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Ten new species of the genus Klinckowstroemia are described from passalid beetles (Coleoptera: Passalidae) collected in eastern and southern Mexico. The new species are Klinckowstroemia bifurcata sp. nov., K. hunteri sp. nov., K. valdezi sp. nov., K. montanoi sp. nov., K. brevimarginata sp. nov., K. rectimarginata sp. nov., K. franckei sp. nov., K. santibanezi sp. nov., K. ballesterosi sp. nov. and K. cristinae sp. nov. In some cases two species of Klinckowstroemia occurred on a single individual beetle and in one case an individual specimen of Odontotaenius zodiacus carried three species of mites: K. hunteri, K. bifurcata and K. valdezi. Eleven species of passalid beetles are new hosts for Klinckowstroemia. The beetle species associated with the most species of mites was Odontotaenius zodiacus, with six species; the other beetle species carried one or two mite species each. Mites were found in three areas of the passalid’s body: the coxae, the mesoepisternum and the humeri. Some species of Klinckowstroemia were found on more than one host species: K. valdezi and K. santibanezi were associated with four and three species of passalids respectively. This work increases the number of described species of the genus Klinckowstroemia from 14 to 24 species. The number of species of passalid beetles associated with this mite genus increased from 22 to 33.

Klinckowstroemia rectimarginata Villegas-Guzman, Pérez & Reyes-Castillo, 2009: 25 — Holotype female (CNAC, 006597), from Paxillus leachi MacLeay, Dos Amates, Municipality Catemaco, Veracruz, Mexico (18°29.35'N, 95°03.58'W, 120 m).

Klinckowstroemia pennula Villegas-Guzman, Reyes-Castillo & Pérez, 2011: 21 — Holotype female (CNAC, 006706), from undescribed Pseudacanthus sp., mountain rain forest, 7 km E San Pedro-San Pablo Ayutla, Ixtlán de Juárez, Oaxaca, Mexico (17°35. 293'N, 96°27. 162'W, 2459 m).

Klinckowstroemia pennula Villegas-Guzman, Reyes-Castillo & Pérez, 2011: 21 — Holotype female (CNAC, 006706), from undescribed Pseudacanthus sp., mountain rain forest, 7 km E San Pedro-San Pablo Ayutla, Ixtlán de Juárez, Oaxaca, Mexico (17°35. 293'N, 96°27. 162'W, 2459 m)., Published as part of LIU, DONG, YI, TIAN-CI, XU, YUN & ZHANG, ZHI-QIANG, 2013, new mite species described during 2007 to 2012 3663, pp. 1-102 in Zootaxa 3663 (1) on page 26, DOI: 10.11646/zootaxa.3663.1.1, http://zenodo.org/record/5630595

LIU, DONG, YI, TIAN-CI, XU, YUN, ZHANG, ZHI-QIANG (2013): new mite species described during 2007 to 2012 3663. Zootaxa 3663 (1): 1-102, DOI: 10.11646/zootaxa.3663.1.1, URL: http://dx.doi.org/10.11646/zootaxa.3663.1.1

Klinckowstroemia brevimarginata Villegas-Guzman, Pérez & Reyes-Castillo, 2009: 19 — Holotype female (CNAC, 006585), from Verres aff. intermedius, oak forest, 101 km Federal highway Puerto Escondido-Oaxaca, Oaxaca, Mexico (16°28.156'N, 97°01.674'W, 1906 m).

Klinckowstroemia oconnori Villegas-Guzman, Reyes-Castillo & Pérez, 2011: 31 — Holotype female (NMNH), Cerro Azul, Panama, Panama (950 m)., Published as part of LIU, DONG, YI, TIAN-CI, XU, YUN & ZHANG, ZHI-QIANG, 2013, new mite species described during 2007 to 2012 3663, pp. 1-102 in Zootaxa 3663 (1) on page 26, DOI: 10.11646/zootaxa.3663.1.1, http://zenodo.org/record/5630595

Klinckowstroemia melissae Villegas-Guzman, Reyes-Castillo & Pérez, 2011: 25 — Holotype female (CNAC, 006729), from undescribed Pseudacanthus sp., mountain rain forest, 7 km. E San Pedro-San Pablo Ayutla, Ixtlán de Juárez, Oaxaca, Mexico (17 35. 293'N, 96 27. 162'W, 2459 m)., Published as part of LIU, DONG, YI, TIAN-CI, XU, YUN & ZHANG, ZHI-QIANG, 2013, new mite species described during 2007 to 2012 3663, pp. 1-102 in Zootaxa 3663 (1) on page 26, DOI: 10.11646/zootaxa.3663.1.1, http://zenodo.org/record/5630595

Klinckowstroemia stilla Villegas-Guzman, Reyes-Castillo & Pérez, 2011: 28 — Holotype female (CNAC, 006739), from Proculus goryi, mountain rain forest, Ejido Benito Juárez, Reserva de la Biosfera Volcán Tacan, Chiapas, Mexico (15°05. 966'N, 92°08. 600'W, 1687 m).

Klinckowstroemia valdezi Villegas-Guzman, Pérez & Reyes-Castillo, 2009: 15 — Holotype female (CNAC, 006558), from Proculejus hirtus (Truqui), pine-oak forest, 1.5 km N Viborillas, municipality Huayacocotla, Veracruz, Mexico (20°31.245'N, 98°29.923'W, 2328 m).

Klinckowstroemia montanoi Villegas-Guzman, Pérez & Reyes-Castillo, 2009: 11 — Holotype female (CNAC, 006538), from Verres aff. intermedius, oak forest, 8 km NW San Miguel Coatlán, Federal highway Cuixtla-San Miguel Coatlán, Oaxaca, Mexico (16°15.064'N, 96°41.629'W, 2170 m)., Published as part of LIU, DONG, YI, TIAN-CI, XU, YUN & ZHANG, ZHI-QIANG, 2013, new mite species described during 2007 to 2012 3663, pp. 1-102 in Zootaxa 3663 (1) on page 26, DOI: 10.11646/zootaxa.3663.1.1, http://zenodo.org/record/5630595

Klinckowstroemia rectimarginata Villegas-Guzman, Pérez & Reyes-Castillo, 2009: 25 — Holotype female (CNAC, 006597), from Paxillus leachi MacLeay, Dos Amates, Municipality Catemaco, Veracruz, Mexico (18°29.35'N, 95°03.58'W, 120 m)., Published as part of LIU, DONG, YI, TIAN-CI, XU, YUN & ZHANG, ZHI-QIANG, 2013, new mite species described during 2007 to 2012 3663, pp. 1-102 in Zootaxa 3663 (1) on page 27, DOI: 10.11646/zootaxa.3663.1.1, http://zenodo.org/record/5630595

Klinckowstroemia franckei Villegas-Guzman, Pérez & Reyes-Castillo, 2009: 26 — Holotype female (CNAC, 006603), from Odontotaenius zodiacus, pine-oak forest, Ejido Atepixco, Municipality Zacualtipan, Hidalgo, Mexico (20°35.466'N, 98°36.358'W, 1972 m)., Published as part of LIU, DONG, YI, TIAN-CI, XU, YUN & ZHANG, ZHI-QIANG, 2013, new mite species described during 2007 to 2012 3663, pp. 1-102 in Zootaxa 3663 (1) on page 26, DOI: 10.11646/zootaxa.3663.1.1, http://zenodo.org/record/5630595

Klinckowstroemia brevimarginata Villegas-Guzman, Pérez & Reyes-Castillo, 2009: 19 — Holotype female (CNAC, 006585), from Verres aff. intermedius, oak forest, 101 km Federal highway Puerto Escondido-Oaxaca, Oaxaca, Mexico (16°28.156'N, 97°01.674'W, 1906 m)., Published as part of LIU, DONG, YI, TIAN-CI, XU, YUN & ZHANG, ZHI-QIANG, 2013, new mite species described during 2007 to 2012 3663, pp. 1-102 in Zootaxa 3663 (1) on page 26, DOI: 10.11646/zootaxa.3663.1.1, http://zenodo.org/record/5630595

Klinckowstroemia hunteri Villegas-Guzman, Pérez & Reyes-Castillo, 2009: 8 — Holotype female (CNAC, 006530), from Odontotaenius zodiacus (Truqui), pine-oak forest, 2 km Mojonera-Tizapan, municipality Zacualtipan, Hidalgo, Mexico (20°38.107'N, 98°35.981'W, 2019 m).

Klinckowstroemia cristinae Villegas-Guzman, Pérez & Reyes-Castillo, 2009: 40 — Holotype female (CNAC, 006649), from Odontotaenius zodiacus, pine-oak forest, 2 km Federal highway Crucero-Tianguistengo, Hidalgo, Mexico (20°40.413'N, 98°40.255'W, 2080 m)., Published as part of LIU, DONG, YI, TIAN-CI, XU, YUN & ZHANG, ZHI-QIANG, 2013, new mite species described during 2007 to 2012 3663, pp. 1-102 in Zootaxa 3663 (1) on page 26, DOI: 10.11646/zootaxa.3663.1.1, http://zenodo.org/record/5630595

Klinckowstroemia ballesterosi Villegas-Guzman, Pérez & Reyes-Castillo, 2009: 39 — Holotype female (CNAC, 006640), from Odontotaenius zodiacus, pine-oak forest, Municipality of Soledad Atzompa, Soledad Atzompa, Veracruz, Mexico (18°44.712'N, 97°40.967'W, 2125 m)., Published as part of LIU, DONG, YI, TIAN-CI, XU, YUN & ZHANG, ZHI-QIANG, 2013, new mite species described during 2007 to 2012 3663, pp. 1-102 in Zootaxa 3663 (1) on page 26, DOI: 10.11646/zootaxa.3663.1.1, http://zenodo.org/record/5630595

Klinckowstroemia valdezi Villegas-Guzman, Pérez & Reyes-Castillo, 2009: 15 — Holotype female (CNAC, 006558), from Proculejus hirtus (Truqui), pine-oak forest, 1.5 km N Viborillas, municipality Huayacocotla, Veracruz, Mexico (20°31.245'N, 98°29.923'W, 2328 m)., Published as part of LIU, DONG, YI, TIAN-CI, XU, YUN & ZHANG, ZHI-QIANG, 2013, new mite species described during 2007 to 2012 3663, pp. 1-102 in Zootaxa 3663 (1) on page 27, DOI: 10.11646/zootaxa.3663.1.1, http://zenodo.org/record/5630595

Klinckowstroemia santibanezi Villegas-Guzman, Pérez & Reyes-Castillo, 2009: 32 — Holotype female (CNAC, 006620), from Proculejus sartorii, pine forest, Municipality of Ixtlán de Juárez, Oaxaca, Mexico (17°31.722'N, 96°30.678'W, 2742 m)., Published as part of LIU, DONG, YI, TIAN-CI, XU, YUN & ZHANG, ZHI-QIANG, 2013, new mite species described during 2007 to 2012 3663, pp. 1-102 in Zootaxa 3663 (1) on page 27, DOI: 10.11646/zootaxa.3663.1.1, http://zenodo.org/record/5630595

Klinckowstroemia bifurcata Villegas-Guzman, Pérez & Reyes-Castillo, 2009: 7 — Holotype female (CNAC, 006514), from Odontotaenius zodiacus (Truqui), pine-oak forest, 2 km Federal Highway Crucero-Tianguistengo, Hidalgo, Mexico (20°40.413'N, 98°40.255'W, 2080 m)., Published as part of LIU, DONG, YI, TIAN-CI, XU, YUN & ZHANG, ZHI-QIANG, 2013, new mite species described during 2007 to 2012 3663, pp. 1-102 in Zootaxa 3663 (1) on page 26, DOI: 10.11646/zootaxa.3663.1.1, http://zenodo.org/record/5630595

Klinckowstroemia multisetillosa Rosano & Huinerse conocía asociada a Proculus mniszechi Kaup en Guatemala. Hemos enonntrado K. multisetillosa asociada a poblaciones alopáticas de P. burmeisteri Kuwert y P- opacipenn/s (Thompson). K. miultisetillosa se conoce de 4 regiones de Guatemala y 1 de Honduras En el presente, parece rjue no hay flujo genético entre esas 5 áreas. Sólo una especie de Proculus es conocida de cada una de las áreas mencionadas. Puesto que este género no vuela, la distribución de acaros entre estas "islas geográficas" puede ser explicada en 2 formas: la primera implica una amplia y antigua disiribución y la segunda implica la distribución de los ácaros por foresia en otras especies de pasalidos que vuelan.

Key to the species of Klinckowstroemia based on females (modified from Rosario & Hunter, 1988). 1. Medial margins of latigynal shields wavy (Fig. 2)............................................................ 2 - Medial margins of latigynal shields straight or pointed, never wavy (Fig. 7 A)...................................... 8 2. Sternal and latigynal setae barbed and/or serrated............................................................. 4 - Sternal and latigynal setae smooth......................................................................... 3 3. All sternal setae of equal length.............................................................. K. simplesetosa - Sternal setae st 2, st 3 and st 4 are microsetae (Fig. 2 G).................................................. K. pennula 4. With four pairs of setae a; st 1 long and barbed, not serrated............................................... K. scotti - With one pair of setae a, st 1 serrated, not barbed............................................................. 5 5. Seta st 1 long (31���40) and serrated, setae st 3 and st 4 are medium and/or microsetae................................. 6 - Seta st 1 medium (21���29) and serrated, setae st 3 and st 4 are short....................................... K. montanoi 6. Setae st 3 of medium length.............................................................................. 7 - Setae st 3 and st 4 are microsetae................................................................... K. valdezi 7. Seta st 4 short................................................................................ K. victoriae - Seta st 4 a microseta (Fig. 4 H).................................................................... K. melissae 8. Sternal seta st 1 barbed or bifurcate........................................................................ 9 - Sternal seta st 1 smooth................................................................................ 19 9. Sternal seta st 1 barbed................................................................................. 10 - Sternal seta st 1 bifurcate........................................................................ K. bifurcata 10. Sternal seta st 2 barbed................................................................................. 11 - Sternal seta st 2 smooth................................................................................. 14 11. Sternal setae st 3 and st 4 smooth.................................................................. K. candidoi - Sternal setae st 3 and st 4 barbed and serrated................................................................ 12 12. Sternal seta st 1 very long, st 2 and st 3 medium; with 6���10 serrated setae on latigynal shields................ K. santibanezi - Sternal setae st 1, st 2 and st 3 are long and serrated........................................................... 13 13. Sternal shield granulated..................................................................... K. ballesterosi - Sternal shield reticulated (Fig. 7 A)............................................................... K. oconnori 14. Mesogynal shield rounded anteriorly.......................................................... K. atramaculata - Mesogynal shield pointed anteriorly...................................................................... 15 15. Medial margins of latigynal shields pointed..................................................... K. tapachulensis - Medial margins of latigynal shields truncate............................................................... 16 16. Seta st 2 short or medium; body more than 900 ��m long...................................................... 17 - Seta st 2 microseta, body less than 900 ��m long............................................................. 18 17. Seta st 1 is the longest of sternal setae............................................................. K. schusteri - Seta st 3 is the longest of sternal setae.............................................................. K. cristinae 18. Latigynal shields with 4���6 setae setae st 3 and st 4 microsetae..................................... K. brevimarginata - Latigynal shields with three setae, setae st 3 and st 4 short......................................... K. rectimarginata 19. Two pairs of setae on latigynal shield............................................................. K. tragardhi - More than two pairs of setae on latigynal shields............................................................ 20 20. Posterior margin of ventral shield concave......................................................... K. concava - Posterior margin of ventral shield truncate................................................................. 21 21. Mesogynal shield extending anteriorly to sternogynal shield; medial margins of latigynal shields pointed............... 22 - Mesogynal shield not extending anteriorly to sternogynal shield; medial margins of latigynal shields truncated........... 26 22. Mesogynal shield extending anteriorly to sternogynal shield slightly............................................ 23 - Mesogynal shield extending anteriorly to sternogynal shield deeply............................................. 24 23. Seta st 1 microseta, st 2 short and simple, hyp 3 serrated................................................ K. franckei - Seta st 1 short and simple, st 2 microseta, hyp 3 simple.................................................. K. hunteri 24. Lateral margins of mesogynal shield concave; all sternal setae of equal length......................... K. multisetillosa - Lateral margins of mesogynal shield straight............................................................... 25 25. Sternal setae medium length, sternal and sternogynal shields reticulated..................................... K. starri - Sternal setae are microsetae, sternal and sternogynal shields granulated (Fig. 6 C).............................. K. stilla 26. Lateral margins of mesogynal shield concave; five or six pairs of setae on latigynal shield.................. K. grabowskii - Lateral margins of mesogynal shield straight; three or four pairs of setae on latigynal shield.......................... 27 27. Sternal seta st 2 shorter than st 3 and st 4; three pairs of setae on latigynal shields.............................. K. reyesi Sternal seta st 2 longer than st 3 and st 4; four pairs of setae on latigynal shields.............................. K. truncata, Published as part of Villegas-Guzm��n, Gabriel A., Reyes-Castillo, Pedro & P��rez, Tila M., 2011, New species of Klinckowstroemia Baker & Wharton, 1952 (Acari: Klinckowstroemiidae), associated with passalid beetles from Mexico and Panama, pp. 19-36 in Zootaxa 2747 on pages 35-36, DOI: 10.5281/zenodo.207115, {"references":["Rosario, R. M. T. & Hunter, P. E. (1988) The genus Klinckowstroemia Tragardh and descriptions of nine new species (Acarina: Trigynaspida: Klinckowstroemiidae). Acarologia, 29, 119 - 136."]}

Klinckowstroemia stilla sp. nov. Female (n = 5) Body oval. Idiosoma, L = 563 (557–595); W = 397 (397–437) (Fig. 5 A). Gnathosoma with different types of setae hyp 1 long and simple, L = 31 (30–34). Seta hyp 2 simple, L = 19. Seta hyp 3 simple, L = 9 (Fig. 6 A), and hyp 4 simple, L = 12 (8–12). Chelicera with excrescences, movable digit with five teeth, two small and three large; fixed digit with six teeth. Most palpal setae simple, but av 1 and av 2 of trochanter are branched and serrate respectively. On femur al 1 serrate. Anterior margin of trochanter palp with blunt process (b) and spine-like process (s). Hyaline hood extending to level of coxa I, seta a 1 short and simple, L = 12 (7–12). Tritosternum base wider than long. Presternal L = 25 (22–25), anterior W = 78 (78–81), shagreened (Fig. 6 D); anterior margin of the shield slightly concave medially, without presternal notch. Sternal seta st 1 micro seta (Fig. 6 E), placed near to anterior margin, and lyriform sternal pore stp 1 near to posterior margin. Sternal shield (Fig. 6 C) L = 47 (37–56) shagreened, sternal seta st 2 microseta placed near to anterior margin, stp 2 below, almost in the centre of the shield. Sternal setae st 3 and st 4 microsetae, near posterior margin of shield. Sternogynal shield L = 31 (31–34), W = 109 (109–124), triangular, posterior margin narrow and slightly pointed at apex. Surface shagreened; stp 3 located closer to anterior margin. Sternogynal apodeme with broad base that reaches to the shield, posterior arms thin, extending posteriorly to level of latigynal and mesogynal condyles. Latigynal shields shagreened, L = 90 (84–90); W = 56, each with an oval pore and four simple setae, medial margin of shield pointed (Fig. 6 B). Latigynal apodeme thick and separate from sternogynal shield. Mesogynal shield L = 81 (81–93), W = 62 (62–81), drop-shaped, extending anteriorly to sternogynal shield (Fig. 6 B), posterior margin rounded and wider, surface shagreened. Mesogynal condyles extending almost to the middle of the shield. Ventral shield, L = 171 (164–171); posterior W = 317 (317–341), shield triangular (Fig. 6 G), truncated posteriorly, surface shagreened, with 12 pairs of small setae (Fig. 6 H). Three pores underneath posterior margin of coxa IV (Fig. 3 A). Metapodal-peritrematal-exopodal shield (MPES) shagreened, with one pair of pores and one simple seta (Fig. 5 A). Anal shield, L = 87 (87–93); anterior W = 217 (217–257), with 5 pairs of simple setae (Fig. 6 H) plus longer antero-anal seta, L = 19 (12–19), two pairs of pores, one near anterior margin, and the other below the anus. Serrate setae of the leg I are: trochanter pv 1; femur pv 1, pv 2, and pv 3; genu I, al 1 and av 1; tibia av 1, and av 2 serrated. Serrate setae on leg II are: femur pv 1; genu av 1, and pv 1. Serrate setae leg III are: trochanter ad 1, and ad 2; femur av 1; genu pv 1. Serrated setae on leg IV are: trochanter pv 1; genu pv 1. Seta pv 1 of femur IV large, L = 36 (34–40) (Fig. 6 F). Male (n = 2) Body oval, similar to female. Idiosoma, L = 531–544, W = 371–384 (Fig. 5 B). Presternal shield, L = 31–34, anterior W = 56–62, shagreened, without presternal notch, anterior margin slightly concave (Fig. 5 B). Sternal seta st 1 microseta is near the middle of the shield. Sternal setae st 2, st 3, and st 4 are microsetae. Sternal shield shagreened, L = 34–43. Genital opening oval, L = 37, W = 50–53. Ventral shield with 30 setae (Fig. 5 B), L = 233, W = 295, shagreened but laterally reticulated, with 2–3 pores near to anterior margin of coxa IV. Anal shield, L = 93, W = 233, with four pairs of simple setae, plus longer antero-anal seta L = 12, with two pores: one near to anterior margin and the other below the anus. Hypostomal seta hyp 1 long and simple, L = 31–32, seta hyp 2 is simple and medium, L = 17–19, while seta hyp 3 is short and simple, L = 7–9. Seta pv 1 serrated on coxa IV, L = 33–34. Diagnosis. The mesogynal shield extending deeply to the sternogynal shield, with its posterior margin wide and rounded; and all the sternal setae, on female and male, are microsetae. Remarks. Klinckowstroemia stilla sp. nov. is similar to K. starri Rosario & Hunter and K. multisetillosa Rosario & Hunter, because they have the mesogynal shield extending deeply to the sternal shield, and the medial margins of latigynal shields pointed. K. stilla (L = 575, W = 409) is smaller than K. starri (L = 738, W = 541) and K. multisetillosa, (L = 936, W = 634); and sternal and sternogynal shields are reticulated in K. starri but shagreened in K. stilla and K. multisetillosa. Furthermore, K. stilla has all the sternal setae as microsetae, while K. starri has medium-size sternal setae, with st 1 serrated and the others simple. Klinckowstroemia multisetillosa has six pairs of anterior setae a and the lateral margins of mesogynal shield concave, but K. stilla only has one pair of setae a and the lateral margins of mesogynal shield straight. Etymology. This name is from Latin stilla (= drop) referring to the shape of the mesogynal shield which is similar a drop of water. Material examined. Proculus goryi: holotype Ƥ (CNAC 006739), 4 Ƥ (CNAC 006740- 6743) and 2 33 (CNAC 006744- 6745) paratypes from Chiapas, Mexico, Ejido Benito Juárez, Reserva de la Biosfera Volcán Tacan (15 0 5. 966 ’ N, 92 0 8. 600 ’ W), 1,687 m., 19 May 2007, mountain rain forest, ex one beetle, E. Chame, P. Reyes- Castillo, and B. Gomez; one female was found on the prosternum, and others in the alcohol. Holotype female, 4 paratype females, and 2 paratype males deposited in the CNAC. Ecology. The species K. stilla was collected on one specimen of Proculus goryi at one locality in Chiapas near the border with Guatemala. Another klinckowstroemiid species has been associated with the genus Proculus, namely Klinckowstroemia multisetillosa from Guatemala and Honduras on three different beetle species (Padilla & Schuster 1994).

Key to the species of Klinckowstroemia based on females (modified from Rosario & Hunter, 1988). 1. Medial margins of latigynal shields wavy (Fig. 2)............................................................ 2 - Medial margins of latigynal shields straight or pointed, never wavy (Fig. 7 A)...................................... 8 2. Sternal and latigynal setae barbed and/or serrated............................................................. 4 - Sternal and latigynal setae smooth......................................................................... 3 3. All sternal setae of equal length.............................................................. K. simplesetosa - Sternal setae st 2, st 3 and st 4 are microsetae (Fig. 2 G).................................................. K. pennula 4. With four pairs of setae a; st 1 long and barbed, not serrated............................................... K. scotti - With one pair of setae a, st 1 serrated, not barbed............................................................. 5 5. Seta st 1 long (31–40) and serrated, setae st 3 and st 4 are medium and/or microsetae................................. 6 - Seta st 1 medium (21–29) and serrated, setae st 3 and st 4 are short....................................... K. montanoi 6. Setae st 3 of medium length.............................................................................. 7 - Setae st 3 and st 4 are microsetae................................................................... K. valdezi 7. Seta st 4 short................................................................................ K. victoriae - Seta st 4 a microseta (Fig. 4 H).................................................................... K. melissae 8. Sternal seta st 1 barbed or bifurcate........................................................................ 9 - Sternal seta st 1 smooth................................................................................ 19 9. Sternal seta st 1 barbed................................................................................. 10 - Sternal seta st 1 bifurcate........................................................................ K. bifurcata 10. Sternal seta st 2 barbed................................................................................. 11 - Sternal seta st 2 smooth................................................................................. 14 11. Sternal setae st 3 and st 4 smooth.................................................................. K. candidoi - Sternal setae st 3 and st 4 barbed and serrated................................................................ 12 12. Sternal seta st 1 very long, st 2 and st 3 medium; with 6–10 serrated setae on latigynal shields................ K. santibanezi - Sternal setae st 1, st 2 and st 3 are long and serrated........................................................... 13 13. Sternal shield granulated..................................................................... K. ballesterosi - Sternal shield reticulated (Fig. 7 A)............................................................... K. oconnori 14. Mesogynal shield rounded anteriorly.......................................................... K. atramaculata - Mesogynal shield pointed anteriorly...................................................................... 15 15. Medial margins of latigynal shields pointed..................................................... K. tapachulensis - Medial margins of latigynal shields truncate............................................................... 16 16. Seta st 2 short or medium; body more than 900 µm long...................................................... 17 - Seta st 2 microseta, body less than 900 µm long............................................................. 18 17. Seta st 1 is the longest of sternal setae............................................................. K. schusteri - Seta st 3 is the longest of sternal setae.............................................................. K. cristinae 18. Latigynal shields with 4–6 setae setae st 3 and st 4 microsetae..................................... K. brevimarginata - Latigynal shields with three setae, setae st 3 and st 4 short......................................... K. rectimarginata 19. Two pairs of setae on latigynal shield............................................................. K. tragardhi - More than two pairs of setae on latigynal shields............................................................ 20 20. Posterior margin of ventral shield concave......................................................... K. concava - Posterior margin of ventral shield truncate................................................................. 21 21. Mesogynal shield extending anteriorly to sternogynal shield; medial margins of latigynal shields pointed............... 22 - Mesogynal shield not extending anteriorly to sternogynal shield; medial margins of latigynal shields truncated........... 26 22. Mesogynal shield extending anteriorly to sternogynal shield slightly............................................ 23 - Mesogynal shield extending anteriorly to sternogynal shield deeply............................................. 24 23. Seta st 1 microseta, st 2 short and simple, hyp 3 serrated................................................ K. franckei - Seta st 1 short and simple, st 2 microseta, hyp 3 simple.................................................. K. hunteri 24. Lateral margins of mesogynal shield concave; all sternal setae of equal length......................... K. multisetillosa - Lateral margins of mesogynal shield straight............................................................... 25 25. Sternal setae medium length, sternal and sternogynal shields reticulated..................................... K. starri - Sternal setae are microsetae, sternal and sternogynal shields granulated (Fig. 6 C).............................. K. stilla 26. Lateral margins of mesogynal shield concave; five or six pairs of setae on latigynal shield.................. K. grabowskii - Lateral margins of mesogynal shield straight; three or four pairs of setae on latigynal shield.......................... 27 27. Sternal seta st 2 shorter than st 3 and st 4; three pairs of setae on latigynal shields.............................. K. reyesi Sternal seta st 2 longer than st 3 and st 4; four pairs of setae on latigynal shields.............................. K. truncata

Klinckowstroemia melissae sp. nov. Female (n = 3) Body oval. Idiosoma, L = 736 (723–736); W = 531 (518–531) (Fig. 3 A). Gnathosoma with different types of setae: hyp 1 long and simple, L = 48 (47–50), longer than other three. hyp 2 serrate, L = 33 (33–34). hyp 3 short and serrate, L = 12 (12–14) (Fig. 4 B). hyp 4 short and simple, L = 11 (9–12). Chelicera with excrescences, movable digit with four teeth, two small and two larger; fixed digit with six teeth. Most palpal setae simple, but av 1 and av 2 of trochanter branched and serrate respectively. Anterior margin of palp trochanter with blunt process (b) and slightly longer spine-like process (s). Hyaline hood extending to level of coxa I, seta a 1 large and serrate, L = 25 (25–27). Tritosternum base wider than long. Presternal shield L = 37 (37–43), anterior W = 99 (96–99), shagreened (Fig. 4 C). Presternal notch shaped like a small square. Sternal seta st 1 serrate (Fig. 4 D), L = 27 (25–32), near to anterior margin; opposite margin has lyriform sternal pore stp 1. Sternal shield reticulated and shagreened (Fig. 4 E), L = 56, near the anterior margin of the shield is stp 2, below it is sternal seta st 2, L = 19 (16–19), serrate and medium. Near the posterior margin of the shield are sternal setae st 3, L = 16 (12–16) serrated, and st 4 is a microseta. Sternogynal shield triangular (Fig. 4 A) L = 47; W = 143, posterior margin narrow and pointed at apex, surface is shagreened and reticulated, stp 3 located closer to anterior margin. Sternogynal apodeme with broad base that reaches to the shield, posterior arms thick, extending posteriorly to level of latigynal and mesogynal condyles. Latigynal shields, L = 109 (102–109); W = 62, each with an oval pore and 4–5 setae (Fig. 4 A), medial margin of shield wavy and medium, this shield reticulated and shagreened. Latigynal apodeme thick and separate from the sternogynal shield. Mesogynal shield, L = 53 (53–56), W = 81, triangular, anterior margin rounded and posterior margin extended (Fig. 4 A), surface shagreened. Mesogynal condyles extending almost to level of shield apex. Ventral shield, L = 186 (186–192); posterior W = 428 (422–443), shield triangular, truncated posteriorly, surface reticulated and shagreened, with 13–14 pairs of small setae. One pore located posterior margin of coxa IV (Fig. 3 A). The metapodal-peritrematal-exopodal shield (MPES) reticulated, with a pair of pores and one simple seta (Fig. 3 A). Anal shield, L = 124 (124–130); anterior W = 360 (341–360), with 7–8 pairs of simple setae (Fig. 4 F) plus longer antero-anal seta, L = 25 (25–27), two pairs of pores, one near anterior margin, and the other below the anus. Serrate setae on leg I are: trochanter al 1; femur I, pv 2, and pv 3; genu pl 1. Serrrate setae on leg II are: femur pv 1; genu pv 1; and tibia al 4. Serrate setae on leg III are: femur av 1; genu av 1, and pv 1; tibia ad 4 and, tarsus ad 3. Serrate setae on leg IV: trochanter al 1, pv 1, and pl 1; femur av 1; genu av 1, pl 1, and pv 1; tibia ad 4; tarsus ad 2, and ad 3. Seta pv 1 of leg femur IV large, L = 28 (25–32). Male (n = 5) Body oval, similar to female. Idiosoma, L = 684–723, W = 506–531 (Fig. 3 B). Presternal shield, L = 50–56, anterior W = 93–96, is shagreened, presternal notch shaped like an inverted triangle, moderately deep (Fig. 3 B). Sternal seta st 1, L = 19–22, serrate, near to anterior margin. Sternal seta st 2, L = 14–19, serrated, seta st 3, L = 7–9 simple, and seta st 4 a microseta. Sternal shield reticulated, L = 62–76 (Fig. 3 B). Genital opening oval, L = 47–50, W = 71–74. Ventral shield with 32–34 setae (Fig. 3 B), L = 248–279, W = 403–419, shagreened, laterally reticulated. One pore near the anterior margin of coxa IV. Anal shield, L = 124–127, anterior W = 326–372, with 6–7 pairs of simple setae, plus longer antero-anal seta L = 25–31; with two pores, one near the anterior margin and the other below the anus. Hypostomal seta hyp 1 is long and simple, L = 47–56, seta hyp 2 is serrate and long, L = 33– 37, while seta hyp 3 is serrate and shorter, L = 9–16. Seta pv 1 serrated on coxa IV, L = 20–31. Diagnosis. Sternal, latigynal, sternogynal, and ventral shields are reticulated, and setae st 1, st 2, and st 3 are serrated. Mesogynal shield with posterior margin wide. Male has sternal shield reticulated and st 1 and st 2 serrated. Remarks. Klinckowstroemia melissae sp. nov. is similar to K. victoriae Rosario & Hunter and K. pennula sp. nov., because these species have latigynal shields with wavy medial margins. These species are distinguished by the genital shields, which in K. melissae are reticulated and K. victoriae are shagreened, and body length and width (K. melissae L = 723–736, W = 518–531; K. victoriae L = 834–872, W = 570–616). K. pennula and K. melissae share the number of setae on latigynal and ventral shields. These species are distinguished by length of sternogynal shield (K. pennula L = 50–62; K. melissae L = 28–34); and setae st 1 is simple in K. pennula, and serrated in K. melissae. Male in K. pennula has anterior margin of presternal shield concave and in K. pennula it is rounded. Etymology. This species is named in honour of Melissa Rodriguez, for her friendship, patience and support the senior author during this research; many thanks Bichita. Material examined. From undescribed Pseudacanthus sp.: holotype Ƥ (CNAC 006729), 4 Ƥ and 5 33 (CNAC 006730- 6738) paratypes from Oaxaca, Mexico, 7 km. E San Pedro-San Pablo Ayutla, Ixtlán de Juárez (17 35. 293 ’ N, 96 27. 162 ’ W), 2,459 m., 20 July 2007, mountain rain forest, ex three beetles, O. Francke, A. Valdez, C. Santibañez, A. Ballesteros, and H. Montaño. Mites were found on femur, near to coxa I, and in the alcohol. Holotype female, 4 paratype females, and 5 paratype males deposited in the CNAC. Ecology. The species K. melissae and K. pennula were found together on three specimens of an undescribed Pseudacanthus sp. Two of these passalids were living in the same log, and the other passalid beetle was collected in the same locality and day. Specimens of K. pennulae were more abundant than K. melissae; this is the first case where two species found on the same host (synhospitality) are almost the same size. In other known cases of synhospitality one mite species is larger than the other (Villegas-Guzman et al. 2009).

Villegas-Guzmán, Gabriel A., Reyes-Castillo, Pedro, Pérez, Tila M. (2011): New species of Klinckowstroemia Baker & Wharton, 1952 (Acari: Klinckowstroemiidae), associated with passalid beetles from Mexico and Panama. Zootaxa 2747: 19-36, DOI: 10.5281/zenodo.207115

Klinckowstroemia stilla sp. nov. Female (n = 5) Body oval. Idiosoma, L = 563 (557���595); W = 397 (397���437) (Fig. 5 A). Gnathosoma with different types of setae hyp 1 long and simple, L = 31 (30���34). Seta hyp 2 simple, L = 19. Seta hyp 3 simple, L = 9 (Fig. 6 A), and hyp 4 simple, L = 12 (8���12). Chelicera with excrescences, movable digit with five teeth, two small and three large; fixed digit with six teeth. Most palpal setae simple, but av 1 and av 2 of trochanter are branched and serrate respectively. On femur al 1 serrate. Anterior margin of trochanter palp with blunt process (b) and spine-like process (s). Hyaline hood extending to level of coxa I, seta a 1 short and simple, L = 12 (7���12). Tritosternum base wider than long. Presternal L = 25 (22���25), anterior W = 78 (78���81), shagreened (Fig. 6 D); anterior margin of the shield slightly concave medially, without presternal notch. Sternal seta st 1 micro seta (Fig. 6 E), placed near to anterior margin, and lyriform sternal pore stp 1 near to posterior margin. Sternal shield (Fig. 6 C) L = 47 (37���56) shagreened, sternal seta st 2 microseta placed near to anterior margin, stp 2 below, almost in the centre of the shield. Sternal setae st 3 and st 4 microsetae, near posterior margin of shield. Sternogynal shield L = 31 (31���34), W = 109 (109���124), triangular, posterior margin narrow and slightly pointed at apex. Surface shagreened; stp 3 located closer to anterior margin. Sternogynal apodeme with broad base that reaches to the shield, posterior arms thin, extending posteriorly to level of latigynal and mesogynal condyles. Latigynal shields shagreened, L = 90 (84���90); W = 56, each with an oval pore and four simple setae, medial margin of shield pointed (Fig. 6 B). Latigynal apodeme thick and separate from sternogynal shield. Mesogynal shield L = 81 (81���93), W = 62 (62���81), drop-shaped, extending anteriorly to sternogynal shield (Fig. 6 B), posterior margin rounded and wider, surface shagreened. Mesogynal condyles extending almost to the middle of the shield. Ventral shield, L = 171 (164���171); posterior W = 317 (317���341), shield triangular (Fig. 6 G), truncated posteriorly, surface shagreened, with 12 pairs of small setae (Fig. 6 H). Three pores underneath posterior margin of coxa IV (Fig. 3 A). Metapodal-peritrematal-exopodal shield (MPES) shagreened, with one pair of pores and one simple seta (Fig. 5 A). Anal shield, L = 87 (87���93); anterior W = 217 (217���257), with 5 pairs of simple setae (Fig. 6 H) plus longer antero-anal seta, L = 19 (12���19), two pairs of pores, one near anterior margin, and the other below the anus. Serrate setae of the leg I are: trochanter pv 1; femur pv 1, pv 2, and pv 3; genu I, al 1 and av 1; tibia av 1, and av 2 serrated. Serrate setae on leg II are: femur pv 1; genu av 1, and pv 1. Serrate setae leg III are: trochanter ad 1, and ad 2; femur av 1; genu pv 1. Serrated setae on leg IV are: trochanter pv 1; genu pv 1. Seta pv 1 of femur IV large, L = 36 (34���40) (Fig. 6 F). Male (n = 2) Body oval, similar to female. Idiosoma, L = 531���544, W = 371���384 (Fig. 5 B). Presternal shield, L = 31���34, anterior W = 56���62, shagreened, without presternal notch, anterior margin slightly concave (Fig. 5 B). Sternal seta st 1 microseta is near the middle of the shield. Sternal setae st 2, st 3, and st 4 are microsetae. Sternal shield shagreened, L = 34���43. Genital opening oval, L = 37, W = 50���53. Ventral shield with 30 setae (Fig. 5 B), L = 233, W = 295, shagreened but laterally reticulated, with 2���3 pores near to anterior margin of coxa IV. Anal shield, L = 93, W = 233, with four pairs of simple setae, plus longer antero-anal seta L = 12, with two pores: one near to anterior margin and the other below the anus. Hypostomal seta hyp 1 long and simple, L = 31���32, seta hyp 2 is simple and medium, L = 17���19, while seta hyp 3 is short and simple, L = 7���9. Seta pv 1 serrated on coxa IV, L = 33���34. Diagnosis. The mesogynal shield extending deeply to the sternogynal shield, with its posterior margin wide and rounded; and all the sternal setae, on female and male, are microsetae. Remarks. Klinckowstroemia stilla sp. nov. is similar to K. starri Rosario & Hunter and K. multisetillosa Rosario & Hunter, because they have the mesogynal shield extending deeply to the sternal shield, and the medial margins of latigynal shields pointed. K. stilla (L = 575, W = 409) is smaller than K. starri (L = 738, W = 541) and K. multisetillosa, (L = 936, W = 634); and sternal and sternogynal shields are reticulated in K. starri but shagreened in K. stilla and K. multisetillosa. Furthermore, K. stilla has all the sternal setae as microsetae, while K. starri has medium-size sternal setae, with st 1 serrated and the others simple. Klinckowstroemia multisetillosa has six pairs of anterior setae a and the lateral margins of mesogynal shield concave, but K. stilla only has one pair of setae a and the lateral margins of mesogynal shield straight. Etymology. This name is from Latin stilla (= drop) referring to the shape of the mesogynal shield which is similar a drop of water. Material examined. Proculus goryi: holotype �� (CNAC 006739), 4 �� (CNAC 006740- 6743) and 2 33 (CNAC 006744- 6745) paratypes from Chiapas, Mexico, Ejido Benito Ju��rez, Reserva de la Biosfera Volc��n Tacan (15 0 5. 966 ��� N, 92 0 8. 600 ��� W), 1,687 m., 19 May 2007, mountain rain forest, ex one beetle, E. Chame, P. Reyes- Castillo, and B. Gomez; one female was found on the prosternum, and others in the alcohol. Holotype female, 4 paratype females, and 2 paratype males deposited in the CNAC. Ecology. The species K. stilla was collected on one specimen of Proculus goryi at one locality in Chiapas near the border with Guatemala. Another klinckowstroemiid species has been associated with the genus Proculus, namely Klinckowstroemia multisetillosa from Guatemala and Honduras on three different beetle species (Padilla & Schuster 1994)., Published as part of Villegas-Guzm��n, Gabriel A., Reyes-Castillo, Pedro & P��rez, Tila M., 2011, New species of Klinckowstroemia Baker & Wharton, 1952 (Acari: Klinckowstroemiidae), associated with passalid beetles from Mexico and Panama, pp. 19-36 in Zootaxa 2747 on pages 28-31, DOI: 10.5281/zenodo.207115, {"references":["Padilla, E. E. & Schuster, J. (1994) Klinckowstroemia multisetillosa Rosario & Hunter (Acarina: Trygynaspida: Klinckowstroemiidae) associated with three species of Proculus Kuwert (Coleoptera: Passalidae). Acta Zoologica Mexicana (n. s.), 61, 1 - 15."]}

Klinckowstroemia melissae sp. nov. Female (n = 3) Body oval. Idiosoma, L = 736 (723���736); W = 531 (518���531) (Fig. 3 A). Gnathosoma with different types of setae: hyp 1 long and simple, L = 48 (47���50), longer than other three. hyp 2 serrate, L = 33 (33���34). hyp 3 short and serrate, L = 12 (12���14) (Fig. 4 B). hyp 4 short and simple, L = 11 (9���12). Chelicera with excrescences, movable digit with four teeth, two small and two larger; fixed digit with six teeth. Most palpal setae simple, but av 1 and av 2 of trochanter branched and serrate respectively. Anterior margin of palp trochanter with blunt process (b) and slightly longer spine-like process (s). Hyaline hood extending to level of coxa I, seta a 1 large and serrate, L = 25 (25���27). Tritosternum base wider than long. Presternal shield L = 37 (37���43), anterior W = 99 (96���99), shagreened (Fig. 4 C). Presternal notch shaped like a small square. Sternal seta st 1 serrate (Fig. 4 D), L = 27 (25���32), near to anterior margin; opposite margin has lyriform sternal pore stp 1. Sternal shield reticulated and shagreened (Fig. 4 E), L = 56, near the anterior margin of the shield is stp 2, below it is sternal seta st 2, L = 19 (16���19), serrate and medium. Near the posterior margin of the shield are sternal setae st 3, L = 16 (12���16) serrated, and st 4 is a microseta. Sternogynal shield triangular (Fig. 4 A) L = 47; W = 143, posterior margin narrow and pointed at apex, surface is shagreened and reticulated, stp 3 located closer to anterior margin. Sternogynal apodeme with broad base that reaches to the shield, posterior arms thick, extending posteriorly to level of latigynal and mesogynal condyles. Latigynal shields, L = 109 (102���109); W = 62, each with an oval pore and 4���5 setae (Fig. 4 A), medial margin of shield wavy and medium, this shield reticulated and shagreened. Latigynal apodeme thick and separate from the sternogynal shield. Mesogynal shield, L = 53 (53���56), W = 81, triangular, anterior margin rounded and posterior margin extended (Fig. 4 A), surface shagreened. Mesogynal condyles extending almost to level of shield apex. Ventral shield, L = 186 (186���192); posterior W = 428 (422���443), shield triangular, truncated posteriorly, surface reticulated and shagreened, with 13���14 pairs of small setae. One pore located posterior margin of coxa IV (Fig. 3 A). The metapodal-peritrematal-exopodal shield (MPES) reticulated, with a pair of pores and one simple seta (Fig. 3 A). Anal shield, L = 124 (124���130); anterior W = 360 (341���360), with 7���8 pairs of simple setae (Fig. 4 F) plus longer antero-anal seta, L = 25 (25���27), two pairs of pores, one near anterior margin, and the other below the anus. Serrate setae on leg I are: trochanter al 1; femur I, pv 2, and pv 3; genu pl 1. Serrrate setae on leg II are: femur pv 1; genu pv 1; and tibia al 4. Serrate setae on leg III are: femur av 1; genu av 1, and pv 1; tibia ad 4 and, tarsus ad 3. Serrate setae on leg IV: trochanter al 1, pv 1, and pl 1; femur av 1; genu av 1, pl 1, and pv 1; tibia ad 4; tarsus ad 2, and ad 3. Seta pv 1 of leg femur IV large, L = 28 (25���32). Male (n = 5) Body oval, similar to female. Idiosoma, L = 684���723, W = 506���531 (Fig. 3 B). Presternal shield, L = 50���56, anterior W = 93���96, is shagreened, presternal notch shaped like an inverted triangle, moderately deep (Fig. 3 B). Sternal seta st 1, L = 19���22, serrate, near to anterior margin. Sternal seta st 2, L = 14���19, serrated, seta st 3, L = 7���9 simple, and seta st 4 a microseta. Sternal shield reticulated, L = 62���76 (Fig. 3 B). Genital opening oval, L = 47���50, W = 71���74. Ventral shield with 32���34 setae (Fig. 3 B), L = 248���279, W = 403���419, shagreened, laterally reticulated. One pore near the anterior margin of coxa IV. Anal shield, L = 124���127, anterior W = 326���372, with 6���7 pairs of simple setae, plus longer antero-anal seta L = 25���31; with two pores, one near the anterior margin and the other below the anus. Hypostomal seta hyp 1 is long and simple, L = 47���56, seta hyp 2 is serrate and long, L = 33��� 37, while seta hyp 3 is serrate and shorter, L = 9���16. Seta pv 1 serrated on coxa IV, L = 20���31. Diagnosis. Sternal, latigynal, sternogynal, and ventral shields are reticulated, and setae st 1, st 2, and st 3 are serrated. Mesogynal shield with posterior margin wide. Male has sternal shield reticulated and st 1 and st 2 serrated. Remarks. Klinckowstroemia melissae sp. nov. is similar to K. victoriae Rosario & Hunter and K. pennula sp. nov., because these species have latigynal shields with wavy medial margins. These species are distinguished by the genital shields, which in K. melissae are reticulated and K. victoriae are shagreened, and body length and width (K. melissae L = 723���736, W = 518���531; K. victoriae L = 834���872, W = 570���616). K. pennula and K. melissae share the number of setae on latigynal and ventral shields. These species are distinguished by length of sternogynal shield (K. pennula L = 50���62; K. melissae L = 28���34); and setae st 1 is simple in K. pennula, and serrated in K. melissae. Male in K. pennula has anterior margin of presternal shield concave and in K. pennula it is rounded. Etymology. This species is named in honour of Melissa Rodriguez, for her friendship, patience and support the senior author during this research; many thanks Bichita. Material examined. From undescribed Pseudacanthus sp.: holotype �� (CNAC 006729), 4 �� and 5 33 (CNAC 006730- 6738) paratypes from Oaxaca, Mexico, 7 km. E San Pedro-San Pablo Ayutla, Ixtl��n de Ju��rez (17 35. 293 ��� N, 96 27. 162 ��� W), 2,459 m., 20 July 2007, mountain rain forest, ex three beetles, O. Francke, A. Valdez, C. Santiba��ez, A. Ballesteros, and H. Monta��o. Mites were found on femur, near to coxa I, and in the alcohol. Holotype female, 4 paratype females, and 5 paratype males deposited in the CNAC. Ecology. The species K. melissae and K. pennula were found together on three specimens of an undescribed Pseudacanthus sp. Two of these passalids were living in the same log, and the other passalid beetle was collected in the same locality and day. Specimens of K. pennulae were more abundant than K. melissae; this is the first case where two species found on the same host (synhospitality) are almost the same size. In other known cases of synhospitality one mite species is larger than the other (Villegas-Guzman et al. 2009)., Published as part of Villegas-Guzm��n, Gabriel A., Reyes-Castillo, Pedro & P��rez, Tila M., 2011, New species of Klinckowstroemia Baker & Wharton, 1952 (Acari: Klinckowstroemiidae), associated with passalid beetles from Mexico and Panama, pp. 19-36 in Zootaxa 2747 on pages 25-28, DOI: 10.5281/zenodo.207115, {"references":["Villegas-Guzman, G. A., Perez, T. M. & Reyes-Castillo, P. (2009) New species of the genus Klinckowstroemia Baker & Wharton from Mexico (Acari: Mesostigamata: Klinckowstroemiidae). Zootaxa, 2248, 1 - 46."]}

Klinckowstroemia oconnori sp. nov. Female (n = 3) Body oval. Idiosoma: L = 781 (781���806); W = 576 (563���576) (Fig. 7 A). Gnathosoma with different hypostomal setae: hyp 1 long and simple, L= 50 (47���54); seta hyp 2 long and serrated, L = 43 (40���43), seta hyp 3 medium and serrated, L = 16 (12���16), and seta hyp 4 medium-sized and serrated, L = 16 (9���16). Chelicera with excrescences, movable digit with four teeth and fixed digit with seven teeth. Palp setae are simple, but av 1 and av 2 on trochanter are branched and serrated, respectively; on femur pd 1, pd 2, and pv 1 are serrated. On genu the setae al 1, al 2, ad 1, pl 1, pd 1 and pv 1 are serrated. Hyaline hood extending to level of coxa I, seta a 1 moderate and simple, L = 19 (19���22). Tritosternum base wider than long. Presternal (Pst) L = 37 (34���37), anterior W= 105 (102���109) shagreened and with line near the posterior margin; near posterior margin is a lyriform sternal pore stp 1. Presternal notch is a small concave depression. Sternal seta st 1, L = 53 (34���53), serrated (Fig. 7 A). Sternal shield L = 59 (56���62), shagreened, with some reticulations (Fig. 7 A). Lyrifissure stp 2 near to anterior margin of the shield; st 2, L = 47, serrated, below. Sternal setae st 3, L = 37, and st 4, L = 40, both setae are large and serrated, and near to posterior margin of the shield. Intact sternal setae present in only one specimen. Sternogynal shield (SgS), L = 68 (62���68); W = 158 (140���158) triangular and shagreened, posterior margin straight and slightly pointed, near to anterior margin is pore stp 3. Sternogynal apodeme thin and short. Latigynal shields shagreened, L = 130 (127���130); W = 65 (62���65), each with an oval pore and four simple setae, medial margin of shield straight. Mesogynal shield, L = 62 (59���62); W = 83. 7, triangular and shagreened, posterior margin wider and slightly pointed at apex. Mesogynal condyles extending almost to level of shield apex. Ventral shield, L = 171 (171���189); posterior W = 419 (403���419), shield triangular and shagreened, truncated posteriorly, with ten pairs of small setae (Fig. 7 A), with two pairs of pores at the level of posterior margin of coxa IV. The metapodal-peritrematal-exopodal shield (MPES) shagreened, with one pair of pores and one simple seta. Anal shield, L = 140 (124���140); anterior W = 341 (316���341), with 4 pairs of simple setae plus longer anteroanal seta, L = 22 (22���25); two pairs of pores, one near the anterior margin, and the other below the anus (Fig. 7 A). Serrate setae on leg I are: trochanter ad 1, al 1 and pv 1; femur: av 3, pv 1 and pv 2; tibia ad 1, ad 2, ad 3, av 1, av 2, pl 1, and pl 2. Serrate setae on Leg II are: femur pv 1; genu pv 1. Serrate setae on leg III: trochanter ad 1 and ad 2; femur av 1; genu av 1, and pv 1; tibia ad 4; tarsus ad 3, and pl 1. Serrate setae on leg IV: femur av 1; genu av 1, and pv 1; tibia ad 4, and al 4; tarsus ad 2 and ad 3 serrated. Seta pv 1 of the femur IV large L= 28 (27���28). Male (n = 5) Body oval, similar to female. Idiosoma L = 742���806, W = 493���563 (Fig. 7 B). Presternal (Pst) L = 31���37, anterior W = 102���109, presternal notch V-shaped, shagreened and with one line near anterior margin (Fig. 8 B) as in female; near to posterior margin is a lyriform sternal pore stp 1. Sternal seta st 1, L = 40���56, long and serrated (Fig. 8 C). Sternal shield shagreened (Fig. 8 D) and reticulated (Fig. 8 E), L = 121���127, with two pores, one near the anterior margin (stp 2) and other near the genital opening (stp 3). Sternal seta st 2 long and serrated, L = 48���50; sternal seta st 3 long and serrated, L = 39���47; sternal seta st 4 medium and serrated, L = 20���33. Genital opening oval, L = 50���62, W = 68���71. Ventral shield shagreened and reticulated, L = 264���285, W= 372���450 (Fig. 8 H), with 30 setae and two pores, one above the anterior margin of coxa IV, and other below coxa IV. Anal shield L = 102���140, anterior W = 322���363 (Fig. 8 G); with four pairs of simple setae plus longer antero-anal seta, L = 12���25, with two pores: one near the anterior margin and another below the anus. Hypostomal setae (Fig. 8 A): hyp 1 long and simple, L = 43���56; seta hyp 3 serrate and long, L = 32���43; seta hyp 2 simple and shorter, L = 11���12. Anterior margin of trochanter palp with blunt process (b) and spine-like process (s) (Fig. 8 F). Diagnosis. The sternal shield in both sexes is reticulated, all sternal setae on the female are long and serrated, and the sternogynal shield extends posteriorly to the mesogynal shield. Males have sternal setae s t 1, st 2, and st 3 long and serrated, and seta st 4 medium and serrated. Remarks. Klinckowstroemia oconnori sp. nov. is similar to K. montanoi Villegas-Guzman et al. because in both sexes the sternal shield is reticulated, but K. montanoi has sternal setae simple and short, in both sexes, whereas K. oconnori has sternal setae long and serrated. K. oconnori is similar to K. ballesterosi because both species have sternal setae long and serrated, but the sternal shield in K. ballesterosi is shagreened, whereas on K. oconnori it is reticulated, and K. oconnori (L= 781���806, A= 563���576) is smaller than K. ballesterosi (L= 896���1088, A= 640���774). Etymology. This species is named name in honour of Barry M. OConnor who collected the passalid hosts of this species and gave us the mites. Material examined. From Odontotaenius striatopunctatus: holotype ��, and 2 33 paratypes from Panama, Panama, Cerro Azul 950 m, 6 May 1991,B. M. OConnor; mites were found in the alcohol. From Petrejoides subrecticornis 2 �� and 2 33 paratypes from Panama, Panama, Cerro Azul 950 m, 6 May 1991, B. M. OConnor; mites were found in the alcohol. Holotype female, and 3 paratypes males deposited in NMNH, one female (CNAC 006746) and a male (CNAC 006747) paratypes are deposited in CNAC. One female was lost when it was being prepared for scanning electron micrographs, but this specimen was considered for description. Ecology. Specimens were found on two species of passalid beetles. The beetles were probably found in the same log, because they were collected at the same locality and on the same day; perhaps, for this reason they share the same species of mites, Published as part of Villegas-Guzm��n, Gabriel A., Reyes-Castillo, Pedro & P��rez, Tila M., 2011, New species of Klinckowstroemia Baker & Wharton, 1952 (Acari: Klinckowstroemiidae), associated with passalid beetles from Mexico and Panama, pp. 19-36 in Zootaxa 2747 on pages 31-35, DOI: 10.5281/zenodo.207115

Klinckowstroemia oconnori sp. nov. Female (n = 3) Body oval. Idiosoma: L = 781 (781–806); W = 576 (563–576) (Fig. 7 A). Gnathosoma with different hypostomal setae: hyp 1 long and simple, L= 50 (47–54); seta hyp 2 long and serrated, L = 43 (40–43), seta hyp 3 medium and serrated, L = 16 (12–16), and seta hyp 4 medium-sized and serrated, L = 16 (9–16). Chelicera with excrescences, movable digit with four teeth and fixed digit with seven teeth. Palp setae are simple, but av 1 and av 2 on trochanter are branched and serrated, respectively; on femur pd 1, pd 2, and pv 1 are serrated. On genu the setae al 1, al 2, ad 1, pl 1, pd 1 and pv 1 are serrated. Hyaline hood extending to level of coxa I, seta a 1 moderate and simple, L = 19 (19–22). Tritosternum base wider than long. Presternal (Pst) L = 37 (34–37), anterior W= 105 (102–109) shagreened and with line near the posterior margin; near posterior margin is a lyriform sternal pore stp 1. Presternal notch is a small concave depression. Sternal seta st 1, L = 53 (34–53), serrated (Fig. 7 A). Sternal shield L = 59 (56–62), shagreened, with some reticulations (Fig. 7 A). Lyrifissure stp 2 near to anterior margin of the shield; st 2, L = 47, serrated, below. Sternal setae st 3, L = 37, and st 4, L = 40, both setae are large and serrated, and near to posterior margin of the shield. Intact sternal setae present in only one specimen. Sternogynal shield (SgS), L = 68 (62–68); W = 158 (140–158) triangular and shagreened, posterior margin straight and slightly pointed, near to anterior margin is pore stp 3. Sternogynal apodeme thin and short. Latigynal shields shagreened, L = 130 (127–130); W = 65 (62–65), each with an oval pore and four simple setae, medial margin of shield straight. Mesogynal shield, L = 62 (59–62); W = 83. 7, triangular and shagreened, posterior margin wider and slightly pointed at apex. Mesogynal condyles extending almost to level of shield apex. Ventral shield, L = 171 (171–189); posterior W = 419 (403–419), shield triangular and shagreened, truncated posteriorly, with ten pairs of small setae (Fig. 7 A), with two pairs of pores at the level of posterior margin of coxa IV. The metapodal-peritrematal-exopodal shield (MPES) shagreened, with one pair of pores and one simple seta. Anal shield, L = 140 (124–140); anterior W = 341 (316–341), with 4 pairs of simple setae plus longer anteroanal seta, L = 22 (22–25); two pairs of pores, one near the anterior margin, and the other below the anus (Fig. 7 A). Serrate setae on leg I are: trochanter ad 1, al 1 and pv 1; femur: av 3, pv 1 and pv 2; tibia ad 1, ad 2, ad 3, av 1, av 2, pl 1, and pl 2. Serrate setae on Leg II are: femur pv 1; genu pv 1. Serrate setae on leg III: trochanter ad 1 and ad 2; femur av 1; genu av 1, and pv 1; tibia ad 4; tarsus ad 3, and pl 1. Serrate setae on leg IV: femur av 1; genu av 1, and pv 1; tibia ad 4, and al 4; tarsus ad 2 and ad 3 serrated. Seta pv 1 of the femur IV large L= 28 (27–28). Male (n = 5) Body oval, similar to female. Idiosoma L = 742–806, W = 493–563 (Fig. 7 B). Presternal (Pst) L = 31–37, anterior W = 102–109, presternal notch V-shaped, shagreened and with one line near anterior margin (Fig. 8 B) as in female; near to posterior margin is a lyriform sternal pore stp 1. Sternal seta st 1, L = 40–56, long and serrated (Fig. 8 C). Sternal shield shagreened (Fig. 8 D) and reticulated (Fig. 8 E), L = 121–127, with two pores, one near the anterior margin (stp 2) and other near the genital opening (stp 3). Sternal seta st 2 long and serrated, L = 48–50; sternal seta st 3 long and serrated, L = 39–47; sternal seta st 4 medium and serrated, L = 20–33. Genital opening oval, L = 50–62, W = 68–71. Ventral shield shagreened and reticulated, L = 264–285, W= 372–450 (Fig. 8 H), with 30 setae and two pores, one above the anterior margin of coxa IV, and other below coxa IV. Anal shield L = 102–140, anterior W = 322–363 (Fig. 8 G); with four pairs of simple setae plus longer antero-anal seta, L = 12–25, with two pores: one near the anterior margin and another below the anus. Hypostomal setae (Fig. 8 A): hyp 1 long and simple, L = 43–56; seta hyp 3 serrate and long, L = 32–43; seta hyp 2 simple and shorter, L = 11–12. Anterior margin of trochanter palp with blunt process (b) and spine-like process (s) (Fig. 8 F). Diagnosis. The sternal shield in both sexes is reticulated, all sternal setae on the female are long and serrated, and the sternogynal shield extends posteriorly to the mesogynal shield. Males have sternal setae s t 1, st 2, and st 3 long and serrated, and seta st 4 medium and serrated. Remarks. Klinckowstroemia oconnori sp. nov. is similar to K. montanoi Villegas-Guzman et al. because in both sexes the sternal shield is reticulated, but K. montanoi has sternal setae simple and short, in both sexes, whereas K. oconnori has sternal setae long and serrated. K. oconnori is similar to K. ballesterosi because both species have sternal setae long and serrated, but the sternal shield in K. ballesterosi is shagreened, whereas on K. oconnori it is reticulated, and K. oconnori (L= 781–806, A= 563–576) is smaller than K. ballesterosi (L= 896–1088, A= 640–774). Etymology. This species is named name in honour of Barry M. OConnor who collected the passalid hosts of this species and gave us the mites. Material examined. From Odontotaenius striatopunctatus: holotype Ƥ, and 2 33 paratypes from Panama, Panama, Cerro Azul 950 m, 6 May 1991,B. M. OConnor; mites were found in the alcohol. From Petrejoides subrecticornis 2 Ƥ and 2 33 paratypes from Panama, Panama, Cerro Azul 950 m, 6 May 1991, B. M. OConnor; mites were found in the alcohol. Holotype female, and 3 paratypes males deposited in NMNH, one female (CNAC 006746) and a male (CNAC 006747) paratypes are deposited in CNAC. One female was lost when it was being prepared for scanning electron micrographs, but this specimen was considered for description. Ecology. Specimens were found on two species of passalid beetles. The beetles were probably found in the same log, because they were collected at the same locality and on the same day; perhaps, for this reason they share the same species of mites

Klinckowstroemia brevimarginata sp. nov. Material examined. Holotype. �� (CNAC 006585), Oaxaca, Mexico, 101 km Federal highway Puerto Escondido-Oaxaca (16 �� 28.156 ' N, 97 ��01.674' W), 1,906 m, 28 June 2006, from Ve r re s aff. intermedius, O. Francke, A. Valdez and H. Monta��o coll., oak forest. Paratypes. 5 �� (CNAC 006586- CNAC 006590), 6 33 (CNAC 006591- CNAC 006596), same data as holotype, from two specimens of Ve r re s aff. intermedius. Holotype female, four paratype female and five paratype males deposited in the CNAC, one female and male paratypes deposited in NMNH. Female (n = 6). Body oval. Idiosoma, L = 762 (736���762), W = 525 (512���532) (Fig. 10 A). Ve n tr al idiosoma. Hyaline hood extending to level of coxa I, seta a 1 moderately long and simple, L = 19 (17���20). Tritosternum base wider than long. Tetartosternum shield L = 31 (28���31), anterior W = 99 (93���99), shagreened, anterior margin rounded, tetartosternal notch like a small cavity. Sternal seta st 1 near anterior margin, moderately long and serrate, L = 19 (16���22), lyriform sternal pore stp 1 near posterior margin. Sternal shield L = 62 (62���65), shagreened (Fig. 11 E). Pore stp 2 near centre of shield, behind it st 2, st 3 and st 4 are simple microsetae, the last two setae located on sides of shield near the corners, close to posterior margin. Sternogynial shield, L = 47 (47���53), W = 143 (140���152) triangular, narrow because mesogynial shield extends anteriorly (Fig. 11 D); posterior margin slightly pointed at apex, surface shagreened, stp 3 near anterior margin. Sternogynial apodeme long and thin, extending to posterior to level of latigynial and mesogynial condyles. Latigynial shields, L = 109 (109���124), W = 71 (68��� 71), each with an oval pore and four to six setae, commonly with five (Fig. 11 D), medial margins of shield short and straight, surface of shield shagreened; latigynial apodeme thin. Mesogynial shield L = 84 (82���93), W = 99 (93���99), triangular, wider posteriorly, slightly pointed at apex, surface shagreened. Mesogynial condyles behind level of apex of shield. Ventral shield, L = 198 (176���198), posterior W= 403 (403���429), triangular (Fig. 11 G), truncated posteriorly, surface shagreened and reticulated laterally, with 13 pairs of small setae. One pore underneath posterior margin of coxa IV. Metapodal-peritremal-exopodal shield with a pair of pores, and a seta, surface reticulated. Anal shield, L = 109 (102���124), anterior W = 288 (288���307), with five pairs of simple setae plus longer antero anal seta (Fig. 13 A), L = 24 (22���25), two pairs of pores, one near anterior margin and the other below the anus. Gnathosoma. Capitular setae (sc) short and serrate, L = 14 (9.3���14). Hypostomal setae hyp 1 simple, L = 50 (50���56), longest. Seta hyp 2 medium length, serrate, L = 31 (31���34); hyp 3 short and simple, L = 12 (12���14) (Fig. 11 C). Chelicera with barbed seta, movable digit bifurcate at tip, with four teeth, fixed digit with six teeth. Most palpal setae simple, except al 1 and al 2 on trochanter are branched and serrate respectively; femoral setae av 1, ad 2, pl 1, pd 1 and pd 2 barbed; genu setae pl 1, pl 2, pd 1, pd 2 and pv 1 barbed. Anterior margin of palp trochanter with blunt process (b), and denticulated process divided (s) (Fig. 11 F). Legs. Most leg setae simple, except: Leg I: trochanter I ad 1, al 1, pl 1 and pv 1 barbed; femur I: pv 1, pv 2 and pv 3 barbed; genu I: av 1 and pv 1 barbed; tibia I: ad 1, ad 2, ad 3, pd 1 and pd 3 barbed. Leg II: femur II: av 1 and pv 1 barbed; genu II: av 1 and pv 1 barbed. Leg III: trochanter III: pv 1 and pl 1 barbed; femur III: av 1 barbed; genu III: av 1 and pv 1 barbed. Leg IV: trochanter IV: pv 1; femur IV: av 1 barbed; genu IV av 1 barbed. Seta pv 1 is large and serrated (Fig. 11 H), L = 32 (31���38). Male (n = 6) Body oval. Idiosoma, L = 736���781, W = 499���544 (Fig. 10 B). Ve n tr al idiosoma. Tetartosternum, L = 31���40, anterior W = 93���102, shagreened, anterior margin round (Fig. 13 B). Tetartosternal notch distinct, anterior margin divided in two parts, with a pore near posterior margin. Sternal seta st 1, L = 9���16, serrate. Sternal shield shagreened, L = 93���102, reticulated laterally (Fig. 13 C). Setae, st 2, st 3 and st 4 are simple microsetae; with two pores, one near anterior margin of shield, one close to genital opening. Genital opening oval, L = 47���53, W = 78���81. Ventral shield, L = 282���307, posterior W = 448, shagreened, with 38 setae and two pores, one at level of anterior margin of coxa IV and the other below coxa IV; this area of shield reticulated laterally (Fig. 10 B). Anal shield shagreened, L = 109���124, anterior W = 333���358, with five pairs of simple setae plus longer antero anal seta, L = 20���25, with two pores, one near anterior margin and the other below anus. Metapodal-peritremal-exopodal shield with a pair of pores, and a seta, reticulated. Gnathosoma. Hypostomal seta hyp 1 long and simple, L = 47���53, hyp 2 serrate and long, L = 31���40, hyp 3 simple and shorter, L = 12. Etymology. This name is from Latin brevis (= small) and marginata (= margin) referring to the size of the medial margins of the latigynial shields. Remarks. This species can be distinguished from all others because the mesogynial shield extends to the sternogynial shield and is wide on posterior margin; the margin of the latigynial shield is straight and reduced, 12 (9���16). Sternal seta st 1 is of moderate length (16���22) and serrate in both sexes and sternal setae st 2, st 3 and st 4 are microsetae and simple. The sternal, genital, ventral and tetartosternum shields are shagreened and the ventral shield is reticulated laterally. The male tetartosternal shield has a round anterior margin, the tetartosternal notch dividing the anterior margin in two, and 38 pairs of setae on the ventral shield. Klinckowstroemia brevimarginata is similar to K. rectimarginata because they have st 1 serrate and st 2 is a microseta; however, K. rectimarginata (813���902) is bigger than K. brevimarginata (736���762) and has three setae on the latigynial shield, while K. brevimarginata has four to six setae. Klinckowstroemia brevimarginata is similar to K. schusteri because it has four setae on the latigynial shield and st 1 serrated, but the former is shorter than the latter and setae st 2, st 3 and st 4 are microsetae, while in K. shusteri setae st 2, st 3 and st 4 are short, the mesogynial shield does not extend anteriorly, and the medial margin of the latigynial shields is long. The specimens were found in decaying trunks at the same locality at Oaxaca, Mexico. The mites were found in the alcohol with two passalids., Published as part of Villegas-Guzman, Gabriel A., P��rez, Tila M. & Reyes-Castillo, Pedro, 2009, New species of the genus Klinckowstroemia Baker & Wharton from Mexico (Acari: Mesostigmata: Trigynaspida: Klinckowstroemiidae), pp. 1-46 in Zootaxa 2248 on pages 19-25, DOI: 10.5281/zenodo.190717

Klinckowstroemiidae Camin & Gorirossi Klinckowstroemiidae Camin & Gorirossi, 1955: 51. Along with their sister-group, the Fedrizziidae, the Klinckowstroemiidae are placed in the Fedrizzioidea (Camin & Gorirossi 1955). Kethley (1977) included the Paramegistidae and Promegistidae in this superfamily, but they were later moved to the Paramegistoidea by Kim (2004) and the original concept of the Fedrizzioidea retained. The Fedrizziidae and Klinckowstroemiidae are very similar and can only be distinguished by the form of the genital shields (Seeman 2007). The distribution is different for each family, since Fedrizziidae are found in Australia, Indonesia, Malaysia, Thailand, Papua New Guinea and Philippines (Seeman 2007), while Klinckowstroemiidae haves been recorded only in the American Continent. Both families have the same hosts, passalid beetles. Diagnosis (modified from Rosario & Hunter 1987) Dorsal shield convex, reticulate, bearing minute setae with no apparent pattern. Palpal trochanter with seta av 1 large, pectinate; seta av 2 pilose. Male jugular shields usually fused, forming tetartosternum, bearing sternal seta st 1 and pore stp 1; female always with tetartosternum. Sternal shield bearing st 2, st 3, st 4 and stp 2. Anal, ventral and metapodal-peritremal-exopodal shields contiguous or fused. Sternogynial, latigynial and mesogynial shields well developed, each with a vaginal apodeme on the internal surface; latigynial shields with setae, mesogynial and sternogynial shields usually without setae. Coxae marginal line present. Femora II���IV with ventral hyaline lobe bearing seta pv 1. Male genital opening oval, wider than long and between coxae II���III., Published as part of Villegas-Guzman, Gabriel A., P��rez, Tila M. & Reyes-Castillo, Pedro, 2009, New species of the genus Klinckowstroemia Baker & Wharton from Mexico (Acari: Mesostigmata: Trigynaspida: Klinckowstroemiidae), pp. 1-46 in Zootaxa 2248 on page 4, DOI: 10.5281/zenodo.190717, {"references":["Camin, J. H. & Gorirossi, F. E. (1955) A revision of the Suborder Meostigmata (Acarina), based on new interpretations of comparative morphological data. Chicago Academy of Sciences Special Publication, 11, 1 - 70.","Kethley, J. B. (1977) A review of the higher categories of Trigynaspida (Acari: Parasitiformes). International Journal of Acarology, 3, 129 - 149.","Kim, C. - M. (2004) Trigynaspida (Acari: Mesostigamta): new diagnosis, classification and phylogeny. Acarologia, 44, 157 - 197.","Seeman, O. D. (2007) Revision of the Fedrizziidae (Acari: Mesostigmata: Fedrizzioidea). Zootaxa, 1480, 1 - 55.","Rosario, R. M. T. & Hunter, P. E. (1987) The family Klinckowstroemiidae Tragardh with descriptions of two new species of Klinckowstroemiella (Acarina: Mesostigmata: Trigynaspida). Acarologia, 28, 307 - 317."]}

Klinckowstroemia franckei sp. nov. Material examined. Holotype. �� (CNAC 006603), Hidalgo, Mexico, Ejido Atepixco, Municipality Zacualtipan (20 �� 35.466 ' N, 98 �� 36.358 ' W), 1,972 m, 22 September 2006, from Odontotaenius zodiacus, pineoak forest, O. F. Francke coll.. Paratypes. 6 �� (CNAC 006604- CNAC 006609), 10 33 (CNAC 006610- CNAC 006619), same data as holotype, from nine specimens of Odontotaenius zodiacus. Holotype, three paratype females and seven paratype males deposited in the CNAC, one female and female paratype each deposited in NMNH, UGA and UMMZ. Female (n = 7). Body oval. Idiosoma, L = 909 (896���941); W = 627 (627���666) (Fig. 14 A). Ve n tr al idiosoma. Hyaline hood extending to level of coxa I, seta a 1 long and slightly barbed, L = 47 (47��� 53). Tritosternum base wider than long. Tetartosternum shield, L = 37, anterior width 124 (121���124); shagreened and reticulated (Fig. 15 E), anterior margin slightly wavy with a small square notch. Simple microseta st 1 near posterior margin of shield (Fig. 17 D), lyriform pore stp 1 near middle of shield, closer to posterior margin. Sternal shield, L = 90 (84���90), shagreened and reticulated (Fig. 17 A). Pore stp 2 near middle of shield, behind it sternal seta st 2, short and simple, L = 7, only seen in one specimen (Fig. 17 A). Setae st 3 is medium length and simple, L = 20 (12���22), st 4 simple and short, L = 9 (6���9) (Fig. 17 A), st 3 and st 4 near corners of shield, close to posterior margin. Sternogynial shield, L = 68 (65���71), W = 171 (140���171) triangular, posterior margin slightly pointed at apex, surface shagreened and reticulated (15 F), stp 3 near anterior margin (Fig. 15 F). Sternogynial apodeme long and thick, extending posteriorly to level of latigynial and mesogynial condyles. Latigynial shields, L = 130, W = 65 (65���71) each with an oval pore and four setae, medial margins of shields pointed, shield shagreened and reticulated; latigynial apodeme thin and long. Mesogynial shield, L = 78 (71.3���78), W = 74 (68���84), triangular, wider posteriorly and pointed at apex, extending slightly anterior to sternogynal shield, surface shagreened, mesogynial condyles under level of shield apex. Ventral shield, L = 242 (242���260); posterior W = 480 (467���512), triangular, truncated posteriorly, surface shagreened (Fig. 17 C), with 12 pairs of small setae (Fig. 14 A). One pore underneath posterior margin of coxa IV. Metapodal-peritremal-exopodal shield (MPES) reticulated, with a pair of pores (Fig. 17 C) and a seta. Anal shield, L = 149 (149���164); anterior W = 416 (390���429), with five pairs of simple setae plus longer antero anal seta, aa, L = 31 (29���32), two pairs of pores, one near anterior margin, another below the anus. Gnathosoma. Capitular setae (sc) short and serrate, L = 14 (12���16). Hypostomal setae hyp 1 longest, simple, L = 65 (62���65); hyp 2 medium length and barbed, L = 37 (36���40); hyp 3 short and slightly serrated, L = 15 (14���16) (Fig. 15 G). Chelicera with barbed seta, movable digit with four teeth, fixed digit with six teeth. Most palpal setae simple, except al 1 and al 2 on trochanter are branched and serrate respectively. Femur setae av 1 and pd 1 barbed, ad 1 serrated. Anterior margin of palp trochanter with blunt process (b) and denticulated process (s) (Fig. 15 H). Legs. Leg setae mostly simple, except: Leg I: trochanter ad 1 is barbed, pv 1 serrated; femur pv 1, pv 2, pv 3 and av 1 barbed; genu av 1 barbed, pv 1 serrated. Leg II, femur av 1 barbed; genu av 1, pv 1 barbed. Leg III: femur av 1 barbed; genu pv 1, av 1 slightly barbed. Leg IV: trochanter pv 1, pl 1 slightly serrated; femur av 1 barbed; genu av 1, pv 1 slightly barbed; tibia al 3 barbed; tarsus al 2 serrated. Seta pv 1 is large and slightly serrated, L = 29 (28���29) (Fig. 17 B). Male (n = 10) Body oval, similar to female. Idiosoma, L = 896���960, W = 621���659 (Fig. 14 B). Ve n tr al idiosoma. Tetartosternum, L = 34���47, anterior W = 118���124, shagreened, anterior margin round. Tetartosternal notch small, dividing anterior shield margin in two parts, stp 1 near posterior margin. Sternal shield shagreened and reticulated, L = 148 (143���152). Sternal seta st 1 simple, L = 16���19, st 2, st 3 and st 4, are microsetae and simple; shield with two pores, stp 2 near anterior margin, stp 3 close to genital opening. Genital opening oval, L = 56���62, W = 78���87. Ventral shield, L = 326���357, posterior W = 486���550, shagreened, reticulated only near anterior margin, with 26���32 setae and two pores, one at level of anterior margin of coxa IV, the other below coxa IV (Fig. 14 B). Anal shield shagreened, L = 140���161, anterior W = 397���429, with six pairs of simple setae plus longer antero anal seta, L = 28���37, with two pores, one near anterior margin and one below anus. Metapodal-peritremal-exopodal shield reticulated with a pair of pores, with seta. Gnathosoma. Hypostomal seta hyp 1 long and simple, L = 62���68, hyp 2 barbed and intermediate, L = 37��� 43, hyp 3 simple and short, L = 12���14. Etymology. This species is named in honour of Oscar F. Francke who collected the many of the passalid beetles that carried Klinckowstroemiidae. Remarks. This species can be distinguished from all others because setae st 1 and st 2 are microsetae, and st 3 is the longest of the sternal setae. The tetartosternal, sternal, mesogynial and latigynial shields are shagreened and reticulated. the medial margins of the latigynial shield are pointed, and there are four small simple setae on this shield. The mesogynial shield has a broad posterior margin and its apex is anteriorly pointed. The male has st 1 medium sized and simple, with a small tetartosternal notch that divides the anterior margin in two parts; the tetartosternum and sternal shield are reticulated and shagreened. The other sternal setae are microsetae. The ventral shield has 26���32 setae, but many of the specimenss have 28. Klinckowstroemia franckei is similar to K. multisetillosa because they have similar size (K. franckei L= 908,W= 627; K. multisetillosa L= 936, W= 634), st 1 is simple, and the medial margin of the latigynial shield is pointed. However, the new species has only one pair of seta a, while K. multisetillosa has six pairs. Furthermore, the latigynial shield of K. franckei has four setae and is larger than that of K. multisetillosa, which has five setae. The posterior margin of the mesogynial shield is broader and more convex in K. multisetillosa than in K. franckei. The capitular seta is also small and simple in K. multisetillosa, but in K. franckei it is twice as long and serrated. The beetles carrying these mites were found in decaying trunks at the same locality; it is possible that some specimens occupied the same trunk as a family group. Almost all mites were found in the alcohol in the vials where the beetles were collected. Three mites were found on the body: one on the mesoepisternum, one on the mesosternellum and another near to coxa III. Each mite was found on a different passalid., Published as part of Villegas-Guzman, Gabriel A., P��rez, Tila M. & Reyes-Castillo, Pedro, 2009, New species of the genus Klinckowstroemia Baker & Wharton from Mexico (Acari: Mesostigmata: Trigynaspida: Klinckowstroemiidae), pp. 1-46 in Zootaxa 2248 on pages 26-32, DOI: 10.5281/zenodo.190717

Klinckowstroemia bifurcata sp. nov. Material examined. Holotype. �� (CNAC 006514), Hidalgo, Mexico, 2 km Federal Highway Crucero- Tianguistengo (20 �� 40.413 ' N, 98 �� 40.255 ' W), 2,080 m, 23 September 2006, pine-oak forest, from Odontotaenius zodiacus (Truqui), O. Francke, A. Valdez and H. Monta��o coll., mites found in the alcohol. Paratypes. 4 �� (CNAC 006515- CNAC 006518), 1 3 (CNAC 006519), same data as holotype, from 3 specimens of Odontotaenius zodiacus; 8 �� (CNAC 006520- CNAC 006527), 2 33 (CNAC 006528, CNAC 006529), Hidalgo, Mexico, 2 km Mojonera-Tizapan, municipality Zacualtipan (20 �� 38.107 ' N, 98 �� 35.981 ' W), 2,019 m, 22 September 2006, from 4 specimens of Odontotaenius zodiacus, O. Francke, A. Valdez and H. Monta��o coll. Mites were on coxa I, mesepisternum and loose in the alcohol. Holotype, 9 paratype females and 3 paratype males deposited in the CNAC, one female paratype each deposited in NMNH, UGA and UMMZ. Female (n = 13). Body oval. Idiosoma: L = 966 (922���1,088), W = 723 (659���762) (Fig. 1 A). Ve n tr al idiosoma. Hyaline hood extending to level of coxa I, seta a 1 large and simple (Fig. 2 G), L = 34 (30���37). Tritosternum base wider than long. Tetartosternum (Tst) L = 40 (40���44), anterior W= 124 (112���127) (Fig. 2 E); near the posterior margin is a lyriform sternal pore stp 1. Tetartosternal notch square, shagreened and with a line near the anterior margin. Sternal seta st 1, L = 18 (16���22), bifurcated (Fig. 2 C). Sternal shield (Fig. 2 B) L = 90 (78���91), stp 2 in centre of shield, below it st 2, L = 11 (9���19), simple. Setae st 3, L = 31 (19��� 40) and st 4, L = 16 (9���22) near posterior margin of shield, both large and simple, close together (Fig. 2 B). Shield shagreened, without reticulations. Sternogynial shield, L = 68 (62���96); W = 192 (167���201) triangular (Fig. 2 F), shagreened, with stp 3 closer to anterior margin. Sternogynial apodeme short. Latigynial shields, L = 130 (124���146), W = 76 (75���84) each with an oval pore and four setae (Fig. 2 F), medial margin of shield straight, extremity blunt. Mesogynial shield, L = 76 (68���84), W = 106 (96���112), triangular, wider anteriorly and slightly pointed at apex, surface shagreened. Mesogynial condyles extending almost to level of shield apex. Ventral shield, L = 269 (248���288), posterior W = 576 (518���608), shield triangular, truncated posteriorly, surface shagreened, with 12 pairs of small setae (Fig. 1 A). One pore at level of posterior margin of coxa IV. Anal shield, L = 192 (167���218); anterior W = 518 (467���576), with six pairs of simple setae plus longer antero anal seta, L = 25 (17���31), two pairs of pores, one near the anterior margin, the other below the anus. Gnathosoma. Capitular seta (sc) short and serrate, 12 (9���16). Hypostomal seta hyp 1 long and simple, 62 (56���71). Seta hyp 2 long and serrate, L = 40 (34���47). Seta hyp 3 short and simple, L = 7 (7���19) (Fig. 2 A). Chelicera with barbed seta, movable digit with four teeth, fixed digit with five teeth. Palp setae are simple, av 1 and av 2 on trochanter branched and serrate respectively; dorsal setae of femur pl 1 and pl 2 serrate and feathery respectively (Fig. 2 A). Anterior margin of palp trochanter with blunt process (b) (Fig. 2 H). Legs. Most setae on leg I serrate, on trochanter ad 1, al 1 and pv 1, genu al 1 and av 1. Serrate setae on leg II: femur av 1 and genu av 1. Serrate setae on leg III: femur av 1, genu pv 1, tibia ad 3 and al 3; on leg IV, genu pv 1, tibia ad 3 and al 3. Other setae simple, except pv 1 on trochanter II���IV lightly barbed (Fig. 2 D). Male (n = 3). Body oval, similar to female. Idiosoma L = 986���1024, W = 691���736 (Fig. 1 B). Ve n tr al idiosoma. Tetartosternum L = 50���53, posterior W = 102���112, tetartosternal notch shallow (Fig. 4 A), shagreened, with one line near anterior margin as in female. Sternal seta st 1, L = 19���24, simple. The other sternal setae are microsetae. Sternal shield shagreened, L = 152���155 (Fig. 4 B). Genital opening oval, L = 59���66, W = 87���94. Ventral shield with 32 setae (Fig. 1 B), L = 363���378, posterior W = 608, shagreened. Anal shield L = 179���217, anterior W = 512���544; with six pairs of simple setae plus longer antero anal seta, L = 20���30. Gnathosoma. Hypostomal seta hyp 1 is long and simple, L = 65���71; seta hyp 2 is serrate and long, L = 43��� 50, while seta hyp 3 is simple and shorter, L = 16���17. Etymology. This name is from Latin bifurcata (= bifurcate) referring to the shape of st 1. Remarks. This species can be distinguished from all others because sternal seta st 1 is bifurcate; and the tetartosternum, sternal and genital shields are shagreened. Sternal setae st 3 and st 4 are slightly serrated, seta st 3 longest. The mesogynial shield is wider on the posterior margin and the latigynial shield has four small simple setae and a pore. Some setae on the palp and leg I are serrated and feathery. The male has sternal seta st 1 simple and the tetartosternum is similar to that of the female. Klinckowstroemia bifurcata is similar to K. reyesi and K. truncata because these three species have the same number of setae on the latigynial, ventral and anal shields. Klinckowstroemia bifurcata (981) is slightly bigger than K. truncata (820) but is slightly smaller than K. reyesi (1009). Klinckowstroemia bifurcata has four setae on the latigynial shield and sternal seta 3 and 4 are serrate as in K. truncata. Sternal seta 3 is longest as in K. reyesi. Klinckowswtroemia bifurcata has the medial margin of the latigynial shield straight and its extremity blunt, while that of K. reyesi and K. truncata is longer and completely straight. The specimens were found in decaying trunks at two different localities in the state of Hidalgo, Mexico, but on the same host (Odontotaenius zodiacus). In these localities we collected three and four passalids carrying K. bifurcata. The mites were usually found in the alcohol, only in one case they were located on the passalid���s body, on coxa I and mesoepisternum. Two or more mite species were found on one beetle: one male of K. victoriae, seven (4 ��, 3 33) of K. bifurcata; and another passalid was found with three species, K. bifurcata; K. hunteri and K. valdezi. This association is known as synhospitality (Eichler 1966)., Published as part of Villegas-Guzman, Gabriel A., P��rez, Tila M. & Reyes-Castillo, Pedro, 2009, New species of the genus Klinckowstroemia Baker & Wharton from Mexico (Acari: Mesostigmata: Trigynaspida: Klinckowstroemiidae), pp. 1-46 in Zootaxa 2248 on pages 7-8, DOI: 10.5281/zenodo.190717, {"references":["Eichler, W. (1966) Two new evolutionary terms for speciation in parasitic animals. Systematic Zoology, 15, 216 - 218."]}

Klinckowstroemia montanoi sp. nov. Material examined. Holotype. �� (CNAC 006538), Oaxaca, Mexico, 8 km NW San Miguel Coatl��n, Federal highway Cuixtla-San Miguel Coatl��n (16 �� 15.064 ' N, 96 �� 41.629 ' W), 2,170 m, 24 June 2006, from Verres aff. intermedius, O. Francke, A. Valdez and H. Monta��o coll., oak forest. Paratypes. 14 �� (CNAC 006539- CNAC 006552), 5 33 (CNAC 006553- CNAC 006557), same data as holotype, from four specimens of Verres aff. intermedius. Holotype, 10 paratype female and four paratype males deposited in the CNAC, one female and one male paratype deposited in NMNH; one female paratype each deposited in UGA and UMMZ. Female (n = 15). Body oval. Idiosoma, L = 742 (704���755), W = 512 (493���538) (Fig. 5 A). Ve n tr al idiosoma. Hyaline hood extending to level of coxa I, seta a 1 moderate and simple, L = 19 (16���18). Tritosternum base wider than long. Tetartosternum (Tst) L = 43 (40���47), anterior W = 93 (87���93), shagreened (Fig. 6 G), with a line near anterior margin, lyriform sternal pore stp 1 near posterior margin. Tetartosternal notch square-ish, sternal seta st 1, L = 22 (21���29), serrate (Fig. 8 F). Sternal shield (Fig. 6 H) L = 50 (43���56), stp 2 near centre of shield, behind it sternal seta st 2, short and simple, L = 6. Setae st 3, L = 6, simple, and st 4, a microseta, near posterior margin of shield. Shield reticulated and shagreened (Fig. 6 H). Sternogynial shield, L = 43 (37���47), W = 152 (143���155), triangular, posterior margin narrow and pointed at apex, surface shagreened and reticulated (Fig. 8 A), stp 3 near anterior margin. Sternogynial apodeme short and thick, extending to posterior level of latigynial and mesogynial condyles. Latigynial shields, L = 99 (96���102); W = 65 (65��� 68) each with an oval pore and five to eight setae, medial margins of shields wavy and short (Fig. 8 A), shields reticulated and surface shagreened; latigynial apodeme thin. Mesogynial shield, L = 62 (59���68); W = 74 (71���77), triangular, wider posteriorly and pointed at apex, surface shagreened. Mesogynial condyles in centre of shield. Ventral shield, L = 211 (195���220); posterior W = 422 (416���435), shield triangular (Fig. 8 D), truncated posteriorly, surface shagreened and reticulated, with 13���14 pairs of small setae (Fig. 5 A). One pore underneath posterior margin of coxa IV. Metapodal-peritremal-exopodal shield with a pair of pores and one simple seta. Anal shield (Fig. 8 E), L = 124 (105���140); anterior W = 352 (320���365), with five pairs of simple setae plus longer anteroanal seta, L = 25 (20���25), two pairs of pores, one near anterior margin and the other below anus. Gnathosoma. Capitular setae (sc) short and serrate, L = 11 (8���12). With three hypostomal setae (Fig. 6 F), hyp 1 long and simple, L = 46 (43���50), longest. Seta hyp 2 medium and serrate, L = 29 (26���32). Seta hyp 3 short and serrate, L = 15 (14���17). Chelicera with barbed seta, movable digit bifurcate at tip, with four teeth; fixed digit with five teeth. Palp setae generally simple, except al 1 and al 2 on trochanter branched and serrate respectively. Femur setae av 1, pl 1, pl 2, pl 3 and pd 1 serrate. Genu setae av 1, pl 1, pl 2, pl 3, pd 1 and pd 2 serrate. Anterior margin of palp trochanter with blunt process (b), denticulated process divided (s) (Fig. 8 B). Legs. Leg setae mostly simple except: Leg I: trochanter I ad 1 and pv 1 serrate; femur I ad 1, ad 2, pv 1 and pv 3 serrate; genu I av 1 and pv 1 serrate. Leg II: femu av 1 and genu av 1 serrate. Leg III: femur av 1; genu III av 1 and pv 1 serrate; trochanter IV pv 1, femur av 1; genu av 1 and pv 1 serrate. Seta pv 1 (Fig. 8 C) is large and serrate, L = 37 (34���38). Male (n = 5) Body oval, similar to female. Idiosoma, L = 704���755, W = 474���518 (Fig. 5 B). Ve n tr al idiosoma. Tetartosternum, L = 40���56, anterior W = 93���124, shagreened. Tetartosternal notch deep, V-shaped. Sternal seta st 1, L = 20���22, simple. Setae st 2, st 3 and st 4 are simple microsetae. Sternal shield shagreened and reticulated (Fig. 5 B), L = 78���81. Genital opening oval, L = 43���47, W = 71���78. Ventral shield, L = 279���285, posterior W = 384���403, shagreened, with 24���26 setae (Fig. 5 B), with two pores, one at level of coxa III and other below posterior margin of coxa IV. Anal shield, L = 124���127, anterior W = 333���352, with five pairs of simple setae plus longer antero anal seta, L = 24���25, with two pores, one near anterior margin and the other below anus. Gnathosoma. Hypostomal seta hyp 1 long and simple, L = 47���50, hyp 3 serrate, L = 31���34, hyp 2 simple and shorter, L = 16���25. Etymology. This species is named in honour of H��ctor Monta��o, who helped to collect almost all the passalid beetles used in this research. Remarks. This species can be distinguished from all others because st 4 is a microseta, st 2 and st 3 are short and simple, st 1 is moderate and serrate and the medial margin of the latigynial shield is wavy. The sternal, latigynial and sternogynial genital shields are shagreened and reticulated, the tetartosternum is only shagreened. The mesogynial shield is wider in the anterior and posterior margins. In the male the tetartosternal notch is deep and distinct, V-shaped, almost reaching half the length of the shield, and there are 24���26 pairs of setae on the ventral shield. Klinckowstemia montanoi is similar to K. scotti and K. valdezi because all the species have wavy medial margins on the latigynial shield, st 1 serrated, and number of setae on anal shield is equal. Klinckowstroemia montanoi (735.6) is longer than K. valdezi (685.9), but shorter than K. scotti (1292). The latigynial shield has eight pilose setae in K. scotti, while K. montanoi have five to eight simple setae simple and K. valdezi has three simple setae. K. montanoi has 13���14 pairs setae on the ventral shield, while K. valdezi and K. scotti have 12 pairs. The specimens were found in decaying trunks only in one locality in the state of Oaxaca, Mexico, on the same host, Ve r re s aff. intermedius. Mites were found in the alcohol, not on the beetles., Published as part of Villegas-Guzman, Gabriel A., P��rez, Tila M. & Reyes-Castillo, Pedro, 2009, New species of the genus Klinckowstroemia Baker & Wharton from Mexico (Acari: Mesostigmata: Trigynaspida: Klinckowstroemiidae), pp. 1-46 in Zootaxa 2248 on pages 11-15, DOI: 10.5281/zenodo.190717

Villegas-Guzman, Gabriel A., Pérez, Tila M., Reyes-Castillo, Pedro (2009): New species of the genus Klinckowstroemia Baker & Wharton from Mexico (Acari: Mesostigmata: Trigynaspida: Klinckowstroemiidae). Zootaxa 2248: 1-46, DOI: 10.5281/zenodo.190717

Klinckowstroemia santibanezi sp. nov. Material examined. Holotype. �� (CNAC 006620), Oaxaca, Mexico, Municipality of Ixtl��n de Ju��rez (17 �� 31.722 ' N, 96 �� 30.678 ' W), 2,742 m, 13 November 2005, pine forest, from Proculejus sartorii, O. Francke, M. Cordova, A. Jaimes and G. Montiel coll. Paratypes. 1 �� (CNAC 006621), 1 3 (CNAC 006622), same data as holotype, from one specimen of Proculejus sartorii; 7 �� (CNAC 006623- CNAC 006629), 4 33 (CNAC 006630- CNAC 006633), Oaxaca, Mexico, Cerro Pel��n, highway Tuxtepec-Oaxaca (17 �� 35.75 ' N, 96 �� 28.41 ' W), 2,240 m, 22 November 1996, evergreen cloud forest, from one specimen of Proculejus brevis, inside log of Persea americana, F. Lorea. coll.; 2 �� (CNAC 006634- CNAC 006635), Oaxaca, Mexico, Municipality of Ixtl��n de Ju��rez (17 �� 31.722 ' N, 96 �� 30.678 ' W), 2,742 m, 13 November 2005, pine forest, from one specimen of Vindex sp. nov., O. Francke, M. Cordova, A. Jaimes and G. Montiel coll.; 3 �� (CNAC 006636- CNAC 006638), 1 3 (CNAC 006639), Oaxaca, Mexico, 8 km SE San Miguel Talea de Castro, Municipality San Miguel Talea de Castro, (17 �� 19.620 ' N, 96 �� 17.403 ' W), 2,082 m, 22 July 2007, evergreen cloud forest, from three specimens of Proculejus brevis, O. Francke, A. Valdez, C. Santiba��ez, A. Ballesteros and H. Monta��o coll. Holotype, ten paratype females and three paratype males deposited in the CNAC, one female and male paratype each deposited in NMNH, UGA and UMMZ. Female (n = 14). Body oval. Idiosoma, L = 1229 (1165���1517); W = 909 (832���1107) (Fig. 16 A). Ve n tr al idiosoma. Hyaline hood extending beyond to level of coxa IV, seta a 1 long and barbed (Fig. 18 C), L = 93 (56���96). Tritosternum base wider than long. Tetartosternum shield, L = 47 (40���53), anterior W = 155 (133���155), shagreened (Fig. 17 F), anterior margin slightly rounded, corners of shield concave, medial notch small, square. Sternal seta st 1 long and barbed (Fig. 17 E), L = 47 (40���61), near centre of shield; lyriform sternal pore stp 1 close to posterior margin of shield. Sternal shield L = 109 (93���115), shagreened. Pore stp 2 near centre of shield, behind and below it st 2, moderately long and serrated, L = 26 (12���26). Setae st 3 moderately long, serrated, L = 19 (19���25), st 4 is serrated and short, L = 9 (9���16) (Fig. 18 B). Setae st 3 and st 4 on sides of shield, close to posterior margin. Sternogynial shield, L = 90 (87���102), W = 233 (217���236) triangular, posterior margin pointed at apex (Fig. 17 G), surface shagreened, stp 3 near anterior margin. Sternogynial apodeme short and thick, extending posteriorly to level of latigynial and mesogynial condyles. Latigynial shields, L = 186 (174���186), W= 93 (87���93) each with an oval pore and six to ten serrated setae, usually eight; medial margins of shields straight, surface shagreened, latigynial apodeme long and thin. Mesogynial shield, L = 74 (71���90), W = 121 (112���140), triangular, wider posteriorly and anteriorly (Fig. 17 G), slightly pointed at apex, surface shagreened, mesogynial condyles at level of shield apex. Ventral shield, L = 285 (264���304), posterior W = 627 (595���678), triangular, truncated posteriorly, surface shagreened and reticulated like a honey-comb, with 11���14 pairs of small setae (Fig. 16 A). One pore underneath posterior margin of coxa IV. Metapodal-peritremal-exopodal shield reticulated, with a pair of pores and three setae. Anal shield, L = 192 (192���242), anterior W = 576 (531���608) (Fig. 18 D), with five to seven pairs of simple setae plus longer antero anal seta, L = 27 (27���45), two pairs of pores, one near anterior margin and the other below the anus. Shield reticulated like ventral shield (Fig. 18 D). Gnathosoma. Capitular setae (sc) medium and barbed, L = 19 (19���25). Hypostomal setae (Fig. 18 A) hyp 1 long and simple, L = 71 (71���87), longest; hyp 2 long and barbed, 59 (57���62); hyp 3 moderately long and barbed, L = 20 (16���20). Chelicera with barbed seta, movable digit with four teeth, fixed digit with six teeth, movable finger divided at tip. Palp setae are simple, except al 1 and al 2 on trochanter are branched and serrate respectively. Femur setae pl 1 barbed, pl 2 and pd 1 serrated; genu seta al 1 barbed. Anterior margin of palp trochanter with a small blunt process (b) and denticulated process (s) (Fig. 18 E). Legs. Most leg setae simple, except: Leg I: trochanter ad 1, av 1, al 1 and pv 1 pilose; genu ad 1, av 1, pl 2 and pv 1 pilose; tibia ad 1, ad 2, ad 3, av 1, av 2, pd 1, pv 2 and pd 3 pilose. Leg II: trochanter ad 1, ad 2, pd 1 and pd 2 serrated; femur av 1, pv 1, pd 1, pd 2 and pd 3 serrated; genu av 1 and pv 1 serrated; tibia ad 3, al 3, pd 3 and pl 3 serrated; tarsus ad 1, ad 2, ad 3, pd 2, pd 3 and pl 2 serrated, av 2 and av 3 slightly serrated. Leg III: trochanter ad 1, ad 2, pd 1, pd 2 and pv 1 serrated; femur ad 1, ad 2, av 1, pd 1 and pl 1 serrated; genu av 1, pl 1 and pd 2 serrated; tibia pd 3 serrated; tarsus av 3 serrated. Leg IV: trochanter pv 1 and pl 1 serrated; femur av 1 serrated; genu av 1, pl 1 and pv 1 serrated; tibia pd 3 and pl 3 barbed. Seta pv 1 is serrated and long, 36 (31���47) (Fig. 17 H). Male (n = 6) Body oval, similar to female. Idiosoma, L = 1242���1448, W = 954���1037 (Fig. 16 B). Ve n tr al idiosoma. Tetartosternum, L = 59���62, anterior W = 149���161, shagreened and reticulated, anterior margin round and middle of shield is compressed (Fig. 18 F) Tetartosternal notch is small and divides anterior margin in two parts, with a pore near posterior margin. Sternal seta st 1, L = 17���25, simple (Fig. 18 F). Sternal shield (Fig. 18 G) shagreened and reticulated, L = 180���195. Sternal setae, st 2 simple, L = 14, st 3 simple, L= 9��� 12), st 4 is short and simple, L = 6, (these setae were found in only two specimens); with two pores, one close to anterior margin of shield and one close to genital opening. Genital opening oval, L = 81���87, W= 121���133 (Fig. 18 G). Ventral shield, L = 422���481, posterior W = 608���672, shagreened and reticulated like a honeycomb, with 34���36 setae; with two pores, one at level of anterior margin of coxa IV and other below level of coxa IV. Anal shield shagreened and reticulated, L = 205���264, anterior W = 544���589; with six pairs of simple setae plus longer antero anal seta, L = 42���48, with two pores, one near anterior margin and the other below anus. Metapodal-peritremal-exopodal shield reticulated with a pair of pores, without seta (Fig. 16 B). Gnathosoma. Hypostomal seta hyp 1 long and simple, L = 80���87; hyp 2 barbed and long, L = 50���68, hyp 3 barbed and medium length, L = 17���25. Etymology. This species is named in honour of Carlos Santiba��ez, who helped to collect almost all the passalid beetles used in this research. Remarks. This species can be distinguished from all others because the body is very large and setae st 1 is long and barbed, the longest of the sternal setae. In females the tetartosternal, sternal, mesogynial and latigynial shields are shagreened, and only the ventral and anal shield are reticulated. The latigynial shield has six to ten serrated setae. The mesogynial shield has broad posterior and anterior margins. The male has st 1 medium length and simple, and the tetartosternal, sternal, ventral and anal shields are reticulated and shagreened. The tetartosternal notch is small and divides the anterior margin into two parts. The ventral shield has 34���36 setae. Klinckowsroemia santibanezi is very similar is similar to K. candidoi, because they both have the hyaline hood beyond coxa IV, st 1 long and barbed, and the ventral and anal shields reticulated like honey-comb. But new the species has all the sternal setae barbed, with six to ten serrated setae on the latigynial shield, and st 3 and st 4 are serrated. In contrast, K. candidoi has three to four simple setae on the latigynial shield and sternal setae st 3 and st 4 are simple. The beetles carrying this species were found in decaying trunks in three different localities and hosts from Oaxaca. Two mites were found on the humerus and metasternum of the hosts, but almost all specimens were found in the alcohol. One passalid beetle had two species of mites, K. santibanezi and K. victoriae., Published as part of Villegas-Guzman, Gabriel A., P��rez, Tila M. & Reyes-Castillo, Pedro, 2009, New species of the genus Klinckowstroemia Baker & Wharton from Mexico (Acari: Mesostigmata: Trigynaspida: Klinckowstroemiidae), pp. 1-46 in Zootaxa 2248 on pages 32-33, DOI: 10.5281/zenodo.190717

Klinckowstroemia valdezi sp. nov. Material examined. Holotype. Ƥ (CNAC 006558), Veracruz, Mexico, 1.5 km N Viborillas, municipality Huayacocotla (20 ° 31.245 ' N, 98 ° 29.923 ' W), 2,328 m, 23 September 2006, from Proculejus hirtus (Truqui), O. Francke, A. Valdez and H. Montaño coll., pine-oak forest. Paratypes. 3 Ƥ (CNAC 0 0 6559 - CNAC 006561), 3 33 (CNAC 006562- CNAC 006564), same data as holotype, from three specimens of Proculejus hirtus; 2 Ƥ (CNAC 006565, CNAC 006566), 13 (CNAC 006567), Hidalgo, Mexico, 2 km Federal highway Crucero-Tianguistengo (20 ° 40.413 ’ N, 98 ° 40.255 ’ W), 2,080 m, 23 September 2006, pine-oak forest, from two specimens of Oileus nonstriatus (Dibb), O. Francke, A. Valdez and H. Montaño coll.; 1 Ƥ (CNAC 006568), 1 3 (CNAC 006569), Hidalgo, Mexico, 2 km Mojonera-Tizapan, municipality Zacualtipan (20 ° 38.107 ' N, 98 ° 35.981 ' W), 2,019 m, 22 September 2006, on one specimen of Odontotaenius zodiacus (Truqui), O. Francke, A. Valdez and H. Montaño coll., pine-oak forest; 7 Ƥ (CNAC 006570- CNAC 006576), 3 33 (CNAC 006577- CNAC 006579), Veracruz, Mexico, 11.9 km N Huayacocotla (20 ° 32.116 ’ N, 98 ° 28.783 ’ W), 11 October 1982, from Proculejus brevis, tropical rain forest, P. Reyes-Castillo coll.; 3 Ƥ (CNAC 006580- CNAC 006582), 2 33 (CNAC 006583, CNAC 006584), Veracruz, Mexico, 16.4 km N Huayacocotla (20 ° 32.116 ’ N, 98 ° 28.783 ’ W), 11 October 1982, from Proculejus brevis, tropical rain forest, P. Reyes-Castillo coll.; Holotype, 13 paratype females and 8 paratype males deposited in the CNAC, one female and male paratype each deposited in NMNH, UGA and UMMZ. Female (n = 17). Body oval. Idiosoma, L = 710 (653–710), W = 486 (448–499) (Fig. 7 A). Ve n tr al idiosoma. Hyaline hood extending to level of coxa I, seta a 1 moderate and simple, L = 16 (13–22). Tritosternum base wider than long. Tetartosternum (Tst) (Fig. 8 G) long 31 (28–31), anterior W = 84 (76–87), shagreened, with a line on each side; near posterior margin is a lyriform sternal pore stp 1. Tetartosternal notch square. Sternal seta st 1 very long and serrated, L = 39 (31–40) (Fig. 8 H). Sternal shield (Fig. 9 A), L = 59 (53– 62), shagreened. Near centre of shield stp 2, behind it seta st 2, this seta was seen in only three specimens, L = 12; in one of them one seta is small and other is very long and serrate. Sternal setae st 3 and st 4 are broken, located near posterior margin of shield (Fig. 9 A). Sternogynial shield L = 40 (31–40), W = 140 (127–146), triangular (Fig. 9 C), narrow because medial margins of latigynial shields are long, posterior margins slightly pointed at apex, surface shagreened, stp 3 near the anterior margin. Sternogynial apodeme short and thin, extending to posterior level of latigynial and mesogynial condyles. Latigynial shields, L = 99 (90–105), W = 62 (56 – 56) each with an oval pore and three setae (Fig. 9 C), medial margins of shields long and wavy, shield shagreened; latigynial apodeme thin. Mesogynial shield, L = 62 (47–62), W = 76 (62–77), triangular, wide posteriorly and slightly pointed at apex (Fig. 9 C), surface shagreened. Mesogynial condyles almost reach level of shield apex. Ventral shield, L = 192 (176–208); posterior W = 397 (352–397), shield triangular (Fig. 9 D), truncated posteriorly, surface shagreened, with 12 pairs of small setae (Fig. 7 A). One pore underneath posterior margin of coxa IV. Metapodal-peritremal-exopodal shield with a pair of pores, without setae, surface reticulated (Fig. 9 D). Anal shield long, 109 (93–115); anterior width, 320 (288–339), with five pairs of simple setae plus longer antero anal seta, L = 22 (19–24), two pairs pores, one near anterior margin and the other below the anus (Fig. 9 E). Gnathosoma. Capitular setae (sc) short and simple, L = 9 (6–14). With three hypostomal setae (Fig. 9 F), hyp 1 long and simple, L = 50 (47–53), longest. Seta hyp 2 medium length and finely serrated, L = 34 (25–34), hyp 3 short and simple, L = 11 (9–14). Chelicera with barbed seta, movable digit bifurcated at the tip, with three teeth; fixed digit with six teeth. Palpal setae simple, except al 1 and al 2 on trochanter branched and serrate respectively. Setae of ad 1 femur serrate (Fig. 9 H); genu setae av 1 and ad 1 serrate. Anterior margin of palp trochanter with blunt process long (b), denticulated process divided (s) (Fig. 9 G). Legs. Most leg setae simple, except: Leg I trochanter ad 1 and pl 1 serrate; femur pv 1 and pv 2 serrate; genu av 1, pl 1 and pv 1 serrate. Leg II: femur av 1 and pv 1 serrate; genu av 1 and pv 1 serrate; tibia pl 3 serrate. Leg III: femur av 1 serrate; genu av 1 and pv 1 serrate; tibia ad 3 serrate; tarsus ad 3 serrate. Leg IV genu pv 1 serrate; tarsus ad 3 serrate. Seta pv 1 on trochanter IV is large and serrate, L = 25 (20–25) (Fig. 9 B). Male (n = 11) Body oval similar to female. Idiosoma, L = 627–691, W = 429–467 (Fig. 7 B). Ve n tr al idiosoma. Tetartosternum, L = 31–37, anterior W = 78–87, shagreened (Fig. 11 B). Tetartosternal notch deep and distinct, 1 / 3 length of shield. Sternal seta st 1 simple, L = 7–10 (Fig. 11 A). Setae st 2, st 3 and st 4 are simple microsetae. Sternal shield shagreened, L = 62–68. Genital opening oval, L = 34–47, W = 53– 62. Ventral shield, L = 273–310, posterior W = 333–352, shagreened, with 28 setae, with two pores, one at level coxa III and other below the posterior margin of coxa IV (Fig. 7 B). Anal shield, L = 93–112, anterior W = 269–288; with five pairs of simple setae plus longer antero anal seta, L = 19–23, with two pores one near anterior margin and the other below anus. Gnathosoma. Hypostomal seta hyp 1 simple, L = 43–53, hyp 3 finely serrate and intermediate in length, L = 21–33, hyp 2 is simple and shorter, L = 9–12,. Etymology. This species is named in honour of Alejandro Valdez, who helped to collect almost all the passalid beetles used in this research. Remarks. The latigynial shield of this species is long and wavy, st 1 is long and serrated; and sternal setae st 3 and st 4 are microsetae. All shields are shagreened. The mesogynial shield has wide anterior and posterior margins. The male has a very deep tetartosternal notch, almost 1 / 3 of the length of the shield, and 28 pairs of setae on the ventral shield. Klinckowstroemia valdezi is similar to K. montanoi and K. scotti because they have wavy medial margins on the latigynial shield, and the same number of setae on the anal shield. The new species can be separated from these two species because st 1 is long and serrated (L = 31–40), the others setae are microsetae and simple. Klinckowstroemia montanoi has st 1 slightly shorter and serrate (L = 21–29), the other sternal setae are short and simple. Klinckowstroemia valdezi is similar to K. scotti because they have 12 pairs of setae a on the ventral shield, but they differ because K. scotti has four pairs of setae on the hyaline hood and the latigynial shield has eight pilose setae, while K. valdezi has one pair of setae a and three simple setae on the latigynial shield. Klinckowstroemia valdezi is similar to K. montanoi because they have one pair of setae a on the hyaline hood and st 3 is a microseta, but K. montanoi has five to eight setae on the latigynial shield and st 1 (23.8) is moderate and serrate, while K. valdezi has three setae on the latigynial shield and st 1 is long (36) and serrate. The specimens were found in decaying trunks, on three beetle species and at different localities: two from the state of Hidalgo and one from the state of Veracruz. The mites were found in the alcohol, but in some cases these were found on the passalid’s body: one mite was on femur III, one on the episternum and another under the elytron. Two Klinckowstroemia species were found on two passalids: the larger K. victoriae (idiosoma ca. 850) and the smaller K. valdezi (idiosoma 563–710). One passalid specimen had three mite species: K. bifurcata, K. hunteri and K. valdezi.

Klinckowstroemia brevimarginata sp. nov. Material examined. Holotype. Ƥ (CNAC 006585), Oaxaca, Mexico, 101 km Federal highway Puerto Escondido-Oaxaca (16 ° 28.156 ' N, 97 °01.674' W), 1,906 m, 28 June 2006, from Ve r re s aff. intermedius, O. Francke, A. Valdez and H. Montaño coll., oak forest. Paratypes. 5 Ƥ (CNAC 006586- CNAC 006590), 6 33 (CNAC 006591- CNAC 006596), same data as holotype, from two specimens of Ve r re s aff. intermedius. Holotype female, four paratype female and five paratype males deposited in the CNAC, one female and male paratypes deposited in NMNH. Female (n = 6). Body oval. Idiosoma, L = 762 (736–762), W = 525 (512–532) (Fig. 10 A). Ve n tr al idiosoma. Hyaline hood extending to level of coxa I, seta a 1 moderately long and simple, L = 19 (17–20). Tritosternum base wider than long. Tetartosternum shield L = 31 (28–31), anterior W = 99 (93–99), shagreened, anterior margin rounded, tetartosternal notch like a small cavity. Sternal seta st 1 near anterior margin, moderately long and serrate, L = 19 (16–22), lyriform sternal pore stp 1 near posterior margin. Sternal shield L = 62 (62–65), shagreened (Fig. 11 E). Pore stp 2 near centre of shield, behind it st 2, st 3 and st 4 are simple microsetae, the last two setae located on sides of shield near the corners, close to posterior margin. Sternogynial shield, L = 47 (47–53), W = 143 (140–152) triangular, narrow because mesogynial shield extends anteriorly (Fig. 11 D); posterior margin slightly pointed at apex, surface shagreened, stp 3 near anterior margin. Sternogynial apodeme long and thin, extending to posterior to level of latigynial and mesogynial condyles. Latigynial shields, L = 109 (109–124), W = 71 (68– 71), each with an oval pore and four to six setae, commonly with five (Fig. 11 D), medial margins of shield short and straight, surface of shield shagreened; latigynial apodeme thin. Mesogynial shield L = 84 (82–93), W = 99 (93–99), triangular, wider posteriorly, slightly pointed at apex, surface shagreened. Mesogynial condyles behind level of apex of shield. Ventral shield, L = 198 (176–198), posterior W= 403 (403–429), triangular (Fig. 11 G), truncated posteriorly, surface shagreened and reticulated laterally, with 13 pairs of small setae. One pore underneath posterior margin of coxa IV. Metapodal-peritremal-exopodal shield with a pair of pores, and a seta, surface reticulated. Anal shield, L = 109 (102–124), anterior W = 288 (288–307), with five pairs of simple setae plus longer antero anal seta (Fig. 13 A), L = 24 (22–25), two pairs of pores, one near anterior margin and the other below the anus. Gnathosoma. Capitular setae (sc) short and serrate, L = 14 (9.3–14). Hypostomal setae hyp 1 simple, L = 50 (50–56), longest. Seta hyp 2 medium length, serrate, L = 31 (31–34); hyp 3 short and simple, L = 12 (12–14) (Fig. 11 C). Chelicera with barbed seta, movable digit bifurcate at tip, with four teeth, fixed digit with six teeth. Most palpal setae simple, except al 1 and al 2 on trochanter are branched and serrate respectively; femoral setae av 1, ad 2, pl 1, pd 1 and pd 2 barbed; genu setae pl 1, pl 2, pd 1, pd 2 and pv 1 barbed. Anterior margin of palp trochanter with blunt process (b), and denticulated process divided (s) (Fig. 11 F). Legs. Most leg setae simple, except: Leg I: trochanter I ad 1, al 1, pl 1 and pv 1 barbed; femur I: pv 1, pv 2 and pv 3 barbed; genu I: av 1 and pv 1 barbed; tibia I: ad 1, ad 2, ad 3, pd 1 and pd 3 barbed. Leg II: femur II: av 1 and pv 1 barbed; genu II: av 1 and pv 1 barbed. Leg III: trochanter III: pv 1 and pl 1 barbed; femur III: av 1 barbed; genu III: av 1 and pv 1 barbed. Leg IV: trochanter IV: pv 1; femur IV: av 1 barbed; genu IV av 1 barbed. Seta pv 1 is large and serrated (Fig. 11 H), L = 32 (31–38). Male (n = 6) Body oval. Idiosoma, L = 736–781, W = 499–544 (Fig. 10 B). Ve n tr al idiosoma. Tetartosternum, L = 31–40, anterior W = 93–102, shagreened, anterior margin round (Fig. 13 B). Tetartosternal notch distinct, anterior margin divided in two parts, with a pore near posterior margin. Sternal seta st 1, L = 9–16, serrate. Sternal shield shagreened, L = 93–102, reticulated laterally (Fig. 13 C). Setae, st 2, st 3 and st 4 are simple microsetae; with two pores, one near anterior margin of shield, one close to genital opening. Genital opening oval, L = 47–53, W = 78–81. Ventral shield, L = 282–307, posterior W = 448, shagreened, with 38 setae and two pores, one at level of anterior margin of coxa IV and the other below coxa IV; this area of shield reticulated laterally (Fig. 10 B). Anal shield shagreened, L = 109–124, anterior W = 333–358, with five pairs of simple setae plus longer antero anal seta, L = 20–25, with two pores, one near anterior margin and the other below anus. Metapodal-peritremal-exopodal shield with a pair of pores, and a seta, reticulated. Gnathosoma. Hypostomal seta hyp 1 long and simple, L = 47–53, hyp 2 serrate and long, L = 31–40, hyp 3 simple and shorter, L = 12. Etymology. This name is from Latin brevis (= small) and marginata (= margin) referring to the size of the medial margins of the latigynial shields. Remarks. This species can be distinguished from all others because the mesogynial shield extends to the sternogynial shield and is wide on posterior margin; the margin of the latigynial shield is straight and reduced, 12 (9–16). Sternal seta st 1 is of moderate length (16–22) and serrate in both sexes and sternal setae st 2, st 3 and st 4 are microsetae and simple. The sternal, genital, ventral and tetartosternum shields are shagreened and the ventral shield is reticulated laterally. The male tetartosternal shield has a round anterior margin, the tetartosternal notch dividing the anterior margin in two, and 38 pairs of setae on the ventral shield. Klinckowstroemia brevimarginata is similar to K. rectimarginata because they have st 1 serrate and st 2 is a microseta; however, K. rectimarginata (813–902) is bigger than K. brevimarginata (736–762) and has three setae on the latigynial shield, while K. brevimarginata has four to six setae. Klinckowstroemia brevimarginata is similar to K. schusteri because it has four setae on the latigynial shield and st 1 serrated, but the former is shorter than the latter and setae st 2, st 3 and st 4 are microsetae, while in K. shusteri setae st 2, st 3 and st 4 are short, the mesogynial shield does not extend anteriorly, and the medial margin of the latigynial shields is long. The specimens were found in decaying trunks at the same locality at Oaxaca, Mexico. The mites were found in the alcohol with two passalids.

Klinckowstroemia hunteri sp. nov. Material examined. Holotype. Ƥ (CNAC 006530), Hidalgo, Mexico, 2 km Mojonera-Tizapan, municipality Zacualtipan (20 ° 38.107 ' N, 98 ° 35.981 ' W), 2,019 m, 22 September 2006, from Odontotaenius zodiacus, O. Francke, A. Valdez and H. Montaño coll., pine-oak forest. Paratypes. 2 Ƥ (CNAC 006531, CNAC 006532), 5 33 (CNAC 006533- CNAC 006537), same data as holotype, from 3 specimens of Odontotaenius zodiacus; 2 Ƥ, Hidalgo, Mexico, 2 km E Acaxochitlán (20 °08.500' N, 98 ° 10.740 ' W), 2,214m, 23 September 2006, from Odontotaenius zodiacus, O. Francke, A. Valdez and H. Montaño coll. pine-oak forest. Holotype female, two paratype females and four paratype males deposited in the CNAC, one female and one male paratypes deposited in NMNH. Female (n = 13). Body oval. Idiosoma, L = 966 (896–1,043); W = 666 (640–672) (Fig. 3 A). Ve n tr al idiosoma. Hyaline hood extending to level of coxa I, seta a 1 large and barbed, L = 42 (31–43). Tritosternum base wider than long. Tetartosternum L = 40 (34–40), anterior W = 124 (124–127), shagreened and reticulated as illustrated (Fig. 4 C). Tetartosternal notch triangular. Sternal seta st 1 simple (Fig. 4 D), L = 6, but broken on all specimens, lyriform sternal pore stp 1 near posterior margin. Sternal shield (Fig. 4 E) L = 87 (59–90), stp 2 near centre of shield, behind it sternal seta st 2, a simple microseta. Setae st 3, L = 20 (12–22) and st 4, L = 7 (6–8) near posterior margin of shield, both simple, close together. Shield reticulated and shagreened. Sternogynial shield (Fig. 4 G) L = 65 (62–68); W = 170 (161–174) triangular, anterior margin narrow and pointed at apex, surface shagreened and reticulated, stp 3 located closer to lateral margins. Sternogynial apodeme with broad base that rises to the shield, posterior arms thick, extending posteriorly to level of latigynial and mesogynial condyles. Latigynial shields, L = 143 (121–143); W = 68 (47–76), each with an oval pore and four setae, medial margin of shield straight and short (Fig. 4 G), shield reticulated, surface shagreened; latigynial apodeme thick and separate from sternogynial shield. Mesogynial shield L = 81 (53– 81), W = 84 (71–93), triangular, wider posteriorly and pointed at apex, anterior margin extending slightly anteriorly to sternogynal shield (Fig. 4 G), surface shagreened. Mesogynial condyles extending almost to level of shield apex. Ventral shield, L = 279 (251–282); posterior W = 544 (518–544), shield triangular, truncated posteriorly, surface shagreened (Fig. 6 B), with 12 pairs of small setae (Fig. 3 A). One pore underneath posterior margin of coxa IV (Fig. 3 A). Metapodal-peritremal-exopodal shield (MPES) with a pair of pores and one simple seta (Fig. 3 A, 6 B). Anal shield, L = 192 (102–198); anterior W = 474 (448–474) (Fig. 4 F), with six pairs of simple setae plus longer antero anal seta, L = 34 (32–34.1), two pairs of pores, one near anterior margin and the other below the anus. Gnathosoma. Capitular setae (sc) short and serrate, L = 20 (14–20). With three hypostomal setae (Fig. 4 H), hyp 1 long and simple, L = 65 (56–68), longest. Seta hyp 2 serrate, L = 40 (40–45). Seta hyp 3 short and simple, L = 12 (12–16). Chelicera with barbed setae, movable digit with four teeth, two small and two large; fixed digit with six teeth, two large and four small. Most palpal setae simple, but av 1 and av 2 of trochanter are branched and serrate respectively. On femur pv 1 is barbed; ad 1, ad 1, pd 1 and pd 2 are serrated. On genu ad 1 and ad 2 are serrate. Anterior margin of palp trochanter with blunt process (b), denticulated process (s) slightly longer (Fig. 6 A). Legs. Most leg setae simple, except: Leg I: trochanter I ad 1, av 1, al 1 and pv 1 serrated; femur I, al 1, pv 1 and pv 3 serrated; genu I, av 1 and pl 1 serrated. Leg II: femur av 1 and genu av 1 serrated. Leg III: femur av 1; genu av 1 and pv 1; tibia ad 3 and al 3 serrated. Leg IV: coxa av 1 and pv 1, tibia ad 4, tarsus ad 2 and ad 3 slightly serrated. Male (n = 4) Body oval, similar to female. Idiosoma, L = 992–1018, W = 717–755 (Fig. 3 B). Ve n tr al idiosoma. Tetartosternum, L = 47–56, anterior W = 124–133, shagreened, with one line near anterior margin (Fig. 6 D). Tetartosternal notch small, square, moderately deep (Fig. 6 D). Sternal seta st 1, L = 9–22, simple (Fig. 6 C). Sternal seta st 2, L = 12, simple, seta st 3, L = 6–11, simple, and st 4 is a microseta. Sternal shield shagreened, L = 153. Genital opening oval, L = 59–65, W = 89–96. Ventral shield with 36 setae, L = 372–388, shagreened, with two pores, one near anterior margin of coxa IV and other below the same coxa (Fig. 3 B). Anal shield, L = 183–217, with six pairs of simple setae, plus longer antero anal seta L = 12–34, with two pores: one near to anterior margin and the other below the anus. Gnathosoma. Hypostomal seta hyp 1 long and simple, L = 65–71, seta hyp 3 serrate and long, L = 43–47, seta hyp 2 simple and shorter, L = 12–16. Legs. Seta pv 1 on coxa IV slightly serrated (Fig. 6 E). Etymology. This species is named in honour of Preston E. Hunter, for his contribution to the study of mites associated with passalids, especially of the family Klinckowstroemiidae. Remarks. This species can be distinguished from all others because the latigynial and sternogynial apodemes are strongly developed and wider, separating the sternogynial and latigynial shields. The mesogynial shield has a wide posterior margin. The tetartosternum, sternal, sternogynial and latyginal shields are reticulated and shagreened. The male has 34 setae on the ventral shield, and the tetartosternum is hexagonal in shape and expanded. Klinckowstroemia hunteri is similar to K. atramaculata, because they have the latigynial shield long and st 1 barbed, but the last species lacks the well-developed latigynial and mesogynial apodemes, has three setae on latigynial, and seta a 1 is simple, while, K. hunteri has four setae on latigynial shield and seta a 1 long and barded. The male of K. atramaculata has 13–15 pair of setae, while the male of K. hunteri with 17 pairs. The species K. bifurcata and K. hunteri were found together on three specimens of Odontotaenius zodiacus. It is possible that these passalids lived in the same log, because they share the same klinckowstroemiids and they were collected at the same locality on the same day. One of these passalid had three different species of the genus Klinckowstroemia. Furthermore seven (5 Ƥ, 2 33) specimens of K. hunteri were collected together with one female of K. victoriae on the same species of passalid.

Klinckowstroemia cristinae sp. nov. Material examined. Holotype. Ƥ (CNAC 006649), Hidalgo, Mexico, 2 km Federal highway Crucero- Tianguistengo (20 ° 40.413 ' N, 98 ° 40.255 ' W), 2,080 m, 23 September 2006, pine-oak forest, from Odontotaenius zodiacus, O. Francke, A. Valdez and H. Montaño coll. Paratypes. 3 Ƥ (CNAC 006650- CNAC 006652), 2 33 (CNAC 006653, CNAC 006654), same data as holotype, from two specimens of Odontotaenius zodiacus; 1 Ƥ (CNAC 006655), Hidalgo, Mexico, 2 km Federal highway Crucero- Tianguistengo (20 ° 40.413 ' N, 98 ° 40.255 ' W), 2,080 m, 23 September 2006, pine-oak forest, from Oileus rimator, O. Francke, A. Valdez and H. Montaño coll. Holotype, three paratype females and two paratype males deposited in the CNAC, one female paratype deposited in NMNH. Female (n = 5). Body oval. Idiosoma, L = 928 (928–1005), W = 672 (672–710) (Fig. 21 A). Ve n tr al idiosoma. Hyaline hood extending to level of coxa I, seta a 1 long and slightly serrated (Fig. 23 A), L = 34 (32–38). Tritosternum base wider than long. Tetartosternum shield (Fig. 22 D), L = 43 (40–43), anterior W = 115 (115–124); shagreened, anterior margin slightly concave, medial notch small, triangular. Sternal seta st 1 moderately long, slightly serrated (Fig. 22 E), L = 20 (20–25), near centre of shield; lyriform sternal pore stp 1 close to posterior margin of shield. Sternal shield, L = 81 (81–93) shagreened. Pore stp 2 near anterior margin of shield, behind and below it is sternal seta st 2, medium length and simple, L = 19 (12–19). Setae st 3 long, slightly serrated, L = 28 (25–37), st 4 is medium length, slightly serrated, L = 16 (16–25). Setae st 3 and st 4 towards sides of shield, close to posterior margin (Fig. 22 C). Sternogynial shield, L = 62, W = 171 (164– 186) triangular, posterior margin pointed at apex, surface shagreened, stp 3 near anterior margin (Fig. 21 A). Sternogynial apodeme long and thin, extending to posterior level of latigynial and mesogynial condyles. Latigynial shields (Fig. 22 C), L = 124 (124–140), W= 78 (74–78) each with an oval pore and three or four simple setae, usually four; medial margins of shields straight and short, shield are shagreened; latigynial apodeme long and thin. Mesogynial shield, L = 74 (71–78), W = 99 (93–102), triangular, wider posteriorly and anteriorly (Fig. 22 C), rounded at apex, surface shagreened, mesogynial condyles below level of shield apex. Ventral shield, L = 245 (245–264), posterior W = 531 (512–557), triangular, truncated posteriorly (Fig. 22 F), surface shagreened, reticulated like a honey-comb (Fig. 22 G), with 12–14 pairs of small setae, usually 14. One pore underneath posterior margin of coxa IV. Metapodal-peritremal-exopodal shield reticulated, with a pore, with two setae. Anal shield, L = 161 (155–177), anterior W = 448 (435–480), with six pairs of simple setae plus longer antero anal seta, L = 29 (29–33), shield shagreened, with two pairs of pores, one near anterior margin and the other below the anus. Gnathosoma. Capitular setae (sc) medium and simple, L = 16 (12–17). Hypostomal setae hyp 1 longest and simple, L = 66 (65–71). Seta hyp 2 long and barbed, 40 (40–45), hyp 3 moderately long and simple, L = 19 (16–19). Chelicera with barbed seta, movable digit with four teeth, two large and two small, fixed digit with five teeth, all similar in size, movable finger divided at the tip. Palpal setae are generally simple, but al 1 (av 1) and al 2 (av 2) on trochanter are branched and serrate respectively. Femur setae pv 1 serrated, al 2, ad 1 and ad 2 barbed. Genu setae al 1 and al 2 are barbed, pl 1 and ad 1 are serrated. Anterior margin of palp trochanter with a blunt process (b) and process (s) (Fig. 22 H). Legs. Most leg setae simple, except: Leg I: trochanter ad 1, al 1 and pv 1 serrated; femur pv 1, pv 2 and pv 3 serrated; genu al 1, av 1 and pv 1 serrated; tibia ad 1, ad 2, ad 3, pd 1 and pd 3 serrated. LegII: femur av 1 and pv 1 serrated; genu av 1 and pv 1 serrated. Leg III: trochanter pv 1 slightly serrated; femur av 1 serrated; genu av 1 and pv 1 serrated; tibia ad 3 serrated. Leg IV: trochanter pl 1 slightly serrated; femur av 1 serrated; genu av 1 and pv 1 serrated; tibia ad 3 barbed. Seta pv 1 is serrated and long, 26 (26–30). Male (n = 3) Body oval, similar to female. Idiosoma, L = 966–1011, W = 698–717 (Fig. 21 B). Ve n tr al idiosoma. Tetartosternum (Tst), L = 47–53, anterior W = 124–127, shagreened (Fig. 23 D), anterior margin concave as in female. Tetartosternal notch small, V-shaped and divides anterior margin in two parts, shield with a pore near posterior margin. Sternal seta st 1, L = 22–25, slightly serrated. Sternal shield shagreened (Fig. 23 B), L = 143–149. Setae st 2, st 3 and st 4, microseta simple; shield with two pores, one near anterior margin of the shield, one below coxa IV. Genital opening oval, L = 62, W= 87–90. Ventral shield, L = 372, posterior W= 544–557; shagreened and reticulated as in female, 34 setae (Fig. 21 B); with two pores, one near anterior margin of coxa IV and one below of coxa IV. Anal shield shagreened, L = 171–177, anterior W = 454; with six pairs of simple setae plus longer antero anal aa seta, L = 31–34, with two pores, one near anterior margin and one below anus. Metapodal-peritremal-exapodal shield reticulated, with a pore, without setae. Gnathosoma. Hypostomal setae (Fig. 23 C), hyp 1 long and simple, L = 65–71, hyp 2 serrated and long, L = 43–47, hyp 3 simple and medium length, L = 19. Etymology. This specific name cristinae is named in honour of Mrs. Cristina Olvera for her hospitality and kindness in hosting the senior author during this research. Remarks. This species can be distinguished from all others because the anterior margin of the tetartosternum shield is concave in both sexes, and sternal seta st 3 is the longest and is slightly serrated. The mesogynial shield is long and wider posteriorly; the latigynial shield is long with the medial margins reduced, all the shields are shagreened, except the ventral shield, which is reticulated. The male has st 2, st 3 and st 4 microsetae and simple; the ventral shield is reticulated and shagreened, and the other shields are only shagreened. The tetartosternal notch is small and V-shaped, and the ventral shield has 34 pairs of simple setae. Klinckowstoemia cristinae is similar to K. schusteri because they have three to four simple setae on the latigynial shield, hyp 2 and hyp 3 are serrated and long and moderatly long respectively, in both species. However, the new species is smaller (idiosoma 953.6 versus 1374), has sternal seta st 3 (29.2) long and slightly serrated, and has shagreened sternogenital shields. In comparison, K. schusteri has several shields reticulated (sternal, sternogynial, mesogynial and latigynial) and st 3 (8) small and simple. Both species have large latigynial shields but that of K. cristinae has reduced medial margins (19.5) compared with K. schusteri. The passalids carrying this species were found in decaying trunks in the same locality at Hidalgo, Mexico, on two hosts, Odontotaenius zodiacus and Oileus rimator. Mites were found in the alcohol, but on O. zodiacus we found a male of K. cristinae on coxa I. On O. rimator we found two other species of klinckowstroemiids, K. scotti and K. cristinae.

Klinckowstroemia bifurcata sp. nov. Material examined. Holotype. Ƥ (CNAC 006514), Hidalgo, Mexico, 2 km Federal Highway Crucero- Tianguistengo (20 ° 40.413 ' N, 98 ° 40.255 ' W), 2,080 m, 23 September 2006, pine-oak forest, from Odontotaenius zodiacus (Truqui), O. Francke, A. Valdez and H. Montaño coll., mites found in the alcohol. Paratypes. 4 Ƥ (CNAC 006515- CNAC 006518), 1 3 (CNAC 006519), same data as holotype, from 3 specimens of Odontotaenius zodiacus; 8 Ƥ (CNAC 006520- CNAC 006527), 2 33 (CNAC 006528, CNAC 006529), Hidalgo, Mexico, 2 km Mojonera-Tizapan, municipality Zacualtipan (20 ° 38.107 ' N, 98 ° 35.981 ' W), 2,019 m, 22 September 2006, from 4 specimens of Odontotaenius zodiacus, O. Francke, A. Valdez and H. Montaño coll. Mites were on coxa I, mesepisternum and loose in the alcohol. Holotype, 9 paratype females and 3 paratype males deposited in the CNAC, one female paratype each deposited in NMNH, UGA and UMMZ. Female (n = 13). Body oval. Idiosoma: L = 966 (922–1,088), W = 723 (659–762) (Fig. 1 A). Ve n tr al idiosoma. Hyaline hood extending to level of coxa I, seta a 1 large and simple (Fig. 2 G), L = 34 (30–37). Tritosternum base wider than long. Tetartosternum (Tst) L = 40 (40–44), anterior W= 124 (112–127) (Fig. 2 E); near the posterior margin is a lyriform sternal pore stp 1. Tetartosternal notch square, shagreened and with a line near the anterior margin. Sternal seta st 1, L = 18 (16–22), bifurcated (Fig. 2 C). Sternal shield (Fig. 2 B) L = 90 (78–91), stp 2 in centre of shield, below it st 2, L = 11 (9–19), simple. Setae st 3, L = 31 (19– 40) and st 4, L = 16 (9–22) near posterior margin of shield, both large and simple, close together (Fig. 2 B). Shield shagreened, without reticulations. Sternogynial shield, L = 68 (62–96); W = 192 (167–201) triangular (Fig. 2 F), shagreened, with stp 3 closer to anterior margin. Sternogynial apodeme short. Latigynial shields, L = 130 (124–146), W = 76 (75–84) each with an oval pore and four setae (Fig. 2 F), medial margin of shield straight, extremity blunt. Mesogynial shield, L = 76 (68–84), W = 106 (96–112), triangular, wider anteriorly and slightly pointed at apex, surface shagreened. Mesogynial condyles extending almost to level of shield apex. Ventral shield, L = 269 (248–288), posterior W = 576 (518–608), shield triangular, truncated posteriorly, surface shagreened, with 12 pairs of small setae (Fig. 1 A). One pore at level of posterior margin of coxa IV. Anal shield, L = 192 (167–218); anterior W = 518 (467–576), with six pairs of simple setae plus longer antero anal seta, L = 25 (17–31), two pairs of pores, one near the anterior margin, the other below the anus. Gnathosoma. Capitular seta (sc) short and serrate, 12 (9–16). Hypostomal seta hyp 1 long and simple, 62 (56–71). Seta hyp 2 long and serrate, L = 40 (34–47). Seta hyp 3 short and simple, L = 7 (7–19) (Fig. 2 A). Chelicera with barbed seta, movable digit with four teeth, fixed digit with five teeth. Palp setae are simple, av 1 and av 2 on trochanter branched and serrate respectively; dorsal setae of femur pl 1 and pl 2 serrate and feathery respectively (Fig. 2 A). Anterior margin of palp trochanter with blunt process (b) (Fig. 2 H). Legs. Most setae on leg I serrate, on trochanter ad 1, al 1 and pv 1, genu al 1 and av 1. Serrate setae on leg II: femur av 1 and genu av 1. Serrate setae on leg III: femur av 1, genu pv 1, tibia ad 3 and al 3; on leg IV, genu pv 1, tibia ad 3 and al 3. Other setae simple, except pv 1 on trochanter II–IV lightly barbed (Fig. 2 D). Male (n = 3). Body oval, similar to female. Idiosoma L = 986–1024, W = 691–736 (Fig. 1 B). Ve n tr al idiosoma. Tetartosternum L = 50–53, posterior W = 102–112, tetartosternal notch shallow (Fig. 4 A), shagreened, with one line near anterior margin as in female. Sternal seta st 1, L = 19–24, simple. The other sternal setae are microsetae. Sternal shield shagreened, L = 152–155 (Fig. 4 B). Genital opening oval, L = 59–66, W = 87–94. Ventral shield with 32 setae (Fig. 1 B), L = 363–378, posterior W = 608, shagreened. Anal shield L = 179–217, anterior W = 512–544; with six pairs of simple setae plus longer antero anal seta, L = 20–30. Gnathosoma. Hypostomal seta hyp 1 is long and simple, L = 65–71; seta hyp 2 is serrate and long, L = 43– 50, while seta hyp 3 is simple and shorter, L = 16–17. Etymology. This name is from Latin bifurcata (= bifurcate) referring to the shape of st 1. Remarks. This species can be distinguished from all others because sternal seta st 1 is bifurcate; and the tetartosternum, sternal and genital shields are shagreened. Sternal setae st 3 and st 4 are slightly serrated, seta st 3 longest. The mesogynial shield is wider on the posterior margin and the latigynial shield has four small simple setae and a pore. Some setae on the palp and leg I are serrated and feathery. The male has sternal seta st 1 simple and the tetartosternum is similar to that of the female. Klinckowstroemia bifurcata is similar to K. reyesi and K. truncata because these three species have the same number of setae on the latigynial, ventral and anal shields. Klinckowstroemia bifurcata (981) is slightly bigger than K. truncata (820) but is slightly smaller than K. reyesi (1009). Klinckowstroemia bifurcata has four setae on the latigynial shield and sternal seta 3 and 4 are serrate as in K. truncata. Sternal seta 3 is longest as in K. reyesi. Klinckowswtroemia bifurcata has the medial margin of the latigynial shield straight and its extremity blunt, while that of K. reyesi and K. truncata is longer and completely straight. The specimens were found in decaying trunks at two different localities in the state of Hidalgo, Mexico, but on the same host (Odontotaenius zodiacus). In these localities we collected three and four passalids carrying K. bifurcata. The mites were usually found in the alcohol, only in one case they were located on the passalid’s body, on coxa I and mesoepisternum. Two or more mite species were found on one beetle: one male of K. victoriae, seven (4 Ƥ, 3 33) of K. bifurcata; and another passalid was found with three species, K. bifurcata; K. hunteri and K. valdezi. This association is known as synhospitality (Eichler 1966).

Klinckowstroemia franckei sp. nov. Material examined. Holotype. Ƥ (CNAC 006603), Hidalgo, Mexico, Ejido Atepixco, Municipality Zacualtipan (20 ° 35.466 ' N, 98 ° 36.358 ' W), 1,972 m, 22 September 2006, from Odontotaenius zodiacus, pineoak forest, O. F. Francke coll.. Paratypes. 6 Ƥ (CNAC 006604- CNAC 006609), 10 33 (CNAC 006610- CNAC 006619), same data as holotype, from nine specimens of Odontotaenius zodiacus. Holotype, three paratype females and seven paratype males deposited in the CNAC, one female and female paratype each deposited in NMNH, UGA and UMMZ. Female (n = 7). Body oval. Idiosoma, L = 909 (896–941); W = 627 (627–666) (Fig. 14 A). Ve n tr al idiosoma. Hyaline hood extending to level of coxa I, seta a 1 long and slightly barbed, L = 47 (47– 53). Tritosternum base wider than long. Tetartosternum shield, L = 37, anterior width 124 (121–124); shagreened and reticulated (Fig. 15 E), anterior margin slightly wavy with a small square notch. Simple microseta st 1 near posterior margin of shield (Fig. 17 D), lyriform pore stp 1 near middle of shield, closer to posterior margin. Sternal shield, L = 90 (84–90), shagreened and reticulated (Fig. 17 A). Pore stp 2 near middle of shield, behind it sternal seta st 2, short and simple, L = 7, only seen in one specimen (Fig. 17 A). Setae st 3 is medium length and simple, L = 20 (12–22), st 4 simple and short, L = 9 (6–9) (Fig. 17 A), st 3 and st 4 near corners of shield, close to posterior margin. Sternogynial shield, L = 68 (65–71), W = 171 (140–171) triangular, posterior margin slightly pointed at apex, surface shagreened and reticulated (15 F), stp 3 near anterior margin (Fig. 15 F). Sternogynial apodeme long and thick, extending posteriorly to level of latigynial and mesogynial condyles. Latigynial shields, L = 130, W = 65 (65–71) each with an oval pore and four setae, medial margins of shields pointed, shield shagreened and reticulated; latigynial apodeme thin and long. Mesogynial shield, L = 78 (71.3–78), W = 74 (68–84), triangular, wider posteriorly and pointed at apex, extending slightly anterior to sternogynal shield, surface shagreened, mesogynial condyles under level of shield apex. Ventral shield, L = 242 (242–260); posterior W = 480 (467–512), triangular, truncated posteriorly, surface shagreened (Fig. 17 C), with 12 pairs of small setae (Fig. 14 A). One pore underneath posterior margin of coxa IV. Metapodal-peritremal-exopodal shield (MPES) reticulated, with a pair of pores (Fig. 17 C) and a seta. Anal shield, L = 149 (149–164); anterior W = 416 (390–429), with five pairs of simple setae plus longer antero anal seta, aa, L = 31 (29–32), two pairs of pores, one near anterior margin, another below the anus. Gnathosoma. Capitular setae (sc) short and serrate, L = 14 (12–16). Hypostomal setae hyp 1 longest, simple, L = 65 (62–65); hyp 2 medium length and barbed, L = 37 (36–40); hyp 3 short and slightly serrated, L = 15 (14–16) (Fig. 15 G). Chelicera with barbed seta, movable digit with four teeth, fixed digit with six teeth. Most palpal setae simple, except al 1 and al 2 on trochanter are branched and serrate respectively. Femur setae av 1 and pd 1 barbed, ad 1 serrated. Anterior margin of palp trochanter with blunt process (b) and denticulated process (s) (Fig. 15 H). Legs. Leg setae mostly simple, except: Leg I: trochanter ad 1 is barbed, pv 1 serrated; femur pv 1, pv 2, pv 3 and av 1 barbed; genu av 1 barbed, pv 1 serrated. Leg II, femur av 1 barbed; genu av 1, pv 1 barbed. Leg III: femur av 1 barbed; genu pv 1, av 1 slightly barbed. Leg IV: trochanter pv 1, pl 1 slightly serrated; femur av 1 barbed; genu av 1, pv 1 slightly barbed; tibia al 3 barbed; tarsus al 2 serrated. Seta pv 1 is large and slightly serrated, L = 29 (28–29) (Fig. 17 B). Male (n = 10) Body oval, similar to female. Idiosoma, L = 896–960, W = 621–659 (Fig. 14 B). Ve n tr al idiosoma. Tetartosternum, L = 34–47, anterior W = 118–124, shagreened, anterior margin round. Tetartosternal notch small, dividing anterior shield margin in two parts, stp 1 near posterior margin. Sternal shield shagreened and reticulated, L = 148 (143–152). Sternal seta st 1 simple, L = 16–19, st 2, st 3 and st 4, are microsetae and simple; shield with two pores, stp 2 near anterior margin, stp 3 close to genital opening. Genital opening oval, L = 56–62, W = 78–87. Ventral shield, L = 326–357, posterior W = 486–550, shagreened, reticulated only near anterior margin, with 26–32 setae and two pores, one at level of anterior margin of coxa IV, the other below coxa IV (Fig. 14 B). Anal shield shagreened, L = 140–161, anterior W = 397–429, with six pairs of simple setae plus longer antero anal seta, L = 28–37, with two pores, one near anterior margin and one below anus. Metapodal-peritremal-exopodal shield reticulated with a pair of pores, with seta. Gnathosoma. Hypostomal seta hyp 1 long and simple, L = 62–68, hyp 2 barbed and intermediate, L = 37– 43, hyp 3 simple and short, L = 12–14. Etymology. This species is named in honour of Oscar F. Francke who collected the many of the passalid beetles that carried Klinckowstroemiidae. Remarks. This species can be distinguished from all others because setae st 1 and st 2 are microsetae, and st 3 is the longest of the sternal setae. The tetartosternal, sternal, mesogynial and latigynial shields are shagreened and reticulated. the medial margins of the latigynial shield are pointed, and there are four small simple setae on this shield. The mesogynial shield has a broad posterior margin and its apex is anteriorly pointed. The male has st 1 medium sized and simple, with a small tetartosternal notch that divides the anterior margin in two parts; the tetartosternum and sternal shield are reticulated and shagreened. The other sternal setae are microsetae. The ventral shield has 26–32 setae, but many of the specimenss have 28. Klinckowstroemia franckei is similar to K. multisetillosa because they have similar size (K. franckei L= 908,W= 627; K. multisetillosa L= 936, W= 634), st 1 is simple, and the medial margin of the latigynial shield is pointed. However, the new species has only one pair of seta a, while K. multisetillosa has six pairs. Furthermore, the latigynial shield of K. franckei has four setae and is larger than that of K. multisetillosa, which has five setae. The posterior margin of the mesogynial shield is broader and more convex in K. multisetillosa than in K. franckei. The capitular seta is also small and simple in K. multisetillosa, but in K. franckei it is twice as long and serrated. The beetles carrying these mites were found in decaying trunks at the same locality; it is possible that some specimens occupied the same trunk as a family group. Almost all mites were found in the alcohol in the vials where the beetles were collected. Three mites were found on the body: one on the mesoepisternum, one on the mesosternellum and another near to coxa III. Each mite was found on a different passalid.

Klinckowstroemia Baker & Wharton, 1952: 135. Type species Klinckowstroemia tragardhi Baker & Wharton by original designation. Diagnosis (modified from Rosario & Hunter 1988) Hyaline hood forming crescent above gnathosoma that may extend posteriorly along lateral margin of dorsal shield, but not surrounding idiosoma; only seta a 1 present on anterior dorsal portion of hood, or more than three pairs of a setae present. Dorsal shield with an even margin, without visible constrictions; posterior portion of dorsal shield without ridges. Anal shield separate from ventral shield. Fused metapodal-peritremalexopodal region separated from ventral shield by metapodal suture. Pedofossae III present. Anteroanal setae longer than other setae on anal shield. Female latigynial shield triangular with pointed, blunt or sinuous medial margins; sternogynial and mesogynial shield without setae. Sternoventral suture present in males. In addition to the type species, the other known species are K. concava Hunter & Butler, 1966, K. truncata Hunter & Butler, 1966, K. grabowskii Chernoff & Pope, 1970, K. tapachulensis Chernoff & Pope, 1970, K. atramaculata Rosario & Hunter, 1988, K. candidoi Rosario & Hunter, 1988, K. multisetillosa Rosario & Hunter, 1988, K. reyesi Rosario & Hunter, 1988, K. schusteri Rosario & Hunter, 1988, K. scotti Rosario & Hunter, 1988, K. simplesetosa Rosario & Hunter, 1988, K. starri Rosario & Hunter, 1988, K. victoriae Rosario & Hunter, 1988., Published as part of Villegas-Guzman, Gabriel A., P��rez, Tila M. & Reyes-Castillo, Pedro, 2009, New species of the genus Klinckowstroemia Baker & Wharton from Mexico (Acari: Mesostigmata: Trigynaspida: Klinckowstroemiidae), pp. 1-46 in Zootaxa 2248 on page 4, DOI: 10.5281/zenodo.190717, {"references":["Baker, E. W. & Wharton, G. W. (1952) An Introduction to Acarology. Macmillan: New York, 465 pp.","Rosario, R. M. T. & Hunter, P. E. (1988) The genus Klinckowstroemia Tragardh and descriptions of nine new species (Acarina: Trigynaspida: Klinckowstroemiidae). Acarologia, 29, 119 - 136.","Hunter, P. E. & Butler, L. (1966) New Klinckowstroemia mites from Costa Rican passalid beetles (Acarina: Klinckowstroemiidae). Journal of the Georgia Entomological Society, 1, 24 - 30.","Chernoff, N. & Pope, R. (1970) Two new mites of the genus Klinckowstroemia from Mexico (Acarina: Klinckowstroemiidae). Proceedings of the Biological Society of Washington, 83, 145 - 154."]}

Klinckowstroemia ballesterosi sp. nov. Material examined. Holotype. �� (CNAC 006640), Veracruz, Mexico, Municipality of Soledad Atzompa, Soledad Atzompa (18 �� 44.712 ' N, 97 �� 40.967 ' W) 2,125 m, 23 March 2007, from Odontotaenius zodiacus, pine-oak forest, O. Francke, A. Valdez, C. Santiba��ez, A. Ballesteros and H. Monta��o coll. Paratypes. 3 �� (CNAC 006641- CNAC 006643), 1 3 (CNAC 006644), same data as holotype, from three specimen of Odontotaenius zodiacus; 2 �� (CNAC 006645, CNAC 006646), Veracruz, Mexico, Tlacolulan, Municipality of Tlacolulan (19 �� 38.508 ' N, 96 �� 59.650 ' W), 1,876 m, 30 September 2006, from one specimen of Odontotaenius zodiacus, oak forest, O. Francke, C. Santiba��ez, A. Valdez and H. Monta��o coll.; 2 �� (CNAC 006647, CNAC 006648), Oaxaca, Mexico, 8 km SE San Miguel Talea de Castro, Municipality of San Miguel Talea de Castro (17 �� 19.620 ' N, 96 �� 17.403 ' W), 2,082 m, 22 July 2007, evergreen cloud forest, from one specimen of Petrejoides recticornis, O. Francke, A. Valdez, C. Santiba��ez, A. Ballesteros and H. Monta��o coll. Holotype, four paratype females and one paratype male deposited in the CNAC, one female paratype each deposited in NMNH, UGA and UMMZ. Female (n = 8). Body oval. Idiosoma, L = 992 (896���1088), W = 742 (640���774) (Fig. 19 A). Ve n tr al idiosoma. Hyaline hood extending to level of coxa I, seta a 1 long and barbed, L = 47 (37���50). Tritosternum base wider than long. Tetartosternum shield, L = 43 (37���43), anterior W = 121 (112���127), shagreened and reticulated (Fig. 18 H), anterior margin with two small protuberances, corners of shield pointed, medial notch small, square. Sternal seta st 1 long and serrated (Fig. 20 A), L = 57 (47���68), near centre of shield; lyriform pore stp 1 close to posterior margin. Sternal shield (Fig. 20 B), L = 87 (74���90) shagreened and reticulated (Fig. 20 C). Pore stp 2 near anterior margin of shield, behind and below it seta st 2, long and serrated, L = 50 (37���54). Setae st 3 long and serrated, L = 47 (35���59), st 4 serrated and moderately long, L = 37 (31���47) (Fig. 20 B). Setae st 3, st 4 on sides of shield, close to posterior margin. Sternogynial shield, L = 50 (47���62), W = 186 (167���189) triangular (Fig. 20 D), posterior margin pointed at apex, surface shagreened, with stp 3 near anterior margin. Sternogynial apodeme long and thick, extending posteriorly to level of latigynial and mesogynial condyles. Latigynial shields, L = 130 (124���140), W= 84 (71���84) (Fig. 20 D) each with an oval pore and four or five simple setae, usually four; medial margins of shields are straight, shields shagreened; latigynial apodeme thin and long. Mesogynial shield, L = 87 (81���96); W = 112 (99���115), triangular (Fig. 20 D), wider posteriorly and anteriorly, apex pointed, surface shagreened, mesogynial condyles below level of shield apex. Ventral shield, L = 248 (223���273), posterior W = 576 (512���627), triangular, truncate posteriorly, surface shagreened, with 14���15 pairs of small setae (Fig. 20 G). One pore underneath posterior margin of coxa IV. Metapodal-peritremal-exopodal shield reticulated, with a pair of pores and two setae. Anal shield, L = 195 (149���198), anterior W = 524 (448���544), shield shagreened (Fig. 19 A), with five pairs of simple setae plus longer antero anal seta, L = 31 (29.��� 35), two pairs of pores, one near anterior margin, another below the anus. Gnathosoma. Capitular setae (sc) small and serrated, L = 7 (7���18). Hypostomal setae hyp 1 long and simple, L = 68 (59���68), longest; hyp 2 long and barbed, 45 (39���47); hyp 3 medium length and barbed, L = 22 (12���22). Chelicera with barbed seta, movable digit with four teeth, two large and two small; fixed digit with six teeth, two large and four small, movable digit divided at the tip. Palp setae are simple, except al 1 and al 2 on trochanter are branched and serrate respectively. Femur setae pl 1, pl 2 and pv 1 serrated, avl 1 barbed; genu seta ad 1 and pl 2 are barbed, pv 1 and ad 2 are serrated. Anterior margin of palp trochanter with a small blunt process (b) and denticulate process (s) (Fig. 20 F). Legs. Most leg setae simple, except Leg I: trochanter ad 1, al 1 and pv 1 barbed; femur pv 1, pv 2 and pv 3 serrated; genu al 1, pl 1 and pv 1 serrated; tibia ad 1, ad 2, ad 3, pd 1 and pd 3 serrated. Leg II: trochanter ad 1 serrated; femur: av 1 and pl 1 serrated; genu: av 1 and pv 1 serrated. Leg III: trochanter pv 1 serrated; femur av 1 and pl 1 serrated; genu av 1 and pv 1 serrated. Leg IV: trochanter pv 1 and p 12 serrated; femur av 1 serrated; genu av 1 and pv 1 serrated. Seta pv 1 is slightly serrated and long, 34 (29���35) (Fig. 20 E). Male (n = 3) Body oval, similar to female. Idiosoma, L = 1005���1018, W = 723���768 (Fig. 19 B). Ve n tr al idiosoma. Tetartosternum, L = 43, anterior W = 118���124, shagreened (Fig. 20 H), anterior margin straight, corners pointed as in female. Tetartosternal notch small, V-shaped, divides anterior margin of shield in two parts, with a pore near posterior margin. Sternal seta st 1, L = 9���16, simple (Fig. 20 H). Sternal shield shagreened (Fig. 22 A), L = 140���149. Sternal setae st 2 a simple microseta, st 3 long and serrated, L = 34���43, st 4 is long and serrated, L = 40���43; shield with two pores, one near anterior margin and one near posterior margin. Genital opening oval, L = 53���59, W= 78���84. Ventral shield L = 307���390, posterior W = 576���640, shagreened, with 34 setae (Fig. 22 B); with two pores, one near anterior margin of coxa IV and one below coxa IV. Anal shield shagreened, L = 180���202, anterior W = 512���563; with five pairs of simple setae plus longer antero anal seta, L = 31���34, and two pores, one near anterior margin and one below anus. Metapodalperitremal-exopodal shield reticulated, with a pair of pores and one seta. Gnathosoma. Hypostomal seta hyp 1 long and simple, L = 68���71, hyp 2 simple and long, L = 42���43, hyp 3 is simple and medium length, L = 14���16. Etymology. This species is named in honour of Alfonso Ballesteros, who helped to collect almost all the passalid beetles used in this research. Remarks. This species can be distinguished from all others because the sternal setae are long and serrated, and seta st 1 is the longest. In females the tetartosternal and sternal shields are shagreened and reticulated, and the other shields are only shagreened. The latigynial shield with has four or five simple setae. The mesogynial shield is broad posteriorly, and the form of the tetartosternal shield is very different to others species. In the male st 3 and st 4 long and serrated, and the tetartosternal, sternal, ventral and anal shields are shagreened. The tetartosternal notch is small, V-shaped, and divides the anterior margin in two parts. The ventral shield has 17 pairs of setae. Klinckowstroemia ballesterosi is similar to K. santibanezi and K. candidoi because they have setae a 1 and st 1 long and barbed, but the new species is smaller (896���1088 versus 1164��� 1517 and ca. 1290 respectively), and all sternal setae are serrated and long, with fourto five simple setae on the latigynial shield. In contrast, in K. santibanezi the sternal setae are barbed and serrated, and there are eight serrated setae on the latigynial shield; and in K. candidoi the sternal setae are serrated and simple and there are three to four simple setae on the latigynial shield. Klinckowstroemia ballesterosi has more setae on the ventral shield (K. ballesterosi 14, K. santibanezi 11���14, K. candidoi 11). In K. ballesterosi the tetartosternum and sternal shield are reticulated, while in K. santibanezi and K. candidoi the ventral and anal shields are reticulated. The beetles carrying this species were found in decaying tree trunks in two different states of Mexico. In Veracruz they were found on Odontotaenius zodiacus and almost all specimens were found in the alcohol, with only one specimen on the humerus of one of the beetles. From Oaxaca it was found with Petrejoides recticornis free in the alcohol., Published as part of Villegas-Guzman, Gabriel A., P��rez, Tila M. & Reyes-Castillo, Pedro, 2009, New species of the genus Klinckowstroemia Baker & Wharton from Mexico (Acari: Mesostigmata: Trigynaspida: Klinckowstroemiidae), pp. 1-46 in Zootaxa 2248 on pages 39-40, DOI: 10.5281/zenodo.190717

Klinckowstroemia santibanezi sp. nov. Material examined. Holotype. Ƥ (CNAC 006620), Oaxaca, Mexico, Municipality of Ixtlán de Juárez (17 ° 31.722 ' N, 96 ° 30.678 ' W), 2,742 m, 13 November 2005, pine forest, from Proculejus sartorii, O. Francke, M. Cordova, A. Jaimes and G. Montiel coll. Paratypes. 1 Ƥ (CNAC 006621), 1 3 (CNAC 006622), same data as holotype, from one specimen of Proculejus sartorii; 7 Ƥ (CNAC 006623- CNAC 006629), 4 33 (CNAC 006630- CNAC 006633), Oaxaca, Mexico, Cerro Pelón, highway Tuxtepec-Oaxaca (17 ° 35.75 ' N, 96 ° 28.41 ' W), 2,240 m, 22 November 1996, evergreen cloud forest, from one specimen of Proculejus brevis, inside log of Persea americana, F. Lorea. coll.; 2 Ƥ (CNAC 006634- CNAC 006635), Oaxaca, Mexico, Municipality of Ixtlán de Juárez (17 ° 31.722 ' N, 96 ° 30.678 ' W), 2,742 m, 13 November 2005, pine forest, from one specimen of Vindex sp. nov., O. Francke, M. Cordova, A. Jaimes and G. Montiel coll.; 3 Ƥ (CNAC 006636- CNAC 006638), 1 3 (CNAC 006639), Oaxaca, Mexico, 8 km SE San Miguel Talea de Castro, Municipality San Miguel Talea de Castro, (17 ° 19.620 ' N, 96 ° 17.403 ' W), 2,082 m, 22 July 2007, evergreen cloud forest, from three specimens of Proculejus brevis, O. Francke, A. Valdez, C. Santibañez, A. Ballesteros and H. Montaño coll. Holotype, ten paratype females and three paratype males deposited in the CNAC, one female and male paratype each deposited in NMNH, UGA and UMMZ. Female (n = 14). Body oval. Idiosoma, L = 1229 (1165–1517); W = 909 (832–1107) (Fig. 16 A). Ve n tr al idiosoma. Hyaline hood extending beyond to level of coxa IV, seta a 1 long and barbed (Fig. 18 C), L = 93 (56–96). Tritosternum base wider than long. Tetartosternum shield, L = 47 (40–53), anterior W = 155 (133–155), shagreened (Fig. 17 F), anterior margin slightly rounded, corners of shield concave, medial notch small, square. Sternal seta st 1 long and barbed (Fig. 17 E), L = 47 (40–61), near centre of shield; lyriform sternal pore stp 1 close to posterior margin of shield. Sternal shield L = 109 (93–115), shagreened. Pore stp 2 near centre of shield, behind and below it st 2, moderately long and serrated, L = 26 (12–26). Setae st 3 moderately long, serrated, L = 19 (19–25), st 4 is serrated and short, L = 9 (9–16) (Fig. 18 B). Setae st 3 and st 4 on sides of shield, close to posterior margin. Sternogynial shield, L = 90 (87–102), W = 233 (217–236) triangular, posterior margin pointed at apex (Fig. 17 G), surface shagreened, stp 3 near anterior margin. Sternogynial apodeme short and thick, extending posteriorly to level of latigynial and mesogynial condyles. Latigynial shields, L = 186 (174–186), W= 93 (87–93) each with an oval pore and six to ten serrated setae, usually eight; medial margins of shields straight, surface shagreened, latigynial apodeme long and thin. Mesogynial shield, L = 74 (71–90), W = 121 (112–140), triangular, wider posteriorly and anteriorly (Fig. 17 G), slightly pointed at apex, surface shagreened, mesogynial condyles at level of shield apex. Ventral shield, L = 285 (264–304), posterior W = 627 (595–678), triangular, truncated posteriorly, surface shagreened and reticulated like a honey-comb, with 11–14 pairs of small setae (Fig. 16 A). One pore underneath posterior margin of coxa IV. Metapodal-peritremal-exopodal shield reticulated, with a pair of pores and three setae. Anal shield, L = 192 (192–242), anterior W = 576 (531–608) (Fig. 18 D), with five to seven pairs of simple setae plus longer antero anal seta, L = 27 (27–45), two pairs of pores, one near anterior margin and the other below the anus. Shield reticulated like ventral shield (Fig. 18 D). Gnathosoma. Capitular setae (sc) medium and barbed, L = 19 (19–25). Hypostomal setae (Fig. 18 A) hyp 1 long and simple, L = 71 (71–87), longest; hyp 2 long and barbed, 59 (57–62); hyp 3 moderately long and barbed, L = 20 (16–20). Chelicera with barbed seta, movable digit with four teeth, fixed digit with six teeth, movable finger divided at tip. Palp setae are simple, except al 1 and al 2 on trochanter are branched and serrate respectively. Femur setae pl 1 barbed, pl 2 and pd 1 serrated; genu seta al 1 barbed. Anterior margin of palp trochanter with a small blunt process (b) and denticulated process (s) (Fig. 18 E). Legs. Most leg setae simple, except: Leg I: trochanter ad 1, av 1, al 1 and pv 1 pilose; genu ad 1, av 1, pl 2 and pv 1 pilose; tibia ad 1, ad 2, ad 3, av 1, av 2, pd 1, pv 2 and pd 3 pilose. Leg II: trochanter ad 1, ad 2, pd 1 and pd 2 serrated; femur av 1, pv 1, pd 1, pd 2 and pd 3 serrated; genu av 1 and pv 1 serrated; tibia ad 3, al 3, pd 3 and pl 3 serrated; tarsus ad 1, ad 2, ad 3, pd 2, pd 3 and pl 2 serrated, av 2 and av 3 slightly serrated. Leg III: trochanter ad 1, ad 2, pd 1, pd 2 and pv 1 serrated; femur ad 1, ad 2, av 1, pd 1 and pl 1 serrated; genu av 1, pl 1 and pd 2 serrated; tibia pd 3 serrated; tarsus av 3 serrated. Leg IV: trochanter pv 1 and pl 1 serrated; femur av 1 serrated; genu av 1, pl 1 and pv 1 serrated; tibia pd 3 and pl 3 barbed. Seta pv 1 is serrated and long, 36 (31–47) (Fig. 17 H). Male (n = 6) Body oval, similar to female. Idiosoma, L = 1242–1448, W = 954–1037 (Fig. 16 B). Ve n tr al idiosoma. Tetartosternum, L = 59–62, anterior W = 149–161, shagreened and reticulated, anterior margin round and middle of shield is compressed (Fig. 18 F) Tetartosternal notch is small and divides anterior margin in two parts, with a pore near posterior margin. Sternal seta st 1, L = 17–25, simple (Fig. 18 F). Sternal shield (Fig. 18 G) shagreened and reticulated, L = 180–195. Sternal setae, st 2 simple, L = 14, st 3 simple, L= 9– 12), st 4 is short and simple, L = 6, (these setae were found in only two specimens); with two pores, one close to anterior margin of shield and one close to genital opening. Genital opening oval, L = 81–87, W= 121–133 (Fig. 18 G). Ventral shield, L = 422–481, posterior W = 608–672, shagreened and reticulated like a honeycomb, with 34–36 setae; with two pores, one at level of anterior margin of coxa IV and other below level of coxa IV. Anal shield shagreened and reticulated, L = 205–264, anterior W = 544–589; with six pairs of simple setae plus longer antero anal seta, L = 42–48, with two pores, one near anterior margin and the other below anus. Metapodal-peritremal-exopodal shield reticulated with a pair of pores, without seta (Fig. 16 B). Gnathosoma. Hypostomal seta hyp 1 long and simple, L = 80–87; hyp 2 barbed and long, L = 50–68, hyp 3 barbed and medium length, L = 17–25. Etymology. This species is named in honour of Carlos Santibañez, who helped to collect almost all the passalid beetles used in this research. Remarks. This species can be distinguished from all others because the body is very large and setae st 1 is long and barbed, the longest of the sternal setae. In females the tetartosternal, sternal, mesogynial and latigynial shields are shagreened, and only the ventral and anal shield are reticulated. The latigynial shield has six to ten serrated setae. The mesogynial shield has broad posterior and anterior margins. The male has st 1 medium length and simple, and the tetartosternal, sternal, ventral and anal shields are reticulated and shagreened. The tetartosternal notch is small and divides the anterior margin into two parts. The ventral shield has 34–36 setae. Klinckowsroemia santibanezi is very similar is similar to K. candidoi, because they both have the hyaline hood beyond coxa IV, st 1 long and barbed, and the ventral and anal shields reticulated like honey-comb. But new the species has all the sternal setae barbed, with six to ten serrated setae on the latigynial shield, and st 3 and st 4 are serrated. In contrast, K. candidoi has three to four simple setae on the latigynial shield and sternal setae st 3 and st 4 are simple. The beetles carrying this species were found in decaying trunks in three different localities and hosts from Oaxaca. Two mites were found on the humerus and metasternum of the hosts, but almost all specimens were found in the alcohol. One passalid beetle had two species of mites, K. santibanezi and K. victoriae.

Klinckowstroemia hunteri sp. nov. Material examined. Holotype. �� (CNAC 006530), Hidalgo, Mexico, 2 km Mojonera-Tizapan, municipality Zacualtipan (20 �� 38.107 ' N, 98 �� 35.981 ' W), 2,019 m, 22 September 2006, from Odontotaenius zodiacus, O. Francke, A. Valdez and H. Monta��o coll., pine-oak forest. Paratypes. 2 �� (CNAC 006531, CNAC 006532), 5 33 (CNAC 006533- CNAC 006537), same data as holotype, from 3 specimens of Odontotaenius zodiacus; 2 ��, Hidalgo, Mexico, 2 km E Acaxochitl��n (20 ��08.500' N, 98 �� 10.740 ' W), 2,214m, 23 September 2006, from Odontotaenius zodiacus, O. Francke, A. Valdez and H. Monta��o coll. pine-oak forest. Holotype female, two paratype females and four paratype males deposited in the CNAC, one female and one male paratypes deposited in NMNH. Female (n = 13). Body oval. Idiosoma, L = 966 (896���1,043); W = 666 (640���672) (Fig. 3 A). Ve n tr al idiosoma. Hyaline hood extending to level of coxa I, seta a 1 large and barbed, L = 42 (31���43). Tritosternum base wider than long. Tetartosternum L = 40 (34���40), anterior W = 124 (124���127), shagreened and reticulated as illustrated (Fig. 4 C). Tetartosternal notch triangular. Sternal seta st 1 simple (Fig. 4 D), L = 6, but broken on all specimens, lyriform sternal pore stp 1 near posterior margin. Sternal shield (Fig. 4 E) L = 87 (59���90), stp 2 near centre of shield, behind it sternal seta st 2, a simple microseta. Setae st 3, L = 20 (12���22) and st 4, L = 7 (6���8) near posterior margin of shield, both simple, close together. Shield reticulated and shagreened. Sternogynial shield (Fig. 4 G) L = 65 (62���68); W = 170 (161���174) triangular, anterior margin narrow and pointed at apex, surface shagreened and reticulated, stp 3 located closer to lateral margins. Sternogynial apodeme with broad base that rises to the shield, posterior arms thick, extending posteriorly to level of latigynial and mesogynial condyles. Latigynial shields, L = 143 (121���143); W = 68 (47���76), each with an oval pore and four setae, medial margin of shield straight and short (Fig. 4 G), shield reticulated, surface shagreened; latigynial apodeme thick and separate from sternogynial shield. Mesogynial shield L = 81 (53��� 81), W = 84 (71���93), triangular, wider posteriorly and pointed at apex, anterior margin extending slightly anteriorly to sternogynal shield (Fig. 4 G), surface shagreened. Mesogynial condyles extending almost to level of shield apex. Ventral shield, L = 279 (251���282); posterior W = 544 (518���544), shield triangular, truncated posteriorly, surface shagreened (Fig. 6 B), with 12 pairs of small setae (Fig. 3 A). One pore underneath posterior margin of coxa IV (Fig. 3 A). Metapodal-peritremal-exopodal shield (MPES) with a pair of pores and one simple seta (Fig. 3 A, 6 B). Anal shield, L = 192 (102���198); anterior W = 474 (448���474) (Fig. 4 F), with six pairs of simple setae plus longer antero anal seta, L = 34 (32���34.1), two pairs of pores, one near anterior margin and the other below the anus. Gnathosoma. Capitular setae (sc) short and serrate, L = 20 (14���20). With three hypostomal setae (Fig. 4 H), hyp 1 long and simple, L = 65 (56���68), longest. Seta hyp 2 serrate, L = 40 (40���45). Seta hyp 3 short and simple, L = 12 (12���16). Chelicera with barbed setae, movable digit with four teeth, two small and two large; fixed digit with six teeth, two large and four small. Most palpal setae simple, but av 1 and av 2 of trochanter are branched and serrate respectively. On femur pv 1 is barbed; ad 1, ad 1, pd 1 and pd 2 are serrated. On genu ad 1 and ad 2 are serrate. Anterior margin of palp trochanter with blunt process (b), denticulated process (s) slightly longer (Fig. 6 A). Legs. Most leg setae simple, except: Leg I: trochanter I ad 1, av 1, al 1 and pv 1 serrated; femur I, al 1, pv 1 and pv 3 serrated; genu I, av 1 and pl 1 serrated. Leg II: femur av 1 and genu av 1 serrated. Leg III: femur av 1; genu av 1 and pv 1; tibia ad 3 and al 3 serrated. Leg IV: coxa av 1 and pv 1, tibia ad 4, tarsus ad 2 and ad 3 slightly serrated. Male (n = 4) Body oval, similar to female. Idiosoma, L = 992���1018, W = 717���755 (Fig. 3 B). Ve n tr al idiosoma. Tetartosternum, L = 47���56, anterior W = 124���133, shagreened, with one line near anterior margin (Fig. 6 D). Tetartosternal notch small, square, moderately deep (Fig. 6 D). Sternal seta st 1, L = 9���22, simple (Fig. 6 C). Sternal seta st 2, L = 12, simple, seta st 3, L = 6���11, simple, and st 4 is a microseta. Sternal shield shagreened, L = 153. Genital opening oval, L = 59���65, W = 89���96. Ventral shield with 36 setae, L = 372���388, shagreened, with two pores, one near anterior margin of coxa IV and other below the same coxa (Fig. 3 B). Anal shield, L = 183���217, with six pairs of simple setae, plus longer antero anal seta L = 12���34, with two pores: one near to anterior margin and the other below the anus. Gnathosoma. Hypostomal seta hyp 1 long and simple, L = 65���71, seta hyp 3 serrate and long, L = 43���47, seta hyp 2 simple and shorter, L = 12���16. Legs. Seta pv 1 on coxa IV slightly serrated (Fig. 6 E). Etymology. This species is named in honour of Preston E. Hunter, for his contribution to the study of mites associated with passalids, especially of the family Klinckowstroemiidae. Remarks. This species can be distinguished from all others because the latigynial and sternogynial apodemes are strongly developed and wider, separating the sternogynial and latigynial shields. The mesogynial shield has a wide posterior margin. The tetartosternum, sternal, sternogynial and latyginal shields are reticulated and shagreened. The male has 34 setae on the ventral shield, and the tetartosternum is hexagonal in shape and expanded. Klinckowstroemia hunteri is similar to K. atramaculata, because they have the latigynial shield long and st 1 barbed, but the last species lacks the well-developed latigynial and mesogynial apodemes, has three setae on latigynial, and seta a 1 is simple, while, K. hunteri has four setae on latigynial shield and seta a 1 long and barded. The male of K. atramaculata has 13���15 pair of setae, while the male of K. hunteri with 17 pairs. The species K. bifurcata and K. hunteri were found together on three specimens of Odontotaenius zodiacus. It is possible that these passalids lived in the same log, because they share the same klinckowstroemiids and they were collected at the same locality on the same day. One of these passalid had three different species of the genus Klinckowstroemia. Furthermore seven (5 ��, 2 33) specimens of K. hunteri were collected together with one female of K. victoriae on the same species of passalid., Published as part of Villegas-Guzman, Gabriel A., P��rez, Tila M. & Reyes-Castillo, Pedro, 2009, New species of the genus Klinckowstroemia Baker & Wharton from Mexico (Acari: Mesostigmata: Trigynaspida: Klinckowstroemiidae), pp. 1-46 in Zootaxa 2248 on pages 8-11, DOI: 10.5281/zenodo.190717

Klinckowstroemia rectimarginata sp. nov. Material examined. Holotype. Ƥ (CNAC 006597), Veracruz, Mexico, Dos Amates, Municipality Catemaco (18 ° 29.35 ’ N, 95 °03.58’ W), 120 m, 15 November 1987, from Paxillus leachi MacLeay, P. Hunter coll. Paratypes. 2 Ƥ (CNAC 006598, CNAC 006599), 2 33 (CNAC 006600, CNAC 006601), Veracruz, Mexico, Coyame Bastonal, Municipality Catemaco (18 ° 23.25 ’ N, 94 ° 56.36 ’ W), 14 November 1987, from Heliscus tropicus, P. Hunter coll.; 1 3 (CNAC 006602), Veracruz, Mexico, Coyame Bastonal, Municipality Catemaco (18 ° 23.25 ’ N, 94 ° 56.36 ’ W), 830–970 m, 14 November 1987, from Ve t u r i u s marilucae Boucher, P. Hunter coll. Holotype, two paratype females and three paratype males deposited in the CNAC. Female (n = 4). Body oval. Idiosoma, L = 864 (813–902), W = 634 (614–691) (Fig. 12 A). Ve n tr al idiosoma. Hyaline hood extending to level of coxa I, seta a 1 moderately long and simple, L = 22 (19–25). Tritosternum base wider than long. Tetartosternum shield (Tst), L = 34 (34–40), anterior W= 115 (105–115); shagreened, anterior margin with two lines (Fig. 13 E), tetartosternal notch like a small cavity in centre of margin. Seta st 1 near shield margin, medium and slightly pilose (Fig. 13 F), L = 25 (19–29), lyriform pore stp 1 near posterior margin. Sternal shield (StS) L = 71 (68–71), shagreened. Pore stp 2 near centre of shield, seta st 2 below and behind it, a simple microseta. Sete st 3, L = 7.9 0.9 (7.4–9.3) and st 4, L = 7 (7–9) near posterior margin of the shield, both small and simple (Fig. 12 A). Sternogynial shield (SgS), L = 53 (53– 56), W = 146 (146–171) triangular, anterior margin slightly pointed at apex, surface shagreened, stp 3 near anterior margin (Fig. 13 G). Sternogynial apodeme long and thin, extending posteriorly to level of latigynial and mesogynial condyles. Latigynial shields, L = 127 (124–130), W = 71 (71– 74) each with an oval pore and three setae, medial margins of shield medium and straight, surface shagreened; latigynial apodeme thin. Mesogynial shield, L = 74 (68–81), W = 99 (93–102), triangular, pointed at apex and slightly wider posteriorly, surface shagreened. Mesogynial condyles behind level of shield apex. Ventral shield, L = 202 (202–223), posterior W = 461 (448–512), shield triangular, truncated posteriorly, surface shagreened, with 12 pairs of small setae (Fig. 12 A). One pore underneath posterior margin of coxa IV. Metapodal-peritremalexopodal shield with a pair of pores and two setae, reticulated. Anal shield, L = 140 (118–146); anterior W = 397 (384–429), with five pairs of simple setae plus longer antero anal seta, aa, L = 19 (19–25), three pairs of pores, two near anterior margin, the other below the anus (Fig. 12 A). Gnathosoma. Capitular setae (sc) short and serrated, 9 (9–12). Hypostomal setae (Fig. 13 D), hyp 1 long and simple, L = 53 (53–56), longest. Seta hyp 2 moderate and barbed, L = 34 (28–34). Seta hyp 3 short and simple, L = 12. Chelicera with barbed seta, movable digit with five teeth, fixed digit with six teeth. Palp setae simple, except al 1 (av 1) and al 2 (av 2) on trochanter branched and serrate respectively. Femur setae av 1, pl 1 and pd 1, serrated, pd 2 pilose. Genu setae pl 1, pl 2, pd 1, pd 2 and av 1 serrated. Anterior margin of palp trochanter with blunt process (b) and denticulated process (s) (Fig. 13 H). Legs. Most leg setae simple, except: Leg I: trochanter I al 1 and pv 1 serrated; femur I pv 1 and pv 3 serrated; genu I pl 1 and pv 1 slightly serrated; tibia I ad 1, ad 2, ad 3, pd 1 and pd 3 serrated. Leg II: trochanter II pv 2 serrated; femur II av 1 and pv 1 slightly serrated; genu II av 1 and pv 1 slightly serrated. Leg III: trochanter pd 1 and pd 2 pilose; femur av 1 slightly serrated; genu av 1, pl 1 and pv 1 barbed; tarsus ad 3 serated. Leg IV: trochanter pv 1 pilose; femur av 1 serrated; genu pv 1 serrated. Seta pv 1 is large and slightly serrated, L = 22 (22–25) (Fig. 15 A). Male (n = 3) Body oval similar, to female. Idiosoma, L = 845–890, W = 614–685 (Fig. 12 B). Ve n tr al idiosoma. Tetartosternum, L = 34–37, anterior W = 109–118, shagreened and reticulated, with four horizontal lines, anterior margin reduced medially but without a notch (Fig. 15 B), pore stp 1 near posterior margin. Sternal seta st 1, L = 6–17, simple (Fig. 15 C). Sternal shield shagreened and reticulated, L = 130–139. Setae, st 2, st 3 and st 4, are simple microsetae; with two pores, one near anterior margin of shield, one close to genital opening (Fig. 15 D). Genital opening oval, L = 53–59, W = 71–81. Ventral shield, L = 301– 316, posterior W = 461–512, shagreened, with 38 setae and two pores, one at level of anterior margin of coxa IV and other below coxa IV. Anal shield shagreened, L = 130–140, anterior W = 384–416, with five pairs of simple setae plus longer antero anal seta, L = 25–28, and two pores, one near anterior margin and one below the anus. Metapodal-peritremal-exopodal shield reticulated, with a pair of pores and two setae (Fig. 12 B). Gnathosoma. Hypostomal seta hyp 1 long and simple, L = 58–59, hyp 2 barbed and intermediate in length, L = 34–40, hyp 3 simple and shorter, L = 7–12. Etymology. This name is from Latin rectus (= straight) and marginata (= margin) referring to the shape of the anterior and posterior margins of the tetartosternal shield. Remarks. This species can be distinguished from all others because the tetartosternal shield is rectangular; the tetartosternal notch is very small in the female and absent in the male. The female has a large genital shield. The sternoventral suture is near the anterior margin of the anal shield. Seta st 1 is long and serrated in the female, and medium length and simple in the male. In the female sternal seta st 2 is a microseta and st 3 and st 4 are short and simple; while in male all sternal setae are microsetae. Seta al 2 (av 2) is large and slightly serrated on both sexes. The male has four horizontal lines on the tetartosternal shield, the sternal shield is reticulated and shagreened, and the ventral shield is shagreened, with 19 pairs of setae. Klinckowstroemia rectimarginata is similar to K. schusteri and K. brevimarginata because they have st 1 and hyp 2 serrated and barbed respectively. In the new species the medial margins of the latigynial shields are straight and reduced (16–22), which is similar to K. brevimarginata; seta st 1 are of medium length (19–29) and serrate but slightly longer in K. rectimarginata than K. brevimarginata. In both species the other sternal setae are simple microsetae, while in K. schusteri the setae are bigger, st 2 and st 3 are simple and st 4 is slightly serrated. The sternal, tetartosternal, ventral and genital shields are shagreened; these are similar in K. brevimarginata and K. rectimarginata, while in K. schusteri they are reticulated. There are four setae on the latigynial shield in K. schusteri, four to six in K. brevimarginata and three in K. rectimarginata. Males present more differences; the tetartosternal shield is reticulated and its anterior margin reduced, and the tetartosternal notch is slightly evident. The tetartosternal, sternal and ventral shields are reticulated and shagreened. The specimens were found in decaying trunks in the same locality from Veracruz, but on three different hosts.

Klinckowstroemia montanoi sp. nov. Material examined. Holotype. Ƥ (CNAC 006538), Oaxaca, Mexico, 8 km NW San Miguel Coatlán, Federal highway Cuixtla-San Miguel Coatlán (16 ° 15.064 ' N, 96 ° 41.629 ' W), 2,170 m, 24 June 2006, from Verres aff. intermedius, O. Francke, A. Valdez and H. Montaño coll., oak forest. Paratypes. 14 Ƥ (CNAC 006539- CNAC 006552), 5 33 (CNAC 006553- CNAC 006557), same data as holotype, from four specimens of Verres aff. intermedius. Holotype, 10 paratype female and four paratype males deposited in the CNAC, one female and one male paratype deposited in NMNH; one female paratype each deposited in UGA and UMMZ. Female (n = 15). Body oval. Idiosoma, L = 742 (704–755), W = 512 (493–538) (Fig. 5 A). Ve n tr al idiosoma. Hyaline hood extending to level of coxa I, seta a 1 moderate and simple, L = 19 (16–18). Tritosternum base wider than long. Tetartosternum (Tst) L = 43 (40–47), anterior W = 93 (87–93), shagreened (Fig. 6 G), with a line near anterior margin, lyriform sternal pore stp 1 near posterior margin. Tetartosternal notch square-ish, sternal seta st 1, L = 22 (21–29), serrate (Fig. 8 F). Sternal shield (Fig. 6 H) L = 50 (43–56), stp 2 near centre of shield, behind it sternal seta st 2, short and simple, L = 6. Setae st 3, L = 6, simple, and st 4, a microseta, near posterior margin of shield. Shield reticulated and shagreened (Fig. 6 H). Sternogynial shield, L = 43 (37–47), W = 152 (143–155), triangular, posterior margin narrow and pointed at apex, surface shagreened and reticulated (Fig. 8 A), stp 3 near anterior margin. Sternogynial apodeme short and thick, extending to posterior level of latigynial and mesogynial condyles. Latigynial shields, L = 99 (96–102); W = 65 (65– 68) each with an oval pore and five to eight setae, medial margins of shields wavy and short (Fig. 8 A), shields reticulated and surface shagreened; latigynial apodeme thin. Mesogynial shield, L = 62 (59–68); W = 74 (71–77), triangular, wider posteriorly and pointed at apex, surface shagreened. Mesogynial condyles in centre of shield. Ventral shield, L = 211 (195–220); posterior W = 422 (416–435), shield triangular (Fig. 8 D), truncated posteriorly, surface shagreened and reticulated, with 13–14 pairs of small setae (Fig. 5 A). One pore underneath posterior margin of coxa IV. Metapodal-peritremal-exopodal shield with a pair of pores and one simple seta. Anal shield (Fig. 8 E), L = 124 (105–140); anterior W = 352 (320–365), with five pairs of simple setae plus longer anteroanal seta, L = 25 (20–25), two pairs of pores, one near anterior margin and the other below anus. Gnathosoma. Capitular setae (sc) short and serrate, L = 11 (8–12). With three hypostomal setae (Fig. 6 F), hyp 1 long and simple, L = 46 (43–50), longest. Seta hyp 2 medium and serrate, L = 29 (26–32). Seta hyp 3 short and serrate, L = 15 (14–17). Chelicera with barbed seta, movable digit bifurcate at tip, with four teeth; fixed digit with five teeth. Palp setae generally simple, except al 1 and al 2 on trochanter branched and serrate respectively. Femur setae av 1, pl 1, pl 2, pl 3 and pd 1 serrate. Genu setae av 1, pl 1, pl 2, pl 3, pd 1 and pd 2 serrate. Anterior margin of palp trochanter with blunt process (b), denticulated process divided (s) (Fig. 8 B). Legs. Leg setae mostly simple except: Leg I: trochanter I ad 1 and pv 1 serrate; femur I ad 1, ad 2, pv 1 and pv 3 serrate; genu I av 1 and pv 1 serrate. Leg II: femu av 1 and genu av 1 serrate. Leg III: femur av 1; genu III av 1 and pv 1 serrate; trochanter IV pv 1, femur av 1; genu av 1 and pv 1 serrate. Seta pv 1 (Fig. 8 C) is large and serrate, L = 37 (34–38). Male (n = 5) Body oval, similar to female. Idiosoma, L = 704–755, W = 474–518 (Fig. 5 B). Ve n tr al idiosoma. Tetartosternum, L = 40–56, anterior W = 93–124, shagreened. Tetartosternal notch deep, V-shaped. Sternal seta st 1, L = 20–22, simple. Setae st 2, st 3 and st 4 are simple microsetae. Sternal shield shagreened and reticulated (Fig. 5 B), L = 78–81. Genital opening oval, L = 43–47, W = 71–78. Ventral shield, L = 279–285, posterior W = 384–403, shagreened, with 24–26 setae (Fig. 5 B), with two pores, one at level of coxa III and other below posterior margin of coxa IV. Anal shield, L = 124–127, anterior W = 333–352, with five pairs of simple setae plus longer antero anal seta, L = 24–25, with two pores, one near anterior margin and the other below anus. Gnathosoma. Hypostomal seta hyp 1 long and simple, L = 47–50, hyp 3 serrate, L = 31–34, hyp 2 simple and shorter, L = 16–25. Etymology. This species is named in honour of Héctor Montaño, who helped to collect almost all the passalid beetles used in this research. Remarks. This species can be distinguished from all others because st 4 is a microseta, st 2 and st 3 are short and simple, st 1 is moderate and serrate and the medial margin of the latigynial shield is wavy. The sternal, latigynial and sternogynial genital shields are shagreened and reticulated, the tetartosternum is only shagreened. The mesogynial shield is wider in the anterior and posterior margins. In the male the tetartosternal notch is deep and distinct, V-shaped, almost reaching half the length of the shield, and there are 24–26 pairs of setae on the ventral shield. Klinckowstemia montanoi is similar to K. scotti and K. valdezi because all the species have wavy medial margins on the latigynial shield, st 1 serrated, and number of setae on anal shield is equal. Klinckowstroemia montanoi (735.6) is longer than K. valdezi (685.9), but shorter than K. scotti (1292). The latigynial shield has eight pilose setae in K. scotti, while K. montanoi have five to eight simple setae simple and K. valdezi has three simple setae. K. montanoi has 13–14 pairs setae on the ventral shield, while K. valdezi and K. scotti have 12 pairs. The specimens were found in decaying trunks only in one locality in the state of Oaxaca, Mexico, on the same host, Ve r re s aff. intermedius. Mites were found in the alcohol, not on the beetles.

Klinckowstroemia ballesterosi sp. nov. Material examined. Holotype. Ƥ (CNAC 006640), Veracruz, Mexico, Municipality of Soledad Atzompa, Soledad Atzompa (18 ° 44.712 ' N, 97 ° 40.967 ' W) 2,125 m, 23 March 2007, from Odontotaenius zodiacus, pine-oak forest, O. Francke, A. Valdez, C. Santibañez, A. Ballesteros and H. Montaño coll. Paratypes. 3 Ƥ (CNAC 006641- CNAC 006643), 1 3 (CNAC 006644), same data as holotype, from three specimen of Odontotaenius zodiacus; 2 Ƥ (CNAC 006645, CNAC 006646), Veracruz, Mexico, Tlacolulan, Municipality of Tlacolulan (19 ° 38.508 ' N, 96 ° 59.650 ' W), 1,876 m, 30 September 2006, from one specimen of Odontotaenius zodiacus, oak forest, O. Francke, C. Santibañez, A. Valdez and H. Montaño coll.; 2 Ƥ (CNAC 006647, CNAC 006648), Oaxaca, Mexico, 8 km SE San Miguel Talea de Castro, Municipality of San Miguel Talea de Castro (17 ° 19.620 ' N, 96 ° 17.403 ' W), 2,082 m, 22 July 2007, evergreen cloud forest, from one specimen of Petrejoides recticornis, O. Francke, A. Valdez, C. Santibañez, A. Ballesteros and H. Montaño coll. Holotype, four paratype females and one paratype male deposited in the CNAC, one female paratype each deposited in NMNH, UGA and UMMZ. Female (n = 8). Body oval. Idiosoma, L = 992 (896–1088), W = 742 (640–774) (Fig. 19 A). Ve n tr al idiosoma. Hyaline hood extending to level of coxa I, seta a 1 long and barbed, L = 47 (37–50). Tritosternum base wider than long. Tetartosternum shield, L = 43 (37–43), anterior W = 121 (112–127), shagreened and reticulated (Fig. 18 H), anterior margin with two small protuberances, corners of shield pointed, medial notch small, square. Sternal seta st 1 long and serrated (Fig. 20 A), L = 57 (47–68), near centre of shield; lyriform pore stp 1 close to posterior margin. Sternal shield (Fig. 20 B), L = 87 (74–90) shagreened and reticulated (Fig. 20 C). Pore stp 2 near anterior margin of shield, behind and below it seta st 2, long and serrated, L = 50 (37–54). Setae st 3 long and serrated, L = 47 (35–59), st 4 serrated and moderately long, L = 37 (31–47) (Fig. 20 B). Setae st 3, st 4 on sides of shield, close to posterior margin. Sternogynial shield, L = 50 (47–62), W = 186 (167–189) triangular (Fig. 20 D), posterior margin pointed at apex, surface shagreened, with stp 3 near anterior margin. Sternogynial apodeme long and thick, extending posteriorly to level of latigynial and mesogynial condyles. Latigynial shields, L = 130 (124–140), W= 84 (71–84) (Fig. 20 D) each with an oval pore and four or five simple setae, usually four; medial margins of shields are straight, shields shagreened; latigynial apodeme thin and long. Mesogynial shield, L = 87 (81–96); W = 112 (99–115), triangular (Fig. 20 D), wider posteriorly and anteriorly, apex pointed, surface shagreened, mesogynial condyles below level of shield apex. Ventral shield, L = 248 (223–273), posterior W = 576 (512–627), triangular, truncate posteriorly, surface shagreened, with 14–15 pairs of small setae (Fig. 20 G). One pore underneath posterior margin of coxa IV. Metapodal-peritremal-exopodal shield reticulated, with a pair of pores and two setae. Anal shield, L = 195 (149–198), anterior W = 524 (448–544), shield shagreened (Fig. 19 A), with five pairs of simple setae plus longer antero anal seta, L = 31 (29.– 35), two pairs of pores, one near anterior margin, another below the anus. Gnathosoma. Capitular setae (sc) small and serrated, L = 7 (7–18). Hypostomal setae hyp 1 long and simple, L = 68 (59–68), longest; hyp 2 long and barbed, 45 (39–47); hyp 3 medium length and barbed, L = 22 (12–22). Chelicera with barbed seta, movable digit with four teeth, two large and two small; fixed digit with six teeth, two large and four small, movable digit divided at the tip. Palp setae are simple, except al 1 and al 2 on trochanter are branched and serrate respectively. Femur setae pl 1, pl 2 and pv 1 serrated, avl 1 barbed; genu seta ad 1 and pl 2 are barbed, pv 1 and ad 2 are serrated. Anterior margin of palp trochanter with a small blunt process (b) and denticulate process (s) (Fig. 20 F). Legs. Most leg setae simple, except Leg I: trochanter ad 1, al 1 and pv 1 barbed; femur pv 1, pv 2 and pv 3 serrated; genu al 1, pl 1 and pv 1 serrated; tibia ad 1, ad 2, ad 3, pd 1 and pd 3 serrated. Leg II: trochanter ad 1 serrated; femur: av 1 and pl 1 serrated; genu: av 1 and pv 1 serrated. Leg III: trochanter pv 1 serrated; femur av 1 and pl 1 serrated; genu av 1 and pv 1 serrated. Leg IV: trochanter pv 1 and p 12 serrated; femur av 1 serrated; genu av 1 and pv 1 serrated. Seta pv 1 is slightly serrated and long, 34 (29–35) (Fig. 20 E). Male (n = 3) Body oval, similar to female. Idiosoma, L = 1005–1018, W = 723–768 (Fig. 19 B). Ve n tr al idiosoma. Tetartosternum, L = 43, anterior W = 118–124, shagreened (Fig. 20 H), anterior margin straight, corners pointed as in female. Tetartosternal notch small, V-shaped, divides anterior margin of shield in two parts, with a pore near posterior margin. Sternal seta st 1, L = 9–16, simple (Fig. 20 H). Sternal shield shagreened (Fig. 22 A), L = 140–149. Sternal setae st 2 a simple microseta, st 3 long and serrated, L = 34–43, st 4 is long and serrated, L = 40–43; shield with two pores, one near anterior margin and one near posterior margin. Genital opening oval, L = 53–59, W= 78–84. Ventral shield L = 307–390, posterior W = 576–640, shagreened, with 34 setae (Fig. 22 B); with two pores, one near anterior margin of coxa IV and one below coxa IV. Anal shield shagreened, L = 180–202, anterior W = 512–563; with five pairs of simple setae plus longer antero anal seta, L = 31–34, and two pores, one near anterior margin and one below anus. Metapodalperitremal-exopodal shield reticulated, with a pair of pores and one seta. Gnathosoma. Hypostomal seta hyp 1 long and simple, L = 68–71, hyp 2 simple and long, L = 42–43, hyp 3 is simple and medium length, L = 14–16. Etymology. This species is named in honour of Alfonso Ballesteros, who helped to collect almost all the passalid beetles used in this research. Remarks. This species can be distinguished from all others because the sternal setae are long and serrated, and seta st 1 is the longest. In females the tetartosternal and sternal shields are shagreened and reticulated, and the other shields are only shagreened. The latigynial shield with has four or five simple setae. The mesogynial shield is broad posteriorly, and the form of the tetartosternal shield is very different to others species. In the male st 3 and st 4 long and serrated, and the tetartosternal, sternal, ventral and anal shields are shagreened. The tetartosternal notch is small, V-shaped, and divides the anterior margin in two parts. The ventral shield has 17 pairs of setae. Klinckowstroemia ballesterosi is similar to K. santibanezi and K. candidoi because they have setae a 1 and st 1 long and barbed, but the new species is smaller (896–1088 versus 1164– 1517 and ca. 1290 respectively), and all sternal setae are serrated and long, with fourto five simple setae on the latigynial shield. In contrast, in K. santibanezi the sternal setae are barbed and serrated, and there are eight serrated setae on the latigynial shield; and in K. candidoi the sternal setae are serrated and simple and there are three to four simple setae on the latigynial shield. Klinckowstroemia ballesterosi has more setae on the ventral shield (K. ballesterosi 14, K. santibanezi 11–14, K. candidoi 11). In K. ballesterosi the tetartosternum and sternal shield are reticulated, while in K. santibanezi and K. candidoi the ventral and anal shields are reticulated. The beetles carrying this species were found in decaying tree trunks in two different states of Mexico. In Veracruz they were found on Odontotaenius zodiacus and almost all specimens were found in the alcohol, with only one specimen on the humerus of one of the beetles. From Oaxaca it was found with Petrejoides recticornis free in the alcohol.