Variability in deer diet and plant vulnerability to browsing among forests with different establishment years of sika deer.

Increased ungulate browsing alters the composition of plant communities and modifies forest ecosystems worldwide. Ungulates alter their diet following changes in availability of plant species; however, we know little about how browse selection and plant community composition change with different stages of deer establishment. Here, we provide insight into this area of study by combining multiple approaches: comparison of the understory plant community, analysis of records of browsing damage, and DNA barcoding of sika deer feces at 22 sites in forests in northern Japan varying in when deer were first established. The coverage of vegetation and number of plant species were only lower at sites where deer were present for more than 20 years, while the difference in plant coverage among deer establishment years varied among plant species. Deer diet differed across establishment years, but was more affected by the site, thereby indicating that food selection by deer could change over several years after deer establishment. Plant life form and plant architecture explained the difference in plant coverage across establishment years, but large variability was observed in deer diet within the two categories. Integrating these results, we categorized 98 plant taxa into six groups that differed in vulnerability to deer browsing (degree of damage and coverage). The different responses to browsing among plant species inferred from this study could be a first step in predicting the short- and long-term responses of forest plant communities to deer browsing.

Environmentally triggered variability in the genetic variance-covariance of herbivory resistance of an exotic plant Solidago altissima.

The variability in the genetic variance-covariance (G-matrix) in plant resistance and its role in the evolution of invasive plants have been long overlooked. We conducted an additional analysis of the data of a reciprocal transplant experiment with tall goldenrod, Solidago altissima, in multiple garden sites within its native range (USA) and introduced range (Japan). We explored the differences in G-matrix of resistance to two types of foliar herbivores: (a) a lace bug that is native to the USA and recently introduced to Japan, (b) and other herbivorous insects in response to plant origins and environments. A negative genetic covariance was found between plant resistances to lace bugs and other herbivorous insects, in all combinations of garden locations and plant origins except for US plants planted in US gardens. The G-matrix of the resistance indices did not differ between US and Japanese plants either in US or Japanese gardens, while it differed between US and Japanese gardens in both US and Japanese plants. Our results suggested that the G-matrix of the plant resistance may have changed in response to novel environmental differences including herbivore communities and/or other biotic and abiotic factors in the introduced range. This may have revealed a hidden trade-off between resistances, masked by the environmental factors in the origin range. These results suggest that the stability of the genetic covariance during invasion, and the environmentally triggered variability in the G-matrices of plant resistance may help to protect the plant against multiple herbivore species without changing its genetic architecture and that this may lead to a rapid adaptation of resistance in exotic plants. Local environments of the plant also have a critical effect on plant resistance and should be considered in order to understand trait evolution in exotic plants.

Insect Herbivory Selects for Volatile-Mediated Plant-Plant Communication.

Plant volatile organic compounds (VOCs) are major vehicles of information transfer between organisms and mediate many ecological interactions [1-3]. Altering VOC emission in response to herbivore damage has been hypothesized to be adaptive, as it can deter subsequent herbivores [4], attract natural enemies of herbivores [5], or transmit information about attacks between distant parts of the same plant [6-9]. Neighboring plants may also respond to these VOC cues by priming their own defenses against oncoming herbivory, thereby reducing future damage [10-12]. However, under which conditions such information sharing provides fitness benefits to emitter plants, and, therefore, whether selection by herbivores affects the evolution of such VOC signaling, is still unclear [13]. Here, we test the predictions of two alternative hypotheses, the kin selection and mutual benefits hypotheses [14], to uncover the selective environment that may favor information sharing in plants. Measuring the response to natural selection in Solidago altissima, we found strong effects of herbivory on the way plants communicated with neighbors. Plants from populations that experienced selection by insect herbivory induced resistance in all neighboring conspecifics by airborne cues, whereas those from populations experiencing herbivore exclusion induced resistance only in neighbors of the same genotype. Furthermore, the information-sharing plants converged on a common, airborne VOC signal upon damage. We demonstrate that herbivory can drive the evolution of plant-plant communication via induction of airborne cues and suggest plants as a model system for understanding information sharing and communication among organisms in general.

Fauna of nocturnal moth species collected in a semi-natural grassland at Kanpu-zan in northern Japan.

Semi-natural grasslands, which house species-rich ecosystems, have rapidly declined since the twentieth century due to land-use practices, such as agricultural intensification and abandonment. Owing to their diversity and known habitat associations, nocturnal moths are considered as one of the most suitable organisms to be studied for assessing the dynamics of species composition as a result of changes in landscape management of semi-natural grasslands. The present study provides the foremost description of nocturnal moth fauna of the semi-natural grassland at Kanpu-zan, northern Japan. Moth population data from 1987 were compared to the data collected in 2018 to evaluate the impact of decline in grasslands on species-richness. During the field sampling in 2018, a total of 226 nocturnal moth species were detected, which was nearly two-thirds of the number of species recorded in 1987, i.e. 396 species. The values obtained in 2018 were found to be nearly constant for different sites. For both periods, it was evident that moth fauna in Kanpu-zan mainly consisted of species that relied on woody plants. Amongst the species which were only recorded in 1987, 107 species were generalists that fed on plants that are commonly distributed in Kanpu-zan. No moth species were recorded that depended upon endangered or extinct plant food sources. Thus, it is unlikely that the decline in the number of moth species in Kanpu-zan was due to the loss in plant food sources. Our results suggest that environmental factors other than food plants may have caused decline and changes in nocturnal moth fauna. More studies on various organism fauna are needed for understanding the conservation of semi-natural grassland, considering that the loss of semi-natural grasslands is one of the major threats to biodiversity.

Parallel environmental factors drive variation in insect density and plant resistance in the native and invaded ranges.

Geographic variation in the traits of a species is shaped by variation in abiotic conditions, biotic interactions, and evolutionary history of its interactions with other species. We studied the geographic variation in the density of the lace bug, Corythucha marmorata, and the resistance of tall goldenrod Solidago altissima to the lace bug herbivory in their native range in the United States and invaded range in Japan. We conducted field surveys and reciprocal transplant experiments to examine what abiotic and biotic factors influence variation in lace bug density, and what ecological and evolutionary factors predict the resistance of the host plant between and within the native and invaded ranges. Lace bug density was higher throughout the invaded range than in the native range, higher in populations with warmer climates, and negatively affected by foliage damage by other insects in both ranges. The higher lace bug density in warmer climates was explained by the shorter developmental time of the lace bugs at higher temperatures. The resistance of S. altissima to lace bugs was higher in populations with lace bugs compared to populations without lace bugs in both native and invaded ranges, indicating that the evolutionary history of the interaction with the lace bugs was responsible for the variation in S. altissima resistance in both ranges. The present study revealed that abiotic and biotic factors, including temperature and other herbivorous insects, can drive the geographic variation in lace bug density, which in turn selects for variation in plant resistance in both in the native and invaded ranges. We conclude that the novel combination of factors such as higher temperature and lower number of other herbivorous insects is responsible for the higher lace bug density in the invaded range than in the native range.

Multiple and mass introductions from limited origins: genetic diversity and structure of Solidago altissima in the native and invaded range.

Understanding the origins and diversity of invasive species can reveal introduction and invasion pathways, and inform an effective management of invasive species. Tall goldenrod, Solidago altissima, is a herbaceous perennial plant native to North America and it has become a widespread invasive weed in East Asian countries. We used microsatellite and chloroplast DNA markers to obtain information on neutral processes and on genetic diversity in native and invaded populations of S. altissima and to infer how it invaded and spread in Japan. We found that introduced (n = 12) and native (n = 20) populations had similar levels of genetic diversity at nuclear SSR loci. Genetic structure analysis indicated that at least two independent colonization events gave rise to current S. altissima populations in Japan. The majority (68%) of the Japanese S. altissima were genetically similar and likely shared a common origin from a single or a small number of populations from the southern USA populations, while the populations in Hokkaido were suggested to arise from a different source. Our results suggest that multiple and mass introductions have contributed to the persistence and rapid adaptation of S. altissima promoting its widespread establishment throughout Japan.

Isolation and characterization of microsatellite loci in the invasive herb Solidago altissima (Asteraceae).

PREMISE OF THE STUDY: Microsatellite markers were developed in the perennial herb Solidago altissima from populations within its introduced range in Japan to assess its population structure and to facilitate tracking of invasion expansion. • METHODS AND RESULTS: Using 454 pyrosequencing, 16 microsatellite primer sets were developed for S. altissima. The primer sets were tested on 70 individuals sampled from three populations in Japan. The primers amplified di- and trinucleotide repeats with five to 25 alleles per locus, and the expected heterozygosity ranged from 0.46 to 0.92. • CONCLUSIONS: These results indicate the utility of primers in S. altissima for future research on a wide range of applications, including tracking of invasion dynamics and investigating population genetics of the species.