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1972 A Description of The Adult and The Nymphal Stages of Schizodactylus Monstrosus Drury Orthoptera

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1972 A Description of The Adult and The Nymphal Stages of Schizodactylus Monstrosus Drury Orthoptera

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Journal of Natural History

ISSN: 0022-2933 (Print) 1464-5262 (Online) Journal homepage: https://www.tandfonline.com/loi/tnah20

A description of the adult and the nymphal stages


of Schizodactylus monstrosus (Drury) (Orthoptera)

Narain Khattar

To cite this article: Narain Khattar (1972) A description of the adult and the nymphal stages of
Schizodactylus�monstrosus (Drury) (Orthoptera), Journal of Natural History, 6:5, 589-600, DOI:
10.1080/00222937200770521

To link to this article: https://doi.org/10.1080/00222937200770521

Published online: 17 Feb 2007.

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J. nat. Hist., 1972, 6:589-600

A description of the adult and the nymphal stages of


Schizodactylas monstrosas (Drury) (Orthoptera)
N A R A I N KHATTAI~
Department of Zoology, Government College, Ajmer, India

Introduction
Schizodactylus monstrosus, first described as the cricket of Bengal, is mainly
found in India, Pakistan, Ceylon and Burma. I t is also commonly known as
the ' Bherwa of Bihar ' and the maize cricket, although it does not belong to
the family of true crickets. The present author has already described much
of the morphology and anatomy of this insect (Khattar 1958, et seq.; K h a t t a r
and Srivastava 1962). These studies confirm its placement in a separate
family Schizodactylidae.
During the course of the present studies, it was with great difficulty that
observations could be made owing to the insect's nocturnal and burrowing
habits. As such, though the entire life-cycle passes through nine nymphal
stages and takes more than a year for its completion, the exact time taken b y
each stage could not be determined. The present studies are therefore,
confined to the description of the various nymphs and adult of this insect.
Numerous papers have been published on the biology and life-history of a
number of different orthopteran insects. A few important ones are those of
Carothers (1923), Criddle (1926, et seq.), Cowan (1929), Faure (1935), Chesler
(1938), Hubbell (1940), Burnett (1951), Katiyar (1953, 1955, 1956, 1961),
Richards, T. (1958), Buttiker and Bunzli (1958), Gabbutt (1959), Hunter-Jones
and Ward (1959), Richards, A. (1961), Barnes (1963), Jago (1963) and Eades
(1964). Without describing their morphological characteristics, Ramsay
(1964) has given an account of the number of moults in 229 species and sub-
species of Orthoptera, while Uvarov (1966) has given an account of the post-
embryonic development of Orthoptera in general.

Observations
Habits and burrow-making
The young and adults, for their habitation, make isolated slanting tunnels
which they rarely leave except at night. They prefer to live in moist sandy
places where tunnelling is easy; they do not survive in dry, hard or water-
logged soil. Once they come out of their burrows, they dig fresh ones.
Throughout its lifecycle, the insect is exclusively carnivorous. This was
confirmed after dissecting open the alimentary canal and studying its contents
of the various nymphs and adult. They feed mainly on ground beetles
(Khattar, previous article).
The tunnels are usually dug at an angle of 60 degrees. The diameter and
length of the tunnel increases with each instar. The diameter of the tunnel
varies from 1 to 1¼ inches. The adults can go as deep as two feet while the
earlier nymphs usually remain near the surface. The tunnel is wider at the

Published online 17 Feb 2007


590 Narain K h a t t a r

bottom, and the insect remains there facing the opening. No side tunnels
branching off the main tunnel have ever been observed.
The digging is done at a very rapid rate with the mandibles which, along
with the protruding labrum, function as an efficient shovel. The fore and
middle legs are employed only to give a firm footing to the insect during the
process of digging. The small heap of sand so dug with the jaws is first
pushed backwards b y the under side of' the body and then thrown out with a
force by the large spurs of hind tibiae. As the insect goes deeper, every time
the dug-in sand is brought to the surface on the hind tibial spurs and thrown
out with a force. When the insect has gone 5 to 6 inches deep (the distance
probably measured with antennae, the tips of which keep touching the opening
of the tunnel), the ejection of the sand is stopped. The brought-up sand is
now left to accumulate. This accumulation of sand over the opening is
usually pressed with the triangular outgrowths of the basitarsomeres of the
hind legs to prevent its falling back into the tunnel. The opening of the tunnel
is thus soon closed, marked b y a projecting large loose plug of sand. The
process of digging continues even after the mouth of the burrow is closed,
observable b y a gradual rise of the plug. This characteristic formation of the
sand plug readily identifies the burrow of this insect. Similar formation of
plugs, but of loose earth, has also been noticed in the case of the burrowing
cricket, Gymnogryllus humeralis b y Singh (1951).
On ground, the insect usually runs very fast, often in small jumps.
Possibly because of their narrow posterior ends and membranous character,
it is unlikely that the wings in the adults are ever used for the purpose of flying.

Description of the adult and nymphs


Adult (fig. 1)
The adult is large (about 4 cm long), robustly built, with a striking appear-
ance due to curious fiat expansions on its tarsi and spirally rolled wings.
Large mandibles and exceptionally large legs give the insect a ferocious look.
The general body colour is creamish yellow with black patches dorsally, and
creamish green ventrally. The head is large, whitish with a green tinge in

FI~. 1. Seh~zodaetygus monstrosus, adult;.


Adult and nymphal stages of Schizodactylus monstrosus 591

front, light ereamish laterally, and with brown patches dorsally. The
antennae are very long (each about 9.6 em) and filiform. The scape is white
with a black dot on its inner side, the rest of the antenna is pale brown. The
large pyriform labrum is whitish in eolour and covers the mandibles in front.
The maxillary and labial palps are quite tong and while the insect is resting,
their tips usually rest on the ground. The former extend anteriorly in front
of the head, and the later laterally to the head. The pronotum is large, well-
developed and rectangular in shape. It is pinkish in colour with light and
dark brown patches dorsally. It is tuberculate middorsally and dorsolaterally.
The tegmina and wings are long, tapering posteriorly and roll into a spiral which
lies over the cerei. The two wings of the same side roll over each other. The
sides of the tegmina turn down abruptly and cover the lateral sides of the abdo-
men. The tegmina are brown dorsally, and light brown laterally. The fore, mid
and hind femora are off-white or light pinkish, with brown patches on the
outer latral sides. The fore and mid tibiae are light green in eolour, a pro-
nouneed characteristic of this insect. The hind tibiae arc light pinkish in
eolour. The number and arrangement of spines and spurs present on the
femora and tibiae of the three pairs of legs are shown in figs. 4 and 5, the des-
eription of which has been already given (Khattar and Srivastava 1962). The
claws on the legs are long, curved and brown in eolour with black apices. The
abdominal tergites are off-white or pale in eolour with dark patches on their
midposterior margins. Ventrally, the abdomen is ereamish green, and bulges
slightly downwards. The eerei are long, flexible, unsegmented, and broad
basally tapering into blunt ends. They are white in eolour, slightly curved
medially and beset with numerous sensory hairs (Khattar, 1965).

Nymphs (Figures 2 and 3)


As already stated, the entire life-cycle of the insect takes more than a
year for its completion, passing through nine nymphal instars. Like adults,
all the instars are burrow-makers, nocturnal and carnivorous in habits, and
live in very specific conditions of loose moist sand near water expanses. While
the first nymphs were hatched from the eggs collected from the burrows, the
other nymphs were collected from the fields from time to time. The following
morphological characteristics were taken into consideration to determine the
various nymphal instars, as shown in table 1.
The various nymphs are almost similar in eolour and bodyform to the adult.
In the various nymphal stages, the body is slightly darker than that of the
adult. The nymphs are usually light brownish dorsally. The dark brown
patches on the abdominal tergites are quite well marked in nymphs in contrast
to those of the adults where they become less dark and more scanty. Ventrally,
the abdomen is characteristically ereamish green and bulges downwards in all
the instars. The head is almost white in front, brown laterally and dorsally,
and with dark patches on the vertex. The antennae are always longer than
the body. Their eolour varies from light brown to off-white. It has not been
possible to determine the rate of growth of the antennae and their exact
measurements, because of their tendency to frequently break off. The average
increase, in length of the antennae, in the final ecdysis is about 14 mm or 17 %.
The pronotum is large well-developed and rectangular in shape. It is
usually light brownish in eolour, with dark brown patches and markings on the
592 Narain K h a t t a r

dorsal side. After the fifth instar, the pronotum exhibits tubercles middorsally
and dorsolaterally. The colour of femora varies from off-white to light brown.
The fore and middle tibiae are characteristically light green or light brownish
green. The tarsi and their outgrowths are usually white. No variation has
been observed in the number and arrangement of the spines and spurs present
on the femora and tibiae of different instars and adult, except for the size and
chitinisation of these structures which increase with each instar. As in adult,

~EP
AS
AS.

,AS
I-
,EP
5.

DLS

ASP~ I
Ep~ ~\ ~t "

Ci-~ 3"
FIG. 2. Last nymphal instar, dorsal view.

the second and third tarsal segments of all the three pairs of legs in all the
nymphs are flexible, and possess characteristic outgrowths, the euplantulae.
The basitarsomeres of the hind legs, in all the instars, possess large triangular
fixed plates which increase in size and become more chitinised in each instar.
B u t for the difference in size, the cerci in the various nymphal instars are
similar to those in the adult.
Developmental changes in the tegmina and wings are gradual and distinct
as described below (fig. 3):
First nymph: Tegmina and wing rudiments are present as small, and
indistinctly differentiated triangular lobes on the lateral sides of the meso-
and meta-thorax.
Table 1
Increase in g r o w t h (in ram) of different structures in various n y m p h a l instars a n d adult of Schizodactylus monstrosus

Structure First Second Third Fourth Fifth Sixth Seventh Eighth :Ninth Adult
Nymph Nymph Nymph Nymph Nymph Nymph Nymph Nymph Nymph

~10-25 11'25C:
Head width 2.75 3.50 4.25 5.30 6.75 7.25 8.60 9.10 ( 11.00
12'00~
Pronotum width 1.05 1.50 1.90 2.50 3.25 3.60 4.20 4.60 5.00 5-75
9~
Fore f e m u r 2.80 3.75 4-80 5.30 7-25 7-80 9.00 9.30 11.00 12.25
Fore tibia 2.25 3.50 4.80 6.30 7.50 8.20 9.75 10.00 11.20 13.50
Fore t a r s u s 1.25 1.80 2.40 2.90 3,60 4-00 4.40 4.80 5-10 6"00 O
Mid f e m u r 3.00 4.25 5.60 7.00 8.50 9.00 10.50 11.00 12.20 14-30 O
Mid tibia 3,20 4.00 4-90 6.80 8.40 8.60 10.00 10.25 10.75 13'75
Mid t a r s u s 0.80 1.25 1.75 2.40 2'80 3'10 3'90 4.00 4.20 4'75
Hind femur 5.25 6.75 8.75 10.25 12.75 13.50 16.00 16.65 18.30 22.00
H i n d tibia 3.80 5.25 6.80 8.30 10.00 10.25 13.00 13.25 14.20 17.00
Hind tarsas 1.30 2.00 2.75 3.80 4.60 4'90 5.50 5.75 6-30 7"80
Cercus 1.10 1.40 2.60 3.00 4-00 4.25 5.25 5-50 5.85 9'00
B o d y length 7.25 11.25 14.30 17.00 23.00 25.00 27.25 30.00 40.00 42.00
594 Narain Kha%tar

Second nymph: The small triangular tegmina and wing rudiments become
separated from the meso- and meta-thoracic segments by distinct ridges. These
lobes point downwards and slightly anteriorly.
Third nymph." There is a slight increase in the size of the tegmina and wing
rudiments, but they remain as triangular lateral flaps of the meso- and meta-
thorax separated by distinct ridges.
MT MTT

TP WP TP WP TP WP TP WP

I II Ill IV
I AS

TP WP TP WP TP WP
v VI VII

TP

" WP

VIII IX wP
Fin. 3. :Developmen~ of pronotum, tegmina and wings in the successive instars (I-IX).

Fourth nymph: The tegmina and wing rudiments become large and fleshy
and remain laterally downwards on the sides of the meso- and meta-thorax.
Traces of longitudinal striations are visible on both the rudiments.
Fifth nymph : Tegmina and wing rudiments enlarge into fleshy, conical pads,
and are directed laterally downwards. Longitudinal striations become more
prominent. Small anal areas become distinct and are partly separated on both
the rudiments.
Sixth nymph: The fleshy rudiments increase further. TegminM rudiments
are directed slightly backwards while those of the wings point laterally down-
wards. The ridges separating the anal areas become more distinct.
Seventh nymph: The tegminal and wing pads become reflected over the
back. The wing pads extend up to the posterior margin of the first abdominal
segment while those of the tegmina reach only up to the posterior margin
of the metathoracic segment. The wing rudiments overlap the basal
posterior margin of the tegminal rudiments.
Adult and nymphal stages ofSchizodactylus monstrosus 595

Y$

C° Do
FI•. 4. A. F o r e t i b i a a n d t a r s u s of a d u l t , d o r s a l view. B. T h e s a m e in v e n t r a l view. C. Mid
t i b i a a n d t a r s u s of a d u l t , d o r s a l view . D. The s a m e in v e n t r a l vi e w .

A. B.
b'~c~. 5. A. H i n d t i b i a a n d t a r s u s of a d u l t , d o r s a l vi e w . B. The s a m e i n v e n t r a l v i e w
596 Narain Khattar

Eighth nymph: Tegminal and wing pads are reflected over the back and
become slightly larger. The tegminal rudiments extend up to the anterior
margin of the first abdominal segment while those of the wings extend beyond
the anterior margin of the second abdominal segment. The tracheation is
more distinct on both the rudiments.
Ninth nymph: The tegminal and wing pads become quite large and cover
the dorsal and lateral sides of the first four abdominal segments. The wing
pads now extend beyond the anterior margin of the fifth abdominal segment
while those of the tegmina extend up to the posterior margin of the fourth
abdominal segment. The wing pads overlap the tegminal pads. Both the
pads become darker in shade dorsally. The tracheation, too, is more distinct.
A description of this tracheation has already been given by Raage (1957).

Discussion
Dyar (1890) was the first investigator to give a formula on the basis of
which the various stages of lepidopterus larvae could be determined. According
to him, the head capsule of these larvae grows in geometrical progression,
increasing in width at each ecdysis by ratio of usually about 1.4, which is
constant for a species. Subsequently, other investigators found this formula
applicable in the case of other insects also. Further, it was also confirmed t h a t
this formula could as well be applied to structures other than the head capsule.
The progression ratio is rarely regular, but the difference is so marginal that it
does not affect the determination of nymphal instar (Richards, 1961).
Again this ratio may not be constant for all the different structures of the
body in the same species. While some of the structures m a y give a reliable
data for determining the instars, the others m a y not. The different structures
usually taken into consideration for determining the nymphal instars among
Orthoptera are, the length of the body, the antennae, the legs, the genitalia,
the pronotum, the tegmina and wings, the number of eye stripes, and a few
less important morphological featm'es such as colouration, etc. In a series
of papers, Criddle (1926, et seq.) took an account of the number of antennal
segments, development of wings, modification of pronotum and length of
hind femora in determining the hopper instars of a few members of Orthoptera.
Mukerji and Batra (1938) and Roonwal (1940, 1946) took into consideration only
the number of eye stripes in determining the nymphs of some Acridids.
In another paper, Roonwal (1952) gives importance to variation in the
number of the antennal segments during the postembryonic development of
Heiroglyphus nigrorepletus and some other Acridids. Jago (1963) has shown in
locusts that while the number of eye stripes is a fairly reliable means for
determining the different instars, the antennae are not. In another investiga-
tion, Burnett (1951) while taking into account the number of eye stripes in the
postembryonic development of Nomadacris septemfasciata states that this
number is not perfectly correlated to the number of instars. Earlier, Chesler
(1938), gave systematic description of the identification of the various nymphs
of Aerididae on the basis of the length of body, hind femora, antennae and wing
rudiments. Cowan (1929) took into account the development of the ovipositor
in females and the subgenital plate in the males as nymph-determining
characters in the Mormon cricket, Anabrus simplex.
Adult and nymphal stages of Schizodactylus monstrosus 597

Similarly, l~ichards (1961), in addition to the development of ovipositor,


takes into account a few other morphological structures such as the length of
the various parts of the legs, the body and cerci, as determining factors.
Clarke (1957) gave emphasis to the body size only, stating, however, that linear
size increases during a stadium as well as at an ecdysis. Coleman (1911),
Coleman and Kannan (1911), Karandikar (1945) and Albrecht (1955) have
shown that the development of tegmina and wings is of special value in deter-
mining the hopper instars.
Details of the burrowing habits of Schizodactylus monstrosus have already
been given in the present studies. Earlier, Ramme (1931) had incorrectly
assumed that the flexible spurs (called paws by him) of the hind tarsi of this
insect help in digging. Later, Carpentier (1953), however, correctly observed
that the digging is done solely with the jaws. Carpentier (1953) also observed
that occasionally the adult Schizodactylus live in open burrows and in such
cases the insects themselves leave the burrows unplugged. The present
investigator too, has come across some such instances, but has not been able
to determine the reason for their preference to live in open burrows. Regarding
the depth of the burrow of the adult insect, Carpentier (1953) has said that it
would depend on what depth the insect reaches water. B u t during all these
years, the present author has not been able to find a single instance of a burrow
going down to the level of water. However, there appears to be a distinct
correlation between the depth of the burrow and the condition of the soil--the
drier the sand, the deeper the burrow and vice versa. The insects are thus
very sensitive to dryness and usually die if kept in a glass jar without moist
sand. They need high moisture content, cool temperature and specific
conditions of sand for their survival. Further, though these insects try to
hide themselves away from direct sunlight, yet, as also observed b y Carpentier
(1953), they are attracted towards night light.
The well developed wings (always longer than the body) of the adult
Schizodactylus are peculiarly rolled up in a spiral at rest. Except for a single
reference b y Carpentier (1953), there is no other account available regarding the
flight-capacity of these insects. On the basis of a personal communication to
this effect from Dr. S. D. B. Agarwal, Carpentier (1953) reported that these
insects are able to fly at night and the flight is rectilinear. The present author,
however, has not seen these insects flying. While experimenting with the
adult insects, in the laboratory and outside, the author has been able to observe
a few facts. While jumping from a height of one foot or s% the insect opens
its wings--apparently for a soft landing. The wings roll back as the insect
lands. When the insect was dropped from heights of 10 to 15 feet, both the
wings opened at right angles to the body axis. At certain times, the wings
would even flutter but were of not much use in either changing the direction
or carrying the insect forward or even for a soft landing. Possibly the
reason for this lack of flying-ability in Schizodactylus is that not only the wings
narrow down abruptly but, also, they are too membranous and delicate to
provide adequate support for the exceptionally heavy body.

Summary
1. The life-cycle of Schizodactylus monstrosus takes more than a year for its
completion and passes through nine nymphal instars.
598 Narain Khattar

2. Like adults, all the nymphs make solitary burrows for their habitation.
They live in very specific conditions of loose moist sand near water expanses.
The tunnels are dug solely with the jaws, at 60-degree angle to the ground, and
with no side tunnels. The depth of the tunnel correspondingly increases
with each instar, and is about 2 feet in the case of adult insect. The diameter
of the tunnel varies from ~ to 11 inches. CharacteristieMly, the nymphs and
adults close the openings of their burrows with projecting plugs of sand.
3. Like adults, all the nymphs are carnivorous and nocturnal in habt%
and shun direct sunlight.
4. The adult insect is large, robustly built and looks ferocious. Save for the
difference in size all the nymphs are almost similar in form and colouration to
the adult, with no different morphological features. The fore and mid tibiae
are characteristically green in colour in all the nymphs and adult.
5. The wings in adult are rolled posteriorly in a peculiar spiral lying over
the cerei. They can hardly be used for the purpose of flight.

Acknowledgments
The present author is greatly indebted to Dr. P. N. Mathur, Principal,
Government College, Ajmer, for his guidance and encouragement. Sincere
thanks are also due to Dr. Tej Singh, Professor of Zoology, Government
College, Ajmer, for providing the research facilities, and the University Grants
Commission, New Delhi, for awarding financial assistance to carry out this work.

Abbreviations
ABD: Abdomen.
ANT: Antenna.
AS: Apical spines.
ASP: Apical spurs.
CE: Cereus.
DLS: DorsolaterM spines.
EP: Euplantulae.
FE: Femm'.
I/f: Head.
LS : Lateral spines.
MT: Mesothorax.
MTT: Metathorax.
PT: Prothorax.
SAS: Subapieal spines.
TI: Tibia.
TP: TegminM pad.
TPL: Triangular plate.
UNG: Unguitraetor.
VAS: Ventroapical spines.
VS: Ventral spines.
WP: Wing pad.
1,2,3,4: First, Second, Third and Fourth tarsal segments.
1AS: First abdominal segment.

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