Systematic Revision Of The Arboreal Neotropical “ Thorellii ” Clade Of Centruroides Marx, 1890, Bark Scorpions (Buthidae C. L. Koch, 1837) With Descriptions Of Six New Species Author Goodman, Aaron M. Graduate School and University Center, City University of New York; Division of Invertebrate Zoology, American Museum of Natural History; Institute for Biodiversity Science and Sustainability, California Academy of Sciences Author Prendini, Lorenzo Arachnology Lab and Scorpion Systematics Research Group, Division of Invertebrate Zoology, American Museum of Natural History Author Francke, Oscar F. Colección Nacional de Arácnidos, Departamento de Zoología, Instituto de Biología, Universidad Nacional Autónoma de México Author Esposito, Lauren A. Graduate School and University Center, City University of New York; Division of Invertebrate Zoology, American Museum of Natural History; Institute for Biodiversity Science and Sustainability, California Academy of Sciences text Bulletin of the American Museum of Natural History 2021 2021-09-16 2021 452 1 93 journal article 2777 10.1206/0003-0090.452.1.1 01c03222-2a9d-47ec-9f63-776920b6a540 0003-0090 5825896 Centruroides yucatanensis , sp. nov. Figures 2 , 4 , 7E, F , 10E, F , 15C , 16C , 17O, R , 18O, R , 19O, R , 20O, R , 21O, R , 22O, R , 23O, R , 24O, R , 25R, O , 42 , 43 , tables 1 , 9 , 10 Centruroides schmidti : Armas, 1996: 25–29 , 98, figs. 1–4 (misidentification), table I. Centruroides sissomi : Vázquez, 1999: 53 , 60–62, fig. 7 (misidentification); Armas et al., 2003: 95 (misidentification, part); Armas, 2006: 4 , 7, fig. 5 (misidentification, part); Teruel et al., 2015a: 8 (misidentification); Delfín-González et al., 2017: 283, 285, table 2 (misidentification); Esposito and Prendini, 2019: 4 , fig. 2 (misidentification); Ponce-Saavedra and Francke, 2019: 3, table 1 (misidentification); Crews and Esposito, 2020: 14 , fig. 11 (misidentification). TYPE MATERIAL : MEXICO : Quintana Roo : Município Benito Juárez : Holotype ♂ ( CNAN T01416 ), paratype ♀ (CNAN T01417 ), 2 juv. ♀ paratypes (CNAN T01418 , T01419 ), Puerto Morelos , Jardín Botánico Alfredo Barrera, 20°50′42.1″N 86°54′12.9″W , 23 m , 4.vii.2007 , G. Montiel, R. Paredes, M. Ramírez, D. Chibras, and G. Bonilla. ETYMOLOGY: The species name refers to the Yucatán Peninsula of southeastern Mexico , where the species occurs. DIAGNOSIS: Centruroides yucatanensis differs from the closely related species, C. chanae and C. hoffmanni , as follows. The carapace, pedipalps, tergites, and metasoma are less infuscate, creating a less mottled appearance, in C. yucatanensis than C. chanae . Less reticulate infuscation is present on the chelicerae of C. yucatanensis than C. chanae . The interocular triangle is less darkly infuscate in C. yucatanensis than C. hoffmanni . The carapace is shorter, its length and width similar, in C. yucatanensis (fig. 7E, F, table 10 ) but longer, its length greater than its width, in C. hoffmanni (fig. 7A, B, table 10 ). The carapace surfaces are more finely granular, the carinae less developed and the sulci broader and shallower in C. yucatanensis than C. hoffmanni . The pedipalp chela manus of the male is less incrassate in C. yucatanensis (fig. 15C) than C. hoffmanni (fig. 15B), with fewer spiniform granules on its prolateral surfaces than in C. chanae and C. hoffmanni (fig. 15A, B). The ventral surfaces of the telotarsi of leg I are more finely and sparsely setose in C. yucatanensis than C. chanae . The pectinal tooth count of the male is lower in C. yucatanensis , usually 13 or 14 (fig. 9E, table 10 ) than C. chanae , usually 17 (fig. 9A, table 10 ) and C. hoffmanni , usually 15 (fig. 9D, table 10 ), and the pectinal teeth are more ovoid. The ventrolateral and ventrosubmedian carinae of mesosomal sternite VII are vestigial, weakly granular in C. yucatanensis , whereas the ventrolateral carinae are distinct, granular, and the ventrosubmedian carinae weakly developed, granular in C. chanae , and the ventrolateral carinae granular, and the ventrosubmedian carinae weakly granular and restricted to the posterior half of the segment in C. hoffmanni . The ventrolateral and ventrosubmedian carinae of the metasomal segments are more pronounced in C. yucatanensis , being slightly serrate on segments I–IV, compared with finely granular to subserrate on I–III and obsolete, smooth on IV in C. chanae . The ventrosubmedian carinae of segments I and II are absent in C. yucatanensis (figs. 18O, R, 19O, R, 20O, R, 21O, R, 22O, R), absent or obsolete in C. chanae (figs. 18I, L, 19I, L, 20I, L, 21I, L, 22I, L), and very pronounced in C. hoffmanni (figs. 18C, F, 19C, F, 20C, F, 21C, F, 22C, F). TABLE 10 Diagnostic ratios (mean/median/mode) for species of the arboreal Neotropical “ thorelli ” clade of Centruroides Marx, 1890 , bark scorpions

Leg I length: carapace length	Metasoma V length: mesosoma length	Metasoma V length: carapace length
Centruroides berstoni , sp. nov.	1.9/1.9/1.9	2.5/2.4/2.0	1.6/1.6/1.9
Centruroides catemacoensis , sp. nov.	1.9/1.9/1.9	2.3/2.2/2.2	1.5/1.5/1.7
Centruroides chanae , sp. nov.	1.9/1.9/1.9	3.0/3.1/3.1	1.7/1.8/1.8
Centruroides cuauhmapan , sp. nov.	1.8/2.1/1.9	2.2/2.1/1.8	1.5/2.1/1.7
Centruroides hamadryas , sp. nov.	1.8/1.9/1.8	1.8/2.8/2.8	1.6/1.7/1.8
Centruroides hoffmanni Armas, 1996	1.8/1.8/1.8	2.3/2.4/1.7	1.6/1.7/1.3
Centruroides rileyi Sissom, 1995	1.9/1.9/1.9	2.2/2.1/2.1	1.5/1.5/1.2
Centruroides schmidti Sissom, 1995	2.0/1.9/1.9	2.2/2.2/2.8	1.5/1.4/1.9
Centruroides yucatanensis , sp. nov.	1.8/1.8/1.8	2.1/2.1/2.0	1.5/1.4/1.3

DESCRIPTION: The following description is based on the holotype male, with differences among other material noted in the section on variation. Coloration: Base color yellow, with extensive infuscation, creating mottled or marbled pattern. Carapace with uniformly infuscate marbling, more densely infuscate medially. Pedipalp chela fingers and manus, dorsal and retrolateral intercarinal surfaces with moderately infuscate marbling; prolateral and ventral intercarinal surfaces mostly immaculate. Legs retrolateral surfaces with infuscate marbling; prolateral surfaces pale, immaculate. Tergites with unformly infuscate mottling, pale stripe medially, blackish spots submedially, and faint, narrow bands laterally. Sternites with faintly infuscate marbling. Metasomal segments uniformly, faintly marbled; segment V and telson markedly infuscate, noticeably darker than preceding segments. Carapace: Shape trapezoidal; anterior width four-fifths of posterior width ( table 10 ); anteromedian sulcus moderately deep, oval; posteromedian sulcus shallow anteriorly, deep posteriorly; median ocular tubercle weakly granular; carinae moderately developed, comprising small to medium-sized granules; lateral ocular and posterosubmedian carinae weakly developed; intercarinal surfaces finely and evenly granular (fig. 10E). Pedipalps: Orthobothriotaxic, Type A; femur dorsal trichobothria with α configuration; pedipalp chela fixed finger, trichobothrium db situated slightly distal to et. Femoral carinae strongly developed, serrate; dorsal intercarinal surface moderately granular; prolateral intercarinal surface with series of large spiniform granules. Patella carinae strongly developed, granular; prolateral intercarinal surface with five or six large subspiniform granules. Chela manus dorsomedian and retrodorsal carinae complete, granular; prodorsal carina absent. Fixed finger, median denticle row comprising eight oblique subrows, each flanked by pro- and retrolateral supernumerary denticles. Movable finger, median denticle row with short terminal row comprising four denticles preceded by eight oblique subrows, each flanked by pro- and retrolateral supernumerary denticles. . mm 5 = bars Scale. aspect Ventral. B . aspect Dorsal. A . habitus,) 01407 T CNAN ( ♂ ,. nov. sp, chanae Centruroides . 40 FIGURE FIGURE 41. Centruroides chanae , sp. nov. , ♀ (CNAN T01405), habitus. A. Dorsal aspect. B. Ventral aspect. Scale bars = 5 mm. Legs: Leg I length 1.78× greater than carapace length ( table 10 ). Telotarsi ventral surfaces sparsely covered with short setae; ungues markedly curved. Pectines: Pectinal plate 1.61× wider than long; posterior margin distinctly rounded; pectinal tooth count 13/14 ( ♂ ) (fig. 8D, table 10 ). Mesosoma: Tergites width similar to carapace posterior width; I and II slightly narrower ( table 10 ). Pretergites surfaces smooth to finely granular. Posttergites surfaces weakly granular; I–VI with dorsomedian carinae absent on I and II, vestigial, granular on III–VI; VII surface weakly granular, dorsomedian carina vestigial, granular, dorsosubmedian and dorsolateral carinae smooth. Sternites III–VI, surfaces smooth; VII surface smooth, ventrolateral and ventrosubmedian carinae smooth. FIGURE 42. Centruroides yucatanensis , sp. nov. , ♂ (CNAN T01416), habitus. A. Dorsal aspect. B. Ventral aspect. Scale bars = 10 mm. FIGURE 43. Centruroides yucatanensis , sp. nov. , ♀ (CNAN T01417), habitus. A. Dorsal aspect. B. Ventral aspect. Scale bars = 10 mm. Metasoma: Metasoma length 2.45× mesosoma length ( table 10 ). Segments longer than wide; increasing in length posteriorly, segment V 2× length of I; ventral carinae vestigial, weakly granular on segments I–IV, other carinae absent or obsolete; lateral intercarinal surfaces sparsely granular on segments I–III, other surfaces smooth (figs. 17–22O). Telson: Vesicle elongate, ovoid; ventral surface shallowly convex; ventromedian carina granular, terminating at subaculear tubercle; subaculear tubercle narrow and angular in lateral aspect, directed toward midpoint of aculeus. Aculeus angled ventrally at slightly less than 90° (fig. 25O, R). Variation: Base coloration varies from light yellow to orange. Adult males and females differ as follows. The pedipalp chela manus is incrassate, with the prodorsal carina spinose, the mesosoma proportionally longer and slenderer, the metasoma 2× longer, with segment V markedly longer (1.86× carapace length), and the telson more elongate, with the vesicle more rounded and bilobed posteriorly, in males (figs. 23O, R, 24O, R, 25O, R, table 10 ). The tegument is more densely infuscate, the prodorsal carina of the pedipalp chela manus finely granular, the pectinal plate produced into a rounded lobe posteriorly, which is punctate and slightly infuscate, metasomal segment V shorter (1.3× carapace length) and the telson shorter and narrower, with the vesicle surfaces less granular, in females (figs. 10E, F, 15–16C, 17O, R, 18O, R, 19O, R, 20O, R, 21O, R, 22O, R, 23O, R, 24O, R, 25O, R, table 10 ). The pectinal tooth count is similar in both sexes ( table 9 ). DISTRIBUTION: Centruroides yucatanensis is endemic to the Yucatán Peninsula. Although presently known from only three localities, two in the state of Yucatán in the north of the peninsula, and a third on the western coast, in the state of Quintana Roo , the distribution of this species was probably more extensive before much of its habitat was destroyed for agriculture and rangeland (fig. 4). ECOLOGY: The localities at which C. yucatanensis has been recorded range in altitude from 23– 38 m . The habitat at these localities varies from low or medium semideciduous broadleaf forest to tall, moist evergreen broadleaf forest, often with a dense understory. Much of the original habitat has been cleared for agriculture and rangeland across the Yucatán Península, and this species appears to be confined to remnant patches of forest. The habitat and habitus are consistent with the arboreal, corticolous ecomorphotype ( Prendini, 2001a ). REMARKS: Armas (1996) identified 11 individuals from the Jardín Botánico Alfredo Barrera, Puerto Morelos, Quintana Roo, as C. schmidti . Meristic data recorded by Armas (1996) for an adult male and three females differed greatly from comparative data for the holotype of C. schmidti collected at Lake Tickamaya, Honduras ( Sissom, 1995 ). Armas (1996) also described C. sissomi , and assigned it to the thorellii group due to its small size. Illustrations and photographs of the holotype female of C. sissomi in the original description indicate that the pectinal plate is not lobed, and subsequent photographs of the holotype by Armas (2006) indicate a densely granular tegument, characters inconsistent with the “ thorellii ” clade. Material examined during the present study confirmed the presence of two sympatric species of Centruroides in the Jardín Botánico Alfredo Barrera, Puerto Morelos. One species, determined to be conspecific with the material previously misidentified as C. schmidti by Armas (1996) , based on meristic data, is a hitherto undescribed species, described herein as C. yucatanensis . The other is presumed to be C. sissomi , based on the fact that the pectinal plate is not lobed and the tegument is densely granular, as well as its dark orange coloration and larger size ( 40 mm ). Other records of C. schmidti and/or C. sissomi from the Yucatán Península ( Vázquez, 1999 ; Teruel et al., 2015a ; Ponce-Saavedra and Francke, 2019 ) are misidentifications of C. yucatanensis . MATERIAL EXAMINED: MEXICO : Quintana Roo : Município Benito Juárez: Puerto Morelos , Jardín Botánico Alfredo Barrera, 20°50′42.1″N 86°54′12.9″W , 38 m , 4.vii.2007 , G. Montiel, R. Paredes, M. Ramírez, D. Chibras, and G. Bonilla, 1 ♂ (AMNH [LP 7597]), 2 juv. ♂ (CNAN SC3984). Yucatán : Município Felipe Carrillo Puerto: Cenote Chac-ha, 3.5 km N and 3 km E of Kalacmul, 20°04′40.3″N 88°08′27.9″W , 23 m , 9.vii.2007 , R. Paredes, D. Chibras, and G. Montiel, 1 ♀ (CNAN SC4004).