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The Phylogeny and Classification of Predaceous Diving Beetles (Coleoptera: Dytiscidae)

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Ecology, Systematics, and the Natural History of Predaceous Diving Beetles (Coleoptera: Dytiscidae)

Abstract

The phylogenetics and higher (family-group) classification of extant members of the beetle family Dytiscidae (Coleoptera), or predaceous diving beetles, is reviewed and reassessed. A phylogenetic analysis of the family is presented based on 168 species of diving beetles and 9 outgroup taxa from Gyrinidae, Noteridae, Amphizoidae, and Paelobiidae. All currently recognized dytiscid subfamilies and tribes are represented, most by multiple genera and species. Data include 104 morphological characters and approximately 6700 aligned bases from 9 DNA sequence fragments from cytochrome c oxidase I (COI) and II (COII), histone III (H3), 16S rRNA (16S), 12S rRNA (12S), arginine kinase (argkin), RNA polymerase II (RNA pol II), elongation factor 1 alpha (Ef1α), and wingless (wnt). Parsimony and Bayesian analyses were conducted. The topology of the parsimony tree (consensus of 13 equally-parsimonious solutions) exhibits numerous anomalies inconsistent with convincing morphological features and the Bayesian results and has, generally, relatively poor bootstrap support for major clades. The Bayesian topology is more consistent with major morphological features and has strong support for most clades, and conclusions are based primarily on this estimate. Major higher-level phylogenetic relationships with strong support include: (1) monophyly of Dytiscidae Leach, (2) Matinae Branden sister to the rest of Dytiscidae, (3) Agabinae Thomson + Colymbetinae Erichson, (4) Hydrodytinae Miller + Hydroporinae Aubé, (5) Dytiscinae Leach + Laccophilinae Gistel + Cybistrini Sharp + Copelatinae Branden, (6) monophyly of the subfamilies Matinae, Colymbetinae, Copelatinae, Coptotominae Branden, Lancetinae Branden, Laccophilinae (including Agabetes Crotch), Agabinae (support weaker than in other subfamilies) and Hydroporinae (monophyly of Hydrodytinae not tested), (7) paraphyly of Dytiscinae with Cybistrini sister to Laccophilinae (with strong support) and this clade sister to other Dytiscinae, and (8) monophyly of both Agabini (Agabus-group of genera) and Hydrotrupini Roughley (Hydrotrupes Sharp and the Platynectes-group of genera). Major conclusions regarding tribes within Hydroporinae include: (1) monophyly of the tribes Vatellini Sharp, Methlini Branden, Hydrovatini Sharp, Hygrotini Portevin, Hyphydrini Gistel (without Pachydrus Sharp) and Bidessini Sharp (including Peschetius Guignot, Hydrodessus J. Balfour-Browne and Amarodytes Régimbart) (monophyly of Laccornini Wolfe and Roughley and Pachydrini Biström, Nilsson and Wewalka not tested), (2) Pachydrini is a problematic, long-branched taxa resolved here as sister to Hydrovatini but with weak support, (3) Hydroporini monophyletic except for Laccornellus Roughley and Wolfe and Canthyporus Zimmermann, (4) Laccornellus and Canthyporus together monophyletic and sister to Hydroporinae except Laccornini. Four groups are resolved within Hydroporini exclusive of Laccornellus + Canthyporus corresponding to the Deronectes-, the Graptodytes-, the Necterosoma- and the Hydroporus-groups of genera. The classification of Dytiscidae is revised with the following taxonomic changes [2014]: (1) Hydrotrupini is recognized as a tribe of Agabinae including the genus Hydrotrupes and the Platynectes-group of genera, (2) the genus Rugosus García is moved from Colymbetinae to Copelatinae, (3) Cybistrini is elevated from tribe rank within Dytiscinae to subfamily of Dytiscidae, (4) Hyderodini Miller is placed as a junior synonym of Dytiscini, (5) Laccornellus and Canthyporus are removed from Hydroporini and placed in their own tribe, Laccornellini, (6) the following family-group names are resurrected from synonymy with Hydroporini and placed as subtribes within Hydroporini, Deronectina Galewski (for the Deronectes-group of genera), Siettitiina Smrž (for the Graptodytes-group of genera), Sternopriscina Branden (for the Necterosoma-group of genera), and Hydroporina (for the Hydroporus-group of genera), (7) Carabhydrini Watts is placed as a junior synonym of Sternopriscina, and (8) Hydrodessus, formerly incerta sedis with respect to tribe, is placed in Bidessini. Each subfamily, tribe and subtribe is diagnosed and its taxonomic history discussed.

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Author information

Authors and Affiliations

  1. Department of Biology and Museum of Southwestern Biology, University of New Mexico, Albuquerque, NM, USA

    Kelly B. Miller

  2. Department of Zoology, Swedish Museum of Natural History, Stockholm, Sweden

    Johannes Bergsten

Authors
  1. Kelly B. Miller
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  2. Johannes Bergsten
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Correspondence to Kelly B. Miller .

Editor information

Editors and Affiliations

  1. School of Biological, Environmental, and Earth Sciences, University of Southern Mississippi, Hattiesburg, MS, USA

    Donald A. Yee

Appendix

Appendix

Morphological and ecological characters and states used in combined phylogenetic analysis of Dytiscidae. Characters used in this analysis are derived from several other recent analyses, which should be consulted for additional description and illustration (Miller 2000, 2001, 2009; Miller and Bergsten 2012; Miller et al. 2006, 2007b, 2009a).

Head

  1. 1.

    Anterolateral margin of eye: (0) Not emarginate, (1) Emarginate.

  2. 2.

    Mandibles: (0) Apically acute, (1) Apically broad.

  3. 3.

    Postocular carina: (0) Absent, (1) Present.

  4. 4.

    Transverse occipital line: (0) Absent, (1) Present.

  5. 5.

    Anterior clypeal groove: (0) Broadly interrupted, (1) Continuous.

  6. 6.

    Anterior clypeal margin: (0) Unmodified, (1) Margin produced or beaded.

  7. 7.

    Fronto clypeal suture: (0) Medially effaced, (1) Entire.

  8. 8.

    Anterior prothoracic glands: (0) Absent, (1) Present.

  9. 9.

    Eyes: (0) Not divided, (1) Divided but contiguous (as in Spanglerogyrus), (2) Completely divided (as in most Gyrinidae).

  10. 10.

    Antennae: (0) Simple, (1) With enlarged scape and pedicel and short, compact flagellum (as in Gyrinidae).

  11. 11.

    Scape: (0) Simple, (1) Medially distinctly constricted (as in Noteridae).

Thorax

  1. 12.

    Scutellum: (0) Not externally visible with elytra closed, (1) Externally visible with elytra closed.

  2. 13.

    Lateral pronotal margin: (0) Unbeaded, (1) Beaded.

  3. 14.

    Longitudinal sublateral basal striae on pronotum: (0) Absent, (1) Present.

  4. 15.

    Prosternum, prosternal process: (0) In same plane as prosternum, (1) Declivous, in different plane from prosternum.

  5. 16.

    Prosternal process: (0) Not reaching metaventrite (1) Reaching metaventrite.

  6. 17.

    Prosternal process: (0) Flat to carinate, (1) With distinct, medial, longitudinal sulcus.

  7. 18.

    Prosternal process apex: (0) Pointed or narrowly rounded, (1) Broad, apically truncate.

  8. 19.

    Metepisternum: (0) Not reaching mesocoxal cavities, separated by mesepimeron (1) Reaching mesocoxal cavities.

  9. 20.

    Transverse metaventral suture: (0) Absent, (1) Present.

  10. 21.

    Small lateral lobe on medial portion of metacoxa: (0) Absent, (1) Present.

  11. 22.

    Metacoxa anterior expansion: (0) Not anteriorly expanded, (1) Anteriorly expanded.

  12. 23.

    Metacoxa and abdominal sternite II: (0) Not fused, (1) Fused.

  13. 24.

    Medial cleft of metendosternite: (0) Narrow, (1) Broad and rounded.

  14. 25.

    Noterid platform: (0) Absent, (1) Present.

  15. 26.

    Retractoris mesothoracis muscle insertion on metendosternite: (0) Deeply concave, distinct from margins, (1) Shallow, flattened.

  16. 27.

    Furcodorsalis metathoracis muscle insertions on anterior rami of metendosternite: (0) Medially, (1) Apically.

  17. 28.

    Basal portion of metendosternite: (0) Broad, lateral margin divergent anteriorly, (1) Narrow, lateral margins parallel.

Abdomen

  1. 29.

    Transverse rugae on dorsum of abdominal segment II: (0) Absent, (1) Present.

  2. 30.

    Speleum: (0) Absent, (1) Present (in Vatellini).

  3. 31.

    Apex of female sternum six: (0) Evenly curved, (1) Medially emarginate.

  4. 32.

    Abdominal tergum VIII: (0) Not modified, (1) Apically acute or acuminate with anteriorly directed processes.

Elytron

  1. 33.

    Anterior oblique epipleural carina: (0) Absent, (1) Present.

  2. 34.

    A field of short impressed striae at base of female elytron: (0) Absent, (1) Present.

  3. 35.

    Elytral apices: (0) Not acuminate, (1) Distinctly acuminate.

  4. 36.

    Elytral apices: (0) Not truncate or sinuate, (1) Truncate and slightly sinuate.

  5. 37.

    Apicoventral elytral setae: (0) Absent, (1) Present.

  6. 38.

    Apicoventral elytral setae: (0) A large region of fine setae, (1) A small region of stiff setae, (2) A linear submarginal series.

Legs

  1. 39.

    Pro- and mesotarsi: (0) Clearly pentamerous, (1) Pseudotetramerous.

  2. 40.

    Apices of male protarsal adhesive setae: (0) Oval to elongate, (1) Round, sucker shaped.

  3. 41.

    Anterior protibal spur in male: (0) Absent, (1) Present.

  4. 42.

    Posterior protibial spur in male: (0) Absent, (1) Present.

  5. 43.

    Protibial hooked spur: (0) Absent, (1) Present (as in Noteridae).

  6. 44.

    Ventral series of setae on mesofemur: (0) Shorter than width of mesofemur, (1) Longer than width of mesofemur.

  7. 45.

    Oblique line of setae on mesotarsomeres: (0) Absent, (1) Present.

  8. 46.

    Posterodorsal series of setae on mesotibia: (0) Simple, (1) Bifid.

  9. 47.

    Posterodorsal series of setae on metatibia: (0) Simple, (1) Bifid.

  10. 48.

    Posterodorsal setae on metatibia: (0) A linear series, slightly curved, (1) A large cluster, (2) A strongly oblique series.

  11. 49.

    Posteroapical setae on meso- and metatibia: (0) Simple, (1) Bifid.

  12. 50.

    Appressed striae on metacoxa: (0) Absent, (1) Present.

  13. 51.

    Metacoxal lines: (0) Not closely approximated medially, (1) Closely approximated.

  14. 52.

    Posteromedial metacoxal lobes: (0) Absent, (1) Present small, (2) Present large rounded.

  15. 53.

    Posteromedial metacoxal rim: (0) Discontinuous medially, (1) Continuous medially.

  16. 54.

    Metacoxal cavities: (0) Separated broadly, (1) Contiguous or closely approximated.

  17. 55.

    Metacoxa: (0) Medial portion not concave, (1) Medial portion concave laterally.

  18. 56.

    Deep, oblique groove on metatrochanter: (0) Absent, (1) Present.

  19. 57.

    Linear series of setae in groove near anteroapical angle of metafemur: (0) Absent, (1) Present.

  20. 58.

    Dorsal series of natatory setae on metafemur: (0) Absent, (1) Present.

  21. 59.

    Apices of metatibial spurs: (0) Simple, (1) Bifid.

  22. 60.

    Anterior metatibial spur: (0) Similar to posterior, (1) Acuminate, broad (Cybistrini).

  23. 61.

    Posteroventral series of setae on metatarsomere I: (0) Absent, (1) Present.

  24. 62.

    Posterodorsal series of setae on metatarsomere I: (0) Absent, (1) Present.

  25. 63.

    Adpressed setae along apicodorsal and apicoventral margins of metatarsomeres I–IV: (0) Absent, (1) Present.

  26. 64.

    Natatory setae along posteroventral margin of metatarsi: (0) Absent on males and females, (1) Present on males, absent on females, (2) Present on males and females.

  27. 65.

    Natatory setae along posteroventral margin of metatibia and tarsomeres: (0) Present or absent on both, (1) Present on metatarsomeres but absent on metatibia in both sexes.

  28. 66.

    Anterodorsal margin of metatarsomeres I–IV: (0) Unlobed, (1) Lobed.

  29. 67.

    Metatarsal claws: 0) Male and female unequal, posterior shorter than anterior, (1) Male and female with claws equal, (2) Male equal, female unequal, anterior shorter than posterior, (3) Male and female unequal, anterior shorter than posterior, (4) Male single, female unequal, anterior shorter than posterior, (5) Male and female each with a single claw.

  30. 68.

    Legs: (0) Not expanded, (1) Meso- and metatibia broad, elongate, apically with elongate extension (as in Spanglerogyrus), (2) Meso- and metatibia short and broad, meso- and metataromeres I–IV very broad, subtriangular (as in most Gyrinidae).

Internal

  1. 69.

    Proventriculus: (0) Without five finger-like lobes on crusher teeth, (1) With five finger-like lobes on crusher teeth.

Male Genitalia

  1. 70.

    Median lobe: (0) Asymmetrical, (1) Bilaterally symmetrical.

  2. 71.

    Ventral sclerite on median lobe: (0) Absent, (1) Present.

  3. 72.

    Lateral lobes: (0) With one segment, (1) With two segments, (2) With three segments.

  4. 73.

    Lateral lobes: (0) Symmetrical, (1) Asymmetrical.

Female Genitalia

  1. 74.

    Genital configuration: (0) carabid-type, (1) noterid-type, (2) amphizoid-type, (3) hydroporine-type, (4) Agaporomorphus-type, (5) dytiscine-type (Miller 2001).

  2. 75.

    Gonocoxal fusion: (0) Not fused, (1) Weakly fused, (2) Completely fused.

  3. 76.

    Anterior apodeme on gonocoxae: (0) absent, (1) present.

  4. 77.

    Gonocoxal shape: (0) Not short and broad, (1) Short and broad.

  5. 78.

    Gonocoxal shape: (0) Not extremely elongate, (1) Extremely elongate.

  6. 79.

    Dorsolateral carina on gonocoxa: (0) Absent, (1) Present.

  7. 80.

    Dorsolateral carina on gonocoxa: (0) Not dentate, (1) Dentate.

  8. 81.

    Gonocoxae: (0) Rounded or not flattened, (1) Strongly laterally flattened.

  9. 82.

    Articulation of laterotergite and gonocoxa: (0) Strongly angled, laterotergite extending posteriorly, (1) Not angled.

  10. 83.

    Laterotergites: (0) Absent, (1) Present.

  11. 84.

    Series of short spinous setae along medial margin gonocoxosternite: (0) Absent, (1) Present.

  12. 85.

    Rami dentation: (0) Not dentate, (1) Weakly dentate, (2) Strongly dentate.

  13. 86.

    Rami configuration: (0) Not modified, (1) Strongly sclerotized, apically fused, anteriorly divergent.

  14. 87.

    Bursa size: (0) Short, (1) Long, slender, slightly twisted.

  15. 88.

    Bursal shape: (0) Various, (1) Flattened, with thick-walled, parallel margins.

  16. 89.

    Bursal gland reservoir: (0) Absent, (1) Present.

  17. 90.

    Bursal sclerotization: (0) Not heavily sclerotized, (1) Heavily sclerotized.

  18. 91.

    Bursal attachment to gonocoxal apparatus: (0) Anteriorly near bases of gonocoxae, (1) Posteriorly near apices of gonocoxae.

  19. 92.

    Receptacle: (0) Absent, (1) Present.

  20. 93.

    Receptacle shape: (0) Not cone shaped, (1) Cone shaped with basal sclerotized ring.

  21. 94.

    Spermathecal and fertilization ducts: (0) Not coiled, (1) Coiled.

  22. 95.

    Spermatheca: (0) Not reduced, (1) Reduced.

  23. 96.

    Spermathecal spine: (0) Absent, (1) Present.

  24. 97.

    Triangular spermathecal process: (0) Absent, (1) Present.

  25. 98.

    Large accessory gland reservoir near base of fertilization duct: (0) Absent, (1) Present.

  26. 99.

    Large accessory gland reservoir on spermatheca: (0) Absent, (1) Present.

  27. 100.

    Spermathecal disc: (0) Absent, (1) Present.

  28. 101.

    Base of fertilization duct: (0) Without modifications, (1) A heavily sclerotized, irregular ring.

  29. 102.

    Gland reservoirs laterally at base of oviduct: (0) Absent, (1) Present.

  30. 103.

    Thick musculature on vagina: (0) Absent, (1) Present.

Habitat

  1. 104.

    Habitat: (0) Not on water surface, (1) found on water surface.

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Miller, K.B., Bergsten, J. (2023). The Phylogeny and Classification of Predaceous Diving Beetles (Coleoptera: Dytiscidae). In: Yee, D.A. (eds) Ecology, Systematics, and the Natural History of Predaceous Diving Beetles (Coleoptera: Dytiscidae). Springer, Cham. https://doi.org/10.1007/978-3-031-01245-7_3

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