Macrobrachium lanchesteri (De Man, 1911)

Chaowvieng, Apisara, Sutcharit, Chirasak, Chanabun, Ratmanee, Srisonchai, Ruttapon, Jeratthitikul, Ekgachai & Siriwut, Warut, 2024, Molecular phylogeny and taxonomic position of Macrobrachium lanchesteri (De Man, 1911), with descriptions of two new species from Thailand (Decapoda, Caridea, Palaemonidae), ZooKeys 1190, pp. 163-193 : 163

Macrobrachium lanchesteri (De Man, 1911)

Figs 3A, B , 4

Palaemon paucidens Lanchester, 1902: 568, pl. 33, fig. 4. Type locality: Singora [Songkhla Province, Thailand]. [Not De Haan (1844) and Hilgendorf (1893)].

Palaemon (Eupalaemon) lanchesteri De Man, 1911: 264. Replacement Name.

Palaemon lanchesteri : Kemp 1918: 257.

Macrobrachium lanchesteri : Suvatti 1937: 49; Suvatti 1967; 139; Holthuis 1950: 139; Holthuis 1980: 96; Johnson 1961: 56; Naiyanetr 1980: 17; Naiyanetr 1992: 17; Naiyanetr 1998: 32; Chong and Khoo 1988: 196; Cai and Ng. 2002: 77; Cai et al. 2004: 586; Wowor et al. 2004: 349; Hanamura et al. 2011: 13.

Macrobrachium lar : Kamita 1966: 138.

Cryphiops (Macrobrachium) lanchesteri : Johnson 1968: 233.

Macrobrachium cf. lanchesteri : Ng 1994: 75, fig. 2.

Material examined.

Topotypes: Songkhla • 11 ♀♀, 3 ♂♂, Wat Pho Klang, Khu Khut, Sathing Phra; 7°25'26.2"N, 100°25'05.8"E; MUMNH MP00216. • 25 ♀♀, 3 ♂♂, Laem Wang, Khu Khut, Sathing Phra; 7°27'12.3"N, 100°25'12.9"E; MUMNH MP00217. • 5 ♀♀, 3 ♂♂, Bang Khiat, Singhanakhon; 7°20'19.2"N, 100°25'36.8"E; MUMNH MP00218.

Additional material.

Chiang Rai • 12 ♀♀, 16 ♂♂ , Chiang Khong Street Market, Wiang, Chiang Khong; 20°15'57.7"N, 100°24'21.7"E; MUMNH MP00219. Chiang Mai • 1 ♀ , Nam Lao, Pa Pae, Mae Taeng; 19°12'16.4"N, 98°40'51.3"E; MUMNH MP00350. Phayao • 5 ♀♀ , 1 ovigerous, 4 ♂♂, Wean, Ban Pa Suk, Chiang Kham; 19°30'04.7"N, 100°16'32.9"E; MUMNH MP00220. • 17 ♀♀, 14 ♂♂, Yom, Ban Duea, Pong; 19°06'24.3"N, 100°15'58.5"E; MUMNH MP00221. Nan • 4 ♀♀ , Mae Mang, Bo Kluea Tai, Bo Kluea; 19°08'12.6"N, 101°09'03.1"E; MUMNH MP00222. • 10 ♀♀, 1 ovigerous, 8 ♂♂, Ban Hua Fai, Sisaket, Na Noi; 18°19'33.4"N, 100°43'25.6"E; MUMNH MP00223. • 1 ovigerous, Huai Pa Sak, Sisaket, Na Noi ; 18°19'33.4"N, 100°43'25.6"E; MUMNH MP00224. • 1 ♀, 1 ♂ , Ban Huai Lao, Chiang Khong, Na Noi; 18°18'42.3"N, 100°54'18.2"E; MUMNH MP00225. • 2 ♀♀, 1 ovigerous, 4 ♂♂, Huai Hin, Sathan, Na Noi; 18°14'21.8"N, 100°41'57.7"E; MUMNH MP00226. • 2 ♀♀, Ban Na Bua , Nong Daeng, Mae Charim; 18°45'12.8"N, 101°00'39.7"E; MUMNH MP00227. • 3 ♀♀, 1 ♂ , Nan River , Tan Chum, Wiang Sa; 19°00'59.5"N, 100°46'47.3"E; MUMNH MP00228 . Lampang • 1 ♀, 2 ♂♂ , Ban Mae Pa, Thoen; 17°38'17.9"N, 99°15'39.6"E; MUMNH MP00229. Mae Hong Son • 3 ♀♀, 8 ♂♂ , Pai, Pang Mu, Mueang Mae Hong Son; 19°19'36.9"N, 97°56'34.8"E; MUMNH MP00230. • 2 ♀♀, 2 ♂♂ , Yuam, Mae Sariang; 18°09'26.0"N, 97°55'37.8"E; MUMNH MP00231. • 8 ♀♀, 4 ♂♂, Nong Pong Sila, Khun Yuam; 18°50'26.4"N, 97°56'21.3"E; MUMNH MP00232. Uttaradit • 3 ♀♀, 5 ♂♂ , Huai Nam Muet, Phak Khuang, Thong Sang Khan; 17°28'21.5"N, 100°22'25.6"E; MUMNH MP00233. • 4 ♀♀ , 1 ovigerous, 3 ♂♂, Klong Tron, Ban Bueng Pra Kot, Thong Sang Khan; 17°35'39.1"N, 100°29'10.3"E; MUMNH MP00234. • 4 ♀♀, 15 ♂♂ , Nam Pat, Fak Tha; 18°00'04.4"N, 100°52'42.7"E; MUMNH MP00235. • 2 ♀♀, Ban Rai Phana Wan, Muang Chet Ton, Ban Khok; 18°08'48.7"N, 101°02'11.0"E; MUMNH MP00236 . Tak • 2 ♀♀, 2 ♂♂ , Ban Klong Haui Sai, Nong Bua Tai, Mueang Tak; 16°46'24.2"N, 99°06'45.0"E; MUMNH MP00237. • 1 ♀, Klong Mae Sot, Phra That Pha Daeng, Mae Sot; 16°42'24.7"N, 98°36'52.1"E; MUMNH MP00238. Phitsanulok • 13 ♀♀, 7 ♂♂ , Khek, Kaeng Sopha, Wang Thong; 16°53'15.4"N, 100°39'13.3"E; MUMNH MP00239. • 7 ♀♀, 3 ♂♂, Ban Bo, Wang Nok Aen, Wang Thong; 16°51'13.6"N, 100°36'43.2"E; MUMNH MP00240. Phetchabun • 16 ♀♀, 5 ♂♂ , Ban Pho Ngam, Tha Phon, Mueang Phetchabun; 16°35'29.5"N, 101°07'41.6"E; MUMNH MP00241. • 4 ♀♀, 2 ♂♂, Si Thep Historical Park, Si Thep; 15°28'19.7"N, 101°08'48.2"E; MUMNH MP00242. • 1 ovigerous, 1 ♂, Nam Khuen Nam Lang, Wang Hin, Wang Pong; 16°24'20.6"N, 100°47'56.5"E; MUMNH MP00243. • 5 ♀♀, 1 ♂, Klong Nam Phung, Hin Hao, Lom Kao; 16°58'47.8"N, 101°12'51.5"E; MUMNH MP00244. Kamphaeng Phet • 4 ♀♀, 2♂♂ , Khlong Khlung, Wang Sai; 16°11'51.7"N, 99°36'53.0"E; MUMNH MP00245. Nong Khai • 4 ♀♀ , 4 ovigerous, 2 ♂♂, Nam Suai, Song Hong, Mueang Nong Khai; 17°45'01.1"N, 102°51'00.5"E; MUMNH MP00246 . Loei • 5 ovigerous, 1 ♂ , Mekong River , Hat Bia, Pak Chom; 18°03'39.4"N, 101°47'51.8"E; MUMNH MP00247. • 3 ♀♀ , 1 ovigerous, 8 ♂♂, Hueang, Na Chan, Chaing Khan; 17°47'13.7"N, 101°34'31.1"E; MUMNH MP00248. • 2 ♀♀ , 1 ovigerous, 4 ♂♂, Klong Nam Man, Na Ho, Dan Sai; 17°19'35.6"N, 101°08'54.5"E; MUMNH MP00249. • 4 ♀♀, Klong Khok Khamin, Khok Khamin, Wang Saphung; 17°10'24.7"N, 101°50'52.1"E; MUMNH MP00250. • 4 ♀♀, 3 ovigerous, 1 ♂, Ban Wang Kum, Dan Sai; 17°07'27.6"N, 101°10'40.7"E; MUMNH MP00251. • 4 ♀♀, 2 ♂♂, Ban Tha Yang, Phu Kradueng; 16°53'47.6"N, 101°53'18.3"E; MUMNH MP00252. Udon Thani • 1 ♀ , Klong Nam Khong, Thap Kung, Nong Saeng; 17°10'01.5"N, 102°46'03.2"E; MUMNH MP00253. • 1 ♀, 2 ovigerous, 2 ♂♂, Huai Yang, Nong Ya Sai, Wang Sam Mo; 16°59'19.2"N, 103°21'50.0"E; MUMNH MP00254. • 2 ♀♀, 2 ovigerous, Huai Yang, Nong Ya Sai, Wang Sam Mo; 16°57'22.9"N, 103°22'19.9"E; MUMNH MP00255. Sakon Nakhon • 2 ♀♀, 3 ♂♂ , Nam Chan, Akat, Akat Amnuai; 17°35'46.1"N, 104°00'21.6"E; MUMNH MP00256. • 13 ♀♀, 4 ♂♂, Klong Lak, Chiang Khruea, Mueang Sakon Nakhon; 17°15'33.7"N, 104°07'00.1"E; MUMNH MP00257. • 2 ♀♀, 1 ♂ , Nong Han, Tha Rae, Mueang Sakon Nakhon; 17°15'08.5"N, 104°09'32.0"E; MUMNH MP00258. • 3 ♀♀, 2 ♂♂ , Klong Un, Khok Phu, Phu Phan; 17°00'22.1"N, 103°54'50.5"E; MUMNH MP00259. Mukdahan • 3 ♀♀ , Klong Bang I, Nong Weang, Nikhom Kham Soi; 16°23'43.8"N, 104°34'33.7"E; MUMNH MP00260. Kalasin • 1 ovigerous, 4 ♂♂ , Bueng Aram, Khlong Kham, Yang Talat; 16°24'21.8"N, 103°20'26.4"E; MUMNH MP00261. • 2 ♀♀, 3 ♂♂, Huai Sathot, Kham Bong, Huai Phueng; 16°41'32.6"N, 103°51'20.1"E; MUMNH MP00262. Maha Sarakham • 6 ♀♀ , 1 ovigerous, 2 ♂♂, Chi River , Tha Tum , Mueang Maha Sarakham; 16°10'58.2"N, 103°27'19.9"E; MUMNH MP00263. • 1 ♀ , Chi River , Koeng , Mueang Maha Sarakham; 16°13'01.1"N, 103°16'44.1"E; MUMNH MP00264 . Chaiyaphum • 6 ♀♀ , Huai I Muet, Khon San; 16°31'34.6"N, 101°39'28.0"E; MUMNH MP00265. • 3 ♀♀, 1 ovigerous, 2 ♂♂, Pha Iang Waterfall, Huai Ton, Mueang Chaiyaphum; 15°57'42.2"N, 101°54'17.8"E; MUMNH MP00266. • 1 ♀, Na Siao, Mueang Chaiyaphum; 15°55'05.1"N, 102°06'13.8"E; MUMNH MP00267. • 2 ♀♀ , 2 ovigerous, Tat Ton, Na Siao, Mueang Chaiyaphum; MUMNH MP00268. Roi Et • 5 ♀♀, 4 ♂♂ , Ban Nong Phue, Chaturaphak Phiman; 15°49'07.7"N, 103°30'36.9"E; MUMNH MP00269. Ubon Ratchathani • 2 ♀♀ , 2 ovigerous, 2 ♂♂, Mekong River , Khong Chiam, Khong Chiam; 15°19'10.3"N, 105°29'53.2"E; MUMNH MP00270. • 4 ♀♀ , Tung Lung, Nong Saeng Yai, Khong Chiam; 15°20'12.2"N, 105°24'02.6"E; MUMNH MP00271. Si Sa Ket • 1 ♀ , Bueng Bun Local Market, Bueng Bun; 15°19'18.6"N, 104°03'01.2"E; MUMNH MP00272. • 2 ♀♀, 1 ovigerous, 2 ♂♂, Khayung, Thung Yai, Kantharalak 14°34'42.8"N, 104°38'47.5"E; MUMNH MP00273. Nakhon Ratchasima • 9 ♀♀ , 1 ovigerous, 3 ♂♂, Lam Takhong, Mu Si, Pak Chong; 14°33'00.7"N, 101°27'34.1"E; MUMNH MP00274. • 2 ♀♀, 1 ♂, Sema, Sung Noen; 14°55'11.0"N, 101°47'53.5"E; MUMNH MP00275. Lopburi • 3 ♀♀ , 1 ovigerous, 6 ♂♂, Lop Buri Local Market, Tha Sala, Mueang Lop Buri; 14°47'25.2"N, 100°40'27.7"E; MUMNH MP00276 . Nakhon Nayok • 1 ♀ , Khao Perm, Ban Na; 14°21'14.5"N, 101°05'06.9"E; MUMNH MP00277. Chachoengsao • 1 ♂ , Nong Prue Mai Kaeo, Wang Yen, Plaeng Yao; 13°33'31.5"N, 101°17'44.0"E; MUMNH MP00278 . Sa Kaeo • 3 ♀♀ , 1 ovigerous, 3 ♂♂, Huai Yang, Thap Rat, Ta Phraya; 14°07'24.9"N, 102°40'03.6"E; MUMNH MP00279. Chanthaburi • 3 ♀♀, 5 ♂♂ , Wang Kra Prae, Soi Dao; 12°58'18.9"N, 102°16'12.5"E; MUMNH MP00280. • 6 ♀♀, 5 ovigerous, 6 ♂♂, Koh Phasuk, Makham; 12°40'48.9"N, 102°12'08.6"E; MUMNH MP00281. • 10 ♀♀, 11 ♂♂, Phra Sathueng, Thap Chang, Soi Dao; 13°12'29.1"N, 102°10'07.0"E; MUMNH MP00282. • 2 ♀♀, 3 ♂♂, Klong I Ngaeo, Sung, Khlung; 12°27'13.8"N, 102°16'56.1"E; MUMNH MP00283. Trat • 1 ♀ , Tha Sen, Khlong Yai; 12°06'53.3"N, 102°43'16.8"E; MUMNH MP00284. Suphan Buri • 6 ♀♀ , 4 ovigerous, 12 ♂♂, Ban Chara Mai, Ban Kong, U Thong; 14°34'51.0"N, 99°52'04.7"E; MUMNH MP00285. • 1 ovigerous, 1 ♂ , Kra Siao, Dan Chang; 14°50'01.9"N, 99°40'38.4"E; MUMNH MP00286 . Kanchanaburi • 1 ♀ , 1 ovigerous, 3 ♂♂, Huai Khayeng, Khayeng, Thong Pha Phum; 14°38'59.0"N, 98°34'31.3"E; MUMNH MP00287. • 1 ♀, Huai I Su, Nong Ri, Bo Phloi; 14°32'33.3"N, 99°22'41.4"E; MUMNH MP00288. • 3 ♀♀, 5 ♂♂, Taphoen, Lat Ya, Mueang Kanchanaburi; 14°08'14.0"N, 99°22'57.8"E; MUMNH MP00289. • 23 ♀♀, 3 ovigerous, 15 ♂♂, Huai Mae Pla Soi, Na Suan, Si Sawat; 14°34'03.4"N, 99°12'13.6"E; MUMNH MP00290. • 2 ♀♀, 5 ♂♂, Klong Phachi, Klon Do, Dan Makham Tia; 13°55'05.2"N, 99°22'59.3"E; MUMNH MP00291. Ratchaburi • 3 ♀♀ , 1 ovigerous, 2 ♂♂, Khok Mu, Ta Nao Si, Suan Phueng; 13°28'29.5"N, 99°15'48.8"E; MUMNH MP00292. Phetchaburi • 2 ♀♀ , Yang Chum, Klat Luang, Tha Yang; 12°46'55.7"N, 99°40'43.7"E; MUMNH MP00293. • 4 ♀♀, 2 ♂♂, Ban Yang Nam Klat Tai, Nong Ya Plong; 13°06'33.3"N, 99°43'22.8"E; MUMNH MP00294. Prachuap Khiri Khan • 2 ♂♂ , Phongprasan, Bang Saphan; 11°12'53.1"N, 99°30'09.6"E; MUMNH MP00295. Ranong • 1 ovigerous, Bang Khun Paeng, Bang Phra Nuea, La-un; 10°04'06.0"N, 98°43'04.1"E; MUMNH MP00296 . Phang Nga • 1 ♂ , Phu Ta Jor, Le, Kapong; 8°46'04.7"N, 98°27'19.8"E; MUMNH MP00297. • 6 ♀♀, 1 ovigerous, 1 ♂, Thung Maphrao, Thai Mueang; 8°32'56.4"N, 98°19'23.4"E; MUMNH MP00298. Phuket • 1 ♂ , Bang Pae, Pa Klok, Thalang; 8°02'17.8"N, 98°24'12.4"E; MUMNH MP00299. Krabi • 9 ovigerous, 1 ♂ , Pak Nam, Mueang Krabi; 8°04'50.7"N, 98°55'07.3"E; MUMNH MP00300. Songkhla • 2 ♀♀, 1 ♂ , Klong Na Thap , Chana; 7°01'20.1"N, 100°43'51.5"E; MUMNH MP00301 .

Diagnosis.

Rostrum straight or proximal convex and distal margin gently upward. Rostrum length reaches beyond end of antennular peduncle and as long as scaphocerite. Rostral formula: 6-10/1-6 teeth including 1-3 distal teeth with small gap separate from rest. Carapace smooth. Epistome bilobed. First pereiopods reaching end of scaphocerite. Second pereiopods long and slender, similar in form and length, exceeding scaphocerite. Fingers covered with scattered setae, slightly shorter than palm. Translucent razor on cutting edge present anteriorly. Proximal quarter of cutting edges with one or two tiny teeth. Carpus cylindrical shape and articulation margin expanded. Carpus 1.5 × longer than chela. Merus subcylindrical. Carpus 1.5 × longer than merus. Third pereiopods long and slender, reaching end of scaphocerite. Dactylus curved distally with short setae. Propodus 2 × longer than dactylus. Propodus with 4-8 pairs of spines distributed along its length and fine setae at articulation. Propodus 2 × longer than carpus. Sixth and seventh thoracic sternites usually smooth. Eighth thoracic sternite with acute median process. First and second pleonal sternites with small median process. Third and fourth pleonal sternites smooth. Fifth pleonal sternite with triangular ridge. Uropodal diaeresis with inner movable spine shorter than outer angle.

Composite description.

Rostrum (Fig. 4B ). Straight or slightly convex proximally and upward distally. Rostrum length exceeding end of antennular peduncle and slightly shorter than scaphocerite. Dorsal margin with 6-10 teeth including 1-3 teeth distally with small gap from rest. Postorbital margin with one or two teeth, reaching to one-fourth of carapace length. Ventral margin with 1-6 teeth, starting from middle to distal margins. Short setae present between rostral teeth.

Cephalon (Fig. 4B ). Well-developed eye. Ocular beak without laterally expanded tip. Cornea longer and broader than stalk. Postantennular carapace margin rounded. Cornea osculum longer than stalk. Antennular peduncle longer than wide with fine setae, basal segment short, second segment shorter than third segment. Stylocerite projection sharp, reaching beyond basal segment. Antennal spine sharp situated below orbital margin. Hepatic spine slightly larger than antennal spine, positioned posteriorly and lower than antennal spine. Scaphocerite with straight margin, distolateral tooth sharp and not reaching end of lamella. Epistome bilobed (Fig. 4C ). Branchiostegal suture starting from carapace margin to behind hepatic spine. Carapace surface smooth.

First pereiopods. Long and slender, reaching end of scaphocerite. Fingers as long as palm, tips with fine setae. Series of setae present at anterior inner part of palm. Carpus slightly longer than merus. Distal articulation of carpus with series of fine setae. Ischium shorter than merus. Scattered setae present on all segments.

Second pereiopods (Fig. 4D ). Long and slender, similar in form and exceeding scaphocerite. Fingers subcylindrical covered with scattered setae. Palm 1.1-1.4 × longer than fingers. Fingers with translucent razor edge present anteriorly and one or two tiny teeth on proximal quarter of cutting edges. Tip of fingers crossed and covered by fine setae (Fig. 4E ). Carpus cylindrical shape and articulation margin expanded. Carpus 1.3-1.5 × longer than chela. Merus subcylindrical. Carpus 1.1-2 × longer than merus. Scattered short setae present on all segments.

Third pereiopods (Fig. 4F ). Long and slender, reaching end of scaphocerite. Dactylus short and curved distally. Propodus with 4-8 pairs of spines along inferior-lateral margin and fine setae at distal articulation, 2 × longer than dactylus. Propodus 2 × longer than carpus. Short setae present on all segments.

Fourth and fifth pereiopods. Long and slender, exceeding scaphocerite. Propodus of fourth pereiopods with 5-10 pairs of spines distributed along its length, 2 × longer than dactylus. Propodus slightly longer than merus. Ischium shorter than merus. Propodus with fine setae at distal articulation. Scattered short setae present on all segments. Propodus of fifth pereiopods with 7-13 pairs of spines distributed along its length and fine setae at distal articulation. Propodus 2 × longer than carpus. Propodus as long as merus. Scattered short setae present on all segments.

Thoracic sternum. Fourth and fifth thoracic sternites with transverse plate. Sixth and seventh thoracic sternites smooth. Eighth thoracic sternite with or without acute median process.

Pleon. Smooth. All pleonal sternites with transverse ridge. First and second pleonal sternites usually with small median process. Third and fourth pleonal sternites smooth. Fifth pleonal sternite with triangular ridge. Preanal carina present, obtuse ridge developed without spine or setae. Ventral margin of pleural tergum with small setae.

Telson (Fig. 4G ). Tapered posteriorly, protruding point on middle margin with lateral spines and few fine setae. Inner spines longer than outer spines. Dorsal surface with two pairs of small spines similar in size.

Uropods (Fig. 4G ). Uropodal diaeresis with inner movable spine, usually shorter than outer angle. Exopods longer than endopods.

Remarks.

The specimen collected in this study generally agrees with the original description in Lanchester (1902), and a subsequent description of the lectotype provided by Chong and Khoo (1988). Previous studies reported that male specimens tended to display the sexual dimorphism with a large body size, tomentose fingers, and minute spinules on all segments (except fingers) of second pereiopods. In this study, only one large male specimen, collected from Loei Province, Thailand, exhibits this characteristic. Typically, both male and female specimens possess fine setae on fingers and scattered setae on surface of second pereiopods. Furthermore, this study also observed two variable characters occurring on the second pereiopods. Firstly, the proportional length and form of second pereiopods were found to be variable in specimens from Krabi population. Their second pereiopods are shown to be prominently long and robust, similar to those of M. sintangense (a common riverine species). The palm margin is laterally inflated and slightly shorter than fingers, and the chela slightly longer than the carpus. Additionally, Chong and Khoo (1988) reported the presence of two tiny teeth on the basal portion of cutting edges of fingers in M. lanchesteri as a diagnostic character. In this study, one or two tiny teeth were present on the cutting edges of fingers and vary among M. lanchesteri populations. Historically, M. lanchesteri was noted to resemble several other species including M. idae , M. peguense (see under remarks of M. panhai sp. nov.), M. sankollii Jalihal, Shenoy & Sankolli, 1988, M. unikarnatakae Jalihal, Shenoy & Sankolli, 1988, and M. sintangense . Further phylogenetic relationships and phylogenetic placement of aforementioned taxa should be tested to elucidate and verify their taxonomic identities.

Macrobrachium lanchesteri has a wide distribution across mainland Southeast Asia and southern China. This species can live in various freshwater ecosystems by inhabiting aquatic vegetation in stagnant freshwater habitats such as ponds, lakes, and paddy fields.

Figures

Figure 3. Living habit of specimens of three Macrobrachium species A, B M. lanchesteri from Bang Khiat, Singhanakhon, Songkhla, Thailand C, D M. rostrolevatus sp. nov. from Bueng Khong Long, Bueng Kan, Thailand E M. panhai sp. nov. from Sri Nakarin Dam, Tha Kradan, Si Sawat, Kanchanaburi, Thailand. Scale bars: 1 cm.

Figure 4. Morphological characteristics of Macrobrachium lanchesteri (A female topotype MUMNH MP 00218 B-H female topotype MUMNH MP 00216.1) A lateral view B carapace C epistome D second pereiopod E teeth between fingers F third pereiopod G uropod and H movable spine at uropodal diaeresis. Scale bars: 1 mm.