A review of host-commensal associations between canestriniid mites (Astigmata: Canestriniidae) and Insecta with keys and descriptions of the new genera

Abstract The data on host-mites associations between Canestriniidae and insects are summarized. Names of many hosts, incorrectly mentioned in the original works, have been corrected. 9 new genera have been described: Balearella, Cercediphotia, Ciprusenia, Ismaielienia, Javanellina, Kuramasenia, Midiphotia, Phaleratusellus and Sumatranella, 24 species are transferred to another or new genera Balearella asilahica, B. balearica, B. samsinaki, Ciprusenia carabicola, C. izabelae, C. kamelskyi, C. sellnicki, C. viviannae, Imaielienia egypti, I.maroccana, Javanellina macgillavryi, Kuramasenia illegalis, K. paavoi, K. pekingensis, K. pictura, K. ramoni, Midiphotia hispanica, M. jureceki, Phaleratusellus alatus, Psedocanestrinia damiani, P. roberti, Sumatranella batocerae, S. elae and S.phoebinae. 370 species of canestriniids belonging to 101 genera were collected from 361 species of hosts belonging to 113 genera. Canestriniidae were found in 12 Insecta families and in 2 Diplopoda families but species of the families Blattidae and Geotrupidae are accidental hosts. The main hosts of Canestriniidae are Cassidinae (Chrysomelidae) (77 canestriniid species), Cetoniinae (Scarabaeidae) (48), Carabinae (Carabidae) (42), Macroliniinae (Passalidae) (35) and Dynastinae (Scarabaeidae) (20). About 58% of the canestriniid genera are associated with one genus of host and about 60% of canestriniid species are associated with one host species. Canestriniids with large host number have also been found: Arrunsithiana battosi (15 host species), Grandiellopsis gambosa (15) and many others. Most canestriniid species found in Asia (109), South America (78), Africa (66), Europe (51), Oceania (33) and Central America (28). Many species of canestriniids were found in Indonesia (47), Brazil (44) and Papua New Guinea (23). Keyes to genera of the Asia, Europe and Oceania are provided. https://doi.org/urn:lsid:zoobank.org:pub:4296DC9D-5430-423D-9C88-E97617E1D7AC


Introduction
Canestriniidae belongs to the least known families of mites among the Astigmata. They are mainly phoretically related to some insect families. However, some of the canestriniids may be parasites. It was established that the sister mite of Vereoxia bogeri Haitlinger, 1995 was extracting hemolymph from their hosts (Okabe et al. 2017). So far, these mites have been found on insects belonging to12 families: Blattidae, Carabidae, Cerambycidae, Chrysomelidae, Elateridae, Geotrupidae, Lucanidae, Passandridae, Passalidae, Scarabaeidae, Tenebrionidae and Zoopheridae (Tables I-XIV). Their presence in Blattidae and Geotrupidae is random. Moreover, one species Diplopodocoptes transkeiensis Fain, 1987 was collected from the Diplopoda (Fain 1987b). This requires confirmation that Diplopoda are in fact permanent hosts of this species. Sometimes is included in the Canestriniidae Sphaegus elseni Fain, 1974 described on the basis of the hypopus stage (Fain 1974), S. elseni was collected in Africa from Sphex furmosus Christ, 1791[=S. unbrosum metallicus (Taschenberg, 1869] (Hymenoptera: Sphecidae). Fain included this species in the family Acaridae and subfamily Rizoglyphinae. Therefore, this species has not been included in the tables. Currently, 370 species of Canestriniidae are known. Except 370 canestriniid species are included in Table 2 subspecies, 18 species determined to the genus, 6 species determined to the family and 4 species determined as near known species (Okabe & Goka 2008). In the future, this number of species can be expected to be at least doubled. The knowledge of their relationship to the hosts is based mainly on taxonomic work. So far, only one paper has been published discussing the relationship of canestriniid mites with Carabidae (Samšiňák 1971). Santiago-Blay and Fain (1994) published list of mites (including canestriniid mites) associated with Chrysomelidae and Haitlinger (2000) introduced a list of canestriniids occurring on Tenebrionidae. Many species need a generic revision. In this work 9 new genera are described and 24 species were transferred to another genus. In paper on Ibizella balearica n. gen. from Balearic Islands (Haitlinger & Šundić 2016) was described new genus. According to International Code on Zoological Nomenclature, generic name created after 1999 without type species designation are unavailable. Therefore, in this work a new genus Balearella is described and the designated typical species B. balearica.
Keys to the genera for females and males of Asia, Europe and Oceania are provided. All species of canestriniid mites are listed in this work, also undetermined species and all their hosts. Many insects were described from museums material labeled mistakenly or with synonyms and sometimes impossible to identify. In this work, if possible, many names were verified. The aim of this work is providing current knowledge about the relationship between canestriniid mites and their hosts.

Material and methods
The list of canestriniid mites and their hosts was prepared based on literature data. In many papers names of insects are improper or distorted. Canestriniid mites whose generic identify is questionable are marked with an asterisk in the tables (*). Also, wrong names of the hosts (synonyms, spelling errors) are in parentheses and with asterisk. Generic status of some species is given variously in the literature. For example, Grandiella conula Summers & Schuster, 1979 was transferred to the genus Ennodithiana Haitlinger, 1993. Later this species was transferred to the genus Teklathiana (Haitlinger 1999a). Haitlinger (1993c) provided the justification for transferring this species to the genus Ennodithiana.The correct name is Ennodithiana conula. Hatohylla rosascostai (Lombardini, 1939) described as Coleoglyphus rosascostai was transferred to the genus Hatohylla Haitlinger, 1990. O'Connor (2004 lists him as Grandiella rosascostai. This species certainly does not belong to this genus. This species differs from Grandiella species in the shape of the opisthosoma and the number of caudal setae. Seee also key to the Neotropical genera (Haitlinger 1999a).
Canestrinia pentodontis (Berlese, 1897) has been described as C. dorcicola var. pentodontis. Because there are no drawings of this species and laconic description its species identify requires confirmation.
Male. Propodosomal plate narrow, sejugal suture weakly developed or absent, adanal suckers present, dorsum without ornamentation or ornamented iin median and posterior part of idiosoma, setae d 2 distinctly longer than setae d 1 . Hysterosoma with only 5 pairs of setae. Ventral side of idiosoma with 6 pairs of setae at posterior margin of idiosoma. Setae cG I distinctly longer than setae mG I. Tarsi I-IV very short.
Etymology. The genus was named after archipelago of the Balearic Islands where holotype was collected.
Male. Dorsum with 7 pairs of setae (series c, d, e, f), idiosoma without ornamentation. All these setae short. Moreover, there are two pairs of long caudal setae h 1 and h 2 and long setae cp. Setae d 1 and d 2 shifted near opisthosoma. Opisthosoma short, much narrower than hysterosoma. Sejugal furrow and adanal suckers present. On the ventral side, at the anal opening, there are 4 pairs of setae (p 1-3, f 2 ) and on posterior margin long setae h 3 . Legs I-IV short.
Etymology. The genus name is based on the name of the Island of Java, where the holotype was collected with the addition word nellina.
Remarks. This genus is similar to the genus Haitlingeria. It differs in ornamentation in the form of shell-like and circular structures, setae e 1 and e 2 blunt ended and setae e 1 about three times longer than the setae e 2 (females). Javanellina gen. nov. is also similar to the genus Kuramasenia gen. nov. It differs in the lack of ornamentation and very short opisthosoma (males) (see also keys).
Kuramasenia gen. nov. Canestrinia: Berlese, 1881 (part) Type species. Kuramasenia pictura (Samšiňák, 1971) (Canestrinia pictura) Female. Idiosoma with lines or cell-shaped structures. Hysterosoma the same width as the opisthosoma or only slightly wider. Idiosoma with 6 pairs of setae (series c, d, e). Setae d 2 more than twice as long as setae d 1. Setae h 1 , h 2 and cp long. Posterior margin of idiosoma almost simple or slightly rounded. Sejugal furrow and setae ve present. Tarsi IV longer than the others.
Male. Opisthosoma clearly narrower than the hysterosoma, slightly narrowed in the middle part. Setae ve present. Idiosoma with 6 pairs of dorsal setae (series c, d, e). Setae d 2 longer than setae d 1. Setae e 2 located just at the posterior margin of idiosma. Setae h 1 , h 2 and cp very long, setae h 3 much shorter. Sejugal furrow and adanal suckers present.Tarsi III and IV longer than tarsi I and II.
Etymology. The genus name is based on the name of Mount Kurama where the holotype was found with the addition word senia.

Midiphotia gen. nov.
Neophotia: Samšiňák, 1971 (part) Type species. Midiphotia jureceki (Samšiňák, 1971) (Neophotia jureceki) Female. Hysterosoma and opisthosoma partially ornamented cell-shaped or longitudinal lines. Sejugal furrow well developed. Propodosomal plate absent. Setae ve present. Dorsal surface with 6 pairs of short setae (serie c, d, e). All of these setae of a similar length. Setae h 1 , h 2 and cp very long, setae h 3 short. Tarsi III and IV longer than tarsi I and II. Opisthosoma in the posterior part only a little narrowed.
Male. Idiosoma not ornamented. Opisthosoma very short and narrow. All dorsal setae from series c, d, e very short and smooth. Setae h 1 and cp very long. Setae ve present. 4 pairs of short setae below the genital region. Sejugal furrow and adanal suckers present. Legs IV thicker than legs I-III.
Etymology. The genus name is based on the name of another genus Photia with the addition midi. This is the last part name of the mountain peak Pic du Midi where the holotype was found.
Midiphotia hispanica (Samšiňák, 1971) comb. nov. (Canestrinia hispanica) Remarks. The new genus differs from all European genera in setae d 1 placed near setae e 1 or at least half the distance between setae c 1 and e 1 (females) (see remarks for Cercediphotia) and is similar to Neophotia (males). It differs in the presence of 2 pairs of long caudal setae and the relatively distant position of adanal suckers from the anal opening (see also keys).

Neophotia Samšiňák, 1971
Type species. Neophotia drvotocka, Samšiňák, 1971 Female. Hysterosoma, opisthosoma amd the ventral part of opisthosoma covered by denticles. Sejugal furrow only in the middle of the idiosoma. Propodosomal plate absent. Setae ve present. Setae c 2 at least twice as long as setae c 1 . Setae d 2 missing. Dorsal surface with 5 pairs of short setae (serie c, d, e). Setae h 1 -h 3 and cp long or very long. 7 pairs of short setae below of genital region. Copulatory tube unusually long. Tarsi III and IV significantly longer than tarsi I and II. Opisthosoma in the posterior part strongly narrowed.
Male. Dorsum without ornamentation. All dorsal setae short (series c, d, e). Only setae h 1 and cp are long. Two pairs of setae at anal opening Setae ve present. Incomplete sejugal furrow and adanal suckers present. Legs IV very thick, much thicker than the legs I-III.
Remarks. This genus differs from other genera of presence on the dorsal and ventral parts of the opisthosoma denticles (females); it is similar to the genus Midiphotia (males). See remarks for this genus.

Percanestrinia Berlese, 1880
Type species. Percanestrinia blaptis (Canestrini & Berlese, 1880) (Alloptes blaptis) Female. Sejugal furrow present. All dorsal setae long or semilong. Setae cp, h 1 and h 2 longer than the rest, setae h 3 short. Dorsum with 7 pairs of setae (below sejugal furrow). Setae c 1 shifted beyond the level of setae d 2 and are located right next to each other. Setae se, cp, h 1 and h 2 with or without spikeketes. Below genital region 6 pairs of setae. Tarsi IV clearly longer than the others.
Males. Opisthosoma short, clearly narrower than the hysterosoma, narrower in the middle part. Setae c 1 and others long or semilong placed as in females. Setae cp, h 1 and h 2 very long, setae h 3 short. Sejugal furrow and adanal suckers present. Tarsi I-IV of similar length.
Remarks. This genus is different from all other genera of shifted setae c 1 beyond the level of setae d 2 and setae c 1 located near setae d 1 (females and males) (see also keys). P. norodomi Haitlinger, 1992. Phaleratusellus gen. nov.
Phaleratus: Nesbitt, 1976 (part) Type species. Phaleratusellus alatus (Summers & Schuster, 1982) (Phaleratus alatus) Male. Sejugal furrow and propodosomal plate absent. Adanal suckers represented by a pair of pits resembling alveoli of microsetae. Dorsal surface with different types of setae: very wide flabelliform, long heavy tubular, short and heavily extended and microsetae. Caudal setae with extended bases. Setae vi short, stout, possibly barbed. Setae ve identifiable as a pair of alveoli having no apparent shafts. Three pairs of minute paranal stae and 1 pair of microsetae on ventrolateral walls of opisthosoma. Legs I-IV slender, all similar in size.
A small ventral seta at the tip of each tarsus stout, spurlike, slightly more prominent on tarsi III and IV.
Etymology. The name of the new genus is based on the name of another genus Phaleratus with the addition word ellus.
Remarks. This genus is similar to the genus Passalophagus Nesbitt, 1976and Phaleratus Nesbitt, 1976. Summers & Schuster (1982a describing the species Phaleratus alatus they stated: the assignment of this new species to Phaleratus Nesbitt is a provisional action which may yield a split-upsome future date, when the finding of kindred species shows how to make a better arrangement. However, the differences between these species are large enough to already now that both species belong to different genera. The genus Phaleratusellus differs from the genus Passalophagus in normal, relatively thin legs I-IV vs. legs I-IV short and thick, sclerotized socket-like cup about the coxa absent vs. sclerotized socket-like cup about the coxa present, three pairs of wide flabelliform setae vs. two pairs of wide flabelliform setae; from Phaleratus it differs in 4 kinds of dorsal setae: very wide flabelliform, relatively long with extended bases, short, barbed with widened tops and microsetae vs. long plumose, long tubular with widened base, microsetae; caudal setae with widened bases and pointed vs. caudal setae with widened bases, tubular, blunt ended. Khaustov & Eidelberg, 2001) Pseudocanestrinia. damiani (Haitlinger, 1989) comb. nov. (C. damiani), P. roberti (Haitlinger, 1990), comb. nov.
Male. Idiosoma oval bears 8-10 of sort dorsal setae according three long caudal setae, located on ventral side or on the posterior margin Sejugal furrow and propodosomal plate absent, adanal suckers present, dorsum without ornamentation. Opisthosoma narrow, extended posteriorly. Posterior margin of opisthosoma almost straight or with appendix. Tarsi 26

R. Haitlinger
III and IV without ambulacra. Ventral part below adanal suckers to 12 short and nude setae. Female. Dorsum is covered with cell-shaped structures. All setae from the c, d, e series are short and of similar length. The setae h 1 , h 2 and h 3 are located on ventral side. Setae p 1 and p 3 are longer than the other setae of the p and ad series.
Etymology. The genus name is based on the name of Island Sumatra where the holotype was collected with addition word nella.
Remarks. The new genus is similar to the genera Gioharattia, Jullongia and Teophyssa. It different from them in: almost the entire of dorsal side of idiosoma is ornamented, setae d and e on tarsi III and IV are very long (females). The new genus Sumatranella differs from all other genera by the lack of ambulacra on tarsi III and IV (males) (see also keys).

Family Blattidae Latreille, 1810
This family contains 3 subfamilies. Only one species of canestriniids was found in subfamily Blattinae Latreille, 1810

Subfamily Blattinae Latreille, 1810
This subfamily contains 23 genera. Only one species Coleopterophagus blattophagus Banks, 1914 was found on Periplaneta americana (Linnaeus, 1758) in Panama (Banks 1914). It is an accidental host, and the species described requires generic revision.

Family Carabidae Latreille, 1802
This family contains 28 subfamilies with about 40,000 species. Canestriniid mites were stated only on species belonging to 2 subfamilies: Carabinae and Harpalinae.46 species of canestriniids from 18 genera were found on 78 host species belonging to 7 genera.

Subfamily Harpalinae Bonelli, 1810
This subfamily contains about 20,000 species. Only 4 species of canestriniids belonging to 4 genera were found on 4 species of hosts from 3 genera May be Megacanestrinia mucronata Trrägårdh, 1906 collected on Carabidae undet. is also associated with Harpalinae (Trägårdh (1906). The species of Harpalinae are exceptionally hosts of canstriniids. Globosophotia bettinae Haitlinger, 1991 was collected from Pterostichus globosus (Fabricius, 1792) in Morocco (Haitlinger 1991f) (Table). It is the only species collected from the genus Pterostichus Bonelli, 1810 which contains more than 1100 species. All 4 mite species were collected in Africa (Table I).

Family Cerambycidae Latreille, 1802
The large family consist about 10 subfamilies, about 428 genera and over 26,000 species. Canestriniidae were collected from species of two subfamilies: Cerambycinae and Lamiinae. Canestriniidae rarely occur on Cerambycidae. So far, 16 species of canestriniid mites beconging to 7 genera have been found on 17 species and 8 genera of Cerambycidae (Tables  XII and Tables XIV). The relationships of canestriniids and cerambycids and their distribution are very A review of host-commensal associations between canestriniid mites (Astigmata: Canestriniidae) and Insecta 29 poorly known. Only 2 species related to cerambycids are known from Africa, 2 from America, 6 from Asia and one from Europe (Summers & Schuster 1982b;Haitlinger 1989dHaitlinger , 1995b (Tables I, III, V, VII  and XIV).

Subfamily Cerambycinae Latreille, 1802
The subfamily includes about 3900 species. Only 3 canestriniid species were collected from 4 species of hosts: Dicanestrinia cerambycis (Canestrini, 1878) (Table  III). On the basis of current research, it can be concluded that Cerambycinae are very rare hosts of canestriniid mites.

Subfamily Lamiinae Latreille, 1825
The subfamily contains over 3052 genera and over 29,258 species (Saha et al. 2013) but 9 canestriniid species were collected only from 14 host species belonging to 5 genera (Summers & Schuster 1982b;Vishnupriya & Mohanasundaram 1988;Haitlinger 1989dHaitlinger , 1995b. Most species (10) belongs to the genus Batocera Castelnau, 1840 (which contains 54 species); 6 species of canestriniids collected from them. They belong to 2 genera: Caestriniella Berlese, 1910 and Sumatranella gen. nov. C. togata Berlese, 1910 was collected from Batocera lineolata Chevrolat, 1852 from Java. Summers & Schuster (1982b) quote it after him, adding a possible second name for this species C. liniolata Chev. Both of these names are wrong. The correct name seems to be B. lineolata. This species was known from China, Japan, Korea and Taiwan and most recently from India (Boyane et al. 2020). Also, Batocera proserpina mentioned by Summers & Schuster (1982b) has the correct name B. wallacei proserpina Thomson, 1865. Probably most of canestriniids found on Lamiinae are associated with only one host. The exception is C. togata found on 3 hosts, S. elae (Haitlinger, 1989) and C. physsana Haitlinger, 1989 found on 2 hosts (Tables VIII and X). Caanestriniids from Lamiinae were collected from Africa (Kenya, Tanzania -2 species), Central America (Panama -3 species of the genus Thopia Summers & Schuster, 1982 from 3 species of hosts) and Asia (India -1 species, Indonesia -5 species, Malaysia 1 species, 4 from the genus Canestriniella and 3 from the genus Sumatranella collected from 10 species of the genus Batocera Castelnau, 1840 and 1 species of the genus Clyster Arrow, 1908 (Tables XII and XIV).

Family Chrysomelidae Latreille, 1802
This family contains 11 subfamilies and about 35,000 species but canestriniids were obtained mainly from Cassidinae, less from Chrysomelinae and exceptionally from Eumolpinae. 85 species of canestriniids and 3 unidentified species from 23 genera were found on 153 species and 8 unidentified species of chrysomelids of 39 genera (Table X).
?? Of the 159 species of Chrysomelidae, 84 species of canestriniids were collected

Family Elateridae Leach, 1815
Elateridae represents one of largest families within Coleoptera and contains about 10,000 species and 19 subfamilies. Canestriniidas were collected only from species belonging to subfamily Semiotinae.

Family Geotrupidae Latreille, 1802
The family has about 340 species in about 50 genera in two subfamilies: Geotrupinae and Taurocerastinae. Only one species of Canestriniidae was collected from Geotrupinae.

Family Scarabaeidae Latreille, 1802
This family contains 19 subfamilies and about 30,000 species. 70 species of canestriniids belonging to 25 genera were found on only 66 host species, 5 determined to genus and from unidentified Cetoniinae and Dynastinae belonging to 3 subfamilies and 42 genera (Tables XI and Tables XIV). The relationship between canestriniids and scarabaeids and their distribution is very poorly known. 42 of canestriniid species are known from Africa, 21 from Asia, 7 from Europe and 8 species from Papua New Guinea.
In the original description of Boetophela ginae Haitlinger, 1989 it was stated that this species was caught in the Sumatra on the Batocera rubus (Linnaeus, 1758) (Cerambycidae) (Haitlinger 1989d). It was based on a wrong label (collected at the museum). This species and all others of this genus are found only in Africa (Haitlinger 1993a).

Subfamily Dynastinae MacLeay, 181
This subfamily contains over 1500 species and 225 genera. 20 canestriniid species from 8 genera were 50 R. Haitlinger collected from 12 dynastid species belonging to 7 genera (Tables XI and XIV) (Haitlinger 1989b, Khaustov & Eidelberg Khaustov & Eidelberg 2001Trach 2006). 3 canestriniid species collected from 2 host species were found in Africa, 9 canestriniid species collected from 8 host species were found in Asia, 2 canestriniid species collected from 2 host species belonging to one genus were collected in Europe and 7 canestriniid species from 3 genera collected from one identified host species and a few unidentified hosts.

Subfamily Pimeliinae Latreille, 1802
The subfamily contains 223 genera and many species. Only 2 dpecies: Boleohylla kiogana Haitlinger, 1991 collected from Psammodes carbonarius Gerstaecker, 1954 in Uganda and Canestrinia remigans Berlese, 1910 collected on Morica planata (Fabricius, 1801) in an unknown place in Africa (Haitlinger (1991f) (Tables I and XIV). Possibly C. remigans belongs to a different genus because species of the genus Canestrinia were not found in Africa and also not found on Tenebrionidae.

Family Glomeridae Leach, 1815
Extensive family with 47 genera. Only one species Diplopodocoptes transkeiensis Fain, 1987 was found from an unidentified host in Republic of South Africa (Fain 1987b) (Tables XIII and XIV).

Family Odontopygidae Attems, 1909
Extensive family with about 55 genera. Only one species Diplopodocoptes transkeiensis was found from an unidentified host in Kenya (Fain 1987b) (Tables  XIII and XIV).
So far only one species of Canestriniidae has been found on Diplopoda from Africa. The relationships between canestriniids and Diplopoda are unclear and requires further research.

Distribution
Canestriniidae occur in all regions of the world, with the exception of their southern and northern ends. In Europe the most northerly were found in Estonia, South Norway (Coleopterophagus albini Haitlinger, 1990 collected from Protaetia metallica Herbst, 1782) and South Sweden [Procericola bourgognei (Oudemans, 1923)] from Carabus coriaceus Linnaeus, 1758] (Lundquist 1985;Haitlinger 2015). In Asia the most northerly were found in Russia: Photia lopatini Khaustov & Eidelberg, 2001 in Sachalina, collected from C. lopatini, P. sibirica Khaustov & Eidelberg, 2001 in Novosibirsk prov., collected from C. schoenherri, Uriophela arieli Haitlinger, 1991 in Amur prov. collected from Hemisodorcus rubrofemoratus and P. polymorpha Samšiňák, 1971 from Japan in Hokkaido, collected from C. gehini and Russia in Vladivostok and vicinity of Ussuri, collected from C.imperialis, C. schoenherri, C. schrenki and C. vestinghoffi. P. polymorpha has been collected from many species of hosts. According to Khaustov and Eidelberg (2001), it could be a collective species. In America the most northerly canestriniids were found in Mexico (in Alaska, Canada and the USA, they have not been found so far); furthest in the south of America they were found in northern Argentina (Haitlinger 1992b).
In Africa, 68 species of canestrinids belonging to 32 genera were found. Most of the canestriniid genera are represented only in Africa. Only Athogavia Haitlinger, 1989 is also known from Asia, Camirohylla Haitlinger, 1991 is known from Europe and Iran, Canestrinia Berlese, 1881, Mesophotia Samšiňák, 1971and Ibizella Haitlinger & Šundić, 2016 are known also from Europe, Coleopterophagus Berlese, 1882 Ismailienia gen. nov. and Percanestrinia Berlese, 1911 are known from Europe or Asia. Canestriniidae were found in only 20 countries so far, from known 59 countries (including France -Majotte, Reunion, Great Britain -Saint Helene, Portugal -Madeira and Spain -Canary Islands). All mite species have been found in only one country except Cetonicola orestesi Hatlinger, 1991 (Ghana, Liberia), and Diplopodocoptes transkeiensis Fain, 1987 (Kenya, Republic of South Africa) ( Table  I). Most species are described from Tanzania (13), Democratic Republic of Congo (8) and Cameroon (7). Canestriniidae were collected from 44 host species belonging to 5 families and 33 genera (Tables I and II); 4 host species identified to genus and 12 identified to family. From 30 host species, only one canestriniid  (Table I). 28 species of canestriniids belonging to 13 genera collected from 29 species of hosts belonging to 3 families and 19 genera have been found in Central America. Most species belong to the genus Bibulothiana Haitlinger, 1993 (4) and 3 species each belong to the genera Ennodithiana Haitlinger, 1993, Phriknodora Summers & Schuster, 1982and Thopia Summers & Schuster, 1982 in the remaining 6 genera 1 or 2 mites species were found. 21 host species belong to the Cassidinae subfamily (Chrysomelidae), one species belongs to the family Elateridae, 3 species to the subfamily Cerambycinae and 3 species to the subfamily Lamiinae (Cerambycidae). Canestriniidae were found in 9 countries, from known 20 countries. Most species were found in Panama (14), Mexico (9) and Costa Rica (7). All canestriniid species were collected from one or two host species with the exception of Apalotacarus alutaceus (Turk, 1948) and Grandiellopsis gambosa (Summers & Schuster, 1979) found on 3 hosts (Table II).

Host specifity at the generic level
Carabidae 12 genera of canestriniids were found in Carabinae. 8 genera are associated with separate host genera: Paraphotia with Calosoma (C. auropunctatum is the only known host of this genus, from about 167 known species), Cercediphotia, Kuramasenia, Mesophotia, Midiphotia, Neophotia, Paracanestrinia, Percanestriniella, Photia and Procericola are associated with Carabus only. P. howdeni, collected from Passalidae, is also included in the genus Photia but its inclusion in this genus is unreasonable and needs to be corrected. Genera Canestrinia and Dicanestrinia are also associated with other genera of hosts, including other families (Cerambycidae, Scarabaeidae, Tenebrionidae). 4 genera of canestriniids were found in Harpalinae. The genus Globosophotia is known only from genus Pterostichus, Megacanestrinia is known only from genus Tefflus (M. mucronata described from unidentified Carabidae probably belonging to the genus Tefflus). The genus Coleoglyphus has been found of different genera of hosts.
Cerambycidaee 2 genera amd 3 species of canestriniids were found in Cerambycinae. The genus Phriknodora is known only from genera Juiaparus and Trachyderes. Dicanestrinia has also been found on Carabidae. The typical species Dicanestrinia cerambycis is described from Cerambyx cerdo. It is the only Canestriniidae species found on the genus Cerambyx among 14 species of Cerambyx. 4 genera and 11 species of canestriniids were found in Lamiinae. The genera Canestriniella and Sumatranella are associated exclusively with the genus Batocera. Anaspistes is associated with Sternotomis and only Thopia was found on Lgocheirus, Neoptychodes, Taeniotes and Cassidinae. and Thopia (2 species from 5) with Polychalca (Polychalca punctatissima common with Grandiellopsis).In the remaining 7 genera this specificity is absent. However, each of these genera has been found outside of also common hosts also species of the genera found only in them. Arrunsithiana are associated with 6 genera hosts, exclusi ely collected from Mesomphalia. The highest number of host species among Chrysomelidae was found (34) including 28 among Stolas (many common to Grandiella and Bibulothiana). Bibulothiana are associated with 4 genera hosts but only they were found on Chelymorpha and Microtenochira, on Crambelea and Omaspides only Ennodithiana was found. Grandiella was found on 10 host genera but exclusively on Anacassis (it is also a host for Olafithiana, but a different species), Botanochara, Eumolpus, Goniochenia and Orexita, Grandiellopsis was collected from 5 host genera, on Carlobruchia and Discomorpha (15 species) (D. conspersiprennis was also found on Ovarrothiana) no other canestriniids were found. Olafithiana was fiund on 4 host genera, for Elytrosphaera and Helocassis only. Ovarrothiana was found on 5 host genera; from Agenysa and Miocalaspis collected exclusively and from Eugenysa including Arrrunsithiana (two hosts are common with Arrunsithiana). 77 canestriniid species were collected drom 149 chrysomeliid species. 122 species of Chrysomelidaae are hosts only for one species of Canestriniidae. 22 species of Chrysomelidae are hosts of 2 species of Canestriniidae and 5 species are hosts for 3 to 6 species of Canestriniidae (Table VI). Only 5 species were collected from a larger host species: Polychalca punctatissima (3), Stolas lebasii (3), Ecchoma clypeata (4), Stolas conspersa (4) and S. chalybaea (6) ( Table VI). The greatest number of host species was found for Arrunsithiana battosi (15) and Grandiellopsis gambosa (15). Also, from many hosts were collected Grandiella tacjannae (13), Olafithiana foliacea (12), Grandiella tetracaudata (9) and Bibulothiana brevispinata (6).

Elateridae
Only one genus of canestriniids is associated with the subfamily Dengrometrinae (Elateridae) 4 Danaithiana species were found on 3 host species of the genus Semiotus.
Lucanidae 7 genera of canestriniids were found in Lucaninae. The genus Noemiphela is exclusively associated with genus Cladognathus. Genera Cooremania, Haitlingeria and Rugoniphela are associated with genus Dorcus. 3 other genera found on Dorcus: Canestrinia, Coleopterophagus and Sandrophela are associated with a few other genera. The genus Uriophela is exclusively associated with Hemisodorcus, Sandrophela is associated with Dorcus, Hexathrius and Odontolabis, Amboinophela is exclusively associated with Proposocoilus. Coleopterophagus was found on Lucanus but is mainly associated with the genus Protaetia (Cetoniinae). 3 genera canestriniids: Canestrinia, Sandrophela and Vereoxia are associated with the genus Odontolabis but only Vereoxia is exclusively associated with this genus.

Passalidae
Knowledge of the host species of Passalidae is very limited. As many as 28 species of canestriniids were collected from unidentified Passalidae. 10 genera of canestriniids were found in 11 genera of Macrolininae.
Cnecoderma Dicronorhina and Smaragdesthes. The rest of the genera have wide relationships with the hosts. Gioharattia (associated with 4 genera of hosts), Paraphagella (5), Albinorattia (6), Boetophela (6), Cetonicola (7). 7 genera of canestriniids are associated with 7 genera of Dynastinae. 5 genera of canestriniids are associated with only one genus of hosts. Hargeinella and Vandeinella are associated with Trichogomphus, Canestrinia with Petodon, Canestriniella is aassociated with Clyster but it is probably accidental host and Oceogavia is associated with Papuana. Athogavia is associated with Oryctes and Trichogomphus but the main hosts are species of the genus Oryctes. Aurilossongia was found on Chalcosoma and Xelotrupes but the main hosts are species of the genus Xelotrupes. 2 genera of canestriniids are associated with 2 genera of Scarabaeinae. Scarabiphotia is associated with undetermined Scarabaeinae and Coleoglyphus is associated with Copris and Heliocopris.
Tenebrionidae 14 genera of canestrinnids were found in Tenebrionidae. 3 genera are associated with 3 genera of hosts in Lamiinae. Saniothiana is exclusively associated with Odontopezus, Boleohylla was found on Metallonotus, Donnelafontia is assocted with Calostegia and Odontopezus.
2 genera canestriniids were found on 2 genera of hosts in the subfamily Pimeliinae. Canestrinia is associated with Morica and Boleohylla is associated with Psammodes. The 8 genera of canestriniids are associated with the 5 genera of Tenebrioninae. Camirohylla is associated with Blaps and Dila, Balearella is associated with Blaps and Gnaptor, Ciprusenia and Ismalienia with Blaps. Hatohylla was found on Nyctobates and Percansetrinia is mainly associated with Blaps but was collected also from Carabinae.

Discussion
Canestriniidae is one of the least known families of mites among Astigmata. 370 species and 101 genera collected from 361 species of hosts (113 genera) have been described so far. Knowledge of the Canestriniidae fauna on different continents is very varied. In Africa, only 66 species from 31 genera were found on 44 host species. 28 species from 13 genera collected on 30 species of hosts were found in Central America, 78 species from 17 genera collected on 119 species of hosts were found in South America, 109 species from 34 genera collected on 120 species of hosts were found in Asia, 49 species from 20 genera collected on 62 species of hosts were found in Europe and 33 species from 12 genera collected on 15 species of hosts were found in Oceania. Should be regarded as the best known a small area of Europe. The Canestriniidae are the least known in Africa, but they are also very poorly researched elsewhere in the world. For example, canestriniids have not yet been found on Cassidinae in Africa, Asia and Oceania. But 138 Cassidinae species are known from Africa, 157 species are known from Asia and 16 species are known from Australia and Papua New Guinea (Borowiec 2009(Borowiec , 2010 (14), Poland (14), Ecuador (13), India (13) and Tanzania (13). Considering the size of the diversity and number of potential hosts as relatively best researched can be considered Panama, Ukraine, Italy and Poland. Canestriniidae were collected from 12 insect families: Blattidae, Carabidae, Cerambycidae, Chrysomelidae, Elateridae, Geotrupidae, Lucanidae, Passalidae, Passandridae, Scarabaeidae, Tenebrionidae and Zopheridae. In addition, one species has been described from the Diplopoda. The presence Ciprusenia sellnicki on Typhoeus typhoeus (Geotrupidae) is certainly coincidental. Also, the presence of Coleopterophagus blattophagus on Blattidae must be considered as accidental. Whether species belonging to Passandridae and Zoopheridae are stable hosts for canestriniids require confirmation. In Elateridae, only some species from the genus Semiotus are hosts for the 60 R. Haitlinger canestriniids. In other insect families many subfamilies have been described. However, Canestriniidae have been found in one to four subfamilies, but usually in one subfamily. Only in Scarabaeidae in the subfamilies Cetoniinae and Dynastinae a similar number of described species of canestriniids was found. In Canestriniidae at the generic level and species level exist clear host specificity. About 58% of the canestriniid genera are associated with one genus of host and about 60% of canestriniid species are associated with one host species.However, there are canestriniid species with very limited host specificity collected from many hosts: Arrunsitiana battosi collected from 15 host species, Grandiellopsis gambosa (15), Grandiella tacjanne (12), Olafithiana foliacea (12), Photia polymorpha (12), A. quadrata (11), Grandiella tetracaudata (10), P. procustidis (8), Albinorattia igori (7), Procericola bourgognei (7), Canistra osculatii (6), Gioharattia maharae (6), Percanestriniella levis (6), Photia chrysocarabi (6), Stolas chalybaea (6).
The host relationships of many species and genera of mites have been discussed in detail above. Some species of mites mentioned in the text and tables require generic revision. There are problems with some canestriniid genera. Of the 23 described species of the genus Canestrinia 6 were transferred to the genus Pseudocanestrinia, 5 to the genus Kuramasenia one to the genus Pseudocanestriniella and one to the genus Balearella, one to the genus Javanellina (Khaustov & Eidelberg 2001;Haiitlinger and  Šundić 2016, this paper). Of the 17 species listed in Table XIV, 5 species require generic revision: C. kacperi, C. manicata, C. microdisca, C. nepalensis and C. remigans. According to Khaustov and Eidelbeg members of the genus Canestrinia are associated with lucanid beetles and scarabaeid beetles of the genus Pentodon. However, 2 species associated with the genus Pentodon have now been transferred to the genus Pseudocanestrinia (P. damiani, P. roberti). Moreover, C. microdisca, C. nepalensis and Coleopterophagus berlesei are associated with Lucanidae. However, it is currently impossible to establish their generic identity due of the laconic description and no drawings. C. manicata caught in Brazil on Chrysomelidae and C. remigans caught in Africa on Tenebrionidae. Both of these species probably do not belong to Canestrinia. Hosts have not been established for C. javensis. It remains to be decided whether they belong to the genus Canestrinia and that indeed no species of Cnestrinia is associated with Carabidae. Also belonging to the genus Coleopterophagus C. blattophagus collected from Blattidae in Panama, C. pulcher and Photia howdeni are unbelievable. The presence of canestriniids on Blattidae is certainly accidental. In addition, species of the genus Coleopterophagus in America they do not occur (species whose genus is questionable are marked with an asterisk in the tables). Based on current knowledge, the main hosts of Canestriniidae should be considered Cassidinae (Chrysomelidae) (42 described canestriniid species were collected from143 species of hosts), Cetoniinae (Scarabaeidae) (48 vs. 50), Carabinae (42 vs.74), Macroliniinae (Passalidae) (35 vs. 18) and Dynastinae (Scarabaeidae) (20 vs. 12).   * Percanestrinia norodomi has also 2 lobes, but genu III with short and enlarged seta ** based on D. huberti and D. knobi, ***based on the description of ..... ......C. dorcicola; Haitlinger (1988c) A key to the European genera of canestriniid mites for females In genera Camirohylla Haitlinger, 1991 and Lombardiniella;Cooreman, 1950  A key to the Asian and Oceanic genera of canestriniid mites for males Genera which are already in the keys for Europe are not included in these keys. In genera: Cnecoderma Summers & Schuster, 1981, Hypopteryx Summers & Schuster, 1981, Noemiphela Haitlinger, 1991, Paramelisia Summers & Schuster, 1982, Shillongia Haitlinger, 1990and Taseopus Summers & Schuster, 1982 Haitlinger, 1990 A key to Asiatic and Oceanic genera of canestriniid mites for females In genera: Hargeinella Haitlinger, 1989, Phaleratosellus gen. n., Sajanophela Haitlinger, 1990, Shillongia Haitlinger, 1990, Uriophela Haitlinger, 1991 Haitlinger, 1989