Folk knowledge of invertebrates in Central Europe - folk taxonomy, nomenclature, medicinal and other uses, folklore, and nature conservation

Background There is scarce information about European folk knowledge of wild invertebrate fauna. We have documented such folk knowledge in three regions, in Romania, Slovakia and Croatia. We provide a list of folk taxa, and discuss folk biological classification and nomenclature, salient features, uses, related proverbs and sayings, and conservation. Methods We collected data among Hungarian-speaking people practising small-scale, traditional agriculture. We studied “all” invertebrate species (species groups) potentially occurring in the vicinity of the settlements. We used photos, held semi-structured interviews, and conducted picture sorting. Results We documented 208 invertebrate folk taxa. Many species were known which have, to our knowledge, no economic significance. 36 % of the species were known to at least half of the informants. Knowledge reliability was high, although informants were sometimes prone to exaggeration. 93 % of folk taxa had their own individual names, and 90 % of the taxa were embedded in the folk taxonomy. Twenty four species were of direct use to humans (4 medicinal, 5 consumed, 11 as bait, 2 as playthings). Completely new was the discovery that the honey stomachs of black-coloured carpenter bees (Xylocopa violacea, X. valga) were consumed. 30 taxa were associated with a proverb or used for weather forecasting, or predicting harvests. Conscious ideas about conserving invertebrates only occurred with a few taxa, but informants would generally refrain from harming firebugs (Pyrrhocoris apterus), field crickets (Gryllus campestris) and most butterflies. We did not find any mythical creatures among invertebrate folk taxa. Almost every invertebrate species was regarded as basically harmful. Where possible, they were destroyed or at least regarded as worth eradicating. However, we could find no evidence to suggest any invertebrate species had suffered population loss as a result of conscious destruction. Sometimes knowledge pertaining to the taxa could have more general relevance, and be regarded as folk wisdom concerning the functioning of nature as a whole. Conclusions The high number of known invertebrate folk taxa suggests that it would be worth conducting further investigations in other areas of Europe.


Background
Traditional knowledge systems about the landscape and the biota have been fundamental for human development since the times of pre-modern and pre-industrial societies in Europe. Humans living in close contact with the landscape as herdsmen and peasants have long possessed unified, systematic knowledge, including folk taxonomies, about phenomena that were of importance to them. The use and management of natural resources was based on centuries-old, often millennia-old ecological experience, on multi-generational knowledge passed down from generation to generation [1,2].
Ethnozoology is the scientific study of the dynamic relationships among people, and animals. Traditional ethnozoological knowledge has great cultural and economical importance. It is widely studied in the tropics and North America (e.g. [3][4][5]), but also in Europe (e.g. [6,7]). Wild animal-based natural resources are often among the key resources local communities depend on [8,9]. A major goal of these communities is to use and manage these resources sustainably (e.g. taboos: [10]; social rules: [11,12]). Long-term sustainability in the use and management of natural resources requires healthy ecosystems, while at the same time, sustainable management often contributes to maintaining the health of ecosystems [13,14].
The knowledge passed by local traditional communities, however, not only serves sustainable use and maintenance of the local community and its environment but may also provide valuable data, information and knowledge to science and conservation. Among the potential benefits of traditional ecological knowledge, it can help science to recognize new species (e.g. [15]), provide data on population sizes and dynamics of species that are difficult to observe [16,17], support the monitoring of ecosystem health, incl. pasture conditions [18,19], and develop efficient conservation managament strategies and practices [20][21][22][23].
There is no reason to imagine that European peasant and herder communities differ fundamentally from native societies in other parts of the world with regard to their ecological knowledge [24]. However, there is scarce information about European folk knowledge of wild invertebrate fauna, including their use in healing and nutrition. Researchers in ethnobiology seldom pay attention to invertebrates in the European context [25]. By contrast, several comprehensive studies have been conducted in other parts of the world. As early as 1887, Stearns published an ethnoconchological work on the use of shells as money among aboriginals of North America [26]. This was actually the first time the prefix "ethno-" was combined with a research field, thus preceding Harshberger's term "ethnobotany", coined in 1895 [27]. Another pioneering study was Henderson's and Harrrington's ethnozoology of the Tewa people in New Mexico. This study gives a full list of animals, including invertebrates, by order and gives their Tewa names as well as their scientific names [28]. In a comprehensive study Bodenheimer [29] reviewed the ethnographical literature of the use of insects as food worldwide. Nowadays there are several important studies available dealing with ethnobiological aspects of invertebrates. We can, for instance, mention Bentley and Rodríguez [30] on the entire invertebrate fauna of Honduras, and Krause et al. [31] on the insect fauna knowledge of the Roviana people (Solomon Islands). Gurung [32] detailed the knowledge of arthropods among Tharu farmers in Nepal, while Hemp [33] described what the peoples living near Mount Kilimanjaro (Tanzania) knew about invertebrates. A particularly impressive ethnozoological study is Morris [34], dealing with the impact of insects and their classification in Malawi folk culture. In addition, the literature on aquatic and coastal-marine invertebrates is particularly rich (e.g. [35][36][37]).
The general experience is that many invertebrate species have specific and relevant benefits or detriments, although the number of locally known folk taxa is higher than this [31]. Some culturally salient invertebrate species may even be important keystone species in the lives of certain communities. The majority of these are coastal-marine invertebrates (e.g. shellfish in British Columbia - [37]; crabs (Ucides cordatus) in Brazil - [38,39]). There are fewer culturally salient species among terrestrial invertebrates, and relatively few species have known folk uses (cf. [32,40]). Keystone species include, among spiders for example, the bird-eating spiders for Afro-Brazilians in Bahia [41], while among lepidopterans there is the Brahmaeid moth on Taiwan [42].
European folk knowledge about invertebrates has, since the nineteenth century, been researched mostly by folklorists and linguists. In 1879-80 the Swedish author Strindberg used a questionnaire to gather valuable data regarding folk names and rhymes connected with the ladybird. His research, using mapping as a method, is a pioneering work in folklore about animals [43]. An encyclopedia was published about Romanian insect folklore, including local names, legends, fables and myths, the role of insects in witchcraft, and beliefs about insects as pests or as omens [44]. Herman published the local names of insects and invertebrate pest species known by Hungarian herders [45]. We can also mention an interesting article on folk knowledge about botflies (Oestridae) found as parasites on domesticated reindeer, published by the ethnographer and linguist Wiklund [46]. This kind of ethnographic folklore-linguistic research tradition continues today in Europe. Wiggen, for instance, inspired by current ethnobiologists, has recently published an exciting study on the traditional names of lower animals in Norway [47]. In European cultures, it is generally quite uncommon to use or consume invertebrates [48,49]. The only invertebrates with any significant ethnobiological literature are for the taxa of snails [50], slugs [51], leeches [52], ladybirds [6], crustaceans [53], oil beetles [54] and head lice [55], but none of these are cultural keystone species. Here we should also mention a small but intriguing study on Sami children's knowledge and use of small invertebrates for amusement and to play with [56]. In 2006, Svanberg [57] published a small book with ethnozoological studies on the human relationship with bumblebees, earthworms, froghoppers, isopods, liver flukes, moonjellies and starfish in Scandinavia and Estonia. There is of course extensive biological literature on pests, but very little detailed documentation of folk knowledge has yet been carried out in Europe [58,59]. We are, however, of the opinion that further data may exist in local languages, in works on ethnography, local history and perhaps even linguistics, but these have not yet entered the international ethnobiological literature (e.g. [60]).
There is also very little Hungarian literature on folk knowledge of invertebrates. Linguistic (dialectic), ethnographic and ethnobiological literature is available concerning 161 invertebrate species in the Sóvidék region in Transylvania [61], 67 taxa along the Danube [62], the beetle taxa Melolontha melolontha, Lucanus cervus and Lytta vesicatoria [63], and the snail species Helix spp. [64]. Sporadic data may also appear in ethnographic and linguistic literature written in the Hungarian language, for example in monographs on farming and forest ethnography, e.g. in Hegyi [65] on Lytta vesicatoria and Melolantha melolantha. To date, nothing has been published in English about the folk knowledge of invertebrates of the Carpathian Basin.
Our article has the objective of presenting the Hungarian folk knowledge of invertebrate species uncovered in three areas of the Carpathian Basin (in Romania, Slovakia, and Croatia), including: 1) a list of folk taxa of invertebrates, 2) their folk biological classifications and nomenclatures, 3) their salient features, and 4) their uses, related proverbs and sayings, and their conservation.
This is the first article in Europe to deal comprehensively with an entire invertebrate fauna. The folk knowledge, nomenclature and uses of 208 taxa are presented in detail. The high number of known folk taxa suggests that it would be worth conducting further investigations in other areas of Europe.

Study areas
We collected data among ethnic Hungarians practising small-scale, traditional agriculture. Our (Fig. 1)). As the people we studied spend a lot of time in the fields and forests during their everyday activities, they still have a close, direct connection to their natural environment. The settlements where the data were collected, each with between 100 and 2500 inhabitants, are characterised by a large amount of abandoned agricultural land, and by ageing populations.
The three study areas are characterised by a moderate continental climate, with a mean annual precipitation of 600-700 mm. The mean annual temperature in the two northern areas is 8-8.5°C (July mean 19°C, January mean −4°C), while in Baranja, further south, it is slightly higher, around 10°C (July mean 21°C, January mean −4°C) [66]. The elevation is 75-90 m.a.s.l. in Baranja, 200-350 m.a.s.l. in Sălaj, and 190-500 m.a.s.l. in Gemer. Gemer and Sălaj typically have closed broadleaved forests (oak), while in Baranja there is a mixture of riparian vegetation, marshland and mixed hardwood gallery forests (oak, ash and elm).

Data collection and analysis
Data was collected in Sălaj in summer 2010, and in Baranja and Gemer in summer 2012. In each area, the objective was to identify and interview local people with the most extensive knowledge. We employed a number of techniques: in Sălaj we first consulted the local Calvinist priest, and then followed the snowball method; in Gemer we also followed the snowball method, but this time starting with the best informants from earlier ethnobotanical researches; in Baranja we collaborated with the local nature conservation warden, István Tórizs, to meet the people who, in the warden's view, had the greatest traditional folk knowledge. In total we interviewed 58 people. The overall average age of the interviewees was 75 years (within a range from 36 to 90 years), and the regional average ages were 78 in Sălaj, 74 in Baranja, and 71 in Gemer. All the informants retained memories of traditional forest use and smallholder farming, and some were still practitioners. 55 of the interviewees were Calvinist.
We conducted indoor interviews recorded on a dictaphone (approximately 88 h of recording), since the presentation of living specimens and direct observation of animals in the wild would have been greatly inconvenient for most of the informants. Prior informed consent was obtained before all the interviews, and ethical guidelines suggested by the International Society of Ethnobiology were followed. We studied "all" invertebrate species or species groups potentially occurring in the vicinity of the settlements under investigation. We placed an average of 12 photos of species of similar habitat and size on a sheet of A4 paper, to give interviewees a sense of the context and relative size of each taxon. In many instances during our preliminary study, the differing scale of the pictures had greatly inhibited recognition. Where ambiguous descriptions occurred, further enquiries of the characteristics of the species in question were made in order to facilitate identification of the animal at the finest possible taxonomic level. Detailed lists of invertebrate taxa documented by zoologists were available for the regions studied or for ecologically similar neighbouring regions (e.g. [67][68][69][70]). We also included a few species that do not occur in the areas under investigation, in order to check the authenticity of local folk knowledge.
In total we collected 3465 individual data records on 208 folk generics and specifics. We also conducted semistructured interviews with the majority of informants and carried out picture sorting, during which they were asked to group species according to their own systems. We used these results to reconstruct the folk taxonomy. Figures depicting taxonomic relations were prepared following the method used by Berlin [71]. Circles drawn in solid lines on these figures indicate scientific taxa (one species, one genus, one order, one family), whereas those drawn in small and large dashes represent, respectively, folk taxa and more inclusive folk categories. When circles of scientific taxa overlap, this indicates that certain scientific taxa were viewed as alike (e.g. "it is a house mouse, but of a different kind"). Inclusive categories were established on the basis of data collected by pile sorting, co-references and direct questions. However, it was not our intention to arrange individual taxa according to Berlin's system of taxonomic levels, since the communities we examined are too heterogeneous for this. For each of the taxa, where possible, we documented the local name (or names), their salient features, their uses, any damage they cause, any personal attitudes expressed towards the taxa (positive, negative or neutral), and related folklore issues. The habitats of the species (see Appendix) were determined based on the interviews, on our own experiences and on the scientific literature.
We have listed our data in tables, and summarised the results broken down according to informant and taxon. We have not carried out a quantitative comparison of the Fig. 1 The study areas in Central Europe knowledge among the three communities, for the data sets have, in many cases, low sample sizes. The differences between the three areas which are important from a qualitative aspect are presented in the chapter on results and discussion. Literal quotations are in italics, and comments by individual interviewees are separated by a slash.

Results and discussion
Folk taxa and unknown taxa The folk knowledge of invertebrates revealed in the areas under investigation was extensive and detailed. Folk generics and specifics were documented for a total of 208 invertebrate folk taxa. The majority of these were Coleoptera, Diptera, Lepidoptera, Arachnida and Hymenoptera, while Myriapoda, Crustacea and Annelida were represented with fewer folk taxa (Fig. 2).
Of the 208 folk taxa, in 135 cases (65 %) they could be identified with one or two scientific species, in 28 cases (13 %) with several (3-6) scientific species, and in 45 cases (22 %) with many (more than 6) scientific species.
Certain species were exceptionally well known, but 37.5 % of the taxa were familiar only to between 1 and 3 people (Fig. 3). With certain species or groups of species, the only informants who knew them were those most likely to encounter them because of their profession or as a result of some special activity (such as fishermen using animals as bait, or herders with livestock parasites). 45 taxa (22 %) were known to almost all the informants.
The 208 folk generics and specifics found is greater than the number of known vertebrate folk taxa ( [72], Ulicsni ined). Compared with four studies that covered entire faunas [28,30,33,73], the ratio of invertebrate to vertebrate taxa in our region was significant (54 % of specific level taxa). Apart from a single exception (bivalves-frogs, see below), the invertebrates were classified into separate supraspecific groups from the vertebrates, although invertebrates did not constitute a unified group, i.e., an inclusive folk taxon with clearly defined boundaries. This distinction is much sharper, for example, among Mongolians [74].
The differences in the fauna of the three different areas seemed to have little effect on the list of local folk taxa. The faunas of the three areas are similar, as they all contain mostly common, generalist species. The proportion of folk taxa that were restricted to just one of the three areas amounted to only 3.8 % (8 species). For this reason, our analyses treated all the data uniformly. Based on our data, the folk taxonomies could also be regardedwith negligible differencesas uniform (major differences are shown in the results and discussed below).
The distribution of knowledge was not even. Only 36 % of the species were known to at least half of the Fig. 4 Folk taxa containing bees (Apidae), as well as mining bees (Andrenidae), wasp species (Vespidae) and family of parasitoidal wasps (Sphecidae). Continuous circles on these figures indicate scientific taxa (one species, one genus, one ordo, one family), whereas small and large dashed circles represent folk taxa and more inclusive folk categories, respectively. The overlap of the circles of scientific taxa indicates that certain scientific taxa are viewed as 'alike'. Prototypic species are marked with undelining. NN means the inclusive taxon is not named informants. There are two possible reasons for this: 1) the erosion of knowledge (e.g. reductions in hand harvesting mean less familiarity with the dwarf velvet mite Microtrombidium pusillum); 2) certain species are linked to particular farming activities, and so are not generally known. A beekeeper, for instance, would have better knowledge of bee pests, a herder would be more familiar with sheep parasites. Such species may be completely unknown to other members of the community.
The reliability of the knowledge was very high. Despite carrying out constant checks using cross-questioning, errors, falsifications and slips of the tongue were only registered in very few cases. It was more likely for respondents to answer that they didn't know information or weren't familiar with species. Due to the general aversion towards the majority of invertebrate species, however, the informants were sometimes prone to exaggeration. A similarly high degree of reliability and low proportion of errors have been experienced in other Central European locations in studies of botanical knowledge [75,76]. For some species (e.g. vine louse, itch mite), there was a high proportion of knowledge that was not based on personal experience.
In line with our expectations (cf. [77,78]), larger species, those occurring more frequently and those with more distinctive morphologies were more widely known. There was also a greater degree of knowledge of species living in habitats closest to the homes of the informants. Animal and human parasites were often exceedingly well-known. Compared with knowledge of vertebrates, the majority of invertebrate taxa were less detailed. At the same time, a quarter of invertebrate taxa were known to an extent which was comparable to that of the best known vertebrate species.
It was surprising to us that so many invertebrate species are known which have, to our knowledge, no economic significance. The reasons for this were not always clear. Human lifestyles have greatly changed, so there is uncertainty concerning how important a given taxon may have been in the past (e.g. the dormouse species' , which were once regularly hunted, but which are not used at all today, [72]. Yet there were other species that we did not expect to be widely known which proved, during the study, to be significant even today. Examples are species that have appeared recently, such as Harmonia axyridis, and species of predatory mites that are particularly small, harmless and can be seen on other insects. Sometimes only the larval form of an animal is known, such as those of the click beetles (Agriotes spp.). In such cases, their place in the taxonomy was less consistent, and often haphazard. The same phenomenon was also often observed in the Sóvidék region (Romania) by Gub [61].
Also surprisingly, informants made no distinction between a significant number of diverse and morphologically easy-to-distinguish lepidopteran species. The hummingbird hawk-moth (Macroglossum stellatarum), with its remarkably unique behaviour, was a relatively frequently seen species. Despite being widely known, astonishingly, it was only given a name in one case, and even this was just the name used within the informant's family.
By comparison, in places where use is made of lepidopteran species (e.g. larvae are eaten in Mexico), up to 67 different species may be known in detail [79]. Species of the order Lepidoptera are an important food source in numerous other regions of the world [80].
We did not find any mythical creatures among invertebrate folk taxa, whereas ethnic Hungarians identify several such animal taxa among vertebrates (e.g. house snake, whistling snake), which are still considered living mythical creatures in the areas under investigation.
With recently settled invasive species or major local invasions of species with a constant lower-level presence, we found that the media played an enhanced role as a source of information. The degree of knowledge sometimes varied greatly, depending on the extent of the invasion, which resulted in some significant differences between the three areas. However, there were only a few species which were known to a varying degree in the three areas (such as the Italian tree cricket, which was more familiar in areas practising viticulture, and Simulium spp., in areas where there had previously been major invasions). Namesmain features and points of interest, unnamed species, modern names Ninety-three percent of folk taxa had their own individual folk names. The proportion of covert categories was low compared to their higher prevalence among, for example, the Matses of Peru [81]. Where the covert categories are concerned, there is a chance that a few further known folk taxa were not identified during our data collection. The descriptive names used in the case of folk specifics most frequently referred to their morphology or their habitat. A few taxa were only named with the name of the inclusive category.
With some of the taxa, the names given to them within the same community were not consistent. Names could be chopped and changed around even in the case of species that were otherwise clearly separated, such as with locusts, grasshoppers and cicadas; all three of these taxa share the ability to jump, but their size and morphology differ. Almost everybody could distinguish between the three taxa, but the names they used were sometimes swapped around. Berlin et al. [82] also found that people agreed closely on the appropriate names for some species and disagreed markedly on the names of other species.
In a few cases, two or more taxa were given an identical name, even though the fact of their separateness as taxa was widely recognised (e.g. Lampyris noctiluca, Lamprohiza splendidula and Cetonia aurata). The first two are glow worms that light up at night, while the third is a bug (rose chafer) that shines beautifully in sunlight. In our experience, if it was necessary to make a distinction between the first two and the third species, then more knowledgeable informants would, in every case, separate them by adding epithets to the name (e.g. nappali szentjánosbogár [daytime Saint John's bug], or: "the one, which is just a féreg"). In everyday speech, however, the context would determine whether the folk specific referred to the first two or to the third species, so there was no need for separate names.
On other occasions, the same folk name was used for completely unrelated and well distinguished taxonomic groups (e.g. bolha [flea]: Pulex irritans -Chaetocnema . The names of folk specifics typically made reference to morphological, habitat and ecological properties. There were also instances of the usefulness of the creature being referred to in its name (jópiókalópióka, 'good leechhorse leech' , Hirudo medicinalis -Haemopis sanguisuga). Larval forms were given separate names in several instances (e.g. Hypoderma bovis, Melolontha melolontha, Pediculus humanus capitis), even if the larva and the imago comprised the same folk taxon.
There were several taxa with multiple names. The firebug (Pyrrhocoris apterus) is a generally known species not only in the areas of our investigation, but generally in regions where Hungarian is spoken [61,62]. The reason for this may be its distinctive behaviour, or perhaps the fact that hordes of them together can be witnessed in early spring (this phenomenon often also serves as the basis for folk weather forecasts). This species was given a wide range of diverse names. This contradicts the earlier observation [77] that smaller species which cause little or no harm, and which also have no benefit, are often not given names, regardless of how common they are. The proliferation of names also contradicts the observations of Fleck et al. [81], which state, roughly, that the more salient a species is, the more uniform its name will be.
There were far fewer instances of modern names or names used by only one family or individual. Some of these names were humorous, such as pizsamás bogár [pyjama beetle] for Leptinotarsa decemlineata, or vízibizigli (paddled boat) for the waterstriders. This phenomenon has been observed, although similarly infrequently, in botanical studies [83].
Names and other types of knowledge could, in certain cases, be a hybrid of traditional and scientific knowledge. However, the overwhelming majority of the knowledge recorded in our study had a traditional, folk background. Only rarely did some names come to light which derived from formal education or from the media (e.g. aranyos virágbogár [golden flower bug] -Cetonia aurata; aranyszemű fátyolka [gold-eyed veil] -Chrysopa perla). It is more common for the official Hungarian scientific names to originate from folk names. The balance in favour of traditional knowledge is stronger for invertebrates than it is for vertebrates [72]. The influence of schooling could only be felt among a few informants and only for a very limited number of species. In Appendix, all the names used by local people which demonstrably originate from "modern" sources (school, media, books, etc.) have been underlined.

Folk taxonomy, folk nomenclature and salient features
The folk taxonomy and nomenclature for the 208 folk taxa are presented in Figs. 3, 4, 5, 6, 7, 8, 9, 10, 11, 12, 13 and 14. Further data (English equivalents, salient features, main habitats and proportion of people who knew the taxon) are contained in Appendix. 16 prototypic species have been recognised, sharing the following features: their names consisted mostly of one simple noun, and within each inclusive taxon they represented the most typical behaviour, were usually the most common species, or could serve as a basis for comparison due to some other feature.
The group containing all the hymenopteran taxa except for ants and gall wasps (Fig. 4) did not have its own separate name. Informants tended to divide this group into three parts: méhek (bees), darazsak (wasps), and dongók (bumblebees), the latter of which had a certain overlap with the méhek (bees) taxon. Prototypic species could only be identified for the first two, more stable groups.
The group called légy (fly) included a significant proportion of true fly (Diptera) species, and not a single group belonging to a different scientific taxon (Fig. 5). The dipteran folk taxa were distinguished primarily according to ecological salience, and secondarily according to morphological salience, into widely known taxa. The dipteran taxa Fannia canicularis, Stomoxys calcitrans and Haematobia irritans were not known to many informants, and could only be partly differentiated, never entirely. This state of uncertainty is reflected in the diagram with overlapping circles. We recorded knowledge of a total of 24 dipteran species, although informants did not include them all and always to the inclusive Diptera category. The sole group to contain a large number of taxa was the one referred to as bogár (beetle or bug, cf. [45]), which totalled 48 folk taxa (Fig. 6). As with many of the inclusive folk taxa, there were no sharp divisions here either. With species that do not strictly belong in the group of beetles there were further instances of the name bogár (beetle) being used, but the species that feature in Fig. 6 are the ones that could be classified with greater certainty in the folk taxon of bogár (beetle). The key attributes for classification were the hardness of the integumentary system and the shape of the species. The most common taxa in this group were those with a hard chitinous covering and those belonging to the scientific order Coleoptera.
The folk prototypic species for the entire group of beetle (bogár) were primarily the black-coloured members of the family of ground beetles (Carabidae). The prototypic species for inclusive taxa with fewer members were the seven-spot ladybird (Coccinella septempunctata), the great silver water beetle (Hydrous piceus), and so on.
There were examples of taxa at two separate levels being given the same name, even though the informants could clearly distinguish between the levels (see vízibogár [water beetle]).
The flea beetles (Chaetocnema spp.) constituted a special case. Here, the complex phenomenon was identified using a single taxon, the combined presence of a Chaetocnema species and an Erwinia bacterium species, which causes leaf dieback that forms a distinctive pattern.
Among ladybirds (Coccinellidae), informants could distinguish 5 or 6 species. The harlequin ladybird (Harmonia axyridis), a recently arrived invasive species, was almost universally known. In the year it appeared, this species was immediately noticed everywhere, and viewed as alien and harmful. The firebug (Pyrrhocoris apterus) is well known in every settlement, and has a wide variety of names (13 different names in the three areas).
Within the inclusive taxon of bogár (beetle), there were also instances of species with markedly different appearances (even to an untrained eye) being classified together. For example, the European mole cricket (Gryllotalpa Fig. 10 Folk taxa containing the larvae of some arthropods gryllotalpa) was associated with the dor beetles (Geotrupes spp.), with the reason given that these species are found close to animal faeces.
The cockchafer (Melolontha melolontha) and its larva appeared in two (sometimes three) separate places within the folk taxonomy. In addition to the separation of the larva and the imago, the caterpillar of the large white butterfly (Pieris brassicae) (and, to a lesser extent, other species of butterfly) as well as its imago were regarded as stages in the ontogenetic development of the cockchafer. This was particularly true of the Sălaj area, although other scattered data [84] indicate that this view is common among much of the ethnic Hungarian population of Transylvania.
The folk taxon containing mostly orthopteran species only differed from the scientific classification in the absence of the European mole cricket (Gryllotalpa gryllotalpa). It did, however, contain the majority of cicadas (Fig. 7). The prototypic species in this taxon was the field cricket (Gryllus campestris). The distinction between this and the Italian tree cricket (Oecanthus pellucens), and therefore the entire classification as well, differed significantly among the different areas (in Sălaj all informants knew the distinction, but only one made the distinction in Gemer).
The harvestmen (Opiliones) and cellar spiders (Pholcidae) are different groups at ordinal level, but the informants treated them as a single folk specific (Fig. 8). The waterstriders (Gerris spp.), although belonging to the Heteroptera, were also included among folk spider species.
There was justification for classifying smaller parasites, plant pests and other similar species together ( Fig. 9), although it was not possible to confine this group within an inclusive taxon that ruled out all uncertainty. The group was heterogeneous in terms of both the scientific taxonomy and the various folk saliences. The number of known species is high, and they were very accurately identified. There was a high number of taxa that had their own prototypic species [species of green aphid, chicken body louse (Menacanthus stramineus), bed bug (Cimex lectularius), head louse (Pediculus humanus capitis), human flea (Pulex irritans)].  Figure 10 shows the majority of the larvae of insect species. This was the most uncertain of the inclusive taxa, and was not regarded as an independent group by many of the informants.
Within the category of ants, there was one clearly defined folk taxon, namely the winged castes of the most diverse species of ant (Fig. 11). Figure 12 shows the ringed worms (Annelida), flatworms (Platyhelminthes) and roundworms (Nematoda). The folk taxonomy of the ringed worms completely mirrored the scientific taxonomy, even at the level of two supraspecific taxa. In the case of the flatworms and roundworms, less information is available.
Within the molluscs, the group of snails and slugs was very clearly defined (Fig. 13). The bivalves sometimes shared associations with other molluscs (in their names, for example), but they were more frequently linked with frogs. Informants whose folk knowledge had suffered from the least amount of erosion almost exclusively regarded bivalves as the eggs of certain frog species (mostly Pelophylax and Rana).
Apart from the overlap with the cockchafer (Melolontha melolontha) and the special classification of lepidopteran caterpillars, the folk taxon of lepidopterans was also quite intact, and largely in agreement with scientific taxonomy (Fig. 14). Two additional folk taxa were included here which are classified differently according to entomologists: the moth flies (Psychodidae) and the lacewings (Chrysopa spp.).
Only a few invertebrate taxa were left out of all inclusive categories. Most of these remained alone during the pile sorting exercises. They could, on very rare occasions, be sorted into one group or other, although inconsistently, and without true conviction. Such taxa included e.g. the Tisa mayfly (Palingenia longicauda) and the froghoppers (Cercopidae) (Fig. 15). Figures 3,4,5,6,7,8,9,10,11,12,13 and 14 show that 90 % of the taxa were embedded in the taxonomy, with generics and specifics dominating. The greatest degree of knowledge was connected to the more inclusive categories (and not to generics or specifics) primarily in the case of spiders, and to a lesser extent the snails, ants and lepidopterans. Berlin et al. [82] argued that biological species differ considerably in their overall distinctiveness from one another, and this differential distinctiveness leads to the formation of folk generic categories of differing degrees of perceptual importance. A significant part of the taxonomic literature, however, is about vertebrates, and the basic principles established in the literature often do not work with invertebrate groups. Among invertebrates, there is greater importance attached, for example, to prototypic species. These play an important role in taxonomic identification [81]. Nevertheless, the prototypic species were often given only brief descriptions by our informants. The reason for this may lie in the fact that these prototypic species were used as the basis for comparison. In such cases, the less typical species were the ones requiring more detailed descriptions, because they are being compared with and differentiated from the prototypic species.
Within a folk specific, we generally found species that were related from a scientific point of view. It was rare to find taxa that were far removed from each other according to scientific classification. As an example of the greatest distance, harvestmen (Opiliones) and cellar spiders (Pholcidae) (2 scientific orders) were identified as a single folk specific. The reason for this is probably because their physical structures are very similar (especially long legs). In line with previous findings [71], folk taxonomic relations were, to a significant extent, based on the morphological appearance of the taxa. The parallel with the scientific taxonomy was therefore surprisingly precise (especially in the case of ringed worms).
One interesting belief resulted in a quite remarkable taxonomy. The connection between bivalves and frogs is generally known in Sălaj, but was only reported by the most knowledgeable informants in Baranja. In the past, this knowledge may have been more widespread here as well.
The connection between the two species is even reflected in the name of the bivalves (békateknő -"frog tub"). We could not find out how or where this belief originated. It is hard to perceive any axis on the bodies of the bivalves, so it could be that they were not regarded as an animal species in their own right for this reason. When touching the body of the bivalves, the experience is similar to touching the slimy skin of a frog, and furthermore, they live in the same habitat. Thanks to the media, and perhaps from speaking to relatives who have been to the seaside, many informants have now heard of seashells. The majority of these were called kagyló (shell), and they were sharply distinguished from freshwater species. More knowledgeable informants said that the seashells were, like their freshwater equivalents, the eggs of frogs. However, slight majority recognised that they are separate species. Several informants could identify tadpoles (one of the common folk names is kutyahal -"dogfish"), although surprisingly they were unaware of their relationship with fully grown frogs. Tadpoles therefore exerted no influence on the supposed link between bivalves and frogs. In Ghimeş (Gyimes, Romania), the tadpoles of the yellowbellied toad (Bombina variegata) are used in veterinary medicine. The connection between the spawn (tojásegg), the tadpole (békapinty, frog carp?) and the mature adult is recognised for all common species of frog occurring there [85].
Sometimes species were classified not (only) according to morphological salient features, but (also) ecological and cultural features (e.g. Geotrupes spp., Gryllotalpa gryllotalpa). In other words, species which are clearly different from each other, even to an untrained eye, could sometimes be placed into the same taxonomical group. In such cases, morphology, the default first priority when making classifications [71], was replaced by ecological differentiation.
A few species were included in the taxonomy which were not universally viewed as animals, with some informants describing them as diseases rather than species of fauna. These included the liver fluke and species of mite. This uncertainty may derive from the small size of the creatures, or from the fact that they are hard to observe. The small size of the animals involved may also be the main reason behind the various explanations given for the origin of "cuckoo spit" (meadow froghopper foam nests), Erwinia infestations of maize, and gossamer. These three phenomena were regarded as structures created by the most diverse range of species, and in the Among the inclusive taxonomic categories, the one known as bogár [beetle] is closest to the "wug" taxon introduced by Brown [86]. "Wug" in the three regions studied included most invertebrate species, such as beetles, butterflies, bees and flies; it did not include molluscs, and only rarely did it also include flatworms, roundworms or ringed worms, so as a category it more or less covered the arthropods. The name férgek [worms], also often used as an inclusive category, was less readily applicable to the folk taxonomy. Sometimes the name was used for wormlike creatures, at other times it was applied to other invertebrate pests. In extreme instances, it even encompassed the house mouse, the wolf, the bear (cf. [63]), or indeed any animal regarded as harmful in any way.

Human uses of invertebrate taxa
A total of 24 invertebrate species were documented as being of direct use to humans ( Table 1). The use of invertebrates in our study areas was much less significant, than the role of plants in nutrition and medicine (e.g. [76,87,88]), or the role of insects in nutrition and medicine in other parts of the world [89,90]. Four species were used for medicinal purposes, 5 species were consumed, 11 were used as bait for fishing, and 2 species were used as playthings. Compared with the tropics (27 medicinal species [91]; more than 200 edible species [92]) this is much lower both in diversity and in terms of the body mass of the invertebrates used.
Andricus hungaricus and Cynips quercusfolii were known to be used for tanning leather, although rarely. Even less commonly, Lytta vesicatoria was mentioned as an aphrodisiac, and Daphnia spp. were used as food for aquarium fish.
Completely new was the discovery that the honey stomachs of black-coloured carpenter bees (Xylocopa violacea, X. valga) were consumed even when there was no shortage of alternative foods. This practice was previously unknown in Europe.
In the last hundred years, the consumption of invertebrates in Europe has traditionally been restricted to just a   (46) Sometime it is protected like a taboo. God' s Lamb. It has a cross on its back.
x 5. 6 Lytta vesicatoria Spanish fly (86) If the rabid dog bit someone, you had to feed nine piece of it to the man. / If you pour (the tincture prepared from the beetle) onto the head of the man, he did not shiver any more.  (20) The poor cooked it. / They were collected in springtime. / They were washed at least ten times. It was scalded and the foot cut off. It was soaked in lukewarm water, in cold water, lukewarm again, a lot of work. / Snails are best before the weeds grow too high.
x 8. 15 Astacus astacus, Astacus leptodactylus European crayfish, Danubian crayfish (48) The old of long time ago caught it, it became red when cooked. / My father caught many on the Rét (a marsh), we cooked them in a big pot. In salty water. / The tail and the nippers are good to eat.
x 9. 8 Araneae e.g. Tegenaria domestica spiders (34) If you cut your feet, you would pick spider net in the stable and covered the cut to heal.
x x x

4
Apis mellifera European honey bee (113) When the bees are gone we will be gone as well because there will be nothing to eat. / Honey is good for a lot of things.
x x x 11. 13 Bivalvia e.g. Anodonta cygnea clams (24) There were many, fed to the pigs. / You would make buttons of it. It is good for bait to catch carp and predatory fish. My husband would know them because he was a fisherman and would collect them.
x 15. 12 Eisenia fetida redworm (12) Not all earthworms would do for angling. This is the best one.
x 16. 12 Aporrectodea dubiosa earthworm species (13) This is harder and it (the fish) can not pull it down (from the hook).
x 17. 6 x few species [93], and in the areas of our investigation, they were only consumed occasionally. The consumption of nádiméz (honey in the tube of the reed) from thatched roofs was quite widespread among children, but the decline of this practice may be due to the gradual replacement of thatching as a roofing material. Molluscs are consumed relatively commonly across the globe [14,26], and this was also true for the three study areas in the past [94]. Surprisingly, the idea of consuming edible snails was mostly rejected as disgusting by the informants, and it was only among the most elderly informants in Baranja that there was any tradition of eating edible snails. The use of Spanish fly (Lytta vesicatoria) was well known, although very few informants had actually seen it used in practice. Its consumption was sometimes linked to superstitious elements such as consuming a "magic number" (9) of beetles placed in palinka (distilled fruit spirit), and mixing them with "randomly" found dog faeces. Blister beetles are used the world over to treat incurable or barely curable illnesses [95], and in our study areas, they were previously used as an antidote to rabies.
We found that leeches were used in four ways: 1) placed on the neck to reduce blood pressure, 2) for treating symptoms of periodontitis, 3) as a painkiller, by increasing the flow of blood as well as from the analgesic entering the bloodstream, 4) as a fishing bait. One species (European medicinal leech -Hirudo medicinalis) has a medicinal effect, while the other (horse-leech -Haemopis sanguisuga) does not. Detailed morphological knowledge was of great importance here. In the Carpathian Basin, it is common for medicinal and non-medicinal plant species also to be given the prefix of orvosi (medicinal) or ló-/kutya-(horse/dog), respectively [75,76].
The use of invertebrate taxa for veterinary medicine was not documented in any of the areas under investigation, although such practices are known in the region. In north-east Romania, for example, spiders are used to treat flatulence in cattle by rubbing the spider into the animal's side [Ulicsni ined.].
Contrary to our expectations, we did not document any current uses for blister beetles or slugs. The use of slugs as a lubricant of cartwheels [51] was not mentioned in our study areas. Based on other data collections, however, this practice was known in the Carpathian Basin [Molnár ined.].

Proverbs and sayings
Invertebrates are featured in a number of proverbs and sayings ( Table 2). During data collection, a total of 30  (50) There was a doctor here when we were kids who had an aquarium and he gave them to the fish. We would go to collect them with a little dipping net.

6
Gryllotalpa gryllotalpa European mole cricket (63) This is a good bait (to angle We are scared that they (Harmonia axyridis) will kill off all of our nice little ladybugs. Oh, those littles. Which is a pity, because they are good. We picked out the reed (from the roof), when we saw that there was reed honey in it. My grandmother was very angry and scored at us because we destroyed the reed roof beehive and we then ate the reed honey. We were kids and made pipe of it. It was a toy. / Oak galls would be used for tanning leather in the past.

Melitaea athalia butterflies (135)
This is indeed not a pest. We were glad to see it before. They are aware of the harm many species do, yet adult individuals are not destroyed.
x Table 2 Proverbs and common sayings referring to Invertebrata Latin name, proper name (serial number in the Appendix)

Proverbs, their meanings and explanations
Coccinella septempunctata seven-spot ladybird (21) We said to it: ladybug, where do you take me to get married? Then we married in the direction where it flew.
Joking comparison frequently quoted to kids.
Said mostly to kids with an affectionate joking gesture because of their attachment.
gossamer air-threads (44) There will be no rain because it stretches.

Vernacular weather forecast.
Microtrombidium pusillum dwarf velvet mite (46) Shine, sun, shine, Jesus' lamb is freezing to death under the gardens. And then the Sun shone.
A superstition wishing to change the weather.
Wisdom based on observations impersonating the species.
Gryllus campestris field cricket (61) You could not put down your clothing in the grass because old people said: the cricket would gnaw a hole in it.
Might be a belief.

Gryllus campestris field cricket (61)
If you draw a cross on the back of a cricket, it would not jump any more.
Fun for kids based on belief.
The firebug (Pyrrhocoris apterus) can be seen in dense masses in springtime.

Lytta vesicatoria
Spanish fly (86) The ash tree is stinky, it will rain.
Prophecy connected to Spanish fly invasion.

Melolontha melolontha cockchafer (88)
If there are too many maybeetles, corn yields will be good.
Maybe vernacular experience or possibly only a belief.
Cerambyx cerdo great capricorn beetle (90) Your moustache stands up like that of a capricorn beetle.
An analogy on the long moustache bending upwards.
Meaning of the proverb: there is no universal truth.
winged ant castes (120) e.g. Tetramorium caespitum When the winged ant comes out, it will rain.

Vernacular weather forecast.
Tetramorium caespitum and similar species pavement ant (124) Be like the ant and work! Ants were considered 'diligent' animals (busy as an ant).
Based on real observation, augmented to mythical heights (9 is a mythical number in Shamanism).

Vespula vulgaris and similar species common wasp (128)
Yellow wasp, small wasp, large wasp, they all scratch on a bunch of grapes.
Pun made of a vernacular observation.

Andricus hungaricus
Hungarian gall wasp (132) My mother made us thrown them away. You must not keep it at the house because brood will not hatch the eggs.
Belief says it prevents brooding of the hen.
Lepidoptera e.g. Melitaea athalia butterflies (135) The superstition was that if you see a yellow butterfly in spring, you would fall ill. If you see a red one, you will remain healthy and fall in love, if a black one, someone would die.
Vernacular prophecy. The yellow butterfly may be Gonopteryx rhamni, red ones may be several other species.
taxa were associated with a proverb or some other folk wisdom (such as weather forecasting, harvest predicting, similarly to the way in which birds, for instance, are associated in many human cultures [96]). Some of these were based on observations of animal behaviour or experience of their population cycles, and so do have some genuine basis in fact (e.g. the swarming patterns of Lytta vesicatoria). Other folk beliefs, however, were probably closer to old wives' tales (e.g. drawing a cross on the back of a field cricket will prevent it from jumping; the presence of Andricus hungaricus prevents hens from brooding). Forecasts of weather phenomena based on the behaviour of various invertebrates (e.g. winged ants mean that rain is coming) occurred frequently. The positive attitude towards the presence of hog lice on swine is probably based on the observation that parasites abandon sick or dead livestock. Gub [61] also found examples of healing involving external animal parasites, a practice that can also be deduced from the same kinds of observation.
Games with the invertebrates, and the ill-treatment of animals were quite widespread in the past, although they were not confined to particular species. Nevertheless, larger and more easily caught species, such as Melolontha melolontha, Lucanus cervus and Oryctes nasicornis, were more likely to fall victim. Gub [61] describes several special games involving the cockchafer and the stag beetle.
In addition to Vallejo and González [55], Gub [61] also mentions the use of head lice in human medicine, especially in treating jaundice. We did not document any similar instances, although this practice may well be widespread, and with further research there is a high chance of finding more such cases.
One belief that made a scattered appearance in the areas under investigation stated that a dragonfly hovering about the water indicated that there was no snake in the water. The name recorded for the dragonfly by Gub [61], kígyópásztor (snake-shepherd) may also derive from this belief.

Invertebrate species that enjoy folk conservation or state protection
Conscious ideas about conserving invertebrates only occurred with a few taxa (see the last column in Table 1). Seven-spot ladybirds, dwarf velvet mites and often spiders were said to enjoy protection, but informants would generallybut not universallyrefrain from harming firebugs, field crickets and most butterflies.
With regard to ladybirds, the tradition of protecting them came from the culture (songs and sayings), but Even the mottled butterfly came from a caterpillar.
You do not necessarily worth more just because of your better appearance or even something ugly may become beautiful one day.

Saturnia pyri giant peacock moth (151)
The boszorkánylepke (witch butterfly) were nailed above the door for superstition.
It was used as a superstitious protection against the Devil.
Drosophila spp. e.g. Drosophila melanogaster fruit flies (159) Fruit flies cause the wine to ferment.
In their opinion the presence of fruit flies cause the wine to ferment.
Drosophila spp. e.g. Drosophila melanogaster fruit flies (159) The man from Vörösmart swallowed the frog; he thought it was a fruit fly.

Mocking a village.
Tipula spp. e.g. Tipula maxima crane flies (160) We stroke the mosquito king to death; there will be no mosquitos now.
The Tipula species which are much greater than biting mosquitos but are related to them are presented by the saying as a kind of king.

Musca domestica housefly (169)
Noah wanted to chase them out from the Bark. He could not. Well, fly, then. He said. And the name stuck.
Folk etymology for the name of the fly. He blames Noah for the existence of flies. Légy in Hungarian also means: be (you should exist).

Pediculus humanus capitis head louse (191)
It' s not a shame to get it, only to keep it.
Educating saying on responsibility.

Haematopinus suis hog louse (193)
You can find a louse only in a good hog.
In their opinion louses occur on healthy pigs only.
It was held that wherever a dragonfly hovers over the water there will be no snakes in it. they were also recognised as useful animals. Many informants knew that they help reduce aphid populations. The taboo about destroying dwarf velvet mites was explained by a few informants as being due to the crossshaped marking on their backs. Many stated that hurting spiders brought bad luck.
Butterflies were respected for their beauty. Here it should be noted that the state protection enjoyed by certain species of butterfly (e.g. Iphiclides podalirius, Inachis io) in Hungary is justified more by their beauty than their rarity.
Field and house crickets were generally left unharmed as a result of their pleasant chirruping and their cultural significance. Surprisingly, most people knew nothing about legal protection for invertebrates.
Also surprisingly, almost every invertebrate species was regarded as basically harmful. Where possible they were destroyed or at least regarded as being worth eradicating. Informants reported little information about the benefits of invertebrates, or did not regard the benefits as significant. Because they are very common, even species that were regarded as useful were not given any protection (for example, fruit flies are believed to aid fermentation). However, we could not find any information to suggest that any invertebrate species had disappeared or become rarer as a result of conscious destruction.
In the areas under investigation, traditional uses of and attitudes towards invertebrates have not revealed any kind of activity that would cause major damage from a nature conservation point of view. The fundamental factors behind this state of sustainability are small-scale farming, which imposes less strain on the environment, and the fact that resources are mostly used locally. Traditional methods of agriculture do without chemicals, so populations of many invertebrate species only began to decline as intensive farming spread (starting in the 1980s).
With the exception of edible snails and in a few cases certain galls the use of invertebrate taxa had remained local, and was therefore sustainable. In areas where the use has spread beyond the locality, for example in Mexico, with invertebrates living in species of Agave [90], or in areas of the Carpathian Basin where edible snails are harvested in big quantity [64], a significant reduction in the prevalence of such species has been experienced. The effects of such destruction have tended to be far more significant with regard to vertebrate taxa ( [7], e.g. predatory mammals and birds).

Folk wisdom related to nature as a whole
Sometimes knowledge pertaining to the taxa could have a more general relevance, and be regarded as folk wisdom concerning the functioning of nature as a whole. The damage caused by the gypsy moth (Lymantria dispar), for example, was regarded as a minor problem, becauseaccording to many informantsmajor damage does not occur by itself "in nature", only as a result of human intervention. A kind of tolerance was exhibited, especially in connection with species that people were fond of whatever reason, or regarded as relatively harmless, in phrases such as "they have to eat too", or "they are also God's creations". The damage caused by such species is often accepted, and regarded as tolerable and natural. Certain instances of "wisdom" appeared not to originate from traditional folk knowledge. The view that "if the bees disappear, then we will disappear too, because there won't be anything to eat" probably springs from the influence of the media.
Folk wisdom in our study areas was fragmentary, probably heavily eroded, and seemed no longer to constitute a unified, systematic world view, or social conventions that impact on everyday behaviour and thinking, as has been described e.g. in connection with the ontology of Native Indian communities in North America [97][98][99].

Conclusions
Despite the fact that our material was gathered only recently, folk knowledge is still alive among Hungarian people in these regions, as are some of the folk uses. We argue, however, that before the dual impact of the market economy and public education became so powerful, Hungarian rural people might have possessed knowledge as deep as that of, for example, the natives of Amazonia. Ethnographic works from the late 19th and early 20th centuries provide the basis for this argument. The high number of known invertebrate folk taxa documented in our three study areas suggests that it would be worth conducting further investigations in other areas of Europe as well.
Local traditional ecological knowledge of invertebrates is highly relevant to helping us understand the mentality and worldview of local people. Understanding local worldviews can be a first step towards developing locally appropriate, culture-specific nature conservation strategies and local school curriculadesperately needed in our globalising world.

Appendix
For each of the taxa, we collected and documented the local name (or names), their salient features, their uses, any damage they cause, any personal attitudes expressed towards the taxa (positive, negative or neutral), and related folklore issues. The habitats of the species were determined based on our own experiences, on the interviews, and on the scientific literature. Literal quotations are in italics, and comments by individual interviewees are separated by a slash.             pécsik, pécsiklégy, bögöly, lólégy (horse légy), bogaraztató légy (make-jump légy) It bites the cows in summer. Horses even more. Greyish. Bite more before rain. / From beginning, mid-July up to mid-August is the season when they attack livestock in big numbers.  This attacks the livestock, cows intensively. They say, it may happen that the cow will perish when bitten. / This fly was here in 1938 and it was drummed out that everybody should take care and not go to the meadows too frequently, because you will get bitten. / We put (dry) manure in a pot and walked around the grandma so she could hoe and this fly did not hurt her. We made smoke of it.