A revision of Cretaceous mantises and their relationships, including new taxa (Insecta, Dictyoptera, Mantodea)

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).