Zoological Indication of Climate Change in the Central Kazakh Steppe Compared to the Middle of the 20th Century Using the Example of Carabid and Tenebrionid Beetles

Abstract

Studies of communities of ground beetles (Carabidae) and black beetles (Tenebrionidae) were conducted in central Kazakhstan along 70° E in typical, dry, and desert steppe subzones in 1976–1978 and 2018. Parallel to that, soil-cover studies were conducted and the climate indicators of the studied areas were compared. Despite a considerable change in climate over 50 years, which has been expressed in a rise in temperatures with a simultaneous increase in precipitation, the basic features of soils have not undergone essential changes, but they keep or get strong potentials for solonetz and solonchak elementary soil processes, which are capable of drastically changing the structure of the soil cover. The generality of local fauna of carabids in 1976–1978 was 48–62% and, at the beginning of 21st century, it decreased to 16% in dry steppe and to 7% in desert steppe. In tenebrionids, which are more adapted to aridity, the faunal similarity decreased from 70–75 to 37% in the typical steppe and increased to 87% in the dry steppe. There are more “southern” subarid species in the communities and fewer relatively “northern” boreal and polyzonal species, which disappear completely in dry and desert steppes. The Tencar index that expresses the ratio of the number of individuals and species of arid tenebrionids and more humic carabids is used as an integral zoo indicator of changes in aridity of the environment. In the typical steppe, the index values are low and change little in the long-term dynamics and along the catena. However, in the dry steppe, Tencar index values increased 5 times in 2018 when compared to the middle of the 20th century and in desert steppe they grew by two orders of magnitude. The trigger for changes in the composition and structure of local communities of carabids and tenebrionids is not a change in atmospheric humidity, but a rise in average annual temperatures, which exceeded the global trend values and activated the salinization of soils, creating the conditions for the desertification of the territory and biota.

Appendices

Appendix 1.

Soil sections in the zonal series of steppe soils of central Kazakhstan, which were observed in 2018

Catena position	Coordinates	Soil	Horizons, cm	Description
Kazakhstan, Akmola Region, environs of the settlement of Shortandy. Virgin steppe site
EL	51.56608° N 71.28065° E	Southern carbonate deeply solonetzic medium-thick chernozem	Adk (0–12)	Light loamy, dry, dark gray, powdery, intertwined with roots of steppe vegetation, loose, soil effervesces from HCl
			Ak (12–43)	Light loamy, fresh, dark gray, slightly lumpy, loose, soil effervesces from HCl
			AVk (43–76)	Light loamy, fresh, gray-brown, tongue-shaped humus streaks on a brown background, compacted, lumpy–nutty prismatic, slightly sticky, soil effervesces from HCl
			Vk (76–103)	Light loamy, fresh, yellowish brown with gray spots of humus, prismatic, dense, soil effervesces from HCl
			Sks (103–…)	Light loamy, fresh, brownish with a whitish tint, structureless, less dense than horizon B, gypsum in the form of small grains, soil effervesces from HCl
TR1	51.56540° N 71.28464° E	Meadow chernozem solonetzic soil	Ad (0–9)	Light loamy, dry, dark gray, powdery, intertwined with roots of steppe vegetation, loose, soil does not effervesce from HCl
			Ak (9–37)	Light loamy, fresh, dark gray, lumpy–nutty, compacted, abundant with plant roots, soil effervesces from HCl
			AVk (37–74)	Light loamy, fresh, uneven grayish brown color due to tongue-shaped humus streaks on a yellowish brown background, lumpy–nutty, carbonate structures in the form of veins along the pores and root passages, compacted, soil effervesces from HCl
			Vk (74–102)	Light loamy, fresh, yellowish brown, prismatic, small dark spots of approximately 0.5–1 mm in size (possibly manganese structures), compacted, soil effervesces from HCl
			Sks (102–…)	Light loamy, fresh, brownish with a whitish tint, gypsum in the form of grains, loose, structureless, soil effervesces from HCl
TR2	51.56681° N 71.29067° E	Deeply carbonate solonetz	Adk (0–7)	Medium loamy, dry, light gray, powdery, loose, intertwined with roots of steppe vegetation, soil effervesces from HCl
			Ak (7–30)	Medium loamy, fresh, spotty brownish gray, slightly lumpy powdery, slightly compacted, rich in carbonates in the form of lines and tubes along root passages, soil effervesces from HCl
			V1k (30–54)	Heavy loamy, fresh, grayish brown, richly colored with humus, nutty–prismatic, abundant with carbonates in the form of lines and tubes along the passages of plant roots, sticky, compacted, soil effervesces from HCl
			V2k (54–77)	Heavy loamy, moist, yellowish brown with humic streaks, sticky, lumpy–prismatic, abundant with carbonates in the form of lines and tubes along the passages of plant roots, compacted, soil effervesces from HCl
Catena position	Coordinates	Soil	Horizons, cm	Description
			Vks (77–95)	Heavy loamy, moist, light brown, prismatic, spots of gypsum, spots of humus are found along the facets of individual structures, stickiness is barely noticeable, carbonates in the form of impregnation, compacted, soil effervesces from HCl
AC	51.56628° N 71.29117° E	Alluvial meadow-marshy soil	Adg (0–21)	A lot of undecomposed organic matter (approximately 40–50%), the mineral part is light loamy, very loose, moist, black with a brown tint and bluish tint, the structure of the mineral part is lumpy, soil does not effervesce from HCl
			Ag (21–79)	Light loamy, very loose, black with a bluish tint, wet, loosely lumpy, abundant with plant roots, bright brown spots, soil does not effervesce from HCl
			Bg (79–…)	Medium loamy, wet, lumpy grainy, black with a brownish tint, slightly compacted, soil does not effervesce from HCl
Kazakhstan, Akmola Region, environs of the settlement of Arykty. Lake depression
EL	50.54415° N 70.46271° E	Dark chestnut solonetzic soil	Ak (0–17)	Light loamy, fresh, powdery, brownish gray, loose, rounded sand grains of white and brown color, plant roots, upper 8 cm are strongly intertwined with plant roots, soil effervesces from HCl from the depth of 5 cm
			V1k (17–32)	Medium loamy, fresh, brownish gray, rounded sand grains of white and brown color, compacted, lumpy–prismatic, small plant roots, soil effervesces from HCl
			V2k (32–70)	Heavy loamy, fresh, nutty–prismatic, very dense, vertical cracks with streaks of humus, abundantly saturated with carbonates (the section wall becomes white upon drying), rounded sand grains of white and brown color, small plant roots, soil effervesces from HCl
			V3k (70–78)	Light loamy, fresh, light brown, indistinctly prismatic, compacted, rounded sand grains of white and brown color, saturated with carbonates, soil effervesces from HCl
			S1ks (78–101)	Light loamy, fresh, brown, structureless, slightly compacted upon drying, single spots of gypsum, rounded sand grains of white and brown color, soil effervesces from HCl
			S2ks (101–124)	Light loamy, fresh, brown, structureless, slightly compacted upon drying, gypsum accumulations in the form of spots and spherical (ball-shaped) formations, rounded sand grains of white and brown color, soil effervesces from HCl
			S3ks (124–…)	Light loamy, fresh, brown, structureless, loose, accumulations of gypsum in the form of spots, carbonates in the form of impregnation, rounded sand grains of white and brown color, soil effervesces from HCl
Catena position	Coordinates	Soil	Horizons, cm	Description
TR1	50. 54271° N 70.46065° E	Meadow–chestnut solonetzic saline soil	Ak (0–20)	Light loamy, dry, loose, upper 2 cm are abundantly intertwined with plant roots, brownish gray, lumpy–powdery, rounded sand grains of white and brown color, soil effervesces from HCl from the depth of 5 cm
			V1k (20–51)	Light loamy, fresh, whitish brown, streaks of humus, lumpy–nutty, carbonates in the form of impregnation, rounded sand grains of white and brown color, small plant roots, compacted, soil effervesces from HCl
			V2skg (51–89)	Medium loamy, fresh, compacted, whitish brown (due to salts and gypsum), nutty–prismatic, rounded sand grains of white and brown color, gypsum accumulations in the form of spots and tubes, salt crystals abundantly appear on the section wall upon drying, individual small roots, carbonates along pores and root passages in the form of small tubes, soil effervesces from HCl
			S1kg (89–124)	Lightly loamy, fresh, brown, rounded sand grains of white and brown color, sand grains are glued together by clay material into conglomerates, salt crystals abundantly appear on the section wall upon drying, structureless, sticky, streaked with small black dots of 0.3–0.5 mm (possibly manganese structures), there are no gypsum structures, soil effervesces from HCl
			S2k (124–…)	Light loamy, sticky, structureless, fresh, brown, rounded sand grains of white and brown color, the horizon contains a layer of sand mixed with loam, there are no gypsum structures, soil effervesces from HCl
TR2	50.54246° N 70.46014° E	Crustal saline solonetz	Ak (0–2)	Light loamy, gray, dry, powdery, intertwined with roots of steppe vegetation, loose, rounded sand grains of white and brown color, soil effervesces from HCl
			V1k (2–18)	Heavy loamy, brownish gray, compacted, nutty–columnar, plant roots, rounded sand grains of white and brown color, carbonates in the form of spots, dry, soil effervesces from HCl
			V2ksg (18–100)	Heavy loamy, wet, brown, sticky, grained, gypsum in the form of spots and tubes, rusty spots, the section wall is covered with salt crystals upon drying, rounded sand grains of white and brown color, loose, soil effervesces from HCl
AC	50.54190° N 70.45928° E	Shor solonchak	Crust (0–2)	Heavy loamy, gray, wet, loose, wall saltpeters, structureless, plastic, sticky, soil effervesces from HCl
			Skg (2–40)	Heavy loamy, brown, wet, loose, wall saltpeters, structureless, plastic, sticky, soil effervesces from HCl
Catena position	Coordinates	Soil	Horizons, cm	Description
Kazakhstan. Karaganda Region, environs of the settlement of Barshino. Lake depression
EL	49.62547° N 69.47282° E	Light-chestnut surface–stony soil	A (0–13)	Sandy loam, medium stony, fresh, light gray-brown, slightly compacted, lumpy powdery fine earth, plant roots, soil does not effervesce from HCl
			V (13–30)	Sandy loam, very stony, brown, denser than the previous one, the amount of solid structures increases sharply, the fine earth is organized into a lumpy–nutty structure, soil does not effervesce from HCl
TR1	49.62322° N 69.47160° E	Meadow chestnut surface–stony soil	Ad (0–5)	Sandy loam, intertwined with roots of steppe vegetation (approximately 40–50% of the horizon), fresh, gray, lumpy powdery fine earth, compacted by roots, there is no skeleton, soil does not effervesce from HCl
			A (5–14)	Sandy loam, fresh, gray, lumpy powdery fine earth, many plant roots, slightly compacted, there is no skeleton, soil does not effervesce from HCl
			AV (14–32)	Sandy loam, medium stony, fresh, brown, plant roots, fresh, lumpy fine earth, slightly compacted, soil does not effervesce from HCl
			VS (32–45)	Sandy loam, very stony, fresh, whitish brown, loosely lumpy fine earth, denser than the previous one, single plant roots, soil does not effervesce from HCl
TR2	49.62237° N 69.47198° E	Crustal surface–stony solonetz	A (0–3)	Sandy loam, slightly stony, dry, gray, lumpy powdery fine earth, a small amount of plant roots, soil does not effervesce from HCl
			V (3–12)	Sandy loam, slightly stony, fresh, grayish brown, columnar, many plant roots, dense, soil does not effervesce from HCl
			Sk (12–36)	Sandy loam, medium stony, structureless fine earth, fresh, brownish brown, single plant roots, less dense than the overlying horizon, soil effervesces from HCl
			Dk (36–…)	Rocky crimson mass mixed with fine earth of the same color, a small amount of small dead plant roots, soil effervesces from HCl
TR3	49.62034° N 39.47446° E	Crustal surface–stony solonetz	A (0–4)	Sandy loam, medium stony, fresh, cracking crust on the surface, loose, powdery fine earth, gray-brown, there are many plant roots, but they do not form turf, soil does not effervesce from HCl
			V (4–30)	Clayey, medium-stony, very dense, lumpy, brownish brown, fresh, dead plant roots, soil does not effervesce from HCl
AC	49.61977° N 69.47681° E	Shor solonchak	Ks (0–1)	Salt crust, soil effervesces from HCl
			Sgkg (1–30)	Clayey, brown, structureless, wet, plant roots are found in the upper part of the horizon, salt crystals appear on the section wall upon drying, soil effervesces from HCl

Appendix 2

Spring population of carabids and tenebrionids on the steppe catenas of central Kazakhstan in 1976–78 and in 2018 (the sum of specimens per 100 trap days). Legends: EL is eluvial, TR is transit, AC is accumulative catena position; L is the latitudinal group of ranges: B is boreal, SH is subboreal humid, SA is subarid, PZ is polyzonal.

A. Subzone of the typical arid steppe, Shortandy
	Beetle species	1976			1978			2018
		positions
		EL	TR	AC	EL	TR	AC	EL	TR	AC
L	Carabidae
B	Agonum fuliginosum	–	–	3	–	–	–	–	–	3
PZ	Agonum thoreyi	–	–	–	–	–	–	–	–	3
B	Agonum viduum	–	–	–	–	–	–	–	–	5
PZ	Amara aenea	2	–	–	–	1	–	–	–	–
B	Amara infima	–	–	–	3	–	–	–	–	–
SA	Amara pastica	1	–	–	–	–	–	–	–	–
B	Amara plebeja	–	–	1	–	–	–	–	–	–
SH	Amara tibialis	–	18	1	–	14	5	–	–	–
PZ	Anisodactylus binotatus	–	–	–	–	–	–	–	–	3
SH	Badister unipustulatus	–	1	–	–	–	5	–	–	–
SA	Bembidion varium	–	–	–	–	–	–	–	–	3
SH	Bembidion biguttatum	–	–	–	–	–	–	–	–	8
B	Bembidion humerale	–	–	11	–	–	–	–	–	–
SH	Bembidion minimum	–	–	–	–	–	–	–	–	5
SH	Bembidion octomaculatum	–	–	35	–	–	30	–	–	–
SA	Bembidion pallidiveste (?)	–	–	6	–	–	9	–	–	–
PZ	Bembidion properans	1	144	2	–	–	4	–	5	–
PZ	Bembidion quadrimaculatum	–	1	–	–	3	–	–	–	–
SH	Bembidion quadripustulatum	–	–	–	–	–	–	–	–	3
B	Bembidion transparens	–	1	41	–	10	6	–	–	–
SH	Bradycellus caucasicus	–	–	60	–	–	7	–	–	–
B	Calosoma investigator	–	–	–	–	2	–	–	–	–
SH	Carabus clathratus	–	–	1	–	–	–	–	–	8
SA	Carabus cribellatus	4	3	–	3	–	–	–	3	–
SA	Carabus estreicheri	1	–	–	–	–	–	–	–	–
PZ	Carabus granulatus	–	–	–	–	–	–	–	–	5
B	Chlaenius nigricornis	–	1	6	–	3	–	–	–	10
SA	Chlaenius spoliatus	–	–	–	–	–	–	–	–	10
PZ	Chlaenius tristis	–	–	–	–	–	–	–	–	5
PZ	Clivina fossor	–	–	–	–	–	–	–	–	5
SH	Curtonotus aulicus	–	–	–	–	–	–	–	–	15
SH	Curthonotus castaneus	–	–	–	–	–	–	–	3	–
SH	Curtonotus convexiusculus	–	–	–	2	–	–	–	–	–
SH	Cymindis angularis	1	–	–	–	2	–	–	–	–
SA	Cymindis lateralis	1	–	–	–	–	–	–	–	–
SA	Dyschiriodes rufipes	3	2	–	28	3	–	–	–	–
–	Dyschiriodes sp.	–	–	–	–	–	–	–	2	–
SH	Elaphrus cupreus	–	–	–	–	–	–	–	–	10
SA	Harpalus amplicollis	5	–	–	3	–	–	–	2	–
SA	Harpalus anxius	–	–	–	–	5	5	–	5	–
SA	Harpalus calathoides	1	–	–	5	2	–	–	–	–
PZ	Harpalus distinguendus	–	–	–	–	2	–	–	–	–
SA	Harpalus modestus	1	–	–	–	–	–	–	–	–
	Beetle species	1976			1978			2018
		positions
		EL	TR	AC	EL	TR	AC	EL	TR	AC
SA	Harpalus politus	–	–	–	3	–	–	–	2	–
SA	Harpalus pusillus (?)	1	–	–	–	–	–	–	–	–
SH	Harpalus smaragdinus	–	1	–	–	–	–	–	8	–
SA	Harpalus subcylindricus	–	–	–	–	–	–	–	3	–
SH	Harpalus tardus	1	–	–	–	–	–	–	–	–
SA*	Harpalus tenebrosus	–	–	–	2	–	–	–	–	–
SA	Microlestes fissuralis	–	–	–	–	–	–	3	–	–
SA	Microlestes maurus	–	–	–	–	–	–	–	10	–
PZ	Microlestes minutulus	33	81	22	25	130	85	–	15	–
SH	Oodes helopioides	–	–	–	–	–	–	–	–	18
SH	Oxypselaphus obscurus	–	–	–	–	–	–	–	–	5
SH	Poecilus cupreus	–	2	29	–	–	4	–	–	3
SH	Poecilus punctulatus	–	18	–	–	2	1	–	–	–
SA	Poecilus sericeus	5	1	–	2	–	–	13	5	–
SA	Poecilus subcoeruleus	–	–	–	–	2	–	–	–	–
PZ	Poecilus versicolor	–	–	–	–	2	4	–	3	–
SA	Pseudotaphoxenus rufitarsis	–	–	–	–	–	–	3	–	–
SH	Pterostichus anthracinus	–	–	–	–	–	–	–	–	25
SH	Pterostichus gracilis	–	–	–	–	–	–	–	–	8
SH	Pterostichus macer	1	5	–	5	16	–	–	–	–
B	Pterostichus minor	–	3	26	–	–	10	–	–	7
PZ	Pterostichus niger	–	–	4	–	–	–	–	3	5
PZ	Pterostichus nigrita	–	–	3	–	–	–	–	–	30
PZ	Pterostichus strenuus	–	–	–	–	–	2	–	–	3
SH	Pterostichus vernalis	–	–	–	–	–	5	–	–	3
PZ	Stenolophus mixtus	–	–	–	–	–	–	–	–	5
SA	Microlestes plagiatus	–	–	–	–	–	10	–	–	–
SH	Syntomus truncatellus	2	17	–	45	13	–	3	–	–
SA	Taphoxenus gigas	1	–	–	–	–	–	3	–	–
	Total number of Carabidae species	65	299	251	126	212	192	25	69	213
	Total number of Carabidae individuals	18	16	16	12	17	16	5	14	27
	Menhinick’s index	2.2	0.9	1.0	1.1	1.2	1.2	1.0	1.7	1.8
	Tenebrionidae
	Blaps halophila	3	5	–	3	2	–	–	–	–
	Blaps lethifera	2	–	–	–	1	–	–	–	–
	Centorus procerus moldaviensis									1
	Crypticus quisquilius	3	–	–	–	15	–	–	–	–
	Gonocephalum granulatum pusillum	12	2	–	16	8	2	5	107	–
	Opatrum riparium	–	36	–	3	82	5	–	–	–
	Opatrum sabulosum	11	–	–	96	18	2	–	23	–
	Pedinus femoralis	6	–	–	17	2	–	5	18	–
	Platyscelis hypolitha	–	–	–	2	–	–	–	–	–
	Tentyria nomas	2	–	–	–	–	–	15	5	–
	Total number of Tenebrionidae species	7	3	–	6	7	3	3	4	–
	Total number of Tenebrionidae individuals	39	43	–	137	198	9	25	153	–
	Menhinick’s index	1.1	0.5	–	0.5	0.6	1.0	0.6	0.3	–

B. Dry steppe, Arykty
	Beetle species	1976			1978			2018
		positions
		EL	TR	AC	EL	TR	AC	EL	TR	AC
L	Carabidae
B	Amara infima	–	–	–	–	7	–	–	–	–
B	Amara littorea	–	–	4	–	–	6	–	–	–
SA	Amara pastica	4	–	–	–	–	–	–	–	–
SH	Amara tibialis	–	–	–	–	14	3	–	–	–
SA	Brachinus hamatus	–	–	–	–	–	3	–	47	40
SH	Broscus semistriatus	–	–	–	–	–	–	–	3	5
SH	Calosoma auropunctatum	–	6	–	–	–	–	–	–	–
SH	Calosoma denticolle	10	–	–	–	–	–	–	–	–
SH	Carabus clathratus	–	10	6	–	3	5	–	–	–
SA	Corsyra fusula	–	–	–	–	–	–	20	–	–
SA	Harpalus anxius	–	–	–	–	–	–	–	3	–
PZ	Harpalus distinguendus	–	–	–	–	8	–	–	–	–
SA	Harpalus sarmaticus	–	–	–	–	–	–	7	–	–
SH	Harpalus smaragdinus	–	–	–	–	–	–	10	–	–
SA	Harpalus subcylindricus	–	–	–	–	–	–	3	–	–
PZ	Microlestes minutulus	–	70	34	6	328	208	–	–	–
SA	Microlestes plagiatus	–	10	6	–	–	–	–	–	–
SA	Microlestes schroederi	–	–	–	–	–	–	7	–	–
SH	Poecilus cupreus	–	–	58	–	–	6	–	–	–
SH	Poecilus punctulatus	–	–	–	–	10	–	–	–	–
SA	Poecilus sericeus	36	–	–	18	–	–	30	7	3
SA	Poecilus subcoeruleus	–	16	6	–	33	3	–	–	–
SH	Pterostichus macer	1	62	60	–	20	23	–	–	–
B	Pterostichus minor	–	–	4	–	7	15	–	–	–
PZ	Pterostichus nigrita	–	–	6	–	–	3	–	–	–
SH	Syntomus truncatellus	–	–	–	–	–	–	3	30	3
SA	Taphoxenus gigas	16	–	–	3	–	–	10	7	–
	Total number of Carabidae species	4	6	9	3	9	10	8	6	4
	Total number of Carabidae individuals	60	174	184	27	430	275	9	97	51
	Menhinick’s index	0.5	0.5	0.7	0.6	0.4	0.6	0.8	0.6	0.6
	Tenebrionidae
	Blaps halophila	40	–	–	52	–	–	30	10	7
	Blaps lethifera	–	–	–	–	–	–	27	3	–
	Centorus rufipes									1
	Crypticus quisquilius	–	3	5	2	5	2	–	–	–
	Gonocephalum granulatum pusillum	–	3	–	26	382	125	–	7	–
	Gonocephalum pygmaeum	7	25	10	–	–	–	–	–	–
	Oodescelis polita	–	5	–	5	3	–	3	3	–
	Opatrum riparium	–	–	–	–	–	–	–	–	–
	Opatrum sabulosum	–	–	–	10	6	3	–	3	–
	Pedinus femoralis	6	–	–	–	–	–	20	3	–
	Platyscelis hypolitha	–	–	–	6	–	–	3	–	–
	Tentyria nomas	234	47	17	167	10	3	460	153	44
	Total number of Tenebrionidae species	4	6	3	7	5	4	6	7	2
	Total number of Tenebrionidae individuals	287	83	32	268	406	133	543	182	51
	Menhinick’s index	0.2	0.7	0.5	0.4	0.3	0.3	0.3	0.5	0.3

C. Deserted steppe, Barshyn (Barshino)
	Beetle species	1976			1978			2018
		positions
		EL	TR	AC	EL	TR	AC	EL	TR	AC
L	Carabidae
SH	Agonum viridicupreum	–	–	30	–	–	8	–	–	–
–	Amara (Bradytus) sp.	–	6	–	–	–	–	–	–	–
SA	Bembidion pallidiveste (?)	–	–	–	–	–	10	–	–	–
SA	Brachinus hamatus	–	–	200	–	–	–	–	–	–
SH	Calosoma auropunctatum	–	–	6	–	–	8	–	–	–
SH	Calosoma denticolle	–	6	–	–	–	–	–	–	4
SA	Carabus bessarabicus	–	–	6	–	16	–	–	8	–
SA	Chlaenius spoliatus	–	–	30	–	–	–	–	–	–
PZ	Chlaenius tristis	–	–	–	–	–	30	–	–	–
SH	Curtonotus convexiusculus	–	10	–	–	–	–	–	–	–
SA	Cymindis lateralis	–	10	60	–	–	–	–	–	–
SA*	Cymindis violacea (?)	30	–	–	4	–	–	–	–	–
SH	Elaphrus cupreus	–	–	–	–	–	60	–	–	–
SA	Harpalus dispar splendens	–	–	–	–	–	–	–	–	8
SA	Harpalus salinus	14	–	–	16	–	–	–	–	–
SH	Harpalus smaragdinus	–	4	–	–	–	–	–	–	–
SA	Harpalus steveni	–	12	–	–	–	–	–	–	–
SA	Microlestes maurus	–	–	–	–	–	–	–	4	–
PZ	Microlestes minutulus	–	–	80	–	–	10	–	–	–
SA	Microlestes plagiatus	–	–	–	–	–	84	–	–	–
SH	Ophonus rufibarbis	–	6	–	–	–	–	–	–	–
SA	Poecilus crenuliger	–	–	–	–	–	–	–	20	–
SH	Poecilus cupreus	–	–	–	–	–	50	–	–	–
SA	Poecilus laevicollis	–	–	–	–	–	–	–	–	8
SA	Poecilus sericeus	–	–	–	–	–	–	–	12	–
SA	Poecilus subcoeruleus	–	70	–	–	58	392	–	–	–
SA	Pseudotaphoxenus tillesii	–	6	–	–	–	–	–	–	–
SH	Pterostichus macer	–	6	24	–	–	16	–	–	–
SA	Taphoxenus gigas	4	–	–	8	–	–	1	4	–
	Total number of Carabidae species	3	10	8	3	2	10	1	5	3
	Total number of Carabidae individuals	48	136	436	28	74	668	1	48	20
	Menhinick’s index	0.4	0.9	0.4	0.6	0.2	0.4	1.0	0.7	0.7
	Tenebrionidae
	Anatolica lata	5	6	–	124	–	–	4	16	–
	Blaps halophila	14	–	–	26	8	–	16	12	24
	Blaps lethifera	–	–	–	–	–	–	8	12	–
	Centorus calcaroides intermedius									1
	Centorus filiformis	–	–	–	–	8	–	–	–	–
	Centorus procerus	–	10	6	–	–	16	–	–	–
	Oodescelis polita	–	70	–	–	50	–	–	4	–
	Opatrium sabulosum	–	–	–	–	8	–	–	8	–
	Pedinus femoralis	10	6	–	8	8	–	–	8	–
	Platyscelis rugifrons	–	–	–	–	–	–	–	12	–
	Platyscelis sp.	200	6	–	54	–	–	–	–	–
	Tentyria nomas	30	6	–	50	8	–	72	20	–
	Total number of Tenebrionidae species	5	6	1	5	6	1	4	8	2
	Total number of Tenebrionidae individuals	308	104	6	262	90	16	100	92	24
	Menhinick’s index	0.3	0.6	0.4	0.3	0.6	0.2	0.4	0.8	0.2

1. * With the transition to the arid group of species.