DOI: 10.5061/dryad.ns1rn8q3k
title: Unilateral exclusion of three leaf beetle species due resource competition for oviposition sites

description: Resource competition is a key factor in shaping community structure.
 However, exclusion due to resource competition likely occurs only in
 immobile herbivorous insects such as scale insects, because free-living
 insects mitigate competition by moving to alternative resources. Given
 that eggs are immobile, their placement is critical for larval survival
 and development, especially in terms of optimal resource consumption,
 making oviposition site selection a crucial factor for females. Therefore,
 competition for oviposition sites may lead to exclusion if resources are
 scarce. This study examined the competitive relationships among three leaf
 beetle species (Coleoptera: Chrysomelidae): Aspidimorpha difformis,
 Aspidimorpha transparipennis, and Laccoptera nepalensis. Competitive
 dynamics were assessed using field surveys, experiments, and simulations.
 Results revealed that habitats for both Aspidimorpha species were less
 than half the size of L. nepalensis habitats. Additionally, both
 Aspidimorpha species avoided oviposition on leaves with the oothecae of
 other species, with A. difformis showing a reduction of up to 0% of
 ootheca number. Simulation results indicated that only A. difformis, which
 experienced oviposition site reductions due to other species, faced
 certain extinction under conditions mimicking natural condition of
 mortality. Our findings suggest that even free-living herbivorous insect
 experiences unilateral exclusion due to resource competition for
 oviposition sites, as observed in the field.Thus, based on our findings,
 even free-living herbivorous insects can be excluded due to interference
 competition during their immobile stage, such as eggs.
description: # Unilateral exclusion of three leaf beetle species due resource
 competition for oviposition sites Data for habitats, fertility, and
 extinction probabilities of three leaf beetle (Coleoptera: Chrysomelidae)
 species, *Aspidimorpha difformis*, *Aspidimorpha transparipennis*, and
 *Laccoptera nepalensis*. All surveys and experiments were conducted in
 Shizuoka Prefecture, Japan. This manuscript has been accepted for
 publication (02/4/2025): Nomura N. and Kasai A. Unilateral exclusion of
 three leaf beetle species due resource competition for oviposition sites.
 Ecological Entomology (accepted). Contact Natsuki Nomura
 ([nomura.natsuki.y5@s.gifu-u.ac.jp](mailto:nomura.natsuki.y5@s.gifu-u.ac.jp)) with any questions. ## Data-specific information These terms are common across multiple CSV files and are explained in this section. 1. Adifformis, Atransparipennis, and Lnepalensis are leaf beetles. This term has been mainly used in columns which named to "Species" or " Replaced_or_coexisted_species”. 2. “Control,” “Mix,” “Intraspecific_replacement,” and “Intraspecific_replacement” are names of treatments in experiment. “control” and “mix” refer to experiments with one and two females of leaf beetles in one experimental equipment, respectively. ”Intraspecific_replacement” indicates that one female was first tested in one experimental setup and then replaced with one female from the other species. In “Intraspecific_replacement,” the method is the same as interspecific replacement, changed females are same species. This term has been mainly used in columns which named to "Treatments”. 3.  In the “control” treatment, there is no replaced or altered species as only one female is tested, therefore, when the treatment is “Control,” “Replaced_or_coexisted_species” was written as “Blank.” Data files include: ### Calculated\_fertilyties.csv A raw data of realized fertility which was calculated 10,000 times using daily egg counts and days of egg laying. Its average and standard deviation were used to population viability analysis. The first cell in each column contains the title, followed by the calculated value. Each title comprises the 1) species to be evaluated, 2) coexisting or replaced insects with 1), and 3) names of treatments. For example, “Adifformis_Atransparipennis_Interspecific_replacement” is isolated to "Adifformis,","Atransparipennis", and Interspecific \_replacement"; thus, it means that potential fecundity of *A. difformis* which was replaced from *A. transparipennis*. ### Egg\_in\_ootheca\_number.csv A raw data of eggs in ootheca of three leaf beetle species with different treatments. The meaning of each column and its title are as follows: * Species (column 1): Name of examined leaf beetle * Replaced_or_coexisted_species (column 2): Name of replaced or coexisted leaf beetle * Treatments (column 3): Name of treatments * Egg_number_in_ootheca (column 4): Number of eggs in ootheca of the species shown in column 1 ### Ootheca\_number\_summarise.csv Raw data of total number of ootheca for each experimental equipment. The meaning of each column and its title are as follows: * Used (column 1): Used to analyses (1 = used, 0 = unused) * No (column 2): The identification number of each experimental equipment * Species (column 3): Name of examined leaf beetle * Replaced_or_coexisted_species (column 4): Name of replaced or coexisted leaf beetle * Treatments (column 5): Name of treatments * Ootheca_number (column 6): Total number of ootheca of the species shown in column 4 * Leaf_number (colmn7): The leaf numbers of prepared host plants * Note (colmn 8): Annotations are written (Prepared insect died = dead, Annotation is none = blank cell) ### Ootheca\_number\_whole.csv Raw data on the number of ootheca per leaf for the three leaf beetle species. This was used to analyze oviposition site preference using a generalized linear mixed model, as well as to compare the number of oothecae per leaf and the proportions of leaves as oviposition sites among the three leaf beetle species. The meaning of each column and its title are as follows: * Data (column 1): Date of data collection * Case_No (column 2): The identification number of each experimental equipment * Leaf_No (column 3): The identification number of each host plant leaf * Species (column 4): Name of examined leaf beetle * Replaced_or_coexisted_species (column 5): Name of replaced or coexisted leaf beetle * Treatments (column 6): Name of treatment * Ootheca_number (column 7): Number of ootheca of the species shown in column 4 * Egg_number_of_pre_replaced_species (column 8): If column 6 is ”Intraspecific replacement” and ”Interspecific replacement”, number of ootheca of the species shown in column 5, elsewhere, blank cell * Leaf_hardness_Newton (column 9): Hardness of leaf lamina * Percent_of_feeding_scars (column 10): % of area of feeding scars per leaves * Used_for_glmm (column11): If column 6 is ”Intraspecific replacement” and ”Interspecific replacement,” the number of ootheca of the species shown in column 5, elsewhere, is a blank cell (used = 1, unused = 0) * Note (column 12): Annotations are written (Prepared insect died = dead, Annotation is none = blank cell) ### Population\_dynamics\_2019\_to\_2020.csv Population dynamics of three species of leaf beetles from 2019 to 2020. This was used to calculate the mortality rates of the three leaf beetle species under field conditions. The meaning of each column and its title are as follows: * Area name (column 1): Name of survey sites * Year (column 2): Survey year * Month (column 3): Survey month * \[Developmental period]_[Species] (column 4-11): Observed number of leaf beetles (e.g. column 4 and row2 shows that five ootheca of *A. difformis* were observed at May, 2019) #### Annotation Host plant communities which all of the three leaf beetle species inhabited were nothing during survey period. Thus, *A. trasnparipennis* in Oya and *A. difformis* in Abekawa water park are blank cell respectively. On the other hand, there are no data from the survey at Abekawa Water Park in July 2020 because the survey site was flooded. Therefore, row 54 has been entered as an NA to indicate that there are no data. ### Survey\_whole.csv Raw data on the presence/absence of ootheca, larvae, pupae, and adults of the three leaf beetle species in 2021. All survey sites were host plant communities of three leaf beetle species in Shizuoka City, Shizuoka Prefecture, Japan. The meaning of each column and its title are as follows: * Id: individual numbers of each survey sites for data management * Species: name of surveyed leaf beetle * Area_name: Name of survey sites * Latitude and Longitude (column4-5): Location information of each survey site * \[Developmental period] \_[Month] (column6-33): Monthly results of each developmental stage (present = 1, absent = 0) * Eggscore-Adultsscore (column 34-37): Frequency of occurrence of each developmental stage at survey sites (&gt;0 = present, 0 = absent) * Habitation (column 38): Results of determining habitat, temporary occurrence site, and non-habitat of leaf beetles at each survey site (1 = habitat, 0.5 = temporary occurrence site, 0 = non-habitat) #### Annotation Habitats of three leaf beetle species (column 38) were determined using results in column 34-37. (All of cell are &gt;1 = habitat, one or more cell is &gt;1 = temporary occurrence site, all of cell are 0 = non-habitat.). ### Vortex\_PVA\_results.csv Raw data of the population viability analysis using VORTEX software. All output results are shown in this CSV file, but only extinction probabilities of metapopulations were used in our study. The meaning of each column and its title are as follows: * Species (column 1): Name of leaf beetles - Replaced_or_coexisted_species (column 2): Name of replaced or coexisted leaf beetle - Treatments (column 3): Name of treatments - Scen.name (column 4): Name of scenarios - Pop.name (column 5): Name of population (Metapopulation is combination of Population 1 and 2.) - Year (column 6): passed years - PExtant (column 7): Average of probabilities of extant - SE.PExtant. (column 8): Standard error of extant probabilities - PExtinct (column 9): Average of probabilities of extinct - SE.PExtinct. (column 10): Standard error of extinction probabilities - det.r (column 11): Deterministic intrinsic per capita population growth rate - stoch.r (column 12): Average of stochastic intrinsic per capita population growth rate - SE.r. (column 13): Standard error of stochastic intrinsic per capita population growth rate - SD.r. (column 14): S tandard deviation of stochastic intrinsic per capita population growth rate - N.extant (column 15): Average population size excluding scenarios excluding scenarios in which extinction occurred - SE.N.extant. (column 16): Standard error of population size excluding scenarios in which extinction occurred - SD.N.extant. (column 17): Standard deviation of population size excluding scenarios in which extinction occurred - N.all (column 18): Average of population size including scenarios in which extinction occurred - SE.N.all. (column 19): Standard error of population size including scenarios in which extinction occurred - SD.N.all. (column 20): Standard deviation of population size including scenarios in which extinction occurred - GeneDiv (column 21): Average of expected heterozygosity - SE.GD. (column 22): Standard error of expected heterozygosity - SD.GD. (column 23): Standard deviation of expected heterozygosity - Inbreed (column 24): Average of observed heterozygosity remaining in the extant populations - SE.Inbr. (column 25): Standard error of observed heterozygosity remaining in the extant populations - SD.Inbr. (column 26): Standard deviation of observed heterozygosity remaining in the extant populations - nAlleles (column 27): Average of alleles remaining within extant population - SE.nAlleles. (column 28): Standard error of alleles remaining within extant population - SD.nAlleles. (column 29): Standard deviance of alleles remaining within extant population - nLethals (column 30): Average of lethal alleles remaining within extant populations - SE.nLethals. (column 31): Standard error of lethal alleles remaining within extant populations - SD.nLethals. (column 32): Standard deviance of lethal alleles remaining within extant populations - TE (column 33): The mean time to first population extinction - SE.TE. (column 34): Standard error of time to first population extinction - SD.TE. (column 35): Standard deviance of time to first population extinction - medianTE (column 36): The median time to extinction (Extinction in &gt; 500 scenarios) - SD.PExtant. (column 37): Standard deviance of probabilities of extant - SD.PExtinct. (column 38): Standard deviance of probabilities of extinct #### Annotation Scen. name (column 4) means that these simulations were performed using different parameters. In the "baseline model", juvenile mortality was calculated based on field survey results, and the annual probability of a population-depleting catastrophes was fixed at 1%. In scenario name of “[value]%_of_catastrophes”, probability of a population-depleting catastrophes was changed from the baseline model according to values. In scenario name of “[value]_of_mortalty”, juvenile mortality was changed from the baseline model according to value.