Eusphinctus

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Eusphinctus is a species-poor South East Asian genus with apparently small colonies. There are only two species of Eusphinctus, both quite similar, and Brown (1975: 75) allowed the possibility that specimens described as Eusphinctus taylori may be just small workers of Eusphinctus furcatus, but he decided not to synonymize them until more specimens are available. (Borowiec 2016)

Identification

Eusphinctus workers from Kerala, India, Photo by Kalesh Sadasivan.

Borowiec (2016) - Worker Eusphinctus workers belong to dorylines with conspicuous gastral constrictions visible between abdominal segments IV, V, and VI. This morphology is also seen in Aenictogiton, certain species of Leptanilloides, Sphinctomyrmex, and Zasphinctus. Eusphinctus is unique in the combination of propodeal spiracle situated low on the sclerite and propodeal lobes present, a large pygidium armed with modified setae, pronotomesopleural suture present, and cinctus of abdominal segment IV simple and not cross-ribbed. This genus is thus far known only from India, Bangladesh, Myanmar, and Thailand and the only lineage with gastral constriction that is currently known to overlap with it is Zasphinctus. In Zasphinctus the pronotomesopleural suture is fused, in Aenictogiton the propodeal spiracle is positioned high and there are no propodeal lobes, Leptanilloides has a reduced and unarmed pygidium, and the neotropical Sphinctomyrmex has 12-segmented antennae and the cinctus on abdominal segment IV smooth. The relative proportions of abdominal segments are also different, with segments IV, V, and VI being about equal in size in Sphinctomyrmex and Zasphinctus, while in Eusphinctus segment IV is the largest of the three.

Male The male of Eusphinctus can be recognized by a combination of 12-segmented antennae, pronounced propodeal lobes, narrow axial helcium, conspicuous constrictions present between abdominal segments IV, V, and VI, costal (C) cell present in the fore wing, submarginal cell closed by Rs·f2–3, R·f3 present past pterostigma, and marginal cell open. Abdominal sternite IX (subgenital plate) in Eusphinctus gradually tapers caudad and has simple, straight spines directed posteriorly. Sphinctomyrmex and Zasphinctus also have constrictions between abdominal segments IV, V, and VI but the former always has 13-segmented antennae and the latter lacks veins C and R·f3 in the fore wing.

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Keys including this Genus

 

Distribution

Indomalayan, known from India, Bangladesh, Myanmar, and Thailand.

Distribution and Richness based on AntMaps

Species by Region

Number of species within biogeographic regions, along with the total number of species for each region.

Afrotropical Region Australasian Region Indo-Australian Region Malagasy Region Nearctic Region Neotropical Region Oriental Region Palaearctic Region
Species 0 0 0 0 0 0 2 0
Total Species 2840 1735 3042 932 835 4378 1740 2862

Biology

A. B. Soans and W. L. Brown collected two colonies of Eusphinctus furcatus in Kottiyoor, Kerala, India. One was located in leaf litter near a rotting log and the other one was found under a stone in a shaded creek bottom. There were about 50 workers in each of the observed nests, and one colony contained two ergatoid gynes (Brown 1975).

Castes

Morphology

Phylogeny

Dorylinae

Lioponera (76 species, 0 fossil species)

Lividopone (1 species, 0 fossil species)

Parasyscia (57 species, 0 fossil species)

Zasphinctus (24 species, 0 fossil species)

Vicinopone (1 species, 0 fossil species)

Simopone (40 species, 0 fossil species)

Tanipone (10 species, 0 fossil species)

Eusphinctus (2 species, 0 fossil species)

Ooceraea (17 species, 0 fossil species)

Syscia (39 species, 0 fossil species)

Eburopone (2 species, 0 fossil species)

Aenictus (226 species, 0 fossil species)

Aenictogiton (7 species, 0 fossil species)

Dorylus (127 species, 0 fossil species)

Cerapachys (5 species, 0 fossil species)

Chrysapace (4 species, 0 fossil species)

Yunodorylus (4 species, 0 fossil species)

Neocerapachys (2 species, 0 fossil species)

Acanthostichus (23 species, 1 fossil species)

Cylindromyrmex (10 species, 3 fossil species)

Sphinctomyrmex (3 species, 0 fossil species)

Leptanilloides (19 species, 0 fossil species)

Neivamyrmex (129 species, 1 fossil species)

Cheliomyrmex (4 species, 0 fossil species)

Labidus (9 species, 0 fossil species)

Eciton (29 species, 0 fossil species)

Nomamyrmex (2 species, 0 fossil species)

See Phylogeny of Dorylinae for details.

Nomenclature

The following information is derived from Barry Bolton's Online Catalogue of the Ants of the World.

  • EUSPHINCTUS [Dorlyinae]
    • Eusphinctus Emery, 1893a: cclxxv. Type-species: Eusphinctus furcatus, by monotypy.
    • Eusphinctus subgenus of Sphinctomyrmex: Emery, 1895k: 456.
    • Eusphinctus revived status as genus: Wheeler, W.M. 1918a: 219.
    • Eusphinctus junior synonym of Sphinctomyrmex: Brown, 1975: 31.
    • Eusphinctus as genus: Borowiec, 2016: 138.

Taxonomic Notes

Borowiec (2016) - Based on morphological and molecular evidence (Borowiec in prep.), I revive Eusphinctus from synonymy with Sphinctomyrmex. The taxonomic history of taxa classified under Sphinctomyrmex is somewhat complicated. Detailed discussions can be found in Wheeler (1918) and Brown (1975), and I only briefly recount the history of taxonomic changes to provide a background for an arrangement proposed here. The genus Sphinctomyrmex was established by Mayr (1866b) based on a single dealate gyne specimen from Brazil. Mayr emphasized the prominent constrictions between abdominal segments present in the specimen in the description, which gave inspiration for the name. Later, Emery (1893a) described a new genus from Myanmar, Eusphinctus, for an ant with similar constrictions. Other species from the Old World followed, described under either name. Wheeler in 1918 decided (after André 1905) to reserve Sphinctomyrmex for all New World forms and further split Eusphinctus into three subgenera, Eusphinctus s. str., Nothosphinctus, and Zasphinctus, according to various combinations of the gyne morphology, number of antennal segments (11 or 12), and presence or absence of eyes in the worker. Brown (1975) discussed the taxonomic history of the genus and all of Wheeler’s characters in detail. He pointed out that the characters used to differentiate these vary and the combinations enumerated by Wheeler do not hold as generic diagnoses with newly discovered species. Brown thus concluded that it was most sensible to synonymize all the genus-level names under Sphinctomyrmex until more evidence, particularly from male morphology, was gathered. However, he allowed for a possibility that two species, Eusphinctus furcatus and Eusphinctus taylori indeed deserved a separate generic status (Brown 1975).

Here I propose a new classification where all the New World species are retained in Sphinctomyrmex, while most of the described Old World forms are relegated to Zasphinctus. The two remaining above mentioned Old World species are separated from Zasphinctus as Eusphinctus. Molecular data shows that all three genera arose independently on the dorylomorph tree. Despite sharing characteristic gastral constrictions, these lineages are also discrete in worker and male morphology (see diagnosis above). Both morphology and molecules support the notion that abdominal constrictions have been independently derived several times in the Dorylinae: in Eusphinctus, Sphinctomyrmex, and Zasphinctus and also in Aenictogiton and some Leptanilloides.

Eusphinctus belongs to a clade that also includes Ooceraea and Syscia (Borowiec, in prep.). The members of this clade share the universal reduction in the number of antennal segments, from 12 to 11 or fewer in the worker caste and from 13 to 12 or fewer in males.

Description

Worker

Borowiec (2016) - Head: Antennae with 11 segments. Apical antennal segment moderately enlarged, broader than and about equal in length to two preceding segments combined. Clypeus without cuticular apron. Lateroclypeal teeth present. Parafrontal ridges reduced. Torulo-posttorular complex vertical. Antennal scrobes absent. Labrum unknown. Proximal face of stipes unknown. Maxillary palps unknown. Labial palps unknown. Mandibles triangular, with teeth. Eyes present, composed of fewer than five ommatidia. Ocelli absent. Head capsule with differentiated vertical posterior surface above occipital foramen. Ventrolateral margins of head without lamella or ridge extending towards mandibles and beyond carina surrounding occipital foramen. Posterior head corners dorsolaterally immarginate. Carina surrounding occipital foramen ventrally present. Mesosoma: Pronotal flange separated from collar by distinct ridge. Promesonotal connection with suture completely fused. Pronotomesopleural suture visible, unfused up to notal surface. Mesometapleural groove weakly impressed. Transverse groove dividing mesopleuron present. Pleural endophragmal pit concavity present. Mesosoma dorsolaterally immarginate. Metanotal depression or groove on mesosoma absent. Propodeal spiracle situated low on sclerite. Propodeal declivity with distinct dorsal edge or margin and rectangular in posterior view. Metapleural gland without bulla visible through cuticle. Propodeal lobes present, well developed. Metasoma: Petiole anterodorsally immarginate, dorsolaterally immarginate, and laterally above spiracle marginate. Helcium in relation to tergosternal suture placed at posttergite and axial. Prora forming a simple U-shaped margin. Spiracle openings of abdominal segments IV–VI circular. Abdominal segment III anterodorsally immarginate and dorsolaterally immarginate. Abdominal segment III more than half size of succeeding segment IV, which is weakly constricted at presegmental portion (uninodal waist). Girdling constriction of segment IV present, i.e. pre- and postsclerites distinct. Cinctus of abdominal segment IV gutter-like, not sculptured. Abdominal segment IV not conspicuously largest segment. Abdominal tergite IV not folding over sternite, and anterior portions of sternite and tergite equally well visible in lateral view. Girdling constriction between pre- and posttergites of abdominal segments V and VI present. Girdling constriction between pre- and poststernites of abdominal segments V and VI present. Pygidium large, with impressed medial field, armed with modified setae, and deeply notched at apex. Hypopygium unarmed. Legs: Mid tibia with single pectinate spur. Hind tibia with single pectinate spur. Hind basitarsus not widening distally, circular in cross-section. Posterior flange of hind coxa not produced as raised lamella. Metatibial gland absent. Metabasitarsal gland absent. Hind pretarsal claws simple. Polymorphism: Monomorphic.

Queen

Borowiec (2016) - Ergatoid, ‘scarcely different in size from the workers’ (Brown 1975) except slightly larger eyes and wider abdominal segment II (petiole). Presence of ocelli unknown.

Male

Borowiec (2016) - Head: Antennae with 12 segments. Clypeus with cuticular apron. Parafrontal ridges absent. Torulo-posttorular complex vertical. Maxillary palps unknown. Labial palps unknown. Mandibles triangular, edentate. Ventrolateral margins of head without lamella or ridge extending towards mandibles and beyond carina surrounding occipital foramen. Carina surrounding occipital foramen ventrally present. Mesosoma: Pronotal flange not separated from collar by distinct ridge. Notauli present. Transverse groove dividing mesopleuron present. Propodeal declivity with distinct dorsal edge or margin. Metapleural gland opening absent. Propodeal lobes present. Metasoma: Petiole anterodorsally immarginate, dorsolaterally immarginate, and laterally above spiracle marginate. Helcium in relation to tergosternal suture placed at posttergite and supraaxial. Prora forming a V-shaped protrusion. Spiracle openings of abdominal segments IV–VI circular. Abdominal segment III more than half size of succeeding segment IV; latter weakly constricted at presegmental portion (uninodal waist). Girdling constriction of segment IV present, i.e. pre- and postsclerites distinct. Cinctus of abdominal segment IV gutter-like, not sculptured. Girdling constriction between pre- and postsclerites of abdominal segments V and VI present. Abdominal segment IV not conspicuously largest segment. Abdominal sternite VII simple. Abdominal sternite IX distally armed with two spines, with lateral apodemes about as long as medial apodeme, directed anteriorly (towards head). Genitalia: Cupula long relative to rest of genital capsule and of approximately equal length on both dorsal and ventral surfaces. Basimere broadly fused to telomere, with sulcus discernable at junction, and ventrally with left and right arms abutting. Telomere gradually tapering toward apex. Volsella narrow, hook-shaped. Penisvalva laterally compressed, rounded at apex. Legs: Mid tibia with single pectinate spur. Hind tibia with single pectinate spur. Posterior flange of hind coxa not produced as raised lamella. Metatibial gland absent. Metabasitarsal glands absent. Hind pretarsal claws simple. Wings: Tegula present, broad, demiovate in shape. Vein C in fore wing present. Pterostigma broad. Abscissa R·f3 absent. Abscissae Rs·f2–3 present, disconnected from Rs+M. Cross-vein 2r-rs present, differentiated from Rs·f4 by presence of Rs·f2–3. Abscissae Rs·f4–5 present, fused in absence of 2rs-m. Abscissa M·f2 in fore wing contiguous with Rs+M. Abscissa M·f4 in fore wing present, reaching wing margin. Cross-vein 1m-cu in fore wing present. Cross-vein cu-a in fore wing present, arising from Cu and distal to, at or near M·f1. Vein Cu in fore wing present, with only Cu1 branch prominent. Vein A in fore wing with abscissae A·f1 and A·f2 present. Vein C in hind wing absent. Vein R in hind wing absent. Vein Sc+R in hind wing present. Abscissa Rs·f1 in hind wing present, shorter than 1rs-m. Abscissa Rs·f2 in hind wing absent. Cross-vein 1rs-m in hind wing fused with M·f1. Vein M+Cu in hind wing present. Abscissa M·f1 in hind wing present. Abscissa M·f2 in hind wing present. Cross-vein cu-a in hind wing present. Vein Cu in hind wing present. Vein A in hind wing with abscissae A·f1 and A·f2 present.

Larva

Borowiec (2016) - Larva not known. Presence of cocoons unknown.

References

  • Borowiec, M.L. 2016. Generic revision of the ant subfamily Dorylinae (Hymenoptera, Formicidae). ZooKeys 608: 1–280 (doi: 10.3897/zookeys.608.9427).
  • Bolton, B. 2003. Synopsis and Classification of Formicidae. Mem. Am. Entomol. Inst. 71: 370pp (page 142, Eusphinctus as junior synonym of Sphinctomyrmex )
  • Borowiec, M.L. 2019. Convergent evolution of the army ant syndrome and congruence in big-data phylogenetics. Systematic Biology 68, 642–656 (doi:10.1093/sysbio/syy088).
  • Brown, W. L., Jr. 1973b. A comparison of the Hylean and Congo-West African rain forest ant faunas. Pp. 161-185 in: Meggers, B. J., Ayensu, E. S., Duckworth, W. D. (eds.) Tropical forest ecosystems in Africa and South America: a comparative review. Wash (page 180, Eusphinctus as junior synonym of Sphinctomyrmex [provisional])
  • Brown, W. L., Jr. 1975. Contributions toward a reclassification of the Formicidae. V. Ponerinae, tribes Platythyreini, Cerapachyini, Cylindromyrmecini, Acanthostichini, and Aenictogitini. Search Agric. (Ithaca N. Y.) 5(1 1: 1-115 (page 31, Eusphinctus as junior synonym of Sphinctomyrmex )
  • Cantone S. 2018. Winged Ants, The queen. Dichotomous key to genera of winged female ants in the World. The Wings of Ants: morphological and systematic relationships (self-published).
  • Donisthorpe, H. 1943g. A list of the type-species of the genera and subgenera of the Formicidae. [part]. Ann. Mag. Nat. Hist. 11(10): 617-688 (page 645, Eusphinctus in Cerapachyinae, Cerapachyini)
  • Emery, C. 1893a [1892]. [Untitled. Introduced by: "M. C. Emery, de Bologne, envoie les diagnoses de cinq nouveaux genres de Formicides".]. Bull. Bimens. Soc. Entomol. Fr. 1892:cclxxv-cclxxvii. (page cclxxv, Eusphinctus as genus)
  • Emery, C. 1895m. Viaggio di Leonardo Fea in Birmania e regioni vicine. LXIII. Formiche di Birmania del Tenasserim e dei Monti Carin raccolte da L. Fea. Parte II. Ann. Mus. Civ. Stor. Nat. 34[=(2(14): 450-483 (page 456, Eusphinctus as subgenus of Sphinctomyrmex)
  • Emery, C. 1911e. Hymenoptera. Fam. Formicidae. Subfam. Ponerinae. Genera Insectorum 118: 1-125 (page 7, Eusphinctus as subgenus of Sphinctomyrmex)
  • Forel, A. 1900f. Les Formicides de l'Empire des Indes et de Ceylan. Part VII. J. Bombay Nat. Hist. Soc. 13: 303-332 (page 328, Eusphinctus as subgenus of Sphinctomyrmex)
  • Forel, A. 1917. Cadre synoptique actuel de la faune universelle des fourmis. Bull. Soc. Vaudoise Sci. Nat. 51: 229-253 (page 239, Eusphinctus as subgenus of Sphinctomyrmex)
  • Wheeler, W. M. 1910b. Ants: their structure, development and behavior. New York: Columbia University Press, xxv + 663 pp. (page 137, Eusphinctus as subgenus of Sphinctomyrmex)
  • Wheeler, W. M. 1918a. The Australian ants of the ponerine tribe Cerapachyini. Proc. Am. Acad. Arts Sci. 53: 215-265 (page 219, Eusphinctus as genus)
  • Wheeler, W. M. 1922i. Ants of the American Museum Congo expedition. A contribution to the myrmecology of Africa. VII. Keys to the genera and subgenera of ants. Bull. Am. Mus. Nat. Hist. 45: 631-710 (page 639, Eusphinctus in Cerapachyinae, Cerapachyini; Eusphinctus as genus)