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Research Article
Preliminary division of not socially parasitic Greek Temnothorax Mayr, 1861 (Hymenoptera, Formicidae) with a description of three new species
expand article infoSebastian Salata, Lech Borowiec§
‡ California Academy of Sciences, San Francisco, United States of America
§ University of Wrocław, Wrocław, Poland
Open Access

Abstract

The division of Greek members of the genus Temnothorax into 17 morphological groups is proposed. Temnothorax aveli species group is reviewed with three species: T. turcicus (Santschi) (North Aegean Islands, Sterea Ellas, Peloponnese and Thessaly), and two species new to science: Temnothorax brackoi sp. nov. (Epirus, Ionian Islands, Macedonia, Peloponnese, western Sterea Ellas, Thessaly, and also Dalmatia in Croatia), and T. messiniaensis sp. nov. (Ionian Islands and Peloponnese); a new species Temnothorax triangularis sp. nov., a member of the Temnothorax nylanderi species group is also described (Sterea Ellas: Euboea Island).

Keywords

Balkan Peninsula, Temnothorax aveli group, Temnothorax nylanderi group, taxonomy

Introduction

The Myrmicinae genus Temnothorax Mayr, 1861 is one of the most speciose in the Mediterranean Region (sensu Vigna Taglianti et al. 1999). Among 279 Palearctic taxa (Bolton 2019), 194 are known from this region (Borowiec 2014, Radchenko et al. 2015, Salata and Borowiec 2015, Csősz et al. 2015, 2018, Galkowski and Lebas 2016, Galkowski and Cagniant 2017, Sharaf et al. 2017, Catarineu et al. 2017, Salata et al. 2018). Most of the recent research performed on this genus was focused on taxa from European and Anatolian parts of the Mediterranean Region. Those areas are considered as most diverse and species-rich, and each further study confirms this assumption. Greece, located at the joint of European and Anatolian subunits, hosts numerous and various Temnothorax taxa and so it is one of the most challenging fields of research. Study on Greek Temnothorax is additionally impeded by a complicated taxonomical history of several taxa and a lack of new material. Moreover, recent studies published by Csősz et al. (2015, 2018) revealed that species considered as a widespread represent in fact complexes of cryptic taxa.

As most of Temnothorax species are morphologically variable, to determine their distribution range and morphological variability, researchers ought to investigate rich material. During several field trips to various places on Greek islands and the mainland, we collected more than 1000 samples of Temnothorax species from 480 localities. The material collected during those expeditions helped us to establish morphological variability within species and divide collected material into several species groups. In most cases, a division into species group is based on species morphology only, and we did not verify if it corresponded with the phylogeny of this genus. This method is a widely accepted tool helping in determining the range of studied material and is commonly practiced (Csősz et al. 2018, Salata and Borowiec 2015, Galkowski and Cagniant 2017, Catarineu et al. 2017). However, we adjusted our species-group definitions to phylogenetical data provided by Prebus (2017) and Csősz et al. (2015).

Some Greek Temnothorax have been already included in studies on species groups of T. nylanderi (Csősz et al. 2015), T. interruptus (Csősz et al. 2018), T. recedens (Salata and Borowiec 2015), and T. muellerianus (Salata and Borowiec 2015). Below we present our division of Temnothorax from Greece, which will be used as a base for further studies and we describe three new species: one of them, Temnothorax triangularis, is a member of recently revised T. nylanderi species group (Csősz et al. 2015). Two others are a widespread Greek species, members of the Temnothorax aveli species group. Because Temnothorax taxa known from Crete were revised recently (Salata et al. 2018), and the majority of its endemic species creates single-species groups unknown from other Greek regions, they were excluded from this paper. Also, socially parasitic members of Temnothorax, often classified in separate genera Chalepoxenus Menozzi and Myrmoxenus Ruzsky, are not included in this study (see discussion in Seifert et al. 2016 and Ward et al. 2016).

Materials and methods

Most of the material was sampled between 2007 and 2019 from sites in different parts of Greece. The main method was direct sampling (hand collecting). Individual specimens and nests were collected on the ground, in leaf litter and rock rubble, under stones, on tree trunks, and in dry twigs of herbs. This method was occasionally supplemented by litter sifting and collecting material with an entomological umbrella. All specimens were preserved in 75% EtOH. Study was supported with material deposited in the collection of G. Bračko (Ljubljana, Slovenia). All studied type specimens of taxa mentioned in differential diagnosis or characteristics are listed below.

Examined specimens are housed in the following collections:

BMNH Natural History Museum, London;

CASC California Academy of Sciences, San Francisco, California, USA;

DBET Department of Biodiversity and Evolutionary Taxonomy, University of Wrocław, Poland;

MHNG Museum d’Historie Naturelle, Geneve, Switzerland;

HNHM Hungarian National History Museum, Budapest, Hungary;

NHMW Natural History Museum, Vienna, Austria;

MNHW Museum of Natural History, University of Wrocław, Wrocław, Poland;

MZLS Museum of Zoology, Lausanne, Switzerland;

NHMB Naturhistorisches Museum Basel, Switzerland;

PWC Petr Werner collection, Prague, Czech Republic;

SMNG Senckenberg Museum für Naturkunde Görlitz, Görlitz, Germany;

ZMHB Museum für Naturkunde, Zentralin-stitut der Humboldt-Universität, Berlin, Germany.

To determine a distribution range and a morphological variability of the new species we compared them with material collected from other Greek regions. Data concerning samples used in the comparison is provided in series of regional checklists (Borowiec and Salata 2012, 2013, 2017a, b, 2018a, b, c, d, e, Bračko et al. 2016) and we see no reason to repeat this information.

Specimens were compared using standard methods of comparative morphology. All measurements were made in μm using a pin-holding stage, permitting rotations around X, Y, and Z axes. A Nikon SMZ18 stereomicroscope was used at a magnification of ×100 for each character. Photographs were taken using a Nikon SMZ 1500 stereomicroscope, Nikon D5200 camera and Helicon Focus software. All given label data of type specimens are in original spelling, presented in square brackets; a vertical bar (|) separates data on different rows and double vertical bars (||) separate labels. Images of type specimens are available online on AntWeb (www.AntWeb.org) and are accessible using the unique CASENT or FOCOL identifying specimen code. If not stated differently material deposited in Museum of Natural History, University of Wrocław (in permanent deposit in the Department of Biodiversity and Evolutionary Taxonomy).

Pilosity inclination degree applies to this used in Hölldobler and Wilson (1990). The adpressed (0–5°) hairs run parallel, or nearly parallel to the body surface. Decumbent hairs stand 10–15°, subdecumbent hair stands 30°, suberect hairs stand 35–45°, and the erect hairs stand more than 45° from the body surface.

Measurements:

EL eye length; measured along the maximum vertical diameter of eye;

EW eye width; measured along the maximum horizontal diameter of eye;

HL head length; measured in straight line from mid-point of anterior clypeal margin to mid-point of posterior margin in full-face view;

HW head width; measured in full-face view directly above the eyes;

PEH petiole height; measured in lateral view, the chord of ventral petiolar profile at node level is the reference line perpendicular to which the maximum height of petiole is measured;

PEL petiole length; measured in lateral view, from anterior corner of subpetiolar process to dorsocaudal corner of caudal cylinder;

PNW pronotum width; maximum width of pronotum in dorsal view;

PPH postpetiole height; measured perpendicularly to a line defined by the linear section of the segment border between dorsal and ventral petiolar sclerite;

PPL postpetiole length; maximum length of the postpetiole measured in lateral view perpendicular to the straight section of lateral postpetiolar margin;

PPW postpetiole width; maximum width of postpetiole in dorsal view;

PSL propodeal spine length; measured from the centre of the propodeal spiracle to the top of the propodeal spine in lateral view;

PW petiole width; maximum width of petiole in dorsal view;

SDL spiracle to declivity length; minimum distance from the centre of the propodeal spiracle to the propodeal declivity;

SL scape length; maximum straight-line length of scape excluding the articular condyle;

WL mesosoma length; measured as diagonal length from the anterior end of the neck shield to the posterior margin of the propodeal lobe.

Indices:

CI HW/HL * 100;

EI1 EW/EL * 100;

EI2 EW/HL * 100;

SI1 SL/HL * 100;

SI2 SL/HW * 100;

MI PNW/WL * 100;

PI PL/PH * 100;

PPI PPL/PPH * 100;

PSI PSL/SDL * 100.

Abbreviations:

g. gyne;

m. male;

w. worker.

We decided to list all other ant species collected from the same localities as species new to science. In our opinion, it provides valuable information about ecosystem structure and species diversity characteristic for habitats preferred by these species.

Type material of species noted in the comparative diagnoses

Temnothorax affinis (Mayr, 1855), syntype (w.): Oesterreich || Type || Leptothorax | affinis | Mayr || 29889 || affinis | Mayr || GBIF-D/FFoCol | 2006 (ZMHB); syntype (w.): Oesterreich || Type || Leptothorax | affinis | Mayr || GBIF-D/FoCol | 2007 (ZMBH), available from photo in AntWeb (FOCOL2006);

Temnothorax aveli (Bondroit, 1918), syntype (w.): Lept. aveli | Bondr. || Type || Leptothorax | Aveli Bondr. | type – Sayat || Sammlung | Dr. F. Santschi | Kairouan || ANTWEB | CASENT0912907 (NHMB), available from photo in AntWeb (CASENT0912907);

Temnothorax artvinensis Seifert, 2006, holotype (w.): ARTVIN – 5km SW Artvin | 100 Mh 1164 | Kiefernwald 50% | Leg. Schulz 27.06.93 TURKEI || Holotype | Temnothorax | artvinense | SEIFERT || GBIF-D/FoCol | 1361 (SMNG), available from photo in AntWeb (FOCOL1361);

Temnothorax graecus (Forel, 1911), lectotype (w.): L. bulgaricus | For. | r. graecus | type Forel | Patras … | … (Forel) || Lectotype | Leptothorax graecus | Forel, 1911 top specimen | det. A. Schulz & M. Verhaagh 1999 || Typus || r. d. graecus | Forel || Coll. | A. Forel || ANTWEB | CASENT0909017 (MHNG), examined; paralectotype (w.): the same pin as lectotype, bottom specimen (MHNG), examined; paralectotypes (2w.): Typus || L. bulgaricus | For. | r. graecus | Forel | type | Amaroussia | p. Athenes (Forel) || coll. | A. Forel (MHNG), examined.

Temnothorax kemali (Santschi, 1934), syntype (w.): Leptothorax | kemali Sant | Type || Izmir | 29.VII.33 | Santschi || Type || Leptothorax | kemali | A. Schulz det || Basel 10 || ANTWEB | CASENT0912952 (NHMB), available from photo in AntWeb (CASENT0912952);

Temnothorax laconicus Csősz, Seifert, Müller, Trindl, Schulz & Heinze, 2015, paratype (w.): Temnothorax laconicus || Temnothorax laconicus | Csosz et al. 2014 | PARATYPE || PARATYPE || GREECE, GRE 2011-0345 | Taygethos Oros, Street to Profiti | Ilias | 36.968N, ; 22.404E, ; 800 m | leg. A. Schulz, 01.05.2011 || ANTWEB | CASENT0914696 (HNHM), available from photo in AntWeb (CASENT0914696);

Temnothorax lagrecai (Baroni Urbani, 1964), paratype (w.): Caltagirone | Sichel leg. || BOSCO S. PIETRE | 15.IV.62 || TYPUS || ANTWEB | CASENT0919741 (NHMW), available from photo in AntWeb (CASENT0919741)

Temnothorax nigriceps (Mayr, 1855), syntype (w.): L. tubero | nigriceps | Mt. Tendre | … || Typus || Coll. | A. Forel ||ANTWEB | CASENT0909038 (MHNG), examined;

Temnothorax rabaudi (Bondroit, 1918), paralectotype (w.): Leptothorax | Rabaudi | Boundr. | type | Gironde || ANTWEB | CASENT0904752 (MSNG), available from photo in AntWeb (CASENT0904752)

Temnothorax sordidulus (Müller, 1923), syntype (w.): No: 175 Types | L. carinthiacus | Bernard || TYPE || 20.4.56 | … || Carinthia | Viktring | Holzel leg. || ANTWEB | CASENT0907632 (MZLS), available from photo in AntWeb (CASENT0907632);

Temnothorax tauricus (Ruzsky, 1902), cotypes, (3w.): L. unifasciatus | … | r. tauricus | Ruzsky | Crimee || Cotypus || v. L. tauricus | Ruzsky || 1288 || Coll. Forel || ANTWEB | CASENT0909049 (MHNG), examined; cotypes (3w.): L. unifasciatus | Latr. | tauricus | Ruzsky | Crimee || 1288 || cotype (MHNG), examined.

Taxonomy

Species groups of Greek Temnothorax

The number of Temnothorax taxa known from Greece is estimated at 59 (including undescribed species) and include recent data published in taxonomic revisions and faunistic papers (Salata and Borowiec 2018). Below we present series of morphological characters that were used in species-groups division. Some of the species groups were defined in former publications and we used those descriptions as a base for our research. This applies to the following cases: bulgaricus species group (Radchenko 1995a); affinis species group, tuberum species group, corticalis species group, and clypeatus species group (Radchenko 1995b); nylanderi species group (Radchenko 1995c, Csősz et al. 2015); exilis species group (Cagniant and Espadaler 1997); interruptus species group (Csősz et al. 2018); and recedens species group (Salata and Borowiec 2015). However, in some cases we adapted those definitions to Greek species and thus they differ from the original definitions. We also list members of each species group known from Greece. Taxa known and described from Crete are not included in this division, as they were recently revised in a separate paper (Salata et al. 2018), and most of them create separate, single species groups, so far known only from this island.

Temnothorax affinis species group: antennae 12-segmented, club not or only slightly darkened, metanotal groove absent, body colouration orange to dark orange with darker head and dark first gastral tergite, propodeal spines very long and thin, straight or only slightly curved, petiole node subangular in profile, head and mesosoma surface moderately sculptured, head completely microsculptured and with more or less developed longitudinal ridges. In Greece only one arboreal species, T. affinis (Mayr).

Temnothorax angustulus species group: antennae 12-segmented, club darkened, metanotal groove feebly marked, body colouration mostly brown to almost black, including head, first gastral tergite mostly brown with yellowish brown basal spot, propodeal spines short to long but thin, straight or slightly curved, petiole node angulate in profile, head and mesosoma surface finely sculptured, head only partly microsculptured, central part more or less shiny, longitudinal ridges diffused visible only in ocular area and sides of head. In Greece only one species known from Mediterranean coniferous forests, T. dessyi (Menozzi).

Temnothorax anodontoides species group: antennae 12-segmented, club darkened, metanotal groove absent, body colouration brown, dark brown to almost black including head, first gastral tergite completely brown to almost black not or only slightly paler at base, propodeal spines very short with broad base, not longer than basal width, petiole node rounded in profile, head and mesosoma surface very strongly sculptured appear partly rugose, head background microsculptured, whole surface with longitudinal ridges or rugosities. In Greece at least three undescribed species known only from high montane localities.

Temnothorax aveli species group: antennae 12-segmented, completely yellow but club sometimes slightly darkened, metanotal groove absent, body colouration mostly yellow including head, sometimes only gena and femora slightly darkened, first gastral tergite with narrow to broad dark posterior band, always with paler spot at base, propodeal spines short to moderately long, with broad base, petiole node rounded in profile, head and mesosoma surface finely sculptured, head almost uniformly, regularly microsculptured, mostly without longitudinal ridges, sometimes only central part of head with narrow area of diffused microreticulation, but head never appears smooth and shiny – in Greece three, revised below, species mostly known from lowland habitats, especially Mediterranean forests and bushes, nestling inside dry stems of trees and twigs of bushes and large herbs– T. brackoi sp. nov., T. messiniaensis sp. nov., T. turcicus (Santschi, 1934), (?)T. tauricus (Ruzsky) (see comments in the redescription of T. turcicus).

Comments: Members of the aveli species group are morphologically similar to those assigned to the tuberum species group. Despite morphological similarities taxa of both groups can be easily distinguished based on nesting and habitat preferences. Species of the aveli species group inhabit lowlands and have nests inside dry stems of trees, bushes and herbs. While Greek species of the tuberum species group occur in the mountain areas and nest in soil, under moss, in rock crevices or debris.

Temnothorax bulgaricus species group: antennae 12-segmented, club usually distinctly darkened, metanotal groove absent, body colouration mostly yellow, sometimes head partly darkened, first gastral tergite with broad dark posterior band, always with yellow spot at base, propodeal spines reduced to triangular tubercle, petiole node rounded in profile, head in central part more or less smooth and shiny, on sides with longitudinal ridges, sometimes almost whole surface of head smooth and shiny or smooth area reduced to a narrow medial stripe, mesosoma laterally with strong sculpture of longitudinal ridges. In Greece two species associated with humid habitats such as stream valleys or bushes inside dark deciduous forests, T. bulgaricus (Forel) and T. nadigi (Kutter).

Temnothorax clypeatus species group: antennae 12-segmented, completely yellow, metanotal groove inconspicuous but well-marked, body colouration ochraceous to reddish brown, first gastral tergite with broad dark posterior band, always with pale spot at base, propodeal spines moderate to long, thorn-shaped, petiole with short peduncle, appears high and bulky in profile, head sculpture from complete, with microreticulate background and more or less developed ridges and costae, to mostly smooth and shiny in central part, only on gena and around eyes with longitudinal ridges, mesosoma laterally with distinct microreticulation and strong reticulate and longitudinal sculpture. In Greece one arboreal species associated with deciduous trees, especially large oaks in sunny habitats, T. clypeatus (Mayr).

Temnothorax corticalis species group: antennae 12-segmented, completely yellow, metanotal groove absent, body colouration ochraceous to light brown, head always more or less darkened, first gastral tergite with broad dark posterior band, always with pale spot at base, propodeal spines from very short, triangular to moderately long needle-shaped, petiole with very short peduncle, appears triangular in profile with more or less angulate node, head sculpture from complete, with more or less reticulate sculpture but shiny background, occasionally in central part of frons reticulation partly diffused, mesosoma dorsally and laterally with distinct microreticulation and often sides of pronotum with longitudinal ridges. In Greece one arboreal species associated with deciduous forests, T. corticalis (Schenck), but occurrence of T. jailensis (Arnoldi) is possible.

Temnothorax exilis species group: antennae 12-segmented, club usually distinctly darkened, metanotal groove absent, body colouration extremely variable, from almost completely yellow to black, often mesosoma paler coloured than head and gaster, first gastral tergite in pale forms with dark posterior band, propodeal spines moderate to long, thin, straight, petiole node angulate in profile, head in central part more or less smooth and shiny, on sides usually with longitudinal ridges, sometimes almost whole surface of head smooth and shiny or smooth area reduced to a narrow medial stripe, mesosoma laterally with strong sculpture of longitudinal ridges. Xerothermophilous species associated with rocky, open and arid habitats; in Greece only T. exilis (Emery) recorded, but based on a high variability of insular populations the real number of species of this group is difficult to estimate and requires further studies.

Temnothorax flavicornis species group: antennae 11-segmented, unicolourous yellow, metanotal groove present, body colouration almost completely yellow to dark yellow, without distinct contrast between colouration of head and mesosoma, first gastral tergite with dark posterior, propodeal spines from long to very long, claw-shaped, from straight to slightly curved, petiole with moderately long peduncle and node angulate, head always with microreticulate background and more or less developed reticulate or costulate sculpture, along middle of head runs stripe with diffused reticulation, more or less smooth and shiny, mesosoma with microreticulate sculpture, often with distinct ridges or costae. One species associated with various arboreal habitats, T. flavicornis (Emery).

Temnothorax graecus species group: antennae 12-segmented, club usually distinctly darkened, metanotal groove absent, body colouration mostly yellow, first gastral tergite with broad dark posterior band, always with yellow spot at base, propodeal spines very short to short, triangular to needle-shaped, petiole node rounded in profile or obtusely angulate, head in central part more or less smooth and shiny, on sides with longitudinal ridges, sometimes almost whole surface of head smooth and shiny or smooth area reduced to a broad medial stripe, mesosoma laterally with moderate sculpture of longitudinal ridges. Associated mostly with moderately humid to arid deciduous forests or mediterranean bushes, collected on rocks and stones; T. aeolius (Forel), T. graecus (Forel), T. smyrnensis (Forel), and at least two undescribed species.

Temnothorax interruptus species group: antennae 12-segmented, club usually distinctly darkened, frontal lobes conspicuously wider than frons, metanotal groove absent or indistinct, body yellow to light brown, gena darker, first gastral tergite with dark posterior band, often interrupted in the middle, propodeal spines very long, thin and curving downwards, petiole node subangulate to obtuse in profile, head with distinct microreticulation and longitudinal ridges, often partly with large reticulate sculpture, mesosoma mostly microreticulate, dorsally and laterally more or less rugose or costulate. In Greece two species associated with open habitats such as rocks and stones overgrown by bushes or limestones on mountain pastures, collected also in deciduous or mixed forests, and occasionally in coniferous forests; T. morea Csősz, Salata & Borowiec and T. strymonensis Csősz, Salata & Borowiec.

Temnothorax kemali species group: antennae 12-segmented, club usually more or less darkened, occasionally whole antennae yellow, metanotal groove absent, body yellow to orange, gena usually darker, first gastral tergite with dark posterior band, propodeal spines long and thin apically often curving downwards, petiole node subangulate to obtuse in profile, head at least in central part without microreticulation, smooth and shiny, only gena and area around eyes with longitudinal ridges, in some specimens sculpture of sides and central part of head more distinct but area between ridges or costae always smooth and shiny. Species associated with Mediterranean herbs and bushes or dry deciduous and coniferous forests, often nestling inside dry stems of herbs; T. kemali (Santschi) and at least one undescribed species.

Temnothorax luteus species group: antennae 12-segmented, club usually more or less darkened, occasionally whole antennae yellow, metanotal groove absent, body yellow to orange, gena sometimes darker, first gastral tergite with dark posterior band, propodeal spines straight, long and thin, petiole node subangulate in profile, head most often without microreticulation, smooth and shiny, only gena and area around eyes with longitudinal ridges. Xerothermophilous species associated with lowland habitats, Mediterranean herbs and bushes; at least one undescribed species.

Temnothorax nylanderi species group: diverse and speciose group, antennae 12-segmented, unicolourous yellow, metanotal groove usually distinct but in some species inconspicuous, body colouration variable, from almost completely yellow to dark brown but usually without distinct contrast between colouration of head and mesosoma, first gastral tergite usually with dark posterior band (except dark species with unicolourous gaster), propodeal spines from very short, triangular to very long, claw-shaped, from straight to distinctly curved, petiole with moderately long peduncle and node from angulate to obtuse in profile, head always with microreticulate background and more or less developed reticulate or costulate sculpture, sometimes along middle of head runs stripe with diffused reticulation, more or less smooth and shiny, mesosoma with microreticulate sculpture, often with distinct ridges or costae. Species associated with various shadowy habitats, nesting in rock, stones, and dry branches inside forests; T. angulinodis Csősz, Heinze & Mikó, T. angustifrons Csősz, Heinze & Mikó, T. ariadnae Csősz, Heinze & Mikó, T. crasecundus Seifert & Csősz, T. crassispinus (Karavaiev), T. helenae Csősz, Heinze & Mikó, T. laconicus Csősz, Seifert, Müller, Trindl, Schulz & Heinze, T. lichtensteini (Bondroit), T. lucidus Csősz, Heinze & Mikó, T. nylanderi (Foerster), T. parvulus (Schenck), T. sordidulus (Müller), T. subtilis Csősz, Heinze & Mikó, T. tergestinus (Finzi), and T. triangularis sp. nov.

Comment: Results presented by Prebus (2017) revealed that the nylanderi species group, as defined by Csősz et al. 2015, is paraphyletic and the position of T. flavicornis within it is unlikely. Therefore, we decided to exclude this species from this group and consider it as the single representative of the flavicornis species group.

Temnothorax recedens species group: antennae 12-segmented, unicolourous yellow to brown, metanotal groove very deep, body colouration variable, from almost completely yellow to dark brown, often mesosoma paler coloured than head and gaster, first gastral tergite in pale forms with dark posterior band, propodeal spines from short, triangular to long and thin, the shape of the needle, straight to slightly curved, petiole with long peduncle and node obtuse in profile, head mostly smooth and shiny, pronotum almost completely shiny, mesonotum and propodeum laterally with microreticulate sculpture, without distinct ridges or costae. Species associated with various arboreal habitats, nesting in rock crevices or under moss; T. antigoni (Forel), T. recedens (Nylander), T. rogeri Emery, and T. solerii (Menozzi).

Comment: With great probability this group is more speciose than it is apparent from current data, especially T. recedens shows high regional variability and wide ecological variance, which suggests that this taxon is a group of cryptic species

Temnothorax rottenbergi species group: very large species, antennae 12-segmented, club darkened or whole antennae dark, metanotal groove inconspicuous, body completely black or distinctly bicoloured with head and gaster brown to black and mesosoma partly to completely red, propodeal spines very long and strong, apex of spines often curving downwards, petiole with long peduncle and globular node, head with strong reticulate sculpture, mesosoma dorsally and laterally with strong, partly reticulate and partly costate sculpture. Xerothermophilous and alpine species associated with rock and stones on open, sunny habitats such as mountain pastures, grasslands or edges of forests. From Greece T. rottenbergi (Emery) and T. semiruber (André) were recorded but occurrence of first species needs confirmation.

Temnothorax tuberum species group: diverse group, antennae 12-segmented, club darkened, metanotal groove absent or indistinct, mesosoma colouration variable, from almost completely yellow to ochraceous, head always darker coloured than mesosoma, in extreme case almost black, first gastral tergite always with dark posterior band, propodeal spines from short, triangular to moderately long but never needle-shaped, from straight to slightly curved apically, petiole with moderately long peduncle and node subangulate in profile, head always with microreticulate background and more or less developed reticulate or costulate sculpture, without smooth and shiny areas, mesosoma with microreticulate sculpture, often with distinct ridges or costae – species associated with various habitats, from open and sunny to shadowy arboreal, usually nesting in rocks or stones, most species were noted also on rocks on mountain pastures – T. melanocephalus (Emery), T. nigriceps (Mayr), T. tuberum (Fabricius), T. unifasciatus (Latreille), and several undescribed taxa.

Comment: Results presented by Prebus (2017) suggest that the tuberum species group (sensu Cagniant and Espadaler 1997) and unifasciatus species group (sensu Bernard 1967) are paraphyletic. In both cases species groups were defined based on West-Mediterranean taxa. Only two Greek members of the tuberum species group were included in analysis presented by Prebus (2017): T. nigriceps and T. unifasciatus and they created a separate cluster. Confirmation, if the tuberum species group as defined by us here is a natural, monophyletic group requires further studies.

A key to Temnothorax species groups known from Greece*

1 Metanotal groove present, distinct to inconspicuous 2
Metanotal groove absent, mesosoma in lateral view evenly convex 7**
2 Antennae 11-segmented T. flavicornis species group
Antennae 12-segmented 3
3 Head and mesosoma almost entirely smooth and shiny; metanotal groove very deep; antennal scape very long and thin T. recedens species group
Head and mesosoma sculptured, sometimes only frons and mesosomal dorsum with reduced or absent sculpture; metanotal groove deep to indistinct; antennal scape short to moderate 4
4 More or less large species; body completely black or distinctly bicoloured with head and gaster brown to black and mesosoma partly to completely red; propodeal spines very long and sharp, apex of spines often curving downwards; petiole with long and more or less thin peduncle and globular node, head with sparse and thick reticulae and, at least on frons, smooth interspaces T. rottenbergi species group
Moderately sized species; body colouration variable, from almost completely yellow to dark brown but without distinct contrast between colouration of head and mesosoma; propodeal spines short to long, straight or only slightly curved; petiole with peduncle short to moderately long, and node from angulate to obtuse in profile; head with sculpture variable, but always fine and dense, interspaces usually with additional microsculpture 5
5 Promesonotal suture distinct; body colouration ochraceous to reddish brown; anterior margin of clypeus with distinct notch T. clypeatus species group
Promesonotal suture absent or indistinct; body yellow to dark brown and never ochraceous or reddish; anterior margin of clypeus without distinct notch 6
6 Antennal club darkened; metanotal groove feebly marked; body colouration more or less uniform, mostly brown to almost black; first gastral tergite with yellowish brown basal spot; propodeal spines short to long but thin, straight or only slightly curved T. angustulus species group
Antennal club unicolourous, yellow; metanotal groove usually distinct but in some species inconspicuous; body colouration variable, from almost completely yellow to dark brown but never uniformly coloured (at least legs and antennae brighter); first gastral tergite with dark posterior band or uniformly coloured; propodeal spines from very short, triangular to very long, claw-shaped, always with wide base T. nylanderi species group
7 Frontal lobes conspicuously wider than frons; first gastral tergite with dark posterior band, often interrupted in the middle; propodeal spines very long, thin and curving downwards T. interruptus species group
Frontal lobes never conspicuously wider than frons; dark posterior band on the first gastral tergite never interrupted in the middle or absent; propodeal spines of a different shape 8
8 Propodeal spines very long and thin and never with wide base 9
Propodeal spines very short to moderately long, always with wide base 11
9 Body orange to dark orange with darker head and first gastral tergite with bright spot on its basal part; head and mesosoma surface moderately sculptured, head completely microsculptured and with more or less developed longitudinal ridges T. affinis species group
Body yellow to orange, gena sometimes darker, first gastral tergite with dark posterior band; head, at least in central part, without microreticulation, smooth and shiny 10
10 Propodeal spines long and thin, apically often curving downwards; head microsculptured, only its central part with microreticulation reduced or absent T. kemali species group
Propodeal spines straight, long and thin; head most often without microreticulation, smooth and shiny T. luteus species group
11 Body colouration uniform, brown, dark brown to almost black; propodeal spines very short with broad base, not longer than basal width; petiole node rounded in profile; head and mesosoma surface very strongly sculptured; alpine species T. anodontoides species group
Body bicoloured or entirely yellow; if body uniformly dark brown to black then propodeal spines small to moderately long, and frons and dorsal surface of mesosoma with reduced sculpture or smooth 12
12 Propodeal spines indistinct, strongly reduced to triangular tubercles; petiolar node obtuse in profile T. bulgaricus species group
Propodeal spines short to moderately long, always distinct; petiolar node from obtuse to angulate in profile 13
13 Head sculpture variable, more or less sparse and fine; at least centre of frons and central part of promesonotal dorsum smooth; propodeal spines moderately long to long; body colouration variable, from almost completely yellow to completely black; also within populations of the same species body colouration variable T. exilis species group
Head with dense and fine sculpture, frons sometimes with sculpture reduced but never smooth; sculpture on mesosoma never reduced or smooth. If head and dorsum of promesonotum with reduced sculpture, then propodeal spines very small; body colouration from mostly yellow to light brown or bicoloured but constant within a single species 14
14 Body distinctly bicoloured, yellow to light brown with head usually darker than mesosoma and gaster with broad dark band 15
Body almost entirely yellow to dark yellow; only head sometimes with slightly darker frons or malar area and gaster usually with narrow dark band 16
15 Antennal club never darkened; propodeal spines from very short, triangular to moderately long needle-shaped, petiole with very short pedicel; head with moderately dense reticulate sculpture but smooth background, occasionally in central part of frons reticulation partly diffused T. corticalis species group
Antennal club darkened; propodeal spines from short, triangular to moderately long but never needle-shaped; petiole with moderately long pedicel; head always with microreticulate background and moderately developed reticulate or costulate sculpture, without smooth and shiny areas T. tuberum species group
16 Propodeal spines short to moderately long with broad base; head and mesosoma surface finely sculptured, head almost uniformly, regularly microsculptured, mostly without longitudinal ridges, sometimes only central part of head with narrow area of diffused microreticulation, but head never appears smooth and shiny; petiolar node angulate in profile T. aveli species group
Propodeal spines very short to short, triangular to needle-shaped; head in central part more or less smooth and shiny, on sides with longitudinal ridges, sometimes almost whole surface of head smooth and shiny or smooth area reduced to a broad medial stripe; petiolar node rounded to subangular in profile T. graecus species group

Review of Greek species of the Temnothorax aveli species group

Temnothorax brackoi sp. nov.

Figs 1, 2, 9, 13

Type material

Holotype: worker (pin), (CASENT0846669): GREECE, Pel. Messinia | 1.8 km E of Saidona, 985 m | 36.88491N, 22.30419E | 20 VI 2016, L. Borowiec || Collection L. Borowiec | Formicidae | LBC-GR02195 (MNHW).

Paratypes: • 11w. (pin), 9w. (EtOH) (CASENT0846607-CASENT0846617): the same nest sample as holotype (DBET); • 11w. (pin), 5w. (EtOH)(CASENT0846618-CASENT0846628): GREECE, Pel. Messinia | 0.8 km SE of Exochori, 535 m, | 36.89582N, 22.27464E | 20 VI 2016, L. Borowiec || Collection L. Borowiec | Formicidae | LBC-GR02184 (DBET, MHNG, CASC, BMNH); • 8w. (pin), 1w. (EtOH) (CASENT0846629-CASENT0846636): GREECE, Pel. Messinia | n. Arachova, 860 m | 37.03922N, 22.21876E | 13 VI 2016, L. Borowiec || Collection L. Borowiec | Formicidae | LBC-GR02029 (DBET); • 2w. (pin) (CASENT0846637-CASENT0846638): GREECE, Pel. Messinia | 1.3 km S of Artemisia, 870 m | 37.08738N, 22.23378E | 17 VI 2016, L. Borowiec || Collection L. Borowiec | Formicidae | LBC-GR02121 (DBET).

Other material

GREECE. Epirus, Preveza: • 4w. (pin) (CASENT0846670-CASENT0846673), 21w. (EtOH): 700 m E of Kanali, oak forest, 39.0638N/20.70621E, 50 m, 2016-08-27, leg. L. Borowiec; • 8w. (pin) (CASENT0846674-CASENT0846681), 14w. (EtOH): n. Kanali, oak forest, 39.05458N/20.70207E, 15 m, 2016-08-28, leg. L. Borowiec; • 27w (EtOH): n. Kanali, pine forest, 39.05388N/20.70118E, 20 m, 2016-08-28, leg. L. Borowiec. Ionian Islands, Cephalonia: • 6w. (pin) (CASENT0846682-CASENT0846687): Avithos Lake, area near a small lake in a moist, shaded valley of a small creek, 38.17293N/20.71233E, 278 m, 2014-06-25, leg. L. Borowiec; • 68w. (EtOH): Avithos Lake, shrubs around small lake, 38.17203N/ 20.71107E, 288 m, 2019-06-10, leg. L. Borowiec; • 29w. (EtOH): Katapodata; roadsides with shrubs, 38.23337N/20.64594E, 100 m, 2019-06-10, leg. L. Borowiec; • 4w. (EtOH): 1.5 km NE of Koulourata, mixed forest on shrubs, 38.20667N/ 20.67715E, 273 m, 2019-06-10, leg. L. Borowiec; • 11w. (EtOH): ancient Same; roadsides with shrubs, 38.2522N/20.66423E, 220 m, 2019-06-10, leg. L. Borowiec; • 13w. (EtOH): rd. Skala-Poros; Mediterranean shrubs, 38.12872N/20.79576E, 5 m, 2019-06-12, leg. L. Borowiec. Ionian Islands, Corfu: • 6w. (EtOH): Ag. Ilias, deciduous forest, 39.79367N/19.87508E, 191 m, 2013-06-10, leg. L. Borowiec; • 5w. (pin) (CASENT0846688-CASENT0846692), 5w. (EtOH): Akr. Kefali n. Agios Stefanos, frygana on sea coast, 39.75154N/19.63272E, 13 m, 2013-06-05, leg. L. Borowiec; • 4w. (EtOH): E of Nymfes, old olive tree plantation, 39.75047N/19.8057E, 179 m, 2013-06-09, leg. L. Borowiec; • 8w. (pin) (CASENT0846693-CASENT0846700), 1w. (EtOH): Klimatia, deciduous forest, 39.74123N/19.78953E, 311 m, 2013-06-06, leg. L. Borowiec; • 1w. (pin) (CASENT0846701): N of Ag. Stefanos, old olive forest, 39.76338N/19.65213E, 88 m, 2013-06-05, leg. L. Borowiec; • 4w. (pin) (CASENT0846702-CASENT0846705): n. Doukades, old olive tree plantation, 39.70075N/19.75055E, 174 m, 2013-06-08, leg. L. Borowiec; • 3w. (pin) (CASENT0846706-CASENT0846708), 1w. (EtOH): n. Vistonas, old olive tree plantation, 39.68549N/19.71453E, 422 m, 2013-06-08, leg. L. Borowiec; • 20w. (pin) (CASENT0846709-CASENT0846728), 15w. (EtOH): Nymfes, deciduous forest, 39.75478N/19.79535E, 162 m, 2013-06-06, leg. L. Borowiec; • 20w. (EtOH): Old Perithia, old mountain village, 39.76159N/19.87412E, 467 m, 2013-06-10, leg. L. Borowiec; • 3w. (pin) (CASENT0846729-CASENT0846731): Strinilas, deciduous forest, 39.73963N/19.84265E, 632 m, 2013-06-07, leg. L. Borowiec. Ionian Islands, Lefkada: • 3w. (pin) (CASENT0846732-CASENT0846734), 2w. (EtOH): Asprogerakata, stream valley with deciduous forest, 38.78278N /20.65262E, 430 m, 2016-09-02, leg. L. Borowiec; • 2w. (pin) (CASENT0846735-CASENT0846736), 8w. (EtOH): Egklouvi, stream valley with deciduous forest, 38.73212N/20.64821E, 690 m, 2016-09-02, leg. L. Borowiec. Ionian Islands, Zakynthos: • 4w. (EtOH): 1 km E of Elies, shrubs along roadsides, 37.90173N/20.68787E, 300 m, 2018-05-06, leg. L. Borowiec; • 18w. (EtOH): 1 km N of Exo Chora, mixed forest, 37.81063N/20.68459E, 430 m, 2018-05-08, leg. L. Borowiec; • 1w. (pin) (CASENT0846737): 1.2 km N of Ano Vasilikos, roadsides along olive plantation and pasture with oak shrubs, 37.72456N/20.97786E, 30 m, 2018-05-05, leg. L. Borowiec; • 128w. (EtOH): 1.2 km N of Vasilikos, roadsides along olive plantation and pasture with oak shrubs, 37.72456N/20.97786E, 30 m, 2018-05-05, leg. L. Borowiec; • 2w. (EtOH): 1.2 km NE of Anafonitria shrubs around burned forests, 37.85489N/20.64124E, 475 m, 2018-05-10, leg. L. Borowiec; • 1w. (pin) (CASENT0846738), 43w. (EtOH): 1.2 km SW of Ano Vasilikos loc. 1, roadsides along olive plantation, 37.72374N/20.95964E, 72 m, 2018-05-05, leg. L. Borowiec; • 1w. (pin) (CASENT0846739), 25w. (EtOH): 1.2 km SW of Ano Vasilikos loc. 2, roadsides along olive plantation, 37.72325N/20.96123E, 75 m, 2018-05-05, leg. L. Borowiec; • 1w. (pin): 1.2 km SW of Skinaria, limestone hills after burned forests, 37.87694N/20.69272E, 375 m, 2018-05-06, leg. L. Borowiec; • 1w. (pin) (CASENT0846740), 13w. (EtOH): 1.4 km E of Ano Volimes, mixed forest, 37.87594N/20.68525E, 430 m, 2018-05-06, leg. L. Borowiec; • 1w. (EtOH): 1.5 km N of Exo Chora: shrubs around meadow, 37.81446N/ 20.68826E, 460 m, 2018-05-08, leg. L. Borowiec; • 38w. (EtOH): 1.7 km NE of Ano Volimes, shrubs around pastures, 37.88627N/20.68456E, 445 m, 2018-05-06, leg. L. Borowiec; • 3w. (EtOH): 1.8 km SW of Volimes, shrubs along roadsides, 37.86472N/20.64234E, 350 m, 2018-05-10, leg. L. Borowiec; • 1w. (EtOH): 1.9 km W of Maries, roadsides in burned forests, 37.818N/20.65556E, 290 m, 2018-05-09, leg. L. Borowiec; • 1w. (pin), 30w. (EtOH): 2.5 km NE of Maries, mixed forest, 37.8299N/20.70151E, 475 m, 2018-05-08, leg. L. Borowiec; • 51w. (EtOH): 3.9 km NE of Maries, shrubs around olive plantation, 37.84202N/20.70938E, 370 m, 2018-05-08, leg. L. Borowiec; • 1w. (pin) (CASENT0846741), 16w. (EtOH): 470 m NE of Orthonies, shrubs along roadsides, 37.85435N/20.69843E, 405 m, 2018-05-10, leg. L. Borowiec; • 1w. (EtOH): 500 m S of Apelati, pine forest, 37.66873N/20.81407E, 160 m, 2018-05-07, leg. L. Borowiec; • 1w. (EtOH): 700 m SW of Koroni, maquis, 37.86582N/20.71753E, 290 m, 2018-05-10, leg. L. Borowiec; • 3w. (EtOH): 750 m S of Volimes, shrubs in cypress forest, 37.86762N/20.66191E, 360 m, 2018-05-10, leg. L. Borowiec; • 1w. (pin) (CASENT0846742), 81w. (EtOH): 800 m SE of Xirokastello, roadsides along olive plantation, 37.73491N/20.95139E, 75 m, 2018-05-05, leg. L. Borowiec; • 10w. (EtOH): 880 m S of Orthonies, shrubs in cypress forest, 37.84462N/20.69843E, 390 m, 2018-05-10, leg. L. Borowiec; • 66w. (EtOH): Ag. Georgiou monastery, shrubs along roadsides, 37.85971N/20.63646E, 330 m, 2018-05-10, leg. L. Borowiec; • 8w. (EtOH): Ag. Joannis, roadsides with shrubs, 37.72924N/20.94553E, 165 m, 2018-05-05, leg. L. Borowiec; • 6w. (EtOH): Argassi, urban area, 37.76182N/20.92704E, 10 m, 2018-05-04, leg. L. Borowiec; • 1w. (EtOH): Livia Mts. loc. 1, shrubs along roadsides, 37.83018N/20.71619E, 600 m, 2018-05-08, leg. L. Borowiec; • 15w. (EtOH): Vrachionas Mts., mountain pastures with shrubs, 37.81798N/ 20.70621E, 670 m, 2018-05-08, leg. L. Borowiec; • 1w. (EtOH): W of Kampi, pine forest, 37.78161N/20.68078E, 165 m, 2018-05-09, leg. L. Borowiec. Macedonia, Pieria: • 4w. (EtOH): Platamonas Castle hill, on stones, soil and herbs, 40.00868N/ 22.59654E, 12 m, 2019-05-11, leg. L. Borowiec; • 1w. (pin) (CASENT0846743): 2 km W of Panteleimonas, roadsides with shrubs, 39.98563N/ 22.59513E, 305 m, 2019-05-15, leg. L. Borowiec; • 20w. (EtOH): road to P. Poroi loc. 1, roadsides with shrubs, 39.97963N/ 22.61563E, 110 m, 2019-05-17, leg. L. Borowiec; • 7w. (EtOH): road to P. Poroi loc. 2, roadsides with shrubs, 39.97627N/ 22.61146E, 185 m, 2019-05-17, leg. L. Borowiec; •81w. (EtOH): road to P. Poroi loc. 3, roadsides with shrubs, 39. 96863N/22.60494E, 260 m, 2019-05-17, leg. L. Borowiec. Peloponnese, Korinthia: • 16w. (pin) (CASENT0846744-CASENT0846759): n. Evrostina, deciduous forest, 38.07386N/22.39388E, 662 m, 2013-09-01, leg. L. Borowiec. Peloponnese, Laconia: • 13w. (pin) (CASENT0846760-CASENT0846772), 2w. (EtOH): Taygetos Mts., 1.5 km SW of Anavryti, coniferous forest, 37.0191N/22.36003E, 990 m, 2016-06-21, leg. L. Borowiec. Peloponnese, Messinia: • 1w. (pin) (CASENT0846773): Taygetos Mts., 0.7 km S of Dyrrachio, coniferous forest, 37.1811N/22.2074E, 800 m, 2016-06-22, leg. L. Borowiec; • 5w. (pin) (CASENT0846774-CASENT0846778): Taygetos Mts., 2 km W of Arachova, stream valley with Platanus forest, 37.0357N/22.1978E, 680 m, 2016-06-13, leg. L. Borowiec; • 1w. (pin) (CASENT0846779): Taygetos Mts., Chora Getson, stream valley with deciduous forest, 36.94779N/22.25466E, 615 m, 2016-06-14, leg. L. Borowiec; • 4w. (pin) (CASENT0846780-CASENT0846783): Taygetos Mts., Karveli, rest area with stream, 37.07591N/22.20633E, 600 m, 2016-06-17, leg. L. Borowiec; • 2w. (pin) (CASENT0846784-CASENT0846785): Tetrazi Mts., 0.5 km E of Vastas, stream valley with deciduous forest, 37.36441N/21.99304E, 895 m, 2016-06-19, leg. L. Borowiec; • 1w. (pin) (CASENT0846786), 2w. (EtOH): Tetrazi Mts., Isaris, pine forest, 37.36644N/22.01516E, 790 m, 2016-06-19, leg. L. Borowiec; • 1w. (pin) (CASENT0846787): Tetrazi Mts., Karnasi, stream valley with Platanus forest, 37.31904N/21.99158E, 460 m, 2016-06-19, leg. L. Borowiec. Sterea Ellas, Aetolia-Acarnania: • 1w. (EtOH): Psila Alonia, stream valley with Platanus forest, 38.96255N/21.19527E, 62 m, 2016-09-04, leg. L. Borowiec. Sterea Ellas, Euboea: • 2w. (EtOH): 1.2 km NW of Gerontas, mediterranean shrubs along roadsides, 38.45885N/23.808E, 405 m, 2018-06-10, leg. L. Borowiec; • 2w. (EtOH): 2.3 km S of Stropones, mixed forest, 38.9933N/23.87807E, 860 m, 2018-06-10, leg. L. Borowiec; • 1w. (EtOH): 300 m NW of Agios, pine forest with mediterranean shrubs, 38.65856N/23.55525E, 600 m, 2018-06-11, leg. L. Borowiec; • 7w. (EtOH): 570 m NW of Drosia, stream valley with mixed forest, 38.61705N/23.59089E, 140 m, 2018-06-11, leg. L. Borowiec. Thessaly, Larissa: • 1w. (pin) (CASENT0846788): Mt. Olympus, 5.3 km E of Olympiada, alpine pastures with shrubs, 40.00989N/22.31096E, 800 m, 2017-05-09, leg. L. Borowiec; • 7w. (EtOH): Mt. Ossa, 2.4 km SE of Karitsa, stream valley in deciduous forest, 39.82632N/22.77557E, 425 m, 2017-05-04, leg. L. Borowiec; • 12w. (EtOH): Mt. Ossa, 600 m SE of Karitsa, stream valley in deciduous forest, 39.84009N/22.76983E, 300 m, 2017-05-04, leg. L. Borowiec; • 3w. (pin) (CASENT0846789-CASENT0846791), 44w. (EtOH): Mt. Ossa, Kokkino Nero, stream valley in urban area, 39.83389N/22.79379E, 3 m, 2017-05-10, leg. L. Borowiec. CROATIA, Podgorje: • 4w. (pin) (CASENT0846792-CASENT0846795), vicinity Sibenj, 9 km S Senj, 0–50 m, 3.6.1997, A. Schulz & K. Vock leg.

Locus typicus

Peloponnese, Taygetos Mts.

Differential diagnosis

Temnothorax brackoi is most similar to T. aveli (Bondroit, 1918), a species widely distributed in the western part of the Mediterranean basin, east to Slovenia. It is similar to T. aveli especially in body shape, petiolar structure and unicolour yellow antennae. Temnothorax aveli differs in microreticulation of head not as regular as in T. brackoi, with slightly shiny surface between sculpture and often with presence of median stripe of diffused microsculpture on frons, while in T. brackoi head sculpture is always homogenous, without areas of diffused microreticulation and dull surface between sculpture. From Greek members of Temnothorax, T. brackoi is most similar to sympatric T. graecus, but differs from it in well-developed sculpture on the whole head surface (in T. graecus central part of frons is more or less shiny, without microreticulation) and larger, distinctly triangular propodeal spines (in T. graecus spines are short, needle-shaped); some specimens of T. brackoi can also be confused with species belonging to the T. tuberum group, but T. brackoi differs from them in low and arched petiolar node and lack of longitudinal striation on frons.

Description

Worker (n = 10): HL: 0.637 ± 0.04 (0.584–0.683); HW: 0.547 ± 0.03 (0.509–0.696); SL: 0.437 ± 0.02 (0.410–0.560); EL: 0.148 ± 0.01 (0.124–0.174); EW: 0.105 ± 0.01 (0.087–0.124); WL: 0.735 ± 0.05 (0.671–0.820); PSL: 0.155 ± 0.015 (0.130–0.174); SDL: 0.111 ± 0.01 (0.096–0.124); PEL: 0.271 ± 0.03 (0.224–0.311); PPL: 0.168 ± 0.01 (0.149–0.186); PEH: 0.208 ± 0.015 (0.186–0.236); PPH: 0.207 ± 0.015 (0.180–0.224); PNW: 0.368 ± 0.02 (0.329–0.401); PLW: 0.170 ± 0.01 (0.149–0.199); PPW: 0.223 ± 0.02 (0.196–0.242); CI: 85.9 ± 1.2 (83.1–87.5); SI1: 68.6 ± 1.6 (66.7–71.3); SI2: 79.9 ± 1.6 (77.1–81.9); MI: 50.0 ± 1.0 (48.4–51.3); EI1: 71.0 ± 5.0 (64.6–78.3); EI2: 16.5 ± 1.4 (14.8–18.75); PI: 130.0 ± 7.9 (120.0–145.5); PPI: 81.4 ± 6.1 (128.2–150.0); PSI: 135.8 ± 12.4 (120.0–155.6).

Colour

Head, antennae, mesosoma, petiole, postpetiole and legs uniformly yellow to dark yellow, in few specimens, frons and femora partly darkened. Gaster yellow, only the first gaster tergite with wide, dark band on its posterior part (Figs 1, 2). Head. Rectangular, 1.16 times as long as wide, lateral surfaces below and above eyes gently convex, posterior edges convex, occipital margin of head straight or slightly concave (Figs 9, 13). Anterior margin of clypeus slightly convex, medial notch absent. Eyes moderate, oval, 1.4 times as long as wide. Antennal scape short, in lateral view slightly curved, 0.69 times as long as length of the head, in apex gradually widened, its base with small, triangular tooth, funiculus long, club 3-segmented (Fig. 9). Surface of scape with very fine microreticulation, shiny, covered with thin, moderate dense, decumbent to suberect setae. Mandibles rounded with thick sparse, longitudinal striae, shiny. Clypeus shiny with thick, sparse, longitudinal striae, area between striae smooth and shiny. Frontal carinae short, not extending beyond frontal lobes. Antennal fossa deep, with sparse, thin, roundly curved striae and dense reticulation. Frontal lobes narrow, smooth with slight, dense longitudinal striation (Fig. 13). Frons, vertex and temples with dense, thick, reticulation, sometimes central surface of frons with additional a few thin, longitudinal, interrupted wrinkles, surface between striation smooth; malar area with irregular, thick, reticulation, space between reticulation smooth or with very sparse microreticulation, shiny; genae with sparser, thank of frons, and thick reticulation, shiny. Frons and vertex with erect, pale, short and thick setae (Fig. 13). Mesosoma. Elongate, approximately twice as long as wide, slightly arched in profile. Metanotal groove absent. Pronotum convex on sides. Propodeal spines short to moderate, triangular, directed upward, base wide, tips sharp (Fig. 2). Whole surface with dense, reticulation, its dorsal surface and lateral surfaces of pronotum with thick and sparse longitudinal striation or reticulation. Area between thick sculpture shiny, smooth or sometimes with sparse, fait microreticulation. Entire mesosoma bearing erect, pale, short and thick setae (Fig. 2). Petiole. In lateral view, with short peduncle, node low, with anterior face flat, and posterior face convex and dorsum convex. Peduncle and petiolar node shiny, with thick, dense reticulation, area between rugae smooth, dorsum with sparser reticulation. Dorsal surface bearing sparse, short, erect setae (Fig. 2). Postpetiole. In lateral view, regularly convex, apical half with gently convex sides (Figs 1, 2), on the whole surface shiny, with thick, dense reticulation, dorsum with sparser reticulation; area between rugae smooth. Dorsal surface bearing sparse, short, semierect to erect setae (Fig. 2). Gaster. Gaster smooth and shiny, bearing erect, thin, pale setae (Figs 1, 2).

Figures 1, 2. 

Worker of Temnothorax brackoi sp. nov. 1 Dorsal 2 Lateral.

Etymology

Named after Gregor Bračko, a Slovenian myrmecologist, for his significant contributions to the studies on ants of the Balkan Peninsula.

General distribution

Greece: Epirus, Ionian Islands, Peloponnese, western Sterea Ellas, Thessaly; Croatia: Dalmatia.

Biology

Specimens of T. brackoi were collected from various localities: sunny and shadowy located on lowlands and highlands (9 - 990 m a.s.l). The species was noted in various habitats, most often on shrubs and trees growing along roadsides and olive plantations, maquis, phrygana, often inside forests (mostly deciduous, occasionally coniferous). We noted also its presence on the leaves of climbing plants growing on walls or trees. Nests were not found, probably like other species of this group, are located inside dry stems of herbs.

The following ant species were recorded in the same areas as T. brackoi:

Epirus, Preveza, 700 m E of Kanali: Aphaenogaster balcanica (Emery), A. subterranea (Latreille), Camponotus dalmaticus (Nylander), C. fallax (Nylander), C. lateralis (Olivier), C. piceus (Leach), C. vagus (Scopoli), Cataglyphis nodus (Brullé), Colobopsis truncata (Spinola), Crematogaster schmidti (Mayr), Dolichoderus quadripunctatus (Linnaeus), Lasius lasioides (Emery), Messor wasmanni Krausse, Myrmecina graminicola (Latreille), Pheidole pallidula (Nylander), Plagiolepis pygmaea (Latreille), Temnothorax bulgaricus (Forel), T. rogeri Emery; near Kanali, oak forest: Aphaenogaster balcanica (Emery), A. subterranea (Latreille), Camponotus dalmaticus (Nylander), C. fallax (Nylander), C. lateralis (Olivier), Cataglyphis nodus (Brullé), Colobopsis truncata (Spinola), Crematogaster schmidti (Mayr), Lasius lasioides (Emery), Pheidole pallidula (Nylander); near Kanali, pine forest: Temnothorax bulgaricus (Forel).

Ionian Islands, Cephalonia, Avithos Lake: Aphaenogaster muelleriana Wolf, Camponotus dalmaticus (Nylander), C. gestroi Emery, C. ionius Emery, C. lateralis (Olivier), Crematogaster schmidti (Mayr), C. sordidula (Nylander), Lepisiota frauenfeldi (Mayr), Liometopum microcephalum (Panzer), Messor wasmanni Krausse, Monomorium monomorium Bolton, Myrmecina graminicola (Latreille), Pheidole pallidula (Nylander), Plagiolepis pygmaea (Latreille), Tapinoma festae Emery, Temnothorax bulgaricus (Forel), T. exilis (Emery), T. laconicus Csősz et al., Tetramorium kephalosi Salata & Borowiec; Katapodata: Camponotus dalmaticus (Nylander), Crematogaster schmidti (Mayr), Lepisiota frauenfeldi (Mayr), Plagiolepis pygmaea (Latreille); 1.5 km NE of Koulourata: Aphaenogaster balcanica (Emery), Camponotus dalmaticus (Nylander), C. kiesenwetteri (Roger), C. lateralis (Olivier), Crematogaster schmidti (Mayr), Lepisiota frauenfeldi (Mayr), Pheidole balcanica Seifert, Plagiolepis pygmaea (Latreille), Temnothorax bulgaricus (Forel), T. leviceps (Emery). T. messiniaensis sp. nov.; ancient Same: Camponotus kiesenwetteri (Roger), C. lateralis (Olivier), Colobopsis truncata (Spinola), Crematogaster ionia Forel, C. sordidula (Nylander), Pheidole balcanica Seifert, Temnothorax bulgaricus (Forel), T. messiniaensis sp. nov.; road Skala-Poros: Camponotus aethiops (Latreille), C. gestroi Emery; C. dalmaticus (Nylander), C. kiesenwetteri (Roger), C. lateralis (Olivier), Colobopsis truncata (Spinola), Crematogaster schmidti (Mayr), C. sordidula (Nylander), Temnothorax messiniaensis sp. nov.

Ionian Islands, Corfu, Ag. Ilias: Aphaenogaster subterranea (Latreille), Camponotus dalmaticus (Nylander), C. lateralis (Olivier), Temnothorax laconicus Csősz et al.; Akr. Kefali near Agios Stefanos: Aphaenogaster balcanica (Emery), Bothriomyrmex communista Santschi, Camponotus aethiops (Latreille), C. dalmaticus (Nylander), C. lateralis (Olivier), C. piceus (Leach), Crematogaster schmidti (Mayr), C. sordidula (Nylander), Lasius illyricus Zimmermann, Lepisiota melas (Emery), L. nigra (Dalla Torre), Messor wasmanni Krausse, Pheidole pallidula (Nylander), Plagiolepis pygmaea (Latreille), Prenolepis nitens (Mayr), Temnothorax laconicus Csősz et al.; E of Nymfes: Aphaenogaster muelleriana Wolf, A. subterranea (Latreille), Camponotus aethiops (Latreille), C. dalmaticus (Nylander), C. lateralis (Olivier), Crematogaster sordidula (Nylander), Dolichoderus quadripunctatus (Linnaeus), Formica gagates Latreille, Lasius bombycina Seifert & Galkowski, L. illyricus Zimmermann, Lepisiota frauenfeldi (Mayr), Temnothorax laconicus Csősz et al.; Klimatia: Aphaenogaster balcanica (Emery), A. muelleriana Wolf, A. subterranea (Latreille), Camponotus aethiops (Latreille), C. dalmaticus (Nylander), C. gestroi Emery, C. lateralis (Olivier), C. heidrunvogtae Seifert, C. piceus (Leach), Cataglyphis nodus (Brullé), Colobopsis truncata (Spinola), Crematogaster schmidti (Mayr), C. sordidula (Nylander), Dolichoderus quadripunctatus (Linnaeus), Formica gagates Latreille, Lasius bombycina Seifert & Galkowski, L. illyricus Zimmermann, L. lasioides (Emery), Lepisiota frauenfeldi (Mayr), L. nigra (Dalla Torre), Messor ibericus Santschi, M. wasmanni Krausse, Myrmecina graminicola (Latreille), Pheidole pallidula (Nylander), Plagiolepis pygmaea (Latreille), P. xene Stärcke, Prenolepis nitens (Mayr), Temnothorax cf. exilis, T. laconicus Csősz et al., T. rogeri Emery, Tetramorium kephalosi Salata & Borowiec.; N of Ag. Stefanos: Aphaenogaster balcanica (Emery), Camponotus aethiops (Latreille), C. fallax (Nylander), C. lateralis (Olivier), C. piceus (Leach), Crematogaster schmidti (Mayr), Lasius bombycina Seifert & Galkowski, L. illyricus Zimmermann, Lepisiota frauenfeldi (Mayr), Messor ibericus Santschi, Pheidole pallidula (Nylander), Plagiolepis pygmaea (Latreille), Solenopsis cf. fugax, Temnothorax clypeatus (Mayr); near Doukades: Camponotus lateralis (Olivier), C. piceus (Leach), Lasius lasioides (Emery); near Vistonas: Camponotus aethiops (Latreille), C. dalmaticus (Nylander), C. piceus (Leach), Crematogaster sordidula (Nylander), Lepisiota melas (Emery), Messor ibericus Santschi, M. wasmanni Krausse, Plagiolepis pygmaea (Latreille), Temnothorax cf. exilis, T. rogeri Emery, Tetramorium kephalosi Salata & Borowiec; Nymfes: Aphaenogaster cf. subterranea, Temnothorax laconicus Csősz et al., T. cf. tuberum, Tetramorium cf. caespitum; Corfu, Old Perithia: Aphaenogaster balcanica (Emery), A. epirotes (Emery), A. muelleriana Wolf, Bothriomyrmex communista Santschi, Camponotus aethiops (Latreille), C. dalmaticus (Nylander), C. gestroi Emery, C. lateralis (Olivier), C. piceus (Leach), Crematogaster schmidti (Mayr), Lasius illyricus Zimmermann, L. lasioides (Emery), Lepisiota frauenfeldi (Mayr), L. melas (Emery), Messor ibericus Santschi, M. wasmanni Krausse, Plagiolepis pygmaea (Latreille), Prenolepis nitens (Mayr), Tapinoma erraticum (Latreille), Temnothorax affinis (Mayr), T. cf. exilis, T. laconicus Csősz et al., T. cf. nigriceps, T. rogeri Emery, Tetramorium cf. caespitum, T. kephalosi Salata & Borowiec; Strinilas: Aphaenogaster balcanica (Emery), A. epirotes (Emery), Bothriomyrmex communista Santschi, Camponotus dalmaticus (Nylander), C. lateralis (Olivier), Formica gagates Latreille, Lasius lasioides (Emery), Plagiolepis perperamus Salata et al., P. pygmaea (Latreille), Ponera coarctata (Latreille), Solenopsis cf. fugax, Tapinoma erraticum (Latreille), T. laconicus Csősz et al., Tetramorium kephalosi Salata & Borowiec, T. cf. punctatum;

Ionian Islands, Lefkada, Asprogerakata: Aphaenogaster balcanica (Emery), A. muelleriana Wolf, Camponotus aethiops (Latreille), C. dalmaticus (Nylander), Crematogaster schmidti (Mayr), Lepisiota frauenfeldi (Mayr), Pheidole cf. pallidula, Prenolepis nitens (Mayr), Tetramorium kephalosi Salata & Borowiec; Egklouvi: Aphaenogaster balcanica (Emery), Camponotus dalmaticus (Nylander), Cataglyphis nodus (Brullé), Crematogaster schmidti (Mayr), Messor wasmanni Krausse, Pheidole cf. pallidula, Temnothorax lichtensteini (Bondroit), T. rogeri Emery;

Ionian Islands, Zakynthos, 1 km E of Elies: Camponotus aethiops (Latreille), C. dalmaticus (Nylander), C. kiesenwetteri (Roger), C. lateralis (Olivier), C. oertzeni Forel, Crematogaster schmidti (Mayr), Messor wasmanni Krausse, Plagiolepis pygmaea (Latreille), Temnothorax cf. tergestinus; 1 km N of Exo Chora: Aphaenogaster balcanica (Emery), Camponotus dalmaticus (Nylander), C. kiesenwetteri (Roger), C. oertzeni Forel, Crematogaster schmidti (Mayr), C. sordidula (Nylander), Lepisiota frauenfeldi (Mayr), Pheidole cf. pallidula, Plagiolepis pygmaea (Latreille), Temnothorax bulgaricus (Forel), T. rogeri Emery, T. cf. tergestinus; 1.2 km N of Ano Vasilikos: Temnothorax exilis (Emery); 1.2 km N of Vasilikos: Aphaenogaster balcanica (Emery), Camponotus aethiops (Latreille), C. dalmaticus (Nylander), C. kiesenwetteri (Roger), C. lateralis (Olivier), Colobopsis truncata (Spinola), Crematogaster schmidti (Mayr), C. sordidula (Nylander), Lepisiota frauenfeldi (Mayr), Plagiolepis pygmaea (Latreille), Temnothorax exilis (Emery), T. graecus (Forel), T. rogeri Emery, T. cf. tergestinus; 1.2 km NE of Anafonitria: Aphaenogaster balcanica (Emery), A. cf. epirotes, Camponotus aethiops (Latreille), C. dalmaticus (Nylander), C. kiesenwetteri (Roger), Crematogaster schmidti (Mayr), Lepisiota frauenfeldi (Mayr), Pheidole cf. pallidula, Plagiolepis pygmaea (Latreille), Temnothorax exilis (Emery), T. muellerianus (Finzi), T. cf. tergestinus; 1.2 km SW of Ano Vasilikos loc. 1: Aphaenogaster balcanica (Emery), Camponotus dalmaticus (Nylander), C. kiesenwetteri (Roger), C. lateralis (Olivier), C. oertzeni Forel, Crematogaster schmidti (Mayr), C. sordidula (Nylander), Lepisiota frauenfeldi (Mayr), Plagiolepis pygmaea (Latreille), Temnothorax exilis (Emery); 1.2 km SW of Ano Vasilikos loc. 2: Aphaenogaster balcanica (Emery), Aphaenogaster cf. epirotes, Camponotus dalmaticus (Nylander), C. lateralis (Olivier), C. oertzeni Forel, Cataglyphis nodus (Brullé), Crematogaster schmidti (Mayr), C. sordidula (Nylander), Lepisiota melas (Emery), Messor wasmanni Krausse, Pheidole cf. pallidula, Plagiolepis pygmaea (Latreille), Temnothorax exilis (Emery); 1.2 km SW of Skinaria: Aphaenogaster balcanica (Emery), Camponotus aethiops (Latreille), C. gestroi Emery, C. ionius Emery, C. kiesenwetteri (Roger), C. oertzeni Forel, Crematogaster sordidula (Nylander), Lepisiota melas (Emery), Messor wasmanni Krausse, Pheidole cf. pallidula, Plagiolepis pygmaea (Latreille), Temnothorax exilis (Emery), T. rogeri Emery, T. cf. tergestinus, Tetramorium diomedeum Emery, T. kephalosi Salata & Borowiec; 1.4 km E of Ano Volimes: Aphaenogaster balcanica (Emery), A. cf. epirotes, A. muelleriana Wolf, Camponotus dalmaticus (Nylander), Crematogaster schmidti (Mayr), Lepisiota frauenfeldi (Mayr), Messor wasmanni Krausse, Pheidole cf. pallidula, Plagiolepis pygmaea (Latreille), Prenolepis nitens (Mayr), Temnothorax bulgaricus (Forel), T. cf. tergestinus, Tetramorium kephalosi Salata & Borowiec; 1.5 km N of Exo Chora: Aphaenogaster cf. epirotes, Camponotus dalmaticus (Nylander), C. kiesenwetteri (Roger), C. lateralis (Olivier), C. oertzeni Forel, Crematogaster schmidti (Mayr), Lepisiota frauenfeldi (Mayr), Messor wasmanni Krausse, Plagiolepis pygmaea (Latreille), Temnothorax bulgaricus (Forel), T. cf. tergestinus, Tetramorium kephalosi Salata & Borowiec; 1.7 km NE of Ano Volimes: Camponotus aethiops (Latreille), C. dalmaticus (Nylander), C. gestroi Emery, C. kiesenwetteri (Roger), C. lateralis (Olivier), Crematogaster schmidti (Mayr), C. sordidula (Nylander), Pheidole cf. pallidula, Temnothorax cf. tergestinus; 1.8 km SW of Volimes: Aphaenogaster balcanica (Emery), Camponotus aethiops (Latreille), C. dalmaticus (Nylander), C. kiesenwetteri (Roger), Crematogaster schmidti (Mayr), C. sordidula (Nylander), Plagiolepis pygmaea (Latreille), Temnothorax cf. tergestinus; 1.9 km W of Maries: Aphaenogaster balcanica (Emery), A. cf. epirotes, Camponotus aethiops (Latreille), C. dalmaticus (Nylander), C. kiesenwetteri (Roger), Crematogaster sordidula (Nylander), Lepisiota melas (Emery), Messor wasmanni Krausse, Pheidole cf. pallidula, Plagiolepis pygmaea (Latreille), Temnothorax cf. tergestinus; 2.5 km NE of Maries: Aphaenogaster balcanica (Emery), A. cf. epirotes, A. subterraneoides Emery, Bothriomyrmex communista Santschi, Camponotus aethiops (Latreille), C. dalmaticus (Nylander), C. kiesenwetteri (Roger), Crematogaster schmidti (Mayr), C. sordidula (Nylander), Plagiolepis pygmaea (Latreille), Temnothorax rogeri Emery, Tetramorium kephalosi Salata & Borowiec; 3.9 km NE of Maries: Camponotus aethiops (Latreille), C. dalmaticus (Nylander), Crematogaster schmidti (Mayr), Prenolepis nitens (Mayr), Temnothorax cf. tergestinus, Tetramorium diomedeum Mayr; 470 m NE of Orthonies: Aphaenogaster balcanica (Emery), Camponotus aethiops (Latreille), C. dalmaticus (Nylander), C. kiesenwetteri (Roger), Crematogaster schmidti (Mayr), C. sordidula (Nylander), Lepisiota melas (Emery), Plagiolepis pygmaea (Latreille), Temnothorax bulgaricus (Forel), T. exilis (Emery), Temnothorax cf. tergestinus; 500 m S of Apelati: Aphaenogaster balcanica (Emery), Camponotus dalmaticus (Nylander), C. lateralis (Olivier), Crematogaster ionia Forel, C. schmidti (Mayr), Messor wasmanni Krausse, Myrmecina graminicola (Latreille), Plagiolepis pygmaea (Latreille), Temnothorax cf. tergestinus; 700 m SW of Koroni: Camponotus aethiops (Latreille), C. kiesenwetteri (Roger), C. lateralis (Olivier), Lepisiota melas (Emery), Pheidole cf. pallidula, Plagiolepis pygmaea (Latreille), Temnothorax exilis (Emery), Temnothorax cf. tergestinus; 750 m S of Volimes: Camponotus dalmaticus (Nylander), C. gestroi Emery, C. lateralis (Olivier), Crematogaster schmidti (Mayr), C. sordidula (Nylander), Messor wasmanni Krausse, Plagiolepis pygmaea (Latreille), Temnothorax bulgaricus (Forel), T. rogeri Emery, T. cf. tergestinus; 800 m SE of Xirokastello: Camponotus dalmaticus (Nylander), C. kiesenwetteri (Roger), C. lateralis (Olivier), C. oertzeni Forel, Colobopsis truncata (Spinola), Crematogaster schmidti (Mayr), Lepisiota melas (Emery), Messor wasmanni Krausse, Plagiolepis pygmaea (Latreille), Temnothorax exilis (Emery), T. cf. tergestinus, Tetramorium cf. punctatum; 880 m S of Orthonies: Aphaenogaster balcanica (Emery), A. cf. epirotes, Camponotus aethiops (Latreille), C. dalmaticus (Nylander), C. kiesenwetteri (Roger), C. lateralis (Olivier), Crematogaster schmidti (Mayr), Lepisiota frauenfeldi (Mayr), L. melas (Emery), Pheidole cf. pallidula, Plagiolepis pygmaea (Latreille), Temnothorax bulgaricus (Forel), T. rogeri Emery, T. cf. tergestinus; Ag. Georgiou monastery: Camponotus aethiops (Latreille), C. dalmaticus (Nylander), Crematogaster schmidti (Mayr), Plagiolepis pygmaea (Latreille); Ag. Joannis: Camponotus aethiops (Latreille), C. kiesenwetteri (Roger), C. lateralis (Olivier), Crematogaster schmidti (Mayr), C. sordidula (Nylander), Pheidole cf. pallidula, Solenopsis cf. lusitanica, Temnothorax graecus (Forel), T. rogeri Emery, T. cf. tergestinus, Tetramorium kephalosi Salata & Borowiec; Argassi: Aphaenogaster balcanica (Emery), Camponotus dalmaticus (Nylander), C. kiesenwetteri (Roger), C. oertzeni Forel, Colobopsis truncata (Spinola), Crematogaster schmidti (Mayr), C. sordidula (Nylander), Lasius alienus (Foerster), Lepisiota melas (Emery), Messor ibericus Santschi, Monomorium monomorium Bolton, Nylanderia jaegerskioeldi (Mayr), Pheidole cf. pallidula, Plagiolepis perperamus Salata et al., P. pygmaea (Latreille), Tapinoma erraticum (Latreille), Temnothorax rogeri Emery, Tetramorium immigrans Santschi, T. cf. punctatum; Livia Mts. loc. 1: Aphaenogaster balcanica (Emery), A. cf. epirotes, Bothriomyrmex communista Santschi, Camponotus aethiops (Latreille), C. kiesenwetteri (Roger), Crematogaster schmidti (Mayr), C. sordidula (Nylander), Messor wasmanni Krausse, Prenolepis nitens (Mayr), Tetramorium cf. punctatum; Vrachionas Mts.: Aphaenogaster balcanica (Emery), Bothriomyrmex communista Santschi, Camponotus aethiops (Latreille), C. dalmaticus (Nylander), C. kiesenwetteri (Roger), C. oertzeni Forel, Crematogaster schmidti (Mayr), Messor wasmanni Krausse, Plagiolepis pygmaea (Latreille), Prenolepis nitens (Mayr), Temnothorax cf. tergestinus, Tetramorium cf. punctatum; s, W of Kampi: Aphaenogaster balcanica (Emery), A. cf. epirotes, A. muelleriana Wolf, Camponotus aethiops (Latreille), Crematogaster schmidti (Mayr), Pheidole cf. pallidula, Plagiolepis pygmaea (Latreille), Solenopsis cf. lusitanica, Temnothorax rogeri Emery.

Macedonia, Pieria, Platamonas Castle hill: Aphaenogaster epirotes (Emery), A. muelleriana Wolf, A. cf. subterranea, Camponotus dalmaticus (Nylander), C. lateralis (Olivier), C. piceus (Leach), Crematogaster ionia Forel, C. sordidula (Nylander), Lasius emarginatus (Olivier), L. turcicus (Santschi), Messor hellenius Agosti & Collingwood, Pheidole balcanica Seifert, Plagiolepis pygmaea (Latreille), Prenolepis nitens (Mayr), Temnothorax lichtensteini (Bondroit); road to P. Poroi loc. 1: Camponotus aethiops (Latreille), C. dalmaticus (Nylander), C. ionius Emery, C. piceus (Leach), Chalepoxenus muellerianus (Finzi), Crematogaster ionia Forel, Dolichoderus quadripunctatus (Linnaeus), Formica gagates Latreille, Lasius emarginatus (Olivier), Messor hellenius Agosti & Collingwood, Plagiolepis pygmaea (Latreille), Tapinoma erraticum (Latreille), Temnothorax cf. exilis, T. graecus (Forel), T. turcicus (Santschi); road to P. Poroi loc. 2: Camponotus aethiops (Latreille), C. dalmaticus (Nylander), C. piceus (Leach), Cataglyphis nodus (Brullé), Crematogaster schmidti (Mayr), Dolichoderus quadripunctatus (Linnaeus); Lasius emarginatus (Olivier), Lepisiota frauenfeldi (Mayr), Pheidole balcanica Seifert, Plagiolepis pygmaea (Latreille), Prenolepis nitens (Mayr), Tapinoma erraticum (Latreille), Temnothorax graecus (Forel), T. turcicus (Santschi), Tetramorium kephalosi Salata & Borowiec; road to P. Poroi loc. 3: Camponotus dalmaticus (Nylander), C. lateralis (Olivier), C. vagus (Scopoli), Crematogaster schmidti (Mayr), Dolichoderus quadripunctatus (Linnaeus), Lasius emarginatus (Olivier), Messor hellenius Agosti & Collingwood, Pheidole balcanica Seifert, Plagiolepis pygmaea (Latreille), Prenolepis nitens (Mayr), Temnothorax bulgaricus (Forel), T. morea Csősz, Salata & Borowiec, T. turcicus (Santschi).

Peloponnese, Korinthia, near Evrostina: Aphaenogaster subterranea (Latreille), Camponotus lateralis (Olivier), Crematogaster schmidti (Mayr), Dolichoderus quadripunctatus (Linnaeus), Lasius emarginatus (Olivier), L. turcicus (Santschi), Ponera coarctata (Latreille), Temnothorax laconicus Csősz et al.

Peloponnese, Laconia, Taygetos Mts., 1.5 km SW of Anavryti: Aphaenogaster balcanica (Emery), A. subterranea (Latreille), Camponotus dalmaticus (Nylander), C. oertzeni Forel, C. piceus (Leach), Cataglyphis nodus (Brullé), Crematogaster ionia Forel, Temnothorax laconicus Csősz et al.

Peloponnese, Messinia, Taygetos Mts., 0.7 km S of Dyrrachio: Camponotus lateralis (Olivier), Crematogaster schmidti (Mayr), Lasius flavus (Fabricius), L. illyricus Zimmermann, Temnothorax helenae Csősz et al.; Taygetos Mts., 0.8 km SE of Exochori: Aphaenogaster balcanica (Emery), Camponotus dalmaticus (Nylander), C. lateralis (Olivier), Crematogaster ionia Forel, Lepisiota frauenfeldi (Mayr), Messor structor (Latreille), M. wasmanni Krausse, Pheidole pallidula (Nylander), Plagiolepis pygmaea (Latreille), Temnothorax cf. bulgaricus, T. exilis (Emery), T. helenae Csősz et al., T. recedens (Nylander), Tetramorium kephalosi Salata & Borowiec; Taygetos Mts., 1.3 km S of Artemisia: Aphaenogaster subterranea (Latreille), Camponotus boghossiani Forel, Crematogaster schmidti (Mayr), Lasius cf. alienus, L. illyricus Zimmermann, Myrmecina graminicola (Latreille), Plagiolepis pygmaea (Latreille), Temnothorax crasecundus Seifert & Csősz, T. helenae Csősz et al.; Taygetos Mts., 1.8 km E of Saidona: Lasius illyricus Zimmermann, Plagiolepis pygmaea (Latreille), Temnothorax crasecundus Seifert & Csősz, Tetramorium cf. caespitum; Taygetos Mts., 2 km W of Arachova: Aphaenogaster cf. muelleriana, Bothriomyrmex communista Santschi, Camponotus boghossiani Forel, C. dalmaticus (Nylander), C. laconicus Emery, Cataglyphis nodus (Brullé), Crematogaster schmidti (Mayr), Lepisiota frauenfeldi (Mayr), Messor wasmanni Krausse, Pheidole pallidula (Nylander), Plagiolepis pygmaea (Latreille), Stigmatomma denticulatum Roger, Tapinoma erraticum (Latreille), Temnothorax cf. bulgaricus, T. laconicus Csősz et al., T. cf. luteus, T. rogeri Emery, Tetramorium kephalosi Salata & Borowiec, Tetramorium cf. punctatum; Taygetos Mts., Arachova: Camponotus aethiops (Latreille), C. laconicus Emery, C. piceus (Leach), Cataglyphis nodus (Brullé), Crematogaster schmidti (Mayr), Dolichoderus quadripunctatus (Linnaeus), Lasius cf. alienus, L. illyricus Zimmermann, Lepisiota frauenfeldi (Mayr), Messor wasmanni Krausse, Pheidole pallidula (Nylander), Plagiolepis pygmaea (Latreille), Tapinoma erraticum (Latreille), Temnothorax cf. bulgaricus, T. parvulus (Schenck); Taygetos Mts., Chora Getson: Aphaenogaster balcanica (Emery), A. cf. muelleriana, Camponotus boghossiani Forel, Crematogaster schmidti (Mayr), Dolichoderus quadripunctatus (Linnaeus), Lasius cf. alienus, L. illyricus Zimmermann, Lepisiota frauenfeldi (Mayr), Pheidole pallidula (Nylander), Plagiolepis pygmaea (Latreille), Temnothorax cf. bulgaricus, T. laconicus Csősz et al., T. recedens (Nylander), T. strymonensis Csősz, Salata & Borowiec; Taygetos Mts., Karveli: Aphaenogaster cf. muelleriana, Crematogaster schmidti (Mayr), Lasius cf. alienus, L. illyricus Zimmermann, Myrmecina graminicola (Latreille), Temnothorax helenae Csősz et al., T. strymonensis Csősz, Salata & Borowiec; Tetrazi Mts., 0.5 km E of Vastas: Aphaenogaster balcanica (Emery), Camponotus dalmaticus (Nylander), C. fallax (Nylander), C. laconicus Csősz et al., C. piceus (Leach), Colobopsis truncata (Spinola), Crematogaster schmidti (Mayr), Formica gagates Latreille, Lasius illyricus Zimmermann, Pheidole pallidula (Nylander), Plagiolepis pygmaea (Latreille), Temnothorax helenae Csősz et al., T. laconicus Csősz et al., T. morea Csősz, Salata & Borowiec, T. rogeri Emery; Tetrazi Mts., Isaris: Aphaenogaster balcanica (Emery), A. subterranea (Latreille), Camponotus dalmaticus (Nylander), C. laconicus Csősz et al., C. lateralis (Olivier), Crematogaster schmidti (Mayr), Dolichoderus quadripunctatus (Linnaeus), Lasius cf. alienus, L. illyricus Zimmermann, Lepisiota frauenfeldi (Mayr), Temnothorax cf. bulgaricus, T. helenae Csősz et al.,; Tetrazi Mts., Karnasi: Aphaenogaster balcanica (Emery), A. cf. muelleriana, Camponotus dalmaticus (Nylander), C. lateralis (Olivier), Crematogaster schmidti (Mayr), Lasius illyricus Zimmermann, Messor hellenius Agosti & Collingwood, Pheidole pallidula (Nylander), Temnothorax cf. bulgaricus, Temnothorax laconicus Csősz et al., T. morea Csősz, Salata & Borowiec, T. rogeri Emery, Tetramorium kephalosi Salata & Borowiec.

Sterea Ellas, Aetolia-Acarnania, Psila Alonia: Aphaenogaster balcanica (Emery), A. cf. muelleriana, A. subterranea (Latreille), Camponotus dalmaticus (Nylander), C. lateralis (Olivier), Cataglyphis nodus (Brullé), Crematogaster schmidti (Mayr), Dolichoderus quadripunctatus (Linnaeus), Lasius illyricus Zimmermann, Messor wasmanni Krausse, Pheidole cf. pallidula, Plagiolepis pygmaea (Latreille), Solenopsis cf. lusitanica, Temnothorax crassispinus (Karavaiev).

Sterea Ellas, Euboea, 1.2 km NW of Gerontas: Aphaenogaster balcanica (Emery), Camponotus aethiops (Latreille), C. dalmaticus (Nylander), C. gestroi Emery, C. kiesenwetteri (Roger), C. laconicus Emery, C. lateralis (Olivier), C. piceus (Leach), Cataglyphis aenescens (Nylander), Crematogaster ionia Forel, C. schmidti (Mayr), Pheidole cf. pallidula, Plagiolepis pygmaea (Latreille), Tapinoma erraticum (Latreille), Temnothorax graecus (Forel), T. recedens (Nylander); 2.3 km S of Stropones: Camponotus aethiops (Latreille), C. piceus (Leach), C. vagus (Scopoli), Cataglyphis nodus (Brullé), Formica fusca Karavaiev, Lasius alienus Förster, L. flavus (Fabricius), Pheidole pallidula (Nylander), Temnothorax crasecundus Seifert & Csősz, T. helenae Csősz et al., T. unifasciatus (Latreille); 300 m NW of Agios: Camponotus aethiops (Latreille), C. dalmaticus (Nylander), Cataglyphis nodus (Brullé), Crematogaster ionia Forel, Dolichoderus quadripunctatus (Linnaeus), Formica fusca Karavaiev, Lasius lasioides (Emery), Pheidole cf. pallidula, Plagiolepis pygmaea (Latreille), Temnothorax lichtensteini (Bondroit), T. parvulus (Schenck), T. recedens (Nylander), T. unifasciatus (Latreille); 570 m NW of Drosia: Aphaenogaster balcanica (Emery), A. subterranea (Latreille), Camponotus dalmaticus (Nylander), C. ionius Emery, C. lateralis (Olivier), C. piceus (Leach), C. vagus (Scopoli), Cataglyphis nodus (Brullé), Crematogaster ionia Forel, C. schmidti (Mayr), Lepisiota frauenfeldi (Mayr), Messor hellenius Agosti & Collingwood, Temnothorax bulgaricus (Forel).

Thessaly, Larissa, Mt. Olympus, 5.3 km E of Olympiada: Aphaenogaster subterranea (Latreille), Camponotus aethiops (Latreille), C. lateralis (Olivier), C. piceus (Leach), Crematogaster schmidti (Mayr), Dolichoderus quadripunctatus (Linnaeus), Formica cunicularia Latreille, F. gagates Latreille, Lasius alienus Förster, L. emarginatus (Olivier), Messor hellenius Agosti & Collingwood, Pheidole pallidula (Nylander), Plagiolepis pygmaea (Latreille), Prenolepis nitens (Mayr), Solenopsis cf. lusitanica, Tapinoma erraticum (Latreille), Temnothorax recedens (Nylander), Tetramorium kephalosi Salata & Borowiec, Tetramorium moravicum Kratochvil; Mt. Ossa, 2.4 km SE of Karitsa: Aphaenogaster subterranea (Latreille), Camponotus lateralis (Olivier), Cataglyphis nodus (Brullé), Crematogaster schmidti (Mayr), Dolichoderus quadripunctatus (Linnaeus), Formica gagates Latreille, Lasius emarginatus (Olivier), Myrmoxenus gordiagini Ruzsky, Plagiolepis pygmaea (Latreille), Temnothorax crasecundus Seifert & Csősz, T. cf. subtilis-helenae, T. tauricus (Ruzsky), T. cf. unifasciatus, Tetramorium moravicum Kratochvil; Mt. Ossa, 600 m SE of Karitsa: Aphaenogaster subterranea (Latreille), Camponotus fallax (Nylander), Colobopsis truncata (Spinola), Lasius emarginatus (Olivier), Plagiolepis pygmaea (Latreille), Prenolepis nitens (Mayr), Temnothorax crasecundus Seifert & Csősz, T. lichtensteini (Bondroit), T. cf. subtilis-helenae, T. cf. unifasciatus; Mt. Ossa, Kokkino Nero: Aphaenogaster subterranea (Latreille), Camponotus aethiops (Latreille), C. dalmaticus (Nylander), C. lateralis (Olivier), Colobopsis truncata (Spinola), Crematogaster schmidti (Mayr), Dolichoderus quadripunctatus (Linnaeus), Formica cunicularia Latreille, Lasius alienus Förster, L. emarginatus (Olivier), Messor hellenius Agosti & Collingwood, Pheidole pallidula (Nylander), Prenolepis nitens (Mayr), Temnothorax tauricus (Ruzsky), T. cf. unifasciatus, Tetramorium cf. caespitum.

Temnothorax messiniaensis sp. nov.

Figs 3–4, 5–6, 10, 12, 14

Type material

Holotype, worker (pin) (CASENT0846796): GREECE, Pel., Messinia | 2 km E of Kalamata, 65 m, | 37.01863N / 22.15626E | 12 VI 2016, L. Borowiec || Collection L. Borowiec | Formicidae | LBC-GR01997 (MNHW).

Paratypes: • 3Q., 7w. (pin) (CASENT0846639-CASENT0846648): the same nest sample as holotype (DBET, BMNH, CASC, MHNG); • 1w. (pin) (CASENT0846649): GREECE, Pel., Messinia | 1.4 km S of Flesiada, 700 m | 37.08964N, 21.76581E | 16 VI 2016, L. Borowiec || Collection L. Borowiec | Formicidae | LBC-GR02091 (DBET); • 1w. (pin) (CASENT0846650): GREECE, Pel., Messinia | Kalamata, old centre, 60 m | 37.04617N, 22.11691E | 11 VI 2016, L. Borowiec || Collection L. Borowiec | Formicidae | LBC-GR02216 (DBET); • 2w. (pin) (CASENT0846651-CASENT0846652): GREECE, Pel., Messinia | Kalamata, railway park, 8m | 37.03157N, 22.11004E | 11 VI 2016, L. Borowiec || Collection L. Borowiec | Formicidae | LBC-GR01989 (DBET); • 2w. (pin) (CASENT0846653-CASENT0846654): GREECE, Pel., Messinia | 0.8 km N of Koromilea, 485 m | 37.16272N, 21.84809E | 16 VI 2016, L. Borowiec || Collection L. Borowiec | Formicidae | LBC-GR02103 (DBET); • 1w. (pin) (CASENT0846655): GREECE, Pel., Messinia | 0.8 km SE of Exochori, 535 m | 36.89582N, 22.27464E | 20 VI 2016, L. Borowiec || Collection L. Borowiec | Formicidae | LBC-GR02658 (DBET); • 1w. (pin) (CASENT0846656): GREECE, Pel., Messinia | 0.8 km W of Eleochori, 481 m 37.03838N, 22.17227E | 13 VI 2016, lL. Borowiec || Collection L. Borowiec | Formicidae | LBC-GR02005 (DBET).

Other material

Greece. Ionian Islands, Cephalonia: • 7w. (EtOH): Avithos Lake, shrubs around small lake, 38.17203N/ 20.71107E, 288 m, 2019-06-10, leg. L. Borowiec; • 3w. (pin), 9w. (EtOH): 1.6 km SW of Digaleto, small gorge with oaks, 38.16558N, 20.67099E, 564 m, 2019-06-11, leg. L. Borowiec; • 1w. (EtOH): 1.8 km SW of Digaleto, pastures with oaks, 38.16593N, 20.66788E, 580 m, 2019-06-11, leg. L. Borowiec; • 7w. (EtOH): Kapandriti vicinity, roadsides with herbs, 38.12913N/20.72447E, 320 m, 2019-06-09, leg. L. Borowiec; • 12w. (EtOH): Katapodata; roadsides with shrubs, 38.23337N/20.64594E, 100 m, 2019-06-10, leg. L. Borowiec; • 1w. (pin) (CASENT0846797), 43w. (EtOH): 800 m S of Kateleios, roadsides with bushes, 38.07066N/20.75329E, 20 m, 2019-06-09, leg. L. Borowiec; • 5w. (EtOH): 1.5 km NE of Koulourata, mixed forest on shrubs, 38.20667N/ 20.67715E, 273 m, 2019-06-10, leg. L. Borowiec; • 48w. (EtOH): Kremmidi, roadsides with bushes, 38.09048N/20,74471E, 285 m, 2019-06-09, leg. L. Borowiec; • 7w. (EtOH): Moni Aprilion, hill with oak forest and rocks, 38.26221N/20.66651E, 220 m, 2019-06-10, leg. L. Borowiec; • 48w. (EtOH): 1 km NW of Pastra, roadsides with bushes, 38.10058N/20.7421E, 300 m, 2019-06-09, leg. L. Borowiec; • 4w. (EtOH): 1.7 km NW of Pastra, pastures, on shrubs, 38.1084N/20.74085E, 300 m, 2019-06-09, leg. L. Borowiec; • 6w. (pin) (CASENT0846798-CASENT0846803): n. Peratata, pine forest on a rocky hill, 38.14058N/20.55038E, 211 m, 2014-06-24, leg. L. Borowiec; • 11w. (EtOH): ancient Same; roadsides with shrubs, 38.2522N/20.66423E, 220 m, 2019-06-10, leg. L. Borowiec; • 3w. (pin) (CASENT0846804-CASENT0846806), 6w. (EtOH): Skala vicinity loc. 1, small gorge with mediterranean shrubs, 38.08178N/20.79275E, 40 m, 2019-06-07, leg. L. Borowiec; • 5w. (pin) (CASENT0846807-CASENT0846811), 44w, 1q (EtOH): Skala vicinity loc. 2, small gorge with mediterranean shrubs, nest inside dry branch of shrub, 38.08221N, 20.79504E, 34 m, 2019-06-07, leg. L. Borowiec; • 61w. (EtOH): rd. Skala-Poros; mediterranean shrubs, 38.12872N/20.79576E, 5 m, 2019-06-12, leg. L. Borowiec. Ionian Islands, Zakynthos: • 3w. (pin) (CASENT0846812-CASENT0846814), 3w. (EtOH): 1 km N of Exo Chora, mixed forest, 37.81063N/20.68459E, 430 m, 2018-05-08, leg. L. Borowiec; • 1w. (pin) (CASENT0846815): 1.2 km N of Vasilikos, roadsides along olive plantation and pasture with oak shrubs, 37.72456N/20.97786E, 30 m, 2018-05-05, leg. L. Borowiec; • 5w. (pin) (CASENT0846816-CASENT0846820), 1w. (EtOH): 1.2 km NE of Anafonitria, shrubs around burned forests, 37.85489N/20.64124E, 475 m, 2018-05-10, leg. L. Borowiec; • 1w. (pin) (CASENT0846821): 1.2 km SE of Loucha, roadsides in cypress forest, 37.78617N/ 20.73706E, 445 m, 2018-05-09, leg. L. Borowiec; • 5w. (pin), 3w. (EtOH): 1.2 km SW of Skinaria, limestone hills after burned forests, 37.87694N/20.69272E, 375 m, 2018-05-06, leg. L. Borowiec; • 2w. (pin) (CASENT0846822-CASENT0846823): 1.4 km S of Lithakia, shrubs along olive plantation, 37.70641N/20.82342E, 225 m, 2018-05-07, leg. L. Borowiec; • 2w. (pin) (CASENT0846824-CASENT0846825): 1.8 km SW of Volimes, shrubs along roadsides, 37.86472N/20.64234E, 350 m, 2018-05-10, leg. L. Borowiec; • 3w. (pin) (CASENT0846826-CASENT0846827), 40w. (EtOH): 1.9 km W of Maries, roadsides in burned forests, 37.818N/20.65556E, 290 m, 2018-05-09, leg. L. Borowiec; • 3w. (pin) (CASENT0846828-CASENT0846830), 19w. (EtOH): 330 m S of Stimies, shrubs around olive plantation, 37.69009N/20.79988E, 245 m, 2018-05-07, leg. L. Borowiec; • 1w. (pin) (CASENT0846831): 470 m NE of Orthonies, shrubs along roadsides, 37.85435N/ 20.69843E, 405 m, 2018-05-10, leg. L. Borowiec; 1w. (pin) (CASENT0846832): 580 m SW of Lithakia, shrubs along roadsides, 37.71491N/20.8242E, 225 m, 2018-05-07, leg. L. Borowiec; • 3w. (pin) (CASENT0846833-CASENT0846835), 4w. (EtOH): 600 m E of Ag. Leon, shrubs in pine forest, 37.77045N/20.72959E, 600 m, 2018-05-09, leg. L. Borowiec; • 2w. (pin) (CASENT0846836-CASENT0846837), 3w. (EtOH): 700 m SW of Koroni, maquis, 37.86582N/20.71753E, 290 m, 2018-05-10, leg. L. Borowiec; • 2w. (pin) (CASENT0846838-CASENT0846839): 800 m SE of Xirokastello, roadsides along olive plantation, 37.73491N/20.95139E, 75 m, 2018-05-05, leg. L. Borowiec; • 3w. (pin) (CASENT0846840-CASENT0846842), 9w. (EtOH): 880 m S of Orthonies, shrubs in cypress forest, 37.84462N/ 20.69843E, 390 m, 2018-05-10, leg. L. Borowiec; • 3w. (pin) (CASENT0846843-CASENT0846845), 2w. (EtOH): Ag. Georgiou monastery, shrubs along roadsides, 37.85971N/20.63646E, 330 m, 2018-05-10, leg. L. Borowiec; • 2w. (pin) (CASENT0846846-CASENT0846847): Ag. Joannis, roadsides with shrubs, 37.72924N/20.94553E, 165 m, 2018-05-05, leg. L. Borowiec; • 1w. (pin) (CASENT0846848): Vrachionas Mts., mountain pastures with shrubs, 37.81798N/20.70621E, 670 m, 2018-05-08, leg. L. Borowiec; Peloponnese, Achaia: • 1w (pin) (CASENT0846849): Kalavrita, 710 m, 38.03342N, 22.10456E, leg. C. Lebas; Peloponnese, Lakonia: • 1w (pin) (CASENT0846850): Mistra vicinity, 378 m, 37.08115N, 22.36545E, leg. C. Lebas.

Terra typica

Greece, Peloponnese, Messinia.

Differential diagnosis

Both T. messiniaensis sp. nov. and T. turcicus (Santschi) are characterised by very long propodeal spines, character strongly distinguishing them from T. brackoi sp. nov. They differ from species of the T. interruptus group in lack of wide frontal lobes and not distinctly triangular propodeal spines; from T. affinis they differ in brighter body colouration and shape of propodeal spines (T. affinis has propodeal spines thin and never triangular, while T. messiniaensis and T. turcicus have propodeal spines more triangular, with wider base); from T. kemali both new species differ in absence of distinctly arched dorsum of petiole node and more triangular shape of propodeal spines, additionally T. messiniaensis differs from T. kemali in not reduced sculpture on frons centre and T. turcicus differs from T. kemali in very thin dark band on posterior part of first gastral tergite. T. messiniaensis differs from T. turcicus in not reduced sculpture on frons centre and wider dark band on posterior part of first gastral tergite. From specimens of T. aveli with long propodeal spines and almost complete microreticulation of head T. messiniaensis differs in less convex mesosoma, longer head, usually darkened gena and propodeal spines directed more upwards.

Description of worker (n = 10): HL: 0.619 ± 0.04 (0.565–0.682); HW: 0.515 ± 0.03 (0.471–0.564); SL: 0.428 ± 0.02 (0.388–0.459); EL: 0.134 ± 0.008 (0.129–0.153); EW: 0.102 ± 0.01 (0.094–0.118); WL: 0.723 ± 0.05 (0.624–0.765); PSL: 0.171 ± 0.02 (0.135–0.188); SDL: 0.109 ± 0.02 (0.082–0.159); PEL: 0.263 ± 0.02 (0.224–0.282); PPL: 0.167 ± 0.007 (0.152–0.176); PEH: 0.207 ± 0.009 (0.188–0.281); PPH: 0.195 ± 0.01 (0.176–0.212); PNW: 0.359 ± 0.03 (0.318–0.400); PLW: 0.162 ± 0.01 (0.141–0.176); PPW: 0.208 ± 0.01 (0.188–0.223); CI: 83.4 ± 2.5 (78.9–86.6); SI1: 69.1 ± 2.3 (65.5–72.5); SI2: 82.9 ± 2.4 (78.6–87.6); MI: 49.7 ± 1.6 (47.6–52.3); EI1: 76.4 ± 6.8 (66.7–90.9); EI2: 16.5 ± 1.1 (14.5–18.2); PI: 127.2 ± 4.5 (118.0–133.3); PPI: 85.9 ± 4.1 (77.7–93.3); PSI: 170.8 ± 16.8 (153.8–200.0).

Colour

Head, antennae, mesosoma, petiole, postpetiole and legs uniformly yellow to ochre, often posterior part of frons, gena and femora partly darkened, occasionally also antennal club slightly darkened. Gaster yellow, only the first gaster tergite with wide, dark band on its posterior part (Figs 3, 4). Head. Rectangular, but slightly longer than in both Greek congeners, 1.2 times as long as wide, lateral surfaces below and above eyes gently convex, posterior edges convex, occipital margin of head straight or slightly concave (Figs 10, 14). Anterior margin of the clypeus slightly convex, medial notch absent. Eyes moderate, oval, 1.31 times as long as wide. Antennal scape short, in lateral view slightly curved, 0.69 times as long as length of the head, in apex gradually widened, its base with small, triangular tooth, funiculus long, club 3-segmented (Fig. 10). Surface of scape with very fine microreticulation, shiny, covered with thin, moderate dense, decumbent setae. Mandibles rounded with thick sparse, longitudinal striae, shiny. Clypeus shiny with thick, sparse, longitudinal striae, area between striae smooth and shiny. Frontal carinae short, not extending beyond frontal lobes. Antennal fossa deep, with irregular, dense rugosity and sometimes with a few thin, roundly curved striae. Frontal lobes narrow, smooth with slight, dense longitudinal striation (Fig. 14). Frons, vertex and temples with dense, thick, reticulation, central surface of frons with longitudinal reticulation and sometimes with additional thin, longitudinal striation, striae sometimes interrupted, surface between striation smooth; malar area with irregular, thick, reticulation, space between reticulation smooth or with very sparse microreticulation, shiny; genae with sparser, than on frons, and thick reticulation, shiny. Frons and vertex with erect, pale, short and thick setae (Fig. 14). Mesosoma. Elongate, approximately twice as long as wide, slightly arched in profile. Metanotal groove absent or slightly marked as in the species from the Temnothorax angustulus group. Pronotum convex on sides. Propodeal spines long, directed upward, with base slightly to moderate wider than apex, tips sharp (Fig. 4). Whole surface with dense, reticulation, sometimes its dorsal surface and lateral surfaces of pronotum and mesonotum with additional thick, sparse to moderate dense longitudinal striation or longitudinal reticulation. Area between thick sculpture shiny, smooth or sometimes with sparse, fine microreticulation (Fig. 4). Entire mesosoma bearing erect, pale, short and thick setae (Figs 3, 4). Petiole. In lateral view, with short peduncle, node moderate high, with anterior face flat, and posterior face convex and dorsum flat or slightly convex. Peduncle and petiolar node shiny, with thick, dense reticulation and sometimes thick, sparse longitudinal wrinkles, area between rugae smooth, dorsum with sparser reticulation. Dorsal surface bearing sparse, short, erect setae (Fig. 4). Postpetiole. In lateral view, regularly convex, apical half with gently convex sides (Figs 3, 4), on the whole surface shiny, with thick, dense reticulation, dorsum with sparser reticulation; area between rugae smooth. Dorsal surface bearing sparse, short, erect setae (Figs 3, 4). Gaster. Gaster smooth and shiny, bearing erect, thin, pale setae (Figs 3, 4).

Figures 3, 4. 

Worker of Temnothorax messiniaensis sp. nov. 3 Dorsal 4 Lateral.

Figures 5, 6. 

Gyne of Temnothorax messiniaensis sp. nov. 5 Dorsal 6 Lateral.

Description of gyne (n = 3): Colour. Head in frontal part yellowish brown to brown, gena and temples yellowish, border between dark and pale parts of head diffused. Antennae uniformly yellow. Pronotum yellow, scutum yellow laterally and yellowish brown anteriorly, without distinct borders between darker and paler parts. Mesosterna brown, propodeum yellowish dorsally and gradually darker ventrally, petiole and postpetiole brown dorsally and yellow ventrally. Legs mostly yellow, mid- and hind femora largely brown centrally. Gastral tergites mostly brown, first tergite with large yellowish spot basally, all tergites with yellowish to yellowish brown posterior margin (Figs 5, 6). Head. Eyes big, oval [EL / HW: 0.30]. Antennal scape short [SL / HW: 42], not reaching occipital margin of head. Clypeus shiny with distinct, longitudinally carinulae, interstices smooth. Antennal fossa deep, with concentric carinae, interspaces smooth. Frontal lobes moderately wide 0.4 times as wide as head width, microreticulate with thick longitudinal costae. Frons shiny, entire surface longitudinally costate, interstices on sides distinctly microreticulate, in central part microreticulation diffused and surface appears partly smooth and shiny. Area above eyes and sides of head microreticulate and longitudinally costate, interstices appear slightly rugulose. Entire head bearing suberect to erect, pale and thin setae. Mesosoma. Pronotum anteriorly with regular microreticulation and on sides microreticulate with thin longitudinal costae. Scutum with dense, regular, thin longitudinal costae and more or less diffused microreticulation between costae, appears slightly shiny. Scutellum mostly with thin longitudinal costae only marrow median part with diffused sculpture, appears shiny (Fig. 5). Metanotum with slight sculpture, rugulose or punctate. Propodeum with distinct sculpture. Propodeal spines medium length [PSL / HW: 0.33], wide at base, triangular, straight, with acute apex. Area above propodeal spines with transverse, thin costae, dorsal surface of spines with longitudinal costae, sides of propodeum with concentric costae only area close to base of spines distinctly mocroreticulate but without costae, interstices between costae distinctly microreticulate, shiny. Area between propodeal spines on sides with longitudinal costae, centrally with distinct microreticulation, area below spines with transverse costae. Anepisternum and katepisternum with longitudinal costae and microreticulated interspaces, only anterior and posterior corners of anepisternum with small smooth and shiny areas, and katepisternum close to ventral margin with diffused costae. Metaepisternum and metakatepisternum, with dense, longitudinal costae and microreticulate interspaces only metakatepisternum close to ventral margin with diffused microreticulation and partly shiny. Dorsal surface of mesosoma with sparse, erect, long, thick and pale setae (Figs 5, 6). Petiole and postpetiole. Shiny anteriorly, dorsal and lateral surface microreticulate with sparse longitudinally costae. Gaster. Smooth and shiny, first tergite on whole surface and rest of tergites posteriorly bearing moderately dense, long, erect setae and sparse, short adhering setae (Figs 5, 6).

Etymology

Named after the historical Greek land of Messinia (Μεσσηνία), Peloponnese, where specimens from the type series were collected.

General distribution

Greece: southern Ionian Islands and Peloponnese.

Biology

Specimens of T. messiniaensis were collected from sunny localities in lowlands and highlands (8 - 670 m a.s.l). The species was noted in various habitats, most often on shrubs growing along roadsides and olive plantations, maquis, phrygana and forests (cypress, deciduous and mixed). We noted also its presence in a park in a centre of a city and, occasionally, in shrubs located in a pine forest located on a rocky hill. Nests were found inside dry stems of various herbs and shrubs. Colonies polygynous.

The following ant species were recorded in the same areas as Temnothorax messiniaensis:

For localities on Ionian Islands: Cephalonia: Avithos Lake, Katapodata, 1.5 NE of Koulourata ancient Same, rd. Skala-Poros, Zakynthos: 1 km N of Exo Chora, 1.2 km N of Vasilikos, 1.2 km NE of Anafonitria, 1.2 km SW of Skinaria, 1.8 km SW of Volimes, 1.9 km W of Maries, 470 m NE of Orthonies, 700 m SW of Koroni, 800 m SE of Xirokastello, 880 m S of Orthonies, Ag. Georgiou monastery, Ag. Joannis, Vrachionas Mts. and Peloponnese, Messinia, Taygetos Mts., 0.8 km SE of Exochori see Temnothorax brackoi.

Ionian Islands, Cephalonia, 1.6 km SW of Digaleto: Aphaenogaster balcanica (Emery), A. muelleriana Wolf, Camponotus aethiops (Latreille), C. dalmaticus (Nylander), Crematogaster schmidti (Mayr), Pheidole balcanica Seifert, Plagiolepis pygmaea (Latreille), Temnothorax bulgaricus (Forel), T. laconicus Csősz et al., T. rogeri Emery, T. strymonensis Csősz, Salata & Borowiec; Tetramorium kephalosi Salata & Borowiec; 1.8 km SW of Digaleto: Aphaenogaster balcanica (Emery), Camponotus aethiops (Latreille), C. dalmaticus (Nylander), Crematogaster sordidula (Nylander), Pheidole balcanica Seifert, Plagiolepis pygmaea (Latreille), Temnothorax laconicus Csősz et al., T. rogeri Emery; Kapandriti vicinity: Aphaenogaster balcanica (Emery), Camponotus dalmaticus (Nylander), C. gestroi Forel, C. kiesenwetteri (Roger), Colobopsis truncata (Spinola), Crematogaster schmidti (Mayr), Plagiolepis pygmaea (Latreille), Temnothorax leviceps (Emery); 800 m S of Kateleios: Camponotus dalmaticus (Nylander), C. lateralis (Olivier), Crematogaster schmidti (Mayr), C. sordidula (Nylander), Lasius alienus (Förster), L. illyricus Zimmermann, Liometopum microcephalum (Panzer), Pheidole balcanica Seifert, Plagiolepis pygmaea (Latreille), Temnothorax graecus (Forel), T. leviceps (Emery), T. cf. unifasciatus; Kremmidi: Aphaenogaster balcanica (Emery), Camponotus aethiops (Latreille), C. gestroi Forel, C. dalmaticus (Nylander), C. kiesenwetteri (Roger), Crematogaster schmidti (Mayr), C. sordidula (Nylander), Pheidole balcanica Seifert; Plagiolepis pygmaea (Latreille), Temnothorax bulgaricus (Forel), T. graecus (Forel), T. leviceps (Emery); Moni Aprilion: Aphaenogaster balcanica (Emery), Camponotus kiesenwetteri (Roger), Pheidole balcanica Seifert, Plagiolepis pygmaea (Latreille), Temnothorax rogeri Emery, T. strymonensis Csősz, Salata & Borowiec; 1.7 km NW of Pastra: Aphaenogaster balcanica (Emery), A. cf. epirotes, Bothriomyrmex communistus Santschi, Camponotus dalmaticus (Nylander), C. gestroi Forel, C. kiesenwetteri (Roger), C. lateralis (Olivier), Crematogaster schmidti (Mayr), C. sordidula (Nylander), Lepisiota frauenfeldi (Mayr), Messor wasmanni Krausse, Plagiolepis pygmaea (Latreille), Tetramorium kephalosi Salata & Borowiec; near Peratata: Aphaenogaster balcanica (Emery), A. muelleriana Wolf, Camponotus aethiops (Latreille), C. dalmaticus (Nylander), C. lateralis (Olivier), Crematogaster schmidti (Mayr), C. sordidula (Nylander), Lasius lasioides (Emery), Messor ibericus Santschi, Pheidole pallidula (Nylander), Plagiolepis pygmaea (Latreille), Temnothorax clypeatus (Mayr); Skala vicinity loc. 1: Aphaenogaster balcanica (Emery), Camponotus aethiops (Latreille), C. dalmaticus (Nylander), C. kiesenwetteri (Roger), Colobopsis truncata (Spinola), Crematogaster sordidula (Nylander), Plagiolepis pallescens Forel, P. pygmaea (Latreille), Proformica oculatissima (Forel) Temnothorax cf. exilis, Tetramorium kephalosi Salata & Borowiec; Skala vicinity loc. 2: Aphaenogaster balcanica (Emery), Camponotus dalmaticus (Nylander), C. kiesenwetteri (Roger), C. lateralis (Olivier), C. oertzeni Forel, Crematogaster schmidti (Mayr), C. sordidula (Nylander), Lasius alienus (Förster), Plagiolepis pygmaea (Latreille), Temnothorax exilis Emery, Tetramorium kephalosi Salata & Borowiec.

Ionian Islands, Zakynthos, 1.2 km SE of Loucha: Aphaenogaster balcanica (Emery), A. cf. epirotes, Bothriomyrmex communista Santschi, Camponotus aethiops (Latreille), C. dalmaticus (Nylander), C. gestroi Emery, C. kiesenwetteri (Roger), C. lateralis (Olivier), Crematogaster schmidti (Mayr), Lepisiota frauenfeldi (Mayr), Messor wasmanni Krausse, Tapinoma erraticum (Latreille), Temnothorax bulgaricus (Forel), T. exilis (Emery), T. graecus (Forel), T. rogeri Emery, T. cf. tergestinus, Tetramorium diomedeum Emery; 1.4 km S of Lithakia: Aphaenogaster balcanica (Emery), Camponotus dalmaticus (Nylander), C. kiesenwetteri (Roger), C. oertzeni Forel, Crematogaster schmidti (Mayr), C. sordidula (Nylander), Lepisiota frauenfeldi (Mayr), L. melas (Emery), Messor wasmanni Krausse, Pheidole cf. pallidula, Plagiolepis pygmaea (Latreille), Temnothorax rogeri Emery, T. cf. tergestinus; 330 m S of Stimies: Aphaenogaster balcanica (Emery), Bothriomyrmex communista Santschi, Camponotus kiesenwetteri (Roger), C. oertzeni Forel, Crematogaster schmidti (Mayr), C. sordidula (Nylander), Messor wasmanni Krausse, Pheidole cf. pallidula, Plagiolepis pygmaea (Latreille), Temnothorax exilis (Emery), T. graecus (Forel); 580 m SW of Lithakia: Aphaenogaster cf. epirotes, Camponotus aethiops (Latreille), C. dalmaticus (Nylander), C. gestroi Emery, C. kiesenwetteri (Roger), C. lateralis (Olivier), Crematogaster schmidti (Mayr), C. sordidula (Nylander), Messor ibericus Santschi, M. wasmanni Krausse, Plagiolepis pygmaea (Latreille), Temnothorax exilis (Emery), T. cf. tergestinus, Tetramorium diomedeum Emery; 600 m E of Ag. Leon: Aphaenogaster balcanica (Emery), A. muelleriana Wolf, Camponotus aethiops (Latreille), C. dalmaticus (Nylander), C. kiesenwetteri (Roger), C. lateralis (Olivier), Crematogaster schmidti (Mayr), Lepisiota frauenfeldi (Mayr), L. melas (Emery), Messor ibericus Santschi, M. wasmanni Krausse, Pheidole cf. pallidula, Plagiolepis pygmaea (Latreille), Temnothorax bulgaricus (Forel), T. rogeri Emery, T. cf. tergestinus, Tetramorium kephalosi Salata & Borowiec.

Peloponnese, Messinia, 2 km E of Kalamata: Aphaenogaster balcanica (Emery), A. cf. muelleriana, Camponotus gestroi Emery, C. ionius Emery, C. kiesenwetteri (Roger), C. laconicus Emery, C. lateralis (Olivier), Crematogaster schmidti (Mayr), C. sordidula (Nylander), Formica clara Forel, Lasius illyricus Zimmermann, L. lasioides (Emery), L. neglectus Van Loon, Boomsma & Andrasfalvy, Lepisiota frauenfeldi (Mayr), Messor wasmanni Krausse, Nylanderia jaegerskioeldi (Mayr), Pheidole pallidula (Nylander), Plagiolepis pygmaea (Latreille), Temnothorax graecus (Forel), T. cf. luteus, T. recedens (Nylander), T. rogeri Emery, Tetramorium cf. caespitum; Egaleo Mts., 1.4 km S of Flesiada: Aphaenogaster balcanica (Emery), A. cf. muelleriana, Camponotus aethiops (Latreille), C. dalmaticus (Nylander), Crematogaster schmidti (Mayr), C. sordidula (Nylander), Lasius lasioides (Emery), Pheidole pallidula (Nylander), Plagiolepis pygmaea (Latreille), Tapinoma erraticum (Latreille), Temnothorax exilis (Emery), T. laconicus Csősz et al., T. morea Csősz, Salata & Borowiec; Kalamata, old centre: Aphaenogaster balcanica (Emery), Crematogaster schmidti (Mayr), Formica clara Forel, Lepisiota frauenfeldi (Mayr), L. melas (Emery), Messor hellenius Agosti & Collingwood, Pheidole pallidula (Nylander), Plagiolepis pygmaea (Latreille), Solenopsis cf. lusitanica, Temnothorax graecus (Forel), Tetramorium cf. hungaricum, Trichomyrmex perplexus Radchenko; Kalamata, railway park: Camponotus lateralis (Olivier), Crematogaster schmidti (Mayr), Formica clara Forel, Lasius illyricus Zimmermann, L. lasioides (Emery), L. neglectus Van Loon, Boomsma & Andrasfalvy, Messor wasmanni Krausse, Pheidole indica Mayr, Plagiolepis pygmaea (Latreille), Temnothorax graecus (Forel), Tetramorium cf. caespitum; Kondovounia Mts., 0.8 km N of Koromilea: Aphaenogaster balcanica (Emery), A. cf. muelleriana, Camponotus dalmaticus (Nylander), C. gestroi Emery, C. laconicus Emery, C. lateralis (Olivier), Cataglyphis nodus (Brullé), Crematogaster schmidti (Mayr), Lasius lasioides (Emery), Lepisiota frauenfeldi (Mayr), Pheidole pallidula (Nylander), Plagiolepis pygmaea (Latreille), Temnothorax cf. bulgaricus, T. exilis (Emery), T. laconicus Csősz et al., T. morea Csősz, Salata & Borowiec, T. recedens (Nylander); Taygetos Mts., 0.8 km W of Eleochori: Aphaenogaster balcanica (Emery), Camponotus gestroi Emery, C. ionius Emery, C. kiesenwetteri (Roger), Lepisiota nigra (Dalla Torre), Messor wasmanni Krausse, Pheidole pallidula (Nylander), Plagiolepis pygmaea (Latreille), Temnothorax exilis (Emery), Tetramorium kephalosi Salata & Borowiec.

Temnothorax turcicus (Santschi, 1934)

Figs 7, 8, 11, 15

Leptothorax turcicus Santschi, 1934: 278.

Type material

Syntype, worker (pin): • [TURKEY]: Izmir | 29.VII.33, Santschi || Type || Sammlung | Dr. F. Santschi | Kairouan | ANTWEB | CASENT0913009 (NHMB).

Other material

Greece. North Aegean, Lesbos: • 2w. (pin) (CASENT0846851-CASENT0846852): n. Ahladeri, 39.15958N/26.29292E, 9 m, 2015-06-10, leg. L. Borowiec. Macedonia, Chalkidiki: • 12w. (pin) (CASENT0846853-CASENT0846864): Holomontas, Taxiarhis vicinity, mountain deciduous forest, in leaf litter, 40.4N/23.51666E, 594 m, 2009-08-30, leg. L. Borowiec; • 2w. (pin) (CASENT0846865-CASENT0846866): Holomontas, Stagira, on wall in deciduous forest, 40.52896N/23.74872E, 539 m, 2009-09-03, leg. L. Borowiec; • 5w. (pin) (CASENT0846867-CASENT0846871): Holomontas, Stagira-Neochori road, on wall in deciduous forest, 40.51666N/23.7E, 512 m, 2009-09-03, leg. L. Borowiec. Macedonia, Kavalas: • 1w. (pin) (CASENT0846872): Nestos river near Komnina, 41.169N/24.6966E, 100 m, 1999-10-10, leg. E. Nikolakakis. Macedonia, Pieria: • 5w. (pin) (CASENT0846873-CASENT0846877), 2w. (EtOH): road to P. Poroi loc. 1, roadsides with shrubs, 39.97963N/22.61563E, 110 m, 2019-05-17, leg. L. Borowiec; • 6w. (EtOH): road to P. Poroi loc. 2, roadsides with shrubs, 39. 97627N/22.61146E, 185 m, 2019-05-17, leg. L. Borowiec; •14w. (pin) (CASENT0846878-CASENT0846891), 40w. (EtOH): road to P. Poroi loc. 3, roadsides with shrubs, 39.96863N/22.60494E, 260 m, 2019-05-17, leg. L. Borowiec. Peloponnese, Arcadia: • 1w. (pin) (CASENT0846892): 3.2 km NW Polidroso, 1000 m, 37.19874N/2257603E, 1000, 2016-06-18, leg. L. Borowiec. Peloponnese, Laconia: • 6w. (pin) (CASENT0846893-CASENT0846898): Parnon Mts., 5 km NE of Karies, 37.324N/22.538E, 1000 m, 2000-04-29, leg. A. Schulz & K. Vock (3w DBET, 3w PW). Sterea Ellas, Euboea: • 1w. (pin) (CASENT0846899): 1 km NE of Amfithea, 38.5519N/23.79546, 200 m, 2018-06-10, leg. L. Borowiec. Thessaly, Larissa: • 1w. (pin) (CASENT0846900): Kato Olimbos Mts, 6.1 km S of Kalipefki, 39.91322N/22.4641E, 855 m, 2017-05-09, leg. L. Borowiec.

Terra typica

Greece, Thessaly, Mt. Ossa.

Differential diagnosis

Differentiation from T. kemali, T. brackoi and T. messiniaensis – see differential diagnosis in T. messiniaensis. Temnothorax turcicus differs from specimens of T. aveli with long propodeal spines in thin dark band on first gastral tergite, head not darker from mesosoma, mesosoma less convex in profile and propodeal spines directed slightly more upwards; from T. lagrecai (Baroni Urbani, 1964), species described and known only from Sicily, differs in petiolar node dorsum flat or slightly convex and distinctly bigger mesosoma size - ML 0.595 ± 0.50 (0.517–0.680) vs. ML = 0.779 ± 0.05 (0.677–0.832).

Redescription

Worker (n = 10): HL: 0.670 ± 0.03 (0.614–0.696); HW: 0.573 ± 0.03 (0.522–0.596); SL: 0.465 ± 0.02 (0.431–0.484); EL: 0.158 ± 0.009 (0.149–0.174); EW: 0.118 ± 0.01 (0.102–0.137); WL: 0.779 ± 0.05 (0.677–0.832); PSL: 0.192 ± 0.005 (0.186–0.199); SDL: 0.118 ± 0.007 (0.106–0.124); PEL: 0.271 ± 0.015 (0.248–0.286); PPL: 0.173 ± 0.01 (0.149–0.186); PEH: 0.206 ± 0.015 (0.186–0.230); PPH: 0.205 ± 0.02 (0.174–0.236); PNW: 0.394 ± 0.02 (0.360–0.422); PLW: 0.169 ± 0.007 (0.161–0.180); PPW: 0.219 ± 0.01 (0.199–0.230); CI: 85.6 ± 1.2 (83.6–86.8); SI1: 69.4 ± 0.6 (68.5–70.2); SI2: 81.1 ± 1.3 (79.2–82.7); MI: 50.7 ± 1.3 (49.3–53.2); EI1: 75.3 ± 8.8 (66.1–91.7); EI2: 17.7 ± 1.3 (16.4–20.0); PI: 132.3 ± 9.2 (120.0–148.4); PPI: 84.8 ± 4.5 (75.8–88.9); PSI: 162.7 ± 7.5 (155.0–176.5).

Colour

Whole body uniformly yellow to dark yellow, sometimes club in darker yellow colouration. Gaster yellow, only the first gaster tergite with very thin, dark band on its posterior part (Figs 7, 8). Head. Oval, 1.16 times as long as wide, lateral surfaces below and above eyes gently convex, posterior edges convex, occipital margin of head straight or slightly convex (Fig. 15). Anterior margin of clypeus slightly convex, medial notch absent. Eyes moderate, oval, 1.34 times as long as wide. Antennal scape short, in lateral view slightly curved, 0.69 times as long as length of the head, in apex gradually widened, its base with small, triangular tooth, funiculus long, club 3 segmented (Figs 12, 15). Surface of scape with very fine microreticulation, shiny, covered with thin, moderate dense, decumbent setae. Mandibles rounded with thick sparse, longitudinal striae, shiny. Clypeus shiny with thick, sparse, longitudinal striae, area between striae smooth and shiny. Frontal carinae short, not extending beyond frontal lobes. Antennal fossa deep, with irregular, dense to sparse, thick rugosity and sometimes with a few thin, roundly curved striae, surface between thick sculpture smooth or with sparse microreticulation. Frontal lobes narrow, smooth with slight, dense longitudinal striation (Fig. 15). Frons, vertex and temples with dense, thick, longitudinal reticulation, central surface of frons and vertex with longitudinal reticulation sparser or reduced, with additional thin, longitudinal striation, striae sometimes interrupted, surface between striation smooth and shiny; malar area with irregular, thick, reticulation, space between reticulation smooth or with very sparse microreticulation, shiny; genae with sparser, than on frons, and thick reticulation, shiny (Fig. 15). Frons and vertex with erect, pale, short and thick setae. Mesosoma. Elongate, 1.98 times as long as wide, slightly arched in profile. Metanotal groove absent. Pronotum convex on sides. Propodeal spines long, directed upward, with base slightly to moderate wider than apex, tips sharp (Fig. 8). Whole surface with dense, reticulation, sometimes its dorsal surface and lateral surfaces of pronotum and mesonotum with additional thick, sparse longitudinal wrinkles. Area between thick sculpture shiny, smooth or sometimes with sparse, fine microreticulation (Fig. 8). Entire mesosoma bearing erect, pale, short and thick setae (Fig. 8). Petiole. In lateral view, with short peduncle, node moderate high, with anterior face straight, and posterior face convex and dorsum flat or slightly convex. Peduncle and petiolar node shiny, with thick, dense reticulation, area between rugae smooth, dorsum with sparser reticulation. Dorsal surface bearing sparse, short, erect setae (Fig. 8). Postpetiole. In lateral view, regularly convex, apical half with gently convex sides (Fig. 8), on the whole surface shiny, with thick, dense reticulation, dorsum with sparser reticulation; area between rugae smooth. Dorsal surface bearing sparse, short, erect setae. Gaster. Gaster smooth and shiny, bearing erect, thin, pale setae (Figs 7, 8).

Figures 7, 8. 

Worker of Temnothorax turcicus (Santschi) 7 Dorsal 8 Lateral.

Figures 9–12. 

Head and antennae 9 Worker of Temnothorax brackoi sp. nov. 10 Worker of Temnothorax messiniaensis sp. nov. 11 Worker of Temnothorax turcicus (Santschi) 12 Gyne of Temnothorax messiniaensis sp. nov.

General distribution

Eastern Austria, Bulgaria, Croatia, Greece: Macedonia, North Aegean Islands, Sterea Ellas, Peloponnese, and Thessaly, Hungary, Slovakia, western Turkey.

Comment

We examined a syntype of Temnothorax tauricus (Ruzsky, 1902) preserved in Forel’s collection (MHNG) and it appears to be very similar to specimens of T. turcicus collected in Greece. The only difference is a slightly darkened antennal club in the syntype specimen of T. tauricus (all studied specimens of T. turcicus have antennae uniformly yellow). We discussed this issue with Alex Radchenko (Kiev, Ukraine) who confirmed that all 17 syntypes of T. tauricus preserved in Karavaiev’s collection (Kiev, Ukraine) also have slightly darkened antennal club. In our opinion this difference could be an infraspecific variation. Within nest samples of Temnothorax messiniaensis, a member of the aveli species group, we observed single specimens with more or less darkened antennal club. Temnothorax tauricus was recorded from Caucasus and southern Ukraine but is sympatric with T. turcicus in Bulgaria and Greece. Temnothorax tauricus have nests in dry stems of herbs, grasses or rarely in soil under stones (Radchenko 2016) and by those preferences reminds species of the T. aveli species group. Clarification of taxonomic relation between those two taxa requires further study based on material collected from the whole distribution range of both species. If our supposition on the conspecifity of both taxa is confirmed, then the name T. tauricus will have priority over the name T. turcicus.

Biology

Specimens collected on shadow localities, from seacoast to 1000 m a.s.l. Foraging workers were observed on herbs in stream valley of tourist resort, valleys with Platanus trees, mountain coniferous forest and mountain pastures close to border of coniferous forest. Nests were not found, probably like other species of this group, are located inside dry stems of herbs.

The following ant species were recorded in the same areas as T. turcicus:

For localities on Macedonia: Pieria, road to P. Poroi loc. 1, Pieria, road to P. Poroi loc. 2, Pieria, road to P. Poroi loc. 2, and Thessaly, Larissa, Mt. Ossa, Kokkino Nero: see Temnothorax brackoi.

North Aegean, Lesbos, near Ahladeri: Camponotus dalmaticus (Nylander), C. lateralis (Olivier), C. sanctus, Crematogaster ionia Forel, Lasius neglectus Van Loon, Boomsma & Andrasfalvy, Monomorium monomorium Bolton, Pheidole pallidula (Nylander), Plagiolepis perperamus Salata et al., Temnothorax bulgaricus (Forel), T. cf. luteus, Tetramorium rhodium Emery.

Peloponnese, Arcadia, 3.2 km NW Polidroso: Aphaenogaster cf. subterranea, Camponotus aethiops (Latreille), C. dalmaticus (Nylander), C. nitidescens Forel, C. vagus (Scopoli), Cataglyphis nodus (Brullé), Crematogaster ionia Forel, Formica cunicularia Latreille, F. fusca Linnaeus, Lasius bomycina Seifert & Galkowski, L. flavus (Fabricius), L. illyricus Zimmermann, Pheidole cf. pallidula, Plagiolepis pygmaea (Latreille), Temnothorax crasecundus Seifert & Csősz, T. helenae Csősz et al., T. laconicus Csősz et al., T. cf. unifasciatus, Tetramorium cf. caespitum.

Sterea Ellas, Euboea, 1 km NE of Amfithea: Camponotus lateralis (Olivier), Cataglyphis nodus (Brullé), Crematogaster schmidti (Mayr), Pheidole cf. pallidula, Plagiolepis pygmaea (Latreille), Temnothorax bulgaricus (Forel), T. recedens (Nylander).

Thessaly, Larissa, Kato Olimbos Mts, 6.1 km S of Kalipefki: Aphaenogaster epirotes (Emery), Bothriomyrmex communista Santschi, Camponotus aethiops (Latreille), C. oertzeni Forel, C. piceus (Leach), Formica cunicularia Latreille, Lasius alienus Förster, Messor mcarthuri Steiner et al., M. wasmanni Krausse, Plagiolepis pygmaea (Latreille), Temnothorax cf. unifasciatus, Tetramorium cf. caespitum.

A key to Greek members of the T. aveli species group

1 Propodeal spines short to moderate (PSI<155), triangular, with wide base (Fig. 2); head on whole frontal surface with regular reticulation, without longitudinal striation, with dull background (Fig. 13) T. brackoi sp. nov.
Propodeal spines long (PSI > 155), thin, with base slightly to moderate wider than base (Figs 4, 8); head regularly reticulate but often with additional thin, longitudinal striation and partly shiny background (Figs 14, 15) 2
2 Central surface of frons with dense, thick, longitudinal reticulation and sometimes with additional thin, longitudinal striation (Fig. 10); first gaster tergite with wide, dark band on its posterior part (Figs 3–4); gena, mid and hind femora often obscured in middle T. messiniaensis sp. nov.
Central surface of frons and vertex with longitudinal reticulation sparser than of rest of head or reduced (Fig. 12); first gaster tergite with very thin, dark band on its posterior part (Figs 7, 8); gena, mid and hind femora never obscured in middle T. turcicus (Santschi)
Figures 13–15. 

Head sculpture 13 Worker of Temnothorax brackoi sp. nov. 14 Worker of Temnothorax messiniaensis sp. nov. 15 Worker of Temnothorax turcicus (Santschi).

Description of a new species of the Temnothorax nylanderi species group

Temnothorax triangularis sp. nov.

Figs 16–17, 18–20, 21, 22, 23–24

Differential diagnosis

Temnothorax triangularis belongs to the T. nylanderi species group. It differs from most of members of this group in uniformly brown body with darker frons and posterior band of first gastral tergite, and absent or very shallow, inconspicuous metanotal groove. There are four other known species with dark body colouration: T. laconicus, T. artvinensis, T. sordidulus, and T. tergestinus. Temnothorax triangularis differs from all of them in extremely shallow metanotal groove and frons lacking reticulation and covered with dense, thick, longitudinal striations, additionally frons centre has sculpture weaker or reduced, it differs also from T. laconicus and T. artvinensis in shorter, triangular propodeal spines.

Etymology

Named after short, triangular propodeal spines.

Type material

Holotype, worker (pin) (CASENT0846901): GREECE, Sterea Ellas, Eubea | 2.4 km SW of Stropones | 38,60327N/23,87E, 1025 m |10 VI 2018, L. Borowiec || Collection L. Borowiec | Formicidae | LBC-GR02682 (MNHW).

Paratypes, 25w., 1Q.(pin) (CASENT0846661-CASENT0846691): the same nest sample as holotype (DBET, BMNH, CASC, MHNG); 5w. (EtOH): the same locality as holotype, collected in litter (DBET); 11w, 1Q (pin) (CASENT0846692-CASENT0846704): GREECE, Sterea Ellas | Eubea |3.7 km SW of Metochi, 830 m | 38.60402N/23.91683E, | 13 VI 2018, L. Borowiec (DBET).

Terra typica

Euboea, Greece.

Other material

GREECE, Sterea Ellas, Euboea: 2w. (EtOH): 1.5 km SW of Koutourla, 38.62838N/23.92772E, 695 m, 2018-06-13, leg. L. Borowiec; 1w.(pin) (CASENT0846902), 2w. (EtOH): 2.3 km S of Stropones, 38.9933N/23.87807E, 860 m, 2018-06-10, leg. L. Borowiec; 2w. (EtOH): 2.7 km SE of Stropones, 38.59851N/23.9085E, 855 m, 2018-06-13, leg. L. Borowiec; 3w. (EtOH): 2.9 km S of Stropones, 38.59133N/23.88562E, 880 m, 2018-06-13, leg. L. Borowiec; 43w. (EtOH): 3.7 km SW of Metochi, 38.60402N/23.91683E, 830 m, 2018-06-13, leg. L. Borowiec.

Description of worker (n = 10): HL: 0.684 ± 0.01 (0.671–0.708); HW: 0.603 ± 0.02 (0.578–0.650); SL: 0.504 ± 0.02 (0.484–0.534); EL: 0.140 ± 0.01 (0.124–0.149); EW: 0.105 ± 0.006 (0.093–0.112); WL: 0.769 ± 0.02 (0.742–0.820); PSL: 0.161 ± 0.02 (0.143–0.183); SDL: 0.119 ± 0.006 (0.112–0.130); PEL: 0.294 ± 0.02 (0.273–0.323); PPL: 0.182 ± 0.008 (0.174–0.199); PEH: 0.245 ± 0.01 (0.236–0.270); PPH: 0.231 ± 0.01 (0.217–0.248); PNW: 0.403 ± 0.01 (0.388–0.435); PLW: 0.177 ± 0.009 (0.168–0.199); PPW: 0.241 ± 0.01 (0.230–0.267); CI: 88.2 ± 2.2 (85.3–92.7); SI1: 73.7 ± 1.5 (71.2–76.4); SI2: 83.6 ± 2.4 (78.9–87.5); MI: 52.4 ± 1.2 (50.8–54.7); EI1: 75.0 ± 4.2 (68.8–85.0); EI2: 15.3 ± 1.0 (13.6–16.7); PI: 120.1 ± 5.4 (108.0–128.8); PPI: 79.0 ± 2.8 (75.0–85.7); PSI: 135.8 ± 12.4 (120.0–155.6).

Colour

Whole body uniformly brown to bright brown, sometimes mesosoma and genae brighter. Legs and antennae bright brown to dark yellow, femora in central part darkened (Figs 16, 17). Head. Oval, 1.14 times as long as wide, lateral surfaces below and above eyes gently convex, posterior edges convex, occipital margin of head straight or slightly concave (Figs 18, 19). Anterior margin of the clypeus slightly convex, medial notch absent. Eyes small, oval, 1.33 times as long as wide. Antennal scape short, in lateral view slightly curved, 0.73 times as long as length of the head, in apex gradually widened, its base with small, triangular tooth, funiculus long, club 3-segmented (Fig. 18). Surface of scape with very fine microreticulation, shiny, covered with thin, moderate dense, decumbent setae. Mandibles rounded with thick sparse, longitudinal striae, shiny. Clypeus shiny with thick, longitudinal striae, area between striae shiny with few longitudinal wrinkles. Frontal carinae short, not extending beyond frontal lobes. Antennal fossa deep, with sparse, thin roundly curved striae, area between striae with sparse and fine reticulation, shiny. Frontal lobes narrow, smooth with slight, dense longitudinal striation (Figs 18, 19). Space between frontal carinae and vertex with dense, thick, longitudinal striation, sparser or reduced in the central part, striae sometimes interrupted, surface between striae smooth and shiny; space between frontal carinae ad eyes, temples and malar area with longitudinal, thick reticulation, space between reticulation smooth or with very sparse microreticulation, shiny; genae with very sparse thick reticulation, partly smooth, always shiny. Frons and vertex with erect, pale, short and thick setae. Mesosoma. Elongate, 1.9 times as long as wide, slightly arched in profile. Metanotal groove absent or very shallow, inconspicuous. Pronotum convex on sides. Propodeal spines short, triangular, with wide base, directed upward, with angulate tips (Fig. 21), only in fewer than 20% of specimens are propodeal spines moderately long, with wide bases and sharp tips (Fig. 22). Lateral surfaces of pronotum with thick and sparse longitudinal striation or reticulation, its dorsal surface with thick, sparse, irregular reticulation; mesonotum and propodeum on the whole surface with thick, denser than on pronotum, irregular reticulation. Area between thick sculpture shiny, smooth or sometimes with sparse, fine microreticulation (Fig. 17). Entire mesosoma bearing erect, pale, short and thick setae (Fig. 17). Petiole. In lateral view, with short peduncle, node high, with anterior face flat to slightly convex, and posterior face and dorsum convex. Peduncle and petiolar node shiny, with thick, dense reticulation, area between rugae smooth, dorsum with sparser reticulation. Dorsal surface bearing sparse, short, erect setae (Figs 21, 22). Postpetiole. In lateral view, regularly convex, apical half with gently convex sides (Figs 21, 22), on the whole surface shiny, with thick, dense reticulation, dorsum with sparser reticulation; area between rugae smooth. Dorsal surface bearing sparse, short, semierect to erect setae. Gaster. Smooth and shiny, bearing erect, thin, pale setae (Figs 16, 17).

Figures 16, 17. 

Worker of Temnothorax triangularis sp. nov. 16 Dorsal 17 Lateral.

Description of gyne (n = 2). Colour. Head brown, temples slightly brighter coloured than frontal parts. Antennae uniformly yellow. Mesosoma, petiole and postpetiole bright brown, legs yellow. First gastral tergite mostly dark brown with yellowish brown spot basally, remaining tergites brown with dark brown posterior margins (Figs 23, 24). Head. Eyes big, almost round [EL / HW: 0.26]. Antennal scape short [SL / HW: 0.76], not reaching occipital margin of head. Clypeus smooth and shiny laterally with diffused, longitudinal carinulae. Antennal fossa deep, rugulose with concentric carinae. Frontal lobes wide 0.43 times as wide as head width, rugulose with thick longitudinal costae, interstices microreticulate (Fig. 20). Frons shiny, entire surface longitudinally costate and rugose, interstices distinctly microreticulate. Area above eyes and sides of head rugulose and partly longitudinally costate, only small area behind eyes smooth and shiny. Entire head bearing erect, pale and thin setae. Mesosoma. Pronotum with thick rugosities in anterior part, sides with thick rugosity and dense longitudinal costae. Surface between rugosities microreticulate. Scutum with dense, thick longitudinal costae and microreticulation between costae but appears shiny. Scutellum laterally with thick longitudinal costae, to the centre costae partly diffused, along middle smooth and shiny area (Figs 23, 24). Metanotum with fine longitudinal costae and microreticulated background. Propodeum with area above propodeal spines with transverse and on sides with longitudinal costae and microreticulated background. Propodeal spines short [PSL / HW: 0.28], triangular, with wide base, straight and angulate apex. Area between and below propodeal spines with distinct microreticulation tends to form transverse ridges. Anepisternum and katepisternum with gentle, dense longitudinal costae. Metaepisternum and metakatepisternum, with dense, longitudinal costae and shiny area close to ventral margin. Surface between costae microreticulate. Dorsal surface of mesosoma with sparse, erect, long, thick and pale setae (Figs 23, 24). Petiole and postpetiole. Microreticulate, the entire surface punctate to rugulose, dorsal surface longitudinally costulate. Gaster. Smooth and shiny, bearing sparse, long, erect setae (Figs 23, 24).

Figures 18–20. 

Temnothorax triangularis sp. nov. 18 Worker head and antennae 19 Worker head sculpture 20 Gyne head and antennae.

General distribution

Greece, Sterea Ellas, Euboea.

Biology

Alpine species. Ants were observed on stones, dry branches, and herbs in coniferous forest, or coniferous forest with an admixture of chestnut. Nests were found in the dry branches of conifers lying on the ground.

The following ant species were recorded in the same areas as T. triangularis:

Sterea Ellas, Euboea, 1.5 km SW of Koutourla: Aphaenogaster subterranea (Latreille), Formica fusca Linnaeus, Lasius alienus Förster, L. brunneus (Latreille), L. illyricus Zimmermann, Pheidole pallidula (Nylander), Plagiolepis pygmaea (Latreille), P. pallescens Forel, Temnothorax helenae Csősz et al., T. lichtensteini (Bondroit), T. unifasciatus (Latreille); 2.3 km S of Stropones: see Temnothorax brackoi; 2.4 km SW of Stropones: Camponotus aethiops (Latreille), Formica fusca Linnaeus, F. sanguinea Latreille, Lasius alienus Förster, L. illyricus Zimmermann, Messor structor (Latreille), Temnothorax crasecundus Seifert & Csősz, T. unifasciatus (Latreille), Tetramorium impurum (Förster); 2.7 km SE of Stropones: Aphaenogaster subterranea (Latreille), A. cf. subterranea, Camponotus aethiops (Latreille), C. oertzeni Forel, Formica fusca Linnaeus, Lasius alienus Förster, L. distinguendus (Emery), Tapinoma erraticum (Latreille), Temnothorax helenae Csősz et al., T. unifasciatus (Latreille);

Figures 21, 22. 

Variation of propodeal spines of Temnothorax triangularis sp. nov.

2.9 km S of Stropones: Camponotus piceus (Leach), C. vagus (Scopoli), Lasius brunneus (Latreille), L. flavus (Fabricius), Myrmica scabrinodis Nylander, Temnothorax helenae Csősz et al., T. unifasciatus (Latreille); 3.7 km SW of Metochi: Aphaenogaster subterranea (Latreille), A. cf. subterranea, Camponotus fallax (Nylander), C. vagus (Scopoli), Lasius illyricus Zimmermann, Temnothorax crasecundus Seifert & Csősz, T. helenae Csősz et al., T. unifasciatus (Latreille).

Note

Although this species has inconspicuous metanotal groove, which tends to disappear in some specimens, we decided to place it in the Temnothorax nylanderi group. General body shape, structure of petiole, unicolourous antennae, head and mesosoma sculpture presented by T. triangularis are very similar to those observed in large species of the group i.e. T. nylanderi (Förster), T. crassispinus (Karavaiev) or T. crasecundus Seifert & Csősz. Moreover, an inconspicuous metanotal groove was observed also in some samples of small species of the nylanderi group, such as T. helenae Csősz, Heinze & Mikó. Usually the depth of metanotal groove is more or less constant within nest samples.

Figures 23, 24. 

Gyne of Temnothorax triangularis sp. nov. 23 Dorsal 24 Lateral.

Figures 25–27. 

Distribution in Greece 25 Temnothorax brackoi sp. nov. 26 Temnothorax messiniaensis sp. nov. 27 T. turcicus (Santschi) denoted by red circles and T. triangularis sp. nov. by black circles.

Acknowledgements

We would like to thank Dr. Bernard Landry (Genève, Switzerland) for providing access to the collection of ants preserved in Muséum d’Histoire Naturelle, Geneve (MHNG). We wish to thank Matthew Prebus and Christophe Galkowski for their helpful comments on early drafts of this article. Thanks to Christophe Galkowski (Saint–Aubin–de–Médoc, France), Enrico Schifani (Palermo, Italy), Antonio Scupola (Museo di Storia Naturale, Verona), and Petr Werner (Prague, Czech Republic) for donations and loans of specimens and their valuable comments on French and Italian species of the Temnothorax aveli species group. Our thanks are also extended to Alex Radchenko (Kiev, Ukraine) for his comments on types of Temnothorax tauricus and possible synonymy of this taxon with T. turcicus. Lech Borowiec thanks Jolanta Świętojańska (University of Wrocław) for her assistance during his field trips.

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1 Crete, as an island, creates a well-defined unit and studies performed on Temnothorax known from the island were published in a separate paper (Salata et al. 2018). The majority of endemic taxa known from the island create a single-species group and the material collected from other Greek islands and regions adjacent to Crete confirms their uniqueness. Therefore, Cretan endemic species of Temnothorax are not included in the key.
2 Some specimens of the T. tuberum species group and T. interruptus species group can have very indistinct metanotal grooves. Usually it applies to single specimens within a nest sample.