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Zookeys 569: 37-51 (2016)

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A new genus of Neelidae (Collembola) from Mexican caves

Vladimir Papac', José G. Palacios-Vargas?

I State Nature Conservancy of the Slovak Republic, Slovak Caves Administration, Cave Care Department, Zeleznitnd 31, 979 01 Rimavskd Sobota, Slovakia 2 Laboratorio de Ecologia y Sistemdtica de Microartrépo- dos, Departamento de Biologia, Facultad de Ciencias, Universidad Nacional Auténoma de México, 045 10 Mexico, DE

Corresponding author: José Palacios Vargas (troglolaphysa@hotmail.com)

Academic editor: L. Deharveng | Received 10 March 2015 | Accepted 8 January 2016 | Published 24 February 2016 Attp://z00bank.ore/DF7C35 95-33DC-49E0-88F 1-64880C804A D8

Citation: Papd¢ V, Palacios-Vargas JG (2016) A new genus of Neelidae (Collembola) from Mexican caves. ZooKeys 569: 37-51. doi: 10.3897/zookeys.569.5984

Abstract

The new genus Spinaethorax, whose proposal is based on specimens of Megalothorax spinotricosus Palacios- Vargas & Sanchez, 1999, is given a new name combination and a redescription. The type species comes from two caves in Campeche State, México. A new combination is also suggested for Megalothorax tonoius Palacios-Vargas & Sanchez, 1999. The new genus is similar to Megalothorax Willem, 1900 and Neelus Folsom, 1896, but it clearly differs from all genera within family Neelidae by a peculiar combination of characters and the presence of some new features, e.g. globular sensillum on Ant. IH, sword-like macro- setae on oral fold. A comparative table and an identification key for all Neelidae genera as well as some

summary tables of antennae chaetotaxy and legs setation for type species are provided.

Keywords

Neelidae, Taxonomy, Mexico

Introduction

The family Neelidae comprises five genera and 41 species in the world. However, only 9 species in 3 genera are known from Mexico (Palacios-Vargas 1997; Palacios-Vargas and Sanchez 1999). This family is mainly a euedaphic group of Collembola, whose members

Copyright V. Papdc, J.G. Palacios-Vargas. This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.

38 Vladimtr Papdé & José G. Palacios-Vargas / ZooKeys 569: 37-51 (2016)

are usually very small (0.3-1.0 mm), they have nor eyes or scales, however, their anten- nae are shorter than their heads. The main differences among the genera of this family are in the structure of forehead setation, antennae, sensory fields and furcula.

Members of this family are cosmopolitan. Therefore, they have been found in vari- ous localities mainly associated with soil and litter at different altitudes from sea shore up to 3,000 m a.s.l. (Garcia-Gémez et al. 2009). They have frequently been recorded from caves, mainly in places with rich organic material. This family has two endemic genera: Acanthoneelidus, with only one species from Europe, and Zelandothorax from New Zeland plus three cosmopolitan genera. Megalothorax is the most diversified with 28 species, Neelus with 6 species Neelides with 5, both widely spread, too. The new genus described herein has two species distributed in Mexican caves.

The most remarkable contribution on this family from Mexico is the work by Bonet (1947), who revised the whole family of Neelidae. The catalog by Palacios-Vargas (1997) records 7 species in this family but some of them have to be revised in the light of new characters. The most recent contribution to the taxonomy of this family from Mexico was that by Palacios-Vargas and Sanchez (1999) who described the two new species that are revised herein.

Materials and methods

The present redescription of Megalothorax spinotricosus Palacios-Vargas & Sanchez, 1999, is based on original slides deposited in Facultad de Ciencias, UNAM. Specimens were obtained from samples of bat guano and soil that were processed by Berlese-Tul- lgren funnels and preserved in 75% ethylalcohol. Slides were mounted using Hoyer’s solution.

Body length was measured on slides excluding antennae and furcula. Lengths of unguis and unguiculus were measured between the most basal (proximal) point and the tip on their inner margins. Besides the common measurements, the ratio “unguis I, II, III (inner margin): Ti. I, II, II width (middle part)” that can be used as additional character, was also included.

We followed nomenclature used in last Neelus revision (Kovaé and Papaé 2010) for labral setae and dental spines. Nomenclature used in the most recent revision of Megalothorax after Schneider and D’Haese (2013) was applied for arrangement of sensory fields, wax rod crypt (wrcl—8) on head, ‘Th. and Abd. sensilla s1, s2, s3, subseg- ments of dens and chaetotaxy of antennae. Forehead chaetotaxy (presence of a0 seta) is applied according to Deharveng (1978) and posterior chaetotaxy of head is used according to Palacios-Vargas and Sanchez (1999).

Abbreviations: Ant.—antennal segment; Th.—thoracic segment; Abd.—abdomi- nal segment; Ti.—tibiotarsus; scx—subcoxae; s.f.—sensory field; wrc—free wax rod

generating crypt; sl, s2, s3, s3’—swollen sensilla; dp—proximal part of dens; dd—dis- tal part of dens; UNAM—Universidad Nacional Auténoma de México.

A new genus of Neelidae (Collembola) from Mexican caves by,

Taxonomy

Spinaethorax gen. n. http://zoobank.org/9E27A3C3-9464-4A6A-94B8-5BB470502D2C

Diagnosis. A genus of the Neelidae Folsom, 1896 with the following diagnostic characters:

Habitus of Neelidae. Small size, about 0.6 mm. Color white. Tegumentary gra- in fine and uniform. Apex of head with sword-like spines, body with several such spines, mainly around sensorial fields. Ant. II and IV fused and Ant. III with small globular sensillum in proximal position. Anterior labral setae R, and R, thick, curved and smooth. Oral fold with 1+1 sword-like macrosetae. Basomedian field of labium furnished with 6+6 setae. Presence of 3 setae around abdominal sensory fields, no E3 spine/setae on dd. Mid abdomen with swollen sensilla s3 and s3’. Base of Abd. IV sternite with 1+1 neosminthuroid setae, smooth and with pointed tip.

Type species. Spinaethorax spinotricosus (Palacios-Vargas & Sanchez, 1999), comb. n.

Redescription. Figs 1-15.

Type material. Holotype: female mounted on slide. Original label: 23/00/1991, Mexico, Campeche, Cueva Xtancumbilxunaan (cave), 29.xii.1996, A. Ruiz and S. Aguilar col., ext. soil. Paratypes: 1 female on slide, 23.viii. 1991, J. G. Palacios col., direct collection and 2 juveniles on slides, the same data as the holotype. Type material deposited at Facultad de Ciencias, UNAM.

Other material. Mexico, Campeche, Cueva Actin Guachapil (cave), 1 male, 1 female and 2 juveniles on slides. Original label: 14/iv/2012, 22.iii.1997, A. Ruiz and S. Aguilar col., ext. guano; 1 female on slide, 29.v. 1997, J. G. Palacios col., ext. guano. Material deposited at Facultad de Ciencias, UNAM.

Diagnosis. Unpaired seta a0 between antennal basis. Labral chaetae R, and R, thick, curved and smooth. Oral fold with 1+1 sword-like macrosetae. Basomedian field of labium with 6+6 setae. Ant. HI with small globular sensillum. Manubrium with 4+4 posterior setae. Mucro with both lamellae serrated and with rounded tip.

Description. For lengths of different character see Table 1. Body length 0.5—0.7 mm. Habitus globular, as other members of Neelidae. No pigmentation, cuticle finely granulated, /inea ventralis without crossing with integumentary channels on ventral head back. Mid and hind Abd. with numerous spine-like microsetae, stouter spines around all sensory fields on body and apex of head.

Head. Head length and width 215 and 145 um, respectively. No eyes. Head with smooth, pointed ordinary setae and spines of different width and length (Fig. 1). Frontal part with ordinary setae (lateral ones longer than axial, 26-30 um, re- spectively 10-16 um), seta a0 present; medial part between posterior s.f. ordinary or slightly spine-like setae (20 sm); posterior part with 3+3 stouter spines IL, —IL, of different lengths (28-42 pm) and 1+1 smaller axial spines IL, (17—21 um), others similar to smaller setae (10-14 um). Labrum with 5,5,4 setae, 4 prelabrals. Pattern of labral setae (Fig. 2) after Massoud and Vannier (1967): a-row: 2R, + 2R,, m-row:

40 Vladimtr Papdé & José G. Palacios-Vargas / ZooKeys 569: 37-51 (2016)

Table |. Lengths (um, mean in parenthesis) of different morphological characters of Spinaethorax spi-

notricosus comb. n. (male, females and juvenile separately).

Body:part Spinaethorax spinotricosus| Spinaethorax spinotricosus comb. n. male comb. n., females comb. n., juvenile Body total 520 530-720 (605) 350-500 (445) Head width 135 133-215 (162) 100-130 (121.2) Head length 170 183-237 (209) 130-167 (154.8) Riabum |B 9-12 (10) 7-8 (7.6) R2 labrum 10 10-14 (11.5) 8-10 (9.3) Antenna 120 125-145 (132.6) 94-112 (106.3) Ant. I 10 11-13 (11.4) 9 Ant. II 23 25-30 (26.4) 18-23 (21.6) Ant. I-IV 87 90-104 (97.5) 67-81 (76) Ant. IV sensillum Sy 16 16-18 (16.6) 10-12 (11.3) Ant IV macrosensilla S 20-32 (23-31) 1424 (14.8—22.6) Tibiotarsus I width 17 17-18 (17.5) 15-17 (16.4) Tibiotarsus IT width le? 16-19 (16.8) 14-17 (16) Tibiotarsus III width 17 16-19 (17.4) 16-18 (16.5) Unguis I Ze 26-33 (28.7) 18-23 (21.2) Unguis Il 24-30 (26.3) 16-21 (19) Unguis III 23 23-30 (25) 15-21 (18.8) Unguiculus I 14 13-16 (14.1) 10-12 (11.2) Unguiculus II 14 14-18 (15.5) 10-12 (11) Unguiculus II 15 14-18 (16.1) G=1a C15) Manubrium 54 56-66 (60.3) 35-45 (42.5) Dens (proximal part, dp) 52 29-36 (32.3) 20-26: (23:3) Dens (distal part, dd) 66-83 (71.2) 41-57 (55.6) Mucro 67 65-87 (73.6) 42-58 (53.4) Ries 6 7-9 (7.7) 5~7 (6.1) Macroseta on oral fold 25 25-30 (28) 18-23 (21) Spines IL, on head 35-42 (37.3) 28-30 (28.8) Spines IL, on head 33 28-36 (31.1) 21-26 (23.9) Spines IL, on head 30 28-33 (29.8) 17-25 (22.5) Spines IL, on head 17-21 (18.1) 12-15 (14)

m + 2r, + 2r, and p-row with 5 ordinary setae (11 pm). Anterior R, and R, slightly thick, smooth and curved, R, (11 pm) longer than R, (9 pm). Medial setae (m-row) equal (11 um), smooth median setae in one line with others. Maxillary palp simple, with 1 enlarged terminal seta (18 um), 1 basal seta (14 um) and 1 sublobal hair (Fig. 2). Basomedian field of labium with 6+6 setae (Fig. 5), median ones slightly longer

A new genus of Neelidae (Collembola) from Mexican caves 4]

3

ae a i | & Lay pe ui

.

(|) AWN

ANNE Re, 8

Figures 1-5. Spinaethorax spinotricosus. | dorsal chaetotaxy of head 2 anterior part of head with labrum, of—oral fold 3 mandible 4 maxilla 5 labium with ventral head back, bA— basolateral field of labium.

42 Vladimtr Papdé & José G. Palacios-Vargas / ZooKeys 569: 37-51 (2016)

(12 um) than others (10 um); basolateral field with 1+1 setae (10 um), oral fold with 1+1 basal setae (8 pm) and 1+1 terminal sword-like macrosetae (25-30 um). Head with 3+3 smooth postmedian setae ventrally (Fig. 5); 2+2 anterior setae equal (16 um); posterior 1+1 seta slightly curved at tip (18 pm). Mandible with 4 apical teeth, medial ones longer (Fig. 3). Maxilla as in Fig. 4.

Thorax and abdomen (Fig. 6). Dorsally with ordinary setae, swollen sensilla s1, s2, s3, s3’, spines of different size (4-6 and 25-45 um), 6+6 wax rods (wrcl, 2, 4, 6-8) with straight setae (3-4 um) and 2+2 (wrc 3 and 5) with thicker and longer straight setae (6-7 um). All wrc placed in small cuticular depressions. Trichobothria or their sockets not observed. ‘Th. II with 3+3 ordinary setae (12-14 um) and 6+6 stouter spines (25-36 pm) around thoracic sensory fields, axial spines smaller (25 um); sensory fields at leg II base with 2+2 ordinary setae (20 um), 3+3 stouter spines, anterior ones longer (45 um) than posterior (26—28 um), 1+1 lateral sensillum s1 (8 um) broadened at tip and 1+1 swollen transparent rod with bifid tip and no base (8 um) above s.f. of leg II. Th. III with 5+5 ordinary setae (6-8 um), 4+4 wrc (wrc 1-4) and several small spine-like microsetae, whose overall number is not seen clearly; at leg II base with 1+1 ordinary seta (16 um), 3+3 stouter spines, medial ones longer (42 pm) than lateral (22 um) and 2+2 wrc (5, 6). Anterior Abd. medially with 1+1 wre 7 and 2+2 swollen sensilla s3 and s3’; hind Abd. with abdominal sensory fields, which are surrounded by 2+2 ordinary setae (12 and 18 ym), 1+1 stouter spines (30 pm), 1+1 swollen sensilla s2 and 1+1 wrc 8 above Abd. s.f. Dorso and dorso-lateral anterior and hind abdomen covered with numerous spine-like microsetae (4—6 pm) arranged as in Figs 6 and 10. Their overall number is not seen clearly. Abd. tergum VI with 3+3 setae (17 um) and 1 unpaired axial seta (14 um). Anal complex with three anal valves, each with on seta (7 um). Abd. VI sternum with 4+4 setae (18-20 um) and 2+2 very small globular structures (1—2 pm) next to anal valve setae. Female genital plate (Abd. V sternum, Fig. 10) with 4+4 setae (8-11 um) and 1+1 axial microsetae (4 um). Male genital plate with 5+5 ordinary setae (12 um) arranged in circle and with 2+2 spine-like setae (7 um) difficult to observe (Fig. 9), laterally surrounded with 4+4 setae (18 um). Abd. IV sternum with 3+3 setae in one row (18—20 um), one seta more laterally and 1+1 distal setae (8 um). Lateral part of Abd. IV sternum with 1+1 short and pointed neosmin- thuroid setae (6 um) (Fig 10).

Appendages. Ant. III and IV not separated (Fig. 11). Length of antennae 140 um, ratio antenna/head = 0.65; length of antennal segments I, IH, III-IV as 11, 26 and 103 um. Ant. I furnished with 3 short setae (8-10 um). Ant. IT with 1 medial seta and 5 apical setae arranged in a whorl. Ant. III organ consists of 2 transparent rods (7 um), 1 leaf-like transparent sensillum Sg (12 um) and spine-like seta (7 um). Proximal part of Ant. III bears 1 globular sensillum Sg (4 um). Ant. IV with 13 curved macrosensilla S finely blunt at tip (24-32 um); subapically with 1 long and thick subapical sensillum Sy (16-18 pm) and with 1 thick shorter apical sensillum Sx broadened at tip (12-14 um); Ant. IV organ like a tiny, hardly visible spine (5 ym); apically with curved setae a and sa apical in position (10 um). Complete chaetotaxy of antennae provided in

Table 2. Setae numbers of legs I-III (Figs 13-15): scx I: 1, 1, 3 (one spine); scx I:

A new genus of Neelidae (Collembola) from Mexican caves 43

:

oe SAS

Figures 6-10. Spinaethorax spinotricosus. 6 thoracic and abdominal chaetotaxy with abdominal sensory field enlarged 7 ventral tube in lateral view 8 tenaculum 9 male genital plate in lateral view, arrow shows

anterior direction 10 female genital plate frontal view, av—anal valve setae, nch—neosminthuroid setae.

44 Vladimtr Papdé & José G. Palacios-Vargas / ZooKeys 569: 37-51 (2016)

Figures | 1-12. Spinaethorax spinotricosus. \1 dorsal Ant. I-IV, with Ant. IV organ enlarged 12 furcula,

posterior view with anterior seta enlarged.

1, 1, 1; coxae: 1, 1, 2; trochantera: 4, 3, 3; femora: 8, 8, 8 and tibiotarsi: 13, 14, 13. Thin meso- or microsetae as in following numbers on leg I: femur with 2; on leg I: femur with 1; on leg III: trochanter with 1, femur with 1. For complete setation of legs see Table 3. Tibiotarsal tenent hairs ordinary, straight and pointed (16-18 um). Unguis narrow, in distal part bended; both unguis and unguiculus unequally long in leg I, Il and III: unguis 31, 27 and 26 um, respectively, unguiculus 14, 15 and 16 um,

A new genus of Neelidae (Collembola) from Mexican caves 45

Figures 13-15. Spinaethorax spinotricosus. 13 leg I 14 leg II 15 leg UI, with detail of each feet complex.

46 Vladimtr Papdé & José G. Palacios-Vargas / ZooKeys 569: 37-51 (2016)

Table 2. Chaetotaxy of antennae in Spinaethorax spinotricosus comb. n.

Spinaethorax spinotricosus comb. n. Chaetae Sg Misc Ant. I 3 0 Ant. II 6 0 2 Sensory organs, Ant. III 15 0 1 spine-like chaeta, 1 globular sensillum Sg, 1 leaf-like sensillum Sg Ant. IV 13 0 Iysxe: Sys) Orsdiattsa

Table 3. Setation of legs in Spinaethorax spinotricosus comb. n.

Spinaethorax spinotricosus comb. n.

Subcoxae I 1 1 3 Subcoxae II 1 1 1 Coxae 1 1 2 Trochantet 3 Tibiorarsus 13

respectively. Length ratio unguis I, II, III / ti. I, U, UI width (31, 27, 26/ 17, 16, 17 um) = 1.8, 1.7, 1.5. Unguis I and III with three auxiliary lamellae la, lp, Bp, unguis II with only lp and Bp (Fig. 14); unguiculus I with small internal tooth on distal part, II and I untoothed and III with bended tip; unguiculi without apical filament and basal lamellae. Ventral tube with 2+2 distal setae and without posterior lobe (Fig. 7). Retinaculum with 3+3 teeth, no setae on corpus (Fig. 8). Furcula well developed (Fig. 12), length of manubrium, dens (dp and dd) and mucro: 56, 31, 71 and 74 um, respectively. Manubrium with 4+4 setae posteriorly, lateral ones shorter (10-15 pm) than axial (16-20 pm). Dens in proximal part (dp) with 2+2 posterior setae, lateral ones (14 pm) shorter than those axial (22 um); distal part (dd) apically with 2+2 broad, blunt lateral spines (9 um) and 1 medial sharp spine (7 um) on anterior side; with 2 external (E1—E2) posteriorly and 2 internal (J1—J2) spines (6—7 um each, distal with short apical filament), and 1 medial, subapical seta (18 um). Mucro with serrated la- mellae and rounded tip. Base of mucro furnished with small scale without base. Mid- dle mucro width 7 um.

Sensory fields (Figs 1, 6, 14 and 15). 6+6 s.f. placed in depressions each with secre- tory rod (10-12 pm), i.e. blunt seta with basal part inserted on cuticle and placed in upper margin of the field. Following arrangement: (a) anterior and posterior field on head (s.f. 1, 15 x 10 and s.f. 2, 20 x 10 um) each with secretory rod and 1 seta on margin (14-18 pm); (b) thoracic field (s.f. 3; 35 x 20 um) with secretory rod, 3 internal spines

A new genus of Neelidae (Collembola) from Mexican caves 47

(4 um) arranged in triangle, 2 external marginal setae (12-14 um) and 6 sword-like spines of different lengths (25-36 um), 3 spines are in anterior position and 3 spines above s.f. in axial position; (c) fields at base of legs II and III (s.f. 4, 5; 20 x 15 um) each with secretory rod, 2 internal spines (4 ym) in s.f. 4 and 1 internal spine (4 um) in s.f. 5. S.f4 with 2 marginal external setae (20 um), 3 stouter sword-like spines, medial ones longer (45 pm) than lateral (26-28 um), 1 lateral sensillum s1 broadened at tip (8 um) and 1 swollen rod with split tip and without base (8 um) above s.f. 4. S.f. 5 with 1 marginal external seta (16 pm), 3 stouter sword-like spines, medial ones longer (42 um) than lateral (22 um); (d) abdominal field (s.f. 6; 30 x 25 um) with secretory rod, 1 internal spine (4 um), 2 marginal ordinary setae (12 and 18 pm), | sword-like spine above s.f. (30 um) and 1 swollen sensillum s2 on the margin of s.f. (6 pm). Wrc 8 is located above Abd. s.f.

Both sexes known.

Etymology. The genus is named after the spine-like setae on thorax and abdomen.

Distribution and ecology. Spinaethorax spinotricosus is currently known from two caves and is putatively spread in the cave systems of Yucatan Peninsula, mainly in places with accumulation of bat guano or other type of rich organic material.

Variation. The young have only one proximal seta on dens.

Spinaethorax tonoius (Palacios-Vargas & Sanchez, 1999), comb. n.

Note. This species, described from a cave in State of Guerrero, shares with new type species of the genus similar generic characters like presence of sword-like macrosetae on oral fold, globular sensillum on Ant. HI., stouter spines on ‘Th., fusion of Ant. HI and IV, 3+3 setae around Abd. s.f., 2+2 setae on proximal part of dens and absence of E3 spine on distal part of dens. The main differences consist in setation of hind Abd. (numerous thickened macrosetae in S. tonoius comb. n. vs. numerous spine-like micro- setae in S. spinotricosus comb. n.), in chaetotaxy of the apex of head (only one spine IL, in S. tonoius comb. n. vs. three spines IL,—IL, in S. spinotricosus comb. n.), in the structure of tenent hairs on Ti. (more developed in S. tonoius comb. n. vs. shorter in S. spinotricosus comb. n.) and in setation around Abd s.f. (absence of axial spine in S. tonoius comb. n. vs. presence of spine in S. spinotricosus comb. n.). Species description is based only on one adult specimen (holotype) and does not allow us to describe it in an appropriate way, along with drawings and measurements.

Discussion. The family Neelidae was for a long time an overlooked group of Col- lembola, mainly due to its small size and lack of diagnostic characters. In spite of the foregoing facts recent years molecular phylogenetic analysis have revealed unexpected diversity within this family (Schneider et al. 2011). Recently Neelus has been revised (Kovac and Papaé 2010) as well as Megalothorax with a redescription of the nominal species Megalothorax minimus (Schneider & D’Haese, 2013). Soon after additional new taxonomical characters in genus Megalothorax were defined (Papa¢ and Kovac 2013). S. spinotrichosus comb. n. and S. tonoius comb. n. share many specific characters.

Viadimtr Papdé & José G. Palacios-Vargas / ZooKeys 569: 37-51 (2016)

48

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A new genus of Neelidae (Collembola) from Mexican caves 49

These species were included in the genus Megalothorax (Palacios-Vargas & Sanchez, 1999), because some features (fusion of Ant. segments II and IV, absence of E3 spine on distal dens) indicated that those specimens belonged to this genus. On the other hand, some characters link those specimens to Neelus (dp with 2+2 setae - except WN. fimbriatus, 1+1 neosminthuroid setae at the base of Abd. IV sternite). Schneider and D’Haese (2013) stated that chaetotaxy of M. spinotricosus clearly differs from the other Megalothorax by the presence of great number of microsetae on Abd. and pointed out that MV. spinotricosus deserved its own genus. On the basis of recent diagnostic features, material of these two species was re-examined arriving to the same conclusion as Schne- ider and D’Haese (2013) that they represent a new genus in the Neelidae. Spinaethorax gen. n. differs from other genera by striking morphological features and combinations, which clearly separate them, e.g. sword-like macrosetae on oral fold, six setae on baso- median field of labium, Ant. III and IV fused, presence of small globular sensillum Sg on Ant. III, three marginal setae around abdominal sensory field, absence of spine E3 on dd and dp with two seate. For comparison with other genera see Table 4.

Identifiaction key to the World genera of Neelidae

The identification key is based on that of Bretfeld (1999).

1 Head and body with well developed sensory fields; R, labrum setae shorter than R,; labrum without apical fringes and split structures; retinaculum with Oreo gO le ee POOLS AR rh tbe eR Racing Senta Sl Rs eRe eT Ale it ba ee mag SAS 2

- Large sensory fields absent; R, labrum setae longer than R,; labrum with apical fringes and split structures; retinaculum with 2+2 teeth... facts Neelides Caroli, 1912, type species Neelides folsomi Caroli, 1912; Italy

2, Dens with short conical spines posteriorly; median labral setae present; a-row Oblabauintwithy4-Or G:setae cals ted ewesuse wees pownebs PanonaU monte teres akensns tense eae 3

Dens with broad triangular spines posteriorly; median labral setae missing; ASEOW, Olva DIU Rae WAIN De SOlAS AaPen sn) Bice Seond Barre scel Balen ee Ral eee, aM Buen: Ue ieee abe Aes Zelandothorax Delamare Deboutteville & Massoud, 1963, type species Megalothorax novozealandiae Salmon, 1944 (New Zealand)

3 partnvoer! UU Re-baY BAYS ale cia ie: k 9 heen aye ec ee we ieee ey kee Pa ver ewer N 4 - nt, lean: Separated wit sutras. wb ee ce we, oe eee ET > 4 Sensory fields of abdomen with 5 marginal setae; 2+2 neosminthuroid setae; basomedian field of labium with 3+3 or 4+4 setae; proximal part of dens with ONO SETA a tosnueatan ec tasdialet shine Rdilas SA Mh SiS Ae tesco Megalothorax

Willem, 1900, type species Megalothorax minimus Willem, 1900; Belgium = Sensory fields of abdomen with 3 marginal setae; 1+1 neosminthuroid setae; basomedian field of labium with 6+6 setae; proximal part of dens with two

naethorax gen. n., type species Spinaethorax spinotricosus comb. n. (Mexico)

50 Vladimtr Papdé & José G. Palacios-Vargas / ZooKeys 569: 37-51 (2016)

5 Sensory fields of abdomen with 2 marginal setae; apex of head without spines; proximal part of dens with two setae (only N. fimriatus with one SEL) Mea rccsteteet ce Miavax, eossauoa oe ema nies Ree cecncen a acca eet. Neelus Folsom, 1896, type species Neelus murinus Folsom, 1896; United States of America - Sensory fields of abdomen with 5 marginal setae; apex of head with blunt spines} proximal jpart of dens wathyonheseta.t cote. ..s.0e. coat. -acphacnneeeeeeedpan eres 6 os 9 Ls dae cols Reels ty hg Acanthoneelidus Bretfeld & Griegel, 2006, type species Acanthothorax pratensis Bretfeld & Griegel, 1999 (Poland)

Acknowledgements

We thank to International Exchange Office UNAM (Direccién General de Cooper- acion e Internacionalizacién) for supporting this project, we also thank the support of CONABIO (Project LE002) for digitalization of the collections data of the Facultad de Ciencias, UNAM made by Isaac Cruz, Maira Montejo and Leonel Pérez. Luis Parra

and Hugo Mejia reviewed the mansucript and gave criticism.

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