Manticora Leaflet

complete and comprehensive monograph of the genusManticora clarifying the nomenclature, characterizingall the species correctly, and presenting a newdeterminat ion key to both males and femalesand a description of one new species

more than 500 color photographs of all types [holotypes,lectotypes, and neotypes] together with photosof large series of specimens [males and females]of all species from different localities as well asphotos of behaviour and localities

chapters covering history of study, biology, behaviour,systematics, native legends and mythology, manticora'sreflection in literature and more

includes distribution maps for all 13 manticoraspecies

over 200 pages and a luxurious hardcover edition

T h e H i s t o r y

For 59 years only one species (M. tuberculata) was known, described by the Swedishentomologist De Geer (1778). But the author did not recognize that it is a cicindelid andincluded it in the genus Carabus L. as C. tuberculatus.The same was done three yearslater by the Danish entomologist Fabricius who was not aware of De Geer's descriptionand 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 asa 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 ofthe present Cape Province was being colonized.At that time the Cape Colony (Kapland)comprised only a small south-western part of the present WesternCape.The eastern and midland parts were not inhabited by colonistsor 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 factthat it was a junior synonym, had been used for many years. It wasKlug in 1849 who began to use the correct name tuberculata andothers followed him.The name M. maxillaris used by Latreille in 1804and 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 byWaterhouse in 1837. As it was based on a female specimen (nowdeposited 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 inmost collections, which made it impossible for authors to comparethe two species.

In 1849 Klug described three more species (M. granulata,M. scabra, M. herculeana) in his monograph, of which only M. scabracan be recognized as valid. The specimens on which Klug based hischaracteristics and descriptions are deposited in ZMBC and markedby the following numbers corresponding to the museum catalogue: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 largefemale 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 describedas M. herculeana is said to be "the interior of Mozambique".According to nomenclaturalrules 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. sicheliiand M. mygaloides, both well defined and justified.The lectotypes of both of them arein 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.

o f S t u d y o f t h e G e n u s

M a n t i c o r a - A M o n o g r a p h o f t h e G e n u s

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Fig. 1 - Manticora tuberculata (De Geer)

28B I O T O P E S A N D T H E I R C H A R A C T E R I S T I C S


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

29B I O T O P E S A N D T H E I R C H A R A C T E R I S T I C S


Fig. 17 - Namibia, Brandberg, habitat of M. gruti

Fig. 18 - Namibia, Otjiwarongo, habitat of M. werneri

Fig. 19 - Namibia, Rehoboth, habitat of M. livingstoni

S Y S T E M A T I C S


Manticora mygaloides THOMSON, 1859

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: MozambiqueTYPE 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 labelsare attached to it: Mozambique - M. mygaloides Thomson (marked TYPEon the reverse) - red label SYNTYPUS - coll. Ehlers V. de Poll - coll.W. HornDEI 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 rightangle, left one only slightly shorter. Left inner tooth of left mandiblelarger than right one. Head large, oval, only finely punctured betweeneyes, often without visible punctures. Pronotum with anterior and lateralparts punctured very finely or without any punctures at all. Lobes ofpronotum round. Elytra wide, cordiform, flat, dull, very finely granulatedon sides of disk and posteriorly, most of disk smooth, gradually slopingbackward. Anterolateral margins of disk distinctly serrated. Elytra ofsmaller specimens narrower, very narrow shape of the smallest specimensreminds of M. tuberculata. Legs and tarsi black. Length 43 - 59 mm.Female: Black, dull. Left inner tooth of left mandible of same lengthas right one. Punctures of head and pronotum as in male. Elytra narrow,only in the largest specimens broader, convex, almost smooth, only

RARITY:Common

HABITAT:Sandy patches in relativelymoist habitats, in woods andamong dense growth of vege-tation.

DISTRIBUTION:R.S.A. (Natal,Transvaal),Mozambique, Zimbabwe, Botswana,Namibia,Angola, Zambia.A widelydistributed species with an apparentdiversity of different populations. For a prospective description of subspecies,more of geografical and morphologicalwork is needed.

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Fig. 86Manticora mmygaloides THOMSON

m, 51 mm, R.S.A., Transvaal, Thabazimbi, 8 NOV 1994, WERNER leg., MARS


f, 44 mm, R.S.A., Transavaal, Thabazimbi, 8 NOV 1994, WERNER leg., SKRB

on the sides of disk and posteriorly slightly granulated. Legs andtarsi black. Length 43 - 55 mm.

ETYMOLOGY:The species was named after its resemblance to the birdspiders of the genus Mygale.

DISCUSSION:After comparing the female holotype of M. latipennisWaterhouse (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 nameM. mygaloides for the real M. latipennis Waterhouse. Now it is clearthat Thomson (description, figures, type material) had in his handsmaterial 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 and prominently granulated elytra, glossy instead of dull and widerthan those of M. mygaloides.This character is especially prominentin small males of both species, small specimens of M. latipennisretain 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 distinctlygranulated at their sides, whereas those of female M. mygaloidesare 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, Maunenv. etc.) As both taxa retain their basic characters in localitieswhere they co-exist, they should be considered separate species.

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 mmand that of females 47 mm, in the middle of the distribution rangein central Zimbabwe (70 km N of Chivhu, Featherstone env., SmrÏleg.) 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 northwesternZambia (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 measured55 mm.This pair is by far the largest of this species known. At thatsize the northern population of M. mygaloides is comparable withthe 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 the size of the population from Kabompo - Mufumbwe. Specimens



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Fig. 88 - M. mygaloides female lying eggs

Fig. 89 - M. mygaloides - larva (third instar)

Fig. 90 - M. mygaloides - larva (third instar)making its tunnel



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from Angola ("Angola - or., 25 km N-S. KAWOYA, G. Schnur leg." and"Angola - Ukuanyama,W. Hauser leg") are of the size of specimens fromR.S.A.The specimens collected in large numbers by SmrÏ in centralZimbabwe (70 km N of Chivhu, Featherstone env.) differ by completesmoothness 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 legagainst the elytra.The stridulation of this species was confirmed fromthe 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 faras I know, stridulation was not observed in any other species of this genus.

Fig. 91 - R.S.A., Transvaal, Thabazimbi, habitatof M. mygaloides



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1 2 3

4 5 6


1 - m, 46 mm, Mozambique, DEIC – LECTOTYPE2 - 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., SKRB4 - 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., HRDY6 - m, 46 mm, R.S.A., Thabazimbi, N. Pr., 16 DEC 2001, M. HRDÝ ET AL. leg., HRDY

(all 1.6x actual size)



Manticora l ivingstoni CASTELNAU, 1863

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 mmlong 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 seriescorresponds perfectly with Castelnau's description (very long and slendermandibles, very glossy, almost f lat 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. andfurther on to W. Horn coll. (now in DEIC).The lectotype was designatedto stabilize the nomenclature of M. latipennis group.

DESCRIPTION: Habitus as in Fig. 123, 124 (1.6x actual size).Medium to large size.Male: Black, sometimes rufous, very glossy. Mandibles very long andvery 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 onlyindistinctly, sometimes not at all. Lobes of pronotum round and lessprolonged posteriorly. Elytra very glossy, wider than in previous species,

RARITY:Not rare

HABITAT:Dry sandy areas with low perennialplants, low grasses, shrubs and scatte-red low trees.

DISTRIBUTION:Central Namibian plateau, vicinity of thelake N'Gami, Namibia - Botswana border.

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Fig. 123Manticora llivingstoni CASTELNAU

m, 50 mm, Namibia, 40 km S of Witvlei/Gobabis,

18 DEC 1999, H. W. GIESS leg., MARS


f, 49 mm, Namibia, Rehoboth, 11 MAR 1999, LÍZLER lgt., MARS



cordiform, gradually sloping posteriorly, disk gradually rising with stri-king depressions in the centre of each elytron. Sutural part perceptiblyconvex. 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 thanright one. Head large, not punctured. Pronotum punctured as in male,lobes of pronotum posteriorly round. Elytra moderately wide, almostparallel-sided, convex, with a depression in the centre of each elytron as inthe male, very glossy, finely granulated on sides of disk and prominentlyon 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. livingstoniCastelnau 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 descriptionand its locality as given by Péringuey (Damaraland, vicinity of lakeN'Gami) are identical with the type series of M. livingstoni and its typelocality.A different species from northern Ovamboland was considerederroneously by Péringuey (1893) to be M. livingstoni.The synonymyof M. mygaloides var. damarensis with M. livingstoni has alreadybeen proposed by W. Horn (after comparing the types) in his letter toPéringuey (Péringuey 1898, p. 305) and in his further works (W. Horn1905, 1910), but regardless of that both names have been generally usedin the sense of Péringuey (1893), not Castelnau (1863).The problem wasmost likely caused by the fact that the type material of M. livingstoniin DEIC was overlooked and considered lost. It has also been omittedin the Döbler's (1973) "Katalog der in Sammlungen des ehemaligenDeutschen 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 asM. damarensis) both are good species.

REMARKS: After a February rainfall near Rehoboth, Namibia, numerousspecimens were captured by Werner, who was also able to observethe 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. 125 - Livingstone, D. (1813 - 1873)



104

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

2

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



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3 4 5

6 7 8


1 - m, 50 mm, Ngami (Botswana), DEIC – LECTOTYPE2 - m, 50 mm, Ngami (Botswana), DEIC – SYNTYPE3 - m, 49 mm, Ngami (Botswana), DEIC – SYNTYPE

4 - m, 54 mm, Namibia, 15 km N Rehoboth, 8 - 9 FEB 1995, WERNER lgt., MARS5 - m, 53 mm, Namibia, 15 km N Rehoboth, 8 - 9 FEB 1995, WERNER lgt., MARS6 - m, 51 mm, Namibia, 15 km N Rehoboth, 8 - 9 FEB 1995, WERNER leg., HRDY7 - m, 52 mm, Namibia, 15 km N Reheboth, 8 - 9 FEB 1995, WERNER leg., KWC8 - m, 51 mm, Namibia, 15 km N Rehoboth, 8 - 9 FEB 1995, WERNER leg., SKRB

(all 1.5x actual size)

B E H A V I O U R & B I O L O G Y


160

spectrum of these beetles. Insects of various orders as well as otherinvertebrates and some vertebrates were offered to them. Oberprielerstates: "The following items didn't elicit hunting and feeding behav-iour in manticoras: lizards, scorpions, spiders, ticks, millipedes andsnails. Woodlice were not readily identified as prey but sometimesseized and eaten when at close range and the beetles were hungry.The following insects, however, were consistently recognized as preyand 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 rutelineScarabeidae and their larvae, mealworms (Tenebrionidae), variousmoths and caterpillars, and f ly larvae. Cockroaches, caterpillars and most other beetle larvae elicited only moderate response frommanticoras in that they were generally not hunted (i.e. their trail

not followed), although the beetles would sizeand eat them readily when detecting them at closerange. Moths (even large ones such as Agriusconvolvuli) were more readily seized and eatenwhen found resting on the ground or on lowvegetation, such as under a light in the morning,even though the numerous scales clearly troubledthe 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 alwaysreleased them after a few attempts. The strongestfeeding response in manticoras was undoubtedlyelicited by crickets, grasshoppers, termites andmelolonthine beetles. When such an insect wasdropped or fell into a terrarium with a numberof hungry manticoras, a frantic scramble wouldfollow, with all beetles rushing around and bitingat everything they encountered, including theirown kind."

The extreme mandibles of males also have a secondary function in mating. Owing to the keensense of smell, a male manticora, after detectingthe feromon of a female, follows her in the samemanner as he pursues prey. When near, he ap-proaches her from the left side. Then he pounceson her, grabs her between prothorax and elytraby his longer right mandible, immediately locking

both of them in a firm grip around lateral coupling grooves at the baseof her prothorax.The grip is so strong that the female is unable to escape.Sometimes she sinks down at once and the mating starts, sometimesshe 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 tostop her. When she is tired, she stops and sinks down, and the matingbegins. After it the female stands up and usually continues her run.The male can rest on the back of the female for several hours, duringwhich many copulations take place.The female often stops and remainsstill in an upright position for a long time. Both beetles are then rigidand 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 backof a female from 4 PM to roughly 11 AM of the following day, that is

Fig. 207 - Male of M. scabra following a female

Fig. 208 - Mating pair of M. scabra

Fig. 209 - Male and female of M. holubi inpostcopulatory amplexus



161

for approximately 19 hours.Then the pairseparated, but after few minutes the malepounced on the female again and a newcopulation took place. If the female isnot ready to mate, she drives the maleoff by the raised left hind leg (my obser-vation) or she turns on her back (Hrd˘,pers. com.).

Sometimes a mating male riding onthe 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 incaptivity. The intruder tries to throw offthe riding male who defends his positionvigorously. It is not so easy as the femalemoves around and continuously changesthe posture of the first male. Observingsuch a fight, I always had the impressionof seeing the duel of an infantrymanwith a cavalryman on a crazy horse.Usually the riding male is able to defendhis right. Only in case the intruder ismuch larger, he sometimes succeeds in his indecent effort.

Once I saw (in the wild) a female of M. holubi taking the maleriding on her back as far as to her burrow. She quickly went into itand the male had to stay outside as the hole was not wide enoughfor both. He did not follow her in but stayed in front of the tunnelguarding it. He either walked impatiently around the entrance or stoodstill close to it. At one moment the female appeared in the openingand the male became agitated. But after a few minutes the femalemoved deeper inside again. After roughly one hour I had to leaveowing to nightfall. The next morning I returned to the spot but the patient suitor was not there. Mostlikely he was already riding his chosenone somewhere in the field.

For laying eggs the female chooses a sandy ground, and by the tip of herelytra and hind legs makes a shallowhole in the soil. Then she slides out the enormous ovipositor and by twomovable hooks at its end penetratesdeeper. She prones in the depression in a declining position, the front part ofher body raised up on front legs, the ovi-positor buried deep in the ground. Shelays only one egg in each hole. The eggis yellowish, of oval shape, and 2 - 3 mmlong (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 enlargesits cell into a tunnel aiming upward

Fig. 210 - Details of male and female M. scabra in postcopulatory amplexus

Fig. 211 - Details of male and female M. scabra in postcopulatory amplexus



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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)



167

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

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Manticora Leaflet