The external morphology of the Mormon cricket (Anabrus simplex Haldeman) with special reference
to exoskeletal factors involved in the penetration of sodium arsenate dust
by Leo W Tannenbaum
A THESIS Submitted to the Graduate Committee in partial fulfillment of the requirements for the
degree of Master of Science in Entomology
Montana State University
© Copyright by Leo W Tannenbaum (1941)
Abstract:
The external morphology of the Mormon cricket was studied In Its entirety In order that exoskeletal
factors Involved In sodium arsenlte penetration might he better understood. Of special Interest were
thorax modifications. The mesothoracic spiracle is unusual because it consists of two separate openings
leading into individual tracheae. Twenty-three figures of the Insect1S anatonyr sure here included.
After trying some thirty plastics and paraffins a paraffin melting at 38-40°C was used to localize
sodium arsenlte penetration. A simple test for cricket normalcy was devised. It was noted that crickets
go through definite stages in their ability to react to simple stimuli after the sodium arsenlte
applications have begun to penetrate. Mne lots of the Mormon cricket were dusted on different parts of
the body to determine if preferential penetration exists. Applications to membranous areas which are
not too smooth resulted in earlier reactions and slightly earlier 50% mortality points. Wide portions of
sclerotlzed areas were not far enough behind in this respect to be significant. It was concluded that
other factors besides degree of eclerotiration, such as position in respect to major organs, were of great
importance and would modify the results. Analysis indicated that time of death and amount of
penetration of sodium arsenite are not closely related. IHB SXTBEKAl MOEPHOLOGY 07 THB MOEMON CMCKBT
(Anabrus simplex Haldeman)
WITH SPBCIAL HBTBEBNCB TO BZOSKELETAL FACTORS INVOLVED IN
THE PENETRATION OF SODIUM ABSBNITB DUST
By
LEO W TANNENBAUM
A THESIS
Submitted to the Graduate Committee
in
p a r tia l fulfillm ent of the requirements
for the degree of
Master of Science in Entomology
at
Montana S tate College
Approved:
In Charge of Major Work
Chairman, Examining Committee
ythairm an, Grathiate Committee
Bozeman, Montana
Jbne1 IgUl
H31%
T /f* -
2-
TABLE OF CONTENTS
Page
A b stract ....................................... ................. ......................................... ..
U
vi vi vn
In trod u ction ..................................................................... ..............................
S e c tio n I , E xternal Morphology ...........................
Review o f L ite ra tu r e on Morphology
Head .......................................... ................. •
Thorax ..........................
Abdomen........................ ........................................................ ..
16
28
A b b reviation s and l e t t e r in g on P la te F igu res ......... ...................
32
P la te s ..........................................
35
S e c tio n I I , The P en etra tio n o f Sodium A r se n ite
through th e e x o sk ele to n ........................
Review o f L ite ra tu r e on P en etra tio n o f
Sodium A r se n ite ................................................
Procedure ...................
W
lj.0
U2
E ffe c t o f D usting th e Abdomen w ith Sodium A r se n ite . . . . . . . .
U9
E ffe c t o f D usting th e Thorax and i t s Appendages
57
................
General C on sid eration s ..........................
61
C onclusions .................................
614-
L ite ra tu r e C ited and C o n s u lt e d .................
66
66124
ABSTRACT
The external morphology o f the Mormon cricket was studied in
i t s en tirety in order th at exoskeletal facto rs involved in sodium arsenite
penetration might he b e tte r understood. Of special in te re st were thorax
modifications. The mesothoracic spiracle i s unusual because i t consists
of two separate openings leading into individual tracheae. Twenty-three
figures of the In sec t1S anatonyr acre here Included,
After try in g some th irty p la stic s and p araffin s a p araffin melting
at 38-40*0. was used to lo calize sodium arsen ite penetration. A simple te s t
for cricket normalcy was devised. I t was noted that crick ets go through
d efin ite stages in th e ir a b ility to react to simple stimuli a f te r the sodium
arsen ite applications have begun to penetrate. Uine lo ts of the Mormon
cricket were dusted on d ifferen t p a rts of the body to determine i f
p referen tia l penetration e x ists. Applications to membranous areas which
are not too smooth resu lted in e a rlie r reactions and slig h tly e a rlie r
50# m ortality points, fid e portions of sclerotized areas were not fa r
enough behind in th is respect to be sig n ific an t. I t was concluded that
other factors besides degree of s c le ro tiration, such as position in respect
to major organs, were of great importance and would modify the re su lts.
Analysis indicated th at time of death and amount of penetration of sodium
arsen ite are not closely related .
-UINTEODUCTIOE .
The Mormon cricket, Anabrus simplex (Eald1) , is an important
pest of several ag ricu ltu ral and range crops in Montana and certain other
Western states#
Control in recent years has consisted, fo r the most
p a rt, in the use of sodium atsen ite as a dust.
of general use throughout those sta te s infested.
This in secticid e has been
Although i t is known to
be a powerful poison l i t t l e i s known eith er of i t s method of penetration
or of the exoskeletal factors involved.
Since penetration does take
place through the exoskeleton, the outer p arts of the insect influence
i t and may well be factors regulating the efficiency of control.
paper has a twofold purpose:
This
(I) a study of the external morphology of
the Mormon cricket so that the exoskeletal areas may be b e tte r understood,
and (2) experimental determination of the b a rrie r the major exoskeletal
areas offer to penetration of sodium arsen ite dust.
To carry out the f i r s t purpose a complete study of the external
morphology was made.
To carry out the second purpose, experiments were
conducted to determine or gather information on the following points:
(I) the possible existence of p a rts of the cricket more susceptible to
penetration than other p a rts, and (2) the rela tio n of amount of penetration
to the time necessary fo r f if ty per cent m ortality.
The author i s greatly indebted for suggestions and criticism to
Dr. H. 3. M ills under whom th is study was conducted. Mr. Ellsworth
Hastings has taken a kind in te re s t in chemical aspects of the work, and
Dr. G. C. Crampton, of the Massachusetts State College, has offered
valued criticism in i t s morphological aspects. Mr. D. J. Pletsch has been
most helpful in matters dealing with illu s tr a tio n s . All these kind
services are greatly appreciated by the author.
-5 -
Section-I
EXfSHNAL MOHPHOLOGFr
No complete morphological study of a decticine Orthopteraa is
available in the lite r a tu r e .
Snodgrass (I909), in discussing the thoracic
s c le rite s of the various Orders, illu s tr a te d three views of the meso thorax
of Anabrus simplex.
The same author (1957) in considering the male g e n ita lia
of Orthopteroid in sects discussed the male term !naila of th is species.
Insects in rela ted groups have been trea ted in greater d e ta il.
(1930) made a d etailed study of the head of Stenopelmatus.
Crampton
The same author
in a series of papers (1915-1938) paid special atten tio n to the phylogenetic
relationships of the O rylloblattidae and other Orthopteroide, and studied
in a comparative fashion the head, mouthparts, and thoracic s c le rite s in
Orthoptera and rela ted orders.
Walker in a series of papers (I919-I933)
dealt with subjects in the same general manner.
Of especial in te re s t to
th is study are th is author's papers (1919b, 1922) on the terminal stru ctu res'
of Orthopteroid in se cts.
mouthparts of Orthoptera.
Tuasa (I920) studied the anatomy of the head and
Oumey (1936) made a complete study o f
Ceuthophilus and i t s phylogenetic position.
Train
The head of Anabrus i s sim ilar in many respects to the heads of
Stenopelmatus and Ceuthophilus.
The marked, broad dorsal roundness and
the smoothness of Stenopelmatus are lacking.
These differences may be
rela ted to the differences in habit since Anabrus does not burrow as does
Stenopelmatus.
In i t s more general c h a ra cte ristic s the head o f in .tru s
is more lik e th at of Ceuthophilus since antennae and eyes are more ce n trally
_6_
as well as more dorsally located.
These c h a ra cte ristic s may he taken for
evidence that Anahms has departed further from the more prim itive type
than has Stenopelmatus.
Head capsule:
When seen from in front as shown in fig . I , the
head capsule of Anahrus i s ohlong-ovoid in shape.
The coronal suture Cs
which represents the stem of the Y—shaped epicranial suture i s p lain ly hut
not strongly demarked.
I t extends cen trally terminating a t a point near
the d is ta l end of a prominent raised process, the fastigium, fas.
The
l a tte r i s homologous with the process.occurring in Llcodla, Paranahrus,
Ceuthophilus and other genera,
fro n tal sutures, which represent the arms
of the T-shaped epicranial suture, are not present in Anahrus.
Immediately
helow the fastigium, fa s, and almost a t rig h t an&les to the lin e of the
coronal suture, Cs, are suture—lik e folds caused hy the close proximity of
antennal sockets and fastigium, fas.
On each side of the coronal suture
p a rie ta ls p ar.
Cs
are the regions known as ••
The areas above and behind the compound eye e are the
temples or teopora te .
cheeks or genae ge.
The regions helow and behind the eyes are the
Above the compound eyes e and extending toward the
p o stoccipital suture pos o f fig . 2 is a suture here considered to he
temporal suture te s .
Because there are no fro n tal sutures the frons f
may he considered to be the area below and between the antennae extending
down to the frontoclypeal suture fee.
Beneath the gena ge and separated
from i t hy the eplstomal sdture eti i s the eubgena or hasimandihulare hm.
The l a t t e r i s separated from the mandible md hy the baeimandibular
membrane bmm.
In the Mormon crick et each epistomal suture es extends
-7 -
mesad meeting i t s fellow on the other side.
The portions of the eplstomal
sutures es between the frons f and the clypeus c comprise the frontoclypeal
suture fcs.
Zn Anabrus coincident with the sutures labelled es are the
fro n tal p its
the roots of invaginations forming the pretentorium g t of
fig . 5» Anterior to the frontoclypeal suture fos i s the clypeus c.
I f the head capsule be viewed from the rear as in fig . 2 the
postgenal suture pgs may be seen to separate the genae ge from the postgenae pge.
The postgenal suture pgs leads upward from the subgena bm and
i s lo s t or discontinued in the occiput.
or buttressed areas.
In Anabrus i t meets two thickened
The lower one lab elled bu i s an easily seen thickened
area between the postgenal ridge pgs and the postocciptal suture pos.
Slightly above th is area and running through the terminus of the postgenal
suture pgs i s the temporal suture tea.
Eyes:
The compound eyes e are not very large but are re la tiv e ly
larg er than, the eyes of Stenopelmatus and CeuthophiIus.
This may be in lin e
with the suggestion of Crastpton (1930) th at there seems to be a marked
correlation of Insects having both small eyes and long se n sitiv e.antennae
in insects with hiding h ab its.
Anabrus with le ss marked hiding habits than
the two above mentioned genera has larg er eyes and re la tiv e ly shorter
antennae than Ceuthophllus.
The o c e lli in Anabrus are v e stig ia l and may be non-functional.
The median ocellus i s present as a lig h t spot below the fastiglum fa s .
The la te r a l o ce lli are present as sim ilar spots a t about the widest point
of the fastiglum as shown in fig . I .
Occipital region:
The posto ccip ital suture pos of fig . 2
separates the postgena pge from the postocciput go.
The postocciput go
-g "bears a p air of processes, the o ccip ital condyles occ upon which the la te r a l
cervical p la tes are a rtic u lated .
The o ccip ital condyles in Anahrus are small
s c le rite s which are folded on th e ir long axes.
They are separated from the
postocciput merely hy grooves. P o sterio r to the suhmentum sm and the post­
occiput £0 i s the membrane of the cervix.
I f th is membrane be removed a
posterior view would be had of the p o sterio r te n to ria l arms poc and the
occip ital foremen ocf of which an an terio r view i s had in fig . 5»
Tentorium:
capsule.
The tentorium i s the endosk eletal portion of the head
I t i s very sim ilar to the type seen in the more generalized
Orthopterold in se cts.
The tentorium i s formed by invaginations of the head
capsule. ‘ I f the labrum, clypeus, and the an terio r portion of the head
capsule be removed from a IO^ KOH preparation the p arts of the tentorium
may be seen as in f ig . 5« The an terio r arms or pretentorium jrt extend
backward from the fro n tal p its f£ of f ig I as invaginations.
The dorsal ,
arms or supratentorium st^ are of a th in and tendonous nature and ris e
an terio r to the body o f the tentorium or eutentorium eu.
Immediately
an terio r to the body of the tentorium eu i s the neural foramen n f.
Through
the neural foramen pass Commissures connecting the brain and the
suboesophageal ganglion.
Extending forward from the gular p its g) seen
in fig . 2 as invaginations are the p o sterio r arms of the tentorium or
parocclput poc.
The p o sterio r arch £ i s also a re su lt of the invagination.
The body of the tentorium eu may be considered as a fusion o f an terio r and
posterio r te n to ria l arms.
In fig . 5 also, may be seen an ental view of the
rear portion of the head capsule.
Besides the added strength given to th is
portion by a thick parocclput poc there is fu rth er b uttressing bu above the
parocclput.
The postgenal process ££ i s also thickened in th a t area within
-9 -
the dotted lin e In the figure.
In the postgenal process
I s the p o st-
genal acetabulum pga a socket fo r the hypo condyle h of the mandible seen
in fig . It. Along the lower edge of the postgenal process i s a thickened
marginal area, the parastomlua p s t.
HMD APPENDAGES
Antennae:
an of fig . 3.
About the base of the antenna is a ring, the antennale
In Stenopelmatus and Ceuthophilus the antennale bears a
median projection fo r piv o tal purposes, the antennlfer.
antennlfer i s lacking.
In Anabrus the
The basiscape of Stenopelmatus and other in sects i s
also lacking in Anabrus.
However, there i s a small "inpushed" tendon-
connection (in the membrane immediately an terio r to the f i r s t antennal
segment) which helps in moving the antennae.
The antennae are long and
thread-like, in most cases slig h tly longer than the body.
The number of antennal
segments i s v a ria b le,in some specimens reaching more than one hundred and
fo rty .
The f i r s t antennal segment o r scape sc i s broad and flatten ed .
Basally i t has projections which aid in pivoting the antennae.
The segment
next to the scape, the pedicel gd, i s more cy lin d rical and i s followed by
the postpedicel ppd.
The postpedicel and a ll other segments d is ta l to i t
are known co llectiv ely as the flagellum.
R a g e lla r segments may be divided
into the short segments or brachymeres, and long segments or dollchomeres.
'
In Anabrus long brachymeres lb , short brachymere eb, and slender dollchomeres
-
-
...
-
-
sd, may be easily distinguished.
Clypeus and labrumt
,
.
-
-
,
.
Intermediate types are also present.
In many in sects the clypeus may be divided
by differences such as coloring into a p o sterio r postclypeus and an an terio r
-1 0 -
anteclypeus.
This division i s apparent only fa in tly in Anabrus,
I f the
lahrom la and clypeus c are inspected from an Inner or hoccal view as in f i g .7
the epipharynx may he seen.
hy fine sensory h a irs.
The greater p a rt of the epipharynx i s covered
Two main divisions may he distinguished, an an terio r
preepipharynx pre and a p o sterio r post epipharynx poe.
correspond to the lahrom and clypeus respectively.
has several small s c le rite s .
These two areas
The epipharyngeal area
The tormae to are a p air of small s c le rite s
situated near the lahro-clypeal la te r a l angles, and thus delim it the lahrum
and clypeus.
Between the lahrum and clypeus and extending transversely
between the tormae i s a small s c le rite , the lntertorm a i t .
Mandihle: Beneath the hasimandlhular membranes bmm of fig . I
are the paired jaws or mandibles md.
The mandible is roughly pyramidal in
shape and the space within i t i s f ille d with tissu e.
and there are three faces.
In fig . 4, a p o sterio r view of the dextral
mandi ble, the re a r face may he seen.
Bie apex of the roughly pyramidal
mandible is the tearing area or gnathapex ga.
mola m is proximad of the gnathapex.
the b itin g edge near the base.
points.
The base i s triangular
The grinding portion, or
A h ru stla or brush hr i s present on
The base of the mandible has two a rtic u lato ry
The p o sterio r artic u lato ry point, the gn&thocondyle or hypo-
condyle h of fig . 4 .f i t s into the postgenal acetabulum pga of the poetgenal
process ££ shown in fig . 5» A nterior a rtic u la tio n for the mandible is
afforded by means of the ginglymus g i.
The glnglymus (shown in fig s . I
and 4) f i t s into a groove in the side of the clypeus.
To the extensor
prominence go (on the outer edge of the mandible near the hypocondyle h)
is joined the extensor tendon et which helps open the mandible.
Near the
-li­
medl an ridge of the mandible, a t I ta base, la the tendon, e c le rite or
gnathite Qi to which ia attached a large flexor tendon f t .
In Anabrns both
the gnathite gn and flexor tendon f t are slig h tly modified from th e ir
homologs in Centhophilns and Stenopelmatns.
The gnathite gn i s not readily
distinguishable as a separate a c le rite and i s fused with the flexor tendon f t .
The flexor tendon i s very.large since i t ia in closing the jaws during chewing
that strength i s most needed.
At i t s base i t extends more than h alf the
distance to the gnathocondyle h.
Extending inwardly, the flexor tendon f t
i s composed of one large p o sterio r portion closely apposed to the base of
three additional narrow branches.
The endognath eg i s an in tern al; shblf at
the base of the mandible.
Maxilla: In fig . 2 there may be seen a p o sterio r view of the
s in is tra ! m axilla.
The labium and hypopharyngeal regions are connected to
the maxilla by means of the basim axillary membrane b.
The basal segment or
cardo of the m axilla is divided by the cardinal suture or cardosuture cds
into a basal basicardo be, and a d is ta l distlcardo do.
D istal to the car do
are the stipes £ and the parastlpes ps which are separated by the p a ra s tip lta l
suture pss.
When the p o sterio r side of the car do and stip es are viewed
entally as in fig . 6 an in tern al ridge, the endocardo ec corresponds to the
cardosuture cds, and another in te rn a l ridge, the endostip es eet corresponds
to the external p a ra s tip lta l suture gss of fig . 2.
The in tern al ridges
o ffer additional support to the maxilla and afford places of attachment fo r
muscles.
The endo stip es est affords attachment fo r the flexor muscles of
the maxilla.
The distlcardo dc i s enlarged by an extruding bulge which
supplies space for_rauscles.
A process not seen externally (unless the
-1 2 -
maxilla Is pulled out) the cardoprocess
car dotendon ctn.
o ffers attachment to the
The extensor muscles act In conjunction with the la tte r
to open the maxilla by pulling i t away, la te ra lly , from the mesal p a rt of
the head.
The condyle acting in th is operation i s the artic u lato ry process
of the basicardo d which pivots externally on a process of the parocciput
poo.
In Anabrus a division of the stip es into basal and d is ta l regions is
not perceptible,
A small region the p alp ife r g f is clearly seen.
the five-segmented maxillary palpus mg.
I t bears
The f i f t h segment of the maxillary
palpus bears a rath er large sen sitiv e area a t i t s tip .
d is ta lly an outer galea and an inner la c in ia I .
The stip es s bears
The galea i s divided
d istin c tly into a b aslgalea bg and a d is tigalea dg.
In Anabrus the baslgalea
is not much wider than the d istig a le a a t th e ir juncture and the l a t t e r is
blunt, not slender and graceful.
In fig . 9» an an terio r or mesal view of
p art of the maxilla, five sensory areas may be seen on the galea.
A large
area a t the tip of the d istig a le a i s obvious; basad of th is and indicated
by fine lin e s are four le ss easily distinguished sensory areas the function
of which is obscure.
At the d is ta l end of the la c in ia I are two tooth-
lik e projections the laciniadentes Id.
A th ird process the lacin u la I i i
i s joined to the la c in ia by means of a socket which can be b e tte r seen
from the an terio r view.
border.
The la c in ia bears a fringe of setae on i t s mesal
In Anabrus there is present also, on the an terio r face of the
la c in ia , a process with probable sensory function which is sim ilar in
appearance to those on the galea.
Labium;
The labium may be seen in fig . 2.
the jparaglossae pgl are attached to the gloss!ger gg.
The glossae g l and
The portion of the
-1 3 -
labium immediately "baead of the gloeaiger gg is the la b ia l s tip ite s 11_.
In Anabrus demarcation of the gloseiger gg from the paragloeeae pgl and
gloesa gl is more easily distinguished on the p o sterio r than on the an terio r
face of the labium.
On the an terio r face as shown in fig . 8, the lin e of
division i s not perceptible.
basad of the glossa g l.
A basal area the basigloesa bgl may be seen
Sutural division between the la b ia l stip e ts li_ and
glossiger gg may be seen as in fig . 2.
The palpiger gg is la te r a l of the s c le rite s 1JL_ and bears the threesegmented la b ia l palpus l£ .
Orampton (1930) has discussed the theories on
homologies of la b ia l and maxillary p a rts mentioned above.
11 and the menturn mn are separated by a membranous area.
d istin c t in Anabrus.
The la b ia l s tip ite s
The rnentua mn is
I t has a notched appearance d ls ta lly .
are attached in te rn a lly muscles which move the labium.
To the mentum
In Anabrus latero —
basal to the mentum are the an terio r arms of the submenturn asm (an terio r
arms of postmentum of Snodgrass) which extend back to the main body of the
submenturn sm.
The large membranous area between the anerior submental arms
and between the mentum and submentum i s the submental mmnbrane.
degrees of membranization.
It
two
The portion between the an terio r arms behind
the dotted lin e shown in fig . 2 i s slig&tly darker and harder than that
portion immediately an terio r to i t .
In insects of th is rath er prim itive
hypopnathous type the gular region i s generally considered to be the rear
portion of the p la te lab elled sm, or altern ativ ely , th at i t has not yet
developed as in the more advanced prognathous type.
situated near the base of the p la te lab elled sm.
The gular p its gjj are
They are the openings to
invaginations which form, the p o sterio r arms of the tentorium poc of fig . 5.
There i s no poetgular p la te as there i s in Stenopelmatus.
Hypopharynx: t The hypopharynx h£ of fig . 10 i s a fleshy tonguelik e structure found in the median ventral p art of the head "between the lahrum
and lahimn.
I t i s very much lik e the hypopharynx of Stenopelmatus.
fig . 10 the hypopharynx h£ is seen- to he composed of several p a rts .
d is tilingua dl i s the region a t the free end.
In
The
I t is sclerotized fo r the most
p art hut portions are very thin as shown in fig . 10.
The hasal region of the
hypopharynx is composed of a dorsal dorsollngua or surlingua si immediately
anterior to the mouth and a ventral membranous basllingua h i.
The salivary
.
duct opens into the hasal hasillngua above the labium.
I n Anabrus on the
side walls of the diet i lingua dl a p a ir of p la te s afford attachment fo r the
hypopharyngeal re tra c to r muscles.
A slender s c le rite the lingualora 11 i s : ,
in the hasal portion of the hypopharynx as i s also the postlingua p i.
-1 5 -
CEEVIX
The cervix Ie the membranous neck region Immediately behind the
head and immediately an terio r to the pro thorax.
In Anabrus the cervical
area Is much larg er than i t appears since normally i t i s p a rtia lly
concealed beneath the an terio r border of the pronotum.
the cervix i s shown to have three p a irs of p la te s.
e c le rite Ic of fig . 13 i s the latero cerv icale.
fellow being on the other side.
In fig s. 13 and Ig
The large la te r a l
I t is one of a p a ir, i t s
The an terio r end of the latero cerv icale Ic
a rtic u la te s with the o cc ip ital condyle occ of fig . 2.
seen in fig . 18 are the ventral cervical s c le rite s vc.
Ventrad o f these as
Dorsad of the
la te r a l cervicales are a p a ir of small dorsal cervical s c le rite s dcs
connected so closely mesally as to suggest th at they are a single central
s c le rite .
-1 6 thorax
The thorax of the Dectlclnae has been l i t t l e Investigated.
Snodgrass (1909) drew three views of the mesothorax of Anabrus.
Most
notable In the anatomy of Anabrus I s th at elongation of the pronotum has
been accompanied not only by prothoracic modification but also by
modification of the p tero thorax as well.
In Anabrus the tegmina have been
so shortened th at in the male they reach only to the middle of the f i r s t
abdominal te rg ite and in the female they are even shorter.
both sexes are represented only by membranous pouches.
The wings in
The thoracic
epiracular apparatus o ffers an in te re stin g modification which may be seen
in certain other in sects of the subfamily.
PROTHORAX
Pronotum:
The pronotum pn of fig . 13 extends back from the
membrane o f the cervix to a point above the middle of the metathorax.
is free dorsally from a point an terio r to the meso thoracic te r gum.
It
The pron­
otum jjn covers the epi sternum eps^and the epimeron epm^ so th at they are
only p a r tia lly v isib le .
In th is study specimens of Anabrus simplex
examined from Montana and eastern Idaho, the pronotum was found to be very
smooth and carinae and sulci were only very fa in tly indicated.
Gurney (1939)
stated that carinae are rare in the genus except in Anabrus loagipes but
suggested the p o ssib ility of longipes being a subspecies of simplex.
As
in other Orthoptero id s, Anabrus has no prothoracic spiracle.
Propleuron:
The f i r s t portion of the propleuron I s a narrow
border-like s c le r ite , the prepectus ppc, which extends v entrally and
-1 7 -
contlnuously through the stern al area. . The dorsal terminus o f the prepectus
ppe Is coincident with th at point a t shich the eplsternum eps^ls overgrown
hy the prono turn pn> The epl sternum eps^ls closely un ited with the pro no turn.
I f the prono turn i s removed the epl sternum i s seen to he almost triangular
in shape and to extend almost to the dorsum.
At i t s hase the epl sternum is
continuous with the precoxale prcx^. The pleu ral ridge p lr i s a mesal .
inflexion of the epl sternum extending along i t s posterior edge.
At i t s
lower portion the pleu ral ridge i s represented externally hy a suture, the
pleural suture p is which separates the episternum eps from the epimeron
epm^.
In the generalised thoracic segment the pleu ral ridge hears an in te rn a l
arm or apophysis which i s closely associated with the ventral apophysis
(furca).
The pleu ral apophysis i s absent in the pro thorax o f Anahrus.
Unlike most Orthopteroids the epimeron epm^ o f Anahrus i s a short s c le rite .
The major portion of the epimeron has been lo s t and as may he seen in fig . 13
only enough of the epimeron ex ists fo r the pronoturn to overlap.
The short­
ness of the epimeron i s a useful adaptation since i t allows fo r more flexible
movement of the prono turn. At the basal extremity of the external portion
of the epi sternum in Anahrus there i s a small s c le rite upon which the
trochantia tn^ p iv o ts.
The small s c le rite i s very probably a portion of
the prim itive coxopleurite, p a rt of which has fused with the anopleurite
to form the p i euro n and p a rt of which has remained a free s c le rite , the
trochantia tn^.
Prosternum:
has!sternum hs;
p o sterio r to the prepectus ppe of fig . IS is the
The furcal p its fup are the outward evidences of stern al
apophyses which form the furca fu^.
The furcal p its fup are inwardly
connected hy a ridge or stem acosta of which the outward evidence i s , in
Anahrus, a groove the stem aeostal suture stc s.
The stem acostal suture
-IS -
dlvides the eusternum into two p a rts.
The an terio r p art is the has!sternum
hs; the posterio r portion i s the sternellum or furcasteraum fs.
The furca
i s useful fo r support of the prothorai and for muscle connection.
Posterior
to the furcasteraum fs^ and separated from the l a t t e r hy a narrow
membranous area i s the spinasternum ssj which in Anabrus i s a very narrow
transverse s c le rite .
The spinasternum ss bears the spinal p i t spp an out­
ward evidence of an in tern al apodeme, the spina sp i.
The spina in Anabrus
is roughly hammer-like in shape.
PTEROTEOBAX
Mesonoturn;
shown.
In fig . 14 the dorsum of the male p tero thorax is
Immediately behind the lntersegmental maabrane is the acrotergite
atgg of the mesothorax.
The antecostal suture acs divides the acro­
te rg ite atgg from the prescutum psc.
by an apophysis or prephragoa gh.
as in the male.
In tern ally the suture i s represented
The prephragma ph of fig . 14 i s shown
In the female the prephragma ph i s shorter.
set i s the area immediately behind the prescutum psc.
scutum i s marked sparingly with setae.
the scutum set i s the scutellum gel.
The scutum
In Anabrus the
The slig h tly raised area behind
Extending poster!ad from the ante-
costal suture to opposite the re a r tip of the scutellum and running along
the side of the processes mentioned above are the maxillary s c le rite s by
means of which the tegmen i s artic u la te d .
These s c le rite s w ill be
discussed under "articu la tio n o f the wings".
In the male Anabrus the
posterior p art of the mesonotum i s folded under immediately behind the
scutellum scl and then refolded to form the postecutellum p sc l.
The f i r s t
-1 9 -
fold, v isib le as the p o sterio r s c le rite of the eunotum i s the p o stte rg ite
the female cricket the postscutellmn i s absent.
The spiracles s^ are considered p art of the te r gain thron^iout
the body.
Anabrus presents a special modification of the mesothoracic
spiracle sp^ of fig , 13 ^diich i s not evident in Staiopelmatus or Ceutho—
phiIus.
In the membrane between the propleuron and mesopleuron of fig . 13
there may be seen a "biforous" sp iracle.
The meso thoracic spiracle of
Anabrus consists of two openings each of which leads to a trachea.
The
larger (posterior) opening is unusual for an Orthopteroid in th a t i t is
guarded by numerous tiny eatae.
This large opening gives immediate entrance
to a large air-sac type of trachea.
guarded by two small s c le rite s ,
The smaller, an terio r opening is
The p o sterio r s c le rite appears to be p art of
the :peritrem all s c le rite of the p o sterio r opening.
gives immediate entrance to a small trachea.
The smaller opening
The tracheae of both entrances
do not lead into a common chamber and apparently are in no way connected.
Although th is i s not a resp irato ry study a b r ie f note on the tracheae
involved w ill not be out of place.
The large a ir-sa c type of trachea is
connected transversely with i t s fellow on the other side of the thorax by
means of a very narrow trachea.
leads d irec tly to the front leg.
From the body of the sac a large trachea
The large trachea i s about one-third the
diameter of the leg i t s e l f and extends down the leg p ast the auditory
apparatus aa of fig . 12.
The smaller (anterior) trachea extends cephalad.
I t bears several branches a t the head.
Midway between the head branches
and the o rific e th is trachea gives o ff a branch sharply in a ventral
direction.
This branch divides several times also; one of the subsidiary
-2 0 -
tranches descends to the front leg and another to the long ventral tracheal
chain.
Metanoturn;
notum.
The metanotmn of Anabrug i s somewhat lik e the meso-
The acro terg ite atg^ i s followed by the prescutum psc, .
prephragma ph, i s much smaller than in the mesonotum.
anterio r to the scutellum sc l.
postscutellum or postnotum.
The
The scutum set is
In Anabrus both sexes lack the metathoracic
To the dorsolateral area, to iriaich i s artic u ­
la ted the wing in the mesothorax, is attached a membranous wing pouch wj.
Mesopleuron?
Snodgrass (1909) drew figures of the in te rn a l and
external mesopleuron of Anabrus simplex.
This author points out the
existence of a preepisternum homologous to th at seen in Qryllus. D jseosteira,
®nd Melanoplus.
The s c le rite in question is separated from what Snodgrass
(1909) called episternum by a suture which runs almost the f u ll length of
the epi sternum.
Gumey (1936) noted the existence of "a lengthwise suture"
on mesopleuron and metapleuron in Ceuthophilus.
These sutures would
appear to be homologous to those sim ilarly placed in Anabrus.
In th is
paper, the preepi sternum w ill be considered merely as p a rt of the
episternum, and th at the episternum epsg i s separated from the epimeron
2E 2 V the pleu ral suture.
The p leu ral suture p is i s represented in te rn a lly
by a pleural ridge which extends down from the wing process wgg of fig . 13
to the pleural coxal process.
The pleural ridge of the mesopleuron
branches near i t s upper extremity and comes forward to support the
epi sternum.
The an terio r portion of the epi sternum i s divided tran s­
versely by a suture.
At the base of the tegmen are two paraptera.
The
one anterio r to the wing process wp^ i s the basalare bag, that p o sterio r
-2 1 -
to the wing process wp^ I s the euhalare sa^« Both are aids In wing
manipulation and are used for muscle attachment.
The pleural apophysis
p la is an invagination forming an in tern al arm continuous with the pleu ral
ridge.
The pleural arm is closely associated a t i t s d is ta l end with the
furca.
The precoxale prcxo of the mesothoracic pleuron is hasally contigu­
ous with the base of the an terio r episternum (preepisternum of Snodgrass).
I t should he noted in fig . 13 th at the precoxale of the mesothorax i s
considerably shorter than the one present on the propleuron.
Metapleuron;
The metapleuron is very much lik e the mesopleuron.
Perhaps the most strik in g difference i s the modification attendant to the
atrophy of the wings.
In Anahrus neith er sex i s able to fly .
The anterior
and posterio r la te r a l edges of the episternum eps, have fused with the
metanotum mtn of fig . 13.
The wing process wgj has not only fused with
the no turn but also has fused with the subalare s a ,.
The basal are b a, is
— J
— J
closer to the epi sternum eps^ (being separated from the l a tte r
suture) than the corresponding p a rts o f the mesothorax.
only
by a
The suture which
divides the episternum eps? into two p arts does not run through the fu ll
length of the la t t e r .
There i s no precoxal bridge in the metapleuron.
Mesosternum;
As shown in fig . 18 the mesothoracic b aslsternum
of Anabrus i s divided into two p a rts, the an terio r b a sisternum abs2 and
the posterio r basi sternum pbs^,.
The an terio r basi sternum absg i s composed
of two roughly quadrate s c le rite s which are projected well below the level
of surrounding membrane.
Two broad spines pro ject below the basal portion
of the posterior basi sternum ^bs2.
Immediately p o sterio r to the spine-
bearing process th ere i s a median trian g u lar impression in the sternum.
-2 2 -
The triangular area hears the roots of three apophyseal processes which
form the furca fu^.
This s c le rite is the double s c le rite furcasternum
plus spinastermun fs s 0.
In tern ally furca and spina are Joined.
The d is ta l
ends of the furca fug are closely associated with the d is ta l ends of the
pleural apophyses p la of f ig . 13.
The spina of the mesosteroum hears one
median and two p a irs of la te r a l projections.
Metasternum:
The metastemum of Anahrus i s divided into an
an terio r has!sternum ahsj and a p o sterio r has!sternum ghsj.
has!sternum ahs^ Ie a single prominent s c le rite .
The an terio r
The p o sterio r h a sisternum
phs^ i s marked hy the presence of two prominent hroad spines.
furcasternum fs^ is united with the p o sterio r hasi sternum phs, .
The
As in
other Orthopteroids except G ryllohlatta the spina i s lacking in the
metastemum.
A rticulation o f the wings;
of ax illa ry s c le rite s .
The tegmlna are artic u lated hy means
In fig . 14 th e ir disposition and number may he
seen as well as the disposition of other p te ra lla in the male thorax.
The
tegula tg i s a small hairy scale-lik e Iohe at the an terio r base of the
tegmen.
Between the tegula t£ and artic u lated with the Costa i s the
humeral p la te hup.
The f i r s t a x illa ry s c le rite lax in Anahrus i s supported
hy an anterio r notal process which i s hent under the main hody of the
no turn so th at a good portion of the a x illa ry i s hidden beneath the no turn
also.
The an terio r end of the f i r s t ax illa ry lax a rtic u la te s with the
complex of co stal, subcostal, and rad ia l veins which have anastamosed a t
the base.
The second a x illa ry 2ax i s hinged mesally to the f i r s t a x illa ry
and the vein complex mentioned above i s flexibly attached to i t s outer side.
The second a x illa ry 2ax may also he seen on the ventral side of the tegmen
In fig . 13 where I t artic u lated with the meso thoracic wing process wp^.
The th ird a x illa ry 3ax, "because of the cen tralizatio n and merging of most
of the important longitudinal veins of the tegmen represents in Anahrus
a fusion of the th ird a x illa ry and the two median p lates which are usually
hasad o f the Media, Cuhitu s, and f i r s t Anal vein.. The remaining tegmen
artic u la tio n represents an in te re stin g interm ediate stage. . I t i s generally
considered th a t the fourth a x illa ry i s a strongly s c le ro tlzed portion of
the posterior notal process which has "become separated from the la tte r .
The posterior apex of the th ird a x illa ry a rtic u la te s with the fourth
a x illa ry when i t i s present and with the p o sterio r notal process when
the fourth a x illa ry i s not present.
ex ists.
In Anahrus an interm ediate condition
The e c le rite lab elled Uax in fig . lU i s separated from the main
p art of the p o sterio r no turn "by a th in ly sclerotized area which i s not,
s tr ic tly speaking, membranous.
The s c le rite labelled Uax may therefore he
considered to be a p art of the p o sterio r notal process.
Since i t i s more
highly sclerotized than the r e s t of the p o sterio r notal process and i s fused
with the p o sterio r apex of the f i r s t a x illa ry lax i t i s here considered to
he an ax illa ry Uax.
The fourth a x illa ry s c le rite Uax a rtic u la te s with the
th ird a x illa ry s c le rite 3 ax.
; In the female Anahrus the tegmen i s more reduced and shows an
even more strongly fused group of a x illa rle e .
All are apparently present
in a less sclerotized sta te than in the male.
Wings;
sexes d iffe r.
I t can he seen in fig s. 22 and 2] the tegmlna of the
The tegmen of the male i s la rg e r.
In th is sex the Costa,
-2 4 -
Sa'bcosta, and Eadlus are present together as large ridge with the several
veins composing suh-rldges.
Very l i t t l e other venation I s recognizable
because the tegmen Is so atrophied.
of the tegmlna.
As In many other strld u lato ry Insects the l e f t tegmen
folds over the rig h t.
surface.
The male Anabrus strld u la te s by means
Both tegmlna bear a f lle - llk e vein on the ventral
S trld u la tion I s accomplished when the tegmlna are vibrated, the
f ile on the ventral surface of the l e f t tegmen being scraped on the veins
near the posterodorsal margin o f the rig h t tegmen.
highly sclerotized than the l e f t .
almost hyaline.
the male.
The rig h t tegmen Is le ss
The median c e ll in the rig h t tegmen is
The female tegmlna are shorter and thicker than those of
Thqy cannot be folded over each other.
All that remains of the
venation in th is sex i s the presence of four longitudinal ridges in a
homologous position to the ridges on the male tegmen.
means of strid u la tio n .
The female has no
All th at remains of the wings in both sexes i s a
membranous pouch Wj of fig s. 13 and 14.
'
'
"Lias
Although the legs could have been considered in the discussion
of the pleural areas, they are here discussed together fo r convenience and
ease o f comparison.
The pro thoracic leg may be seen in la te r a l view in
H g. 12 and I t s basal a rtic u la tio n may be seen in fig s . 13 and 18.
fig . 13 the coxa
Cx1
i s seen to have two means of a rtic u la tio n .
In
The
f i r s t i s the pleu ral coxal process (a projection o f the pleuron); the
second i s the trochantin tn^.
The coxa cx of fig . 12 i s rath er broad.
I t bears a basal border the basicoxlte box which i s set o ff from the main
-2 5 -
d ls ta l portion of the coxa by a suture (the basicostal suture).
Bie
basicoxite box in the foreleg of Anabrus bears one small fro n to -la te ra l process
and two meeal knob-like processes which are probably sensory in nature.
The more la te r a l s tria te d process may be seen in fig . 12.
The mesal knob­
lik e processes bear tin y setae and may be seen in fig . 13 where the coxa
Cxl has been pushed rearward for purposes of illu s tr a tio n .
one very large spine and numerous small setae.
The coxa bears
The main body of the coxa
bears a small invagination (indicated by the dark p it in f ig . 12) which is
used for muscle connection.
The coxa a rtic u la te s with the trochanter t r
by means of two fu lc ra l points k, k which are seen best in f ig . 13.
Near
the fu lc ra l points the coxa i s strengthened by in te rn a l ridges which are
represented externally by sutures which cut o ff small s c le rite s from the
main coxal walls.
membrane.
tin y setae.
■
■
-'v -
The coxa and trochanter are connected loosely by
The trochanter t r and femur fe are joined firm ly.
B0th bear
The femur fe of fig . 12 i s ridged and f la t beneath.
.
The mesal
-
ridge of the femur bears several spines.
Both la te r a l and mesal ridges
usually bear a p a ir of small apical spines.
The tib ia ti_ is quadrate-
cylindrical in cross section and of uniform thickness. The under side i s
'i
** ■'
'
,
somewhat flatten ed . Bie tib ia t l bears a variable number of spurs. Bie
usual arrangement i s six ventral p a irs, a laterodorsal row o f fiv e, and
two or three on the medio dorsal surface.
Near the base of the tib ia t i
of fig . 12 there may be seen a s l i t labelled aa.
the mesal side by another s l i t .
apparatus.
This s l i t i s matched on
The two together comprise the auditory
Below the auditory apparatus aa there is a p a ir of small
impressions one of which may be seen in fig . 12.
Bie impressions are
—26—
deeper than fa in t tlh la l grooves shlch lead into them.
i s four-segmented.
The tarsu s ta r
The four segments are loosely joined and In fig . 12
have "been pulled apart to show th e ir a rtic u la tio n to t e t t e r advantage.
The f i r s t ta rsa l segment i s the longest and broadest; the fourth Is almost
as long and i s the most slender.
tilo te d .
The th ird ta rsa l segment i s d istin c tly
H g. 11 i s a ventral view of the p re ta r sue of the foreleg.
pretarsus i s attached to the fourth ta rs a l segment ta^.
The
The ungoitrac to r
p la te u tr when pulled by the tendon ten moves the unguis.
Mesothoraclc le g ; As in the foreleg the coxa cxg of fig s. 13
and 16 a rtic u la te s on a pleu ral coxal process and on the trochantin tn .
—2
Like the f i r s t trochantin, the trochantin tn^ i s rath er hoot-like in shape.
I ts basal end a rtic u la te s with the p i ear on a t the terminus of the suture
which divides the episternum eps^ in to two p a rts , and thus i s closely
associated with the p o sterio r end of the precoxale prcxg.
The membrane
between the base of the coxa and the trochantin i s marked by a narrow lin e lik e sclero tizatio n which delim its the inner attachment o f the basicoxite
hex and is indicated in fig . I J by a lig&t dotted lin e .
bex bears knob-like processes as in the foreleg.
The basicoxite
The pleural a rtic u la r
socket pas of fig . 16 a rtic u la te s with the pleuron and divides the
basicoxite into two p a rts.
meron mer.
points.
The p o sterio r portion of the basicoxite is the
The trochanter t r a rtic u la te s loosely with the coxa a t two
The trochanter has an Internalbgsicosta (represented by an external
suture) which gives i t a marginal s c le rite .
This proximal s c le rite bears
a small setae-bearing process sim ilar to the areas present on the
basicoxite.
The trochanter t r and femur fe are Joined firm ly.
i s grooved beneath and bears a variable number of spines.
The femur
The tib i a t i i s
—27—
shaped very much lik e the tih ia of the foreleg and hears a variable
number of spurs.
The four-segmented tarsus ta r Is attached to the tib ia
and bears the claws or ungues ung.
Metathoraclc le g : A. la te r a l view of the hind leg may be seen in
fig . 15.
The hind leg is very large and i s not drawn to the scale of the
other legs for th is reason.
poBteroventral side.
The coxa cx i s marked by a suture on i t s
Posteroventral views o f th is and the other two coxae
may be seen in fig . 18.
of the mesothoracic leg.
In most respects the coxa i s very much lik e that
The trochanter t r is very small and when not
pulled out as in fig . 15 can hardly be seen.
firmly to the femur fe.
The trochanter i s joined
At the base of the femur a b asicostal suture
separates a marginal s c le rite from the remainder of th is leg segment.
femur bears a variable number of spines.
spihes and spurs.
form plantulae p in .
The
The tib i a t i bears numerous
The p u lv illu s of the f i r s t ta rs a l segment i s divided to
-2 8 -
ABDcam
The abdomen of Anabras la essen tially sim ilar to those of other
Tettigoniidae.
Walker (1919) discussed female terminal abdominal
structures of Orthopteroids in a manner which affords easy comparison
with homologous structures in the Mormon crick et.
Snodgrass (1937) stated
that Angbrus and other Decticinae exhibit some important p h allic modification.
Abdominal segments:
in shape.
The abdomen of Anabrus i s rather cylindrical
As shown in fig . 19 there are ten abdominal te rg ite s t^ 1Q, and
nine s te m ite s S1^ in the male.
The female as shown in fig . 20 has ten
abdominal te rg ite s and eight ste rn lte s .
In both sexes the membranes
between the te rg ite s are almost as wide as the te rg ite s themselves.
Normally most of th is membrane i s hidden in the telescoping of the abdomi­
nal segments.
The sternum, however, does not reveal a sim ilar equality of
s c le rite and membrane.
Therefore, i f the abdomen of Anabrus were drawn
out fu lly i t would appear quite lik e a sem i-circle.
The ste rn lte s are
very small and the membranous pleu ral area veiy large.
The membranous
pleural area is composed of dorsal, p leu ral, and ventral membrane.
The
te rg ite s and s te m ite s are d is tin c t.
There are eight p airs of abdominal spiracles 8£ in
^ f r brua'
n i &re located in the membrane.
An abdominal spiracle consists
of two small s c le rite s which guard the opening to the trachea.
TERMINAL STRUCTURES OE mat.-b
Tergites;
S tarting with the eighth te rg ite tg of fig . 19 the
te rg ite s are smaller than those preceding i t .
Except fo r a tiny median
-2 9 -
dorsal area the an terio r border of the tenth te rg ite t ^ i s almost
en tirely hidden from external view both by the p o sterio r edge of the
ninth te rg ite and. by membrane.
ary modification.
The tenth te rg ite t ^ presents a second­
The median portion i s of a membranous nature as shown
in fig s. 19 and 21.
Sternlte s t
The ninth s te m lte s^ of fig s. 19 and 21 i s larger
than the other s te m ite s .
This ste rn !te i s the subgenital p la te .
divided transversely by a suture.
ste m lte bears s ty li sty .
I t is
The p o sterio r portion of the ninth
In prim itive insects such as G rylloblatta the
s ty li stg are borne by coxites.
The ninth s te m lte in Anabrus i s
considered, therefore, to bear un ited ninth coxites.
Gurney (1936) pointed
out that s ty li are not present in Ceuthophilus but are present in
Bhaphidophora, Neoconocephalus, and Scudderla.
Cercus? The male cercus i s broad unsegmented, hooked apically
and with a tooth directed mesally.
Cowan (1929) used the cercus to
iden tify the in sta rs of Anabrus simplex.
Gumey (1939) pointed out that
the size and angle a t which the tooth p rojects from the main body of the
cercus may be used to separate A. simplex and longipes from cerclata since
in the l a t t e r the inner tooth i s large and p ro jects at rig h t angles.
Anal s c le r lte s : About the anal opening a of fig . 21 are three
s c le rite s.
The suprarenal p la te sap i s often called the epiproct by
system atists and was so called by Snodgrass (1937)•
However, the former
term was applied to the tenth te rg ite by Crampton (1929, 1931) as Gurney
(1936) pointed out.
On each side of the anal opening a and below the
supra-anal p la te sap are the paraprocts ppt which are well defined
-3 0 -
s c le rlte s in Ana~brus.
P tola:
Snodgrass (1937) discussed the penis of Anahrus simplex.
The tettig o n io id type of p h allic structure hears in a modified form the
three p airs of p h allic lohes usually present in the Orthoptera.
la te ra l p h allic Iohe Ig l may he seen in fig . 19.
The
Beneath the la te r a l p h allic
Iohe the posterior edge of the ventral p h allic Iohe v l may he seen.
The
dorsal p h allic Iohe is hidden in te rn a lly and cannot he seen in fig . 19.
I t divides the p h allic cavity in tiro acting as the floor of the upper
cavity.
The ventral p h allic Iohe i s the floor of a ventral cavity or
spermatophore sac.
The sc le ro tic arms of the phallus re s t upon the top
of the dorsal p h allic lohe. ,
TSBMIKAL STBUCTtJBB 07 IBiALB
general features:
The most marked differences from the abdominal
s c le r itea of the male are seen sta rtin g at the eighth segment.
The eighth
and ninth te rg ite s tg and t^ of fig . 20.d iffe r in shape from the homologous
male te rg ite s .
of the male.
female.
The tenth te rg ite t
i s rath er sim ilar to the tenth te rg ite
There are only eight complete abdominal ste rn ite s in the
The v alv ife r v f i s a p a rt of the ninth abdominal ste rn ite .
The
eighth abdominal s te rn ite Sg a rtic u la te s with the lower edge of the
eighth te rg ite t^ .
The cercus ce of fig . 20 i s slender and not broad and
hook-like as th at of the male shown in fig . 19.
The supra-anal p la te sap
and paraprocts ppt are sim ilar to those of the male.
Ventral valves? The valves of the adult female Anahrus f i t
together tig h tly to form a sword-like ovipositor which i s as long as the
-3 1 -
abdomen.
The ventral valve vv of fig , 20 la fused a t I t s "base with the
taslvalvula hv. The v alv ifer vf i s a small s c le rite to which the
dorsoproxlmal edge of the ventral valve vv, and the dorsal valve dv are
attached.
The ventral valve hears a membrane on the proximal th ird of
i t s mesal surface.
Dorsal valves: The dorsal valve dv i s broader than the ventral
valve and lik e i t i s attached to the v alv ife r vf.
The dorsal valve bears
two in tern al apophyses, the superior apophysis sup and the in fe rio r
apophys i s ia , for tendon attachment.
Inner valves;
and ventral valves.
The inner valve Iv of fig . 20 f i t s in to the dorsal
The ramus r i s an arm of the ven tral valve Tqr which
i t i s connected to the body.
the two inner valves.
The pons valvorum gv i s a bridge connecting
The inner valves are attached dorsally to the
superior intervalvula si and vent ra lly to the in fe rio r in te rv a l vula inv.
-3 2 ABBESVIATIONS MD LlTTlBINa QN PLATS HfflJBES
a
a t3
acs
an
ant
asm
atg
ax
axe
L
La
Lc
Lex
Lg
Lgl
LI
Lm
Lmm
Lr
Ls
Lu
Lv .
anal opening
auditory apparatus
anterior Lasietenmm
antecostal suture
an termale
antenna
anterior arms of suLmentum
acro terg ite, p re te rg ite
a x illa ry s c le rite
a x illa ry cord
Lasimaxillary memLrane
Lasalare
Lasicardo
Lasicoxite
Lasigalea
Lasiglossa
Lasillngua
suLgena, LaslmandiLulare
LaslmandiLular memLrane
L rustia
Lasisternum
Luttressed area
Lasivalvula
c
c da
ce
cp
cs
ctn
cx
cxc
clypeus
car dosuture
cercus
cardoprocess
coronal suture
cardotendon
coxa
coxal cavity
d
dc
dcs
dg
dl
dv
articu lato ry process of Lasicardo
disticardo
dorsal cervical s c le rite
d lstig alea
d lstilin g u a
dorsal valve
e
ec
ep
eps
epm
eg
compound eye
endocardo
extensor prominence
episternum
epimeron
endognath
es
esp
est
et
cu
epistomal suture
en d o stlp ital process
endostipes
extensor tehd(m
eutentorium or Lody of tentorium
f
fas
fee
fe
tp
fs
f ss
ft
Ai
Aip
frons
fastigium
frontoclypeal suture
femur
fro n ta l p it
Aircastemum
Aircastemum + spinastemum
flexor tendon
Airca
Aircal p it
ga
ge
gg
gi
gl
gn
gp
gs
gnathapex
gena
glossiger
ginglymus
glossa
gnathite
gular p it
gnathal suture or impress
h
hypocondyle or gnathocondyle
hypopharynx
hup humeral p la te
ia
it
Inv
Iv
in fe rio r apophysis
in te r to rma
po sitio n of in fe rio r intervalvula
inner valve
k
A ilcral point of leg segment
I
la
IL
Ic
Id
II
II
Ip
Ip l
III
la c in ia
IaLrum
long Lrachymere
latero cerv icale
laciniadentes
IaL ial s tip ite s
lingualora
IaL ial palpus
la te r a l p h allic IoLes
lacin u la
-3 3 -
m
md
mer
mn
men
mp
mtn
mol a
mandible
meron
menturn
mesonotum
maxillary palpus
metanotum
nf
neural foramen
occ occip ital condyle
ocf o ccip ital foramen
p
par
pas
pbs
pgs
pd
pf
pg
pga
pge
pgl
pgs
ph
pi
p la
pin
p lr
pn
po
poc
poe
pos
PP
ppc
ppd
I'. .
ppt
pra
prex
pre
ps
Psc
Pscl
pss
p et
pt
posterio r te n to ria l arch
p a rie ta l
pleural a rtic u la r sock (o f coxa)
posterio r has!sternum
postgenal suture
pedicel
pal&lfer
palpiger
postgenal acetabulium
poatgena
paraglossa
postgenal suture
prephragma
postlingua or linguatendoh
pleural apophysis
plan tu la
pleural ridge
pronotum
postocciput
parocciput or p o sterio r arms of
tentorium
postepipharynx
postoccip ltal suture
postgenal process
p r^ e c tu s
postpedicel, th ird
antennal segment
pareproct
prealare
precoxale
preepipharynx
parastipes
prescutum
postscutellum
paras tip i ta l suture
parastorn!urn
pretentorium or an terio r arms of
tentorium
p tg p o stte rg ite
pv pons Yalvorum
r
ramus of inner valve
s
Bi_g
sa
sap
sb
sc
scl
set
sd
si
si
sm
smm
sp
spi
spp
ss
st
stcs
sty
sup
stip es
abdominal ste m ite*
subalare
suprarenal p la te
short brachymeres
scape
s cut ellum
scutum
slender dolichomeres
p o sitio n of superior intervalvula
suprallngua, dorsolingua
submentum
submental membrane
spiracle
^pina
spinal p it
spinasternunu <.
supratentorium
stem acostal suture
stylus
superior apophysis
ti_ioabdominal te rg ite
tai| d ls ti tarsus
ta r tarsus
te
tempua, temple
ten tendon
te s temporal suture
tg tegula
ti
tib ia
tn tr o chantin
to
torma
tr
trochanter
ung unguis
u tr u n g u itra c tir p la te
vc
vl
w
vf
ventral cervical s c le rite
ventral p h allic lobe
ventral valve
v alv ifer
Vg
Wj
meeothoracic wing or tegmen
metathoracic wing pouch
-34Aaabrus simplex
P late I
Iig . I .
Fig. 2.
Frontal view o f the head.
P osterior view of back of head.
Hg. 3* Antenna.
H g. 4.
P osterior view of dextral mandible.
H g. 5«
Anterior view of tentorium and neighboring p a rts.
H g. 6.
Inner surface of cardo and stip es.
H g. 7.
Inner or buccal view of labrum and clypeus.
H g. 8.
Anterior view of gloseae and paraglossae.
H g. 9.
Anterior view of la c ln la and galea.
H g. 10.
L ateral view of hypopharynx and upper and lower lip , with
most of head capsule removed.
-3 5 pla ts
I
I pH
Hj
I/Tl K/ S i
Up. 3
M
Iii
-3 6 -
Anabrus simplex
P late II
rig . n .
Ventral view of pretarsus of s in ls tr a l foreleg.
H g. 12 .
la te r a l view of foreleg.
H g. 13.
Lateral view of thorax.
H g. 14.
Dorsal view of male pterothorax.
H g. 15 .
Lateral view o f rear leg.
H g. 16 .
Lateral view of mesothoraclc leg.
Hg. 17.
Lateral view of base of re a r tib ia .
H g. 18.
Ventral view of thorax,
thorax.
-37PLATE II
f
-3 8 -
Anabras simplex
P late I I I
H g. 19.
la te r a l view of adult male abdominal sc le rite e .
H g. so.
Lateral view of adult female terminal e c le rite s.
H g. SI.
Dorsoposterior view of adult male terminal s c le rlte s .
H g. 22.
Female tegmen.
H g. 23.
Male tegnen.
-3 9 pla te h i
F > .1 9
R f . 20
n F '21
R f 23
-UoS e c tio n I I
I
THS PBEBTBATION OP SODIUM AESSNITS
TEEOUOS THB BXDSKSLBTGN
REVIEW OP IITEEATUEE
Sodium arsen ite has "been used both as a stomach and a contact
poison.
I t was not u n til 1923» however, th at reference was made concern­
ing i t s p o s s ib ilitie s in the l a t t e r method of k illin g in se cts.
Previous
to th a t year i t s efficiency as a stomach poison especially in b a its was ?
well known. Mally (1923) found th at sodium arsen ite when applied by
means of hand bellows as a dust to the antennae o f in sects k ille d them
even though i t was not Ingested.
Granovsky (1926) summarized the studies of several Russian
sc ie n tists.
Sovdarg, one of the in v estig ato rs, dusted and sprayed migratory
grasshoppers with d isodium acid arsen ite (HaH AsOj)* and d lsodium acid
arsenate (Ha2H AsOl4) .
Death occurred even thou^i the in sects were given
no opportunity to ingest the poison.
Sovdarg pointed out th at the spray
acted upon the in sects more rapidly than the dust and th at i f humidity
were higher the arsenicala reacted more rapidly.
Granovsky (1926) noted th a t grasshoppers which had crawled
through poison b a it in which sodium arsen ite was an ingredient often
were found with the ventral p o sterio r thorax and an terio r abdomen in a
disintegrated condition.
Other p arts of the body showed no signs of
Injury * Noting the strength of the in secticid e th is author suggested
the use of hydrated lime as a d iluent.
Shotwell and Cowan (1928) were f i r s t to te s t sodium arsen ite (in
the form of spray as well as dust) on the Mormon cric k et.
*- Probably Ha2H AsOj.
In te s ts on th is
insect and the le sse r migratory grasshopper, Melanoplns mexl caaus
(Sanssnre), excellent re su lts were obtained as fa r as 200 feet from the
point of application.
The poison was also effectiv e when used as a b a rrie r.
These workers reported injurious action by the in secticid e on several
plants.
Cowan (1929) and Cowan and McCampbell (1929) noted th a t sodium
and calcium arsen ites in combination with hydrated lime are very effective
as a means of controlling the Mormon cricket in a l l in s ta rs .
Hot only
did m ortality remain h i b u t crops could be b e tte r protected by the use
of such a diluent.
Zing and Rutledge (1932) stated that irre g u la r re su lts were obtained
iAen a cloud of sodium arse n ite dust was blown through a cage containing
Locusta migratoriodes (Rch. and Zrm.).
X series of experiments were carried
on by these authors consisting of applications of dust to several p a rts of
the lo cu sts' anatomy.
Ho i l l effects were observed a f te r two days.
In
another experiment the locusts were allowed to f lu tte r in a cloud o f dust
and ingestion was prohibited.
These locusts died and since arsenic was
recovered from dissected tracheae the authors conclude that sodium arsen ite
probably has the tracheal system as i t s point of penetration.
O'Kane and Olover (1935) placed arsenic-containing wax c e lls on
the body of Perlplaneta americana ( I . ) .
Arsenic was recovered from v ita l
in tern al organs, proving th at penetration took place th ro n g the integument.
Both sodium arsen ite and arsenlous acid penetrated, possibly with the aid
of a surface flu id .
P aralysis did not re su lt from use of the arsenic.
Schweiss and Burge (1935) reported th at sodium arsen ite and
-4 2 -
Iio e , in combination, set uplaa i r r i ta t io n which i s followed "by cleaning
of the affected p arts and subsequent ingestion of the poison.
Olover (1936) • using a wax c e ll on P eriplaneta aaerlcana, found
that both arsenious oxide and sodium areenite penetrated the Integument.
Sodium areenite was found to penetrate the more rapidly of the two.
At
high concentrations arsenic was recovered in a l l tissu es and was voided
through the digestive tr a c t.
Concentration of arsenic in the roach varied
with the area of the c e ll.
-
. -
Hastings and pepper (1939) reported on ce rtain in e rt diluents
used in conjunction with sodium areenite on the Mormon crick et.
insecticid e was found to be both a contact and a stomach .poison.
of 50^ m ortality was reached e a rlie r a t higher temperatures.
The
The point
Adults varied
more with temperature changes than did the nymphs in re la tio n to the 50^
m ortality point.
PBOCBIXJHS
The procedure adopted in th is study may be divided into three
p arts: (I) the general handling of crickets* (2) the dusting of crick ets
with sodium areenite to discover whether some p a rts of the exoskeleton
are more susceptible to penetration than other p a rts , and ()X the chemical
analysis of the crick ets a f te r death to determine whether a rela tio n
ex ists between the amount of arsenious acid recovered, the time fo r 50^6
m ortality, and the exoskeletal area treated .
I . General treatm ent:
During the study crick ets were collected from the Infested areas
south of B illin g s.
Collections were made several times and the collections
of three dates were used.
This was made necessary "by the fa c t th at adult
crickets w ill not remain in good h ealth when held in cap tiv ity .
After co llectio n the crick ets were brought to Bozmmn and placed
in a cage out of doors.
During th is period they were fed with a lfa lfa ,
clover, and dandelion p lan ts.
Apparently normal crickets were then brought
indoors and kept in a small cage w ithin a temperature cabinet where the
temperature remained constant a t 27*0. and the humidity remained constant
a t approximately 53#»
The crick ets remained in the temperature cabinet
for about 36 hours before being trea ted with the in secticid e.
During th is
period the Insects were fed regularly but they were not fed a f te r the
application of the dust.
I I . The d u s tin g o f th e c r i c k e t s :
Since p re fe re n tia l penetration (of the same m aterial) through
differen t exoskeletal areas was one of the objects of the study i t was
decided to dust ce rtain p a rts of the exo skeleton with chemically pure
l|l
meta sodium arsen ite and leave other p arts free of the poison.
Because
Anabrus simplex is a large and active insect i t was recognized th at merely
placing the poison upon the desired lo c a lity would insure neith er that
the poison would remain on th a t p a rt of the body, nor th at i t would not
be ingested a fte r cleaning with the mouthparts.
In order to insure more
exactness in these m atters the crick ets were (I) f itte d with an appliance
to prevent ingestion, and (2) a means of lo calizin g the poison was perfected.
The cricket was prevented from ingesting the poison by the use
of a "collar" of the type used by Hastings and Pepper (1939).
♦-Baker’s C.P. meta sodium arsen ite (HaAsOg).
The "collar"
when f itte d over the head of the cricket prevented cleaning of the dusted
areas ty mouthparts#
Local!zation of the poison to the area to which i t
was applied was d if fic u lt.
Since the insect could move even with the
c o lla r in place, and would tw ist i t s body, the wax c e ll method as used
by O1Kane and Olover (1935) was found im practical for general use on body
areas of the Mormon crick et.
I t was decided, therefore, to develop a th in
coating of some m aterial which could be placed on the areas immediately
adjacent to those areas dusted.
tested and rejected.
Some th irty glues and p la s tic s were
I t was found that water soluble animal glues usually
did not adhere to the body of the cricket or were not ample protection.
Glues with a powerful organic solvent usually k ille d or in ju red the in sects
Several of these glues o r p la s tic s and several waxes necessitated use of
high temperatures during application.
The protective coating used in most cases in th is experiment
was a paraffin melting a t 38° to UO0 C. Because application in quantity
a t th is temperature could not but be eith er injurious or upsetting to the
normal physiological a c tiv ity of the crick et, the p araffin was applied by
means of a melting apparatus.
in p a ra lle l.
used.
A group of e le c tric sockets were connected
To complete the c irc u it, two th in wires (Alumel 27) were
The wires were connected by tw isting together and soldering, and
th e ir ends were file d to a fin e, needle-like p o in t.
Close to th e ir point
of connection each of the wires was sheathed in a narrow glass tube six
inches in length.
The glass tubes were fused and then taped together.
The grade of wire used offered enough resistance to the e le c tric current
to re su lt in a ris e in temperature which could be controlled by the
-45number of lamps and th e ir wattage placed In the sockets.
The warm pointed
Mid of the melting pencil was dipped Into the so lid p araffin and then
"brought close to the ezoskeletal portion to be coated.
Since the point was
very fine the p araffin could be applied in very minute q uantities which
would solidify immediately a fte r contact with the crick et.
Throughout the en tire experiment care was taken not to heat eith er
the wax or the pencil to a temperature which would be harmful to the crick et.
Application of p araffin was done in each case with the aid of a hand lens.
A fter application and careful te stin g to see i f the crick et was "apparently
normal", the insect was dusted on the exposed portion by means of a small
brush containing several fin e b r is tle s .
The c o lla r referred to above was
then adjusted and an id e n tific a tio n number w ritten on it* .
Crickets th at
had been so treated were placed in 5-inch p e trie dishes th at had been so
divided by cardboard p a rtitio n s as to allow equal spaces for four crick ets.
Ihen the four crick ets were in th e ir places the p e trie dish was covered
with nettin g to prevent th e ir escape, and placed in the same temperature
cabinet from which they had been taken p rio r to dusting.
Nine dusting experiments (series I to IX) were carried out.
In
table I the series are lis te d by number and th e ir treatment with C.P. meta
sodium arsenite (Na AsOj (without diluent) i s indicated.
•-Check lo ts of undusted crick ets were kept under the same conditions.
The check lo ts consisted of one unwaxed group wearing c o lla rs, one waxed
group wearing c o lla rs, and a small group th at had not been handled. In
no case was there any m ortality in the check lo ts before 100$ m ortality
had been reached in 4he dusted group.
-46T able I . —A p p lic a tio n o f sodium a r s e n l t e to v a r io u s p a r t s o f e x o sk e le to n
an d d i s p o s i t i o n o f p a r a f f i n
Series
No.
I
II
III
IV
V
VI
VII
V III
IZ
Sodium arsen lte dust applied
to;
dorsal abdominal membranes
I I to V III inclusive
P araffin applied to:
abdominal te rg lte s II to
VII inclusive
membranes before and
behind the te rg lte s
38-40*0. +
small amount
of beeswax to
make i t secur
ventral abdominal membranes
II to VIII inclusive
ventral abdominal
p lates
38-40» C.
ventral abdominal p la tes
I I to V II Inclusive
ventral abdominal
membranes
38-4o* 0.
la te r a l membranes of
abdomen
border of te rg lte s
38-40*C.
above and border o f
abdomen immediately
below pleu ral
membranes, also femora
of p o sterio r legs
pronotal shield
border of pronotal
shield, also femora
of f i r s t 2 p airs
of legs
membrane beneath pronotal
shield
te rg lte s before and
behind the membranes
w*
Type of
p araffin
$8-40* C.
38-40*0.
- -
thoracic pleura not
including propleura
femora of la s t two
p airs of legs
38-4o*c.
ta rs i of a l l the legs
tib ia e of legs
38-4o»c.
-47I t may be of in te re s t to note the method employed for Judging
the crickets to be normal or abnormal and the method of determining
m ortality.
I t was found th at an apparently normal cricket recently taken
from the fie ld reacts in a c h a ra c te ristic manner to certain stim uli.
A
sudden movement toward i t by an object (such as a pencil) two or three
inches from i t s eyes usually re su lts in a rapid re tre a t or a hasty back­
ward movement accompanied by continued playing of the antennae.
A normal cricket also can climb to the top of the cage and can
easily climb a cardboard held a t a 45» angle.
regain i t s feet i f turned on i t s back.
I t w ill, moreover, immediately
Lack o f the la s t reaction or more
than one of the preceding was taken as a sign of abnormality.
Observations of Mormon crick ets dusted with sodium arsen ite
revealed that th e ir a c tiv itie s and reactions underwent certain and
d efin ite change in a rath er regular order.
Perhaps the f i r s t sign of
abnormality i s a general sluggishness and slowing up of some o f the regular
responses.
This may be called "subnormality".
The second stage i s the
evidently great d iffic u lty encountered in returning to the normal position
from an inverted one.
This may be called "slig h t lo ss of coordination".
The th ird stage i s indicated by in a b ility to recover the upright position
and may be referred to as "loss o f coordination".
The fourth stage which
may be called "complete lo ss of coordination" may be recognized as th at
point in which the cricket w ill respond to stim uli by a mere twitching
and l i t t l e , i f any, organized response.
Death was taken to be the point
a t which crickets showed no movement and no v is ib le reaction to prodding
with a dissecting needle.
-46Throughout the study crick ets were used only I f they showed the
"normal reactions".
Check lo ts of crickets underwent the same te s ts .
After "being dusted and placed In the temperature cabinet the
crickets were examined a t hourly In terv als and th e ir condition noted.
Since each cricket was numbered I t s condition could be followed from hour
to hour and noted on a record sheet of the sample type shown In table I I .
below.
In th is tab le conditions are indicated by symbols*.
Table I I . —Sample data sheet.
____ ______ _________ _
Cricket ■I
number
I
2 3 4 p
AN SH
I
SLC
2
AH SH
SLC
AH SN
SLC
3
• 4
AH
SN SLC
AH
SN
SLC
5
6
AH
SN
SLC
Series I
Hours sifter dusting
b
8 9 10 11 12 13 14 15 1& i t
CLC
D
LC
LC
CLC
D
CLC
LC
D
LC
CLC
D
CLC
LC
D
CLC
LC
D
♦-Definition of abbreviations
AH - apparently normal
SH - subnormal
SLC - slig h t loss of coordination
LC - lo ss of coordination
CLC - complete loss of coordination
D - dead
I I I . Quantitative determination of sodium arsen lte;
When the crick ets were adjudged dead they were taken from the
p e trie dish and washed carefu lly to remove the sodium arsen lte which re­
mained on the outer surface.
The crick ets (with co llars s t i l l In place)
were then dried in an oven a t 85°C.
The in sects were divided Into ten groups of fiv e each according
to the order in point of time in which they died.
Thus the f i r s t five to
die were analyzed as a group, group I of series I ; the second fiv e to die,
group 2 of series I were analyzed as a group also .
In most cases the f i r s t
two g ro u p s to d ie ( group I and group 2) and th e l a s t two g ro u p s to d ie
(g ro u p 9 and group 10) w ere ta k e n f o r a n a l y s i s i n o r d e r t h a t th e ex trem es
m ig h t h e t e s t e d .
When more th a n f i v e c r i c k e t s d ie d i n th e same h o u r f i v e
c r i c k e t s f o r th e group w ere ch o sen a t random from th o s e h a v in g d ie d t h a t
h o u r.
I n some c a s e s b e c a u se o f te c h n i c a l d i f f i c u l t i e s o r d i s t r i b u t i o n o f
m o r t a l i t y , g ro u p s 3 and U o r 7 a n d 8 w ere ta k e n i n p r e f e r e n c e to th e
extrem e g ro u p s.
I n one s e r i e s sev e n o f th e te n g ro u p s w ere ta k e n i n o r d e r
to compare b e t t e r th e tim e o f m o r t a l i t y i n t h e g ro u p s w ith th e amount o f
a r s e n io u s o x id e d e te rm in e d .
The m ethod o f a n a l y s i s u s e d was a m o d if ic a tio n
o f th e Q u tz e lt m ethod, a s g iv e n i n A .O .A .C ., f i f t h e d i t i o n ( ig h o ) .
MTBCT OF IXJSTINQ THE ABDOMffl WITH SODIUM AHSfflITS
A. E f f e c t o f D u stin g th e Dorsum
A d u lt c r i c k e t s c o l l e c t e d a t th e same tim e w ere d iv id e d i n t o two
g ro u p s .
The f i r s t s e r i e s was d u s te d w ith sodium a r s e n i t e on membranes
I I t o V III i n c l u s i v e o f th e d o r s a l abdomen.
The sec o n d s e r i e s was d u s te d
on t h e t e r g i t e s I I to V II i n c l u s i v e o f th e abdomen.
I t was in te n d e d b y
t h i s means to d e te rm in e w h e th er o r n o t th e t e r g i t e s o r membranes o f th e
d o r s a l abdomen p e r m itte d g r e a t e r p e n e t r a t i o n o r r e s u l t e d i n a more r a p id
m o rta lity .
One more membrane was d u s te d th a n th e number o f t e r g i t e s
i n o r d e r to have a more e q u a l t o t a l a r e a .
I t may b e n o te d i n p a s s in g t h a t
th e membranes a r e n o t v e r y much s m a lle r th a n th e t e r g i t e s .
The g r e a t e s t
d if f e r e n c e l i e s i n amount o f ex p o sed s u r f a c e s in c e th e membranes a r e
fo ld e d i n b e n e a th th e s c l e r o t i z e d a r e a s .
I n d u s tin g th e membranes th e
abdomen o f th e c r i c k e t was b e n t g e n t l y so t h a t th e e n t i r e membranes c o u ld
-5 0 -
t# dusted.
,
In a comparison of exoskeletal areas for the purpose of th is
study I t is necessary to consider time required for reaction to the poison
as well as quantity of penetration.
The f i r s t two se rie s revealed l i t t l e
time difference in passing through the stages shown "by poisoned c ric k ets.
Tables I I I , IV and V show the manner in which the series reached these
stages.
The time necessary to reach 50# m ortality for series I and II
may "be seen in figure 24.
I t w ill he seen read ily from th is figure
the differences are. merely a matter of minutes.
th at
In view of these fa c ts
i t may he said th a t differences in speed of action are very small.
In comparing the amount of penetration shown hy quantitative
analysis two points were considered!
(I) the possible existence of a re la tio n
between time of m ortality within a series and the amount o f sodium arsen ite
th at penetrated; (2) the possible existence of a re la tio n between sim ilar
times of m ortality in d ifferen t se rie s and the amount of sodium arsen ite
found in the crick ets.
In the f i r s t series the f i r s t and second groups of fiv e and the
la s t group of five to die were analysed by modified Qutseit method.
From
the second series the f i r s t and second and the ninth and ten th groups of
five were selected fo r analysis, as outlined in the procedure above.
Table VI shows the re su lts of analysis.
In the f i r s t series comparison of groups I , 2, and 10 show no
d efin ite re la tio n between amount of penetration and the time of m ortality.
In series II the f i r s t ten to die show about three per cent more arsenious
oxide than do the la s t ten crick ets to die.
I t would appear then that in
each of the f i r s t two series (w ithin the series) time o f death and amount
-5 1 -
fable I I I .—Per cent loss of coordination
fim e i n
hours
I
2
3
4
5
6
7
8
9
10
11
12
S
15
I
18
54
80
92
96
II
III
24
2
80 86
98 100
100
100
-
IV
V
TI
4
0
4 86
12 96
36 96
70 100
VII
V III
0
10
48
82
96
0.00
6.67
17.35
24.00
29.33
0
4
16
34
52
98
98
100
33.33
48.00
58.67
70.40
72.67
62
68
80
88
90
80.00
90
92
92
96
100
86
98
98
100
C1 C9
0
0
3 30
60 95
85 100
95
IX
0.00
3.33
70.00
90.00
96.67
100
-
—
^2—
Table IV.—Per cent complete loss of coordination
Time in
hours
I
2
3
4
5
I
4
26
46
66
80
6
7
8
9
10
84
92
100
11
12
13
14
15
16
17
18
19
20
II
0
12
56
92
92
III
0
4
82
98
98
100 100
IV
0
0
0
4
20
V
I0
4
60
78.
90
VI
0
0
6
26
6s
42 96 ' 92
60 98 96
74 98 98
82 100 100
94
94
94
98
94
100
VII VIII
0.00
0
0.00
0
4.00
4
16
10.67
18.67
30
IX
0.00
0.00
0.00
36.67
63.33
30.67
54.67
66.67
70.67
78.67
34
48
56
68
78
93.33
96.67
85.33
90.67
92.00
92.00
93*33
84
86
86
C1
O
O
10
4o
65
80 90
90 95
95 100
95
95
83.33
96.67
96.67
96.67
96.67
100
96.67
88 96.67
94 100.00
96.00
96
96.00
96
97.33 98
98.67 100
100.00
Op
6
0
0
60
85
•
-5 3 Table V .—Per cen t m o r ta lity
Time I n
Ci
0
0
G?
0
5
0
20
o
50
65
2
3
U
5
I
0
2
12
16
22
II
0
0
0
8
14
III
0
0
2
4
18
IV
0
0
0
2
V
0
0
4
14
18
VI
0
0
0
2
24
VIII
VII
0.00
0
0
0.00
0
0.00
0
0.00
L6
0.00
IX
0.00
0.00
0.00
0.00
10.00
6
7
8
9
10
28
40
48
62
72
24
34
54
70
78
38
60
74
88
94
0
32
0 50
0 . 64
8 70
12 78
30
58
70
82
86
4.00
8.00
14.67
28.00
37.33
18
22
29
26
34
16.67
26.67
46.67
53.33
56.67
11
12
13
IU
15
76
86
90
94
98
84 96
88 100
92
96
98
90 92
94 100
98
42.67
50.67
62.67
68.00
72.00
44
52
56
75
66
70.00
83.33
90.00
93.33
97.67
16
17
18
19
20
98
100
77.33
84.00
88.00
90.67
94.67
80
100.00
95
95
95
100
94.67
96.00
97.33
100.00
90
92
92
98
hours
I
21
22
23
24
100
:o
20
28
44
54
64
66
74
78 100
78
84
90
98
100
82
86
88
88
o
0
90
10
100
do
65
----- -
5ERIE5 I
SERIES E
PERCENT
MORTALITY
-5 4 -
TIME IN HOURS
F ig. 24.
I
M o rta lity curves fo r dust a p p lic a tio n in
S e r ie s I (d o r sa l abdominal membranes) and
S e r ie s II (abdominal t e r g i t e s ) .
-5 5 -
o f sodium arsenlte that penetrated seem to have no d irect relationship.
In comparing series I and I I fo r evidence of differences in ra te
of penetration, i t w ill he seen th a t crick ets dying in the same time in the
two series cannot be sig n ifican tly compared since, as has Just been shown,
no relationship ex ists in a given series between time of death and amount
o f penetration.
Comparison by averages of crick ets in the two groups
shows th at series I absorbed .206 mg. of poison per gram of dry body welgjht
while series I I absorbed .228 mg. of poison.
The differences here are not
very significant because they are within the realm of experimental erro r.
B. Effect o f Busting the Pleura and Venter
of the Abdomen
Crickets th at were collected from south of B illin g s on the same
date were taken to Bozeman and handled as described in the procedure.
In
th is second batch of crick ets three groups of f i f t y crickets each were
dusted on three d ifferen t p a rts of the abdomen; the ventral membranes,
the ventral p la te s, and the la te r a l membranes.
Those series dusted on
ventral and la te r a l membranes (se rie s I I I and V) show a marked sim ilarity
in reaction.
I if ty per cent of the crickets in both series reached f i r s t
lo ss of coordination between the f i r s t and second hours a fte r application.
F ifty per cent in both se rie s reached the point o f complete lo ss of
coordination between the second and th ird hours, and 50$6 m ortality in
the two series took place about one h a lf hour ap art, as shown in fig . 2$.
Crickets in series IV were dusted on the ventral p la te s.
This
series took one of the longest o f any in the study to reach f if ty per cent
m ortality.
The reason fo r th is may be ascribed p a r tia lly to the smaller
-5 6 -
series
H
SERIES H
y— s
----- 0 SERES 3E
PERCENT
0
TIME IN HOURS
F ig. 25.
M o rta lity curves fo r a p p lic a tio n in S e r ie s I I I
(v e n tr a l abdominal membranes), S e r ie s IV
(abdominal s t e r n i t e s ) , and s e r ie s V ( la t e r a l
abdominal membranes).
—57a r e a in v o lv e d and th e f a c t t h a t th e d u s t may h e more e a s i l y c la im e d by
g r a v ity i n t h i s p o s i t i o n .
However, th e l a t t e r re a s o n sh o u ld b e a p p lic a b le
to s e r i e s I I I ( th e s e r i e s i n w hich th e v e n t r a l membranes w ere d u s te d ) an d
t h i s s e r i e s r e q u ir e d th e l e a s t amount o f tim e to re a c h th e p o i n t o f $0#
m o rta lity .
I t w ould a p p e a r th e n t h a t th e l a t e r a l and v e n t r a l membranes
p e rm it more r a p i d r e a c t i o n b o th b e c a u se o f d i f f e r e n c e s i n a r e a a n d q u a l i t i e s
o f th e membrane.
The q u a n t i t a t i v e a n a ly s is o f c r i c k e t s i n each o f th e th r e e
s e r i e s , a s shown i n t a b l e T I , r e v e a l s a m inor tr e n d to w ard g r e a t e r
p e n e tr a ti o n among c r i c k e t s w hich l i v e d lo n g e r w ith in t h e i r own g ro u p s.
X com parison o f tim e r e q u ir e d f o r 50$ m o r t a l i t y i n th e th r e e s e r i e s , f i g . 25,
and a v e ra g e amount o f p e n e t r a t i o n t i l l d e a th , t a b l e T I , shows a g a in th e la c k
o f r e l a t i o n s h i p betw een th e tw o.
D if f e r e n c e s i n th e d is ta n c e from v i t a l
o rg a n s a s w e ll a s th e s i z e o f th e a r e a s a r e v e ry p ro b a b ly th e e x p la n a tio n
f o r th e f a c t t h a t c r i c k e t s d u s te d on l a t e r a l abdom inal membranes a b so rb e d
th e m ost p o is o n i n a lm o st th e l e a s t tim e .
The f a c t t h a t th e v e n t r a l
membranes a r e c lo s e to th e m ain n e rv e tr u n k m ig h t e x p la in th e f a c t t h a t
a q u ic k r e a c t i o n was shown to t h e p o is o n (w hich h a s b een c o n s id e re d p a r ­
t i a l l y a n e rv e t i s s u e p o is o n ) , b u t t h a t p e n e t r a t i o n m ight n o t h av e b een
e s p e c ia lly g re a t.
The tim e f o r th e f i r s t r e a c t i o n to th e p o is o n may b e
seen f o r a l l th e s e r i e s i n t a b l e I I I . . F u r th e r r e a c t i o n s may b e se e n i n
t a b l e s IT and T.
BFFBGT OF DUSTING THB THOBAX AND ITS APPENDAGES
From a b a tc h o f c r i c k e t s c o l l e c t e d on th e same day f o u r g ro u p s
o f c r i c k e t s w ere i s o l a t e d and t e s t e d to d e te rm in e th e e f f e c t o f sodium
a r s e n i t e on th e th o r a x and i t s ap p en d ag es.
The f o u r s e r i e s c o n s i s t e d o f
-5 8 -
Table VI.—Quantitative data on penetration
Group
2
45
117
)
307 )
142 )
III
3
:
V
VI
VII
VIII
II
I
3 .15
2.8275 - .1400 .
4
16
2.9939
.375
5
17
2.5297
.750
6
18
3.1015
.375
7
19 - 2.7793
.375
I 24
2.7025
I.125
2
25
2.9779
.750
9
26
2.8086
1.050
10
27
2.9034
1.250
I 28
1.6205
0.250
2 29
1.853
.250
8
30A 1.9625
.250
9
30B I.S196
.250
10
31
2.0666
.175
I . 20
1.7434
0.375
2 21
2.06l6
.250
9 23
2.0776
.175
10 22
1.8752
.175
1 32
2.1941
0.375'
2 33 . 1.8995
.400
9
34
2.4401
.150
I 26
2.4152
. 675
2 37
2.3394
.400
7 38
2.3930
.500
10
39
2.3217
.500
.080
.075
.150
.075
.075
.225
.150
.210
.250
.050
.050
.050
.050
.035
. 075
.050
.035
.035
.075
.080
.030
.135
.080
.100
.100
.142 )
.125 ) .162
.296 )
.121 )
.135 )
.416 )
.252 ) .368
.374 )
.431 )
.154 )
.135 )
.127 ) .159
.137 , )
.085 )
.215 )
.121 ) .128
.084 )
.093; )
.171 ) .
.211 ) .150
.062 )
. 279 )
.171 ) .218
.209 )
.215 )
-5 9 -
applications on the pronotal shield (series V II)« the sides o f the thorax
not including the propleura (se rie s V III), and the ta r s i (se rie s IX).
The application of sodium arsen ite on the pronotal shield was
localized "by the use o f p araffin as a thin edge around the shield.
P araffin was also placed on the tib i a of those in sects on which the ta r s i
were dusted and on the femora o f those having the pleura dusted.
Ho
p araffin was used on those crick ets which were treated with sodium arsen ite
on the membrane beneath the pronotal shield,
,
The m ortality curves of those se rie s on which applications
were made on the membrane beneath the pronotal shield and on the thoracic
pleura show a rath er marked sim ila rity .
The p oints of 50$ m ortality are
only about twenty minutes apart, as may be seen in fig . 26.
These two
series are in sharp contrast with the series VI and IX, crick ets dusted
on the pronotal shield and on the ta r s i.
point for the four curves may be noted.
By looking a t fig . 26 the midU
Series VI and IX are about two
hours apart a t the point of 50$ m ortality.
The reason most plausible
for series IX to have a more f la t m ortality curve than VI may be the
distance ta r s i are found from v ita l centers.
Careful examination o f the
pleura of the thorax and the membrane beneath the pronotal shield reveals
a common facto r.
Both are smooth and o ffer fewer crevices and le ss
opportunity fo r adhesion and subsequent penetration.
The pronotal shield
on the other hand supplies a wide" penetrable surface and therefore yielded
an early 50$ m ortality
Comparison o f the time required for 50$ m ortality with the
quantity of penetration (tab le VI) reveals th a t in the groups te sted no
-6 o -
• ---- • SERIES ST
o— o SERIES IX
PERCENT
MORTALI
e— -e SERIES Vll
e----- « SCRIES
TIME IN HOURS
P ig. 26.
M o rta lity curves fo r s e r ie s VI (p ro n o ta l s h ie ld ) ,
s e r ie s VII (membrane beneath p ro n o ta l s h i e l d ) ,
s e r ie s VIII (p te r o th o ra c ic p le u r a ) , and s e r ie s IX
(ta r s i).
-S i-
r e l a t i o n seemed to e x i s t -"between th e p o in t o f 50$ m o r t a l i t y and th e
r e l a t i v e amount o f sodium a r s e n l t e d e te rm in e d p e r u n i t o f d ry "body w e ig h t.
Q W SBU
j
CONSIDERATIONS
S in c e t h e c r i c k e t s w ere t e s t e d i n g ro u p s which w ere c o l l e c t e d a t t h r e e
d i f f e r e n t d a te s , th e p r o b a b i l i t y e x i s t s t h a t a l l t h r e e g ro u p s d i f f e r e d
from one a n o th e r i n many r e s p e c t s .
Itor t h i s re a s o n th e s e r i e s h av e b een
com pared o n ly w ith o t h e r s e r i e s w hich were c o l l e c t e d a t th e same d a te .
S in c e i t i s o f I n t e r e s t , how ever, to compare a l l th e g ro u p s w ith
each o t h e r some "common den o m in ato r" i s n e c e s s a r y .
The f a c t o r t h a t m ight
be u s e d f o r a t e n t a t i v e com pariso n I n t h i s c a s e was th e t e s t i n g o f a l l
c r i c k e t g ro u p s w ith one ty p e o f a p p l i c a t i o n an d th e n w e ig h tin g t h a t group
w ith th e v a lu e o f i t s 50$ r e a c t i o n p o i n t when com pared w ith t h e v a lu e o f
th e f i r s t group so t e s t e d .
B ie s e r i e s u s e d a s a t e s t s e r i e s was s e r i e s I I .
T ab le V r e v e a ls th e tim e i n h o u rs f o r 50$ m o r t a l i t y d e riv e d i n th e
o rd in a r y manner ( ta k e n from th e m o r t a l i t y c u rv e s , f i g s . 2 ^ -2 6 ) .
was ta k e n a s a b a s i s o f co m p ariso n .
S e r ie s I I
In t h i s s e r i e s th e t e r g l t e e o f th e
abdomen were d u s te d and th e p o i n t o f 50$ m o r t a l i t y , 7 .7 h o u rs c o n s id e re d
a s th e com parison u n i t .
Twenty c r i c k e t s from each su c c e e d in g b a tc h w ere
d u s te d and t r e a t e d e x a c tly a s was s e r i e s I I .
Each 50$ r e a c t i o n p o i n t
C1 , C2 o f t a b l e s I I I , IV , V t h e r e a f t e r c o u ld b e w e lg h te d ^ a c c o rd in g t o th e
v a lu e o b ta in e d i n com parison w ith th e sm all gro u p a c t i n g a s a "common
d e n o m in a to r".
I t m ust be p o in te d o u t t h a t no a tte m p t a tj p r e c is e n e s s i n
com parison i s c la im e d i n t h i s f a s h io n , b u t t h a t a g e n e r a l m ethod o f
com parison may th u s be draw n.
By t h i s m eans, i f m o r t a l i t y I 8 c o n s id e re d ,
—
62—
series I I I , IV, and V might "be given a ratin g of 83$ strength of the f i r s t
two series and the th ird collection "batch might "be rated 104$ in the same
manner.
VII.
The time fo r 50$ m ortality might then "be considered as in table
-62Table V I I ,— Time fo r 50^ m ortality on a comparative basis
Series
Ho.
I
II
III
IV
V
VI
V II
V III
IX
Time fo r 5Qf>
m ortal!ty
8.0
7.7
6.8
13.8
7.4
6.8
12.4
12.0
9.0
Adjusted time
fo r 50$ m ortality
8.0
7.7
5.7
11.6
6.2
7.1
12.9
12.5
9.4
-64COKGLUSIOHS
The most strik in g aspects of the morphology of Anahnxs are the
modifications of the thorax.
The mesothoracic sp iracle presents an unusual,
i f not hith erto unknown, modification in in sect anatomy.
This spiracle may
he of in te re st from the aspect of Mormon crick et control.
The Mormon cricket shows a d efin ite and orderly change in i t s
reactions once penetration of sodium arsen ite begins.
from any of the stages through which i t passes.
I t never recovers
Inspection of tables I I I ,
IV, and V reveals th at in general those series reaching the point of 50$
m ortality also reached the points of 50$ la the several stages of reaction
in a p a ra lle l manner.
Time required for 50$ m ortality and the amount of sodium arsen ite
th at penetrates in th a t time apparently have l i t t l e , i f any, relationship.
Membranes have, in general, a tendency to permit a greater speed of reaction.
This i s modified to a large extent by the proximity of the area to Important
organs, the q u a litie s aiding or hindering adhesion, the position of the
area in regard to i t s action as a b a rrie r to gravity, and the size of the
area.
Thus series I , I I I , V, and IX are e ith e r membranous or contain much
membrane.
They, therefore, showed early reactions to the poison and reach­
ed the point of 50$ m ortality.
Series VI, in which a large sclerotlzed
area, the pronotum, was dusted, reacted slowly a t f i r s t but reached the
point of 50$ m ortality almost as quickly as the speediest o f the membranous
areas.
The smoothly membranous se rie s, series VII, required one of the
longest 50$ m ortality points since i t offered l i t t l e opportunity for
adhesion.
The thoracic pleura and abdominal e te ra ite s which are sclerotlzed.
-6 5 -
and vhlch are rath er smooth, give l i t t l e aid to adhesion, "both required
a lengthy period to reach the point of 50$ m ortality.
Dusting of the
abdominal te rg lte s may have resulted in an early reaction because o f the
size of the areas and the proximity to the main circulatory system.
-6 6 -
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N578
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