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Forest succession endangers the special ant fauna of abandoned quarries along the River Meuse Wallonia, Belgium.

The user has requested enhancement of the downloaded file. All in-text references underlined in blue are added to the original document and are linked to publications on ResearchGate, letting you access and read them immediately.

Forest Succession Endangers the Special Ant Fauna of Abandoned Quarries along the River Meuse Wallonia, Belgium Wouter Dekoninck,1,2,3 Frederik Hendrickx,1,2 Michel Dethier,4 and Jean-Pierre Maelfait2,5 Abstract the nature value of the ant fauna of these sites, we propose In Western Europe, old abandoned mining sites and quar- a so-called habitat preference approach, wherein each ries are often of high biodiversity and conservation value species is assigned to one of the following three habitat due to the presence of a number of endangered species.

In preference categories: 1 eurytopic, 2 bound to wet the southern part of Belgium Wallonia , many ancient shadow-rich habitats, or 3 bound to dry open habitats.

In , gered in Belgium and of high conservation value. The pro- we collected 26 ant species by pitfall trapping in four of portion of the total number of captured specimens these quarries.

In addition to common ones, several rare included in the latter habitat preference category group is species, usually associated with mesomorphic to xeromor- strongly reduced as scrub and tree encroachment advan- phic grasslands, were found in high numbers.

Quarries ces. This proportion can therefore be used as a proxy to undergoing forest succession were dominated by eurytopic monitor the effects of management measures that prevent species and by species typical of wet shadowy places, further forest succession.

Therefore, we suggest a management that halts further Key words: conservation value, diversity, forest succes- forest succession of open mesomorphic and xeromorphic sion, Formicidae, quarry restoration, refuges, xeromorphic habitat patches in these quarries.

To assess and monitor grasslands. Introduction ; Belgium: Bizoux et al. As the natural have caused many drastic local changes to soil, vegetation, habitats of many of these endangered species have been and landscape, both in Belgium and elsewhere in Western considerably reduced due to a range of human activities, Europe.

These two industries, together with rock extrac- these old mining sites are the only remaining refuges for tion, were responsible for many quarries and mining sites.

This led to the often abundant mainly consists of species that prefer xerothermic habitats. An adequate conservation and restoration management of these quarries that not only considers species number but 1 also includes the habitat affinities of the most endangered Department of Entomology, Royal Belgian Institute of Natural Sciences, Vautierstraat 29, B Brussels, Belgium species needs further implementation.

As restoration 2 Terrestrial Ecology Unit, Ghent University, K. Ledeganckstraat 35, B management of these types of sites is now one of the Ghent, Belgium 3 Address correspondence to W.

Dekoninck, email wouter. In Australia Majer ; Steiner et al. This is particularly so when ; Hoffmann et al.

This allows reliable comparisons of Koch ; Redi et al. All specimens have been identified using Seifert Different stages of forest succession were represented in Voucher specimens are deposited in the personal collec- the sampling design.

The aims of this paper were as follows: 1 To explore which factors are the main drivers of ant Habitat Preferences assemblage structure and diversity in these quarries.

Based on its habitat preference in Flanders Dekoninck 2 To evaluate the effect of the present state of forest et al.

Besides four generalist species—here called eurytopic spe- cies EU , which have no preference for any particular hab- Methods itat but occur in all kinds of disturbed habitat patches—all other species are habitat specialists.

Within this last Description of the Sites group, we can distinguish species from wet closely vege- In Belgium, the only remaining mining activities are the tated, shadow-rich habitats WCH and species from extraction of clay, sand, and gravel in the northern part of open dry habitats ODH.

One species Tetraponera alla- Belgium Flanders and the quarrying of limestone, dolo- borans could not be placed in one of these categories as it mite, slate, porphyries, and marble in the southern part of is an accidentally introduced species from Southeast Asia Belgium Wallonia Diels In Wallonia, most of the Ward ; Dekoninck et al.

We sampled 12 sites distributed over four ancient quarries. Their general and vegetation characteristics, The variation in species assemblage composition among history, surface, and duration of uninterrupted forest suc- the sites was analyzed using an indirect gradient analysis, cession are summarized in Table 1.

Except for site T4 where detrended correspondence analysis DCA. DCA is an grassland was being grazed by sheep until , the sampled indirect multivariate technique that positions samples quarries were left unmanaged after excavation stopped.

Default settings were used, that is, detrending by 26 Sampling and Identification segments and nonlinear rescaling.

Pitfalls with a diameter of 9. Only the 23 most abun- traps were emptied fortnightly from April until October dantly caught species were included in the analysis to Although the use of pitfall trapping to collect ants in avoid bias caused by accidental immigrants and vagrants Western and Central Europe has been criticized by some e.

Description of the sites with characterization of vegetation and afforestation stage. Lat A. We therefore opted to calculate the average number composition than species richness alone.

Moreover, multiple sites were sampled in Spearman rank order correlations of the surface area of most quarries, which induces spatial dependence in the data the sampled sites and S, H, and E were carried out in that does not allow us to treat them as independent data order to take species—area relationships into account.

All species, their scaled and ranked abundance per site, and their habitat preference category are given in Table 2. The maximum number of species 18 was found on the xeromorphic site E1 in Engis.

In all xeromorphic and other grassland vegetations except T4 , more than nine species were found. Low evenness in the open thermophilic sites resulted from the dominance of one or two species in these sites.

Lasius niger was one of the dominant species in C2, C3, and E1, and Myrmica sabuleti was dominant in C3, T2, and T3.

In the sites with spontaneous establishment of forest, diver- Figure 2. Species are pre- decreased. The scores of the sites are species richness increased.

However, evenness showed no indicated as filled dots open vegetation or filled triangles forested similar tendency in the forest sites. In these sites, evenness sites.

Abundances of species typical for xeromeso- the sites with S, H, and E were not significant. Sites do not cluster per quarry, and the forest wet grasslands, were also significantly correlated with axis sites Group A were separated from the open thermo- 1.

The dense grassland site T4 where sheep grazing stop- ped 10 years ago, clustered together with E1 and C2, near the group of sites undergoing forest succession.

The ant community of site T4 was dominated by Formica fusca and the eurytopic Lasius niger and Myrmica rubra. Habitat Preference of the Different Assemblages and the Habitat Preference Approach In the ancient quarries with spontaneous establishment of Figure 1.

In some cases, restoration has been very successful and shows that these man-made and sometimes polluted sites are able to recover.

The above-mentioned findings are confirmed by the present study, showing that abandoned and recovering quarries in Western Europe can harbor a diverse and in- teresting ant fauna that is very valuable in terms of nature conservation.

This is most pronounced for the sites with dry open vegetation where species from heathland and mesomorphic to xeromorphic grasslands can constitute Figure 3.

The number of species S and the habitat preference of the a considerable part of the ant fauna. These species with a species collected in each site with WCH, DOH, and EU species.

In group B however species from ODHs Open or Closed Vegetation in the Ancient Quarries? Our results show that forest succession during natural The relative worker abundances for the three habitat recovery of ancient quarries has a deleterious effect on preference categories are shown in Figure 4.

This habitat the ant fauna both its species richness and the occurrence preference approach shows that the relative abundance of and abundance of xeromorphic and hence threatened DOH specimens is much lower in the sites undergoing for- species.

Other studies also reported on the high species est succession. The relative abundance of DOH specimens richness of ant assemblages of open habitats such as in site T4 was intermediate.

This was in the relative importance of ants of the DOH category to explained by the high exposure of these habitats to direct species of the WCH and EU category.

Open ancient quarries along the river Meuse in Belgium started ancient quarries with higher solar radiation allow less Figure 4. Moorish — obs.

James Morison , after his separation from the United Secession Church. Morn , morn, n. Morse , mors, n. Morse Mort , mort, n.

Mosa , the river Meuse, Gr. Doublet Mussulman. On the 11th Simbirsk fell, and on 8 October Samara. The Whites fell back eastwards to Ufa and Orenburg.

In Omsk the Russian Provisional Government quickly came under the influence — then the dominance — of its new War Minister, Rear-Admiral Kolchak.

The members of the Directory were arrested and Kolchak proclaimed the "Supreme Ruler of Russia". By mid-December White armies had to leave Ufa, but they balanced this failure with a successful drive towards Perm , which they took on 24 December.

In February the Red Army overthrew the White Russian-supported Kokand autonomy of Turkestan. British support of the White Army provided the greatest threat to the Red Army in Central Asia during Great Britain sent three prominent military leaders to the area.

One was Lt. Bailey , who recorded a mission to Tashkent, from where the Bolsheviks forced him to flee. Another was Gen. Malleson , leading the Malleson Mission , who assisted the Mensheviks in Ashkhabad now the capital of Turkmenistan with a small Anglo-Indian force.

However, he failed to gain control of Tashkent, Bukhara and Khiva. The third was Maj. Dunsterville, who the Bolsheviks drove out of Central Asia only a month after his arrival in August The first regional congress of the Russian Communist Party convened in the city of Tashkent in June in order to build support for a local Bolshevik Party.

In July two Left SR and Cheka employees, Blyumkin and Andreyev, assassinated the German ambassador, Count Mirbach. In Moscow a Left SR uprising was put down by the Bolsheviks, using Cheka military detachments.

Lenin personally apologized to the Germans for the assassination. Mass arrests of Socialist-Revolutionaries followed.

Estonia cleared its territory of the Red Army by January This rendered possible another threat to the Red Army—one from Gen. Yudenich, who had spent the summer organizing the Northwestern Army in Estonia with local and British support.

In October he tried to capture Petrograd in a sudden assault with a force of around 20, men. The attack was well-executed, using night attacks and lightning cavalry maneuvers to turn the flanks of the defending Red Army.

Yudenich also had six British tanks, which caused panic whenever they appeared. The Allies gave large quantities of aid to Yudenich, who, however, complained that he was receiving insufficient support.

By 19 October Yudenich's troops had reached the outskirts of the city. Some members of the Bolshevik central committee in Moscow were willing to give up Petrograd, but Trotsky refused to accept the loss of the city and personally organized its defenses.

Trotsky himself declared, "It is impossible for a little army of 15, ex-officers to master a working-class capital of , inhabitants. Trotsky armed all available workers, men and women, ordering the transfer of military forces from Moscow.

Within a few weeks the Red Army defending Petrograd had tripled in size and outnumbered Yudenich three to one. At this point Yudenich, short of supplies, decided to call off the siege of the city and withdrew, repeatedly asking permission to withdraw his army across the border to Estonia.

However, units retreating across the border were disarmed and interned by order of the Estonian government, which had entered into peace negotiations with the Soviet Government on 16 September and had been informed by the Soviet authorities of their 6 November decision that, should the White Army be allowed to retreat into Estonia, it would be pursued across the border by the Reds.

Following the Treaty of Tartu most of Yudenich's soldiers went into exile. Former Imperial Russian and then Finnish Gen. Mannerheim planned an intervention to help the Whites in Russia capture Petrograd.

However, he did not gain the necessary support for the endeavour. Lenin considered it "completely certain, that the slightest aid from Finland would have determined the fate of [the city]".

The British occupied Murmansk and, alongside the Americans , seized Arkhangelsk. With the retreat of Kolchak in Siberia, they pulled their troops out of the cities before the winter trapped them in the port.

The remaining White forces under Yevgeny Miller evacuated the region in February At the beginning of March the general offensive of the Whites on the eastern front began.

Ufa was retaken on 13 March; by mid-April, the White Army stopped at the Glazov — Chistopol — Bugulma — Buguruslan —Sharlyk line.

Reds started their counteroffensive against Kolchak's forces at the end of April. The Red 5th Army, led by the capable commander Tukhachevsky , captured Elabuga on 26 May, Sarapul on 2 June and Izevsk on the 7th and continued to push forward.

Both sides had victories and losses, but by the middle of summer the Red Army was larger than the White Army and had managed to recapture territory previously lost.

Following the abortive offensive at Chelyabinsk, the White armies withdrew beyond the Tobol. In September a White offensive was launched against the Tobol front, the last attempt to change the course of events.

However, on 14 October the Reds counterattacked, and thus began the uninterrupted retreat of the Whites to the east. On 14 November the Red Army captured Omsk.

Kolchak lost control of his government shortly after this defeat; White Army forces in Siberia essentially ceased to exist by December.

Retreat of the eastern front by White armies lasted three months, until mid-February , when the survivors, after crossing Lake Baikal, reached Chita area and joined Ataman Semenov 's forces.

The Cossacks had been unable to organise and capitalise on their successes at the end of By they had begun to run short of supplies.

Consequently, when the Soviet counteroffensive began in January under the Bolshevik leader Antonov-Ovseenko , the Cossack forces rapidly fell apart.

The Red Army captured Kiev on 3 February General Denikin's military strength continued to grow in the spring of During several months in winter and spring of , hard fighting with doubtful outcomes took place in the Donbass , where the attacking Bolsheviks met White forces.

At the same time Denikin's Armed Forces of South Russia AFSR completed the elimination of Red forces in the northern Caucasus and advanced towards Tsaritsyn.

At the end of April and beginning of May the AFSR attacked on all fronts from the Dnepr to the Volga, and by the beginning of the summer they had won numerous battles.

French forces landed in Odessa but, after having done almost no fighting, withdrew on 8 April By mid-June the Reds were chased from the Crimea and the Odessa area.

Denikin's troops took the cities of Kharkov and Belgorod. At the same time White troops under Wrangel's command took Tsaritsyn on 17 June On 20 June Denikin issued his Moscow directive, ordering all AFSR units to prepare for a decisive offensive to take Moscow.

Although Great Britain had withdrawn its own troops from the theatre, it continued to give significant military aid money, weapons, food, ammunition and some military advisers to the White Armies during Major Ewen Cameron Bruce of the British Army had volunteered to command a British tank mission assisting the White Army.

He was awarded the Distinguished Service Order [61] for his bravery during the June battle of Tsaritsyn for single-handedly storming and capturing the fortified city of Tsaritsyn, under heavy shell fire in a single tank; this led to the capture of over 40, prisoners.

After the capture of Tsaritsyn, Wrangel pushed towards Saratov but Trotsky, seeing the danger of the union with Kolchak , against whom the Red command was concentrating large masses of troops, repulsed his attempts with heavy losses.

When Kolchak's army in the east began to retreat in June and July, the bulk of the Red Army, free from any serious danger from Siberia, was directed against Denikin.

Denikin's forces constituted a real threat and for a time threatened to reach Moscow. The Red Army, stretched thin by fighting on all fronts, was forced out of Kiev on 30 August.

Kursk and Orel were taken, on 20 September and 14 October, respectively. Vladimir Sidorin continued north towards Voronezh , but there Semyon Budyonny 's cavalrymen defeated them on 24 October.

This allowed the Red Army to cross the Don River , threatening to split the Don and Volunteer Armies.

Fierce fighting took place at the key rail junction of Kastornoye, which was taken on 15 November; Kursk was retaken two days later.

The high tide of the White movement against the Soviets had been reached in September By this time Denikin's forces were dangerously overextended.

The White front had no depth or stability—it had become a series of patrols with occasional columns of slowly advancing troops without reserves. Lacking ammunition, artillery and fresh reinforcements, Denikin's army was decisively defeated in a series of battles in October and November The Red Army recaptured Kiev on 17 December and the defeated Cossacks fled back towards the Black Sea.

While the White armies were being routed in Central Russia and the east, they had succeeded in driving Nestor Makhno's anarchist Black Army formally known as the Revolutionary Insurrectionary Army of Ukraine out of part of southern Ukraine and the Crimea.

Despite this setback, Moscow was loath to aid Makhno and the Black Army and refused to provide arms to anarchist forces in Ukraine. The main body of White forces, the Volunteers and the Don Army, pulled back towards the Don, to Rostov.

The smaller body Kiev and Odessa troops withdrew to Odessa and the Crimea, which it had managed to protect from the Bolsheviks during the winter of — By February the British government had pulled its military forces out of Central Asia.

For a time Central Asia was completely cut off from Red Army forces in Siberia. During this conference a regional bureau of Muslim organisations of the Russian Bolshevik Party was formed.

The Bolshevik Party continued to try to gain support among the native population by giving them the impression of better representation for the Central Asian population and throughout the end of the year were able to maintain harmony with the Central Asian people.

Communication difficulties with Red Army forces in Siberia and European Russia ceased to be a problem by mid-November Due to Red Army successes north of Central Asia, communication with Moscow was re-established and the Bolsheviks were able to claim victory over the White Army in Turkestan.

In the Ural-Guryev operation of —, the Red Turkestan Front defeated the Ural Army. During winter , Ural Cossacks and their families, totaling about 15, people, headed south along the eastern coast of the Caspian Sea towards Fort Alexandrovsk.

Only a few hundred of them reached Persia in June During the winter —20, the Orenburg Army retreated to Semirechye in what is known as the Starving March , as half of the participants perished.

By the beginning of the main body of the Armed Forces of South Russia was rapidly retreating towards the Don, to Rostov.

Denikin hoped to hold the crossings of the Don, then rest and reform his troops, but the White Army was not able to hold the Don area, and at the end of February started a retreat across Kuban towards Novorossiysk.

Slipshod evacuation of Novorossiysk proved to be a dark event for the White Army. Russian and Allied ships evacuated about 40, of Denikin's men from Novorossiysk to the Crimea, without horses or any heavy equipment, while about 20, men were left behind and either dispersed or captured by the Red Army.

Following the disastrous Novorossiysk evacuation, Denikin stepped down and the military council elected Wrangel as the new Commander-in-Chief of the White Army.

He was able to restore order to the dispirited troops and reshape an army that could fight as a regular force again. This remained an organized force in the Crimea throughout After Moscow's Bolshevik government signed a military and political alliance with Nestor Makhno and the Ukrainian anarchists, the Black Army attacked and defeated several regiments of Wrangel's troops in southern Ukraine, forcing him to retreat before he could capture that year's grain harvest.

Stymied in his efforts to consolidate his hold, Wrangel then attacked north in an attempt to take advantage of recent Red Army defeats at the close of the Polish—Soviet War of — The Red Army eventually halted this offensive, and Wrangel's troops had to retreat to Crimea in November , pursued by both the Red and Black cavalry and infantry.

Wrangel's fleet evacuated him and his army to Constantinople on 14 November , ending the struggle of Reds and Whites in Southern Russia.

After the defeat of Wrangel, the Red Army immediately repudiated its treaty of alliance with Nestor Makhno and attacked the anarchist Black Army; the campaign to liquidate Makhno and the Ukrainian anarchists began with an attempted assassination of Makhno by Cheka agents.

Anger at continued repression by the Bolshevik Communist government and at its liberal use of the Cheka to put down anarchist elements led to a naval mutiny at Kronstadt in March , followed by peasant revolts.

Red Army attacks on the anarchist forces and their sympathisers increased in ferocity throughout In Siberia, Admiral Kolchak's army had disintegrated.

He himself gave up command after the loss of Omsk and designated Gen. Grigory Semyonov as the new leader of the White Army in Siberia.

Not long after this Kolchak was arrested by the disaffected Czechoslovak Corps as he traveled towards Irkutsk without the protection of the army, and turned over to the socialist Political Centre in Irkutsk.

Six days later this regime was replaced by a Bolshevik-dominated Military-Revolutionary Committee. On 6—7 February Kolchak and his prime minister Victor Pepelyaev were shot and their bodies thrown through the ice of the frozen Angara River, just before the arrival of the White Army in the area.

Remnants of Kolchak's army reached Transbaikalia and joined Semyonov's troops, forming the Far Eastern army. With the support of the Japanese army it was able to hold Chita, but after withdrawal of Japanese soldiers from Transbaikalia, Semenov's position became untenable, and in November he was driven by the Red Army from Transbaikalia and took refuge in China.

The Japanese, who had plans to annex the Amur Krai , finally pulled their troops out as Bolshevik forces gradually asserted control over the Russian Far East.

On 25 October Vladivostok fell to the Red Army, and the Provisional Priamur Government was extinguished. In Central Asia, Red Army troops continued to face resistance into , where basmachi armed bands of Islamic guerrillas had formed to fight the Bolshevik takeover.

The Soviets engaged non-Russian peoples in Central Asia, like Magaza Masanchi , commander of the Dungan Cavalry Regiment, to fight against the Basmachis.

The Communist Party did not completely dismantle this group until General Anatoly Pepelyayev continued armed resistance in the Ayano-Maysky District until June The regions of Kamchatka and Northern Sakhalin remained under Japanese occupation until their treaty with the Soviet Union in , when their forces were finally withdrawn.

The results of the civil war were momentous. Soviet demographer Boris Urlanis estimated the total number of men killed in action in the Civil War and Polish—Soviet War as , , in the Red Army, , White armies and Poles and the total number of military personnel dead from disease on both sides as , During the Red Terror , estimates of Cheka executions range from 12, to 1.

William Henry Chamberlin suspected that there were about 50, Some ,—, Cossacks were killed or deported during Decossackization , out of a population of around three million.

At the end of the Civil War the Russian SFSR was exhausted and near ruin. The droughts of and , as well as the famine , worsened the disaster still further.

Disease had reached pandemic proportions, with 3,, dying of typhus in alone. Millions more also died of widespread starvation, wholesale massacres by both sides and pogroms against Jews in Ukraine and southern Russia.

By there were at least 7,, street children in Russia as a result of nearly ten years of devastation from World War I and the civil war.

The Russian economy was devastated by the war, with factories and bridges destroyed, cattle and raw materials pillaged, mines flooded and machines damaged.

The industrial production value descended to one-seventh of the value of and agriculture to one-third. According to Pravda , "The workers of the towns and some of the villages choke in the throes of hunger.

The railways barely crawl. The houses are crumbling. The towns are full of refuse. Epidemics spread and death strikes—industry is ruined. War Communism saved the Soviet government during the Civil War, but much of the Russian economy had ground to a standstill.

The peasants responded to requisitions by refusing to till the land. The exchange rate with the US dollar declined from two rubles in to 1, in With the end of the war, the Communist Party no longer faced an acute military threat to its existence and power.

However, the perceived threat of another intervention, combined with the failure of socialist revolutions in other countries—most notably the German Revolution —contributed to the continued militarisation of Soviet society.

Although Russia experienced extremely rapid economic growth [90] in the s, the combined effect of World War I and the Civil War left a lasting scar on Russian society and had permanent effects on the development of the Soviet Union.

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