Who Described the Species?
Around 60 individuals have described species in the Cicadellinae since Linnaeus (1758a). However, around 65% of the total currently valid species (2,300 species) have been described by just 7 individuals. It is worth presenting their work in more detail.
Walker, Francis (1808 – 1874)
Francis Walker was born in Southgate, England on 1 July 1809 and died at Wanstead, England October 5 1874. Walker was employed by the BMNH as a curator between 1844 and 1873. He described almost 20,000 new insect species, but, unfortunately, he was, sometimes a careless taxonomist, often describing the same species more than once under different specific names. The British Museum paid him 1 shilling for each new species and 1 pound for each new genus. He is best known for his catalogues of Orthoptera, Neuroptera, Homoptera, Diptera, Lepidoptera and Hymenoptera. Upon his death one (anonymous) obituary read: “More than twenty years too late for his scientific reputation, and after having done an amount of injury to entomology almost inconceivable in its immensity, Francis Walker has passed from among us.” Walker described 222 Cicadellinae species of which 138 are considered valid. Despite his generally poor taxonomic reputation, Carvalho & Webb (2005) describe in detail the labelling of Walker specimens, which has generally led to confident selection of type specimens of the species he described.
Signoret, Victor Antoine (1816-1889)
Victor Signoret was born in Paris, France 6 April 1816 and died in Paris on April 3 1889. Signoret was a qualified pharmacist and medical doctor and practised in Paris. He started his natural history studies with Coleoptera but moved his interests to Hemiptera. He published around 130 scientific papers, mostly on Hemiptera. He was a founder of scale insect taxonomy and made slide preparations of specimens in order to study their morphology (Ben-Dov & Matile-Ferrero, 1995). These authors also indicated that notebooks from his later studies were stored in the Natural History Museum Paris. These show Signoret to be a superb artist and his pencil sketches of Heteroptera (especially Cydnidae) and leafhoppers are quite remarkable. Among the major achievements of his studies on Auchenorrhyncha is the ‘Revue icongraphique des Tettigonides’, published in the Annales de la Société entomologique de France’ in parts between 1853-1855.
Signoret reviewed the known species and described 246 species (of which 211 are presently considered valid). The highlight of the work is the 19 beautifully illustrated colour plates of the dorsal view of 310 species. It is interesting that, as noted by Young (1968a), the colour of the plates varies between different copies. An indexed, uncoloured set of plates is in the library MNHM. All species were treated in the genus Tettigonia (despite some other genera being available) and Signoret indicated in the text the collection from which the specimens originated.
Upon his death in 1889, his collection of around 30,000 specimens was sold to the Vienna Museum of Natural History. There were 3000 species of Auchenorrhyncha represented by 8,300 specimens and among these were many of the specimens used in the preparation of the monograph. The specimens were re-labelled by Anton Handlirsch (1865-1935), who was in charge of the Hemiptera collection at that time. Each specimen has a printed label ‘coll Signoret’ and a handwritten locality, and a second label with a handwritten species name and printed ‘det Signoret’ (where appropriate). It is no longer clear in what way the specimens were labelled by Signoret. The Vienna collection was studied by Young in 1962-63 and he designated lectotypes for those species that he felt able to agree were eligible (Young & Beier, 1963a). Significant numbers of Signoret specimens are also to be found in der Humboldt Museum für Naturkunde, Berlin. Here it seems the specimens do contain Signoret determination labels.
Distant, William Lucas (1845- 1922)
William Distant was born in London on 12 November 1845 and died in London on 4 February 1922. He had a keen interest in natural history from an early age and he developed an interest in the taxonomy of the Coleoptera, Lepidoptera and Hemiptera. In his later life he concentrated almost completely on the Hemiptera and especially the Cicadidae. He was appointed as a part-time assistant at the BMNH, London in April 1899, where he worked on the arrangement of the Hemiptera for two or three days a week until ill health made it impossible for him to continue. He published the results of his studies at the Museum in a series of papers ‘Rhynchotal Notes’ in the ‘Annals and Magazine of Natural History’. He also contributed to Volume 1 of the Homoptera of the Biologia Centrali-Americana and the Hemiptera volumes of the Fauna of British India. Distant published several hundred titles and many of them were divided into parts. His bibliography was compiled by Dolling (1991).
Distant described 147 cicadelline species (79 Old World and 68 New World) of which 104 species are currently accepted. The majority of the type material is deposited in the BMNH, London.
Fowler, W. W. (1849 – 1923)
William Fowler was born in Gloucester, England in January 1849 and died in Reading on 3 June 1923. Canon Fowler’s father was a Vicar and after taking his degree at Oxford William Fowler was ordained a priest and eventually appointed as Canon in Lincoln Cathedral. He was drawn in his leisure time to the study of insects and especially Coleoptera and wrote ‘The Coleoptera of the British Islands’ which appeared in 5 volumes between 1887 and 1891. Despite its age this book is still highly sought after for the study of Coleoptera in Britain. His interests were not confined to British entomology and he wrote the Hemiptera-Homoptera sections of the ‘Biologia Centrali-Americana’, which appeared between 1894-1909. This work also contains some beautiful coloured plates (which are available online at http://www.sil.si.edu/digitalcollections/bca/). He described 139 (113 valid) Cicadellinae species and the majority of these are housed in London, but some others are in Vienna and Brussels (Wilson 2008). Young (1965c) selected and designated lectotypes of Fowler’s species in the course of preparation of his monographs.
Melichar, Leopold (1856-1924)
Leopold Melichar (1856–1924) was born in Brno, Moravia, Czech Republic on 5 December 1856 and died in Brno on 2 September 1924. For most of his career Melichar was a medical doctor in Vienna. He studied medicine in Prague and graduated in 1881 and first worked in Prague before moving to Vienna in 1888, where he worked as a high official with the Ministry of Health. An interest in Homoptera was suggested to him by Ladislav Duda and by 1896 he had published work on the central European fauna. His work on non-European Homoptera started when he studied specimens collected in Sri Lanka by the Czech entomologist Uzel (Melichar 1903b). In 1912 he retired to Brno to allow more time to be spent of entomology, but during WWI he became chief of a Red Cross Hospital on Brno. During his life he was an enthusiastic collector and travelled to North Africa, Spain and elsewhere in the Mediterranean region. On his death his insect collection was passed to the Moravian Museum in Brno where it is currently housed.
Melichar was the first since Victor Signoret (1816–1889) to attempt a monographic treatment of the Cicadellinae, but no part was published during his life. As well as his own collection, he also examined specimens from other European museums, especially from Budapest. His manuscript was sent to Dr. Géza Horváth at the Hungarian Natural History Museum for publication in the Annales Musei Nationales Hungarici where four parts appeared between 1924 and 1932.
However, in 1951, following questions about the completion of the series and the remaining manuscript, Dr. Vilmos Székessy published a further part in Melichar’s name (Melichar 1951a). In these monographs, seventy-seven new cicadelline genera and approximately 200 taxa in the species-group were described. These species-group taxa include many ‘varieties’, which were correctly interpreted as available names of subspecific rank by Young. Unfortunately, the work was entirely without illustrations, characters were based solely on external morphology, and the majority of his new genera, although validly published according to the nomenclatural rules of the International Commission of Zoological Nomenclature (ICZN 1999), lacked formal descriptions and type designations. China (1927d) dealt with preoccupied generic names from Melichar (1926a) and later (in 1938d) designated type species for several of Melichar’s genera based on the remaining unpublished manuscript.
Publication details of the 5 parts of the Melichar Cicadellinae monograph. All in the Annales Musei Nationales Hungarici.
Part 1 XXI (21) pp. 195-243 1924a
Part 2 XXII (22) pp. 329-410 1925a
Part 3 XXIII (23) pp. 273-394 1926a
Part 4 XXVII (27) pp. 285-328 1932a
Part 5 (n. s.) I pp. 72-111 1951a
During his European study visit in 1962-63 David Young spent some weeks in Brno and fully examined the Melichar collection and subsequently designated lectotypes for Melichar species in Brno (Young & Lauterer 1966a). Further Melichar lectotype designations were made by Lauterer & Schröder (1970a) Young also worked extensively on the specimens and the Cicadellinae collection is now fully re-curated by Pavel Lauterer and is the best represented after Raleigh, London and Washington. A visit to HNHM, Budapest in 2006 enabled Michael Wilson to study the Cicadelline collection and an inventory of the Melichar species in that museum (Wilson & Takiya, 2007).
Jacobi, Arnold (1870 - 1948)
Arnold Jacobi was born 31 January 1870 in Leipzig, Germany and died 16 June 1948 in Dresden. He studied Natural History at the University of Leipzig from 1890-95 and subsequently taught for several years before gaining a position in 1899 as scientific technician in the Biology department of the Agriculture & Forestry faculty at ‘Kaiserliches Gesundheitsamt’ in Berlin. In 1903 he applied for a position at the ‘Königl. Forstakademie’ in Tharandt (Dresden). He became Director of the ‘Königl. Zoologisches & Arthropolgisch-Ethnographisches Museum’ in Dresden in 1906. He held this position until he retired in 1936.
Jacobi was interested in many aspects of natural history and published widely on various groups. He published on Auchenorrhyncha for over 40 years (from 1902 to 1944), mostly on tropical groups, and in total described 628 species in 38 papers. Among these he described 68 Cicadelline species of which 52 are currently valid species. Many specimens he studied came from the Dresden-based company O. Staudinger & A. Bang-Haas, who sourced material especially from South America. Given the localities and collectors some of the same material was also studied by Melichar, with whom Jacobi was in contact as well with E. Schmidt.
The majority of the Jacobi type material is preserved in Dresden but some is also found in Bonn and in Berlin. Emmrich (1994) provides a list of Auchenorrhyncha papers published by Jacobi. During the course of his European visit in 1963-4 Young was sent specimens of species described by Jacobi from Dresden and most of these were designated as lectotypes by Young & Lauterer (1964a). However, the complete syntype series of each species was not examined by Young but this was rectified by Emmrich (1973) who listed all specimens. The type series of some species from Jacobi (1944a) are divided between Bonn and Dresden.
Young, David Allan (1915- 1991)
David Young was born May 26 1915, Wilkinsburg, Pennsylvania, USA and died June 8 1991, Kentucky. He studied entomology at Cornell University and received his M.S. in 1942. Following war service, he took an instructorship in the Department of Biology at the University of Louisville until 1948, after which he entered the University of Kansas to study for his Ph.D., which he attained in 1950, under the direction of R. H. Beamer. From 1950 to 1957 he worked at the USNM employed by the US Department of Agriculture. In 1957, Young accepted a position as an associate professor at North Carolina State University, where he continued his systematic research on leafhoppers (begun with Beamer) and administered the NCSU Insect Collection. Young (1958a) published a synopsis of the U.S. species of Homalodisca (Proconiini) and, after his promotion to Professor in 1961, spent the academic year in Europe (1962-1963) visiting a number of European museums where the large majority of species, which had been described by European workers were deposited. This visit enabled Young to study most of the types of Cicadellinae described by European workers and the taxonomic and nomenclatorial research became the basis of the three monographs he produced on the group (Young 1968a, 1977a & 1986a).). We now take it for granted that we take a laptop computer with us to visit museums, in order to take notes on specimens examined, perhaps we may also have a database and illustrations to assist our studies. None of this was available to Young during his year-long visit. Young published a series of papers designating lectotypes based on these studies: Stockholm, Sweden (mostly Stål species) (Young 1963a); Berlin, Germany (Young 1964a); Eberswalde, Germany (Breddin species) (Young 1965b); London, U.K. (Walker, Distant and Fowler species) (Young 1965c); Copenhagen, Denmark and Kiel, Germany (Fabricius species) (Young 1965d); Dresden, Germany (Jacobi species) (Young & Lauterer, 1964a); Brno, Czech Republic (Melichar species) (Young & Lauterer 1966a); Warsaw, Poland (Schmidt species) (Young & Nast 1963a); Budapest, Hungary (Young & Soós 1964a); Paris, France (Young 1974a). The political situation in parts of Europe made it difficult or impossible to visit certain museums. There were a number of cicadelline species (around 150 species) that Young either failed to place in a genus (especially unique female types) or failed to locate during his studies. He listed these at the end of each of the monographs (Young 1977a, 1986a) as genera and species of uncertain position.
The format of Young’s monographs was established in his 1968 study of the tribe Proconiini (Young 1968a) and continued with studies on Cicadellini (Young 1977a, 1986a). In general most Young’s lectotype designations were made prior to publication (see above and references), but frequently, taxonomic notes on these designations were given in the monographs. Young stated (1977a: 2), that for reason of time constraints, he did not list museums from where he examined specimens of previously described species, or provide detailed descriptions. He did usually figure the characters of the male genitalia or figured the head markings (often of a lectotype or topotype). Also in his monographs he did not correct mis-identifications in the literature and many localities for some previously described species will have been based on mis-identifications.
Full details and descriptions were only provided for new species. For each genus a list of species with synonyms and country records was given. The treatment of each genus in his monographs may be considered a partial revision (except where the genus and all species were newly described). Inevitably it is now clear that further work is necessary to provide a full revision in some cases, either from the availability of additional specimens or the discovery of type material unknown to Young.
There were about 150 cicadelline species that Young either failed to place in a genus (especially unique female types) or failed to locate during his studies. He listed these at the end of each of the monographs (Young 1977a, 1986a) as genera and species of uncertain position. We have followed this arrangement in the present work but cross-referenced if a species has been placed to genus or otherwise discussed in subsequent studies.
It should be noted that in his monographs Young clearly designated the holotype but not paratypes by name, listing the latter either as additional material (Young, 1977a) or ‘holotype and..... (Young, 1986a) Nevertheless, the fact that he gave a range of measurements indicated that he intended that these specimens be part of the type series and therefore paratypes (ICZN 1999 article 72.4.5). Some museum curators have labelled these additional specimens from the type series, returned after study, as paratypes, which allows easy recognition of the specimens that Young studied. In those collections where additional specimens from the same localities (and perhaps with the same data as those studied by Young) might be added under a named species, however, there is the risk that the type series, other than the holotype (the name bearing type), may not be easily recognised in the future.