Validation of the status of a species with high CO 1 and low nuclear genetic divergences: the scab mite Caparinia ictonyctis stat. res. (Acariformes: Psoroptidae) parasitizing the African hedgehog Atelerix albiventris Author Bochkov, Andre V. Author Klimov, Pavel B. Author Kim, Da-Hee Author Skoracki, Maciej text Zootaxa 2019 2019-01-15 4544 4 523 547 journal article 27657 10.11646/zootaxa.4544.4.4 3f253e3e-777e-465d-841d-a67e3b645d1a 1175-5326 2618505 3739EAED-83E4-4B3C-A81A-E2120DDAA85C Caparinia tripilis ( Michael, 1889 ) ex Erinaceus europaeus : 1 male and 1 tritonymph, Spain , Madrid Province , San Fernando de Henares , 40°25'25''N , 3°31'57''W , 28 June 1977 , coll. M. Portus ( IRSNB ) ; 1 male , 3 females , 4 tritonymphs, 2 larvae , the Netherlands , Nijmegen , 51°50'33''N , 5°51'10''E , 29 August 1967 , coll. F.S. Lukoschus ( IRSNB ) ; 6 males , 2 females , and 2 tritonymphs, Belgium , Flemish Brabant Province , Leefdaal , 50°50'51''N , 4°35'21''E , 11 July 1979 , coll. F. Puylaert ( IRSNB ) ; 1 female , Poland , Kraków , 50°5'N , 19°55'E , 17 July 1966 , coll. Zurowski ( IRSNB ) ; 20 males , 20 females , 10 tritonymphs, 10 protonymphs, and 10 larvae , Russia , St. - Petersburg Province , Gatchina District , near Taytsy village , 59°40'00''N , 30°07'00''E , September 2015 , coll. A.V. Bochkov ( ZISP , AVB 17-0305 - 001 ) [many additional specimens are preserved in alcohol or were used for scanning electron microscopy and molecular analysis]; 10 males , 10 females , 10 tritonymphs, 10 protonymphs, and 10 larvae , Russia , St. Petersburg Province , no further data, 0 5 March 1988 , coll. I. Skiba ( ZISP AVB 14-0505 - 008 ) . ex Erinaceus roumanicus : 10 males , 10 females , 10 tritonymphs, 10 protonymphs, and 10 larvae , Russia , North Caucasus, Caucasus National Reserve, Khosta forest, 23 June 1977 , coll. H. Dubinina (ZISP). Note: the original label says Erinaceus europaeus , however, based on the locality, the host name should be E. roumanicus (Heet al. 2012). Caparinia ictonyctis ( Lawrence, 1955 ) ex Atelerix albiventris : 20 males , 20 females , 10 tritonymphs, 10 protonymphs, and 10 larvae , South Korea , South Leolla Province, Gwangju , pet shop, February 2010 , coll. S.-S. Shin (ZISP AVB 14-0505-004) [many additional specimens are preserved in alcohol or were used for scanning electron microscopy and molecular analysis]; 6 males and 1 female , USA, Maryland , Baltimore Zoo , directly transported from Togo , 4 February 1986 , coll. E. Neely ( IRSNB ) . ex Ictonyx striatus : 3 males and 3 females ( syntypes of Caparinia ictonyctis Lawrence, 1955 ), South Africa , KwaZulu-Natal, Pietermaritzburg , April , 1954, coll. R.F. Lawrence ( KZNM 5767 ); 1 male and 1 female , [original label in French: Ictonyx striatus erythreae , Egypte, Saukin], other data unknown ( IRSNB ). On the original label by A. Fain , the geographical data are probably not quite correct. Ictonyx striatus erythreae is absent in Egypt but occurs in neighbouring Sudan ( Wilson and Reeder 2005 ). The locality “Saukin” is absent in Egypt and according to Geonames database probably is El Qadarif city, 14°02'05''N , 35°23'00''E , Al Qadarif state, Sennar Province , Sudan . Caparinia setifera ( Mégnin, 1880 ) 2 males , 1 female and 2 tritonymphs ( syntypes ) from Hyaena hyaena , [coll. Hist. Nat. Paris ], no other data ( IRSNB ) ; 1 male , 11 females , and 1 tritonymph from same host, South Africa , KwaZulu-Natal , Hluhluwe- Umfolozi Game Reserve , 28°02'21''S , 32°03'31''E , 9 November 1973 , coll. M.E. Keep ( IRSNB ) . Caparinia erinacei Fain, 1962 1 male and 1 female ( paratypes ) from Atelerix frontalis , South Africa , Transvaal [now Gauteng ], Pretoria Zoo, March 1959 , coll. F. Zumpt (MRAC); 2 females and 2 males from “hedgehog”, Tanzania , Arusha Region , Serengeti National Park, 14 September 1971 , coll. T. Mcharo (IRSNB). The undetermined “hedgehog” from Tanzania recorded as host of this species ( Fain 1975 ) is probably Atelerix albiventris , because among African hedgehogs only this species inhabits Tanzania ( He et al. , 2012 ). Caparinia algirus Fain and Portús, 1979 Female ( holotype ), 3 males and 3 females ( paratypes ) from Atelerix algirus , Spain , Bolearic Islands, Formentera Island, Pitiusas, 38°55'50''N , 1°21'45''E , no date, coll. M. Portus (IRSNB); 2 males and 2 males ( paratypes ) from same host, Spain, Catalonia , Barcelona, 02°17'N , 41°41'E , no date, coll. M. Portus (IRSNB). Caparinia lophiomys Fain, 1975 12 males and 9 females ( paratypes ) from Lophiomys imhausi , Somalia , May 1916 , no other data (IRSNB). Morphological descriptions. In the analyses, key, and descriptions, the idiosomal chaetotaxy follows Griffiths et al. (1990) with modifications of Norton (1998) for coxal setae. The leg chaetotaxy follows Grandjean (1939) . All measurements are given in micrometers (µm) and were taken as follows: body length = total length from the anterior extremity of the palps to the posterior border of the body, including the lobar membranes in males; body width = width at the level of setae cp ; length of dorsal shields = maximum length, measured along the median line of the shields; length of male opisthosomal lobes = measured from the anterior edge of the opisthosomal cleft to the posterior end of the lobe; length of the posterior legs = length from the most basal point of the trochanter to the apex of the tarsus, excluding pretarsus. Drawings were made with a Leica microscope equipped with differential interference contrast (Nomarski) optics and a camera lucida. Fine mite structures (mites from Atelerix albiventris , ZISP AVB 14-0505-004) were examined with a scanning electron microscope Quanta 250. Mites were put in 96% ethanol for 24 hours, transferred to hexamethyldisilazane for 10 minutes, and then dried and sputtered with platinum. DNA amplification, sequencing, and alignment. We sequenced individual specimens of Caparinia from Atelerix albiventris and Erinaceus europaeus for 6 genes: two nuclear ribosomal RNA genes, 18S and 28S rDNA; three nuclear protein-coding genes: elongation factor 1alpha100E (EF1-α), signal recognition particle protein 54k (SRP54), Hsc70-5 heat shock protein cognate 5 (HSP70); and one mitochondrial protein-coding gene (CO1). CO1 was sequenced from 14 specimens for Caparinia ex Atelerix albiventris (all were identical) and 2 specimens of Caparinia tripilis ex Erinaceus europaeus . Previously published amplification and sequencing protocols were used ( Bochkov et al. , 2014 ; Klimov & OConnor, 2008 , 2013 ; Knowles & Klimov, 2011 ). GenBank accession numbers are as follows: MG766225-MG766259, MG766261 - MG766269 (Table 1). The sequence of 18S of Caparinia from Erinaceus europaeus (MG766260) was identified as a gregarine (an endoparasitic protozoan) and, therefore, was excluded from further analyses. Domain D4 of 28S rDNA was also excluded because our standard protocol TABLE 1 . Collection data and GenBank accession numbers for select taxa of the scab mite family Psoroptidae (id=unique numbers for DNA extraction vouchers; field number = lot-based museum UMMZ accession numbers for vouchers and co-vouchers).

id Mite species	Host	Country	filed number	18S	28S	EF1-α	SRP54	HSP70	CO1
1647 Caparinia ictonyctis	Atelerix albiventris	SOUTH KOREA	BMOC 13-0508-003	KF891891	MG766268	MG766261	MG766266	MG766227	KF891934
1660 Chorioptes bovis	Ovis aries	ICELAND	BMOC 13-0810-007	KF891892	KF891899	KF891904	KF891910	KF891916	KF891935
1664 Chorioptes sweatmani	Alces alces	SWEDEN	BMOC 13-0810-010	KF891893	KP406736	KF891905	KF891911	KF891917	KF891936
2034 Caparinia tripilis	Erinaceus europaeus	RUSSIA	BMOC 16-0825-012	-*	MG766269	MG766262	MG766264	MG766226	MG766233
2035 Caparinia tripilis	Erinaceus europaeus	RUSSIA	BMOC 16-0825-013	-	-	-	-	-	MG766234
597 Psoroptes ovis	Oryctolagus cuniculus	USA	BMOC 05-0105-001	JQ000241	JQ000549	JQ000850	JQ001154	JQ001469	-
597 Psoroptes ovis	Oryctolagus cuniculus	USA	BMOC 05-0105-001	JQ000241	JQ000549	JQ000850	JQ001154	JQ001469	KJ957822
- Chorioptes bovis	Bison bonasus	POLAND	-	GQ864308	-	-	-	-	-
- Psoroptes ovis	Oryctolagus cuniculus	CHINA	-	-	-	-	-	-	KJ957822
- Psoroptes ovis	Oryctolagus cuniculus	CHINA	-	FJ907505	-	-	-	-	FJ907499
- Psoroptes ovis	Ovis aries	UNITED KINGDOM	-	-	-	-	-	-	BQ835080
- Psoroptes ovis	Ovis aries	UNITED KINGDOM	-	-	-	-	-	-	FR748605

*Gregarine sequence amplified, GenBank accession number MG766260. produced superimposed sequences. Sequences of rDNA were aligned in Mesquite ver. 3.31 ( Maddison & Maddison, 2016 ) using a previously established secondary structure model ( Klimov & OConnor, 2008 ); alignment of other loci was unambiguous. Voucher and co-voucher mite specimens are deposited in the University of Michigan Museum of Zoology, Ann Arbor, Michigan under the following accession numbers: BMOC 13-0508-003 (AD1647): Caparinia ictonyctis , ex Atelerix albiventris ; BMOC 16-0825-012 (AD2034), BMOC 16-0825-013 (AD2035): Caparinia tripilis , ex Erinaceus europaeus . Genetic distances. Following tradition, we use Kimura 2-parameter (K2P) genetic distances ( Hebert et al. , 2003 ), but see ( Srivathsan & Meier, 2012 ) for criticism. Distances were calculated in PAUP* ver. 4.0a (build 158) ( Swofford, 2016 ) using the default settings.