Cardiocondyla, Emery, 1869

Seifert, Bernhard, 2023, A revision of the Palaearctic species of the ant genus Cardiocondyla Emery 1869 (Hymenoptera: Formicidae), Zootaxa 5274 (1), pp. 1-64 : 8-11

Key to the species with outdoor occurrence in the Palaearctic

In order to comprise all 34 species with outdoor occurrence in the Palaearctic, the key and the supplementary information SI1 and SI2 additionally included nine species of species groups with other zoogeographic origin. These species either extend their range into the southern Palaearctic or are globally dispersing tramp species potentially occurring in each locality with sufficiently warm climate. The complete taxonomic names of these added species are Cardiocondyla minutior Forel 1899 , C. tjibodana Karavajev 1935 and C. goa Seifert 2003 (all three belonging to the C. minutior species group), C. emeryi Forel 1881 (belonging to the C. emeryi species group), C. fajumensis Forel 1913 and C. unicalis Seifert 2003 (both belonging to the C. shuckardi species group), C. wroughtonii (Forel 1890) and C. obscurior Wheeler 1929 (both belonging to the C. wroughtonii species group) and C. mauritanica Forel 1890 (belonging to the C. nuda species group).

The key presented below almost thoroughly used dichotomous decision schedules for cases requiring combinations of multiple characters and in the same time it aspired to achieve a low error rate inevitably resulting in tedious procedures. Lamenting does not help here and those who think that simple answers can be given for difficult problems bark up the wrong tree. The finally best approach is here using a software providing linear discriminant functions (LDA), measuring the full set of morphometric characters and running the investigated samples as wild-cards within the data space of the hypothesis-providing supplementary file SI2. It is convenient to run the analysis with primary (raw) measurements instead with ratios. A good check of the result is then comparing with the images that are given for any species in four aspects and to consider geographic distribution. The low error rates reported in the key cannot be achieved for such tiny ants without high-resolution optical systems, a good measuring stage and a careful realization of character definitions.

1a Anterior postpetiolar sternite laterally more prominent than in median part—as result an imagined cross section shows a deeply concave ventral margin of the sternite. The lateral protrusions appear in frontolateral view as dents ( Fig. 113 ). C. wroughtonii group............................................................................................... 2

1b Anterior postpetiolar sternite in median part more prominent or at equal level as the lateral surface—as result an imagined cross section shows a convex or straight ventral margin of the sternite................................................ 3

2a With all measurements in mm, discriminant 34.21* CW-59.33 *PoOc-177.83* EYE +72.15* SL-113.56 * FRS-106.46 *SP+140.69*PpW-14.15 <0 [error 0% in 37 nest sample means of two workers measured]. 1 st gaster segment in a majority of samples with absent or weakly developed blackish pigmentation. Worldwide tramp species. Outdoor occurrence confirmed for the Middle East. Figs. 105–108 ...................................................... wroughtonii

2b Discriminant> 0 [0% error in 65 nest sample means of two workers]. 1 st gaster segment in a majority of samples with strongly developed blackish pigmentation. Worldwide tramp species. Outdoor occurrence confirmed for southern Spain and the Middle East. Figs.109–113 ............................................................................. obscurior

3a True foveolae on vertex dorsal of the eyes completely absent. Instead the bases of pubescence hairs are placed in the center of flat tubercles or flat pits of only 4–10 µm diameter, giving a finely punctate surface appearance at lower magnifications ( Figs. 89 , 97 ). Postocular distance large, PoOc/CL 0.416 –0.467. Frons usually very wide, FRS/CS> 0.288 (except C. tibetana , here 0.280 ± 0.006). Propodeal spines reduced to blunt dents. Metanotal depression deep. C. stambuloffii group.............. 4

3b Bases of pubescence hairs on vertex dorsal of the eyes placed in the center of foveolae or cuticular meshes of> 10 µm diameter ( Figs. 9 , 25 , 117 )—if foveolae reduced ( verdensis n. sp.), than PoOc/CL <0.394. Frons narrower, FRS/CS <0.288. Metanotal depression varying from nearly absent to deep. Propodeal spines variable......................................... 8

4a Eye larger, EYE/CS>0.235. Frons narrower, FRS/CS <0.295. Waist segments lower, PeH/CS <0.343, PpH/CS <0.283. Southern and northern margin of Tarim Basin. Figs. 101–104 ............................................ tibetana

4b EYE/CS <0.235, FRS/CS> 0.295, PeH/CS>0.343, PpH/CS <0.283............................................ 5

5a Posterior 40% of vertex surface completely smooth except for tiny pits of 4–5 µm around the bases of pubescence hairs just visible with high-resolution stereomicroscopy. Pubescence longer. With all measurements in mm, discriminant 334.68*PLG-45.456*CW+88.43*PoOc-66.63*EYE -3.60> 2.3. South Kazakhstan. Figs. 98–100 ........................... gibbosa

5b Posterior part of vertex not completely smooth, at least with weak rugulae and small tubercles around the bases of pubescence hairs. Discriminant <2.3............................................................................... 6

6a Westpalaearctic, known range extending east to the western shores of Caspian Sea. With all measurements in mm, discriminant 44.95*CW+54.68*PoOc+31.14*SL-130.25*SPBA-79.81*PpW +40.84*PeH+90.89*PpH -28.33 <0.4 [0% error in 82 individuals]. Figs. 86–89 ...................................................................... stambuloffii

6b Central Palaearctic, known range extending from western coast of Caspian Sea east to Mongolia. Discriminant> 0.4 [0% error in 59 individuals]..................................................................................... 7

7a With all measurements in mm, discriminant 120.6*PoOc-29.53*SL+109.54*PeW-109.42*PeH-157.97*PLG-7.70 <0.7 [0% error in 37 individuals]. Figs. 94–97 ............................................................ rolandi n. sp.

7b Discriminant> 0.7 [0% error in 22 individuals]. Figs. 90–93 ......................................... koshewnikovi

8a Propodeal spines reduced to obtusely angled corners ( Fig. 115 ). Postpetiole rather narrow, PpW/CS <0.46, in dorsal view never suggestively hexagonal. With all measurements in mm, discriminant 48.06*PoOc+27.66*SL-56.96*PpW+119.07*MGr-13.79> 0 [error 0% in 17 individuals].......................................................................... 9

8b Propodeal spines not reduced to obtusely angled corners. If spines occasionally reduced to blunt corners, then EYE/CS> 0.221 and PpW/CS>0.450. Discriminant <0 [error 0% in 1140 individuals].......................................... 10

9a Head elongated CL/CW> 1.16 ( Fig. 114 ). Dorsum of promesonotum and lateral pronotum with more developed microsculpture ( Figs. 115, 116 ). Head sculpture stronger, foveolae with regular margin ( Fig. 117 ). Egypt and Saudi Arabia.......................................................................................................... fajumensis

9b Head very short CL/CW <1.16 ( Fig. 114 ). Dorsum of promesonotum and lateral pronotum nearly glabrous ( Figs. 119, 120 ). Head sculpture weaker, foveolae sometimes with irregular margin ( Fig. 121 ). Iran............................. unicalis

10a Frons very narrow, FRS/CS 0.196 —0.228 ( Fig. 122 ), postpetiole with a bulging sternite and high ( Fig.123 .), PpH/CS 0.306 –0.358. With all measurements in mm, discriminant 128.52*FRS+42.56*PeW-143.94*PpH+1.616 <0 [error 0% in 145 specimens]. Confirmed outdoor occurrence in S Iberia, N Africa and Middle East. Figs. 122–125 .................. emeryi

10b Frons wider and postpetiole without a strongly bulging sternite. Discriminant> 0 [error 0% in 863 specimens].......... 11

11a Small, CW <420 mm. Metanotal depression very weak or absent, spines well developed, postpetiole much lower than petiole and with a completely flat sternite ( Fig. 127 ). Vertex with deeply impressed foveolae ( Fig.126 ). All surfaces of head, mesosoma and waist matt ( Fig. 128 ). Outdoor occurrence confirmed for Egypt and Saudi Arabia........... minutior , tjibodana & goa These species are inseparable by single or few characters. It is recommeded to measure CL, CW, SL, EYE, FRS, SP, PpW, PeH, PpH and PLG and then running the test specimens as wild-cards in a LDA against the data given in supplementary information SI2. The total classification error in 210 worker individuals in SI2 is 2.4%. Users who refrain from doing this time consuming and challenging separation are recommended to name this collective cluster........................... minutior aggr.

11b Larger, CW> 420 mm. Minute specimens of C. mauritanica may match the character combination of 11a but differ by shorter spines and a thinner petiolar peduncle (compare Figs. 127 and 130 )............................................ 12

12a Outlines of postpetiole in dorsal view approximately hexagonal ( Fig. 131 ). Metanotal depression shallow (MGr/CS 2.14±0.52). Propodeal spines developed as short dents ( Fig. 130 ). Gastral pubescence dense (sqPDG 3.72 ± 0.28). Postocular distance large (PoOc/CL 0.447 ± 0.008, Fig. 129 ). Microsculpture on vertex strongly reticulate ( Fig. 132 ). With MGr, PoOc and SP in mm and sqPDG in µm, discriminant 136.0*MGr-25.0*PoOc*73.99*SP+1.81*sqPDG-7.2322 <0 [error 0% in 151 specimens]. Mediterranean and South Temperate zone of the Palaearctic.......................................... mauritanica .

12b Outlines of postpetiole in dorsal view not approximately hexagonal. Metanotal depression usually deeper, propodeal spines more developed, gastral pubescence often more dilute, postocular distance usually smaller. Microsculpture on vertex different. Discriminant> 0 [error 0.3% in 619 specimens]............................................................ 13

13a Postocular distance extremely small (PoOc/CL <0.340). Petiole height and postpetiole width very low. With all measurements in mm, discriminant 30.96*PoOc-38.98*CL+17.12*PeH+26.77*PpW+ 7.196 <0 [error in 4 specimens 0%]. Kuweit. Figs. 62–65 ....................................................................................... opistopsis

13b Postocular distance larger (PoOc/CL 0.342 –0.469]. Petiole height and postpetiole width higher. Discriminant>0 [error 0% in 612 specimens]...................................................................................... 14

14a Head surface between inner eye margin and paramedian vertex strongly microrugulose ( Fig. 69 ). Dorsal promesonotum longitudinally microrugulose ( Fig. 68 ).All surfaces of head, mesosoma and petiole matt due to more developed microsculpture. Dorsal profile of mesonotum straight, contrasting the clearly convex dorsal profile of propodeum, metanotal depression feebly suggested ( Fig. 67 ). With all measurements in mm, discriminant 95.88*EYE-164.99*MGr-12.035> 0. Yemen. Figs. 66–69 ............................................................................................... rugulosa

14b Body surfaces less microrugulose, often shiny. Dorsal profile of mesosoma different. Discriminant <0................ 15

15a Microsculpture of head with irregular foveolar margins ( Fig. 73 ). Additionally differing from members of the elegans and ulianini group by larger eye (EYE/CS 0.265 –0.274] and from members of the batesii group by longer tergite pubescence (PLG/ CS 6.84–6.94). Afghanistan. Figs. 70–73 ............................................................ kushanica

15b Foveolae on vertex not with irregular margins. Character combination different................................... 16

16a Foveolae on vertex extremely small (dFOV 7.0–9.1 µm). Tergite pubescence very dilute (sqPDG 5.88–5.92). Scape and head much elongated (SL/CS 0.854 –0.867, CL/CW 1.238 –1.256). All body surfaces very shiny. Island endemic of Cape Verde, Figs.82–85 ............................................................................... verdense n. sp.

16b Foveolae much larger (dFOV 12–21 µm). Tergite pubescence often more dense. Scape and head usually shorter......... 17

17a With sqPDG in µm and all other measurements in mm, discriminant 201.57*PLG-0.32*sqPDG-40.86*CL-10.32*PoOc+70.74 *SL+23.72*SPBA-24.27*SP+26.48*PeW+12.96*PpW-73.42*PeH-8.61> 0 [error 0% in 177 individuals]. C. elegans group................................................................................................... 18

17b Discriminant <0 [error 0% in 322 individuals]. C. batesii & C. ulianini group.................................... 21

18a Occurring from Iberia accross S France to Italy. Contact zone with dalmatica in the border region of NE Italy and Slovenia. With all measurements in mm, discriminant 56.7*SP+156*PeW-61.9*PoOc-126.2*PeH-48.2*PpW+126.6*MGr+17.86>0 [error 0% in 81 two-specimen nest samples]. Figs. 6–9 ................................................... elegans

18b Occurring farther east. If found in NE Italy or Slovenia, discriminant <0........................................ 19

19a Combination of larger foveloar diameter on vertex, lower postpetiolar height, longer propodeal spines and higher pubescence distance on gaster tergites. With dFOV, sqPDG in µm and SP, PpH in mm, discriminant 1.623*sqPDG+0.718*dFOV+96.14*SP -62.63*PpH-15.233> 2 [error 0% in 13 specimens]. S Georgia, NE Turkey, Iran, Afghanistan. Figs. 18–21 ...... brachyceps

19b Foveolar diameter smaller, postpetiolar height larger, spines shorter, pubescence distance smaller. Discriminant <2 [error 1.4% in 72 specimens]..................................................................................... 20

20a Bases of propodeal spines more approached, petiole narrower and higher, surface of propodeum glabrous, propodeal spines very acute, petiole high with a very steep anterior profile of the node. With all measurements in mm, discriminant 163.8*PeH- 29.1*PeW-134.63*SPBA-5.68>2 [error 0% in 6 specimens]. Asia Minor. Figs. 14–17 ............... dalmaticoides n. sp.

20b Bases of propodeal spines more distant, petiole wider and lower, surface of propodeum less glabrous, propodeal spines less acute, petiole lower with a less steep anterior profile of the node. Discriminant <2 [error 0% in 59 specimens]. From N Italy across the whole Balkans, Cyprus and Asia Minor east to the Iran. Northern range border in Austria, Slovakia and Hungary at 48°N. Figs. 10–13 .............................................................................. dalmatica

21a With all measurements in mm, discriminant 32.44*CW-103.36*CL+134.03*PoOc-44.05*EYE+23.08*SL-41.77*SPBA+48.21*PpW-0.135 <0 [error 0% in 98 individuals]. Species of the C. batesii group..................... 22

21b Discriminant> 0 [error 0.4% in 241 individuals]. Species of the C. ulianini group................................. 25

22a Frontal carinae notably diverging frontad, FL/FR>1.109. Postocular distance large, PoOc/CL>0.408. Metanotal depression very shallow, MGr <2.5 %. Jordan. Figs. 74–77 ...................................................... tenuifrons

22b Frontal carinae less diverging, FL/FR <1.110. Postocular distance smaller, PoOc/CL <0.409. Metanotal depression deeper, MGr usually> 2.5 %................................................................................. 23

23a Petiole massive, ratio PeW /PpW> 0.57. With all measurements in mm, discriminant 152.578*PeW-69.53*PpW+2.092*SL+3 0.63*PoOc-10.57> 2.5 [error 0% in 4 individuals]. Only known from two sites in Asia Minor. Figs. 58–61 ....... semirubra

23b Petiole less massive, ratio PeW/PpW in 97% of individuals <0.57. Discriminant <2.5 [error 0% in 132 individuals. Figs. 54–57............................................................................................. 24

24a With all measurements in mm, discriminant 63.22*CL-63.18*PoOc+55.89*SL+64.64*PeW-184.93*PeH-93.82*PpH-11.19 <0 [error 0% in 37 specimens]. Morocco, Tunisia, Algeria, Spain. Figs. 54–57 .................................. batesii

24b Discriminant> 0 [error 0% in 95 specimens]. Portugal, Morocco, Algeria, Tunisia, Egypt, Cyprus, Turkey, Middle East, Iran eastwards to 56°E. Color polymorphic. Figs. 78–81 ....................................................... nigra

25a Frons very narrow, FRS/CS 0.225; propodeal spines acute but very short, SP/CS 0.071; postocular distance small, PoOc/CL 0.403; postpetiole wide PpW/CS 0.552. With all measurements in mm, discriminant 160.18*FRS+41.17*PoOc-45.99*PpW- 15.23 <0. Israel. Figs. 34–37 ..................................................................... gallilaeica

25b Frons wider; propodeal spines longer; postocular distance larger; postpetiole often narrower. Discriminant> 0 [error 0% in 240 specimens]......................................................................................... 26

26a Combination of large propodeal spine base distance, small eye, large postocular distance, broad and high postpetiole. With all measurements in mm, discriminant 44.74*PoOc-167.14*EYE+161.96*SPBA-56.07*PpW+31.77*PpH+0.543>0 [error 0% in 179 specimens]...................................................................................... 27

26b Propodeal spine base distance smaller, eye larger, postocular distance smaller, postpetiole narrower and lower. Discriminant <0 [error 0% in 61 specimens]........................................................................... 29

27a With sqPDG in µm and all other measurements in mm, discriminant 163.54*PoOc+0.755*sqPDG-57.89* CL-58.26 *CW+197. 78* EYE +47.39* FRS-54.85 *SP+55.08*PeW-109.15*PeH+53.45*PpH-178.59* PLG-4.19 > 0.5 [error 0% in 84 specimens]. S Balkans and Asia Minor. Figs. 42–45 ............................................................... bulgarica

27b Discriminant <0.5 [error 0% in 95 specimens]............................................................. 28

28a Microsetae on clypeus shorter ( Fig. 46 ), pubescence on all body parts shorter, foveolae on vertex less densely packed ( Fig.49 ), postpetiole wider, petiole node in lateral view more symmetric ( Fig.47 ), dorsum of mesosoma and waist shiny and mesosoma frequently darker. With dFOV in µm and all other measurements in mm, discriminant 0.965*dFOV+164.79*EYE-38.64*PpW+123.49* PLG +77.67*MGr-32.49 <0 [error 2.9% in 69 specimens]. SE Balkans, Asia Minor, Israel, Caucasus, Caspian region , Iran, Afghanistan, Kyrgistan. Figs, 46–49.............................................. sahlbergi

28b Microsetae on clypeus longer ( Fig. 50 ), pubescence on all body parts longer, foveolae on vertex more densely packed ( Fig. 53 ), postpetiole narrower, petiole node in lateral view more asymmetric ( Fig. 51 ), dorsum of mesosoma and waist less shiny and mesosoma frequently lighter. Discriminant> 0 [error 7.7% in 26 specimens]. Georgia, Iran. Figs 50–53 ........... persiana

29a Vertex with rather large, well-demarcated, bicoronate and densely-packed foveolae and in anteromedian part with well-developed longitudinal sculpture( Figs. 38, 41 ). Pubescence on gaster tergites more dense. With dFOV, sqPDG in µm and SPBA in mm, discriminant 2.559*sqPDG-0.472*dFov+15.117*SPBA-4.857 <0 [error 0% in 10 specimens]. Eqypt , Israel. Figs. 38–41 ........................................................................................ israelica

29b Vertex with smaller, less demarcated and more spaced foveolae and with very weak or absent longitudinal microsculpture. Pubescence on gaster tergites less dense. Discriminant> 0 [error 0% in 51 specimens]............................. 30

30a Postpetiole and frons wider, pubescence hairs on gaster tergites shorter. With all measurements in mm, discriminant 93.21* FRS +27.62*PpW-406.34* PLG-6.246 > 0 [error 0% in 44 specimens]. Continuously distributed from the S Ukraine (32°E) eastwards to N Xinjang (88°E). Figs. 22–25 ..................................................... ulianini

30b Postpetiole and frons narrower, pubescence hairs on gaster tergites longer. Discriminant 93.21* FRS +27.62*PpW-406.34* PLG-6.246 <0 [error 0% in 7 specimens]..................................................................... 31

31a Postocular distance and postpetiole height much larger (PoOc/CL> 0.423, PpH/CS> 0.260). Foveolae on vertex mediad of eye with diffuse margins ( Fig. 29 ). Anterior face of petiole node moderately sloping down ( Fig. 27 ). SE Kazakhstan. Figs. 26–29 .............................................................................................. littoralis

31b Postocular distance and postpetiole height smaller (PoOc/CL <0.423, PpH/CS <0.260). Foveolae on vertex mediad of eye more strongly demarcated and bicoronate ( Fig. 33 ). Anterior face of petiole node steeper ( Fig. 31 ). Iranian Caspi region. Figs. 30–33 .................................................................................. caspiense n. sp.

Figures

FIGURES 109–113. Cardiocondyla obscurior; Fig. 109: head in dorsal view; Fig. 110: lateral view; Fig. 111: dorsal view; Fig. 112: head surface between inner eye margin and paramedian vertex; Fig. 113: frontolateral aspect of waist segments. Galapagos: Santa Cruz, 2012.11.19

FIGURES 105–108. Cardiocondyla wroughtonii; Fig. 105: head in dorsal view; Fig. 106: lateral view; Fig. 107: dorsal view; Fig. 108: head surface between inner eye margin and paramedian vertex. Malaisia: Lundu, 2007.05.27

FIGURES 86–89. Cardiocondyla stambuloffii; Fig.86: head in dorsal view; Fig. 87: lateral view (flipped horizontally); Fig. 88: dorsal view; Fig. 89: head surface between inner eye margin and paramedian vertex. Georgia: river Podshori, 2010.08.27

FIGURES 94–97. Cardiocondyla rolandi n. sp., holotype; Fig. 94: head in dorsal view; Fig. 95: lateral view; Fig. 96: dorsal view; Fig. 97: head surface between inner eye margin and paramedian vertex. China: Oasis Yengisar, 2004.09.03

FIGURES 6–9. Cardiocondyla elegans; Fig. 6: head in dorsal view; Fig. 7 lateral view; Fig. 8: dorsal view; Fig. 9: head surface between inner eye margin and paramedian vertex. France: Brehemont-1.1 km E, 2010.07.15.

FIGURES 22–25. Cardiocondyla ulianini; Fig. 22: head in dorsal view; Fig. 23: lateral view; Fig. 24: dorsal view; Fig. 25: head surface between inner eye margin and paramedian vertex. Kyrgistan: 42.44.41N, 75.49.50E, 2000.07.30-91.

FIGURES 114–117. Cardiocondyla fajumensis; Fig. 114: head in dorsal view; Fig. 115: lateral view; Fig. 116: dorsal view; Fig. 117: head surface between inner eye margin and paramedian vertex. Egypt: Hurghada, 2008.04.16

FIGURES101–104.Cardiocondylatibetana, paratype;Fig.101:headin dorsalview; Fig.102:lateral view (flippedhorizontally); Fig. 103: dorsal view; Fig. 104: head surface between inner eye margin and paramedian vertex (flipped horizontally). China: Tarim: Ceele Station, 1966.08.26

FIGURES 98–100. Cardiocondyla gibbosa, lectotype; Fig. 98 head in dorsal view; Fig. 99: lateral view; Fig. 100: dorsal view. Kazakhstan: Suzak, 1923.07.03

FIGURES 90–93. Cardiocondyla koshewnikovi; Fig. 90: head in dorsal view; Fig. 91: lateral view (flipped horizontally); Fig. 92: dorsal view; Fig. 93: head surface between inner eye margin and paramedian vertex. Kazakhstan: Sassy Kol, 2001.08.07

FIGURES 118–121. Cardiocondyla unicalis, holotype; Fig. 118: head in dorsal view; Fig. 119: lateral view; Fig. 120: dorsal view; Fig. 121: head surface between inner eye margin and paramedian vertex. Iran: Ma’amulam, 1973.08.06

FIGURES 122–125. Cardiocondyla emeryi; Fig. 122: head in dorsal view; Fig. 123: lateral view; Fig. 124: dorsal view; Fig. 125: head surface between inner eye margin and paramedian vertex. Angola: Candelela, 2022.02.11

FIGURES 126–128. Cardiocondyla minutior; Fig. 126: head in dorsal view; Fig. 127: lateral view; Fig. 128: dorsal view. Malaisia: Melina Beach-Paya, 2007.06.03

FIGURES 129–132. Cardiocondyla mauritanica; Fig. 129: head in dorsal view; Fig. 130: lateral view; Fig. 131: dorsal view; Fig. 132: head surface between inner eye margin and paramedian vertex. Angola: Luanda, 2022.02.0

FIGURES 62–65. Cardiocondyla opistopsis, holotype; Fig. 62: head in dorsal view; Fig. 63: lateral view (flipped horizontally); Fig. 64: dorsal view; Fig. 65: head surface between inner eye margin and paramedian vertex. Kuweit: Burgan, 1988

FIGURES 66–69. Cardiocondyla rugulosa, holotype; Fig. 66: head in dorsal view; Fig. 67: lateral view (flipped horizontally); Fig. 68: dorsal view; Fig. 69: head surface between inner eye margin and paramedian vertex. Yemen: Sanaa, 1991.05

FIGURES 70–73. Cardiocondyla kushanica, paratype; Fig. 70: head in dorsal view; Fig. 71: lateral view (flipped horizontally); Fig. 72: dorsal view; Fig. 73: head surface between inner eye margin and paramedian vertex. Afghanistan: Darountah, 1958.01.21

FIGURES 82–85. Cardiocondyla verdensis n. sp., holotype; Fig. 82: head in dorsal view; Fig. 83: lateral view; Fig. 84: dorsal view; Fig. 85: head surface between inner eye margin and paramedian vertex. Cape Verde: Sao Nicolao, 2003.07.21

FIGURES 18–21. Cardiocondyla brachyceps; Fig. 18: head in dorsal view; Fig. 19: lateral view; Fig. 20: dorsal view; Fig. 21: head surface between inner eye margin and paramedian vertex. Iran: Fars, 1997.09.19

FIGURES 14–17. Cardiocondyla dalmaticoides n. sp., holotype; Fig. 14: head in dorsal view; Fig. 15: lateral view, postpetiole and gaster in full lateral view shown upper left; Fig. 16: dorsal view, postpetiole and gaster in full dorsal view shown lower left; Fig. 17: head surface between inner eye margin and paramedian vertex. Turkey: Reyhauli-2 km N, 1993.06.09

FIGURES 10–13. Cardiocondyla dalmatica, Fig. 10: head in dorsal view; Fig. 11: lateral view; Fig. 12: dorsal view; Fig. 13: head surface between inner eye margin and paramedian vertex. Hungary: Balassagyarmat, 2007.06.31

FIGURES 74–77. Cardiocondyla tenuifrons, holotype; Fig. 74: head in dorsal view; Fig. 75: lateral view (flipped horizontally); Fig. 76: dorsal view; Fig. 77: head surface between inner eye margin and paramedian vertex (flipped horizontally). Jordan: Abdallah, 1996.03.30

FIGURES 58–61. Cardiocondyla semirubra, paratype; Fig. 58: head in dorsal view; Fig. 59: lateral view; Fig. 60: dorsal view; Fig. 61: head surface between inner eye margin and paramedian vertex. Turkey: Sanliurfa-20 km S, 1993.06.12

FIGURES 54–57. Cardiocondyla batesii; Fig. 54: head in dorsal view; Fig. 55: lateral view; Fig. 56: dorsal view; Fig. 57: head surface between inner eye margin and paramedian vertex. Spain: Padul-4.5 km WNW, 2001.04.18

FIGURES 78–81. Cardiocondyla nigra; Fig. 78: head in dorsal view; Fig. 79: lateral view; Fig. 80: dorsal view; Fig. 81: head surface between inner eye margin and paramedian vertex (flipped horizontally). Cyprus: Linda Evangelista, 2003.03

FIGURES 34–37. Cardiocondyla gallilaeica, holotype; Fig. 34: head in dorsal view; Fig. 35: lateral view (flipped horizontally); Fig. 36: dorsal view; Fig. 37: head surface between inner eye margin and paramedian vertex. Israel: Sedé Eliyyahu, 1983.09.06

FIGURES 42–45. Cardiocondyla bulgarica; Fig. 42: head in dorsal view; Fig. 43: lateral view; Fig. 44: dorsal view; Fig. 45: head surface between inner eye margin and paramedian vertex. Bulgaria: Burgas-7 km S, 2003.09.30

FIGURES 46–49. Cardiocondyla sahlbergi; Fig. 46: head in dorsal view; Fig. 47: lateral view; Fig. 48: dorsal view; Fig. 49: head surface between inner eye margin and paramedian vertex. Russia: Barkhan Sarykum, 2017.04.21

FIGURES 50–53. Cardiocondyla persiana, paratype; Fig. 50: head in dorsal view; Fig. 51: lateral view; Fig. 52: dorsal view; Fig. 53: head surface between inner eye margin and paramedian vertex. Iran: Shiraz-7 km NE, 1997.09.18

FIGURES 38–41. Cardiocondyla israelica; Fig. 38: head in dorsal view; Fig. 39: lateral view (flipped horizontally); Fig. 40: dorsal view; Fig. 41: head surface between inner eye margin and paramedian vertex. Egypt: Ebn Salam Mansora, 2002.11.07

FIGURES 26–29. Cardiocondyla littoralis; Fig. 26: head in dorsal view; Fig. 27: lateral view; Fig. 28: dorsal view; Fig. 29: head surface between inner eye margin and paramedian vertex. Kazakhstan: Sassy Kol, 2001.08.07.

FIGURES 30–33. Cardiocondyla caspiense n. sp., holotype; Fig. 30: head in dorsal view; Fig. 31: lateral view; Fig. 32: dorsal view; Fig. 33: head surface between inner eye margin and paramedian vertex. Iran: Miankaleh, 2004.07.23