|Formica nasuta, now Proformica nasuta|
|Based on Ward et al. 2016.|
The ant genus Proformica Ruzsky, 1902 is composed of 25 species (Bolton 2014) restricted to dry and open environments such as steppes, mountain meadows and Mediterranean seashores (Agosti 1994). It is endemic to the Palaearctic region, with a disjunct distribution. A first area extends from eastern Europe to eastern Asia and contains most of the species, and a second area, much more limited in species number and distribution, occurs at the southwestern tip of Europe (Portugal, Spain and southern France). (Galkowski et al. 2017)
Distribution and Richness based on AntMaps
Galkowski et al. (2017) - Queen reproductive status, nest census. Excavation of nests of the two taxa revealed the same general structure: the entrance opens directly at the ground surface, sometimes under a small stone; a vertical gallery of 10–20 cm leads to a small chamber where males can be found when present; then, the gallery goes down obliquely and reaches a final chamber, about 50 cm below ground level, where queens are present. Secondary galleries, lateral (perpendicular) to the principal one, may be present and lead to chambers. Repletes, i.e. workers with inflated gaster serving as stores of liquid food, were found in colonies of both taxa. Colonies had one to many queens that appeared to be actively reproducing (mated, with numerous mature oocytes and yellow bodies). Workers, even the largest, always had fewer than 3 ovarioles per ovary and never had a spermatheca. In contrast, apterous and winged queens had a spermatheca and many more ovarioles per ovary (~ 10).
• Antennal segment count 12 • Antennal club absent • Palp formula 6,4 • Spur formula 1 simple, 1 simple • Eyes present • Scrobes absent • Caste marked size-variation • Sting absent
The following information is derived from Barry Bolton's New General Catalogue, a catalogue of the world's ants.
- PROFORMICA [Formicinae: Formicini]
- Proformica Ruzsky, 1902d: 13 [as subgenus of Formica]. Type-species: Formica nasuta, by monotypy.
- [Proformica also described as new by Ruzsky, 1903b: 303.]
- Proformica subgenus of Formica: Wheeler, W.M. 1922a: 699; Emery, 1925b: 242; Creighton, 1950a: 457.
- Proformica raised to genus: Bondroit, 1918: 40; Kuznetsov-Ugamsky, 1927b: 26; Dlussky, 1969a: 218
Galkowski et al. (2017) - The taxonomy of the genus Proformica is complicated and in need of revision. The situation is particularly complex in the eastern area, with currently 23 species reported. In Western Europe, two distinct zoogeographical areas can be distinguished, the Iberian Peninsula and southern France, which are separated by a barrier formed by the Pyrenees mountain range. Three described species are currently recorded for the Iberian Peninsula (Collingwood 1976), but at least six forms are recognized by ant taxonomists (Xavier Espadaler, Barcelona, pers. comm.) and substantial morphological variation within each form makes species delimitation difficult. In contrast, only one described species, P. nasuta, has been recorded for southern France (Proformica ferreri may also be present in the French part of the Pyrenees).
Proformica nasuta is the type species for the genus Proformica and was described from Beaucaire, France. The concept of this species is unclear. For instance, variation in the number of erect hairs on the mesosoma, a character commonly used in the taxonomy of Proformica, has been interpreted either as mere intraspecific variation (Espadaler & Cagniant 1987), or as an indication that the name P. nasuta actually covers two taxa (Santschi 1925; Collingwood & Yarrow 1969). Populations of species of Proformica are small, inconspicuous and patchily distributed, and the species are often considered rare. As a consequence, the genus is poorly represented in institutional collections and most taxonomic work is based on few specimens, rendering the accurate perception of intra-specific variation difficult. Moreover, the type specimen of P. nasuta has not been located. Having not been found in the most likely candidate collections and not explicitly referred to in the literature, it is presumably lost.
Nylander (1856) described P. nasuta, the type species of the genus, from Beaucaire. Our analyses assigned workers from the type locality and from two other localities within a radius of 10 km (Jonquières and Tarascon) a single taxon. They lack erect hairs on the mesosoma, agreeing with the description of Proformica nasuta by Nylander as “nuda”. Although the type is presumably lost (as it could not be found in the following collections: Nylander (Helsinki) (Radchenko 2007), Forel (Geneva), Emery (Genoa), Bondroit (Brussels) and Santschi (Basel)), we are confident that the nest samples we collected in Beaucaire and in the surrounding area correspond to the species described by Nylander.
Consequences for the taxonomy of Proformica - There has been much confusion in the concept of the taxon P. nasuta.. None of the taxonomic studies published after the description of the species by Nylander (1856) made reference to the type, which was collected in Beaucaire (France). Forel (1886) described the worker and the queen from specimens collected in Orange (France) and sent samples to many of his colleagues throughout Europe and Russia. We examined the specimens from Orange in the Forel, Emery and Bondroit collections, and we collected new samples from the same locality in 2011. All differ markedly from those of Beaucaire and belong to the new species we describe in this study, P. longipilosa. All the taxonomic studies after 1886 used the samples from Orange, or descriptions of them, as a reference for P. nasuta. These studies described P. nasuta as having long erect hairs and sparse pubescence (e.g., Ruzsky 1905; Emery 1909; Wheeler 1913; Bondroit 1918; Santschi 1925; Bernard 1968; Dlussky 1969; Collingwood 1976; Agosti & Collingwood 1987), two characters that are typical of P. longipilosa. Other specimens from southeastern France (Plateau de Caussols, Tourettes-sur-Loup) have also been used as references for P. nasuta (Collingwood 1956; Stumper 1957; Dlussky 1969), but they come from an area that we now recognise as belonging to the range of P. longipilosa, and are thus likely distinct from P. nasuta. This mistake has been perpetuated so that the actual conception of P. nasuta refers to P. longipilosa. A consequence of this is that all reports of P. nasuta since 1886, including all those from eastern Europe and Asia, are probably erroneous.
- Agosti, D. 1991. Revision of the oriental ant genus Cladomyrma, with an outline of the higher classification of the Formicinae (Hymenoptera: Formicidae). Syst. Entomol. 16: 293-310 (page 295, Proformica in Formicinae, Formica genus group)
- Agosti, D. 1994b. The phylogeny of the ant tribe Formicini (Hymenoptera: Formicidae) with the description of a new genus. Syst. Entomol. 19: 93-117 (page 112, diagnosis, review of genus)
- Ashmead, W. H. 1905c. A skeleton of a new arrangement of the families, subfamilies, tribes and genera of the ants, or the superfamily Formicoidea. Can. Entomol. 37: 381-384 (page 384, Proformica in Formicinae, Lasiini)
- Bolton, B. 2003. Synopsis and Classification of Formicidae. Mem. Am. Entomol. Inst. 71: 370pp (page 130, Proformica in Formicinae, Formicini)
- Bondroit, J. 1918. Les fourmis de France et de Belgique. Ann. Soc. Entomol. Fr. 87: 1-174 (page 40, Proformica as genus.)
- Bondroit, J. 1918. Les fourmis de France et de Belgique. Ann. Soc. Entomol. Fr. 87: 1-174 (page 40, Proformica in Formicinae, Formicini)
- Creighton, W. S. 1950a. The ants of North America. Bull. Mus. Comp. Zool. 104: 1-585 (page 457, Proformica as subgenus of Formica)
- Dlussky, G. M. 1969a. Ants of the genus Proformica Ruzs. of the USSR and contiguous countries (Hymenoptera, Formicidae). Zool. Zh. 48: 218-232 (page 218, 231, Proformica raised to genus; Key to species of former U.S.S.R. and contiguous countries.)
- 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 687, Proformica as subgenus of Formica)
- Emery, C. 1925d. Hymenoptera. Fam. Formicidae. Subfam. Formicinae. Genera Insectorum 183: 1-302 (page 242, Proformica in Formicinae, Formicini; Proformica as subgenus of Formica)
- Forel, A. 1917. Cadre synoptique actuel de la faune universelle des fourmis. Bull. Soc. Vaudoise Sci. Nat. 51: 229-253 (page 250, Proformica in Camponotinae, Formicini; Proformica as subgenus of Formica)
- Kuznetsov-Ugamsky, N. N. 1927b. Zur Kenntnis der Gattung Proformica Ruzsky. Soc. Entomol. Stuttg. 42: 26-28 (page 26, Proformica as genus)
- Kuznetsov-Ugamsky, N. N. 1928a. Beiträge zur Ameisenfauna Mittelasiens. I. Die Gattung Proformica, Ruzsky 1903. Zool. Anz. 75: 7-23 (page 7, Proformica as genus)
- Ruzsky, M. 1902d. Material on the ant fauna of the Caucasus and the Crimea. Protok. Obshch. Estestvoispyt. Imp. Kazan. Univ. 206(su suppl: 1-33 (page 13, Proformica as subgenus of Formica)
- Ruzsky, M. 1903b. Essay on the myrmecofauna of the Kirghiz steppe. Tr. Rus. Entomol. Obshch. 36: 294-316 (page 303, Proformica also described as new)
- Wheeler, W. M. 1910b. Ants: their structure, development and behavior. New York: Columbia University Press, xxv + 663 pp. (page 143, Proformica as subgenus of Formica)
- 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 699, Proformica in Formicinae, Formicini; Proformica as subgenus of Formica)