , Motozu, Asago-cho, Asago-gun, 20 May 1991 ( TT), seven males. CJ, 760 m alt., Nakusa-jinja Shrine, Mt Myoken-san, Youka-cho, Yabugun, 20 June 1989 ( NT), two males, two females. CJ, along NR 9 , Yutani, Onsencho, Mikata-gun, 19 May 1991 collected ( TT)— cultured— 27 May 1991 xed, three males, two females; 20 June 1991 xed, seven males, eight females. CJ, near skiing ground, 550 m alt., Mihara-kôgen Highlands, Onsen-cho, Mikata-gun, 2 August 1989 ( NT), four females. CJ, Hachifuse-kôgen Highlands, near skiing ground, Sekinomiya-cho, Yabu-gun, 20 July 1989 ( NT), one male. Takitani, Shingu-cho, Ibo-gun, 23 August 1995 ( SY), one female. CJ, Senbon, near JR Senbon Station, Shingu-cho, Ibo-gun, 28 April 1991 collected ( TT)—cultured— 8 June 1991 xed, form a, eight males, three females; form b, three males, two females. Nara Prefecture: CJ, Harimichi, Sakurai-shi, 24 May 1989 ( TT), three males, four females. Type of Fontaria tonominea (see synonymy), September 1895 (Suck). Wakayama Prefecture: type of Fontaria (Japonaria) marmorata (see synonymy), date and collector unknown. CJ, along NR 424 , Shimotsuno, Kainan-shi, 30 April 1993 ( TT), two males, four females. CJ, Hiyamizu, Kainan-shi, 16 April 1991 ( TT), three males, two females. Locality and collector as above, 30 April 1993, six males. CJ, along NR 42 , Ikeda, Hidaka-cho, Hidaka-gun, 6 April 1991 ( TT), form a, three males, seven females; form b, one male, one female. Hisagi, Hikigawa-cho, Nishimuro-gun, 18 July 1982 (H. Minato), ve males. Kokashi-tôge Pass, 300 m alt., Ezumi, Susami-cho, Nishimuro-gun, 10 August 1995 (Y. Nishikawa), one male. Kamiôno, Hongû-cho, Higashimuro-gun, 5 August 1996 ( SY), two females. Kawayu-onsen Spa, Hongûcho, Higashimuro-gun, 16 August 1997 ( SY), one female. Tottori Prefecture: CJ, Kaji—Ôdori-tôge Pass, 770 m alt., along woodland path, Wakasa-cho, Yazu-gun, 25 July 1989 ( NT), eight males, ve females. Oshika-kei Ravine, Misasa-cho, Tôhaku-gun, 22 August 1992 ( SY), one male. Namariyama, Misasa-cho, Tôhakugun, 22 August 1992 (H. Minato), one male. CJ, 570 m alt., north-eastern slope of Mt Hotokegasen, Misasa-cho, Tôhaku-gun, 21 September 1988 ( NT), four females. Sasa sp. grassland, Kagamiganari, 940 m alt., Mt Daisen, Daisen-cho, Saihaku-gun, 27 August 1987 ( NT), ve males, six females. Along Daisen-natsuyama-tozand ô (mountain trail), 1570 m alt., Mt Daisen, Daisen-cho, Saihaku-gun, 18 September 1992 ( NT), one male. Daisen-ji Temple, under rotten wood, 780 m alt., Daisen-cho, Saihaku-gun, 3 May 1992 ( NT), two females. Chamaecyparis obtusa forest, Mt Mihira-san, 660 m alt., Kôfu-cho, Hino-gun, 3 August 1989 ( NT), four males, ve females. Cryptomeria japonica and Chamaecyparis abtusa mixed forest, 550 m alt., Hijihara—Nakamura, Nichinan-cho, Hino-gun, 3 August 1989 ( NT), two males, one female. Shimane Prefecture: Shishidani, Sanbe experimental plantation of Shimane University, Mt Sanbe-san, Ôda-shi, 15 September 1996 collected (N. Kaneko)— cultured— May 1997 xed, nine males, two females. Nishinohara, Mt Sanbe-san, Ôda-shi, 28 May 1988 (N. Kaneko), four males, three females. Ôtani, Sanbe experimental plantation of Shimane University, Tonbara-cho, Iishi-gun, 27 October 1993 (N. Kaneko), three males, two females. Western foot of Mt Sanbesan, Ôda, Ôda-shi, 28 October 1993 (N. Kaneko), one male, ve females. Kawamotomachi, Ôchi-gun, October 1987, collector unknown, two males. CJ, 610 m alt., near Dôga-tôge Tunnel, Hikimi-cho, Mino-gun, 19 July 1990 ( NT), one male, one female. Okayama Prefecture: CJ, Kumai-tôge Pass, Chûka-son, Maniwa-gun, 26 October 1993 ( NT), three males, four females. Inubasari-tôge Pass, 530 m alt., Shimohiruzen,

2014 one Yatsuka-son female. CJ, Maniwa-gun, 440 m alt.,, Mitano 28 October, Yatsuka-son 1990 (M,. Maniwa-gun Murakami and, 16 NT July), 1991 two males ( NT) ,,

February one gun woodland, male 24 September. path Cryptomeria to Makibanoyakata 1993 japonica (NT) , and two, south-east Chamaecyparis males, three of Mt females obtusa Kajiko-san. CJ mixed, 430, Kume-cho forest m alt,. 460, Unekita, Kumem alt.,,

13 Nakaseijitsu, Yoshii-cho, Akaiwa-gun, 5 May 1993 ( NT), one male, ve females .

09 CJ, 260 m alt., Hatakeganari, Kamogawa-cho, Mitsu-gun, 5 May 1993 ( NT) , two

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females. Hiroshima Prefecture: Sasa sp. grassland, 1080 m alt., Tsukimigaoka skiing at ground, Mt Dogo-san, Hiba-gun , 14 July 1989 ( NT), four males, six females. CJ ,

Bath

]

(Kajiyadoko NT), two females, west of. Cryptomeria Taishaku-kyo japonica , 560 m and alt., Chamaecyparis Tôjô-cho, Hiba-gun obtusa , 7 mixed August forest 1990,

of 800 m alt., Tawarabaragoshi, Takano-cho, Hiba-gun, 6 August 1990 ( NT), one

University

male three dawa Takano-cho, females Pass three,, 630 females. Hiba-gun Chamaecyparis m alt.., CJ, Takano-cho 6, August 640 obtusa m 1990 alt,., Hiba-gun forest Tamaidawa-jinja ( NT,), 770 one, 6 m female August alt., Kusan-dawa. Shrine CJ 1990, 630, (Tamai-dawa NT m alt) Pass,., three Tamanuki-, Takanomales Pass,,

[

by cho, Hiba-gun, 6 August 1990 ( NT), one male, one female. Cryptomeria japonica

Downloaded alt 440

and Kuchiwa-cho., m Fukayama Chamaecyparis alt., Chiyoda-cho, Hiba-gun, Mt obtusa Tenguishi-san,, Yamagata-gun 6 August mixed, 1990 forest Geihoku-cho, 5 (, NT August 710), m one, 1990 alt Yamagata-gun male., (Kaneohara NT. CJ),, two Nijudani-tôge, males 4 skiing August. CJ ground, 760 Pass 1990 m ,, ( NT), four males, two females . CJ, start of mountain trail, Mt Kanmuri-san, 740 m alt., Saioto, Geihoku-cho , Yamagata-gun , 4 August 1990 ( NT), one female . Fagus crenata forest, 990 m alt., Mt Garyu-san, Geihoku-cho, Yamagata-gun , 19 July 1990 ( NT), eight males, six females . CJ, 980 m alt., Uchiguro-tôge Pass, Togauchi-cho, Yamagata-gun , 19 July 1990 ( NT), three males, two females . CJ, 810 m alt., Kanmuri-kôgen Highlands, Yoshiwa-mura, Saeki-gun , 18 July 1990 ( NT), six males, four females . Yamaguchi Prefecture: CJ, 880 m alt., Mt Rakan-san, Nishiki-cho, Kuga-gun , 18 July 1990 ( NT), one male . CJ, Hibari-tôge Pass, 260 m alt., Asahison, Abu-gun , 5 May 1995 ( NT), two males . Tokushima Prefecture: Ôtaki, Wakimachi, Mima-gun , 13 October 2000 (K. Kawano), one female . CJ, Saga — Kurimizaka, Sanagouchi-son, Myôdô-gun , 9 June 1991 ( TT), one male, one female . CJ, Ichinose, Sanagouchi-son, Myôdô-gun , 18 September 1992 ( TT), one male, one female . CJ, Okunono, 320 m alt., Sanagouchi-son, Myôdô-gun , 30 April 1992 collected ( TT)— cultured— 1 May 1992 xed, one male, three females . CJ, Mt Shôzanjisan, 850 m alt., Kamiyama-cho, Myozai-gun , 29 July 1999 (M. Ogawa), one male .

2014 February 13 09: 23 at] Bath of University [by Downloaded CJ, Mt Kôtsu-san, 210 m alt., Yamakawa-cho, Oe-gun, 9 May 1991 ( TT) collected— cultured— 9–20 May 1991 xed, four males, seven females. CJ, Mt Kôtsu-san, 210–320 m alt., Yamakawa-cho, Oe-gun, 9 May 1991 ( TT), six males. CJ, Mt Kôtsusan, 320 m alt., Yamakawa-cho, Oe-gun, 9 May 1991 collected ( TT)—cultured— 10 June 1991 xed, one male, one female. Cryptomeria japonica and Chamaecyparis obtusa mixed forest, Mt Ryuô-san, 990 m alt., Mima-cho, Mima-gun, 3 October 1990 ( NT), two females. CJ, Fujiu, Mima-cho, Mima-gun, 5 September 1992 ( TT), two males, seven females, six 7S males, four 7S females. Mt Tairyuji-san, Anan-shi, 5 September 1993 (H. Minato), one male. Jyurokucho, Kitajisando, Mt Tairyujisan, Anan-shi, 6 October 2000 (K. Kawano), one female. CJ, near Jigen-ji Temple, 510 m alt., Kamikatsu-cho, Katsuura-gun, 6 July 1991 ( TT), one male, one female. CJ, near Unpen-ji Temple, 840 m alt., Mt Unpenji-san, Ikeda-cho, Myoshi-gun, 11 June 1988 ( NT), three females. Locality as above, 25 May 1989 ( TT), one female. Locality as above, 5 August 1999 ( SY), one male. Fujiyu, about 400 m alt., Shitsukawa, Ikeda-cho, Miyoshi-gun, 23 August 1993 ( TT), one male. CJ, near Tsurugisan skiing ground, along prefectural road 22, 1350 m alt., Ichiu-son, Mimagun, 10 June 1993 ( TT), two males. Abies homolepis forest, Mt Miune, along mountain trail, 1490–1590 m alt., Higashiiyayama-son, Miyoshi-gun, 1 July 1992 (M. Kamada), 41 males, 14 females. CJ, Toyanotakidani, Tonono, near Koboke, Yamashiro-cho, Miyoshi-gun, 2 October 1996 ( TT), two females, one 7S male. CJ, Mt Jinkichimori, 1000–1100 m alt., along mountain trail, Kitô-son, Naka-gun, 29 August 1999 ( SY), six males, two females. CJ, 330 m alt., along prefectural road 296, 1.2 km of junction with NR 193 , near Futonoji-bashi Bridge, Kaikawatanikawa, Kaikawa, Kaminaka-cho, Naka-gun, 1 October 2000 ( TT), 11 males, three females. Kagawa Prefecture: Nagaodani, Sangawa-cho, Ôkawa-gun, 28 July 1994 ( SY), two males, three females. CJ, camping ground, 700 m alt., Mt Ryuô-zan, Shionoe-cho, Kagawa-gun, 3 October 1990 ( NT), one female. Sugiô-jinja Shrine, Kawaoku, Kotonami-cho, Nakatado-gun, 25 May 1997 ( SY), one male, two females. Ôsako, Kotonami-cho, Nakatado-gun, 25 September 1993 ( SY), one male. Motoshino, Saita-cho, Mitoyo-gun, 23 May 1996 ( SY), one male. Kotohira-jinja Shrine, Kotohira-cho, Nakatado-gun, 23 May 1994 ( SY), one male. Ehime Prefecture: Nanata, Kawanoe-shi, 8 May 1993 ( SY), one male, one 7S female. Shimohachimanjinja Shrine, Shimotani, Kawanoe-shi, 8 May 1993 ( SY), three males, one female. Shimotani, Kawanoe-shi, 3 May 1993 ( SY), one female. Near Sanpeki-kyo Bridge, Sanpeki-kyo Ravine, Saijo-shi, 17 September 1992 ( SY), one female. Near Ômiya-jinja Temple, Nishinokawa, Saijo-shi, 17 September 1992 ( SY), one male. Along Higashinokawadani River, other side of Kôchihachiman-jinja Shrine, Higashinokawa, Saijo-shi, 17 September 1992 ( SY), one female. CJ, Fukumi-dera Temple, Mt Fukumi-san, Shigenobu-cho, Onsen-gun, 15 August 1991 ( NT), one male, six females. CJ, 400–500 m alt., experimental plantation of Ehime University, Ôino, Matsuyama-shi, 27 September 1999 (Y. Nishikawa), two males, one female. Cryptomeria japonica and Chamaecyparis obtusa mixed forest, Ehime Forest Park, Lake Ôtani, Iyo-shi, 3 May 1994 ( NT), one male. CJ, Sakura-suidô, Seizui, Hiromicho, Kitauwa-gun, 26 May 1989 ( TT), one male. Yamagamiou-jinja Shrine, Hiromicho, Kitauwa-gun, 18 April 1987 ( TT), two females. Kashima Is., Nishiumi-cho, Minamiuwa-gun, 19 April 1987 ( TT), three males, four females. Kochi Prefecture: Mt Senbon-san, along mountain trail, Umaji-mura, Aki-gun, 31 October 1998 ( TT), one female. Mishima-jinja Shrine, Yusuhara-cho, Takaoka-gun, 20 August 1994 ( SY), one male. Ichinomataonsen-keikok u Ravine, Taisho-machi, Hata-gun,

25 September 1999 ( SY) , one female. Mt Imano-san , along mountain trail, Miharamura, Hata-gun , 26 September 1999 ( SY) , one female. Asanabetodorokinotak i Water Fall , Ichinose Saga-cho, Hata-gun , 13 June 1999 ( SY) , one female. Akitatenmangû Shrine , Nakamura-shi , 25 September 1999 ( SY) , one female. Mt Kokuzô-san, Sakawa-cho , Takaoka-gun , 10 June 1995 ( SY) , one female, one 7S female. Along Imose-gawa River, Hahajima , Okinoshima Is., Sukumo-shi , 20 August 1996 ( SY) , one male, one female. Fukuoka Prefecture: male holotype of Japonaria egregia (NSMT-My 160; see synonymy) , 21 April 1960 (A. Haga). Kawachi, Yahatahigashi-ku , Kitakyushu-shi , 19 October 1965 (A. Haga), one male paratype of J. egregia (NSMT-My 160). Kumamoto Prefecture: CJ, along NR 445, 580 m alt., Hayakusu, Tomochi-machi, Shimomashiki-gun , 30 May 1989 ( TT) , one male, one female. CJ, Momigi—Yamaingiri , along woodland path, 1060 m alt., Izumimura, Yatsushiro-gun , 13 August 1993 collected ( TT)— cultured— 3 September 1993

2014 forest xed,, nine 930 males m alt.,, 13 Yuyama-tôge females. Cryptomeria Pass , Mizukami-mura japonica and, Chamaecypari Kuma-gun, 22 s obtusa August mixed 1989

February (1999 machi NT), (, T one Minamiamabe-gun. Sota female), pitfall. Oita trap Prefecture, 23, one August male: Mt 1989. CJ Tsurumi-dake , ( NT Mt), Shakuma-san one, male Beppu-shi , one, 350 female, 16 m – alt 17. CJ., October , Yayoi- 240 m

13 alt., Ôhara, Ôhira, Ume-machi, Minamiamabe-gun, 23 August 1989 ( NT), three

09 males, two females. Miyazaki Prefecture: CJ, 1000 m alt., Iiboshi-tôge Pass, Mt

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Iiboshi-san, Shiiba-son, Higashiusuki-gun, 22 August 1989 ( NT), one female .

at Literature records. There are many literature records for P. tonominea , e.g.

] Nishikawa and Murakami (1991), Kuwabara (1993) and Minato (1968).

Bath of Species of uncertain status

University Japonaria terminalis Takakuwa Parafontaria, 1942 : terminalis 39, gures 1 (, Takakuwa 2, type (s) (, YM 1942, not) examined) destroyed

[in re in 1945 by air raid on Matsuyama-shi, type locality (see Remarks): Shiroyama,

by Kagoshima-shi, Kagoshima Prefecture; Takakuwa, 1954: 80, gure 85; Miyosi, 1959: 86,

Downloaded Parafontaria pl Remarks Distribution. 5, gures terminalis . No. 69, Kyushu 70 specimens:. Murakami: Kagoshima of, 1993 this: species 100 Prefecture. have. been collected since the original description. Many attempts by the author to collect this species at the type locality, Kagoshima-shi and its vicinity have failed. The gures of the original description clearly show that this species is very similar to P. erythrosoma , which is distributed in Ibaraki and Tochigi Prefectures, about 1000 km from the type locality of P. terminalis . These facts suggest that P. terminalis is conspeci c with P. erythrosoma and that the type locality of P. terminalis was mislabelled. It is hence conjectured that the correct type locality of P. terminalis is in the distributional range of P. erythrosoma or its adjacent region.

Results of cladistic analysis

The cladistic analysis yielded 33 equally parsimonious cladograms with a tree length of 17 steps, a consistency index of 0.941, a homoplasy index of 0.059, a retention index of 0.964 and a rescaled consistency index of 0.908. The strict and majority rule (50%) consensus trees identi ed two and eight clades, respectively,

2014 February 13 09: 23 at] Bath of University [by Downloaded

Strict