Cataglyphis Foerster, 1850

Agosti, Donat, 1990, Review and reclassification of Cataglyphis (Hymenoptera, Formicidae), Journal of Natural History 24, pp. 1457-1505 : 1461-1465

Cataglyphis Foerster

Cataglyphis Foerster, 1850:493 . Type-species: Cataglyphis fairmairei Foerster, 1850:493 [Junior synonym of Formica bicolor Fabricius, 1793: 356 ; by monotypy.

Monocombus Mayr, 1855: 493 . Type-species: Formica viatica Fabricius, 1787: 308 ; by monotypy. [Synonymized by Mayr, 1861: 44; re-erected as subgenus Cataglyphis (Monocombus) by Santschi, 1929a: 30.] Synonymy here reconfirmed.

Formica dubdiviso 3 Nylander, 1856: 59 [see note 1 below].

Myrmecocystus Wesmael, 1838 sensu Forel, 1878: 372 , p.p [see note 2 below]; Myrmecocystus, Lomnicki, 1925: 164 , misapplication of name.

Paraformica Forel, 1915: 95 (footnote). [As subgenus of Formica .] Type-species: Formica (Proformica) emmae Forel, 1909: 381 ; by monotypy [raised to genus by Emery, 1925: 260]; new combination and new status as subgenus of Cataglyphis (Paraformica) by Santschi, 1929a: 30. Syn. nov.

Machaeromyrma Forel, 1916: 441 [as subgenus of Cataglyphis ]. Type-species: Formica bombycina Roger, 1859: 232 ; by original designation. [Synonymy by Urbani, 1969: 218.] Eomononocombus Arnoldi, 1968: 1815 [as subgenus of Cataglyphis ]. Type-species: Myrmecocystus albicans st. cinnamomea Santschi, 1929a: 61 ; by original designation. [Synonymy by Hôlldobler and Wilson, 1990: 18.]

Note 1: Nylander (1856: 59) synonymized Cataglyphis and Monocombus with Formica , but the proposed name, subdiviso 3, is unavailable. Nevertheless, the diagnosis he gave is accurate and corresponds with the actual diagnosis of Cataglyphis

Note 2. Emery and Forel (1879) made this synonymy without explanations; possibly because Roger (1862: 254) stated that Myrmecocystus melligerus 'zeigt, mit Ausnahme des, bis zur Starke einer grossen Erbse aufgetriebenen Hinterleibes, vollkommene generische Uebereinstimmung mit Cataglyphis viatica \ Curiously enough, 1 year later Roger (1863:12) synonymized the older genus Myrmecocystus with the later published Cataglyphis (and therefore a junior synonym of Myrmecocystus ), a situation that was cleared up by Emery and Forel (1878: 372).

Diagnosis

Workers and females: Formicine ants with the following diagnostic characters.

1. Mandibles large triangular, a row of five teeth which become smaller from apical to basal (Fig. 6). In the bombycinus group sometimes up to seven teeth and large workers with a falcate mandible (Figs 4 and 5); in kurdistanicus large workers with an elongate mandible with a row of denticles; longitudinally striate.

2. Macrochaetae on the stipes of the maxilla.

3. Often a distinct psammophore formed at least by the macrochaetae on the stipes of the maxillae and on the inner apical margin of the mandibles; sometimes with long curved hairs on the third to fifth segment of the maxillary palps.

4. First maxillary palp segment flat.

5. Eyes located distinctly behind the midlength of the head.

6. Propodeal spiracles slit-shaped.

7. Wings (female caste) with a straight frontal side and very sparse, short pubescence (length of the hairs space between the hairs).

Males: Formicine ants with the following diagnostic characters.

1. Body of the same size as the females.

2. Ocelli and eyes only slightly enlarged in comparison to those in the female caste.

3. Stipes with a median appendage (Fig. 28).

4. The toothless zone on the sagittated side ventral of the ergot stretched (Fig. 63).

5. Front wing short in comparison to the alitrunk (WI> 50).

6. Dorsum of alitrunk black or bicoloured with two lateral bands and a semicircular spot cranially.

Distribution and biology

Cataglyphis species are distributed exclusively in the palaearctic realm, from Portugal to Mongolia and the Ganges valley in the East, from the Coté d’Ivoire and southern Sudan to Czechoslovakia, from sea level to over 3000 m. They avoid the forests, but live within the gaps of forested areas in Rosaceae forest of central Asia or the steppic forests, and can be found between sand dunes of the Sahara or the Chotts. Most species are found in steppic or Mediterranean-type habitats.

Summaries of the biology and physiology are to be found in Dlussky (1981), Wehner (1982, 1987), and Wehner et al. (1983).

Phylogeny of Cataglyphis

The taxonomic and systematic problems in Cataglyphis are mainly to be found at species level. Thus it is easier to place specimens in a species-group or complex than a species. Species-groups, as they are defined here, are taxa for which at least one autapomorphy can be found which is not merely a change in length or width.

The character states of male genitalia characters which are included in the analysis (Table 1) are the following:

1. Subgenital plate short, SPI <125 (0); long, SPI> 125 (1).

2. Subgenital plate bilobed, sometimes with an emarginate median part (0) (Figs 13, 18, 19, 25-27); distally emarginate, without lateral lobes (1) (Fig. 17); trilobed (2) (Figs 14-16, 20-24).

3. Squamula and stipes confluent (0) (Figs 33-39); squamula overlapping stipes caudally (1) (Figs 28-30, 32,40-42).

4. Stipes without a mediobasal appendage (0) (Figs 29, 30, 40-42); with a mediobasal appendage, separated from the stipes by a carinae (1) (Figs 28, 33-39); or confluent (2) (Figs 31, 32).

5. Volsella straight (0) (Figs 48-52); volsella curved (1) (Figs 53-62).

6. Sagitta with serrated face ventrally [if not serrated, then ventral face simple] (0) (Figs 64, 66-74); sagitta with serrated face laterally (1) (Figs 63, 65).

7. Median fold pointing towards the basal end of the ventral face (0) (Figs 63-65); towards anterior edge of the ventral face (1) (Figs 66, 67); towards the anterodorsal appendage (2) (Figs 68-77).

8. Anterodorsal appendage absent (0) (Figs 63-67); present (1) (Figs 68-77).

Only characters indicating relationships are presented. Synapomorphies and autapomorphies are given in Fig. 1. The characters are explained in the text.

9. Sagitta with anterior (ventral) part depressed (0) (Figs 63 and 64); a quadrangular to rectangular shield (1) (Figs 64, 66, 67); elongated (2) (Figs 68-77).

The analysis of the data, using mhennig* of hennig86 and unordered, unweighted characters, revealed two equally parsimonious trees (17 steps; ci = 76, ri = 85), of which the one chosen matched best with characters of the female castes (Fig. 1). Proformica , as the supposed sister-group of Cataglyphis (Agosti, in preparation) has been chosen as the outgroup. All trees showed the main division at node 16, which is also supported by the appearance of a squamiform petiole in clade 14 and a nodiform petiole and elongated third and fourth maxillary palp segment in clade 15. Clade 15 corresponds with the former subgenus Cataglyphis sensu strictu.

Within clade 14 the emmae group and the bombycinus group are always sister-group and together they are the sister-group of the pallidus group.

Finally the sister-group relation altisquamis groupj cursor group has been favoured because otherwise the autapomorphies of the altisquamis group are two reversals (characters 7 and 9) and the lateral serrated face of the sagitta (character 6), which is the autapomorphy of the sister-group altisquamis group I cursor group in the proposed tree, is a homoplasy. This treatment best matches the assumed relationship based on female castes.

The sister-group of the pallidus group has always been bombycinus group and emmae group.

This system does not support Santschi’s subgenera ( Santschi, 1929a), because his subgenus Monocombus (= altisquamis, cursor and pallidus groups) is polyphyletic.

It is hoped that further analysis, including characters of the female castes and species as terminal taxa, will reduce the number of trees (Agosti, unpublished) and furthermore make Cataglyphis available for zoogeographic studies.