Insect Systematics and Diversity, (2022) 6(1): 5; 1–20
https://doi.org/10.1093/isd/ixab026
Research
Molecular Phylogenetics, Phylogenomics, and Phylogeography
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UCE Phylogenomics Resolves Major Relationships
Among Ectaheteromorph Ants (Hymenoptera: Formicidae:
Ectatomminae, Heteroponerinae): A New Classification
For the Subfamilies and the Description of a New Genus
G. P. Camacho,1,2,3,4,8, W. Franco,1, M. G. Branstetter,5 M. R. Pie,6, J. T. Longino,7,
T. R. Schultz,2 and R. M. Feitosa1
1
Programa de Pós-Graduação em Entomologia, Departamento de Zoologia, Universidade Federal do Paraná, 82590-300, Curitiba,
Brazil, 2Department of Entomology, National Museum of Natural History, Smithsonian Institution, 20560, Washington, DC, USA,
3
Department of Entomology, Institute for Biodiversity Science and Sustainability, California Academy of Sciences, 94118, San
Francisco, CA, USA, 4Center for Integrative Biodiversity Discovery, Leibniz Institute for Evolution and Biodiversity Science, Museum
für Naturkunde, 10115, Berlin, Germany, 5U.S. Department of Agriculture, Agricultural Research Service (USDA-ARS), Pollinating
Insects Research Unit, Utah State University, 84322, Logan, UT, USA, 6Programa de Pós-Graduação em Zoologia, Departamento de
Zoologia, Universidade Federal do Paraná, 82590-300, Curitiba, Brazil, 7Department of Biology, University of Utah, 84112, Salt Lake
City, UT, USA, and 8Corresponding author, e-mail: gabieco.camacho@gmail.com
Subject Editor: Jeffrey Sosa-Calvo
Received 7 July 2021; Editorial decision 19 October 2021
Abstract
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, Alfaria status revived, Boltonia
Camacho and Feitosa new genus, Ectatomma, Gnamptogenys, Heteroponera, Holcoponera status revived,
Poneracantha status revived, Rhytidoponera, Stictoponera status 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.
Key words: phylogenomics, ultraconserved elements, Formicidae, Ectatomminae, Heteroponerinae
Ants are a globally diverse lineage of eusocial aculeate wasps and
represent one of the great success stories of evolution, being the
richest and most ecologically dominant group among all social insects (Hölldobler and Wilson 2008). The taxonomy and internal
phylogeny of Formicidae have been significantly stabilized in recent
decades due to extensive study of ant systematics on a global scale
(Baroni Urbani et al. 1992, Bolton 1995, Brady et al. 2006, Moreau
et al. 2006, Ward 2014). Many of the findings from these studies
have shown a great congruence between existing morphological
and molecular hypotheses, such as the monophyly of the subfamily
Proceratiinae and the recognition of the subfamily Paraponerinae as
a distinct lineage among poneroid ants (Ouellette et al. 2006). Other
findings, however, highlight the need for a better understanding
of the ancestral morphology and biology of ants. The subfamilies
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1
2
Fig. 1. In lateral view, workers of the Ectatomminae genera, showing
the morphological diversity within the clade. (A) Acanthoponera
mucronata (CASENT0173540), (B) Alfaria minuta (CASENT0281213), (C)
Ectatomma planidens (CASENT0173379), (D) Gnamptogenys acuminata
(USNMENT00441095), (E) Heteroponera panamensis (CASENT0106021),
(F) Holcoponera ammophila (CASENT0281512), (G) Poneracantha mecotyle
(CASENT0281530), (H) Rhytidoponera metallica (CASENT0172345), (I)
Stictoponera biroi (CASENT0172380), (J) Typhlomyrmex rogenhoferi
(CASENT0173390). See Fig. 3 for images of Boltonia microps. Images by
April Nobile, Estella Ortega, Michael Branstetter, Zach Lieberman, and Jeffrey
Sosa-Calvo; available from www.antweb.org (Antweb 2021).
(Amblyoponinae)). However, most of these studies were limited to
analyzing only a relatively low number of mitochondrial and nuclear
genes, sequenced using traditional Sanger sequencing methods (except
for Blaimer et al. 2015, Branstetter et al. 2017, and Borowiec 2019).
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Ectatomminae and Heteroponerinae (commonly referred to as
ectaheteromorphs) are important in this regard. Although possessing
morphological and behavioral traits thought to be plesiomorphic for
ants as a whole (Baroni Urbani 1989, Hölldobler and Wilson 1990,
Keller 2000, Bolton 2003, Ward and Brady 2003), the two subfamilies are part of the large formicoid clade, in which they are sister
to the highly derived Myrmicinae (Brady et al. 2006, Moreau et al.
2006, Ouellette et al. 2006, Branstetter et al. 2017).
The ectaheteromorphs contain 302 described ant species (Bolton
2021) distributed across most tropical and subtropical regions of the
world, with a substantial number of species also occurring in hot temperate environments (Camacho and Feitosa 2015, Feitosa 2015). Species
live and forage in the soil and vegetation and are known to nest underground, in rotting logs, in leaf litter, or in trees, with colony sizes ranging
from a few dozen to a few hundred workers. Ectaheteromorph workers
vary morphologically, from large ants with robust bodies and welldeveloped compound eyes to tiny and totally blind (Fig. 1). They also
range from possessing very short to very long appendages. The cuticle
varies from coarsely sculptured to polished and shiny. Coloration can be
drab or highly conspicuous (Camacho and Feitosa 2015, Feitosa 2015).
The clade has a disjunct distribution, occurring in the Neotropical
region (with a minor extension into the southern Nearctic) and in the
Australian and Indomalayan regions (Janicki et al. 2016). Currently, the
subfamily Ectatomminae is divided into two tribes: (1) Ectatommini,
composed of the genera Ectatomma Fr. Smith (Hymenoptera:
Formicidae), exclusive to the Neotropical region, Rhytidoponera
Mayr (Hymenoptera: Formicidae), occurring only in the Australian
region, and Gnamptogenys Roger (Hymenoptera: Formicidae),
present in the Neotropical, Nearctic, Indomalayan, and Australasian
regions; and (2) Typhlomyrmecini, composed of the single genus
Typhlomyrmex Mayr (Hymenoptera: Formicidae), which is strictly
Neotropical in distribution. Heteroponerinae contains a single
tribe, Heteroponerini, which includes the genus Acanthoponera
Mayr (Hymenoptera: Formicidae), strictly Neotropical, and
Heteroponera Mayr (Hymenoptera: Formicidae), which has a disjunct
distribution between the Neotropical and Australian regions. The
enigmatic genus Aulacopone Arnoldi (Hymenoptera: Formicidae),
known only from two collection events in Azerbaijan, including the
type locality, is currently incertae sedis in the subfamily. The taxonomic limits of the ectaheteromorph genera have been relatively stable
since they were originally proposed and there have been numerous
species-level treatments within individual genera (e.g., Ward 1980,
Ward 1984, Lattke 1995, Lattke 2004, Nettel-Hernanz et al. 2015,
Camacho et al. 2020). There have been multiple attempts to understand relationships among the genera using morphology alone (Emery
1911, Brown 1965, Lattke 1994, Keller 2011, Feitosa 2015) but the
results have been contradictory or poorly supported. The monophyly
of genera has also never been formally tested using molecular data.
The incorporation of molecular biology into phylogenetic inference has greatly advanced understanding of ant evolution and
ecological success. Several studies investigated the early evolution
and diversification of ants (Brady et al. 2006, Ouellette et al. 2006,
Moreau and Bell 2013), resolving most of the relationships among
subfamilies (Branstetter et al. 2017, Borowiec et al. 2019). Among
the 17 subfamilies of Formicidae, internal phylogenetic relationships
have been extensively studied using molecular data in only ten, accounting for 94% of the described species diversity within the family
(Ward and Brady 2003 (Myrmeciinae); Ward et al. 2010 (Aneuretinae
and Dolichoderinae); Schmidt 2013 (Ponerinae); Brady et al. 2014,
Borowiec 2019 (Dorylinae); Ward et al. 2015 (Agroecomyrmecinae
and Myrmicinae); Chomicki et al. 2015 (Pseudomyrmecinae); Blaimer
et al. 2015, Ward et al. 2016 (Formicinae); Ward and Fisher 2016
Insect Systematics and Diversity, 2022, Vol. 6, No. 1
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Insect Systematics and Diversity, 2022, Vol. 6, No. 1
Methods
Taxon Sampling
Our dataset comprised 135 individuals belonging to 130 species
of ectaheteromorph ants (Supp Table S1 [online only]). The only
genus we could not sample was Aulacopone, which is a monotypic
genus known only from its holotype (collected in 1929 and currently
lost) and by another specimen that was collected in 1936, which
was coated with gold-palladium for scanning electron microscopy
long ago. We maximized the sampling breadth by including at least
one representative from each biogeographic region in which a genus
occurs and by sampling across morphologically disparate groups
within genera. In addition, we included 15 taxa to serve as closely
related outgroups from six ant subfamilies (Myrmicinae, Dorylinae,
Pseudomyrmecinae, Formicinae, Myrmeciinae, and Dolichoderinae)
(Supp Table S1 [online only]) belonging to the formicoid clade of
ants (sensu Brady et al. 2006). Trees were rooted using Dorylinae.
The total sample comprised 73 species of Gnamptogenys (77 terminals), 13 species of Heteroponera (14 terminals), four species of
Acanthoponera (four terminals), three species of Typhlomyrmex
(three terminals), 26 species of Rhytidoponera (26 terminals), and
11 species of Ectatomma (11 terminals). All specimens included in
this study were collected in accordance with local regulations and all
necessary permits were obtained. Voucher specimens have been deposited at the Entomological Collection Padre Jesus Santiago Moure
of the Federal University of Paraná (DZUP), Brazil; at the John
T. Longino personal collection (JTLC), University of Utah, Salt Lake
City, UT, USA; and at the Smithsonian Institution National Museum
of Natural History (NMNH/USNM), Washington, DC, USA.
Morphological Data
We examined the external morphology of adult forms to produce
diagnostic information for the formal and informal groupings proposed in this study (Supp Table S2 [online only]), following the terminology traditionally used for myrmecological revisions (Keller
2011). For the surface sculpturing, we followed the terminology proposed by Harris (1979). The type material was examined in person
or by photographs, when available at www.antweb.org (Antweb
2021). Taxonomic history for the species follows Bolton (2021).
Molecular Data Collection
DNA was extracted destructively or non-destructively from adult
workers using a DNeasy Blood and Tissue Kit (Qiagen, Valencia,
CA, USA). We quantified DNA for each sample using a Qubit fluorometer (High sensitivity kit, Life Technologies, Inc.) and sheared
5.7– 271 ng (x̄ = 47 ng) of DNA to a target size of approximately
600 bp by sonication (Qsonica). The sheared DNA was used as input
for a modified genomic DNA library preparation protocol (Kapa
Hyper Prep Library Kit, Kapa Biosystems) that incorporated ‘withbead’ cleanup steps (Fisher et al. 2011) and a generic SPRI substitute
(Rohland and Reich 2012), “speedbeads” hereafter, as described by
(Faircloth et al. 2015). We used TruSeq-style dual indexing adapters
during adapter ligation (Glenn et al. 2019), and PCR-amplified 50%
of the resulting library volume. After rehydrating and purifying reactions, we combined groups of ten libraries at equimolar ratios into
enrichment pools having final concentrations of 153–178 ng/μL.
We enriched each pool using a set of 9,898 custom-designed
probes (MYcroarray, Inc., now Arbor Biosciences) targeting 2,524
UCE loci specific for ants (Branstetter et al. 2017, ‘hym-v2-antspecific’). We followed library enrichment procedures for the
MYcroarray MYBaits kit (Blumenstiel et al. 2010), except we used
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Phylogenomic methods, in contrast, can efficiently generate
hundreds to thousands of loci for phylogenetic inference, allowing
for the resolution of previously intractable phylogenetic problems and providing increased confidence (Johnson et al. 2013,
Blaimer et al. 2015, Branstetter et al. 2017). Phylogenomic approaches can increase the number of characters available hundreds
to thousands of times, which can reduce stochastic error for phylogenetic inference (Delsuc et al. 2005) and help to overcome phylogenetic conflict among gene trees (Camacho et al. 2019). Among
alternative phylogenomic markers, ultraconserved elements (UCEs)
are ideal for the study of evolutionary relationships at different time
scales (Faircloth et al. 2015). Enrichment of UCE loci has been used
to investigate issues involving older phylogenetic divergences for
various vertebrates (Crawford et al. 2012, Faircloth et al. 2013a;
McCormack et al. 2013), several insect groups (Faircloth et al. 2015,
Blaimer et al. 2016a), and ants (Blaimer et al. 2015, Branstetter et al.
2017). The technique is also useful for understanding relationships
at the species and population level (Smith et al. 2013, Ješovnik et al.
2017; Ströher et al. 2019, Branstetter and Longino 2019, Longino
and Branstetter 2020, Prebus 2021). UCE enrichment is effective
even for poorly preserved specimens with degraded DNA (Blaimer
et al. 2016b), and the cost is relatively low compared to other DNAsequencing methods (Branstetter et al. 2017).
Previous studies have supported the monophyly of the
ectaheteromorphs and their placement near the Myrmicinae.
They have long been thought to be closely related to the subfamily Myrmicinae, based on morphology (Brown 1958, Bolton
2003). Feitosa (2015) discovered ten diagnostic characters for the
ectaheteromorph group, providing morphological support for
monophyly. Early molecular datasets supported the monophyly of
the ectaheteromorphs, but estimates of their placement relative to
other subfamilies were uncertain (Brady et al. 2006, Moreau et al.
2006). Using UCEs, Branstetter et al. (2017) found the first strong
evidence that ectaheteromorphs were a sister clade to Myrmicinae,
the most diverse subfamily of ants. The study, however, focused on
relationships among subfamilies and included UCE data for only
four ectaheteromorph species. Thus, this and all previous molecular
studies have been based on very limited taxon sampling within the
ectaheteromorphs.
Here, we use UCE data to reconstruct the phylogeny of
ectaheteromorph ants and improve ectaheteromorph systematics. To
do so, we assembled a data set of 2,127 UCE loci by means of target
enrichment and multiplexed sequencing of 135 ectaheteromorph
taxa, greatly expanding the taxon sampling of Branstetter et al.
(2017). We selected taxa to contain a broad representation of species across genera and were able to include six of the seven currently valid ectaheteromorph genera and many of the species groups
within genera. Our detailed objectives were to (1) use phylogenomic
information and dense taxon sampling to test the monophyly of subfamilies, tribes, and genera within the ectaheteromorphs; (2) resolve
phylogenetic relationships among lineages; and (3) use the results to
improve the ectaheteromorph classification at all taxonomic levels.
Based on the phylogenetic results and morphology, and in order to establish an evolutionary classification in which higher taxa are monophyletic, we: (i) synonymize Heteroponerinae under Ectatomminae;
(ii) describe one new genus, Boltonia Camacho and Feitosa gen.n.
(Hymenoptera: Formicidae); (iii) revive the genera Alfaria Emery
(Hymenoptera: Formicidae), Holcoponera Mayr (Hymenoptera:
Formicidae), Poneracantha Emery (Hymenoptera: Formicidae), and
Stictoponera Mayr (Hymenoptera: Formicidae) from synonymy; and (iv) provide an illustrated identification key for the
Ectatomminae genera.
4
Processing and Alignment of UCE Data
The sequencing facility demultiplexed and converted raw data from
BCl to FASTQ format using BCL2FASTQ (available at http://support.
illumina.com/downloads/bcl2fastq_conversion_software_184.html).
We trimmed the demultiplexed FASTQ data output for adapter contamination and low-quality bases using Illumiprocessor (Faircloth
2013b), which is a wrapper program around TRIMMOMATIC
(Bolger et al. 2014). All further data processing described in the following relied on scripts within the PHYLUCE v1.5. package. We
computed summary statistics on the data using the get_fastq_stats.
py script, and assembled the cleaned reads using the assemblo_trinity.
py wrapper around the program Trinity (v2013-02-25) (Grabherr
et al. 2011). Average sequencing coverage across assembled contigs
was calculated using get_trinity_coverage.py. To identify assembled
contigs representing enriched UCE loci from each species, speciesspecific contig assemblies were aligned to the ant-specific hym-v2
bait file (Branstetter et al. 2017) using match_contigs_to_probes.py
(min_coverage = 50, min_identity = 80), and sequence coverage statistics (avg, min, max) for contigs containing UCE loci were calculated
using get_trinity_coverage_for_uce_loci.py. Subsequently, we used get_
match_counts.py to query the relational database containing matched
probes created in the previous step, in order to generate a list of UCE
loci shared across all taxa. This list of UCE loci was then used in the
get_fastas_from_match_counts.py script to create FASTA files for each
UCE locus, which contain sequence data for taxa present at that particular locus (Supp Table S3 [online only]). We aligned all data in all
these FASTA files using MAFFT (Katoh et al. 2009) through seqcap_
align_2.py (min-length = 20, no-trim). Following alignment, we further
trimmed our alignment using a wrapper script (get_gblocks_trimmed_
alignment_from_untrimmed.py) for Gblocks (Castresana 2000) using
the following settings: b1=0.5, b2=0.5, b3=12, b4=7.We then used get_
only_loci_with_min_taxa.py to filter the initial set of alignments to include only loci with data for more than 75% of taxa (>112 of 150) or
90% of taxa (>135 of 150). These are referred to as the 75p-matrix
and 90p-matrix, respectively (Supp Table S4 [online only]).
Phylogenetic Inference
We performed a set of sensitivity analyses of our dataset, by
employing both concatenated and species-tree analyses on the different data matrices, and also by recoding the nucleotide data to
RY-characters. This set of sensitivity analysis was performed to allow
for assumptions that differ from those used in the primary analysis
and to check the robustness of the results.
For the concatenated analyses, we used the Sliding-Window Site
Characteristics based on Entropy method (SWSC-EN; Tagliacollo
and Lanfear 2018) to partition the UCE data for phylogenetic analysis. This method breaks UCE loci into three regions, corresponding
to the right flank, core, and left flank. The theoretical underpinning
of the approach comes from the observation that UCE core regions
are conserved, whereas the flanking regions become increasingly more
variable (Faircloth et al. 2012). After running the SWSC-EN algorithm,
the resulting data subsets were analyzed using PARTITIONFINDER2
(Lanfear et al. 2012). For this analysis we used the rclusterf algorithm,
AICc model-selection criterion, and the GTR+G model of sequence
evolution. Using the SWSC-EN partitioning scheme and concatenated
matrices, we inferred phylogenetic relationships of ectaheteromorphs
with the likelihood-based program IQ-TREE v1.5.5 (Nguyen et al.
2015). For the analysis we selected the ‘-spp’ option for partitioning
and the ”-m MFP” option for ModelFinder (Kalyaanamoorthy et al.
2017) to select the best model of sequence evolution. To assess branch
support, we performed 1,000 replicates of the ultrafast bootstrap approximation (UFB) (Minh et al. 2013, Hoang et al. 2018). Additionally,
we performed matched-pair tests of symmetry to test the assumptions
of stationarity and homogeneity for the partition scheme. We used
the ‘—symtest-remove-bad’ option on IQ-TREE v.2.1.3 to remove all
‘bad’ partitions (pvalue cutoff = 0.050) and continued the analysis with
the remaining ‘good’ partitions, as described by Naser-Khdour et al.
(2019). The resulting best-fit partitioning scheme included 1,427 data
subsets (245 ‘bad’ partitions removed) for the 75p-matrix and 902
data subsets (147 ‘bad’ partitions removed) for the 90p-matrix and
had a significantly better log likelihood than alternative partitioning
schemes (75p-matrix: SWSC-EN-symtest: -13272290.208; SWSC-EN:
-16,476,333.842; By Locus: -16,773,830.932; Unpartitioned:
-16,912,745.749; 90p-matrix: SWSC-EN-symtest: -8903417.276;
SWSC-EN:
-10,811,172.113;
By
Locus:
-11,010,444.832;
Unpartitioned: -11,093,438.532). We also recoded the nucleotides to
RY-characters for both matrices in an attempt to reduce possible negative effects caused by base composition heterogeneity or saturation
(Phillips and Penny 2003). For these support measures, values ≥ 95%
signal were regarded as well supported in this study.
For species-tree analyses, we used the SWSC-EN partitioning
scheme to estimate gene trees for the 2,180 UCE loci in the 75p-matrix
and the 1,351 UCE loci in the 90p-matrix, since partitioning the
UCE loci can improve gene-tree resolution (Freitas et al. 2021). Each
partitioned gene tree reconstruction was done with IQ-TREE using
the ”-m MFP” option for ModelFinder for the best model fit with
1000 UFB replicates. We also contracted very low support branches
(e.g., below 10% bootstrap support) from gene trees, since Zhang
et al. (2018) showed that this can improve accuracy in species tree
estimation. Species-tree analyses with local posterior probability
support values were performed in ASTRAL-III (Zhang et al. 2018)
using the manipulated gene trees as input.
Finally, given that there was relatively low support for some of
the inferred nodes (see results), we explicitly explored the level of
support of each locus for competing topologies. First, we obtained
gene trees for all 2,520 loci in our dataset without partitioning, as
well as the mean ultrafast-bootstrap support and GC content for the
corresponding locus. We then counted how many gene trees supported each competing topology using the testMono function in
‘ape’ (Paradis and Schliep 2019) in R v.3.6.3. (R Core Team 2020).
This also allowed us to test if a given topology was supported by
loci with biased base composition and/or low signal (i.e., low mean
average bootstrap support across all nodes). All the above phylogenetic analyses were performed on the Smithsonian Institution’s HighPerformance Computing Cluster (SI/HPC).
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a 0.1X concentration of the standard MYBaits concentration and
added 0.7 μL of 500 μM custom blocking oligos designed against our
custom sequence tags. We ran the hybridization reaction for 24 h at
65 °C, subsequently bound all pools to streptavidin beads (MyOne
C1; Life Technologies), and washed bound libraries according to a
standard target enrichment protocol (Faircloth et al. 2012). We used
the with-bead approach for PCR recovery of enriched libraries as described in Faircloth et al. (2012). We combined 15 μL of streptavidin
bead-bound, enriched library with 25 μL HiFi Ready Mix (Kapa
Biosystems), 5 μL of Illumina TruSeq primer mix (2.5 μM each),
and 5 μL of ddH2O. We purified resulting reactions using 1.0X
speedbeads, and we rehydrated the enriched pools in 22 μL EB.
We quantified 2 μL of each enriched pool using a Qubit fluorometer (broad range kit). Enriched DNA samples were sequenced on
four Illumina HiSeq 2500 lanes (2x125bp v4 chemistry) at the High
Throughput Genomics Lab at the University of Utah. All of the UCE
laboratory work was conducted at the University of Utah.
Insect Systematics and Diversity, 2022, Vol. 6, No. 1
Insect Systematics and Diversity, 2022, Vol. 6, No. 1
Data Availability
All phylogenetic datasets are available in the Dryad data repository under https://doi.org/10.5061/dryad.sxksn034j. Raw sequence
data files have further been submitted to NCBI’s Sequencing Read
Archive (BioProject PRJNA668430) (Supp Table S6 [online only]).
Nomenclature
Results
UCE Capture Statistics
An average of 45,784 contigs with a mean length of 420.2 bp were
assembled by Trinity after adapter- and quality-trimming of raw
reads, with an average contig coverage of 7.3X (Supp Table S3 [online only]). From the bulk set of contigs, we extracted an average of
2,126 UCE loci per sample and these had a mean contig length of
722.3 bp and average coverage of 39.2X. The 75p-matrix retained
2,180 loci, which provided 1,205,560 bp of sequence data, 558,013
informative sites, and only 10.05% missing data. The 90p-matrix retained 1,351 loci, generating 792,650 bp of sequence data, of which
368,562 were informative, with 7.1% missing data. For additional
sequencing and assembly information see Supplementary Material
(Supp Tables S3 and S4 [online only]).
Phylogenetic Results
Our concatenated, RY-recoded, and species-tree analyses recovered
highly congruent topologies for Ectatomminae, with only a few
incongruences at the genus and species levels. Analysis of the concatenated 90p-matrix recovered a highly resolved phylogeny for
the ectaheteromorphs with most nodes displaying maximum UFB
support (Fig. 2). Only a few nodes were recovered with a UFB
score lower than 95%, mainly involving interspecific relationships
among closely related species within a genus (Fig. 2). For the 90pRY concatenated analysis, we also recovered a highly resolved
phylogeny with high support, but with some differences in generic
relationships from the 90p-matrix, most notably the paraphyly of
Heteroponera in relation to Acanthoponera (Supp Fig. S1 [online
only]). The 75p-matrix analysis recovered results very similar results to those of the 90p-RY analysis. Relationships among species were congruent, except for the position of Heteroponera
sp._GPC22 (Supp Fig. S2 [online only]). The 75p-RY concatenated
analysis was also mostly congruent with the 90p-RY dataset, but
recovered some conflicting relationships between Poneracantha,
Alfaria, and Holcoponera (Supp Fig. S3 [online only]). The species
trees estimated by ASTRAL-III closely matched the topology estimated by the 90p-matrix concatenated analysis of nucleotide data,
with most nodes showing maximum local posterior probability
(LPP) support values (Supp Figs. S4 and S5 [online only]). All of the
results discussed below refer to the 90p-matrix concatenated tree,
except where noted, since this was the topology with the highest
likelihood value and because the completeness of the matrix minimizes the effect of missing data.
The ectaheteromorphs, as currently defined, encompass two
different subfamilies. We found strong support for the monophyly
of both subfamilies (heteroponerines: UFB = 100, LPP = 1;
ectatommines: UFB = 100, LPP = 1) (Fig. 2, Supp Figs. S1–S5
[online only]) and for the sister-group relationship between them
(UFB = 100; LPP = 1) (Fig. 2, Supp Figs. S1–S5 [online only]). We also
recovered the ectaheteromorphs (heteroponerines + ectatommines)
as the sister clade of the Myrmicinae.
The heteroponerines include the genera Heteroponera and
Acanthoponera. Acanthoponera was recovered as monophyletic in
all analyses (UFB = 100; LPP = 1) (Fig. 2, Supp Figs. S1–S5 [online
only]). Heteroponera, in contrast, was recovered as paraphyletic with
respect to Acanthoponera, with a single species, Boltonia microps
(Borgmeier) new combination (formerly classified as Heteroponera),
clearly separated from the other species of Heteroponera and sister
to all other Heteroponerinae with maximum support (UFB = 100;
LPP = 1) (Fig. 2, Supp Figs. S1–S5 [online only]). The remaining
Heteroponera species were recovered as a monophyletic group in the
concatenated analyses of the 90p-matrix (UFB = 76) (Fig. 2, Supp
Fig. S2 [online only]) and in the species-tree analyses, although with
low support (90p-matrix: LPP = 0.49; 75p-matrix: LPP = 0.86) (Supp
Figs. S4 and S5 [online only])). Analsyes of the 75p-matrix, as well
as of both RY-coded matrices, recovered Heteroponera monticola
Kempf and Brown, a South American species, as sister to a clade
comprising Acanthoponera and Heteroponera (UFB = 100) (Supp
Figs. S1–S3 [online only]).
Ectatommini, as classified here, comprises eight extant genera,
all of them included in our analyses, and together they formed a
clade with maximum support in all analyses (UFB = 100; LPP = 1)
(Fig. 2, Supp Figs. S1–S5 [online only]). Within the tribe the reciprocally monophyletic genera Rhytidoponera (UFB = 100; LPP = 1) and
Ectatomma (UFB = 100; LPP = 1) formed a clade in the concatenated
and species-tree analyses (UFB =100; LPP = 1) (Fig. 2, Supp Figs. S2,
S4, and S5 [online only]), as well as for the 75p-matrix converted to
RY-coding (UFB = 98) (Supp Fig. S3 [online only]), and the clade was
recovered as sister to all other ectatommines. For the 90p-matrix
converted to RY-coding, Ectatomma was recovered as sister to all
remaining ectatommines with full support, and Rhytidoponera was
sister to the remaining genera (UFB = 97) (Supp Fig. S1 [online only]).
The genus Gnamptogenys was found to be paraphyletic with
respect to Typhlomyrmex, with full support (UFB = 100; LPP = 1)
(Fig. 2, Supp Figs. S1–S5 [online only]) and, consequently, a series
of independent clades within Gnamptogenys are here redefined
as different genera (Fig. 2). The Indomalayan genus Stictoponera
status revived (formerly the coxalis, laevior, and taivanensis groups
of Gnamptogenys sensu Lattke 2004) was recovered as a single
clade with maximum support (UFB = 100; LPP = 1) (Fig. 2, Supp
Figs. S1–S5 [online only]). The genus Poneracantha status revived,
a lineage comprised mainly of species specialized in preying on
myriapods and diplopods (formed mostly by species representing
the rastrata group of Gnamptogenys sensu Lattke 1995), was recovered with maximum support (UFB = 100; LPP = 1) (Fig. 2, Supp
Figs. S1–S5 [online only]). The very distinctive Alfaria status revived
(formerly the minuta group of Gnamptogenys sensu Brandão and
Lattke, 1990) formed a clade also recovered with maximum support
(UFB = 100; LPP = 1) (Fig. 2, Supp Figs. S1–S5 [online only]). Our
analyses also recovered a clade formed by Holcoponera status revived (UFB = 100; LPP = 1) comprising Australasian, Indomalayan,
and Neotropical species (most of the species of the striatula group of
Gnamptogenys sensu Lattke (1995) and the albiclava and epinotalis
groups of Gnamptogenys sensu Lattke (2004)) (Fig. 2, Supp Figs.
S1–S5 [online only]). We recovered, with maximum support, a
monophyletic Typhlomyrmex including two small-sized species formerly assigned to Gnamptogenys (T. reichenspergeri (Santschi) and
T. lavra (Lattke)) (BS = 100; LPP = 1) (Fig. 2, Supp Figs. S1–S5 [online only]). The discovery of this clade is a very surprising result of
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This paper and the nomenclatural act(s) it contains have been registered in Zoobank (www.zoobank.org), the official register of the
International Commission on Zoological Nomenclature. The LSID
(Life Science Identifier) number of the publication is: urn:lsid:zoobank.
org:pub:55B5ECCD-6C6E-4721-B094-ADDC2275CEE6
5
6
Insect Systematics and Diversity, 2022, Vol. 6, No. 1
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Fig. 2. Phylogeny of the subfamily Ectatomminae based on phylogenomic analyses of the UCE 90% complete data set (150 taxa). Figure is based on IQ-Tree besttree searches with ultrafast bootstrap (UFB) frequencies of less than 100% mapped onto the respective nodes. UFB searches consisted of 1000 replicates. The
eleven larger ectatommine lineages are indicated. Branch color indicates the biogeographical range of the species.Taxa marked with asterisk (*) were classified in
Gnamptogenys prior to this revision and those with double asterisk (**) were included in Heteroponera prior to this revision. See Supplementary material for the
75% complete matrix (Supp Fig. S1 [online only]). Ant photos show heads in frontal view of, from top to bottom: Gnamptogenys acuminata (USNMENT00441095),
Typhlomyrmex rogenhoferi (CASENT0004700), Holcoponera striatula (CASENT0106042), Alfaria simulans (CASENT0603729), Poneracantha rastrata
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Insect Systematics and Diversity, 2022, Vol. 6, No. 1
Gene Support for Alternative Topologies
Despite the large dataset used in the present study, some nodes
showed relatively low support or were incongruent between different datasets (see red dots in Fig. 2), which could indicate either
low or conflicting phylogenetic signals. To explore these possibilities,
we looked at the support of gene trees for competing topologies. We
found that, of all 2,520 gene trees, 620 recovered Heteroponera as
monophyletic (including H. monticola) (Supp Fig. S6 [online only],
N1), and 526 recovered H. monticola as sister to a clade formed by
the remaining Heteroponera + Acanthoponera (Supp Fig. S6 [online only], N2). Regarding the sister group relationship between
Rhytidoponera and Ectatomma, 16 gene trees recovered both genera
as sister groups (Supp Fig. S6 [online only], N3), while 31 recovered
Ectatomma as sister to all other Ectatommini genera (Supp Fig. S6
[online only], N4). Lastly, 454 gene trees recovered Holcoponera
as sister to Alfaria (Supp Fig. S6 [online only], N5), while 412 gene
trees recovered Poneracantha as sister to Holcoponera (Supp Fig.
S6 [online only], N6). These results indeed support the existence of
considerable incongruence among loci for those particular nodes.
For each of those problematic nodes, gene trees mostly supported
two alternative topological hypotheses, with significantly less support for other topologies. The favored topologies have significantly
higher (p < 0.05) mean bootstrap values, and the loci supporting
the favored topologies have significantly lower GC content than loci
supporting alternative topologies. But relatively low numbers of loci
support the favored topologies, and in each case the majority of
loci do not support the favored topologies. Among the favored topologies there is nearly equal evidence supporting the alternatives, in
terms of both bootstrap support and GC content.
Discussion
Phylogenomics Resolves Relationships Among
Ectatomminae Genera
Our concatenated and species-tree analyses recovered a well resolved
and highly congruent phylogeny for Ectatomminae, while identifying
possible incongruences that need to be further investigated (Figs. 2,
Supp Figs. S1–S5 [online only]). These results, based on our 2,520
UCE loci dataset, are congruent with prior research that suggests that
having a greater number of loci is beneficial (Borowiec et al. 2015,
Branstetter et al. 2017), although it remains unclear how many loci
are necessary to resolve phylogenetic relationships. However, it has
long been recognized that simply increasing the amount of data can
exacerbate systematic bias in phylogenetic estimation (Phillips et al.
2004, Philippe et al. 2011, Borowiec et al. 2015) and that to improve
phylogenetic inference data quality is key (Borowiec et al. 2015).
We showed that, despite the incongruencies found among different
datasets for some nodes (see red dots in Fig. 2), all alternative topologies are supported by good-quality data with strong phylogenetic
signal. However, recoding nucleotides to RY characters suggests that
composition bias may be contributing to support for nodes where
gene-tree incongruence is pervasive (Supp Fig. S6 [online only]).
RY-coding reduces such biases and increases the signal on internal
branches relative to external, increasing phylogenetic signal in mitochondrial genome data (Phillips and Penny 2003). Nevertheless,
using RY-coding reduces the dataset size and, as shown in Supp Fig.
S6 (online only), nodes that are incongruent between the nucleotide
and RY-character data are supported by relatively few loci, which
may suggest that dataset size may be important for resolving phylogenetic relationships in Ectatomminae. If loci are discordant, it is expected that numerous additional markers are necessary to generate a
robust tree, allowing for an amplification of phylogenetic signal with
the increase of the amount of data (Camacho et al 2019).
Previous research has shown that taxonomic balance within a
data set has a large impact on phylogenetic results (Branstetter et al.
2017), emphasizing the importance of both broad taxonomic sampling (i.e., covering taxonomic disparity and geographic coverage)
and taxonomic evenness across samples (i.e., having comparable
samples sizes among the groups, according to their diversity). The fact
that we recover alternative hypotheses for some nodes may suggest
that a larger sampling of those groups might shed light on their relationships in the future. Despite the fact that our phylogeny includes a
broad representation of Heteroponera, the addition of H. inca to the
phylogeny could help elucidate the position of H. monticola, since
both species seem to be morphologically similar and possibly closely
related. Regarding the relationship among Rhytidoponera and
Ectatomma, even though the 26 species of Rhytidoponera included
(CASENT0281223), Stictoponera biroi (CASENT0281519), Rythidoponera metallica (CASENT0172345), Ectatomma lugens (USNMENT00445341), Heteroponera
brounii (CASENT0172105), Acanthoponera mucronata (CASENT0173540), and Boltonia microps (CASENT0173544). Images by April Nobile, Jeffrey Sosa-Calvo,
Zach Lieberman, Will Ericson, Michael Branstetter, and Estella Ortega; available from www.antweb.org (Antweb 2021).
Downloaded from https://academic.oup.com/isd/article/6/1/5/6514765 by Universidade Federal do Paraná user on 25 January 2022
our study, since these former Gnamptogenys species were thought
to be closely related to Holcoponera (former striatula group sensu
Lattke (1995)) due to their remarkable morphological similarities
shared with other small-sized Holcoponera species (i.e., H. mina
(Brown), H. haytiana (Wheeler and Mann), and H. relicta (Mann)).
The genus Gnamptogenys was recovered as a clade consisting of
species from the sulcata, concinna, and mordax groups (sensu Lattke
1995) with full support (UFB = 100; LPP = 1) (Fig. 2, Supp Figs.
S1–S5 [online only]). The concinna group (UFB = 100; LPP = 1), also
recognized by Lattke (1995), contains two of the largest species in
the genus, G. concinna (Smith) and G. haenschi (Emery). Although
morphologically quite different, the differences are probably due
to microhabitat differences (G. concinna has large eyes and bright
color and is a canopy ant; G. haenschi has small eyes and drab color
and occurs on the ground and in litter samples, and occasionally
under rotten wood). The sulcata and mordax groups (UFB = 100;
LPP = 1) recognized here each contain multiple species and only partially correspond to Lattke’s (1995) concepts for these groups (Fig.
2, Supp Figs. S1–S5 [online only]).
Regarding relationships among genera within what was previously Gnamptogenys, Stictoponera was recovered as sister to all
other lineages in all analyses (UFB = 100, LPP = 1) (Fig. 2, Supp Figs.
S1–S5 [online only]). Gnamptogenys and Typhlomyrmex were recovered as sister groups in all analyses with full support (UFB = 100,
LPP = 1) (Fig. 2, Supp Figs. S1–S5 [online only]). Holcoponera as
sister to Alfaria was recovered by the concatenated (90p-matrix and
75p-matrix) and the 75p-matrix species-tree analyses with full support (UFB = 100; LPP = 1) (Fig. 2, Supp Figs. S2 and S5 [online only]),
and with lower support by the 90p-matrix converted to RY-coding
(UFB = 78; Supp Fig. S1 [online only]) and the 90p-matrix species
tree (LPP = 0.85; Supp Fig. S4 [online only]), which also recovered
Poneracantha as sister to both genera with full support (UFB = 100,
LPP = 1) (Fig. 2, Supp Figs. S1–S5 [online only]). The 75p-matrix
converted to RY-coding, in contrast, recovered Poneracantha as sister
to Holcoponera (UFB = 93), and Alfaria as sister to both genera with
full support (UFB = 100) (Supp Fig. S3 [online only]).
8
in our phylogeny represents a broad sampling for the genus, there
are 104 species currently described and a more complete phylogeny
for the genus could provide increased support. Similarly, a larger
sampling of Holcoponera, Alfaria, and Poneracantha species could
elucidate the relationships among those genera. Nevertheless, our
results recover a robust and fully resolved topology that we discuss
below through an in-depth discussion of the morphological hypotheses available for the group.
We propose taxonomic changes for the subfamily that improve ant
systematics, i.e., by ensuring that formally named taxa are monophyletic, while simultaneously keeping names fairly stable. At the
subfamily level, our decision to synonymize Heteroponerinae
under Ectatomminae is not based on the monophyly of these
groups, since both are reciprocally monophyletic as currently circumscribed, and their sister-group relationship has been broadly
discussed. Historically, the close relationship between both groups
has been supported by morphological (Brown 1958, Bolton 2003,
Ward 2007, Keller 2011) and molecular data (Brady et al. 2006,
Moreau et al. 2006, Moreau and Bell 2013, Branstetter et al. 2017).
However, morphology can be misleading, especially when defining
the diagnostic characters for the groups separately. When describing
Heteroponerinae, Bolton (2003) stated that there is no unequivocal
apomorphy for the subfamily, suggesting a number of characters
that could have this status. Feitosa (2015) investigated the phylogeny
of Heteroponerinae using morphological data, testing the characters suggested by Bolton (2003), as well as by several others, and
also could not identify any apomorphy for the group. However, in
his work, Feitosa included species of Ectatomminae as outgroups
and his analysis suggested at least ten diagnostic characters for the
clade comprising both Ectatomminae and Heteroponerinae. For
this reason, we reclassify all ectaheteromorph ants as members of
a single subfamily, ensuring the monophyly criterion that already
applies to all other ant subfamilies but, most importantly, providing a clear diagnosis for the subfamily based on morphological
synapomorphies.
Regarding taxonomic changes at the tribal level, our aim is to
keep the classification stable. In this sense, the new combination of
the tribe Heteroponerini and the synonymy of Typhlomyrmecini
are made to ensure the correct placement of the former, and the
monophyly of Ectatommini in the case of the latter. At a generic
level within the tribe Heteroponerini, the paraphyly of Heteroponera
is a striking result, unpredicted by morphology, with B. microps appearing as a separately diverging lineage. This result is congruent
with the previous hypotheses of Borgmeier (1957) and Feitosa
(2015), which suggested that the diagnostic characters for this
species are highly divergent from the morphological patterns for
Heteroponera, but its placement as a separate genus is supported
here for the first time. Similarly, the position of H. monticola, recovered as sister to all the other Heteroponera species, as well as the
recovery of two separate clades, the first comprising H. carinifrons
(from Chile) as sister to the Australasian species and the second comprising the remaining Neotropical species, are also entirely new evolutionary hypotheses for the genus, with strong implications for its
biogeographical history.
This phylogenetic scenario suggests that the common ancestor
of Heteroponerini morphologically resembled a modern member of
Acanthoponera, with a relatively large body, prominent spines, well
developed eyes, and long palps. An early lineage probably split off
and evolved to occupy the epigaeic and hypogaeic strata of the environment, maybe displaced by an emerging dominant lineage of ants
(e.g., Myrmicinae). This now cryptic early lineage of heteroponerines
has undergone a drastic reduction of body size, appendages, and eyes,
as we can see in the extant Boltonia. Later, a second divergence event
separated two lineages of Heteroponera and adaptation for living
in the ground was repeated. In this second process, H. monticola
and H. inca retained several plesiomorphic traits, also related to
Acanthoponera, but the remaining Heteroponera gradually lost these
characters as they made their way to the soil and morphologically
converged on Boltonia in the reduction of appendages and body size.
This scenario is supported by the presence of tarsal teeth and lobes
in Acanthoponera, traits strictly related to arboreal habits that were
lost in the remaining lineages of heteroponerine adapted to nesting
and foraging in the ground (Feitosa 2015). Our results regarding relationships among species in Heteroponerini shed new light on the
study of their morphological evolution. We believe that, in order to
ensure the stability of the classification, to best understand the evolution of this group, and to make the most significant contribution
to ant systematics, the assessment of relationships among the species should combine both molecular and morphological approaches.
Unfortunately, the genus Aulacopone was not included in our analysis due to the unavailability of specimens and difficulties of collecting in its type locality. The genus is monotypic and was collected
only twice in the 1920/30s, with the only known specimen currently
metal-coated, making recovery of DNA information from the pinned
specimen a risk to the only specimen available. The distribution of
this genus is singular within the Ectatomminae, being the only group
to occur in the Palearctic region. Aulacopone is said to share several
morphological similarities with the other heteroponerines (Brown
1958, Taylor 1980, Lattke 1994, Bolton 2003, Feitosa 2015), but
its position among the Ectatomminae is still not well defined due to
the impossibility of examining important characters in the previous
phylogenetic study (Feitosa 2015).
The eight genera that comprise Ectatommini are shown to form
a well-supported clade, a result that is congruent with previous
morphological hypotheses for the group (Bolton 2003, Ward 2007,
Keller 2011), although these works considered the four genera as
previously defined. In the molecular phylogenies published so far,
only one or a few specimens of each genus were included, limiting
their conclusions regarding the relationships among them (Brady
et al. 2006, Moreau et al. 2006, Moreau and Bell 2013, Branstetter
et al. 2017). Given these limitations, this is the first molecular
study that aimed to investigate the genus-level relationships in
Ectatomminae. A fairly novel result, the sister-group relationship between Ectatomma and Rhytidoponera, is congruent with previous
morphological hypotheses by Keller (2000, 2011) and was suggested
by other broad-scale molecular phylogenies of Formicidae that did
not focus specifically on these groups (Brady et al. 2006, Moreau
et al. 2006, Moreau and Bell 2013). Brown (1958) noticed some
similarities between the two genera, noting similarities in wing venation and male genitalia and absence of a metacoxal spine (present
in Holcoponera, Gnamptogenys, and Stictoponera). Also, Brown
(1958) called attention to similarities between Ectatomma workers
and those of the largest species of Rhytidoponera. Our results are
the first to include broad species-level representatives of those genera
and our results shed light on the evolution of these groups.
Perhaps the most strikingly novel result in our study is the strong
support for the paraphyly of the former Gnamptogenys in relation to Typhlomyrmex. This result was never previously predicted
by any morphological or molecular study. Historically, the position
of Typhlomyrmex relative to the Ectatommini was first addressed
by Emery (1911), but Brown (1965) later placed the genus in its
own tribe, Typhlomyrmecini, considering it to be closely related to
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Taxonomy of Ectatomminae Revisited
Insect Systematics and Diversity, 2022, Vol. 6, No. 1
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Insect Systematics and Diversity, 2022, Vol. 6, No. 1
Gnamptogenys rastrata group and considered them to be closer to
Holcoponera than to the present definition of Gnamptogenys based
on the presence of triangular mandibles, long and typically sculptured scapes, the convex clypeal lamella, and the well-developed
metacoxal tooth, a result that is also recovered by our molecular
data. Lattke (1995) also recovered the sulcata and mordax groups
as sister groups, with the concinna group as closely related to
them, but not monophyletic. We obtained similar results, except
for the monophyly of the concinna group, and redefine the sulcata,
concinna, and mordax groups as a smaller, strictly Neotropical
Gnamptogenys. Finally, the sister-group relationship between the
species T. reichenspergeri, T. lenis, and T. lavra and the remaining
Typhlomyrmex is a result never predicted by morphology and, in
fact, the phylogenetic distance among those species is similar to the
distance among other genera. Those species have in common absent or reduced eyes, with less than 15 ommatidia; promesonotal
suture well marked, totally interrupting dorsal mesosomal sculpture;
propodeal spiracle separated from declivity margin by a distance
longer than its diameter; metacoxal dorsum unarmed; and petiole
pedunculate. T. reichenspergeri, T. lenis, and T. lavra lack a welldefined antennal club and a prominent anteroventral process on
the petiole. We chose to combine those species into Typhlomyrmex
based on these shared diagnostic characteristics, in the interest of a
more stable classification.
Additional work is necessary because we strongly believe that the
molecular phylogenetic data should be combined with the study of
morphological characters that are diagnostic for the newly defined
genera and for the new generic combinations, so that the final classification can be functional and useful to any researcher studying specimens in the laboratory or in the field. In this study, we demonstrate
that UCE data provide a robust source of phylogenomic data for
the Ectatomminae ants. Morphological evolution, interpreted with
reference to our resulting phylogeny, has produced diagnostic characters for defining taxonomic groups. We believe that the phylogenetic framework and the new classification proposed here provides a
solid foundation for the further study of Ectatomminae taxonomy
and systematics, as well as for reconstructing the morphological evolution of the genera, species groups, and species that it comprises.
Taxonomic Account
In order to erect a phylogenetic classification for the subfamily, with
monophyletic tribes and subfamilies (Ward 2011), we propose a
number of higher-level taxonomic changes. New and revived combinations include the junior synonyms of the species names listed
below. Author and year of publication for all genus and species
names can be found in AntCat (http://antcat.org/). The tribal and
generic classifications of ectaheteromorphs are here modified to
achieve consistency with our molecular phylogenetic results. We
maintain the existing classification as far as possible, while striving
to ensure that all recognized tribes are monophyletic. Genera known
only from fossils are indicated with a dagger; most of these are unplaced to tribe and are treated as incertae sedis within the subfamily.
Ectatomminae Emery
= Heteroponerinae Bolton new synonym
Diagnosis: Presenting the characters of ‘poneromorph’ subfamilies
described by Bolton (2003: p.40). Clypeus broadly inserted between frontal lobes (Bolton 2003); anterior clypeal margin with a
narrow lamellar apron (Bolton 2003). Torulus not completely fused
to frontal lobe (Bolton 2003). Antenna with 12 segments (Bolton
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the Amblyoponini. Lattke (1994) suggested that the similarities of
Typhlomyrmecini with Ectatommini required further exploration
and Bolton (2003) most recently considered the Typhlomyrmecini to
be a member of Ectatomminae. However, rather than forming a separate tribe in Ectatomminae, it now appears that this group of ants is
a highly derived lineage among the former species of Gnamptogenys
with a distinctive cryptic morphology.
The paraphyly of Gnamptogenys in relation to Typhlomyrmex
provided two different alternatives for the taxonomic treatment of
the genera in Ectatomminae, the first being the synonymization of
Gnamptogenys under Typhlomyrmex, since the latter is the oldest
available name. However, we recognize the importance of the name
Gnamptogenys within the myrmecological literature and, with a nomenclatural gender change from feminine (Gnamptogenys) to masculine (Typhlomyrmex) for most species, this would not be the most
parsimonious treatment. The second possibility, chosen here, involved
reviving available names for the different clades recovered in our phylogeny, considering the similar phylogenetic distances between those
clades and between other Ectatomminae genera, and the strong diagnostic morphological characters recovered for each of the lineages.
The availability of generic names for each of those clades shows that
hypotheses for those groups were once presented, but morphological
data were not sufficient to define them at the time, and they were
later synonymized under Gnamptogenys (Brown 1958). With our molecular dataset we recovered each clade with strong support and, by
reciprocal illumination, defined the morphological characters that separate each genus from any other genus in Ectatomminae.
The generic status of Holcoponera, Stictoponera, and Alfaria
were subjects of long and arduous inquiry into the myrmecological
literature since they were first proposed as subgenera of Ectatomma
in the case of the first two, or as a genus, in the case of Alfaria.
Brown (1958) found no basis for maintaining the generic status of
those names, but divided Gnamptogenys into four groups, namely
the Gnamptogenys group, the Stictoponera group, the Holcoponera
group, and the Alfaria group. Brown considered Holcoponera to be
a well-defined genus based on its more compact, dorsally convex
mesosoma with a marked promesonotal suture interrupting the
sculpture and on the form of the petiolar node, as well as by characters of wing venation and larval hairs. However, when analyzing
the similarities between the species Typhlomyrmex reichenspergeri,
Holcoponera relicta, and Holcoponera mina, he considered the lack
of gastric sculpture in T. reichenspergeri as evidence against its placement in a separate genus. In our study, we recovered T. reichenspergeri
as sister to Typhlomyrmex and relatively distant from Holcoponera
and we found that Holcoponera is not a strictly Neotropical genus
because it also includes Indomalayan and Australian species formerly described as Rhopalopone and Wheeleripone. Brown considered it impossible to define the genus Stictoponera because of
dissimilarities among the Old World species. We resolve the problem
by showing that Old World species fall into two independent clades,
one within Holcoponera. Brown considered the genus Alfaria to be
the most distinct of the ectatommine genera but felt that A. striolata
cast doubts on its generic status due to the less inflated second gastric
segment and to its sculpture, which is similar to that of Stictoponera.
Our genomic data, however, show that Alfaria forms a distinct clade
among the Ectatomminae and, even though A. striolata was not included in the phylogeny, the presence of an expanded frontal carina
suggests that this species placement is correct.
The current definition of the genus Poneracantha is a novel result,
as this was proposed as a monotypic subgenus to contain the highly
divergent P. bispinosa. However, Lattke (1995) proposed that the
specialized millipede predators that belong to this genus formed the
10
Tribes: Ectatommini and Heteroponerini
Incertae sedis: †Canapone, †Electroponera, †Pseudectatomma.
Notes: In spite of the reciprocal monophyly of the subfamilies
Ectatomminae and Heteroponerinae, the morphological evidence
strongly suggests that all ectaheteromorph genera could be combined into a single subfamily Ectatomminae, which is the oldest
available name. Ectatomminae, as defined here, presents a combination of 10 diagnostic characters that can be used to differentiate
those ants from any other ant subfamily, making the identification of
those groups more accessible.
Tribe Ectatommini Emery
= Stictoponerini Arnol’di
= Typhlomyrmecini Emery new synonym
Diagnosis (Females): Ectatommine ants of small to large size
(head width 0.44-2.84mm, head length 0.56-3.8mm). Antennal
scrobe usually absent. Eye absent to well-developed (Bolton 2003).
Acetabulum of antennal socket apparatus spherical (Keller 2011);
accessory chamber of antennal socket present (Keller 2011). Labial
palp with two palpomeres (Keller 2011). Promesonotal suture fused
and immobile to complete and flexible (Bolton 2003). Ventral flap
on metapleural gland opening present (Keller 2011). Metacoxal
cavity open (Bolton 2003). Petiolar sternite fused with tergite over its
entire length (except in Rhytidoponera) (Bolton 2003, Keller 2011);
laterotergites of petiole indistinct to absent.
Genera: Alfaria status revived, Ectatomma, Gnamptogenys,
Holcoponera status revived, Poneracantha status revived,
Rhytidoponera, Stictoponera status revived, and Typhlomyrmex.
Alfaria Emery status revived
= Opisthoscyphus Mann new combination
combination, piei new combination, simulans revived combination,
striolata revived combination, and vriesi new combination (and the
junior synonyms soror new combination, carinata revived combination, emeryi revived combination, mus revived combination,
panamensis revived combination, pneodonax new combination,
scabrosus new combination, and bufonis revived combination).
Distribution: Exclusively Neotropical, from southern Mexico to
northern Argentina.
Notes: Alfaria is a very morphologically distinct lineage among the
Ectatommini, given the extreme anterior curvature of the gaster
in profile. In fact, these ants are usually mistakenly identified as
Proceratium Roger, 1863, due to the impressive convergence in this
character. We here resurrect the name Alfaria, firstly proposed by
Emery (1896) and synonymized under Gnamptogenys by Brown
(1958), to comprise the species previously included in the minuta
group of Gnamptogenys sensu Brandão and Lattke (1990). All
Alfaria species can be identified using the work of Camacho et. al.
(2020) under the previous combination in Gnamptogenys.
Ectatomma Smith
Type Species: Ectatomma tuberculatum (Olivier)
Diagnosis (Females): Occipital lobe absent. Antennal club absent.
Palp formula 2,2. Pronotum usually with two or three tubercles.
Mesonotum prominent and clearly differentiated from propodeum,
separated by a deep transverse suture. Promesonotal suture well
marked, interrupting or not the dorsal mesosomal sculpture.
Propodeal spiracle elliptical or slit-shaped and separated from the
declivous face of propodeum by a distance longer than its diameter.
Apex of protibia with a stout seta close to the strigil base; dorsum of
posterior coxa without projections.
Species: brunneum, confine, edentatum, gibbum, goninion, †gracile,
lugens, muticum, opaciventre, parasiticum, permagnum, planidens,
ruidum, suzanae, tuberculatum, and vizottoi.
Distribution: Exclusively found in the New World, from USA
(Texas) to Argentina (Buenos Aires).
Notes: Ectatomma are among the most conspicuous elements of the
ant fauna in Neotropical ecosystems. Currently, the most comprehensive work including an identification key for the species in the
genus is the revision by Kugler and Brown (1982). However, this
work does not include the species Ectatomma parasiticum Feitosa
and Fresneau, in Feitosa et al. (2008), E. suzannae Almeida (1986)
and E. vizottoi Almeida (1987).
Type Species: Alfaria simulans Emery
Diagnosis (Females): Head subquadrate; occipital lobe usually
present; frontal carina broadly expanded laterad; row of stout setae
on base of foretarsus opposite to strigil present; promesonotal suture
absent to lightly impressed, never interrupting dorsal mesosomal
sculpture; petiolar spiracle facing directly ventrad and sunken within
a pit; second gastral (IV abdominal) sternite usually strongly reduced, so that the gaster is directed ventrally and anterad.
Species: caelata new combination, falcifera new combination, fieldi
new combination, minuta revived combination, petiscapa new
Gnamptogenys Roger
= Commateta Santschi
= Emeryella Forel
= Tammoteca Santschi
Type Species: Gnamptogenys sulcata (Smith)
Diagnosis (Females): Head subquadrate to elongate. Mandible
subtriangular to subfalcate. Occipital lobe absent. Antennal club
absent. Palp formula 2,2 to 3,2. Pronotum unarmed and without
tubercles. Promesonotal suture feebly impressed to absent, never
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2003). Pronotum with the humeral corners angled, forming a distinct delimitation between the anterior and lateral margins (Lattke
2004). Antero-ventral angle of pronotum triangular (Feitosa 2015).
Pretarsus without arolium (Lattke 2004). Petiole pedunculate (Keller
2011). Petiolar node as wide or wider than long (Feitosa 2015).
Subpetiolar process very well developed, occupying more than onethird of the ventral portion of the petiolar sternite (Feitosa 2015).
Helcium projecting from about midheight of the anterior face of
abdominal segment III. Abdominal segment IV presclerites separated from the rest of segment by a constriction or slight thickening
(Lattke 1994). Fourth abdominal tergite arched and larger than the
sternite, giving the segment a curved appearance (Keller 2011).
Insect Systematics and Diversity, 2022, Vol. 6, No. 1
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Insect Systematics and Diversity, 2022, Vol. 6, No. 1
interrupting dorsal mesosomal sculpture, sometimes with a small pit
frequently situated medially on a weakly impressed promesonotal
suture. Mesonotum not prominent, forming a continuous line with
the propodeum, separated by a transverse suture. Propodeal spiracle
oval or rounded, separated from the declivous face of propodeum
by a distance longer or shorter than its diameter. Apex of protibia
without a stout seta close to the strigil base; dorsum of posterior
coxae frequently with a lobe or spine.
Distribution: Exclusively found in the New World, from USA
(Texas) to Argentina (Buenos Aires), with one species occurring in
Cuba.
Notes: In the new concept proposed here, Gnamptogenys is now
restricted to the species from the previous sulcata, concinna, and
mordax groups (sensu Lattke 1995), considering that G. sulcata
is the type-species of the genus. All except one of the species of
Gnamptogenys can be identified using the work of Camacho et. al.
(2020). Gnamptogenys rugimala, a newly described species, can be
identified using the paper by Marcineiro and Lattke (2020).
Holcoponera Mayr status revived
= Mictoponera Forel
= Rhopalopone Emery
= Spaniopone Wheeler and Mann
= Wheeleripone Mann
Type Species: Holcoponera striatula (Mayr)
Diagnosis (Females): Head wider posterad than anterad; mandible triangular with striae or rugulae on frontal surface; anterior
clypeal margin convex; scape usually surpassing vertexal margin;
eye slightly behind cephalic midlength; promesonotal suture frequently well marked, totally interrupting dorsal mesosomal sculpture; propodeal spiracle close to the declivous face of propodeum;
propodeum unarmed; anterior prosternal process broadly concave
medially; metacoxal dorsum always with a denticle or lobe; petiolar
node high; anteroventral postpetiolar process relatively wide; second
gastric segment only slightly arched ventrally.
Species: acuta revived combination, albiclava new combination,
ammophila new combination, andina new combination, aspera new
combination, aterrima new combination, atrata new combination,
auricula new combination, avus new combination, bisulca new combination, brunnea new combination, crenaticeps new combination,
cribrata new combination, dichotoma new combination, ejuncida
new combination, epinotalis new combination, extra new combination, gentryi new combination, gracilis revived combination,
haytiana new combination, ilimani new combination, latistriata
new combination, lucida new combination, luzonensis new combination, major new combination, malaensis new combination, mina
revived combination, moelleri revived combination, nigrivitrea new
combination, pernambucana revived combination, pilosa new combination, pittieri new combination, pleurodon revived combination,
Distribution: Neotropical, Indomalayan, and Australasian.
Notes: In the new classification proposed here the available name
Holcoponera is resurrected from synonymy under Gnamptogenys to
include most of the species of the striatula group sensu Lattke (1995),
and the albiclava and epinotalis groups sensu Lattke (2004). The only
species from the former striatula group not included in Holcoponera
are lavra, lenis, and reichenspergeri, which were transferred to
Typhlomyrmex in this study. The Neotropical species of Holcoponera
can be identified using the work of Camacho et. al. (2020). Oriental
species can be identified using the key in Lattke (2004).
Poneracantha Emery status revived
= Barbourella Wheeler
= Parectatomma Emery
Type Species: Poneracantha bispinosa (Emery)
Diagnosis (Females): Head subquadrate or wider anterad than
posterad in frontal view; anterior clypeal margin usually straight;
frontal surface of mandible usually striate or rugulose; scape usually
surpassing vertex; promesonotal suture feebly impressed to absent,
never interrupting dorsal mesosomal sculpture; metanotal suture
well impressed; propodeum usually armed with denticles or spines;
petiolar node low; subpetiolar process shape variable, usually projecting anterad but sometimes subquadrate; metacoxal teeth generally present, usually acicular; second gastric segment slightly arched
ventrally.
Species: banksi new combination, bispinosa revived combination,
†brunoi, cuneiforma new combination, enodis new combination,
ingeborgae new combination, insularis new combination, lanei new
combination, laticephala new combination, lineolata new combination, lucaris new combination, mecotyle new combination, mediatrix new combination, menozzii revived combination, perspicax
new combination, rastrata new combination, semiferox new combination, triangularis new combination, and wilsoni new combination (and the junior synonyms schubarti new combination, trigona
new combination, aculeaticoxae new combination, and triangularis
richteri new combination).
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Species: acuminata, alfaroi, andersoni, annulata, biquetra,
boliviensis, bruchi, †casca, concinna, continua, curvoclypeata,
ericae, †europaea, falcaria, fernandezi, flava, haenschi, hartmani,
horni, interrupta, kempfi, †levinates, lucaris, mordax, nana, †pristina, regularis, rimulosa, † rohdendorfi, rugimala, rumba, schmitti,
siapensis, stellae, sulcata, tortuolosa, transversa, and volcano.
porcata revived combination, preciosa new combination, relicta
new combination, sila new combination, solomonensis new combination, striatula new combination, and strigata new combination
(and the junior synonyms dammermani new combination, diehlii
new combination, moelleri splendens revived combination, teffensis
revived combination, teffensis concinna revived combination, emeryi
revived combination, vidua revived combination, magnifica revived
combination, striatula angustipleura new combination, arcuata revived combination, brasiliensis revived combination, brasiliensis
calcarata revived combination, brasiliensis mayri revived combination, curtula revived combination, curutlum paulina new combination, curtulum stolli new combination, curtulum vollenweideri
revived combination, emeryi recta revived combination, rustica
revived combination, simplicoides revived combination, striatula
angustiloba new combination, striatula antillana revived combination, striatula obscura new combination, wasmanni revived
combination, wasmanni isthmica revived combination, wheeleri
revived combination, concentrica new combination, satzgeri new
combination, simplex revived combination, simplex spurium new
combination).
12
Distribution: Exclusively Neotropical, occurring from Guatemala to
Uruguay, and in the Caribbean islands of Hispaniola and Lesser Antilles.
Notes: Here we revive the name Poneracantha from synonymy
under Gnamptogenys to include all the species representing the previous rastrata group sensu Lattke (1995). All Poneracantha species
can be identified using the work of Camacho et. al. (2020) under the
previous combination in Gnamptogenys.
Type Species: Rhytidoponera araneoides (Le Guillou)
Diagnosis (Females): Occipital lobe frequently present. Antennal club
absent. Palp formula 2,2 to 3,2. Pronotum unarmed. Mesonotum
not prominent, forming a continuous line with the propodeum, separated by a transverse suture. Promesonotal suture well marked, totally interrupting dorsal mesosomal sculpture. Propodeal spiracle oval
or rounded, separated from the declivous face of propodeum by a
distance longer than its diameter. Apex of protibia with a stout seta
close to the strigil base; dorsum of posterior coxa without projections.
Species: abdominalis, acanthoponeroides, aciculata, aenescens,
anceps, aquila, araneoides, arborea, aspera, atropurpurea, aurata,
barnardi, barretti, borealis, carinata, celtinodis, cerastes, chalybaea,
chnoopyx, clarki, confusa, convexa, cornuta, crassinodis, cristata,
croesus, depilis, dubia, enigmatica, eremita, ferruginea, flavicornis,
flavipes, flindersi, foreli, foveolata, fulgens, fuliginosa, †gibsoni,
greavesi, gregoryi, haeckeli, hanieli, hilli, impressa, incisa, inops,
inornata, insularis, †kirghizorum, koumensis, kurandensis, laciniosa,
lamellinodis, laticeps, levior, litoralis, luteipes, maledicta, maniae,
mayri, metallica, micans, mimica, mirabilis, nexa, nitida, nitidiventris,
nodifera, nudata, numeensis, opaciventris, peninsularis, pilosula,
pulchella, punctata, punctigera, punctiventris, purpurea, reflexa,
reticulata, rotundiceps, rufescens, rufithorax, rufiventris, rufonigra,
scaberrima, scabra, scabrior, socrus, spoliata, strigosa, subcyanea,
tasmaniensis, taurus, tenuis, terrestris, trachypyx, turneri, tyloxys,
versicolor, victoriae, violacea, viridis, †waipiata, wilsoni, and yorkensis.
Distribution: Exclusively Australasian.
Notes: This speciose ectatommine genus could be considered an ecological equivalent of Ectatomma in the Australian region. The most
recent taxonomic tools for the identification of Rhytidoponera species include the papers by Ward (1980, 1984) and Heterick (2009).
Species. bicolor revived combination, biloba new combination,
binghamii revived combination, biroi revived combination, bulbopila
new combination, chapmani new combination, coccina new combination, coxalis revived combination, crassicornis revived combination, delta new combination, dentihumera new combination,
fistulosa new combination, fontana new combination, gabata new
combination, gastrodeia new combination, grammodes new combination, helisa new combination, hyalina new combination, lacunosa
new combination, laevior revived combination, leiolabia new combination, macretes new combination, meghalaya new combination,
menadensis revived combination, nanlingensis new combination,
niuguinensis new combination, ortostoma new combination,
palamala new combination, panda revived combination, paso new
combination, pertusa new combination, polytreta new combination,
posteropsis revived combination, quadrutinodules new combination,
rugodens new combination, scalpta new combination, sichuanensis
new combination, sinensis new combination, sinhala new combination, taivanensis revived combination, toronates new combination, and treta new combination (and the junior synonyms banana
new combination, bicolor minor new combination, borneensis revived combination, costata revived combination, costata pinealis revived combination, costata simalurensis revived combination, costata
unicolor revived combination, laevior avia revived combination,
kalabit new combination, menadensis obscura revived combination,
parva revived combination, rugosa wallacei revived combination,
spiralis revived combination, and stylata revived combination)
Distribution: Oriental region, into South-East Asia, including
southern China, covering the Sundas and Melanesia all the way to
Fiji, including the Philippines.
Notes: Our phylogenomic results suggest that the Indomalayan
Stictoponera species represent a separate evolutionary lineage,
not strictly related to the other Australasian lineages in the subfamily. We here resurrect the name Stictoponera from synonymy
under Gnamptogenys in order to accommodate the species previously included in the coxalis, laevior, and taivanensis groups
of Gnamptogenys (Lattke 2004, Chen et al. 2017). These species comprise a well-supported clade forming the sister group of
Gnamptogenys in the new sense. Its placement as sister to the former
Gnamptogenys is congruent with Lattke (2004), who predicted it
based on morphological features and morphological phylogenetic
analysis.
Typhlomyrmex Mayr
Type Species: Typhlomyrmex rogenhoferi Mayr
Stictoponera Mayr status revived
Type Species: Stictoponera coxalis (Roger)
Diagnosis (Females): Occipital lobe present. Antennal club absent.
Palp formula 3,2. Pronotum usually unarmed, occasionally with
humeral projections. Mesonotum not prominent, forming a continuous line with the propodeum, separated by a transverse suture.
Promesonotal suture absent to feebly impressed, never interrupting
the dorsal mesosomal sculpture. Propodeal spiracle oval to rounded
and separated from the declivous face of propodeum by a distance
longer than its diameter. Apex of protibia without a stout seta close
to the strigil base; apex of meso- and metatibia with two spurs;
dorsum of posterior coxae frequently with a lobe or spine.
Diagnosis (Females): Head subquadrate; antennal club sometimes
well-defined and formed by 3 or 4 segments; cephalic vertex mostly
smooth and shining, sometimes presenting faded striae or rugulae;
eye absent or reduced, with less than 15 ommatidia; promesonotal
suture well marked, totally interrupting dorsal mesosomal sculpture;
propodeal spiracle separated from declivity margin by a distance
longer than its diameter; metacoxal dorsum unarmed or at most
with a small lobe or denticle; petiole pedunculate, sometimes with a
prominent anteroventral process.
Species: clavicornis, foreli, lavra new combination, lenis new combination, major, meire, prolatus, pusillus, reichenspergeri new combination, and rogenhoferi.
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Rhytidoponera Mayr
= Chalcoponera Emery
Insect Systematics and Diversity, 2022, Vol. 6, No. 1
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Distribution: Exclusively Neotropical, occurring from Mexico to
Argentina (Buenos Aires).
Genera: Acanthoponera,
Heteroponera.
Notes: Most species of Typhlomyrmex can be identified using the key
of Lacau et al. (2008), while T. lavra, T. lenis, and T. reichenspergeri
(formerly included in the striatula group of Gnamptogenys) can be
identified using Camacho et al. (2020).
Acanthoponera Mayr
Tribe Heteroponerini Bolton new combination
Boltonia
new
genus,
Type Species Acanthoponera mucronata (Roger)
Diagnosis (Females): Ants of comparatively medium size (head
width 0.90–1.61, head length 1.00–1.75). Mandible triangular.
Palp formula 6,4. Frontal lobe reduced, only partially covering
antennal insertions. Antennal club with four antennomeres.
Antennal scrobe deeply impressed. Eye well-developed, with
clear limits between ommatidia. Propodeum with a pair of welldeveloped spines. Tarsal claw with conspicuous preapical teeth
and a basal lobe. Petiole with a long posterodorsal projection.
Anterior face of abdominal segment III with an arched carina
above the helcium.
Species: goeldii, minor, mucronata, and peruviana.
Distribution: Exclusively Neotropical, from southern Mexico to
northern Argentina.
Fig. 3. Worker of Boltonia microps in A) frontal view; B) dorsal view; and C) lateral view. Images by April Nobile (CASENT0173544); available from www.antweb.
org (Antweb 2021).
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Diagnosis: Ectatommine ants of small to medium size (head width
0.42–1.61 mm, head length 0.53–1.75 mm); cephalic dorsum with
a longitudinal carina extending from anterior margin of clypeus to
posterior margin of head (Bolton 2003); antennal scrobe present
(Bolton 2003); eye present; acetabulum of antennal socket apparatus
hemispherical (Keller 2011); accessory chamber of antennal socket
absent (Keller 2011); labial palp with three or four palpomeres
(Bolton 2003, Keller 2011); promesonotal suture complete and flexible (Bolton 2003); ventral flap on metapleural gland opening absent (Keller 2011); metacoxal cavity closed (Bolton 2003); petiolar
sternite articulated with tergite over its entire length (Bolton 2003,
Keller 2011); laterotergites of petiole present.
Aulacopone,
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Insect Systematics and Diversity, 2022, Vol. 6, No. 1
Notes: Acanthoponera represents a lineage of arboreal and nocturnal Neotropical ants. Most species can be identified using the
work of Feitosa and Prada-Achiardi (2019) for the Colombian
fauna.
ommatidia. Propodeum unarmed. Tarsal claw simple, without conspicuous preapical teeth nor a basal lobe. Petiole unarmed. Anterior
face of abdominal segment III without an arched carina above the
helcium.
Aulacopone Arnol’di
Species: microps.
Type Species: Aulacopone relicta Arnol'di
Distribution: Exclusively Neotropical, from Costa Rica to northern
Argentina and southern Brazil.
Species: relicta.
Distribution: The only known specimens were collected in
Azerbaijan in mountainous forests.
Notes: The genus is only known from two queens collected in
Azerbaijan. The first specimen was collected in 1929 in Alazapin
on the border with Iran, and later designated as the holotype by
Arnol’di and deposited at the Zoological Institute of the Russian
Academy of Sciences. The second specimen was collected in 1936,
also by Arnol’di in the same country, in the region of Khachmaz,
and later deposited in his personal collection at the Institute of
Evolutionary Animal Morphology in Moscow. However, the holotype has been missing since 1979 and has not been examined for any
study other than the original description. The second specimen was
coated in gold-palladium for the study of its external morphology
using scanning electron microscopy by Taylor (1980).
Notes: We here propose the new genus Boltonia to accommodate
a single species, B. microps (Borgmeier 1957), formerly a member
of Heteroponera. This species represents a divergent lineage at the
base of Heteroponerini and is the sister-group of all the remaining
species in the tribe. The genus name is an homage to Barry Bolton,
legendary ant taxonomist and author of Bolton’s Catalogue of Ants
of the World, which is the very foundation of all taxonomic papers
published in myrmecology since 1994.
Zoobank LSID: urn:lsid:zoobank.org:act:7F2A5019-E237-47758CE1-CCE4E124431C
Heteroponera Mayr
= Anacanthoponera Wheeler
= Paranomopone Wheeler
Type Species: Heteroponera carinifrons Mayr
Type Species: Boltonia microps (Borgmeier) new combinationtion
(Fig. 3)
Diagnosis (Females): Ants of comparatively small to medium size
(head width 0.63-1.37; head length 0.72-1.54). Mandible triangular.
Palp formula 3,2 to 4,3. Frontal lobe expanded, completely covering
antennal insertions. Antennal club with three antennomeres.
Antennal scrobe shallowly to deeply impressed. Eye well-developed
to reduced, with clear limits between ommatidia. Propodeal spine
absent to well-developed. Tarsal claw simple, without conspicuous
preapical teeth (except in H. dolo and H. robusta) nor a basal lobe.
Petiole with or without posterodorsal projections. Anterior face of
abdominal segment III with an arched carina above the helcium.
Diagnosis (Females): Ants of comparatively small size (head width
0.42-0.53, head length 0.53-0.67). Mandible subfalcate. Palp
formula 3,2. Frontal lobe expanded, completely covering antennal
insertions. Antennal club with three antennomeres. Antennal scrobe
absent. Eye drastically reduced, without conspicuous limits between
Species: angulata, brounii, carinifrons, crozieri, darlingtonorum,
dentinodis, dolo, ecarinata, flava, georgesi, imbellis, inca, inermis,
leae, lioprocta, majeri, mayri, monteithi, monticola, panamensis,
pendergrasti, relicta, rhodopygea, robusta, trachypyx, viviennae, and
wilsoni.
Boltonia Camacho and Feitosa new genus
Fig. 4. Dorsal view of head, showing: A) Cephalic median longitudinal carina present, extending from the anterior clypeal margin to the vertex (Acanthoponera
minor—CASENT0178699); B) Cephalic median longitudinal carina not extending from the anterior clypeal margin to the vertex (Ectatomma tuberculatum—
CASENT0173380); C) Cephalic median longitudinal carina absent (Holcoponera striatula—CASENT0173386). Photos by April Nobile; available from www.
antweb.org (Antweb 2021).
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Diagnosis (Queens): Ants of comparatively medium size. Mandibles
subfalcate. Median portion of clypeus modified, raised as a short,
blunt triangular point projecting from the antennal insertions to the
mandible. Frontal lobe expanded, extending from the clypeal posterior margin to the vertex. Antennal scrobe wide and deep.
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Insect Systematics and Diversity, 2022, Vol. 6, No. 1
Distribution: Neotropical, from Nicaragua to southern Chile; and
Australian, including New Zealand.
Notes: Identification tools for the species of Heteroponera include
the works of Feitosa and Prada-Achiardi (2019) for the Colombian
fauna and Taylor (2011; 2015) for Australian groups.
Key to the Ectatomminae Genera
-
Cephalic median longitudinal carina absent or not extending from the anterior clypeal margin to the vertex (Fig.
2(1). Median portion of clypeus modified, raised as a short, blunt
triangular point projecting from the antennal insertions to the
mandible. Antennal scrobe wide and very deep (exclusively
Paleartic) (known only by queens) ..................... Aulacopone
-
Median portion of clypeus not raised, not or only to a small
extent covering the mandible. Antennal scrobe deep to absent
............................................................................................ 3
3(2). Tarsal claws with a prominent basal lobe and a long preapical
tooth. Propodeum armed with prominent spines (exclusively
Neotropical) .................................................. Acanthoponera
Fig. 5. Lateral view of pronotum, showing: A) Pronotal tubercles present; mesonotum prominent, separated from propodeum by a deep transversal suture
(Ectatomma tuberculatum—CASENT0173380); B) Pronotal tubercles or projections absent; mesonotum not prominent, forming a continuous profile with
propodeum (Holcoponera striatula—CASENT0173386). Photos by April Nobile; available from www.antweb.org (Antweb 2021).
Fig. 6. Dorsal view of pronotum, showing: A) Pronotum and mesonotum separated by a distinct suture (Rhytidoponera abdominalis—CASENT0281333); B)
Pronotum and mesonotum continuous with a discrete groove (Gnamptogenys stellae—CASENT0281227). Photos by Cerise Chen (A) and Estella Ortega (B)
available from www.antweb.org (Antweb 2021).
Fig. 7. Frontal view of head, showing: A) Expanded frontal lobes (Alfaria falcifera—CASENT0179971); B) Occipital lobes absent (Gnamptogenys continua—
CASENT0173383). Photos by Erin Prado (A) and April Nobile (B); available from www.antweb.org (Antweb 2021).
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1. Cephalic median longitudinal carina present, extending from
the anterior clypeal margin to the vertex (Fig. 4A). Metapleural
gland orifice simple, directed posteriorly or laterally (tribe
Heteroponerini) .................................................................. 2
4B, C). Metapleural gland orifice forming an oblique curved
slit bounded below by a convex rim of cuticule that directs
the orifice dorsally to posterodorsally (tribe Ectatommini)
..................................................................................... 5
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-
Tarsal claws simple, without a prominent basal lobe or
preapical tooth. Propodeum generally angled or with small
rhomboidal teeth at most .................................................... 4
4(3). In frontal view, mandible subfalcate, with around four teeth
on the masticatory margin. Antennal scrobe absent. Eye drastically reduced, without conspicuous limits between ommatidia (exclusively Neotropical) ................................ Boltonia
-
-
Pronotum unarmed and without tubercles. Mesonotum not
prominent, forming a continuous profile with the propodeum
(Fig. 5B). Apex of anterior tibia in outer lateral view without
a seta close to the spur base; if seta present, then species distribution is exclusively Australasian ....................................... 6
6(5). In dorsal view, pronotum and mesonotum always separated
by a distinct suture, so that each tergite forms a separate plate
Fig. 8. Lateral view of gaster, showing: A) Second gastral (IV abdominal) sternite not strongly reduced in relation to the tergite; dorsal profile of gaster gently
convex, so that the apex of gaster is only discretely directed ventrally (Gnamptogenys acuminata—USNMENT00441095); B) Second gastral (IV abdominal)
sternite strongly reduced in relation to the tergite; dorsal profile of gaster extremely convex, so that the gaster is strongly directed ventrally and anterad (Alfaria
minuta—CASENT0281213). Photos by Jeffrey Sosa-Calvo (A) and Estella Ortega (B); available from www.antweb.org (Antweb 2021).
Fig. 9. Dorsal view of mesosoma, showing: A) Promesonotal suture absent (Gnamptogenys acuminata—USNMENT00441095); B) Promesonotal suture feeble,
never interrupting dorsal mesosomal sculpture (Poneracantha banksi—INBIOCRI001281007); C) Promesonotal suture well marked, totally interrupting dorsal
mesosomal sculpture (Holcoponera moelleri—CASENT0173384). Photos by Jeffrey Sosa-Calvo (A), Estella Ortega (B), and April Nobile (C); available from www.
antweb.org (Antweb 2021).
Fig. 10. Lateral view of gaster, showing: A) Second gastric segment (IV abdominal) relatively straight (Gnamptogenys acuminata—USNMENT00441095);
B) Second gastric segment (IV abdominal) slightly arched ventrally (Poneracantha mecotyle—CASENT0281530). Photos by Jeffrey Sosa-Calvo (A) and Zach
Lieberman (B); available from www.antweb.org (Antweb 2021).
Downloaded from https://academic.oup.com/isd/article/6/1/5/6514765 by Universidade Federal do Paraná user on 25 January 2022
In frontal view, mandible subtriangular, with six to eight
teeth on the masticatory margin. Antennal scrobe shallowly
to deeply impressed. Eye well-developed, with clear limits between ommatidia (Neotropical and Australian) .....................
........................................................................ Heteroponera
5(1). Pronotum usually with 2 or 3 tubercles. Mesonotum prominent and clearly differentiated from propodeum, separated
by a deep transverse suture (Fig. 5A). Apex of anterior tibia
in outer lateral view with a seta close to the spur base (exclusively Neotropical) .............................................. Ectatomma
17
Insect Systematics and Diversity, 2022, Vol. 6, No. 1
(Fig. 6A). Dorsum of posterior coxa never with a lobe or
spine. Petiolar sternite tightly attached but never fused to the
tergite (exclusively Australasian) ................... Rhytidoponera
-
In dorsal view, pronotum and mesonotum usually continuous
or separated by a discrete groove (Fig. 6B); if a well-impressed
promesonotal suture is present (as in Holcoponera and
Typhlomyrmex), then the dorsum of posterior coxa frequently
with a lobe or spine. Petiolar sternite fused with tergite over its
entire length ........................................................................ 7
7(6). Frontal lobes strongly expanded, entirely covering the antennal insertions (Fig. 7A). Second gastral (IV abdominal)
sternite strongly reduced in relation to the tergite; dorsal
profile of gaster extremely convex, so that the gaster is usually
strongly directed ventrally and anterad (Fig. 8A). Exclusively
Neotropical. .............................................................. Alfaria
-
Frontal lobes less developed, only partially covering the antennal insertions (Fig. 7B). Second gastral (IV abdominal)
sternite not strongly reduced in relation to the tergite; dorsal
profile of gaster gently convex, so that the apex of gaster is
directed ventrally or posteriorly (Fig. 8B). .......................... 8
8(7). Promesonotal suture feebly impressed to absent, never
interrupting dorsal mesosomal sculpture (Fig. 9A, B) ..........9
-
Promesonotal suture well marked, totally interrupting dorsal
mesosomal sculpture (Fig. 9C) .......................................... 11
Strictly Indomalayan species .......................... Stictoponera
9(8).
-
Strictly Neotropical species ........................................... 10
10(9). Propodeum rarely armed with denticles or spines. Metacoxal
teeth present or absent. Second gastric segment (IV abdominal) relatively less curved (Fig. 10A) ......... Gnamptogenys
-
Propodeum usually armed with denticles or spines. Metacoxal
teeth generally present. Second gastric segment (IV abdominal) relatively more curved (Fig. 10B) ........... Poneracantha
11(8). Eye absent or reduced. Propodeal spiracle separated from
posterior face of propodeum by a distance longer than its
diameter (Fig. 11A). Metacoxal dorsum unarmed or at most
with a small lobe or denticle ...................... Typhlomyrmex
-
Eye well developed to reduced. Propodeal spiracle close to
posterior face of propodeum (Fig. 11B). Metacoxal dorsum
always with a denticle or lobe ....................... Holcoponera
Supplementary Data
Supplementary data are available at Insect Systematics and Diversity
online.
Acknowledgments
This work would not have been possible without the myrmecological collections supported at the Museu de Zoologia da Universidade de São Paulo, the
Comissão Executiva do Plano da Lavoura Cacaueira, the Instituto Nacional de
Pesquisas da Amazônia, and the Coleção Entomológica Padre Jesus Santiago
Moure. This work was also made possible by the infrastructure provided by
the Departamento de Zoologia at the Universidade Federal do Paraná, Brazil.
The important scientific contributions proposed here were only made possible
by consistent and effective funding of those institutions. In addition to our
gratitude to those institutions, we would also like to thank Antweb.org for
making images of the type and non-type specimens available for this study.
We thank Patrícia Stroher, André Olivotto, Bonnie Blaimer, Ana Jesovnik, Jeffrey Sosa-Calvo, Mike Lloyd, Dietrich Gotzek, and Eugenia Okonski for assistance with lab work, bioinformatic analyses, and logistics; Alan Andersen
for the donation of specimens of Rhytidoponera; Milan Janda for the donation of specimens of Gnamptogenys from Asia; and Stefan Cover and David
Lubertazzi for access to the MCZ collection. This research was supported by
the Partnerships for Enhanced Engagement in Research (PEER) Science project
3-188, Cycle 3, funded by the U.S. National Academies of Sciences, Engineering and Medicine and the United States Agency for International Development
(USAID). GPC, WF, and RMF were supported by the Conselho Nacional de
Desenvolvimento Científico e Tecnológico (CNPq) (PhD. Fellowship and grants
140338/2014-4, 141234/2018-0, and 301495/2019-0, respectively). MGB and
JTL were supported by National Science Foundation grants DEB-1354996 and
1932405. TRS was supported by U.S. National Science Foundation grants DEB
1456964, 1654829, and 1927161. GPC was also supported by the Peter and
Carmen Buck Pre-doctoral fellowship at the National Museum of Natural History, Smithsonian Institution. USDA is an equal opportunity provider and employer. The authors declare that there are no conflicts of interest.
Author Contributions
GPC: Conceptualization; Data curation; Formal Analysis; Funding acquisition; Investigation; Methodology; Project administration; Software; Validation; Visualization; Writing – original draft; Writing – review & editing. WF:
Data curation; Formal analysis; Investigation; Writing - review & editing.
MGB: Data curation; Formal analysis; Investigation; Software; Writing – review & editing. MRP: Conceptualization; Resources; Supervision; Writing
– review & editing. JTL: Resources; Formal analysis; Funding acquisition;
Writing – review & editing. TRS: Funding acquisition; Resources; Supervision; Writing – review & editing. RMF: Conceptualization; Data curation;
Formal analysis; Funding acquisition; Investigation; Methodology; Resources;
Supervision; Writing – review & editing.
Downloaded from https://academic.oup.com/isd/article/6/1/5/6514765 by Universidade Federal do Paraná user on 25 January 2022
Fig. 11. Lateral view of propodeum, showing: A) Propodeal spiracle separated from declivity margin by a distance longer than its diameter (Typhlomyrmex
lavra); B) Propodeal spiracle close to the declivous face of propodeum (Holcoponera relicta—USNMENT00412058). Photos by Gabriela Camacho (A) and Jeffrey
Sosa-Calvo; available from www.antweb.org (Antweb 2021).
18
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