Volume 2 Chapter 4-8 - Invertebrates - Molluscs PDF

Glime, J. M. 2017. Invertebrates: Molluscs. Chapt. 4-8. In: Glime, J. M. Bryophyte Ecology. Volume 2. Bryological Interaction.
4-8-1
Ebook sponsored by Michigan Technological University and the International Association of Bryologists. Last updated 26 September 2017 and
available at <http://digitalcommons.mtu.edu/bryophyte-ecology2/>.
CHAPTER 4-8
INVERTEBRATES: MOLLUSCS
TABLE OF CONTENTS
Gastropoda: Snails and Slugs............................................................................................................................. 4-8-2

Reproduction................................................................................................................................................ 4-8-3
Mating and the Love Dart ..................................................................................................................... 4-8-4
Egg and Larval Development ............................................................................................................... 4-8-5
Bryophyte Interactions................................................................................................................................. 4-8-6
Abundance ................................................................................................................................................... 4-8-6
Adaptations .................................................................................................................................................. 4-8-7
Confusing the Predator ......................................................................................................................... 4-8-7
Jumping to Escape ................................................................................................................................ 4-8-7
Keeping It Small ................................................................................................................................... 4-8-8
Conical Shape ....................................................................................................................................... 4-8-9
Avoiding Desiccation............................................................................................................................ 4-8-9
No Shell – Slugs ................................................................................................................................. 4-8-10
In Search of Food....................................................................................................................................... 4-8-11
Low Palatability? ................................................................................................................................ 4-8-12
Low Nutritional Quality? .................................................................................................................... 4-8-14
Food for Some .................................................................................................................................... 4-8-14
An Avoidance of Gametophores? ....................................................................................................... 4-8-16
Deterrents to Herbivory ...................................................................................................................... 4-8-20
Digestibility ........................................................................................................................................ 4-8-21
Role in Bryophyte Competition with Lichens .................................................................................... 4-8-21
Palatable Gametophytes...................................................................................................................... 4-8-21
Aquatic Grazing.................................................................................................................................. 4-8-21
Bryophyte Antifeedants ............................................................................................................................. 4-8-22
Dispersal Agents ........................................................................................................................................ 4-8-23
Bryophytes as Home .................................................................................................................................. 4-8-28
Epiphytic............................................................................................................................................. 4-8-32
Calcareous Areas ................................................................................................................................ 4-8-33
Bogs and Mires ................................................................................................................................... 4-8-35
Aquatic................................................................................................................................................ 4-8-37
Plant Protectors .......................................................................................................................................... 4-8-38
Mussels (Bivalve Molluscs) .............................................................................................................................. 4-8-38
ECHINODERMATA........................................................................................................................................ 4-8-38
Summary ........................................................................................................................................................... 4-8-39
Acknowledgments............................................................................................................................................. 4-8-39
Literature Cited ................................................................................................................................................. 4-8-39
4-8-2 Chapter 4-8: Invertebrates: Molluscs
CHAPTER 4-8
INVERTEBRATES: MOLLUSCS
Figure 1. Slug on a Fissidens species. Is it eating, or just a casual visitor? Photo by Janice Glime.
The most familiar of the bryophyte inhabitants among of terrestrial taxa restricted to limestone areas. Slugs
the molluscs are the snails and slugs, but you will see that (Figure 3), on the other hand, lack shells and exhibit no
some bivalves also have an interesting relationship with external twists. Instead they have a thin calcareous plate
bryophytes. embedded in the mantle.
Mollusca are considered to be bilaterally symmetrical Unlike the marine snails, terrestrial gastropods lack an
(like humans) (Pratt 1935), but they seem to push the operculum to cover the shell opening. Instead, they use a
definition to the limit. In bivalves, that is not too difficult calcified slime (epiphragm; Figure 4) for protection in
to understand, but in snails the twisted body and shell seem hibernation or aestivation. The respiratory pore (Figure
to twist the definition as well; even organs normally paired, 3) is on the right side of the body, and closes to keep out
like kidneys, are not paired (Figure 2). water in aquatic species or to prevent desiccation under dry
conditions on land. Both aquatic and terrestrial gastropods
Gastropoda: Snails and Slugs have lungs, necessitating return to the surface for aquatic
Most terrestrial and freshwater snails (Pulmonata) members to get air. Aquatic members have only one pair
have spiral shells and these may be taller than the diameter of non-retractile tentacles, whereas land-dwellers have two
of the opening (elongate/conical; Figure 26) or shorter pairs and both are retractile. Aquatic species have an eye at
(Figure 145) (Pratt 1935). The inside body is also a spiral, the base of each tentacle; the land snails have their eyes on
but it is not the same spiral as the one of the shell. This the tips of the rear pair of tentacles.
internal spiral affects the digestive system as well. With its Most gastropods eat algae and plants, which they
mouth to the ground, the snail is infamous for the scrape with the radula (Figure 5), but a few are
positioning of the anus above the mouth on the right side of carnivorous. The radula is made of chitin with rows of
the head (Figure 2). minute calcareous teeth. And if you thought bryophytes
In snails, the mantle secretes a shell, and this requires used minute characters for identification, snail
calcium carbonate. For this reason, you will find a number identification is often based on these teeth!
Chapter 4-8: Invertebrates: Molluscs 4-8-3
Figure 2. Snail, showing its major internal and external parts. Note the dart sac from which the love dart is ejected. Image from
Wikimedia Creative Commons.
Figure 3. Great Red Slug, Arion rufus, dark form, Bishop

Middleham Quarry Nature Reserve, Co Durham. Note the large
respiratory pore on the mantle of this sometimes moss dweller. Figure 5. Pomacea canaliculata mouth showing radula.
This snail can travel nearly 0.5 km in search of more suitable Photo by S. Ghesquiere, through Wikimedia Commons.
conditions (Sandelin 2012). Photo by Brian Eversham, with
permission. Reproduction
Most terrestrial snails and slugs are simultaneous
hermaphrodites, mutually exchanging gametes during
copulation. This is not true for land-dwelling prosobranch
snails (including the Pomatiidae, Aciculidae,
Cyclophoridae, and others) – families that have separate
sexes (Wikipedia 2012b). The prosobranch snails are the
ones that have an operculum that can be used to cover the
opening when they retreat into the shells.
Some land snails are sequential hermaphrodites,
being first male, then female (Nordsieck 2012b). Others,
such as Arianta arbustorum (Helicidae; Figure 6), a moss-
dwelling snail, have a mechanism that prevents sperm cells
from fertilizing the snail's own egg cells before they reach
Figure 4. Helix pomatia epiphragm. Photo by Hannes the sperm pouch of the mate. In the aquatic Lymneidae,
Grobe, through Wikimedia Commons. snails can reproduce using unfertilized eggs, permitting
them to multiply rapidly in a new location and causing

invasive species problems when they are introduced as
aquarium pets.
The reproductive anatomy of the snail is a bit peculiar,
with the penis and vagina everting from near the head
(Figure 7-Figure 8). In the hermaphrodites, the penes wrap
around each other, sometimes extending to great lengths
(Figure 9).
Figure 9. Slugs mating, demonstrating the very long penes.

Photo through Wikimedia Commons.
Mating and the Love Dart

The mating process is a combination of love and war
(Figure 10). The dart, or more than one in some species, is
Figure 6. Arianta arbustorum on a bed of mosses and leafy made of calcium carbonate, chiton, or cartilage (Figure 11).
liverworts. Photo ©Roy Anderson, with permission. During mating, each snail tries to inject this "dart" into the
other snail (Figure 12) (Koene & Chase 1998a; Chase &
Blanchard 2006). It might be more appropriate to call this
a dagger because it is injected by a thrust, not a shot or a
throw. The first mating of a snail stimulates the production
of the dart, so it cannot be used until the second mating.
Once used, it requires time to generate a new one.
Figure 7. Helix pomatia head during mating. Redrawn from

Johannes Meisenheimer, through Wikimedia Commons.
Figure 10. Roman snails (Helix pomatia) in full foot contact

during mating. This process of contact of foot, lips, and tentacles
can take up to 20 hours. Photo through Wikimedia Commons.
Figure 8. Helix pomatia head after mating, showing both

male and female parts of this simultaneous hermaphrodite. Figure 11. Love dart of the snail Monachoides vicinus.
Redrawn from Johannes Meisenheimer, through Wikimedia Photo by Joris M. Koene and Hinrich Schulenburg through
Commons. Wikimedia Commons.
Egg and Larval Development

Most gastropods lay eggs, with only a few species
bearing live young. In aquatic snails, development of the
larva occurs as a planktonic stage once it leaves the egg,
but in terrestrial pulmonate snails, development is
completed within the egg. Some snails (e.g. Clausiliidae)
exhibit ovoviviparity, wherein the larvae emerge inside the
mother's body and emerge from "her" body as juvenile
snails (Nordsieck 2012b). This practice permits these
snails to live in dry areas where external eggs could not
survive the desiccation. Some species of the oviparous
(egg-laying) species, such Arion flagellus (Figure 13), lay
their eggs under or among bryophytes (Figure 14).
Figure 12. Mating garden snails (Helix aspera) with love

dart in snail on right, just above the antenna of the left snail.
Photo by Eynar through Wikimedia Commons.
Figure 13. Arion flagellus on a sheet of mosses. Photo ©
But what does the dart accomplish? Early hypotheses Roy Anderson <habitas.org.uk>, with permission.
considered it to be a "gift of calcium" to help in the
development of the eggs. Leonard (1992) used a
theoretical model to support the hypothesis that the love
dart induced the partner to act as a male, hence insuring
that the thruster would also be fertilized. Koene and Chase
(1998a, b) used an experimental approach to disprove the
long-held hypothesis of a "gift of calcium."
Through the work of Koene and Chase (1998 a, b), the
role of this dart has become clearer. It carries with it a mix
of hormones that help to move the sperm cells toward the
sperm pouch where they are stored until fertilization
(Koene & Chase 1998a, b). This is accomplished by
causing changes in the structure of the copulatory canal
leading ultimately to the sperm pouch. These changes
increase the chances, often doubling them, that sperm from
that mating snail are successful in fertilizing eggs, since it
Figure 14. Arion flagellus eggs in Oaks Wood, Cambourne,
is likely that the partner will have multiple mating events. Cambridgeshire, UK. Note the bit of moss beside the eggs and on
But the dart, preferably aimed at the foot, can miss its ideal the eggs – remnants of the cover that previously protected them.
target and land in a less desirable location, like the base of Photo by Brian Eversham, with permission.
the antenna. When that happens, the snail is no longer able
to retract or extend the antenna. Richter (1972) found that the banana slug (Ariolimax
Each partner goes through gyrations apparently in an columbianus, Figure 15) laid 3-4 mm eggs under moss
attempt to avoid being recipient of the love dart, or at least where soil conditions were neither excessively wet nor dry.
to avoid receiving it in an undesirable location. So far, Placing eggs under mosses and other loose substrata may
be an energy-saving strategy for some species. Bauer
Leonard's (1992) hypothesis of stimulating the partner to
(1994) considered the behavior of some snails that dig
carry out its male role does not seem to have been tested holes to be an investment in parental care, but incurring
experimentally, but with the mix of hormones it could still an energy cost. Other than these preparations, snails do not
be a viable part of the story. It appears that this love dart, tend their eggs or hatchlings. Ariolimax californicus
although not understood at the time, could have been the (Figure 16) also may occur under bryophytes (Peggy
basis for the story regarding Cupid's arrow (Chase 2010). Edwards, pers. comm.).
Figure 15. Ariolimax columbianus on moss. Photo by Bill

Leonard, with permission.
Figure 18. Eremarionta immaculata in the Riverside

Mountains, CA, USA. Photo by William D. Wiesenborn, with
permission.
But not all snails and slugs find the bryophyte

substrate attractive. Some actually avoid its rough surface.
Nevertheless, trails of slime (Figure 19) are not unusual,
and we have little insight into the reasons why some find it
inviting while others find it repulsive.
Figure 16. Ariolimax californicus exiting a mat of mosses, a

suitable location for laying eggs. Photo coourtesy of Peggy
Edwards.
Bryophyte Interactions
Glistening trails of pearly mucous (Figure 17) criss-
cross mats and turfs of green, signalling the passing of
snails and slugs on the low-growing bryophytes (Figure 1).
In California, the white desert snail Eremarionta
immaculata (Figure 18) is more common on lichens and
mosses than on other plant detritus and rocks (Wiesenborn
2003). Wiesenborn suggested that the snails might find Figure 19. Snail or slug trails on Dicranum viride on big
more food and moisture there. Are these molluscs simply maple trunk. Photo courtesy of Betsy St. Pierre.
travelling from one place to another across the moist moss
surface, or do they have a more dastardly purpose (as Abundance
hunters) for traversing these miniature forests?
Snails can sometimes occur in significant numbers in
moss habitats. Their need for a moist environment (Pratt
1935) would seemingly attract snails to the mosses as a
moist substrate. Quantitative information on snails and
slugs among bryophytes is scarce, and often only mentions
that bryophytes are abundant in the habitat (e.g. Nekola
2002).
The study by Grime and Blythe (1969) is helpful in
understanding numbers and dynamics of moss-dwelling
snail populations, but we need many more studies. They
found average morning populations of up to 8.5 per 100 g
dry weight of moss in early September for the copse snail
Arianta arbustorum (Figure 20) at Winnats Pass in
Derbyshire, England. In collections totalling 82.4 g of
moss, they examined snail populations in a 0.75 m2 plot
each morning on 7, 8, 9, & 12 September 1966. Arianta
Figure 17. Lehmannia valentiana with its slime trail on a arbustorum numbered 0, 7, 2, and 6 on those days,
moss (upper right) in Swavesey, Cambridgeshire, UK. Photo by respectively, with weights of 0.0, 8.5, 2.4, and 7.3 per 100
Brian Eversham, with permission. g dry mass of moss. This was surpassed only by those on
Urtica dioica (stinging nettle) reaching 14.4 and Jumping to Escape

Mercurialis perennis (dogs mercury) reaching 16.2.
Nevertheless, it takes a lot of dry moss to make 100 g. A second adaptation to avoid predation is to "jump."
Jumping slugs (Hemphillia; Figure 22) don't actually jump.
Instead, when they are approached by a predator snail or
other predator, they tighten their muscles, coil up, and
straighten rapidly, flopping around on their substrate until
they are free of it, and fall. This effects a rapid motion that
looks like a jump (Leonard 2011). This activity also breaks
the slime trail, facilitating their freedom to "jump." The
slow-moving predator snails don't have a chance. Leonard
says these slugs are potentially successful dispersers of
fungal spores. I would think that would work for
dispersing bryophytes as well, for spores, asexual
structures, and fragments.
Figure 20. The moss-dwelling copse snail, Arianta

arbustorum. Photo © Roy Anderson <habitas.org.uk>, with
permission.
Nighttime activity by many snails is likely to be

greater than that during the day, and little snails may
actually seek refuge in mosses during the day (Grime &
Figure 22. Hemphillia glandulosa, the warty jumping slug,
Blythe 1969). Furthermore, snails like Arianta on moss. This and the above photo illustrate the variability in its
arbustorum (Figure 20) typically climb, often to a coloration. Photos by Kristiina Ovaska, with permission.
considerable height, to obtain food. Bryophytes just don't
fit as a refuge for larger snails, so the behavior of the larger In Canada, some of these Hemphillia (Figure 21-
Arianta arbustorum may not reflect that of the small Figure 24) species seem safe from extinction due to
snails. sufficient abundance, but others are endangered due to
increasing patchiness of suitable habitats (Leonard &
Adaptations Ovaska 2003). The 1994 NW Forest Plan regulates ground
disturbance activities on federal lands in northern
Confusing the Predator California to Washington, protecting "survey and manage"
In the Pacific Northwest, USA, unusual jumping slugs species, including several species of jumping slugs,
in the genus Hemphillia (Figure 21-Figure 24) prefer Hemphillia. Hemphillia dromedarius (dromedary snail;
coarse woody debris or moss mats on decaying logs Figure 23-Figure 24) is officially threatened in both Canada
(Leonard & Ovaska 2003). They have some remarkable and the United States, where it lives in the state of
adaptations for their log habitats. One such adaptation Washington. Legal protection of these slugs can help in the
appears to be to confuse their predators by smearing their protection of mosses in these areas. However, the Bush
slime trail (Figure 17). administration was not sympathetic to this protection and it
could be lost at any time with a change in administrative
philosophy. Perhaps the novelty of its jumping behavior
will increase public interest and sympathy and lead to its
protection in yet another way.
Figure 21. Hemphillia glandulosa, the warty jumping slug,

on moss. This and the following photo illustrate the variability in Figure 23. Hemphillia dromedarius, the dromedary jumping
its coloration. Photo by Kristiina Ovaska, with permission. slug. Photo by Kristiina Ovaska, with permission.
Szlavecz (1986) determined that snail size plays an

important role in their behavior, including food searching.
Although one might think that larger animals need to eat
more, it seems that the larger Monadenia hillebrandi
mariposa (Figure 27) instead spends more time crawling
and less time feeding, permitting it to travel farther.
Although it prefers leaf litter, it consumes mosses as well
(Figure 28). This snail lives in cool, mossy forests and
sometimes hibernates among mosses, including thick moss
on a bigleaf maple branch (Sandelin 2012).
Figure 24. Eggs of Hemphillia dromedarius, the dromedary

jumping slug. Photo by Kristiina Ovaska, with permission.
Keeping It Small
If you want to go clambering among the bryophytes, it
helps to be small (Figure 26). One would expect that size
would also constrain movement among the bryophytes and
restrict larger snails to the surface. But some tiny snails
actually occur fairly deep within the bryophyte mat. Such
is the elongate snail captured by Jan-Peter Frahm deep Figure 27. Monadenia hillebrandi, a consumer of the
within a cushion of Distichium capillaceum (Figure 25). mosses Rhytidiadelphus sp. and Grimmia trichophylla. Photo
by John Slapcinsky, through Creative Commons.
Figure 25. Distichium capillaceum with a snail nestled deep

within the cushion. Photo by Jan-Peter Frahm, with permission.
Truncatellina cylindrica (Figure 26) is another very

small snail. Where it lives at Groomsport, Down, UK, it
occurs in yellow dunes among mosses and the roots of
vegetation on drier, sunny slopes (Anderson 1996).
Figure 26. Truncatella cylindrica on Tortula sp. Note the Figure 28. Laboratory selection of foods by the snail
small size of this conical snail. Photo by Stefan Haller, with Monadenia hillebrandi mariposa. Upper: all data combined.
permission. Lower: juveniles vs adults. Redrawn from Szlavecz 1986.
Monadenia fidelis (Figure 29) lives in dry forests as

well as prairie wetlands where its presence is indicative of
an unburned prairie (Severns 2005). Loubser et al. (2005)
found it associated with nearby mosses in 33% of their
samples. But like many observations of animals with
bryophytes, this may mean that they need bryophytes in
their habitat, that they prefer the same habitats as
bryophytes, or that the relationship is coincidental – the
bryophytes are near something they need. In this case,
mosses are one of its winter hibernating sites, where they
hibernate under mosses in crotches of maple trees
(Monadenia 2016).
Figure 31. Cochlicopa lubricella, moss snail, on mosses.

Photo © Roy Anderson <habitas.org.uk>, with permission.
Turton (1840) also reported another tiny conical snail,

Ena obscura (Figure 32), from mosses and under stones.
But this snail has another way to be elusive from would-be
predators. It covers itself with mud or debris, rendering it
nearly invisible by hiding the shiny shell (The Great Snail
Hunt 2012), but might it also provide a means of
controlling water loss or temperature?
Figure 29. Monadenia fidelis (Pacific sideband snail) on

mosses. Photo by Walter Siegmund through Wikipedia
Commons.
Conical Shape
The terrestrial conical snails, or at least the smaller of
these snails, seem to be more suited to traversing the
internal spaces of bryophytes. Cochlicopa lubrica (Figure
30) and Cochlicopa lubricella (Figure 31), moss snails,
have been known from mosses for a long time. In 1840
Turton reported these snails from mosses and grass on the
ground and under stones in the British Isles.
Figure 32. Ena obscura, a snail that lives in forests or on

walls, under stones and moss (Turton 1840) in the Sulehay,
Northants, UK. It covers itself with mud as camouflage. Photo
by Roger S. Key, with permission.
Avoiding Desiccation
Bryophytes remain moist long after their epiphytic and

rock substrata, and even those on dry soil can become
moist, collecting fog or light rainfall that never reaches the
soil. Hence, they can become a refuge for snails and slugs
seeking moisture. Such is often the case for the banana
slug, Ariolimax columbianus (Figure 15), in the Pacific
lowlands, USA. This slug leaves its moist cover on a
Figure 30. Cochlicopa lubrica on mosses. Photo by moss-covered fallen log to forage at night, then returns to
Malcolm Storey, through Creative Commons. the moss (Sandelin 2012). Taking advantage of the
moisture at night, this slug can travel nearly 0.5 km in among bryophytes. Some snails remain dormant for as
search of more suitable conditions. many as five or six years. Boss suggests that the ability to
The large (up to 13-15 cm) bryophyte-dwelling slug hibernate and aestivate may play a strong role in the
Arion ater (Figure 33-Figure 35) forms a ball by expansion of geographic range, speciation, and extinction.
contracting its body and humping up (Figure 34) (Sandelin The European snail species Fruticicola fruticum
2012). That reduces its surface area and thus reduces water (=Eulota fruticum, Bradybaena fruticum; Figure 36)
loss. It can also twist on itself to reduce exposed surface hibernates from October until a time in spring when the
area (Figure 35). This twisting ability is probably also weather is suitable for it to become active (Künkel 1928).
helpful as it climbs moss setae and feeds on the capsules. It accomplishes this hibernation in dead moss or it may
burrow into the ground with its aperture facing upward.
Figure 33. Black form of Arion ater in an extended position.

Photo by David Perez, through GNU Free Documentation.
Figure 36. Fruticicola fruticum with Polytrichum nearby.

Photo by Michael Becker, through Wikimedia Commons.
No Shell – Slugs
Slugs can be somewhat common on bryophytes and
seem to have the same adaptations as snails. Their only
advantage would seem to be greater flexibility due to the
absence of a hardened and bulky shell, but that brings with
it a greater chance for desiccation. For many species, being
small helps in permitting them to hide from predators and
to maneuver among the bryophytes (Figure 37).
Figure 34. Black form of Arion ater forming a ball by

contracting and humping up. Photo by Emőke Dénes, through
Wikimedia Commons
Figure 37. Keeled slug (Tandonia budapestensis), common

inhabitant of mosses such as this Leucolepis in the Pacific
Northwest, USA. Photo courtesy of Jeri Peck.
The Limacidae is a family of slugs, and both common

genera (Deroceras, Limax) have members that have been
Figure 35. Arion ater juvenile contracting on itself. Photo found among mosses. In the sub-Antarctic Marion Island,
© Roy Anderson <habitas.org.uk>, with permission the slug Deroceras panormitanum (Figure 38; originally
described as the separate species D. caruanae) lives in
Bryophytes can offer the snails and slugs yet another moist bryophyte communities as well as on decaying
means to escape drought and extreme heat or cold. These bryophytes (Smith 1992). With a totally exposed body,
gastropods can hibernate in cold temperatures or aestivate slugs in such harsh environments can find shelter and
in heat or drought (Boss 1974), and this sometimes occurs moisture among the bryophytes.
Figure 38. Deroceras panormitanum on what appears to be

a species of the moss Campylopus. Photo © Roy Anderson
<habitas.org.uk>, with permission.
Brain Eversham (pers. comm. 21 March 2012) tells me

that the yellow slugs, Limax flavus (=Limacus flavus;
Figure 39) and L. maculatus (Figure 40), live mainly on
old walls in Britain, where, like many snails, they are night
active. They feed primarily on lichens and algae, but will
graze on dead plant material if they run out of lichens.
They don't generally eat leafy mosses, but they will browse
on the capsules. He has observed Tortula muralis (Figure Figure 41. Tortula muralis, a species whose capsules serve
41) and Grimmia pulvinata (Figure 42) with the setae as food for species of Limax. Photo by Michael Lüth, with
permission.
remaining but all the capsules nibbled off. He suggests that
the capsules and spores are more nutritious or more
digestible than the leaves and stems.
Figure 39. Limax flavus on a bed of mosses. Photo © Roy

Anderson <habitas.org.uk>, with permission.
Figure 42. Grimmia pulvinata with capsules and awns.

Photo by Michael Lüth, with permission.
In Search of Food
As just described for two species of Limax, snails and
slugs may browse on bryophytes. They have a rasping
tongue (radula) that destroys the epidermis of
tracheophytes (Grime & Blythe 1969), but what does it do
to moss leaves only one cell thick? Apparently in some
cases it makes mosses potential food (Szlavecz 1986), and
Figure 40. Limax maculatus on moss at Bridge House, enables some gastropods to consume even the tough
Swavesey, UK. Photo by Brian Eversham, with permission. capsule (Davidson & Longton 1987, Davidson et al. 1990).
Guy Brassard reported to me that Stéphane Leclerc has

taken a picture of a slug in Quebec, Canada, eating a
Buxbaumia aphylla (Figure 43-Figure 44) capsule!
Michael Lüth (Bryonet 23 September 2017) observed and
photographed a slug grazing on the capsule of Buxbaumia
viridis (Figure 45). Dave Kofranek reports tasting it – it
tastes like cucumbers (Bryonet 24 September 2017).
Figure 45. Buxbaumia viridis with slug eating capsule.

Low Palatability?
Often it appears that the palatability index for
bryophytes is low (Jennings & Barkham 1975).
Figure 43. Buxbaumia aphylla that are immature and have
Furthermore, snails and slugs seem to be less interested in
not been eaten. Photo by Štĕpán Koval, with permission. grazing things with awns than those without. Robin
Stevenson (pers. comm. January 2008) has seen Bryum
argenteum (Figure 46-Figure 47) that is completely grazed
over, but never observed such grazing on an awned
Grimmia species (Figure 42). Could it just be that there is
no nutrition in an awn, or do they have trouble gliding
across the furry tips of leaves?
Figure 46. Bryum argenteum, a moss with no awns and a

Figure 44. Buxbaumia aphylla that may have been damaged food source for snails and slugs. Photo by Michael Lüth, with
by a herbivore. Photo by Janice Glime. permission.
shown for Grimmia pulvinata in Figure 48 (Bryonet 12

January 2008). On the other hand, Frank Greven (Bryonet
13 January 2008) has seen this pattern as a result of grazing
by isopods (wood lice). Robin Stevenson (pers. comm. 14
January 2008) agrees that isopods might be deterred by the
awns, causing them to eat in such a pattern. But in this
case, after climbing up a bridge coping, the snail or
whatever might have found that this moss provided the best
choice available.
Figure 47. Bryum argenteum showing lack of awns. Photo

from UBC website, with permission from Shona Ellis. Figure 49. Grimmia trichophylla in Bretagne (Brittany),
France, showing somewhat less imposing awns than those of
But awns, even in Grimmia pulvinata (Figure 48), Grimmia pulvinata. Photo by Michael Lüth, with permission.
may not deter all snails (Figure 48). Szlavecz (1986) was
able to identify the awned Grimmia trichophylla (Figure
49) in the feces of the California snail, Monadenia
hillebrandi mariposa (Figure 27) and also demonstrated
that the spine tips of the tracheophyte Selaginella hansenii
(Hansen's spikemoss; Figure 50) did not deter feeding or
crawling. Perhaps it depends on the density of the hair tips,
since Grimmia trichophylla (Figure 49) and S. hansenii
(Figure 50) have much less dense hairs than G. pulvinata
(Figure 48), and on the particular species and size of snail
or slug. On the other hand, it appears that the slugs are able
to graze the lower margins of a clump, apparently resting
on the substrate without the need to traverse the awns
(Figure 48).
Figure 50. Selaginella hansenii, a spine-tipped tracheophyte

eaten by the snail Monadenia hillebrandi mariposa. Photo by J.
E. (Jed) and Bonnie McClellan © California Academy of
Sciences, with permission.
Figure 48. Grimmia pulvinata exhibiting grazing that

girdles the base of the clump in a pattern typical of snail or slug
grazing, but also known for isopods. Photo by Robin Stevenson,
with permission.
Michael Lüth has observed snails grazing on

Orthotrichum (Figure 51) and Terry McIntosh has seen Figure 51. Orthotrichum urnigerum, member of a genus
slugs grazing on other bryophytes, with both observers known to be grazed by snails. Photo by Michael Lüth, with
indicating that the damage to the moss was similar to that permission
Low Nutritional Quality?

That rasping tongue is not always enough to
accomplish the task of obtaining nutrients from mosses.
Oyesiku and Ogunkolade (2006) experimented with snails
and the moss Bryoerythrophyllum campylocarpum. In
laboratory experiments, snails (Limicolaria aurora; Figure
52) gained the most weight when fed Bryoerythrophyllum
campylocarpum paste. Snails that had only unground moss
actually lost weight. Those in the field experiment
(restricted to B. campylocarpum) either lost weight or
remained the same. Fecal matter of field snails had
fragments of moss that had lost chlorophyll from their cells
as well as that of abundant algae and Cyanobacteria.
Presence of snails on the moss was seasonal from April Figure 54. Syntrichia laevipila on bark. Photo by Jonathan
until October, when moisture and lower temperature of the Sleath, with permission.
moss may have provided favorable habitat. This
experiment suggests that in this case the snail was unable to Algae growing on mosses, especially in the aquatic
penetrate the cells of the moss, making it an unlikely food habitat, could be a prominent source of food for gastropods.
source in nature. Rather, the researchers suggest that snails In the Negev Desert, adult desert snails (Sphincterochila
most likely use moss as a moist and cool habitat. zonata) fed exclusively on algae on the soil surface,
creating an algal turnover of 142 kg hectare-1, despite being
active for only 8-27 days in winter during the rainy period
(Shachak & Steinberger 1980). Other Negev Desert snails
feed on the mosses themselves. Sphincterochila boissieri
(Figure 55) feeds on shrubs there, but its feces indicate that
it also feeds on the moss Tortula atrovirens (=Desmatodon
convolutus; Figure 56) (Yom-Tov & Galun 1971). This is
a snail that has color morphs of brown and white, but they
apparently don't affect its temperature (Yom-Tov 1971;
Slottow et al. 1993). However, their rodent predators
choose more brown than white snails, enough to be
significantly different (Slottow et al. 1993).
Figure 52. Shell of Limicolaria aurora. Photo by David G.
Robinson, USDA APHIS PPQ at Bugwood.org, through public
domain.
Food for Some

Clearly for some slugs and snails there are bryophytes
that do indeed seem palatable. Ochi (1960) reported that
the thallose liverwort Conocephalum conicum (Figure 53)
served as food for a slug. Merrifield (2000) found evidence
of heavy grazing on epiphytic bryophytes, particularly the
moss Syntrichia laevipila (Figure 54), of Oregon white
oaks (Quercus garryana) in the Willamette Valley, Oregon,
USA, and considered that either springtails or slugs were Figure 55. Sphincterochila boissieri, a species that is known
likely responsible. She considered that the abundance of to eat Tortula atrovirens in the Negev desert. Photo by Mark A.
gemmae on S. laevipila may be a response to this grazing. Wilson, through Creative Commons.
Figure 53. Conocephalum conicum showing feeding Figure 56. Tortula atrovirens, a moss that is eaten by the
damage upper middle) by something, perhaps a slug. Photo by Negev Desert snail, Trochoidea seetzeni. Photo by Des
John Hribljan, with permission. Callaghan, with permission.
Szlavecz (1986) examined feeding preferences in 31

individuals of the snail Monadenia hillebrandi mariposa
(Figure 27). Collections of field feces indicated that they
consumed the mosses Rhytidiadelphus sp. (Figure 57) and
Grimmia trichophylla (Figure 58) in nature, among other
things. In the lab, they preferred shrub and bay litter over
mosses, but preferred mosses and lichens over grasses and
pine litter. More green moss than brown occurred in the
feces, whereas brown material was more common from
consumed tracheophytes (Figure 59).
Figure 57. Rhytidiadelphus squarrosus, a member of a Figure 59. Comparison of green and brown portions of plant
genus that has been found in feces of the snail Monadenia material eaten by the snail Monadenia hillebrandi mariposa.
hillebrandi mariposa. Photo by Michael Lüth, with permission. Modified from Szlavecz 1986.
Figure 58. Grimmia trichophylla showing awns. Photo by

Michael Lüth, with permission.
Grime and Blythe (1969) found bryophytes in the feces Figure 60. Cepaea nemoralis, banded snail juvenile at Old
of four species of snails out of the six examined from Sulehay Forest, UK, a species that lives in a mossy habitat but
Winnats Pass, Derbyshire, England, on 13 October. But apparently does not eat them. Photo by Brian Eversham, with
then, tracheophyte foods often become less nutritious as the permission.
plants prepare for winter. Studies by Chatfield (1973),
Williamson & Cameron (1976), and Richter (1976) In the tropical montane rainforest of Brazil, those
indicate that at least juvenile snails might do best on a small, flattened snails in the Charopidae (Figure 62) eat
mixed diet. But for Cepaea nemoralis (Figure 60-Figure bryophytes (Maciel-Silva & dos Santos 2011). Both
61), it appears that even though mosses are part of their Canalohypopterygium tamariscinum (syn. =
habitat, they are seldom part of the diet (Williamson & Hypopterygium tamarisci; Figure 63) and Lopidium
Cameron 1976). concinnum (Figure 64) had evidence of leaf herbivory,
mostly in the beginning of the rainy season (September to

December). A species of snail in the Charopidae and a
moth larva in the Geometridae were the culprits. Using an
index of damage (ID) in 2007, 2008, Maciel-Silva and dos
Santos found that C. tamariscinum had higher damage
(68%, 35%) than L. concinnum (38%, 23%) in these two
years (Figure 65). These rates were lower than those for
tracheophytes. They found no correlation with phenols,
proteins, or the ratio between them (Figure 65).
Figure 63. Canalohypopterygium tamariscinum, a food

source for Charopidae. Photo by Niels Klazenga, with
permission.
Figure 61. Cepaea nemoralis, a species that lives in a mossy

habitat but apparently does not eat them. Photo by Stefan Haller,
with permission.
Figure 64. Evidence of Charopidae herbivory on Lopidium

concinnum from an Atlantic Forest, Brazil. Photo by Adaises
Maciel-Silva and Nivea Dias dos Santos, with permission.
Figure 62. Charopidae feeding on Lopidium concinnum Figure 65. Charopidae and Geometridae damage to
from an Atlantic Forest, Brazil. Photo by Adaises Maciel-Silva mosses in 10 colonies of plants. Image from Adaises Maciel-
and Nivea Dias dos Santos, with permission. Silva and Nivea Dias dos Santos.
An Avoidance of Gametophores?
Davidson and Longton (1985, 1987; Davidson 1988,
1989) reported that several species of generalist slugs
consumed bryophytes. In some cases, the protonema
(threadlike stage that develops from moss spore) is readily
consumed (Grime 1979). In Great Britain, capsules and
protonemata of several mosses [Brachythecium rutabulum
(Figure 66), Mnium hornum (Figure 67-Figure 68), and
Funaria hygrometrica (Figure 69)] were eaten
preferentially to leafy gametophores by slug species in the
genus Arion (Figure 70) (Davidson & Longton 1987;
Davidson et al. 1990). Cambs (2012) found that the slug
Limax maculatus (Figure 40) likewise would eat capsules,
but the leafy parts seemed to serve only as an emergency
food. It appears that some may even eat calyptrae Figure 68. Mature capsules of Mnium hornum. Photo by
(covering over capsule; Figure 71). Ferulic acid, present Janice Glime.
in shoots but absent in young capsules of Mnium hornum,
is a phenolic compound that is only released after severe
hydrolysis. Its antibiotic role as an antifungal agent (Sarma
& Singh 2003) and in antiherbivory (Seigler 1983; Smith
2011) may contribute to this preference for capsules, as
discussed below. Davidson and coworkers found that older
capsules with spores were less preferred than the green
ones (Figure 72; Davidson & Longton 1987; Davidson et
al. 1990).
Figure 69. Capsules of Funaria hygrometrica – potential

snail food. Photo by Michael Lüth, with permission.
Figure 66. Slug eating capsules of Brachythecium. Note the

number of setae that are missing capsules. Photo by Janice
Glime. Figure 70. Arion rufus on mosses in a woodland above
Poole's Cavern, Buxton, UK. Photo by Brian Eversham, with
permission.
Figure 67. Young, green capsules of Mnium hornum that

are preferred by Arion slugs. Photo by Michael Lüth, with Figure 71. Slug on moss calyptra, apparently finding
permission. something to eat. Photo courtesy of Sarah Lloyd.
Figure 75. Arion subfuscus, a slug known to consume

Mnium hornum. Photo by Gary Bernon, USDA APHIS at
Bugwood.org, through public domain.
Presence of moss cells of Brachythecium rutabulum

(Figure 76) and Mnium hornum (Figure 77-Figure 78) in
the feces of previously starved Arion suggest that the leafy
mosses are not digested well (Davidson et al. 1990). On
Figure 72. Relative damage by slugs (Arion spp.) of the other hand, all three species of slugs named above
sporophyte stages of two species of bryophytes. n=300-500 at readily consumed Funaria hygrometrica (0.4-6.5 mg wet
day 0. LCI = late calyptra stage; EOI = early operculum intact; weight per slug; Figure 69) in overnight feeding trials. The
LOI = late operculum intact; OF = operculum fallen; EF = empty
importance of mosses as food may rest with the organisms
and fresh. Redrawn from Davidson et al. 1990.
living on the mosses – fungi, bacteria, protozoa, rotifers,
The slugs consumed only trivial amounts of etc., making indigestibility of the mosses inconsequential.
Brachythecium rutabulum shoots (Figure 66; Davidson
1989). Mnium hornum (Figure 77) was also ignored, but
after 5-7 days of starvation Arion rufus (10-15cm long;
Figure 73) and A. subfuscus (5-7 cm long; Figure 75) ate
significant quantities of shoots of this species. The garden
slug Arion hortensis (Figure 74) still ignored the moss
even after 7 days of starvation.
Figure 76. Brachythecium rutabulum cells as they might be

seen in feces. Photo by Tom Thekathyil, with permission.
Figure 73. Arion rufus on a bed of mosses. Photo by Jean

Bisetti, with permission.
Figure 77. Mnium hornum shoots – a species that was

Figure 74. Arion hortensis s.s. at Bridge House, Swavesey, ignored in experiments until the slugs were starved. Photo by
UK. Photo by Brian Eversham, with permission. Janice Glime.
Figure 80. Buxbaumia viridis capsules. Note that the leafy

Figure 78. Mnium hornum leaf tip cells, what one might see part belongs to another species of moss. Photo by Adolf Ceska,
in feces. Photo by Bob Klips, with permission. with permission.
It is perhaps not surprising that snails eat the capsules

of Splachnum (Figure 79). This genus has odors that
attract flies, so they may serve as attractants to gastropods
as well.
Figure 81. Buxbaumia aphylla showing exposed green

spores in the capsule that has split open. Photo by Janice Glime.
Slugs also eat hornworts (Anthocerotophyta; Figure

82). Bisang (1996) reported that they especially eat the
green sporophytes.
Figure 79. Snail on setae of Splachnum capsules in Alaska,
eating capsules. Photo courtesy of Blanka Shaw.
Indirect evidence suggests that slugs and snails graze

capsules of Buxbaumia viridis (Gordon Rothero, Birds
feeding on moss capsules, Bryonet-l, 10 April 2003; Figure
80). Stark (1860) relayed a story of the ill fate of collected
specimens of Buxbaumia aphylla (bug-on-a-stick moss;
Figure 81) on their journey from Scotland to England. A
slug had inadvertently been included in the package and it
managed to destroy their prized specimens. On the other
hand, B. aphylla can fool you. After repeated observations
with my graduate student, Chang-Liang Liao, we have
discovered in the field that what appeared to me to be
grazing on capsules of Buxbaumia aphylla is really only Figure 82. Phaeoceros carolinianus, a hornwort with
the splitting of the capsule top as it dries (Figure 81), and mostly green sporophytes, a food source for slugs. Photo by
that this occurs on nearly every capsule. Michael Lüth, with permission.
Deterrents to Herbivory
Longton (pers. comm. 1996) has speculated that
phenolic compounds that protect the leafy gametophytes
deter herbivory, especially on perennials. This could
account for greater herbivory on the annual Funaria
hygrometrica (Figure 83) than on perennial
Brachythecium rutabulum (Figure 66) or Mnium hornum
(Figure 77). The phenolic compounds in the latter two
species were released only after severe hydrolysis, leading
Davidson et al. (1990) to suspect that the phenolic acids
might be tightly bound to cellulose in the cell wall. The
greater palatability of the F. hygrometrica supports the
general theory that perennials invest more resources in
defense against herbivory than do annuals such as F.
hygrometrica.
Figure 83. Young sporophytes of Funaria hygrometrica

before spores form. Photo by Michael Lüth, with permission.
Given the choice of capsules or vegetative material,

both Arion rufus (Figure 3, Figure 70, Figure 73) and A.
subfuscus (Figure 84) preferred immature capsules (see
Figure 85 with a slug on immature capsules of Leucolepis
acanthoneuron) of all three mosses, with Mnium hornum
(Figure 77) being top choice (Davidson 1989). Setae were
generally ignored, but A. subfuscus did occasionally eat M. Figure 85. Slug browsing on immature capsule of the moss
hornum and Brachythecium rutabulum (Figure 66) setae. Leucolepis acanthoneuron. Photo from UBC website, with
All three slugs also ate protonemata in the laboratory, and permission.
for B. rutabulum and Funaria hygrometrica (Figure 83) Davidson and Longton (1987) suggested that Arion
the protonemata were eaten just as much by A. rufus and hortensis (Figure 74) was restricted by the physical
A. subfuscus as were immature capsules. In fact, dry structure of the capsule to consuming developing spores
weight consumption exceeded that of immature capsules. from broken capsules in Polytrichum commune (Figure
Young shoots were also eaten, but less readily. 86); no spores were eaten from unbroken capsules. When
approaching Mnium hornum (Figure 77), the slugs would
withdraw their tentacles, then retreat, suggesting some sort
of chemical deterrent; they behaved similarly in the
presence of extracts from the capsule. It is likely that
hydroxycinnamic and phenolic acids in this species and in
Brachythecium rutabulum (Figure 66) provided this
chemical protection against herbivory (Davidson et al.
1989). Stems of both species were apparently protected by
ferulic and possibly m- and p-coumaric acids bound in the
cell walls of the shoots (Davidson et al. 1989), explaining
the preference of the slugs for capsules. On the other hand,
when moss extracts were placed on communion wafers, the
slugs ate them more readily, suggesting that chemistry
alone was not the likely deterrent (Anonymous 1987;
Figure 84. Arion subfuscus, a slug that prefers immature Davidson et al. 1990). Rather, some physical feature of the
capsules. Photo by Sanja 565658, through Creative Commons. mosses, perhaps the cell wall, deterred these slugs.
removed all the plants by dining on them, leaving behind

only a stump and a slime trail (Figure 88). This was a
research station, so Callaghan needed to find a way to
discourage the slugs. Suggestions from Bryonetters
included sprinkling ground glass around the study area
(Michael Richardson, Bryonet 10 June 2011); putting out
cups of beer to attract and drown the slugs or putting curry
powder or other hot substance around the mosses (Janice
Glime, Bryonet 10 June 2011); copper rings that are
effective in gardens and could be made with a coil of wire
(David Bell, Bryonet 10 June 2011).
Figure 86. Polytrichum commune capsules showing the

persistent hairy calyptra and waxy capsule that is only eaten by
snails when the capsule is broken. Photo by Michael Lüth, with
permission.
Digestibility Figure 87. Hookeria lucens in healthy condition. Photo by

So what did the slugs derive from the consumed Des Callaghan, with permission.
mosses? When they consume preferred foods such as
lettuce leaf or carrot root, the resulting feces contain
macerated, partially pigmented tissue (Davidson 1989).
When they consumed bryophytes, on the other hand, large
pieces of leaf, whole leaves, and even stem pieces remained
intact. Most cells still contained green chloroplasts.
Evidently the moss did little more than fill the gut. Even
the preferred capsules were poorly digested, with capsule
wall fragments, opercula, and peristome teeth remaining.
Mature spores seemed unharmed, but immature spores
seemed to have experienced some digestion, appearing
broken, colorless, and shrivelled. Likewise, the
protonemata seemed to be digestible, resembling the lettuce
and carrots in being macerated and colorless or brown.
Caution must be used in conducting laboratory Figure 88. Temperature/humidity data logger with Hookeria
experiments with food choices. Jennings and Barkham lucens eaten by slugs. Photo by Des Callaghan, with permission.
(1975) found that bryophytes all gave low palatability
scores when six species of slugs, including the three in the Annie Martin (Bryonet 11 June 2011) is a professional
Davidson (1989) study, had a choice of foods. The wider gardener and described her experience in trying to
range of choices in the field may permit them to avoid the eliminate slugs. She suggested putting salt on the head (if
less palatable bryophytes. put on the tail the slug continues to live and eat). Her
Role in Bryophyte Competition with Lichens experience with beer is that it just keeps on attracting snails
night after night, even though many of them drown, so it is
Rosso and McCune (2003) found that molluscs on an ineffective waste of money. Brown mulch seems to
shrubs in the Pacific Northwest, USA, exhibited significant provide a favorable habitat, so she eliminated it, a
herbivore activity on the lichens. Bryophytes, on the other technique that worked, but isn't relevant for discouraging
hand, had little change in cover between stems in snails on mossy rocks.
exclusions and those available for herbivory. It appears
that the mollusc herbivory on lichens (Boch et al. 2011) Aquatic Grazing
may benefit the bryophytes by contributing to the
Grazing by gastropods (slugs and snails) can be so
successful competition of the bryophytes over the lichens severe as to define distribution of a bryophyte species.
in the understory of these forests. Lohammar (1954) found that in northern Europe Fissidens
Palatable Gametophytes fontanus (Figure 89) was absent in lakes where Fontinalis
antipyretica (Figure 90) was also absent. Gerson (1982)
Des Callaghan (Bryonet 10 June 2011) reports slugs suggested that scarcity of Fissidens in some places is due
feasting on the gametophytes of Hookeria lucens (Figure to snail grazing. In the presence of Fontinalis, this smaller
87) near a stream. In only six days they completely moss lives among the Fontinalis fronds where it is
presumably protected from snail grazing by the inedible

forest of Fontinalis surrounding it and the density of the
Fontinalis stems.
Figure 91. Ricciocarpos natans, a species with

molluscicidal properties, floating on the water surface. Photo by
Janice Glime.
Figure 89. Fissidens fontanus, a moss that seems to be
vulnerable to snail grazing except where it is protected by
Fontinalis species. Photo by Michael Lüth, modified by Janice Bryophyte Antifeedants
Glime, with permission.
Based on the foregoing discussion, it appears that at
least some bryophytes are able to discourage browsing by
slugs (Frahm & Kirchhoff 2002). Alcohol extracts of the
moss Neckera crispa (Figure 92) and leafy liverwort
Porella obtusata (Figure 93) have antifeedant activity
against the slug Arion lusitanicus (Figure 94). Extracts of
0.5% dry weight of the moss had low activity, whereas
those from the liverwort exhibited moderate activity at only
0.05%. At 0.25% the antifeedant activity of Porella
obtusata was complete. It is likely that this activity is not
specific for slugs and may discourage insects, bacteria, and
fungi as well.
Figure 90. Fontinalis antipyretica, a moss that apparently

protects the smaller Fissidens from grazing by snails. Photo by
Bernd Haynold, through Wikimedia Commons.
It may be that in the aquatic habitat the snail effect on Figure 92. Neckera crispa, a moss that has antifeedant
some bryophytes is much greater than in the terrestrial activity against the slug Arion lusitanicus. Photo by Michael
habitat. But it is not necessarily all bad. Steinman (1994) Lüth, with permission.
opined that snail grazing could account for the apparent
unresponsiveness of epiphytes following phosphorus On the other hand, Arion lusitanicus (Figure 94), also
enrichment in a woodland stream in Tennessee, USA, known as the murder slug, easily eats the thallose liverwort
where bryophytes were prominent. And some bryophytes Marchantia polymorpha (Figure 95) (Nils Cronberg,
seem prepared to fight back. The thallose liverwort Bryonet 7 April 2016). Cronberg has observed this species
Ricciocarpos natans (Figure 91) exhibits molluscicidal feeding on Marchantia and has noticed that as the slug had
properties that are active against the snail carrier of invaded the wetland, Marchantia polymorpha had
schistosomiasis (Wurzel et al. 1990). disappeared in parallel with the invasion.
secretion can be sticky. It is therefore no surprise that these

animals have dispersal abilities.
Figure 93. Porella obtusata. Photo by Jan-Peter Frahm,

with permission.
Figure 96. Snails such as this one traversing epiphytic

mosses in Japan may be effective dispersal agents. Photo by
Janice Glime.
Slugs are able to disperse the brood branches of

Dicranum flagellare (Figure 97) (Kimmerer & Young
1995). These tiny branches become entrapped in the
secretions and are deposited in the ensuing slime trail.
Kimmerer and Young found that these can be transported at
least 23 cm from the colony, although the mean distance in
their study was only 3.7 cm.
Figure 94. Arion lusitanicus, a slug that traverses mosses,
but finds Neckera crispa and Porella obtusata unpalatable. Photo
by Mogens Engelund, through Wikipedia Commons.
Figure 97. Dicranum flagellare showing the tight flagellate

branches that can be dispersed by slugs. Photo by Janice Glime.
Figure 95. Marchantia polymorpha showing a nibbled And it appears that the secretion increases the ability
thallus on the upper left, about 1/3 down and 1/3 over from the
of the propagule to adhere to its substrate without affecting
corner. It also has a tear that is not likely the result of herbivory.
Photo by James K. Lindsey, with permission. the germination rate. In fact, experiments by Davidson
(1989) suggest that passage of spores through the slug's
digestive system may enhance germination success. All
Dispersal Agents plates containing mature spores from slug (Arion spp.;
It appears that slugs are not all bad in the bryophyte Figure 94) fecal pellets produced shoots, whereas only 80%
world and may instead be a necessary vector for some of the plates with uneaten mature Mnium hornum (Figure
propaguliferous taxa (Stolzenburg 1995). Slugs and snails 67-Figure 68) spores and 70% of those with uneaten
(Figure 96) leave a trail of mucous as they go, and as you Brachythecium rutabulum (Figure 98) spores produced
well know if you have handled these molluscs, this shoots.
Figure 100. Arion vulgaris feces with bryophytes and other

Figure 98. Brachythecium rutabulum, for which the spores material in it. Photo courtesy of Manfred Türke.
germinate better if they have passed through the gut of a slug
(Arion). Photo by Michael Lüth, with permission.
For those snails and slugs that nibble on spores, one

might assume that not all spores end up inside them.
Unless they have perfect aim with that huge foot, their
somewhat clumsy feeding method is undoubtedly going to
render some spores as passengers in the mucous on the
foot. Sooner or later, these will be deposited in a new
location.
The ability of snails and slugs to glide across
bryophytes and to climb setae to capsules suggests that
these animals may be important as dispersal agents. But
how widespread are herbivory and dispersal among Figure 101. Arion vulgaris bryophyte from slug feces.
bryophytes that temporarily host these slow-moving Photo courtesy of Manfred Türke.
animals?
Although we know that bryophyte spores reach the
mollusc gut, experiments are needed to see if spores
expelled in feces are able to colonize successfully.
Davidson (1989) found that Brachythecium rutabulum
(Figure 98) and Mnium hornum (Figure 77) spores eaten
by Arion species actually germinated better than controls.
Manfred Türke sent me images of mosses in the feces
of the slug Arion vulgaris (Figure 99). I was amazed at the
size of the fragment of moss in the feces (Figure 100-
Figure 101). This is a potential means for dispersal, but the
various species of bryophytes must be tested for viability.
Digestive enzymes and extreme pH could damage the moss
cells. On the other hand, the pathogenic fungi Phytophora
spp. (Figure 102) survive as both oospores and filaments
and are viable after passing through the digestive system of
this slug species (Telfer et al. 2015). This was
demonstrated by culturing the feces on agar.
Figure 102. Phytophthora parasitica zoosporangia, a genus
that survives passage through the gut of Arion vulgaris. Photo by
Tashkoskip, through Creative Commons.
To provide additional information on the potential

dispersal ability of slug feces, Boch et al. (2013) fed
capsules of four bryophyte species [Bryum pallescens
(Figure 103), Funaria hygrometrica (Figure 69),
Leptobryum pyriforme (Figure 104), Pellia endiviifolia
(Figure 105)] to three slug species [Arion vulgaris (Figure
99), A. rufus; Figure 3, Figure 70, Figure 73), Limax
Figure 99. Arion vulgaris, a slug that eats mosses, cinereoniger (Figure 106)]. Among the 117 bryophyte
potentially dispersing them. Photo by Dilian Georgiev through samples, 51.3 % of the spore cultures had germination
Creative Commons. following gut passage.
Figure 106. Limax cinereoniger on a mat of moss. Photo

by Michal Maňas through Creative Commons.
Boch et al. (2013) found that germination rates did not

Figure 103. Bryum pallescens with capsules. Spores of this differ among the bryophyte species, but the species of slug
species pass through the guts of several slugs and retain their had strong effects. Among these three slugs, Limax
viability. Photo by David T. Holyoak, with permission. cinereoniger (Figure 106) ate the lowest percentage of the
bryophytes provided, and even correcting for that, they had
the lowest percentage of feces samples (12.9%) producing
protonemata. On the other hand, 76% of those of Arion
vulgaris (Figure 99) and 74% of those of Arion rufus
(Figure 3, Figure 70, Figure 73) produced protonemata
(Figure 107).
Figure 104. Leptobryum pyriforme with capsules. Spores

are able to pass through the guts of at least some slugs and remain
viable. Photo by Michael Lüth, with permission.
Figure 107. Comparison of spore germination from
bryophytes cultured from the feces of three species of slugs.
White bars = Arion rufus; light grey bar = Arion vulgaris, dark
grey bar = Limax cinereoniger. Redrawn from Boch et al.
(2013).
Türke et al. (2013) provide evidence that slugs do

indeed disperse fragments of mosses by consuming spores
and fragments. For tracheophyte seeds, they suggested an
average of 5 m dispersal distance, exceeding the typical
less than 1 m in dispersal by ants. In some slugs, the seeds
are destroyed in the digestive tract, but in other cases they
remain viable propagules.
Boch et al. (2015) discussed several ways that slugs
benefit bryophytes. Their herbivory on tracheophytes
(lignified vascular plants) permits more light to reach the
low-growing bryophytes. But they also crawl across
Figure 105. Pellia endiviifolia with sporophytes. The spores bryophytes and some eat the bryophytes. This puts them in
of this species are able to pass through the gut of several slug the position to disperse spores, fragments, and other
species and remain viable. Photo by Janice Glime. propagules.
Nevertheless, documentation of the effect of the slugs

on the bryophyte community is meager. Boch and
coworkers (2015) designed a factorial common garden
experiment to determine some of the effects of slugs on the
bryophyte vegetation. They collected sporophytes of 11
native and 1 invasive bryophyte species [Barbula
convoluta (Figure 108), Brachythecium rutabulum
(Figure 98), Brachythecium velutinum (Figure 109),
Bryum sp. (Figure 103), Campylopus introflexus (Figure
110), Ceratodon purpureus (Figure 111), Funaria
hygrometrica (Figure 69), Leptobryum pyriforme (Figure
112), Marchantia polymorpha (Figure 95), Phascum
cuspidatum (Figure 113), Plagiomnium affine agg.
(Figure 114), Pohlia sp. (Figure 115)], representing 8
families. They used three enclosure treatments: slugs
previously fed with bryophyte sporophytes, slugs that had
not been fed sporophytes, no slugs. The researchers
demonstrated that bryophyte cover increased in 21 days
from 1.4% to 3.9% in plots where slugs had been fed, an Figure 110. Campylopus introflexus with capsules. Photo
increase that was 2.8 times higher than in the other two by Michael Lüth, with permission.
treatments. After eight months, the species richness was
2.6X higher (5.8 vs 2.2) than in the other treatments. The
researchers concluded that the slugs contributed to
increasing bryophyte cover and diversity by reducing the
dominance of tracheophytes. The early increase in cover in
the enclosures with slugs fed sporophytes suggests that
they also accomplish dispersal.
Figure 111. Ceratodon purpureus with young capsules,

showing the normal proliferation. Photo by Michael Lüth, with
permission.
Figure 108. Barbula convoluta with capsules. Photo by

Kristian Peters, with permission.
Figure 109. Brachythecium velutinum with unopened Figure 112. Leptobryum pyriforme with numerous
capsules. Photo by Michael Lüth, with permission. immature capsules. Photo by Michael Lüth, with permission.
Tetraplodon. However, we need observations of feeding to

determine the identity of the herbivores.
Figure 113. Phascum cuspidatum with unopened capsules.

Figure 116. Tetraplodon mnioides with one capsule eaten

by an unidentified herbivore. Photo by Michael Lüth, with
permission.
Lüth (2010) suggested that the pre-dispersal stage of

the capsules on Splachnaceae are likely to attract
herbivores that differ from the flies that spread the spores.
At this earlier stage, the capsules have a different odor from
that during the dispersal stage. This odor lasts for only a
short time and is therefore often missed by field biologists.
On Bryonet (26 August 2016), Lüth explained that
Splachnum ampullaceum smells like Vaccinium
oxycoccos and occurs in the same habitats, often blending
with these cranberries. And Tetraplodon mnioides (Figure
Figure 114. Plagiomnium affine with developing capsules. 116) smells like Vaccinium myrtillus. Although not all
Photo by Jan-Peter Frahm, with permission.
evolutionary successes are linked to adaptation, it makes
one wonder if these early odors are adaptive to facilitate a
longer dispersal and subsequent deposition in dung,
although one might assume that would require a larger
mammal, not a slug.
Figure 115. Pohlia nutans with immature capsules. Photo

by Michael Lüth, with permission.
When the question of bryophyte dispersal by slugs

arose on Bryonet, Scott Redhead (Bryonet 26 August 2016)
suggested that this might even occur in the Splachnaceae. Figure 117. Splachnum ampullaceum sporophytes with a
To that suggestion, Michael Lüth posted an image of cranberry of similar color to the right. Photo by Michael Lüth,
Tetraplodon mnioides (Figure 116) showing one uneaten with permission.
capsule and one that had been removed by an animal,
possibly a slug, documenting his own observations of I think most people would consider dispersal by snails
capsule herbivory. Christian Schröck (Bryonet 26 August and slugs to be distance-limited. But perhaps, with the help
2016) likewise observed grazed capsules in Voitia and of birds, this is not so limited. Kawakami et al. (2003)
demonstrated that the Japanese White-eyes (Zosterops Killdeer feet out of water. But the likelihood that an
japonicus; Figure 118) and the Brown-eared Bulbuls aquatic snail is carrying bryophyte spores is small due the
(Hypsipetes amaurotis; Figure 119) are birds that eat rarity of capsules. Nevertheless, if a wetland snail has
snails. In fact, five species of snails are able to remain in similar behavior, it has a better chance of having consumed
their shells and appear in the feces. If these snails had spores from wetland mosses.
eaten moss spores, those spores might be transported a
considerable distance, yet be viable in the gut of the snail.
It is probably a rare event. Lots of questions remain in this
relationship, but the scenario brings up interesting
hypotheses.
Figure 120. Charadrius vociferus, a species that disperses

snails on its feet. Photo by Andrew C, through Creative
Commons.
One additional factor determining the suitability of a

slug for spore (or fragment) dispersal is the habitat where
feces are likely to be deposited. Researchers have made the
first steps in understanding the role of slugs in bryophyte
dispersal, but much remains to be explored.
Figure 118. Zosterops japonicus, a bird that passes intact Bryophytes as Home
snails through the gut. Photo by Dick Daniels, through Creative
Commons. Because of their small movement space, bryophytes
can serve as safe sites for smaller snails. Birds can be
significant consumers of snails, particularly during
migration (Shachak & Steinberger 1980), and bryophytes
can make the snails less conspicuous, if not hiding them
completely. In terrestrial habitats, arachnids such as
spiders and daddy-long-legs (Opiliones) are also predators
on snails (Nyffeler & Symondson 2001). While some
spiders can probably navigate the spaces within the moss
mat, it seems unlikely that most mature daddy-long-legs
could manage without getting caught. In addition to the
arachnids, carabid beetles prey on terrestrial gastropods
(Symondson 2004). Some of these beetles use a pump
mechanism to extract the gastropod remains from its shell.
Even snails are predators on slugs. The shell of the
snail makes navigation among the bryophyte branches
more difficult, potentially making the bryophytes a refuge
for the smaller of vulnerable slugs.
Figure 119. Hypsipetes amaurotis, a bird that passes intact In a study of bryophyte inhabitants in the Bükk
snails through the gut. Photo by Nubobo, through Creative Mountains of Hungary, Varga (2008) found the tiny
Commons. gastropods Punctum pygmaeum (Figure 121) and Pupilla
Malone (1965) discovered another possibility, muscorum (Figure 150) among the terrestrial mosses
exemplified by the Killdeer (Charadrius vociferus; Figure Plagiobryum zieri (Figure 122), Hypnum cupressiforme
120). Malone found two species of freshwater snails (Figure 123), and Tortella tortuosa (Figure 124). Standen
attached to the feet of the Killdeer. These were able to (1898) found Punctum pygmaeum from moss shakings.
remain attached and viable long enough to effect dispersal. From my own observations, it appears that snails and slugs
The snail Galba obrussa was able to survive 14 hours on are common on and even in bryophyte clumps, but finding
documentation on the use of bryophytes by these small

species evades even the aggressive Google search.
Figure 124. Tortella tortuosa in Europe. Photo by Michael

Lüth, with permission.
The European snails Azeca goodalli (Figure 125),

Euconulus fulvus (Figure 126), Columella edentula
(Figure 127), Discus (subgen Goniodiscus) rotundatus
Figure 121. The tiny Punctum pygmaeum on Ena (Figure 128), Lauria cylindracea (Figure 129-Figure 130,
montanum, both on a moss. Photo by Stefan Haller, with
permission. Vertigo pusilla (Figure 131), and Vitrina pellucida (Figure
132) live among mosses, among other substrata
(Cloudsley-Thompson & Sankey 1961). Carychium
tridentatum (Figure 133), Discus rotundatus, Cepaea
hortensis (Figure 134), Oxychilus navarricus (formerly O.
helveticus; Figure 135), and several rare species of
Aegopinella (formerly in Retinella) [A. pura (Figure 136),
A. nitidula (Figure 137-Figure 138)] are known under
mossy brick rubble (Verdcourt 1954). Clausilia bidentata
(10-11 mm; Figure 139) is also rare, but can be found
under moss. Standen (1898) reported on Clausilia rugosa
(Figure 140) swarming on mossy walls in the UK and
feeding on mosses and lichens. Standen (1898) found the
snail Acme lineata on a patch of the thallose liverwort
Marchantia sp. (Figure 95).
Figure 122. Plagiobryum zieri, a moss that supports the

gastropods Punctum pygmaeum and Pupilla muscorum. Photo
by Michael Lüth, with permission.
Figure 125. Azeca goodalli shell. Photo by Francisco

Welter Schultes, through Creative Commons.
Figure 126. Euconulus fulvus. Photo by Brian Eversham,

Figure 123. Slug on Hypnum. Photo by Janice Glime. with permission.
Figure 131. Vertigo pusilla on bark. Photo © Roy Anderson

Figure 127. Columella edentula. Photo © Roy Anderson

Figure 132. Vetrina pellucida on bark. Photo © Roy

Figure 128. Discus rotundatus on moss. Photo by

Christophe Quintin, through Creative Commons.
Figure 133. Carychium tridentatum on moss-covered

branch. Photo © Roy Anderson <habitas.org.uk>, with
permission.
Figure 129. Lauria cylindracea on bark. Photo by

Figure 130. Lauria cylindracea, whose small size can be

seen in comparison to this seed. Photo by Christophe Quintin, Figure 134. Cepaea hortensis venturing into one of the
through Creative Commons. Pottiaceae mosses. Photo by Stefan Haller, with permission.
Figure 138. Aegopinella nitidula showing shell coils. Photo

by Brian Eversham, with permission.
Figure 135. Oxychilus navarricus on the moss
Rhytidiadelphus squarrosus. Photo © Roy Anderson
Figure 139. Clausilia bidentata on moss. Photo by

Figure 136. Aegopinella pura on leaf litter. Photo © Roy

Figure 140. Clausilia rugosa on bark, a species that eats

mosses and lichens. Photo by O. Gargominy, through Creative
Commons.
Eucobresia diaphana (Figure 141) lives in humid,

cool places on mountains and in forests of Europe, where it
is likely to encounter mosses, as seen in Figure 141 (Welter
Figure 137. Aegopinella nitidula on moss. Photo © Roy Schultes 2012b), but other than this picture, I can't verify
Anderson <habitas.org.uk>, with permission. what use it might make of them.
Figure 143. Wainuia urnula, a tiny night-active New

Zealand endemic snail that traverses mosses, as shown here.
Photo by Andrew Spurgeon, with permission.
Figure 141. Eucobresia diaphana on a species of the moss

Tortula. Photo by Stefan Haller, with permission.
On the South Pacific Kermadec Islands, Iredale (1913)

remarked that in dry weather one must look for the snails
among the mosses, where they hide from the dryness. He
commented that they are quite variable in choice of trees,
with one bole producing a dozen or more while the next
half dozen adjoining trees disclose none.
Not surprisingly, new species still lurk amid the Figure 144. Wainuia edwardi, member of a genus that lives
bryophytes. Efford (1998) found a new species of the among mosses. Photo by James W. Atkinson, with permission.
carnivorous New Zealand endemic genus Rhytida (Figure
142), and reported observations by others of R. patula and Epiphytic
R. meesoni perampla crawling on mosses and tree trunks Wiesenborn (2003) observed snails in the Riverside
at night. These and other New Zealand snails often fall Mountains of California and found that the active snails
prey to introduced predators. Wainuia urnula (Figure preferred epiphytic mosses (Figure 145) and lichens
143), another night-active snail on mosses, tree trunks, and compared to plant detritus and four sizes of rocks as
rocks, was readily eaten by possums, rats, and hedgehogs habitat. They suggested that the epiphytes could provide
in captivity. Efford (2000) found that 82% of the 315 W. these snails with food or moisture. Tree bark soon
urnula snails examined had an unusual food in the feces becomes a desert after the rain dries up, but mosses remain
and gut – terrestrial amphipods. Its relative, W. edwardi moist much longer, permitting the snails to be active longer
and to search there for food where other small invertebrates
(Figure 144), did not consume amphipods, and no other
likewise take refuge from desiccation.
gastropod is known to consume them. The adaptation for
consuming amphipods appeared to be largely behavioral,
although there were some differences in the teeth.
Figure 142. Rhytida otagoensis, member of a carnivorous

genus that has some moss-dwellers. Image by James Atkinson, Figure 145. Monachoides incarnatus on bark where it often
with permission. encounters bryophytes. Photo by Stefan Haller, with permission.
Tropical islands, especially Hawaii, are particularly The slug Prophysaon vanattae (scarletback
vulnerable to invasive species. With all the visitor traffic taildropper; Figure 148) is one of those slugs that seems to
and import/export business, hitchhikers easily reach the find a safe site under mosses on trees on Vancouver Island,
islands. Snails are among these, and may be one of the Canada (Kristiina Ovaska, pers. comm. 30 June 2009). But
causes of the apparent extinction of the bird called Po'ouli it also hangs on epiphytic moss mats in the moist deciduous
(Melamprosops phaeosoma; Figure 146) (Mountainspring forest there and may even lay eggs there (Figure 149).
et al. 1990). This native Hawaiian bird is especially Pilsbry (1948) suggested that the pupillid snail
adapted to feeding on land snails and insects on branches Bothriopupa variolosa in eastern North America might
and under mosses, lichens, and bark. Its toes are large and prefer mossy rocks and trees.
are used for prying up moss and bark to acquire tree snails.
The bill is stout, withstanding the force needed for
manipulating the snails. Its demise is due largely to
increased activity and habitat modification by feral pigs,
avian disease, and possible gene pool impoverishment due
to low numbers. But it also suffers competition for food by
the introduced garlic snail (Oxychilus alliarius; Figure
147), a native of northwestern Europe (Welter Schultes
2012a) that emits a garlic odor when it is disturbed. This
species is likewise a moss-dweller of mountain slope
forests. It feeds on living and dead plant tissue, but it also
consumes small snails and the eggs of other snails and
slugs (Oxychilus 2011).
Figure 148. Prophysaon vanattae, the scarletback
taildropper, can be found hiding under mosses. Photo by Kristiina
Ovaska, with permission.
Figure 146. Poʻouli (Melamprosops phaeosoma) on a

mossy branch. Note the sturdy beak used to pry loose bark or
crush snails found under bryophytes. Photo through Wikimedia
Commons.
Figure 149. Prophysaon vanattae with eggs on a moss.
Photo by Kristiina Ovaska, with permission.
Calcareous Areas
Because of the need for calcium to make the shell,
many snails are dependent on limestone habitats to obtain
this important resource. Hence, this is a good place to look
for snails on mosses growing there.
Pupilla muscorum (Figure 150) is named for its
occurrence among mosses in Great Britain, although it also
occurs under stones and in leaf litter (Ehrmann 1956). This
tiny (3-4 mm high shell) moss snail often prefers
calciferous ground, but others describe it as indifferent to
limestone content (Nordsieck 2012a). These snails are
ovoviviparous. The eggs can survive over winter inside
the female's body and are laid in the favorable conditions of
spring. At that point, it is not the eggs that must survive
because the juveniles usually hatch during oviposition.
Figure 147. Oxychilus alliarius on moss on bark. Photo © Pupilla triplicata (Figure 151) is likewise a moss dweller
Roy Anderson <habitas.org.uk>, with permission. in Hungary and elsewhere (Deli et al. 2002).
Figure 150. Pupilla muscorum. Photo by Malcolm Storey,

through Creative Commons.
Figure 153. Pomatias elegans at Cheddar, Somerset, UK.
Photo by Roger S. Key, with permission.
Trochulus (formerly Trichia) plebeia (Figure 154)

occurs in wet mossy areas by springs in limestone areas
(Gilbert et al. 2005). Trochulus villosus (Figure 155) lives
in the German Alps and requires high moisture (Welter
Schultes 2010), making bryophytes useful for maintaining
that moisture. This strange genus of snails has hairs on its
shell that help to hold it against wet surfaces (Gilbert et al.
2005). I don't have any indication that these hairs offer any
particular help for living among bryophytes, but if they
have any tactile properties, they could help keep it from
getting stuck between branches by warning that the passage
was getting too narrow.
Figure 151. Pupilla triplicata, a European moss dweller.

Photo by O. Gargominy, through Creative Commons.
Another tiny conical snail (2-3 mm) of calcareous

areas is Acicula fusca (Figure 152) in moss on chalk cliffs
at Ballycastle, and on chalk underlying basalt at Black
Head, Antrim, UK (Anderson 1996). And Pomatias
elegans (Figure 153) occurs on mosses in limestone areas
in the Burren, County Clare, UK (Platts et al. 2003).
Figure 154. Trochulus plebeia, a hairy snail, at Sugley
Wood, UK. Photo by Brian Eversham, with permission.
Figure 152. Acicula fusca, a tiny snail that lives among

mosses on chalk cliffs. Photo © Roy Anderson <habitas.org.uk>, Figure 155. Trochulus villosus on mosses in Germany.
with permission. Photo by Stefan Haller, with permission.
The European family Clausiliidae, known as door

snails, derive their name from the "sliding door" that covers
the opening of the shell (Wikipedia 2012a). This
calcareous door is known as a clausilium, hence the family
name. It permits the snail to retreat into its shell and seal it
off against predators. Cochlodina laminata (Figure 156),
the plaited door snail, lives "between mosses" as well as
leaf litter, but may also be found climbing trees in
deciduous forests and montane pine forests (Welter
Schultes 2012b). Clausilia dubia (Figure 157) is a
calciphilic inhabitant of humid, shady rocks and old walls,
but also lives on tree trunks "full of moss." Michael
Proctor (pers. comm. 23 April 2016) informed me that this
species is very common on Carboniferous limestone in
Yorkshire Dales, UK, in the bryophyte and lichen habitats.
Macrogastra ventricosa (Figure 158), the ventricose door
snail, lives in places with plentiful mosses on the forest
floor or on tree trunks, mostly in the mountains (Welter Figure 158. Macrogastra ventricosa on moss. Photo by J.
C. Schou, Biopix, through Creative Commons.
Schultes 2012b). Macrogastra attenuata (Figure 159)
lives between moss-covered rocks as well as on stones,
rocks, and leaf litter in montane forests.
Figure 159. Macrogastra attenuata, a species of moss-

covered rocks in montane forests of Europe. Photo by Niels
Sloth, with permission.
Vertigo meramecensis (Meramac River snail), unlike a

number of other members of the genus, is a strict calciphile
(Nekola & Coles 2010). It is a species of special concern
that lives in Iowa and Missouri, USA, and dwells in
decomposed leaf litter of moss-covered ledges and shaded
Figure 156. Cochlodina laminata on bark where it appears
to be grazing mosses. Photo by Andrew Dunn, through Creative carbonate cliffs, among other places.
Commons. Bogs and Mires
True bogs are acid, poor fens are acid, intermediate
fens have intermediate pH levels, and rich fens are basic.
For a snail, that pH range is an important consideration in
choice of habitat because of the need for calcium in
forming a shell. Because of this relationship, most
malacologists have considered Sphagnum (Figure 160)
peatlands, heathlands, and pine forests as unsuitable
habitats for snails and consequently have poor snail
biodiversity (Karlin 1961; Kerney & Cameron 1979;
Horsák & Hájek 2003).
In fact, Nekola (2010) found that highly and even
moderately acidic sites had significantly (P<0.000000005)
lower richness and abundance than did neutral and
Figure 157. Clausilia dubia with moss. Photo by O. calcareous habitats. Nevertheless, the typical acid site
Gargominy, through Creative Commons. supported 5-10 species.
But some snails actually thrive in the low pH of bogs

and other acid habitats. One such snail is Vertigo malleata
(Figure 161), an extreme calcifuge. The degree to which
snails have been overlooked in these habitats is exemplified
by finding this new species in 60 sites out of 100 acid sites
investigated from Maine to Florida, USA (Coles & Nekola
2007). In the bogs it was found primarily in leaf litter on
top of the Sphagnum (Figure 160). Nekola (Jeff Nekola,
pers. comm. 16 April 2012) informed me that Vertigo
malleata was virtually absent in the Sphagnum itself,
occurring only where there was leaf litter on top of the
Sphagnum. It would be interesting to watch its behavior if
it is placed amid the Sphagnum. Is it avoiding Sphagnum,
or seeking food only found among the litter? In more
northern locations, V. cristata (Figure 162) or V. perryi
may be present in bogs, but again, they only occur in the
leaf litter, not among the Sphagnum (Jeff Nekola, pers
comm. 16 April 2012). Vertigo cristata is likewise Figure 162. Vertigo cristata, a species that lives on leaf
litter, but not Sphagnum, in bogs. Photo from BIO Photography
common in pine and spruce forests, heaths, and Sphagnum Group, Biodiversity Institute of Ontario, through Creative
peatlands (Nekola & Coles 2010). Commons.
It appears that other snails that live in bogs and poor

fens likewise typically avoid the Sphagnum (Figure 160).
Like Vertigo malleata (Figure 161), Gastrocopta
tappaniana occurs in decomposing leaf litter of fens,
pocosins, and Sphagnum bogs (Nekola & Coles 2010).
Even Vertigo perryi, a resident on the sides of Sphagnum
hummocks, occurs on sedge leaf litter there. And Vertigo
ventricosa (Figure 163) occurs in well-decomposed
graminoid and broadleaf plant litter in the Sphagnum
peatlands and poor fens.
Figure 160. Sphagnum blanket bog, a habitat that does not

provide enough calcium for snails. Photo from Creative
Commons.
Figure 163. Vertigo ventricosa, a species of litter in

peatlands. Photo from BIO Photography Group, Biodiversity
Institute of Ontario, through Creative Commons.
Slugs have much less need for that important element

– calcium (Ca). In boggy habitats, these gastropods would
seem to have little choice but to travel across bryophytes
Figure 161. Vertigo malleata from Lewis Ocean Bay, South (Stanisic 1996). Deroceras laeve (Figure 164) is among
Carolina, USA. This snail lives in Sphagnum peatlands, but the slugs that traverse the complicated topography of bogs
avoids the Sphagnum, living in patches of leaf litter on top of it. and mires. But their specific relationships to the
Photo by Jeff Nekola, with permission. bryophytes seems unknown. On the other hand, another
member of the genus, Deroceras reticulatum (Figure 165), limestone stream, she found 17 snails among the
is a ubiquitous slug, but Anderson (2010) points out that bryophytes, but she found none in the acid stream. Moss
raised and blanket peat or exposed ground above 300 m are inhabitants in the limestone stream included Ancylus
the only habitats where it is not likely to be found. Hence, fluviatilis (Figure 166) and a species of Planorbis (Figure
it appears that physiological differences are important in 167). She pointed out that these molluscs were only
separating these slugs. occasionally found among the mosses.
Figure 164. Deroceras laeve (marsh slug) at Flitwick Moor,

Bedfordshire, UK. Photo by Brian Eversham, with permission. Figure 166. Ancylus fluviatilis showing its close adherence
to the substrate. Photo by Mauro Mariani, through Wikimedia
Commons.
Figure 165. Deroceras reticulatum on a bed of mosses (not

Sphagnum). Photo © Roy Anderson <habitas.org.uk>, with
permission.
Aquatic
In streams, it is likely that snails find mosses as a safe
site from the current. Habdija et al. (2004) rarely found
any gastropods on bryophytes at velocities of greater than Figure 167. Planorbarius corneus. Photo © Roy Anderson
70 cm s-1, whereas oligochaetes became more abundant at <habitas.org.uk>, with permission.
higher velocities. Flow rates are much slower within the
moss mats, thus providing a haven for feeding where the Invasive species such as the carnivorous Euglandina
current is unlikely to dislodge the snails and slugs. This rosea (Figure 168), a native of tropical North America, can
also provides them protection from predators such as fish have severe effects on native snail species elsewhere
(mostly), ducks, shore birds, and amphibians (Pennak (Kinzie 1992). In Hawaii, this species has endangered the
1953). aquatic endemic (Hawaii only) lymnaeid snails due to its
Frost (1942) found a strong difference in gastropod seek and capture behavior. The few surviving individuals
inhabitants among bryophytes between an acid and an are primarily restricted to streamside seeps or damp mosses
alkaline stream in her River Liffey survey in Ireland. In the and liverworts covering rocks near waterfalls.
Figure 168. Euglandina rosea, an invasive carnivore. Photo

through Wikimedia Commons.
Plant Protectors
Not all slugs and snails seem to share a love of
bryophyte habitats. As already noted, some seem to avoid
them. Heinjo During has shared with me a story that
unravelled in the Netherlands, published by Bart van
Tooren (1990). To quote van Tooren, an increasing Figure 169. Sphaerium corneum on an aquatic plant. Photo
number of Linum (flax) seedlings correlates with an © Roy Anderson <habitas.org.uk>, with permission.
increasing number of bryophytes and other plants.
Presumably, the slugs that were eating the seedlings would
not traverse the bryophytes to get to these vulnerable young
plants. They experimented by comparing plots with >70%
cover of bryophytes with those having <20% cover. Their
results were complicated by superimposing treatments of
added water and/or NPK nutrients. In the control plots (no
additions), the survival of Linum (flax) seedlings was
greatest in plots with low bryophyte cover. However, in all
three treatments at Vrakelberg the survival was greatest in
plots with >70% bryophyte cover, whereas at Laamhel the
addition of water plus nutrients was the only treatment that
resulted in a large shift to greater survival with high
bryophyte cover.
Although van Tooren (1990) was unable to Figure 170. Pisidium amnicum. Photo © Roy Anderson
demonstrate significant effects of bryophytes in his 1990 <habitas.org.uk>, with permission.
study, he and his coworkers did find them on the same
slope in the 1981 study (Keizer et al. 1985). Bryophytes
under the growing conditions of that year significantly
reduced mortality of the tracheophytes Linum catharticum
and Carlina vulgaris. Apparently, bryophytes may serve as
deterrents to slugs in some years when weather conditions
might otherwise encourage herbivory, but provide little
support for them in years when nutrients and/or water
availability are different. Such interactions between
species that change with the weather require further
investigation.
Mussels (Bivalve Molluscs)

Mussels are not common bryophyte inhabitants, but
can occasionally occur there in aquatic environs. Frost Figure 171. Leptodictyum riparium, a moss known to grow
(1942) found Sphaerium corneum (Figure 169) and four on freshwater mussels. Photo by Michael Lüth, with permission.
species of Pisidium (Figure 170) among the mosses in the
limestone stream in her River Liffey, UK, survey, but their ECHINODERMATA
typical niches were elsewhere in the stream.
Some bivalve molluscs and other organisms can I refuse to create a chapter for this marine phylum, but
actually turn the relationship around and provide a home one observation is interesting enough to note here. Claudio
for the bryophytes. Yes, some of these animals actually Delgadillo-Moya (pers. comm. 30 March 2016) reported to
have mosses growing on them. Neumann and Vidrine me that a student who is working on sea urchins has found
(1978) found Fissidens fontanus (Figure 89) and moss tissue in the gut of one and leafy liverwort fragments
Leptodictyum riparium (Figure 171) growing on in another! There is no bryophyte known to be marine, but
freshwater mussel shells. some do tolerate sea spray and live near the water. Most
likely one of these, no, two of these, fell into the water or digestive system and survive, thus adding another form
washed in from a stream or river. Resourceful urchin! of dispersal.
Gastropods can be common among epiphytes,
avoid acid habitats, and abound in limestone habitats.
Tiny mussels are able to live among bryophytes in
Summary aquatic habitats. Fissidens fontanus and Leptodictyum
Snails and slugs (gastropods) have often been riparium can live on the shells.
observed on bryophytes. They are adapted to land with Echinoderms generally have no association with
a calcified slime epiphragm to cover the shell opening bryophytes, but if a bryophyte falls into the marine
and respiratory pore in the body. A radula of many water it may occasionally be eaten.
teeth permits them to scrape their food. Reproduction
is mostly by simultaneous hermaphroditism. This
may be facilitated by a love dart that facilitates
movement of sperm cells to the sperm pouch by
Acknowledgments
injecting hormones. Larvae develop within the egg in Bryonetters have been wonderful in making their
most so that the gastropods are typically oviparous. A photographs available to me and seeking photographs from
few are known to deposit eggs in mosses. others. Paul Davison has been helpful in providing
The white desert snail, Eremarionta immaculata, suggestions and offering images. And a long time ago
is common on bryophytes and seems to prefer them as a Allen Neumann sent me a specimen of a clam shell with
habitat. The copse snail, Arianta arbustorum is a Fissidens fontanus growing on it. Numerous
night-active inhabitant. More quantitative studies have photographers and malacologists have been helpful in
shown that some slugs and snails prefer bryophytes. providing images and information. Michael Lüth's
More active snails might be found at night, whereas photographs are a valued contribution. I thank all those
tiny snails might take refuge in the bryophytes during photographers who have made their images available
the day. through the public domain.
Adaptations include "jumping" (Hemphillia), small
size, conical snail, hibernation/estimation, and no shell
(slugs). Snails might use them as a safe site to escape Literature Cited
spiders, daddy-long-legs, and beetles, whereas other
Anderson, Roy. 1996. Species Inventory for Northern Ireland
predators may lurk among the bryophytes. In streams,
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birds, and amphibians.
Anderson, Roy. 2010. Deroceras (Deroceras) reticulatum (O. F.
Bryophyte leafy plants and capsules can serve as Müller 1774) Grey field slug. MolluscIreland. Accessed 18
food for snails and slugs, but some of these molluscs April 2012 at
seem to avoid leaves with awns. Nutritional quality <http://www.habitas.org.uk/molluscireland/species.asp?ID=7
may be poor in some, and some have antiherbivore 4>.
compounds that interfere with development, digestion, Anonymous. 1987. Molluscs prefer manna to mosses. New
and palatability. In some cases the moss structure is Scient. 116: 25.
such that the snails actually lose weight, whereas moss Bauer, B. 1994. Parental care in terrestrial gastropods. Cell.
paste fosters a weight gain. But the gastropods may Molec. Life Sci. 50: 5-14.
gain their nutrition from adhering algae and Berlinger, M., Fischer, M., Prati, D., Knop, E., Nentwig, W.,
Cyanobacteria. In some cases protonemata and green Türke, M., and Boch, S. 2012. Fern and bryophyte
capsules are preferred to leafy plants. Fissidens endozoochory by slugs. submitted to Oecologia.
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And some leafy mosses are palatable.
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But some slugs won't eat the moss even when they Lichen endozoochory by snails. PLoS One 6(4):e18770.
have been starved for 7 days. They have even been
Boch, S., Berlinger, M., Fischer, M., Knop, E., Nentwig, W.,
observed retreating from a moss. Various phenolic
Türke, M., and Prati, D. 2013. Fern and bryophyte
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eat some bryophyte species. Ricciocarpos natans has
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molluscicidal properties that are effective against snail Endozoochory by slugs can increase bryophyte establishment
vectors of schistosomiasis. and species richness. Oikos 124: 331-336.
The moss may not offer any nutrition. Intact cells
Boss, K. J. 1974. Oblomovism in the Mollusca. Trans. Amer.
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Because of their mucous trail, slugs and snails are post101.html>.
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Glime, J. M. 2017. Invertebrates: Molluscs. Chapt. 4-8. In: Glime, J. M. Bryophyte Ecology. Volume 2. Bryological Interaction.

Gastropoda: Snails and Slugs............................................................................................................................. 4-8-2

Figure 3. Great Red Slug, Arion rufus, dark form, Bishop

them to multiply rapidly in a new location and causing

Figure 9. Slugs mating, demonstrating the very long penes.

Mating and the Love Dart

Figure 7. Helix pomatia head during mating. Redrawn from

Figure 10. Roman snails (Helix pomatia) in full foot contact

Figure 8. Helix pomatia head after mating, showing both

Egg and Larval Development

Figure 12. Mating garden snails (Helix aspera) with love

Figure 15. Ariolimax columbianus on moss. Photo by Bill

Figure 18. Eremarionta immaculata in the Riverside

But not all snails and slugs find the bryophyte

Figure 16. Ariolimax californicus exiting a mat of mosses, a

Urtica dioica (stinging nettle) reaching 14.4 and Jumping to Escape

Figure 20. The moss-dwelling copse snail, Arianta

Nighttime activity by many snails is likely to be

Figure 21. Hemphillia glandulosa, the warty jumping slug,

Szlavecz (1986) determined that snail size plays an

Figure 24. Eggs of Hemphillia dromedarius, the dromedary

Figure 25. Distichium capillaceum with a snail nestled deep

Truncatellina cylindrica (Figure 26) is another very

Monadenia fidelis (Figure 29) lives in dry forests as

Figure 31. Cochlicopa lubricella, moss snail, on mosses.

Turton (1840) also reported another tiny conical snail,

Figure 29. Monadenia fidelis (Pacific sideband snail) on

Figure 32. Ena obscura, a snail that lives in forests or on

Bryophytes remain moist long after their epiphytic and

Figure 33. Black form of Arion ater in an extended position.

Figure 36. Fruticicola fruticum with Polytrichum nearby.

Figure 34. Black form of Arion ater forming a ball by

Figure 37. Keeled slug (Tandonia budapestensis), common

The Limacidae is a family of slugs, and both common

Figure 38. Deroceras panormitanum on what appears to be

Brain Eversham (pers. comm. 21 March 2012) tells me

Figure 39. Limax flavus on a bed of mosses. Photo © Roy

Figure 42. Grimmia pulvinata with capsules and awns.

Guy Brassard reported to me that Stéphane Leclerc has

Figure 45. Buxbaumia viridis with slug eating capsule.

Figure 46. Bryum argenteum, a moss with no awns and a

shown for Grimmia pulvinata in Figure 48 (Bryonet 12

Figure 47. Bryum argenteum showing lack of awns. Photo

Figure 50. Selaginella hansenii, a spine-tipped tracheophyte

Figure 48. Grimmia pulvinata exhibiting grazing that

Michael Lüth has observed snails grazing on

Low Nutritional Quality?

Food for Some

Szlavecz (1986) examined feeding preferences in 31

Figure 58. Grimmia trichophylla showing awns. Photo by

mostly in the beginning of the rainy season (September to

Figure 63. Canalohypopterygium tamariscinum, a food

Figure 61. Cepaea nemoralis, a species that lives in a mossy

Figure 64. Evidence of Charopidae herbivory on Lopidium

Figure 69. Capsules of Funaria hygrometrica – potential

Figure 66. Slug eating capsules of Brachythecium. Note the

Figure 67. Young, green capsules of Mnium hornum that

Figure 75. Arion subfuscus, a slug known to consume

Presence of moss cells of Brachythecium rutabulum

Figure 76. Brachythecium rutabulum cells as they might be

Figure 73. Arion rufus on a bed of mosses. Photo by Jean

Figure 77. Mnium hornum shoots – a species that was

Figure 80. Buxbaumia viridis capsules. Note that the leafy