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Data from: Functional monocentricity with holocentric characteristics and chromosome-specific centromeres in a stick insect
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Data from: Transitions between phases of genomic differentiation during stick-insect speciation
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Data from: Divergent dynamics of sexual and habitat isolation at the transition between stick insect populations and species
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Data from: Selection on a genetic polymorphism counteracts ecological speciation in a stick insect
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Data from: Old World and New World Phasmatodea: phylogenomics resolve the evolutionary history of stick and leaf insects
Phasmatodea comprises over 3,000 extant species and stands out as one of the last remaining insect orders for which a robust, higher-level phylogenetic hypothesis is lacking. New research suggests that the extant diversity is the result of a surprisingly recent and rapid radiation that has been difficult to resolve with standard Sanger sequence data. In order to resolve the early branching events of stick and leaf insects, we analyzed transcriptomes from 61 species, including 38 Phasmatodea species comprising all major clades and 23 outgroup taxa, including all other Polyneoptera orders. Using a custom-made ortholog set based on reference genomes from four species, we identified on average 2,274 orthologous genes in the sequenced transcriptomes. We generated various sub-alignments and performed maximum-likelihood analyses on several representative datasets to evaluate the effect of missing data and matrix composition on our phylogenetic estimates. Based on our new data, we are able to reliably resolve the deeper nodes between the principal lineages of extant Phasmatodea. Among Euphasmatodea, we provide strong evidence for a basal dichotomy of Aschiphasmatodea and all remaining euphasmatodeans, the Neophasmatodea. Within the latter clade, we recovered a previously unrecognized major New World and Old World lineage, for which we introduce the new names Oriophasmata tax. nov. ("Eastern phasmids") and Occidophasmata tax. nov. ("Western phasmids"). Occidophasmata comprise Diapheromerinae, Pseudophasmatinae, and Agathemera, whereas all remaining lineages form the Oriophasmata, including Heteropterygidae, Phylliinae, Bacillus, Lonchodidae (Necrosciinae + Lonchodinae), Clitumninae, Cladomorphinae, and Lanceocercata. We furthermore performed a divergence time analysis and reconstructed the historical biogeography for stick and leaf insects. Phasmatodea either originated in Southeast Asia or in the New World. Our results suggest that the extant distribution of Phasmatodea is largely the result of dispersal events in a recently and rapidly diversified insect lineage rather than the result of vicariant processes.
FIGURE 6 in Megacraniinae-The Palm Stick Insects: A new subfamily of Old World Phasmatodea and a redefinition of Platycraninae Brunner v. Wattenwyl, 1893 (Phasmatodea: "Anareolatae")
FIGURE 6. Details of terminalia of ♂♂: A. Megacrania spina Hsiung, 2007, ventral view; B. Megacrania spina Hsiung, 2007, caudal view of anal segment (tergum X); C. Ophicrania conlei Gottardo, 2011, ventral view; D. Platycrana viridana (Olivier, 1792), ventral view.
FIGURE 10 in Megacraniinae-The Palm Stick Insects: A new subfamily of Old World Phasmatodea and a redefinition of Platycraninae Brunner v. Wattenwyl, 1893 (Phasmatodea: "Anareolatae")
FIGURE 10. Details of Platycrana viridana (Olivier, 1792), Platycraninae: Platycranini: A. Terminalia of ♀, lateral view: Kei Islands [coll. FH, No. 0173-5]; B. Terminalia of ♀, dorsal view: Kei Islands [coll. FH, No. 0173-5]; C. Terminalia of ♀, ventral view: Kei Islands [coll. FH, No. 0173-6]; D. Terminalia of ♂, lateral view: Philippines, Panay Island, Mt. Nangtud [coll. FH, No. 0173-13]; E. Terminalia of ♂, dorsal view: Philippines, Panay Island, Mt. Nangtud [coll. FH, No. 0173-13]; F. Terminalia of ♂, ventral view: Philippines, Panay Island, Mt. Nangtud [coll. FH, No. 0173-13]; G. Head, pro- and mesothorax of ♂ in lateral view: Philippines, Panay Island, Mt. Nangtud [coll. FH, No. 0173-13]; H. Head, pro- and mesothorax of ♀ in lateral view: Kei Islands [coll. FH, No. 0173-3]; J. Head, pro- and mesothorax of ♀ in lateral view: Kei Islands [coll. FH, No. 0173-6].
FIGURE 3 in Megacraniinae-The Palm Stick Insects: A new subfamily of Old World Phasmatodea and a redefinition of Platycraninae Brunner v. Wattenwyl, 1893 (Phasmatodea: "Anareolatae")
FIGURE 3. Habitus of Ophicrania Kaup, 1871 species ♂♂ (Megacraniinae subfam. nov.): A. Ophicrania vittipennis (Stål, 1875): Philippines, Leyte Island, Mahaplag [coll. FH, No. 0726-12]; B. Ophicrania capito (Westwood, 1859): Borneo, Sabah, Crocker Range, Mt. Trus Madi 1200m [coll. FH, No. 0039-2]; C. Ophicrania sp. nov.: Borneo, Kalimantan, Malinau [coll. FH, No. 0832-1]; D. Ophicrania sagittarius Bresseel & Bushell, 2009: Philippines, Mindanao Island, Agusan del Sur, Rosario [coll. FH, No. 1131-1].
FIGURE 9 in Megacraniinae-The Palm Stick Insects: A new subfamily of Old World Phasmatodea and a redefinition of Platycraninae Brunner v. Wattenwyl, 1893 (Phasmatodea: "Anareolatae")
FIGURE 9. Habitus of Platycrana viridana (Olivier, 1792), Platycraninae: Platycranini Brunner v. Wattenwyl, 1893: A. ♀ do- solateral view: Kei Islands [coll. FH, No. 0173-3]; B. ♀ dorsolateral view: Kei Islands [coll. FH, No. 0173-6]; C. ♀ dorsolateral view: Philippines, Panay Island, Mt. Nangtud [coll. FH, No. 0173-11]; D. ♀ dorsolateral view: Philippines, Panay Island, Mt. Nangtud [coll. FH, No. 0173-11].
FIGURE 13. Supposed relationships between Phasmatidae s in Megacraniinae-The Palm Stick Insects: A new subfamily of Old World Phasmatodea and a redefinition of Platycraninae Brunner v. Wattenwyl, 1893 (Phasmatodea: "Anareolatae")
FIGURE 13. Supposed relationships between Phasmatidae s. str. (= Lanceocercata), including the subordinate Megacraniinae subfam. nov., and the sister group Platycraninae sensu nov. with the two contained tribes Platycranini and Stephanacridini. The tree shown is merely meant to visualize the estimated relationships here suggested. It does not restrict to defining apomorphies for each clade but also uses key-features that are helpful for distinction. Although the results of Robertson et al. (2018, Fig. 6) and Simon et al. (2019, Fig. 2), both based on molecular data, are incorporated the above tree cannot be interpreted as a strict phylogenetic cladogram.
FIGURE 12 in Megacraniinae-The Palm Stick Insects: A new subfamily of Old World Phasmatodea and a redefinition of Platycraninae Brunner v. Wattenwyl, 1893 (Phasmatodea: "Anareolatae")
FIGURE 12. Eggs of Megacraniinae subfam. nov. (A-H), Platycraninae: Platycranini (J-K) and Platycraninae: Stephanacridini (L-M): A. Megacrania batesii Kirby, 1896, dorsolateral view: NE-Australia, Queensland, Cape Tribulation [coll. FH, No. 0175-E]; B. Megacrania tsudai Shiraki, 1933, dorsolateral view: Taiwan, Pindang County, Gio-Pon [coll. FH, No. 0493-E]; C. Megacrania brocki Hsiung, 2002 dorsolateral view: Indonesia, Prov. Maluku, Buru Island [coll. FH, No. 1012-E]; D. Erastus apalamnus Rehn, 1904, dorsal view: Indonesia, Prov. Maluku, Obi Island [coll. FH, No. 1212-E]; E. Erastus apalamnus Rehn, 1904, dorsal view: Indonesia, Prov. Maluku, Obi Island [coll. FH, No. 1212-E]; F. Ophicrania nigroplagiatus var. (Redtenbacher, 1906), dorsolateral view: Philippines, Luzon Island [coll. FH, No. 0206-E]; G. Graeffea crouanii (Le Guillou, 1841), dorsolateral view: captive reared from Melanesia, Fiji, Suva [coll. FH, No. 0204-E2]; H. Ophicrania apterus var. (Redtenbacher, 1908), dorsolateral view: Papua New Guinea, Morobe Prov., Aseki [coll. FH, No. 0222-E1], J. Platycrana viridana (Olivier, 1792), dorsal view: Philippines, Panay Island, Mt. Nangtud [coll. FH, No. 0173-E1]; K. Platycrana viridana (Olivier, 1792), lateral view: Philippines, Panay Island, Mt. Nangtud [coll. FH, No. 0173-E1]; L. Hermarchus pythonius var. (Westwood, 1859), dorsolateral view: captive reared from Fiji Islands [coll. FH, No. 0144-E1]; M. Macrophasma lyratus (Redtenbacher, 1908), dorsolateral: West Papua, Prov. Papua Barat, Fak-Fak [coll. FH, No. 1081-E].
FIGURE 8 in Megacraniinae-The Palm Stick Insects: A new subfamily of Old World Phasmatodea and a redefinition of Platycraninae Brunner v. Wattenwyl, 1893 (Phasmatodea: "Anareolatae")
FIGURE 8. Terminalia of ♀♀ of Megacraniinae subfam. nov.: A. Erastus apalamnus Rehn, 1904, lateral view (with egg): Indonesia, Prov. Maluku, Obi Island [coll. FH, No. 1212-5]; B. Graeffea crouanii (Le Guillou, 1841), lateral view: Melanesia, Tonga, Tongatapu [coll. FH, No. 0204-2]; C. Megacrania sp. nov., lateral view: Papua New Guinea, Morobe Prov., Aseki [coll. FH, No. 1014-2]; D. Megacrania phelaus (Westwood, 1859), lateral view: Solomon Islands, Maramasike Island prope Malaita [coll. FH, No. 0174-7]; E. Megacrania tsudai Shiraki, 1933, ventral view: Taiwan, Pindang County, Gio-Pon [coll. FH, No. 0493-1]; F. Megacrania phelaus (Westwood, 1859), dorsal view: Solomon Islands, Maramasike Island prope Malaita [coll. FH, No. 0174-7].
FIGURE 11 in Megacraniinae-The Palm Stick Insects: A new subfamily of Old World Phasmatodea and a redefinition of Platycraninae Brunner v. Wattenwyl, 1893 (Phasmatodea: "Anareolatae")
FIGURE 11. Habitus and details of members of Platycraninae: Stephanacridini Günther, 1953: A. Macrophasma oreitrephes (Günther, 1929) ♀: Papua New Guinea, Morobe Prov., Aseki [coll. FH, No. 0107-2]; B. Hermarchus insignis (Kaup & Heyden, 1871) ♀: captive reared from 'Australian Islands' [coll. FH, No. 0144-13]; C. Eucarcharus sp. ♂: captive reared from Philippines, Mindanao Island, Prov. Bukidnon, Mt. Apo [coll. FH, No. 0630-7]; D. Hermarchus insignis (Kaup & Heyden, 1871) terminalia of ♀, lateral view: captive reared from 'Australian Islands' [coll. FH, No. 0144-13]; E. Hermarchus insignis (Kaup & Heyden, 1871) terminalia of ♀, dorsal view: captive reared from 'Australian Islands' [coll. FH, No. 0144-13]; F. Hermarchus insignis (Kaup & Heyden, 1871) terminalia of ♀, ventral view: captive reared from 'Australian Islands' [coll. FH, No. 0144-13]; G. Stephanacris draconius Hennemann & Conle, 2006 terminalia of ♀, dorsal view: Papua New Guinea, Morobe Prov., Wau [coll. FH, No. 0377-2]; H. Hermarchus pythonius var. (Westwood, 1859) terminalia of ♂, lateral view: captive reared from Fiji Islands [coll. FH, No. 0144-2]; J. Hermarchus pythonius var. (Westwood, 1859) terminalia of ♂, ventral view: captive reared from Fiji Islands [coll. FH, No. 0144-2]; K. Stephanacris globiceps Redtenbacher, 1908 terminalia of ♂, dorsal view: West P- pua, Arfak Mountains, Manokwari, Maibri village, 670m [coll. FH, No. 1000-1]; L. Phasmotaenia lanyuhensis Huang & Brock, 2001 terminalia of ♂, ventral view: Taiwan, Lanyuh Island [coll. FH, No. 0616-2].
FIGURE 7 in Megacraniinae-The Palm Stick Insects: A new subfamily of Old World Phasmatodea and a redefinition of Platycraninae Brunner v. Wattenwyl, 1893 (Phasmatodea: "Anareolatae")
FIGURE 7. Terminalia of ♂♂ of Megacraniinae subfam. nov.: A. Megacrania spina Hsiung, 2007, lateral view: West Papua, Prov. Papua Barat, Fak-Fak [coll. FH, No. 1083-1]; B. Megacrania spina Hsiung, 2007, dorsal view: West Papua, Prov. Papua Barat, Fak-Fak [coll. FH, No. 1083-1]; C. Megacrania spina Hsiung, 2007, ventral view: West Papua, Prov. Papua Barat, FakFak [coll. FH, No. 1083-1]; D. Megacrania sp. nov., lateral view: Papua New Guinea, Morobe Prov., Aseki [coll. FH, No. 10143]; E. Megacrania sp. nov., dorsal view: Papua New Guinea, Morobe Prov., Aseki [coll. FH, No. 1014-3]; F. Graeffea leveri (Günther, 1937), lateral view: captive reared from Solomon Islands, Russell Island NW of Guadalcanal [coll. FH, No. 0635-1]; G. Ophicrania capito (Westwood, 1859), dorsal view: Borneo, Sabah, Crocker Range, Mt. Trus Madi 1200m [coll. FH, No. 0039-2]; H. Ophicrania conlei Gottardo, 2011, dorsal view: Philippines, Mindoro Island, Mt. Halcon [coll. FH, No. 0611-1]; J. Ophicrania sp. nov.: Borneo, Kalimantan, Malinau [coll. FH, No. 0832-1].
FIGURE 1 in Megacraniinae-The Palm Stick Insects: A new subfamily of Old World Phasmatodea and a redefinition of Platycraninae Brunner v. Wattenwyl, 1893 (Phasmatodea: "Anareolatae")
FIGURE 1. Habitus of of Megacrania Kaup, 1871 species (Megacraniinae subfam. nov.): A. Megacrania phelaus (Westwood, 1859) ♀: Solomon Islands, Maramasike Island prope Malaita [coll. FH, No. 0174-7]; B. Megacrania phelaus (Westwood, 1859) ♂: Solomon Islands, Maramasike Island prope Malaita [coll. FH, No. 0174-8]; C. Megacrania spina Hsiung, 2007 ♂: West Papua, Prov. Papua Barat, Fak-Fak [coll. FH, No. 1083-1].
FIGURE 5 in Megacraniinae-The Palm Stick Insects: A new subfamily of Old World Phasmatodea and a redefinition of Platycraninae Brunner v. Wattenwyl, 1893 (Phasmatodea: "Anareolatae")
FIGURE 5. Details of head and prothorax in ventral view (Megacraniinae subfam. nov.). A. Graeffea meridionalis Günther, 1932 ♂: Solomon Islands, Malaita Island, Haburarumu Uru, 100-250m [coll. FH, No. 1195-1]; B. Erastus apalamnus Rehn, 1904 ♀: Indonesia, Prov. Maluku, Obi Island [coll. FH, No. 1212-5].
FIGURE 4 in Megacraniinae-The Palm Stick Insects: A new subfamily of Old World Phasmatodea and a redefinition of Platycraninae Brunner v. Wattenwyl, 1893 (Phasmatodea: "Anareolatae")
FIGURE 4. Details of head and prothorax, A, B, D-G dorsal view, C dorsolateral view, H-J lateral view (Megacraniinae subfam. nov.): A. Megacrania brocki Hsiung, 2002 ♀: Indonesia, Prov. Maluku, Buru Island [coll. FH, No. 1012-3]; B. Megacrania sp. nov. ♀: Papua New Guinea, Morobe Prov., Aseki [coll. FH, No. 1014-2]; C. Megacrania spina Hsiung, 2007 ♂: West Papua, Prov. Papua Barat, Fak-Fak [coll. FH, No. 01083-1]; D. Erastus apalamnus Rehn, 1904 ♂: Indonesia, Prov. Maluku, Obi Island [coll. FH, No. 1212-5]; E. vittipennis (Stål, 1875): Philippines, Leyte Island, Mt. Balocaue [coll. FH, No. 0726-8]; F. Ophicrania capito (Westwood, 1859): Borneo, Sabah, Crocker Range [coll. FH, No. 0039-1]; G. Graeffea meridionalis Günther, 1932 ♂: Solomon Islands, Malaita Island, Haburarumu Uru, 100-250m [coll. FH, No. 1195-1]; H. Erastus apalamnus Rehn, 1904 ♀: Indonesia, Prov. Maluku, Obi Island [coll. FH, No. 1212-5]; J. Megacrania phelaus (Westwood, 1859) ♂: captive reared from Solomon Islands, Malaita Island [coll. FH, No. 0174-10]. Arrow indicates the opening of the prothoracic defensive glands.
FIGURE 2 in Megacraniinae-The Palm Stick Insects: A new subfamily of Old World Phasmatodea and a redefinition of Platycraninae Brunner v. Wattenwyl, 1893 (Phasmatodea: "Anareolatae")
FIGURE 2. Habitus of Graeffea Brunner v. Wattenwyl, 1868 and Ophicrania Kaup, 1871 species (Megacraniinae subfam. nov.): A. Graeffea meridionalis Günther, 1932 ♂: Solomon Islands, Malaita Island, Haburarumu Uru, 100-250m [coll. FH, No. 1195-1]; B. Graeffea crouanii (Le Guillou, 1841) ♀: Melanesia, Tonga, Tongatapu [coll. FH, No. 0204-1]; C. Ophicrania nigroplagiatus var. (Redtenbacher, 1906) ♀: Philippines, Luzon Island [coll. FH, No. 0206-1]; D. Ophicrania apterus var. (Redtenbacher, 1908) ♀: Papua New Guinea, Morobe Prov., Aseki [coll. FH, No. 0222-1].
FIGURE 21. Habitat and live insects. A in Studies on Neotropical Phasmatodea XXIV: Andeocalynda n. gen., a new genus of Andean stick insects, with the descriptions of nine new species from Colombia and Ecuador (Phasmatodea: "Anareolatae": Diapheromeridae: Diapheromerinae)
FIGURE 21. Habitat and live insects. A. Moist and cloudy dwarf forest in El Cajas National Park, near Cuenca at ca. 3100 m; B. ♀ PT of Andeocalynda decorata n. sp. at the type-locality [coll. FH, No. 0519-1]; C. Mating couple of Andeocalynda tenuis n. sp. at the type-locality, the ♀ feeding on the unidentified host plant [coll. OC]; D. ♂ PT of Andeocalynda tenuis n. sp. on one of it's unidentified host plants [coll. FH, No. 0518-2].
FIGURE 22 in Studies on Neotropical Phasmatodea XXIV: Andeocalynda n. gen., a new genus of Andean stick insects, with the descriptions of nine new species from Colombia and Ecuador (Phasmatodea: "Anareolatae": Diapheromeridae: Diapheromerinae)
FIGURE 22. Map of NW-South America (Colombia and Ecuador) showing all currently known records of Andeocalynda n. gen. marked by yellow dots. The small number of records in total and the big gaps within the distributional range show our still fractional knowledge of the distribution of this new genus.
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