<oological j'ournal of-the Linnean Sociely ( 1984), 80: 3 6 9 4 2 0
The association of oribatid mites with lichens
EDMUND L. SEYD F.L.S.
Department of ~ o o l o g y ,The Manchester Museum, University of Manchester
AND
MARK R. D. SEAWARD F.L.S.
School of Environmental Science, University of Bradford, West Yorkshire
Received December 1982, accepted f o r publication August 1983
Oribatei (Acari, Cryptostigmata), are found in a variety of terrestrial habitats, and many are
associated with lichens; the relationship ranges from casual to highly dependent. Eighty-three
species associated with lichens have been surveyed, and a tentative classification, based on their
ecological requirements, is presented: Group A consists of species restricted to lichens as a biotope,
though occasionally occurring as accidentals in other habitats; Group B consists of species which
while preferring lichens as a habitat and feeding source are also adapted to existence on other plants
(though in some cases their immatures may be lichen-restricted); Group C consists of species which,
though frequently found on lichens, are equally common in other biotopes, particularly mosses, and
must be regarded as much more generalized in their feeding habits. Certain aspects of oribatidlichen specificity are discussed. The importance of oribatid-lichen associations from the point of
view of soil fertility and energetics is emphasized.
KEY \VORDS:
soild energetirs
Ecology-systematics
~
association list (Oribatei)
-
oribatid-lichen specificin
CONTENTS
Introduction
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Lichen-grazing by oribatid mites .
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Culture-feeding observations .
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List of Oribatei associated with lichens .
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A, B and C groups of lichen-associated oribatid
Group A species .
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Sup-group Aa specks .
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Group B speries
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Group C species .
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Ori ha tid-lichrn specificity .
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Discussion.
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Acknowledgements
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37 1
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INTRODUCT J
Oribatid mites are most commonly found in mosses, plant litter, humus and
soil but there is an increasing body of evidence which demonstrates that in
0024-4082/84/040369
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369
0 1984 T h r Linnean Sorirtv of London
370
E. L. SEYD AND M. R. D. SEAWARD
addition they are frequently found in lichen samples. They have also been
collected from fungi and algae, and from various parts of higher plants. In
general they can be said to be particularly abundant where decaying plant
material has sufficient moisture content and/or is penetrated by fungal mycelia
( Cloudsley-Thompson, 1958).
The present paper reviews current knowledge of the association of these mites
with lichens, with particular reference to their feeding habits. In general surveys
of the feeding habits of Oribatei, Hughes (1959), and Evans, Sheals &
Macfarlane (1961) suggest that the majority feed either by cropping the mycelia
and fruiting bodies of fungi, which grow on partially decaying debris, or they
feed directly on the debris itself. Kranz & Lindquist (1979) divide the Oribatei
into microphytophages which feed on the microflora (fungi, yeasts, bacteria) of
decaying plant material, and macrophytophages which feed on plant tissues,
including lichens, although Luxton ( 1972) believes that generally speaking the
food of macrophytophages needs to be softened and decayed by fungi before it
can be eaten. He also makes the point that while some Oribatei feed only
microphytophagously and others macrophytophagously, there are also many (if
not the majority of forms) which combine both these main types of feeding and
these he has termed panphytophages.
The importance of investigating the association of oribatid mites with both
micro- and macroflora lies in the fact that these mites play an important role in
soil energetics and soil fertility (Kates & Runkel, 1948; Waksman, 1952;
Ghilarov, 1963, 1965; Berthet, 1964; Witkamp & Crossley, 1966; Tarman, 1974;
Krivolutzky, 1976; Crossley, 1977; Reichle, 1977; Lebrun, 1979). The Oribatei
contribute to humus formation not only by the direct consumption and break
up of fallen plant material and of fungal-decayed litter, but also by eating and
distributing fungi and their spores (Forsslund, 1939; Witkamp, 1960; Kevan,
1962; Woodring & Cook, 1962; Macfadyen, 1964, 1968; Luxton, 1966a;
Wallwork, 1972a; Mitchell & Parkinson, 1976; Lindquist et a/., 1979). While the
generally accepted view is that the most important role of oribatid mites in soil
fertility is to act as secondary decomposers by conditioning the organic debris
for action by the primary microbial decomposers, Luxton (1972) believes that it
is now clear that some oribatids act as primary decomposers themselves.
Within the totality of degrading and decomposing plant biomass, lichens may
seem to occupy a relatively unimportant place. It has to be remembered,
however, that not only do lichens form a major part of the vegetational cover in
certain habitats (e.g. alpine lichen heaths) but that they also form an important
element of what Wallwork (1976) has termed ‘suspended soils’, using this term
in its broadest sense. Thus, together with fungi and mosses, lichens are present
on rocky outcrops and trees, and some of this ‘suspended soil’, as it degrades,
eventually becomes part of the main soil biomass itself.
Where lichens, together with moss, partially envelop dead wood, they clearly
hasten the decay of this substrate, though, of course, the oribatids present may
be feeding not only on the lichens and mosses but also on the decomposing wood
or, more usually, on the microflora and fauna associated with it. An interesting
study of the decaying log of an oil palm tree by Lasebikan (1977), in which he
sampled the arthropod fauna at three separate stages of the log’s decomposition,
showed that the most numerous group of arthropods in all three samples were
oribatid mites.
THE ASSOCIATION OF ORIBATID MITES WITH LICHENS
37 1
Furthermore, lichens are important to soil fertility through their role in
mineral element transfer. In a study of five forested ecosystems, Pike (1978)
found that lichens accounted for up to 10% of the annual above-ground
elemental turnover; the largest component being nitrogen. A large part of this
elemental transfer from lichens to other ecosystems takes place through leaching,
bacterial incorporation and non-cellular particle formation. However, Lawrey
( 1980) believes that lichen-herbivore activity, including that of oribatid mites,
can result in a significant transfer of essential mineral elements in terrestrial
ecosystems.
Many other invertebrates, of course, besides oribatid mites are associated with
lichens, and these associations have been reviewed by Gerson (1973) and by
Gerson & Seaward (1977).
LICHEN GRAZING BY ORIBATID MITES
However close the association of certain species of Oribatei with lichens may
be, this does not necessarily mean that these mites are feeding on the lichen
material itself. It is possible that they are using the lichens for shelter or for
feeding on the associated microflora. Any observations on the grazing of lichens
by Oribatei which have been made are therefore important.
Laundon ( 1967) found Ameronothrus maculatus (Michael) burrowing under the
cortex of a number of lichens and completely eating away the centre of the
thallus, leading him to believe that the survival of certain lichens of the
Caloplacetum heppianae DR. union was threatened in places by this mite. Schulte
(1976) reported groups of these mites grazing on the soredia of crustose lichens.
Gilbert (1976) found Xanthoria parietina (L.) Th.Fr. heavily infested by mites on
the Flannan Isles, West Scotland: Ameronothrus maculatus (Michael) and
Phauloppia lucorum (C. L. Koch) were both abundant, while a third mite,
Trichoribates trimaculatus (C. L. Koch) was present in small numbers. Grazing by
these mites was sufficient to have caused visible deterioration of the lichen
material. Gilbert (pers. comm.) also found an infestation by Ameronothrus
maculatus (Michael) on the very rare lichen Lecanora achariana A.L.Sm. in the Lake
District, and though he did not see the mites actually grazing on the lichen
material, the latter was damaged, presumably by the mites.
Phauloppia Lucorum (C. L. -Koch) is extremely common on lichens, the
association having been reported by numerous acarologists and lichenologists.
MacNeill (1966) investigated an infestation of lichens by this mite and observed
both adults and nymphs eating the lichen by breaking off pieces along the edges
of the thalli and excavating material from the flat surfaces.
Zopf (1907) considered that as many as 29 species of lichens, mostly the larger
foliose and fruticose kinds, were eaten by Oribatei, and he found many examples
of thallial lobes which were deformed by their depredations. Paulson &
Thompson (191 1, 1913) concluded that Oribatei must be regarded as the chief
pests of lichens and that they seemed to have a predilection for the mature
apothecia. Smith (1921) also noted that it is the apothecial discs and soredia
which are taken first, these evidently being the choicest portions; she quotes de
Lesdain as saying that the thallus round the base of the perithecia of Verrucaria
spp. is frequently eaten away, leaving the perithecia solitary and difficult to
determine.
372
E. L. SEYD AND M. R. D. SEAWARD
Sowter (1971) found several species of Parmelia from a small island on
Ullswater infested and eaten by oribatid mites; one gathering had been so badly
eaten that it was not possible to determine the lichen species reliably.
Grandjean (1948a, 1950a) has investigated the association of a population of
Camisia segnis (Hermann) with lichens in which it deposited its eggs. Excretion of
a substance by the female during oviposition caused the lichen thallus to swell
and grow over the egg. The larvae and nymphs fed on the lichen when it was
wet but were unable to do so when it was hard and dry. This is probably true
for microarthropods in general, and Metz (1971) found that the moisture
content of the substrate determines to a large degree the number of
microarthropods present. Smith (1921) also stressed the fact that mites only eat
lichens when they are moist, and Travt (1963) found that adults of
Ommatocepheus ocellatus (Michael), which feed within the thallus, are only able to
do so when the lichens had been moistened; as long as the lichens remained dry,
the mites stayed inactive in their burrows.
Grandjean (1951) also studied the habitats of Dometorina plantivaga (Berlese).
Cells which are excavated in the lichen thallus were found to contain either a
larva or a numph; their whole development takes place within the cell, and both
nymphs and adults feed on the lichen. Another species, Pirnodus detectidens
Grandjean, which feeds exclusively on lichens, occurs mostly on saxicolous forms
(Travt, 1963). Travt (1969a) observed the damage caused to lichens by two
mites, Dometorina plantivaga insularis Travt and Pirnodus soyeri Travt, relatively
large areas of the lichen surface being entirely destroyed. Jacot (1934) also
witnessed the feeding of the oribatid Scapheremaeus petrophagus (Banks) on
saxicolous lichens.
Finally, it is worth noting that Grandjean (1948a) believed that mites may
not only feed on lichens but may also hide under the thalli during winter, while
Peake & James (1967) suggested that the activities of Oribatei, as well as slugs
and snails, may be important in the dissemination of crustaceous lichens.
CULTURE-FEEDING OBSERVATIONS
Direct observation of the grazing of lichens by oribatids is difficult, but
selective feeding experiments in culture cells in the laboratory can provide useful
additional data on feeding habits. Such experiments, followed in some cases by
analysis of gut contents and faecal pellets, have been carried out by a number of
workers (Michael, 1884; Grandjean, 1947, 1948a, 1950a; Taberly, 1952/3;
Wallwork, 1958; Woodring & Cook, 1962; Woodring, 1963, 1965; Berthet,
1964; Rockett & Woodring, 1966; Luxton, 1966a, 1972;Jalil, 1969; Littlewood,
1969; Lebrun, 1970; Shereef, 1971, 1972, 1976; Saichuae, Gerson & Henis,
1972; Tadros, 1973, 1979; Nannelli, 1974; Haq & Prabhoo, 1976; Mitchell &
Parkinson, 1976; Anderson, 1977; Behan & Hill, 1978; Schatz, 1979).
Some interesting points have arisen from these studies. For example,
Littlewood (1969), who used algae as a laboratory food, made the discovery
that the algae had to become contaminated with micro-organisms before feeding
would take place. Wallwork (1958), who offered the mites he cultured a variety
of foods, and supported his observations by analysis of gut contents and faecal
material and by observations in the field,
found that some mites were “selective
__
feeders” and others “non-selective”. He also makes the point that larvae,
THE ASSOCIATION OF ORIBATID MITES WITH LICHENS
373
nymphs and adults may vary in their feeding habits and that therefore more
than one stage in the life cycle should be considered.
The significance of the results of selective feeding experiments in the
laboratory in order to determine feeding preferences of any particular species of
Oribatei is questionable. As Behan & Hill (1978) point out, it is difficult to
extrapolate from laboratory observations to actual feeding habits in the field.
Nevertheless, Luxton ( 1972) feels that while laboratory studies are necessarily
somewhat artificial, they have provided useful information. Lebrun (1979), on
the other hand, doubts whether laboratory studies on diet choice give much
reliable information as to nutritional specificity, (a view with which Mitchell
& Parkinson (1976) agree) and he feels that the best indicators are analyses of
gut contents and observations in the field. The trouble is, as Woodring (1963)
notes, that a given species in likely to feed on different foods in different habitats
and areas, so that one cannot always assume, with certain notable exceptions,
that the food in the gut is the only kind of food on which the species feeds or
that the species will be found only where that particular food exists.
We would certainly agree with Woodring & Cook (1962) that where a species
is always or nearly always found in a very restricted habitat, as is the case in
lichens, it can be assumed to feed exclusively or at least primarily on the
material which makes up just about 1 0 0 ~ of
o its environment. However, even in
such cases some caution is necessary, as Anderson (1975) found evidence that
oribatid mites exhibit marked seasonal changes in the foods they ingest, and
Bhattacharyya (1962) points out that during periods of scarcity oribatids may
be forced to live on less palatable material than that on which they normally
feed. Because of these and other studies it is difficult to dispute the view of Haq
& Prabhoo (1976) that the food habits of Oribatei, if they are suspected of being
selective feeders, have to be studied under both field and laboratory conditions
before it is absolutely safe to assign them to any particular category.
LIST OF ORIBATEI ASSOCIATED WITH LICHENS
The following list of lichen-associated species of Oribatei has been compiled
from the zoogeographical and ecological studies of many authors in which
catalogues of species are given. Such catalogues usually give the biotopes in
which the species have been collected and in which they are normally found.
These data have been supplemented with observations which have been made
on lichen-grazing by oribatid mites and by artificial feeding experiments carried
out on Oribatei reared in culture cells.
The mites which appear on this list can conveniently be divided into three
groups. Group A consists of species which are restricted to lichens as a biotope,
though they may occasionally be found in other habitats as ‘accidentals’. The
evidence in favour of some of these forms being lichen-restricted is, however,
based on only a single record or on very few records and such forms have been
placed in a sub-group Aa. Group B consists of species which, while preferring
lichens as a habitat and feeding-source, are also found living and feeding on
other plants, though in some cases their immatures may be lichen-restricted.
Finally Group C consists of those species, which though frequently found on
lichens, are also equally common in other biotopes, particularly mosses, and
must therefore be regarded as much more generalized in their feeding habits.
3 74
E. L. SEYD AND M. R. D. SEAWARD
The classification and taxonomy of the Oribatei used throughout this paper
follows that of Balogh (1972), and the nomenclature of lichens is mainly
according to Hawksworth, James & Coppins ( 1980).
Family Phthiracaridae
Phthiracarus crinitosimilis Willmann, 1939
Family Nothridae
Nothrus simplex Banks, 1895
Family Camisiidae
Camisia invenusta Michael, 1888
Camisia segnis (Hermann, 1804)
Family Trhypochthoniidae
Trhypochthonius cladonicola (Willmann, 1919)
Trhypochthonius montanus van der Hammen, 1956
Family Hermanniidae
Hermannia reticulata Thorell, 187 1
Hermannia scabra (L. Koch, 1878)
Family Liodidae
Liodes theleproctus (Hermann, 1804)
Poroliodes farinosus (C. L. Koch, 1840)
Family Plateremaeidae
Phereliodes wehnckei (Willmann, 1930)
Family Cepheidae
Conoppia palmicinctum (Michael, 1884)
Ommatocepheus ocellatus (Michael, 1882)
Family Eremaeidae
Eremaeus oblongus C. L. Koch, 1836
Tricheremaeus serratus (Michael, 1885)
Family Zetorchestidae
Strenzkea depilata TravC, 1966
Family Metrioppiidae
Macquarioppia striata (Wallwork, 1963)
Metrioppia helvetica Grandjean, 1931
Family Carabodidae
Carabodes areolatus Berlese, 1916
Carabodes labyrinthicus (Michael, 1879)
Carabodes marginatus (Michael, 1884)
Carabodes minusculus Berlese, 1923
Carabodes willmanni Bernini, 1975
Family Tectocepheidae
Tectocepheus sarekensis Tragsrdh, 1910
Family Oppiidae
Oppia crozetensis (Richters, 1907)
Family Autognetidae
Autogneta penicillum Grandjean, 1960
Family Hydrozetidae
Hydrozetes capensis Engelbrecht, 1974
Family Ameronothridae
Ameronothrus bilineatus (Michael, 1888)
Aa
Aa
B
B
B
Aa
C
C
C
C
C
C
B
C
B
C
C
C
C
B
C
B
B
C
C
B
B
C
THE ASSOCIATION OF ORIBATID MITES WITH LICHENS
Ameronothrus lapponicus Dalenius, 1963
Ameronothrus lineatus (Thorell, 1871)
Ameronothrus maculatus (Michael, 1884)
Family Podacaridae
Alaskozetes antarcticus (Michael, 1903)
Halozetes belgicae (Michael, 1903)
Halozetes crogetenjis (Richters, 1908)
Halozeles intermedius Wallwork, 1963
Halozetes marionensis Engelbrecht, 1974
Family Cymbaeremeidae
Cyrnbaeremaeus cymba (Nicolet, 1885)
Cymbaeremaeus marginalis Banks, 1895
Scapheremaeus petrophagus (Banks, 1906)
Family Micreremidae
Micreremus brevipes (Michael, 1888)
Family Licneremaeidae
Licneremaeus discoidalis Willmann, 1930
Family Scutoverticidae
Lamellovertex caelatus (Berlese, 1895)
Provertex delamarei TravC, 1962
Provertex mailloli TravC, 1964
Scutovertex minutus (C. L. Koch, 1836)
Family Zetomotrichidae
Ghilarouus hispanicus Subias & PQez-Iiiigo, 1977
Family Oripodidae
Cryptorebatula tazjhanensis Jacot, 1934
Pirnodus detectidens Grandjean, 1956
Pirnodus soyeri Travi., 1969
Family Oribatulidae
Dometorina plantivaga insularis TravC, 1969
Dometorina plantivaga plantivaga (Berlese, 1895)
Dometorina plantivaga saxicola Grandjean, 1951
Eporibatula gessneri Willmann, 1931
Lucoppia (Phauloppia) nemoralis Berlese, 1916
Maudheimia petronia Wallwork, 1962
Maudheimia wilsoni Dalenius & Wilson, 1958
Oribatula exudans TravC, 1961
Oribatula parisi Travi., 1961
Oribatula saxicola Halbert, 1920
Oribatula thalassophila Grandjean, 1935
Oribatula venusta Berlese, 1908
Phauloppia coineaui Travk, 1961
Phauloppia knoep@eri Travi., 1961
Phauloppia lucorum (C. L. Koch, 1841)
Phauloppia saxicola TravC, 1961
Ggoribatula frisiae (Oudemans, 1900)
agoribatula frisiae insularis TravC, 1961
Family Haplozetidae
Peloribater barbatus Aoki, 1977
Peloribates nishinoi Aoki. 1977
375
Aa
C
Aa
C
C
C
C
Aa
C
Aa
A
C
C
C
A
A
C
C
Aa
A
A
A
A
A
B
B
Aa
B
C
Aa
Aa
C
C
B
Aa
B
A
C
Aa
Aa
Aa
376
E. L. SEYD AND M. R. D. SEAWARD
Family Chamobatidae
Chamobates cuspidatus (Michael, 1884)
Family Ceratozetidae
Calyptozetes sarekensis (Trag2rdh, 1910)
Ceratozetes shiranensis Aoki, 1976
Diapterobates notatus (Thorell, 1871)
Humerobates rostrolamellatus Grandjean, 1936
Sphaerozetes arcticus Hammer, 1952
Family Mycobatidae
Allomycobates lichensis Aoki, 1976
Mycobates parmeliae (Michael, 1884)
Family Mochlozetidae
Mochlozetes penetrabilis Grandjean, 1930
Family Oribatellidae
Adoribatella punctata Woolley, 1967
Oribatella calcarata (C. L. Koch, 1836)
Family Tegoribatidae
Lepidozetes singularis (Berlese, 1910)
Tegoribates bryophilus Woolley, 1965
Family Achipteriidae
Parachipteria petiti TravC, 1960
Pseudachipteria magnus (Sellnick, 1928)
Family Parakalummidae
Porokalumma rotunda (Wallwork, 1963)
Family Galumnidae
Centroribates uropygium Grandjean, 1928
C
B
Aa
C
B
C
Aa
B
B
C
C
C
C
C
C
C
C
An analysis of this list shows that the number of oribatid families and genera
containing lichen-associated species is not large in relation to the total number
of such families and genera. Thus only 27% of the known oribatid families are
represented in the list and 12% of the genera. The number of species of Oribatei
in the list expressed as a percentage of the total number of oribatid species is, of
course, very small indeed.
Another point emerging from an analysis of the list is that certain families are
better represented by lichen-associated genera than others. This is true, for
example, of the Oribatulidae and to a lesser extent of other families such as the
Ceratozetidae. The same is true of some of the genera so far as lichen-associated
species are concerned. Thus genera such as Carabodes, Ameronothrus, Halozetes,
Dometorina, Oribatula and Phauloppia are represented by several lichen-associated
species, whereas many genera are represented by only a single species. A final
point is that certain genera-Provertex, Pirnodus, Dometorina, Oribatula, Phauloppia,
Peloribates-are represented mainly by A or B Group species.
A, B AND C GROUPS OF LICHEN-ASSOCIATED ORIBATID SPECIES
Before dealing in detail with the individual species appearing on the list we
wish to emphasize one or two points. Firstly, the division of lichen-associated
oribatid mites into A, B and C groups is purely a provisional one and should be
regarded simply as a useful framework on which future research can be based.
Moreover it is in large part a division based on subjective criteria, since there is
THE ASSOCIATION OF ORIBATID MITES WITH LICHENS
37 7
an almost inevitable bias in a literature survey which in turn is derived from
field sampling techniques concentrating on a single type of microhabitat, in this
case one or more thalli, possibly of a single lichen species. This is least true of
Group A forms, where there is considerable evidence to support the conclusion
that these are lichen-restricted, or at the very least intimately associated with
lichens, though several of the Aa forms may have to be placed in Group B at a
future date as more data become available.
While the status of several of the Group B forms seems fairly secure, there are
many others which are clearly borderline cases. Some, such as Ommatocepheus
ocellatus, possibly ought to be transferred to Group A, and others placed in
Group C. The same is true for Group C species, some of which might more
properly be classed in Group B and others which should perhaps be deleted
from the list altogether. The whole question of whether a species should be
regarded as generalized in its food habits simply because it is found in a wide
variety of biotopes is really impossible to answer without detailed investigations,
including laboratory studies, of each individual species, in order to determine
whether it is microphytophagous, macrophytophagous or panphytophagous. I t
was, for example, Michael’s (1884) opinion that oribatid mites frequent mosses
not because they are feeding on the material of the moss itself but rather on the
fungi and lichens which grow in and around the mosses.
In no sense then must our three groups be regarded as definitive categories
but rather as the authors’ subjective estimations of the most probable status of
the mites in question, i.e. the relative closeness of their associations with lichens.
The second point we wish to make concerns the taxonomy of many of the
mite and lichen species, which is difficult and complex, and the present
preferred taxon may carry with it one or more synonyms. Where a synonym has
been used for a species in any publication referred to in the text, this synonym is
placed in square brackets after the name and date of the author’s publication to
enable the interested reader to find the reference to that species with the
minimum loss of time. It is perhaps worth noting here that quite apart from
such synonymic difficulties, there are other taxonomic difficulties which arise
purely from wrong identifications. In the case of the Acari, Bernini (1979) has
pointed out that in some cases the supposed widespread distribution of a species
has been incorrectly assessed because of such taxonomic mistakes. It is obvious
that such mistakes must equally affect the soundness of some of the views
expressed in the present study as to the extent to which a particular oribatid
species is closely or not so closely associated with lichens. In fact Cancela da
Fonseca (1970) has gone so far as to suggest that because of widespread
incorrect identifications, the “systematics of type specimens” should be replaced
by a “systematics of type populations” in ecological work!
In the detailed discussion which follows of each species on the list, the species
within each group are arranged in alphabetical order.
Group A species
Dometorina plantivaga plantivaga (Berlese, 1895)
Detailed information about the bionomics of this species has been provided by
Grandjean ( 195I ) , who created the new genus Dometorina, and by Travi. ( 1961,
378
E. L. SEYD AND M. R. D. SEAWARD
1963). TravC ( 1969a) has separated the original species, D. plantivaga (Berlese),
into three subspecies-D. plantivaga plantivaga (Berlese), G. plantivaga insularis
TravC and D. plantivaga saxicola Grandjean.
Grandjean (1951) studied the species in the region of PCrigueux (Dordogne)
and Travi. (1961, 1963) in the Massane Forest and the environs of Banyuls-surMer. In the Massane, TravC (1963) found that the species was most abundant
in lichens encrusting the branches of oak and maple trees at the edges of the
forest. The lichens provide both protection and a source of food to the mites.
As Grandjean was the first to observe, the adult mites are to be found beneath
the thalli of foliaceous lichens on the branches of trees or wandering along the
branches. The immatures excavate cells beneath the thalli of the foliaceous
lichens and feed on the crustaceous lichens adhering to the bark. When the cell
becomes too small for the animal it excavates the foliaceous lichen or even the
bark itself.
The openings of the cells are closed by the mites’ droppings, which are stuck
to each other and to supporting struts constructed across the openings by a
sticky substance excreted by the mites, which are able to repair the ‘plugs’ to
their cells if these are damaged.
Dometorina plantivaga plantivaga has been recorded in lichens by a number of
other workers. AndrC (1975) found that the species constituted 50% of the mites
in a sample of Lecidella elaeochroma (Ach.) Choisy [Lecidea limitata (Scop.) Gray]
and 25% in Lecanora conizaeoides Nyl. ex Crombie. In further investigations,
AndrC (1976a, 1979) has reported the species from both foliaceous and
crustaceous lichens. Gjelstrup ( 1978b) collected the species both in arboricolous
and saxicolous lichens in Denmark, and Gjelstrup & Srachting (1979) extracted
specimens from Ramalina siliquosa (Huds.) A.L.Sm. on Bornholm. Hull (1914b,
1916) [Oribatula plantivaga] found that the species was abundant in lichens on a
cliff wall at Whitley Bay, and Halbert (1923) [Oribatula plantivaga] collected it
near Dublin, though unfortunately he gives no details as to habitat. Rajski
(1968) reported five specimens from lichens on apple trees in Poland. Though
the species is predominantly arboricolous (TravC, 1963; Niedbala, 1969;
Wallwork, 1976) it has on occasion been found also in saxicolous lichens
(Sellnick, 1949, 1960; Gjelstrup, 1978b).
However the species has also been recorded from non-lichen biotopes. Lebrun
(1976), though collecting it from lichens on bark, also found it in bark algae; he
regards the species as microphytophagous (Lebrun, 1971). Most of the nonlichen records have been from mosses (Schweizer, 1922, 1948; Willmann, 1928,
1931a; Franz, 1954), and Berlese (1895a, 1899, 1913a) stated that it lived in
moss throughout Italy. In view of the great weight of evidence that this is a
lichen-restricted species, certainly in its immature stages, it seems most likely
that where it has been reported from moss this has included lichen material, or
that in many cases adults have wandered into non-lichen biotopes. It has
already been noted that the adults are in the habit of wandering about on the
branches of trees. Lions (1975) found a low density of species in forest soil and
litter; he suggests (Lions, 1978) that such specimens have got there by falling
from the trees above. Iturrondobeitia & Subias (1981) also recorded specimens
from non-lichen biotopes in forests. Bonnet, Cassagnau & Travi. (1975)
collected it in moss growing on a rock in a forest and in the surrounding soil.
One further point requires clarification. Van der Hammen ( 1952) [Eporibatula
THE ASSOCIATION OF ORIBATID MITES WITH LICHENS
'379
plantivaga], who collected the species in moss on trees, says that it appears to be
common on orange trees in southern Europe and that the Oudemans Collection
contains a number of specimens from imported oranges, The fact is that there is
an oribatid species, Siculobata sicula (Berlese), which is closely related to
D. plantivaga plantiuaga, and which is very common on orange and lemon trees.
The record by Bayoumi (1979) of D.plantivaga from an orange orchard in the
Nile Delta is quite possibly Siculobata sicula.
Dometorina plantiuaga insularis TravC, 1969
Travi: (1969a) found this subspecies living in a crustaceous lichen species on
the iles Salvages, which lie between Madeira and the Canary Islands. It was
living in the lichens together with another oribatid, Pirnodus soyeri Travt, which
was unusual since in his work on the Oribatei of the Massane, Pyrtntes
Orientales Travt ( 1963) found the oribatid population of crustaceous lichens
was always monospecific.
Dometorina plantivaga insularis is saxicolous and the crustaceous lichen in which
it lives, Buellia subcanescens R.-G.Wern., is severely damaged by the mite. While
the mites are immature the damage is not so apparent but as they grow the
lichen is progressively eaten away and often all that remains is a thin crust
covered to a greater or lesser extent by their droppings and the open and
abandoned cells of the mites. The mites nourish themselves both within and on
the surface of the lichen, and Travi. (1969a) was able to observe the exact
method of feeding. Both the nymphs and adults are unable to feed when the
lichen material is dry.
Dometorina plantiuaga saxicola Grandjean, 1951
This subspecies was collected by Grandjean (1951) from a thin covering of
lichens on rocks. He collected this subspecies four times, twice in Switzerland
(Goschenen and Andermatt) at 1300 m and 1500 m, and twice in Corsica at
similar altitudes in the Col de Bavella, Vizzavona. Like the preceding
subspecies, D . plantivaga insularis, it appears to be saxicolous only. Until there is
contrary evidence we are assuming that the subspecies is lichen-restricted.
Phauloppia saxicola Travt, 1961
Travt (1961, 1963) consistently collected this species from saxicolous lichens,
particularly in habitats exposed to the sun. It was present in lichens from the
Banyuls-sur-Mer littoral and also in the Massane region of the PyrCnCes
Orientales. It was especially abundant in lichens from Font-Romeu and the
Bouilliouses in the Haute PyrtnCes.
Pirnodus detectidens Grandjean, 1956
Grandjean (1956) obtained two adults of this new species and genus by
brushing rocks in a wood to the north of Collioure (PyrCnCes Orientales). These
rocks had a thin covering of lichens. He collected another adult and three
nymphs from soil with tufts of grass but he believed that these had fallen from
the lichen-covered rocks.
Travi. (1958, 1960, 1963) was able to show that Pirnodus detectidens is restricted
to lichens as a biotope. It appears to be known only from the PyrbnCes
380
E. L. SEYD AND M. R. D. SEAWARD
Orientales and Travt found it was very abundant in the Massane Forest region
and he also collected it near Arles-sur-Tech in the Vallespir and near Banyulssur-Mer.
The species, which has exceptional sexual dimorphism, inhabits certain
crustaceous lichens, of which, in particular, it prefers (Pertusaria rupicola (Fr.)
Harm.). Travk, in fact, never found a specimen of this lichen without the mite
being present, and he believed that the mites not only obtained from the lichens
an abundant source of food but also protection against unfavourable climatic
factors (wind, rain, sun) and predators.
The larvae, nymphs and females of the mite (males are much less frequent)
move very slowly and do not make long journeys. They live in cells excavated in
the lichens where these adhere to rocks and normally do not leave these cavities
unless disturbed. They are able to repair ruptures in the cells with their faeces
and Travk (1 958) was able to make a nymph repair ruptures by destroying each
of its reparations. I n any one cavity several specimens and eggs are to be found.
I n his extensive work on the ecology and biology of the oribatids of the
Massane Forest, Trav6 (1963) showed that P. detectidens occurs only on
saxicolous lichens, never on arboricolous ones.
Pirnodus soyeri Travk, 1969
Travt (1969a) found this species living in a crustaceous lichen species on the
lies Salvages and as already mentioned it shares this habitat with the oribatid
Dometorina plantivaga insularis Travk. The lichen, Buellia subcanescens, inhabited by
the two mites, is severely damaged, as has already been described. I t must be
assumed that Pirnodus soyeri is as culpable as D.plantivaga insularis for this
destructive action on the lichen, but owing to the presence of much smaller
numbers of the former species, Travt’s observations were confined to the
activities of D . plantivaga insularis.
Pirnodus soyeri is interesting in that it exhibits an exceptional sexual
dimorphism (TravC, 1969b), which is also characteristic of the other lichenrestricted species of the genus, P . detectidens.
Provertex delamarei Travt, 1962
Travk (1962) first collected this new species in the Massane at 700 m from
foliaceous lichens growing on rocks. All the samples except one were from the
Massane in the Abberes, but a single specimen was collected at 1400 m near
Ayguatebia in the canton of Orlette. All the specimens were found in lichens
except one from mosses.
Further details concerning the habitat preferences of this species were given
by TravC (1963). The species was almost entirely restricted to lichens in which it
was abundant. The rocks on which the lichens containing the species were found
were in exposed southern-facing positions. While the great majority of specimens
were taken from lichens a t a number of different sites, there were three moss
samples in which a few specimens were present.
It appears the Provertex delamarei is a rare species and is only found in a
restricted and rather special habitat. We are placing the species in Group A in
view of the fact that the great majority of the specimens have been collected
from lichens, and we are assuming that the few specimens found in mosses have
strayed there accidentally.
T H E ASSOCIATION OF ORIBATID MITES W I T H LICHENS
?A3 1
Another species of the genus Provertex discovered by Travk also inhabits
lichens. The type species of the genus, P. kiihnelti MihelEiE, 1959, from Austria,
on the other hand, does not appear to have been collected from lichens.
Provertex mailloli Travk, 1964
This species was collected from foliaceous lichens covering rocks a few
kilometres from Banyuls-sur-Mer (TravC, 1964). The collection consisted of 18
adults, 40 nymphs and 12 larvae. A further male and female were obtained from
lichens and mosses growing on a rocky ledge at the other end of the Pyrknkes
Orientales in the Carlitte massif near the Porteille de la Grave at about 2300 m.
Until there is evidence to the contrary, it must be assumed that P . mailloli is
lichen-restricted.
Scapheremaeus petrophagus (Banks, 1906)
In 1906 Banks described a new species of oribatid, Scutovertex petrophagus,
which he had collected from rocks taken from the Traghanic Falls near Ithaca,
New York. Banks found that the surface of the rock was pitted and within each
pit was a mite, which he believed excavated the rock. He assumed that there
must be some tiny plant organism in the pits on which the mites fed.
Jacot (1934) visited the site where Banks made his discovery and found that
as the waves broke against the massive blocks of limestone, the rock surface
became moist enough to support an extensive growth of Thelidium auruntii
(Massal.) Krempelli, its black oval fruiting bodies projecting somewhat when
ripe. Jacot found specimens of Scapheremaeus petrophagus feeding on the fruiting
bodies at night and spending the day in cavities in the thallus.
I t is strange that Banks did not observe the lichens. Presumably the mites not
only shelter in the lichens as Jacot describes but also during the day within
crevices in the rock, which Banks believed that the mites excavated.
Sub-group Aa species
Allomycobates lichensis Aoki, 1976
Aoki (1976) investigated the oribatid fauna of a number of plant zones on the
eastern rocky slope of the Mt. Shirame volcano, Japan, where sulphurous acid
gas spouts through cracks in the rocks.
The zone nearest the crater contains only the lichen Cladonia theiophila Asah.,
which is strongly sulphur-resistant, and 163 specimens of A. lichensis were
collected from the lichen. The dominant plant in the next zone is Pobgonum
cuspidatum, which contained no specimens of the mite. In the zone beyond that,
Cladonia theiophila was again present and the collection from the lichen yielded
3936 specimens of A . lichensis, whereas other oribatid specimens were present in
very small numbers. Beyond this zone came three more vegetation zones in
which lichens were not present and A. lichensis was present only in the last zone,
290 specimens being collected. This zone, however, consisted of several different
forms of tree and it is perhaps possible that arboreal lichens were present from
which the mites originated. In view of the very large numbers of specimens
382
E. L. SEYD AND M. R. D. SEAWARD
collected from lichens in two of the zones, we are provisionally placing this
species in Group A.
Ameronothrus maculatus (Michael, 1884)
The first record of this species was by Michael (1884) [Scutovertex maculatus],
who found it in lichens at Land’s End, Cornwall. Since then it has been
collected mostly in lichens on the sea shore or at least on coastal regions; it
appears on the list of Evans et al. (1961) of oribatids charactkristic of the orange
lichen zone of rocky shores.
Halbert ( 1920) [Scutovertex maculatus] collected it under lichens on exposed
rock surfaces at Malahide on the coast near Dublin. Gilbert (1976) recorded it
from foliose lichens (Xanthoria parietina) on the Flannel Islands, Outer
Hebrides, and Gjelstrup & Sachting (1979) from Ramalina siliquosa on Bornholm
in the Baltic. Grandjean (1958) found it was present in many collections
obtained by brushing the surface of rocks covered with thin lichens on the coast
of Trkgastel, Brittany. Sellnick (1928, 1960) also describes it as a coastal species
though in a paper on the oribatid fauna of the Swedish coast he states that the
six specimens he collected were not from lichens (Sellnick, 1949). Other workers
who have recorded the species from the sea shore are Karppinen (1966), who
does not mention lichens as one of the biotopes, and Schulte (1974), who found
the species in the supralittoral and inland areas near the coast. In a later paper
(Schulte, 1976), while implying that A . maculatus can feed on algae, he classes it
as a lichen-dweller and describes it living in the thalli of crustose lichens of the
coast and inland, grazing in isolated groups on the soredia. Traglrdh (1931),
although describing the species as a coastal lichen-dweller, obtained his
specimens from moss on a stone. There are records in the collections of the
British Museum (Natural History) of specimens from the Farne Islands,
Northumberland, and from the lichen, Lichinu pygmaea (Light F.) Agardh, from
Cardigan Bay and from Wembury in Devon. In the Oudemans Collection (see
van der Hammen, 1952) there is a record of the species from drift accumulation
along the shore at Zeeburg.
The species is, however, not restricted to coastal lichens. Laundon (1967)
reported depredations of various species of the association Caloplacetum heppianae
DR. by this mite in various London churchyards. Gilbert (pers. comm.) found it
in Lecanora achariana, growing on oak trees beside a tarn on Helvellyn, and noted
that the lichen was damaged, presumably by the mites. The British Museum
(Natural History) has a record from a lichen thallus at Beckenham, Kent, and
Hull (1914a) [Scutovertex maculatus] found it under stones in the Derwent Valley.
It is perhaps worth mentioning that Hull (1918) described a subspecies,
S. maculatus v. pseudomaculatus, from the wall lichen, Xanthoria parietina [Physci
parietina] .
The species could be regarded as a borderline case between Groups A and B,
for while Schuster (1979) is in no doubt that the main food of the species is
lichen and Wallwork (1976) classes it as a lichenophage, there are as we have
seen a few records from non-lichen biotopes. Oudemans ( 1900) [Scutovertex
maculatus] for example, records specimens from Oude Pekela, Netherlands, in
dried chicory roots and the British Museum (Natural History) has a record from
a house in Blandford, Dorset. Clearly more work on this species is required
before it can be assigned definitely to Group A rather than Group B. A survey of
THE ASSOCIATION OF ORIBATID MITES WITH LICHENS
383
the ecology of the Ameronothridae is given by Schulte (1976) and by Schulte,
Schuster & Schubart (1975).
Ameronothrus lapponicus Dalenius, 1963
Dalenius (1963b) stated that this species has only once been found in the
Tornetrask territory of Swedish Lapland on pine trees but he makes no mention
of lichens as the biotope. Schulte (1976), however, states that A. lapponicus is a
lichenophage, living in the thalli growing on decaying conifers of the arctic tree
line and that the mites depend on lichens for food (see also Schuster, 1979).
Ceratozetes shiranensis Aoki, 1976
This is a second oribatid mite species collected by Aoki (1976) from the plant
zones on the eastern slope of the M t Shirane volcano.
The species was present, together with Allomycobates lichensis, in the lichen
Cladonia theiophila occupying the first zone. There were, however, only 25
specimens in the lichen sample from this zone as against 163 specimens of
A. lichensis. In the third zone there were only two specimens in the lichen sample
as against nearly 4000 specimens of A. lichensis. However, except for a single
specimen in one of the non-lichen zones, Ceratozetes shiranensis was otherwise
absent from such zones and for this reason we are including the species in
Group A.
Cryptoribatula taishanensis Jacot, 1934
Nineteen specimens of this species were picked out of crustose lichens which
were encrusted on large boulders near Erhu Miao, Taishan, Shantung at about
3000 feet. The mites were completely enclosed in the lichens Uacot, 1934).
More information is needed on the habitat of this mite but the likelihood of it
being a true lichenophage is increased by the fact that, as Grandjean (1956)
points out, it is related to Pirnodus detectidens, which is a lichen-restricted form.
Qmbaeremaeus marginalis Banks, 1895
Banks ( 1895) [Eremaeus manginalis] collected numerous specimens of this species
under lichens on the bark of apple trees at Sea Cliff, New York. In his treatise
on the Acarina, Banks (1904) says it occurs under lichens on trees in eastern
U.S.A. Whether this statement refers only to the 1895 collection or whether
further collections were made is not clear.
Halozetes marionensis Engelbrecht, 1974
Numerous adults, nymphs and larvae were collected by Engelbrecht ( 1974a)
from lichens on an exposed beach covered with smooth boulders on Marion
Island, South Africa.
Maudheimia petronia Wallwork, 1962
This is an antarctic species. Five specimens were found by Wallwork i1962) in
saxicolous lichens.
Nothrus simplex Banks, 1895
According to Banks (1904), this species is present in lichens on dry rocks in
North America.
384
E. L. SEYD AND M. R . D. SEAWARD
Oribatula parisi Travt, 1961
TravC (1961) collected this new saxicolous species “au pic des 4 Termed’ from
Pamelia saxatilis (L.) Ach. Martinez (1981) collected specimens a t the base of
and on the trunk of an oak tree on which were growing mosses and lichens.
Oribatula saxicola Halbert, 1920
This species is included in the list of Evans et al. (1961) of species
characteristic of the orange lichen zone of the sea shore. Its inclusion in this list
appears to be based on Halbert’s records of the species from the orange lichen
zone of Malahide Island and of Howth. Halbert (1920) collected the species at
Malahide from dry or slightly damp rocky flakes, but a t Howth he found it
under lichens growing on rocks on the shore.
Peloribates barbatus Aoki, 1977
This species was collected by Aoki (1977) from lichens growing on tombstones
in Ichihara-shi, Central Japan.
Peloribates nishinoi Aoki, 1977
As with the preceding species Aoki (1977) found this species in lichens
growing on tombstones at the same site.
Phauloppia knoep8eri Travt, 1961
L.-P. Knoepffler collected a considerable number of adults and immatures of
this species from lichens at an altitude of 80 m near Cape Saint-Florent in
Corsica (Travt, 1961). We are not aware of any other recordspofChis spTGei
Phthiracarus crinitosimilis Willmann, 1939
Willmann (1939a) collected the species from lichens growing on spruce trees
and from the bark of these trees in the Glatzer Schneeberg of Poland. Willmann
also found specimens in moss including Sphagnum but he believed that these
species may have fallen here from the trees above.
Trhypochthonius montanus van der Hammen, 1956
This species was collected by van der Hammen (1956) from Cladonia on alpine
heath at an altitude of about 1600 m in the Wissel Lake region of Dutch New
Guinea.
agoribatula frisiae insularis TravC, 1961
Travt (1961) collected specimens of this subspecies near the village of Soller,
Majorca, by brushing a tree trunk. Three other samples of lichens from volcanic
rocks containing the subspecies came from the jles Columbretes, Spain.
agoribatula frisiae insularis appears to have been the most abundant of the
oribatid species in the samples. The Oribatei were few in numbers probably
because the climatic conditions are extremely rigorous, the islands suffering
severe gales in winter and long periods of drought in the summer.
This subspecies is provisionally placed in Group A until further data become
available. agoribatula frisiae (Oudemans, 1916) was also collected by TravC
T H E ASSOCIATION OF ORIBATID MITES W I T H LICHENS
383
(1961) from a number of the lichen samples but this species is normally found in
numerous non-lichen habitats (see Group C ) .
Group B species
Autogneta penicillum Grandjean, 1960
This species was taken by Grandjean (1960) in the neighbourhood of Dax,
France, from the bark of a pine stump overgrown with lichens and he also found
it in rotten wood. The gut contents consisted of fragments of mycelia and spores.
Calyptoretes sarekensis (Tragirdh, 1910)
Tragdrdh ( 1910) [ Oribata sarekensis] found this species in lichens, mosses and
under stones in the Sarekgebirge of Swedish Lapland. Thor (1929) stated that
the species was plentiful in lichens in the subalpine regions of Norway; he also
collected it on Spitzbergen and Bear Island in lichens, litter and mosses (Thor,
1930a). He describes the species as a lichenophage.
Hammer (1937, 1944, 1946, 1952b) discovered that C. sarekensis was very
common in Greenland in dry biotopes with an under-vegetation of lichens, and
she collected specimens in northern Canada (Hammer, 1952a) from vegetation
with which lichens were associated. Strenzke, Lesse & Denis (1955), who also
reported the species from Greenland, collected the majority of specimens in
Cladonia although some were found in other biotopes. Strenzke (1952) had
earlier described the species as one characteristic of lichen-covered habitats.
Dalenius (1960, 1962a, 1962b, 1963a, 1963b) studied the Oribatei of the
Tornetrask territory of Swedish Lapland. He believed C. sarekensis to be
predominantly a lichen-Seeder and he found it was very abundant in lichenheath (low alpine belt), although it was equally abundant in stony soil of the
high alpine belt.
Travi: (1963) collected the species in the Hautes Pyrknkes from several
different types of lichens and mosses and also from lichens in the subalpine zone.
He also collected it from lichens and mosses on the C6te d'Azur near St
Raphael and at Yice near the village of St Agnks. Solh0y (1975) found that the
species was present among the mites he extracted from samples he took both
above and below ground on lichen-heath in southern Norway.
Seyd (1979) has reported the presence of this arctic-alpine form from the
summits of mountains throughout the British Isles. In the majority of the
localities his samples were a mixture of mosses and lichens, but in the case of the
samples from Kinder Scout, Derbyshire (Seyd, 1962) and Moel Hebog,
Snowdonia (Seyd, 1968) separate samples were taken and C. sarekensis was
present only in the lichen sample. O n Burtness Combe, Lake District [Seyd,
1966) and the Cnicht, Snowdonia (Seyd, 1981), where separate samples were
also taken, the species was present in the moss as well as the lichen sample.
However, in the latter collection, while C. sarekensis formed only 3.6", of the
oribatid specimens present in the moss sample, the species formed 20.5O,, of the
lichen sample specimens.
From this review it would appear that the species has a decided preference for
lichens as a biotope. Nevertheless it is clearly not restricted to this habitat and
both Tuxen (1943) in Iceland and Karppinen (1967) on Spitzbergen and in
Norway (Karppinen, 1971) have reported it only from non-lichen biotopes. As
17
386
E. L. SEYD AND M. R. D. SEAWARD
Hammer (1944) observes, it does seem to have adapted itself to live on various
plants as well as lichens.
Camisia invenusta Michael, 1888
Michael (1888) only mentions lichens as the habitat for this species. Halbert
(1915, 1920) found it under lichens on rocks on the sea shore at Howth,
Ireland, and Evans et al. (1961) include it in their list of orange lichen zone
species. However the species is by no means restricted to coastal areas. It was
collected by Seyd (1966, 1968, 1981) from lichens and mosses on mountain
summits in the Lake District and Snowdonia, which is of some interest since
TravC (1960, 1963) recorded it from lichens and mosses in the Massane,
PyrCnCes Orientales, and in the Hautes Pyrtntes. TravC found that it was rarer
in arboricolous lichens and mosses than in saxicolous ones. Sellnick & Forsslund
(1955) collected it from Parmelia centrifuga and moss in Sweden, and Sellnick
(1908b) from moss on the Faroe Islands. Other non-lichen records are those of
Schweizer (1922) from humus in Switzerland, Csiszar & Jeleva (1962) from
moss in Bulgaria, Polderman (1974) from a salt marsh in the Netherlands, and
Bernini (1971) from several non-lichen biotopes in Italy. The species is a rare
one and several of the records are from high altitudes.
Cumisia segnis (Hermann, 1804)
Grandjean (1948a, 1950a) studied the habits of this species in lichens growing
on the branches of a maple tree by the door of his house near Ptrigneux. He was
also able to study the activities of the mites in culture cells, in which lichen
material was used as the food source.
Grandjean (1936a) had earlier obtained his specimens of C. segnis by beating
and brushing the trunks, branches and leaves of trees. I n some cases he obtained
specimens from moss and debris under trees, but since he regarded the species as
arboricolous, he considered that they had fallen there from the trees above.
Willman (1928, 1933, 1952) collected it in both ground and corticolous
lichens, and AndrC (1979) also reports it from the latter. Sengbusch (1957)
found it in lichens from a scarlet oak. Travk (1960) obtained one specimen on
lichens and three in moss in the Massane. He later collected specimens by
beating conifers: from plants and soil on a large rock; from rocks covered with
liverworts and lichens in woods at Banyuls-sur-Mer, and from lichens and
mosses in the Hautes Pyrenkes (Travi., 1963). Gjelstrup (1978a) and Subias
(1980) took single specimens from lichens and mosses, and Seyd (1981) reported
seven specimens from lichens and some from moss from a mountain summit in
Snowdonia.
It may seem surprising in view of these records and in view of the fact that
Travk (1963), on the basis of Grandjean’s studies, includes the species in his
group of lichenophagous forms, that we do not place C. segnis in Group A.
There is, however, a long list of authors who have collected the species from
such diverse habitats as moss including Sphagnum, leaf litter, humus, pineneedles, under bark and in soil (Berlese, 1885a [Nothrus segnis]; Michael, 1888
[ibid.];TragHrdh, 1910 [ibid.];Hull, 1914a [ibid.];Halbert, 1915 [ibid.];Sellnick,
1908a, 1928, 1929; Thor, 1937; Franz, 1943; van der Hammen, 1952; Strenzke,
1952; Tarman, 1955; Sellnick & Forsslund, 1955; Karppinen, 1955; Schweizer,
1922, 1948, 1956; Willmann, 1931a, 1956; Kunst, 1957, 1958; Dalenius, 1950,
T H E ASSOCIATION OF ORIBATID MITES W I T H LICHENS
387
1960; Halkkova & Kunst, 1960; Tarras-Wahlberg, 1961; Seyd, 1962; Block,
1965; Luxton, 1966b; Rajski, 1967; Bernini, 1971; Fujikawa, 1972; Webb, 1972;
Usher, 1975a, b; Schatz, 1979; Christensen, 1980). In culture experiments
Littlewood (1969) was able to feed her specimens on algae and Schuster (1956)
regards C. segnis as a non-specialist feeder because the greater part of its
droppings are not exactly definable.
It might still be argued that C. segnis is an arboricolous lichenophage, which
has fallen from trees into such habitats, but by no means all of the above
biocoenoses were from woods. Seyd (1962), for example, recorded the species
from moss on the top of Kinder Scout, where there have been no trees for a long
time. Aoki (1973), in fact, describes the species as a “wandering form” which
lives in the soil and litter as well as on trees.
Carabodes labyrinthicus (Michael, 1879)
Michael (1884) [ Tegeocranus labyrinthicus] stated that the nymphs of this
species are common in lichens and mosses on walls, and Hull (1915, 1916) also
recorded it from lichens and mosses. Halbert (19 15) [ Tegeocranus labyrinthicus]
found it under lichens on the sea shore at Howth. Willmann (1933, 1939a, 1952,
1956), while recording it from lichens on the bark of trees, also recorded it many
times from moss including Sphagnum (Willmann, 1928, 1929, 1931a, b, 1937,
1939a, 1942a, 1943, 1954, 1956). He also found it in moss in the water of springs
(Willmann, 1923). Hammer (1952a, 1972) collected it from lichens and moss on
a rock and on a beech stump. Tarras-Wahlberg (1952) recorded it from lichens
and Sphagnum, and Strenzke (1952), who collected it from lichens and mosses,
described it as a species characteristic of lichen-covered biotopes. S o l h ~ y( 1975)
found it both above and below ground in lichen-heath and Steigen, S o l h ~ y&
Gyllenberg (1975) reported it as being very common in the lichen and litter
layers of lichen-heath, constituting 40% of the total mite biomass. Schatz (1979)
also obtained specimens from lichen heath. Dalenius (1960) lists it among the
lichen-heath species, although he found it many times in other habitats as well
(Dalenius, 1950, 1960). Sellnick & Forsslund ( 1953), Karppinen ( 1962) and
Seyd (1962, 1981) have all recorded the species from lichens, and Gjelstrup &
Sochting (1979) found that it was abundant in Ramalina siliquosa.
Several authors besides Willmann have taken the species in arboricolous
lichens. Thus, van der Hammen (1952) has records from lichens on oak trees
although he also found it in forest litter and moss, and Pschorn-Walcher &
Gunhold (1957) note its presence in lichens and mosses on trees in parks. Andrt.
(1976a, 1979) also has records of the species from arboricolous lichens and in
one such lichen, Lecanora conizaeoides, C. labyrinthicus constituted 50% of the mites
present (Andrk, 1975). Gjelstrup (1979) found it was the dominant oribatid
mite in lichens on beech and birch trees but this author notes that it is
saxicolous as well as arboricolous (Gjelstrup, 1978b). Travk f 1960, 1963j also
found it in both saxicolous and arboricolous lichens in the Pyrenees.
Besides the records already noted from moss, including Sphagnum, there are a
considerable number of further records from these two habitats, as well as from
litter, humus, soil, dead leaves, pine needles, etc. (Hull, 1914a, 1915; Sellnick &
Forsslund, 1953; Franz, 1943, 1954; Hammer, 1944, 1946, 1955; Schweizer,
1922, 1948, 1956; Kunst, 1957, 1958; Tarman, 1958; Sellnick, 1928, 1929, 1960;
Halgskovi & Kunst, 1960; Tarras-Wahlberg, 1961; Lebrun, 1965a; Rajski,
388
E. L. SEYD AND M. R. D. SEAWARD
1968; Ptrez-Ihigo, 1971; Bernini, 1971; Engelmann, 1972; Cancela de Fonseca,
1975; Steigen et al., 1975; Athias-Henriot & Cancela d a Fonseca, 1976;
Karppinen, 1956a, b, 1957, 1958a, c, 1966, 1972, 1977; Cgluggr & Vasiliu, 1980).
I n Lebrun’s (1971) opinion C. labyrinthicus is certainly a macrophytophage, an
opinion with which Schatz (1979) agrees. Nordberg (1936) found the species
was very common in the nests of many different types of bird, which is not
surprising when specimens are common in the plant material from which the
nests are made. Wiiniewski (1965) found a high density of the species in ant
hills, which Rajski (1968) suggests is due to the accumulation of raw humus in
such habitats.
Carabodes minusculus Berlese, 1923
Willmann (1928) collected over 100 specimens of this species from Cladonia on
moorland, and in his work on the Oribatei of Germany (Willmann, 1931a) he
gives the biotope for this form as lichens in wood, heath and moorland. Other
workers who have reported it in lichens are: Tarras-Wahlberg (1952) from
Sweden; Schweizer ( 1956) from the Swiss National Park; Karppinen ( 1958c,
1971) from Norway and Finland; Block (1966a) from the hagg lip of an eroding
blanket bog in Westmorland; Seyd (1962, 1966, 1968) from mountain peaks in
Britain; Travi: (1956, 1963) from the Pyrkntes; Bernini (1976a) from Italy;
Gjelstrup (1978a) from the Faroe Islands; Gjelstrup & S ~ c h t i n g(1979) in
Ramalina siliquosa from Bornholm Island and Tarman (1973) in the Triglav
National Park. I n several of these records other habitats are mentioned as well,
and a number of authors, including some of the above, have collected the
species in a wide variety of non-lichen habitats such as moss including Sphagnum,
litter, humus, soil and grass from woods and heathlands (Willmann, 1942a;
Schweizer, 1948; Strenzke, 1952; Evans, 1952; Franz, 1943, 1954; Kunst, 1957;
Sengbusch, 1957; Hal6skova & Kunst, 1960; Frank & Zivkovitch, 1960; Block,
1965; Luxton, 1966b; Karppinen, 195613, 1966; Rajski, 1968; Bernini, 1971;
Pkrez-Iiiigo, 1971; Dalenius, 1950, 1960; Webb, 1972; Engelmann, 1972;
Solhray, 1976; Mahunka, 1977; Lions, 1978). There is even a record by
Wiiniewski (1965) from an ant’s nest. Lebrun (1971) classes the species as a
macrophytophage.
It might appear from this review that C. minusculus does not have a preference
for lichens as a biotope and should be placed in Group C. Yet the suspicion
remains with us that given the choice between lichens and other plant material
the species prefers the former. This view is based partly on Willmann (193l a )
giving lichens as the normal habitat for the species and on Rajski’s (1968)
statement that it is often to be found in dry lichens. Strenzke (1952), too, states
that it is frequently to be found in lichens, although in fact his collection was
made in humus. Moreover Seyd (1981), in a study of the oribatid fauna of a
Snowdonia mountain peak, collected samples of moss and lichens from a small
area and while the moss sample contained only three specimens of C. minusculus,
the lichen sample contained over 1000.
Carabodes willmanni Bernini, 1975
Bernini (1975) collected this new species from 28 different localities. It was
abundant in lichens at a few sites but most he collected in mosses. However
Bellido (1978, 1979), in a study of the species in the Massif Forest of Paimpont
THE ASSOCIATION OF ORIBATID MITES WITH LICHENS
389
( Ille-et-Vilaine) , found 191 specimens in 13 samples of saxicolous lichens and
11 790 specimens in 15 samples of Cladonia, while samples from an Erica biotope
had only 2 1 12 specimens. I turrondobeitia & Subias ( 1981 ) collected specimens
from forest habitats in Spain.
Bernini (1975) also showed that some of the records of Carabodes minusculus by
several authors were really records of C. willmanni. These records include those
by van der Hammen (1952), who described the species as abundant in mosses
and lichens; Strenzke (1952), who found it in the humus of woods and heaths;
Sellnick & Forsslund (1953), who recorded it as abundant in Cladonia; and
Perez-Ifiigo (1971),who recorded it from the Spanish fauna in humus and moss.
Bernini (1976a) also showed that there were other specimens in some of the
European collections which had been incorrectly identified and from this
material, as well as from his own collections, he was able to describe five
new species of Carabodes. Two of these, C. schatzi Bernini and C. dissimilis Bernini,
were collected from lichens and a year later Bernini (1977a) described yet
another new species, C. grandjeani Bernini, a single specimen of which he found
in lichens near Siena, Italy. I t may well be that these three species will also find
their place in Groups A or B as further data on their habitats accumulates.
Eporibatula gessneri Willmann, 1931
Willmann (1939a, 1956) collected the species in the Glatzer Schneeberg of
Poland from lichens and the bark of trees and in some non-lichen biotopes,
though he states that it is characteristic of tree and ground lichens.
However Willmann (1942b) also collected the species in moss in a mineral
spring and Pax & Willmann (1937) had found it earlier on pieces of wood,
stones and moss in one of the waterfalls of the Glatzer Schneeberges. In fact the
original description of the species (Willmann, 1931b) was based on specimens he
had collected from moss and submerged plants at the edges of lakes in the
Altvatergebirge of Czechoslovakia.
There is a record by Winkler (1954) from a spruce forest in the Jeseniky
mountains of Czechoslovakia but the species is certainly rare. In Poland,
however, it is not endemic to the Sudety mountains as Pax (1944) supposed,
since Rajski 11968) collected it in the Poznan district.
Hydrozetes capensis Engelbrecht, 1974
Engelbrecht (1974b) collected this species in South Africa from six different
biotopes. One sample of dripping-wet lichens and mosses from a canal contained
100 specimens and a second sample from wet lichens contained 50 specimens.
The other four samples from underneath shrubs, among dry leaves beneath
poplars, among wet decomposed grass and among wet reeds contained only
single or very few specimens.
Humerobates rostrolamellatur Grandjean, 1936
There is some confusion over the nomenclature ofthis species and as a result
of a critical survey of earlier descriptions by Jacot (1931) and Grandjean
(1936a), the latter author found it necessary to rename the species. (See also van
der Hammen (1952) for a list of synonyms.)
Grandjean ' 19364 described the species as arboricolous. 'Thus Willmann
(1931a) [Humerobates fungorum (L.)] recorded large numbers on the bark of fruit
390
E. L. SEYD AND M. R. D. SEAWARD
trees feeding on fungi and lichens, although he also collected specimens in moss
and conifer needles. Oudemans (1905) [Murciu humerulis (Herm.)] and Voigts &
Oudemans (1905) [Notarpis humeralis Herm.] found specimens under bark, and
Oudemans (1916) [Murcia humeralis (Herm.)] collected the species on the bark of
fruit trees and believed it to be feeding on algae and fungal hyphae and spores.
Arboricolous collections of the species by Lions (1966) and Lebrun (1976) were
made from lichens, though the latter author also found it in bark algae, and
Murphy & Bulla (1973), who found it was prevalent on various kinds of fruit
trees, consider it feeds on algae and other lower plants.
Travi. (1956) also found H . rostrolumellutus to be common on trees in lichens
and mosses, but he also collected it from saxicolous lichens (Travt, 1958). In his
study of the oribatid fauna of the Massane, PyrknCes Orientales, Travi. (1963)
collected it from both arboricolous and saxicolous lichens and mosses, and at
Banyuls-sur-Mer he found it in great numbers in lichens. Seaward (unpub.
data) found a species-poor lichen community on Sambucus heavily infested with
this oribatid.
On the other hand, the following authors have collected the species from a
variety of non-lichen biotopes such as soil, moss, decaying leaves, litter, etc.:
Berlese ( 1883) [ Oributes humeralis (Herm.)]; Michael ( 1884) [Oribatu lapidaria
Lucas]; Oudemans (1896) [Oribata humeralis (Herm.)]; Oudemans (1900)
[Notuspis lupidurius (Lucas)]; Sellnick ( 1928) [Humerobutes humerulis (Herm.)];
Franz (1954); Willmann (1956); Luxton (1966b); Bernini (1969); Webb (1972);
Phez-Ifiigo (1975). Strenzke (1952) recorded it from woods, meadows, moss on
walls, as well as under the bark of trees, and van der Hammen (1952) in moss
from a number of different habitats and even in the nest of a spider! Subias
(1980) collected it in turf in Spain. Jalil (1969) was able to maintain a culture of
the species on a diet consisting of algae. This species appears to be a borderline
case between Groups B and C.
Lucoppiu (Phuuloppia) nemorulis Berlese, 1916
Berlese (1916) collected the type specimens from moss but most of the other
records of this species have been from lichens. Schweizer ( 1956) [Phuuloppia
nemorulis] recorded it from lichens on a stone in the Swiss National Park, while
the records given by Willmann (1933, 1939a, 1956) [Oribata nemoralis] are from
arboricolous lichens. In fact Willmann (1933) picks out the species as one which
is particularly characteristic of tree lichens. Willmann ( 1954) [Oribatu nemorum]
did find the species in soil but says that it is usually found in lichens and moss
growing on trees. Franz (1954) collected it in woodland litter, and Csiszhr &
Jeleva (1962) from moss on tree-trunks in Bulgaria. It appears on Balogh’s
(1943) Hungarian list but the author gives no habitat.
Maudheimia wilsoni Dalenius & Wilson, 1958
This species is known only from two localities in the Antarctic, both in
Western Droning Maud Land (Dalenius & Wilson, 1958). One is Passat and the
other Ekberget, the latter being unique in that it is the southern-most locality
where wintering land animals have been found.
The mites were found in large numbers under stones and pieces of rock,
presumably sheltering from the severe climate. The vegetation consists only of
lichens, mosses and algae, on which it must be assumed the mites feed. There is
THE ASSOCIATION OF ORIBATID MITES WITH LICHENS
39 1
an abundance and luxuriance of foliaceous, fruticolous and crustaceous lichens,
often covering large areas, particularly at Passat. Mosses, on the other hand,
occur in small scattered patches and algae only in connection with the organic
debris of birds.
It is a pity that the authors were not able to carry out laboratory feeding
experiments to determine whether the mites preferred lichens or mosses as a
diet. They note that only a few mites were found in mosses, so that one must
assume that most were feeding on the abundant lichens, but there is no
reference to lichens having been collected and processed for the mites.
Mochlozetes penetrabilis Grandjean, 1930
Grandjean (1930) collected this new species from the branches and trunks of
trees and from dead wood in the neighbourhood of Colon (Panama). He also
obtained specimens in Colombia and Venezuela. Many years later Grandjean
(1959) obtained further specimens from grass in Trinidad, and an examination
of the excreta showed that this contained mainly mycelial filaments and an
abundance of brown spores, which Grandjean believed were from lichens.
Mycobates parmeliae (Michael, 1884)
Michael ( 1884) [Oribata parmeliae] recorded this species only from Xanthoria
parietina [Parmelia parietina] at Land's End, Cornwall, and Colloff (in litt.)
extracted 17 adults of this mite from the same species of lichen on a rocky shore
at the Lizard. Halbert ( 1915, 1920) [Oribata parmeliae] found it was common
under lichens on rocks on the sea shore at Howth, Ireland. Sellnick '1949)
collected i t in lichens o n the Swedish coast. As in the case of Camisia invenusta
(Mich.), it appears on the orange-lichen zone list of Evans et al. (1961 j but like
this species i t is by no means confined to maritime locations.
Tarman (1973) found specimens in soil and lichens growing on a scree in the
Triglav National Park, but he also found it in the soil and litter of a pine forest.
Gjelstrup (1978b) recorded the species in Denmark from arboricolous and
saxicolous lichens and Gjelstrup & S ~ c h t i n g(1979) collected it from Ramalina
siliquosa on the Island of Bornholm in the Baltic. There is a record by IVillmann
(1956) from C'etraria islnndica (L.) Ach. from the Spieglitzer Schneeberg of
Poland, and this author gives lichens and mosses as the biotopes for this species
in his survey of German oribatids (Willmann, 1931a), as does Sellnick i1928) in
his study of European forms.
However, several authors have found the species in litter, dead leaves, moss
including Sphagnum, soil, etc. (Franz, 1943, 1954; Hammer, 1952a; kIihelEiE,
1957; Kunst, 1957, 1958; Karppinen, 1958b, 1971, 1977; Engelmann 1972;
Fujikawa, 1972, 1974; Lions, 1975, 1978). Schweizer (1956) collected it in moss
i n the Swiss Xational Park, though earlier he had also recorded it there from
lichens (Schweizer, 1948;).
Travi. (1960, 1963) found M . parmeliae in many of his samples from the
Massane, Pyrknkes Orientales. The majority of these samples were arboricolous
and saxicolous lichens, but in a few cases small numbers of specimens were
recorded from mosses and liverworts. Travi: (1963), in fact, classes the species as
a lichenophage and believes it to be restricted to lichens in its immature
stages, whereas the adults are capable of feeding on other plant material, a point
stressed also by Wallwork ( 1976).
392
E. L. SEYD AND M. R. D. SEAWARD
Ommatocepheus ocellatus (Michael, 1882)
Travt (1956, 1963) studied this species in the Massane, Pyrtntes Orientales,
where he collected it from various species of lichens and also from liverworts.
The species is to be found both on saxicolous and arboricolous lichens but it has
a preference for the latter. He also recorded it from Ortdon, Hautes Pyrtnkes, in
lichens on a rock.
It was only on the corticolous lichen, Pertusaria albescens (Huds.) Choisy &
Werner [Pertusaria globulzfra], that Travt (1963) was able to observe the mites
feeding and then only when the lichens became saturated after rain. Both adults
and nymphs remain quiescent under dry conditions. Travt classes the species as
a lichenophage, but leaves open the question as to whether it is lichen-restricted
in its diet. Wallwork (1976) also classes it as a lichenophage.
Michael (1882) described the species from material he had collected on
lichens at Land’s End. The only other record from Britain appears to be that of
Seyd (1981), who collected one specimen from lichens at 687 m in Snowdonia,
N. Wales, which is of some interest in view of Travt’s record from the Hautes
Pyrtntes. Willmann (1931a) gives not only lichens but also moss as the biotope
of 0. ocellatus, while Sellnick (1928) gives only moss. Tarman (1973), Mahunka
(1974) and Lions (1978) have found the species in forest litter and soil, Csiszar
& Jeleva (1962) in moss on a cliff, and Ptrez-Iiiigo (1975) in soil on Tenerife.
Bernini (1971) found it in Juniper and Vaccinium cushions and in meadowland at
1000 m on Mt. Reatini, Italy.
Phauloppia coineaui Travt, 1961
This species was collected by Travt (1961, 1963) from the Massane region
and from the Hautes Pyrtnkes, but it was rare. It was nearly always found in
saxicolous and arboricolous mosses and lichens in the forests of Sainte-Baume
and Fontainebleau.
Gjelstrup & Sochting (1979) found that this species was very abundant in
Ramelina siliquosa on Bornholm in the Baltic. Moreover it was discriminating in
its lichen host, avoiding closely related species. Apparently the loose medullary
structure of R. siliquosa provides the mites with the best environment for their
development, both from the point of view of food and protection. Colloff (in litt.)
extracted 12 adults of this mite from a 2 g sample of Ramalina siliquosa and 35
adults and nymphs from a 12 g (wet wt) sample of Xanthoria parietina from two
British rocky shore sites. Bernini (1971), however, found the species in nonlichen habitats and the records of Lions (1975, 1978, 1979) have been from
forest humus.
Phauloppia lucorum (C. L. Koch, 1841)
There has been considerable confusion over the nomenclature of this species,
mainly because Berlese’s (1892) description of the Oppia lucorum of Koch (1841)
really refers to the Notaspis burrowsii of Michael (1890). It is Berlese’s (1895b)
Oppia conformis which is synonymous with Koch’s Zetes lucorum. The situation is
not made any easier by the fact that Willmann (1931a) used the name Oribata
geniculatus (L.) for this species and there are several other names in its synonymy.
Its taxonomy has been unravelled by Grandjean (1950b) and van der Hammen
(1952) but in view of the confusion we cannot be certain that every record of
this species which we list is in fact a record of Phauloppia lucorum.
'['HE ASSOCIATION OF ORIBATID MITES WITH LICHENS
393
Phauloppia lucorum is not only extremely common in lichens but also very
abundant in them. Seaward (1974) found it swarming in lichens, just as nearly
100 years ago Michael (1888) [Notaspis lucorum] reported it swarming in lichens
at Land's End. Fieldwork data obtained by Colloff (in lilt.) from four maritime
sites in England and Scotland contain the following observations: 53 adults were
extracted from 0.5 g (wet wt) of Xanthoriaparietina at one site and 187 adults and
46 nymphs were extracted from 12 g (wet wtj of the same species at another site;
a 2 g sample of Ramalina siliquosa from a further site contained 53 adults which
were associated with the blackened bases of the thalli, and although several
pieces were dissected, no mites could be found in the medulla (cf. Gjelstrup &
Smhting, 1979).
Hull (1914a, 1915) [Notaspis lucorum] collected it in moss as well as lichens,
but he notes that it usually feeds on lichens on trees and walls and reports
shaking out specimens like dust from lichens on the branches of trees (Hull,
1916). Other authors who have collected the species either from arboricolous or
saxicolous lichens are: Oudemans (1900) [Eremaeus lucorum]; Andrt i1925);
Sellnick (1928) [Phauloppia conformis (Berl.)]; Cooreman (1941) [Lucoppia
lucorum]; Dalenius (1950) [Orzbata geniculatus (L.)]; Strenzke (1952); van der
Hammen (1952); Pschorn-Walcher (1953); Willmann (1931a, 1933, 1937,
1939a, 1952, 1956) [ Oribata geniculatus L.]; Schweizer ( 1956) [Phauloppia conformis
Berl.]; Knulle (1957); Pschorn-Walcher & Gunhold (1957) [Phauloppia conformis
Berl.]; Kunst ( 1959); Travt (1958, 1960, 1961, 1963); Rajski (1968); Tarman
(1973); Gilbert (1976); Andri: (1976a, b); Gjelstrup (l978a, b); Gjelstrup &
Smhting (1979). MacXeill (1966) observed both adults and nymphs of
P. lucorum breaking off and eating pieces of lichen thallus.
In view of these records it might seem more reasonable to place the species in
Group A. However several of the above authors have collected the species from
moss as well as lichens and other workers have reported the species only from
non-lichen biotopes, usually moss (Koch, 1841 [Zetes lucorum]; Berlese, 1895b
[Oppia conformis Berl.]; Voigts & Oudemans, 1905 [Eremaeus lucorum]; Halbert,
1907, 1915 [Notaspis lucorum]; Franz 1954 [Phauloppia conformis Berl.]; TarrasWahlberg, 196 1 ; Csiszar & Jeleva, 1962; Bernini, 1971; Polderman, 1974; Bonnet
et al., 1975; Iturrondobeitia & Subias, 1981). Willmann (1942a) [Oribata
geniculatus (L.)] recorded the species from mineral springs and believed that the
mites were feeding on fungal and bacterial spores in the skin of mould on the
surface of the water. However in further records from mineral springs Willmann
(1950) suggested they might be accidental strays from the surrounding
vegetation, a view which Pax (1948), who had found the species in sulphur
springs, appears to have shared. Karppinen ( 1 9 5 8 ~ )[Oribata geniculatus L.] was
surprised to collect it from the ground in a Myrtillus dominatd habitat, since as
he says it is above all found in arboricolous mosses and lichens. Nordberg ( 1936)
found that the species was very common in the nests of birds, which is not
surprising for a mite which is abundant in lichens and mosses. He also notes its
occasional mass appearance in houses, an occurrence which Oudemans ( 1900)
also mentions.
Travi: (19631 does not include this species in his section on lichenophagous
forms and until more work has been done on its feeding habits it seems best to
place it in Group B. It is possible, nevertheless, that it may eventually, at least in
its early stages, be found to be lichen-restricted in its diet. Grandjean (1948b),
394
E. L. SEYD AND M. R. D. SEAWARD
for instance, states that it lives on the bark of trees, on rocks and on walls,
provided that their surfaces are covered with lichens. He points out that
P. lucorum is an active climbing mite and that it is not surprising if isolated
specimens are found in diverse habitats. Oudemans (1929) [Oribata geniculatus
(L)] states that it has been popularly known as the vagabond or wandering
mite, and he also notes records from lichens by other authors.
Trhypochthonius cladonicola (Willmann, 1919)
Willmann ( 1919) first described this species from specimens he collected in
Cladonia from four German localities, and in his list of the Oribatei of Germany
(Willmann, 1931a ) the only biotope given is Cladonia of heath and moorland. I n
a later study Willmann (1933) says that he never found it in any habitat other
than Cladonia. Other workers who have reported the species from Cladonia
include Dalenius ( 1950), van der Hammen ( 1952), Sellnick ( 1928, 1960),
Karppinen (1962) and Gjelstrup (1978b). Kniille (1957), however, mentions
other biotopes besides Cladonia, and Tarras-Whalberg (1952) and Strenzke
(1952) also specify moss as well as Cladonia as habitats, though the latter notes
that the species is characteristic of lichen-covered biotopes. Franz (1943) found
one specimen in Nardetum but he believed that it got there from neighbouring
lichens.
O n the other hand in some publications by several of these authors, nonlichen biotopes such as forest soil, moss and Sphagnum are the only ones given
(Willmann, 1928, 1942a; Sellnick, 1929; Karppinen, 1958c, 1966; Dalenius,
1960; Tarras-Wahlberg, 1961) . Schweizer ( 1948) collected a single specimen in
poor meadowland in the Swiss National Park.
Tricheremaeus serratus (Michael, 1885)
Michael (1885, 1888) collected this species from several places in Britain
(Warwickshire, Westmorland, Cumberland, Staffordshire, N. Wales), yet we
have been unable to find any other author who has recorded the species from
this country. Moreover the only record from abroad that we have been able to
trace is that of TravC (1960, 1963) from the PyrCnCes Orientales.
Michael (1888) observed that the species is usually to be found among lichens
in dry places and TravC’s (1963) specimens were obtained by sweeping lichens,
mosses and liverworts.
The species bears a close resemblance to 1.nemossensis Grandjean which
Grandjean (1963) collected in the neighbourhood of Mont DorC (Puy-de-Dome)
from moss on the trunk of a birch.
Group C species
Adoribatella punctata Woolley, 1967
Woolley (1967) collected specimens of this new species in moss near Deadman
Pass, Colorado, and in lichens from Cameron Pass.
Alaskozetes antarcticus (Michael, 1903)
TragArdh (1907a, b) [Notasp; antarctica] found this antarctic species in lichens
as well as in moss, under stones, on damp sand and in birds’ nests. Dalenius &
Wilson (1958) [Halozetes antarctica] give its habitat as mosses and lichens and it
THE ASSOCIATION OF ORIBATID MITES WITH LICHENS
395
seems likely that Richters’ ( 1907) [Notaspis antarctica] specimens were collected
from these two habitats.
Block (1977, 1980) and Young & Block (1980a, b) give its habitat as crustose
lichens, algae, intertidal debris, penguin guano, seal wallows and birds’ nests,
and they comment that it is principally a scavenger feeding on detritus, mostly
of vertebrate origin. Covarrubias (1968) found it in similar habitats and he
notes that its lowest average densities are in mosses and lichens, but on Signy
Island it appears to be common in lichens as in dry moss (Goddard, 1979).
Travi. (1981b) records it from lichens and algae from the Kerguelen
Archipelago.
Ameronothrus bilineatus (Michael, 1888)
Michael (1888) [Scutovertex bilineatus] found this species in large numbers on
algae or crawling on weed and rocks at the bottom of freshwater pools at Bull
Bay, Anglesea. It is certainly a sea-shore form (Hull, 1916 [Scutovertex bilineatus] ;
Sellnick, 1928, 1960; Schulte, 1976) but it appears to have been collected
mainly from algae and shore detritus. However Halbert (1920) [Scutovertex
bilineatus] found it under limestone flakes at Malahide, where there were
encrusting lichens, and on this account it figures in the list of Evans et al. (1961)
of intertidal orange-lichen zone species.
Ameronothrus lineatus (Thorell, 1971 )
This is not only a littoral species but also a predominantly arctic and subarctic one. Thorell’s (187 1 ) [Eremaeus lineatus] original description was based on
specimens from Spitzbergen, and it has been recorded there by: Troussart
(1894) [Eremaeus lineatus]; Hull (1922) [Scutouertex lineatus] in moss; Thor (1934)
in coprophilous lichens; Block (196613) in Dryas litter and by Karppinen 1,1967)
in dry tundra. There are records, too, from Greenland (Graverson, 1931;
Hammer, 1937, 1944); Canada (Hammer, 1952a) on grassy coastal meadows
submerged at high tide; and from Novaya Zemlya, Bear Island and Siberia
(Koch, 1878 [Eremaeus lineatus]; TragArdh, 1900, 1905 [Scutovertex lineatus]).
Sellnick (1908b) recorded it in moss from the Faroe Islands and from nonlichen habitats on the Swedish coast, and Karppinen (1966) in algae on the
Finnish coast. However, in Norway, Thor (1937) again found it was common in
coprophilous lichens. Hull (1916) [Scutovertex lineatus] recorded it as far south as
the coasts of Anglesea and Ireland.
Schulte (1974, 1976), in his study of the Ameronothridae of the marine
littoral, found the species in saxicolous lichens and he classes it as a lichendweller. However it also figures as an algal-dweller along with A . marinus,
A . bilineatus, A . schusteri and A . schubarti.
Carabodes areolatus Berlese, I9 16
Schweizer (1948, 1956) found the species was abundant in lichens in the Swiss
National Park but he also collected it in moss and under stones and wood.
Willmann (1933, 1956) reported it from lichens, and Bojcova (1931 ) found large
numbers in Cladonia. Rajski (1968) describes it as a forest species, and Sengbusch
(1957) collected it from lichens on oak trees and from moss in Virginia. Dalenius
(1960) recorded it from lichen heath in Swedish Lapland but he also found it in
heath and meadow woodland.
396
E. L. SEYD AND M. R. D. SEAWARD
There are several records from moss including Sphagnum, litter, soil, humus,
etc. (Berlese, 1916; Sellnick & Forsslund, 1953; Franz, 1943, 1954; Willmann,
1931a, 1954; Tarman, 1955; Kunst, 1958; Sellnick, 1928, 1929, 1960;
Karppinen, 1956b, 1971; Bernini, 1971; Wauthy, 1976; Lions, 1978).
Carabodes marginatus (Michael, 1884)
The following authors have collected this species from lichens: Hull ( 1915);
Schweizer (1948); Tarrras-Wahlberg (1952); Sellnick & Forsslund (1953);
Willmann (1933, 1939a, 1956); Knulle (1957); TravC (1963); Hammer (1972);
Gjelstrup (1979a); Seyd (1962, 1981). Other biotopes are included in several of
these records.
It is questionable, however, whether C. maginatus should be included in a list
of lichen-associated Oribatei, since the records from moss, Sphagnum, grass, litter,
decaying leaves, humus, soil, pine needles, etc. is a very long one: Michael
(1884); Oudemans (1896); Hull (1914a); Schweizer (1922); Pax & Willmann
(1937); Willmann (1923, 1928, 1931a, 1938, 1942a, 1943); Strenzke (1952); van
der Hammen (1952); Franz (1943, 1954); Tarman (1958); Kunst (1958);
HalBskovA & Kunst (1960); Dalenius (1960); Sellnick (1908a, 1928, 1929,
1960); Tarras-Wahlberg (1961); Block (1965); Lebrun (1965a); Rajski (1968);
Engelmann (1972); Webb (1972); Pande & Berthet (1975); Solhray (1976);
Lions (1978). Nearly all the records of Karppinen (1956a, b, 1957, 1958a, c,
1962, 1966, 1971, 1977) are from non-lichen biotopes, although in a few cases
lichens formed a part of the ground cover. The species appears to be
particularly common in woodland.
Centroribates uropygium Grandjean, 1928
Grandjean (1928) collected this new species from a diversity of habitats of the
Sierra Morena, Spain, but it was particularly common in mosses and lichens.
Chamobates cuspidatus (Michael, 1884)
This is a common species, Hull (1916) describing it as ubiquitous in Britain.
Hull (1914a) [Oribates cuspidatus] did collect it from lichens, as well as other
biotopes, and his is not the only record from this habitat. Hammer (1937, 1944,
1946), for example, collected the majority of her specimens in Canada from
lichens. Schweizer (1948) found specimens in a lichen crust on moss, as well as
in other habitats, and other authors who have recorded the species from lichens
are Willmann (1956), Sengbusch (1957) and AndrC (1976a).
There are a number of authors, however, who have recorded the species only
from non-lichen habitats: (Oudemans, 1900 [Notaspis cuspidatus]; Hull, 1918;
Willmann, 1931a, 1932; Delenius, 1950; van der Hammen, 1952; Franz, 1954;
Sellnick, 1960; Lebrun, 1965a, b; Solhray, 1976; Gjelstrup, 1978a; Hammer,
1977). Michael (1884) [Oribatu cuspiduta], who first described the species, stated
that moss was its chief habitat. This again is one of those species which perhaps
should not be classed as lichen-associated.
Conoppia palmicinctum (Michael, 1884)
As with Humerobates rostrolamellatus and Phauloppia lucorum, there has been
confusion over the taxonomy of this species, owing to the fact that Berlese’s
(188513) Oppia microptera appears to be synonymous with Michael’s (1884)
THE ASSOCIATION OF ORIBATID MITES WITH LICHENS
397
Leiosoma palmicinctum (see Grandjean, 1936b; Rafalski, 1966). Moreover Berlese
(1908), who created the new genus Conoppia, with Oppia microptera as the type,
went on to create another new genus Phyllotegeus, with Leiosoma palmicinctum as
the type (Berlese, 1913b), not realising that the specimens of C. microptera were
really nymphs of P. palmicinctum. (For details see Grandjean, 1936b.) Not all
authors, however, have accepted the synonymy, Willmann ( 1939c) believing
they are two distinct species, a view accepted by Balogh (1943) in his list of
Hungarian species.
Michael ( 1884) [Leiosoma palmicinctum] found that larvae and nymphs of the
species appeared to be restricted to lichens at Land’s End, though he remarks
that the adults are of a “more wandering disposition”. He was able to rear a
specimen from egg to adult in a culture cell containing lichen material as a food.
Turk ( 1972) [Phyllotegeus palmicinctum] obtained specimens of the species from
lic,hens and moss on a stone wall at Cambourne, Cornwall.
The European records, however, have come from a variety of biotopes other
than lichens, such as moss, forest litter, soil and under stones, most of them
appearing under the name of either Conoppia microptera or Phyllotegeus palmicinctum
(Willmann, 1931 a, 1938, 1 9 3 9 ~ Franz,
;
1943, 1954; Schweizer, 1956; Rafalski,
1966; Ptrez-Ifiigo, 1970, 1972; Bernini, 1971; Tarman, 1973; Subias &
Iturrondobeitia, 1977; Iturrondobeitia & Subias, 1981). Several of these records
were from woodland litter, which leaves open the possibility that some of the
specimens originated from lichens growing on trees.
QmbaeremaeuJ cymba (Nicolet, 1885)
Rajski (1968) states that this is an arboricolous species, which is found
accidentally in litter and soil. Certainly most, though not all, of the records have
been either from trees or in forest habitats (Michael, 1879, 1888; Berlese, 1886,
1898; Hull, 1916; Niedbala, 1969 and many of the authors listed below). The
species appears to be present always in small numbers.
Lichen records are provided by: Willmann ( 1933), Grandjean ( 1948a,
1950a); Strenzke ( 1952) who describes it as characteristic of lichen-covered
biotopes; Schweizer (1948, 1956); Sellnick (1960); TravC (1956, 1963) who
recorded it from many localities in the PyrCnCes Orientales, mainly from lichens;
Andri: (1976a, 1979) and Gjelstrup (1979). Franz (1954), though he collected
the species in the soil, litter and moss of woods, notes that its normal habitat is
arboreal mosses and lichens.
Non-lichen records are provided by: Oudemans ( 1900a); Schweizer 922)
Andri: ( 1925); Sellnick ( 1 92 1, 1928, 1929); Willmarin ( 193 1a ); Thor 937)
Franz (1943); Dalenius (1950); van der Hammen (1952); MihelEiE 95 7)
972)
Kunst (1957, 1958); Lebrun (1965a) b); Bernini (1971); Engelmann
Karppinen (1958c, 1972); PCrez-Iiiigo (1971, 1975); Mahunka (1974, 977)
Lions (1978); Iturrondobeitia & Subias (1981) and Martinez 11981).
Diapterobates notatus (Thorell, 1871)
This is an arctic species, its range extending from Alaska through Canada,
Jan Mayen, Swedish Lappland, Spitzbergen, Bear Island, Franz Josef Land to
Siberia. ‘Thor 1930a, 1934) [Murcia nolata] found it on Spitzbergen in lichens,
mosses and Salix vegetation, and Niedbala (1971) collected it in large numbers
in the litter from lichens, mosses and phanerograms. Seniczak & Plichta ( 1 978)
398
E. L. SEYD AND M. R. D. SEAWARD
found that a t Hornsund, Spitzbergen, the species constituted a high percentage
of the oribatid fauna of two lichen habitats and one of Saxzfraga, with lesser
numbers in a moss habitat. Summerhayes & Elton (1928) [Sphaerozetes notatus]
recorded it from localities in Spitzbergen rich in lichen species as well as mosses
and phanerograms, and Karppinen’s ( 1967) [ Trichoribates notatus] samples from
dry tundra may well have included lichen material.
Block (1966b), on the other hand, collected it in Dryas litter on Spitzbergen,
and Hammer (1952a) [ Trichoribates notatus] from moss in Canada, and from
humus and leaves in Canada (Hammer, 1955). I n Alaska she collected it from a
bog, as well as in dry leaves between moss cushions (Hammer, 1967a).
Bohnsack’s ( 1973) Alaskan samples containing the species consisted of tundra
turf. The records of other authors from Spitzbergen and other parts of the
species’ range do not give habitat data (Thorell, 1871; Koch, 1878; TragHrdh,
1900, 1905, 1910; Hull, 1922).
Eremaeus oblongus C . L. Koch, 1836
Both Michael (1879, 1888) [Notaspis oblonga] and Hull (1916) described this
species as being generally distributed and common.
Lichen records include those of Hull (1914a, 1915) [Notaspis oblonga];
Schweizer (1948); Dalenius (1950); Willmann (1939a, 1956); Sengbusch (1957);
Travt (1960, 1963); Seyd (1966); Rajski (1967); Tarman (1973) and AndrC
(1979).
Non-lichen records (moss, litter, soil, under bark and stones, conifer needles
etc.) are provided by Hull (1918) [Notaspis oblonga]; Schweizer (1922); Sellnick
(1908a, 1928, 1929); Irk (1939); Hammer (1944); Strenzke (1952); van der
Hammen ( 1952); Franz ( 1943, 1954); Willmann ( 1931a, 1932, 1942a, 1954);
Tarman (1955); Lebrun (1965b); Karppinen (1957, 1958c, 1966); Niedbala
(1969); Bernini (1971) and Bonnet et al. (1975).
Many of these lichen and non-lichen records are either corticolous or
saxicolous in woodland areas, but this is by no means always the case. Block
(1965) found the species on high moorland and Seyd ( 1966) on a Lakeland
peak, and Rajski (1967) says it occurs in open terrain in mosses and lichens. Pax
& Willmann (1937) recovered it from moss in waterfalls, and Nordberg (1936)
from birds’ nests.
Ghilarovus hispanicus Subias & Ptrez-Iiiigo, 1977
Subias & Pkrez-Iiiigo (1977) collected 35 specimens of this species in the
Sierra de Cazorla at altitudes between 800 and 1400 m. The specimens were all
taken from rocks, and the authors comment that the species lives in mosses and
lichens and also in the debris which accumulates in the rock crevices (see also
Subias, 1980).
Halozetes belgicae (Michael, 1903)
This antarctic species was found by TragHrdh (1907a, b) [Notaspis belgicae] in
lichens under stones and in lichens and mosses and Dalenius & Wilson (1958)
[Petorgunia belgicae] give its biotopes as lichens and mosses. The records of
Wallwork (1963, 1970, 1972b, c) and of Covarrubias (1968), however, add
other habitats to these such as algae, grass, the faeces of marine mammals and
birds, and birds’ nests. Block (1979) recorded the species from the undersides of
THE ASSOCIATION OF ORIBATID MITES WITH LICHENS
399
rocks, and Goddard (1979) reported it present on Signy Island in small numbers
under stones on dry moss and lichens. Travi: (1981b) reports its presence in
lichens and algae from the Kerguelen Archipelago.
Halozetes crazetensis (Richters, 1908)
TravC (1976),who found the species abundant on the Kerguelen Archipelago,
described it (Travk, 1979) as essentially saxicolous and says that it lives in
lichens and mosses, although in a later paper (Travt, 1981b) he found that its
greatest density was in moss. Watson (1967) also gives its habitat as lichens on
rocks and cave walls, but he adds other biotopes such as algae, mosses, plant
litter and nest material. Wallwork (1963) also mentiones lichens in his list of
biotopes on Macquarie Island, but on Campbell Island (Wallwork, 1966),
Crozet Island iwallwork, 1972b) and South Georgia (Wallwork, 1972c) lichens
do not figure among a variety of habitats mentioned.
Halozetes intermedius Wallwork, 1963
There was exceptional density of this species in crustaceous lichens of the
intertidal zone of the Kerguelen Archipelago (Travi., 198 1b) . I t appears to feed
on algae as well as lichens and Wallwork (1963) found it in several non-lichen
habitats.
Hermannia reticulata Thorell, 187 1
The taxonomy and distribution of this species have recently been reviewed by
Seyd (1981). It is an arctic and subarctic species with a number of southern
‘outliers’ in Britain and Europe but, unlike Calyptozetes sarekensis, not all of these
are montane sites.
The species has been collected from lichens and mosses on Bear Island (Thor,
1930b), Spitzbergen (Niedbala, 1971) and the Faroe Islands (Gjelstrup, 1978a),
and Hammer (1952a) collected it in Canada from Cussiope heath and lichen
heath, both of which are rich in lichens, particularly the latter, and Haarlov
(1942) also found it on Cassiope heath in Greenland. Behan & Hill (1978) regard
the species as a panphytophage in the North American arctic.
O n the other hand there are records from localities in Greenland Uclrgensen,
1934; Hammer, 1944, 1946, 1952b), Spitzbergen (Thor, 1930a; Block, 1966b),
the Faroes (TragHrdh, 1931), Norway (Willmann, 1929; Thor, 1937) and
Swedish Lappland (Woas, 1978) which do not mention lichens as part of the
plant habitat. Moreover, so far as the southern ‘outliers’ are concerned, Hull
(1918) [Hernmunnia quadriserzata Banks] found it in dead wood, Halbert (1920)
on a green algal-like weed, and Andri. (1925) and Oudemans (1927) under
stones, although both Michael (1888) and Seyd (1981) recorded it from lichens
as well as mosses. Several records of this mite are from coastal sites, and Evans
et al. (1961) include it in their list of species characteristic of the orange lichen
zone of the sea shore. Colloff (in litt.) has recently extracted adults of this mite
from Anaptychia fusca, Lecanora atra and Xanthoria parietina from a Cornish
maritime site.
Hermannia scabra (L. Koch, 1878)
This species also has a wide distribution in arctic and subarctic regions.
Koch’s (1878) specimens were from Novaya Zemlya and Siberia, and TragHrdh
400
E. L. SEYD AND M. R. D. SEAWARD
(1900, 1905) also found it on Bear Island. Banks (1923) [Lohmannia scabra]
recorded it from the Pribilof Islands, Graversen (1931) from Greenland, and
Hammer (1944, 1955) from Greenland and Alaska. Behan & Hill (1978)
describe it as a North American panphytophage. Like H. reticulate this species is
known from Britain and Europe and though predominantly a littoral form, it
has been recorded from inland sites as well.
Halbert (1915, 1920) found it under limestone flakes in the orange lichen
zone of the sea shore, but it was also present in the Peluetia zone and in the old
nests of birds. Hull (1914b) collected it in lichens, mosses and grass on the coast,
and Sellnick (1949) from lichens and other biotopes on the coast of Sweden; it
also appears on the orange lichen zone list of Evans et al. (1961). Hammer’s
(1944, 1955) specimens in Greenland and Alaska were from lichens and moss,
and Gjelstrup & Srachting (1979) recorded it from the lichen Ramalina siliquosa.
Michael (1888) [Nothrus nodosa Michael] collected it a t coastal sites from lichens,
mosses and algae, and from lichens at an inland site.
However, other authors have recorded the species only from non-lichen
biotopes, usually moss, both a t coastal sites and inland (Sellnick, 1908a; Hull,
1914a, 1915, 1916; Willmann, 1931a, 1937; Franz, 1954; Pschorn-Walcher &
Gunhold, 1957; Sengbusch, 1957; Dalenius, 1950, 1960; Karppinen, 1971).
Voigts & Oudemans (1 905) found specimens under bricks in Bremen, and van
der Hammen (1952) says that Oudemans specimens were collected on the shore,
but he gives no details of habitat. Schweizer (1926) obtained specimens from
saltwater springs.
Woas (1980) has recently described a new species of Hermannia, H. intermedia
Woas, from lichens and Enteromorpha in the intertidal zone of the Gironde,
France.
Lamellovertex caelatus (Berlese, 1895)
A1though Berlese’s original description of this species was from specimens
found in moss at Desenzano (Lake Garda) and Schweizer (1922) collected it in
moss in the Swiss Alps, Bernini (1976b) found it in lichens on branches lying on
the ground a few km from Siena.
Lepidozetes singularis (Berlese, 1910)
This is an alpine species, Willmann (1931a) collecting it from moss in the
Black Forest, from lichens and from the bark of a pine in the Glatzer
Schneeberg range (Willmann, 1939a); Franz (1954) in woodland litter, conifer
needles and grass heathland in the Alps and Schweizer (1948) in a lichen crust
in the Swiss National Park. Schatz (1979), who collected specimens in grass
heathland and lichen heath in the Hochgebirge of the Tirol, carried out culturefeeding observations and concluded that the species was a panphytophage.
Travi. (1963) found it in moss, lichens and liverworts of the Massane, Pyrknkes
Orientales and in arboricolous lichens of the Hautes PyrCntes. C5lug5r &
Vasiliu (1980) recorded it from forest biotopes and lichens were not present in
the habitats from which Bernini (1971) collected it.
Licneremaeus discoidalis Willmann, 1930
Willmann (1930) recorded this new species from Guatemala from
arboricolous mosses and lichens. Grandjean ( 1931a) described a new species,
THE ASSOCIATION OF ORIBATID MITES WITH LICHENS
40 1
L. exornatus, from the trunks and branches of trees in Venezuela but he does not
say whether these supported mosses and lichens.
Liodes theleproctus (Hermann, 1804)
According to TravC (1960) this is both a saxicolous and arboricolous species.
Travt (1963) collected it in these habitats in the Pyrtntes Orientales from
lichens, mosses and liverworts, always in small numbers. Grandjean (1936a) also
found it on the bark of trees with a covering of mosses and lichens and on
lichen-covered rocks. Several other authors have collected it from moss, often on
trees, and from woodland litter and humus (Michael, 188% INorthrus
theleproctus]; Schweizer, I922 [Neoliodes theleproctus]; Franz, 1954 [Peroliodes
theleproctus]; Sellnick, 1908a, 1960; Mahunka, 1974; Lions, 1978; Iturrondobeitia
& Subias, 1981). Halbert (1907) found it on the undersides of stones among
heather on Lambay Island.
Macquarioppia slriata (Wallwork, 1963)
Watson (1967) collected this species from lichens in a cave on Macquarie
Island, as well as from litter and algae, but Wallwork (1970, 197213) does not
mention lichens in his list of habitats for this species. O n the other hand, among
the biotopes in which Travt (1979) found the species, were crustaceous and
foliaceous lichens from the Kerguelen Archipelago.
Metrioppia helveticn Grandjean, 193 1
Grandjean (1931b) found this species in the environs of Andermatt at 1600 m
in mosses and lichens. The collection also contained humus, and Grandjean
suggests that the mites may feed partly on the humus and partly on the
cryptogams.
Micreremus brevzpes (Michael, 1888)
Michael (1888) [Errmaeus brevipes] found that this species was abundant on
the foliage of trees, especially oaks, in Epping Forest and the New Forest. Van
der Hammen f 1952) also collected it from oaks, and Rajski (l968), although he
found the highest frequency of the species was in pine woods, notes that oak
trees are its usual habitat. Tarras-Wahlberg (1961), however, found it was
present in pine sweepings.
Franz (1954) collected the species in woodland litter and soil, but he observes
that corticolous lichens and mosses are its normal biotopes. Grandjean f 1950a),
Lebrun (1976, and Andri. (l976a) all found it was present in corticolous lichens.
Sdlnick (19281 and Willmann (1931a) give mosses and lichens as habitats.
Niedbala (1969) collected it from the branches of trees in the environs of
Poznan, but in Spitzbergen (Niedbala, 1971) he found i t in the litter of mosses,
lichens and vascular plants. Halaskova & Kunst (1960) obtained specimens
from bark lichens. Travi. (1963) records only a few specimens from the
Massane one from corticolous lichens and a few from plants and soil.
Other records, mostly from forest moss or litter, are: Sellnick (1929); Csiszar
& Jeleva (1962); Lebrun (1965bj; Bernini (1969); Mahunka (1974); Bonnet
et al. (1975); Karppinen (1958c, 1972, 1977); Iturrondobeitia & Subias (1981)
and Martinez (1981). Christensen (1980) found a few specimens in dead wood.
18
402
E. L.SEYD AND M. R. D. SEAWARD
Oppia crozetensis (Richters, 1907)
Watson (1967) reported this species from Macquarie Island in lichens, litter,
algae and nest material, and TravC (1979) includes crustaceous and foliaceous
lichens among the biotopes in which it is found in Kerguelen Archipelago.
Wallwork (1963, 1970, 1972b, c), on the other hand, in his papers on Pacific
island Oribatei, does not include lichens in his list of biotopes in which he found
the species.
Oribatella calcarata (C. L. Koch, 1836)
This species is synonymous with Michael’s (1884) Oribata quadricornuta. Van
der Hammen ( 1952) prefers to make Oribatella quadricornuta (Michael) the senior
synonym on the grounds that Koch’s description is insufficient, but most authors
have followed Willmann’s (1931a) use of the name Oribatella calcarata (C. L.
Koch) as the senior synonym.
Evans et al. (1961) include 0. calcarata in their list of species characteristic of
the intertidal orange lichen zone, no doubt on the basis of Halbert’s (1920)
[Oribata quadricornuta] record of specimens under stones in that zone. Willmann
(1939b, 1952) also collected specimens from coastal areas, though not from
lichens, and he also reported it from moss, including Sphagnum, and humus from
inland localities (Willmann, 1931a, 1938, 1939a, 1956). Koch ( 1836) [Oribates
calcarata] described the species from specimens collected in woodland, and
several authors have also found it in woods (Michael, 1884; van der Hammen,
1952; Franz, 1954; Dalepius, 1960; Lebrun, 1965a, 1965b; CBlugk- & Vasiliu,
1980). Schweizer (1948) found it in a lichen crust, as well as in moss and D v a s
and Erica heath, and AndrC (1976a, 1979) [Oribatella quadricornuta] gives records
from corticolous lichens.
Lebrun (1971) regards this form as a non-specialized feeder, and many
authors have recorded it from habitats such as moss, litter and soil (Franz, 1943;
Dalenius, 1950, 1960; Schweizer, 1956; MihelEiE, 1957; Frank & Zivkovitch,
1960; Csiszk & Jeleva, 1962; Karppinen, 1966; Luxton, 1966b; Engelmann,
1972; Bernini, 1971, 1977b; Gjelstrup, 1978a).
Oribatula exudans TravC, 1961
Travi. (1961, 1963) collected this species from the Massane area, PyrCnCes
Orientales, the Hautes Pyrtnkes and in one sample from the area around
Banyuls-sur-Mer. It was abundant in samples of mosses and lichens, but he also
obtained some specimens from phanerogams, and two soil samples contained the
species. An examination of the faecal pellets of 0. exudans showed the presence of
pollen grains, fungal spores, mycelia and portions of thallus.
Oribatula thalassophila Grandjean, 1935
Grandjean (1935) found this species on rocks covered with barnacles on the
Brittany coast. He also collected it in algae in the supra-littoral zone, but Schuster
(1979) states that it is a lichen feeder.
Oribatula venusta Berlese, 1908
Several records of this species, originally described by Berlese (1908) from
Norway, have been from the sea shore and it is included in the orange lichen
zone list of Evans et al. (1961). Halbert (1920) found it under stones on the
Mayo coast in the orange lichen zone, and in lichens and moss in sandhills at
THE ASSOCIATION OF ORIBATID MITES WITH LICHENS
40 3
Portmarnock. Niedbala ( 197 1) [agoribatula uenusta] recorded it from lichens,
moss and phanerogams on the sea shore of Spitzbergen, and Karppinen (1967)
also collected it from dry tundra and grassy fellside on Spitzbergen.
Dalenius (1960) collected specimens on the shore of the Tornetrask Territory,
Swedish Lappland, but he also found it in a variety of other habitats including
forests, meadows, heaths and bogs. He recorded it too from southern Sweden
from various biotopes (Dalenius, 1950). Franz (1943, 1954) collected it in moss,
humus, soil and woodland litter in the Alps, and Pax & Willmann (1937) from
stones, wood and moss in waterfalls of the Glatzer Schneeberg range. Fujikawa
(1972) also found the species in non-lichen biotopes.
Parachipteria petiti Travk, 1960
This species was collected by TravC (1960, 1963) from several species of
Parmelia in the PyrhCes Orientales, and also from mosses and hepatics. Travi:
found that it was most abundant in moss.
Phereliodes wehnckei (Willmann, 1930)
Willmann ( 1930) [Cymbaeremaeus wehnckei] recorded this species from
Guatemala from arboricolous mosses and lichens. Grandjean ( 1931a) collected
specimens in Venezuela but he gives no biotope for them. The new genus
Phereliodes was later created for this species by Grandjean (1964).
Porokalumma rotunda (Wallwork, 1963)*
TravC (1981b) states that this species is present in lichens and algae of the
supra-littoral zone of the Kerguelen Archipelago.
Poroliodes farinosus (C. L. Koch, 1840)
Gjelstrup (197813) states that the most frequent habitat for this species is
lichens, particularly lichens growing on trees and rocks, and Gjelstrup &
Smhting ( 1979) obtained specimens from Ramalina siliquosa on Bornholm.
Sellnick (1949) collected one specimen from lichens on the Swedish coast.
Most of TravC’s (1963) specimens were taken from saxicolous and
arboricolous mosses and hepatics, but Parmelia is also noted as one biotope. Both
Grandjean (1936a) and Karppinen (1966) reported the species present in several
non-lichen habitats, although Parmelia does figure in the latter author’s list of
habitats.
Pseudachipteria magnus (Sellnick, 1928)
This is an ‘alpine’ species recorded from high ground in Europe, Britain and
Scandinavia, in most cases from moss (Sellnick 1928 [Notaspis magnus];
Willmann, 1931a [Notaspis magnus]; Schweizer, 1948 [Notaspis magnus]; Franz,
1954 [Parachipteria magna]; Sellnick, 1960 [Parachipteria magna]; Karppinen, 1971
[Achipteria magna]; Bonnet et al. (1975); Subias, 1980; Seyd, 1966, 1968, 1981).
This is a doubtful Group C species and we are including it here since Travti
*This is the last of the antarctic species to appear on our list. Several other antarctic species have been found in
habitats that include lichens, but they are not included in the List because their habitats are still not sufficiently
researched. The antarctic species in general present a problem in that there are many confusions and
uncertainties regarding their taxonomy (Travt 1981a). A review of the antarctic species of Oribatei is given by
Wallwork (1973).
404
E. L. SEYD AND M. R. D. SEAWARD
(1960, 1963) found it in saxicolous and arboricolous lichens and mosses of the
Massane Forest and also in lichens and mosses of the Hautes Pyrtntes.
Scutouertex minutus (C. L. Koch, 1836)
Several authors have collected this species in lichens (Dalenius, 1950;
Pschorn-Walcher, 1953; Pschorn-Walcher & Gunhold, 1957; Knolle, 1957).
Both Willmann (1931a) and Sellnick (1960) give moss as well as lichens as the
biotopes of this species and a number of collections have been made from
habitats in which lichens do not figure (Strenzke, 1952; van der Hammen, 1952;
Franz, 1943, 1954; Tarman, 1955; Schweizer, 1956; Frank & Zivkovitch, 1960;
Csiszar & Jeleva, 1962; Nied bala, 1969; Karppinen, 1966, 196 7, 1971 ) .
Sphaerozetes arcticus Hammer, 1952
Hammer (1952a) found this species was common in lichens, moss and plant
cover in Northern Canada, and she collected it again from lichens and moss in
Alaska (Hammer, 1955).
Strenekea depilata Travt, 1966
Travt (1966) recorded this new species from the Massane Forest near
Banyuls-sur-Mer. He collected it from foliaceous lichens on rocks and from
liverworts and lichens on the trunk of a beech. He gives its habitat as mosses,
lichens and liverworts on rocks and trees.
Tectocepheus sarekensis Tragirdh, 1910
This is one of those species which should perhaps have been deleted from our
list. We are tentatively including it in Group C on the basis of lichen records by
Andrk (1975, 1976a, 1979), Travt (1963), and Gjelstrup & Sochting (1979).
Schatz (1979), who regards the species as a panphytophage on the basis of
culture-feeding observations, found it was present in numerous habitats in the
Tirol but that it was the dominant species in alpine lichen heath. There are
many authors, however, who have recorded it from such diverse habitats as
moss, Sphagnum, litter, grass, leaves and soil (Tragirdh, 1910; Willmann, 1931a;
Franz, 1954; Kunst, 1957; Tarman, 1958; Frank & Zivkovitch, 1960; Dalenius,
1960; Karppinen, 1956a, 1958a, c, d, 1967; Travt, 1963; Murphy & Jaiil, 1964;
Lebrun, 1965b; Rajski, 1968; Niedbala, 1969; Oltrec, 1975; Ptrez-Iiiigo, 1971,
1974, 1975; Mahunka, 1974, 1976; Athias-Henriot & Cancela da Fonseca, 1976;
Zyromska-Rudzka, 1976; Bayoumi, 1979; Iturrondobeitia & Subias, 1981;
Martinez, 1981) .
Tegoribates bryophilus Woolley, 1965
Woolley (1965) collected this new species in Colorado in moss and litter from
Mt. Meeker, and in lichens from the Cameron Pass.
cZygoribatulafrzsias (Oudemans, 1900)
TravC (1961, 1963) collected many of his specimens of this species from
saxicolous lichens of the Mediterranean littoral. He did, however, obtain one
specimen from lichens at Font-Romeu, Hautes PyrtnCes, and other specimens
from soil in the valley of the Baillaury. Other records of this species have,
however, been from habitats in which lichens have not been present. Oudemans
THE ASSOCIATION OF ORIBATID MITES WITH LICHENS
#I5
( 1900) [Eremaeus frisiae]; Oudemans ( 1916) [ Oribatula frisiae]; Sellnick ( 1928);
Willmann (1 93 1a) [Oribatula (agoribatula) frzszae] and van der Hammen (1952),
all obtained their specimens from mosses, and some of Travk’s (1963) specimens
at Banyuls-sur-Mer came from moss as well as lichens. Evans (1952) collected
his specimens in soil.
This concludes the list of Group C species. The point must be made, however,
that there are many other species which have turned up in lichen samples and
which acarologists and lichenologists may therefore come across from time to
time. However, the evidence that any of these species are lichen-associated in
the sense used in this review is weak, and the majority of them may well be
‘accidentals’. Nevertheless we give a short list in alphabetical order of some of
the additional species most likely to occur in lichen samples, and some of them
may of course eventually be classed as Group C forms as more evidence
accumulates:
Banksinoma lanceolata (Michael)
Belba tatrica (Kulcz)
Brachychthonius immaculatus Forsslund
Brachychthonius lapponicus (Trag2rdh)
Brachychthonius perpusillus Berlese
Ceratozetes thienemanni Willmann
Carabodes ajinis Berlese
Carabodes scymnus Hull
Euzetes globulus (Nicolet)
Hemileus initialis (Berlese)
Liebstadia similis (Michael)
Mycobates punctatus Hammer
Niphocephalus nivalis delamari Travk
Nothrus borussicus Sellnick
Oppia ornata (Oudemans)
Oppia quadicarinata (Michael)
Oribatula tibialis (Nicolet)
Scheloribales latipes (C. L. Koch)
Scheloribates pallidulus (C. L. Koch)
Suctobelba grandis europaea Willmann
Tectocepheus velatuJ (Michael)
Trhypochthonius tectorum (Berlese)
Trichoribates trimaculatus (C. L. Koch)
cZygoribatula exarata (Berlese)
,?’ygoribatula exilis IlNicolet)
I n the studies by Hammer (1961, 1962, 1966, 1967b, 1968) of the oribatid
fauna of Peru, Chile and New Zealand, she describes many new species, several
of which she found in biotopes that included lichens. The same is true of her
studies on the Oribatei of Northern Canada (Hammer, 1952a) and Alaska
(Hammer, 1955). However, in many cases only single specimens or very few
specimens were taken and the lichen were always mixed with other vegetation,
chiefly moss, but also liverworts, leaves, litter, etc. Clearly a great deal more
work on these mites is required before any of them can be regarded as lichenassociated.
406
E. L. SEYD AND M. R. D. SEAWARD
ORIBATID-LICHEN SPECIFICITY
Lichens are symbiotic, being composed of a fungal (mycobiont) and an algal
(phycobiont) component. Essentially they can be classified into three growth
forms, crustose, foliose and fruticose, but there are many intermediates. Each
growth form is characterized by a particular arrangement of cortical, algal and
medullary tissues and by different degrees of attachment to the substrate. The
morphology of a mature lichen thallus is almost entirely dictated by the fungus;
the algae occupy only 5% of the volume (9% by weight). Less than 30 genera
of algae are known to enter into lichen symbioses, and in temperate regions, the
unicellular green alga Trebouxia is found in about 70% of lichen species, and the
filamentous green alga Trentepohlia in about 10%; a similar percentage contain
blue-green algae, the commonest genus being Nostoc.
It would be interesting to review those lichen species most commonly
associated with oribatids, and indeed to establish specific relationships between
the two associates, in terms of (a) the protection and shelter provided by the
particular growth form, and (b) the palatability of one or both of the symbionts.
The greatest problem involved in undertaking such a survey stems from a lack of
understanding of the taxonomy of one of the associates by the researcher:
acarologists giving scant attention to the lichens, and lichenologists merely
recording the infestation of specific lichens by unidentified mites. However, our
detailed survey of literature sources has revealed considerable basic information
from which tentative conclusions can be drawn.
It would appear that lichen growth forms in a wide range of habitats are
exploited by oribatids of Groups A and B, but obviously the highly crustose
lichens (e.g. Caloplaca citrina, Candelariella aurella, Lecanora dispersa) will only
provide a food-source; however, perithecia of certain crustose pyrenocarpous
lichens may additionally provide shelter, or at least provide a site in which to
lay eggs. O n the other hand, oribatids of Group C, despite their
panphytophagous feeding habit, would appear to be mainly associated with
Cladonia spp. (the only exception being an ill-defined habitat of Parmelia in
Travt, 1963), and are therefore probably restricted to terricolous (including
mossy and peaty) habitats. Such habitats would present little difficulty to
oribatids with catholic feeding habits, since a wider range of biotopes than could
be found for example on lichen-dominated tree-barks, rocks and man-made
substrates, would be readily available.
The protective value of lichens requires further investigation; for example, the
larvae of Mycobates parmeliae are bright orange, blending with the colour of its
associated lichen Xanthoria parietina, but the adults are more abundant in
bryophytes. Concealment, camouflage and mimicry are further considered in
Gerson & Seaward (1977).
With the exception of the highly specific requirements of certain marine
species exploiting Lichzna pygmaea (with a blue-green alga), all oribatids on
named lichens investigated in our survey are associated with lichens containing
green algae of the Protococcaceae, and with only two exceptions, the
phycobiont is the genus Trebouxia. Until detailed investigations to establish foodpreference(s) of oribatids, similar to those on psocids (Broadhead, 1958), have
been undertaken, we can only speculate as to the palatability of Trebouxxia.
Certain oribatids are known to prefer the lichen thallus, others the asci and
THE ASSOCIATION OF ORIBATID MITES W I T H LICHENS
40 7
spores. The value of the mycobiont as a food-source needs further investigation,
since it may be (a) the sole diet, (b) part of a mixed diet including algae, or (c)
a substitute when the preferred algal diet has been depleted.
Lichens produce many unusual secondary products not found in other plants.
To date several hundred lichen substances have been determined (Culberson,
1969, 1970; Culberson, Culberson & Johnson, 1977), many of which have been
found to be specific to particular genera or indeed species. T h e resistance of
most lichens to attack by insects and micro-organisms may be correlated with
the presence of specific substances. The biological role of these lichen substances
is as yet undetermined, they appear to have no basic metabolic function (cf.
Fraenkel, 1959), but they may in part aid specific oribatid feeders in lichenlocating, or they behave as repellents to grazers (cf. Rundel, 1978), which would
otherwise be inimical to thallial development, thereby enhancing the
competitiveness of one lichen species over another. Through selection, lichenspecific oribatid species have become dependent on smell and taste of these
secondary plant substances, but there is no experimental evidence to date to
show that lichens attract particular feeding and mimicking species (Gerson &
Seaward, 1977). However, panphytophages (non-specific feeders) are considered to possess ecological advantages. T h e association of oribatids with specific
lichens could prove advantageous for the dispersal of spores and propagules
which may be transported on body surfaces or in the gut; culture experiments
have shown that 140/, of faecal pellets contained viable fungal spores (Behan &
Hill, 1978). Soredia of Lepraria incana have been shown to be present on the
dorsal hairs of Tricheremaeus serratus. The distances travelled by oribatids (see
Berthet, 1964) will naturally be significant in lichen dispersal.
Several mechanisms have been proposed to account for the widely reported
lichen resistance to invertebrates, including the specific effects of lichen acids,
the protective gelatinous covering of some lichens, and the presence of chelating
agents in many lichens (Gerson, 1973). The effect of lichen acids is probably
made up of several factors, such as solubility, reduced palatability, outright
toxicity and possible indirect toxicity mediated through the antibiotic effect of
some acids on the symbiotic (gut) microflora of certain grazers (Gerson &
Seaward, 1977). The whole subject of ‘protection from grazers’ is still full of
contradictions: for example, substances belonging to particular chemical groups
serve as protectants to some lichen species but not to others. O u r survey of the
chemical products of lichen species known to be associated with oribatids has
failed to reveal any correlations that would elucidate specific chemical
protections (cf. Gerson & Seaward, 1977: table 111). Larger numbers of both
oribatid and lichen associates will need to be surveyed before such patterns, if
they exist, can be revealed. O u r investigations of numerous genera, such as
Purmelia, have shown no particular lichen substance(s) to be repellent to
oribatids. Why, for example, does P. saxatilis support numerous species in a
variety of habitats whereas P. omphulodes has not been found to be associated
with oribatids, yet both contain similar products (atranorin, lobaric acid,
salazinic acid) ?
As pointed out in Gerson & Seaward (1977), there is a lack of experimental
data about most aspects of lichen-invertebrate associations, and oribatid studies
are no exception. Attention should be drawn to the few instances where both
associates have been named; Travk (1963) provides the most detailed data, but
408
E. L. SEYD AND M. R. D. SEAWARD
even here it is difficult to correlate the specificity of each organism when lists of
oribatids are set against lists of lichens. Only when authors accurately name
both members of the association can some attempt be made to determine
feeding and sheltering specificity. Such observations can then be supported by
experimentation to determine the effect of lichen substances on oribatid grazers
(specificity, mode of action, tolerance and possibly detoxification), the role of
oribatids in lichen dispersal, and the energetics of oribatid faunas of lichens.
DISCUSSION
It is inevitable that in a review of work extending back over 100 years very
many relevant papers will have been missed. The literature concerning even this
restricted field of research is so vast that it would be quite impossible to follow
up all the reference. As Balogh (1972) has written in the Introduction to his list
of the oribatid genera of the world “Like every other specialised branch of
science, Oribatid literature also becomes increasingly difficult for a
comprehensive survey as time passes. Old works are hardly obtainable and
recent publications appear scattered in often obscure technical journals all over
the five continents”. Exactly the same point has been emphasized by Strenzke
(1952).
This said, we doubt if even a completely comprehensive survey of the
literature would have resulted in much alteration to the number and character
of the species listed in Groups A and B, though there would certainly have been
additions to the far less important Group C list. I n any case the main purpose of
this review is to provide a working list of species to which lichenologists and
acarologists can refer when they find specimens of Oribatei in their lichen
samples in order that they can decide whether it is worth following up this
particular association in greater detail.
One fact which must become obvious from this review is that the greater part
of the data concerning oribatid-lichen associations is extremely superficial. As
mentioned in the section on ‘oribatid-lichen specificity’, this is largely because
lichenologists in general have seldom been interested in the micro-arthropod
fauna present in their samples and those that have been interested have not
always had the mites identified. Equally, acarologists have rarely bothered to
get the lichens in their samples identified down to genera, let alone species. Time
and time again acarologists (and regrettably this is true of the senior author of
this paper) have simply referred to their lichen material as ‘lichen’ or at the
most to one or two lichen genera such as Cladonia and Parmelia. Of course there
are important exceptions to this, but work in the future on these associations
should be such that in any further review of this field of research this particular
section becomes the dominant one.
If we are ever to substantially increase our knowledge and understanding of
these associations, then what will be required are both field and laboratory
studies in depth of each particular oribatid species found in association with
particular species of lichens. We shall need many more studies along the lines of
that of Wallwork (1958), who studied the feeding behaviour of some forest
Acarina, in which faecal material and gut contents of mites from field
populations were analysed and related as far as possible to these populations.
This work is certainly difficult, and in carrying it out the dangers of
T H E ASSOCIATION OF ORIBATID MITES W I T H LICHENS
409
extrapolating from laboratory observations to actual feeding habits in the field
must always be borne in mind, as we emphasized in the section on ‘Culturefeeding observations’. Even in those cases which Woodring & Cook (1967) had
in mind, where an oribatid species is always or nearly always found in a very
restricted habitat, laboratory feeding studies should if possible be carried out
together with gut-content analysis. Such studies would certainly result in a
clearer-cut separation between Group A, Aa and B forms.
We also need studies in depth on how far any particular species is using its
lichen-substrate for shelter rather than food, or equally for both purposes, and
also studies on the extent to which any particular species is a microphytophage,
macrophytophage or panphytophage.
Finally we make no apology for returning to the point which was emphasized
in our ‘Introduction’, namely the importance such studies have in arriving at a
better understanding of the relationship of oribatids to soil fertility, a point of
view supported in particular by Kubiena (1955), Luxton (1972), Behan & Hill
(1978) and Lebrun (1979). We are, however, only too well aware of the
difficulty and complexity of the task, for as Waliwork (1976) has said “The soil
and its associated habitats are immensely complex, from the ecological point of
view. We are beginning to understand the extent of the microhabitat diversity
that occurs here, but there is still much more to be learned about the reaction of
species populations to this diversity”.
ACKNOWLEDGEMENTS
We wish to thank M r K. H . Hyatt, Department of Zoology, British Museum
(Natural History) for enabling one of us (ELS) to consult the excellent
Arachnology Section library, especially the extensive reprint collection on the
Oribatei, and also for providing working space during the early months of 1980
and 1981. The General Library and Departmental Libraries (Zoology, Botany,
Entomology, Geology) of the Museum were also used on many occasions and we
wish to thank their staff for their assistance and cooperation. We should also like
to express our grateful thanks to M r D. Macfarlane, Commonwealth Institute of
Entomology, for his interest and help throughout the preparation of this work
and for allowing us to draw on his encyclopaedic knowledge of the literature
and taxonomy of the Oribatei, and to M r M. J. Colloff, Department of
Zoology, University of Glasgow, for permission to quote some of his unpublished
field data.
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