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Fig. 10 in Messor erwini sp. n., a hitherto cryptic harvester ant in the Iberian Peninsula

Fig. 10. Messor capitatus major (A, C, E, G, H; collection code #19646) and minor (B, D, F, I, J; collection code #19643) workers. Lateral view of body (A, B); dorsal view of body (C, D); head in full-face view (E, F); frontal triangle in larger magnification (G, I); lateral view of petiole in larger magnification (H, J).

Fig. 7 in Messor erwini sp. n., a hitherto cryptic harvester ant in the Iberian Peninsula

Fig. 7. Dendrogram solution for the morphometric data of Iberian Messor species. Sample information in the dendrogram given as follows: final species hypothesis followed by a five-digit sample code applied by the genetic lab separated by a hyphen. Two columns of rectangles represent results of the partitioning algorithms 'hclust' and 'kmeans'. Different colors distinguish species. Messor barbarus: black, M. erwini sp. n.: green, M. capitatus: blue, M. bouvieri: red.

Fig. 5 in Messor erwini sp. n., a hitherto cryptic harvester ant in the Iberian Peninsula

Fig. 5. Phylogenetic analysis based on COI of Messor barbarus, M. bouvieri, M. capitatus, and the hitherto unidentified M. erwini sp. n. with Aphaenogaster iberica, M. lobognathus, and M. chamberlini as outgroups, using the Neighbor-Joining method based on Kimura 2-parameter. The percentage of replicate trees in which the associated taxa clustered together in the bootstrap test (1000 replicates) are shown next to the branches. The analysis was conducted in MEGA 7.0.26 (Felsenstein 1985; Saitou and Nei 1987; Nei and Kumar 2000; Kumar et al., 2016).

Fig. 1 in Messor erwini sp. n., a hitherto cryptic harvester ant in the Iberian Peninsula

Fig. 1. Sampling sites. (A) Iberian peninsula; (B) Catalonia. Occurrences of Messor barbarus (yellow, subclades 1 and 2 as defined by Romiguier et al., 2017), M. bouvieri (red), M. capitatus (blue), and the hitherto unidentified M. erwini sp. n. (green) depicted; at the focal study area Castellbell i el Vilar, M. barbarus, M. bouvieri, and the unidentified Messor co-occur (Google Earth Pro 2019, Data SIO, NOAA, U.S. Navy, NGA, GEBCO, Image Landsat/Copernicus).

Fig. 6 in Messor erwini sp. n., a hitherto cryptic harvester ant in the Iberian Peninsula

Fig. 6. Phylogenetic analysis based on COI of Messor barbarus, M. bouvieri, M. capitatus, and the hitherto unidentified M. erwini sp. n. with Aphaenogaster iberica, M. lobognathus, and M. chamberlini as outgroups, using the Maximum Likelihood method based on the General Time Reversible model. The percentage of replicate trees in which the associated taxa clustered together in the bootstrap test (1000 replicates) are shown next to the branches. The analysis was conducted in MEGA 7.0.26 (Nei and Kumar 2000; Kumar et al., 2016).

Fig. 3 in Messor erwini sp. n., a hitherto cryptic harvester ant in the Iberian Peninsula

Fig. 3. Method of Evanno et al. (2005) implemented on the STRUCTURE output data. L(K): mean ± standard deviation (SD) (left) and DeltaK (right). The analysis was conducted in STRUCTURE Harvester (Earl and von Holdt 2012).

Fig. 9 in Messor erwini sp. n., a hitherto cryptic harvester ant in the Iberian Peninsula

Fig. 9. Messor bouvieri major (A, C, E, G, H; collection code #19624) and minor (B, D, F, I, J; collection code #19620) workers. Lateral view of body (A, B); dorsal view of body (C, D); head in full-face view (E, F); frontal triangle in larger magnification (G, I); lateral view of petiole in larger magnification (H, J).

Fig. 4 in Messor erwini sp. n., a hitherto cryptic harvester ant in the Iberian Peninsula

Fig. 4. Bar plot of the Bayesian cluster analysis based on microsatellites genotyped in 286 individuals. The number of clusters is 4: Messor barbarus (yellow cluster), M. bouvieri (red), M. capitatus (blue), and the hitherto unidentified M. erwini sp. n. (green). The y axis shows the percentage of assignment to a certain cluster. The analysis was conducted in STRUCTURE 2.3.4 (Pritchard et al., 2000).

Fig. 8 in Messor erwini sp. n., a hitherto cryptic harvester ant in the Iberian Peninsula

Fig. 8. Messor barbarus major (A, C, E, G, H; collection code #19610) and minor (B, D, F, I, J; collection code #19612) workers. Lateral view of body (A, B); dorsal view of body (C, D); head in full-face view (E, F); frontal triangle in larger magnification (G, I); lateral view of petiole in larger magnification (H, J).

Fig. 2 in Messor erwini sp. n., a hitherto cryptic harvester ant in the Iberian Peninsula

Fig. 2. Position of morphometric characters on head in frontal view, including scape and frontal triangle, and on mesosoma in lateral and dorsal view; see Table 4 for definition of abbreviations and measurement of characters.

Fig. 11 in Messor erwini sp. n., a hitherto cryptic harvester ant in the Iberian Peninsula

Fig. 11. Messor erwini sp. n. major (A, C, E, G, H; holotype; collection code #19603e) and minor (B, D, F, I, J; paratype; collection code #19603d) workers. Lateral view of body (A, B); dorsal view of body (C, D); head in full-face view (E, F); frontal triangle in larger magnification (G, I); lateral view of petiole in larger magnification (H, J).

Fig. 3. Messor hodnii n in Révision taxonomique du genre Messor (Hymenoptera, Formicidae) au Maghreb et description de Messor hodnii sp. n., une nouvelle espèce de fourmi trouvée en Algérie

Fig. 3. Messor hodnii n. sp., tête avec psammophore.

Fig. 4. Site d in Révision taxonomique du genre Messor (Hymenoptera, Formicidae) au Maghreb et description de Messor hodnii sp. n., une nouvelle espèce de fourmi trouvée en Algérie

Fig. 4. Site d'échantillonnage aux abords de Chott El Hodna.

Fig. 1. Messor hodnii n in Révision taxonomique du genre Messor (Hymenoptera, Formicidae) au Maghreb et description de Messor hodnii sp. n., une nouvelle espèce de fourmi trouvée en Algérie

Fig. 1. Messor hodnii n. sp., corps vue de profil.

Fig. 2. Messor hodnii n in Révision taxonomique du genre Messor (Hymenoptera, Formicidae) au Maghreb et description de Messor hodnii sp. n., une nouvelle espèce de fourmi trouvée en Algérie

Fig. 2. Messor hodnii n. sp., tête vue de face.

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Data from: Turning one into five: integrative taxonomy uncovers complex evolution of cryptic species in the harvester ant Messor "structor"

Seed harvesting ants are ecosystem engineers that shape vegetation, nutrient cycles, and microclimate. Progress in ecological research is, however, slowed down by poor species delimitation. For example, it has not been resolved to date, how many species the European harvester ant Messor "structor" (Latreille, 1798) represents. Since its first description, splitting into additional taxa was often proposed but not accepted later on due to inconsistent support from morphology and ecology. Here, we took an iterative integrative-taxonomy approach – comparing multiple, independent data sets on the same sample – and used traditional morphometrics, Wolbachia symbionts, mitochondrial DNA, amplified fragment length polymorphism, and ecological niche modelling. Using the complementarity of the data sets applied, we resolved multiple, strong disagreements over the number of species, ranging from four to ten, and the allocation of individuals to species. We consider most plausible a five-species hypothesis and conclude the taxonomic odyssey by redescribing Messor structor, M. ibericus Santschi, 1925, and M. muticus (Nylander, 1849) stat.rev., and by describing two new species, M. ponticus sp.n. and M. mcarthuri sp.n. The evolutionary explanations invoked in resolving the various data conflicts include pronounced morphological crypsis, incomplete lineage-sorting or ongoing cospeciation of endosymbionts, and peripatric speciation – these ants' significance to evolutionary biology parallels that to ecology. The successful solution of this particular problem illustrates the usefulness of the integrative approach to other systematic problems of comparable complexity and the importance of understanding evolution to drawing correct conclusions on species' attributes, including their ecology and biogeography.