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The standard latitudinal diversity gradient (LDG), in which species richness decreases from equator to pole, is a pervasive pattern observed in most organisms. Some lineages, however, exhibit inverse LDGs. Seemingly problematic, documenting and studying contrarian groups can advance understanding of LDGs generally. Here, we identify one such contrarian clade and use a historical approach to evaluate alternative hypotheses that might explain the group's atypical diversity pattern. We focus on the biogeographical conservatism hypothesis (BCH) and the diversification rate hypothesis (DRH).

Global.

Ants (Hymenoptera: Formicidae: Stenammini).

We examined the shape of the LDG in Stenammini by plotting latitudinal midpoints for all extant, described species. We inferred a robust genome‐scale phylogeny using UCE data. We estimated divergence dates using beast2 and tested several biogeographical models inBioGeoBEARS. To examine diversification rates and test for a correlation between rate and latitude, we used the programs BAMM and STRAPP, respectively.

Stenammini has a skewed inverse LDG with a richness peak in the northern temperate zone. Phylogenomic analyses revealed five major clades and several instances of non‐monophyly among genera (Goniomma,Aphaenogaster). Stenammini and all its major lineages arose in the northern temperate zone. The tribe originated ~51 Ma during a climatic optimum and then diversified and dispersed southward as global climate cooled. Stenammini invaded the tropics at least seven times, but these events occurred more recently and were not linked with increased diversification. There is evidence for a diversification rate increase in HolarcticAphaenogaster + Messor, but we found no significant correlation between latitude and diversification rate generally.

Our results largely support the BCH as an explanation for the inverse latitudinal gradient in Stenammini. The clade originated in the Holarctic and likely became more diverse there due to center‐of‐origin, time‐for‐speciation and niche conservatism effects, rather than latitudinal differences in diversification rate.

 

Although savanna woody encroachment has become a global phenomenon, relatively little is known about its effects on multiple dimensions and levels of savanna biodiversity.

Using a combination of field surveys, a species‐level phylogeny, and functional metrics drawn from a morphological dataset, we evaluated how the progressive increase in tree cover in a fire‐suppressed savanna landscape affects the taxonomic, functional, and phylogenetic diversity of neotropical ant communities, at both the alpha and beta levels. Ants were sampled along an extensive tree cover gradient, ranging from open savannas to forests established in former savanna areas.

Variation in tree cover had a significant influence on all facets of diversity at the beta level, whereas at the alpha level tree cover variation affected the taxonomic and functional but not the phylogenetic diversity of the ant communities.

In general, ant community responses to variation in tree cover were largely non‐linear as differences in taxonomic alpha diversity and in the taxonomic, functional, and phylogenetic composition of the sampled communities were often much stronger at the savanna/forest transition than at any other part of the gradient. This indicates that savanna ant communities switch rapidly to an alternative state once the savanna turns into forest.

Ant communities in the newly formed forest areas lacked many of the species typical of the savanna habitats, suggesting that the maintenance of a fire suppression policy is likely to result in a decrease in ant diversity and in the homogenisation of the ant fauna at the landscape scale.

 

Uncovering the evolutionary history of the subfamilies Ectatomminae and Heteroponerinae, or ectaheteromorphs, is key to understanding a major branch of the ant tree of life. Despite their diversity and ecological importance, phylogenetic relationships in the group have not been well explored. One particularly suitable tool for resolving phylogeny is the use of ultraconserved elements (UCEs), which have been shown to be ideal markers at a variety of evolutionary time scales. In the present study, we enriched and sequenced 2,127 UCEs from 135 specimens of ectaheteromorph ants and investigated phylogeny using a variety of model-based phylogenomic methods. Trees recovered from partitioned maximum-likelihood and species-tree analyses were well resolved and largely congruent. The results are consistent with an expanded concept of Ectatomminae that now includes the subfamily Heteroponerinae new synonym and its single tribe Heteroponerini new combination. Eleven monophyletic groups are recognized as genera: Acanthoponera, Alfariastatus revived, Boltonia Camacho and Feitosa new genus, Ectatomma, Gnamptogenys, Heteroponera, Holcoponerastatus revived, Poneracanthastatus revived, Rhytidoponera, Stictoponerastatus revived, and Typhlomyrmex. The new phylogenetic framework and classification proposed here will shed light on the study of Ectatomminae taxonomy and systematics, as well as on the morphological evolution of the groups that it comprises.

 

Nylanderia(Emery) is one of the world's most diverse ant genera, with 123 described species worldwide and hundreds more undescribed. Fifteen globetrotting or invasive species have widespread distributions and are often encountered outside their native ranges. A molecular approach to understanding the evolutionary history and to revision ofNylanderiataxonomy is needed because historical efforts based on morphology have proven insufficient to define major lineages and delimit species boundaries, especially where adventive species are concerned. To address these problems, we generated the first genus‐wide genomic dataset ofNylanderiausing ultraconserved elements (UCEs) to resolve the phylogeny of major lineages, determine the age and origin of the genus, and describe global biogeographical patterns. Sampling from seven biogeographical regions revealed a Southeast Asian origin ofNylanderiain the mid‐Eocene and four distinct biogeographical clades in the Nearctic, the Neotropics, the Afrotropics/Malagasy region, and Australasia. The Nearctic and Neotropical clades are distantly related, indicating two separate dispersal events to the Americas between the late Oligocene and early Miocene. We also addressed the problem of misidentification that has characterized species‐level taxonomy inNylanderiaas a result of limited morphological variation in the worker caste by evaluating the integrity of species boundaries in six of the most widespreadNylanderiaspecies. We sampled across ranges of species in theN. bourbonicacomplex (N. bourbonica(Forel) + N. vaga(Forel)), theN. fulvacomplex (N. fulva(Mayr) + N. pubens(Forel)), and theN. guatemalensiscomplex (N. guatemalensis(Forel) + N. steinheili(Forel)) to clarify their phylogenetic placement. Deep splits within these complexes suggest that some species names – specificallyN. bourbonicaandN. guatemalensis– each are applied to multiple cryptic species. In exhaustively samplingNylanderiadiversity in the West Indies, a ‘hot spot’ for invasive taxa, we found five adventive species among 22 in the region; many remain morphologically indistinguishable from one another, despite being distantly related. We stress that overcoming the taxonomic impediment through the use of molecular phylogeny and revisionary study is essential for conservation and invasive species management.

 

Fungi shape the diversity of life. Characterizing the evolution of fungi is critical to understanding symbiotic associations across kingdoms. In this study, we investigate the genomic and metabolomic diversity of the genus Escovopsis , a specialized parasite of fungus-growing ant gardens. Based on 25 high-quality draft genomes, we show that Escovopsis forms a monophyletic group arising from a mycoparasitic fungal ancestor 61.82 million years ago (Mya). Across the evolutionary history of fungus-growing ants, the dates of origin of most clades of Escovopsis correspond to the dates of origin of the fungus-growing ants whose gardens they parasitize. We reveal that genome reduction, determined by both genomic sequencing and flow cytometry, is a consistent feature across the genus Escovopsis, largely occurring in coding regions, specifically in the form of gene loss and reductions in copy numbers of genes. All functional gene categories have reduced copy numbers, but resistance and virulence genes maintain functional diversity. Biosynthetic gene clusters (BGCs) contribute to phylogenetic differences among Escovopsis spp., and sister taxa in the Hypocreaceae. The phylogenetic patterns of co-diversification among BGCs are similarly exhibited across mass spectrometry analyses of the metabolomes of Escovopsis and their sister taxa. Taken together, our results indicate that Escovopsis spp. evolved unique genomic repertoires to specialize on the fungus-growing ant-microbe symbiosis. 

Abstract The ant genus Nylanderia Emery has a cosmopolitan distribution and includes 150 extant described species and subspecies, with potentially hundreds more undescribed. Global taxonomic revision has long been stalled by strong intra- and interspecific morphological variation, limited numbers of diagnostic characters, and dependence on infrequently collected male specimens for species description and identification. Taxonomy is further complicated by Nylanderia being one of the most frequently intercepted ant genera at ports of entry worldwide, and at least 15 globetrotting species have widespread and expanding ranges, making species-level diagnoses difficult. Three species complexes (‘bourbonica complex’, ‘fulva complex’, and ‘guatemalensis complex’) include globetrotting species. To elucidate the phylogenetic positions of these three complexes and delimit species boundaries within each, we used target enrichment of ultraconserved elements (UCEs) from 165 specimens representing 98 Nylanderia morphospecies worldwide. We also phased the UCEs, effectively doubling sample size and increasing population-level sampling. After recovering strong support for the monophyly of each complex, we extracted COI barcodes and SNPs from the UCE data and tested within-complex morphospecies hypotheses using three molecular delimitation methods (SODA, bPTP, and STACEY). This comparison revealed that most methods tended to over-split taxa, but results from STACEY were most consistent with our morphospecies hypotheses. Using these results, we recommend species boundaries that are conservative and most congruent across all methods. This work emphasizes the importance of integrative taxonomy for invasive species management, as globetrotting occurs independently across at least nine different lineages across Nylanderia. 

Abstract The evolutionary history of fungus-farming ants has been the subject of multiple morphological, molecular phylogenetic, and phylogenomic studies. Due to its rarity, however, the phylogenetic position, natural history, and fungal associations of the monotypic genus Paramycetophylax Kusnezov have remained enigmatic. Here we report the first excavations of colonies of Paramycetophylax bruchi (Santschi) and describe its nest architecture and natural history. Utilizing specimens from these collections, we generated ultraconserved-element (UCE) data to determine the evolutionary position of Paramycetophylax within the fungus-farming ants and ribosomal ‘fungal barcoding’ ITS sequence data to identify the fungal cultivar. A maximum-likelihood phylogenomic analysis indicates that the genus Paramycetophylax is the sister group of the yeast-cultivating Cyphomyrmex rimosus group, an unexpected result that renders the genus Cyphomyrmex Mayr paraphyletic. A Bayesian divergence-dating analysis indicates that Paramycetophylax diverged from its sister group around 36 mya (30–42 mya, HPD) in the late Eocene-early Oligocene, a period of global cooling, expansion of grasslands, and large-scale extinction of tropical organisms. Bayesian analysis of the fungal cultivar ITS gene fragment indicates that P. bruchi practices lower agriculture and that the cultivar grown by P. bruchi belongs to the Clade 1 group of lower-attine fungi, a clade that, interestingly, also includes the C. rimosus-group yeast cultivars. Based on these results, we conclude that a better understanding of P. bruchi and its fungal cultivar, including whole-genome data, is critical for reconstructing the origin of yeast agriculture, a major transition in the evolution of fungus-farming ants. 

The fungus-growing ant Mycetomoellerius (previously Trachymyrmex ) zeteki (Weber 1940) has been the focus of a wide range of studies examining symbiotic partners, garden pathogens, mating frequencies, and genomics. This is in part due to the ease of collecting colonies from creek embankments and its high abundance in the Panama Canal region. The original description was based on samples collected on Barro Colorado Island (BCI), Panama. However, most subsequent studies have sampled populations on the mainland 15 km southeast of BCI. Herein we show that two sibling ant species live in sympatry on the mainland: Mycetomoellerius mikromelanos Cardenas, Schultz, & Adams and M . zeteki . This distinction was originally based on behavioral differences of workers in the field and on queen morphology ( M . mikromelanos workers and queens are smaller and black while those of M. zeteki are larger and red). Authors frequently refer to either species as “ M . cf. zeteki ,” indicating uncertainty about identity. We used an integrative taxonomic approach to resolve this, examining worker behavior, chemical profiles of worker volatiles, molecular markers, and morphology of all castes. For the latter, we used conventional taxonomic indicators from nine measurements, six extrapolated indices, and morphological characters. We document a new observation of a Diapriinae (Hymenoptera: Diapriidae) parasitoid wasp parasitizing M . zeteki . Finally, we discuss the importance of vouchering in dependable, accessible museum collections and provide a table of previously published papers to clarify the usage of the name T . zeteki . We found that most reports of M . zeteki or M . cf. zeteki —including a genome—actually refer to the new species M . mikromelanos .