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See discussions, stats, and author profiles for this publication at: https://www.researchgate.net/publication/262465763 Coadaptación entre los ácaros (Arachnida: Klinckowstroemiidae) y coleópteros Passalidae (Insecta: Coleoptera) ARTICLE in REVISTA DE BIOLOGIA TROPICAL · JUNE 2012 Impact Factor: 0.52 · DOI: 10.15517/rbt.v60i2.3941 CITATION 1 READS 25 4 AUTHORS, INCLUDING: Oscar Francke Universidad Nacional Autónoma de México 88 PUBLICATIONS 408 CITATIONS SEE PROFILE Pedro Reyes-Castillo Institute of Ecology INECOL 46 PUBLICATIONS 100 CITATIONS SEE PROFILE Available from: Pedro Reyes-Castillo Retrieved on: 05 February 2016
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Seediscussions,stats,andauthorprofilesforthispublicationat:https://www.researchgate.net/publication/262465763

Coadaptaciónentrelosácaros(Arachnida:Klinckowstroemiidae)ycoleópterosPassalidae(Insecta:Coleoptera)

ARTICLEinREVISTADEBIOLOGIATROPICAL·JUNE2012

ImpactFactor:0.52·DOI:10.15517/rbt.v60i2.3941

CITATION

1

READS

25

4AUTHORS,INCLUDING:

OscarFrancke

UniversidadNacionalAutónomadeMéxico

88PUBLICATIONS408CITATIONS

SEEPROFILE

PedroReyes-Castillo

InstituteofEcologyINECOL

46PUBLICATIONS100CITATIONS

SEEPROFILE

Availablefrom:PedroReyes-Castillo

Retrievedon:05February2016

Accepted by B. Halliday: 14 Dec. 2010; published: 24 Jan. 2011

ZOOTAXAISSN 1175-5326 (print edition)

ISSN 1175-5334 (online edition)Copyright © 2011 · Magnolia Press

Zootaxa 2747: 19–36 (2011) www.mapress.com/zootaxa/ Article

19

New species of Klinckowstroemia Baker & Wharton, 1952 (Acari: Klinckowstroemiidae), associated with passalid beetles from Mexico and Panama

GABRIEL A. VILLEGAS-GUZMÁN1, PEDRO REYES-CASTILLO2 & TILA M. PÉREZ1 1Colección Nacional de Ácaros, Departamento Zoología, Instituto de Biología, Universidad Nacional Autónoma de México. Apartado Postal 70-153, Ciudad Universitaria, 04510 México, D. F.. E-mail: [email protected], [email protected] 2Instituto de Ecología, A. C. Apartado Postal 63, 91000 Xalapa, Veracruz, México. E-mail: pedro. [email protected]

Abstract

Four new species of the genus Klinckowstroemia are described, all obtained from passalid beetles collected in Mexico andPanama. The species are: Klinckowstroemia pennula, sp. nov., K. melissae sp. nov., K. stilla sp. nov., and K. oconnori sp.nov. We found two instances where different mite species inhabited the same host, K. pennula, and K. melissae on Pseu-dacanthus sp., and K. pennula and K. candidoi on Oileus bifidus. This is the first record of the genus Klinckowstroemiafrom Panama. An updated key for the 24 species of Klinckowstroemia is presented.

Key words: mites, Passalidae, Central America, synhospitality, phoresy

Introduction

Mites of the genus Klinckowstroemia are distributed principally in Mesoamerica, where they occur exclusively onbeetles of the family Passalidae (bessbugs) (Hunter & Butler 1966; Chernoff & Pope 1970; Rosario & Hunter1987; Villegas-Guzman et al. 2009). Eighteen of the 24 species in the genus are found in Mexico (Table 1). Theseare associated with 32 species of passalid beetles belonging to 14 different genera. Four species of klinckowstro-emiids have been found on beetles (bessbugs) from Central America. Following the taxonomic revision of thegenus Klinckowstroemia (Villegas-Guzman et al. 2009), we found more material of four new species associatedwith bessbugs, and they are described below.

Material and methods

Bessbugs were collected in and under decaying logs and put individually in vials with 80% ethanol. The specimenswere taken to the Colección Nacional de Ácaros (CNAC) and mites removed with fine-tipped forceps under a ster-eomicroscope. The klinckowstroemiid mites were separated into vials with 80% ethanol and some of them clearedwith lacto-phenol solution and mounted in Hoyer’s fluid. Passalid beetles are deposited at the Colección Ento-mológica, Instituto de Ecología, Xalapa (IEXA), and that collection was also checked to search for klinckowstro-emiid and other mites.

Slide-mounted specimens were examined with a Nikon microscope using differential contrast and phase con-trast microscopy. Measurements are given in micrometres and are reported in the text as holotype, followed by theminimum and maximum of paratypes values given in parentheses (only one measurement is given where no varia-tion was observed). Abbreviations used in the descriptions are L = length, W = width. Leg chaetotaxy follows See-man (2007). Nomenclature for structures like setae and shields follows Rosario & Hunter (1987). The drawingswere made using a drawing tube, and scanning electron micrographs of gold coated specimens were made using aHitachi S-2460M microscope. The photographs were edited using software Adobe Photoshop version 10. 0. Abbre-viations used in illustrations: hyp1–4 = hypostomal setae 1–4; st1–4 = sternal setae; av1 = first anteroventral seta of

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palp trochanter; av2 = second anteroventral seta of palp trochanter; Sgs = sternogynal shield; Lgs = latigynal shield;Mgs = mesogynal shield; GO = genital opening of male; b = palp trochanter blunt process; s = palp trochanter spinelike or denticulate process; Pst = presternal shield, VS = ventral shield; MPES = metapodal-exopodal-peritrematalshield; Sts = sternal shield, aa = antero-anal seta; stp1–3 sternal pore. Setae less than 5 micrometres long are con-sidered microsetae.

Specimens are deposited in the Colección Nacional de Ácaros, Instituto de Biología, UNAM (CNAC); theNational Insect and Mites Collection, National Museum of Natural History (NMNH) at The Smithsonian Institu-tion; the Georgia Museum of Natural History, University of Georgia (UGA); and the Museum of Zoology, Univer-sity of Michigan (UMMZ).

TABLE 1. Species of the genus Klinckowstroemia associated with passalid beetles. *Species found in Mexico.

Species Hosts Localities

K. atramaculata Rosario & Hunter, 1988 Passalid not identified Puntarenas, Costa Rica

K. ballesterosi* Villegas-Guzman et al., 2009

Odontotaenius zodiacus and Heliscus tropicusPetrejoides recticornis

Soledad Atzompa and Tlacolulan, VeracruzSan Miguel Talea de Castro, Oaxaca

K. bifurcata* Villegas-Guzman et al., 2009 Odontotaenius zodiacus and Proculejus hirtus

Tianguistengo and Zacualtipan, Hidalgo

K. brevimarginata* Villegas-Guzman et al., 2009

Verres aff. intermedius Puerto Escondido, Oaxaca

K. candidoi* Rosario & Hunter, 1988 Oileus herosPseudacanthus sp.

Xicotepec de Juárez, PueblaTlapacoyan, Veracruz

K. concava* Hunter & Butler, 1966 Passalus sp. Verres sp. Veturius sp.

San Vito, Costa RicaArroyo Claro, Veracruz. Bastonal, Veracruz

K. cristinae* Villegas-Guzman et al., 2009 Odontotaenius zodiacus and Oileus rima-tor

Tianguistengo, Hidalgo

K. franckei* Villegas-Guzman et al., 2009 Odontotaenius zodiacus Zacualtipan, Hidalgo

K. grabowskii* Chernoff & Pope, 1970 Undulifer incisusOileus rimator

Cerro San Felipe, OaxacaGuerrero

K. hunteri* Villegas-Guzman et al., 2009 Odontotaenius zodiacus Zacualtipan and Acaxochitlán, Hidalgo

K. melissae*, this paper Pseudacanthus sp. San Pedro-San Pablo Ayutla, Oaxaca

K. montanoi* Villegas-Guzman et al., 2009

Verres aff. intermedius San Miguel Coatlán, Oaxaca

K. multisetillosa Rosario & Hunter, 1988 Proculus mniszechi and P. opacipennis P. burmeisteri

Baja Verapaz, Cerro San Gil, La Unión and Sierra de las Minas, Guate-malaEl Portillo, Honduras

K. oconnori, this paper Odontotaenius striatopunctatus y Petrejoides subrecticornis

Cerro Azul, Panamá

K. pennula*, this paper Pseudacanthus sp. San Pedro-San Pablo Ayutla, Oaxaca

Vindex agnoscendus Santiago Comaltepec, Oaxaca

Oileus bifidus San Miguel Talea de Castro, Oaxaca

K. rectimarginata* Villegas-Guzman et al., 2009

Paxillus leachi, Heliscus tropicus and Veturius marilucae

Catemaco, Coyame and Bastonal, Veracruz

K. reyesi* Rosario & Hunter, 1988 Petrejoides sp. Huautla de Jiménez, Oaxaca

K. santibanezi* Villegas-Guzman et al., 2009

Proculejus sartorii, P. brevis, P. nudicostis and Vindex sp.

San Miguel Talea de Castro, Cerro Pelón and Ixtlán de Juárez, Oaxaca

continued next page

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Zootaxa 2747 © 2011 Magnolia Press · 21KLINCKOWSTROEMIA FROM MEXICO AND PANAMA

Klinckowstroemia pennula sp. nov.

Female (n = 15)Body oval. Idiosoma: L = 800 (717–851), W = 570 (586–576) (Fig. 1A). Gnathosoma: seta hyp1 long and simple, 56 (50–56). hyp2 medium and serrate, L = 34 (28–34). hyp3 short and

simple, L = 9 (6–9) (Fig. 2A). hyp4 short and serrate, 6 (6–12), chelicera with excrescences, movable digit withfive teeth and fixed digit with seven teeth. Palp setae simple, but av1 and av2 on trochanter branched and serrate,respectively. Anterior margin of palp trochanter with blunt process (b) slightly rounded, and denticulate process (s)divided in some specimen.

Hyaline hood extending to level of coxa I, seta a1 medium and simple, L = 12 (12–17). Tritosternum basewider than long. Presternal shield (Pst) L = 43 (36 –43), anterior W= 102 (90–102) (Fig. 2D); near the posteriormargin is a lyriform sternal pore stp1. Presternal notch square, shagreened, with a line near to each lateral margin.Sternal seta st1, L = 17 (9–17), simple (Fig. 2C).

Sternal shield (Fig. 2B) L = 65 (53–71), stp2 placed at 1/3 the length of the shield, st2 below stp2, setae st2, st3,and st4 are micro setae, setae st3 and st4 near posterior margin and close to each other. Shield is shagreened, with-out reticulations.

Sternogynal shield, L = 19 (15–25); W = 155 (135–158) reduced, shagreened, with stp3 closer to anterior mar-gin. Sternogynal apodeme short. Latigynal shields, L = 109 (90–109), W = 68 (65–76) shagreened, each with anoval pore and 4–5 setae (Fig. 2E), medial margin of shield wavy. Mesogynal shield, L = 53 (43–62), W = 68 (62–78), triangular, rounded anteriorly, surface shagreened. Mesogynal condyles extending almost to level of shieldapex.

TABLE 1. (continued)

Species Hosts Localities

K. schusteri Rosario & Hunter, 1988 Petrejoides sp. Huehuetenango, Guatemala

K. scotti* Rosario & Hunter, 1988 Pruculejus sp. Agua Blanca and Tenango de Doria, Hidalgo

K. simplesetosa* Rosario & Hunter, 1988 Verres hageniVerres corticicolaHeliscus tropicusPassalus (Pertinax) punctatustriatusVerres corticicola

Dos Amates, VeracruzSan Andrés Tuxtla, Veracruz“Los Tuxtlas” E. B. T., VeracruzBalzapote, VeracruzLaguna Escondida, Veracruz

K. starri Rosario & Hunter, 1988 Passalus (Pertinax) sp. Passalus jansoniVerres hageniVeturius cirratusVeturius tuberculifrons

Costa Rica

K. stilla*, this paper Proculus goryi Reserva de la Biosfera Volcán Tacaná, Chiapas

K. tapachulensis* Chernoff & Pope, 1970 Passalus interruptusPopilius eclipticus

Tapachula, Chiapas

K. tragardhi Baker & Wharton, 1952 Undetermined Passalid Surinam

K. truncata Hunter & Butler, 1966 Passalus sp. Pseudacanthus sp.

La Selva, Costa Rica

K. valdezi* Villegas-Guzman et al., 2009 Proculejus hirtus and P. brevisOileus nonstriatus and Odontotaenius zodiacus

Huayacocotla, VeracruzTianguistengo and Tizapan, Hidalgo

K. victoriae* Rosario & Hunter, 1988 Oileus herosOgyges laevissimusPublius agassizi

Xicotepec de Juárez, PueblaQuetzaltenango, GuatemalaPuerto Parada, Guatemala

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FIGURE 1. Klinckowstroemia pennula sp. nov. (A) Body female ventral view, scale = 200 µm. (B) Body male ventral view,scale = 200 µm.

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Zootaxa 2747 © 2011 Magnolia Press · 23KLINCKOWSTROEMIA FROM MEXICO AND PANAMA

FIGURE 2. Klinckowstroemia pennula sp. nov. Female: (A) Gnathosoma. (B) Presternal and seta st1. (C) Seta st1. (D) Sternalshield. (E) Genital shield. (F) Seta pv1 on leg IV. Male: (G) Sternal shield. (H) Presternal and seta st1.

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Ventral shield, L = 248 (231–257), posterior W = 431 (360–450), shield triangular, truncated posteriorly, sur-face shagreened, with 12–14 pairs of small setae (Fig. 1A). One pore located below posterior margin of coxa IV.The metapodal-peritrematal-exopodal shield (MPES) reticulated, with a pair of pores and one simple seta.

Anal shield, L = 127 (96–140); anterior W = 341 (252–372), with 6–7 pairs of simple setae plus longer antero-anal seta, L = 29 (22–29), two pairs of pores, one near the anterior margin, the other below the anus (Fig. 1A).

Serrate setae on leg I are: trochanter ad1, av1, al1 and pv1; femur pv1, pv2, and pv3; genu al1, av1, and pv1.Serrate setae on leg II are: trochanter pd1; femur pv1, and genu av1, av1, and pv1. Serrate setae on leg III: tro-chanter pd2, and pv1; femur av1, and pl1; genu pv1; tibia ad4 and al4. Serrate setae on leg IV: trochanter pv1, andpl1; femur av1; genu av1, and pv1. Seta pv1 of femur IV large, L = 31 (25–31) (Fig. 2F).

Male (n = 11)Body oval, similar to female. Idiosoma L = 742–851, W = 499–576 (Fig. 1B). Presternal shield, L = 43–65,

anterior W = 93–115, anterior margin concave without presternal notch (Fig. 2H), shagreened. Sternal seta st1, L =11–16, serrate. The other sternal setae are microsetae. Sternal shield shagreened, L = 62–71 (Fig. 2G). Genitalopening oval, L = 43–50, W = 65–78. Ventral shield with 36 setae (Fig. 1B), L = 347–375, posterior W = 357–467,shagreened on the anterior region, and reticulated below coxa IV. Anal shield L = 118–133, anterior W = 279–357;with 5 pairs of simple setae plus longer antero-anal seta, L = 20–30. Hypostomal seta hyp1 is long and simple, L =50–56; seta hyp2 is medium and serrated, L = 25–34, while seta hyp3 is simple and short, L = 6–11.

Diagnosis. Sternogynal shield is very much reduced (15–24. 8), and the latigynal shields are bigger than onother species (width 65. 1–77. 5), also sternal setae st2, st3, and st4 are microsetae. Mesogynal shield is rounded onanterior margin. Male has presternal shield expanded anteriorly, sternal seta st1 in middle of the shield; ventralshield in posterior region reticulated.

Remarks. Klinckowstroemia pennula sp. nov. is similar to K. victoriae Rosario & Hunter and K. melissae sp.nov., because these species have latigynal shields with wavy medial margins. These species are distinguished bythe number of the setae on the latigynal shields (4–5 K. pennula and K. melissae, three for K. victoriae), bodylength and width (K. pennula L 717–851, W 486–576; K. melissae, L 723–736, W 518–531; K. victoriae L 834–872, W 570–616) and length of the medial margin of the latigynal shield (K. pennula 43–62, and K. melissae 28–39). Additionally, the presternal shield of male K. pennula has a concave anterior margin, slightly longer than on K.melissae (Tp 43–65, Tm 50–56).

Etymology. This name is from Latin pennula (= little wing) referring to the shape of the latigynal shield. Material examined. From undescribed Pseudacanthus sp.: holotype ♀ (CNAC006706), 1 ♀ (CNAC006707)

and 2 ♂♂ (CNAC006708-6709) paratypes from Oaxaca, Mexico, 7 km E San Pedro-San Pablo Ayutla, Ixtlán deJuárez (17 35. 293’ N, 96 27. 162’ W), 2,459 m, 20 July 2007, mountain rain forest, ex one beetle, O. Francke, A.Valdez, C. Santibañez, A. Ballesteros, and H. Montaño. Mites were found on humeri, and in the alcohol. Fromundescribed Pseudacanthus sp. 3♀♀ (CNAC006710- CNAC006712) and 1 ♂ (CNAC006713) paratypes fromOaxaca, Mexico, 7 km E San Pedro-San Pablo Ayutla, Ixtlán de Juárez (17 35. 293’ N, 96 27. 162’ W), 2,459 m, 20July 2007, mountain rain forest, ex one beetle, O. Francke, A. Valdez, C. Santibañez, A. Ballesteros, and H. Mon-taño. Mites were found on coxa I, and in the alcohol. From Vindex agnoscendus 1♀ (CNAC006714) and 1♂(CNAC006715) paratypes from Oaxaca, Mexico, 10 km NE Mirador Santiago Comaltepec, Ixtlán de Juárez (1735. 293’ N, 96 27. 162’ W), 2,188 m, 16 June 2007, mountain rain forest, ex one beetle, O. Francke, A. Valdez, C.Santibañez, A. Ballesteros, and H. Montaño. Mites were found in the alcohol. From Proculejus sp. 2♀♀(CNAC006716, CNAC006717) paratypes from Oaxaca, Mexico, 10 km NE Mirador Santiago Comaltepec, Ixtlánde Juárez (17 35. 293’ N, 96 27. 162’ W), 2,188 m, 16 June 2007, mountain rain forest, ex one beetle, O. Francke,A. Valdez, C. Santibañez, A. Ballesteros, and H. Montaño. Mites were found in the alcohol. From Pseudacanthussp. nov. 1♀ (CNAC006718) and 2 ♂♂ (CNAC006719, CNAC006720) paratypes from Oaxaca, Mexico, 7 km ESan Pedro-San Pablo Ayutla, Ixtlán de Juárez (17 35. 293’ N, 96 27. 162’ W), 2,459 m, 20 July 2007, mountainrain forest, ex one beetle, O. Francke, A. Valdez, C. Santibañez, A. Ballesteros, and H. Montaño. Mites were foundon femur, coxa II, and in the alcohol. From undescribed Pseudacanthus sp. 1♀ (CNAC006721) and 2 ♂♂(CNAC006722, CNAC006723) paratypes from Oaxaca, Mexico, 7 km E San Pedro-San Pablo Ayutla, Ixtlán deJuárez (17 35. 293’ N, 96 27. 162’ W), 2,459 m, 20 July 2007, mountain rain forest, ex one beetle, O. Francke, A.Valdez, C. Santibañez, A. Ballesteros, and H. Montaño. Mites were found in the alcohol. From Oileus bifidus 1♀(CNAC006724) and 1 ♂ (CNAC006725) paratypes from Oaxaca, Mexico, Camino a El Porvenir, San MiguelTalea de Castro, (17 18. 160’ N, 96 14. 423’ W), 1,870 m, 22 July 2007, mountain rain forest, ex one beetle, O.

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Zootaxa 2747 © 2011 Magnolia Press · 25KLINCKOWSTROEMIA FROM MEXICO AND PANAMA

Francke, A. Valdez, C. Santibañez, A. Ballesteros, and H. Montaño. Mites were found on mentum, near the eyes,and in the alcohol. From Oileus bifidus 1♀ (CNAC006726) and 1 ♂ (CNAC006727) paratypes from Oaxaca, Mex-ico, Camino a El Porvenir, San Miguel Talea de Castro, (17 18. 160’ N, 96 14. 423’ W), 1,870 m., 22 July 2007,mountain rain forest, ex one beetle, O. Francke, A. Valdez, C. Santibañez, A. Ballesteros, and H. Montaño. Miteswere found on prosternum and in the alcohol. From Oileus bifidus 1♀ (CNAC006728) paratype from Oaxaca,Mexico, Camino a El Porvenir, San Miguel Talea de Castro, (17 18. 160’ N, 96 14. 423’ W), 1,870 m, 22 July2007, mountain rain forest, ex one beetle, O. Francke, A. Valdez, C. Santibañez, A. Ballesteros, and H. Montaño.Mites were found on prosternum. Holotype female, 11 paratype females, and 8 paratype males deposited in theCNAC, one female and one male paratype deposited in NMNH, UGA, and UMMZ respectively.

Ecology. The passalid beetles were found in decaying trunks at three different localities in the state of Oaxaca,Mexico, and the mites were present on four different hosts. On the undescribed Pseudacanthus sp. (three speci-mens living in the same trunk), were two species of mite, K. pennula and K. melissae. The passalid beetle Oileusbifidus (five specimens found inhabiting the same trunk) also had two species of mites, K. pennula and K. candidoi.In both cases we considered that the host beetles belonged to the same “family or social group of passalid beetlesbecause they live in the same log, and for this reason we can find two species of klinckowstroemiids on the variousindividuals of the same social group; if they did not inhabit the same galleries they would probably not share thesame phoretic mite species. The mites were located on the passalids’ body, on the coxa II, femur, prosternum, nearthe eyes and on the mentum; we also found many mites in the alcohol. The beetles were collected in mountain rainforest in a range of altitude of 1,870 to 2,459 m.

Klinckowstroemia melissae sp. nov.

Female (n = 3)Body oval. Idiosoma, L = 736 (723–736); W = 531 (518–531) (Fig. 3A). Gnathosoma with different types of setae: hyp1 long and simple, L = 48 (47–50), longer than other three. hyp2

serrate, L = 33 (33–34). hyp3 short and serrate, L = 12 (12–14) (Fig. 4B). hyp4 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 av1 and av2 of trochanter branched and serrate respectively. Anterior margin of palptrochanter with blunt process (b) and slightly longer spine-like process (s).

Hyaline hood extending to level of coxa I, seta a1 large and serrate, L = 25 (25–27). Tritosternum base widerthan long. Presternal shield L = 37 (37–43), anterior W = 99 (96–99), shagreened (Fig. 4C). Presternal notchshaped like a small square. Sternal seta st1 serrate (Fig. 4D), L = 27 (25–32), near to anterior margin; opposite mar-gin has lyriform sternal pore stp1.

Sternal shield reticulated and shagreened (Fig. 4E), L = 56, near the anterior margin of the shield is stp2, belowit is sternal seta st2, L = 19 (16–19), serrate and medium. Near the posterior margin of the shield are sternal setaest3, L = 16 (12–16) serrated, and st4 is a microseta.

Sternogynal shield triangular (Fig. 4A) L = 47; W = 143, posterior margin narrow and pointed at apex, surfaceis shagreened and reticulated, stp3 located closer to anterior margin. Sternogynal apodeme with broad base thatreaches 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. 4A), medial margin ofshield wavy and medium, this shield reticulated and shagreened. Latigynal apodeme thick and separate from thesternogynal shield. Mesogynal shield, L = 53 (53–56), W = 81, triangular, anterior margin rounded and posteriormargin extended (Fig. 4A), 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, sur-face reticulated and shagreened, with 13–14 pairs of small setae. One pore located posterior margin of coxa IV(Fig. 3A). The metapodal-peritrematal-exopodal shield (MPES) reticulated, with a pair of pores and one simpleseta (Fig. 3A).

Anal shield, L = 124 (124–130); anterior W = 360 (341–360), with 7–8 pairs of simple setae (Fig. 4F) plus lon-ger 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 al1; femur I, pv2, and pv3; genu pl1. Serrrate setae on leg II are: femurpv1; genu pv1; and tibia al4. Serrate setae on leg III are: femur av1; genu av1, and pv1; tibia ad4 and, tarsus ad3.

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FIGURE 3. Klinckowstroemia melissae sp. nov. (A) Body female ventral view, scale = 200 µm. (B) Body male ventral view,scale = 200 µm.

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Zootaxa 2747 © 2011 Magnolia Press · 27KLINCKOWSTROEMIA FROM MEXICO AND PANAMA

FIGURE 4. Klinckowstroemia melissae sp. nov. Female: (A) Genital shield. (B) Gnathosoma. (C) Presternal and seta st1. (D)Seta st1. (E) Sternal shield. (F) Anal shield. Male: (G). Presternal and seta st1. (H) Sternal shield.

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VILLEGAS-GUZMÁN ET AL.28 · Zootaxa 2747 © 2011 Magnolia Press

Serrate setae on leg IV: trochanter al1, pv1, and pl1; femur av1; genu av1, pl1, and pv1; tibia ad4; tarsus ad2, andad3. Seta pv1 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. 3B). Presternal shield, L = 50–56,

anterior W = 93–96, is shagreened, presternal notch shaped like an inverted triangle, moderately deep (Fig. 3B).Sternal seta st1, L = 19–22, serrate, near to anterior margin. Sternal seta st2, L = 14–19, serrated, seta st3, L = 7–9simple, and seta st4 a microseta. Sternal shield reticulated, L = 62–76 (Fig. 3B). Genital opening oval, L = 47–50,W = 71–74. Ventral shield with 32–34 setae (Fig. 3B), L = 248–279, W = 403–419, shagreened, laterally reticu-lated. One pore near the anterior margin of coxa IV. Anal shield, L = 124–127, anterior W = 326–372, with 6–7pairs of simple setae, plus longer antero-anal seta L = 25–31; with two pores, one near the anterior margin and theother below the anus. Hypostomal seta hyp1 is long and simple, L = 47–56, seta hyp2 is serrate and long, L = 33–37, while seta hyp3 is serrate and shorter, L = 9–16. Seta pv1 serrated on coxa IV, L = 20–31.

Diagnosis. Sternal, latigynal, sternogynal, and ventral shields are reticulated, and setae st1, st2, and st3 are ser-rated. Mesogynal shield with posterior margin wide. Male has sternal shield reticulated and st1 and st2 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 bythe 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. melissaeshare the number of setae on latigynal and ventral shields. These species are distinguished by length of sternogynalshield (K. pennula L = 50–62; K. melissae L = 28–34); and setae st1 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 thesenior author during this research; many thanks Bichita.

Material examined. From undescribed Pseudacanthus sp.: holotype ♀ (CNAC006729), 4 ♀♀ and 5 ♂♂(CNAC006730-6738) paratypes from Oaxaca, Mexico, 7 km. E San Pedro-San Pablo Ayutla, Ixtlán de Juárez (1735. 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. Hol-otype 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 undescribedPseudacanthus sp. Two of these passalids were living in the same log, and the other passalid beetle was collected inthe same locality and day. Specimens of K. pennulae were more abundant than K. melissae; this is the first casewhere two species found on the same host (synhospitality) are almost the same size. In other known cases of syn-hospitality one mite species is larger than the other (Villegas-Guzman et al. 2009).

Klinckowstroemia stilla sp. nov.

Female (n = 5)Body oval. Idiosoma, L = 563 (557–595); W = 397 (397–437) (Fig. 5A). Gnathosoma with different types of setae hyp1 long and simple, L = 31 (30–34). Seta hyp2 simple, L = 19. Seta

hyp3 simple, L = 9 (Fig. 6A), and hyp4 simple, L = 12 (8–12). Chelicera with excrescences, movable digit with fiveteeth, two small and three large; fixed digit with six teeth. Most palpal setae simple, but av1 and av2 of trochanterare branched and serrate respectively. On femur al1 serrate. Anterior margin of trochanter palp with blunt process(b) and spine-like process (s).

Hyaline hood extending to level of coxa I, seta a1 short and simple, L = 12 (7–12). Tritosternum base widerthan long. Presternal L = 25 (22–25), anterior W = 78 (78–81), shagreened (Fig. 6D); anterior margin of the shieldslightly concave medially, without presternal notch. Sternal seta st1 micro seta (Fig. 6E), placed near to anteriormargin, and lyriform sternal pore stp1 near to posterior margin.

Sternal shield (Fig. 6C) L = 47 (37–56) shagreened, sternal seta st2 microseta placed near to anterior margin,stp2 below, almost in the centre of the shield. Sternal setae st3 and st4 microsetae, near posterior margin of shield.

Sternogynal shield L = 31 (31–34), W = 109 (109–124), triangular, posterior margin narrow and slightlypointed at apex. Surface shagreened; stp3 located closer to anterior margin. Sternogynal apodeme with broad basethat reaches to the shield, posterior arms thin, extending posteriorly to level of latigynal and mesogynal condyles.

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FIGURE 5. Klinckowstroemia stilla sp. nov. (A) Body female ventral view, scale = 200 µm. (B) Body male ventral view, scale= 200 µm.

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FIGURE 6. Klinckowstroemia stilla sp. nov. Female: (A) Gnathosoma. (B) Genital shield. C) Sterna shield. (D) Presternal andseta st1. (E) Seta st1. (F) Seta pv1 on leg IV. (G) Ventral shield. (H) Anal shield.

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Latigynal shields shagreened, L = 90 (84–90); W = 56, each with an oval pore and four simple setae, medial marginof shield pointed (Fig. 6B). 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. 6B), posterior marginrounded 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. 6G), truncated posteri-orly, surface shagreened, with 12 pairs of small setae (Fig. 6H). Three pores underneath posterior margin of coxaIV (Fig. 3A). Metapodal-peritrematal-exopodal shield (MPES) shagreened, with one pair of pores and one simpleseta (Fig. 5A).

Anal shield, L = 87 (87–93); anterior W = 217 (217–257), with 5 pairs of simple setae (Fig. 6H) plus longerantero-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 pv1; femur pv1, pv2, and pv3; genu I, al1 and av1; tibia av1, and av2 ser-rated. Serrate setae on leg II are: femur pv1; genu av1, and pv1. Serrate setae leg III are: trochanter ad1, and ad2;femur av1; genu pv1. Serrated setae on leg IV are: trochanter pv1; genu pv1. Seta pv1 of femur IV large, L = 36(34–40) (Fig. 6F).

Male (n = 2) Body oval, similar to female. Idiosoma, L = 531–544, W = 371–384 (Fig. 5B). Presternal shield, L = 31–34,

anterior W = 56–62, shagreened, without presternal notch, anterior margin slightly concave (Fig. 5B). Sternal setast1 microseta is near the middle of the shield. Sternal setae st2, st3, and st4 are microsetae. Sternal shield sha-greened, L = 34–43. Genital opening oval, L = 37, W = 50–53. Ventral shield with 30 setae (Fig. 5B), 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 anteriormargin and the other below the anus. Hypostomal seta hyp1 long and simple, L = 31–32, seta hyp2 is simple andmedium, L = 17–19, while seta hyp3 is short and simple, L = 7–9. Seta pv1 serrated on coxa IV, L = 33–34.

Diagnosis. The mesogynal shield extending deeply to the sternogynal shield, with its posterior margin wideand 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 Rosa-rio & Hunter, because they have the mesogynal shield extending deeply to the sternal shield, and the medial mar-gins 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 inK. stilla and K. multisetillosa. Furthermore, K. stilla has all the sternal setae as microsetae, while K. starri hasmedium-size sternal setae, with st1 serrated and the others simple. Klinckowstroemia multisetillosa has six pairs ofanterior setae a and the lateral margins of mesogynal shield concave, but K. stilla only has one pair of setae a andthe lateral margins of mesogynal shield straight.

Etymology. This name is from Latin stilla (= drop) referring to the shape of the mesogynal shield which issimilar a drop of water.

Material examined. Proculus goryi: holotype ♀ (CNAC006739), 4 ♀♀ (CNAC006740-6743) and 2 ♂♂(CNAC006744-6745) paratypes from Chiapas, Mexico, Ejido Benito Juárez, Reserva de la Biosfera Volcán Tacan(15 05. 966’ N, 92 08. 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, 4paratype 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 nearthe 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).

Klinckowstroemia oconnori sp. nov.

Female (n = 3)Body oval. Idiosoma: L = 781 (781–806); W = 576 (563–576) (Fig. 7A). Gnathosoma with different hypostomal setae: hyp1 long and simple, L= 50 (47–54); seta hyp2 long and ser-

rated, L = 43 (40–43), seta hyp3 medium and serrated, L = 16 (12–16), and seta hyp4 medium-sized and serrated, L= 16 (9–16).

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FIGURE 7. Klinckowstroemia oconnori sp. nov. (A) Body female ventral view, scale = 200 µm. (B) Body male ventral view,scale = 200 µm.

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Zootaxa 2747 © 2011 Magnolia Press · 33KLINCKOWSTROEMIA FROM MEXICO AND PANAMA

FIGURE 8. Klinckowstroemia oconnori sp. nov. Male: (A) Gnathosoma. (B) Presternal and seta st1. (C) Seta st1. (D) Sternalshield. (E) Reticulations on sterna shield. (F) Palp trochanter. (G) Anal shield. (H) Ventral shield.

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Chelicera with excrescences, movable digit with four teeth and fixed digit with seven teeth. Palp setae are sim-ple, but av1 and av2 on trochanter are branched and serrated, respectively; on femur pd1, pd2, and pv1 are serrated.On genu the setae al1, al2, ad1, pl1, pd1 and pv1 are serrated.

Hyaline hood extending to level of coxa I, seta a1 moderate and simple, L = 19 (19–22). Tritosternum basewider than long. Presternal (Pst) L = 37 (34–37), anterior W= 105 (102–109) shagreened and with line near theposterior margin; near posterior margin is a lyriform sternal pore stp1. Presternal notch is a small concave depres-sion. Sternal seta st1, L = 53 (34–53), serrated (Fig. 7A).

Sternal shield L = 59 (56–62), shagreened, with some reticulations (Fig. 7A). Lyrifissure stp2 near to anteriormargin of the shield; st2, L = 47, serrated, below. Sternal setae st3, L = 37, and st4, L = 40, both setae are large andserrated, 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 marginstraight and slightly pointed, near to anterior margin is pore stp3. Sternogynal apodeme thin and short. Latigynalshields shagreened, L = 130 (127–130); W = 65 (62–65), each with an oval pore and four simple setae, medial mar-gin of shield straight. Mesogynal shield, L = 62 (59–62); W = 83. 7, triangular and shagreened, posterior marginwider 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, truncatedposteriorly, with ten pairs of small setae (Fig. 7A), with two pairs of pores at the level of posterior margin of coxaIV. 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 antero-anal seta, L = 22 (22–25); two pairs of pores, one near the anterior margin, and the other below the anus (Fig. 7A).

Serrate setae on leg I are: trochanter ad1, al1 and pv1; femur: av3, pv1 and pv2; tibia ad1, ad2, ad3, av1, av2,pl1, and pl2. Serrate setae on Leg II are: femur pv1; genu pv1. Serrate setae on leg III: trochanter ad1 and ad2;femur av1; genu av1, and pv1; tibia ad4; tarsus ad3, and pl1. Serrate setae on leg IV: femur av1; genu av1, and pv1;tibia ad4, and al4; tarsus ad2 and ad3 serrated. Seta pv1 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. 7B). Presternal (Pst) L = 31–37, ante-

rior W = 102–109, presternal notch V-shaped, shagreened and with one line near anterior margin (Fig. 8B) as infemale; near to posterior margin is a lyriform sternal pore stp1. Sternal seta st1, L = 40–56, long and serrated (Fig.8C). Sternal shield shagreened (Fig. 8D) and reticulated (Fig. 8E), L = 121–127, with two pores, one near the ante-rior margin (stp2) and other near the genital opening (stp3). Sternal seta st2 long and serrated, L = 48–50; sternalseta st3 long and serrated, L = 39–47; sternal seta st4 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. 8H), with 30 setaeand two pores, one above the anterior margin of coxa IV, and other below coxa IV. Anal shield L = 102–140, ante-rior W = 322–363 (Fig. 8G); with four pairs of simple setae plus longer antero-anal seta, L = 12–25, with twopores: one near the anterior margin and another below the anus. Hypostomal setae (Fig. 8A): hyp1 long and simple,L = 43–56; seta hyp3 serrate and long, L = 32–43; seta hyp2 simple and shorter, L = 11–12. Anterior margin of tro-chanter palp with blunt process (b) and spine-like process (s) (Fig. 8F).

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 st1, st2, and st3long and serrated, and seta st4 medium and serrated.

Remarks. Klinckowstroemia oconnori sp. nov. is similar to K. montanoi Villegas-Guzman et al. because inboth 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 spe-cies have sternal setae long and serrated, but the sternal shield in K. ballesterosi is shagreened, whereas on K. ocon-nori 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 ofthis species and gave us the mites.

Material examined. From Odontotaenius striatopunctatus: holotype ♀, and 2 ♂♂ paratypes from Panama,Panama, Cerro Azul 950 m, 6 May 1991,B. M. OConnor; mites were found in the alcohol. From Petrejoides subre-cticornis 2♀♀ and 2 ♂♂ paratypes from Panama, Panama, Cerro Azul 950 m, 6 May 1991, B. M. OConnor; miteswere found in the alcohol. Holotype female, and 3 paratypes males deposited in NMNH, one female (CNAC006746) and a male (CNAC 006747) paratypes are deposited in CNAC. One female was lost when it was beingprepared for scanning electron micrographs, but this specimen was considered for description.

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Ecology. Specimens were found on two species of passalid beetles. The beetles were probably found in thesame log, because they were collected at the same locality and on the same day; perhaps, for this reason they sharethe same species of mites

Discussion

As in the previous study, we found individual bessbugs with two different species of klinckowstroemiids: Klinck-owstroemia pennula cohabiting with K. melissae on Pseudacanthus sp., and K. pennula with K. candidoi on Oileusbifidus; both cases were found in the state of Oaxaca, Mexico. We found 18 passalid beetles with two species ofKlinckowstroemia each, the species with more cases are: Odontotaenius zodiacus and Oileus bifidus, both with fivecases; in those ten beetles we found eight different species of Klinckowstroemia. Synhospitality appears to occurwhen populations of mites overlap geographically but only one species of passalid host occurs in that zone; there-fore, the mites must use the same host.

This is the first record of the genus Klinckowstroemia in Panama. In that country only Similantennurellaspinata Rosario, 1988, Klinckowstroemiella sexisetosa, Rosario & Hunter, 1987, and Antennurella trouessarti Ber-lese, 1904 had been collected before. With this new record Panama becomes the only known country where thefour genera of the family Klinckowstroemiidae co-exist. This family has been little studied, and the principal prob-lem apparently is the low frequency of occurrence of these mites on passalids. According to Rosario (1988), out of100 specimens beetles examined by her, only one specimen carried klinckowstroemiids. In the last five years wecollected and checked more than 1,150 passalid beetles from Mexico; of these 758 carried mites, and only 168were klinckowstroemiids (14. 6%, rate of infestation). This data illustrates the low frequency of this family ofmites. The number of klinckowstroemiid varies from 1 to 40 mites per passalid host. We consider it important tocontinue collecting passalid beetles to look for mites, specially klinckowstroemiids, because in countries like Bra-zil, Colombia, and Venezuela, there are no records of klinckowstroemiids associated with passalid beetles,although they are expected to be here inasmuch as that both the hosts and the phoronts are of Gondwanan origin(Kim, 2004).

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. 7A) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 82. Sternal and latigynal setae barbed and/or serrated. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4- Sternal and latigynal setae smooth. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 33. All sternal setae of equal length . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . K. simplesetosa- Sternal setae st2, st3 and st4 are microsetae (Fig. 2G) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ..K. pennula4. With four pairs of setae a; st1 long and barbed, not serrated. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .K. scotti - With one pair of setae a, st1 serrated, not barbed . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .55. Seta st1 long (31–40) and serrated, setae st3 and st4 are medium and/or microsetae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .6- Seta st1 medium (21–29) and serrated, setae st3 and st4 are short . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .K. montanoi6. Setae st3 of medium length . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .7- Setae st3 and st4 are microsetae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . K. valdezi7. Seta st4 short . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . K. victoriae- Seta st4 a microseta (Fig. 4H) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .K. melissae8. Sternal seta st1 barbed or bifurcate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9- Sternal seta st1 smooth . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 199. Sternal seta st1 barbed. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10- Sternal seta st1 bifurcate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .K. bifurcata10. Sternal seta st2 barbed. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11- Sternal seta st2 smooth. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1411. Sternal setae st3 and st4 smooth . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .K. candidoi- Sternal setae st3 and st4 barbed and serrated. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1212. Sternal seta st1 very long, st2 and st3 medium; with 6–10 serrated setae on latigynal shields . . . . . . . . . . . . . . . .K. santibanezi- Sternal setae st1, st2 and st3 are long and serrated . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1313. Sternal shield granulated. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .K. ballesterosi - Sternal shield reticulated (Fig. 7A). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . K. oconnori14. Mesogynal shield rounded anteriorly . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .K. atramaculata

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- Mesogynal shield pointed anteriorly. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1515. Medial margins of latigynal shields pointed. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .K. tapachulensis- Medial margins of latigynal shields truncate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1616. Seta st2 short or medium; body more than 900 µm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 17- Seta st2 microseta, body less than 900 µm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1817. Seta st1 is the longest of sternal setae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . K. schusteri- Seta st3 is the longest of sternal setae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .K. cristinae18. Latigynal shields with 4–6 setae setae st3 and st4 microsetae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . K. brevimarginata- Latigynal shields with three setae, setae st3 and st4 short . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .K. rectimarginata19. Two pairs of setae on latigynal shield. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .K. tragardhi- More than two pairs of setae on latigynal shields . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2020. Posterior margin of ventral shield concave . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .K. concava- Posterior margin of ventral shield truncate. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2121. 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. . . . . . . . . . . 2622. Mesogynal shield extending anteriorly to sternogynal shield slightly . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 23- Mesogynal shield extending anteriorly to sternogynal shield deeply . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2423. Seta st1 microseta, st2 short and simple, hyp3 serrated . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . K. franckei- Seta st1 short and simple, st2 microseta, hyp3 simple . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .K. hunteri24. Lateral margins of mesogynal shield concave; all sternal setae of equal length . . . . . . . . . . . . . . . . . . . . . . . . .K. multisetillosa- Lateral margins of mesogynal shield straight . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2525. Sternal setae medium length, sternal and sternogynal shields reticulated. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . K. starri- Sternal setae are microsetae, sternal and sternogynal shields granulated (Fig. 6C).. . . . . . . . . . . . . . . . . . . . . . . . . . . . . K. stilla26. 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. . . . . . . . . . . . . . . . . . . . . . . . . . 2727. Sternal seta st2 shorter than st3 and st4; three pairs of setae on latigynal shields . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . K. reyesi Sternal seta st2 longer than st3 and st4; four pairs of setae on latigynal shields. . . . . . . . . . . . . . . . . . . . . . . . . . . . . .K. truncata

Acknowledgments

We express our sincere thanks to the members of Colección Nacional de Ácaros and Colección Nacional de Arác-nidos, who collected the specimens and made them available for study. Dr. Oscar Francke kindly reviewed themanuscript. We also thank Drs. Ronald Ochoa (NMNH), Barry OConnor (UMMZ), and Dac Crossley (UGA) wholent holotypes and paratypes of Klinckowstroemia for this study, and M. S. Berenit Mendoza for taking the photo-graphs with the scanning electron microscope. Two anonymous reviewers kindly reviewed the manuscript andmade significant recommendations for its improvement. The senior author was supported by Conacyt and Posgradoen Ciencias Biológicas de la UNAM.

References

Baker, E.W. & Wharton, G.W. (1952) An Introduction to Acarology. Macmillan: New York, 465 pp. Chernoff, N. & Pope, R. (1970) Two new mites of the genus Klinckowstreomia from Mexico (Acarina: Klinckowstroemiidae).

Proceedings of the Biological Society of Washington, 83, 145–154. Hunter, P.E. & Butler, L. (1966) New Klinckowstroemia mites from Costa Rican passalid beetles (Acarina: Klinckowstroemii-

dae). Journal Georgia Entomological Society, 1, 24–30. Kim, C.-M. (2004) Trigynaspida (Acari: Mesostigmata): new diagnosis, classification, and phylogeny. Acarologia, 44, 157–

197. Padilla, E.E. & Schuster, J. (1994) Klinckowstroemia multisetillosa Rosario & Hunter (Acarina: Trygynaspida: Klinckowstro-

emiidae) associated with three species of Proculus Kuwert (Coleoptera: Passalidae). Acta Zoológica Mexicana (n. s. ), 61,1–15.

Rosario, R.M.T. (1988) The genus Antennurella Berlese and descriptions of two new species of Similantennurella gen. nov.(Acarina: Trigynaspida: Klinckowstroemiidae). Acarologia, 29, 329–337.

Rosario, R.M.T. & Hunter, P.E. (1987) The family Klinckowstroemiidae Trägårdh with descriptions of two new species ofKlinckowstroemiella (Acarina: Mesostigmata: Trigynaspida). Acarologia, 28, 307–317.

Rosario, R.M.T. & Hunter, P.E. (1988) The genus Klinckowstroemia Trägårdh and descriptions of nine new species (Acarina:Trigynaspida: Klinckowstroemiidae). Acarologia, 29, 119–136.

Seeman, O.D. (2007) Revision of the Fedrizziidae (Acari: Mesostigmata: Fedrizzioidea). Zootaxa, 1480, 1–55. Villegas-Guzman, G.A., Pérez, T.M. & Reyes-Castillo, P. (2009) New species of the genus Klinckowstroemia Baker & Wharton

from Mexico (Acari: Mesostigamata: Klinckowstroemiidae). Zootaxa, 2248, 1–46.

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