Taxonomic Position of the Genus Puto Signoret (Homoptera: Coccinea: Pseudococcidae)
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ZOOSYSTEMATICA ROSSICA, 21(1): 97–111 25 JULY 2012 Taxonomic position of the genus Puto Signoret (Homoptera: Coccinea: Pseudococcidae) and separation of higher taxa in Coccinea Таксономическое положение рода Puto Signoret (Homoptera: Coccinea: Pseudococcidae) и разделение высших таксонов Coccinea I.A. GAVRILOV-ZIMIN & E.M. DANZIG И.А. ГАВРИЛОВ-ЗИМИН, Е.М. ДАНЦИГ I.A. Gavrilov-Zimin & E.M. Danzig, Zoological Institute, Russian Academy of Sciences, 1 Universitetskaya Emb., St Petersburg, 199034, Russia. E-mail: coccids@gmail.com Taxonomic characters of the families Pseudococcidae Cockerell, 1898, Putoidae Beadsley, 1969, Pennygullaniidae Koteja et Azar, 2008 and two superfamilies of Coccinea are discussed in view of phylogenetic meaning of these characters. It is shown that Putoidae Beadsley, 1969 is an unavailable name (nomen nudum); moreover, Putoidae sensu Williams et al., 2011 as a separate family considered in the superfamily Orthezioidea (= an informal group of “archeo- coccids”), is a taxonomic paradox that lies outside of both cladistic and evolutionary concepts in taxonomy and ignores the majority of well-known and carefully proven facts. There is not a single unique apomorphic character in the monotypic family Putoidae; all characters of the genus Puto Signoret, 1875 are plesiomorphies of all scale insects or synapomorphies with other mealybugs or with the closely related Ceroputo Šulc, 1898. Ceroputo is accepted here as a sub- genus of Puto. The monotypic fossil family Pennygullaniidae is considered by us a new subjec- tive synonym of Pseudococcidae. The taxonomic and nomenclatural problems connected with the higher taxa of scale insects and other groups of Homoptera are briefly discussed. Таксономические признаки семейств Pseudococcidae Cockerell, 1898, Putoidae Beadsley, 1969, Pennygullaniidae Koteja et Azar, 2008 и двух надсемейств подотряда Coccinea об- суждаются в свете филогенетического значения этих признаков. Показано, что Putoidae Beadsley, 1969 является непригодным названием (nomen nudum). Более того, Putoidae sensu Williams et al., 2011, принимаемое как отдельное семейство внутри надсемейства Orthezioidea (= неформальная группа “археококциды”), представляет собой таксономи- ческий парадокс, лежащий за рамками как кладистической, так и эволюционной кон- цепций в современной систематике и противоречащий большинству хорошо известных и тщательно изученных фактов. Монотипное семейство Putoidae не имеет ни одной уни- кальной апоморфной черты; все изученные признаки рода Puto Signoret, 1875 являются либо общекокцидными плезиоморфиями, либо синапоморфиями с другими мучнисты- ми червецами и близкородственным Ceroputo Šulc, 1898. Таксон Ceroputo принимается здесь как подрод Puto. Монотипное вымершее семейство Pennygullaniidae рассматрива- ется нами как новый субъективный синоним семейства Pseudococcidae. Кратко обсуж- даются также таксономические и номенклатурные проблемы, связанные с названиями высших таксонов кокцид и родственных групп Homoptera. Key words: mealybugs, scale insects, Hemiptera, Homoptera, Coccoidea, Orthezioidea Ключевые слова: мучнистые червецы, кокциды, Hemiptera, Homoptera, Coccoidea, Orthezioidea © 2012 Zoological Institute, Russian Academy of Scienсes 98 I.A. GAVRILOV-ZIMIN & E.M. DANZIG. TAXONOMIC POSITION OF THE GENUS PUTO INTRODUCTION practice (see works of E. Danzig, J. Koteja, E. Podsiadlo, R. Jashenko, I. Gavrilov and According to the paper of Williams et al. others)]: Aphidinea, Coccinea, Aleyrodin- (2011), recently published in Zootaxa, two ea, Psyllinea, Cicadinea. The International similar nominal genera of mealybugs, Puto Code of Zoological Nomenclature (below: Signoret, 1875 and Ceroputo Šulc, 1898, the Code) does not regulate now the taxo- which have been considered for many years nomic names higher than family-group as subjective synonyms, must not only be names. We follow the principle introduced distinguished as two separate genera but by Rohdendorf (1977) and consider the must be placed in two different families and suborder names as the family-group ones even in two different superfamilies of Coc- with their own coordination. cinea. Unfortunately, we cannot accept the Probably, it will be better to accept stan- opinion of the cited authors; below, we dis- dardized typified names for all animal higher cuss all the known taxonomic characters of taxa as it is done now for botanical names or Puto in comparison with other scale insects was accepted for insects by paleontologists and in view of evolutionary significance of (History of Insects, 2002). In this case all these characters. proboscidian insects will be covered by the Before further discussion, we need to name Cimicidea Laicharting, 1781. Howev- clarify our concept of the taxonomic posi- er, in view of numerous disputes about the tion of scale insects as a whole and related future of the Code we avoid accepting typi- groups, because: 1) it is not possible to con- fied names for taxa higher than suborders. sider the questions of higher classification The name Hemiptera Linnaeus, 1758, within Coccinea without consideration of frequently used (often under pressure from relationships of scale insects with other pro- editors of journals) in coccidological litera- boscidian insects; 2) many different classifi- ture as an order name for all the groups of cations have been proposed for hemipteroid proboscidian insects, is not accepted by us insects, especially in the last years [see, for in this sense, because: 1) this name was used example, the reviews of Brożek et al. (2003), by C. Linnaeus for proboscidians + thrips Forero (2008), Kluge (2010a, b)]; 3) there (Fig. 1); therefore it is an older synonym for are some nomenclatural problems connect- Condylognatha Börner, 1904; 2) for many ed with names of higher taxa of scale insects years until now, this name has been used by and other hemipteroids; 4) two principally numerous authors for true bugs (Heterop- different approaches to higher classification tera) only; 3) there are at least two sepa- of hemipteroid insects (cladistic and evolu- rate orders (Heteroptera and Homoptera) tionary) are present in the literature. within the “order Hemiptera” accepted by It seems that the oldest non-typified different modern authors. A similar taxo- name for all scale insects is Gallinsecta De nomic situation exists with the well-known Geer, 1776 (Kluge, 2000, 2010a, b). But, and widely used name Rhynchota Burmeis- instead of this name (and several other non- ter, 1835, which originally covered not only typified old names), during the entire his- proboscidians but also Siphunculata. More- tory of coccidology different typified names over, this name is preoccupied by Rhyncho- have been preferred by coccidologists: Coc- ta Billberg, 1820 (= Aphaniptera Kirby et cidae, Coccoptera, Coccoidea, Coccomor- Spence, 1815) (Kluge, 2010a). The oldest pha, Coccinea, Coccina, etc. We recognize name which covers all the recently accept- the scale insects and other homopterans ed proboscidian insects (and only them) as suborders of the order Homoptera, and is Arthroidignatha Spinola, 1850 (Kluge, use the special ending “-nea” for all typified 2000, 2010a, b). Although this name has suborder names in Homoptera [following not been used in entomological literature Pesson (1951) and internal coccidological for many years, in the modern difficult taxo- © 2012 Zoological Institute, Russian Academy of Scienсes, Zoosystematica Rossica 21(1): 97–111 I.A. GAVRILOV-ZIMIN & E.M. DANZIG. TAXONOMIC POSITION OF THE GENUS PUTO 99 Fig. 1. Copy of p. 343 from “Systema Naturae” by C. Linnaeus (1758). nomic and nomenclatural situation with the filter chamber of the digestive tract as well names Hemiptera and Rhynchota we prefer as ability to produce honeydew can be con- to use Arthroidignatha for all proboscidian sidered as synapomorphies of Homoptera insects. (Hamilton, 1981; Lambdin, 2001; D’Urso, As for the widely known and frequently 2002; present paper). We do not see any rea- discussed order name Homoptera Latreille, son to ignore these characters and prefer the 1810, we do not see serious reasons to re- probable morphological synapomorphies of ject it. It originally covered all probocidians Hemelytrata (see, for example, Emeljanov, without true bugs but with the addition of 1987) or believe the untestable data of mo- thrips. However, all other authors used this lecular cladograms based on a small number name in its modern composition, i.e. with- of studied taxa (Campbell et al., 1995; von out thrips. The notion about paraphyletic Dohlen & Moran, 1995; others; see also our content of Homoptera auct. [for review, see comments below, in the section “The data of for example, von Dohlen & Moran (1995) or DNA sequencing”). A detailed historical re- Gullan (1999)] is merely a hypothesis that vision of different phylogenetic reconstruc- considers some facts and ignores others. Ac- tions of proboscidian insects was given by cording to the cladistic point of view, the Brożek et al. (2003) and by Forero (2008) problem boils down to considering synapo- and will not be repeated here. In any case, morphies of the Hemelytrata Fallen, 1829 whether further investigations support the (Cicadinea+Coleorrhyncha+Heteroptera) hypothesis about the paraphyly of the Ho- in contrast to synapomorphies of Homop- moptera or not, it cannot be a reason to reject tera. Some authors (for example, Gullan, the taxon Homoptera. 1999) even affirm that Homoptera is char- The cladistic dogma about rejecting acterized by plesiomorphic characters only. paraphyletic