Two new genera and two new species of Mantophasmatodea (Insecta, Polyneoptera) from Namibia

Abstract Two new species and two new genera (Pachyphasma, Striatophasma) of Mantophasmatodea are described from Namibia. Pachyphasma brandbergense is endemic to the Brandberg massif; Striatophasma occupies an extensive area south of the region inhabited by Mantophasma. Phylogenetic analyses (see Predel et al. in press) suggest a sistergroup relationship of Striatophasma and the South African Austrophasmatidae.


Introduction
When Mantophasmatodea was described in 2002 (Klass et al. 2002), 88 years had passed since the last discovery of a new extant insect order (Grylloblattodea by Walker 1914). Hence the interest in this new taxon was tremendous. Within a few years studies on morphology, genetics, behaviour and peptides of Mantophasmatodea were published (e.g. Klass et al. 2003, Dallai et al. 2003, Predel et al. 2005, Beutel and Gorb 2008, Eberhard and Picker 2008. Simultaneously the number of described species as well as the geographic distribution increased: being originally described with one species from Namibia and one from southern Tanzania, a remarkable species diversity was found, particularly in South Africa (Picker et al. 2002, Klass et al. 2003, Eberhard and Picker 2008. Up to now a total of 16 extant species in ten genera have been described (Klass et al. 2002, Klass et al. 2003, Zompro et al. 2003, Zompro and Adis 2006, Eberhard et al. 2011. The phylogenetic relationships within Mantophasmatodea as well as the taxonomic status of several genera have been subject of debate (Klass et al. 2003, Zompro et al. 2003, Zompro 2005, Zompro and Adis 2006, Damgaard et al. 2008, Eberhard et al. 2011) However, the separation and morphological description of species is sometimes difficult especially when based on few specimens. In a separate manuscript (Predel et al. in press) peptide mass fingerprints in combination with peptide sequencing were used to test the intraordinal phylogeny of Southern African Mantophasmatodea. Figure 1 shows a simplified version of the cladogram retrieved by Predel et al. (in press). Their analyses resulted in two monophyletic lineages: one containing all South African Austrophasmatidae (Austrophasma Klass et al., 2003, Hemilobophasma Klass et al., 2003, Karoophasma Klass et al., 2003, Lobatophasma Klass et al., 2003, Namaquaphasma Klass et al., 2003and Viridiphasma Eberhard et al., 2011 and the Namibian genus Striatophasma gen. n. while the second clade contains all remaining Namibian species which are currently grouped in Mantophasmatidae. Within Mantophasmatidae Praedatophasma Zompro & Adis, 2002+ Tyrannophasma Zompro et al., 2003and Mantophasma Zompro et al., 2002 n. were retrieved as sistergroups. The present article describes these two new genera and species discovered by Predel et al. (in press Figure 1. Simplified tree (midpoint-rooted) from a Bayesian phylogenetic analysis of peptide hormone sequences from southern African Mantophasmatodea (adapted from Predel et al. in press).

Methods
The terminology for the head and thorax follows Seifert (1995). The terms for abdominal structures are those used by Klass et al. (2003). The coloration refers to living specimens. Species descriptions are based on a designated holotype but all available specimens were taken into account in order to assess the intraspecific variation.
The information for the specimens is given in a standard manner, i.e., locality, geographic coordinates, elevation, date of collection (month indicated in lower case Roman numerals), habitat information, collector, depository, and preparation. Female (♀) and male (♂) symbols indicate the sex.
The following measurements are a selection from the proposed standards of Zompro et al. (2003) [abbreviations used by Zompro et al. (2003) in parentheses]: total length (a), length of pronotum (b), width of pronotum (c), length of mesonotum (b), width of mesonotum (c), length of metanotum (b), width of metanotum (c), length of the head (d), width of the head (f ), head width over eyes (g), width between eyes (h), length of eye (i) and width of eye (j). Specimens were examined under a Zeiss Stemi SV11 with a calibrated ocular micrometer. A Leica MZ 12,5 with a camera lucida was used for line drawings.
A pair of each newly described species was dehydrated using increasing steps of ethanol up to 100%, dried at the critical point (Emitech K850 critical point dryer) and subsequently sputter-coated (Emitech K500). Scanning electron microscopy was performed on a Philips XL30 ESEM using a special specimen holder (Pohl 2010). Parts of each species are illustrated in the standard views of dorsal, lateral and ventral. The head is also in frontal view (frons being vertically) and the terminalia are in caudal view.
Peptide mass fingerprints and sequences were obtained by direct profiling of neuroendocrine tissues from single specimens using MALDI-TOF mass spectrometry as described in Predel et al. (in press.) The specimens referred below along with the abbreviations used in the text will be deposited in the following collections: NMNW -National Museum of Namibia, Windhoek, Namibia; ZFMK -Zoologisches Forschungsinstitut und Museum Alexander Koenig, Bonn, Germany; ZMUB -University of Bergen, Zoological Museum, Bergen, Norway.

Taxonomy
Striatophasma gen. n. urn:lsid:zoobank.org:act:8B1618EC-5FBD-4C51-9301-7E0076EA742D http://species-id.net/wiki/Striatophasma Description and diagnosis. Striatophasma gen. n. is placed as sistergroup to all remaining South African Austrophasmatidae sensu Klass et al. (2003) based on peptide hormone sequences (Predel et al. in press). Except for Austrophasma gansbaaiense and Viridiphasma clanwilliamense (Eberhard et al. 2011), it can be easily distinguished from the Austrophasmatidae sensu Klass et al. (2003) by its greenish colour. From all South African Austrophasmatidae, Striatophasma is separated by the lack of a butterfly-shaped dark median spot below the antennal base, genae that protrude from the compound eyes, the presence of spination in most body parts and its distribution in Namibia. From other Namibian mantophasmatodeans it can be distinguished by the presence of a brown dorso-median longitudinal stripe in males in combination with greenish body colouration. Asymmetric male genitalia and compound eyes that do not protrude from the genae distinguish Striatophasma from the East African Tanzaniophasma.
Type species. Striatophasma naukluftense sp. n. Other included species. None thus far. Etymology. The generic group name Striatophasma is a composition from the Latin word striatus meaning striped and the ending -phasma which is commonly used to term mantophasmatodeans. The gender is neuter.
Wings: completely absent. Abdomen: longer than thorax and head combined; bright green, meso-dorsal brown longitudinal stripe, enlarging posteriorly; populations of Naukluft mountains, Nauchas and Remhoogte differ in the shape of the stripe (Fig. 6). Tergites with lateral longitudinal brown stripe. Abdomen covered with setae. Abdominal tergum I same width as metathorax; terga slightly broadening towards tergum VIII, terga IX and X narrowing again. Pleura bright green. Sterna bright green with median brown stripe. Male terminalia (Fig. 7): tergum IX bright green, mesal brown stripe; shorter than tergum VIII, postero-ventral border with dorsal bend in lateral view. Tergum X bright green, mesal brown stripe, roof-shaped in lateral view. Subgenital plate (sternite IX) green with brown-reddish areas; process of subgenital plate broad, dorsal part arch-shaped when seen from posterior, broadly emarginated dorsally. Cerci one segmented, base green, distally brown-reddish; densely covered with setae; diameter mesally round, uniformly curved, slightly narrowed towards the apex; dorsal projec- Head (Fig. 8): moss green, dorso-median brown stripe absent; labrum, tip of lacinia and labial palps yellow; compound eyes brownish with reddish stripe. Mouthparts similar to male. Compound eyes prominent, kidney-shaped, dorso-laterally positioned, smaller than in the male, twice as long as high. Interoccular distance ca. 1.5 times the diameter of one eye, on vertex smaller than on ventral eye margin. Temporal ridge distinct. Distance between antennal sockets ca. 1.5 times the diameter of one antennal socket. Thorax ( Fig. 9): moss green with dark green meso-dorsal stripe; thoracic nota with lateral brown dots. Legs: reddish-green, with brown dots where setae emerge; covered with setae; two ventro-median rows of spikes present on tibia and femur; spikes on tibia black.
Abdomen: moss green, dark green meso-dorsal stripe representing dorsal vessel, lateral margins of terga whitish. Tergum I as broad as metanotum. Pleura and sternites moss green.
Etymology. The species is named after the type locality, the Naukluft Mountains. Comments. Specimens were mainly found in dwarf shrubs. Adult specimens are obviously not associated with grass tussocks.
Pachyphasma gen. n. urn:lsid:zoobank.org:act:E43A1051-968A-4DFE-BB94-EDD0AB742B07 http://species-id.net/wiki/Pachyphasma Description and diagnosis. Based on peptide hormone sequences, Pachyphasma gen. n. is placed as sistergroup to Mantophasma/Sclerophasma (Predel et al. in press). The clade Pachyphasma gen. n. + Mantophasma/Sclerophasma was determined as sistergroup to Tyrannophasma + Praedatophasma. Pachyphasma gen. n. can be distinguished from all other mantophasmatodeans by an abdomen that is shorter than the thorax and a metanotum that is broader than the abdominal tergum I. Pachyphasma was found in the same biotope/habitat as Tyrannophasma gladiator, but on flowering bushes of Compositae while Tyrannophasma mainly inhabits grass tussocks but never these bushes.
The new genus is characterized by several distinct neuropeptide sequences, e.g. periviscerokinin-1 (EAAGLIAFPRTamide) (Predel et al. in press). The respective mass signal ([M+H] + : 1144.6) can be detected in preparations of abdominal perisympathetic organs of males/females/nymphs. Type species. Pachyphasma brandbergense sp. n. Other included species. None thus far. Etymology. the generic group name Pachyphasma is a composition from the Greek word pachys meaning thick and the ending -phasma which is commonly used to term mantophasmatodeans. The gender is neuter.
Paratypes NAMIBIA: Brandberg Plateau, south of Königstein: four males and eight females; same data as holotype; one male and two females in ethanol, one pair in copula in ethanol (NMNW); one male and two females in ethanol (ZMUB); two females in ethanol and one male and one female critical point dried, sputter coated with gold (ZFMK). Two juveniles and six heavily damaged specimens (1 juvenil, 2♀, 3♂) were excluded from the type-series (collection R. Predel).
Description male. Measurements (male holotype followed by paratypes in parentheses, critical point dried male and male in copula not measured, in mm): total length Head (Fig. 13): nearly triangular when seen frontally, orthognathous. Bright green; thin dorso-median brown longitudinal stripe; broader lateral brown longitudinal stripes posterior of the compound eyes; vertex, frons and subgena with reddish areas; head capsule with brown spots where setae emerge; compound eyes grey with reddish stripe; mouthparts including palpi and labrum greenish. Head capsule slightly broader than thorax; sparsely covered with setae. Compound eyes kidney-shaped, prominent, globular, about 1.5 times as long as high; interoccular distance ca. the length of eye, on vertex larger than on ventral eye margin; ocelli absent. Coronal and frontal suture as well as frontoclypeal and temporal ridge indistinct; subgenal ridge distinct, with bend ventrally of compound eye; distance between antennal sockets same as diameter of one socket; anterior tentorial pits dorsomesally of anterior articulation of mandible. Frons with three bulges, two ventromedial of antennal sockets, one in between the antennal sockets. Clypeus trapezoid, with four long setae dorsally; oval sclerite in between clypeus and labrum. Labrum greenish, anteriorly rounded, flat sparsely covered with setae. Maxilla of orthopteroid type, sparsely covered with setae; maxillary palp five-segmented, green, palpomeres two to five covered with setae, palpomere one and two short, nearly as wide as long, palpomere three 2.5 times longer than wide, palpomere four and five ca. twice as long as wide. Labial palp three segmented, green. Scape and pedicel bright green. Scape conical, at base as long as wide. Pedicle half as wide as scape, twice as long as wide. Flagellum yellow-greenish; nearly as long as the entire animal; 25 flagellomeres. Thorax (Fig. 14): compact. Bright green; thin meso-dorsal brown stripe that ends in the anterior third of metanotum; lateral edges of all three nota with thick brown stripe; lateral margin of nota white; pleurae bright green with reddish areas. Completely covered with setae; brown spots where setae emerge along entire thorax. Pronotum nearly squared; overlapping head and mesonotum. Mesonotum not as wide as pronotum, nearly squared. Metanotum narrower than mesonotum; narrowing posteriorly. Coxae large. Legs: bright green. Completely covered with setae; brown spots where setae emerge. Prothoracic leg more massive than meso-and metathoracic leg. Tarsus with 5 brownish tarsomeres; proximal four with euplantulae; large arolium present.
Wings: completely absent. Abdomen: shorter than thorax: enlarging posteriorly; completely covered with setae. Bright green; lateral brown longitudinal stripes on tergites. Abdominal tergum I narrower than metanotum; terga slightly broadening towards tergum VIII; terga IX and X narrowing again. Pleura bright green with yellowish stripe. Sterna bright green.
Abdomen: bright green, dark green meso-dorsal longitudinal stripe; pleura bright green with yellowish longitudinal stripe; sternites bright green.
Etymology. The species is named after the type locality, the Brandberg massif.

Discussion
The discovery of the two new genera of Mantophasmatodea results from extensive collecting activities during the last years (Fig. 19). With it, it can be expected that, at least in Namibia, the majority of higher taxa are now known. It is particularly remarkable, that within the large distributional range of Mantophasma, additional mantophasmatodean taxa have not been found (see Predel et al. in press). Pachyphasma brandbergense may represent a phylogenetic relict which survived only on the Brandberg Massif, an isolated biotope outside the currently accessible range of Mantophasma. At the plateau of the Brandberg massif, Pachyphasma brandbergense occurs sympatrically with  the much larger Tyrannophasma gladiator. Nevertheless, both species occupy different ecological niches. In contrast to Pachyphasma gen. n., the newly described Striatophasma gen. n. represents a widely distributed taxon. Its northernmost known distributional limit seems to be equivalent to the southern limit of Mantophasma (Predel et al. in press). The general distribution of these taxa suggests that Striatophasma is better adapted to conditions of lower rainfall and scattered vegetation. In the field, different morphotypes from different localities of Striatophasma were found with a variable development of the brown dorsal stripe in males (Fig. 6).
Thus far, the monophyly of Pachyphasma and Striatophasma is supported by distinct morphological features and a number of specific peptide hormone sequences which clearly separate these taxa from all other (Predel et al. in press). Most peptide hormones are synthesized in the central nervous system and are released via neurohemal organs which accumulate these messenger molecules. In insects, the ganglia of the different tagmata (head, thorax, abdomen) have each their specific external release sites with tagma-specific peptide hormones which can be analysed by mass spectrometry (see Predel et al. 2005). It is beyond the scope of this manuscript and also beyond our current state of knowledge to present a morphology-based diagnostic key for the separation of all mantophasmatodeans. More data, particularly about within-species variation, are required to address this problem. For the differentiation of the major lineages of Mantophasmatodea, however, mass fingerprints of peptide hormones provide unambiguous information (see Predel et al. 2005, in press) which can be used in a dichotomic key. This character set is already studied in detail with respect to Mantophasmatodea and can be obtained from single specimens (males, females or larvae). It is thus possible to assign newly collected specimens to previously analyzed taxa simply and rapidly and we encourage entomologists to contact one of us in case of problems in obtaining such data. Alternatively, we offer to perform immediate and free of charge analyses for the collector if living or frozen specimens can be provided (we are also working on the development of methods to analyze insects preserved in ethanol).