ORIBATID MITES (ORIBATEI, ACARIFORMES) IN BIRD FEATHERS ...

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Acta Zoologica Lituanica, 2004, Volumen 14, Numerus 1 ISSN 1648-6919

ORIBATID MITES (ORIBATEI, ACARIFORMES) IN BIRD FEATHERS: NON-PASSERINES Dmitri A. KRIVOLUTSKY1, Natalia V. LEBEDEVA

2, 3

1

Institute of Parasitology of the Russian Academy of Sciences, Lenin pr. 33, 119071 Moscow, Russia. E-mail: [email protected] Branch of Murmansk Marine Biological Institute of the Kola Research Centre of the Russian Academy of Sciences, Chekhov pr., 41, 344006 Rostov-on-Don, Russia. E-mail: [email protected] 3 Rostov State University, 148 Pushkinskaya St., 344006 Rostov-on-Don, Russia 2 Azov

Abstract. The authors have established that oribatid mites (Acariformes, Oribatei) are transmitted by birds. More than 2,100 birds of 150 species from different ecological groups were surveyed. 180 species of live oribatid mites were collected from bird feathers. Some oribatid mite species live in bird feathers constantly. The authors assume that oribatid mites feed not on the particles of the skin, but on fungi inhabiting bird feathers, as all oribatid mites are mycophagous. Oribatid mites were found in different forms of their lifecycle. It has been established that they reproduce in bird feathers. Transmission of these microarthropods by birds may be a principle cause of their distribution, especially in the remote Arctic areas, on the Oceanic islands, and in the oases of deserts. Key words: oribatid mites, bird feathers

INTRODUCTION Transmission of oribatid mites (Acariformes, Oribatei) by birds is a well-known phenomenon revealed by the authors of this article in 19972002. It has radically changed the knowledge of factors determining geographical distribution of these mass (from 10 to 400 thousand specimens per 1 m3) soil inhabitants Though the relationship between birds and oribatid mites inhabiting their feathers has been studied already for many years, not a single research paper has reported about the study results. The authors of the last review Mites and birds: diversity, parasitism and co-evolution (Proctor & Owens 2000) describe findings of 2,500 species of oribatid mites of 40 families found in bird feathers. However, they do not name any oribatid family or species. This is amazing because oribatid mites have been not only found in nests of birds, but also investigated (Borisova 1968; Wasylik 1971, 1973; Erik & Glukhova 1974). Some species of oribatid mites have been investigated quite thoroughly (Borisova 1969, 1978; Gembitsky & Andreichikova 1969; Gembitsky 1970; Yaroshenko & Kharchenko 1972; Lebedeva et al. 1998, 1999; Tryjanowski et al. 1999). Biogeographical peculiarities of the distribution of soil microarthropods, a mass group inhabiting all types of soil is still unknown. That oribatid mites may be transmitted by water and wind, with vegetable remains and soil, it is common knowledge. It is insufficient for the explanation of the distribution of microarthropods, especially in the

Arctic, the Antarctic, and the Oceania (Krivolutsky 1995). In the biota of the Arctic and the Antarctic, species diversity is particularly poor, and it is impossible to investigate it throughout the year. Soil microfauna, a diverse world of organisms that are less than 1.0 mm long and abundant in the soils of the Arctic zone and tundra, is one of the groups available for research there. Representatives of that microfauna, microarthropods in particular, may be found in great quantities even in the tiniest patches of soil, assemblages of moss, lichen, and rocks. Special attention should be paid to the research into the ways of their distribution in the Arctic and the Antarctic. It is known that some species of microarthropods may be transmitted by water and wind, with vegetable detritus and soil, with driftwood and by ice floes. However, in most cases that knowledge is insufficient for the explanation of mechanisms of distribution of small arthropods, especially in the Arctic, the Antarctic, and the Oceania (Krivolutsky 1995). Birds, too, play a significant role in the distribution of microarthropods; the more so that blood-sucking Izodides and Gamasidae, which are also found in remote ocean islands of all the latitudes, have been revealed in the nests and feathers of polar birds for a long time. Hammer (1965) described a population of Mucronothrus nasalis Tragardh, a European marsh species found in New Zealand, but it was not discovered how that species found its way to that island country. Possibility of its transmission of this species to this locality by birds was not considered.

Oribatid mites in bird feathers

Three species of oribatid mites were found in the feathers of Hazel Grouse (Bonasa bonasia) by one of the authors of this article during collection of ectoparasites of birds in the Perm Province (Russia) in 1959. However, most scientists have not taken this into consideration. In 1977, Hydrozetes lacustris, oribatid mites inhabiting several continents, were detected in the stomach of Red-necked Phalarope (Phalaropus lobatus) from the Chukotsk Peninsula. It is quite possible that if not the oribatid mites themselves, then their eggs are transmitted by birds at least at short distances equal to the distances of their diurnal passages during seasonal migration. Hypotheses about more distant transmissions of oribatid mites carried out by birds in this way are hardly possible because the chemical medium in birds stomachs of and the time of the presence of food in their intestinal tract do not allow mites to survive there, so, they cannot be transmitted at such long distances. Birds make long-distance passages, including oceanic birds, waterfowl, and birds inhabiting areas near water bodies, are of special interest for the researchers of distribution of oribatid mites because they make the most distant passages, sometimes through entire continents. In addition, on their passage routes and in their nesting and wintering grounds birds keep to biotopes that are similar according to conditions of wetness, therefore, oribatid mites can be easily naturalised there. The current paper is a sequel to a series of our studies on invertebrates, partially oribatid mites, which are interrelated ecologically with birds during different stages of their lifecycle (Lebedeva et al. 1998, 1999; Krivolutsky & Lebedeva 1999). Studies carried out by the authors of this paper in 19982002 revealed that there is one more earlier unknown factor determining the distribution of microarthropods: constant presence of different groups of mites (Oribatei, Gamasidae, Acaridia, Trombidiformes, and Prostigmata), collembolans, Diptera larvae, staphylinids, etc. in bird feathers.

MATERIAL

AND METHODS

A total of 2,100 carcasses of birds belonging to 150 species was investigated. Over 180 species of oribatid mites in different stages of their lifecycle were found in bird feathers. The birds were obtained during special ornithological expeditions. In addition, carcasses of birds shot in the Sheremetyevo Airport (Moscow), in the zoological gardens of Moscow, Rostov, and Odessa, were analysed. Carcasses of birds that perished during the winter when oribatid mites could not penetrate their plumage from the surface of soil were also collected. Quite a great number of birds were collected on the

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coasts of the Barents and Black Seas after storms, or they were shot by hunters. The main regions of material collection were the coast of the Sea of Azov (Delta of the River Don, brackish lagoon (46°N, 38°E) on the eastern coast of the Sea of Azov), the coastal area of the Black Sea (environs of Odessa), the Kostroma, Tver and Moscow Provinces, Karelia, the coast of the Barents Sea (environs of settlement Dalnya Zelentsy), Novaya Zemlya, Spitsbergen, Gdañsk Peninsula, the lower reaches of the River Volga, Belarus (Homel Province), the Autonomous Republic of Komi, the Far East, Kaliningrad Region, and Abkhazia. The majority of microarthropods was recovered from the feathers of birds using common termo-eclectors of Berlese-Tullgren. There skins of birds (feathers downward) or those of small mammals, such as mice, American minks, and Norway rats, as well as patches of skin (10 × 10 cm2) from the abdominal part of the body of dogs and goats were kept for two days. It should be taken into account that oribatid mites and collembolans were collected from the skins of all the investigated specimens of mammals, even from those of mice and rats kept in vivaria and from the floor litter of those vivaria. Because oribatid mites were found only in the feathers of 80% of birds (less than 40% in Passeriformes), special research was conducted in order to reveal the most effective method of oribatid collection. Consequently, an experiment was carried out on Tree Sparrows (Passer montanus) of the Moscow Province: preliminary-refrigerated solid carcasses of birds, disembowelled and salted from inside carcasses of birds, and the patches of skin taken from the carcasses were investigated separately. It was found out that when the skins of birds and mammals were used for the collection of microarthropods, distinctly incomplete collections occurred with respect to the most diverse and interesting animals: epifauna, or inhabitants of the soil surface, litter, and plants. Presumably, while taking out the skin, most of those large microarthropods were lost. In addition, we revealed that mircroarthropods survive in bird feathers as long as two years of their storage in refrigerators at a temperature ranging from -18 to -20°C. We investigated the feathers of bird species the list of which is presented in the Annex I. The English and Latin names of bird species are presented according to Behme and Flint (1994). Oribatid mites inhabiting feathers of birds of prey and owls Practically, all birds of prey inhabiting regions of Russia are protected by law. Thus, to investigate oribatid mites inhabiting their feathers is quite a difficult task, because only specimens killed in accidents (collisions

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Krivolutsky, D.A., Lebedeva, N.V.

with high-voltage wires, lighthouses, utility poles, planes, cars, etc.) were available for research. Consequently, we could investigate only some accidental specimens of birds of prey. Among them, there were also a few specimens of hawks and harriers shot by hunters during spring and autumn hunting seasons. In total, 53 specimens of 19 species of birds of prey and owls were investigated: Honey Buzzard (Pernis apivorus), Black Kite (Milvus migrans), Hen Harrier (Circus cyaneus), Marsh Harrier (Circus aeruginosus), Goshawk (Accipiter gentilis), Sparrowhawk (Accipiter nisus), Common Buzzard (Buteo buteo), White-tailed Eagle (Haliaeetus albicilla), Peregrine (Falco peregrinus), Saker Falcon (Falco cherrug), Hobby (Falco subbuteo), Red-footed Falcon (Falco vespertinus), Lesser Kestrel (Falco naumanni), Kestrel (Falco timunculus), Long-eared Owl (Asio otus), Short-eared Owl (Asio flammeus), Tengmalms Owl (Aegolius funereus), Tawny Owl (Strix aluco), and Ural Owl (Strix uralensis). Despite the great size of those birds, oribatid mites in

their feathers and nests were very sparse. Carabodes forsslundi, Phthiracarus sp., Brachychthonius berlesei, and Oppia splendens were found in the feathers of White-tailed Eagle; Chamobates cuspidatus in those of Ural Owl; Opiella nova and Carabodes areolatus Tengmalms Owl; Oppia unicarinata Long-legged Buzzard (Buteo rufinus); Malaconothrus egregius, Carabodes labyrinthicus, Ceratozetes cisalpinus, Tropacarus carinatus, and Phthiracarus globosus Lesser Kestrel; Oppia splendens Red-footed Falcon; Liochthonius sellnicki, Tectocepheus velatus, and Trichoribates trimaculatus Peregrine; Nothrus palustris, Tectocepheus velatus, Trhypochthonius tectorum, Oppia unicarinata, Tropacarus carinatus, and Opiella nova Marsh Harrier; and O. nova Hen Harrier. We did not find any oribatid mites in the feathers of quite a large number of birds (Red-footed Falcons, Sparrowhawks, Kestrels, etc.). The list of birds of prey inhabited by oribatid mites most abundantly is presented in Table 1.

Table 1. Oribatid mites in bird feathers of prey and owls.

Black Kite

Sparrowhawk

Goshawk

Kestrel

Long-eared Owl

Shorteared Owl

Total

1 Number of birds Hypochthonius rufulus Liochchthonius sellnicki Phthiracarus borealis Tropacarus carinatus Steganacarus striculus Nothrus palustris Camisia spinifer Platynothrus peltifer Trhypochthonius tectorum Malaconothrus egregius Hermannia reticulata H. scabra Poroliodes farinosus Belba sp. Metabelba sp. Cepheus cepheiformis Eremaeus oblongus Gustavia microcephala Carabodes areolatus Tectocepheus velatus Autogneta willmanni Oppia fallax

Common Buzzard

Oribatid mite species

Honey Buzzard

Bird species

2 4 1 1

3 8 1 1 1 1 1 9 2

4 4 3 1 2

5 6 1 1 1 1 1 1 1 1

6 5 1 1 1 1 2

7 3 1 1 1 1 1

8 6 1 2 1 1

9 3 1 1 1

10 39 2 6 1 1 1 2 1 2 1 2 1 2 1 1 2 2 2 1 3 14 2 1

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Table 1 continued

1 Oppia maritima Oppia minus Oppia ornata Oppia splendens Oppia unicarinata Opiella nova Quadropia quadricarinata Suctobelba trigona Suctobelbella acutidens Peloribates longipilosus Zygoribatula exilis Liebstadia similis Scheloribates latipes Scheloribates laevigatus Neoribates roubali Diapterobates humeralis Fuscozetes fuscipes Melanozetes mollicomus Trichoribates trimaculatus Chamobates cuspidatus Minunthozetes pseudofusiger Parachipteria punctata Pergalumna nervosa Number of oribatid mites

2 1 5 2 1 6

3 1 1 7 1 1 1 14

4 1 2 6

It is obvious that the feathers of birds of prey are sparsely inhabited by oribatid mites. This could be possibly explained by the dryness of their skin and high body temperature, which is unacceptable for soil microarthropods. It should be noted that preimaginal stages of oribatid mites, as well as other representatives of another mass group of microarthropods (Collembola), were actually absent in bird feathers of prey. Among nocturnal bird species, the greatest amount of oribatid mites was found in Long-eared Owl and Shorteared Owl (Table 1), whereas the rest of the species occurred singly. Opiella nova and Carabodes areolatus were found in the feathers of Tengmalms Owl, Chamobates cuspidatus, Diapterobates notatus, and Scheloribates laevigatus in those of three Ural Owls. No oribatid mites were detected among eight microarthropods found in the feathers of Tawny Owl. Thus, 52 species of mature oribatid mites were collected from the feathers of birds of prey. We should point out also a great diversity of relatively large oribatid mites belonging to epifauna (Hermannia, Nothrus, Camisia, Platynothrus, Belba, Gustavia, Cepheus, Trichoribates, Neoliodes, Fuscozetes, Parachipteria, Pergalumna, and Neoribates), which was not found in bird feathers

5 1 1 10

6 1 2 3 1 1 1 1 1 2 1 15

7 1 1 1 2 1 10

8 1 1 1 1 8

9 2 1 5

10 1 1 1 5 4 12 1 6 1 1 1 1 2 2 1 2 1 1 3 1 2 2 1 46

from other taxonomic groups. Small oribatid mites (Oppia, Suctobelbella, and Liochthonius) made in total about 30% of all the specimens found in feathers. In addition, immature specimens were not found at all. Among the oribatid species found in bird feathers, not a single species was found at least in 50% of bird species. That means that they inhabited the feathers of birds of prey accidentally. Oribatid mites inhabiting nests of birds of prey and owls deserve special attention as, in our opinion, they find their way from the nests to the feathers of incubating birds (they also get into the feathers of birds from soil and trees during their nights resting). We investigated the population of microarthropods in 53 nests of 19 species of birds of prey and owls: Black Kite, Saker Falcon, Peregrine Falcon, Hobby, Merlin (Falco columbarius), Red-footed Falcon, Kestrel, Goshawk, Common Buzzard, Rough-legged Buzzard (Buteo lagopus), Long-legged Buzzard, Spotted Eagle (Aquila clanga), Barred Owl (Tyto alba), Eagle Owl (Bubo bubo), Ural Owl, Long-eared Owl, Snowy Owl (Nyctea scandiaca), and Scops Owl (Otus scops). Nests (n = 6) of Saker Falcon, Eagle Owl, Scops Owl, Spotted Eagle, Red-footed Falcon, and Kestrel were

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collected in the Zoological Garden of Odessa. On the basis of our research, we assumed that oribatid mites and other microarthropods could penetrate the clean nest building material from the feathers of birds because only collembolans were found in the soils and litter of aviaries. It is even more interesting that oribatid mites were found there: Tectocepheus velatus and Banksinoma lanceolata were found in the nest of an Eagle Owl and Haplochthonius simplex (a species that had never been found in Ukraine before) in that of a Saker Falcon. In the nests of a Saker Falcon and Scops Owls, larvae of the Pseudoscorpiones were found. In the remaining nests from the Zoological Garden, no oribatid mites were found. It should be noted that oribatid mites were not found in 32 of the 53 investigated nests. However, all those 32 nests had been built earlier by Corvids (Rook (Corvus frugilegus), Magpie (Pica pica) and Carrion Crow (Corvus corone corone)). As they were high in trees, they could be dried well enough during the long drought of the summer of 2002, which made the environment there for microarthropods unfavourable. Quite a small number of oribatid mites were recorded in large nests of desert birds of prey. For instance, in the nest of Saker Falcon from Mongolia, only nymphs of Nothrus and Camisia and mature individuals of Protoribates sp., Opiella nova, and Oppia sp. were found. In the nest of Long-legged Buzzard, Tectocepheus sp., Passalozetes sp., and Phthiracarus sp. were detected. In the nest of Black Kite built high in a tree in the Tver Province (Russia), only two species of oribatid mites occurred: Opiella nova and Minunthozetes pseudofusiger. In three nests of Ural Owl built high in trees in Tver Province, only one specimen of Fosseremaeus laciniatus was recorded, and no oribatid mites occurred in the nests of Great Grey Owl (Strix nebulosa) and Goshawk built in one of the parks in Moscow. However, they contained just single specimens of collembolans and Gamasidae. In eight nests of Long-eared Owl located mostly in the Odessa Province (Ukraine), a single specimen of Chamobates cuspidatus was found, and in 10 nests of Red-footed Falcon, only one specimen of Trichoribates trimaculatus occurred. All the six investigated nests of Kestrel did not contain any oribatid mites. Single specimens of oribatid mites were found also in the nests of some other birds of prey: Punctoribates punctum (two nests of Scops Owl) and Diapterobates notatus (two nests of Peregrine Falcon). More oribatid mites were obtained from the nests built on the ground in tundra: Oppia unicarinata, Oribatula tibialis, Damaeus sp., Tectocepheus velatus, and Microtritia sp. in the nests of Snowy Owl; Hemileius initialis, Ceratoppia bipilis, Diapterobates notatus, Oribatula tibialis, and Carabodes areolatus in those of Rough-legged Buzzard.


Generalised data on the fauna of oribatid mites in the nests of several groups of birds of prey mentioned-above are presented in Table 2. Table 2. Oribatid mites in the nests of birds of prey and owls (+ denotes presence of oribatid mites).

Oribatid mite species Buzzards Falcons Owls Haplochthonius simplex + Phthiracarus sp. + Steganacarus striculus + Nothrus sp. + Camisia sp. + Damaeus sp. + Fosseremaeus laciniatus + Ceratoppia bipilis + Carabodes areolatus + Tectocepheus velatus + + Opiella nova + Oppia unicarinata + Oppia sp. + Banksinoma lanceolata + Passalozetes sp. + Oribatula tibialis + + Hemileius initialis + Protoribates sp. + Diapterobates notatus + + Trichobates trimaculatus + Chamobates cuspidatus + Punctoribates punctum + Achipteria sp. + Microtritia sp. + Number of nests 5 22 19 Thus, oribatid mites in the nests of birds of prey are not numerous. We collected oribatid mites in subarid regions during the warm season of the year. We may assume that those collections would have been more numerous if they had been done in the North or in a desert, where birds build nests on the ground and also during prolonged wet periods. So, this study should be continued. The study carried out by our colleagues from Belgium (Fain et al. 1993), who found about 700 oribatid mites belonging to 33 species of 24 families in the nests of Eagle Owl, may be an example. Their findings included the following nine species of oribatid mites: Nothrus sp., Trhypochthonius tectorum, Suctobelbella subtrigona, Tectocepheus sarekensis, Micreremus brevipes, Trichoribates trimaculatus, Galumna lanceata, Oribatula tibialis, and Dometorina plantivaga. The collected material demonstrates that the number of

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oribatid mites inhabiting nests of birds of prey is relatively small. They are mostly hemiedaphic forms with some admixture of epifauna; inhabitants of soil apertures are found there only singly.

in Table 4. Oribatid mite species found in bird feathers varied from 0 to 30. Oribatid mites were not recorded in seven specimens of investigated bird species (in two specimens of Black-headed Gull, three specimens of Coot (Fulica atra), one specimen of Shoveler (Anas clypeata), and one specimen of Arctic Skua (Stercorarius parasiticus). None of the oribatid mites found in the feathers of this group of birds were detected in 50% of bird species, and 33 of 34 oribatid species were found in them only singly. The greatest number of the specimens of oribatid mites was found in the feathers of Great Egret (65 specimens of 12 species) and Cormorant (Phalacrocorax carbo; 42 specimens of 18 species) from the delta of the Don River. In the feathers of Goosander (Mergus merganser), Scheloribates latipes and one species of Brachichthonius were observed. Many oribatid mites found in bird feathers of this species are common in the middle latitude of Russia. Among them, there are many small, capable of inhabiting soil

Oribatid mites in the feathers of waterfowl and birds inhabiting aquatic ecosystems 364 specimens of 66 species and nymphs (Table 3) and 311 specimens of 77 species and nymphs (Table 4) of oribatid mites were found in the feathers of waterfowl and birds that are closely related to marine ecosystems, respectively. Data about oribatid mites detected in the feathers of fullfledged Night Heron (Nycticorax nycticorax) nestlings (n = 5), which contained one specimen of Tropacarus pulcherrimus, as well as a nymph of an oribatid mite found in a young Bittern (Botaurus stellaris) and Peloribates longipilosus revealed in the feathers of Swan Goose (Anser cygnoides) have not been included

1 Number of birds Palaeacarus sp. Hypochthonius rufulus Brachychthonius sp. Liochthonius evansi Liochthonius sellnicki Phthiracarus borealis Phthiracarus nitens Phthiracarus sp. Steganacarus striculus Oribotritia loricata Nothrus palustris Camisia biurus Camisia horrida Camisia sp. Platynothrus peltifer Trhypochthoniellus setosus Trhypochthonius tectorum Malaconothrus egregius Nanhermannia coronata

2 3 -

3 3 1 1 1

4 6 1 1 1 -

5 1 -

6 4 1 1 -

7 2 3 4 1 1 1

8 4 1 1 1 2 1 2 9 -

9 7 1 2 1 -

10 11 12 13 14 15 16 3 2 1 3 1 1 35 1 1 - 1 5 - 1 - 1 - 2 -

Total

Moorhen (Gallinula chloropus) Spotted Crake (Porzana porzana) Black-headed Gull (Larus ridibundus)

Coot (Fulica atra)


Great Crested Grebe (Podiceps cristatus) Red-necked Grebe (Podiceps grisegena) Grey Heron (Ardea cinerea) Pond Heron ( Ardeola grayii) Little Egret (Egretta garzetta) Great Egret (Egretta alba) Mute Swan (Cygnus olor) Mallard (Anas platyrhynchos) White-eyed Pochard (Aythya nyroca) Shelduck (Tadorna tadorna) Tufted Duck (Aythya fuligula)

Table 3. Oribatid mites in the feathers of waterfowl and birds related to aquatic ecosystems.

17 83 1 3 1 1 1 1 1 9 6 1 2 2 1 3 13 1 1 1 2

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Table 3 continued

1 Nanhermannia sellnicki Hermannia reticulata Hermannia sp. Poroliodes farinosus Licnodamaeus pulcherrimus Belba sp. Epidamaeus sp. Metabelba sp. Eremobelba geographica Eremaeus oblongus Adoristes poppei Carabodes areolatus Carabodes femoralis Carabodes marginatus Tectocepheus velatus Autogneta tragardhi Concogneta delacarlica Multioppia sp. Oppia maritima Oppia ornata Oppia splendens Oppia translamellata Oppia unicarinata Oppia sp. Oppiella nova Quadroppia quadricarinata Suctobelba trigona Suctobelbella acutidens Suctobelbella hammeri Suctobelbella tatarica Suctobelbella sp. Banksinoma lanceolata Oribella paoli Limnozetes rugosus Oribatula tibialis Zygoribatula exilis Scheloribates laevigatus Trichoribates trimaculatus Chamobates lapidarius Minunthozetes pseudofusiger Mycobates parmeliae Punctoribates punctum Achipteria coleoptrata Parachipteria punctata Tegoribates latirostres Galumna sp. Pergalumna nervosa Oribatei nymphs (no determ.) Total number of mites

2 1 1 1 1 4 4

3 1 4

4 1 1 1 1 7

5 3 1

6 1 1 1 1 1 6

7 8 9 10 11 12 13 14 15 16 1 - 2 - 1 - 1 - 2 1 1 1 1 - 1 - 1 - 1 5 - 2 - 1 5 - 1 - 1 - 1 1 11 5 - 2 1 - 2 - 1 1 1 - 1 - 1 - 1 - 1 - 2 1 4 - 1 1 - 1 - 1 - 1 - 1 16 2 - 2 11 2 2 - 1 - 6 2 2 - 1 1 1 - 5 - 1 - 1 4 - 7 - 4 - 2 - 1 1 - 1 - 1 1 2 2 3 - 1 2 - 1 - 5 - 1 - 1 - 1 2 - 1 - 1 - 2 4 - 6 - 2 - 2 2 1 - 1 1 - 1 3 - 4 - 7 23 24 11 11 4 3 4 1 7 22

17 1 2 1 3 1 2 1 2 1 1 1 15 1 5 24 1 3 1 1 3 8 1 2 20 24 5 3 5 1 1 4 13 1 1 3 11 6 2 2 3 1 3 10 4 1 4 1 20 66

33


1 Number of birds Palaeacarus histricinus Palaeacarus sp. Hypochthonius rufulus Phthiracarus borealis Phthiracarus logneus Phthiracarus sp. Steganacarus striculus Eupthiracarus sp. Oribotritia loricata Nothrus silvestris Nothrus palustris Camisia biurus Camisia spinifer Camisia sp. Platynothrus peltifer Platynothrus punctatus Trhypochthonius tectorum Malaconothrus egregius Trimalaconothrus tardus Nanhermannia nana Nanhermannia coronata Hermannia reticulata Poroliodes farinosus Licnodamaeus pulcherrimus Belba corynopus Belba sp. Metabelba pulverrulenta Metabelba sp. Eremaeus fovelatus Eremaeus oblongus Ceratopia bipilis Xenillus tegeocranus Carabodes areolatus Carabodes forsslundi Carabodes labyrinthicus Carabodes marginatus Carabodes subarcticus Tectocepheus velatus Autogneta longilamellata Concogneta delacarlica

2 1 1 5 1 1 1 9 -

3 1 1 1 -

4 12 2 1 1 1 1 2 1 1 9 1 1 1 12 1 1

5 2 -

6 12 1 1 1 5 1 4 1 2 1

7 2 -

8 7 2 1 3 1 -

9 5 4 2 1 2 -

10 11 12 13 14 15 1 21 2 9 5 2 1 - 1 - 1 - 1 - 1 - 1 - 2 - 1 1 - 1 - 1 - 1 - 1 1 5 -

Total


Red-throated Diver (Gavia stellata) Black-throated Diver (Gavia arctica) Cormorant (Phalacrocorax carbo) Barnacle Goose (Branta leucopsis) Common Eider (Somateria mollissima) Stellers Eider (Polysticta stelleri) Herring Gull (Larus argentatus) Great black-backed Gull (Larus marinus) Common Gull (Larus canus) Kittiwake (Rissa tridactyla) Marsh Tern (Sterna anglica) Fulmar (Fulmarus glacialis) Little Auk (Alle alle) Guillemot (Uria aalge)

Table 4. Oribatid mites in the feathers of marine birds.

16 80 2 1 1 1 4 9 6 1 1 2 1 2 1 1 4 2 1 4 1 1 6 1 2 1 1 1 1 1 1 2 1 9 10 2 3 3 1 34 1 2

34


Table 4 continued

1 Caleremaeus monilipes Oppia fallax v. obsoleta Oppia maritima Oppia ornata Oppia splendens Oppia translamellata Oppia unicarinata Oppia sp. Oppiella nova Quadroppia quadricarinata Suctobelba trigona Suctobelba sp. Suctobelbella hammeri Suctobelbella acutidens Suctobelbella subcornigera Suctobelbella sp. Banksinoma lanceolata Banksinoma setosa Micreremus brevipes Scutovertex minutus Oribatula tibialis Zygoribatula exilis Odontocepheus elongatus Liebstadia similis Scheloribates laevigatus Neoribates roubali Trichoribates trimaculatus Chamobates cuspidatus Chamobates lapidarius Minunthozetes semirufus Minunthozetes pseudofusiger Punctoribates punctum Achipteria coleoptrata Parachipteria punctata Galumna sp. Pergalumna nervosa Pilogalumna tenuiclava Oribatei nymphs (no determ.) Total number of mites

2 1 1 4 3 1 9 11

3 1 3

4 1 2 2 1 5 2 1 1 5 2 1 1 2 3 1 3 30

apertures (infauna) species (for example, 12 species of the genera Palaeacarus, Liochthonius, Oppia, Oppiella, Quadroppia, Zygoribatula, and Minunthozetes). Findings of adults and nymphs of Palaeacarus histricinus, an oribatid species common in the woods of Eurasia but rare elsewhere, in the feathers of Cormorant were quite unexpected. Nothrus palustris, Eremaeus sp., Liebstadia similes, and Suctobelbella subcornigera were found in six nests of Cormorant built at the height of 1215 m

5 2 1 1 3

6 1 1 2 1 1 1 2 1 1 1 1 2 19

7 1 1 2

8 1 1 5

9 10 11 12 13 14 15 - 1 1 - 4 - 1 - 1 - 1 13 - 2 - 1 - 1 - 1 - 1 3 1 1 - 1 4 - 1 1 - 1 - 1 - 1 1 - 1 1 - 3 1 2 3 - 4 1 15 4 19 1 4 2 2

16 1 1 1 4 12 2 3 1 30 5 6 3 5 1 6 2 2 1 2 1 4 1 5 12 1 1 3 1 1 5 2 4 9 1 2 2 39 74

from the ground in the delta of the Don River. However, they were not found in the feathers of this species, though recorded in those of other bird species. In some birds species (all specimens of Crested Coot (Fulica cristata), Arctic Skua, one specimen of Shoveler, three specimens of Mallard (Anas platyrhynchos), and most of the specimens of Black-headed Gull), oribatid mites were absent, except for some single specimens of Prostigmata, Gamasidae, and Collembola, which are


common soil microarthropods. On the whole, the number and diversity of oribatid mites in the feathers of waterfowl or birds related to aquatic ecosystems were low. The most frequent species were Tectocepheus velatus (found in 16 bird species), Oppiella nova, Oppia splendens (found in 12 bird species), Carabodes areolatus, Zygoribatula exilis, Phthiracarus sp. Steganacarus striculus (found in six bird species), and Achipteria coleoptrata (found in five bird species). Some findings are interesting from the faunistic point of view. For example, Odontocepheus elongatus, quite a rare Mediterranean oribatid species in Russia, was found in the feathers of Cormorant; Eremaeus fovelatus, a rare Arctic species, was found in the feathers of Kittiwake (Rissa tridactyla); Caleremaeus monilipes, a rare West European species, was found in the feathers of Red-throated Diver (Gavia stellata) and Kittiwake. Apparently, this group of birds requires special studies on the carriage of oribatid mites in their feathers. However, it is evident that oribatid mites are constant inhabitants there and that they are quite diverse. That should be taken into consideration while investigating the biogeography of soil microarthropods. The data obtained are, however, insufficient for a thorough biogeographical analysis, though it allows to discuss some interesting factors. First, it is the constant presence of such hygrophilous primitive species as Paleacarus histricinus, Liochthonius sp., Brachychthonius sp., Hypochthonius rufulus, Nothrus silvestris, and Malaconothrus egregious in the feathers of waterfowl and birds related to aquatic ecosystems. Second, the fact that Chamobates lapidarius, Minunthozetes semirufulus, Caleremaeus monilipes, and Odontocepheus elongatus, which are rare in the European part of Russia, but common in Central and West Europe, have also been recorded in the feathers of the above-mentioned birds. It is possible that those mites penetrated the plumage of birds in their wintering sites. Abundance of oribatid mites (mature specimens, nymphs, and larvae) in the feathers of Red-throated Diver, a migrant species from the Kostroma Province, should also be considered. Red-throated Diver is a species with a circumpolar breeding range and migrating to the southern seas (including the Black Sea and the Caspian Sea). The presence of abundant and diverse communities or assemblages of oribatid mite species in the feathers of seasonal migrants during their passage period and transmission of these mites at the distances from hundreds to thousands of kilometers can be taken for granted. Third, the collection of oribatid mites in bird feathers related to aquatic ecosystems is quite peculiar compared with similar groups in birds of other ecological groups.

35

Hemiedaphic species of oribatid mites (Tectocepheus velatus, Zygoribatula exilis, Minunthozetes pseudorufulus, etc.) and species of epifauna (Xenillus tegeocranius, Ceratoppia bipilis, Achipteria coleoptrata, Camisia sp., Steganacarus striculus, Phthiracarus sp., Galumna sp., etc.) are continuously present in the feathers of waterfowl. Sometimes they even reproduce there, which can be proved by the presence of larvae and nymphs in the feathers. However, inhabitants of soil apertures, one of the most massive groups among the soil inhabitants (Oppia, Oppiella, Suctobelbella, and Quadroppia), were found only singly, and their preimaginal forms were not found at all. The greatest diversity of oribatid mites was recorded in birds during their breeding period and one or two months after it: from two to three species in a specimen. However, in most cases, 12 species of oribatid mites were found in the feathers of 46% of investigated birds and in the feathers 14% of birds they were not found at all. The latter were mostly nestlings and fledglings. Some interesting findings of quite a diverse fauna of collembolans were revealed in the feathers of 20% of birds (13 and, sometimes, 56 or even 14 specimens per bird). Among them were Prostigmata mites (mostly Scutacaridae and Pyemotidae) found in 20% of birds. In 10% of birds, free-living Gamasidae and, Rhodacaridae, which are especially characteristic of soils and nests of birds, were found. We have revealed unambiguously that oribatid mites, as well as other groups of microarthropods, including collembolans, constantly inhabit bird feathers. In addition we found many immature forms of oribatid mites (especially Trhypochthonius tectorum, Tectocepheus velatus, and Diapterobates notatus, Carabodes marginatus), i.e. larvae and nymphs. On the basis of these findings we may conclude that some species of oribatid mites constantly inhabit bird feathers, which is not a temporary phenomenon occurring during the breeding or nestling-raising periods. The time of material collection on the coast of the Barents Sea was not optimal, because most of the birds had moulted and therefore a large number of small parasites and symbionts inhabiting their feathers were lost. We did not find any oribatid mites in the feathers of Arctic Skua. However, they contained quite a great number of collembolans and Prostigmata mites. Caleremaeus monilipes, Eremaeus fovelatus, Carabodes labirinticus, Tectocepheus velatus, Liebstadia similis, Minunthoretes semirufus, Phthiracarus logneus, Parachipteria punctata, and Pilogalumna tenuiclava were found in the feathers of gulls from the coast of East Murman. Eremaeus sp. and Carabodes subarcticus were found on the skins of Guillemots (Uria aalge) together with the larvae of Diptera, Gamasidae, and small beetles.

36

Oppia fallax and Carabodies areolatus as well as nymphs of several species of oribatid mites were collected from a Common Eider (Somateria mollissima). Thus, oribatid mites, collembolans, insect larvae, Gamasidae, and Prostigmata mites are constant inhabitants in the feathers of Arctic birds. These birds may be also considered as a major factor determining distribution of microarthropods in the most remote islands of the Arctic Ocean. To confirm this conclusion, we may present another, though indirect, evidence: analysis of the microarthropod fauna found in the nests of birds in the Arctic (it differs significantly from the inhabitants of the soils of tundra and the Arctic) revealed that it was transmitted to the Arctic islands by birds. We had at our disposal nests of several bird species from Novaya Zemlya and Northern-Western part of Cola peninsula. In a nest of Bewicks Swan (Cygnus bewickii) built in Belushye village, we found Liochthonius sp., Tectocepheus velatus, Oppia unicarinata, Quadroppia quadricarinata, Oppia minus, Oribatula tibialis, Diapterobates notatus, and Eniochthonius minutissimus. Eremaeus oblongus, Chamobates sp., Ch. lapidaricus, Tectocepheus velatus, Carabodes subarcticus, and Eniochthonius minutissimus together with the nymphs of Camisiidae and Damaeudae were found in a nest of Iceland Gull (Larus glaucoides). The latter mite family, as well as the species Eniochthonius minutissimus and Oppia minus, had not been found in the Arctic before, and their presence there may be related to their transmission by birds. Nests of Common Eiders and gulls contained an entire community of oribatid mites that were absent in the surrounding soils of tundra, where Krivolutsky (1966) found 56 species of oribatid mites in 1966: Eniochthonius minutissimus, Nothrus silvestrus, Nanhermannia coronata, Hermannia reticulata, Suctobelbella subcornigera, Carabodes forsslundi, Caleremaeus monilipes, Chamobates lapidarius, Autogneta longilamellata, Oribella castanea, Gustavia microcephala, Ameronothrus lineatus, Minunthozetes pseudofusiger, Minunthozetes semirufus, Fuscozetes fuscipes, Parachipteria punctata, Eupelops plicatus, and Pilogalumna tenuiclava (19 species in all). Most of them, as we may see from the above-presented data, were found in bird feathers as well, but they were absent in the soils of tundra. Thus, an assumption could be made that those oribatid mites were transmitted by birds into their nests and then, quite possibly, to the soil. We can undoubtedly conclude that birds are transmitters of microarthorpods. It is very likely that they bring them to Northern-Western part of Cola peninsula not from the central or southern parts of Russia, but from the West European coastal regions. We have come to


this conclusion considering that such species as Carabodes forsslundi, Licneremaeus licnophorus, Caleremaeus monilipes, Minunthozetes semirufus, and Oribella castanea, which are very common in England, Holland, and Sweden, but very rare in the European north of Russia, have been found there. Barren soils of Arctic islands are poor in collembolan species. There the ground is ice-covered during the major part of the year. Nevertheless, oribatid mites and other microarthropods are found in the soils of the most remote Arctic islands. Their species composition is fairly diverse there, so their distribution cannot be accidental. According to MacArthurs insular biogeography, the rate of such a process is well-known. It is difficult to understand how soil microarthorpods have reached remote Arctic islands located hundreds and thousands of kilometers away from the continents, surmounting vast obstacles of water and ice of the Arctic Ocean. This could have occurred just as a result of microarthropod distribution, and the latter just aided by birds capable of covering long distances (thousands of kilometers or more) between islands and continents during their passages. The phenomenon of oribatid mites inhabiting bird feathers enables to explain a lot of unknown issues concerning the biogeography of oribatid mites and other soil microarthropods and to reveal mechanisms of their distribution on the islands of the Arctic Ocean. We investigated the feathers of birds shot in the Spitsbergen archipelago, in the environs of Barentsburg, Ice and Green fiords. The birds were shot by the staff members of Murmansk Marine Biological Institute in JulyAugust 2001, during their expeditions to the Spitsbergen archipelago. 30 specimens of the following five bird species were studied as possible transmitters of microarthropods: Fulmar (Fulmarus glacialis), Little Auk (Alle alle), Kittiwake, and Arctic Tern (Sterna paradisaea). Thirteen species of oribatid mites (19 specimens) were found in the above-mentioned birds: Steganacarus striculus, Nothrus palustris, Platynothrus punctatus, Malaconothrus egregius, Oppia sp., Oppia translamellata, Oppia splendens, Oppiella nova, Suctobelbella sp., Suctobelbella subcornigera, S. hammeri, Liebstadia similes, and Parachipteria punctata. The feathers of birds shot on Novaya Zemlya in 2002 were inhabited by the following oribatid mite species: Suctobelba sp., Suctobelbella hammeri, Oppia splendens, O. unicarinata, and Quadroppia quadricarinata. In addition, they contained Acaridia, Collembola, and Prostigmata. All of them had been absent in the soils of Novaya Zemlya before (Gilyarov & Krivolutsky 1975; Karppinen & Krivolutsky 1982; Krivolutsky & Kalyakin 1993). In all bird species, except Arctic Tern, adult oribatid mites were found. In the feathers of terns, only nymphs of oribatid mites occurred. The most primitive and large


oribatid mites (Platynothrus and Nothrus) were found only in their nymphal form. Only three species of oribatid mites from our list had been described earlier from the soils of Spitsbergen: Suctobelbella subcornigera, Oppiella nova, and Oppia translamellata. The other had been recorded from further south: Scandinavia, Kola Peninsula, etc. (Karppinen & Krivolutsky 1982; Krivolutsky 1995). Liebstadia similes and Parachipteria punctata were found in the feathers of Kittiwake in the coastal area of the Barents Sea near Murmansk. All Gamasidae found in bird feathers are soil predators of different families of the non-parasitic group. In the feathers of Fulmar and Kittiwake, also Prostigmata and Trombidiidae of the family Bdellidae, recorded earlier in the soils of Spitsbergen, were detected. In the feathers of all the investigated bird species, Acaridia and Collembola were revealed. Oribatid mites in the feathers of waders and crakes This group of birds is especially important for our studies because of its extremely distant migrations. Its role in the transmission of oribatid mites may be very significant. Despite of our possession of quite a limited number of species and specimens of sandpipers, the material collected enabled estimation of principle peculiarities of the transmission of oribatid mites by species of this group of birds. A total of 58 specimens of birds belonging to 16 species was investigated: Woodcock (Scolopax rusticola), Greenshank (Tringa nebularia), Redshank (Tringa totanus), Dunlin (Calidris alpina), Green Sandpiper (Tringa ochropus), Ringed Plover (Charadrius hiaticula), Marsh Sandpiper (Tringa stagnatilis), Ruff (Philomachus pugnax), Terek Sandpiper (Xenus cinereus), Oystercatcher (Haematopus ostralegus), Common Snipe (Gallinago gallinago), Little Stint (Calidris minuta), Purple Sandpiper (Calidris maritima), Red-necked Phalarope, Lapwing (Vanellus vanellus), and Corncrake (Crex crex). No oribatid mites were found in the feathers of 15 specimens of those bird species; the rest specimens contained 71 specimens of oribatid mites (49 mature individuals and 22 larvae and nymphs) belonging to 38 species. In addition to oribatid mites, bird feathers contained 15 collembolans, seven Gamasidae, nine Prostigmata, five Acaridia, and four larvae of Diptera. In some bird species shot in small numbers (13) just single specimens of oribatid mites were detected: Liochthonius sellnicki (found in Marsh Sandpiper); Suctobelba trigona (Ruff); Chamobates lapidarius (Common Snipe); Suctobelba trigona (Greenshank); Oppiella nova, Conchogneta delacarlica, Tectocepheus velatus (Little Stint); Hydrozetes palustris (in the contents of the stomach of Red-necked

37

Phalarope); Gymnodamaeus bicostatus, Chamobates cuspidatus and Punctoribates punctum (Lapwing); Oppiella nova and Oppia ornata (Terek Sandpiper); single specimens of Green Sandpiper and Oystercatcher contained no oribatid mites at all, whereas Redshank (three specimens) just larvae. In the rest five species of birds, quite a diverse population of oribatid mites was present (Table 5). Of 30 mite species, however, 22 were found just singly in the feathers of 37 investigated birds; six oribatid species were found twice; and only two species (Oppiella nova and Oppia unicarinata) were detected in three bird species. Table 5 shows that as to numbers of oribatid mites found in the feathers of waders, they are very scarce. As to species, their collection is accidental. However, they are found constantly and, relatively, in large numbers. It should be noted that only nymphs and larvae of such large representatives of oribatid mites as Ceratoppia bipilis, Nothrus palustris, and Diapterobates notatus were discovered. Some species of oribatid mites found in the feathers of waders were very interesting from the faunistic point of view. This is true of the findings of Sphaerochthonius splendidus and Licnodamaeus pulcherrimus in the feathers of a Corncrake from the Homel Province. These are the species confined to the Mediterranian region, which had not been found in the south of Belarus and the north of Ukraine in natural ecosystems before. In the feathers of Lapwing from the Moscow Province, mature Gymnodamaeus bicostatus was found. Earlier we used to find only nymphs of this species. All the findings of oribatid mites in the waders from the North deserve attention, because most of them were found in that region for the first time. This is true of oribatid mites collected from the feathers of Purple Sandpiper from the Spitsbergen archipelago, Ringed Plover, and Dunlin from Northern-Western part of Cola peninsula (the northern coast of the Kola Peninsula; Table 5). The findings of Hypochthonius rufulus, Tropacarus carinatus, and Chamobates schuetzi are especially interesting, because they had not been found so far to the north during earlier pedobiological-zoological studies. Fosseremaeus laciniatus, a rare species in the Moscow Province, also is an interesting finding recorded in the feathers of Corncrake. The collected study material about oribatid mites from the nests of waders was rather sparse. We investigated three nests of Lapwing and four nests of Black-winged Stilt (Himantopus himantopus) from the Krasnodar Province as well as one nest of Ringed Plover from the Odessa Province. Those birds build their nests on the ground. In the nests of Lapwing, we found one specimen of Oppia minus and Malaconothrus egregious at

38


Table 5. Oribatid mites in the feathers of some waders and the Corncrake.

Oribatid mite species Number of birds Palaeacarus hystricinus Hypochthonius rufulus Sphaerochthonius splendidus Tropacarus carinatus Nothrus palustris Platynothrus punctatus Trhypochthonius sp. Malaconothrus egregius Hermannia reticulata Licnodamaeus pulcherrimus Belba sp. Fosseremaeus laciniatus Ceratoppia bipilis Carabodes areolatus Tectocepheus velatus Oppia unicarinata Oppia neerlandica Oppiella nova Conchogneta delacarlica Suctobelbella hammeri Suctobelbella acutidens Suctobelbella sp. Banksinoma lanceolata Zygoribatula exilis Liebstadia similes Diapterobates notatus Chamobates schuetzi Punctoribates punctum Achipteria sp. Parachipteria punctata NN Oribatei, LL Oribatei Total number of species

Corncrake 5 1 1 1 1 1 1 1 1 1 1 1 3 11

Ringed Plover 3 1 1 1 1 1 1 1 5 7

a time and 90 specimens of collembolans. In the nests of Black-winged Stilt, we discovered one specimen of Carabodes areolatus, four specimens of Oppiella nova, and 169 collembolans. The nest of Ringed Plover was empty. It is likely that the main sources of the microarthropods inhabiting bird feathers were not nests, but the superficial layers of soil. Thus, the fauna of oribatid mites in the feathers of waders and Corncrake can be distinguished for its originality: their population is small in numbers, species composition accidental. Other microarthropods are even more numerous. For such a small sample of birds,

Bird species Woodcock

Dunlin

9 1 1 1 1 1 1 1 3 7

10 1 1 1 1 1 1 1 1 5 4 9

Total number Purple of specimens Sandpiper 10 37 1 1 1 1 1 2 1 1 1 1 1 1 1 1 2 1 2 1 3 1 1 3 1 1 2 1 2 1 1 1 1 1 1 1 1 6 15 6 30

the number of oribatid mites is, however, quite significant, and the role of waders in the distribution of this group of mites is undeniable. Small numbers of microarthropods in the feathers of waders and their absence in 27% of birds show that these birds are capable of cleaning effectively their feathers from symbionts and ectoparasites. This is of great importance for waders building nests on the ground. It is also possible that dwelling and foraging in shallows of water bodies and a constant contact with water enables these birds to clean their plumage even more effectively from microarthropods.

39


Oribatid mites in the feathers of Galliformes Galliformes, birds closely related to the soil, are also interesting from this point of view. Oribatid mites were found though singly, in the feathers of domestic hens reared in the aviaries of pheasants and in Japanese

Quails (Coturnix coturnix japonica). In the feathers of domestic hens, Heminothrus targionii, Adoristes poppei, Oppia unicarinata, Oppiella nova, and Diapterobates notatus were detected. In those of two wild pheasants, Suctobelba trigona was found. In the

Table 6. Oribatid mites in the feathers of Galliformes.

Oribatid mite species Total number of bird specimens Hypochtonius rufulus Phthiracarus globosus Steganacarus striculus Camisia sp. Nanhermannia coronata Hermannia reticulata Hermannia gibba Licnodamaeus pulcherrimus Epidamaeus kamaensis Hypodamaeus riparius Eremobelba geographica Eremaeus oblongus Adoristes ovatus Carabodes areolatus Carabodes marginatus Odontocepheus elongates Tectocepheus velatus Autogneta willmanni Oppia falax Oppia minus Oppia splendens Oppia unicarinata Oppiella nova Quadroppia quadricarinata Suctobelba trigona Suctobelbella subcornigera Oribella paoli Oribatula tibialis Zygoribatula exilis Liebstadia humerata Liebstadia similis Scheloribates laevigatus Diapterobates notatus Trichoribates incisellus Chamobates lapidarius Minunthozetes pseudofusiger Mycobates sp. Punctoribates punctum Oribatella berlesei Parachipteria punctata Total number of oribatid species

Hazel Grouse 28 2 2 1 1 3 1 1 2 1 1 1 3 2 1 1 3 1 1 13 1 1 10 2 1 1 2 3 1 2 29

Bird species Black Grouse 6 5 1 1 1 2 1 2 14 5 2 2 1 12

Capercaillie 6 4 1 1 5 1 1 2 7 3 3 1 5 1 1 1 1 9 1 1 1 20

Total 40 11 4 2 1 3 1 1 1 1 4 1 1 1 8 1 2 1 1 4 1 1 5 34 4 1 3 1 20 1 1 4 1 1 1 5 12 1 1 1 4 40

40

plumage of a Willow Grouse (Lagopus l. lagopus) from the Gdansk peninsula, Oppia unicarinata was recorded. No oribatid mites occurred in all the 10 investigated Grey Partridges (Perdix perdix) from the south of Ukraine and the Rostov Province. It should be taken into account, however, that oribatid mites were collected during a long drought. Abundant and diverse fauna of oribatid mites was recorded in the feathers of Hazel Grouse, Black Grouse (Tetrao tetrix), and Capercaillie (Tetrao urogallus) from taiga: 152 mature oribatid mites of 40 species were collected from 40 specimens of birds (Table 6). However, oribatid mites were absent in the feathers of five birds (12.5%). Composition of oribatid species in the feathers of Galliformes from taiga was quite similar. This can be explained by the fact that they were collected in one limited area the environs of the village Manturovo in the Kostroma Province. In the European part of taiga, the fauna of oribatid mites has been investigated quite thoroughly. Thus, the findings of a large number of southern species (common in the zonal types of soils to the south of broad-leaved forests) in the taiga are especially interesting to us. They are Licnodamaeus pulcherrimus, Odontocepheus elongates, Autogneta willmanni, Liebstadia humerata, and Eremobelba geographica. Since large Galliformes are residents, it is very unlikely that they are transmitters of oribatid mites. These birds could have collected the above-mentioned oribatid species from the patches of soil directly open to the sunlight, or from the nests. Both the premises should be investigated by oribatologists in the nearest future. Other bird species During the investigation some orders of birds were obtained in single specimens. The investigation showed that their plumage was inhabited by the following species of soil microarthropods: Cuckoo (Cuculus canorus) (Licnodamaeus pulherrimus); Black Woodpecker (Dryocopus martius) (Tectocepheus velatus, Quadroppia quadricarinata, Chamobates lapidarius, Platynotthrus peltifer (N2), and Nothrus palustris (N2)); Great Spotted Woodpecker (Dendrocopus major) (Oppiella nova); Three-toed Woodpecker (Picoides tridactylus) (Carabodes areolatus); Green Woodpecker (Picus viridis) (Oppiella nova and Carabodes areolatus); Nightjar (Caprimulgus europaeus) (four specimens) (Oppiella nova Chamobates cuspidatus, Tectocepheus velatus, and Hermannia sp.); Demoiselle Crane (Anthropoides virgo) (one specimen from Kalmykia) (Hypochthonius rufulus, Tectocepheus velatus, Carabodes femoralis, Chamobates cuspidatus, Tropacarus carinatus, Tropacarus pulcherrimus,


Oribotritia loricata, Steganacarus applicatus, Phthiracarus logneus, and Phthiracarus globosus). Rock Dove (Columba livia) (159 specimens from the Moscow Province) were also investigated for oribatid mites. About 30 oribatid mites belonging to 16 species (Liochthonius sellnicki, Trhypochtonius tectorum, Trhypochthoniellus excavatus, Malaconothrus egregius, Nothrus palustris (LL), Tectocepheus velatus, Carabodes areolatus, C. marginatus (imago, LL larvae, N1 nymph of first stage, N2 nymph of second stage, N 3 nymph of third stage), Suctobelbella subcornigera, Suctobelba trigona, Oppia ornata, O. minus, O. unicarinata, O. tuberculata, Oppiella nova, and Oribatula tibialis) were disclosed in their feathers. The abundance of oribatid mites in the nests of doves was very low: just nymphs of Tectocepheus velatus, Diapterobates notatus, and Suctobelba sp. were found (in total, five specimens).

REFERENCES Behme, R.L. and Flint, V.E. 1994. Dictionary of animal names in five languages. Birds. Latin, Russian, English, German, French. Moscow: Russky Yazyk, Russo. [Áåìå, Ð.Ë., Ôëèíò, Â.Å. 1994. Ïÿòèÿçû÷íûé ñëîâàðü íàçâàíèé æèâîòíûõ. Ïòèöû. Ëàòèíñêèéðóññêèé-àíãëèéñêèé-íåìåöêèé-ôðàíöóçñêèé. Ìîñêâà: Ðóññêèé ÿçûê, Ðóññî.] Borisova, V.I. 1968. On the knowledge of the fauna of Sand Martin (R. riparia), House Martin (D. urbica) and Swallow (H. rustica) TASSR. Natural resources of the VolgaKama region. Animal kingdom: 162179. [Áîðèñîâà, Â.È. 1968. Ê ïîçíàíèþ ôàóíû ãíåçä áåðåãîâîé (R. riparia), ãîðîäñêîé (D. urbica) è äåðåâåíñêîé (H. rustica) ëàñòî÷åê ÒÀÑÑÐ. Ïðèðîäíûå ðåñóðñû Âîëæñêî-Êàìñêîãî êðàÿ. Æèâîòíûé ìèð: 162179.] Borisova, V.I. 1969. On the knowledge of the fauna in the nests of some bird species inhabiting the shore of Kuybishev reservoir. Issues of the formation of the littoral biocenoses of reservoirs: 125140. [Áîðèñîâà, Â.È. 1969. Ê ïîçíàíèþ ôàóíû ãíåçä íåêîòîðûõ âèäîâ ïòèö ïîáåðåæüÿ Êóéáûøåâñêîãî âîäîõðàíèëèùà. Âîïðîñû ôîðìèðîâàíèÿ ïðèáðåæíûõ áèîöåíîçîâ âîäîõðàíèëèù: 125140.] Borisova, V.I. 1978. On the structure of nest-hole censuses of swallows. Parasitology 12 (5): 337382. [Áîðèñîâà, Â.È. 1978. Ê ñòðóêòóðå ãíåçäîâî-íîðíûõ öåíîçîâ ëàñòî÷åê. Ïàðàçèòîëîãèÿ 12 (5): 337382.] Erik, V.V. and Glukhova, V.M. 1974. About the parasite fauna of the nests in the Curonian Spit. Parasitological collection of the Institute of Zoology of the Academy of Sciences of the USSR 26: 161183. [Ýðèê, Â.Â.,

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Ãëóõîâà, Â.Ì. 1974. Î ïàðàçèòîôàóíå ïòè÷üèõ ãíåçä ðàéîíà Êóðøñêîé êîñû. Ïàðàçèòîëîãè÷åñêèé ñáîðíèê Çîîëîãè÷åñêîãî Èíñòèòóòà Àêàäåìèè Íàóê ÑÑÑÐ 26: 161183.] Fain, A., Vangeluwe, D. and Wauthy, G. 1993. Observations on mites inhabiting nests of Bubo bubo (L.) (Strigiformes, Strigidae) in Belgium. Belgian Journal of Zoology 123 (11): 326. Gembitsky, A.S. 1970. Mites nest inhabitants in Belarus. II Conference on Acarology. Abstracts 1: 132133. [Ãåìáèöêèé, À.Ñ. 1970. Êëåùè îáèòàòåëè ãíåçä â Áåëîðóññèè. II àêàðîëîãè÷åñêîå ñîâåùàíèå. Òåçèñû äîêëàäîâ 1: 132133.] Gembitsky, A.S. and Andreichikova, E.I. 1969. Oribatid mites inhabitants of bird nests in the territory of Belarus. Issues of soil zoology: Materials of the III All-Union Conference: 5051. [Ãåìáèöêèé, À.Ñ., Àíäðåé÷èêîâà, Å.È. 1969. Ïàíöèðíûå êëåùè îáèòàòåëè ãíåçä ïòèö íà òåððèòîðèè Áåëîðóññèè. Ïðîáëåìû ïî÷âåííîé çîîëîãèè: Ìàòåðèàëû III Âñåñîþçíîãî ñîâåùàíèÿ: 5051.] Gilyarov, M.S. and Krivolutsky, D.A. (eds) 1975. The key to soil mites: Sarcoptiformes. Moscow: Nauka. [Ãèëÿðîâ, Ì.Ñ., Êðèâîëóöêèé, Ä.À. (ðåä.) 1975. Îïðåäåëèòåëü îáèòàþùèõ â ïî÷âå êëåùåé: Sarcoptiformes. Ìîñêâà: Íàóêà.] Hammer, M. 1965. Are low temperatures a species preserving factor? Illustrated by oribatid mite Mucronothrus nasalis (Willmann). Acta University Lundensis, Lund 2 (2): 110. Karppinen, E. and Krivolutsky, D.A. 1982. List of oribatid mite (Acarina, Oribatei) of Northern palaearctic region. I. Europe. Acta Entomologica Fennica 41: 118. Krivolutsky, D.A. (ed.) 1995. Oribatid mites. Moscow: Nauka. [Êðèâîëóöêèé, Ä.À. (ðåä.) 1995. Ïàíöèðíûå êëåùè. Ìîñêâà: Íàóêà.] Krivolutsky, D.A. and Kalyakin, V.N. 1993. Microfauna of soils in the ecological control on Novaya Zemlya. Vol. 2. In: P.V. Bojarsky (ed.) Proceedings of the complex marine Arctic expedition, pp. 125131. Moscow. [Êðèâîëóöêèé, Ä.À., Êàëÿêèí, Â.Í. 1993. Ìèêðîôàóíà ïî÷â â ýêîëîãè÷åñêîì êîíòðîëå íà Íîâîé Çåìëå. Íîâàÿ Çåìëÿ. Ò. 2. Ïîä ðåäàêöèåé Ï.Â. Áîÿðñêîãî. Òðóäû Ìîðñêîé àðêòè÷åñêîé êîìïëåêñíîé ýêñïåäèöèè, ññ. 125131. Ìîñêâà.] Krivolutsky, D.A. and Lebedeva, N.V. 1999. Transmission of soil microarthropods by birds. Little Bustard 4: 23 24. [Êðèâîëóöêèé, Ä. À., Ëåáåäåâà, Í. Â. 1999. Ðàñïðîñòðàíåíèå ïî÷âåííûõ ìèêðîàðòðîïîä ïòèöàìè. Ñòðåïåò 4: 2324.] Lebedeva, N.V., Shakhab, S.V. and Aksenova, M.M. 1998. Composition of microarthropods from the nests of some Passeriformes. Caucasian ornithological bulletin 10:

8386. [Ëåáåäåâà, Í.Â., Øàõàá, Ñ.Â., Àêñåíîâà, Ì.Ì. 1998. Ñîñòàâ ìèêðîàðòðîïîä èç ãíåçä íåêîòîðûõ âèäîâ îòêðûòîãíåçäÿùèõñÿ âîðîáüèíûõ ïòèö. Êàâêàçñêèé îðíèòîëîãè÷åñêèé âåñòíèê 10: 8386.] Lebedeva, N.V., Shakhab, S.V., Aksenova, M.M. and Markitan, L.V. 1999. New data on the oribatid mites inhabiting bird nests. Caucasian ornithological bulletin 11: 118223. [Ëåáåäåâà, Í.Â., Øàõàá, Ñ.Â., Àêñåíîâà, Ì.Ì., Ìàðêèòàí, Ë.Â. 1999. Íîâûå ìàòåðèàëû ïî îðèáàòèäàì â ãíåçäàõ ïòèö. Êàâêàçñêèé îðíèòîëîãè÷åñêèé âåñòíèê 11: 118223.] Proctor, H. and Owens, I. 2000. Mites and birds: diversity, parasitism and coevolution. Tree 15 (9): 358364. Tryjanowski, P., Baraniak, E., Bajoczyk, R., Gwiazdowicz, D., Konwerski, S., Olsczanowski, Z. and Szymkowiak, P. 1999. Arthropods in Red-backed Shrike (Lanius collurio) nests. The Ring 21 (1): 70. Wasylik, A. 1971. Nest types and the abundance of mites. Ecologia Polska 19 (39): 689699. Wasylik, A. 1973. The mites (Acaroides) inhabiting the nests of the tree sparrow (Passer montanus). Ecologia Polska 21 (52): 869899. Yaroshenko, N.N. and Kharchenko, V.I. 1972. Oribatid mites (Acariformes, Oribatei) in the nests of birds in the Donetsk Province. Bulletin of Zoology 3: 2023. [ßðîøåíêî, Í.Í., Õàð÷åíêî, Â.È. 1972. Ïàíöèðíûå êëåùè (Acariformes, Oribatei) ãíåçä ïòèö íà òåððèòîðèè Äîíåöêîé îáëàñòè. Âåñòíèê çîîëîãèè 3: 2023.]

ORIBATIDINËS

ERKËS

(ORIBATEI, ACARIFORMES) NEÞVIRBLINIAI

PAUKÐÈIØ PLUNKSNOSE:

D.A. Krivolutsky, N.V. Lebedeva

SANTRAUKA Ðio straipsnio autoriai patvirtina, kad paukðèiai perneða oribatidines erkes (Oribatei). Tirta 150 rûðiø paukðèiai (2100 vienetø) ið skirtingø ekologiniø grupiø. Jø plunksnose aptiktos 180 rûðiø oribatidinës erkës. Kai kurios ið jø paukðèiø plunksnose gyvena nuolat. Autoriai mano, kad oribatidinës erkës minta ne paukðèiø odos dalelëmis, o grybais, esanèiais jø plunksnose, nes visos oribatidinës erkës yra mikofagai. Oribatidinës erkës buvo aptiktos ávairiose jø gyvenimo stadijose. Nustatyta, kad jos paukðèiø plunksnose gali daugintis. Tai, kad paukðèiai gali bûti ðiø mikroartropodø perneðëjais, gali turëti lemiamos reikðmës jø paplitimui itin atokiose vietovëse (Arktikoje, Okeanijos salose bei dykumø oazëse). Received: 24 January 2003 Accepted: 8 March 2004

42


Annex I. List of oribatid mites found in bird feathers

1. 2. 3.

4. 5. 6. 7. 8.

9. 10. 11. 12. 13. 14. 15. 16. 17. 18. 19.

20.

21. 22. 23. 24. 25.

Family PALAEACARIDAE Grandjean, 1932 Paleacarus hystricinus Trägåtdh, 1932 Cormorant, Corncrake Paleacarus sp. Trägåtdh, 1932 Ferruginous Duck (Aythya nyroca), Common Gull Family HYPOCHTHONIIDAE Berlese, 1910 Hypochthonius rufulus C.L. Koch, 1835 Red-throated Diver, Great Egret, Goshawk, Black Grouse, Capercaillie, Hazel Grouse, Demoiselle Crane, Dunlin Family SPHAEROCHTHONIIDAE Grandjean, 1947 Sphaerochthonius splendidus (Berlese, 1904) Corncrake Family BRACHYCHTHONIIDAE Balogh, 1943 Brachychthonius berlesei Willmann, 1928 White-tailed Eagle Brachychthonius sp. Berlese, 1910 Little Egret, Goosander Liochthonius evansi (Forsslund, 1958) Mute Swan Liochthonius sellnicki (Thor, 1930) Mallard, Black Kite, Common Buzzard, Peregrine, Marsh Sandpiper, Rock Dove, Long-eared Owl Family PHTHIRACARIDAE Perty, 1841 Phthiracarus borealis (Trägåtdh, 1910) Mute Swan, Sparrowhawk, Kittiwake Phthiracarus globosus (C.L. Koch, 1841) Lesser Kestrel, Black Grouse, Capercaillie, Hazel Grouse, Demoiselle Crane Phthiracarus ligneus Willmann, 1931 Demoiselle Crane, Great Black-backed Gull Phthiracarus nitens (Nicolet, 1855) Ferruginous Duck Phthiracarus sp. Perty, 1841 Cormorant, Grey Heron, Mallard, White-tailed Eagle, Spotted Crake, Black-headed Gull, Great Black-backed Gull, Kittiwake Steganacarus applicatus (Sellnick, 1920) Demoiselle Crane Steganacarus striculus C.L. Koch, 1836 Red-throated Diver, Great Egret, Little Egret, Tufted Duck, Kestrel, Capercaillie, Hazel Grouse, Little Auk Tropacarus carinatus (C.L. Koch, 1844) Marsh Harrier, Common Buzzard, Lesser Kestrel, Demoiselle Crane, Ringed Plover Tropacarus pulcherrimus Berlese, 1885 Night Heron, Demoiselle Crane Family EUPHTHIRACARIDAE Jacot, 1930 Euphthiracarus sp. Ewing, 1917 Cormorant Family ORIBOTRITIIDAE Grandjean, 1954 Oribotritia loricata Rathke, 1799 Black-throated Diver, Red-necked Grebe, Demoiselle Crane Family NOTHRIDAE Berlese, 1896 Nothrus palustris C.L. Koch, 1839 Great Egret, Mute Swan, Marsh Harrier, Sparrowhawk, Corncrake, Purple Sandpiper, Kittiwake, Rock Dove, Long-eared Owl, Black Woodpecker Nothrus silvestris Nicolet, 1855 Red-throated Diver, Cormorant Family CAMISIIDAE Oudemans, 1900 Camisia biurus (C.L. Koch, 1839) Mute Swan, Herring Gull Camisia horrida (Hermann, 1804) Mute Swan Camisia spinifer (C.L. Koch, 1835) Mute Swan, Sparrowhawk, Kittiwake Camisia sp. von Heyden, 1826 Mute Swan, Hazel Grouse, Black-headed Gull


43

Annex I continued

26. 27. 28. 29. 30. 31. 32. 33. 34. 35. 36. 37. 38. 39. 40. 41. 42.

43. 44.

45. 46. 47. 48. 49. 50. 51.

Heminothrus targionii (Berlese, 1855) Platynothrus peltifer (C.L. Koch, 1939)

Domestic hen Great Egret, Grey Heron, Mute Swan, Goshawk, Sparrowhawk, Black-headed Gull, Kittiwake, Black Woodpecker Platynothrus punctatus (C.L. Koch, 1979) Fulmar, Purple Sandpiper, Little Auk Family TRHYPOCHTHONIIDAE Willmann, 1931 Trhypochthoniellus excavatus (Willmann, 1919) Rock Dove Trhypochthoniellus setosus (Willmann, 1928) Mallard Trhypochthonius tectorum (Berlese, 1896) Black-necked Grebe (Podiceps nigricolis), Cormorant, Marsh Harrier, Rock Dove, Short-eared Owl Trhypochthonius sp. Berlese, 1904 Corncrake Family MALACONOTHRIDAE Berlese, 1916 Malaconothrus egregius Berlese, 1904 Red-throated Diver, Cormorant, Grey Heron, Common Eider, Goshawk, Lesser Kestrel, Corncrake, Purple Sandpiper, Rock Dove, Long-eared Owl Trimalaconothrus tardus (Michael, 1888) Common Eider Family NANHERMANNIIDAE Sellnick, 1928 Nanhermannia coronata Berlese, 1913 Black-necked Grebe, Great Egret, Common Eider, Hazel Grouse Nanhermannia nana (Nicolet, 1855) Common Eider Nanhermannia sellnicki Forsslund, 1958 Great Egret Family HERMANNIIDAE Sellnick, 1928 Hermannia gibba (C.L. Koch, 1839) Black Grouse Hermannia reticulata Thorell, 1871 Mute Swan, Kestrel, Hazel Grouse, Dunlin, Great Black-backed Gull Hermannia scabra (C.L. Koch, 1839) Black Kite, Sparrowhawk Hermannia sp. Nicolet, 1855 Spotted Crake, Nightjar Family LIODIDAE Grandjean, 1954 Poroliodes farinosus (C.L. Koch, 1839) Ferruginous Duck, Sparrowhawk, Black-headed Gull Family GYMNODAMAEIDAE Grandjean, 1954 Gymnodamaeus bicostatus (C.L. Koch, 1835) Lapwing Family LICHNODAMAEIDAE Grandjean, 1954 Licnodamaeus pulcherrimus (Paoli, 1908) Black-throated Diver, Great Egret, Hazel Grouse, Corncrake, Cuckoo Family DAMAEIDAE Berlese, 1896 Belba sp. Heyden, 1826 Little Egret, Ferruginous Duck, Goshawk, Woodcock, Kittiwake Belba corynopus (Hermann, 1804) Common Eider, Black Grouse Epidamaeus sp. Bulanova-Zachvatkina, 1957 Great Egret Hypodamaeus riparius (Nicolet, 1855) Black Grouse, Hazel Grouse, Long-eared Owl Metabelba sp. Grandjean, 1936 Great Egret, Kestrel, Black-headed Gull, Shorteared Owl Metabelba pulverrulenta (C.L. Koch, 1839) Cormorant Family CEPHEIDAE Berlese, 1896 Cepheus cepheiformis (Nicolet, 1855) Sparrowhawk, Common Buzzard

44


Annex I continued

52. 53. 54. 55. 56. 57. 58. 59. 60. 61. 62.

63. 64. 65. 66. 67. 68. 69.

70. 71. 72. 73.

Family DAMAEOLIDAE Grandjean, 1954 Fosseremaeus laciniatus (Berlese, 1905) Corncrake Family EREMOBELBIDAE Balogh, 1961 Eremobelba geographica Berlese, 1908 Black-necked Grebe, Hazel Grouse Family EREMAEIDAE Sellnick, 1928 Eremaeus foveolatus (Hammer, 1952) Kittiwake Eremaeus oblongus (C.L. Koch, 1835) Common Buzzard, Kestrel, Hazel Grouse, Blackheaded Gull, Guillemot Eremaeus sp. C.L. Koch, 1836 Guillemot Family GUSTAVIIDAE Oudemans, 1900 Gustavia microcephala (Nicolet, 1855) Honey Common Buzzard Family LIACARIDAE Sellnick, 1928 Adoristes ovatus (C.L. Koch, 1839) Hazel Grouse Adoristes poppei (Oudemans, 1906) Spotted Crake, domestic hen Family PELOPPIDAE Balogh, 1943 Ceratoppia bipilis (Hermann, 1804) Corncrake, Ringed Plover, Herring Gull Family XENILLIDAE Woolley et Higgins, 1966 Xenillus tegeocranus Hermann, 1804 Cormorant Family CARABODIDAE C.L. Koch, 1837 Carabodes areolatus Berlese, 1916 Cormorant, Great Egret, Little Egret, Grey Heron, Mallard, Common Eider, Honey Common Buzzard, Sparrowhawk, Common Buzzard, Capercaillie, Hazel Grouse, Spotted Crake, Dunlin, Black-headed Gull, Guillemot, Rock Dove, Tengmalms Owl, Green Woodpecker, Three-toed Woodpecker Carabodes femoralis (Nicolet, 1855) Demoiselle Crane, Spotted Crake Carabodes forsslundi Sellnick, 1953 Cormorant, Common Eider, White-tailed Eagle Carabodes labyrinthicus (Michael, 1879) Lesser Kestrel, Herring Gull Carabodes marginatus (Michael, 1884) Red-throated Diver, Cormorant, Pond Heron, Shelduck, Capercaillie, Black-headed Gull, Rock Dove Carabodes subarcticus Trägåtdh, 1902 Guillemot Odontocepheus elongatus (Michael, 1879) Cormorant, Hazel Grouse Family TECTOCEPHEIDAE Grandjean, 1954 Tectocepheus velatus (Michael, 1880) Red-throated Diver, Cormorant, Great Egret, Little Egret, Grey Heron, Mute Swan, Shelduck, Mallard, Tufted Duck, Common Eider, Black Kite, Marsh Harrier, Goshawk, Common Buzzard, Peregrine, Kestrel, Hazel Grouse, Demoiselle Crane, Little Stint, Dunlin, Woodcock, Black-headed Gull, Herring Gull, Great Black-backed Gull, Common Gull, Kittiwake, Rock Dove, Nightjar, Black Woodpecker Family AUTOGNETIDAE Grandjean, 1960 Autogneta longilamellata (Michael, 1885) Cormorant Autogneta tragardhi Forsslund, 1947 Ferruginous Duck Autogneta willmanni (Dyrdowska, 1929) Common Buzzard, Hazel Grouse Conchogneta delacarlica (Forsslund, 1947) Cormorant, Great Egret, Mute Swan, Ferruginous Duck, Common Eider, Little Stint, Dunlin


45

Annex I continued

74. 75. 76. 77. 78. 79. 80. 81.

82. 83. 84.

85. 86.

87.

88.

89.

Family CALEREMAEIDAE Grandjean, 1965 Caleremaeus monilipes (Michael, 1882) Red-throated Diver, Kittiwake Family OPPIDAE Grandjean, 1954 Multioppia sp. Hammer, 1961 Spotted Crake Oppia falax (Paoli, 1908) Cormorant, Common Eider, Capercaillie, Hazel Grouse, Short-eared Owl Oppia maritima Willmann, 1929 Black-headed Gull, Long-eared Owl Oppia minus (Paoli, 1908) Black Grouse, Rock Dove, Long-eared Owl Oppia neerlandica (Oudemans, 1900) Ringed Plover Oppia ornata (Oudemans, 1900) Red-throated Diver, Mute Swan, Sparrowhawk, Coot, Terek Sandpiper, Rock Dove Oppia splendens (C.L. Koch, 1840) Red-throated Diver, Cormorant, Great Egret, Barnacle Goose, Mute Swan, Common Eider, Goshawk, Sparrowhawk, Common Buzzard, White-tailed Eagle, Red-footed Falcon, Hazel Grouse, Moorhen, Coot, Black-headed Gull, Great Black-backed Gull, Kittiwake, Short-eared Owl Oppia translamellata (Willmann, 1923) Fulmar, Black-headed Gull, Kittiwake Oppia tuberculata Bulanova-Zachvatkina, 1964 Rock Dove Oppia unicarinata (Paoli, 1908) Barnacle Goose, Mallard, Honey Common Buzzard, Marsh Harrier (Circus aeruginosus), Goshawk, Common Buzzard, Long-legged Common Buzzard, Willow Grouse, Black Grouse, Capercaillie, Hazel Grouse, Corncrake, Ringed Plover, Purple Sandpiper, Black-headed Gull, Whiskered Tern (Chlydonias hybridus), Rock Dove Oppia sp. C.L. Koch, 1836 Black-throated Diver, Great Egret, Mute Swan, Ferruginous Duck, Purple Sandpiper Oppiella nova (Oudemans, 1902) Red-throated Diver, Great Crested Grebe, Fulmar, Cormorant, Great Egret, Little Egret, Mute Swan, Shelduck, Mallard, Common Eider, Marsh Hawk, Marsh Harrier, Goshawk, Common Buzzard, Black Grouse, Capercaillie, Hazel Grouse, Ringed Plover, Terek Sandpiper, Little Stint, Purple Sandpiper, Woodcock, Black-headed Gull, Herring Gull, Great Black-backed Gull, Kittiwake, Rock Dove, Longeared Owl, Short-eared Owl, Tengmalms Owl, Nightjar, Green Woodpecker, Great Spotted Woodpecker, domestic hen Family QUADROPPIIDAE Balogh, 1983 Quadroppia quadricarinata (Michael, 1885) Cormorant, Great Egret, Grey Heron, Barnacle Goose, Common Eider, Stellers Eider, Goshawk, Capercaillie, Hazel Grouse, Black Woodpecker Family SUCTOBELBIDAE Jacot, 1938 Suctobelba trigona (Michael, 1888) Cormorant, Mute Swan, Mallard, Ferruginous Duck, Honey Common Buzzard, Common Buzzard, Pheasant (Phasianus colchicus), Hazel Grouse, Greenshank, Ruff, Kittiwake, Rock Dove Suctobelba sp. Paoli, 1908 Common Eider

46


Annex I continued

90. 91. 92. 93. 94.

95. 96. 97. 98. 99. 100. 101. 102. 103.

104. 105.

106. 107.

108. 109. 110. 111. 112. 113. 114. 115.

Suctobelbella acutidens (Forsslund, 1941) Suctobelbella hammeri (Krivolutsky, 1966)

Goshawk, Woodcock, Black-headed Gull Mute Swan, Common Eider, Stellers Eider, Dunlin, Kittiwake Suctobelbella subcornigera (Forsslund, 1941) Fulmar, Capercaillie, Rock Dove Suctobelbella tatarica (Krivolutsky, 1968) Mute Swan Suctobelbella sp. Jacot, 1937 Cormorant, Great Egret, Common Eider, Corncrake, Purple Sandpiper, Great Black-backed Gull Family THYRISOMIDAE Grandjean, 1954 Banksinoma lanceolata (Michael, 1885) Mute Swan, Coot, Woodcock, Black-headed Gull Banksinoma setosa Rjabinin, 1974 Common Eider Oribella paoli Oudemans, 1913 Mute Swan, Capercaillie Family LIMNOZETIDAE Grandjean, 1954 Limnozetes rugosus (Sellnick, 1923) Great Crested Grebe Family MICREREMIDAE Grandjean, 1954 Micreremus brevipes (Michael, 1888) Common Eider Family SCUTOVERTICIDAE Grandjean, 1954 Scutovertex minutus (C.L. Koch, 1836) Common Eider, Great Black-backed Gull Family HAPLOZETIDAE Grandjean, 1936 Peloribates longipilosus Csiszar, 1962 Swan Goose, Goshawk Family ORIBATULIDAE Thor, 1929 Oribatula tibialis (Nicolet, 1855) Cormorant, Great Egret, Ferruginous Duck, Black Grouse, Capercaillie, Hazel Grouse, Coot, Rock Dove Zygoribatula exilis (Nicolet, 1855) Great Egret, Mute Swan, Mallard, Ferruginous Duck, Kestrel, Capercaillie, Spotted Crake, Dunlin, Black-headed Gull, Great Black-backed Gull, Kittiwake Family PROTORIBATIDAE J.Balogh et P.Balogh, 1884 Liebstadia humerata Sellnick, 1928 Capercaillie Liebstadia similis (Michael, 1888) Goshawk, Black Grouse, Hazel Grouse, Purple Sandpiper, Great Black-backed Gull, Kittiwake Family SCHELORIBATIDAE Grandjean, 1953 Scheloribates laevigatus (C.L. Koch, 1835) Cormorant, Ferruginous Duck, Hazel Grouse, Blackheaded Gull, Great Black-backed Gull, Kittiwake Scheloribates latipes (C.L. Koch, 1844) Cormorant, Goosander, Black Kite, Common Common Buzzard Family PARAKALUMMIDAE Grandjean, 1936 Neoribates roubali (Berlese, 1910) Common Eider, Kestrel Family CERATOZERIDAE Jacot, 1925 Ceratozetes cisalpinus Berlese, 1908 Lesser Kestrel Diapterobates humeralis (Hermann, 1804) Honey Common Buzzard Diapterobates notatus (Thorell, 1871) Capercaillie, Woodcock, domestic hen Fuscozetes fuscipes (C.L. Koch, 1844) Honey Common Buzzard Melanozetes mollicomus (C.L. Koch, 1839) Kestrel Trichoribates incisellus (Kramer, 1897) Hazel Grouse Trichoribates trimaculatus (C.L. Koch, 1835) Grey Heron, Peregrine, Kestrel, Black-headed Gull, Long-eared Owl


47

Annex I continued

116. 117. 118. 119. 120. 121. 122.

123. 124. 125. 126.

127. 128. 129. 130.

Family CHAMOBATIDAE Thor, 1938 Chamobates cuspidatus (Michael, 1884) Cormorant, Common Eider, Goshawk, Demoiselle Crane, Lapwing, Ural Owl, Nightjar Chamobates lapidarius (Lucas, 1849) Mallard, Black Grouse, Capercaillie, Hazel Grouse, Snipe, Black-headed Gull, Black Woodpecker Chamobates schuetzi (Oudemans, 1902) Dunlin Family MYCOBATIDAE Grandjean, 1954 Minunthozetes pseudofusiger (Schweizer, 1922) Red-throated Diver, Cormorant, Great Egret, Shelduck, Goshawk, Capercaillie, Hazel Grouse, Common Gull Minunthozetes semirufus (C.L. Koch, 1835) Kittiwake Mycobates parmeliae (Michael, 1884) Great Crested Grebe, Capercaillie Punctoribates punctum (C.L. Koch, 1839) Cormorant, Mute Swan, Tufted Duck, Hazel Grouse, Corncrake, Lapwing, Great Black-backed Gull Family ORIBATELLIDAE Jacot, 1925 Oribatella berlesei (Michael, 1898) Capercaillie Family ACHIPTERIIDAE Thor, 1929 Achipteria coleoptrata (Linnaeus, 1758) Red-throated Diver, Cormorant, Great Egret, Mallard, Common Gull Achipteria sp. Berlese, 1885 Woodcock Parachipteria punctata (Nicolet, 1855) Cormorant, Mute Swan, Goshawk, Kestrel, Black Grouse, Capercaillie, Hazel Grouse, Ringed Plover, Dunlin, Black-headed Gull, Great Black-backed Gull, Kittiwake Family GALUMNIDAE Jacot, 1925 Galumna sp. Heyden, 1928 Great Egret, Mute Swan, Black-headed Gull Pergalumna nervosa (Berlese, 1914) Cormorant, Great Egret, Common Buzzard, Great Black-backed Gull Pilogalumna tenuiclava Grandjean, 1956 Great Black-backed Gull Nymphs of Oribatei mites Red-throated Diver, Great Crested Grebe, Cormorant, Little Bittern (Ixobrychus minutus), Little Egret, Mute Swan, Mallard, Tufted Duck, Common Eider, Corncrake, Ringed Plover, Redshank, Dunlin, Woodcock, Black-headed Gull, Herring Gull, Great Black-backed Gull, Kittiwake, Whiskered Tern, Arctic Tern