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Heat-Cold Dialectic in the Activity of Proformica Longiseta, a Thermophilous Ant Inhabiting a High Mountain (Sierra Nevada, Spain)

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Heat-Cold Dialectic in the Activity of Proformica Longiseta, a Thermophilous Ant Inhabiting a High Mountain (Sierra Nevada, Spain) Int J Biometeorol (1998) 41:175–182 © ISB 1998 ORIGINAL ARTICLE &roles:Ignacio Fernández-Escudero · Alberto Tinaut Heat-cold dialectic in the activity of Proformica longiseta, a thermophilous ant inhabiting a high mountain (Sierra Nevada, Spain) &misc:Received: 1 August 1997 / Accepted: 27 October 1997 &p.1:Abstract The activity of the thermophilous ant Profor- Cerdá and Retana 1988; Fellers 1989; North 1993) and, mica longiseta has been studied in a Mediterranean high- in other cases, temperature (Harkness 1977; Hölldobler mountain environment. An analysis has been made of the and Taylor 1983; Curtis 1984a; Cloudsley-Thompson biotic and abiotic variables involved, the location of and 1989) or light intensity (Gano and Rogers 1983; Roseng- conditions surrounding the activity, as well as the strate- ren and Fortelius 1986; Cerdá and Retana 1989). Besides gies used by the ant to remain active at high tempera- these factors, which might be the most obvious ones, bi- tures. The results of this study indicate that the maxi- otic factors may emerge, superimposed over them, which mum activity occurs during the middle hours of the day may control the rhythm of the activity. These latter fac- and that the variable which most influences daily activity tors include: fluctuation in food availability (Whittford is temperature, especially at the soil surface. With re- and Etterschank 1975; Cros et al. 1992); biological cycle spect to the biotic variables, the availability of food and and larval presence (Bosch et al. 1987; Cros et al. 1992); the demand for food by the larvae strongly correlate with competition or conflict with simultaneously foraging activity. The time outside the nest is usually spent in veg- species (Bernstein 1979; Retana et al. 1988); interfer- etation searching for food. This activity continues even ence of myrmecophagous (anteaters) species (Mehlop when the temperature of the soil surface exceeds 58°C. and Scott 1983); and compromises between predator To tolerate these temperatures, the ant not only has a pressure and heat stress (Wehner et al. 1992). high resistance to heat (critical thermal maxi- Most of these studies have focused on arid or desert mum = 51.1°C), but also increases the speed of its move- areas but not at high altitude, as in the present paper. ments and resorts to thermal refuges. The present work Mountain biotopes differ markedly from low arid or de- contributes data on the biology of this highly thermophil- sert areas, primarily because sharp decreases in atmo- ous species, which is also capable of tolerating a long, spheric pressure set in motion a series of peculiarities. hard winter and then developing within a short vegeta- The principal characteristic, the reduction in dust and tive period. haze particles, results in lower air humidity and therefore increased insulation and radiation. All of these factors &kwd:Key words High mountain · Activity · Temperature · exert considerable impact on the ecology, morphology Resistance to heat · Ants&bdy: and physiology of alpine fauna, particularly arthropods (Mani 1990; Sømme and Block 1991). The Mediterranean mountain is unique among high- Introduction altitude biotopes because its latitudinal position increas- es total radiation and decreases air humidity as well as The activity pattern of ant species arises from the inter- rainfall. This creates a cold, arid environment with spe- action of various factors (Heatwole and Muir 1989). In cial characteristics (Table 1) such as: (1) a range of daily some cases, “ambient conditions” are held to be the gen- temperature variations exceeding that of more northerly eral factors responsible for this pattern (Ayre 1958; and wetter mountains (the Alps); (2) a rainfall rate dur- ing the vegetative period (May–September) similar to I. Fernández-Escudero (✉)1 · A. Tinaut that of arid environments. Furthermore, the Sierra Neva- Departamento de Biología Animal y Ecología, Facultad de Ciencias, Universidad de Granada, E-18071 Granada, da, although arid, differs from deserts, at least hot de- Spain serts, in having: (1) a thermal contrast between the soil Present address: surface and the air which is more extreme than in de- 1 Department of Genetics, Uppsala Genetic Center, serts; (2) low night-time temperatures, even in summer. University of Uppsala, Box 7003, S-75007 Uppsala, Sweden&/fn-block: All these characteristics make the Sierra Nevada more 176 Table 1 Climatic data for de- sert areas and high mountain&/tbl.c:&tbl.b: Place Average Average Annual average Annual temperature (°C) temperature (°C) temperature precipitation January July (°C) (mm) Turanicoa –6.9 to 4.8 27.1–32.3 11.0–17.1 82–244 Kazahjastana –13.6 to 6.0 25.1–28.6 7.5–12.1 103–184 Dzungariaa –22.0 to –15.2 23.9–25.4 4.3–6.8 96–247 Gobia –18.1 23.3 3.8 107 Pei-Shana –15.6 to –9.8 22.6–24.6 4.3–9.6 42–74 Deserts of Tibet and Pamira –20.0 8.0–12.0 – 103 Great Basina –2.0 20.4 7.8–13.2 158–419 Chihuahuab 12.2 26.6 21.1 204.3 Saharac 11.8 35.1 22.8 32.1 Sierra Nevada (2500 m)d –2.5 14.0 3.3 In summer 142 Alps (2500 m)e –9.5 5.2 –2.7 1552 Pyrenees (1714 m)f –1.6 13.4 5.2 In summer 635 Place Average diurnal Difference air temperature- Lowest temperature aWest 1983, Orlovsky 1994 range ground temperature in July b Morafka et al. 1981 (°C) (°C) (°C) c Delye 1968 d Tinaut 1979, Prieto 1983 Sierra Nevada (2500 m)d 7.7 40 5 e Mani 1968, 1990 Alps (2500 m)e 5– 6 f Font Tullot 1983 Saharag – 18 15.5 g Wehner 1989 Tunisiah –2115 h Heatwole and Muir 1979&/tbl.b: similar to a cold desert than to other European high The objective of this investigation was to study the mountains or other arid zones. activity pattern of a Mediterranean high-mountain ant Under these conditions, it might be expected that the species and its relation to abiotic and biotic factors, in an rhythms of biological activity, as well as habitat use, attempt to resolve the apparent contradiction of a ther- would contrast clearly with those in arid or hot zones; mophilic ant inhabiting a cold environment. that is, while one of the prime aims of animals in arid zones is heat loss, in the high mountain it is heat gain. Thus, in arid and hot areas, some ant species change Materials and methods their daily activity patterns over the year, becoming noc- turnal during the summer, e.g. Tapinoma nigerrimum in Natural history of Proformica longiseta the Mediterranean area (Cerdá and Retana 1988) or P. longiseta Collingwood, a small [3.3 (SEM 5) mm; n = 30) allo- Messor arenarius in the pre-Sahara steppe (Heatwole and metric and polymorphic ant (Fernández-Escudero et al. 1994), is Muir 1989). In addition, two groups of species, diurnal endemic to the Sierra Nevada and neighbouring mountains be- tween 1900 m and 2800 m in altitude. The nests are found under (heat-adapted) and nocturnal (cold-adapted), inhabit most stones which serve as an efficient heat source (Fernández-Escu- deserts, as in Anatolia (Baroni-Urbani and Aktac 1981) dero et al. 1993; Tinaut and Fernández-Escudero 1993). The P. or the Sahara (Heatwole and Muir 1989; Heatwole 1991). longiseta populations typically lodge between 155 and 1200 indi- On the other hand, Somme and Block (1991) stated viduals. This ant feeds mainly on nectar and occasionally on dead insects (Fernández-Escudero and Tinaut, in press a). It inhabits the that in alpine habitats arthropod activity can be noctur- driest areas of these mountains, where it is the most abundant ant nal, a situation that these authors assumed to represent (83.16%; Tinaut 1979). The other ant species sharing the area in- an adaptive advantage, perhaps by reducing the risk of clude Tetramorium caespitum L., Tapinoma nigerrimum Nyl., overheating in melanistic species, or by maintaining a Leptothorax tristis Bondroit, L. tuberum Fabricius and Formica lemani Bondroit. corporal water balance or avoiding predators. In a high The genus Proformica inhabits arid steppes from Central Asia Mediterranean mountain the temperature difference be- to the Iberian Peninsula and thus occupies cold as well as hot plac- tween the day and night or between the presence or ab- es. In the Iberian Peninsula, two other species, P. f e r re r i Bondroit sence of sun is higher than in an alpine mountain; there- and P. nasuta Nylander, live in a savannah-like grassland but only fore, the compromise is between the heat gain during the at altitudes of no more than 1000 m (Tinaut 1981; Cros 1986). night or sun absence and heat loss during the middle of the day. Study area However, despite the importance of the ants in these The present study was carried out at the head of the San Juan River mountainous habitats (Pisarski 1978; Gómez and Zamo- (2400 m in altitude) in the Sierra Nevada (Granada, Spain). The ra 1992), few studies have focused on how these insects study area covers the dry places in this valley, where the most rep- manage to thrive under such harsh climatic conditions resentative vegetation includes Genista versicolor Boiss., Juniperus (Heinze and Hölldobler 1994) and virtually nothing is communis subp. alpina (Suker) Celak, Hormatophylla spinosa (L.) Kupler and Thymus serpilloides Bory. Of the vegetal cover, roughly known about the activity pattern of high-mountain ants. 16% is shrub, 27% grass, while 57% is bare ground. The soil is In addition, the high mountain probably does not have very poor and siliceous. This zone belongs climatically to the At- the broad spectrum of ant species found in the hot desert.
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