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Stephen Sharnoff
Updated January 30, 1998
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A snail or slug has chewed its way across this lichen on a tree trunk, leaving a series of characteristic scraping marks behind. |
| Photograph copyright Stephen/Sylvia Sharnoff | |
INTRODUCTION
Many species of invertebrates live on and
among lichens,
using them for concealment, shelter, or food. Over half of the
orders of insects have associations with lichens. There are
striking cases of insects mimicking lichens, the most well known
being that of the peppered moth (Biston betularia) on
lichen-covered tree bark (Richardson, 1974). In popular accounts
of the evolution of moth coloration following the industrial
revolution it is not always made clear that the reason tree bark
became dark in industrial areas was not primarily from soot
deposition, but was mainly due to the disappearance of light-
colored lichens from the trunks and branches (see Bishop, et. al.
1975). As the air in Great Britain has become cleaner, lichens
have grown back, and the moths with pale, lichen-like, pre-
industrial coloration have increased their numbers relative to
the darker ones.
Lichens are a part of many food webs which include
invertebrates and their predators. For example small lichen-
eating arthropods, such as mites or barklice, may be preyed upon
by spiders (which sometimes camouflage their nests with lichen)
which are in turn eaten by shrews which themselves are snatched
up by owls or hawks. In an striking example from northern
Sweden, birds, invertebrates, and lichens were all found to be
functionally connected, and all were impacted by forestry practices
(Pettersson, et. al 1995). The authors found that Natural forests had
significantly greater invertebrate diversity than managed forests and
nearly five times as many invertebrates per branch...The number and
biomass of invertebrates were related to the number of lichens,
even after controlling for sampling location and branch size.
Other studies have implicated forestry in the decline of non-
migratory passerine birds in northern Europe through the
destruction and fragmentation of forests, but our study indicates
that it may also reduce foraging habitat quality through a
reduction in lichen abundance.
Many lichen species contain bitter compounds that may
discourage feeding by invertebrates. Lawrey (1986) states, "It
appears...that lichen palatability to invertebrate herbivores is,
at least in part, based on chemical defense, and that chemical
defense is related to nutritional status. As yet, however, very
little is known about the nutritional quality of lichens or the
nutritional requirements of invertebrates that consume lichens."
Invertebrates that are considered pests from the human
economic point of view, such as gypsy moths, may be deterred by
lichens' anti-herbivore chemical defenses (Blewitt and Cooper-
Driver 1990). In the case of the hemlock tree looper moth, which
lays its eggs in lichens in North America, the lichens can be
part of the problem.
Invertebrates often carry bits of lichen with them as they
go about their business; they are considered to be important
agents of lichen dispersal (Bailey 1976).
A short summary, drawn primarily from Gerson and Seaward
1977 but also from Lawrey 1987 and Richardson 1974, of the major
groups known to use lichens follows. Except for listing
terrestrial invertebrates first, the order follows Gerson and
Seaward.
TERRESTRIAL INVERTEBRATES
Class Insecta
Bristletails (Thysanura) and springtails (Collembola)
include many species which eat lichens, and some proturans, tiny
segmented wingless insects, (Protura) are found in lichens. Hale
(1972) found a lichen community in Maryland that had been at
least 50% destroyed by thousands of springtails (Hypogastrura
packardi). The insects were then reared in the laboratory on an
exclusively lichen diet.
Barklice (Psocoptera) are "voracious lichen feeders" (Gerson
and Seaward 1977). Richardson (1974) remarks, "An area of old
oak woods was recently found in East Lothian, Scotland, in which
the tree trunks over an area of 5,000 sq. meters presented a
grotesque appearance, being covered with the white dying remains
of [lichen species] Evernia prunastri and Hypogymnia physodes.
The bark louse responsible was identified as Reuterella
helvimacula...). Of the eight most common pscocid species that
live on larch trees in northern England, two (Elipsocus
melachlani and Reuterella helvimacula) feed primarily on lichen.
Many Orthoptera, from among the katydids (Tettigoniidae) and
grasshoppers (several families) as well as stick insects
(Phasmatodea) mimic lichens. E.S. Ross (personal communication)
has observed a preying mantis (order Mantodea) camouflaged as a
lichen; a similar example was found in the Congo by Farb (1963).
Some webspinners (Embioptera) eat lichens; they live in silk
tunnels which they construct. Examples have been reported from
the Galapagos Islands (Ross 1966) and Australia (Anon. 1970).
Butterflies and moths (Lepidoptera) have "the richest
association with lichens" (Gerson and Seaward, 1977). There are
many examples of mimicry, in fact many moth names reflect the
resemblance, such as the "Pale lichen moth" (Crambidia pallida)
and the "Powdered lichen moth" (Bruceia pulverina). Some
lepidopteran species live in lichens, and numerous species of
caterpillars eat lichens. Bagworms (Psychidae) in particular
include many species that are lichen feeders. A noctuid moth,
Zanclognatha theralis, in Tennessee, was discovered to both feed
on Usnea strigosa and to look remarkably like the lichen (Sigal
1984).
Many beetles (Coleoptera) live on or in lichens. An amazing
example from New Guinea is that of the weevil (Gymnopholus
lichenifer) whose back is typically covered with a whole
community of living lichens. The lichens function as camouflage
for the beetle, and they support a diverse group of other
invertebrate species including "nematodes, rotifers, psocids and
mites" (Richardson, 1974).
Some lacewing larvae (Neuroptera) cover themselves with
lichens for camouflage. In the southeastern region of the United
States, these may be frequently seen on the trunks of hardwood
trees, noticeable as bits of "wiggling lichen" moving across the
bark (personal observation).
Some examples are known of flies (Diptera) associated with
lichens, and a number of studies have found ant (Hymenoptera)
nests built underneath lichen mats; Bailey (1970) observed the
dispersal of lichen propagules by ants.
Some species of Termites (Isoptera) in the genus
Hospitalitermes feed exclusively on lichens (Kalshoven 1958).
Class Arachnida
The Oribatei, or oribatid mites (Acari: Cryptostigmata),
also called beetle or moss mites, live on lichens in great
numbers and in varying degrees of dependence on them. These
mites play important roles in soil energetics and soil fertility
(Seyd and Seaward 1984). At least one species deposits its eggs
in lichens; it secretes a substance which causes the lichen to
swell and grow over the eggs, protecting them. Oribatid mites
are also the largest category of lichen feeder (Lawrey, 1987).
There is an extensive literature on the association between
oribatid mites and lichens; for a thorough review and large
bibliography see Seyd and Seaward, 1984.
Prostigmatid mites have also been reported to feed on
lichens. Dense populations of Protostigmatids have been found in
Antarctic lichens. Gall mites (Eriophyoidea) form galls in
lichens. Astigmatid mites have been associated with lichens in
the subantarctic islands. Anyone who has spent substantial time
observing lichens in North America will have noticed the frequent
occurrence of bright red mites on lichen thalli.
Some spiders (Araneae) disguise their webs with lichens, and
the giant crab spider of Trinidad constructs nests of lichens in
the genus Usnea. Many spiders live in tundra lichens.
There have been a few reports of diplopods (millipedes)
eating lichens, and of isopods and amphipods (small crustaceans
such as pill bugs and beach fleas) associated with lichens.
Many snails and slugs (class gastropoda) feed on lichens.
It is not uncommon to find trails left in a lichen by the
mouthparts of snails (Peake and James, 1967). This feeding
influences lichen community composition and competition between
lichen species (Froberg, et.al, 1993). In Oregon it was found
that lichen cover on shrub stems increased when slugs were
excluded (Rosso and McCune, 1997). In the Negev Desert, in
Israel, two species of snails that eat lichens growing under the
surface of limestone rocks ("endolithic lichens") were discovered
to be converting rock to soil at the amazing rate of 0.7 to 1.1
metric tons per hectare per year (Schachak, et. al 1987). The
action was due to the fact that the snails pass significant
amounts of rock through their digestive tracts in the process of
consuming the lichen. In addition, the snails were taking
nitrogen from the lichens and leaving it behind in the new soil;
this was found to be a principal component of the nitrogen cycle
of this desert." (Jones and Shachak, 1990).
Aquatic Fauna
Many types of protozoa have been collected from lichens, but
none are known to be specifically associated with lichens. They
remain on lichens in cyst form while dry, and become active when
wet. Among protozoans, the ciliate genus Colpoda is often
collected in lichens.
Nematodes have been associated with lichens, and can reach
high numbers in them, but what functional connection, if any,
they have to lichens is not known.
Rotifers are common on lichens, and one was found feeding on
a lichen (Pyatt 1968).
Oligochaeta in the family Enchytraeidae have been collected
in large numbers from coastal lichens, and some were raised
successfully on a diet of lichens from the genus Cladonia.
Tardigrades are very common in lichens, and some may require
or prefer lichen habitats. It is not known if Tardigrades feed
on lichens, or just use them as habitat.
MARINE INVERTEBRATES
There are rich lichen floras in the intertidal zone in many
places, and they are accompanied by a correspondingly complex set
of invertebrates. Lichens such as Verrucaria spp. are grazed by
many marine animals including limpets (Patella spp.), periwinkles
(Littorina) and bivalves. Marine invertebrates often use lichens
as shelter; Colman (1940) found up to 16,050 individuals of Laaea
rubra, a bivalve mollusk in 100g wet weight of lichen (Fletcher
1980). Pyrenocollema spp., among other lichens, also live on the
shells of a number of marine organisms, e.g. barnacles.
Barnacles apparently compete with lichens for space (Fletcher
1973); when either organism is well established, the other cannot
easily intrude. A crustacean isopod, Campecopea hirsuta was
found by Weiser (1963) to feed extensively on the lichen Lichina
pygmaea; it was also its preferred habitat.
As with terrestrial invertebrates, marine ones are food for
other organisms; for example, the rock pipit (Anthus spinoletta
spinoletta) (L.) was observed pecking out animals from lichen
covered swards in the supralittoral zone (Fletcher 1973).
BIBLIOGRAPHY
note: This bibliography omits most of the available references
to oribatid mites' association with lichens; a list may be found
in Seyd and Seaward, 1984.
Ali, S.M., M.N. Farooqui and S. Tejpal. 1970. A report of
Laimaphelenchus penardi (Steine, 1914) Fil. Sch. Stek. 1941
(Nematoda: Aphelenchoididae) from India. Marathwada Univ. J.
Sci. 9: 43-44.
Andre, H. 1975. Observations sur les Acariens corticoles de
Belgique. Notes Rech. Fondation Univ. Luxemburgeoise 4: 1-31.
Andre, H. 1979. Notes on the ecology of corticolous epiphyte
dwellers. 1.The mite fauna of fruticose lichens. Recent Advances
in Acarology 1: 551-557.
Andre, H.M. 1985. Associations between corticolous
microarthropod communities and epiphytic cover on bark.
Holarctic Ecology 8: 113-119.
Anonymous. 1970. The Insects of Australia. University Press,
Melbourne.
Aoki, J.-I. 1977. Two new Peloribates species (Acari:
Oribatida) collected from lichens growing on tombstones in
Ichihara-shi, central Japan. Annotationes Zoologicae Japonenses
50: 187-190.
Argue, C.W. 1971. Some terrestrial tardigrades from New
Brunswick, Canada. Can. J. Zool. 49: 401-415.
Bachmann, E. 1929. Tiergallen auf Flechten. Arch. Protistenk.
66: 61-103.
Bailey, R.H. 1970. Animals and the dispersal of soredia from
Lecanora conizaeoides Nyl. ex Cromb. Lichenologist 4: 256.
Bailey, R.H. 1976. Ecological aspects of dispersal and
establishment in lichens. In Lichenology: Progress and Problems.
(D.H. Brown, D.L. Hawksworth and R.H. Bailey, eds.), pp. 215-247.
Academic Press, London and New York.
Baker, E.W. and A.E. Pritchard. 1953. A guide to the spider
mites of cotton. Hilgardia 22: 203-234.
Barkman, J.J. 1958. Phytosociology and Ecology of Cryptogamic
Epiphytes. Van Gorcum: Assen, Netherlands.
Barrows, E.M. 1990. Sex ratio, parasitism and hosts of an
everglades population of Prochalia pygmea (Lepidoptera:
Psychidae). Florida Entomologist 73(4): 672-673.
Baur, A., Baur, B. and L. Froberg. 1992. The effect of lichen
diet on growth rate in the rock-dwelling land snails Chondrina
clienta (Westerlund) and Balea perversa (Linnaeus). J. Moll.
Stud. 58: 345-347.
Beique, R. and A. Francoeur. 1966. Les fourmis d'une pessiere a
Cladonia (Hymenoptera: Formicidae). Nature Can. 93 : 99-106.
Benedetti, R. 1973. Notes on the biology of Neomachilis
halophila on a California sandy beach (Thysanura: Machilidae).
Pan-Pacif. Entomol. 49: 246-249.
Bengston, S.A., A. Fjellberg and T. Solhoy. 1974. Abundance of
tundra arthropods in Spitsbergen. Entomologica Scand. 5: 137-
142.
Bishop, J.A. 1975. Moths, melanism and clean air. Scientific
American 232: 90-99.
Bishop, J.A., L.M. Cook, J. Muggleton and M.R.D. Seaward. 1975.
Moths, lichens and air pollution along a transect from Manchester
to North Wales. J. Applied Ecology 12: 83-98.
Blewitt, M.R. and G.A. Cooper-Driver. 1990. The effects of
lichen extracts on feeding by gypsy moths (Lymantria dispar).
The Bryologist 93(2): 220-221.
Boardman, M., R.R. Askew, and L.M. Cook. 1974. Experiments on
resting site selection by nocturnal moths. J. Zool. Lond. 172:
343-355.
Borrer, D.J. and D.M. DeLong. 1971. An Introduction to the
Study of Insects 3rd ed. Holt, Rinehart and Winston, New York.
Branch, G.M. and M.L. Branch. 1981. Experimental analysis of
intraspecific competition in an intertidal gastropod, Littorina
unifasciata. Aust. J. Marine and Freshwater Res. 32: 573-589.
Brightman, F.H. 1965. Insect on lichens. Lichenologist 3(1):
154.
Broadhead, E. 1958. The psocid fauna of larch trees in northern
England--an ecological study of mixed species populations
exploiting a common resource. J. Anim. Ecol. 27: 217-263.
Broadhead E. and I.W.B. Thornton. 1955. An ecological study of
three closely related psocid specis. Oikos 6: 1-50.
Broadhead, E. and A. Wapshere. 1966. Mesopsocus populations on
larch in England--the distribution and dynamics of two closely-
related coexisting species of Psocoptera sharing the same food
resource. Ecological Monographs 36: 327-388.
Bullock, J.A. 1966. Observations on the fauna of Pulau Tioman
and Pulau Tulai. 9. Introductory report on the terrestrial
arthropods. Bull. Natn. Mus. St. Singapore 34: 104-128.
Byazrov, L.G., E.F. Martyonva and L.N. Medvedev. 1976.
Nogokhvoski (Collembola) v lishainikovykh sinuziyakh Khangaya
(MNR). Byull. Mosk. Obshch. Ispyt. Prir., Otdel Biol. 81: 66-73.
Clarke, C.A. and P.M. Sheppard. 1966. A local survey of the
distribution of the industrial melanic forms in the moth Biston
betularia and estimates of the selective values of these in an
industrial environment. Proc. R. Soc., ser B 165: 424-439.
Coker, P.D. 1967. Damage to lichens by gastropods.
Lichenologist 3(3): 428-429.
Coleman, M.J. and H.B.N. Hynes. 1970. The life histories of
some Plecoptera and Ephemeroptera in a southern Ontario stream.
Can. J. Zool. 48: 1333-1339.
Colloff, M.J. 1983. Oribatid mites associated with marine and
maritime lichens on the island of Great Cumbrae. Glasg. Nat. 20:
347-359.
Colloff, M.J. 1984. Notes on two lichenophagous oribatid mites
from Ailsa Craig (Acari: Cryptostigmata). Glasgow Naturalist 20:
457.
Colman, J.S. 1940. On the faunas inhabiting intertidal
seaweeds. J. Mar. Biol. Assoc. U.K. 24: 129-183.
Cott, H.B. 1957. Adaptive Coloration in Animals Methuen,
London.
Cunningham, D.D. 1907. Plagues and Pleasures of Life in Bengal.
Murray, London.
De Boer, S. 1975. Breeding Nudaria mundana L. (Lep.,
Arctiidae). Ent. Ber., Amst. 35(12): 181-182.
Dev, H.N. 1964. Preliminary studies on the biology of the Assam
thrips, Scirtothrips dorsalis Hood on tea. Indian J. Ent. 26:
184-194.
Emmerich R., I. Giez, O.L. Lange and P. Proksch. 1993. Toxicity
and antifeedant activity of lichen compounds against the
polyphagous herbivorous insect Spodoptera littoralis.
Phytochemistry 33: 1389-1394.
Elton, C.S. 1966. The Pattern of Animal Communities. Methuen &
Co. Ltd, London. (see pps 122, 179, 182-184, 208-209, 355.)
Espoz, C., G. Guzman, and J.C. Castilla. 1995. The lichen
Thelidium litorale on shells of intertidal limpets: a case of
lichen-mediated cryptic mimicry. Marine Ecology Progress Series
119: 191-197.
Essig, E.O. 1958. Insects and Mites of Western North America.
Macmillan, New York.
Evans, G.O., J.G. Sheals, and D. Macfarlane. 1961. The
Terrestrial Acari of the British Isles. Vol. 1. British Museum
(Natural History), London.
Fain, A. 1975. Acariens recoltes par le Dr. J. Trave aux Iles
Subantartiques. I. Familles Saproglyphidae et Hyadesidae.
Acarologia 16: 684-708.
Fletcher, A. 1973. The ecology of marine (littoral) lichens on
some rocky shores of Anglesey. Lichenologist 5: 368-422
Fletcher, A. 1980. Marine and maritime lichens of rocky shores:
their ecology, physiology and biological interactions. Pages
789-842 in The Shore Environment, vol.2: Ecosystems. J.H.
Price, D.E.G. Irvine and W.F. Farnham, eds. The Systematics
Assoc. Special Volume No. 17B, Academic Press.
Forbes, W.T.M. 1960. Lepidoptera of New York and Neighboring
States, Part 4. Cornell Univ. Agric. Exp. Sta. Memoir 371. 188
pp.
Froberg, L., A. Baur, and B. Baur. 1993. Differential herbivore
damage to calcicolous lichens by snails. Lichenologist 25(1):83-
95.
Gadea, E. 1964a. Sobre la nematofauna muscicola y liquenicola
de las islas Pitiusas. Publ. Inst. Biol. Apl. 37: 73-93.
Gadea, E. 1964b. El poblamiento animal liquenicola en pequenos
islotes del Mediterraneo espanol. Bol. Real Soc. Espan. Hist.
Natur., Secc. Biol. 62: 333-336.
Gadea, E. 1974. Nematodes liquenicolas de Columbretes.
Miscelenea Zool. 3(4): 3-8.
Garrett, R.M. 1974. A species of Chrysopa of special interest
to lichenologists. Revue Bryol. Lichen. 40: 283-286.
Gerson, U. 1968. Five tydeid mites from Israel (Acarina:
Prostigmata). Israel J. Zool. 17: 191-198.
Gerson, U. 1971. The mites associated with citrus in Israel.
Israel J. Ent. 6: 5-22.
Gerson, U. 1973. Lichen-arthropod associations. Lichenologist
5: 434-443.
Gerson U. and M.R.D. Seaward. 1977. Lichen-invertebrate
associations. Pages 69-119 in M.R.D. Seaward, ed., Lichen
Ecology. Academic Press, London.
Giez, I., O.L. Lange and P. Proksch. 1994. Growth retarding
activity of lichen subtances against the polyphagous herbivorous
insect Spodoptera littoralis. Biochem. Syst. Ecol. 22: 113-120.
Gilbert, O.L. 1971. Some indirect effects of air pollution on
bark-living invertebrates. J. Appl. Ecol. 8: 77-84.
Gilbert, O.L. 1976. A lichen-arthropod community.
Lichenologist 8: 96-97.
Gjelstrup, P. and U. Sochting. 1979. Cryptostigmatid mites
(Acarina) associated with Ramalina siliquosa (Lichenes) on
Bornholm in the Baltic. Pedobiologia 19: 237-245.
Graham, A. 1955. Molluscan diets. Proc. Malac. Soc. Lond. 31:
144-159.
Grandjean, F. 1950. Observations ethologiques sur Camisia
segnis (Herm.) et Platynothris peltifer (Koch) (Acariens). Bull.
Mus. Hist. Nat. Paris. 224-231.
Green, J. 1951. The feeding habits of Cylindronotus
laevioctostriatus (Goeze) (Col., Tenebrionidae) and its larva.
Ent. Mon. Mag. 87: 19.
Gressitt, J.L. 1965. Flora and fauna on backs of large Papuan
moss-forest weevils. Science 150: 1833-1835.
Gressitt, J.L. 1966. Epizoic symbiosis: the Papuan weevil genus
Gymnopholus (Leptopiinae) symbiotic with cryptogamic plants,
oribatid mites, rotifers and nematodes. Pacif. Insects 8: 221-
280.
Gressitt. J.L. 1967b. Notes on arthropod populations in the
Antarctic Peninsula South Shetland Islands -South Orkney Islands
area. In Entomology in Antarctica (J.L. Gressitt, ed.) pp. 307-
320. Washington, D.C.
Gressitt, J.L. 1970. Papuan weevil genus Gymnopholus: second
supplement with studies in epizoic symbiois. Pacif. Insects 12:
753-762.
Gressitt, J.L. 1977. Symbiosis runs wild on the backs of high-
living weevils. Smithsonian 7(11): 135-140.
Gressitt, J.L. and J. Shoup. 1967a. Ecological notes on free-
living mites in North Victoria Land. In J.L. Gressitt, ed.
Entomology of Antarctica. American Geophysical Union, Washington.
pp. 307-320.
Grummann, V.J. 1960. Die Cecidien auf Lichenen. Bot. Jb. 80:
101-144.
Hale, M.E. 1972. Natural history of Plummers Island, Maryland.
XXI. Infestation of the lichen Parmelia baltimorensis Gyel. &
For. by Hypgastrura packardii Folsom (Collembola). Proc. Biol.
Soc. Washington 85: 287-296.
Hale, W.G. 1963. The Collembola of eroding blanket bog. In
Soil Organisms (J. Doeksen and J. van der Drift, eds), pp. 406-
413. North-Holland, Amsterdam.
Harding, D.J.L. and R.A. Stuttard. 1974. Microarthropods. In
Biology of Plant Litter Decomposition (C.H. Dickinson and G.J.F.
Pugh, eds.), Vol.II, pp. 489-532. Academic Press, London and New
York.
Hawksworth, D.L. 1991. Linnaeus, the first report of
lichenophagy in psychid moths, and the identity of Lichen
candelarius. The Lichenologist 23:92.
Healy, I.N. 1967. The population metabolism of Onychiurus
procampatus Gisin (Collembola). In Progress in Soil Zoology
(O.Graff and J.E. Satchell, eds) pp. 127-134. North-Holland,
Amsterdam.
Henderson, A. and D.J. Hackett. 1986. Lichen and algal
camouflage and dispersal in the psocid nymph Trichadenotecnum
fasciatum. Lichenologist 18(2): 199-200.
Hesbacher, S., B. Baur, A. Bauer and P. Proksch. 1995.
Sequestration of lichen compounds by three species of terrestrial
snails. J. Chem. Ecol. 21(2): 233-246.
Hesbacher, S., I. Giez, G. Embacher, K. Fiedler, W. Max, A.
Trawoger, R. Turk, O.L. Lange and P. Proksch. 1995.
Sequestration of lichen compounds by lichen-feeding members of
the Arctiidae (Lepidoptera). J. Chem. Ecol. 21(12): 2079-2089.
Holden, M. and M.V. Tracey. 1950. A survey of enzymes that can
break down tobacco-leaf components. 2. Digestive juice of Helix
on defined substances. Biochem. J. 47: 407-414.
Holland, W.J. 1968. The Moth Book. Dover Publications, Inc.,
New York.
Holloway, B.A. 1970. A new genus of New Zealand Anthribidae
associated with lichens (Insecta: Coleoptera). N. Z. J. Sci. 13:
435-446.
Hughes, A.M. and P.J. Tillbrook. 1966. A new species of
Calvolia (Acaridae, Acarina) from the South Sandwich Islands.
Bull. Br. Antarct. Surv. 10: 45-53
James, P.W. and A. Henssen. 1976. The morphological and
taxonomic significance of cephalodia. In Lichenology: Progress
and Problems (D.H. Brown, D.L. Hawksworth and R.H. Bailey, eds.)
pp. 27-77. Academic Press, London.
Jones, C.G. and M. Schachak. 1990. Fertilization of the desert
soil by rock-eating snails. Nature 346: 839-841.
Kalshoven, L.G.E. 1958. Observations on the black termites,
Hospitalitermes sp., of Java and Sumatra. Insectes Soc. 5: 9-30.
Kettlewell, H.B.D. 1956. Further selection experiments on
industrial melanism in the Lepidoptera. Heredity 10: 287-301.
Kettlewell, H.B.D. 1959. Brazilian insect adaptations.
Endeavour 18: 200-210.
Kettlewell, H.B.D. 1961. The phenomenon of industrial melanism
in Lepidoptera. A. Rev. Ent. 6: 245-262.
Kettlewell, H.B.D. 1973. The Evolution of Melanism. Oxford
University Press, Oxford.
Kimmell, R.G. and P.A. Meglitsch. 1969. Notes on Iowa
tardigrades. Proc. Iowa Acad. Sci. 76: 454-462.
Kim, K.C. 1993. Biodiversity, conservation and inventory: why
insects matter. Biodiversity and Conservation 2: 191-214.
Kohlmeyer, J. and B. Volkmann-Kohlmeyer. 1988. Halographis
(Opegraphales), a new endolithic lichenoid from corals and
snails. Canad. J. bot. 66: 1138-1141.
Kuhnelt, W. 1976. Soil Biology with Special Reference to the
Animal Kingdom. 2nd. Ed. Faber. London.
Lagerspetz, K. 1953. Biocoenological notes on the Parmelia
saxatilis -Dactylochelifer latreillei community of seashore
rocks. Arch. Soc. Zool. Bot. Fenn. Vanamo 7: 131-142.
Lamb, I.M. The occurrence of a rotifer on the surfaces of
apothecia of Xanthoria parietina. 1968. Lichenologist 4(1): 74-
75.
Laminger, H. 1971. Uber das Vorkommen von Schalenamoeben
(Rhizopoda, Testacea) in alpinen Flechten. Zool. Anz. 186: 335-
337.
Laundon, J.R. 1967. A study of the lichen flora of London.
Lichenologist 3: 277-327.
Laundon, J.R. 1971. Lichen communities destroyed by psocids.
Lichenologist 5: 177.
Lawrey, J.D. 1980a. Calcium accumulation by lichens and
transfer to lichen herbivores. Mycologia 72: 586-594.
Lawrey, J.D. 1980b. Correlations between lichen secondary
chemistry and grazing activity by Pallifera varia. The
Bryologist 83(3): 328-334.
Lawrey, J.D. 1983a. Lichen herbivore preference: a test of two
hypotheses. Am. J. Bot. 70: 1188-1194.
Lawrey, J.D. 1983b. Vulpinic and pinastric acids as lichen
antiherbivore compounds: contrary evidence. Bryologist 86: 365-
369.
Lawrey, J.D. 1984. Role of lichen compounds in herbivore
defence. Pages 232-247 in Biology of Lichenized Fungi by J.D.
Lawrey, Praeger Publishers.
Lawrey, J.D. 1986. Biological role of lichen substances.
Bryologist 89(2): 111-122.
Lawrey, J.D. 1987. Nutritional ecology of lichen/moss
arthropods. Pages 209-233 in Nutritional Ecology of Insects,
Mites, Spiders, and Related Invertebrates. F. Slansky and J.G.
Rodriquez, eds., John Wiley & Sons.
Lawrey, J.D. 1989. Lichen secondary compounds: Evidence for a
correspondence between the antiherbivore and anti-microbial
function. Bryologist 92: 326-369.
Leinaas, H.P. and L. Somme. 1983. Adaptations in Xenyella
maritima and Anurophorus laricis to lichen habitats of alpine
rocks. Oikos 43: 197-206.
Leinaas, H.P. and A. Fjellberg. 1985. Habitat structure and
life history strategies of two partly sympatric and closely
related lichen feeding Collembola species. Oikos 44: 448-458.
Lindsay, D.C. 1978. The role of lichens in Antarctic
ecosystems. Bryologist 81: 268-276.
Llano, G.A. 1956. Utilization of lichens in the Arctic and
Subarctic. Econ. Bot. 10: 367-392.
Luxton, M. A re-description of Cryptognathus lagena Kramer 1879
(Acari: Prostigmata: Cryptognathidae). Acarologia 14: 591-594.
MacNeill, N. 1966. Mites (Acari) on lichen. Ir. Nat. J. 15:
242-243.
Madssen, H. 1940. A study of the littoral fauna of northwest
Greenland. Meddr. Gronland. 124: 1-24.
Malcolm, W.M. and J.P. Hanks. 1973. Landing-site selection and
searching behaviour in the micro-lepidopteran Agonopteryx
pulvipenella. Anim. Behav. 21: 45-48.
Mani, M.S. 1962. Introduction to High Altitude Entomolgy.
Methuen, London.
Mani, M.S. 1964. The Ecology of Plant Galls. W. Junk, The
Hague.
Marcus, E. 1959. Tardigrada. In Fresh-water biology. (W.T.
Edmonson, ed.) pp. 508-521. Wiley, New York.
Matuura, A., Y. Yamamoto and I. Hattori. 1973. Early Stages of
Japanese Moths. Vol. 1. Hoikusha Publishing Co., Ltd., Osaka,
Japan. 238 pp.
Maynard, E.A. 1951. A Monograph on the Collembola or Springtail
Insects of New York State. Ithaca.
McCarthy P.M. and J.A. Healy. 1978. Dispersal of lichen
propagules by slugs. Lichenologist 10: 131-134.
McDonogh, R.S. 1939. The habitat, distribution and dispersal of
the psycid moth, Luffia ferchaultella, in England and Wales. J.
Anim. Ecol. 8: 10-28.
Meininger, C.A. 1981. The influence of urban climate on the
tardigrade species inhabiting mosses and lichens. M.S. Thesis,
University of Cincinnati, Cincinnati, Ohio. 109 pp.
Meininger, C.A., G.W. Uetz, and J.A. Snider. 1985. Variation in
epiphytic microcommunities (tardigrade-lichen-bryophyte
assemblages) of the Cincinnati, Ohio area. Urban Ecology 9: 45-
61.
Michael, A.D. 1883-1887. British Oribatidae. Ray Society,
London.
Morton, J.E. 1954. The crevice faunas of the upper intertidal
zone at Wembury. J. Mar. Biol. Ass. U.K. 33: 187-224.
Morton, J.E., A.D. Boney, and E.D.S. Corner. 1957. The
adaptations of Lasaea rubra (Montagu), a small intertidal
lamellibranch. J. Mar. Biol. Ass. U.K. 36: 383-405.
Nielson, C.O. 1963. Laminarinases in soil and litter
invertebrates. Nature 199: 1151-1161.
Nielson, C.O. Nematoda. In Soil Biology (A. Burges and F. Raw,
eds.) pp. 197-211. Academic Press, London and New York.
Nosek, J. 1973. The European Protura, their Taxonomy, Ecology
and Distribution, with Keys for Determination. Museum d'Histoire
Naturelle, Geneve.
Owen, G. 1966. Feeding. In Physiology of Mullusca (K.M.
wilbur and C.M. Younge, eds.), Vol. II, pp. 1-51. Academic
Press, New York and London.
Paulson, R. and P.G. Thompson. 1913. Report on the lichens of
Epping Eorest (second paper). Essex Naturalist 17: 90-104.
Peake, J.F. and P.W. James. Lichens and mollusca. 1967. The
Lichenologist 3(3):425-427.
Pettersson, R.B., J.P. Ball, K.-E. Renhorn, P.-A. Esseen and K.
Sjoberg. 1995. Invertebrate communities in boreal forest
canopies as influenced by forestry and lichens with implications
for passerine birds. Biological Conservation 74: 57-63.
Plitt, C.C. 1934. A lichen-eating snail. Bryologist 37: 102-
104.
Prinzing, A. 1997. Spatial and temporal use of microhabitats as a key strategy for the colonization of tree bark by Entomobrya nivalis L. (Collembola: Entomobryidae), pp. 453-476 in N.E. Stork, J. Adis, and R.K. Didham (eds.), Canopy Arthropods. Chapman & Hall, London.
Prinzing, A. and H.P. Wirtz. 1997. The
epiphytic lichen, Evernia prunastri L., as a habitat for arthropods: shelter
from desiccation, food-limitation and indirect mutualism. pp. 477-494 in
N.E. Stork, J. Adis, and R.K. Didham (eds.), Canopy Arthropods. Chapman
and Hall, London.
Pyatt, F.B. 1968. The occurrence of a rotifer on the surfaces
of apothecia of Xanthoria parietina. Lichenologist 4: 74-75.
Reutimann, P. and C. Scheidegger. 1987. Importance of lichen
secondary products in food choice of two oribatid mites (Acari)
in an alpine meadow ecosystem. J. Chem. Ecol. 13: 363-370.
Richardson, D.H.S. 1974. The Vanishing Lichens. Hafner Press
(Division of Macmillan Pub. Co., New York).
Ridley, H.N. 1930. The Dispersal of Plants Throughout the
World. Reeve and Co., Ashford, England.
Roberts, D. and D. Zimmer. 1990. Microfaunal communities
associated with epiphytic lichens in Belfast. Lichenologist
22(2): 163-171.
Ross, E.S. 1966. A new species of Embioptera from the Galapagos
Islands. Proc. Calif. Acad. Sci. 4th series, vol. XXXIV (12):
499-504.
Rosso, A.L. and B. McCune. 1997. Herbivory by mollusks--should
an epiphyte ecologist care? Northwest Science Abstracts, March,
1997.
Rundel, P.W. 1978. The ecological role of secondary lichen
substances. Biochemical Systematics and Ecology. 6: 157-170.
Schmid, G. 1929. Endolithische Kalkflechten und Schneckenfrass.
Biologisches Zentralblatt 49: 28-35.
Schuster, R.O. and A.A. Grigarick. 1966. Tardigrada from the
Galapagos and Cocos Islands. Proc. Calif. Acad. Sci. 34: 315-
328.
Seaward, M.R.D. 1965. Lincolnshire psocids. Trans.
Lincolnshire Nat. Union. 16: 99-100.
Seaward, M.R.D. 1974. A note on Phauloppia lucorum C.L. Koch
(Acari: Oribatei) and lichens. Lichenologist 6: 126-127.
Seaward, M.R.D. 1984. Contribution of lichens to ecosystems.
Pages 107-129 in CRD Handbook of Lichenology, Vol.II. M. Galun,
ed. CRC Press, Inc., Boca Raton, Florida.
Seguy, E. La Biologie des Dipteres. Lechevalier, Paris.
Seyd, E.L. and M.R.D. Seaward. 1984. The association of
oribatid mides with lichens. Zoological Journal of the Linnean
Society 80: 369-420.
Shachak, M., C.G. Jones and Y. Granot. Herbivory in rocks and
the weathering of a desert. Science 236: 1098-1099.
Shorrocks, B., J. Marsters, I. Ward, and P.J. Evennett. 1991.
The fractal dimension of lichens and the distribution of
arthropod body lengths. Functional Ecology 5: 547-460.
Siddiqi, M.R. and D.L. Hawksworth. 1982. Nematodes associated
with galls on Cladonia glauca, including two new species.
Lichenologist 14(2): 175-184.
Sigal, L.L. 1984. Of lichens and lepidopterons. The Bryologist
87(1): 66-68.
Skorepa, A.C. and A.J. Sharp. Lichens in "packets" of lacewing
larvae (Chrysopidae). Bryologist 74(3): 363-364.
Slansky, F. Jr. 1979. Effect of the lichen chemicals atranorin
and vulpinic acid upon feeding and growth of larvae of the
yellow-striped armyworm, Spodoptera ornithogalli. Environmental
Entomology 8(5): 865-868.
Slocum, R.D. and J.D. Lawrey. 1976. Viability of the epizoic
lichen flora carried and dispersed by green lacewing (Nodita
pavida) larvae. Can. J. Bot. 54: 1827-1831.
Smith, A.L. 1921. Lichens. Cambridge University Press,
Cambridge, England.
Sochting, U. and P. Gjelstrup. 1985. Lichen communities and the
associated fauna on a rocky shore on Bornholm in the Baltic.
Holarctic Ecology 8: 66-75.
Southwood, T.R.E. and D. Leston. 1959. Land and Water Bugs of
the British Isles. Warne, London.
Sowter, F.A. 1971. Mites (Acari) and lichens. Lichenologist 5:
176.
Stehr, F.W., ed. 1991. p. 136 in Immature Insects Vol. 2.
Kendall Hunt, Pubs. (reference to lichen-camouflaged lacewing
larvae.)
Stebaev, I.V. 1963. Die Veranderung der Tierbevolkerung der
Boden im Laufe der Bodenentwicklung auf Felsen und auf
Verwitterungsprodukten im Wald-Wiesenlandschaften des Sud-Urals.
Pedobiologia 2: 265-309. (in Russian)
Stephenson, N.L. and P.W. Rundel. 1979. Quantitative variation
and the ecological role of vulpinic acid and atranorin in the
thallus of Letharia vulpina (Lichenes). Biochem. Syst. Ecol. 7:
263-267.
Stephenson, T.A. and A. Stephenson. 1972. Life Between
Tidemarks on Rocky Shores. Freeman, San Francisco.
Stout, J.D. and D.W. Heal. 1967. Protozoa. In Soil Biology (A.
Burges, and F. Raw, eds.), pp.149-195. Academic Press, New York
and London.
Strong, J. 1967. Ecology of terrestrial arthropods at Palmer
Station, Antarctic Peninsula. In J.L. Gressitt, ed. Entomology
of Antartica. American Geophysical Union, Washington. pp. 357-
371.
Stubbs, C.S. 1987. Corticolous lichen ecology: distribution
patterns and lichen-invertebrate interactions. M.S. Thesis,
University of Maine, Orono.
Stubbs, C.S. 1989. Patterns of distribution and abundance of
corticolous lichens and their invertebrate associates on Quercus
rubra in Maine. Bryologist 92(4): 453-460.
Stubbs, C.S. 1992. Lichen dispersal by invertebrates. Ph.D.
Dissertation. University of Maine, Orona.
Stubbs, C.S. 1995. Dispersal of soredia by the oribatid mite,
Humerobates arborea. Mycologia 87(4): 454-458.
Thom, R.H. and G. Iffrig. 1985. p. 69 in Directory of Misouri
Natural Areas. Pub. by Missouri Natural Areas Committee (MO
Dept. of Conservation and Dept. of Nat. Res. (reference to
grasshopers camouflaged by lichen-covered rocks.)
Thompson, J.C. 1960. Ciliated protozoa from tree bourne mosses
and lichens. J. Protozool. 7(supp.): 17.
Tillbrook, P.J. 1967. The terrestrial invertebrate fauna of the
maritime Antarctic. Phil. Trans. R. Soc. B. 252: 261-278.
Trave, J. 1963. Ecologie et biologie des Oribates (Acariens)
saxicoles et arboricoles. Vie Milieu (Suppl.) 14: 1-267.
Trave, J. 1969. Sur le peuplement des lichens crustaces des
Iles Salvages par les Oribates (Acariens). Rev. Ecol. Biol. Sol.
6: 239-248.
Turner, B.D. and E. Broadhead. 1974. The diversity and
distribution of psocid populations on Mangifera indica L. in
Jamaica and their relationship to altitude and micro-epiphytic
diversity. J. Anim. Ecol. 43: 173-190.
Uetz, G.W., C.A. Meininger, and J. Snider. 1980. Effect of air
pollution on micro-communities of invertebrates in lichens and
mosses. Rep. RP 1224-2 submitted to Electric Power Research
Institute, Palo Alto, California. 130 pp.
Watson, G., J.J.Davis, and W.C. Hanson. 1966. Terrestrial
Invertebrates. In Environment of the Cape Thompson Region,
Alaska (N.J. Wilimovsky and J.N. Wolf, eds.), pp. 565-584. U.S.
Atomic Energy Commission.
Weber, W.A. 1974. Two lichen-arthropod associations in
Australia and New Guinea. Lichenologist 6: 168.
Wessels, D.C.J. and L.A. Wessels. 1991. Erosion of biogenically
weathered Clarens sandstone by lichenophagous bagworm larvae
(Lepidoptera; pyschidae). Lichenologist 23(3): 283-291.
Wessels, D.C.J., L.A. Wessels, and W.H. Holzapfel. 1979.
Preliminary report on lichen-feeding Coleoptera occurring on
Teloschistes capensis in the Namib Desert, South West Africa.
Bryologist 82: 270-273.
Wielgolaski, F.E. 1975. Functioning of Fennoscandian tundra
ecosystems. In Fennoscandian Tundra Ecosystems Part. 2: Animals
and Systems Analysis (F.E. Wielgolaski, ed.), pp. 300-326.
Springer Verlag, Berlin.
Wieser, W. 1963. Adaptations of two intertidal isopods. II.
Comparison between Campecopea hirsuta and Naesa bidentata
(Sphaeromatidae). J. Mar. Biol. Assoc. U.K. 43: 97-112.
Woodring, J.P. 1963. The nutrition and biology of saprophytic
Sarcoptiformes. Adv. Acarol. 1: 89-111.
Woodring, J.P. and E.F. Cook. 1962. The biology of Ceratozetes
cisalpinus, Scheloribates laevigatus, and Oppia neerlandica
(Oribatei) with a description of all stages. Acarologia 4: 101-
137.
Yom-Tov, Y. and M. Galun. 1971. Note of feeding habits of the
desert snails Sphincterochila boissieri Charpentier and
Trochoidea (Xerocrassa) seetzeni Charpentier. Veliger 14: 86-88.
Young, S.R. and W. Block. 1980. Some factors affecting
metabolic rate in an Antarctic mite. Oikos 34: 178-185.
Zagulyayev, A.K. 1970. Two new primitive species of
lichenophagous moths (Lepidoptera, Tineidae) from the wet forests
of Azerbaidzhan. Entomological Review. 49(3): 408-411.
Zopf, W. 1896. Zur biologischen Bedeutung der Flechtensauren.
Biol. Centralbl. 16: 593-610.
Zopf, W. 1907. Biologische und morphologische Beobachtungen an
Flechten. III. Durch tierische Eingriffe hervorgerufene
Gallenbildungen an Vertreten der Gattung Ramalina. Ber. Dsch.
Bot. Ges. 25: 233-237.
Zukal, H. 1895. Morphologische und biologische Untersuchungen
uber die Flechten. Acad. Wiss. Math.-Naturw. Cl. Wien 104(1):
1303-1395.
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