Integrated species delimitation and conservation implications of an endangered weevil Pachyrhynchus sonani (Coleoptera: Curculionidae) in Green and Orchid Islands of Taiwan

Oceanic islands are productive habitats for generating new species and high endemism, which is primarily due to their geographical isolation, smaller population sizes and local adaptation. However, the short divergence times and subtle morphological or ecological divergence of insular organisms may obscure species identity, so the cryptic endemism on islands may be underestimated. The endangered weevil Pachyrhynchus sonani Kôno (Coleoptera: Curculionidae: Entiminae: Pachyrhynchini) is endemic to Green Island and Orchid Island of the Taiwan‐Luzon Archipelago and displays widespread variation in coloration and host range, thus raising questions regarding its species boundaries and degree of cryptic diversity. We tested the species boundaries of P. sonani using an integrated approach that combined morphological (body size and shape, genital shape, coloration and cuticular scale), genetic (four genes and restriction site‐associated DNA sequencing, RAD‐seq) and ecological (host range and distribution) diversity. The results indicated that all the morphological datasets for male P. sonani, except for the colour spectrum, reveal overlapping but statistically significant differences between islands. In contrast, the morphology of the female P. sonani showed minimum divergence between island populations. The populations of P. sonani on the two islands were significantly different in their host ranges, and the genetic clustering and phylogenies of P. sonani established two valid evolutionary species. Integrated species delimitation combining morphological, molecular and ecological characters supported two distinct species of P. sonani from Green Island and Orchid Island. The Green Island population was described as P. jitanasaius sp.n. Chen & Lin, and it is recommended that its threatened conservation status be recognized. Our findings suggest that the inter‐island speciation of endemic organisms inhabiting both islands may be more common than previously thought, and they highlight the possibility that the cryptic diversity of small oceanic islands may still be largely underestimated.     

Captive breeding of two insular populations of Pachyrhynchus sarcitis (Coleoptera: Curculionidae) from Lanyu and Babuyan Islands

Endemic species of oceanic islands are vulnerable due to their geographical isolation and small population sizes. For endangered island species, captive breeding program is an important strategy for conservation and sustainable management. Pachyrhynchus sarcitis is a flightless weevil decorated with colourful markings and distributed exclusively on several islets of Southeast Asia including Lanyu, Ludao, and Babuyan Islands. The life history of threatened Pachyrhynchus species of these islands was poorly known. This study reared P. sarcitis from Lanyu and Babuyan Islands for the first time in the laboratory using their host plant (Leea guineensis). The two island populations showed significant differences in instar numbers, in which the weevils from Lanyu Island had a higher instar number regardless the length of developmental duration. The adult body size of both sexes of the Babuyan population became smaller under laboratory condition; whereas Lanyu population in the laboratory did not show this trend. The differences in larval development might suggest local adaptation to the host plants or other life history associated characteristics which requires further research.     

Helicobacter pylori and hepatitis A virus infection in school-aged children on two isolated neighborhood islands in Taiwan

Background. The transmission routes of Helicobacter pylori and hepatitis A virus (HAV) infections have been extensively discussed in previous literature. However, whether H. pylori and HAV shared the same transmission pattern or not remains unclear. Lower socioeconomic status was recognized as a consistent risk factor to both infections. However, whether fecal‐oral transmission was a risk factor to both infections is still under debate. Materials and Methods. In 1996, we conducted a cross‐sectional study to evaluate the seroprevalence of antibody to H. pylori and HAV among the randomly selected school‐aged children (age between 13 and 15) on Green Island (n = 91) and Lanyu Island (n = 138) (two isolated neighborhood islands near Taiwan Main Island). Results. The seroprevalence of H. pylori and HAV on the Green Island were 82.4% and 5.5%, respectively. The seroprevalence of H. pylori and HAV on Lanyu Island were 71.0% and 90.6%, respectively. H. pylori seroprevalence of all children and the subgroup of 13‐year‐olds was significantly lower on Lanyu Island than Green Island. However, it was not significantly different in subgroups of 14‐ and 15‐year‐olds. HAV seroprevalence was significantly higher on Lanyu Island than Green Island among all children and in each age subgroup. The correlation of H. pylori infection and HAV infection did not demonstrate significant linear correlation on both islands. Conclusions. In conclusion, H. pylori and HAV infections in school‐aged children of 13–15 years of age on Green Island and Lanyu Island did not demonstrate significant correlation. The results of this study imply that H. pylori and HAV may share different transmission routes of infection.     

Conservation of Hipparchia autonoe (Esper) (Lepidoptera: Nymphalidae), Natural Monument in South Korea

Hipparchia autonoe, designated Natural Monument no. 458, is a species inhabiting Mt. Halla on Jeju Island, which is the only habitat of H. autonoe in South Korea. Recently, this species is in danger of extinction as the population has been remarkably reduced because its host plants and nectar plants are gradually disappearing due to plant succession in its habitat. On the climbing road of Jeju Island Eorimok, H. autonoe used to be observed at and above 1300 m a.s.l. However, since Sasa quelpaertensis has recently expanded its habitat up to 1400 m, H. autonoe is now observed at and above 1500 m. In Mongolia, the population of H. autonoe seems quite stable as there are dense populations of a host plant and nectar plants. Accordingly it is judged that we can maintain a stable community for support of H. autonoe if we successfully manage the host plant and nectar plant communities by controlling the density of S. quelpaertensis using biological and physical control methods. Comparative analysis of DNA barcode region of COI (658 bp) was done to check the homogeneity and the genetic diversity of H. autonoe collected in Korea and Mongolia. The DNA sequence difference among individuals collected in South Korea was 0.0–0.2%, and in Mongolia 0.0–1.4%. This confirmed a 0.06–1.2% barcoding gap. We believe that this result will provide basic information useful to guide conservation of H. autonoe in South Korea.     

High frequency of extrapair copulation with low level of extrapair fertilization in the Lanyu scops owl Otus elegans botelensis

The genetic mating system is known for only a few species of owls. Most of them are genetically monogamous. The Lanyu scops owl Otus elegans botelensis breeds in high density in the forests of Lanyu (Orchid Island), southeast of Taiwan. Because extra‐pair copulations (EPCs) have frequently been observed, we suspected a high degree of extra‐pair fertilizations (EPFs). Through intensive field observations we quantified both within‐pair and extra‐pair copulations of this owl, and used a set of microsatellite loci to assign the parentage of 200 offspring from 108 families collected between 1999 and 2004. Among copulations for which we could identify both participants, 19.77% were EPC. However, no EPC was observed in the period just before egg laying. We found only two cases of parentage mismatch among 108 broods. The first case was most likely a case of brood usurpation, which would be the first case reported in the Strigidae. The second case was an EPF resulting in one offspring. Our study found that parentage mismatch in the Lanyu scops owls to be extremely low, therefore reproductive success can be estimated accurately by simply counting chicks in nests.     

Ustilago aldabrensis, a new species from Seychelles, and two other smut fungi on Dactyloctenium

A new smut fungus Ustilago aldabrensis on the grass Dactyloctenium ctenoides is described and illustrated from Aldabra Island in the Seychelles archipelago. It is compared with similar species known on the genus Dactyloctenium. A further two smut fungi infecting this host plant genus, U. dactyloctenii-gigantei and U. idonea are discussed. U. dactyloctenii-gigantei is reported for the first time from Nigeria on a new host plant, D. aegyptium. U. idonea is redescribed, and its nomenclature and geographical distribution are clarified. A key to smut fungi infecting species of Dactyloctenium is provided.     

Holocene uplifted coral reefs in Lanyu and Lutao Islands to the southeast of Taiwan

Lanyu and Lutao Islands to the southeast of Taiwan are located in the northern extension of the Luzon Arc. Crustal deformation of these islands provides a key to understand the collision of the Luzon Arc against Taiwan. To clarify the style and the rate of vertical movement during the Holocene, uplifted coral reefs fringing these two islands were investigated. Living corals were also investigated for comparison with fossil corals. It was found that Isopora palifera lives dominantly in the upper slope of the present-day fringing coral reefs in Lanyu Island at an average depth of 101 ± 46 cm (one standard deviation) below mean sea level. Using I. palifera as an accurate indicator of paleo-sea levels, Holocene relative sea-level changes were reconstructed. Lanyu Island has been uplifted continuously at a rate of 2.0 mm yr⁻¹, at least during the late Holocene from 2,269 cal. yr BP to the present. Lutao Island has been uplifted at an average rate of 1.2 mm yr⁻¹, since at least 5,749 cal. yr BP, although it is unclear whether the uplift was continuous. The present observations, combined with the GPS displacement field and deep crustal structure, suggest that the continuous uplift is related to aseismic slip on the Longitudinal Valley Fault.     

Description of a new species of the genus Anochetus Mayr (Hymenoptera: Formicidae) from Orchid Island,Taiwan

A trap-jaw ant Anochetus lanyuensis sp. nov., belonging to the graeffei species group Brown, 1978, is described on the basis of worker and gyne castes from Orchid Island (Lanyu), Taiwan. A. lanyuensis sp. nov. is easily distinguishable from the other species in the graeffei species group by the large size of its eye, longitudinal striae on the pronotal disc, and concave dorsal margin of the petiolar node in the frontal view.     

Modelling the climate suitability of green carpenter bee (Xylocopa aerata) and its nesting hosts under current and future scenarios to guide conservation efforts

Due to local extinction, the endangered green carpenter bee (Xylocopa aerata) has a disjunct distribution in the southeast of Australia. The species relies on dead softwood from a small selection of plant species for making its nests. Habitat fragmentation, combined with deleterious fire events, is thought to have negatively impacted on nesting substrate availability and recolonisation chances. Here, we use MaxEnt algorithm to model both the current distribution and the effect of climate change scenarios on the distribution of both X. aerata and four plant species that provide most of its nesting substrate: Banksia integrifolia, B. marginata, Xanthorrhoea arborea and Xanthorrhoea semiplana subsp. tateana. The annual mean temperature is the strongest climatic predictor of the distribution of X. aerata and its host plants. The modelled distribution of the bee under current climatic conditions indicates that climatic factors are unlikely to cause local extinctions. In all future scenarios, suitable areas for X. aerata and each of its nesting hosts are expected to contract towards the southeast of mainland Australia. The suitability of Kangaroo Island for the bee and its current local current host species is maintained in all scenarios, while Tasmania will become increasingly suitable for all species. The Grampians National Park in western Victoria, where the bee species were last seen outside of its current range (in the 1930s), is predicted to remain suitable for X. aerata and several host plants under all scenarios. Therefore, this relatively large area of native vegetation may be a good case study for re‐introduction as part of future conservation efforts.     

Induction of host‐plant preference in Brontispa longissima (Gestro) (Coleoptera: Chrysomelidae)

The coconut hispine beetle, Brontispa longissima, is a serious invasive pest of coconut palm (Cocos nucifera) in Southeast Asia and the Pacific region. Recently, however, it has been reported that on Ishigaki Island, Japan, where C. nucifera is rare, the endemic palm Satakentia liukiuensis serves as a regular host. To explore this phenomenon, we examined the preferences of the beetle for the two palm hosts by choice tests and measurements of feeding amount using four different groups: naïve adults obtained from a colony reared on C. nucifera (NC); naïve adults from a colony reared on S. liukiuensis (NS); wild adults collected on C. nucifera on Ishigaki Island (WC); and wild adults collected on S. liukiuensis on Ishigaki Island (WS). Both NC and NS consumed significantly more C. nucifera leaves than S. liukiuensis leaves and were observed more often on the leaves of C. nucifera. These results suggest that B. longissima has an innate preference for C. nucifera. In contrast, field‐collected adults (WC, WS) initially showed preferences for the species on which they were collected, but after exposure to the other host plant for 2 weeks, their preferences for that host increased. These results suggest that the feeding experience of B. longissima adults affected their subsequent host preferences. This flexible host preference may have facilitated the beetle’s successful establishment in novel Japanese habitats where the original host plant was rare.     

Karyotypes of two rare species of hynobiid salamanders from Taiwan,Hynobius sonani (Maki) and Hynobius formosanus Maki (Urodela)

Two salamanders endemic to Taiwan, Hynobius sonani and Hynobius formosanus, are similar to Japanese hynobiids in having 2n = 58 chromosomes and by the absence of a medium-sized telocentric pair. H. sonani has 6 metacentric and 10 telocentric microchromosome pairs, while H. formosanus has 5 metacentric and 11 telocentric microchromosome pairs. The karyotype of H. sonani is nearly identical to that of the Japanese hynobiid salamander H. boulengeri. Karyophylogenetic relationships among forms of mountain-stream type hynobiids of Taiwan and Japan are discussed.This paper is dedicated to late Mr. Tadao Yamamoto who died on November 2, 1985.     

First detection of the fruit fly Bactrocera synnephes (Diptera: Tephritidae) on Iriomote Island, south-western Japan

Bactrocera synnephes, a fruit fly species common in Taiwan, was first detected in Japan in 2006, when several male adults were trapped at Kanokawa, an uninhabited coastal area of Iriomote Island, Okinawa (200 km from Taiwan). To examine potential colonization by B. synnephes, we collected fruit of the wild gourd Diplocyclos palmatus (Cucurbitaceae), a host of the fruit fly, as well as other cucurbitaceous species, in Kanokawa and other areas of Iriomote Island. No B. synnephes infestation was found on any cucurbitaceous plants. Based on these results, we conclude that B. synnephes had very recently invaded Iriomote Island, possibly from Taiwan, but had failed to colonize. Bactrocera synnephes may have entered Iriomote Island by natural or human‐induced means, but a human‐induced invasion pathway is less likely in light of the non‐pest status of the species in Taiwan. The disappearance of B. synnephes from Iriomote Island may be related to the loss of host fruit as a result of tropical cyclones. Although B. synnephes is not regarded as a pest elsewhere, the pest status of the species is unclear in Okinawa, where no fruit fly pests currently attack commercial cucurbitaceous fruit. We must remain alert for possible re‐invasion by B. synnephes.     

Comparison of the life history strategies of three Dysdercus true bugs (Heteroptera: Pyrrhocoridae), with special reference to their seasonal host plant use

The life history strategy and seasonal host plant use of three Dysdercus bugs (D. cingulatus, D. poecilus and D. decussatus) were compared based on 2 years of twice monthly observations on Ishigaki‐jima Island (24°N, 124°E) in the southernmost part of Japan. Dysdercus poecilus reproduced almost year round, exclusively on Sida rhombifolia, which bears fruit and/or seeds almost year round. Dysdercus cingulatus reproduced successively on various malvaceous and bombacaceous plant species, according to their seasonal fruiting cycles. Its robust reproduction was observed on Hibiscus makinoi during winter months and on Chorisia speciosa and Bombax ceiba in early summer, whereas small‐scale reproduction was observed on various malvaceous plant species during the summer and autumn months. Dysdercus decussatus reproduced on Hibiscus tiliaceus and Thespesia populnea during the summer months, when these host plants bear abundant fruit and/or seeds, and its adults formed conspicuous aggregations without copulation on the underside of the leaves of those plants from November to the subsequent May. Each Dysdercus species showed a species‐specific life history strategy according to the differences in the phenology of their host plants.     

Host plant preference and performance of the sibling species of butterflies <Emphasis Type="Italic">Leptidea sinapis</Emphasis> and <Emphasis Type="Italic">Leptidea reali</Emphasis>: a test of the trade-off hypothesis for food specialisation

A large proportion of phytophagous insect species are specialised on one or a few host plants, and female host plant preference is predicted to be tightly linked to high larval survival and performance on the preferred plant(s). Specialisation is likely favoured by selection under stable circumstances, since different host plant species are likely to differ in suitability—a pattern usually explained by the “trade-off hypothesis”, which posits that increased performance on a given plant comes at a cost of decreased performance on other plants. Host plant specialisation is also ascribed an important role in host shift speciation, where different incipient species specialise on different host plants. Hence, it is important to determine the role of host plants when studying species divergence and niche partitioning between closely related species, such as the butterfly species pair Leptidea sinapis and Leptidea reali. In Sweden, Leptidea sinapis is a habitat generalist, appearing in both forests and meadows, whereas Leptidea reali is specialised on meadows. Here, we study the female preference and larval survival and performance in terms of growth rate, pupal weight and development time on the seven most-utilised host plants. Both species showed similar host plant rank orders, and larvae survived and performed equally well on most plants with the exceptions of two rarely utilised forest plants. We therefore conclude that differences in preference or performance on plants from the two habitats do not drive, or maintain, niche separation, and we argue that the results of this study do not support the trade-off hypothesis for host plant specialisation, since the host plant generalist Leptidea sinapis survived and performed as well on the most preferred meadow host plant Lathyrus pratensis as did Leptidea reali although the generalist species also includes other plants in its host range. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Probable long distance dispersal of Leptinella plumosa Hook.f. to Heard Island: habitat, status and discussion of its arrival

During the 2003–2004 austral summer the number of vascular plant species recorded from Heard Island rose from 11 to 12 with the discovery of one small plant of Leptinella plumosa Hook.f. (Asteraceae), an indigenous subantarctic species. It is described and its habitat, likely status and possible means of arrival on the island are discussed. We conclude that the species probably arrived by natural means with a seabird as its most likely dispersal vector. The life history and biology of L. plumosa indicates its likely persistence on Heard Island.     

The role of ecological availability and host plant characteristics in determining host use by the bogus yucca moth Prodoxus decipiens

1. Specialisation in host plant use is strongly correlated with speciation in many plant‐feeding insects. Specialised taxa, however, could be restricted in host range due to limits in ecological availability of host plant species rather than trade‐offs in using alternate host species. 2. Moths in the genus Prodoxus are extreme specialists on Yucca and speciation is closely tied to host plant shifts. However, many Yucca ranges are allopatric. This study examined whether the bogus yucca moth Prodoxus decipiens is limited in host range because of biogeographic factors or due to differences in the characteristics of host plant species. 3. In a common garden, local P. decipiens moths that use Yucca filamentosa were exposed to individuals of five Yucca species, two that are known hosts of P. decipiens in other parts of its range and three that are used by its sister species, Prodoxus quinquepunctellus. 4. Local moths were attracted to flowers of all Yucca species and females attempted oviposition in the flowering stalks of all species. However, larvae successfully completed development to diapause in only one of the five host plant species. Larval development on non‐natal Yucca species was significantly reduced compared with the local host. 5. The results suggest that differences in host plant characteristics among Yucca species would result in strong natural selection during a host shift. Thus, specialisation in host plant use is probably due to trade‐offs involved with using novel host plant species as well as ecological availability.     

Study of COI sequences from endemic New Zealand aphids highlights high mitochondrial DNA diversity in Rhopalosiphina (Hemiptera: Aphididae)

Focussed searches were made across New Zealand between 2013 and 2016, for endemic aphids from the Schizaphis (Rhopalosiphina) genus, which is currently represented by two putative, undescribed species from the endemic host plants Aciphylla and Dracophyllum. Cytochrome c oxidase I (COI) gene sequences (48 in total) from the Schizaphis were analysed together with those from a broader collection of New Zealand endemic aphids that has been assembled since the year 2000. The bulk of the Schizaphis belonged to two clusters corresponding to the host plant genera. Two aphids from central North Island Dracophyllum represented a much diverged lineage without clear affiliations to other New Zealand Schizaphis. Inter-population variation in the New Zealand Schizaphis was high compared with that seen in international studies of Aphidinae and among populations of other endemic New Zealand Aphidina. Within Schizaphis from Dracophyllum, geography played an apparent role in genetic structuring, with populations from Taranaki (North Island) and especially Mt Lyford (South Island) being divergent from those on the South Island main divide. Two distinct lineages of Schizaphis, which co-occurred at some sites, were found on Aciphylla. Our sequence comparisons, including GMYC analyses, indicated up to five New Zealand Schizaphis lineages, and two newly discovered endemic Aphis species from the host plants Clematis and Hebe.     

Allozyme variations in Michelia formosana (Kanehira) Masamune(Magnoliaceae), and the inference of a glacial refugium in Taiwan

  Michelia formosana is distributed islandwide in Taiwan and also occurs in the Ryukyu Archipelago, Japan. Allozyme genetic variability in M. formosana (Magnoliaceae) was investigated using five polymorphic loci (Pgm-1, Idh-1, Mr-1, Skdh-1, and Ppo-3) from five enzyme systems. The average value of expected heterozygosity (H_e) describing within-population variation was 0.241. The overall F_is (0.0736) indicates a significant deficiency of heterozygotes at the population level. This positive value of F_is is mainly contributed from populations found at Pinglin, Chienshi, Wufeng, and the Ryukyus, all located at northern latitudes, and is also caused by locus Ppo-3. Among-population variation, F_st, accounted for 10.6% of the total heterozygosity and deviates significantly from zero. This populational differentiation agrees well with that of general tropical woody species outcrossed by animals. The population on Lanyu (Orchid Island) has some morphological differences from the plants native to Taiwan, but this was not reflected in the cluster analysis using the unweighted pair group method. Pgm-1d, however, is the diagnostic characteristic for the plants growing on Lanyu as it is entirely absent from all other populations of M. formosana. About 18% of the alleles of this study show clinal geographical variation and were found to be significantly related to latitudinal gradients throughout the species range. This, however, is not a linear relation but a curve with a peak form which is exactly the same as the relationship between expected heterozygosities of populations and latitude. This observation suggests that Nantou County, in central Taiwan could be a glacial refugium for genetic diversity of M. formosana. This inference is further supported by analysis of published studies on seven widely spread plant species in Taiwan. Location of this glacial refugium is probably related to the first emergence of the Central Mountain Ridge millions of years ago that naturally becamed the major window to receive genetic diversity from the Asian mainland. Received: 16 April 2000 / Accepted: 12 May 2000     

Pollination mutualism between a new species of the genus Colocasiomyia de Meijere (Diptera: Drosophilidae) and Steudnera colocasiifolia (Araceae) in Yunnan, China

A new species of the genus Colocasiomyia de Meijere (Diptera: Drosophilidae) was discovered from inflorescences of Steudnera colocasiifolia K. Koch (Araceae) in Yunnan, China. The new species is described as Colocasiomyia steudnerae Takenaka and Toda, sp. nov., and we investigated the reproductive ecology of both the fly and the plant species. This fly species reproduces in the inflorescences/infructescences of the plant, and depends almost throughout its entire life cycle on the host plant. The fly species is the most abundant flower visitor for S. colocasiifolia and behaves intimately with the flowering events, suggesting that it is the unique and most efficient pollinator for the host plant. Bagging (insect‐exclusion) treatment of inflorescences resulted in no fruits. These findings strongly suggest that intimate pollination mutualism has evolved between the fly and the host plant, as are known in other Colocasiomyia flies and Araceae plants. One notable feature of this system is that the new species almost monopolizes the host‐plant inflorescence as a visitor, without any cohabiting Colocasiomyia species. In comparison to other cases where two Colocasiomyia species share the same inflorescence and infructescence of Araceae host plants for reproduction by separating their breeding niches microallopatrically between the staminate (upper male‐flower) and the pistillate (lower female‐flower) regions on the spadix, C. steudnerae exhibits a mixture of stamenicolous and pistillicolous breeding habits.