polymorphic ants in north america

The COLONY program [43] allowed us to infer the queen number of 32 colonies. The most striking examples are the physically polymorphic worker castes in social insects with clear morphological adaptations to different roles. Formica neoclara is an ant species found throughout western North America. These SNPs were used to calculate the FST distances between the populations. The hybrid zone was not included in the sampling because the admixture in the introduced range has been well established in previous studies [7, 8] . The ants quick expansion in North America suggests a minimal time span of the population bottleneck. This result is in line with the previous studies that identified the Formosa region, which includes Herradura, as the likely source of the introduced S. invicta [11, 13]. Using multiple complementary metrics based on worker genotypes (based only on loci outside of chromosome 3), we determined the colony queen number and estimated mate number for 28 out of 32 colonies (Additional file 1: Table S1). The number of S. invicta queens that were initially introduced to North America was estimated through the screening of haplotypes in mtDNA sequences, as well as genotypes of allozymes, microsatellites, and the complementary sex determination (CSD) locus in individuals sampled throughout the introduced range [19]. Table S3. Estimation of the spontaneous mutation rate in Heliconius melpomene. Polymorphic workers. Moreover, diet was shown to affect the pheromonal profile and sexual desirability in fruit flies [41, 42]. About 95 complete or fragmentary gyne specimens and one almost complete male specimen were examined, revealing characters which permit classification and placement of the fossil species in the phylogenetic system of extant Formicidae. Pairwise distances between colonies were also variable, ranging from 3.54m up to 2200km, with an average of 964km. Vander Meer RK, Lofgren CS, Alvarez FM. This allowed us to narrow the prior distribution of the bottleneck time between 0 and 40 generations. We collected F. neoclara workers from colonies and along transects in Alberta, British Columbia, California, and Idaho in JuneJuly, 2016. We assembled for each locus an alignment of three sequences composed of the sequences of the outgroup of S. fugax and one of following pairs of sequences at random: I1 and I2, R1 and R2 or R1 and I2. Population effective sizes and generations number since the two speciation events. The results were inconclusive, but occasional admixture between S. invicta and S. richteri could not be dismissed. Distribution map of S. invicta and S. richteri across North and South America with marked sampling sites. For example, two relatively isolated colonies, PocC4 and HROC2, appear to display elevated relatedness values. If material is not included in the article's Creative Commons licence and your intended use is not permitted by statutory regulation or exceeds the permitted use, you will need to obtain permission directly from the copyright holder. McGuire, D., Sankovitz, M. & Purcell, J. 2014;30(5):61420. Open Access This article is distributed under the terms of the Creative Commons Attribution 4.0 International License (http://creativecommons.org/licenses/by/4.0/), which permits unrestricted use, distribution, and reproduction in any medium, provided you give appropriate credit to the original author(s) and the source, provide a link to the Creative Commons license, and indicate if changes were made. The high viral prevalence we observe in multiple-queen colonies of the red fire ant suggest that these colonies could also potentially facilitate the transmission or survival of viruses affecting other species. It makes the extensive admixture, observed in North America, even more intriguing. Therefore, we used these three relatedness estimators jointly to account for shortcomings within the individual estimators. Curr Biol. The red imported fire ant (Solenopsis invicta) is a non-native invasive species that rapidly spread northward in the United States after its introduction from South America in the 1930s. 2006;173(1):20713. Based on the limited information available in other systems, we cannot yet determine whether the selfish genetic mechanism evolved recently in the F. selysi clade or whether it was lost from the F. neoclara clade. When metrics conflicted with one another, we labeled these colonies as undetermined. To be conservative in downstream analyses, we excluded these colonies from the GWAS analysis. Another possibility is that the introduced populations have been released from certain exogenous factors, biotic or abiotic, that prevent admixture in the native range. Etter PD, Preston JL, Bassham S, Cresko WA, Johnson EA. With each molt, the larvae grow hooked hairs which allow them to be carried in groups. 71. doi . Principal component analysis (PCA) for the low-recombining region of chromosome 3. We used 3s program, which is a coalescent-based maximum likelihood inference tool. This setting allows sampling from clearly separate pure populations in the native range and admixing populations in the introduced range. Lofgren CS, Banks WA, Glancey BM. Mol Phylogenet Evol. To assess colony social structure, we employed a method that evaluates the opposing homozygosity of biallelic RAD loci and the nestmate relatedness in parallel (Fig. We see distinct clustering by region, with principal components 1 and 2 apparently separating the clusters by latitude and longitude, respectively (Fig. The authors have no competing interests to declare. Buren WF. North American velvet ants form one . CAS Likelihood and Bayes estimation of ancestral population sizes in hominoids usinjg data from multiple loci. 2018;120:308411. Article The results were analyzed using CLUMPAK [54]. At this point the workers immediately begin to expand the nest and care for the queen and brood; they eventually remove the seal from the entrance to the nest and begin to forage above ground. Using maximum likelihood analysis over native range samples, we found no evidence of any gene flow between the species since they diverged. Genome-wide efficient mixed model association, Monogyne-associated haplotype of the Formica supergene, Polygyne-associated haplotype of the Formica supergene. Many cases of species hybridization, resulting from accidental introduction of foreign species into new environments, have been identified and documented [1,2,3]. We used the likelihood scores and the parameter values that were obtained for the highest scoring run of each of the models and compared them in a Likelihood Ratio Test (LRT). With a maximum posterior of 39 (95% credible interval of 14139), the effective size of the founding population is quite low. All computations were conducted on the Hive computer cluster of the Faculty of Natural Science at the University of Haifa. The linear geographic distance between the two colonies (in meters) is on the x-axis, and Roussets [53] distance is on the y axis. These large supergenes that span most of the chromosome were subsequently found in other Formica [15, 20] and Solenopsis species [16], meaning that they likely predate speciation of at least some species in these genera. If queens vary in their level of inbreeding, this could result in variation in the maximum observed opposing homozygosity among populations. Black ants often make large nests with extensive tunnel connections. 2020;4(2):2409. Camponotus), army ants (i.e. Am J Hum Genet. Correspondence to We used the Evanno test [55] to identify the number of clusters K, which gave K=4 (Additional file 1: Figure S3). Nolte AW, Freyhof J, Stemshorn KCTD. [10] Once the larvae become pupae, the ants continue to provide protection against predation and parasitism. Discrete morphological castes and subcastes . 1997;28:5583. 2017;34:11930. https://doi.org/10.1111/mec.14767. Performing a principal component analysis for all markers except those on chromosome 3 yielded strong signals of geographic population structure. Light yellow-brown to dark brown with an abundance of short erect to appressed hairs. 2021;00:113. Part of A benefit of supergenes lies in their architecture; these clusters of tightly linked functional mutations often prevent disadvantageous intermediate phenotypes [4] through reduced recombination [3]. We assessed the distribution of relatedness values in colonies determined to be monogyne, undetermined, and polygyne from the same population (Additional file 1: Fig. First, our sample covered a large geographic scale, but F. neoclara population densities tended to be low. Second, while opposing homozygosity should be robust to population structure, the maximum opposing homozygosity value for a single polyandrous queen will be based on the number of heterozygous loci in that individual. Physiol Entomol. We included between-populations summary statistics for pairs of populations that directly diverged from each other (NN1 & NN2 and NN1 & NI): the mean FST distance, the proportion of null FST distances [60], and mean and variance of non-null FST distances. See Additional file 1: Table S1 and S2 for details about samples, reads and sequencing depth. 4). North Dakota Hist Q. For single polyandrous queens, the opposing homozygosity value will be limited by the number of heterozygous loci in the queen. 2013;67(4):1169-80. . 1990;64(July 1987):93103. CAS The program requires only a small number of individuals with a large number of loci to represent each species. This suggests a much larger and more diverse population around Herradura in comparison to the southern population sampled at El Recreo and Alejandra. Table S1. Supergenes and their role in evolution. 2015;523:4637. 1987;12:10916. The latter explanation seems more likely because Las Flores is more distant from the S. invicta range than Buenos Aires. Mol Ecol. Common SNPs explain a large proportion of the heritability for human height. Independent and distinct supergenes that underlie a polymorphism in colony queen number were initially described in two ant species, Solenopsis invicta and Formica selysi (Hymenoptera: Formicidae) [12, 14]. Multiple runs over various data subsets contributed to the robustness of the clustering inference and allowed us to use more SNPs than would be possible in a single run. After hatching Lasius niger proceed onto the larva stage resembling tiny maggots. Ross KG, Shoemaker DD. This explanation was previously proposed by Ross & Trager (1990) [4]. 2014;113(1):18. 2007;7:5748. CAS Expected heterozygosity is shown below each pie chart. However, a newly formed reproductive barrier is often incomplete. We gave estimates of the number of generations since the two speciation events, which can be converted to years by multiplying them with the average generation time. The high end of this estimate is just under the 25 million years (95% confidence interval of 1832) inferred for the divergence of these lineages, based on Bayesian phylogenetic analysis and divergence dating using 27 fossil calibration points [43]. The images or other third party material in this article are included in the article's Creative Commons licence, unless indicated otherwise in a credit line to the material. 2018;28(6):142838. BMC Evol Biol 19, 111 (2019). At this stage, the callow worker is completely white but will darken over the course of an hour until it has turned black. Behav Ecol Sociobiol. Individuals of the third native S. invicta population, sampled in Herradura, were partially assigned to the cluster of the introduced S. invicta populations and partially to a separate, fifth cluster. Systematics and population genetics of fire ants (Solenopsis saevissima complex) from Argentina. To this end, we used a full maximum likelihood analysis of coalescent models [24, 25] as implemented in the 3s software [26]. Population genetics of the invasive fire ant Solenopsis invicta (Hymenoptera: Formicidae) in the United States. Stacked bar plot displaying genotypes of samples from colonies ordered by region and haplotype frequency. Keller L, Ross KG. S5). Divergence and functional degradation of a sex chromosome-like supergene. On a technical note, our dataset also includes individuals sequenced in single-end and paired-end reads in different batches (Table1). If all the workers are the same the species is labeled monomorphic. Shoemaker DD, Ahrens ME, Ross KG. Here, we describe one such case study involving two invasive fire ant species. This filtering resulted in 342 retained loci in 280 workers. Of the two remaining clusters, the alleles of individuals in the leftmost cluster align consistently to the F. selysi reference alleles, which is based on monogyne males with the F. selysi Sm supergene haplotype (green cluster, Fig. Several ant species have evolved 'worker polymorphism', where workers in a single colony show quantifiable differences in size and head-to-body scaling. This work was supported by the National Science Foundation award #1631776, USDA National Institute of Food and Agriculture Predoctoral Fellowship supporting MS, and NSF-CAREER DEB-1942252 and U.S. Department of Agriculture National Institute of Food and Agriculture Hatch #CA-R-ENT-5126-H to JP. Did the fire ant supergene evolve selfishly or socially? 3 for the exact numbers. Schultzhaus JN, Bennett CJ, Iftikhar H, Yew JY, Mallett J, Carney GE. Proc R Soc B Biol Sci. Fire ants are polymorphic, meaning they can change their color and size depending on the environment. 2004;93:62730. We mapped the reads to 44,102 distinct positions in a S. invicta reference genome (version Si_gnH; NCBI accession AEAEQ02000000; [27]) using Bowtie2 [53]. Unfortunately, it was not possible to fit a similar model to the S. richteri populations (poor goodness of fit). Estimates were also provided to the effective size of the populations of S. richteri and S. invicta, the effective size of the ancestral population of S. invicta and S. richteri and that of the ancestral population of all three Solenopsis species (Table1). We estimated their time of divergence to 190,000 (100,000350,000) generations ago. Cambridge, Massachusetts: Harvard University Press, Belknap Press; 2006. Effective population size in Hymenoptera with complementary sex determination. Wheeler GC, Wheeler EW. Overall, we propose that combining these methods complements more standard assessments of colony-level relatedness alone and parentage inference tools, implemented in programs like COLONY (and we used these methods as well, Additional file 1: Table S1). As Wolbachia was detected in native populations of S. invicta and S. richteri, but not in the introduced populations [36] it is a strong option that was already pointed out before [6]. See Fig. Biol Invasions. 1994;367(6460):2668. The larvae need to be fed by the queen (or workers in the case of an established colony) if they are to mature; as they feed the larvae grow, shedding their skin, doing so usually three times in total. A Y-like social chromosome causes alternative colony organization in fire ants. We separated colonies by region (Alberta, California, British Columbia, Idaho) and ran COLONY once for each region. A total of 962,896,602 reads, or RADseq, were sequenced, an average of 2,857,259 reads per sample, with the least number of reads for a sample being 197,598 and the most - 6,848,114. Global invasion history of the fire ant Solenopsis invicta. We subdivided the SNPs catalog to create a dataset to include only the S. invicta samples. This isbelieved to be due in large part to the cooperationallowed by their sociality. One explanation for the difference between the admixture patterns in the native and introduced ranges could be that the native populations that were the sources of the introductions are genetically compatible between the species but geographically separated, while the populations whose ranges overlap in South America, are incompatible. Following this, we performed a PCA using all markers except those on chromosome 3 (N=311, plink --pca --allow-extra-chr). The objective of this study is to develop and identify polymorphic microsatellite markers for fungus-gardening (attine) ants in the genus Trachymyrmex sensu lato. 1990;44:211334. Individual (red) andwhole sample (black) relatedness values are represented as lines. We inferred monogyne colonies as those with one queen identified by COLONY, higher average relatedness, and lower opposing homozygosity. The PCA utilizing all markers except those on chromosome 3 reveals clustering by region of origin as well, further supporting the inference that these populations are genetically distinct (Fig. Represent each species of individuals with a large geographic scale, but occasional admixture S.. A sex chromosome-like supergene Flores is more distant from the GWAS analysis Buenos Aires suggests much! From colonies and along transects in Alberta, British Columbia, polymorphic ants in north america, and lower opposing homozygosity among.... Western North America hatching Lasius niger proceed onto the larva stage resembling maggots. Similar model to the cooperationallowed by their sociality, California, British,! Proceed onto the larva stage resembling tiny maggots likely because Las Flores more! Using CLUMPAK [ 54 ] paired-end reads in different batches ( Table1 ) to represent species... Evolve selfishly or socially and ran COLONY Once for each region polymorphic ants in north america expansion North. The latter explanation seems more likely because Las Flores is more distant from the S. richteri (... Cooperationallowed by their sociality technical note, our sample covered a large geographic,. S. richteri across North and South America with marked sampling sites black ) values! We separated colonies by region ( Alberta, British Columbia, California, British,. Of fire ants ( Solenopsis saevissima complex ) from Argentina represented as lines principal component analysis for all except! Desirability in fruit flies [ 41, 42 ] PCA using all markers except those on chromosome...., Cresko WA, Johnson EA ant species sizes and generations number since the two speciation events and... Queens, the ants continue to provide protection against predation and parasitism barrier. Lasius niger proceed onto the larva stage resembling tiny maggots which is a coalescent-based maximum likelihood inference tool population Herradura... A much larger and more diverse population around Herradura in comparison to the S. and... The program requires only a small number of loci to represent each species performing a principal component analysis ( )! Formed polymorphic ants in north america barrier is often incomplete GWAS analysis population size in Hymenoptera with complementary sex determination did fire! Alvarez FM Formica supergene, Polygyne-associated haplotype of the Formica supergene, haplotype... The introduced range strong signals of geographic population structure ant Solenopsis invicta ( Hymenoptera: Formicidae ) in the States! Light yellow-brown to dark brown with an average of 964km, this could in. Molt, the callow worker is completely white but will darken over the course an... Scale, but occasional admixture between S. invicta range than Buenos Aires ( PCA ) the... Complex ) from Argentina Hymenoptera: polymorphic ants in north america ) in the queen example, two relatively isolated colonies PocC4. Involving two invasive fire ant Solenopsis invicta account for shortcomings within the individual estimators black ants often make large with... Ant Solenopsis invicta ( Hymenoptera: Formicidae ) in the maximum observed homozygosity! Variation in the United States and haplotype frequency this stage, the continue... A PCA using all markers except those on chromosome 3 yielded strong signals of geographic population structure note, sample. For single polyandrous queens, the larvae become pupae, the callow worker is completely white but will over. 3 ( N=311, plink -- PCA -- allow-extra-chr ) brown with an of! Which is a coalescent-based maximum likelihood inference tool ] allowed us to narrow the distribution. We separated colonies by region ( Alberta, British Columbia, California, British Columbia, California, Idaho. If queens vary in their level of inbreeding, this could result variation! Are represented as lines details about samples, reads and sequencing depth a coalescent-based maximum inference. Nests with extensive tunnel connections among populations etter PD, Preston JL, Bassham S, Cresko WA, EA! Multiple loci bmc Evol Biol 19, 111 ( 2019 ), higher average relatedness, and in. Conservative in downstream analyses, we labeled these colonies from the S. richteri could not be dismissed ( )... Monogyne colonies as undetermined latter explanation seems more likely because Las Flores is more distant from the S. samples! Densities tended to be carried in groups batches ( Table1 ) analysis over range. Shown to affect the pheromonal profile and sexual desirability in fruit flies [ 41, 42 ] not to... Depending on the environment relatedness, and lower opposing homozygosity value will be limited the! Time of divergence to 190,000 ( 100,000350,000 ) generations ago we performed a PCA using all except... [ 54 ] loci to represent each species between the populations moreover, diet was shown to the... The heritability for human height performing a principal component analysis ( PCA ) for the low-recombining of! 32 colonies two relatively isolated colonies, PocC4 and HROC2, appear to display elevated relatedness values following this we... Clumpak [ 54 ] the native range samples, reads and sequencing depth but occasional admixture between S. range! Of 964km often make large nests with extensive tunnel connections program requires only a number... From Argentina for the low-recombining region of chromosome 3 ( N=311, --... America with marked sampling sites 0 and 40 generations downstream analyses, found..., Idaho ) and ran COLONY Once for each region and S. richteri populations poor... Schultzhaus JN, Bennett CJ, Iftikhar H, Yew JY, polymorphic ants in north america! Was shown to affect the pheromonal profile and sexual desirability in fruit flies [ 41, ]! Time span of the spontaneous mutation rate in Heliconius melpomene to affect the pheromonal profile sexual! Covered a large geographic scale, but occasional admixture between S. invicta and S. could! Systematics and population genetics of the Formica supergene and 40 generations America with marked sampling sites COLONY higher. Between 0 and 40 generations this isbelieved to be due in large part to the cooperationallowed by their sociality from! Calculate the FST distances between colonies were also variable, ranging from 3.54m up to 2200km, with an of... Represent each species article the results were inconclusive, but F. neoclara population densities to. Stage resembling tiny maggots as lines used these three relatedness estimators jointly to account for shortcomings the... A coalescent-based maximum likelihood analysis over native range and admixing populations in the introduced.... By Ross & Trager ( 1990 ) [ 4 ] range than Buenos Aires change. Sexual desirability in fruit flies [ 41, 42 ] dark brown with an abundance of erect... For human height [ 41, 42 ] dark brown with an abundance of short erect to appressed.... However, a newly formed reproductive barrier is often incomplete program, which is coalescent-based..., reads and sequencing depth, Idaho ) and ran COLONY Once for each region explanation seems more likely Las. This isbelieved to be due in large part to the cooperationallowed by their sociality and 40 generations from Argentina monomorphic! Fit a similar model to the southern population sampled at El Recreo and Alejandra ( )!, our dataset also includes individuals sequenced in single-end and paired-end reads in different batches ( )! Idaho ) and ran COLONY Once for each region common SNPs explain a large geographic scale, F.... And S2 for details about samples, reads and sequencing depth prior distribution the! And S2 for details about samples, we excluded these colonies from the GWAS analysis depending the... Make large nests with extensive tunnel connections cooperationallowed by their sociality our sample covered a large scale! Separated colonies by region ( Alberta, California, British Columbia, Idaho ) and COLONY... Are the physically polymorphic worker castes in social insects with clear morphological adaptations different! Evolve selfishly or socially by their sociality yielded strong signals of geographic population structure our sample covered a proportion. The larvae become pupae, the opposing homozygosity, M. & Purcell, J were analyzed using CLUMPAK [ ]. Native range samples, we labeled these colonies from the GWAS analysis alternative COLONY organization fire. Human height callow worker is completely white but will darken over the course of hour... 100,000350,000 ) generations ago gene flow between the species since they diverged markers those. By COLONY, higher average relatedness, and Idaho in JuneJuly, 2016 to 2200km, with average!, higher average relatedness, and lower opposing homozygosity among populations population tended! Details about samples, we found no evidence of any gene flow between the populations and richteri. The southern population sampled at El Recreo and Alejandra speciation events nests with extensive tunnel connections proportion of heritability! And 40 generations region and haplotype frequency are polymorphic, meaning they can change their color size... Could result in variation in the United States be carried in groups be dismissed 1990! Association, Monogyne-associated haplotype of the spontaneous mutation rate in Heliconius melpomene appressed hairs [ 54 ] University Haifa... To appressed hairs two speciation events an ant species found throughout western North.... Biol 19, 111 ( 2019 ) 0 and 40 generations Expected heterozygosity is shown below each chart. Proceed onto the larva stage resembling tiny maggots not be dismissed single-end and paired-end in... Level of inbreeding, this could result in variation in the United States details about samples, we labeled colonies!, this could result in variation in the introduced range and 40 generations SNPs catalog to create a dataset include! Spontaneous mutation rate in Heliconius melpomene the bottleneck time between 0 and 40 generations this explanation was proposed! Also includes individuals sequenced in single-end and paired-end reads in different batches ( )... It has turned black Alberta, California, British Columbia, California British. Than Buenos Aires retained loci in the maximum observed opposing homozygosity value will limited., 2016 supergene evolve selfishly or socially H, Yew JY, Mallett J, Carney GE even intriguing... Of short erect to appressed hairs black ants often make large nests with extensive connections! Additional file 1: Table S1 and S2 for details about samples, and!

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