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Parasites of Phasmida

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Abstract

Parasites are recorded from several species of phasmids from Borneo. Keywords: Phasmida, Parasite, Ectoparasite, Endoparasite, Arachnida, Mite, Nematode, Mermithid, Acacus sarawacus, Asceles margaritatus, Carausius abbreviatus, C. sanguineoligatus, Dares ulula, Haaniella echinata, H. grayi, H. dehaani, H. saussurei, Hoploclonia gecko, Lonchodes jejunus, Sarawak, Sabah, Borneo.
The Entomologist -112
(l),
37-42 (1993)
Parasites of Phasmida
P.
E.
BRAGG,
8 Cornwall Avenue, Beeston Rylands, Nottingham NG9 1NL,
UK.
Keywords: Phasmida, Parasite, Ectoparasite, Endoparasite, Arachnida, Mite, Nema-
tode, Mermithid, Acacus sarawacus, Asceles margaritatus, Carausius abbreviatus,
C.
sanguineoligatus, Dares ulula, Haaniella echinata,
H.
grayi, H. dehaani,
H.
saussurei,
Hoploclonia gecko, Lonchodes jejunus, Sarawak, Sabah, Borneo.
Introduction
The
occurrence
of
parasites
in
stick insects has been noted by a number of authors.
There are undoubtedly numerous bacteria, viruses and fungi which parasitize the Phas-
mida, although few
of
these have been recorded (Bedford, 1976; Casimer, 1956; Tho-
mas & Poinar, 1973). Surprisingly there are also very
few
records of animals which are
parasitic on the Phasmida.
The
vast majority of those recorded are insects
of
the orders
Diptera and Hymenoptera. This
paper
reports parasites
in
several species of phasmids
from Borneo; there are no previous records of phasmid parasites from this area.
Previous Records
Bedford's (1978) review of the phasmids gives the insect parasites of Phasmida as be-
longing to the families: Tachinidae, Ceratopogonidae, (Diptera); Eupelmidae, Chrysi-
didae and Cleptidae (Hymenoptera). In addition to the insects listed by Bedford, King
(1867) noted the occurrence of 'Ichneumon
fly
larvae'
in
the adult females of Anisomor-
pha buprestoides (Stoll) and
'a
minute species of Ichneumon
fly,
probably of the Chalci-
didae'
in
the ova
of
this species. Bedford's review also overlooked two papers which
mention some of the non-insect parasites of phasmids, those by Henry (1937), and
Casimer (1956).
Ticks (Arachnida: Acarina) and Microsporidia (Protozoa: Cinidospora) are listed
in
Carlberg's (1986) review. Carlberg's reference
to
ticks
is
based on a photograph by
Workman (1978) which shows mites
of
the genus Lepus on A. buprestoides.
The
only
written references to mites are from Australia and the USA.
The
first occurred
in
Time-
ma californica Scudder, an immature mite on the thorax (Henry, 1937).
The
second
is
the occurrence
of
red mites, Allidosoma froggatti (Oudemans), found on Podacanthus
wilkinsoni Macleay (Casimer, 1956).
The
third
is
a brief mention of a 'possibly parasitic
mite' on the tarsus
of
a nymph of Acrophylla sp. (Daniels 1987). Some confusion could
arise
due
to the use of the common name 'stick-ticks' which has been used for the dipter-
an family Ceratopogonidae (eg. Wirth, 1971).
The
only parasitic worms recorded are those of the phylum Nematomorpha, which
parasitize the nymphs and adults
of
Podacanthus wilkinsoni Macleay (Campbell and
Hadlington, 1967), and
one
example of an adult female Timema callfornica Scudder
which was parasitized
by
a nematode (Henry, 1937).
37
P.
E.
Bragg
Harman
(1987) states that records
of
parasites
of
Phasmida
are
largely
of
Neotropical
origin.
There
are
however
quite
a
number
of
records for
the
Australian region, including
an
attempt
at biological control
of
phasmids using parasites
in
Fiji (Paine, 1968).
There
appear
to be no records
of
parasites
of
Phasmida
for the
Oriental
region; the closest
record
is
for
Ceratopogonidae
on
Buru
Island
(Edwards,
1926).
New Records
The
following new records
are
the results
of
collections
of
phasmids
made
over
four
years in various
parts
of
Borneo.
The
purpose
of
these collections was to try to establish
the phasmids in culture, so live specimens were
brought
back to Britain.
At
the time
of
capture
several specimens were found to have ectoparasites
and
subsequently several
other
specimens
were
found to
be
suffering from
endoparasites.
The
first two collections,
in
1988
and
1989, were restricted to
Sarawak
and little notice
was
taken
of
ectoparasites, the sole concern being to remove
them
in
order
to increase
their chances
of
survival. In 1990 collections were
made
on Mt. Kinabalu in
Sabah
and
at
various locations
in
Sarawak.
In
1991
a collection
of
phasmids was
made
as
part
of
the
University
of
Brunei DarussalamJRoyal Geographical Society Brunei Rainforest Pro-
ject
at
Kuala Belalong
in
the
Temburong
district
of
Brunei;
a collection was also
made
in
Sarawak.
During
the 1990
and
1991
collections
more
notice was
taken
of
the
occurrence
of
parasites.
A)
Ectoparasites
In 1989 ectoparasites were found
on
specimens
of
Haaniella grayi grayi (Westwood).
Although not closely
examined,
they were
apparently
reddish-brown
coloured
mites
which were found
on
the
antennae
and
leg joints
of
the phasmids.
They
occurred
on
at
least half
of
the
15
specimens collected
near
kampong
Bengoh,
30 miles south east
of
Kuching. Affected specimens were found on two
mountains
in
the
area
and
also in the
farmland
at
the kampong.
The
mites were
found
on
both males
and
females, from third
instar nymphs up to
and
including
the
adult stage. All affected specimens had several
mites
and
one
large female
had
about
20 mites. In 1990 and 1991 a
number
of
infested
specimens
of
H.
g.
grayi were collected on Mt. Serapi, Sarawak. In 1991,
of
24 exami-
ned, all
but
one
were found
to
be carrying brownish-red mites,
the
exception was a first
instar nymph.
The
infested individuals included all sizes
of
insects from first instar to
adult.
At
Poring Springs on Mt. Kinabalu in 1990, two
out
of
four specimens
of
Haaniella
echinala
Redtenbacher
were found to have several mites
attached
to
the
abdominal pits.
A third infested specimen
of
this species was
one
of
five found
near
Kapit,
Sarawak,
also
in
1990.
Not
all
of
the
specimens were closely
examined
in
1990
and
it
is
likely
that
more
than
three
ofthe
specimens were infested. In 1991,
at
Kuala Belalong,
Brunei,
21
speci-
mens
of
this species were
examined
and all specimens, which included adults
and
first
instar nymphs, were found to be infested.
38
Parasites
of
Phasmida
In
1991
six specimens
of
Haanie//a saussurei Kirby were collected
in
Sarawak,
one
from Simunjan and five from the outskirts
of
Kuching; all
were
found to be carrying
brown mites.
Thirteen
specimens
of
Haanie//a dehaani (Westwood) were collected
on
Mt. Serapi,
Sarawak in 1991
and
all were found to be carrying brown mites.
On
Mt. Serapi,
Sarawak,
in
1990, seven female
and
four males
of
Dares u/u/a (West-
wood) were collected.
One
female was found with
one
red coloured mite on the
upper
surface
of
the
abdomen.
In August 1990 I accompanied
Mr
c.L.
Chan
of
Kota Kinabalu, Sabah to Mt. Kina-
balu Park
Headquarters.
Four
nights were spent collecting phasmids in the
area
around
the
Headquarters.
During this time
about
twenty specimens
of
a wingless form
of
Asce-
/es
margaritatus
Redtenbacher
were found to be infested with small brown mites. r re-
moved and preserved a
number
of
these mites which affected about
50%
of
this species.
The
mites occurred
on
the
thorax,
abdomen,
legs and
antennae
of
the
Phasmids. Most
infested specimens had several mites
and
several specimens had
more
than ten mites
on
them.
The
mites affected adults and nymphs
of
both
sexes.
The
mites
on
adult males
were
concentrated
around
the
base
of
the
rudimentary wings.
Two
other
affected species
of
the sub-family Necrosciinae
were
also found near Kina-
balu Park
H.Q.
One
male Asce/es dorsa/is
Redtenbacher
had a single red coloured mite
attached to
the
base
of
the fore wing, two females
of
the same species showed no signs
of
parasitism. A total
of
11
Acacus sarawacus (Westwood) were collected near the
Park
H.Q.;
one
adult female was found to have a red mite
on
the
upper
surface
of
the
abdo-
men.
B) Endoparasites
One
adult female specimen
of
Lonchodes jejunus
(Brunner),
collected at
Damai,
Sara-
wak
in
December
1987,
became
listless within a day
of
arrival
in
Britain.
The
following
day a mermithid larva was found
in
the cage. During the next few days the insect became
more
torpid
and
more
mermithid larvae were found.
The
insect was found to
be
dead,
having produced six
worms
over
a period
of
seven days.
These
worms were all off-white
in
colour,
Imm
diameter
and in excess
of
25cm long, the largest being 36cm.
The
insect
has a body length
of
15cm.
The
only
other
phasmid brought back alive from
Damai
was
another
adult female L. jejunus. This laid 59 eggs during the following
three
weeks and
was then discovered
dead
with a single 30cm worm
in
the cage. Several eggs
were
hatch-
ed successfully
and
gave rise to a sexual culture.
None
of
the
second generation
were
affected by parasites.
A single adult female Carausius abbreviatus
(Brunner)
was captured at Bengoh, Sara-
wak
in
July 1989. a
few
days later
it
was found almost
dead
on
the
bottom
of
its cage.
Lying nearby was a living Mermithid larva approximately 20cm
in
length and
Imm
diameTer.
The
in~ec[
and worm have
been
deposited
in
the Sarawak
Museum,
Kuching.
In August 1990 at Mt. Kinabalu Park a female Carausius sanguineo/igatus
(Brunner),
one
of fvur collected, was fuund
dead
two days after capture;
in
the
cage a single living
39
P.
E.
Sragg
mermithid larva was found.
In
August
1991,
at
Kuala
Belalo:lg
Brunei,
one
male,
of
the
21
specimens
of
H.
echinata which
were
collected, died
and
produced
a S4cm long mermithid larva a few
days
after
being caught.
In
January
1988 a pair
of
Hoploclonia gecko
(Westwood)
were collected in at
Bako
National
Park,
Sarawak.
In early
March,
twelve weeks after
capture
the female
became
listless,
and
approximately
eight
worms
were
found
around
the ovipositor
and
anus.
These
worms were
between
Imm
and
2mm
in
length
but
they were not closely exami-
ned.
They
may
have
been
nematodes
or
insect larvae
but
their size
and
general propor-
tions suggests
that
they
were
probably
insect larvae.
The
phasmids were being
reared
in
a room which was
unheated,
apart
from sufficient
to
keep
the
temperature
from falling
below
S°c.
The
low
temperature
and
the fact
that
other
cultures have
never
shown
signs
of
similar infestations, suggests
that
these
parasitic
worms
may have
been
of
Sarawak
origin.
Discussion
The
endoparasites
of
L. jejunus,
C.
sanguineoligatus,
C.
abbreviatus and H. echinata
appeared
to be identical.
The
specimen from
C.
sanguineoligatus has
been
identified by
Dr
DJ.
Gibson
of
the
NHM,
as a mermithid larva
(Nematoda:
Mermithidae).
The
emergence
of
the largest endoparasites so soon
after
capture
suggests that
the
stress
of
travel, change in climate
or
change
of
the host's
diet,
may have caused the
parasites
to
emerge
early.
If
this were not the case I would have
expected
some
further
evidence
of
infestations
in
specimens which lived much longer. Specimens
of
L. jejunus
lived for up
to
six
months
after
capture
and
other
species lived for
up
to
one
year, but
all
the worms were found within two weeks
of
capture
and
usually within a few days.
As
might have
been
expected,
endoparasites
occurred
in
the
older
phasmids, these having
had longer to
encounter
a parasite.
The
rate
of
parisitism seems high; 19%
of
the 200 specimens from Mt. Serapi
were
found
to
be affected
in
1991.
During
1990
more
attention
was paid
to
the possibility
of
ectoparasites
and
the
number
of
affected species which were noticed implies
that
some
may have
been
overlooked at
Bengoh
in
1989
where
at least
S%
of
the 180 specimens
were affected.
However
it should
be
noted
that collections were
made
at
ground
level
and
therefore a disproportional
number
of
Heteropteryginae
were found.
The
Heterop-
teryginae
are
generally slow moving,
ground
dwelling insects and
the
freqency
of
para-
sitism
in
the
more
mobile subfamilies
appears
to
be much lower.
Many
of
the fully
winged phasmids seem to live mainly in
the
canopy
and
are
not frequently
encountered;
although I have collected
over
one
hundred
specimens only
one
has
been
parasitized,
the
male Asceles dorsalis from Sabah.
In
1991
special
attention
was paid
to
specimens
of
Haaniella
spp.,
the
results strongly
suggest
that
parasitism by mites
is
normal for this genus.
This
genus
is
essentially
ground
dwelling
and
therefore likely to be
in
frequent
contact
with mites in the leaf litter.
40
Parasites
of
Phasmida
Sununary
Parasitism
of
phasmids by mites and
nematodes
appears
to
be
more
widespread
than
is
suggested by existing records. Parasites have
been
found in twelve species from
Borneo
which have not previously
been
recorded
as hosts.
Members
of
the genus Haaniella are
usually infested by parasitic mites.
Acknowledgements
I wish
to
express
thanks
to Virginia
Cheese
man
(entomological livestock supplier) for
providing
the
collapsable cages used
during
the
1989
and
subsequent
collections
and
to
c.L.
Chan
for his hospitality during my 1990 visit.
The
1991
expedition was partly
supported
by a
grant
from
the
Royal Entomological Society.
References
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G.O.
(1976) Description
and
development
of
the eggs
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of
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G.O.
(1978) Biology
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U.
(1986) Phasmida: A biological review (Insecta). Zoologisher Anzeiger,
216: 1-18.
Campbell,
K.
G.
&
Hadlington,
P. (1967)
The
biology
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the
three
species
of
phasmatids
which
occur
in plague
numbers
in forests
of
south
eastern
Australia. Foresl1y
Commission N.S.
W.
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M. (1956)
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muria violescens Leach. Bulletin
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94-95.
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(1926)
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A.
(1987)
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Phasmids -'Stick-ticks' -Biting midges
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family
Ceratopogonidae
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L.M.
(1937) Biological
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Timema californica
Scudder
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Timemidae).
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C.B.
(1867)
Un
titled
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Paine,
R.
W. (1968) Investigations for
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Graeffea
crouanii
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Guillou).
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567-604.
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P.
E.
Bragg
Thomas, G.M. & Poinar,
G.O.
(1973) Report of diagnoses
of
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1972. Hilgardia, 42: 261-359.
Wirth, W. W. (1971) A review
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pomyia subgenus Microhelea parasitic on walking stick insects (Diptera: Cerato-
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42
... In Insects of Australia, Key (1970) mentions mites and mermithid nematodes as parasites; Young (2009) records parasitic flies as common parasites in the USA, and recently Mirzaee et al. (2021) recorded parasitic mites in Iran. The absence of any records of mantid parasites in Borneo may due to a low level of collecting and a lack of reporting; parasitism in stick insects (Insecta: Phasmida) was unknown in Borneo until I reported it in 11 species (Bragg 1993). Notes: Key to femoral spines: s = small, L = large, t = tubercule (where a spine would normally be expected). ...
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The stick-insect Graeffea crouanii (Le Guillou) is a pest of coconuts of local and sporadic importance in the south Pacific and there have been recent outbreaks on Taveuni Island, in the Fiji group. As there appear to be virtually no parasites of the nymphal stages, a preliminary search was made in 1960 for parasites of other palm-feeding Phasmatids in Melanesia. This revealed the presence of Tachinidae parasitising species of the genera Ophicrania and Megacrania , and in 1963–64 these were studied in the Solomon Islands. The Tachinidae comprise at least two species of Mycteromyiella: M. laetifica (Mesnil) attacking both O. leveri Günth. and a species of Megacrania in the western Solomons, and M. phasmatophaga Crosskey attacking 0. leveri and some other Phasmatid hosts on Guadalcanal. The early stages of both species of Mycteromyiella are briefly described and compared, with notes on their bionomics. There was no evidence of any egg parasite attacking Ophicrania in the Solomons. O. leveri , which is very closely related to G. crouanii , has never caused significant damage to coconuts in the Solomon Islands, except on the small island of Savo, from which its Tachinid parasites appear to be absent. It is concluded that Mycteromyiella , especially M. laetifica , which appears to be fairly specific, may be an important factor in the control of O. leveri in the Solomons. The principal hosts of O. leveri are the sago palm ( Metroxylon salomonense ) and species of arecoid palms in the forest, on all of which the insect has better scope for concealment than on coconuts. Collections of nearly 6,000 examples of O. leveri from sago palm on Kolombangara island, in the western Solomons, in August 1963-February 1964 showed an average parasitism by Mycteromyiella laetifica of 28 per cent. Eggs of the Tachinid are laid on nymphs of all instars as well as on adults. The host-survival rate was about 30 per cent, for nymphs and 50 per cent, for adults. A small number of parasitised specimens of O. leveri from Kolombangara were released on Savo, but there was no evidence six months later that M. laetifica had become established there. Breeding trials at Honiara showed that O. leveri could be reared successfully in captivity but not M. laetifica , which shows reluctance to mate in cages and has a narrow range of environmental tolerance in the pupal stage, in which a mortality of at least 70 per cent, seems unavoidable under the conditions practicable for transportation of this stage by air. Between October 1963 and March 1964 nearly 960 puparia of M. laetifica were sent to Fiji. About half of them were used for breeding trials, which showed that the Tachinid could be reared through G. crouanii in captivity but could not be maintained. The rest were released on Taveuni, but a further outbreak of G. crouanii during 1965 yielded no evidence that M. laetifica had become established. Material of M. phasmatophaga , which has a more restricted choice of environment than M. laetifica , but also a somewhat greater potentiality for killing its host, was obtained by exposing O. leveri on seedling coconut palms planted in the forest at Honiara. Quantities were insufficient for transmission to Fiji; 150 parasitised hosts were released on Savo but samples of O. leveri collected there six months later gave no indication of its establishment. Despite this initial failure, it is considered that Mycteromyiella could bring about the control of G. crouanii in Fiji and other affected Pacific islands, and the means by which this might be achieved are discussed.
Phasmida: A biological review (Insecta)
  • U Carlberg
Carlberg, U. (1986) Phasmida: A biological review (Insecta). Zoologisher Anzeiger, 216: 1-18.
The biology of the three species of phasmatids which occur in plague numbers in forests of south eastern Australia
  • K G Campbell
  • P Hadlington
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