Manticora Leaflet

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 Fig. 1 - Manticora tuberculata (De Geer) by 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 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. latipennis Waterhouse, M. livingstoni Castelnau and M. holubi sp. n.

Fig. 86

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

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 eyes, often without visible punctures. Pronotum with anterior and lateral parts punctured very finely or without any punctures at all. Lobes of pronotum round. Elytra wide, cordiform, flat, dull, very finely granulated 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

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RARITY: Common

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

DISTRIBUTION:

Fig. 87

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

R.S.A. (Natal,Transvaal), Mozambique, Zimbabwe, Botswana, Namibia,Angola, Zambia.A widely distributed species with an apparent diversity of different populations. For a prospective description of subspecies, more of geografical and morphological work is needed.

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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. latipennis 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 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 specimens of M. mygaloides are very narrow. The elytra of male M. latipennis 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. latipennis 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. 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 collector 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

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

from Angola ("Angola - or., 25 km N-S. KAWOYA, G. Schnur leg." and "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 impression 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

Fig. 123

Manticora livingstoni CASTELNAU m, 50 mm, Namibia, 40 km S of Witvlei/Gobabis, 18 DEC 1999, H. W. GIESS leg., MARS

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. livingstoni 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 to 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 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,

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RARITY: Not rare

HABITAT: Dry sandy areas with low perennial plants, low grasses, shrubs and scattered low trees.

Fig. 124

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

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

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cordiform, gradually sloping posteriorly, disk gradually rising with striking 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 previous 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 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 elevated to a species status (Werner/Wiesner 1994, Mare‰ 1995). Now it is considered a junior synonym of M. livingstoni, but the above mentioned 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.

Fig. 125 - Livingstone, D. (1813 - 1873)

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

2

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5

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

Fig. 208 - Mating pair of M. scabra

Fig. 209 - Male and female of M. holubi in postcopulatory amplexus

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 behaviour 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 presence evoked in manticoras varied: crickets, mole crickets, grasshoppers and locusts, termites, cockroaches, melolonthine and ruteline Scarabeidae and their larvae, mealworms (Tenebrionidae), various moths and caterpillars, and f ly larvae. Cockroaches, caterpillars and most other beetle larvae elicited only moderate response from 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 Tenebrionidae (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 approaches her from the left side. Then he pounces on her, grabs her between prothorax and elytra 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 running, 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 observed a male (M. holubi from Rundu, Namibia) riding on the back of a female from 4 PM to roughly 11 AM of the following day, that is

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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 observation) or she turns on her back (Hrd˘, pers. com.). Sometimes a mating male riding on the back of a female is attacked by another male and a fight follows. I saw it several 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. 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 owing to nightfall. The next morning I returned to the spot but 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 ovipositor 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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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. holubi

Mating of M. scabra

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

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