Pandinoides duffmackayi, Prendini, 2016

Prendini, Lorenzo, 2016, Redefinition And Systematic Revision Of The East African Scorpion Genus Pandinoides (Scorpiones: Scorpionidae) With Critique Of The Taxonomy Of Pandinus, Sensu Lato, Bulletin of the American Museum of Natural History 2016 (407), pp. 1-66 : 22-34

Pandinoides duffmackayi , sp. nov.

Figures 1C , 2 , 10–16 ; tables 2, 3

Scorpio cavimanus: Pocock, 1888: 249 (part); Pocock, 1896: 431, 432 (part).;

Pandinus cavimanus: Kraepelin, 1899: 121 , 122 (part); 1913: 183 (part); Werner, 1936: 185, 186 (part); Probst, 1973: 327 (part).

Pandinus (Pandinoides) cavimanus: Lamoral and Reynders, 1975: 564 (part); Fet, 2000: 468 (part); Kovařík, 2009: 1, 50, 52, 114 (part), table 3 (part), figs. 286, 290 [misidentification].

HOLOTYPE: ♂ ( AMNH), TANZANIA: Aru- sha 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) . Rifl 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 x 25.2 mm in red sand, dry Aca- cia / Commiphora , 1 ♀ ( NMK 232 old 75), dug from hole 14.6 x 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 pro- and 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 compris- ing 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:

I have seen two specimens of [ Scorpio cavimanus ]—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.... The specimen from Kilima-Njaro is smaller and slightly less granular than the other.

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.

Figures

FIGURE 1. Pandinoides Fet, 1997, habitus in life. A, B. Pandinoides cavimanus (Pocock, 1888) adults seized from shipment for exotic pet trade by U.S. Fish and Wildlife Service (A); ♀ (AMNH), imported to the United States from Tanzania (B). C. Pandinoides duffmackayi, sp. nov., paratype ♂ (AMNH), imported to U.S. from Tanzania. D. Pandinoides militaris (Pocock, 1900), ♂, Tsavo, Kenya. Photos courtesy Randy Mercurio (B, C) and Corwin Trent (D).

FIGURE 2. Map plotting known distributions of Pandinoides cavimanus (Pocock, 1888) (circles), Pandinoides duffmackayi, sp. nov. (triangles), and Pandinoides militaris (Pocock, 1900) (squares) based on data from the literature and the present contribution. White symbols denote type localities. Question marks denote unconfirmed, dubious records.

FIGURE 10. Pandinoides duffmackayi, sp. nov., habitus, dorsal (A, C) and ventral (B, D) aspects. A, B. Holotype ♂ (AMNH), C, D. Paratype ♀ (AMNH), Kikatitti, Tanzania. Scale bars = 10 mm.

FIGURE 11. Pandinoides duffmackayi, sp. nov., carapace, dorsal aspect (A, C), and sternum, genital opercula, and pectines, ventral aspect (B, D). A, B. Holotype ♂ (AMNH), C, D. Paratype ♀ (AMNH), Kikatitti, Tanzania. Scale bars = 2.5 mm.

FIGURE 12. Pandinoides duffmackayi, sp. nov., holotype ♂ (AMNH), Kikatitti, Tanzania, dextral pedipalp patella, retrolateral (A) and ventral (B) aspects. Scale bar = 2.5 mm.

FIGURE 13. Pandinoides duffmackayi, sp. nov., holotype ♂ (AMNH), Kikatitti, Tanzania, dextral pedipalp chela, retrolateral (A), ventral (B), and prolateral (C) aspects. Scale bar = 2.5 mm.

FIGURE 14. Pandinoides duffmackayi, sp. nov., paratype ♀ (AMNH), Kikatitti, Tanzania, dextral pedipalp chela, retrolateral aspect. Scale bar = 2.5 mm.

FIGURE 15. Pandinoides duffmackayi, sp. nov., paratype ♂ (AMNH), Kikatitti, Tanzania, legs I–IV, tibiae, basitarsi and telotarsi, ventral aspect (A–D). Scale bar = 1 mm.

FIGURE 16. Pandinoides duffmackayi, sp. nov., holotype ♂ (AMNH), Kikatitti, Tanzania, metasoma and telson, dorsal (A), lateral (B), and ventral (C) aspects. Scale bar = 2.5 mm.