Introduction
Lacewings in a broad sense – the Neuropterida – is a small superorder of holometabolous insects that contains ca. 6,500 extant species and is distributed worldwide. The superorder is comprised of three orders, snakeflies (Raphidioptera), alderflies and their relatives (Megaloptera), and lacewings in a narrow sense (Neuroptera).
Geographical and Geopolitical Divisions of Albania
Albania is located in southeastern Europe in the southwestern part of the Balkan Peninsula. From a biogeographical point of view, Albania is composed of two biogeographic regions: the Mediterranean and the Alpine. The country, which contains high mountains, hills, and coastal and river plains, provides a variety of habitats for numerous plant and animal species. Protected natural areas in Albania include 15 national parks, which cover a surface area of 1,177 km2, or approximately 4.1% of the territory of the country.
In 2000 Albania was divided into 12 counties (Figure 1), which replaced the districts into which the country was formerly divided. The counties are used in the Neuropterida of Albania fauna project because they represent the current de facto first-order geopolitical subdivisions of the country.
| 1 – Berat |
| 2 – Dibër |
| 3 – Durrës |
| 4 – Elbasan |
| 5 – Fier |
| 6 – Gjirokastër |
| 7 – Korçë |
| 8 – Kukës |
| 9 – Lezhë |
| 10 – Shkodër |
| 11 – Tirana |
| 12 – Vlorë |
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History of the Study of Albanian Neuropterida
The first mention of a neuropterid species from Albania dates to the end of the 19th century, when Albarda (1891) recorded Ornatoraphidia flavilabris (Costa) from the country in his review of the then-known snakefly species of the world. Early in the 20th century, Penther (1914) reported on a zoological collecting trip to the border region between present-day Montenegro and Albania. Among other insects collected, one of the most interesting finds was a male snakefly specimen that was described five decades later as a new species. The type locality of that species, which is now known to be in Albania, has in the past been incorrectly attributed to Montenegro (Aspöck & Aspöck 1964; Penther 1914; Devetak & Rausch 2016). Three decades after Albarda’s work, Pongrácz (1923) published the first overview of the neuropterid fauna of Albania. In his list of lacewings then known from the territory that now comprises Albania, Kosovo and Montenegro, one raphidiopteran and 11 neuropteran species were recorded from localities now located within Albania. Navás (1923, 1932) subsequently added three additional species to the fauna.
In the middle of the 19th century, Capra (1945) documented the occurrence of six neuropteran species in Albania, including the description of a new antlion species, Cueta albanica, which is today considered a junior synonym of Cueta lineosa. The decade of the 1960’s was a particularly important period in the development of knowledge about the Albanian neuropterid fauna. Steinmann (1963, 1964) reported two raphidiopteran species from the country (one of which was subsequently determined to have been collected in Kosovo, which now lies outside of Albanian territory). Zelený (1964) published the second overview of the Albanian neuropterid fauna (reporting 24 species), which was based primarily on the collections of the 1961 entomological expedition to Albania organized by the German Entomological Institute. In the same year, Bartoš (1964) reported two additional raphidiopteran species for the country, and Aspöck & Aspöck (1964) described the snakefly Phaeostigma thaleri as a new species from Albanian material.
During the second half of the 20th century several papers containing miscellaneous data on Albanian raphidiopterans were published by Austrian neuropterists H. Aspöck, U. Aspöck and H. Rausch (Aspöck & Aspöck 1965, 1966, 1974; Aspöck et al. 1973, 1976, 1980, 1989, 1991). The description of the only Balkan species of the family Nevrorthidae, Nevrorthus apatelios, which is known from Albania (Aspöck et al. 1977, Jones & Devetak 2009), also dates from this period. The publications of this period were very important for clarifying many of the taxonomic problems that were then present among the species of the Balkan fauna.
In the 21st century, a new era of the study of Albanian lacewings has started. The catalogue of Palearctic Neuropterida by Aspöck et al. (2001) listed thirty-one neuropteran and four raphidiopteran species from Albania. In a review of Balkan owlflies, Popov (2004) reported two species of that family from the country, and the first alderfly species was recently reported from the country (Dvořák 2016). Between 2007 and 2016 several expeditions were carried out by a number of neuropterists with the goal of improving knowledge of the neuropterid fauna of Albania. In the period from 2007 to 2014 field biologists from the Hungarian Natural History Museum (Budapest) participated in multiple collecting trips to the Balkan countries, including Albania. Sziráki (2014) presented new Albanian occurrence records of Raphidioptera based on the results of those trips. Between 2012 and 2014 three Albanian expeditions were organized by Slovenian zoologists from the Department of Biology of the University of Maribor, which resulted in a number of papers on the fauna (Devetak & Janžekovič 2012; Devetak et al. 2012, 2013; Klokočovnik et al. 2014, 2016; Podlesnik et al. 2016). And finally, from 2012 to 2016, Austrian entomologists Hubert and Renate Rausch conducted three expeditions to several Balkan countries, including Albania. The results of the Slovenian and Austrian expeditions, together with all previous literature records, were summarized by Devetak & Rausch (2016) in the third comprehensive overview of Albanian Neuropterida.
Neuropterida of Albania
The current number of valid neuropterid species know from Albania and each of its counties is shown in Table 1. The dedicated neuropterological expeditions undertaken during the last few years have revealed a rich fauna, which currently comprises ca. 15% Raphidioptera, 1% Megaloptera, and 84% Neuroptera species (Table 2). Zoogeographically, the Albanian fauna can be divided into four groups of species based upon their overall distributions: Eremial (few species), Mediterranean, European, and “widely distributed” (Palearctic, Euro-Siberian, and Euro-Mediterranean) (Aspöck et al. 2001). It is expected that additional species in the Mediterranean and widely-distributed groups will be discovered in Albania in the future. A few exceptional regional discoveries may also be expected, such as the spectacular recent rediscovery of the snakefly species Phaeostigma thaleri, one hundred years after its original discovery and fifty years after its first description. Females of P. thaleri have now been collected for the first time and larvae and adults have been reared from eggs of captive females (Rausch, unpublished).
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Discussion
Current knowledge of the Albanian lacewing fauna
Species of special interest include the antlions Cueta lineosa and Myrmeleon hyalinus, the Albanian records of which represent the northernmost known localities for these species on the Balkan Peninsula. Despite the fact that the Albanian neuropterological fauna is well investigated relative to the neighboring countries of Montenegro, Kosovo, and North Macedonia (Table 2), one can expect the future discovery of additional species within its borders. We estimate that approximately three quarters of the lacewing species that actually live in Albania have so far been recorded. Higher taxa that are currently not reported, but which are expected to occur within the country include spongillaflies (Sisyridae) and spoonwings (Nemopteridae). It is expected that these additional families will be found in grassland habitats in eastern Albania (spoonwings) and along rivers and natural lakes (spongillaflies), habitats that are currently under sampled. Additionally, mountain habitats have not been explored sufficiently. Among the countries that neighbor Albania, Greece has the largest number of neuropterid species (Table 2). Its higher biodiversity may be explained by its larger size (ca. four times the size of Albania), its geographical position as the southernmost Balkan country, its diverse terrain (particularly its numerous islands), and its eastern border abutting Turkey, which provides geographic continuity to the diverse neuropterid fauna of the Middle East.
Threats to the Neuropterida of Albania
Lowland river ecosystems contain fine sand microhabitats that are preferred by antlion larvae (e.g. Cueta lineosa, Myrmeleon inconspicuus). Planned dams and associated hydroelectric power stations threaten Europe’s last wild rivers. Current plans to build a series of dams along the Vjosa and Osumi rivers would jeopardize local lowland species, including antlions, and alter the river ecosystem. In other areas of the country, such as Divjake Karavasta National Park, tourism is the main threat to antlions (Myrmeleon hyalinus, Synclisis baetica, Acanthaclisis occitanica) through recreational disturbance of the sand dune ecosystems in which the larvae live.
Methods
In addition to records from literature and natural history collections, the faunal lists presented here contain records of specimens collected by the author and his associates and colleagues (see Acknowledgments) that have not previously been published. Adult neuropterids were collected using an insect net or were picked by hand from surfaces near a light source. Antlion larvae were excavated from their pits using a spoon or collected by sifting the substrate. Specimens were preserved either in ethanol or dry (pinned or pointed) and most are deposited in the author’s collection in Maribor.