Redefinition And Systematic Revision Of The East African Scorpion Genus Pandinoides (Scorpiones: Scorpionidae) With Critique Of The Taxonomy Of Pandinus, Sensu Lato Author Prendini, Lorenzo Scorpion Systematics Research Group, Division of Invertebrate Zoology, American Museum of Natural History text Bulletin of the American Museum of Natural History 2016 2016-09-21 2016 407 1 66 http://www.bioone.org/doi/10.1206/0003-0090-407.1.1 journal article 7634 10.1206/0003-0090-407.1.1 cf139a31-bcd0-4bed-b887-67bd8f9b7f12 0003-0090 4612737 958953B4-BEA7-4528-8D54-C431CED61A40 Pandinus ( P. ) imperator (C.L. Koch, 1841 ) Buthus imperator C.L. Koch, 1841: 1 , 2, pl. CCLXXXIX, fig. 695. Heterometrus roeseli Simon, 1872: 53 , 54, pl. VI, fig. 4 (first synonymized by Thorell, 1893: 377 ), syn. nov. Pandinus africanus Thorell, 1876b: 128 , 202, 203 (synonymized by Fet, 2000: 466 ). Scorpio simoni Becker, 1880: 137–140 , pl. II, fig. 1 (synonymized by Thorell, 1893: 377 ). Pandinus ( P. ) camerounensis Lourenço, 2014: 143–149 , table 1, figs. 12–19, syn. nov. TYPE MATERIAL : Buthus imperator : [types lost; type locality unknown]. Heterometrus roeseli : [types lost or not designated; type locality probably in coastal Guinea ( Lourenço, 2014: 142 )]. Pandinus ( P. ) camerounensis : Holotype ♂ ( ZMH ), 3 ♂ , 2 ♀ paratypes ( MNHN , ZMH ), Sanguéré-Djoi / Kismatari / Djalingo regions (ca. 09°23.229′N– 013°50.068′E ), Cameroon , viii.2011 – xi.2012 , P. Prudent , cotton and tomato fields, some collected in termite mounds. REMARKS: Lourenço (2014) revalidated Pandinus roeseli ( Simon, 1872 ) , originally synonymized by Thorell (1893) and described a new species of Pandinus , P. camerounensis , in a contribution he presumably considered to be “serious research.” The justification for Lourenço’s (2014: 143 , 147, 149) decisions was based on the following argument (italics added): If the trichobothrial patterns for both P. dictator and P. gambiensis are clearly diagnostic, a number of semi-distinct populations from Western Africa can be associated to P. imperator (Lourenço & Cloudsley-Thompson 1999) . The study of some Pandinus recently collected in the north of Cameroon (in a zone of transition between the Sahel and Savannas) showed a similar trichobothrial pattern but exhibited other distinctive characters to that of ‘ P. imperator ’ collected from areas of rain forests. For this reason, I decided to explore the morphology of the hemispermatophores. This character has been largely used in many similar groups with great success (e.g. Lamoral, 1979 , Lourenço 1987 , 2009 ), but was largely neglected in the study of Pandinus . I also extended this analysis to male specimens of the Coastal region of Guinea (cf. P. roeseli ) and to specimens from south of Togo / Ghana , ‘a priori’ cospecific with P. imperator . The results are presented in the key below (see also Figures 8–12 ). Since the hemispermatophore structure is quite distinct for these three populations, P.roeseli is reestablished here as a valid species, and a new species is described from the North of Cameroon . Further studies will be necessary to clarify the status of all West-African populations and future molecular investigation would contribute greatly to this subject …. Pandinus camerounensis sp. n. , is here distinguished from Pandinus imperator ( Koch, 1841 ) and from Pandinus roeseli ( Simon, 1872 ) , the two most geographically related species of the genus, mainly by their global size and structure of the hemispermatophores. The first two species are bigger in size, reaching total lengths of 150 to 180 mm . The structure of hemispermatophores are quite distinct in the three species (see Figs 8–12 ). In P. imperator the distal lamina is weakly curved and the basal portion larger than the distal one; a tubercular structure is present on its apex. In P. roeseli the distal lamina is shorter than in the other two species and globally large. Lourenço’s (2014: 145 , 147) “serious” analysis appears to be based on five specimens, in addition to the six comprising the type series of P. camerounensis : a male P. roeseli from Kindia, Guinea ; a male P. imperator from SE Notsé, southern Togo ; a female P. imperator from Edéa, southern Cameroon ; and two female P. imperator from Eboname [actually, Ebomane], northern Gabon . Aside from the small sample size, the identification of the alleged P. imperator specimens from Cameroon and Gabon is questionable, because there are no credible records of this species east of the Massif de l’Adamaoua, between Cameroon and Nigeria ( Prendini, 2004 ). Neither Rossi (2014d: 49 , fig. 1), who claimed that data were included from Lourenço (2014) , nor Kovařík (2011: 15 , fig. 42) before him plotted a single point for P. imperator in southern Cameroon or northern Gabon in the maps each author reproduced from Prendini’s (2004: 255) fig. 13. The putatively diagnostic difference in total body length between P. camerounensis ( 95–110 mm ) vs. P. imperator and P. roeseli ( 150–180 mm ), largely meaningless given the small sample size, breaks down when the raw data presented by Lourenço (2014: 147) are examined more closely. The specimen of P. roeseli from Kindia is cited as 115 mm in total length, the specimen of P. imperator from Notsé, 155 mm , and the specimen from Edéa, almost certainly referable to P. dictator , 163 mm . An assessment of size variation among the material examined for the present study suggests the existence of a north-south cline in body size, associated with an aridity gradient, across the entire range of P. imperator , such that individuals in the arid north, e.g., Mouvielo, Burkina Faso (NM 14004), are on average smaller than those in the humid south, e.g., Kakum National Park, Ghana (AMNH), rendering indistinct any populations thought to be “semi-distinct” based on the examination of a handful of specimens. The putatively diagnostic differences in hemispermatophore structure between P. imperator and P. roeseli described by Lourenço (2014: 147 , 149), which cannot be discerned from the superficial illustrations presented, must also be disregarded given the lack of quantification and, especially, the small sample size (although unspecified, it may be concluded that only three hemispermatophores were examined, one from each putative species). Finally, in response to Lourenço’s (2014) comment regarding the need for molecular investigation, it may be noted that ongoing research has revealed low levels of genetic divergence between small-bodied and large-bodied populations of P. imperator from the rainforests and Sahel regions of West Africa, supporting the morphological evidence of a sin- gle panmictic population (L. Prendini, unpublished data). In view of the above, H. roeseli is hereby returned to synonymy with P. imperator and P. camerounensis newly synonymized therewith: Heterometrus roeseli Simon, 1872 = Pandinus ( P. ) imperator (C.L. Koch, 1841 ) , syn. nov. ; Pandinus ( P. ) camerounensis Lourenço, 2014 = Pandinus ( P. ) imperator (C.L. Koch, 1841 ) , syn. nov. ADDITIONAL MATERIAL EXAMINED: 1 ♂ (AMNH). West Coast Africa, G.A. Perkins, 1 ♀ (MCZ). W. Africa, 1997, 1 juv. ♀ (NHRM [JF 68]), 1996, G. Alroth, 1 ♀ (NHRM [JF 69]). BENIN : Alibori Dept.: Banikoara Commune : Chutes de Koudou, ‘W’ Park, 11°40.43′N 003°18.51′E , 31.v.2005 , V. Vignoli and S. Tchibozo, 229 m , 1 subad. ♂ (AMCC [LP 4837]), 1.vi.2005 , V. Vignoli and S. Tchibozo, 258 m , 1 ♀ (AMCC [LP 4833]). Karimama Commune : near Djona on Alibory River, 11°40′N 002°50′E , v.1964 , J.A. MacKallor, 1 juv. ♂ (USNM). Atakora Dept.: Tanguiéta Commune : Pendjari Park, 11°27.07 ′N 001°34.02′E , 8.vi.2005 , V. Vignoli & S. Tchibozo, 171 m , 1 juv. ♂ (AMCC [LP 4832]). Borgou Dept.: Bembèrèkè Commune : Riviere Dere, Forêt Classée d’Ouenou-Benou, Gando village, 10°12.09′N 002°39.05′E , 3.vi.2005 , V. Vignoli and S. Tchibozo, 420 m , 1 ♀ (AMCC [LP 4836]). Collines Dept.: Dassa-Zoumè Commune : ‘La caverne,’ 07°46.32′ N 002°11.12′E , 13.vi.2005 , V. Vignoli and S. Tchibozo, 187 m , 1 ♀ (AMCC [LP 4834]), 13.vi.2005 , V. Vignoli and S. Tchibozo, 187 m , 1 juv. (AMCC [LP 4835]). Savé Commune : Ferme Founfoun à 1 km de Savacon [Savakon, 07°15′N 002°04′E ], 30.iii.1997 , D. Meirte, sous bois mort, 1 subad. ♂ (MRAC 208.379). BURKINA FASO : Sud-Ouest Region: Bougouriba Prov. : Mouviélo [ 10°46′N 003°08′W ], Upper Volta, s/p Diébougou [ 10°58′N 003°15′W ], 30.iv.1980 , Dr Prost, de nuit au sol [at night on ground], 1 ♀ (NM 14004). CÔTE D’IVOIRE: Sismos, 13.ii.1963 , J.F. Jezequel, 1 ♂ (MNHN RS 4149). SNB, 24.iii.1963 , pres. Cpt. Tenko, J.F. Jezequel, 1 ♂ (MNHN RS 4150). Abidjan Autonomous Distr: Adiopo Doumé [ 05°20′N 004°07′W ], 17.viii.1966 , 1 ♀ (MRAC 130.708); Ivory Coast, Banco Forest [ 05°22′N 004°03′W ], ii.1989 , J. Visser, 2 ♀ (SAM C4509, C4510), 1 subad. ♂ (SAM C4508); Bingerville, 05°21′N 003°53′W , 1962, J. Decelle, 2 ♂ (MRAC 123.726), xii.1963 , J. Decelle, 1 ♂ (MRAC 126.978). Comoé Distr.: Indénié-Djuablin Region : Appouessou, Forêt Classée Bossematie, Eco. Forest, 06°35′N 003°28′W , pitfall, Station 2E, 18.iii.1994 , R. Jocqué and N. Séabé, 1 juv. ♀ (MRAC 205.479), Station 2B, 3.v.1994 , R. Jocqué and N. Séabé, 1 juv. ♀ (MRAC 205.480), Station 2C, 9.iv.1995 , Jocqué and Tanoh, 1 juv. ♀ (MRAC 205.309), Station 2B, 9.iv.1995 , Jocqué and Tanoh, 1 juv. ♀ (MRAC 205.310). Lagunes Distr.: Agnéby- Tiassa Region : Lamto, station écologique, 06°13′N 005°02′W , 2.iv.1963 , J.F. Jezequel, dans une case, 1 ♂ (MNHN RS 4148). Montagnes Distr.: Cavally Region : Toyébli [Tiobli, 06°37′N 008°29′W ], 2–3.viii.1966 , Verheyen and Thys v.d. Audenaerde, 7 ♂ , 13 ♀ (MRAC 130.717), 29–30.viii.1966 , Verheyen and Thys v.d. Audenaerde, 11 ♂ , 9 ♀ (MRAC 130.718); Village km 7 route Toulépleu–Liberia [ 06°32′N 008°30′W ], 2.viii.1966 , Verheyen and Thys v.d. Audenaerde, 16 ♂ , 6 ♀ (MRAC 130.719), 20 ♂ , 9 ♀ (MRAC 130.720). Tonkpi Region : Flampleu, 07°17′N 008°03′W , 20–24.vii.1966 , W. Verheyen and D. Thys van den Audenaerde, 2 juv. ♂ (MRAC 131.048, 131.049). Sassandra- Marahoué Distr.: Marahoué Region : Danangoro, 07°11′N 005°56′W , iii.1977 , P.M. Elsen, 1 juv. ♂ (MRAC 160.473). Vallée du Bandama Distr.: Gbêkê Region : Bouaké Sessénoua (forêt), 07°41′N 005°09′W , 9.vii.1962 , J.F. Jezequel, 1 ♀ (MNHN RS 4202), 1964, J.F. Jezequel, galeries forestieres et case, 1 ♂ (MNHN RS 4207), 1 ♂ , 1 ♀ (MNHN RS 4208), 23.iii.1964 , J.F. Jezequel, 1 ♂ (MNHN RS 4203). Yamoussoukro Autonomous Distr: Kossou, 06°57′N 004°58′W , 13.iv.1975 , R. Jocqué, 1 subad. ♂ (MRAC 160.528), 13.v.1975 , R. Jocqué, 2 juv. ♂ (MRAC 160.529), v.1975 , R. Jocqué, 2 juv. ♂ , 1 juv. ♀ (MRAC 161.991). Vallée du Bandama Distr.: Gbêkê Region : Bouaké, 07°41′N 005°02′W , vi.1977 , P.M. Elsen, 1 juv. ♀ (MRAC 160.472), v.1977 , P.M. Elsen, 1 juv. ♂ (MRAC 160.474). Hambol Region: Chaussée de Badika[ha], riv. Bandama blanc, 09°12′N 005°10′W , 22.viii.1985 , G. Teugels, 1 juv. ♂ (MRAC 168.783). GHANA : F. Cotterill, 2 ♂ , 1 subad. ♂ , 1 juv. ♂ (NHMZ 13/27). Imported for pet trade: x.1997 , ex D. Taylor, 1 ♂ (AMCC 101703 [LP 1601]). Brong- Ahafo Region: Mive, Brongh Ahafo, 26. iv.1991 , Tyle, 1 ♂ (NHRM [JF 48]). Central Region : Cape Coast Distr. : Cape Coast [ 05°06′N 001°14′W ], Univ. Cape Coast Collection, J. Boggs, 1 ♂ , ♀ , 5 juv. (AMNH); Cape Coast, 15.vi.1969 , 1 subad. ♂ (USNM). Twifo/ Heman/Lower Denkyira Distr. : Kakum National Park: 05°21.359′N 001°23.016′W , 24.i.2014 , S.M. Mwangi, 654 ft , UV in deep burrow, 1 subad. ♀ , 2 juv. ♂ , 10 juv. ♀ (AMNH); Abrafu village, 05°20.175′N 001°22.689′W , 27.i.2014 , S.M. Mwangi, 403 ft , 1 ♂ , 2 subad. ♀ , 1 juv. ♀ (AMNH), 3 juv. ♂ , 2 juv. ♀ (AMNH), 1 juv. ♂ (AMCC [LP 12326]); 05° 20.480 ′ N 001°22.661′W , 27.i.2014 , S.M. Mwangi, 540 ft , at night with UV, 1 ♂ , 2 subad. ♂ , 2 subad. ♀ (AMNH). Kakum Regeneration Forest, Abrafu village, 05°18.918′N 001°22.552′W , 28.i.2014 , S.M. Mwangi, 451 ft , 1 ♂ , 3 subad. ♂ , 3 subad. ♀ , 1 juv. ♀ (AMNH). Eastern Region : East Akim Distr. : Tafo [ 06°13′N 000°22′W ], residential area, in garden of bungalow, at base of dead plantain, excavated chambers in soil terrarium, iii–iv.1946, R.G. Donald, 2 ♂ , 1 juv. (AMNH). Volta Region : South Dayi Distr. : Wegbe [ 07°07′N 000°27′E ], Togoland, W. Innes, 1899, 1 ♀ (SAM 6353). Western Region: Jomoro Distr. : Ankasa National Park, 05°12.928′N 002°39.557′W , 30.i.2014 , S.M. Mwangi, 228 ft , collected in casava farm, killed by farmer, 1 ♂ (AMNH). Juabeso-Bia Distr. : Bia National Park (Kunkumso Park): 06°37.327′N 003°03.342′W , 3.ii.2014 , S.M. Mwangi, 823 ft , during day in cocoa farm adjacent to park, 1 ♀ , 1 subad. ♂ , 3 juv. ♂ (AMNH), 1 ♀ , 17 juv. ♂ , 10 juv. ♀ (AMNH), 1 juv. ♂ (AMCC [LP 12325]); near Appa So Twin wells, 06°36.526′N 003°04.449′W , 3.ii.2014 , S.M. Mwangi, 733 ft , at night with UV near the burrow, 1 subad. ♂ (AMNH); camp 15, 06°32.665′N 003°01.965′W , 3.ii.2014 , S.M. Mwangi, 767 ft , on ground, killed by villager in forest, 1 ♂ (AMNH). LIBERIA : Colonization Society, J.O. Wilson, 1 ♂ (USNM 30467). Bong Co.: Fuamah Distr. : Dobli Island [ 06°53′N 010°23′W ], Bequaert, 1 ♂ (MCZ). Jorquelleh Distr. : Gibanga [Gbanga, 07°00′N 009°28′W ], Harvard Exped., 3 ♂ , 2 ♀ , 1 subad. ♂ , 1 juv. ♂ (MCZ). Grand Bassa Co.: Distr. 4 : Ganta [ 05°38′N 009°48′W ], 1932, 2 ♀ (AMNH 32332). Gbarpolu Co.: Belleh Distr. : Bell[e] Yella [Beliyela, 07°23′N 010°00′W ], iii.1940 , W.M. Mann, Smithsonian Institution- Firestone Expedition, 1 ♂ (USNM). Grand Gedeh Co.: Tchien Distr. : Zwedru (Tchien) [ 06°04′N 008°08′W ], Eastern Prov., J.J. Baldwin Jr., 1 ♂ (USNM 177246); Zwedru [ 06°01′N 008°09′W ], 6.vi.1947 , Jensen, 1 juv. ♀ (USNM). Margibi Co.: Gibi Distr. : Gibi [ 06°40′N 010°00′W ], 1940, W.M. Mann, Smithsonian Institution-Firestone Expedition, 1 ♂ , 1 subad. ♂ (USNM). Montserrado Co.: Careysburg Distr. : Mt. Coffee [ 06°30′N 010°36′W ], 1896, O.F. Cook, 5 ♂ , 1 ♀ (USNM), 1899–1900, Rev. George P. Goll, 1 ♀ (USNM). Greater Monrovia Distr. : Monrovia [ 06°18′N 010°48′W ], 1 ♂ (MCZ), 29.vii.1963 , R.N. Nilson, 2 ♂ (CAS), 06°19′N 010°48′W , 1975, W.G. Johnson, 8 ♂ , 5 ♀ (MRAC 147.335). Nimba Co.: Sanniquelleh-Mahn Distr. : Oldtown Gobonwea, 225 mi from Monrovia, 40 miles E Mt. Nimba [ 07°33′N 008°37′W ], Charles D. Miller III, at night, with aid of light, they frequent rocky outcrops in the bush, frequently come out during the day after heavy rains, 2 ♂ , 2 ♀ (AMNH), 4 ♂ , 3 ♀ (USNM); Yekepa Nimba, 07°35′N 008°32′W , 15.iii.1980 , M. Louette and P. Rigaux, 1 ♂ (MRAC 155.160). NIGERIA : x.1996 , 1 ♂ (NHRM [JF 49]). Kaduna State : Kaduna Local Government Area ( LGA ): Kaduna [ 10°36′N 007°27′E ], N Nigeria, ii.1958 , W. McDonald, 1 ♀ (AMNH); Randa [ 09°07′N 008°29′E ], N Nigeria, Rev. Judd, 1 ♂ , 1 ♀ , 1 subad. ♀ (AM 4657). Kogi State : Idah LGA : Iyale, 60 miles S [N] Idah [ 07°39′N 007°17′E ], Kwara State, i.1970 , L. Dick, 1 juv. ♀ (AMNH). Nassarawa State : Keana LGA : Keana [ 08°08′N 008°48′E ], N Nigeria, 2 ♀ (AM 4008, 4050), 1 subad. ♂ (AM 4268). Lafia LGA : Wana [ 08°50′N 008°28′E ], N Nigeria, 1 ♂ , 2 ♀ (AM 8481), 1 juv. ♂ (AM 6644). Ogun State : Odeda LGA : Oloke Meji [ 07°25′N 003°32′E ], Ibadan, 2 ♀ (USNM). Oyo State : Ibadan LGA : Ibadan, 07°14′N 003°50′E , ix.1968 , S. Afolabi Toye, 1 ♂ , 1 ♀ (MRAC 134.308). Plateau State : Jos LGA : Jos, 09°55′N 008°54′E , 7–26.iv.1963 , Meussen and Bouguiaux, 1 juv. ♀ (MRAC 123.774), x–xii.1965, E. Bot Gwong, 1 juv. ♂ , 2 juv. ♀ (MRAC 130.639). SIERRA LEONE : Northern Region: Port Loko Distr. : Pepel Town, 08°35′N 013°03′W , ix.1976 , D. Olu-Pitt, 1 juv. ♂ (MRAC 148.502). Western Region: Western Area Urban Distr. : Freetown, 08°30′N 013°15′W , Pepel Town, iv.1977 , D. Olu-Pitt, 1 juv. ♀ (MRAC 159.087). TOGO : 1 ♂ , 3 ♀ , 1 juv. (AMNH); viii.2004 , B.R. Tomberlin, 1 ♀ (AMCC [LP 3082]). Centrale Region : Soutouboua Prefec. : Fazao [ 08°42′N 000°46′E ], 20–24.viii.1969 , F. Puylaert, 1 ♂ , 2 ♀ (MRAC 200.957 ex 135.994). Kara Region : Assoli Prefec. : Bafilo, Aledje [Alédjo], 09°15′N 001°12′E , 19.vii.1969 , F. Puylaert, 1 subad. ♀ (MRAC 136.007). Doufelgou Prefec. : Niamtougou, 09°46′N 001°06′E , 21–24.vii.1969 , F. Puylaert, 2 ♂ , 1 ♀ (MRAC 136.005). Maritime Region : Lacs Prefec. : Togoville, 06°14′N 001°29′E , 11–17.ix.1969 , F. Puylaert, 1 ♂ (MRAC 136.006), 1 ♀ (MRAC 200.958 ex 135.995), 2 juv. ♂ , 2 juv. ♀ (MRAC 136.008). Plateaux Region : Ogou Prefec. : Atakpamé, Kolekepe [Kolokopé], 07°28′N 001°19′E , 26–31.viii.1969 , F. Puylaert, 1 ♂ (MRAC 136.004); Badou, 07°37′ N 000°37′E , 17.vii.1968 , W. Verheyen and coll., 1 ♂ (MRAC 134.644). Savanes Region : Kpendjal Prefec. : Nanergou, 10°55′N 000°09′E , 17.viii.1968 , W. Verheyen and coll., 7 ♂ , 11 ♀ (MRAC 134.642). ERRONEOUS RECORDS: Kenya , 1 ♂ (AMNH). Lake Mary, 5 mi. SW Sanford, Seminole Co., Florida , U.S.A. , v.1984 , sent by D. Viekers, Univ. Central Florida , Orlando, 1 ♂ (AMNH 32816).