Trophobioses between ants and hemipterans (PDF)
by Dirk Mezger, antbase.net
Ants are by far the most dominant arthropods in tropical rainforests in terms of biomass and abundance, and they govern
key functions in their ecosystems (Beattie, 1985; Davidson 1997). Most of them are omnivore because for a solely predacious
life style, there is not enough prey. Many of them depend on trophobioses with hemipterans. Ants gather sugar and amino
acid containing exudates from these plant sap sucking insects. This can be an advantage for both ants and hemipterans because
the hemipterans are protected against predators and parasitoids, and mould is less likely when the sugar containing exudates
are removed by ants. Even the plants can benefit from these trophobioses when herbivorous insects are removed by ants.
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Fig. 1 Coreid bug associated with Crematogaster modiglianii
on climbing bamboo (Dinochloa trichogona) |
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Fig. 2 Delphacids associated with Lophomyrmex cf. bedoti
on climbing bamboo (D. trichogona) |
The rainforests of Borneo are generally very species rich and trophobiotic associations are no exception. In a few square
kilometres of mature rainforests of Danum Valley and adjacent areas covered with low secondary vegetation, 57 ant species
can be found together with 61 species of hemipterans (Blüthgen et al, 2006). In the understorey of primary forests, a species
of climbing bamboo (
Dinochloa trichogona) is very common, harbouring a third of all trophobioses of the forest
understorey (Mezger & Blüthgen in prep.). On this plant hemipterans are very diverse; there are associations between ants and
coreid bugs (fig. 1), delphacids cicads (fig. 2), scale insects and mealy bugs. No other plant is the host of such a diverse
hemipteran community. More that 25 species of ants visit these trophobionts, from small Myrmicinae like
Lophomyrmex
and tiny
Crematogaster Orthocrema) sp., to common species of
Camponotus like
C. arrogans
and
Crematogaster modiglianii to several
Dolichoderus species. At night, sometimes large
Dinomyrmex
gigas come on the bamboo in order to get honey dew from
Notobitus, a large coreid.
Climbing plants like
Spatholobus are also important host plants for some hemipterans visited by ants. Especially
some species of cicadellids and membracids can be found on these plants. In a few cases there are visited by
Crematogaster
modiglianii and
Camponotus rufifemur at the same time. This ant species live together in so called parabioses
(Menzel et al. in prep.). That means that these ant species share a nest in a tree hole.
On seedlings of large trees like
Parashorea (Dipterocarpaceae), trophobioses can be found often (fig. 3). For these
seedlings the trophopbioses are especially important because they seem to be vulnerable to herbivores and the ants can protect
them.
Most of the trophobioses
are found in the canopy of the large trees but they are out of sight.
Observing the large trails of ants along stems of the large trees one
can assume that there a lots of plant sup sucking insects on the tree.
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Fig 3 Larval stages of an unidentified cicadellid together with Lophomyrmex
cf. bedoti on a Parashorea seedling |
Some hemipterans live inside the nests of tree living ants (Weißflog 2001). These ants, e.g. species of
Camponotus
(
C. Karavaievia-group),
Polyrhachis or
Monomorium, built nest of silk and plant particles underside
of a leaf so the hemipterans, mostly coccids, get a maximum of protection and ants' source of honey dew is protected against
competitive ant species. Some ants which have nests inside hollow stems of living plants keep their hemipterans inside these
stems.
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Dolichoderus herdsmen ants are only found in South East Asia. These ants have a very close association with their
hemipteran partner (Dill et al. 2002). These ants are associated with a special group of mealy bugs (Allomyrmococini).
The mealy bugs are continuously guarded, brought to the biwak nest of these ants at night time and carried to their host
plants which include a broad array of plants including the climbing bamboo (fig. 4) mentioned above. In order to transport
their hemipteran partner to suitable host plants these ants have a nomadic lifestyle (Maschwitz & Hänel 1985). These
associations is very specific, every of these ant species has its own mealy bug species.
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Fig. 4 Dolichderus maschwitzi (herdsmen ants) together with Promyrmococcus on D. trichogona
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When leaving the forest and looking to areas covered with low secondary vegetation, there is less diversity of hemipterans.
Many areas are covered by the invasive weed
Chromolema odoratum. On these plant originally from Central America,
only two species of aphid (
Aphis spinocola and
A. gossypi), and sometimes one species of mealy bug and a
coccid can be found. When these areas are not too far away from the forest, quite a lot of ants from all subfamilies can be
found on these associations if no aggressive invasive ants are present. When these invasive species like e.g.
Anoplolepis
longipes have already occupied the site, only few other ant species visit trophobioses nearby them.
Literature
Beattie AJ (1985) The evolutionary ecology of ant-plant mutualisms. Cambridge University Press, New York. 182 pp
Blüthgen N, Mezger D, Linsenmair KE (2006) Ant-hemipteran trophobioses in a Bornean rainforest -diversity, specificity
and monopolisation. Insectes Sociaux, in press
Davidson DW (1997) The role of resource imbalances in the evolutionary ecology of tropical arboreal ants.
Biol. J. Linn. Soc. 61: 153-181
Dill M, Williams DJ, Maschwitz U (2002) Herdsmen ants and their mealybug partners. Abh. Senckenberg. Naturforsch.
Ges. 557: 1-373
Maschwitz U, Hänel H. (1985) The migrating herdsman Dolichoderus (Diabolus) cuspidatus:
an ant with a novel mode of life. Behav. Ecol. Sociobiol. 17: 171-184
Weißflog A (2001) Freinestbau von Ameisen (Hymenoptera: Formicidae) in der Kronenregion feuchttropischer Wälder Südostasiens.
Dissertation. University of Frankfurt, Frankfurt/Main, Germany
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