RESUMEN
Type specimens of 42 taxa of Phasmatodea (including probable type specimens of 14 taxa) have been located in the Eidgenössisches Technisches Hochschulzentrum, Zürich. The species are listed alphabetically, with the number of specimens, sex and locality data. Some minor taxonomic changes are proposed: 1 new combination, 1 revised combination, 2 lectotype designations.
Asunto(s)
Neoptera , Animales , SuizaRESUMEN
The concept that complex ancestral traits can never be recovered after their loss is still widely accepted, despite phylogenetic and molecular approaches suggest instances where phenotypes may have been lost throughout the evolutionary history of a clade and subsequently reverted back in derived lineages. One of the first and most notable examples of such a process is wing evolution in phasmids; this polyneopteran order of insects, which comprises stick and leaf insects, has played a central role in initiating a long-standing debate on the topic. In this study, a novel and comprehensive time tree including over 300 Phasmatodea species is used as a framework for investigating wing evolutionary patterns in the clade. Despite accounting for several possible biases and sources of uncertainty, macroevolutionary analyses consistently revealed multiple reversals to winged states taking place after their loss, and reversibility is coupled with higher species diversification rates. Our findings support a loss of or reduction in wings that occurred in the lineage leading to the extant phasmid most recent common ancestor, and brachyptery is inferred to be an unstable state unless co-opted for nonaerodynamic adaptations. We also explored how different assumptions of wing reversals probability could impact their inference: we found that until reversals are assumed to be over 30 times more unlikely than losses, they are consistently inferred despite uncertainty in tree and model parameters. Our findings demonstrate that wing evolution is a reversible and dynamic process in phasmids and contribute to our understanding of complex trait evolution. [Dollo's law; Phasmatodea; phylogenetic comparative methods; polyneoptera; reversals; wing.].
Asunto(s)
Evolución Biológica , Alas de Animales , Animales , Insectos/genética , FilogeniaRESUMEN
The genus Pterinoxylus Serville, 1838 is redescribed and revised at the species level. It is distributed throughout most of Central America, the northern half of South America and also has one species on the Lesser Antilles. Detailed descriptions, notes on intraspecific variability and illustrations are provided for all six known species. Keys are presented to distinguish males, females and eggs. Two new species are described from Costa Rica: P. cocoense n. sp. from both sexes and the eggs and P. speciosus n. sp. from both sexes. The female of P. perarmatus (Redtenbacher, 1908) is described and illustrated for the first time, as are the eggs of the type-species P. eucnemis Serville, 1838 and P. perarmatus (Redtenbacher, 1908). The external morphology of all species shows considerable intraspecific variability, which is discussed and illustrated. While P. cocoense n. sp. is an endemic and the only stick insects that has so far become known from Cocos Island a small island some 550 km off the Costa Rican Pacific coast, all other species appear to have fairly wide distributional ranges. Maps show the distributions of all six known species. Type-specimens of the two newly described species are deposited in the collections of MNCR-A (Costa Rica) and Zoologisches Museum und Universität, Hamburg, Germany (ZMUH).
Asunto(s)
Neoptera , Masculino , Femenino , Animales , Costa RicaRESUMEN
Phasmatodea species diversity lies almost entirely within its suborder Euphasmatodea, which exhibits a pantropical distribution and is considered to derive from a recent and rapid evolutionary radiation. To shed light on Euphasmatodea origins and diversification, we assembled the mitogenomes of 17 species from transcriptomic sequencing data and analysed them along with 22 already available Phasmatodea mitogenomes and 33 mitogenomes representing most of the Polyneoptera lineages. Maximum Likelihood and Bayesian Inference approaches retrieved consistent topologies, both showing the widespread conflict between phylogenetic approaches and traditional systematics. We performed a divergence time analysis leveraging ten fossil specimens representative of most polyneopteran lineages: the time tree obtained supports an older radiation of the clade with respect to previous hypotheses. Euphasmatodea diversification is inferred to have started ~ 187 million years ago, suggesting that the Triassic-Jurassic mass extinction and the breakup of Pangea could have contributed to the process. We also investigated Euphasmatodea mitogenomes patterns of dN, dS and dN/dS ratio throughout our time-tree, trying to characterize the selective regime which may have shaped the clade evolution.
Asunto(s)
Genoma Mitocondrial , Insectos/clasificación , Insectos/genética , Filogenia , Animales , Composición de Base/genética , Teorema de Bayes , Calibración , Fósiles , Variación Genética , Funciones de Verosimilitud , Factores de TiempoRESUMEN
The present paper describes 16 new species and one new genus from French Guiana and numerous taxonomic changes are proposed prior to the publication of a comprehensive guide to the Phasmatodea of French Guiana. The following 16 new species are described and illustrated: Phanocles procerus n. sp., Phanocloidea lobulatipes n. sp., Cladomorphus guianensis n. sp., Hirtuleius gracilis n. sp., Parastratocles rosanti n. sp., Parastratocles fuscomarginatus n. sp., Paraprisopus apterus n. sp., Paraprisopus multicolorus n. sp., Agrostia longicerca n. sp., Isagoras similis n. sp., Paragrostia brulei n. sp., Prexaspes globosicaput n. sp., Prexaspes guianensis n. sp., Dinelytron cahureli n. sp., Prisopus clarus n. sp. and Prisopus conocephalus n. sp.. The new genus Paragrostia n. gen. is established for the newly described Paragrostia brulei n. sp. and Paragrostia flavimaculata (Heleodoro, Mendes Rafael, 2017) n. comb. the latter of which is here transferred from Agrostia Redtenbacher, 1906. Fifty-six new combinations are proposed with species transferred to other genera: Bacteria pallidenotata Redtenbacher, 1908, is transferred to Phanocloidea Zompro, 2001 (n. comb.); Bacteria maroniensis Chopard, 1911 is transferred to Phanocles Stål, 1875 (n. comb.); Cladomorphus gibbosus (Chopard, 1911) is transferred to Hirtuleius Stål, 1875 (n. comb.); Stratocles soror Redtenbacher, 1906, Parastratocles lugubris (Redtenbacher, 1906) and Parastratocles cryptochloris (Rehn, 1904) are transferred to Brizoides Redtenbacher, 1906 (n. comb.); Stratocles xanthomela (Olivier, 1792), Stratocles forcipatus Bolívar, 1896 and Stratocles tessulatus (Olivier, 1792) are transferred to Parastratocles (n. comb.); Olcyphides cinereus (Olivier, 1792), Perliodes affinis Redtenbacher, 1906, Perliodes nigrogranulosus Redtenbacher, 1906, Perliodes sexmaculatus Redtenbacher, 1906, Isagoras rugicollis (Gray, 1835), Isagoras sauropterus Rehn, 1947, Brizoides viridipes (Rehn, 1905) and Brizoides graminea Redtenbacher, 1906 are transferred to Agrostia Redtenbacher, 1906 (n. comb.); Agrostia flavimaculata Heleodoro, Mendes Rafael, 2017 is transferred to Paragrostia n. gen. (n. comb.); Isagoras affinis Chopard, 1911, Isagoras chocoensis Hebard, 1921, Isagoras metricus Rehn, 1947 and Isagoras schraderi Rehn, 1947 are transferred to Tenerella Redtenbacher, 1906 (n. comb.); Xerosoma glyptomerion Rehn, 1904 is transferred to Isagoras Stål, 1875 (n. comb.); Isagoras venosus (Burmeister, 1838), Paraphasma paulense Rehn, 1918 and Paraphasma quadratum (Bates, 1865) are transferred to Prexaspes Stål, 1875 (n. comb.); Prexaspes (Prexaspes) cneius (Westwood, 1859) is transferred to Tenerella Redtenbacher, 1906 (n. comb.); Prexaspes lateralis (Fabricius, 1775) is transferred to Paraphasma Redtenbacher, 1906 (n. comb.); Isagoras santara (Westwood, 1859) and Prexaspes olivaceus Chopard, 1911 are transferred to Periphloea Redtenbacher, 1906 (n. comb.); Dinelytron agrion Westwood, 1859 is transferred to Paraprisopus Redtenbacher, 1906 (n. comb.); Anarchodes atrophicus (Pallas, 1772) is transferred to Ignacia Rehn, 1904 (n. comb.); Planudes asperus Bellanger Conle, 2013, Planudes brunni Redtenbacher, 1906, Planudes cortex Hebard, 1919, Planudes crenulipes Rehn, 1904, Planudes funestus Redtenbacher, 1906, Planudes melzeri Piza, 1937, Planudes molorchus (Westwood, 1859), Planudes paxillus (Westwood, 1859), Planudes perillus Stål, 1875, Planudes pygmaeus (Redtenbacher, 1906) and Planudes taeniatus Piza, 1944 are transferred to Isagoras Stål, 1875 (n. comb.); Prisopoides atrobrunneus Heleodoro Rafael, 2020, Prisopoides brunnescens Heleodoro Rafael, 2020, Prisopoides caatingaensis Heleodoro Rafael, 2020 and Prisopoides villosipes (Redtenbacher, 1906) are transferred to Prisopus Peletier de Saint Fargeau Serville, 1828 (n. comb.); Melophasma antillarum (Caudell, 1914), Melophasma brachypterum Conle, Hennemann Gutiérrez, 2011, Melophasma colombianum Conle, Hennemann Gutiérrez, 2011 and Melophasma vermiculare Redtenbacher, 1906 are transferred to Paraprisopus Redtenbacher, 1906 (n. comb.); Prexaspes (Elasia) ambiguus (Stoll, 1813), Prexaspes (Elasia) brevipennis (Burmeister, 1838), Prexaspes (Elasia) pholcus (Westwood, 1859), Prexaspes (Elasia) viridipes Redtenbacher, 1906 and Prexaspes (Elasia) vittata (Piza, 1985) are transferred to Prexaspes Stål, 1875 (n. comb.). Twenty-six new synonymies are established: Perliodes Redtenbacher, 1906 and Chlorophasma Redtenbacher, 1906 are synonymised with Agrostia Redtenbacher, 1906 (n. syn.); Chlorophasma Redtenbacher, 1906 is synonymised with Agrostia Redtenbacher, 1906 (n. syn.); Elasia Redtenbacher, 1906 is synonymised with Prexaspes Stål, 1875 (n. syn.); Prisopoides Heleodoro Rafael, 2020 is synonymised with Prisopus Peletier de Saint Fargeau Serville, 1828 (n. syn.); Melophasma Redtenbacher, 1906 is synonymised with Paraprisopus Redtenbacher, 1906 (n. syn.); Bacteria crassipes Chopard, 1911 is synonymised with Bacteria pallidenotata Redtenbacher, 1908 (n. syn.); Perliodes grisescens Redtenbacher, 1906 and Metriophasma (Metriophasma) pallidum (Chopard, 1911) are synonymised with Agrostia cinerea (Olivier, 1792) (n. syn.); Perliodes nigrogranulosus Redtenbacher, 1906 and Metriophasma (Metriophasma) ocellatum (Piza, 1937) are synonymised with Isagoras rugicollis (Gray, 1835) (n. syn.); Isagoras chopardi Hebard, 1933 is synonymised with Tenerella cneius (Westwood, 1859) (n. syn.); Isagoras proximus Redtenbacher, 1906 is synonymised with Isagoras glyptomerion (Rehn, 1904) (n. syn.); Chlorophasma hyalina Redtenbacher, 1906 is synonymised with Agrostia graminea (Redtenbacher, 1906) (n. syn.); Isagoras nitidus Redtenbacher, 1906 is synonymised with Anisa flavomaculatus (Gray, 1835) (n. syn.); Prexaspes acuticornis (Gray, 1835) is synonymised with Prexaspes servillei (Gray, 1835) (n. syn.); Prexaspes nigromaculatus Chopard, 1911 is synonymised with Periphloea santara (Westwood, 1859) (n. syn.); Prexaspes (Elasia) janus Kirby, 1904 is synonymised with Paraphasma maculatum (Gray, 1835) (n. syn.); Prexaspes dictys (Westwood, 1859) is synonymised with Prexaspes brevipennis (Burmeister, 1838) (n. syn.); Parastratocles aeruginosus Redtenbacher, 1906: 107 is synonymised with Parastratocles forcipatus Bolívar, 1896 (n. syn.); Parastratocles carbonarius (Redtenbacher, 1906: 106) is synonymised with Parastratocles lugubris (Redtenbacher, 1906) (n. syn.); Prisopus spinicollis Burmeister, 1838, Prisopus spiniceps Burmeister, 1838 and Prisopus cornutus Gray, 1835 are synonymised with Prisopus ohrtmanni (Lichtenstein, 1802) (n. syn.); the genus Planudes Stål, 1875 is synonymised with Isagoras Stål, 1875 (n. syn.); Pseudophasma annulipes (Redtenbacher, 1906) is synonymised with Pseudophasma blanchardi (Westwood, 1859) (n. syn.); Ignacia appendiculatum (Kirby, 1904) is synonymised with Anarchodes atrophicus (Pallas, 1772) (n. syn.). Isagoras obscurum Guérin-Méneville, 1838 is shown to have been erroneously synonymised with Isagoras rugicollis (Gray, 1835) and is here re-established as a valid species (rev. stat.). Pseudophasma castaneum (Bates, 1865) is re-established as a valid species here (rev. stat.). Paraprisopus Redtenbacher, 1906 and the entire tribe Paraprisopodini are transferred to Pseudophasmatidae: Pseudophasmatinae (n. comb.). Lectotypes are designated for Perliodes grisescens Redtenbacher, 1906, Isagoras plagiatus Redtenbacher, 1906.Neotypes are designated for Agrostia cinerea (Olivier, 1792), Prexaspes ambiguus (Stoll, 1813), Prisopus horridus (Gray, 1835) and Prisopus sacratus (Olivier, 1792).
Asunto(s)
Escarabajos , Heterópteros , Animales , Guyana Francesa , NeopteraRESUMEN
The new genus Andeocalynda n. gen. is described and the supposed relationships are discussed. The genus is restricted to the mountainous Andean regions of Ecuador and Colombia. Nine new species are described and illustrated: A. aspericollis n. sp., A. banosense n. sp., A. brevicercata n. sp. and A. lojaense n. sp. from Ecuador based on the males only, A. decorata n. sp., A. mutica n. sp., A. densegranuloa n. sp. and A. tuberculata n. sp. from Ecuador as well as A. putumayoense n. sp. from Colombia based on the male and female. The eggs of the two new Ecuadorian species A. decorata n. sp. and A. tenuis n. sp. are described and illustrated. Two species, previously attributed to other genera are transferred to Andeocalynda n. gen., this is: A. carrikeri (Hebard, 1919) from the genus Bacteria Berthold, 1827 (n. comb.) and A. comis (Bates, 1865) from the genus Clonistria Stål, 1875 (n. comb.). As a result, Andeocalynda n. gen. currently includes eleven known species.
Asunto(s)
Neoptera , Distribución Animal , Animales , Colombia , Ecuador , Femenino , MasculinoRESUMEN
The neotropical genus of stick insects, Phantasca Redtenbacher, 1906, was recently revised by Hennemann et al. (2018). However, the females and eggs of several species remain unknown, including all five species recorded from Brazil. The female and egg of Phantasca phantasma (Westwood, 1859) are here described for the first time, based on material available at the Museu de Zoologia da Universidade de São Paulo, Brazil (MZSP). A set of measurements of males is also given, in addition to the measurements of the holotype presented in Hennemann et al. (2018). This complement to the description of P. phantasma is particularly important because it is the type species of the genus (designated by Zompro, 2001), therefore providing additional grounds for future taxonomic decisions involving Phantasca.
Asunto(s)
Distribución Animal , Insectos , Animales , Brasil , Femenino , MasculinoRESUMEN
The areolate Oriental family Heteropterygidae Kirby, 1893 is critically reviewed and the results of the present study contradict the arrangement suggested by Zompro (2004), but in most aspects agree with a molecular study presented by Whiting et al (2003) and a phylogenetic study presented by Bradler (2009). The family is critically discussed and new hypotheses are presented for the phylogeny and intra-familiar relationships, placing the subfamily Dataminae Rehn & Rehn, 1939 as the basalmost clade of Heteropterygidae. The subfamilies Obriminae Brunner v. Wattenwyl, 1893 and Heteropteryginae Kirby, 1893 together represent the sister-group of Dataminae. Arguments and a tree are presented to support this hypothesis. New diagnoses and lists of genera are provided for all three subfamilies contained in Heteropterygidae, along with keys to distinguish between them. The subfamily Obriminae is critically reviewed and the distinction between the three tribes Obrimini Brunner v. Wattenwyl, 1893, Eubulidini Zompro, 2004 and Miroceramiini Zompro, 2004 introduced by Zompro (2004) is shown to be poorly supported. While Obrimini sensu Zompro, 2004 is generally accepted (but now also contains genera that were placed in Eubulidini or Miroceramiini by Zompro (2004)), the tribes Eubulidini and Miroceramiini are not supported. A new arrangement is introduced, which is based on morphological characters neglected or overlooked by Zompro (2004) but were partly discussed by Bradler (2009). The genus Mearnsiana Rehn & Rehn, 1939 is removed from Miroceramiini and transferred to Obrimini. The genera Eubulides Stål, 1877, Heterocopus Redtenbacher, 1906, Theramenes Stål, 1875 and Stenobrimus Redtenbacher, 1906 are removed from Eubulidini and also transferred to Obrimini. Consequently, Eubulidini is synonymised with Obrimini (n. syn.). Miroceramiini is a monotypical tribe and only includes the Wallacean genus Miroceramia Günther, 1934. The new tribe Tisamenini n. trib. is established for the three basal genera Tisamenus Stål, 1875, Ilocano Rehn & Rehn, 1939 and Hoploclonia Stål, 1875 all of which were placed in Eubulidini by Zompro (2004). The latter genus differs from the other two genera by the morphology of the female genitalia, which is unique amongst the entire family. Three generic groups are recognized within Obrimini, the Obrimus-group, Stenobrimus-group and Theramenes-group. Keys are presented to distinguish between the three tribes now contained in the Obriminae, i.e. Obrimini, Tisamenini n. trib. and Miroceramiini. The genus Hennobrimus Conle, 2006 is synonymised with Mearnsiana Rehn & Rehn, 1939, based on the fact that the type-species of both genera are conspecific (n. syn.). Hennobrimus hennemanni Conle, 2006, the type-species of Hennobrimus, and Trachyaretaon manobo Lit & Eusebio, 2005 are synonymised with Mearnsiana bullosa Rehn & Rehn, 1939, the type-species of Mearnsiana (n. syn.). Theramenes dromedarius Stål, 1877 from the Philippines is removed from synonymy with the Wallacean Theramenes olivaceus (Westwood, 1859) and re-established as a valid species (rev. stat.). The subfamily Heteropteryginae Kirby, 1896 is revised at the species-level and a new diagnosis is presented. Keys to the two genera and all 16 known species are provided along with new descriptions, differential diagnoses, lists of examined material, detailed information on the known distributions, measurements and illustrations of the insects and eggs. The intra-subfamiliar and intra-generic relationships are discussed and a cladogram is presented. Heteropteryginae contains two genera: Heteropteryx Gray, 1835 (Type-species: Phasma dilatatum Parkinson, 1798) and Haaniella Kirby, 1896 (Type-species: Phasma (Heteropteryx) muelleri de Haan, 1842). The distribution of this subfamily is restricted to Sundaland with the exception of a single species that is found in Vietnam. All other species are distributed in Borneo, Sumatra, the Mentawai Islands, Singapore, Peninsular Malaysia and Thailand. Heteropteryginae contains the largest and most striking members of the entire family Heteropteryginae, some of which are amongst the heaviest insects known. The subfamily is characterized by apomorphies such as the presence of wings, having a tympanal area (= stridulatory organ) in the basal portion of the alae, straight profemora, strongly shortened tarsi, lack of rough sensory-areas on the prosternum and typically X-shaped micropylar plate of the eggs. The sister-group of Heteropteryginae is represented by the Obriminae, with which it shares a beak-like secondary ovipositor in the females and presence of a medio-apical spine on the area apicalis. Both features are synapomorphies of Heteropteryginae + Obriminae. The genus Haaniella Kirby, 1904 contains 16 known species, five of which are newly described herein. The genus Miniopteryx Zompro, 2004 (Type-species: Haaniella parva Günther, 1944) is synonymised with Haaniella on the basis that the distinguishing feature mentioned in the original description is a character that is frequently found throughout the genus (n. syn.). The type-species H. parva Günther, 1944 is automatically retransferred to Haaniella (rev. stat.). Haaniella aculeata n. sp. from western Sumatra is described from the male. Haaniella macroptera n. sp. from Singapore and the Johor state in southern Peninsular Malaysia is described from both sexes and the eggs. Haaniella gintingi n. sp. from Central Sumatra is described from both sexes and the eggs and Haaniella kerincia n. sp. from Western Sumatra is described from the insects only, the eggs being still unknown. One new species, Haaniella gorochovi n. sp., is the only representative of the genus and subfamily Heteropteryginae known from Vietnam and both sexes as well as the eggs are described. Haaniella erringtoniae (Redtenbacher, 1906) is endemic in Peninsular Malaysia, here removed from synonymy with H. muelleri (de Haan, 1842) and re-established as a valid species (rev. stat.). The Sumatran Haaniella glaber (Redtenbacher, 1906) is removed from synonymy with H. muelleri (Haan, 1842) and re-established as a valid species (rev. stat.). Leocrates glaber Redtenbacher, 1906 and Haaniella muelleri simplex Günther, 1944 are removed from synonymy with H. muelleri (Haan, 1842) (rev. stat.) and synonymised with H. glaber. Haaniella mecheli (Redtenbacher, 1906) and H. rosenbergii (Kaup, 1871) are removed from synonymy with H. muelleri (Haan, 1842) and re-established as valid species (rev. stat.). Haaniella erringtoniae novaeguineae Günther, 1934 and Haaniella muelleri var. b. (Haan, 1842) are synonymized with H. rosenbergii (Kaup, 1871) (n. syn.). The type-species Haaniella muelleri (Haan, 1842) is shown to be a fairly rare species that is restricted to Sumatra. All subsequent records of H. muelleri from outside Sumatra and references to captive breeding of stock originating from Peninsular Malaysia in Europe relate to H. erringtoniae (Redtenbacher, 1906). The previously unknown males and eggs of H. rosenbergii (Kaup, 1871) as well as the previously unknown females and eggs of H. parva Günther, 1944 are described and illustrated for the first time. Based on morphological characters of the insects and eggs three distinct species-groups are recognized within Haaniella. The muelleri species-group contains nine species that are distributed throughout Sumatra, the Mentawei Islands, Singapore and Peninsular Malaysia. These are characterized by the smooth ventral surface of the meso- and metafemora and lemon-shaped eggs which entirely lack the setae seen in the two other species-groups. The grayii species-group comprises four species, two of which are endemic in Borneo, one endemic in Sumatra and the fourth species being the only known representative of the subfamily in Vietnam. These species are characteristic for the prominent pair of spines on the abdominal tergites II-IV of males and long apically multidentate epiproct of females. The echinata species-group contains three exceptionally Bornean species, which are characterized by the long and apically pointed subgenital plate of females, which clearly projects beyond the epiproct, as well as the sub-basal lateral tooth of the anal segment of males. The muelleri species-group is sister to the remainder two species-groups. Heteropteryx Gray, 1853 is a monotypical genus and only contains the type-species H. dilatata (Parkinson, 1798), which is found throughout Peninsular Malaysia, Thailand, Sumatra and Northeastern Borneo. This genus differs from Haaniella by the strongly conically elevated head, which posteriorly projects over the anterior margin of the pronotum, females being bright green or yellow in colour with plain and translucent pink alae and having distinct spines on the abdominal tergites, and males having a strongly shortened mesothorax and dull pink alae. Lectotypes are designated for Haaniella parva Günther, 1944, Heteropteryx echinata Redtenbacher, 1906, Heteropteryx saussurei Redtenbacher, 1906 and Heteropteryx scabra Redtenbacher, 1906 to guarantee stability of these names. Information on the habitats, host-plants, biology, life cycle, parasitism and captive breeding of the species of Heteropteryginae is presented and a list summarising all taxonomic changes presented herein.
Asunto(s)
Insectos/clasificación , Distribución Animal , Estructuras Animales/anatomía & histología , Estructuras Animales/crecimiento & desarrollo , Animales , Asia , Tamaño Corporal , Ecosistema , Europa (Continente) , Femenino , Insectos/anatomía & histología , Insectos/genética , Insectos/crecimiento & desarrollo , Masculino , Tamaño de los Órganos , FilogeniaRESUMEN
The anareolate New World subfamily Cladomorphinae Bradley & Galil, 1977 is reviewed and keys to the six tribes currently included are presented; these are: Cladomorphini Bradley & Galil, 1977, Cladoxerini Karny, 1923, Cranidiini Günther, 1953, Pterinoxylini n. trib., Hesperophasmatini Bradley & Galil, 1977 and Haplopodini Günther, 1953 rev. stat.. New diagnoses are presented for all these tribes and possible relationships within Cladomorphinae are discusssed. Morphology of the genitalia and egg-structures indicate Cladomorphinae as presently treated to be polyphyletic. Two subordinate groups are recognized within present Cladomorphinae, which differ considerably in numerous morphological characters of the insects and eggs. The first group and here regarded as Cladomorphinae sensu stricto is formed by the mostly South American Cladomorphini + Cranidiini + Cladoxerini, while the second group is formed by the predominantly Caribbean Hesperophasmatini + Pterinoxylini n. trib. + Haplopodini. Members of the first group (= Cladomorphini sensu stricto) share the dorsally carinate basitarsus in which the two dorsal carinae are melted with another, increasingly elongated gonapophyses VIII of females which are noticeably longer than gonapophyses IX and lamellate as well as strongly displaced medioventral carina of the profemora. Cranidiini + Cladomorphini share the strongly elongated and filiform gonapophyses VIII and presence of gonoplacs in the females, specialized poculum of males and presence of a median line in the eggs. Cranidiini differs from all other tribes of Cladomorphinae by the entirely unarmed legs of both sexes, distinctly broadened and leaf-like body and prominent longitudinal keel of the mesosternum of females, prominently enlarged poculum and spinulose phallus of males as well as the conspicuous narrowing of the posteromedian gap of the internal micropylar plate of the eggs and noticeably separated median line. Cladomorphini is characteristic for the specialized vomer and poculum of males and distinct opercular structures of the eggs. Certain representatives of Cladomorphini indicate relationships to the "Phanocles-group" of Diapheromerinae: Diapheromerini, hence Cladomorphini as presently treated may be paraphyletic. The exclusively South American Cladoxerini (= Baculini n. syn.) differs from the other two tribes of Cladomorphinae sensu stricto by the distinctly serrate profemora of both sexes and conspicuously shortened antennae of females, which consist of less than 30 segments and are much shorter than the profemora in females. Genital morphology, such as the elongated gonapophyses VIII and presence of gonoplacs in females, as well as the lamellate medioventral carina of the profemora indicate close relation to Cladomorphini. Cranidiini appears to be the sister-taxon of Cladomorphini + Cladoxerini. The tribe Baculini Günther, 1953 is synonymised with Cladoxerini (n. syn.), on the basis that the type-genera of both tribes are congeneric, with Baculum Saussure, 1861 being a junior synonym of Cladoxerus St. Fargeau & Audinet-Serville, 1827 (n. syn.). The genus Tersomia Kirby, 1904 is removed from Hesperophasmatini and transferred to Cladoxerini. Wattenwylia Toledo Piza, 1938 is removed from Pachymorphinae: Gratidiini and transferred to Cladoxerini. A detailed new diagnosis is presented for Cranidiini along with a detailed differentiation and the tribe is shown to be monotypical, only containing its type-genus Cranidium Westwood, 1843. All Caribbean genera subsequently added to Cranidiini are removed and transferred to Haplopodini rev. stat.. The three tribes Hesperophasmatini + Pterinoxylini n. trib. + Haplopodini rev. stat. are closely related and might form a monophyletic clade within Cladomorphinae sensu lato. They differ from Cladomorphinae sensu stricto by the short gonapophyses VIII and reduced gonoplacs of females, unspecialized poculum of males and lack of a micropylar line in the eggs. Haplopodini Günther, 1953 is re-established (rev. stat.) and comprises almost exclusively Caribbean genera previously placed in Hesperophasmatini by Bradley & Galil (1977) or Cranidiini by Zompro, (2004). Aploploides Rehn & Hebard, 1938, Diapherodes Gray, 1835, Haplopus Burmeister, 1838 and Paracranidium Brock, 1998 were misplaced in Cranidiini and are transferred to Haplopodini. On the basis of numerous morphological characters of the insects and eggs Hesperophasmatini is removed from Pseudophasmatidae: Xerosomatinae and re-transferred to its previous position in the subfamily Cladomorphinae sensu lato. A detailed newdiagnosis of Hesperophasmatini is presented, but is only provisional since the true diversity is as yet only fractionally known. The lack of a gula distinguishes Hesperophasmatini from all other tribes. The genus Laciphorus Redtenbacher, 1908 is removed from Hesperophasmatini and transferred to Diapheromeridae: Diapheromerinae: Diapheromerini. The new tribe Pterinoxylini n. trib. is established to contain only the type-genus Pterinoxylus Audinet-Serville, 1838. It is closely related and perhaps the sister taxon of Hesperophasmatini, with which it shares the presence of rough sensory areas on the probasisternum and profurcasternum. It differs from Hesperophasmatini and Haplopodini by the presence of a tympanal region (= stridulatory organ) in the alae of females and the alveolar eggs, which possess peripheral opercular and polar structures. Haplopodini is likely to be the sister group of Pterinoxylini n. trib. + Hesperophasmatini. The tribe Haplopodini rev. stat. is revised at the species level and comprises eight almost exclusively Caribbean genera, four of which are newly described. All eight genera now contained in Haplopodini are described in detail, differentiated from their closest relatives and their relationships and systematic position within Haplopodini are discussed. Keys and maps showing their distributions are presented along with a discussion of the distributional patterns. Detailed descriptions, differential diagnoses, synonymic listings, illustrations, material listings and measurements are given of all 26 currently known species and subspecies of Haplopodini. Four new genera are described within Haplopodini. The monotypical Apteroplopus n. gen. (type-species: Dyme grosse-tuberculata Brunner v. Wattenwyl, 1907) from Honduras is the only taxon of the tribe represented in Central America. It is only known from the male which differs from all other genera by being entirely apterous. Cephaloplopus n. gen. (type-species: Cephaloplopus pulchellus n. sp.) and Parhaplopus n. gen. (type-species: Haplopus cubensis Saussure, 1868) occur only on Hispaniola and Cuba. Both are closely related to Haplopus Burmeister, 1838 but in addition to having noticeably different eggs, both genera differ from Haplopus in several morphological characters. The monotypical Venupherodes n. gen. (type-species: Platycrana venustula Audinet-Serville, 1838) is endemic to Cuba, and in females being apterous resembles the second exclusively Cuban genus Aploploides Rehn & Hebard, 1938. It however differs from all other members of Haplopodini by the laterally expanded mesonotum of females, which overlaps the mesopleurae, as well as the morphology of the eggs. Two species-groups are recognized within Diapherodes Gray, 1835. The gigantea species-group comprises the species from the Lesser Antilles, which are: D. angulata (Fabricius, 1793), Diapherodes dominicae (Rehn & Hebard, 1938), D. gigantea gigantea (Gmélin, 1789), D. gigantea saintluciae n. ssp. and Diapherodes martinicensis Lelong & Langlois, 2005. The three species of the jamaicensis species-group, which are D. achalus (Rehn, 1904), D. jamaicensis (Drury, 1773) and D. laevicollis Redtenbacher, 1906, are restricted to the two Greater Antillean islands Jamaica and Puerto Rico. Haplopus Burmeister, 1838 is the most widely distributed genus being represented on all islands of the Greater Antilles except Jamaica, and also in the Virgin Islands, Bahamas, Florida Keys, Dry Tortugas and as far southwest as the Cayman Islands and Swan Islands. Nine new species and one new subspecies are described: Cephaloplopus alope n. sp. and Haplopus sobrinus n. sp. from Cuba, Cephaloplopus euchlorus n. sp., Cephaloplopus laetus n. sp., Cephaloplopus pulchellus n. sp., Haplopus brachypterus n. sp., Haplopus intermedius n. sp. and Parhaplopus navarroi n. sp. from Hispaniola, Haplopus woodruffi n. sp. from Cayman Brac (Cayman Islands) and Diapherodes gigantea saintluciae n. ssp. from Saint Lucia. Seven of these are described from both sexes but Cephaloplopus alope n. sp. and Haplopus sobrinus n. sp. are only known from the females and Cephaloplopus laetus n. sp. only from the males. The previously unknown males of Diapherodes angulata (Fabricius, 1793), Diapherodes laevicollis Redtenbacher, 1908, Haplopus bicuspidatus de Haan, 1842 and Parhaplopus cubensis (Saussure, 1868) as well as the previously unknown female of Parhaplopus evadne (Westwood, 1859) n. comb. are described and illustrated for the first time. Descriptions and illustrations of the eggs of eleven species are presented: Cephaloplopus euchlorus n. sp., Cephaloplopus pulchellus n. sp., Diapherodes achalus (Rehn, 1904), Diapherodes dominicae (Rehn & Hebard, 1938), Diapherodes gigantea gigantea (Gmélin, 1789), Diapherodes martinicensis Lelong & Langlois, 2005, Diapherodes jamaicensis (Drury, 1773), Haplopus bicuspidatus de Haan, 1842, Haplopus micropterus St. Fargeau & Audinet-Serville, 1825, Parhaplopus navarroi n. sp. and Venupherodes venustula (Audinet-Seville, 1838) n. comb.. Type specimens of the newly described taxa are deposited in the collections of ANSP, NHMUK, IIBZ, FSCA, MCZC, MNHN and USNM. Six species are transferred to other genera (n. comb.): Bacteria grossetuberculata (Brunner v. Wattenwyl, 1907) to Apteroplopus n. gen. (ABSTRACT TRUNCATED)
Asunto(s)
Insectos/anatomía & histología , Insectos/clasificación , Distribución Animal/fisiología , Animales , Femenino , Insectos/fisiología , Masculino , Especificidad de la Especie , Indias OccidentalesRESUMEN
The tribe Phasmatini Gray, 1835 predominantly includes very striking stick insects of remarkable size, most of which are characteristic for their large and often colourful wings. The tribe represents roughly half of the Giant Stick Insects of Wallacea, a subregion in Eastern Indonesia comprising thousands of islands that are separated by deep water straits from the continental islands to the west (Sundaland: Borneo, Java and Sumatra) and East (New Guinea). Within Wallacea the Phasmatini are represented by four genera, namely Anchiale Stål, 1875, Eurycnema Audinet-Serville, 1838, Paracyphocrania Redtenbacher, 1908 and Phasma Lichtenstein, 1796. Currently eight distinct species are known to occur in the Wallacea. Two of these are newly described in the present paper , this is Anchiale buruense sp. n. and Paracyphocrania major sp. n.. Keys are provided for the distinction of the Phasmatini taxa of Wallacea and information on the overall distribution of the four genera is presented, which reveals a derivation of the tribe from the Australian region. The tribe Phasmatini itself is briefly characterized and discussed. Brief characterizations of the four genera are provided along with complete lists of the species currently contained. Anchiale Stål, 1875 is represented in the Wallacea by two species. A. maculata (Olivier, 1792) is widely distributed throughout Wallacea except for Sulawesi and shows considerable intraspecific variability. Both sexes and the eggs are illustrated and the eggs formally described for the first time. The synonymies of A. maculata are clarified. A. stolli Sharp, 1898 and A. confusa Sharp, 1898 from New Britain and the Solomon Islands are shown to be errorneous synonyms of A. maculata and are here re-established as valid species (stat. rev.). A. caesarea Redtenbacher, 1908 is removed from the genus Acrophylla Gray, 1835 and transferred to Anchiale (comb. n.). The new species A. buruense sp. n. from the island of Buru (Maluku Islands) is described and illustrated based on both sexes and the egg. Eurycnema Audinet-Serville, 1838 has two species within the boundaries of Wallacea. E. nigrospinosa Redtenbacher, 1908 is only known from the Kei Islands in the southeastern portion of Wallacea and otherwise found on New Guinea. Both sexes and the eggs are illustrated. E. versirubra Audinet-Serville, 1838 is found on some of the Lesser Sunda Islands and otherwise distributed throughout Java, Sumatra and SE-Borneo. The records from Java, Sumatra and southeast Borneo are estimated to be artificial. Females of this species are dimorph and occur in two distinct colour morphs, which differ by the distinct colouration of the undersides of the tegmina and alae. E. versirubra colour morph versirubra has two forms, the insects either being bright green or dull yellow with the ventral surfaces of the tegmina and alae bright red. E. versirubra colour morph versifasciata, has the ventral surfaces of the tegmina and alae yellow, the insects themselves being pale bluish green to turquoise in colour. Only the bright green form appears to be natural with the other colour-forms caused by parthenogenetic reproduction in captivity. Both sexes, the eggs and the two aforementioned colour-morphs are illustrated. Paracyphocrania Redtenbacher, 1908 contains two known species and is endemic to Sulawesi and the nearby island of Peleng. The new species Paracyphocrania major sp. n. from Peleng is described and illustrated based on the female and egg. The male remains as yet unknown. The previously unknown male of P. lativentris Redtenbacher, 1908 is described for the first time. Both sexes and the eggs are illustrated. Vasilissa tecticollis Redtenbacher, 1908 is shown to have been misinterpreted previously, here removed from Paracyphocrania and shown to be a synonym of the Australian Tropidoderus rhodomus McCoy, 1882 (syn. n.). Hence, the type-locality of V. tecticollis originally given as "Philippines" is definitely wrong, which proves the tribe Phasmatini is not represented in the Philippines at all. Phasma Lichtenstein, 1796 is represented in Wallacea with two distinct species and has one further species on New Guinea. A key is provided to distinguish between the three known species of Phasma. Ph. gigas (Linnaeus, 1758) is widely distributed throughout Wallacea and found on almost all major islands, although the historic records from Sulawesi deserve evaluation. Ph. gigas is most certainly not present on New Guinea, with all New Guinean records actually referring to Ph. reinwardtii (de Haan, 1842). Ph. gigas exhibits remarkable intraspecific variability in the colouration and certain morphological features of the insects, with several of these variations appearing to be peculiar to certain localities and islands. The range of variation as well as both sexes and the eggs are illustrated. Papuanoidea straleni Werner, 1930 from New Guinea has erroneously been synonymised with Ph. gigas and is here shown to be a synonym of Ph. reinwardtii (de Haan, 1842) (syn. n.). The second Wallacean representative of the genus, Ph. marosense Hennemann, 1998, is endemic to Sulawesi. The previously unknown male is described for the first time and both sexes and the eggs are illustrated. Holotypes of the two newly described species, Anchiale buruense sp. n. and Paracyphocrania major sp. n., are deposited in the State Zoological Collections Munich, Germany (ZSMC).
Asunto(s)
Insectos/clasificación , Distribución Animal , Estructuras Animales/anatomía & histología , Estructuras Animales/crecimiento & desarrollo , Animales , Australia , Tamaño Corporal , Borneo , Ecosistema , Femenino , Indonesia , Insectos/anatomía & histología , Insectos/crecimiento & desarrollo , Nueva Guinea , Tamaño de los Órganos , Reino UnidoRESUMEN
The defensive secretion of Parectatosoma mocquerysi, a walkingstick insect from Madagascar, was determined to contain glucose, water, and a new monoterpene, parectadial, (4S)-(3-oxoprop-1-en-2-yl)cyclohex-1-enecarbaldehyde. Here, we describe the elucidation of parectadial's molecular structure and absolute configuration via microsample NMR technology, GC-MS, CD, chiral GC-FID, and synthesis from enantiomerically pure (S)- and (R)-perillaldehyde. This work demonstrates the value of walkingstick insects as sources of new bioactive compounds and provides an analytical framework for identifying such substances.