Pandinoides cavimanus (Pocock, 1888) Figures 1A, B, 2, 3–9; table 1

Type Material: Holotype ♂ (BMNH 1863.66) [examined], TANZANIA: Tabora Prov.: Uyui Distr.: Central Africa, between the center and Umyamuezi [04°59′S 033°10′E] and 5° to 7°S latitude, Captain Speke.

Fet (2000: 468) mistakenly listed two syntypes and Kovařík (2003: 149) erroneously designated one, BMNH 1863.66 from Umyamuezi, the lectotype (also see Kovařík, 2009: 58). Despite mentioning two specimens in the original description of Scorpio cavimanus, Pocock (1888: 249) explicitly designated one as the type: “I have seen two specimens … one (dried) brought from Kilima-Njaro by Mr. M. J. Jackson; the other, which, being preserved in spirit of wine, I have selected as the type, brought by Capt. Speke from Umyamuezi.”

Diagnosis: Pandinoides cavimanus may be separated from P. duffmackayi, sp. nov., and P. militaris, as follows. Pandinoides cavimanus is usually brownish black (fig. 1A, B), rarely reddish brown, in color whereas P. duffmackayi, sp. nov., is reddish brown to reddish black (fig. 1C) and P. militaris, yellowish brown (fig. 1D) to reddish brown. The interocular and circumocular surfaces of the carapace are sparsely and finely granular anterior to the median ocular tubercle, along the median longitudinal sulcus, and often on the frontal lobes in P. cavimanus (fig. 4A, C), but smooth, or nearly so, in P. duffmackayi, sp. nov. (fig. 11A, C), and smooth or with a few granules anterior to the ocular tubercle in P. militaris (fig. 18A, C). The metasoma of P. cavimanus is 52%–56% (♂) and 49%–53% (♀) of total length, the summed lengths of segments IV and V, 115%–130% (♂), 107%–120% (♀) of carapace length (table 1), whereas the metasoma of P. duffmackayi, sp. nov., is 49%–53% (♂) and 46%–49% (♀) of total length, the summed lengths of segments IV and V, 105%–116% (♂) and 91%–110% (♀) of carapace length (tables 2, 3), and the metasoma of P. militaris is 46%–52% (♂) and 42%–50% (♀) of total body length, the summed lengths of segments IV and V, 98%–117% (♂) and 87%–105% (♀) of carapace length (tables 4, 5). The dorsal surfaces of metasomal segments I–IV in the male are finely and sparsely granular in P. cavimanus (fig. 9A) but smooth in P. duffmackayi, sp. nov. (fig. 16A), and P. militaris (fig. 23A). The ventral surface of the telson vesicle of the male bears obsolete carinae, each comprising isolated spiniform granules, restricted to the anterior half or third, in P. cavimanus (fig. 9B, C), whereas the vesicle is smooth or nearly so in P. duffmackayi, sp. nov. (fig. 16B, C), and bears distinct carinae, each comprising prominent spiniform granules, extending its entire length, in P. militaris (fig. 23B, C). The proximal lobe on the pedipalp chela fixed finger of the adult male is slightly smaller than the medial lobe, creating a moderate gap proximally between the fixed and movable fingers, when closed in P. cavimanus (fig. 6A), whereas the proximal lobe is similar in size to the medial lobe, creating little to no gap proximally between the fingers, when closed in P. duffmackayi, sp. nov. (fig. 13A), and it is vestigial, much smaller than the medial lobe, or absent, creating a prominent gap proximally between the fingers, when closed in P. militaris (fig. 20A).

Based on unpublished genetic data, P. cavimanus is most closely related to P. duffmackayi, sp. nov., from which it may be further separated as follows. Pandinoides cavimanus is larger, with total adult body length 87–111 mm, carapace length 15–18 mm and pedipalp chela length 21–29 mm (table 1), than P. duffmackayi, sp. nov., with total adult body length 60–78 mm, carapace length 10–13 mm and chela length 14–19 mm (tables 2, 3). Pandinoides cavimanus bears 83–91 trichobothria on the pedipalp, with 48–55 trichobothria on the patella, including 31–37 in the v series, whereas P. duffmackayi, sp. nov., bears 74–82 trichobothria on the pedipalp, with 39–45 trichobothria on the patella, including 22–28 in the v series. The pectinal tooth count of P. cavimanus, with 14 or 15, usually 14 (♂) and 13 or 14, usually 13 (♀), is higher than that of P. duffmackayi, sp. nov., with 11–13, usually 12 (♂) and 10–12, usually 11 (♀). The prolateral t and st macrosetae of basitarsi III and IV are spiniform in P. cavimanus (fig. 8C, D), whereas t and st are setiform on III, and t setiform and st spiniform on IV in P. duffmackayi, sp. nov. (fig. 15C, D).

Pandinoides cavimanus has often been confused with P. militaris due to their similar size and high trichobothrial counts, but may be separated from the latter as follows. The carapace is less compressed, dorsoventrally, in P. cavimanus (fig. 4A, C) than in P. militaris (fig. 18A, C). The median ocular tubercle is posteromedial, its distance from the anterior carapace margin 51%–56% of the carapace length in P. cavimanus (table 1), but medial, its distance from the anterior carapace margin 46%–52% of the carapace length, in P. militaris (tables 4, 5). The ventrosubmedian carinae on metasomal segment IV are distinct, costate in P. cavimanus (fig. 9C), but obsolete, granular in P. militaris (fig. 23C). The chela manus of P. cavimanus is broader, the length along the retroventral carina 56%–63% (♂) and 58%–67% (♀) of the width, and more convex, the height 59%–73% (♂) and 66%–76% (♀) of the width, than the manus of P. militaris, in which the length along the retroventral carina is 64%–76% (♂) and 62%–72% (♀) of the width, and the height 49%–68% (♂) and 64%–73% (♀) of the width. The chela fixed finger of P. cavimanus arises abruptly from the manus and is wider proximally (figs. 6A, 7) than that of P. militaris, which arises more gradually from the manus (figs. 20A, 21). The chela fingers and distal surfaces of the manus are moderately (♂) to sparsely (♀) setose in P. cavimanus (fig. 1B), but densely (♂) to moderately (♀) setose in P. militaris (fig. 1D). Pandinoides cavimanus bears 31–35 trichobothria on the pedipalp chela, including 7–10 in the V series, whereas P. militaris bears 33–36 trichobothria on the chela, including 9–12 in the V series. The angle of the first proximal median lamella (scape) of the pecten is smaller (more acute), especially in the female, of P. cavimanus (fig. 4B, D), compared with P. militaris (fig. 18B, D).

Redescription: The following redescription supplements the original description by Pocock (1888) and is based on the material examined.

Total Length: Adult medium, maximum length, measured from anterior margin of carapace to tip of aculeus, 106 mm (104–111 mm, n = 4) (♂), 95 mm (87–105 mm, n = 7) (♀) (table 1).

Color: Chelicerae, dorsal surfaces bicolored, proximal three-quarters of manus dorsal surface sparsely infuscate, paler than carapace and densely infuscate distal quarter of manus dorsal surface and fingers. Carapace, tergites, metasoma, telson, pedipalp trochanter, femur, patella pro- and retrolateral surfaces, chela fingers, and legs dorsal and retrolateral surfaces entirely infuscate, uniformly dark brownish black; pedipalp patella ventral surface and chela manus dorsal, lateral, and ventral surfaces entirely infuscate but slightly paler, dark maroon. Coxosternal region, sternites, and legs, prolateral and ventral surfaces predominantly infuscate, paler, dark olive-brown with maxillary lobes darker, and leg tibia, basitarsus, and telotarsus paler than femur and patella. Genital opercula and pectines, immaculate, uniformly pale cream.

Chelicerae: Movable finger, prodistal (ventral) and retrodistal (dorsal) teeth unequal, retrodistal tooth considerably smaller than prodistal tooth, aligned longitudinally and not opposable. Fingers and manus, proventral surfaces, with long, dense vestiture of macrosetae. Coxae, prodorsal surfaces without stridulatory setae (scaphotrix); promedian surfaces without chemoreceptive lamelliform setae (trichocopae).

Carapace: Anterior width of posterior width, 72% (69%–73%, n = 4) (♂), 71% (66%–76%, n = 7) (♀); posterior width of length, 99% (97%–101%, n = 4) (♂), 98% (91%–104%, n = 7) (♀) (table 1). Three pairs of lateral ocelli, anterior pairs larger, facing anteriorly, separated by distance equal to width of ocellus from smaller posterior pair, facing posteriorly. Median ocelli slightly larger than anterior pairs of lateral ocelli, distance between ocelli equal to or greater than width of ocellus; median ocular tubercle situated posteromedially, distance from anterior carapace margin 54% (52%–56%, n = 4) (♂), 54% (51%–56%, n = 7) (♀) of carapace length (table 1). Anterior carapace margin with deep median notch (fig. 4A, C); with small median projection (epistome); without median depression; rostrolateral margin without distinct notch next to posterior lateral ocelli. Median longitudinal sulcus narrow, suturiform; continuous from median notch to interocular sulcus; with obsolete anterior furcation; without anterocular depression. Median ocular tubercle raised, superciliary carinae higher than ocelli, not extended anteriorly or posteriorly. Interocular sulcus present. Circumocular depressions completely encircling median ocular tubercle, converging anteriorly. Posteromedian and posteromarginal sulci deep. Paired mediolateral and posterolateral sulci shallow. Circumocular depressions completely encircling median ocular tubercle, converging anteriorly. Posteromedian and posteromarginal sulci deep. Paired mediolateral and posterolateral sulci shallow. Median longitudinal suture continuous from median notch to median ocular tubercle, equally strong along entire length; not extending to anterior margin of carapace, terminating at or posterior to median notch. Anterior furcated sutures obsolete. Anterocular furcated sutures absent. Interocular suture present, slender. Posterior sutures present but less distinct than median longitudinal and interocular sutures, converging on median ocular tubercle; connected anteriorly to posterior furcations of interocular suture and extending anteriorly beyond median ocular tubercle. Interocular and circumocular surfaces sparsely and finely granular anterior to median ocular tubercle, along median longitudinal sulcus, and often on frontal lobes (fig. 4A, B). Anterolateral, mediolateral and posterolateral surfaces uniformly finely granular, more coarsely and densely so in male. Posteromedian surfaces smooth.

Pedipalps: Femur width of length, 53% (52%–53%, n = 4) (♂), 57% (48%–62%, n = 7) (♀) (table 1). Retrodorsal carina distinct, comprising several spinform granules; more strongly developed than prodorsal carina. Dorsomedian carina vestigial, reduced to prominent granule demarcated by conspicuous macroseta. Dorsal secondary carina absent. Prodorsal carina obsolete, comprising isolated subspiniform granules. Promedian carina distinct, comprising row of spiniform granules (several demarcated by conspicuous macrosetae), oriented diagonally between prodorsal and proventral carinae. Proventral carina distinct, costate-granular, more strongly developed than retroventral carina. Ventromedian, secondary accessory, and retroventral carinae absent. Retromedian carinae absent, indicated only by macrosetal rows. Prodorsal intercarinal surfaces with scattered granules; other intercarinal surfaces smooth. Patella width of length, 45% (42%–47%, n = 4) (♂), 44% (42%–47%, n = 7) (♀) (table 1). Dorsal surface convex, dorsomedian carina dorsal to plane of retrodorsal carina, obsolete, costate (fig. 5A). Retrodorsal and retromedian carinae absent. Retroventral carina obsolete, costate (fig. 5B). Promedian carina obsolete, costate, demarcated by conspicuous macroseta; anterior process absent. Other carinae absent. Intercarinal surfaces smooth. Chela short, broad, base of fixed finger arising abruptly from manus (figs. 6A, 7); manus, height of width, 66% (59%–73%, n = 4) (♂), 61% (56%–63%, n = 7) (♀); length along ventroexternal carina of width, 61% (56%–63%, n = 4) (♂), 62% (58%–67%, n = 7) (♀); length along ventroexternal carina of length movable finger, 52% (50%–57%, n = 4) (♂), 57% (53%–60%, n = 7) (♀) (table 1). Dorsomedian carina obsolete, except proximal to base of fixed finger, distinct, costate. Dorsal secondary, subdigital, retromedian, retrolateral secondary and secondary accessory carinae absent. Digital carina absent, except proximal to base of fixed finger, obsolete costate. Retroventral and ventromedian carinae distinct, costate; retroventral carina more pronounced and projecting ventral to plane of ventromedian carina (fig. 6B). Proventral and promedian carinae obsolete, granular, each indicated by prominent macroseta. Prodorsal carina absent, but indicated by prominent macroseta. Chela moderately (♂) to sparsely (♀) setose on fingers and distally on manus. Manus, retrodorsal surface with (adult ♂) or without (♀, immature stages) marked concave depression at base of fixed finger (figs. 6A, 7), predominantly smooth proximally with shallow, anastomosing granules distally; retrolateral surfaces shallowly granular; ventral intercarinal surface smooth; prolateral intercarinal surfaces predominantly smooth, with scattered spiniform granules dorsally and distally. Fixed and movable fingers, prolateral intercarinal surfaces finely granular, retrolateral intercarinal surfaces predominantly smooth (♂) or finely granular (♀); median denticle rows each with six enlarged retrolateral denticles (including terminal denticle), proximal three retrolateral denticles on fixed finger, and proximal three (♀) or second and third most proximal (♂) retrolateral denticles on movable finger situated on lobes; fixed finger with proximal lobe, similar in size (♀, immatures) or slightly smaller than medial lobe (adult ♂); movable finger (adult ♂) without proximal lobe, with median lobe markedly more pronounced than other lobes, and with correspondingly well-developed notch in fixed finger; moderate gap proximally between fingers, when closed (adult ♂); terminal denticles of fingers interlocking unevenly when closed, movable finger displaced to exterior; distinct notch near tip of fixed finger to accommodate terminal denticle of movable finger.

Trichobothria: Neobothriotaxic major, Type C, with the following segment totals (n = 25; table 1): femur, 3 (1 d, 1 i, 1 e); patella, 52 (48–55): 2 d, 1 i, 34 (31–37) v, 14 (14 or 15) e, usually comprising 3 et, 2 est, 2 em, 2 esb, 5 eb; chela, 33 (31–35), manus, 21 (19–22), comprising 2 D, 10 E, 9 (7–10) V; fixed finger, 12 (11–13), comprising 4 d, 4 e, 4 (3–5) i (figs. 5–7). Total count of trichobothria per pedipalp: 87 (83–91). Femur, i situated on dorsal surface. Patella, d₂ situated on dorsomedian carina, slightly closer to d1 than to i. Chela, distance et–est ca. half distance est–esb; distance est–esb greater than half distance esb–eb; est aligned with dst.

Legs: First pair of maxillary lobes (coxapophyses) tapering anteriorly, longer than and encircling second pair. Stridulatory organs, comprising “rasp” (granular tubercles) and “scraper” (stridulatory setae or scaphotrix), present on opposing surfaces of coxae of pedipalps and first pair of legs, respectively. Legs acarinate. Femora, patellae, and tibiae, pro- and retrolateral surfaces each with scattered setiform macrosetae. Tibiae, prolateral surfaces, without spiniform macrosetae; I and II, retrolateral surfaces, usually each with two spiniform (t, st) macrosetae; III and IV, retrolateral surfaces, without spiniform macrosetae. Basitarsi I–IV, prolateral pedal spurs present (fig. 8); retrolateral pedal spurs absent; retrolateral margins similar, unmodified, rounded; I and II, dorsoventrally compressed, retrolateral margin produced into bladelike edge, III and IV terete, retrolateral margin unmodified, rounded. Basitarsi, pro- and retrolateral surfaces, each with microsetae, scattered long and short setiform macrosetae, and spiniform macrosetae, more numerous on I and II than III and IV. Basitarsi, spiniform macrosetae, I, retrolateral: t, st (polymorphic), sb; retroventral: t; proventral: t, st; II, retrolateral: t, st (polymorphic), sb; retroventral: t; proventral: t, st; III, retrolateral: t, sb; retroventral: t; proventral: t, st; prolateral: t, st; IV, retrolateral: t; retroventral: t; proventral: t, st; prolateral: t, st. Telotarsi I–IV short, stout, and distally broadened in dorsal and lateral views. Laterodistal lobes rounded. Dorsomedian lobes approximately equal to laterodistal lobes; each terminating in single setiform macroseta. Telotarsi, pro- and retrolateral surfaces, each with scattered microsetae and setiform macrosetae; I and II, proand retrolateral surfaces, in addition with two or more, dense brushlike rows of long to very long setiform macrosetae. Telotarsi each with pro- and retroventral rows of spiniform macrosetae, two of which are inserted on laterodistal lobes; macrosetal counts in pro- and retroventral rows equal on telotarsi I–IV, 3 and 4 (4 or 5) (n = 25), respectively (table 1). Telotarsal ungues short, curved; equal on telotarsi I and II, subequal on III and IV.

Sternum: Shape subpentagonal (fig. 4B, D). Median longitudinal sulcus shallow anteriorly, deep and narrow posteriorly.

Genital Operculum: Genital opercula suboval, completely divided longitudinally, partially overlapping, genital papillae present (♂, fig. 4B); subcordate, fused, genital papillae absent (♀, fig. 4D).

Hemispermatophore: Lamelliform, with complex, folded capsule and accessory distal lobe protruding between articular suture and distal lobe (hook). Distal lamina with distal crest truncate, unfolded.

Pectines: Distal edge reaching past distal edge of coxa IV but not reaching to distal edge of trochanter IV (♂, fig. 4B) or almost reaching distal edge of trochanter IV (♀, fig. 4D). First proximal median lamella (scape) of each pecten with mesial margin obtusely angular, greater than 90° but less than 180°, and devoid of teeth in proximal 19% (14%–23%, n = 4) (♂) or 24% (17%–31%, n = 7) (♀) of prolateral margin (table 1). Pectinal teeth straight and elongate (♂) or shorter and curved (♀); tooth count, 14/14 (13–15/14–15, n = 4) (♂), 13/13 (12–14/12–14, n = 7) (♀). Fulcra smooth proximally but densely setose (microsetae only) distally.

Mesosoma: Tergites each with shallow pair of submedian depressions and obsolete median carina. Pretergites smooth and glabrous. Posttergites smooth and glabrous anteromedially, densely and coarsely granular posteromedially and laterally (♂) or smooth and glabrous medially and anterolaterally, sparsely and coarsely granular posterolaterally (♀). Sternites IV–VI, each with paired longitudinal depressions prolateral to spiracles, absent on VII. Surface, sternites III–VII, smooth; VII, with paired, costate ventrosubmedian and ventrolateral carinae, without posteromarginal carina. Sternite VII, length of width, 52% (44%–56%, n = 4) (♂), 56% (47%–69%, n = 7) (♀) (table 1).

Metasoma and Telson: Metasomal segments I–V progressively increasing in length, decreasing in width; segment V, width of segment I width, 66% (64%–68%, n = 4) (♂), 66% (64%–68%, n = 7) (♀) (table 1). Metasoma robust; width of length, segment I, 93% (90%–95%, n = 4) (♂), 91% (89%–93%, n = 7) (♀); II, 78% (74%–81%, n = 4) (♂), 80% (76%–86%, n = 7) (♀); III, 68% (65%–73%, n = 4) (♂), 69% (60%–77%, n = 7) (♀); IV, 55% (52%–57%, n = 4) (♂), 58% (52%–65%, n = 7) (♀); V, 43% (39%–48%, n = 4) (♂), 43% (40%–45%, n = 7) (♀). Telson vesicle, width of metasomal segment V, width, 111% (105%–115%, n = 4) (♂), 97% (87%–107%, n = 7) (♀); enlarged (♂), globose, height of length, 62% (58%–66%, n = 4) (♂), 62% (57%–65%, n = 7) (♀); dorsal surface flat, ventral surface evenly curved. Aculeus relatively short, strongly curved, length of vesicle length, 32% (31%–34%, n = 4) (♂), 37% (33%–40%, n = 7) (♀). Length, metasoma and telson, of total length, 54% (52%–56%, n = 4) (♂), 51% (49%–53%, n = 7) (♀). Dorsosubmedian carinae, segments I–IV, distinct, complete, costategranular, posterior spiniform granules obsolete; V, absent (fig. 9A). Dorsolateral carinae, segments I–V, distinct, complete, costate-granular on I–IV, granular on V. Median lateral carinae, segment I, obsolete, granular, reduced to posterior two-thirds, II–V, absent, demarcated by macroseta at posterior margin on II–IV and near anterior margin on V (fig. 9B). Ventrolateral and ventrosubmedian carinae more strongly developed on segments I and II than on segments III and IV (fig. 9C). Ventrolateral carinae, segments I–IV, distinct, complete, costate on I–III, costate-granular on IV; V, distinct, complete, diverging posteriorly, comprising subspiniform granules, terminal granule similar in size to preceding granules. Ventrosubmedian carinae, segments I–IV, distinct, complete, costate; V, vestigial, each reduced to discontinuous row of spiniform granules, demarcated by macrosetae. Ventromedian carina, segment V, comprising single row of spiniform granules, unmodified posteriorly. Ventral surface, lateral aspect, segment IV, shallowly convex. Anal arch, segment V, dorsal carina, costate; ventral carina comprising subspiniform granules. Dorsal and lateral intercarinal surfaces, segments I–IV, finely and sparsely granular; V, smooth. Ventral intercarinal surfaces, segments I–IV, smooth; V with scattered granules. Telson vesicle, dorsal and lateral surfaces smooth; ventral surface with four obsolete longitudinal carinae, each comprising isolated spiniform granules, restricted to anterior half or third of vesicle.

Geographical Variation: All except three specimens examined were dark brownish black in color. One specimen with an indefinite locality in “East Africa” (BMNH) and two of the five male specimens labelled “Ugogo district, German East Africa” (BMNH 1914.7.3.3–7) were reddish brown in color, resembling P. militaris.

Ontogenetic Variation: Immature males and females closely resemble each other, and adult females, in general morphological features, but can be sexed by examination of the pectines and genital operculum. Males acquire secondary sexual characters in the final instar. In the specimens examined for this study, sexual maturity was determined by the presence of a marked concave depression in the retrodorsal surface of the pedipalp chela manus, at the base of the fixed finger in males (fig. 6A) and a fully developed ovariuterus or the gravid condition in females.

Sexual Dimorphism: The characters of primary external sexual dimorphism are the undivided genital operculum, which opens in a single flap in the female (fig. 4D), compared with the two unconnected sclerites, which open independently and cover a pair of genital papillae in the male (fig. 4B). Additionally, males possess larger pectines (fig. 4B, D) with a greater number of pectinal teeth (table 1). The most obvious secondary sexual characters observed in adult males, compared with adult females and juveniles of both sexes, concern the modifications of the pedipalp chela: a marked concave depression is present in the retrodorsal surface of the manus, at the base of the fixed finger; the proximal lobe of the movable finger is absent; the median lobe of the movable finger is markedly more pronounced than the other lobes; and a moderate gap is present proximally between the fingers, when closed (fig. 6A).

Distribution: Pandinoides cavimanus appears to be endemic to Tanzania (fig. 2). Records from Kenya are referable to P. duffmackayi, sp. nov., or P. militaris, those from Ethiopia or Somalia, including the (immature) female on the cover of Kovařík (2009), to P. militaris. Records of P. cavimanus from northeastern Tanzania, including Kilimanjaro (Pocock, 1888), Arusha (Werner, 1936), and Naberera (Kovařík, 2009) are referable to P. duffmackayi, sp. nov. Most of the known locality records of P. cavimanus occur in the central part of Tanzania, in the Dodoma, Iringa, Shinyanga, and Singida provinces. Records from the eastern shore of Lake Tanganyika and Bukoba on the western shore of Lake Victoria/Nyanza (Kraepelin, 1913; Birula, 1915b) await confirmation. This species does not occur in the northeast of the Democratic Republic of Congo (formerly Zaïre), as stated by Kovařík (1998, 2009) and Fet (2000), nor in Mozambique, as stated by Kovařík (1998).

Ecology: The known locality records of P. cavimanus occur in arid savannah, dominated by Acacia Mill. and Commiphora Jacq. at 1010–1120 m elevation. Most specimens, for which data are available, were excavated from burrows during the day. Pandinoides cavimanus is fossorial, constructing burrows in compacted clayey to sandyloam soils. It is allopatric with P. militaris and parapatric with P. duffmackayi, sp. nov., and Pandinurus viatoris (Pocock, 1890), but sympatric with another scorpionid, Opistophthalmus boehmi (Kraepelin, 1896).

Conservation: This species is protected in the Ruaha National Park.

Remarks: Pocock (1888: 249) noted that: “The specimen from Kilima-Njaro is smaller and slightly less granular than the [type] from Umyamuezi.” The smaller specimen is conspecific with P. duffmackayi.

In discussing the differences in coloration between P. exitialis and P. gregoryi, which Kovařík (2003) mistakenly synonymized with the former, Kovařík (2009 : 55)remarked that:

Whereas it is indeed true that color may vary both within and among conspecific populations of some scorpion species, e.g. Hadrurus arizonensis Ewing, 1928, each case must be assessed on its own merits. Variation in color pattern (rather than color itself) is often both consistent and congruent with variation in other morphological (and indeed, genetic) characters, as described by Kovařík (2009) in the closely related P. exitialis and P. gregoryi, and the three closely related species of Pandinoides covered in the present contribution. Valid taxa, e.g., P. gregoryi and P. militaris, may be mistakenly synonymized when differences in coloration are dismissed out of hand without detailed investigation of other characters.

Additional Material Examined: 1 ♂ (NM 18769). East Africa, 1 ♂ (BMNH). TANZANIA: Imported for pet trade: 2 ♂ (AMNH), 1 ♀ (AMCC [LP 3250]), 1 ♀ (AMNH), [leg] (AMCC [LP 3259]), ix.2002, ex R. MacInnes, 1 subad. ♀ (AMNH), [leg] (AMCC [LP 2161]), vii.2007, ex J. Huff, 1 ♂ (AMCC [LP 7084]), iii.2008, ex A. Tietz, 1 subad. ♀ (AMNH), [pedipalp] (AMCC [LP 8271]). Ugogo district, German East Africa, Dr. E.J. Baxter, 5 ♂ (BMNH 1914.7.3.3-7), 1 ♀, 1 subad. ♂ (BMNH 1914.7.3.8–9). Dodoma Prov.: Dodoma Distr.: Dodoma [06°10′S 035°45′E], G.E.A., 8.xii.1918, A. Loveridge, 2 ♂(MCZ 15519), 1 subad. ♀, 4 juv. (MCZ 15513), 1 ♂ (BMNH 1923.11.8.9); Dodoma, brought in by dealer, 1 ♂, 1 juv. ♂ (ZMUC); Mapinduzi Village, Dodoma region [06°11′S 035°45′E], ix.2006, J. Beraducci, found in ground, 3 ♂, 3 ♀, 2 subad. ♂, 1 subad. ♀ (AMNH), 1 juv. ♀ (AMCC [LP 8918]), x.2006, J. Beraducci, 1 ♂, 3 ♀, 1 subad. ♂ (AMNH). Kongwa Distr.: Kongwa [06°12′S 036°25′E], 21.iv.1918, A. Loveridge, 1 ♀ (MCZ 15522). Mpwapwa Distr.: Mpapua [Mpwapwa, 06°21′S 036°29′E], 1899, 1 ♂ (SAM 5111 ex ZMH). Iringa Prov.: Iringa Distr.: Ruaka [Ruaha] National Park [07°40′S 034°52′E], 1979, K.M. Howell, 1 ♂ (MRAC 159.373). Singida Prov.: Manyoni Distr.: Ikikuyu [Kikuyu, 05°52′S 035°04′E], 12.ii.1923, A. Loveridge, 1 ♂, 1 subad. ♂ (MCZ 15514), 1 ♂, 1 juv. ♂ (MCZ 15518), 1.xi.1923, A. Loveridge, 1 ♂ (MCZ 15521), 12.xi.1923, A. Loveridge, 1 juv. ♂ (MCZ 15516); Kikuyu, Dodoma, 21.xii.1929, A. Loveridge, 2 ♂, 1 ♀ (MCZ 15515).

Erroneous Records: Florida, U.S.A., New Bold's 5–10, Clubmont (introduction), vii.1965, University of Florida Collections, 1 ♂ (FSCA). Stanley Falls [Tshopo Prov., Democratic Republic of Congo], 00°30′N 025°12′E, 1 ♂ (MRAC 4669). Based on the known localities, the record from Stanley Falls is assumed to be a labeling error.

Pandinoides duffmackayi, sp. nov. Figures 1C, 2, 10–16; tables 2, 3

Holotype: ♂ (AMNH), TANZANIA: Arusha Prov.: Arumeru Distr.: Majengo Village, Kikatitti [03°23′S 036°57′E], near Kilimanjaro International Airport, ix.2006, J. Beraducci, found in holes in the earth.

Paratypes: KENYA: Eastern Prov.: Machakos Distr.: Athi yu Mawe [01°27′S 036°59′E], C.S. Betton, 1 ♂ (BMNH 1899.7.31.1); Nairobi, Athi River, 01°27′S 036°59′E, xi.1945, 1 ♀ (NMK 231 old 6). Rift Valley Prov.: Kajiado Distr.: near Hunters Lodge [02°12′S 037°40′E], 1977, Mrs. Fordyce, 1 ♀ (NMK 234 old 107); Kajiado town, 01°51′S 036°47′E, J.P.E.C. Darlington, 1 ♀, 2 1st instars (NMK 557); Kiseria Ngong [Kiserian, 01°24′S 036°49′E], 15.iii.1983, J. Sylvester, 1 ♀ (NMK 230); Namanga, 13 mi. N, 02°24′S 036°50′E, 4350 ft, 19.ix.1976, A. Duff-Mackay, dug from hole 10 × 25.2 mm in red sand, dry Acacia/Commiphora, 1 ♀ (NMK 232 old 75), dug from hole 14.6 × 31.3 mm in red sand, dry Acacia/Commiphora, 1 ♂ (NMK 233 old 76); Ngong hills, NW of, 01°21′S 036°35′E, 5800 ft, 17.x.1976, A. Duff-Mackay, dug out of multiple-entrance burrow, lava rocks, Acacia, lileshwa [Tarchonanthus camphoratus L.], 1 ♀ (NMK 235 old 91), 2 ♂, 1 ♀ (NMK 236 old 92), dug from burrow, lava rocks, Acacia, lileshwa, 1 ♀, 3 juv. ♂, 3 juv. ♀ (NMK 237 old 93), 1 subad. ♂ (NMK 238 old 90); Ngong hills, SW of, 01°23.5′S 036°35′E, 8.ix.1982, R. Drewes and A. Duff-Mackay, dug from nest 225 mm underground, 80 mm diam. by 25 mm height, length large burrow 535 mm, 3 small holes and 1 large, new diggings outside and inside blocked most of way, food: segments of millipede, open ground near brush, sparse grass heavily grazed, 2 juv. ♂, 6 juv. ♀ (NMK 239 old 289). TANZANIA: Imported for pet trade: 3 ♂ (AMNH), [leg] (AMCC [LP 3258]), 1 ♀ (AMNH), 1 ♀ (AMCC [LP 3249]), xi.1997, ex F. Somma, 1 ♂, 1 juv. ♀ (AMCC [LP 1600]), 3.x.2007, ex A. Tietz, 1 juv. ♂ (AMCC [LP 7294]). Arusha Prov.: Arumeru Distr.: Kikatitti [03°23′S 036°57′E], near Kilimanjaro International Airport, x.2006, J. Beraducci, 2 ♂, 1 subad. ♂, 5 juv. ♀ (AMNH), 1 juv. ♀ (AMCC [LP 8335]); Majengo Village, Kikatitti, near Kilimanjaro International Airport [03°23′S 036°57′E], ix.2006, J. Beraducci, found in holes in the earth, 2 ♂, 3 ♀, 1 juv. ♂, 3 juv. ♀ (AMNH). Kilimanjaro Prov.: Kilimanjaro [03°04′S 037°22′E], M.J. Jackson, 1 ♂ (BMNH 1887.147).

Etymology: The specific epithet is a patronym in honor of Alexander Duff-Mackay (1939–2003), a herpetologist described as “one of the most knowledgeable biologists in East Africa” (Schiøtz, 2003: 1) who worked at the National Museums of Kenya, Nairobi from 1964 to 1995. Duff-Mackay was very interested in scorpions (Spawls et al., 2004: 7) and, using ultraviolet light detection and traditional methods, collected many large series in Kenya, including most of the type series of the new species, recording detailed notes about their habitat and ecology that have greatly aided our understanding of the taxonomy, distribution, and ecology of the scorpions of East Africa.

Diagnosis: Pandinoides duffmackayi, sp. nov., may be separated from P. cavimanus and P. militaris, as follows. Pandinoides duffmackayi, sp. nov., is reddish brown to reddish black in color (fig. 1C), whereas P. cavimanus is usually brownish black (fig. 1A, B), rarely reddish brown, and P. militaris, yellowish brown (fig. 1D) to reddish brown. Pandinoides duffmackayi, sp. nov., is smaller, with total adult body length 60–78 mm, carapace length 10–13 mm and chela length 14–19 mm (tables 2, 3), than P. cavimanus, with total adult body length 87–111 mm, carapace length 15–18 mm and pedipalp chela length 21–29 mm (table 1), and P. militaris, with total adult body length 95–123 mm, carapace length 17–19 mm, and chela length 25–35 mm (tables 4, 5). The ventral surface of the telson vesicle of the male is smooth or nearly so in P. duffmackayi, sp. nov. (fig. 16B, C), but bears obsolete carinae, each comprising isolated spiniform granules, restricted to its anterior half or third, in P. cavimanus (fig. 9B, C), and distinct carinae, each comprising prominent spiniform granules, extending its entire length, in P. militaris (fig. 23B, C). Pandinoides duffmackayi, sp. nov., bears 74–82 trichobothria on the pedipalp, with 39–45 trichobothria on the patella, including 22–28 in the v series, whereas P. cavimanus bears 83–91 trichobothria on the pedipalp, with 48–55 trichobothria on the patella, including 31–37 in the v series, and P. militaris bears 84–93 trichobothria on the pedipalp, with 47–55 trichobothria on the patella, including 30–38 in the v series. The proximal lobe on the pedipalp chela fixed finger of the adult male is similar in size to the medial lobe, creating little to no gap proximally between the fixed and movable fingers, when closed in P. duffmackayi, sp. nov. (fig. 13A), whereas the proximal lobe is slightly smaller than the medial lobe, creating a moderate gap proximally between the fingers, when closed in P. cavimanus (fig. 6A), and it is vestigial, much smaller than the medial lobe, or absent, creating a prominent gap proximally between the fingers, when closed, in P. militaris (fig. 20A). The pectinal tooth count of P. duffmackayi, sp. nov., with 11–13, usually 12 (♂) and 10–12, usually 11 (♀), is lower than that of P. cavimanus, with 14 or 15, usually 14 (♂), and 13 or 14, usually 13 (♀), and P. militaris, with 14–16, usually 14 (♂), and 13–15, usually 14 (♀). The prolateral t and st macrosetae are setiform on basitarsi III, and t setiform and st spiniform on IV, in P. duffmackayi, sp. nov., whereas t and st are spiniform on II and IV in P. cavimanus and P. militaris.

Based on unpublished genetic data, P. duffmackayi, sp. nov., is most closely related to P. cavimanus, from which it may be further separated as follows. The interocular and circumocular surfaces of the carapace are smooth, or nearly so, in P. duffmackayi, sp. nov. (fig. 11A, C), but sparsely and finely granular anterior to the median ocular tubercle, along the median longitudinal sulcus, and often on the frontal lobes in P. cavimanus (fig. 4A, C). The metasoma of P. duffmackayi, sp. nov., is 49%–53% (♂) and 46%–49% (♀) of total body length, the summed lengths of segments IV and V, 105%–116% (♂) and 91%–110% (♀) of carapace length (tables 2, 3), whereas the metasoma of P. cavimanus is 52%–56% (♂) and 49%–53% (♀) of total body length, the summed lengths of segments IV and V, 115%–130% (♂) and 107%–120% (♀) of carapace length (table 1). The dorsal surfaces of metasomal segments I–IV in the male are smooth in P. duffmackayi, sp. nov. (fig. 16A), but finely and sparsely granular in P. cavimanus (fig. 9A).

Pandinoides duffmackayi, sp. nov., may be further separated from P. militaris as follows. The carapace is less compressed, dorsoventrally, in P. duffmackayi, sp. nov. (fig. 11A, C), than in P. militaris (fig. 18A, C). The median ocular tubercle is posteromedial, its distance from the anterior carapace margin 51%–56% of the carapace length in P. duffmackayi, sp. nov. (tables 2, 3), but medial, its distance from the anterior carapace margin 46%–52% of the carapace length in P. militaris (tables 4, 5). The ventrosubmedian carinae on metasomal segment IV are distinct, costate in P. duffmackayi, sp. nov. (fig. 16C), but obsolete, granular in P. militaris (fig. 23C). The chela manus of P. duffmackayi, sp. nov., is broader, the length along the retroventral carina 56%–70% (♂) and 57%–70% (♀) of the width, and more convex, the height 58%–77% (♂) and 59%–94% (♀) of the width, than the manus of P. militaris, in which the length along the retroventral carina is 64%–76% (♂) and 62%–72% (♀) of the width, and the height 49%–68% (♂) and 64%–73% (♀) of the width. The chela fixed finger of P. duffmackayi, sp. nov., arises abruptly from the manus and is wider proximally (figs. 13A, 14) than that of P. militaris, which arises more gradually from the manus (figs. 20A, 21). The chela fingers and distal surfaces of the manus are moderately (♂) to sparsely (♀) setose in P. duffmackayi, sp. nov. (fig. 1C), but densely (♂) to moderately (♀) setose in P. militaris (fig. 1D). Pandinoides duffmackayi, sp. nov., bears 31–34 trichobothria on the pedipalp chela, including 7–10 in the V series, whereas P. militaris bears 33–36 trichobothria on the chela, including 9–12 in the V series. The angle of the first proximal median lamella (scape) of the pecten is smaller (more acute), especially in the female, of P. duffmackayi, sp. nov. (fig. 11B, D), compared with P. militaris (fig. 18B, D).

Description: The following account is based on the type material. Only characters that differ from P. cavimanus are described.

Total Length: Adult medium, maximum length, measured from anterior margin of carapace to tip of aculeus, 70 mm (60–75 mm, n = 8) (♂), 69 mm (61–78 mm, n = 9) (♀) (tables 2, 3).

Color: Chelicerae, dorsal surfaces bicolored, proximal three-quarters of manus dorsal surface sparsely infuscate, paler than carapace and densely infuscate distal quarter of manus dorsal surface and fingers. Tergites, sternite VII, metasoma, telson and pedipalp chela fingers densely infuscate, brownish black, telson similar to metasoma; carapace lateral and posterior surfaces, pedipalp trochanter, femur, patella proand retrolateral surfaces infuscate but paler, dark reddish brown; carapace interocular and circumocular surfaces, pedipalp patella ventral surface and chela manus dorsal, lateral and ventral surfaces immaculate, markedly paler, light reddish brown. Coxosternal region, sternites III–VI and legs faintly infuscate, yellowish brown, with maxillary lobes darker. Genital opercula and pectines immaculate, uniformly pale cream.

Carapace: As for P. cavimanus, except as follows. Anterior width of posterior width, 74% (68%–81%, n = 8) (♂), 70% (65%–76%, n = 9) (♀); posterior width of length, 92% (83%–96%, n = 8) (♂), 96% (90%–106%, n = 9) (♀) (tables 2, 3). Median ocular tubercle situated posteromedially, distance from anterior carapace margin 54% (51%–56%, n = 8) (♂), 53% (51%–55%, n = 9) (♀) of carapace length (tables 2, 3). Interocular, circumocular, and posteromedian surfaces smooth or nearly so (fig. 11A, C). Anterolateral and mediolateral surfaces sparsely and coarsely granular; posterolateral surfaces sparsely and finely granular.

Pedipalps: As for P. cavimanus, except as follows. Femur width of length, 57% (53%–66%, n = 8) (♂), 55% (50%–66%, n = 9) (♀) (tables 2, 3). Retrodorsal carina obsolete, granular; more strongly developed than prodorsal carina. Prodorsal carina obsolete, comprising few rounded granules. Promedian carina distinct, comprising row of spiniform granules (several demarcated by conspicuous macrosetae), oriented diagonally between prodorsal and proventral carinae. Patella width of length, 45% (40%–58%, n = 8) (♂), 44% (37%–54%, n = 9) (♀) (tables 2, 3). Chela short, broad, base of fixed finger arising abruptly from manus (figs. 13A, 14); manus, height of width, 71% (58%–77%, n = 8) (♂), 75% (59%–94%, n = 9) (♀); length along ventroexternal carina of width, 61% (56%–70%, n = 8) (♂), 62% (57%–70%, n = 9) (♀); length along ventroexternal carina of length movable finger, 56% (48%–64%, n = 8) (♂), 54% (48%–60%, n = 9) (♀) (tables 2, 3). Dorsomedian carina obsolete, except proximal to base of fixed finger, distinct, costate. Digital carina absent, except proximal to base of fixed finger, obsolete, costate-granular. Prodorsal, promedian and proventral carinae absent, each indicated by prominent macroseta. Chela moderately (♂) to sparsely (♀) setose on fingers and distally on manus. Manus, retrodorsal surface predominantly reticulate proximally with shallow, anastomosing granules distally; retrolateral surfaces shallowly granular; ventral intercarinal surface smooth; prolateral surfaces predominantly smooth, except for few scattered spiniform granules distally. Fixed and movable fingers, pro- and retrolateral intercarinal surfaces finely and sparsely granular (♂) to smooth (♀); median denticle rows each with six enlarged retrolateral denticles (including terminal denticle), proximal three retrolateral denticles on fixed and movable fingers situated on lobes; fixed finger with proximal lobe similar in size to medial lobe; movable finger (adult ♂) with proximal lobe, with median lobe slightly more pronounced than other lobes, and with correspondingly well-developed notch in fixed finger; little to no gap proximally between fingers, when closed (adult ♂).

Trichobothria: As for P. cavimanus, except as follows. Neobothriotaxic major, Type C, with the following segment totals (n = 34; tables 2, 3): femur, 3 (1 d, 1 i, 1 e); patella, 43 (39–45): 2 d, 1 i, 25 (22–28) v, 14 (14 or 15) e, usually comprising 3 et, 2 est, 2 em, 2 esb, 5 eb; chela, 32 (31–34), manus, 20 (19–22), comprising 2 D, 10 E, 8 (7–10) V; fixed finger, 12 (12–13), comprising 4 d, 4 e, 4 (4 or 5) i (figs. 12–14). Total count of trichobothria per pedipalp: 78 (74–82). Chela, distance et–est half to greater than half distance est–esb; est proximal to or aligned with dst.

Legs: As for P. cavimanus, except as follows. Basitarsi, spiniform macrosetae, I, retrolateral: t, sb; retroventral: t; proventral: t, st; II, retrolateral: t, sb; retroventral: t; proventral: t, st; III, retrolateral: t, sb; retroventral: t; proventral: t, st; IV, retrolateral: t; retroventral: t; proventral: t, st; prolateral: st (fig. 15). Telotarsi, macrosetal counts in pro- and retroventral rows equal on I–IV, 3 (2 or 3) and 4 (3 or 4) (n = 34), respectively (tables 2, 3).

Pectines: As for P. cavimanus, except as follows. Distal edge reaching past distal edge of coxa IV but not reaching to distal edge of trochanter IV (♂, fig. 11B) or not reaching to distal edge of trochanter IV (♀, fig. 11D). First proximal median lamella (scape) of each pecten with mesial margin obtusely angular, greater than 90° but less than 180°, and devoid of teeth in proximal 19% (12%–24%, n = 8) (♂) or 30% (22%–35%, n = 9) (♀) of prolateral margin (tables 2, 3). Pectinal tooth count, 12/13 (11–13/12–13, n = 8) (♂), 11/11 (10–12/10–12, n = 9) (♀).

Mesosoma: As for P. cavimanus, except as follows. Posttergites smooth and glabrous medially and anterolaterally, sparsely (♂) to very sparsely (♀), and finely granular posterolaterally. Sternite VII, length of width, 60% (47%–70%, n = 8) (♂), 59% (46%–87%, n = 9) (♀) (tables 2, 3).

Metasoma and Telson: As for P. cavimanus, except as follows. Metasomal segments I–V progressively increasing in length, decreasing in width; segment V, width of segment I width, 66% (59%–71%, n = 8) (♂), 66% (58%–73%, n = 9) (♀) (tables 2, 3). Metasoma robust; width of length, segment I, 107% (95%–126%, n = 8) (♂), 111% (101%–130%, n = 9) (♀); II, 86% (80%–95%, n = 8) (♂), 95% (85%–105%, n = 9) (♀); III, 78% (72%–88%, n = 8) (♂), 80% (72%–88%, n = 9) (♀); IV, 65% (61%–69%, n = 8) (♂), 68% (59%–77%, n = 9) (♀); V, 48% (45%–51%, n = 8) (♂), 52% (43%–68%, n = 9) (♀). Telson vesicle, width of metasomal segment V, width, 107% (95%–118%, n = 8) (♂), 94% (80%–107%, n = 9) (♀); enlarged (♂), globose, height of length, 64% (55%–72%, n = 8) (♂), 66% (54%–86%, n = 9) (♀); dorsal surface flat, ventral surface evenly curved. Aculeus relatively short, strongly curved, length of vesicle length, 35% (30%–38%, n = 8) (♂), 38% (35%–41%, n = 9) (♀). Length, metasoma and telson, of total length, 50% (49%–53%, n = 8) (♂), 47% (46%–49%, n = 9) (♀). Median lateral carinae, segments I–III, obsolete, granular, reduced to posterior half of segment, IV and V, absent demarcated only by macroseta at posterior margin on IV and near anterior margin on V (fig. 16B). Ventrosubmedian carinae, segments I–IV, complete, costate, distinct on I and II, obsolete on III and IV; V, vestigial, each reduced to discontinuous row of spiniform granules, demarcated by macrosetae (fig. 16C). Dorsal intercarinal surfaces, segments I–V, smooth (fig. 16A). Lateral intercarinal surfaces, segments I–V, sparsely granular (♂) to smooth (♀). Telson vesicle, dorsal and lateral surfaces smooth; ventral surface smooth or nearly so.

Geographical Variation: No noticeable variation.

Ontogenetic Variation: As for P. cavimanus, except that immature stages are often markedly more infuscate than adults, in some cases entirely so.

Sexual Dimorphism: As for P. cavimanus, except as follows. Pandinoides duffmackayi, sp. nov., is the least sexually dimorphic of the three species, the adult male differing from the adult female as follows. The concave depression in the retrodorsal surface of the pedipalp chela manus, at the base of the fixed finger, is shallower than in the other species. The proximal lobe of the movable finger is present and the median lobe of the movable finger only slightly more pronounced than the other lobes, such that little to no gap is present proximally between the fixed and movable fingers, when closed (fig. 13A).

Distribution: Pandinoides duffmackayi, sp. nov., is endemic to Kenya and Tanzania (fig. 2). The known locality records fall within a fairly small area in southwestern Kenya (Eastern and Rift Valley provinces) and northeastern Tanzania (Arusha and Kilimanjaro provinces). Records attributed to P. cavimanus from this area, including Kilimanjaro (Pocock, 1888), Arusha (Werner, 1936), and Naberera (Kovařík, 2009) are referable to P. duffmackayi, sp. nov.

Ecology: The known locality records of P. duffmackayi, sp. nov., occur in arid savannah, dominated by Acacia and Commiphora or Leleshwa (Tarchonanthus camphoratus), with sparse grass at 1325–1770 m elevation. Most specimens for which data are available were excavated from burrows during the day. Pandinoides duffmackayi, sp. nov., constructs burrows up to 55 cm long and 25 cm deep in compacted clayey soil with pumice to consolidated red sandy loam. This species is parapatric with the other two congeners. It may be sympatric with Pandinurus exitialis (Pocock, 1888) in the Namanga area of Kenya.

Conservation: This species appears to be protected in the Ngong Hills Nature Reserve near Nairobi, Kenya.

Remarks: Pocock (1888: 249) first noted differences in size and granulation between an individual of this species and the holotype of P. cavimanus when stating:

However, with only two specimens available for study, neither in good condition, Pocock (1888) was apparently unable to appreciate the significance of the variation.

Erroneous Record: Port Herald [Nsanje], Nyasaland [Malawi], Dr. J.E.S. Old, 1 ♂ (BMNH 1914.4.7.1). Based on the known localities of the other material examined, this specimen is probably mislabeled.

Pandinoides militaris Pocock, 1900, stat. rev. Figures 1D, 2, 17–23; tables 4, 5

FIGURE 17.

Pandinoides militaris (Pocock, 1900), habitus, dorsal (A, C) and ventral (B, D) aspects. A, B. ♂ (AMNH), C, D. ♀ (AMNH), Kishushe, Kenya. Scale bars = 10 mm.

FIGURE 18.

Pandinoides militaris (Pocock, 1900), carapace, dorsal aspect (A, C), and sternum, genital opercula, and pectines, ventral aspect (B, D). A, B. ♂ (AMNH), C, D. ♀ (AMNH), Kishushe, Kenya. Scale bars = 5 mm.

FIGURE 19.

Pandinoides militaris (Pocock, 1900), ♂ (AMNH), Kishushe, Kenya, dextral pedipalp patella, retrolateral (A) and ventral (B) aspects. Scale bar = 5 mm.

FIGURE 20.

Pandinoides militaris (Pocock, 1900), ♂ (AMNH), Kishushe, Kenya, dextral pedipalp chela, retrolateral (A), ventral (B), and prolateral (C) aspects. Scale bar = 5 mm.

FIGURE 21.

Pandinoides militaris (Pocock, 1900), ♀ (AMNH), Kishushe, Kenya, dextral pedipalp chela, retrolateral aspect. Scale bar = 5 mm.

FIGURE 22.

Pandinoides militaris (Pocock, 1900), ♂ (AMNH), Kishushe, Kenya, legs I–IV, tibiae, basitarsi and telotarsi, ventral aspect (A–D). Scale bar = 2.5 mm.

FIGURE 23.

Pandinoides militaris (Pocock, 1900), ♂ (AMNH), Kishushe, Kenya, metasoma and telson, dorsal (A), lateral (B), and ventral (C) aspects. Scale bar = 5 mm.

Type Material: Holotype subad. ♀ (BMNH 1897.11.10.4–5) [examined], ETHIOPIA: Oromia Regional State: Borena Administrative Zone: El Dere [03°52′N 039°46′E] near Aimola [ca. 04°05′30″N 040°21′15″E, Mandera Distr., North Eastern Prov., Kenya], Boran country, Somaliland, 23.iii.1895, A. Donaldson Smith.

In the original description of P. militaris, Pocock (1900b: 61) listed two specimens from the BMNH: “Aimola in the Boran Country (Donaldson Smith); also Ndi, on the Weiss Road inland from Mombasa (C. Steuart Betton).” Pocock (1897b: 397) had previously mentioned “three female examples … of which … only one, the smallest, is labeled with an exact locality… near Aimola in the Boran country, 3,000 ft. 23.3.95” and prior to that “two mutilated males obtained by Dr. Gregory at Kinani and a place 4 miles to the south of it,” which Pocock (1896: 431) determined to be P. cavimanus. All these specimens were examined for the present study and determined to be conspecific. In spite of the availability of two adult males from Kinani, two adult females from Somaliland with no precise locality data, a subadult female from Aimola, and a juvenile male from Ndi, Pocock (1896, 1897b, 1900b: 61) based the description of P. militaris solely on the subadult female from Aimola (actually, El Dere), specifically stating the measurements were “of [the] type.” No mention was made of more than one type. Therefore, the specimen from Aimola is the holotype of P. militaris, the specimen from Ndi was mistakenly listed as a paratype (Fet, 2000: 468; Kovařík, 2003: 149), and the subsequent designation of the specimen from Aimola as lectotype (Kovařík, 2009: 50; also see Rossi, 2015a: 50, fig. 11) was unjustified.

Based on the original label, the type locality is “El Dere” not “Aimola” and, according to the maps of A. Donaldson Smith's expedition route (Donaldson Smith, 1896: 121; 1897, map sheet 3), El Dere is in southern Ethiopia whereas Aimola, to the northeast, is in northeastern Kenya, near the Ethiopian border, not in Somalia as generally assumed (e.g. Fet, 2000; Kovařík, 2003, 2009).

Diagnosis: Pandinoides militaris may be separated from P. cavimanus and P. duffmackayi, sp. nov., by the following combination of characters. Pandinoides militaris is yellowish brown (fig. 1D) to reddish brown in color, whereas P. cavimanus is usually brownish black (fig. 1A, B), rarely reddish brown, and P. duffmackayi, sp. nov., reddish brown to reddish black (fig. 1C). The carapace is more compressed, dorsoventrally, in P. militaris (fig. 18A, C) than in P. cavimanus (fig. 4A, C) and P. duffmackayi, sp. nov. (fig. 11A, C). The median ocular tubercle is medial, its distance from the anterior carapace margin 46%–52% of the carapace length in P. militaris (tables 4, 5), but posteromedial, its distance from the anterior carapace margin 51%–56% of the carapace length in P. duffmackayi, sp. nov. (tables 2, 3), and P. cavimanus (table 1). The ventrosubmedian carinae on metasomal segment IV are distinct, costate in P. cavimanus (fig. 9C) and P. duffmackayi, sp. nov. (fig. 16C), but obsolete, granular in P. militaris (fig. 23C). The ventral surface of the telson vesicle of the male bears distinct carinae, each comprising prominent spiniform granules, extending its entire length in P. militaris (fig. 23B, C), whereas the vesicle bears obsolete carinae, each comprising isolated spiniform granules, restricted to its anterior half or third in P. cavimanus (fig. 9B, C), and is smooth or nearly so in P. duffmackayi, sp. nov. (fig. 16B, C). The chela manus of P. militaris is narrower, the length along the retroventral carina 64%–76% (♂) and 62%–72% (♀) of the width, and less convex, the height 49%–68% (♂) and 64%–73% (♀) of the width, than the manus of P. cavimanus, in which the length along the retroventral carina is 56%–63% (♂) and 58%–67% (♀) of the width, and the height 59%–73% (♂) and 66%–76% (♀) of the width, and that of P. duffmackayi, sp. nov., in which the length along the retroventral carina is 56%–70% (♂) and 57%–70% (♀) of the width, and the height 58%–77% (♂) and 59%–94% (♀) of the width. The chela fixed finger of P. militaris arises more gradually from the manus, and is narrower proximally (figs. 20A, 21), than the fingers of P. cavimanus (figs. 6A, 7) and P. duffmackayi, sp. nov. (figs. 13A, 14), which arise abruptly from the manus. The chela fingers and distal surfaces of the manus are densely (♂) to moderately (♀) setose in P. militaris (fig. 1D), but moderately (♂) to sparsely (♀) setose in P. cavimanus (fig. 1B) and P. duffmackayi, sp. nov. (fig. 1C). The proximal lobe on the pedipalp chela fixed finger of the adult male is vestigial, much smaller than the medial lobe, or absent, creating a prominent gap proximally between the fixed and movable fingers, when closed in P. militaris (fig. 20A), whereas the proximal lobe is similar in size to the medial lobe, creating little to no gap proximally between the fingers, when closed in P. duffmackayi, sp. nov. (fig. 13A), and it is slightly smaller than the medial lobe, creating a moderate gap proximally between the fingers, when closed in P. cavimanus (fig. 6A). Pandinoides militaris bears 33–36 trichobothria on the pedipalp chela, including 9–12 in the V series, whereas P. cavimanus and P. duffmackayi, sp. nov., bear 31–35 and 31–34 trichobothria on the chela, respectively, including 7–10 in the V series. The angle of the first proximal median lamella (scape) of the pecten is greater (more obtuse), especially in the female, of P. militaris (fig. 18B, D), compared with P. cavimanus (fig. 4B, D) and P. duffmackayi, sp. nov. (fig. 11B, D).

Pandinoides militaris has often been confused with P. cavimanus due to their similar size and high trichobothrial counts, but may be separated from the latter as follows. The circumocular surfaces of the carapace are smooth or with a few granules anterior to the ocular tubercle in P. militaris (fig. 18A, C), but sparsely and finely granular anterior to the median ocular tubercle, along the median longitudinal sulcus, and often on the frontal lobes in P. cavimanus (fig. 4A, C). The metasoma of P. militaris is 46%–52% (♂), 42%–50% (♀) of total body length, the summed lengths of segments IV and V, 98%–117% (♂), 87%–105% (♀) of carapace length, whereas the metasoma of P. cavimanus is 52%–56% (♂) and 49%–53% (♀) of total body length, the summed lengths of segments IV and V, 115%–130% (♂), 107%–120% (♀) of carapace length. The dorsal surfaces of metasomal segments I–IV in the male are smooth in P. militaris (fig. 23A), but finely and sparsely granular in P. cavimanus (fig. 9A).

Pandinoides militaris may be further separated from P. duffmackayi, sp. nov., as follows. Pandinoides militaris is larger, with total adult body length 95–123 mm, carapace length 17–19 mm and pedipalp chela length 25–35 mm (tables 4, 5), than P. duffmackayi, sp. nov., with total adult body length 60–78 mm, carapace length 10–13 mm and chela length 14–19 mm (tables 2, 3). Pandinoides militaris bears 84–93 trichobothria on the pedipalp, with 47–55 trichobothria on the patella, including 30–38 in the v series, whereas P. duffmackayi, sp. nov., bears 74–82 trichobothria on the pedipalp, with 39–45 trichobothria on the patella, including 22–28 in the v series. The pectinal tooth count of P. militaris, with 14–16, usually 14 (♂), and 13–15, usually 14 (♀), is higher than that of P. duffmackayi, sp. nov., with 11–13, usually 12 (♂), and 10–12, usually 11 (♀).The prolateral t and st macrosetae of basitarsi III and IV are spiniform in P. militaris (fig. 22C, D), whereas t and st are setiform on III, and t setiform and st spiniform on IV in P. duffmackayi, sp. nov. (fig. 15C, D).

Redescription: The following redescription supplements the original description by Pocock (1900b) and is based on the material examined. Only characters that differ from P. cavimanus are described.

Total Length: Adult medium, maximum length, measured from anterior margin of carapace to tip of aculeus, 115 mm (99–123 mm, n = 7) (♂), 104 mm (95–112 mm, n = 10) (♀) (tables 4, 5).

Color: Chelicerae, dorsal surfaces bicolored, proximal three-quarters of manus dorsal surface with sparsely reticulate infuscation, markedly paler than carapace and densely infuscate distal quarter of manus dorsal surface and fingers. Pedipalp chela fingers densely infuscate, brownish black; carapace lateral and posterior surfaces, tergites, sternite VII, metasoma, telson and pedipalp trochanter, femur, patella pro- and retrolateral surfaces infuscate but paler, dark yellowish brown to olive brown, telson paler than metasoma; carapace interocular and circumocular surfaces, telson, pedipalp patella ventral surface, chela manus dorsal, lateral and ventral surfaces and legs immaculate, markedly paler, light yellowish brown; legs dorsal and retrolateral surfaces becoming paler distally, tibia, basitarsus, and telotarsus paler than femur and patella. Coxosternal region and sternites III–VI immaculate, yellowish. Genital opercula and pectines immaculate, uniformly pale cream.

Carapace: As for P. cavimanus, except as follows. Anterior width of posterior width, 67% (62%–72%, n = 7) (♂), 68% (65%–72%, n = 10) (♀); posterior width of length, 101% (91%–109%, n = 7) (♂), 102% (97%–105%, n = 10) (♀) (tables 4, 5). Median ocular tubercle situated medially, distance from anterior carapace margin 50% (47%–51%, n = 7) (♂), 49% (46%–52%, n = 10) (♀) of carapace length (tables 4, 5). Interocular and circumocular surfaces smooth or with few granules anterior to ocular tubercle (fig. 18A, C). Anterolateral, mediolateral, and posterolateral surfaces finely and sparsely granular, more coarsely and densely so in male. Posteromedian surfaces smooth or nearly so.

Pedipalps: As for P. cavimanus, except as follows. Femur width of length, 54% (48%–59%, n = 7) (♂), 55% (51%–60%, n = 10) (♀) (tables 4, 5). Retrodorsal carina obsolete, granular; more strongly developed than prodorsal carina. Prodorsal carina obsolete, comprising few rounded granules. Promedian carina distinct, comprising row of subspiniform granules (several demarcated by conspicuous macrosetae), oriented diagonally between prodorsal and proventral carinae. Patella width of length, 42% (39%–44%, n = 7) (♂), 43% (41%–46%, n = 10) (♀) (tables 4, 5). Chela fairly short, broad, base of fixed finger arising gradually from manus (figs. 20A, 21); manus, height of width, 59% (49%–68%, n = 7) (♂), 69% (64%–73%, n = 10) (♀); length along ventroexternal carina of width, 69% (64%–76%, n = 7) (♂), 66% (62%–72%, n = 10) (♀); length along ventroexternal carina of length movable finger, 51% (47%–55%, n = 7) (♂), 53% (49%–55%, n = 10) (♀) (tables 4, 5). Dorsomedian carina obsolete. Digital carina absent, except proximal to base of fixed finger, obsolete, granular to costate-granular. Proventral and promedian carinae obsolete, granular, each indicated by prominent macroseta. Prodorsal carina absent, but indicated by prominent macroseta. Chela densely (♂) to moderately (♀) setose on fingers and distally on manus. Manus, retrodorsal surface predominantly smooth (♂) to reticulate (♀) proximally with shallow, anastomosing granules distally; retrolateral surfaces shallowly granular; ventral intercarinal surface smooth; prolateral intercarinal surfaces predominantly smooth, with scattered spiniform granules dorsally and distally. Fixed and movable fingers, pro- and retrolateral intercarinal surfaces finely and sparsely granular; median denticle rows each with six enlarged retrolateral denticles (including terminal denticle), proximal three (♀) or second and third most proximal (♂) retrolateral denticles on fixed and movable fingers situated on lobes; fixed finger proximal lobe, similar in size to medial lobe (♀, immatures), vestigial, much smaller than medial lobe, or absent (adult ♂); movable finger (adult ♂) without proximal lobe, median lobe markedly more pronounced than other lobes, and with correspondingly well-developed notch in fixed finger; prominent gap proximally between fingers, when closed (adult ♂).

Trichobothria: As for P. cavimanus, except as follows. Neobothriotaxic major, Type C, with the following segment totals (n = 34; tables 4, 5): femur, 3 (1 d, 1 i, 1 e); patella, 51 (47–55): 2 d, 1 i, 34 (30–38) v, 14 (13–15) e, usually comprising 3 et, 2 est, 2 em, 2 esb, 5 eb; chela, 35 (33–36), manus, 22 (21–24), comprising 2 D, 10 E, 10 (9–12) V; fixed finger, 13 (12 or 13), comprising 4 d, 4 e, 5 (4 or 5) i (figs. 19–21). Total count of trichobothria per pedipalp: 89 (84–93). Chela, distance et–est half to less than half distance est–esb; est aligned with or distal to dst.

Legs: As for P. cavimanus, except as follows. Basitarsi, spiniform macrosetae, I, retrolateral: t, st, sb, b (polymorphic); retroventral: t; proventral: t, st; II, retrolateral: t, st, sb, b (polymorphic); retroventral: t; proventral: t, st; III, retrolateral: t, sb; retroventral: t; proventral: t, st; prolateral: t, st; IV, retrolateral: t; retroventral: t; proventral: t, st; prolateral: t, st (fig. 22). Telotarsi, macrosetal counts in pro- and retroventral rows equal on I–IV, 3 (1–4) and 4 (2–5) (n = 34), respectively (tables 4, 5). Telotarsal ungues subequal on telotarsi I–IV.

Pectines: As for P. cavimanus, except as follows. Distal edge reaching past distal edge of coxa IV but not reaching to distal edge of trochanter IV (♂, fig. 18B; ♀, fig. 18D). First proximal median lamella (scape) of each pecten with mesial margin obtusely angular, greater than 90° but less than 180° (♂) or sublinear (♀), and devoid of teeth in proximal 20% (14%–24%, n = 7) (♂) or 23% (10%–31%, n = 10) (♀) of prolateral margin (tables 4, 5). Pectinal tooth count, 14/15 (14–15/14–16, n = 7) (♂), 14/14 (14/13–15, n = 10) (♀).

Mesosoma: As for P. cavimanus, except as follows. Posttergites smooth and glabrous medially and anterolaterally, sparsely and finely granular posterolaterally. Sternite VII, length of width, 61% (49%–68%, n = 7) (♂), 56% (48%–67%, n = 10) (♀) (tables 4, 5).

Metasoma and Telson: As for P. cavimanus, except as follows. Metasomal segments I–V progressively increasing in length, decreasing in width; segment V, width of segment I width, 65% (58%–73%, n = 7) (♂), 63% (61%–67%, n = 10) (♀)(tables 4, 5). Metasoma robust; width of length, segment I, 95% (85%–121%, n = 7) (♂), 102% (90%–130%, n = 10) (♀); II, 80% (77%–83%, n = 7) (♂), 89% (79%–118%, n = 10) (♀); III, 73% (68%–79%, n = 7) (♂), 77% (70%–101%, n = 10) (♀); IV, 61% (53%–67%, n = 7) (♂), 65% (60%–85%, n = 10) (♀); V, 45% (38%–50%, n = 7) (♂), 50% (41%–55%, n = 10) (♀). Telson vesicle, width of metasomal segment V, width, 103% (87%–118%, n = 7) (♂), 90% (81%–110%, n = 10) (♀); enlarged (♂), globose, height of length, 59% (54%–70%, n = 7) (♂), 55% (49%–63%, n = 10) (♀); dorsal surface flat, ventral surface evenly curved. Aculeus relatively short, strongly curved, length of vesicle length, 33% (27%–37%, n = 7) (♂), 37% (33%–42%, n = 10) (♀). Length metasoma and telson, of total length, 49% (46%–52%, n = 7) (♂), 46% (42%–50%, n = 10) (♀). Dorsosubmedian carinae, segments I–IV, distinct, complete, granular, posterior spiniform granules obsolete; V, absent (fig. 23A). Dorsolateral carinae, segments I and II, obsolete, complete, granular; III–V, distinct, complete, granular. Median lateral carinae, segment I, obsolete, costate, restricted to posterior third; II–V, absent, demarcated by macroseta at posterior margin on I–IV and near anterior margin on V (fig. 23B). Ventrolateral carinae, segments I–IV, distinct, complete, costate on I–III, granular on IV; V, distinct, complete, comprising subspiniform granules, diverging posteriorly, terminal granule similar in size to preceding granules (fig. 23C). Ventrosubmedian carinae, segments I–III, distinct, complete, costate; IV, obsolete, complete, granular (♂) or costate-granular (♀); V, vestigial, each reduced to discontinuous row of spiniform granules, demarcated by conspicuous macrosetae. Ventromedian carina, segment V, comprising single row of spiniform granules, breaking up into numerous granules posteriorly. Dorsal and lateral intercarinal surfaces, segments I–V, smooth. Ventral intercarinal surfaces, segments I–III, smooth; IV and V, with scattered granules. Telson vesicle, dorsal and lateral surfaces smooth; ventral surface with four distinct longitudinal carinae, each comprising prominent spiniform granules, extending entire length of vesicle.

Geographical Variation: Specimens from the south of the distribution are usually yellowish brown in color, those from the north are reddish brown.

Ontogenetic Variation: As for P. cavimanus, except that immature stages are often markedly more infuscate than adults.

Sexual Dimorphism: As for P. cavimanus, except as follows. Pandinoides militaris is the most sexually dimorphic of the three species. The most obvious secondary sexual characters observed in adult males, compared with adult females and juveniles of both sexes, concern the modifications of the pedipalp chela: a marked concave depression is present in the retrodorsal surface of the manus, at the base of the fixed finger, the proximal lobe of the fixed finger is vestigial, much smaller than medial lobe, or absent, the proximal lobe of the movable finger is absent, the median lobe of the movable finger is markedly more pronounced than the other lobes, and a prominent gap is present proximally between the fingers, when closed (fig. 20A).

Distribution: Recorded from Ethiopia (Oromia Regional State), Kenya (Coast, Eastern, North Eastern and Rift Valley provinces) and Somalia (Gedo, Jubbada Hoose, Shabeellaha Dhexe and Shabeellaha Hoose regions). All records attributed to P. cavimanus from Somalia and most from Kenya, e.g., Kinani (Pocock, 1896) and Nguni, N of Ngomeni (Kovařík, 2009), are referable to P. militaris (fig. 2). The presence of P. militaris in Ethiopia is based on the type locality and an additional record, Arigalgalu [04°25′N 039°55′E], reported by Moriggi (1941: 94). Its presence in Tanzania remains to be confirmed, but unpublished reports of P. cavimanus from the vicinity of the Usambara Mountains may be referable to P. militaris. The alleged occurrence of P. militaris in Sudan (Lamoral and Reynders, 1975; Kovařík, 1998, 2002, 2009; Fet, 2000) is dubious. It was based on Roewer's (1943) record from “Dufile, Lado district, Sudan,” which is in the Moyo District of northern Uganda [03°33′N 031°57′E] and remains to be confirmed.

Ecology: The known locality records of P. militaris occur in arid savannah, dominated by Acacia and Commiphora, or riverine woodland dominated by Salvadora at 240–700 m elevation. Although several adult males were collected on the surface at night with UV light detection, most specimens for which data are available, were excavated from burrows during the day. Pandinoides militaris constructs burrows up to 75 cm long in red sandy-loam soils. A few specimens have been collected in or under rotten logs. This species is allopatric with P. cavimanus and parapatric with P. duffmackayi, sp. nov., but sympatric with Pandinurus gregoryi (Pocock, 1896), P. exitialis, and at least one species of Pandinops Birula, 1913. It appears to differ ecologically from P. exitialis, which has been collected under stones, and P. gregoryi, which has been collected under the peeling bark of fallen and standing trees.

Conservation: This species has been recorded in at least three protected areas in Kenya: the Kora National Park, Samburu Game Reserve and Tsavo West National Park.

Remarks: Pocock (1897b: 397, 398) regarded three specimens collected in Somaliland by A. Donaldson Smith as conspecific with S. bellicosus, stating:

Pocock's (1897b: 397) statement should actually read “the carapace in the latter being shorter than the 4th and 5th caudal segments, whereas in the former it is longer.”

Pocock (1898: 498) subsequently listed the specimen from Ndi (Weiss Road Camp), inland from Mombasa, as “? Scorpio bellicosus”:

Following Kraepelin's (1899) rediagnosis of P. bellicosus, Pocock (1900b: 62) realized that the specimens previously determined to that species (Pocock, 1897b, 1898), were misidentified:

Pocock (1900b: 61) described these specimens as P. militaris, diagnosing that species from P. cavimanus on the basis of the characters Pocock (1897b, 1898) had used to separate P. bellicosus from the latter:

Pocock's (1900b: 61) description of the holotype from El Dere near Aimola agrees with the redescription presented above:

In spite of these consistent differences (and many others mentioned in the diagnosis, above), Kovařík (2002: 20) synonymized P. militaris with P. cavimanus, based solely upon an appeal to authority: “I have examined the types of both species and conclude that Pandinus militaris Pocock, 1900, is a junior synonym of Pandinus cavimanus (Pocock, 1888).” This synonymy was repeated by Kovařík (2003, 2009) and perpetuated by Rossi (2015a). In view of the abundant differences described herein, P. militaris is hereby revalidated.

Additional Material Examined: KENYA: 1 subad. ♂ (NMK 259), 1 ♀ (NMK 260), Ritcher?, 1 ♂ (NMK 258), iii.1969, imported, 1 ♀ (AMNH), 1994, ex M. Scharmach, pet trade, 1 ♀ (AMNH [LP 341]). Coast Prov.: Taita Taveta Distr.: Jora, Kasigau [03°35′S 038°36′E], vii.1965, R. Dio, 1 ♂ (NMK 240); Kishushe [03°17′S 038°18′E], 25.vii–8.viii.2008, S. Mwangi, collected in burrows, 2 ♂, 5 ♀, 1 subad. ♂, 17 juv. ♂, 16 juv. ♀ (AMNH), 1 juv. ♂ (AMCC [LP 8879]); Voi, Segala region [Sagala, 03°29′S 038°36′E], xii.1971, K. Werner, 3 ♀ (NHRM [JF 58]); Tsavo West National Park: Ndi [Ndii, 03°14′S 038°30′E], Weiss Road Camp, inland from Mombasa, C.S. Betton, collected during construction of Mombasa-Uganda railway, 1 juv. ♂ (BMNH); Tsavo, Taita Discovery Centre, 03°25′S 038°46′E, 27.iii.2000, R. Jocqué and C. Warui, Acacia-Commiphora forest, by hand on ground, 1 ♂ (MRAC 209.615), night catch around center, 2 ♂ (MRAC 209.644, 210.006). Tana River Distr.: Kora National Park: Kampiya Ndovu [00°05′S 038°46′E], 30.iii.1984, Kilamuntu, 1 subad. ♂ (NMK 241); Kora Base Camp, 2.viii.1983, Kora Research Project, 1 ♂, 1 ♀, 1 subad. ♀ (NMK 242); near Kora expedition camp, 20.viii.1983, A. Duff-Mackay, in nest with multiple entrances, in Salvadora bush, 1 juv. ♂, 2 juv. ♀ (NMK 243), 21.viii.1983, A. Duff-Mackay, in rotten log on ground, 2 juv. ♂ (NMK 244); Kora National Reserve, 26.ii.1984, A. Duff-Mackay, 1 ♀, 1 juv. ♂ (NMK 245 old 323); Kora National Reserve, near base camp, vii.1983, M. Ritchie, M. Collins, J. Muhangani and M. Coe, 2 ♀, 2 juv. ♂ (NMK 246); near Kora, S bank Tana River, 780 ft, 5.viii.1976, A. Duff-Mackay, three found together in one burrow, 4 entrances, 3 blocked, transition riverine forest, dry Acacia/Commiphora, 1 subad. ♂ (NMK 253 old 8); Nzoka, Kora reserve [00°07′S 038°46′E], 2.ii.1983, Kora Research Project, in burrow, 1 ♀, 6 juv. ♂, 2 juv. ♀ (NMK 558). Tana River, between coast and Kamega [Kamuga, 00°04′S 038°11′E], xi.1892, Chanler's Expedition, 1 ♂ (USNM 57-275), 1 ♂ (USNM 57-316). Eastern Prov.: Machakos Distr.: viii–ix.1976, Mutisya, specimens died in snake park, 1 subad. ♂, 1 subad. ♀, 1 juv. ♀ (NMK 257 old 68). Tsavo West National Park: Lugard Falls, Athi River [03°03′S 038°41′E], iii.1937, Hitchens, 3 juv. ♀ (NMK 247). Makueni Distr.: Mtito Andei [02°41′S 038°10′E], vi.1950, E.C. Swete-Kelly, 2 ♂ (NMK 248, 249), 1 ♀, 1 subad. ♂ (NMK 250); Tsavo West National Park: Kinani [Kenani, 02°51′S 038°20′E], J.W. Gregory, 1 ♂ (BMNH 1893.11.9.4); Kinani, 4 mi. S [02°55′S 038°23′E], J.W. Gregory, 1 ♂ (BMNH 1893.11.9.5); near Kenani River, 02°48′S 038°18′E, 2200 ft, 13.ix.1976, A. Duff-Mackay, dug from burrow, single entrance, Acacia/Commiphora on red sand, newly sloughed, 1 ♂ with exuvium (NMK 251 old 71), six specimens dug together from hole with five entrances, Acacia/Commiphora on red sand, debris from burrow collected, 1 juv. ♂, 1 juv. ♀ (NMK 252 old 72). Kitui Distr.: near Mitamisyi, 00°31′S 038°24′E, 2300 ft, 7.viii.1976, A. Duff-Mackay, in burrow with single elliptical entrance, in red sand, Acacia/Commiphora woodland, 1 subad. ♂ (NMK 254 old 11). Marsabit Distr.: North Horr, 8 km N, 03°23′N 037°05′E, 2600 ft, 11.vii.1987, A. Duff-Mackay, burrow straight 28 ins. and 2 ins. deep, 1 subad. ♂ (NMK 255 old 341). Meru Distr.: Mweru River, 13.viii.1909, S.M. Allen, 1 ♀ (MCZ 15512). Rift Valley Prov.: Narok Distr.: Northern Guaso Nigero [Ewaso Ngiro] River [01°09′S 035°46′E], 1909, Heller, 1 subad. ♂ (USNM). Samburu Distr.: Samburu Game Reserve [00°35′N 037°32′E], campsite 5 or 6, 1976, L. Williams, under tent, one jumped off bush (lighter) and one of same type found on log on fire, sympatric with Pandinurus exitialis [NMK 561 old 101], 1 subad. ♂ (NMK 256 old 101). SOMALIA: Somaliland, A. Donaldson Smith, 2 ♀ (BMNH, 1897.11.10.4–5); Somalia, imported for pet trade: iii.2008, ex A. Tietz, 1 ♂ (AMNH [LP 10206]), [leg] (AMCC [LP 8273]), 1 ♀, 1 subad. ♂ (AMNH). Gedo Region: Juba [Jubba] River, C.L. Chevallier, 1 ♂ (BMNH 12.10.15.1). Shabeellaha Hoose Region: Afgooye Distr.: Afgoi [Afgoye/Afgooye], 15 km S [02°02′N 045°01′E], 25.vii.1961, C. Gans, under stump, 1 ♂ (MCZ 15517).

Pandinus (P.) imperator (C.L. Koch, 1841)

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):

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 single 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).

Pandinurus (P.) sudanicus Hirst, 1911

Type Material: Pandinus exitialis sudanicus: Holotype ♀ (BMNH), Gebel Mts., S of Obeid [North Kordofan], Sudan. Pandinurus (P.) prendinii: Holotype ♀ (HNHM 1443), Messina [Musina], Transvaal [Limpopo Province], South Africa, 1976, A. Héjja.

Distribution: The two new records from South Sudan presented here, while a considerable distance south of the type locality and Kovařík's (2012: 12) additional record (Lagowa, Kordofan)—both situated in Sudan, not South Sudan, as stated by Kovařík (2012)—are clearly conspecific with the latter based on similar size, coloration, macrosculpture (anastomosing tubercles) on the retrodorsal surface of the pedipalp chela manus in the adult male and female, and counts of pedipalp trichobothria and pro- and retroventral spiniform macrosetae on the telotarsi.

Remarks: Pandinus exitialis sudanicus Hirst, 1911, was synonymized with Pandinus magrettii Borelli, 1901, by Birula (1927: 85). Kovařík (2012: 3, 6) revalidated the taxon and elevated it to the rank of species, a move that appears to be reasonable, based on material examined for the present contribution. The retrodorsal surface of the pedipalp chela manus in the adult male and female of P. sudanicus comprises anastomosing tubercles, as in P. magrettii, rather than conspicuously pointed granules, as in P. exitialis, and the medial lobe on the chela movable finger of the adult male is considerably larger than the other lobes on the finger, as in P. exitialis, which is not the case in P. magrettii.

Rossi (2015a: 20, 21) described Pandinurus (P.) prendinii on the basis of a single female, allegedly from Messina in the Limpopo Province of South Africa, noting its close morphological similarity to P. sudanicus including, for example, the macrosculpture on the retrodorsal surface of the pedipalp chela manus in the adult female, which comprises anastomosing tubercles, evident in Rossi's (2015a: 53) figures 32 and 33. Two other workers (F. Kovařík, in 1996; B.F. Striffler, in 2002) previously examined the specimen and identified it as “Pandinus sp.,” probably on account of the erroneous locality data. The meristic data (total body length, pedipalp trichobothrial counts, pectinal tooth counts and counts of spiniform macrosetae in the pro- and retroventral rows of the telotarsi) listed as diagnostic for P. prendinii by Rossi (2015a: 20) fall entirely within the ranges given for P. sudanicus two pages later (table 6). The suggestion that P. prendinii may represent a relictual population of Pandinus, sensu lato in southern Africa, and that, because the putative type locality is 5000 km from the known localities of P. sudanicus, this could justify its recognition as distinct from the latter rests entirely on the grossly mistaken assumption that the locality data are trustworthy, an assumption doubted by others (Kovařík, 1997). Despite extensive surveys of scorpions throughout southern Africa, including the vicinity of Messina, by the author, there are no credible records of Pandinus, sensu lato, from south of the Zambezi River, as noted by Prendini et al. (2003; also see Prendini, 2005), and especially not south of the Limpopo River. In view of the evidence, the following new synonym is presented: Pandinurus (P.) prendinii Rossi, 2015 = Pandinurus (P.) sudanicus (Hirst, 1911), syn. nov.

TABLE 6

Putatively diagnostic meristic data for species of Pandinurus Fet, 1997, provided by Rossi (2015a): total body length (mm); counts of trichobothria in V series of pedipalp chela manus; pectinal tooth counts; counts of spiniform macrosetae in pro- and retrolateral rows of telotarsi I–IV.

Additional Material Examined: SOUTH SUDAN: Boma State: Boma National Park [06°29′N 033°55′E], base camp, 8.iv.1983, T. Tear, 2 ♂ (AMNH). Eastern Equatoria State: Torit [04°25′N 032°34′E], Equatoria, Anglo-Egyptian Sudan, 2000 ft, xii.1949, H. Hoogstraal, 2 ♂, 6 ♀, 2 subad. ♂, 3 juv. ♂, 2 juv. ♀ (FMNH). SUDAN: Blue Nile State: Ingessana Hills, SW of Roseires, 11°27′N 033°59′E, xii.1968, J.L. Cloudsley-Thompson, 1 ♀ (MRAC 134.601).

Pandinurus (Pandiborellius) percivali (Pocock, 1902)

Type Material: Pandinus percivali: Holotype ♀ (BMNH 1899.12.2.5) [examined], Al-Khaur, Abian country, 80 mi. E of Aden, Arabia [Yemen]. Pandinurus (Pandicaporiaccous) janae: Holotype ♀ [immature] (HNHM 1442), paratype ♀ (ARPC 0253 ex HNHM), Wadi Zabid, Yemen, xi.1969, A. Szalay-Marzsó.

Remarks: Rossi (2015a) based P. janae and the new monotypic subgenus into which it was placed, on a single female from the general vicinity of the type locality of P. percivali, the species to which it was previously determined by other workers (F. Kovařík, in 1996; B.F. Striffler, in 2002). Rossi (2015a: 31) mistakenly regarded the specimen from Wadi Zabid to be adult, an assumption that formed the basis of his conclu sion that it was misidentified by Kovařík (1997, 2009) and his justification for its description as a new species and subgenus:

It is, however, clear from Rossi's (2015a: 36, 57, table 1) meristic data and figures 67–73, that the holotype of P. janae is immature and, based on the distinctive shape of the pedipalp chela and the prominent costate carinae on the manus, as well as the metasoma and habitus, that it is indeed conspecific with P. percivali as F. Kovařík and B.F. Striffler surmised. The putatively diagnostic differences in total body length between P. janae and P. percivali may be dismissed on account of the immaturity of the holotype of the former. The pedipalp trichobothrial and pectinal counts of P. janae are close to those of P. percivali (table 6). In the absence of other significant differences, the lower counts of spiniform macrosetae in the pro- and retroventral rows of the telotarsi of P. janae cannot justify its recognition as distinct from P. percivali, because the sample sizes (n = 2 for P. janae; n = 1 for P. percivali) are far too small to assess the variation, and anomalous telotarsal setal counts are not uncommon in scorpionids, based on thousands of specimens examined by the author. In view of the evidence, the following new synonym is presented: Pandinurus (Pandicaporiaccous) janae Rossi, 2015 = Pandinurus (Pandiborellius) percivali (Pocock, 1902), syn. nov.

Pandinurus (Pandipalpus) viatoris (Pocock, 1890)

Type Material: Scorpio viatoris: Holotype ♂ (BMNH 1890.4.15.1) [examined], “East Africa” [probably Malawi or Zambia]. Pandinurus (Pandipalpus) bartolozii: Holotype ♂ (MZUF 1059), allotype ♀ (MZUF 1456), Kolwezi, Shaba, Democratic Republic of the Congo (D.R.C.), ii.1990, D. Mosca. Pandinurus (Pandipalpus) flagellicauda: Holotype ♂ (MZUF 1457), allotype ♀, 4 ♀ paratypes (MZUF 1458), 1 ♂, 1 ♀ paratypes (ARPC 0251, 0252 ex MZUF), Kolwezi, Shaba, D.R.C., ii.1990, D. Mosca. Pandinurus (Pandipalpus) lorenzoi: Holotype ♂ (ARPC 0024), Morogoro, Kigulunyembe, Mt. Uluguru, Tanzania, 18-31.iii.2008. Pandinurus (Pandipalpus) pantinii: Holotype ♂ (MSNB), Balaka, Malawi, xii.2004, E. Ferrario. Pandinurus (Pandipalpus) pygmaeus: Holotype ♀ [subad.] (MSNB), Likasi, S.-E. Shaba, D.R.C., ii.1986, K. Tshikamba.

The type locality of P. viatoris is unknown, but the holotype described by Pocock (1888) probably originated from former British colonial territory, either Northern Rhodesia (now Zambia) or Nyasaland (now Malawi), rather than former German colonial territory (Tanganyika, now Tanzania) as suggested by Rossi (2015a: 65, fig. 125). There are at present no confirmed records of P. viatoris from Kenya (former British East Africa).

Distribution: Prendini et al. (2003: 234) discussed putative records of P. viatoris from Zimbabwe, demonstrated that these records actually occur in Zambia, and noted that there are no credible records of Pandinus south of the Zambezi River. Kovařík (2012: 19, fig. 64) did not cite Prendini et al. (2003) and erroneously plotted three points for P. viatoris south of the Zambezi River, one in the extreme north of Zimbabwe (presumably, the specimen erroneously labelled “Mashonaland” [ZMB 35310]) and two in Mozambique, corresponding to the approximate locations of Tete and Beira, where this species does not appear to occur based on surveys by the author. Rossi (2015a: 29) discussed the reliability of Kovařík's (2012) Zimbabwean record, but apparently missed the point about the Zambezi River, as he did not mention the other two localities, and went on to describe a new Pandinurus species allegedly from Messina, in the Limpopo Province of South Africa (see above).

Remarks: Rossi (2015a) described five new species in a new subgenus Pandipalpus, three based on singletons, among them an immature specimen, two from the same locality in the Democratic Republic of the Congo, and one from a locality nearby. With one exception, the meristic data (total body length, pedipalp trichobothrial counts, pectinal tooth counts and counts of spiniform macrosetae in the pro- and retroventral rows of the telotarsi) listed as diagnostic for the five species by Rossi (2015a: 23–25, 27, 28, 36, 37, table 1, 2) overlap entirely with one another and with the ranges given for P. viatoris (reproduced here in table 6) with which they are evidently conspecific. The putatively diagnostic difference in total body length between P. pygmaeus and the other species may be dismissed on account of the immaturity of the holotype of the former. As with P. janae, discussed above, Rossi (2015a: 29) mistakenly regarded the holotype of P. pygmaeus to be adult and this assumption formed the basis of his justification for its recognition as a new species: “Pandinurus (Pandipalpus subgen. n.) pygmaeus n. sp. ha dimensioni minori, 83 mm, contro 100–125 mm.” It is, however, clear from Rossi's (2015a: 28, 36, 61, table 1) meristic data and figures 99–105 that the holotype of P. pygmaeus is immature and that it is conspecific with P. viatoris. The holotypes of P. bartolozii and P. flagellicauda, originally part of the same series of specimens, were previously examined by F. Kovařík (in 1997), considered conspecific, and identified as P. viatoris (Kovařík and Whitman, 2005: 114; Kovařík, 2012: 21). Curiously, Rossi (2015a: 25) appears to have regarded the sympatric occurrence of these specimens as evidence for their distinctiveness, despite the lack of data supporting any possible ecological difference:

In view of the evidence, the following new synonyms are presented: Pandinurus (Pandipalpus) bartolozii Rossi, 2015 = Pandinurus (Pandipalpus) viatoris (Pocock, 1890), syn. nov.; Pandinurus (Pandipalpus) flagellicauda Rossi, 2015 = Pandinurus (Pandipalpus) viatoris (Pocock, 1890), syn. nov.; Pandinurus (Pandipalpus) lorenzoi Rossi, 2015 = Pandinurus (Pandipalpus) viatoris (Pocock, 1890), syn. nov.; Pandinurus (Pandipalpus) pantinii Rossi, 2015 = Pandinurus (Pandipalpus) viatoris (Pocock, 1890), syn. nov.; Pandinurus (Pandipalpus) pyg maeus Rossi, 2015 = Pandinurus (Pandipalpus) viatoris (Pocock, 1890), syn. nov. Ongoing research has revealed low levels of genetic divergence between populations of P. viatoris in Malawi and Tanzania, further supporting the conclusion that P. viatoris is a single, panmictic species (L. Prendini, unpublished data).

Additional Material Examined: 1 subad. ♀ (AMNH). DEMOCRATIC REPUBLIC OF CONGO: Haut-Katanga Prov.: Lukafu, 10°31′S 027°33′E, 1930, G.F. de Witte, 1 ♀ (MRAC 61.608 old 23.806), 6–22.xii.1930, G.F. de Witte, 3 juv. ♀ (MRAC 23.970–972), 1 juv. ♂, 2 juv. ♀ (MRAC 23.973–975); Monasterie N.O. de Béthanie [04°24′S 019°19′E], Katanga, 10°00′S 027°00′E, 1947, R.R. Soeurs, 1 ♂ (MRAC 57.722). Haut-Lomami Prov.: Bukama, 09°12′S 025°51′E, 1937, P. Brien, 1 ex. (MRAC 168.782); Kabongo, 07°20′S 025°35′E, 1952, Dierkx, 1 ♀ (MRAC 73.615); Kamina [08°44′S 024°59′E], Luabo [09°03′S 024°47′E], 1949, Ecole Normale, 1 ♀ (MRAC 65.801); Kisanga [06°51′S 024°10′E], Kele [06°55′S 023°10′E], Katanga, 1959, R.F.A. van Oost, 5 ex. (MRAC 207.425). Lualaba Prov.: Muema, 10°25′S 024°49′E, 1927, A. Bayet, 3 ♂, 2 juv. (MRAC 130.812), 3 ex. (MRAC 130.813). Zilo [10°30′S 025°28′E], Shaba, v.1991, 1 ♂ (NHRM [JF, 57]). Maniema Prov.: Misosa [?Musasa, 03°53′S 026°49′E], vii.1939, H.J. Bredo, 13 ex. (MRAC 57.597–610), 19 ex. (MRAC 57.611–630), 19 ex. (MRAC 57.631–650). North Kivu Prov.: Beni, 00°29′N 029°28′E, Lisfrane, 1 ♂, 1 ♀ (MRAC 57.743). Tanganyika Prov.: Tanganika, J. Hecq, 1 ♂ (MRAC 130.814); Baudouinville [Kirungu], 07°02′S 029°47′E, iii.1954, H. Bomans, 1 juv. (MRAC 78.968). KENYA: Rift Valley Prov.: Narok Distr.: S Guaso Nejiro, Toita plains, 1 subad. ♀ (USNM). MALAWI: Southern (Blantyre) Region: Mangochi Distr.: Fort Johnston [Mangochi, 14°29′S 035°16′E], Sir Alfred, 1 ♂ (TM 17462 ex AM 2179); Mangochi [14°28′S 035°16′E], Rift Valley 1 ♂ (AMNH); Monkey Bay [14°03′S 034°55′E], 14.vii.1975, D. Eccles, 1 ♀ (AMNH [AH 204]), 3 juv. ♂ (AMNH [AH 191, 192, 230]), 2 juv. ♀ (AMNH [AH 229, 231]), 19.vi.1975, D. Eccles, found in burrow with mother, 1 juv. ♂, 1 juv. ♀ (AMNH), 23.xii.1993, C.R. Owen, 1 juv. ♂, 1 juv. ♀ (NHRM [JF 140]); Monkey Bay, ca. 1 km S on road S128 to Mangochi, 14°06′25″S 034°55′01″E, 13.xii.2007, L. Prendini & W.R. Schmidt, 485 m, mesic savanna (Miombo Woodland) on flat plain with low granite/dolerite koppies (plains in between), coarse sandy-loam soils, UV light detection on warm, dark (overcast), humid night after rain, sitting motionless on soil surface near base of tree, 1 ♂ (AMNH), [leg] (AMCC [LP 8025]). Zomba Distr.: Chinsewu Village, 20.5 km W Zomba on Namadidi Road, 15°19′20″S 35°11′28″E, 11–14. xii.2007, L. Prendini, W.R. Schmidt & R. Mbaya, 584 m, mesic savanna (Miombo Woodland) remnants on flat plain in old lands (subsistence agriculture/hoed fields of maize) in between freshly-hoed lands, well-drained sandy/clayey-loam soil, excavated from burrows in open ground and at base of bushes, burrows often multi-entranced, with several individuals (mixed sex and age) occupying same burrow, 13 ♂, 14 ♀, 6 subad. ♂, 9 subad. ♀, 21 juv. ♂, 20 juv. ♀ (AMNH), 2 juv. ♀ (AMCC [LP 8026]). MOZAMBIQUE: Chifurubasi, iv–v.1905, W. Tiesler, 1 ♂ (ZMB 1092/05). Nampula Prov.: Nampula Distr.: Nampula Mountain [Nampula, 15°07′S 39°16′E, iii.1976, K. Groseh, 1 ♀ (NM 10984). Zambézia Prov.: Gurué Distr.: Lioma [15°10′S 036°48′E], 22.ii.2001, C.R. Owen, 1 ♀ (NHRM [JF 110]), 1 subad. ♂ (NHRM [JF 112]), 28.ii.2001, C.R. Owen, 1 ♀ (NHRM [JF 111]). TANZANIA: Imported for pet trade, probably Serengeti area, 1994, ex M. Scharmach, 2 ♂, 1 subad. ♀, 1 juv. ♂ (AMNH [LP 332]), 2 ♂ (AMNH [LP 337]), 1 ♂ (AMCC 101704 [LP 1557]). Arusha Prov.: Karatu Distr.: Karatu, 03°21.177′S 035°48.871′E, 6.viii.2015, M. Roppo, 1264 m, 1 juv. [leg] (AMCC [LP 13453]). Dodoma Prov.: Kondoa Distr.: Kath Mission, Kwa Mtoro [05°14′S 035°26′E], Ussandawi, i–ii.1912, Dr. E. Obst, 1 ♂, 2 ♀, 1 subad. ♀ (ZMH). Lindi Prov.: Kilwa Distr.: Mbarawala Plateaux, 09°02.374′S 039°07.206′E, 29.ii–4.iii.2008, P. Hawkes, 270 m, 1 ♂ (AMNH). Liwale Distr.: Liwale [09°08′S 038°17′E], iv.1948, C.P.J. Iomides, 1 ♂ (NMK 295). Mara Prov.: Serengeti Distr.: Serengeti, 1 ♀ (NMK 298), x.1949, 1 ♂ (NMK 296), ii.1959, 1 juv. ♂ (NMK 297 old 42). Seren geti National Park, Wandamu Kopjes, 02°29.441′S 034°55.367′E, 1.viii.2015, M. Roppo, 1523 m, 1 juv. [leg] (AMCC [LP 13452]). Rukwa Prov.: Mpanda Distr.: Lake Tanganyika, Smithsonian Institution African Expedition, 1 ♂ (USNM 156/396); Mpanda town, 06°30′S 031°30′E, xii.1980, K.M. Howell, 1 ♀ (MRAC 159.310). Singida Prov.: Singida Distr.: Mangasini [04°53′S 034°09′E], 14.xii.1929, A. Loveridge, 2 ♂, 1 ♀ (MCZ 15573); Usandawi, 1929–1930, Fliegner, 1 ♀ (ZMB 146D/33); Usandawi, Mangasini [04°53′S 034°09′E], 13.xii.1929, A. Loveridge, 1 juv. ♀ (MCZ 15572). ZAMBIA: Eastern Prov.: Chipata Distr.: Chief Sayiri area, 30 mi. from Fort Jameson [Chipata, 13°38′S 032°39′E], 6–18.i.1965, V.J. Wilson, 2 ♂, 2 subad. ♂, 2 subad. ♀, 1 juv. ♂ (NM 9090); Fort Jameson, V.L. Wilson, 2 ♂, 2 ♀ (NM 18772), i.1965, V.J. Wilson, 1 ♂ (TM 17461). Northern Prov.: Mpulungu Distr.: Mpulungu, 08°46′S 031°08′E, 1994, P.D. Plisnier, 1 ♂ (MRAC 209.525).

Erroneous Record: Mashonaland [Zimbabwe], 1893, R. Bartels, 1 ex. (ZMB 35310 old 1247/1911).