The Beetle Tree of Life Reveals that Coleoptera Survived End-Permium Mass Extinction to Diversify During the Cretaceous Terrestrial Revolution

Here we present a phylogeny of beetles (Insecta: Coleoptera) based on DNA sequence data from eight nuclear genes, including six single-copy nuclear protein-coding genes, for 367 species representing 172 of 183 extant families. Our results refine existing knowledge of relationships among major groups of beetles. Strepsiptera was confirmed as sister to Coleoptera and each of the suborders of Coleoptera was recovered as monophyletic. Interrelationships among the suborders, namely Polyphaga (Adephaga (Archostemata, Myxophaga)), in our study differ from previous studies. Adephaga comprised two clades corresponding to Hydradephaga and Geadephaga. The series and superfamilies of Polyphaga were mostly monophyletic. The traditional Cucujoidea were recovered in three distantly related clades. Lymexyloidea was recovered within Tenebrionoidea. Several of the series and superfamilies of Polyphaga received moderate to maximal clade support in most analyses, for example Buprestoidea, Chrysomeloidea, Coccinelloidea, Cucujiformia, Curculionoidea, Dascilloidea, Elateroidea, Histeroidea and Hydrophiloidea. However, many of the relationships within Polyphaga lacked compatible resolution under maximum-likelihood and Bayesian inference, and/or lacked consistently strong nodal support. Overall, we recovered slightly younger estimated divergence times than previous studies for most groups of beetles. The ordinal split between Coleoptera and Strepsiptera was estimated to have occurred in the Early Permian. Crown Coleoptera appeared in the Late Permian, and only one or two lineages survived the end-Permian mass extinction, with stem group representatives of all four suborders appearing by the end of the Triassic. The basal split in Polyphaga was estimated to have occurred in the Triassic, with the stem groups of most series and superfamilies originating during the Triassic or Jurassic. Most extant families of beetles were estimated to have Cretaceous origins. Overall, Coleoptera experienced an increase in diversification rate compared to the rest of Neuropteroidea. Furthermore, 10 family-level clades, all in suborder Polyphaga, were identified as having experienced significant increases in diversification rate. These include most beetle species with phytophagous habits, but also several groups not typically or primarily associated with plants. Most of these groups originated in the Cretaceous, which is also when a majority of the most species-rich beetle families first appeared. An additional 12 clades showed evidence for significant decreases in diversification rate. These clades are species-poor in the Modern fauna, but collectively exhibit diverse trophic habits. The apparent success of beetles, as measured by species numbers, may result from their associations with widespread and diverse substrates – especially plants, but also including fungi, wood and leaf litter – but what facilitated these associations in the first place or has allowed these associations to flourish likely varies within and between lineages. Our results provide a uniquely well-resolved temporal and phylogenetic framework for studying patterns of innovation and diversification in Coleoptera, and a foundation for further sampling and resolution of the beetle tree of life

ZooKeys, unlocking Earth’s incredible biodiversity and building a sustainable bridge into the public domain: From “print-based” to “web-based” taxonomy, systematics, and natural history. ZooKeys Editorial Opening Paper

This is the publisher's version, also available electronically from http://zookeys.pensoft.net/articles.php?id=1929.Publishing taxonomic and systematics studies in the digital era faces major challenges and requires new approaches, many of which are currently stimulating spirited discussions amongst taxonomists and systematists. New amendments to the International Code of Zoological Nomenclature are expected to regulate electronic publishing of new taxa and create a standard form for their registration (ZooBank). Responding to a perceived need, this editorial announces establishment of ZooKeys – a new online and print journal in zoological taxonomy and systematics, which aims to quickly respond and adapt to the newest developments in taxonomic publishing. Open Access is accepted as mandatory for ZooKeys. Th e rationale for and concept of ZooKeys is discussed in detail

The Beetle Tree of Life Reveals the Order Coleoptera Survived End Permain Mass Extinction to Diversify During the Cretaceous Terrestrial Revolution

Here we present a phylogeny of beetles (Insecta: Coleoptera) based on DNA sequence data from eight nuclear genes, including six single-copy nuclear protein-coding genes, for 367 species representing 172 of 183 extant families. Our results refine existing knowledge of relationships among major groups of beetles. Strepsiptera was confirmed as sister to Coleoptera and each of the suborders of Coleoptera was recovered as monophyletic. Interrelationships among the suborders, namely Polyphaga (Adephaga (Archostemata, Myxophaga)), in our study differ from previous studies. Adephaga comprised two clades corresponding to Hydradephaga and Geadephaga. The series and superfamilies of Polyphaga were mostly monophyletic. The traditional Cucujoidea were recovered in three distantly related clades. Lymexyloidea was recovered within Tenebrionoidea. Several of the series and superfamilies of Polyphaga received moderate to maximal clade support in most analyses, for example Buprestoidea, Chrysomeloidea, Coccinelloidea, Cucujiformia, Curculionoidea, Dascilloidea, Elateroidea, Histeroidea and Hydrophiloidea. However, many of the relationships within Polyphaga lacked compatible resolution under maximum-likelihood and Bayesian inference, and/or lacked consistently strong nodal support. Overall, we recovered slightly younger estimated divergence times than previous studies for most groups of beetles. The ordinal split between Coleoptera and Strepsiptera was estimated to have occurred in the Early Permian. Crown Coleoptera appeared in the Late Permian, and only one or two lineages survived the end-Permian mass extinction, with stem group representatives of all four suborders appearing by the end of the Triassic. The basal split in Polyphaga was estimated to have occurred in the Triassic, with the stem groups of most series and superfamilies originating during the Triassic or Jurassic. Most extant families of beetles were estimated to have Cretaceous origins. Overall, Coleoptera experienced an increase in diversification rate compared to the rest of Neuropteroidea. Furthermore, 10 family-level clades, all in suborder Polyphaga, were identified as having experienced significant increases in diversification rate. These include most beetle species with phytophagous habits, but also several groups not typically or primarily associated with plants. Most of these groups originated in the Cretaceous, which is also when a majority of the most species-rich beetle families first appeared. An additional 12 clades showed evidence for significant decreases in diversification rate. These clades are species-poor in the Modern fauna, but collectively exhibit diverse trophic habits. The apparent success of beetles, as measured by species numbers, may result from their associations with widespread and diverse substrates - especially plants, but also including fungi, wood and leaf litter - but what facilitated these associations in the first place or has allowed these associations to flourish likely varies within and between lineages. Our results provide a uniquely well-resolved temporal and phylogenetic framework for studying patterns of innovation and diversification in Coleoptera, and a foundation for further sampling and resolution of the beetle tree of life.Facultad de Ciencias Naturales y Muse

The beetle tree of life reveals that Coleoptera survived end-Permian mass extinction to diversify during the Cretaceous terrestrial revolution

Here we present a phylogeny of beetles (Insecta: Coleoptera) based on DNA sequence data from eight nuclear genes, including six single-copy nuclear protein-coding genes, for 367 species representing 172 of 183 extant families. Our results refine existing knowledge of relationships among major groups of beetles. Strepsiptera was confirmed as sister to Coleoptera and each of the suborders of Coleoptera was recovered as monophyletic. Interrelationships among the suborders, namely Polyphaga (Adephaga (Archostemata, Myxophaga)), in our study differ from previous studies. Adephaga comprised two clades corresponding to Hydradephaga and Geadephaga. The series and superfamilies of Polyphaga were mostly monophyletic. The traditional Cucujoidea were recovered in three distantly related clades. Lymexyloidea was recovered within Tenebrionoidea. Several of the series and superfamilies of Polyphaga received moderate to maximal clade support in most analyses, for example Buprestoidea, Chrysomeloidea, Coccinelloidea, Cucujiformia, Curculionoidea, Dascilloidea, Elateroidea, Histeroidea and Hydrophiloidea. However, many of the relationships within Polyphaga lacked compatible resolution under maximum-likelihood and Bayesian inference, and/or lacked consistently strong nodal support. Overall, we recovered slightly younger estimated divergence times than previous studies for most groups of beetles. The ordinal split between Coleoptera and Strepsiptera was estimated to have occurred in the Early Permian. Crown Coleoptera appeared in the Late Permian, and only one or two lineages survived the end-Permian mass extinction, with stem group representatives of all four suborders appearing by the end of the Triassic. The basal split in Polyphaga was estimated to have occurred in the Triassic, with the stem groups of most series and superfamilies originating during the Triassic or Jurassic. Most extant families of beetles were estimated to have Cretaceous origins. Overall, Coleoptera experienced an increase in diversification rate compared to the rest of Neuropteroidea. Furthermore, 10 family-level clades, all in suborder Polyphaga, were identified as having experienced significant increases in diversification rate. These include most beetle species with phytophagous habits, but also several groups not typically or primarily associated with plants. Most of these groups originated in the Cretaceous, which is also when a majority of the most species-rich beetle families first appeared. An additional 12 clades showed evidence for significant decreases in diversification rate. These clades are species-poor in the Modern fauna, but collectively exhibit diverse trophic habits. The apparent success of beetles, as measured by species numbers, may result from their associations with widespread and diverse substrates – especially plants, but also including fungi, wood and leaf litter – but what facilitated these associations in the first place or has allowed these associations to flourish likely varies within and between lineages. Our results provide a uniquely well-resolved temporal and phylogenetic framework for studying patterns of innovation and diversification in Coleoptera, and a foundation for further sampling and resolution of the beetle tree of life.This is the publisher’s final pdf. The published article is copyrighted by the author(s) and published by John Wiley & Sons, Ltd on behalf of Royal Entomological Society. The published article can be found at: http://onlinelibrary.wiley.com/journal/10.1111/%28ISSN%291365-311

Taxonomy based on science is necessary for global conservation

Peer reviewe

Reducing the environmental impact of surgery on a global scale: systematic review and co-prioritization with healthcare workers in 132 countries

Abstract Background Healthcare cannot achieve net-zero carbon without addressing operating theatres. The aim of this study was to prioritize feasible interventions to reduce the environmental impact of operating theatres. Methods This study adopted a four-phase Delphi consensus co-prioritization methodology. In phase 1, a systematic review of published interventions and global consultation of perioperative healthcare professionals were used to longlist interventions. In phase 2, iterative thematic analysis consolidated comparable interventions into a shortlist. In phase 3, the shortlist was co-prioritized based on patient and clinician views on acceptability, feasibility, and safety. In phase 4, ranked lists of interventions were presented by their relevance to high-income countries and low–middle-income countries. Results In phase 1, 43 interventions were identified, which had low uptake in practice according to 3042 professionals globally. In phase 2, a shortlist of 15 intervention domains was generated. In phase 3, interventions were deemed acceptable for more than 90 per cent of patients except for reducing general anaesthesia (84 per cent) and re-sterilization of ‘single-use’ consumables (86 per cent). In phase 4, the top three shortlisted interventions for high-income countries were: introducing recycling; reducing use of anaesthetic gases; and appropriate clinical waste processing. In phase 4, the top three shortlisted interventions for low–middle-income countries were: introducing reusable surgical devices; reducing use of consumables; and reducing the use of general anaesthesia. Conclusion This is a step toward environmentally sustainable operating environments with actionable interventions applicable to both high– and low–middle–income countries

Elaphidion

Key to species of Elaphidion of Puerto Rico and the Virgin Islands 1. Third and fourth antennomeres spined on both internal and external apical angles..................................... 2 1 ’ Third and fourth antennomeres spined only on internal angle..................................................................... 3 2. Prosternal process parallel-sided, rounded apically, with caudad apex at median; length 30 mm or greater; Puerto Rico ............................................................................................................................ E. michelii Ivie 2 ’ Prosternal process widened and truncate apically, sometimes emarginate medially, median not most caudad point; length 25 mm or less; widespread .................................................. E. conspersum (Newman) 3. Integument dark chocolate brown to black, with patches of white pubescence on head, pronotum and elytra; emargination between apical elytral spines obliquely truncate to weakly bisinuate so that sutural spine is shorter .................................................................................................................... E. irroratum (L.) 3 ’ Integument light brown to dark chocolate brown, with ochre to golden pubescence; emargination between apical elytral spines simply arcuate, both spines long, although outer may be longer................................. 4 4. Metafemoral apex spiniform, spine longer than basal metatibial width; antennomere 4 short relative to 5; apical spine of male antennomere 3 greater than 0.5 × length of antennomere 4; apical spine of female antennomere 3 subequal to or longer than length of antennomere 4; setae of pronotal and elytral disk broader, rounded-blunt at apex ............................................................................................ E. mayesae Ivie 4 ’ Metafemoral apex dentiform, spine subequal to basal metatibial width; antennomere 4 longer relative to 5; apical spine of male antennomere 3 less than 0.5 × length of antennomere 4; apical spine of female antennomere 3 distinctly shorter than antennomere 4; setae of pronotal and elytral disk narrow, acutely attenuate at apex.................................................................................................................................................... 5 5. Lighter in color, setae generally distributed on elytral disc, allowing derm to show through between, always visible just mesad humeral umbone and between striae at level of metacoxa (Figs 15, 17); punctures of basal portion of elytral disc smaller (Fig. 19); Northern Virgin Islands ........ E. pseudonomon Ivie 5 ’ Darker in color, setae clumped in dense patches, completely hiding derm, giving a distinct patterned appearance, density always visible just mesad humeral umbone and between striae at level of metacoxa (Figs 14, 16); punctures of basal portion of elytral disc larger (Fig. 18); St. Croix and Lesser Antilles...... ................................................................................................................................ E. glabratum (Fabricius)Published as part of Ivie, Michael A. & Schwengel-Regala, Michelle L., 2007, The Elaphidion Audinet-Serville of the Puerto Rican Bank: new species, distributions, taxonomic corrections, and a key to species (Coleoptera: Cerambycidae: Elaphidiini), pp. 55-68 in Zootaxa 1503 on pages 62-63, DOI: 10.5281/zenodo.17714

Elaphidion glabratum Fabricius

Elaphidion glabratum (Fabricius) (Figs 16, 18 20) Stenocorus glabratus Fabricius, 1775: 180. Elaphidion glabratum: Ivie, 1985: 310. Monné and Giesbert, 1995: 54. Chalumeau and Touroult, 2005: 101. Monné and Hovore, 2005: 65. [For a complete pre- 1985 synonymy, see Ivie 1985.] A complete discussion of the tortured nomenclatural history of this species was given by Ivie (1985). DIAGNOSIS: The small size (8–19 mm), cinerous to golden pubescence, and relatively long unispinose antennomere 4 will distinguish this species and E. pseudonomon from all other Elaphidion in the area. Although long series of these two closely-related species are obviously distinct to the naked eye when placed next to each other, actual diagnoses of E. glabratum and E. pseudonomon are difficult, and require careful use of details. From E. glabratum, E. pseudonomon, can be distinguished by the male genitalia (see Ivie 1985), somewhat lighter color, details of the antennal configuration and setation. The antennal characters require detailed measurements. In male E. glabratum antennomeres 3 and 4 are subequal (with 4 only rarely more than 1 mm shorter than 3), and 5 is longer (by 1–3 mm) than 3 (i.e. antennomere 4 is greater than 30 % total length of 3 + 4 + 5). In male E. pseudonomon, antennomere 3 and 5 are subequal, with 5 no more than 1 mm longer than 3, while 4 is at least 2 mm shorter than 3 (i.e. antennomere 4 is less than 30 % total length of 3 + 4 + 5). In females of both species, antennomere 4 is consistently shorter than 3, and in E. pseudonomon it is slightly more so, but the difference is difficult to quantify. In female E. pseudonomon, antennomere 4 is usually less that 28 % of the total 3 + 4 + 5, while in E. glabratum it is usually greater than 28 %. For unassociated females, the allopatric distribution is helpful, with E. pseudonomon occurring on the Virgin Islands that lie on the Puerto Rican Bank, while E. glabratum occurs from St. Croix and its satellites into the Lesser Antilles. The elytral setation and punctation are also different, and helpfully not sexually dimorphic. The setae of E. pseudonomon (Figs 17, 19) are more golden and individually larger than in E. glabratum (Figs 16, 18), covering the elytra and pronotum more uniformly, but not being dense enough to completely obscure the underlying cuticle. The specimen therefore looks uniformly lighter colored to the unaided eye because of the uniform mix of the color of setae and cuticle. In fresh specimens, the humerus is covered in setae (Fig. 17), and at mid-elytron (at the level of the metacoxa) the setae uniformly cover the entire disc except for 2 narrow, indistinct strial lines (Fig. 19). Older rubbed specimens of E. pseudonomon may exhibit bare spots, but never to the extent seen in E. glabratum, and the remaining setal patches are always sparse enough to see through to the surface. This characteristic is always distinguishable at a point just mesad the humeral umbone (Fig. 17), and on the mid-point of the elytron at the level of the metacoxa (Fig. 19). Greasy specimens may appear darker, but this is an obvious artifact of preservation. The setae in E. glabratum are cinerous and narrower, and form dense patches that at least in part completely obscure the cuticle (Figs 16, 18), leaving other areas bare except for the single seta associated with each puncture. The difference in setal density of these patches relative to the condition in E. pseudonomon is always distinguishable at a point just mesad the humeral umbo (Fig. 16), and on the mid-point of the elytron at the level of the metacoxa (Fig. 18). The overall effect of this condition is more areas of unobscured cuticle showing to the naked eye, thus making the fresh specimen look darker. Lastly, post-humeral punctures of E. pseudonomon are usually slightly smaller in diameter, making them seem less dense (Fig. 21). In E. glabratum the post-humeral punctures of the elytra are slightly larger (Fig. 20), although there seems to be about the same number of actual elytral punctures in both species. DISTRIBUTION: St. Croix, Buck Is. (near St. Croix, WIBF), Saba (WIBF), St. Martin, St. Barthelémy, St. Eustatius, Nevis, Antigua, Montserrat, and Guadeloupe, and probably St. Christopher, Dominica and St. LuciaPublished as part of Ivie, Michael A. & Schwengel-Regala, Michelle L., 2007, The Elaphidion Audinet-Serville of the Puerto Rican Bank: new species, distributions, taxonomic corrections, and a key to species (Coleoptera: Cerambycidae: Elaphidiini), pp. 55-68 in Zootaxa 1503 on pages 61-62, DOI: 10.5281/zenodo.17714

Elaphidion conspersum Newman

Elaphidion conspersum Newman (Fig. 14) Elaphidion conspersum Newman, 1841: 110. Ivie, 1985: 309. Browne et al., 1993: 44. Monné and Giesbert, 1995: 54. Chalumeau and Touroult, 2005: 96. Monné and Hovore, 2005: 65. Valentine & Ivie 2005: 280. [For a complete pre- 1985 synonymy, see Ivie 1985.] DIAGNOSIS: The bispinose antennae and pronotum with the median callus broadly connected to the base of the pronotum will distinguish this species from all others in the region. Length 18–25 mm. DISTRIBUTION: Bahama Is. (Crooked, Long, Great Exuma, New Providence, Nassau [WIBF] San Salvador [WIBF]); Cuba, Hispaniola, Puerto Rico, St. Thomas, St. John, Guana, St. Croix, Anegada [WIBF], St. Martin, Guadeloupe (Grande-Terre, Basse-Terre, Désirade), Curação, Bonaire. BIOLOGY: Chalumeau and Touroult (2005) list Hippomane manchinella L. as a host of this species.Published as part of Ivie, Michael A. & Schwengel-Regala, Michelle L., 2007, The Elaphidion Audinet-Serville of the Puerto Rican Bank: new species, distributions, taxonomic corrections, and a key to species (Coleoptera: Cerambycidae: Elaphidiini), pp. 55-68 in Zootaxa 1503 on page 56, DOI: 10.5281/zenodo.17714