Sheltered life beneath elytra: three new species of Eutarsopolipus (Acari, Heterostigmatina, Podapolipidae) parasitizing Australian ground beetles Author Katlav, Alihan Hawkesbury Institute for the Environment, Western Sydney University, Locked Bag 1797, Penrith, NSW 2751, Australia Author Hajiqanbar, Hamidreza Department of Entomology, Faculty of Agriculture, Tarbiat Modares University, 14115 - 336 Tehran, Iran Author Riegler, Markus Hawkesbury Institute for the Environment, Western Sydney University, Locked Bag 1797, Penrith, NSW 2751, Australia & Queensland Museum, PO Box 3300, South Brisbane 4101, Australia text Parasite 2021 Paris, France 2021-11-05 28 75 1 21 http://dx.doi.org/10.1051/parasite/2021069 journal article 10.1051/parasite/2021069 1776-1042 PMC8570142 34738903 12524440 7CFD69B7-066F-41F7-B37F-53C7CEDDFE39 Eutarsopolipus chlaenii Katlav & Hajiqanbar n. sp. ( Figs. 10–11 ) urn:lsid:zoobank.org:act: 25276820-D40C-4F2F-AAA2- E68575A38719 Type material: Total material recovered : ♀ ( n = 4), larval ♀ ( n = 16), ex. under elytra, on the base of membranous hind wing of specimens of Chlaenius flaviguttatus Macleay, 1825 ( Coleoptera : Carabidae : Harpalinae: Chlaeniini) ( Fig. 13 ). Three out of four collected host specimens found parasitised. Beetles specimens were collected at three independent events on 24 Feb 2020 , 26 Feb 2020 , and 28 Feb 2020 . Holotype : adult female ( ANIC 52-003965), ex. under elytra, on the base of membranous hind wing of C. flaviguttatus ; Coll. Shams Paryav; 24 Feb 2020 . Paratypes : adult female ( n = 3), larval female ( n = 5), same data as holotype ( 24 Feb 2020 , 26 Feb 2020 , and 28 Feb 2020 ). Type locality : Loc. Vines Drive, Hawkesbury Campus , Western Sydney University , Richmond , NSW, 33°36 ļ 45.6 ļļ S 150°44 ļ 40.2 ļļ E . Deposition of material : The holotype , one adult female and 2 larval female paratypes are deposited at ANIC ( ANIC 52-003965 - 68 ). 1 adult female and 2 larval female paratypes are deposited at QM ( QMS 117009-10 , 117042 ) . The remaining paratypes ( TMU SP-20200224 , 1–3 ) , 11 non-type larval females and the host beetle specimen are deposited at AC-DE-TMU. Etymology : The species epithet “ chlaenii ” refers to the generic name of the carabid host beetle Chlaenius flaviguttatus . Authorship : Note that the authors of the new taxon are different from the authors of this paper; Article 50.1 and Recommendation 50A of International Code of Zoological Nomenclature [ 24 ]. Adult female ( Fig. 10 ) (n = 4) Gnathosoma ( Figs. 10a–10b ). Length 45 (43–45), width 42 (40–42); cheliceral stylets length 28 (29–30); pharynx length 12 (12–13), pharynx width 12 (12–13); ch 15 (15–16), pointed; su 6 (5–6), needle-like; distance between setae ch–ch 27 (27–29), su–su 17 (16–17). Idiosoma ( Figs. 10a–10b ). Length 230 (225–240), width 185 (165–185). Idiosomal dorsum ( Fig. 10a ). Respiratory system (stigmata and tracheae) present, stigmata stalked; all dorsal setae pointed; prodorsal plate (PrS) with setae v 1 6 (5–6), setae v 2 vestigial, setae sc 1 5 (5–6), sc 2 42 (38–41). Plate C setae c 1 7 (6–7), c 2 5 (6–6). Plate D setae d 5 (5–6); cupuli ia evident, anterolaterad setae d . Plate EF setae f 7 (6–7); cupuli im evident, anterolaterad setae f . Plate H not evident, setae h 7 (7–8). Distances between setae: v 1 – v 1 35 (34–37), v 2 – v 2 42 (41–44), v 1 – v 2 13 (13–14), sc 1 – sc 1 61 (57–60), v 1 – sc 1 18 (18–19), sc 2 – sc 2 62 (58–61), sc 1 – sc 2 39 (38–40), c 1 – c 1 61 (57–64), c 1 – c 2 48 (42–46), d–d 59 (57–58), f–f 37 (33–36). Idiosomal venter ( Fig. 10b ). All coxal plates smooth; all coxal setae pointed; ap1–2 and appr well developed, ap2 reaching to appr; apsej absent; coxisternal field I with setae 1 a 3 (2–2); alveoli of vestigial setae 1 b not evident; coxisternal field II with 2 a 2 (2–2); alveoli of vestigial setae 2 b evident; coxisternal field III with subequal setae 3 a 7 (7–8) and 3 b 7 (8–8). Distances between setae: 1 a –1 a 19 (20–22), 2 a –2 a 27 (25–26), 3 a –3 b 19 (24–26). Legs ( Figs. 10c–10e ). Setal formula for legs I–III (femurtarsus): 2-0-6(+ φ )-8(+ ω ), 0-0-4-6(+ ω ), 0-0-4-6. Ambulacrum I with a well-developed sickle-shaped claw, ambulacrum II–III each with a pair of well-developed claws. Leg I ( Fig. 10c ): femur, d 3 (2–2), slightly thickened, seta l ļ 16 (15–16) thick and blunt-ended; tibia with φ 5 (5–5) baculiform, d 28 (29–31), l ļ 4 (4–5), l ļļ 3 (3–3), v ļ 5 (5–6) stiff, v ļļ 9 (8–9), seta k 5 (5–6); tarsus I, ω 3 (3–4) digitiform, eupathidial setae tc ļ 9 (8–9) and tc ļļ 10 (9–10) distinctly blunt-ended, pl ļ 9 (8–9), setae u ļļ 2 (2–3), pv ļ 3 (2–2) and pv ļļ 2 (2–2) subequal, seta s 5 (5–5) spine-like, with a blunt tip, p ļ 1 (1–2). Leg II. ( Fig. 10d ): tibia, d 8 (7–8), l ļ 7 (6–7), v ļ 11 (10–11), v ļļ 6 (6–6); tarsus, ω 4 (3–4) digitiform, tc ļ 7 (6–7), setae u ļ 6 (6–6) and tc ļļ 5 (5–6) spine-like, pl ļļ 17 (17–18), pv ļļ 3 (3–3), u ļļ 2 (2–2). Leg III ( Fig. 10e ): tibia, setae d 7 (7–8), l ļ 6 (6–7), v ļ 11 (10–12), v ļļ 6 (5–6); tarsus, tc ļ 10 (10–10), setae u ļ 6 (6–7) and tc ļļ 6 (5–6) spine-like, pl ļļ 16 (15–16), pv ļļ 3 (3–3), uļļ 2 (2–2). 14 Table 2. Comparison of selected characters (range of measurements if available) of all the described species of the leytei species group in the genus Eutarsopolipus (male is unknown for E. leytei ); species abbreviated as E. pulcher n. sp. ( Ep ), E. leytei ( El ) , E. dastychi ( Ed ) , E. orpheus ( Eo ) , E. biuncatus ( Eb ) , and E. janus ( Ej ) .

Life stage	Female		Male	Larval female
Character	Ep	El	Ed	Eo	Eb	Ej	Ep	Ed	Eo	Eb	Ej	Ep	El	Ed	Eo	Eb	Ej
Gn. L.	42–50	78	47–50	50–57	44–46	52–57	25–36	30–33	30–32	28–31	28–37	29–32	56	37–40	32–38	30–34	31–34
Ch. S. L.	44–51	68	47–48	23–28	35–38	45–51	17–19	23–26	15	17–18	25–28	28–34	54	35–40	21–27	25–27	36–42
Setae ch	18–25	20	27–30	10–18	14–22	25–35	8–12	12–15	m	1–2	3–4	20–25	20	30–36	31–35	19–23	26–33
Setae su	12–14	3	22	11–15	8–12	15–20	9–10	9–10	6–7	4–6	8–9	10–12	3	15–18	6–7	8–10	12–15
Setae v 1	11–14	5	36–45	17–26	8–10	19–24	m	5–10	m	m–2	4–9	11–13	m	42–50	26–32	12–14	15–18
Setae sc 1	9–13	11	25	17–29	7–9	9–11	m	5–13	m	m	m–2	10–12	10	25–28	28–33	5–6	8–10
Setae sc 2	51–60	62	58–70	47–59	15–20	36–40	34–46	60–70	38–40	2–3	27–34	62–72	78	94–101	90–95	51–52	70–80
Setae c 1	4–6	12	18–19	8–10	5–7	8–9	m	10	m	m	2–3	7–8	10	18–22	14–18	5–6	7–10
Setae c 2	4–6	9	17–21	9–18	4–6	8–10	m	5–7	m	m	2	5–7	9	17–21	10–15	3–4	7–10
Setae d	4–5	12	18–21	11–18	4–5	9–10	m	8–10	m	m	m–2	6–7	9	20–23	15–17	4–5	7–9
Setae f	5	8	22	10–11	4–5	7–9	m	3–4	m	m	m–2	7–9	8	16–18	8–12	4–5	6–8
Setae h 1	–	–	–	–	–	–	m	m	m	m	m	64–67	148	65–90	55–64	65–70	100
Setae h 2	–	–	–	–	–	–	–	–	–	–	–	1–2	67	m	m	3–4	4–6
Setae 1 a	5–6	~4	5–6	4–5	5–6	6–7	m	2–5	m	2–3	4–5	5–7	~2	8–10	6–8	2–3	4–5
Setae 2 a	5–7	~4	5	3–4	4–5	6–7	m	3–4	m	2–3	4–5	4–6	~2	7–10	8–9	3–4	4–5
Setae 3 a	4–5	8	9	2–3	5–6	6–9	m	~2	m	3	5–6	5–6	4	10–13	9–11	4–5	7–8
Setae 3 b	5–6	4	7	3	4–5	6–7	m	~4	m	2–3	5–6	5	2	10–12	9–10	3–4	5–6
Sol. Ta I ω	4–5	~2	4–5	5–6	3–4	4–5	4–5	5–7	4	3	3–4	4–5	~2	4–5	4–5	3–4	4–5
Sol. Ti I φ	7–9	7	10	7–8	6–7	6–8	6–8	5–6	5–6	4–5	6–7	7–8	7	7–9	8–9	6–7	7–8

A. Katlav et al.: Parasite 2021, 28 , 75 Figure 9. Key to the world species of Eutarsopolipus in the leytei group (based on adult females). Male (Unknown) Larval female ( Fig. 11 ) (n = 5) Gnathosoma ( Figures 11a–11b ). Length 39–46, width 38–40; cheliceral stylets length 29–32; pharynx length 13–15, pharynx width 10–11; ch 17–19 pointed; su 3–4 needle-like; distance between setae ch–ch 22–26, su–su 12–14. Idiosoma ( Figs. 11a–11b ). Length 165–195, width 125–145. Idiosomal dorsum ( Fig. 11a ). All dorsal setae needle-like except sc 2 attenuate; PrS with setae v 1 6–7, v 2 vestigial, sc 1 6–7, sc 2 95–98. Plate C setae c 1 5–6, c 2 5–6. Plate D setae d 6–7; cupuli ia anterolaterad setae d . Plate EF setae f 7–8; cupuli im anterolaterad setae f . Plate H situated ventrally with setae h 1 97–101 and h 2 22–24. Distances between setae: v 1 – v 1 24–26, v 2 – v 2 44–47, v 1 – v 2 12–14, sc 1 – sc 1 59–62, v 1 – sc 1 20–22, sc 2 – sc 2 57–58, sc 1 – sc 2 40–42, c 1 – c 1 83–86, c 1 – c 2 21–23, d–d 34–36, f–f 28–29. Idiosomal venter ( Fig. 11b ). All coxal plates smooth; all coxal setae tiny and pointed; ap1–2 and apsej evident; coxisternal fields I–II each divided from its pair, with setae 1 a 1–1; alveoli of setae 1 b not evident; coxisternal field II with 2 a 2–3; alveoli of setae 2 b not evident; coxisternal field III widened, with setae 3 a 7–9 and 3 b 7–8 subequal. Distances between setae: 1 a –1 a 22–34, 2 a –2 a 28–30, 3 a –3 b 21–22. Legs ( Figs. 11c–11e ). Setal formula for legs I–III (femurtarsus): 2-0-6(+ φ )-7(+ ω ), 0-0-4-6(+ ω ), 0-0-4-6. Ambulacrum I with a small bifid claw, ambulacrum II–III each with a pair of tiny claws. Leg I ( Fig. 11c ): femur, d microseta, seta l ļ 4–5 stiff; tibia, φ 3–4 baculiform, d 17–21, seta l ļ 4–5 slightly thickened, l ļļ 2–3, v ļ 3–4, seta v ļļ 4–5 slightly thickened and blunt-ended, seta k 2–3; tarsus, ω 3–4 digitiform, eupathidial setae tc ļ 7–8 and tc ļļ 7–8 subequal, distinctly blunt-ended, pl ļ 6–6, setae pv ļ 1–2, pv ļļ 2–3, seta s 4–5 blunt spur-like, p ļ 1–1; u ļļ not visible. Leg II. ( Fig. 11d ): tibia, d 4–5, l ļ 6–7, v ļ 6–7, v ļļ 4–6; tarsus, ω 2–3 digitiform, tc ļ 5–7, setae u ļ 4–5 and tc ļļ 5–6 blunt spur-like, pl ļļ 10–12, pv ļļ 2–2, u ļļ 2–2. Leg III ( Fig. 11e ): tibia, d 5–6, l ļ 6–7, v ļ 6–8, v ļļ 3–5; tarsus, tc ļ 6–7, setae u ļ 5–6 and tc ļļ 6–7 blunt spur-like, pl ļļ 10–13, pv ļļ 3–4, u ļļ 1–1. Differential diagnosis The new species belongs to a subgroup of the myzus species group that shares a combination of the following characters in adult females: ambulacrum I claw well-developed, idiosoma without lateral bulges or posteriorly without wrinkled lobes, shield C not divided, femur I seta l ļ developed (not microseta), and cheliceral stylets less than 35 µm long [ 13 ]. This assemblage includes E. chlaenii n. sp. and four other species: E . steveni Khaustov, 2010 , E. anichtchenkoi , Hajiqanbar & Mortazavi, 2012 , E . gombrooni Hajiqanbar & Mortazavi, 2019 , and E . oconnori Hajiqanbar & Mortazavi, 2019 . Among these species, E. chlaenii n. sp. is more similar to E. anichtchenkoi and E . oconnori by having setae h and f subequal. However, it is readily distinguishable from E. anichtchenkoi by having developed setae v 1 , sc 1 , 1 a and 2 a (adult female with microsetae v 1 , sc 1 , 1 a and 2 a in E. anichtchenkoi ), sc 2 almost five times longer than h 1 (adult female with sc 2 at least nine times longer than h 1 in E. anichtchenkoi ) and tarsus III with six setae (tarsus III with seven setae in E. anichtchenkoi ). Eutarsopolipus chlaenii n. sp. also differs from E . oconnori by having shorter distances c 1 - c 1 , d-d , f-f in the adult female (64, 59, 37 vs. 101, 113, 86, respectively, in E . oconnori ) and longer setae su and h 1 and cheliceral stylets in larval females (101, 4, 32, vs. 61, m, 26, respectively, in E . oconnori ). All the important characters among these five species of the myzus species group are compared for all life stages (excluding E. chlaenii n. sp. with unknown male) in Table 3 . Among adult females of the myzus species group with a strong claw on ambulacrum I, lateral bulges or posterior wrinkled lobes and entire shield C, six species have short cheliceral stylets (less than 35 µm long). The key to this subgroup is presented in Figure 12 .