Neomantodea. Grimaldi (2003: 44 and fig. 27; Fig. 28) introduced the name Neomantodea for a taxon comprising all extant Mantodea including †Ambermantis from New Jersey Cretaceous amber as their sistergroup. He named the following diagnostic characters for this grouping:
“Midfemur lacking spines; claval furrow straight or only slightly arched; fore tibia with long apical spur; fore femur with patch of small scales on mesal surface” (Grimaldi 2003: 44).
However, the claval furrow is strongly arched at least in Metallyticus (e.g. Wieland 2008a:
fig. 14; Béthoux & Wieland 2009: figs. 1a, 2a, 6a; Fig. 336), but also in some “higher”
Manto-Blepharodes
Fig. 27: Phylogeny of Empusidae propo-sed by Roy (2004a: fig. 24). Arrow indi-cates Idolomorphini sensu Roy 2004a, see Tab. 1 for Idolomorphini sensu Ehrmann 2002 (see chapter 4.3.74).
State of knowledge - Further systematic hypotheses 41
dea, for instance in Deroplatys lobata (see Béthoux & Wieland 2009: fig. 12e, f). Furthermore, there are small but distinct articulated spines on the mid- and hind femora of Chaeteessa (see characters 83, 84; Figs. 277-284). Additionally, the current state of knowledge is that Chaetees-sa does not have a tibial spur (e.g. Beier 1968a: 4; Salazar 2005: 269; but see character 68 and discussion in chapter 4.6.5). Therefore possibly none of these characters represents an autapo-morphy supporting the Neomantodea hypothesis. The scale-like nature of the grooming setae may be a synapomorphic character. However, as fossil species were not included in any other phylogenetic study and are not included in the present analysis except for comparative reasons, this hypothesis cannot be examined in further detail.
Eumantodea. Grimaldi (2003: 44 and fig. 27; Fig. 28) named the taxon containing all extant mantodeans Eumantodea, for which he listed the following characteristics: “All living mantises, which have fore femur with discoidal spines”. Although the current state of knowledge is that Metallyticus does not exhibit discoidal spines, Grimaldi (2003: tab. 3) encoded the discoidal spines as present for Metallyticus. However, the results herein (characters 54, 55 and discussion in chapter 4.6.4) indicate the maintenance of at least one discoidal spine in adult Metallyticus.
The lack of discoidal spines was encoded as “missing” or “unknown” for the fossil taxa in Grimaldi (2003). However, several of the specimens are (possibly early instar) nymphs that may exhibit much more slender and fragile spines than the adults (see also chapter 4.6.7 for postembryonic changes in the foreleg morphology).
The term Eumantodea was adopted by Svenson & Whiting (2009; Fig. 4A) and is certainly suitable to distinguish the extant Mantodea from their stemgroup ancestors.
Mantoidea. Mantoidea has initially been used as a synonym for Mantodea, for instance by Handlirsch (1903a, b). Vickery & Kevan (1983: 218, 219) introduced the superfamily Mantoi-dea comprising all MantoMantoi-dea except for Chaeteessa and Metallyticus.
Roy (1999: 39) used Mantoidea in a more restricted sense which has been adopted by sub-sequent workers, encompassing the traditional Acanthopidae, Hymenopodidae, Tarachodidae, Liturgusidae, Mantidae, Toxoderidae, Thespidae, Iridopterygidae, Sibyllidae, and Empusidae of the current classification sensu Ehrmann (2002: 372 ff.). Grimaldi (2003: fig. 27; character labels 22 and 23 addressed incorrectly, corrected in Grimaldi & Engel 2005: fig. 7.60; Fig. 28 herein) and Grimaldi & Engel (2005: fig. 7.60), used Beier’s (1968a) classification as a working hypothesis. They listed several putative autapomorphies for this grouping. Among them are the metathoracic hearing organ (“cyclopean ear”; see chapter 4.1.2.: introduction of “metathoracic hearing organ” and characters 130 and 131 for further detail; Yager & Svenson 2008) and the prothorax being “long” (Grimaldi & Engel 2005: tab. 7.3) or “2-20 times as long as wide”
(Grimaldi 2003: 40). However, as established by Yager & Svenson (2008: fig. 6), there are plenty of mainly Neotropical genera currently assigned to Mantoidea that exhibit the non-au-ditory MSMT-morphology of the metathoracic hearing organ (see Yager & Svenson 2008 and
“metathoracic hearing organ” in chapter 4.1.2), i.e. that are (primarily) deaf (Yager & Svenson
State of knowledge - Further systematic hypotheses 42
2008: 557). This would contradict the assumed single origin of tympanal hearing in Mantodea as found by Yager & Svenson (2008) and Sven-son & Whiting (2009).
As far as the second potential apomor-phy for Mantoidea is concerned, Svenson &
Whiting (2004a, 2009) and Yager & Svenson (2008) found support for a secondarily short-ened prothorax in several groups, among them Paraoxypilinae, Amorphoscelinae and Ere-miaphilidae. As it has been generally assumed that short prothoraces in Mantodea represent the plesiomorphic state, this further contradicts the Mantoidea-hypothesis. Therefore, the two autapomorphies suggested by Grimaldi (2003) and Grimaldi & Engel (2005) are doubtful (see also character 28).
Ware et al. (2008: figs. 2, 3) did not find sup-port for Mantoidea in their analyses.
None of the other studies allowed an ap-proach to the putative monophyly of Mantoi-dea, mostly due to a small taxon sample (e.g.
Klass 1995, 1997; Klass & Meier 2006), due to biogeographical restriction of the taxon sample (Jantsch 1999) or the lack of a subsequent phylogenetic analysis of the generated morphological data (Wieland 2006).
Mantomorpha. Klass (1995: 186; Fig. 1A) named the group containing all extant Mantodea except for Mantoida Mantomorpha. He listed 11 apomorphic morphological characters of mem-branous and sclerotized structures of the male genital apparatus for the group (Klass 1995: 187, 189). This was supported in several morphological studies (Klass 1995, 1997; Klass & Meier 2006). A basal dichotomy between Mantoida and the remaining Mantodea was also found in sev-eral molecular studies (e.g. Svenson & Whiting 2004a; Inward et al. 2007; Lo et al. 2007; Ware et al. 2008; Yager & Svenson 2008). The molecular studies often did not include Chaeteessa and/
or Metallyticus or they did not have suitable tissue for gaining optimal DNA sequences (Yager &
Svenson 2008). Ware et al. (2008) included all three taxa and found Mantoida to be the sistergroup of all remaining Mantodea. Svenson & Whiting (2009) who were able to gain better sequences from the DNA of Chaeteessa found the latter to be the sistergroup of the remaining Mantodea.
Mantidea. Klass (1995: 186; Fig. 1A) named all extant Mantodea except for Mantoida and Chae-teessa Mantidea. Six apomorphic morphological characters were listed (Klass 1995: 187, 189).
† Baissomantis
Fig. 28: Phylogeny of Mantodea including fossils found by Grimaldi (2003: fig. 27). Taxon names of extant Mantodea refer to classification of Beier (1968a), see chapter 2.1.4 for details.
Neomantodea Eumantodea Mantoidea
State of knowledge - Palaeontological record 43
Artimantodea. Svenson & Whiting (2009: node 7; Fig. 4A) introduced the name Artimantodea for all extant Mantodea excluding Chaeteessa and Mantoida (and possibly Metallyticus). This name would be synonymous with Mantidea sensu Klass (1995: 186) if Metallyticus is included in Artimantodea (situation unclear, see discussion of Mantoida, Chaeteessa and Metallyticus in this chapter 2.1.4; Fig. 4A). Autapomorphic characters for Artimantodea (if Metallyticus falls outside of Artimantodea) have not been mentioned.
Cernomantodea. Svenson & Whiting (2009: node 48; Fig. 4A) recovered many of the phylo-genetic affinities found by Yager & Svenson (2008). One of the most important morphological traits is the metathoracic hearing organ (see also discussion of Hymenopodidae in this chapter, characters 130 and 131 in chapter 4.1.2 and the corresponding introduction). All Cernomanto-dea (“perceptive MantoCernomanto-dea”) have one or two ears on the meta- and mesothorax, respectively.
In conclusion it can be stated that the monophyly of many of the families and subfamilies of the traditional classification is not supported by the molecular and morphological studies. So far, the monophyly of Empusidae, Toxoderidae, Sibyllidae, Amorphoscelinae, Paraoxypilinae, Acanthopidae (Stenophylla not studied), including its subgroups Acontistinae and Acanthopi-nae, OxyothespiAcanthopi-nae, and PhyllotheliiAcanthopi-nae, has been corroborated. The basal split in Mantodea is currently still under discussion.