Christopher H. C. Lyal

Family-group names in Coleoptera (Insecta)

Abstract We synthesize data on all known extant and fossil Coleoptera family-group names for the first time. A catalogue of 4887 family-group names (124 fossil, 4763 extant) based on 4707 distinct genera in Coleoptera is given. A total of 4492 names are available, 183 of which are permanently invalid because they are based on a preoccupied or a suppressed type genus. Names are listed in a classification framework. We recognize as valid 24 superfamilies, 211 families, 541 subfamilies, 1663 tribes and 740 subtribes. For each name, the original spelling, author, year of publication, page number, correct stem and type genus are included. The original spelling and availability of each name were checked from primary literature. A list of necessary changes due to Priority and Homonymy problems, and actions taken, is given. Current usage of names was conserved, whenever possible, to promote stability of the classification. New synonymies (family-group names followed by genus-group names): Agronomina Gistel, 1848 syn. nov. of Amarina Zimmermann, 1832 (Carabidae), Hylepnigalioini Gistel, 1856 syn. nov. of Melandryini Leach, 1815 (Melandryidae), Polycystophoridae Gistel, 1856 syn. nov. of Malachiinae Fleming, 1821 (Melyridae), Sclerasteinae Gistel, 1856 syn. nov. of Ptilininae Shuckard, 1839 (Ptinidae), Phloeonomini Ádám, 2001 syn. nov. of Omaliini MacLeay, 1825 (Staphylinidae), Sepedophilini Ádám, 2001 syn. nov. of Tachyporini MacLeay, 1825 (Staphylinidae), Phibalini Gistel, 1856 syn. nov. of Cteniopodini Solier, 1835 (Tenebrionidae); Agronoma Gistel 1848 (type species Carabus familiaris Duftschmid, 1812, designated herein) syn. nov. of Amara Bonelli, 1810 (Carabidae), Hylepnigalio Gistel, 1856 (type species Chrysomela caraboides Linnaeus, 1760, by monotypy) syn. nov. of Melandrya Fabricius, 1801 (Melandryidae), Polycystophorus Gistel, 1856 (type species Cantharis aeneus Linnaeus, 1758, designated herein) syn. nov. of Malachius Fabricius, 1775 (Melyridae), Sclerastes Gistel, 1856 (type species Ptilinus costatus Gyllenhal, 1827, designated herein) syn. nov. of Ptilinus Geoffroy, 1762 (Ptinidae), Paniscus Gistel, 1848 (type species Scarabaeus fasciatus Linnaeus, 1758, designated herein) syn. nov. of Trichius Fabricius, 1775 (Scarabaeidae), Phibalus Gistel, 1856 (type species Chrysomela pubescens Linnaeus, 1758, by monotypy) syn. nov. of Omophlus Dejean, 1834 (Tenebrionidae). The following new replacement name is proposed: Gompeliina Bouchard, 2011 nom. nov. for Olotelina Báguena Corella, 1948 (Aderidae). Reversal of Precedence (Article 23.9) is used to conserve usage of the following names (family-group names followed by genus-group names): Perigonini Horn, 1881 nom. protectum over Trechicini Bates, 1873 nom. oblitum (Carabidae), Anisodactylina Lacordaire, 1854 nom. protectum over Eurytrichina LeConte, 1848 nom. oblitum (Carabidae), Smicronychini Seidlitz, 1891 nom. protectum over Desmorini LeConte, 1876 nom. oblitum (Curculionidae), Bagoinae Thomson, 1859 nom. protectum over Lyprinae Gistel 1848 nom. oblitum (Curculionidae), Aterpina Lacordaire, 1863 nom. protectum over Heliomenina Gistel, 1848 nom. oblitum (Curculionidae), Naupactini Gistel, 1848 nom. protectum over Iphiini Schönherr, 1823 nom. oblitum (Curculionidae), Cleonini Schönherr, 1826 nom. protectum over Geomorini Schönherr, 1823 nom. oblitum (Curculionidae), Magdalidini Pascoe, 1870 nom. protectum over Scardamyctini Gistel, 1848 nom. oblitum (Curculionidae), Agrypninae/-ini Candèze, 1857 nom. protecta over Adelocerinae/-ini Gistel, 1848 nom. oblita and Pangaurinae/-ini Gistel, 1856 nom. oblita (Elateridae), Prosternini Gistel, 1856 nom. protectum over Diacanthini Gistel, 1848 nom. oblitum (Elateridae), Calopodinae Costa, 1852 nom. protectum over Sparedrinae Gistel, 1848 nom. oblitum (Oedemeridae), Adesmiini Lacordaire, 1859 nom. protectum over Macropodini Agassiz, 1846 nom. oblitum (Tenebrionidae), Bolitophagini Kirby, 1837 nom. protectum over Eledonini Billberg, 1820 nom. oblitum (Tenebrionidae), Throscidae Laporte, 1840 nom. protectum over Stereolidae Rafinesque, 1815 nom. oblitum (Throscidae) and Lophocaterini Crowson, 1964 over Lycoptini Casey, 1890 nom. oblitum (Trogossitidae); Monotoma Herbst, 1799 nom. protectum over Monotoma Panzer, 1792 nom. oblitum (Monotomidae); Pediacus Shuckard, 1839 nom. protectum over Biophloeus Dejean, 1835 nom. oblitum (Cucujidae), Pachypus Dejean, 1821 nom. protectum over Pachypus Billberg, 1820 nom. oblitum (Scarabaeidae), Sparrmannia Laporte, 1840 nom. protectum over Leocaeta Dejean, 1833 nom. oblitum and Cephalotrichia Hope, 1837 nom. oblitum (Scarabaeidae).

Phylogeny and classification of the Psocodea, with particular reference to the lice (Psocodea: Phthiraptera)

ABSTRACT. Apomorphies that have been proposed for the Psocodea, Psocoptera, Phthiraptera and superfamilial groups within the Phthiraptera are enumerated and evaluated. The Psocodea and Phthiraptera are considered to be holophyletic, but the sister‐group of the Phthiraptera lies within the Psocoptera. Within the Phthiraptera the Anoplura and Rhyncophthirina form a holophyletic group whose sister‐group is the Ischnocera, and the Amblycera is the sister‐group of this assemblage. The common ancestor of the Phthiraptera is suggested to have been parasitic, and all lice are believed to have evolved under environmental constraints similar to those operating today. On the evidence provided by host relationships the origin of the lice is dated as the Cretaceous, but the host of the ancestor of the order is not identified. The lice of marsupials in South America and Australia are not considered to comprise a holophyletic group.

Elytro-tergal stridulation in weevils (Insecta: Coleoptera: Curculionoidea)

Many Curculionidae, Scolytidae and Platypodidae stridulate by rubbing a plectrum on tergite VII against a subapical file on the elytral underside. The file on the right elytron may be smaller than that on the left, or absent; on either elytron it usually comprises a series of parallel ridges. The plectrum generally consists of one or more pair of setiferous tubercles, although other structures are sometimes employed. This stridulatory system (‘type 1’) is found in members of 14 curculionid subfamilies and in Scolytidae. Platypodidae have a similar and probably homologous organ, although the morphology of the file differs somewhat from that found elsewhere and the plectrum is a simple flange. Within Curculionidae three other elytro-tergal stridulatory systems have been found. Firstly, in some Cossoninae and Molytinae, the file is a row of tubercles on tergite VII and the plectrum is a ridge on the elytron (‘type 2’); this is newly reported here. Secondly, females of Ithyporini, some Camptorhinini and some ...

DNA profiling of host–herbivore interactions in tropical forests

1. The diversity of insects in tropical forests remains poorly known, in particular regarding the critical feeding associations of herbivores, which are thought to drive species richness in these ecosystems.

Impacts of logging on density-dependent predation of dipterocarp seeds in a South East Asian rainforest

Much of the forest remaining in South East Asia has been selectively logged. The processes promoting species coexistence may be the key to the recovery and maintenance of diversity in these forests. One such process is the Janzen–Connell mechanism, where specialized natural enemies such as seed predators maintain diversity by inhibiting regeneration near conspecifics. In Neotropical forests, anthropogenic disturbance can disrupt the Janzen–Connell mechanism, but similar data are unavailable for South East Asia. We investigated the effects of conspecific density (two spatial scales) and distance from fruiting trees on seed and seedling survival of the canopy tree Parashorea malaanonan in unlogged and logged forests in Sabah, Malaysia. The production of mature seeds was higher in unlogged forest, perhaps because high adult densities facilitate pollination or satiate pre-dispersal predators. In both forest types, post-dispersal survival was reduced by small-scale (1 m2) conspecific density, but not by proximity to the nearest fruiting tree. Large-scale conspecific density (seeds per fruiting tree) reduced predation, probably by satiating predators. Higher seed production in unlogged forest, in combination with slightly higher survival, meant that recruitment was almost entirely limited to unlogged forest. Thus, while logging might not affect the Janzen–Connell mechanism at this site, it may influence the recruitment of particular species.

Seed-feeding beetles of the weevil tribe Mecysolobini (Insecta: Coleoptera: Curculionidae) developing in seeds of trees in the Dipterocarpaceae

Species comprising two monophyletic groups of Alcidodes sensu lato (crassus group and dipterocarpi group) feed on fruits of the ecologically and economically important tropical timber family Dipterocarpaceae. Fifteen Alcidodes species are newly described from several thousand specimens of seed-feeding beetles reared from south-east Asian dipterocarp hosts (Dipterocarpoidae) and four other species are revised. Morphological keys are provided for identifying these 19 taxa. Weevil associations are provided for 70 species of Dipterocarpaceae in five genera (16 Dipterocarpus spp., four Dryobalanops spp., six Hopea spp., 39 Shorea spp. and five Vatica spp.). These records relate primarily to specimens reared from seeds in Borneo and in Peninsular Malaysia, but all known previous host records are also included (from India, Andaman Islands, Thailand, Laos, Vietnam and the Philippines) and assessed. These validated and documented host associations refute general assertions that (1) seed-eating beetles are host-specific in the tropics and (2) seed-eating beetles found on dipterocarp hosts are broad generalists across species in several genera of mast-fruiting Dipterocarpaceae. Most Alcidodes species exploit seeds from several dipterocarp species. Some Alcidodes species occur across broad geographic regions with several host species from a dipterocarp genus, other species have been found only on Bornean Hopea or Dipterocarpus species. Others, such as the two previously undescribed species feeding on Vatica species, may be restricted to separate sections of the host genus. Within an extensively sampled and diverse interspecific mast-fruiting dipterocarp community, no Alcidodes species was observed to feed on sympatric species across genera within the Dipterocarpaceae. Additional host records and specimens of Alcidodes spp. throughout the range of the Dipterocarpaceae in south and southeast Asia are required to further define these clades and the tribe Mecysolobini in general as well as to resolve the complex host associations and distributions documented.

Phylogeny of the Oxycoryninae sensu lato (Coleoptera : Belidae) and evolution of host-plant associations

Phylogenetic relationships among the genera of the subfamily Oxycoryninae and other belids (Curculionoidea) were reconstructed by cladistic analysis using 21 terminals and 98 characters: 62 from imaginal morphology, 33 from larval morphology and three biological characters relating to host plants and larval feeding habits. Terminal taxa represent all extant genera of Oxycoryninae, two genera of each of the three tribes of Belinae plus two outgroup taxa used to root the tree. New information on the larvae and biology of the metrioxenines is used in phylogenetic reconstruction. In accord with the single optimal cladogram obtained, a revised classification of the Oxycoryninae is proposed. The subfamily is classified into three tribes (Oxycorynini, Metrioxenini and Aglycyderini), with the tribe Oxycorynini further classified into three subtribes (Oxycraspedina Marvaldi & Oberprieler, subtr. nov., Oxycorynina and Allocorynina) and the tribe Metrioxenini into two subtribes (Metrioxenina and Afrocorynina ( = Hispodini, syn. nov.)). Larval and adult unambiguous synapomorphies defining each clade are identified. Tracing the evolution of biological traits from the phylogenetic estimate indicates that drastic shifts to phylogenetically distant host plants occurred from the ancestral belid association with conifers. Structural, chemical and/or ecological similarities of the plant organs consumed apparently had a major influence in the colonisation of different plant taxa by this group of weevils.

Morphology and systematic significance of sclerolepidia in the weevils (Coleoptera: Curculionoidea)

Abstract Sclerolepidia are minute specialized structures found only alongthe metepisternal suture in the weevil subfamilies Scolytinae, Cossoninae, Baridinae and some tribes in the Cryptorhynchinae, Conoderinae, Molytinae, Curculioninae and Cyclominae of the Curculionidae. Their presence has been used to establish relationships at subfamily level in the Curculionidae, although some of these are questionable, because of misperceptions of the distribution of the structure across taxa. Within the Curculionidae sclerolepidia are probably homologous, but provide limited information aiding establishment of sister‐group relationships between subfamilies, and multiple loss is likely. Analogous structures occur in the Erirhinidae and the Eucnemidae, but not elsewhere in the Coleoptera. Structural evidence shows sclerolepidia to be derived from scales. Their morphology allows division into ‘squamiform’, ‘digitate’, ‘peg‐like’ and ‘plumose’ types. Squamiform sclerolepidia are only slightly modified from the ancestral scale and are probably the plesiomorphic type, with modification probably occurring independently a number of times. Taxonomic characters and phylogenetic information are provided by distribution of different types within the weevils at species, genus and higher levels. Numerous systematic changes were identified as necessary during this project from the study of the sclerolepidia; 27 genera are transferred at tribal level and these changes are listed in an appendix. The functions of sclerolepidia probably differ across the Curculionidae, and may include sensing the relative positions of the metepisternum and metasternum during flight, and diffusion of secreted wax or other chemicals.

Co-evolution of trichodectid lice (Insecta: Phthiraptera) and their mammalian hosts

The cladogram of Trichodectidae produced by Lyal (1985) is interpreted, with reservations, as a phylogeny and is compared to such host phylogenies as are available. While the predominant pattern of louse relationships is broadly congruent with host relationships, indicating either phyletic tracking or a mixture of phyletic tracking and host-group-limited resource tracking by the lice, a substantial proportion (20·7%) of speciation events of the lice could only be explained by invoking secondary infestation of distantly related hosts by the lice. The ancestor of the Trichodectidae almost certainly parasitized a eutherian, but the identity of the eutherian group is unclear. The primary host association of the Neotrichodectinae was with an American rodent, from which resource tracking onto Mephitinae and then Mustelinae, Procyonidae, Bradypodidae and Melinae took place. The Trichodectinae had an ancestral association with the ancestor of the Feloidea, with subsequent secondary infestations of Mustelidae, Can...

DNA-based taxonomy of larval stages reveals huge unknown species diversity in neotropical seed weevils (genus Conotrachelus): relevance to evolutionary ecology

High diversity in tropical phytophagous insects may be linked to narrow host specificity and host shifts, but tests are complicated by incomplete taxonomy and difficulties in food source identification. Specimens of the highly diverse New World genus Conotrachelus (Coleoptera: Curculionoidea) were reared from >17,500 fruits (seeds) at six Central American rain forests. Interception traps were used for comparison with assemblages flying in the forest. Mitochondrial cox1 and the nuclear 28S genes were sequenced for 483 larval and adult specimens. A Yule-Coalescent technique was used to group cox1 sequences into putative species (17 from traps, 48 from rearing). Cox1 sequences of 24 species from museum collections provided matches for three species from traps and no match for the reared species. Inga (Fabaceae) was the predominant host among 15 other genera and 67% of the weevils were monophagous. A three gene tree (cox1, rrnL, 28S) recovered four well-supported clades feeding on Inga confirmed by phylogenetic community analyzes that showed phylogenetic conservation of host plant utilization. This suggests that host shifts are not directly involved in speciation, while the broad taxonomic host range and the evolutionary repeated shifts still contribute to the high species richness in Conotrachelus. The DNA-based approach combining species delimitation and phylogenetic analysis elucidated the evolutionary diversification of this lineage, despite insufficient taxonomic knowledge. Conotrachelus is an example of the diverse tropical groups that require DNA-based taxonomy to study their evolutionary ecology.