GeolCarp_Vol65_No4_285

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GEOLOGICA CARPATHICA

, AUGUST 2014, 65, 4, 285—292 doi: 10.2478/geoca-2014-0019

Introduction

The order Mantodea comprises  ~ 2400 living species (Otte et
al. 2013) but its distribution and diversification patterns dur-
ing  the  Cretaceous  remain  poorly  known.  These  predatory
insects are mostly relatively large in size and possess rapto-
rial  forelegs  and  cursorial  mid  and  hind  legs,  and  exhibit
sexual cannibalism of males during mating (e.g. Beier 1968;
Grimaldi 2007). Predatory mantises probably evolved during
the Late Jurassic (Vršanský 2002, 2005b) and radiated in the
Early Cenozoic (Gratshev & Zherikhin 1993; Grimaldi 2003,
2007).  Mantodea  can  either  represent  the  sister  group  to
Blattaria + Isoptera (and Blattaria is paraphyletic in respect to
Isoptera)  (Grimaldi  2007)  or  they  independently  originated
from the cockroach family Liberiblattinidae (Vršanský 2002,
2004, 2010). The family Chaeteessidae belongs to the primi-
tive Mantodea and includes one recent and six fossil genera
(Zherikhin  2002,  2005b).  The  extinct  genus  Cretophotina
has  four  described  species  (Gratshev  &  Zherikhin  1993;
Vršanský 2002; Grimaldi 2003) and Chaeteessa has five re-
cent neotropical species plus one fossil species from the Oli-
gocene  to  Miocene  Dominican  amber  (Grimaldi  2003;
Rondón et al. 2007).

Two mantodeans were reported from the Lower Cretaceous

Crato Formation, including Santanmantis axelrodi Grimaldi,
2003 and one unnamed mantis (Grimaldi 2003, 2007; Hörnig
et al. 2013). The aim of this paper is to describe another new
mantodean species from the Lower Cretaceous Crato Forma-
tion and to discuss the implications of this finding for Creta-
ceous distribution patterns and phylogeny of mantodeans.

Material and methods

The specimen described here is deposited in the Staatliches

Museum für Naturkunde Stuttgart, Germany (SMNS 67583).

New Lower Cretaceous basal mantodean (Insecta) from the

Crato Formation (NE Brazil)

SHIH-WEI LEE

National Museum of Marine Science & Technology, No. 367 Pei-Ning Road, Zhongzheng District, Keelung 20248, Taiwan;

leesw@mail.nmmst.gov.tw

(Manuscript received December 23, 2013; accepted in revised form June 5, 2014)

Abstract: Mantodea are very rare in the fossil record. 28 fossil species are reported since the earliest occurrence of
mantodeans in the Upper Jurassic (Tithonian). Here, I describe Cretophotina santanensis n. sp. from the Aptian (Lower
Cretaceous) Crato Formation of Chapada do Araripe (northeastern Brazil). This species is characterized by long an-
tenna and primitive raptorial forelegs. Morphological characters shared with the living genus Chaeteessa would support
its assignment to the family Chaeteessidae. The tropical occurrence of the Early Cretaceous genus Cretophotina in
Gondwana, together with occurrences of the genus Chaetessa from subtropical and temperate zones of Laurasia, im-
plies that members of the family Chaeteessidae achieved nearly cosmopolitan distribution during the Early Cretaceous.

Key words: Fossil insects, Mantodea, Chaeteessidae, Cretophotina, Chaeteessa, Mesozoic, Aptian.

Fig. 1. Location of the Crato Formation (7°19’60” S and
39°50’00” W).

The drawings were made on the dry specimen using a camera
lucida attached to a Leica MZ125 stereo microscope and digi-
tized by scanning. Photographs were made with a Leica
DFC490 digital macro camera. The figures have been edited
with the Adobe Photoshop™ CS3 imaging software.

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The Crato Formation is exposed in the north-central part

of the Chapada do Araripe (Fig. 1), a large plateau in north-
eastern Brazil (Martill & Bechly 2007). The limestones from
this  location  contain  highly  abundant  plants,  invertebrates
and vertebrates (Menon & Martill 2007). All fossils occur in
laminated  limestones  of  the  Nova  Olinda  Member  of  the
Crato  Formation  that  correspond  to  the  Aptian  stage
( ~ 120 Myr)  (Martill  &  Heimhofer  2007).  Chapada  do
Araripe was located at 10° to 15° S during the Early Creta-
ceous and still within the tropics (Martill 2007). The deposi-
tional  environment  was  represented  by  a  lake  or  lagoon.
Hypotheses  range  between  freshwater  lake  and  brackish  to
hypersaline lagoon with varying depth and extension (Martill
& Bechly 2007). Among the fossils, insects dominate both in
abundance and taxonomic diversity. More than 300 species,
representing  over  20  insect  orders  have  been  described,
among which the terrestrial adult orthopterans, hemipterans,
ephemeropterans  and  odonates  (Menon  &  Martill  2007;
Bechly 2007; Heads & Leuzinger 2011).

The nomenclature for fore and hind wings is based on Com-

stock & Needham (1898), Kukalová-Peck (1991) and Nel &
Roy (1996). Abbreviations used: C – Costa, Sc – Subcosta,
R – Radius, Rs – Radius sector, M – Media, Cu – Cubitus
(A – anterior, P – posterior), A – Anal vein and the nomen-
clature used for cockroaches, RA + RP + MA would be R + RS.

Systematic paleontology

Order: Mantodea Burmeister, 1838

The traditional classification of mantises by Beier (1968)

recognizes eight families: the “basal” mantises with five fami-
lies  (Chaeteessidae,  Mantoididae,  Metallyticidae,  Amorpho-
scelididae  and  Eremiaphilidae);  and  the  superfamily
Mantoidea  with  three  families  (Empusidae,  Hymenopodidae
and Mantidae), that radiated during the Early Cenozoic (Beier
1968; Grimaldi 2003; Grimaldi & Engel 2005).

The EDNA fossil insect database (active December 15

2013) catalogues 28 fossil mantid species. Mantodeans prob-
ably  originated  from  the  cockroach  family  Liberiblattinidae
during  the  Middle  or  Late  Jurassic  (Vršanský  2002).  The
phylogenetic analyses indicate that mantises first diversified
during  the  Cretaceous,  and  explosively  diversified  (as  the
Mantoidea) in the Cenozoic (e.g. Grimaldi 2007). Vršanský
(2002)  described  Juramantidae  with  the  earliest  mantodean
Juramantis  initialis,  which  occur  also  in  diverse  amber  (P.
Vršanský, personal communication 2013). To date, the Crato
Formation has yielded two species of mantises. One is an un-
named specimen SMNS 66528 (Fig. 2) and the second spe-
cies  represents  the  primitive  Santanmantis  axelrodi
Grimaldi,  2003  (Bechly  et  al.  2001;  Grimaldi  2003,  2007;
Grimaldi  &  Engel  2005);  AMNH  1957  from  the  American
Museum of Natural History, New York, USA; SMNS 66519,
66677, 66678, 66679 and 66680 (Fig. 3).

Family: incertae sedis

Species: Santanmantis axelrodi Grimaldi, 2003

Fig. 3

The monotypic genus Santanmantis Grimaldi, 2003 in-

cludes S. axelrodi. According to Grimaldi (2003, 2007) and
Grimaldi  &  Engel  (2005),  it  is  a  relatively  small  mantis,
body  9.5—10.5 mm  long.  Habitus  similar  to  the  living  basal
mantises,  with  notably  short  abdomen  and  body,  pronotum
nearly  square.  Antenna  about  the  same  length  as  the  body.
Fore  and  hind  wings  extended  well  beyond  the  apex  of  the
abdomen.  Forewings  partially  sclerotized,  pseudovein  (a
very well developed claval suture) is larger than in any other
living  or  extinct  mantis.  The  primitive  raptorial  forelegs
were folded under the thorax, and the mid and hind legs were
long, gracile and with retained cursorial function.

Family: Chaeteessidae Handlirsch, 1920 sensu Gratshev &

Zherikhin, 1993

Type species: C. filata Burmeister, 1838 (Fig. 4)

Fig. 2. Unnamed mantodean in the Crato Formation. Fore- and hind
wings unknown; SMNS 66528.

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The family seems to be paraphyletic (Gratshev &

Zherikhin 1993; Grimaldi 1997).

D i a g n o s i s : (modified after Gratshev & Zherikhin 1993):

Small  or  medium-sized  mantids  with  cylindrical  body.  Head
freely movable, without excrescence; antennae long, filiform.
Prothorax  short,  without  lateral  expansion.  Forewings  fully
developed;  costa  field  narrow,  not  widened  basally;  interca-
lary  veins  present,  at  least  between  some  longitudinal  veins;
Sc usually  moderately  long,  multibranched;  R  almost  always
multibranched  apically;  RS indistinguishable;  M with  two  or
three branches, M5 indistinguishable among transverse veins;
CuA at least four-branched; clavus large, distinctly separated.
Hind wings fully developed, without colour patterns; R usu-
ally with short apical branches;   RS long, simple; M two or
three  branched,  M5  slightly  oblique,  transverse;  anterior
branch of A2 distinct; anal region moderately large. Fore fem-
ora  narrow,  with  two  ventral  rows  of  rigid  setae;  fore  tibiae
without terminal hook, with two rows of rigid setae, setae of
apical pair larger than others, symmetrical; tarsal articulation
between  them.  Mid  and  hind  legs  without  projections,  tarsi

five-segmented. Abdomen without projections; cerci long,
multisegmented, not widened apically.

C o m p o s i t i o n : Chaeteessa Burmeister, 1838 (Recent,

South  America);  Lithophotina  Cockerell,  1908  see  Sharov
(1962)  (Oligocene,  North  America);  Arvernineura  Piton,
1940 see Nel & Roy (1996) (Paleocene, West Europe); Cre-
tophotina  Gratshev  &  Zherikhin,  1993  (Cretaceous,  Asia);
Vitimophotina  Gratshev  &  Zherikhin,  1993  (Lower  Creta-
ceous,  Siberia);  Baissomantis  Gratshev  &  Zherikhin,  1993
(Lower  Cretaceous,  Siberia);  Kazakhophotina  Gratshev  &
Zherikhin, 1993 (Upper Cretaceous, Turonian);  Megaphotina
Gratshev  &  Zherikhin,  1993  (Oligocene,  Russia)  a  species
within uncertain genus for nymph in Upper Cretaceous am-
ber  from  Siberia  Chaeteessites  minutissimus  Gratshev  &
Zherikhin, 1993 and probably one amber specimen from Ja-
pan  (Upper  Cretaceous,  Northeast  Japan;  deposited  in  Kuji
Amber  Museum,  Japan;  brief  reported  by  National  Geo-
graphic News, April 25, 2008).

R e m a r k s : Diagnostic characters Chaeteessa filata, Cre-

tophotina tristriata, and C. santanensis n. sp. are shared with

the present specimen, supporting the
categorization of the genus Cretophotina
which is close to Chaeteessa within
Chaeteessidae.

Genus: Cretophotina Gratshev &

Zherikhin, 1993

Type Species: C. tristriata Gratshev &

Zherikhin, 1993

According to the previous studies

(Gratshev  &  Zherikhin  1993;  Vršanský
2002;  Grimaldi  2003),  the  genus  was
known exclusively on the basis of wings
from the Cretaceous of Asia. Character-
istics  as  follows:  costal  field  distinctly
wider  than  field  between  ScP  and  RA;
RP+MA  apically  with  five  to  eight  ter-
minal  branches;  MP  with  two  to  three
branches;  CuA  with  six  to  ten  terminal
branches;  CuP  branch  separated  from
main  stem  and  distinctly  curved.
Pseudovein is present.

C o m p o s i t i o n : C. tristriata Grat-

shev  &  Zherikhin,  1993  (Lower  Creta-
ceous, Siberia);  C. mongolica  Gratshev
&  Zherikhin,  1993  (Lower  Cretaceous,
Mongolia);  C.  serotina  Gratshev  &
Zherikhin, 1993 (Upper Cretaceous, Ka-
zakhstan) and C. selenginesis Vršanský,
2002 (Lower Cretaceous, Mongolia).

Fig. 5. 1—2 – Wings of Arvernineura insig-
nis Piton, 1940, France, Paleocene. 3 – Cha-
eteessa valida Perty, 1833, Living species,
female: A – left forewing, B – right
forewing (after Nel & Roy 1996).

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Fig. 6. Cretophotina santanensis n. sp., holotype; SMNS 67583.

Cretophotina santanensis n. sp.

Figs. 6—8

H o l o t y p e : SMNS 67583.
D e r i v a t i o n o m i n i s : Named after the city of Santana

do Cariri near the type locality.

S t r a t u m t y p i c u m : Lower Cretaceous, Upper Aptian,

Nova Olinda Member of Crato Formation.

L o c u s t y p i c u s : Chapada do Araripe, vicinity of Nova

Olinda, southern Céara, North-East Brazil.

D i a g n o s i s : Assigned to Cretophotina on the basis of

thorax that lacks elongation, with primitive raptorial forelegs
with  retained  cursorial  function,  without  Chaeteessa-similar
spines. C. santanensis differs from other members of the fami-
ly Chaeteessidae in having antenna of moderate width, elong-
ated  ~ 1½  time  as  long  as  forewing.  Pseudovein  moderate.
Forewing very similar to the Chaeteessa valida Perty, 1833
(see  Nel  &  Roy  1996)  (Fig. 5),  but  differs  in  RP + MA  and
CuA more branched. M (MP) extremely reduced.

Fig. 7. C. santanensis n. sp., holotype; SMNS 67583; Head and tho-
rax part.

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D e s c r i p t i o n : Antenna filiform, 31 mm long and with

more than 110 segments. Head 2 mm long and 4 mm wide is
moderately  wider  than  the  thorax  (5 mm  long  and  3 mm
wide) from dorsal view and with big compound eyes located
laterally.  Pronotum  slightly  longer  than  wide,  ending  in  the
anterior margin of thorax and seems a little bit shorter than
living  Chaeteessidae.  Raptorial  foreleg  without  preserved
spines,  coxa  robust,  8 mm  long,  femur  8.5 mm  tibia  8 mm
and tarsus segmented, about 5 mm long. Forewing elongated
and nearly ellipse, 23 mm long and 6 mm wide. Pseudovein
slightly  longer  than  other  Cretophotina  and  Chaeteessa.
Sc about ½ of whole wing length, subcostal area much nar-
rower than costal area. RA long and origin in about anterior
1/3  level  of  wing.  RP + MA  (fused)  with  eight  branches.
MP  extremely  reduced  and  with  three  branches.  CuA  with
six branches. It is possible, that the first branch of CuA fused
with  MP3  (discussed  below).  CuP  is  adjacent  and  parallel
with CuA6. Anal with at least three (possible four) branches.

Discussion

Although mantodeans originated in the Tithonian (Vršanský

2002), they were still rare during the earliest Cretaceous. For
example,  they  were  not  found  at  Yixian  (Liang  et  al.  2010)
and  at  Chernovskie  Kopi  (Barna  2014).  The  superfamily
Mantoidea  radiated  during  the  Early  Cenozoic,  leading  to  a
high diversity of extant species (e.g. Grimaldi & Engel 2005).
The living genus Chaeteessa with five extant and one extinct
species is restricted to the Southern America (Grimaldi 2003;
Rondón et al. 2007). In addition to the morphologically simi-
lar and phylogenetically related genus Cretophotina from the
tropical  Crato  Formation,  Cretophotina  occurred  in  Laurasia
in  warm  subtropical  and  warm  temperate  zones  of  Siberia,
Mongolia  and  Kazakhstan  (Zherikhin  et  al.  1999;  Vršanský
2002;  Grimaldi  2003).  The  geographical  distribution  of  the
family Chaeteessidae in the Early Cretaceous thus seems to be
widely cosmopolitan.

Cretophotina santanensis has a long antenna, longer than

1½ of whole wing length. This character is a unique symple-
siomorphy with cockroaches. The new species shows a typi-
cal mantid forewing and Chaeteessidae body, with thorax

short  and  not  as  slim  as  modern  mantises  (see  Figs. 6—7).
The  genus  Chaeteessa  has  a  unique  arrangement  of  thick,
dark foreleg spines, virtually forming a basket and primitively
lacks a feature found in all other living mantises (Grimaldi &
Engel  2005).  The  forelegs  of  C.  santanensis  are  primitive
raptorial, which means that they lack spines and retain their
cursorial function, so that they resemble the most basal Jan-
tarimantidae  and  Juramantidae.  Specimen  SMNS  67583
forewing shows no colouration so that it is similar to Asian
Cretophotina  with  very  little  colouration.  In  analogy  to
cockroaches,  the  partially  colourless  wings  could  mean
comparatively open habitat (Vršanský et al. 2009).

Colouration was extensive in the genus Santanmantis. In C.

santanensis  the  forewing  has  the  first  branch  of  CuA  fused
with  MP3,  which  is  designated  as  a  deformation.  This  new
species is thus also similar to most basal mantodeans with de-
formations,  including  the  earliest  mantodean  Juramantis  ini-
tialis (Vršanský 2005b). C. santanensis forewing is similar to
the forewing of Chaeteessa valida Perty, 1833 (Fig. 5.3), and
differs by RP+MA and CuA being more branched and by the
origin of CuP being much closer to CuA6 (in C. valida and C.
filata the origin of CuP is much deeper). C. santanensis differs
from  C.  tristriata  by  the  forewing  ScP  being  much  shorter
(shorter than ½ of wing length, while in C. tristriata ScP long-
er  than  2/3  of  whole  wing  length),  by  RP+MA  with  more
branches (up to eight branches, while in C. tristriata RP+MA
with  five  branches)  and  CuP  much  straight.  C.  santanensis
forewing differs from Chaeteessa filata Burmeister, 1838 and
Lithophotina floccosa Cockerell, 1908 by having the long RA.
The forewing of this specimen is larger than C. valida (about
17.3 mm long), C. filata (about 17 mm long) while length of
the  Cretophotina  selenginesis  Vršanský,  2002  (about  20 mm
long), is 23 mm long.

It is difficult to interpret the phylogenetic relationships of

C.  santanensis  (nevertheless,  no  plesiomorphic  characters
are  observed).  Ten  short  fossil  mantodean  specimens  were
found so far in the Crato formation, corresponding to an ex-
tremely  small  proportional  abundance  of  this  order  among
several dozens of thousand of fossil insect reported from the
Crato  Formation  (Bechly  2010).  This  rarity  contrasts  with
the  dominance  of  predatory  cockroaches  Raphidiomimidae
in the Jurassic (Vishniakova 1973; Liang et al. 2009, 2012)

Fig. 8. C. santanensis n. sp., holotype; SMNS 67583; Forewing.

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and Mutoviidae in the Upper Permian of Isady in north-west-
ern Russia (Vršanský & Aristov 2012). These mantodeans
and also predatory Eadiidae (Vršanský 2009) are missing in
the Crato Formation.

Conclusions

Cretophotina santanensis n. sp. is described in America

for the first time and has a close relationship to living Chae-
teessa in tropical Southern America. The new species shows
a basel mantodeans type with some symplesiomorphy to
cockroaches.

Acknowledgments:  I  am  thankful  to  Dr.  Günter  Bechly
(Stuttgart) and Prof. Dr. James Nebelsick (Tübingen) for the
patient instruction and general help. I thank Dr. Karin Wolf-
Schwenninger  (Stuttgart)  for  technical  support.  I  am  most
grateful to Dr. Peter Vršanský (Bratislava, Moscow) for his
help  and  critical  comments.  Finally  I  want  to  thank  the
Staatliches Museum für Naturkunde, Stuttgart, Germany for
generously  providing  a  working  space  and  allowing  me  to
access the studied material.

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