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USA (1870s-1970s)Diapheromera femorata
AUSTRALIA (1880s-1980s)Didymuria violescensPodacanthus wilkinsoniAnchiale austrotessalata
FIJI (1920s-Present)Graeffea crouanii
CHINA (1980s-Present)Various spp.
MAJOR INFESTATIONS
“…eggs and fecal pellets are dropped to the ground in great numbers, producing
a pattering sound, like rain, accompanied by a peculiar seething
sound of thousands of jaws chewing the leaves.”
- Helfer (1963)
REVIEW OF DIAPHEROMERA
Wisconsin Outbreak (1959-60s)
“Within seconds after a person stopped under or near a tree, walkingsticks
dropped onto and crawled upon his body so that even to an experienced
entomologist it was an uncomfortable, creepy feeling.”
Oatman (1965)
REVIEW OF DIAPHEROMERA
PODACANTHUS WILKINSONI
Right: Female, Kennedy HarrisBelow: Male (LT,PLT), ANIC (Paul Brock)Phasmida Species File
Readshaw (1965) A Theory of Phasmatid Outbreak Release
Campbell (1974) Factors Limiting the Distribution and Abudance of the Three Species of Phasmatids (Phasmatodea: Phasmatidae) Which Occur in Plague Numbers in forests of South-Eastern Australia
Readshaw (1990) Phasmatid Outbreaks Revisiting
CLASSIC LITERATURE
Outbreaks are infrequent
Semi-voltine life cycle
Inability of predators and parasitoids to contain infestation
High fecundity
Slow migration
READSHAW’S CRITERIA
KENTROMORPHISM
Individuals reared at diff erent densities have diff erent colouration and morphology
Potentially analogous (homologous?) to density dependence in plaguing Orthoptera?
How general is this theory?
Take the model of Australian outbreaks
Apply it to other outbreaks
See what breaks
TESTING THE THEORY OF PHASMATID OUTBREAK RELEASE
Expanded by Campbell to be require an absence of forest catastrophe (mainly fi re) for a period of 10-80 years
No forest fi res associated with D. femorata outbreaks…
… but they always seem to occur in plantations or second growth forest
Conclusion:Outbreaks are linked to habitat disturbance, whether that be natural (forest fi re) or anthropogenic (plantations)
OUTBREAKS ARE INFREQUENT
Readshaw predicted this was important, as infestation levels only occur every other year, preventing a rapid escalation in predators/parasites
D. femorata is generally uni-voltine. Some Chinese pest species are up to tri-voltine.
Conclusion:May be important in specific in cases, but more generally phasmid outbreaks happen without this condition.
SEMI-VOLTINE LIFE CYCLE
Myna bird, introduced to Fiji to control Graeffea crouaniiGerald McCormack
PREDATORS AND PARASITOIDS
Exova tunana, parasitoid of Eurycantha insularisKimsey, Dewhurst & Nyaure (2013)
PREDATORS AND PARASITOIDS
If there is an infestation – the stick insects are not being predated or parasitised at a suffi cient rate to control the outbreak.
True by definition.
PREDATORS AND PARASITOIDS
All species involved lay many times more eggs than would be needed if no predators or parasitoids were present.
This is true for all Phasmid species.
Facilities outbreaks when population rise in phasmids is not immediately followed by a rise in predators and parasites.
Need to look at why this occurs. Given what else we know habitat disturbance is likely (forest fi res; altering of the forest ecosystem; movement of phasmids into plantations)
HIGH FECUNDITY
Diapheromera is wingless – movement confined to walking.
Gravid females of Australian species can glide – limited fl ight. Reports show that lighter males disperse more rapidly than females. Concentration of eggs beneath females.
Outbreaks generally occur in a very confined area, that spreads over several years and may merge with others to form widespread damage.
SLOW MIGRATION
Not known in any phasmids besides the Australian ones mentioned by Key.
The Australian species come from forest that is subject to natural forest fi res, this could be an evolutionary adaption to the eff ects of these.
KENTROMORPHISM
Disturbance of forest (either natural or anthropogenic)
Reduction in controlling species allows phasmid population to rise more rapidly than their predators
High fecundity and slow migration creates intense population rises on limited scales (at least initially)
Rejected ideas:Semi-voltine life cycle
Kentromorphism
GENERAL THEORY OF OUTBREAKS