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Selected species of Camponotus (Figs. 14–17) and New Caledonia Colobopsis (Figs. 18–19), full-face view of head of minor worker. 14, Camponotus (Pseudocolobopsis) claviscapus, Costa Rica (CASENT0249388); 15, C. (Myrmentoma) decipiens, Texas, USA (CASENT0249367); 16. C. (Tanaemyrmex) chloroticus, Fiji (CASENT0171139); 17, C. (Myrmamblys) pulchellus_cf, New Caledonia (CASENT0280237); 18, Colobopsis indet. (CASENT0280248); 19, Colobopsis camela (CASENT0280242). Images from AntWeb (www.antweb.org); photographers Will Ericson (14–15), Eli Sarnat (16) and Shannon Hartman (17–19).
Source publication
The classification of the ant subfamily Formicinae is revised to reflect findings from a recent molecular phylogenetic study and complementary morphological investigations. The existing classification is maintained as far as possible, but some tribes and genera are redefined to ensure monophyly. Eleven tribes are recognized, all of which are strong...
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For a long time, New Caledonia was considered a continental island, a fragment of Gondwana harbouring old clades that originated by vicariance and so were thought to be locally ancient. Recent molecular phylogenetic studies dating diversification and geological data indicating important events of submergence during the Paleocene and Eocene (until 3...
Citations
... We also provide detailed annotations for our synopsis of fossil Camponotini (see the "Notes"). Finally, we point out that all future studies on fossils that may possibly be associated with Camponotus or Camponotini should critically evaluate the morphological evidence for placement in any of the extant genera particularly in reference to Ward et al. (2016) for workers and Ward and Boudinot (2021) for workers and alates (the wing venation characters apply equally to males and queens). A recent work which ignored these studies is Takahashi and Aiba (2023), which misidentified multiple specimens as Camponotus. ...
... Note 2. Camponotus and the tribe Camponotini more broadly is one of the most challenging taxonomic puzzles in the Formicidae, and not merely due to the massive size of these taxa (1084 valid species and 411 valid subspecies are currently attributed to Camponotus at the date of writing, Bolton 2023). Although some genera in the tribe are reasonably identifiable based on external morphology (e.g., Ward et al. 2016), others, such as the fundamental distinction between Colobopsis-which is sister to all other Camponotini-and the hyperdiverse Camponotus is challenging even with extant material in hand and under the microscope (Ward and Boudinot 2021). For these reasons, we substantially revise the fossil system of Camponotus in order to meet the twin aims of: (1) cleaning up the useless species names attributed to Camponotus, and (2) discouraging uncritical use of these fossils for macroevolutionary analysis (e.g., Klimeš et al. 2022). ...
... All of these fossils could be considered unidentifiable to species, hence invalid, but are here treated as incertae sedis in Camponotus to highlight their existence. Critically, because of the lack of morphological information, it is possible that a number of these taxa belong to other genera of Camponotini (see Ward et al. 2016 and. Reexamination of the original material is necessary in all cases. ...
As the only direct records of the history of evolution, it is critical to determine the geological source of biota-bearing fossils. Through the application of synchrotron-radiation micro-computed tomography (SR-µ-CT), Fourier-transformed infrared-spectroscopy (FT-IR), visual evaluation of ultraviolet fluorescence (UV-VS), radiocarbon dating (¹⁴C quantification), and historical sleuthing, we were able to identify and sort 161 (83 Baltic amber, 71 Copal and 7 Kauri gum pieces) individually numbered and largely mislabeled pieces of East African Defaunation resin (~145 years old) and copal (~390 years old), as well as Baltic amber (~35 million years old) from the Phyletisches Museum collection. Based on this collection, we define two new species: ‡Amphientomum knorrei Weingardt, Bock & Boudinot, sp. nov. (Psocodea: Amphientomidae, copal) and †Baltistena nigrispinata Batelka, Tröger & Bock, sp. nov. (Coleoptera: Mordellidae, Baltic amber). For selected taxa, we provide systematic reviews of the fossil record, including: Amphientomidae, for which we provide a key to all species of Amphientomum, extant and extinct, and recognize the junior synonymy of Am. ectostriolatum Li, 2002 (an unjustified emendation) under Am. ectostriolate Li, 1999 (syn. nov.); the fossil ant genus †Yantaromyrmex and the clades Dorylinae, Plagiolepidini, Camponotus, Crematogaster, and Pheidole (Formicidae); the Nevrorthidae (Neuroptera); and Doliopygus (Coleoptera: Curculionidae: Platypodinae). We synonymize Palaeoseopsis Enderlein, 1925 with Amphientomum Pictet, 1854, syn. nov. and transfer one species from Amphientomum, forming Lithoseopsis indentatum (Turner, 1975), comb. nov. To prevent the uncritical usage of unidentifiable fossils attributed to Camponotus for macroevolutionary analysis, we transfer 29 species to the form genus †Camponotites Steinbach, 1967, which we consider to be most useful as incertae sedis in the Formicinae. We treat †Ctt. ullrichi (Bachmayer, 1960), comb. nov. as unidentifiable hence invalid stat. nov. We also transfer †Ca. mengei Mayr, 1868 and its junior synonym †Ca. igneus Mayr, 1868 to a new genus, †Eocamponotus Boudinot, gen. nov., which is incertae sedis in the Camponotini. Concluding our revision of Camponotus fossils, we transfer †Ca. palaeopterus (Zhang, 1989) to Liometopum (Dolichoderinae), resulting in †L. palaeopterumcomb. nov. and the junior synonymy of †Shanwangella Zhang, 1989, syn. nov. under Liometopum Mayr, 1861. Because the type specimens of the genera †Palaeosminthurus Pierce & Gibron, 1962, stat. rev. and †Pseudocamponotus Carpenter, 1930 are unidentifiable due to poor preservation, we consider these taxa unidentifiable hence invalid stat. nov. To avoid unsupported use of the available fossils names attributed to Crematogaster for divergence dating calibration points, we transfer three species to a new collective taxon that is incertae sedis in Myrmicinae, †Incertogaster Boudinot, gen. nov., forming †In. aurora (LaPolla & Greenwalt, 2015), †In. praecursor (Emery, 1891), comb. nov., and †In. primitiva (Radchenko & Dlussky, 2019), comb. nov. Finally, we transfer †Ph. cordata (Holl, 1829) back to Pheidole, and designate a neotype from our copal collection based on all available evidence. All new species plus the neotype of ‡Ph. cordata are depicted with 3D cybertypes from our µ-CT scan data. We introduce the convention of a double dagger symbol (‡) to indicate fossils in copal or Defaunation resin, as these may yet be extant. To further contextualize our results, we provide a discussion of amber history and classification, as well as the Kleinkuhren locality, to which multiple specimens were attributed. We conclude with conspecti on key biological problems and increasing potential of µ-CT for phylogenetic paleontology.
... Formicinae is a large and successful group distributed globally across a wide range of terrestrial habitats comprising about 3030 described species. This subfamily comprises well known taxa like wood ants (formica), carpenter ants (Camponotus), weaver ants (Oecophylla) honey pot ants ( Myrmecocystus) and about fifty other genera (Ward et al. 2016). In the study area, Oecophylla was the dominant genera followed by Paratrechina, Camponotus and Anoplolepis (Fig. 3). ...
Importance of ants in our ecosystem is well recognized. Abundance and stability make the ant population one of the most successful insect groups in the ecosystem. Present investigation was carried out to assess the ant diversity in Kuthuparamba region, Kannur District in Kerala state. Results showed 13 genera of ants representing three subfamilies. The subfamily Myrmicinae was found to be dominant with six genera followed by Formicinae with four genera and Ponerinae with three genera.
... Some of the collected specimens of each species were kept in 96% ethanol for future molecular studies, and some were kept in 75% ethanol for identification and taxonomic studies. Species identified to species level using identification keys such as Collingwood and Agosti (1996), Ward et al. (2016), Khalili-Moghadam et al. (2021), Salata et al. (2021). Additionally, the identification of some samples has been carried out with the help of Dr. Jonathan Romiguier and Yannick Juve from Université de Montpellier. ...
... Formicinae is one of the most diverse ant subfamilies, comprising about 3,030 species widely distributed around the world (Ward et al., 2016). ...
In recent years, the insect microbiome has become the focus of many actinomycete researchers in their search for novel bioactive compounds with members of the order Hymenoptera at the forefront of the revolution. Hymenoptera encompasses all bees, wasps, ants, and sawflies and is the third largest insect order by species richness. Additionally, Hymenoptera is the most diverse insect order in terms of ecological roles, behaviors, and social systems, thus making it an ideal starting point in the search for symbiotic actinomycetes. The aim of this review is to summarize current knowledge on hymenopteran associations with actinomycetes including information on interactions between actinomycetes and hymenopterans, isolation, and screening methodologies, as well as novel actinomycete species and natural products discovered between early 2013 and 2023. A total of 19 new species were discovered within this time period, with the genus Streptomyces being represented by 11 species while the remaining 8 belonged to rare actinomycetes genera. In addition, 35 novel compounds were reported from hymenopteran-associated actinomycetes within the same time period with the majority originating from Streptomyces strains. The reported novel compounds exhibit a range of biological activities including antibacterial, antifungal, anticancer, anti-enzymatic, and antiproliferative activity, as well as cytotoxicity.
... The subfamily Dorylinae was represented by one species only. Formicinae is known as a highly adaptive taxon to any terrestrial environment which explain its current vast global distribution (Ward et al., 2016). On the other hand, Myrmicinae was more common in undisturbed habitat like forest. ...
... The morphology of the species complexes where the species are grouped is doubtful. Ward et al. (2016) revised the classification of the genus and raised the subgenus Colobopsis Mayr, 1861 andDinomyrmex Ashmead, 1905 to the genus level and relegated the genera Forelophilus Kutter, 1931 andPhasomyrmex Stitz, 1910 to subgenera under Camponotus. ...
Two new species, Camponotus sholensis sp. nov. and Camponotus meghalayaensis sp. nov. are described from India and redescriptions of four species (C. habereri Forel, 1911, C. keihitoi Forel, 1913, C. quadrinotatus Forel, 1886 and C. simoni Emery, 1893) new to India are provided. We also recorded and described an unidentified form ‘Camponotus sp. 101’ that does not correspond to any species already known in India. An identification key supplemented with digital images of the known species of the genus is also provided.
... However, a relatively small proportion of ant species have estimates of genetic relatedness in the literature; in particular, the ant subfamily Formicinae is diverse, with 51 extant genera and 3030 described species (Ward et al. 2016), but only a small fraction of genera and species within this complex have estimates of genetic relatedness in the literature (Appendix 1). As such, researchers that want to study population genetics, adaptation, or speciation in understudied eusocial species may desire estimates of kinship and/or relatedness of their samples. ...
... For each of 51 extant genera in this ant family (Ward et al. 2016), we searched Google Scholar with the search terms '[genus]' and 'relatedness' including only results that contained both terms. We manually scanned search results for those potentially relevant to within-colony relatedness for each genus. ...
Understanding a haplodiploid species’ social structure and quantifying relatedness among individuals are both important when designing sampling schemes or identifying potential biases in population genetics studies. However, it is not always possible to accurately identify social structure of study species in the field, or to collect large numbers of individuals from a single colony to estimate relatedness with methods that rely on accurate estimation of allele frequencies. Here, we assessed the utility of allele-frequency-free inference of relationships in haplodiploid ant colonies, while using limited sample sizes. Using genome-wide single-nucleotide polymorphism data, we measured intracolony relatedness and kinship estimates consistent with full-sister relationships among workers in three Nearctic species: Camponotus herculeanus, C. laevissimus, and C. modoc. Notably, the allele-frequency-free inference of relationships clearly demonstrated these full-sister relationships without ambiguity; this result suggests the utility of these methods for identifying closely related individuals in population genetics studies of haplodiploid organisms. We additionally performed a literature review of relatedness estimates in the subfamily Formicinae both as a compiled resource and to place our results in context within this larger clade of ants. Our results suggestive of Camponotus colonies founded by a lone singly mated queen are consistent with previously published relatedness estimates in the genus Camponotus that have generally shown high intracolony relatedness.
... Blochmannia species are obligate intracellular endosymbionts of ants and were first discovered in the genus Camponotus in 1887 by Friedrich Blochmann (18). Preceding phylogenetic studies imply that these endosymbionts were first transferred horizontally from a group of secondary symbionts of mealybugs to the first common ancestor of the Camponotini about 51 million years ago (19)(20)(21). ...
All ants of the species rich genus Camponotus (carpenter ants) possess the obligate intracellular bacterial mutualist Blochmannia. We tested the relevance of the endosymbiont Blochmannia for offspring rearing using cross-fostering experiments between Camponotus sp. colonies and subcolonies (worker groups), which were either treated with antibiotics to remove Blochmannia or untreated. Our antibiotic treatment reduced the level of Blochmannia endosymbionts in eggs, larvae and workers significantly. Corroborating previous results, we found that eggs from treated colonies had a significantly reduced probability to develop into larvae and almost zero probability to become adults. Surprisingly, subcolonies treated with antibiotics had a significantly higher success in raising their own and foreign eggs from treated and untreated colonies than untreated subcolonies. This might indicate that the Blochmannia symbiosis entails significant costs for the host in terms of brood rearing, i.e., symbiont-free workers are more successful in brood rearing than untreated workers. If confirmed, this would be a rare case where the costs of a symbiosis can be empirically measured and quantified. Alternatively, the antibiotic treatment increased as a side effect the brood rearing effort of workers leading to the differences in brood rearing success of treated workers. But even if that would be the case, it still indicates that workers that have either lost or have a significantly reduced number of endosymbionts can still raise brood from antibiotic treated and untreated colonies better than untreated workers. Thus Blochmannia, although crucial for brood development, may reduce the amount of brood a colony can raise.
... Blue squares represent dulosis (n = 10), yellow stars represent inquilinism (n = 57), and grey circles represent temporary social parasitism (n = 24). Economo et al., 2019;Fischer et al., 2020;Maruyama et al., 2008;Mezger & Moreau, 2016;Moreau, 2008;Prebus, 2017;Ward et al., 2010Ward et al., , 2015Ward et al., , 2016, but for social parasites without available historical biogeography data, we assumed the origin event happened in the biogeographic region that the species currently occurs in (see Appendix S3 for comments on decisions and references). ...
Aim
One of the most consistent global biogeographic patterns is the latitudinal diversity gradient where species richness peaks within the equatorial tropics and decreases towards the poles. Here, we explore the global biogeography of socially parasitic ant species, which comprises the most diverse group of social parasites in the Hymenoptera. We test the biogeographic hypothesis that ant social parasites are distributed along an inverse latitudinal diversity gradient (iLDG) by peaking in diversity outside of the equatorial tropics, which would contrast with the biogeographic pattern observed in free‐living, non‐parasitic ant species.
Location
Global.
Taxon
Ants (Hymenoptera: Formicidae).
Methods
We assembled a comprehensive biogeographic dataset consisting of 6001 geographic distribution records for all 371 taxonomically described socially parasitic ant species. We used phylogenetic and taxonomic studies to estimate the number of independent evolutionary origins of ant social parasitism to directly compare species richness with the number of species representing independent evolutionary origins of social parasitism across a latitudinal gradient. In addition, we compared ant social parasite diversity across biogeographic regions using rarefaction to account for different sampling efforts. Finally, we tested for a correlation between latitude and the proportion of ant social parasite species within regional ant faunae.
Results
The geographic distribution records and the inferred 91 independent evolutionary origins of socially parasitic life histories in ants show that both species richness and the number of species representing independent evolutionary origins of social parasitism peak in the northern hemisphere outside of the equatorial tropics. Based on rarefaction curves, northern latitude regions harbour the most ant social parasite species, but the diversity of independent evolutionary origins is not significantly different between northern and southern hemispheres. The proportion of ant social parasite species within regional faunae is tightly correlated with latitude only in the northern hemisphere.
Main conclusions
The iLDG of ant social parasites contrasts with the biogeographic pattern observed in free‐living, non‐parasitic ant species and appears to be driven by large species radiations as well as by the presence of specialized life histories exclusive to the northern hemisphere.
... The remaining species belong to the Formicinae, where C. j. and F. j. are more closely related to each other than to the two Lasius spp. (Fig. 7A; Ward, 2014;Ward et al., 2016). Spontaneous beacon aiming has also been reported for a number of other ant species belonging to these three subfamilies (Fig. 7A). ...
... Fig. 7. Beacon aiming as an ancestral behavioral response. (A) Schematic representation of the phylogenetic relationships of the ant species investigated and ant species for which spontaneous beacon aiming has been described in the literature (based on Ward, 2014 andWard et al., 2016). Black lines: no significant orientation to beacon (BP-), *: low path straightness (PS-). ...
Many insects, including ants, are known to respond visually to conspicuous objects. In this study, we compared orientation in an arena containing only a black target beacon as local information in six species of ants of widely varying degree of phylogenic relatedness, foraging strategy, and eye morphology (Aphaenogaster, Brachyponera, Camponotus, Formica, and two Lasius spp.), often found associated in similar urban anthropogenic habitats. Four species of ants displayed orientation toward the beacon, with two orienting toward it directly, while the other two approached it via convoluted paths. The two remaining species did not show any orientation with respect to the beacon. The results did not correlate with morphological parameters of the visual systems and could not be fully interpreted in terms of the species' ecology, although convoluted paths are linked to higher significance of chemical signals. Beacon aiming was shown to be an innate behavior in visually naive Formica workers, which, however, were less strongly attracted to the beacon than older foragers. Thus, despite sharing the same habitats and supposedly having similar neural circuits, even a very simple stimulus-related behavior in the absence of other information can differ widely in ants but is likely an ancestral trait retained especially in species with smaller eyes. The comparative analysis of nervous systems opens the possibility of determining general features of circuits responsible for innate and possibly learned attraction toward particular stimuli.
