Reconstructing the nonadaptive radiation of an ancient lineage of ground‐dwelling stick insects (Phasmatodea: Heteropterygidae)

Stick and leaf insects (Phasmatodea) are large terrestrial herbivorous arthropods known for masquerading as plant parts such as bark, twigs and leaves. Their evolutionary history is largely shaped by convergent evolution associated with adaptive radiations on geographically isolated landmasses that have repeatedly generated ground-dwelling ecomorphs. The members of one lineage, however, the Oriental Heteropterygidae, are morphologically rather uniform, and have a predominantly ground-dwelling lifestyle. The phylogeny of Heteropterygidae that comprises approximately 130 described species is controversial and remains uncertain. In particular, the systematic position of the giant Jungle Nymph Heteropteryx dilatata, whose males are capable of flight and exhibit the most plesiomorphic wing morphology among extant phasmatodeans, is of major interest to the scientific community. Here, we analysed a set of seven nuclear and mitochondrial genes to infer the phylogeny of Heteropterygidae covering the group's overall diversity. The divergence time estimation and reconstruction of the historical biogeography resulted in an ancestral distribution across Sundaland with long distance dispersal events to Wallacea, the Philippines and the South Pacific. We were able to resolve the relationships among the three principal subgroups of Heteropterygidae and revealed the Dataminae, which contain entirely wingless small forms, as the sister group of Heteropteryginae + Obriminae. Within Heteropteryginae, Haaniella is recovered as paraphyletic in regard to Heteropteryx. Consequently, Heteropteryx must be considered a subordinate taxon deeply embedded within a flightless clade of stick insects. Within Obriminae, the Bornean Hoploclonia is strongly supported as the earliest diverging lineage. Based on this finding, we recognize only two tribes of equal rank among Obriminae, the Hoplocloniini trib. nov. and Obrimini sensu nov. Within the latter, we demonstrate that previous tribal assignments do not reflect phylogenetic relationships and that a basal splitting event occurred between the wing-bearing clade Miroceramia + Pterobrimus and the remaining wingless Obrimini. The Philippine genus Tisamenus is paraphyletic with regard to Ilocano hebardi, thus, we transfer the latter species to Tisamenus as Tisamenus hebardi comb. nov. and synonymize Ilocano with Tisamenus. We discuss character transformations in the light of the new phylogenetic results and conclude that the current taxonomic diversity appears to be mainly driven by allopatry and not to be the result of niche differentiation. This radiation is thus best described as a nonadaptive radiation

Changes to the Fossil Record of Insects through Fifteen Years of Discovery

The first and last occurrences of hexapod families in the fossil record are compiled from publications up to end-2009. The major features of these data are compared with those of previous datasets (1993 and 1994). About a third of families (>400) are new to the fossil record since 1994, over half of the earlier, existing families have experienced changes in their known stratigraphic range and only about ten percent have unchanged ranges. Despite these significant additions to knowledge, the broad pattern of described richness through time remains similar, with described richness increasing steadily through geological history and a shift in dominant taxa, from Palaeoptera and Polyneoptera to Paraneoptera and Holometabola, after the Palaeozoic. However, after detrending, described richness is not well correlated with the earlier datasets, indicating significant changes in shorter-term patterns. There is reduced Palaeozoic richness, peaking at a different time, and a less pronounced Permian decline. A pronounced Triassic peak and decline is shown, and the plateau from the mid Early Cretaceous to the end of the period remains, albeit at substantially higher richness compared to earlier datasets. Origination and extinction rates are broadly similar to before, with a broad decline in both through time but episodic peaks, including end-Permian turnover. Origination more consistently exceeds extinction compared to previous datasets and exceptions are mainly in the Palaeozoic. These changes suggest that some inferences about causal mechanisms in insect macroevolution are likely to differ as well

The Phasmatodea and Raptophasma n. gen., Orthoptera incertae sedis, in Baltic Amber (Insecta: Orthoptera).

The stick insects (Orthoptera: Phasmatodea) in Baltic amber are revised. A new family of Areolatae, Archipseudophasmatidae n. fam., is introduced based on the genus Archipseudophasma n. gen., with the type-species A. phoenix n. sp., differing from the closely related two families, Heteronemiidae and Pseudophasmatidae, in the strongly elongated third segment of the antennae and the fully developed tegmina, projecting beyond the abdomen. It includes two subfamilies, of which only one is named. The second is based on nymphs only, which are useless to describe as new taxa. Pseudoperla lineata PICTET & BERENDT, 1854, represents a new genus of Archipseudophasmatinae: Balticophasmatini n. trib., Balticophasma n. gen., and is not a synonym of Pseudoperla gracilipes PICTET & BERENDT, 1854, as stated by HAGEN in GERMAR & BERENDT in BERENDT (1856: 39). Electrobaculum Sharov, 1968, is the only genus of Pseudophasmatidae: Pseudophasmatinae: Electrobaculini n. trib. Several of the specimens sent to the author for study are actually not phasmids but Orthoptera incertae sedis represented by Raptophasma n. gen., with the type-species R. kerneggeri n. sp. Even though this genus from the first glance looks quite similar to Mantodea, it appears to show more features characteristic of Phasmatodea

Note on Pseudodiacantha macklottii (de Haan, 1842)

The type species of Lamachus Stål, 1877, Lamarchinus Uvarov, 1940, Lopaphus Redtenbacher, 1908 (not Lopaphus Westwood, 1859), Lopaphodes Karny, 1923, Orxines Stål, 1875, Pseudodiacantha Redtenbacher are reviewed, with the result that Lamachus and Lamarchinus are found to be new synonyms of Orxines and that the correct name of one of the most famous phasmids has to be corrected into Pseudodiacantha macklottii (de Haan, 1842

SEM-Studie des Eies von Bacteria nodulosa REDTENBACHER, 1908, und die Erstbeschreibung des Weibchens (Phasmatodea: Heteronemiidae, Heteronemiinae, Heteronemiini)

SEM-study of the egg of Bacteria nodulosa REDTENBACHER, 1908, and the first description of the female (Phasmatodea: Heteronemiidae, Heteronemiinae, Heteronemiini). The female and the egg of Bacteria nodulosa REDTENBACHER, 1908, are described for the first time and the male is redescribed. A lectotype is designated. The egg was examined using a scanning electron microscope

A replacement name for Microphasma Zompro, 1999

Microphasma Zompro, 1999, a genus of Phasmatodea, is preoccupied by Microphasma Woltereck, 1909 (Crustacea). Miniphasma n.n. is introduced as a replacement nam

Note on Pseudodiacantha macklottii (de Haan, 1842)

The type species of Lamachus Stål, 1877, Lamarchinus Uvarov, 1940, Lopaphus Redtenbacher, 1908 (not Lopaphus Westwood, 1859), Lopaphodes Karny, 1923, Orxines Stål, 1875, Pseudodiacantha Redtenbacher are reviewed, with the result that Lamachus and Lamarchinus are found to be new synonyms of Orxines and that the correct name of one of the most famous phasmids has to be corrected into Pseudodiacantha macklottii (de Haan, 1842