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On the Taxonomy of Chigger Mites (Acariformes: Trombiculidae) Parasitizing Birds in Thailand and Malaysia, with the Description of a New Species

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11 September 2023

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13 September 2023

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Abstract
Chigger mites were collected from 65 bird species in different regions of Thailand and Malaysia. In total, 21 species were recorded. The previously unknown fauna of chiggers parasitizing shorebirds of Thailand included one new species, Neacariscus (Whartonacarus) ... n. sp. as well as Neacariscus (Neacariscus) pluvius (Wharton, 1945) previously recorded only in Oceania; Neacariscus (Whartonacarus) shiraii (Sasa, Kano & Obata, 1952) (known from Oceania and Japan); and Neacariscus (Whartonacarus) sulae (Oudemans, 1910) and Schoengastia archaea (Taufflieb, 1960), both described from West Africa. Toritrombicula kirhocephales Goff, 1982 described from Papua New Guinea was for the first time recorded in Asia (Thailand). Neoschoengastia gallinarum (Hatori, 1920) and Helenicula comata (Womersley, 1952) were for the first time recorded in Thailand. Ascoschoengastia lorius (Gunther, 1939) was for the first time recorded in Malaysia. Finally, Ericotrombidium cosmetopode (Vercammen-Grandjean & Langston, 1971) described from free larvae, was for the first time recorded on a host (bird Pellorneum ruficeps Swainson); and Leptotrombidium miculum (Traub & Audy, 1954) was for the first time recorded on a bird host. New host species were recorded for the large part of the collected chigger species.
Keywords: 
Subject: Biology and Life Sciences  -   Parasitology

1. Introduction

Southeast Asia is one of the best-studied regions of the world with regard to the taxonomy of chigger mites. The leading countries of this region are Malaysia, with 202 recorded chigger species, and Thailand, with 156 species (Stekolnikov 2021a). However, the chigger species specific to some groups of hosts, such as bats or birds, are under-explored even in such territories. In general, extensive investigations focused on bird chiggers are rare, as compared with chiggers from mammals (Trnka et al. 2022). Among the recent works on Southeast Asian chiggers, a paper based on the collections from Vietnam should be consulted (Kaluz et al. 2016), in which three species of Neoschoengastia Ewing, 1929 (including two new to science), Odontacarus audyi (Radford, 1946), Helenicula scanloni (Domrow & Nadchatram, 1964), Neotrombicula elegans Schluger, 1966, and six species of Leptotrombidium Nagayo et al., 1916 were recorded on seven bird species. Noteworthy is that all of them, with the exception of H. scanloni, were unknown previously in Vietnam, despite the fact that this country is the third most studied for chiggers in Southeast Asia, after Malaysia and Thailand (Stekolnikov 2021a).
Our collections performed over the timeframe of 2020 – 2022 obtained a large number of chiggers from birds across Thailand and Malaysia (1,531 individual hosts of 65 species), including one new trombiculid species and a series of unexpected new geographic and host records. Here, we provide a list of these chigger-host records compared with published data, alongside morphological descriptions of our specimens including one new species and notes on the genus Neacariscus and its constituent subgenera. The purpose of this work is to serve as foundation for quantitative ecological studies of bird-chigger relationships in Southeast Asia that will be presented in a separate publication; thus, we limit our discussion to qualitative findings in the current study.

2. Material and Methods

2.1. Approvals for bird sampling

In Peninsular Malaysia, the animal handling work was subjected to the approvals and guidelines from the Universiti Malaya Institutional Animal Care and Use Committee (Permission No: G8/07052020/09012020-01/R), and a wildlife research permit was obtained from the Department of Wildlife and National Parks (DWNP, Permit No. W-00064-15-21). Separate permits were obtained from the Sarawak Forest Department, National Parks and Nature Reserves, East Malaysia (Permit No. SFC.810-4/6/1 – 033 for collection of biological resources and Permit No. WL 13/2022 for permission to enter parks). In Thailand, the study was approved by the Faculty of Tropical Medicine Animal Care and Use Committee (FTM-ACUC, Certification No. 011/2020E) and the Department of National Park Wildlife and Plant Conservation (DNP), Ministry of Natural Resources and Environment (permission letter No. 0907.4/21997). The study was also approved by the Animal Welfare and Ethical Review Body of the University of Liverpool with reference nos. AWC0219 (Malaysia) and AWC0179 (Thailand).

2.2. Study sites and bird trapping

Bird trapping was conducted across eight provinces of Thailand (Nan, Nakhon Sawan, Kanchanaburi, Samut Sakhon, Phetchaburi, Rayong, Trang and Satun) and seven states of Malaysia (Selangor, Pahang, Terengganu, Kedah, Perak, Johor and Sarawak). Precise sampling locations are listed in Table 1 and Table 2, with GPS co-ordinates provided in the Results. Names of birds follow the IOC World Bird List (Gill et al. 2023). Each location was sampled for 1 – 6 days in proportion to the scale of the site and bird abundance, and some locations were sampled on more than one occasion. Multiple strategies were applied to the capture of different bird species, with expert assistance from DWNP in Malaysia and DNP in Thailand. Passeriformes, Coraciiformes, Piciformes, Podargiformes, Strigiformes, Gruiformes, and Cuculiformes were trapped using mist nets, which were checked every 15 to 20 minutes depending on the abundance of birds in the area of deployment. In Malaysia, Galliformes were captured using large scoop nets at night from 20:00 h to 00:00 h; whereas in Thailand, domestic chickens (only sampled in Ban Huai Muang village, Nan province) were brought by their owners for examination by the field team. Finally, waders (Charadriiformes) in Thailand were captured by cannon netting at diurnal high-tide roosts. In Thailand, birds were identified with reference to Treesucon & Limparungpatthanakij (2018) and Lekagul & Round (1991); whereas in Malaysia, the key reference work was Robson (2018).

2.3. Chigger collection and examination

Trapped birds were handled gently during examination for the presence of chigger mites. Chiggers attached to the birds were removed carefully using fine forceps and placed in tubes containing 70% ethanol. These were labelled individually according to the number assigned to each of the bird hosts.
A survey of free-living chiggers in the environment was also conducted at Ban Huai Muang village after chigger infestations on chickens were confirmed. Briefly, square black plastic plates were deployed on the ground in a formation of six grids in different habitats inside and around the village (Kundin et al. 1966). The plates were carefully examined for chiggers within 10 minutes following deployment; the free-living chiggers were removed using a fine paintbrush and transferred into 80% ethanol.
With the aid of a dissecting microscope, ~10 – 20% of chiggers from each host or environmental sample were selected by differences in size and appearance as subsamples representative of the chigger species composition. These were then washed in sterile water to remove ethanol, arranged in a dorsoventral position on a glass slide with a drop of Berlese fluid (TCS Bioscience Ltd, UK), and flattened with a round coverslip. Slides were allowed to dry at room temperature before microscopic examination. Initial identification (usually to subgenus) was performed using a combination of brightfield and autofluorescence microscopy (Kumlert et al. 2018). A list of the material from Thailand is given in Table 1 and from Malaysia in Table 2.
A proportion of the microscope slides was shipped to the Zoological Institute of the Russian Academy of Sciences (ZIN, Saint Petersburg, Russia) and examined by AAS under a Leica DM 2500 compound microscope (Leica Microsystems GmbH, Wetzlar, Germany) using differential interference contrast (DIC). Microphotographs were taken by means of a Leica DMC 4500 digital camera. Morphological drawings were prepared using a drawing tube. Measurements were taken using an ocular micrometer, on an MBI-3 microscope (LOMO plc, Saint Petersburg, Russia) with phase contrast optics. Identification of chigger mites to species was performed with the use of taxonomic revisions published by Nadchatram (1967a, b), Nadchatram & Traub (1971), Vercammen-Grandjean & Langston (1976), Stekolnikov (2013), and other sources cited below, and also by direct comparison with previously identified specimens from the collection of ZIN.
Holotype and paratypes of the new species are deposited in ZIN. Most of the Malaysian material was returned to the Universiti Malaya (UM); the material from Thailand is mainly deposited in the University of Liverpool (UoL).

3. Results

Genus Neacariscus Vercammen-Grandjean, 1960
Type species — Acariscus pluvius Wharton, 1945, by original designation.
Diagnosis — SIF = 7BS(7B)-N-2-3(2)111.1000; fPp = B/B/NNN or B/N/NNN; fCx = 1.1.(1 – 3); fSt = 2.2(4); scutum trapezoidal, with pronounced anterolateral shoulders, with dense puncta; flagelliform sensilla (trichobothria) branched in distal part, nude, or covered by cilia; eyes 2 + 2, often very large; solenidion (ω) of palpal tarsus rod-like or long and slender; parasubterminala (z) nude.
Remarks — The genus Acariscus Ewing, 1943 was a heterogeneous group. Its type species, Trombicula flui van Thiel, 1930, was synonymized with Eutrombicula batatas (Linnaeus, 1758) by Michener (1946). Thus, Acariscus became a synonym of Eutrombicula Ewing, 1938. Subsequently, Vercammen-Grandjean (1960) described a new genus Neacariscus comprising two subgenera – the nominative, with the type species Acariscus pluvius, and Whartonacarus Vercammen-Grandjean, 1960, with the type species Trombicula thompsoni Brennan, 1953. This work did not include lists of species. Loomis (1966) considered Whartonacarus as a subgenus of Toritrombicula Sasa, Hayashi & Kawashima, 1953; he included in it five species. At the same time, he synonymized Neacariscus with Toritrombicula and placed A. pluvius in the nominative subgenus of the latter. However, Vercammen-Grandjean & Langston (1976) disagreed with that decision and restored the generic status of Neacariscus, with three subgenera – the nominative (one species, A. pluvius), Whartonacarus (eight species), and Loomiscus Vercammen-Grandjean & Langston, 1976 (one species). Brennan & Goff (1977) included Whartonacarus in their key to genera as a genus, without a discussion. Later many authors used this name as generic (Domrow & Lester 1985; Hoffmann 1990; Mertins et al. 2009; Stekolnikov & González-Acuña 2015). However, none of them discussed the status of Neacariscus. We note the following:
1. Both Neacariscus pluvius and all species of Whartonacarus significantly differ from Toritrombicula by a) a trapezoidal scutum with prominent anterolateral shoulders vs. a subrectangular scutum without anterolateral shoulders; b) two-pronged palpal claw vs. three-pronged; c) nude parasubterminala (z) vs. branched; d) presence of mastitarsala. This complex of traits definitely constitutes a difference at the generic level.
2. All species of Whartonacarus differ from N. pluvius by the presence of palpal subterminala (ζ) and by the presence of three genualae I (σ) vs. two. These differences can at most justify the subgeneric status.
Therefore, we follow the system of Vercammen-Grandjean & Langston (1976), which included the genus Neacariscus with three subgenera – Neacariscus, Whartonacarus, and Loomiscus. The latter subgenus includes one species, Neacariscus (Loomiscus) dupliseta (Loomis, 1966), which differs from the species of Neacariscus (Whartonacarus) by the presence of multiple additional nude setae on the genuae of all legs. The subgenus Neacariscus (Whartonacarus) includes the following species: N. (W.) anous (Wharton, 1945), N. (W.) chaetosa (Brennan & Jones, 1961), N. (W.) floridensis (Mertins, 2009) (in Mertins et al. 2009), n. comb., N. (W.) nativitatis (Hoffmann-Sandoval, 1950), N. (W.) oceanica (Brennan & Amerson, 1971), N. (W.) shiraii (Sasa, Kano & Obata, 1952), N. (W.) sulae (Oudemans, 1910), and N. (W.) thompsoni (Brennan, 1953).
Subgenus Neacariscus (Neacariscus) Vercammen-Grandjean, 1960
Type species — Acariscus pluvius Wharton, 1945, by original designation.
Diagnosis — SIF = 7B-N-2-2111.1000; fPp = B/B/NNN; fCx = 1.1.1; fSt = 2.2; flagelliform sensilla (trichobothria) branched in distal part; eyes 2 + 2, of normal size; solenidion (ω) of palpal tarsus long and slender.
Neacariscus (Neacariscus) pluvius (Wharton, 1945)
(Figure 1A)
Distribution and hosts — This species was described from Bougainville Isl. (Solomon Islands archipelago) and from Guam (Mariana Islands), ex Anous stolidus (L.), Anous tenuirostris (Temminck) (Charadriiformes: Laridae), Pluvialis dominica (Statius Müller) (Charadriiformes: Charadriidae), and Tringa incana (J.F. Gmelin) (syn. Heteroscelus incanus) (Charadriiformes: Scolopacidae). Loomis (1966) also reported N. pluvius from Ulithi Atoll (Caroline Islands), ex Tringa incana and Sterna sumatrana Raffles (Charadriiformes: Laridae). Here this species is for the first time recorded in Asia (Thailand). Charadrius leschenaultii and Xenus cinereus are new host species.
Material examined — Three larvae (ZIN 17963 – 17965) ex two Charadrius leschenaultii and one Xenus cinereus, THAILAND, Trang province, Samran Beach, 7.201348 N, 99.562105 E and 7.209399 N, 99.557854 E, 17 and 19 March 2021, coll. S. Koosakulnirand.
Subgenus Neacariscus (Whartonacarus) Vercammen-Grandjean, 1960
Type species — Trombicula thompsoni Brennan, 1953, by original designation.
Diagnosis — SIF = 7BS-N-2-3111.1000; fPp = B/B/NNN or B/N/NNN; fCx = 1.1.(1 – 3); fSt = 2.2(4); flagelliform sensilla (trichobothria) branched in distal part, nude, or covered by cilia; eyes 2 + 2, often very large; solenidion (ω) of palpal tarsus rod-like or long and slender.
Neacariscus (Whartonacarus) ... Stekolnikov, n. sp.
(Figure 1B,C, Figure 2A–E, Figure 3A,C–E and Figure 4)
Diagnosis — SIF = 7BS-N-2-3111.1000; fPp = B/N/NNN; fCx = 1.1.1; fSt = 2.4; PL > AL ≥ AM; fD = 2H-11(13)-11(10)-(8 – 12)-(7 – 10)+(0 – 3); DS = 42 – 49; V = 64 – 84; NDV = 107 – 133; Ip = 1642 – 1697; eyes 2 + 2, very large; flagelliform sensilla (trichobothria) with few branches in distal part; sensillary bases anterior to level of PL; solenidion (ω) of palpal tarsus long and slender; leg tarsi with multiple sclerite bars. Standard measurements are given in Table 3.
Description of larva (based on holotype, three paratypes ZIN 17983 – 17985 and three additional specimens ZIN 17976 – 17978)
Idiosoma (Figure 1B,C and Figure 2A–E) — Eyes 2 + 2, very large; 42 – 49 sharply-pointed dorsal idiosomal setae (including two clearly separated humeral setae) moderately covered with thin barbs; 1st posthumeral row (C excluding humeral setae) with 11 (in five specimens) or 13 (in two specimens) setae, 2nd row (D) with 11 (in four specimens) or 10 (in three specimens) setae, 3rd (E) row with 8 – 12 setae, 4th row (F) with 7 – 10 setae, remaining caudal setae 0 – 3 in number; in holotype, fD = 2H-11-11-10-10; two sternal setae between coxae I and four sternal setae between coxae III; 64 – 84 ventral setae; NDV = 107 – 133.
Gnathosoma (Figure 3C–E) — Cheliceral blade with tricuspid cap; cheliceral base densely punctate; gnathobase (infracapitulum) densely punctate, bears one pair of branched gnathocoxal (tritorostral) setae; galeal (deutorostral) seta nude; palpal claw (odontus) with two prongs, main inner and additional smaller outer; palpal femur, genu, and tibia with puncta; palpal femoral seta thick, covered with long branches; palpal genual and tibial setae thin, nude; palpal tarsus with seven branched setae, nude subterminala (ζ) and very long, pointed, curved tarsala (ω).
Scutum (Figure 1B and Figure 3A) —Trapezoidal, with pronounced anterolateral shoulders, with dense puncta; anterior margin sinuous, lateral margins concave, posterior margin almost straight in middle part and obliquely cut at edges; AM placed at level of AL; sensillary (trichobothrial) bases placed anterior to level of PL (PSB – P-PL = 4 – 10 µm, mean 8); PL > AL ≥ AM; all scutal setae barbed similarly to dorsal idiosomal setae; sensilla (trichobothria) flagelliform, with ca. four branches in distal part.
Legs (Figure 4) — All legs 7-segmented (with divided femur), with one pair of claws and claw-like empodium; leg tarsi with multiple sclerite bars. Leg I: coxa with one branched seta (1B); trochanter 1B; basifemur 1B; telofemur 5B; genu 4B, three genualae (σ), microgenuala (κ); tibia 8B, 2 distal tibialae (φ) in tandem, microtibiala (κ); tarsus 22B, tarsala (ω), famulus (ε) distal to tarsala, subterminala (ζ), nude parasubterminala (z), pretarsala (ζ). Leg II: coxa 1B; trochanter 1B; basifemur 2B; telofemur 4B; genu 3B, genuala (σ); tibia 6B, two tibialae (φ) in tandem; tarsus 18B, tarsala II (ω), famulus (ε) behind tarsala, pretarsala (ζ). Leg III: coxa 1B; trochanter 1B; basifemur 2B; telofemur 3B; genu 3B, long genuala (σ); tibia 6B, long tibiala (φ); tarsus 16B, mastitarsala in middle part of segment.
Type material — Larval holotype (ZIN 17980, BTRA045) ex Calidris tenuirostris, THAILAND, Trang province, Koh Libong, 7.243892 N, 99.451315 E, 16 March 2021, coll. S. Koosakulnirand; four paratypes (ZIN 17983 – 17986) ex four Calidris tenuirostris, THAILAND, Trang province, Samran Beach, 7.209399 N, 99.557854 E, 19 March 2022, coll. S. Koosakulnirand.
Additional material — Three larvae (ZIN 17976 – 17978) ex two Calidris tenuirostris and one Charadrius lechenaultii, THAILAND, Satun province, Thung Sabo, 7.042112 N, 99.671153 E, 14 March 2021, coll. S. Koosakulnirand.
Two larvae (UoL) ex one Calidris tenuirostris, THAILAND, Trang province, Koh Libong, 7.243892 N, 99.451315 E, 16 March 2021, coll. S. Koosakulnirand (same data as for holotype); nine larvae (UoL) ex four C. tenuirostris (6) and two Xenus cinereus (3), THAILAND, Trang province, Samran Beach, 7.209399 N, 99.557854 E, 19 March 2022, coll. S. Koosakulnirand; one larva (UoL) ex Calidris ruficollis, THAILAND, Trang province, Samran Beach, 7.201348 N, 99.562105 E, 15 March 2021, coll. S. Koosakulnirand; six larvae (UoL) ex three C. tenuirostris, THAILAND, Satun province, Thung Sabo, 7.042112 N, 99.671153 E, 14 March 2021, coll. S. Koosakulnirand.
Etymology — The species epithet ... refers to the Andaman Sea, on the shore of which the type locality is situated.
Differential diagnosis — The new species is similar to Neacariscus (Whartonacarus) shiraii but differs from it in exhibiting fewer idiosomal setae (DS = 42 – 49 vs. 77 – 92; NDV = 107 – 133 vs. 162 – 190); the rows of dorsal idiosomal setae are unpaired (only few additional setae are present posterior to the central part of 1st row); in its longer scutum with more prominent posterior margin (ASB = 39 – 42 vs. 33 – 36, PSB = 23 – 27 vs. 19 – 21, SD = 63 – 66 vs. 52 – 57, and P-PL = 14 – 20 vs. 11 – 13); and in longer legs (Ip = 1642 – 1697 vs. 1381 – 1550 and TaIIIL = 178 – 189 vs. 160 – 162).
Neacariscus (Whartonacarus) shiraii (Sasa, Kano & Obata, 1952)
(Figure 2F,G and Figure 3B)
Diagnosis — SIF = 7BS-N-2-3111.1000; fPp = B/N/NNN; fCx = 1.1.1; fSt = 2.4; PL > AL ≥ AM (PL > AM > AL); fD = 2H-23-24(20)-18(19)-13(11)-(0 – 2); DS = 77 – 92; V = 85 – 98; NDV = 162 – 190; Ip = 1381 – 1550; eyes 2 + 2, very large; flagelliform sensilla (trichobothria) with few branches in distal part; sensillary bases anterior to level of PL; solenidion (ω) of palpal tarsus long and slender; leg tarsi with multiple sclerite bars. Standard measurements are given in Table 3.
Distribution and hosts — This species was described from Tokyo (Japan), ex Pluvialis dominica (syn. Charadrius dominicus). Later it was recorded on Heron Isl. (Australia, Queensland), ex Limosa lapponica (L.) (Charadriiformes: Scolopacidae); on Guam, ex P. dominica; on Okinawa Isl. (Japan), ex Sterna dougalli Montagu (Charadriiformes: Laridae); and in Mexico (Sonora), ex Haematopus palliatus Temminck (Charadriiformes: Haematopodidae) (Vercammen-Grandjean & Langston 1976). Here it is for the first time recorded in Thailand and on Xenus cinereus.
Material examined — Two larvae (ZIN 17981, 17982) ex two Xenus cinereus, THAILAND, Trang province, Samran Beach, 7.209399 N, 99.557854 E, 19 March 2022, coll. S. Koosakulnirand.
Remarks — The form of N. (W.) shiraii from Mexico, with DS = 44 and V = 44 (Loomis 1966; Mertins et al. 2009), probably belongs to some other species.
Neacariscus (Whartonacarus) sulae (Oudemans, 1910)
(Figure 1D)
Diagnosis — SIF = 7BS-N-2-3111.1000; fPp = B/N/NNN; fCx = 1.1.1; fSt = 2.2; PL >> AL = AM; fD = 2H-10-10-6-2-2; DS = 32 – 36; V = 50 – 65; NDV = 86 – 101; Ip = 1181 – 1206; eyes 2 + 2, of moderate size; flagelliform sensilla (trichobothria) with short branches in distal part; sensillary bases anterior to level of PL; solenidion (ω) of palpal tarsus long and slender. Standard measurements are given in Table 4.
Distribution and hosts — This species was described from a single specimen (holotype) collected in an unknown locality of West Africa, ex Morus capensis (Lichtenstein) (syn. Sula capensis) (Suliformes: Sulidae). Zumpt (1961) referred to the host as Morus bassanus (L.). Here it is for the first time recorded in Asia and on Pluvialis squatarola.
Material examined — Two larvae (ZIN 17962, 18168) ex Pluvialis squatarola, THAILAND, Trang province, Koh Libong, 7.243892 N, 99.451315 E, 16 March 2021, coll. S. Koosakulnirand.
Remarks — The species was initially described briefly (Oudemans 1910): a fully illustrated redescription was published two years later (Oudemans 1912). Fuller (1952) and Vercammen-Grandjean & Langston (1976) also examined and redescribed the holotype. According to the latter authors, fD of the holotype is 2H-8-8-8-4-4-2, which seems different from the fD in our material (2H-10-10-6-2-2). However, comparison of the drawing made by Oudemans (1912, fig. A1) with our specimens shows that the divergence could be caused by a different mode of preparation and, probably, unequal levels of mite engorgement.
Fuller (1952) described the ventral palpal tibial seta of the holotype as branched, which agrees with the drawing made by Oudemans (1912, fig. A4). However, Vercammen-Grandjean & Langston (1976) drew and described this seta as nude, as in our material. The scutum on their drawing (pl. 282) bears multiple transverse striae, which were not mentioned by Oudemans (1912) and Fuller (1952). They are also absent in our specimens. Therefore, our material may represent a new species close to N. sulae. Additional collections in Africa and one more re-examination of the N. sulae holotype would be desirable to reach a final conclusion.
Genus Toritrombicula Sasa, Hayashi & Kawashima, 1953
Type species — Trombicula (Toritrombicula) hasegawai Sasa, Hayashi & Kawashima, 1953, by original designation.
Diagnosis — SIF = 7B-N(B)-3-2111.0000; fPp = B/B/NNN or B/B/NNB; fCx = 1.1.1; fSt = 2.2; scutum subrectangular, without prominent anterolateral shoulders, with puncta; flagelliform sensilla (trichobothria) branched in distal part; eyes 2 + 2, anterior pair often very large; solenidion (ω) of palpal tarsus rod-like; parasubterminala (z) branched.
Toritrombicula densipiliata (Walch, 1922)
Distribution and hosts — Indonesia, Malaysia, Papua New Guinea, Philippines, Taiwan, Thailand, ex 21 bird species (Stekolnikov 2021a). Arachnothera longirostra, Batrachostomus stellatus, Cacomantis sepulcralis, Copsychus malabaricus, Cyanoptila cyanomelana, Cyornis tickelliae, Ficedula dumetoria, Geokichla sibirica, Lanius tigrinus, Larvivora cyane, Malacocincla abbotti, Pellorneum malaccense, Malacocincla sepiaria, Malacopteron cinereum, Malacopteron magnum, Mixornis bornensis, Turdinus macrodactylus, Pellorneum capistratum, Pellorneum ruficeps, Cyornis brunneatus, Cyanoderma erythropterum, Stachyris maculata, Terpsiphone atrocaudata, Tricholestes criniger, and Turdus obscurus are new host species.
Material examined — Two larvae (ZIN 18002, 18003) ex Geokichla citrina and Geokichla sibirica, THAILAND, Rayong province, Koh Mun Nai, 12.612384 N, 101.687574 E, 10 March 2021, coll. S. Koosakulnirand; one larva (ZIN 17935) ex Mixornis bornensis, MALAYSIA, Sarawak State, Sematan Pueh village, 1.831126 N, 109.708966 E, 23 March 2022, coll. P. Rajasegaran.
Five larvae (UM) ex one Pellorneum ruficeps (1), one Cyornis tickelliae (1), two Larvivora cyane (2), and one Malacocincla abbotti (1), MALAYSIA, Kedah State, Langkawi Isl., Gunung Machinchang, 6.429010 N, 99.729852 E, 16 – 17 March 2022, coll. P. Rajasegaran; 41 larvae (UM) ex one Pellorneum nigrocapitatum (1), one Calyptomena viridis (1), one Ficedula dumetoria (3), one Macronus ptilosus (2), one Pellorneum malaccense (2), one Malacocincla sepiaria (1), one Malacopteron cinereum (2), one Malacopteron magnum (1), one Turdinus macrodactylus (1), five Philentoma pyrhoptera (11), two Stachyris maculata (2), three Stachyris poliocephala (12), and one Terpsiphone atrocaudata (1), MALAYSIA, Pahang State, Krau Wildlife Reserve, 3.596982 N, 102.183190 E, 16 – 20 February 2021, coll. P. Rajasegaran; 30 larvae (UM) ex two Actenoides concretus (3), one Arachnothera longirostra (1), one Batrachostomus stellatus (1), five Copsychus malabaricus (11), one Lanius tigrinus (1), two Larvivora cyane (2), three Philentoma pyrhoptera (4), and two Cyornis brunneatus (6), MALAYSIA, Perak State, Behrang Forest Reserve, E101.578552, N3.732175, 19 – 24 September 2022, coll. P. Rajasegaran; two larvae (UM) ex Anthipes solitaris and Turdus obscurus, MALAYSIA, Perak State, Larut Hill, 4.867858 N, 100.779824 E, 9 – 11 March 2022, coll. P. Rajasegaran; four larvae (UM) ex Calyptomena viridis, Copsychus malabaricus, Cyanoptila cyanomelana, and Philentoma pyrhoptera, MALAYSIA, Sarawak State, Gunung Gading National Park, Lundu, 1.692098 N, 109.845157 E, 23 – 26 March 2022, coll. P. Rajasegaran; one larva (UM) ex Mixornis bornensis, MALAYSIA, Sarawak State, Sematan Pueh village, 1.831126 N, 109.708966 E, 23 March 2022, coll. P. Rajasegaran; one larva (UM) ex Cacomantis sepulcralis and two larvae (UM) ex one Tricholestes criniger, MALAYSIA, Selangor State, Ulu Gombak Forest Reserve, 3.325987 N, 101.752747 E, 10 – 12 February 2021, coll. P. Rajasegaran; seven larvae (UM) ex four Larvivora cyane (4), two Philentoma pyrhoptera (2), and one Stachyris poliocephala (1), MALAYSIA, Terengganu State, Pasir Raja Forest Reserve, 4.790517 N, 102.996835 E, 22 – 27 October 2021, coll. P. Rajasegaran; 33 larvae (UM) ex 11 Copsychus malabaricus, MALAYSIA, Terengganu State, Redang Isl., 5.780867 N, 103.006183 E, 8 – 13 October 2022, coll. P. Rajasegaran.
Toritrombicula kirhocephales Goff, 1982
(Figure 5A)
Diagnosis — SIF = 7B-N-3-2111.0000; fPp = B/B/NNN; fCx = 1.1.1; fSt = 2.2; PL > AL > AM; fD = 2H-10(11)-8(10)-(6 – 8)-4-2; DS = 32 – 37; V = 26 – 34; NDV = 58 – 68; Ip = 1026 – 1246; eyes 2 + 2, of moderate size; flagelliform sensilla (trichobothria) with short branches in distal part; sensillary bases anterior to level of PL. Standard measurements are given in Table 5.
Distribution and hosts — This species was described ex Pitohui kirhocephalus (R.P. Lesson & Garnot) (Passeriformes: Oriolidae) from Papua New Guinea (East Sepik province). Here it is for the first time recorded in Asia (Thailand) and on all hosts mentioned below.
Material examined — Six larvae (ZIN 18004 – 18009) ex one Larvivora cyane (1), two Pachycephala cinerea (2), one Pycnonotus conradi (1), and one Phylloscopus tenellipes (2), THAILAND, Rayong province, Koh Mun Nai, 12.612384 N, 101.687574 E, 9 – 11 March 2021, coll. S. Koosakulnirand; one larva (ZIN 18010) ex L. cyane, THAILAND, Kanchanaburi province, W Mahidol University Campus, 14.13111 N, 99.14863 E, 10 December 2021, coll. S. Koosakulnirand; two larvae (ZIN 18011, 18012) ex Ficedula albicilla and Pycnonotus conradi, THAILAND, Kanchanaburi province, 5 km NW Mahidol University Campus, 14.15452 N, 99.12069 E, 13 December 2021, coll. S. Koosakulnirand.
19 larvae (UoL) from same localities, same and additional host individuals of the same species plus one larva (UoL) ex Cyornis tickelliae, THAILAND, Kanchanaburi province, W Mahidol University Campus, 14.13111 N, 99.14863 E, 10 December 2021, coll. S. Koosakulnirand; one larva (UoL) ex Pycnonotus aurigaster, THAILAND, Kanchanaburi province, 5 km NW Mahidol University Campus, 14.15452 N, 99.12069 E, 13 December 2021, coll. S. Koosakulnirand.
Remarks — Our specimens deviate from the original description (Goff 1982) in more numerous idiosomal setae (fD = 2H-10(11)-8(10)-8(7)-4-2 vs. 2H-10-8-6-4-2, DS = 34 – 37 vs. 32, V = 31 – 34 vs. 26, NDV = 65 – 68 vs. 58) and slightly longer legs (Ip = 1145 – 1246 vs. 1026 – 1045, TaIIIL = 112 – 121 vs. 98). We estimate this difference as a case of intraspecific variation.
Toritrombicula kirhocephales differs from T. densipiliata in exhibiting fewer idiosomal setae (two humeral setae vs. four; 1st posthumeral row simple vs. double; DS = 34 – 37 vs. 50 – 65; NDV = 58 – 65 vs. about 104) (Goff 1982; Nadchatram 1967b; Vercammen-Grandjean & Langston 1976).
Toritrombicula uphami Nadchatram, 1967
(Figure 5B)
Distribution and hosts — Described ex Pitta sordida (Statius Müller) (Passeriformes: Pittidae) in Malaysia. Cyornis tickelliae, Enicurus ruficapillus, and Pellorneum nigrocapitatum are new host species.
Material examined — Two larvae (ZIN 18172, 18173) ex Enicurus ruficapillus and Pellorneum nigrocapitatum, MALAYSIA, Terengganu State, Pasir Raja Forest Reserve, 4.790517 N, 102.996835 E, 24 October 2021, coll. P. Rajasegaran; one larva (ZIN 18169) ex Cyornis tickelliae, MALAYSIA, Kedah State, Langkawi Isl., Gunung Machinchang, 6.429010 N, 99.729852 E, 15 March 2022, coll. P. Rajasegaran.
Genus Neoschoengastia Ewing, 1929
Type species — Schoengastia americana Hirst, 1921, by original designation.
Diagnosis — SIF = 7B(7BS)-B-3-(2–3)111.0(1)000; cheliceral blade with tricuspid cap; scutum trapezoidal, with anterolateral shoulders, covered by puncta and cuticular striations around sensillary bases, with almost straight, concave or bilobate posterior margin; sensillary bases far anterior to PL; usually AL > PL; sensilla pyriform or globose, covered with setules; eyes 2 + 2; parasubterminala (z) branched; onychotriches frequently present.
Neoschoengastia longipes Nadchatram, 1967
(Figure 5C)
Diagnosis — SIF = 7BS-B-3-3111.0000; fPp = B/B/BNB; fCx = 1.1.3; fSt = 2.2; AL ≥ PL > AM; fD = 2H-6-6-4-6-4-2 (6-6-4-4-6, 6-6-6-6-4, and other similar variants); Ip = 1140 – 1294; DS = 28 – 31; V = 19 – 25; NDV = 50 – 54; onychotriches present on claws and empodium. Standard measurements are given in Table 6.
Distribution and hosts — THAILAND, ex 14 bird species (Stekolnikov 2021a). Alcedo atthis, Calliope calliope, Larvivora cyane, and Otus sunia are new host species.
Material examined — Two larvae (ZIN 17946, 17947) ex Alcedo atthis and Larvivora cyane, THAILAND, Rayong province, Koh Mun Nai, 12.612384 N, 101.687574 E, 9 March 2021, coll. S. Koosakulnirand; six larvae (ZIN 17948 – 17953) ex four Copsychus malabaricus (4), Cyornis tickelliae (1), and L. cyane (1), THAILAND, Kanchanaburi province, SW Mahidol University Campus, 14.11806 N, 99.15253 E, 8 – 9 December 2021, coll. S. Koosakulnirand; one larva (ZIN 17954) ex C. malabaricus, THAILAND, Kanchanaburi province, W Mahidol University Campus, 14.13111 N, 99.14863 E, 10 December 2021, coll. S. Koosakulnirand.
A great many (42) larvae (UoL) from the same localities, same and additional individuals of the same host species, plus 13 larvae ex four C. malabaricus (7), two C. tickelliae (2), and three L. cyane (4), THAILAND, Kanchanaburi province, W Mahidol University Campus, 14.134225 N, 99.1374365 E, 11 December 2021, coll. S. Koosakulnirand; nine larvae ex two Calliope calliope (4), one L. cyane (3), and one Otus sunia (2), THAILAND, Kanchanaburi province, NW Mahidol University Campus, 14.154772 N, 99.1193296 E, 12 December 2021, coll. S. Koosakulnirand.
Remarks — Neoschoengastia longipes belongs to the group of species close to N. americana, which is characterized by the presence of three setae on coxa III (fCx = 1.1.3). Nadchatram (1967a) compared N. longipes with Neoschoengastia entomyza Womersley, 1952 and Neoschoengastia thomasi (Radford, 1946). The main difference between N. longipes plus N. thomasi and N. americana is the presence of six setae in first posthumeral row vs. eight. Our material on N. longipes also differs from two specimens of N. americana collected in Paraguay (Stekolnikov et al. 2022) by NDV = 50 – 54 vs. 61 – 62, AP = 34 – 41 vs. 28 – 31, and AL = 59 – 74 vs. 47. The difference by AL and, to a lesser extent, by AP agrees with the morphometric data on N. americana provided by Domrow (1974). However, the mean values of measurements from Malaysian specimens identified as N. americana by Domrow & Nadchatram (1960) differed from ours. In addition, the specimens examined by Domrow & Nadchatram (1960) were characterized by the presence of two additional sternal setae (fSt = 2.4. vs. 2.2). Probably, they belong to a new and still undescribed species. Neoschoengastia entomyza, a species described from Australia, was synonymized with N. americana by Domrow (1974), but we believe that these two taxa require more detailed comparison. Thus, presence of N. americana outside the American continent remains questionable.
Neoschoengastia thomasi was described incompletely, without characters of gnathosoma and legs (Radford 1946). According to the redescription prepared by Nadchatram (1967a) on the base of the type specimen deposited in the British Museum (Natural History), London (currently, Natural History Museum), N. thomasi differs from N. americana and N. longipes by the branched lateral palpal tibial seta (fPp = B/B/BbB vs. B/B/BNB). The redescription was illustrated with a drawing of the scutum lacking all setae except for one AL and one PL (Nadchatram 1967a, fig. 17). Surprisingly, the cuticular striations in the medial part of the scutum were drawn as transverse, whereas they are longitudinal in all above species. As noted by Domrow & Lester (1985), this drawing rather corresponds to Neoschoengastia struthidia Womersley, 1952. Fernandes & Kulkarni (2003) based their redescription of N. thomasi on existing literature. Unfortunately, the size of the N. thomasi type series and the place of deposition of other type specimens is unknown. Therefore, it remains a dubious species.
Among our N. longipes material, three measured specimens (ZIN 17951, 17953, 17954) from Copsychus malabaricus collected 9 – 10 December 2021 differ from five other measured specimens by a lesser NDV (50 vs. 53 – 54) smaller scutum (AW = 46 – 50 vs. 51 – 58; SB = 26 – 28 vs. 29 – 35; SD = 44 – 47 vs. 49 – 56), and shorter legs (Ip = 1197 – 1210 vs 1222 – 1294; TaIIIL = 122 – 128 vs. 131 – 139). We provisionally consider this as a case of intraspecific variability.
Neoschoengastia solitus Nadchatram, 1967
Distribution and hosts — Malaysia, Thailand, ex 11 bird species (Stekolnikov 2021a).
Material examined — Three larvae (ZIN 17955 – 17957) ex one Geokichla citrina, THAILAND, Rayong province, Koh Mun Nai, 12.612384 N, 101.687574 E, 10 March 2021, coll. S. Koosakulnirand.
Neoschoengastia gallinarum (Hatori, 1920)
Distribution and hosts — China, Malaysia, Taiwan, Vietnam, ex 24 bird species and subspecies, and from two species of rats and a hare (Stekolnikov 2021a). Here this species is for the first time reported from Thailand. Argusianus argus, Lophura rufa, and Polyplectron inopinatum are new hosts.
Material examined — Four larvae (ZIN 17958 – 17961) ex one Gallus gallus domesticus, THAILAND, Nan province, Ban Huai Muang village, 19.139995 N, 100.718956 E, 23 December 2021, coll. S. Koosakulnirand; two larvae ex one Argusianus argus and 14 larvae ex five Polyplectron malacense, MALAYSIA, Johor State, Jemaluang Wildlife Conservation Centre, 2.291356 N, 103.852966 E, 22 February 2022, coll. P. Rajasegaran; 11 larvae ex five Gallus gallus domesticus, MALAYSIA, Johor State, Kota Tinggi plantation, 2.030233 N, 103.866035 E, 21 – 23 June 2022, coll. P. Rajasegaran; 24 larvae ex two Lophura rufa (15), three Polyplectron inopinatum (8), and one Polyplectron malacense (1), MALAYSIA, Perak State, Sungkai Wildlife Conservation Centre, 4.064302 N, 101.366232 E, 9 January and 8 March 2021, coll. P. Rajasegaran; nine larvae ex seven Gallus gallus domesticus, MALAYSIA, Selangor State, Bestari Jaya village, 3.378008 N, 101.410224 E, 12 April 2021, coll. P. Rajasegaran.
Genus Schoengastia Oudemans, 1910
Type species — Thrombidium vandersandei Oudemans, 1905.
Diagnosis — SIF = 7BS-N-3(2)-3(2)111.(0 – 2)000; cheliceral blade serrate on its dorsal edge, with tricuspid cap; scutum usually subpentagonal, with convex posterior margin; AL > PL > AM, sensillary bases situated not far apart and close to level of PLs; sensilla globose, covered with setules; eyes 2 + 2.
Schoengastia archaea (Taufflieb, 1960)
(Figure 5D)
Diagnosis — SIF = 7BS-N-3-2111.0000; fPp = B/B/NBB; fCx = 1.1.1; fSt = 2.2; AL ≥ PL >> AM; fD = (49 – 60)-(26 – 30)+(56 – 72); DS = 138 – 159; V = 89 – 100; NDV = 227 – 254; Ip = 1024 – 1127; cheliceral blade with four dorsal hooks; eyes 2 + 2, very large; scutum as long as wide, with prominent anterolateral shoulders, with greatly projected posterior margin concave in center; entire scutum covered with puncta, its half posterior to sensillary bases also covered with distinct transverse striations; sensilla (trichobothria) drop-shaped, covered with setules; sensillary bases posterior to level of PL; leg tarsi with multiple sclerite bars; S1 in distal part of segment, f1 behind S1, f2 behind S2. Standard measurements are given in Table 7.
Distribution and hosts — This species was described from Senegal (Rufisque), ex Sterna hirundo L. (Charadriiformes: Laridae) (Taufflieb 1960). Here it is for the first time recorded in Asia (Thailand) and on all hosts mentioned below.
Material examined — Six larvae (17966 – 17971) ex two Charadrius mongolus (2), one Calidris ruficollis (1), two Calidris tenuirostris (2), and one Xenus cinereus (1), THAILAND, Trang province, Samran Beach, 7.201348 N, 99.562105 E and 7.209399 N, 99.557854 E, 15 – 17 March 2021, 19 March 2022, coll. S. Koosakulnirand; four larvae (ZIN 17972 – 17975) ex three Numenius arquata, THAILAND, Trang province, White Dragon Spine Beach (7.2984973 N, 99.4206218 E), 20 March 2022, coll. S. Koosakulnirand.
12 larvae (UoL) from same localities, same and additional host individuals of the same species.
Remarks — This species is similar to Schoengastia galapa Stekolnikov, 2021 described ex marine iguana Amblyrhynchus cristatus Bell (Squamata: Iguanidae) from Galápagos Islands; this similarity has not been noted before. Both species stand out among Schoengastia due to the numerous idiosomal setae (NDV ca. 200 or more), presence of two genualae I (σ) vs. three or more, presence of anterolateral shoulders of the scutum, and the presence of transverse cuticular striations on the scutum posterior to sensillary bases. Such striations do not reach the level of sensillary bases in all other species of Schoengastia (Stekolnikov 2021b). Schoengastia galapa differs from S. archaea by fPp = N(b)/N/NNB vs. B/B/NNB, much shorter legs (Ip = 733 – 795 vs. 1019 – 1127) without additional sclerite bars, presence of mastitarsalae, f1 distal to S1 vs. proximal, smaller anterior pair of eyes, fewer idiosomal setae (NDV = 197 – 206 vs. 227 – 256), and by a wider scutum (AW = 46 – 48 vs. 34 – 38, PW = 62 – 66 vs. 51 – 56).
Genus Ascoschoengastia Ewing, 1946
Ascoschoengastia lorius (Gunther, 1939)
Distribution and hosts — Australia, China, Papua New Guinea, Thailand, Vietnam, ex nine mammal hosts and four bird species (Stekolnikov 2021a). Here this species is for the first time recorded in Malaysia and on Meiglyptes tukki.
Material examined — 12 larvae (UM) ex one Meiglyptes tukki, MALAYSIA, Pahang State, Krau Wildlife Reserve, 3.596982 N, 102.183190 E, 17 February 2021, coll. P. Rajasegaran.
Genus Blankaartia Oudemans, 1911
Blankaartia acuscutellaris (Walch, 1922)
Distribution and hosts — Eastern Europe, Spain, Tropical Africa, Central, South, Southeast and East Asia, mainly on water birds, also on mammals of different orders, including humans, and two records on chameleons (Stekolnikov 2021a). Lewinia striata is a new host species.
Material examined — One larva (ZIN 18170) ex Lewinia striata, MALAYSIA, Sarawak State, Sematan Pueh village, 1.831126 N, 109.708966 E, 22 March 2022, coll. P. Rajasegaran.
One larva (UM) with same data.
Genus Ericotrombidium Vercammen-Grandjean, 1966
Ericotrombidium cosmetopode (Vercammen-Grandjean & Langston, 1971)
(Figure 6)
Diagnosis — SIF = 7BS-B-3-2111.0000; fPp = B/B/NNB; fCx = 1.1.1; fSt = 2.2; PL > AL > AM (PL > AM ≥ AL); fD = 2H-8-6-6-4-2 (8-6-6-4-4-2); DS = 28 – 32; V = 27 – 28; NDV = 55 – 60; Ip = 666 – 770; dorsal idiosomal setae tetrapectinate, with thin barbs; scutum with large puncta, with bilobate posterior margin; SB at level of PL; sensilla with cilia in medial part and ca. 13 branches in distal third; f1 distal to S1, f2 distal to S2; S2 thin, pointed. Standard measurements are given in Table 8.
Distribution and hosts — This species was described from free larvae found in a ground hole, Malaysia, Selangor State, Ulu Langat Forest Reserve. Pellorneum ruficeps is the first known host of this species.
Material examined — One larva (ZIN 17937) ex Pellorneum ruficeps, MALAYSIA, Pahang State, Fraser’s Hill, 3.725630 N, 101.716365 E, 22 October 2021, coll. P. Rajasegaran.
Remarks — As compared with the original description, our specimen has longer legs and a slightly reduced number of idiosomal setae (Table 8).
Genus Eutrombicula Ewing, 1938
Eutrombicula wichmanni (Oudemans, 1905)
Distribution and hosts — Australia, Myanmar, Brunei, China, Guam, Indonesia, Japan, Malaysia, Papua New Guinea, Philippines, Taiwan, Thailand, Vietnam, ex many species of mammals from different orders (including humans), reptiles, and birds (Stekolnikov 2021a).
Material examined — Three larvae (ZIN 17997 – 17999) collected by the black plate technique, THAILAND, Nan province, Ban Huai Muang village, 19.140834 N, 100.718315 E, 26 December 2021, coll. Kamonchanok Bunmee; 19.1391337 N, 100.7195905 E, 25 December 2021, coll. Rawadee Kumlert; 19.1378833 N, 100.7193387 E, 27 December 2021, coll. Kamonchanok Bunmee; two larvae (ZIN 18000, 18001) ex two Gallus gallus domesticus, same place, 19.139652 N, 100.718019 E and 19.140026 N, 100.719794 E, 23 December 2021, coll. S. Koosakulnirand.
A great many (113) larvae (UoL) ex Gallus gallus domesticus, THAILAND, Nan province, Ban Huai Muang village, different collection sites, 23 – 24 December 2021, coll. S. Koosakulnirand; 2 larvae (UoL) ex Gallus gallus domesticus, THAILAND, Nan province, Ban Santisuk village, 19.131006 N, 100.698332 E, 24 December 2021, coll. S. Koosakulnirand.
Genus Helenicula Audy, 1954
Helenicula comata (Womersley, 1952)
Distribution and hosts —China, India, Philippines, ex Rattus tanezumi (Stekolnikov 2021a). Here this species is for the first time recorded in Thailand and on Gallus gallus domesticus.
Material examined — One larva (ZIN 17989) collected by the black plate technique, THAILAND, Nan province, Ban Huai Muang village, 19.1392019 N, 100.7199339 E, 25 December 2021, coll. Rawadee Kumlert; three larvae (ZIN 17990 – 17992) collected by the black plate technique, same place, 19.1386161 N, 100.7162079 E, 26 December 2021, coll. Kamonchanok Bunmee.
Three larvae (UoL) ex Gallus gallus domesticus, THAILAND, Nan province, Ban Huai Muang village, 19.139696 N, 100.720205 E, 24 December 2021, coll. S. Koosakulnirand.
Genus Leptotrombidium Nagayo, Miyagawa, Mitamura & Imamura, 1916
Leptotrombidium deliense (Walch, 1922)
Distribution and hosts — Australia, Cambodia, China, India, Indonesia, Laos, Malaysia, Maldive Islands, Myanmar, Nepal, Papua New Guinea, Philippines, Sri Lanka, Taiwan, Thailand, Vietnam, ex a wide range of mammal hosts (including humans), birds, occasionally on reptiles and arthropods (Stekolnikov 2021a).
Material examined — One larva (ZIN 17936) ex Pellorneum ruficeps, MALAYSIA, Pahang State, Fraser’s Hill, 3.725630 N, 101.716365 E, 22 October 2021, coll. P. Rajasegaran; one larva (ZIN 18171) ex P. ruficeps, MALAYSIA, Kedah State, Langkawi Isl., Gunung Machinchang, 6.429010 N, 99.729852 E, 16 March 2022, coll. P. Rajasegaran.
Leptotrombidium imphalum Vercammen-Grandjean & Langston, 1976
Distribution and hosts — Sri Lanka, China, India, Malaysia, Myanmar, Pakistan, Taiwan, Thailand, ex 19 species of mammals from four orders and ex one unidentified bird (Stekolnikov 2021a). Amaurornis phoenicurus and Lewinia striata are new host species.
Material examined — Two larvae (ZIN 17933, 17934) ex Amaurornis phoenicurus and Lewinia striata, MALAYSIA, Sarawak State, Sematan Pueh village, 1.831126 N, 109.708966 E, 22 March 2022, coll. P. Rajasegaran.
Leptotrombidium miculum (Traub & Audy, 1954)
Distribution and hosts — Malaysia, Thailand, ex five species of rodents (Stekolnikov 2021a). Here this species is for the first time recorded on a bird host.
Material examined — One larva (ZIN 17932) ex Pellorneum ruficeps, MALAYSIA, Kedah State, Langkawi Isl., Gunung Machinchang, 6.429010 N, 99.729852 E, 17 March 2022, coll. P. Rajasegaran.
Genus Odontacarus Ewing, 1929
Odontacarus audyi (Radford, 1946)
Distribution and hosts — India, Malaysia, Thailand, Vietnam, on 22 bird species (Stekolnikov 2021a). Cacomantis sepulcralis, Cyornis tickelliae, Eumyias thalassinus, Geokichla citrina, Geokichla sibirica, Larvivora cyane, Philentoma pyrhoptera, Cyornis brunneatus, Stachyris nigriceps, and Stachyris poliocephala are new host species.
Material examined — Three larvae (ZIN 17938 – 17940) ex Geokichla citrina, Geokichla sibirica and Larvivora cyane, THAILAND, Rayong province, Koh Mun Nai, 12.612384 N, 101.687574 E, 10 – 11 March 2021, coll. S. Koosakulnirand; two larvae (ZIN 17941, 17943) ex L. cyane and Cyornis tickelliae, THAILAND, Kanchanaburi province, Mahidol University Campus, 14.1288777 N, 99.1611407 E, 8 – 9 December 2021, coll. S. Koosakulnirand; one larva (ZIN 17942) ex L. cyane, THAILAND, Kanchanaburi province, SW Mahidol University Campus, 14.11806 N, 99.15253 E, 9 December 2021, coll. S. Koosakulnirand; two larvae (ZIN 17944, 17945) ex L. cyane and Copsychus malabaricus, THAILAND, Kanchanaburi province, W Mahidol University Campus, 14.13111 N, 99.14863 E, 10 December 2021, coll. S. Koosakulnirand; one larva (ZIN 18167) ex Gallus gallus domesticus, THAILAND, Nan province, Ban Santisuk village, 19.131006 N, 100.698332 E, 23 December 2021, coll. S. Koosakulnirand.
Fifty-three larvae (UoL) from same localities of Kanchanaburi province, same and additional host individuals of the same species, plus 11 larvae ex one Copsychus saularus (2), one Mixornis gularis (1), two Pycnonotus conradi (4), and two Rhipidura javanica (4), THAILAND, Kanchanaburi province, Mahidol University Campus, 14.1288777 N, 99.1611407 E, 9 December 2021, coll. S. Koosakulnirand; nine larvae (UoL) ex one Mixornis gularis (2), two Pycnonotus conradi (4), and one Rubigula flaviventris (3), THAILAND, Kanchanaburi province, W Mahidol University Campus, 14.13111 N, 99.14863 E, 10 December 2021, coll. S. Koosakulnirand; 53 larvae (UoL) ex three Copsychus malabaricus (7), four Cyornis tickelliae (15), four Larvivora cyane (15), one Pycnonotus aurigaster (2), four Pycnonotus conradi (8), and two Pycnonotus finlaysoni (6), THAILAND, Kanchanaburi province, W Mahidol University Campus, 14.134225 N, 99.1374365 E, 11 December 2021, coll. S. Koosakulnirand; nine larvae (UoL) ex one Dicrurus paradiseus (1), one Larvivora cyane (3), one Calliope calliope (3), one Ficedula albicula (1), and one Pycnonotus conradi (1), THAILAND, Kanchanaburi province, NW Mahidol University Campus, 14.154772 N, 99.1193296 E, 12 – 13 December 2021, coll. S. Koosakulnirand.
Three larvae (UM) ex Pellorneum ruficeps, Geokichla citrina, and Larvivora cyane, MALAYSIA, Kedah State, Langkawi Isl., Gunung Machinchang, 6.429010 N, 99.729852 E, 16 – 17 March 2022, coll. P. Rajasegaran; one larva (UM) ex P. ruficeps, MALAYSIA, Pahang State, Fraser’s Hill, 3.725630 N, 101.716365 E, 22 October 2021, coll. P. Rajasegaran; five larvae (UM) ex Cacomantis sepulcralis, Copsychus malabaricus, Eumyias thalassinus, Stachyris nigriceps, and Stachyris poliocephala, MALAYSIA, Selangor State, Ulu Gombak Forest Reserve, 3.325987 N, 101.752747 E, 9 – 12 February 2021, coll. P. Rajasegaran; 16 larvae (UM) ex one C. malabaricus (1), four L. cyane (5), four Philentoma pyrhoptera (4), one Cyornis brunneatus (2), and four S. poliocephala (4), MALAYSIA, Terengganu State, Pasir Raja Forest Reserve, 4.790517 N, 102.996835 E, 23 – 27 October 2021, coll. P. Rajasegaran.
Genus Parascoschoengastia Vercammen-Grandjean, 1960
Parascoschoengastia heynemani (Nadchatram & Upham, 1966)
Distribution and hosts — China and Malaysia, ex Halcyon pileata (Boddaert), Alcedo atthis bengalensis J.F. Gmelin, Alcedo peninsulae, and Alcedo meninting Horsfield (Coraciiformes: Alcedinidae) (Stekolnikov 2021a). Actenoides concretus is a new host species.
Material examined — Four larvae (UM) ex one Actenoides concretus, MALAYSIA, Pahang State, Krau Wildlife Reserve, 3.596982 N, 102.183190 E, 18 February 2021, coll. P. Rajasegaran; four larvae (UM) ex one Alcedo peninsulae, MALAYSIA, Sarawak State, Gunung Gading National Park, Lundu, 1.692098 N, 109.845157 E, 22 March 2022, coll. P. Rajasegaran.
Characteristics of infestation
Thailand. In total, 754 hosts were examined; 199 (26.4%) were found parasitized by chiggers (2,340 specimens); 440 chigger specimens from 140 hosts were slide-mounted and identified (Table 1). There were 30 hosts (22%) bearing more than one chigger species, considering only those cases confirmed by the identification of slide-mounted chiggers. One individual of Geokichla citrina from Koh Mun Nai was parasitized by three species (N. solitus, O. audyi, and T. densipiliata). The pairs of co-occurring species were N. longipes and O. audyi (19 cases), N. longipes and T. kirhocephales (3), N. (W.) ... and S. archaea (3), O. audyi and T. kirhocephales (2), O. audyi and T. densipiliata (1), and E. wichmanni and H. comata (1).
Malaysia. In total 777 hosts were examined; 129 (16.6%) were found parasitized by chiggers; 243 chigger specimens from 124 hosts were slide-mounted and identified (Table 2). One Pellorneum ruficeps from Fraser’s Hill bore three species (E. cosmetopode, L. deliense, and O. audyi); whereas four hosts bore pairs of species—O. audyi and T. densipiliata (2), L. miculum and O. audyi (1), and L. imphalum and B. acuscutellaris (1).

4. Discussion

The specificity of chigger mites for different orders of birds is a trait that can be noted first of all when considering our results. Thus, our materials from Thailand demonstrate three clearly different sets of chigger species connected with birds from different orders occurring in different habitats (Table 9). Shorebirds (order Charadriiformes) were parasitized by four species of Neacariscus and by Schoengastia archaea. Birds inhabiting forests (mainly Passeriformes) bore two species of Neoschoengastia (N. longipes and N. solitus), two Toritrombicula spp., and Odontacarus audyi. Domestic chicken (Galliformes) were parasitized by Eutrombicula wichmanni, Neoschoengastia gallinarum, Helenicula comata, and O. audyi. Thus, only the latter species was found on birds belonging to two of these ecological (and taxonomic) groups.
In the materials from Malaysia (Table 10), we can see that Galliformes, represented in this collection by four species of pheasants in addition to domestic chickens (Table 2), were the hosts of a sole chigger species, Neoschoengastia gallinarum. The chigger species composition on Passeriformes was similar to that of Thailand, and included two species of Toritrombicula and O. audyi. Other recorded species were rare. Finally, Parascoschoengastia heynemani was found only on two species of kingfisher (Coraciiformes: Alcedinidae), in two localities from different provinces. Noteworthy is that previous records of this species were on hosts from the same family (Stekolnikov 2021a).
Since chigger mites are temporary parasites, their distribution is expected to be determined by the natural conditions of the biotopes suitable both for their free-living postlarval stages and for their hosts. However, the size of actually recorded host ranges for different chigger species can be highly variable and dependent, theoretically, on the number of host species in some localities and on the modes of chigger activity; for example, preferred habitats where they lie in wait for their hosts (ground surface, grass, tree branches, etc.). These aspects of their behavior are poorly known: however, the connection of P. heynemani with kingfishers suggests this chigger species is nidicolous, occupying the burrows in which kingfishers nest. Similarly, A. lorius larvae and post-larval stages were found together in tree-hollow nests of psittaform birds in Queensland, Australia, indicating that the entire lifecycle of this species can be nidicolous (Shaw et al. 2010). Such ecological habits could explain the infestation of the buff-necked woodpecker (M. tukki) by A. lorius in Malaysia, as this host species also nests in tree hollows.
The separate set of chigger species from the shorebirds in Thailand was the most unexpected discovery during our investigation. Among the five chigger species collected from the birds inhabiting the seashore of southern Thailand, Schoengastia archaea and Neacariscus sulae were previously known only from their type localities in West Africa. Neacariscus pluvius was previously recorded only in Oceania, and N. shiraii from Oceania and Japan. One species of this genus was described above as new.
According to the literature, species of the genus Neacariscus parasitize mainly aquatic birds (Vercammen-Grandjean & Langston 1976; Mertins 2009). One species, N. thompsoni, in addition to the birds, was found on a fish-eating bat. Only two species, N. chaetosa and N. nativitatis, are known from lizards in Central and South America; while N. shiraii was occasionally collected on rats. However, where the collection sites of these species are known, they tend to be located in coastal areas accessible to seabirds (Mertins 2009; Stekolnikov & González-Acuña 2015). Schoengastia archaea was also described from a species of Charadriiformes. We assume that the fauna of chiggers inhabiting seashores and parasitizing the birds associated with these biotopes is mostly uniform throughout the tropical zone of the world. Probably, Schoengastia galapa, described from the Galápagos Islands belongs to the same species complex, although it was collected only from a marine iguana (Stekolnikov 2021b). Its presence on seabirds seems probable.
Migrations of birds could facilitate widespread dispersal of the bird chiggers, such as S. archaea and N. sulae. Varma (1964) discussed this possibility in detail. In part, the wide range of Blankaartia acuscutellaris, which extends from Southeast Asia to Western Europe and Africa, could be connected with the fact that principal hosts of this species are birds associated with wetlands (although it has been frequently collected from other hosts, including humans) (Mąkol & Korniluk 2017; Trnka et al. 2022). Recently, a disjunctive area of distribution, consisting of three isolated parts—Eastern Europe, Iran, and Vietnam—was established for Neotrombicula elegans (Shamsi et al. 2020). Since in Vietnam this species was found on three species of birds, including Larvivora sibilans (Swinhoe), which was noted as a rare vagrant to Europe (Clement & Rose 2015), these authors hypothesized that dissemination by birds could be the cause of the scattered range of N. elegans.
We add that alternative migration routes within the same population of birds can result in the presence of greatly disjunct wintering sites, both latitudinally and longitudinally. Thus, the wintering sites of the little ringed plovers Charadrius dubius Scopoli (Charadriiformes: Charadriidae) breeding in south Sweden were identified as far apart as West Africa and India using geolocators (Hedenström et al. 2013). Similarly, Terek sandpiper (X. cinereus), which we found to be parasitized with three Neacariscus spp. and S. archaea in Thailand, has a vast breeding range across the taiga from Finland to eastern Siberia. Its wintering grounds extend from eastern and southern Africa, the Middle East, southern Asia, and Australia, with vagrants appearing as far afield as the Americas (White et al. 2006). Provided that the breeding sites of some bird species are suitable to support the life cycle of chiggers (at least for part of the year), variability of migration routes could thus result in extensive meridional dissemination of chiggers over vast distances.
The chigger species composition on the Passeriformes in the study areas significantly differed from that of the seven passerine bird species examined by Kalúz et al. (2016) in north Vietnam. Among the 12 chigger species found by these authors, only one (O. audyi) was also recorded in our study. Moreover, they found no species of Toritrombicula, whereas we recorded three, and this genus was rather frequent in our collections (Tables 9, 10). In Thailand, we collected two species of Neoschoengastia from passerines whereas three other species from the same genus were reported by Kalúz et al. (2016). In contrast, we did not find this genus on Passeriformes in the Malaysian localities. Finally, we collected three species of Leptotrombidium from Passeriformes in Malaysia, but none in Thailand, while Kalúz et al. (2016) found six other Leptotrombidium species. Evidently, the fauna of bird chiggers in forest biotopes of Southeast Asia is highly variable and requires further investigations, including quantitative analysis of host-chigger networks using the data presented here, which will be the subject of a separate publication.

Acknowledgements

We thank Dr Nikita Chernetsov (Zoological Institute, Saint Petersburg, Russia) for useful information on bird migrations, and Dr Somying Thunhikorn and Dr Kirana Noradechanon of the DNP for facilitating access to national parks in Thailand. Elsewhere in Thailand we also acknowledge the assistance of Mr Jirut Khamaye and the field research team during sampling in the Bueng Boraphet Non-Hunting Area and Mr Suthep Jualaong, Director, Eastern Marine and Coastal Resources Center (EMCOR), Department of Marine and Coastal Resources, on Ko Mun Nai. In Malaysia, we were very grateful for the support of the field team from the DWNP (Ismail Hj. Mamat, Abdul Rahman bin Ahmad, Mohd Hairol Mat Zin, Adzri bin Azmi, Stewart Angin, and Suhaimi bin Mansor); research assistants Nuramirah Diyanah binti Mohd Johan and Siti Nurul Izzah binti Mohd Azami from Universiti Malaya; and Dr Mohammad Saiful bin Mansor’s field team from Universiti Kebangsaan Malaysia. This research was supported by a Royal Society International Collaboration Award (ICA\R1\191058) awarded to BLM and SA; a Mahidol-Liverpool PhD scholarship awarded to SK; the Higher Institution Centre of Excellence (HICoE), Universiti Malaya; and the Ministry of Science and Higher Education of the Russian Federation (122031100263-1, to AAS).

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Figure 1. Neacariscus (Neacariscus) pluvius (Wharton, 1945), specimen ZIN 17965: A – anterior dorsal aspect of idiosoma. Neacariscus (Whartonacarus) ... Stekolnikov, n. sp., holotype: B – anterior dorsal aspect of idiosoma; C – sternal area. Neacariscus (Whartonacarus) sulae (Oudemans, 1910), specimen ZIN 18168: D – anterior dorsal aspect of idiosoma. Scale bars: 50 μm.
Figure 2. Neacariscus (Whartonacarus) ... Stekolnikov, n. sp.: A – additional specimen ZIN 17976, dorsal aspect of idiosoma; B – additional specimen ZIN 17976, arrangement of ventral idiosomal setae; C – holotype, humeral seta; D – holotype, central seta of 2nd posthumeral row; E – holotype, preanal seta. Neacariscus (Whartonacarus) shiraii (Sasa, Kano & Obata, 1952), specimen ZIN 17981: F – dorsal aspect of idiosoma; G – arrangement of ventral idiosomal setae. Abbreviations: H – humeral seta; u – uropore (anus); V – ventral idiosomal setae. Scale bars: A, B, F, G – 100 μm; C – E – 50 μm.
Figure 3. Neacariscus (Whartonacarus) ... Stekolnikov, n. sp.: A holotype, scutum and eyes; C – additional specimen ZIN 17976, cheliceral blade; D – paratype ZIN 17984, ventral aspect of idiosoma; E – paratype ZIN 17984, dorsal aspect of idiosoma. Neacariscus (Whartonacarus) shiraii (Sasa, Kano & Obata, 1952), specimen ZIN 17981: B – scutum and eyes. Abbreviations: AL – anterolateral scutal seta; AM – anteromedian scutal seta; Cb – cheliceral base; Cs – cheliceral blade; Ga – galeal (deutorostral) seta; gns – gnathocoxal (tritorostral) seta; ic – infracapitulum (gnathobase, gnathocoxa); PC – palpal claw (odontus); PF – palpal femur; PG – palpal genu; PL – posterolateral scutal seta; PTa – palpal tarsus; PTi – palpal tibia; S – sensillum (trichobothrium); S0 – palpal tarsala (ω); ζ – palpal subterminala (ζ). Scale bars: A, B – 50 µm; C – E – 20 µm.
Figure 4. Neacariscus (Whartonacarus) ... Stekolnikov, n. sp., holotype: A – leg I tarsus; B – leg I genu and tibia; C – leg I trochanter, basifemur, and telofemur; D – leg II tarsus; E – leg II genu and tibia; F – leg II trochanter, basifemur, and telofemur; G – leg III tarsus; H – leg III genu and tibia; I – leg III trochanter, basifemur, and telofemur. Abbreviations: f1 – microtarsala I (ε); f2 – microtarsala II (ε); ga – genuala I (σ); gm – genuala II (σ); gp – genuala III (σ); Mt – mastitarsala; pST – parasubterminala (z); PT' – pretarsala I (ζ); PT" – pretarsala II (ζ); S1 – leg tarsala I (ω); S2 – leg tarsala II (ω); ST – subterminala (ζ); ta – tibiala I (φ); tm – tibiala II (φ); tp – tibiala III (φ); µga – microgenuala (κ); µta – microtibiala (κ). Scale bar: 50 μm.
Figure 5. Toritrombicula kirhocephales Goff, 1982: A – specimen ZIN 18009, anterior dorsal aspect of idiosoma. Toritrombicula uphami Nadchatram, 1967: B – specimen ZIN 18169, anterior dorsal aspect of idiosoma. Neoschoengastia longipes Nadchatram, 1967: C – specimen ZIN 17947, dorsal aspect of idiosoma. Schoengastia archaea (Taufflieb, 1960): D – specimen ZIN 17967, dorsal aspect of idiosoma.
Figure 6. Ericotrombidium cosmetopode (Vercammen-Grandjean & Langston, 1971), specimen ZIN 17937: A –anterior dorsal aspect of idiosoma; B – scutum. Scale bars: 50 μm.
Table 1. Material from Thailand. Mahidol Univ. – collection sites NW, W, and SW of Mahidol University campus; Albino Dragon – Albino Dragon Spine beach.
Table 1. Material from Thailand. Mahidol Univ. – collection sites NW, W, and SW of Mahidol University campus; Albino Dragon – Albino Dragon Spine beach.
Province Locality Habitat Month Host species Host order Host family Hosts Slides
Kanchanaburi Mahidol Univ. Highland December Dicrurus paradiseus (L.) Passeriformes Dicruridae 1 1
Kanchanaburi Mahidol Univ. Highland December Calliope calliope (Pallas) Passeriformes Muscicapidae 2 7
Kanchanaburi Mahidol Univ. Highland December Copsychus malabaricus (Scopoli) Passeriformes Muscicapidae 15 61
Kanchanaburi Mahidol Univ. Highland December Copsychus saularis (L.) Passeriformes Muscicapidae 1 4
Kanchanaburi Mahidol Univ. Highland December Cyornis tickelliae Blyth Passeriformes Muscicapidae 11 42
Kanchanaburi Mahidol Univ. Highland December Ficedula albicilla (Pallas) Passeriformes Muscicapidae 2 2
Kanchanaburi Mahidol Univ. Highland December Larvivora cyane (Pallas) Passeriformes Muscicapidae 14 64
Kanchanaburi Mahidol Univ. Highland December Pycnonotus aurigaster (Vieillot) Passeriformes Pycnonotidae 2 3
Kanchanaburi Mahidol Univ. Highland December Pycnonotus conradi (Finsch) Passeriformes Pycnonotidae 11 21
Kanchanaburi Mahidol Univ. Highland December Pycnonotus finlaysoni Strickland Passeriformes Pycnonotidae 2 6
Kanchanaburi Mahidol Univ. Highland December Rubigula flaviventris (Tickell) Passeriformes Pycnonotidae 1 3
Kanchanaburi Mahidol Univ. Highland December Rhipidura javanica (Sparrman) Passeriformes Rhipiduridae 2 4
Kanchanaburi Mahidol Univ. Highland December Mixornis gularis (Horsfield) Passeriformes Timaliidae 2 3
Kanchanaburi Mahidol Univ. Highland December Otus sunia (Hodgson) Strigiformes Strigidae 1 2
Nan Huai Muang Village December Gallus gallus domesticus (L.) Galliformes Phasianidae 30 122
Nan Ban Santisuk Village December Gallus gallus domesticus Galliformes Phasianidae 2 3
Rayong Koh Mun Nai Island March Alcedo atthis (L.) Coraciiformes Alcedinidae 1 7
Rayong Koh Mun Nai Island March Larvivora cyane Passeriformes Muscicapidae 2 8
Rayong Koh Mun Nai Island March Pachycephala cinerea (Blyth) Passeriformes Pachycephalidae 3 12
Rayong Koh Mun Nai Island March Phylloscopus tenellipes Swinhoe Passeriformes Phylloscopidae 1 2
Rayong Koh Mun Nai Island March Pycnonotus conradi Passeriformes Pycnonotidae 2 2
Rayong Koh Mun Nai Island March Geokichla citrina (Latham) Passeriformes Turdidae 1 5
Rayong Koh Mun Nai Island March Geokichla sibirica sibirica (Pallas) Passeriformes Turdidae 1 3
Satun Thung Sabo Shore March Charadrius leschenaultii Lesson Charadriiformes Charadriidae 1 1
Satun Thung Sabo Shore March Calidris tenuirostris (Horsfield) Charadriiformes Scolopacidae 3 6
Trang Albino Dragon Shore March Numenius arquata (L.) Charadriiformes Scolopacidae 3 8
Trang Koh Libong Shore March Pluvialis squatarola (L.) Charadriiformes Charadriidae 1 2
Trang Koh Libong Shore March Calidris tenuirostris Charadriiformes Scolopacidae 2 4
Trang Samran beach Shore March Charadrius leschenaultii Charadriiformes Charadriidae 2 2
Trang Samran beach Shore March Charadrius mongolus Pallas Charadriiformes Charadriidae 2 4
Trang Samran beach Shore March Calidris ruficollis (Pallas) Charadriiformes Scolopacidae 2 3
Trang Samran beach Shore March Calidris tenuirostris Charadriiformes Scolopacidae 8 16
Trang Samran beach Shore March Xenus cinereus (Güldenstädt) Charadriiformes Scolopacidae 6 7
Table 2. Material from Malaysia. Behrang – Behrang Forest Reserve; Bestari Jaya – Bestari Jaya village; Gunung Gading – Gunung Gading National Park, Lundu; Jemaluang – Jemaluang Wildlife Conservation Centre; Kota Tinggi – Kota Tinggi plantation; Krau – Krau Wildlife Reserve; Machinchang – Gunung Machinchang, Langkawi Isl.; Pasir Raja – Pasir Raja Forest Reserve; Pueh – Pueh village, Sematan; Sungkai – Sungkai Wildlife Conservation Centre; Ulu Gombak – Ulu Gombak Forest Reserve.
Table 2. Material from Malaysia. Behrang – Behrang Forest Reserve; Bestari Jaya – Bestari Jaya village; Gunung Gading – Gunung Gading National Park, Lundu; Jemaluang – Jemaluang Wildlife Conservation Centre; Kota Tinggi – Kota Tinggi plantation; Krau – Krau Wildlife Reserve; Machinchang – Gunung Machinchang, Langkawi Isl.; Pasir Raja – Pasir Raja Forest Reserve; Pueh – Pueh village, Sematan; Sungkai – Sungkai Wildlife Conservation Centre; Ulu Gombak – Ulu Gombak Forest Reserve.
Province Locality Month Host species Host order Host family Hosts Slides
Johor Jemaluang February Argusianus argus (L.) Galliformes Phasianidae 1 2
Johor Jemaluang February Polyplectron malacense (Scopoli) Galliformes Phasianidae 5 14
Johor Kota Tinggi June Gallus gallus domesticus Galliformes Phasianidae 5 11
Kedah Machincang March Cyornis tickelliae Blyth Passeriformes Muscicapidae 2 2
Kedah Machincang March Larvivora cyane (Pallas) Passeriformes Muscicapidae 3 3
Kedah Machincang March Malacocincla abbotti Blyth Passeriformes Pellorneidae 1 1
Kedah Machincang March Pellorneum ruficeps Swainson Passeriformes Pellorneidae 3 4
Kedah Machincang March Geokichla citrina (Latham) Passeriformes Turdidae 1 1
Pahang Fraser’s Hill October Pellorneum ruficeps Passeriformes Pellorneidae 1 3
Pahang Krau February Actenoides concretus (Temminck) Coraciiformes Alcedinidae 1 4
Pahang Krau February Calyptomena viridis Raffles Passeriformes Calyptomenidae 1 1
Pahang Krau February Terpsiphone atrocaudata (Eyton) Passeriformes Monarchidae 1 1
Pahang Krau February Ficedula dumetoria (Wallace) Passeriformes Muscicapidae 1 3
Pahang Krau February Malacocincla sepiaria (Horsfield) Passeriformes Pellorneidae 1 1
Pahang Krau February Malacopteron cinereum Eyton Passeriformes Pellorneidae 1 2
Pahang Krau February Malacopteron magnum Eyton Passeriformes Pellorneidae 1 1
Pahang Krau February Pellorneum nigrocapitatum (Eyton) Passeriformes Pellorneidae 1 1
Pahang Krau February Pellorneum malaccense (Hartlaub) Passeriformes Pellorneidae 1 2
Pahang Krau February Turdinus macrodactylus (Strickland) Passeriformes Pellorneidae 1 1
Pahang Krau February Cyanoderma erythropterum (Blyth) Passeriformes Timaliidae 1 1
Pahang Krau February Macronus ptilosus Jardine & Selby Passeriformes Timaliidae 1 2
Pahang Krau February Stachyris maculata (Temminck) Passeriformes Timaliidae 2 2
Pahang Krau February Stachyris poliocephala (Temminck) Passeriformes Timaliidae 3 12
Pahang Krau February Philentoma pyrhoptera (Temminck) Passeriformes Vangidae 5 11
Pahang Krau February Meiglyptes tukki (R.P. Lesson) Piciformes Picidae 1 12
Perak Behrang September Actenoides concretus Coraciiformes Alcedinidae 2 4
Perak Behrang September Lanius tigrinus Drapiez Passeriformes Laniidae 1 1
Perak Behrang September Copsychus malabaricus (Scopoli) Passeriformes Muscicapidae 5 11
Perak Behrang September Cyornis brunneatus (H.H. Slater) Passeriformes Muscicapidae 2 6
Perak Behrang September Larvivora cyane Passeriformes Muscicapidae 2 2
Perak Behrang September Arachnothera longirostra (Latham) Passeriformes Nectariniidae 1 1
Perak Behrang September Philentoma pyrhoptera Passeriformes Vangidae 3 4
Perak Behrang September Batrachostomus stellatus (Gould) Podargiformes Podargidae 1 1
Perak Larut Hill March Anthipes solitaris (S. Müller) Passeriformes Muscicapidae 1 1
Perak Larut Hill March Turdus obscurus J.F. Gmelin Passeriformes Turdidae 1 1
Perak Sungkai Jan., March Lophura rufa (Raffles) Galliformes Phasianidae 2 15
Perak Sungkai January Polyplectron inopinatum (Rothschild) Galliformes Phasianidae 3 8
Perak Sungkai January Polyplectron malacense Galliformes Phasianidae 1 1
Sarawak Gunung Gading March Alcedo peninsulae Laubmann Coraciiformes Alcedinidae 1 4
Sarawak Gunung Gading March Calyptomena viridis Passeriformes Calyptomenidae 1 1
Sarawak Gunung Gading March Copsychus malabaricus Passeriformes Muscicapidae 1 1
Sarawak Gunung Gading March Cyanoptila cyanomelana (Temminck) Passeriformes Muscicapidae 1 1
Sarawak Gunung Gading March Philentoma pyrhoptera Passeriformes Vangidae 1 1
Sarawak Pueh March Amaurornis phoenicurus (Pennant) Gruiformes Rallidae 1 1
Sarawak Pueh March Lewinia striata (L.) Gruiformes Rallidae 1 3
Sarawak Pueh March Mixornis bornensis Bonaparte Passeriformes Timaliidae 1 2
Selangor Bestari Jaya April Gallus gallus domesticus Galliformes Phasianidae 7 9
Selangor Ulu Gombak February Cacomantis sepulcralis (S. Müller) Cuculiformes Cuculidae 2 2
Selangor Ulu Gombak February Copsychus malabaricus Passeriformes Muscicapidae 1 1
Selangor Ulu Gombak February Eumyias thalassinus (Swainson) Passeriformes Muscicapidae 1 1
Selangor Ulu Gombak February Tricholestes criniger (Blyth) Passeriformes Pycnonotidae 1 2
Selangor Ulu Gombak February Stachyris nigriceps Blyth Passeriformes Timaliidae 1 1
Selangor Ulu Gombak February Stachyris poliocephala Passeriformes Timaliidae 1 1
Terengganu Pasir Raja October Copsychus malabaricus Passeriformes Muscicapidae 1 1
Terengganu Pasir Raja October Cyornis brunneatus Passeriformes Muscicapidae 1 2
Terengganu Pasir Raja October Enicurus ruficapillus Temminck Passeriformes Muscicapidae 1 1
Terengganu Pasir Raja October Larvivora cyane Passeriformes Muscicapidae 7 9
Terengganu Pasir Raja October Pellorneum nigrocapitatum Passeriformes Pellorneidae 1 1
Terengganu Pasir Raja October Stachyris poliocephala Passeriformes Timaliidae 5 5
Terengganu Pasir Raja October Philentoma pyrhoptera Passeriformes Vangidae 5 6
Terengganu Redang Isl. October Copsychus malabaricus Passeriformes Muscicapidae 11 33
Table 3. Morphometric (AW–dmt, μm) and meristic (DS–NDV) traits of Neacariscus (Whartonacarus) shiraii (Sasa, Kano & Obata, 1952) and Neacariscus (Whartonacarus) ... Stekolnikov, n. sp. Abbreviations: AW – distance between anterolateral scutal setae; PW – distance between posterolateral scutal setae; SB – distance between sensillary (trichobothrial) bases; ASB – distance from the level of sensillary bases to extreme anterior margin of scutum; PSB – distance from the level of sensillary bases to extreme posterior margin of scutum; SD – length of scutum (ASB + PSB); P-PL – distance from the level of posterolateral scutal setae (PL) to extreme posterior margin of scutum; AP – distance between antero- and posterolateral scutal seta on one side; AM – length of anteromedian scutal seta; AL – length of anterolateral scutal setae; PL – length of posterolateral scutal setae; S – length of sensilla (trichobothria); H – length of humeral setae; Dmin – length of the shortest dorsal idiosomal seta; Dmax – length of the longest dorsal idiosomal seta; Vmin – length of the shortest ventral idiosomal seta; Vmax – length of the longest ventral idiosomal seta; pa – length of leg I (including coxa, excluding claws); pm – length of leg II (including coxa, excluding claws); pp – length of leg III (including coxa, excluding claws); Ip – sum of leg lengths (pa + pm + pp); TaIIIL – length of leg tarsus III; TaIIIW – width of leg tarsus III; dmt – distance between the base of leg III tarsus and base of mastitarsala; DS – number of dorsal idiosomal setae (excluding scutal); V – number of ventral idiosomal setae (excluding coxal and sternal); NDV = DS + V.
Table 3. Morphometric (AW–dmt, μm) and meristic (DS–NDV) traits of Neacariscus (Whartonacarus) shiraii (Sasa, Kano & Obata, 1952) and Neacariscus (Whartonacarus) ... Stekolnikov, n. sp. Abbreviations: AW – distance between anterolateral scutal setae; PW – distance between posterolateral scutal setae; SB – distance between sensillary (trichobothrial) bases; ASB – distance from the level of sensillary bases to extreme anterior margin of scutum; PSB – distance from the level of sensillary bases to extreme posterior margin of scutum; SD – length of scutum (ASB + PSB); P-PL – distance from the level of posterolateral scutal setae (PL) to extreme posterior margin of scutum; AP – distance between antero- and posterolateral scutal seta on one side; AM – length of anteromedian scutal seta; AL – length of anterolateral scutal setae; PL – length of posterolateral scutal setae; S – length of sensilla (trichobothria); H – length of humeral setae; Dmin – length of the shortest dorsal idiosomal seta; Dmax – length of the longest dorsal idiosomal seta; Vmin – length of the shortest ventral idiosomal seta; Vmax – length of the longest ventral idiosomal seta; pa – length of leg I (including coxa, excluding claws); pm – length of leg II (including coxa, excluding claws); pp – length of leg III (including coxa, excluding claws); Ip – sum of leg lengths (pa + pm + pp); TaIIIL – length of leg tarsus III; TaIIIW – width of leg tarsus III; dmt – distance between the base of leg III tarsus and base of mastitarsala; DS – number of dorsal idiosomal setae (excluding scutal); V – number of ventral idiosomal setae (excluding coxal and sternal); NDV = DS + V.
Neacariscus shiraii Neacariscus ... n. sp. (n = 7)
ZIN 17981 ZIN 17982 Holotype* Specimen from Guam** Range Mean Holotype
AW 84 74 81 80 77-81 78 77
PW 95 88 89 92 86-95 91 90
SB 29 29 30 28 25-30 28 28
ASB 36 36 35 33 39-42 40 40
PSB 21 20 20 19 23-27 25 25
SD 57 56 55 52 63-66 65 65
P-PL 13 11 - - 14-20 17 18
AP 37 34 33 32 35-44 40 41
AM 45 54 61 - 45-50 47 45
AL 56 49 50 46 45-51 50 50
PL 92 83 82 84 55-83 72 76
S - - 82 90 87-92 89 90
H 104 103 93 64 81-94 86 94
Dmin 63 67 - 54 56-68 60 59
Dmax 88 88 70 72 68-77 74 76
Vmin 49 49 38 40 35-41 38 41
Vmax 79 81 - 60 61-70 64 63
pa 533 513 - 480 558-581 572 574
pm 472 455 - 416 497-515 509 508
pp 545 518 - 485 580-607 589 583
Ip 1550 1487 - 1381 1642-1697 1670 1665
TaIIIL 162 160 - - 178-189 181 182
TaIIIW 22 22 - - 22-23 22 23
dmt 74 74 - - 64-81 72 76
DS 80 77 92 84 42-49 45 44
V 85 85 98 96 64-84 70 74
NDV 165 162 190 180 107-133 115 118
Notes: * – after original description, cited by Vercammen-Grandjean & Langston (1976); ** – slide NAMRU 2-697, after the same work.
Table 4. Morphometric (AW – dmt, μm) and meristic (DS – NDV) traits of Neacariscus (Whartonacarus) sulae (Oudemans, 1910). Abbreviations as in Table 3.
Table 4. Morphometric (AW – dmt, μm) and meristic (DS – NDV) traits of Neacariscus (Whartonacarus) sulae (Oudemans, 1910). Abbreviations as in Table 3.
ZIN 17962 ZIN 18168 Holotype*
AW 83 86 88
PW 99 97 104
SB 37 34 40
ASB 34 35 32
PSB 20 21 20
SD 54 56 52
P-PL - 16 -
AP 33 34 32
AM - 36 ca. 35
AL - 52 38
PL 85 88 83
H 99 - 94
Dmin - 68 60
Dmax - 85 92
Vmin - 38 42
Vmax - 59 60
pa 391 405 386
pm 365 380 355
pp 425 421 404
Ip 1181 1206 1145
TaIIIL 115 121 -
TaIIIW 20 20 -
dmt - 35 -
DS 36 32 36
V 65 56 50
NDV 101 88 86
Note: * – after Vercammen-Grandjean & Langston (1976).
Table 5. Morphometric (AW – TaIIIW, μm) and meristic (DS – NDV) traits of Toritrombicula kirhocephales Goff, 1982 (n = 5). Abbreviations as in Table 3.
Table 5. Morphometric (AW – TaIIIW, μm) and meristic (DS – NDV) traits of Toritrombicula kirhocephales Goff, 1982 (n = 5). Abbreviations as in Table 3.
Range Mean Type series, range*
AW 65-74 70 69-77
PW 79-90 84 84-89
SB 22-27 25 25-28
ASB 32-36 34 27-33
PSB 16-18 17 16-18
SD 49-54 51 -
P-PL 9-14 11 -
AP 32-40 35 31-33
AM 38-43 41 43-45
AL 46-51 48 41-52
PL 54-71 65 51-58
H 50-64 57 54-64
Dmin 32-36 34 37
Dmax 54-68 61 54
Vmin 32-33 33 29
Vmax 54-56 55 47
pa 396-443 416 355-361
pm 333-365 347 309-316
pp 405-437 416 362-368
Ip 1145-1246 1186 1026-1045
TaIIIL 112-121 115 98
TaIIIW 18-20 19 18
DS 34-37 35 32
V 31-34 32 26
NDV 65-68 67 58
Note: * – After Goff (1982).
Table 6. Morphometric (AW – TaIIIW, μm) and meristic (DS – NDV) traits of Neoschoengastia longipes Nadchatram, 1967 (n = 8). Abbreviations as in Table 3.
Table 6. Morphometric (AW – TaIIIW, μm) and meristic (DS – NDV) traits of Neoschoengastia longipes Nadchatram, 1967 (n = 8). Abbreviations as in Table 3.
Range Mean Original description*
AW 46-58 53 51, 53
PW 62-90 77 80, 86
SB 26-35 30 32, 31
ASB 20-27 23 21, 23
PSB 24-29 26 24, 26
SD 44-56 49 -
P-PL 7-16 10 -
AP 34-41 38 34, 33
AM 35-43 38 31, 35
AL 59-74 65 69, 68
PL 54-70 59 59, 57
S (length) 38-38 38 36, 34
S (width) 20-22 21 19, 19
H 48-58 52 49
Dmin 27-34 29 -
Dmax 49-56 52 44-46
Vmin 25-28 26 -
Vmax 31-41 36 30-34
pa 403-437 421 410-460
pm 360-398 373 357-370
pp 425-461 441 380-400
Ip 1197-1294 1234 1140-1230
TaIIIL 122-139 131 140
TaIIIW 14-18 16 14
DS 28-31 30 30
V 19-25 22 -
NDV 50-54 52 -
Note: * – Holotype and mean of 11 specimens; single unspecified value or range for some characters; after Nadchatram (1967).
Table 7. Morphometric (AW – TaIIIW, μm) and meristic (DS – NDV) traits of Schoengastia archaea (Taufflieb, 1960) (n = 7). Abbreviations as in Table 3.
Table 7. Morphometric (AW – TaIIIW, μm) and meristic (DS – NDV) traits of Schoengastia archaea (Taufflieb, 1960) (n = 7). Abbreviations as in Table 3.
Range Mean Original description*
AW 34-36 35 38
PW 51-56 53 56
SB 23-25 24 23
ASB 26-29 28 24
PSB 29-31 29 33
SD 56-58 57 57
P-PL 27-34 31 -
AP 17-21 19 19
AM 23-25 25 25
AL 56-65 60 58
PL 56-61 58 61
S (length) 27-30 28 29
S (width) 14-15 15 -
Dmin 20-22 20 25
Dmax 28-34 32 29
Vmin 20-24 21 22
Vmax 24-30 27 25
pa 371-401 386 362
pm 297-329 311 299
pp 351-396 372 358
Ip 1024-1127 1069 1019
TaIIIL 101-110 105 100
TaIIIW 15-19 16 -
DS 138-159 148 160
V 89-100 94 96
NDV 227-254 241 256
Note: * – After Taufflieb (1960).
Table 8. Morphometric (AW – TaIIIW, μm) and meristic (DS – NDV) traits of Ericotrombidium cosmetopode (Vercammen-Grandjean & Langston, 1971). Abbreviations as in Table 3.
Table 8. Morphometric (AW – TaIIIW, μm) and meristic (DS – NDV) traits of Ericotrombidium cosmetopode (Vercammen-Grandjean & Langston, 1971). Abbreviations as in Table 3.
New specimen Holotype* Mean (n = 4)*
AW 60 62 62
PW 75 72 71
SB 23 23 22
ASB 28 26 26
PSB 13 12 13
SD 41 38 38
P-PL 14 - -
AP 24 23 22
AM 33 42 39
AL 35 39 38
PL 43 47 47
S 59 64 64
H 38 42 43
Dmin 33 38 37
Dmax 41 46 44
Vmin 24 32 31
Vmax 40 44 43
pa 274 236 235
pm 234 206 206
pp 263 238 241
Ip 770 680 682
TaIIIL 74 - -
TaIIIW 14 - -
DS 28 32 32
V 27 28 28
NDV 55 60 60
Note: * – After Vercammen-Grandjean and Langston (1971, 1976).
Table 9. Distribution of chiggers by host orders in Thailand. Names of localities as in Table 1. Sum of the column ‘Hosts’ is larger than the total number of host individuals examined due to the co-parasitization of one host with different chigger species.
Table 9. Distribution of chiggers by host orders in Thailand. Names of localities as in Table 1. Sum of the column ‘Hosts’ is larger than the total number of host individuals examined due to the co-parasitization of one host with different chigger species.
Host order Hosts Chigger species Slides Province Locality Habitat Month
Charadriiformes 4 Neacariscus ... n. sp. 9 Satun Thung Sabo Shore March
Charadriiformes 3 Schoengastia archaea 8 Trang Albino Dragon Shore March
Charadriiformes 1 Neacariscus sulae 2 Trang Koh Libong Shore March
Charadriiformes 12 Neacariscus ... n. sp. 17 Trang Samran beach Shore March
Charadriiformes 2 Neacariscus shiraii 2 Trang Samran beach Shore March
Charadriiformes 3 Neacariscus pluvius 3 Trang Samran beach Shore March
Charadriiformes 8 Schoengastia archaea 14 Trang Samran beach Shore March
Coraciiformes 1 Neoschoengastia longipes 5 Rayong Koh Mun Nai Island March
Passeriformes 4 Neoschoengastia longipes 8 Rayong Koh Mun Nai Island March
Passeriformes 1 Neoschoengastia solitus 3 Rayong Koh Mun Nai Island March
Passeriformes 3 Odontacarus audyi 3 Rayong Koh Mun Nai Island March
Passeriformes 2 Toritrombicula densipiliata 3 Rayong Koh Mun Nai Island March
Passeriformes 6 Toritrombicula kirhocephales 17 Rayong Koh Mun Nai Island March
Passeriformes 51 Odontacarus audyi 140 Kanchanaburi Mahidol Univ. Highland December
Passeriformes 6 Toritrombicula kirhocephales 8 Kanchanaburi Mahidol Univ. Highland December
Passeriformes 29 Neoschoengastia longipes 73 Kanchanaburi Mahidol Univ. Highland December
Strigiformes 1 Neoschoengastia longipes 2 Kanchanaburi Mahidol Univ. Highland December
Galliformes 29 Eutrombicula wichmanni 115 Nan Huai Muang Village December
Galliformes 1 Helenicula comata 3 Nan Huai Muang Village December
Galliformes 1 Neoschoengastia gallinarum 4 Nan Huai Muang Village December
Galliformes 1 Eutrombicula wichmanni 2 Nan Ban Santisuk Village December
Galliformes 1 Odontacarus audyi 1 Nan Ban Santisuk Village December
Table 10. Distribution of chiggers by host orders in Malaysia. Names of localities as in Table 2. Sum of the column ‘Hosts’ is larger than the total number of host individuals examined due to the co-parasitization of one host with different chigger species.
Table 10. Distribution of chiggers by host orders in Malaysia. Names of localities as in Table 2. Sum of the column ‘Hosts’ is larger than the total number of host individuals examined due to the co-parasitization of one host with different chigger species.
Host order Hosts Chigger species Slides Province Locality Month
Coraciiformes 1 Parascoschoengastia heynemani 4 Pahang Krau February
Coraciiformes 1 Parascoschoengastia heynemani 4 Sarawak Gunung Gading March
Coraciiformes 2 Toritrombicula densipiliata 4 Perak Behrang September
Cuculiformes 1 Odontacarus audyi 1 Selangor Ulu Gombak February
Cuculiformes 1 Toritrombicula densipiliata 1 Selangor Ulu Gombak February
Galliformes 6 Neoschoengastia gallinarum 16 Johor Jemaluang February
Galliformes 5 Neoschoengastia gallinarum 11 Johor Kota Tinggi June
Galliformes 6 Neoschoengastia gallinarum 24 Perak Sungkai January, March
Galliformes 7 Neoschoengastia gallinarum 9 Selangor Bestari Jaya April
Gruiformes 1 Blankaartia acuscutellaris 2 Sarawak Pueh March
Gruiformes 2 Leptotrombidium imphalum 2 Sarawak Pueh March
Passeriformes 1 Ericotrombidium cosmetopode 1 Pahang Fraser’s Hill October
Passeriformes 1 Leptotrombidium deliense 1 Pahang Fraser’s Hill October
Passeriformes 1 Leptotrombidium deliense 1 Kedah Langkawi March
Passeriformes 1 Leptotrombidium miculum 1 Kedah Langkawi March
Passeriformes 3 Odontacarus audyi 3 Kedah Langkawi March
Passeriformes 1 Odontacarus audyi 1 Pahang Fraser’s Hill October
Passeriformes 4 Odontacarus audyi 4 Selangor Ulu Gombak February
Passeriformes 14 Odontacarus audyi 16 Terengganu Pasir Raja October
Passeriformes 5 Toritrombicula densipiliata 5 Kedah Langkawi March
Passeriformes 21 Toritrombicula densipiliata 41 Pahang Krau February
Passeriformes 14 Toritrombicula densipiliata 25 Perak Behrang September
Passeriformes 2 Toritrombicula densipiliata 2 Perak Larut Hill March
Passeriformes 4 Toritrombicula densipiliata 4 Sarawak Gunung Gading March
Passeriformes 1 Toritrombicula densipiliata 2 Sarawak Sematan Pueh March
Passeriformes 1 Toritrombicula densipiliata 2 Selangor Ulu Gombak February
Passeriformes 6 Toritrombicula densipiliata 6 Terengganu Pasir Raja October
Passeriformes 11 Toritrombicula densipiliata 33 Terengganu Redang Isl. October
Passeriformes 1 Toritrombicula uphami 1 Kedah Langkawi March
Passeriformes 2 Toritrombicula uphami 2 Terengganu Pasir Raja October
Piciformes 1 Ascoschoengastia lorius 12 Pahang Krau February
Podargiformes 1 Toritrombicula densipiliata 1 Perak Behrang September
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