Salt lakes of La Mancha (Central Spain): A hot spot for tiger beetle (Carabidae, Cicindelinae) species diversity

Abstract

The tiger beetle assemblage of the wetlands of La Mancha (central Spain) comprises nine species: Calomera littoralis littoralis, Cephalota maura maura, Cephalota circumdata imperialis, Cephalota dulcinea, Cicindela campestris campestris, Cicindela maroccana, Cylindera paludosa, Lophyra flexuosa flexuosa, and Myriochila melancholica melancholica. This assemblage represents the largest concentration of tiger beetles in a single 1º latitude / longitude square in Europe. General patterns of spatial and temporal segregation among species are discussed based on observations of 1462 specimens registered during an observation period of one year, from April to August. The different species of Cicindelini appear to be distributed over space and time, with little overlapping among them. Three sets of species replace each other phenologically as the season goes on. Most of the species occupy drying or dried salt lakes and salt marshes, with sparse vegetation cover. Spatial segregation is marked in terms of substrate and vegetation use. Calomera littoralis and Myriochila melancholica have been observed mainly on wet soils; Cephalota circumdata on dry open saline flats; Cephalota dulcinea and Cylindera paludosa in granulated substrates with typical halophytic vegetation; Cephalota maura is often present in man-modified areas. Cephalota circumdata and Cephalota dulcinea are included as species of special interest in the list of protected species in Castilla-La Mancha. Conservation problems for the Cicindelini assemblage arise from agricultural activities and inadequate use of sport vehicles. Attempts at restoring the original habitat, supressing old semi-industrial structures, may affect the spatial heterogeneity of the lakes, and have an effect on Cicindelinae diversity.

Keywords: Behaviour; Coleoptera; Conservation; Habitat selection; Lakes; Phenology; Salt marshes.

Figure 1.

Map of the Iberian Peninsula showing the location of the study area. The inset shows a detail of the 27 marshes and small lakes included in the Life Project “La Mancha Wetlands” (1–27), and some other monitored marshes and lakes (28–32). Numbers represent the sampled areas: TOLEDO: Lillo: 1 Laguna de El Longar 2 Laguna del Altillo Chica 3 Laguna del Altillo Grande 4 Laguna de la Albardiosa; Miguel Esteban: 31 La Laguna de Miguel Esteban; Villacañas: 5 Laguna Larga 6 Laguna de Tirez 7 Laguna de Peña Hueca; Quero: 8 Laguna del Taray 9 Laguna Grande 29 Cerro de San Cristóbal; Villafranca de Los Caballeros: 10 Laguna Chica 11 Laguna Grande 12 Laguna de la Sal 32 Gigüela marshes. CIUDAD REAL: Alcázar de San Juan: 13 Laguna de los Carros 14 Laguna de Pajares/los Pájaros 15 Laguna de las Yeguas 16 Laguna del Camino de Villafranca 17 Laguna de la Veguilla; Campo de Criptana: 18 Laguna de Salicor 30 Arroyo de San Marcos; Pedro Muñoz: 19 Laguna del Retamar 20 Laguna del Pueblo/de la Vega 21 Laguna de Navalafuente 22 Laguna de Alcahozo. CUENCA: Mota del Cuervo: 23 Laguna de Manjavacas 24 Laguna de Sánchez Gómez 25 Laguna de la Dehesilla; Las Mesas/Las Pedroñeras: 26 Laguna de Alcahozo Chico 27 Laguna del Taray Chico. El Hito: 28 Laguna del Hito.

Figure 2.

Representative specimens of tiger beetles from La Mancha wetlands A Calomera littoralis littoralis (La Dehesilla, Cuenca) B Cicindela campestris campestris (La Dehesilla, Cuenca) C Cicindela maroccana (La Sal, Toledo) D Cylindera paludosa (El Longar, Toledo) E Lophyra flexuosa flexuosa (El Pardo, Madrid) F Myriochila melancholica melancholica (Tirez, Toledo). Photographs by MG-P.

Figure 3.

Representative specimens of Cephalota from La Mancha wetlands. A Cephalota (Taenidia) dulcinea (Sánchez Gómez, Cuenca) B Cephalota (Cassolaia) maura (Arroyo San Marcos, Ciudad Real) C Cephalota (Taenidia) circumdata imperialis (Tirez, Toledo). Photographs by MG-P.

Figure 4.

Maps showing the location of species of Cicindelinae found in the wetlands of La Mancha. Numbers correspond to Fig. 1. A Calomera littoralis littoralis B Cephalota (Cassolaia) maura C Cephalota (Taenidia) circumdata imperialis D Cephalota (Taenidia) dulcinea E Cicindela campestris campestris F Cylindera paludosa.

Figure 5.

Representative habitats of Cicindelinae in La Mancha. A Laguna de la Sal (Toledo) in May. Salt flats occupied by Calomera littoralis littoralis B Laguna de Peña Hueca (Toledo) in April. Wet salt flats, with gypsum outcrops, occupied by Calomera littoralis littoralis; when totally dry, in summer, occupied by Cephalota circumdata imperialis C Laguna de Las Yeguas (Ciudad Real) in June. Salt flats, occupied by Calomera littoralis littoralis and Cephalota circumdata imperialis; and halophytic prairies occupied by Cephalota dulcinea and Cylindera paludosa D La Laguna de Miguel Esteban (Toledo) in July. Salt flats, occupied by Calomera littoralis littoralis; and halophytic prairies with “albardinal” occupied by Cephalota maura maura E Laguna del Camino de Villafranca (Ciudad Real), with flamingos, in July. Sapropel shores occupied by Calomera littoralis littoralis F Laguna de Alcahozo (Ciudad Real) in June. Salt flats, occupied by Calomera littoralis littoralis and Cephalota circumdata imperialis; and open areas near dry halophytic prairies occupied by Cephalota dulcinea and also by Cephalota circumdata imperialis. Photographs by N. Percino and MG-P.

Figure 6.

Representative habitats of Cicindelinae in La Mancha. A Channel at Laguna de las Yeguas (Ciudad Real) in May. Wet areas occupied by Calomera littoralis littoralis and Cicindela campestris; when drier, occupied by Cephalota maura maura B Road side pools and halophytic vegetation in Laguna de Manjavacas (Cuenca) in April. Calomera littoralis littoralis was the only species observed C Wells and deep pools in Laguna de Manjavacas (Cuenca) in June. Cephalota maura maura is frequent in these structures D Laguna del Pueblo de Pedro Muñoz (Ciudad Real) in July. Mud areas amongst hydrophytic matts, occupied by Calomera littoralis littoralis and Myriochila melancholica melancholica E Temporary pools at Laguna del Longar (Toledo), in July. Calomera littoralis littoralis, Myriochila melancholica melancholica and Cephalota maura co-occcur at the sides F Laguna del Taray Chico (Cuenca) in May. Trails and non-halophytic prairies near the fresh water lake, occupied by Cicindela campestris campestris. Photographs by N. Percino and MG-P.

Figure 7.

Observations of Calomera littoralis littoralis at Laguna de Manjavacas (left) and La Dehesilla (upper right). Colours indicate the month in which observations were made (see legend). Note seasonal changes in specimen’s location as the water front retreats or following changes in humidity of the soil. Blue squares correspond to the habitat shown in Fig. 6b.

Figure 8.

Frequency of observation of Cicindelinae during the sampling period (April to August), all sites pooled. From top to bottom: Calomera littoralis littoralis, Cephalota maura maura, Cicindela campestris campestris, and Myriochila melancholica melancholica.

Figure 9.

Frequency of observation of Cicindelinae during the sampling period (April to August), all sites pooled. From top to bottom: Cephalota circumdata imperialis, Cephalota dulcinea, and Cylindera paludosa.

Figure 10.

Behavioural observations on Cephalota circumdata imperialis. A, B A few specimens of Cephalota circumdata located at the shade of a rock in the middle of the exposed salt flat C Individual photographed on the wet mud under the remains of a plastic bag, also in the salt flat. This behaviour is atypical for the species, which is generally active at full sunlight in mid summer in the area. The observations were made the 13th of June, at mid-day, the first date in which activity of the species was recorded. It is possibly that the specimens had just eclosed and wait in the shade while hardening their integument. Photographs by MG-P.

Figure 11.

Characteristic habitat of the La Mancha endemic tiger-beetle Cephalota (Taenidia) dulcinea. A Trails at laguna del Taray (Toledo) B Trails and halophytic vegetation in Laguna de Pajares (Ciudad Real), where it was found in company of Cephalota maura maura C Trails and “albardinal” in Laguna del Altillo Chica (Toledo), where it co–occurs with Cylindera paludosa D Open areas amongst halophytic vegetation at laguna de La Sal (Toledo), Calomera littoralis littoralis and Cicindela maroccana occur in this habitat at the end of the spring E Halophytic vegetation at Laguna de Peña Hueca (Toledo), in this spot, Calomera littoralis littoralis and Cicindela maroccana are present at the end of the spring F Halophytic vegetation and open areas with salty granulated soils at Laguna de Sánchez Gómez (Cuenca). Photographs MG-P.

Figure 12.

Behavioural observations on Cephalota dulcinea A Specimen of Cephalota dulcinea located in a trail devouring a winged hymenopteran. Note the number of insect remains located behind the specimen (ants, other Hymenoptera and Coleoptera), suggesting that the spot is used as a hunting post B A couple of Cephalota dulcinea involved in courship C A recently killed specimen of Cephalota dulcinea in a trail, about to be seized by ants D Trail located near laguna de Pajares with a large colony of Cephalota dulcinea, Cephalota maura maura and scattered Cylindera paludosa. We hypothesize that these trails, despite of being surrounded by unfavourable habitat, might be used as dispersal corridors between lakes, facilitating the maintenance of metapopulation systems. Photographs by MG-P.

Figure 13.

Conservation problems for tiger beetles in La Mancha, and remains of old semi-industrial activities which provide additional structural heterogenity A Saline pools at Laguna de Peña Hueca (Toledo) occupied at different seasons by Calomera littoralis littoralis, Cephalota dulcinea, Cephalota circumdata imperialis and Cephalota maura maura B Trails on salty soils at Laguna de la Dehesilla (Cuenca), where Calomera littoralis littoralis, Cicindela campestris campestris and Cephalota dulcinea are present along different periods of the year C Old channel in Laguna del Longar (Toledo), where Cephalota maura maura, Cicindela campestris and Cylindera paludosa co-occur D Saline pool at Laguna de Tirez (Toledo), where Calomera littoralis littoralis and Myriochila melancholica melancholica co-occur, while populations of Cephalota dulcinea and Cephalota circumdata imperialis are established not far from this point E Effect of sheep along the shores of the saline Laguna Grande de Quero (Toledo), in this spot Calomera littoralis littoralis was present F Vineyards at the edge of Laguna de Alcahozo Chico (Cuenca), note the diverse halophytic vegetation and nearby open areas where Calomera littoralis littoralis, Cicindela campestris campestris, Cylindera paludosa, and Lophyra flexuosa flexuosa live in close proximity. Photographs by N. Percino and MG-P.

Figure 14.

Location of seven species of Cicindelinae at Laguna del Longar (Lillo, Toledo). Colour dots indicate the spatial position of each specimen (often, more than one specimen are presented under a single dot) (see legend). Note the spatial segregation of most species. Dots in lower right area, where Myriochila melancholica melancholica was found, correspond to the habitat shown in Fig. 6e. Dots in the upper left area, where Cicindela campestris campestris was present, correspond to the habitat shown in Fig. 13c.