Erforschungsgeschichte der Phasmatodea

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1758- ca. 1830

Carl von LINNÉThe bibliographical history of the Phasmatodea reaches as far back as to the mid of the 18th century. The first species were described by the famous Swedish naturalist Carl von LINNÉ (1707-1778) in the tenth edition of his well known “Systema Naturae” of 1758 where he combined all Orthopterous insects in the genus Gryllus. Later LINNÉ separated the Phasmatodea and Mantodea from Gryllus and united them in the genus Mantis, which was overtaken by Johann Christian FABRICIUS (1745-1808) in his “Entomologica Systematica” of 1793. Caspar STOLL („Représentation des Spectres ou Phasmes, des Mantes, des Sauturelles, des Grillons, des Criquets et des Blattes des quatre Parties du Monde“, 1789-1815) and A.A.H. LICHTENSTEIN (1802) discussed the status of an own family for the Phasmatodea which was first established in the „Verzeichnis der Käfer Preußens“ by I.C.W. ILLIGER (1798) and FABRICIUS later works (1798). In the first edition of his great work STOLL (1788) provided beautiful hand colorated 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 A.G. OLIVIER in his „Encyclopédie Méthodique, Histoire Naturelle, Insectes“ of 1792. Most certainly 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.



Tafel von WestwoodIt was until 1833 that George Robert GRAY published the first book which was exceptionally dealing with Phasmatodea („ The entomology of Australia, in a series of monographs. The monograph of the genus Phasma.“). Only two years later, in 1835, GRAY published the first monography of the order („Synopsis of the species of insects belonging to the family of Phasmidae“) which already listed 134 described species and a great number of new generic names. Further, updated monographies of the order were published in 1838 by Hermann BURMEISTER in his series “Handbuch der Zoologie” and J.G. AUDINET-SERVILLE in the even more extensive”Histoire naturelle des insectes”.
In his work of 1842 Willy De HAAN provided detailed descriptions of numerous new species mainly from the Oriental region, most of which he illustrated on six beautiful hand colorated plates. The author interpreted the generic names established by GRAY (1833 & 1835) as subgenera of Phasma LICHTENSTEIN.Brunner
The most important and extensive publication of the 19th century dealing with Phasmatodea is undoubtly John Obadiah WESTWOOD´s „Catalogue of Orthopterous Insects in the Collection of the British Museum“ of 1859 which already listed 39 genera, comprising 471 species in two main groups – 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 showing a wide range of species make it remain an
Westwood invaluable work for the identification of phasmids. Only a few years later (1859-1870) H. de SAUSSURE published several papers describing another large number of new genera and species and in 1865 another 52 new species were described by Henry Walter BATES. All of these works followed the classification proposed by BURMEISTER (1838).
Caspar STÅL (1875) was the first to establish a new classification of the Phasmatodea and defined new criterias for a natural system of the order which he revised two times within only one year. The most important and extensive of his three works is most certainly his “Recensio Orthopterorum” of 1875 which already listed 95 genera. The paper was not only of great importance for the systematization of the order as it first introduced the terms “areola apicalis” and “segmentum medianum”, which are still of great systematical value, 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 wheter the insects posses an “areola apicalis tibiarum” or not, but did not name these groups.
In 1893 Carl BRUNNER v. WATTENWYL (1823-1914) revised the classification of the order yet again and established a system which did however not resist the strongly increasing number of described taxa and became more and more useless quite rapidly.




In addition to several earlier papers describing numerous new taxa, KIRBY published his „A synonymic catalogue of Orthoptera“ in 1904, which included new names, taxonomic changes, reference to type specimens deposited in the Natural History Museum London and the designation of a type-species for each genus. However KIRBY´s 1904 catalogue did not receive the recognition it warranted.Redtenbacher Tafel
At the beginning of the 20th century C. BRUNNER v. WATTENWYL and Joseph REDTENBACHER (1856-1926) started with their extensive research for Redtenbachera monograhic work of the order. The result of this cooperation was the outstanding monograph, published in three parts between 1906 and 1908 entitled “Die Insektenfamilie der Phasmiden” which is in Latin with a little German. The monograph recognized 1899 species and is still the most extensive, monographical study of the order. 27 excellent black and white plates include detailed drawings of almost all the genera described. Unfortunately much confusion arises when comparing this outstanding work with KIRBY´s 1904 catalogue, which was completely ignored by BRUNNER v. WATTENWYL & REDTENBACHER (1906-1908). The authors named STÅL´s two main groups “Areolatae” and “Anareolatae”and the classification presented generally arranged several equally placed “tribes” next to another and which were mainly distinguished by the length relations of the metanotum and median segment.
The first half of the 20th century has seen very few revisions, but mostly papers dealing with defined geographical regions or local faunas which are neither geographically or taxonomically complete. In this period numerous new taxa were described by e.g. J. CARL, Andrew Nelson CAUDELL, Klaus GÜNTHER, Morgan HEBARD und John A.G. REHN. REHN & REHN´s (1938) study of the tribe Obrimini of the Philippine Islands is the only work dealing comprehensively with any larger taxa. Except for some nomenclatorial changes suggested by J.J. KARNY in 1923 the classification of the Phasmatodea did not see any substantial changes till 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 into a few families and renaming the “Areolatae” and “Anareolatae” as the families Phyllidae and Phasmatidae. Later authors generally followed KARNY´s classification (e.g. HANDLIRSCH, 1930, BRUES & MELANDER, 1932 & CHOPARD, 1949). 



after 1950

It was until 1953 that GÜNTHER published his important paper „Über die taxonomische Gliederung und die geographische Verbreitung der Insektenordnung der Phasmatodea“, which provided detailed keys up to the subfamilies. Furthermore GÜNTHER divided his subfamilies into several tribes on which he commented but did not provide any further determinating tools for. Since BRUNNER v. WATTENWYL & REDTENBACHER´s (1906-1908) monograph GÜNTHER´s work is the first fundamental step towards a new classification of the order, followed by keys by BEIER (1957 & 1958).
The next important paper on classification is J.C. BRADLEY & Bella S. GALIL´s „The taxonomic arrangement of the Phasmatodea with keys to the subfamilies and tribes“ (1977) which is still widely used but generally a slightly updated translation of GÜNTHER´s work. Based on the differential characters used by GÜNTHER, the publication provides keys to all the tribes as well – as a result, keys to all 6 families, 17 subfamilies and 35 tribes of Phasmatodea are provided for the first time. Because of the exceptional work with 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 and opinions, but mostly without apparent justification.
Although quite inaccurate the classification of BRADLEY & GALIL (1977) is still widely used by entomologist´s working with the taxonomy of Phasmatodea. However, it´s most remarkable use was to provide a basic tool for identifaction to new interested entomologist´s which resulted in the fact that the first enthusiast´s began to work with Phasmatodea seriously in the early 1980´s.
Most of the more recent paper however deal with sigle taxa or the study of local faunas, important authors being e.g.: Phil E. BRAGG, Paul D. BROCK, Oskar V. CONLE, Burghard HAUSLEITHNER, Frank H. HENNEMANN, Francis SEOW-CHOEN und Oliver ZOMPRO. Especially BRAGG has done extensive studies on the Phasmid fauna of Borneo and published numerous papers (1992-2000) describing many new taxa. His beautilul 2001 book “Phasmids of Borneo” covers more than 700 pages and will be a standard work for everyone working with Oriental Phasmatodea for many years to come. For the first time the book includes revisions of certain genera and taxa in a monographical order. Since the end of the 20th century various entomologist´s have started publishing detailed revisions of different genera or tribes, which gives hope to a rapid increasing of our knowledge about these fascinating insects. Important authors are e.g. P.E.BRAGG, Ariel CAMOUSSEIGHT, O.V. CONLE, F.H. HENNEMANN, Vernon R. VICKERY and O. ZOMPRO.



The eggs of Phasmatodea – history of the Ootaxonomy

Kaup TafelThe first Phasmid egg was illustrated in 1798 by John PARKINSON alongside with the, for that time, very detailed original description of Heteropteryx dilatata („Description of the Phasma Dilatatum“).
It was however until 1871 when KAUP published a paper (“Über die Eier der Phasmiden”) which was exceptionally dealing with the eggs of Phasmatodea and in which he provided descriptions and illustrations of 26 different species. He 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 posses similar eggs...]“ and concluded that the eggs of Phasmatodea might be an important character for the distinction of species. Furthermore the author stated: „später vielleicht die Arten durch die Eier schneller unterscheiden lernt als durch die Tiere selbst. […in the future the distinction of species may proove to be more easier and relieble examining their eggs instead of the insects themselves.]“. But due to only little egg-material was available for examination during the early 20th century, KAUP´s brilliant suggestions were mostly ignored. It was until the second half of the 20th century that live material of different species was imported to Europe for breeding purposes which rapidly raised the number eggs available for examination. This lead to KAUP´s ideas being first used as a basis for further studies after remaining generally ignored for almost 100 years. Authors which have so since then done more detailed studies on the eggs of Phamatodea and used the ootaxonomy for the systematization of taxa are e.g. B. HAUSLEITHNER, O.V. CONLE, F.H. HENNEMANN, John T.C. CLARK-SELLICK (= CLARK) and O. ZOMPRO.
A terminology for the description of Phasmid egg morphology was established by CLARK (1976). Subsequently (1997-1998) CLARK-SELLICK publisehed three extensive papers dealing with the eggs of Phasmatodea and discussding their taxonomic importance for the order. One of these papers (1997) includes a key and illustrations of the eggs of 131 genera. In the 1998 paper the author discussed the form and structure of the micropylar plate and its use for the systematic of the order. Additionally CLARK-SELLICK mentioned many interesting observations but did not draw any taxonomic consequences.
The important and leading works of CLARK-SELLICK (1976-1998) have greatly improved our knowledge about the ootaxonomy of Phasmatodea, but still the eggs of many genera and species have remained unknown.



The three first described species of Phasmatodea:

1. Gryllus (Mantis) gigas LINNÉ, 1758 [Gültiger Name: Phasma gigas (LINNÉ, 1758)]

2. Gryllus (Mantis) siccifolius LINNÉ, 1758 [Gültiger Name: Phyllium siccifolium (LINNÉ, 1758)]

3. Gryllus (Mantis) phthisicus LINNÉ, 1758 [Gültiger Name: Pseudophasma phthisicum (LINNÉ, 1758)]





AUDINET-SERVILLE, J.G. (1838): Histoire Naturelle des Insectes. Orthoptères. Paris.

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.

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.

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, London, 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, London, 23: 203-228.

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.

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. (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.

ILLIGER, J.K.W. (1798): Verzeichniss der Käfer Preussens. Halle, Johann Jacob Gebauer.

KARNY, H.H. (1923): Zur Nomenklatur der Phasmoiden. Treubia, 3(2): 230-242.

KAUP, J.J. (1871): Über die Eier der Phasmiden. Berliner Entomologische Zeitung, 15: 17-24, pl. 1.

KEVAN, D.K. McE. (1982): Phasmatoptera. In: Parker, S.F. [Ed.]: Synopsis and Classification of Living Organisms, Vol. 2. McGraw Hill, New York. pp. 379-382.

KIRBY, W.F. (1904): A Synonymic Catalogue of Orthoptera, Vol. 1. British Museum, London. 501 pp.

LICHTENSTEIN, A.A.H. (1796): Catalogus Musei zoologici ditissimi Hamburgi, d III Februar 1796. Auctionis lege distrahendi. Section 3. Hamburg.

LICHTENSTEIN, A.A.H. (1802): A dissertation on two natural genera hitherto confounded under the name of Mantis. Transactions of the Linnean Society, London, 6(1): 1-39, plates 1 & 2.

LINNÉ, C. (1758): Systema Naturae per regna tria nature, secundum classes, ordines, genera, species, cum characteribus, differents, synonymus, loci. 10th Ed., 1, [IV+], 34 pp. Holmiae.

OLIVIER, A.G. (1792): Mante. In: Encyclopédie Méthodique. Histoire Naturelle. Insectes. Panckoucke, Paris, Vol. 7. pp. 619-643.

OTTE, D. & BROCK, P.D. (2005): Phasmida Species File. Catalog of the Stick and Leaf Insects of the world. 2nd Edition. The Insect Diversity Association at the Academy of Natural Sciences, Phildelphia. 414 pp.

PARKINSON, J. (1798): Description of the Phasma dilatatum. Transactions of the Linnean Society, London, 4(1): 190-192, pl. 18.

STÅL, C. (1875): Recensio Orthopterorum. Revue critique des Orthoptères decrits par Linné, de Geer et Thunberg. P.A. Norstedt & Söner, Stockholm. 105 pp.

STOLL, C. (1813): Représentation des Spectres ou Phasmes, des Mantes, des Sauturelles, des Grillons, des Criquets et des Blattes des quatre Parties du Monde. L'Europe, L'Asia, L'Afrique et L'Amerique; ressemblées et déscrites. Amsterdam. [Note: This work was published in Dutch and French and in two parts, pages 1-56 and plates 1-18 in 1788, and the remainder posthumously in 1813. The first part of 1788 did not use latinized binominal names]

WESTWOOD, J.O. (1859): Catalogue of Orthopterous Insects in the collection of the British Museum. Part 1. Phasmidae. British Museum, London, 196 pp, 40 plates.