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LICHENS AND INVERTEBRATES: A BRIEF REVIEW AND BIBLIOGRAPHY

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.


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