Isometrus maculatus ( De Geer, 1778 )

Kovařík, František, Lowe, Graeme, Ranawana, Kithsiri B., Hoferek, David & Š, V. A., 2016, Scorpions of Sri Lanka (Scorpiones Buthidae, Chaerilidae, Scorpionidae) with description of four new species of the genera Charmus Karsch, 1879 and Reddyanus Vachon, 1972, stat n, Euscorpius 220, pp. 1-133 : 35-40

Isometrus maculatus ( De Geer, 1778)

( Figs. 13 , 127–137 , 196 , 221–222 , 237–238 , 242, 252 , 403–404 , 555 , 557 , 559 )

Scorpio maculatus De Geer, 1778: 346 .

Isometrus maculatus: Kraepelin, 1899: 66 .

Isometrus (Isometrus) maculatus: Vachon, 1972: 169– 180 , figs. 1–13, 15, 17, 19; Fet & Lowe, 2000: 147; Lourenço & Huber, 2002: 266; Teruel & Kovařík, 2012: 88–91, figs. 4, 27, 36–37, 46, 1 94–207, 566– 569; Kovařík & Ojanguren, 2013: 182–184, 347, 355–356, figs. 1251–1255, 1264, 1368–1375, 1376– 1383 (reference and synonymes list until 2013); Veronika & al., 2013: 75, figs. 1, 15–20, tab. 1.

TYPE LOCALITY AND TYPE REPOSITORY. “ Suriname and Pennsylvania ”; NHRS.

S RI LANKAN MATERIAL EXAMINED. Sri Lanka , Northern Province , Mannar District , Madhu Road, 08°48'26.3"N 080°10'26"E, 90 m a.s.l. (Locality 15 CH, Fig. 584 ), 24. –25.IV.2015, 1♂ 1♀, FKCP, 1♀ 2juvs., UPSL, leg. Kovařík et al.; Northern Province, Mannar District, Marichchukkaddi env, border of Wilpattu National Park , 08°33'32.3"N 079°56'51"E, 7 m a.s.l. (Locality 15CI, Fig. 585 ), 25. –26.IV.2015, 1♂ 1♀ ( Figs. 129–130 , 132, 134 , 196 , 222 , 238 , 404 ) 1 juv., FKCP, leg. Kovařík et al.; Southern Province, Matara District, Kekanadura village , 05°58'28.2"N 080°36'20.5"E, 40 m a.s.l. (Locality 15CP, Figs. 593–594 ), 30.IV.2015, 1♂ ( Figs. 127–128 , 131, 133 , 221 , 237 , 242, 252 , 403 , 555, 557, 5 59), FKCP, leg. Kovařík et al.; Eastern Province, Ampara District, Lahugala-Kitulana National Park , 06°52' 46"N 081°43'21.8"E, 40 m a.s.l. (Locality 15CR, Fig. 596 ), 3.–4.V.2015, photos only, leg. Kovařík et al.

DIAGNOSIS. Total length 30–75 mm. Females usually reach ca. 45 mm. Manus of pedipalp very thin, in males its width equals that of patella and femur. Pedipalps and legs yellow, with spots. Manus of pedipalps yellow with several spots, fingers dark. Mesosomal segments lightcolored. First (basal) middle lamella of female pecten quadrangular. Posterior margin of sternite V straight (females) to very slightly convex medially (males). Subaculear tooth spinoid. Pectinal teeth number 15–20 (1 7–20 in the examined Sri Lanka specimens).

COMMENTS ON DISTRIBUTION. I. maculatus has been regarded as cosmopolitan (see Fet & Lowe, 2000: 149) and records of this species are so numerous that a complete listing would be unreasonably long. A list of specimens which the first author examined until 2003 is published in Kovařík (2003: 3). Additonal records are given in Kovařík & Ojanguren (2013: 182–184). Records peaked in the earlier era of wooden sailing ships and harbors full of wooden barracks, both ideal niches for these scorpions. For that reason between 1758 and ca. 1940 this species was reported from many countries and large harbors including Hamburg in Europe. With the transition to modern building materials and modes of travel, reports of this species have greatly diminished, and in many places have ceased altogether. Apparently, relatively few of the old invasive populations were able to persist without being resupplied by a constant influx of new individuals via human transport. The distribution of I. maculatus in southeast Asia is not well documented, but it is possibly indigenous to India and/ or Sri Lanka because of the presence there of its most closely related species: I. thurstoni and I. thwaitesi . Lourenço & Huber (2002: 266) suggested that Sri Lanka was the original home of this species before cosmotropical dispersal by human activity, because Sri Lanka "is the only region in the world where I. maculatus is found in really wild conditions inland." This was pure speculation, unsupported by any data. Firstly, their assertion implies that the authors know all species of scorpions present in every "wild condition inland" everywhere else on the globe. Secondly, their logic is flawed because although certain exotic species may fail to penetrate endemic 'wild' ecosystems of host countries, this is by no means the rule. There are many counterexamples where invasive species have become established in endemic ecosystems and displaced native fauna or flora, causing major negative impacts. However, Veronika et. al. (2013: 75) cited this speculation as fact, claimimg that I. maculatus is an "Endemic species in Sri Lanka ". A proper test of this hypothesis may require comparative DNA studies of widespread I. maculatus populations.

Figures

Figures 12–15: Maps showing the known geographical distribution of the Sri Lankan species.

Figures 127–130: Isometrus maculatus. Figures 127–128. Male from locality 15CP in dorsal (127) and ventral (128) views. Figures 129–130. Female from locality 15CI in dorsal (129) and ventral (130) views.

Figures 131–134: Isometrus maculatus. Figures 131, 133. Male from locality 15CP, chelicerae, carapace and tergites I–III (131) and sternopectinal region and sternite III (133). Figures 132, 134. Female from locality 15CI, chelicerae, carapace and tergites I–III (132) and sternopectinal region and sternite III (134).

Figures 135–137: Isometrus maculatus, female from locality 15CH (135) with newborns before first ecdysis (136), and with juveniles after first ecdysis (137).

Figures 193–200: Distal segments of leg IV, retroventral view of Sri Lankan Buthidae and Chaerilidae genera. Buthoscorpio sarasinorum, male from locality 15CF (193), Charmus laneus, male from locality 15CO (194), Hottentotta tamulus, male from locality 15CK (195), Isometrus maculatus, female from locality 15CI (196), Isometrus thwaitesi, female from locality 15CO (197), Lychas srilankensis, male from locality 15CJ (198), Reddyanus basilicus, male from locality 15CS (199), and Chaerilus ceylonensis, male from locality 15CD (200).

Figures 209–224: Metasomal segments II–III, lateral view of Sri Lankan Reddyanus and Isometrus species. Figures 209–210. Reddyanus basilicus, male (209) and female (210) from locality 15CR. Figures 211–212. R. besucheti, male holotype (211) and female from locality 15CG (212). Figures 213–214. R. ceylonensis sp. n., male holotype (213) and female paratype from locality 15CI (214). Figures 215–216. R. jayarathnei sp. n., male (215) and female (216) paratypes. Figures 217–218. R. loebli, male (217) and female (218) from locality 15CG. Figures 219–220. R. ranawanai sp. n., male holotype (219) and female paratype (220). Figures 221–222. Isometrus maculatus, male from locality 15CP (221) and female from locality 15CI (222). Figures 223– 224. I. thwaitesi, male from locality 15CH (223) and female from locality 15CO (224).

Figures 225–240: Pedipalp, dorsal view of Sri Lankan Reddyanus and Isometrus species. Figures 225–226. Reddyanus basilicus, male (225) and female (226) from locality 15CR. Figures 227–228. R. besucheti, male from locality 15CD (227) and female from locality 15CG (228). Figures 229–230. R. ceylonensis sp. n., male holotype (229) and female paratype from locality 15CI (230). Figures 231–232. R. jayarathnei sp. n., male (231) and female (232) paratypes. Figures 233–234. R. loebli, male (233) and female (234) from locality 15CG. Figures 235–236. R. ranawanai sp. n., male holotype (235) and female paratype (236). Figures 237–238. Isometrus maculatus, male from locality 15CP (237) and female from locality 15CI (238). Figures 239–240. I. thwaitesi, male from locality 15CH (239) and female from locality 15CO (240).

Figures 241–259: Pedipalp chela dorsal (241), movable finger (242–250), and fixed finger (251–259) of Sri Lankan Isometrus, Lychas, and Reddyanus species. Figures 241–242, 252. Isometrus maculatus, female from locality 15CO (241) and male from locality 15CP (242, 252). Figures 243–244, 253. I. thwaitesi, male from locality 15CH (243) and female from locality 15CO (244, 253). Figures 245, 254. Reddyanus basilicus, male from locality 15CS. Figures 246, 255. R. besucheti, male from locality 15CD. Figures 247, 256. R. ceylonensis sp. n., male holotype. Figures 248, 257. R. jayarathnei sp. n., male paratype. Figures 249, 258. R. loebli, male from locality 15CG. Figures 250, 259. R. ranawanai sp. n., male holotype. Figure 251. Lychas srilankensis, male from locality 15CO.

Figures 403–429: Telson, lateral view of Sri Lankan Buthidae and Chaerilidae species. Figures 403–404. Isometrus maculatus, male from locality 15CP (403) and female from locality 15CI (404). Figures 405–406. I. thwaitesi, male from locality 15CH (405) and female from locality 15CO (406). Figures 407–408. Lychas srilankensis, male (407) and female (408) from locality 15CJ. Figures 409–410. Reddyanus basilicus, male (409) and female (410) from locality 15CR. Figures 411–412. R. besucheti, male holotype (411) and female from locality 15CG (412). Figures 413–414. R. ceylonensis sp. n., male holotype (413) and female paratype (414) from locality 15CI. Figures 415–416. R. jayarathnei sp. n., male (415) and female (416) paratypes. Figures 417–418. R. loebli, male (417) and female (418) from locality 15CG. Figures 419–420. R. ranawanai sp. n., male holotype (419) and female paratype (420). Figures 421–422: Buthoscorpio sarasinorum, male from locality 15CF (421) and female from locality 15CG (422). Figures 423–424. Charmus laneus, male (423) and female (424) from locality 15CO. Figures 425–426. Charmus saradieli sp. n. male paratype (425) (MHNG) and female holotype (426). Figurs 427. Hottentotta tamulus, male from locality 15CK (427). Figures 428–429. Chaerilus ceylonensis, male (428) and female (429) from locality 15CD.

Figures 555–570: Key characters for distinguishing males of Sri Lankan Isometrus and Reddyanus. Figures 555–556, 565– 566. Pedipalp chela dorsal view, I. maculatus from locality 15CP (555), I. thwaitesi from locality 15CH (556), R. basilicus, holotype (565), and R. besucheti, holotype (566). Figures 557–558. Sternopectinal area, I. maculatus from locality 15CP (557), and I. thwaitesi from locality 15CH (558). Figures 559–560, 567–570. Sternite V, I. maculatus from locality 15CP (559), I. thwaitesi from locality 15CH (560), R. besucheti, from locality 15CF (567), R. ceylonensis sp. n., holotype (568), R. jayarathnei sp. n., paratype (569), and R. ranawanai sp. n., paratype (570). Figures 561–564. Metasomal segment II lateral (561–562) and pedipalp patella dorsal (563–564), R. loebli from locality 15CG (561, 563) and R. besucheti, from locality 15CF (562, 564).

Figures 582–584: Figures 582–583. Locality 15CG, Sri Lanka, Central Province, Matale District, Habarana, Wananiwahana Resort, 07°59'25.8"N 080°43'24.6"E, 280 m a.s.l. During night collecting on 23.–24.IV.2015 in Wananiwahana resort garden (UV detection) we recorded Buthoscorpio sarasinorum (running on the land among leaves), Reddyanus besucheti (on the land among leaves) and Reddyanus loebli (hidden under scales of barks of standing tree trunks, see the blue arrows on the Fig. 583). On the locality we recorded night temperature 28.4 ºC– 24 ºC (minimum temperature) and night humidity varied between 55% and 87%. Figure 584. Locality 15CH, Sri Lanka, Northern Province, Mannar District, Madhu Road, 08°48'26.3"N 080°10'26"E, 90 m a.s.l. During night collecting on 24.–25.IV.2015 (UV detection) we recorded Isometrus maculatus and Isometrus thwaitesii sympatrically (running on branches and trunks of trees, also siting on leaves one to four metres high), Reddyanus ceylonensis sp. n. (on the land among leaves), Reddyanus loebli (hidden under scales of barks of standing tree trunks).

Figures 585–587: Figure 585. Locality 15CI, Sri Lanka, Northern Province, Mannar District, Marichchukkaddi env, border of Wilpattu National Park, 08°33'32.3"N 079°56'51"E, 7 m a.s.l. During night collecting on 25.–26.IV.2015 (UV detection) we recorded Isometrus maculatus and Isometrus thwaitesii sympatrically (running on branches and trunks of trees, also sitting on leaves one to four metres high), Reddyanus ceylonensis sp. n. (on the land among leaves; the type locality), Reddyanus loebli (hidden under scales of bark of standing tree trunks). Near to the locality we recorded temperature between 37.6 ºC (maximum daytime temperature and 26.2 ºC (minimum nighttime temperature) shortly before sunrise. Humidity varied between 67% and 81%. Figures 586–587. Locality 15CJ, Sri Lanka, Northern Province, Jaffna District, 09°42'51.6"N 080°04'44.8"E, 19 m a.s.l. During night collecting on 26.–27.IV.2015 (UV detection) we recorded Lychas srilankensis (on the land usually under branches, leaves or in their vicinity), Heterometrus swammerdami (inside the termite constructions or inside heaps of garden "waste") and Heterometrus gravimanus inside burrows in open terrain (Fig. 587). The females had 30 to 60 cm long oblique burrows once or twice curved. A female shortly after maturity ecdysis was in the burrow together with a male; another female was in the burrow together with three juveniles which had their own narrow burrows separated under surface from the motherś burrow; a juvenile after the third ecdysis had a proper burrow 30 cm long oblique and once curved. There also lives Isometrus maculatus. On the locality we recorded night temperature 31.6 ºC– 27.2 ºC (minimum temperature) and night humidity varied between 60% and 87%.

Figures 592–594: Figure 592. Locality 15CO, Sri Lanka, North Central Province, Puttalam District, Eluwankulam, 08°17'15"N 079°50'38.7"E, 38 m a.s.l. During night collecting from 10.30 p.m. on 28.IV.2015 to 4.00 a.m. on 29.IV.2015 (UV detection) we recorded Charmus laneus (running on the ground among leaves), Isometrus thwaitesii (running on branches of bushes, also sitting on leaves), Lychas srilankensis (on the ground under branches, leaves or in their vicinity), Reddyanus ceylonensis sp. n. (on the ground), Chaerilus ceylonensis (inside oblique usually not curved burrows 15 to 30 cm long) and Heterometrus swammerdami (inside long burrows). On the locality we recorded between 10.00 p.m. and 4.00 a.m. temperature 25.9 ºC and humidity between 58% and 70%. On the locality we recorded 6 scorpion species and genera of all three Sri Lankan families. Figures 593–594. Locality 15CP, Sri Lanka, Southern Province, Matara District, Kekanadura village, 05°58'28.2"N 080°36'20.5"E, 40 m a.s.l. During night collecting on 30.IV.2015 (UV detection) we recorded Isometrus maculatus (inside a house, the species occupy the roof of the house, Fig. 593) and Heterometrus serratus (inside almost straight burrows in a ground wall). On the locality we recorded night temperature 28.4 ºC– 25.2 ºC (minimum temperature) and night humidity varied between 63% and 89%.

Figures 595–596: Figure 595. Locality 15CQ, Sri Lanka, Uva Province, Monaragala District, Monaragala, 06°52'30.7"N 081°21'17"E, 288 m a.s.l. During night collecting on 2.–3.V.2015 (UV detection) we recorded Reddyanus loebli (under scales of bark and under bark) and Heterometrus serratus (females with juveniles inside oblique burrows). On the locality we recorded a temperature of 33.1 ºC at 5.00 p.m., which gradually dropped to 24.6 ºC (minimum temperature) before sunrise. Humidity during the night varied between 61% and 84%. Figure 596. Locality 15CR, Sri Lanka, Eastern Province, Ampara District, Lahugala Kitulana National Park, 06°52'46"N 081°43'21.8"E, 40 m a.s.l. During night collecting on 3.–4.V.2015 (10.00 p.m. to 01.00 a.m.) we recorded Isometrus maculatus (running on branches, also sitting on leaves on trees), Reddyanus basilicus (on the ground among leaves) and Reddyanus loebli (hidden under scales of bark of standing tree trunks). On the locality we recorded between 10.00 p.m. on 3.V. and 01.00 a.m. on 4.V.2015 temperature 28 ºC, which gradually dropped to 26.9 ºC and humidity 73%.