Aneuretinae

Aneuretinae Temporal range: 60.2–0 Ma K Pa Eo Ol Mi Pl Tiffanian to late Paleocene – Recent
Aneuretus simoni
Scientific classification
Kingdom:	Animalia
Phylum:	Arthropoda
Class:	Insecta
Order:	Hymenoptera
Family:	Formicidae
Subfamily:	Aneuretinae Emery, 1913
Diversity
1 genera 1 species 9 fossil genera 13 fossil species

More diverse in its fossil forms, there is but one extant species of this subfamily. Aneuretus simoni is only known from Sri Lanka.

Identification

Sting (that may not be visible), single petiole with a long, narrow anterior peduncle, propodeum armed with a pair of spines. Major tibial spur of hind leg simple or with a few minute barbules. Palp forumula 3,4. Only known from Sri Lanka. In the field workers resemble small yellow Pheidole minors in their morphology. Behaviorally they have a tendency to keep their long petiole folded up against their propodeum with their gaster slightly elevated.

Males Boudinot (2015) - The male of Aneuretus simoni is uniquely identified by the exceedingly long and thin petiolar peduncle and the unpetiolated third abdominal segment. The species is further identified by the following combination of characters: oblique mesopleural sulcus present; seven closed cells present on forewing; jugal lobe absent; abdominal segment IV without cinctus between pre- and postsclerites; abdominal sternum IX unpronged and edentate; telomere extending anteroventrad basimere. Additional characters for distinguishing A. simoni from the Dolichoderinae and Formicinae are indicated in the subfamily key couplets 19 and 20.

Apomorphies of Aneuretinae

Based on Bolton (1990):

(i) Helcium fused in posterior foramen of abdominal segment 2. (Plesiomorphically very mobile in Dolichoderinae.)

(ii) Postsclerites of abdominal segment 3 reduced. (Plesiomorphically large in Dolichoderinae.)

(iii) Petiole with a long anterior peduncle. (Plesiomorphically sessile or subsessile in Dolichoderinae.)

See images of genera within this subfamily

Keys including this Subfamily

• Key to Subfamilies, Males

Statistics

Extant Taxa

Tribes	Valid Genera	% World Genera	Invalid Genera	Valid Species/Subsp.	% World Species	Invalid Species/Subsp.
2	1	0.2%	0	1	0.01%	1

Fossil Taxa

Fossil Genera	% World Fossil Genera	Valid Fossil Species/Subsp.	% World Fossil Species/Subsp.
9	5.2%	13	1.44%

Fossils known from: Baltic amber, Baltic Sea region, Europe (Bartonian, Middle to Late Eocene), Bembridge Marls, Isle of Wight, UK (Priabonian, Late Eocene), Bolshaya Svetlovodnaya, Sikhote-Alin, Russia (Priabonian, Late Eocene), Burmese amber, Kachin State, Myanmar (Early Cenomanian, Late Cretaceous), Florissant, Colorado, United States (Late Eocene), Foremost Formation amber, Alberta, Canada (Campanian, Late Cretaceous), Green River Formation, Colorado, United States (Lutetian, Middle Eocene), Paskapoo Formation, Alberta, Canada (Tiffanian to late Paleocene), Sakhalin amber, Ukraine (Thanetian, Paleocene).

List of Tribes and Genera

Tribes

Aneuretini • Pityomyrmecini

Extant Genera

Aneuretus

Fossil Genera

Aneuretellus • Britaneuretus • Burmomyrma • Cananeuretus • Mianeuretus • Napakimyrma • Paraneuretus • Pityomyrmex • Protaneuretus

Notes

Boudinot (2015) - The sole extant member of the Aneuretinae, A. simoni, is restricted to Sri Lanka, and is the survivor of a lineage which has a somewhat diverse fossil record (LaPolla et al. 2013). The subfamily is of considerable interest as it is sister to the Dolichoderinae (Brady et al. 2006). Eight fossil genera are ascribed to the Aneuretinae based on the work of several authors (e.g., Dlussky & Rasnitsyn 2009). Some taxa, based on workers, are definitely members of the Aneuretinae, i.e., †Paraneuretus and †Protaneuretus from Baltic amber (37–42 My; Wheeler 1915; LaPolla et al. 2013), while others are less certain, i.e., †Pityomyrmex (also from Baltic amber, Wheeler 1915; placed in Aneuretinae by Dlussky & Rasnisyn 2009) and †Aneuretellus (Sakhalin amber, 56–59 My; Dlussky 1988; LaPolla et al. 2013). The impression-fossil taxa †Britaneuretus (see Antropov et al. 2014) and †Mianeuretus (see Carpenter 1930) may not be members of the Aneuretinae. Because of the occurrence of definitive aneuretines in Baltic amber, it will be critical to carefully study the reproductives occurring in these fossils to determine whether any may be placed in the Aneuretinae.

Two fossil “aneuretine” taxa are worth discussing specifically. The affinities of †Burmomyrma (~98 My, Burmese amber; Dlussky 1996; LaPolla et al. 2013) and †Cananeuretus (78–19 My, Canadian amber; Engel & Grimaldi 2005; LaPolla et al. 2013) with Aneuretus simoni are uncertain. The description and illustration of †Burmomyrma in Dlussky (1996) provide no characters which support a relationship of the fossil taxon with Aneuretus; the diagnosis includes one extreme autapomorphy and several characters which are pleisiomorphic for the family or are broadly shared among several subfamilies. The character combination indicated by Dlussky (1996) to assign †Burmomyrma to the Aneuretinae is weak, especially given that Aneuretus has complete (“ancestral”) wing venation while †Burmomyrma lacks almost all vein abscissae. Placement of †Burmomyrma within the Leptanillinae, and indeed other aculeate hymenopteran families, cannot be ruled out. No taxonomic action is taken here, however.

†Cananeuretus, on the other hand, cannot be so easily considered distantly related to the Aneuretinae. The Grassy Lake deposit of Canadian amber includes representatives of the Sphecomyrminae, Ectatomminae, and critically, the Dolichoderinae (LaPolla et al. 2013). While the placement of the fossil dolichoderine †Chronomyrmex (see McKellar et al. 2013) in the Leptomyrmecini (sensu Ward et al. 2010) is debatable, co-occurrence of these subfamilies in this deposit suggests the placement of †Cananeuretus is plausible. As the diagnosis of the Aneuretinae provided here and previously (Wilson et al. 1956; Bolton 2003) is based largely on pleisiomorphic characters, other characters should be considered. For example, future studies of Canadian amber should be sensitive to specific traits occurring in Aneuretus and extant Dolichoderinae. Aneuretus shares, among other characters, a deep median notch on the anterior clypeal margin and fine serrations intercalated among larger denticles on the masticatory mandibular margin, both of which occur in the Tapinomini, the tribe sister to the remaining Dolichoderinae (Ward et al. 2010). Reconsideration of the fossil record of Aneuretinae will be valuable for improving our concepts of both the Aneuretinae and Dolichoderinae.

Biology

Flight Period

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Flight Months for Genera within this Subfamily:

Taxon	Month
Aneuretellus
Aneuretus
†Aneuretellus
†Britaneuretus
†Burmomyrma
†Cananeuretus
†Mianeuretus
†Napakimyrma
†Paraneuretus
†Pityomyrmex
†Protaneuretus
Britaneuretus
Burmomyrma
Cananeuretus
Mianeuretus
Napakimyrma
Paraneuretus
Pityomyrmex
Protaneuretus

Morphology

Karyotype Data

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Known Haploid Counts: No results.

Haploid Count Details: No results.

Known Diploid Counts: No results.

Diploid Count Details: No results.

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

• ANEURETINAE [subfamily of Formicidae]
  • Aneuretini Emery, 1913a: 6. Type-genus: Aneuretus Emery, 1893a: cclxxv.

Taxonomic History

• Aneuretini as tribe of Dolichoderinae: Emery, 1913a: 6.
• Aneuretinae as subfamily of Formicidae: Clark, 1951: 16 (footnote); Wilson, et al. 1956: 93; Wheeler, G.C. & Wheeler, J. 1972a: 40; Snelling, R.R. 1981: 400; Dlussky & Fedoseeva, 1988: 78; Bolton, 1990c: 1361; Hölldobler & Wilson, 1990: 16; Shattuck, 1992b: 201; Baroni Urbani, et al. 1992: 315; Shattuck, 1994: 1; Bolton, 1994: 15; Bolton, 1995b: 9.
• Aneuretinae as junior synonym of Dolichoderinae: Baroni Urbani, 1989: 147.
• Aneuretinae as formicomorph subfamily of Formicidae: Bolton, 2003: 18, 79.
• Aneuretinae as formicoid subfamily of Formicidae: Brady, et al. 2006: 18173; Moreau, et al. 2006: 102.
• Aneuretinae as formicoid dolichoderomorph subfamily of Formicidae: Ward, 2007a: 556.

Taxonomic References

Forel, 1895e: 461 (diagnosis); Bingham, 1903: 290 (diagnosis); Emery, 1913a: 6 (diagnosis); Wilson, et al. 1956: 93 (diagnosis, review of subfamily and genus); Eisner, 1957: 453 (proventriculus morphology); Wheeler, G.C. & Wheeler, J. 1972a: 40 (diagnosis); Wheeler, G.C. & Wheeler, J. 1976b: 60 (larvae, review and synthesis); Wheeler, G.C. & Wheeler, J. 1985: 258 (synoptic classification); Bolton, 1990c: 1361 (morphology, status); Shattuck, 1992b: 201 (higher classification, phylogeny); Baroni Urbani, et al. 1992: 315 (phylogeny); Bolton, 1994: 15 (diagnosis, synoptic classification); Bolton, 1995a: 1047 (census); Dlussky & Rasnitsyn, 2002: 414 (diagnosis, wingless impression fossils); Bolton, 2003: 18, 79 (diagnosis, synopsis); Brady, et al. 2006: 18173 (phylogeny); Moreau, et al. 2006: 102 (phylogeny); Keller, 2011: 1 (morphology, phylogeny); Dlussky & Perfilieva, 2014: 433 (British Eocene species key); Boudinot, 2015: 49 (diagnosis).

References

• Baroni Urbani, C. 1989. Phylogeny and behavioural evolution in ants, with a discussion of the role of behaviour in evolutionary processes. Ethol. Ecol. Evol. 1:137-168.
• Bolton, B. 1990e. Army ants reassessed: the phylogeny and classification of the doryline section (Hymenoptera, Formicidae). J. Nat. Hist. 24:1339-1364.
• Bolton, B. 2003. Synopsis and classification of Formicidae. Mem. Amer. Entomol. Inst. 71:1-370.
• Borowiec, M.L., Moreau, C.S., Rabeling, C. 2020. Ants: Phylogeny and Classification. In: C. Starr (ed.), Encyclopedia of Social Insects (doi:10.1007/978-3-319-90306-4_155-1).
• Boudinot, B.E. 2015. Contributions to the knowledge of Formicidae (Hymenoptera, Aculeata): a new diagnosis of the family, the first global male-based key to subfamilies, and a treatment of early branching lineages. European Journal of Taxonomy. 120:1-62. doi:10.5852/ejt.2015.120
• Clark, J. 1951. The Formicidae of Australia. 1. Subfamily Myrmeciinae: 230 pp. CSIRO, Melbourne. [(31.xii).1951.]
• Emery, C. 1913a [1912]. Hymenoptera. Fam. Formicidae. Subfam. Dolichoderinae. Genera Insectorum. 137:1-50.