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New genera and species of Ptiliini (Coleoptera: Ptiliidae) with a tribal revision and key to genera
Abstract
The list of genera in the polyphyletic tribe Ptiliini, used as a dumping ground for difficult genera of Ptiliidae by several generations of Coleopterists, is brought up to date with recent additions and omissions, and figures and a key to the genera are provided to aid their determination. 29 new species and 3 new genera are added. It was hoped to be able to determine some generic groups within the tribe but that work must depend on further cladistical and DNA research in the future. The new genera are Cingulum gen.n. , Iloptila gen.n. and Numa gen.n. and the new species are Actidium cooteri sp.n. , A. minimum sp.n. , A. nigrum sp.n. , Cingulum orsippus gen.n. sp.n. , Dipentium bicolor sp.n. , D. punctissimum sp.n. , D. spinosum sp.n. , Gomyella intricata sp.n. , G. nicoya sp.n., G. profunda sp.n. , G. tripla sp.n. , Greensladella cicra sp.n. , G. similata sp.n. , Iloptila modica gen.n. sp.n. , Micridium circulatum sp.n. , M. dembickyi sp.n. , M. hirsutum sp.n. , M. juara sp.n. , M. novum sp.n. , Millidium karnatakense sp.n ., Numa carmen gen.n. sp.n. , Ptiliodes kanchiporam sp.n. , Ptiliola bennetti sp.n. , P. nigra sp.n. , P. peruviensis sp.n. , Ptilium lisae sp.n. , P. longum sp.n. , P. pallidulum sp.n. and P. piceum sp.n.
Ptiliidae (featherwing beetles) is a group of minute staphylinoid beetles with a scarce fossil record. Here a second member of the Mesozoic genus Kekveus Yamamoto et al. is reported from mid-Cretaceous Burmese amber, with detailed morphology obtained through confocal microscopy. Kekveus brevisulcatus Li, Yamamoto, Newton & Cai sp. nov. shares with K. jason Yamamoto et al. the unpaired medial pronotal fovea and narrowly separated transverse metacoxae, but can be separated from the latter based on its less elongate body, shorter pronotal foveae, and much weaker transverse depression on the head. Our phylogenetic analyses support the discheramocephalin affinity of Kekveus , although its relationship with other members of Discheramocephalini cannot be confidently resolved.
This paper examines the described species of Cissidium and adds 77 new ones based on a study of material in the Manchester Museum, UK, the Natural History Museum, London, UK and in the possession of Dr Yoshihiro Sawada, Japan. The new taxa are: Cissidium advena sp. nov., C. amazonicum sp. nov., C. amieuense sp. nov., C. angulatum sp. nov., C. angusi sp. nov., C. apterum sp. nov., C. arcuatum sp. nov., C. aristophanousi sp. nov., C. besucheti sp. nov., C. bifoveolatum sp. nov., C. bilineatum sp. nov., C. bomjesus sp. nov., C. brachypterum sp. nov., C. ceylonicum sp. nov., C. clareae sp. nov., C. cryptophagoides sp. nov., C. davaoense sp. nov., C. deanei sp. nov., C. delicatum sp. nov., C. dewhurst sp. nov., C. dividuum sp. nov., C. dybasi sp. nov., C. eciton sp. nov., C. ecuadoriense sp. nov., C. faustum sp. nov., C. flavum sp. nov., C. franzi sp. nov., C. fraternum sp. nov., C. glabratum sp. nov., C. greensladei sp. nov., C. houailou sp. nov., C. ibicarense sp. nov., C. impressum sp. nov., C. inexspectatum sp. nov., C. insulare sp. nov., C. kolombangaricum sp. nov., C. lamington sp. nov., C. lawrencei sp. nov., C. loebli sp. nov., C. logunovi sp. nov., C. losbanos sp. nov., C. lucidulum sp. nov., C. luctuosum sp. nov., C. mahleri sp. nov., C. mindanao sp. nov., C. misellum sp. nov., C. modestum sp. nov., C. montanum sp. nov., C. murphyi sp. nov., C. mussardi sp. nov., C. noumeae sp. nov., C. obscenum sp. nov., C. obscurum sp. nov., C. obsoletum sp. nov., C. opacum sp. nov., C. orami sp. nov., C. peruviense sp. nov., C. pilosellum sp. nov., C. pinense sp. nov., C. plaumanni sp. nov., C. puncticolle sp. nov., C. riparium sp. nov., C. robustum sp. nov., C. rogeri sp. nov., C. seditiosum sp. nov., C. semicalvum sp. nov., C. similatum sp. nov., C. steeli sp. nov., C. subangulatum sp. nov., C. subfaustum sp. nov., C. sueae sp. nov., C. trangse sp. nov., C. upiense sp. nov., C. visendum sp. nov., C. waginicum sp. nov., C. werneri sp. nov. and C. yoruba sp. nov. The division of the genus is discussed and five species groups are established. The majority of the new species are the first records of Cissidium for the 28 countries represented.
The higher-level systematics of featherwing beetles (Coleoptera: Ptiliidae), the smallest non-parasitic insects and abundant in the world’s forests, is poorly known. Based on larval morphology, the systematic relationships and internal classification are revised and analysed from a comparative, evolutionary and phylogenetic perspective. We introduce 37 morphological characters of all instars and of strong phylogenetic signal in a first-ever maximum parsimony phylogenetic family analysis, providing support for the erection of Nossidiinae, subfam. nov., keeping Cephaloplectinae within Ptiliidae and sinking Acrotrichini as a tribe into a revised subfamily Ptiliinae. Full homology of characters between Ptiliidae and its sister taxon Hydraenidae is demonstrated and miniaturisation effects on the larval exoskeleton reviewed. Keeping generalised mouthparts adapted to sapro-mycophagy through geological time made three evolutionarily independent invasion events of marine littorals possible in Ptiliidae. Keys to larval instars, subfamilies, tribes and species are presented for the first time, supplemented by model descriptions of multiple hierarchical levels (species, genus, tribe, subfamily, family), introducing many phylogenetically informative characters. The conservative larval morphology/chaetotaxy proves particularly useful for analysis on the family-group taxon level, as exemplified by caudal urogomphi, a key character in ptiliid systematics. The revised nomenclature and classification improves systematic predictability and better reflects natural relationships within Ptiliidae.
Erro gen. n. a new genus of Ptiliidae (Coleoptera) with the new species impiger sp. n. and angolensis sp. n. is described and figured based on material collected in Angola and Madagascar. Pending further work on the phylogeny of the Ptiliidae the new genus is placed in the subfamily Ptiliinae where it appears to occupy a place close to the tribe Discheramocephalini.
The six recognised species of the genus Micridium Motschulsky, 1869: M. vittatum (Motschulsky, 1845) M. halidaii (Matthews, 1868), M. angulicolle (Fairmaire, 1858), M. lineatum (Le Conte, 1863). M. rhodeanum (Casey, 1924) and M. groehni Polilov & Perkovsky, 2004 are reviewed and eleven new species: M. attenboroughi sp.n., M. boliviense sp.n., M. elegans sp. n., M. foveatum sp.n., M. inornatum sp.n., M. johnsoni sp.n., M. newtoni sp.n., M. oweni sp.n., M. proprium sp.n., M. quadridens sp.n. and M. thayerae sp.n. are added. The new species are the first records of the genus from South America and Madagascar. The morphological characteristics separating Micridium from other genera in the tribe Ptiliini are discussed and the genus synonymised with the closely related monotypic genus Micridina Johnson, 1969 with its single species M. hilli Johnson, 1969. The synonymy of Dilinium Casey, 1924 with Micridium is confirmed and the type species D. rhodeanum Casey, 1924 re-described. Micridium groehni, described from a single specimen in Baltic amber, does not appear to belong to Micridium as defined here. A key to the new species is provided.
The recently described and originally monotypic genus Discheramocephalus Johnson, 2007 from the Solomon Islands is revised. Six new species are described, illustrated and keyed: Discheramocephalus brucei sp. n. (Cameroon), D. elisabethae sp. n. (Cameroon), D. mikaeli sp. n. (Tanzania), D. stewarti sp. n. (Bolivia), D. jarmilae sp. n. (Bolivia), D. minutissimus sp. n. (Indonesia). Adults of D. minutissimus have a body length of about 400-426 μm, which is at the lower limit among non-egg-parasitoid insects. Evidence is provided that an egg size large enough to produce a viable larva is the main factor limiting miniaturisation of female insects. Females and males of egg-parasitoids are able to overcome the 400 μm threshold and reach limits of 180 μm and 130 μm, respectively. Brain size is likely the second most important factor limiting miniaturisation in insects.
The smallest beetles and the smallest non-parasitic insects belong to the staphylinoid family Ptiliidae. Their adult body length can be as small as 0.325 mm and is generally smaller than 1 mm. Here we address the phylogenetic relationships within the family using formal analyses of adult morphological characters and molecular data, and also a combination of both for the first time. Strongly supported clades are Ptiliidae + Hydraenidae, Ptiliidae, Ptiliidae excl. Nossidium, Motschulskium and Sindosium, Nanosellini, and a clade comprising Acrotrichis, Smicrus, Nephanes and Baeocrara. A group comprising Actidium, Oligella and Micridium + Ptilium is also likely monophy-letic. Limulodes + Cephaloplectus, strongly supported as a clade (Cephaloplectus included only in morphological analysis), are deeply nested within Ptiliidae in the morphology-only non-weighted and in the molecular analyses, but placed as sister to the remaining Ptiliidae (excl. Nossidium, Motschulskium and Sindosium) after successive reweighting. We propose that Ptiliidae should be taxonomically divided into two subfamilies: the most recently established Nossidiinae and Ptiliinae, the latter currently composed of seven tribes: Acrotrichini, Cephaloplectini, Discheramocephalini, Nanosellini, Ptenidiini, Ptiliini and Ptinellini, although the monophyly and taxonomic status is still uncertain in some cases. Important morphological innovations evolved in the stem group of Hydraenidae and Ptiliidae, including partly internalized mouthparts suitable for saprophagy and sporophagy, a labral-mandibular locking device, a specific elytral locking mechanism with elongated alacristae, wings with fringes of hairs, and a very unusual spermathecal pump. A complex feature of Ptiliidae linked with miniaturization is the transformation of the alae into "feather wings", with a highly efficient flight mechanism unique in beetles. Nano-sellini include the smallest ptiliid species and display features correlated with extremely small body size, such as simplification of the endoskeleton of the head and thorax, far-reaching reduction of the wing venation, and a reduced number of antennomeres.
This three-volume series represents a comprehensive treatment of the beetles of Australia, a relatively under-studied fauna that includes many unusual and unique lineages found nowhere else on Earth. Volume 1 contains keys to all 117 beetle families found in Australia, and includes over 1100 illustrations of adults, larvae and anatomical structures. This volume is based in part on Lawrence & Britton’s out-of-print Australian Beetles, but is fully updated and expanded. The biology and morphology for all major beetle lineages is described and illustrated, along with anatomical terms which clarify the characters and terminology used in the keys; few other resources for beetle identification include such a detailed morphological background. A chapter on the fossil record is also included, and family sections provide full descriptions of adults and larvae, including the world distribution of each family. The revised identification keys (currently recognised as one of the most valuable keys worldwide) will aid quarantine agents, biologists and students in identifying members of the most species-rich order of animals.
Ptiliola flammifera (Młynarski) is recognised as a species distinct from P. kunzei (Heer). Apart from differences in the structure of the aedeagus, both species can be separated by the pronotal surface and the apical fringe of the elytra. The male of P. flammifera carries a distinct tuft of hairs on the metaventrite, absent in P. kunzei. Currently, P. flammifera is only known from Poland and the Netherlands. A key for the identification of all three Palaearctic Ptiliola species is presented. A lectotype is designated for Trichopteryx kunzei Heer, 1841.
2,000 Ptiliidae collected in the North and South Islands of New Zealand in 1983/1984 by Peter Hammond of the Natural History Museum, London, are determined to 34 species, four of which are new to the country. Because there are very few previous records, most from the Auckland district of North Island, the Hammond collection provides much new distributional data. The three new species: Nellosana insperatus sp. n., Notoptenidium flavum sp. n., and Notoptenidium johnsoni sp. n., are described and figured; the genus Ptiliodes is moved from Acrotrichinae to Ptiliinae, and Ptenidium formicetorum Kraatz recorded as a new introduction. Information is provided to aid separation of the new species from those previously recorded.
Ptiliola flammifera (Młynarski) is recognised as a species distinct from P. kunzei (Heer). Apart from differences in the structure of the aedeagus, both species can be separated by the pronotal surface and the apical fringe of the elytra. The male of P. flammifera carries a distinct tuft of hairs on the metaventrite, absent in P. kunzei. Currently, P. flammifera is only known from Poland and the Netherlands. A key for the identification of all three Palaearctic Ptiliola species is pre-sented. A lectotype is designated for Trichopteryx kunzei Heer, 1841.
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