Temnothorax crassispinus

Temnothorax crassispinus is a forest species which typically lives in cavities in fallen twigs and acorns; colonies usually number from a few dozen to about 200 workers (Mitrus, 2019). Zryanin & Zryanina (2007) found it nesting in wood fragments within deciduous and pine forests. Apparently some nests were mixed with Temnothorax nylanderi although specific details of their relation is unknown. In Greece it is uncommon, recorded from Epirus, Macedonia, Peloponnese and Sterea Ellas. Temnothorax crassispinus shares habitat and nest preferences with Temnothorax crasecundus. In Aetolia-Acarnania, a single nest was found in a dry branch of fir inside a fir forest (Borowiec & Salata, 2021).

Identification
Csösz et al. (2015) - A member of the nylanderi species-complex. Temnothorax crassispinus may be confused with other long-spined species treated in this revision: Temnothorax angulinodis, Temnothorax laconicus, Temnothorax lichtensteini and Temnothorax parvulus. Temnothorax angulinodis clearly differs from T. crassispinus by its sharply angulate petiolar node in lateral view (72–82°). In T. crassispinus, the frontal profile and the truncate dorsum of the petiole meet in an obtuse angle (100–115°). The deviation of the propodeal spines from longitudinal mesosomal axis (in lateral view) helps to separate T. crassispinus (32–42°) from T. laconicus and T. lichtensteini (20–25°). Temnothorax parvulus differs from T. crassispinus in the surface sculpturing on the head dorsum. If the samples are dust-covered, other measures can also help: T. crassispinus is considerably larger (CS), has a higher petiolar node (SPST/CS), and a wider head (CL/CWb) than T. parvulus.

Temnothorax crassispinus shares most of its main characteristics, shape and surface sculpturing, with its siblings, Temnothorax crasecundus and Temnothorax nylanderi. The simple propodeal spine length ratio (SPST/CS) helps to separate nest samples of T. crassispinus from those of T. nylanderi without error, but the same character overlaps between nest sample means of T. crassispinus and Temnothorax crasecundus. The shortest discriminant formula (D4) that separates T. crassispinus from T. crasecundus with a classification success rate 95% in single individuals and 97% in nest sample means is D4 = +0.0392*SL -0.0746*SPST +0.0933*SPL -0.0295*SPWI -7.2179.

Distribution
This species is distributed from the Balkans to Central Europe. The distributional area of Temnothorax crassispinus lies between the ranges of its two parapatric relatives, Temnothorax nylanderi in the West and Temnothorax crasecundus in the East.

Distribution based on Regional Taxon Lists
Palaearctic Region: Albania, Austria, Belarus, Bosnia and Herzegovina, Bulgaria, Croatia, Czech Republic, Estonia, Germany, Greece, Hungary, Italy, Latvia, Lithuania, Montenegro, Poland, Republic of Macedonia, Republic of Moldova, Romania, Russian Federation, Serbia, Slovakia, Slovenia, Ukraine.

Biology
Mitrus et al. (2017) - common and widely distributed in Central and Eastern Europe (Czechowski, 2012). This ant species lives in light coniferous and mixed forests (Seifert, 2007; Czechowski et al., 2012). Colonies of the ant are small, typically ranging from a few dozen to several hundreds of workers (Seifert, 2007; Czechowski et al., 2012). However, the density of colonies locally can be high, and the average number of inhabited nest sites could be higher than 2 nests m−2 (Strätz & Heinze, 2004; Białas et al., 2011), and the ant plays a considerable role in seed dispersal (Fokuhl et al., 2012).

Temnothorax crassispinus colonies form nests in cavities in logs plus in acorns and small sticks situated in the litter layer (Seifert, 2007; Czechowski et al., 2012). They can use more than a single nest site in the summer, i.e., exhibit seasonal polydomy. If potentially good nest sites are covered by leaves or overgrown by herbaceous plants or bushes, they are rarely used by ants (S. Mitrus, pers. obs.). Such overshadowing could affect, e.g. temperature and moisture (Ruckli et al., 2013). In forests, where the ant typically lives, different herb species could form dense stands and significantly change environmental conditions at a local scale. In southern Poland, balsams (Impatiens spp.) are a plant species that typically form such stands, and they could overshadow potentially good nest sites for the ant.

The maximum observed foraging distance of workers of T. crassispinus was 162 cm, with an average of 52 cm (Fokuhl et al., 2012).

A field experiment examined nest use in artificial nests placed in and out of areas with native Impatiens noli-tangere and the naturalized introduced species I. parviflora, both forest flowing balsam species. The ant was found to be more frequent in places without balsam cover. Nests within balsam areas were smaller and were more frequently found without queens.

Nomenclature

 *  crassispinus. Leptothorax (Leptothorax) nylanderi var. crassispina Karavaiev, 1926f: 69 (w.q.) UKRAINE.
 * Combination in Temnothorax: Bolton, 2003: 271.
 * Subspecies of nylanderi: Karavaiev, 1927c: 266 (in key).
 * Junior synonym of nylanderi: Karavaiev, 1934: 137; Bolton, 1995b: 237; Radchenko, 1995d: 3.
 * Status as species: Radchenko, 2000: 44; Czechowski, et al. 2002: 50; Seifert, 2007: 239; Czechowski, et al. 2012: 148; Seifert & Csösz, 2015: 40; Csösz, Heinze & Mikó, 2015: 38 (redescription).
 * Senior synonym of slavonicus: Radchenko, 2000: 44; Csösz, Heinze & Mikó, 2015: 38.
 * slavonicus. Leptothorax nylanderi subsp. slavonicus Seifert, 1995: 4, figs. 5, 6, 10, 15, 17, 20 (w.q.) GERMANY.
 * Status as species: Seifert, 1996b: 127, 229.
 * Junior synonym of crassispinus: Radchenko, 2000: 44; Csösz, Heinze & Mikó, 2015: 38.

Worker
Csösz et al. (2015) - Body color: brown; yellow. Body color pattern: mesosoma, antenna and legs, waist and anterior region of 1st gastral tergite lighter than head and posterior region of gaster. Antenna color pattern: clava concolorous funicle. Absolute cephalic size: 544–688 μm (mean = 623, n = 42). Cephalic length vs. Maximum width of head capsule (CL/CWb): 1.100–1.180 (mean = 1.140). Postocular distance vs. cephalic length (PoOc/CL): 0.379–0.403 (mean = 0.390). Postocular sides of cranium contour frontal view orientation: converging posteriorly. Postocular sides of cranium contour frontal view shape: strongly convex. Vertex contour line in frontal view shape: straight. Vertex sculpture: main sculpture parallel costate, ground sculpture areolate. Genae contour from anterior view orientation: converging. Gena contour line in frontal view shape: convex. Gena sculpture: rugoso-reticulate with areolate ground sculpture. Median region of antennal rim vs. frontal carina in frontal view structure: not fully overlapped by frontal carina. Concentric carinae laterally surrounding antennal foramen count: present. Eye length vs. absolute cephalic size (EL/CS): 0.247–0.268 (mean = 0.256). Frontal carina distance vs. absolute cephalic size (FRS/CS): 0.362–0.399 (mean = 0.377). Longitudinal carinae on median region of frons count: present. Longitudinal carinae on medial region of frons shape: not forked. Smooth median region on frons count: absent. Antennomere count: 12. Scape length vs. absolute cephalic size (SL/CS): 0.756–0.811 (mean = 0.784). Facial area of the scape absolute setal angle: 0–15°. External area of the scape absolute setal angle: 30°. Ground sculpture of submedian area of clypeus: smooth. Median carina of clypeus count: present. Lateral carinae of clypeus count: present. Median anatomical line of propodeal spine angle value to Weber length in lateral view: 32–42°. Spine length vs. absolute cephalic size (SPST/CS): 0.288–0.356 (mean = 0.329). Minimum spine distance vs. absolute cephalic size (SPBA/CS): 0.282–0.339 (mean = 0.312). Maximum spine distance vs. absolute cephalic size (SPWI/CS): 0.362–0.421 (mean = 0.389). Apical spine distance vs. absolute cephalic size (SPTI/CS): 0.342–0.397 (mean = 0.366). Maximum mesosoma width vs. absolute cephalic size (MW/CS): 0.509–0.662 (mean = 0.626). Metanotal depression count: present. Metanotal depression shape: shallow. Dorsal region of mesosoma sculpture: rugulose with areolate ground sculpture. Lateral region of pronotum sculpture: areolate ground sculpture, main sculpture forked costate. Mesopleuron sculpture: areolate ground sculpture superimposed by dispersed rugulae. Metapleuron sculpture: areolate ground sculpture superimposed by dispersed rugulae. Frontal profile of petiolar node contour line in lateral view shape: concave. Dorsal profile of petiolar node contour line angle value to frontal profile of petiole contour line in lateral view: 100–115°. Anterodorsal rim of petiole count: absent medially. Dorsal profile of petiolar node contour line in lateral view shape: slightly convex. Dorsal region of petiole sculpture: ground sculpture areolate, main sculpture dispersed rugose; ground sculpture areolate, main sculpture absent. Dorso-caudal petiolar profile contour line in lateral view shape: straight; concave. Dorsal region of postpetiole sculpture: ground sculpture areolate, main sculpture dispersed rugose; ground sculpture areolate, main sculpture absent.

Type Material
Csösz et al. (2015) - Syntype workers of Leptothorax nylanderi var. crassispina Karavajev, 1926: Golossev near Kiev, Leg. Karawajew, “No. 3057 Col. Karawajew” (7##, CASENT0914703) [UKR:Golossev-crassispinus-TYPE].

Paratype workers of Leptothorax nylanderi slavonicus Seifert, 1995: Germany, Kr. Görlitz, Hutberg Schönau-Berzdorf, 19.03.1993. Seifert (4## ) [GER:Hutberg-slavonicus-TYPE].

References based on Global Ant Biodiversity Informatics

 * Babik H. 2011. Ants of Botanical and Zoological Gardens of Warsaw (Poland). Entomologica romanica 16: 53.
 * Borowiec L. 2014. Catalogue of ants of Europe, the Mediterranean Basin and adjacent regions (Hymenoptera: Formicidae). Genus (Wroclaw) 25(1-2): 1-340.
 * Borowiec L., and S. Salata. 2012. Ants of Greece - Checklist, comments and new faunistic data (Hymenoptera: Formicidae). Genus 23(4): 461-563.
 * Borowiec L., and S. Salata. 2017. Ants of the Peloponnese, Greece (Hymenoptera: Formicidae). Polish Journal of Entomology 86: 193-236.
 * Borowiec M. L. 2009. New data on the occurrence of formicoxenine ants (Hymenoptera: Formicidae) in Poland. Wiadomo?ci Entomologiczne 28: 237-246.
 * Borowiec M. L. 2009. Nowe dane o rozmieszczeniu mrówek (Hymenoptera: Formicidae) z plemienia Formicoxenini w Polsce. Wiadomosci Entomologiczne 28: 237-246.
 * Bracko G. 2007. Checklist of the ants of Slovenia (Hymenoptera: Formicidae). Natura Sloveniae 9: 15-24
 * Bracko G., H. C. Wagner, A. Schulz, E. Gioahim, J. Maticic, and A. Tratnik. 2014. New investigation and a revised checklist of the ants (Hymenoptera: Formicidae) of the Republic of Macedonia. North-Western Journal of Zoology 10(1): 10-24.
 * Bracko, G. 2006. Review of the ant fauna (Hymenoptera:Formicidae) of Croatia. Acta Entomologica Slovenica 14(2): 131-156.
 * Bracko, G. "New species for the ant fauna of Slovenia (Hymenoptera:Formicidae)." Natura Sloneniae 5 (1) (2003): 17-25.
 * Bracko, G. "Review of the ant fauna (Hymenoptera: Formicidae) of Croatia." Acta Entomologica Slovenica Vol 14 st (2006): 131-156.
 * Bujan J., A. Brigic, Z. Sedlar, and R. Sostaric. 2015. Progressive vegetation succession of fen habitats promotes the lack of habitat specialist ants. Insectes Sociaux 62: 415-422.
 * Cagniant, H. Les peuplements de fourmis des forêts algériennes: écologie, biocénotique, essai biologique. Universite de Toulouse, 1973.
 * Csosz S., B. Marko, K. Kiss, A. Tartally, and L. Galle. 2002. The ant fauna of the Ferto-Hansag National Park (Hymenoptera: Formicidae). In: Mahunka, S. (Ed.): The fauna of the Fert?-Hanság National Park. Hungarian Natural History Museum, Budapest, pp. 617-629.
 * Csősz S, Heinze J, and I. Mikó. 2015. Taxonomic synopsis of the Ponto-Mediterranean ants of Temnothorax nylanderi species-group. PLoS ONE 10(11): e0140000. doi:10.1371/journal.pone.0140000
 * Csősz S., B. Markó, and L. Gallé. 2001. Ants (Hymenoptera: Formicidae) of Stana Valley (Romania): Evaluation of the effectiveness of a myrmecological survey.  Entomologica Romanica 6 : 121-126.
 * Csősz S., B. Markó, and L. Gallé. 2011. The myrmecofauna (Hymenoptera: Formicidae) of Hungary: an updated checklist. North-Western Journal of Zoology 7: 55-62.
 * Czechowski W., A. Radchenko, W. Czechowska and K. Vepsäläinen. 2012. The ants of Poland with reference to the myrmecofauna of Europe. Fauna Poloniae 4. Warsaw: Natura Optima Dux Foundation, 1-496 pp
 * Dubovikoff D. A., and Z. M. Yusupov. 2018. Family Formicidae - Ants. In Belokobylskij S. A. and A. S. Lelej: Annotated catalogue of the Hymenoptera of Russia. Proceedingss of the Zoological Institute of the Russian Academy of Sciences 6: 197-210.
 * Dvorak, L., P. BOGUSCH, I. MALENOVSKÝ, P. BEZDÌÈKA, K. BEZDÌÈKOVÁ, K. HOLÝ, P. LIKA, J. MACEK, L. ROLLER, M. RÍHA et al. "Hymenoptera of Hády Hill, near the city of Brno (Czech Republic), collected during the Third Czech-Slovak Hymenoptera meeting." Acta Musei Moraviae, Scientiae biologicae (Brno) 93 (2008): 53-92.
 * Fiedler, K., F. Kuhlmann, B. C. Schlick-Steiner, F. M. Steiner and G. Gebauer. 2007. Stable N-isotope signatures of central European ants  assessing positions in a trophic gradient. Insectes Sociaux 54(4):393-402.
 * Foitzik S., and Heinze, J. 1999. Non-random size differences between sympatric species of the ant genus Leptothorax (Hymenoptera: Formicidae). Entomologia Generalis 24: 65-74.
 * Gallé L., B. Markó, K. Kiss, E. Kovács, H. Dürgő, K. Kőváry, and S. Csősz. 2005. Ant fauna of Tisza river basin (Hymenoptera: Formicidae).  In: Gallé, L. (szerk.): Vegetation and Fauna of Tisza River Basin I. Tiscia Monograph Series 7; Szeged, pp. 149-197.
 * Ihnatiuk O. A., and S. V. Stukalyuk. 2015. Degradation changes in the structure of multispecies associations of ants in urbanized areas. Russian Journal of Ecology 46(1): 109–115.
 * Karaman C., K. Kiran, and V. Aksoy. 2014. New records of the genus Strumigenys Smith, 1860 (Hymenoptera, Formicidae) from Black Sea region of Turkey. Trakya University Journal of Natural Sciences, 15(2): 59-63.
 * Karaman M. G. 2011. A catalogue of the ants (Hymenoptera, Formicidae) of Montenegro. Podgorica: Catalogues 3, Volume 2, Montenegrin Academy of Sciences and Arts, 140 pp.
 * Kiran K., and C. Karaman. 2012. First annotated checklist of the ant fauna of Turkey (Hymenoptera: Formicidae). Zootaxa 3548: 1-38.
 * Kiss K., and K. Fetyko. 2008. Notes about the ant fauna (Hymenoptera: Formicidae) of the Tur valley. In. Sike T., Márk-Nagy J. (eds), The flora and fauna of the Tur river natural reserve, Biharean Biologist, II, Oradea, pp.71-76.
 * Korlevic, A. "Prilozi fauni hrvatskih opnokrilaca." Glasn. Hrv. Narav. Dr. 5 (1890): 189-250.
 * Lapeva-Gjonova A., and K. Kiran. 2012. Ant fauna (Hymenoptera, Formicidae) of Strandzha (Istranca) Mountain and adjacent Black Sea coast. North-western journal of Zoology 8(1): 72-84.
 * Lorinczi G. 2011. Density and spatial pattern of nests in sub-Mediterranean ground-dwelling ant communities (Hymenoptera: Formicidae). Community Ecology 12(1): 51-57.
 * Majzlan O., and P. Devan. 2009. Selected insect groups (Hymenoptera, Neuroptera, Mecoptera, Raphidioptera) of the Rokoš Massif (Strážovské vrchy Mts.). Rosalia (Nitra), 20, p. 63–70.
 * Marko B. 1999. New ant taxa (Hymenoptera: Formicidae) in the Romanian fauna. Entomol. Rom. 4: 95-98.
 * Marko B. 2008. Ants (Hymenoptera: Formicidae) of the Sand Dunes of Foieni Protected Area and Its Surroundings (Satu Mare County, Romania), and a New Species for the Romanian Fauna. Acta Scientiarum Transylvanica 16(3): 87-99.
 * Markó B., B. Sipos, S. Csősz, K. Kiss, I. Boros, and L. Gallé. 2006. A comprehensive list of the ants of Romania (Hymenoptera: Formicidae). Myrmecologische Nachrichten 9: 65-76.
 * Markó B., and S. Csősz. 2002. Die europäischen Ameisenarten (Hymenoptera: Formicidae) des Hermannstädter (Sibiu, Rumänien) Naturkundemuseums I.: Unterfamilien Ponerinae, Myrmicinae und Dolichoderinae. Annales Historico-Naturales Musei Nationalis Hungarici 94: 109-121.
 * Mitrus S. 2015. Unexpected high overwintering survival rate of the cavity-dwelling ant Temnothorax crassispinus (Hymenoptera: Formicidae). Entomologica Fennica 26: 194-200.
 * Müller, G. "Le formiche della Venezia Guilia e della Dalmazia." Bollettino della Società Adriatica di Scienze Naturali in Trieste 28 (1923): 11-180.
 * Nemet E., Z. Czekes, I. Tausan, and B. Marko. 2012. Contribution to the knowledge of the myrmecofauna of the Cefa Nature Park (North-Western Romania). Acta Scientiarum Transylvanica 20(1): 61-72.
 * Pusch K., J. Heinze, and S. Foitzik. 2006. The influence of hybridization on colony structure in the ant species Temnothorax nylanderi and T. crassispinus. Insectes Sociaux 53: 439-445.
 * Pusch, K., J. Heinze and S. Foitzik. 2006. The influence of hybridization on colony structure in the ant species Temnothorax nylanderi and T. crassispinus. Insectes Sociaux 53:439-445
 * Rzeszowski K., H. Babik, W. Czechowski, and B. Marko. 2013. Ants (Hymenoptera: Formicidae) of Chelmowa Gora in the Swietokrzyski National Park. Fragmenta Faunistica 56(1): 1-15.
 * Salata S. 2014. Ants (Hymenoptera: Formicidae) of the National Park of the Stołowe Mts. Przyroda Sudetow 17: 161-172.
 * Salata S. 2014. Mrówki (Hymenoptera: Formicidae) Parku Narodowego Gór Sto?owych. PRZYRODA SUDETÓW t. 17: 161-172.
 * Salata S., and L Borowiec. 2017. Species of Tetramorium semilaeve complex from Balkans and western Turkey, with description of two new species of (Hymenoptera: Formicidae: Myrmicinae). Annales Zoologici (Warsaw) 62:279–313.
 * Salata S., and L. Borowiec. 2019. Preliminary division of not socially parasitic Greek Temnothorax Mayr, 1861 (Hymenoptera, Formicidae) with a description of three new species. ZooKeys 877: 81-131.
 * Santschi, F. 1934. Fourmis du Sahara central. Memoires de la Société d'Histoire Naturelle d'Afrique du Nord 4: 165-177.
 * Santschi, F. "Formicides nouveaux de l'Afrique Mineure (4e note)." Bulletin de la Société d' Histoire naturelle de l' Afrique du Nord 2 (1911): 11-14.
 * Saure C. 2005. Rote Liste und Gesamtartenliste der Bienen und Wespen (Hymenoptera part.) von Berlin mit Angaben zu den Ameisen. In: DER LANDESBEAUFTRAGTE FÜR NATURSCHUTZ UND LANDSCHAFTSPFLEGE / SENATSVERWALTUNG FÜR STADTENTWICKLUNG (Hrsg.): Rote Listen der gefährdeten Pflanzen und Tiere von Berlin. CD-ROM.
 * Schlick-Steiner B. C., and F. M. Steiner. 1999. Faunistisch-ökologische Untersuchungen an den freilebenden Ameisen (Hymenoptera: Formicidae) Wiens. Myrmecologische Nachrichten 3: 9-53.
 * Schlick-Steiner, B.C., F.M. Steiner, K. Moder, A. Bruckner, K. Fiedler and E. Christian. 2006. Assessing ant assemblages: pitfall trapping versus nest counting (Hymenoptera, Formicidae). Insectes Sociaux 53:274-281
 * Seifert B. 1995. Two new Central European subspecies of Leptothorax nylanderi (Förster, 1850) and Leptothorax sordidulus Müller, 1923 (Hymenoptera: Formicidae). Abhandlungen und Berichte des Naturkundemuseums Görlitz 68(7): 1-18.
 * Seifert B., and S. Csösz. 2015. Temnothorax crasecundussp. n.  a cryptic Eurocaucasian ant species (Hymenoptera, Formicidae) discovered by Nest Centroid Clustering. ZooKeys 479: 3764.
 * Slipinski P., B. Marko, K. Rzeszowski, H. Babik, and W. Czechowski. 2014. Lasius fuliginosus(Hymenoptera: Formicidae) shapes local ant assemblages. North-Western Journal of Zoology 10(2): 404-412.
 * Somogyi A. A., G. Lorinczi, J. Kovacs, and I. E. Maak. 2017. Structure of ant assemblages in planted poplar (Populus alba) forests and the effect of the common milkweek (Asclepias syriaca). Acta Zoologica Academiae Scientiarum Hungaricae 63(4): 443–457.
 * Steiner F. M., S. Schödl, and B. C. Schlick-Steiner. 2002. Liste der Ameisen Österreichs (Hymenoptera: Formicidae), Stand Oktober 2002. Beiträge zur Entomofaunistik 3: 17-25.
 * Strätz, M. and J. Heinze. 2004. Colony structure and sex allocation ratios in the antTemnothorax crassispinus. Insectes Sociaux 51(4):372-377.
 * Stukalyuk S. V. 2015. Structure of the ant assemblages (Hymenoptera, Formicidae) in the broad-leaved forests of Kiev. Entomological Review 95(3): 370–387.
 * Stukalyuk S. V. 2017. Stratification of the ant species (Hymenoptera, Formicidae) in the urban broadleaf woodlands of the city of Kiev. Entomological Review 97(3): 320-343.
 * Tartally A. 2009. Data on the ant fauna of Gy?r?f? (SW-Hungary) (Hymenoptera: Formicidae). Natura Somogyiensis 13: 155-158.
 * Tausan I. 2014. Temnothorax parvulus (Schenck, 1852)(Hymenoptera: Formicidae) in Romania. Brukenthal, Acta Musei 9: 511-514.
 * Tausan I., J. Dauber, M. R. Tricia, and B. Marko. 2017. Succession in ant communities (Hymenoptera: Formicidae) in deciduous forest clear-cuts – an Eastern European case study. European Journal of Entomology 114: 92-100.
 * Tausan I., M. M. Jerpel, I. R. Puscasu, C. Sadeanu, R. E. Brutatu, L. A. Radutiu, and V. Giurescu. 2012. Ant fauna (Hymenoptera: Formicidae) of Sibiu County (Transylvania, Romania). Brukenthal. Acta Musei 7(3): 499-520.
 * Tausan I., and B. Marko. 2009. Comparative analysis of ant communities (Hymenoptera: Formicidae) in the surroundings of Sibiu (Romania). Brukenthal. Acta Musei 4(3): 635-644.
 * Ticha K., and P. Stys. 2002. Monogyny in Leptothorax slavonicus (Hymenoptera: Formicidae). Acta Soc. Zool. Bohem. 66: 151-160.
 * Vogrin, V. "Prilog fauni Hymenoptera - Aculeata Jugoslavije." Zast. Bilja 31(suppl.) (1955): 1-74.
 * Wiezick M. 2008. Ant communities (Hymenoptera: Formicidae) of forest-steppe habitats at southern and South-Western slopes of Plesivecka a Planina plateau at Slovensky Kras Karst. Natura Carpatica 49: 85-94.
 * Wiezik M., A. Wieziková, and L. Ma?áková. 2010. Ant assemblages (Hymenoptera: Formicidae) of typical habitats at Boky National Nature Reserve. Naturae Tutela 14(1): 91-98.
 * Wlodarczyk T. 2010. Ant species composition in relation to forest associations in Szczecin Landscape Park. Polish Journal of Entomology 79: 91-99.