Amblyoponinae

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Amblyoponinae
Temporal range: 47–0 Ma Lutitian, Middle Eocene – Recent
Amblyopone australis
Scientific classification
Kingdom: Animalia
Phylum: Arthropoda
Class: Insecta
Order: Hymenoptera
Family: Formicidae
Subfamily: Amblyoponinae
Forel, 1893
Diversity
9 genera
146 species
1 fossil genera
7 fossil species

The subfamily Amblyoponinae contains 11 genera which are found throughout most of the world, although they are more common in tropical and forested regions. It is represented in Australia by five genera which are most abundant and diverse in coastal and heavily forested areas, the subfamily being largely absent from the dry interior. Of these five, two (Amblyopone and Prionopelta) are found essentially world-wide, one (Mystrium) is found in Africa and from India to Australia, one (Myopopone) occurs from India to Australia while the last (Onychomyrmex) is restricted to Far North Queensland. Some species can be locally common and regularly encountered while others are rarely seen.

Nests are in soil under rocks or logs or in rotten wood. Colonies are often composed of numerous small nests scattered over a small area. In some cases large central chambers are absent and the colony consists of many small satellite nests containing a few workers together with a small amount of brood (larvae and pupae).

In most genera workers are cryptic predators in soil and leaf litter and are seldom seen foraging on the surface of the ground. Some species show a strong preference for centipedes while others will feed on a range of soft-bodied arthropods. It is common in some groups for workers to bring their larvae to food sources rather than attempt to move especially large prey back to the nest. When disturbed most will move slowly underground or remain motionless.

One genus, Onychomyrmex, differs significantly from others in this subfamily. These uncommon ants have an army-ant life style. They alternate between temporary bivouacs in protected sites on the ground and semi-permanent nests in soil or in rotten logs. Groups of workers forage by forming distinct raiding columns across the surface of the ground and some species are known to be specialist predators on centipedes.


Photo Gallery

  • Stigmatomma oregonense Quincy, California. Photo by Alex Wild.

Identification

The petiole with distinct front and top faces but without a separate rear face, the rear section of the petiole being attached to the gaster by its entire height. The upper surfaces of the petiole and gaster separated by at most a shallow impression. The upper plate of the second segment of the gaster (second gastral tergite) only weakly arched and located in the middle of the gaster, the rear-most part of the gaster pointing rearward or directly down. The broad attachment of the petiole to the gaster combined with the presence of a relatively straight gaster will separate species of Amblyoponinae from those of other subfamilies.

Boudinot (2015) - The supraaxial helcium serves to distinguish both female and male Amblyoponinae, excluding Apomyrma and Opamyrma, from the majority of the Formicidae. Besides occurrence in the amblyoponines, the supraaxial state of the helcium is only developed in Acanthostichus (Dorylinae) and male Proceratium (Proceratiinae), and weakly in the workers of Martialis (Martialinae) and the male of Tatuidris (Agroecomyrmecinae). Males of three amblyoponine genera (Bannapone, Concoctio, and Opamyrma) remain unknown.

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Keys including this Subfamily

Keys to Genus in this Subfamily

Distribution

Distribution and Species Richness based on AntMaps

Statistics

Extant Taxa

Tribes Valid Genera % World Genera Invalid Genera Valid Species/Subsp. % World Species Invalid Species/Subsp.
1 9 1.7% 13 146 0.96% 49

Fossil Taxa

Fossil Genera % World Fossil Genera Valid Fossil Species/Subsp. % World Fossil Species/Subsp.
1 0.6% 7 0.77%

Fossils known from: Baltic amber, Baltic Sea region, Europe (Bartonian, Middle to Late Eocene), Bolshaya Svetlovodnaya, Sikhote-Alin, Russia (Priabonian, Late Eocene), Chon-Tyz mine, Naryn Province, Kyrgyzstan (Middle Miocene), Messel, Germany (Lutetian, Middle Eocene), Radoboj, Croatia (Burdigalian, Early Miocene), Shanwang, China (Early Miocene).

List of Tribes and Genera

Tribes

Extant Genera

Fossil Genera

Notes

Boudinot (2015) - The Amblyoponinae was only recently split from the Ponerinae sensu lato (Bolton 2003) and has been recovered in all molecular phylogenies as poneroids, with uncertain relationship to the Proceratiinae and the remainder of the group (Brady et al. 2006; Moreau & Bell 2013; Ward 2014). Two main clades are recovered in the Amblyoponinae, termed the XMAS (Xymmer, Myopias, Adetomyrma, Stigmatomma) and OCP (Onychomyrmex, Concoctio, Prionopelta) clades (Yoshimura & Fisher 2012a). Apomyrma, in its original description (Brown et al. 1971), was proposed to be closely related to the Amblyoponinae (then Amblyoponini), a contention supported by Wheeler & Wheeler (1985) and Hölldobler & Wilson (1990). Other authors demurred, placing the genus in its own tribe (Apomyrmini) in the Ponerinae sensu lato (Dlussky & Fedoseeva 1988) and in the Leptanillinae (Bolton 1990b; Kugler 1992), and eventually in its own subfamily, Apomyrminae (Baroni Urbani et al. 1992; Bolton 1994, 2003). The “Apomyrma” sequences used by Saux et al. (2004) were contaminated (P.S. Ward, pers. comm.), but fortuitously their transfer of Apomyrma to the Amblyoponinae was supported by subsequent studies (Brady et al. 2006; Moreau & Bell 2013). This classification is followed here.

Biology

Flight Period

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Morphology

Karyotype Data

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Known Haploid Counts: 9, 16, 19, 22, 24.

Haploid Count Details: 16 (Taxon: Mystrium camillae), 19 (Taxon: Stigmatomma reclinatum), 22 (Taxon: Amblyopone), 24 (Taxon: Amblyopone australis), 9 (Taxon: Fulakora cleae).

Known Diploid Counts: 18, 32, 38, 44, 48.

Diploid Count Details: 18 (Taxon: Fulakora cleae), 32 (Taxon: Mystrium camillae), 38 (Taxon: Stigmatomma reclinatum), 44 (Taxon: Amblyopone), 48 (Taxon: Amblyopone australis).

Phylogeny

Formicoidea
Camelomecia clade
(unplaced to family)

Camelomecia

Camelosphecia

Formicidae              
Stem Formicidae

Unplaced to subfamily (Baikuris, Cretomyrma, Dlusskyidris)

Clade Sphecomyrmines

Haidomyrmecinae

Sphecomyrminae

Zigrasimeciinae

Clade Antennoclypeata

Brownimeciinae

Crown Formicidae              

Leptanillinae

Martialinae

Amblyoponinae - see relationships

Apomyrminae

Proceratiinae

Ponerinae - see relationships

Agroecomyrmecinae

Paraponerinae

Dorylinae - see relationships

Aneuretinae

Dolichoderinae - see relationships

Myrmeciinae - see relationships

Pseudomyrmecinae - see relationships

Formicinae - see relationships

Ectatomminae - see relationships

Myrmicinae - see relationships

See Phylogeny of Formicidae for details.

Nomenclature

  • AMBLYOPONINAE [subfamily of Formicidae]
    • Amblyoponinae Forel, 1893a: 162. Type-genus: Amblyopone Erichson, 1842: 260.

Taxonomic History

  • Amblyoponinae as subfamily of Formicidae: Forel, 1893a: 162 [Amblyoponinae]; Forel, 1895b: 110 [Amblyoponeridae]; Clark, 1951: 15 (in key) [Amblyoponinae]; Bolton, 2003: 41 (revived status).
  • Amblyoponinae as poneromorph subfamily of Formicidae: Bolton, 2003: 41, 153.
  • Amblyoponinae as poneroid subfamily of Formicidae: Ouellette, et al. 2006: 365; Brady, et al. 2006: 18173; Moreau, et al. 2006: 102; Ward, 2007a: 555; Ward & Fisher, 2016: 689.
  • Amblyoponinae as senior synonym of Apomyrmini: Saux, et al. 2004: 466.
  • [Amblyoponeridae: incorrect subsequent spelling by Forel, 1895b: 110.]

Taxonomic References

Emery, 1895j: 766 (diagnosis); Wheeler, W.M. 1910g: 134 (diagnosis); Emery, 1911d: 21 (diagnosis, genera, key); Emery, 1911d: 96 (subtribe Onychomyrmicini diagnosis); Forel, 1917: 235 (synoptic classification); Wheeler, W.M. 1922a: 640, 758, 1006 (genera key, Afrotropical, Malagasy catalogues); Brown, 1960a: 146 (diagnosis, all genera revision, key); Wheeler, G.C. & Wheeler, J. 1976b: 48 (larvae, review and synthesis); Wheeler, G.C. & Wheeler, J. 1985: 256 (synoptic classification); Terayama, 1989b: 345 (Taiwan fauna); Hölldobler & Wilson, 1990: 9 (synoptic classification); Brandão, 1991: 389 (Neotropical fauna, synoptic classification); Bolton, 1994: 164 (synoptic classification); Ward, 1994: 167 (tribe diagnosis and discussion); Bolton, 1995a: 1042 (census); Bolton, 1995b: 9 (catalogue); Hashimoto, 1996: 353 (abdominal structure); Schoeters, et al. 1999: 3 (venom gland structure); Xu, 2000a: 300 (Malesian genera key); Bolton, 2003: 41, 153 (diagnosis, synopsis); Saux, et al. 2004: 466 (phylogeny, diagnosis); Ouellette, et al. 2006: 359 (phylogeny); Brady, et al. 2006: 18173 (phylogeny); Moreau, et al. 2006: 102 (phylogeny); Ward, 2007a: 555 (classification); Fernández & Arias-Penna, 2008: 30 (Neotropical genera key); Terayama, 2009: 93 (Taiwan genera key); Keller, 2011: 1 (morphology, phylogeny); Yoshimura & Fisher, 2012a: 11 (Malagasy genera males key); General & Alpert, 2012: 70 (Philippines genera key); Sarnat & Economo, 2012: 22 (Fiji Is species key); Boudinot, 2015: 17 (diagnosis); Baccaro, et al. 2015: 66, 120 (Brazil genera key, text); Esteves & Fisher, in Delabie, et al. 2015: 13 (amblyoponines of neotropics); Ward & Fisher, 2016: 689 (phylogeny); Fisher & Bolton, 2016: 39 (diagnosis).

Forel 1893 37:162

Taxonomic Notes

Boudinot (2015):

Synapomorphies

The Amblyoponinae was diagnosed by Bolton (2003) based primarily on the female castes, while more recently Yoshimura & Fisher (2012a) diagnosed the males for the Malagasy region. Apomyrma and Opamyrma violate most of these characters; these violations are noted below. Synapomorphies of the Amblyoponinae from the two aforementioned resources are as follows, with respective pleisiomorphies presented in brackets:

1. Dentiform clypeal setae present on anterior clypeal margin (all adult castes) (note 1). [Dentiform clypeal setae absent.]

2. Metapleural gland orifice directed more-or-less posterodorsally (female castes) (note 2). [Metapleural gland orifice directed laterally.]

3. Helcium supraaxial, thus petiole situated high on abdominal segment III, petiole without distinct posterodorsal face and abdominal tergum III without distinct anterodorsal face (all adult castes) (note 3). [Helcium infraaxial.]

4. Petiole very broadly attached to abdominal segment III (all adult castes) (note 4). [Petiole narrowly attached to abdominal segment III.]

5. Helcial sternite very wide (all adult castes) (note 4). [Helcial sternite narrow.]

6. Abdominal segment IV tergosternal fusion present (all adult castes) (note 5). [Abdominal segment IV tergosternal fusion absent.]

7. Basivolsella with ventroapical process, near bases of cuspis and digitus (male) (note 6). [Apical process of basivolsella absent.]

Notes on synapomorphies

1. Generally present in males, although these setae may be difficult to ascertain or absent in very small species. Not present in any other extant ant taxon, although present in most †Sphecomyrminae. Apomyrma and Opamyrma workers and Apomyrma males lack dentiform clypeal setae (the male of Opamyrma is unknown); rather, the workers have dentiform setae on the labrum. The male of Apomyrma has a very reduced labrum which lacks dentiform setae.

2. Males with or, more often, without metapleural gland orifice. The metapleural gland of Apomyrma was not evaluated in this study due to insufficient magnification.

3. This corresponds to the third, fifth, and sixth amblyoponine synapomorphies of Bolton (2003). Apomyrma workers and males have infraaxial helcia, while the worker of Opamyrma has an axial helcium.

4. Petiole very narrowly attached in worker Apomyrma and Opamyrma. The petiolar conformation of the male of Apomyrma, though broad, still differs from that observed in Amblyoponinae (see note 5).

5. Reversed in Adetomyrma female castes and variable in males (Yoshimura & Fisher 2012a).

6. Character from Yoshimura & Fisher (2012a) and confirmed here via dissection of Amblyopone, Myopopone, Onychomyrmex, Paraprionopelta, and New World Stigmatomma.

References