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subspecies AND varieties in the same paper (implication of different “ranks”), and the varieties were based on slightly different traits of coloration. These facts unambiguously reveal that Ohaus’ varieties were proposed for infrasubspecific entities. Infrasubspecific entities are not regulated by the ICZN. Ohaus (1904b) described the new species P. nigrosternalis and P. puncticollis with varieties in the same publication. The specimens used in the descriptions of the varieties are excluded from the type series of the nominal species-groups taxon in these cases. Article 72.4.1 of the ICZN states that specimens referred to by name as “distinct variants” are excluded from the type series. Designation of Lectotypes and Neotypes Article 74.7.3 of the ICZN requires that designations of lectotypes after 1999 must “contain an express statement of the taxonomic purpose of the designation” (International Commission on Zoological Nomenclature 1999). Lectotypes are designated in the genus Platycoelia in order to preserve the stability of nomenclature by selecting one specimen as the sole, namebearing type of the taxon. The lectotype specimen serves to tie the published name to an actual specimen and as a reference standard for the taxon. Lectotypes were selected for the following names: Callichloris baessleri Ohaus, C. bocki Ohaus, C. helleri Ohaus, C. signaticollis Burmeister, Leucopelaea baronis Ohaus, Melolontha flavostriata Latreille, P. abdominalis Ohaus, P. alternans Erichson, P. boliviensis Blanchard, P. burmeisteri Arrow, P. burmeisteri Ohaus, P. chrysotina Ohaus, P. confluens Ohaus, P. flavoscutellata Ohaus, P. forcipalis Ohaus, P. gaujoni Ohaus, P. hirta Ohaus, P. humeralis Bates, P. inflata Ohaus, P. inflata tucumana Ohaus, P. intermedia Ohaus, P. laevis Burmeister, P. limbata Ohaus, P. lutescens Blanchard, P. marginata Burmeister, P. mesosternalis Ohaus, P. nervosa Kirsch, P. nigrocauda Bates, P. nigrosternalis Ohaus, P. occidentalis Ohaus, P. olivacea Blanchard, P. parva Kirsch, P. MONOGRAPHIC REVISION OF PLATYCOELIA 25 pomacea Erichson, P. pomacea brasiliensis Ohaus, P. prasina Erichson, P. pulchrior Ohaus, P. puncticollis Ohaus, P. quadrilineata Burmeister, P. rufosignata Ohaus, P. scutellata Guérin-Méneville, P. simplicior Ohaus, P. steinheili Ohaus, P. tschudii Ohaus, P. unguicularis Ohaus, P. valida Burmeister, P. variolosa Ohaus, P. wallisi Ohaus. I consider that lectotypes are necessary for these names due to the long history of taxonomic confusion of some species and names in this genus. It is particularly critical because early authors did not explicitly state how many specimens where in the original type series. Information about collecting locality and date, collector, and type depository were also often omitted. This makes it impossible to determine with a high degree of accuracy what and where the ENTIRE type series is. If lectotypes are not designated, then the possibility will remain that “rogue” syntypes will surface and destabilize the application of the name to that taxon. Platycoelia, and many other groups of scarabs, have had many species described based on mixed series of specimens later considered to represent multiple species. The ICZN requires that a designation of a neotype “is validly designated when there is an exceptional need and only when that need is stated expressly” (Article 75.3). As with the lectotype designations, two neotypes are designated in the genus Platycoelia in order to preserve the stability of nomenclature by selecting one specimen as the sole, namebearing type of the taxon when the original name-bearing type specimen(s) were lost or destroyed. The neotype specimen serves to tie the published name to an actual specimen and as a reference standard for the taxon. Other qualifying conditions for designating valid neotypes in section 75.3 of the code are satisfied in the discussions and descriptions of the individual species. A neotype was selected for Callichloris alticola Gutiérrez and Callichloris laelaps Gutiérrez. I consider that neotypes are necessary for this names due to the history of taxonomic confusion of species and names in this genus. Until revisionary work is done on long-neglected groups such as Platycoelia, the taxonomy and classification
26 BULLETIN OF THE UNIVERSITYOF NEBRASKA STATE MUSEUM are “complex zoological problems” and there is doubt surrounding the identities of all species and names. Species Concept I utilize the Phylogenetic Species Concept outlined by Wheeler and Platnick (2000): species are the smallest aggregation of populations diagnosable by a unique combination of character states. Each species description is a scientific hypothesis. The number of specimens examined (given at the beginning of each description) is the number of replicates studied. Character states are used to support the hypothesis that all the individuals examined are of the same species. The different species can be recognized by a unique combination of character states because they are isolated genetically from other species (in sexual organisms such as Platycoelia). Population genetics has demonstrated the inevitability of character state fixation due to various forces, including genetic drift and natural selection. Different character states will become fixed in different species due to genetic isolation. As is generally true, the strength of the hypothesis (species description) increases with the number of replicates (specimens) examined. Not all species are equally diagnosable. Some are easily recognized by examining one individual with a unique set of characters (for example a male with radically different parameres), and some must be proposed only after many individuals from different populations are examined (when species have recently diverged or have not changed drastically since their divergence so fixed character states are difficult to recognize). These analyses must be done in the context of other closely related species. Examining intraspecific variation within closely related species is an important tool in distinguishing fixed character states from phenotypic traits that vary within a population or species. When traits are mistaken for fixed character states, the number of species can be overestimated. No species description (hypothesis) is accurate for all of the individuals in a species. There are sometimes anomalous individuals that have unusual traits or have been malformed during development. Size and coloration of some species many vary widely from year to year, depending on the amount of rain, food availability, temperature, etc. Some populations also may exhibit phenotypic variation of certain traits at different levels from other populations. There is also the possibility of some low level of interspecific breeding and gene flow between closely related species. Ideally, a taxonomist would examine large series of specimens from single collecting events and from different geographic areas within the distribution of a given species to get a full appreciation of the intraspecific variation. Unfortunately, this is often a luxury we do not have. The opposing forces of overwhelming species richness of insects and disappearing habitats, research dollars, and expertise make career-long taxonomic studies of limited groups impractical. Because all proposed species are scientific hypotheses, examining more specimens and specimens from different localities and populations can test their validity. I expect that some of my Platycoelia species designations will have to be slightly modified as more specimens become available. I have not seen a compelling argument for maintaining subspecific names in the context of the phylogenetic species concept. In fact, subspecies concepts all seem subjective and reliant on the whims of the experts on the given group to determine what populations are deserving of the status. To me, this harks back to the days before cladistic analysis when taxonomic authorities would present intuitive phylogenies as fact. I am not opposed, on principle, to naming subspecies, but I have yet to find an objective set of criteria for determining whether different populations warrant subspecific status. Therefore, I do not recognize subspecies as valid taxa in this work. Ohaus (1904b) proposed two subspecific names within Platycoelia. Types of each were examined and determined to be synonymous with the nominal species.
Page 2 and 3: Bulletin of the University of Nebra
Page 4 and 5: C O N T E N T S Abstract...........
Page 6 and 7: LIST OF FIGURES Fig. 1. Platycoelia
Page 8 and 9: Bulletin of the University of Nebra
Page 10 and 11: Members of the genus Platycoelia De
Page 12 and 13: alone to divide up the rutelines (a
Page 14 and 15: eing the most recent authority on t
Page 16 and 17: els Amongst the Great Andes of the
Page 18 and 19: tion. Zischka was a German who live
Page 20 and 21: Puncture size: large = 0.17 mm or l
Page 22 and 23: 26. Pronotal surface sparsely punct
Page 24 and 25: Fig. 11. Platycoelia alternans male
Page 26 and 27: MONOGRAPHIC REVISION OF PLATYCOELIA
Page 28 and 29: Fig. 33. Platycoelia rufosignata me
Page 30 and 31: CNCI Canadian National Collection o
Page 34 and 35: PHYLOGENETIC PATTERNS AND BIOGEOGRA
Page 36 and 37: metaunguitractor plate usually with
Page 40 and 41: larvae, and Paucar-Cabrera and Smit
Page 50 and 51: Erichson (1847) did not state how m
Page 52 and 53: species]; Ohaus 1918:178 [catalog l
Page 58 and 59: Platycoelia marginata and Platycoel
Page 60 and 61: tarsomere and metatarsomere 5 with
Page 62 and 63: 5. Platycoelia meridensis new speci
Page 64 and 65: in apical half. Protarsomere 4 with
Page 68 and 69: 7. Platycoelia prasina Erichson, 18
Page 70 and 71: written and typeface), e) “PLATYC
Page 72 and 73: mentum. Apex of mentum with triangu
Page 74 and 75: 1972:302 [catalog listing as subspe
Page 76 and 77: green to yellow; parameres with the
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duced tooth. Apex of mentum with tr
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Mnisnech,” “Dr Ohaus Vidit 1903
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VARIATION. Male (n=19). Length 19.2
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“Platycoelia forcipalis cotype Oh
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males with internomedial stridulato
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MONOGRAPHIC REVISION OF PLATYCOELIA
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tures moderate with dark coloration
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gins. Ventrally with dark brown met
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19. Platycoelia flavostriata (Latre
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20. Platycoelia nigrosternalis Ohau
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LOCALITY DATA. 66 specimens examine
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Calloodes nitidissimus Lea, 1919 (C
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