Acari: Mesostigmata: Rhodacaridae

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Zootaxa 4363 (3): 409–420 http://www.mapress.com/j/zt/ Copyright © 2017 Magnolia Press

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ISSN 1175-5326 (print edition)

ZOOTAXA

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https://doi.org/10.11646/zootaxa.4363.3.6 http://zoobank.org/urn:lsid:zoobank.org:pub:0E67BB90-5833-4C44-B352-2AF232F67847

Two new species of Afrodacarellus (Acari: Mesostigmata: Rhodacaridae) from Brazil

and a key to the world species of the genus MARCIA D. DOS SANTOS1, RAPHAEL C. CASTILHO2, GILBERTO J. DE MORAES3 & EDMILSON S. SILVA4 1

Centro de Ciências Agrárias, Universidade Federal de Alagoas, 57100-000 Rio Largo, Alagoas, Brazil. E-mail: [email protected] 2 Departamento de Fitossanidade, UNESP/FCAV, 14884-900 Jaboticabal, São Paulo, Brazil. E-mail: [email protected] 3 CNPq Researcher, Departamento de Entomologia e Acarologia, ESALQ–Universidade de São Paulo, 13418–900 Piracicaba, São Paulo, Brazil. E-mail: [email protected] 4 Laboratório de Entomologia e Acarologia, Universidade Federal de Alagoas, 57309-005 Arapiraca, Alagoas, Brazil. E-mail: [email protected]

Abstract Within a project to determine the edaphic predatory mites from the Brazilian Alagoas state, two new species of Afrodacarellus Hurlbutt (Rhodacaridae) were found, namely Afrodacarellus alagoensis Santos & Castilho n. sp. and Afrodacarellus xucurukariri Santos & Castilho n. sp.. These are here described based on the morphology of adult females. A key for the separation of females of the 31 recognisable world species of Afrodacarellus is provided, including the newly described species. Key words: edaphic mite, Rhodacaroidea, taxonomy

Introduction Rhodacaridae Oudemans is a widespread mite group living in the soil and in accumulation of decaying organic matter. Species of this group have been generally reported in the literature as predators (Walter et al., 1988; Castilho et al., 2009; Castilho et al., 2015). This family is relatively small, with about 155 known species, of which slightly over 30 belong to Afrodacarellus Hurlbutt, the most diverse genus (Castilho et al., 2012; Abo-Shnaf et al., 2013; Rueda-Ramirez et al., 2013; Castilho et al., 2015; Duarte et al., 2016). Afrodacarellus was established by Hurlbutt (1974) as an offshoot of Afrogamasellus Loots & Ryke, in an extensive study of species from Tanzania then considered to belong to the latter genus. The Tanzanian species were separated in two groups, one of which with epistome containing a stalked anteromedian extension, long and narrow genital shield and posterolateral rod-like seta at the base of tarsus I at least one-third as long as the segment. This group was considered to constitute a new genus, then named by him as Afrodacarellus. A full characterisation of the genus was given by Castilho et al. (2012). Most Afrodacarellus species were described from Africa (Castilho et al., 2012). Exceptions are six species described from South America: three from Argentina (Karg, 1977; 1979), two from Ecuador (Karg, 2003) and one from Venezuela (Karg & Schorlemmer, 2009). A survey was recently conducted in the northeastern Brazilian Alagoas state in an effort to determine the local edaphic predatory mites. The aims of this study are to describe two new Afrodacarellus species found in that survey and to provide a key to help the separation of the world species of this genus.

Material and methods Soil and litter samples were collected in different parts of Alagoas. They were taken to the laboratory of Accepted by B. Halliday: 4 Aug. 2017; published: 12 Dec. 2017

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Universidade Federal de Alagoas (Arapiraca campus) where mites were extracted using Berlese funnels. The extracted mesostigmatid mites were mounted in Hoyer’s medium and later identified to families. The rhodacarids collected were examined under phase (Leica, DMLB) and interference (Nikon, Eclipse 80i) contrast microscopes to compare with the original descriptions and redescriptions of mites of this family. Given that they were identified as Afrodacarellus, but did not fit the description of the species of this genus, they were described as new species. Taxonomically relevant structures were illustrated with the use of a digital camera attached to the interference contrast microscope and measured with the use of a graded ocular. Setal nomenclature is based on Lindquist & Evans (1965), as adapted to the rhodacarids by Castilho & Moraes (2010). Leg chaetotaxy is based on Evans (1963). For each structure, the mean and the corresponding range (for variable measurements) are given in micrometres.

Results Genus Afrodacarellus Hurlbutt, 1974 Afrodacarellus Hurlbutt, 1974: 589. Type species: Afrodacarellus femoratus Hurlbutt, 1974, by original designation. Afrogamasellus (Foliogamasellus) Karg, 1977: 345 (Synonym by Castilho et al. 2012). Type species: Afrogamasellus camaxiloensis Loots, 1969, by original designation. Afrogamasellus (Foliogamasellus).—Karg, 2003: 26; Karg & Schorlemmer, 2009: 65. Afrogamasellus (Latogamasellus) Karg, 1977: 345 (Synonym by Castilho et al. 2012). Type species: Afrogamasellus squamosus Karg, 1977, by original designation. Afrogamasellus (Latogamasellus).—Karg, 2003: 26; Karg & Schorlemmer, 2009: 65. Latogamasellus.—Antony, 1986: 126.

Afrodacarellus alagoensis Santos & Castilho n. sp. Diagnosis of adult female. Anteromedian extension of epistome flanked by two pairs of anterolateral extensions; opisthonotal shield with 19 pairs of smooth setae (R5 absent); seta J2 at least as long as distance between its base and base of J3; seta J4 at least 1.3 times as long as distance between its base and base of J5; without plates laterad of genital shield; ventrianal shield with six pairs of setae (including Jv1) in addition to circumanal setae; unsclerotised cuticle laterad of ventrianal shield with two pairs of setae (Jv4 and Jv5); with a pair of rounded (bearing a pore-like structure) and a pair of elongate metapodal platelets, not fused with ventrianal shield. Adult female (Figs 1–11) Five specimens measured. Gnathosoma. Fixed cheliceral digit 20 (20–22) long, with five teeth in addition to apical tooth and a discreet setiform pilus dentilis (Fig. 1); movable cheliceral digit 24 (23–25) long, with three teeth in addition to apical tooth; antiaxial and dorsal lyrifissures as well as dorsal seta distinct, the latter stubby. Arthrodial process of chelicera brush-shaped. Number of setae on palp trochanter-tarsus: 2–5–6–14–15; setae al of palp femur and al1 and al2 of palp genu distinctly stouter than other setae of same respective segments (Fig. 2) (al2 markedly more ventral and only slightly posteriad of al1); apotele 3-tined. Anteromedian extension of epistome flanked by two pairs of anterolateral extensions and few irregular denticles, the inner slightly shorter than the median extension and slightly longer than the outer, both distally variously shaped (acute, blunt or expanded, smooth or denticulate) (Fig. 3). Deutosternum delimited by lateral lines, with eight transverse lines, the basalmost smooth and others with 4–6 denticles each (Fig. 4); smooth lines projecting outward from deutosternum between the first and second transverse lines and between the second and third transverse lines. Internal malae indistinct. Corniculus horn-like, about twice as long as its basal width. Seta h3 about in longitudinal line with h1 and posteromediad of h2. Measurements of setae: h1 10 (8–10), h2 9 (8–11), h3 6 (5–7), sc 11 (10–11); all setae aciculate and smooth. Dorsal idiosoma (Fig. 5). Idiosoma 260 (255–273) long and 136 (125–158) wide at widest level. Podonotal and opisthonotal shields separate. Podonotal shield smooth, except for few striae laterad of j6 and z5, and a line parallel to posterior margin, which delimits a narrow and smooth posterior band; with a delineated marginal strip containing the insertions of j1, z1, r2, r3, r5 and r6; 130 (125–135) long and 129 (126–134) wide at widest level; with 22 pairs of setae (j1–j6, z1–z6, s1–s6, r2, r3, r5 and r6), four pairs of distinguishable lyrifissures, one pair of

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distinguishable pores and two pairs of scleronoduli between j5 and j6. Unsclerotised cuticle along lateral margins of podonotal shield with a pair of setae (r4). Opisthonotal shield reticulate, with a line parallel to anterior margin delimiting a narrow and smooth anterior band; with a delineated marginal strip containing the insertions of R1–R4; 129 (118–138) long and 111 (108–119) wide at widest level; with 19 pairs of setae (J1–J5, Z1–Z5, S1–S5 and R1– R4), eleven pairs of distinguishable lyrifissures and a pair of pores. Unsclerotised cuticle along lateral margins of opisthonotal shield with one pair of distinguishable lyrifissures. Measurements of setae: j1 9 (7–10), j2 13 (12–14), j3 16 (15–17), j4 17 (16–17), j5 16 (15–17), j6 16 (16–17), z1 6 (5–7), z2 14 (9–16), z3 16 (14–18), z4 17 (16–18), z5 20 (18–21), z6 21 (20–21), s1 6 (5–7), s2 11 (9–16), s3 17 (16–17), s4 21 (20–22), s5 23 (20–26), s6 24 (22–25), r2 23 (21–26), r3 16 (15–17), r4 7 (6–9), r5 19 (15–21), r6 21 (20–22), J1 29 (27–30), J2 29 (27–30), J3 27 (25– 30), J4 30 (26–32), J5 28 (27–30), Z1 31 (30–33), Z2 32 (30–35), Z3 32 (30–33), Z4 30 (26–35), Z5 24 (20–26), S1 31 (30–32), S2 31 (30–32), S3 31 (30–33), S4 30 (30–32), S5 27 (25–30), R1 24 (21–25), R2 24 (23–25), R3 22 (21–22), R4 24 (23–25). All dorsal idiosomal setae aciculate and smooth, r6 distinctly stouter. Most podonotal setae shorter than distance to subsequent seta of same series; opisthonotal setae at least as long as distance between their base and base of the subsequent setae of same series. Ventral idiosoma (Fig. 6). Base of tritosternum 10 long and 6 (5–7) wide proximally (Fig. 7); laciniae 37 (30– 40), separated for about 50% of their total length, pilose. Sternal shield reticulate laterally and smooth centrally; region anteriad of first pair of lyrifissures (iv1) lightly sclerotised and punctate; posterior margin concave; approximately 63 (61–65) long from center of anterior margin of punctate and lightly sclerotised region to center of posterior margin, and 57 (55–59) wide between coxae II and III; with four pairs of setae and three pairs of lyrifissures; seta st1 inserted in punctate and lightly sclerotised region of sternal shield. Discrete section of endopodal plate arched, extending from posterior region of coxa III to mid region of coxa IV. Genital shield reticulate and with a punctate band along straight posterior margin; extending posteriorly behind coxae IV; without plates laterad of genital shield; distance between st5–st5 32 (31–34). Lyrifissure iv5 on unsclerotised cuticle, posterolaterad of st5. Ventrianal shield reticulate; except for a slender smooth band along anterior margin; 103 (98– 112) long and 99 (95–102) wide at widest level; with six pairs of setae (Jv1–Jv3, Zv1–Zv3). Unsclerotised cuticle laterad of ventrianal shield with two pairs of setae (Jv4 and Jv5) and two distinguishable pairs of lyrifissures. Peritreme extending anteriorly to posterior region of coxa II (r3). Peritrematic plate narrow, fused anteriorly to dorsal shield at level of r2, extending posteriorly as a narrow strip behind coxa IV; lyrifissure and pore next to coxa III and lyrifissures and pore behind stigma not distinguishable. Exopodal plate fragmented into subtriangular platelets between coxae II–III and III–IV. With a pair of rounded and a pair of elongate metapodal platelets, the first bearing a pore-like structure. Measurements of setae: st1 14 (12–15), st2 12 (11–14), st3 11 (11–14), st4 11 (11– 12), st5 12 (11–12), Jv1 14 (13–15), Jv2 15 (14–17), Jv3 23 (21–25), Jv4 28 (26–30), Jv5 28 (27–31), Zv1 13 (12– 15), Zv2 12 (11–13), Zv3 11 (10–12), para-anal 20 (18–21), postanal 24 (23–26). All setae aciculate and smooth, except Zv2 and Zv3, forked distally. Spermathecal apparatus. Not distinguishable. Legs. (Figs 8–11). Lengths: I 228 (213–243), II 162 (143–173), III 133 (130–135), IV 174 (170–176). Chaetotaxy: I coxa 0 0/2 0/0 0, trochanter 1 0/1 1/2 1, femur 2 3/1 2/3 2, genu 2 3/2 3/1 2, tibia 2 3/2 3/2 2; II coxa 0 0/1 0/1 0, trochanter 1 0/1 0/2 1, femur 2 3/1 2/2 1, genu 2 3/1 2/1 2, tibia 2 2/1 2/1 2; III coxa 0 0/1 0/1 0, trochanter 1 1/1 0/2 0, femur 1 2/1 1/0 1, genu 2 2/1 2/1 1, tibia 2 1/1 2/1 1; IV coxa 0 0/1 0/0 0, trochanter 1 1/1 0/ 2 0, femur 1 2/1 2/0 0, genu 2 2/1 3/1 1, tibia 2 1/1 3/1 2; tarsi II–IV: 18, 18, 17. Seta al2 of femur II stubby. Several setae of tarsi II and III, ad2 and pd1 of femur IV, distinctly stouter than other setae of same respective segments. Seta pd3 of basitarsus IV distinctly longer than other basitarsal setae. Pretarsus I absent; pretarsi II–IV similar in shape and length, each consisting of an elongate ambulacral stalk, a pair of strongly sclerotised claws and three rounded pulvillar lobes. Adult male. Not found. Material examined. Holotype female from soil at the base of Mimosa tenuiflora (Willd.) Poir (Fabaceae) at Arapiraca, Alagoas state, Brazil, March 24, 2012; one paratype female from soil at the base of Cupania oblongifolia Mart. (Sapindaceae) at Teotônio Vilela, Alagoas state, August 8, 2012; three paratype females from soil at the base of Tapirira guianensis Aublet (Anacardiaceae) at Teotônio Vilela, Alagoas state, November 15, 2011. Types collected by M.E. Duarte and J.C. Santos, and deposited in the Mite Reference Collection of Departamento de Entomologia e Acarologia, Escola Superior de Agricultura “Luiz de Queiroz” (ESALQ), Universidade de São Paulo (USP), Piracicaba, state of São Paulo, Brazil. AFRODACARELLUS FROM BRAZIL

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FIGURES 1–7. Afrodacarellus alagoensis Santos & Castilho n. sp. Female. 1. Chelicera; 2. Palp femur and genu; 3. Epistome; 4. Hypostome and palp trochanter; 5. Dorsal idiosoma; 6. Ventral idiosoma; 7. Tritosternum. Lyrifissures enlarged for improved visibility.

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FIGURES 8–11. Afrodacarellus alagoensis Santos & Castilho n. sp. Female. 8. Leg I; 9. Leg II; 10. Leg III; 11. Leg IV.

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Etymology. The specific name alagoensis is derived from Alagoas, the state where the type specimens were collected. Remarks. Afrodacarellus alagoensis is most similar to A. minutus Hurlbutt, 1974, A. pocsi Hurlbutt, 1974 and A. msituni Hurlbutt, 1974. Females of all of these species differ from the new species by having a single pair of opisthogastric setae off the ventrianal shield. In addition, females of A. minutus and A. pocsi differ by having sternal shield totally smooth, while female of A. minutus has 20 pairs of opisthonotal setae (R5 present).

Afrodacarellus xucurukariri Santos & Castilho n. sp. Diagnosis of adult female. Anterior region of epistome slightly acuminate, denticulate, with median denticle slightly longer than other denticles; podonotal shield smooth centrally; opisthonotal shield with 20 pairs of smooth setae and without punctate band along anterior margin; seta J2 at most 0.8 times as long as distance between its base and base of J3; seta J4 at most 0.7 times as long as distance between its base and base of J5; deutosternum narrow (distance h3–h3 about three times the length of distalmost transverse deutosternal line); without plates laterad of genital shield; ventrianal shield with five pairs of setae (including Jv1) in addition to circumanal setae; each side with a rounded and an elongate metapodal platelet, the former laterad of anterior end of the latter. Adult female (Fig. 12–23) Five specimens measured. Gnathosoma. Fixed cheliceral digit 25 (24–25) long, with five teeth in addition to apical tooth and a discreet setiform pilus dentilis (Fig. 12); movable cheliceral digit 25 (24–25) long, with two teeth in addition to apical tooth; antiaxial and dorsal lyrifissures as well as dorsal seta distinct, the latter thin. Arthrodial process of chelicera brush-shaped. Number of setae on palp trochanter–tarsus: 2–5–6–14–15; setae al1 and al2 of palp genu spatulate and smooth (Fig. 13) (al2 markedly more ventral and only slightly posteriad of al1); apotele 3-tined. Anterior region of epistome slightly acuminate, with margin denticulate, with median denticle slightly longer than other denticles (Fig. 14). Deutosternum narrow (distance h3–h3 over three times length of distalmost transverse deutosternal line), delimited by lateral lines, with eight transverse lines (Fig. 15), each with four (distalmost) or three denticles (others). With four pairs of smooth lines laterad of deutosternum, at level of fourth, sixth, seventh and eight transverse deutosternal lines (from distalmost). Internal malae fimbriate, separated from each other, shorter than corniculi, the latter horn-like, about three times as long as its basal width. Seta h3 about in longitudinal line with h1 and posteromediad of h2. Measurements of setae: h1 15 (10–16), h2 10 (8–12), h3 7 (6–9), sc 11 (11– 12); all setae aciculate and smooth. Dorsal idiosoma (Fig. 16). Idiosoma 289 (285–296) long and 168 (153–188) wide at widest level. Podonotal and opisthonotal shields separate. Podonotal shield reticulate laterally and smooth centrally, without transverse line next and parallel to posterior margin; with a delineated marginal strip containing the insertions of j1, z1, r2, r3, r5 and r6; 130 (127–133) long and 142 (140–145) wide at widest level; with 22 pairs of setae (j1–j6, z1–z6, s1–s6, r2, r3, r5 and r6), four pairs of distinguishable lyrifissures, three pairs of distinguishable pores and two pairs of scleronoduli between j5 and j6. Unsclerotised cuticle along lateral margins of podonotal shield with a pair of setae (r4). Opisthonotal shield reticulate and with a transverse line parallel to anterior margin delimiting a narrow anterior band smooth over most of its surface; with a delineated marginal strip containing the insertions of Z5, S5 and R1–R5; 145 (140–152) long and 120 (118–125) wide at widest level; with 20 pairs of setae (J1–J5, Z1–Z5, S1– S5 and R1–R5), ten pairs of distinguishable lyrifissures and two pairs of distinguishable pores. Measurements of setae: j1 11 (10–12), j2 10 (9–11), j3 9 (8–9), j4 10 (8–10), j5 8 (7–10), j6 9 (7–10), z1 5 (4–6), z2 11 (9–11), z3 10 (9–12), z4 10 (9–10), z5 9 (8–11), z6 9 (8–10), s1 9 (8–9), s2 9 (6–9), s3 11 (10–12), s4 11 (10–12), s5 11 (10–12), s6 10 (7–11), r2 13 (10–14), r3 14 (12–15), r4 7 (6–9), r5 10 (7–10), r6 13 (11–15), J1 8 (8–10), J2 8 (8–9), J3 8 (7–9), J4 9 (8–9), J5 10 (9–12), Z1 9 (8–10), Z2 9 (8–10), Z3 10 (9–11), Z4 11 (10–12), Z5 14 (12–16), S1 8 (7–10), S2 8 (7–9), S3 10 (10–11), S4 11 (10–12), S5 14 (13–16), R1 9 (8–10), R2 10 (9–10), R3 10 (9–12), R4 12 (11–13), R5 12 (11–15). All dorsal idiosomal setae aciculate, smooth; and much shorter than distance to subsequent setae of the same series. Ventral idiosoma (Fig. 17). Base of tritosternum 10 (9–10) long and 7 (6–7) wide proximally (Fig. 18); laciniae 35 (30–43), separated for about 70% of their total length, pilose. Sternal shield reticulate laterally and smooth centrally; posterior margin slightly concave; region anterior to the first pair of lyrifissures (iv1) lightly sclerotised and punctate; approximately 70 (69–75) long from center of anterior margin of punctate and lightly

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FIGURES 12–18. Afrodacarellus xucurukariri Santos & Castilho n. sp. Female. 12. Chelicera; 13. Palp genu; 14. Epistome; 15. Hypostome and palp trocanter; 16. Dorsal idiosoma; 17. Ventral idiosoma; 18; Tritosternum. Lyrifissures enlarged for improved visibility. AFRODACARELLUS FROM BRAZIL

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FIGURES 19–22. Afrodacarellus xucurukariri Santos & Castilho n. sp. Female. 19. Leg I; 20. Leg II; 21. Leg III; 22. Leg IV.

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sclerotised region to center of posterior margin and 60 (55–65) wide between coxae II and III; with four pairs of setae and three pairs of lyrifissures; seta st1 inserted in well sclerotised region of sternal shield. Discrete section of endopodal plate arched, extending from posterior region of coxa III to posterior region of coxa IV. Genital shield with few striae and a punctate band along straight posterior margin; extending posteriorly behind coxae IV; without plates laterad of genital shield; distance between st5–st5 45 (43–46). Lyrifissure iv5 on unsclerotised cuticle, laterad of st5. Ventrianal shield reticulate, except for a smooth band along anterior margin, 104 (102–108) long and 90 (70–98) wide at widest level; with five pairs of setae (Jv1–Jv3, Zv1 and Zv2) in addition to circumanal setae and one pair of marginal lyrifissures. Unsclerotised cuticle along lateral margins of ventrianal shield with two pairs of setae (Jv4 and Jv5) and two distinguishable pairs of lyrifissures. Peritreme extending anteriorly to posterior region of coxa II (region between r2 and r3). Peritrematic plate narrow, fused anteriorly to dorsal shield at level of r2; extending posteriorly as a narrow strip behind coxa IV; with one distinguishable lyrifissure and one distinguishable pore next to coxa III, and with two lyrifissures and one pore behind stigma. Exopodal plate fragmented into subtriangular platelets between coxae II–III and III–IV. With a pair of rounded and a pair of elongate metapodal platelets, the former with a pore-like structure and laterad of anterior end of the latter. Measurements of setae: st1 15 (14–15), st2 14 (12–15), st3 13 (12–15), st4 12 (11– 14), st5 14 (13–15), Jv1 12 (12–13), Jv2 14 (12–15), Jv3 13 (12–15), Jv5 15 (12–16), Zv1 11 (10–12), Zv2 12 (11–12), Zv3 11 (10–11), para-anal 15 (14–16), postanal 17 (15– 19). All ventral idiosomal setae aciculate and smooth. Spermathecal apparatus. Not distinguishable. Legs. (Figs 19–22) Lengths: I 260 (248–298); II 175 (163–188); III 143 (130–145); IV 198 (198–200). Chaetotaxy: I coxa 0 0/2 0/0 0, trochanter 1 0/11/2 1, femur 2 3/1 2/3 2, genu 2 3/2 3/1 2, tibia 2 3/2 3/2 2; II coxa 0 0/1 0/1 0, trochanter 1 0/1 0/2 1, femur 2 3/1 2/2 1, genu 2 3/1 2/1 2, tibia 2 2/1 2/1 2; III coxa 0 0/1 0/1 0, trochanter 1 1/1 0/2 0, femur 1 2/1 1/0 1, genu 2 2/1 2/1 1, tibia 2 1/1 2/1 1; IV coxa 0 0/1 0/0 0, trochanter 1 1/1 0/ 2 0, femur 1 2/1 2/0 0, genu 2 2/1 3/1 1, tibia 2 1/1 3/1 2; tarsi II–IV: 18, 18, 17. Setae av2 of tarsus II, pv2 of tarsus III, pv of genu and tibia IV, and pv2 of tarsus IV differing from other setae of the same segments by being distally divided. Seta av of genu II discreetly serrate. Pretarsus I absent; pretarsi II–IV similar in shape and length, each consisting of an elongate ambulacral stalk, a pair of strongly sclerotised claws and three rounded pulvillar lobes. Adult male. Not found. Material examined. Holotype female from litter at the base of Byrsonima crispa Juss. (Malpighiaceae), Palmeira dos Índios, Alagoas, Brazil, (09º23'44" S, 36º37'48" W), July 1, 2013; four paratype females from soil at the base of Spondias purpurea L. (Anacardiaceae) (09º22'929"S and 36º37'751"W), May 7 and July, 27 2013, all at Palmeira dos Índios, Alagoas, Brazil. All specimens collected by M.D. dos Santos and deposited in the Mite Reference Collection of the Departamento de Entomologia e Acarologia, Escola Superior de Agricultura "Luiz de Queiroz" (ESALQ), Universidade de São Paulo (USP), Piracicaba, state of São Paulo, Brazil. Etymology. The specific name xucurukariri is the name of a Brazilian Indian tribe from Alagoas state. Remarks. Afrodacarellus xucurukariri is most similar to A. myersi Loots, 1969, but females of the latter species differ by being about 50% smaller and having most dorsal shield setae proportionally shorter; by having wider deutosternum (distance h3–h3 about twice the length of distalmost transverse line), rounded and elongate metapodal platelets of each side about in line, and setae av2 of tarsus II, pv2 of tarsus III and pv2 of tarsus IV undivided.

Key to the world species of Afrodacarellus, based on adult females The key subsequently provided is an update of that provided by Hurlbutt (1974), with the inclusion of species described after that publication and of species transferred to that genus by Castilho et al. (2012). Aside from the new species here described and from A. citri, previously reported from Brazil (Castilho & Moraes, 2010), the key was prepared based solely on literature information. This is certainly a limitation of the key, given that descriptions may in some cases not be sufficiently accurate. Thus, characters more prone to intraspecific variations were avoided in the key. Afrodacarellus succinctus (Berlese, 1916) was not included in the key because of insufficient information about this species in the literature.

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Ventrianal shield with four pairs of opisthogastric setae in addition to circumanal setae; dorsal shield setae feather-shaped. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Afrodacarellus bipilosus (Karg, 1979); Argentina Ventrianal shield with more than four pairs of opisthogastric setae in addition to circumanal setae; dorsal shield setae not feather-shaped . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2 Seta Jv1 off ventrianal shield (but next to its anterior margin) . . . . . . . . . . . . .Afrodacarellus euungulae (Karg, 2003); Ecuador Seta Jv1 on ventrianal shield. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3 First two pairs of ventrianal setae in about a transverse line . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4 Second pair of ventrianal setae well posteriad of first pair . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8 Anterior margin of epistome with a relatively short medial spine-like projection, without lateral extensions; movable cheliceral digit with two teeth . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5 Anterior margin of epistome with three relatively elongate, denticulate projections; movable cheliceral digit with five teeth . 6 Distance h3–h3 about twice the length of distalmost transverse deutosternal line; rounded and elongate metapodal platelets of each side about in line; av2 of tarsus II, pv2 of tarsus III and pv2 of tarsus IV undivided . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Afrodacarellus myersi (Loots, 1969); South Africa Distance h3–h3 over three times the length of distalmost transverse deutosternal line; rounded metapodal platelet laterad of anterior end of elongate metapodal platelet; av2 of tarsus II, pv2 of tarsus III and pv2 of tarsus IV distally divided . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Afrodacarellus xucurukariri Santos & Castilho n. sp.; Brazil Setae of the J, Z and S series respectively about 20, 30 and 30 µm long; with a single pair of elongated metapodal platelets . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .Afrodacarellus furculatus (Karg, 1979); Argentina Setae of the J, Z and S series about 13–22 µm long; with a pair of elongate and a pair of ellipsoidal to rounded metapodal platelets . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7 Podonotal shield with 22 pairs of setae (s1 and r1 absent); opisthonotal shield imbricate; sternal shield reticulate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Afrodacarellus squamosus (Karg, 1977); Argentina Podonotal shield with 23 pairs of setae (r1 absent); opisthonotal shield reticulate; sternal shield smooth . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Afrodacarellus unospinae (Karg, 2003); Ecuador Anterior margin of epistome with medial projection spine-like . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9 Anterior margin of epistome with medial projection distally expanded and with few terminal tines . . . . . . . . . . . . . . . . . . . . 12 With a pair of relatively large, boomerang-shaped plates laterad of st5, bearing iv5 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Afrodacarellus citri (Loots, 1969); South Africa Without large, boomerang-shaped plates laterad of st5 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10 Anterior margin of epistome with medial projection sided by a pair of similar projections; ventrianal shield oval, without anterolateral corners. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Afrodacarellus lubalensis (Loots, 1969); Angola Anterior margin of epistome with medial projection sided only by denticles; ventrianal shield subpentagonal, with distinct anterolateral corners . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11 Base of anteromedian extension of epistome narrow (ratio length: basal width about 5:1); on each side, rounded metapodal platelet laterad of elongate metapodal platelet . . . . . . . . . . . . . . . . . . . . . . . . Afrodacarellus mongii (Hurlbutt, 1974); Tanzania Base of anteromedian extension of epistome broad (ratio length: basal width about 1:1); on each side, rounded metapodal platelet in line with elongate metapodal platelet . . . . . . . . . . . . . . . . . . . . . . . .Afrodacarellus bakeri (Hurlbutt, 1974); Tanzania Opisthonotal shield smooth. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 13 Opisthonotal shield reticulate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 14 Opisthonotal shield with depression behind J4; ventrianal shield reticulate . . . . . Afrodacarellus pili Hurlbutt, 1974; Tanzania Opisthonotal shield without depression behind J4; ventrianal shield smooth . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Afrodacarellus longipodus Hurlbutt, 1974; Tanzania With a single pair of tiny rounded metapodal platelets. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 15 With a pair of elongate and a pair of ellipsoidal to rounded metapodal platelets, which may be fused to each other or discrete . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 17 Anterior margin of sternal shield concave; anterior margin of epistome with a pair of short spine-like lateral projections (almost half as long as medial extension) separated from each other by a distance corresponding to over twice their length . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Afrodacarellus lunguensis (Ryke & Loots, 1966); Congo Anterior margin of sternal shield straight; anterior margin of epistome denticulate, with denticles of about uniform length and distinctly shorter than half the length of the medial projection; if a pair of denticles is slightly longer than others, the former are close together. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 16 With a pair of spur-like hypostomal processes posterolaterad of sc . . . . . .Afrodacarellus leleupi (Ryke & Loots, 1966); Congo Without spur-like hypostomal processes. . . . . . . . . . . . . . . . . . . . . . . . . Afrodacarellus kivuensis (Ryke & Loots, 1966); Congo Most dorsal shield setae lightly spatulate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Afrodacarellus reticulatus (Loots, 1969); Angola No dorsal shield setae spatulate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 18 Metapodal platelets of each side fused to each other . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 19 Metapodal platelets of each side separate from each other . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 20 Opisthonotal shield partially reticulate; with five pairs of opisthogastric setae on ventrianal shield in addition to circumanal setae (Zv3 off the shield) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Afrodacarellus machadoi (Loots, 1969); Angola Opisthonotal shield with transverse lines (except for scant diagonal lines); with six pairs of opisthogastric setae on ventrianal shield in addition to circumanal setae (Zv3 on the shield) . . Afrodacarellus filofissus (Karg & Schorlemmer, 2009); Venezuela Setae J1 and J2 distinctly shorter than distance between their bases and bases of J2 and J3, respectively . . . . . . . . . . . . . . . . 21 Setae J1 and J2 at least as long as distance between their bases and bases of J2 and J3, respectively . . . . . . . . . . . . . . . . . . . 26

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21. 22.

23. 24.

25.

26. 27.

28.

29.

30.

Podonotal shield extensively reticulate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 22 Podonotal shield at most with scant reticulation near posterolateral corners . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 23 Anterior margin of well sclerotised part of sternal shield concave; with two conspicuous lines starting near center of anterior margin of the shield and running posteriorly, deflecting abruptly laterally behind st2. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .Afrodacarellus concavus Hurlbutt, 1974; Tanzania Anterior margin of well sclerotised part of sternal shield about straight; without such line . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .Afrodacarellus ruwenzoriensis (Loots, 1969); Uganda Basitarsus IV with four setae (pl3 present) . . . . . . . . . . . . . . . . . . . . . . . . . Afrodacarellus camaxiloensis (Loots, 1969); Angola Basitarsus IV with three setae (pl3 absent) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 24 Seta J3 longer than distance between its base and base of J4; J1 over four-fifths as long as distance between its base and base of J2. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Afrodacarellus mossi Hurlbutt, 1974; Tanzania Seta J3 shorter than distance between its base and base of J4; J1 less than two–thirds as long as distance between its base and base of J2. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 25 Femur IV with seven setae, including two ventral setae; ventrianal shield with six pairs of setae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .Afrodacarellus ngorongoroensis Hurlbutt, 1974; Tanzania Femur IV with six setae; ventrianal shield with five pairs of setae . . . . . . . Afrodacarellus femoratus Hurlbutt, 1974; Tanzania Seta pd2 of tarsus III barbed; scleronoduli distinctly behind level of z5 . . . Afrodacarellus novembus Hurlbutt, 1974; Tanzania Seta pd2 of tarsus III not barbed; scleronoduli about in transverse line with z5. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 27 Anteromedian extension of tectum over three times as long as spine-like extentions laterad to it; podonotal shield with minute punctation . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Afrodacarellus pocsi Hurlbutt, 1974; Tanzania At least closest anterolateral extensions of tectum over half as long as anteromedian extension; podonotal shield smooth. . . 28 Lines laterad of deutosternum at level between sixth and seventh transverse deutosternal lines denticulate; extensions adjacent to anteromedian extension of tectum at least three times as long as more lateral extensions. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Afrodacarellus lupangaensis Hurlbutt, 1974; Tanzania Lines laterad of deutosternum at level between sixth and seventh transverse deutosternal lines smooth; extensions adjacent to anteromedian extension of tectum not over twice as long as more lateral extensions . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 29 Podonotal shield with 20 pairs setae (R5 present); setae Zv2 and Zv3 distally split; sternal shield totally smooth . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .Afrodacarellus minutus Hurlbutt, 1974; Tanzania Podonotal shield with 19 pairs setae (R5 absent); setae Zv2 and Zv3 distally split or not; sternal shield reticulate laterally and smooth centrally . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 30 Opisthogastric integument with eight setae, two of which off ventrianal shield; setae Zv2 and Zv3 distally split. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Afrodacarellus alagoensis Santos & Castilho n. sp.; Brazil Opisthogastric integument with seven setae, one of which off ventrianal shield; setae Zv2 and Zv3 not split distally . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Afrodacarellus msituni Hurlbutt, 1974; Tanzania

Discussion Karg (1977) ignored the description of Afrodacarellus by Hurlbutt (1974), and divided Afrogamasellus (including the species that had been transferred by the latter author to Afrodacarellus) into four subgenera, namely Afrogamasellus (Podalogamasellus), Afrogamasellus (Jugulogamasellus), Afrogamasellus (Foliogamasellus) and Afrogamasellus (Latogamasellus). The first two subgenera referred truly to Afrogamasellus, but the others were synonymised by Castilho et al. (2012) under Afrodacarellus. Acceptance of the latter two subgenera is not considered appropriate in the present publication, because they were not adequately characterised in the original descriptions. One of the diagnostic characteristics proposed by Karg (1977) to separate the subgenera, insertion of st1, does not seem reliable because of the continuous variation between two extremes: distinctly in the punctate area anteriad of the sclerotised shield (typical of Afrogamasellus (Foliogamasellus)) to distinctly in the sclerotised shield (typical of Afrogamasellus (Latogamasellus)). In several Afrodacarellus species, st1 is very close to the anterior margin of the sclerotised sternal shield (on or off it), hampering their classification at subgeneric level. An addition characteristic used by Karg (1977) to separate the subgenera, shape of epistome, was not adequately stated. The author mentioned Afrogamasellus (Foliogamasellus) to have a “complex epistome, with anteromedian extension frequently leaf-shaped”. The original description of the type species of the subgenus, A. camaxiloensis, shows five anterior extensions, the median of which distally enlarged and denticulate. Except for A. citri, all other species included by Karg (1977) in that subgenus have similar anteromedian extension, but the number of other anterior extensions is quite variable, some having none. Several species with st1 off the sclerotised sternal shield have epistome either with a single anterior extension (though with anterolateral margin denticulate) or anteromedian extension acuminate. In characterising Afrogamasellus (Latogamasellus), Karg (1977) mentioned the epistome to have 1–3 anteromedian extensions, without specifying the shape of the median extension. The AFRODACARELLUS FROM BRAZIL

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species included by the author in this subgenus include A. squamosus, the type species, whose medial extension is similar to that of the type species of Afrogamasellus (Foliogamasellus).

References Abo-Shnaf, R.I.A., Castilho, R.C. & Moraes, G.J. (2013) Two new species of Rhodacaridae (Acari: Mesostigmata) from Egypt and a key to the species of the family from the Mediterranean region. Zootaxa, 3718 (1), 28–38. https://doi.org/10.11646/zootaxa.3718.1.2 Antony, L.M.M.K. d'A. (1986) A Phylogenetic Analysis of the Rhodacaroidea (Acari: Mesostigmata). PhD thesis, Ohio State University, Columbus, x + 262 pp. Berlese, A. (1916) Centuria seconda di Acari nuovi. Redia, 12, 125–177. Castilho, R.C. & Moraes, G.J. (2010) Rhodacaridae mites (Acari: Mesostigmata: Rhodacaroidea) from the state of São Paulo, Brazil, with descriptions of a new genus and three new species. International Journal of Acarology, 36, 387–398. https://doi.org/10.1080/01647951003766970 Castilho, R.C., Moraes, G.J. & Halliday, B. (2012) Catalogue of the mite family Rhodacaridae Oudemans, with notes on the classification of the Rhodacaroidea (Acari: Mesostigmata). Zootaxa, 69, 1–69. Castilho, R.C., Venancio, R. & Narita, J.P.Z. (2015) Mesostigmata as biological control agents, with emphasis on Rhodacaroidea and Parasitoidea. In: Carrillo, D., Moraes, G.J. & Peña, J.E. (Eds.), Prospects for biological control of plant feeding mites and other harmful organisms. Springer, Florida, pp. 1–31. https://doi.org/10.1007/978-3-319-15042-0_1 Castilho, R.C., Moraes, G.J., Silva, E.S. & Silva, L.O. (2009) Predation potential and biology of Protogamasellopsis posnaniensis Wisniewski & Hirschmann (Acari: Rhodacaridae). Biological Control, 48, 164–167. https://doi.org/10.1016/j.biocontrol.2008.10.004 Duarte, A.F., Castilho, R.C., Cunha, U.S. & Moraes, G.J. (2016) A new species of Binodacarus (Acari: Mesostigmata: Rhodacaridae), with a new characterisation of the genus. Systematic & Applied Acarology, 21, 1194–1201. https://doi.org/10.11158/saa.21.9.4 Evans, G.O. (1963) Observations on the chaetotaxy of the legs in the free–living Gamasina (Acari: Mesostigmata). Bulletin of the Britsh Museum of Natural History, 10, 275–303. https://doi.org/10.5962/bhl.part.20528 Hurlbutt, H.W. (1974) The Afrogamasellus Loots & Ryke and Afrodacarellus n. gen. (Acarina: Rhodacaridae) of Tanzania. Acarologia, 15, 565–615. Karg, W. (1977) Zur Kenntnis einiger Milbengattungen der Rhodacaridae Oudemans, 1902 (Acarina, Parasitiformes). Teil 1. Zoologische Jahrbucher, Abteilung fur Systematik, Oekologie und Geographie der Tiere, 104, 327–351. Karg, W. (1979) Zur Kenntnis einiger Milbengattungen der Rhodacaridae Oudemans, 1902 (Acarina, Parasitiformes). Teil 2. Zoologische Jahrbucher, Abteilung fur Systematik, Oekologie und Geographie der Tiere, 106, 197–213. Karg, W. (2003) Die Raubmilbengattungen Afrogamasellus Loots et Ryke und Oloopticus Karg mit zwei neuen Arten-Ein Beitrag zur Evolution der Bodenmilben (Acarina, Gamasina). Abhandlungen und Berichte des Naturkundemuseums Görlitz, 75, 23–33. Karg, W. & Schorlemmer, A. (2009) New insights into predatory mites (Acarina, Gamasina) from tropical rain forests with special reference to distribution and taxonomy. Zoosystematics and Evolution, 85, 57–91. https://doi.org/10.1002/zoos.200800016 Lindquist, E.E. & Evans, G.O. (1965) Taxonomic concepts in the Ascidae, with a modified setal nomenclature for the idiosoma of the Gamasina (Acarina: Mesostigmata). Memoirs of the Entomological Society of Canada, 47, 5–66. https://doi.org/10.4039/entm9747fv Loots, G.C. (1969b) The tetrastigma species group of the genus Afrogamasellus (Acari: Rhodacaridae) from Central Africa. Revue de Zoologie et de Botanique Africaines, 79, 359–385. Rueda-Ramirez, D., Castilho, R.C. & Moraes, G.J. (2013) Mites of the superfamily Rhodacaroidea (Acari: Mesostigmata) from Colombia, with a key for the world species of Desectophis Karg (Ologamasidae). Zootaxa, 3734 (5), 521–535. https://doi.org/10.11646/zootaxa.3734.5.2 Walter, D.E., Hunt, H.W. & Elliott, E.T. (1988) Guilds or functional groups? An analysis of predatory arthropods from a shortgrass steppe soil. Pedobiologia, 31, 247–260.

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