Research Article |
Corresponding author: Sebastian Salata ( sdsalata@gmail.com ) Academic editor: Marek Borowiec
© 2019 Sebastian Salata, Ana Carolina Loss, Celal Karaman, Kadri Kiran, Lech Borowiec.
This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Citation:
Salata S, Loss AC, Karaman C, Kiran K, Borowiec L (2019) Review of the Camponotus kiesenwetteri group (Hymenoptera, Formicidae) in the Aegean with the description of a new species. ZooKeys 899: 85-107. https://doi.org/10.3897/zookeys.899.46933
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Based on recently collected material, the Camponotus kiesenwetteri group is redefined, and its members known from the Aegean region are diagnosed. Camponotus schulzi sp. nov. is described from İzmir Province, Turkey. Camponotus nadimi Tohmé, 1969 syn. nov. is proposed as a junior synonym of Camponotus libanicus André, 1881 and Camponotus kiesenwetteri cyprius Emery, 1920 syn. nov. as a junior synonym of Camponotus kiesenwetteri (Roger, 1859). A key to workers of species of the C. kiesenwetteri group is provided. Niche modeling analyses are used to account for species habitat suitability across the Aegean region.
Aegean Region, carpenter ants, Myrmentoma, new synonym, niche modelling, taxonomy
The genus Camponotus Mayr, 1861 with 1041 valid species and 454 valid subspecies is one of the most speciose within Formicidae. Members of this genus are distributed throughout the world, including the Arctic. However, unquestionably Camponotus reaches the highest diversity in the tropics (
In the two last decades, the majority of studies on Mediterranean Camponotus focused on the Aegean region. Several recent publications show that this region is diverse and rich in taxa endemic to some islands (
The Camponotus kiesenwetteri group comprises several taxa of the subgenus Myrmentoma Forel, 1912 distributed almost exclusively in the Aegean. Only C. libanicus André, 1881 and C. aktaci Karaman, 2013 extend their distribution range to Asia Minor and the Near East. For the first time, the group was defined by
Specimens deposited in the Department of Biodiversity and Evolutionary Taxonomy, University of Wrocław, Poland and the Entomological Museum of Trakya University, Edirne, Turkey were collected between 1991 and 2019 from sites in different parts of the Aegean region. The dominant method was direct sampling (hand collecting). Individual specimens were collected on the ground and tree trunks and from low vegetation. Nests always were located in the soil, most often under trees. All specimens were preserved in 75% EtOH. The study was also supported by material deposited in the Natural History Museum of Crete (Iraklion, Greece), the Muséum d’Historie Naturelle, Genève, and samples collected by Petr Werner (Prague, Czechia). Photos were taken using a Nikon SMZ 1500 stereomicroscope, Nikon D5200 photo camera, and Helicon Focus software. All given label data are in the original spelling, presented in square brackets; a vertical bar (|) separates data on different rows and double vertical bars (||) separate labels. Type specimens’ photographs are available online on AntWeb (https://www.AntWeb.org) and are accessible using the unique CASENT or FOCOL identifying specimen code.
Examined specimens are housed in the following collections:
DBET Department of Biodiversity and Evolutionary Taxonomy, University of Wrocław, Poland;
EMTU Entomological Museum of Trakya University, Edirne, Turkey;
PW Petr Werner collection, Prague, Czechia;
ZMHB Museum für Naturkunde der Humboldt-Universität, Berlin, Germany.
Pilosity inclination degree follows that used in
Measurements: all measurements are given in mm.
HL head length; measured in a 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;
SL scape length; maximum straight-line length of scape;
PW pronotum width; maximum width of pronotum in dorsal view;
PRL propodeum length; measured in lateral view, from metanotal groove to posterior-most point of propodeum;
PRW propodeal width; maximum width of propodeum in dorsal view;
PTH petiole height; the chord of ventral petiolar profile at node level is the reference line perpendicular to which the maximum height of petiole is measured, measured in lateral view;
PTW petiole width; maximum width of the petiolar node in lateral view;
WL Weber’s length; measured as diagonal length from the anterior end of the neck shield to the posterior margin of the propodeal lobe.
Ratios:
CI cephalic index, HL/HW;
SI scape index, SL/HL;
PI petiole index, PTH/PTW.
Habitat suitability for species was estimated by niche modeling using Maxent 3.4.1 (
Camponotus aegaeus Emery, 1915
Camponotus aktaci Karaman, 2013
Camponotus boghossiani Forel, 1911
= Camponotus boghossiani stenoticus Emery, 1915 (= Camponotus kiesenwetteri angustatus Forel, 1889 not Camponotus angustata (Latreille, 1798))
Camponotus kiesenwetteri (Roger, 1859)
= Camponotus kiesenwetteri cyprius Emery, 1920 syn. nov.
Camponotus libanicus André, 1881
= Camponotus libanicus sahlbergi Forel, 1913
= Camponotus nadimi Tohmé, 1969 syn. nov.
Camponotus nitidescens Forel, 1889
Camponotus schulzi sp. nov.
Metanotal groove absent or shallow; propodeal dorsum relatively flat, propodeal declivity deeply concave, posterior protrusions absent or weakly to well developed; body densely punctate, appears dull (only C. nitidescens and C. schulzi have sculpture partially reduced on the lateral sides of mesosoma); the whole body bearing short to long, thick, pale and erect setae, and additional short appressed microsetae; head, mesosoma, and gaster uniformly blackish-brown to black (only C. aktaci has gaster yellowish-brown); polymorphic species.
All known species have similar biological preferences and were most often collected in warm and arid habitats within coniferous forests, especially pine forests. Less frequently they were observed in oak forest, woodland-meadow ecotones, xerothermic meadows, suburban areas with maquis, pastures with shrubs, olive plantations, river bank, orchards, occasionally in rocky gorges with deciduous trees. However, records from open habitats most often were located in the vicinity of trees, especially pine trees. Nests were located in soil, usually sandy, under trees, most often between roots, under small stones, less frequently under big stones. The only observed nest of C. nitidescens was located in a cracked rock wall on a roadside in oak forest under a loose piece of rock. Workers were active all day with the highest activity at dusk. Both major and minor workers were most often found on trunks and branches of coniferous trees, less often on the ground or litter.
Most of the records located in the European mainland came from areas below 700 m a.s.l. and only C. nitidescens is known exclusively from sites located between 1100 and 1700 m a.s.l. However, on Crete, specimens of C. kiesenwetteri were also found in area above 1000 m a.s.l., and the highest record comes from Trocharis peak in Lasithi province (2131 m a.s.l.). Members of the group known from Turkey manifest more alpine preferences. According to label data, the new species Camponotus schulzi was collected at the site located at an altitude of 1150–1500 m. Also C. aktaci is known almost exclusively from montane habitats located above 1000 m a.s.l.
1 | Mesosoma in lateral view forms a regular arch; metanotal groove absent (Figs |
2 |
– | Mesosoma in lateral view with shallow metanotal groove (Figs |
4 |
2 | Legs mostly yellowish to reddish-brown, gaster yellowish-brown. Setation of head, mesosoma, and gaster short and sparse (Figs |
C. aktaci Karaman |
– | Legs and gaster mostly brown to black. Setation of head, mesosoma, and gaster long and dense (Figs |
3 |
3 | Petiolar scale thin, PI > 1.50 (Figs |
C. aegaeus Emery |
– | Petiolar scale thick, PI < 1.42 (Figs |
C. libanicus André |
4 | Posterior margin of propodeum with well developed, lateral dentate protrusions (Figs |
C. kiesenwetteri (Roger) |
– | Posterior margin of propodeum without or with weakly developed, indistinct protrusions (Figs |
5 |
5 | Surface of mesosoma more strongly sculptured, reticulate and granulate with more or less dull background; posterior margin of propodeum sometimes with weakly-developed, indistinct protrusion (Figs |
C. boghossiani Forel |
– | Surface of mesosoma weaker sculptured, especially sides of mesosoma appear more or less shiny; posterior margin of propodeum without protrusions (Figs |
6 |
6 | Base of antennal scape with extension (Fig. |
C. schulzi sp. nov. |
– | Base of antennal scape without extension (Fig. |
C. nitidescens Forel |
Camponotus (Orthonotomyrmex) libanicus var. aegaea
Emery, 1915: 4, figs 1, 2 (s.w.q.m.). Syntype workers, queen, Isola Rodi, Greece (Festa) (
Head, mesosoma, and gaster uniformly blackish-brown to black; metanotal groove absent; propodeum without posterior protrusion; body densely punctate, appears dull; base of scape without extension; whole body bears long, thick, pale, dense and erect setae, and short appressed microsetae; petiolar scale thin (PI > 1.50).
Greece: North Aegean Islands, South Aegean Islands (Dodecanese), Central Macedonia, Eastern Macedonia and Thrace; Turkey: Adana, Afyon, Antalya, Aydın, Balıkesir, Bilecik, Bursa, Çanakkale, Denizli, Diyarbakır, Elazığ, İzmir, Kırklareli, Kütahya, Manisa, Muğla, Sakarya, Uşak, and Yalova. The species was also recorded from North Macedonia (
Almost completely blackish-brown to black body and regularly arched (in lateral view) mesosoma cluster this species with Camponotus libanicus. At first glance both species look extremely similar and the most relevant character distinguishing both taxa is the shape of petiolar scale. Camponotus aegaeus has the scale thin (PI > 1.50) with a feebly convex anterior surface, while in C. libanicus the scale is thick (PI < 1.42) with a strongly convex anterior surface. Both species appear to be vicariant taxa with a more westerly distribution of C. aegaeus and more a easterly distribution of C. libanicus (Figs
Camponotus aktaci Karaman, 2013: 37, figs 1, 7 (w.). Holotype worker, Akcatekir Village, (37°21’N, 34°49’E), 1300 m a.s.l., Adana, Turkey (EMTU) [holotype and paratypes personally investigated].
Head and mesosoma uniformly black, gaster and legs yellowish-brown; metanotal groove absent; propodeum without posterior protrusion; body densely punctate, appears dull; base of scape without extension; whole body bears short, thin, pale, sparse and erect setae, and short appressed microsetae; petiolar scale thick.
Turkey: Adana, Bingöl, Diyarbakır, Elazığ, Malatya, Muğla.
Mostly yellowish-brown gaster and legs and short and sparse setation of head, mesosoma and gaster distinctly separates this species from other members of the Camponotus kiesenwetteri group. Temperature seasonality contributed most to the distribution model. Niche modeling showed highly suitable areas matching species known distribution at Eastern Anatolian deciduous forests but also additional areas in the central Anatolian steppe region, where there are no current occurrence records for the species. However, the westernmost record from Muğla Province is located in an area of low habitat suitability.
Camponotus boghossiani
Forel, 1911: 357 (s.w.). Syntype workers, Lesbos, Greece (
=Camponotus boghossiani var. stenotica Emery, 1915: 7 (=Camponotus kiesenwetteri angustatus Forel, 1889: 261, not Camponotus angustata (Latreille, 1798));
Head, mesosoma, and gaster uniformly black; metanotal groove present, shallow; propodeum without or with indistinct bulge-like protrusions; body densely punctate, appears dull; base of scape without extension; whole body bears long, thick, pale, dense and erect setae, and short appressed microsetae; petiolar scale thick.
Greece: North Aegean Islands, Crete (Heraklion), South Aegean Islands (Cyclades, Dodecanese), Peloponnese (Messinia); Turkey: Antalya, Balıkesir, Çanakkale, Denizli, Karaman, Kütahya, Muğla, and Uşak.
Density of sculpture slightly differs within this species and populations from Peloponnese and Aegean Islands are slightly more sculptured than populations from western Turkey. Camponotus boghossiani is most similar to C. nitidescens and C. schulzi and differs from them in the stronger sculpture of the mesosoma and gaster which, at first glance, appears very dull. While in both relatives the sculpture is slightly diffused and the surface is at least partly shiny. Camponotus kiesenwetteri has a similarly sculptured body surface but differs in having the posterior margin of the propodeum more or less excavate and forming well-developed, lateral dentate protrusions while in C. boghossiani the posterior margin of the propodeum is straight, without protrusions. Isolated specimens of C. kiesenwetteri, with posterior margin of propodeum very shallowly excavate, at first glance look very similar to specimens of C. boghossiani but can be easily be separated by having an antennal scape with a distinct basal extension, while in C. boghossiani the base of the antennal scape has no extension. Precipitation of the wettest quarter was the variable that contributed the most to the distribution model. High suitable areas are indicated especially along the coast of Turkey, Cyprus and Crete.
Formica (Hypoclinea) kiesenwetteri Roger, 1859: 241 (w.). Syntype workers, Greece (ZMHB) [Syntype workers images of Formica (Hypoclinea) kiesenwetteri examined, AntWeb, FOCOL2486 and FOCOL2487, photos by Christiana Klingenberg, available on https://www.AntWeb.org].
=Camponotus kiesenwetteri var. cypria Emery, 1920: 26 (w.) syn. nov. Syntype worker, Cyprus (
Head, mesosoma, and gaster uniformly black; metanotal groove present, shallow; propodeum with distinct dentate protrusions; body densely punctate, appears dull; base of scape with extension; whole body bears long, thick, pale, dense and erect setae, and short appressed microsetae; petiolar scale thick.
Greece: Attica, North Aegean Islands, South Aegean Islands (Cyclades, Dodecanese), Central Greece, Crete (Chania, Heraklion, Lasithi, Rethymno), Ionian Islands, Central Macedonia, Eastern Macedonia and Thrace, Peloponnese; Cyprus; Turkey: Balıkesir, İzmir and Muğla.
The species can be easily separated by the following combination of characters: strongly sculptured body, mesosoma with metanotal groove and posterior margin of propodeum with distinct dentate protrusions, and antennal scape with distinct basal extension. Camponotus nitidescens and C. schulzi both differ in having a partly shiny body, and C. boghossiani differs in having a propodeum without apical protrusions and an antennal scape without basal extension.
Camponotus kiesenwetteri cyprius was described by
Camponotus (Orthonotomyrmex) libanicus
André, 1881: 54, pl. 3, figs 14, 15 (w.). Syntype worker, Lebanon (
=Camponotus (Orthonotomyrmex) libanicus r. sahlbergi Forel, 1913: 435 (s.w.);
=Camponotus (Myrmentoma) nadimi Tohmé, 1969: 6, figs 3, 4 (s.w.) syn. nov. [types unavailable].
Head, mesosoma, and gaster uniformly black; metanotal groove absent; propodeum without posterior protrusion; body densely punctate, appears dull; base of scape without extension; whole body bears long, thick, pale, dense and erect setae, and short appressed microsetae; petiolar scale thick (PI < 1.42).
The species is known from Lebanon (
Camponotus libanicus belongs to the species with mesosoma evenly convex in profile, not interrupted by the metanotal groove. It is very similar to C. aegaeus and differs by having a thick petiolar scale with PI < 1.42, which in C. aegaeus is thinner at PI > 1.50. See also comments in C. aegaeaus.
In the description of C. nadimi from Lebanon,
Camponotus kiesenwetteri nitidescens
Forel, 1889: 260 (w.) Syntype workers, Kefalonia, Greece (
Head, mesosoma, and gaster uniformly brownish-black to black; metanotal groove present, shallow; propodeum without protrusions; body punctate, mesosoma with sculpture reduced and its lateral sides at least partially shiny; base of scape without extension; whole body bears long, thick, pale, dense and erect setae, and short appressed microsetae; petiolar scale thick.
Greece: Ionian Islands (Cephalonia) Peloponnese (Lakonia and Messinia), Western Greece (Aetolia-Acarnania).
Camponotus nitidescens together with C. schulzi are well distinguished from other species of the C. kiesenwetteri group in the partly reduced sculpture of the mesosoma and gaster with, at least, the lateral sides of mesosoma partly shiny. However, the sculpture is never as shiny as in members of related members of the Camponotus lateralis group. Solar radiation was the variable that contributed the most to the distribution model. Although the known distribution is restricted to the western area of the Aegean region, highly suitable areas are indicated in Crete, northeast coast of Turkey, coast of Syria and Lebanon.
Holotype : major worker (CASENT0876000): Turkey |Bozdag Mountain | 38.3277N, 28.1112E || 1150–1500 mH | 10.05.2003 | leg. A. Schulz (DBET); paratypes: 2 major workers, 5 minor workers (CASENT0876001–CASENT0876007): the same data as holotype (DBET, PW, EMTU).
Head, mesosoma, and gaster uniformly black; metanotal groove present, shallow; propodeum without protrusions; body punctate, mesosoma with sculpture reduced and its lateral sides at least partially shiny; base of scape with extension; whole body bears long, thick, pale, dense and erect setae, and short appressed microsetae; petiolar scale thick.
Measurements. Major worker (n = 3): HL: 1.827 (1.78–1.92), HW: 1.72 (1.63–1.82), SL: 1.59 (1.52–1.65), WL: 2.343 (2.27–2.44), PW: 1.22 (1.16–1.27), PRL: 0.657 (0.64–0.68), PRW: 0.43 (0.42–0.44), PTH: 0.40 (0.38–0.41), PTW: 0.293 (0.27–0.32), CI: 1.041 (1.028–1.055), SL/HW: 0.926 (0.889–0.982), PTH/PTW: 1.367 (1.281–1.413); minor worker (n = 5): HL: 1.31 (1.13–1.46), HW: 1.03 (0.94–1.29), SL: 1.297 (1.21–1.41), WL: 1.83 (1.65–2.02), PW: 0.96 (0.86–1.08), PRL: 0.58 (0.52–0.64), PRW: 0.34 (0.32–0.39), PTH: 0.397 (0.35–0.48), PTW: 0.307 (0.27–0.38), CI: 1.192 (1.132–1.241), SI: 1.185 (1.093–1.287), PI: 1.297 (1.263–1.333). Body colouration. Head, mesosoma and petiolus black, gaster from brownish-black to black. Legs brown to black, trochanters as dark as femora (Figs
Named after Andreas Schulz, a German amateur myrmecologist and naturalist, who extensively explored the Aegean region and collected valuable material, including the specimens of C. schulzi sp. nov.
Camponotus schulzi sp. nov. is distinctly polymorphic, the largest major workers 1.5 times longer than the smallest minor workers. Within the C. kiesenwetteri group, together with C. boghossiani, C. kiesenwetteri, and C. nitidescens, it forms a distinct complex characterized by a shallow but distinct metanotal groove. Camponotus boghossiani and C. kiesenwetteri differ from C. schulzi in the matt body with strong and non-reduced sculpture on the whole head, mesosoma, and gaster (Figs
Camponotus schulzi sp. nov. is a member of the subgenus Myrmentoma. Currently, there are 24 species and one subspecies of this subgenus known from the eastern part of the Mediterranean.
The Camponotus lateralis group is the most speciose and represented by 12 species: C. anatolicus Karaman & Aktaç, 2013, C. atricolor (Nylander, 1849), C. candiotes Emery, 1894, C. dalmaticus (Nylander, 1849), C. ebneri Finzi, 1930, C. heidrunvogtae Seifert, 2019, C. hirtus Karaman & Aktaç, 2013, C. honaziensis Karaman & Aktaç, 2013, C. lateralis (Olivier, 1792), C. piceus (Leach, 1825), C. rebeccae Forel, 1913, and C. staryi Pisarski, 1971. In the most recent revision of the group (
The Camponotus fallax group contains six species and one subspecies – C. abrahami Forel, 1913, C. fallax (Nylander, 1856), C. gestroi Emery, 1878, C. gestroi creticus Forel, 1886, C. kurdistanicus Emery, 1898, C. tergestinus Müller, 1921, and C. vogti Forel, 1906. The group is characterized by a small to moderate body size, regularly arched mesosoma sometimes with shallow concavity between mesonotum and propodeum, straight to angular dorsal surface of propodeum, shiny surface of mesosoma and gaster, and short and never dense pubescence hairs on gaster.
The Camponotus kiesenwetteri group as defined here comprises seven species and can be divided into two groups. The first one consists of species lacking a metanotal groove and includes C. aegaeus, C. libanicus, and C. aktaci. The second group is created by taxa with shallow but distinct metanotal groove: C. boghossiani, C. kiesenwetteri, C. nitidescens, and C. schulzi. Most of the members of the kiesenwetteri group have an exclusively Aegean distribution. However, based on the distribution patterns of C. libanicus, and C. aktaci more records of members of this group are expected from the Near East. In fact, all species but C. kiesenwetteri showed large areas of suitable habitats in the east portion of the Aegean region.
We would like to thank Dr. Bernard Landry (Genève, Switzerland) for providing access to the collection of ants preserved in Muséum d’Histoire Naturelle, Genève (