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11-27-2004

The Revised Classification for : What the Hell is Going On?

Brett C. Ratcliffe University of Nebraska-Lincoln, [email protected]

Mary Liz Jameson University of Nebraska State Museum, [email protected]

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Ratcliffe, Brett C. and Jameson, Mary Liz, "The Revised Classification for Scarabaeoidea: What the Hell is Going On?" (2004). Papers in Entomology. 25. https://digitalcommons.unl.edu/entomologypapers/25

This Article is brought to you for free and open access by the Museum, University of Nebraska State at DigitalCommons@University of Nebraska - Lincoln. It has been accepted for inclusion in Papers in Entomology by an authorized administrator of DigitalCommons@University of Nebraska - Lincoln. Published in SCARABS 15 (November 2004), pp. 3-10. Published by Coleopterists Society; online at http://www-museum.unl.edu/research/entomology/Scarabs-Newsletter.htm

The Revised Classification for Scarabaeoidea: What the Hell is Going On? by Brett C. Ratcliffe and Mary Liz Jameson Systematics Research Collections W436 Nebraska Hall, University of Nebraska Lincoln, NE 68588-0514 bratcliff[email protected] and [email protected]

Considering the turmoil and vast changes in the classification of the superfamily Scarabaeoidea during the last 20 years, particularly in North America, we were asked to provide an update for the readers of Scarabs wherein we offer our perspectives. Much of what follows is extracted from our scarabaeoid introduction in American (Jameson and Ratcliffe 2002). By the time this overview is printed, there may have been more changes in the classification because of the rapidly accumulating evidence supporting new hypotheses.

These rapid changes are a result of intensified study of the family groups using both traditional morphological evidence combined Brett and Mary Liz with increasingly insightful The superfamily Scarabaeoidea molecular studies. While possibly is a large, diverse, cosmopolitan disruptive now, these new studies group of beetles. As a personal are exciting because, for the aside (and, of course, with no first time, we are establishing bias), these are probably the finest the higher classification of the beetles in the world. Scarabaeoids Scarabaeoidea based on evidence are adapted to most habitats, and facts rather than intuition. and they can be fungivores, This research confirms many of herbivores, necrophages, our hypotheses of classification coprophages, saprophages, and Ed. Note: Dave Hawks is but also clearly refutes others. sometimes carnivores. They are conducting DNA studies Be on the lookout for future widely distributed around the on the Scarabaeoidea publications by Team Scarab and globe, even living in the Arctic David Hawks! in burrows. Some scarabs exhibit parental care and sociality. Page 3 Some are myrmecophilous, is not resolved and continues termitophilous, or ectoparasitic. to be debated. The hierarchical Many possess extravagant horns, level of families and subfamilies others are able to roll into a within the Scarabaeoidea is in compact ball, and still others disarray and remains unresolved. are highly armored for inquiline In most U.S. literature prior to life. A very few are occasionally the 1970s (e.g., Arnett 1968), the agricultural pests that may Scarabaeoidea included three destroy crops (even beetles have families: Passalidae, Lucanidae, to eat!) while others are used in and . This three- the biological control of dung family system of classification was and dung flies. Scarabaeoids are the “traditional” North American popular beetles due to their large system and had several practical size, bright colors, and interesting and conceptual advantages. First, natural histories. Early Egyptians it recognized the shared, derived revered the scarab as a god, characters that unite subfamilies Jean-Henri Fabre Jean-Henri Fabre studied their within the family Scarabaeidae. behavior, and Charles Darwin Second, it provided a classification used observations of scarabs in his system that allowed easy retrieval theory of sexual selection. of hierarchical information based on the fact that subfamilies were What characterizes a part of the family Scarabaeidae scarabaeoid? (e.g., life history, morphology, larval type). Phylogenetic research The antennal club is lamellate, the indicates, however, that the family prothorax is often highly modified Scarabaeidae (in the traditional for burrowing (with large coxae, sense) is not a monophyletic usually with concealed trochantins group. Accordingly, most workers and closed cavities), the protibia now follow the 12-family system is usually dentate with a single established by Browne and Scholtz spur, the wing venation is reduced (1995, 1999) and Lawrence and and with a strong intrinsic spring Newton (1995). This system mechanism for folding, tergite 8 places emphasis on the differences forms a true pygidium and is not that separate taxa rather than concealed by tergite 7, there are the similarities that unite them. four Malpighian tubules, and larvae Whereas families, subfamilies, are scarabaeiform (cylindrical, c- and tribes in the staphylinoids and shaped). curculionoids are being combined because of shared characters (thus What is the current status of the increasing efficient data retrieval), classification? the scarabaeoids are being split into numerous families because Monophyly of the superfamily of supposed differences (thus, in Scarabaeoidea is well-founded our view, decreasing information and undisputed (Lawrence and retrieval, at least in the short term). Britton 1991). The sister group The debate concerning scarabaeoid Page 4 for the Scarabaeoidea, however, classification systems illustrates the weak phylogenetic foundation of approximately 21,000 species are the superfamily. This problem is in the subfamilies Melolonthinae, the result of a number of factors Dynastinae, Rutelinae, and including (1) lack of thorough Cetoniinae (the “higher” scarabs). study of all scarabaeoid taxa, (2) Only a few phylogenetic analyses lack of diagnostic characters for have addressed relationships of all taxa, (3) lack of phylogenetic pleurostict subtribes, genera, or study of all taxa, (4) prevailing species (Ratcliffe 1976; Ratcliffe and philosophies regarding categorical Deloya 1992; Jameson 1990, 1996, levels, and (5) emphasis in 1998; Jameson et al. 1994; Krell research on the less speciose 1993; Montreuil 2000; Paucar 2003; groups of scarabaeoids and lack Smith 2003), and only one analysis of research on the more speciose has been conducted to address groups (such as the subfamilies tribal or subfamilial relationships of Scarabaeidae including the (Browne and Scholtz 1999). Melolonthinae, Rutelinae, Dynastinae, Aphodiinae, and Historically, the superfamily Cetoniinae). Scarabaeoidea was divided into Charles Darwin two generalized groups based Within the Scarabaeoidea there on the position of the abdominal is a disparity in the knowledge spiracles; the Laparosticti and between less speciose basal Pleurosticti. Pleurostict scarabs lineages and the more speciose were characterized by having groups of “higher” Scarabaeidae. most of the abdominal spiracles For example, the family situated on the upper portion of the includes approximately sternites (Ritcher 1969; Woodruff 300 species in four genera. 1973) and included taxa whose Excellent revisionary, larval, and adults feed on leaves, flowers and phylogenetic studies are available pollen, and whose larvae feed for this group (Baker 1968; Scholtz primarily on roots and decaying 1982, 1986, 1990, 1991, 1993; wood. Laparostict scarabs, on the Scholtz and Peck 1990). Excellent other hand, were characterized monographs are also available for by having most of the abdominal the approximately 600 species of spiracles located on the pleural Geotrupidae (Howden 1955, 1964, membrane between the tergites 1979, 1985a-b, 1992; Howden and sternites (Ritcher 1969) and and Cooper 1977; Howden and included taxa whose adults and Martínez 1978) and the Trogidae larvae feed on dung, carrion, (Vaurie 1955), and these provide hides, and feathers. The position the foundation for addressing of the spiracles, however, is not a relationships within this group. consistent character (Ritcher 1969), In comparison, the family and, in recent years, subfamilies Scarabaeidae (sensu Lawrence and tribes that were once included and Newton 1995) includes in the Laparosticti have been raised approximately 91% of the species to higher taxonomic status (family (ca 27,800) of Scarabaeoidea. and subfamily, respectively). Within the Scarabaeidae, Page 5 The composition of the characters or suites of characters. Scarabaeoidea remains a topic of debate. Lawrence and Newton Underlying the classification (1995) proposed 13 families (12 problem is, of course, the fact that found in the Nearctic, Belohinidae we are dealing with constructs is Madagascan), and Scholtz that are 200 years old and that and Browne (1996) and Browne pre-date evolutionary theory. and Scholtz (1995, 1998, 1999) Linnaean classifications were proposed 13 families (all Nearctic, based on overall morphological including Bolboceratidae; similarity rather than shared, Belohinidae was not addressed). We derived characters. Thus, some follow, with some hesitation, the groups within the scarabaeoids system of Lawrence and Newton are not monophyletic lineages; (1995) and treat the Scarabaeoidea instead, they are groups that were as including 12 Nearctic families created historically because they (11 of which were previously superficially resembled each other. considered subfamilies of the family Our system of classification needs Scarabaeidae, and one of which was to convey information and concepts previously considered a subfamily and allow for easy retrieval of A “New” Scarab of the Lucanidae). Our reluctance information. Whether a certain to accept elevation of some new taxon is classified at the level of families within the Scarabaeoidea family or subfamily may be trivial stems from the fact that: 1) there if we can continue to convey the have been no comprehensive needed information. We remain taxonomic treatments of all higher apprehensive that the trend of categories of scarabaeoids (families elevation to many families within and subfamilies) and, 2) there the Scarabaeoidea will result, at are few comprehensive, rigorous, least in the short term, in a net loss phylogenetic analyses of higher in retrievability of information. scarabaeoid groups and, thus, a lack of synapomorphic characters Despite the considerable that establish a basis for uniform debate, phylogenetic analyses of familial and subfamilial levels. We scarabaeoid higher categories prefer to see clades delimited by are on-going and their results shared derived characters before bring us closer to understanding the elevation of certain taxa to relationships of the groups. A family level. Despite our reluctance preliminary “total evidence” to accept this classification system, phylogenetic analysis of 13 we have little basis for disputing families of Scarabaeoidea the validity of current taxonomic (excluding Belohinidae, including conclusions other than the fact Bolboceratidae) and most of the that some of these taxonomic subfamilies was conducted using conclusions have been based on 134 adult and larval characters narrow taxonomic frame-works (Brown and Scholtz 1999). Results (only scarab taxa from certain of this analysis showed that geographic regions rather than all the superfamily Scarabaeoidea Page 6 scarab groups) or based on few is comprised of three major lineages: the glaresid lineage relationships of all of these taxa. that consists of only the family However, most hypotheses Glaresidae; the passalid lineage generally consider the Aphodiinae that consists of two major lines- and Scarabaeinae as the sister -a glaphyrid line (containing group to the Melolonthinae, Glaphyridae, Passalidae, Dynastinae, Rutelinae, and Lucanidae, Diphyllostomatidae, Cetoniinae. The former Trichiinae Trogidae, Bolboceratidae, and and Valginae are now considered Pleocomidae), and a geotrupid tribes of the Cetoniinae. line (containing Geotrupidae, Ochodaeidae, Ceratocanthidae, The family Scarabaeidae is and Hybosoridae); and the scarab sometimes referred to as the family lineage (containing Aphodiinae, Melolonthidae, especially by some Scarabaeinae, Orphninae, of the Latin American workers. Melolonthinae (sensu lato), In this usage, the family includes Rutelinae, Dynastinae, and the subfamilies Melolonthinae, Cetoniinae). Euchirinae, Phaenomeridinae, Dynastinae, Cetoniinae, The past thirty years have seen Glaphyrinae, and Systellopodinae many changes and debates in (Endrödi 1966) whereas the And Another... the classification of the family Scarabaeidae refers to everything Scarabaeidae. In the “traditional” else except Passalidae, Lucanidae, North American system, the and Trogidae. This classification category Scarabaeidae has been is not in wide use today and treated as including the all is incorrect. The family group scarabaeoid families except the names Rutelinae and Dynastinae Passalidae and Lucanidae. Old were established by MacLeay in World scarab workers have tended 1819, and the family group name to split the Scarabaeidae into Melolonthinae was established by several families. Samouelle in 1819. However, the family group name Cetoniinae was While the debate continues, we established a few years earlier in follow Lawrence and Newton 1815 by Leach. Thus, the family (1995) and consider the family group name Cetoniidae has priority Scarabaeidae to include the over Melolonthidae. Therefore, if subfamilies Aphodiinae, one wants to consider all of these Scarabaeinae, Melolonthinae, subfamilies in the same family Dynastinae, Rutelinae, and (exclusive of Scarabaeinae, which Cetoniinae. Several smaller was established by Latreille in subfamilies that are not present 1802), then the valid name would in the Nearctic region are also be Cetoniidae! Accordingly, included in the Scarabaeidae: the family name Scarabaeidae Orphninae, Phaenomeridinae, (including Melolonthinae, Pachypodinae, Allidiostomatinae, Scarabaeinae, Dynastinae, Dynamopodinae, Aclopinae, Cetoniinae, etc.) is the correct and Euchirinae. No phylogenetic family group name for these taxa analyses have addressed the and not Melolonthidae. Page 7 At the family level, classification Summary of the Families and of the world Scarabaeidae is Subfamilies of Scarabaeoidea of variably known. The classification the United States, Canada, and of the world Dynastinae is Neartic Mexico fairly well established due to the work of Endrödi (1985). Most Melolonthinae, Rutelinae, Lucanidae: Lucaninae, Nicaginae, and Cetoniinae remain poorly Syndesinae known taxonomically, and many Diphyllostomatidae New World genera cannot be Passalidae reliably identified. Classification Glaresidae of the Scarabaeinae (Hanski and Trogidae Cambefort 1991) and Aphodiinae Pleocomidae (Dellacasa 1987, 1988a, 1988b, Geotrupidae: Bolboceratinae, 1991, 1995) are fairly well Geotrupinae established. The of the Ochodaeidae: Ochodaeinae, North American scarab beetles Chaetocanthinae is relatively stable although Hybosoridae no one volume is available for Ceratocanthidae identification. Regional works Glaphyridae are sometimes the best sources Scarabaeidae: Aphodiinae, for identification of Nearctic Scarabaeinae, Melolonthinae, scarab beetles. The family Rutelinae, Dynastinae, Cetoniinae Scarabaeidae includes about 91% of all scarabaeoids and includes about 27,800 species worldwide. Acknowledgment Within the Scarabaeidae, the Aphodiinae and Scarabaeinae This project was supported by an include approximately 6,850 NSF/PEET grant (DEB 0118669) to species worldwide (about 22% M. L. Jameson and B. C. Ratcliffe. of scarabaeoids and 25% of Scarabaeidae). The subfamilies Orphninae, Melolonthinae, Literature Cited Dynastinae, Rutelinae, and Cetoniinae include approximately ARNETT, R. H., JR. 1968. The Beetles of the United States. A Manual for Identification. The 20,950 species (about 69% American Entomological Institute, Ann Arbor, of scarabaeoids and 75% of MI. 1,112 pp. Scarabaeidae). BAKER, C. W. 1968. Larval taxonomy of the Troginae in North America with notes on bi- ologies and life histories (Coleoptera: Scarabae- idae). United States National Museum Bulletin, 279: 1-79.

BROWNE, D. J. and C. H. SCHOLTZ. 1995. Phylogeny of the families of Scarabaeoidea (Co- leoptera) based on characters of the hindwing articulation, hindwing base and wing venation. Systematic Entomology, 20: 145-173. Page 8 BROWNE, J. and C. H. SCHOLTZ. 1998. HOWDEN, H. F. 1955. Biology and taxonomy Evolution of the scarab hindwing articulation of North American beetles of the subfam- and wing base: A contribution toward the ily Geotrupinae with revisions of the genera phylogeny of the Scarabaeidae (Scarabaeoidea: Bolbocerosoma, Eucanthus, Geotrupes and Coleoptera). Systematic Entomology, 23: 307- Peltotrupes (Scarabaeidae). Proceedings of the 326. United States National Museum, 104: 151-319.

BROWNE, J. and C. H. SCHOLTZ. 1999. A HOWDEN, H. F. 1964. A Catalog of the Co- phylogeny of the families of Scarabaeoidea leoptera of America North of Mexico. Family (Coleoptera). Systematic Entomology, 24: Scarabaeidae. Subfamily: Geotrupinae. United 51-84. States Department of Agriculture, Agriculture Handbook 529-34a. 17 pp. DELLACASA, M. 1987. Contribution to a world-wide catalogue of Aegialiidae, Apho- HOWDEN, H. F. 1979. A revision of the diidae, Aulonocnemidae, Termitotrogidae Australian Blackburnium Boucomont (Coleoptera Scarabaeoidea). Memorie della (Coleoptera: Scarabaeidae: Geotrupinae). Società Entomologica Italiana, 66: 1-455. Australian Journal of Zoology, Supplementary Series, 72:1-88. DELLACASA, M. 1988a. Contribution to a world-wide catalogue of Aegialiidae, Apho- HOWDEN, H. F. 1985a. A revision of the South diidae, Aulonocnemidae, Termitotrogidae American genus Parathyreus Howden and (Coleoptera Scarabaeoidea) Part II. Memorie Martínez (Coleoptera: Scarabaeidae: Geotrupi- della Società Entomologica Italiana, 67: 1-229. nae). Coleopterists Bulletin, 39: 161-173.

DELLACASA, M. 1988b. Contribution to a HOWDEN, H. F. 1985b. A revision of the South world-wide catalogue of Aegialiidae, Apho- American species in the genus Neoathyreus diidae, Aulonocnemidae, Termitotrogidae Howden and Martínez (Coleoptera, Scarabaei- (Coleoptera Scarabaeoidea). Addenda et dae, Geotrupinae). Contributions of the Ameri- corrigenda (First note). Memorie della Società can Entomological Institute, 21: 1-95. Entomologica Italiana, 67: 291-316. HOWDEN, H. F. 1992. A revision of the Aus- DELLACASA, M. 1991. Contribution to a tralian genera Eucanthus Westwood, Bol- world-wide catalogue of Aegialiidae, Apho- bobaineus Howden and Cooper, Australobolbus diidae, Aulonocnemidae, Termitotrogidae Howden and Cooper and Gilletus Boucomont (Coleoptera Scarabaeoidea). Addenda et cor- (Scarabaeidae: Geotrupinae). Invertebrate Tax- rigenda (Second note). Memorie della Società onomy, 6: 605-717. Entomologica Italiana, 70: 3-57. HOWDEN, H. F. and J. B. COOPER. 1977. DELLACASA, M. 1995. Contribution to a The generic classification of the Bolboceratini world-wide catalogue of Aegialiidae, Apho- of the Australian Region, with descriptions of diidae, Aulonocnemidae, Termitotrogidae four new genera (Scarabaeidae: Geotrupinae). (Coleoptera Scarabaeoidea). Addenda et cor- Australian Journal of Zoology, Supplementary rigenda (Third note). Memorie della Società Series, 50: 1-50. Entomologica Italiana, 74: 159-232. HOWDEN, H. F. and A. MARTÍNEZ. 1978. A ENDRÖDI, S. 1966. Monographie der Dy- review of the New World genus Athyreus Ma- nastinae (Coleoptera, Lamellicornia) I. Teil. cLeay (Scarabaeidae, Geotrupinae, Athyreini). Entomologische Abhandlungen Museum für Contributions of the American Entomological Tierkunde, 33: 1-460. Institute, 15: 1-70.

ENDRÖDI, S. 1985. The Dynastinae of the JAMESON, M. L. 1990. Revision, phylogeny World. Dr. W. Junk Publisher, London. 800 pp. and biogeography of the genera Parabyrsopolis Ohaus and Viridimicus (new genus) (Coleop- HANSKI, I. and Y. CAMBEFORT. 1991. Dung tera: Scarabaeidae: Rutelinae). Coleopterists Beetle Ecology. Princeton University Press, Bulletin, 44: 377-422. Princeton, NJ. 481 pp. JAMESON, M. L. 1996. Revision and phylogeny of the Neotropical genus Cnemida (Coleoptera: Scarabaeidae: Rutelinae). Insecta Mundi, 10: 285-315.

Page 9 JAMESON, M. L. 1998. Phylogenetic analysis of RATCLIFFE, B. C. and A. C. DELOYA. 1992. the subtribe Rutelina and revision of the Rutela The biogeography and phylogeny of Hologym- generic groups (Coleoptera: Scarabaeidae: netis (Coleoptera: Scarabaeidae: Cetoniinae) Rutelinae: Rutelini). Bulletin of the University of with a revision of the genus. Coleopterists Bul- Nebraska State Museum, 14: 1-184. letin, 46: 161-202.

JAMESON, M. L. and B. C. RATCLIFFE. 2002. RITCHER, P. O. 1969. Spiracles of adult Scara- Series Scarabaeiformia Crowson 1960, Super- baeoidea (Coleoptera) and their phylogenetic family Scarabaeoidea Latreille 1802, pp. 1-5. In, significance. I. The abdominal spiracles. Annals Arnett, R. H., M. Thomas, P. E. Skelley, and J. H. of the Entomological Society of America, 62: Frank (eds.), American Beetles, Volume 2. CRC 869-880. Press, Boca Raton, FL. 861 pp. SCHOLTZ, C. H. 1982. Catalogue of the world JAMESON, M. L., B. C. RATCLIFFE and M. A. Trogidae (Coleoptera: Scarabaeoidea). Republic MORÓN. 1994. A synopsis of the Neotropical of South Africa, Department of Agriculture and genus Calomacraspis Bates with a key to larvae Fisheries, Entomology Memoire, 54: 1-27. of the American genera of Rutelini (Coleoptera: Scarabaeidae: Rutelinae). Annals of the Entomo- SCHOLTZ, C. H. 1986. Phylogeny and system- logical Society of America, 87: 43-58. atics of the Trogidae (Coleoptera: Scarabae- oidea). Systematic Entomology, 11: 355-363. KRELL, F.-T. 1993. Phylogenetisch-system- atische Revision des Genus Temnorhynchus SCHOLTZ, C. H. 1990. Phylogenetic trends Hope, 1837 (Coleoptera: Scarabaeoidea: in the Scarabaeoidea (Coleoptera). Journal of Melolonthidae: Dynastinae: Pentodontini). 1. Natural History, 24: 1027-1066. Teil: Phylogenetische Analyse, mit Anmerkunge zur phylogenetisch-sytematischen Methodolo- SCHOLTZ, C. H. 1993. Descriptions of larvae gie. Beitrage zur Entomologie, 43: 237-318. of African Trogidae (Coleoptera), with implica- tions for the phylogeny of the family. African LAWRENCE, J. F. and E. B. BRITTON. 1991. Entomology, 1: 1-13. Coleoptera. The of Australia, 2nd edition, Volume 1, , pp. 543-683. Melbourne SCHOLTZ, C. H. and J. BROWNE. 1996. Poly- University Press, Carlton. phyly in the Geotrupidae (Coleoptera: Scara- baeoidea): a case for a new family Bolbocerati- LAWRENCE, J. F. and A. F. NEWTON. 1995. dae. Journal of Natural History, 30: 597-614. Families and subfamilies of Coleoptera (with selected genera, notes, references and data on SCHOLTZ, C. H. and PECK. 1990. Description family-group names), pp. 779-1006. In J. Paka- of a Polynoncus Burmeister larva, with implica- luk and S. A. Slipínski (eds.), Biology, Phylogeny, tions for phylogeny of the Trogidae (Coleop- and Classification of Coleoptera. Papers Cel- tera: Scarabaeoidea). Systematic Entomology, ebrating the 80th Birthday of Roy. A. Crowson. 15: 383-389. Muzeum i Instytut Zoologii PAN, Warszawa. 1,092 pp. SMITH, A. B. T. 2003. A monographic revision of the genus Platycoelia Dejean (Coleoptera: MONTREUIL, O. 2000. Cladistic systematics of Scarabaeidae: Rutelinae: Anoplognathini). the genus Amphimallon (Coleoptera: Scara- Bulletin of the University of Nebraska State baeidae: Melolonthinae). European Journal of Museum 15: 1-202. Entomology 97: 253-270. VAURIE, P. 1955. A revision of the genus Trox PAUCAR, A. 2003. Systematics and phylogeny in North America. Bulletin of the American of the genus Epectinaspis Blanchard (Coleop- Museum of Natural History, 106: 1-89. tera: Scarabaeidae: Rutelinae) and description of a new genus of Anomalini from Mexico. WOODRUFF, R. E. 1973. The scarab beetles of Coleopterists Society Monographs 2: 1-60. Florida (Coleoptera: Scarabaeidae). Part. I. The Laparosticti (subfamilies: Scarabaeinae, Apho- RATCLIFFE, B. C. 1976. A revision of the genus diinae, Hybosorinae, Ochodaeinae, Geotrupi- Strategus (Coleoptera: Scarabaeidae). Bulletin nae, Acanthocerinae). of Florida of the University of Nebraska State Museum, 10: and Neighboring Land Areas, 8: 1-220. 93-204.

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