complete and comprehensive monograph of the genus

Manticora clarifying the nomenclature, characterizing

all the species correctly, and presenting a new
determination key to both males and females
and a description of one new species

more than 500 color photographs of all types [holotypes,

lectotypes, and neotypes] together with photos
o f large series of specimens [males and females]
of all species from different localities as well as
photos of behaviour and localities

chapters covering history of study, biology, behaviour,

systematics, native legends and mythology, manticora's
reflection in literature and more

includes distribution maps for all 13 manticora

species

over 200 pages and a luxurious hardcover edition

The History
of Study of the Genus

19

For 59 years only one species (M. tuberculata) was known, described by the Swedish
entomologist De Geer (1778). But the author did not recognize that it is a cicindelid and
included it in the genus Carabus L. as C. tuberculatus. The same was done three years
later by the Danish entomologist Fabricius who was not aware of De Geer's description
and described it once more under the name Carabus maxillosus. In the same year (1781)
the species was described for the third time by Thunberg, who was the first to treat it as
a tiger beetle and included it in the genus Cicindela L. under the name C. gigantea.
These three entomologists worked with material of the same origin (Cape of Good Hope)
not knowing about each other. In the second half of the 18th century only the western part of
the present Cape Province was being colonized. At that time the Cape Colony (Kapland)
comprised only a small south-western part of the present Western
Cape. The eastern and midland parts were not inhabited by colonists
or visited by naturalists.
The genus Manticora was established by Fabricius in 1792.
The name given to the species by him (maxillosa), in spite of the fact
that it was a junior synonym, had been used for many years. It was
Klug in 1849 who began to use the correct name tuberculata and
others followed him.The name M. maxillaris used by Latreille in 1804
and Fischer in 1821 for the species was taken as an incorrect spelling
of M. maxillosa.
The second species of the genus, M. latipennis, was described by
Waterhouse in 1837. As it was based on a female specimen (now
deposited in BMNH), it later caused a serious nomenclatural confusion
(see SYSTEMATICS section of this book).
The third species (M. tibialis), coming from the eastern part
of the present Cape Province, was described in 1848 by Boheman.
In spite of the fact that it is very easily distinguishable from
M. tuberculata, the two were for some time confused. Most likely,
this was caused by the extreme rarity of Manticora specimens in
most collections, which made it impossible for authors to compare
the two species.
In 1849 Klug described three more species (M. granulata,
M. scabra, M. herculeana) in his monograph, of which only M. scabra
can be recognized as valid. The specimens on which Klug based his
characteristics and descriptions are deposited in ZMBC and marked
by the following numbers corresponding to the museum catalogue: Fig. 1 - Manticora tuberculata (De Geer)
No. 1 - M. tuberculata (in fact M. tibialis), No. 2 - M. scabra,
No. 3 - M. granulata, No. 4 - M. latipennis, No. 5 - M. herculeana. M. granulata is a large
female of M. tibialis. M. scabra and M. herculeana are smaller and large specimens
of the same species from different localities. The material of M. scabra came in many
specimens from Inhambane env., Mozambique, and the locality of the specimens described
as M. herculeana is said to be "the interior of Mozambique". According to nomenclatural
rules the name M. scabra, which was published three pages earlier, has priority and is
the valid name of the species.
The next two new species were described by Thomson in 1859. They were M. sichelii
and M. mygaloides, both well defined and justified. The lectotypes of both of them are
in DEIC.
In 1863 Laporte de Castelnau added three new species (M. dregei, M. ludovici and
M. livingstoni), but only M. livingstoni (lectotype in DEIC) can be recognized as valid.

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Fig. 14 - R.S.A., N Concordia,

habitat of M. skrabali

Fig. 15 - R.S.A., E Concordia,

habitat of M. skrabali

Fig. 16 - Namibia, Brandberg,

habitat of M. gruti

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Fig. 17 - Namibia, Brandberg,

habitat of M. gruti

Fig. 18 - Namibia, Otjiwarongo,

habitat of M. werneri

Fig. 19 - Namibia, Rehoboth,

habitat of M. livingstoni

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M a n t i c o r a m y g a l o i d e s T H O M S O N, 1 8 5 9

Monographie des Cicindelides I, p. 7, 8, 66, pl. 2, fig. 3, 4

Castelnau 1863. Fleutiaux 1892. Horn W. 1905, 1910, 1926. Mare‰ 1976,
1978, 1995, 1997, 2000.Wiesner 1992.Werner/Wiesner 1994.
Oberprieler/Arndt 2000.Werner 2000.
TYPE LOCALITY: Mozambique
TYPE REPOSITORY: DEIC (Lectotype, Fig. 92/1)
DESIGNATION OF THE LECTOTYPE: I designated as lectotype (Fig. 92/1)
one syntype in DEIC. It is a male 46 mm long. The following 5 labels
are attached to it: Mozambique - M. mygaloides Thomson (marked TYPE
on the reverse) - red label SYNTYPUS - coll. Ehlers V. de Poll - coll.W. Horn
DEI Eberswalde.The red label "LECTOTYPUS J. Mare‰ 2000" was added.
The lectotype was designated to stabilize the nomenclature, namely
to make clear the differences among M. mygaloides Thomson, M. lati-
pennis Waterhouse, M. livingstoni Castelnau and M. holubi sp. n.
DESCRIPTION: Habitus as in as in Fig. 86, 87 (1.8x actual size).
Medium size.
Male: Dull black. Mandibles long and narrow, right one bent at right
angle, left one only slightly shorter. Left inner tooth of left mandible
larger than right one. Head large, oval, only finely punctured between
Fig. 86 eyes, often without visible punctures. Pronotum with anterior and lateral
Manticora m ygaloides THOMSON parts punctured very finely or without any punctures at all. Lobes of
m, 51 mm, R.S.A., Transvaal, Thabazimbi, pronotum round. Elytra wide, cordiform, flat, dull, very finely granulated
8 NOV 1994, WERNER leg., MARS on sides of disk and posteriorly, most of disk smooth, gradually sloping
backward. Anterolateral margins of disk distinctly serrated. Elytra of
smaller specimens narrower, very narrow shape of the smallest specimens
reminds of M. tuberculata. Legs and tarsi black. Length 43 - 59 mm.
Female: Black, dull. Left inner tooth of left mandible of same length
as right one. Punctures of head and pronotum as in male. Elytra narrow,
only in the largest specimens broader, convex, almost smooth, only

equator

RARITY:
Common

HABITAT:
Sandy patches in relatively
moist habitats, in woods and
among dense growth of vege-
tation.

DISTRIBUTION:
R.S.A. (Natal,Transvaal),
Mozambique, Zimbabwe, Botswana,
Namibia,Angola, Zambia.A widely
distributed species with an apparent
diversity of different populations. For
Fig. 87 a prospective description of subspecies,
Manticora m ygaloides THOMSON more of geografical and morphological
work is needed.
f, 44 mm, R.S.A., Transavaal, Thabazimbi,
8 NOV 1994, WERNER leg., SKRB

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on the sides of disk and posteriorly slightly granulated. Legs and

tarsi black. Length 43 - 55 mm.
ETYMOLOGY:The species was named after its resemblance to the bird
spiders of the genus Mygale.
DISCUSSION:After comparing the female holotype of M. latipennis
Waterhouse (female) and the lectotype of M. mygaloides Thomson,
I have to disprove the synonymy of these taxons. Thomson (1859,
Nota p. 66) declared the synonymy of M. scabra Klug and M. lati-
pennis Waterhouse (which is not so) and suggested the new name
M. mygaloides for the real M. latipennis Waterhouse. Now it is clear
that Thomson (description, figures, type material) had in his hands
material different from M. latipennis Waterhouse, which deserves
a separate species status. Also Castelnau (1863) and Péringuey
(1893) regarded these two taxa as separate species. In spite
of being roughly the same size, M. latipennis has more densely Fig. 88 - M. mygaloides female lying eggs
and prominently granulated elytra, glossy instead of dull and wider
than those of M. mygaloides.This character is especially prominent
in small males of both species, small specimens of M. latipennis
retain the shape of elytra of the large ones, whereas small speci-
mens of M. mygaloides are very narrow. The elytra of male M. lati-
pennis are very glossy with more prominent granules on the disk,
whereas the elytra of M. mygaloides are almost smooth and dull.
Elytra of female M. latipennis are very wide, very glossy and distinctly
granulated at their sides, whereas those of female M. mygaloides
are much narrower, smooth and dull. Both taxa are widely distributed,
in some places their distributions overlap (Natal, Transvaal,
Zimbabwe), while other localities are inhabited by only one of them
(M. mygaloides in Mozambique, northern Namibia, southern
and western Angola, southern, central and western Zambia), M. lati-
pennis in several Botswana localities, e. g. Linokana, Serowe, Maun
env. etc.) As both taxa retain their basic characters in localities
where they co-exist, they should be considered separate species. Fig. 89 - M. mygaloides - larva (third instar)
REMARKS: As was already pointed out by Péringuey (1893), this
species varies much in size. In the south (Transvaal - Thabazimbi,
Werner/Lízler leg.) the length of the males does not exceed 50 mm
and that of females 47 mm, in the middle of the distribution range
in central Zimbabwe (70 km N of Chivhu, Featherstone env., SmrÏ
leg.) the maximum length of males is 52 mm and that of females
49 mm. These data are based on more than 30 specimens from
each locality. The real giants of this species live in northwestern
Zambia (Kabompo - Mufumbwe, NW Province, DEC 2001,
Werner/Lízler/Hrd˘ leg.). The only male captured there reached
the length of 59 mm and the largest of several big females measured
55 mm. This pair is by far the largest of this species known. At that
size the northern population of M. mygaloides is comparable with
the largest specimens of M. scabra. One month later, a local collec-
tor sent about 200 specimens of this species from Mufumbwe env.,
situated roughly 110 km NE from Kabompo. They were still large
(the length of the largest male 54 mm), but did not approach Fig. 90 - M. mygaloides - larva (third instar)
the size of the population from Kabompo - Mufumbwe. Specimens making its tunnel

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Fig. 91 - R.S.A., Transvaal, Thabazimbi, habitat from Angola ("Angola - or., 25 km N-S. KAWOYA, G. Schnur leg." and
of M. mygaloides "Angola - Ukuanyama, W. Hauser leg") are of the size of specimens from
R.S.A. The specimens collected in large numbers by SmrÏ in central
Zimbabwe (70 km N of Chivhu, Featherstone env.) differ by complete
smoothness of the elytra and mild gloss. But as stated above, more
geographical and morphological information is needed to ascertain
the possibility of a subspecies status.
Interesting information was provided in November 1994 by Werner
(pers. comm.). In the vicinity of Thabazimbi he heard one specimen
of this species stridulate, according to him most likely by rubbing its leg
against the elytra.The stridulation of this species was confirmed from
the same spot 4 years later by Lízler (pers. comm.), who had the impres-
sion that it was caused by rubbing the prothorax against the elytra.As far
as I know, stridulation was not observed in any other species of this genus.

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1 2 3

4 5 6

Fig. 92
Manticora m ygaloides THOMSON
1 - m, 46 mm, Mozambique, DEIC – LECTOTYPE
2 - m, 50mm, R.S.A., Transvaal, near Thabazimbi, 14 - 17 NOV 1997, LÍZLER leg., MARS
3 - m, 47 mm, R.S.A., Transvaal, Thabazimbi, 6 - 8 NOV 1994, WERNER leg., SKRB
4 - m, 43 mm, R.S.A., Transvaal, near Thabazimbi, 14 - 17 NOV 1997, LÍZLER leg., MARS
5 - m, 46 mm, R.S.A., Thabazimbi, N. Pr., 16 DEC 2001, M. HRDÝ ET AL. leg., HRDY
6 - m, 46 mm, R.S.A., Thabazimbi, N. Pr., 16 DEC 2001, M. HRDÝ ET AL. leg., HRDY
(all 1.6x actual size)

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M a n t i c o r a l i v i n g s t o n i C A S T E L NAU, 1 8 6 3

Rev. Zool. (2) XV, p. 71

Fleutiaux 1892. Horn W. 1905, 1910, 1926, 1932. Mare‰ 1976.
Wiesner 1992. Oberprieler/Arndt 2000.
TYPE LOCALITY: Vicinity of lake N'Gami, Damaraland (Namibia)
TYPE REPOSITORY: DEIC (Lectotype, Fig. 128/1)
SYNONYM: M. mygaloides var. damarensis Péringuey, 1893.
M. damarensis Werner/Wiesner 1994,
Mare‰ 1995, 1997, Werner 2000
DESIGNATION OF THE LECTOTYPE: I designated as a lectotype (Fig.
128/1) a male from the type series of 5 syntypes in DEIC. It is 50 mm
long and the following 6 labels are attached to it: N'Gami - M. living-
stoni Castelnau (markes "TYPE" on the reverse) - red label SYNTYPUS
- coll. Ehlers V. de Poll - TYPE coll.W. Horn - Coll.W. Horn DEI Eberswalde.
A red label "LECTOTYPUS J. Mare‰ 2000" was added. The type series
corresponds perfectly with Castelnau's description (very long and slender
mandibles, very glossy, almost flat elytra with very feeble granulation,
occurrence at localities in the vicinity of lake N'Gami and Damaraland).
Part of Castelnau's cicindelid collection went to Van de Poll coll. and
further on to W. Horn coll. (now in DEIC).The lectotype was designated
Fig. 123
to stabilize the nomenclature of M. latipennis group.
Manticora livingstoni CASTELNAU
m, 50 mm, Namibia, DESCRIPTION: Habitus as in Fig. 123, 124 (1.6x actual size).
40 km S of Witvlei/Gobabis, Medium to large size.
18 DEC 1999, H. W. GIESS leg., MARS
Male: Black, sometimes rufous, very glossy. Mandibles very long and
very slender, left one only slightly shorter, right one bent at right angle.
Left inner tooth of left mandible larger than right one. Head large,
not punctured, anterior and lateral parts of pronotum punctured only
indistinctly, sometimes not at all. Lobes of pronotum round and less
prolonged posteriorly. Elytra very glossy, wider than in previous species,

equator

RARITY:
Not rare

HABITAT:
Dry sandy areas with low perennial
plants, low grasses, shrubs and scatte-
red low trees.
Fig. 124 DISTRIBUTION:
Manticora livingstoni CASTELNAU Central Namibian plateau, vicinity of the
f, 49 mm, Namibia, Rehoboth, lake N'Gami, Namibia - Botswana border.
11 MAR 1999, LÍZLER lgt., MARS

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cordiform, gradually sloping posteriorly, disk gradually rising with stri-

king depressions in the centre of each elytron. Sutural part perceptibly
convex. Lateral and posterior parts of disk losely and finely granulated,
centre smooth. Outer margin of disk serrated less distinctly than in pre-
vious species. Legs and tarsi black. Length 46 - 57 mm.
Female: Black, very glossy. Left inner tooth of left mandible larger than
right one. Head large, not punctured. Pronotum punctured as in male,
lobes of pronotum posteriorly round. Elytra moderately wide, almost
parallel-sided, convex, with a depression in the centre of each elytron as in
the male, very glossy, finely granulated on sides of disk and prominently
on posterior incline, centre of disk smooth. Margin of disk not serrated.
Legs and tarsi black. Length 43 - 50 mm.
ETYMOLOGY: The species was named in honor of David Livingstone,
a famous English explorer.
DISCUSSION: After examination of the type material of M. livingstoni
Castelnau and M. mygaloides var. damarensis Péringuey (1893, p. 13)
I have to conclude that the latter is a junior synonym of the former.
The type of M. mygaloides var. damarensis (in DEIC), its description
and its locality as given by Péringuey (Damaraland, vicinity of lake
Fig. 125 - Livingstone, D. (1813 - 1873)
N'Gami) are identical with the type series of M. livingstoni and its type
locality. A different species from northern Ovamboland was considered
erroneously by Péringuey (1893) to be M. livingstoni. The synonymy
of M. mygaloides var. damarensis with M. livingstoni has already
been proposed by W. Horn (after comparing the types) in his letter to
Péringuey (Péringuey 1898, p. 305) and in his further works (W. Horn
1905, 1910), but regardless of that both names have been generally used
in the sense of Péringuey (1893), not Castelnau (1863).The problem was
most likely caused by the fact that the type material of M. livingstoni
in DEIC was overlooked and considered lost. It has also been omitted
in the Döbler's (1973) "Katalog der in Sammlungen des ehemaligen
Deutschen Entomologischen Institutes aufbewahrten Typen - IX".
Later, after an extensive study M. mygaloides var. damarensis was ele-
vated to a species status (Werner/Wiesner 1994, Mare‰ 1995). Now it
is considered a junior synonym of M. livingstoni, but the above men-
tioned studies demonstrated that M. holubi sp. n. (which was treated
as M. livingstoni) and M. livingstoni Castelnau (which was treated as
M. damarensis) both are good species.
REMARKS: After a February rainfall near Rehoboth, Namibia, numerous
specimens were captured by Werner, who was also able to observe
the behaviour and biology of this species (Werner, pers. comm.).
Beyers collected several specimens of M. livingstoni together with
M. werneri near Gobabis, Namibia.

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Fig. 126 - Namibia, W Rehoboth, habitat of M. livingstoni 1

Fig. 127 - Namibia, N Rehoboth, habitat of M. livingstoni

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3 4 5

6 7 8

Fig. 128
Manticora l ivingstoni CASTELNAU
1 - m, 50 mm, Ngami (Botswana), DEIC – LECTOTYPE
2 - m, 50 mm, Ngami (Botswana), DEIC – SYNTYPE
3 - m, 49 mm, Ngami (Botswana), DEIC – SYNTYPE
4 - m, 54 mm, Namibia, 15 km N Rehoboth, 8 - 9 FEB 1995, WERNER lgt., MARS
5 - m, 53 mm, Namibia, 15 km N Rehoboth, 8 - 9 FEB 1995, WERNER lgt., MARS
6 - m, 51 mm, Namibia, 15 km N Rehoboth, 8 - 9 FEB 1995, WERNER leg., HRDY
7 - m, 52 mm, Namibia, 15 km N Reheboth, 8 - 9 FEB 1995, WERNER leg., KWC
8 - m, 51 mm, Namibia, 15 km N Rehoboth, 8 - 9 FEB 1995, WERNER leg., SKRB
(all 1.5x actual size)

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spectrum of these beetles. Insects of various orders as well as other

invertebrates and some vertebrates were offered to them. Oberprieler
states: "The following items didn't elicit hunting and feeding behav-
iour in manticoras: lizards, scorpions, spiders, ticks, millipedes and
snails. Woodlice were not readily identified as prey but sometimes
seized and eaten when at close range and the beetles were hungry.
The following insects, however, were consistently recognized as prey
and readily eaten, although the intensity of the response their pre-
sence evoked in manticoras varied: crickets, mole crickets, grasshop-
pers and locusts, termites, cockroaches, melolonthine and ruteline
Scarabeidae and their larvae, mealworms (Tenebrionidae), various
moths and caterpillars, and f ly larvae. Cockroaches, caterpillars
Fig. 207 - Male of M. scabra following and most other beetle larvae elicited only moderate response from
a female
manticoras in that they were generally not hunted (i.e. their trail
not followed), although the beetles would size
and eat them readily when detecting them at close
range. Moths (even large ones such as Agrius
convolvuli) were more readily seized and eaten
when found resting on the ground or on low
vegetation, such as under a light in the morning,
even though the numerous scales clearly troubled
the beetles. Hard beetles such as native Tenebrio-
nidae (genera Psammodes, Phanerotomea, Dichtha,
Zophosis) were also recognized as food and seized,
but manticora could never crush these and always
released them after a few attempts. The strongest
feeding response in manticoras was undoubtedly
elicited by crickets, grasshoppers, termites and
melolonthine beetles. When such an insect was
dropped or fell into a terrarium with a number
of hungry manticoras, a frantic scramble would
follow, with all beetles rushing around and biting
at everything they encountered, including their
own kind."
The extreme mandibles of males also have
a secondary function in mating. Owing to the keen
sense of smell, a male manticora, after detecting
the feromon of a female, follows her in the same
manner as he pursues prey. When near, he ap-
proaches her from the left side. Then he pounces
on her, grabs her between prothorax and elytra
Fig. 208 - Mating pair of M. scabra
by his longer right mandible, immediately locking
both of them in a firm grip around lateral coupling grooves at the base
of her prothorax.The grip is so strong that the female is unable to escape.
Sometimes she sinks down at once and the mating starts, sometimes
she runs around with the male riding on her back. When she is run-
ning, the male tries to catch some steady object by his hind legs to
stop her. When she is tired, she stops and sinks down, and the mating
begins. After it the female stands up and usually continues her run.
The male can rest on the back of the female for several hours, during
which many copulations take place.The female often stops and remains
still in an upright position for a long time. Both beetles are then rigid
and motionless like statues. The antennae of the female aim upward,
whereas those aim of the male downward. In a terrarium, I have ob-
served a male (M. holubi from Rundu, Namibia) riding on the back
Fig. 209 - Male and female of M. holubi in of a female from 4 PM to roughly 11 AM of the following day, that is
postcopulatory amplexus

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for approximately 19 hours.Then the pair

separated, but after few minutes the male
pounced on the female again and a new
copulation took place. If the female is
not ready to mate, she drives the male
off by the raised left hind leg (my obser-
vation) or she turns on her back (Hrd˘,
pers. com.).
Sometimes a mating male riding on
the back of a female is attacked by anot-
her male and a fight follows. I saw it seve-
ral times observing M. scabra and
M. mygaloides in the wild as well as in
captivity. The intruder tries to throw off
the riding male who defends his position
vigorously. It is not so easy as the female
moves around and continuously changes
the posture of the first male. Observing
such a fight, I always had the impression
of seeing the duel of an infantryman
with a cavalryman on a crazy horse.
Usually the riding male is able to defend
his right. Only in case the intruder is
much larger, he sometimes succeeds in his indecent effort. Fig. 210 - Details of male and female
M. scabra in postcopulatory amplexus
Once I saw (in the wild) a female of M. holubi taking the male
riding on her back as far as to her burrow. She quickly went into it
and the male had to stay outside as the hole was not wide enough
for both. He did not follow her in but stayed in front of the tunnel
guarding it. He either walked impatiently around the entrance or stood
still close to it. At one moment the female appeared in the opening
and the male became agitated. But after a few minutes the female
moved deeper inside again. After roughly one hour I had to leave Fig. 211 - Details of male and female
owing to nightfall. The next morning I returned to the spot but M. scabra in postcopulatory amplexus
the patient suitor was not there. Most
likely he was already riding his chosen
one somewhere in the field.
For laying eggs the female chooses
a sandy ground, and by the tip of her
elytra and hind legs makes a shallow
hole in the soil. Then she slides out
the enormous ovipositor and by two
movable hooks at its end penetrates
deeper. She prones in the depression
in a declining position, the front part of
her body raised up on front legs, the ovi-
positor buried deep in the ground. She
lays only one egg in each hole. The egg
is yellowish, of oval shape, and 2 - 3 mm
long (M. holubi). Mass ovipositions
by manticora females were witnessed
by SníÏek (M. holubi in Rundu env.,
9 - 11 FEB) and by a Werner/Lízler/Hrd˘
expedition (M. mygaloides in NW Zambia,
mid-December).
After hatching, the little larva enlarges
its cell into a tunnel aiming upward

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Mating of M. scabra Mating of M. holubi

Female of M. scabra defending herself against a male’s attempt to mate by lying on her back

Fig. 219 - Mating pair of M. scabra

Fig. 220 - M. scabra (female)

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Fig. 221 - Fighting males of M. scabra

Fig. 222 - Defence position of an attacked manticora

(M. scabra - female)

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Female of M. mygaloides laying eggs

Fig. 223 - Female of M. mygaloides laying eggs

Fig. 224 - Larva of M. mygaloides - third instar

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Fig. 225 - Larva of M. mygaloides - third instar

Fig. 226 - Larva of M. mygaloides burrowing its tunnel

Manticora - A Monograph of the Genus