Sheltered life beneath elytra: three new species of ...
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Sheltered life beneath elytra: three new species ofEutarsopolipus (Acari, Heterostigmatina, Podapolipidae)parasitizing Australian ground beetles
Alihan Katlav1,* , Hamidreza Hajiqanbar2,a, Markus Riegler1 , and Owen D Seeman3
1 Hawkesbury Institute for the Environment, Western Sydney University, Locked Bag 1797, Penrith, NSW 2751, Australia2 Department of Entomology, Faculty of Agriculture, Tarbiat Modares University, 14115-336 Tehran, Iran3 Queensland Museum, PO Box 3300, South Brisbane 4101, Australia
Received 21 June 2021, Accepted 4 October 2021, Published online 5 November 2021
Abstract β In this study, we conducted a summer sampling of carabid beetles in eastern Australia to identify theirassociated parasitic mites. Here, we describe three new species of the genus Eutarsopolipus from under the elytra(forewings) of three native carabid species (Coleoptera: Carabidae): Eutarsopolipus paryavae n. sp. (pterostichi group)from Geoscaptus laevissimus Chaudoir; Eutarsopolipus pulcher n. sp. (leytei group) from Gnathaphanus pulcher(Dejean); and Eutarsopolipus chlaenii n. sp. (myzus group) from Chlaenius flaviguttatus Macleay. We further providean identification key of the world species of pterostichi and leytei species groups as well as closely related species ofthe myzus group possessing similar characters including short cheliceral stylets. The significant diversity ofEutarsopolipus recovered here suggests that the current knowledge about Australian podapolipid mites (speciallyEutarsopolipus) is still in its infancy and deserves further study.
Key words: Acari, Biodiversity, Trombidiformes, Carabidae, Symbiosis, Systematics.
Resume β Vivre Γ lβabri sous les Γ©lytres : trois nouvelles espΓ¨ces dβEutarsopolipus (Acari, Heterostigmatina,Podapolipidae) parasitant des carabes australiens. Dans cette Γ©tude, nous avons effectuΓ© un Γ©chantillonnageestival de carabes dans lβest de lβAustralie pour identifier leurs acariens parasites associΓ©s. Nous dΓ©crivons troisnouvelles espΓ¨ces du genre Eutarsopolipus sous les Γ©lytres (ailes antΓ©rieures) de trois espΓ¨ces de carabes indigΓ¨nes(Coleoptera : Carabidae) : Eutarsopolipus paryavae n. sp. (groupe pterostichi) de Geoscaptus laevissimus Chaudoir,Eutarsopolipus pulcher n. sp. (groupe leytei) de Gnathaphanus pulcher (Dejean) et Eutarsopolipus chlaenii n. sp.(groupe myzus) de Chlaenius f laviguttatus Macleay. Nous fournissons en outre une clΓ© dβidentification des espΓ¨cesmondiales des groupes dβespΓ¨ces pterostichi et leytei ainsi que des espΓ¨ces Γ©troitement apparentΓ©es du groupemyzus possΓ©dant des caractΓ¨res similaires, y compris des stylets chΓ©licΓ©raux courts. La diversitΓ© importante desEutarsopolipus collectΓ©s ici suggΓ¨re que les connaissances actuelles sur les acariens podapolipidΓ©s australiens(en particulier Eutarsopolipus) en sont encore Γ leurs balbutiements et mΓ©ritent une Γ©tude plus approfondie.
Introduction
Beetles are among themost successful animals on the planet,accounting for about 25% of described species [10, 46]. Theirsuccess is partly attributed to their modified, sclerotizedforewings, known as elytra, that protect their body against phys-ical damage, desiccation, predation and thermal stress, enablingthem to occupy a wide range of ecological niches [33, 48]. Thesubelytral space serves as a suitable microhabitat for a broadrange of organisms such as mites, pseudoscorpions and
nematodes that occupy this niche temporarily or perma-nently [6, 36, 37]. Some mites have evolved to be permanentectoparasites in the subelytral spaces of beetles, imbibing beetlehaemolymph using piercing stylets [2, 7]. This parasitic associ-ation sometimes occurs in one part of a miteβs life cycle. Forexample, in Parasitengona (Acariformes: Prostigmata), larvaeare parasites of many insects and are sometimes found underthe elytra of terrestrial and aquatic beetles, while the nymphsand adults are free-living predators of immature stages of smallarthropods [51, 52]. However, some taxa represent evolutionarytransitions from phoresy towards parasitism, as in a fewcanestrinioid mites (Astigmata) in which deutonymphs remainphoretic on the thoracic venter of some carabid beetles, whereasthe other stages (feeding stages) are subelytral parasites of thesame hosts [15, 49]. Some other groups are real parasites with
*Corresponding author: [email protected]. Hamidreza Hajiqanbar died on October 18, 2021, at the age of 48,when this paper was in press. This article is dedicated to the memory ofDr. Hamidreza Hajiqanbar, who cherished research as his life joy andmade a major contribution to the world's insect-associated mites.
Parasite 28, 75 (2021)οΏ½A. Katlav et al., published by EDP Sciences, 2021https://doi.org/10.1051/parasite/2021069
Available online at:urn:lsid:zoobank.org:pub:7CFD69B7-066F-41F7-B37F-53C7CEDDFE39www.parasite-journal.org
This is an Open Access article distributed under the terms of the Creative Commons Attribution License (https://creativecommons.org/licenses/by/4.0),which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited.
OPEN ACCESSRESEARCH ARTICLE
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their abbreviated life stages all occurring on the hostβs bodysurface [47]. Such subelytral parasitic associations with beetleshave evolved independently in many Astigmata (e.g. mostmembers of Canestrinioidea), Mesostigmata (e.g. severalmembers of Gamasina) and Prostigmata (e.g. several membersof Raphignathina and Heterostigmatina) [14, 16, 35, 38].
The cohort Heterostigmatina (Acariformes: Prostigmata) isa large group of morphologically diverse mites, among whichnumerous species are associated with arthropods [49]. Somespecies are subelytral symbionts of various beetles with theirassociations varying from facultative or obligate phoresy toparasitoidism or parasitism [25, 28, 30, 31]. Several speciesare potential biocontrol agents against pest beetles. For exam-ple, the species of the families Pyemotidae and Acarophenaci-dae are known as insect ectoparasitoids, with the former mostlyattacking juvenile stages of bark beetles and stored-productbeetles and the latter egg ectoparasitoids of various beetlefamilies [8, 25, 29, 50].
All members of the family Podapolipidae are specializedobligate external (and rarely internal) parasites of various insects[18], among which at least 20 genera are subelytral ectoparasitesof different beetle families, mainly Carabidae, Chrysomelidae,Coccinellidae, and Scarabaeidae [21, 23, 38, 45]. These mitesare sexually transmitted, i.e. the motile stages of the mite (larvaeor adult females) move from one host individual to anotherduring copulation [17]. Parasitisation with these mites cannegatively affect host fitness. For example, in some ladybirds,individuals parasitised with Coccipolipus suffer lower fecundityand egg viability [17] and sometimes reduced longevity [40].Beyond this, these mites canmodify host sexual and behaviouraltraits to boost their transmission success among individual hosts[1]. For example, in the milk weed leaf beetle, males parasitizedby Chrysomelobia tend to more frequently contact other males,and are more successful in mating competition compared tounparasitised males; and this facilitates the miteβs higher trans-mission rate [1].
Four genera of Podapolipidae are exclusively associatedwith carabid beetles: Dorsipes (22 species), Eutarsopolipus(99 species), Ovacarus (3 species) and Regenpolipus (5 species)[11, 13, 19, 26, 27, 44]. Apart from Ovacarus, which is anendoparasite of the reproductive tracts of some carabids, the restare subelytral ectoparasites [11]. Species of Eutarsopolipus areversatile in morphology and are currently grouped into tenspecies groups [42]. Most of the species are specific to a singlehost species. However, a few parasitize more than one hostspecies [41] or more rarely more than one genus [26], yet thepossibility of them being cryptic species remains untested.More interestingly, in some cases more than one species canparasitize one host species [42] and sometimes they are special-ized to different microhabitats such as the elytral cavity, onhindwings or on the dorsal abdomen of their host [39].
Australia is anticipated to harbour rich Eutarsopolipus faunagiven its large diversity of carabid beetles [5]. This is inferredfrom small sampling efforts that have recently been conductedin some regions, and yet that discovered a considerable numberof new species [31, 41β44]. Here, we describe three new speciesof Eutarsopolipus belonging to three different species groups(leytei, myzus, pterostichi) from three native Australian carabidbeetles, raising the total number of Australian Eutarsopolipus
to 30 species. All these species were recovered following aminimal sampling effort at one site, again corroborating thehypothesis that Australia is home to diverse podapolipid faunaawaiting discovery.
Materials and methods
Carabid host beetles were collected at night on the ground,near an outdoor LED solar light lamp in Richmond, New SouthWales, in February 2020. The subelytral area of the beetles(preserved in 75β80% ethanol) was subsequently examinedfor mite infestation. Mite specimens were cleared in a mixtureof Nesbittβs fluid and a small amount of glycerine slidemounted in Hoyerβs medium. Mite morphology was studiedusing a light microscope (Olympus BX51) equipped with phasecontrast illumination. Mites from Queensland specimens of thecarabid host Gnathaphanus pulcher were removed from driedbeetles as described in Seeman [42] and examined using aNikon 80i microscope equipped with differential interferencecontrast. All measurements are given in micrometres forholotypes and the range of measurements for five selected para-types (in parentheses), if available. Distances between setaewere measured from the base of one seta to the other; setae withtheir acetabulum remnant only were categorised as vestigialsetae and those with their setae not extending past the acetabu-lum as microsetae (m). Terminology and setal notation wereadapted from Lindquist [32]. The species group assignmentfollows that of Seeman [42]. Host beetles were all identifiedwith the help of Geoff Monteith.
Abbreviations
ap apodemappr prosternal apodemeapsej sejugal apodemeQM Queensland Museum, QLD, AustraliaANIC Australian National Insect Collection; Canberra,
ACT, AustraliaAC-DE-TMU The Acarological Collection, Department of Ento-
mology, Faculty of Agriculture, Tarbiat ModaresUniversity, Tehran, Iran
Results
Family Podapolipidae Ewing, 1922Genus Eutarsopolipus Berlese, 1913Type species: Tarsopolipus lagenaeformis Berlese, 1911,
by original designation.Species group: pterostichi β Key characters of the group
based on adult female: stigmata and tracheae absent; genuaIIβIII without setae [42].
Eutarsopolipus paryavae Katlav & Hajiqanbarn. sp. (Figs. 1β3)
urn:lsid:zoobank.org:act:36B8618D-FA09-474C-B4C3-2613DD962A5B
Type material: Total material recovered: $ (n = 4),# (n = 15), larval $ (n = 4), ex. under elytra, on the base ofmembranous hind wing of one specimen of Geoscaptus
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Figure 1. Eutarsopolipus paryavae n. sp. (adult female). (a) Body dorsum; (b) body venter; (c) right leg I; (d) ventral view of tarsus I;(e) right leg II; (f) right leg III. All legs in dorsal view.
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Figure 2. Eutarsopolipus paryavae n. sp. (male). (a) Body dorsum; (b) body venter; (c) right leg I; (d) right leg II; (e) right leg III. All legs indorsal view.
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laevissimus Chaudoir, 1855 (Coleoptera: Carabidae: Scaritinae).Holotype: adult female (ANIC 52-003953), ex. under elytra, onthe base of membranous hind wing of G. laevissimus;
Coll. Shams Paryav; 11 Feb 2020. Paratypes: adult female(n = 3), male (n = 5) and larval female (n = 4), same data asholotype.
Figure 3. Eutarsopolipus paryavae n. sp. (larval female). (a) Body dorsum; (b) body venter; (c) right leg I; (d) right leg II; (e) right leg III. Alllegs in dorsal view.
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Type locality: Loc. Vines Drive, Hawkesbury Campus,Western Sydney University, Richmond, NSW, 33οΏ½36045.600 S150οΏ½44040.200 E.
Deposition of type material: The holotype, one adult female,2 male and 2 larval female paratypes are deposited at ANIC(ANIC 52-003953-58). 1 adult female, 2 males and 1 larvalfemale paratypes are deposited at QM (QMS 117000-04). Theremaining paratypes (TMU SP-20200211, 1β3), 10 non-typemales and the host beetle specimen are deposited at AC-DE-TMU.
Etymology: The new species is named after the firstauthorβs mother, Shams Paryav, the collector of the host beetlesamples, in gratitude of her immense engagement in materialcollections.
Authorship: Note that the authors of the new taxon aredifferent from the authors of this paper; Article 50.1 andRecommendation 50A of International Code of ZoologicalNomenclature [24].
Description
Adult female (Fig. 1) (n = 4)
Gnathosoma (Figs. 1aβ1b). Length 68 (72β86), width 58(60β68); cheliceral stylets length 66 (63β68); pharynx length14 (15β18), pharynx width 13 (13β14); ch 19 (21β26), su3 (4β5); distance between setae chβch 34 (40β42), suβsu17 (19β22).
Idiosoma (Figs. 1aβ1b). Length 300 (390β475), width250 (295β335).
Idiosomal dorsum (Fig. 1a). All dorsal setae needle-likeexcept attenuating setae sc2; prodorsal plate (PrS) with setaev1 5 (4β5), v2 vestigial, sc1 8 (7β8), sc2 36 (35β38). Plate Csetae c1 8 (9β10), c2 9 (9β11). Plate D setae d 9 (8β9); cupuliia anterolaterad setae d. Plate EF setae f 7 (7β8); cupuli imanterolaterad setae f. Plate H not evident; setae h 12 (9β11).Distances between setae: v1βv1 30 (31β35), v2βv2 61 (62β69),v1βv2 19 (21β25), sc1βsc1 74 (76β87), v1βsc1 27 (27β31),sc2βsc2 103 (113β125), sc1βsc2 49 (51β56), c1βc1 85 (91β99),c1βc2 49 (58β63), dβd 101 (104β107), fβf 67 (70β74), h1βh111 (14β18).
Idiosomal venter (Fig. 1b). All coxal plates smooth; allcoxal setae tiny needle-like; ap1β2 well developed, both reach-ing to appr; coxisternal field I with setae 1a 3 (3β4) and coxis-ternal field II with 2a 4 (3β4); alveoli of setae 1b and 2b notevident; coxisternal field III with setae 3a 7 (7β8) slightlylonger than 3b 5 (5β6). Distances between setae: 1aβ1a25 (26β35), 2aβ2a 30 (31β45), 3aβ3b 21 (23β28).
Legs (Figs. 1cβ1e). Setal formula for legs IβIII (femur-tarsus): 2-0-5(+u)-7(+x), 0-0-4-6, 0-0-4-5. Ambulacrum I withwell-developed sickle-shaped claw, ambulacrum IIβIII eachwith a pair of tiny claws. Leg I (Fig. 1c): femur, d microseta,slightly thickened, seta l0 1 (m-1); tibia, u 5 (6β7) clubbed,d 33 (29β35), l0 4 (4β5), l00 4 (4β6), v0 3 (4β5) and v00 5 (5β6)slightly thickened, seta k absent; tarsus, x 3 (3β4) digitiform,eupathidial setae tc0 14 (13β16) and tc00 14 (14β15) distinctlyblunt-ended, pl0 3 (3β5), pl00 5 (5-6), setae pv0 2 (2β3) andpv00 2 (2) slightly thickened, seta s 6 (6β7) blunt spur-like,u00 and p0 not evident. Leg II (Fig. 1d): tibia, d 14 (10β13),l0 4 (4β5), v0 5 (5β6), v00 4 (4β4); tarsus, tc0 5 (5β6), setae
u0 6 (7β8) and tc00 6 (6β7) blunt spur-like, pl00 28 (21β25),pv00 4 (4β5), u00 2 (2). Leg III (Fig. 1e): tibia, d 8 (7β9),l0 5 (4β4), v0 5 (5β6), v00 5 (5); tarsus, tc0 5 (5), setae u0 6 (6β7)and tc00 6 (6) blunt spur-like, pl00 20 (22β24), pv00 3 (4β4).
Male (Fig. 2) (n = 5)
Gnathosoma (Figs. 2aβ2b). Length 33β36, width 32β33;cheliceral stylets length 23β26; pharynx length 9β10, pharynxwidth 7β8; ch 17β21, su 3β4; distance between setae chβch25β26, suβsu 12β14.
Idiosoma (Figs. 2aβ2b). Length 145β210, width 120β130.Idiosomal dorsum (Fig. 2a). All dorsal setae short (except
sc2) and pointed; PrS with setae v1 2β3, v2 vestigial, sc1 4β6,setae sc2 52β65 attenuate. Plate CD with seta c1 4β5, c2 6β7,d 5β6; cupuli ia anterior to setae d. Plate EF setae f 3β4; cupuliim anterolaterad setae f. Genital capsule length 31β34, width25β30, situated posterior to margin of EF, setae h1 barelyvisible in few specimens. Distances between setae: v1βv118β22, v2βv2 40β43, v1βv2 18β20, sc1βsc1 50β55, v1βsc120β22, sc2βsc2 61β65, sc1βsc2 37β41, c1βc1 36β45, c1βc229β40, dβd 38β40, fβf 21β25.
Idiosomal venter (Fig. 2b). All coxal plates smooth; allcoxal setae pointed; ap1β2 and apsej well developed, all fusedwith appr; coxisternal field I with setae 1a 2, alveoli 1b notevident; coxisternal field II with 2a 3β3, alveoli 2b evident;coxisternal field III with setae 3a 5β6 slightly longer than3b 4β4. Distances between setae: 1aβ1a 15β19, 2aβ2a23β27, 3aβ3b 19β20.
Legs (Figs. 2cβ2e). Setal formula for legs IβIII (femur-tarsus): 2-0-5(+u)-8(+x), 0-0-4-6, 0-0-4-5. Ambulacrum I withwell-developed claw, ambulacrum IIβIII each with a pair of tinyclaws. Leg I (Fig. 2c): femur, d microseta, slightly thickened,seta l0 2β2 thickened; tibia, u 4β5 clubbed, d 24β26, l0 3,l00 1β2, v0 2β3, v00 3β4, seta k absent; tarsus, x 2 tiny, cone-shaped; eupathidial setae tc0 10β12 and tc00 11β12 distinctlyblunt-ended, setae pl0 3β4 and pl00 3β4 slightly blunt-ended,pv0 2, pv00 2β2, seta s 4β5 blunt spur-like, u00 1β2, seta p0 notevident. Leg II. (Fig. 2d): tibia, d 5β7, l0 3β5, v0 3β4, v00 2β3;tarsus, seta tc0 4β5, slightly blunt-ended; setae u0 5β6 andtc005β6 blunt spur-like, pl00 19β20, pv00 2β3, u00 1β2. Leg III(Fig. 2e): tibia, d 5β6, l0 3, v0 3, v00 3β3; tarsus, tc0 3β4 slightlyblunt-ended, setae u0 6β7 and tc00 5β6 blunt spur-like, pl00 18β20,pv00 3.
Larval female (Fig. 3) (n = 4)
Gnathosoma (Figs. 1aβ1b). Length 35β39, width 41β42;cheliceral stylets length 30β33; pharynx length 10β12, pharynxwidth 9β11; ch 21β24, su 3β4; distance between setae chβch35β38, suβsu 16β17.
Idiosoma (Figs. 3aβ3b). Length 220β235, width 135β175.Idiosomal dorsum (Fig. 3a). All dorsal setae pointed; PrS
with setae v1 3β4, v2 vestigial, sc1 6β7, sc2 65β75. Plate C setaec1 6β9, c2 7β9. Plate D setae d 6β8; cupuli ia anterolaterad setaed. Plate EF setae f 7β8; cupuli im anterior to setae f. Plate Hsituated ventrally with setae h1 130β140, h2 29β32. Distancesbetween setae: v1βv1 13β16, v2βv2 39β40, v1βv2 20β23, sc1βsc1 51β53, v1βsc1 26β27, sc2βsc2 53β55, sc1βsc2 40β41, c1βc149β53, c1βc2 29β32, dβd 28β29, fβf 28β31.
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Idiosomal venter (Fig. 3b). All coxal plates smooth; allcoxal setae tiny needle-like; ap1 and apsej well developed, bothfusing to appr; ap2 not reaching to appr; coxisternal field I withsetae 1a 2β3, alveoli 1b not evident; coxisternal field II with2a 2β3, alveoli 1b not evident; coxisternal field III with setae3a 5β6 and 3b 5β6 subequal. Distances between setae: 1aβ1a17β22, 2aβ2a 24β28, 3aβ3b 24β27.
Legs (Figs. 3cβ3e). Setal formula for legs IβIII (femur-tarsus): 2-0-5(+u)-8(+x), 0-0-4-6, 0-0-4-5. Ambulacrum I withwell-developed bifid claw with blunt tips, ambulacrum IIβIIIeach with a pair of barely discernible claws. Leg I (Fig. 3c):femur, d microseta, slightly thickened, seta l0 2β3 slightly thick-ened; tibia, u 5β5 baculiform, d 27β30, setae l0 4 and l00 2β3slightly blunt-ended, v0 2β3, v00 4β5, seta k absent; tarsus, x2β2 cone-shaped with blunt tip, eupathidial setae tc0 10β12and tc00 9 distinctly blunt-ended, pl0 3β4, pl00 4β5, setae pv0
2β2 and pv00 2β3 slightly thickened, seta s 5 blunt spur-like,u00 2, seta not evident. Leg II. (Fig. 3d): tibia, d 10β12,l0 4β5, v0 4β5, v00 3β4; tarsus, tc0 5β5, setae u0 5β7 and tc00
5β7 blunt spur-like, pl00 20β23, pv00 2β3, u00 2. Leg III(Fig. 3e): tibia, d 9β10, l0 5β6, v0 5, v00 2β3; tarsus, tc0 4β5,setae u0 6β7 and tc00 5β6 blunt spur-like, pl00 20β23, pv00 2β3.
Differential diagnosis
Within the pterostichi species group, the new species ismost similar to E. fischeri Husband, 1998 and E. teteri Husband& Husband, 2009 in having ambulacra II and III with a pair ofclaws each and ambulacra I with one claw and femur I with twosetae. However, it differs from both species in having cheliceralstylets longer than 60 (vs. shorter than 40 in both species), setaeh1 9β12 (absent in E. teteri and microsetae in E. fischeri) andseta k on tibia I absent (seta k on tibia I present in both species).The setal counts alone mask further differences. In E. paryavaeand E. fisheri, the setae on femur I are the tiny setae d and l0, butin E. teteri seta l0 is absent and v00 is present. Another importantdifference is the absence of a solenidion on tarsus II, which ispresent in E. teteri and probably present in E. fischeri (presentin male and larva, absent or obscured in females). All theimportant characters among these three species are comparedfor all life stages in Table 1 and a key to the world speciesof the pterostichi group of Eutarsopolipus (based on adultfemales) is presented in Figure 4.
Species group: leytei β Key characters of the group basedon adult females: stigmata and tracheae present; ambulacralclaws IIβIII present; genu IIβIII with setae [42].
Eutarsopolipus pulcher Hajiqanbar & Seemann. sp. (Figs. 5β8)
urn:lsid:zoobank.org:act:33189535-1D7E-4118-B46D-7EA9AE8E0207
Type material. Total material recovered: adult female(n = 12), male (n = 6), larval female (n = 13), ex. under elytra,on the base of membranous hind wing of specimens ofGnathaphanus pulcher (Dejean, 1829) (Coleoptera: Carabidae;Harpalinae). Four out of ca. 160 examined host specimensfound parasitized (4% prevalence). Samples were collectedat four independent events on 14 Feb 2020, 26 Feb 2020,
30 Feb 2020 and 3 March 2020. Holotype: adult female (ANIC52-003959), ex. under elytra, on the base of membranoushind wing of G. pulcher; Coll. Shams Paryav; 14 Feb 2020.Paratypes: adult female (n = 5), male (n = 5), larval female(n = 5), same data as holotype.
Type locality: Loc. Vines Drive, Hawkesbury Campus,Western Sydney University, Richmond, NSW, 33οΏ½36045.600 S150οΏ½44040.200 E.
Deposition of types: The holotype, one adult female, 2 maleand 2 larval female paratypes are deposited at ANIC (ANIC 52-003959-54). 2 adult female, 2 males and 1 larval femaleparatypes are deposited at QM (QMS 117005-10). The remain-ing paratypes (TMU SP-20200214, 1β3) and the host beetlespecimen are deposited at AC-DE-TMU.
Other material examined: adult female (n = 21), male(n = 4), larval female (n = 3), ex. under elytra, on the base ofmembranous hind wing of G. pulcher (host registration numberT137238), Loc. βFeez Creekβ property entrance, QLD,21οΏ½5104000 S 148οΏ½1401900 E; Coll. S. Wright & C. Burwell;9 Mar 2005 (QMS 117011-38). Adult female (n = 1), larvalfemale (n = 2), same data except different beetle (host registra-tion number T137239) (QMS 117039-41).
Etymology: The new species name βpulcherβ is adoptedafter the species name of the carabid host beetle G. pulchermeaning βbeautifulβ in Latin that is associated with thebeautiful metallic colouration patterns of elytra in this beetle.Furthermore, this epithet has a proper relevance to the beautifultrifurcate setae u0 on tarsi IIβIII in adult females of the new mitespecies.
Authorship: Note that the authors of the new taxon aredifferent from the authors of this paper; Article 50.1 andRecommendation 50A of International Code of ZoologicalNomenclature [24].
Description
Adult female (Fig. 5) (n = 6)
Gnathosoma (Figs. 5aβ5b). Length 45 (42β50), width 40(34β41); cheliceral stylets length 51 (44β48); pharynx length18 (16β18), pharynx width 14 (13β16); ch 18 (19β25), su 13(12β14); distance between setae chβch 24 (23β26), suβsu 14(14β16).
Idiosoma (Figs. 5aβ5b). Length 350 (270β415), width 215(180β290).
Idiosomal dorsum (Fig. 5a). All dorsal setae pointed;prodorsal plate (PrS) with setae v1 13 (11β14), v2 vestigial,sc1 11 (9β13), sc2 57 (51β60). Plate C setae c1 6 (4β6), c2 6(4β6). Plate D setae d 5 (4β5); cupuli ia evident, anterolateradsetae d. Plate EF setae f 5 (4β5); cupuli im evident, anterolateradsetae f. Plate H and setae h1 not evident. Distances betweensetae: v1βv1 46 (46β49), v2βv2 51 (49β59), v1βv2 34 (32β34),sc1βsc1 76 (74β80), v1βsc1 27 (27β29), sc2βsc2 93 (88β99),sc1βsc2 70 (68β71), c1βc1 81 (75β92), c1βc2 54 (49β54), dβd113 (102β112), fβf 83 (73β93).
Idiosomal venter (Fig. 5b). All coxal plates smooth; allcoxisternal setae tiny needle-like; ap1β2 and apsej welldeveloped, reaching to appr; coxisternal field I with setae 1a5 (5β6); alveoli of vestigial setae 1b evident; coxisternalfield II with 2a 5 (5β7); alveoli of vestigial setae 2b evident;
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Figure 4. Key to the world species of Eutarsopolipus in the pterostichi group (based on adult females).
Table 1. Comparison of selected characters (range of measurements if available) of E. paryavae n. sp. (Ep) with two closely related species ofthe pterostichi species group: E. fischeri (Ef) and E. teteri (Et). Dashes (β) denote absence of the character. Characters for which the data areobscured/not available in the original descriptions are given as question marks (?). Letters v and m indicate vestigial and mirosetae,respectively. Abbreviations: L. (length), S. (stylet), Gn. (gnathosoma), Gen. cap. (genital capsule), Ch. (chelicera), Sol. (solenidion), Ta(tarsus), Ti (tibia), Fe (femur).
Life stage Female Male Larval female
Character Ep Ef Et Ep Ef Et Ep Ef Et
Gn. L. 68β86 45β48 45 33β36 26β29 29β32 35β39 31 35β38Ch. S. L. 63β68 33β34 38 23β26 19β22 22 30β33 22 32β34Setae ch 19β26 17β19 15 17β21 2β4 5 21β24 17 20β25Setae su 3β5 5β6 14 3β4 2β3 8 3β4 3 10β12Setae v1 4β5 5β6 10 2β3 2 ? 3β44 5 15Setae sc1 7β8 6β7 9 4β6 2 ? 6β7 3 9β10Setae sc2 35β38 32β42 59 52β65 48 38β45 65β75 74 80Setae c1 8β10 3β5 5 4β5 2 ? 7β9 3 8β12Setae c2 9β11 5β6 7 6β7 2 ? 7β9 4 10Setae d 8β9 3β5 7 5β6 ? ? 6β8 4 10Setae f 7β8 5 5 3β4 ? ? 7β8 4 7β10Setae h1 9β12 m β β β β 130β140 43 60β62Setae h2 β β β β β β 29β32 16 5β7Setae 1a 3β4 2 m 2 v mβ3 2β3 m 4Setae 2a 3β4 2β3 7 3 v 3 2β3 m 5β6Setae 3a 7β8 4β5 ? 5β6 v ? 5β6 3 ?Setae 3b 5β6 7 3 4 ? m 5β6 4 5Gen. cap. L. β β β 31β34 25β38 38 β β β
Gen. cap. W. β β β 25β30 27β30 32 β β β
Sol. Ta I x 3β4 3 5 2 2β3 5 2 3 4β5Sol. Ti I u 5β7 7β10 7 4β5 7β8 5β7 4β5 6 8β13Sol. Ta II x β ? 5 β 2β3 5 β 3 5Fe I seta v00 β β 15 β β 10β18 β β 10Fe I seta l0 mβ1 ~3β4 β 2 ~1 β 2β3 ~2 β
Ta III seta pl00 20β24 15β17 15 18β20 12 10β15 20β23 13 15β18
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Figure 5. Eutarsopolipus pulcher n. sp. (adult female). (a) Body dorsum; (b) body venter; (c) right leg I; (d) ventral view of tarsus I; (e) rightleg II; (f) right leg III. All legs in dorsal view.
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coxisternal field III with setae 3a 4 (4β5) and 3b 6 (5β6).Distances between setae: 1aβ1a 31 (29β35), 2aβ2a 34(36β42), 3aβ3b 26 (22β27).
Legs (Figs. 5cβ5e, 6). Setal formula for legs IβIII (femur-tarsus): 3-2-6(+u)-8(+x), 0-1-4-6 (+x), 0-1-4-6. AmbulacrumI with sickle-shaped claw, ambulacrum IIβIII each with a pairof well-developed claws. Leg I (Fig. 5c): femur, d microseta,seta l0 15 (14β16) pointed and thickened, subequal to v00 15(12β15); genu with tiny setae l0 2 (2β2) and l00 1 (1β2); tibiawith u 8 (7β9) baculiform, d 29 (25β29), l0 11 (9β11), l00 9(7β9), v0 5 (5β6) stiff, v00 14 (13β16), seta k 8 (8β10); tarsusI, x 5 (4β5) digitiform, eupathidial setae tc0 14 (12β15) andtc00 15 (13β15) distinctly blunt-ended, pl0 11 (11β13), pl00 15(13β17), setae pv0 3 (3β3) and pv00 2 (2β3) subequal, seta s 6(6β7) modified and thickened, p0 2 (2) slightly thickened. LegII. (Fig. 5d): genu, l0 2 (2); tibia, d 17 (15β17), l0 9 (7β9),v0 14 (12β14), v00 15 (13β19); tarsus II, x 4 (3β4) digitiform,tc0 5 (5β7), setae u0 8 (7β8) spine-like and trifurcate, tc00 6 (6β7)blunt spur-like, pl00 13 (12β13), pv00 2 (2β3), u00 2 (2). Leg III(Figs. 5e, 6): genu, l0 2 (2β2); tibia, d 17 (15β18), l0 9 (7β9),v0 14 (13β14), v00 17 (15β18); tarsus III, tc0 5 (5β7), setae u0 8(7β8) spine-like and trifurcate (Fig. 6), tc00 6 (5β6) bluntspur-like (Fig. 5f), pl00 14 (12β14), pv00 2 (2β3), u00 14 (12β14).
Male (Fig. 7) (n = 5)
Gnathosoma (Figs. 7aβ7b). Length 25β36, width 23β27;cheliceral stylets length 17β19; pharynx length 9β10, pharynx
width 6β8; ch 8β12; su 9β10; distance between setae chβch17β20, suβsu 12β13.
Idiosoma (Figs. 7aβ7b). Length 140β160, width 105β115.Idiosomal dorsum (Fig. 7a). All setae on dorsum microsetae
(except sc2); PrS with setae v2 vestigial, setae sc2 34β46 atten-uate and pointed. Plate CD with cupuli ia anterior to setae d.Plate EF setae with cupuli im anterolaterad setae f. Genital cap-sule length 23β30, width 28β33, situated posterior to margin ofEF, setae h1 barely visible on genital capsule. Distancesbetween setae: v1βv1 18β19, v2βv2 30β33, v1βv2 18β19, sc1βsc144β47, v1βsc1 17β18, sc2βsc2 42β45, sc1βsc2 26β28, c1βc139β43, c1βc2 25β29, dβd 29β34, fβf 19β21.
Idiosomal venter (Fig. 7b). All coxal plates smooth; allventral setae on coxal area microsetae; ap1-2 well developed,fused with appr, apsej weekly developed, not reaching appr;alveoli of setae 1b on coxisternal field I evident; on coxister-nal field II alveoli of setae 2b evident. Distances between setae:1aβ1a 17β19, 2aβ2a 22β24, 3aβ3b 18β20.
Legs (Figs. 7cβ7e). Setal formula for legs IβIII (femur-tarsus): 2-2-6(+u)-8(+x), 0-1-4-6(+x), 0-1-4-6. Ambulacrum Iwith a small claw, ambulacrum IIβIII each with a pair of smallclaws. Leg I (Fig. 7c): femur, setae d and l0 microsetae; genu,setae l0 and l00 microsetae; tibia, u 6β8 baculiform, d 21β25,l0 and l00 microsetae, v0 1, v00 12β14, seta k 3β5; tarsus, x 4β5digitiform; eupathidial setae tc0 8β10 and tc00 10β11 distinctlyblunt-ended, setae pl0 8β9 and pl00 10β12, seta pv0 1β1 stiffand blunt-ended, pv00 2β2, seta s 4β5 blunt spur-like, p0 1β1.Leg II. (Fig. 7d): genu, l0 1; tibia, l0 2, d 13β15, v0 11β14, v00
11β14; tarsus, x 4β5 thickened and digitiform, seta tc0 4β5,slightly blunt-ended, u0 4β6 spine-like and bifurcate, tc00 4β5blunt spur-like, pl00 9β11, pv0 2, pv00 2, u00 1. Leg III (Fig. 7e):genu, l0 1; tibia, d 10β13, l0 1, v0 1, v00 12β14; tarsus, tc0 5β7 stiffand slightly blunt-ended, setae u0 5β6 spine-like and bifurcate,tc00 4β5 blunt spur-like, pl00 9β10, pv00 1, u00 1.
Larval female (Fig. 8) (n = 5)
Gnathosoma (Figs. 8aβ8b). Length 29β32, width 24β29;cheliceral stylets length 28β34; pharynx length 10β13, pharynxwidth 7β9; ch 20β25; su 10β12; distance between setae chβch15β18, suβsu 10β11.
Idiosoma (Figs. 8aβ8b). Length 125β145, width 95β110.Idiosomal dorsum (Fig. 8a). All dorsal setae needle-like
except sc2 which is long and attenuate; PrS with setae v1 11β13,v2 vestigial, sc1 10β12, sc2 62β72. Plate C setae c1 7β8, c2 5β7.Plate D setae d 6β7; cupuli ia anterolaterad setae d. Plate EFsetae f 7β9; cupuli im anterior to setae f. Plate H not evident;setae h1 64β66, h2 mβ2. Distances between setae: v1βv125β28, v2βv2 35β37, v1βv2 22β25, sc1βsc1 56β60, v1βsc123β26, sc2βsc2 44β48, sc1βsc2 30β32, c1βc1 25β28, c1βc231β34, dβd 22β25, fβf 25β30.
Idiosomal venter (Fig. 8b). All coxal plates smooth; allcoxal setae tiny needle-like; ap1β2 well developed, both fusingto appr; apsej not evident; coxisternal field I with setae 1a 5β7;alveoli of setae 1b on coxisternal field I evident; coxisternalfield II with 2a 4β6; alveoli of setae 2b evident; coxisternal fieldIII with setae 3a 5β6 and 3b 5 subequal. Distances betweensetae: 1aβ1a 17β21, 2aβ2a 16β22, 3aβ3b 17β19.
Legs (Figs. 8cβ8e). Setal formula for legs IβIII (femur-tarsus): 3-2-6(+u)-8(+x), 0-1-4-5(+x), 0-1-4-5. Ambulacrum I
Figure 6. Phase-contrast micrograph of tarsus III in Eutarsopolipuspulcher n. sp. (adult female) representing modified trifurcate seta u0
and spur-like seta tc00.
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Figure 7. Eutarsopolipus pulcher n. sp. (male). (a) Body dorsum; (b) body venter; (c) right leg I; (d) right leg II; (e) right leg III. All legs indorsal view.
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with a small bifid claw, ambulacrum IIβIII each with a pair ofsmall claws. Leg I (Fig. 8c): femur, d microseta, seta l0 2slightly thickened, v00 10β11; genu, l0 2, l00 1β1; tibia, u 7β8
baculiform and bent, d 28β30, setae l0 8β9 and l00 9β12, v0
4β5, v00 10β12, seta k 3β4; tarsus, x 4β5 digitiform, eupathidialsetae tc0 8β9 and tc00 9β10 distinctly blunt-ended, pl0 9β10,
Figure 8. Eutarsopolipus pulcher n. sp. (larval female). (a) Body dorsum; (b) body venter; (c) right leg I; (d) right leg II; (e) right leg III. Alllegs in dorsal view.
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pl00 12β14, setae pv0 1β1 and pv00 1β2, seta s 3β5 blunt spur-like,p0 2. Leg II. (Fig. 8d): genu, l0 2; tibia, d 13β18, l0 10β11, v0
10β13, v00 13β15; tarsus, x 4β5 digitiform, tc0 3β4, setae u0 5and tc00 4β5 blunt spur-like, pl00 9β11, pv00 2, u00 not evident.Leg III (Fig. 8e): genu, l0 2; tibia, d 14β15, l0 8β12, v0 7β11,v00 11β12; tarsus, tc0 6β7, setae u0 5β6 and tc00 5β6 blunt spur-like, pl00 10β11, pv00 2, u00 not evident.
Differential diagnosis
This new species is unique in Eutarsopolipus by havingtrifurcate setae u0 on tarsi IIβIII. However, among species withsimple claws on legs I (unlike E. biuncatus Seeman, 2021 andE. janus Seeman, 2021 with bifurcate claws on legs I), it is mostsimilar to E. leytei Husband & Raros, 1989 with femur I seta lΚΉvery short, not reaching genual base in adult females; but itis readily distinguishable from this species by longer setae v111β14 (mβ5 in E. leytei) and shorter cheliceral stylets beingat most 51 in E. pulcher n. sp. vs. 68 in E. leytei.
The new species further differs from E. dastychi with setaev1 longer than ch and setae c1, c2, d and f shorter than 8 in adultfemales (vs. setae v1 shorter than ch and setae c1, c2, d and flonger than 15 in adult females of E. dastychi). The male ofE. pulcher n. sp. resembles that of E. orpheus with all ventraland dorsal setae (except sc2) being microsetae, but it differsfrom E. orpheus with setae ch longer than 8 (ch microsetaein male of E. orpheus). The larval female of E. pulcher n. sp.is similar to E. orpheus with h1 shorter than 70 and h2 shorterthan 2, but it is readily distinguishable from E. pulcher n. sp. byshorter setae sc1, sc2, c1, c2, d, 3a and 3b (Table 2). All theimportant characters among the species of leytei group are com-pared for all life stages (excluding E. leytei with unknown male)in Table 2 and keys to the world species (based on adultfemales) are presented in Figure 9.
Species group: myzus β Key characters of the group basedon adult females: stigmata and tracheae present; ambulacralclaws IIβIII present; genu IβIII without setae; femur I withtwo setae [42].
Eutarsopolipus chlaenii Katlav & Hajiqanbarn. sp. (Figs. 10β11)
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 hindwing 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 eventson 24 Feb 2020, 26 Feb 2020, and 28 Feb 2020. Holotype:adult female (ANIC 52-003965), ex. under elytra, on the baseof membranous hind wing of C. flaviguttatus; Coll. ShamsParyav; 24 Feb 2020. Paratypes: adult female (n = 3), larvalfemale (n = 5), same data as holotype (24 Feb 2020, 26 Feb2020, and 28 Feb 2020).
Type locality: Loc. Vines Drive, Hawkesbury Campus,Western Sydney University, Richmond, NSW, 33οΏ½36045.600 S150οΏ½44040.200 E.
Deposition of material: The holotype, one adult femaleand 2 larval female paratypes are deposited at ANIC (ANIC52-003965-68). 1 adult female and 2 larval female paratypesare deposited at QM (QMS 117009-10, 117042). The remain-ing paratypes (TMU SP-20200224, 1β3), 11 non-type larvalfemales and the host beetle specimen are deposited atAC-DE-TMU.
Etymology: The species epithet βchlaeniiβ refers to thegeneric name of the carabid host beetle Chlaenius flaviguttatus.
Authorship: Note that the authors of the new taxon aredifferent from the authors of this paper; Article 50.1 andRecommendation 50A of International Code of ZoologicalNomenclature [24].
Adult female (Fig. 10) (n = 4)
Gnathosoma (Figs. 10aβ10b). Length 45 (43β45), width 42(40β42); cheliceral stylets length 28 (29β30); pharynx length 12(12β13), pharynx width 12 (12β13); ch 15 (15β16), pointed;su 6 (5β6), needle-like; distance between setae chβch 27 (27β29),suβsu 17 (16β17).
Idiosoma (Figs. 10aβ10b). Length 230 (225β240), width185 (165β185).
Idiosomal dorsum (Fig. 10a). Respiratory system (stigmataand tracheae) present, stigmata stalked; all dorsal setae pointed;prodorsal plate (PrS) with setae v1 6 (5β6), setae v2 vestigial,setae sc1 5 (5β6), sc2 42 (38β41). Plate C setae c1 7 (6β7),c2 5 (6β6). Plate D setae d 5 (5β6); cupuli ia evident, anterolat-erad setae d. Plate EF setae f 7 (6β7); cupuli im evident, antero-laterad setae f. Plate H not evident, setae h 7 (7β8). Distancesbetween setae: v1βv1 35 (34β37), v2βv2 42 (41β44), v1βv213 (13β14), sc1βsc1 61 (57β60), v1βsc1 18 (18β19), sc2βsc262 (58β61), sc1βsc2 39 (38β40), c1βc1 61 (57β64), c1βc2 48(42β46), dβd 59 (57β58), fβf 37 (33β36).
Idiosomal venter (Fig. 10b). All coxal plates smooth; allcoxal setae pointed; ap1β2 and appr well developed, ap2reaching to appr; apsej absent; coxisternal field I with setae1a 3 (2β2); alveoli of vestigial setae 1b not evident; coxisternalfield II with 2a 2 (2β2); alveoli of vestigial setae 2b evident;coxisternal field III with subequal setae 3a 7 (7β8) and 3b7 (8β8). Distances between setae: 1aβ1a 19 (20β22), 2aβ2a27 (25β26), 3aβ3b 19 (24β26).
Legs (Figs. 10cβ10e). Setal formula for legs IβIII (femur-tarsus): 2-0-6(+u)-8(+x), 0-0-4-6(+x), 0-0-4-6. Ambulacrum Iwith a well-developed sickle-shaped claw, ambulacrum IIβIIIeachwith a pair ofwell-developed claws. Leg I (Fig. 10c): femur,d 3 (2β2), slightly thickened, seta l0 16 (15β16) thick and blunt-ended; tibia with u 5 (5β5) baculiform, d 28 (29β31), l0 4 (4β5),l00 3 (3β3), v0 5 (5β6) stiff, v00 9 (8β9), seta k 5 (5β6); tarsus I, x 3(3β4) digitiform, eupathidial setae tc0 9 (8β9) and tc00 10 (9β10)distinctly blunt-ended, pl0 9 (8β9), setae u00 2 (2β3), pv0 3 (2β2)and pv00 2 (2β2) subequal, seta s 5 (5β5) spine-like, with a blunttip, p0 1 (1β2). Leg II. (Fig. 10d): tibia, d 8 (7β8), l0 7 (6β7), v0 11(10β11), v00 6 (6β6); tarsus, x 4 (3β4) digitiform, tc0 7 (6β7),setae u0 6 (6β6) and tc00 5 (5β6) spine-like, pl00 17 (17β18),pv00 3 (3β3), u00 2 (2β2). Leg III (Fig. 10e): tibia, setae d 7 (7β8),l0 6 (6β7), v0 11 (10β12), v00 6 (5β6); tarsus, tc0 10 (10β10), setae u0
6 (6β7) and tc00 6 (5β6) spine-like, pl00 16 (15β16), pv00 3 (3β3),u00 2 (2β2).
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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. 42β50 78 47β50 50β57 44β46 52β57 25β36 30β33 30β32 28β31 28β37 29β32 56 37β40 32β38 30β34 31β34Ch. S. L. 44β51 68 47β48 23β28 35β38 45β51 17β19 23β26 15 17β18 25β28 28β34 54 35β40 21β27 25β27 36β42Setae ch 18β25 20 27β30 10β18 14β22 25β35 8β12 12β15 m 1β2 3β4 20β25 20 30β36 31β35 19β23 26β33Setae su 12β14 3 22 11β15 8β12 15β20 9β10 9β10 6β7 4β6 8β9 10β12 3 15β18 6β7 8β10 12β15Setae v1 11β14 5 36β45 17β26 8β10 19β24 m 5β10 m mβ2 4β9 11β13 m 42β50 26β32 12β14 15β18Setae sc1 9β13 11 25 17β29 7β9 9β11 m 5β13 m m mβ2 10β12 10 25β28 28β33 5β6 8β10Setae sc2 51β60 62 58β70 47β59 15β20 36β40 34β46 60β70 38β40 2β3 27β34 62β72 78 94β101 90β95 51β52 70β80Setae c1 4β6 12 18β19 8β10 5β7 8β9 m 10 m m 2β3 7β8 10 18β22 14β18 5β6 7β10Setae c2 4β6 9 17β21 9β18 4β6 8β10 m 5β7 m m 2 5β7 9 17β21 10β15 3β4 7β10Setae d 4β5 12 18β21 11β18 4β5 9β10 m 8β10 m m mβ2 6β7 9 20β23 15β17 4β5 7β9Setae f 5 8 22 10β11 4β5 7β9 m 3β4 m m mβ2 7β9 8 16β18 8β12 4β5 6β8Setae h1 β β β β β β m m m m m 64β67 148 65β90 55β64 65β70 100Setae h2 β β β β β β β β β β β 1β2 67 m m 3β4 4β6Setae 1a 5β6 ~4 5β6 4β5 5β6 6β7 m 2β5 m 2β3 4β5 5β7 ~2 8β10 6β8 2β3 4β5Setae 2a 5β7 ~4 5 3β4 4β5 6β7 m 3β4 m 2β3 4β5 4β6 ~2 7β10 8β9 3β4 4β5Setae 3a 4β5 8 9 2β3 5β6 6β9 m ~2 m 3 5β6 5β6 4 10β13 9β11 4β5 7β8Setae 3b 5β6 4 7 3 4β5 6β7 m ~4 m 2β3 5β6 5 2 10β12 9β10 3β4 5β6Sol. Ta I x 4β5 ~2 4β5 5β6 3β4 4β5 4β5 5β7 4 3 3β4 4β5 ~2 4β5 4β5 3β4 4β5Sol. Ti I u 7β9 7 10 7β8 6β7 6β8 6β8 5β6 5β6 4β5 6β7 7β8 7 7β9 8β9 6β7 7β8
14A.Katlav
etal.:
Parasite2021,
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Male (Unknown)
Larval female (Fig. 11) (n = 5)
Gnathosoma (Figures 11aβ11b). Length 39β46, width38β40; cheliceral stylets length 29β32; pharynx length 13β15,pharynx width 10β11; ch 17β19 pointed; su 3β4 needle-like;distance between setae chβch 22β26, suβsu 12β14.
Idiosoma (Figs. 11aβ11b). Length 165β195, width 125β145.Idiosomal dorsum (Fig. 11a). All dorsal setae needle-like
except sc2 attenuate; PrS with setae v1 6β7, v2 vestigial, sc16β7, sc2 95β98. Plate C setae c1 5β6, c2 5β6. Plate D setae d6β7; cupuli ia anterolaterad setae d. Plate EF setae f 7β8; cupuliim anterolaterad setae f. Plate H situated ventrally with setae h197β101 and h2 22β24. Distances between setae: v1βv1 24β26,v2βv2 44β47, v1βv2 12β14, sc1βsc1 59β62, v1βsc1 20β22,sc2βsc2 57β58, sc1βsc2 40β42, c1βc1 83β86, c1βc2 21β23, dβd34β36, fβf 28β29.
Idiosomal venter (Fig. 11b). All coxal plates smooth;all coxal setae tiny and pointed; ap1β2 and apsej evident;coxisternal fields IβII each divided from its pair, with setae1a 1β1; alveoli of setae 1b not evident; coxisternal field IIwith 2a 2β3; alveoli of setae 2b not evident; coxisternalfield III widened, with setae 3a 7β9 and 3b 7β8 subequal.Distances between setae: 1aβ1a 22β34, 2aβ2a 28β30, 3aβ3b21β22.
Legs (Figs. 11cβ11e). Setal formula for legs IβIII (femur-tarsus): 2-0-6(+u)-7(+x), 0-0-4-6(+x), 0-0-4-6. Ambulacrum Iwith a small bifid claw, ambulacrum IIβIII each with a pair oftiny claws. Leg I (Fig. 11c): femur, d microseta, seta l0 4β5 stiff;tibia, u 3β4 baculiform, d 17β21, seta l0 4β5 slightly thickened,l00 2β3, v0 3β4, seta v00 4β5 slightly thickened and blunt-ended,seta k 2β3; tarsus, x 3β4 digitiform, eupathidial setae tc0 7β8
and tc00 7β8 subequal, distinctly blunt-ended, pl0 6β6, setaepv0 1β2, pv00 2β3, seta s 4β5 blunt spur-like, p0 1β1; u00 notvisible. Leg II. (Fig. 11d): tibia, d 4β5, l0 6β7, v0 6β7, v00 4β6;tarsus, x 2β3 digitiform, tc0 5β7, setae u0 4β5 and tc00 5β6 bluntspur-like, pl00 10β12, pv00 2β2, u00 2β2. Leg III (Fig. 11e): tibia,d 5β6, l0 6β7, v0 6β8, v00 3β5; tarsus, tc0 6β7, setae u0 5β6 andtc00 6β7 blunt spur-like, pl00 10β13, pv00 3β4, u00 1β1.
Differential diagnosis
The new species belongs to a subgroup of the myzus speciesgroup that shares a combination of the following characters inadult females: ambulacrum I claw well-developed, idiosomawithout lateral bulges or posteriorly without wrinkled lobes,shield C not divided, femur I seta l0 developed (not microseta),and cheliceral stylets less than 35 lm long [13]. This assemblageincludes E. chlaenii n. sp. and four other species: E. steveniKhaustov, 2010, E. anichtchenkoi, Hajiqanbar & Mortazavi,2012, E. gombrooni Hajiqanbar & Mortazavi, 2019, andE. oconnori Hajiqanbar & Mortazavi, 2019. Among these spe-cies, E. chlaenii n. sp. is more similar to E. anichtchenkoi andE. oconnori by having setae h and f subequal. However, it isreadily distinguishable from E. anichtchenkoi by having devel-oped setae v1, sc1, 1a and 2a (adult female with microsetae v1,sc1, 1a and 2a in E. anichtchenkoi), sc2 almost five times longerthan h1 (adult female with sc2 at least nine times longer than h1in E. anichtchenkoi) and tarsus III with six setae (tarsus III withseven setae in E. anichtchenkoi). Eutarsopolipus chlaenii n. sp.also differs from E. oconnori by having shorter distances c1-c1,d-d, f-f in the adult female (64, 59, 37 vs. 101, 113, 86, respec-tively, in E. oconnori) and longer setae su and h1 and cheliceralstylets in larval females (101, 4, 32, vs. 61, m, 26, respectively,
Figure 9. Key to the world species of Eutarsopolipus in the leytei group (based on adult females).
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Figure 10. Eutarsopolipus chlaenii n. sp. (adult female). (a) Body dorsum; (b) body venter; (c) right leg I; (d) ventral view of tarsus I; (e) rightleg II; (f) right leg III. All legs in dorsal view.
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in E. oconnori). All the important characters among these fivespecies of the myzus species group are compared for all lifestages (excluding E. chlaenii n. sp. with unknown male) inTable 3. Among adult females of the myzus species group with
a strong claw on ambulacrum I, lateral bulges or posteriorwrinkled lobes and entire shield C, six species have short che-liceral stylets (less than 35 lm long). The key to this subgroupis presented in Figure 12.
Figure 11. Eutarsopolipus chlaenii n. sp. (larval female). (a) Body dorsum; (b) body venter; (c) right leg I; (d) right leg II; (e) right leg III. Alllegs in dorsal view.
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Table 3. Comparison of selected characters (range of measurements if available) of five closely related species of the myzus species group inEutarsopolipus (male is unknown for E. chlaenii n. sp.): E. chlaenii n. sp. (Ec), E. steveni (Es), E. anichtchenkoi (Ea), E. gombrooni (Eg), andE. oconnori (Ea).
Life stage Female Male Larval female
Character Ec Es Ea Eg Eo Es Ea Eg Eo Ec Es Ea Eg Eo
Gn. L. 43β45 31β34 47β50 43β50 50 21β22 26β29 24ββ26 25 39β46 22β24 30β35 23 26β36Ch. S. L. 28β30 28β31 30β35 29β32 34 14β15 12β13 13β16 13 29β32 19β20 17β19 18 16β26Setae ch 15β16 17β19 12β13 14β16 14 6β7 8β10 4 5 17β19 15β17 12β13 14 11β14Setae su 5β6 6β7 4β6 5β7 5 3β4 2β3 2 m 3β4 7β8 2 7 mSetae v1 5β6 5β6 m 6β7 8 3β4 m 3β4 m 6β7 3β4 6β7 5 5β7Setae sc1 5β6 5β6 m 8 8 3β4 m 3β4 m 6β7 3β4 5β6 5 7β8Setae sc2 38β42 26β28 37β45 18β21 45 35β37 39β45 26β31 17 95β98 41β47 78β79 43 85β93Setae c1 6β7 6β8 4β5 5β9 9 4β5 m mβ3 m 5β6 4β5 7β9 5 6β7Setae c2 5β6 6β7 4β5 7β8 9 4β5 m 4β5 m 5β6 4β5 6 4 6β7Setae d 5β6 6β7 4β5 6β7 8 4β5 m mβ4 m 6β7 4β5 9 3 7Setae f 6β7 7β8 4β5 7β8 8 4β4 m mβ3 m 7β8 6β7 9 6 5β8Setae h1 7β8 24β26 3β4 13β16 9 β β β β 97β101 70β75 140β172 89 57β61Setae h2 β β β β β β β β β 22β24 24β27 20β21 20 12β20Setae 1a 2β3 2β3 m 2β3 m v m m m 1 2 m m mSetae 2a 2 3β4 m 3β4 m 2 m 1β2 m 2β3 3 m 2 mSetae 3a 7β8 ~8 4β5 8β9 9 3β4 m 3β4 m 7β9 10 3 10 5β9Setae 3b 7β8 8β9 4β5 8 11 4 m 4β5 m 7β8 6 4 9 6β8Gen. cap. L. β β β β β 21β22 26β29 31β34 18 β β β β β
Gen. cap. W. β β β β β 21β22 25β28 34β35 24 β β β β β
Sol. Ta I x 3β4 3β4 2 3 3 3β4 3 3β4 6 3β4 3β4 2β3 ~2 2β3Sol. Ti I u 5 4β5 5 4β5 5 4β5 4 3β5 8 3β4 4β5 4 4 4β5Sol. Ta II x 3β4 3β4 2 3 3 3β4 2 3β4 6 2β3 3β4 3 3 3β4Fe I seta d 2β3 m m m ~1 m m m m m m m m mFe I seta lβ 15β16 ~14 12β13 11β13 13 m ~1 ~1 4 4β5 m ~3 2 4β5Ti I seta d 28β31 ~20 ~22 17β26 18 ~17 ~22 15β16 28 17β21 ~18 ~18 ~20 ~19
Figure 12. Key to closely related species of myzus group (based on adult females) possessing similar characters including short cheliceralstylets (<35 lm long).
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Discussion
Among all Eutarsopolipus, leytei is apparently the mostprimitive group that represents the putative plesiomorphies ofa well-developed tracheal system as well as retention of genualIβIII setae (2-1-1) and all femoral I setae (3 setae). Conversely,the pterostichi group with a missing tracheal system andgenual IβIII setae (0-0-0), reduction of femoral I setation(2 setae) and sometimes reduction/absence of ambulacral claws[as in E. echinatus, 43] may be relatively more derivative thanthe other Australian Eutarsopolipus [31, 43]. However, themyzus group, possessing a combination of plesiomorphies(well-developed tracheal system and ambulacral claws) andsome apomorphies [reduction of femoral I setation (2 setae)and absence of genual IβIII setae (0-0-0)], may hold an interme-diate position. It is surprising that in our study such consider-able species diversity was detected in a single locationfollowing a minimal sampling effort preformed across fewerthan three weeks. This may substantiate the previously heldnotion that Australia exhibits diverse Eutarsopolipus fauna witha wide gradient of morphological variations [42]. Despite a fewsporadic studies on Australian Eutarsopolipus, six out of the tenknown species groups that exist across the world (includingochoai, megacheli and secundus) have so far been recordedfrom Australia ([31, 42, 44], present study). However, the richdiversity of Australian carabid beetles may posit the idea thatthe current knowledge about their associated Eutarsopolipusmites is still in its infancy; therefore, more extensive faunisticstudies in different regions could potentially lead to the discov-ery of enormous diversity in Eutarsopolipus.
With the description of E. chlaenii, this study reports themyzus group for the first time in Australia, thereby extendingits distribution to Oceania, and beyond the previously recordedHolarctic, Afrotropical and Oriental realms [12, 22]. About halfof the species of this group (13/25) are parasites of carabids ofthe genus Chlaenius Bonelli [12, 22]. Furthermore, the findingof E. pulcher n. sp. from G. pulcher is the second record of theleytei group from a native carabid of the genus GnathaphanusMacleay, 1825 (tribe Harpalini). Recently, a study in the same
location found another species, E. orpheus from under theelytra of Gnathaphanus melbournensis (Castelnau, 1867),probably suggesting more specific association of the leyteigroup with carabids of Gnathaphanus. This carabid genus isapparently native to the Australasian and Oriental regions andrepresents more than 15 species in Australia [4] with G. pulcherand G. melbournensis being highly abundant in easternAustralia [3]. It is interesting, however, that the only Palearcticrepresentative of the leytei group, E. dastychi, was found fromCalathus of the carabid tribe Sphodrini [20] which is phyloge-netically diverged from the carabid tribe Harpalini. This kind ofcounterintuitive host range is even more profound among themyzus and pterostichi groups, both of which are associated withcarabids of the two distantly related subfamilies, Harpalinae andScaritinae [26, 42], suggesting that several episodes of hostswitching may have contributed to the evolution of their hostassociations.
Carabid beetles are generalist predators that feed on avariety of small invertebrates including important agriculturalpests and thus serve as important biocontrol agents [34].However, their ecological interactions are often hard to predict[9]. It is unknown how the parasitic role of Eutarsopolipusmites can shape the ecology and evolution of carabids, yetincorporation of such information may contribute to modelspredicting interaction networks of carabids for future biocontrolprograms.
Conflict of interest
The authors declare that they do not have any conflict ofinterest.
Acknowledgements. We sincerely thank Geoff Monteith (formerSenior Curator of Entomology at the Queensland Museum) for hisassistance in identifying some of the host beetles. Special thanksto the first authorβs mother, Shams Paryav for her kind assistancein filed insect collection. AK was supported by a Western SydneyUniversity Postgraduate Research Award. OS was partially fundedby the Australian Biological Resources Study (grant RG18-02) andby the Queensland Museum.
References
1. Abbot P, Dill LM. 2001. Sexually transmitted parasites andsexual selection in the milkweed leaf beetle, Labidomeraclivicollis. Oikos, 92, 91β100.
2. Athias-Binche F. 1993. From phoresy to parasitism: theexample of mites and nematodes. Research and Reviews inParasitology-Revista, 53, 73β79.
3. Atlas of Living Australia. 2020. https://bie.ala.org.au/species/urn:lsid:biodiversity.org.au:afd.taxon:7bbbcf53-08e7-4c98-8373-211849be892c; accessed 10 March 2020.
4. Australian Faunal Directory. 2020. https://biodiversity.org.au/afd/taxa/Gnathaphanus; accessed 10 March 2020.
5. Baehr M, Will K. 2019. Carabidae Latreille, 1802, inβAustralian Beetlesβ: Archostemata, Myxophaga, Adephaga,Polyphaga, Vol. 2, SlipinskΓ A, Lawrence JF, Editors. CSIROPublishing: Melbourne, Vic., Australia. p. 61β217.
6. Bartlow AW, Agosta SJ. 2021. Phoresy in animals: review andsynthesis of a common but understudied mode of dispersal.Biological Reviews, 96, 223β246.
Figure 13. Eutarsopolipus chlaenii n. sp. under the elytra of thehost beetle Chlaenius flaviguttatus Macleay, 1825, localized on theproximal portion of the hostβs hindwing, with view of the miteβsengorged female (yellow colour) producing eggs (milky colour).
A. Katlav et al.: Parasite 2021, 28, 75 19
![Page 20: Sheltered life beneath elytra: three new species of ...](https://reader034.fdocuments.us/reader034/viewer/2022050415/6270ef1363a8375213003b34/html5/thumbnails/20.jpg)
7. Bochkov AV, OConnor BM, Wauthy G. 2008. Phylogeneticposition of the mite family Myobiidae within the infraorderEleutherengona (Acariformes) and origins of parasitism ineleutherengone mites. Zoologischer Anzeiger, 247, 15β45.
8. Cross EA, Moser JC. 1971. Taxonomy and biology of somePyemotidae (Acarina: Tarsonemoidea) inhabiting bark beetlegalleries in North American conifers. Acarologia, 13, 47β64.
9. De Heij SE, Willenborg CJ. 2020. Connected carabids: Networkinteractions and their impact on biocontrol by carabid beetles.BioScience, 70, 490β500.
10. Gullan PJ, Cranston PS. 2014. The insects: an outline ofentomology, 3rd ed. John Wiley & Sons.
11. Hajiqanbar H, Husband RW, Kamali K, Saboori A, Kamali H.2007. Ovacarus longisetosus n. sp. (Acari: Podapolipidae) fromAmara (Paracelia) saxicola Zimm. (Coleoptera: Carabidae) andnew records of Coccipolipus, Dorsipes, Eutarsopolipus andTarsopolipus from Iran. International Journal of Acarology, 33,241β244.
12. Hajiqanbar H, Mortazavi A. 2012. First record of the myzusspecies group (Acari: Podapolipidae: Eutarsopolipus Berlese,1911) from Asia, with the description of two new speciesparasitising carabid beetles. Systematic Parasitology, 83, 189β202.
13. Hajiqanbar H, Mortazavi A, Khaustov A. 2019. Two newspecies of Eutarsopolipus (Acari: Prostigmata: Podapolipidae)parasitizing Syntomus lateralis (Coleoptera: Carabidae) fromIran. Zootaxa, 4647, 154β167.
14. Halliday B. 2019. The enemy of my parasite is my friend: thepossible role of predatory mites as biological control agents of pestbeetles in soil. International Journal of Acarology, 45, 189β196.
15. Houck MA, OConnor BM. . 1991. Ecological and evolutionarysignificance of phoresy in the Astigmata. Annual Review ofEntomology, 36, 611β636.
16. Hunter PE, Rosario RM. 1988. Associations of Mesostigmatawith other arthropods. Annual Review of Entomology, 33,393β417.
17. Hurst GD, Sharpe RG, Broomfield AH, Walker LE, MajerusTM, Zakharov IA, Majerus ME. 1995. Sexually transmitteddisease in a promiscuous insect, Adalia bipunctata. EcologicalEntomology, 20, 230β236.
18. Husband RW. 1984. Dilopolipus, Panesthipolipus, Peripolipusand Stenopolipus, new genera of Podapolipidae (Acarina) fromthe Indo-Australian Region. International Journal of Acarology,10, 251β269.
19. Husband RW. 2007. Regenpolipus hexastichus n. sp. (Acari:Podapolipidae), parasite of Anthia hexastichum Gerstaecker(Coleoptera: Carabidae), from Kenya. International Journal ofAcarology, 33, 327β332.
20. Husband RW, Khaustov AA. 2004. A new species of Eutar-sopolipus (Acari: Podapolipidae) from Calathus fuscipes(Coleoptera: Carabidae) from Ukraine. International Journal ofAcarology, 30, 329β333.
21. Husband RW, Kurosa K. 2000. Two new genera and a newspecies of mites (Acari: Podapolipidae) associated with weevils(Coleoptera: Curculionidae) in Argentina. International Journalof Acarology, 26, 247β255.
22. Husband RW, Kurosa K. 2013. Eutarsopolipus asiaticus sp. nov.(Acari: Podapolipidae), subelytral parasite of Chlaenius costigerChaudoir (Coleoptera: Carabidae) from Japan. Systematic &Applied Acarology, 18, 61β70.
23. Husband RW, Raros LA. 1989. New species of Podapolipusand a new species of Eutarsopolipus (Acari: Podapolipidae)from the Philippines. Philippine Entomologist, 7, 525β536.
24. International Code of Zoological Nomenclature. 1999. TheInternational Trust for Zoological Nomenclature (ed.), London.
25. Kaliszewski M, Athias-Binche F, Lindquist EE. 1995. Parasitismand parasitoidism in Tarsonemina (Acari: Heterostigmata) andevolutionary considerations. Advances in Parasitology, 35,335β367.
26. Katlav A, Hajiqanbar H. 2018. First description of male andlarval female of parasitic mite Eutarsopolipus abdominis (Acari:Podapolipidae) with redescription of the adult female. Journal ofParasitology, 104, 1β9.
27. Katlav A, Hajiqanbar H, Talebi AA. 2014. Dorsipes caspiusn. sp. (Acari: Podapolipidae), a subelytral parasite of Pterostichuscaspius (Menetries) (Coleoptera: Carabidae) with notes on hostrange of the genus and the distribution of the platysmae group.Systematic Parasitology, 89, 117β132.
28. Katlav A, Hajiqanbar H, Talebi AA. 2014. First record of thegenus Acanthomastix Mahunka, 1972 (Acari: Dolichocybidae)from Asia, with the description of a new species. InternationalJournal of Acarology, 40, 7β14.
29. Katlav A, Hajiqanbar H, Talebi AA. 2015. First record of thegenus Aethiophenax (Acari: Acarophenacidae) from Asia,redefinition of the genus and description of a new species.Journal of Asia Pacific Entomology, 18, 389β395.
30. Katlav A, Hajiqanbar H, Talebi AA. 2015. Pseudopygmephorellusmazandaranicus sp. nov. (Acari: Heterostigmata: Pygmephoridae),phoretic on scarabaeid dung beetles (Coleoptera: Scarabaeidae)from Iran. Zootaxa, 3919, 100β110.
31. Katlav A, Riegler M, Seeman OD. 2020. Tiny hitchhikers andparasites: a review of Australian heterostigmatic mites (Acari:Prostigmata) associated with insects, with description of threenew species. Austral Entomology, 59, 401β421.
32. Lindquist EE. 1986. The world genera of Tarsonemidae (Acari:Heterostigmata): a morphological, phylogenetic, and systematicrevision, with a reclassification of family-group taxa in theHeterostigmata. Memoirs of the Entomological Society ofCanada, 118, 1β517.
33. Linz DM, Hu AW, Sitvarin MI, Tomoyasu Y. 2016. Functionalvalue of elytra under various stresses in the red flour beetle.Tribolium castaneum. Scientific Reports, 6, 34813.
34. Lovei GL, Sunderland KD. 1996. Ecology and behavior ofground beetles (Coleoptera: Carabidae). Annual Review ofEntomology, 41, 231β256.
35. Mortazavi A, Hajiqanbar H, Lindquist EE. 2018. A new familyof mites (Acari: Prostigmata: Raphignathina), highly specializedsubelytral parasites of dytiscid water beetles (Coleoptera:Dytiscidae: Dytiscinae). Zoological Journal of the LinneanSociety, 184, 695β749.
36. Moser JC, Konrad H, Kirisits T, Carta LK. 2005. Phoretic mitesand nematode associates of Scolytus multistriatus and Scolytuspygmaeus (Coleoptera: Scolytidae) in Austria. Agricultural andForest Entomology, 7, 169β177.
37. Poinar GO Jr, Curcic BP, Cokendolpher JC. 1998. Arthropodphoresy involving pseudoscorpions in the past and present. ActaArachnologica, 47, 79β96.
38. Regenfuss H. 1968. Untersuchungen zur morphologie, system-atik und oekologie der Podapolipidae (Acarina: Tarsonemini).Zeitschrift fΓΌr wissenschaftliche Zoologie, 177, 183β282.
39. Regenfuss H. 1972. Γber die Einnischung synhospitalerParasitenarten auf dem WirtskΓΆrper: Untersuchungen anektoparasitischen Milben (Podapolipidae) auf laufkΓ€fern(Carabidae). Journal of Zoological Systematics and Evolution-ary Research, 10, 44β65.
40. Schroder RF. 1982. Effect of infestation with Coccipolipusepilachnae Smiley (Acarina: Podapolipidae) on fecundity andlongevity of the Mexican bean beetle. International Journal ofAcarology, 8, 81β84.
20 A. Katlav et al.: Parasite 2021, 28, 75
![Page 21: Sheltered life beneath elytra: three new species of ...](https://reader034.fdocuments.us/reader034/viewer/2022050415/6270ef1363a8375213003b34/html5/thumbnails/21.jpg)
41. Seeman OD. 2019. New species of Eutarsopolipus (Trombid-iformes: Podapolipidae) from the pterostichine genera Castel-naudia and Trichosternus (Coleoptera: Carabidae) in Australia.Zootaxa, 4717, 206β230.
42. Seeman OD. 2019. Two new species of Australian Eutarsopoli-pus (Acariformes: Podapolipidae) from Nurus medius (Coleop-tera: Carabidae). Zootaxa, 4647, 134β153.
43. Seeman OD. 2020. Unusual new species of Australian Eutar-sopolipus (Acariformes: Podapolipidae) from Clivina (Coleop-tera: Carabidae). Annales Zoologici, 70, 425β438.
44. Seeman OD. 2021. Contrasting species diversification ofEutarsopolipus (Acariformes: Podapolipidae) on Castelnaudiaand Notonomus (Coleoptera: Carabidae). Zootaxa, 4971, 1β74.
45. Seeman OD, Nahrung HF. 2013. Two new species ofChrysomelobia Regenfuss, 1968 (Acariformes: Podapolipidae)from Paropsis charybdis StΓ₯l (Coleoptera: Chrysomelidae).Systematic Parasitology, 86, 257β270.
46. Stork NE, McBroom J, Gely C, Hamilton AJ. 2015. Newapproaches narrow global species estimates for beetles, insects,and terrestrial arthropods. Proceedings of the National Academyof Sciences, 112, 7519β7523.
47. Trach VA, Khaustov AA. 2007. A review of mites of the familyCanestriniidae (Acari, Astigmata) associated with the leaf
beetles (Coleoptera, Chrysomelidae) in Ukraine with descriptionof a new species of the genus Pseudamansia. Vestnik Zoologii,41, 267β272.
48. van de Kamp T, Riedel A, Greven H. 2016. Micromorphologyof the elytral cuticle of beetles, with an emphasis on weevils(Coleoptera: Curculionoidea). Arthropod Structure & Develop-ment, 45, 14β22.
49. Walter DE, Lindquist EE, Smith IM, Cook DR, Krantz GW.2009. Order Trombidiformes, in A manual of acarology, 3rd edn.Krantz GW, Walter DE, Editors. Texas Tech University Press:Texas, USA. p. 233β420.
50. Walter DE, Seeman OD. 2017. A new species of Parac-arophenax (Acariformes: Acarophenacidae) with a new meansof phoretic attachment. International Journal of Acarology, 43,329β335.
51. Wohltmann A, GabryΕ G, Mazkol J. 2006. Acari: Acari:Terrestrial Parasitengona inhabiting transient biotopes, inSΓΌΓwasserfauna von Mitteleuropa, Chelicerata: Araneae/AcariI. Spektrum. Gerecke R, Editor. Elsevier: MΓΌnchen. p. 158β240.
52. Zawal A, PeΕ‘iΔ V. 2018. The diversity of assemblages of watermites from Lake Skadar and its catchment, in The Skadar/Shkodra Lake environment. PeΕ‘iΔ V, Karaman GS, KostianoyAG, Editors. Springer: Berlin.
Cite this article as: Katlav A, Hajiqanbar H, Riegler M & Seeman OD. 2021. Sheltered life beneath elytra: three new species ofEutarsopolipus (Acari, Heterostigmatina, Podapolipidae) parasitizing Australian ground beetles. Parasite 28, 75.
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