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, according to hirn, eating is the most important activity for an insect, and he restruetured the entire ínsect c1assification systern on this basis. Not until Edition XIII of Systema Naturae (Gmelin 1790) was the usage accepted of having more than two characters for defining genera. After Linnaeus, other ímportant characters appear: bent or geniculate antennae (as used by Leach [1815] to separate lucanids and passalids) and the position of the spiracles (as used by Burmeister [1842]). Burmeister (1842) was also thc first person to propase a general classification of insects based on the type of metamorphosis. Subsequently, Erichson (1847) introduced a detailed characterbased classification system for scarabs, which is the basis for the one still used today. Erichson (1847) establishcd two lines, Pleurosticti and Laparosticti. The first group 'lilas characterised by having the abdominal spirac1es on the dorsal part of the ventral segments, whereas the second group has them situated on the intersegrnental membrane that joins the dorsal and ventral abdominal segmenta and are, therefore, covered by the elytra. Erichson (1848) was also the first author to propase antennae as olfactory organs. Finally, Lacordaire (1856) recognized such important characters as transverse eoxae and the laek of foretarsi. The second practice in the establishment of a c1assification system was the inerease in the number of groups. As airead y mentioned, Linnaeus (1735) recognized the existence oftwo genera, Scarabaeus and Lucanus. Fabrieius (1775) recognized six genera, and Latreille (1797) recognized thirteen genera. A third trend in the pre-Darwinian era was the grouping of genera based on shared charaeters. Linnaeus (1735) began creating higher-level taxa, but he did 22 COLEOPTERISTS SOCIETY MONOGRAPH NUMBER 5, 2006
not recognize them as entities. For instance, in 1735 he established a group of genera characterized by having truncate antennae: Scarabaeus, Dermestes, Cassída, Coccinella, Gyrinus, Necydalis, and Attelabus, Fabricius (1775) established for the first time a group of genera defined by the character antennis clava lame/atta and incorporated into ir the following genera: Lucanus, Scarabaeus, Trox, Melolontha, Trichius and Cetonia. However, it was with Latreille (1797) that an important step was taken. He was the first author to establish higher groups in thc lamellicorns by recognizing two (unnamed) families: "Family I" (representing the stag and bess beetles" and "Family II" (encompassing the scarab beetles). Duméril (1800) named these higher-Ievel groups "Serricomes" or "Priocéres" and "Lamellicornes" or "Pétalocéres", respectively. By 1802, Latreille proposed three higher-Ievel groups: "Coprophages", "Géotrupins", and "Scarabéides". This trend continued, and Leach (1815) recognized four "tribes": "Coprides" including families Coprida and Aphodida, "Scarabaeidaes" with no included family, "Geotrupides" including families Geotrupida and Cetonida, and "Lucanides" including families Lucanida and Passalida. However, Latreille (1817) lumped everything into one group, the lamellicorns. By thc end of this first period, Lacordaire (1856) recognized 13 higher-Ievel groups. Whereas Leach (1815) called the more inclusive groups "tri bes" and divided them into "families", Lacordaire (1856) called the more inclusive groups "familles" and divided them into "tribus". Within the "Pectinicornes" he recognized the "Lucanides" and "Passalides", and within the "Lamellicomes" he recognized four groups within the "légion Pleurostictiques": "Mélolonthides", "Rutélides", "Dynastides", "Céronides", and seven groups within the "Iégion Laparostictiques": "Coprides", "Aphodiides", "Orphnides", "Hybosorides", "Géotrupides", "Trogides", and "Glaphyrides". This classification, which was proposed 150 years ago, is still large1y used by taxonomists today. Seemingly idiosyncratic ciassifications were also proposed, as was the ease of MacLeay (1819). MacLeay invented the quinarian system (a system based on five elements) and believed the systern reflected "the natural order given by God." Thus, each family was composed of five genera that containcd five species each, so each family was formed by 25 speeies. The most prominent feature of the systern was its cireularity. "The Circular Systern". also known as "Quinarism" or "Quinarian Systern", a name derived from the specia1 significance thought to be possessed by the number five, held that all organisms have been created by one plan, and this plan is founded on the principle of a series of affinities returning unto thernselves, whieh can only be represented by a eircle. Following this pattem, MacLeay (1819) established the "Lamellicornes Petalocera Saprophaga" which included tho following five families: Geotrupidae, Scarabaeidae, Aphodiidae, Trogidae, and Dynastidae, and the "Lamellicornes Petalocera Thalerophaga" which included five families: Rutelidae, Cetoniidae, Glaphyridae, Melolonthidae, and Anoplognathidae. Samouelle (1819) in his Use -fuí Campendium (published in June) already applied the suffix "ídae" for family narnes; MacLeay's work (published in November of that same year) fol1owed and popularized this same usage. In a relatively short period oftime (1735-1858) a great development ofhigherIevel searab systematics took place. The number and type of characters multiplied quickly frorn one to many. Groupings of taxa evolved from a loose connection of genera to complicated systerns full of hierarchies and families. Key authors like Linnaeus, Fabrieius, and Latreille founded our classification system with their COLEOPTERISTS SOCIETY MONOGRAPH NUMBER 5, 2006 23
ideas and knowledge. By 1856, over 150 years ago, a classification system was established that any modern specialist can recognize.
Post-Darwinian and Pre-Hennigian: 1859-1949 Darwinian ideas of evolution were launched in 1859 with the publication of "The Origin of Species' (Darwin 1859), and this set the stage fOI a new method of understanding biodiversity. Darwin. through his theory of evolution by cornmon descent, supplied the explanation ofwhy the members of a taxon are more similar to each other than they are to members of other taxa (Mayr and Ashlock 1991). Even so, Frorn 1859 to 1949 the general c1assification ofthe Scarabaeoidea did not change rnuch relative to earlier changes presented by Burmeister (1842, 1844, 1847) or Lacordaire (1856). At the family level, Darwinian ideas did not seern to have strong influences with the exception of the appearance of the first scarab evolutionary trees proposed by Sharp and Muir (1912) and Paulian (1945). However, searabaeoid c1assifieations beearne more cornplicated at the generic and family levels due to the enormous accumulation of new genera and species. The expansion of knowledge was driven by the exploration of distant parts of the world through the auspices of economic policies of expanding colonial powers. The landslide of new species resulted in thc creation of new genera to accommodate them without giving too much thought to the organization of upper hierarchies. This phenomenon is still evident in groups like the Melolonthinae, Rute1inae, and Cetoniinae, which are particu1arIy speciose groups. A descriptive approach was the obvious logical priority, whereas the integration of all this information became a Iong-term objective after the descriptive work was finished. A valiant attempt, in the best Linnaean tradition, at concentrating all the described species in a monumental catalog is the Coleopterorum Catalogus (19]0-1974), cornpiled by nine European specialists under the coordination of Sigmund Schenkling. During this period several new approaches appeared, such as the wellsurnmarized attempt by Mulsant and Rey (1871), which used larval characters to establish the classification, and comparative studies of male genitalia (Sharp and Muir 1912) to establish evolutionary relationships among scarabaeoids. C1assifications proposed prior to 1859, based 00 the grouping of similar species, continued to be accepted in the post-Darwiriian era probab1y because similar species are usualIy descendants from a common ancestor (Mayr ] 982; Mayr and Ashlock 1991), and, as Darwin stressed again and again, "a11 true c1assification is genealogical" (1859: 420). Although Darwinian views of evo1ution were set into motion during this time, there was not a means of methodologically employing and testing new ideas in classification.
Post-Hennigian: 19511--2006 FroID ]950 to 2006, the descriptive trend increased, and taxonomists sought help from other disciplines like mathernatics, statistics, genetics, and molecular biology in order to explain and describe diversity. The description process became more analytical, and paradigms shifted from evolutionary systematics to phylogenetic systematics. New information, like cytology, food habits, ecologica1-geographica1 distributions, and DNA sequencing were employed. This entire process was enonnously aided by the development of computers. However, the new paradigrn in systematic methodology for this period is I Hennig's (1950) j 24 COLEOPTERISTS SOCIETY MONOGRAPH NUMBER 5, 2006 concept of the shared possession of uniquely derived characters and its application to the practice of classification. During the 19508, most studies relied on intuition and non-repeatable methods, but they were based on the morphology of the groups under study. It was not until Endródi (1966) that a "primitive" phylogenetic analysis was presented for the first time based on concepts similar to those proposed by Wagner (1961) for the phylogeny of ferns. However, Endródi's five-family classification (Scarabaeidae, Melolonthidae, Trogidae, Passalidae and Lucanidae) was not adopted. Similarly, Medvedev's (1976) and Iablokoff-Khnzorian's (1977) classifications, which were based on comparative analyses, were not generally accepted. Paulian's (1941) and Balrhasar's (1963) classifications were based on a multip1ication of families by e1evating the rank of subfamilies to families rather than analyses. This 1ast approach is still being emulated in Europe (Martín-Piera 2000). Studies by the South African schoo1 are the only ones that uti1ized a solid, rnethodo1ogica1 base. Scho1tz and collaborators (e.g., Scholtz 1990; Scholtz and Chown 1995; Browne and Scholtz 1995, 1998, 1999; Grebennikov and Scholtz 2004; Grebennikov el al. 2004) have undertaken a comprehensive morphological research program complernented by bio1ogical and paleontological information that proposes a scarabaecid phylogeny of 13 families, although the latest study (Scholtz and Grebennikov 2005) considcrs 14 families. The c1assification proposed by Lawrence and Newton (1982, 1995) is basically the same as the c1assification of Brown and Scholtz (1995), except that the former do not recognize thc Bolboceratidae. Jameson and Ratcliffe (2002) have used the Lawrence and Newton system (1995) (a1beit with sorne hesitation on their part) as the basis for their study of the North American Scarabaeoidea. However, there are still problems with these post-Hennigian analyses in solving the Scarabaeinae, Aphodiinae, Melolonthinae, Dynastinae, Rutelinae, and Cetoniinae, groups that in many cases are taxonomic "catch-alls".
Future of Scarabaeoid Classification We have witnessed during the last decade a strong and renewed impetus in the c1assification of the Scarabaeoidea. Phylogenetic analysis has emerged as the tool of choice for recent studies. However, an ec1ectic approach to c1assification should not be discarded, and other methods should always be used (e.g., Pretorius and Scholtz 2001) in order to obtain independent support, comparison, or validation for a particular classification. Onc of the most prevalent problems currently in systematics is that phylogenetic analyses are being conducted on paraphyletic groups (e.g., limited genera from a tribe or taxa from a specific geographic area) that purport to depict evolution of a natural group. Clearly, this is not what a natural system endeavors ro build, a systern that reflects most closely the actual relationship of the species. However, the definition of a natural classification is still open for debate as can be gleaned from Mayr's change in altitnde. Mayr and Ashlock (1991: 264-265) first proposed that an evolutionary systematics approach had an advantage over a phylogenetic approach. However, Mayr (1997: 93) indicated that the "two approaches can continue to exist side by side". Conversely, Schuh (2000) presented a different opinion and considered that a phylogenetic approach is presently thc method that dominates systematic thinking. A c1assification of the Scarabaeoidea requires analysis of the totality of the groups, a truly Herculean task, that is probably beyond the reach of a single I COLEOPTERISTS SOCIETY MONOGRAPH NUMBER 5, 2006 25 persono Perhaps the future of such an endeavor lies in joining [orces in an all- j encompassing effort to establish a classification of the Scarabaeoidea. This would require the forrnation of an international consortium of specialists where each person wouId participate in a standardized effort to create such a classification. One can cite as 1 possible benefits of a "global" scarab c1assification a more efficient transmission of knowledge worldwide, comprehensive analyses of widely distributed groups and avoidance of establishing paraphyletic taxa, access to rare or difficult material, use of a 1 unified set of characters, etc. This synthesis of knowledge would certainly allow the resolution of many problems that is not currently possible. 1
Acknowledgments I I would like to thank the library personnel of the following universities and institutions for their help in locating references: Harvard University (Cambridge, MA), National Museum of Natural History (Washington, D.C.), Martin-Luther-Universitat (Halle-Wittenberg), Universitát des Saarlandes (Saarbrücken), Technische Universitát Dresden (Dresden). Institut für Geschichte der Medizin, Naturwissenschaft und Technik, Brnst-Haeckel-Haus. Friedrich-Schiller-Universitát (Jena), Deutsches Bntomologisches Instituí (BberswaldeFinow), and Hurnboldt-Universitát (Berlín). T am also grateful to the following persons for having sent me additional bibliographic materials: Jorge E. Llorente (Mexico City); Brett C. Ratcliffe (Lincoln, NE); Henry F. Howden (Ottawa), and Patrick Amaud (París). l am also grateful to Brett Ratcliffe, Mary Liz Jameson, Sara Thomton, Jane Yeomans, and Stephanie Larson for their careful revision of the text. Last but not least, J would Iike to thank the invaluable help and comments given by three anonymous reviewers.
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