Cataglyphis cf. savignyi (Dufour, 1862)

A. Ionescu & P. - A. Eyer, 2016, Notes on Cataglyphis Foerster, 1850 of the bicolor species-group in Israel, with description of a new species (Hymenoptera: Formicidae), Israel Journal Of Entomology 46, pp. 109-131 : 126-129

Cataglyphis cf. savignyi (Dufour, 1862)

(Figs 17, 29–31)

Recognition: Cataglyphis cf. savignyi is similar to C. niger , but in ten out of 20 sampled localities all medium and large specimens display the typical ‘ savignyi ’ color pattern. In other localities most ants are black: in the Hazeva samples four out of 22 ants have red head, of the 49 ants from the northern Coastal Plain only two have brownish-red head and mesosoma, while the 31 ants from Mikhmoret and Palmahim and 23 ants from the central Negev are completely black. All 51 examined specimens from 11 localities in Egypt have red head. This species has raised mesonotum over pronotum in 74% (n=182) of examined specimens, a higher frequency than in C. niger and the ants from Egypt (37 %, n=51). C. cf. savignyi shows a wide range of inter-population variability for most of the investigated characters. Scape and midfemur length of Israeli ants in most of the sampled localities is similar to that in C. cf. savignyi from Tunisia and the Nile region in Egypt, but is signiFcantly longer in ants from Nir Yizhaq, Mishor Yamin and Sinai ( Table 2 ). Likewise, the propodeum of ants from Nir Yizhaq, Mishor Yamin and Sinai is signiFcantly higher than in other places ( Table 2 ). Sculpture in most Israeli ants is similar to C. niger and C. cf. savignyi from Sinai, but gaster is laterally shiny in samples from the northern Coastal Plain, Mikhmoret and Modi’in, and in C. cf. savignyi from Tunisia and the Nile region in Egypt. Pilosity is variable among localities. For example, the pubescence on the convex face of scape is appressed to subappressed on distal ¼ of length in ants from central Negev, similar to C. cf. savignyi from Tunisia, the Nile Delta and SE Sinai, but subappressed to decumbent, with occasional 1–2 erect hairs on distal ¾ of length in ants from Tel ’Arad, although the erect hairs are fewer than in C. isis , and shorter than in C. nigripes . In C. cf. savignyi from Israel 48 out of 176 examined specimens display 1–4 setae on propodeum dorsum, but the frequency of ants with setae on propodeum dorsum is higher in Palmahim, Modi’in and Tel ’Arad (25/54) than in the northern Coastal Plain and central Negev (5/62), and Egypt (2/51). Overall, ants from the southern Coastal Plain seem similar to ants from northern Sinai and the Nile region, while ants from the central Negev resemble ants from central and SE Sinai.

Measurements: TL=5.3–12.8, HL=1.50–3.22, HW=1.31–3.16, EL=0.43–0.88, iOD=0.94–2.42, OcClD=0.71–1.51, SL=1.74–3.36, F1L=0.49–0.91, F2L=0.25– 0.51, ML=2.38–4.61, MW=0.92–2.05, PrL=0.82–1.68, PrH=0.22–0.55, PNL= 0.46–0.96, PNW=0.29–0.70, PNH=0.25–0.57, PNA=63–81°, mFmL=1.95–4.22, hTbL=2.44–5.22 (n=200).

Material examined: Israel: Bezet, 525, 15.iii.2014, T. Reiner-Brodetzki (10☿, 1Ƌ); Nahariyya, 2012, R. Zeltzer (11☿); Bustan HaGalil, 2012, R. Zeltzer (17☿); ’Akko, 3.xi.2010, R. Zeltzer (8☿); Mikhmoret, 2012, R. Zeltzer (20☿); Ra’ananna, 10.v.1944, H. Bytinski-Salz (1♀, 2Ƌ); Palmahim, 2012, M. Saar (20☿); Modi’in, 3.iii.2011, R. Zeltzer (21☿); Suseya, 15.iv. 2016, A.L.L. Friedman (1☿); Tel ’Arad, 26.ii.2009, A. Ionescu (3☿), 10.iv.2011, R. Zeltzer (4☿), #1, 10.iv.2011, R. Zeltzer (4☿), #2, 10.iv.2011, R. Zeltzer (4☿), #3, 10.iv.2011, R. Zeltzer (1☿), #4, 10.iv.2011, R. Zeltzer (4☿), #5, 10.iv.2011, R. Zeltzer (6☿); Ziqim East, #1, 2012, R. Zeltzer (3☿), #2, 2012, R. Zeltzer (3☿), #3, 2012, R. Zeltzer (2☿), #4, 2012, R. Zeltzer (2☿); Horbat Mamshit, 28.iv.19[??], H. Bytinski-Salz (1♀, 3Ƌ); Hazerim, 3.iv.2011, R. Zeltzer (3☿), #1, 3.iv.2011, R. Zeltzer (1☿), #2, 3.iv.2011, R. Zeltzer (4☿), #3, 3.iv.2011, R. Zeltzer (5☿); Bor Mashash, 20.iv.2005, G. Kunikov (1☿, 2Ƌ); Holot Mashash, 14.iv.2003, A. Mozer (3☿, 2Ƌ); Sede Boqer, 20.vi.1974, B. Mordechai (3☿, 1♀), 16.v.1974, J. Morad (1Ƌ); Nir Yizhaq, 3.iv.2011, R. Zeltzer (12☿,1Ƌ); Yeroham, 11.iv.19[??], H. Bytinski-Salz (1Ƌ); Mishor Rotem, 26.ii.2009, A. Ionescu (3☿); Mishor Yamin, 10.iv.2011, R. Zeltzer (16☿); Qadesh Barnea’, 29.v.1970, H. Bytinski-Salz (1☿); Holot ’Agur, 3.iii.2016, I. Renan (4☿); Hazeva, 25.v.1992, A. Ionescu (1☿, 1♀, 1Ƌ), 15.i.1994, Shapira (4☿), 25.i.1994, Shapira (8☿), 6.ii.1994, Shapira (8☿), 22.iii.1994, A. Ionescu (2☿, 1♀, 1Ƌ). Egypt: Quassasin, 16.xii.1945 (5☿); Saqqara (Sakkara), 12.ii.1981, J. Kugler (1☿); Dendera, 15.ii.1981, J. Kugler (2☿); RaFah, 26.vi.1973, B. Shalmon (2☿) (desertorum sensu Santschi); W. Garandel, 28.iii.1969, J. Kugler (6☿); Mitla, 25.v.1971, H. Bytinski- Salz (3☿), 13.iv.1973, D. Furth (4Ƌ); Abu Ageilah, 6.iv.1968, Ch. Saudler (3☿); Oasis Feran, 25.v.1971, H. Bytinski-Salz (1☿); Djebel Katarina, 6.iv.1968, Ch. Saudler (3☿); Nabq, 4.iv.1968, Ch. Saudler (5☿); Ein Khadidja [En Chadjiah] (70 km SSW Nabq), 23.iv.1968, H. Schweiger (4☿).

C. cf. savignyi from northern Africa: Tunisia: Kairouan, Dr. Santschi (2☿), identiFed by Santschi as var. desertorum Forel (junior synonym of savignyi ).

C. oasium (sensu Santschi): Egypt: Alexandria, 10.xi.1937 (3☿); Assiut, 14.ii.1981, J. Kugler (5☿); Komombo, 15.ii.1981, J. Kugler (2☿); Wadi Feran, 18.v.1970, H. Bytinski-Salz (2☿). Iran: Shiraz Env., 16–21.v.1972, H. Bytinski-Salz (2☿).

C. nigripes : Iran: Shiraz Env., 16–21.v.1972, H. Bytinski-Salz (19☿, 3Ƌ).

C. abyssinica (sensu Santschi): Senegal: ‘309’ (1☿).

Distribution: The species was reported as widely distributed from the northern fringe of the Sahara desert, from the Atlantic to northern Sinai and the Arabian Peninsula (Collingwood & Agosti 1996).

Notes: The Santschi’s (1929) key for the bicolored specimens of C. cf. savignyi from Israel will identify them as ‘ savignyi ’, with the exception of the samples from Tel ’Arad that key out to C. st. saharae Santschi. The ants from Egypt will split into ‘ savignyi ’ and ‘ oasium ’ (the samples from Alexandria, Assiut, Komombo and Wadi Feran); however, using this key, all identiFcations of ants from Israel and Sinai will be dubious, except for the ants from Modi’in, because of the accentuated gaster sculpture. Using the identiFcation key of Collingwood and Agosti (1996) all C. cf. savignyi specimens from Israel and Egypt with bright-red mesosoma and/or low propodeum (Fig. 17) and petiole will key out to C. abyssinicus or C. saharae (sensu Collingwood in Sharaf et al. 2015).

A particular difFculty in deFning the niger species-complex stems from the uncertainty related to the identity of C. savignyi . The description of Formica savignyi by Dufour (1862) is based on two illustrations of a worker and a male from Egypt by J. Savigny, and on specimens in the Dufour’s collection considered by him as similar to the illustrations. The described workers were collected in the region of the Pyramids (Egypt) and in Algeria; and the single male was collected in Oran (Algeria). Santschi (1929) assumed that Dufour’s types had been lost and designated new types from Egypt and Libya (AntWeb CASENT0912229) that best Ft the original description, and provided a new diagnosis. However, Agosti (1990) retrieved two workers from ‘Gizeh’ and one worker and one male from ‘North Africa’ labeled savignyi in Dufour’s collection in the Muséum National d’Histoire Naturelle, Paris. The workers belong to the mauritanicus speciescomplex (AntWeb CASENT0915501) and the male to the niger species-complex, the latter being selected as the lectotype of C. savignyi (Agosti 1990) .

It is probable that the lectotype of C. savignyi is related to the bicolor group populations from northern Africa that were thoroughly investigated by morphometric, biochemical and molecular means by Wehner et al. (1994) and Knaden et al. (2005). These populations are genetically distinct from the Israeli C. cf. savignyi , and in the molecular phylogenies reconstructed in Knaden et al. (2012) and Aron et al. (2016) C. savignyi is paraphyletic. Because the name C. savignyi should be reserved for populations from which the type was selected, it seems that C. cf. savignyi from Israel needs renaming. C. cf. savignyi from Israel, C.

abyssinica from Egypt and C. niger are part of the same clade, but the taxonomic importance of the genetic divergence between C. niger and these other populations (Aron et al. 2016) is yet unknown; and, therefore, a nomenclatural change under the present state of knowledge of these ants seems inadvisable.

We will refer to C. niger and C. cf. savignyi from Israel collectively as the C. niger -complex because of the following: (i) C. niger cannot be reliably distinguished from black C. cf. savignyi from the Coastal Plain region; (ii) there are too few samples of C. cf. savignyi from NE Africa and the Middle East, that have been investigated by molecular means for conFdent assessment of the phylogenetic relationships between these ants; (iii) it is unclear, which ant population the name C. savignyi should be attributed to.

Discernment of species using traditional identiFcation keys or morphometrics within the niger species-complex is very difFcult for ants from Israel, if indeed there is more than one species. The primary cause of the inability to produce an identiFcation key based on binary characters for use in faunistic studies, etc., which discriminates between niger and savignyi , is the bewildering variability of morphological characters of individuals within and among populations of C. cf. savignyi . This difFculty extends also to discrimination between C. cf. savignyi and other species, e.g. dark colored C. israelensis in the contingency areas between the species, or C. abyssinicus from Egypt (Knaden et al. 2014). Nevertheless, since C. cf. savignyi and C. israelensis are parapatric in Israel, the following key characters in conjunction with the known geographical distribution can be useful.

Tables

Table 2: Least square means ± 95 % conFdence interval at covariate mean, and sample size for four variables per sampling localities grouped by taxon and geographic region; see ‘ Materials and Methods’ for full names of variables and Fig. 1 for numbers and names of geographic regions.

Locality, no. of geogr. region Montfort & Daverat, 1, 5 Hula & ’Ammi’ad, 1b, 2 ’Atlit, 4 Hof haBonim, 4 Allon, 12 Tel Barukh, 8	SL ln(HW)=0.74 2.47±0.03, 22 2.49±0.04, 14 2.54±0.03, 18 2.54±0.04, 12 2.63±0.03, 21 2.51±0.04, 11	PrH PrL=1.12 0.30±0.01, 17 0.32±0.01, 13 0.33±0.01, 29 0.35±0.01, 17 0.29±0.01, 24 0.34±0.01, 12	PNH PNL=0.71 0.42±0.02, 10 0.42±0.02, 13 0.43±0.01, 29 0.44±0.01, 15 0.43±0.02, 14 0.41±0.02, 12	PNA PNL=0.73 71.4±2.4o, 8 72.4±2.1o, 11 77.3±1.4o, 27 78.1±1.8o, 15 74.4±1.9o, 14 71.6±2.1o, 12	mFmL MW=1.54 3.06±0.04, 22 3.03±0.04, 14 3.03±0.03, 30 3.01±0.04, 17 3.07±0.03, 26 2.99±0.05, 12
Yafo, 8	2.52±0.02, 39	0.36±0.01, 42	0.40±0.01, 22	71.0±1.5o, 22	3.02±0.03, 43
Ziqim West, 9	2.53±0.04, 13	0.35±0.01, 13	0.37±0.02, 13	69.9±2.1o, 12	3.03±0.04, 18
Nahariyya, 4	2.52±0.04, 11	0.35±0.01, 11	0.38±0.02, 8	67.0±2.4o, 6	3.01±0.04, 14
’Akko, 4	2.46±0.03, 22	0.35±0.01, 15	0.39±0.02, 10	70.4±2.9o, 8	3.01±0.03, 25
Mikhmoret, 8	2.59±0.03, 17	0.35±0.01, 22	0.42±0.01, 18	72.3±1.8o, 15	3.02±0.04, 22
Palmahim, 9	2.58±0.03, 19	0.35±0.01, 11	0.42±0.02, 10	71.0±2.3o, 10	3.12±0.04, 20
Ziqim East, 9	2.58±0.04, 10	0.33±0.02, 10	0.37±0.02, 9	70.1±2.5o, 8	3.06±0.05, 10
Nir Yizhaq, 9	2.67±0.04, 9	0.38±0.01, 11	0.40±0.02, 8	68.7±2.5o, 8	3.24±0.05, 12
Modi’in, 10	2.56±0.04, 10	0.34±0.01, 14	0.39±0.02, 9	70.1±2.4o, 9	3.07±0.04, 21
Tel ’Arad, 15	2.58±0.03, 15	0.35±0.01, 22	0.42±0.01, 19	71.2±1.7o, 18	3.20±0.03, 25
Hazerim, 15	2.57±0.04, 11	0.36±0.02, 8	0.38±0.02, 5	71.3±4.1o, 3	3.13±0.05, 13
Mishor Yamin, 17	2.70±0.03, 15	0.38±0.01, 15	0.39±0.02, 14	70.3±2.1o, 12	3.24±0.04, 16
Hazeva, 14	2.73±0.03, 16	0.33±0.01, 22	0.39±0.01, 22	70.8±1.5o, 22	3.15±0.04, 21
Nile reg., Egypt	2.53±0.04, 11	0.37±0.01, 18	0.39±0.02, 14	72.0±2.0o, 13	3.12±0.04, 20
Sinai, Egypt	2.71±0.04, 11	0.39±0.01, 24	0.38±0.01, 16	69.9±1.6o, 20	3.23±0.03, 30