Parabuthus Muelleri, and Implications for the Phylogeny of Parabuthus (Scorpiones: Buthidae)

AMERICAN MUSEUM NOVITATES Friday May 02 2003 03:12 PM novi 03331 Mp_1 Allen Press • DTPro System File # 01TQ

PUBLISHED BY THE AMERICAN MUSEUM OF NATURAL HISTORY CENTRAL PARK WEST AT 79TH STREET, NEW YORK, NY 10024 Number 3408, 24 pp., 7 ®gures, 4 tables May 22, 2003

Discovery of the Male of Parabuthus muelleri, and Implications for the Phylogeny of Parabuthus (Scorpiones: Buthidae)

LORENZO PRENDINI1

ABSTRACT The male of Parabuthus muelleri Prendini, 2000 is described, based on a specimen discovered in the Alexis Harington Scorpion Collection (recently acquired by the American Museum of Natural History). This is only the third known specimen of P. muelleri. The holotype and paratype are both female. The male presents several character states, including the lobate condition of the pectinal proximal median lamella and pedipalp chelae that are not incrassate, that are uncommon in male Parabuthus Pocock, 1890. These character states, previously scored with missing entries in a cladistic character matrix for Parabuthus species, are now added and a reanalysis of Parabuthus phylogeny, resulting in new insights about the phylogenetic position of P. muelleri, is presented. Lectotypes are designated for four northeastern African species of Parabuthus.

INTRODUCTION and 8 species occurring in northeast Africa and the Arabian Peninsula (Prendini, 2001a). All Parabuthus Pocock, 1890 is an exclusively but six of the southern African species have Old World genus of scorpions, 1 of 82 genera been reported from Namibia (Lamoral, 1979), in the diverse, cosmopolitan family Buthidae with four being endemic to that country, of (Fet and Lowe, 2000; KovarÏõÂk, 2001, 2002). which P. muelleri Prendini, 2000 is the most The genus displays a classic ``arid corridor'' recent addition. pattern of distribution (Balinsky, 1962), with At the time of its description, P. muelleri 20 species occurring in southwestern Africa was known from only two adult female spec-

1 Assistant Curator, Division of Invertebrate Zoology, American Museum of Natural History. e-mail: lorenzo@ amnh.org

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imens. A thorough search through the collec- digital imaging system. Measurements were tions of the National Museum of Namibia made with Mitutoyo digital calipers. Color (Windhoek), the South African Museum designation follows Smithe (1974, 1975, (Cape Town), the Transvaal Museum (Pre- 1981), trichobothrial notation follows Va- toria), and the Natal Museum (Pietermaritz- chon (1974), and mensuration follows Stahn- burg), all with extensive holdings of sorted ke (1970) and Lamoral (1979). Morphologi- and unsorted Parabuthus material from cal terminology follows Couzijn (1976) for southern Africa, revealed no additional spec- the segmentation of legs, Hjelle (1990) and imens. The absence of adult male specimens Sissom (1990) for the segmentation of ped- prevented several important characters per- ipalps, and Stahnke (1970), Lamoral (1979), taining to sexual dimorphism and the male Sissom (1990), and Prendini (2000a, 2001a) genitalia from being described for P. muel- for remaining features. leri, and these characters also had to be As in previous papers (Prendini, 2000a, scored with missing entries in a previously 2001a), the terms used by other authors on published cladistic character matrix for Par- the southern African scorpion fauna (East- abuthus species (Prendini, 2001a). wood, 1977; Lamoral, 1977, 1979; Fitz- A single adult male specimen has since Patrick, 1994) for certain metasomal carinae been discovered in the Alexis Harington have been replaced with terms implying spe- Scorpion Collection (recently acquired by the ci®c homology statements between carinae American Museum of Natural History). This on segment V and those on the preceding is only the third known specimen of P. muel- segments. The term ``ventral'' (segments I± leri, but it is suf®cient to update the diag- V) is replaced with ``ventrosubmedian'' (seg- nosis and description of the species, and pro- ments I±IV only) and ``ventromedian'' (seg- vides additional diagnostic characters to dis- ment V only) and the terms ``dorsal'' (seg- tinguish it from the sister species, P. capensis ments I±IV only) and ``dorsal accessory'' (Ehrenberg, 1831). The male specimen pre- (segment V only) are replaced with ``dorso- sents several character states, including the submedian''. lobate condition of the pectinal proximal median lamella and pedipalp chelae that are not Cladistic Analysis incrassate, that are uncommon in male Par- abuthus. These character states are now add- The present analysis is based on the pre- ed to the cladistic character matrix for Par- viously published morphological data matrix abuthus species (Prendini, 2001a), and a re- for relationships among the species of Par- analysis of Parabuthus phylogeny, resulting abuthus (Prendini, 2001a), to which the in new insights about the phylogenetic po- states of characters 9±12, 17, and 23, previ- sition of P. muelleri, is presented. ously scored with missing entries for the male of P. muelleri, have now been added MATERIALS AND METHODS (table 1; appendix 2). The matrix comprises 51 characters, 9 coded into multistates and Material, Photography, and Terminology 44 coded into binary states, scored for 27 The single adult male specimen of P. species. Multistate characters were treated as muelleri originates from the Alexis Haring- unordered, i.e., nonadditive (Fitch, 1971). ton Scorpion Collection (AH), which is now Trees were rooted using the outgroup deposited in the American Museum of Nat- method (Watrous and Wheeler, 1981; Farris, ural History (AMNH). Consult appendix 1 1982; Nixon and Carpenter, 1993). As in the for the repositories of other material exam- previous analysis, an exemplar species from ined for the cladistic analysis, where the full each of two Afrotropical buthid genera, collection data (previously unpublished) are Grosphus Simon, 1888, from Madagascar, provided and lectotypes designated for four and Uroplectes Peters, 1861, from southern northeastern African species. and central Africa, were included as out- Photographs of P. muelleri were taken in group taxa on the basis of morphological and visible light as well as under long-wave ul- molecular evidence that these genera are traviolet light using a Microptics ML1000 most closely related to Parabuthus (Pocock, AMERICAN MUSEUM NOVITATES Friday May 02 2003 03:12 PM novi 03331 Mp_3 Allen Press • DTPro System File # 01TQ

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TABLE 1 Distribution of 53 Characters Among 25 Species of the Genus Parabuthus Pocock, 1890 The ®rst two taxa are outgroups. Refer to appendix 1 for character list. Character states are scored 0 to 2, ? (unknown), ± (inapplicable) or * (polymorphic).

1890; Kraepelin, 1908; Werner, 1934; Pren- (hold 10,000 trees in memory; hold 10 start- dini and Wheeler, in prep.). ing trees in memory; perform TBR branch- Character data were edited, cladograms swapping on 100 random-addition repli- prepared, and character optimizations con- cates). Additional swapping on up to 1000 ducted using WinClada, vers. 0.9.9ϩ (Nixon, trees that are up to 5% longer than the short- 1999). Ambiguous optimizations were re- est trees (command jump 50;) was per- solved using accelerated transformation formed to help the swapper move between (ACCTRAN) or Farris optimization, which multiple local optima (``islands'' sensu Mad- favors reversals over parallelisms to explain dison, 1991). Finally, trees found with this homoplasy (Farris, 1970; Swofford and Mad- command were again swapped with TBR, dison, 1987, 1992) and therefore maximizes using the command max*; to retain only op- homology (Griswold et al., 1998). Three au- timal trees. tapomorphies (characters 4, 21, and 50) were Successive approximations character excluded from all analyses; hence, tree sta- weighting (Farris, 1969) and implied char- tistics are calculated from phylogenetically acter weighting (Goloboff, 1993, 1995) were informative characters only (Bryant, 1995). conducted to assess the effects of weighting Characters were not weighted a priori. against homoplasious characters, and the re- Analyses with equal weighting were con- sultant topologies were compared with the ducted using NONA vers. 2.0 (Goloboff, topology obtained by analysis with equal 1997a), according to the following command weights (see Prendini, 2000b, 2001a). Suc- sequence: hold10000; hold/10; mult*100; cessive weighting, using the squared consis- AMERICAN MUSEUM NOVITATES Friday May 02 2003 03:12 PM novi 03331 Mp_4 Allen Press • DTPro System File # 01TQ

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TABLE 2 Summary of Statistical and Topological Diferences Among the Most Parsimonious Trees (MPTs) MPTs were obtained by analysis with equal weights (EW), successive weights (SW), and implied weights (IW) with six values for the concavity constant (k), arranged in order of decreasing ®tness. Unweighted length is reported for the SW tree. Letters A and B refer to alternative topologies for node A (®g. 1).

tency index (CI) as a weighting function ues for the concavity constant were moderate (Goloboff, 1991), was implemented with to mild (i.e., k ϭ 3±6; table 2). Although NONA by invoking the swt.run ®le (com- topologically identical, the MPTs obtained mand sequence: run swt.run hold10000; by the analyses with implied weights were hold/10; mult*100; jump50; max*;). Pee- 3±7% ®tter than the MPTs obtained by anal- Wee version 2.6 (Goloboff, 1997b) was used ysis with equal weights, while the MPT re- for analyses with implied weighting, apply- trieved under k ϭ 5 was also one step shorter. ing the command sequence: hold1000; hold/ In contrast, under strong concavity (k ϭ 1± 10; mult*100; jump50; max*;. Analyses 2), analyses with implied weights located with implied weighting investigated the use two MPTs, each three steps longer and 5± of six values for the concavity constant, k, 12% less ®t than the MPTs obtained by anal- spanning the input range permitted by Pee- ysis with equal weights (table 2). These trees Wee (command: conc N;). differed from the topology in ®gure 1 with The relative degree of support for each respect to the species comprising node ``A'', node in the tree obtained with equal weight- for which the alternative arrangements were ing was assessed with branch-support or de- as follows: (P. mossambicensis (P. kraepelini cay indices (Bremer, 1988, 1994; Donoghue ((P. raudus ϩ P. schlechteri)((P. transvaal- et al., 1992). Branch support indices up to icus ϩ P. villosus)(P. capensis (P. muelleri ®ve extra steps (setting the maximum num- (P. calvus ϩ P. pallidus ϩ P. planicau- ber of trees held in memory to 10,000) were da))))))); (P. mossambicensis (P. kraepelini calculated with NONA by means of the fol- ((P. raudus ϩ P. schlechteri)(P. transvaal- lowing command sequence: h10000; bsup- icus (P. villosus (P. capensis (P. muelleri (P. port 5;. calvus ϩ P. pallidus ϩ P. planicauda)))))))). The MPTs obtained by analysis with im- RESULTS AND DISCUSSION plied weights under k ϭ 1±2 are longer and Analysis of the 50 informative characters less ®t than the MPTs obtained by the re- in NONA located a single most parsimonious maining analyses, and hence they are consid- tree (MPT) with equal weights (table 2; ®g. ered to be suboptimal. The alternative topol- 1). The same topology was retrieved in the ogy, obtained by weighting regimes that min- analysis with successive weights and also in imized length as well as those that maximized the analyses with implied weights when val- ®t (table 2), is instead regarded as optimal. AMERICAN MUSEUM NOVITATES Friday May 02 2003 03:12 PM novi 03331 Mp_5 Allen Press • DTPro System File # 01TQ

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Unambiguously optimized synapomorphies TYPE MATERIAL: Holotype & (NMNW are indicated on this topology in ®gure 1, 1854), Namibia: Hardap Region: MaltahoÈhe which also provides branch support values for District: Farm Onis 8, 82 km from Sesriem

nodes. The length, ®t (fi), consistency indices, to Naukluft, 24Њ22.46ЈS, 16Њ13.17ЈE, 1260 retention indices, and ®nal successive weights m, 7.i.1998, L. Prendini and E. Scott. Para- of informative characters on this topology are type & (SAMC C4514), Namibia: Karas Re- listed in table 3. gion: LuÈderitz District: Farm Plateau 38, near As might be expected, the arrangement of Aus, 26Њ40.62ЈS, 16Њ31.85ЈE, 1550 m, relationships among the species of Parabu- 30.xii.1997, L. Prendini and E. Scott. thus retrieved in the present analyses is al- DIAGNOSIS: Parabuthus muelleri falls in a most identical to that obtained previously group of species also including P. calvus, P. (Prendini, 2001a), as are the major ®ndings. capensis, P. pallidus, and P. planicauda. Monophyly of the genus Parabuthus is again This species is morphologically most easily supported, but monophyly of the disjunct confused with P. capensis, with which it southern African versus northeastern African shares the following combination of charac- and Arabian species is not. The optimal to- ters: metasomal segments I and II, stridula- pology presented here differs from that pub- tory region extended anteriorly beyond an- lished previously only in relationships terodorsal edge of segment, giving a steplike among the ®ve species comprising node ``B'' appearance in lateral aspect; metasomal seg- (®g. 1): P. calvus, P. capensis, P. muelleri, ments II±IV, dorsosubmedian carinae with P. pallidus, and P. planicauda. Previously, distal spiniform granules more pronounced the relationships among these species were than preceding granules; metasomal segment retrieved as follows by the majority of anal- IV, dorsosubmedian carinae medially discon- yses, including those deemed optimal: ((P. tinuous, median lateral carina continuous and capensis ϩ P. muelleri)(P. pallidus (P. cal- distinct; metasomal segment V, dorsosub- vus ϩ P. planicauda))). All previous analy- median carinae distinct with sharp, spiniform ses supported the (P. capensis ϩ P. muelleri) or subspiniform granules, and dorsolateral group. In contrast, all present analyses re- carinae distally obsolete. Parabuthus muel- trieved the following arrangement of these leri and P. capensis can be separated from species: (P. capensis (P. muelleri (P. calvus all other Parabuthus on the basis of the fol- ϩ P. pallidus ϩ P. planicauda))). Thus, lowing character: metasomal segment II, and whereas a sister-group relationship between to a lesser extent III, with posterodorsal edge P. capensis and P. muelleri was identi®ed by elevated and slightly curved forward medi- previous analyses, the addition of six char- ally, forming a subtriangular V-shape. acter states for the male of P. muelleri indi- Although morphologically similar, P. cates that this species actually shares a more muelleri can be separated from P. capensis recent common ancestor with P. calvus, P. by several characters. The movable ®nger of pallidus, and P. planicauda than with P. ca- the pedipalp chela (adult male and female) is pensis. The sister-group relationship between curved ventrally in P. muelleri, such that the P. calvus and P. planicauda, retrieved by proximal dentate margin is distinctly emar- most, but not all previous analyses, is no lon- ginate when the ®ngers are closed (i.e., a ger supported either. proximal ``gap'' is evident). The emarginate condition occurs in the male of several Par- SYSTEMATICS abuthus species (e.g., P. granulatus, P. ka- FAMILY BUTHIDAE C. L. KOCH, 1837 laharicus, and P. laevifrons), but it is uncom- GENUS PARABUTHUS POCOCK, 1890 mon in female Parabuthus, and does not oc- Parabuthus muelleri Prendini, 2000 cur in the male or female of P. capensis.In addition, P. muelleri has a more slender me- Parabuthus muelleri Prendini, 2000a: tasoma, in which the median width:length 32±38, percentage for metasomal segments I±V (n ®gs. 1±9, table 2. ϭ 3) is 80.5% (75±86%), 78% (75±81%), Parabuthus muelleri: Prendini, 2001a: 17; 2001b: 76% (72±80%), 69.5% (65±74%), and 55% 137. (53±57%), compared with the metasoma of AMERICAN MUSEUM NOVITATES Friday May 02 2003 03:12 PM novi 03331 Mp_6 Allen Press • DTPro System File # 01TQ

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P. capensis, in which the median width: ther in the description of sexual dimorphism, length percentage (n ϭ 6) is 86% (77±95%), below. 73% (78±88%), 92% (79±105%), 73% (67± DESCRIPTION: The following description of 79%), and 58% (53±63%). Metasomal seg- the male (AMNH [AH 3991]) supplements ment III is usually broader than segments I the previous descriptions (Prendini, 2000a) and II in P. capensis, but this is not the case of the holotype female (NMNW 1854) and in P. muelleri. Parabuthus muelleri is further paratype female (SAMC C4514). distinguished by the unusual shape of the tel- Color: Carapace, chelicerae, tergites, ster- son, which differs from all known Parabu- nites, and metasomal segments I±III: Cinna- thus species in the presence of a distal mon no. 123A. Metasomal segments IV, V ``bulge'' and a very short, sharply curved and telson: Burnt Sienna no. 132. Pedipalps aculeus. The median percentage of aculeus and legs: Clay Color no. 123B. Pectines: length:telson length in P. muelleri (n ϭ 3) is Chamois no. 123D. Metasomal segments IV, 30% (26±34%), compared with 39% (36± V and telson are distinctly darker than seg- 42%) in P. capensis (n ϭ 6). The two species ments I±III, whereas pedipalps and legs are may also usually be distinguished by the rel- distinctly paler than carapace, mesosoma, ative positions of the trichobothria on the metasoma, and telson (®gs. 2, 3). ®xed ®nger of the chela: eb and esb are lo- Carapace: Carapace with sulci, without cated proximal to the basal dentate margin of carinae, and covered entirely by uniform, the ®xed ®nger in P. muelleri, whereas eb is coarse granulation, becoming coarser on in- located proximal to the basal dentate margin, terocular and posterolateral surfaces. Anteri- and esb is located distal to it, in P. capensis. or margin of carapace procurved; posterior However, this character has been found to be margin straight. Five pairs of lateral ocelli. Median ocelli considerably larger than lateral polymorphic in P. capensis: in populations ocelli, situated anteromedially (®g. 4). Ocu- from the eastern part of the distributional lar tubercle with pair of smooth superciliary range, esb is also located proximal to the bas- carinae, protruding slightly above median al dentate margin (Prendini, 2000a). ocelli. Anteromedian furrow shallow; pos- Two characters of the newly described teromedian furrow shallow anteriorly, be- male can also be used to separate P. muelleri coming deeper posteriorly; posterolateral fur- from P. capensis, notably the lobate condi- rows shallow, wide, curved; posteromarginal tion of the pectinal proximal median lamella furrow narrow, deep. and pedipalp chelae that are not incrassate Chelicerae: Movable ®nger with distal ex- (i.e., not sexually dimorphic). The absence of ternal and distal internal teeth equal, oppos- sexual dimorphism of the pedipalp chelae is able. Ventral aspect of ®ngers and manus a particularly obvious diagnostic difference with long, dense macrosetae. Fixed ®nger between P. muelleri and P. capensis,in with a pair of denticles on the ventral sur- which the pedipalp chelae are markedly di- face. morphic (the pedipalp chela manus of the Sternum: Subtriangular (®g. 3). Median male is incrassate, whereas that of the female longitudinal furrow Y-shaped, shallow ante- is slender). Both characters are discussed fur- riorly, deep and narrow posteriorly.

← Fig. 1. The optimal tree obtained by analysis under weighting regimes that maximized ®t and min- imized length. This topology was retrieved by analyses with equal weights, successive weights, and implied weights under k ϭ 3±6 (table 2). Zero-length branches are collapsed. This topology also cor- responds to the majority rule (Ͼ50%) consensus of MPTs obtained by the eight analyses in which weighting regime and multistate character transformation were varied (table 2). Solid bars indicate uniquely derived apomorphic character states, whereas empty bars indicate parallel derivations of apomorphic states under ACCTRAN optimization. The number above each bar gives the character number, whereas the number below gives the character state. Branch-support values of nodes are provided below branches. Refer to appendix 2 for character descriptions. AMERICAN MUSEUM NOVITATES Friday May 02 2003 03:12 PM novi 03331 Mp_8 Allen Press • DTPro System File # 01TQ

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TABLE 3

Length (steps), Fit (fi), Consistency Indices (CIs), and Retention Indices (RIs) of 50 Informative Characters Scored Among 25 Species of the Genus Parabuthus Pocock, 1890 Final weights obtained with succesive weighting (SW) are also reported.

Pedipalps: Pedipalps covered in short ma- and a large proximal granule, ¯anked by an crosetae (®gs. 7±9). Femur ®nely and uni- inner and an outer accessory granule; chela formly granular; pentacarinate, all carinae dis- ®ngers each with a terminal denticle. tinct, granular, except for internomedian ca- Trichobothria: Orthobothriotaxic, type A, rina, comprising spiniform granules. Patella ␣ con®guration, with the following segment ®nely and uniformly granular; carinae absent totals (®gs. 7±12): femur, 11 (5 dorsal, 4 in- or obsolete; dorsointernal and ventrointernal ternal, 2 external), patella, 13 (5 dorsal, 1 carinae each comprising a row of granules internal, 7 external), and chela, 15 (8 manus, proximally; internomedian carina comprising 7 ®xed ®nger). Total number of trichobothria a large spiniform granule, proximally, and a per pedipalp, 39. Chela with eb located prox- few smaller granules, distally. Chela smooth; imal to basal dentate margin of ®xed ®nger carinae absent. Chela short, slender, length and esb located just distal; dt almost level along ventroexternal carina 28% greater than with et; db equidistant between est and esb.

chela width and 30% greater than chela Patella with esb2 slightly distal to esb1. Fe- height; length of movable ®nger 45% greater mur with d2 on proximointernal side of dor- than length along ventroexternal carina. Chela sointernal carina; d3 distal to d2; d4 equidis- ®xed ®nger slightly curved dorsally and mov- tant between d3 and d5. able ®nger slightly curved ventrally, such that Mesosoma: Pre-tergites smooth and shiny, proximal dentate margin emarginate when ®n- granular along posterior margins. Post-ter- gers are closed (®g. 7). Dentate margins of gites entirely coarsely granular, granulation chela ®ngers each with 11 oblique granular becoming coarser distally (®g. 2); I±VII each rows, each comprising 4±6 small granules with a weakly developed, granular median AMERICAN MUSEUM NOVITATES Friday May 02 2003 03:12 PM novi 03331 Mp_9 Allen Press • DTPro System File # 01TQ

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Figs. 2, 3. Parabuthus muelleri Prendini, 2000, male (AMNH [AH 3991]), habitus. 2. Dorsal aspect. 3. Ventral aspect. Scale bar ϭ 5 mm. AMERICAN MUSEUM NOVITATES Friday May 02 2003 03:12 PM novi 03331 Mp_10 Allen Press • DTPro System File # 01TQ

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carina; VII additionally with distinct pairs of segment III narrow, virtually obsolete, con- costate granular dorsosubmedian and dorso- sisting of a few granules in the proximal lateral carinae, and with well-developed third of the segment; segment II, and to a stridulatory region between dorsosubmedian lesser extent III, with posterodorsal edge el- carinae, consisting of round to slightly cres- evated and slightly curved forward medially, cent-shaped granules reaching the posterior forming a subtriangular V-shape (®g. 5). Me- margin. Sternites entirely smooth, except for tasoma densely covered with long macrose- posterolateral surfaces of sternite VII, which tae, especially on the ventral surface of the are sparsely granular; lateral and distal mar- telson. Metasomal segments I±IV each with gins each with a row of sparsely distributed 10 carinae, but segment IV with ventrosub- macrosetae; sternite VII with weakly devel- median and median lateral carinae distinct oped pairs of costate ventrosubmedian and only in the proximal half of the segment; ventrolateral carinae. segment V with seven carinae, including a Pectines: First proximal median lamella of single, obsolete granular ventromedian cari- each pecten suboval, mesially enlarged and na, a pair of distinct ventrolateral carinae, a lobate (®g. 3). Pectinal teeth: 36/36. pair of dorsolateral carinae, distinct only in Genital operculum: Completely divided the proximal half of the segment, and a pair longitudinally. Genital papillae present. of dorsosubmedian carinae reduced to a few Legs: Tibia III and IV with spurs; retro- prominent rounded or subspiniform granules lateral margins with scattered macrosetae. medially. Metasomal segments I±V with dor- Basitarsi I and II only slightly compressed sosubmedian carinae converging distally in dorsoventrally, retrolateral margins each with segment I, subparallel in segments II±V; ven- dense row of long, ®ne macrosetae. Telotarsi trolateral carinae converging distally in seg- each with paired ventrosubmedian rows of ments I±III, subparallel in segment IV, di- ®ne macrosetae. Telotarsal laterodistal lobes verging in segment V. All metasomal carinae truncated; median dorsal lobes extending to costate granular to granular, except for ven- ungues. Telotarsal ungues short, distinctly trosubmedian and ventrolateral carinae of curved, and equal in length. segment I, which are costate to costate gran- Metasoma and telson: Metasomal seg- ular. Metasomal segments II±IV with distal ments I±V width/length ratio progressively granules of dorsosubmedian carinae very decreasing, width percentage of length 86% slightly enlarged, rounded; segments II and for I, 81% for II, 80% for III, 68% for IV, III with distal granules of ventrosubmedian and 53% for V (table 4). Telson oval, globose carinae and, to much a lesser extent, ventro- (®g. 6), height 66% of length, with ¯attened lateral carinae, enlarged, obtuse, and elevat- dorsal surface and rounded ventral surface; ed; segment V with subdistal granules of vesicle not signi®cantly narrower than me- ventrolateral carinae enlarged into laterally tasomal segment V, width 90% of metasomal compressed, lobate processes. Telson with a segment V. Metasoma entirely granular, ex- distal ``bulge'' and a very short, sharply cept for ventromedian surfaces of segments curved aculeus (®g. 6); aculeus length per- I and II, and dorsomedian surfaces of seg- centage of vesicle length, 35%. ments IV, V and telson. Metasomal segments Hemispermatophore: A hemispermato- I and II each with a well-developed stridu- phore was dissected from the male, con®rm- latory region on the dorsomedian surface, ing that it is adult. The hemispermatophore consisting of round to slightly crescent- is typical of other species of Parabuthus, shaped ®ne granules extending to the poste- characteristically ¯agelliform, with pars rec- rior margin (®g. 5); stridulatory region of ta parallel to axis of distal lamina (®g. 13).

← Figs. 4±6. Parabuthus muelleri Prendini, 2000, male (AMNH [AH 3991]), diagnostic characters. 4. Carapace. 5. Dorsal aspect of segments I and II, showing dorsomedian stridulatory region and subtriangular V-shape curvature at posterodorsal edge. 6. Lateral aspect of segments IV, V and telson. Scale bars ϭ 1 mm. AMERICAN MUSEUM NOVITATES Friday May 02 2003 03:12 PM novi 03331 Mp_12 Allen Press • DTPro System File # 01TQ

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Figs. 7±9. Parabuthus muelleri Prendini, 2000, male (AMNH [AH 3991]), distribution of trichobothria and macrosetae on the dextral pedipalpal chela. 7. Dorsal aspect. 8. Ventral aspect. 9. Internal aspect. Scale bar ϭ 1 mm.

Sexual dimorphism: Unlike most species 1977; Lamoral, 1977, 1979, 1980; Newlands of Parabuthus, in which the pedipalp chela and Martindale, 1980; Prendini, 2001a), the manus of the adult male is noticeably incras- adult male of P. muelleri is not sexually di- sate (bulbous or swollen), compared with the morphic in this respect. The chela width: more slender manus of the adult female (Po- length and height:length ratios of 72% and cock, 1889, 1890, 1902; Kraepelin, 1899, 70%, respectively, in the male do not differ 1908; Purcell, 1898, 1899, 1901; Werner, signi®cantly from the equivalent ratios of 1916; Hewitt, 1913, 1915, 1918; Eastwood, 74% and 69%, respectively, in the female. AMERICAN MUSEUM NOVITATES Friday May 02 2003 03:12 PM novi 03331 Mp_13 Allen Press • DTPro System File # 01TQ

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Figs. 10±12. Parabuthus muelleri Prendini, 2000, male (AMNH [AH 3991]), distribution of trichobothria and macrosetae on the dextral pedipalpal patella and femur. 10. Dorsal aspect of patella. 11. External aspect of patella. 12. Dorsal aspect of femur. Scale bars ϭ 1 mm.

Parabuthus calvus, P. nanus, P. pallidus, imal median lamellae of the pectines, a char- and P. planicauda are the only other species acter observed in the female of most Para- of the genus in which the pedipalp chelae of buthus speciesÐP. granulatus and P. kala- the adult male are not sexually dimorphic. haricus are exceptionsÐbut otherwise only The absence of sexual dimorphism of the in the male of P. calvus and P. planicauda pedipalp chelae provides an additional ob- (Purcell, 1898, 1901; Pocock, 1902; Krae- vious diagnostic character for separating P. pelin, 1908; Hewitt, 1918; Werner, 1934; muelleri from the closely related and mor- Lawrence, 1955; Eastwood, 1977; Lamoral, phologically similar P. capensis. The male of 1979; Sissom, 1994; Prendini, 2001a). In P. muelleri also displays weakly lobate prox- other respects, the differences between the AMERICAN MUSEUM NOVITATES Friday May 02 2003 03:12 PM novi 03331 Mp_14 Allen Press • DTPro System File # 01TQ

14 AMERICAN MUSEUM NOVITATES NO. 3408

TABLE 4 Meristic Data for Holotype Female (NMNW 1854), Paratype Female (SAMC C4514), and Newly Discovered Male (AMNH [AH 3991]) of Parabuthus muelleri Prendini, 2000 Male Parabuthus capensis (Ehrenberg, 1831) from Vanrhynsdorp, Western Cape Province, South Africa (SAMC C4565) included for comparison. Measurements follow Stahnke (1970), Lamoral (1979), and Prendini (2000a). AMERICAN MUSEUM NOVITATES Friday May 02 2003 03:12 PM novi 03331 Mp_15 Allen Press • DTPro System File # 01TQ

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Fig. 13. Parabuthus muelleri Prendini, 2000, male (AMNH [AH 3991]), hemispermatophore, ental aspect. Scale bar ϭ 1 mm.

male and female of P. muelleri are typical of pacted, chalky soil, calcrete nodes, and do- other species of Parabuthus. The male ex- lomite rocks. The holotype was excavated hibits genital papillae, a greater pectinal from the burrow of a scorpionid, Opisto- tooth count, increased granulation and seta- phthalmus opinatus (Simon, 1888), which it tion (particularly of the pedipalps, metasoma, had preyed on, and was also syntopic with and telson), and is proportionally more slen- O. scabrifrons Hewitt, 1918. The paratype der than the female. was found sitting motionless on a stone at DISTRIBUTION: Parabuthus muelleri is en- night and was syntopic with the ischnurid demic to Namibia (®g. 14) and presently re- Hadogenes tityrus (Simon, 1888), the scor- corded from two neighboring localities in the pionids Opistophthalmus gigas Purcell, 1898 LuÈderitz District (Karas Region) and a third and O. scabrifrons, and also with Parabuthus locality, ca. 260 km north, in the MaltahoÈhe granulatus and P. villosus. According to A. District (Hardap Region). Despite the large Harington's collecting notes, the newly de- distance between the ®rst two localities and scribed male specimen was collected in an the third, the habitat in which the specimens area with P. villosus and P. laevifrons, with were collected (at the two localities for the latter probably being a misidenti®cation which habitat data are available) was re- of P. stridulus. markably similar (Prendini, 2000a). The ADDITIONAL MATERIAL EXAMINED: ( specimens were each found in an area of lev- (AMNH [AH 3991]), Namibia: Karas Region: el ground, with chalky soil, calcrete nodes, LuÈderitz District: locality uncertain, probably and dolomitic rocks. The localities at which Aus [26Њ41ЈS, 16Њ15ЈE], A. Harington. Unfor- they were collected both occur along an ex- tunately, the collection locality data for this tensive ridge of dolomite, extending from the specimen are ambiguous. Harington suggests Huib-Hoch Plateau in the south (Plateau is two possible localities, the other being Wor- situated at the northern end) to the Naukluft tel (Khomas Region: Windhoek District: mountains in the north (Onis is ca. 20 km 23Њ08ЈS, 17Њ10ЈE). Aus is clearly the more south of the Naukluft). Thus, the two local- plausible of the two alternatives, given its ities are fairly close to the northern and close proximity to Plateau, the collection lo- southern edges of this geological formation. cality of the paratype female. Indeed, it is It is not known whether P. muelleri is re- quite possible that the male specimen also stricted to this formation, but the occurrence originated from Plateau, where Harington of two of the three known specimens in such collected extensively and also received ma- similar, albeit distantly located habitats sug- terial donated by the owners of the property, gests that the species may occur in similar H. and W. Erni. The possibility that the male habitats throughout the intervening areas specimen was collected at Plateau also seems (Prendini, 2000a). more likely in view of the considerably dif- ECOLOGY: The holotype and paratype of P. ferent habitats at Aus and Plateau, which muelleri were collected in a region of com- have few scorpion species in common, de- AMERICAN MUSEUM NOVITATES Friday May 02 2003 03:12 PM novi 03331 Mp_16 Allen Press • DTPro System File # 01TQ

16 AMERICAN MUSEUM NOVITATES NO. 3408

Fig. 14. The known distribution of Parabuthus muelleri Prendini, 2000 (Ⅵ), which is endemic to Namibia. Contour interval ϭ 600 m.

spite their proximity. The Aus Mountains are Parabuthus data matrix have already been part of a granitic formation completely sep- acknowledged in my previous papers, but I arated from the dolomitic Huib-Hoch Plateau thank them all again here. I also reiterate my by a sandy plain approximately 10 km across appreciation to Jack Harington, Lucian Har- at the narrowest point. No scorpions associ- ington, and Eone de Wet for transferring ated with rocky habitats in either the Aus Alexis Harington's scorpion collection to the Mountains or the Huib-Hoch Plateau have AMNH, to the Scott family for accommo- been collected on this plain, which probably dation, assistance, and congenial company represents a signi®cant barrier to their dis- during the sorting and packing of the collec- persal, as has been demonstrated elsewhere tion in Johannesburg, and to Randall T. for lithophilous and lapidicolous scorpion Schuh for expediting the ®nancial aspects of species (Newlands, 1972; Prendini, 2001b). bringing the collection to New York. This is the fourth paper incorporating material from ACKNOWLEDGMENTS Alexis' tremendous collection and it certain- The people and institutions that assisted in ly is not the last. I thank Roy Larimer for the acquisition of the holotype and paratype assistance with the photography in this paper, of P. muelleri and in the compilation of the Steve Thurston for preparing the photograph- AMERICAN MUSEUM NOVITATES Friday May 02 2003 03:12 PM novi 03331 Mp_17 Allen Press • DTPro System File # 01TQ

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ic plates, Randall T. Schuh, Victor Fet, and (Arachnida: Scorpionida). Transactions of the an anonymous reviewer for commenting on Zimbabwe Scienti®c Association 68: 7±14. an earlier draft of the manuscript, and Lee Goloboff, P.A. 1991. Homoplasy and the choice Herman for assistance with the editorial pro- among cladograms. Cladistics 7: 215±232. cess at the AMNH. Goloboff, P.A. 1993. Estimating character weights during tree search. Cladistics 9: 83±91. Goloboff, P.A. 1995. Parsimony and weighting: a REFERENCES reply to Turner and Zandee. Cladistics 11: 91± Balinsky, B.I. 1962. Patterns of animal distribu- 104. tion on the African continent. Annals of the Goloboff, P.A. 1997a. NONA, version 2.0. Com- Cape Provincial Museums 2: 299±309. puter software and documentation. New York: Birula, A.A. 1915. Arachnologische BeitraÈge. VI. American Museum of Natural History. UÈ ber die nordostafrikanischen Formen von Goloboff, P.A. 1997b. Pee-Wee, version 2.6. Parabuthus liosoma (Hemp. et Ehr.). Revue Computer software and documentation. New Russe d'Entomologie 15: 131±146. York: American Museum of Natural History. Bremer, K. 1988. The limits of amino acid se- Griswold, C.E., J.A. Coddington, G. Hormiga, quence data in angiosperm phylogenetic recon- and N. Scharff. 1998. Phylogeny of the orb- struction. Evolution 42: 795±803. web building spiders (Araneae, Orbiculariae: Bremer, K. 1994. Branch support and tree stabil- Deinopoidea, Araneoidea). Zoological Journal ity. Cladistics 10: 295±304. of the Linnean Society 123: 1±99. Bryant, H.N. 1995. Why autapomorphies should Hewitt, J. 1913. The Percy Sladen Memorial Ex- be removed: a reply to Yeates. Cladistics 11: pedition to Great Namaqualand, 1912±1913. 381±384. Records and descriptions of the Arachnida of Couzijn, H.W.C. 1976. Functional anatomy of the the collection. Annals of the Transvaal Museum walking-legs of Scorpionida with remarks on 4: 146±159. terminology and homologization of leg seg- Hewitt, J. 1915. New South African Arachnida. ments. Netherlands Journal of Zoology 26: Annals of the Natal Museum 3: 289±327. 453±501. Hewitt, J. 1918. A survey of the scorpion fauna Donoghue, M.J., R.G. Olmstead, J.F. Smith, and of South Africa. Transactions of the Royal So- J.D. Palmer. 1992. Phylogenetic relationships ciety of South Africa 6: 89±192. of Dipsacales based on rbcL sequence data. Hjelle, J.T. 1990. Anatomy and morphology. In Annals of the Missouri Botanical Garden 79: G.A. Polis (editor), The biology of scorpions: 333±345. 9±63. Stanford, CA: Stanford University Press. Eastwood, E.B. 1977. Notes on the scorpion fauna KovarÏõÂk, F. 2001. Catalog of the scorpions of the of the Cape. Part 2. The Parabuthus capensis world (1758±1998) by V. Fet, W.D. Sissom, G. (Ehrenberg) species-group; remarks on taxon- Lowe, and M. Braunwalder (New York Ento- omy and bionomics (Arachnida, Scorpionida, mological Society, 2000: 690 pp.). Discussion Buthidae). Annals of the South African Muse- and supplement for 1999 and part of 2000. Ser- um 73: 199±214. ket 7: 78±93. Farris, J.S. 1969. A successive approximations ap- KovarÏõÂk, F. 2002. Co noveÂhousÏtõÂruÊ v roce 2000. proach to character weighting. Systematic Zo- AkvaÂrium TeraÂrium 45: 55±61. [in Czech] ology 18: 374±385. Kraepelin, K. 1899. Scorpiones und Pedipalpi. In Farris, J.S. 1970. Methods for computing Wagner F. Dahl (editor), Das Tierreich 8: 1±265. Berlin: trees. Systematic Zoology 19: 83±92. R. FriedlaÈnder und Sohn Verlag. Farris, J.S. 1982. Outgroups and parsimony. Sys- Kraepelin, K. 1908. Skorpione und Solifugen. In tematic Zoology 31: 328±334. L.G. Schultze (editor), Forschungsreise im Fet, V., and G. Lowe. 2000. Family Buthidae C. Westlichen und Zentralen SuÈdafrika, Ausge- L. Koch, 1837. In V. Fet, W.D. Sissom, G. fuÈhrt in den Jahren 1903±1905 1: 247±282. Lowe, and M.E. Braunwalder (editors), Catalog Jena: Fischer. of the scorpions of the world (1758±1998): 55± Lamoral, B.H. 1977. Parabuthus kalaharicus,a 286. New York: New York Entomological So- new species of scorpion from the Kalahari ciety. Gemsbok National Park in the Republic of Fitch, W.M. 1971. Toward de®ning the course of South Africa (Buthidae, Scorpionida). Koedoe evolution: minimum change for a speci®c tree 20: 101±107. topology. Systematic Zoology 20: 406±416. Lamoral, B.H. 1979. The scorpions of Namibia FitzPatrick, M.J. 1994. A checklist of the Para- (Arachnida: Scorpionida). Annals of the Natal buthus Pocock species of Zimbabwe with a re- Museum 23: 498±783. description of P. mossambicensis (Peters, 1861) Lamoral, B.H. 1980. Two new psammophile spe- AMERICAN MUSEUM NOVITATES Friday May 02 2003 03:12 PM novi 03331 Mp_18 Allen Press • DTPro System File # 01TQ

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cies and new records of scorpions from the Prendini, L. 2000a. A new species of Parabuthus northern Cape Province of South Africa Pocock (Scorpiones: Buthidae), and new re- (Arachnida: Scorpionida). Annals of the Natal cords of Parabuthus capensis (Ehrenberg), Museum 24: 201±210. from Namibia and South Africa. Cimbebasia Lawrence, R.F. 1955. Solifugae, scorpions and Pe- 16: 201±214. dipalpi, with checklists and keys to South Af- Prendini, L. 2000b. Phylogeny and classi®cation rican families, genera and species. Results of of the superfamily Scorpionoidea Latreille the Lund University Expedition in 1950±1951. 1802 (Chelicerata, Scorpiones): an exemplar In B. HanstroÈm, P. Brinck, and G. Rudebeck approach. Cladistics 16: 1±78. (editors), South African animal life 1: 152±262. Prendini, L. 2001a. Phylogeny of Parabuthus Uppsala: Almqvist and Wiksells. (Scorpiones, Buthidae). Zoologica Scripta 30: Maddison, D. 1991. The discovery and impor- 13±35. tance of multiple islands of most parsimonious Prendini, L. 2001b. Substratum specialization and trees. Systematic Zoology 40: 315±328. speciation in southern African scorpions: the Newlands, G. 1972. A description of Hadogenes Effect Hypothesis revisited. In V. Fet and P.A. lawrencei sp. nov. (Scorpiones) with a checklist Selden (editors), Scorpions 2001. In Memoriam and key to the South West African species of Gary A. Polis: 113±138. Burnham Beeches, the genus Hadogenes. Madoqua (II) 1: 133± UK: British Arachnological Society. 140. Probst, P.J. 1973. A review of the scorpions of Newlands, G., and C.B. Martindale. 1980. The East Africa with special regard to Kenya and buthid scorpion fauna of Zimbabwe-Rhodesia Tanzania. Acta Tropica 30: 312±335. with checklists and keys to the genera and spe- Purcell, W.F. 1898. Descriptions of new South Af- cies, distributions and medical importance rican scorpions in the collection of the South (Arachnida: Scorpiones). Zeitschrift fuÈr Ange- African Museum. Annals of the South African wandte Zoologie 67: 51±77. Museum 1: 1±32. Nixon, K.C. 1999. WinClada, version 0.9.9ϩ. Purcell, W.F. 1899. New South African scorpions Computer software and documentation. Avail- in the collection of the South African Museum. able at: http://www.cladistics.com. Annals of the South African Museum 1: 433± Nixon, K.C., and J.M. Carpenter. 1993. On out- 438. groups. Cladistics 9: 413±426. Purcell, W.F. 1901. On some South African Pocock, R.I. 1889. Notes on some Buthidae, new Arachnida belonging to the orders Scorpiones, and old. Annals and Magazine of Natural His- Pedipalpi, and Solifugae. Annals of the South tory, ser. 6, 3: 334±351. African Museum 2: 137±225. Pocock, R.I. 1890. A revision of the genera of Sissom, W.D. 1990. Systematics, biogeography scorpions of the family Buthidae, with de- and paleontology In G.A. Polis (editor), The scriptions of some South-African species. Pro- biology of scorpions: 64±160. Stanford, CA: ceedings of the Zoological Society 1890: 114± Stanford University Press. 141. Sissom, W.D. 1994. Descriptions of new and Pocock, R.I. 1895. On the Arachnida and Myri- poorly known scorpions of Yemen (Scorpiones: opoda obtained by Dr. Anderson's collector Buthidae, Diplocentridae, Scorpionidae). Fauna during Mr. T. Bent's expedition to the Hadra- of Saudi Arabia 14: 3±39. maut, South Arabia, with a supplement upon Smithe, F.B. 1974. Naturalist's color guide sup- the scorpions obtained by Dr. Anderson in plement. New York: American Museum of Nat- Egypt and the Eastern Soudan. Journal of the ural History, xiiiϩ229 pp. Linnean Society 25: 292±316. Smithe, F.B. 1975. Naturalist's color guide. New Pocock, R.I. 1899. Solifugae, Scorpiones, Chilop- York: American Museum of Natural History, oda and Diplopoda (Appendix C). In A. Don- unpaginated. aldson Smith, Through unknown African coun- Smithe, F.B. 1981. Naturalist's color guide. Part tries. The First Expedition from Somaliland to III. New York: American Museum of Natural Lake Lamu: 392±407. London: Edward Ar- History, 37 pp. nold. Stahnke, H.L. 1970. Scorpion nomenclature and Pocock, R.I. 1902. A contribution to the system- mensuration. Entomological News 81: 297± atics of scorpions. I. Some corrections in no- 316. menclature. II. Notes on some species of Par- Swofford, D.L., and W.P. Maddison. 1987. Re- abuthus contained in the British Museum. III. constructing ancestral character states under Descriptions of some new and old species. An- Wagner parsimony. Mathematical Biosciences nals and Magazine of Natural History, ser. 7, 87: 199±229. 10: 364±380. Swofford, D.L., and W.P. Maddison. 1992. Parsi- AMERICAN MUSEUM NOVITATES Friday May 02 2003 03:12 PM novi 03331 Mp_19 Allen Press • DTPro System File # 01TQ

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mony, character-state reconstructions, and evo- Watrous, L.E., and Q.D. Wheeler. 1981. The out- lutionary inferences. In R.L. Mayden (editor), group comparison method of character analysis. Systematics, historical ecology, and North Systematic Zoology 30: 1±11. American freshwater ®shes: 186±283. Palo Werner, F. 1916. UÈ ber einige Skorpione und Alto, CA: Stanford University Press. Gliederspinnen des Naturhistorischen Museums Vachon, M. ``1973'' [1974]. EÂ tude des caracteÁres in Wiesbaden. JahrbuÈcher des Nassauischen utiliseÂs pour classer les familles et les genres Vereins fuÈr Naturkunde 69: 79±97. de scorpions (Arachnides). 1. La trichobothri- Werner, F. 1934. Scorpiones, Pedipalpi. In H.G. otaxie en arachnologie. Sigles trichobothriaux Bronn (editor), Klassen und Ordnungen des et types de trichobothriotaxie chez les scorpi- Tierreichs 5, IV, 8, Lief. 1±2, Scorpiones, 1± ons. Bulletin du MuseÂum National d'Histoire 316. Leipzig: Akademische Verlaggesells- Naturelle (Paris), ser. 3, 140: 857±958. chaft. Vachon, M. 1979. Arachnids of Saudi Arabia. Scorpiones. Fauna of Saudi Arabia 1: 30±66.

APPENDIX 1

TAXA EXAMINED FOR CLADISTIC ANALYSIS OF juv &, same data (BMNH 1893.1.11.48±54); 2 &, PARABUTHUS POCOCK, 1890 SOMALIA, Somali Coast, Berbera, H.M. Phipson (BMNH 1895.6.1.45). This species, with two sub- Depositories for specimens examined are ab- species (one nominotypical), has not been revised breviated as follows: AMNH, American Museum since the original description, and the status of the of Natural History (New York, NY); AH, Alexis subspecies, Parabuthus granimanus fuscicauda Harington Collection (lodged at AMNH); BMNH, Caporiacco, 1947, is unknown. The two speci- The Natural History Museum (London, UK); mens referred to by Pocock (1895: 312) as ``types NMSA, Natal Museum (Pietermaritzburg, South of ( and &'' are hereby designated as lectotype Africa); NMNW, National Museum of Namibia and paralectotype. Based on a comparison of Po- (Windhoek, Namibia); SAMC, South African Mu- cock's type specimens, the nominotypical subspe- seum (Cape Town, South Africa); TMSA, Trans- cies is suspected to be conspeci®c with P. hunteri. vaal Museum (Pretoria, South Africa); USNM, The types of P. granimanus and P. hunteri share US National Museum of Natural History, Smith- the apomorphic state of characters 7 and 12 and sonian Institution (Washington, DC). Taxonomic differ only in the darker coloration, and reduced notes are provided and lectotypes designated for granulation and setation of the metasoma, of P. the northeastern African and Arabian species of granimanus. The two specimens of P. granimanus which type material was examined. from ``Somali Coast'' (also discussed in Pocock's description) are indistinguishable from the types OUTGROUPS of Parabuthus hunteri Pocock, 1895. Nonetheless, 1. Grosphus madagascariensis (Gervais, 1843): I have refrained from providing a formal synon- 10 (,3&, MADAGASCAR, Nosy Lava ymy, pending a more detailed revision of the Par- (AMNH). abuthus species from northeastern Africa and 2. Uroplectes triangulifer (Thorell, 1876): (, Arabia. &, SOUTH AFRICA, Northern Prov., Potgieters- 4. Parabuthus heterurus Pocock, 1899: Lecto- rus, 24Њ11ЈS, 29Њ01ЈE, iv.1934, R.F. Lawrence type &, paralectotype (, SOMALIA, Schebegh (SAMC B8227); (, SOUTH AFRICA, Eastern River [Shebeli River], Dr A. Donaldson-Smith Cape Prov., Karoo Nature Reserve, Graaff-Reinet, (BMNH 1897.11.10.9±10); &, SOMALIA, Bur- 32Њ12ЈS, 24Њ28ЈE, 8±9.ix.1987, S. van Noort ao, 2.v.1938, E.F. Peck (BMNH 1939.3.16.17± (SAMC C3754); 2 &, SOUTH AFRICA, Eastern 18); (, SOMALIA, Burao, 8.v.1947, W.A. Mac- Cape Prov., Alicedale, 25.i.1979, A. Harington fadyeu (BMNH 1949.12.19.5). This species, with (AMNH); 2 &, SOUTH AFRICA, North West two subspecies (one nominotypical), has not been Prov., Lichtenburg, 27.vii.1975, A. Harington revised since its description but can be separated (AMNH). from the closely related P. granimanus and P. hunteri on the basis of characters 7, 12, and 46. The taxonomic status of the subspecies, Parabu- NORTHEASTERN AFRICAN AND ARABIAN PARABUTHUS thus heterurus stefaninii Caporiacco, 1927, is 3. Parabuthus granimanus Pocock, 1895: Lec- presently unknown. Pocock's (1899) two syntypes totype (, paralectotype &, SOMALIA, Zaila were examined and are hereby designated as lec- [Zeyla], E.W. Oates (BMNH 1893.1.11.48±54); totype and paralectotype. AMERICAN MUSEUM NOVITATES Friday May 02 2003 03:12 PM novi 03331 Mp_20 Allen Press • DTPro System File # 01TQ

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5. Parabuthus hunteri Pocock, 1895: Lectotype 1201); &, SOUTH AFRICA, Western Cape Prov., &, paralectotype (, SUDAN, Duroor, 60 mi N of Betjesfontein [Biesjesfontein], 31Њ10ЈS, 17Њ53ЈE, Suakin, D. Anderson (BMNH 1894.11.2.52±60); 1898, M. Bergh (SAMC 2228); &, SOUTH AF- paralectotype (, same data (BMNH 1894.11.2.41± RICA, Western Cape Prov., Knersvlakte, N of 50); paralectotype (, SUDAN, Suakin, 7.ix1892, Vanrhynsdorp, 31Њ37ЈS, 18Њ44ЈE, 1999, M. de Ja- Dr B. Penton to D. Anderson (BMNH ger (SAMC C4615); &, SOUTH AFRICA, North- 1892.12.22.1). This species, which has not been ern Cape Prov., Paulshoek, E of Garies, 30Њ22ЈS revised since the original description, is suspected 18Њ16ЈE, i.1997, S. Todd (AMNH). to be conspeci®c with the nominotypical subspe- 10. Parabuthus capensis (Ehrenberg, 1831): (, cies of P. granimanus (see above). One of Po- NAMIBIA, Karas Region, Boomrivier [Fish River cock's (1895) 38 syntypes has been selected as Canyon National Park], 28Њ01ЈS, 17Њ04ЈE, 13± the lectotype of P. hunteri. The remaining speci- 26.ix.1992, E. Marais (NMNW 1509); 5 (,2&, mens, referred to in Pocock's (1895) description, 2 juv &, SOUTH AFRICA, Eastern Cape Prov., are hereby designated as paralectotypes. Graaff-Reinet and Kruidfontein, 8 mi from Graaff 6. Parabuthus leiosoma (Ehrenberg, 1828): (, Reinet, 32Њ22ЈS, 24Њ36ЈE, ix.1902, J. Paynter &, KENYA, Sabuk Retreat, Rift Valley, 2.vi.1999, (SAMC 12010); &, juv (, SOUTH AFRICA, H. Herren (AMNH); subadult &, 3 juv (, 6 juv Western Cape Prov., Table View, Cape Town, &, KENYA, Nguruman, Rift Valley, vi±vii.1998, 33Њ49ЈS, 18Њ29ЈE, 29.iv.1986, Louw (SAMC J. Lazell (USNM); (, &, juv (, SOMALIA C1618); 2 (,3&, 4 subadult (, 2 subadult &,3 (SAMC 409.60); (, &, SOMALIA, 1899, ex juv (, 2 juv &, SOUTH AFRICA, Western Cape Hamburg Mus. (SAMC 4060); &, juv (, juv &, Prov., Port Nolloth, 29Њ17ЈS, 16Њ51ЈE, iv.1972, J. YEMEN, Aden, 1899, Oates and Shopland Visser (NMSA 10358); (, 2 subadult &, subadult (SAMC 6343). Three subspecies (one nominotyp- ( [black form], SOUTH AFRICA, Western Cape ical) are currently recognized, of which P. leio- Prov., Laaiplek, 32Њ46ЈS, 18Њ10ЈE, ix.1976, G. soma leiosoma was reviewed by Vachon (1979) McLachlin (SAMC C74). and subsequently redescribed by Sissom (1994). 11. Parabuthus distridor Lamoral, 1980: Para- The taxonomic validity of Parabuthus leiosoma types 12 (,2&, juv (, juv &, SOUTH AFRICA, abyssinicus Pocock, 1901 and P. leiosoma dmi- Northern Cape Prov., Richtersveld, sandy ridge, 8 trievi Birula, 1903, both from Ethiopia, is dubi- ous. Neither has been reviewed since Birula km S Springklipberg, 28Њ40ЈS, 16Њ53ЈE, (1915). Probst (1973) neglected to mention sub- 21.ii.1979, B. Lamoral (NMSA 11305); paratype species in his brief treatment of P. leiosoma from (, same data (NMSA 11436); paratype &, same East Africa. data (NMSA 11435); 2 (, &, SOUTH AFRICA, 7. Parabuthus pallidus Pocock, 1895: Lecto- Northern Cape Prov., 25 km E of Port Nolloth at type (, paralectotype &, KENYA, Mombasa, Mr turnoff to Wolfberg, iii.1997, L. Prendini and E. Last, purchased of H. Grose-Smith (BMNH Scott (SAMC C4604). 1890.3.15.10±11); 2 &, KENYA, N Turkana, 12. Parabuthus gracilis Lamoral, 1979: Para- Lake Rudolf Rift Valley Exped., 1934, N. Fuchs types 5 (,2&, 5 juv, NAMIBIA, Erongo Region, (BMNH); 3 &, KENYA, Sabuk Retreat, Rift Val- Messum Crater area, 21Њ16ЈS, 14Њ13ЈE, ley, 2.vi.1999, H. Herren (AMNH). This species 26.iii.1976, B. Lamoral (NMSA 10848); para- was brie¯y reviewed by Probst (1973). Pocock's types 3 (, &, 4 juv (, NAMIBIA, Kunene Re- (1895) two syntypes were examined and are here- gion, MoÈwebaai, 4 km N, 19Њ19ЈS, 12Њ40ЈE, by designated as the lectotype and paralectotype. 29.iii.1976, B. Lamoral and L. Ferguson (NMSA 10859). 13. Parabuthus granulatus (Ehrenberg, 1831): SOUTHERN AFRICAN PARABUTHUS 9 (,2&, juv (, juv &, NAMIBIA, Karas Region, 8. Parabuthus brevimanus (Thorell, 1876): 20 Berseba, 10 km S, 26Њ07ЈS, 17Њ46ЈE, 27.ii.1976, (,5&, 2 subadult (, 2 juv (, 8 juv &, NAMIB- B. Lamoral (NMSA 10731); 4 (,2&, 3 subadult IA, Erongo Region, Farm Kranzberg 59, 21Њ58ЈS, &, 9 juv, NAMIBIA, Kunene Region, Farm Vrede 15Њ39ЈE, 23.iii.1976, B. Lamoral and L. Ferguson 719, 20Њ23ЈS, 14Њ14ЈE, 31.iii.1976, B. Lamoral (NMSA 10819); 3 (, &, juv &, SOUTH AFRI- and L. Ferguson (NMSA 10836). CA, Northern Cape Prov., Richtersveld, Spring- 14. Parabuthus kalaharicus Lamoral, 1977: bokvlakte, 28Њ23ЈS, 17Њ04ЈE, 20±21.ii.1973, B. Paratypes 9 (, NAMIBIA, Karas Region, Farm Lamoral (NMSA 10442). Sterkstroom 320, 25Њ43ЈS, 19Њ19ЈE, 19.iii.1969, 9. Parabuthus calvus Purcell, 1898: Holotype B. Lamoral (NMSA 10947); paratypes 2 &, (, SOUTH AFRICA, Northern Cape Prov., On- SOUTH AFRICA, Northern Cape Prov., Kalahari der Bokkeveld, Bokkeveld Mountains, 31Њ20ЈS, Gemsbok National Park, Twee Rivieren, 26Њ30ЈS, 19Њ04ЈE, Calvinia, 1897, M. Schlechter (SAMC 20Њ35ЈE, iii.1970, B. Lamoral (NMSA 10946); AMERICAN MUSEUM NOVITATES Friday May 02 2003 03:12 PM novi 03331 Mp_21 Allen Press • DTPro System File # 01TQ

2003 PRENDINI: PARABUTHUS MUELLERI 21

paratypes 3 ( (SAMC C213), Twee Rivieren, 21. Parabuthus nanus Lamoral, 1979: Para- 1960±1970, le Riche family and staff. types (, &, NAMIBIA, Karas Region, Farm Tsi- 15. Parabuthus kraepelini Werner, 1902: &, rub 13, 26Њ52ЈS, 16Њ02ЈE, 3.iii.1976, B. Lamoral NAMIBIA, Kunene Region, Etosha National (NMSA 10772); paratype (, SOUTH AFRICA, Park, Aus, 2±6.iii.1969, B. Lamoral and R. Day Northern Cape Prov., Goodhouse, 21 km S, (NMSA 10022); 2 &, NAMIBIA, Kunene Region, 29Њ04ЈS, 18Њ06ЈE, 29±31.i.1973, B. Lamoral Etosha National Park, Gemsbokvlakte, 4.iii.1969, (NMSA 10703); paratypes (, &, SOUTH AFRI- B. Lamoral and R. Day (NMSA 10019); subadult CA, Northern Cape Prov., Goodhouse, 10 km S, (, NAMIBIA, Hardap Region, Farm Kangas 371, 28Њ55ЈS, 18Њ14ЈE, 11.ii.1979, B. Lamoral (NMSA 23Њ36ЈS, 17Њ03ЈE, 14.iii.1976, B. Lamoral (NMSA 11304). 10850); 2 (,2&, NAMIBIA, Otjozondjupa Re- 22. Parabuthus planicauda (Pocock, 1889): gion, Okahandja, 24.ix.1994, I. Engelbrecht Lectotype &, SOUTH AFRICA, Dr Quain (SAMC C4605). (BMNH 1870.26); 6 &, SOUTH AFRICA, West- 16. Parabuthus kuanyamarum Monard, 1937: ern Cape Prov., Ashton, 33Њ49ЈS, 20Њ03ЈE, i.1914, 3 &, BOTSWANA, N Khwaai and Lechwee W.F. Purcell (SAMC B1748); 4 (,6&, 2 subadult camps, 18Њ40ЈS±19Њ00ЈS, 23Њ00ЈE±23Њ45ЈE, 16± (, juv &, SOUTH AFRICA, Eastern Cape Prov., 20.xi.1979, B. Lamoral (NMSA 13972); 13 (, &, Brakkloof, Grahamstown, 33Њ14ЈS, 26Њ23ЈE, 7 juveniles, NAMIBIA, Hardap Region, Farm 1897, J. White (SAMC 1734); 15 (,28&, Ghobab 381, 23Њ26ЈS, 17Њ21ЈE, 12.iii.1976, B. La- SOUTH AFRICA, Eastern Cape Prov., Graaff- moral (NMSA 10813). Reinet and Kruidfontein, 8 mi from Graaff-Re- 17. Parabuthus laevifrons (Simon, 1888): 2 (, inet, 32Њ22ЈS, 24Њ36ЈE, ix.1902, J. Paynter &, juv &, NAMIBIA, Karas Region, Farm Ort- (SAMC 12008); (, &, SOUTH AFRICA, West- mansbaum 120, 28Њ19ЈS, 18Њ43ЈE, 26±28.i.1973, ern Cape Prov., Stilbaai, 8±12.xi.1940, V. Fitzsi- B. Lamoral and L. Ferguson (NMSA 10509); (, mons (TMSA 8520). 4 &, 2 subadult &, 3 juv (, 3 juv &, NAMIBIA, 23. Parabuthus raudus (Simon, 1888): 8 (,5 Karas Region, Tses, dune strip SE, 25Њ53ЈS, &, subadult &, 3 juv (, juv &, NAMIBIA, Kho- 18Њ10ЈE, 23±24.ii.1973, B. Lamoral and K. Porter mas Region, Farm Frischgewaagd 289, 22Њ32ЈS, (NMSA 10521). 17Њ50ЈE, 20.iii.1976, B. Lamoral (NMSA 10817); 18. Parabuthus mossambicensis (Peters, 1861): &, 2 subadult (, SOUTH AFRICA, Northern (, MOZAMBIQUE, Gorongoza, 22.ii.1971, G. Cape Prov., Kalahari Gemsbok National Park, Vasse (NMSA 10088); (, SOUTH AFRICA, Mata Mata, 3 miles N, 24.iv.1970, B. Lamoral Northern Prov., Messina Nature Reserve, (NMSA 10931); 3 (,2&, SOUTH AFRICA, xii.1993, L. Prendini and K.M.A. Prendini Northern Cape Prov., Richtersveld, Swaartpoort (SAMC C4606); 3 (, SOUTH AFRICA, North- near Ochta Diamond Mine, 28Њ07ЈS, 16Њ56ЈE, ern Prov., Farm Rochdale 700, i.1996, L. Prendini ii.1974, R. Faber (NMSA 10924). and J. Laing (SAMC C4607); 3 &, SOUTH AF- 24. Parabuthus schlechteri Purcell, 1899: &, RICA, Northern Prov., Waterpoort, i.1996, L. subadult (, subadult &, NAMIBIA, Karas Re- Prendini and J. Laing (SAMC C4608). gion, Farm Tsirub 13, 26Њ52ЈS, 16Њ02ЈE, 19. Parabuthus muelleri Prendini, 2000: Holo- 3.iii.1976, B. Lamoral (NMSA 10730); (, &, type &, NAMIBIA, Hardap Region, Farm Onis 8, NAMIBIA, Karas Region, Keetmanshoop, Farm 82 km from Sesriem to Naukluft, 24Њ22.46ЈS, Noachabeb, 7±12.i.1972, (NMSA 11406); (, &, 16Њ13.17ЈE, 1260 m, 7.i.1998, L. Prendini and E. SOUTH AFRICA, Northern Cape Prov., Agge- Scott (NMNW 1854); paratype &, NAMIBIA, neys, 6±8.xii.1997, L. Prendini, G.J. MuÈller, K. Karas Region, Farm Plateau 38, near Aus, Rostoll, and J. du Plessis (SAMC C4609); (, 26Њ40.62ЈS, 16Њ31.85ЈE, 1550 m, 30.xii.1997, L. SOUTH AFRICA, Western Cape Prov., Ga- Prendini and E. Scott (SAMC C4514); (, NA- mkaskloof Nature Reserve, 33Њ31ЈS, 21Њ37ЈE, MIBIA, Karas Region, locality uncertain, proba- 21.ii.1997, M. de Jager (SAMC C4610). bly Aus, 26Њ41ЈS, 16Њ15ЈE, A. Harington (AMNH 25. Parabuthus stridulus Hewitt, 1913: Holo- [AH 3991]). type (, NAMIBIA, LuÈderitzbucht, South West 20. Parabuthus namibensis Lamoral, 1979: Ho- Africa [Karas Region, 26Њ35ЈS, 15Њ10ЈE], lotype &, paratype (, NAMIBIA, Erongo Region, 26.xi.1912 (TMSA 1868); (,2&, NAMIBIA, Cape Cross, 5 km N, 21Њ43ЈS, 13Њ56ЈE, Karas Region, Agate Beach, LuÈderitz, ii.1973, B. 25.iii.1976, B. Lamoral and L. Ferguson (NMSA Lamoral (NMSA 10573); &, NAMIBIA, Erongo 2184); (, NAMIBIA, Erongo Region, RoÈssing, Region, Cape Cross, 5 km N, 21Њ43ЈS, 13Њ56ЈE, Lower Ostrich Gorge, 22Њ30ЈS, 14Њ58ЈE, 2± 25.iii.1976, B. Lamoral and L. Ferguson (NMSA 3.xi.1985, Irish and Rust (NMNW 896); &, juv 10907); (, NAMIBIA, Erongo Region, Cape (, same data, except 8.v±5.vi.1984, E. Grif®n Cross, 5 km N, 21Њ43ЈS, 13Њ56ЈE, 25.iii.1976, B. (NMNW 845). Lamoral and L. Ferguson (NMSA 10904); (, sub- AMERICAN MUSEUM NOVITATES Friday May 02 2003 03:12 PM novi 03331 Mp_22 Allen Press • DTPro System File # 01TQ

22 AMERICAN MUSEUM NOVITATES NO. 3408

adult (, NAMIBIA, Karas Region, Kolmanskop, SOUTH AFRICA, Northern Prov., Langjan Na- 12 km E LuÈderitz, 26Њ43ЈS, 15Њ17ЈE, iii.1973, C.J. ture Reserve, i.2000, L. Prendini and E. Scott Coetzee (NMSA 10501); (, &, NAMIBIA, Karas (SAMC C4613). Region, Farm Plateau 38, near Aus, 26Њ38.63ЈS, 27. Parabuthus villosus (Peters, 1862): sub- 16Њ30.77ЈE, 30.xii.1997, L. Prendini and E. Scott adult (, NAMIBIA, Kunene Region, Farm Groot- (SAMC C4611). berg 191, 19Њ46ЈS, 14Њ15ЈE, 2.iv.1976, B. Lamoral 26. Parabuthus transvaalicus Purcell, 1899: and L. Ferguson (NMSA 10913); (,3&, NA- Lectotype &, paralectotype &, SOUTH AFRICA, MIBIA, Karas Region, Farm Plateau 38, 26Њ40ЈS, ``Transvaal'' (SAMC 3003); &, SOUTH AFRI- 16Њ30ЈE, 29.ii.1976, B. Lamoral (NMSA 10805); CA, Northern Prov., Dendron, Soutpansberg, &, NAMIBIA, Kunene Region, Kamanjab, 3 km 18.iii.1970, (NMSA 11449); &, SOUTH AFRI- W, 1 9 Њ37ЈS, 14Њ48ЈE, 5.iv.1976, B. Lamoral and CA, Northern Prov., Kruger National Park, Pafuri, L. Ferguson (NMSA 10833); &, NAMIBIA, Ku- 22Њ27ЈS, 31Њ17ЈE, 18.x.1980, L. Braack (NMSA nene Region, Sesfontein, 3 km N clinic, 19Њ07ЈS, 13899); (, &, SOUTH AFRICA, Northern Prov., 13Њ36ЈE, 3.iv.1976, B. Lamoral (NMSA 10738); Mphakane, S, granite koppies 1 km from turnoff 2 (, SOUTH AFRICA, Northern Cape Prov., Pel- to Munnik, 23Њ32.20ЈS, 29Њ42.42ЈE, 29.xii.1999, la pumpstation, iii.1997, L. Prendini and G.J. L. Prendini and E. Scott (SAMC 4612); (, MuÈller (SAMC C4614).

APPENDIX 2

CHARACTERS AND CHARACTER STATES USED FOR 8. Chela movable ®nger, length compared with CLADISTIC ANALYSIS OF PARABUTHUS POCOCK, length of manus (measured along ventroexternal 1890 carina), in &: long (length ®nger/length carina: Character states were scored 0±2, ? (unknown), 1.70±2.00) (0); short (length ®nger/length carina: ± (inapplicable), or * (polymorphic). Multistate Ϯ1.50) (1). characters were treated as unordered (nonaddi- 9. Chela manus, shape in adult (, compared tive). Three autapomorphies, indicated by ², were with adult &: similar (0); sexually dimorphic (1); excluded from all analyses. Refer to table 1 for unknown (?). data matrix. 10. Chela ®xed ®nger, shape in adult (: straight or slightly curved dorsally such that proximal GENERAL dentate margin linear when ®ngers are closed (0); strongly curved dorsally, proximal dentate margin 1. Adult general size: large, carapace length distinctly emarginate when ®ngers are closed (1); 6.5±17.0 mm (0); small, carapace length 2.5±5.0 unknown (?). mm (1). 11. Chela movable ®nger, shape in adult (: 2. Color of carapace, mesosoma, and metaso- straight, proximal dentate margin linear when ®n- ma: pale yellow to light brown (0); dark brown to gers are closed (0); curved ventrally, proximal black (pedipalps and legs may be dark or pale) dentate margin distinctly emarginate when ®ngers (1); polymorphic (*). are closed (1); unknown (?). 3. Color pattern of metasoma: metasomal seg- 12. Chela ®xed and movable ®ngers with basal ments I±V and telson uniformly colored (0); me- lobe in adult (: absent (0); present (1); unknown tasomal segments III±V and telson infuscated (?). (i.e., darker than segments I and II) (1); polymorphic (*). TRICHOBOTHRIA

CARAPACE 13. Pedipalp femur, position of e1: level with or distal to d5 (0); almost halfway between d4 and d5 4. ²Carapace dorsoventrally compressed: ab- (1).

sent (0); present (1). 14. Pedipalp patella, position of esb2: distinctly 5. Granulation of median ocular tubercle ((, &) distal to esb1 (0); level with or slightly distal to and surrounding surfaces (&): entire (0); smooth esb1 (1). areas (1). 15. Chela ®xed ®nger, position of dt: in line with or distal to et (0); proximal to et (1). PEDIPALPS PECTINES 6. Pedipalps, setation: setose (0); smooth (1). 7. Chela manus, surface: smooth (0); granular 16. Proximal median lamella of pectines, in &: (1). arcuate (0); enlarged and lobate (dilate) (1). AMERICAN MUSEUM NOVITATES Friday May 02 2003 03:12 PM novi 03331 Mp_23 Allen Press • DTPro System File # 01TQ

2003 PRENDINI: PARABUTHUS MUELLERI 23

17. Proximal median lamella of pectines, in (: segment (0); rounded, extended beyond antero- subrectangular (0); weakly lobate (dilate) (1); dorsal edge of segment (1); inapplicable (±). strongly lobate (dilate) (2); unknown (?). 33. Metasomal segment I, extent of stridulatory region (if present) in dorsal aspect: terminating at LEGS anterodorsal edge of segment (0); extended for- wards in V-shape onto anterior surface (1); inap- 18. Legs IV, length: moderately long, not reach- plicable (±). ing to posterior edge of metasomal segment III 34. Metasomal segment II, extent of stridula- (0); very long, reaching past posterior edge of me- tory region (if present) in dorsal aspect: reaching tasomal segment III (1). posterodorsal margin (0); not reaching postero- 19. Basitarsi of legs I and II, macrosetal combs: dorsal margin (1); inapplicable (±); polymorphic absent (0); weakly developed (1); strongly devel- (*). oped (2). 35. Metasomal segment II, nature of stridula- 20. Basitarsi of legs I and II, laterally expand- tory region (if present): ®ne to coarse granules ed: absent (0); present (1). (0); horizontal ridges (1); inapplicable (±). 21. ²Basitarsi of legs III and IV, prolateral sur- 36. Metasomal segments II and III, posterodor- faces with dense tufts of macrosetae: absent (0); sal edge: straight (0); anteromedially curved in a present (1). V-shape (1). 22. Telotarsal ungues, relative length: equal (0); 37. Metasomal segments IV and V, lateral in- subequal (1). tercarinal surfaces: granular (0); smooth (1). 38. Metasomal segments I±IV, dorsosubmedian HEMISPERMATOPHORE carinae: present (0); absent (1). 39. Metasomal segment IV, dorsosubmedian ca- 23. Hemispermatophore, pars recta: parallel to rinae (if present): continuous (0); discontinuous axis (0); S-shaped (1); unknown (?). (1); inapplicable (±). 40. Metasomal segments II±IV, distal spiniform MESOSOMA granules of dorsosubmedian carinae (if present), size relative to preceding granules: equally devel- 24. Sternites, surface: smooth (0); punctate (1). oped (0); noticeably more pronounced (1); inap- 25. Sternite III, ``pit'' organ at proximal apex: absent (0); present (1). plicable (±). 26. Sternite VII, carinae: present (0); absent (1). 41. Metasomal segment IV, ventrosubmedian and ventrolateral carinae: present and continuous to edge of segment (0); present but ventrosub- METASOMA median carinae becoming obsolete distally (1); 27. Metasomal segments I±V and telson, seta- absent (2). tion: virtually asetose (0); sparsely to moderately 42. Metasomal segment IV, median lateral ca- setose (1); very densely setose (2). rina: absent to proximally obsolete (0); continuous 28. Metasomal segments, width relative to but poorly developed (1); continuous and distinct length: much narrower (length IV/width IV: 1.7± (2). 2.11) (0); slightly narrower (length IV/width IV: 43. Metasomal segment V, dorsosubmedian ca- 1.2±1.5) (1). rinae: absent (0); present, poorly developed with 29. Metasomal segments, width from I±IV: be- blunt, rounded granules (1); present, distinct with coming narrower distally, metasomal segment I sharp, spiniform granules (2). wider than segment IV (0); becoming wider dis- 44. Metasomal segment V, dorsolateral carinae: tally, metasomal segment I narrower than segment absent, except for a few proximal granules (0); IV (1). distally obsolete (1); continuous to distal edge of 30. Metasomal segments I±III, stridulatory re- segment (2). gion on dorsal surface: absent from I±III (0); 45. Metasomal segment V, ventrolateral cari- strongly developed on I±III (1); strongly devel- nae: converging distally (0); subparallel to diverg- oped on I and II, weakly developed to absent on ing distally (1). III (2). 46. Metasomal segment V, distal half of ventro- 31. Metasomal segment I, shape of stridulatory lateral carinae: with spinose processes (0); with region (if present) in dorsal aspect: narrow, par- lobate processes (1). allel-sided (0); broad, rounded anteriorly, with 47. Metasomal segment V, ventrosubmedian ca- posterior constriction (1); inapplicable (±). rinae: absent or indistinct from surrounding gran- 32. Metasomal segments I, and to a lesser ex- ules (0); distinct (1). tent II, shape of stridulatory region (if present) in 48. Metasomal segment V, ventromedian cari- lateral aspect: truncated at anterodorsal edge of na: present (may be indistinct) (0); absent (1). AMERICAN MUSEUM NOVITATES Friday May 02 2003 03:12 PM novi 03331 Mp_24 Allen Press • DTPro System File # 01TQ

24 AMERICAN MUSEUM NOVITATES NO. 3408

TELSON BEHAVIOR 49. Telson vesicle, width relative to width of 52. Diurnal retreat: hides under rocks (0); bur- metasomal segment V: approximately equal (0); rows under rocks (1); burrows in open ground (2); considerably narrower (1). unknown (?); polymorphic (*). 50. ²Telson vesicle, dorsoproximal surface: 53. Foraging strategy: sit-and-wait (0); errant very shallowly excavated along longitudinal half (1); unknown (?). (0); deeply excavated (1). 51. Telson aculeus, shape: gently curved (0); abruptly bent (1).

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a This paper meets the requirements of ANSI/NISO Z39.48-1992 (Permanence of Paper).