Taxonomic review of the Cataglyphis livida complex (Hymenoptera, Formicidae), with a description of a new species from Iran

Abstract Cataglyphis ficisp. nov., a member of the Cataglyphis livida complex, is described from the Estahban county of south-western Iran. The species is characterized by small body size and yellow to yellowish red body coloration with distinctly infuscated legs. Additionally, the taxonomic status of all known members of the Cataglyphis livida complex is updated. Cataglyphis lutea Pisarski, 1967, stat. rev. is raised to the species level and Cataglyphis viaticoides (André, 1881) is proposed as a senior synonym of Cataglyphis livida bulgarica Atanassov, 1982, syn. nov. and Cataglyphis albicans mixtus (Forel, 1895), syn. nov. Finally, a provisional key to members of the livida complex is provided.


Introduction
The ant genus Cataglyphis Foerster, 1850 currently includes 112 valid species and subspecies (Bolton 2020) distributed mostly in the semideserts and deserts of the Palearctic Region and the sub-Saharan area, India, and Pakistan. Cataglyphis species are among The work presented here is a contribution to studies on members of the C. livida complex. We list an updated synopsis of members of this complex and provide a provisional key to their identification. Additionally, we describe Cataglyphis fici sp. nov., a new member of the C. livida complex, based on material recently collected from Iran.

Materials and methods
Investigated specimens were collected in fig orchards located in Estahban city, Fars Province, Iran and are part of the material gathered for a scientific project conducted by the second author. The city is placed 1730 m a.s.l and is characterized by a dry climate, with a yearly precipitation amount of 224 millimeters and summer temperatures frequently exceeding 25.0 °C.
The dominant method was direct sampling (hand collecting). Individual specimens were collected on the ground and preserved in 75% EtOH. Photographs 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 (www.AntWeb.org) and are accessible using the unique CASENT identifying specimen code.
Examined specimens are housed in the following collections: MNHW Museum of Natural History, University of Wrocław, Poland, in temporary deposit by Department of Biodiversity and Evolutionary Taxonomy, University of Wrocław, Poland; MHNG Muséum d'Historie Naturelle, Genève, Switzerland.

Measurements:
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 at the center of the eyes; SL scape length; maximum straight-line length of scape excluding the basal condylar bulb; 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 propodeum 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; HFL hind femur length; measured on dorsal side from trochanter to apex of femur.

Diagnosis of workers of the Cataglyphis livida complex
Small body size (WL < 3.0 mm); colonies with monomorphic workers, lacking distinct major or soldier castes; petiole nodiform with angled dorsal outline and short peduncle; body subtly microsculptured and shiny; body uniformly yellow to red (never brown to black) or bicolored with entirely to partially black gaster. Distribution: from Morocco to Asia Minor and the Middle East, in semideserts, deserts and rocky open areas such as dry hills or coastal cliffs. Note 1. Agosti (1990), as the first one, noticed that the data from labels of type specimens of C. viaticoides did not correspond with the original description of the species. This problem was later investigated and clarified by Bračko et al. (2016). Based on evidence gathered and discussed by the authors, the definition of the livida complex proposed by Agosti (1990) was modified to accommodate C. viaticoides.
Note 2. Based on its description, C. albicans fezzanensis Bernard is characterized by the presence of polymorphic worker caste, and additional study on the type specimen * Due to ambiguous description of this species it is tentatively assigned to the livida complex.
indicated that its body sculpture is stronger and less shiny than in other members of the livida complex. Based on this data, we decided not to list this species as a member of this complex.
Provisional key to the Cataglyphis livida complex Diagnosis. Cataglyphis fici is a member of the Cataglyphis albicans group and can be separated from all species clustered in the cinnamomea and fortis complexes and most of members of the albicans complex by yellow to yellowish red body coloration; while other species have body completely brown to black. From bicolored species of the albicans complex it differs in smaller body size (WL < 3 mm) and presence of infuscated to * Note. The taxonomic status of C. arenaria, C. argentata, and C. aurata is unclear and requires further studies. Due to the lack of distinct morphological differences between types of these taxa and their ambiguous descriptions, we decided to list all three species in one couplet. For more details, see comments in the species accounts. yellowish brown mid and hind legs. However, C. fici is most similar to species included in the C. livida complex and can be easily separated based on at least mid and hind legs partly to mostly infuscated to yellowish brown. In contrast, all remaining members of the C. livida complex have legs uniformly colored and always in the same shade as mesosoma and head. Additionally, C. fici differs from C. lutea, C. arenaria, C. argentata, C. aurata, and C. livida in gaster darker than head and mesosoma sometimes infuscated. Color. Head, mesosoma and petiolar scale from yellow to yellowish red, in the darkest specimens sides of mesonotum and propodeum indistinctly infuscated, gaster in the palest specimens mostly yellow with infuscated two apical sternites (Fig. 6), in dark specimens yellowish at base then gradually infuscated apically, with dark brown apical tergites and sternites (Fig. 5). Forelegs in the palest specimens completely yellow, mid and hind legs with brown femora and tibiae and yellowish tarsi, in dark specimens all legs at least partly infuscated. Usually fore coxa mostly brown with reddish spots of  diffused borders laterally, fore femora mostly brown with yellowish apices, fore tibiae yellowish brown and fore tarsi yellowish; mid and hind femora dark brown, tibiae yellowish brown to brown, tarsi yellowish to yellowish brown (Figs 2, 5). Antennal scape yellow, funicles in the palest specimens slightly darker than scape, yellowish basally and yellowish brown apically, in dark specimens only first segment of funiculus yellowish, remaining segments gradually yellowish brown to dark brown. Head. Subrectangular, approximately 1.05 × as long as wide, sides below eyes almost parallel, above eyes gently convex, occipital margin convex (Fig. 3). Anterior margin of the clypeus convex, with small median emargination, clypeal anterior margin with a row of short, white setae and additional six long, white setae, the longest as long as 0.7 × length of clypeus. Whole surface of clypeus densely microreticulate with shiny background, covered with very sparse and short, adpressed hairs. Eyes large, oval, approximately 1.4 × as long as wide. Frontal carinae short, not extending beyond frontal lobes, interocular area without shiny line or carina and with a pair of long white setae. Antennal fossa shallow, microreticulate with shiny background. Whole head surface finely microreticulate with shiny background, occipital part of the head and are behind eyes with reduced sculpture and shinier, covered with extremely sparse, indistinct, adpressed hairs. Ocellar region with a pair of moderately long white setae, occipital part of head with 2-6 long, white erect setae, underside only with a pair of long, white setae close to lateral margin of head. Antennal scape moderately long, in frontal view almost straight, approximately 1.1 × as long as length of the head; thin, in apex only slightly and gradually widened; its base without tooth. Funiculus long, first funicle segment elongated, approximately 0.8 × as long as segments II and III combined, and 1.7 × as long as segment II (Fig. 3). Surface of scape densely microsculptured; shiny to indistinctly opalescent; covered with strong, moderately dense, decumbent setae. Mandibles rounded, only in basal part smooth and shiny, apical ¾ with deep grooves, surface shiny with several long yellow setae, masticatory margin with four4 large teeth. Mesosoma. Long, 2.1 × as long as wide; metanotal groove shallow (Fig. 2). Pronotum convex on sides (Fig. 1). In lateral view promesonotum slightly arched in profile; propodeum positioned lower than promesonotum, moderately convex in lateral view; propodeal spiracle strongly elongated and slit-shaped, approximately 4.2 × as long as wide (Fig. 2). Whole mesosoma opalescent, with dense, fine microreticulation and shiny background; covered with extremely sparse and short adpressed microsetae, on sides of pro-and mesonotum appears almost hairless, only anterior part of pronotum, posterior angles of mesonotum and propodeum with sparse setosity. Pro-and mesonotum without erect setae, propodeum without erect setae or in its posterior part with one or two short, white, erect setae. Scale. In form of a short cuneiform node, in lateral view almost trapezoidal with very short peduncle. Anterior face close to base distinctly convex, posterior face slightly concave, top of scale in lateral view obtusely rounded, without erect setae (Fig. 4). In anterior and posterior view top margin of scale without emargination. Surface of petiole distinctly microreticulate and shiny. Gaster. With fine transverse microreticulation and striation and very shiny background. Whole surface of gaster with hardly visible, extremely short, sparse, adpressed microsetae, tergites I and II without erect setae, tergites III and IV with a pair of long white setae centrally, each gastral sternite with 2-4 long, white, erect setae. Legs. Dorsal and lateral surface of femora and tibiae covered with sparse, white adpressed setae. Ventral surface of femora and tibiae with rows of elongate, white, erect spiniform setae.
Diagnosis. Whole body yellow, only gaster sometimes with indistinctly infuscated apex; mesosoma, posterior part of the head and coxae covered with a layer of silvery hair. Distribution. North Africa region, from Mauritania to Jordan. Based on photographs available on AntWeb specimens from Arabian Peninsula probably refer to C. lutea.
Note. Cataglyphis arenaria was separated from C. livida and C. viaticoides based on the presence of a thick layer of silvery hair on mesosoma and posterior part of the head. Two years after the original description of C. arenaria, Karavaiev (1911) described Myrmecocystus albicans ssp. lividus var. aurata, which was later validated by Menozzi (1932) as trinominal. The latter species also was separated from C. livida based on presence of silvery hair on its body. Probably, Karavaiev was unaware of the existence of C. arenaria during his work on C. aurata. Study of type specimens and descriptions of both C. arenaria and C. aurata did not provide any characters useful in separating these two species. Thus, we conclude that both taxa could be conspecific (see also note in Cataglyphis argentata (Radoszkowsky, 1876) and C. aurata Menozzi, 1932). However, this hypothesis requires verification based on larger material collected from the whole area of their distribution, supported with studies on male genitalia, and genetic analyses.

Diagnosis.
Whole body yellow, only gaster sometimes with indistinctly infuscated apex; mesosoma, body covered with a layer of silvery hair.
Distribution. Egypt. Note. Type specimens of this species are considered lost and, as suggested by Agosti (1990), due to ambiguous description of this species, its assignation to the livida complex is tentative. The silvery hair mentioned in the description can suggest an affiliation of C. argentata with the bombycina or laevior complexes. If C. argentata is a member of the livida complex, then its description could indicate that it is probably conspecific with two other North African taxa: C. arenaria and C. aurata. If this assumption is correct, then the name C. argentata has priority over C. arenaria and C. aurata. Menozzi, 1932 Cataglyphis (Cataglyphis) albicans aurata Menozzi, 1932: 95 [first available use of Myrmecocystus albicans ssp. lividus var. auratus Karavaiev, 1911. Syntype worker, Assuan, Egypt (MHNG) [syntype worker images examined, AntWeb, CASENT0911100, photos by Zach Lieberman, available on AntWeb.org].

Cataglyphis aurata
Diagnosis. Whole body yellow, only gaster sometimes with indistinctly infuscated apex; mesosoma, posterior head and coxa covered with a layer of silvery hair. Distribution. North Africa. Probably records from Asia Minor refer to Cataglyphis lutea.
Note. Cataglyphis aurata was separated from C. livida based on the presence of a thick layer of silvery hair on its body. Probably, Karavaiev, during his work on C. aurata, was unaware of the existence of C. arenaria, another species described from the North African region characterized by the same feature. Study on type specimens and descriptions of both C. arenaria and C. aurata did not provide any characters useful separating these two species. Thus, we conclude that they could be conspecific. However, this hypothesis requires verification based on larger material collected from the whole area of their distribution, supported with studies on male genitalia, and genetic analyses. See also note in C. argentata. Diagnosis. Whole body yellow, only gaster sometimes with indistinctly infuscated apex; mesosoma and coxa covered with a layer of silvery hair. Distribution. Unknown. Due to mislabeling of type specimens of C. livida and C. viaticoides, both species were wrongly interpreted, and most of their historic records require verification. Based on available material, we can confirm its presence in Egypt, coastal parts of Israel, Syria, and Antalya Province in Turkey.

Cataglyphis livida (André, 1881)
Note. A study on type specimens of C. livida revealed that this species could be easily separated from most members of the livida complex based on the presence of a layer of silvery hair on propodeum and katepisternum, and lack of these on posterior head. Lack of comment on this feature in the original description combined with mislabeling of type specimens (see Bračko et al. 2016) caused confusion, leading to the long-lasting misinterpretation of C. livida. Pisarski, 1967 (Emery 1906), later validated by Pisarski (1967) as a subspecies of C. livida, and finally considered as its junior synonym (Radchenko 1997). A study on type specimen revealed that C. lutea distinctly differs from C. livida in lack of a layer of silvery hair on mesosoma, and its distribution does not overlap with confirmed records of C. livida. Thus, we decided to raise it to the species status. AntCat resources indicated that, except type locality, C. lutea is also known from Aran va Bidgol, Maranjab, Iran (CDA000106) and Saudi Arabia (CASENT0906455). Diagnosis. Head and mesosoma uniformly yellowish red to reddish yellow, gaster entirely or mostly dark; thin layer of silvery hair limited to propodeum. Distribution. Balkans and Asia Minor. Note. Radchenko (1997), based on confusion related to the type labels of C. livida and C. viaticoides (see Bračko et al. 2016), considered C. livida bulgarica and C. albicans mixtus as junior synonyms of C. livida. Results presented by Bračko et al. (2016) clarified that the only member of the livida complex with entirely or mostly black gaster is C. viaticoides and thus Cataglyphis livida bulgarica Atanassov, 1982 and Cataglyphis albicans mixtus (Forel, 1895) should be considered as its junior synonyms.