Maternal Care under Large Clutches with Small Eggs: The Evolution of Life History Traits in Shield Bugs.

AbstractPatterns in the correlated evolution of parental care and life history traits are long established but controversial. Although parental care is related to large egg size in many taxa, conflicting results have also been reported. To test the evolutionary relationships between parental care and life history traits, we performed phylogenetic comparative analyses using shield bugs (Heteroptera: Acanthosomatidae), in which maternal guarding of eggs and young has repeatedly evolved. Our analyses revealed that female body size affected reproductive resource allocation. Contrary to the expectations of current theories, the acquisition of maternal care was associated with small eggs, large clutches, and large egg resource allocation. There was a greater trade-off between egg size and clutch size in caring species than in noncaring species. Egg and hatchling developmental rates were not correlated with egg size but were slower in caring species than in noncaring species. Analyses of evolutionary transitions suggest that the establishment of large clutches, small eggs, and large egg resource allocation preceded the evolution of maternal care. To our knowledge, this is the first study clarifying the evolution of parental care linked with small eggs in invertebrates.

New genera and new species of remarkably large-sized or uniquely-shaped mirine plant bugs from Taiwan (Heteroptera: Miridae: Mirinae).

New taxa are proposed and described for several large-sized or uniquely-shaped mirine plant bugs from Taiwan or the northeastern Oriental Region. Two new genera, Rhopalimiris n. gen. and Taiwanocapsus n. gen. are described to accommodate three new species, Rhopalimiris amplissimus n. sp., R. gagai n. sp. and Taiwanocapsus luteocordatus n. sp., which are diagnosed and described. In addition, new species of Eocalocoris Miyamoto & Yasunaga and Eurystylus Stål, namely Eo. gotohi n. sp. and Eu. yixuanae n. sp., are also described. An updated checklist of the Taiwanese Mirini is also provided.

A revision of Scutellera (Hemiptera: Heteroptera: Scutelleridae).

The genus Scutellera Lamarck, 1801 (Hemiptera: Heteroptera: Scutelleridae: Scutellerinae: Scutellerini) is revised. Three species are recognized: Scutellera nepalensis (Westwood, 1837), with two subspecies, S. nepalensis nepalensis and S. nepalensis amethystina (Germar, 1839), stat. nov.; S. perplexa (Westwood, 1837); and S. spilogastra (Walker, 1867), comb. nov. (=S. brevirostris Breddin, 1909, syn. nov.). The included species are redescribed, illustrated, keyed, their bionomics and distribution are reviewed. Lectotypes are designated for Tectocoris nepalensis Westwood, 1837, Calliphara amethystina Germar, 1839, S. lanius Stl, 1854, and Callidea spilogastra Walker, 1867. Scutellera perplexa is recorded for the first time from Nepal, Vietnam and Laos. Scutellera holosericea Dohrn, 1863, is removed from Scutellera and transferred to Procilia Stl, [1865], resulting in a formal combination Procilia holosericea (Dohrn, 1863), comb. nov.; the species is potentially a junior subjective synonym of the Afrotropical P. morgani (White, 1839); its type locality, Java, is considered as erroneous.

Zhengica cornuta gen. nov., sp. nov., a new genus and species of Pentatomidae from East Asia (Hemiptera: Heteroptera).

Zhengica cornuta gen. nov., sp. nov. (Hemiptera: Heteroptera: Pentatomidae: Pentatominae: Pentatomini) from Shanxi, China, is described, illustrated, and its phylogenetic affinities are discussed. The genus is recognized as a generalized member of the clade of Pentatoma Olivier, 1789 and related genera, its phylogenetically closest relative is probably Cervicoris Hsiao Cheng, 1977.

Two mitochondrial genomes of Taiwanese rhinoceros beetles, Oryctes rhinoceros and Eophileurus chinensis (Coleoptera: Scarabaeidae).

Two mitochondrial genomes of the dynastine beetles, Oryctes rhinoceros (Linnaeus, 1758) and Eophileurus chinensis (Faldermann, 1835), were assembled via high-throughput sequencing (HTS). Each of the mitogenomes has 37 genes, showing standard gene order and annotation as the other insects, except for the transfer genes, presenting tQ-tI-tM order. To examine their phylogenetic positions, 118 public mitogenomes of Scarabaeidae were used to infer a ML tree. Overall, our scarabaeid phylogeny reveals clear relationships with high nodal supports, and the two rhinoceros beetles are both grouped with the subfamily Dynastinae. The feeding habit of the two clades seems to represent coprophagous and phytophagous types. However, polyphyletic relationships were observed in the subfamily Melolothinae and in the tribes of Onthophagini and Oniticellini. Further systematic revision is needed.

Redefinition of two little known mirine plant bug genera Babacoris and Paramiridius (Hemiptera: Heteroptera: Miridae: Mirinae).

A long neglected plant bug genus Babacoris Miyamoto, 1994 (Mirinae: Mirini), proposed for a single Taiwanese species, B. striatus Miyamoto, is redefined and redescribed as a valid genus, based on recent rediscovery of the type species from Hengchun, Taiwan; the neotype is designated for B. striatus. The genus is now presumed to be closely related to Paramiridius Miyamoto Yasunaga which also has been poorly known since the original description. A species recently described from Laos, Babacoris laomontanus (Oh, Yasunaga Lee) comb. n., is transferred from Paramiridius. An updated checklist of the tribe Mirini in Taiwan is incorporated, with proposition of a new combination, Adelphocorisella minutum (Poppius) comb. n., having been left in Megacoelum Fieber as incertae sedis.

The genus Alphocoris in the Indomalaya (Hemiptera: Heteroptera: Scutelleridae).

The Indomalayan species of the genus Alphocoris Germar, 1839 (Hemiptera: Heteroptera: Scutelleridae: Odontotarsinae: Odontotarsini) are revised. Three species, A. caudatus Rédei, Tsai Jindra, sp. nov. (India: Maharashtra), A. naso Rédei Tsai, sp. nov. (India: Goa), and A. asper Rédei, Tsai Jindra, sp. nov. (India: Goa and Maharashtra), are recognized. The type material of A. lixoides Germar, 1839 (type locality: Senegal) is documented; previous records of this species from India and Pakistan are considered as based on misidentifications, the species is restricted to the Afrotropical Region.

A revision of Graptophara (Hemiptera: Heteroptera: Scutelleridae).

The Indomalayan genus Graptophara Stål, 1865 (Hemiptera: Heteroptera: Scutelleridae: Scutellerinae) is redescribed and revised, and its systematic position is discussed. Two species, G. reynaudii (Guérin-Méneville, 1834) (the type species, distributed in the Sundaic Region) and G. anomala sp. nov. (southern India) are recognized, redescribed or described, and illustrated. A lectotype is designated for Callidea pulchra Westwood, 1837, now a junior synonym of G. reynaudii.

Taxonomic review of the plant bug subfamily Isometopinae for Taiwan and Japanese Southwest Islands, with descriptions of new taxa (Hemiptera: Heteroptera: Miridae: Isometopinae).

The fauna of plant bug subfamily Isometopinae in Taiwan and Japanese Southwest (Nansei) Islands is reviewed. Twenty-five species are recognized, including two new species of Myiomma Puton, M. austroccidens sp. nov. and M. kentingense sp. nov., which are herein diagnosed and described. In addition, Isometopus yehi Lin, 2004 is synonymized with I. bipunctatus Lin; Isometopidea yangi Lin is transferred to Kohnometopus; and a substitute name, Alcecoris linyangorum, is proposed for A. formosanus (Lin Yang) (= a junior secondary homonym of Alcecoris formosanus Lin). An annotated checklist, with updated distributional record and biological information, is provided for all treated taxa. A new tribe Sophianini is proposed for two genera, Alcecoris and Sophianus, characterized principally by the conspicuously modified antennal structures. An additional new species, Alcecoris cochlearatus sp. nov., found during examination of related Oriental specimens, is described from the Malay Peninsula.

Lectotype designations and taxonomic corrections on Neotropical Scutelleridae described by G. Breddin (Hemiptera: Heteroptera).

Lectotypes are designated and documented for the following species and infrasubspecific taxa: Dystus villosus Breddin, 1904; Lobothyreus breviceps Breddin, 1914; Pachycoris torridus (Scopoli, 1772) var. laetissimus Breddin, 1906 (originally proposed as var. laetissima); P. torridus var. moestissimus Breddin, 1906 (originally proposed as var. moestissima); Polytes speculiger Breddin, 1914. The identities of the taxa in concern are clarified. The following new junior subjective synonyms are proposed: Pachycoris torridus (Scopoli, 1772) = P. torridus var. laetissimus Breddin, 1906, syn. nov., = P. torridus var. moestissimus Breddin, 1906, syn. nov.

Maternal care in Acanthosomatinae (Insecta: Heteroptera: Acanthosomatidae)--correlated evolution with morphological change.

BACKGROUND: Maternal care (egg-nymph guarding behavior) has been recorded in some genera of Acanthosomatidae. However, the origin of the maternal care in the family has remained unclear due to the lack of phylogenetic hypotheses. Another reproductive mode is found in non-caring species whose females smear their eggs before leaving them. They possess pairs of complex organs on the abdominal venter called Pendergrast's organ (PO) and spread the secretion of this organ onto each egg with their hind legs, which is supposed to provide a protective function against enemies. Some authors claim that the absence of PO may be associated with the presence of maternal care. No study, however, has tested this hypothesis of a correlated evolution between the two traits. RESULTS: We reconstructed the molecular phylogeny of the subfamily Acanthosomatinae using five genetic markers sequenced from 44 species and one subspecies with and without maternal care. Eight additional species from the other two acanthosomatid subfamilies were included as outgroups. Our results indicated that maternal care has evolved independently at least three times within Acanthosomatinae and once in the outgroup species. Statistical tests for correlated evolution showed that the presence of maternal care is significantly correlated with the secondary loss or reduction of PO. Ancestral state reconstruction for the node of Acanthosoma denticaudum (a non-caring species in which egg smearing with developed POs occurs) and A. firmatum (a caring species with reduced POs) suggested egg smearing was still present in their most recent common ancestor and that maternal care in A. firmatum has evolved relatively recently. CONCLUSIONS: We showed that maternal care is an apomorphic trait that has arisen multiple times from the presence of PO within the subfamily Acanthosomatinae. The acquisition of maternal care is correlated with the reduction or loss of PO, which suggests an evolutionary trade-off between the two traits resulting from physiological costs. This prediction also implies that presence of maternal care can be highly expected for those groups lacking behavioral data, which invariably also lack the organ. No secondary loss of maternal care was detected in the present tree. We suggest that the loss of maternal care may be suppressed due to the vulnerability of the PO-free condition, which thus maintains maternal care.

The identity of Acanthosoma vicinum, with proposal of a new genus and species level synonymy (Hemiptera: Heteroptera: Acanthosomatidae).

The identity of Acanthosoma vicinum Uhler, 1861 (type species of the monotypic genus Grossaria Kumar, 1974) is clarified based on reexamination of the lectotype. The following new combination and new subjective synonymies are proposed: Elasmucha Stål, 1864 = Grossaria Kumar, 1974, syn. nov.; Elasmucha vicina (Uhler, 1861), comb. nov. (transferred from Acanthosoma) = Elasmucha dorsalis (Jakovlev, 1876), syn. nov. Reversion to the senior name E. vicina is considered to be undesirable, therefore, in order to preserve stability, no nomenclatural changes are proposed in this paper, but an application has simultaneously been submitted to the International Commission on Zoological Nomenclature to give the specific name dorsalis precedence over the specific name vicinum.

Redefinition of Acanthosoma and taxonomic corrections to its included species (Hemiptera: Heteroptera: Acanthosomatidae).

The genus level diagnostic characters of Acanthosoma Curtis, 1824, Anaxandra Stål, 1876, and Sastragala Amyot & Serville, 1843 (Hemiptera: Heteroptera: Acanthosomatidae) are discussed. The synonymy of Acanthosoma and Anaxandra proposed by Kumar (1974) is supported. Acanthosoma and Sastragala are redefined and recognized as valid genera. The identities of various species of Acanthosoma are clarified based on their type materials, their diagnostic characters are discussed and several misidentifications in previous works are corrected. The following new or reinstated combinations and new subjective synonymies are proposed: Acanthosoma haemorrhoidale haemorrhoidale (Linnaeus, 1758) = Acanthosoma proximum Dallas, 1851, syn. nov. = Acanthosoma difficile Dallas, 1851, syn. nov. = Acanthosoma dubium Dallas, 1851, syn. nov.; Acanthosoma firmatum (Walker, 1868), comb. nov. (transferred from Sastragala) = Acanthosoma giganteum Matsumura, 1913, syn. nov.; Acanthosoma murreanum (Distant, 1900), comb. nov. (transferred from Sastragala) = Acanthosoma acutangulata Liu, 1979, syn. nov. = Sastragala neoelongata Ahmad & Moizuddin, 1990, syn. nov.; Acanthosoma rufispinum (Distant, 1887), comb. nov. (transferred from Sastragala) = Sastragala minuta Ahmad & Moizuddin, 1990, syn. nov.; Acanthosoma tauriforme (Distant, 1887) = Anaxandra longispina Liu, 1987, syn. nov.; Acanthosoma hampsoni (Distant, 1900), comb. nov. (transferred from Sastragala); Acanthosoma labiduroides Jakovlev, 1880 = Acanthosoma coralliferum Horváth, 1889, syn. nov. = Acanthosoma zanthoxylum Hsiao & Liu, 1977, syn. nov.; Acanthosoma forfex Dallas, 1851 = Acanthosoma distinctum Dallas, 1851, syn. nov.; Acanthosoma rufescens Dallas, 1851, comb. restit. = Acanthosoma elongatum Dallas, 1851, syn. nov. = Anaxandra hamata Reuter, 1881, syn. nov.; Acanthosoma nigricorne Walker, 1868 = Acanthosoma nigrospina Hsiao & Liu, 1977, syn. nov.; Acanthosoma alaticorne Walker, 1868, comb. restit. = Anaxandra laticollis Hsiao & Liu, 1977, syn. nov. New combinations resulting from synonymization of Anaxandra with Acanthosoma: Acanthosoma sichuanense (Liu, 1980), comb. nov., A. montanum (Liu, 1987), comb. nov. Species transferred to Sastragala Amyot & Serville, 1843: Sastragala nigrolineata (Stål, 1876), comb. nov.; S. sigillata (Stål, 1876), comb. nov.; S. versicolor Distant, 1910, comb. restit.; S. yunnana (Hsiao & Liu, 1977), comb. nov.; to Elasmostethus Fieber, 1860: Elasmostethus singhalensis (Distant, 1902), comb. nov. Lectotypes are designated for the following species: Acanthosoma: A. difficile, A. distinctum, A. dubium, A. elongatum, A. forfex, A. proximum; Anaxandra: A. hamata, A. nigrocornuta Reuter, 1881, A. tauriformis; Sastragala: S. murreeana, S. rufispina. A neotype is designated for Anaxandra montana. New country records: Acanthosoma alaticorne and A. rufispinum: Nepal; A. forfex: Burma (Myanmar); A. labiduroides: Burma (Myanmar) and Vietnam; A. montanum: Laos and Vietnam; A. murreeanum: India, Thailand; A. nigricorne: Nepal, Burma (Myanmar) and Vietnam; A. rufescens: Pakistan, Nepal, Taiwan, Burma (Myanmar), Thailand, Laos and Vietnam.

The soapberry bug, Jadera haematoloma (Insecta, Hemiptera, Rhopalidae): First Asian record, with a review of bionomics.

The soapberry bug, Jadera haematoloma (Herrich-Schäffer, 1847) (Insecta: Hemiptera: Heteroptera: Rhopalidae: Serinethinae), a species native in tropical and subtropical regions of the New World and accidentally introduced to Hawaii, is reported for the first time from Asia (Taiwan). This record represents the first occurrence of the species in Asia. Stable populations composed of hundreds of specimens were found in seven localities of Kaohsiung City and one locality in Tainan City, and a single specimen was observed in Chiayi County. Aggregating adults and larvae fed in large numbers on the sapindacean plants Cardiospermum halicacabum L. and Koelreuteria elegans (Seem.) A. C. Smith ssp. formosana (Hayata) F. G. Meyer. Diagnostic characters of adults and larvae of Jadera haematoloma are discussed. A review of its bionomics and a bibliography are provided. Initial observations on the populations in southern Taiwan are presented. The species is potentially invasive, and further extension of its range is anticipated in Southeast Asia.