Research History of Phasmatodea

The three first described species of Phasmatodea:

1. Gryllus (Mantis) gigas LINNÉ, 1758 [Valid name: Phasma gigas (LINNÉ, 1758)]

2. Gryllus (Mantis) siccifolius LINNÉ, 1758 [Valid name: Phyllium (Phyllium) siccifolium (LINNÉ, 1758)]

3. Gryllus (Mantis) phthisicus LINNÉ, 1758 [Valid name: Pseudophasma phthisicum (LINNÉ, 1758)]

1758-1830

Carl von Linné

Carl von Linné

The research and bibliographical history of the Phasmatodea reaches as far back as to the mid-18th century. The first three species (see above) were described by the famous Swedish naturalist and inventor of binary nomenclature Carl von LINNÉ (lat. Carolus Linnaeus, 1707-1778) in the 10th edition of his "Systema Naturae" of 1758, where he combined all Orthopterous insects in the genus Gryllus. Subsequently, LINNÉ separated the current Phasmatodea and Mantodea from Gryllus and united them in the genus Mantis, which was overtaken by the Denish zoologist Johann Christian FABRICIUS (1745-1808) in his "Entomologica Systematica” of 1793. Caspar STOLL (ca. 1725-1791) and Anton August Heinrich LICHTENSTEIN (1753-1816) discussed the status of their own family for the Phasmatodea, which was first established in the "Verzeichnis der Käfer Preußens" by Johann Karl Wilhelm ILLIGER in 1798 and the 1798 work of FABRICIUS. In the 1st edition of 1788 of his great work "Représentation des Spectres ou Phasmes, des Mantes, des Sauturelles, des Grillons, des Criquets et des Blattes des quatre Parties du Monde" STOLL provided beautiful hand-coloured plates showing several species of Phasmatodea, but he did not use the binominal system of nomenclature proposed by LINNÉ (1758) and thus left his species without scientific names. Subsequently, several Latinized names for the species illustrated by STOLL were proposed by Guillaume Antoine OLIVIER in his "Encyclopédie Méthodique, Histoire Naturelle, Insectes" of 1792. Not being aware of OLIVIER's 1792 publication, LICHTENSTEIN (1796 & 1802) proposed Latinized names for STOLL's species as well, most of which are synonymous with the names introduced by OLIVIER.

1830-1900

Plate by Westwood, J.O.

Plate by Westwood, J.O.

It was not until 1833 that the English zoologist George Robert GRAY (1808-1872) published the first book that exceptionally dealt with Phasmatodea, titled "The Entomology of Australia", in a series of monographs. The monograph of the genus Phasma.". In 1835, GRAY published his "Synopsis of the species of insects belonging to the family of Phasmidae", the first monograph of the order which already listed 134 described species and a great number of new generic names. Two updated monographs of the order were published shortly thereafter in 1838 by the well-known German zoologist Hermann Carl Conrad BURMEISTER in his series "Handbuch der Entomologie" and the French entomologist Jean Guillaume AUDINET-SERVILLE in his extensive and famous "Histoire naturelle des insectes". In his 1842 work "Bijdragen to de kennis der Orthoptera" in the book "Verhandelingen over de Natuurlijke Geschiedenis der Nederlansche Overzeesche Bezittingen" Willy De HAAN presented detailed descriptions of numerous new species mainly from the Oriental region, most of which he illustrated on six beautiful hand-coloured plates. The author interpreted all generic names previously established as subgenera of Phasma LICHTENSTEIN, 1796.

The most important and extensive publication of the 19th century dealing with Phasmatodea is undoubtedly the well-known "Catalogue of Orthopterous Insects in the Collection of the British Museum" of 1859 by the famous English entomologist and archaeologist John Obadiah WESTWOOD (1805-1893). This monograph already listed 39 genera and 471 species in two main groups, that were divided into adults winged or wingless. Many species were described for the first time and especially the very detailed descriptions and excellent 48 black and white plates that show a wide range of species make it a very important and invaluable work for the taxonomy of phasmids. Between 1859 and 1870 the Swiss zoologist and mineralogist Henri Louis Frédéric de SAUSSURE published several papers that described another fairly large number of new genera and species and another 52 new species were described by the famous English naturalist and explorer Henry Walter BATES in a single publication in 1865. All of these works followed the classification proposed by BURMEISTER (1838).

The Swedish entomologist Carl STÅL (1833-1878) established a new classification of the Phasmatodea and defined important new criteria for a more natural system of the order, which he revised two times within only one year. The most important and extensive of his three works is his "Recensio Orthopterorum" of 1875, which already listed 95 genera of Phasmatodea. This particular paper was not only of great importance for the systematization of the order because it first recognised and introduced the "areola apicalis" and "segmentum medianum", which are still of great importance for the phasmatodean phylogeny, but also for the very detailed descriptions of genera and species. STÅL was the first to divide the Phasmatodea into two large groups according to whether the insects possess an "areola apicalis tibiarum" or not, but did not name these groups.

In 1893 the well-known Swiss naturalist Karl Friedrich BRUNNER v. WATTENWYL (1823-1914) revised the classification of the order yet again and established a new system, that however was not capable of resisting the strongly growing number of taxa and became outdated quite rapidly.

Further important authors that have described various new taxa of Phasmatodea in the late 19th century include: Ignacio BOLÍVAR (1895, 1896, 1897), Ermanno GIGLIO-TOS (1893, 1894, 1895, 1897, 1898), Ferdinand KARSCH (1896, 1898), Johann Jakob KAUP (1866, 1871), William Forsell KIRBY (1844, 1889, 1891, 1896) and James WOOD-MASON (1873, 1875, 1876, 1877, 1878, 1879).

1900-1950

Plate by Redtenbacher, J.

Plate by Redtenbacher, J.

In addition to some earlier papers that described various new Phasmatodea, the English entomologist William Forsell KIRBY (1844-1912) published his 1st part of "A synonymic catalogue of Orthoptera" in 1904. This paper is very important for the taxonomy of the order because it included numerous new names, taxonomic changes, reference to type specimens deposited in the Natural History Museum London (NHMUK) and the designation of a type-species for each genus. Unfortunately, however, KIRBY's 1904 catalogue did not receive the recognition it warranted by subsequent workers, which has caused considerable confusion throughout the taxonomy of the Phasmatodea.

At the beginning of the 20th century, Karl Friedrich BRUNNER v. WATTENWYL and the Austrian entomologist Josef REDTENBACHER (1856-1926) started with their extensive research for a monographic work of the order. The result of this cooperation was the outstanding monograph, published in three parts between (1906, 1907 and 1908) entitled "Die Insektenfamilie der Phasmiden", which is in Latin with a little German. The monograph recognized 1899 species and still is the most extensive, monographical study of the order. The 27 excellent black and white plates include detailed drawings of almost all the genera known at that time. The main shortcoming of this outstanding work, however, is, that KIRBY's taxonomically important 1904 catalogue was completely ignored by BRUNNER v. WATTENWYL & REDTENBACHER (1906-1908), a fact that has led to numerous inaccuracies and misinterpretations of genera. The authors named the two subgroups recognised by STÅL (1875) "Areolatae" and "Anareolatae" and the classification presented in the monograph generally arranged several equally placed "tribes" next to the other, and mostly distinguished between these by the length relations of the metanotum and median segment.

The first half of the 20th century has seen very few revisions, but a great number of papers of a variable extent that either deal with defined geographical regions and local faunas, but none of which is neither geographically nor taxonomically complete, study the material from particular collections or expeditions or describe single new taxa. A considerable number of new taxa have been described during this period and descriptions and localities have become increasingly more accurate and detailed. Some authors that have published numerous new taxa are e.g.: Ignacio BOLÍVAR (1912, 1919, 1922), J. CARL (1913, 1915), Andrew Nelson CAUDELL (1902, 1903, 1904, 1905, 1906, 1908, 1913, 1914), Lucien CHOPARD (1911, 1938, 1955), Heinrich Wolfgang Ludwig DOHRN (1910), Ermanno GIGLIO-TOS (1910, 1912, 1914), Robert Walter Campbell SHELFORD (1913), Tokuichi SHIRAKI (1911), Yngve SJÖSTEDT (1909, 1924) and Salvador de TOLEDO PIZA Jun. (1936, 1938, 1939, 1943, 1944, 1946). The German entomologist Klaus GÜNTHER (1907-1975) was the most prolific author of the early 20th century and published numerous papers on various faunas including Borneo, New Guinea, the Pacific Ocean or Sulawesi that described a large number of new genera and species. The most extensive of his papers is the 1928 publication "Die Phasmoiden der Deutschen Kaiserin Augusta-Fluss-Expedition 1912/13", which described two new genera, 27 new species and five new subspecies from Papua New Guinea. The two American entomologists Morgan HEBARD (1887-1946) and James Abram Garfield REHN (1881-1965) deserve special mention for publishing numerous papers about Phasmatodea either on their own or together. Most of these deal with taxa from North, Central and South America or the Caribbean but REHN also published several papers that include African and Oriental taxa (e.g. 1904, 1906, 1912). REHN published two papers together with his brother John J. W. REHN, one in 1933 that dealt with leaf insects of the genus Phyllium, and a study of the tribe Obrimini of the Philippine Islands in 1938, which is the only work dealing comprehensively with any larger taxa.

Except for some nomenclatorial changes suggested by the Austrian entomologist Heinrich Hugo KARNY in 1923, the classification of the Phasmatodea did not see any substantial changes until the mid of the 20th century. The changes suggested by KARNY are mainly restricted to uniting the equally placed "tribes" established by BRUNNER v. WATTENWYL & REDTENBACHER (1906-1908) into a few families and renaming the "Areolatae" and "Anareolatae" as Phyllidae and Phasmatidae. Subsequent authors generally followed KARNY's classification (e.g. HANDLIRSCH, 1930; BRUES & MELANDER, 1932; CHOPARD, 1949).

1950 - 1980

It was not until 1953 that Klaus GÜNTHER published his important paper entitled "Über die taxonomische Gliederung und die geographische Verbreitung der Insektenordnung der Phasmatodea", which presented a new classification of the Phasmatodea and provided characterizations and keys to all families and subfamilies. GÜNTHER subdivided his subfamilies into tribes on which he commented but did not provide any further determinating keys. Since BRUNNER v. WATTENWYL & REDTENBACHER's (1906-1908) monograph GÜNTHER's work is the first fundamental step towards a new and more natural classification of the order. Further, more detailed keys were published by Max BEIER in 1957 and 1958.

The next paper of importance for the classification of Phasmatodea, although generally a slightly updated translation of GÜNTHER's work is "The Taxonomic Arrangement of the Phasmatodea with Keys to the Subfamilies and tribes" of 1977 by John C. BRADLEY & Bella S. GALIL. This approach to classification was widely used until quite recently. Based on the characteristics summarized by GÜNTHER, the publication also provided keys to all the tribes and as a result, keys to all 6 families, 17 subfamilies and 35 tribes of Phasmatodea were provided for the first time. This author’s work was exceptionally based on literary sources and as BRADLEY died whilst working on the translation, the work includes numerous inaccuracies, errors and misspellings and is still far away from a natural system of the order. KEVAN (in PARKER, 1982) suggested several taxonomic changes mostly without apparent justification, which however have largely remained ignored by subsequent authors.

Although inaccurate in various aspects the most remarkable use of the classification proposed by BRADLEY & GALIL (1977) was to provide a basic tool for identification to forthcoming entomologists. This resulted in the first enthusiasts began to work with Phasmatodea seriously in the early 1980's. Except for the aforementioned work on the phasmatodean classification, there were only very few descriptions of new taxa and few taxonomic efforts.

After 1980

The first author to publish on the taxonomy and ootaxonomy ( see below) of Phasmatodea and who described several new species mostly based on captive-reared material between 1983 and 1992 is the Austrian worker Burghard HAUSLEITHNER. A large number of papers dealing with the biology and habits of various phasmids have been published by Ulf CARLBERG (Denmark), who from 1983 to 1994 also published a series of eight research papers that list the bibliography of Phasmatodea entitled "Bibliography of Phasmida (Insecta). I to VIII". Between 1992 and 2008 the English worker Philip E. Bragg did extensive research on the phasmid fauna of Borneo and published a large number of papers that dealt with their taxonomy and in total described over 50 new species and several new genera. His massive 2001 book "Phasmids of Borneo" described another 19 new species, two subspecies and one new genus. For the first time since the monograph of BRUNBER v. WATTENWYL & REDTENBACHER (1906-1908) BRAGG’s book includes revisions of certain genera and taxa in a monographical order. Most other publications of the late 20th century dealt with single taxa, the study of local faunas or provided important catalogues of type-material of Phasmatodea from various museum collections around the world. The most prolific authors were: Paul D. BROCK (England), Baoling CAI (China), Ariel CAMOUSSEIGHT (Chile), Shuchun CHEN (China), Oskar V. CONLE (Germany), Yunheng HE (China), Frank H. HENNEMANN (Germany), Shengli LIU (China), Francis SEOW-CHOEN (Singapore), Vernon R. VICKERY (Canada), J. J. Wang (China) and Oliver ZOMPRO (Germany). The only noteworthy scientific books on Phasmatodea in the 1990s are Paul D. BROCK’s well-known "Stick-Insects of Britain, Europe and the Mediterranean" of 1991 and "Stick and leaf insects of Peninsular Malaysia and Singapore" of 1999 and the beautifully illustrated "The Stick Insects of New Zealand" of 1991 by John T. SALMON. A considerable and, if compared to the preceding 50 years, a remarkably growing number of new taxa have been described by the mentioned authors in the late 20th century.

Since the beginning of the 21st century, an increasing number of more extensive taxonomic and phylogenetic papers but also books about Phasmatodea have been published and the number of new taxa that are being described frequently increased. Paul D. BROCK has done extensive work, especially on the fauna of Australia and described a good number of new species and genera, some of them in cooperation with Jack HASENPUSCH. Their book "The Complete Field Guide to Stick and Leaf Insects of Australia" was published in 2009. Other publications of Paul D. BROCK have dealt with the Phasmatodea of South Africa and New Zealand. The prolific Singaporean worker Francis SEOW-CHOEN has specialized in the Phasmatodea of Sundaland and has recently published three beautifully illustrated books: "A Taxonomic Guide to the Stick Insects of Borneo" in 2016, "A Taxonomic Guide to the Stick Insects of Borneo. Volume II" in 2017 and "A taxonomic guide to the stick insects of Singapore" in 2017. The two very prolific Chinese workers Shuchun CHEN and Yungeng HE published their extensive monographic book "Phasmatodea of China" in 2007, which described 76 new species. Their work on the Chinese fauna is continued by George Wai Chun HO, who has published numerous research papers on Phasmatodea from China and Vietnam and the 2013 book "Stick Insects of Hong Kong". Since 2013 extensive research on the phasmid fauna of Vietnam has been carried out by two Belgian entomologists Joachim BRESSEEL and Jérome CONSTANT, who have already published several papers that describe new taxa. The French author Nicolas CLIQUENNOIS has published a good number of papers dealing with the taxonomy and phylogeny of the phasmids of the Mascarenes and Madagascar, which have resulted in the descriptions of several new taxa. Sparked by a comprehensive study on leaf insects of the genus Phyllium by HENNEMANN et al. (2009), the American entomologist Royce T. CUMMING has published several papers on the family Phylliidae that have described several new species and even a new genus. In addition to various major papers on Oriental Phasmatodea (e.g. 2008, 2009, 2015, 2016) the two prolific German authors Frank H. HENNEMANN and Oskar V. CONLE are conducting extensive research on the Phasmatodea of the New World (North, Central and South America and the Caribbean) in a continuous series of research papers entitled "Studies on Neotropical Phasmatodea" started in 2005 (e.g. 2008, 2009, 2016, 2018). Books by these authors include "The Stick Insects of Colombia" of 2011 which described 74 new species and four new genera.

In addition to studies on the biology, systematics and taxonomy of Phasmatodea, several workers have recently begun to seriously study the biogeography and molecular phylogeny of the order. First publications have mainly focused on biogeographically restricted taxa such as the Mediterranean Phasmatodea (e.g. SCALI et al., 2003, 2012, 2013), the New Zealand stick insects (e.g. TREWICK et al., 2005, 2008; BUCKLEY et al., 2008, 2010) or the genus Timema in the Southern United States (LAW & CRESPI, 2002a,b; SCHWANDER et al., 2011). Still, the classification of the Phasmatodea is far away from a natural phylogeny and the early evolutionary history, such as the true relationships between subfamilies and other higher lever taxa is still obscure. However, our knowledge has been dramatically improved with several recent phylogenetic analyses of molecular (WHITING et al., 2003; BUCKLEY et al., 2009, 2010; KÔMOTO, et al., 2011, 2012) and morphological (BRADLER, 2009; Friedmann et al., 2012) datasets. In general, relationships between stick insects are more congruent with geographical distributions rather than with traditional classifications, but still, a much denser taxon sampling will be necessary for reconstructing a proper phylogeny of the Phasmatodea.

The eggs of Phasmatodea – history of the ootaxonomy.

Plate by Kaup.

Plate by Kaup.

The first phasmid egg was illustrated in 1798 by John PARKINSON alongside the very detailed original description of Heteropteryx dilatata in a paper entitled "Description of the Phasma Dilatatum".

It was not until 1871 when the German entomologist Johann Jakob KAUP (1803-1873) published "Über die Eier der Phasmiden", which was the first paper exceptionally dealing with the eggs of Phasmatodea and provided descriptions and illustrations of the eggs of 26 different species. KAUP wrote: "Obgleich ich bis jetzt nur eine verschwindend kleine Zahl von Eiern untersuchen konnte, so glaube ich doch den Schluss wagen zu dürfen, dass verwandte Species auch ähnliche Eier besitzen.... [Although I have so far examined only a very small number of eggs I feel able to claim that related species possess similar eggs...]" and concluded that the eggs of Phasmatodea might be an important character for the distinction of species. Furthermore, KAUP stated: "später vielleicht die Arten durch die Eier schneller unterscheiden lernt als durch die Tiere selbst. […in the future the distinction of species may prove to be easier and reliable examining their eggs instead of the insects themselves.]". But, since only very limited egg material was available for examination during the early 20th century, KAUP's brilliant investigations and suggestions were largely ignored. It was not until the second half of the 20th century that live material of different species was imported into Europe for breeding purposes and eggs of a growing number of species became available for examination. After remaining largely ignored for almost 100 years, this fortunate development in Europe has caused KAUP's hypotheses to be rediscovered and first used as a basis for further studies in the eggs of phasmatodeans. Authors, who have since then done more detailed studies on the eggs of Phasmatodea and have used the ootaxonomy for the systematization and differentiation of taxa are e.g. John T. C. CLARK-SELLICK (= CLARK), Burghard HAUSLEITHNER, Oskar V. CONLE, Frank H. HENNEMANN and Oliver ZOMPRO.

A detailed terminology for the description of the phasmid egg morphology was established by CLARK (1976). Subsequently, CLARK-SELLICK published three extensive papers (1997-1998) that dealt with the eggs of Phasmatodea and discussed their importance for the classification and taxonomy of the order. One of the two 1997 papers included a key and illustrations of the eggs of 131 genera. In his 1998 paper, the author discussed the shape and structure of the micropylar plate and its use for the systematics of the order. Throughout his papers, CLARK-SELLICK mentioned many interesting observations and provided numerous hypotheses but never drew any taxonomic consequences. The important works of CLARK-SELLICK published between 1976 and 1998 have greatly improved our knowledge of the ootaxonomy of Phasmatodea, but still, the eggs of many genera and species remain unknown.

Literature

The following list of references is not complete. It does not list all publications mentioned in the text above and is only a selection of the most important papers and books. Full citings and reference lists of all authors mentioned above can be found in the online Phasmida Species File

  • AUDINET-SERVILLE, J.G. (1838): Histoire Naturelle des Insectes. Orthoptères. Libraire Encyclopédique de Roret, Paris, 18. 776 pp.
  • BATES, H. W. (1865): Descriptions of fifty-two new species of Phasmidae from the collection of Mr. W. Wilson Saunders, with remarks on the family. Transactions of the Linnean Society of London, 25(1): 321-359.
  • BEIER, M. (1957): Orthopteroidea. Ordnung: CheleutopteraCrampton
  • BRADLER, S. (2009): Phylogenie der Stab- und Gespentschrecken (Phasmatodea). Species, Phylogeny and Evolution, 2: 3-19.
  • BRADLEY, J.C. & GALIL, B.S. (1977): The taxonomic arrangement of the Phasmatodea with keys to the subfamilies and tribes. Proceedings of the Entomological Society of Washington, 79(2): 176-208.
  • BRAGG, P.E. (2001): Phasmids of Borneo. Natural History Publications (Borneo), Kota Kinabalu. 772 pp.
  • BROCK, P.D. (1999): Stick and leaf insects of Peninsular Malaysia and Singapore. Malaysian Nature Society, Kuala Lumpur. 222 pp.
  • BROCK, P.D. & HASENPUSCH, J. (2009): The Complete Field Guide to Stick and Leaf Insects of Australia. CSIRO Publishing, Collingwood, Victoria. 204 pp.
  • BRUNNER v. WATTENWYL, C. (1893): Révision du système des Orthoptères et description des espèces rapportées par M. Leonardo Fea de Birmanie. Annali de Museo Civico di Storia Naturale Giacomo Doria, Genova, (2)13(33): 76-101 & plates 2-4.
  • BRUNNER v. WATTENWYL, C. & REDTENBACHER, J. (1906-1908): Die Insektenfamilie der Phasmiden, Parts 1-3. Verlag Wilhelm Engelmann, Leipzig. 589 pp., 37 plates.
  • BURMEISTER, H. (1838): Handbuch der Entomologie, II. Berlin, pp. 553-589.
  • BUCKLEY, T.R., ATTANAYAKE, D., PARK, D., RAVINDRAN, S., JEWELL, T.R. & NORMARK, B.B. (2008): Investigating hybridization in the parthenogenetic New Zealand stick insect Acanthoxyla (Phasmatodea) using single-copy nuclear loci. Molecular Phylogenetics and Evolution, 48: 335-349.
  • BUCKLEY, T.R., ATTANAYAKE, D., & BRADLER, S. (2009): Extreme convergence in stick insect evolution: phylogenetic placement oft he Lord Howe Island tree lobster. Proceedings oft he Royal Society B., 276: 1055-1062.
  • BUCKLEY, T.R., ATTANAYAKE, D., NYLANDER, J.A.A. & BRADLER; S. (2010): The phylogenetic placement and biogeographical origins oft he New Zealand stick insects (Phasmatodea). Systematic Entomology, 35: 207-225.
  • CAMOUSSEIGHT, A. (1995): Revision taxonomia del genero Agathemera (Phasmatodea: Pseudophasmatidae). Revista Chilena de Entomología, 22: 35-53.
  • CARL, J. (1913): Phasmides nouveaux ou peu connus du Muséum de Genève. Revue Suisse de Zoologie, 21(1): 1-55, plate 1.
  • CARL, J. (1915): Phasmiden von Neu-Caledonien und den Loyalty-Inseln. In: SARASIN, F. & ROUX, J. [Eds.]: Forschungen in Neu-Caledonia. Zoologie, (2)2(9): 173-194.
  • CARLBERG, U. (1983): Bibliography of Phasmida (Insecta). I. 19760-79. Spixiana, 6(1): 27-43.
  • CARLBERG, U. (1985): Bibliography of Phasmida (Insecta) II (1960-69). Beiträge zur Entomologie, Berlin, 35(1): 3-12.
  • CARLBERG, U. (1986): Bibliography of Phasmida (Insecta) III: 1950-59. Beiträge zur Entomologie, Berlin, 36(2): 255-260.
  • CARLBERG, U. (1987a): Bibliography of Phasmida (Insecta) - IV. 1940-49. Beiträge zur Entomologie, Berlin, 37(1): 197-202.
  • CARLBERG, U. (1987b): Bibliography of Phasmida (Insecta). VI. 1980-84. Spixiana, 10(2): 147-156.
  • CARLBERG, U. (1988): Bibliography of Phasmida (Insecta) – V (1930-39). Beiträge zur Entomologie, Berlin, 38(1): 277-287.
  • CARLBERG, U. (1994a): Bibliography of Phasmida (Insecta). VII. 1985-1989. Spixiana, 17(2): 179-191.
  • CARLBERG, U. (1994b): Bibliography of Phasmida (Insecta) - VIII. Supplement I (1930-1984). Beiträge zur Entomologie, Berlin, 44(1): 234-250.
  • CHEN, S.C. & HE, Y. (2007): Phasmatodea of China. China Forestry Publishing House, 476 pp.
  • CLARK-SELLICK, J.T. (1997a): The range of capsule morphology within the Phasmatodea and its relevance to the taxonomy of the order. Italian Journal of Zoology, 64: 97-104.
  • CLARK-SELLICK, J.T. (1997b): Descriptive terminology of the phasmid egg capsule, with an extended key to the phasmid genera based on egg structure. Systematic Entomology, 22: 97-122.
  • CLARK-SELLICK, J.T. (1998): The micropylar plate of the eggs of Phasmida, with a survey of the range of plate form within the order. Systematic Entomology, 23: 203-228.
  • CONLE, O.V., HENNEMANN, F.H. & GUTIÉRREZ, Y. (2013): The Stick Insects of Colombia. A catalogue and bibliography with the descriptions of four new genera and 74 new species. Books on Demand GmbH, Norderstedt. 406 pp.
  • CONLE, O.V., HENNEMANN, F.H. & PEREZ-GELABERT, D. (2008): Studies on neotropical Phasmatodea II: Revision of the genus Malacomorpha Rehn, 1906, with the descriptions of seven new species (Phasmatodea: Pseudophasmatidae: Pseudophasmatinae). Zootaxa, 1748: 1-64.
  • DOHRN, H. (1910): Beitrag zur Kenntnis der Phasmiden. Entomologische Zeitung Stettin, 71: 397-414.
  • FABRICIUS, J.C. (1793): Entomologia systematica emendata et aucta. Orthoptera. Vol. 2. Hafniae [Copenhagen], 519 pp.
  • FABRICIUS, J.C. (1798): Supplementum Entomologiae Systematicae. Havniae [Copenhagen], 572 pp.
  • GIGLIO-TOS, E. (1910): Fasmidi esotici del R. Museo zoologico di Torino e del Museo civico di Storia naturale di Genova. Bolletino dei Musei di Zoologa ed Anatomia comparara della Royale Università di Torino, 25(625): 1-57.
  • GRAY, G.R. (1833): The Entomology of Australia, Part 1. The monograph of the genus Phasma. London, 28 pp., plates 1-8.
  • GRAY, G.R. (1835): Synopsis of the species of insects belonging to the family of Phasmidae. Longman, Rees, Orme, Green & Longman, London. 48 pp.
  • GÜNTHER, K. (1929): Die Phasmoiden der Deutschen kaiserin Augusta-Fluß-Expedition 1912/13. Ein beitrag zur Kenntnis der Phasmoidenfauna Neuguineas. Mitteilungen aus dem Zoologischen Museum in Berlin, 14: 60-746, plates 1-6.
  • GÜNTHER, K. (1953): Über die taxonomische Gliederung und die geographische Verbreitung der Insektenordnung der Phasmatodea. Beiträge zur Entomologie, Berlin, 3: 541-563.
  • HAAN, W. de (1842): Bijdragen tot de Kennis der Orthoptera. Verhandelingen over de natuurlijke Geschiedenis der Nederlandsche overzeesche Bezittingen. In: TEMMINCK, C.J. [Ed.]: Verhandelingen Zoologie, Vol. 2: 95-138.
  • HENNEMANN, F.H. & CONLE, O.V. (2008): Revision of Riental Phasmatodea: the tribe Pharnaciini Günther, 1953, including the description oft he world’s longest insect, and a survey oft he family Phasmatidae Gray, 1835 with keys tot he subfamilies and tribes (Phasmatodea: „Anareolatae“: Phasmatidae). Zootaxa, 1906: 1-316.
  • HENNEMANN, F.H., CONLE, O.V., BELLANGER, Y., LELONG, P. & JOURDAN, T. (2018): Studies on Neotropical Phasmatodea XVII: Revision of Phantasca Redtenbacher, 1906, with the descriptions of sex new species (Phasmatodea: Diapheromeridae: Diapheromerinae). European Journal of Taxonomy, 435: 1-62.
  • HENNEMANN, F.H., CONLE, O.V., BROCK, P.D. & SEOW-CHOEN, F. (2016): Revision of the Oriental subfamily Heteropteryginae Kirby, 1896, with a re-arrangement oft he family Heteropterygiade and the descriptions of five new species of Haaniella Kirby, 1904 (Phasmatodea: Areolatae: Heteropterygidae). Zootaxa, 4159(1): 1-129.
  • HENNEMANN, F.H., CONLE, O.V., GOTTARDO, M. & BRESSEEL, J. (2009): On certain species of the genus Phyllium Illiger, 1798, with proposals for an intra-generic systematization and the descriptions of five new species from the Philippines and Palawan (Phasmatodea: Phylliidae: Phylliinae: Phylliini). Zootaxa, 2322: 1-83.
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