AMERICAN MUSEUM Novitates
PUBLISHED BY THE AMERICAN MUSEUM OF NATURAL HISTORY CENTRAL PARK WEST AT 79TH STREET, NEW YORK, NY 10024
Number 3412, 47 pp., 27 figures, 3 tables July 28, 2003
A Revision of Cretaceous Mantises and Their Relationships, Including New Taxa (Insecta: Dictyoptera: Mantodea)
DAVID GRIMALDI!
ABSTRACT
All genera of Cretaceous mantises are reviewed, and diagnoses of some are revised based on re-examination of type specimens. Five new Mantodea are described from Cretaceous deposits on four continents, including: concretions in limestone from the Santana Formation of northeast Brazil (Aptian, 120 Ma), inclusions in amber from the Raritan Formation of New Jersey, USA (Turonian, 90 Ma), and in amber from undetermined formations of Lebanon (Barremian, 125 Ma) and northern Myanmar (Burma) (approximately early Cenomanian to late Albian, 100 Ma). Prior to this, virtually all of the oldest mantises were from five Creta- ceous localities in Eurasia. New Mantodea are Santanmantis axelrodi, n. gen., n. sp. (Brazil); Ambermantis wozniaki, n. gen., n. sp. (New Jersey); Jersimantis burmiticus, n. sp. (Myanmar); and Burmantis asiatica and B. lebanensis, n. gen. and n. spp. (Myanmar and Lebanon, re- spectively). The first two are based on adults, the last three on nymphs. Cladistic analysis of 26 morphological characters and 20 taxa, including living families and well-preserved fossils, indicates that Cretaceous mantises are phylogenetically basal to all living species and do not belong to the most basal living families Chaeteessidae, Mantoididae, and Metallyticidae. The classification of Cretaceous Mantodea is revised, which includes Santanmantidae, n. fam. and Ambermantidae, n. fam. Stratigraphic and cladistic ranks of taxa, with now improved fossil sampling, indicate that the order Mantodea is relatively recent like Isoptera (termites), with an origin no earlier than Late Jurassic. Superfamily Mantoidea, comprising three families and 95% of the Recent species in the order, radiated in the Early Tertiary to produce the exuberance of forms seen today.
' Curator, Division of Invertebrate Zoology, American Museum of Natural History. Adjunct Professor: Department of Ecology and Evolutionary Biology, City University of New York; Department of Entomology, Cornell University; Department of Ecology, Evolutionary, and Environmental Biology, Columbia University. e-mail: grimaldi@amnh.org
Copyright © American Museum of Natural History 2003 ISSN 0003-0082
2 AMERICAN MUSEUM NOVITATES
INTRODUCTION
Few insects have captured the popular imagination like butterflies, certain beetles, and mantises. Sexual cannibalism of the male mantis by his mate, for example, though of- ten exaggerated, actually is a frequent con- sequence of their voracious and indiscrimi- nate predatory behavior. The common green mantises of temperate regions, like Mantis religiosa L., do not reflect the true diversity of the nearly 2300 described species in the order, the great proportion of them occurring in tropical regions. Tropical species mimic, for example, twigs, leaves, flowers, and even ants, which not only prevents detection by other predators but better allows them to am- bush prey. As for any impressive group of organisms, an understanding of the relation- ships and origins of a group can provide unique insight into the evolution of special- ized life histories, such as predation. In this respect, perhaps the least appreciated aspect of mantis biology is the fact they are Dic- tyoptera.
Despite remarkable disparity in structure and habits, roaches, termites, and mantises comprise an indisputably monophyletic group, the Dictyoptera. This is based on mo- lecular (Wheeler et al., 2001) and morpho- logical features (Kristensen, 1975, 1991; Klass, 1997, 1998a, 1998b). Among the more distinctive morphological features de- fining the group are a perforated tentorium, a reduced to highly vestigial ovipositor large- ly or entirely hidden in a vestibulum, and eggs laid in a case, the ootheca. Exact rela- tionships of mantises to one of the other two dictyopteran orders, though, is controversial, and three of the four possible hypotheses have been proposed: (1) Mantodea (Blattaria + Isoptera): Hennig (1981), Klass (1997, 1998a, 1998b). (2) (Mantodea + Blattaria) Isoptera: Boudreaux (1979), Thorne and Car- penter (1992), Kambhampati (1996), Wheel- er et al. (2001), Vrsansky (2002), Vrsansky et al. (2002). (3) Mantodea + Isoptera + Blattaria (unresolved): Kristensen (1991), Grimaldi (1997).
Thus far no one has hypothesized a sister- group relationship of the two most morpho- logically modified orders, Mantodea + Is- optera. This ambiguity of relationships has
NO. 3412
been due in part to a common assumption that each order is monophyletic. While Is- optera and Mantodea are each clearly mono- phyletic, traditional and more recent evi- dence is compelling for a sister-group rela- tionship between termites and certain roaches (Cryptocercidae) (Klass, 1998a, 1998b; Lo et al., 2000), rendering the Blattaria paraphy- letic. Such paraphyly, though, has been dis- puted (Grandcolas, 1994, 1996, 1999; Grand- colas and D’Haese, 2001).
Despite the highly modified morphology of mantises (discussed in detail below), in several important respects they are basal to living roaches and all termites. For example, the main appendages comprising the ovipos- itor, the gonapophyses and gonoplacs, are least reduced in mantises among the three or- ders. In basal termites and living roaches the Ovipositor is entirely concealed within the vestibulum; in more derived termites, com- prising 99% of the species, the ovipositor is essentially lost. In mantises the ovipositor plesiomorphically protrudes from the vestib- ulum. Also, with a few exceptions (including some extinct species), mantises plesiomorph- ically have three well-developed ocelli; liv- ing roaches and all termites have only the lateral ocelli, with the median one being ex- tremely vestigial or lost altogether (Gn some termites and roaches all ocelli are lost). The wing venation of mantises, too, is more gen- eralized in some respects than in termites and roaches. Thus, it is highly unlikely that man- tises are closely related to any particular group of extant roaches.
Roaches are renowned for their antiquity because of fossils from the Carboniferous (e.g., Carpenter, 1992), an age more than twice that of the oldest known fossils of ter- mites (Thorne et al., 2000) and mantises (this report) from the Cretaceous. Great disparity in ages of the three orders was reconciled by proposals that Paleozoic and Lower Meso- zoic fossils are not true roaches but a para- phyletic assemblage of “‘roachoids,”’ or stem-group Dictyoptera, plesiomorphically possessing a long ovipositor (Hennig, 1981; Grimaldi, 1997). In this respect, Isoptera, Mantodea, and modern roaches are consid- ered derived from some extinct lineage of these roachoids, perhaps in the Jurassic.
The present report explores the earliest
2003
known history of mantises. Unfortunately, phylogenetic relationships among mantises have barely been explored, the most com- monly used classification is based largely on the work of Beier (1968). Beier divided the Mantodea into essentially eight families, which are adopted for purposes of this report:
Chaeteessidae: monogeneric (Chaeteessa), occur- ring in Neotropical forests, with very distinc- tive forefemoral and tibial spines.
Mantoididae: monogeneric (Mantoida), also oc- curring in Neotropical forests.
Metallyticidae: monogeneric (Metallyticus), col- orful and metallic forms in Asian forests.
Amorphoscelididae: 15 genera, all Old World (Af- rica, Mediterranean, Asia, Australia), with very distinctive femoral spines and spurs.
Eremiaphilidae: two genera (Eremiaphila, Heter- onutarsus), which are stout-bodied, long-legged forms that are apterous and brachypterous, liv- ing in deserts of northern Africa and the Middle East.
Empusidae, Hymenopodidae, Mantidae: 380 gen- era comprising the superfamily Mantoidea and 95% of all species. Comprises a great diversity of forms in most habitats. Some of the 21 sub- families of Mantidae are sometimes given fam- ily-level status.
Because this report focuses on the rela- tionships of early, Mesozoic fossils, the phy- logenetic hypothesis presented here focuses on the basal relationships of mantises, nota- bly families exclusive of the superfamily Mantoidea. Relationships within this diverse group require separate study. Also compro- mising the study of early mantis evolution is the scarcity of fossils, Mesozoic or other- wise, and many of these until now have been merely wings or fragments thereof. Table 1 is a summary of the known fossil mantises.
There are 17 species-level taxa of Creta- ceous Mantodea, and only about 10 Tertiary ones are known, though the available Tertia- ry specimens are not as well studied and their diversity is much greater. For example, there are approximately eight species of mantises in Miocene Dominican amber alone (D. Gri- maldi, unpubl.), most of them nymphs of Mantidae and all undescribed. A Jurassic mantis was recently described on the basis of a very fragmentary wing, specifically just the clavus (Vrsansky, 2002), which, contrary to the original claim, does not possess features specific to Mantodea. The presence of many
GRIMALDI: EARLY MANTISES 3
crossveins of Juramantis initialis Vrskansky, for example, is a feature widespread in Blat- todea from the Paleozoic to the Recent. The large number of known Cretaceous mantises is due mostly to the study by Gratshev and Zherikhin (1993), who described 12 species from productive deposits in central and east- ern Eurasia. Of the 17 Cretaceous taxa now known, 10 are compressions or impressions in rocks, and only one is not Eurasian but is from the Cretaceous of Brazil. The Brazilian Cretaceous fossils described herein are the only complete mantises preserved in rock from any geological period; Cretomantis lar- valis (from the Zaza Formation of Siberia and described herein) is an apparent exuvium of a nymph, so it is missing wings. Three of the rock fossil taxa are simply too fragmen- tary for any meaningful assessment of rela- tionships. These are Amorphoscelites sharo- vil, Kazakhophotina corrupta, and Vitimipho- tina corrugata, all described by Gratshev and Zherikhin (1993). The first is a foreleg and the last two are merely fragments of wings.
Specimens in amber have finer preserva- tion, typical of this medium (Grimaldi, 1996), but amber biases toward the preser- vation of smaller organisms, in this case nymphal mantises. Of the eight Cretaceous specimens in amber, five are nymphs. Chae- teessites and Electromantis in Santonian- aged amber from the Taymyr peninsula of northern Siberia have just the anterior por- tions of the body preserved. A new genus is described herein for two nearly complete nymphs in mid-Cretaceous amber from Myanmar and in Early Cretaceous amber from Lebanon. Jersimantis luzzii and a new species of this genus from Burmese amber are nymphs preserved in their entirety. Two New Jersey amber specimens are portions of adults; one a portion of a wing, the other comprised of wings, pronotum, and dorsal surface of the head. Finally, the finest pre- served specimen from the Mesozoic is a small adult in New Jersey amber, Amber- mantis, described herein.
Despite fragmentary specimens and a meager fossil record, the mantis fossils can provide powerful means for interpretation of evolutionary history when studied in a phy- logenetic context (e.g., Smith, 1994). With such an approach, the significance and infor-
+: AMERICAN MUSEUM NOVITATES NO. 3412 TABLE 1 Known Fossil Mantodea Age Taxon Matrix Parts Location Epoch/Series (Ma) Ref.# CRETACEOUS Ambermantis wozniaki> amber adult New Jersey Turonian 90 1 Amorphoscelites sharovi shale foreleg Siberia Valang-Haut. 135 2 Baissomantis maculata shale wings Siberia Valang-Haut. 135 2 Baissomantis picta shale wings Siberia Valang-Haut. 135 2 Burmantis asiatica> amber nymph Myanmar Cenomanian 100 1 Burmantis lebanensis® amber nymph Lebanon Barremian 125 1 Chaeteessites minutissimus amber nymph Siberia Santonian 85 2 Cretomantis larvalis shale nymph Siberia Valang-Haut. 135 2 Cretophotina mongolica shale wing Mongolia Barrem.-Aptian 125 2 Cretophotina serotina shale wing Kazakhastan Turonian 90 2 Cretophotina tristriata shale wings Siberia Valang-Haut. 135 2 Electromantis sukatshevae amber nymph Siberia Santonian 85 2 Jersimantis burmiticus> amber nymph Myanmar Cenomanian 100 1 Jersimantis luzzit amber nymph New Jersey Turonian 90 3 Kazakhophotina corrupta shale wing Kazakhastan Turonian 90 Z Santanmantis axelrodi> limestone adult Brazil Aptian 115 l Vitimiphotina corrugata shale wing Siberia Valang.-Haut. 135 Z TERTIARY
Archaeopllebia enigmatica shale wing France Paleocene 60 4 Arverineura insignis shale wings France Paleocene 60 4 Chaeteessa sp. amber adult Dom. Republic Miocene 20 5 Lithophotina floccosa shale wings Colorado Eocene 45 6 Mantidae spp. amber nymphs Dom. Republic Miocene 20 5 Mantidae spp. amber nymphs Baltic Region Eocene 40 5 Mantoida sp. amber adult Dom. Republic Miocene 20 5 Megaphotina sichotensis shale wing Russia Oligocene 35 2 Prochaeradodis enigmaticus shale wings France Paleocene 60 4
# The most recent, comprehensive reference for the fossils is provided, not necessarily the original reference where descriptions were made. References: 1, this paper; 2, Gratshev and Zherikhin, 1993; 3, Grimaldi, 1997; 4, Nel and Roy, 1996; 5, D. Grimaldi,
unpubl.; 6, Sharov, 1962. b Species described herein.
mation content of preserved venational char- acters can be determined, and, in conjunction with chronology, ages of lineages can be bet- ter extrapolated. Zherikhin (2002: 276), in the most recent review of mantis fossils, stat- ed ‘“‘The oldest known fossils [in the Creta- ceous] may well represent the early stage of mantid evolution’’, which contrasts with the estimates of a Late Paleozoic age proposed by Carpenter (1992) and Hennig (1981). With new specimens and this analytical ap- proach, questions like the following can be addressed: Is the apparent young age of the order due to a gap in the fossil record of 100 million years or more, or did mantises evolve
only about 150 Ma? The answer depends in part on how one defines a mantis.
MATERIALS AND METHODS
SOURCES, PREPARATION, AND STUDY OF NEW MATERIAL
Specimens newly reported here derive from three sources: Lower Cretaceous (Ap- tian-aged) limestone from the Santana For- mation, Ceara, Brazil; and mid-Cretaceous ambers from the Raritan Formation of central New Jersey, USA (Turonian), and undefined formations in northern Burma (Myanmar). Additional, previously reported and de-
2003
scribed specimens derive from various de- posits as specified later.
Santana Formation fossils include a great diversity of vertebrates and arthropods (Maisey, 1991) and are renowned for their preservation (Grimaldi and Maisey, 1990; Martill, 1988). The rich insect deposits de- rive from the Crato Member of the Santana Formation, near the town of Nova Olinda in Ceara, northeastern Brazil. Though the Crato Member has not been dated palynologically, its lithology indicates a probable Aptian age. The fossils, including arthropods, are pre- served as concretions of iron hydroxide and apatite, so they have relief that is lifelike or nearly so. They have also preserved remark- able details, including soft tissues such as striations of muscle myofibrils. A matrix of soft, fine-grained limestone facilitates prep- aration with acid digestion, using a weak so- lution of acetic acid (2% or less), but this technique is complicated by the intricacy of arthropods. With too much acid digestion, fine structures like spines, antennae, and wing veins will disintegrate.
To avoid possible damage from prepara- tion, AMNH 1957 was scanned using ultra high resolution computerized X-ray tomog- raphy (UHR CT) to observe critical ventral structures obscured by matrix. The three-di- mensional preservation of the fossils, in a matrix with substantially lower density than the concretion, allows the use of this tech- nique. The technique has been described and used very successfully for large insects and small vertebrates preserved in amber (Gri- maldi et al., 2000a).
CT scanning used an ACTIS 200/225 Mi- crofocus System (Bio-Imaging Research, Lincolnshire, IL), operating at 150 kV of X- ray energy. Five contiguous image stacks were collected in volume CT mode (cone beam) at a slice thickness of 14 wm. The final reconstructed image resolution was 512 x 512 pixels. For three-dimensional recon- struction the original dataset was cropped us- ing Scion Image (Scion Corporation, Fred- erick, MD). Isosurface rendering was done using Imaris Surpass 3.1 (Bitplane AG, Zu- rich), and volume rendering used Voxblast 3.0 (Vaytek Inc., Fairfield, IA). A rotating, three-dimensional image of AMNH 1957 can be viewed on www.amnh.org/science. While
GRIMALDI: EARLY MANTISES S,
the resolution was sufficient to determine the size and position of the forelegs and the pres- ence of spines (i.e., their bases), the structure of foreleg spines could not be seen.
Amber specimens from the Cretaceous of central New Jersey, USA, derive from the palynologically-dated Raritan Formation (Turonian). Mantodean specimens reported here come from two closely situated outcrops of equivalent age, in the towns of Sayreville and East Brunswick, Middlesex County. These outcrops have yielded an impressive diversity of fossilized organisms, including various plants; the oldest mushrooms; the oldest true tardigrade, definitive ants, and a parasitiform mite (an argasid soft tick); a plethora of arthropods; and two extremely rare flowers (reviewed in Grimaldi et al., 2000b). Like most amber deposits, amber from the Raritan Formation was deposited in brackish water lagoons and deltas, and in this case the amber was produced by a forest of Cupressaceae in a warm temperate or sub- tropical environment (Grimaldi et al., 2000b).
Burmese amber has recently been redis- covered (e.g., Grimaldi et al., 2002), with the only fossiliferous collection of the material having been assembled 80 years ago and re- siding at the Natural History Museum (NHM), London. Historically and presently, the material derives from northern Myanmar, Kachin state, though identity and stratigra- phy of the deposits have been confused. Many popular reports indicate this amber is Eocene or younger, which has been attributed to its redeposition in younger deposits. Re- cent re-study of arthropod inclusions in the NHM collection, though, indicates a Creta- ceous age (Rasnitsyn and Ross, 2000; Zher- ikhin and Ross, 2000). Study of the AMNH collection indicates an age that is probably mid-Cretaceous, perhaps Cenomanian or Tu- ronian (Grimaldi et al., 2002). Most recently, dating based on ammonites and pollen indi- cates an age of Late Albian for Burmese am- ber (Cruikshank and Ko, 2002). Moreover, this amber has preserved a biota quite dis- tinct from the prolific deposits of Cretaceous amber in Canada, Siberia, northern Spain, and New Jersey, probably due to its age, pa- leogeographic isolation, and its formation in a distinctly tropical paleoenvironment. Bur-
6 AMERICAN MUSEUM NOVITATES
mese amber was formed by a conifer, per- haps Metasequoia, and is probably the most biotically diverse Cretaceous amber deposit (Grimaldi et al., 2002).
Lebanese amber deposits are the only ones to prolifically yield the oldest insect inclu- sions, approximately Barremian in age. Other deposits of Lower Cretaceous amber occur, including ones as old as Lebanese amber, but none are so prolific or diverse. Though known for decades (Schlee and Dietrich, 1970), only recently has a systematic study of Lebanese amber been made (Azar, 2000). Dozens of deposits occur throughout Leba- non. Indeed, the formations yielding this am- ber occur in Jordan and Israel (the “‘Levan- tine Amber Belt’’), though insects are known in amber only from the former of the two. The source of this amber has popularly been reported as araucarian even though the ex- tinct family Cheirolepidiaceae is the most likely candidate (Azar, 2000). Lebanese am- ber has yielded a diverse arrray of arthropod orders and families, with many representing the oldest definitive records of their group (Azar, 2000). This amber is far more brittle and fractured than any other amber, so prep- arations must be done very carefully.
Inclusions in amber are best observed by grinding and polishing a flat surface close to the inclusion, which reduces most distortion or obscurity from the amber matrix. Unfor- tunately, unlike soft Dominican amber or hard Baltic amber, most Cretaceous ambers are fractured and brittle, so in order to best observe inclusions the material must be treat- ed prior to trimming and polishing. Pieces from the AMNH collection were embedded in an inert, optically high-quality epoxide resin under vacuum, before any trimming and polishing. The procedure was described in Nascimbene and Silverstein (2000). This technique fills in cracks that otherwise may split through an inclusion during preparation or which obscure observation with reflective surfaces. Even the hard and remarkably du- rable Burmese amber, which is easily pol- ished, is permeated by fractures, so its prep- aration and study is greatly improved by ep- oxy vacuum-embedding. Lastly, epoxy em- bedding protects the amber from the normal disintegration that occurs via long-term ex-
NO. 3412
posure to oxygen and other atmospheric con- ditions (Grimaldi, 1993).
MORPHOLOGICAL TERMINOLOGY
Names of morphological structures gen- erally follow Snodgrass (1935). For wing ve- nation terminology the system of Kukalova- Peck (1991), used by Nel and Roy (1996), was not used. Snodgrass’s (1935) venation system, which is a refinement of the classic Comstock-Needham system, was used in- stead. Good justification for use of the Snod- grass system specifically for Mantodea was presented in an early study on mantis wing venation (Smart, 1956). In fact, Smart pre- sented compelling evidence for interpretation of the CuP veins in roaches and mantises. Another important feature of mantis wings, besides venation, is the presence or absence of a thin, sclerotized area obliquely running near the basal branches of veins M and Cu. This feature has been given several names, with the one used here being “‘pseudovein’’.
A hallmark feature of mantises is the pair of raptorial prothoracic legs armed with modified setae. To clarify ambiguity about these various kinds of setae, the following terminology is used throughout this paper (see fies):
Setae are socketed, hairlike, unsclerotized struc- tures, being usually long and fine.
Scales are socketed, flattened setae which can be fan-shaped, lanceolate, or paddle-shaped, with the thickest portion always being several times the width of the base. Ribbing that typically occurs in all setae are particularly well defined on scales.
Spines are socketed, sclerotized structures, slight- ly to considerably thicker than setae.
Spinules are socketed or apparently unsocketed structures that are very short and stout and have a fine tip.
Spurs are heavily sclerotized, thick spines that have the basal articulation virtually fused to the surrounding cuticle. Spurs often sit atop a tu- bercle. These are not to be confused with the structures called spurs in Diptera, with are spines in the membrane of the tibial-tarsal joint.
Acronyms throughout the paper refer to the following repositories:
AMNH American Museum of Natural History, New York
2003
ae
setae scales spines JS d do TS spicules spur
Fig. 1. Significant setal modifications on man- tis forelegs and terminology used in this study.
PIN Paleontological Institute of the Russian Academy of Sciences, Moscow Staatliches Museum fiir Naturkunde, Stuttgart
SMNS
SYSTEMATICS
One goal of this work is to examine the relationships of Mesozoic and living mantis- es, and a revised classification reflecting these relationships is presented at the end of this paper. Taxonomy and definitions of taxa are treated first, in alphabetical order of gen- era.
GRIMALDI: EARLY MANTISES 7
Ambermantis, new genus
DIAGNOsIs: Spination of forefemur similar to Mantoida, except that Ambermantis ple- siomorphically lacks discoidal spines. De- rived features are the following: extremely long foretarsi, forebasitarsus longer than for- etibia. Extremely long hindlegs, with length of hindtibia plus tarsi equal to length of body; cerci long, with 20 segments.
TYPE SPECIES: A. wozniaki, n. sp., Creta- ceous of New Jersey.
INCLUDED SPECIES: Monotypic.
EtyMo_Locy: Directly from amber, itself a derivative of ambra (L.), in reference to the mode of preservation of three known speci- mens, all adults.
Ambermantis wozniaki, new species Figures 2a, 3, 4
Archimantis zherikhini Vrsansky, 2002a: 6 (mis- identification of specimen AMNH NJ90cc: see comments below).
Jantarimantis zherikhini (Vrsansky), 2002b: 1 (re- placement name for preoccupied Archimantis Saussure).
DIAGNOsIs: As for genus.
DESCRIPTION: Taken largely from holotype specimen, which is nearly complete. Body length of holotype Gncluding cerci) 15 mm, some coloration patterns preserved. Holotype specimen only missing portions of antennae, left mid- and hindlegs. Species identity of paratypes based on similar wing venation. Paratype specimen (AMNH NJ-90cc: fig. 2b, 4c) has wings, most of the pronotum, and dorsal part of the head preserved, with total body length (from front of head to tip of wings, excluding antennae) of 13 mm, fore- wing length 9.5 mm. Specimen of a forewing exposed on surface of the amber may belong to this species; it was studied using an SEM (fig. 4a, b).
Head: Considerably wider (by approxi- mately1.5X) than pronotum; in frontal view overall shape triangular, with broad vertex and narrow oral region. Eyes very large, with broad frontal field, inner margins close to scape. Eyes egg-shaped in frontal view, nar- row end ventrally; eyes round in lateral view. All 3 ocelli present, very close, large; median one slightly smaller and lying nearly between scapes. Pair of small, blackish, ovate stig-
8 AMERICAN MUSEUM NOVITATES NO. 3412
K “ae 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. ec. Jersimantis burmiticus,
holotype AMNH Bul170, in Burmese amber. d. Burmantis asiatica, holotype, in Burmese amber. For scales, refer to illustrations of specimens.
2003
GRIMALDI: EARLY MANTISES 9
{ lateral
Figs.
wf Rass “right cercus
es 1.0mm
/ S styli d.
1.0mm
labrum
r mandible
iE galea j-<—— maxillary palp
Ut aT
lacinia labial palp
Ambermantis wozniaki, new species, holotype (AMNH NJ1085), in New Jersey Cretaceous
amber. a. Habitus of entire animal, oblique left lateral view. b. Left foreleg, ventral view of femur with folded tibia and basitarsomere. c. Foretibia, showing spination of mesal edge. d. Frontal view of head.
e. Male genitalia.
mata occur just below median ocellus and between scapes. Antenna flagellate and ex- tremely long, longer than body (16 mm); fla- gellomeres filiform and gradually tapering in diameter apicad. Mouthparts fully discern- able. Clypeus relatively shallow, depth half that of labrum. Labrum roughly triangular in shape, with shallow lobe in middle. Left mandible, pair of well-developed galeae and laciniae present. Maxillary palp 5-segment- ed, total length quite long, approximately equal to length from tip of labrum to base of antenna. Labial palp 3-segmented, approxi- mately half the length of maxillary palp, api- cal palpomere with tapered, darkened tip.
Thorax: Pronotum with dark maculations, length approximately 1.2—1.3X the width, with distinct transverse groove and scattered pimples on dorsal surface. Lateral margins of pronotum turned downward, covering per- haps half of pleura; posterior margin slightly upturned. Forelegs with coxa and femur hav- ing maculated pattern, as figured. Forecoxa long, length 0.6 that of femur. Forefemur stoutest of femora, distal half 0.4 thickness of basal half of femur. Spination of forefemur seen best in left leg of holotype. Ventral sur- face of forefemur with 2 rows of spines; a row of 5 spines on lateral edge, distal spine half the size of others; row of 10 spines on
10 AMERICAN MUSEUM NOVITATES
NO. 3412
Fig. 4. Ambermantis wozniaki, new species, in New Jersey amber. a, b. Scanning electron micro- graph of an isolated, imprinted wing on the surface of the amber (a), with diagram of the venation (bd). c. Paratype, AMNH NJ90cc, a cast/imprint of the dorsal half of the specimen on the surface of the
amber; the ventral half was missing.
mesal edge, slightly smaller than lateral spines. Forefemur with groove to receive api- cal spur of tibia, and patch of fine setulae (the ‘‘brush’’) on mesal surface near distal end; another patch of fine, but longer setulae on ventromesal surface. Foretibia 0.6
length of femur; with large apical spur hav- ing basal articulation to tibia barely discern- able, length of spur 0.2 length of entire tib- ia. Forebasitarsus articulated on tibia consid- erably preapically, articulation point near penultimate spine. Foretibia with 2 rows of
2003
spines on ventral (adpressed) surface, lateral row of 4 spines and mesal row of 7 spines. Tibial spines are thin and sharp, gradually shorter proximad. Foretarsi with dark band- ing, very long, longer than forefemur; fore- basitarsomere longer than tibia, with small apical spur. Right midleg of holotype mostly preserved; coxa and base of femur with dark maculation; midtibia slightly longer than midbasitarsomere. Midfemur stout, nearly as thick as forefemur. Hindlegs extremely long; tibia and tarsus equal in length to body, hind- tibia equal to length of hindfemur + tro- chanter, with small apical spur. Hindfemur slender, with small, sharp, slightly curved spiniform seta projecting from apex. Hind- femur and tibia with dark banding at each end.
Wings long and narrow, extended to tip of abdomen but with cerci projecting. Forewing with dark maculations (not illustrated), length approximately 4.5x< the width. Hindwing not visible. Venation derived from holotype and paratype AMNH NJ90cc. Fore- wing: Sc long, approximately 0.7 length of wing, with 9 crossveins joining to C; R with single, short dichotomous fork at apex; M with 3 significant branches (best seen in NJ90cc), first fork near middle of wing, sec- ond fork between first one and wing apex; Cu, with 5—7 main branches, a long inter- calary vein present between each; vein 1V (first vannal vein [Smart, 1956]) present, curved, not fused to Cu,; CuP fused to Cu,. Numerous short crossveins present.
Abdomen: Cerci long, with 20 segments, segments longer apicad. Subgenital plate typ- ical, pair of short styli on its posterior mar- gin; ventral lobe large, apical process typical of Mantodea.
TYPE SPECIMENS: Holotype: male, AMNH NJ1085, NEW JERSEY: Middlesex County, Sayreville, Raritan Formation, Upper Creta- ceous (Turonian) (Grimaldi et al., 2000b), collected by Joseph Wozniak. Specimen is in a piece of turbid, light yellow amber (fig. 2a), which had been trimmed to a size just slight- ly larger (18 X 8 X 6 mm) than the mantis, for optimal viewing. Still, some portions were obscured by turbidity or by milky froth coating parts of the specimen. The amber piece had deep cracks in it, so it was embed- ded in epoxy after trimming, then re-embed-
GRIMALDI: EARLY MANTISES I<k
ded. All epoxy was trimmed within 1 mm of the surface of the amber. Paratype: AMNH NJ90cc, NEW JERSEY: Middlesex County, East Brunswick, Raritan Formation (Grimal- di et al., 2000b). This is a partial specimen comprised of a dorsal impression of head and wings (figs. 2b, 4c), and it is one of over 30 inclusions found in a remarkable piece of amber, including the oldest definitive mush- rooms (Grimaldi et al., 2000).
ETYMOLOGY: Patronym, in honor of the collector and donor of the beautiful holotype specimen, Joseph Wozniak. This is the larg- est insect preserved in New Jersey amber, though portions of what were larger insect specimens also occur in this amber.
COMMENTS: The holotype specimen (AMNH NJ1085) was mentioned and figured in Gri- maldi et al. (2000b: fig. 43g) and is the best- preserved mantis from the Mesozoic. It super- ficially resembles Mantoida, but lacks impor- tant synapomorphies, such as the discoidal spines that occur on the forefemur of virtually all living mantises. Spination of the forefemur, in fact, is relatively simple. Ambermantis also has several odd features, perhaps the most striking being the extremely long hindlegs. In most mantises, even extremely gracile ones, the length of the hindtibia + tarsus is generally about one-half the body length. Eremiaphila is exceptional because the body is very short and stout and the legs stiltlike, an extreme special- ization for dwelling in sandy habitats. No other mantis has hindlegs as long as Ambermantis. Less exceptional but still distinctive are the long cerci in Ambermantis. The number of cer- cal segments in Ambermantis is 20 and in Chaeteessa 23—25, but in most other mantises there are 8—12, including the most basal fossil forms, where the cerci are preserved (in Eve- miaphila the cerci are quite reduced). How- ever, there are species in the Mantidae that also have many cercal segments (i.e., Theopompel- la, 26-28; Choerododis, 20—22).
Jantarimantis zherikhini (Vrsansky) was very recently described as a mantis on the basis of two incomplete specimens in New Jersey amber in the AMNH (he originally used the preoccupied generic name Archi- mantis) (Vrsansky, 2002a, b). The two spec- imens are not even in the same order. What Vrsansky called specimen “M1” (the holo- type of J. zherikhini) is actually a roach of
12 AMERICAN MUSEUM NOVITATES
the extinct, unusual family Umenocoleidae. A small series of completely preserved umenocoleid adults in New Jersey amber are in the AMNH collection, so all of these, in- cluding the holotype of Jantarimantis zheri- khini, will be treated in another paper. What Vrsansky labelled ““AMNH M2” and as “‘ad- ditional material’ (not even as a paratype) is specimen AMNH NJ90cc, properly de- scribed herein and named as a paratype of Ambermantis wosniaki. During Vrsansky’s 1995 visit to the AMNH, the completely pre- served holotype of Ambermantis wozniaki had not yet been discovered.
Genus Amorphoscelites Gratshev and Zherikhin
Amorphoscelites Gratshevy and Zherikhin, 1993: 163. Type Species: A. sharovi Gratshev and Zherikhin, 1993: 163 (Early Cretaceous, Rus- sia). By original designation.
DIAGNOsIS: Poorly known genus based on a single isolated foreleg (PIN 3064/8586), originally defined by Gratshev and Zherikhin on the basis of the following significant fea- tures: coxa long; femur fairly stout, 3 times as long as broad (length, including trochan- ter, 4.6 mm), inner surface sculptured and with flat tubercles, ventrally with two longi- tudinal rows of small spines. Tibia fairly long (ength, including apical spur 0.7 length of femur), inner surface with numerous, minute denticles or spicules; apical spur large, with tarsus articulated at its base.
INCLUDED SPECIES: Monospecific.
COMMENTS: The specimen is clearly man- todean but far too incomplete for phyloge- netic analysis or classification. Structure of the foreleg differs from Amorphoscelidae by this family possessing a small spine in the middle of the femur and with a very long tibial spur, and often possessing few if any other spines (even minute ones). Amorphos- celites has no such spine preserved, numer- ous fine spines or spicules, and the foretibia is of standard size for mantises.
Genus Baissomantis Gratshev and Zherikhin
Baissomantis Gratshev and Zherikhin, 1993: 159. Type Species: B. picta Gratshev and Zherikhin,
NO. 3412
1993: 159 (Early Cretaceous, Russia). By orig- inal designation.
DIAGNOSIS: Known only as isolated wings from the Cretaceous of Russia, and defined by Gratshev and Zherikhin largely on the ba- sis of the following significant features: R with 1 or 2 branches, ending at costal margin just beyond Sc; RS separate from R, multi- branched; M with 2 or 3 branches; Cu, strongly arched. Wings with patterning. Sub- sequent study by the author indicates a pseu- dovein is absent.
INCLUDED SPECIES: B. picta; B. maculata Gratshev and Zherikhin, 1993 (figs. 5a, 6).
COMMENTS: The absence of a pseudovein, though not mentioned by Gratshev and Zher- ikhin, is highly significant, and would make this genus plesiomorphic to true mantises.
Burmantis, new genus
DIAGNOSIS: Distinguished from other gen- era known as nymphs in amber (Chaetees- sites, Electromantis, Jersimantis) most read- ily 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 fore- tibia; at least midocellus present (these two features observed only for the type speci- mens)
TYPE SPECIES: B. asidtica, n. sp.
INCLUDED SPECIES: B. asiatica, B. lebanen- sis, N. Sp.
EtymMo.Locy: 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
D1AGNosIs: Differs from Burmantis leba- nensis aS given in the diagnosis of that spe- cies, below.
DESCRIPTION: Based on a single specimen,
2003
Fis5.
GRIMALDI: EARLY MANTISES 13
waa” Sey ae ie SS + ie te “eat " ae * i = | o- eee PNP conae i ee Tat | ae ‘ Sst _ “3
a re iy ES Pap ST hy mj Fe. e b, 5 es - " = he 2 . js # | 4
\ .
Wings of Eurasian Cretaceous mantises. a. Baissomantis picta Gratshev and Zherikhin, PIN
1989/2486. b, ce. Cretophotina spp. b. Cretophotina tristriata paratype, PIN 1989/2487. ¢. Cretophotina tristriata holotype, PIN 3064/8585. Original photographs.
14 AMERICAN MUSEUM NOVITATES NO. 3412
pseudovein
Cretophotina tristriata
forewing
Cun r CuP
Baissomantis
maculatus hindwing
2003
which is a nymphal exuvium. Since the cu- ticle is cleared, and with the thin amber prep- aration made, it was possible to study the specimen under compound microscopy at 100. Portions of the body are collapsed and difficult to reconstruct, but even coloration patterns and microscopic structures like sen- silla are observable. Head: Eyes large, but only partially preserved. Median ocellus pre- sent but lateral ones not apparent (perhaps a preservational artifact). Frons slightly bul- bous; 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 G.e., slight asymmetry) not pos- sible. Labial palps preserved, 3-segmented; maxilla preserved, lacinia sharp and sclero- tized, toothlike. Maxillary palps not pre- served or apparent. Antenna long, flagellate; scape with a thin sclerite in the socket mem- brane ventrally; pedicel rounded apically; fla- gellomere | 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 ante- pronotum 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 scler- ites. 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 ven- trolateral margin. All trochanters small. For- efemur 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-
<
GRIMALDI: EARLY MANTISES 1 Be,
face of bulbous base of forefemur, and ven- tral surface of basal half of forefemur, with dense, fine, erect pubescence. Ventrolateral edge of forefemur with 3 long, stiff, sclero- tized spines, their lengths approximately equal to width of femur; each spine on a low tubercle. Ventromesal edge of forefemur, dis- tal 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 pre- sent. Mesal surface of forefemur with patch of scattered spicules (“‘brush’’) near middle; spicules are slightly thickened, but not scal- iform as in all living Mantodea (i.e., fig. 8a). Foretibia with 2 ventral rows of spines on distal two-thirds of tibia; mesal row with 8 thicker spines, lengths of which gradually in- creased 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 tib- ia that projects beyond tarsal articulation (fig. 8a), as occurs in all Mantodea except Chae- teessa. 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 remain- ing tarsomeres. Length of hindtibia and tar- sus 4.8 mm, approximately same length as body exclusive of cerci.
Abdomen: Short, broad, tergites with mi- nute, spiculelike setulae. Cerci well devel- oped, 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
Fig. 6. Wings and venation of selected Mantodea from the Lower Cretaceous of Russia. Cretopho- tina tristriata (forewing and portion of hindwing), and Baissomantis maculatus (fore- and hindwings).
Original drawings.
16 AMERICAN MUSEUM NOVITATES NO. 3412
J pedicel
*)<—|abrum
mandible b.
Fig. 7. Burmantis asiatica, new species (holotype). a. General outline of body. b. Frontal view of face.
2003 GRIMALDI: EARLY MANTISES 17
basitarsomere
femoral brush
ventromesal spines
ventrolateral spines
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.
18 AMERICAN MUSEUM NOVITATES
near Tanai and Myitkyina. The specimen is in a clear yellowish piece of amber contain- ing scattered bits of debris, stellate tri- chomes, two staphylinoid beetles, and a ber- othid neuropteran. The piece is slightly rect- angular, 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 local- ity of the fossil.
COMMENTS: Exquisite preservation of the forelegs reveals a tibial spination that is ple- siomorphic: there are no discoidal spines, the setulae of the forefemoral brush are not par- ticularly 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 re- spect: a long forebasitarsomere (character 13, below).
Burmantis lebanensis, new species Figure 9
DrAGNosis: 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 spi- culelike setulae; cerci shorter and with 9—10 (vs. 12) segments, and without very long se- tae 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. asia- tica; first flagellomere long, length about equal to 4—5 other, basal flagellomeres. Length of flagellomeres gradually and great- ly increased distad. Thorax: Pronotum too distorted to reconstruct shape, but it and sev- eral other sclerites covered with small irreg- ular 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 sur- face with dense, fine pubescence on proximal half; femur with two ventral rows of spines, ventromesal row with 5 thick, short, sclero-
NO. 3412
tized spines alternating with 4 smaller, less sclerotized ones; ventrolateral row with 3 long spines, a minute one distally. Forefe- moral brush present (seen vaguely in dorsal view of specimen), but details (i.e., number and shape of scales) not visible. Only prox- imal 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 fair- ly stout; midtibia very thin, length equal to that of midfemur. Length of midtarsi equal to length of midtibia; length of hindtibia ap- proximately 1.6 length of midtibia. Abdo- men: 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 Bcharré, collected by An- toun Estephan (Early Cretaceous, approxi- mately Barremian). The amber piece is clear, transparent yellow; it was embedded in ep- oxy and trimmed to separate one piece con- taining a scelionid wasp, the other containing the mantis nymph and another parasitoid wasp.
ETYMOLOGY: Referring to Lebanon, the source country of the Lower Cretaceous am- ber.
COMMENTS: Foreleg structure of this spe- cies 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 [Santon- ian]). By original designation.
2003
19
GRIMALDI: EARLY MANTISES
LP NN
/ i.
)
ff
4 bho {> iy ops /
My YAY) h, wh i
He
wo S\ SN
rot
coxa
trochanter
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.
20 AMERICAN MUSEUM NOVITATES
DIAGNOSIS: Known only as a partial nymph in Cretaceous amber from northern Siberia, defined mostly on the basis of dis- tinctive 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 “‘col- lective’? 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 am- ber, preserving details of the foreleg (fig. 11). Preservation of the specimen does not allow observation of ocelli, so their presence is un- confirmed. The forefemoral brush is appar- ently absent, but this is difficult to be certain of given preservation of the specimen.
Genus Cretomantis Gratshev and Zherikhin
Cretomantis Gratshev and Zherikhin, 1993: 161. Type Species: C. larvalis Gratshev and Zheri- khin, 1993: 161 (Early Cretaceous, Russia). By original designation.
DIAGNOSIS (revised from Gratshev and Zherikhin): The genus is based on the com- pression of a single but complete nymph, which apparently is an exuvium (PIN 3064/ 8511, holotype, figs. 12, 13). A stout-bodied nymph with forefemora stout and apparently having a ventral furrow, with furrow bor- dered mesally with row of 8—12 spines and 3—4 short spines or spicules. Mesal row of femoral spines preserved as small, rounded mounds, probably small tubercles that were the bases of spines. Foretibia short (0.7X length of femur) and stout, with large apical spine or spur (basal articulation obscure); with mesal row of 8—10 stout spines, lateral row of 5 smaller spines. Forebasitarsus ex- tremely short, ca. 0.20 length of foretibia. Midfemur with 2 ventral rows of spicules or minute spines; hindfemur short and _ stout, only 1.2 length of foretibia. Hindtibia only ca. 1.1 length of hindfemur. Cerci short, approximately same size as styli.
INCLUDED SPECIES: Monotypic.
NO. 3412
COMMENTS: Several aspects of the original diagnosis were found to require some revi- sion (Gratshev and Zherikhin, 1993: 161): only one apical tibial spine/spur is preserved, so “‘apical pair [of spines] strongly differ- entiated”’ cannot be corroborated; the large, apical spine of the tibia is not “‘placed be- yond tarsal articulation’’; and the foretarsi are not longer than the tibia.
Head structures are difficult to determine, which may be due to the specimen being an exuvium with a crumpled cuticle. Three ter- minal abdominal appendages are preserved in the specimen, a pair of finer ones with at least 5 segments, and a slightly thicker one (presumably one of a pair). Distinguishing styli from cerci is ambiguous.
Genus Cretophotina Gratshev and Zherikhin
Cretophotina Gratshev and Zherikhin, 1993: 150. Type Species: C. tristriata Gratshev and Zher- ikhin, 1993: 150 (Early Cretaceous, Siberia). By original designation.
DIAGNOsIS: Known only on the basis of wings from the Cretaceous of Eurasia, de- fined originally by Gratshev and Zherikhin on the following significant features: costal field distinctly wider than field between Sc and R; R apically with 5—8 terminal branch- es; M with 2—3 branches; CuA with 6—10 terminal branches, posterior branch separated from main stem; Cu, distinctly curved. Re- examination by myself indicates the pseu- dovein is present.
INCLUDED SPECIES: C. tristriata (figs. 5c, 6); C. mongolica Gratshev and Zherikhin, 1993; and C. serotina Gratshev and Zheri- khin, 1993.
COMMENTS: Gratshev and Zherikhin (1993) omitted mention of a very important feature in Cretophotina: the presence of a short pseudovein near the basal forks of M and Cu, (figs. 5b, c; 6), found in all mantis wings save Baissomantis. The paratype of Cretophotina tristriata (PIN 1989/2487) and the holotype (PIN 3064/8585) (fig. 6) have venational differences that strongly suggest different species. The paratype has M 3- branched (vs. 2), the 2™ vein of Cu, with a short, apical branch (vs. none), the basal branching of Cu, dichotomous (vs. apparent-
2003 GRIMALDI: EARLY MANTISES mal
Fig. 10. Photomicrographs of holotype of Chaeteessites minutissimus Gratshev and Zherikhin, a partial nymph in Siberian amber, holotype PIN 3311/603. a. Entire specimen. b. Detail of foretibia. Original photographs.
22 AMERICAN MUSEUM NOVITATES
Fig. 11.
ly pectinate in the holotype), and CuP is complete (vs. incomplete).
Genus Electromantis Gratshev and Zherikhin
Electromantis Gratshev and Zherikhin, 1993: 162. Type Species: E. sukatshevae Gratshev and Zherikhin, 1993: 163 (Late Cretaceous, Sibe- ria). By original designation.
DIAGNOSIS: Known only as partial remains of a nymph in amber (fore- and midlegs, ven- tral portions of head and thorax)(PIN 3631/ 7), with the following distinctive features: forefemur incrassate (greatest width 0.3 x the length), with pubescent ventral furrow, bor- dered by 2 rows of fine spines; foretibia short, length (excluding apical spine) 0.5X length of femur, with one large and one smaller apical spines. Larger apical tibial spine 0.6X< length of tibia; smaller apical spine 0.6 length of larger one; apical spines
NO. 3412
basitarsomere
spines
fore femur
0.1mm
Drawing of Chaeteessites minutissimus holotype.
at apex of tibia (no projection beyond artic- ulating bases). Foretibia with 2 rows of ap- proximately 7 small spines, increased in size distad. Forebasitarsomere slender, slightly shorter than tibia.
INCLUDED SPECIES: Monotypic.
COMMENTS: Proportions of the forefemur and foretibia, and spination of each, distin- guish this genus from Amorphoscelites, Bur- mantis, Cretomantis, and Jersimantis.
Genus Jersimantis Grimaldi
Jersimantis Grimaldi, 1997: 6. Type Species: J. luzzii Grimaldi, 1997: 6 (mid-Cretaceous [Tu- ronian] of New Jersey) (fig. 14 herein). By orig- inal designation.
DIAGNOsIS (emended): Plesiomorphically as in Chaeteessites minutissimus, with fore- femur having ventral row of 3—4 long, fine, stiff setae (no spines); apex of foretibia with two spines (one large, one small), having
2003 GRIMALDI: EARLY MANTISES 23
~*~ * - * e * i a = 1 ‘ * -. fi Pf # é «* 3, - —_ 4 Pe ? ; | 4 J ~ * = . r. \ 7 oT] he | « \o “ { é * ~ Fj /? Pa - = 7 &@ a hall a ™ “iy age q 2 ee a. "* “ | i] — (| a , € are; ¥ | bp . ian, J r > =~ a a Ma - ne i. , % : a 3 re bas Te! —e “f . > . ee Nas =a = 2 Li. =« +" “* ] # &
Fig. 12. Photomicrograph (original) of Cretomantis larvalis Gratshev and Zherikhin, holotype (PIN 3064/8511).
well-defined articulation points but no spur Jersimantis burmiticus, new species
at the apex of a tibial extension. Differs from Figures 2c, 15
Chaeteessites by lacking a medial row of for-
etibial spinules (instead there are just fine, Diacnosis: Differs from J. luzzi by having
stiff setae); differs from Amorphoscelites, bulbous vertex with finely reticulate (vs. Cretomantis, and Electromantis by having a smooth) surface; pronotum with pair of low slender forefemur and by spination of the paramedian ridges (vs. none); stiff foretibial foreleg. Apomorphically with vertex bul- setae thicker; ventral surface of forefemur bous, ocelli absent. without fine, dense setulae; cerci with 10
24 AMERICAN MUSEUM NOVITATES
fore bias “
{
apical spine | 7 fore femur ——*;
bases of spines
Fig. 13.
segments (vs. 3) that are highly differentiated (described below).
DESCRIPTION: Based entirely on the unique, nymphal specimen, which is completely pre- served save for distal flagellomeres (lost at surface of amber). Body length 3.50 mm, in- cluding cerci. Specimen is observable dor- sally, ventrally, and frontally. Portions of the cuticle are transparent, allowing observation
NO. 3412
Drawing (original) of Cretomantis larvalis holotype, showing details.
of some usually microscopic or concealed features (1.e., absence of forefemoral brush).
Head: Broad, approximately 1.7 width of pronotum. Eyes large, exophthalmic, with large frontal field. Vertex bulbous, having fine pattern of reticulations on surface; ocelli absent. Mouthparts largely obscured, all but basal segments of palps lost.
Thorax: Pronotum comparatively small
2003 GRIMALDI: EARLY MANTISES 25
Fig. 14. Nymph in New Jersey amber, Jersimantis luzzii holotype (AMNH NJ425) (from Grimaldi, 1997).
26 AMERICAN MUSEUM NOVITATES
(saan 9 eee | FI 0.50 mm
Fie. 15: mid-Cretaceous amber from Myanmar.
for mantises, length only slightly more than (1.25X) the width; transverse groove pre- sent; with pair of slight, paramedian ridges extended approximately three-quarters the pronotal length. Pronotum with sides curved slightly downward, though not extended along pleura. Legs: Robust, hind pair lon- gest (approximately 1.7X length of fore- legs). Foreleg with coxa small, forefemur stout (though no thicker than midfemur); tibia and tarsi slender. Forefemur approxi- mately 1.5 the length of foretibia, 1.3 the length of foretarsi. Forefemur with ven- tro-lateral row of 4 long, fine, stiff setae; no thick setae or spines, nor dense patch of fine setae on inner surface. Foretibia with two ventral rows of sharp, stiff setae; setae on inner row thicker, spiculate, approximately 6 in row; apex of tibia with pair of thick, stiff, long setae with well-defined articula- tion. Inner foretibial spine longest, length approximately 3X the width of tibia. All legs with 5 tarsomeres, tarsomere 4 with pulvillar lobe extended ventrally 0.4x length of pretarsus. Midleg with short, fine setae; tibia with pair of fine apical setae on
NO. 3412
Jersimantis burmiticus, new species (holotype), AMNH Bul70, oblique dorsal view, in
ventral surface, length of longest seta twice the width of tibia. Hind legs long and slen- der, lengths of femur and tibia equal.
Abdomen: Only 8 segments visible. Pair of short styli between cerci, attached to ter- minal sternite. Styli extended slightly past midlength of first cercal segment. Cerci very distinctive: 10 segments, basal segment large, nearly one-third length of cercus, with whorl of 4 long, fine, stiff setae at apex; apical 9 segments with basal one largest, having whorl of 4 small setae at apex; distal 8 segments small, tapered apicad to fine point.
TYPE SPECIMEN: Nymph, AMNH Bul70, MYANMAR: Kachin, from mines near Tan- ail, ex: Leeward Capital Corp. 1999. The specimen is in a piece of dark, transparent amber, 15 X 14 X 7 mm, which contains 9 other arthropod inclusions: | Cecidomyidae, 1 Psychodidae, 2 Chimeromyia (Diptera), 2 Auchenorrhyncha, 1 Coleoptera, 2 larvae. The mantis nymph is slightly distorted by dorsoventral compression of the body and frontal compression of the head.
ETYMOLOGY: From Burma (Myanmar).
2003
Genus Kazakhophotina Gratshev and Zherikhin
Kazakhophotina Gratshev and Zherikhin, 1993: 156. Type Species: K. corrupta Gratshev and Zherikhin, 1993: 156 (Upper Cretaceous, Ka- zakhstan). By original designation.
DIAGNOsIS: Known only as a portion of a wing (PIN 2383/150), defined originally by Gratshev and Zherikhin on basis of the fol- lowing most significant features: costal field (between Sc and C) distinctly wider than subcostal one (between Sc and R); no inter- calary veins between Sc and R; R with 6 apical branches; M 2-branched; CuA with 4 or more branches.
INCLUDED SPECIES: Monotypic.
COMMENTS: The unique specimen on which the genus is based is too incomplete and distorted to include in a phylogenetic analysis and classification.
Santanmantis, new genus
DIAGNOsIS: A primitive type of mantis with tips of wings apomorphically extended well beyond apex of the abdomen (by more than one-third the wing length); venation re- duced, 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 distinc- tive is the very long pseudovein: instead of a sclerotized area restricted to the basal fork of M and Cu,, it is a tubular vein extending from this region through veins CuA,, 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 oviposi- tor) 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.
GRIMALDI: EARLY MANTISES 27
Santanmantis axelrodi, new species Figures 16—24
DrtAGNnosis: As for the genus, given above.
DESCRIPTION: Gross aspects of ventral structures were observed using HRCT scans of the holotype specimen (figs. 18, 19). Mea- surements of various parts are given in table 2. Specimens from the SMNS (Staatliches Museum fiir Naturkunde, Stuttgart) have pro- visional numbers.
Head: Antennae filiform, at least basal 8— 10 flagellomeres with lengths 2.5 the width; scape and pedicel small. Eyes large, situated frontally and somewhat laterally, with a large postoccipital space. Distance be- tween eyes wide, equal to width of eye. Ocelli present, but seen in only one specimen (SMNS 172). Head hypognathous, mouth- parts (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—0.75 X its width (as seen in AMNH 1957, SMNS 112, and 174), the surface evenly covered with fine punc- tations (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 para- types, appears due to preservational differ- ences. Forelegs observed using HRCT on ho- lotype: held frontally, tibiae and femora fold- ed against each other, femoro-tibial 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 foreti- bia apparently with a spur, though cannot discern whether the spur has a well-defined articulation (1.e., fig. 19). Forecoxae not vis- ible. 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 hind- tibiae, but row of at least 4 ventral spines occur on midfemur (visible dorsally). Fore- wings 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,, CuP, and anal veins and
28 AMERICAN MUSEUM NOVITATES NO. 3412
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.
2003 GRIMALDI: EARLY MANTISES
1.0 mm a5 _spronotum . foreleg amid leg
mid femoral
crop contents
vid
& 7
forewing.
Ly *
Fig. 17. Illustrated rendering of Santanmantis axelrodi, holotype, with detail of terminalia.
30 AMERICAN MUSEUM NOVITATES NO. 3412
Fig. 18. High-resolution 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 param- eters.
2003
GRIMALDI: EARLY MANTISES a1
Fig. 19. of complete specimen, exposing more of the long, slender hindlegs. Right: a more detailed, surface- rendered 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 pm), not the actual preservation. The bases of some spines on the foretibia and femur are visible.
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 homon- omous, though anal regions (1.e., presence of expansive fan on hindwing) were not pre- served. 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, with 4 main branches, bases of 2 most proximal branches very close. Claval furrow at CuA, 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,; A with two main branches. Only a portion of
High-resolution CT scans of holotype of Santanmantis axelrodi. Left: a volume-rendering
hindwing tip was preserved (SMNS 112: fig. 2S),
Abdomen: Relatively short and stout, length approximately 1.3 the width. Con- tents 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 seg- ments (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
aD AMERICAN MUSEUM NOVITATES NO. 3412
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.
2003
Fig: 2-1,
GRIMALDI: EARLY MANTISES 33
c
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. Giinter Bechly (SMNS).
complete specimen, though the wing vena- tion 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 para- types, 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 por- tions of forewings excellent, showing signif- icant 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-
34 AMERICAN MUSEUM NOVITATES
NO. 3412
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.
tiful specimen with dorsal surface preserved; forewings spread and nearly completely pre- served; hindwings folded and covered be- neath abdomen. Pronotum preserved (fig. 22b); portions of right midfemur exposed and most of right hindtibia and tarsus. Ab- domen well preserved, though cerci barely discernable.
Paratype, SMNS 113 (fig. 21c): A beau- tifully 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 evi- dent. Dorsal portions of head damaged. Pro- thorax appears to be saddle-shaped with an- terior edge emarginate. A portion of what ap- pears to be a midleg protrudes from under
2003 GRIMALDI: EARLY MANTISES 35
pseudovein
* Pompe. £ oy : Meee oe neat
:
ee Ape nnes re egse tens paovnsrnsecensssssneeennernenscee ne eee nee ns hae 3 5%
? ae
eneee
: seo i aevonpessaanccan? .
:
au
.
.
; ek cee an crete eosenesnetagenen ons sandege es t } 3
* * .
Byevee
dee
anne see .
aes ‘ pane eaenee ee seces teens ase
ee eset ef.
. Pe mereee ome masncene mee . oe
agh et acen secur ccees eatsatre .
oe ‘t . segestess ap teens wee WN oe ooen ’
AY o . H soe e esa ee et of i 3 3 : davon aera geben sea oo ovens ns s ones : or Se ee ae anaes tees. Caan ail < Hy 7 4 ’ ~ . : natn te . 3 . : . * : = : \ 4 ° 5 : . : : 8 on ‘ 3 ast 2d : : H 3 : wet : 5 2 a etenet™ 3 = 3 : . J ee z > z o . . : = . . 3 3. % 3 * . 8 * . 5 3 H e + * ¥ *. : . * * 3 : ‘ 4 -}————--4 :
hindwing SMNS 112 ea
: : Y
2
wee teneeys . ry . | ‘eae Em STARE ET s a } i :
Hy ae aee evergetons se eee oe .
: ee 7eete tt vee caneetignas me eed
AOC TOE TOUSEN EEO NES MOUNTS Mea anesen cenetnnencenem ett eesaes ‘ . POO ee en ereenatteenn ane
: Sesevessceces a=,
Cre
be, Pet nies Cod Sees spe tg 4 iy
ey ea ceveaeestene eanen typeeseeeey :
ees ta . . to he rel TE
Re pi eae SOR ES ete asthe angie on ogeh veges ¥ bar hs 1) sus
forewing
SMNS 115
Fig. 23. Wing venation of Santanmantis axelrodi paratypes, showing slight variation.
36 AMERICAN MUSEUM NOVITATES
1.0mm
CuP
NO. 3412
“pseudovein"
Fig. 24. Left wing of Santanmantis axelrodi paratype, AMNH 1956.
left forewing. Portions of what is probably left hindleg are exposed, including distal por- tion of femur; a long, thin tibia; and tarsi. Small spines occur along one edge of tarsi and apical portion of tibia. Forelegs not vis- ible; probably folded beneath head and pron- otum, buried in matrix.
Paratype, SMNS 115 (figs. 21a, 23): Headless specimen with ventral surface ex-
posed; right forewing is spread, revealing ve- nation. 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 (prob- ably 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)
AMNH SMNS 1957 1956 112 113 114 115 172
Body length (exclusive of cerci) 11.3 10.16 9.7 - 10.2 - 9.5 Pronotum width 2.4 2.35 2.0 - - - 22 Pronotum length 1.8 Ea 1.4 1.5 - - 1.5 Head width 2.5 2.64 2:2 - 2.4 - 2.7 Forewing length 12.0 - 11.8 13.0 - 12.0 11.1 Forewing width (greatest) - 2.83 3.1 3.5 - 3.2 3.1 Hindwing length 11.4 - 11.3 13.0 - - 10.8 Length wings extended past tip of abdomen 4.5 3.67 5.5 - 3.5 - 4.6 Legs
Midfemur length - 2.53 - - - 3.1 -
Midtibia length - 4.00 - - - 4.1 -
Midtarsus length - - - - - 29 -
Hindfemur length 4.63 - - - - 2a -
Hindtibia length 4.88 - 4.1 4.6 - - -
Hindtarsus length - - 2.3 - - - - Cercus length 1.4 - - - - - 1.0
4 Structures hidden in matrix, measured from HRCT images.
2003
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 reveal- ing a deep claval furrow [fig. 22f]). Left fore- wing outstretched, left hindwing and right fore- and hindwings folded over abdomen. Abdomen well preserved, including left cer- cus. 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 San- tana fossil insects and his generosity to the AMNH.
DISCUSSION: The holotype and five para- types 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 Cre- taceous mantis species. Santanmantis lacks synapomorphies distinct to all living mantis- es, including Mantoida and Chaeteessa, as given in the diagnosis. Santanmanitis 1s dis- tinct from Baissomantis (L. Cretaceous, Eur- asia), which has more dichotomous branch- ing in R (vs. pectinate), more branches in CuA, (5 or 6, vs. 4), a complete CuP (figs. 23, 24), and no pseudovein. The two groups, though, have distinct plesiomorphic similar- ities, particularly the strongly arched claval furrow—a condition intermediate between roaches and more derived mantises. The ex- tremely long wings (or, conversely, a very short abdomen) in Santanmantis are unusual, as most fully winged Mantodea and Blatto- dea have the tips of the wings extended to the apex of the abdomen or slightly beyond. In Santanmantis the wings extend well be- yond the abdominal apex by more than one- third the wing length. This condition is in-
GRIMALDI: EARLY MANTISES 37
termediate between what is found in Isoptera and the rest of the Dictyoptera.
A preserved and full crop in the type spec- imen offered an apparent opportunity to con- firm if the diet of Santanmantis was indeed predatory. Crop contents of mineralized in- sect fossils are sometimes well preserved (e.g., Krassilov and Rasnitsyn, 1999), and this was especially expected for this speci- men given the preservation of relief, of tis- sues, 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 DSM-1 SEM, in order to scrutinize for frag- ments 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 rec- ognized in these samples. Though Santan- mantis 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 Zher- ikhin, 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 fea- tures: wing with extensive dark patterns; R with single apical fork; M 2-branched, 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.
PHYLOGENY OF BASAL MANTISES
Characters used for a cladistic analysis were external, comprising features of the head, wings, and legs. Spination of the fore- legs accounts for significant characters; for- tunately, there is little ontogenetic change be- tween nymphs and adults in mantis spination. A more exhaustive search for characters
38 AMERICAN MUSEUM NOVITATES
would involve the male and female genitalia (i.e., LaGreca, 1954; Klass, 1997, 1998a) and even internal features (Klass, 1998b, for the proventriculus). Preliminary results indicate that there is significant variation in shape and structure of sclerites of the prothoracic and cervical region of Mantodea (D. Grimaldi, unpubl. data). Characters gleaned from this portion of the body must await a more de- tailed comparative study since most of these characters would not be observable in fossils, particularly because they are so intricate. The extreme asymmetry in dictyopteran male genitalia, for example, has even led to con- siderable controversy about homologous structures in this part of the body (e.g., Grandcolas, 1996; Klass, 2001), though the work by Klass on innervation and muscula- ture of genitalic structures has helped to clar- ify problems. Accurate identification of gen- italic features among a broad array of man- tises is a very large project out of scope for the present one. Moreover, many characters seen in the earliest mantis fossils (e.g., pres- ence/absence of forefemoral brush, well-de- veloped and strongly curved claval furrow) represent variation that does not occur in liv- ing mantises.
CHARACTERS
1. Blattodean-type discoid pronotum is re- duced, not covering the head. Plesiomorph- ically, it is as occurs in most living and Paleozoic roaches, which is large enough to shield most of the head or even some- times the whole head in dorsal view.
2. Forelegs raptorial, spiny, and folded under the thorax at rest, with associated movable forecoxa. Plesiomorphically, the forelegs are not differentiated from the others and are used in walking.
3. Eyes large, exophthalmic, with a large frontal field. Plesiomorphically, the eyes do not occupy the entire lateral surface of the head, nor are they bulging with a large frontal area, and they usually have the fronto-mesal margin emarginate.
4. Loss or great reduction of the claval fur- row, wherein vein CuA, runs in the teg- minous forewing. Plesiomorphically, this furrow is very distinctive and well devel- oped (e.g., figs. 20c; 22c, f).
5. Midfemur without spines along its length. Plesiomorphically, the midfemur has spines, as in roaches and in Santanmantis.
6.
10.
11.
NO. 3412
Mid and hindlegs are long and slender and are the only legs involved in walking, or at least appear structurally so. Plesiomorphi- cally, all three pairs of legs are involved in locomotion, and the mid- and hindlegs are not particularly longer or more slender than the fore pair.
. Claval furrow in the forewing is not
arched, at best it is slightly curved and of- ten straight. The plesiomorphic situation is found in most modern roaches (some, like Plectopterinae, have lost this feature), in Paleozoic roachoids, and some of the most basal, Cretaceous mantises (e.g., figs. 5, 6, 20, 22-24).
. At the base of the hindwing is a small but
fully formed crossvein, r-cu. Plesiomorph- ically this vein is absent. As would be ex- pected, this character is virtually impossi- ble to see in fossils. Only two Cretaceous fossils have the basal portions of the hindwings preserved (Cretophotina tris- triata and Baissomantis maculatus, fig. 6), but preservation of the r-cu crossvein in both is ambiguous or obscure.
. In the region near the middle of veins M
and Cu of the forewing is an oblique, thick- ened (and sometimes pigmented) structure, called the “‘pterostigma’’ by some authors, or ‘‘pseudovein’” (Nel and Roy, 1996) (figs. 4c, 5b, c; 6; 20c; 23, 24). This struc- ture is actually a thickened, sclerotized area of the wing membrane and not a vein. It is not homologous to the true pterostigma in insects, which is located at the apex of the radial vein near the wing margin and usu- ally has more discrete edges. In the Early Cretaceous Santanmantis from Brazil the pseudovein is very long, extending from the basal forks of M and Cu and through CuP and nearly to the margin of the anal lobe. Contrary to comments by Nel and Roy (1996), the pseudovein is not “‘spe- cial’? to Chaeteessa, but is present in var- ious forms throughout Mantodea. Plesio- morphically, this structure is absent. Since it is a sclerotized structure, its apparent ab- sence in compression-fossil wings is usu- ally not ambiguous.
Forewing with vein R simple or at best with 2 main branches. Plesiomorphically, the radial field is large, comprised of 3—4 main branches of R. Metallyticus is the only extant genus with 3 branches of R; presumably this is a reversal.
Pronotal shape is square or rectangular, with sides usually down turned (saddle- shaped). Plesiomorphically, the pronotum
2003
Figs. 25.
GRIMALDI: EARLY MANTISES 39
I SOF SE eS eee a 160mm"
500m
Scanning electron micrographs of tibial spur (a, c) and femoral brush (b, d) in a “‘lower”’
mantis (Amorphoscelis: a, b), and a “‘higher’’ mantis (Pseudocreobotra: c, d). Note the barely discern-
able suture between the spur and tibia in c (in detail).
12
13.
is discoid (rounded or nearly so), as occurs in most extant roaches and Paleozoic roachoids. Caution must be used in observ- ing this feature. The first specimen of San- tanmantis studied here (the holotype) had an apparently discoid pronotum, but sub- sequent study of additional specimens re- vealed the pronotum to be slightly to dis- tinctly quadrate, so the shape of the holo- type’s pronotum was probably slightly de- formed.
Forefemur with a patch of fine, short setae or spinules on inner surface, nearer the dis- tal end (the forefemoral ‘“‘brush’’) (fig. 25). Plesiomorphically, the brush is absent. Forebasitarsomere long, its length equal to or greater than (sometimes considerably so) the length of the foretibia (without the spur). Plesiomorphically, the length of this basitarsomere is 0.5—0.7X the foretibial length, as is found in Chaeteessa and some of the Cretaceous mantises, though some
14.
15.
16.
Is.
Mantoidea have secondarily evolved a re- duction of this segment.
Forefemur with discoidal spines present, usually 3—4 located on the ventral surface at the proximal end and between the mesal and lateral rows of spines. Plesiomorphi- cally, these are absent.
Forefemur with a lateral row of 4—5 spines and a mesal row of generally 8 or more similar spines. This is a fairly conservative arrangement in Mantodea, with the most notable exceptions being cases of extreme modification. Amorphoscelis, for example, has lost most of the forefemoral spines. Plesiomorphically, spines do not occur, with the femur armed merely with stiff se- tae, as in Burmantis, Chaeteessites, and Jersimantis.
Foretibia with a large apical, articulated spine, or a spur on the inner surface. Ple- siomorphically the spine or spur are absent. Foretibia with a distinctive, long, apical
40
18.
19:
20.
pay
22.
23:
24.
AMERICAN MUSEUM NOVITATES
spur (sometimes called the “‘claw’’) on the inner surface. This spur is typically heavily sclerotized, with its articulated base barely discernable (fig. 25c) and situated at the apex of an extension of the tibia that pro- jects well beyond the tibial-tarsal joint. Ple- siomorphically, the spur is absent, or there is a large, spinelike, articulated seta in this spot, sometimes also with a smaller one on the outside surface.
Hind wing with vein 1V (A1) lost (Smart, 1956). Plesiomorphically, it is present, as is found in the living genera Chaeteessa and Metallyticus.
All three ocelli are lost. Plesiomorphically, all three ocelli are present (as in most Man- todea or basal Neoptera), or at least the two lateral ocelli (as in roaches and most ter- mites). Care must be taken in assessing this feature in nymphal mantises, as ocelli are minute and often obscured.
Forewing with vein PCu incomplete (Smart, 1956), not extended to wing mar- gin. Plesiomorphically, it is complete, as is found in Mantoida, Metallyticus, and Bais- somantidae. Some Hymenopodidae and Mantidae have a complete CuP, but this is obviously a reversal of the groundplan state of an incomplete CuP found among virtually all living mantises.
Females with wings reduced or absent. Ple- siomorphically, the females have wings as well developed as in males.
A metathoracic hearing organ is present that is morphologically unique in insects (Yager and Hoy, 1986; Yager, 2000). The structure occurs ventrally, with the external opening being a slit between the hindcox- ae. The tympana of the hearing organ are comprised of a pair of drop-shaped areas of cuticle recessed into the groove, which oppose each other. Mantises are auditory cyclops and tone deaf, distinguishing nei- ther directionality nor frequency of sound. Plesiomorphically, the groove and tympana exist, but specialized tracheal sacs and sen- silla that magnify and transduce the sounds are absent (Yager, 2000).
Pronotum is elongate, with a length 2—20X its width. Plesiomorphically, the length is barely longer than the width.
Cerci are long, with at least 20 segments. Plesiomorphically there are 8—15 segments, as found in roaches and most other man- tises. Some Mantodea have significant re- duction of the cerci (e.g., Eremiaphila, no doubt an adaptation for the extreme con- ditions of its habitat); some Mantidae have
NO. 3412
more than 20, but this is clearly secondar- ily derived.
25. The setae in a patch or brush on the fore- femur are flattened and scalelike, which oc- curs in all living Mantodea. The plesiom- orphic state, where the setae are only slightly thickened, occurs in Burmantis asiatica (fig. 8a), observation of which is a result of unusually good preservation. In this fossil the brush is microscopically vis- ible, and the setae are barely flattened and least modified among all other mantises. Scanning electron microscopy of brush se- tae in various living mantises indicates that the setae are always flattened and scaliform (fig. 25b, d), or have a shape that is feath- erlike.
26. The forefemur has a row of 5 ventromesal spines and 3 long ventrolateral spines. This is a feature that occurs in Burmantis spe- cies. Plesiomorphically, the forefemur has just stiff, sharp setae (as in Jersimantis) or numerous spines arranged as in Mantoida, Chaeteessa, and Ambermantis.
CLADISTIC ANALYSIS
Table 3 is a matrix of 26 morphological characters for 20 living and fossil taxa (min- imum possible steps of 25, maximum possi- ble steps 80). Half of the cells (263 of 520), have missing entries, denoted by a ’’?’’. Only two cells of living species have missing en- tries. These involve Amorphoscelis and refer to the spination of the forelegs, which is ex- tremely reduced in this genus. As expected, virtually all missing entries involve fossil taxa, but there is a very uneven distribution of these among the fossils. Rock fossils av- eraged 17 missing entries (range of 10—20), with the fewest being in the completely pre- served Santanmantis. Amber fossils aver- aged 10 missing entries (4—14), with the few- est in Ambermantis, preserved in entirety as an adult.
Cladistic analysis used the phylogenetic program WINONA, version 2.0 (Goloboff, 1999), run with a PC having a 256-MHz pro- cessor and 40-GB memory. Also used was PAUP version 4 (Swofford, 2002), run on a MacG4 computer with dual processors. Hav- ing half of the matrix with missing entries was computationally intensive and signifi- cantly complicated the analyses. Analyses in WINONA, for example, yielded more than
2003
GRIMALDI: EARLY MANTISES
41
TABLE 3 Matrix of Taxa and Characters for Cladistic Analysis
@
N RR Wk f bP O1 ~] CO Fr NR NO Ot NO ON
Santanmantis axelrodi Baissomantis spp. Cretophotina spp. Jersimantis burmiticus Jersimantis luzzii Chaeteessites minutissi Burmantis asiatica Burmantis lebanensis Cretomantis larvalis Electromantis sukatshev Ambermantis wozniaki Arverineura insignis Chaeteessa spp. Mantoida spp. Amorphoscelis spp. Metallyticus sp. Eremiaphila spp. Empusidae spp. Hymenopodidae Mantidae
PRPPRPRPPrEP Ew PPR EE PPE Pw wR] be PPP PRP PP PPP RPP PPP PE wy wa Plo PPP PPP PrRPw PrP vy PPP RP Pw ve Pl PPP PPR PPP Pw ww vwvrw PHO] eB PPP PPPP Pw PRPOREP WRF ww oly PRPRPPPRP Pw Pw PRP Pw PP wv Plan PRPRPPPRPEP PPR www vw vw OOo] Aa PPPPRPRPP RP www vw vow vo wv vw wv
10,000 most parsimonious (MP) trees, with 35 steps. The strict consensus tree was com- pletely unresolved. PAUP analyses were run with the characters ordered and unordered, in each case using 10 replicate analyses with each one having the input of matrix data ran- domized. Searches for MP trees were termi- nated after 49,600 trees were found (memory could store no more). The strict consensus of the MP trees was completely unresolved, no doubt a result of the many missing charac- ters, but still very unusual given that signif- icant phylogenetic structure exists for the liv- ing families that does not conflict with other parts of the phylogeny.
Two majority-rule (MR) trees were ob- tained, one for ordered characters and the other for unordered characters, both with sig- nificantly resolved topology (fig. 26). Both MR trees yielded poorly supported group- ings, such as the following:
1. Santanmantis + Cretomantis (the former a complete adult, the latter a nymph, with very few shared features);
2. Baissomantis as part of a basal pentatomy, even though its wings lack a distinctive syn-
PRPPPEP PEPE PPP aww wv vv Por lw
PRPRPPORPPPP Pay ww ww vn OoOolor PRPPPPP EP Py PPP P RP PEP yw Pl PPR PPP PPP PP wy Pw wv PRP wy OO Ww ww PRPRPPPRPPROWPPOWrF OOO w ww PPRRPRPwWrPRPwW ODD DCO COW vw PRPPRFPPwrFP Pw PPRPODOOO Ow ww PPRPPRPREP RRP wa PPP wi PRP PE vo Pl oaPRe PRPPrPPrPPRPOWrF COON DOC OW ww PRPPPRPOPPHFP ON WN ww vv vw vv Rw TODO COD CON ONnwvnvwwPrRPwwolorR PRPPPOFORP Pw wwwwww vw OorRPlon PRrPPPORODON WN Vw wv vvww wv OoOlrFn PRPrFRODOOON Nv wv vw vv wvvwy PPRODOCONDODO ODOC COON WOlWwn PDODDDADOFRFWNEFH WN OOOWNOOWNWO!]AN PPP RPPrPEFP Pw Powwow oow ww TODO COCO ON ONOrFPKHPOOW ww
apomorphy (character 9, the pseudovein) common to all other winged mantises;
3. Ambermantis as part of a basal polytomy of living families, even though Ambermantis lacks a derived venational feature of living families (character 10) and distinctive fea- tures of foreleg spination found in the Chae- teessidae, or in all other mantises (character 14, discoidal spines).
Thus, significant aspects of the MR trees were not compelling. An alternative, pre- ferred cladogram was constructed by hand, some portions of which are in agreement with the MR trees, whereas other portions differ considerably (fig. 27). This preferred tree was based on some inference regarding the probability of the existence of synapo- morphies not directly observed in some fos- sils. The more inclusive the synapomorphy, and the more exclusive (i.e., the more re- cently derived) the taxon, the higher the probability that the taxon possessed the syn- apomorphy (or secondarily lost the feature). For example, though Cretophotina is known only as wings, it is a reasonable hypothesis that this genus possessed raptorial forelegs
NO. 3412
AMERICAN MUSEUM NOVITATES
42
SNWISSHNUIW SONSSAAJOCYY
sisuauege] SIJUBULING eoneise syueuung
IZZN| syuewisief SNOIVIULING SnueWUISIEf sijuewossieg
snoijAl|e}a\ -
eplojuey| sepiiudeiweig sepljeosoyudiowy SeEpue|| eepipodouswApy eepisndwy BINOULIBAIY esseajeeuyD SIJUBLUISQUIYY SIJUBWOIJOS]A eunoydojaiy SIJUBLUOJOID) suewuewues dnoi6}no
SNWISSI]NUIL SOJSSOQJOLYD
sisuguega] sijueULNg eoneise snueuuing
zznj snuewisies SnoiiuUng snuewuisief sijuewossieg
Sepiue|| eepipodouswAH eepisndwy sepi|iydelwiasy Sepljeosoydiowy snoizA|e}a\|
66
UNORDERED
eplojuey\ esseajeeyy BINSULOAIY SIJUBLUJEqUYY SIJUBLUOJ}N9/9 euljoydojaiy SIJUBLUOJOID sljuewUeUeS dnou6\no
ORDERED
0.81) of Cretaceous genera and living families of Mantodea, based on the matrix in table 3. Numbers
Fig. 26. Majority-rule cladograms (with characters ordered and unordered) (L = 36, CI = 0.72; RI at nodes are bootstrap values.
2003 GRIMALDI: EARLY MANTISES 43 S % 2 2 = o = ii] o n & = my S) 22 £ o» «4 2 2 2 g & = z = = og 2 FS 4H FE = ” oO A, cM FR =“ 5s © S$ € & o i = 2 G = = @ a2) 2 oc c So wn eo © 69 £ - a oo EG af ee FE SEEZCES BEE SH ES SEEZse SBS SER Ss FS SSFEEESEE 2eepS ER Eee SER ESE SE EES i] = = = — oD geese to@aananwoag Ge SO SSadwt w rs ?+— MANTOIDEA 44 I 23 a 22 19 26 13 21 15? ~-ae 15 7 20 12.7 efeeeeenLenee fennel a2? 14 <¢—— EUMANTODEA 25 a ee sesso Ah 17 <—— NEOMANTODEA 5 15 9 12 % 16 st 11 > 3 <@— MANTODEA . -a Fig. 27. Preferred cladogram for basal relationships of the Mantodea. Ambiguities in character dis-
tribution are indicated by shading. Further work will focus on Eumantodea. See text for discussion.
(character 2) since more basal genera pos- sessed raptorial forelegs. In other words, Cretophina possessesed certain derived, more exclusive features (i1.e., reduction of claval furrow, presence of pseudovein, char- acters 4 and 9, respectively) highly correlated with having raptorial forelegs. Likewise, it is a reasonable hypothesis that Cretophotina had a forefemoral brush, as all but the three most basal genera possess this feature. Con- versely, given that the mantis ear (character 22) is present in the Mantoidea but not in basal living families, it is a reasonable hy- pothesis that the ear did not occur in Creta- ceous mantises since they are even more bas- al on the basis of independent evidence.
Assumptions like these are not accommodat- ed in parsimony or majority-rule analyses, but they may be warranted in that they result
in a testable hypothesis, but more important- ly one that appears predictive. Certainly, the strict consensus tree, an unresolved bush, is erroneous. Strictly interpreted, the consensus trees depict a simultaneous origin of all man- tis groups, even though 135 million years of continuous evolution separates the oldest ge- nus from recent ones. Disciples of strict con- sensus methods could, alternatively, argue that the data prevent a more resolved hy- pothesis, but this is tautological and is even contradicted by the high bootstrap values in some clades of the MR trees. As new fossils are gradually discovered, or more completely preserved specimens of described species, there will be significant revision to the pre- ferred phylogenetic tree. Some phylogenetic structure is required for interpretation of the Cretaceous fossils, so the one hypothesis is
+t AMERICAN MUSEUM NOVITATES
offered here simply as the best available working hypothesis.
One potentially problematic aspect of this analysis is the different criteria used in the taxonomy of compression fossil and amber fossil specimens. Only body structures were available for the nymphs in amber, and only venation for the wings preserved in rocks. Thus, it is possible that there may be some synonymy between a few of the taxa in rocks from the Cretaceous of Eurasia (Baissoman- tis and Cretophotina) and ones known as nymphs in amber (Burmantis, Chaeteessites, Cretomantis, Jersimantis), a possibility that could only be resolved with the discovery of completely preserved adults. It is doubtful, though, that there is significant synonymy here, but any such synonymy would have lit- tle effect on the consistent occurrence of bas- al most mantises in the Cretaceous (see be- low). Fortunately, too, some stability to the systematics of Cretaceous Mantodea is pro- vided by exquisite preservation of complete, adult Santanmantis and Ambermanitis.
Despite ambiguities with the phylogenet- ics, new evidence does not support the clas- sification of Cretaceous mantises proposed by Gratshev and Zherikhin (1993):
Family Chaeteessidae: genus Cretophotina Kazakhaphotina Vitimiphotina Chaeteessites (tentatively) (+ Arverineura, Chaeteessa, Lithophotina, Megaphotina: Tertiary/Recent). Family Baissomantidae: Baissomantis Family Cretomantidae: Cretomantis Electromantis Family Amorphoscelidae: Amorphoscelites (+ extant genera)
The following revised classification is pro- posed, in which Baissomantis (Baissomanti- dae) is considered a sister group to the true mantises, order Mantodea:
ORDER MANTODEA: Pronotum quadrate, sad- dle-shaped, not covering head; forelegs spined, raptorial, foretibia with large apical spine or spur, with freely moving forecoxae; mid- and hindlegs long, slender, used in walking; forewing with pseudovein.
NO. 3412
Family Santanmantidae, new family: as defined for type genus. Genus Santanmantis Genera Incertae Sedis: Amorphoscelites, Bur- mantis, Chaeteessites, Cretophotina, Electro- mantis, Jersimantis, Kazakhaphotina, Vitimi- Pphotina SUBORDER NEOMANTODEA (new): Midfemur lack- ing spines; claval furrow straight or only slightly arched; foretibia with long apical spur; forefemur with patch of small scales on mesal surface. Family Ambermantidae, new family: as defined for type genus. Genus Ambermantis INFRAORDER EUMANTODEA (new): All living man- tises, which have forefemur with discoidal spines. Families: Chaeteessidae, Mantoididae, Metal- lyticidae, Amorphoscelidae, Eremiaphilidae, and superfamily Mantoidea.
The two new families are basal to living families but they also possess unique apo- morphies. Baissomantidae is still defined on the basis of plesiomorphic features and, though related to mantises on the basis of wing venational characters, it is not a mantis in the sense defined here. One could propose families for each or most of the Cretaceous genera, but given that characters preserved in the various fossils are not all comparable, it would be imprudent to formally propose any new higher taxonomic names. The two new families described are for taxa that are preserved as complete adults.
Gorochov (2001: 357) recently comment- ed that “it cannot be excluded that Ponop- terix and possibly Umenocoleus are represen- tatives of a second-oldest branch of Mantina [Mantodea], having a shape of head and pronotum as well as almost raptorial fore- legs, similar to mantises.’’ These two genera are roaches of the Cretaceous family Umen- ocoleidae. Neither Ponopterix nor Umeno- coleus are known to have raptorial forelegs, either “‘almost’’ or actually. Also, the very highly modified, tegminous forewings and hindwing venation, the broad abdomen, short mid- and hindlegs, and other features pre- clude their close relationship to mantises.
CONCLUSIONS
There appears to be no relationship be- tween age and cladistic rank among Creta- ceous Mantodea. This can be due to prob- lems in the phylogenetic hypothesis, to in-
2003
adequate taxon and character sampling, or to both. It is assumed that, given sufficient stratigraphic sampling and accuracy of phy- logenetic hypotheses, such a correlation would appear. What is highly significant, though, is the fact that Cretaceous mantises are consistently basal phylogenetically to liv- ing families, the Eumantodea. The Creta- ceous mantises possess plesiomorphic fea- tures of venation, the forefemoral brush, forefemoral spines, and/or the foretibial spur. Clearly, mantises are in nascent stages of their evolution in the Cretaceous. True man- tises, complete with raptorial forelegs and other diagnostic features like the pseudovein, probably appeared in the Late Jurassic and almost certainly no earlier. In this regard, Zherikhin’s (2002) view on the relatively re- cent age of Mantodea is accurate, and esti- mates of Paleozoic mantises are extremely excessive.
Independent evidence suggests that the su- perfamily Mantoidea is actually quite young, perhaps even entirely Tertiary in age. The group is defined by, among other features, the distinctive cyclopean ear (Yager, 2000), which is sensitive to sounds at 25—50 Hz, or the region of bat echolocations. Stereotyped evasive behavior of flying mantises to bat calls indicate that the ear is probably an ad- aptation for avoidance of bats during night- time flight (Yager, 2000). Thus, Mantoidea is a group probably not much older than the microchiropteran (insectivorous) bats, which appear suddenly in Lower Eocene strata of Europe, North America, Australia, and Af- rica (reviewed in Simmons and _ Geisler, 1998) and further diversified later in the Eo- cene. Basal relationships of the earliest mi- crochiropteran fossils (Simmons and Geisler, 1998) suggest a Paleocene origin of these mammals, and almost certainly no older. The oldest mantoidean is Prochaerododis enig- maticus, from the Paleocene of France (Nel and Roy, 1996), which is based on a portion of a rather distinctive forewing.
Though the Cretaceous mantises have not helped to reveal a close blattodean relative, the revised fossil record provided here, I think, helps clarify the origins and earliest radiation of the mantises. Cretaceous mantis- es were probably superficially very similar to most species of the basal families Chaetees-
GRIMALDI: EARLY MANTISES 45
sidae, Metallyticidae, and Mantoididae: small, brown, stout-bodied predators, cryptic and scuttling among leaves on the forest floor or on tree trunks. Origins and radiations of the superfamily Mantoidea in the Early Ter- tiary led to the array of spectacular mantises found today.
ACKNOWLEDGMENTS
I am indebted to many people who con- tributed directly and indirectly to this project. First, Dr. Herbert Axelrod and Mr. Joseph Wozniak generously donated to the AMNH the wonderful specimens of Santanmantis and Ambermantis, respectively. Mr. Robert Goelet, trustee and Chairman Emeritus of the AMNH, provided funds for the acquisition of Burmese and Lebanese amber fossils. Dr. Giinther Bechly (Staatliches Museum fiir Na- turkunde, Stuttgart) kindly provided excel- lent photographs of SMNS 113 and 172 (te- produced here) and checked on several char- acters in these specimens (currently on ex- hibit and not loanable). He also provided a loan of the other three Santana fossil mantis- es from their collection. Dr. Alexandr Ras- nitsyn (Arthropod Laboratory of the Pale- ontological Institute, Russian Academy of Sciences, Moscow) was an extremely helpful and gracious host during my visit in August 2002, and he very kindly loaned important specimens. I relied on the electron micro- scope and photomicrographic skills of Mr. Tam Nguyen, Senior Scientific Assistant at the AMNH. Biolmaging Resources (Lincol- nshire, IL) generously donated HRCT scans of AMNH 1957, files from which Ms. An- gela Klaus (Director of the AMNH Interde- partmental Laboratories) reconstructed for imaging. Dr. Valerie Schawaroch ran the ma- trix in PAUP on her MacGé4 and provided spirited discussion on the interpretation of re- sults. Lastly, Mr. Paul Nascimbene (AMNH Curatorial Specialist) prepared the amber specimens with his usual finesse. A U.S. Na- tional Science Foundation grant (DBI- 9987372) generously supported curation of the collection of fossil insects at the AMNH, and some aspects of research on the collec- tion. Commentaries provided by Drs. Klaus Klass, Michael Engel, and Erich Tilgner greatly improved the manuscript.
46 AMERICAN MUSEUM NOVITATES
REFERENCES
Azar, D. 2000. Les Ambres Mésozoiques du Li- ban. Doctoral diss., Université de Paris-Sud. Bechly, G. 2001. Die faszinierende Evolution der Insekten. Stuttgarter Beitrage zur Naturkunde
(C) 49: 1-96.
Beier, M. 1968. Mantodea (Fangheuschrecken). Handbuch der Zoologie, vol. IV. Band: Ar- thropoda, 2 Halfte: Insecta. Zweite Auflage: 1— 47.
Boudreaux, H.B. 1979. Arthropod phylogeny, with special reference to the insects. New York: Wiley.
Carpenter, FM. 1992. Superclass Hexapoda. Part R, Arthropoda 4, volumes 3 and 4. Treatise on Invertebrate Paleontology. Boulder, CO: Geo- logical Society of America.
Cruikshank, D., and K. Ko. 2002. Geology of an amber locality in the Hukawng Valley, northern Myanmar. Journal of Asian Earth Sciences 21: 441-455.
Goloboff, P. 1999. Winona, version 2.0, Program and documentation. Tucuman, Argentina.
Gorochov, A.V. 2001. The most interesting finds of orthopteroid insects at the end of the 20% century and a new recent genus and species. Journal of Orthoptera Research 10: 353-367.
Grandcolas, P. 1994. Phylogenetic systematics of the subfamily Polyphaginae, with the assign- ment of Cryptocercus Scudder, 1862 to this tax- on (Blattaria, Blaberoidea, Polyphagoidea). Systematic Entomology 19: 145-158.
Grandcolas, P. 1996. The phylogeny of cockroach families: a cladistic appraisal of morpho-ana- tomical data. Canadian Journal of Zoology 74: 508-527.
Grandcolas, P. 1999. Reconstructing the past of Cryptocercus (Blattaria: Polyphagidae): phylo- genetic histories and stories. Annals of the En- tomological Society of America 92: 303-307.
Grandcolas, P., and C. D’Haese. 2001. The phy- logeny of cockroach families: is the current molecular hypothesis robust? Cladistics 17: 48— 55%
Gratshev, V.G., and V. Zherikhin. 1993. New fos- sil mantids (Insecta, Mantida [sic]). Paleonto- logical Journal 27: 148-165.
Grimaldi, D. 1993. The care and study of fossil- iferous amber. Curator 36: 31-49.
Grimaldi, D. 1996. Amber, window to the past. New York: Abrams/American Museum of Nat- ural History, 215 pp.
Grimaldi, D. 1997. A fossil mantis (Insecta: Man- todea) in Cretaceous amber of New Jersey, with comments on the early history of the Dictyop- tera. American Museum Novitates 3204: 1-11.
Grimaldi, D., M.S. Engel, and P. Nascimbene.
NO. 3412
2002. Fossiliferous Cretaceous amber from Burma (Myanmar): its rediscovery, biotic di- versity, and paleontological significance. Amer- ican Museum Novitates 3361: 1-71.
Grimaldi, D., and J. Maisey. 1990. Introduction. In D. Grimaldi (editor), Insects from the San- tana Formation, Lower Cretaceous, of Brazil. Bulletin of the American Museum of Natural History 195: 5-14.
Grimaldi, D., T. Nguyen, and R. Ketcham. 2000a. Ultra-high-resolution X-Ray computed tomog- raphy (UHR CT) and the study of fossils in amber. Jn D. Grimaldi (editor), Studies on fos- sils in amber, with particular reference to the Cretaceous of New Jersey: 77-91. Leiden: Backhuys.
Grimaldi, D., A. Shedrinsky, and T. Wampler. 2000b. A remarkable deposit of fossiliferous amber from the Upper Cretaceous (Turonian) of New Jersey. Jn D. Grimaldi (editor), Studies on fossils in amber, with particular reference to the Cretaceous of New Jersey: 1-76. Leiden: Back- huys.
Hennig, W. 1981. Insect phylogeny. Translated and edited by Adrian C. Pont, with revisionary notes by Dieter Schlee and others. New York: John Wiley, 514 pp.
Kambhampati, S. 1996. Phylogenetic relation- ships among cockroach families inferred from mitochondrial 12S rRNA gene sequence. Sys- tematic Entomology 21: 89-98.
Klass, K-D. 1997. The external male genitalia and the phylogeny of Blattaria and Mantodea. Bon- ner Zoologische Monographien 42: 1-341.
Klass, K-D. 1998a. The ovipositor of Dictyoptera (Insecta): homology and ground-plan of the main elements. Zoologischer Anzeiger 236: 69-101.
Klass, K-D. 1998b. The proventriculus of the Di- condylia, with comments on evolution and phy- logeny in Dictyoptera and Odonata (Insecta). Zoologischer Anzeiger 237: 15—42.
Klass, K-D. 2001. Morphological evidence on blattarian phylogeny: “‘phylogenetic histories and stories’? (Insecta, Dictyoptera). Deutsche Entomologische Zeitschrift 48: 223-265.
Krassilov, V.A., and A.P. Rasnitsyn. 1999. Plant remains from the guts of fossil insects: evolu- tionary and paleoecological inferences. Pro- ceedings of the First Palaeoentomological Con- ference, Moscow 1998: 65-72. Bratislava: AMBA Projects International.
Kristensen, N.P. 1975. The phylogeny of hexapod “orders.” A critical review of recent accounts. Zeitschrift fiir Zoologische Systematik und Ev- olutionforchung 13: 1—44.
Kristensen, N.P. 1991. Phylogeny of extant hexa- pods. Jn CSIRO (editor), The Insects of Aus-
2003
tralia, 2 ed.: 125-140. Ithaca, NY: Cornell University Press.
Kukalova-Peck, J. 1991. Fossil history and the evolution of hexapod structures. In CSIRO (ed- itor), The Insects of Australia, 2" ed.: 141-179. Ithaca, NY: Cornell University Press.
LaGreca, M. 1954. Sulla struttura morfologica dell’apparato copulatore dei Mantodei. Annali dell’ Istituto Superiore di Scienze e Lettere di Santa Chiara, Napoli 1953/ 4: 1-28.
Lo, N., G. Tokuda, H. Watanabe, H. Rose, M. Slaytor, K. Maekawa, C. Bandi, H. Noda. 2000. Evidence from multiple gene sequences indi- cates that termites evolved from wood-feeding cockroaches. Current Biology 10: 801-804.
Maisey J. (editor) 1991. Santana fossils: an illus- trated atlas. Neptune City, NJ: TRH Publica- tions, 459 pp.
Martill, D. 1988. Preservation of fish in the Cre- taceous Santana Formation of Brazil. Palaeon- tology 31: 1-18.
Nascimbene, P, and H. Silverstein. 2000. The preparation of fragile Cretaceous ambers for conservation and study of organismal inclu- sions. In D. Grimaldi (editor), Studies on fossils in amber, with particular reference to the Cre- taceous of New Jersey: 93-102. Leiden: Back- huys.
Nel, A., and R. Roy. 1996. Revision of the fossil “‘mantid’’? and ‘‘ephemerid’”’ species described by Piton from the Palaeocene of Menat (France)(Mantodea: Chaeteessidae, Mantidae; Ensifera: Tettigonioidea). European Journal of Entomology 93: 223-234.
Rasnitsyn, A.P., and A. Ross. 2000. A preliminary list of arthropod families present in the Bur- mese amber collection at The Natural History Museum, London. Bulletin of the Natural His- tory Museum, London (Geology) 56: 21—24.
Schlee, D., and H-D. Dietrich. 1970. Insekten- fiihrender Bernstein aus der Unterkreide des Lebanon. Neues Jahrbuch fiir Geologie und Pa- laontologie Monatshefte 1970: 40—50.
Sharov, A.G. 1962. Redescription of Lithophotina floccosa Cock. (Manteodea) with some notes on the manteod wing venation. Psyche 69: 102— 106.
Simmons, N.B., and J.A. Geisler. 1998. Phyloge- netic relationships of I/caronycteris, Archaeo- nycteris, Hassianycteris, and Palaeochiropte- ryx to extant bat lineages, with comments on the evolution of echolocation and foraging
GRIMALDI: EARLY MANTISES 47
strategies in Microchiroptera. Bulletin of the American Museum of Natural History 235: 1— 182.
Smart, J. 1956. On the wing-venation of Chae- teessa and other mantids (Insecta: Mantodea). Proceedings of the Zoological Society of Lon- don 127: 545-553.
Smith, A.B. 1994. Systematics and the fossil rec- ord: documenting evolutionary patterns. Lon- don: Blackwell.
Snodgrass, R.E. 1935. Principles of insect mor- phology. New York: McGraw-Hill, 667 pp. Swofford, D. 2002. PAUP, Phylogenetic Analysis Using Parsimony, version 4 Sunderland, MA:
Sinauer Associates.
Thorne, B.L., and J.M. Carpenter. 1992. Phylog- eny of the Dictyoptera. Systematic Entomology 17: 253-268.
Thorne, B.L., D. Grimaldi, and K. Krishna. 2000. Early fossil history of the termites. Jn T. Abe, M. Higashi, and D.E. Bignell (editors), Ter- mites: Evolution, sociality, symbioses, ecology: 77-93. Dordrecht: Kluwer Academic Publish- ing.
Vrsansky, P. 2002. Origin and early evolution of mantises. AMBA Projekty 6(1): 16 pp.
Vrsansky, P, V.N. Vishniakova, and A.P. Rasnit- syn. 2002. Order Blattida Latreille, 1810. The Cockroaches (= Blattodea Brunner von Wat- tenvill, 1882). In A.P. Rasnitsyn and D.L.J. Quicke (editors), History of insects: 263-270. Dordrecht: Kluwer Academic Publishers.
Wheeler, W.C., M. Whiting, Q.D. Wheeler, and J.M. Carpenter. 2001. The phylogeny of extant hexapod orders. Cladistics 17: 113-169.
Yager, DD. 2000. Hearing. In E R. Prete, H. Wells, P. H. Wells, and L. E. Hurd (editors), The Praying Mantids: 93-113. Baltimore: Johns Hopkins University Press.
Yager, D.D., and R.R. Hoy. 1986. The cyclopean ear: a new sense for the praying mantis. Sci- ence 231: 727-729.
Zherikhin, V.V. 2002. Order Mantida Latreille, 1802. The Mantises (= Mantodea Burmeister, 1838). In A.P. Rasnitsyn and D.L.J. Quicke (ed- itors), History of insects: 273-276. Dordrecht: Kluwer Academic Publishers.
Zherikhin, V.V., and A.J. Ross. 2000. A review of the history, geology and age of the Burmese amber (Burmite). Bulletin of the Natural His- tory Museum, London (Geology) 56: 3-10.
Recent issues of the Novitates may be purchased from the Museum. Lists of back issues of the Novitates and Bulletin published during the last five years are available at World Wide Web site
http://library.amnh.org. Or address mail orders to: American Museum of Natural History Library, Central Park West at 79th St., New York, NY 10024. TEL: (212) 769-5545. FAX: (212) 769- 5009. E-MAIL: scipubs @amnh.org
This paper meets the requirements of ANSI/NISO Z39.48-1992 (Permanence of Paper).