Association of uropodid, prodinychid, polyaspidid, antenno- phorid, sejid, microgynid, and zerconid with ants

PEKKA T. LEHT]NF,N

Lehtinen. Pekka T.: Association of uropodid, prodinvchid, prolyaspidid, antennophorid, sejid. microgynid, and zerconid mitcs with ants. [Myrgaster inom kvatsterfamil jema Uropodidae, Prodinychidae, Polyaspididae. , Sejidae, Microgynidae och Zcrconidac.]- Ent. Tidskr. 108: 13-20. Unei, Sweden 1987. ISSN 0013-886x,

The obligatorily myrmecophilous fauna of North Europe include one species of. Anten- nophorus (Antennophoridae) and about 20 species of Uropodidae, representing the sub- families Oplitinae ( Oplitis and Urodiscel/a), Trachyuropo dinae (Urotrachytes and U rojanetia), and a few species in other subfamilies (Phaulodinychus in Uropodinaeland Uroobovella, Oodinychtts, ar,d Trematurella in Trematurinae). Polyaspidid taxa are represented only by deviating populatbns of Dipolyaspis trsracezs. while various generally non-myrmecophilous prodinychids have invaded ant nests locally (Prodinychus flagelliger, Dinychus carinatus, D. arcuutut). and in some case formed huge populations (Trachyxenura pyriformis, Dinychus septentionalit). Many zerconids are numerous in ant nests, but only Prozercon traegardhi anda new species from SW Finland show a distinct preference. Morphological adaptations concerning mouthparts and larval design for ant symbiosis are found in Antennophoridae and in the uropodid subfamilies Oplitinae and Trachyuropodinae. These two subfamilies share only a few parallel adaptations, and they have evolved from differ- ent uropodid groups. Most myrmccophilous species of Nonh Europe prefer a single ant host. Nests of Formicinae, esp. Ltr;ius (Chthonolastus, Cqutolasius & s.str.), Dendrolasius, Csmponotus, and Formica are distinctly preferred to Myrmicinae (except Tetamorium\. Both ants and mites have seemingly isolated populations, which could be classified as species in stalu nascendi. Many of these mite populations with different ant hosts are also allopatric ard here trcatcd as subspecies. All European species of the Oplitis ovalala-group live in myr- micine nests ( Ierramoium, Messor, and Myrmica). A detailed analysis of the of thc uropodid guests of different populations of Formica er.recra most probably will help in the taxonomic reevaluation of the latter. The names used in this paper are based on a recent revision of these groups, including also the checking of all Berlese types.

Pekko T. Lehtinen, Zoological Museum, Department of Biology, University of Turku, S F-205 00 T urk u, Finland.

Introduction Canestrini & R. Canestrini (1882), G. Canestrini The presence of a ich mite fauna in the nests of & Berlese (1884), G. Canestrini (1884), Michael various European ants was wcll documented by (1891 & 1894 a & b), Karpelles (1891), Moniez numerous authors ca. 100 years ago and first men- (1892 & 1894), Wasmann (1894, 1897 a-b, 1898, tioned by Forel (1874) without identification of I tt99 & 1902), Leonardi ( 1895 & 1896), Trouessart the mitcs. Michael (1894b) and Wasmann (1899) (1U96 & 1902), Janet (ltt97 a & b), Karawaicw suggcsted that many of these mites are dependent (1906), and Kneissl (1907, 1908). on certain ant species or at least strongly prefer Although myrmecophilous species of mites their nests as the microhabitat. wcre described by many acarologists, the main Basic data for myrmecophilous Uropodoidea opus is Berlese's Mirmecophili (1904b), at was presented by Haller (1877 & 1882), Lubbock least as far as the Uropodina is concerned - the (ltt8l ). Berlese (1881, 1882-1892, 1904 a & b), G. main group discussed here. Later, Donisthorpe 14 Pekka T. Lehinen

(1927: British Isles), Balogh (1938: Hungary), single stone or any other limited area may include Storkan (1940) and Pecina (1980: Czecho- small nests of many Myrmica, Leptothorax, and slovakia), Greim (1952: S Germany) and Wis- lasius spp. In this way, thc guests of different ant nicwski (1979c: Poland), have published locallists species may become mixed in the sample. of myrmecophilous mites. North European myr- Secondly. many authors have listed the host mecophilous uropodids and polyaspidids havc species of ditferent mites without any quantitative been recentlv described by Greim (in Hirschmann information. Let's take an illuminating examplc. 1957), Hirschmann & Zirngicbl-Nicol (1961), If we have an old large anthill of Formica exsecta. Wisniewski (1979a. 1980a & b), Pecina (191t0), throughout crowded with the very striking, nicc and Hirschmann (1984). Krasinskaya (1961) red uropodid Urojunetia coccinea (500-15(X) speci- studied the tife historics of some of the species dis- mens per liter), it is likcly that single specimens cussed here. are accidentally transported by the ants to the The European uropodids have not been neighbourhood. When we then find single speci- taxonomically revised so far. Names used here are mens of U. coccinea within the closeby anthill of based on a recent. as yet unpublished revision that Formica aquilonia or under the stones. where Ior- also includes checking of all Berlese types of thc mica ftt^sca or Lasiu.c riger happen to have their mite groups discussed here. ncsts. these observations don't prove that the The published information about North Euro- species belong to the local host species of U. coc- pean myrmecophilous mites is scanty, and the cinea. Third, dead specimens of many majority of the common widespread species were species are actively transported to the nest hy the never reported from any of the North European ants. countries. Thor (1900) dcscribed a new species, Uropotla from Norway. It has not formicarum Dispersal and feeding of myrmecophilous mites bccn cited by Central European authors and it is hcrc synomymize d with Oodinychus ovalis (C . L. The mode of living and dispersal of myrmecophil- Koch). Triigirdh (1942) described a new uropodid ous mites is realized according to three differcnt species. Trematttrella stylifera, and stated (1945) strategies. that it is associated with a species of the Formica 1. Phoretic dispersal of adult specimens, Adult ruft-group from Sweden. Hc also proposed a new specimens of Oplitis and Urodiscella are regularly family, Trematurcllidac (1944), for the species. attached to thc protibial comb of their host ant, TrigArdh (1943) also presented a review, where both workers and alates. This behaviour was first thc family Antennophoridae was discussed in dcpicted by Janet (1897a). Adult specimens of morc detail, however, with most examples from are regularly attached below or on tropical and other non-European genera. Hc the head of the ants, and are dispersed in this way. mentioned the phoretic dispersal of uropodid Both the oplitine species and Antennophorus deutonymphs, but exemplified from other insects. make use of this close attachment for their feeding mainly Coleoptera. No mesostigmatid mite from and are more generally classified as commensals. n€sts is previously known from Finland. What they eat is subject to opinions, but Oplitis ^nt "I r.c cvaluation of thc descriptions of the associ- spp. are usually regardcd mainly as feeding of the ations between different species of mites with the minute skin particlcs and other organic debris ant spccies were affected by the obscurity of both combed by their brushlike, strongly specialized mitc and ant taxonomy at the time of the pionccrs mouth parts. of this field (Leonardi 1896, Wasmann 1t199, 2. Phoretic dispercal by deulonymphs. Many Donisthorpe 1927). Thc current specific concepts adults of myrmecophilous and non-myrmecophil- of the Formica rufa grorp date back only to Yar- ous species of the uropodid subfamilies Uro- row (1955), and, in my opinion, there are still podinae and Trematurinae and the nominate many unsolved problems. The yellow species of polyaspidid subfamily are never attached to in- /,asius have quite often been misidentified, not sects. Most of them, however, have a phoretic only by acarologists, but also by myrmecologists. deutonymph that is firmly attached to a flying in- There are also two additional sources of error in sect to secure an effective dispersal. the published data about the association between Phaulodinychus hamulifer, the oddJooking guest mite and ant specics. First, a sample from under a of Lasius niger, throughout covered by club- Association of mites with ants 15 shaped strong setae, can easily be collected by Synocoets were originatly defined as guests tol- sweep-netting at the swarming of its host. Work- erated by ants, while symphiles as guests actively ers. most probably. play no role in the dispersal of and friendly treated by ants. Much additional data this species. The mode of dispersal of the myr- about reciprocal relations of different arthropod mecophilous populations of Dipolyaspis testaceus groups have been accumulated since Wasmann's and Oodinltchus spatuliferus has not been ob- time, and the relationships are best described in served, but most probably their deutonymphs arc general terms of cocvolution. phoretic on swarming Camponotus herculeanus. The term myrmecophilous is now used as a gen- These specialized populations most Iikely repre- eral term for organisms distinctly preferring ant sent evolving new taxa, although no undisputed nests, but this term doesn't describe thc mode of morphological differences are present. association in detail. All authors that have made Manv species of these subfamilies have two exact observations about the habits of mites in ant types of deutonymphs, phoretic and sessile. The nests, have regarded most ofthem as commensals. common uropodid species of various special habi- Our knowledge today is not sufficient for the final tats. Oodittychus ovalis, uses various insects for classification of symbionts and pure commensals dispersal by its phoretic deutonymphs, but sessilc among the myrmccophilous mites, but most prob- deutonvmphs are often found in large numbers to- ably all groups of mites that are obligatory myr- gether with adults and protonymphs in a suitable mecophilous (Antennophorus, Oplitis, Urotlis- h abita t. cella, Urotrachytes, Urojanetia and Phaul- A special group consists of the myrmecophilous odinychus (Uropolyaspis) spp., Oodinychus species ofthc genus Uroseius. Adults ofthis genus spatuliferus & Uroobovella obovata) are sym- are rarely collected on mammal carcasses, while bionts. A part of the facultative myrmccophilous phoretic deutonymphs are relatively common in species may also belong to this category. some of the non-myrmecophilous species. For All zerconids, preferring to live in or regularly Uroseius koe hleri Wisniewski, 1979, onlv a few prescnt in ant nests live there probably simply be- deutonymphs are known from an anthill of For- cause these microhabitats are rich in organic deb- mica polyctena in Poland and from two anthills of ris or afford an optimal microclimate for hiberna- F. rufa in southern Finland. tion. Many of thesc mitcs most probably are fungi- 3. Neither adults nor deutonymps of ohligatory vores as many other debris living ones. Hyphae of myrmecophilous groups are regularly attached to numerous species of fungi are regularly present in anls. Uroobovella obovata is an example of this ant nests, and they are most probably as important strategy, other can be found in the subfamily to the myrmecophilous mites as the ants them- Trachyuropodinae. selves are. Species of Urotrachytes, Urojanetia, and Uroobovella obovata are often found in huge The mite fauna of difrerent ant species numbers within ant nests! whilc solitary specimens are found in the surroundings outside the nests. Most Finnish species of myrmecophilous mites are Most probably they are accidentally attached to associated with only one or two host species (Tab. the workers of ants. and this is sufficient for their 1). Their more detailed distribution will be discus- dispersal. The same is true for all zerconid, sejid, scd elsewhere (Lehtinen. in preparation). Old ob- and prodinychid species. present in ant nests. servations from othcr European countries were summarized without revision of neither ant nor mite species by Bernard (1968). These data cannot Categories of myrmecophilous mites be compared with the data in Tab. 1. Wasmann (1894) first classificd myrmecophilous Members of Lasius s.lat. have the highest into synoects, synocoets, symphiles number of mites associated with a single species of (: m)'rmecoxenes), and, ectoparasires. These ant. In addition. the genus Antennophorus with categories wcrc discussed in detail by Donisthorpe five European specics is completely restricted to (1927: xv-xxiii). The first and last categories are nests of Lcsius, and the oplitine generaOplitis and not represented in the mite groups discussed here, Urodiscella are nearly so restricted. Thc nests of although uropodids as parasites wcre still listed by Lasius (Chthonolasius) are quite difficult to find, Sellnick (1939). as they are 3M0 cm deep in the ground (L. mixtus 16 Pekku T. Lehtinen

Tab. 1. Checklist of Finnish myrmccophilous mites; , Uropodina, Sejina & Zerconina. The column A givcs the type of relationship as: (l) dependent on onc/two closely related ant speices, (2) abundant in ant nests, but several common hosts, (3) occasionallv very abundant in anthills. (4) myrmecophilous populations

Mite species Host species

Aniennophorina Camin & Gorirossi. 1955 Anlennophoridae Berlese, 1892 A nlerutop ho rus pubescerrs Wasmann. 1897 Lasius umbralus Uropodina Kramer. 1881 Uropodidee Kramer. 1881 Oplitinae Hirschmann & Zirngiebl-Nicol, 1962 Oplitis pandata (Michael, 1894) Lasius niger O. pandaru (Michael. 1894)ssp. n. "A" Formica rula (inland), f. e.rrccla (inland bogs) O. pandam (Michael. 1894) ssp. n. "B" Camponoltu herculeanas (SW) O. uillosel/a (Berlese- 1904) Lasius llavus (Ostrobothnia) O. villosella ssp. n. Lasius meridionalis O. ouarr./a (tserlese. 1903) Teiramodum caespitum O. stamncriGretm in Hirschmann 1957 Dendrolasius fu liginosus Urodisce llu ( U ro pectinis) philocten o (lanet. I 897) Lasiusmixtus U. (Uropectiniu) wasm anni (Kneissl, 1907) Lasiusflavus (S Finland) U. (Urodiscellu) uloplzora (Berlese, 1903) Dendrolasius fuliginosus & Lasius mixlus Trachyuropodinae Berlese, l9l7 Urotrochytes lbtmicaria ( Lubbock. l88l ) Lasiusflavus Uroj ane tiu coccinea (Nlichael, 1891 ) Fomicr exsecta (meadows & archipelago), F. sazguinea, [;. (Servi- formica) spp . U. excavata (Wasmann, 1899) Telr8morium caespitum U. wasmanniana (Berlese. 1903) Lasius mixtus U. hirschrnanni (Pecina, 1980) Lasius meridionalis Uropodinae Kramer. 1881 Uroseius koehleriW isniewski, 1979 (D only) Formica rufa M ic rocy I li ha min ima (Kramer, I 81,12) I Phaulodinychns ( P haulo dinychus) sp . n. I Camponotus herculeanus P. (Uropolyaspis) hamulikr (Michael. 1894) t Lasius niger P. ( U.) spin

Trcmalurinae tserlese. I 917 O odittyc hus spotuliJeras (Moniez. I 892) ssp. bectwirhi Wisniewski, 1979 4 Camponotus herculeanus & C. vagas O. ovalis (C. L. Koch,1839) 22 Formico, Camp onotus, La.riu;r- spp. I pi duro p odo di a lveolara (Hirschmann & Z. -Nicol. t96r)2 2 Dendn usius luIiginosus. Lasius mcn- dionalis & 1,. Jlavus I. interstructura (Hirschmann & Z.-Nicol. l96l) '!2 I Formica pratensis Trematurella elegans (Berlese, 191 6) 2 3 Fonnico polvctena, L. niger Uro dias pis tcct a (Kramer. 1876) 5l 2 Uroobovclla obovara (Canestrini & Berlese, 1884) 1 1 Formica flsca, Lasius niger Dinychopsis canla (Hull, 1918) 2 | l.'ormica polyctena

Prodinychidae Berlese, I 9 I 7 Trac hy xenura pyriforzri (Berlesc. I 920) 3 I Formicaexsecta, F. aquilonia, D. fuliginosus P rodi ny chus (A I lodi nychus) flagel liger (Berlese, l9l0) 32 3 Fonnicafusca Diny chus a rcuatus (Tragardh. 1943) 3 3 Formica pol.yctena D, septentrionalis (Triigirdh, 1943) 32 3 Formica polyctena D. ca r inut us Be rlese, 1903 2 1 Losius & Myrmicaspp. Associati

Mite species Host species

Polyaspididae Berlesc, l9 I 3 Discourella (D.) morlesta (I-eonardi. 1899) 4 I 2 Camponons ligniper da, Formica fusca, Lasius 'frachytes aeg1ota (C. L. Koch, l84l) 5 I 2 all ants T. ninima Trigitrdh. l91O 5 I 2 Formica rufu-gr

Sejina TriigArdh. I 93tt SejidaeBerlese,lS95 1836 2 Formica polyctena ^Sejas togatu.r C. L. Koch, 2 Microgynidae Triigdrdh, 1942 M icrosejus t runcicola T r i\gir dh, 1942 2 3 Formica aquilonia, F. rufa, F. fusca, Leptothorax s?p. ZerconinaTragArdh. 1946 Zerconidae Canesrini. l89l Para z e rcon (Para ze rc on) radi atus (Berlese. 1910) 5 I 2 allants Pro ze rco n trae ga rdhi (Halbert. 1915) 2 3 Lasius flavus, L. niger P. lochiSellnick.l943 5 2 all ants Zercon solenites Haarlav , \942 5 3 Formico spp . Z. lindrothiLundqvist & Johnston, 1986 5 3 Lasius spp., Fonnico fusca Z. spatulatus(C.L. Koch, 1939) 3 I all southern ants Z. sp. n. (aff. Z. spatulatus) 2 I Camponotus ligniperdu. Losius niger. L. Jlavus Z. airiosusTriigirdh, 1910 5 3 Formica, Lasius, Camponotus, Myrmica & Leptothot0x spp.

& L. umbrdus) or thc species itself is rare in southern Finland are inhabited bv Urotrachfies (L. meridionalis). Only onc nest of each of these formicarius and Urodiscella n'usmunni, but a few species was checked in Finland for this study. re- isolated populations of L. Jlavus in the small is- sulting in four taxa present only in the nest of lands of thc Bothnian Gulf havc 0 . villosella asthe L. rnixttrs'. namely Urodiscella philoctena, oplitinc guest. U. wasmunni may be dependent Phaulodinychus spinosulus, Urojanetia w sman- alsct on IJ. formicarius, as it has never been found niana, an

floor of a since long unused barn in the Seitsemi- Hypoaspis myrmecophilus and Cosmolaelaps nen National Park, central Finland. The barn had cuneifer. Locally common zerconids (Prozercon been used to store grass from the surrounding kochi, Parazercon radiattts, Zercon curiosus, Z. fields. A new species of Phaulodinychus was spatulatus and, Z- so/enrles, the latter only in north- found in this ncst together with the only Finnish crn and central Finland) may be abundant- inland population ol Ephippiochthonius tet- Uropodids are most often represented only by the rachelatus, a pseudoscorpionid common in the non-hostspecific Oodinychus ovalis. In E Finland, outer and middlc zoncs of the SW archipclago. however, Oplitis punduta ssp. "A" may be locally The second ncst was found in the Saaristomeri Na- abundant in anthills of F. rula, and the Finnish tional Park (SW archipclago). This huge nest in- deutonymphs ol Uroseius koehleri are all from cluded the remaining spccialist species of C. fter- nests of that species. An anthill of F. polyctena culearurs: Oplitis pandata n.ssp. "B". Oodinychus from Kankaanpdii, inland of W Finland, had the sparuliferus bcckwithi and Dipolyaspis testaceus highest density of uropodid mites, almost 7000 criocephali. Wisniewski & Hirschmann (19U3) de- specimens in a sample of less than two liters. The scribed a species Dinycllrrs camponoti based only species composition was quite exceptional, with on larval stage. As long as most larvae of the Di- Ditryclrus septentrionalis and Sejus togatus ir.huge rryclrus spp.. preferring decaying wood, have numbers, Dinychus arcuatus and Dinychopsis never been described. it is impossible to know rutula among the abundant species, together with with which spccics it will bc synonymized, but the the very abundant myrmecophilous Oplitis pan- strongest candidalcis D.t'arinatus Berlese, a rcgu- ddtu spp. "A", and a fcw Ootlinychus oyalrs and lar but not abunclant spccies in SW-Finland. Urojanetia coccineu. A rich sample of lre- Old records from othcr parts of Europe arc thc muturella elegans was collcctcd from an anthill of least reliablc among Fornr'r'a spp.. The most intcr- f. polyctena- esting specics is f'. (()tptoformica) exseoa. lt is Anthills of F. polyctena havc been thoroughly the main host ol Urojanetia cocclnea for the island investigated in Poland (Wisniewski 1979a, b, ctc). and coastal populations and for the meadow popu- and most Polish uropodid species are recorded lations with flat anthills in the inland. The inland from these. Unfortunatel)', the nests of other bog populations of F. exsecta usually have quite species of the F. rufa group are practically unin- rich populations of a new subspecies of Oplilis vestigated in the same area, and generally no in- pandata. These populations of F. exserta often formation has been given as concerns numbers of build high. relatively narrow anthills, and the pos- spccimens of diffcrcnt mitos. A dircct comparison sibility docs cxist that wc arc dealing with differcnt of thc guests of this spccics bccomes difficult also ant taxa, as is now gcncrally accepted for tho tradi- l;ccausc the concept of F. polyctena may bc tional F. rufu. Scvcral local non-myrmecophilous slightlv different in thcsc two countries. specics of Uropodidac and Zerconidae and numc- Ncsts of Formica (ServiJitrmit'a) ilsr:a undcr rous othcr nresostigmatid. prostigmatid, tar- stones and in thc soil rnostly sharc the myr- sonemid, astigmatid. and oribatid mites are often mecophilous species Uroobovella obot,ata and present in old anthills of the southern type of F. e-rr- common zerconids with nests of Lasius niger in secta. This seems to be the most common "nat- similar habitats. The nests of l. (Sen,iformica) in ural" habitat of Traclty-tcnura p,vriforntis, a com- decaying tree stumps are preferred localities for mon species in man-made habitats rich in organic the r are P rodiny c hus fl age ll ige r in E Finland. This debris. is repeatedly confirmed from nests of F. /&sca, but Nests of the other species of the P. exsecto-group apparently many .rf thc trce trunks wcrc inhabited have not bccn sufficicntly invcstigatcd. Most of l'ty F. gagatoides. P. J'lagelliger and Serviformica them may sxceptionally bc accepted as thc host of spp. prefer decaying tree trunks of birch. willow, U. coccinea. Apart from this group, nests of F: aspen, and alder in virgin forests, and it is hard to (Raptiforntica) sanguirtea and its slavcs (Servifor- dccidc whether the association is due to common ,r1ica spp.) sometimes have a rich population of U. preference of this special habitat of decaying wood coccinea. or of true reciprocal dcpcndcncc. Most nests of Formiut aquilonia, F. rufu, F. The mite fauna of the ncsts of Finnish Myr- polyctena, aruJ F. prutensis are verv rich in a fcw micinae is generally poor, with the exception of common spccics of Laclaptidae, espccially 'l'etramorium caespitttttt. Large nests of this Association of mites with ants 19 species are normally inhabited by both of its References specialists, Urojanetia excavato and Oplitis Athias-Binche. F., 1980. Contribution a la connaissance des libres (Arachnides: Anactino- ovatula. Uropodides triches) de quelques ecosystdmes forestiers euro- A large number of nests of Leptothorax acer- peens. - PremiCre partie, Morphologie, systdmatique vorum, Myrmica scabinodis, M. rubra, M . et biologie. - Thesis Univ. Pierre et Marie Curie, ruginodis, and many nests of other Finnish species Paris, 80 pp. Baker, E. & G. Wharton, 1952. An introduction to of Myrmica ar.d Leptothorax have been investi- acarology, The Macmillan Company, New York. 465 gated by Dr. Michael Saaristo and the author. pp. They regularly include specimens of common Balogh, J. 1938. Magyarorszdg hangyabolyban 6lo at- mesostigmatid species from their surroundings kdir6l 1. -Fol. Entomol. Hungar.3: 105-109. Berlese. A., 1881. Indagini sulla metamorfosi di alcuni (P ro z e rc o n koc hi, Parazerco n radiatus, Trachyte s acari insetticoli. - Atti R. Ist. ven. Sci. Lett. Arti (5) aegrota, Zercon spp. ). Uropodid species are, how- 7:747-.751. ever. exceptional, and if so, in low numbers 1882-1903. . in Acari Myriapodi et (Oodinychus ovalis, Uroobovella obovata, Tre- Scorpiones hucusque in ltalia repefta. 143 pp & 11 pl. Padova. maturella elegans). 1904 a. Diagnosi di alcune nuove specie di Acari ita- A reliable comparison of lhc ant hosts of differ- - liani. mirmecofili e liberi. -Zool. Anz.27: 12-28. ent mite species in different parts of Europe is pos- - 1904 b. Illustrazione iconografica degli Acari Mir- sible only when verified data from other parts of mecofili. - Redia 1:2994?4. Bernard, F., 1968: Les Foulmis (Hymenoptera, For- availahle. Europe have become micidae) d'Europe occidentale et septentrionale. - The species composition of the mite groups dis- Faune de l'Europe et du Bassin Mediterranden 3: 1 cussed here in ant nests is now relatively well 411. known in Finland and, obviously, also for the Canestrini, G., 1884. Acari Nuovi o poco noti. - AttiR. Istit. ven. Sci. Leu. A i(6)2; 8-10. whole Europe, we know very of northcrn but little Canestrini, G. & A. Berlese, 1884. Sopra alcune nuove about thc rclationships of these species with their specie di Acari ltaliani. - Atti Soc. Ven.-Trent. Sci. ant hosts. Nat. 9: 175- 182. Canestrini, G. & R. Canestrini, 1882. Acari italiani nuovi o poco noti. - Atti Real. Ist. Ven. Sci. Lelt. Ani (5) 8: 913-930 + Pl. VIII-X. 1927. The guests of British ants. Their Taxonomic position of Oplitinae and Donisthorpe, H. . habits and life histories. George Routledge & Sons. Trachyuropodinae London, 244 pp. Forel, A. 1874, Les Fourmis de la Suisse. -N. Denkschr. Trachyuropodinac is often united with the Op- Schweiz. Gcs. Naturw., Ziirich 26: iv + 452 pp. litinae (Hirschmann & Zirngiebl-Nicol 1962. etc) Greim, E., 1952. Die Okologie der Amcisenmilben to a singlc suhfamily. The rcason for this is their Frankens. - Thesis Univ. Erlangen, 168 pp. similar. brushlike mouthparts and common reduc- Haller. G. 1877. Antcnnophorus uhlmanni, ein neuer Gamaside. Arch. Naturg. 43: 57-62 + Pl. V- of the dorsal larval platcs. Apart from thesc - tions I882. Die Milben als Parasiten der Wirbellosen, similarities, however, most of their structures arc - insbesondere Arthropoden. - 32 -. Halle. quite dissimilar and they can be independently de- Hirschmann. W., 1957. Rumpfbehaarung un Riicken- rived from two non-myrmecophilous groups of fliichen. (G. P. 1). - Acarologie 1: 3-20 + 26 Pl. 1984. Die latotutuli-Gruppe, eine ncue Adulten- Uropodidae. These are the Oplitinae which are Gruppe der Ganggattung Oplitis. Teilgiinge, Stadien minute, semiglobular, with smooth non-sculp- von 8 neucn Oplitis-Arten aus Kamerun tured plates as found in Uroobovella and Uroster- (Trachyuropodini, Oplitinae). - Acarok)gie -31 (G. nella. and. the Trachyuropodinac which are largc, P.476): 156-162. Hirschmann, W. & I. Zirngiebl-Nicol, 1961. Die Gat- flattened oval, brightly red, and with extremely tung Trichouropoda Berlese 1916. - Acarologie 4: l- distinct scufptured plates as in Trachycylliba 34 + Pl. 1+. (= Dist'opoma auct.). Trachyuropoda and, Dis- - 1962. Uropodiden. Die Gattung Uroobovella Berlese Lopoma aoct. were also placed in the same group 1903 nov. comb. Tcilgiinge von Nenteria nov. comb. Erstversuch der Aufstcllung eines Gangsystems der b_v- (1952) Baker & Wharton and Athias-Binche Uropodiden aufgrund der Gnathosoma-Unterseite (1980). ln my opinion. the similarities in the und Chelicere. - Acarologie 5 (G. P. 6): 57 80. PI. mouth parts are simply due to parallel adaptions 2211. to similar feeding habits, and the larval reduction Janet, C., 1897 a. Etudes sur les Fourmis, les Guepes et les Abeilles: 38-47. (in Rapports des animaux myr- is merely a parallel advanced evolution between mecophiles avec les fourmis. H. Ducortieux. groups. Limoges, 98 pp.) 20 Pekka T. Lehrinen

- 1897 b. Sur les rapports de l'Antennophorus - Berlin. uhlmanni avecle Lasius mixtus Nyl. -C. R. S. Acad. 1897 a. Uber einige myrmekophile Acarinen I. - Sci. 124:583-585- ZooL Anz.20: l7tll73. Karawaiew. W., 1906. Weitere Beobachtungen iiber - 1897 b. Ubcr einige myrmekophile Acarinen II. Arten der Gattung Antennophorus. - Mem. Soc. Zool. Anz,. 20: -146--350. Nat. Kiew 20: 209-229. 1898. Uberdic Grste von Tetramorium cacspitum. - Karpelles. L., 1891: Bausteine zu einer Acarofauna Un- - Verslag ol. 53. Somervergad. Nedcrl. Entomol. garns. - Math, Naturwiss. Mitt. Akad. 25: 80-134. Ver.: 6t)-{5. Kneissl, L., 1907. Uroobovella wasmanni Kneissl. Eine 1899. Weitere Nachtrage zum Verzeichniss der neue myrmckophile Milbc. - Zcitschr. Wiss. Insek- Ameisengdste von Holldndisch Limburg. - Tijdschr. tenbiol. 3: 190-191. Entomol. 12:158-171. 1908. Nachtrag zur Bcschrcibung von Urodiscella 1902. Zur Kenntniss der myrmekophilen Anten- - wasmanni mit Aufstellung eincr neuen Varietat U. - nophorus und anderen auf Ameisen und Termiten philoctena var. schmitzi. - Zeitschr. Wiss. Insektcn- reitende Acarincn. - Znol. Anz.25:66-76. biol.l:226-229. Wisniewski. 1., 1979 a. Die Wander - und Dauer- Krasinskaya. A.. 1961. Morfo-biologiceskie osoben- nymphe von Trichouropoda elegans (Kramer 1882) nosti postembrialnog razvitia uropodnych klescej und die Dauernymphe von Trichouropoda spatuli- lcningradskoi oblasti. - Parazit. Sborn. 20: 108 146. fera (Moniez 1892) aus Polen (Trichouropodini, Leonardi, G., 1895. Intorno ad alcune nuove specie di Uropodinae.)- Acarologie 25 (G.P. 312): 37 38, Pl. Acari italiani ecc. - Atti Soc. Vcn.-Trent. Sci. Nat. 5: fig. 34 & Pl. 6: fig, 35. 2: I-t}. 1979 Stadicn von 2 neuen (Uroseius)- 12) - b. Uroseius - 1896. Notizie intorno agli acaroidei vive.ti nci for- Arten aus Ameisennestern Polens (Uropodini. micai. in G. Canestrini: Acarofauna ltaliana 7: l- Uropodinac). - Acarologie 25 (G.P. 316):4346, Pl. 38. VI- Lubbock. J., ltl8l. Observations on ants. bees, and 19'79 c.Ztr Kennlnis der Uropodidenfauna Polens. - wasps. - J. Linn. Soc. London, Zool, 15; 38G(Znd - Acarologie 26 (O. P.339): 6&-74. ed. 1894 as a book). 1980 a. Gang einer neuen mit Trichouropoda spatuli- Michacl, A. , l89l . On the association of Gamasids with fera (Moniez 1892) verwandten Art aus Ameisen- ants. - Proc. Zool. Soc. Lond. l89l:638. nestern Polens (Trichouropodini, Uropodinae). 1894 a. Notes on the Uropodinae. -J. R. Microsc. Acarologie 27 (G.P. 342):8-10 + Pl. I:fig.2. - Soc. 189,1:2tt9 319. Pl. VI-VIl. 1980 b. Stadium ciner neuen -Art aus 1894 b.'Ihe President's address: The progress and - Polen. - Acarologie 27 (G.P. -350): lGIT + Pl. 2: fig. present state of our knowledge of the Acari. - Trans. Il. Linn. Soc. Lond.. Zool. 1894: l8 33. Wisniewski. J. & W. t{irschmann. lg8J. Stadium einer Monicz. C,, 1892. Mdmoirs sur quelques Acariens et neuen Dinychus-Art aus Ameisennest in Polen. - Thysanourcs parasites ou conrntensaux des fourmis, Acarologie 30 (C.P. 13tl): 133-134. - Rev. Biol. Nord France 4 ( l0); 1-12. Yarrow, T. 1955:'l'he British ants allied to Formica rufa I894. Sur quelques arthropodcs trouves dans des L. (Hymenoptera, Formicidae): -'frans. Soc. Brit. - Fourmilicrcs. - Rev. Biol. Nord France 6 (6): -201 -. Entomol. 12: l-4ti. Pecina. P.. 1980. Additional data on several Czecho- slovak members of the subfamily Trachyuropodinac Berlese. 1928 (Uropodidae. Mesostigmata)- - Acta Sammanfattning Univ. Carol.. Biol. 197ti: 357 388. Sellnick. M.. 1939. Milben als Parasiten von Insekten. - En checklist 6vcr myrgester inom kvalsterfamil- Verh, VII lnt. Kongr. Entonrol., Bertin, 1938 2: jerna Uropodidae, Prodinychidae, Polyaspididae, t300- 1307. Antennophoridae, Sejidae, Microgynidae och SrorkJn. J.. 1940. Myrmecophilni Acari z Vdst. fech. Zerconidae i Finland presenteras (Tab. 1). I Tab. Cs. Zool. Sprrl. Praha R: ltr6-171. Thor. S.. 19fi). Milben als Ameisenfeinde. - Nyt Mag. I listas iven v:irdarler bland myrorna och typen av Naturvidensk. 37 : 3-l 5377 . forh6llande anges som: (tabellkod A1) beroende Triigirdh. I., 1942. Uber die Trematurini Berlese. - av en eller tve nArbesliktade myrarter. (A2) van- Z