Tuesday, July 22, 2025

[Entomology • 2025] A Revised Classification of the Assassin Bugs (Hemiptera: Heteroptera: Reduviidae) based on combined analysis of phylogenomic and morphological data

 

Assassin bug diversity and examples of unique morphological and behavioural adaptations found across the family.

in Masonick, Knyshov, Gordon, Forero, Hwang, ... et Weirauch, 2025. 
  
Abstract
Assassin bugs (Hemiptera: Reduviidae Latreille) comprise not only one of the largest radiations of predatory animals (22 subfamilies; >6,800 spp.) but also include the medically important kissing bugs (Triatominae Jeannel). Reduviidae are morphologically diverse, engage in an astounding array of predatory strategies and have evolved some of the most unique anti-predator and stealth techniques in the animal kingdom. While significant progress has been made to reveal the evolutionary history of assassin bugs and revise their taxonomy, the non-monophyly of the second largest assassin bug subfamily, Reduviinae Latreille, remains to be addressed. Leveraging phylogenomic data (2,291 loci) and 112 morphological characters, we performed the first data- and taxon-rich (195 reduvioid taxa) combined phylogenetic analysis across Reduvioidea and reconstructed morphological diagnostic features for major lineages. We corroborated the rampant polyphyly of Reduviinae that demands substantial revisions to the subfamilial and tribal classification of assassin bugs. Our new classification for Reduviidae reduces the number of subfamilies to 19 and recognizes 40 tribes. We describe three new subfamilies to accommodate distantly related taxa previously classified as Reduviinae (Heteropinae subfam. nov.Nanokeralinae subfam. nov., and Pasirinae subfam. nov.). Triatominae sensu nov. are expanded to include closely related predatory reduviine genera. Cetherinae Jeannel, Chryxinae Champion, Pseudocetherinae Villiers, Salyavatinae Amyot & Serville and Sphaeridopinae Amyot & Serville are treated as junior synonyms of Reduviinae sensu nov. Epiroderinae Distant are synonymized with Phimophorinae Handlirsch sensu nov. and Bactrodini Stål stat. nov. are reclassified as a tribe of Harpactorinae Amyot & Serville. Psophidinae Distant is treated as a valid subfamily. This new classification represents a robust framework for future taxonomic and evolutionary research on assassin bugs.

Keywords: ancestral state reconstruction, Cimicomorpha, phylogenetic systematics, revised classification, taxonomy, true bugs
 
Assassin bug diversity and examples of unique morphological and behavioural adaptations found across the family.
(a) Holoptilinae: Ptilocnemus femoralis Horváth (© Martin Lagerwey, Australia); (b) Phymatinae: Phymata americana Melin male mate guarding female (© Marie-Lise Beaudin, Canada); (c) Peiratinae: Rasahus arcuiger (Stål) (© sandralamberts, Peru);
(d) Psophidinae stat. nov.Korinchocoris insolitus Miller (© Joshua Wong, Singapore); (e) Ectrichodiinae: nr. Ectrichodia Lepeletier and Serville communal feeding of adults and nymphs on a millipede (© Nick Ramsey, Equatorial Guinea); (f) Emesinae: Ghilianella sp. Spinola (© Thomas Shahan, Belize);
(g) Stenopodainae (© Damien Brouste, New Caledonia); (h) Triatominae sensu nov.Zelurini trib. nov.Zelurus festivus (Stål) (© sandralamberts, Peru); (i) Triatominae sensu nov.: Rhodniini: Rhodnius sp. Stål nymph engorged on blood-meal (© César Favacho);
(j) Phimophorinae sensu nov.Physoderes sp. Westwood (© Nick Porch, Republic of Mauritius); (k) Harpactorinae sensu nov.: Ectinoderini: Amulius sp. Stål forelegs coated with resin (© Julius Simonelli, Indonesia); (l) Harpactorinae sensu nov.: Bactrodini stat. nov.Bactrodes sp. Stål (© Stephen_WV, Brazil),
 (m) Reduviinae sensu nov.: Reduviini stat. nov.sensu nov.Reduvius sonoraensis Usinger (© James Bailey, USA); (n) Reduviinae sensu nov.: Acanthaspidini stat. nov.sensu nov.Inara flavopicta Stål nymph with corpse backpack (© budak, Singapore); (o) Reduviinae sensu nov.: Salyavatini stat. nov.Salyavata macmahanae van Doesburg and Brailovsky (© Tom Murray, Honduras).

Habitus plate of newly described reduviid subfamilies and other revived taxa.
(a)–(b) Heteropinae subfam. nov. (a) Heteropinus mollis (UCR_ENT 00115568). (b) Platymicrus sp. (UCR_ENT 00129721).
(c) Nanokeralinae subfam. nov.Hadrokerala major (UCR_ENT 00052179).
(d)–(e). Pasirinae subfam. nov. (d) Pasira sp. (UCR_ENT 00052217). (e) Pasiropsis sp. (UCR_ENT 00052178).

(f) Triatominae sensu nov.: Opisthacidiini trib. nov.: Opisthacidius sp. (UCR_ENT 00012957). (g) Phimophorinae sensu nov.Nalata squalida (UCR_ENT 00002748).
(h)–(j) Reduviinae sensu nov.: Lenaeini stat. nov. (h) Pseudozelurus arizonicus (UCR_ENT 00004573). (i) Dyakocoris vulnerans (UCR_ENT 00052204). (j) Tiarodes versicolor (UCR_ENT 00052171).

(k) Reduviinae sensu nov.: Euphenini stat. nov., sensu nov.Australocleptes sp. (AMNH_PBI 00218999). (l)–(m). Reduviinae sensu nov.: Psyttalini trib. nov. (l) Varus flavoannulatus (UCR_ENT 00004574). (m) Cerilocus nero (UCR_ENT 00129676)
(n)–(o). Reduviinae sensu nov.: Acanthaspidini stat. nov.sensu nov. (n) Plynoides sp. (UCR_ENT 00129678). (o) Eriopreda feai (UCR_ENT 00129696).


Paul K. Masonick, Alex Knyshov, Eric R. L. Gordon, Dimitri Forero, Wei Song Hwang, Rochelle Hoey-Chamberlain, Tatiana Bush, Stephanie Castillo, Madison Hernandez, Jamie Ramirez, Samantha Standring, Junxia Zhang and Christiane Weirauch. 2025. A Revised Classification of the Assassin Bugs (Hemiptera: Heteroptera: Reduviidae) based on combined analysis of phylogenomic and morphological data. Systematic Entomology. DOI: doi.org/10.1111/syen.12646 [17 July 2024]