Erthesina fullo (Hemiptera: Heteroptera: Pentatomidae): old and new stories of an invasive stink bug.

The Yellow-Spotted Stink Bug Erthesina fullo (Thunberg, 1783) (Pentatomomorpha: Pentatomoidea: Pentatomidae: Pentatominae: Halyini), native in East and South-East Asia, was recently discovered introduced in both Europe (Albania, 2017) and South America (Brazil: So Paulo, 2020). We report two cases of interception of living specimens in the Czech Republic. The first specimen (Zln, 2021) was found in a sealed container with shoes imported from China, the second specimen (Prague, 2023) was recieved in a parcel sent from Poland. We also present the interesting story of Erthesina fullo in Japan, where the species was introduced to Dejima Island in Nagasaki (Kyushu) already before 1775; at that time Dejima was the only place in Japan where Dutch and Chinese ships were allowed to trade during the isolationist Edo period (16031868). One old name, also proposed based on specimens originating from Japan, Cimex aloes Lichtenstein, 1796, syn. nov., is proposed a junior subjective synonym of E. fullo. The invasion of Erthesina fullo in Japan seems to be the oldest exactly localised invasion in Heteroptera. The taxonomy of Erthesina Spinola, 1837 is discussed, the genus currently includes 9 species. The genus is in urgent need of revision and the resulting problems with identification of its species, including E. fullo, are pointed out. Distribution of all Erthesina species is briefly reviewed.

Dimorphopterus blissoides (Hemiptera: Heteroptera: Blissidae): recent spreading of a neonative species in the Pannonian Basin.

Dimorphopterus blissoides (Baerensprung, 1859) (Hemiptera: Heteroptera: Blissidae) is recorded for the first time from Slovakia and Iran. We provide also additional records of D. blissoides from Bulgaria, Croatia, Greece, Hungary and Romania, and review the distribution of the species. The species is currently spreading in the Pannonian Basin, reaching its northern distribution limit in southern Slovakia. We confirm its development on Phragmites australis, with preference to the loose outer leaf sheaths of the galls of Lipara lucens Meigen, 1830 (Diptera: Chloropidae), where the specimens develop during summer as well as hibernate during winter.

Diversification of the phytophagous lineages of true bugs (Insecta: Hemiptera: Heteroptera) shortly after that of the flowering plants.

More than 95% of phytophagous true bug (Hemiptera: Heteroptera) species belong to four superfamilies: Miroidea (Cimicomorpha), Pentatomoidea, Coreoidea, and Lygaeoidea (all Pentatomomorpha). These iconic groups of highly diverse, overwhelmingly phytophagous insects include several economically prominent agricultural and silvicultural pest species, though their evolutionary history has not yet been well resolved. In particular, superfamily- and family-level phylogenetic relationships of these four lineages have remained controversial, and the divergence times of some crucial nodes for phytophagous true bugs have hitherto been little known, which hampers a better understanding of the evolutionary processes and patterns of phytophagous insects. In the present study, we used 150 species and concatenated nuclear and mitochondrial protein-coding genes and rRNA genes to infer the phylogenetic relationships within the Terheteroptera (Cimicomorpha + Pentatomomorpha) and estimated their divergence times. Our results support the monophyly of Cimicomorpha, Pentatomomorpha, Miroidea, Pentatomoidea, Pyrrhocoroidea, Coreoidea, and Lygaeoidea. The phylogenetic relationships across phytophagous lineages are largely congruent at deep nodes across the analyses based on different datasets and tree-reconstructing methods with just a few exceptions. Estimated divergence times and ancestral state reconstructions for feeding habit indicate that phytophagous true bugs explosively radiated in the Early Cretaceous-shortly after the angiosperm radiation-with the subsequent diversification of the most speciose clades (Mirinae, Pentatomidae, Coreinae, and Rhyparochromidae) in the Late Cretaceous.

Joppeicus paradoxus (Hemiptera: Heteroptera: Joppeicidae): a new alien species in the European Union?

Joppeicus paradoxus Puton, 1881 is the only representative of the family Joppeicidae. This tiny predaceous bug of around 3 mm long has a resemblance to members of the family Anthocoridae sensu lato. Herein we present an up-to-date distribution of the species, recording it for the first time in the Canary Islands (Fuerteventura Island) and in Yemen (Socotra Island). We provide pictures of the dorsal and lateral habitus of the species and discuss its current distribution. The collection circumstances suggest probably native occurrence in Socotra, but in Fuerteventura it was found in an urban environment with a fair chance it was introduced due to human activity. Details on the types of J. paradoxus are also provided.

Bizarre morphology in extinct Eocene bugs (Heteroptera: Pentatomidae).

Newly discovered fossil bugs (Insecta: Hemiptera: Heteroptera: Pentatomidae) from the Eocene of Messel (Germany) and Green River (North America) exhibit an exaggerated morphology including prominent spiny humeral and anterolateral angles of the pronotum and a spiny lateral abdominal margin. Especially the humeral angles are unique; they consist of expansive, rounded projections with strong spines, which is a rare trait among pentatomids. A hypothesis for the function of this extreme morphology is defence against small vertebrate predators, such as birds or reptiles. The same protuberances also produce a disruptive effect camouflaging the specimen in its environment and provide additional protection. Therefore, the extreme morphology provides primary as well as secondary anti-predator defence. The morphology of Eospinosus peterkulkai gen. et sp. nov. and E. greenriverensis sp. nov. resembles that of Triplatygini, which today occur exclusively in Madagascar, as well as that of Discocephalinae or Cyrtocorinae, which today occur in the Neotropics. Due to a lack of conclusive characters, it cannot be excluded that the fossil species may represent a case of remarkable convergence and are not related to either taxon. Phylogenetic analyses using parsimony as well as Bayesian algorithms confirmed that the new genus is a member of Pentatomidae, but could not solve its phylogenetic relationships within Pentatomidae.

Halyomorpha picus (Hemiptera: Heteroptera: Pentatomidae): first confirmed record from Pakistan and two new junior synonyms.

We provide the first confirmed record of Halyomorpha picus (Fabricius, 1794) (Hemiptera: Heteroptera: Pentatomoidea: Pentatomidae: Pentatominae: Cappaeini) from Pakistan: Islamabad Capital Territory, and provide habitus photographs and electron scanning micrographs of the male genitalia of the voucher specimens. Two species previously described from Pakistan are found to fit within the morphological variability of H. picus and the following two junior subjective synonyms are proposed: Halyomorpha picus (Fabricius, 1794) = Halyomorpha punjabensis Ahmad Kamaluddin, 1977, syn. nov., = Halyomorpha azhari Ahmad Zaidi, 1989, syn. nov. The record of Halyomorpha scutellata Distant, 1879, from Pakistan by Sharif et al. (2020) is based on misidentification of a species of Neohalys Ahmad Perveen, 1982 (Pentatominae: Halyini) and excluded from Pakistan fauna. Based on the analysis of female genitalia figures provided by Gadalla (2004), the record of H. picus from Egypt represents a misidentification of H. halys Stl, 1855, which extends the distribution of that invasive species to North Africa.

External structures of the metathoracic scent gland efferent system in the true bug superfamily Pyrrhocoroidea (Hemiptera: Heteroptera: Pentatomomorpha).

Pyrrhocoroidea represents an important group of true bugs (Insecta: Hemiptera: Heteroptera) which includes fire bugs, cotton stainers and other taxa widely used in experimental studies or known as pests. However, the morphology and phylogeny of Pyrrhocoroidea have been only poorly studied so far. Here, structures of the external scent efferent system of the metathoracic scent glands are examined in 64 out of 71 currently valid genera of Pyrrhocoroidea and scanning electron micrographs are provided for most taxa. Several characters are revealed which define each of the three higher taxa within Pyrrhocoroidea: Larginae (small auriculate peritreme lacking manubrium and median furrow; metathoracic spiracle never surrounded by evaporatorium), Physopeltinae (large, widely open ostiole; large peritremal disc with manubrium [new term], lacking median furrow; mace-like mycoid filter processes of equal shape and size on both anterior and posterior margins of metathoracic spiracle), and Pyrrhocoridae (elongate auriculate peritreme with deep median furrow). Within Pyrrhocoridae, three main types (A, B and C) of the external scent efferent system are distinguished, differring in the amount of reductions. The findings are interpreted in the context of phylogenetic hypotheses available for Pyrrhocoroidea and their close relatives, Coreoidea and Lygaeoidea. An updated identification key to the families and subfamilies of Pyrrhocoroidea applicable for both sexes is provided.

On species of the genus emMacrosaldula/em (Hemiptera: Heteroptera: Saldidae) of the Palaearctic Region, with description of emM. graziae/em sp. nov.

In this paper, we present new data on the genus Macrosaldula Leston Southwood, 1964 based on material held in the collections of the National Museum (Prague, Czech Republic) and the Zoological Institute of the Russian Academy of Sciences (St. Petersburg, Russia). Macrosaldula graziae sp. nov., from South Uzbekistan and Tajikistan, is described. Among the Central Asian species of the genus, it is close to M. tadzhika (Kiritshenko, 1912) by the presence of long erect setae on the body dorsum, but the latter species differs in the smaller body size, the presence of light spots on the corium, and in the structure of the male genitalia. In extremely dark specimens of M. jakowleffi (Reuter, 1891), the hemelytra are opaque, with short erect setae. Siberian M. rivularia (J. Sahlberg, 1878), M. simulans Cobben, 1985 and the Far Eastern M. koreana (Kiritshenko, 1912) and M. violacea Cobben, 1985 are distinguished from the new species by the short pubescence on the dorsum. We provide new distributional data for M. clavalis Cobben, 1985 (Georgia), M. jakowleffi (Reuter, 1891) (China: Xinjiang: Altai Mts.-first record), M. miyamotoi Cobben, 1985 (Japan: Honshu), M. nivalis (Lindberg, 1935) (Afghanistan-first record, Kyrgyzstan, Tajikistan), M. scotica (Curtis, 1835) (Georgia: Adzharia; Russia: Murmansk District), and M. tadzhika (Afghanistan-first record, Tajikistan, Uzbekistan). The record of M. scotica from Uzbekistan is disconsidered, and the two specimens are referred to M. graziae sp. nov.

The genera emAgathocles/em and emSurenus/em (Hemiptera: Heteroptera: Pentatomidae): tribal reassessment, redescription, new synonyms, and description of two new species.

The genus Surenus Distant, 1901 (Hemiptera: Heteroptera: Pentatomoidea: Pentatomidae: Pentatominae: Halyini) is revisited and found to be a junior subjective synonym of the genus Agathocles Stål, 1876 (currently Pentatominae: Rolstoniellini). The genus Agathocles and its type species, Agathocles limbatus Stål, 1876, are redescribed. Lectotype of Surenus normalis Distant, 1901 (= Agathocles normalis (Distant, 1901) comb. nov.) is designated and the unknown male of the species is described. Agathocles yunnanensis Zhang Lin, 1984, syn. nov., is considered junior subjective synonym of A. limbatus. Two new species are described: Agathocles flavipes sp. nov. from India (Andhra Pradesh, Karnataka, Maharashtra, and Tamil Nadu) and A. joceliae sp. nov. from Malaysia (Kelantan, Perak). The new species differ from their congeners mainly by the morphology of mandibular plates, length of antennomeres I, IIa and IIb, body length, and structure of male genitalia. Agathocles dubius Distant, 1921 is transferred to the genus Caystrus Stål, 1861 (Pentatominae: Caystrini) based on examination of its holotype with the resulting new combination: Caystrus dubius (Distant, 1921), comb. nov. One new combination is proposed, Paramecocoris ruficornis (Fieber, 1851), comb. nov. (from preoccupied Paramecus Fieber, 1851), and its type locality is clarified as Tenasserim (south Myanmar). Gender agreement and authorship of the name Riazocoris niger Ahmad Afzal, 1977 in Ahmad et al. (1977: 161) are corrected and status of its name bearing type is clarified as lectotype. The following new distribution records are given: A. limbatus from Cambodia, China (Guangxi, Tibet), Laos and Thailand, A. normalis, Caystrus obscurus (Distant, 1901a) and Critheus lineatifrons Stål, 1869 from Laos, Amasenus corticalis Stål, 1863 from Cambodia, Indonesia (E Kalimantan), Laos, Myanmar and Thailand, and Rolstoniellus boutanicus (Dallas, 1849) from Vietnam. Based on characters of external morphology and genitalia, the genus Agathocles is compared with representatives of the genera Halys Fabricius, 1803 (Halyini), Caystrus (Caystrini), Laprius Stål, 1861 (Myrocheini), and Exithemus Distant, 1902 (currently in Rolstoniellini). As a result, the genus Agathocles is here transferred to the tribe Caystrini. The genus Kyrtalus Van Duzee, 1929 is tentatively placed in Myrocheini based on the presence of sulcate mesosternum and femora provided with teeth.

Myxobdella socotrensis sp. nov., a new parasitic leech from Socotra Island, with comments on the phylogeny of Praobdellidae (Hirudinida: Arhynchobdellida).

The Socotra Archipelago (Yemen) is remarkable for unique ecosystems and high endemism, for which it is often referred to as the "Galápagos of the Indian Ocean". Here we describe a new parasitic leech Myxobdella socotrensis sp. nov. from Socotra, the largest island of the archipelago. The new species was found in a freshwater spring attached to the endemic crab Socotrapotamon socotrensis (Hilgendorf, 1883) (Crustacea: Potamidae). Based on its morphology, ecology and a phylogenetic analysis of DNA sequence data (12S, 18S, 28S and COI gene markers), the new leech species is classified into the highly diversified family Praobdellidae (Hirudinida: Arhynchobdellida), distributed in tropical regions of Africa, Asia and Central and South America. The praobdellid leeches are known to infest mammalian mucous membranes, however, some taxa are associated with other hosts, namely crabs, amphibians and birds. By its morphology, the new species fits quite well in the current concept of the genus Myxobdella Oka, 1917. However, the monophyly of Myxobdella was not supported here by molecular data, pointing at the need of a more comprehensive systematic revision of the genus and family. As far as known, Myxobdella socotrensis sp. nov. is endemic to the Socotra Island, but more data are needed to understand its evolutionary origin, biology and distribution.

A review of the fossil taxa originally assigned to Pyrrhocoridae (Hemiptera: Heteroptera).

The four described fossil taxa originally assigned to Pyrrhocoridae (Hemiptera: Heteroptera) are reviewed. Mesopyrrhocoris fasciatus Hong Wang, 1990 (correction of gender agreement) from the Lower Cretaceous of Laiyang Basin, Shandong, China, was reclassified as Cimicomorpha incertae sedis by Shcherbakov (2008), an opinion confirmed here. The status of Dysdercus cinctus Scudder, 1890 and Dysdercus unicolor Scudder, 1890 from the Eocene of Florissant, Colorado, USA, and their placement in Pyrrhocoridae, are doubtful. Pyrrhocoris rottensis nom nov. (= Pyrrhocoris tibialis Statz Wagner, 1950) from the Upper Oligocene of Rott, Germany, is reclassified here as Lygaeoidea incertae sedis due to the presence of ocelli in the fossil. As a result, currently there is no fossil taxon which can be placed in Pyrrhocoroidea with certainty. The extant Pyrrhocoris tibialis Stl, 1874 is confirmed as junior subjective synonym of P. sibiricus Kuschakewitsch, 1866.

Morphological diversity of the metathoracic spiracle in the Lygaeoidea (Hemiptera: Heteroptera).

Spiracles are the openings in the exoskeleton of insects through which air enters into the respiratory system that is formed by a series of tubes called tracheae. They are primarily located on the abdomen, but can also occur on the thorax, including the metathorax. An insect metathoracic spiracle is usually composed of an external opening and a more internal filter apparatus. We propose new terminology for these structures, and we explore the value in their use in taxonomic and phylogenetic studies within the true bug infraorder Pentatomomorpha, with emphasis on the superfamily Lygaeoidea (Insecta: Hemiptera: Heteroptera). These structures were studied using scanning electron microscopy. Two types of metathoracic spiracle external openings were recognized: a narrow opening (type N), which is slit-like; and a wide opening (type W), with internal fine structures located between the mesothoracic and metathoracic margins of the interpleural suture clearly visible. The filter apparatus in the Pentatomomorpha consists of modified mushroom bodies of the metathoracic scent gland evaporatorium, for which the term mycoid filter processes is proposed. Eight different types of mycoid filter processes, and an unmodified microsculpture type (a type with usual cuticular microsculpture) and filter setae can be found on the anterior or posterior margins of the metathoracic spiracle. We believe the wide opening (type W) to be the plesiomorphic character state in the Pentatomomorpha, with multiple, independent transformations leading to the narrow opening in Lygaeoidea. Considerable variability in the structure of the spiracle opening (in Lygaeoidea), and in the structure of the mycoid filter processes (in Pentatomomorpha) was detected. Overall, we found the morphology of these structures to be of limited value concerning the taxonomy or for determining phylogenetic relationships of the higher taxa (families) of Pentatomomorpha, but they may be useful as additional evidence for taxonomic and phylogenetic studies at the generic and perhaps the tribal levels.

Aphelonotus schuhi sp. nov., the first s--pecies of the family Pachynomidae from Puerto Rico (Hemiptera: Heteroptera).

Aphelonotus schuhi sp. nov. (Hemiptera: Heteroptera: Pachynomidae: Aphelonotinae) is described from Puerto Rico based on a single male collected by flight interception trap. The new species is morphologically similar to A. taino Schuh, Weirauch Grillo, 2015 and A. xenos Schuh, Weirauch Grillo, 2015, being distinguished from them by the combination of the presence of a stout subapical spine on the protibia and the characteristic shape of the parameres. An informal A. xenos species-group is proposed to accommodate these three species.

Insect trypanosomatids in Papua New Guinea: high endemism and diversity.

The extreme biological diversity of Oceanian archipelagos has long stimulated research in ecology and evolution. However, parasitic protists in this geographic area remained neglected and no molecular analyses have been carried out to understand the evolutionary patterns and relationships with their hosts. Papua New Guinea (PNG) is a biodiversity hotspot containing over 5% of the world's biodiversity in less than 0.5% of the total land area. In the current work, we examined insect heteropteran hosts collected in PNG for the presence of trypanosomatid parasites. The diversity of insect flagellates was analysed, to our knowledge for the first time, east of Wallace's Line, one of the most distinct biogeographic boundaries of the world. Out of 907 investigated specimens from 138 species and 23 families of the true bugs collected in eight localities, 135 (15%) were infected by at least one trypanosomatid species. High species diversity of captured hosts correlated with high diversity of detected trypanosomatids. Of 46 trypanosomatid Typing Units documented in PNG, only eight were known from other geographic locations, while 38 TUs (~83%) have not been previously encountered. The widespread trypanosomatid TUs were found in both widely distributed and endemic/sub-endemic insects. Approximately one-third of the endemic trypanosomatid TUs were found in widely distributed hosts, while the remaining species were confined to endemic and sub-endemic insects. The TUs from PNG form clades with conspicuous host-parasite coevolutionary patterns, as well as those with a remarkable lack of this trait. In addition, our analysis revealed new members of the subfamilies Leishmaniinae and Strigomonadinae, potentially representing new genera of trypanosomatids.

High Prevalence and Endemism of Trypanosomatids on a Small Caribbean Island.

We describe the monoxenous trypanosomatids parasitizing true bugs and flies on the island of Curaçao. Out of 248 examined true bugs belonging to 17 species, 93 individuals were found to be infected (overall 38% prevalence) by at least one trypanosomatid species (referred to as typing units; TUs). Out of 80 flies, six were infected. All detected trypanosomatids were compared based on their 18S rRNA sequences with TUs parasitizing bugs and flies described from mainland South America, allowing us to assess their diversity and distribution. Besides Leptomonas pyrrhocoris and Leptomonas seymouri, two known species of the subfamily Leishmaniinae, our analysis revealed six new TUs falling into the groups 'jaculum', Blastocrithidia and Herpetomonas. Moreover, two new members of the genus Phytomonas and three new TUs belonging to the monophyletic group designated as 'new clade II' sensu Mol. Phylogenet. Evol, 69, 255 (2013) were isolated. The detected trypanosomatids were characterized by moderate diversity (13 TUs) species richness. Out of nine and four TUs from the heteropteran and dipteran hosts, respectively, 11 TUs have not been encountered before. Although a sampling bias may partially affect the comparison between trypanosomatid communities on Curaçao and the mainland, the high proportion of unique TUs from the former location suggests that the prominent role of islands in increasing the global diversity of macroscopic organisms may also extend to their protistan parasites.

Review of the Neotropical genus Rhyncholepta with descriptions of three new species-group taxa (Hemiptera, Heteroptera, Pentatomidae).

The genus Rhyncholepta Bergroth, 1911 (Hemiptera: Heteroptera: Pentatomidae: Pentatominae: Chlorocorini) is redescribed and five species-group taxa are recognized, keyed, their diagnostic characters illustrated, and the distribution reviewed. Among the five taxa, two species and one subspecies are recognized as new: Rhyncholeptagrandicallosagrandicallosa Bergroth, 1911 (Brazil, Ecuador, French Guiana, Guyana, Peru, Suriname), Rhyncholeptagrandicallosacentroamericana subsp. n. (Belize, Costa Rica, Guatemala, Mexico, Panama), Rhyncholeptahenryi sp. n. (French Guiana), Rhyncholeptameinanderi Becker & Grazia-Vieira, 1971 (Bolivia, Brazil, Ecuador, Peru), and Rhyncholeptawheeleri sp. n. (Guyana). The structure of the male genital capsule was found to be the only reliable character for identifying species-group taxa. For this reason, a simultaneous application has been submitted to the International Commission on Zoological Nomenclature to set aside the non-informative female lectotype of Rhyncholeptagrandicallosagrandicallosa and replace it with the male neotype suggested herein. Based on the available label data and our field experience, most of the specimens were collected by various types of light traps in or near dense forests. Adults can be collected throughout the year.

Psallusthomashenryi sp. n. and Psalluslucanicus from Turkey (Hemiptera, Heteroptera, Miridae).

Psallus (Psallus) thomashenryisp. n. (Hemiptera: Heteroptera: Miridae: Phylinae: Phylini: Phylina) is described from southern Anatolia, Turkey. Illustrations of the dorsal habitus and male genitalia are provided. Its habitus is similar to other uniformly orange species of the subgenus Psallus Fieber, 1858, particularly P.asthenicus Seidenstücker, 1966 from which it can be easily distinguished by the combination of extremely small size (2.3 mm in both sexes) and different morphology of the vesica. Psallus (Psallus) lucanicus Wagner, 1968 is recorded for the first time from Turkey. Psallus (Psallus) aurora (Mulsant & Rey, 1852) is removed from the list of Turkish fauna based on a reevaluation of the voucher specimen. An updated checklist of the species of Psallus known to occur in Turkey is provided. The relevance of Anatolia and the Syro-anatolian-transcaucasian region in the Palearctic distribution of Psallus is discussed. The westernmost record of another mirid, Plagiognathusmarivanensis Linnavuori, 2010, is provided.

Water striders (Heteroptera: Gerromorpha: Gerridae) of Romania with an update on the distribution of Gerris gibbifer and G. maculatus in southeastern Europe.

The Romanian fauna comprises two species of Aquarius Schellenberg, 1800, eight species of Gerris Fabricius, 1794 and one species of Limnoporus Stål, 1868, and we hereby update the distribution and provide insights on the phenology and ecology of all eleven species in this country. We furthermore update the distribution of the two closely related species Gerris gibbifer Schummel, 1832 and G. maculatus Tamanini, 1946 in southeastern Europe. Gerris maculatus is recorded for the first time from Hungary, Montenegro and Slovenia, and the first detailed localities from Romania and Serbia are given. All bibliographic records of G. gibbifer from Romania, Macedonia and Serbia are based on misidentification and this species is thus excluded from the faunal lists of these countries. Both G. gibbifer and G. maculatus occur in Croatia, Hungary, Ukraine, and probably Slovenia.

Phylogeny and the colourful history of jewel bugs (Insecta: Hemiptera: Scutelleridae).

Members of the family Scutelleridae (Heteroptera: Pentatomomorpha: Pentatomoidea) are also called shield bugs because of the greatly enlarged scutellum, or jewel bugs because of the brilliant colours of many species. All scutellerids are phytophagous, feeding on various parts of their host plants. Due to lack of obvious synapomorphies and the failure to apply rigorous phylogenetic methods, the higher classification of Scutelleridae has been disputed for more than 150 years. Here we reconstructed a phylogeny of Scutelleridae based on complete sequences of 18S and 28S nuclear rDNAs and all 13 protein-coding genes of the mitochondrial genome, with the sampled taxa covering all of the currently recognized subfamilies. The monophyly of Scutelleridae was confirmed by the congruence of the results of analyses conducted using Bayesian inference, maximum likelihood and maximum parsimony. The phylogenetic relationships among subfamilies were well resolved for the first time. Furthermore, time-divergence studies estimated that the time of origin of Scutelleridae was in the Early Cretaceous (142.1-122.8 Ma), after the origin of the angiosperms. The diversification between the extant subfamilies of Scutelleridae and within the subfamilies occurred from the late Palaeocene to the late Miocene, simultaneously with the rise of the major groups of angiosperms and other phytophagous insects.