WORLD BLACKFLIES (DIPTERA: SIMULIIDAE): A COMPREHENSIVE REVISION OF THE TAXONOMIC AND GEOGRAPHICAL INVENTORY [2009] Peter H. Adler1 & Roger W. Crosskey2 1Department of Entomology, Soils & Plant Sciences, 114 Long Hall, Clemson University, Clemson, South Carolina, 29634-0315, USA [email protected] 2Department of Entomology, The Natural History Museum, Cromwell Road, London SW7 5BD, UK [email protected] 2009 Inventory Page 1 WORLD BLACKFLIES (DIPTERA: SIMULIIDAE): A COMPREHENSIVE REVISION OF THE TAXONOMIC AND GEOGRAPHICAL INVENTORY [2009] Peter H. Adler & Roger W. Crosskey Introduction to the 2009 Revision The present revision of the Inventory of the world’s Simuliidae continues our intent to provide yearly, fully updated electronic revisions of the World Inventory, which originally was issued in paper format by Crosskey (1988). The current revision, thus, includes all information known to us to have been published before 1 January 2009. The purpose and format of this inventory remains the same as for previous revisions. Because each yearly update replaces previous web revisions, we begin with this revision, to retain the introductions to previous web-based revisions. The Introduction to the 2008 revision, therefore, can be found at the end of the current year’s Introduction but before the References, which include literature from all revisions, beginning with 2008. We also have retained all nomenclatural changes (e.g., new combinations and new synonyms) and all notes from previous web revisions, beginning with 2008. These past changes and notes are summarized at the end of the “Notes for the current (2009) inventory”. In this most recent revision of the Inventory, a total of 2072 formally named species (2060 living and 12 fossil) are listed as valid (Table 1), representing a net gain of 45 valid species since the 2008 inventory revision. Of the 26 valid extant genera, the genus Simulium includes just over 80% of the species. Twelve of the extant genera contain four or fewer extant species, and in time, might be shown to nest within other genera, requiring that their names be synonymized. Although the number of simuliid species being described continues to increase, we are concerned about the growing number of unavailable names. In particular, failure to designate a type specimen and to name its depository when describing a new species renders the name unavailable, according to Articles 16.4.1 and 16.4.2 in the current (fourth) edition of the International Code of Zoological Nomenclature, published in 1999 and effective from 1 January 2000. Additionally, new species continue to be described using existing names in the same genus, resulting in primary homonyms and rendering the most recent (junior) names invalid. Explanatory information on the Inventory text The explanatory information detailed here is drawn verbatim from the 2004 inventory of Crosskey & Howard, except where modified to reflect changes in our approach and updates to the current inventory revision. (1) Taxonomic information Scope The Inventory covers nominal taxa of blackflies known to us to have been described before 1 January 2009. Fossil taxa are included if they are unequivocally Simuliidae, and their palaeontological status is highlighted by enclosure of the entries in square brackets; the geological period is included as part of the provenance data. The following fossils that have been associated with the Simuliidae are excluded on grounds that their assignment to the family is clearly erroneous: Liauningius robustus (Hong, 1982) (Chironomidae), Mesasimulium lahaigouense Zhang, 1986 (family uncertain), Pseudosimulium humidum (Westwood in Brodie, 1845) (Rhagionidae), Simulia pasithea Heyden, 1870 (? Ceratopogonidae), Simulia terribilis Förster, 1891 (? Ceratopogonidae), Simulidium priscum Westwood, 1854 (Rhagionidae), Simulium ? ventricum Lin, Yao, Xiang & Xia, 1988 (family uncertain). Taxa designated in the literature informally, e.g. by letters, numbers, place names, chromosomal diagnostic features, etc., are recorded and are entered where appropriate in the text in light of present knowledge. However, the simuliid literature is enormous and we do not guarantee that every informal handle for a taxon has been unearthed. The literature is replete with cases where unidentifiable specimens have been given no particular handle and simply referred to as ‘sp.’ (e.g. ‘Simulium sp.’). We have not attempted to record these or guess at the identities of species to which they might refer when there is no other information. Subspecies and complexes The subspecies category has not been widely used in the Simuliidae and when it has, the concept has usually borne little relation to the ‘Mayrian’ geographically based concept of a subspecies. Its main use has been in various works on the Palaearctic fauna by I.A. Rubtsov and summarised in Rubtsov & Yankovsky (1988). However, Yankovsky (1995) has since ceased to recognise these Rubtsovian subspecies as having separate validity and has disposed of most of them as straight synonyms. In the Inventory we list no subspecies as such but indicate where names are still sometimes applied to subspecies by a few authors in contemporary Inventory Page 2 literature by suffixing the names with ‘(ssp.)’ where they are listed in the appropriate species entry. It is now long established that some apparently single species (morphospecies) are complexes of biologically separate sibling species, these usually defined on chromosomal criteria and conveniently called cytospecies. Where a morphospecies is known to be such an aggregate of sibling species the suffix ‘(complex)’ has been given immediately following the valid species/author/date statement. In many such complexes the precise status of the constituents is often undetermined (i.e. they are not necessarily always of definite specific status) and they still lack formal scientific nomenclature. To deal with this situation we have suffixed each such name based on chromosomal characterisation with the inert term ‘(cytoform)’: this is to be read as a non-committal umbrella for entities both of undetermined status (cytotypes) and of more or less completely definite sibling status (cytospecies). The equivalents ‘molecular form’ and ‘morphoform’ have been used for entities based, respectively, on gene sequences and hard-parts morphology but still only informally named. Inventory format Scientific (Latinised) names entering into formal nomenclature are ranged to the left-hand margin if valid, but indented if they are synonyms. Valid names are alphabetical but synonyms are in chronological order to show their relative priorities. Vernacular ‘names’ are similarly ranged left, if on current knowledge, they could pertain to valid species but are indented if they are synonyms. Vernacularly named entities are listed chronologically (alphabetically within year) at the end of the appropriate entry, after the scientific names. Occasionally, when cyto-, molecular, or morphoforms have been more finely divided, the subdivisions are indented under the more inclusive form. For example, ‘Kibwezi’ is a cytoform of S. damnosum, which has been subdivided into two cytoforms, ‘Kibwezi A1’ and ‘Kibwezi A2’. Similarly, when one form is known to be equivalent to another form, it is indented. For example, the ‘Cypress Hills’ cytoform is considered equivalent to cytoform ‘AF’, which in turn is a subdivision of cytoform ‘X’ within S. craigi. Taxonomic rank and validity or invalidity of names are distinguished by lettering style: generic and subgeneric names are in capitals (bold type when valid and lightface italics when synonyms), and specific names in lower case letters (bold type when valid and lightface italics when synonyms). In species-group headings, the specific name is highlighted by the use of capitals, e.g. ‘ORNATUM species-group’. In the very rare situation where a species appears to be ‘good’ but its name is at present technically invalid, that name is ranged left and printed in lightface italics to show that it cannot be validly used; see below under ‘Homonyms’ where such cases refer to specific names that are unreplaced homonyms. Nomenclature Spellings, authorship, dating, availability status, and other relevant aspects accord with the requirements of the current (fourth) edition of the International Code of Zoological Nomenclature published in 1999 and effective from 1 January 2000. It is noted particularly that, in accordance with articles 31.2 and 34.2, the spellings of specific names conform to the gender of the generic names with which they are combined; the gender of the subgenus (if any) is not relevant. Nomenclatural annotations. Appropriate annotations are given when necessary, for instance when a name as published is unavailable (e.g. a nomen nudum or proposed for a variety or form after 1960) or was published as a replacement name for a preoccupied (junior) homonym, or the name has the technical status of an incorrect original or incorrect subsequent spelling. Where a name has been the subject of an official Opinion of the International Commission on Zoological Nomenclature, the Opinion number is given accompanied where relevant by the word ‘List’ or ‘Index’, the former referring to the Official List of Specific Names in Zoology and the latter to the Official Index of Rejected and Invalid Names in Zoology. (There is one instance of ‘Gen. List’ (i.e. Cnetha), this referring to the Official List of Generic Names in Zoology.) Homonyms. It is necessary to call attention to a terminological