Future risk of the maize orange leafhopper,Cicadulina bipunctata,and maize wallaby ear symptom in temperate Japan

Since the 1980s, the maize orange leafhopper, Cicadulina bipunctata, has been gradually expanding its range in east Asia associated with global warming. This leafhopper induces maize wallaby ear symptom (MWES) on young maize plants and has become a threat to forage maize production in southern parts of temperate Japan since around 2000. In this study, using predictions of future temperature and precipitation calculated from Atmosphere–Ocean Coupled General Circulation Models, the future risk of C. bipunctata expansion and MWES occurrence in Japan (spatial resolution: 1 km²) was predicted. A nominal logistic regression analysis showed a significant contribution of cumulative low temperature during winter to the current distribution of C. bipunctata. The range of C. bipunctata was predicted to expand northward, particularly in Kyushu, Shikoku and the southern part of Honshu after the 2060s. Predicted intensification of MWES would reduce the efficacy of maize cultivars that are currently tolerant to MWES, in southern Kyushu in the 2020s, and in most parts of Kyushu, Shikoku and southwestern Honshu in the 2060s. These results suggest the need for measures to counter further expansion of C. bipunctata and improvement of current tolerant maize cultivars.     

Host Feeding by an Herbivore Improves the Performance of Offspring

Herbivores have developed diverse strategies to manipulate host plants for their own benefits. The gall induction by the maize orange leafhopper Cicadulina bipunctata is different from that by other gall-inducing insects in that the galls are induced not on feeding sites but appear on distant, newly developing leaves. In addition, adult C. bipunctata are highly mobile and seldom feed on gall tissue that they have induced. These mean that the gall induction by C. bipunctata is unlikely to contribute to the fitness of the inducer itself. The objective of this study was to determine whether manipulation of the host plant by this leafhopper has a subsequent benefit to offspring. Adults feeding on maize seedlings caused a partial change in the glucose content and a remarkable change in the accumulation of free amino acids in the gall tissue. Increases in emergence and developmental rates were observed in nymphs feeding on gall tissue induced by prior adult feeding. Such improvements were not evident in nymphs feeding on a C. bipunctata-resistant variety, which rarely displays galls after C. bipunctata feeding, nor on maize seedlings previously foraged by another leafhopper, Psammotettix striatus. The results indicate that gall tissue induced by adult C. bipunctata contribute to better performance of its offspring through improvement of the nutritional components of host plants.     

Cold tolerance of the maize orange leafhopper,Cicadulina bipunctata

Cicadulina bipunctata was originally distributed in tropical and subtropical regions of the Old World. This leafhopper recently expanded its distribution area to southern parts of temperate Japan. In this study, factors affecting the overwintering ability of C. bipunctata were examined. A series of laboratory experiments revealed that cold acclimation at 15 °C for 7 days enhanced the cold tolerance of C. bipunctata to the same level as an overwintering population, adult females were more tolerant of cold temperature than adult males, and survival of acclimated adult females was highly dependent on temperature from −5 to 5 °C and exposure duration to the temperature. The temperature of crystallization of adult females was approximately −19 °C but temperatures in southern temperate Japan rarely dropped below −10 °C in the winter, indicating that overwintering C. bipunctata adults in temperate Japan are not killed by freezing injury but by indirect chilling injury caused by long-term exposure to moderately low temperatures. An overwintering generation of C. bipunctata had extremely low overwinter survival (<1%) in temperate Japan; however, based on winter temperature ranges, there are additional areas amenable to expansion of C. bipunctata in temperate Japan.     

Estimation of climatic factors relating to occurrence of the maize orange leafhopper, Cicadulina bipunctata

The maize orange leafhopper, Cicadulina bipunctata is a serious pest of forage maize in East and Southeast Asia. In temperate Japan, the occurrence of this leafhopper fluctuates widely among years. Here, we examined effects of climatic factors (temperature, precipitation and sunlight) on the occurrence of C. bipunctata. Seasonal occurrence of adult C. bipunctata in a census field from July to August, when forage maize was most susceptible to the pest, could be described by a simple exponential function with two parameter: estimated density of C. bipunctata on 1 July (N ₀) and intrinsic rate of natural increase (r) for each year. Forward stepwise multiple regression analysis using seasonal occurrence data from 2004 to 2009 detected positive contributions of average temperatures in the previous December and February and a negative contribution of total precipitation during the previous winter to N ₀. The analysis also indicated that average temperature in July of the current year and N ₀ contributed positively and negatively to r, respectively. These results indicated that high temperature and little precipitation during winter and high temperature in early summer induced high occurrence of C. bipunctata in summer. A prediction model based on these factors supported the actual seasonal occurrence in 2010, suggesting that this prediction model is applicable to C. bipunctata forecasting. The prediction model indicated that further global warming in the future is likely to cause further outbreaks of C. bipunctata.     

Phytohormones Related to Host Plant Manipulation by a Gall-Inducing Leafhopper

The maize orange leafhopper Cicadulina bipunctata (Hemiptera: Cicadellidae) induces galls characterized by growth stunting and severe swelling of leaf veins on various plants of Poaceae. Previous studies revealed that galls are induced not on feeding site but on distant, newly extended leaves during the feeding, and strongly suggested that some chemicals injected by the leafhopper affect at the leaf primordia. To approach the mechanism underlying gall induction by C. bipunctata, we examined physiological response of plants to feeding by the leafhopper. We performed high-throughput and comprehensive plant hormone analyses using LC-ESI-MS/MS. Galled maize leaves contained higher contents of abscisic acid (ABA) and trans-Zeatin (tZ) and lower contents of gibberellins (GA₁ and GA₄) than ungalled maize leaves. Leafhopper treatment significantly increased ABA and tZ contents and decreased GA₁ and GA₄ contents in extending leaves. After the removal of leafhoppers, contents of tZ and gibberellins in extending leaves soon became similar to the control values. ABA content was gradually decreased after the removal of leafhoppers. Such hormonal changes were not observed in leafhopper treatment on leaves of resistant maize variety. Water contents of galled leaves were significantly lower than control leaves, suggesting water stress of galled leaves and possible reason of the increase in ABA content. These results imply that ABA, tZ, and gibberellins are related to gall induction by the leafhopper on susceptible variety of maize.     

Oviposition site preference and developmental performance of a gall‐inducing leafhopper on galled and non‐galled host plants

Oviposition preferences of herbivorous insects affect offspring performance. Both positive and negative links between oviposition preference and offspring performance have been reported for many species. A gall‐inducing leafhopper, Cicadulina bipunctata Melichar (Hemiptera: Cicadellidae), feeds on various Poaceae plants and induces galls of enhanced nutritional value for their offspring. Although gall induction by C. bipunctata improves nymphal performance, the oviposition preference of females between galled and non‐galled host plants is still unclear. In this paper, the nymphal performance and oviposition and feeding‐site preference of C. bipunctata were investigated using galled wheat, Triticum aestivum L., and non‐galled barley, Hordeum vulgare L., as host plants. The survival rate of C. bipunctata on wheat was significantly higher than on barley. In the choice test, significantly more eggs were laid into barley, whereas the number of eggs deposited on both hosts was not significantly different in the no‐choice test. The number of settling individuals per leaf area was not significantly different between wheat and barley, suggesting no clear preference for oviposition between these plants. Experience as a nymph with a growing host did not affect oviposition preference as adult female. The inconsistent correspondence between offspring performance and oviposition preference of C. bipunctata may reflect the high mobility of nymphs and/or differences in leaf area between host plants. The results indicate that the previous finding that oviposition preference and offspring performance are not always positively correlated in herbivorous insects is applicable to gall‐inducing insects.     

Clarification of the Identity of the Tea Green Leafhopper Based on Morphological Comparison between Chinese and Japanese Specimens

Tea green leafhopper is one of the most dominant pests in major tea production regions of East Asia. This species has been variously identified as Empoasca vitis (Goëthe), Jacobiasca formosana (Paoli) and Empoasca onukii Matsuda in Mainland China, Taiwan and Japan, respectively. Recent study of DNA sequence data suggested that treatment of this pest as different species in these three adjacent regions is incorrect and that they were a single species; but the correct scientific name for the species has remained unclear. Consistent with the prior molecular evidence, morphological study shows that the male genital characters of Chinese specimens are the same as those of specimens from Japan, so the correct scientific name of tea green leafhopper in China is Empoasca (Matsumurasca) onukii Matsuda.     

Egg parasitoids (Hymenoptera: Mymaridae and Trichogrammatidae) of okra leafhopper,Amrasca biguttula (Ishida) (Hemiptera: Cicadellidae), in southern Japan

The okra leafhopper, Amrasca (Sundapteryx) biguttula (Ishida) (Hemiptera: Cicadellidae), is an economically important pest of okra, Abelmoschus esculentus (L.) Moench (Malvaceae), in Japan. During 2019 and 2022 surveys for its egg parasitoids (Hymenoptera) were conducted on Kyushu and Ryukyu islands in southern Japan. Five species of Mymaridae, Anagrus (Anagrus)?atomus (L.), A. (A.) japonicus Sahad, A. (A.) turpanicus Triapitsyn and Hu, Arescon enocki (Subba Rao and Kaur), and Stethynium empoascae Subba Rao, as well as one of Trichogrammatidae, Pseudoligosita nephotetticum (Mani), were recorded from eggs of A. biguttula on okra in Japan. Of these, A. (A.) turpanicus, identified using a combination of molecular and morphological methods, is newly recorded from Japan; A. (A.)?atomus, A. (A.) turpanicus and P. nephotetticum are for the first time reported from eggs of okra leafhopper.     

Wild host plants of four spider mite species (Acari: Tetranychidae) infesting fruit crops in Okinawa

To determine the wild host plants of four major spider mite species infesting fruit crops grown in the subtropical Asian region, we collected and identified mites from non-crop plants throughout the islands of Okinawa, southwestern Japan. Although the two mango pests, Oligonychus coffeae and O. biharensis, are polyphagous, they did not share any wild host species in the field, indicating that their source plants are completely separate in Okinawa. Several major wild hosts were determined for Eutetranychus africanus, a pest of papaya and citrus. Its host range partially overlapped with that of O. biharensis. The citrus red mite, Panonychus citri, was very rare on non-crop plants in Okinawa, suggesting that it maintains its population mainly on the citrus trees in the area. These results are of great significance when considering vegetation control as part of the integrated management of these pest mites. During our survey, two non-pest species, Panonychus caglei (new to Japan) and Oligonychus gotohi (new to Okinawa), were also found.     

Attributes ofCalamagrostis langsdorffii,C. sachalinensis and their intermediates in Shikoku

The internal structure of theCalamagrostis langsdorffii-sachalinensis complex in Shikoku was examined on the basis of a number of new collections. Chromosome counts disclosed the occurrence of the following entities: 6X and 8X, as well as some aneuploids, ofC. longsdorffii; 6X ofC. sachalinensis; 6X and 8X of the intermediates between these two species. The examination of pollen of these collections and the herbarium specimens showed that all of them may be apomictic in seed formation and that the sexual tetraploids of the pertinent complex may not be distributed in Shikoku. Details of morphological features of the collections were analyzed, and the intermediacy in morphology of the group “intermediate” was confirmed by the distance diagram method. The collections ofC. langsdorffii from Shikoku generally showed an awn longer than that of the collections from Honshu and Hokkaido. On the basis of these and some other observations, relationships of the strains extant in Shikoku were discussed. For the derivation of the present-day composition of the pertinent complex, an hypothesis which is founded on the presumption of a past occurrence of sexual tetraploids of bothC. langsdorffii andC. sachalinensis in Shikoku has been advanced.     

Did maize domestication and early spread mediate the population genetics of corn leafhopper?

Investigating how crop domestication and early farming mediated crop attributes, distributions, and interactions with antagonists may shed light on today's agricultural pest problems. Crop domestication generally involved artificial selection for traits desirable to early farmers, for example, in creased productivity or yield, and enhanced qualities, though invariably it altered the interactions between crops and insects, and expanded the geographical ranges of crops. Thus, some studies suggest that with crop domestication and spread, insect populations on wild crop ancestors gave rise to pestiferous insect populations on crops. Here, we addressed whether the emergence of corn leafhopper (Dalbulus ma id is) as an agricultural pest may be associated with domestication and early spread of maize (Zea mays mays). We used AFLP markers and mitochondrial COI sequences to assess population genetic structuring and haplotype relationships among corn leafhopper samples from maize and its wild relative Zea diploperennis from multiple locations in Mexico and Argentina. We uncovered seven corn leafhopper haplotypes contained within two haplogroups, one haplogroup containing haplotypes associated with maize and the other containing haplotypes associated with Z. diploperennis in a mountainous habitat. Within the first haplogroup, one haplotype was predominant across Mexican locations, and another across Argentinean locations;both were considered pestiferous. We suggested that the divergence times of the maize-associated haplogroup and of the "pestiferous" haplotypes are correlated with the chronology of maize spread following its domestication. Overall, our results support a hypothesis positing that maize domestication favored corn leafhopper genotypes preadapted for exploiting maize so that they became pestiferous, and that with the geographical expansi on of maize farming, corn leafhopper colonized Z. diploperennis, a host exclusive to secluded habitats that serves as a refuge for archaic corn leafhopper genotypic diversity. Broadly, our results help explain the extents to which crop domestication and early spread may have mediated the emergence of today's agricultural pests.     

Phylogeography of Rhabdias spp. (Nematoda: Rhabdiasidae) collected from Bufo species in Honshu,Shikoku, and Kyushu,Japan including possible cryptic species

The genus Rhabdias Stiles & Hassall, 1905 comprises lung parasites of amphibians and reptiles worldwide. In Japan, 9 species have been recorded, including Rhabdias incerta Wilkie, 1930 which has been reported only in Bufo species. In this study, to assess the diversity of R. incerta, we performed molecular analyses of Rhabdias species sampled from three species/subspecies of Japanese toads namely Bufo japonicus, B. japonicus formosus, and B. torrenticola, collected in various regions of Honshu, Shikoku, and Kyushu, Japan. DNA sequence divergence was compared using mtDNA (COI) and nuclear DNA (28S) to identify possible cryptic species. Morphological analysis was performed through light microscopy and scanning electron microscopy (SEM). The results revealed that Bufo spp. serve as hosts for at least three Rhabdias species. Morphologically, most samples were identified as R. incerta but with a longer body and larger buccal cavity than originally described. Rhabdias incerta appears to be specific to the genus Bufo and is further subdivided into two or three phylogroups based on subspecies divisions and biogeography of their host. Some Rhabdias specimens collected in this study resemble R. tokyoensis Wilkie, 1930, parasitic in hosts from the order Caudata, which suggests host switching. Both molecular and morphological analyses suggested the presence of undescribed and cryptic Rhabdias species within toads collected in Japan. This study was the first to molecularly characterize Rhabdias species in Japan, including novel sequences of R. incerta and two undescribed species.     

Climate Change and Phenology: Empoasca fabae (Hemiptera: Cicadellidae) Migration and Severity of Impact

Climate change can benefit individual species, but when pest species are enhanced by warmer temperatures agricultural productivity may be placed at greater risk. We analyzed the effects of temperature anomaly on arrival date and infestation severity of potato leafhopper, Empoasca fabae Harris, a classic new world long distance migrant, and a significant pest in several agricultural crops. We compiled E. fabae arrival dates and infestation severity data at different states in USA from existing literature reviews and agricultural extension records from 1951–2012, and examined the influence of temperature anomalies at each target state or overwintering range on the date of arrival and severity of infestation. Average E. fabae arrival date at different states reveal a clear trend along the south-north axis, with earliest arrival closest to the overwintering range. E. fabae arrival has advanced by 10 days over the last 62 years. E. fabae arrived earlier in warmer years in relation to each target state level temperature anomaly (3.0 days / °C increase in temperature anomaly). Increased temperature had a significant and positive effect on the severity of infestation, and arrival date had a marginal negative effect on severity. These relationships suggest that continued warming could advance the time of E. fabae colonization and increase their impact on affected crops.     

A new loach, Cobitis shikokuensis (Teleostei: Cobitidae), from Shikoku Island, Japan

A new species of spinous loach, Cobitis shikokuensis, is described based on 297 specimens from Shikoku Island, Japan. The new species was formerly known as the Shikoku group of Cobitis takatsuensis. It can be distinguished from other species of Cobitis and closely related genera by a combination of the following characters: dorsal fin with 6 branched soft rays; anal fin with 5 branched soft rays; one brownish streak across eye from the tip of nose, no streak on cheek; a black spot smaller than eye diameter near the dorsal corner of the caudal fin base; 3–5 small brownish speckles on ventral side of caudal peduncle; high caudal peduncle with well-developed fleshy keels on dorsal and ventral side; a lamina circularis at base of dorsal part of pectoral fin absent; first branched soft ray of pectoral fin broad in males; pectoral soft rays widely branched from the approximate midpoint; last anal fin ray with 2 elements; interorbital width 11.2–17.1% of head length.     

846. TRICYRTIS MACRANTHA

Tricyrtis macrantha Maxim. (Liliaceae), an endemic species to Japan, is illustrated. It is distributed in a small area in Shikoku (Kochi Prefecture), and grows on limestone cliffs. Tricyrtis macrantha and its allied species, T. macranthopsis Masam., distributed in Kii Peninsula in western Honshu, and T. ishiiana (Kitag. & T. Koyama) Ohwi & Okuyama, distributed in Pacific coast side of central Honshu, comprise one natural group.     

Cover Caption

The green leafhopper, Empoasca vitis Göthe, is a pest of grapevine and other crops in Europe and Asia. The pair formation, here described for the first time, relies on vibrational communication and starts when the female replies to a male call and a duet is established. It then continues through two different behavioral stages: Location and Courtship. The acquired knowledge can be used to develop a vibrational mating disruption technique for pest population control (see pages 869–882). Photo by Umberto Salvagnin.     

What Happened to <Emphasis Type="Italic">Nezara viridula</Emphasis> (L.) in the Americas? Possible Reasons to Explain Populations Decline

Once abundant in the Americas, the southern green stink bug Nezara viridula (L.) has gradually declined in numbers. Until the 1990s, it was considered the main pest of major crops such as soybean, Glycine max (L.) Merrill, particularly in southern Brazil and southern USA, as well as Argentina, Uruguay, and other countries. In the past 15+ years, a dramatic population decrease was observed to the point of now being considered a secondary pest in these referred countries. In this article, we list and discuss possible reasons which explain the decline in N. viridula population in the Americas. These factors include the following: (1) the steady increase of herbicides used in no-tillage/multiple cropping systems affecting potential hosts such as weeds in crop fields and nearby natural vegetation; (2) the change in cultivation systems, mostly in the Neotropics, favoring other species more adapted to exploit crops in modern day agriculture; (3) competition among several species of stink bugs that colonize major crops; (4) the growing impact of several species of egg parasitoids, some of them laboratory reared and released in crop fields, and other natural enemies (parasitoids and predators); and (5) the impact of global climate change affecting its distribution and biology.     

Effects of an invading coccinellid on native coccinellids in an agricultural landscape

Seven native coccinellid species inhabited alfalfa, corn, and small grain fields in eastern South Dakota prior to invasion and establishment of Coccinella septempunctata L. Six species occurred in all crops, however, Adalia bipunctata (L.) occurred only in corn. The structure of native coccinellid communities differed significantly for years prior to compared with years after establishment of C. septempunctata in fields of the three agricultural crops. Differences in community structure were accounted for mainly by reduced abundance of two species, C. transversoguttata richardsoni Brown and Adalia bipunctata (L.). Annual abundance of C. transversoguttata richardsoni averaged 20–32 times lower during post-invasion years than in years prior to invasion, depending on crop; while annual abundance of A. bipunctata averaged 20 times lower in corn after invasion. Addition of C. septempunctata to the community did not result in a significant increase in total abundance of coccinellids in the crops. Coccinellid abundance in agricultural crops may be limited by the total abundance of prey or by the availability of other requisites in the landscape as a whole. Therefore, introduction of a new species, while resulting in reductions in native species populations, may not increase total coccinellid abundance, and may therefore have no net effect on biological control of aphid pests.