Andrea Lucky’s research while affiliated with University of Florida and other places

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Publications (82)


Defining the decline: a glossary relevant to insect decline
  • Article
  • Full-text available

May 2025

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89 Reads

Journal of Insect Science

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Gagandeep Brar

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Erin Cadwalader

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[...]

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Ross Winton

Insects are declining in abundance and species richness, globally. This has broad implications for the ecology of our planet, many of which we are only beginning to understand. Comprehensive, large-scale efforts are urgently needed to quantify and mitigate insect biodiversity loss. Because there is broad interest in this topic from a range of scientists, policymakers, and the general public, we posit that such endeavors will be most effective with precise and standardized terms. The Entomological Society of America is the world's largest association of professional entomologists and is ideally positioned to lead the way on this front. We provide here a glossary of definitions for biodiversity loss terminology. This can be used to enhance and clarify communication among entomologists and others with an interest in addressing the multiple overlapping research, policy, and outreach challenges surrounding this urgent issue.

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Distribution of attitude toward insects score in the EntoEdu v2 sample. A Overall distribution of attitude score. B Attitude score by country affiliation; scores and distribution were similar for Czech and US students. C Attitude score by study domain; Biology student scores were higher than other domains, with significant differences from students in Humanities & Social Sciences and in Technology & Engineering (no significant difference between Biology and Non-Biological Sciences)
Distribution of knowledge about insects score in the EntoEdu v2 sample. A Overall distribution of knowledge score. B Knowledge score by country affiliation; Czech student scores were higher than US student scores. C Knowledge score by study domain. Biology students scores were higher than other domains, with significant differences from students in Humanities & Social Sciences and in Technology & Engineering (no significant difference between Biology and Non-Biological Sciences)
Distribution of knowledge about insects score by age/grade level in Czechia. First-year university student scores were higher than those of pupils in the final year of middle school (i.e., US 9th grade equivalent)
Classical Test Theory analysis results for the 50 items in the knowledge section, visually grouped by the four entomology knowledge categories used in this instrument (modified from Pearson et al., 2007): (1) Life History, (2) Concepts, (3) Identification, and (4) Techniques. Red bars represent item difficulty (higher bar indicates greater proportion of correct responses); blue bars indicate Upper–Lower Index (ULI) discrimination ability of each item (higher bar indicates better discrimination ability). The black line represents the commonly used acceptability threshold of 20% used to identify potentially problematic items
The item-person (Wright) map, based on a 2 parameter logistic Item Response Theory model, provides a classification of respondents’ ability on the left (i.e., Latent trait: entomological knowledge), where a higher position indicates an individual with greater knowledge. The difficulty of each numbered instrument item is plotted on the right, with the most difficult items at the top and the least difficult at the bottom

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Quantifying ento-literacy: development and validation of an international insect-focused attitude and knowledge survey instrument

February 2025

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72 Reads

International Journal of STEM Education

Background In an era of precipitous insect declines, effective entomology education is especially needed to support firsthand knowledge of nature. Understanding what students know and feel about insects is instrumental to teaching and curriculum development. This study describes the development and validation of a new survey instrument, EntoEdu, measuring ‘entomology literacy’, based on attitude and knowledge, in a cross-cultural context. For the survey validation we use data collected from students in Czechia (CZ), a country known for its entomophilia, and the United States of America (US) to demonstrate the utility of this survey and to address the questions: how do entomology attitude and knowledge differ across national affiliation and study domain, and how are entomology attitude and knowledge correlated in the context of these differences? Results Based on responses from 635 first-year college students, we demonstrate high reliability and evidence of validity of the EntoEdu instrument. Factor analysis supports five independent attitudinal categories within the instrument: Intention to Engage with Insects, Attitude toward Behavior, Control Belief, Hobby, and Disgust. In this study population, average attitude scores did not differ with nationality, whereas knowledge scores were higher in CZ than in the US. In both countries, attitude and knowledge scores were higher among biology students than those in other study domains, and attitude and knowledge were positively correlated. Conclusions The EntoEdu instrument, based on globally recognizable insect taxa, ecology, and behavior, has been developed for broad utility in assessing attitudes toward and knowledge of insects at the post-secondary level, with potential for use at both lower (K-12) and higher (advanced university) levels. The instrument is presented here in two language versions and can be translated into additional languages for comparison of results across test populations in additional countries. In our initial test population we find attitude and knowledge to be correlated, both of which are influenced by nationality, with Czechs more knowledgeable about insects than their US counterparts. We anticipate that this instrument will facilitate entomology assessment to help tailor biology education programs to students’ actual, rather than assumed, entomology knowledge and attitudes, and for tracking change over time.


Nylanderia of the World, Part IV: Taxonomic contributions to the American Clade I of New World Nylanderia (Hymenoptera: Formicidae)

January 2025

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208 Reads

Zootaxa

Globally, potentially hundreds of Nylanderia species remain undescribed, hidden within several broadly distributed complexes of morphologically cryptic species. By integrating phylogenomics, geography, and morphology, we describe eight new Nylanderia species from southern Mexico and Mesoamerica, increasing the total number of known species in the genus to 131. In the Americas, Nylanderia is divided into two distantly related clades: American Clade I (AC1) and American Clade II (AC2). Within AC1, Nylanderia austroccidua (Trager) was originally described as a widespread and morphologically variable species distributed from Utah to Costa Rica. This species was diagnosed by a slight concavity in the anterior face of the pronotum and varying degrees of fine cuticular microsculpturing across the body that causes blue cuticular iridescence under microscopic examination. Using Ultraconserved Elements (UCEs) for molecular phylogenetic analysis, we found that taxa matching the original description of N. austroccidua are paraphyletic with respect to Nearctic Nylanderia species. We also found that AC1 includes a Neotropical subclade extending into Mesoamerica, the distribution of which overlaps with AC2, which is exclusively Neotropical. Along with an updated description of N. austroccidua, we also describe the following new species belonging to clade AC1: N. breviscapa, sp. nov., N. contraria, sp. nov., N. lazulina, sp. nov., N. luceata, sp. nov., N. mendax sp. nov., N. mosaica sp. nov., N. polita sp. nov., and N. usul, sp. nov. A dichotomous key and images of the worker caste of these species are included and, where available, images of queens and males are provided.


Evaluating UCE Data Adequacy and Integrating Uncertainty in a Comprehensive Phylogeny of Ants

January 2025

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173 Reads

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2 Citations

Systematic Biology

While some relationships in phylogenomic studies have remained stable since the Sanger sequencing era, many challenging nodes remain, even with genome-scale data. Incongruence or lack of resolution in the phylogenomic era is frequently attributed to inadequate data modeling and analytical issues that lead to systematic biases. However, few studies investigate the potential for random error or establish expectations for the level of resolution achievable with a given empirical dataset and integrate uncertainties across methods when faced with conflicting results. Ants are the most species-rich lineage of social insects and one of the most ecologically important terrestrial animals. Consequently, ants have garnered significant research attention, including their systematics. Despite this, there has been no comprehensive genus-level phylogeny of the ants inferred using genomic data that thoroughly evaluates both signal strength and incongruence. In this study, we provide insight into and quantify uncertainty across the ant tree of life by utilizing the most taxonomically comprehensive Ultraconserved Elements dataset of ants to date, including 277 (81%) of recognized ant genera from all 16 extant subfamilies, and representing over 98% of described species. We use simulations to establish expectations for resolution, identify branches with less-than-expected concordance, and dissect the effects of data and model selection on recalcitrant nodes. Simulations show that hundreds of loci are needed to resolve recalcitrant nodes on our genus-level ant phylogeny. This demonstrates the continued role of random error in phylogenomic studies. Our analyses provide a comprehensive picture of support and incongruence across the ant phylogeny, while offering a more nuanced depiction of uncertainty and significantly expanding generic sampling. We use a consensus approach to integrate uncertainty across different analyses and find that assumptions about root age exert substantial influence on divergence dating. Our results suggest that advancing the understanding of ant phylogeny will require not only more data but also more refined phylogenetic models. We also provide a workflow for identifying under-supported nodes in concatenation analyses, outline a pragmatic way to reconcile conflicting results in phylogenomics, and introduce a user-friendly locus selection tool for divergence dating.


Fig. 5. Maximum likelihood phylogeny of American Nylanderia Clade I reconstructed using the 90% complete SWSC partitioned matrix. The inset on the left shows the position of this major clade in the full phylogeny. Nodal support (SH-aLRT/UFBoot) is displayed only for nodes with <95% support. Silhouettes on the right represent gonopod shapes of species (to scale).
Fig. 7. Nylanderia fulva complex phylogenies and gonopods: A) ML phylogeny reconstructed using IQ-TREE 2 on the 90% complete SWSC partitioned UCE matrix, with open red circles indicating nodes that were not recovered with strong support by any measure (SH-aLRT and UFBoot <95%); B) multispecies coalescent phylogeny reconstructed from 1,555 gene trees using ASTRAL-III on the 90% complete UCE matrix, with red dots indicating nodes that were not strongly recovered with strong support (Local Posterior Probability <95%); and C) 3D models of N. fulva COI clades I-III and N. pubens gonopods rendered from nano-CT scans. See Supplementary Table S4 for collection and scan data for each CT-scanned specimen. Taxa in blue represent N. fulva clade I, those in yellow represent N. fulva clade II, those in red represent N. fulva clade III, and those in purple represent N. pubens.
Fig. 8. Two-dimensional geometric morphometric analysis of gonopods of 16 Neotropical Nylanderia species: A) boxplot comparing centroid sizes; and B) gonopod morphospace using the 2 highest scoring principal components (PC01 and PC02), with TPS grids aligned along the x and y axes.
Fig. 10. Deformation grids for the first 3 principal components (PC01-PC03) from three-dimensional geometric morphometric analysis of N. fulva complex gonopods. Grids on the left represent dorsal view, and grids on the right represent endal view. For each principal component, the top grids represent the minimum value, and the bottom grids represent the maximum value. Lollipop graphs on the right depict the amount of shape variation associated with each landmark for each principal component.
Ultraconserved element sequencing statistics
“Picking up signals” in male genital morphospace and integrating phylogenomics to delimit Neotropical Nylanderia Emery species (Hymenoptera: Formicidae)

October 2024

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281 Reads

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1 Citation

Insect Systematics and Diversity

Genital morphology, a cornerstone in taxonomy that predates Linnaeus’s Systema Naturae, is vital for species delimitation. However, the widely accepted paradigm that genitalia are taxonomically informative lacks robust testing between closely related species, and supporting evidence is often limited to taxonomic literature in which genitalia are assumed a priori to be species-specific. The cosmopolitan ant genus Nylanderia Emery includes 123 described species, with most in the Neotropics still undescribed. Workers are often morphologically cryptic, and males are rare in collections but required for morphological delimitation. Using Ultraconserved Elements (UCEs) from 236 samples, including 53 Neotropical Nylanderia species, we reconstructed a phylogenetic framework to compare the genitalia (gonopods) of males collected alongside workers. We used geometric morphometrics on images of slide-mounted genitalia from 16 species and nano-CT scans of Nylanderia fulva (Mayr) and Nylanderia pubens (Forel) genital capsules and interpreted results considering phylogeny under maximum likelihood and the multispecies coalescent. We found strong morphological and molecular support for 2 distantly related American clades, identifiable by gonopod shape, with significant differences observed among most species. Three previously reported COI clades of N. fulva were not supported as monophyletic, nor were their gonopods significantly different. However, N. pubens was supported as distinct by all phylogenetic and 3DGM results. Our findings emphasize the importance of male genitalia for delimiting species boundaries and revising Neotropical Nylanderia. Given their importance, particularly in morphologically cryptic taxa, we recommend a greater focus on linking male and worker phenotypes, which can be facilitated through comprehensive nest series collection.


Evaluating UCE data adequacy and integrating uncertainty in a comprehensive phylogeny of ants

July 2024

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410 Reads

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3 Citations

While some relationships in phylogenomic studies have remained stable since the era of Sanger sequencing, many challenging nodes elude resolution, even with genome-scale data. As early studies grappled with random error and insufficient information, incongruence or lack of resolution in phylogenomics is generally associated with inadequate modeling of biological phenomena combined with analytical issues leading to systematic biases. Few phylogenomic studies, however, explore the potential for random error or establish an expectation of what level of resolution should be expected from a given empirical dataset. In presenting incongruent results, phylogeneticists face a choice of providing a diverse array of results from different approaches or a single preferred tree, with few attempting to integrate uncertainties across methods. Recent phylogenetic work has uncovered many well-supported and often novel relationships, as well as more contentious findings, across the phylogeny of ants. Ants are the most species-rich lineage of social insects and among the most ecologically important terrestrial animals. As a result, they have attracted much research, including regarding systematics. To date, however, there has been no comprehensive genus-level phylogeny of the ants inferred using genomic data combined with an effort to evaluate signal and incongruence throughout. Here we provide deeper insight into and quantify uncertainty across the ant tree of life. We accomplish this with the most taxonomically comprehensive Ultraconserved Elements dataset to date, including 277 (81%) of recognized ant genera from all 16 extant subfamilies, representing over 98% of described species-level diversity. We use simulations to establish expectations for resolution, identify branches with less-than-expected concordance, and dissect the effects of data and model selection on recalcitrant nodes. We also construct a consensus tree integrating uncertainty from multiple analyses. Simulations show that hundreds of loci are needed to resolve recalcitrant nodes on our genus-level ant phylogeny, even under a best-case scenario of known model parameters and without systematic bias. This demonstrates that random error continues to play a role in phylogenomics. Our analyses provide a comprehensive picture of support and incongruence across the ant phylogeny, and our consensus topology is congruent with a recent phylogenomic study on the subfamily-level, while rendering a more realistic picture of uncertainty and significantly expanding generic sampling. We use this topology for divergence dating and find that assumptions about root age have significant impact on the dates inferred. Our results suggest that improved understanding of ant phylogeny will require both more data and better phylogenetic models. We also provide a workflow to identify under-supported nodes in concatenation analyses, outline a pragmatic way to reconcile conflicting results in phylogenomics, and introduce a user-friendly locus selection tool for divergence dating.


Distribution of S. invicta, D. bureni, P. obscurithorax, and C. venustula in Florida. S. invicta (shaded light red, data from USDA-APHIS-PPQ https://www.aphis.usda.gov/plant_health/plant_pest_info/fireants/downloads/federal-imported-fire-ant-quarantine.pdf), D. bureni (gray stripes, data from Deyrup (2016)), P. obscurithorax (filled circles, data from GBIF https://doi.org/10.15468/dl.bc4w7h) and C. venustula (open circles, data from GBIF https://doi.org/10.15468/dl.npp4zj). The black star shows the location of the collection site for this study. The distribution of the introduced ants, C. emeryi and B. obscurior is in Supplementary Material 2.
Ant community composition in a citrus grove reveals eastern expansion in Florida of the South American big-headed ant (Hymenoptera: Formicidae)

March 2024

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73 Reads

Florida Entomologist

Florida has become a worldwide hotspot for introduced and invasive ants (Hymenoptera: Formicidae). Furthermore, studies have shown that non-native ants support other invasive insects in Florida, such as the Asian citrus psyllid, Diaphorina citri Kuwayama (Hemiptera: Liviidae), which is the vector of Candidatus Liberibacter asiaticus, the causal pathogen of citrus greening, the worst citrus disease in the world. The Asian citrus psyllid establishes beneficial interactions with invasive, introduced, and native ants. In this study, we described the ant diversity in a citrus grove in south Florida as a first step to assess ant-Asian citrus psyllid interactions for future studies. During the summer of 2018, 25 pitfall traps were set in a citrus grove. The red imported fire ant, Solenopsis invicta (Buren), was present in all the traps (100 %), and 19 traps (76 %) had Dorymyrmex bureni (Trager), the pyramid ant, which is native to the southeast United States. Two introduced ants, Cardiocondyla emeryi (Forel) and Brachymyrmex obscurior (Forel), were found in 15 (60 %) and 11 (44 %) traps, respectively. The South American big-headed ant, Pheidole obscurithorax (Naves), was found in 14 traps (56 %). This introduced ant has been found to co-exist with S. invicta, and to our knowledge, this is the first report of the species in this eastern part of peninsular Florida. Finally, Cardiocondyla venustula (Wheeler), an introduced ant from Africa, was found in one of the traps. Although, this ant is found in south Florida, it seems uncommon, and this is its first report in this county. Our study highlights the continued spread of introduced and invasive ants in Florida.


Diel Patterns Hold Promise as an Ecological Trait for Ants

February 2024

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86 Reads

Sociobiology

Ecological traits have flourished in insect-based studies, resulting in a substantial and growing list of measurable traits. One trait that will likely become more attractive as data quality and curation improve are the diel patterns of insect activities. Diel patterns in ants can help better understand vital ecological processes such as competition and invasion biology. Because diel activity has the potential to be an informative trait in ants, we assessed the diel designations of foraging ants across the literature to quantify and assess the variation and sampling extent of this particular trait. We collected diel designations from 104 peer-reviewed scientific articles and quantified these data across important and documented ecological traits. We found that a disproportionate amount of solitary foraging ants were primarily diurnal foragers relative to ants that cooperatively forage. Our data show that diel patterns in foraging vary widely within and across ant genera. Importantly, we highlight the undersampling of this crucial ecological trait, which currently limits its utility. Our efforts highlight the importance of assessing an ecologically important trait’s landscape of reported data.



Fig. 2: Photomicrograph of Leptomyrmex neotropicus replete specimen BALDR-0155 preserved in Miocene-age Dominican amber. Lateral view with distended gaster visible. Scale = 1 mm.
Fig. 3: X-ray computed tomography images of Leptomyrmex neotropicus. (A) Lateral view of replete L. neotropicus worker specimen BALDR 0155. (B) Lateral view of a non-replete L. neotropicus worker (AMNHDR-13-85 modified from Barden & al. 2017). (C) Z-stack cross section of specimen BALDR 0155, head and gaster denoted by dotted line in sub-panel A; h c = head cuticle, h v = voidspace of head, m = amber matrix, g c = gaster cuticle, g v = voidspace of gaster. Scale = 1 mm.
The first fossil replete ant worker establishes living food storage in the Eocene

May 2023

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202 Reads

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1 Citation

Myrmecological News

Worker specialization extends the behavioral and ecological repertoire of ant colonies. Specialization may relate to colony defense, brood care, foraging, and, in some taxa, storage. Replete workers swell the crop and gaster to store liquid food, which can be accessed by other colony members through trophallaxis. This storage ability, known as repletism, has independently evolved across several ant lineages, but the temporal history of this trait has not yet been investigated. Here, we describe the first fossil replete in the extinct species Leptomyrmex neotropicus Baroni Urbani, 1980 preserved in Miocene-age Dominican amber. Together with new evidence of repletism in L. neotropicus' extant sister species, Lep-tomyrmex relictus Boudinot & al., 2016, we reconstruct the pattern of acquisition and descent in this storage-linked trait. Our ancestral-state reconstruction suggests that Leptomyrmex acquired replete workers in the Eocene and may therefore represent the earliest instance of so-called "honeypot" ants among all known ants, both living and extinct.


Citations (45)


... 3 .Explore: Exploratory data analysis is evident in multiple studies. Borowiec et al. [27] uses simulations to explore and establish expectations for phylogenetic resolution in ant species. Marshall et al. [28] analyzes regional changes in cortical thickness, sulcal depth, surface area, and volume to explore adolescent brain development. ...

Reference:

Feasibility Analysis of Data Science Methodologies in Architectural Design
Evaluating UCE Data Adequacy and Integrating Uncertainty in a Comprehensive Phylogeny of Ants
  • Citing Article
  • January 2025

Systematic Biology

... as sister to all other Basiceros with both morphology only and combined morphological and molecular datasets ( figure 5, electronic supplementary material, figures S2 and S4-S9). The topologies of our reconstructions largely mirror results by Probst et al. [26], even with our inclusion of additional in-and outgroup taxa, while some relationships outside of Basiceros are distinct from Hanisch et al. [48,61]. Relationships among the six genera were recovered with variable support values, particularly the clade containing Rhopalothrix, Protalaridris and Octostruma (electronic supplementary material, figures S3-S9). ...

Evaluating UCE data adequacy and integrating uncertainty in a comprehensive phylogeny of ants

... Dominican amber is dated to the mid-Miocene and preserves over a thousand species of Neotropical arthropods in high fidelity [14]. This lagerstätte has informed new understanding of relict distributions [15], as well as patterns of faunal turnover [16][17][18] and phenotypic history [19]. The ants of Hispaniola are ecologically and morphologically diverse, both within amber and living today [16,20]. ...

The first fossil replete ant worker establishes living food storage in the Eocene

Myrmecological News

... Hypothenemus eruditus (Fig. 6) belongs to one of the most diverse scolytine genera in the world, with more than 220 species currently described (Wood 2007;Vega et al. 2015;Huang et al. 2016;Johnson et al. 2020). With more than 70 recognized synonyms (Vega et al. 2015), H. eruditus likely represent a complex of several closely related species whose morphological identification is extremely difficult. ...

Bark Beetle Hypothenemus eruditus Westwood, 1836 (Insecta: Coleoptera: Curculionidae: Scolytinae): EENY-664/IN1147, 10/2016

EDIS

... In the case of butterfly larvae, ants demonstrate "tending" behaviour, protecting the larvae, whilst the larvae provide food for the ants by secreting a liquid high in sugar and amino acids. Hill et al. (2022) investigated the former, finding that predation of late instars of the Miami blue larvae (Cyclargus thomasi bethunebakeri) did not occur when the tending Florida Carpenter Ant (Camponotus floridanus) was present, whereas almost all were predated when the tending ant was absent. Female Synargis calyce, among others, lay their eggs on plants The similarity between CHC profiles of the larvae and their respective host plants varied between 60 and 82%, with certain compounds only seen groups of caterpillars reared on specific plant species. ...

Protective Benefits of Tending Ants to a Critically Endangered Butterfly

Journal of Insect Science

... It primarily attacks unhealthy, damaged or felled trees, but it can also infest healthy stands (Merkl and 1992). This pest mainly targets trunks and larger branches, rather than roots near the ground, and it can attack both slender and stout sections (Sobel et al., 2015). Additionally, Xyleborus affinis has known to attack over two hundred economically important plant species, including Theobroma cacao, Mangifera indica and Saccharum officinarum (Biedermann, 2020). ...

Ambrosia Beetle Xyleborus affinis Eichhoff, 1868 (Insecta: Coleoptera: Curculionidae: Scolytinae): EENY 627/IN1094, 6/2015
  • Citing Article
  • August 2015

EDIS

... One occurred in the late-Miocene ~ 9 mya in the lineage leading to the largest Tetraopes clade, currently including 16 species. This colonization coincides with a global decrease in temperature and humidity during the Late Miocene, which gave rise to postglacial dispersal northward in some taxa [37][38][39] . The second colonization event to northern Mexico and the southwestern US involved a lineage that gave rise to T. skillmani. ...

Phylogeography and population genetics of a widespread cold‐adapted ant, Prenolepis imparis
  • Citing Article
  • July 2022

... To make matters worse, molecular data are lacking for most eurytomid genera in public databases, making it difficult for non-experts to identify specimens accurately. This is partly due to the low success rate in amplifying the typical barcoding fragment using universal primers (e.g., Li et al., 2010;Zhang et al., 2014), although recently, primer sets with higher success rates have been designed specifically for eurytomids (Jafari et al., 2023;Zhang et al., 2022). ...

Delimiting the cryptic diversity and host preferences of Sycophila parasitoid wasps associated with oak galls using phylogenomic data
  • Citing Article
  • June 2022

... In contrast, other oil-feeding ants, e.g. Pharaoh ants, Monomorium pharaonis L. and little fire ants, Wasmannia auropunctata (Roger), do not accept some fire ant baits, presumably due to the active ingredient and/or formulation (Montgomery et al. 2015;Oi et al. 2022;Williams and Vail 1993). The objective of this study was to identify currently available fire ant baits that are not fed upon by nontarget ants. ...

Response of Wasmannia auropunctata (Hymenoptera: Formicidae) to Water-Soaked Imported Fire Ant Baits
  • Citing Article
  • June 2022

Florida Entomologist

... beast2. org/, Jones et al. 2015) in BEAST2 for its demonstrated effectiveness in species boundaries validation (Busschau et al. 2019;Klimov et al. 2019;Prebus 2021;Williams et al. 2022). The estimated number of putative species in STACEY ranges from one to the number of putative clusters specified. ...

Phylogenomic Delimitation of Morphologically Cryptic Species in Globetrotting Nylanderia (Hymenoptera: Formicidae) Species Complexes

Insect Systematics and Diversity