THE JOURNAL OF COMPARATIVE NEUROLOGY 409:325–338 (1999)

Thoracic and Prothoracic Leg

Neuromuscular System of the Praying
Mantid, Sphodromantis lineola
(Burmeister)
MOLLY BULLARO AND FREDERICK R. PRETE*
DePaul University, Department of Biological Sciences, Chicago, Illinois 60614

ABSTRACT
Historically, praying mantids have attracted attention because of their dramatic prey
capture behavior, loosely termed the strike. However, little is known about the neuromuscular
organization that underpins the behavior. Although once thought to be quite stereotyped,
recent data indicate that the strike is quite plastic and can be aimed accurately within a
relatively large three-dimensional space. Hence, successful prey capture requires the
integration of (1) visual information, indicating prey has been recognized; and (2) propriocep-
tive information, indicating head and prothorax (i.e., visual field) position and initial leg
positions. This study was undertaken as part of a larger program examining how such sensory
information is integrated with the appropriate motor systems. Our goals were (1) to describe
the gross thoracic and foreleg neuromuscular system of Sphodromantis lineola and (2) to
identify the soma locations of the motor neurons associated with the largest leg nerve, N4,
which travels the length of each leg. We found that the thoracic and foreleg neuromusculature
of S. lineola are similar but not identical to what is known about just three other species of
mantid, and that motor neuron somata associated with N4 are arranged in stereotypical,
bilaterally symmetrical groups as they are in other orthopteroids, suggesting that this is a
general organizational feature of the insect CNS. J. Comp. Neurol. 409:325–338, 1999.
r 1999 Wiley-Liss, Inc.

Indexing terms: insect; orthoptera; ganglia, invertebrate; motor neurons

Historically, the praying mantids have drawn people’s
attention because of their especially large raptorial fore-
legs and dramatic method of prey capture (Prete and
Wolfe, 1992). However, despite this long-standing interest
and the identification of almost 2,000 species, little is
known about their basic biology compared with some other
orthopteroid insects (but see, Prete et al., 1999).
Studies of mantid prey capture have generally focused
on the mechanics or kinematics of one or both of the two
component behaviors: the forward, upward, and/or lateral
displacement of the body by the meso- and metathoracic
legs (the lunge), or the rapid grasping movement of the
raptorial forelegs (the strike) (e.g., Copeland, 1975; Cor-
rette, 1990; Prete et al., 1990; Cleal and Prete, 1996; Prete
and Cleal, 1996). Analyses have revealed that these two
components are independent and variable, depending on
prey location (Copeland, 1975; Cleal and Prete, 1996; Prete
and Cleal, 1996; Prete and Hamilton, 1999). Such plastic-
ity suggests that successful prey capture depends on the
integration of at least three classes of sensory information:
(1) visual information, regarding the characteristics of the
visual stimulus; (2) proprioceptive information, regarding
the relative location of the visual field (i.e., head and
prothorax positions); and (3) proprioceptive information,
regarding initial leg positions.
Little is yet known about the neuromuscular organiza-
tion that underpins mantid prey capture. The only ger-
mane data are contained in three studies, one on the
musculoskeletal system of Stagmomantis carolina (Le-
vereault, 1938), one on the gross thoracic neuroanatomy of
Tenodera aridifolia sinensis (Liske et al., 1989), and one on
the prothoracic leg neuromusculature of Hierodula mem-
branacea, which includes a cursory description of the
Grant sponsor: University Research Council, DePaul University; Grant
sponsor: Faculty Research and Development, DePaul University; Grant
sponsor: Department of Biological Services, DePaul University.
*Correspondence to: Frederick R. Prete, DePaul University, Department
of Biological Sciences, 2325 North Clifton Ave., Chicago, IL 60614.
E-mail: f-prete-1@alumni.uchicago.edu or frprete@ibm.net
Received 26 August 1998; Revised 19 January 1999; Accepted 4 February
1999

r 1999 WILEY-LISS, INC.

326
locations of a few prothoracic motor neurons (Gray and
Mill, 1985).
This study was undertaken as a part of a larger research
program examining the ways in which visual and proprio-
ceptive information are integrated with the motor systems
that control prey capture behavior. Our specific goals were
(1) to describe the gross thoracic and foreleg neuromuscu-
lar system of Sphodromantis lineola and compare that
with what is known about the three other mantid species,
and (2) to identify the soma locations of the motor neurons
associated with the largest leg nerve, N4, which travels
the entire length of each of the mantid’s legs.
MATERIALS AND METHODS
Work was done on healthy, laboratory-reared, adult,
female S. lineola selected arbitrarily from an ongoing
colony (for culturing methods, see Prete and Mahaffey,
1993). Drawings of the gross thoracic neuromusculature
were done by hand based on dissections of 10 specimens
killed with an overdose of CO2 and fixed in 95% ethanol.
Dissections were done in ethanol under a wide-field stereo-
microscope (Nikon) and, when needed, tissue contrast was
enhanced with methylene blue. Drawings based on the
dissections were digitized (SuperVISTA S-12 Scanner,
UMAX DATA), stored to disc (Power Macintosh 7600), and
converted to computer graphics (Canvas 5.0.3, Deneba
Systems, Inc.).
Leg motor neurons were identified by backfilling from
the cut ends of live nerves submerged in wells containing
either cobalt chloride or horseradish peroxidase (HRP).
After set-up, mantids were placed in humid containers
overnight at approximately 20°C. Cobalt chloride prepara-
tions were partially dissected to expose the thoracic gan-
glia and placed in saline (159 mM NaCl, 10 mM KCl, 4.5
mM CaCl2, 0.1 mM NaH2PO, pH 7.2), to which two to three
drops of 20% Na4SO3 were added. After 5 minutes, the
thoracic ganglia and nerves of interest were dissected free
under saline. The tissue was dehydrated in a series of
ethanol baths, cleared in methyl salicylate, and perma-
nently mounted between coverslips. HRP preparations
were partially dissected and fixed in 10% phosphate
buffered formalin (pH 7.4) for at least 2 hours. Thoracic
ganglia and nerves of interest were dissected free and
washed several times in 0.5% Triton X-100 in phosphate
buffered saline (pH 7.4). The tissue was reacted with DAB
(Peroxidase Substrate Kit, Vector Laboratories, Inc., Burl-
ington, CA), washed several times in phosphate buffered
saline, dehydrated in a series of ethanol baths, cleared in
xylene, and permanently mounted between coverslips.
Filled cells were drawn with a camera lucida and/or
appropriately magnified confocal images were digitized
(Power Macintosh 7600; software: VideoShop 3.0.2, Avid
Technology) and stored to disk for subsequent analysis
(software: Canvas; Adobe Photoshop 3.0.5, Adobe Systems,
Inc.). Composite drawings were based on a slide library of
20 preparations.
We have done our best to integrate our anatomic data
with that previously published on mantids. To that end, we
have used numbering schemes that are as consistent as we
could make them with those previously used. Unfortu-
nately, in some cases we have had to choose between
absolute consistency and simplicity, and our choices have
not always been ideal. For the most part, we have followed
the muscle numbering system of Levereault (1938), the
M. BULLARO AND F.R. PRETE
prothoracic leg nerve numbering system of Gray and Mill
(1985), and the thoracic nervous system numbering sys-
tem of Liske et al. (1989).
RESULTS
The thoracic nervous system of the mantid includes
three discrete ganglia—the prothoracic (P or T1), mesotho-
racic (Ms or T2), and metathoracic (Mt or T3) ganglion–
connected by a pair of ventral nerve cords, or connectives.
Five bilaterally symmetrical pairs of nerves emerge from
the lateral edges of each ganglia (N1–N5, numbered from
anterior to posterior, Figs. 1, 2), and a sixth, bilaterally
symmetrical pair emerges from the ventral surface of each
(N6, Fig. 9). In addition, an unpaired median nerve
emerges from the posterior, dorsal surface of each ganglion
and lies between the two connectives (PN7, Fig. 1A; MsN7,
MtN7c, Fig. 2).
The first three abdominal ganglia (A1–3) are fused with
the metathoracic ganglion, and give rise to three addi-
tional pairs of bilaterally symmetrical nerves (MtN8a-c),
which emerge from the posterior lateral region of the
ganglion (Fig. 2). Fusions of these four ganglia have been
found in all Mantidae studied (Nesbitt, 1941; Gray and
Mill, 1985; Liske et al., 1989). Two posterior dorsal median
unpaired nerves that serve the first three segments of the
abdomen (MtN7a, MtN7b, Fig. 2) emerge from the dorsal
midline of the metathoracic ganglion.
Major nerves and innervation patterns
Table 1 summarizes the ganglion of origin and muscles
innervated by the main thoracic nerves, and Table 2
compares the numbering systems for the prothoracic nerves
used by Gray and Mill (1985) for H. membranacea, Liske et
al. (1989) for T. a. sinensis, and that used here.
Nerve 1. Nerves 1 (N1) of the pro-, meso-, and
metathoracic ganglia have similar distribution patterns in
each segment. Each nerve has both anterior and posterior
branches some of which fuse with nerves of the same or of a
different thoracic ganglion (Fig. 1).
Prothoracic ganglion. PN1 sends one branch anteri-
orly (PN1A), which fuses with the ipsilateral connective
just before the connective enters the head capsule (Fig. 1A,
arrow 1). Anterior to that point, a smaller (probably
recurrent) nerve from the subesophogeal ganglion emerges
from the connective and innervates the ventral rotator of
the head (Rn-SOG). In H. membranacea and T. a. sinensis,
PN1A fuses with a recurrent nerve from the subesophogeal
ganglion rather than with the connective (Gray and Mill,
1985; Liske et al., 1989). A posterior branch, PN1B, gives
rise to PN1B1 and PN1B2, which project anteriorly, and
PN1B3, which projects posteriorly. Nerve PN1B1 gives
rise to PN1B1a which fuses with a branch of PN2 (Fig. 1A,
arrow 2). Nerve PN1B1b emerges rostral to PN1B1a (Fig.
1B), travels medially and then posteriorly, and fuses with
the anterior branch of MsN1B and a lateral branch of PN7
(Fig. 1A, arrow 3). Nerve PN1B1 continues to project
anteriorly and innervates the protractor of the posterior
lateral cervical sclerite, retractor of the anterior lateral
cervical sclerite, and the lateral dorsal rotator of the head
(Liske et al., 1989). Nerve PN1B2 innervates the muscles
of the thoracic spiracle (an opening in the cuticle continu-
ous with a trachea; Fig. 1A,B). Branch PN1B3 innervates
the tergal levator and tergal protractor of the prothorax
(Liske et al., 1989; Fig. 1A,B).
THORACIC NEUROMUSCULAR SYSTEM OF S. lineola 327

Fig. 1. All figures represent female Sphodromantis lineola. ing scheme based on Liske et al. (1989). P, prothoracic; Ms, mesotho-
A: Prothoracic ganglion and associated nerves (dorsal view); racic; Mt, metathoracis; N, nerve; Rn-SOG, recurrent nerve of the
B,C,D: Enlargements of PN1, PN2, and PN3, respectively (dorsal subesophageal ganglion. Scale bar ⫽ 0.5 mm.
view). Arrows 1–4 indicate plexuses. Thoracic nervous system number-
 Fig. 2. Meso- and metathoracic ganglia and associated nerves (dorsal view). Arrows 1–5 indicate
plexuses. Conventions as in Figure 1. Scale bar ⫽ 0.5 mm.
THORACIC NEUROMUSCULAR SYSTEM OF S. lineola 329

TABLE 1. Ganglion of Origin and Muscles Innervated by the Main

Thoracic Nerves in Sphodromantis lineola1
Ganglion Muscles
Prothoracic
PN1 2, 3, 7a–b, 35, 37
PN2 1a–b, 9a–c, 19a–b, 26
PN3 4, 11, 9a, 23b
PN4 23a, 24, 25, 28a–b, 29, 31a–b, 32, 33, 34a–b
PN5 10a–b, 12, 15, 16, 17, 20, 24, 26
PN6 Target not identified
Mesothoracic
MsN1 36, 39, 40, 41, 52, 53, 54
MsN2 48, 61, 62a–b, 64
MsN3 60, 66, 71b
MsN4 71a, 72, 73, 77, 79, 80, 81, 82
MsN5 49, 50, 51, 65, 70
MsN6 Target not identified
Metathoracic
MtN1 55, 57, 58, 89, 90
MtN2 94, 95a–c, 96
MtN3 93, 98, 101, 103b
MtN4 104, 105, 109, 111, 112, 113, 114
MtN5 94, 95, 96
MtN6 Target not identified
1Muscles numbered per Levereault (1939). P, prothoracic; Ms, mesothoracic; Mt,

metathoracic; N, nerve.

TABLE 2. Comparison of the Numbering Systems Used for the

Prothoracic Nerves of Sphodromantis lineola, Tenodera aridifolia sinensis
(Liske et al., 1989), and Hierodula membranacea (Gray and Mill, 1985)1
S. lineola T. a. sinensis H. membranacea
PN1 Pro N1 N1 Fig. 3. Right prothoracic leg (ventral view to show details of
PN2 Pro N2 N2 femoral spines). A: Coxa; B: trochanter; C: femur; D: femoral-tibial
PN3 Pro N3 N4 joint; E: tibia; F: tarsus. Adapted from Roy (1999), with permission
PN4 Pro N4 N5A
PN5 Pro N5 N6
from The Johns Hopkins University Press. Scale bar ⫽ 5 mm.
PN6 Pro N6 N3
PN7 Pro N7 Not identified
Not identified N/A N5F8
Not identified N/A N5F9 Prothoracic and prothoracic leg musculature. The
PN4E1 N/A Not identified
PN4E2 N/A Not identified prothoracic leg includes six segments, the coxa, which is
PN4D1a N/A N5D2–7 connected to the thorax, the trochanter, the femur, the
1P, prothoracic; Ms, mesothoracic; Mt, metathoracic; N, nerve; N/A, not applicable; Pro, femoral-tibial joint, the tibia, and the tarsus (Fig. 3A–F,
prothoracic. respectively). The thoracic muscles associated with the leg
and the muscles of the prothoracic-coxal joint in S. lineola
are similar to those in other mantids (Levereault, 1938;
Gray and Mill, 1985; Liske et al., 1989), and, likewise, can
Mesothoracic ganglion. A number of plexuses occur in be classified as promotors, remotors, abductors, adductors,
the posterior prothorax. The fusions include (1) MsN1A anterior rotators, or posterior rotators (Fig. 4).
with the ipsilateral connective (Fig. 1A, arrow 4), (2) Promotors. The tergal coxal promotor (19b on Fig. 4)
MsN1A2 with MsN1B (Fig. 2, arrow 4), (3) MsN1B with and pleural coxal promotor (19a on Fig. 4) move the coxa
the ipsilateral branch of PN7 and recurrent nerve PN1B1b anteriorly. They originate on the anterior lateral portion of
(Fig. 1A, arrow 3), and (4) nerve MsN1A2 with MsN1A1b the tergum and on the anterior surface of the episternum,
(Fig. 2, arrow 2). Nerves MsN1B1 and MsN1B2 innervate respectively, and insert on the anterior edge of the coxal
the sternal levator of the prothorax and the sternal rim. Electrophysiologic recordings done in our laboratory
occlusor, and the dilator of the thoracic spiracles, respec- confirmed that these promoters fire if, and only if, a strike
tively. Branch MsN1C bifurcates and innervates the dorsal occurs (Gonka, Laurie, and Prete, manuscript submitted
oblique muscles that extend from the metathoracic tergum for publication).
to the posterior area of the mesothoracic tergum (Fig. 2; Remotors. The lateral and posterior coxal remotors
Levereault, 1938; Liske et al., 1989). are the primary remotors of the coxa. The former (not
Metathoracic ganglion. Another plexus occurs in the shown) originate on the lateral tergum, posterior to the
posterior mesothorax: nerve MtN1A fuses with the ipsilat- supracoxal inflexion (S on Fig. 4). The posterior coxal
eral connective (Fig. 2, arrow 3), and MtN1A2, MtN1B1, remoter (10b on Fig. 4) originates on the midlateral
and recurrent nerve MsN1A2 fuse with the ipsilateral tergum. As in H. membranacea, these remotors insert on a
branch of MsN7 (Fig. 2, arrow 1). tendon that originates from the posterior-lateral portion of
The main targets of the second branch of MtN1B are the the basal coxal rim.
tergal protractor of the mesothorax, and the dilator and Abductors and adductors. Four adductors and four
occlusor of the second thoracic spiracle (Liske et al., 1989). abductors insert on the trochantin. The latter function as
Nerve MtN1C innervates the retractors of the first abdomi- slow coxa promotors in mantids (Gray and Mill, 1985). The
nal tergum, the dorsal oblique muscles, which extend from anterior coxal abductor (16 on Fig. 4) originates on the
the metathoracic tergum to the anterior lateral region of lateral portion of the tergum, behind the supracoxal
the first abdominal tergum. The tiny branch MtN1C2 infolding and the intermediate coxal abductor (not shown)
fuses with MtN2B1 (Fig. 2, arrow 5). originates posterior to it. The posterior lateral coxal abduc-
330
Fig. 4. Prothoracic muscles associated with the prothoracic leg and
the innervation patterns of PN2. For clarity, the prefix ‘‘PN2’’ has been
eliminated from the body of this drawing, and in Figures 4–8, lines
representing nerves are 1.5⫻ thicker than they appear. Muscle
tor (14 on Fig. 4) lies very close to the intermediate
abductor (making them difficult to distinguish), and the
posterior medial coxal abductor (17 on Fig. 4) originates on
the medial lateral tergum. Each of these muscles insert on
the proximal tip of the basal coxal rim behind the coxal
articulation. A fifth pleural coxal abductor (20 on Fig. 4)
inserts posterior to the supracoxal infolding and medial to
the anterior abductor.
Three muscles primarily responsible for adduction of the
coxa insert on the tip of the trochantin. The anterior tergal
coxal adductor (9a on Fig. 4) originates from the median
tergum anterior to the supracoxal infolding, the intermedi-
ate tergal coxal adductor (9b on Fig. 4) originates on the
median tergum, and the posterior tergal coxal adductor (9c
on Fig. 4) originates posterior to the intermediate adduc-
tor. The fourth sternal coxal adductor originates on the
ventral furcal apophysis and inserts on the anterior basal
coxal rim (not shown).
Rotators. The anterior coxal rotator (not shown) origi-
nates on the tip of the furcasternite and inserts on the
dorsal coxal rim. The tergal posterior coxal rotator (13 on
Fig. 4) originates on the mid-lateral tergum posterior to
the leg base and inserts on the posterior coxal rim. The
sternal posterior coxal rotator (not shown) originates on
the dorsal region of the furcal apophysis and also inserts
on the posterior coxal rim.
Nerve 2. Nerve 2 (N2), second in size only to N4,
emerges ventral-laterally from each thoracic ganglia. It
has two main branches both of which are mixed nerves.
The first runs anteriorly and dorsal-laterally, and the
second runs posteriorly and ventral-laterally and travels
toward the leg. The branching patterns of N2 are the same
in all three thoracic segments (Figs. 1A,C, 4).
M. BULLARO AND F.R. PRETE
numbers (bold type) are based on the scheme of Levereault (1938), and
prothoracic leg nerve numbers are based on the scheme of Gray and
Mill (1985). Scale bar ⫽ 2 mm.
Prothoracic ganglion. The anterior branch PN2B gives
rise to PN2B1, PN2B2, and PN2C. The latter gives rise to
PN2C1 in the rostral prothorax, which innervates the
lateral dorsal rotator of the head and carries afferents from
the cervical hairplates. Branch PN2B1 travels anterior-
laterally and gives rise to PN2B1a-c and PN2B1f. Nerve
PN2B1a and b travel dorsally and innervate the pleural
coxal promotor (19a on Fig. 4) and tergal coxal promotor
(19b on Fig. 4), respectively. Nerve PN2B1c travels anteri-
orly and bifurcates into PN2B1d and e, which innervate
the ventral tergal sensilla and the tergal coxal promotor,
respectively. In some cases, PN2B2 arises from the same
point as does PN2C, but in the other cases, it arises more
distally as shown in Figure 4.
This variability also occurs in T. a. sinensis (Liske et al.,
1989). Nerve PN2B2 travels anterior-laterally (behind 9a
in Fig. 4), and then posteriorly giving off a number of small
branches that innervate the anterior tergal coxal adductor
(9a on Fig. 4), the medial and lateral tergum, the interme-
diate tergal coxal adductor (9b on Fig. 4), and the posterior
tergal coxal adductor (9c on Fig. 4). Liske, et al. (1989)
described a ‘‘loop formation’’ in T. a. sinensis in which one
branch of PN2 fused both with itself and PN5. This was not
seen in S. lineola.
Nerve PN2A, travels ventral-laterally into the coxal
cavity and gives off several mixed branches. Nerves
PN2A1–3, innervate dorsal coxal cuticular sensilla and
PN2A4–6 innervate the trochanteral flexor (26 on Fig. 5)
(Fig. 5A,B). Nerve PN2A4a, which fuses with PN2A, has
not been seen in other mantids. Finally, PN2A7 travels to
the distal coxa and fuses with PN4 (Fig. 5A,B arrows 1).
Meso- and metathoracic ganglion. The branching pat-
terns of MsN2 and MtN2 are similar to those in the prothorax
THORACIC NEUROMUSCULAR SYSTEM OF S. lineola 331

Fig. 5. Muscles and associated nerves of the left procoxa (dorsal view). A: Prothoracic nerves (PN)2A
and PN3A. B: Nerves PN4 and PN5A. The prefix ‘‘PN’’ has been eliminated from the body of the drawings.
Scale bar ⫽ 2 mm.

(Figs. 1A, 2). We have only tentatively identified the
targets of N2 in the meso- and metathorax, but they
appear to be the same as those in T. a. sinensis. The
anterior branch, MsN2B, gives rise to MsN2B1–2 and
MsN2C, which innervate the tergosternal muscle of the
mesothorax, epipleuritellar productor of the pleurellite,
and the tergal abductor of the pleurellite. Nerve MtN2B
innervates the tergal abductor of the pleurellite and the
tergal retractor of the epipleurite. These four nerves also
innervate the promotor of the coxa in T. a. sinensis, but we
did not see this in S. lineola (Liske et al., 1989). In both
the meso- and metathoracic ganglion, MsN2A travels
toward the coxal cavity and ultimately fuses with N4 in
the distal coxa. We saw a fusion between MtN1C2 and
MtN2B1, which has not been reported in other species
(Fig. 2, arrow 5).
332
Fig. 6. Muscles of the left trochanter and associated innervation patterns of prothoracic nerve (PN) 4
(dorsal view). The prefix ‘‘PN4’’ has been eliminated from the body of the drawing. Scale bar ⫽ 1 mm.
In T. a. sinensis, N2 exits all three thoracic ganglia from
points corresponding to 10 and 2 o’clock (Liske et al.,
1989). However, in S. lineola, MtN2 exits the ganglion
from positions corresponding to 9 and 3 o’clock.
Nerve 3. Nerve 3 (N3) is small compared with other
laterally branching primary thoracic nerves. It emerges
ventral-laterally, bifurcates almost immediately, and both
branches travel toward the coxa (Figs. 1A,D, 5).
Prothoracic ganglion. Anterior branch PN3B turns
dorsally and innervates the anterior sternal rotator and
the lateral rotator of the head (not shown). Posterior
branch PN3A enters the coxa. Nerve PN3A1 innervates
the proximal regions of the thoracic trochanteral extensors
(23b on Fig. 5), and PN3A2a and b innervate the distal
thoracic trochanteral extensor and anterior tergal coxal
adductor (9a on Fig. 4), respectively. According to Liske et
al. (1989), the second branch of PN3A also innervates the
sternal adductor of the coxa in T. a. sinensis, but we did not
see it do so in S. lineola.
Meso- and metathoracic ganglion. The branching pat-
tern of N3 is similar in all thoracic segments. Again, the
meso- and metathoracic targets appear to be the same as
in T. a. sinensis (Liske et al., 1989). Branches MsN3A and
B innervate the sternal productor of the pleurellite, the
anterior rotator of the coxa, and the thoracic trochanteral
extensor. Similarly, MtN3A and B innervate the sternal
productor of the pleurellite, the anterior rotator of the
coxa, the thoracic trochanteral extensors, and the adduc-
tor of the coxa.
Nerve 4. Nerve 4 (N4) is the largest nerve emanating
from the thoracic ganglia, and it is the major nerve of the
leg. It emerges dorsal-laterally and posteriorly and runs
ventral-laterally. Before entering the coxa, it gives rise to a
number of small branches the targets of which are not
known. As noted, in the distal coxa N4 fuses with the
terminal branch of N2 in each thoracic segment (Figs. 1A,
2, 5B).
M. BULLARO AND F.R. PRETE
Prothoracic ganglion. Just after entering the coxa,
PN4 gives rise to PN4A1, which innervates sensilla in the
proximal ventral cuticle. Branches PN4A2 and 3 innervate
the coxal trochanteral extensor (23a on Fig. 5), which
originates on the ventral coxa and inserts on the distal
extensor apodeme. Nerve PN4A3b1 innervates the poste-
rior accessory trochanteral extensor (not shown), a small
muscle that originates on the distal ventral coxa and
inserts on the base of the main trochanteral extensor
apodeme. Nerve PN4A3c1 innervates the anterior coxal
trochanteral extensor (24 on Fig. 5), which has the same
origin and insertion as the posterior accessory trochan-
teral extensor.
As PN4 travels toward the trochanter, it gives rise to
PN4B and C (Fig. 6). Nerves PN4B1 and 2 innervate the
anterior trochanter cuticular sensilla, and the femoral-
tibial chordotonal organs, respectively. Nerve PN4C1 trav-
els toward the posterior trochanter and coxal-trochanter
joint. It gives off a number of as yet unidentified small
branches which appear to innervate cuticular sensilla.
Nerve PN4C2 travels through the trochanter and termi-
nates proximal to the femur. It innervates the femoral
reductor (28a, 28b on Fig. 6) which originates on the
anterior ventral trochanter and inserts on the anterior
basal rim of the femur. This muscle is identified as a
femoral remoter in S. carolina (Levereault, 1939), but is
considered to be a reductor in H. membranacea and S.
lineola. Finally, PN4C3 innervates the trochanteral tibial
flexor (31 on Fig. 6), a muscle not described in S. carolina.
It originates on the anterior wall of the trochanter and
inserts on the flexor apodeme of the tibia.
Nerve PN4B extends into the femur (Fig. 7). Branch
PN4B3 innervates sensilla on the proximal femur, perhaps
those associated with the movable femoral spines (e.g.,
Prete, 1990). Nerve PN4C also extends into the femur and
fuses with PN4B just distal to PN4B3 (Fig. 7, arrow 1).
After the fusion, the nerve is called PN4D (Gray and Mill,
THORACIC NEUROMUSCULAR SYSTEM OF S. lineola
Fig. 7. Muscles of the left profemur and associated innervation patterns of prothoracic nerves (PN) 4B
and C (dorsal view). The prefix ‘‘PN4’’ has been eliminated from the body of the drawing. Scale bar ⫽
2 mm.
1985). Branch, PN4C4, emerges just after the fusion and
travels anteriorly. In H. membranacea, it gives rise to a
number of small sensory nerves at its distal end (Gray and
Mill, 1985). However, we were only able to identify one
consistently, PN4C4a. In addition, PN4C4 yields a number
of small branches (e.g., PN4C4b, PN4C4b1, PN4C4c) that
innervate the distal tibial extensor (29 on Fig. 7) which
originates on the posterior femur and inserts on the flexor
apodeme. A more proximal branch, PN4C5, travels ven-
trally through the tibial extensor and innervates the
femoral pretarsal flexor (not shown), which originates on
the proximal, ventral femur.
As noted, the fusion of PN4B and PN4C gives rise to
PN4D which, in turn, gives rise to a number of mixed
nerves that travel throughout the femur. The largest,
PN4D1, runs along the ventral femur through the parallel
and pennate tibial flexors. The parallel flexor (31b on Fig.
7) originates on the proximal ventral femur and inserts on
the distal ventral flexor apodeme. The pennate flexor (31a
on Fig. 7) originates from the ventral femur and inserts on
that portion of the apodeme not occupied by the parallel
fibers (Fig. 7).
Nerve PN4D1a1 gives rise to seven thin branches (PN4D
2–7 of Gray and Mill, 1985) that innervate cuticular
sensilla and, in H. membranacea, the pennate tibial flexor
muscle (31a on Fig. 7). Nerves PN4D1a2 and PN4D1a3
innervate the distal regions of the parallel and pennate
femoral tibial flexors, respectively. More proximately,
PN4D1b innervates the parallel femoral tibial flexor.
Before entering the femoral-tibial joint, PN4D1a gives rise
to PN4D1a4, which innervates cuticular sensilla (Fig. 8A).
At the mid-point of the femoral-tibial joint, PN4D1a
bifurcates into PN4E and PN4F both of which travel into
the tibia (Fig. 8A). This differs from H. membranacea in
which PN4E and PN4F arise in the distal femur (Gray and
Mill, 1985).
Nerve PN4E is a sensory nerve that travels unbranched
through the femoral-tibial joint. Again, this differs from H.
333
membranacea, in which branches of PN4E innervate
muscles at the proximal (but not the distal) region of the
tibia (Gray and Mill, 1985). Nerve PN4F is a mixed nerve
with only one fine branch, PN4F1, that innervates the
proximal tibial pretarsal flexor (34b on Fig. 8), which
originates on the proximal, dorsal tibia apodeme and
inserts on the ventral joint (Fig. 8A). Once in the tibia,
PN4E travels toward the tarsus giving off a number of fine
branches (not shown) the targets of which have not been
determined. In the distal tibia, it gives rise to PN4E1 and
PN4E2 which, with PN4E, innervate dorsal and anterior
tibia cuticular sensilla (Fig. 8B).
Nerve PN4F sends a number of small mixed nerves
throughout the tibia. It also gives rise to PN4F2 in the
proximal tibia which innervates anterior tibia sensilla
(Fig. 8B). Nerve PN4F3 innervates the distal tibial pretar-
sal flexor (34a on Fig. 8), which originates on the distal
anterior tibia and inserts on the apodeme of the unguitrac-
tor plate.
Nerve PN4F gives rise to several small nerves: PN4F5a
and b, PN4F6a and b, and PN4F7a and b (Fig. 8B).
Branches labeled ‘‘a’’ innervate the tarsal extensor (32 on
Fig. 8), which originates on the dorsal distal tibia and
inserts on the dorsal, basal tarsal rim. Branches labeled ’b’
innervate the tarsal flexor (33 on Fig. 8), which originates
in the ventral anterior tibia and inserts on the ventral,
basal tarsal rim. These patterns vary slightly from those in
H. membranacea (Gray and Mill, 1985): We found that
PN4E1 and PN4E2 innervate the tarsal extensor and
flexor muscles which they do not in H. membranacea, and
two nerves that were found in H. membranacea, PN5F8
and PN5F9, were not found in S. lineola. Finally, PN4F4
extends into the tarsus which has no muscles. Hence, it is
assumed the nerve is strictly sensory (Gray and Mill,
1985).
Meso- and metathoracic ganglion. The branching pat-
terns N4 and muscle targets within the leg are the same in
334
Fig. 8. A,B: Muscles and innervation patterns of prothoracic nerves (PN) 4D1a and PN4E and F in the
left profemoral-tibial joint and protibia, respectively (dorsal view). The prefix ‘‘PN4’’ has been eliminated
from the body of the drawings. Scale bars ⫽ 2 mm.
each thoracic segment. Table 1 indicates the specific
muscles innervated by MsN4 and MtN4.
Nerve 5. Nerve 5 (N5) emerges ventral-laterally and
posteriorly from the thoracic ganglia. Branch N5A is a
mixed nerve that travels into the coxa, and N5B gives rise
to nerves that innervate the coxal, tergo-pleural, and
pleural-sternal muscles (Figs. 1A, 2, 5B).
Prothoracic ganglion. Nerve branches PN5A1–3 inner-
vate the sternal posterior rotator of the coxa (not shown),
cuticular sensilla in the thoracic-coxal joint, and the
trochanteral flexor (26 on Fig. 5), respectively (Fig 5B).
More distally, a number of fine unnamed branches also
innervate the trochanteral flexor. The terminal portion of
the main nerve (PN5A4 of Gray and Mill, 1985) gives rise
to several branches that innervate the anterior accessory
trochanteral extensor (24 on Fig. 5). Liske et al. (1989)
described a fusion between a branch of the anterior portion
of PN5A and PN2 in T. a. sinensis, but we did not see this in
S. lineola. Branch PN5B runs dorsal-posteriorly from the
ganglion (Fig. 1A), bifurcates, and then gives rise to a
number of nerves that innervate the coxal remotors,
rotators, abductors, and the tergopleurellar muscles as
they do in H. membranacea (Gray and Mill, 1985).
Meso- and metathoracic ganglia. Nerve 5 has similar
distribution patterns in the meso- and metathorax but we
have not unequivocally identified its targets. They appear,
however, to be the same as those in T. a. sinensis: MsN5A
and B innervate the flexor of the wing, tergopleural
muscles, remoter of the coxa, and depressor of the wing;
MtN5A and B innervate the epipleuritellar productor of
the pleurellite, tergal abductors, and the promotor of the
coxa (Liske et al., 1989).
Nerve 6.
Pro-, meso-, and metathoracic ganglia. Nerve 6 (N6) is
a short nerve originating midlaterally on the ventral
surface of the thoracic ganglia (Fig. 9). The anterior
branch, N6A, travels ventrally toward the basisternite
M. BULLARO AND F.R. PRETE
giving off a number of smaller branches; the posterior
branch, N6B, travels toward the furcasternite. Extensive
work on this nerve has been done by the Yager lab. All of
the peripheral branches of N6 carry information from
cuticular mechanoreceptors. The posterior branch inner-
vates hairs of the basisternum, preepisternum, and, espe-
cially, the membranous cuticle at the junction of the body
segments (for leg position proprioception) (Yager, personal
communication). In the metathorax there is little membra-
nous cuticle because of the modifications associated with
the ear and, there, the posterior branch carries auditory
information (Yager and Hoy, 1987; Yager and Scaffidi,
1993; Yager, 1996; Yager, personal communication). Al-
though N6 is probably just sensory (Liske et al., 1989;
Yager, personal communication), we tentatively identified
two very small branches of N6B (not shown) that appear to
innervate an unidentified ventral thoracic muscle.
Nerve 7.
Pro- and mesothoracic ganglion. Nerve N7 (N7) origi-
nates dorsally from the posterior end of the first two
thoracic ganglia (Figs. 1A, 2). After leaving the ganglion, it
gives rise to two branches each of which fuse with ipsilat-
eral branches of N1 from the same and the next posterior
ganglion (Fig. 1A, arrow 3; Fig. 2, arrow, 1). Nerve PN7
was not identified in H. membranacea (Gray and Mill,
1985).
Metathoracic ganglion. In mantids, abdominal gan-
glia 1–3 are fused with the metathoracic ganglion. Unfortu-
nately, all nerves emanating from this compound ganglion
have been named metathoracic nerves and, for the sake of
clarity, we have retained this nomenclature. Three median
unpaired nerves are associated with the metathoracic
ganglion (Fig. 2). Nerve MtN7c is similar to PN7 and
MsN7 in that it emerges mid-dorsally from the posterior
ganglion between the two ventral connectives but it projects
posteriorly into the anterior fourth abdominal ganglion.
Nerves MtN7a and b give rise to a pair of lateral branches
THORACIC NEUROMUSCULAR SYSTEM OF S. lineola
Fig. 9. A–C: Nerve 6 of the pro- (T1), meso- (T2), and metathoracic (T3) ganglion, respectively (ventral
view). Conventions as in Figure 1. Scale bar ⫽ 0.5 mm.
Fig. 10. A: Photograph of horseradish peroxidase–stained sensory
afferents projecting from the left mesothoracic leg into the mesotho-
racic ganglion through nerves MsN2A and MsN4 (dorsal view).
that supply the first abdominal segment. Branches MtN7b1
and 2 fuse with the ipsilateral MtN8c (described below)
posterior to the fourth abdominal ganglion (not shown). In
contrast, Mt7b1 and Mt7b2 fuse with the ipsilateral
MtN8b in T. a. sinensis (Liske et al., 1989).
Nerve 8.
Metathoracic ganglion. Fusion of the first three ab-
dominal ganglia with the metathoracic ganglion results in
three additional pairs of lateral nerves, MtN8a-c (Fig. 2).
These supply the first three abdominal segments. In T. a.
sinensis, all three nerves emerge from the metathoracic
ganglion (Liske et al., 1989). However, in S. lineola, MtN8c
emerges from the ventral connectives posterior to the
ganglion.
Location of N4 motor neuron somata and leg
sensory afferents
Backfills of N4 consistently revealed a single, stereotypi-
cal, bilaterally symmetrical cluster of neurons (the N4
335
Similarly organized afferents were seen in all three thoracic ganglia
(see Fig. 11). B: Camera lucida drawing of the afferents pictured in A.
Conventions as in Figure 1. Scale bar ⫽ 0.5 mm.
motor group) in each of the three thoracic ganglia similar
to that in the metathoracic ganglion of the grasshopper
(Siegler and Pousman, 1990a,b). The somata are in close
proximity to one another in the ventral cortex and their
primary neurites enter the neuropil (more dorsally) in a
discrete bundle that has a characteristic shape and loca-
tion in each ganglion. Each group of neurons and its
primary neurite bundle were unambiguously identifiable
across individuals.
Backfills of both N2A and N4 also revealed the sensory
afferents traveling through these nerves from the legs
(Fig. 10A,B). Arrangements of the afferent terminal
branches were similar in each ganglion and consistent
across individuals. The location and extent of the area
covered by the afferent terminals within each ganglia
suggest that they intermingle with, and synapse on the
neurites of the N4 motor groups, providing a point for
sensory-motor integration. Dashed lines in Figure 11A,C,G
outline the N2A-N4 afferent terminals in each ganglia,
336
Fig. 11. Camera lucida drawings of N4 motor group cells in
the thoracic ganglia (dorsal views). A,B: Prothoracic ganglion (T1).
C–F: Mesothoracic ganglion (T2). G,H: Metathoracic ganglion (T3).
Arrows 1 (A–E, G–H): The approximate points at which the primary
neurite bundles enter the neuropil. Arrows 2 (D, E): The medial-most
M. BULLARO AND F.R. PRETE
members of the N4 group in T2; F: Camera lucida drawing of the
medial-most members of the N4 group stained with horseradish
peroxidase; dashed lines in A, C, and G: The area covered by the leg
sensory afferent terminals in each ganglion (see Fig. 10). Scale bar ⫽
0.5 mm in G,H (applies to A–H).
THORACIC NEUROMUSCULAR SYSTEM OF S. lineola
suggesting the point at which the N4 motor group neurite
bundles invade the neuropil (Fig. 11A–E,G,H, arrow 1).
Prothoracic ganglion. The PN4 motor group con-
tains approximately 20 cell bodies, although there may be
more. Siegler and Pousman (1990b) reported approxi-
mately 24 somata in what appears to be the comparable
group in the grasshopper metathoracic ganglia. Figure
11A,B are drawings of representative backfills depicting
the somata locations of approximately half of the cells in
the group. As did Siegler and Pousman (1990a,b), we
identified these cells as members of the same group based
on the characteristic shape and location of their primary
neurite bundle. The primary neurites of the group travel
dorsal-medially and, after entering the neuropil, bend
posterior-laterally and exit the ganglion through anterior
PN4 (Fig. 11B arrow 3).
The sizes of the somata in the PN4 motor group vary
considerably and, as in the grasshopper, the locations of
the largest cells were more consistent than were those of
the smaller (Siegler and Pousman, 1990a,b). Although we
have not as yet identified the specific targets of individual
cells, based on the anatomy of PN4, they must innervate
the muscles identified in Table 1.
Mesothoracic ganglion. The characteristics of the
MsN4 motor group are virtually the same as those of the
PN4 group. Again, we identified approximately 20 cell
bodies of varying sizes, the positions of the largest cells
were most consistent, and all primary neurites gathered
into a distinctively shaped and consistently positioned
bundle that projected through anterior N4 (Fig. 11C–E
depict representative backfills). However, each MsN4 mo-
tor group contains one cell consistently positioned more
posterior-medially than the rest (Fig. 11D,E, arrow 2).
Figure 11F shows a pair of these cells backfilled with HRP
in a single mantid. Note that their neurites branch within
the area occupied by the N2A-N4 sensory afferent termi-
nals.
Metathoracic ganglion. Data collected on the meta-
thoracic ganglion were consistent with those on the rostral
two except that for an unknown reason, we found it more
difficult to backfill into this ganglion even within the same
animal. Hence, we were not able to identify consistently as
many somata in the MtN4 motor group as we were in the
pro- and mesothoracic groups (only approximately 15).
However, we assume that the number will turn out to be
comparable to the other two groups and, in all other
respects, the MtN4 group appeared to be similar to the
PN4 and MsN4 groups (Fig. 11G,H).
DISCUSSION
Nesbitt (1941) originally divided the orthopteroid orders
into two groups based largely on the differences in their
somatogastic nervous systems. One group included the
Blattidae (roaches), Mantidae (mantids), and Isoptera
(termites), and the other included the Phasmatidae (walk-
ing sticks), Grylloblattidae (crickets), and Saltatoria (grass-
hoppers). Generally, this division is still acknowledged
although not without some controversy. More recently,
these two presumably monophyletic groups—the Blattop-
teriformia and the Orthopteroidea, respectively—have
come to include, respectively, the Notoptera (rock crawl-
ers), Dermaptera (earwigs), Isoptera (termites), Blattodea
(cockroaches), and Mantodea (mantids), on the one hand,
and the Caelifera (grasshoppers), Ensifera (katydids and
337
crickets), and Phasmatodea (stick insects), on the other
hand (e.g., Hennig, 1981; Baccetti, 1987; but also see
Boudreaux, 1979; Vickery and Kevan, 1985; Kristensen,
1995). Although the relationship between the rock crawl-
ers and earwigs, and the termites, cockroaches, and man-
tids remain unclear, the latter three constitute a seem-
ingly well-founded group, the Blattopteroidea (order
Dictyoptera) (Hennig, 1981; Balderson, 1991; Rentz, 1991).
Although the taxonomic relationships within the Blattop-
teroidea is still controversial, molecular techniques sug-
gest that mantids and roaches are sister groups (Kambham-
pati, 1995) that shared a common ancestor some 30 million
years ago (Gillot, 1980; Balderson, 1991); the precise
status of termites is still unresolved (but see Thorne and
Carpenter, 1992; Kristensen, 1995).
Neuroanatomic data collected on the orthopteroid or-
ders, including that presented here, demonstrate that a
number of fundamental organizational characteristics have
been conserved across the taxa. For instance, in grasshop-
pers, crickets, cockroaches, and mantids each of the three
thoracic ganglia give rise to five lateral nerves divisible
into two groups. The anterior group (N1 and N2) inner-
vates the tergal promotors, anterior rotators, the pleural
abductors, the tergosternal muscles, the basalar/subalar
sclerites, and the axillary muscles in the meso- and
metathorax. The posterior group (N3–N5) innervates the
tergal remotors, the sternal remotors or posterior rotators,
the sternal adductors, and the ventral longitudinal muscles
(Nesbitt, 1941; Gray and Mill, 1985; Liske et al., 1989).
The single exception to this organization is that in locusts
the posterior group innervates the sternal adductors (Gray
and Mill, 1985).
A number of plexuses posterior to the pro- and mesotho-
racic ganglia appear in grasshoppers, katydids, crickets,
cockroaches, walking sticks, termites, and mantids. Fur-
ther, the recurrent nerves that interconnect each of the
thoracic ganglia are present in most orthopteroids, as are
unpaired median nerves that emerge from the dorsal
surface of each thoracic ganglion. When it occurs, the
unpaired median nerve fuses with N1 of the adjacent
posterior ganglion and innervates the spiracle muscles
(Nesbitt, 1941).
The pro- and mesothoracic ganglia of S. lineola are
virtually the same size as they are in grasshoppers,
crickets, cockroaches, and termites (Nesbitt, 1941). In
contrast, both the absolute and relative size of the metatho-
racic ganglion differs throughout the orthopteroid orders,
depending on the number of abdominal ganglia that have
fused with it. For example, in Diapheromera femorata
(walking stick), Blaberus cranifer (cockroach), and Termop-
sis angusticollis (termite), only one abdominal ganglion is
fused; in Gryllus assimilis (cricket) two abdominal ganglia
are fused (Nesbitt, 1941); and in T. a. sinensis, H. membra-
nacea, M. religiosa, S. lineola (mantids), and Schistocerca
americana (grasshopper), three abdominal ganglia are
fused (Gray and Mill, 1985; Kerry and Mill, 1986; Liske et
al, 1989; Siegler and Pousman, 1990a,b). Finally, in man-
tids, PN1–5 innervate the basal and intrinsic muscles of
the prothoracic leg in a pattern similar to that seen in
cockroaches and grasshoppers (Nijenhuis and Dresden,
1955a,b; Ewer, 1957; Pipa and Cook, 1959; Campbell,
1961; Gray and Mill, 1985).
Although elusive when neurons are classified individu-
ally (Burrows, 1996), interesting organizational common-
alities emerge when groups of neurons are identified in
338
terms of the particular bundle in which their primary
neurites travel (Siegler and Pousman, 1990a,b). We have
identified only one, perhaps serially homologous, motor
group associated with N4 that is similar in soma number
and distribution, and neurite bundle shape and location to
the (apparently) corresponding group in grasshoppers (cf.,
Siegler and Pousman, 1990a, Fig. 3). To the extent that
such groups reflect fundamental developmental processes
(Siegler and Pousman, 1990a,b; Burrows, 1996), the simi-
larity between S. lineola and the grasshopper suggest a
conserved developmental strategy across orthopteroid or-
ders.
ACKNOWLEDGMENTS
We thank the editors and anonymous reviewers for their
thoughtful comments, criticisms, and helpful suggestions.
F.R.P. was supported by a University Research Council
grant and a Faculty Research and Development grant.
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