Takako Toma

Rapid Adaptive Evolution of Photoperiodic Response during Invasion and Range Expansion across a Climatic Gradient

Understanding the mechanisms of adaptation to spatiotemporal environmental variation is a fundamental goal of evolutionary biology. This issue also has important implications for anticipating biological responses to contemporary climate warming and determining the processes by which invasive species are able to spread rapidly across broad geographic ranges. Here, we compare data from a historical study of latitudinal variation in photoperiodic response among Japanese and U.S. populations of the invasive Asian tiger mosquito Aedes albopictus with contemporary data obtained using comparable methods. Our results demonstrated rapid adaptive evolution of the photoperiodic response during invasion and range expansion across ∼15° of latitude in the United States. In contrast to the photoperiodic response, size-based morphological traits implicated in climatic adaptation in a wide range of other insects did not show evidence of adaptive variation in Ae. albopictus across either the U.S. (invasive) or Japanese (native) range. These results show that photoperiodism has been an important adaptation to climatic variation across the U.S. range of Ae. albopictus and, in conjunction with previous studies, strongly implicate the photoperiodic control of seasonal development as a critical evolutionary response to ongoing contemporary climate change. These results also emphasize that photoperiodism warrants increased attention in studies of the evolution of invasive species.

Aedes (Finlaya) japonicus (Diptera: Culicidae), a Newly Recognized Mosquito in the United States: Analyses of Genetic Variation in the United States and Putative Source Populations

Abstract Introduction of potential disease vectors into a new geographic area poses health risks to local human, livestock, and wildlife populations. It is therefore important to gain understanding of the dynamics of these invasions, in particular its sources, modes of spread after the introduction, and vectorial potential. We studied the population genetics of Aedes (Finlaya) japonicus japonicus (Theobald), an Asian mosquito that was recognized for the first time in the United States in 1998. We examined patterns of genetic diversity using random amplified polymorphic DNA and sequences of ND4 of mtDNA by comparing samples from populations spanning the range of this mosquito in Japan (six samples) and the United States (nine samples) as well as specimens intercepted in New Zealand in 1999. We found geographically differentiated populations in Japan, indicating limited gene flow even on small spatial scales. In the United States, we found evidence of significant genetic differentiation between samples from New York, Connecticut, and New Jersey and those from mid-Pennsylvania and Maryland. We were unable to pinpoint the source location(s) in Japan, although some of the U.S. samples are genetically close to samples from south Honshu and western Kyushu. Further studies should include samples from Korean populations. Distinct genetic signatures in U.S. populations undergoing expansion suggest the possibility of local increases in genetic diversity if and where they meet.

Molecular Phylogenetics of Aedes japonicus, a Disease Vector That Recently Invaded Western Europe, North America, and the Hawaiian Islands

ABSTRACT We used two mitochondrial loci (nicotinamide adenine dinucleotide dehydrogenase subunit 4 and cytochrome oxidase II) and a nuclear locus (28S-D2 spacer) for a total of 1337 bp to evaluate the relationships among the four subspecies of Aedes (Finlaya) japonicus Theobald. Ae. j. japonicus was recently introduced into the United States and has been expanding rapidly. We also included in our analysis a morphologically very closely related species, Aedes (Finlaya) koreicus Edwards, as well as three more distantly related species: Aedes (Finlaya) togoi Theobald, Aedes (Finlaya) hatorii Yamada, and Aedes (Aedimorphus) vexans Meigen. We found that the four subspecies in the Ae. japonicus complex are genetically quite distinct but seem to form a monophyletic group that surprisingly also includes Ae. koreicus, suggesting the need for a taxonomic reconsideration of the group. We also found that the two southern subspecies are more closely related to each other than to any of the remaining subspecies or to Ae. koreicus and may indicate an ancient north—south split of the lineage. Considering the overlap between Ae. j. japonicus and Ae. koreicus, but the stronger association between Ae. koreicus and humans, we are surprised it also has not expanded from its original range. As a proactive reaction to this possibility, we designed and tested a DNA-based rapid assay to differentiate Ae. koreicus from some of the species with which it may be confused in the United States. These Aedes are putative vectors of several important viral encephalitides.

Biological control of container-breeding mosquitoes, Aedes albopictus and Culex quinquefasciatus, in a Japanese island by release of Toxorhynchites splendens adults

Abstract. To control container‐breeding mosquitoes in the small island of Minnajima (0.56km2), northern Okinawa, Japan, laboratory‐reared adults (aged 7–10 days) of Toxorhynchites splendens (Palawan strain), a mosquito with predatory larvae, were released repeatedly during 1984, 1986 and 1987.

Identification of Culex vishnui subgroup (Diptera: Culicidae) mosquitoes from the Ryukyu Archipelago, Japan: development of a species-diagnostic polymerase chain reaction assay based on sequence variation in ribosomal DNA spacers.

Abstract TheCulex vishnuisubgroup includes three important vectors of Japanese encephalitis virus,Culex tritaeniorhynchusGiles,Cx. pseudovishnuiColless, andCx. vishnuiTheobald, all of which occur in the Ryukyu Archipelago, Japan. Although these three species have been shown to be vectors of JE virus in many areas of Southeast Asia, it is not yet known what role each plays in the transmission of the virus in this region. Reliable identification of adult, field-collected specimens is a critical component in epidemiological studies of virus transmission. Mosquitoes in theCx. vishnuisubgroup can be reliably identified in the larval stage. However, because females of these species are very similar, it is difficult to distinguish among them using morphology. We developed a polymerase chain reaction (PCR) assay for the identification of these species. Three species-specific primers were developed for the PCR assay based on a comparative analysis of the nucleotide sequence of the first internal transcribed spacer (ITS1) in the ribosomal DNA gene array. The primers, CT2REV, CP1REV, and CV1REV were designed to amplify a single DNA fragment each fromCx. tritaeniorhynchus,Cx. pseudovishnui,andCx. vishnui,respectively, when paired with a single forward primer that is complementary to the highly conserved 18S rDNA gene. The amplified fragments were separated easily and identified on an agarose gel to facilitate species identification.

HABITATS AND DISTRIBUTION OF ANOPHELES SINENSIS AND ASSOCIATED ANOPHELES HYRCANUS GROUP IN JAPAN

ABSTRACT Mosquito collections were carried out in August 2002 and July 2003 in Japan. Anopheles sinensis of the Hyrcanus Group, Myzomyia Series of Anopheles, was the most common species encountered. The distribution and habitats of 5 Anopheles Hyrcanus Group species are included. Eight species belonging to Aedes, Culex, and Uranotaenia were found associated with An. sinensis in rice paddies and a variety of other larval habitats.

Laboratory evaluation of Toxorhynchites splendens (Diptera: Culicidae) for predation of Aedes albopictus mosquito larvae

Abstract. Biology of the mosquito Toxorhynchites splendens (Wiedemann) was studied in the laboratory to provide baseline data for using the predatory larvae of this species against those of Aedes albopictus (Skuse) in a biological control programme. The mean incubation time of Tx.splendens eggs was 43.8 h and the time required for newly‐hatched larvae to initiate predation was 2.5 h. Mean numbers of prey larvae consumed and killed by each Tx. splendens larva totalled 389 and 345 respectively. The larval period of Tx.splendens was not significantly different for rearing individually or in groups of nine, with equal prey density, and duration of larval development was proportional to prey density. In mass rearing, larval cannibalism was usually observed during days 1–3 post‐eclosion. The incidence of cannibalism decreased sharply on the fourth day after hatching when some larvae became fourth‐instar. Adult female Tx.splendens usually commenced oviposition on day 4 after emergence. The number of eggs laid daily increased on day 7 and the peak oviposition of 6.3 eggs/female/day occurred on day 11. When oviposition containers were provided only intermittently, gravid females of Tx.splendens scattered most of their eggs on the dry floor of the cage. Viability of eggs laid by females aged 4–14 days was high (60–90%) but decreased to less than 40% as the females aged.

Investigation of the aedes (Stegomyia) flavopictus complex (Diptera: Culicidae) in Japan by sequence analysis of the internal transcribed spacers of ribosomal DNA.

Abstract Aedes (Stegomyia) flavopictus Yamada is widely distributed in Japan and Korea. The species comprises three subspecies based on current morphological taxonomy: Aedes f. flavopictus in the Palearctic region of Japan, Ae. f. downsi Bohart & Ingram from Amami and Okinawa Islands, and Ae. f. miyarai Tanaka, Mizusawa & Ingram from Ishigaki and Iriomote Islands of the Ryukyu Archipelago. These subspecies designations are based on observations of a combination of several morphological characters, none of which, by itself is diagnostic for discriminating among the three subspecies. To further study the relationships in this group, we examined the nucleic acid sequence divergence in the internal transcribed spacer regions (ITS1 and ITS2) of the ribosomal DNA gene array of Ae. flavopictus individuals collected at five sites from three geographic regions in Japan. Analysis of sequence data by distance and maximum parsimony methods produced phylogenetic trees that showed separation of the specimens into three major clades, corresponding to both subspecies and geographic region. These results were consistent with and support the current classification and geographic distribution of the three subspecies.

Susceptibility of the Mosquitoes Anopheles minimus, An. sinensis, and An. saperoi (Diptera: Culicidae) from the Ryukyu Archipelago, Japan, to the Rodent Malaria Plasmodium yoelii nigeriense

Abstract The susceptibility of three anopheline mosquitoes, Anopheles minimus Theobald, An. sinensis Wiedemann, and An. saperoi Bohart & Ingram, from the Ryukyu Archipelago to the rodent malaria, Plasmodium yoelii nigeriense was examined to find new vectors other than An. stephensi Liston for rodent malaria studies in the laboratory. The survival rate of the mosquitoes after feeding on mice infected with P. y. nigeriense was also examined. The Beech strain of An. stephensi from India was compared with An. minimus from Ishigaki Island, and An. sinensis and An. saperoi from Okinawa Island. Oocysts were first found on day 3 after feeding on mice infected with P. y. nigeriense in An. stephensi, on day 4 in An. minimus and An. saperoi, and day 6 in An. sinensis. From 8 to 14 d after feeding on malaria-positive mice, oocysts were present in 97.2–100% of An. stephensi, 85.7–100% of An. saperoi, 20–74.1% of An. minimus, and 12.5–13.3% of An. sinensis. The duration of oocyst occurrence in An. saperoi was 55 d, the longest among the anopheline mosquitoes used in this study. On day 8 after feeding, sporozoites were found in the salivary glands and heads of all the mosquitoes tested. From the 10th to 16th d, sporozoites were present in the salivary glands of 14.9% (range, 9.1–28.0%) of An. minimus, 47.3% (40.7–58.1) of An. saperoi, and 96.2% (94.1–97.2) of An. stephensi, but were absent in An. sinensis. Anopheles saperoi could be an excellent vector of P. y. nigeriense because it has comparatively high susceptibility and high longevity even after feeding on infected mice.

New Records of Mosquito Species for Different Islands of the Ryukyu Archipelago, Japan

Abstract During an 11-year study period (1999–2010) on the islands of the Ryukyu Archipelago, Japan, 20 mosquito species were identified. The results revealed new records of species hitherto not reported on these islands. This contribution updates the mosquito fauna of the various islands of the Ryukyu Archipelago.