Chrysomelobia matsuzawai sp. nov. and Chrysomelobia nipponica sp. nov. (Acari: Podapolipidae) are described from Gonioctena rubripennis Baly (Coleoptera: Chrysomelidae) collected in Japan. This is the first record of the genus Chrysomelobia Regenfuss, 1968 in Asia. Adult females of the type species for Chrysomelobia, Chrysomelobia mahunkai Regenfuss, 1968, were recollected from female specimens infesting a Gonioctena sp. in Germany. An updated key to all species of Chrysomelobia is provided.
Mites in the family Podapolipidae are common parasites of a number of families of Coleoptera, and less commonly on Blattodea and Orthoptera (e.g., Regenfuss 1968; Husband 1990; Husband & OConnor 2003). The family Podapolipidae is represented by one species each on Heteroptera (Kurosa & Husband 1994) and Hymenoptera (Husband & Sinha 1970) and has not been found on Diptera, Lepidoptera, Odonata or any aquatic insects. In most instances, each family of parasitized Coleoptera will have genera of Podapolipidae that are associated with that family only. A conspicuous exception is the genus Podapolipus Rovelli & Grassi, 1888 which is found on beetles in four families as well as on insects in the orders Blattodea and Orthoptera (e.g., Husband 1986). Previous reports of Podapolipidae on Chrysomelidae have been recorded as species in the genus Chrysomelobia Regenfuss, 1968 (Parobia Seeman & Nahrung, 2003) (Regenfuss 1968; Eickwort 1975; Drummond et al. 1984; Fain 1987; Haitlinger 1989; Houck 1992; Moraes et al. 1999; Husband & Moraes 1999; Seeman & Nahrung 2003, 2005, 2013; Husband & OConnor 2004; Seeman 2008). The single record of a podapolipid mite that is not a Chrysomelobia but is a parasite of a chrysomelid beetle is Cassidopohpus physonotae Husband & OConnor, 2014, a parasite of Physonota alutacea Beheman (Husband & OConnor 2014). Twenty-one species of Chrysomelobia have been reported from Australia (14), Africa (2), Europe (1), and the Americas (4) on leaf beetles belonging to the subfamily Chrysomelinae (Seeman & Nahrung 2013). The record of a single female specimen of C. donati Haitlinger, 1989 from a cercopid hemipteran is considered accidental. Chrysomelobia nipponica sp. nov. and Chrysomelobia matsuzawai sp. nov. are the first Chrysomelobia species described from Asia.
Materials and methods
Examination of chrysomelid beetles representing primarily the subfamily Chrysomelinae, by Kazuyoshi Kurosa over a number of years, yielded mites belonging to the genus Chrysomelobia (Podapolipidae). Mites were removed from the abdominal tergites and under the elytra. Mites mounted on slides in Hoyer's mounting medium were placed on a heated drying tray for five days and ringed with red insulating varnish.
Measurements were taken with a Zeiss compound phase contrast microscope with a stage micrometer. Measurements are given in micrometers (pm). Alveolar vestiges of setae are designated as v. Microsetae, designated as m, are no longer than the diameter of their setal alveoli. Other terminology is based on Lindquist (1986).
The holotypes are deposited in the National Museum of Nature and Science, Tsukuba, 306-0005, Japan (NSMT). Paratypes of males, larvae and females are housed with the holotypes excepting some female, male and larval paratypes that are placed in the following museums: the A.J. Cook Arthropod Research Collection, Michigan State University, East Lansing, Michigan (CARC); The Acarology Laboratory, Museum of Biodiversity, The Ohio State University, Columbus, Ohio (OSAL); United States National Museum of Natural History, Washington, D.C. (NMNH) (mite collection housed in the USDA Systematic Entomology Laboratory, Beltsville, Maryland); Queensland Museum, South Brisbane, Australia (QMBA); Tarbiat Modares University, Tehran, Iran (TMUI); University of Michigan Museum of Zoology, Ann Arbor, Michigan (UMMZ); Tyumen State University, Tyumen, Russia (TSUR) and Zoological Museum, University of Hamburg, Hamburg, Germany (ZMH).
Description of new species
Diagnosis. All life stages. Tibia I with seta k, tarsus I with seven setae and one solenidion, setae tc′ and tc″ with blunt tips. Adult female: trachea shorter than setae v1, setae v1 slender, setae sc2 bulbous, setae c1 bulbous, setae c2 long, slender, setae e shorter than v1. Coxal setae la, 2a and 3b bulbous, apodemes I, II meeting sternal apodeme. Leg I with one claw. Femur II with minute setae d and conspicuous l′. Tibia IV with a pair of long setae, tarsus IV with a single long seta. Adult male: shield C, D, EF with 4 pairs of setae, c1, d, e minute, c2 developed, 5 long; setae c1 posterior to plane of setae c2. Plate C, D, EF with row of setae d slightly anterior to row of setae e. Genital capsule posterodorsal, shield C, D, EF with broadly concave posterior margin, setae ps1 not evident; tibiae I, II, III with spine-like setae, femur II with minute setae d and longer l′, leg IV enlarged basally, convex lateral margin, about 2/3 length of leg III, tibia III setae v″ nearly 1/2 width of idiosoma, tibia IV setae v′, v″ shorter than seta d. Legs I, II, III with two claws. Three tarsus IV setae plus a curved claw. Larva: dorsal gnathosomal setae nearly 1/2 length of dorsal gnathosomal setae in adult females.
Female (Figs. 1, 2, n=18)
Gnathosoma. Length 60–70, width 54–63 (Table 1). Cheliceral stylets 50–57 (one 46). Pharynx width 10–12. Setae ch 17–27, su 5–10. Palps longer than wide, three segmented, su-su 18–19.
Idiosoma. Length 258–319, width 208–240, setae v1 24–30, positioned on narrowed anterolateral margin of prodorsal shield and immediately posterior to stigmata, v 2 v. Setae sc2 bulbous, length 10–12, width 7–9. Idiosomal plate lengths: PD 98, C 70, D 50–58, EF 45–52; widths PD 208, C 210–220, D 190, EF 103–123, setae c1 bulbous, length 8–9, width 4–6, c2 90–127, d 7–8 (one 12), e 7–10, h1 15–25, h1-h1 53–56. Cupule ia anterolateral to setae d, cupule im anterolateral to setae e. Stigmata at anterolateral margin of prodorsal shield. Trachea length 25–28, width 5, branching not evident. Distance between setae v 1-v1 70–82, sc2-sc2 43–54, c1-c1 86–93, c1-c2 31–37, v1-sc2 38–55, v2-sc2 8–10. Venter with apodemes II meeting sternal apodeme. Coxal setae la bulbous 12 long, 7 wide; 2a bulbous, 10 long, 8 wide, 2b v, 3a 5–8, 3b bulbous, 10 long, 6 wide; 4b 6. Distance between setae la-la 41, 2a-2a 72, 3a-3a 82, 3b-3b 129.
Legs. Femur I setae l′ thick 18–20, d m, v″ 10–16, tibia I with setae l′ m, d 65–75, ϕ 11–14, k thin 12–13. Tarsus I setae tc′, tc″ eupathidial (blunt), tc′ 22–28, tc″ 20–30, solenidion ω 9–11. Femur II setae l′ 9–10, d 2. Tibia II l′ 11–13, d 20–22, v′ 28, v″ 43. Tarsus II setae pl′ 9–14, tc′ 43-46, pl″ 53, u′ 10–12, pv″ 8–9. Tibia III setae l′ 7–10, d 42, v′ 18, v″ 30. Tarsus III setae tc′ 48, pl′ 11–14, pl″ 50, u′ 8–12, pv″ 5–7. Femur and genu IV fused. Tibia IV setae v′, v″ and tarsi IV tc′ exceed 200. Tibia and tarsus IV separate. Setation for femur, genu, tibia, tarsus I, II, III, IV: 30-6(+1)-7(+1), 2-0-4-5, 0-0-4-5, 0-0-2-1.
Male (Figs. 3, 4, n=1)
Gnathosoma. Length 50, width 51. Cheliceral stylets 33, pharynx width 10, setae ch 9, su 6, su-su 19, palp length 15.
Idiosoma. Length 250, width 207, setae v1 2, v2 m, setae sc2 3, c1 m, c2 5, d m, distance between setae v1-v1 32, v2-v2 30, sc2-sc2 66, c1-c1 60, c1-c2 41, d-d 32, e-e 78. Genital capsule posterodorsal, length 35, width 50, two internal lobes interpreted as setae ps2 length10, aedeagus small. Venter with apodemes II almost reaching sternal apodeme. Coxal setae minute.
Legs. Femur I setae l′ 6, v″ 8, d m, femur II setae l′ 10, d m, no femora III, IV setae. No genua I, II, III, IV setae. Tibia I solenidion ϕ 11, slender adjacent seta k 10. Tibia I setae v′ spine-like, tibiae II, III setae l′ spine-like 10, tibia I, II, III setae d 22, 18, 15. Tarsi I setae pl′ 15, tc′ 23, tc″ 30, solenidion ω 10, setae pl″ 22, s 12, pv′ 3, pv″ 8. Tibia III setae v″ 140. Ambulacra I, II, III with two stout claws. Setation for femur, genu, tibia, tarsus of legs I, II, Ill, IV: 3-0-6 (+1)-7(+1), 2-0-4-5, 0-0-4-5, 0-0-3-3 + claw. Thickness of fused femur and genu IV 33.
Larval female (Fig. 5, n=5 exoskeletons containing adult females)
Gnathosoma. Length 43-52, width 37–48. Cheliceral stylets 45–53, pharynx width 8–10. Setae ch 9, su 9, su-su 15.
Idiosoma. Length 270–370, width 195–265, setae v1 m, v2 v, sc2 3, c1 m, c2 3, h1 55, distance setae v1-v130, v2-v2 40, sc2-sc2 63, v2-sc2 20. Distance between setae c1-c2 32, h1-h1 8.
Legs. Femur I setae l′ 5, d m, v″ 6, no genua I, II, III, IV setae. Tibia I solenidion ϕ 6, k 2, v′ 5, v″ 5, d 5. Tarsus I setae tc′ 12, tc″ 14, solenidion ω 5, setae pl″ 6. Femur II setae l′ 6, d 3. Tibia II setae v″ 26, tibia III setae v″ 70. Ambulacra I, II, III with two stout claws.
Etymology. The species is named for Dr. Haruo Matsuzawa, specialist in Chrysomelidae, who provided many potential host beetles that yielded Chrysomelobia mites for this study. The species name is a noun in the genitive case.
Type material. All specimens from Gonioctena rubripennis Baly (Coleoptera: Chrysomelidae). Holotype: adult female (Kurosa Collection No. 3321-3(3/7), Shiromana, Okutama, Tokyo, Japan, 3 May 1980, coll. K. Kurosa, deposited with the type host in the National Museum of Nature and Science, Tsukuba, Japan (NSMT). Paratypes: 5 females, 1 male, same data as holotype (KCN 3321-1 to 3321-8); 3 females, Mineoka, Kamogawa City, Chiba Pref., Japan, 4 June 1978, coll. J. Okuma; 2 females (1 slide), Mt. Odamiyama, Oda-cho, Ehime Pref., Japan, coll. E. Yamamoto; 5 females inside of exoskeletons of larval females, Kamafuga Dam, Miyaga Pref., Japan, 27 April 1995, coll. unknown; 2 females, Bizen-shi, Okayama Pref., Japan, 7-9 1989, coll. unknown. One female paratype each is deposited at CARC, OSAL, NMNH, QMBA, TMUI, TNAU, UMMZ and ZMH. The balance of paratypes is deposited with the holotype (NSMT). The balance of type hosts is deposited in UMMZ.
Differential diagnosis. The new species appears closely related to C. gimlii (Seeman & Nahrung, 2005), but differs by having females with the alveolar vestige of seta v2 situated close to seta sc2 and setae tc′-tc″ on tarsus I eupathidial (alveolar vestige of seta v2 midway between setae v1 and sc2 and setae tc′-tc″ with tapering tips in C. gimlii); and by having males without alveolar vestiges of setae sc1 and lacking extremely long setae on tibia III (alveolar vestiges of setae sc1 present and tarsus III with a very long attenuate seta in C. gimlii).
Remarks. The six species of Chrysomelobia from the Western Hemisphere (4 spp.) and Africa (2 spp.) all have setae on genua I-II, and of those six species, only C. donati lacks setae on genu III. These setal losses place C. matsuzawai sp. nov. within the radiation of 14 Australian species, plus the European species C. mahunkai Regenfuss 1968, that all lack setae on genua I-III. The distinctive bulbous setae present in several species of Chrysomelobia are expressed variously and help define species groups, as indicated in Seeman (2008). The bulbous setae in C. matsuzawai sp. nov. are sc2, c1, la, 2a and 3b, which is the same as mites in the gimlii species group (C. gimlii, C. orthagoriscus Seeman, 2008, C. pagurus Seeman, 2008). Thus, C. matsuzawai sp. nov. is similar to these species, but differs from the other species of the gimlii species group in the following features. In female C. matsuzawai sp. nov., the vestige of seta v2 is situated close to seta sc2 and setae tc′-tc″ on tarsus I are eupathidial, i.e., blunt-tipped. In species of the gimlii species group, seta v2 is in a more typical position midway between setae vl and sc2 and setae tc′-tc″ are not eupathidial, having tapering tips.
Chrysomelobia matsuzawai sp. nov. also differs from all other Australian species, excepting C. lipsettae Seeman, 2008, by having broad tracheae (width 5). The thin trachea that do not anastomose may be a synapomorphy for species of Chrysomelobia that infest eucalypt-feeding Paropsini; the host of C. lipsettae feeds on Acacia (Fabaceae), and C. lipsettae was hypothesized by Seeman (2008) to be a species intermediate between the Australian (+C. mahunkai) and the American and African species of Chrysomelobia.
Maximum measurements in micrometers (μm) for Chrysomelobia mahunkai (mah), C. nipponica sp. nov. (nip), C. matsuzawai sp. nov. (mat), C. gimlii (gim), C. pagurus (pag), C. orthagoriscus (ort), C. captivus (cap), C. alleni (all), C. lipsettae (lip). Males and larvae of C. mahunkai have not been reported.
Diagnosis. All life stages. Tibia I with seta k, tarsus I with seven setae and one solenidion, setae tc′ and tc″ with slender tips. Adult female: tracheae broad and long, anastomosing distally, setae c1 slender, setae c2 long, setae e as long as setae vl. Coxal setae la, 4a slender, 2a and 3b bulbous, femora I, II with seta l′ and minute setae d. Tibia IV with a pair of long setae, tarsus IV with a single long seta. Adult male: setae v 1, v2, d, e minute, setae sc2, cl, c2 short, two times diameter of setal acetabulum. Genital capsule posterodorsal, wider than long, setae h1, h2 minute, setae ps1 minute, setae ps2 internal, lobular. Femur I setae l′ 10, d m, v″ 10, femur II setae l′ 4, d m. Tibiae I, II, III without spine-like setae. Tibia III setae l″ shorter than tibia III setae d. Legs I, II, III with two claws. Larva: gnathosomal setae su near 1/2 length setae ch. Setae v1, v2, c1 minute, setae c2, sc2 short, two times diameter of setal acetabulum.
Female (Figs. 6, 7, n=16)
Gnathosoma. Length 60–67, width 63–76 (Table 1). Cheliceral stylets 60–66. Pharynx width 12–13. Setae ch 37–41, su 12–17. Distance between setae su-su 19–24. Palps longer than wide, two segmented.
Idiosoma. Length 275–300, width 222–289, with prodorsal shield narrow anteriorly, setae v1 25–35, v2 v, sc2 19–22, c1 7–10, c2 120–140, d 8–10, e 28–32, h1 20–25. Idiosomal plate lengths: prodorsal plate 90–98, C 70–72, D 58–60, EF 40–60; widths PD 178–180, C 223–228, D 118–130, EF 112–130, setae c1 in line with c2. Stigmata at anterolateral prodorsal shield. Length of broad trachea leading from stigmata near 80 and anastomosing distally. Stigmata-stigmata 63–70, setae v1-v1 70–76, v2-v2 55–59, sc2-sc2 112–114, c1-c1 55–58, d-d 50–52, e-e 91–100, h1-h1 56–62. Venter with apodemes II not meeting sternal apodeme. Coxal setae la slender 8–12, 2a bulbous 5–6, 3a slender 10–16, 3b bulbous 4–5, 4b 10–17. Distance between la-la 62–66, 2a-2a 106–110, 3a-3a 50-53, 3b-3b 104–112.
Legs. Femur I setae l′ thick 19–22, d m, v″ 20–29, tibia I d 82–90, ϕ 12–13, k 11–13. Tarsus I setae tc′ 35–42, tc″ 30–39, ω 10–12. Ambulacrum I with one claw. Femur II setae l′ 10-13, d m, tibia II l′ 11–20, d 39–48, v′ 22–30, v″ 34–36. Tarsus II setae pl′ 13–14, tc′ 45–53, tc″ 30-37, u′ 9–12, pv″ 6–12. Tibia III setae l′ 8–12, d 34–37, v′ 20–30, v″ 34–37, tarsus III setae pl′ 14-19, tc′ 39–48, tc″ 30–40, u′ 9–10, pv″ 7–10. Tibia IV setae v′, v″ and tarsus IV setae tc′ exceed 250. Setation for femur, genu, tibia, tarsus I, II, III, IV: 3-0-6(+1)-7(+1), 2-0-4-5, 0-0-4-5, 0-0-2-1.
Male (Figs. 8, 9, n=1)
Gnathosoma. Length 50, width 60. Cheliceral stylets 41, pharynx width 10, setae ch 19, su 7, distance su-su 21, palp length 12.
Idiosoma. Length 300, width 230, setae v1 m, v2 m, sc2 4, c1 3, c2 10, d, e, f m. Distance between setae v1-v1 32, v1-sc2 49, sc2-sc2 75, c1-c1 33, c2-c2 114, c1-c2, d-d 18, e-e 63. Genital capsule posterodorsal, length 35, width 50.
Legs. Femur I setae l′, d m, v″ 10, femur II setae l′ 4, d m. No genua I, II, III, IV setae. Tibia I setae l′ 3, d 30, ϕ 10, k 10, v1 thick 6, v″ 17, l″ 3. Tarsus I setae tc′ 27, tc″ 31, ω 8, pl′ 13, pv′ 3, s 8, pv″ 5, pl″ 13. Femur II setae l′ 4, d m, tibia II setae l′ 6, d 19, v1 18, v″ 19, tarsus II setae pl′ 8, tc′ 34, pl″ 25, pv′ 2, u′ 6, pv″ 6. Tibia III seta l′ 16. Thickness of mid femur IV 20. Ambulacra I, II, III with two claws. Setation for femur, genu, tibia, tarsus I, II, III: 3-0-6(+1)-7(+1), 2-0-4-5, 0-0-4-5, 0-0-2-2+ claw.
Larval female (Figure 10, n=1, exoskeleton very pale)
Gnathosoma. Length 49, width 52. Cheliceral stylets 50, pharynx width 11. Setae ch 12, su 7, su-su 9.
Idiosoma. Length 300, width 230. Setae v1 m, v2 m, sc2 4, c1 m, c2 4, h1 m. Distance v1-v1 29, v2-v2 34, v2-sc2 25, sc2-sc2 58, c1-c2 20.
Legs. Femur I seta l′ 4, d m, v″ 10. Tibia I l′ m, d 28, ϕ 10, k 8, v′ 4, v″ 12, l″ m. Tarsus I tc′ 22, tc″ 22, ω 9, pl′ 9, pv′ 3, s 5, pv″ 3, pl″ 8. Femur II seta l′ 5, d m, Tibia II l′ 3, d 7, v′ 8, v″ 14. Tarsus II tc′ 16, pl″ 23, u′ 8, pv″ 4. Tibia III l′ 2, v′ 12, v″ 17. Tarsus III pl′ 10, pl″ 20, tc′ 12, u′ 5, pv″ 3. Setation for femur, genu, tibia, tarsus I, II, III: 3-0-6(+1)-7(+1), 2-0-4-5, 0-0-4-5.
Etymology. The specific name nipponica is an adjective derived from the country of origin, Japan (Nippon).
Type material. All specimens from Gonioctena rubripennis Baly (Coleoptera: Chrysomelidae). Holotype: adult female (RWH25 VIII 2015-2), Katsuura, Chiba Pref., Japan, 22–24 IV 1989, coll. K. Kurosa, deposited with the type host in the National Museum of Nature and Science, Tsukuba, Japan. Paratypes: 1 male, 12 females, 1 larva, same data as holotype; 2 females, Hongo-Cho, Aizu-Wakamatsu, Fukushima Pref., Japan, 26–27 V 1999, coll. unknown; 1 male, Minakami-machi, Gunma Pref., Japan, 1 IV 1999, coll. unknown; 1 female, Fujikawachi, Umemachi, Saiki-shi, Oita Pref., Japan, 3 VI 2012, coll. S. Sasaki; 1 female, Nano-shi, Yamagata Pref. Japan, 23 VI 1999, coll. unknown. 1 female paratype is deposited at each of the following CARC, OSAL, NMNH, QMBA, TMUI, TNAU, TSUR, UMMZ, ZMH. Balance of paratypes is deposited with the holotype (NSMT). Balance of type hosts deposited in UMMZ.
Differential diagnosis. The new species appears closely related to C. mahunkai, but differs by having females with one ambulacral claw (two in all other Chrysomelobia) and seta d on femur II (absent in C. nipponica sp. nov.).
Remarks. The host genus for both new species, Gonioctena (Coleoptera: Chrysomelidae), is also a host for the type species of Chrysomelobia, C. mahunkai. New adult female specimens of C. mahunkai were collected from an unspecified locality in Germany from Gonioctena sp. and the holotype female was also examined. An illustration of the holotype female loaned by Dr. Hieronymus Dastych of the University of Hamburg, Germany is provided (Fig. 11). Previously, the species was recorded from a single female collected from Tansey beetle Chrysolina graminis (L., 1758) (= Chrysomela graminis). Of the two new species, C. matsuzawai sp. nov. is not closely related to C. mahunkai, but in contrast, C. nipponica sp. nov. shares several similar character states with C. mahunkai. These similarities are the bulbous coxal setae 2b and 3b, dorsal setae and seta la unmodified, the female tibia IV with two setae and tarsus IV with one terminal seta, and the broad tracheae that anastomose distally. With the exception of the last character state, these states are also shared with some Australian species, particularly the husbandi species group. Female C. mahunkai and C. nipponica sp. nov. are distinct from the husbandi species group by having well-developed setae la (they are minute in the husbandi species group). Male C. nipponica sp. nov. differ from the husbandi species group by their large leg IV that bears a terminal claw (leg IV is diminutive and lacks a claw in the husbandi species group). Female C. nipponica sp. nov. are distinguished from C. mahunkai by the presence of two ambulacral claws on leg I in C. mahunkai (one in all other Chrysomelobia) and the presence of seta d on femur II in C. nipponica sp. nov. (absent in C. mahunkai). The male and larval stages for C. mahunkai remain unknown, so cannot be compared with C. nipponica sp. nov. , but we anticipate males and larvae of C. matsuzawai sp. nov. and C. nipponica to be similar. The divided plate C in the larva is absent in all Australian species but is present in C. eickworti Husband & OConnor, 2004, C. labidomerae Eickwort, 1975 and C. peruviensis Husband and Moraes, 1999 and may be present in C. nipponica sp. nov.
Key to species of Chrysomelobia
1. Female & male: at least 1 seta on genua I, II and IV and femur IV 2
- Female & male: setae absent on genua I-IV and femur IV 7
2(1). Female: genu I with 3 setae; femur II with 1 seta; femur III without setae C. donati Haitlinger
- Female & male: genu I with 4 setae; femur II with 3 setae; femur III with 2 setae 3
3(2). Female: coxal seta 4b absent. Male: tarsus I without setae ft′ and ft″ C. elytrosphaerae Fain
- Female: coxal seta 4b present. Male: tarsus I with at least l ft seta 4
4(3). Female: genu IV with 2 setae (v″ present). Male: with 4 prodorsal setae or vestiges of setae (sc1 present); post-genital shield posterior to genital capsule expansive; fused telofemur-genu IV with 2 setae C. eickworti Husband & OConnor
- Female: genu IV with 1 seta (v″ absent). Male: with 3 prodorsal setae or vestiges of setae (sc1 absent); post-genital shield posterior to genital shield elongate; fused telofemur-genu IV with 1 seta 5
5(4). Female: Cheliceral stylets 37–46. Male: idiosomal plates reticulate; tibia III, seta v″ short (< 50) C. oneili Moraes, Husband & Lofego
- Female: Cheliceral stylets > 50. Male: idiosomal plates smooth; tibia III, seta v″ long (> 70) 6
6(5). Female: seta h1 30–40. Male: ventral gnathosomal setae 18–23; seta sc2 close to margin of prodorsal shield; genu IV with 1 seta (v′ present) C. peruviensis Husband & Moraes
- Female: seta h1 17–19. Male: ventral gnathosomal setae 10–13; seta sc2 well within margin of prodorsal shield; genu IV without setae (v′ absent) C. labidomerae Eickwort
7(1). Female: tarsus IV with 2 or 3 long terminal setae. Male: seta c2 minute 8
- Female: tarsus IV with 1 long terminal seta. Male: seta c2 developed, > 3 long 14
8(7). Female: seta sc2, c2, la, 2a and 3b slender 9
- Female: seta sc2, c2, la, 2a and 3b bulbous 10
9(8). Female & male: seta 3a absent C. vafer Seeman
- Female & male: seta 3a present C. verecundus Seeman
10(8). Female & male: tibia II lacking seta l′. Female: tibia and tarsus IV partially or completely fused. Male: dorsal shield C-D-E divided or with folds marking a weak division; tarsus IV, setae u′ and pv″ absent C. armstrongi Seeman
- Female & male: tibia II with seta l′. Female: tibia and tarsus IV separate. Male: dorsal shield C-D-E entire; tarsus IV, setae u′ and pv″ present 11
11(10). Female & male: tibia IV with 1 seta (v″ present) C. alipilus (Seeman & Nahrung)
- Female & male: tibia IV without setae (v″ absent) 12
12(11). Female: tarsus IV with 3 terminal setae; setae d and e < 40 C. nahrungae Seeman
- Female: tarsus IV with 2 terminal setae; setae d and e > 45 13
13(12). Female: intercoxal setae further apart (la-la 37, 2a 62–65). Male: tarsus IV with claw, u′, and 3 setae (minute seta pv′ absent). Larva: setae sc2 27–31 and c2 25–29 long C. alleni Seeman & Nahrung
- Female: intercoxal setae closer together (la-la 26–31, 2a 47–51). Male: tarsus IV with claw, u′, and 4 setae (minute seta pv′ present). Larva: setae sc2 13–18 and c2 13–18 long C. aquariolus Seeman
14(7). Female: coxal setae 2a and 3b slender C. lipsettae Seeman
- Female: coxal setae 2a and 3b bulbous or minute 15
15(14). Female: seta la slender; tracheae broad, anastomosing distally 16
- Female: seta la minute or bulbous; tracheae narrow, not anastomosing distally 17
16(15). Female: ambulacra with 2 claws; femur II without seta d C. mahunkai Regenfuss
- Female: ambulacra with 1 claw; femur II with seta d C. nipponica sp. nov.
17(15). Female: seta sc2, c1 and la bulbous. Male: tarsus IV with terminal claw, with 3–4 setae 18
- Female: seta sc2 and c1 slender; seta la minute. Male: tarsus IV lacking terminal claw, with 1–2 setae 21
18(17). Female: vestigial seta v 2 closely associated with seta sc2; tarsus I setae tc′-tc″ eupathidial (blunt-tipped). Male: plate C-D-EF with 5 pairs of setae; post-sternal apodeme well-developed C. matsuzawai sp. nov.
- Female: vestigial seta v 2 not closely associated with seta sc2, about half way between setae v′ and sc2; tarsus I setae tc′-tc″ not eupathidial (tips tapered). Male: plate C-D-EF with 4 pairs of setae; post-sternal apodeme developed or absent 19
19(18).Female: seta sc2 and c1 5–7 long, 4–5 wide, with mediolateral projection 4–5 long (if broken then obvious stub present). Male: tibia II, seta v″ 54–58, tarsus III, seta tc′ 43–47 C. orthagoriscus Seeman
- Female: seta sc2 8–12 long, 5–6 wide; seta c1 7–10 long, 5.5–7 wide, mediolateral projection absent or a minute stub. Male: tibia II, seta v″ either < 40 or > 80 long, tarsus III, seta tc′ either < 35 or > 50 long 20
20(19).Female: distance between setae v1-sc2 52–60, v2-sc2 26–31. Male: seta c2 10–13; tibia II, seta v″ 34–39; tarsus III, seta tc′ 50–56 C. gimlii (Seeman & Nahrung)
- Female: distance between setae v 1-sc2 40–44, v 2 -sc2 12–21. Male: seta c2 5–9; tibia II, seta v″ 80–140; tarsus III, seta tc′ 31–34 C. pagurus Seeman
21(17).Female: setae 2a and 3b bulbous 22
- Female: no coxal setae bulbous . 23
22(21).Female: setae 2a and 3b 5–6 long, 3–4 wide; distance between h1-h1 31–41. Male: tarsus II, seta tc″ < 50 C. husbandi (Seeman & Nahrung)
- Female: setae 2a and 3b 6–7.5 long, 4.5–5 wide; distance between h1-h1 22–29. Male: tarsus II, seta tc″ > 80 C. captivus (Seeman & Nahrung)
23(21).Female & male: femur II without setae C. cubile Seeman
- Female & male: femur II with minute seta 24
24(23).Female: seta v 2 vestigial but distinct; setae h1 length 33–43 C. intrusus Seeman & Nahrung
- Female: seta v 2 absent; setae h1 length 15–19 C. lawsoni (Seeman & Nahrung)
We are thankful for assistance with obtaining potential host Chrysomelidae for Podapolipidae and advice by Barry OConnor and Mark O'Brien of the University of Michigan Museum of Zoology, Ann Arbor, Michigan and for technical assistance from Adrian College librarians Richard Geyer, David Cruse and Noelle Keller.Our cordial thanks are also due to the following Japanese entomologists: Takeshi Matsuzawa, Jun Okuma, Shigeo Sasaki and Eiji Yamamoto.