BZN Volume 61, Part 1, 31 March 2004

Comments

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Comments with the following titles were published on 31 March 2004 in Volume 61, Part 1 of the Bulletin of Zoological Nomenclature

 

Copies of these Comments can be obtained free of charge from the Executive Secretary, The International Commission on Zoological Nomenclature, c/o The Natural History Museum, Cromwell Road, London SW7 5BD, U.K. (e-mail: iczn@nhm.ac.uk).

Comments on the draft proposal to emend the Code with respect to trace fossils
(Proposal, see BZN 60: 141-142, 215-216)

(1) J. Genise 1, M. Bertling 2, S.J. Braddy 3, R.G. Bromley 4, R. Mikuláš 5, K.S.S. Nielsen 4, A.K. Rindsberg 6, M. Schlirf 7 and A. Uchman 8 (see p. 39 for addresses)

  In the recent proposal to clarify the meaning of the term ichnotaxon, Bertling et al. (BZN 60: 141-142) suggested that the Glossary definition of ‘work of an animal’ should be emended. To supplement the proposed definition of trace fossil, the draft included clearly worded lists of structures that qualify as trace fossils and otherwise. In the proposal, fossil cocoons, pupal cases, plant galls and spider webs are considered to be animal products instead of true trace fossils. However, a vigorous general discussion in the ichnologic community in July and August 2003 (Skolithos, 2003) resulted in a consensus that these terms include some structures that qualify as trace fossils, suggesting the need to refine the wording further. Such refinement is proposed herein to reduce ambiguity between trace and body fossils, delineating a sharper boundary around the ichnological realm.
  Cocoon is used in the invertebrate literature with different meanings. Many insect cocoons are structures built of silk and different amounts of other materials to oviposit or protect larvae and pupae (e.g. Chapman, 1982). Pallichnus Retallack, 1984, Fictovichnus Johnston et al., 1996, Rebuffoichnus Roselli, 1939, and Teisseirei Roselli, 1985 are available ichnotaxa for pupal chambers (or cocoons or pupal cases) found in paleosols (Genise et al., 2002). Fossil wasp cocoons have also been recorded but not named (e.g. Bown et al., 1997). Eleven ichnogenera are attributed to trichopteran (caddisfly) cases made with silk and various coarse materials in aquatic environments (e.g. Sukatcheva, 1982, 1999). The ichnotaxonomy of caddisfly cases is well developed. The first ichnogenus was erected by Bosc (1805) and, at present, this is one of the few ichnotaxonomic arrangements that utilizes ichnosubgenera (Sukatcheva, 1982). These cocoons involve behaviour (weaving) and modification of substrate (silk and other materials), thus qualifying as true trace fossils.
  In contrast, the so-called ‘cocoons’ of clitellates (e.g. Manum et al., 1991) and puparia of dipterans (e.g. Chapman, 1982) are just secretions and tanned larval cuticles, respectively; they involve neither behaviour nor modification of substrate. These more likely qualify as body fossils. Thus, we consider all kinds of constructed cocoons (e.g. woven cocoons, caddisfly cases) as trace fossils, whereas simply secreted cocoon-like structures and larval cuticles, such as clitellate ‘cocoons’ and dipteran puparia, are body fossils.
  Similarly, the proposal ruled out spider webs as traces because they were considered, along with eggs and pearls, as secretions (Bertling et al., 2003). However, in spider webs and egg cocoons, the producer weaves the silk in a second step into a construction (i.e. a trace; e.g. Foelix, 1982). Fossil silk threads from spider webs are surpassingly rare in the fossil record (Poinar, 1998); the oldest examples have recently been reported from Early Cretaceous Lebanese amber (Zschokke, 2003). Cocoons are also sometimes preserved in amber (Poinar, 1998). One ichnogenus of caddisfly cases, Secrindusia, is basically composed of silk with few if any clastic particles (Sukatcheva, 1999). Indeed, the morphology of fossil trichopteran cases shows a continuum from structures composed almost entirely of silk to those in which silk is only sparingly used to cement other materials. It is impossible to demark a boundary between traces and non-traces based on the amount of silk incorporated within a structure. Regarding the proposed definition of trace fossil, in constructed webs, nets and cocoons, silk is considered to be the substrate that is modified by the producer.
  In addition, the substrate to which these silk structures are attached may also be modified. Trichopteran silk nets and cases may play an important role in the deposition of travertine (Drysdale, 1999; Leggitt & Loewen, 2002). Silky webs, nets and cocoons are true traces, not comparable to eggshells or pearls, which are un-reworked secretions. Nor are they comparable with agglutinated foraminiferan tests, which incorporate sand grains actively on the cell wall. These tests may fully match the producer morphology, and as such they are considered body fossils.
  Plant galls may be initiated by viruses, fungi, other plants and invertebrates, especially arthropods. Accordingly, diverse kinds of galls exist that may involve the abnormal production of organs or tissues, which in turn may be patternless or show a repeated size and shape (Scott et al., 1992). Ichnologically, it is important to consider that in insect-made galls, the larva grows by feeding from the plant tissues, pupates and emerges as an adult. In doing so, it produces a boring, pellets, a pupal chamber and an exit hole, all of them true traces that have been recorded in fossil leaves and stems (e.g. Scott et al., 1992; Labandeira & Phillips, 1996). In other cases, galls may preserve only the plant reaction tissue and as such may not qualify as trace fossils. The oviposition, plus the larval boring, pellets, pupation chamber and exit hole together make up a composite trace fossil inside the reaction tissue, similar to other traces in leaves, but with disproportionate development of reaction tissue. Besides, all recognized traces of phytophagy, not just galls, produce reaction tissue (Scott et al., 1992; Labandeira, 1998). It is impossible to rule out traces involved in galls, if only because they are surrounded by particularly developed reaction tissues. Moreover, the gall inducer often directly controls the growth, shape and consistency of the reaction tissue (even by DNA transfer). It is difficult in this case to decide where the behaviour of the gall inducer starts to be replaced by mere ‘secretion’ by the plant. Hence, the term gall should be discarded because it is ichnologically ambiguous, whereas more properly defined plant reaction tissue may be included as a clear example of a structure that does not qualify as a trace fossil.
  In conclusion, the proposal by Bertling et al. (2003) is affirmed though slightly modified. The Code must be clear, and thus has to define ichnotaxon well. We propose to define an ichnotaxon as the name of a trace fossil (including burrows, borings and etchings, tracks and trackways, coprolites, gastroliths, regurgitaliths, nests, woven cocoons, spider webs, leaf mines, some type of galls, bite and gnaw structures).

Additional references
Bosc, M.L. 1805. Note sur un fossil remarquable de la montagne de Saint-Gerand-le Puy entre Moulins et Roanne, Département de l’Allier, appèle l’Indusie tubuleuse. Journal des Mines, 17: 397-400.
Bown, T.M., Hasiotis, S.T., Genise, J.F., Maldonado, F. & Brouwers, E.M. 1997. Trace fossils of Hymenoptera and other insects and paleoenvironments of the Claron Formation (Paleocene and Eocene), southwestern Utah. United States Geological Survey Bulletin, 2153: 42-58.
Chapman, R.F. 1982. The Insects. Structure and Function. 919 pp. Harvard University Press, Cambridge, Massachusetts.
Drysdale, R.N. 1999. The sedimentological significance of hydropsychid caddis-fly larvae (Order: Trichoptera) in a travertine-depositing stream: Louie Creek, northwest Queensland, Australia. Journal of Sedimentary Research, 69: 145-150.
Foelix, R.F. 1982. Biology of Spiders. 306 pp. Harvard University Press, Cambridge, Massachusetts.
Genise, J.F., Sciutto, J.C., Laza, J.H., González, M.G. & Bellosi, E. 2002. Fossil bee nests, coleopteran pupal chambers and tuffaceous paleosols from the Late Cretaceous Laguna Palacios Formation, central Patagonia (Argentina). Palaeogeography, Palaeoclimatology, Palaeoecology, 177: 215-235.
Johnston, P.A., Eberth, D.A. & Anderson, P.K. 1996. Alleged vertebrate eggs from Upper Cretaceous redbeds, Gobi Desert, are fossil insect (Coleoptera) pupal chambers: Fictovichnus new ichnogenus. Canadian Journal of Earth Sciences, 33: 511-525.
Labandeira, C.C. 1998. Early history of arthropod and vascular plant associations. Annual Review of Earth and Planetary Sciences, 26: 329-377.
Labandeira, C.C. & Phillips, T.L. 1996. A Carboniferous insect gall: Insight into early ecologic history of the Holometabola. Proceedings of the National Academy of Sciences, 93: 8470-8474.
Leggitt, V.L. & Loewen, M.A. 2002. Eocene Green River Formation “Oocardium tufa” reinterpreted as complex arrays of calcified caddisfly (Insecta: Trichoptera) larval cases. Sedimentary Geology, 148: 139-146.
Manum, S.B., Bose, M.N. & Sawyer, R.T. 1991. Clitellate cocoons in freshwater deposits since the Triassic. Zoologica Scripta, 20: 347-366.
Poinar, G. 1998. Trace fossils in amber: a new dimension for the ichnologist. Ichnos, 6: 47-52.
Retallack, G.J. 1984. Trace fossils of burrowing beetles and bees in an Oligocene paleosol, Badlands National Park, South Dakota. Journal of Paleontology, 58: 571-592.
Roselli, F.L. 1939. Apuntes de geología y paleontología uruguaya. Sobre insectos del Cretácico del Uruguay o descubrimiento de admirables instintos constructivos de esa época. Boletín de la Sociedad Amigos de las Ciencias Naturales ‘Kraglievich-Fontana’, 1: 72-102.
Roselli, F.L. 1985. Paleoicnología: nidos de insectos fósiles de la cubertura Mesozoica del Uruguay. Publicaciones del Museo Municipal de Nueva Palmira, 1(1): 1-56.
Scott, A.C., Stephenson, J. & Chaloner, W.G. 1992. Interaction and coevolution of plants and arthropods during the Palaeozoic and Mesozoic. Philosophical Transactions of the Royal Society London, B, 335: 129-165.
Skolithos. 2003. Archives of SKOLITHOS@LISTSERV.REDIRIS.ES, Forum on trace fossils. RedIRIS. URL http://listserv.rediris.es/archives/ skolithos.html, accessed October 3, 2003.
Sukatcheva, I.D. 1982. The historical development of the Order Trichoptera. Proceedings of the Palaeontological Institute of the Russian Academy of Sciences, 197: 1-111.
Sukatcheva, I.D. 1999. The Lower Cretaceous caddisfly (Trichoptera) case assemblages. Proceedings of the First Palaeoentomological Conference, Moscow, 1998: 163-165.
Zschokke, S. 2003. Spider-web silk from the Early Cretaceous. Nature, 424: 636-637.

(2) M. Bertling 2, A.K. Rindsberg 6, M. Schlirf 7, J.K. Nielsen 4, R. Mikuláš 5, J. Genise 1, K.S.S. Nielsen 4, A. Uchman 8 and R.G. Bromley 4 (see p. 39 for addresses)

  Bertling et al. (BZN 60: 141-142) have proposed various changes to a new edition of the Code, which were commented on by Tubbs (BZN 60: 215-216). His comments show that major arguments obviously have not been put in a way fully comprehensible for the non-ichnologist. Among other things, he states it is not the case ‘that the Code draws a distinction between fossilized tracks and other “works” such as galls, coprolites and nests’ (para. 7). This necessitates two replies. In a separate note, Genise et al. (see (1) above) explain why some kinds of galls and cocoons must not be considered traces or trace fossils, respectively. This should prohibit apodictic declarations such as ‘names based on fossilized galls, cocoons, etc. are ichnotaxa’ (see BZN 60: 215-216). And in the current comment, we address other arguments raised by Tubbs that need further consideration.
  A crucial issue is the relationship of the terms ‘work of an animal’, ‘ichnotaxon’ and ‘trace fossil’. Tubbs admits that ‘confusion perhaps arises from the Glossary’ (para. 3) but nonetheless argues in favour of retaining the wording of the Code and creates the impression that the current wording covers all ichnological nomenclatural needs. This is not the case. The Bertling et al. proposal originated from the need felt by ichnologists to revise the obsolete definitions in the Code that have been carried forward from earlier editions. Tubbs argues that the revised definitions differ from those given in the Code, which is true and intended. This statement is his main argument for rejecting the proposal. As biologists and geologists, we fully understand the difficulty that non-specialists have in dealing with names that are based ultimately on living behavior rather than genetic material. However, as specialists who have debated these topics for several years, we consider that we have now defined the objects of our study in the most parsimonious way.
  Further on, Tubbs states that ‘all these fossils [galls, cocoons, etc.] are commonly called trace fossils’ (para. 2). Apart from the fact that very few papers have ever been published on fossil galls and cocoons, introducing ‘common’ usage of specialist terms into a legislative work that aims at maximum clarity is counterproductive. In addition, some structures that are commonly considered the work of animals (and other organisms) are not considered trace fossils, e.g. soils and biologically induced or destroyed bedding. Against this background, Bertling et al. tried to clarify the terms ‘work of an animal’, ‘ichnotaxon’ and ‘trace fossil’ precisely because their Code definitions are obsolete. The seemingly most commonly understandable term, ‘work of an animal’, is the most ambiguous, especially as it has hardly been used in the last few decades of ichnological literature. Based on the perception that the Code should not contain ambiguous expressions, Bertling et al. favoured eliminating the obsolete and unscientific, thus unnecessary term, ‘work of an animal’ from the Code. Tubbs cites the proposal in a misleading way in stating that ‘Bertling et al. propose . . . to define ‘work of an animal’ as trace fossils . . . as well as secretions such as eggs . . .’, as it was not the aim of the proposal to define the term ‘work of an animal’ but to explain its ambiguity. Tubbs’s comment also shows why terms of common usage should be avoided. Trace fossils and their names (ichnotaxa), on the other hand, are well defined, even though some specialists in marginal fields (e.g., fossil eggs) have idiosyncratic ideas about ichnology. It would be of little help therefore to add a Glossary entry ‘trace fossil’, as offered by Tubbs (para. 7) without deleting ‘work of an animal’ from the text.
  A different issue is the question whether the Code covers ichnotaxa of non-animal origin. In the eyes of the ichnological community, it was a welcome sign of progress in the current edition of the Code to see ‘animals’ in Article 1.2.1 changed to ‘organisms’. How else should one deal with ichnotaxa whose producers cannot be assigned to a kingdom? Dozens of ichnotaxa of questionable producer assignment have been erected under the provisions of the Code and are used by active and respected ichnologists (e.g., Radtke, 1991; Vogel et al., 2000). These trace fossils may have been produced by protists, fungi, chlorophytes or cyanobacteria, i.e. they may be of animal origin or not. Tubbs denies this situation encountered in daily ichnological work in stating ‘if the agent is known not be an animal the Code does not apply’. Following this personal interpretation of Article 1.2.1, an ichnotaxon would be subject to the Code as long as its producer would be unknown; it would fall outside the provisions of the Code as soon as the non-animal origin of the trace fossil could be demonstrated. This argument can be considered unrealistic because the producer can never be identified with certainty. There will always be a chance that extinct or unknown organisms have produced structures that look like those nowadays resulting from different life activities.
  Even more dramatic is Tubbs’s misconception that ‘fossilized works of animals’ and ‘trace fossils’ are synonymous’ (para. 3). This statement, which is demonstrably erroneous, is a circular argument based on the current wording of the Code. There are many more trace fossils than those of animal origin but neither the botanical nor the bacterial Code contains provisions for ichnotaxa. This means that an ichnotaxon going back to an unknown or non-animal producer currently has no ‘legal’ standing. For these reasons, we adhere to our opinion that any trace fossils irrespective of their origin should be covered by the zoological Code.
  Some other points are uncontested. We understand the reasoning provided by Tubbs not to revoke Article 1.3.6 as originally proposed. Also, ichnologists will gladly accept the clarification that ichnofamilies require typification and that ichnofamilies do not compete with biotaxa.

Additional references
Radtke, G. 1991. Die mikroendolithischen Spurenfossilien im Alt-Tertiär W-Europas und ihre palökologische Bedeutung. Courier Forschungsinstitut Senckenberg, 138: 1-150; Frankfurt.
Vogel, K., Gektidis, M., Golubic, S., Kiene, W.E. & Radtke, G. 2000. Experimental studies on microbial bioerosion at Lee Stocking Island, Bahamas and One Tree Island, Great Barrier Reef, Australia: implications for paleoecological reconstructions. Lethaia, 33: 190-204.

(1) CONICET- Museo Paleontológico Egidio Feruglio (9100) Trelew, Chubut, Argentina
(2) Geological and Palaeontological Institute, University of Münster, Corrensstr. 24, D-48149 Münster, Germany
(3) Department of Earth Sciences, University of Bristol, Bristol BS8 1PJ, U.K.
(4) Geological Institute, øster Voidgade 10, DK-1350 Copenhagen K, Denmark
(5) Institute of Geology, Academy of Sciences of the Czech Republic, Rozvojová 135, CZ-16500 Praha 6, Czech Republic
(6) Geological Survey of Alabama, P.O. Box 869999, Tuscaloosa, AL 35486-6999, U.S.A.
(7) Institute for Palaeontology, Pleicherwall I, D-97070 Wurzburg, Germany
(8) Institute of Geological Sciences, Jagiellonian University, ul. Oleandry 2a, PL-30 063 Krakow, Poland.

Comment on the neotypification of protists, especially ciliates (Protozoa, Ciliophora)
(General Article; see BZN 59: 165-169; 60: 48-49, 143, 216-217)

S. A. Revets
Koninklijk Belgisch Instituut voor Natuurwetenschappen, Departement Paleontologie, Vautierstraat 29, B1000 Brusse, Belgium

  I am against the proposal in this article that Article 75.3.6 should be waived in relation to ciliates, other protists and small Metazoa. Successive editions of the Code have regarded the designation of neotypes as unusual acts, admissible only in exceptional circumstances. Therefore, rules governing the designation of a neotype are numerous and stringent, designed to ensure that a neotype will come as close as possible to the original concept of a nominal species.
Foissner proposes to relax the rules governing the designation of neotypes for Ciliophora and other groups of protists, and possibly even small Metazoa. As far as I am concerned, this proposal is unacceptable because it goes against both the letter and the spirit of the Code. It threatens the universality of the Code by trying to create a special interest group. Very few, if any, taxonomists have the luxury of working with a group that is free of nomenclatural problems. If we were to accept the logic of Foissner's proposal, there is a danger that each generation would be entitled to throw out the types prepared with ‘outdated’ techniques and allow ‘authoritative’ redescriptions to be made and supported by new neotypes. Advances in techniques and methods are valuable aids for taxonomy and accompanying nomenclature, they cannot be allowed to steer or control taxonomy and nomenclature. Relaxing the requirements of the neotypification process for ciliates while maintaining these same requirements for all other organisms would destroy the universality of the Code. Furthermore, it carries the very real possibility of students of other groups also making applications to waiver or to relax other requirements of the Code to facilitate their endeavours. That would destroy the universality and the authority of the Code.

Comment on the proposed precedence of Ovula gisortia Passy, 1859 over Cypraea coombii J. de C. Sowerby in Dixon, 1850 (Mollusca, Gastropoda)
(Case 3220; see BZN 59: 173-175; 60: 218-220)

Jean-Michel Pacaud
Muséum national d’Histoire naturelle, Laboratoire de Paléontologie UMR 8569 CNRS, 8 rue Buffon, F-75005 Paris, France

I disagree with the point of view of Todd (BZN 60: 218-220) on the following points:

1. The type material of Cypraea coombii only contains the complete specimen figured by Sowerby in Dixon (1850) and several remains. The specimen no. 5 mentioned by Schilder corresponds to Gisortia tuberculosa (Duclos, 1825) from the Ypresian of the Paris basin. I also stress that Edwards recorded a specimen which was originally deposited in the Museum of Bowerbank, but which was destroyed. This specimen possessed a large callosity on the dorsal face that differed from that of the type of G. coombii. This feature makes G. coombii closer to G. gisortiana than G. tuberculosa, as supposed by the authors who had erroneously regarded G. coombii as a variety of G. tuberculosa.
2. Todd disagreed with the use of the application of Article 81.2.3 of the Code for reasons which are unclear. When he referred to the geographic and stratigraphic ranges of G. coombii he only confirmed that the stratigraphic ranges of G. coombii and G. gisortiana are identical (Early Lutetian). In addition the range and rarity of the specimens cannot be used to discuss of the validity of the names. G. coombii is of course a rare species, but G. gisortiana is also rare in France.

3. Todd regarded the work by Schilder (1930) on Gisortia as the most complete treatment. Luc Dolin (world specialist of the cypraeids) and I do not consider that this work is the best work for the systematic treatment of Gisortia. Considering Gisortia and Megalocypraea (= Gisortia), Schilder discussed 35 taxa of which only five have a preserved shell; thirty taxa are preserved as internal moulds lacking useful morphological characters. This treatment is totally absurd in introducing many species, for which distinctive characters at generic level (e.g. terminal folds, fossula) as well as specific level are missing and only exist when the shells are well preserved. Thus, it appears that in his systematic treatment, Schilder discussed one of the rare specimens of G. coombii which has the shell preserved. Moreover, we also point out that the synonymy list presented by Schilder is very short.

4. Among six reasons given by Schilder for the systematic treatment of the family, Todd remarks that two (Schilder’s numbers 4 and 6) are relevant to the current application. Reason no. 4 (many species are known from only one or a few specimens) cannot be used as an argument against the synonymy of G. coombii and G. gisortiana. Moreover, this argument effectively favours the splitting of species. If Reason no. 6 (most writers have had no opportunity to examine original specimens) is pertinent for previous authors, it is not so in our case. We have examined the type specimens of all species attributed to Gisortia: tuberculosa, coombii, gisortiana, pterophora and chevallieri. The other type specimens are useless internal moulds. In addition, we also have the opportunity to check the type specimens of the subgenus Vicetia for further comparison. Consequently, I consider that we are able to give an accurate statement of the synonymy of G. coombii and G. gisortiana.

5. The short biometric analysis based on preserved shells (see figure below) justifies the taxonomic distinction between G. gisortiana and G. tuberculosa, and also demonstrates that G. coombii is closest to G. gisortiana. Also, for qualitative characters, the species strongly varies and G. coombii could be easily included in the range of variation of G. gisortiana. The callosities mentioned by Todd cannot be regarded as discriminant characters. The French specimens bear either very thick or very thin callosities and some specimens lack callosities. We have not only compared the available specimens using the size, but we have also used the variation of qualitative characters.

6. In our original manuscript we stated that Gisortia gigantea should be regarded as a nomen dubium. However, this point was not included in the published application. For further information, I stress here that Conus gigantea Keferstein, 1828 and Conus gigantea Quenstedt, 1836 are nomina nuda (see Schilder & Schilder, 1971). Strombus giganteus Goldfuss, 1841 (based on an internal mould) should be regarded as a nomen dubium as well as S. giganteus (see Shalfhaütl, 1863). Unfortunately, Dolin & Dolin (1983) regarded the taxon G. gisortiana as a synonym of G. gigantea, but we now regard this nomenclatural act as an error. Todd cited this work in taking the example of Vredenburg (1927). Dolin and I recognise that the taxon G. gisortiana was often considered as a synonym of G. gigantea or that G. gigantea was regarded as a distinct species. In addition, Schilder (1927) erected three species of Gisortia including four subspecies of G. gigantea based on internal moulds. What is the scientific credibility of these taxa?

7. In conclusion, I hope that our proposition is not as premature and unsubtantiated as suggested by Todd. Regarding the occurrence of the taxa, it is also unrealistic to believe that a study based on numerous specimens can be made, only a few specimens being discovered in 180 years. Thus we do not understand why Todd disagreed with our proposal to apply Article 23.2 in order to conserve the taxon G. coombii.

Measurements of Gisortia tuberculosa (black diamond); G. gisortiana (white circles) and G. coombii (black circle).

Comment on the proposed conservation of usage of the specific names Libellula aenea Linnaeus, 1758 (currently Cordulia aenea) and L. flavomaculata Vander Linden, 1825 (currently Somatochlora flavomaculata; Insecta, Odonata) by the replacement of the lectotype of L. aenea with a newly designated lectotype
(Case 3253; see BZN 60: 272-274)

G. Dévai
c/o Am Liebfrauenbusch 3, D-26655 Westerstede, Germany

  I support this application fully. It is a completely convincing and acceptable approach to providing nomenclatural stability.

Comment on the proposed precedence of Bolboceras Kirby, 1819 (July) (Insecta, Coleoptera) over Odonteus Samouelle, 1819 (June)
(Case 3097; see BZN 59: 246-248, 280-281; 60: 303-311)

Brett C. Ratcliffe
Systematics Research Collections, W436 Nebraska Hall, University of Nebraska, Lincoln, NE 68588-0514, U.S.A.

  I support the application to use Bolboceras Kirby 1819 (July) instead of Odonteus Samouelle 1819 (June) because of prevailing usage of the name Bolboceras. The name Odonteus has not been used since it was first proposed except by Krell in 1990, who suggested that Odonteus should be used since it had priority. To do so, however, would not only destabilize the nomenclature for the often cited genus name Bolboceras, but it would also decrease the flow of information that is normally associated with the name Bolboceras. Moreover, Bolboceras is also the type genus for the tribe BOLBOCERATINI and subfamily BOLBOCERATINAE. It would be confusing to use Odonteus instead when referring to the type genus of the family group names. The second volume of ‘American Beetles’, published in 2002, will be the standard for North American Coleoptera classification for the next several decades, and Bolboceras is used in this work (with reference to the current application to the Commission).

M.L. Jameson
University of Nebraska State Museum, W436 Nebraska Hall, Lincoln, Nebraska 68588-0514, U.S.A.

H.F. Howden
Canadian Museum of Nature, P.O. Box 3443, Station ‘D’, Ottawa, Canada K1P 6P4

  Comments by Krell et al. (BZN 60: 303-311) on the proposed precedence of Bolboceras Kirby over Odonteus Samouelle clearly demonstrates that three names are currently being used for one taxon, thus creating confusion within the literature. In our proposal (BZN 59: 246-248), we seek stability and universality in nomenclature. Krell et al. provide a substantial list of literature that references the names Bolboceras Kirby, Odonteus Auctorum, or Odontaeus Dejean for the same taxon. In our proposal, we made the case that preservation of the name Bolboceras would lend the greatest nomenclatural stability based on prevailing usage. Worldwide, the name Bolboceras has been used extensively in the literature for over 180 years (e.g. Klug, 1845, pp. 36-56; Lacordaire, 1856, p. 142; Boucomont, 1912, pp. 7-14; Curtis, 1829, p. 259; Schaeffer, 1906, pp. 249, 253; Paulian, 1959, p. 44; Benasso, 1971, p. 133; Nikolaev, 1987, pp. 27-28; Barbero & Cavallo, 1999, p. 70). The name Odonteus was first brought to the attention of taxonomists by Krikken (1978). Krell (1990) subsequently synonymized Bolboceras under Odonteus. Krell (1990) proposed that the Principal of Priority be implemented, and that the name Odonteus should be used instead of Bolboceras. Based on Krikken (1978), a few authors in Europe implemented use of Odonteus prior to its formal synonymy (Nikolaev, 1980; Jessop, 1986; Shirt, 1986).
  Although Krell et al. provide a long list of literature to support their opinion, the overwhelming majority of this literature cannot be taken into account for two reasons: (1) the definition of prevailing usage according to Article 23.9.6 and (2) the status of actions of the Commission according to Articles 80 and 82.1. As stated in Article 23.9.6, the mere listing of the name in an index, abstracting publication, nomenclator, or list of names must not be taken into account in determining prevailing usage. Thus, most references provided by Krell et al. must be eliminated from their argument (e.g., Gürlich et al., 1995; Alexandrovitch et al., 1996; Hansen, 1996; López-Colón et al., 1996; Rössner, 1996; Telnov et al., 1997). In addition, when a case is under consideration by the Commission, the prevailing usage (Bolboceras) is to be maintained until the ruling of the Commission is published (Articles 80 and 82.1). Based on these Articles and the Case acknowledgement date (December 1998, BZN 55: 205), several references provided by Krell et al. must be eliminated from their argument because they postdate the Case (e.g. Nádai & Merkl, 1999; Martín-Piera & López-Colón, 2000; Rheinheimer, 2000; Carpaneto et al., 2001; Geiser, 2001; Jaszay, 2001; Krell, 2001; Lo Cascio, 2001; Ballerio, 2002; Frank & Konzelmann, 2002; Schaefer, 2002)
 Krell et al. argue that the name Odontaeus Dejean is not a separate generic name but is a subsequent incorrect spelling of Odonteus Samouelle. The name Odontaeus (with or without the correct author attributed) further confounds nomenclatural stability. The name (with or without the correct author attributed) often appears in the literature (e.g. Boucomont, 1902; Wallis, 1928; Paulian and Baraud, 1982; Zunino, 1984; Baraud, 1992; Bunalski, 1999). Krell (1990) stated that Odonteus should be given priority over Bolboceras because of the orthographical similarity in spelling of Odonteus and Odontaeus, thus easing the transition to a new generic name. However, these names are not a reflection of a subsequent incorrect spelling for two reasons. First, Samouelle (1819) attributed Odonteus to Köppe, whereas Dejean (1821) attributed Odontaeus to Megerle. If these names were the same, then the authors would have attributed the name to the same individual. Second, Samouelle included only Scarabaeus mobilicornis Fabricius in his description, whereas Dejean included several previously described species as well as Scarabaeus mobilicornis Fabricius. The inclusion by Dejean of many previously described species shows that the concepts for Odonteus Samouelle and Odontaeus Dejean were different.
  Usage of Odonteus Samouelle (or other authors), 1819 causes further nomenclatural confusion due to its homonym, Odonteus Agassiz, 1838. The name Odonteus Agassiz was in prevailing usage (e.g. Blot, 1988) until Krell (1991) noted that Odonteus Samouelle had nomenclatural priority. Based on the Principles of Priority and Homonymy, Krell (1991) proposed the replacement name Odonteobolca Krell, 1991 for Odonteus Agassiz. Confusion with these homonyms further adds to nomenclatural instability that would result from precedence of Odonteus Samouelle over Bolboceras Kirby as proposed by Krell et al.
  Krell et al. suggest that the type species designation for Bolboceras Kirby is unequivocal. Their proposal to the Commission to designate Scarabaeus quadridens Fabricius, 1781 as the type species for the genus would create a junior synonym (Indobolbus Nikolajev, 1979) and would further disturb nomenclatural stability. Contrary to the opinion of Krell et al., Kirby did not declare ‘explicitly that he used exclusively B. quadridens to describe the genus’. Kirby (1821) stated that ‘details of Bolboceras were taken from B. quadridens’ in reference, we think, to the detailed drawings of the mouthparts of B. quadridens that accompany the dorsal habitus drawing of Bolboceras australasiae Kirby. The image that accompanies Kirby’s description shows a dorsal habitus of B. australasiae with the mouthparts of both B. australasiae and B. quadridens surrounding the image. Thus, Kirby’s statement does not unequivocally establish the type species for Bolboceras, and there is ample ambiguity regarding Kirby’s ‘intention’ (as Krell et al. assert) that Scarabaeus quadridens Fabricius, 1781 is the type for the genus. Curtis (1829, p. 259) unequivocally established the type species of Bolboceras Kirby as Scarabaeus mobilicornis Fabricius (by subsequent designation).
In summary, Krell et al. do not demonstrate stability or universality in the usage of Odonteus Samouelle. Their proposal to the Commission to designate Scarabaeus quadridens Fabricius, 1781 as the type species for Bolboceras Kirby further destabilizes usage because the type species has already been clearly designated (Scarabaeus mobilicornis Fabricius, 1775). Their proposal for precedence of Odonteus Samouelle over Bolboceras Kirby further confounds nomenclatural stability because of confusion with the junior homonym, Odonteus Agassiz, 1835, and the name Odontaeus Dejean. Nomenclatural stability and universality would be achieved by conserving the usage of Bolboceras Kirby, a name that has been prevalent for over 180 years. We stick by the proposals made to the Commission in our application.

Additional references

Blot, J. 1988. La representation schématique des relations existant entre le squelette axial et la squelette interne des nageoires impaires chez un grand nombre de Téléostéens (formule Blot), premet de rectifier une erreur de determination et de mettre un evidence la croissance allométrique des rayons épineux des nageoires impaires dans l’espèce Acanthonemus subaureus (de Blainville) de l’Eocène inférieur de Bolca (Italie du Nord). Compte Rendu de l’ Academie des Sciences, Paris, sér. II, 307: 1131-1136.
Lacordaire, J.T. 1856. Histoire Naturelle des Insectes. Genera des Coléoptères, vol. 3. 594 pp. Librairie Encyclopédique de Roret, Paris.
Schaeffer, C. 1906. On Bradycinetus and Bolboceras of North America, with notes on other Scarabaeidae. Transactions of the American Entomological Society, 32: 249-260.

Comments on the proposed conservation of Lius Deyrolle, 1865 (Insecta, Coleoptera)
(Case 3194; see BZN 60: 132-134)

(1) Svatopluk Bílý
Department of Entomology, National Museum, Kunratice 1, 14800 Praha 4, Czech Republic

I support this application.

(2) Mark G. Volkovitsh
Zoological Institute of the Russian Academy of Sciences, Universitetskaya nab. 1, 199034 St. Petersburg, Russia

I support this application.

(3) Mark Yu. Kalashian
Institute of Zoology, National Academy of Armenia, ul. Sevaka 7, Erevan, Armenia

I support this application.

(4) Allen Sundholm
Turrella, Sydney, N.S.W., Australia

I support this application.

Comments on the proposed precedence of the specific names Acmaeodera oaxacae Fisher, 1949 and Polycesta deserticola Barr, 1974 (Insecta, Coleoptera) over those of Acmaeodera philippinensis Obenberger, 1924 and Polycesta aruensis Obenberger, 1924 respectively
(Case 3257; see BZN 60: 124-126)

(1) Svatopluk Bílý
Department of Entomology, National Museum, Kunratice 1, 14800 Praha 4, Czech Republic

  I do not support this application. Incorrect locality data that do not reflect real distribution are not acceptable grounds for reversal of precedence in relation to these names.

(2) Mark G. Volkovitsh
Zoological Institute of the Russian Academy of Sciences, Universitetskaya nab. 1, 199034 St. Petersburg, Russia

  I do not support this application. It does nothing to improve nomenclatural stability.

(3) Mark Yu. Kalashian
Institute of Zoology, National Academy of Armenia, ul. Sevaka 7, Erevan, Armenia

  I do not support this application.

(4) Allen Sundholm
Turrella, Sydney, N.S.W., Australia

  I support this application.

Comments on the proposed conservation of usage of the names Phymaturus Gravenhorst, 1837 and Lacerta palluma Molina, 1782 (currently Phymaturus palluma; Reptilia, Sauria) by designation of a neotype for Lacerta palluma
(Case 3225; see BZN 60: 38-41, 58, 313-314)

(1) Darrel Frost
Division of Vertebrate Zoology, American Museum of Natural History, Central Park West at 79th Street, New York, NY 10024-5192, U.S.A.

  I support this application. The lizard in question has a substantial literature outside of systematics and so stabilizing the name Phymaturus palluma will have a salutary effect on communication among biologists.

(2) Richard Etheridge and Jay M. Savage
Department of Biology, San Diego State University, San Diego, California 92182-4614, U.S.A.

  In response to the comments by Veloso et al. in BZN 60: 313-314, we make the following comments. The facts relating to the various names in this case are not at issue. However, Phymaturus palluma and Callopistes maculatus were consistently and universally applied in the manner supported by our proposal from 1837 until 1985, as acknowledged by Veloso et al. in the first paragraph of their comment to the Commission. As their remaining remarks make clear, the 1985 paper by Cei & Lescure (see also Lescure & Cei, 1991; Veloso et al., 2000) ushered in a two decade period of instability in the previous long accepted usage of the involved names. In contrast, our proposal contributes to stability by preserving over 150 years of prevailing usage by unambiguously establishing the application of the generic and specific names to those taxa with which they have been long associated.

Comments on the proposed conservation of usage of the specific name Palaeortyx phasianoides Milne-Edwards, 1869 (Aves, Galliformes) by the designation of a neotype
(Case 3266; see BZN 60: 211-214)

(1) Bradley C. Livezey
Section of Birds, Carnegie Museum of Natural History, 4400 Forbes Avenue, Pittsburg, PA 15213-4080, U.S.A.

  I fully support this application.

(2) Zlatozar Boev
Fossil and Recent Birds Department, National Museum of Natural History, Bulgarian Academy of Sciences, 1 Blvd. Tsar Osvoboditel, 1000 Sofia, Bulgaria

  I support this application.

(3) Zygmunt Bochenski
Institute of Systematics and Evolution of Animals, Polish Academy of Sciences, 31-016 Krakow, Slawkowska 17, Poland

  I support this application.

(4) Peter Ballmann
Am Nordpark 1b, 50733 Köln, Germany

  I recommend that this petition to the Commission be accepted without amendments.

(5) Gareth Dyke
Department of Zoology, University College Dublin, Belfield, Dublin 4, Ireland

  I totally agree that the usage of the name Palaeortyx phasianoides should be maintained in the ornithological literature by the designation of a neotype. Confusion surrounding this issue has been needlessly created by Mlíkovský (2000) and is clearly based on misidentifications and taxonomic inaccuracies. This action will go far to avoid further confusion with regard to the nominal species Palaeortyx phasianoides (Gruiformes), Mionetta blanchardi (Milne-Edwards, 1863) (Anseriformes) and Ameripodius alexis Mourer-Chauviré, 2000 (Galliformes).

Comment on the proposed conservation of the specific name of Vespertilio nanus Peters, 1852 (currently Pipistrellus nanus; Mammalia, Chiroptera)
(Case 3240; see BZN 60: 42-44, 314-315)

Dieter Kock
Forschungsinstitut Senckenberg, Senckenberg-Anlage 25, D-60325 Frankfurt am Main, Germany

  I do not support the application to suppress the name Vespertilio pipistrellus africanus Rüppell, 1842 for the African banana bat. This taxon as currently understood is apparently not monotypic as documented by published research work. Until a comprehensive revision of the complex has been achieved, untimely nomenclatural actions are liable to create further instabilities.