Project 3448: C. Weirauch, R. T. Schuh, G. Cassis, W. C. Wheeler. 2019. Revisiting habitat and lifestyle transitions in Heteroptera (Insecta: Hemiptera): insights from a combined morphological and molecular phylogeny. Cladistics. 35 (1):67-105.
Abstract
Heteroptera, the true bugs, are part of the largest clade of non‐holometabolous insects, the Hemiptera, and include > 42 000 described species in about 90 families. Despite progress in resolving phylogenetic relationships between and within infraorders since the first combined morphological and molecular analysis published in 1993 (29 taxa, 669 bp, 31 morphological characters), recent hypotheses have relied entirely on molecular data. Weakly supported nodes along the backbone of Heteroptera made these published phylogenies unsuitable for investigations into the evolution of habitats and lifestyles across true bugs. Here we present the first combined morphological and molecular analyses of Heteroptera since 1993, using 135 taxa in 60 families, 4018 aligned bp of ribosomal DNA and 81 morphological characters, and various analytical approaches. The sister‐group relationship of the predominantly aquatic Nepomorpha with all remaining Heteroptera is supported in all analyses, and a clade formed by Enicocephalomorpha, Dipsocoromorpha and Gerromorpha in some. All analyses recover Leptopodomorpha + (Cimicomorpha + Pentatomomorpha), mostly with high support. Parsimony‐ and likelihood‐based ancestral state reconstructions of habitats and lifestyles on the combined likelihood phylogeny provide new insights into the evolution of true bugs. The results indicate that aquatic and semi‐aquatic true bugs invaded these habitats three times independently from terrestrial habitats in contrast to a recent hypothesis. They further suggest that the most recent common ancestor of Heteroptera was predacious, and that the two large predominantly phytophagous clades (Trichophora and Miroidea) are likely to have derived independently from predatory ancestors. We conclude that by combining morphological and molecular data and employing various analytical methods our analyses have converged on a relatively well‐supported hypothesis of heteropteran infraordinal relationships that now requires further testing using phylogenomic and more extensive morphological datasets.Read the article »
Article DOI: 10.1111/cla.12233
Project DOI: 10.7934/P3448, http://dx.doi.org/10.7934/P3448
| This project contains | Matrices |
|---|---|
Download Project SDD File  | Total scored cells: 10796 Total media associated with cells: 0 Total labels associated with cell media: 0 |
| Characters | |
| Total characters: 81 Total characters with associated media: 0 Total characters with media with labels: 0 Total character states: 219 Total character states with associated media: 0 Total character states with media with labels:0 Total unordered/ordered characters:80/1 |
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MorphoBank Project 3448
MorphoBank Project 3448
- Creation Date:
04 April 2019 - Publication Date:
19 April 2019 - Project views: 16116
- Matrix downloads: 9
This research
supported by
Authors' Institutions
- University of California Riverside
- American Museum of Natural History
- University of New South Wales
Members
| member name | taxa |
specimens |
media | chars | character
| cell scorings (scored, NPA, "-") | cell
| rules | ||||||
| MorphoBank Curator Project Administrator | 135 | 1 | 1 | 81 | 0 | 0 | 10796 (10504, 0, 292) | 0 | 0 | 0 | ||||
| Gerasimos Cassis Full membership | 0 | 0 | 0 | 0 | 0 | 0 | 0 (0, 0, 0) | 0 | 0 | 0 | ||||
| Randall Schuh Full membership | 0 | 0 | 0 | 0 | 0 | 0 | 0 (0, 0, 0) | 0 | 0 | 0 | ||||
| Christiane Weirauch Full membership | 0 | 0 | 0 | 0 | 0 | 0 | 0 (0, 0, 0) | 0 | 0 | 0 | ||||
| Ward Wheeler Full membership | 0 | 0 | 0 | 0 | 0 | 0 | 0 (0, 0, 0) | 0 | 0 | 0 | ||||
Taxonomic Overview for Matrix 'M25988' (135 Taxa)
| taxon | unscored cells |
scored cells |
no cell support |
NPA cells |
"-" cells | cell images | labels on cell images |
member access |
| [1] Aeneolamia contigua Last Modified in 04/04/19 | 0 | 77 | 77 | 0 | 4 | 0 | 0 | 5 |
| [2] Locris rubra Last Modified in 04/04/19 | 0 | 77 | 77 | 0 | 4 | 0 | 0 | 5 |
| [3] Tropidocephala sp Last Modified in 04/04/19 | 0 | 77 | 77 | 0 | 4 | 0 | 0 | 5 |
| [4] Amerzanna peruana Last Modified in 04/04/19 | 0 | 78 | 78 | 0 | 3 | 0 | 0 | 5 |
| [5] Putoniessa rivularis Last Modified in 04/04/19 | 0 | 77 | 77 | 0 | 4 | 0 | 0 | 5 |
| [6] Cicadatra atra Last Modified in 04/04/19 | 0 | 77 | 77 | 0 | 4 | 0 | 0 | 5 |
| [7] Hackeriella veitchi Last Modified in 04/04/19 | 2 | 72 | 72 | 0 | 7 | 0 | 0 | 5 |
| [8] Peloridium hammonniorum Last Modified in 04/04/19 | 2 | 72 | 72 | 0 | 7 | 0 | 0 | 5 |
| [9] Aenictopecheidae sp Last Modified in 04/04/19 | 2 | 76 | 76 | 0 | 3 | 0 | 0 | 5 |
| [10] Systelloderes sp Last Modified in 04/04/19 | 1 | 77 | 77 | 0 | 3 | 0 | 0 | 5 |
| [11] Enicocephalinae sp Last Modified in 04/04/19 | 2 | 76 | 76 | 0 | 3 | 0 | 0 | 5 |
| [12] Enicocephalinae nig 137 Last Modified in 04/04/19 | 2 | 76 | 76 | 0 | 3 | 0 | 0 | 5 |
| [13] Enicocephalus sp cf sturmi ecu 150 Last Modified in 04/04/19 | 2 | 76 | 76 | 0 | 3 | 0 | 0 | 5 |
| [14] Ceratocombus australiensis Last Modified in 04/04/19 | 3 | 76 | 76 | 0 | 2 | 0 | 0 | 5 |
| [15] Ceratocombus vagans 247 Last Modified in 04/04/19 | 3 | 76 | 76 | 0 | 2 | 0 | 0 | 5 |
| [16] Cryptostemma sp Last Modified in 04/04/19 | 3 | 77 | 77 | 0 | 1 | 0 | 0 | 5 |
| [17] Cryptostemma sp peru 251 Last Modified in 04/04/19 | 3 | 77 | 77 | 0 | 1 | 0 | 0 | 5 |
| [18] Hypselosoma hickmani Last Modified in 04/04/19 | 3 | 77 | 77 | 0 | 1 | 0 | 0 | 5 |
| [19] Pateena polymitarior Last Modified in 04/04/19 | 3 | 77 | 77 | 0 | 1 | 0 | 0 | 5 |
| [20] Voragocoris schuhi 254 Last Modified in 04/04/19 | 3 | 77 | 77 | 0 | 1 | 0 | 0 | 5 |
| [21] Chinannus sp cr 80 Last Modified in 04/04/19 | 3 | 77 | 77 | 0 | 1 | 0 | 0 | 5 |
| [22] Ptenidiophyes mirabilis cr 89 Last Modified in 04/04/19 | 3 | 75 | 75 | 0 | 3 | 0 | 0 | 5 |
| [23] Mesovelia mulsanti Last Modified in 04/04/19 | 3 | 75 | 75 | 0 | 3 | 0 | 0 | 5 |
| [24] Macrovelia horni Last Modified in 04/04/19 | 7 | 71 | 71 | 0 | 3 | 0 | 0 | 5 |
| [25] Microvelia sp Last Modified in 04/04/19 | 0 | 77 | 77 | 0 | 4 | 0 | 0 | 5 |
| [26] Aquarius remigis Last Modified in 04/04/19 | 0 | 77 | 77 | 0 | 4 | 0 | 0 | 5 |
| [27] Abedus indentatus Last Modified in 04/04/19 | 0 | 77 | 77 | 0 | 4 | 0 | 0 | 5 |
| [28] Ranatra sp Last Modified in 04/04/19 | 0 | 77 | 77 | 0 | 4 | 0 | 0 | 5 |
| [29] Nerthra adspersa Last Modified in 04/04/19 | 0 | 78 | 78 | 0 | 3 | 0 | 0 | 5 |
| [30] Megochterus occidentalis Last Modified in 04/04/19 | 0 | 78 | 78 | 0 | 3 | 0 | 0 | 5 |
| [31] Corixidae sp Last Modified in 04/04/19 | 0 | 77 | 77 | 0 | 4 | 0 | 0 | 5 |
| [32] Usingerina moapensis Last Modified in 04/04/19 | 0 | 77 | 77 | 0 | 4 | 0 | 0 | 5 |
| [33] Hydrotrephes sp Last Modified in 04/04/19 | 0 | 77 | 77 | 0 | 4 | 0 | 0 | 5 |
| [34] Notonectidae sp peru Last Modified in 04/04/19 | 0 | 77 | 77 | 0 | 4 | 0 | 0 | 5 |
| [35] Pleidae sp peru Last Modified in 04/04/19 | 0 | 77 | 77 | 0 | 4 | 0 | 0 | 5 |
| [36] Salda sp Last Modified in 04/04/19 | 0 | 80 | 80 | 0 | 1 | 0 | 0 | 5 |
| [37] Saldula brevicornis Last Modified in 04/04/19 | 0 | 80 | 80 | 0 | 1 | 0 | 0 | 5 |
| [38] Valleriola sp Last Modified in 04/04/19 | 4 | 76 | 76 | 0 | 1 | 0 | 0 | 5 |
| [39] Leotichius sp Last Modified in 04/04/19 | 4 | 76 | 76 | 0 | 1 | 0 | 0 | 5 |
| [40] Aphelonotus fraterculus 1586 Last Modified in 04/04/19 | 7 | 74 | 74 | 0 | 0 | 0 | 0 | 5 |
| [41] Camarochilus 2720 Last Modified in 04/04/19 | 7 | 74 | 74 | 0 | 0 | 0 | 0 | 5 |
| [42] Phymata pacifica Last Modified in 04/04/19 | 0 | 79 | 79 | 0 | 2 | 0 | 0 | 5 |
| [43] Macrocephalus barberi Last Modified in 04/04/19 | 0 | 79 | 79 | 0 | 2 | 0 | 0 | 5 |
| [44] Tapeinus sp Last Modified in 04/04/19 | 3 | 76 | 76 | 0 | 2 | 0 | 0 | 5 |
| [45] Zelurus sp Last Modified in 04/04/19 | 0 | 79 | 79 | 0 | 2 | 0 | 0 | 5 |
| [46] Triatoma protracta Last Modified in 04/04/19 | 0 | 79 | 79 | 0 | 2 | 0 | 0 | 5 |
| [47] Apiomerus lanipes Last Modified in 04/04/19 | 0 | 76 | 76 | 0 | 5 | 0 | 0 | 5 |
| [48] Rhynocoris segmentarius Last Modified in 04/04/19 | 0 | 76 | 76 | 0 | 5 | 0 | 0 | 5 |
| [49] Stenopoda sp Last Modified in 04/04/19 | 0 | 79 | 79 | 0 | 2 | 0 | 0 | 5 |
| [50] Racelda sp Last Modified in 04/04/19 | 0 | 79 | 79 | 0 | 2 | 0 | 0 | 5 |
| [51] Stenolemoides arizonensis Last Modified in 04/04/19 | 0 | 78 | 78 | 0 | 3 | 0 | 0 | 5 |
| [52] Loricula elegantula Last Modified in 04/04/19 | 6 | 74 | 74 | 0 | 1 | 0 | 0 | 5 |
| [53] Joppeicus paradoxus Last Modified in 04/04/19 | 4 | 76 | 76 | 0 | 1 | 0 | 0 | 5 |
| [54] Himacerus apterus Last Modified in 04/04/19 | 0 | 80 | 80 | 0 | 1 | 0 | 0 | 5 |
| [55] Nabis ferus Last Modified in 04/04/19 | 0 | 80 | 80 | 0 | 1 | 0 | 0 | 5 |
| [56] Nabis apicalis Last Modified in 04/04/19 | 0 | 80 | 80 | 0 | 1 | 0 | 0 | 5 |
| [57] Prostemma hilgendorfii Last Modified in 04/04/19 | 0 | 80 | 80 | 0 | 1 | 0 | 0 | 5 |
| [58] Lasiochilus japonicus Last Modified in 04/04/19 | 6 | 74 | 74 | 0 | 1 | 0 | 0 | 5 |
| [59] Lasiochilus luceonotatus Last Modified in 04/04/19 | 6 | 74 | 74 | 0 | 1 | 0 | 0 | 5 |
| [60] Lipokophila eberhardi Last Modified in 04/04/19 | 7 | 73 | 73 | 0 | 1 | 0 | 0 | 5 |
| [61] Lyctocoris beneficus Last Modified in 04/04/19 | 6 | 74 | 74 | 0 | 1 | 0 | 0 | 5 |
| [62] Anthocoris confusus Last Modified in 04/04/19 | 0 | 80 | 80 | 0 | 1 | 0 | 0 | 5 |
| [63] Orius horvathi Last Modified in 04/04/19 | 0 | 80 | 80 | 0 | 1 | 0 | 0 | 5 |
| [64] Tetraphleps aterrimus Last Modified in 04/04/19 | 0 | 80 | 80 | 0 | 1 | 0 | 0 | 5 |
| [65] Cimex lectularius Last Modified in 04/04/19 | 3 | 74 | 74 | 0 | 4 | 0 | 0 | 5 |
| [66] Curalium cronini Last Modified in 04/04/19 | 5 | 73 | 73 | 0 | 3 | 0 | 0 | 5 |
| [67] Baclozygum brachypterum Last Modified in 04/04/19 | 4 | 75 | 75 | 0 | 2 | 0 | 0 | 5 |
| [68] Thaumastocorinae thai 2846 Last Modified in 04/04/19 | 4 | 75 | 75 | 0 | 2 | 0 | 0 | 5 |
| [69] Discocoris drakei Last Modified in 04/04/19 | 4 | 75 | 75 | 0 | 2 | 0 | 0 | 5 |
| [70] Xylastodoris luteolus Last Modified in 04/04/19 | 4 | 75 | 75 | 0 | 2 | 0 | 0 | 5 |
| [71] Corythucha sp Last Modified in 04/04/19 | 0 | 78 | 78 | 0 | 3 | 0 | 0 | 5 |
| [72] Diplocysta sp Last Modified in 04/04/19 | 0 | 78 | 78 | 0 | 3 | 0 | 0 | 5 |
| [73] Eteoneus angulatus Last Modified in 04/04/19 | 0 | 78 | 78 | 0 | 3 | 0 | 0 | 5 |
| [74] Myiomma sp Last Modified in 04/04/19 | 0 | 79 | 79 | 0 | 2 | 0 | 0 | 5 |
| [75] Monalocoris americanus Last Modified in 04/04/19 | 0 | 78 | 78 | 0 | 3 | 0 | 0 | 5 |
| [76] Campyloneura virgula Last Modified in 04/04/19 | 0 | 78 | 78 | 0 | 3 | 0 | 0 | 5 |
| [77] Dicyphus pallidicornis Last Modified in 04/04/19 | 0 | 78 | 78 | 0 | 3 | 0 | 0 | 5 |
| [78] Tupiocoris sp Last Modified in 04/04/19 | 0 | 78 | 78 | 0 | 3 | 0 | 0 | 5 |
| [79] Phallospinophylus setosus Last Modified in 04/04/19 | 0 | 78 | 78 | 0 | 3 | 0 | 0 | 5 |
| [80] Plagiognathus chrysanthemi Last Modified in 04/04/19 | 0 | 78 | 78 | 0 | 3 | 0 | 0 | 5 |
| [81] Rhinacloa forticornis Last Modified in 04/04/19 | 0 | 78 | 78 | 0 | 3 | 0 | 0 | 5 |
| [82] Semium hirtum Last Modified in 04/04/19 | 0 | 78 | 78 | 0 | 3 | 0 | 0 | 5 |
| [83] Pilophorus discretus Last Modified in 04/04/19 | 0 | 78 | 78 | 0 | 3 | 0 | 0 | 5 |
| [84] Lopidea bullata Last Modified in 04/04/19 | 0 | 78 | 78 | 0 | 3 | 0 | 0 | 5 |
| [85] Pseudopsallus angularis Last Modified in 04/04/19 | 0 | 78 | 78 | 0 | 3 | 0 | 0 | 5 |
| [86] Capsus ater Last Modified in 04/04/19 | 0 | 78 | 78 | 0 | 3 | 0 | 0 | 5 |
| [87] Cyphopelta modesta Last Modified in 04/04/19 | 0 | 78 | 78 | 0 | 3 | 0 | 0 | 5 |
| [88] Irbisia brachycera Last Modified in 04/04/19 | 0 | 78 | 78 | 0 | 3 | 0 | 0 | 5 |
| [89] Lygus lineolaris Last Modified in 04/04/19 | 0 | 78 | 78 | 0 | 3 | 0 | 0 | 5 |
| [90] Trigonotylus sp Last Modified in 04/04/19 | 0 | 78 | 78 | 0 | 3 | 0 | 0 | 5 |
| [91] Deraeocoris brevis Last Modified in 04/04/19 | 0 | 78 | 78 | 0 | 3 | 0 | 0 | 5 |
| [92] Calisius hackeri Last Modified in 04/04/19 | 1 | 78 | 78 | 0 | 2 | 0 | 0 | 5 |
| [93] Dysodius sp Last Modified in 04/04/19 | 1 | 78 | 78 | 0 | 2 | 0 | 0 | 5 |
| [94] Isodermus planus Last Modified in 04/04/19 | 1 | 78 | 78 | 0 | 2 | 0 | 0 | 5 |
| [95] Mezira sayi Last Modified in 04/04/19 | 1 | 78 | 78 | 0 | 2 | 0 | 0 | 5 |
| [96] Anischys luteovarius Last Modified in 04/04/19 | 0 | 80 | 80 | 0 | 1 | 0 | 0 | 5 |
| [97] Stauralia chloracantha Last Modified in 04/04/19 | 0 | 80 | 80 | 0 | 1 | 0 | 0 | 5 |
| [98] Lestonia haustorifera Last Modified in 04/04/19 | 0 | 78 | 78 | 0 | 3 | 0 | 0 | 5 |
| [99] Parastrachia japonensis Last Modified in 04/04/19 | 0 | 80 | 80 | 0 | 1 | 0 | 0 | 5 |
| [100] Cydninae sp 3 Last Modified in 04/04/19 | 0 | 80 | 80 | 0 | 1 | 0 | 0 | 5 |
| [101] Brochymena sp Last Modified in 04/04/19 | 0 | 80 | 80 | 0 | 1 | 0 | 0 | 5 |
| [102] Oechalia schellenbergii Last Modified in 04/04/19 | 0 | 80 | 80 | 0 | 1 | 0 | 0 | 5 |
| [103] Poecilometis sp Last Modified in 04/04/19 | 0 | 80 | 80 | 0 | 1 | 0 | 0 | 5 |
| [104] Tholosanus sp Last Modified in 04/04/19 | 0 | 80 | 80 | 0 | 1 | 0 | 0 | 5 |
| [105] Austrotichus rugosus Last Modified in 04/04/19 | 0 | 79 | 79 | 0 | 2 | 0 | 0 | 5 |
| [106] Choerocoris variegatus Last Modified in 04/04/19 | 0 | 79 | 79 | 0 | 2 | 0 | 0 | 5 |
| [107] Coleotichus costatus Last Modified in 04/04/19 | 0 | 79 | 79 | 0 | 2 | 0 | 0 | 5 |
| [108] Thaumastella namaquensis Last Modified in 04/04/19 | 0 | 80 | 80 | 0 | 1 | 0 | 0 | 5 |
| [109] Urochela luteovaria Last Modified in 04/04/19 | 0 | 80 | 80 | 0 | 1 | 0 | 0 | 5 |
| [110] Urostylis westwoodi Last Modified in 04/04/19 | 0 | 80 | 80 | 0 | 1 | 0 | 0 | 5 |
| [111] Henicocoris monteithi Last Modified in 04/04/19 | 1 | 78 | 78 | 0 | 2 | 0 | 0 | 5 |
| [112] Trisecus sp Last Modified in 04/04/19 | 0 | 79 | 79 | 0 | 2 | 0 | 0 | 5 |
| [113] Kleidocerys sp Last Modified in 04/04/19 | 0 | 80 | 80 | 0 | 1 | 0 | 0 | 5 |
| [114] Lygaeus equestris Last Modified in 04/04/19 | 0 | 80 | 80 | 0 | 1 | 0 | 0 | 5 |
| [115] Oncopeltus fasciatus Last Modified in 04/04/19 | 0 | 80 | 80 | 0 | 1 | 0 | 0 | 5 |
| [116] Malcus elongatus Last Modified in 04/04/19 | 0 | 80 | 80 | 0 | 1 | 0 | 0 | 5 |
| [117] Pachygrontha sp Last Modified in 04/04/19 | 0 | 80 | 80 | 0 | 1 | 0 | 0 | 5 |
| [118] Mcateella sp Last Modified in 04/04/19 | 0 | 80 | 80 | 0 | 1 | 0 | 0 | 5 |
| [119] Neoneides muticus Last Modified in 04/04/19 | 0 | 80 | 80 | 0 | 1 | 0 | 0 | 5 |
| [120] Pronotacantha annulata Last Modified in 04/04/19 | 0 | 79 | 79 | 0 | 2 | 0 | 0 | 5 |
| [121] Ischnodemus sp Last Modified in 04/04/19 | 0 | 80 | 80 | 0 | 1 | 0 | 0 | 5 |
| [122] Cymus sp Last Modified in 04/04/19 | 0 | 80 | 80 | 0 | 1 | 0 | 0 | 5 |
| [123] Germalus sp Last Modified in 04/04/19 | 0 | 80 | 80 | 0 | 1 | 0 | 0 | 5 |
| [124] Remaudiereana inornata Last Modified in 04/04/19 | 0 | 81 | 81 | 0 | 0 | 0 | 0 | 5 |
| [125] Laryngodus cervantes Last Modified in 04/04/19 | 0 | 81 | 81 | 0 | 0 | 0 | 0 | 5 |
| [126] Leptocorisa acuta Last Modified in 04/04/19 | 0 | 80 | 80 | 0 | 1 | 0 | 0 | 5 |
| [127] Cletus sp Last Modified in 04/04/19 | 0 | 80 | 80 | 0 | 1 | 0 | 0 | 5 |
| [128] Aulacosternum nigrorubrum Last Modified in 04/04/19 | 0 | 80 | 80 | 0 | 1 | 0 | 0 | 5 |
| [129] Narnia femorata Last Modified in 04/04/19 | 0 | 80 | 80 | 0 | 1 | 0 | 0 | 5 |
| [130] Maevius indecorus Last Modified in 04/04/19 | 0 | 80 | 80 | 0 | 1 | 0 | 0 | 5 |
| [131] Harmostes sp Last Modified in 04/04/19 | 0 | 80 | 80 | 0 | 1 | 0 | 0 | 5 |
| [132] Dysdercus fuscomaculatus Last Modified in 04/04/19 | 0 | 79 | 79 | 0 | 2 | 0 | 0 | 5 |
| [133] Probergrothius sexpunctatus Last Modified in 04/04/19 | 0 | 79 | 79 | 0 | 2 | 0 | 0 | 5 |
| [134] Largus rufipennis Last Modified in 04/04/19 | 0 | 79 | 79 | 0 | 2 | 0 | 0 | 5 |
| [135] Physopelta melanoptera Last Modified in 04/04/19 | 0 | 79 | 79 | 0 | 2 | 0 | 0 | 5 |
Project views
| type | number of views | Individual items viewed (where applicable) |
| Total project views | 16116 | |
| Project overview | 1256 | |
| Media views | 2418 | Media search (2060 views); M662094 (358 views); |
| Matrix views | 835 | Morphological matrix used for the phylogenetic analyses NEXUS file. (121 views); Matrix landing page (714 views); |
| Taxon list | 9149 | |
| Bibliography | 491 | |
| Views for media list | 766 | |
| Specimen list | 1196 | |
| Documents list | 5 |
Project downloads
| type | number of downloads | Individual items downloaded (where applicable) |
| Total downloads from project | 164 | |
| Project downloads | 155 | |
| Matrix downloads | 9 | Morphological matrix used for the phylogenetic analyses NEXUS file. (9 downloads); |