Complex inter‐island colonization and peripatric founder speciation promote diversification of flightless Pachyrhynchus weevils in the Taiwan–Luzon volcanic belt

Aim

We investigated the spatial and temporal patterns of diversification among colourful and flightless weevils, the Pachyrhynchus orbifer complex, to test the stepping‐stone hypothesis of colonization across the Taiwan–Luzon volcanic belt.

Location

Southeast Asia.

Methods

The phylogeny of the P. orbifer complex was reconstructed from a multi‐locus data set of mitochondrial and nuclear genes using maximum likelihood in RAxML and Bayesian inference in MRBAYES. Likelihood‐based tests in CONSEL were used to evaluate alternative tree topologies. Divergence times were estimated in beast based on a range of mutation rates. Ancestral range and biogeographical history were reconstructed using Bayesian binary MCMC (BBM) methods in RASP and in BioGeoBEARS. Demographic histories were inferred using the extended Bayesian skyline plot (EBSP). Species boundaries were tested using BPP.

Results

The phylogeny of the P. orbifer complex indicated strong support for seven reciprocally monophyletic lineages grouped by current island boundaries (Camiguin, Fuga, Dalupiri, Calayan, Babuyan, Orchid and Yaeyama Islands), except for a sister Green + Itbayat lineage. Complex and stochastic colonization of P. orbifer was inferred to have involved both northward and southward directions with short‐ and long‐distance dispersal events, which are strongly inconsistent with the strict stepping‐stone hypothesis. Divergence time estimates for all extant island lineages (<1 Myr of Middle Pleistocene) are much more recent than the geological ages (22.4–1.7 Myr) and subaerial existence (c. 3 Myr) of the islands. The statistically delimited seven cryptic species imply that the diversity of Pachyrhynchus from small peripheral islands continues to be largely under‐estimated.

Main conclusions

The non‐linear, more complex spatial and temporal settings of the archipelago and stochastic dispersal were probable key factors shaping the colonization history of the P. orbifer complex. Speciation of the P. orbifer complex may have occurred only between islands, indicating that peripatric speciation through the founders of stochastic dispersals was the major evolutionary driver.     

Ninety-eight new species of Trigonopterus weevils from Sundaland and the Lesser Sunda Islands

The genus Trigonopterus Fauvel, 1862 is highly diverse in Melanesia. Only one species, Trigonopterus amphoralis Marshall, 1925 was so far recorded West of Wallace’s Line (Eastern Sumatra). Based on focused field-work the fauna from Sundaland (Sumatra, Java, Bali, Palawan) and the Lesser Sunda Islands (Lombok, Sumbawa, Flores) is here revised. We redescribe Trigonopterus amphoralis Marshall and describe an additional 98 new species: Trigonopterus acuminatus sp. n., Trigonopterus aeneomicans sp. n., Trigonopterus alaspurwensis sp. n., Trigonopterus allopatricus sp. n., Trigonopterus allotopus sp. n., Trigonopterus angulicollis sp. n., Trigonopterus argopurensis sp. n., Trigonopterus arjunensis sp. n., Trigonopterus asper sp. n., Trigonopterus attenboroughi sp. n., Trigonopterus baliensis sp. n., Trigonopterus batukarensis sp. n., Trigonopterus bawangensis sp. n., Trigonopterus binodulus sp. n., Trigonopterus bornensis sp. n., Trigonopterus cahyoi sp. n., Trigonopterus costipennis sp. n., Trigonopterus cuprescens sp. n., Trigonopterus cupreus sp. n., Trigonopterus dacrycarpi sp. n., Trigonopterus delapan sp. n., Trigonopterus dentipes sp. n., Trigonopterus diengensis sp. n., Trigonopterus dimorphus sp. n., Trigonopterus disruptus sp. n., Trigonopterus dua sp. n., Trigonopterus duabelas sp. n., Trigonopterus echinatus sp. n., Trigonopterus empat sp. n., Trigonopterus enam sp. n., Trigonopterus fissitarsis sp. n., Trigonopterus florensis sp. n., Trigonopterus foveatus sp. n., Trigonopterus fulgidus sp. n., Trigonopterus gedensis sp. n., Trigonopterus halimunensis sp. n., Trigonopterus honjensis sp. n., Trigonopterus ijensis sp. n., Trigonopterus javensis sp. n., Trigonopterus kalimantanensis sp. n., Trigonopterus kintamanensis sp. n., Trigonopterus klatakanensis sp. n., Trigonopterus lampungensis sp. n., Trigonopterus latipes sp. n., Trigonopterus lima sp. n., Trigonopterus lombokensis sp. n., Trigonopterus merubetirensis sp. n., Trigonopterus mesehensis sp. n., Trigonopterus micans sp. n., Trigonopterus misellus sp. n., Trigonopterus palawanensis sp. n., Trigonopterus pangandaranensis sp. n., Trigonopterus paraflorensis sp. n., Trigonopterus pararugosus sp. n., Trigonopterus parasumbawensis sp. n., Trigonopterus pauxillus sp. n., Trigonopterus payungensis sp. n., Trigonopterus porcatus sp. n., Trigonopterus pseudoflorensis sp. n., Trigonopterus pseudosumbawensis sp. n., Trigonopterus punctatoseriatus sp. n., Trigonopterus ranakensis sp. n., Trigonopterus relictus sp. n., Trigonopterus rinjaniensis sp. n., Trigonopterus roensis sp. n., Trigonopterus rugosostriatus sp. n., Trigonopterus rugosus sp. n., Trigonopterus rutengensis sp. n., Trigonopterus saltator sp. n., Trigonopterus santubongensis sp. n., Trigonopterus sasak sp. n., Trigonopterus satu sp. n., Trigonopterus schulzi sp. n., Trigonopterus sebelas sp. n., Trigonopterus sembilan sp. n., Trigonopterus sepuluh sp. n., Trigonopterus seriatus sp. n., Trigonopterus serratifemur sp. n., Trigonopterus setifer sp. n., Trigonopterus silvestris sp. n., Trigonopterus singkawangensis sp. n., Trigonopterus singularis sp. n., Trigonopterus sinuatus sp. n., Trigonopterus squalidus sp. n., Trigonopterus sumatrensis sp. n., Trigonopterus sumbawensis sp. n., Trigonopterus sundaicus sp. n., Trigonopterus suturalis sp. n., Trigonopterus syarbis sp. n., Trigonopterus telagensis sp. n., Trigonopterus tepalensis sp. n., Trigonopterus tiga sp. n., Trigonopterus trigonopterus sp. n., Trigonopterus tujuh sp. n., Trigonopterus ujungkulonensis sp. n., Trigonopterus variolosus sp. n., Trigonopterus vulcanicus sp. n., Trigonopterus wallacei sp. n.. All new species are authored by the taxonomist-in-charge, Alexander Riedel. Most species belong to the litter fauna of primary wet evergreen forests. This habitat has become highly fragmented in the study area and many of its remnants harbor endemic species. Conservation measures should be intensified, especially in smaller and less famous sites to minimize the number of species threatened by extinction.     

Overwintering strategy of two weevils infesting three gorse species: When cold hardiness meets plant-insect interactions

The cold hardiness of two closely related weevil species, Exapion ulicis and E. lemovicinum was studied in relation to their life cycles. These two seed-eating weevils reproduce on Ulex plant species with different fruiting phenologies. E. ulicis lays eggs in spring and overwinters as an adult while E. lemovicinum lays eggs in autumn and overwinters as a larva. Adult weevils were collected in natural populations of Brittany (Western France) and characterized with morphological and molecular tools before experiments. We showed that both weevil species exhibited low supercooling points (SCPs) with mean seasonal values below −17 °C. Fresh mass, moisture content and sex were not correlated to supercooling ability. Weevils died upon freezing and the lower lethal temperatures (LLT) were within the range of SCP, indicating that both species are freezing intolerant. Comparison between species for SCP, LLT and survival to exposure at −8 °C in winter showed a higher cold resistance for E. ulicis than for E. lemovicinum. In addition, the seasonal evolution of cold hardiness differed depending on the species. These features suggest that response to cold of weevils is linked to their life cycles, and thus to the life history of their host plants.     

Contributions à la réorganisation systématique des Peritelini ouest-paléarctiques. IX. Révision du genre Meira Jacquelin du Val 1852 (Coleoptera : Curculionidae : Entiminae)

The genus Meira Jacquelin du Val 1852 is revised, four lectotypes and a neotype are designated, a new synonymy is proposed and four new species from France – one from Alpes-Maritimes (Meira alpina n. sp.), two from Var (M. echinoides n. sp. and M. teloniensis n. sp.), one from Vaucluse (M. germanni n. sp.) and one from Italy, from Abruzzo (M. osellai n. sp.) are described,. Synonyms, bibliographic and ecological data are provided. Whole-body photos of the types of the old species and of the four new species, drawings of systematically important characters, keys and distribution maps are also presented.     

Two species of adventive weevil (Coleoptera: Curculionidae) from Europe,hitherto unrecorded from New Zealand

Two curculionid weevils, Orthochaetes setiger (Beck, 1817) and Exomias pellucidus (Boheman, 1834) are recorded in New Zealand for the first time. The former has a wide distribution through the eastern South Island, while the latter has so far only been located in a single suburban garden in Dunedin. Both species are polyphagous and flightless. Although neither is expected to cause notable economic damage, their potential to invade native ecosystems makes them worthy of further investigation.     

New species of the subfamily Conoderinae (Coleoptera: Curculionidae) in Dominican amber

Eulechriops argyrosoman. sp. (Curculionidae: Conoderinae: Lechropini) and Geratozygops platysoman. sp., Geratozygops stenosoman. sp. and Geratozygops arsinotusn. sp. (Curculionidae: Conoderinae: Zygopini) are described from Dominican amber. The small size, nearly parallel sides of the pronotum and silvery sheen distinguish Eulechriops argyrosoma from extant members of the genus, which have not been recorded from Hispaniola. Size and rostral and pronotal characters separate the three Geratozygops species from the single species (Geratozygops atropos) previously described from Dominican amber.http://www.zoobank.org/urn:lsid:zoobank.org:pub:89BC7F43-6C1A-4A6A-839B-CE94C7F6F140     

Introduced invertebrates are important prey for a generalist predator

Concern over biological invasions has drawn increased attention to the impacts of introduced predators or competitors, but not to the importance of introduced prey. North American forests are rich in introduced invertebrates, including species that represent relatively novel taxonomic or trophic guilds and show biased distributions among forest types. We analysed the diets of red‐backed salamanders, Plethodon cinereus, from three upland and three lowland forests to determine whether introduced prey are important contributors to geographical or temporal variation in salamander food resources. We found several introduced species were volumetrically important salamander prey, and were responsible for resource differences between forest types and much of the seasonal fluctuation in food resources in both forest types. In lowland forests, rain had a stronger effect on salamander predation on non‐native earthworms than native taxa, creating more dynamic resource fluctuations in resource levels than was observed in upland forests where earthworms were absent. With one exception, predation on non‐native species was positively associated with predation on native species, suggesting non‐native prey have added to salamander resources rather than replaced salamander predation on native taxa. We hypothesize that the novel resource gradients created by non‐native prey introductions are contributing to patterns of geographical and temporal phenotypic variation among salamander populations.     

Phylogenetic reassessment of the Exophthalmus genus complex (Curculionidae: Entiminae: Eustylini,Geonemini)

The monophyly of the Neotropical entimine weevil genus Exophthalmus Schoenherr, 1823 (Curculionidae: Entiminae: Eustylini Lacordaire) is reassessed. Exophthalmus presently includes more than 80 species, approximately half of which are restricted to either the Caribbean archipelago or the continental Neotropics. The taxonomic composition and position of Exophthalmus have been subject to longstanding disagreements; in particular, authors have questioned the relationship of Exophthalmus to other Caribbean genera such as Diaprepes Schoenherr, 1834 (Eustylini) and Lachnopus Schoenherr, 1840 (Geonemini Gistel), as well as to the speciose Central and South American genera Compsus Schoenherr, 1823, Eustylus Schoenherr, 1842, and Exorides Pascoe, 1881 (all Eustylini), among others. The present study scrutinizes these traditional perspectives, based on a cladistic analysis of 143 adult morphological characters and 90 species, representing 30 genera and seven tribes of Neotropical entimine weevils. The character matrix yielded eight most‐parsimonious cladograms (length = 239 steps; consistency index = 66; retention index = 91), with mixed clade support that remains particularly wanting for some of the deeper in‐group divergences. The strict consensus supports the existence of a paraphyletic Geonemini ‘grade’ that includes Lachnopus and related Caribbean genera such as Apotomoderes Dejean, 1834, followed by a monophyletic Eustylini in‐group clade. Within the latter, a monophyletic South American Eustylini clade – including Compsus, Eustylus, Exorides, and related genera – is sister to a major clade that contains a ‘grade’ of heterogeneous and often misclassified Caribbean members of the Eustylini, Geonemini (Tetrabothynus Labram & Imhoff, 1852 and Tropirhinus Schoenherr, 1823), and Tanymecini Lacordaire (Pachnaeus Schoenherr, 1826), as well as two major clades: one with the majority of Central American Exophthalmus species, and the other with most Caribbean members of Exophthalmus. The Central American Exophthalmus clade is paraphyletic with respect to Chauliopleurus Champion, 1911 (Geonemini) and Rhinospathe Chevrolat, 1878 (Phyllobiini Schoenherr). The Caribbean clade, in turn, contains two subclades: i.e. (1) the Greater Antillean Exophthalmus s.s. clade, including the type species Exophthalmus quadrivittatus (Olivier, 1807); and (2) the primarily Lesser Antillean Diaprepes. The latter genus is therefore nested within Central American and Caribbean species of a highly paraphyletic Exophthalmus, yet may be rendered monophyletic if several Lesser Antillean Exophthalmus species are (re‐)assigned to Diaprepes. The results thus provide a suitable basis for a revision of all Exophthalmus species, and furthermore suggest that historical biographic factors, including colonization via temporary continental Neotropics‐to‐Caribbean land connections, were important in the evolution of major eustyline lineages. Based on these preliminary insights, the following taxonomic and nomenclatural adjustments are made. Compsoricus gen. nov. is erected to accommodate two Puerto Rican species erroneously assigned to Compsus: i.e. the herein designated type species Compsoricus maricao comb. nov. and Compsoricus luquillo comb. nov. Eustylus dentipes comb. nov. is transferred from Compsus. Diaprepes marginicollis Chevrolat, 1880 is reinstated from synonymy under Exophthalmus. Lastly, the following five transfers are proposed: (1) Chauliopleurus Champion, 1911, from Geonemini to Eustylini; (2) Tetrabothynus Labram & Imhoff, 1852, from Geonemini to Eustylini; (3) Tropirhinus Schoenherr, 1823, from Geonemini to Eustylini; (4) Rhinospathe Chevrolat, 1878, from Phyllobiini to Eustylini; and (5) Pachnaeus Schoenherr, 1826, from Tanymecini to Eustylini. © 2012 The Linnean Society of London, Zoological Journal of the Linnean Society, 2012, 164 , 510–557.     

Two new species of Larinus Dejean from North Africa (Coleoptera: Curculionidae), with a key to the striped Larinus (s.str.)

Two new sibling species, Larinus synthesys n. sp. from Algeria, and Larinus boroveci n. sp. from Algeria and Morocco, are described. Both species are tentatively assigned to the subgenus Larinus based on resemblance to species having longitudinal stripes on the elytra and rostrum. L. synthesys n. sp. is the smallest species in Larinus (s. str.) and is closely related to L. boroveci n. sp. The lectotype of Larinus cardopatii Lucas, 1847 is designated and redescribed. An illustrated key to the striped Larinus (s. str.) is given.