REVIEW

FORWARD INTO THE PAST

Review of Taxa, Dicky Yu, (see enclosed notice)

by David Wahl American Entomological Institute Gainesville, Florida


In terms of accessibility and ease of study, ichneumonids have the odds stacked against them. The sheer size of the family, with at least 60,000 species worldwide, has discouraged many potential workers. Then there are the various historical factors: the 1869 Förster names; the proliferation of names in the late 19th - early 20th centuries that created a situation in which it was almost impossible to identify extra-Holarctic species; the Opinions of the International Zoological Commision on Zoological Nomenclature (ICZN) on certain crucial type species; Townes' use of an idiosyncratic family-group nomenclature at odds with that used by other zoologists; and finally the lack of a catalogue for the important fauna of the Western Palaearctic. One would expect that a database of the world ichneumonid fauna would be a tremendously useful tool for organising information for systematists and applied workers. Unfortunately, the database reviewed here is instead a step backward in ichneumonid studies and must be placed with the hindrances listed above. It is one of the most confusing and destabilizing taxonomic creations that I've examined. There are three major strikes against this work: 1) it was put together by a non-systematist without any input from specialists; 2) it uses a species nonmenclature that is not in accordance with the International Code of Zoological Nomenclature (henceforth referred to as "the Code"), and 3) it is riddled with errors and inconsistencies. I shall deal with each one of these in turn.

1) Decisions are made and species assigned according by naive interpretation of recent literature, rather than by the author's judgement or informed scholarly opinion by advisors who are more cognizant of the situation. When two nomenclatures or taxonomic interpretations are at odds, Yu has followed the most recently published paper, rather than querying specialists (TAXA workbook, p. 1). He is "...not trying to create a 'catalogue' which would be a declaration of what I think the taxonomy of the group should be, but a system that reflects the published information available. If this means the system may yo-yo as taxonomists argue back and forth through their publications, so be it." (Yu, pers. comm.). This database is a de facto catalogue, however although the author may not choose to call it such, especially for other hymenopterists and applied workers. I know of no taxonomic listing of similar breath and aims that has not been the work of a specialist and has similarly reflected the compiler's specialized knowlidge. For biological control specialists and other scientists with similar needs, this is one of the biggest strikes against TAXA. Applied workers need consistent and reliable nomenclature. As identifiers, we in the systematics profession strive towards this goal by examining the arguments for the application of a name and then sticking by them until such time as new evidence is brought forth. This concern is absent in TAXA and the use of this database by non-systematists is against their own best interests.

This deliberate blindness toward argumentation and evidence has led to egregious blunders. Two TAXA examples will suffice. A listing of the Pimplini from the United States gives species of both Pimpla and Coccygomimus. Any competent specialist knows that these are one and the same genus, with the name preference depending upon whether one follows the ICZN and its Opinions. As far as I can make out, if a species was originally described in Pimpla, it is kept there even if treated as Coccygomimus by later authors; if described in Coccygomimus but never placed in Pimpla, it remains in Coccygomimus. Another example is in Ichneumon. The numerous Western Palaearctic species have never been authoritatively revised and a wide array of various ichneumonine genera (or even non-ichneumonines) lurk under the name. I printed out a list of 271 species and subspecies of German Ichneumon. After checking a variety of references that dealt with the genus as it is perceived today, I found that the true affiliation of 169 names is simple unknown; these include many species by Christ, Schrank, and other early authors which have not been dealt with since their description. The most prudent course of action would be to create a separate category of "Ichneumon incertae sedis" for all names not dealt with in modern publications. One may be sure that non-specialists will interpret these species as all belonging in the genus and thus produce conflated and misleading figures for various purposes.

2) Perhaps the single most damning aspect of this database is Yu's new system of species nomenclature, which he discusses at length in Yu (1993). He rightly points out that the recombinations of species under different genera, coupled with the necessity to conform with Latin grammar, creates homonymy that will interfere with computerized information retrieval systems. He is also right in stating that the original binomen is the only stable anchor that can correctly associate name variations. Unfortunately, he proposes that the rules of nomenclature be changed so that the original binomen becomes a permanent part of the scientific name, leading to such unwieldy monstrosities as Stirexephanes signatus Torgea formosana Uchida, 1926. Even worse, the system has been implemented in the database as if this was accepted procedure. This is all needless. I've discussed the situation with various systematists who are putting together large taxonomic databases and they point out that changing the Code is unecessary: relational database programs can incorporate the original binomen as background keys that are invisible to the searcher but will associate homonyms. TAXA, however, is not a relational database and the unwieldy association of current combinations with original binomens is Yu's solution. Basically, the zoological community is being asked to make unnecessary nomenclatural changes due to deficient programming.

3a) The endings of junior synonyms are changed, as an automatic function of TAXA, to the gender of the genus under which they are currently placed. This is not required by the Code nor followed in any other work of this nature. Valid names are not exempt from this either, as Benjaminia maurus is changed to Benjaminia maura even though the name was put forth as a noun in apposition (Wahl, 1989). While on the topic of specific names, I note that misspellings of specific epithets are rather common.

b) Species names proposed as "variety" or "form" are not listed as such in synonymies. An example is Neotheronia lineata var. concolor Krieger, 1905, which is listed simply as Neotheronia lineata concolor. Some names proposed as varieties after 1961, such as Alomya debellator var. flavotibiata Constantineanu, 1965, or A. d. var. nigrita Constantineanu, 1965, are listed as subspecies. Neither one is available under the Code (Article 45, e-g). In the Acaenitinae, Lissonta ornatus Hellén, 1937 and trochanterus Hellén, 1937 are listed as junior synonyms of Coleocentrus excitator; as both were proposed as aberrations (see Horstmann, 1990: 183 for a discussion of Hellén's aberrations), they are not available (article 45, e-f). I randomly came across these examples while checking taxa of personal interest and no doubt many more exist.

c) Ninety-six holotypes of the American Entomological Institute are erroneously listed as belonging to the Smithsonian Institution. The publication discussing the situation (Townes & Townes (1983)) was not examined.

d) Carlson (1979:738-739) listed 21 Nearctic species as unplaced to subfamily or genus, 15 of which had the type missing. Yu has uncritically assigned the latter to various genera. For example, Limneria illepida Cresson is placed in Olesicampe since Limneria is a junior synonym of the that genus. Asymmictus nigrofrons Davis is placed in Alloplasta on similar grounds. It is imprudent to place species in this manner and they should have been treated in a manner similar to Carlson's.

e) Townes (1971) listed 11 fossil ichneumonid genera, three of which were assigned to subfamilies. Since then, six more genera have been described, placed either in their own subfamilies (Tanychorinae and Pherhombinae) or in the tryphonine tribes Tryphonini, Oedemopsini, and Idiogrammatini. With the exception of Pherhombus, Yu has placed them all in the Ephialtitidae, which is peculiar considering that he cites no publication that would justify such action (and there are none to my knowledge). The fossil ichneumonid subfamily Tanychorinae (Rasnitsyn, 1980) is not even listed. Ephialtitidae and Ichneumonomidae are place in the the Ichneumonoidea, even though Rasnitsyn (1988) excluded these taxa from the superfamily.

(The following sections list errors discovered while randomly checking subfamilies and taxa of personal interest. The reader should keep in mind that many more undoubtedly exist)

f) ACAENITINAE. All authors have considered Leptocoenites to be in the Coleocentrini, with the exception of Gupta (1987) who placed it in the Acaenitini. Cursory examination of specimens reveals Gupta's placement to be a mistake. Such an anomalous record should have been referred to a specialist for clarification if the compiler was not qualified to investigate; instead, error is perpetuated.

g) ANOMALONINAE. Theriini Viereck, 1918 is not listed as a synonym of Gravenhorstiini (treated as a subfamily in TAXA.

h) BANCHINAE. Lissonotini, which is used here, is a junior synonym of Atrophini (Gauld 1984). Archoprotus porteri Bréthes, whose type is lost, was treated by Townes & Townes (1966) as a possible atrophine. Yu has placed it, with no explanation, in the Banchini.

i) CAMPOPLEGINAE. This subfamily has more than its share of mistakes. Similar to the case of Nearctic Pimpla mentioned above, Nearctic Dusona has been split into Dusona and Delopia: species from this area are in Dusona (if one accepts Gauld (1984) on the matter), but not in both! Even though Gupta (1987) rejected the record of Microcharops taitica from Tahiti as an error, it is listed here as from that island. It is with the tribes, however, that massive mistakes are made. A maximum of five tribes are usually recognized (Townesian names in brackets): Nesomesochorini, Hellwigiini, Cymodusini, Campoplegini [= Porizontini], and Limneriini [= Macrini]. Yu has managed to use six tribes, suing both Campoplegini and Porizontini. Campoplegini is used in the traditional sense (consisting of Campoplex and related genera; Gupta (1987) places Neolophron and Eucaphila here, whereas Yu places them in his Porizontini). Porizontini is used as well, however, and the contents of Limneriini are split between the two tribes! Related genera are placed in different tribes *Meloboris and Nepiera, various genera of the Hyposoter complex, Diadegma and Enytus) contrary to all previous usage.

j) ICHNEUMONINAE. The mistakes in the Campopleginae find their match in thsi subfamily. 1) Family- group names are incorrect. If Alomya is to be placed with Phaeogenes, the tribal name is Alomyini, as Townes & Townes (1951: 276) made a first-revisor decision in between the two. Yu uses Diller's (1981) subtribes and leaves out the subtribe Phaeogenina, leaving a large number of genera floating without subtribal affiliation. Gyrodontina should take precedence over Cratichneumonina. Simarly, heresiarchini is the correct name for Protichneumonini if Heresiarches and its relatives are kept within this tribe (as Yu does). 2) I was surprised to find Amblyteles recorded from North America, thinking that the genus contained only one species, the Palaearctic armatorius Förster. The nearctic species in question, Amblyteles crudosus Cresson, was placed in the genus by Townes (1945) although Carlson (1979) had deliberately left it as unplaced in Amblytelina. Ichneumonine taxonomy has advanced considerably since 1945 and unpublished notes of Townes in the AEI indicate that it is a species of Diphyus. A search of TAXA found 111 species were recorded in Amblyteles. These turned out to be mostly species described in the late 19th-early 20th century by Rodow, Berthomieu, Tischbein, and other European authors when Amblyteles was a considerably more diffuse concept than today (the tratement by Constantineanu (1959) and Perkins (1960) is in the old style). As in Ichneumon, prudence should have dictated Yu's actions. 3) Acanthojoppini and Joppocryptini are kept separate, although Heinrich (1977: 268) clearly indicated tha Acanthojoppini was the junior synonym. 4) Heinrich (1977) and Carlson (1979) assigned the nearctic Ichneumonini to subtribes. With no revisionary work on this fauna since these publication, why was the subtribal association so badly mishandled? Homotherus, Anisopygus, Limonethe, Ectopimorpha, Stenobarichneumon and a host of other genera are in the Ichneumonina; they should be placed in Gyrodontina. In particular, Stenobarichneumon, Vulgichneumon, Richtichneumon, Virgichneumon, Carinodes and Paraditremops form a well-defined complex centred about Melanichneumon, which was correctly placed in Gyrodontina. Errors of this type can be found all over -- Afrotrogus and Pepsijoppa are placed in the Trogina, instead of the Callajoppina where Heinrich (1967) placed them. The most inexplicable action, however, is the placement of 19 Neotropical genera in the Ichneumonina in the absence of any publication. As Heinrich (1967) did not propose subtribes in Ichneumonini until after Townes & Townes (1966), the neotropical ichneumonin fauna was not assigned to subtribes. Needless to say, the same errors afflicting the Melanichneumon complex mentioned earlier apply to this fauna as well. 5) In the Alomyini, both Phaeogenes and Tycherus are used: researchers who use Tycherus do so in the sense of Diller (1981), restricting Phaeogenes to semivulpinus Gravenhorst and its relatives. As used in TAXA, Phaeogenes contains numerous species that should be placed in Tycherus. Even if one does not agree with the separation of Tycherus, the situation here is guaranteed to please no one. 6) The publication dates of Heinrich's monographic series on Ethiopian ichneumonines are incorrect (see note on this subject, this issue of Ichnews); those given in Townes & Townes (1973) should have been followed.

k) LABENINAE. Labiini is used instead of Groteini, even though it is a junior synonym of the latter. The Clasini are retained in this subfamily, even though Gauld (1983) demonstrated that they do not belong here and should be regarded as a tribe of Phygadeuontinae.

l) OXYTORINAE. Oxytorinae, although otherwise seeming to conform to the restricted definition in Wahl (1990), includes Epitropus, Fetialis, and Pantomima. These clearly belong in the Orthocentrinae (Helictinae of authors); Yu was apparently confused by their appearance in a paper dealing with Oxytorinae in a less restricted sense (van Rossem, 1990).

m) PHYGADEUONTINAE. The unfortunate decision was made to follow the few authors (Gupta, 1970, 1987; Jonathan & Gupta, 1973) who elevated the tribes to separate subfamilies (viz., Phygadeuontinae s.s., Hemigastrinae, Mesosteninae). As pointed out by Gauld (1984: 101), the tribes are closely related and no other authors recognize this action.n) PIMPLINAE. Eremochila is placed with the Pimplini (sensu Fitton & Gauld (1976) -- due, I suppose, to confusion over the conflicting tribal nomenclature in the subfamily.

o) TATOGASTRINAE. Wahl (1990) is cited as providing a larval figure for Tatogaster. As stated in the paper, the larva is unknown.

p) TRYPHONINAE. Gupta (1988) restricted the Eclytini to Eclytus, and this was maintained in Townes et al (1992). Thymaridini is a junior synonym of Oedemopsini (Carlson, 1979). Yet in TAXA, oedemopsin genera are randomly distributed between Thymaridini and Oedemopsini. Acaenitellus is in Thymaridini, even though Gupta (1988) explicitly put it in Oedemopsini. Eclytus is also in Thymaridini. As far as I can make out, the genera in Oedemopsini are those mentioned in Townes et al (1992); thymaridin genera are those not mentioned (with the exception of Thymaris which apparently stayed with the tribal name.

Some final comments relating to appearance and utility of TAXA: The presentation of data on screen is rather primitive, below the standards of other taxonomic databases I've examined. In particular, synonyms are presented in an unhelpful format. Arrangement is by the categories "synonym" and "junior synonym" and then within each category numerically by taxon number. Wort of all, past synonyms are listed even though those names have been transferred out of the taxon under examination. Using Campoplex as an example, Sinophorus (with its junior synonym Eulimneria) and Sesioplex are listed as synonyms (even though they are recognized elsewhere as valid genera), along with Phaedroctonus (a junior synonym of Porizon and listed as such under that genus). This convoluted practice serves only to confuse the unwary who lack literature resources. Finally, about 7 megabytes of scanned digitized images are included. After installation of TAXA, the first priority of the user should be to dump the image files. The images are so crude that no really usable information is presented. I have a VGA screen and about all I could say was "well, yes, this looks like a campoplegine..."

After all the errors of commission and omission listed above, and the reader should remember that these were found after only cursory examination, the question arises whether TAXA is of any utility. Indeed it is, but to only about 15-20 people worldwide. These are ichneumonid specialists who will find the list of literature exceedingly useful (which it is!) or will generate lists of species for various taxa or areas. I found it useful for finding overlooked species in various groups of my current research interest. Western European workers will appreciate it as a crude substitute for a catalog of that area. BUT: only these people will have the expertise and collection/literature resources to filter out the errors that affect this database. The benefits of TAXA to a few are far outweighed by its potential harm to many. TAXA cannot be used as a reliable guide to current ichneumonid taxonomy. Unless Yu puts the species nomenclature into accordance with current practice and brings an advisory panel of specialists on board to compensate for his unfamiliarity with ichneumonids, TAXA will remain a bountiful fount of bad scholarship to plague ichneumonid systematics for years to come. I cannot recommend its purchase to hymenopterists or the general entomological community.

I wish to thank the following persons for input during the preparation of the review: John Barron, Ian Gauld, Gary Gibson, Klaus Horstmann, Charles Porter, Mike Sharkey, Larry Spears, and Chris Thompson.)


Literature Cited
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