Molecular analysis reveals a new cryptic species in a limpet Lottia kogamogai (Patellogastropoda: Lottiidae) from Japan Author Nakayama, Ryo Author Sasaki, Takenori Author Nakano, Tomoyuki text Zootaxa 2017 4277 2 237 251 journal article 32828 10.11646/zootaxa.4277.2.4 9ab7ddd1-eb2d-416d-af4c-6fb576cd0a22 1175-5326 809682 572E426E-59FD-452A-A5FF-54C39C1E0FBF Lottia goshimai n. sp. Collisella heroldi : Niu et al . 1992 , figs 2A–C. Lottia kogamogai : Sasaki & Okutani, 1994b , pl. 2a–c. Lottia kogamogai : Fukui & Kashio, 2012 , p. 22, pls 3–2. Lottia cf. kogamogai : Kristof et al . 2016 , p. 2, fig. 1a. Type materials. Holotype ( UMUT-RM 32483 ), Usujiri , Hakodate , Hokkaido , Japan , 10.8 x 8.2 x 3.8 mm . Selected paratypes : Paratype 1 (NSMT-Mo78956) ; Paratype 2 (NSMT-Mo78957); Paratype 3 (UMUT-RM32492); Paratype 4 (UMUT-RM32471), Paratype 5 (UMUT-RM32496); Paratype 6 (UMUT-RM32494); Paratype 7 (UMUT- RM32479). Remaining paratypes are also deposited at UMUT (UMUT RM3232467-32505). Etymology. This species is dedicated to Prof. Seiji Goshima, Faculty of Fisheries Sciences, Hokkaido University, Japan , who has greatly contributed to biological studies of the new species ( Niu & Fuji 1989 ; Niu et al . 1992 , 1994a , b , 1997 , 1998 ). Shell ( Figures 2 , 3 ). Shell moderately thick, aperture oval. Apex antero-dorsally directed and positioned at anterior fourth. Anterior and ventral slopes nearly straight, posterior slope convex. Exterior surface sculptured by crowded, rough, regularly arranged radial riblets. Exterior colour light brown with 12 brown radial rays. Interior margin alternating light brown and dark brown; intermediate area white; central area except apex stained dark chocolate brown; apex area white. Animal. Dorsal part of mantle margin fringed with light brown. Head and visceral mass dark beige. Ventral surface creamy white. FIGURE 2. Shell morphology of Lottia species. A – H. Lottia goshimai n. sp. A. Holotype, on rocks, UMUT-RM32483, Usujiri, Hakodate, Hokkaido (Loc. F); 10.8 x 8.4 x 3.7 mm. B. Paratype 1, on rocks, NSMT-Mo78956, Karakuwa, Kesennuma, Miyagi (Loc. R); 12.7 x 10.6 x 6.2 mm. C. Paratype 2, on rocks, NSMT-Mo78957, Usujiri, Hakodate, Hokkaido (Loc. F); 15.3 x 12.0 x 6.7 mm. D. Paratype 3, on rocks, UMUT-RM32492, Shirakami, Matsumae, Hokkaido (Loc. D); 14.4 x 11.6 x.7.4 mm. E. Paratype 4, on rocks, UMUT-RM32471, Takashima, Otaru, Hokkaido (Loc. B); 8.1 x 5.9 x 3.5 mm. F. Paratype 5, on rocks, UMUT-RM32496, Asamushi, Aomori, Aomori (Loc. I); 8.7 x 6.8 x 3.1 mm. G. Paratype 6, on Septifer virgatus , UMUT- RM32494, Shirakami, Matsumae, Hokkaido (Loc. D); 8.9 x 7.0 x 5.6 mm. H. Paratype 7, on Septifer virgatus , UMUT- RM32479, Esan, Hakodate, Hokkaido (Loc. G); 9.6 x 7.8 x 5.0 mm. I. Lottia luchuana , on rocks, UMUT-RM32514, Onna, Okinawa (Loc. r); 10.5 x 8.3 x 4.7 mm. J. Lottia kogamogai , on rocks, UMUT-RM32417, Banda, Tateyama, Chiba (Loc. Y); 11.8 x 9.5 x 4.0 mm. K. Lottia lindbergi , on rocks, UMUT-RM32509, Kaminokuni, Hiyama, Hokkaido (Loc. C); 14.3 x 11.0 x 5.6 mm. L. Lottia tenuisculpta , on rocks, UMUT-RM32506, Takashima, Otaru, Hokkaido (Loc. B); 11.6 x 9.2 x 5.1 mm. Scale bar = 1 cm. FIGURE 3. Shell sculpture of Lottia goshimai n. sp. and Lottia kogamogai A. Lottia goshimai n. sp. , UMUT-RM32480, Esan, Hakodate, Hokkaido, Japan (Loc. G). B. Lottia kogamogai , UMUT-RM32427, Jogashima, Miura, Kanagawa Pref., Japan (Loc Z). Scale bar = 1mm. FIGURE 4. Radula morphology of Lottia goshimai n. sp. and Lottia kogamogai with morphological variation related to their habitats. A – D. Lottia goshimai n. sp. A, B. Lottia goshimai n. sp. on rocks. UMUT-RM32493, Shirakami, Matsumae, Hokkaido, Japan (Loc. D). C, D. Lottia goshimai n. sp. on the shell of Septifer virgatus . UMUT-RM32484, Usujiri, Hakodate, Hokkaido, Japan (Loc. F). E – H. Lottia kogamogai . E, F. Lottia kogamogai on rocks. UMUT-RM32455, Isozakihana, Shingu, Fukuoka Pref., Japan (Loc. k). G, H. Lottia kogamogai on the shell of Hormomya mutabilis . Cape Ooarai, Ooarai, Ibaraki Pref., Japan (Loc. U). Scale bar = 50 µm. Radula ( Figure 4 ). First lateral teeth relatively short with pointed cusp. Second lateral teeth long with slightly rounded cusp. Third lateral teeth broad with beak-like cusps. Marginal teeth small, rounded. Type locality. Usujiri , Hakodate , Hokkaido , Japan ( 41°56'11"N , 140°57'00"E ). Distribution. Northern part of Japan from Erimo, Hokkaido to Karakuwa, Miyagi on the Pacific coast and Monbetsu, Hokkaido to Tsuruoka, Yamagata , Vostok Bay, Russia in the Japan Sea. Habitat. This species lives in the middle to lower intertidal zone, and is often seen among the small barnacle Chthamalus challengeri and on the purplish mussel Septifer virgatus . Remarks. Although the new species is morphologically similar to L. kogamogai , it is distinguished by having rough, fine and regularly-arranged radial riblets ( Fig. 3 ). The radulae differ in terms of the morphology of the lateral teeth. Lottia goshimai n. sp. has short first lateral teeth, a long and slightly rounded cusp on the second lateral teeth, and slightly beak-like third lateral teeth ( Figs. 4 A–D). Lottia kogamogai has long and rounded first lateral teeth, a broad and pointed cusp on the second lateral teeth, and short, broad third lateral teeth ( Figs. 4 E–H). Intraspecific variation was observed within L. goshimai n. sp. and L. kogamogai in shell and radular morphogy. The individuals living on the shell of Septifer virgatus have taller, darker shells and shorter second lateral teeth than those from rocks ( Figs. 2 , 4 ). The reproductive cycle of the species (as Collisella heroldi ) was examined by Niu & Fuji (1989) . According to this study, the reproductive season is August to October, and size at maturity is about 5 mm . The Lottia kogamogai complex is clearly divided according to geographical distribution of each species: Lottia goshimai n. sp. , L. kogamogai and L. luchuana are limited to northern Japan , southern Japan and tropical Japan respectively ( Fig. 5 ). Although L. tenuisculpta and L. lindbergi have a wide distribution along temperate Japan ( Sasaki & Okutani 1994b ), these two species were not analyzed intensively in this paper. FIGURE 5. The geographical distribution of three Lottia species with warm sea currents around Japan. The record of Lottia goshimai n.sp. from Russian coast is from Kristof et al. (2016). FIGURE 6. Phylogenetic tree of Bayesian analysis based on COI data (661 bp). The numbers above each branch are NJ bootstrap/MP bootstrap/Bayesian posterior probabilities. Three species of Nipponacmea were used as outgroup. TABLE 1. Specimen anđ localitie analyzeđ in thi tuđy. * mean the pecimen ob erveđ in rađular morphology anđ hell culpture. UMUT: The Univer ity Mu eum, The Univer ity Tokyo. NSMT: National Mu eum, Nature anđ Science, Tokyo. Specie locality ID in Fig. 1 GenBank Voucher Lottia kogamogai Sa aki & Okutani, 1994 Banđa, Tateyama, Chiba Pref., Japan ( type locality)* K320 Y LC137940 UMUT-RM 32416 Banđa, Tateyama, Chiba Pref., Japan ( type locality)* K321 Y LC137941 UMUT-RM 32417 Banđa, Tateyama, Chiba Pref., Japan ( type locality)* K322 Y LC137942 UMUT-RM 32418 Cape Unoko , Kitaibaraki , Ibaraki Pref., Japan K583 S LC138001 UMUT-RM 32419 Cape Ooarai , Ooarai , Ibaraki Pref., Japan K571 U LC137997 UMUT-RM 32420 Cape Ooarai , Ooarai , Ibaraki Pref., Japan * K572 U LC137998 UMUT-RM 32421 Cape Inubou , Cho hi, Chiba Pref., Japan K578 V LC138000 UMUT-RM 32422 Cape Yahata , Kat uura, Chiba Pref., Japan K489 W LC137991 UMUT-RM 32423 Cape Yahata , Kat uura, Chiba Pref., Japan K490 W LC137992 UMUT-RM 32424 Cape Yahata , Kat uura, Chiba Pref., Japan K566 W LC137996 UMUT-RM 32425 Joga hima, Miura , Kanagawa Pref., Japan * K336 Z LC137945 UMUT-RM 32426 Joga hima, Miura , Kanagawa Pref., Japan * K337 Z LC137946 UMUT-RM 32427 Joga hima, Miura , Kanagawa Pref., Japan K338 Z LC137947 UMUT-RM 32428 Joga hima, Miura , Kanagawa Pref., Japan K351 Z LC137948 UMUT-RM 32429 Joga hima, Miura , Kanagawa Pref., Japan * K503 Z LC137994 UMUT-RM 32430 Mitsui hi, Manazuru , Kanagawa Pref., Japan K316 a LC137938 UMUT-RM 32431 Yokata Port , Toyama , Toyama Pref., Japan K33 L LC137916 UMUT-RM 32432 Yokata Port , Toyama , Toyama Pref., Japan K249 L LC137935 UMUT-RM 32433 Yokata Port , Toyama , Toyama Pref., Japan K251 L LC137936 UMUT-RM 32434 Akagurizaki , Ooi , Fukui Pref., Japan K244 N LC137933 UMUT-RM 32435 Akagurizaki , Ooi , Fukui Pref., Japan K246 N LC137934 UMUT-RM 32436 Morozaki , Minamichita , Aichi Pref., Japan K29 b LC137915 UMUT-RM 32437 Morozaki , Minamichita , Aichi Pref., Japan K208 b LC137930 UMUT-RM 32438 Rinkai , Shirahama , Wakayama Pref., Japan K84 e LC137919 UMUT-RM 32439 Ku himoto, Wakayama Pref., Japan K3 đ LC137909 UMUT-RM 32440 Ku himoto, Wakayama Pref., Japan K4 đ LC137910 UMUT-RM 32441 ……continued on the next page Specie locality ID in Fig. 1 GenBank Voucher TABLE 1. (Continueđ) Ku himoto, Wakayama Pref., Japan K145 đ LC137923 UMUT-RM 32442 Ku himoto, Wakayama Pref., Japan K146 đ LC137924 UMUT-RM 32443 Ku himoto, Wakayama Pref., Japan K152 đ LC137925 UMUT-RM 32444 Ku himoto, Wakayama Pref., Japan * K157 đ LC137926 UMUT-RM 32445 Urađome , Iwami , Tottori Pref., Japan * K423 O LC137979 UMUT-RM 32446 Cape Hane , Kochi Pref., Japan K 66 g LC137917 UMUT-RM 32447 Go hikinohama, U a, Kochi Pref., Japan K258 h LC137937 UMUT-RM 32448 Omuronohama , Okit u, Kochi Pref., Japan K 28 i LC137914 UMUT-RM 32449 Tat uku hi, Kochi Pref., Japan * K21 j LC137912 UMUT-RM 32450 Tat uku hi, Kochi Pref., Japan K174 j LC137927 UMUT-RM 32451 T uou, Nagato , Yamaguchi Pref., Japan * K364 P LC137949 UMUT-RM 32452 T uou, Nagato , Yamaguchi Pref., Japan K366 P LC137950 UMUT-RM 32453 T uou, Nagato , Yamaguchi Pref., Japan K367 P LC137951 UMUT-RM 32454 I ozakihana, Shingu , Fukuoka Pref., Japan * K70 k LC137918 UMUT-RM 32455 Ura hiro, Nobeoka , Miyazaki Pref., Japan K334 o LC137943 UMUT-RM 32456 Ura hiro, Nobeoka , Miyazaki Pref., Japan K335 o LC137944 UMUT-RM 32457 Cape Sata , Minamio umi, Kago hima Pref., Japan K 26 m LC137913 UMUT-RM 32458 Cape Sata , Minamio umi, Kago hima Pref., Japan K 230 m LC137931 UMUT-RM 32459 Cape Sata , Minamio umi, Kago hima Pref., Japan K 231 m LC137932 UMUT-RM 32460 Hamakanaya, Futt u, Chiba Pref. Japan L945 X LC138005 UMUT-RM 32461 Kaino, Kihoku, Mie Pref., Japan L158 c LC138003 UMUT-RM 32462 Kaino, Kihoku, Mie Pref., Japan L159 c LC138004 UMUT-RM 32463 Kamogaura, Wajima, I hikawa Pref., Japan L1147 M LC138007 UMUT-RM 32464 Kamogaura, Wajima, I hikawa Pref., Japan L1148 M LC138008 UMUT-RM 32465 Kamogaura, Wajima, I hikawa Pref., Japan L1149 M LC138009 UMUT-RM 32466 Lottia goshimai n. sp. Oumu , Monbet u, Hokkaiđo , Japan K434 A LC137983 UMUT-RM 32467 Oumu, Monbet u, Hokkaiđo , Japan K439 A LC137984 UMUT-RM 32468 Cape Erimo, Erimo, Hokkaiđo , Japan K522 E LC137995 UMUT-RM 32469 ……continued on the next page Specie locality ID in Fig. 1 GenBank Voucher ……continued on the next page TABLE 1. (Continueđ)

Taka hima, Otaru, Hokkaiđo, Japan*	K368	B	LC137952	UMUT-RM32470
Taka hima, Otaru, Hokkaiđo, Japan	K373	B	LC137953	UMUT-RM32471
Taka hima, Otaru, Hokkaiđo, Japan	K375	B	LC137954	UMUT-RM32472
Kaminokuni, Hiyama, Hokkaiđo, Japan	K384	C	LC137961	UMUT-RM32473
Kaminokuni, Hiyama, Hokkaiđo, Japan	K385	C	LC137962	UMUT-RM32474
Kaminokuni, Hiyama, Hokkaiđo, Japan*	K386	C	LC137963	UMUT-RM32475
Kaminokuni, Hiyama, Hokkaiđo, Japan	K387	C	LC137964	UMUT-RM32476
E an, Hakođate, Hokkaiđo, Japan*	K394	G	LC137966	UMUT-RM32477
E an, Hakođate, Hokkaiđo, Japan	K395	G	LC137967	UMUT-RM32478
E an, Hakođate, Hokkaiđo, Japan	K397	G	LC137968	UMUT-RM32479
E an, Hakođate, Hokkaiđo, Japan	K398	G	LC137969	UMUT-RM32480
E an, Hakođate, Hokkaiđo, Japan*	K401	G	LC137970	UMUT-RM32481
E an, Hakođate, Hokkaiđo, Japan*	K404	G	LC137972	UMUT-RM32482
U ujiri, Hakođate, Hokkaiđo, Japan	K407	F	LC137973	NSMT-Mo78957
U ujiri, Hakođate, Hokkaiđo, Japan*	K409	F	LC137974	UMUT-RM32483
U ujiri, Hakođate, Hokkaiđo, Japan*	K413	F	LC137975	UMUT-RM32484
U ujiri, Hakođate, Hokkaiđo, Japan*	K414	F	LC137976	UMUT-RM32485
U ujiri, Hakođate, Hokkaiđo, Japan*	K415	F	LC137977	UMUT-RM32486
U ujiri, Hakođate, Hokkaiđo, Japan	K98	F	LC137920	UMUT-RM32487
Tateno, Hokuto, Hokkaiđo, Japan*	K103	H	LC137921	UMUT-RM32488
Tateno, Hokuto, Hokkaiđo, Japan	K106	H	LC137922	UMUT-RM32489
Shirakami, Mat umae, Hokkaiđo, Japan	K378	D	LC137956	UMUT-RM32490
Shirakami, Mat umae, Hokkaiđo, Japan	K379	D	LC137957	UMUT-RM32491
Shirakami, Mat umae, Hokkaiđo, Japan	K380	D	LC137958	UMUT-RM32492
Shirakami, Mat umae, Hokkaiđo, Japan	K381	D	LC137959	UMUT-RM32493
Shirakami, Mat umae, Hokkaiđo, Japan	K382	D	LC137960	UMUT-RM32494
A amu hi, Aomori Pref., Japan	K480	I	LC137985	UMUT-RM32495
A amu hi, Aomori Pref., Japan*	K481	I	LC137986	UMUT-RM32496

TABLE 1. (Continueđ) Specie locality ID in Fig. 1 GenBank Voucher A amu hi, Aomori Pref., Japan * K482 I LC137987 UMUT-RM 32497 A amu hi, Aomori Pref., Japan * K483 I LC137988 UMUT-RM 32498 A amu hi, Aomori Pref., Japan * K484 I LC137989 UMUT-RM 32499 Ot uchi, Iwate Pref., Japan * K205 Q LC137928 UMUT-RM 32500 Ot uchi, Iwate Pref., Japan K206 Q LC137929 UMUT-RM 32501 Ot uchi, Iwate Pref., Japan K496 Q LC137993 UMUT-RM 32502 Karakuwa, Ke ennuma, Miyagi Pref., Japan * K318 R LC137939 NSMT-Mo78956 Karakuwa, Ke ennuma, Miyagi Pref., Japan K429 R LC137980 UMUT-RM 32503 Karakuwa, Ke ennuma, Miyagi Pref., Japan K431 R LC137981 UMUT-RM 32504 Imaizumi, T uruoka, Yamagata Pref., Japan K574 J LC137999 UMUT-RM 32505 Lottia tenuisculupta Sa aki & Okutani , 1994 Taka hima, Otaru , Hokkaiđo , Japan * K369 B LC137953 UMUT-RM 32506 Urađome, Iwami, Tottori Pref., Japan K417 O LC137978 UMUT-RM 32507 Kamekawahamađa, Beppu, Ohita Pref., Japan K433 n LC137982 UMUT-RM 32508 Lottia lindbergi Sa aki & Okutani , 1994 Kaminokuni , Hiyama , Hokkaiđo , Japan * K388 C LC137965 UMUT-RM 32509 Hirai o, Hitachinaka, Ibaraki Pref., Japan L173 T LC137971 UMUT-RM 32510 Tat uku i, Kochi Pref., Japan K20 j LC137911 UMUT-RM 32511 Lottia luchuana ( Pil bry, 1901) Ka ari, Amami , Kago hima Pref., Japan L965 p LC138006 UMUT-RM 32512 Ogimi, Okinawa , Okinawa , Japan L131 q LC138002 UMUT-RM 32513 Onna, Okinawa , Japan * K486 r LC137990 UMUT-RM 32514