Ants of Greece

AntWiki: The Ants --- Online

This webpage is based on Borowiec and Salata, 2022. This definitive, contemporary review of the ants of Greece was a major step towards making this ant fauna one of the better known in the Mediterranean region. The fauna will come into increasingly sharper focus as studies are published detailing the genera and species not included in the 2022 review, i.e., ants from the subfamily Myrmicinae. Information from such studies should be added to this page as they become available. If you know of a published study about Greece's ants that has not been incorporated here, feel free to add it - using your own Antwiki editor account!.

Below you will find information about Greece's geographic and ecological setting, a history of myrmecological research, and details about the subfamilies and genera that are present. There is a list of Greek ant species on a separate webpage, and information about each of the ants listed can be found on their respective Antwiki species pages. Comparative information and some short keys are given below under the generic listings, as are links to pages with keys for genera that have many Greek species.

Not included here is information from the 2022 review regarding various problematic species-complexes and undescribed species. The defining and highlighting of these taxonomic problems by Borowiec and Salata provides a guide for future taxonomic research in Greece.

Introduction

Greece physical map - provided by: Free world maps: http://www.freeworldmaps.net/europe/greece/map.html

Greece is a country of the Balkans, located in Southeastern Europe, bordered to the north by Albania, North Macedonia and Bulgaria; to the east by Turkey, and is surrounded to the east by the Aegean Sea, to the south by the Cretan and the Libyan Seas, and to the west by the Ionian Sea which separates Greece from Italy. The extreme points of Greece are: North: Ormenio village (41°45’41’’ N, 26°13’15’’ E), South: Gavdos island (34°48’11’’ N, 24°07’25’’ E), East: Strongyli island (36°06’17’’ N, 29°38’39’’ E), West: Othonoi island (39°51’11’’ N, 19°22’41’’ E). The country consists of a mountainous, peninsular mainland jutting out into the Mediterranean Sea at the southernmost tip of the Balkans, and two smaller peninsulas projected from it: the Chalkidiki and the Peloponnese, which is joined to the mainland by the Isthmus of Corinth. Greece also has many islands (1,200 to 6,000, depending on the minimum size to take into account, the number of inhabited islands is variously cited as between 166 and 227), of various sizes, the largest being Crete, Euboea, Lesbos, Rhodes, Chios, Kefalonia, Corfu, Limnos and Thassos; groups of smaller islands include the northern Aegean Islands, Cyclades, Dodecanese, Ionian Islands and North Sporades. According to the CIA World Factbook, Greece has 13,676 kilometers of coastline, the largest in the Mediterranean Basin.

Zoogeographically Greece is entirely placed in Mediterranean Province but it is also influenced by the Anatolian, Pontic and Iranian-Turanian faunas. The great distinctiveness of fauna of the eastern part of the Mediterranean basin causes that this region is often recognized as Eastern Mediterranean Subregion. The complex geological structure, numerous isolated mountain ranges, rich vegetation and partial location on numerous islands cause that individual regions of Greece differ greatly in biodiversity and demonstrate a high degree of endemism.

History

The very first publication focused on Greek ants, with description of new taxa, was a report from an expedition to the Peloponnese published by Brullé (1833). In the chapter "Famille de Formicaires" he noted 8 species of ants including two new to science: Formica nodus (now Cataglyphis nodus) and Formica pallidinervis (now synonym of Camponotus lateralis). Due to numerous taxonomic changes, the current status of the remaining 6 species mentioned in this study (Formica herculeana, F. rufa, F. cunicularia, F. nigra, Myrmica rubra and Atta capitata) is diffifficult to verify. Shortly after the mid-nineteenth century, Lucas (1854), in a work on arthropods from Crete, noted from this island Myrmica rubra (current status unclear, probably Aphaenogaster rugosoferruginea) and Formica pubescens (current status unclear, probably a Camponotus species). The next publicaton with descriptions of new taxa from Greece was Roger’s (1859) paper on some ants from the Mediterranean area. He designated Greece as terra typica for Atta splendida (now Aphaenogaster splendida), Formica aerea (now Proformica aerea), and Atta striola. The current status of the latter taxon is unclear as its type series consisted of specimens from Spain and Greece and most likely contained two different species. This name is usually treated as a junior synonym of Aphaenogaster gibbosa but Greek specimens probably represent Aphaenogaster epirotes. Additionaly, Roger described from Zakynthos Formica (Hypoclinea) kiesenwetteri (now Camponotus kiesenwetteri) and Stigmatomma denticulatum. He also noted from Zakynthos Formica (Hypoclinea) frauenfeldi (now Lepisiota frauenfledi) and Leptothorax recedens (now Temnothorax recedens), from Parnassos Mts. Formica truncata (now Colobopsis truncata), and generally from Greece Atta semipolita (current status unclear, probably Aphaenogaster subterranea or A. subterraneoides),Atta testaceopilosa (current status unclear, probably Aphaenogaster balcanica), and Formica cunicularia.

A number of works focused on Greek ants was authored by Auguste Forel (1848-1931). Forel published a number of important taxonomic and faunistic papers on ants from various parts of the world, including papers dedicated entirely to the ants of Greece (Forel 1886 b, 1889) which included descriptions of 10 new taxa. However, new taxa described from this country were included also in his other papers (Forel 1910, 1911 a, 1913 c). In the second half of the nineteenth century, Carlo Emery (1848-1925) joined the significant myrmecological researchers. His contribution to studies on Greek ants covers an important review on Cretan ants (Emery 1894 a) with descriptions of four new taxa (including endemic Aphaenogaster cecconii and Aphaenogaster simonellii), a paper with descriptions of Monomorium creticum and Aphaenogaster epirotes (Emery 1895), a note on ants of Cephalonia (Emery 1901) with description of two new taxa, and a paper on ants from Rhodes with descriptions of five new taxa (Emery 1915). A number of new taxa described from Greece can also be found in his other revisions (Emery 1869, 1906, 1908, 1922, 1925 a, 1925 b). Until the Second World War, Bruno Finzi (1897-1941) and Carlo Menozzi (1802-1943) significantly contributed to studies on the ants of the Greek fauna. The first researcher enriched the list of taxa known from Greece with a number of species (Finzi 1928, 1930 b, 1939). The second researcher deepened the knowledge on biodiversity of ants of the Dodecanese, describing the endemic to Greece or Aegean: Acropyga paleartica, Aenictus rhodiensis, Messor carpathous, Solenopsis crivellarii, Strongylognathus silvestrii, Temnothorax dessyi, and Temnothorax solerii. Felix Santschi (1872-1940), an extremely prolific myrmecologist having a reputation of taxonomic splitter, described from Greece Aphaenogaster sporadis and Messor hellenius, and four infrasubspecific taxa (Santschi 1926, 1927, 1929 c, 1933, 1934). After the Second World War, the first work on Greek ants was a note on species collected in Rhodes (Hamann & Klemm 1976), followed by two faunistic notes published by Legakis (1983, 1984).

Two milestones in studies on the ants of the Balkan Peninsula were the species key published by Agosti & Collingwood (1987) and annotated list of the ants of Greece compiled by Legakis (2011). Although both papers contain a number of errors, uncritically cite questionable taxa and list synonymic names as good taxa from Greece, they undoubtfully provided a good reference point for intensive study of ants from this region. Meanwhile, Greek fauna was continuously enriched with new discoveries, i.e. Douwes et al. (1988) described endemic Epimyrma adlerzi, Seifert (1992, 2003) described endemic Myrmica pelops and Lasius karpinisi, Schulz (1994) described endemic Aphaenoagaster graeca, and Mei (1998) described endemic Lasius myrmidon (now Metalasius myrmidon). Aditionally, Buschinger and Douwes (1993) reviewed parasitic anst of Greece and Collingwood (1993) provided some insights on ant biodiversity of some Greek islands.

The first step to organize the knowledge on ants in Greece was a critical checklist of ants by Borowiec & Salata (2012) published along with a supplement (Borowiec & Salata 2013). Our team continued faunistic research in various regions of Greece, also in cooperation with researchers from Greece, Slovenia and Turkey (Borowiec & Salata 2014 a, 2017 a, b, 2018 a, b, c, d, 2021 a, b, 2022, Borowiec et al. 2021, Bračko et al. 2016, Salata et al. 2020 a). Establishing the status of a number of taxa from Greece required taxonomic revisions and comparative studies with fauna from other regions (Salata & Borowiec 2015 a, b, c, 2017, 2018 a, b, 2019 a, b, Csösz et al. 2018, Salata et al. 2018 a, b, 2019 a, b, 2020 b, 2021). As a result of this work, a number of new species have been described from Greece and a number have been reported from Greece for the first time. Finally, important additions to the fauna of Greece, with descriptions of the new species, were published by Boer (2013), Seifert & Galkowski (2016) and Seifert (2019, 2020 a).

The Physical and Ecological setting

Although Greece is almost entirely situated within the Mediterranean region, it is characterized by a large number of habitats due to the huge horizontal and vertical diversity. Small lowlands are placed near the coast, and these are the Salonica, Thracian, Thessalian and Argolic Plains. As much as 81% of its territory is covered by mountains with an average altitude of 1,200–1900 m above sea level, which usually stretch longitudinally. On the western part, an extension of the Dinaric Mountains stretch from the border with Albania in a form of series of separate longitudinal massifs. Its highest peak, Mount Smolikas in the Pindos massif, reaches 2637 m above sea level. However, the highest mountain areas lie in the central-eastern part of the country, where the Olympus massif rises with the highest peak, Mitikas, 2,917 m above sea level. Mountainous landscape dominates also the Peloponnese, with the highest peak Ilias reaching 2404 m above sea level, and Aegean islands. The Cretan landscape is defined by large and high mountain ranges crossing from west to east and its highest peak, Idi, reaches 2456 m above sea level. Despite harsh and snowy winters, the highest mountain peaks of the Peloponnese are covered with mountain steppe habitats. Most likely it is due to warm summers characterized with high solar insolation and high temperatures which create conditions favorable for this habitat. Thus, the vegetation characteristic for the alpine zone does not occur on these massifs.

Mountains of the western Greece are limestone and sandstone, and were formed during the Alpine orogeny. The north-eastern Greek mountains, including the Olympus massif, were created during the Hercynian orogeny and are predominantly made of metamorphic rocks. Numerous karst formations can be found on the Peloponnese. Greece is a seismically active area and its insular mountains are predominantly volcanic. However, nowadays there is only one active volcano, Kajmeni, located on Santorini.

The soil of Greece is mostly rocky and of little use for agriculture. The decomposition of limestone in the mountain valleys creates sediments of alkaline soil that is used mainly for pasture. Only in the lowlands of Macedonia, Thrace, Attica and Thessaly alluvial soils are fertile and have agricultural potential. In northern Greece, rice cultivation developed on an extensive river alluvia.

A Mediterranean climate, with long, hot and dry summers and mild and humid winters, dominates most of the Greek territory. However, the latitudinal span of Greece is 6 °, which makes the climatic differences in the southern and northern parts of the country noticeable. The mountains located on western Greece and the Peloponnese experience quite heavy snowfall in winter. While the Greek islands and the eastern mainland’s coast have relatively high and stable temperatures thanks to the Aegean Sea influences. The northern part of Greece, where dominantes a more continental climate, is in general cooler and has higher precipitation than in the rest of the country. As a result, the vegetation of Thrace and Macedonia is richer than in other regions. The average temperature in these regions in January is around 5-7 ° C, in July around 25-27 ° C, and the average annual temperature of Northern Greece is 16 ° C. While in Central and Southern Greece the average temperature ranges from 8-10 ° C in winter, 27-28 ° C in summer, and the annual average is 18 ° C. On the southernmost islands (e. g. Rhodes and Crete), winters are even milder (average temperature 13 ° C), summers are hotter (average temperature 29 ° C), and the annual average temperature on these islands exceeds 19 ° C. Rainfall varies across the country. In the Ionian Islands, annually 1000 mm and more drops, sometimes even up to 2000 mm. In the Aegean islands, annual rainfall reaches 1000 mm. Quite different values prevail in mainland Greece. In the Pindos mountains to the east, average rainfall is 400 mm, in the lowlands of Thrace and Sterea Ellas, 384 mm of rain is at most. The summer drought lasts from two months in the west to five in the east. Due to the lack of rainfall and the generally high temperatures in summer, forest fires are among the frequent catastrophic events.

The mountainous landscape of Greece keeps the rivers short and full of rapids and waterfalls. Due to the uneven distribution of rainfall, the rivers in the country are characterized by a high variability of the water level. In the south, many small rivers dry up during the summer. The two large rivers running through Balkans have their estuaries on the territory of Greece: Evros and Strimonas. The other big rivers that run entirely through the territory of the country are: Haliacmon (297 km long), and Pineios (217 km long). There are only two large lakes within the borders of Greece. Lake Prespa, with an area of 288 km², is divided between Greece, Albania and North Macedonia. Whereas Lake Doiran, with an area of 41 km², is shared with North Macedonia. Additionally, there are around 30 smaller natural lakes, but some of them are largely drained or are in the final stage of overgrowing. There are also approximately 13 artificial lakes created as a result of the partitioning of the rivers, that range in size from 2.4 to 80.6 km².

Due to the long lasting activity of humans, the landscapes of Greece are severely transformed, and a large part of this country has been deprived of its original forest land. The warmest regions have vast areas covered with evergreen shrubs and macchia dwarf shrubs, which are referred to as phrygana. Due to pastoralism, phryganas are rarely compact and most often are divided by pastures overgrown with various herbal plants. Phryganas can form more compact habitats on islands, where goat breeding has been restricted. Intensive pastoralism on the lower mountain areas prompt formation of grassy and extensive pastures that are overgrown with shrubs on their peripheries. Whereas, mountain steppes-like habitats are formed on high mountain grazed plateaus. Olive trees are cultivated in many parts of Greece. They can take the form of intensive plantations characterized with poor vegetation or less intensive mosaic of trees mixed with the frigana scrub. The other commonly cultivated trees are figs and citruses.

Coastlines can be formed of cliffs overgrown with herbs or flat alluvia, which are often transformed into salinas. Forests are mainly found in the mountains. The large part of them consist of heavily exploited secondary forests. However, enclaves of natural deciduous forests and fir forests are preserved in the north-eastern Greece and in the higher parts of the mountains respectively. The arid lowlands on the mainland and islands are covered with pine forests. Because coniferous forests are prone to wildfires, wastelands in various stages of succession are common habitats on Greek lowlands. The moist habitats may be occupied by secondary mixed and deciduous forests, while stream valleys are commonly overgrown with gallery forests dominated by plane trees.

Subfamilies

The subfamilies of ants that occur in Greece are listed below, along with subfamily specific generic and species information.

There is a key to the subfamilies of Greek ants.

Aenictinae

One genus and one species.

Aenictus

Useful identification keys, revisions and taxonomic papers: Aktaç et al. 2004.

Diagnosis. Body linear; clypeus reduced; antennae 10-segmented, stout; antennal scape short, shorter than width of head; antennal sockets horizontal, in the plane of the transverse axis of head; palp formula 2, 2; eyes absent; promesonotal suture absent; propodeum unarmed; propodeal lobes present; waist of two separated segments; pygidium very small, reduced to a narrow U-shaped sclerite.

Comparative remarks: Aenictus rhodiensis is a distinct species and the only representative of army ants in this region. From members of Amblyponinae and Ponerinae subfamilies it differs in two segmented pedicel and shiny body surface. The most similar are members of the genus Leptanilla, but A. rhodiensis differs in mesosoma lacking promesonotal suture, absence of significant constriction on mesosoma in dorsal view, and in 10-segmented antennae. While Leptanilla species have distinct promesonotal suture, their mesosoma in dorsal view has a significant constriction, and they have 12-segmented antennae. Aenictus rhodiensis, with length above 2.5 mm, is also distinctly larger than any species of the Mediterranean Leptanilla which are considered as one of the smallest ants and usually their bodies are shorter than 2 mm.

Amblyoponinae

One genus with two species.

Stigmatomma

Useful identification keys, revisions and taxonomic papers: Baroni Urbani 1978.

Diagnosis. Body linear; clypeus small, anterior margin with a row of teeth; antennae 12-segmented, stout; antennal scape short, shorter than width of head; mandibles very elongate, masticatory margin with large, often bispinose teeth; palp formula 4,3 or 5,3; eyes very small, often look as a single small dot or reduced; promesonotal suture present; propodeum unarmed; waist of one large segment; first gastral segment separated from subsequent segments by a deep suture.

Stigmatomma Key / Comparative remarks:

  • Palpal formula 5:3, metasternum with small spiniforn process, eyes small but well visible . . . . . Stigmatomma impressifrons - differs from S. denticulatum in larger body size, with total length above 5.5 mm (in S. denticulatum below 4.5 mm), palpal formula 5:3 (in S. impressifrons 4:3), metasternum with spiniform process (absent in S. denticulatum), head slightly wider than long and eyes small but well visible.
  • Palpal formula 4:3, metasternum without spiniforn process, eyes rudimental reduced to the one ommatidium, almost invisible . . . . . Stigmatomma denticulatum - differs from S. impressifrons in smaller body size, with total length below 4.5 mm (in S. impressifrons above 5.5 mm), palpal formula 4:3 (in S. impressifrons 5:3), metasternum without spiniforn process (present in S. impressifrons) and eyes reduced to a single ommatidium, hardly visible.

Dolichoderinae

Five genera.

Key to the genera of Greek Dolichoderinae

Bothriomyrmex

Two species.

Useful identification keys, revisions and taxonomic papers: Seifert 2012 a.

Diagnosis. Monomorphic, very small, ML below 0.70; mandibles with 5-6 teeth; palp formula 4:3; antennae 12-segmented, no antennal club; antennal scape short, shorter than width of head, without projected setae; antennal sockets close to the posterior clypeal margin; eyes small, almost circular, placed distinctly in front of the middle of head; ocelli absent; mesothorax softly constricted in the middle; mesonotal groove shallow or only in form of mesonotal suture, metapleural gland present; propodeum unarmed; propodeal spiracle circular; petiole in form of a narrow erect scale; acidopore absent, apex of gaster only with transverse slit.

Bothriomyrmex Key / Comparative remarks:

  • Pubescence of first gaster tergite short, tergite surface well visible. Clypeal width small, ClyW/CS 0.736-0.771 . . . . . Bothriomyrmex corsicus - differs from B. communista in sparser and shorter appressed pubescence of gastral tergites which makes the surface of tergites well visible from under the hair. It has also broader clypeus with clypeal index 0.736-0.771.
  • Pubescence of first gaster tergite long, tergite surface less visible. Clypeal width small, ClyW/CS 0.773-0.820 . . . . . Bothriomyrmex communista - differs from B. corsicus in very dense and long appressed pubescence of gastral tergites which makes the surface of tergites hardly visible from under the hair. It has also narrower clypeus with clypeal index ClyW/CS 0.773-0.820.

A single literature record of Bothriomyrmex syria (as per Antmaps) suggests a third Bothriomyrmex species is present in Greece. It is unclear if this identification is correct or if this was a misidentified specimen of one of the two species noted above.

Dolichoderus

One species.

Diagnosis. Monomorphic, moderately large, ML above 1.0 mm; mandibles with 5-6 teeth; palp formula 4:3; antennae 12-segmented, no antennal club; antennal scape short, shorter than width of head, without projected setae; antennal sockets close to the posterior clypeal margin; eyes large, almost circular, placed in front of the middle of head; ocelli absent; mesothorax softly constricted in the middle; mesonotal groove deep, metapleural gland present; propodeum unarmed but with excavate posterior margin; propodeal spiracle circular; petiole in form of a thick erect scale; acidopore absent, apex of gaster only with transverse slit.

Comparative remarks: A very distinct species. Its body structure with very coarse punctation of head, very deep mesonotal groove, propodeum in profile with sharply angulate posterior angles, and characteristic coloration with black head, red mesosoma and black gaster with white maculae at base of first two gastral tergites is unique combination of characters. In the field, workers of D. quadripunctatus may appear similar to bicoloured minor workers of Colobopsis truncata which is often observed foraging on the same tree trunks or branches.

Linepithema

One invasive species.

Useful identification keys, revisions and taxonomic papers: Wetterer et al. 2009.

Diagnosis. Monomorphic, moderately large, ML below 1.2; mandibles with 12-18 teeth; palp formula 6:4; antennae 12-segmented, no antennal club; antennal scape elongate, longer than width of head, without projected setae; antennal sockets close to the posterior clypeal margin; eyes large, almost circular, placed distinctly in front of the middle of head; ocelli absent; mesothorax softly constricted in the middle; metapleural gland present; propodeum unarmed; propodeal spiracle circular; pectinate tibial spurs on meso- and metatibiae; petiole in form of a narrow erect scale, slightly inclined anteriorly; acidopore absent, apex of gaster only with transverse slit.

Comparative remarks: Distinct species. Linepithema humile is similar to the members of the genus Tapinoma but differs in eyes placed distinctly in front of the middle of head (in Tapinoma in the middle), clypeus without median emargination, only with a broad shallow concavity (in Tapinoma with median emargination) and petiolar scale prominent and well visible from above and in profile (in Tapinoma petiolar scale is very small, hidden under produced anteriorly first gastral tergite).

Liometopum

One species.

Useful identification keys, revisions and taxonomic papers: Del Toro et al. 2009. Diagnosis. Polymorphic, with strongly marked size-variation; mandibles with 7-10 teeth along masticatory margin, 3-5 teeth along basal margin; palp formula 6:4; antennae 12-segmented, no antennal club; antennal scape short, shorter than width of head; antennal sockets close to the posterior clypeal margin; eyes large, almost circular, placed on frontal surface distinctly in front of the middle of head with 90-140 ommatidia; ocelli present in major workers; metanotal groove reduced to a suture with the mesonotum and propodeum forming a continuous uninterrupted surface. All surfaces covered in dense pubescence; metapleural gland present; propodeum unarmed; propodeal spiracle circular; single tibial spurs on meso- and metatibiae; petiole in form of a narrow erect scale, hidden under anterior slope of first gaster tergite; acidopore absent, apex of gaster only with transverse slit.

Comparative remarks: A very distinct species. Its bicolored body and mesosoma regularly convex in profile are unique characters in the subfamily Dolichoderinae. In the field, members of this species look like large, bicolored workers of Lasius emarginatus or Lasius illyricus but both members of the subfamily Formicinae well differ in conical propodeum and deep mesonotal groove.

Tapinoma

Four species, and possibly two others.

Useful identification keys, revisions and taxonomic papers: Seifert 1984, 2012 b.

Diagnosis. Usually monomorphic; metanotal groove shallow; antennal segments 8, 11, 12; antennal club: absent to gradual; palp formula: 6,4; Total number of dentas 5-20(+); spur formula: 1 simple-pectinate, 1 simple-pectinate; Eyes with more than100 ommatidia; scrobes absent; pronotal spines, mesonotal spines, propodeal spines,petiolar spines: absen; monomorphic; metaplural gland present.

See Borowiec and Salata (2022) for details about the Greek morphospecies, Tapinoma cf. erraticum_BALC.

Formicinae

Fourteen genera.

Key to the genera of Greek Formicinae

Acropyga

One species.

Diagnosis. Monomorphic; mandibles with 5-9 teeth, tooth 3 smaller than tooth 4; palp formula 5:4, 4:3, 3:3; antennae 11-segmented; antennal scape short, without projected setae but with decumbent pubescence; antennal sockets very close to the posterior clypeal margin; eyes very small, placed in front of the midlength of head; mesothorax not constricted immediately behind the pronotum; mesonotum without stout setae; propodeum lacking projected setae; gaster without stout projected setae; metapleural gland present; propodeum unarmed; propodeal spiracle circular; tibial spurs on meso- and metatibia absent; petiole in form of unarmed scale; acidopore with a seta-fringed nozzle.

Comparative remarks: Acropyga paleartica resembles a very small specimens of the subgenus Cautolasius (=Lasius) but differs in 11-segmented antennae, very shallow mesonotal groove, short antennae and eyes placed distinctly in front of mid length of head. In field, workers may resemble members of Solenopsis because of their very small size but differs in characters of the subfamily Formicinae (pedicel with one segment in shape of scale and apex of gaster with circular nozzle-like acidopore, fringed with setae).

Anoplolepis

One (putative) species.

Diagnosis. Monomorphic; mandibles with 6-9 teeth, tooth 3 smaller than tooth 4; palp formula 6:4; antennae 11-segmented; antennal scape very long, with projected setae; antennal sockets close to the posterior clypeal margin; eyes large, placed behind the midlength of head; mesothorax constricted immediately behind the pronotum; mesonotum without projected setae; propodeum lacking projected setae; gaster without projected setae; metapleural gland present; propodeum unarmed; propodeal spiracle circular; tibial spurs on meso- and metatibia simple; petiole in form of unarmed node; acidopore with a seta-fringed nozzle.

Biological notes. In tropical countries this species is noted from open areas with less friable soils. It is one of the most common ant species which tends honeydew-producing hemipterans in Indonesia. In many tropical and subtropical countries, it is a well-known pest species, because it protects aphids and coccids which injure tropical crops. In this role it has been rated a secondary agricultural pest. It is unlikely that this species will acclimatize permanently in Greece and, if the record from Fauna Europea is based on some reliable data, it can only be treated as an accidental and temporary introduction.

Comparative remarks: Anoplolepis gracilipes: distinctive, a very slim and elongate body, extremely long and slim legs, very elonagted, 11-segmented antennae, big eyes and unarmed propodeum distinguish it well from other members of the subfamily Formicinae.

A Greek record of this introduced species is doubtful and needs confirmation.

Camponotus

Twenty-seven species in three subgenera.

Useful identification keys, revisions and taxonomic papers: Radchenko 1996 b, Ionescu-Hirsch 2010, Karaman & Aktaç 2013, Seifert 2019, Salata et al. 2019 b.

Diagnosis. Body not linear from small to very large, often polymorphic; clypeus broad with convex anterior margin, not armed with teeth; antennae 12-segmented, elongate; antennal scape moderately to very elongate, from slightly shorter to distinctly longer than width of head; antennal sockets in distance from posterior margin of clypeus; mandibles moderately elongate, masticatory margin with spiniform denticles of unequal size; palp formula 5,4 or 6,4; eyes large; promesonotal suture present, from shallow to deep; propodeum unarmed, but sometimes angulate or bituberculate; waist of one segment in form of moderately thick to thin scale; first gastral segment not separated from subsequent segments by a deep suture; mid and hind tibiae each with one pectinate spur; acidopore with a seta-fringed nozzle.

Key to subgenera of Camponotus of Greece

Key to Camponotus species of the subgenus Camponotus of Greece

Key to Camponotus species of the subgenus Myrmentoma of Greece

Key to Camponotus species of the subgenus Tanaemyrmex of Greece

Cataglyphis

Five species.

Useful identification keys, revisions and taxonomic papers: Radchenko 1998.

Diagnosis. Monomorphic to polymorphic or with marked-size variation, moderately large to very large. Mandibles with 5-7 teeth; palp formula 6:4; antennae 12-segmented, no antennal club; antennal scape elongate, distinctly longer than width of head, sometimes with projected setae; antennal sockets close to the posterior clypeal margin; eyes large, circular to elongate, placed slightly behind the midlength of head; ocelli present; mesothorax constricted in the middle; metapleural gland present; propodeum unarmed; propodeal spiracle elongate; simple tibial spurs on meso- and metatibiae; petiole in form of unarmed scale or node; acidopore with a seta-fringed nozzle.

Key to Cataglyphis of Greece

Colobopsis

one species

Useful identification keys, revisions and taxonomic papers: Radchenko 1996 b.

Diagnosis. Body not linear from small to moderately large, strongly polymorphic, major workers with strongly modified head; clypeus broad with convex anterior margin, not armed with teeth; antennae 12-segmented, moderately elongate; antennal scape shorter than width of head; antennal sockets in distance from posterior margin of clypeus; mandibles moderately elongate, masticatory margin with spiniform denticles of unequal size; palp formula 6,4; eyes large; promesonotal suture present, from shallow to deep; propodeum unarmed but angulate; waist of one segment in form of thin scale; first gastral segment not separated from subsequent segments by a deep suture; mid and hind tibiae each with one pectinate spur; acidopore with a seta-fringed nozzle.

Colobopsis truncata - Colobopsis and Camponotus are the only Greek genera with antennal insertions placed distinctly behind clypeal margin. Colobopsis by many years was treated as a subgenus of Camponotus but molecular studies showed that it represents a separate line within the subfamily and should be treated as distinct genus (Ward et al. 2016). Colobopsis differs from Camponotus in a special polymorphism manifested by sharply truncated heads in which the anterior truncated surface of the head in the major workers and the females is distinctly marginate and in which even the mandibles have a sharp external ridge separating an anterior from a latero-ventral face. In the members of Camponotus with distinct polymorphism the difference is manifested only in body size and different proportions of head but never in special structure of head. Colobopsis truncata differs also from members of the genus Camponotus in straight frontal carinae, slightly converging to the front (Camponotus has frontal carinae arched, distinctly converging to the front). In the field, minor workers of Colobopsis truncata are similar to Dolichoderus quadripunctatus especially due to pale spots on gaster and similar behavior. Both arboricolous species are observed foraging on tree trunks and do not show aggressive behavior towards each other. In Greek populations, minor workers of C. truncata usually have dark colored, reddish brown to brown mesosoma while workers of D. quadripunctatus usually has uniformly red mesosoma.

Formica

Fifteen species in four subgenera

Useful identification keys, revisions and taxonomic papers: Seifert 2000, 2002, 2018, Seifert & Schultz 2009, 2021.

Diagnosis. Monomorphic but sometimes with visible body size-variation, moderately large to large, mandibles with 7-10 teeth, third smaller than second and fourth; palp formula 6:4; antennae 12-segmented, no antennal club; antennal scape moderately elongate, longer than width of head, without projected setae; antennal sockets close to the posterior clypeal margin; eyes large, elongate oval, placed behind the midlength of head; ocelli present; mesothorax moderately constricted in its anterior half; metapleural gland present; propodeum unarmed; propodeal spiracle elongate oval; petiole in form of unarmed scale; fore tibia with broad, apical pectinate spur, mid and hind tibia with single spurs apically; acidopore with a seta-fringed nozzle.

Key to subgenera of Formica of Greece

Key to Formica species of the subgenus Formica of Greece (4 species)

Key to Formica species of the subgenus Serviformica of Greece (8 species)

Subgenus Coptoformica (2 species) - Key / Comparative remarks:

  • Erect seta on the surface of gaster tergites start with the first or second tergite, sometimes only as a row of setae at the posterior margin of the tergite. Microsetae on eyes at least in fraction of colony clearly protruding above ommatidiae (use magnification 60x or more) . . . . . Formica exsecta - Formica exsecta and F. bruni are the only Coptoformica species recorded from Greece. They look very similar and for distinguishing characters see comparative remarks under F. bruni.
  • Erect seta on the surface of gaster tergites start with the third or fourth tergite. Microsetae on eyes usually not protruding above ommatidiae (use magnification 60x or more) . . . . . Formica bruni - Formica bruni and F. exsecta are the only Coptoformica species recorded from Greece. Formica exsecta differs in microsetae on eyes at least in fraction of colony clearly protruding above ommatidiae (in F. bruni these microsetae are short, not protruding above ommatidiae). Aditionally, F. bruni has first two or three gastral tergites lacking erected setae on dorsal surface and their posterior margin is lacking a row of setae. While F. exsecta has at least second gastral tergite with few setae close to posterior margin and sometimes dorsal surfaces of first and second tergites have also erected setae. However, in Greece, was observed hitherto only “rubens” morph of F. exsecta which is lacking erected setae on dorsal surface of first two gastral tergites.

Subgenus Raptiformica (one species)

Formica sanguinea differs from all large Greek Formica species with not emarginated occipital margin of head in presence of median emargination on anterior margin of clypeus.

Lasius

Thirty-two species in five subgenera.

Useful identification keys, revisions and taxonomic papers: Seifert 1988, 1992, 2020.

Diagnosis. Body not linear from small to moderately large, not polymorphic; clypeus broad with convex anterior margin, not armed with teeth; antennae 12-segmented, moderately elongate without club; antennal scape slightly shorter to slightly longer than width of head; antennal insertions close to posterior margin of clypeus; mandibles moderately elongate, masticatory margin with spiniform denticles of unequal size; palp formula 6,4; eyes large; promesonotal suture present, shallow; propodeum unarmed but often convex; waist of one segment in form of thin scale; first gastral segment not separated from subsequent segments by a deep suture; mid and hind tibiae each with one spur.

Taxonomical note. Recently Boudinot et al. (2021), based on molecular data, described Metalasius – a new ant genus that consists of one extant species Metalasius myrmidon and one fossil species †Metalasius pumilus. Metalasius myrmidon is known only from Greece and was originally described as Lasius myrmidon. As the revision of Lasius was published recently we couldn’t implement proposed changes in our monograph. Metalasius differs from Lasius in eyes situated in anterior half of head as measured in full-face view (eyes situated in posterior half of head in Lasius), reduced metapleural gland orifice with opening directed posteriorly (metapleural gland orifice small to very large and opening laterally as well as posteriorly in Lasius), propodeal spiracle situated in lower half of propodeum in profile view (propodeal spiracle situated at or above midheight of propodeum in Lasius), antennomere III broader than long (usually longer than broad in Lasius), and hypostoma lacking carina along lateral margin (carina present in Lasius).

Key to Metalasius and subgenera of Lasius of Greece - use this key to determine which of the following keys to use to identify a specimen to species

Key to Lasius species of the subgenus Lasius of Greece (17 species)

Key to Metalasius and Lasius species of the subgenus Cautolasius of Greece (3 species)

Key to Lasius species of the subgenus Chthonolasius of Greece (11 species)

Lasius of the subgenus Austrolasius Key / Comparative remarks:

  • More hairy species, fore femora and tibiae on lateral surface with numerous suberected to erected setae . . . . . Lasius carniolicus - no records from Greece but may be present

Both Greek members of the subgenus Austrolasius are very similar and difficult to identification without comparative materials. Lasius carniolicus is generally more hairy and setose with higher number of semierect hair and erected setae on both ventral and dorsal sides of head and lateral surface of femora and tibia (usually > 11 semierect hair and erected setae of length 10 μm while in L. reginae the number of semierect hair and erected setae is usually < 7). For more detailed morphometric differences see Seifert (2018 p. 127).

  • Less hairy species, fore femora and tibiae on lateral surface without or with only a few suberected to erected setae . . . . . Lasius reginae

See comparative remarks above for Lasius carniolicus.

Leptanillinae

One genus and an undetermined number of species; at least three distinct morphospecies.

Leptanilla

Useful identification keys, revisions and taxonomic papers: Baroni Urbani 1977, Kugler 1986, Scupola & Ballarin 2009, Ward & Sumnich 2012, Griebenow 2020.

Diagnosis. Minute ants, pedicel with two distinct segments; antennae 12-segmented, stout; antennal scape short, shorter than width of head; frontal lobes absent so that antennal sockets are fully exposed and located very close to the anterior margin of the head; eyes absent; promesonotal suture present, deeply impressed and articulated; propodeum unarmed; propodeal lobes absent; pygidium larger and conspicuous, not armed with teeth or spine; sting present.

Note. Ward & Sumnicht (2012) recorded the genus Leptanilla from Greece based on a series of 14 males taken from a blue pan trap set in the open woodland on the island of Rhodes (Dodecanese). Molecular and morphological data suggest that these specimens represent three distinct morphotypes. The poor state of the Leptanilla taxonomy hinders identification of the Greek male specimens. So far, eighteen species of Leptanilla have been described from the Mediterranean area and additionally seven unnamed species were noted only from males. Eleven of these are based on workers (or workers and queens), six are known from males only, and there is no Mediterranean species with both sexes and worker caste described (Ward & Sumnicht 2012). Because males are usually collected separately from the workers, a parallel taxonomy has developed for the two castes (Bolton 1990).

Biological notes. Little known. In Spain a complete colony was found during the excavation of a Leptothorax nest and numerous workers were collected in large soil samples composed of moist sand with a low organic content from slopes of small seasonal water courses (Barandica et al. 1994, Lopez et al. 1994). Males of Leptanilla are usually caught using light traps or blue pan traps.

See Borowiec and Salata 2022 for notes about the morphospecies and Greek Leptanilla taxonomy.

Ponerinae

Three genera.

Key to the genera of Greek Ponerinae

Cryptopone

One species.

Diagnosis. Body linear; clypeus narrow with convex anterior margin, not armed with teeth; antennae 12-segmented, stout; antennal scape short, shorter than width of head; mandibles moderately elongate, masticatory margin with spiniform denticles of equal size; palp formula 2,2; eyes very small, often look as a single small dot; promesonotal suture present; propodeum unarmed; waist of one segment in form of thick scale; first gastral segment not separated from subsequent segments by a deep suture; mid and hind tibiae each with one pectinate and one spiniform spur.

Cryptopone ochracea, at first glance, is similar to the species of Ponera and Hypoponera but differs in mandibles with 6-7 strong dents of approximately equal size, mid and hind tibiae each with two spurs, median spur large and pectinate and lateral spur much smaller and not pectinate. In both relative genera mandibles have more than 8 dents and denticles and the 1-3 apical dents are often somewhat larger than the following 8-13 dents, and mid and hind tibiae have only one pectinate spur. Body and legs vestiture in C. ochracea is longer and denser than in both related genera with several erected setae on first two gastral tergites and tibiae.

Hypoponera

Two species.

Useful identification keys, revisions and taxonomic papers: Seifert 2003, Bolton & Fisher 2011, Seifert 2013.

Diagnosis. Body linear; clypeus narrow with convex anterior margin, not armed with teeth; antennae 12-segmented, stout; antennal scape elongate, slightly shorter than width of head; mandibles elongate, masticatory margin with spiniform denticles of unequal size; palp formula 1,2; eyes very small, with at most 5 ommatidia; promesonotal suture present but shallow; propodeum unarmed; waist of one segment in form of thick scale; petiolar base with rounded lobe, anteriorly without transparent window; first gastral segment not separated from subsequent segments by a deep suture; mid and hind tibiae each with one pectinate spur.

Biological Note: H. punctatissima is less common than H. eduardi and was recorded only from anthropogenic habitats.

Hypoponera Key / Comparative remarks:

  • Scape reaching to hind margin of head, SL/CW > 0.88. Petiole in profile significantly higher and narrower. Mesopleuron completely covered with carinulate sculpture . . . . . Hypoponera eduardi
  • Scape not reaching to hind margin of head, SL/CW < 0.88. Petiole in profile significantly lower and thicker. Mesopleuron smooth . . . . . Hypoponera punctatissima

Both Greek members of the genus Hypoponera are superficially similar and differ in subtle characters. Hypoponera eduardi looks less shiny and has mesopleuron completely covered with carinulate sculpture while H. punctatissima looks slightly shiny and has mesopleuron smooth. Hypoponera eduardi has slightly longer scapi reaching to hind margin of head, SL/CW > 0.88 and petiole in profile significantly higher and narrower, approximately 1.7 times as high as wide. While in H. punctatissima scape is not reaching to hind margin of head, SL/CW < 0.88 and petiole in profile is significantly lower and thicker, approximately 1.5 times as high as wide.

Ponera

Two species.

Useful identification keys, revisions and taxonomic papers: Csősz & Seifert 2003.

Diagnosis. Linear ants, pedicel with one distinct segments; palp formula 2,2; antennae 12-segmented, stout; mandibles stoutly triangular; clypeus simple, without extended lobes; eyes minute or rudimental; antennal scape short, shorter than width of head, funicle gradually incrassate with an apical club; frontal lobes absent so that antennal sockets are fully exposed and very close to the anterior margin of the head; eyes absent; promesonotal suture present, deeply impressed and articulated; propodeum unarmed; propodeal lobes absent; petiolar base in profile with a circular translucent “window” anteriorly; pygidium larger and conspicuous, not armed with teeth or spine; a strong sting is present.

Biological Notes. Ponera coarctata is more ubiquistic than P. testacea and usually prefers shady habitats, especially in lowlands. Ponera coarctata is an ubiquistic forest species, noted from luminous pine forests, mountain deciduous and coniferous forests, stream valleys with plane forests, occasionally from open habitats like mountain pastures but from altitude above 900 m. Nesting always under stones. Collecting sites come from an altitude 88-1400 m. Ponera testacea prefers open, warm and dry habitats such as warm forests and pastures with oaks.

Ponera Key / Comparative remarks:

  • Petiolar node in profile higher and narrower, petiole width to high of petiolar node 1.062-1.260. On average larger CW 0.52-0.60. Body color from yellow to almost black . . . . . Ponera coarctata
  • Petiolar node in profile lower and thicker, petiole width to high of petiolar node 1.105-1.422. On average smaller CW 0.47-0.56. Body color usually from yellow to red, only occasionally partly brown . . . . . Ponera testacea

Both members of Ponera are superficially similar and difficult to identify. Only dark brown specimens of P. coarctata can be distinguished because Greek populations of Ponera testacea are never completely brown. Pale forms of both species differ mainly in shape of petiole, higher and narrower in P. coarctata (petiole width to high of petiolar node 1.062-1.260, see Seifert 2018 p. 80) and lower and thicker in P. testacea (petiole width to high of petiolar node 1.105-1.422). A dark form of P. testacea described as Ponera coarctata var. lucida Emery, 1898 at first glance is very similar to P. coarctata and differs only in structure of petiole and slightly smaller size. This form was recorded mainly from the eastern part of the distribution range of P. testacea (Turkey, including western part, and Azerbaijan) but has never been found in Greek materials (Csősz et al. 2022).

Proceratiinae

One genus with three species.

Proceratium

Useful identification keys, revisions and taxonomic papers: Baroni Urbani & de Andrade 2003.

Diagnosis. Body not linear, dorsal profile forms arch; clypeus small, armed with an anterior projection; antennae 12-segmented, stout; antennal scape short, shorter than width of head; palp formula 2, 2 or 4, 3; eyes very small, often look as a single small dot; promesonotal suture absent; propodeum unarmed; waist of one segment; first gastral segment separated from subsequent segments by a deep suture.

Additional Resources

References

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