Research Article |
Corresponding author: Mario Garcia-Paris ( mparis@mncn.csic.es ) Academic editor: Pavel Stoev
© 2016 Paula Rodriguez-Flores, Jorge Gutierrez-Rodriguez, Ernesto F. Aguirre-Ruiz, Mario Garcia-Paris.
This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Citation:
Rodríguez-Flores PC, Gutiérrez-Rodríguez J, Aguirre-Ruiz EF, García-París M (2016) Salt lakes of La Mancha (Central Spain): A hot spot for tiger beetle (Carabidae, Cicindelinae) species diversity. ZooKeys 561: 63-103. https://doi.org/10.3897/zookeys.561.6042
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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 M. melancholica have been observed mainly on wet soils; C. circumdata on dry open saline flats; C. dulcinea and C. paludosa in granulated substrates with typical halophytic vegetation; C. maura is often present in man-modified areas. Cephalota circumdata and C. 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.
Coleoptera , Habitat selection, Phenology, Behaviour, Conservation, Salt marshes, Lakes
The geomorphology of the Iberian Peninsula has been modelled through a long and complex paleogeographic history, including the ancient emergence of the western territories, complex Miocene reorganization of the tectonic plates with active orogenic periods and incorporation of a large portion of the Betic–Rifean massif, and the effect of the Pleistocene glacial periods (
The Central Iberian Plateau, is an almost flat elevated plain which occupies most of the central area of the Peninsula. The Iberian Plateau, is delimited by mountain chains of different height all around its perimeter, and crossed transversally by the Central System Mountain Chain, which divides it into two sub-regions: a northern one, mostly conformed by the Duero River Basin, located in the Spanish Autonomous Region of Castilla – León, and a southern one crossed by the Tagus and Guadiana river systems, in the Spanish Autonomous Region of Castilla – La Mancha (
Flat, sun exposed areas surrounded by halophytic vegetation, create a typical habitat for tiger beetles (Cicindelinae) (
The tiger beetle fauna of the Iberian Peninsula is considered well-known in relation to the knowledge on other taxonomic groups.
Previous reports on the Cicindelinae of La Mancha lacustrine areas (
In this study, we monitored, from April to August, the presence and activity of tiger beetles in more than 30 small lakes, salt marshes, and gypsum flats in La Mancha region. Most of these locations (27) were monitored as part of a Life Project: La Mancha Wetlands (LIFE+10 NAT/ES/000563), aimed at restoring the espartal or albardinal (Lygeum spartum (L.) Kunth) grasslands and salt flats, by reclaiming Mediterranean salt steppes (Limonietalia), designated as a priority habitat (1510) by Habitats Directive 92/43/EEC, and other halophytic formations in the La Mancha wetlands SCI and SPA zones. Most of the locations studied are protected under the laws of the Autonomous Region of Castilla – La Mancha (Spain), and included in the Nature 2000 European Union conservation program (http://ec.europa.eu/environment/nature/natura2000/db_gis/).
Our main objective was to determine the relevance of the lacustrine environments of La Mancha region for tiger beetle diversity and conservation. During the study we found that the number of species inhabiting the region had been underestimated. In comparison to previously published European locality records, La Mancha region harbours the largest number of species of tiger beetles present in a single square (one degree latitude x longitude). This finding prompted us to undertake the following tasks: (1) inventory the tiger beetle species inhabiting each lake or marsh, (2) identify the phenology for each species during the period of study, (3) gather information on general landscape use for each species, and (4) evaluate threats and possible conservation needs, based on observed abundances and observations on human activities around the lakes.
The study was carried on the 27 marshes and small lakes included in the Life Project La Mancha Wetlands, all located in the traditional demarcation of La Mancha, which include portions of Ciudad Real, Cuenca, and Toledo provinces (Fig.
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.
Localities monitored, geographic coordinates and sampling dates for each locality.
Wetland | Town and province | Coordinates | Date |
---|---|---|---|
Laguna de El Longar | Lillo (Toledo) | 39°42'8.00"N, 3°19'17.86"W | 15-VI-2012, 17-VI-2012, 13-IV-2014, 23-V-2014, 13-VI-2014, 11-VII-2014, 2-VIII-2014, 9-VIII-2014, 23-VIII-2014 |
Laguna del Altillo Chico | Lillo (Toledo) | 39°42'4.57"N, 3°18'11.38"W | 23-V-2014, 7-VI-2014, 13-VI-2014, 2-VIII-2014 |
Laguna del Altillo Grande | Lillo (Toledo) | 39°41'33.10"N, 3°18'1.03"W | 17-V-2014, 23-V-2014, 7-VI-2014, 13-VI-2014 |
Laguna de la Albardiosa | Lillo (Toledo) | 39°39'38.75"N, 3°17'32.35"W | 23-V-2014 |
Laguna Larga | Villacañas (Toledo) | 39°36'17.45"N, 3°19'1.58"W | 26-IV-2014, 17-V-2014, 23-V-2014, 13-VI-2014, 20-VI-2014, 27-VI-2014, 9-VIII-2014, 23-VIII-2014 |
Laguna de Tirez | Villacañas (Toledo) | 39°32'6.85"N, 3°21'26.73"W | 17-VI-2012, 13-IV-2014, 17-V-2014, 20-VI-2014, 27-VI-2014, 2-VIII-2014, 23-VIII-2014, 24-X-2014 |
Laguna de Peña Hueca | Villacañas (Toledo) | 39°30'50.58"N, 3°20'24.98"W | 13-IV-2014, 26-IV-2014, 17-V-2014, 23-V-2014, 13-VI-2014, 27-VI-2014, 11-VII-2014, 2-VIII-2014, 9-VIII-2014, 23-VIII-2014, 24-X-2014 |
Laguna del Taray | Quero (Toledo) | 39°31'48.62"N, 3°18'59.79"W | 26-IV-2014, 7-VI-2014, 21-VI-2014 |
Laguna Grande | Quero (Toledo) | 39°29'59.25"N, 3°15'12.42"W | 26-IV-2014, 24-V-2014, 21-VI-2014, 28-VI-2014, 12-VII-2014 |
Laguna Chica | Villafranca de los Caballeros (Toledo) | 39°27'51.61"N, 3°19'56.61"W | 26-IV-2014, 20-VI-2014, 27-VI-2014 |
Laguna Grande | Villafranca de los Caballeros (Toledo) | 39°27'3.42"N, 3°20'14.67"W | 26-IV-2014, 20-VI-2014, 27-VI-2014, 11-VII-2014, 2-VIII-2014, 9-VIII-2014 |
Laguna de la Sal | Villafranca de los Caballeros (Toledo) | 39°26'14.71"N, 3°19'54.51"W | 26-IV-2014, 24-V-2014, 11-VII-2014 |
Laguna de los Carros | Alcázar de San Juan / Quero | 39°28'12.39"N, 3°15'45.30"W | 8-VI-2012, 26-IV-2014, 24-V-2014, 21-VI-2014 |
Laguna de Pajares | Alcázar de San Juan (Ciudad Real) | 39°27'15.47"N, 3°12'20.83"W | 25-IV-2014, 17-V-2014, 7-VI-2014 |
Laguna de las Yeguas | Alcázar de San Juan (Ciudad Real) | 39°25'0.58"N, 3°16'58.85"W | 26-IV-2014, 17-V-2014, 24-V-2014, 20-VI-2014, 27-VI-2014, 11-VII-2014, 27-VII-2014, 2-VIII-2014, 9-VIII-2014 |
Laguna del Camino de Villafranca | Alcázar de San Juan (Ciudad Real) | 39°24'52.88"N, 3°15'30.63"W | 26-IV-2014, 11-VII-2014, 27-VII-2014, 2-VIII-2014, 9-VIII-2014 |
Laguna de la Veguilla | Alcázar de San Juan (Ciudad Real) | 39°23'45.37"N, 3°14'24.14"W | 26-IV-2014, 11-VII-2014, 8-VIII-2014, 9-VIII-2014 |
Laguna de Salicor | Campo de Criptana (Ciudad Real) | 39°28'0.24"N, 3°10'24.78"W | 8-VI-2012, 13-IV-2014, 17-V-2014, 6-VI-2014, 7-VI-2014 , 28-VI-2014, 8-VIII-2014 |
Laguna de Retamar | Pedro Muñoz (Ciudad Real) | 39°25'42.06"N, 2°58'20.98"W | 25-IV-2014, 21-VI-2014 |
Laguna del Pueblo | Pedro Muñoz (Ciudad Real) | 39°24'39.52"N, 2°56'47.20"W | 25-IV-2014, 6-VI-2014, 28-VI-2014, 12-VII-2014, 8-VIII-2014 |
Laguna de Navalafuente | Pedro Muñoz (Ciudad Real) | 39°25'31.02"N, 2°55'44.17"W | 25-IV-2014 |
Laguna de Alcahozo | Pedro Muñoz (Ciudad Real) | 39°23'27.46"N, 2°52'31.21"W | 25-IV-2014, 16-V-2014, 6-VI-2014, 21-VI-2014 , 8-VIII-2014 |
Laguna de Manjavacas | Mota del Cuervo (Cuenca) | 39°25'0.21"N, 2°51'55.06"W | 25-IV-2014, 16-V-2014, 21-VI-2014, 28-VI-2014, 12-VII-2014, 8-VIII-2014 |
Laguna de Sánchez Gómez | Mota del Cuervo (Cuenca) | 39°25'55.79"N, 2°50'22.26"W | 25-IV-2014, 16-V-2014, 6-VI-2014, 21-VI-2014, 28-VI-2014, 8-VIII-2014, |
Laguna de la Dehesilla | Mota del Cuervo (Cuenca) | 39°25'20.28"N, 2°50'25.51"W | 16-V-2014, 6-VI-2014, 20-VI-2014, 8-VIII-2014 |
Laguna de Alcahozo chico | Mota del Cuervo (Cuenca) | 39°23'26.99"N, 2°52'32.64"W | 25-IV-2014, 16-V-2014, 6-VI-2014, 12-VII-2014 |
Laguna del Taray Chico | Las Mesas / Las Pedroñeras (Cuenca) | 39°24'30.19"N, 2°42'55.98"W | 25-IV-2014, 16-V-2014, |
Laguna de El Hito | Montalbo / El Hito (Cuenca) | 39°52'4.22"N, 2°41'34.02"W | 15-VI-2014 |
Cerro San Cristóbal | Quero (Toledo) | 39°29'16.10’’N 3°15'58.62’’W | 8-VI-2012 , 24-V-2014, 21-VI-2014 |
Arroyo de San Marcos | Campo de Criptana (Ciudad Real) | 39°27'13.66’’N 3°09'37.29’’W | 8-VI-2012, 17-V-2014, |
La Laguna | Miguel Esteban (Toledo) | 39°32'19.15’’N 3°08'03.70’’W | 12-VII-2014 |
Gigüela marshes | Villafranca de los Caballeros (Toledo) | 39°26'09.36’’N 3°18'52.00’’W | 24-V-2014 |
The temporary lakes of La Mancha are a group of wetlands spread across the territory of the Upper Guadiana Basin. According to
These singularities – fluctuating saline lakes or hyper-saline lakes, with salt concentrations that can exceed those of seawater, and vegetation adapted to fairly restrictive conditions – make these wetlands and their surroundings particularly interesting for biodiversity conservation. These areas harbour a large number of endemic or endangered halophytic plants [i.e. Limonium carpetanicum Erben, Limonium dichotomum(Cav.) Kuntze Revis., Limonium squarrosum Erben, Gypsophila tomentosa L., Lygeum spartum, Microcnemum coralloides (Loscos & J.Pardo) Font Quer] and aquatic plants [Lamprothamnium papulosum (K.Wallroth) J.Groves, Tolypella salina R. Corillion, Althenia orientalis (Tzvelev) P.García-Murillo and Talavera, and Ruppia drepanensis Tineo ex Guss.] (
The lakes of La Mancha and the protective steppes that surround them, provide extensive ecosystem services (
Tiger beetle sampling and monitoring was designed to cover the period in which most saline lakes are totally or partially dry, from April to September. Monitoring in 2014 consisted in ½ to 1 hour walks for visual census in favourable habitats within marshes and lakes, including halophytic vegetation, shores of lakes, denuded salt flats, dirt roads with or without saline surfaces, draining channel banks, and margins of cultivated fields. Walks were carried out by one or two people (occasionally more) (MG-P, PCR-F, and collaborators mentioned in the acknowledgments). In particular situations (e.g. behavioural observations) walks took longer than 1 hour. At a first stage, we established transects to evaluate abundance and population density, but populations moved or changed locations to nearby areas along the monitoring season, rendering transects unsuitable. As a consequence, sampling effort and dates changed across patches, so statistical analyses of tiger beetle densities were no longer adequate. This particular situation is quite different from that observed in other studies (
Most species of cicindelids in the region are easy to identify de visu without the need of seizing the specimens. Only Cicindela campestris and Cicindela maroccana are difficult to differentiate at a distance. In those cases, and in any other particularly difficult situation, specimens were captured with an entomological hand-net. Captured specimens were released on site, except for a few per location that were kept as vouchers or reference, or for future confirmation of identification based on molecular analyses (C. campestris and C. maroccana difficult cases). Specimens captured are preserved dry or in ethanol at the
Each sampled area was visited between 1 and 11 times during the period of study (Table
Sampling was not adequately designed to monitor species with late winter and early spring activity, such as C. campestris, C. maroccana, and possibly L. flexuosa that could be already active around completely water- filled lakes. Therefore, presence and abundances recorded for these three species are not representative of their actual temporal and spatial yearly distribution.
Unpredictable climate changes have a strong influence on the phenology and patterns of activity of tiger beetles. Consequently, in these current years in which rain and temperature patterns are often erratic, the results obtained can only be considered representative of the season studied. Climate changes may have a particularly strong effect on the species that develop most of their activity in dry lake basins, which may suffer strong delays or acceleration during the drying period.
Each specimen was geo-referenced and the general types of soil and vegetation were recorded. Categories used for soil structure were: salt crust (coarse or sheet salt layers on lake basins), salt patches in trails (small salt flats outcropping on patches on trails), granulated salty soils (areas with a thick layer of non-crystalline salty soils, usually gritty or dusty); compact soils in flat trails, compact soils in the banks of channels, saprobe shores (soils formed by decomposed organic matter in lake basins with heavy avian populations), and muds (wet soils with lower salt concentration) (Table
Number of specimens of tiger beetles found on each of the different substrate and vegetation types considered. See text for details.
Soil type / Species | C. littoralis | C. maura | C. dulcinea | C. circumdata | C. campestris | C. paludosa | M. melancholica |
---|---|---|---|---|---|---|---|
Salt crust | 227 | 15 | 5 | 82 | 1 | ||
Granulated salty soils | 116 | 12 | 279 | 19 | 45 | 41 | |
Salt patches in trails | 20 | 153 | 24 | ||||
Trails | 11 | 87 | 7 | ||||
Banks | 27 | 13 | 1 | ||||
Sapropel shores | 52 | ||||||
Mud | 166 | 11 | 13 | 35 | |||
Vegetation type / Species | |||||||
Denuded areas | 274 | 63 | 113 | 83 | 12 | 4 | 10 |
Halophytic prairies | |||||||
open | 132 | 15 | 289 | 18 | 21 | 28 | |
dense | 2 | 11 | 33 | 8 | |||
Non halophytic prairies | |||||||
open | 47 | 5 | |||||
Albardinal | |||||||
open | 122 | 26 | |||||
Shore hidrophytic vegetation | |||||||
open | 106 | 1 | 3 | 1 | 25 | ||
dense | 6 | 4 |
Categories used for vegetation cover were: denuded (areas without vegetation) or vegetated. We split vegetated areas into halophytic and non-halophytic prairies. Within the halophytic category, two main groups were selected: albardinal, open or dense, dominated by Lygeum spartum (considered apart because of its ecological singularities and because it is the main target of the conservation and restoration programs); and other halophytic prairies, open or dense (which include all types of prairies located on salt soils, from the temporarily flooded edges of the saline basins, to the never flooded margin of the lakes. Their vegetation include vegetal communities dominated mainly by Salicornia ramossisima J.Woods, Limonium Mill., Suaeda vera Forssk. ex J.F.Gmel., Salsola vermiculata L., Microcnemum coralloides (Loscos & J. Pardo) Font Quer, Plantago maritima L., Suaeda splendens (Pourr.) Gren. & Godr., Frankenia pulverulenta L., Hordeum marinum Huds., Sphenopus divaricatus (Gouan) Reichenb., Lolium rigidum (Gaudin) Weiss ex Nyman, and Bolboschoenus maritimus (L.) Palla at the shore (
The non-halophytic category, open or dense, includes vegetation not necessarily associated with salty soils, either on relatively dry soils [dominated in the area by Scirpus holoschoenus L., Cirsium monspessulanus (L.) Hill., Plantago major L., Trifolium fragiferum L., Cynodon dactylon (L.) Pers.] (
Presence and number of tiger beetles observed at each locality visited is recorded in Table
Observations on local geographic distribution, general landscape selection (soil structure and vegetation types) and adult activity are described in the following paragraphs, sorted by species.
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.
No. | Lake name / Species | C. littoralis | C. maura | C. circumdata | C. dulcinea | C. campestris | C. maroccana | C. paludosa | L. flexuosa | M. melancholica |
---|---|---|---|---|---|---|---|---|---|---|
1 | Laguna de El Longar | 133 | 8 | 3 | 39 | 9 | 8 | 6 | ||
2 | Laguna del Altillo Chico | 2 | 43 | 12 | 16 | |||||
3 | Laguna del Altillo Grande | 11 | 27 | 16 | ||||||
4 | Laguna de la Albardiosa | |||||||||
5 | Laguna Larga | 49 | 1 | |||||||
6 | Laguna de Tirez | 2 | 21 | 35 | 9 | 4 | 4 | |||
7 | Laguna de Peña Hueca | 15 | 1 | 4 | 30 | 2 | ||||
8 | Laguna del Taray | 11 | 1 | |||||||
9 | Laguna Grande | 22 | ||||||||
10 | Laguna Chica | |||||||||
11 | Laguna Grande | 68 | 8 | 1 | ||||||
12 | Laguna de la Sal | 2 | 6 | 2 | ||||||
13 | Laguna de los Carros | 68 | ||||||||
14 | Laguna de Pajares | 10 | 40 | 3 | ||||||
15 | Laguna de las Yeguas | 22 | 1 | 6 | 66 | 1 | 8 | |||
16 | Laguna Camino de Villafranca | 48 | 13 | 2 | 11 | |||||
17 | Laguna de la Veguilla | |||||||||
18 | Laguna de Salicor | 11 | 1 | 10 | ||||||
19 | Laguna de Retamar | |||||||||
20 | Laguna del Pueblo | 51 | 14 | |||||||
21 | Laguna de Navalafuente | 1 | ||||||||
22 | Laguna de Alcahozo | 4 | 12 | 10 | ||||||
23 | Laguna de Manjavacas | 25 | 20 | |||||||
24 | Laguna de Sánchez Gómez | 4 | 95 | 1 | ||||||
25 | Laguna de la Dehesilla | 52 | 30 | 12 | 2 | |||||
26 | Laguna de Alcahozo Chico | 42 | 28 | 4 | 3 | |||||
27 | Laguna del Taray Chico | 3 | ||||||||
28 | Laguna de El Hito | 29 | ||||||||
29 | Cerro San Cristóbal | 23 | ||||||||
30 | Arroyo de San Marcos | 3 | 4 | 1 | ||||||
31 | La Laguna de Miguel Esteban | 9 | 30 | |||||||
32 | Gigüela marshes | 7 |
Calomera l. littoralis (Fig.
Maps showing the location of species of Cicindelinae found in the wetlands of La Mancha. Numbers correspond to Fig.
Representative habitats of Cicindelinae in La Mancha. A Laguna de la Sal (Toledo) in May. Salt flats occupied by Calomera l. littoralis B Laguna de Peña Hueca (Toledo) in April. Wet salt flats, with gypsum outcrops, occupied by C. l. littoralis; when totally dry, in summer, occupied by Cephalota circumdata imperialis C Laguna de Las Yeguas (Ciudad Real) in June. Salt flats, occupied by C. l. littoralis and C. c. imperialis; and halophytic prairies occupied by Cephalota dulcinea and Cylindera paludosa D La Laguna de Miguel Esteban (Toledo) in July. Salt flats, occupied by C. l. littoralis; and halophytic prairies with “albardinal” occupied by Cephalota m. maura E Laguna del Camino de Villafranca (Ciudad Real), with flamingos, in July. Sapropel shores occupied by C. l. littoralis F Laguna de Alcahozo (Ciudad Real) in June. Salt flats, occupied by C. l. littoralis and C. c. imperialis; and open areas near dry halophytic prairies occupied by C. dulcinea and also by C. c. imperialis. Photographs by N. Percino and MG-P.
Representative habitats of Cicindelinae in La Mancha. A Channel at Laguna de las Yeguas (Ciudad Real) in May. Wet areas occupied by Calomera l. littoralis and Cicindela campestris; when drier, occupied by Cephalota m. maura B Road side pools and halophytic vegetation in Laguna de Manjavacas (Cuenca) in April. C. l. littoralis was the only species observed C Wells and deep pools in Laguna de Manjavacas (Cuenca) in June. Cephalota m. maura is frequent in these structures D Laguna del Pueblo de Pedro Muñoz (Ciudad Real) in July. Mud areas amongst hydrophytic matts, occupied by C. l. littoralis and Myriochila m. melancholica E Temporary pools at Laguna del Longar (Toledo), in July. Calomera l. littoralis, Myriochila m. 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 C. c. campestris. Photographs by N. Percino and MG-P.
Most populations of C. littoralis changed location within the same locality during the period of study. We followed these intra-populational movements at some lakes. At Laguna de Manjavacas, specimens were active in late April in the salty marshes surrounding the main, totally water- filled, salt lake. As the lake started to dry out, specimens appeared along the shoreline, and disappeared completely from the dried out salty marshes. Subsequently, and until the complete drying out of the main lake, specimens of C. littoralis, moved from nearby peripheral vegetated areas towards central flat drying areas (Fig.
Observations of Calomera l. 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.
Most specimens were located on salt crust (40.5%), in open denuded areas (48.8%) (Table
Specimens of C. littoralis were present in the area all over the study period (Fig.
Abundance of C. littoralis is highly variable across the study area (Table
Calomera littoralis was occasionally found in sympatry with C. campestris in late spring (La Dehesilla, Camino de Villafranca Channel, Alcahozo Chica), with C. maroccana (La Sal), and with C. circumdata (Altillo Chica, Tirez, Peña Hueca), but strict syntopy, sharing a unique patch, was only observed with M. melancholica at the end of summer (El Longar, Tirez, Camino Villafranca, and Pedro Muñoz), and occasionally with C. maura (El Longar, Camino de Villafranca Channel) and with L. flexuosa (Alcahozo Chica) in summer. When present, C. littoralis is generally the dominant species, except in extreme drying-out situations at the lake basins or channels, when C. circumdata or C. maura might be temporally dominant.
Calomera littoralis is a conspicuous and very active species, displaying its activity in plain sunlight. When approached, they run fast, and when disturbed they are able to start a quite long and sustained, relatively elevated, flight, sometimes changing direction in mid- flight. Flight is never directed to vegetated areas. At some localities (Manjavacas, Peñahueca, Alcahozo Chico) many specimens remain quiet among the sparse vegetation of the basin shore, running towards the central open areas when disturbed. One specimen was observed at night, hidden, but alert, in the crevices of a thick salt crust layer (Peña Hueca).
Cephalota m. maura (Fig.
Most specimens were located on salt patches in trails (20.8%), and on banks and other compact soils (39.6%). According to
Adult specimens of C. maura were observed through most of the study period (Fig.
Cephalota m. maura was occasionally found in strict syntopy with C. campestris in late spring (El Longar, Salicor and San Cristóbal channels), with C. dulcinea and C. paludosa (road to Pajares) in early summer, with C. littoralis in mid and late summer (Laguna Grande de Villafranca, El Longar, Camino Villafranca channel), and with M. melancholica at the end of summer (El Longar).
Specimens of C. m. maura are very active, displaying most of its activity in plain sunlight, although some specimens were observed resting on the ground between the twigs of Scirpus at mid-day, near the water (La Laguna de Miguel Esteban, Laguna Grande de Villafranca). When approached, they run fast, displaying a short low flight when threatened, often directed towards vegetated areas, or even into the water surface, from where they reach the shore.
Cephalota circumdata imperialis (Fig.
Specimens of C. circumdata were present in the area during summer, from late June to August (Fig.
Population densities of C. circumdata are variable, but normally low. Single specimens were often found alone, far from any other specimen, and when close, they were usually relatively isolated, and well distributed along favourable habitat. C. circumdata was occasionally found in sympatry with C. littoralis (Altillo Chica, Tirez, Peña Hueca, Salicor), and with C. dulcinea at the start of summer (El Longar, Las Yeguas, Alcahozo).
Most specimens were observed displaying their activity in plain sunlight, in agreement with previous observations of circadian activity of C. circumdata leonschaeferi in Italy (
Behavioural observations on Cephalota circumdata imperialis. A, B A few specimens of C. 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.
Cephalota dulcinea (Fig.
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 C. m. maura C Trails and “albardinal” in Laguna del Altillo Chica (Toledo), where it co–occurs with C. paludosa D Open areas amongst halophytic vegetation at laguna de La Sal (Toledo), Calomera l. 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, C. l. littoralis and C. 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.
Active adults were observed from late spring to midsummer (Fig.
Changes in population extension and local positioning were marked in some lakes, particularly at Los Carros, Sánchez Gómez and La Dehesilla, while in other areas the specimens occupy constantly at least a portion of the locality (Tirez, Peña Hueca, Las Yeguas). At Los Carros, La Dehesilla and Las Yeguas, large concentrations of specimens were present along dirt roads and trails in late spring (Fig.
Cephalota dulcinea was occasionally found in sympatry with C. circumdata, but strict syntopy, sharing a unique patch, was only observed with C. paludosa at the end of spring in open flats within vegetated areas, and occasionally with C. maura along trails. When present, C. dulcinea is generally the dominant species, and it was the only species found in some small lakes (Los Carros).
Cephalota dulcinea is an active species, displaying its activity in plain sunlight. When approached, they run fast and fly relatively short distances, sometimes changing direction in mid- flight. Flight can be directed to vegetated areas, but usually they fly farther along trails or saline flats. Occasionally, we found some specimens resting in the shade of objects, at midday (El Longar). Preying behaviour was observed towards ants, other small hymenopterans, and small dipterans. Some specimens seem to keep a hunting spot for a period, as suggested by the insect remains scattered around them (Fig.
Behavioural observations on Cephalota dulcinea A Specimen of C. 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 C. dulcinea involved in courship C A recently killed specimen of C. dulcinea in a trail, about to be seized by ants D Trail located near laguna de Pajares with a large colony of C. dulcinea, C. m. maura and scattered C. 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.
Cicindela c. campestris (Fig.
Specimens of C. campestris were present in the area in late spring (Fig.
Population abundances of C. campestris are highly variable (Table
Cicindela campestris is an active but not conspicuous species in the area, displaying its activity in small patches and trails amongst vegetation. When approached, they run along the trails or into vegetation, and when flying, they show a tendency to land on vegetation. Copulatory behaviour was observed during most of the study period.
Cicindela maroccana (Fig.
Specimens of C. maroccana were present in the area during late spring. According to previous studies, the species is also present in early spring (
Cylindera paludosa (Fig.
Specimens of C. paludosa were present from late spring to midsummer (Fig.
Cylindera paludosa was occasionally found in sympatry with C. dulcinea, around salt flats and vegetated saline marshes (Altillo Chica, El Longar, Las Yeguas, La Dehesilla), with C. campestris in late spring (Salicor), and with C. maura (El Longar channel, Pajares trail). Cylindera paludosa is generally the dominant species in salt marshes, and often it is the only species present in marshes of limited extension. Cylindera paludosa is an inconspicuous but very active species, displaying its activity in plain sunlight. When approached, they fly readily towards the nearby vegetated areas, rendering its observation difficult.
Lophyra f. flexuosa (Fig.
We found the specimens in late spring (mid May 2014, and early June 2012). Despite successive samplings, the observations were made in a single occasion. These observations are puzzling, because, according to the literature and personal observations, the species has a relatively long period of activity, from April to July, in other areas of Castilla – La Mancha and nearby Madrid (
35 specimens of Myriochila m. melancholica (Fig.
The period of activity of M. melancholica was restricted to the end of the study period, from late June to early August (Fig.
Frequency of observation of M. melancholica in the area is low, with only a few specimens found per visit. Myriochila melancholica was always and only found in syntopy with Calomera littoralis. In these cases C. littoralis was the dominant species. Myriochila melancholica displayed its activity in plain sunlight, but it is also attracted to artificial lights (
The tiger beetle taxocenosis in La Mancha wetlands is composed of 9 species (Table
The cicindelid assemblage at La Mancha wetlands could be even more complex, since two additional species of tiger beetles, not found in the study area, inhabit the region of Castilla – La Mancha. Cicindela lagunensis has been located in sandy soil areas, relatively close to La Mancha wetlands (
Such a concentration of cicindelid beetles, 9 species in an area of less than 1º latitude/longitude square, is likely the result of a combination of both historical (paleogeographic) and ecological factors. The presence of one endemic species, C. dulcinea, suggests that, on the one hand, the region has maintained a relative level of historical isolation with respect to other Iberian areas. On the other hand, some areas acting as sanctuaries of diversity (sensu
The diversity of environments available for tiger beetles at La Mancha wetlands, also contributes to the maintenance of such a high number of cicindelids, since most of the species seem to occupy particular ecological niches within the area (Fig.
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 C. l. littoralis, C. dulcinea, C. circumdata imperialis and C. m. maura B Trails on salty soils at Laguna de la Dehesilla (Cuenca), where C. l. littoralis, C. c. campestris and C. dulcinea are present along different periods of the year C Old channel in Laguna del Longar (Toledo), where C. m. maura, C. campestris and C. paludosa co-occur D Saline pool at Laguna de Tirez (Toledo), where C. l. littoralis and M. m. melancholica co-occur, while populations of C. dulcinea and C. 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 C. l. littoralis was present F Vineyards at the edge of Laguna de Alcahozo Chico (Cuenca), note the diverse halophytic vegetation and nearby open areas where C. l. littoralis, C. c. campestris, C. paludosa, and L. f. flexuosa live in close proximity. Photographs by N. Percino and MG-P.
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 M. m. melancholica was found, correspond to the habitat shown in Fig.
Habitat partitioning among species in La Mancha is evident at temporal and spatial scales. On the temporal scale, peaks of activity differ markedly among species (Figs
Cephalota dulcinea, C. circumdata, M. melancholica, and possibly C. maroccana, concentrate adult activity in a short period of time, in a single adult emergence period (Figs
Overlap in habitat use is maximal for M. melancholica and C. littoralis when M. melancholica is present, as previously reported by
All other species show little overlap in habitat use (see Fig.
Populations of some species remained on the same spot throughout their whole period of activity, from the earliest observations to the latest. Amongst those, C. circumdata, C. paludosa and C. maura are the most representative, but also some populations of C. dulcinea remained at the same location during their whole active period. Populations of C. littoralis and some of C. dulcinea, moved to different patches within the same location throughout their period of activity. Populations of C. littoralis shifted their location following the retreat of water during the continuous drying-out of lakes and pools (Fig.
Apparent location shifts of C. dulcinea were observed in dry lakes surrounded by trails and dirt roads, particularly at Las Yeguas and Los Carros, but also in other lakes on a smaller scale. Soon after the start of their activity, we observed large concentrations of specimens of C. dulcinea along trails and roads around salt lakes, and less numerous at the salt basin shore in the typical halophytic prairies used by the species throughout the area. The number of specimens located on roads and trails declined as the season advanced, and completely disappeared well before the end of the general activity period of the species (Fig.
These observations, coupled with the absence of C. dulcinea at Laguna de Salicor, where it was found in 2012, suggest that, as a working hypothesis, the whole C. dulcinea deme might function as a metapopulational system, with high probabilities of local extinction matched by frequent recolonization events (
Populations of tiger beetles in La Mancha seems to present a relatively healthy status. Population densities are locally high for C. l. littoralis, C. maura, C. dulcinea, C. paludosa, and C. c. campestris, a set of species with a relatively extensive area of occupation at La Mancha wetlands. Cephalota circumdata imperialis and M. m. melancholica, present less dense populations, and also a more restricted local distribution, but they are nonetheless well distributed across the area. Lophyra f. flexuosa and C. maroccana, two widespread species in Central Spain (
Most of the studied lakes and marshes are legally protected (http://ec.europa.eu/environment/nature/natura2000/db_gis/). Conservation efforts over the last decade have been directed at restoring the original ecological system, and also at eliminating the remains of human activity, particularly the removal and restoration of drainage channels, removal of urban waste, closure and restoration of village dumpsters, control of artificial flooding, and, more recently, restoration of native vegetation (the albardinal and the Limonium prairies), restoration of adjacent agricultural lands around the lakes, removal of salt- related semi-industrial activities, and road tracks removal and restoration (Fig.
We have detected a series of threats that may affect the whole cicindelid (and other salt- associated animals) community. These threats are mostly derived from current agricultural practices around marshes, non-adequate use of trails and dirt roads, and insufficient regulation of water quality. Other threats might be derived from the restoration process itself, particularly the removal of abandoned semi-industrial infrastructures and trails.
Ploughing and cultivation of cereal, legumes, garlic or grapes, in fields adjacent to the lakes (Fig.
Restoration practices might have a significant impact on the larger saline lakes that were subjected in the past to semi-industrial activities, predominantly salt extraction. Salt extraction industries modified the salt flats by creating evaporation pools, scattered deep pits, and some channels (Figs
Closure and restoration of trails and roads around the lakes might pose a threat for species such as C. dulcinea, that consistently use them for hunting and mating. Further studies are required to evaluate the role of these man-made structures for the species, and particularly whether they have any impact on larval development.
As general recommendations, we propose that the current practices of governmental buying of agricultural land around the lakes are absolutely necessary, both to avoid potentially dangerous agricultural practices and to create a buffer zone from other activities such as sheep grazing. Trails and minor roads must be closed for high risk sport practices (quads and 4x4 vehicles), while they can be used for other less aggressive activities. We recommend the maintenance of trails around lakes until their effect on populations of C. dulcinea is properly evaluated. Old semi-industrial structures, evaporation pools, pits and channels, may be retained to increase structural diversity of the lakes. Finally, and especially considering the possible metapopulation system of C. dulcinea, we consider necessary the implementation of legal protection for all salt marshes and saline lakes, of the occupation area of C. dulcinea, including all minor entities that could act as migrational steps during recolonization events.
Our data support the status of protected species for C. circumdata, a definite indicator of well-preserved salt lake flats, with relatively low population densities, and C. dulcinea, an endemism to the salt marshes and lakes of La Mancha. A species appearing as a good indicator of protected halophytic prairies (Limonium, Lygeum) is C. paludosa, but we do not consider necessary additional specific protection for this species.
We are indebted to Nohemí Percino, Melinda Hoffman, Rebecca Shi, Michaël Moes, Pilar López García-Gallo, Diushi Keri, Zahra Saberi, Hamid R. Ghanavi, Gonzalo García Martín and Jorge Espiñeira, for their help during some of the long sampling days, catching and counting tiger beetles under the relentless summer sun. We thank Pablo Pichaco for the additional information supplied. Also, thanks to Pablo Pichaco, Mercedes París and Manuel Sánchez Ruiz for their hospitality and generosity. PCRF and MGP thank Pablo Pichaco and Marcos Toribio for their companionship and sense of humour during nocturnal sampling, Carlos Pérez and Amanda del Río (Global Nature) for their support. We especially thank Julia G. Paris and Eva Nozal for the English revision of the manuscript. We thank Radomir Jaskuła for his very useful comments on the paper. The Consejería de Medio Ambiente de Castilla-La Mancha provided us with all the needed help, including permits for the capture and observation of tiger beetles. This study was possible thanks to the support from the Fundación Global Nature through the European Life European Community Project Humedales de La Mancha (LIFE10 NAT/ES/000563), Spanish Ministry of Agriculture, Food and Environment and to the facilities provided by the Museo Nacional de Ciencias Naturales (
Appendix
Data type: Occurrence
Explanation note: Field observations.