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Handbuch der Zoologie/Handbook of Zoology.
Band/Volume IV Arthropoda: Insecta.
Teilband/Part 38 Coleoptera, Beetles.
Volume 1: Morphology and Systematics
(Archostemata, Adephaga, Myxophaga, Polyphaga partim).
R.G. Beutel & R.A.B. Leschen (eds.).
Walter de Gruyter: Berlin- N.Y., 1-567
Thanks to M.G. Volkovitsh the Editorial Board obtained a possibility to get to know the volume of the training series on zoology, where a review of current publications on general problems of coleopterology as well as some divisions on particular families of beetles are given; each of families included in the volume is supplied not only with a review of literature, but also with a rather detailed information on composition and size of the family, distribution, bionomics, ecology, structure of different instars of living circle, phylogeny and taxonomy. Divisions on many families were prepared by leading experts working with them. In total 38 authors participated in preparation of the volume, including 2 native ones: V.V. Grebennikov, currently working in Canada, and M.G. Volkovitsh from ZIN. Most general chapters were prepared by the editors or with their authorship (R.G. Beutel è R.A.B. Leschen) and, therefore, they allow to conclude on the viewpoint of the editors on many aspects of coleopterology and science at all. A review of every division needs a lot of place and time, and, therefore, the reviewer found a solution to restrict his survey with some important points in the contents of the book very rich of information, without a care to pay an equal attention to every group and every chapter. It should be noted that many families, included in the volume, got most probably a more grounded rank in comparison with other interpretations of ranks and group composition (the rank was mostly borrowed from the publication by Lawrence & Newton, 1995). Particularly, the treatment of the family Hydrophilidae, according to which Helophorinae, Epimetopinae, Georissinae, Hydrochinae and Spercheinae are included there as subfamilies, seems to be quite reasonable in contrast to the opinion of some other specialists, who prefer to regard these groups as isolated families. The wide understanding of the family Leiodidae united the subfamilies Chelovinae, Platypsillinae etc., perhaps, is also rather advisable. At the same time the hierarchical composition and correlation of ranks of Scarabaeoidea, some superfamilies of Elateroidea as well as taxonomical composition and ranks among Cucujoidea and Tenebrionoidea are needed in an essential improvement. The editors mostly follow the compilation papers by J.F. Lawrence, even in cases when it is not evidently reasonable. For example, it is assumed the position of Agnathus decoratus (Germar, 1818), which it has in paper by Lawrence & Newton, 1995, but not other, not less reasonable interpretations (see a review on our site “What family Agnathus decoratus is to be placed in ?” : \rus\AGNATQUE.HTM). Although the volume editors claim that they included in their list also the families, proposed after Lawrence et al., 1999, some vexatious gaps in it were found. It is possible to admit, for example, that the editors did not accept the proposal of S.B. Kazantzev to erect the group of the genus Dexoris Waterhouse, 1878 up to the rank of the family - Dexoridae Kleine, 1933 (Kazantsev, 2003), but it is not understandable why in this list does the subfamily Maynipeplinae Kirejtshuk, 1998 lack (which was included in Lawrence et al., 1999). Thus, it is unknown which name was taken to synonymize the last of the mentioned names. It seems to be reasonable to add to the list, proposed by the editors, all synonyms for families and subfamilies, which should allow the readers to understand their interpretation correctly.
Not large, but enough profound chapter, devoted to molecular research and prepared by A.P. Vogler (pp. 17-22), instead of numerous paradox conclusions, merits a particular attention. It presents a generation of all data, obtained after comparison of nucleotide sequences. The author considers possibilities of certain genes for solution of certain tasks. For example, comparison of sequences of the mitochondrial nucleotides [Cytochrome Oxydasa I, Cytochrome b, 16S rRNA] gives good results when studying forms of the same genus, while the nuclear 18S rRNA is acceptable for study at family level. The author has available information on nucleotide sequences of 2 662 species from 123 families (if 167 families are recognized in the world fauna), at that he noted an irregular distribution of these data among groups: there are 400 species of the genus Carabus and more than 250 species of Scolytidae, while representation of other groups is considerably smaller. 28S rRNA and genes “wingless” were used with good results to examine relation among subfamily Harpalinae. On base of comparison of DNA, for example, a conclusion on monophyly as Geadephaga as Hydradephaga is made, although according to the author opinion the latter were 3 times settling the swimming in water (it is a paradox “Important monograph on water beetles of China”). At first how is it possible to make such a conclusion for the “monophyletic” group – whether it is logical to suppose that water forms of a monophyletic group several times secondarily lost their ability to active swimming (particularly if an hypotheses of water origin of the suborder in general and fusion of metacoxae with metasternum as an adaptation to swimming are recognized). A.P. Vogler arranged the families examined in an original dendrogram, where the suborder Myxophaga is located closer to the base (at Archostemata) than subordera Adephaga and Polyphaga. It is interesting that the author came to conclusion on the “monophyly” of Staphyliniformia and Scarabaeiformia, which are in accordance with that by M. Hansen (1991, 1995). It is also interesting that A.P. Vogler regards that Scirtoidea should be consider as a “sister” group to rest Elateriformia, and the superfamilies Buprestoidea and Dascilloidea are more secluded from 3 other superfamilies (Elateroidea, Cantharoidea and Byrrhoidea); at that, according to his opinion, the water elateriformian families are not “monophyletic”. The conclusions that the supefamily Cucujoidea is “paraphyletic” in relation to Cleroidea and many families of Cucujoidea fall between Lymexyloidea and Tenebrionoidea as well as the families Cryptophagidae, Latrididae, Phalacridae and Corylophidae are disposed between Tenebrionoidea, while the family Kateretidae and family Trogossitidae are shown at a base of the division of the superfamily Cleroidea look like particularly paradox.
In general division on the suborder Adephaga prepared by one of the editors (R.G. Beutel) and I. Ribera (pp. 53-55) a rather considerable place is given to comparison of molecular data (sequences of DNA). It is here noted that crawling water beetles (Haliplidae) demonstrate an especial position by both morphologically and according to their structure of DNA, in the authors viewpoint the “sister” relation of crawling water beetles and other Adephaga are “not very well supported”. They also pointed out on a peculiar view on position of this family, proposed by A.G. Ponomarenko (1977 and D.H. Kavanaugh (1986), although without consideration of it. The rather important fact to understand this group seems to be a presence in the palaeontological chronicle of palaeoendemic family Triaplidae (Middle and Upper Triassic) represented by the forms very similar to crawling water beetles. It is most probable that this extinct group is phyletically linked with recent Haliplidae. There are some essential circumstances, important for understanding of origin and placement of these families of Adephaga. Firstly, their recent species as, probably, the Triassic ones, in contrast to other predaceous groups of Adephaga, are and were mostly algophagous. Secondly, the abdominal base of their imaginal abdomen differs from that of other Adephaga in the incomplete consolidation of basal ventrites. If to recognize more close relation between Adephaga and Myxophaga, and take into consideration of the recent description of Meruidae (Spangler & Steiner, 2005), the assumption that the crawling water beetles are present a group rather ancient and maintaining many archaic features, including in peculiarities of trophics (Kirejtshuk, 1991), namely the algophagy reminds of the trophics of Meru phyllisae Spangler & Steiner, 2005 and representatives of suborder Myxophaga rather than that of all other groups of Adephaga.
The chapter on the predaceous diving beetles (Dytiscidae) written by M. Balke (pp. 90-116) is one of most detailed in the book and most multifarious in illustrations. The remarkable scheme of distribution of ecological preferences in main genera of all subfamilies (Matinae, Laccophilinae, Coptotomonae, Copelatinae, Hydrodytinae, Hydroporinae, Lancetinae, Colybbetinae, Agabinae and Dytiscinae) and tribes, which accompanies a review on bionomics and ecology of the family, has engaged reader attention, at that the data on geographical distribution of groups of the family are given in a very general view. On description of the phylogeny and taxonomy, the taxonomic interpretation accepted by the author is brought with many cladograms and molecular argumentation for phylogenetic interpretation, but the known historical data are not drawn at all.
Another chapter very detailed is the division on ground beetles (Carabidae: E. Arndt, R.G. Beutel, K. Will – pp. 119-146) which are rather numerous and diverse, and, therefore, they are also considered according to subfamilies (Paussinae, Gehringiinae, Omophroninae, Cicindelinae, Carabinae, Hiletinae, Loricerinae, Elaphrinae, Megadopinae, Siagoninae, Scaritinae, Trechinae, Harpalinae, Pseudomorphinae and Brachininae). Description of their phylogeny is essentially reduced to comparison of structural and molecular peculiarities, elaborated according to cladistic methodology without consideration of achievement of classic carabidology. The authors made some certain conclusions on historical development of the family Carabidae and its phylogeny without any reference to comparatively rich data on history of this family. Many families, including almost all small families of both Adephaga and the rest subordera are concisely considered in accordance with a certain scheme of the editors, although the authors not infrequently are rather deviating from this scheme and from the general taxonomical interpretation, accepted by the editors.
The editors did not include the superfamily Scarabaeoidea in the infraorder or series Staphyliniformia, because in their viewpoint, these groups have the only reliable synapomorphy, i.e. lack of metacoxal plate, but the rest features, which are supposed to be homologous, cannot be interpreted as a sequence of common ancestry. The chapter on water scawenger beetles (Hydrophilidae: M. Archangelsky, R.G. Beutel & A. Kumarek - pp. 158-183) is written in a rather detail. It has excellent illustrations and it is looked into level of subfamily (Helophorinae, Epimetopinae, Georessinae, Hydrochinae, Spercheinae, Horelophinae, Horelophopsinae, Hydrophilinae, Hydrophilinae and Sphaeridiinae) and the last two subfamilies into tribes - Hydrophilinae: Sperchopsini, Berosini, Chaetarthriini, Anacaenini, Laccobiini and Hydrophilini; and Sphaeridiinae: Rygmodiini, Coelostomatini, Protosternini, Omicrini, Megasternini and Sphaeridiini. The peculiarities of diverse bionomics of adults and larvae are considered in enough detail. Clown beetles (Histeridae: P.W. Kovarik & M.S. Caterino - pp. 190-121) are also considered according to subfamilies (Abraeinae, Chlamydopsinae, Hataeriinae, Histerinae, Niponiinae, Onthophilinae, Saprininae, Tribalinae and Trypanaeinae) and tribes, for each of them the list of genera and distribution of each genus are given. Diverse binomics and ecological adherences of clown beetles are described in details. The families Agyrtidae (pp. 261-269) and Leiodidae (pp. 269-280) prepared by A.F. Newton are rather laconic and informative, with detailed discussion of phylogenetic schemes, elaborated on base of the cladistic analysis. The chapter on Scydmaenidae (S.T.O. Keepe - pp. 280-288) includes among other things a complete list of genera of recent fauna with indications of distribution for each genus and also an atlas adults represented main groups of the family. Review of the family Staphylinidae enormous by number and variety as well as uniting according to the authors calculation 47 744 species occupies a rather considerable part of book (M.K. Thayer – pp. 296-344). This review is supplied by the list of subfamilies (including the subfamily Pselaphinae in the omaliin lineage) with indication of number of genera and number of species for each of them, excellent and detailed description of peculiarities of bionomics and ecology of different groups, interesting scheme on types of trophics in each subfamily, and also by the scheme on distribution of these subfamilies in different ecological conditions and different geographical zones. A rather large and important part of this chapter is represented by a detailed discussion of phylogenetic relation, which, however, is not accompanied by references on palaeontological materials as well as list of literary resources (14 pages).
One of largest and interesting division is devoted to the infraorder or series Scarabaeiformia, which is recognized by author of this division (C.H. Scholtz and V.V. Grebennikov - pp. 345-425), instead of this isolation are contested by many researchers. In contrast to the chapters on many other groups, the data on extinct faunas are used for Scarabaeiformia, including only one superfamily. Among various hypotheses the authors prefer the hypothesis on trophic dependence of this group on fungi in detritus, grounding their opinion by the table of trophics of recent groups borrowed from the publication by C.H. Scholtz and S.L. Chown (1995). However, it is thought the hypotheses on connections of ancestors of Scarabaeoidea with dedaying wood (Richter, 1958; Iablokoff-Khnzoryan, 1977; Kirejtshuk, 1989 etc.) should be recognized as more advisable, at that the mode of life and type of trophics of stag beetles (Lucanidae) and bess beetles (Passalidae) to a certain extant are similar to those of ancestors, although the mode of life of the latter has been essentially changed in connection with a secondarily developed sociality. A more close connection of both active stages of living circle with decaying wood is more probable than other types of connections admissible for ancestors. It happened most probably during the Middle Jurassic (see Taxonomic list of fossil beetles of suborder Scarabaeina (Part 2)) and at least in the Upper Jurassic Scarabaeoidea reached a considerable representation in the fauna. Further changes of trophics and mode of life were connected with that, probably, first adults and then larvae were mastering other environment and substrates of feeding, at that the decaying organic matter of plant origin was remaining a preferable substance for larval feeding. The initial adherence of ancestors of the group can be to a certain instant traced in feeding of larvae of Lethrus spp. and dung beetles (Aphodiinae and Scarabaeinae) consuming the not completely digested food of herbivorous vertebrates. Besides, in this division the argumentation of phylogenetic relations of Scarabaeoidea is considered in details, the viewpoints not shared by the authors are discussed rather correctly. If the general part of Scarabaeoidea is given in details and with a great number of illustrations, the reviews of families are rather concise, but proper scarab beetles (Scarabaeidae) are accompanied by surveys of all the subfamilies included in it.
Families of Elateriformia included in the 1st volume are considered mainly rather briefly, although nearly each family are supplied by excellent illustrations. However, the chapters on the families Elmidae (pp. 471-496) and Dryopidae (ñòð. 496-508) prepared by J. Kodada and M.A. Jach are rather detailed and well illustrated by excellent pictures of structural peculiarities of adults and larvae. Besides, the chapter on the first family indeed consists of two divisions devoted to the subfamily Larinae (Potamophilinae) and subfamily Elminae. Both have the list of recent genera with indication of number of species and general distribution for each genus, detailed description of bionomics of adults and particularly larvae and also a comparatively small part, where the phylogeny and taxonomy are considered. This is thought reasonable because of shortage of materials from extinct faunas of these groups. The family Psephenidae (C.-F. Lee, M.A. Jach and R.G. Beutel – pp. 521- 533) presented in recent fauna by 272 species and divided into four subfamilies (Eubrianacinae, Eubriinae, Psepheninae è Psephenoidinae) is also supplied by a detailed list of genera with indication number of species and general distribution for each genus, excellent illustrations, detailed description of bionomics of adults and particularly larvae, but the part devoted to phylogeny includes a detailed cladistic comparison of particular groups included in this family, and also comparison of them with other families, in particular with the families Elmidae and Lutrochidae.
It is difficult to prepare a so capacious book without any defect. It stands out that the editors usually tried to include the last references, preferring literary resources based on compilations. As a result, the inaccuracies in interpretations or even ambiguities in references can be found in different parts of the handbook. It particularly concerns the questions of evolution, phylogeny and palaeontology in the general divisions. The editors give preference to cladistic interpretations of the last years, at times pointing out that the remainder, that do not fit their schemes, does not merit any mention. The reader of these divisions gets an impression that almost all achievements of coleopterology became possible only thanks to contributions of some few specialists, working under an inspiring influence of the cladistic ideology. The following examples can be given as characteristic. R.G. Beutel pointed out that the phylogenetic relation accepted in the book (Coleoptera, Neuropterida=Neuropteroidea) was “proposed” by W. Hennig (1969), Mickoleit (1973) etc. (p. 1). Indeed the hypothesis accepted by the editors was formed in first turn due to publications A. Lamåere (1917), A.V. Martynov (1925, 1938) and R.A. Crowson (1955, 1960). The editors with reference to the publication R.G. Beutel and F. Haas (2002) declare that the important peculiarity of Polyphaga is the presence of cervical sclerites (p. 152), but it does not mention the thoughts of R.A. Crowson (1975) on connection of presence of these sclerites in different Neuroptera and Polyphaga among Coleoptera, and also on connection of absence of them in other subordera with the level of mobility of head and character of junction between the head and prothoracic segment.
The rather strange peculiarity of the editors viewpoint is to ignore fossil records because of many gaps in them and, on the other hand, to create a “phylogeny” only on base of data from recent groups without taking into consideration that the real phylogeny is scarcely a linear process and, therefore, kinds of correlations between recent species and groups could be rather different from those in the past. Also the groups were different in the past and they lived in different conditions. The editor’s efforts rather bear a resemblance to attempts to “reconstruct a phylogeny” of Tetrapoda, having casted aside all historical data from extinct faunas of this group. Historical facts can put the reader to confusion, particularly in connection with the statement that the editor recognized that he (Beutel, 1997) defined “synapomorphies” and proposed “sister relation” of the main coleopterous groups (it can be remarked that all of these things were done almost without consideration of palaeontological literature, but with declaration that fossils “not sufficient for a reliable systematic placement” – p. 6). For example, the “ovipositor unpaired, acuminate” is regarded as an apomorphy of Tshekardocoleidae (p. 2), while palaeontologists suppose that females of this family, could be, were maintaining the external ovipositor (plesiotypical feature of not only Coleoptera, but also Neuropterooidea), which was completely retracted inside the abdomen of the females of their descendants (Ponomarenko, 1969). With a reference to palaeontologists Ross & Jarzembovski (1993) (but not palaeocoleopterists) it was declared in the book that representatives of Taldycupedidae got distinct epipleura (p. 3). However, even the eldest Tshekardocolidae are characterized by a fold along vein SC, i.e. epipleura, about which was written by palaeocoleopterists (including in Ponomarenko, 1969) and which are an important diagnostic peculiarity of the order as a whole. According to the editor of this volume A.G. Ponomarenko in the book mentioned above pointed out that family Catiniidae could be phyletically linked as ancestors (“stemlineage”) with the suborder Myxophaga (p. 43), the editor also casted doubt on validity of such a link because A.G. Ponomarenko gave no “a specific justification for this placement”. But such link was not mentioned in the monograph by A.G. Ponomarenko at all. The editor suppose that A.G. Ponomarenko regarded Schizophoriformian Archostemata as aquatic groups because of “smooth surface, stream-lined body shape, hook-like structures of fore leg for possible attachment to or location on water plant” (p. 7), and “especially by the fact, that many of these fossils were found in freshwater deposits” (p. 7). These above mentioned features are on responsibility by the editor, but the Ponomarenko’s viewpoint was mostly grounded by presence of the “schiza” in groups of Schizophoriformia, which was interpreted as an adaptation to be in water and fixation of elytra and abdominal sides and for protection of this underelytral space from penetration of water. Probably it would be important to note that most terrestrial fossils are accessible thanks to sedimentation organics in water basins and study of them in lacustrine limestone. However, it would be a wrong conclusion to regard everything recovered in this substrate as related to inhabitance in water. With a reference to another palaeontologist Labandeira (1993) (but not palaeocoleopterist) the editor pointed out the appearance of the family Cupedidae in the Lower Permian (“Sakmarian”), although palaeocoleopterists and coleopterists, who study evolution and phylogeny, know that this family recorded not earlier than the Triassic, and R.A. Crowson (1981) joined the appearance of this group with the earliest traces of wood boring. The strange sentence inserted by the editor is on page 6: “Character transformations of immature stages have probably played a minor role in early evolution of Coleoptera”. The list of such kind of inexactitudes, unfortunately, can be extended.
It is impossible not to note that the appeared handbook is a rather considerable generalization of enormous materials, which were obtained by researchers of the previous century. It reflects the imagination on many groups from the viewpoint of specialists following the cladistic approach in systematics and phylogenetis popular during the last decades, but frequently ignoring other viewpoints. In this connection this book cannot be left unremarked and should be regarded as a turning-point in development of conception on this insect order. The Editorial Board of the site wishes the editors of the series a success in the preparation of the next volume, at that it would be wanted to see in this volume diversity of approaches and viewpoints, and also a correct usage of data from fossil records.
January 2005.
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