INTRODUCTION
The genus Alloptes Canestrini, 1879 is one of about 23 genera that belong to the family Alloptidae Gaud, 1957 and contains about 46 species (Gaud and Atyeo, 1996; Mironov and Palma, 2006). This genus has been found on flight feathers and wing-coverts of birds from order Charadriiformes(Mironov and Palma, 2006). The genus Alloptes has the following diagnostic characteristics: (1) vertical seta is absent; (2) setae kT of tibiae III and setae d of tarsi II, III are absent; (3) in males, opisthosomal lobes are fused by medial margins into a single lobe; and (4) in females, lateral crest of tarsi III, IV are absent (Vasyukova and Mironov, 1991; Gaud and Atyeo, 1996; Mironov, 1996, 1998).
The genus Alloptes is divided into four subgenera: Alloptes s. str. Gaud, 1972, Apodalloptes Gaud, 1972, Conuralloptes Gaud, 1972, Sternalloptes Kivganov and Mironov, 1992 (Gaud, 1972; Kivganov and Mironov, 1992). These four subgenera are classified based on the setae structure of the anterior legs and opisthosomal chaetotaxy (Gaud, 1972; Kivganov and Mironov, 1992; Mironov, 1998; Mironov and Palma, 2006).
In the present work, we found four unrecorded species [A. (Apodalloptes) orthogramme, A. (Conuralloptes) limosae, A. (C.) procerus and A. (Sternalloptes) fauri] of feather mite during investigations of undiscovered invertebrate species in Korea. Here, we provide morphological description and illustrations of these feather mites with host information, and partial sequences of the mitochondrial cytochrome c oxidase subunit I(COI) as DNA barcodes.
MATERIALS AND METHODS
Feather mite specimens were obtained from the flight feathers and wing-coverts of the following four charadriiform bird species, i.e., black-tailed godwit Limosa limosa, blacktailed gull Larus crassirostris, common sandpiper Actitis hypoleucos, and whimbrel Numenius phaeopus. The carcasses of birds were donated or obtained by present authors from the Chungnam Wild Animal Rescue Center (three waders) or found on dead (the black-tailed gull). The collected mites were preserved directly in 95% ethyl alcohol. The mite specimens were cleared in 10% lactic acid for 24 hours and then mounted on microscope slides using PVA (PVA stock solution 56%, lactic acid 22%, and phenol 22%) as the mounting medium (Downs, 1943). The specimens were photographed using a microscopic digital camera (Leica, Wetzlar, Germany). Terms and measurements followed Gaud and Atyeo (1996), and Norton (1998). All examined specimens were deposited in the National Institute of Biological Resources (NIBR) and Inha University, Incheon, Korea.
DNA sequencing
DNA was extracted from individual mites using a LaboPass Tissue Genomic DNA Isolation Kit Mini (Cosmogenetech Inc., Seoul, Korea) according to the manufacturer’s instructions. PCR amplification, purification, and sequencing were performed as described by Han et al.(2016).
SYSTEMATIC ACCOUNTS
Order Sarcoptiformes Canestrini, 1891
Family Alloptidae Gaud, 1957
Genus Alloptes Canestrini, 1879
Subgenus Apodalloptes Gaud, 1972
Alloptes(Apodalloptes) orthogramme Gaud and Mouchet, 1957 (Figs. 1, 2)
Alloptes orthogramme: Gaud and Mouchet, 1957: 496, fig. 1B; Zumpt, 1961: 241.
Alloptes (Apodalloptes) orthogramme: Gaud, 1972: 60; Vasyukova and Mironov, 1991: 84, fig. 60.
Fig. 1. Alloptes (Apodalloptes) orthogramme, male. A, Dorsal idiosoma; B, Venteral idiosoma. ps, pregenital sclerites. Scale bars: A, B=0.1 mm.
Fig. 2. Alloptes (Apodalloptes) orthogramme, female. A, Dorsal veiw; B, Dorsal idiosoma; C, Ventral idiosoma; D, Opisthosomal lobes. Scale bars: A=0.2 mm; B, C=0.1 mm; D=0.05 mm.
Material examined. 1♂, 1♀, Korea: Chungcheongnam-do, Cheongyang-gun, Jeongsan-myeon (36°22ʹ34ʺN, 126°56ʹ 38ʺE), 12 Sep 2014, collected under a stereomicroscope from flight feathers on the wings of common sandpiper A. hypoleucos by Han Y.-D.
Description. Male (Fig. 1): Idiosoma size 265 ×110 μm (length×width). Prodorsal shield (Fig. 1A): Posterior margin straight, surface without ornamentation, length 63 μm along midline, width of posterior part 64 μm. Hysteronotal shield (Fig. 1A): Anterior margin straight, surface without ornamentation, length 178 μm from anterior margin to bases of setae ps1, width 70 μm at anterior part. Length 6 μm between prodorsal and hysteronotal shields. Setae d2 situated on hysteronotal shield, bases of this setae without incisions. Setae h2 cylindrical, without extensions in the basal half. Distance between dorsal setae: se :se 75 μm, c2 :d2 47 μm, d2 :ps1 105 μm. Subhumeral setae c3 lanceolate. Opisthosoma gradually narrowed to posterior end. Terminal lamella with tree pairs of festoons, teeth-shaped. Sternum (Fig. 1B): Epimerites I fused into a Y-shape. Pregenital sclerite (ps) fused in to a Y-shape and not connected to inner ends of epimerites IIIa. Coxal setae 3a located anterior to setae 4b. Genital arch 19×21 μm (length×width). Setae 4a situated posterior to setae g. Small sclerites present at posterior margins of genital arch. Adanal shields inverted L-shaped, without protrusion at anterior-lateral margins. Setae 4a surrounded by irregular sclerite. Length of genital-anal field 89 μm. Distance between ventral setae: 4b :3a 8 μm, 4b :g 31 μm, 4b: 4a 47 μm, g : ps3 20 μm, ps3:ps1 50 μm, 4a: 4a 52 μm. Setae mG of legs I and II short, spine-like with acute apex. Leg IV short and barely exceeding the end of opisthosomal lobe, 90 μm in length.
Female (Fig. 2): Idiosoma 320 ×125 μm (length ×width) (Fig. 2A). Prodorsal shield (Fig. 2B): Mostly shaped as in male, length 73 μm along midline, width 85 μm. Hysteronotal shield (Fig. 2B): Anterior margin straight, length 220 μm from anterior end to bases of setae h3, width 60 μm at anterior part. Distance 24 μm between prodorsal and hysteronotal shields. Setae h1 located anterior to setae e2. Opisthosomal lobes shorted, terminal cleft as inverted U-shaped. Distance between dorsal setae: c2 :d2 68 μm, d2 : e2 100 μm, e2 :h2 30 μm, h2 :h3 10 μm, h2 : h2 45 μm, h3 :h3 23 μm. Supranal concavity oval, divided from terminal cleft. Sternum (Fig. 2C, D): Epimerites I fused. Epigynum bowshaped, 24×48 μm (length×width). Leg I and II as in the male. Ambulacral disc IV slightly beyond the level of setae f2.
Remarks. Alloptes (A.) orthogramme was originally described by Gaud and Mouchet (1957) based on specimens collected from A. hypoleucos in Cameroun. Thereafter, the ventral hysterosoma of this species was redescribed by Vasyukova and Mironov (1991) from A. hypoleucos in the Sakha Republic (=Yakutia Republic) of Russia.
Alloptes(A.) orthogramme is very similar to A. (A.) curtipes Trouessart, 1885 regarding the external traits. However, A.(A.) orthogramme can be clearly distinguished from A.(A.) curtipes by the following characteristics in males: (1) total body length is less than 350 μm; (2) setae 4a are situated posterior to setae g; and (3) setae h2 are cylindrical-shaped without extensions in the middle part (Gaud, 1972; Vasyukova and Mironov 1991). The morphology of the Korean specimen was consistent with the description and illustrations provided by Gaud and Mouchet (1957), and Vasyukova and Mironov (1991).
Host. This species was found on a flight feather in the wing of common sandpiper, A. hypoleucos.
Distribution. Cameroun (Gaud and Mouchet, 1957; Zumpt, 1961; Gaud, 1972), Russia (Vasyukova and Mironov, 1991), Korea (this study).
Deposition. NIBR No. NIBRIV0000754007-0000754008.
Molecular data. The COI sequence was obtained from single individual and deposited in GenBank with accession number of MK456598.
Subgenus Conuralloptes Gaud, 1972
Alloptes(Conuralloptes) limosae Dubinin, 1951 (Figs. 3, 4)
Alloptes gambettae limosae: Dubinin, 1951: 239, fig. 65.
Alloptes (Conuralloptes) limosae: Gaud, 1972: 66, figs. 26, 29; Vasyukova and Mironov, 1991: 98, fig. 72.
Fig. 3. Alloptes (Conuralloptes) limosae, male. A, Dorsal veiw; B, Dorsal idiosoma; C, Ventral idiosoma. ps, pregenital sclerites. Scale bars: A-C=0.1 mm.
Fig. 4. Alloptes (Conuralloptes) limosae, female. A, Dorsal veiw; B, Dorsal idiosoma; C, Ventral idiosoma; D, Opisthosomal lobes. Scale bars: A=0.2 mm; B, C=0.1 mm; D=0.05 mm.
Material examined. 6♂♂, 3♀♀, Korea: Chungcheongnam-do, Asan-si, Tangjeong-myeon (36°48ʹ58ʺN, 126°2ʹ 45ʺE), 18 May 2017, collected using dissecting microscope from flight feathers on the wings of black-tailed godwit L. limosa by Han Y.-D.
Description. Male (Fig. 3): Idiosoma 290-300×135-155 μm (length×width). Prodorsal shield (Fig. 3B): Posterior margin concave, length 74-82 μm along midline, width of posterior part 90-95 μm. Hysteronotal shield (Fig. 3B): Anterior part slightly concave, lateral margins with small incision at bases of setae d2, length 188-198 μm from anterior margin to bases of setae ps1, width 70-72 μm at anterior part. Distance 10-17 μm between prodorsal and hysteronotal shields along midline 10-17 μm. Distance between dorsal setae: se :se 98-105 μm, c2:d2 37-45 μm, d2 :ps1 118-125 μm. Subhumeral setae c3 lanceolate. Opisthosoma tapered to posterior end. Terminal lamella with 6 dentations, incision between inner pair slit-shaped. Setae h2 cylindrical-shaped, without any expanded on basal half (Fig. 3A). Sternum (Fig. 3C): Epimerites I fused into a Y-shape. Pregneital sclerites (ps) connected to inner ends of epimerites IIIa and paragenital arch, divided from each other. Length of genital-anal field 97-105 μm. Setae 4b situated on the same level of setae 3a. Genital arch 19-20×17-21 μm (length×width). Setae 4a surround by irregular sclerite. Distance between ventral setae: 4b :g 37-44 μm, 4b: 4a 65-67 μm, g :ps3 24-26 μm, ps3:ps1 59-67 μm, 4a : 4a 90-97 μm. Setae mG of legs I and II spine-like with acute and blunted apex, respectively. Length of leg IV 153-155 μm.
Female (Fig. 4): Idiosoma 325-335×120-140 μm(length×width). Prodorsal shield (Fig. 4B): Mostly shaped as in male, length 87-90 μm along midline, width 87-91 μm. Hysteronotal shield (Fig. 4B): Anterior margin straight, length 225- 237 μm from anterior end to bases of setae h3, width 56-61 μm at anterior part. Distance 14-17 μm between prodorsal and hysteronotal shields. Setae h1 located anterior to setae e2. Opisthosomal lobes well developed, terminal cleft as inverted U-shaped. Distance between dorsal setae: c2:d2 55- 62 μm, d2 : e2 92-101 μm, e2 : h2 40-43 μm, h2 : h3 14-17 μm, h2 :h2 55-59 μm, h3 :h3 32-37 μm. Supranal concavity oval, divided from terminal cleft. Sternum (Fig. 4C, D): Epimerites I fused. Epigynum bow-shaped, 23-27×40-43 μm (length×width). Leg I and II as in the male. Ambulacral disc IV extending to or slightly beyond the level of setae h2.
Remarks. Alloptes(C.) limosae was originally described by subspecies of A. gambettae limosae by Dubinin (1951) based on specimens collected from L. limosa in Russia. Thereafter, this species was redescribed by several mite taxonomists with detailed morphological description and illustrations (Gaud, 1972, Vasyukova and Mironov, 1991).
Alloptes (C.) limosae is highly similar to A. (C.) procerus Gaud, 1972 regarding external traits. However, A. (C.) limosae can be clearly distinguished from A.(C.) procerus by the following characteristics in males: (1) distance between prodorsal and hysteronotal shields is less than 20 μm; (2) setae g are located closer to setae 4b than setae ps3; and (3) setae h2 are cylindrical-shaped without enlargement in basal half (Gaud, 1972; Vasyukova and Mironov, 1991). The morphology of Korean specimens was consistent with the description and illustrations provided by Gaud (1972), and Vasyukova and Mironov (1991). However, unlike the description by Gaud (1972), the bases of setae g in all Korean males are closer to setae ps3 than to setae 4b. Although there is no description in Vasyukova and Mironov (1991), the bases of setae g in the illustration are closer to setae ps3 than to setae 4b. Therefore, we consider that setae g are closer to setae ps3 than setae 4b.
Host. This species was found on a flight feather in the wing of the black-tailed godwit L. limosa
Distribution. Morocco (Gaud, 1972), Russia (Dubinin, 1951; Vasyukova and Mironov, 1991), Korea (this study).
Deposition. NIBR No. NIBRIV0000812910, NIBRIV 0000843152-0000843159.
Molecular data. The COI sequences were obtained from three individuals and deposited in GenBank with accession numbers of MK456599-456601.
3. Alloptes(Conuralloptes) procerus Gaud, 1972 (Figs. 5, 6)
Alloptes(Conuralloptes) procerus: Gaud, 1972: 64, figs. 26b, 27c, 29a; Vasyukova and Mironov, 1991: 98.
Fig. 5. Alloptes (Conuralloptes) procerus, male. A, Dorsal veiw; B, Dorsal idiosoma; C, Ventral idiosoma. ps, pregenital sclerites. Scale bars: A-C=0.1 mm.
Fig. 6. Alloptes (Conuralloptes) procerus, female. A, Dorsal idisoma; B, Ventral idiosoma. Scale bars: A, B=0.1 mm.
Material examined. 2♂♂, 3♀♀, Korea: Chungcheongnam-do, Taean-gun, Anmyeon-eup (36°36ʹ4ʺN, 126°21ʹ 58ʺE), 1 May 2014, collected under a dissecting microscope from covert feathers on the wings of whimbrel, N. phaeopus by Han Y.-D.
Description. Male (Fig. 5): Idiosoma 305×140-150 μm (length×width). Prodorsal shield (Fig. 5B): Posterior margin concave, length 71-73 μm along midline, width of posterior part 90 μm. Hysteronotal shield (Fig. 5B): Anterior part slightly concave, lateral margins with small incision at bases of setae d2, setae d2 on margin of these incisions, length 203-205 μm from anterior margin to bases of setae ps1, width 65-69 μm at anterior part. Distance 19-22 μm between prodorsal and hysteronotal shields. Subhumeral setae c3 lanceolate. Opisthosoma gradually narrowed to posterior end. Terminal lamella with tree pairs of festoons, incision between inner pair slit-shaped. Setae h2 cylindrical-shaped, slightly enlarged and flattened in middle part (Fig. 5A). Distance between dorsal setae: se :se 98 μm, c2 :d2 37-39 μm, d2 : ps1 125-35 μm. Sternum (Fig. 5C): Epimerites I fused into a Y-shape. Pregneital sclerites (ps) connected to inner ends of epimerites IIIa and paragenital arch, separated from each other. Length of genital-anal field 118-113 μm. Coxal setae 4b located slightly posterior to 3a. Genital arch 21×22 μm (length×width). Setae 4a surrounded by irregular sclerite. Distance between ventral setae: 3a:4b 5-9 μm, 4b: g 18-23 μm, 4b:4a 57 μm, g:ps3 28-32 μm, ps3:ps1 72-76 μm, 4a : 4a 91-95 μm. Setae mG of legs I and II spine-like with acute and blunted apex, respectively. Length of legs IV 155 μm.
Female (Fig. 6): Idiosoma 355-365×120-145 μm(length× width). Prodorsal shield (Fig. 6A): Mostly shaped as in male, length 68-71 μm along midline, width 84-86 μm. Hysteronotal shield (Fig. 6A): Anterior margin straight, length 240-248 μm from anterior end to bases of setae h3, width 54 μm at anterior part. Distance 20-27 μm between prodorsal and hysteronotal shields. Setae h1 located anterior to setae e2. Opisthosomal lobes well developed, terminal cleft as inverted U-shaped. Distance between dorsal setae: c2 : d2 64-66 μm, d2 : e2 103-105 μm, e2 :h2 42-46 μm, h2 :h3 21-22 μm, h2 : h2 64-66 μm, h3 :h3 34-37 μm. Supranal concavity oval, incompletely separated from terminal cleft. Sternum (Fig. 6B): Epimerites I fused. Epigynum bow-shaped, 25-28×43-47 μm (length×width). Leg I and II as in the male. Ambulacra of legs IV not extending to the level of setae h2.
Remarks. Alloptes (C.) procerus was originally described by Gaud (1976) based on specimens collected from N. phaeopus in Cameroun and Mozambique. Thereafter, this species was recorded by Vasyukova and Mironov (1991) from N. phaeopus in Russia.
Alloptes (C.) procerus is very similar to A. (C.) limosae Dubinin, 1951 regarding external traits. However, A. (C.) procerus can be clearly distinguished from A. (C.) limosae by the following characteristics: (1) setae g are situated at middle of setae 4b and ps3 in males; (2) outer sclerites of termial lobe are almost straight in males; (3) distance between prodorsal and hysteronotal shields is more than 30 μm in females; and (4) supranal concavity is incompletely separated from terminal cleft in females(Gaud, 1972).
The Korean specimens were highly similar to Gaud (1972), although differences were found in the following characteristics: in males, setae g are closer to setae 4b than setae ps3; and in females, distance between prodorsal and hysteronotal shields is 22-27 μm. We considered these difference to be an intraspecific variation.
Host. This species was found on a flight feather in the wing of the whimbrel, N. phaeopus.
Distribution. Cameroun, Mozambique (Gaud, 1972), Russia (Vasyukova and Mironov, 1991), Korea (this study).
Deposition. NIBR No. NIBRIV0000843147-0000843151.
Molecular characteristics. The COI sequences were obtained from three individuals and deposited in GenBank with accession numbers of MK456602-MK456604.
Subgenus Sternalloptes Kivganov and Mironov, 1992
Alloptes(Sternalloptes) fauri Gaud, 1957 (Figs. 7, 8)
Alloptes fauri: Gaud, 1957: 111, figs. 1D, 2C.
Alloptes(Conuralloptes) fauri: Gaud, 1976: 13, figs. 2b, 3b, 4b; Vasyukova and Mironov, 1991: 89, fig. 64.
Alloptes(Sternalloptes) fauri: Kivganov and Mironov, 1992: 199.
Fig. 7. Alloptes (Sternalloptes) fauri, male. A, Dorsal idisoma; B, Ventral idisoma. ps, pregenital sclerites. Scale bars: A, B=0.1 mm.
Fig. 8. Alloptes (Sternalloptes) fauri, female. A, Dorsal idiosoma; B, Ventral idisoma. Scale bars: A, B=0.1 mm.
Material examined. 10♂♂, 9♀♀, Korea: Gyeongsangbukdo, Ulleung-gun, Dokdo (37°14ʹ20ʺN, 131°52ʹ3ʺE), 16 Apr 2014, collected using dissecting microscope from flight feathers in the wing of the black-tailed gulls, L. crassirostris by Han Y.-D.
Description. Male (Fig. 7): Idiosoma 340-365×195-215 μm (length ×width). Prodorsal shield (Fig. 7A): Posterior margin concave, length 73-78 μm along midline, width of posterior part 103-110. Hysteronotal shield (Fig. 7A): Anterior part slightly concave, lateral margins with small incision at bases of setae d2, length 230-238 μm from anterior margin to bases of setae ps1, width 95-115 μm at anterior part. Length 25-50 μm between prodorsal and hysteronotal shields. Subhumeral setae c3 narrowly lanceolate. Opisthosoma gradually narrow and slightly expanded at the end. Terminal lamella with 6 dentations, incision between inner pair slot-shaped. Setae h2 cylindrical-shaped, without enlargement in basal half. Distance between dorsal setae: se :se 93-10 μm, c2 :d2 55-57 μm, d2 :ps1 145-158 μm. Sternum (Fig. 7B): Epimerites I fused into a Y-shape. Pregenital sclerites (ps) Y-shaped, connected to inner ends of epimerites IIIa and paragenital arch. Genital arch 15- 20×18-20 μm (length×width). Coxal setae 4b located anterior to 3a. Setae 4a surrounded by irregular sclerite. Distance between ventral setae: 3a : 4b 11-20 μm, 4b: g 48-55 μm, 4b: 4a 66-75 μm, g:ps3 32-38 μm, g:h2 118-129 μm, ps3 :ps1 82-94 μm, 4a :4a 100-110 μm. Setae mG of legs I and II spine-like with acute and blunted apex, respectively. Length of legs IV 195-213 μm, tarsus IV 49-53 μm in length. Solenidion φ of tibia IV about 1.6-2.0 times length of tarsus IV.
Female (Fig. 8): Idiosoma 350-395×150-175 μm(length× width). Prodorsal shield (Fig. 8A): Mostly shaped as in male, length 70-75 μm along midline, width 90-105 μm. Hysteronotal shield (Fig. 8A): Anterior margin straight, length 233-260 μm from anterior end to bases of setae h3, width 65-78 μm at anterior part. Setae h1 located slightly anterior to setae e2. Supranal concavity oval, divided from terminal cleft. Opisthosomal lobes well developed, terminal cleft as inverted U-shaped. Anterior end of supranal concavity not extending to the level of setae e2. Length 50-53 μm between anterior end of supranal and posterior ends of opisthosomal lobes. Distance between dorsal setae: se :se 82-96 μm, c2 :d2 60-69 μm, d2: e2 105-118 μm, e2 :h2 32- 40 μm, h2 :h3 19-22 μm, h2 :h2 53-59 μm, h3 :h3 27-35 μm. Sternum (Fig. 8B): Epimerites I fused. Epigynum bowshaped, 22-26×50-58 μm (length×width). Leg I and II as in the male. Ambulacral discs of legs IV reaching to the level of setae h2.
Remarks. Alloptes (S.) fauri was originally described by Gaud (1957) based on specimens collected from the lesser black-backed gull Larus fuscus in Morocco. Thereafter, this species was redescribed by Vasyukova and Mironov (1991) with detailed morphological description and illustrations of ventral hysterosoma from L. fuscus in Russia.
Alloptes (S.) fauri is highly similar to A. (S.) bisetatus Haller, 1882 regarding external traits. However, A. (S.) fauri can be clearly distinguished from Alloptes (S.) bisetatus by the following combination characteristics: (1) distance between setae g and h2 are 2.5-2.7 times longer than the that between setae 4b and g in males; (2) setae c3 are equal to or slightly longer than 30 μm in males; (3) anterior end of supranal concavity does not extend to the level of setae e2 in females; (4) distance between anterior margin of supranal concavity and posterior end of opisthosomal lobe is less than 60 μm in females (Gaud, 1976). The morphology of Korean specimens was consistent with the original descriptions and illustrations of Gaud (1976).
Host. This species was found on flight feathers in the wing of the black-tailed gull, L. crassirostris.
Distribution. Morocco (Gaud, 1957), South Africa (Gaud, 1976), Russia (Dubinin, 1951; Vasyukova and Mironov, 1991), Korea (this study).
Deposition. NIBR No. NIBRIV0000812912, NIBRIV 0000843160-0000843177.
Molecular characteristics. The COI sequences were obtained from two individuals and deposited in GenBank with accession numbers of MK456605 and MK456606.
ACKNOWLEDGMENTS
The authors wish to thank Jin-Ho Jang (Chungnam Wild Animal Rescue Center, Korea), Prof. Keeseon S. Eom and Dr. Seongjun Choe (Chungbuk National University School of Medicine, Korea) for sample collection, and Dr. Sergey Mironov (Zoological Institute, Russian Academy of Sciences, Russia) for advice and help in identifying the species.
This work was supported by grants from the National Institute of Biological Resources (NIBR), funded by the Ministry of Environment (MOE) of the Republic of Korea (NIBR20161201, 201701201) and Inha University
참고문헌
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