Insects and pathogenic fungi collected in its native habitats have been tested for their biocontrol potential under quarantine. Among the fungi, Ramularia destructiva, Nectria galligena, and an undescribed species of Cryophonectria have been found to cause damage to M. faya in the field in its native habitats and have been, or are being tested under quarantine in Hawaii. Also, a leaf spotting species of Septoria, tentatively referred to S. myricae, is known to infect Myrica cerifera (waxmyrtle) in the eastern U. S. and was found capable of infecting M. faya as well, on which it can cause defoliation.
Branch galls (left, from Maderia) and
cankers (right, from the Azores), and large trunk cankers, were frequently observed.
The cankers were found to be associated with a Nectria sp., referred to N.
galligena. Whereas no fungus fruiting bodies were found associated with the
galls, it was considered likely that these were caused by Nectria as well.
Although N. galligena itself has a broad host range, it was not found on other
woody species in areas where infection of M. faya was common. The disease
on M. faya may be caused by a host-specific form of N. galligena,
although the fungus has not been evaluated experimentally.
A perithecium (sexual fruiting body) of Nectria
from bark canker tissue (left). The Cylindocarpon state (imperfect state)
of Nectria formed in culture (right).
The fungus Ramularia destructiva frequently
was found attacking growing shoots of M. faya throughout its habitats.
Diseased terminals typically curled around, sometimes in a complete circle. The
fungus produced white sporulating material on the diseased tissue surface as shown
(right). Whereas the disease caused by Ramularia was usually not severe,
occasionally it was found to cause significant impact (middle), and more rarely to
severely attack the entire plant, as shown on the young M. faya (right) in the
Canary Islands. The pathogen appeared to be specific to M. faya, as it was
not found on any other plant.
In addition to its occurrence on shoots, R.
destructiva was occasionally observed infecting M. faya inflorescences in
the Azores.
Ramularia destructiva was
cultured in pure culture (left). Conidia (spores) of R. destructiva
(right), with characteristic dark thickenings at the points where they were attached to
conidiophores or to each other. Other than observations of its activity in the
field, this fungus has not yet been tested for its potential as a biocontrol agent.
Among the insects, or insect damaged observed on M. faya in Madeira
was a shoot tip mining caterpillar that hollowed out the pith of developing shoots (arrow)
and destroyed the apical growing point, and a fruit boring caterpillar which destroyed the
seed. Both types of damage are thought to be possibly caused by the same insect, Carposina
atlanticella. More extensive work with potential insect biocontrol agents was
carried out by entomologists and is reported elsewhere (Lutzow-Felling et al., 1995).
A species, apparently undiscribed, of the fungus Cryphonectria was
found in the Azores causing trunk cankers of M. faya. Cankers caused by
this species were not found on other trees, although neighboring chestnut trees were found
with cankers infected by the related C. parasitica, the fungus responsible for
widespread and devastating chestnut blight in the eastern U. S. The fungus fruits on
the surface of cankered bark, as shown on an inoculated M. faya stem in
quarantine (left), producing long, sinuous spore tendrils as viewed with a hand lens
(right).
In hosts-specificity tests in quarantine in Hawaii, cut stem
sections of several woody species were inoculated with Cryphonectria sp.
The fungus sporulated profusely on M. faya (left) and did not sportulate on most
other species tested. However, sporulation was observed on sections of kolea (Myserine
lessertiana) in the family Myrsinaceae, an endemic Hawaiian tree. Such a
reaction was unexpected since these families are not closely related to one another.
Although inoculation of cut stem or branch sections may provide some indication of
susceptibility or lack thereof, the ability of the fungus to attack an intact, living
plant in the field is still a matter of question.
In addition to potential biocontrol agents from the native habitats of M. faya, other fungi are under investigation:
A constant, usually low level of twig dieback, caused
by Botyrosphaeria ribis, occurs through the M. faya population in
Hawaii. B. ribis has been reported in Hawaii on several woody
hosts.
Significant numbers of mature fruit of M. faya in
Hawaii has been observed to become commonly infected with Botrytis cinerea,
a pathogen already present in the environment (Duffy and Gardner, 1994). This
reduction in viable seed may, in effect, be a "spontaneous biocontrol" to which
little previous attention has been given, but without which the invasive ability of M.
faya would be even greater than it is at present.
A species of the leaf-spotting fungus Septoria, at present referred
to S. myricae, is becoming a problem in commercial plant nurseries in the
southeastern U. S. where waxmyrtle ( Myrica cerifera) is grown for use
in landscaping. The fungus becomes a problem mainly under condtions of sprinkler
irrigation. Leaves with several leaf spots typically fall from the plant. The
fungus was tested on M. faya and found to be capable of attacking this species as
well. It is currently under consideration in Hawaii as a biocontrol agent.
Host range testing on plants from more than 20 families has indicated that the
fungus is limited to the genus Myrica. Under permit, the fungus has been
released to the field, where inoculation trials are underway.
Microscopic characteristics of Septoria myricae: a pycnidial
conidioma (left), and conidia (right).
Inoculum of S. myricae is prepared in a kaolin formulation for
storage. It is applied by suspension in an aqueous sucrose solution, and applied to
foliage of M. faya in the field with a commercial back pack power sprayer.
In recent years, an apparent spontaneous die-off of M. faya has been observed in Hawaii Volcanoes National Park. The condition was typically fast-acting, quickly killing previously vigorous trees. Whereas M. faya appeared most susceptible to the die-off, it became obvious that other plant species, including the endemic ohia, were attacked as well. Several possible explantations for the condition were investigated, eventually leading to the feeding activity of the two-spotted leafhopper (Sophonia rufofascia), first reported in Hawaii in 1988. The insect is thought to have originated in China. Thus far, it has demonstrated an exceptionally wide host range in Hawaii, attacking a large number of unrelated crops, ornamentals, and native species. The damage to its hosts is thought to be caused by a potent feeding toxin that is injected during feeding, rather than by a transmitted pathogen, such as a phytoplasma, as had been earlier postulated.
M. faya trees killed by the two-spotted leafhopped in Hawaii
Volcanoes National Park.