Burmantis, GRIMALDI, 2003

GRIMALDI, DAVID, 2003, A Revision of Cretaceous Mantises and Their Relationships, Including New Taxa (Insecta: Dictyoptera: Mantodea), American Museum Novitates 3412, pp. 1-48 : 12-20

Burmantis , new genus

DIAGNOSIS: Distinguished from other genera known as nymphs in amber ( Chaeteessites , Electromantis , Jersimantis ) most readily by the distinctive foreleg structure: femur with ventromesal row of 5–6 stout, short spines, alternating with shorter ones; 3 long spines on ventrolateral edge; with dense, fine pilosity in ventral furrow. Forefemoral brush present, but setae not scalelike. Foretibia with mesal row of thick spines increasing in size distad (fine setae laterally); apex of tibia with two long, thick, spinelike setae, but not spurlike (observed only in type species). Forebasitarsomere slightly longer than foretibia; at least midocellus present (these two features observed only for the type specimens)

TYPE SPECIES: B. asiatica , n. sp.

INCLUDED SPECIES: B. asiatica , B. lebanensis , n. sp.

ETYMOLOGY: Directly from Burma, former name of the country of Myanmar, from where the type species derives; and mantis, a typical suffix in the order .

Burmantis asiatica , new species

Figures 2d , 7 , 8

DIAGNOSIS: Differs from Burmantis lebanensis as given in the diagnosis of that species, below.

DESCRIPTION: Based on a single specimen,

which is a nymphal exuvium. Since the cuticle is cleared, and with the thin amber preparation made, it was possible to study the specimen under compound microscopy at 100X. Portions of the body are collapsed and difficult to reconstruct, but even coloration patterns and microscopic structures like sensilla are observable. Head: Eyes large, but only partially preserved. Median ocellus present but lateral ones not apparent (perhaps a preservational artifact). Frons slightly bulbous; frontoclypeal suture well developed. Clypeus and labrum preserved (as figured); mandibles well developed, with heavily sclerotized teeth (dentition of right mandible figured), comparison between left and right mandibles (i.e., slight asymmetry) not possible. Labial palps preserved, 3­segmented; maxilla preserved, lacinia sharp and sclerotized, toothlike. Maxillary palps not preserved or apparent. Antenna long, flagellate; scape with a thin sclerite in the socket membrane ventrally; pedicel rounded apically; flagellomere 1 long, its length greater than that of scape + pedicel; basal flagellomeres very short and compact (lengths less than width), gradually lengthened apicad, with lengths 3– 4X the width.

Thorax: Crushed and distorted in places; pronotum difficult to reconstruct, but antepronotum apparently split away from rest of pronotum, and most of pronotum is split in half. Pronotum was apparently quadrate in shape, with mottled coloration, and possesses minute, sharp, spiculelike setulae scattered over surface. Similar setulae scattered over surface of wing pads, less so on other sclerites. Legs: Very well preserved. Forecoxa relatively short, length approximately twice the greatest width; articulation of forecoxa to prothorax is broad, seemingly with modest mobility; forecoxa with small knob on ventrolateral margin. All trochanters small. Forefemur large, basal third slightly inflated and bulbous, width of femur gradually tapered apicad. Base of each femur with small patch of 20–25 minute sensilla. Ventromesal sur­ face of bulbous base of forefemur, and ventral surface of basal half of forefemur, with dense, fine, erect pubescence. Ventrolateral edge of forefemur with 3 long, stiff, sclerotized spines, their lengths approximately equal to width of femur; each spine on a low tubercle. Ventromesal edge of forefemur, distal to bulbous base, with row of 18 short, spinelike setae; basalmost one (number 1) and numbers 3, 5, 7, and 9 thick; spines 2, 4, 6, 8, and 10–18 approximately half the thickness of others. No discoidal spines present. Mesal surface of forefemur with patch of scattered spicules (‘‘brush’’) near middle; spicules are slightly thickened, but not scaliform as in all living Mantodea (i.e., fig. 8a). Foretibia with 2 ventral rows of spines on distal two­thirds of tibia; mesal row with 8 thicker spines, lengths of which gradually increased distad, apicalmost spine nearly 3X width of tibia. Row of ventrolateral spines thinner, only apical spine large. Large apical spines of foretibia not situated on lobe of tibia that projects beyond tarsal articulation (fig. 8a), as occurs in all Mantodea except Chaeteessa . Forebasitarsomere length slightly less than foretibia. More distal foretarsomeres poorly preserved or lost. Midfemur stout, width twice that of fore­ or hindfemora, with longitudinal ventral groove; dorsal apex of midfemur with a short spine. Hindlegs long; femur slightly longer than tibia, apex of hindfemur ventrally incised and dorsally with short spine. Apex of hindtibia with pair of short spines ventrally; dorsally with small lobe. Hind basitarsomere longer than remaining tarsomeres. Length of hindtibia and tarsus 4.8 mm, approximately same length as body exclusive of cerci.

Abdomen: Short, broad, tergites with minute, spiculelike setulae. Cerci well developed, with broad base and tapered apicad to fine point; approximately 12 segments, most with long, fine setae (as figured for apical segments). No genitalic structures visible.

TYPE SPECIMEN: Holotype is a nymph, AMNH, MYANMAR: Kachin, amber mines near Tanai and Myitkyina. The specimen is in a clear yellowish piece of amber containing scattered bits of debris, stellate trichomes, two staphylinoid beetles, and a berothid neuropteran. The piece is slightly rectangular, 10 X 14 mm, and was trimmed and polished to 3 mm thickness and parallel to the plane at which the body of the insect is preserved.

ETYMOLOGY: Referring to the Asian locality of the fossil.

COMMENTS: Exquisite preservation of the forelegs reveals a tibial spination that is plesiomorphic: there are no discoidal spines, the setulae of the forefemoral brush are not particularly dense or scaliform in shape, and the tibial spines are not particularly large or thick (in the extant basal genus Chaeteessa these spines virtually form a basket). The fossil is apomorphic to Chaeteessa and Chaeteessites minutissimus in at least one important respect: a long forebasitarsomere (character 13, below).

Burmantis lebanensis , new species

Figure 9

DIAGNOSIS: Differs from B. asiatica by having fewer (4, vs. 10) small spines on the forefemur alternating among thick ones; pronotum and some other sclerites covered with small tubercles, instead of minute spiculelike setulae; cerci shorter and with 9–10 (vs. 12) segments, and without very long setae apically.

DESCRIPTION: Based entirely on a single nymphal exuvium. Head: Eyes large, but proportions not preserved, nor are ocelli. Mandibles heavily sclerotized, but dentition not visible. Scape and pedicel as in B. asiatica ; first flagellomere long, length about equal to 4–5 other, basal flagellomeres. Length of flagellomeres gradually and greatly increased distad. Thorax: Pronotum too distorted to reconstruct shape, but it and several other sclerites covered with small irregular tubercles (wing pads are smooth). Legs: Foreleg: Most of right one preserved; left one lost. Coxa very short, with deep mesal incision; femur tapered distad, ventral surface with dense, fine pubescence on proximal half; femur with two ventral rows of spines, ventromesal row with 5 thick, short, sclero­ tized spines alternating with 4 smaller, less sclerotized ones; ventrolateral row with 3 long spines, a minute one distally. Forefemoral brush present (seen vaguely in dorsal view of specimen), but details (i.e., number and shape of scales) not visible. Only proximal half of tibia preserved, bearing 6 spines increasing in length distad. Right midleg and hindfemur preserved, plus portion of left mid­ and hindleg. Mid­ and hindfemora fairly stout; midtibia very thin, length equal to that of midfemur. Length of midtarsi equal to length of midtibia; length of hindtibia approximately 1.6X length of midtibia. Abdomen: Largely lost or crumpled. Pair of styles is present; cerci fairly short, with a thick base and tapered to a fine point. Total number of cercal segments not discernable (basal ones obscured).

HOLOTYPE: AMNH L26 , in amber from LEBANON: near Bcharre´, collected by Antoun Estephan (Early Cretaceous, approximately Barremian). The amber piece is clear, transparent yellow; it was embedded in epoxy and trimmed to separate one piece containing a scelionid wasp, the other containing the mantis nymph and another parasitoid wasp.

ETYMOLOGY: Referring to Lebanon, the source country of the Lower Cretaceous amber.

COMMENTS: Foreleg structure of this species and B. asiatica leaves little doubt about their close relationship. The forefemur has similar proportions, with a depressed ventral surface having dense, fine setulae mostly on the basal half. There are two rows of spines, one on the ventromesal edge, the other on the ventrolateral edge. The ventromesal row has 5 strong, sclerotized, short spines, each separated by smaller, less sclerotized spines. The ventrolateral row has three long spines at the middle of the femur. Only the basal half of the tibia is preserved in B. lebanensis , but the spination that is preserved is very similar to that of B. asiatica .

Genus Chaeteessites Gratshev and Zherikhin

Chaeteessites Gratshev and Zherikhin, 1993: 157 . Type Species: C. minutissimus Gratshev and Zherikhin, 1993: 157 (Siberian amber [Santonian]). By original designation.

DIAGNOSIS: Known only as a partial nymph in Cretaceous amber from northern Siberia, defined mostly on the basis of distinctive spination of forelegs: Foretibia with two ventral rows of spines, ones in mesal row thicker, apex of tibia with pair of large spines but neither of them a spur nor situated on a process of the tibia that extends past the tarsal joint; femur ventrally with 3 long, very fine setae (no spines), no discoidal spines.

INCLUDED SPECIES: Monotypic.

COMMENTS: Gratshev and Zherikhin (1993: 157) originally defined the genus as a ‘‘collective’’ one for ‘‘chaeteessids of uncertain generic placement’’. The unique specimen is comprised of the anterior third of a nymph (fig. 10a, b) in a small chip of Siberian amber, preserving details of the foreleg (fig. 11). Preservation of the specimen does not allow observation of ocelli, so their presence is unconfirmed. The forefemoral brush is apparently absent, but this is difficult to be certain of given preservation of the specimen.

Figures

Fig. 2. Photomicrographs of Mantodea in Cretaceous ambers. a. Ambermantis wozniaki, new species, holotype AMNH NJ1085, in New Jersey amber. b. Paratype, ibid., AMNH NJ90cc. c. Jersimantis burmiticus, holotype AMNH Bu170, in Burmese amber. d. Burmantis asiatica, holotype, in Burmese amber. For scales, refer to illustrations of specimens.

Fig. 7. Burmantis asiatica, new species (holotype). a. General outline of body. b. Frontal view of face.

Fig. 8. Burmantis asiatica, new species (holotype). a. Portion of left foreleg, including femur, tibia, and basitarsomere. b. Wing pad. c. Apex of hindfemur. d. Apex of hindtibia. e. Cercus.

Fig. 9. Burmantis lebanensis, new species, holotype AMNH L26. a. Portions of pronotum, showing surface structure. b. Forefemur and basal portion of tibia. Area within dashed line is typical location of brush, which is obscure here. c–e. Apices of midtibia (c), hindtibia (d), and hindbasitarsomere (e). f. Left cercus and pair of styles.