taxonID	type	description	language	source
700187A3FFD7FFC5FCA3FD7CB1DCF808.taxon	description	(Figs 4 – 39)	en	Xuan, Qiang, Prendini, Lorenzo, Engel, Michael S., Cai, Chenyang, Huang, Diying (2025): Extinct scorpion family Chaerilobuthidae from Mid-Cretaceous Burmese amber reinterpreted as subfamily of extant family Pseudochactidae (Chelicerata: Scorpiones). Zoological Journal of the Linnean Society 203 (1): 1-60, DOI: 10.1093/zoolinnean/zlae169, URL: https://doi.org/10.1093/zoolinnean/zlae169
700187A3FFD7FFC5FCA3FD7CB1DCF808.taxon	diagnosis	Diagnosis: The species of Chaerilobuthus share many characters with the extant family Pseudochactidae, notably: sternum pentagonal (Figs 12 B, 15 B, 20 E, 23 B, 26 B, 28 B, 33 B, 34 D, 37 F); cheliceral moveable finger basal teeth absent; moveable finger ventral surface with serrula (Figs 6 C, 12 G, 18 F, 23 C, 29 B, 34 B, 37 B); median denticle rows of pedipalp chela fingers comprising oblique subrows (Figs 13 C, 15 E, 29 F, 33 C); pedipalps with Type D trichobothrial pattern, β configuration; trichobothrium d 2 situated on femur dorsal surface, angle formed by d 1, d 3, and d 4 opening toward prolateral surface (Figs 7, 10, 13, 16, 21, 24, 26, 31, 35, 38); legs with prolateral and retrolateral pedal spurs present, legs III and IV with tibial spurs absent (Figs 6 I – L, 7 F – I, 12 K, L, 15 G, 18 C, 20 J, K, 23 J, K, 29 G – J, 33 C, D, G, H, K, 34 I – L, 37 D), and telotarsi each with two rows of ventrosubmedian spinules (Figs 6 J – L, 9 G, I, 23 J, K, 29 H, J); metasomal segment V with paired ventrosubmedian carinae (Figs 5 F, 9 N, 20 L, 33 I, J, 35 I); telson without subaculear tubercle (Figs 5 F, 9 N, 14 B, 18 G, 19 B, 33 I, J, 37 I). The extinct genus may be distinguished from other pseudochactid scorpions by the following combination of characters: carapace anterior margin sublinear, anterolateral margins entire, posterolateral margins angular, slanting, posteromedian margin slightly recurved (Figs 6 A, 9 A, 12 A, 15 A, 20 B, 23 A, 26 A, 28 A, 34 A, 37 A); carapace circumocular sutures absent (Figs 6 A, 9 A, 12 A, 15 A, 20 B, 23 A, 26 A, 28 A, 34 A, 37 A); median ocelli absent and one pair of large posterolateral major lateral ocelli present (Figs 6 A, 9 A, 12 A, 15 A, 18 D, 20 A, B, 23 A, 26 A, 28 A, 29 A, 33 A, 34 A, C, 37 A, C); pedipalp femoral trichobothria d 3 and d 4 situated closer to midpoint of segment (Figs 7 A, 10, 13 A, 16, 24 A, 26 D, 31, 35 C, J) and d 3 – d 4 axis directed slightly away from retrodorsal carina rather than parallel to it (β configuration) (Figs 7 A, 10, 13 A, 16, 24 A, 26 D, 31, 35 C, J); pectines with low tooth counts (five to eight teeth) (Figs 6 D, 12 B, 15 D, 18 I, 20 G, 23 F, 26 B, 29 C, 33 B, 34 G, 37 H). Additionally, two new characters, considered diagnostic for Chaerilobuthus, are shared with the extant pseudochactid, Qianxie solegladi (Tang, 2022: 7, 11, figs 10 43 – 46): leg I maxillary lobes (coxapophyses) anterior margins with serrula (Fig. 38); leg basitarsi with pair of spinule rows distally (Figs 6 I – L, 9 F – I, 12 H, L, 15 G, 18 C, 20 J, K, 23 K, 29 G – J, 33 G, H, 34 I – L).	en	Xuan, Qiang, Prendini, Lorenzo, Engel, Michael S., Cai, Chenyang, Huang, Diying (2025): Extinct scorpion family Chaerilobuthidae from Mid-Cretaceous Burmese amber reinterpreted as subfamily of extant family Pseudochactidae (Chelicerata: Scorpiones). Zoological Journal of the Linnean Society 203 (1): 1-60, DOI: 10.1093/zoolinnean/zlae169, URL: https://doi.org/10.1093/zoolinnean/zlae169
700187A3FFD7FFC5FCA3FD7CB1DCF808.taxon	discussion	Remarks: Results of morphological phylogenetic analysis justify the transfer of Chaerilobuthidae to Pseudochactidae, as Chaerilobuthinae, stat. nov.; the following new synonyms: Chaeriloiurus Lourenço, 2020 = Chaerilobuthus Lourenço, 2016, synon. nov., and Serratochaerilobuthus Lourenço, 2024 = Chaerilobuthus Lourenço, 2016, synon. nov.; and the following new combinations: Chaerilobuthus brigittemuellerae (Lourenço and Velten, 2020), comb. nov. and Chaerilobuthus schmidti (Lourenço and Velten, 2024), comb. nov ..	en	Xuan, Qiang, Prendini, Lorenzo, Engel, Michael S., Cai, Chenyang, Huang, Diying (2025): Extinct scorpion family Chaerilobuthidae from Mid-Cretaceous Burmese amber reinterpreted as subfamily of extant family Pseudochactidae (Chelicerata: Scorpiones). Zoological Journal of the Linnean Society 203 (1): 1-60, DOI: 10.1093/zoolinnean/zlae169, URL: https://doi.org/10.1093/zoolinnean/zlae169
700187A3FFD0FFCDFF39F9DAB710F909.taxon	description	(Figs 4 – 10)	en	Xuan, Qiang, Prendini, Lorenzo, Engel, Michael S., Cai, Chenyang, Huang, Diying (2025): Extinct scorpion family Chaerilobuthidae from Mid-Cretaceous Burmese amber reinterpreted as subfamily of extant family Pseudochactidae (Chelicerata: Scorpiones). Zoological Journal of the Linnean Society 203 (1): 1-60, DOI: 10.1093/zoolinnean/zlae169, URL: https://doi.org/10.1093/zoolinnean/zlae169
700187A3FFD0FFCDFF39F9DAB710F909.taxon	materials_examined	Type material: Holotype: one juv. (sex unknown) (personal collection of KlausPeter Brucksch, Kuranda, Australia), Cretaceous (Cenomanian) burmite: MYANMAR: Kachin State (precise locality unknown).	en	Xuan, Qiang, Prendini, Lorenzo, Engel, Michael S., Cai, Chenyang, Huang, Diying (2025): Extinct scorpion family Chaerilobuthidae from Mid-Cretaceous Burmese amber reinterpreted as subfamily of extant family Pseudochactidae (Chelicerata: Scorpiones). Zoological Journal of the Linnean Society 203 (1): 1-60, DOI: 10.1093/zoolinnean/zlae169, URL: https://doi.org/10.1093/zoolinnean/zlae169
700187A3FFD0FFCDFF39F9DAB710F909.taxon	diagnosis	Diagnosis: The following combination of characters is diagnostic for C. bruckschi: carapace anterior margin sublinear (Figs 6 A, 9 A); lateral ocelli markedly protruding (Figs 6 A, 9 A), pectines each with five or six teeth (Fig. 6 D); pedipalp chela relatively slender, ratio of chela length: chela manus width less than 5 (Fig. 10); ratio of metasomal segment IV: V length less than 2 (Figs 5 C, D, E, F, 9 K, L, M, N); metasomal segment V dorsolateral carinae slightly serrate (Fig. 9 M); telson vesicle slightly bulbous (Figs 5 E, F, 9 M, N), aculeus moderately long and curved, base narrow (Figs 5 E, F, 9 M, N).	en	Xuan, Qiang, Prendini, Lorenzo, Engel, Michael S., Cai, Chenyang, Huang, Diying (2025): Extinct scorpion family Chaerilobuthidae from Mid-Cretaceous Burmese amber reinterpreted as subfamily of extant family Pseudochactidae (Chelicerata: Scorpiones). Zoological Journal of the Linnean Society 203 (1): 1-60, DOI: 10.1093/zoolinnean/zlae169, URL: https://doi.org/10.1093/zoolinnean/zlae169
700187A3FFD0FFCDFF39F9DAB710F909.taxon	description	Description: Based on the material examined (NIGP 201161, 201162). Carapace: Anterior margin with few, short microsetae; anteromedian depression narrow, shallow; anterosubmedial depressions absent; lateral ocular carinae absent; posteromedian and posteromarginal sulci distinct; posterior margin slightly recurved medially; surfaces almost smooth to sparsely covered by microsetae (Figs 4 A, 6 A, 9 A). Median ocular tubercle situated anteromedially; median ocelli absent; one pair of large posterolateral major lateral ocelli present (Fig. 6 A). Coxosternal region: Surface smooth, sparsely covered in macrosetae (Fig. 5 B). Sternum not visible. Chelicerae: Fixed finger and manus, prolateral and ventral surfaces each with long, dense setae (Fig. 6 B, C). Moveable finger markedly overlapping fixed finger such that cheliceral dentition partly visible; moveable finger with long dorsal distal (dd) denticle, small dorsal subdistal denticle, stout dorsal median denticle, long ventral distal (vd) denticle, and three or four small ventral accessory (va) denticles; fixed finger with long distal denticle and small subdistal denticle. Moveable finger with prominent serrula comprising moderately long spinules in proximal three-quarters. Pedipalps: Segments gracile (Figs 7, 10). Femur with four carinae evident; prodorsal, proventral, and retrodorsal carinae distinct, serrate; retroventral carinae smooth, costate. Patella with six carinae evident; prolateral surface, dorsoventral ‘ vaulted’ projection (‘ anterior process’) moderately developed with moderately developed pair of apophyses or tubercles (‘ patellar spurs’), each possessing one macroseta; retrodorsal, retromedian, and retroventral carinae distinct, smooth, and costate; prodorsal carina incomplete, slightly granular, and obsolete proximally. Chela relatively gracile (CL / CW = 5.83, Appendix 2); manus globose; chela with five carinae evident; digital carina distinct on fixed finger and manus; prodorsal carina moderately developed, comprising several spiniform granules; promedian carina distinct, comprising five or six prominent spiniform granules; proventral carinae obsolete, comprising few isolated granules; retroventral carina obsolete, smooth, and costate; dorsomedian, dorsal secondary, subdigital, retromedian and ventromedian carinae absent or obsolete. Chela finger dentition not visible. Trichobothrial pattern Type D, β configuration. Femur with 11 trichobothria visible, six d, two i and three e trichobothria, trichobothria d 1, d 2, d 4 and d 6 noticeably smaller than others (‘ petite’); trichobothrium e 1 situated proximal to trichobothrium d 3; trichobothrium e 3 situated dorsal to trichobothrium d 6 (Figs 7 A, 10). Patella with nine trichobothria visible, including three d, one i, and five e trichobothria; trichobothrium i 1 situated distal to trichobothrium d 3 (Figs 7 A, 10). Chela with 12 trichobothria visible; fixed finger with two d, two e, and three i trichobothria; manus with four E and one V trichobothria; trichobothrium Et 1 situated distally on manus, almost aligned with, or proximal to, moveable finger condyle; trichobothrium eb situated on fixed finger, slightly distal to moveable finger condyle; trichobothrium esb situated medially to subdistally on fixed finger; trichobothrium V 2 situated distally on manus and near retroventral carina (Figs 7 B – D, 10). Legs: Tibia lateral and ventral surfaces each with scattered macrosetae (Figs 6 I – L, 9 F – I). Basitarsi each with distinct pedal spurs and pair of spinule rows, comprising c. 10 relatively long spinules, distally. Ungues moderately long, distinctly curved; dactyl pronounced, pointed. Pectines: Three marginal lamellae and four median lamellae (Figs 6 D, 9 J); surface sparsely covered in macrosetae; longitudinal suture between marginal and median lamellae distinct. Fulcra absent or obsolete. Teeth rounded; distal three-quarters of each tooth occupied by sensillar field. Genital operculum: Not visible, covered by fissures and bubbles. Mesosoma: Pretergites surfaces almost smooth (Figs 5 A, 6 E, 9 D); post-tergites I – VI surfaces finely granular, acarinate (Figs 5 A, 6 E, 9 D); tergite VII surface sparsely granular with four distinct carinae (paired dorsosubmedian and dorsolateral carinae) in posterior half of segment (Fig. 6 H). Sternites surfaces smooth, with scattered macrosetae; posterior margins broadly recurved; respiratory spiracles (stigmata) small, ovoid to round (Figs 6 F, 9 E). Metasoma: Segments I – V progressively increasing in length, I wider than long, others longer than wide (Figs 4, 8). Segments I – V lateral and ventral surfaces each with few, short macrosetae (Figs 5 C – F, 6 H, 9 K – N); I – IV dorsal surfaces each with distinct depression (Figs. 6 H, 9 K, L). Segments I – III each with six distinct carinae (paired dorsosubmedian, dorsolateral and ventrolateral carinae), IV and V each with eight distinct carinae (paired dorsosubmedian, dorsolateral, ventrolateral and ventrosubmedian carinae); dorsosubmedian carinae distinct on all segments, markedly serrate, and converging posteriorly; dorsolateral carinae distinct, markedly serrate on segments I – III, moderately serrate on segments IV and V; ventrolateral carinae distinct, moderately serrate on segments I – IV, weakly serrate on V; ventrosubmedian carinae absent on segments I – III, obsolete on IV, distinct, moderately serrate, and converging posteriorly on V (Figs 5 C – F, 6 H, 9 K – N). Telson: Dorsal surface flat; lateral and ventral surfaces relatively smooth, with several macrosetae (Figs 5 E, F, 6 G, 9 M, N); subaculear tubercle absent; prominent, moderately long subaculear setal pair.	en	Xuan, Qiang, Prendini, Lorenzo, Engel, Michael S., Cai, Chenyang, Huang, Diying (2025): Extinct scorpion family Chaerilobuthidae from Mid-Cretaceous Burmese amber reinterpreted as subfamily of extant family Pseudochactidae (Chelicerata: Scorpiones). Zoological Journal of the Linnean Society 203 (1): 1-60, DOI: 10.1093/zoolinnean/zlae169, URL: https://doi.org/10.1093/zoolinnean/zlae169
700187A3FFD0FFCDFF39F9DAB710F909.taxon	materials_examined	Material examined: MYANMAR: Kachin State: Noije Bum near HukawngValley, preciselocalityunknown, Cretaceous (Cenomanian) burmite: 1 juv. (sex unknown), dextral pedipalp patella and chela ab - sent (NIGP 201161), 1 juv. (sex unknown) (NIGP 201162).	en	Xuan, Qiang, Prendini, Lorenzo, Engel, Michael S., Cai, Chenyang, Huang, Diying (2025): Extinct scorpion family Chaerilobuthidae from Mid-Cretaceous Burmese amber reinterpreted as subfamily of extant family Pseudochactidae (Chelicerata: Scorpiones). Zoological Journal of the Linnean Society 203 (1): 1-60, DOI: 10.1093/zoolinnean/zlae169, URL: https://doi.org/10.1093/zoolinnean/zlae169
700187A3FFD9FFD2FCB1F91CB721F833.taxon	description	(Figs 11 – 16, 39)	en	Xuan, Qiang, Prendini, Lorenzo, Engel, Michael S., Cai, Chenyang, Huang, Diying (2025): Extinct scorpion family Chaerilobuthidae from Mid-Cretaceous Burmese amber reinterpreted as subfamily of extant family Pseudochactidae (Chelicerata: Scorpiones). Zoological Journal of the Linnean Society 203 (1): 1-60, DOI: 10.1093/zoolinnean/zlae169, URL: https://doi.org/10.1093/zoolinnean/zlae169
700187A3FFD9FFD2FCB1F91CB721F833.taxon	materials_examined	Type material: Holotype: 1 juv. (sex unknown) (Senckenberg Museum, Frankfurt, Germany), Cretaceous (Cenomanian) burmite: MYANMAR: Kachin State (precise locality unknown).	en	Xuan, Qiang, Prendini, Lorenzo, Engel, Michael S., Cai, Chenyang, Huang, Diying (2025): Extinct scorpion family Chaerilobuthidae from Mid-Cretaceous Burmese amber reinterpreted as subfamily of extant family Pseudochactidae (Chelicerata: Scorpiones). Zoological Journal of the Linnean Society 203 (1): 1-60, DOI: 10.1093/zoolinnean/zlae169, URL: https://doi.org/10.1093/zoolinnean/zlae169
700187A3FFD9FFD2FCB1F91CB721F833.taxon	diagnosis	Diagnosis: The following combination of characters is diagnostic for Chaerilobuthus complexus: carapace anteromedian margin aligned with anterolateral margins (Figs 11 A, 14 A); carapace anterior margin slightly depressed submedially (Figs 12 A, 15 A); lateral ocelli protruding (Figs 12 A, 15 A); sternum relatively narrow (Figs 17 B, 20 B); pedipalp chela robust, ratio of chela length: chela manus width less than 5 (Figs 13 C, 16); pedipalp chela fixed finger median denticle row comprising c. seven oblique subrows (Figs 13 C, 15 E); pectines each with five teeth (Figs 12 B, 15 D); mesosoma relatively wide, tergite VII width much greater than length (Fig 11 E); metasomal segment I width much greater than length (Fig. 12 I, J).	en	Xuan, Qiang, Prendini, Lorenzo, Engel, Michael S., Cai, Chenyang, Huang, Diying (2025): Extinct scorpion family Chaerilobuthidae from Mid-Cretaceous Burmese amber reinterpreted as subfamily of extant family Pseudochactidae (Chelicerata: Scorpiones). Zoological Journal of the Linnean Society 203 (1): 1-60, DOI: 10.1093/zoolinnean/zlae169, URL: https://doi.org/10.1093/zoolinnean/zlae169
700187A3FFD9FFD2FCB1F91CB721F833.taxon	description	Description: Based on the material examined (NIGP 200655, 201160). Carapace: Anterior carapace margin with few, short microsetae; anteromedian depression narrow, shallow; anterosubmedial depressions absent; lateral ocular carinae absent; posteromedian and posteromarginal sulci obsolete; surfaces almost smooth to sparsely covered in microsetae (Figs 12 A, 15 A). Median ocular tubercle situated anteromedially; median ocelli absent; one pair of large posterolateral major lateral ocelli present. Coxosternal region: Surface smooth, sparsely covered in macrosetae (Figs 12 B, 39 D). Leg I maxillary lobes (coxapophyses) distal margins slightly curved, dilate anteriorly and adorned with serrula comprising c. 14 – 16 spinules (Fig. 39 D). Sternum pentagonal; lateral margins sublinear, parallel; ventral surface flat, without distinct concave region or median sulcus; posteromedian depression wide, shallow; posterior margin slightly recurved (Figs 12 B, 15 B). Chelicerae: Fixed finger and manus, prolateral and ventral surfaces each with numerous long, dense macrosetae (Fig. 12 C, G). Moveable finger markedly overlapping fixed finger (Fig. 12 D) such that cheliceral dentition partly visible; moveable finger with long dorsal distal (dd) denticle, small dorsal subdistal denticle, stout dorsal median denticle and long ventral distal (vd) denticle; fixed finger with long distal denticle and small subdistal denticle (Figs 18 E, F, 20 C). Moveable finger with prominent serrula, comprising moderately long spinules in proximal three-quarters (Fig. 12 C, G). Pedipalps: Segments moderately robust; surfaces sparsely covered by short microsetae and few macrosetae (Figs 11, 13, 14, 16). Femur with three carinae evident; prodorsal and proventral carinae distinct, serrate, prodorsal carinae with small apophysis proximally (Figs 13 A, 15 F). Patella with four carinae evident; prolateral surface, dorsoventral ‘ vaulted’ projection (‘ anterior process’) distinct with prominent pair of apophyses or tubercles (‘ patellar spurs’), each possessing one macroseta; retrodorsal and retroventral carinae smooth, costate (Figs 13 B, 15 I). Chela relatively robust; manus globose; chela with six carinae evident; digital carina incomplete, distinct on fixed finger, obsolete on manus; dorsomedian carina reduced to few granules proximally on manus; prodorsal carina distinct, comprising several strong spiniform granules; promedian and proventral carinae obsolete, each comprising few isolated granules; retroventral carina obsolete, smooth, and costate (Figs 13 C, D, 15 H); dorsal secondary, subdigital, retromedian and ventromedian carinae absent or obsolete. Fixed and moveable fingers median denticle rows each comprising seven oblique, slightly imbricate subrows, separated by large serrate prolateral and retrolateral denticles (Figs 13 C, 15 E). Trichobothrial pattern Type D, β configuration. Pedipalp femur with 10 trichobothria visible, five d, two i, and three e trichobothria, trichobothria d 1, d 2, d 4, and d 6 noticeably smaller than others (‘ petite’); trichobothrium e 1 situated proximal to trichobothrium d 3; trichobothrium e 3 vertically aligned with trichobothrium d 6 (Figs 13 A, 16). Patella with four trichobothria visible, including three d and one i trichobothria; trichobothrium i 1 situated distal to trichobothrium d 3 (Figs 13 B, 16). Chela with eight trichobothria visible; fixed finger with two d, two e and one i trichobothria; manus with two E and one V trichobothria; trichobothrium Et 1 situated distally on manus, almost aligned with or proximal to moveable finger condyle; trichobothrium eb situated on fixed finger, slightly distal to moveable finger condyle; trichobothrium esb situated medially on fixed finger; trichobothrium V 2 situated distally on manus and near retroventral carina (Figs 13 C, D, 16). Legs: Tibia lateral and ventral surfaces each with scattered macrosetae (Figs 12 K, L, 15 G). Basitarsi each with pair of spinule rows, comprising seven or eight relatively long spinules, distally (Figs 12 H, L, 15 G). Ungues moderately long, distinctly curved; dactyl pronounced, pointed (Figs 12 K, L, 15 G). Pectines: Basal piece long, narrow (Figs 12 B, 15 D). Three marginal lamellae and four median lamellae; surface sparsely covered in macrosetae; longitudinal suture between marginal and median lamellae distinct. Fulcra absent or obsolete. Teeth rounded, distal half of each tooth occupied by sensilla field; peg sensilla very long, columnar with distinct pore. Genital operculum: Two opercular sclerites narrow and completely separated, as long as pectinal plate (Fig. 12 B). Mesosoma: Pretergites surfaces almost smooth (Fig. 12 E); post-tergites I – VI surfaces finely granular, acarinate (Fig. 12 E); tergite VII surface sparsely granular with four moderately developed carinae (paired dorsosubmedian and dorsolateral carinae) in posterior half of segment (Fig. 11 A). Sternites surfaces smooth, with scattered macrosetae; posterior margins broadly recurved; respiratory spiracles (stigmata) small, ovoid, oblique (Figs 12 F, 15 C). Metasoma: Segments I – III progressively increasing in length, I wider than long, others longer than wide (Fig. 12 I, J). Segments I – III lateral and ventral surfaces each with few, short macrosetae; dorsal surfaces each with distinct depression. Segments I – III each with six distinct carinae (paired dorsosubmedian, dorsolateral and ventrolateral carinae); dorsosubmedian carinae distinct, markedly serrate; dorsolateral carinae distinct, moderately serrate; ventrolateral carinae obsolete on segment I, moderately developed on II and III.	en	Xuan, Qiang, Prendini, Lorenzo, Engel, Michael S., Cai, Chenyang, Huang, Diying (2025): Extinct scorpion family Chaerilobuthidae from Mid-Cretaceous Burmese amber reinterpreted as subfamily of extant family Pseudochactidae (Chelicerata: Scorpiones). Zoological Journal of the Linnean Society 203 (1): 1-60, DOI: 10.1093/zoolinnean/zlae169, URL: https://doi.org/10.1093/zoolinnean/zlae169
700187A3FFD9FFD2FCB1F91CB721F833.taxon	materials_examined	Material examined: MYANMAR: Kachin State: Noije Bum near Hukawng Valley, precise locality unknown, Cretaceous (Cenomanian) burmite: one juv. (sex unknown), mesosomal segments IV, V and telson absent (NIGP 200655), one juv. (sex unknown), sinistral pedipalp absent (NIGP 201160).	en	Xuan, Qiang, Prendini, Lorenzo, Engel, Michael S., Cai, Chenyang, Huang, Diying (2025): Extinct scorpion family Chaerilobuthidae from Mid-Cretaceous Burmese amber reinterpreted as subfamily of extant family Pseudochactidae (Chelicerata: Scorpiones). Zoological Journal of the Linnean Society 203 (1): 1-60, DOI: 10.1093/zoolinnean/zlae169, URL: https://doi.org/10.1093/zoolinnean/zlae169
700187A3FFC1FFD8FF3AF895B73EFB37.taxon	description	(Figs 17 – 21, 39)	en	Xuan, Qiang, Prendini, Lorenzo, Engel, Michael S., Cai, Chenyang, Huang, Diying (2025): Extinct scorpion family Chaerilobuthidae from Mid-Cretaceous Burmese amber reinterpreted as subfamily of extant family Pseudochactidae (Chelicerata: Scorpiones). Zoological Journal of the Linnean Society 203 (1): 1-60, DOI: 10.1093/zoolinnean/zlae169, URL: https://doi.org/10.1093/zoolinnean/zlae169
700187A3FFC1FFD8FF3AF895B73EFB37.taxon	materials_examined	Type material: Holotype: one juv. ♀ (personal collection of Patrick Müller, Käshofen, Germany), Cretaceous (Cenomanian) burmite: MYANMAR: Kachin State (precise locality unknown).	en	Xuan, Qiang, Prendini, Lorenzo, Engel, Michael S., Cai, Chenyang, Huang, Diying (2025): Extinct scorpion family Chaerilobuthidae from Mid-Cretaceous Burmese amber reinterpreted as subfamily of extant family Pseudochactidae (Chelicerata: Scorpiones). Zoological Journal of the Linnean Society 203 (1): 1-60, DOI: 10.1093/zoolinnean/zlae169, URL: https://doi.org/10.1093/zoolinnean/zlae169
700187A3FFC1FFD8FF3AF895B73EFB37.taxon	diagnosis	Diagnosis: The following combination of characters is diagnostic for C. enigmaticus: carapace anteromedian margin slightly posterior to anterolateral margins (Figs 17 A and 18 D); lateral ocelli not protruding, macular (Figs 18 D, 20 A); pedipalp chela moderately robust, manus globose, ratio of chela length: chela manus width less than 4 (Figs 18 A, B, 21 A); pectines each with five teeth (Figs 18 I, 20 F, G); ratio of metasomal segment IV: V length greater than 2 (Figs 18 A, B, 20 H, I, L); metasomal segment V dorsosubmedian carinae slightly serrate (Fig. 20 L); telson vesicle bulbous, aculeus very short and moderately curved (Figs 18 G, J, K, 19).	en	Xuan, Qiang, Prendini, Lorenzo, Engel, Michael S., Cai, Chenyang, Huang, Diying (2025): Extinct scorpion family Chaerilobuthidae from Mid-Cretaceous Burmese amber reinterpreted as subfamily of extant family Pseudochactidae (Chelicerata: Scorpiones). Zoological Journal of the Linnean Society 203 (1): 1-60, DOI: 10.1093/zoolinnean/zlae169, URL: https://doi.org/10.1093/zoolinnean/zlae169
700187A3FFC1FFD8FF3AF895B73EFB37.taxon	description	Description: Based on the material examined (NIGP 201158, 201159). Carapace: Posterior width less than length (L / PW = 1.16, Appendix 2). Anterior margin with few, short microsetae; anteromedian depression not visible; anterosubmedial depressions absent; lateral ocular carinae absent; posteromedian and posteromarginal sulci obsolete; surfaces almost smooth to sparsely and finely granular (Fig. 20 A, B). Median ocular tubercle situated anteromedially; median ocelli absent; one pair of large posterolateral major lateral ocelli present. Coxosternalregion: Surfacesmooth, sparselycoveredinmacrosetae (Figs 20 E, 39 B, C). Leg I maxillary lobes (coxapophyses) distal margins slightly curved, dilate anteriorly and adorned with serrula comprising c. 15 denticles (Fig. 39 B, C). Sternum pentagonal; lateral margins sublinear, parallel; ventral surface flat, without distinct concave region or median sulcus; posteromedian depression wide, shallow (Figs 17 A, 20 E). Chelicerae: Fixed finger and manus, prolateral and ventral surfaces each with numerous long, dense macrosetae (Fig. 18 E, F). Cheliceral dentition partly visible, moveable finger with long dorsal distal (dd) denticle, small dorsal subdistal denticle, stout dorsal median denticle, long ventral distal (vd) denticle and four or five small ventral accessory (va) denticles; fixed finger with long distal denticle and small subdistal denticle (Figs 18 E, F, 20 C). Moveable finger with prominent serrula comprising long spinules in proximal three-quarters (Fig. 18 F). Pedipalps: Segments moderately robust; surfaces sparsely covered by short microsetae and few macrosetae (Figs 18 A, B, 21 A). Femur with four carinae evident; prodorsal and proventral carinae distinct, serrate; retrodorsal and retroventral carinae smooth, costate (Figs 18 A, 21 A, C). Patella with five carinae evident; prodorsal carina obsolete, comprising several tubercles; prolateral surface, dorsoventral ‘ vaulted’ projection (‘ anterior process’) distinct with prominent pair of apophyses or tubercles (‘ patellar spurs’), each possessing one macroseta; retrodorsal and retroventral carinae smooth, costate (Figs 18 A, B, 21 A, D). Chela relatively robust (CL / CW = 3.22 or 3.55, Appendix 2); manus globose; chela with six carinae evident; digital carina complete, distinct, granular medially, becoming costate proximally and distally; dorsomedian carina reduced to few granules at base of fixed finger, becoming obsolete proximally on manus; prodorsal carina distinct, comprising several prominent spiniform granules; promedian and proventral carinae obsolete, each comprising few isolated granules; retroventral carina obsolete; dorsal secondary, subdigital, retromedian and ventromedian carinae absent or obsolete (Figs 18 A, B, 21 A, B, E, F). Only fixed finger distal denticle subrows visible, oblique, slightly imbricate and separated by large serrate prolateral and retrolateral denticles (Fig. 21 A). Trichobothrial pattern Type D, β configuration. Femur with three trichobothria visible: two d and one i trichobothria (Fig. 21 C). Patella with six trichobothria visible, including three d, one i, and two e trichobothria; trichobothrium i 1 situated distal to trichobothrium d 3 (Fig. 21 A, D). Chela with 10 trichobothria visible; fixed finger with two d and one i trichobothria; manus with four E and one V trichobothria; trichobothrium Et 1 situated distally on manus, almost aligned with or proximal to moveable finger condyle; trichobothrium eb situated on fixed finger, slightly distal to moveable finger condyle; trichobothrium esb situated medially on fixed finger; trichobothrium V 2 situated distally on manus and near retroventral carina. Legs: Tibia lateral and ventral surfaces each with scattered macrosetae (Fig. 18 C, 20 J, K). Basitarsi each with pair of spinule rows, comprising c. 10 relatively long spinules, distally. Telotarsi each with slightly irregular pair of ventrosubmedian spinule rows, comprising relatively long spinules. Ungues moderately long, distinctly curved; dactyl pronounced, pointed. Pectines: Basal piece long, narrow (Fig. 20 F). Three marginal lamellae and four median lamellae; surface sparsely covered in macrosetae; longitudinal suture between marginal and median lamellae distinct (Figs 18 I, 20 G). Fulcra absent or obsolete. Teeth rounded, distal three-quarters of each tooth occupied by sensillar field; peg sensilla very long, columnar (Fig. 20 G). Genital operculum: Two opercular sclerites narrow and completely separated, as long as pectinal plate; surfaces with several microsetae (Fig. 20 E). Mesosoma: Pretergites surfaces almost smooth (Fig. 19 A); post-tergites I – VI surfaces finely granular, acarinate (Fig. 19 A); tergite VII surface sparsely granular, with four slightly serrate carinae (paired dorsosubmedian and dorsolateral carinae) in posterior half of segment (Fig. 20 H). Sternites surfaces smooth, with scattered macrosetae; posterior margins broadly recurved; respiratory spiracles small, ovoid, oblique (Figs 18 H, 20 D). Metasoma: Segments I – V progressively increasing in length, I wider than long, others longer than wide (Fig. 18 J, K, 20 H, I, L). Segments I – V lateral and ventral surfaces each with few, short macrosetae (Fig. 18 J, K, 20 H, I, L); I – IV dorsal surfaces each with distinct depression (Figs. 18 K, 20 H). Segment I with six distinct carinae (paired dorsosubmedian, dorsolateral, and ventrolateral carinae), segments II – V each with eight distinct carinae (paired dorsosubmedian, dorsolateral, ventrolateral, and ventrosubmedian carinae); dorsosubmedian carinae distinct on all segments, markedly serrate on II – IV, moderately serrate, converging posteriorly on I and V; dorsolateral carinae distinct, moderately serrate on segments I, II and V, slightly serrate on III and IV; ventrolateral carinae distinct, moderately serrate on segments I – IV, markedly serrate on segment V; ventrosubmedian carinae absent on segment I, obsolete on II and III, distinct, moderately serrate on IV and V, and converging posteriorly on V (Fig. 18 J, K, 20 H, I, L). Telson: Vesicle bulbous; dorsal surface flat; lateral and ventral surfaces relatively smooth with several macrosetae; (Figs 18 G, 19 A, B); subaculear tubercle absent; prominent subaculear setal pair, moderately long. Aculeus short, shallowly curved; base narrow, less than one-third vesicle width.	en	Xuan, Qiang, Prendini, Lorenzo, Engel, Michael S., Cai, Chenyang, Huang, Diying (2025): Extinct scorpion family Chaerilobuthidae from Mid-Cretaceous Burmese amber reinterpreted as subfamily of extant family Pseudochactidae (Chelicerata: Scorpiones). Zoological Journal of the Linnean Society 203 (1): 1-60, DOI: 10.1093/zoolinnean/zlae169, URL: https://doi.org/10.1093/zoolinnean/zlae169
700187A3FFC1FFD8FF3AF895B73EFB37.taxon	materials_examined	Material examined: MYANMAR: Kachin State: Noije Bum near HukawngValley, preciselocalityunknown, Cretaceous (Cenomanian) burmite: two juv. (sex unknown) (NIGP 201158, 201159).	en	Xuan, Qiang, Prendini, Lorenzo, Engel, Michael S., Cai, Chenyang, Huang, Diying (2025): Extinct scorpion family Chaerilobuthidae from Mid-Cretaceous Burmese amber reinterpreted as subfamily of extant family Pseudochactidae (Chelicerata: Scorpiones). Zoological Journal of the Linnean Society 203 (1): 1-60, DOI: 10.1093/zoolinnean/zlae169, URL: https://doi.org/10.1093/zoolinnean/zlae169
700187A3FFCCFFDFFCA3FB58B69CFB28.taxon	description	(Figs 22 – 24)	en	Xuan, Qiang, Prendini, Lorenzo, Engel, Michael S., Cai, Chenyang, Huang, Diying (2025): Extinct scorpion family Chaerilobuthidae from Mid-Cretaceous Burmese amber reinterpreted as subfamily of extant family Pseudochactidae (Chelicerata: Scorpiones). Zoological Journal of the Linnean Society 203 (1): 1-60, DOI: 10.1093/zoolinnean/zlae169, URL: https://doi.org/10.1093/zoolinnean/zlae169
700187A3FFCCFFDFFCA3FB58B69CFB28.taxon	materials_examined	Type material: Holotype: one juv. (sex unknown) (Geologisch-Paläontologisches Institut, University of Hamburg, Germany: 11037 - 1 GPIH 4566), Cretaceous (Cenomanian) burmite: MYANMAR: Kachin State (precise locality unknown).	en	Xuan, Qiang, Prendini, Lorenzo, Engel, Michael S., Cai, Chenyang, Huang, Diying (2025): Extinct scorpion family Chaerilobuthidae from Mid-Cretaceous Burmese amber reinterpreted as subfamily of extant family Pseudochactidae (Chelicerata: Scorpiones). Zoological Journal of the Linnean Society 203 (1): 1-60, DOI: 10.1093/zoolinnean/zlae169, URL: https://doi.org/10.1093/zoolinnean/zlae169
700187A3FFCCFFDFFCA3FB58B69CFB28.taxon	diagnosis	Diagnosis: The following combination of characters is diagnostic for C. gigantosternum: carapace relatively long (L / Pw = 1.25); carapace anteromedian margin slightly convex and anterosubmedian margin slightly concave (Figs 22 A, 23 A); carapace posterior margin linear; lateral ocelli not protruding, macular (Fig. 23 A); sternum relatively wide; cheliceral moveable finger with distinct serrula; pedipalp chela slender, manus elongate, with ratio of chela length: chela manus width greater than 4 (Fig. 24); pectines relatively large, each with five teeth (Fig. 23 F); ratio of metasomal segment IV: V length less than 2 (Fig. 23 H, I).	en	Xuan, Qiang, Prendini, Lorenzo, Engel, Michael S., Cai, Chenyang, Huang, Diying (2025): Extinct scorpion family Chaerilobuthidae from Mid-Cretaceous Burmese amber reinterpreted as subfamily of extant family Pseudochactidae (Chelicerata: Scorpiones). Zoological Journal of the Linnean Society 203 (1): 1-60, DOI: 10.1093/zoolinnean/zlae169, URL: https://doi.org/10.1093/zoolinnean/zlae169
700187A3FFCCFFDFFCA3FB58B69CFB28.taxon	description	Description: Based on the material examined (NIGP 200654). Carapace: Anterior margin of carapace with few, short microsetae; anteromedian depression not visible; anterosubmedial depressions absent; lateral ocular carinae distinct; posteromedian and posteromarginal sulci not distinct; surfaces almost smooth to sparsely and finely granular (Fig. 23 A). Median ocular tubercle situated anteromedially; median ocelli absent; one pair of large posterolateral major lateral ocelli present. Coxosternal region: Surface smooth, sparsely covered in few macrosetae (Fig. 23 B). Distal margins of leg I maxillary lobes (coxapophyses) slightly curved and dilate anteriorly, adorned with serrula comprising c. 13 spiny denticles. Sternum pentagonal; lateral margins sublinear, parallel, ventral surface flat, without distinct concave region or median sulcus; posteromedian depression wide, shallow. Chelicerae: Fixed finger and manus, prolateral and ventral surfaces each with numerous macrosetae. Cheliceral dentition partly visible, moveable finger strongly overlapping fixed finger; moveable finger with long dorsal distal (dd) denticle, small dorsal subdistal denticle, stout dorsal median denticle and long ventral distal (vd) denticle; fixed finger with long distal denticle. Moveable finger with prominent serrula, comprising c. 10 long spinules and extending almost entire length of ventral surface (Fig. 23 C). Pedipalps: Segments gracile (Figs 22, 23). Femur with three carinae evident; prodorsal, proventral and retrodorsal carinae distinct, smooth and costate. Patella with three carinae evident; retroventral carinae distinct, smooth and costate; prolateral surface, dorsoventral ‘ vaulted’ projection (‘ anterior process’) distinct, with obsolete pair of apophyses or tubercles (‘ patellar spurs’), each possessing one macroseta. Chela relatively slender (CL / CW = 4.37, Appendix 2); manus elongate; chela with three carinae evident; digital carina complete, distinct, granular medially, becoming costate proximally and distally; prodorsal carina distinct, smooth; retroventral carina obsolete, smooth and costate; other carinae absent or obsolete. Chela finger dentition not visible. Trichobothrial pattern Type D, β configuration (Fig. 24). Femur with 10 trichobothria visible, five d, three e, and two i trichobothria, trichobothrium e 1 situated proximal to trichobothrium d 3; trichobothrium e 3 situated dorsal to trichobothrium d 6. Patella with eight trichobothria visible, including three d, one i, and four e trichobothria; trichobothrium i 1 situateddistaltotrichobothrium d 3. Chelawith 12 trichobothria visible; fixed finger with two d, two e, and three i trichobothria; manus with four E and one V trichobothria; trichobothrium Et 1 situated distally on manus, almost aligned with or proximal to moveable finger condyle; trichobothrium eb situated on fixed finger, slightly distal to moveable finger condyle; trichobothrium esb situated medially on fixed finger; trichobothrium V 2 situated distally on manus and near retroventral carina. Legs: Tibial lateral and ventral surfaces each with scattered macrosetae (Fig. 23 J, K). Basitarsi each with pair of spinule rows, comprising c. 12 relatively long spinules, distally. Telotarsi each with slightly irregular pair of ventrosubmedian spinule rows, comprising relatively long spinules. Ungues moderately long, distinctly curved; dactyl pronounced, pointed. Pectines: Basal piece not visible. Three marginal lamellae and four median lamellae; surface sparsely covered in macrosetae; longitudinal suture between marginal and median lamellae distinct (Fig. 23 E, F). Fulcra absent or obsolete. Teeth rounded, distal three-quarters of each tooth occupied by sensillar field; peg sensilla very long and columnar. Genital operculum: Opercular sclerites not visible. Mesosoma: Pretergites surfaces almost smooth (Fig. 23 D); post-tergites I – VI surfaces finely granular, acarinate (Fig. 23 D); tergite VII surface sparsely granular with four carinae (paired dorsosubmedian and dorsolateral carinae) in posterior half of segment (Fig. 23 D). Sternites surfaces smooth, with scattered macrosetae; respiratory spiracles (stigmata) small, ovoid to round (Fig. 23 E). Metasoma: Segments I – V progressively increasing in length, I wider than long, others longer than wide (Fig. 23 H, I). Segments I – V lateral and ventral surfaces each with few, short macrosetae; I – IV dorsal surfaces each with distinct depression. Segments I – III each with six distinct carinae (paired dorsosubmedian, dorsolateral, and ventrolateral carinae), IV and V each with eight distinct carinae (paired dorsosubmedian, dorsolateral, ventrolateral, and ventrosubmedian carinae); dorsosubmedian and dorsolateral carinae distinct, slightly serrate; ventrolateral carinae distinct, costate-granular; ventrosubmedian carinae obsolete.	en	Xuan, Qiang, Prendini, Lorenzo, Engel, Michael S., Cai, Chenyang, Huang, Diying (2025): Extinct scorpion family Chaerilobuthidae from Mid-Cretaceous Burmese amber reinterpreted as subfamily of extant family Pseudochactidae (Chelicerata: Scorpiones). Zoological Journal of the Linnean Society 203 (1): 1-60, DOI: 10.1093/zoolinnean/zlae169, URL: https://doi.org/10.1093/zoolinnean/zlae169
700187A3FFCCFFDFFCA3FB58B69CFB28.taxon	materials_examined	Material examined: MYANMAR: Kachin State: Noije Bum near Hukawng Valley, precise locality unknown, Cretaceous (Cenomanian) burmite: first instar juv. (sex unknown) (NIGP 200654). Hypothesized to be in first instar based on small size and weak development of carinae and sulci.	en	Xuan, Qiang, Prendini, Lorenzo, Engel, Michael S., Cai, Chenyang, Huang, Diying (2025): Extinct scorpion family Chaerilobuthidae from Mid-Cretaceous Burmese amber reinterpreted as subfamily of extant family Pseudochactidae (Chelicerata: Scorpiones). Zoological Journal of the Linnean Society 203 (1): 1-60, DOI: 10.1093/zoolinnean/zlae169, URL: https://doi.org/10.1093/zoolinnean/zlae169
700187A3FFCBFFE0FCA5FB6DB6D4FC13.taxon	description	(Figs 25 – 26, 39)	en	Xuan, Qiang, Prendini, Lorenzo, Engel, Michael S., Cai, Chenyang, Huang, Diying (2025): Extinct scorpion family Chaerilobuthidae from Mid-Cretaceous Burmese amber reinterpreted as subfamily of extant family Pseudochactidae (Chelicerata: Scorpiones). Zoological Journal of the Linnean Society 203 (1): 1-60, DOI: 10.1093/zoolinnean/zlae169, URL: https://doi.org/10.1093/zoolinnean/zlae169
700187A3FFCBFFE0FCA5FB6DB6D4FC13.taxon	materials_examined	Type material: Holotype: one juv. ♀ (personal collection of Hans-Georg Müller, Gelsenkirchen, Germany), Cretaceous (Cenomanian) burmite: MYANMAR: Kachin State (precise locality unknown).	en	Xuan, Qiang, Prendini, Lorenzo, Engel, Michael S., Cai, Chenyang, Huang, Diying (2025): Extinct scorpion family Chaerilobuthidae from Mid-Cretaceous Burmese amber reinterpreted as subfamily of extant family Pseudochactidae (Chelicerata: Scorpiones). Zoological Journal of the Linnean Society 203 (1): 1-60, DOI: 10.1093/zoolinnean/zlae169, URL: https://doi.org/10.1093/zoolinnean/zlae169
700187A3FFCBFFE0FCA5FB6DB6D4FC13.taxon	diagnosis	Diagnosis: Thefollowingcombinationofcharactersisdiagnosticfor C. hansgeorgmuelleri: carapace length less than posterior width (Fig. 26 A); carapace anterior margin slightly depressed submedially (Fig. 26 A); lateral ocelli protruding (Fig. 26 A); sternum relatively wide (Fig. 26 B); pectines each with five teeth (Fig. 26 B); pedipalp patella trichobothrium i aligned with trichobothrium d 3 (Fig. 26 F); pedipalp chela robust, manus globose, ratio of chela length: chela manus width less than 3 (Fig. 26 C).	en	Xuan, Qiang, Prendini, Lorenzo, Engel, Michael S., Cai, Chenyang, Huang, Diying (2025): Extinct scorpion family Chaerilobuthidae from Mid-Cretaceous Burmese amber reinterpreted as subfamily of extant family Pseudochactidae (Chelicerata: Scorpiones). Zoological Journal of the Linnean Society 203 (1): 1-60, DOI: 10.1093/zoolinnean/zlae169, URL: https://doi.org/10.1093/zoolinnean/zlae169
700187A3FFCBFFE0FCA5FB6DB6D4FC13.taxon	description	Description: Based on the material examined (NIGP 201164). Carapace: Length less than posterior width (L / PW = 0.94). Anteromedian depression narrow, shallow; anterosubmedial depressions absent; lateral ocular carinae absent; posteromedian and posteromarginal sulci distinct; surfaces almost smooth to sparsely and finely granular (Fig. 26 A). Median ocular tubercle situated anteromedially; median ocelli absent; one pair of large posterolateral major lateral ocelli present. Coxosternal region: Surface smooth, sparsely covered in macrosetae (Figs 25 B, 39 G). Leg I maxillary lobes (coxapophyses) distal margins slightly curved, dilate anteriorly and adorned with serrula comprising c. 15 – 25 denticles (Fig. 39 G). Sternum broad, pentagonal; lateral margins sublinear, parallel; ventral surface flat, without distinct concave region or median sulcus; posteromedian depression wide, shallow (Fig. 26 B). Chelicerae: Fixed finger and manus, prolateral and ventral surfaces each with numerous long, dense macrosetae (Fig. 25 A). Cheliceral dentition not clearly visible. Pedipalps: Segments moderately robust; surfaces sparsely covered by short microsetae (Fig. 25 A). Femur with three carinae evident; dorsalsurfacewithseveralcoarsegranules; prodorsalandproventral carinae distinct, serrate; retrodorsal carinae distinct, smooth, and costate (Fig. 26 D). Patella with five carinae evident; prodorsal carina distinct, costate-granular; prolateral surface, dorsoventral ‘ vaulted’ projection (‘ anterior process’) distinct with moderately developed pair of apophyses or tubercles (‘ patellar spurs’), each possessing one macroseta; retrodorsal and retroventral carinae smooth, costate (Figs 25 B, 26 F). Chela relatively robust (CL / CW = 2.62, Appendix 2); manus globose; chela with six carinae evident; digital carina complete, distinct; dorsomedian carina moderately distinct, costate at base of fixed finger to medially on manus, becoming obsolete proximally on manus; prodorsal carina distinct, comprising several prominent spiniform granules, promedian and proventral carinae obsolete, each comprising few isolated granules; retroventral carina obsolete; dorsal secondary, subdigital, retromedian, and ventromedian carinae absent or obsolete (Figs 25 A, B, 26 C). Chela finger dentition not visible. Trichobothrial pattern Type D, β configuration. Femur with 11 trichobothria, five d, three i, and three e trichobothria; trichobothrium e 1 situated proximal to trichobothrium d 3; trichobothrium e 3 situated dorsal to trichobothrium d 6 (Fig. 26 D). Patella with 10 trichobothria, including three d, one i, and two e trichobothria; trichobothrium i 1 aligned with trichobothrium d 3 (Fig. 26 F). Chela with 12 trichobothria; fixed finger with two d, three i, and two e trichobothria; manus with four e and one v trichobothria; trichobothrium Et 1 situated distally on manus, almost aligned with, or proximal to, moveable finger condyle; trichobothrium eb situated on fixed finger, slightly distal to moveable finger condyle; trichobothrium esb situated medially on fixed finger; trichobothrium V 2 situated distally on manus and near retroventral carina (Fig. 26 C). Pectines: Basal piece not visble (Fig. 26 B). Three marginal lamellae and four median lamellae; longitudinal suture between marginal and median lamellae distinct. Fulcra absent or obsolete. Teeth rounded, peg sensilla not visible. Genital operculum: Two opercular sclerites narrow and completely separated (Fig. 26 B).	en	Xuan, Qiang, Prendini, Lorenzo, Engel, Michael S., Cai, Chenyang, Huang, Diying (2025): Extinct scorpion family Chaerilobuthidae from Mid-Cretaceous Burmese amber reinterpreted as subfamily of extant family Pseudochactidae (Chelicerata: Scorpiones). Zoological Journal of the Linnean Society 203 (1): 1-60, DOI: 10.1093/zoolinnean/zlae169, URL: https://doi.org/10.1093/zoolinnean/zlae169
700187A3FFCBFFE0FCA5FB6DB6D4FC13.taxon	materials_examined	Material examined: MYANMAR: Kachin State: Noije Bum near Hukawng Valley, precise locality unknown, Cretaceous (Cenomanian) burmite: one subad. (sex unknown), mesosoma and metasoma absent (NIGP 201164).	en	Xuan, Qiang, Prendini, Lorenzo, Engel, Michael S., Cai, Chenyang, Huang, Diying (2025): Extinct scorpion family Chaerilobuthidae from Mid-Cretaceous Burmese amber reinterpreted as subfamily of extant family Pseudochactidae (Chelicerata: Scorpiones). Zoological Journal of the Linnean Society 203 (1): 1-60, DOI: 10.1093/zoolinnean/zlae169, URL: https://doi.org/10.1093/zoolinnean/zlae169
700187A3FFF4FFE7FC52FC7DB38FF983.taxon	description	(Figs 27 – 31, 39)	en	Xuan, Qiang, Prendini, Lorenzo, Engel, Michael S., Cai, Chenyang, Huang, Diying (2025): Extinct scorpion family Chaerilobuthidae from Mid-Cretaceous Burmese amber reinterpreted as subfamily of extant family Pseudochactidae (Chelicerata: Scorpiones). Zoological Journal of the Linnean Society 203 (1): 1-60, DOI: 10.1093/zoolinnean/zlae169, URL: https://doi.org/10.1093/zoolinnean/zlae169
700187A3FFF4FFE7FC52FC7DB38FF983.taxon	materials_examined	Type material: Holotype: one juv. (sex unknown) (personal collection of KlausPeter Brucksch, Kuranda, Australia), Cretaceous (Cenomanian) burmite: MYANMAR: Kachin State (precise locality unknown).	en	Xuan, Qiang, Prendini, Lorenzo, Engel, Michael S., Cai, Chenyang, Huang, Diying (2025): Extinct scorpion family Chaerilobuthidae from Mid-Cretaceous Burmese amber reinterpreted as subfamily of extant family Pseudochactidae (Chelicerata: Scorpiones). Zoological Journal of the Linnean Society 203 (1): 1-60, DOI: 10.1093/zoolinnean/zlae169, URL: https://doi.org/10.1093/zoolinnean/zlae169
700187A3FFF4FFE7FC52FC7DB38FF983.taxon	diagnosis	Diagnosis: The following combination of characters is diagnostic for C. knodelorum: carapace anterior margin sublinear (Figs 28 A, 29 A); lateral ocelli not protruding, macular (Fig. 29 A); cheliceral finger ventral surface with serrula extending almost entire length (Fig. 29 B); pedipalp chela relatively wide, ratio of chela length: width less than 4 (Figs 28 C, F, 29 E); leg telotarsi ventrosubmedian rows with very long spinules (Fig. 29 G, H, J); pectinal lamellae relatively wide, pectines each with five, relatively large teeth (Fig. 29 C); mesosoma long, ratio of mesosoma: carapace length greater than 2.5.	en	Xuan, Qiang, Prendini, Lorenzo, Engel, Michael S., Cai, Chenyang, Huang, Diying (2025): Extinct scorpion family Chaerilobuthidae from Mid-Cretaceous Burmese amber reinterpreted as subfamily of extant family Pseudochactidae (Chelicerata: Scorpiones). Zoological Journal of the Linnean Society 203 (1): 1-60, DOI: 10.1093/zoolinnean/zlae169, URL: https://doi.org/10.1093/zoolinnean/zlae169
700187A3FFF4FFE7FC52FC7DB38FF983.taxon	description	Description: Based on the material examined (NIGP 200657). Carapace: Anterior margin with few, short microsetae; anteromedian depression narrow, shallow; anterosubmedial depressions absent (Fig. 28 A); lateral ocular carinae absent; posteromedian and posteromarginal sulci distinct (Fig. 30 A); posterior margin slightly recurved medially (Figs 28 A, 29 A); surfaces almost smooth to sparsely and finely granular (Figs. 28 A). Median ocular tubercle situated anteromedially; median ocelli absent; one pair of large posterolateral major lateral ocelli present. Coxosternal region: Surface smooth, sparsely covered in macrosetae (Fig. 28 B). Leg I maxillary lobes (coxapophyses) distal margins slightly curved and dilate anteriorly, adorned with serrula (Fig. 39 E). Sternum pentagonal; lateral margins sublinear, parallel; posterior margin slightly recurved; ventral surface flat, without distinct concave region or median sulcus; posteromedian depression relatively wide, shallow (Fig. 30 B). Chelicerae: Cheliceral dentition partly visible; moveable finger with long dorsal distal (dd) denticle, small dorsal subdistal denticle, stout dorsal median denticle and long ventral distal (vd) denticle; fixed finger with long distal denticle and small subdistal denticle (Fig. 28 A). Moveable finger with prominent serrula, comprising moderately long spinules and extending almost entire length of ventral surface (Fig. 29 B). Pedipalps: Segments gracile; surfaces finely granular (Fig. 27). Femur with three carinae evident; prodorsal carina distinct, markedly serrate; retrodorsal carina distinct, slightly serrate; proventral carina smooth, costate (Figs 28 F, 31). Patella with six carinae evident; prolateral surface, dorsoventral ‘ vaulted’ projection (‘ anterior process’) distinct with obsolete pair of apophyses or tubercles (‘ patellar spurs’), each possessing one macroseta; retrodorsal, retromedian, and retroventral carinae distinct, smooth and costate; prodorsal carina obsolete (Figs 29 D, 31). Chela relatively robust (CL / CW = 3.75, Appendix 2); manus globose; chela with five carinae evident; digital carina incomplete, distinct on fixed finger, obsolete on manus; prodorsal carina distinct, markedly serrate; promedian carina distinct, comprising several prominent spiniform granules distally; retroventral carina obsolete, smooth and costate (Figs 28 C, F, 29 E, 31); dorsomedian, dorsal secondary, subdigital, proventral, retromedian and ventromedian carinae absent or obsolete. Fixed and moveable fingers median denticle rows visible distally, each comprising oblique, slightly imbricate subrows of c. 13 small denticles, separated by large serrate prolateral and retrolateral denticles (Fig. 29 F). Trichobothrial pattern Type D, β configuration (Fig. 31). Pedipalp femur with four trichobothria visible, two d and two e trichobothria. Patella with nine trichobothria visible, including two d, one i, and six e trichobothria; trichobothrium i 1 situated distal to trichobothrium d 3. Chela with 11 trichobothria visible; fixed finger with one d, two e, and three i trichobothria; manus with four E and one V trichobothria; trichobothrium Et 1 situated distally on manus, almost aligned with or proximal to moveable finger condyle; trichobothrium eb situated on fixed finger, slightly distal to moveable finger condyle; trichobothrium esb situated submedially to slightly distally on fixed finger; trichobothrium V 2 situated distally on manus and near retroventral carina. Legs: Tibia lateral and ventral surfaces each with scattered macrosetae (Fig. 29 H – J). Basitarsi each with prominent pedal spurs (Fig. 29 G – J) and pair of spinule rows, each comprising c. 10 relatively long spinules, distally (Fig. 29 G – J). Ungues moderately long, distinctly curved; dactyl pronounced, pointed (Fig. 29 H – J). Pectines: Pectines very short, not extending beyond coxa of leg IV (Fig. 29 C, 30 B). Three marginal lamellae and four median lamellae; surface sparsely covered in macrosetae; longitudinal suture between marginal and median lamellae distinct. Fulcra absent or obsolete. Teeth elongate, distal three-quarters of each tooth occupied by sensillar field. Genital operculum: Two opercular sclerites relatively wide, as long as pectinal plate, and completely separated (Figs 28 B, 30). Mesosoma: Pretergites surfaces almost smooth; post-tergites I – VI surfaces finely granular, acarinate (Figs 27 A, 28 A); sinistral dorsosubmedian and dorsolateral carinae of tergite VII visible (Fig. 27 A). Sternites surfaces smooth, with scattered macrosetae; posterior margin of sternite I broadly recurved (Fig. 27 B); respiratory spiracles (stigmata) not visible. Metasoma: Segments I – IV progressively increasing in length, I wider than long, others longer than wide (Figs 27 A, B, 28 D, E). Segments I – IV lateral and ventral surfaces each with few, short macrosetae; dorsalsurfaceseachwithdistinctdepression. Segments I and II each with six distinct carinae (paired dorsosubmedian, dorsolateral, and ventrolateral carinae), III and IV each with eight distinct carinae (paired dorsosubmedian, dorsolateral, ventrolateral, and ventrosubmedian carinae); dorsosubmedian carinae distinct, crenulate, and converging posteriorly on segments I – IV; dorsolateral carinae distinct, slightly serrate on segments I – III, moderately serrate on IV; ventrolateral carinae distinct, slightly serrate on segments I – IV; ventrosubmedian carinae absent on segments I and II, obsolete on III and IV.	en	Xuan, Qiang, Prendini, Lorenzo, Engel, Michael S., Cai, Chenyang, Huang, Diying (2025): Extinct scorpion family Chaerilobuthidae from Mid-Cretaceous Burmese amber reinterpreted as subfamily of extant family Pseudochactidae (Chelicerata: Scorpiones). Zoological Journal of the Linnean Society 203 (1): 1-60, DOI: 10.1093/zoolinnean/zlae169, URL: https://doi.org/10.1093/zoolinnean/zlae169
700187A3FFF4FFE7FC52FC7DB38FF983.taxon	discussion	Remarks: The specimen examined possesses a relatively small pair of pectines, unlike the larger pectines of the holotype specimen, suggesting that it may be female.	en	Xuan, Qiang, Prendini, Lorenzo, Engel, Michael S., Cai, Chenyang, Huang, Diying (2025): Extinct scorpion family Chaerilobuthidae from Mid-Cretaceous Burmese amber reinterpreted as subfamily of extant family Pseudochactidae (Chelicerata: Scorpiones). Zoological Journal of the Linnean Society 203 (1): 1-60, DOI: 10.1093/zoolinnean/zlae169, URL: https://doi.org/10.1093/zoolinnean/zlae169
700187A3FFF4FFE7FC52FC7DB38FF983.taxon	materials_examined	Material examined: MYANMAR: Kachin State: Noije Bum near Hukawng Valley, precise locality unknown, Cretaceous (Cenomanian) burmite: one juv. [♀?], metasomal segment IV and telson absent (NIGP 200657).	en	Xuan, Qiang, Prendini, Lorenzo, Engel, Michael S., Cai, Chenyang, Huang, Diying (2025): Extinct scorpion family Chaerilobuthidae from Mid-Cretaceous Burmese amber reinterpreted as subfamily of extant family Pseudochactidae (Chelicerata: Scorpiones). Zoological Journal of the Linnean Society 203 (1): 1-60, DOI: 10.1093/zoolinnean/zlae169, URL: https://doi.org/10.1093/zoolinnean/zlae169
700187A3FFF3FFEDFF2EF99EB1FFFA27.taxon	description	(Figs 32 – 39)	en	Xuan, Qiang, Prendini, Lorenzo, Engel, Michael S., Cai, Chenyang, Huang, Diying (2025): Extinct scorpion family Chaerilobuthidae from Mid-Cretaceous Burmese amber reinterpreted as subfamily of extant family Pseudochactidae (Chelicerata: Scorpiones). Zoological Journal of the Linnean Society 203 (1): 1-60, DOI: 10.1093/zoolinnean/zlae169, URL: https://doi.org/10.1093/zoolinnean/zlae169
700187A3FFF3FFEDFF2EF99EB1FFFA27.taxon	materials_examined	Type material: Holotype: one juv. (sex unknown) (Geologisch-Paläontologisches Institut, University of Hamburg, Germany: 11037 - 2 GPIH 4567), Cretaceous (Cenomanian) burmite: MYANMAR: Kachin State (precise locality unknown).	en	Xuan, Qiang, Prendini, Lorenzo, Engel, Michael S., Cai, Chenyang, Huang, Diying (2025): Extinct scorpion family Chaerilobuthidae from Mid-Cretaceous Burmese amber reinterpreted as subfamily of extant family Pseudochactidae (Chelicerata: Scorpiones). Zoological Journal of the Linnean Society 203 (1): 1-60, DOI: 10.1093/zoolinnean/zlae169, URL: https://doi.org/10.1093/zoolinnean/zlae169
700187A3FFF3FFEDFF2EF99EB1FFFA27.taxon	diagnosis	Diagnosis: The following combination of characters is diagnostic for C. serratus: carapace anteromedian margin aligned with anterolateral margins (Figs 33 A, 34 A, 37 A); lateral ocelli not protruding, macular (Figs 34 A, C, 37 C); pedipalp chela slender, ratio of chela length: chela manus width greater than 4 (Figs 35 A, E, 38); pectines each with five teeth (Figs 34 G, 37 H); ratio of metasomal segment IV: segment V length less than 2 (Figs 33 I, J, 37 G); metasomal segment V dorsosubmedian carinae distinctly serrate (Fig. 32 I, J); telson vesicle slightly elongated, not bulbous, aculeus relatively long and moderately curved, base narrow (Figs 33 I, J, 37 I).	en	Xuan, Qiang, Prendini, Lorenzo, Engel, Michael S., Cai, Chenyang, Huang, Diying (2025): Extinct scorpion family Chaerilobuthidae from Mid-Cretaceous Burmese amber reinterpreted as subfamily of extant family Pseudochactidae (Chelicerata: Scorpiones). Zoological Journal of the Linnean Society 203 (1): 1-60, DOI: 10.1093/zoolinnean/zlae169, URL: https://doi.org/10.1093/zoolinnean/zlae169
700187A3FFF3FFEDFF2EF99EB1FFFA27.taxon	description	Description: Based on the material examined (NIGP 200656, 201163). Carapace: Posterior width less than length (L / PW = 1.19 or 1.24, Appendix 2). Anteromedian depression narrow, shallow; anterosubmedial depressions absent; lateral ocular carinae distinct; posteromedian and posteromarginal sulci distinct; surfaces almost smooth to sparsely and finely granular (Figs 32 A, 33 A, 34 A, 37 A). Median ocular tubercle situated anteromedially; median ocelli absent; one pair of large posterolateral major lateral ocelli present. Coxosternal region: Surface smooth, sparsely covered in macrosetae (Figs 33 B, 34 D, 37 F). Leg I maxillary lobes (coxapophyses) distal margins not rounded, slightly dilated anteriorly and adorned with serrula comprising 13 or 14 denticles (Fig. 39 A). Sternum pentagonal; lateral margins sublinear, parallel; ventral surface flat, without distinct concave region or median sulcus; posteromedian depression wide, shallow (Figs 32 B, 33 B, 34 D, 37 F). Chelicerae: Fixed finger and manus, prolateral and ventral surfaces each with numerous long, dense macrosetae (Figs 34 B, 37 B). Cheliceral dentition partly visible, moveable finger with long dorsal distal (dd) denticle, long ventral distal (vd) denticle, and three or four small ventral accessory (va) denticles; fixed finger with long distal denticle. Moveable finger with prominent serrula, comprising short spinules in proximal two-thirds. Pedipalps: Segments gracile; surfaces sparsely covered by short microsetae and few macrosetae (Figs 35 A, E, 38 B, E). Femur with four carinae evident; prodorsal and proventral carinae obsolete, crenulate; retrodorsal and retroventral carinae smooth, costate; ventral surface sparsely and finely granular (Figs 35 C, G, I, J, 38 A, F, I). Patella with five carinae evident; prodorsal carina obsolete, comprising several tubercles; prolateral surface, dorsoventral ‘ vaulted’ projection (‘ anterior process’) moderately developed with prominent pair of apophyses or tubercles (‘ patellar spurs’), each possessing one macroseta; retrodorsal and retroventral carinae smooth, costate (Figs 35 A, D, E, H, 38 C, G, J). Chela relatively slender (CL / CW = 4.94 – 5, Appendix 2); manus globose; chela with six carinae evident; digital carina complete, distinct, granular medially, becoming costate proximally and distally; dorsomedian carina reduced to few granules at base of fixed finger, becoming obsolete proximally on manus; prodorsal, promedian, and proventral carinae obsolete, each comprising few isolated granules; retroventral carina distinct, disjunct proximally; dorsal secondary, subdigital, retromedian, and ventromedian carinae absent or obsolete (Figs 35 A, B, E, F, 38 D, H, K). Chela finger dentition partly visible, fixed and moveable fingers median denticle rows each comprising oblique, slightly imbricate subrows of c. 12 denticles (first subrow with five denticles), separated by large, serrate prolateral and retrolateral denticles (Fig. 33 C). Trichobothrial pattern Type D, β configuration (Figs 35, 38). Femur with 10 trichobothria, four d, three i, and three e trichobothria; trichobothria d 1, d 2, d 4, and d 6 noticeably smaller than others (‘ petite’); trichobothrium e 1 situated proximal to trichobothrium d 3; trichobothrium e 3 situated dorsal to trichobothrium d 6 (Fig. 35 A, C, E, G, I, J). Patella with 10 trichobothria, including three d, one i, and six e trichobothria; trichobothrium i 1 situated distal to trichobothrium d 3 (Fig. 35 A, D, E, H). Chela with 12 trichobothria; fixed finger with three i, two e, and two d trichobothria; manus with four E and one V trichobothria; trichobothrium Et 1 situated distally on manus, almost aligned with or proximal to moveable finger condyle; trichobothrium eb situated on fixed finger, slightly distal to moveable finger condyle; trichobothrium esb situated medially on fixed finger; trichobothrium V 2 situated distally on manus near retroventral carina; macroseta situated medially on manus and near retroventral carina (Fig. 35 A, B, E, F). Legs: Tibia lateral and ventral surfaces each with scattered macrosetae (Figs 33 D, G, H, K, 37 D). Basitarsi each with pair of spinule rows, comprising 10 – 13 relatively short spinules, distally (Fig. 34 I – L). Telotarsi each with slightly irregular pair of ventrosubmedian spinule rows, comprising relatively long spinules (Fig. 33 K, L). Ungues moderately long, distinctly curved; dactyl pronounced, pointed (Figs 34 I – L, 37 D). Pectines: Basal piece long, narrow with shallow median sulcus (Figs 33 B, 34 D, 37 H). Three marginal lamellae and four median lamellae; surface sparsely covered in macrosetae; longitudinal suture between marginal and median lamellae distinct (Figs 33 B, 34 G, 37 H). Fulcra absent or obsolete. Teeth rounded, distal three-quarters of each tooth occupied by sensillar field; peg sensilla relatively long, columnar (Fig. 34 G, 37 H). Genital operculum: Two opercular sclerites narrow and completely separated, as long as pectinal plate (Figs 33 B, 34 D, 37 F). Mesosoma: Pretergites surfaces almost smooth (Figs 33 A, E, 37 E); post-tergites I – VI surfaces finely granular, acarinate (Fig. 34 E, 37 E); tergite VII surface sparsely granular with four serrate carinae (paired dorsosubmedian and dorsolateral carinae) in posterior half of segment (Fig. 34 H). Sternites surfaces smooth, with scattered macrosetae; posterior margins broadly recurved (Figs 33 F, 34 F); respiratory spiracles (stigmata) small, ovoid to round (Fig. 34 F). Metasoma: Segments I – V progressively increasing in length, I wider than long, others longer than wide (Figs 33 I, J, 34 H, 37 G). Segments I – V lateral and ventral surfaces each with few, short macrosetae; I – III dorsal surfaces each with distinct depression. Segment I with six distinct carinae (paired dorsosubmedian, dorsolateral, and ventrolateral carinae), segments II – V each with eight distinct carinae (paired dorsosubmedian, dorsolateral, ventrolateral, and ventrosubmedian carinae); dorsosubmedian carinae distinct on all segments, serrate throughout length, converging posteriorly; dorsolateral carinae distinct, serrate on segments I – III, smooth, costate on IV and V; ventrolateral carinae obsolete on segments I and II, distinct, slightly granular on III – V; ventrosubmedian carinae absent on segment I, obsolete on II and III, distinct, serrate on IV and V. Telson: Vesicle elongate; dorsal surface relatively smooth; lateral and ventral surfaces finely and densely granular; (Figs 33 I, J, 37 I); ventral surface with several macrosetae; subaculear tubercle absent; prominent subaculear setal pair. Aculeus moderately long, shallowly curved; base moderately wide, less than half vesicle width.	en	Xuan, Qiang, Prendini, Lorenzo, Engel, Michael S., Cai, Chenyang, Huang, Diying (2025): Extinct scorpion family Chaerilobuthidae from Mid-Cretaceous Burmese amber reinterpreted as subfamily of extant family Pseudochactidae (Chelicerata: Scorpiones). Zoological Journal of the Linnean Society 203 (1): 1-60, DOI: 10.1093/zoolinnean/zlae169, URL: https://doi.org/10.1093/zoolinnean/zlae169
700187A3FFF3FFEDFF2EF99EB1FFFA27.taxon	materials_examined	Material examined: MYANMAR: Kachin State: Noije Bum near Hukawng Valley, precise locality unknown, Cretaceous (Cenomanian) burmite: two juv. (sex unknown) (NIGP 200656, 201163).	en	Xuan, Qiang, Prendini, Lorenzo, Engel, Michael S., Cai, Chenyang, Huang, Diying (2025): Extinct scorpion family Chaerilobuthidae from Mid-Cretaceous Burmese amber reinterpreted as subfamily of extant family Pseudochactidae (Chelicerata: Scorpiones). Zoological Journal of the Linnean Society 203 (1): 1-60, DOI: 10.1093/zoolinnean/zlae169, URL: https://doi.org/10.1093/zoolinnean/zlae169
