Temnothorax species groups

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The following is based on Cagniant and Espadaler (1997), Csösz, Heinze, and Mikó (2015), Csősz, Salata, and Borowiec (2018), Gonzalez (2021), Radchenko (1995a,b,c), Salata and Borowiec (2015, 2019), and Salata, Środoń, and Borowiec (2023).

Temnothorax is one of the most speciose ant genera and much more taxonomic research is needed to describe every species and to understand how they are all related to one another.

Our current understanding of the Palearctic Temnothorax fauna suggests there remains a lot of descriptive and revisionary research to be done. Prebus (2017), in a worldwide phylogenetic study of Temnothorax, looked at a relatively small number of specimens from the Palearctic. He concluded the Palearctic Temnothorax are mostly a discrete set of species when compared to Temnothorax from other regions. Prebus also arranged the Palearctic Temnothorax species he examined into four sub-clades but this study was not meant to test the validity of past Palearctic species groups or to eastablish a rigorous new framework of species groups for the region. The groups and complexes given here are more working from the bottom up, looking to find how existing and new species fit together into different groupings from examining specimens of existing species and describing new species. It is likely it will be some time before there emerges a comphrensive, stable understanding of the taxonomy and phylogenetics of the Palearctic Temnothorax. Until then, the groupings given below and those presented in older publications will change as more studies are completed.

One final consideration is to understand this groupings are incomplete and subject to change. The lists of species associated with each group are, in many cases, only representative of members studied from a particular region and do not include species from other parts of the Palearctic.

Temnothorax affinis species group

Radchenko (1995c); synopsis by Salata and Borowiec (2019): 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 there is only one arboreal species.

Radchenko also placed only the namesake species in this group.

Temnothorax angustulus species group

Salata and Borowiec (2019): 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, Temnothorax dessyi.

Temnothorax anodontoides species group

Salata and Borowiec (2019): 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

Salata and Borowiec (2019): 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 species mostly known from lowland habitats, especially Mediterranean forests and bushes, nestling inside dry stems of trees and twigs of bushes and large herbs.

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

Radchenko (1995a); synopsis by Salata and Borowiec (2019): 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.

Temnothorax clypeatus species group

Radchenko (1995c); synopsis by Salata and Borowiec (2019): 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.

Temnothorax corticalis species group

Radchenko (1995c); synopsis by Salata and Borowiec (2019): 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 T. corticalis, an arboreal species associated with deciduous forests, is present.

Temnothorax exilis species group

Cagniant and Espadaler (1997); synoposis by Salata and Borowiec (2019): 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 Temnothorax exilis 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

Salata and Borowiec (2019): 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.

Results presented by Prebus (2017) revealed that the nylanderi species group, as defined by Csösz, Heinze, and Mikó (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.

The Temnothorax flavicornis species-complex was elevated to a species-group by Salata and Borowiec (2019), as noted above. Csösz, Heinze, and Mikó (2015) originally diagnosed the species complex as follows: Workers of the lone species of the complex can be distinguished from those of other (nylanderia species group) complexes treated in this revision by the combination of the following salient features: antennae 11-segmented, yellow to light brown color, head rectangular, significantly longer than broad head (CL/CWb [1.226, 1.299]), sculpture of head dorsum dull: with smooth, or inconspicuously areolate ground sculpture combined with longitudinally rugulose or reticulate main sculpture; long to very long propodeal spines (SPST/CS [0.303, 0.420]), deviating from longitudinal axis of mesosoma by 40–45°; petiolar node in lateral view with a concave frontal profile meeting conspicuously developed truncate dorsum in an obtuse angle (105–115°) with a narrowly rounded transition, without a conspicuous sharp fronto-dorsal ridge on the petiolar node.

Temnothorax graecus species group

Based on Salata, Środoń, and Borowiec 2023.

Key to Temnothorax graecus species-group from Greece

Temnothorax interruptus species group

Csősz, Salata, and Borowiec (2018); synopsis by Salata and Borowiec (2019): 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.

Temnothorax kemali species group

Salata and Borowiec (2019): 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.

+ at least one undescribed species

Temnothorax luteus species group

Salata and Borowiec (2019): 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.

+ at least one undescribed species

Temnothorax nylanderi species group

Radchenko (1995h), Csösz, Heinze, and Mikó (2015)

Temnothorax angulinodis species complex

Csösz, Heinze, and Mikó (2015): Workers of the Temnothorax angulinodis species-complex can be distinguished from those of other complexes treated in this revision by the combination of the following salient features: brown to black color; longer than broad head (CL/CWb [1.171, 1.222]), sculpture of head dorsum shiny: with inconspicuously areolate ground sculpture combined with parallel costulate main sculpture; petiolar node in lateral view with a straight or weakly concave frontal profile meeting dorso-caudal plate in an acute angle (72–82°) with a sharp ridge, in dorsal view appearing as conspicuous semicircular anterior-lateral rim; long propodeal spines (SPST/CS [0.332, 0.369]), deviating from longitudinal axis of mesosoma by 32–38°. This peculiar species-complex consists of a single species and is only known from the Peloponnese peninsula.

Temnothorax angustifrons species complex

Csösz, Heinze, and Mikó (2015): Workers of the Temnothorax angustifrons species-complex can be distinguished from those of other complexes by the combination of the following salient features: light yellow to light brown color; moderately longer than broad head (CL/CWb [1.135, 1.254]), sculpture of head dorsum shiny, with inconspicuously areolate ground sculpture combined with feeble costulate main sculpture; short to moderately long propodeal spines (SPST/CS [0.159, 0.267]), deviating from longitudinal axis of mesosoma by 47–52°; petiolar node in lateral view with a weakly concave frontal profile meeting dorso-caudal plate in an obtuse angle (95–105°) with a moderately sharp ridge, in dorsal view appearing as a visible (occasionally inconspicuous) anterior-lateral rim.

Members of this species-complex are known to occur in Turkey and Crete. The occurrence of two samples found in Greece may be ascribed to, dispersal by human activities.

Temnothorax lichtensteini species complex

Csösz, Heinze, and Mikó (2015): Workers of the Temnothorax lichtensteini species-complex can be distinguished from those of other complexes treated in this revision by the combination of the following salient features: dirty yellow to brown color; longer than broad head (CL/CWb [1.181, 1.267]), sculpture of head dorsum dull: with areolate ground sculpture combined with longitudinally rugulose or ruguloso-reticulate (occasionally less conspicuous) main sculpture; long to very long propodeal spines (SPST/CS [0.324, 0.429]), deviating from longitudinal axis of mesosoma by 20–25°; petiolar node in lateral view with a concave frontal profile meeting truncate dorsum in a right angle to an obtuse angle (110–120°) with a narrowly rounded transition, without a conspicuous sharp fronto-dorsal ridge on the petiolar node.

Colonies of this species-complex occur in Southern Europe from Spain to Bulgaria and in Turkey.

Temnothorax nylanderi species complex

Csösz, Heinze, and Mikó (2015): Workers of the Temnothorax nylanderi species-complex can be distinguished from workers of other complexes treated in this revision by the combination of the following salient features: yellow to light brown color (Peloponnese populations occasionally dark brown to black); slightly longer than broad head (CL/CWb [1.100, 1.196]), sculpture of head dorsum dull: with areolate ground sculpture combined with parallel costulate main sculpture; moderately long to long propodeal spines (SPST/CS [0.253, 0.356]), deviating from longitudinal axis of mesosoma by 32–42°; petiolar node in lateral view with a concave frontal profile meeting truncate dorsum in a right angle to an obtuse angle (88–115°) with a narrowly rounded transition, without a conspicuous sharp fronto-dorsal ridge on the petiolar node.

This species-complex is widely distributed in Europe and Turkey.

Temnothorax parvulus species complex

Csösz, Heinze, and Mikó (2015): Workers of the Temnothorax parvulus species-complex can be distinguished from those of other complexes treated in this revision by the combination of the following salient features: dirty yellowish to brownish color; slightly longer than broad head (CL/CWb [1.147, 1.242]), sculpture of head dorsum dull: with uniformly areolate ground sculpture combined with inconspicuous (or the lack of) main sculpture; short to long propodeal spines (SPST/CS [0.205, 0.331]), deviating from longitudinal axis of mesosoma by 38–42°; petiolar node in lateral view with a concave frontal profile meeting truncate dorsum in a right angle to an obtuse angle (100–110°) with a narrowly rounded transition, without a conspicuous sharp fronto-dorsal ridge on the petiolar node.

Members of this species-complex are known to occur in Europe, the Caucasus, and Turkey.

Temnothorax sordidulus species complex

Csösz, Heinze, and Mikó (2015): Workers of the Temnothorax sordidulus species-complex can be distinguished from those of other complexes treated in this revision by the combination of the following salient features: brown to black color; slightly longer than broad head (CL/CWb [1.143, 1.278)], sculpture of head dorsum dull: with areolate ground sculpture combined with conspicuous parallel costulate or irregular reticulate main sculpture; moderately long to long propodeal spines (SPST/CS [0.220, 0.335]), deviating from longitudinal axis of mesosoma by 40–50°; petiolar node in lateral view with a concave frontal profile meeting occasionally indistinct truncate dorsum in an obtuse angle (110–120°) with a narrowly rounded transition, without a conspicuous sharp fronto-dorsal ridge on the petiolar node.

Members of this species-complex are known to occur in Turkey and Crete. The sporadic occurrence of two samples in Greece may be ascribed to anthropochory.

Temnothorax recedens species group

Salata and Borowiec (2015, 2019): 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.

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

Temnothorax rottenbergi species group

Salata and Borowiec (2019): 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. Greece: T. semiruber is present and T. rottenbergi has been recorded but needs confirmation.

Temnothorax tuberum species group

Radchenko (1995c); synopsis by Salata and Borowiec (2019): 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.

+ 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): Temnothorax nigriceps and Temnothorax 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.

Notes for other studies that need to be incorporated here

The following from Gonzalez 2021 needs to be edited and incorporated into what is given above for this group/complex.

Progress has been made establishing species groups and species complexes among the Palearctic Temnothorax.

One of these is the Temnothorax sordidulus species-complex, which was originally conceived by Seifert (2006) and included species characterized by the presence of a distinct mesonotal depression; petiolar shape showing a high node with a small dorsal plane; head, scape and mesosoma more elongated than in the T. nylanderi group (CL/CW 1.08–1.22, SL 0.75–0.81, ML/CS 1.180–1.27 [metrics of Seifert]); sculpture on vertex without the densely arranged, regular and linear carinulae of the T. nylanderi group; spines usually shorter and more erect than the T. nylanderi and the T. parvulus / lichtensteini group; spine axis in lateral view deviating by 40–55 ° from longitudinal axis of mesosoma; coloration of head and gaster always rather dark, varying from medium orange brown to blackish brown, never light yellowish as frequently found in the T. parvulus / lichtensteini group; and mesosoma varying from distinctly lighter or as dark as head and mesosoma. Seifert originally recognized five species in the sordidulus species-complex: T. artvinensis Seifert, 2006, T. melas (Espadaler et al., 1984), T. schoedli Seifert, 2006, T. sordidulus (Müller, 1923) and T. saxonicus (Seifert, 1995). In a revision of the Ponto-Mediterranean T. nylanderi species-group, Csösz, Heinze, and Mikó (2015) gave a definition of the T. sordidulus species-complex that helped separate it from other complexes in the T. nylanderi species-group: “workers of the Temnothorax sordidulus species-complex can be distinguished from those other complexes treated in this revision by the combination of the following salient features: brown to black color; slightly longer than broad head (CL/CWb [1.143, 1.278)], sculpture of head dorsum dull: with areolate ground sculpture combined with conspicuous parallel costulate or irregular reticulate main sculpture; moderately long to long propodeal spines (SPST/CS [0.220, 0.335]), deviating from longitudinal axis of mesosoma by 40–50°; petiolar node in lateral view with a concave frontal profile meeting occasionally indistinct truncate dorsum in an obtuse angle (110–120°) with a narrowly rounded transition, without a conspicuous sharp fronto-dorsal ridge on the petiolar node” (metrics of Csösz, Heinze, and Mikó 2015). Csösz, Heinze, and Mikó (2015) included four species in the T. sordidulus species-complex: T. artvinensis, T. schoedli, T. sordidulus and T. tergestinus(=T. saxonicus according to the authors). In the present study, the T. sordidulus species-complex is characterized by having an elongated head (HL/HWb typically >1.20; metrics defined below), dull surface due to reticulated ground sculpture, distinct metanotal groove, medium to long propodeal spines and high petiole with an anterior concave face and short posterior face.

Beyond Iberia, members of the complex include T. artvinensis and T. schoedli endemic to the Black Sea coast, T. melas endemic to Corsica and the more widespread T. sordidulus and T. tergestinus (Finzi, 1928) (Seifert, 1995;Seifert, 2006; Csösz, Heinze, and Mikó 2015). Temnothorax sordidulus occurs from Turkey to SE France, and T. tergestinus occurs from Bulgaria to SE France (Csősz 2001; Borowiec 2014). In the present paper, four Iberian species match the concept of this complex based on morphometric data; three are previously described endemic species (T. conatensis Galkowski & Lebas, 2016, T. ibericus [Menozzi, 1922] and T. platycephalus [Espadaler, 1997]) and a fourth is a new species from South Iberia described here (T. estel sp. nov.). The taxonomy of the four species is reviewed and an identification key is provided.

References

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