Santanmantis, 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 : 27-37

Santanmantis , new genus

DIAGNOSIS: A primitive type of mantis with tips of wings apomorphically extended well beyond apex of the abdomen (by more than one­third the wing length); venation reduced, such that vein M has only 2 main branches (vs. 3 or 4 found in other primitive mantises) and only 4 main branches of vein CuA (vs. generally 5 or more). Most distinctive is the very long pseudovein: instead of a sclerotized area restricted to the basal fork of M and Cu 1, it is a tubular vein extending from this region through veins CuA 2, CuP, and anal veins and nearly reaching margin of anal lobe. The genus possesses the following combination of plesiomorphic characters: prothorax short; pronotum wider than long, nearly discoid; at least middle femur (and probably hindfemur) with ventral row of spines; mid­ and hindlegs long and thin; forewings tegminous (at least the proximal half), as in roaches, with 4 main branches off vein R, CuP vein (claval furrow) deep and strongly curved; genitalia (possibly ovipositor) protruding from terminal segments (not internal).

TYPE SPECIES: S. axelrodi , new species.

INCLUDED SPECIES: Monotypic.

ETYMOLOGY: From Santana Formation ( Brazil), the provenance of the type specimen and species .

Santanmantis axelrodi , new species

Figures 16–24

DIAGNOSIS: As for the genus, given above.

DESCRIPTION: Gross aspects of ventral structures were observed using HRCT scans of the holotype specimen (figs. 18, 19). Measurements of various parts are given in table 2. Specimens from the SMNS (Staatliches Museum für Naturkunde, Stuttgart) have provisional numbers .

Head: Antennae filiform, at least basal 8– 10 flagellomeres with lengths 2.5X the width; scape and pedicel small. Eyes large, situated frontally and somewhat laterally, with a large postoccipital space. Distance between eyes wide, equal to width of eye. Ocelli present, but seen in only one specimen ( SMNS 172 ). Head hypognathous, mouthparts (mandibles, labrum) narrow compared to dorsal region of head .

Thorax: Short, prothorax not lengthened as in more derived mantises. Pronotum wider than long, its length 0.70–0.75X its width (as seen in AMNH 1957 , SMNS 112 , and 174), the surface evenly covered with fine punctations (perhaps sockets of lost hairs), with two slightly raised areas. Variation in the shape of the pronotum, from nearly discoid in the holotype to quadrate in some paratypes, appears due to preservational differences. Forelegs observed using HRCT on holotype: held frontally, tibiae and femora fold­ ed against each other, femoro­tibial joint barely reaching to level of posterior margin of eyes, presence of spines on either one or both segments suggested by HRCT, though details not discernable. Apex of each foretibia apparently with a spur, though cannot discern whether the spur has a well­defined articulation (i.e., fig. 19). Forecoxae not visible. Mid­ and hindlegs long and slender; proportions as given in table 2. Midcoxae not visible, but hindcoxae (observed with HRCT) small, situated medially, contiguous. No spines apparent on hindfemora or hindtibiae, but row of at least 4 ventral spines occur on midfemur (visible dorsally). Forewings tegminous (especially basal half), long and narrow, extend well past apices of cerci. Pseudovein uniquely long among mantises: a tubular vein extending from this region through veins CuA 2, CuP, and anal veins and nearly reaching margin of anal lobe. Wing lengths slightly longer than total length of body with cerci and exclusive of antennae (body length/forewing length = 0.82–0.94); wing length approximately four times the width (table 2). Fore­ and hindwings homonomous, though anal regions (i.e., presence of expansive fan on hindwing) were not preserved. Forewing venation: Vein Sc long, ends at level of middle of wing; R pectinate, with 5–6 main branches, including an apical fork (some branches are forked). Vein M is a simple fork, its base proximal to the end of Sc. Cu 1 with 4 main branches, bases of 2 most proximal branches very close. Claval furrow at CuA 2 well developed, being strongly arched and defined in relief (e.g., figs. 20c, 22c, f). CuP incomplete, distally shortened, with free end not joining CuA 2; A with two main branches. Only a portion of hindwing tip was preserved ( SMNS 112 : fig. 23) .

Abdomen: Relatively short and stout, length approximately 1.3X the width. Contents of a distended crop and portions from midgut were preserved in two specimens ( AMNH 1957 and SMNS 115 ) (see below). Cerci typically blattoid, well developed, 1.05–1.37 mm long and tapered apicad to fine point; with approximately 10 visible segments (best seen in left cercus of holotype), each segment with long fine setae. Ovipositor (gonapophyses, gonoplacs) protrudent, but short and broad; flanked by pair of small, triangular subgenital plates and with two pairs of small, mounded areas dorsally .

TYPE AND OTHER SPECIMENS: All are from Brazil: Ceara´, Crato Member of the Santana Formation (Aptian: Lower Cretaceous).

Holotype, AMNH 1957 (figs. 16–19): A complete specimen, though the wing venation of this specimen is not as well preserved as in AMNH 1956 , SMNS 112 , 113 , and 115. Proportions of various body structures indicate it is the same species as the other specimens. HRCT scanning of the holotype further revealed features not seen in the paratypes, particularly of the head and forelegs.

Paratype, AMNH 1956 (figs. 20, 24): A beautifully preserved, complete adult with forewings spread but hindwings folded over the abdomen. Apical third of forewings lost, probably because they are membranous; preservation of remaining, sclerotized portions of forewings excellent, showing significant relief. Pronotum subdiscoid; portions of femur and tibia of right foreleg exposed (but not revealing spines), as are portions of mid­ and hindlegs .

Paratype, SMNS 112 (fig. 22a): A beau­ tiful specimen with dorsal surface preserved; forewings spread and nearly completely preserved; hindwings folded and covered beneath abdomen. Pronotum preserved (fig. 22b); portions of right midfemur exposed and most of right hindtibia and tarsus. Abdomen well preserved, though cerci barely discernable .

Paratype, SMNS 113 (fig. 21c): A beautifully displayed adult with the forewings spread, revealing virtually all of the forewing venation (fig. 23). A color photograph of the specimen is in an exhibition catalog ( Bechly, 2001), where the specimen was identified as a chaeteessid. Portions of hindwing venation preserved, though no diagnostic details evident. Dorsal portions of head damaged. Prothorax appears to be saddle­shaped with anterior edge emarginate. A portion of what appears to be a midleg protrudes from under left forewing. Portions of what is probably left hindleg are exposed, including distal portion of femur; a long, thin tibia; and tarsi. Small spines occur along one edge of tarsi and apical portion of tibia. Forelegs not visible; probably folded beneath head and pronotum, buried in matrix .

Paratype, SMNS 115 (figs. 21a, 23): Headless specimen with ventral surface ex­ posed; right forewing is spread, revealing venation. What appears to be the left fore­ and hindwings are spread out, but overlapping venation makes venation difficult to discern. Abdomen broad, filled with material (probably ingestate); portions of legs preserved: right hindleg (femur and tibia only), base of left hindleg; left midleg (femur + tibia + tarsus). What appears to be left midfemur has

TABLE 2 Measurements of Santanmantis axelrodi Specimens (in mm)

row of at least 8 short, ventral spines. Most of thorax is scraped away, so the pronotum is not preserved, nor are forelegs.

Paratype, SMNS 172 (fig. 22d–f): Dorsal surface is exposed; wings poorly preserved (venation barely discernable, though revealing a deep claval furrow [fig. 22f]). Left forewing outstretched, left hindwing and right fore­ and hindwings folded over abdomen. Abdomen well preserved, including left cercus. Best portions of specimen are head and pronotum (fig. 22e) .

SMNS 114 : A complete adult (fig. 21b) with ventral surface exposed and wings fold­ ed, so venation not preserved. The forelegs appear to have a short, stout femur and tibia, rather different from the HRCT scans of the holotype, which is why this specimen was not assigned as a paratype of the species. Its apparent pedunculate eyes may be due to the matrix lying over the central front portion of the face .

ETYMOLOGY: Patronym in honor of Dr. Herbert Axelrod, for his interest with Santana fossil insects and his generosity to the AMNH.

DISCUSSION: The holotype and five paratypes clearly represent a new genus of basal mantis, not placement in the basal living family Chaeteessidae ( Bechly, 2001: 56) (see cladistic analysis, below). These specimens represent one of two superbly preserved Cretaceous mantis species. Santanmantis lacks synapomorphies distinct to all living mantises, including Mantoida and Chaeteessa , as given in the diagnosis. Santanmantis is distinct from Baissomantis (L. Cretaceous, Eurasia) , which has more dichotomous branching in R (vs. pectinate), more branches in CuA 1 (5 or 6, vs. 4), a complete CuP (figs. 23, 24), and no pseudovein. The two groups, though, have distinct plesiomorphic similarities, particularly the strongly arched claval furrow—a condition intermediate between roaches and more derived mantises. The extremely long wings (or, conversely, a very short abdomen) in Santanmantis are unusual, as most fully winged Mantodea and Blattodea have the tips of the wings extended to the apex of the abdomen or slightly beyond. In Santanmantis the wings extend well beyond the abdominal apex by more than onethird the wing length. This condition is in­ termediate between what is found in Isoptera and the rest of the Dictyoptera.

A preserved and full crop in the type specimen offered an apparent opportunity to confirm if the diet of Santanmantis was indeed predatory. Crop contents of mineralized insect fossils are sometimes well preserved (e.g., Krassilov and Rasnitsyn, 1999), and this was especially expected for this specimen given the preservation of relief, of tissues, and even cellular structures in Santana fossils ( Grimaldi and Maisey, 1990; Martill, 1988). Small fragments of the crop contents were studied using the AMNH Zeiss DSM­1 SEM, in order to scrutinize for fragments of plant or animal remains. If present, fragments of plant or arthropod cuticle would have been preserved, as these are particularly durable, but no biological structure was recognized in these samples. Though Santanmantis was clearly predatory (and possibly a scavenger as well), the crop of this specimen may have been filled with soft tissues.

Genus Vitimiphotina Gratshev and Zherikhin

Vitimiphotina Gratshev and Zherikhin, 1993: 154 . Type Species: V. corrupta Gratshev and Zherikhin, 1993: 155 (Early Cretaceous, Russia). By original designation.

DIAGNOSIS: Known only as portions of wings (PIN3064/8587, 3064/419), defined originally by Gratshev and Zherikhin on the basis of the following most significant features: wing with extensive dark patterns; R with single apical fork; M 2­branched, close to R but then strongly divergent; CuA with 6 apical branches.

INCLUDED SPECIES: Monotypic.

COMMENTS: The incomplete specimens on which the genus is based are too poorly known to include in a phylogenetic analysis and classification.

Figures

Fig. 16. Photomicrographs of Santanmantis axelrodi, new species, holotype (AMNH 1957), in Early Cretaceous limestone from the Santana Formation of Brazil. a. Dorsal view of cleaned specimen. b. Detail of head and pronotum. c. Detail of bases of wings. d. Detail of abdomen, showing the crop contents in relief.

Fig. 17. Illustrated rendering of Santanmantis axelrodi, holotype, with detail of terminalia.

Fig. 18. High­resolution CT scans of the holotype of Santanmantis axelrodi (AMNH 1957), showing various views of the anterior half. Top: Completely ventral (left, to oblique ventral, right). Middle: Completely lateral (left) to oblique lateral (right) (note great compression of the specimen). Bottom: Dorsal view, oblique (left) to completely dorsal (right). See text for description of methods and parameters.

Fig. 19. High­resolution CT scans of holotype of Santanmantis axelrodi. Left: a volume­rendering of complete specimen, exposing more of the long, slender hindlegs. Right: a more detailed, surfacerendered view of the ventral surface of the anterior half (cf. fig. 18). The rounded topography of the specimen and lack of spines and other fine structures are due to the resolution of imaging (14 µm), not the actual preservation. The bases of some spines on the foretibia and femur are visible.

Fig. 20. Paratype of Santanmantis axelrodi, AMNH 1956. a. Complete specimen, dorsal view. b. Detail of head and pronotum. c. Detail of left forewing. The membranous apical half of both forewings were not preserved.

Fig. 21. Santana Formation Mantodea in the SMNS. a. Paratype of Santanmantis axelrodi, SMNS 115. b. SMNS 114, probably a different species (see text). c. Paratype of Santanmantis axelrodi, SMNS 113. Photos of b and c are courtesy of Dr. Günter Bechly (SMNS).

Fig. 22. Santanmantis axelrodi paratypes. a–c. SMNS 112, showing dorsal habitus (a), detail of head and pronotum (b), and base of forewing with claval furrow (c). d–f. SMNS 172, showing habitus (d), detail of head and pronotum (e), and base of forewing (f) with raking light that depicts claval furrow in relief.

Fig. 23. Wing venation of Santanmantis axelrodi paratypes, showing slight variation.

Fig. 24. Left wing of Santanmantis axelrodi paratype, AMNH 1956.