Towards a temporal framework for "inordinate fondness": reconstructing the macroevolutionary history of beetles (coleoptera).
Key words: Evolution, molecular phylogenetics, nodal support, timetree.
Coleoptera comprise the largest order of animals with more than 350,000 named species. They are part of the food web in nearly every nonmarine habitat, and play numerous and important ecological roles, for example as consumers, pollinators, and decomposers. Here I review our current understanding of higher-level (particularly subordinal) relationships in beetles based on DNA sequence data. I also elaborate upon a previous review of beetle molecular chronograms (timetrees) (McKenna and Farrell, 2009) by adding newly published data and discussing how phylogenetic methods have been used to reconstruct timing and patterns of ecological and taxonomic diversification in beetles.
The 16 superfamilies and 168 families of extant beetles are separated into four suborders: Adephaga (~35,000 species), Archostemata (~35 species), Myxophaga (~65 species), and Polyphaga (~315,000 species) (Lawrence and Newton, 1995; Arnett and Thomas, 2000; Arnett et al., 2002; Beutel and Leschen, 2005). Adephaga are largely predators and their earliest known fossils are from the early Triassic (Grimaldi and Engel, 2005). The abdominal sternite II of Adephaga is divided by the metacoxae. Archostemata feed on decaying wood as larvae and pollen as adults. The earliest known fossil Archostemata are from the late Permian (H6rnschemeyer, 2005). Archostemata have the labrum fused to the head capsule and roll the tips of their wings under the elytra. Myxophaga are aquatic or semiaquatic and feed on algae (Beutel and Leschen, 2005). They are not definitively known from the fossil record, but possible relatives are known from the Permian (e.g., Ponomarenko, 1969; Lawrence and Newton, 1982). Myxophaga are highly specialized for aquatic life. Adults respire via a plastron, and larvae have tracheal gills. Polyphaga exhibit diverse feeding habits, but most consume living plants or dead and decaying plant parts (Arnett and Thomas, 2000; Arnett et al., 2002). The earliest known polyphagan fossils are from the early Triassic (Grimaldi and Engel, 2005). Polyphaga have presternal cervical sclerites, and the propleura is internalized.
Relationships among the four suborders of beetles remain unsettled (e.g., see Crowson, 1960; Lawrence and Newton, 1982; Lawrence et al., 1995). However, an arrangement with Archostemata as sister to all other beetles, Myxophaga and Polyphaga as sister groups, and Adephaga as sister to Myxophaga and Polyphaga, is favored by recent morphological studies (Beutel and Haas, 2000; Beutel, 2005). Autapomorphies of Coleoptera include elytra with meso- and metathoracic locking devices, close connection of exposed sclerites, reduced abdominal sternite I, and invagination of terminal abdominal segments (Beutel and Haas, 2000). Characters supporting the monophyly of Coleoptera excluding Archostemata include the absence of the mesothoracic discriminal line and katepisternal joint (and other transformations of the thoracic sclerites), internalized or absent metathoracic trochantin, and the presence of a bending zone in the hindwing. Adult Myxophaga and Polyphaga are characterized by the rigid connection of the meso- and metathoracic ventrites, and the fusion of protrochantin and propleura (Beutel and Haas, 2000).
SUBORDINAL RELATIONSHIPS AND TIMETREES
The first study focused on reconstructing relationships in beetles using molecular data (Howland and Hewitt, 1995) sampled a 400 bp piece of cytochrome oxidase I from 37 beetle species in 15 families, with representatives from two of the four beetle suborders (Table l). The resulting neighbor joining tree was not well resolved and provided relatively little new insight into beetle phylogeny. Shull, et al. (2001) published the first comprehensive molecular phylogeny for beetles, with taxa sampled from ali four suborders. They analyzed nearly complete small subunit ribosomal RNA (18S) sequences from 48 beetle species in 19 families (taxon sampling was focused on Adephaga), and 4 neuropterid outgroups. Most of their extensive analyses recovered Adephaga and Polyphaga as sister groups, and the suborders Myxophaga and Archostemata grouped together as sister to a clade comprised of Adephaga and Polyphaga. However, in some analyses Polyphaga arose from within Hydradephaga.
Caterino, et al. (2002) reconstructed beetle phylogeny using 18S sequences from 25 beetle species in 24 families representing all four suborders. They also sampled a diverse array of 46 other insects. When Coleoptera and each of its constituent suborders were constrained to be monophyletic in a parsimony (MP) analysis, Archostemata was recovered in a position sister to the remaining beetles, Adephaga and Polyphaga were sister groups, and Myxophaga was recovered in a position sister to Adephaga and Polyphaga. Without constraints, Coleoptera were rendered paraphyletic by the inclusion of Diptera and Strepsiptera.
Caterino, et al. (2005), although focused on reconstructing relationships within Staphylinifor mia and between Staphyliniformia and Scarabaeiformia, sampled 18S sequences from 25 beetles, including representatives of ali four suborders. Analyses under Bayesian (BI) and maximum likelihood (ML) inference recovered Adephaga and Polyphaga as sister groups, and Archostemata alone or in combination with Myxophaga as the closest relative of ali other beetles. Parsimony analysis recovered Myxophaga in a position sister to Polyphaga. Vogler (2005) analyzed 18S sequences from 795 beetle taxa in 123 families. In the resulting most parsimonious tree, the single exemplar of Archostemata (Distocupes) was recovered in a position sister to the remaining beetles, Adephaga and Polyphaga were sister groups, and Myxophaga was sister to the clade comprised of Adephaga and Polyphaga.
Hughes, et al. (2006) used expressed sequence tags to generate phylogenetic markers (66 genes; matrix 28.6% complete) for 14 beetle species, which included representatives of ali four suborders. Analysis under MP, ML, and BI recovered similar phylogenetic trees. When trees were rooted with Archostemata, Myxophaga and Polyphaga were sister groups, with Adephaga as their sister group. Supertree analyses yielded generally less resolution, and favored the placement of Myxophaga within Polyphaga.
Hunt, et al. (2007) published the most extensively taxon-sampled phylogeny of beetles to date. Their study included nearly complete 18S sequences for 1,880 taxa, and sequences from mitochondrial 16S rRNA and cytochrome oxidase subunit I (COI) for nearly half of these. In the trees resulting from analysis under MP and BI, Adephaga and Polyphaga were sister groups, together sister to Myxophaga plus Archostemata. Hunt, et al. (2007) also published the first molecular timetree for beetles, based on BI analysis of a 340-taxon subset of their 1,880-taxon tree. The age of the ingroup was fixed at 285 Ma in an "all compatible" version of the resulting consensus tree, and seven fossil age constraints were used to calibrate the tree and date internal nodes under penalized likelihood (PL) in the program r8s (Sanderson, 2002, 2003). Average ages and 95% confidence intervals were reported for 13 selected clades in the resulting chronogram (1). Most families (> 100) were found to originate in the Jurassic. To investigate the role of herbivory in beetle diversification, Hunt, et al. (2007) compared species richness between clades feeding on living plants and their sister clades with hosts other than living plants. They also performed a more restrictive analysis limiting the comparison to clades feeding on angiosperms versus those with other hosts (including gymnospermous plants). Based on these analyses, they concluded that the success of beetles is "explained neither by exceptional net diversification rates nor by a predominant role of herbivory and the Cretaceous rise of angiosperms." As an alternative, they proposed that the species richness of beetles is due to "high survival of lineages and sustained diversification in a variety of niches" (Hunt et al., 2007).
Wild and Maddison (2008) evaluated the phylogenetic performance of 9 nuclear genes (alpha-spectrin (AS), RNA polymerase H (RNA Pol II), topoisomerase I (TOP1), arginine kinase (AK), carbamoylphosphate synthase domain (CAD), enolase, phosphoenolpyruvate carboxykinase (PEPCK), wingless (WG), and 28S) sequenced from 31 beetles in 18 genera representing all suborders except Myxophaga (but focused on Adephaga), and 2 outgroups. Performance was evaluated by comparing the topologies obtained by analysis under BI to a "presumably known" test phylogeny in which uncertain subordinal relationships were intentionally left unresolved. Bayesian and MP analyses of the concatenated nine-gene data set recovered Archostemata and Adephaga as sister groups, together sister to Polyphaga. This was the first higher-level molecular phylogenetic study of beetles to include extensive data from nuclear protein-coding genes. Maddison, et al. (2009) used 18S and 28S DNA sequences and data from the nuclear protein-coding gene WG to reconstruct the phylogeny of Adephagan beetles. They sampled exemplars from all four suborders of beetles, including 60 species of Adephaga, 4 genera representing 2 families of Archostemata (more Archostemata than in any other study to date; Cupes capitatus, Priacma serrata, Tenomerga cinetea, and Micromalthus debilis), 3 Myxophaga, and 10 Polyphaga. Bayesian analysis of the combined data recovered Archostemata and Myxophaga as sister groups, and Adephaga and Polyphaga as sister groups.
McKenna and Farrell (2009) analyzed nearly complete 18S sequences from 955 beetle genera, including representatives of 134 beetle families. Analysis under ML inference recovered Adephaga and Polyphaga as sister groups, together sister to Myxophaga plus Archostemata. Nonparametric rate smoothing (NPRS) (Sanderson, 1997) implemented in r8s was used to generate an ultrametric tree from the ML topology. Six fossils and each of two alternative maximum constraints on the age of Holometabola (separately applied) were used to calibrate the tree and date internal nodes, resulting in the most extensively taxon-sampled molecular timetree for beetles to date. The split between Myxophaga + Archostemata and the clade comprised of the suborders Adephaga and Polyphaga was estimated to have occurred - 269 265 Ma (mean 266.8 Ma). By comparison, Hunt, et al. (2007) fixed the age of this split at 285 Ma. McKenna and Farrell (2009) estimated that the Adephaga-Polyphaga split occurred ~ 269265 Ma (mean 266.4 Ma), just slightly later than Hunt, et al. (2007), who estimated this split to have occurred ~ 277 Ma. Divergence times were not evaluated below the subordinal level in McKenna and Farrell (2009) due to the lack of well-supported resolution at lower taxonomic levels in Polyphaga and beyond. The results of Hunt, et al. (2007) and McKenna and Farrell (2009) indicate that the four living suborders of beetles diverged in the Permian, when early amniotes, conifers, and others groups of terrestrial organisms, including other insects (Labandeira and Sepkoski, 1993), were also diversifying. In a recent series of related studies, mitochondrial (mt) genomes were used to estimate the higher-level phylogeny of beetles (Sheffield et al., 2008, 2009; Song et al., 2010; Pons et al., 2010). Mitochondrial genomes have also been used to estimate the higher-level phylogeny of Neuropterida and their relationship to other holometabolous insects, including nine beetles in the suborders Adephaga and Polyphaga (Cameron et al., 2009). In addition to addressing relationships, these studies report on the effects of systematic bias contributed by base compositional heterogeneity and among-site rate variation on phylogeny reconstruction. Base compositional heterogeneity and among-site rate variation are problematic for phylogeny reconstruction because they can produce misleading estimates of topology and branch lengths and misleading bipartition posterior probabilities, and may cluster unrelated taxa based on convergent base content rather than evolutionary history (e.g., Lemmon et al., 2009; Song et al., 2010). Song, et al. (2010), with data from 24 beetles representing all 4 suborders (and 7 outgroups), included more beetles than any other of the aforementioned studies employing mt genomes. They presented numerous phylogenetic trees, e.g., some resulting from analyses designed specifically to overcome systematic bias, and others resulting from more traditional analytical methods. Their MP "reference phylogeny" based on amino acid sequences, and determined to be the least likely to violate phylogenetic assumptions, recovered Myxophaga (1 exemplar; Sphaerius) and Adephaga (3 exemplars) as sister groups, these together sister to Polyphaga (19 exemplars). Archostemata (1 exemplar; Tetraphalerus) was the sister group to all other Coleoptera. Although the monophyly of Adephaga and Polyphaga were well supported, none of the aforementioned subordinal relationships were supported by bootstrap values [greater than or equal to] 80%. Pons, et al. (2010) used mt genomes (2 newly sequenced) to reconstruct the phylogeny of beetles and to estimate nucleotide substitution rates for mitochondrial protein-coding (MPC) genes. Analysis of the nucleotide sequences of the 13 MPC's under BI recovered Polyphaga (10 exemplars) + Archostemata (1 exemplar; Tetraphalerus) as a clade, and also Adephaga (2 exemplars) + Myxophaga (2 exemplars).
Studies focused primarily on reconstructing relationships in Holometabola (e.g., Whiting et al., 1997, 2002a,b; Wheeler et al., 2001) using 18S and/or 28S DNA sequence data give some insight into subordinal relationships in beetles. For example, Whiting (2002a), used 18S sequences from 47 beetles, including representatives of all 4 suborders, and a diverse array of 100 other holometabolous insects. Parsimony analysis recovered a paraphyletic Coleoptera and Adephaga, with the single archostematan sampled (Distocupes) recovered within Myxophaga (represented by Hydroscapha and Torridincola).
Several recent molecular phylogenetic studies that focused on Holometabola or other higherlevel groups of insects indicate that Strepsiptera are closely related to beetles, if not beetles themselves (2) (Misof et al., 2007; Wiegmann et al., 2009a,b; Ishiwata et al., 2010; Longhorn et al., 2010; McKenna and Farrell, 2010). Longhorn, et al. (2010) evaluated relationships in Holometabola with a focus on the phylogenetic placement of Strepsiptera using data from 27 nuclear ribosomal proteins for 22 holometabolous insects, including 7 beetles and 2 Strepsiptera. Their taxon sample lacked representatives from the beetle suborder Archostemata and the supra-ordinal group Neuropterida (Orders Megaloptera, Neuroptera and Raphidioptera); thought to be the sister group to Coleoptera alone or to Coleoptera + Strepsiptera (Weigmann et al., 2009a,b). Analysis under BI recovered the single adephagan (Cicindela) in a position sister to Polyphaga. The single myxophagan (Sphaerius) and the two Strepsiptera were recovered in various positions in the tree depending on the nucleotide-coding scheme and method of phylogenetic inference employed (whether ML, BI, or MP). However, the most convincing placement was as sister group to Coleoptera. Regardless, Strepsiptera were recovered as close relatives of beetles under most coding schemes and analytical methods.
McKenna and Farrell (2010) used DNA sequences from 9 nuclear genes (elongation factor-1[alpha] (EF-1[alpha]), alanyl-tRNA synthetase (AATS), CAD, 6-phosphogluconate dehydrogenase (PGD), sans fille (SNF), triosephosphate isomerase (TPI), RNA Pol II, 28S, and 18S) to reconstruct the phylogeny of Holometabola with a focus on determining the phylogenetic placement of Strepsiptera. Their taxon sample was comprised of 32 exemplars representing all orders of Holometabola, including 8 beetles in as many families (and representing all 4 suborders), 2 Strepsiptera, and 2 hemimetabolous insect outgroups. Analysis under ML and BI recovered the single exemplars of Archostemata and Myxophaga together in a position sister to Adephaga, and these three suborders sister to Polyphaga + Strepsiptera. The same relationships were recovered when rDNA were excluded from ML and BI analyses, except for Strepsiptera, which were recovered in a position sister to Neuropterida and therefore outside of Coleoptera. Strepsiptera have not been included in any published analyses along with both a comprehensive sample of beetle suborders and an extensive sample of beetle families.
OTHER HIGHER-LEVEL TIMETREES
Molecular chronograms for beetles that focused on relationships at the level of series or superfamily and were calibrated with information from the geological record to produce a timetree are so far available only for the cucujiform superfamilies Chrysomeloidea (Farrell, 1998; Gomez-Zurita et al., 2007) and Curculionoidea (Farrell, 1998; McKenna et al., 2009). Farrell (1998) used MP to reconstruct the phylogeny of Phytophaga (Chrysomeloidea and Curculionoidea) from 18S sequences for 115 species, and 212 morphological characters. The resulting most parsimonious trees showed basal conifer- and cycad-feeding lineages in both superfamilies, consistent with their proposed status as ancient host associations. The age of each major constituent group, estimated from the fossil record, was used to prepare an estimated timetree for Phytophaga from the strict consensus tree. Five independent contrasts were made of sister groups where one of the groups was associated with gymnospermous plants and the other with angiosperms, all yielding a positive difference in favor of the hypothesis that feeding on angiosperms is associated with enhanced taxonomic diversity (Farrell, 1998).
Gomez-Zurita, et al. (2007) contended that available phylogenetic information and molecular clock calibrations were insufficient to conclude that the taxonomic diversity of Phytophaga can be attributed to "co-radiation with angiosperms." To revisit this subject, they obtained estimated phylogenetic trees for 167 taxa in the superfamily Chrysomeloidea (but focused on Chrysomelidae) from ML and MP analyses of data from 16S, 18S, and 28S. Penalized likelihood implemented in the program rSs was used to generate an ultrametric tree from the ML topology. Minimum age constraints were imposed based on two fossils, and the vicariant split between Palearctic and Nearctic Timarchini (Chrysomelidae). The resulting timetree was consistent with a Late Cretaceous origin for Chrysomelidae, and subsequent taxonomic diversification over the course of the Cenozoic--considerably later than proposed by Farrell (1998). Gomez-Zurita, et al. (2007) therefore argued that chrysomelid beetles radiated long after the origin of angiosperms and that their diversification was driven by repeated radiation on a pre-existing diverse resource, rather than ancient host associations.
McKenna, et al. (2009) reconstructed the evolutionary history of diversification in weevils using DNA sequence data from 6 genes (EF-1[alpha], CAD, AK, 28S, 18S, and COI) for 135 genera representing all families and subfamilies of Curculionoidea. Divergence times derived from the combined molecular and fossil data were coestimated with phylogeny using the Bayesian relaxed molecular clock method (Drummond et al., 2006) in the program BEAST (3). These analyses indicated diversification into most modern families occurred on gymnosperms in the Jurassic. Colonization of angiosperms appears to have occurred during the Early Cretaceous, with massive taxonomic diversification not beginning until the midCretaceous, when angiosperms first rose to widespread floristic dominance. The authors proposed that these and other evidence are consistent with a deep and complex history of coevolution between weevils and angiosperms (McKenna et al., 2009).
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DISCUSSION AND CONCLUSIONS
Most studies of beetle molecular phylogeny with extensive taxon sampling have relied largely or solely on DNA sequences from the small subunit ribosomal RNA (18S) (e.g., Shull et al., 2001; Vogler, 2005; Hunt et al., 2007; McKenna and Farrell, 2009), and most of these studies have recovered, at least under some analytical conditions, Adephaga and Polyphaga as sister groups, and Archostemata alone or in combination with Myxophaga as the closest relative of all other beetles (Fig. 1). Despite the relative consistency of subordinal relationships recovered in these studies, they fail to recover extensive compatible and well-supported resolution below the subordinal level, particularly in the suborder Polyphaga, the most species rich of the four beetle suborders. The extensively taxon-sampled study of Hunt, et al. (2007), with data from 3 genes, is a step in the right direction. However, it too was based largely on data from 18S. Data from 16S and COI were available for slightly less than half of the taxa.
Due to limited gene and nucleotide sampling, missing data, and a corresponding lack of well-supported resolution in studies to date, existing molecular phylogenies for the order Coleoptera based largely or solely on 18S should be viewed as tentative. The recent 9-gene phylogeny of Wild and Maddison (2008) and the 6-gene phylogeny of McKenna, et al. (2009) (limited to Curculionoidea) recovered considerable well-supported resolution, and suggest that similarly large gene samples may contribute to resolving relationships in other major beetle radiations, and with sufficient taxon sampling, perhaps even across the entirety of the order Coleoptera.
On account of the current lack of strong nodal support for the interrelationships and internal relationships of most groups of beetles, it is difficult to justify detailed evaluation of the timing and causes of ecological and taxonomic diversification in most beetle groups, let alone across the entire order Coleoptera. However, methods for estimating relationships and/or node ages from molecular (and other) data have matured considerably over the approximately 15 years that have elapsed since publication of the first significant attempt at reconstructing beetle phylogeny using DNA sequence data (Howland and Hewitt, 1995). Further, recent studies of the beetle fossil record, e.g., by Oberprieler, et al. (2007) and Krell (2006), are contributing to a better understanding of timing and patterns in the appearance of major groups of beetles. Consequently, future studies that (1) obtain data from multiple molecular markers for a broad cross-section of beetles, neuropterids and Strepsiptera, (2) incorporate the latest information from the beetle fossil record, and (3) use appropriate and statistically rigorous methods for estimating beetle phylogeny and divergence times will undoubtedly contribute further and more robust insights into beetle phylogeny and macroevolution, including factors contributing to the apparent success of the order.
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Wild, A. L. and D. R. Maddison. 2008. Evaluating nuclear protein-coding genes for phylogenetic utility in the Coleoptera. Molecular Phylogenetics and Evolution 48: 877-891.
Received 24 September 2010; accepted 17 March 2011.
(1) McKenna and Farrell (2009) reported estimated divergence times for ali family-level groups in the timetree published by Hunt, et al. (2007).
(2) See McKenna and Farrell (2010) for a discussion of possible placements within or near beetles.
(3) Despite the popularity of BI in studies of beetle molecular phylogeny, Bayesian methods for co-estimating node ages and phylogeny have not yet been applied to the study of major beetle radiations outside of Curculionoidea.
DUANE D. MCKENNA (1)
Department of Biological Sciences, University of Memphis, Memphis, TN 38152, USA
(1) Email address for correspondence: dmckenna@ memphis.edu
Table 1. Studies reviewed. DNA sequence Authors Taxonomic focus data used Howland & Hewitt Coleoptera COI 1995 Whiting et al. 1997 Holometabola 18S, 28S Farrell1998 Chrysomeloidea, 18S Curculionoidea Shull et al. 2001 Adephaga 18S Wheeler et al. 2001 Hexapoda 18S, 28S Whiting 2002a Holometabola 18S Whiting 2002b Holometabola 18S Caterino et al. 2002 Coleoptera 18S Caterino et al. 2005 Staphyliniformia 18S Scarabaeiformia Vogler 2005 Coleoptera 18S Hughes et al. 2006 Coleoptera 66 RP genes Gomez-Zurita et al. Chrysomeloidea 18S, 28S, 16S 2007 Hunt et al. 2007 Coleoptera 18S, 16S, COI Misof et al. 2007 Hexapoda 18S Wild & Maddison Coleoptera 28S, AK, AS, CAD, 2008 Enolase, PEPCK, RNA Pol II, TOPI, WG Sheffield et al. 2008 Coleoptera mt genomes Cameron et al. 2009 Neuropterida mt genomes Maddison et al. 2009 Adephaga 18S, 28S, WG McKenna & Farrell Coleoptera 18S 2009 McKenna et al. 2009 Curculionoidea EF-1[alpha], AK, 28S, 18S, 16S, COI Sheffield et al. 2009 Coleoptera mt genomes Wiegmann et al. Holometabola AATS, CAD, PGD, 2009a SNF, TPI, RNA Pol II Wiegmann et al. Holometabola AATS, CAD, PGD, 2009b SNF, TPI, RNA Pol II Ishiwata et al. 2010 Insecta DNA polymerase delta, RNA Pol II Longhorn et al. 2010 Holometabola 27 RP genes McKenna & Farrell Holometabola EF-l[alpha], AATS, 2010 CAD, PGD, SNF, TPI, RNA Pol II, 28S, 18S Pons et al. 2010 Coleoptera mt genomes Song et al. 2010 Coleoptera mt genomes Beetle suborders Strepsiptera Authors sampled * included? Timetree? Howland & Hewitt Ad, Po No No 1995 Whiting et al. 1997 Ad, Ar, Po Yes No Farrell1998 N/A N/A Yes Shull et al. 2001 All 4 No No Wheeler et al. 2001 Ad, Po Yes No Whiting 2002a All 4 Yes No Whiting 2002b Ad, My, Po Yes No Caterino et al. 2002 All 4 Yes No Caterino et al. 2005 All 4 No No Vogler 2005 All 4 No No Hughes et al. 2006 All 4 No No Gomez-Zurita et al. N/A N/A Yes 2007 Hunt et al. 2007 All 4 No Yes Misof et al. 2007 Ad, Po Yes No Wild & Maddison Ad, Ar, Po No No 2008 Sheffield et al. 2008 All 4 No No Cameron et al. 2009 Ad, Po No No Maddison et al. 2009 All 4 No No McKenna & Farrell All 4 No Yes 2009 McKenna et al. 2009 N/A N/A Yes Sheffield et al. 2009 Ar, Po No No Wiegmann et al. Po Yes Yes 2009a Wiegmann et al. Po Yes Yes 2009b Ishiwata et al. 2010 Ad, Po Yes No Longhorn et al. 2010 Ad, My, Po Yes No McKenna & Farrell All 4 Yes No 2010 Pons et al. 2010 All 4 No No Song et al. 2010 All 4 No No * Ad = Adephaga, Ar = Archostemata, My = Myxophaga, Po = Polyphaga
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|Author:||McKenna, Duane D.|
|Date:||Jan 1, 2011|
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