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Biology of South African Heel-Walkers, with Special Proc. Proc. Ar thropod. Embryo l. Soc. ]pn. 39 ，15-21 (2004) 15 @2 ∞4 Ar thropodan Embryological Society of ]apan ISSN ISSN 1341-1527 Biology Biology of South Mrican Heel-walkers ，with Special Reference to Reproductive Biology (Insecta: (Insecta: Mantophasmatodea) 1 2 Koji Koji TOJ0 )， Ryuichiro MACHIDA )，Klaus ・Dieter KLASS 3) 4 and Mike D. PICKER ) 1) 1) Japan Society for the Promotion of Science σSPS) ，Chiyoda-ku ，To kyo 102-8471 ， Japan; Developmental Mechanisms Laboratory ，Developmental Biology Department ，National lnstitute of Ag robiological Sciences ，Owashi ，Ts ukuba ，lbaraki 305-8634 ，J~ 戸G時 Current Current address: Depart ηtent of Biology ，Faculty of Science ， Shinshu University ，Asahi ，Matsumoto ， Nagano 390-8621 ， Japa 叩 lnstitute 2) lnstitute of Biological Sciences ，University of Ts ukuba ，Ts ukuba ，lbaraki 305-85刀 ， J~仰向 Current Current address: Sugadaira Mon 仰 ze Research Cente r， Universi かof Ts ukuba ，Sanada ， Nagano 386-2201 ， Japan Saatliche 3) Saatliche Naturhis 初rische Sa 抑制l抑留en Dresden ，Museum fur Tierkunde ，Konigsbrucker Landstrase 159 ，A. B. Meyer Bau ， D-Oll09 Dresden ， Germany の ZoologyD ψartment ，University of Cape Town ，Rondebosch 7700 ， Cape Town ， South Africa E-mail: E-mail: [email protected] 必ぽ刀 Abstract Abstract The biology of South Af rican heel-walkers was described ， with special reference to the reproductive ，i. e. ，pre- mating ， mating and ovipositing behaviors. The heel-walkers have apparently adapted their life cycle to the short rainy season ， to survive in a harsh environment. They survive the dry season as eggs underground ， and the first instar nymphs hatch at the beginning of the rainy season ， and grow quickl y， attaining maturity in about three weeks. Mating in in heel-walkers starts after an act of communication known as “drumming ，" and continues for many hours at the bottom bottom of plants (e ・g. ， about three days in KarooPhasma biedouw 仰 sis). Mated females search carefully for a suitable site site to oviposit ， and then deposit a hard pod containing 10 to 12 eggs ， just below the surface of the soi l. Females d巴posit egg pods several times at intervals of three or four days. Af ter the reproductive season at the end of the short rainy rainy season ，their life cy c1 e ends in due course. The life cycle of South Af rican heel-walkers is univoltine. Introduction Introduction In In May 2002 ，a new insect group “ order Mantophasmatodea ，" named after the Mantodea (praying mantises) and Phasmatodea Phasmatodea (walking sticks) ，was established by Klass et al. (2002) ， based on thre 巴 old museum specimens described as as three different species: 1) an ethanol preserved female described as Mantophasma zφわra (type species of the new genus ， the new family and the new order) collected in Namibia in 1909 ， 2) a dried male described as M. subsolana collected collected in Tanzania in 1951 [Klass et al. (2003) have re c1 assified this species as Tanzani φhasma subsolana of a new familyτanzaniophasmatidae familyτanzaniophasmatidae (cf. Table 1)] ， and 3) a 45-million-year 司 old Baltic amber fossil described as Ra 戸toph αsma kerneggeri. kerneggeri. The establishment of this new insect order comes 88 years after the last description of th 巴 order Grylloblattodea Grylloblattodea (= Notoptera) in 1914 (cf. Walk 巴r， 1914). At At around the time of publication ，living mantophasmatodean populations were discovered in Namibia (e. g. ， the highlands highlands around M t. Brandberg ， Erongo Provinc 巴; cf. Adis et al. ， 2002; Zompro et al.， 2002) and South A 企ica (a broad area area of the Western Cape Province including the Namaqualand and part of the Northern Cape Provinc 巴; cf. Picker et al.， 2002). 2002). We have started their taxonomical and biological studies (Klass et al.， 2003; Picker et al. ，2003). Members of the 16 16 K. TO]OETAL Table Table 1 Cl assification of rnernbers of the order Mantophasrnatodea (after Klass et al.. 2003; Zornpro et α1. ， 2003). Family Family Genus Species 苛 ])e locality Reference Collection site 本 加fantophasmatidae MantoPh αsma z φ， 'hyra Brandberg (N amibia) Klass et al. (2002 ，2003) ， Zompro et al. (2002 ，2003) SclerOPhas 刑 α pa 開 szsens 日 Paresisberg (N amibia) Kl ass et al. (2003) 2 τanzaniophasmatidae τanzaniophasmatidae Tanzaniothas 削 α subsolana Uf ipa Dish (百 nzania) Kl ass et al. (2002 ，2003). Zompro et al. (2002 ，2003) Austrophasmatidae Austrophasmatidae Austro 戸hasma caled 側 ens 日 Caledon (South Afr ica) Klass et al. (2003) 4 Aぃ gansbam 由1515 Gansbaai (South Afr ica) Klass et al. (2003) 5 A. A. ra 羽 so 間 ille 悶悶 Rawsonville (South Afr ica) Klass et al. (2003) 6 Loboph 四州 問 delinghuysensis Redelinghuis (South Afr ica) Kl ass et al. (2003) 7 H 四niloboPh α幻叫削ontague 悶 zs Montagu (South Afr ica) Klass et al. (2003) 8 f五aroophasm σ biedouu 側 SlS Biedouw (South A企ica) Klass et al. (2003) 9 K botterkloofensis Botterkloof Pass (South Afr ica) Klass et al. (2003) 10 N 酔叩quaph 府間σ 叩 Mψe 悶 is 0' okiep (South A止ica) Klass et al. (2003) 11 Described Described mantophasrnatodean taxa of uncertain placement Praed 包oph 国間α 削 araisi Karasberg CNarnibia) Zompro et al. (2002 ，2003) ，Kl ass et al. (2003) 12 Rゆtophas 附 a kerneggeri* * Baltic area (Lithuania) Klass et al. (2002 ，2003) ， Zompro et al. (初 02 ，20 日3) ηrannoPh σsma gladiator Brandberg (N amibia) Zompro et al. (2003) *Refer *Refer to numbers in Fig. 1 '" '" A* 45-million-year-old fossil species collected 立orn Baltic amber order Mantophasmatodea grow to a length of 2-3 cm ，without wings and ocelli ，and walk on their “heels"(= th 巴 fused first first to third tarsomeres in each leg) whereas insects usually walk on their “toes" (= pretarsus + distal parts of tarsus). tarsus). Af ter this unique and diagnostic style of walking ，we hav 巴 proposed the common name “heel-walkers" (Machida and Tojo ， 2003; Picker et al.， 2003; も'I o and Machida ， 2003). So fa r， four living species from Namibia and eight from South Afr ica have been described by Zompro et al. (2002 ，2003) and Klass et al. (2002 ， 2003). Klass et al. (2003) have not only described the new South Afr ican heel-walkers ，but reclassified all speci 巴s hitherto descr ・ibed (Table 1). According to Klass et al. (2002 ，2003) ，the new order Mantophasmatodea are phenotypically orthopteroid insects ， 。 1 4' 4' 5 Fig. Fig. 1 Distribution of rnantophasrnatodean species. For the locality and rnantophasrnatodean narnes ， refer toτable 1. These rnaps are reconstructed frorn previous studies (K1 ass et al. ，2∞>2， 2003; Picker et al. ， 2002; Zornpro et al.， 2002 ， 2003) plus sorne new data (open circles) frorn Ot 町 2003 field trip. REPRODUCTIVE BIOLOGY OF MANTOPHASMATODEA 17 with with features which allow one to differentiate them from the dictyopteroid insect orders Mantodea ， Blattodea and Isoptera ， and from the other orthopteroid orders Phasmatodea ，Orthoptera ，Grylloblattodea ， Plecoptera and Dermaptera. Dermaptera. Twenty-one unique synapomorphic characters of the new order were discussed by Klass et a l. (2003). On the the other hand ，Dallai et al. (2003) examined the relationships of heel-walkers with other orthopteroid taxa based on a cladistic cladistic analysis of the comparative morphology of sperm ， and supported that praying mantises are the sister group of heel-walkers . However ， heel-walkers also share many morphological characters with members of other orthopteroid taxa ，and their phylogenetic position still remains controversia l. We have started studying the comparative embryology and reproductive biology of Mantophasmatodea ， to clarify arguments pertaining to the groundplan and its evolutionary transition ，and reconstructing the phylogeny of orthopteroids orthopteroids (cf. Machida et al.， 2004; Tsutsumi et al.， 2004). In the first part of our studies ，we outline the biology ， with with special reference to reproduction ， mainly of South Afr ican heel-walkers. Observation and Discussion We had been observing the heel-walkers' behavior in the field 仕om winter in 2002 and 2003. At the same time ，we had had also observed their behavior ， with special reference to reproduction ， in the laboratory. Heel-walkers were collected in in South Afri ca from a specific succulent desert area known as “Namaqualand" (or “karoo ，"“ succulent karoo ウ or the cape cape floral plains including a type of plain called the “Fynbos" (cf. Table 1; Fig. 1) ， in winter to spring (仕 om early August to mid September) of 2002 and 2003. Nymphs and adults were kept at room temperature in separate plastic cases cases to prevent cannibalism. Fruit flies as food and a little water were provided every day . A mature male and female ， a pair who had display ed mutual “drumming" pre-mating behavior (see the “Mating " section) ，were reared together in a case ， and checked for mating behavior and copulation. Copulated females were moved to a separate plastic case with soil soil collected 仕om the fi e ld for oviposition. L~俳冷 Cりycieた'e olS お'out ， 幼hAj， 斤ri 沈1κ ca 仰悶 hee ωel South Af企 nκcan heel-walkers mostly liv e in a speci 凶f白ic succulent desert area called the Namaqualand ，and on plains known as the Fynbos. Namaqualand ，where KarooPhasma biedouwensis ，K botterklo o， 伽 sis and NamaquaPhasma ookiepensis ookiepensis are found ，is a desert area spreading over southern Namibia and part of South A 企ica (Fig. 1). In the region ， however ， rains from June to late August (or till early September) ， the “ rainy season ，" transform the land into a spectacular spectacular flowering garden of succulent plants in spring (l ate August and September). However ， the land reverts to its its desert state in the summer season from late September to April or May of the next year. In the dry season ， the daily difference difference between high and low temperatures is as much as 20-30 0C.
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