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History of Scarabaeoid Classification

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HISTORY OF SCARABAEOID CLASSIFICATION BERT KOHLMANN Reprinted from COLF.OPTERISTS SaCIETY MONOGRAf'H No. 5 Supplement lo COLEOPTERISTS BLLLETL\ Vol. 60, Decernber 2006 Made in Uníted States 01 America l Coleopterists Society Monograph Number 5:]9-34. 2ClO6. HISTORY OF SCARABAEDID CLASSIFICATlON BERT KOHLMA::-lN I Universidad EARTH, Apdo. 4442-1000, San José, COSTA RICA [email protected] Abstract A historie analysis of the c1assification of Scarabaeoidea is presented. The analysis comprises a historie comparison of the type and number of characters of the genera and their grouping into higher-Ievel taxa. The analysis is divided into three time periods, including a review of 70 publications 00 scarab systematics (beginning with Linnaeus). The first period (1735-1858) encornpasses an era prior to the publication of Darwin's The Orígtn of Species, and it is, therefore, a classificarion process arguably free of any evolutionary influences. The second period (1859-1949) inc1udes a comparative synthesis on the ciassifícation of the Scarabaeoidea based on faunistic and taxonomic works. The third period (1950-2006) summarizes and analyzes classifications influenced by phylogeneric theories and is based on monographs, faunal studies, keys for regional fauna, papers on cornparative rnorphology, and studies specifically devoted to the understanding of evoluüonary relationships and processes in the Scarabaeoidea. A large problem conceming the classification and phylogeny still rernains because rnost studies do not consider all the diversity of the Scarabaeoidea in a single analysis. Scarabs havc always attracted the attention of observers. In ancient times, the Egyptians worshipped the dung-rolling beetle Scarabaeus, and they eonsidered it to be the personification of Khepri, the Sun-Beetle God (Cambefort 1994). Later, in Linnaean times, the group received much attention (Linnaeus 1735, 1758; Geoffroy, 1762; Scopoli, 1763; Fabricius, 1775, 1792; Degeer, 1783; Olivier, 1789; Gmelin, 1790; Latreille, 1796) thus building a long and solid tradition of c1assification. The act of c1assifying individuals and objects is a natural activity of the human mind. In arder to accomplish this, groups of individual s or objects are established that possess common characteristics. and each group is named individually. Thus, c1assifying, undertaken many times in an implicit way, allows us to synthesize infonnation and obtain a global vision of thc ensemble. The placement of an entity in a group allows us to determine 1 its characteristics, predict its behavior, and know how to treat it. The c1assification process began early in the history of biology, as exemplified by the work of Aristotle (Nelson and ¡ Platnick 1981; Llorente 1990; Jahn 2000). According to Mayr (1969), a c1assification is a communication system, and the best system is one that combines thc greatest amount ofinformation and retrieves it with the greatest ease. It is precisely within this frame of 1 reference that 1 examine this process by analyzing the development of classification systems and concepts within the Scarabaeoidea. In this review, 1 divide the history of scarabaeoid c1assification into three time periods (Kohlmann 1984; Kohlmann and Morón 2003), including 70 pub1icarions on scarab systematics (beginning with Linnaeus [1735]) (Table 1). The analysis comprises a historie comparison of the type and number of characteristics of the genera and their grouping into higher-level taxa. The first period (1735-1858) encompasses an era prior to the publication of Darwin's The Origin o/ Species (1859) and it is, therefore, a c1assification process arguably free of any 19 20 COLEOPTERISTS SOCIETY MONOGRAPH NUMBER 5. 2006 evolutionary influences. The second period (1859-1949) ineludes a comparative synthesis on thc ciassification of the Scarabaeoidea based on faunistic and taxonomic works. The possible influence of Darwin's theories is discussed. New rnorphological characters were used in addition to the characters traditionalIy used. However, family-level characters and their application were largeJy unchanged since the middle of the 19th century. Relationships between groups were depicted as dendrograms by Sharp and Muir (1912) and Paulian (1945) bu! without c1ear references to evolutionary theories. Classifications of this period were more complex than thc preceding ones because of the addition of nearly 11,000 new species discovered during 90 years of world exploration. For the third period (1950--2006), 1 summarize and analyze c1assifications that were influenced by Hennig's (1950) seminal phylogenetic work. This section of the analysis is based 00 monographs, faunal studies, keys for regional faunas, papers on comparative morphology, and studies specifically devoted to the understanding of evolutionary relationships and processes in the Scarabaeoidea. Hennig's proposals were mostly ignored until the publication, in 1966, of the revised English version of his 1950 work. One can detect the influence of evolutionary theories on these studies as well as a progressive reflection of relationships in the proposed c1assifications, mainly through the use of cladistic and phenetic approaches. More recently, molecular approaches have also been used to examine the classification and the evolution of scarabs. Still, the main problem remains that most studies do not consider al l thc diversity of the Scarabaeoidea in one, single analysis. Historical, economic, and sociopolitical factors have influenced the development of systematics during thc 20th century. American authors have usually supported classifications that go against excessively multiplying the number of families in Scarabaeoidea, although Iately they have adopted the 13-family classification of Lawrence and Newton (1995). European authors have tended to favor c1assifications with as many as 25 families, as proposed by Paulian (1988) and Ba1thasar (1963). Classification studies undertaken by Clarke Scholtz and his collaborators have a stronger methodological basis (e.g., Scho1tz 1990; Browne and Scho1tz 1995), and Lawrence and Newton (1995) accepted their proposal of 13 families, with the exception of splitting of Geotrupidae. Lawrence and Newton's (1995) classification still has problems to be solved in the family Scarabaeidae involving the subfamilies Scarabaeinae, Aphodiinae, Melolonthinae, Dynastinae, Rutelinae, and Cetoniinae. However, Jameson and Ratcliffe (2002) pro po sed this c1assification as a contemporary option to frame the study of the Scarabaeoidea. Pre-Darwinian: 1735-1858 Undoubtedly, the initial history of the systematics of scarabaeids is strongly intertwined with the great contributions that Linnaeus made to the classification systcm in general and his group definition in particular. Linnaeus (1735) provided a secure footing to scarab c1assification by recognizing as the essential character of the genus Scarabaeus the existence of an antenna in the form of a "cleft club". He also mentioned as a character the absence of horns, but he eliminated this character in the following editions when he realized this was incorrect. He described the genus Lucanus as having mobile, hard, branched horns and capitate, foliaceous antennae. Curiously enough, and without giving any explanation in subsequent editions, he inc1uded the stag beetle in Scarabaeus, and it was Scopoli (1763) who later resurrected Lucanus which Linnaeus later reaccepted. COLEOPTERISTS SOCIETY MO~OGRAPH NUMBER 5, 2006 21 The name Scarabaeus has an obscure origino According to Cambefort (1994), two words exist in Greek to Dame scarabs: K~PGt~O; (crab, locust) usually designates thc stag beetles and K:l:u8ctpo,; (scarab) the sacred beetles. These words were used by Aristotle 1 (Cambefort 1994). At the same time, these words show consonan tal similarities with the Sumerian word kharub (smal1 beast, ant, larva) and the Egyptian word kheprer OI khepri (scarab). In his Tenth Edition of the Systema Naturae, Linnaeus (1758) established a hierarchical structure for classifying and a system for binomial nomenclature. Thus, a solid frame was established for the taxonorny of the Scarabaeoidea. The first practice, initiated by Linnaeus, 'lilas based on thc use of a single character and then slowly increased the number of characters in order to define a genus. Linnaeus (1735) characterized Scarabaeus by its lamellate antennae, and, in 1758 (Linnaeus 1758), he proposed a second character, the presence of dentate protibiae, to unify species in the genus. This marked thc start of the abandonment of strict adherence to the Aristotelian philosophy of classification, where only single characters were used to defme the "esscncc' of a genus. This character-multiplying practice was reinforced by Fabricius (1775) and Degeer (1783), and Latreille (1796) greatly expanded upon it. Contrary to Mayr's opinion (1982), which considered Linnaeus to be the first person to establish a diagnosis, it was Latreille who first established the conceptual difference between a description and a diagnosis, although he referred to this latter term as "habitual characters". At the same time that the practiee of using multiple eharaeters took place, a search for characters that might better reflect the "natural order" and "essencc" of genera began. Accordingly, Linnaeus (1735) and tben Scopoli (1763) used thc form of the antenna as an essential character for the formation of groups. Later, Fabricius (1775) proposed that mouthparts represent a more natural way of establishing classificaticns, because,
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    Review of Literature and the Biology of the Australian Ceratocanthidae (Insecta: Coleoptera) by Dr Trevor J

    Calodema Volume 7 (2006) Review of literature and the biology of the Australian Ceratocanthidae (Insecta: Coleoptera) by Dr Trevor J. Hawkeswood* *PO Box 842, Richmond, New South Wales, 2753, Australia (www.calodema.com) Hawkeswood, T.J. (2006). Review of literature and the biology of the Australian Ceratocanthidae (Insecta: Coleoptera). Calodema, 7: 8-11. Abstract: The literature pertaining to the Australian Ceratocanthidae (Coleoptera) is reviewed. The information available indicates that very little is known about the three Australian species, all of which occur in Queensland. Introduction The family Ceratocanthidae ( = Acanthoceridae) is found in the tropics and subtropics of the world and is represented by about 40 genera and 320 species (Ballerio, 2000; Howden & Gill, 2000; Grebennikov et al., 2002) but this number is likely to rise in the coming years as a result of undescribed species being discovered (Grebennikov et al., 2002). The Ceratocanthidae exhibit a pantropical distribution, with few species living in the temperate regions of the Americas, South Africa and south-east Asia (Grebennikov et al., 2002). Taxonomically, it is widely considered as a separate family of the superfamily Scarabaeoidea and appears to be more closely related to the Hybosoridae and Ochodaeidae (Lawrence & Newton, 1982; Browne & Scholtz, 1999; Grebennikov et al., 2002, 2004; Grebennikov & Scholtz, 2004) and the Trogidae (Crowson, 1955). Ceratocanthid beetles are characterised by having a strongly convex body which allows the mouth-parts, abdomen and parts of the legs to be completely concealed when they roll themselves into a pill-shaped form or little compact ball, hence the common name of pill scarab beetle or ball beetle for the group.