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Corresponding author: Andrea Buffagni ( buffagni@irsa.cnr.it ) Academic editor: Ben Price
© 2025 Andrea Buffagni, Carlo Belfiore.
This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Citation:
Buffagni A, Belfiore C (2025) Biodiversity of Italian freshwaters: an updated checklist of mayfly species (Ephemeroptera) as a starting point for the next taxonomic (r)evolution. ZooKeys 1239: 257-280. https://doi.org/10.3897/zookeys.1239.147826
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The study of biodiversity and ecosystems must be based on detailed knowledge of animal and plant organisms. European strategies for the protection of biodiversity and the restoration of natural environments advocate in-depth studies where knowledge is most lacking or fragmentary, in order to adequately assess relevant changes. Italian Ephemeroptera have not been the subject of specific taxonomic studies for approximately 30 years. This paper presents a list of the species currently thought to occur in Italy, which amounts, on the basis of morphology, to 106 species. Of these, approximately one-fifth are endemic species, demonstrating the importance of the Italian territory for European and Mediterranean biodiversity. The main critical aspects are discussed with reference to both the recent history of the study of Italian Ephemeroptera and the prospects for future development. The general picture shows that i) many species are insufficiently studied and known; ii) there are probably several new species to be described; iii) several endemic species described in the last decades or centuries require confirmation of validity and/or presence. Specific studies on mayflies will need to focus on these aspects in order to increase our knowledge of natural systems and enable their appropriate protection and restoration.
Endemic species, identification uncertainty, Italy, Nature Restoration Law, taxonomy
Where species-level organism identifications are available, evidence indicates that global changes are driving widespread extinctions at unprecedented rates across all groups of organisms (
The most recent comprehensive lists of Italian Ephemeroptera date back to 2003 (
As part of its Biodiversity Strategy, the EU has enacted the Nature Restoration Law (EU 2024/199), introducing new regulations to achieve “the long-term and sustainable recovery of biodiverse and resilient ecosystems [...] through the restoration of degraded ecosystems.” Member States are required to implement effective, area-based restoration measures to cover at least 20% of the EU’s land area—including freshwater ecosystems—by 2030, and to restore all ecosystems in need of intervention by 2050. However, any strategy aimed at protecting biodiversity or restoring natural areas and ecosystems will be ineffective unless underpinned by robust knowledge of the organisms that form the foundation of biodiversity assessments. Comprehensive understanding of the animal and plant species within a given environment is critical for evaluating its current status, detecting changes driven by global or local factors, and assessing the success of restoration measures. With this in mind, Italy has set up the National Biodiversity Future Center (NBFC:
The primary objective of this paper is to provide a complete list of Ephemeroptera species currently believed to occur in Italy, based on traditional morphological methods. Additionally, it aims to offer a brief description of the most interesting or controversial cases related to certain species. This work is intended to serve as a foundation for future revisions and insights derived from an integrative taxonomy approach. In other words, the goal is to establish a “baseline” or “time zero” that will allow for comparison with the results of ongoing and future research on Italian Ephemeroptera. Furthermore, in order to conduct effective revisions using barcoding and other genetic techniques, it is essential to first establish a starting point based on the conclusions reached by experts during the past decades using traditional approaches. This will help to ensure that any new findings are consistent with existing taxonomic knowledge and with the observed distribution of taxa. Given the high proportion of endemic species in Italy (e.g.,
To compile the species list, we relied exclusively on morphological information and descriptions. Reports and potential advancements from molecular biology (e.g., barcoding or metabarcoding) were intentionally excluded unless they aligned with both genetic and morphological approaches, contributed to a morphological description of the collected or reported taxa, and were published by specialists in the order (e.g.,
The species list presented here is primarily based on the foundational work of
With a few rare exceptions, no indication is given of the extinction risk of the species and its IUCN category. In fact, it is not possible to provide reliable information at this stage, given the obvious need for taxonomic revision.
The book by
List of Ephemeroptera species currently believed to be present in Italy based on morphology. The information is given overall for Italy and for each of the six macro-areas defined in the work to describe the distribution patterns of the species. The fourth column indicates whether the species is considered inquirenda, while the sixth column indicates whether it is endemic. At the bottom of the table, the number of species found overall and of endemic species is given, for Italy and for each of the macro-areas. The percentage of endemic species is also given, both in general terms per macro-area, and indicating the percentage of species found only in each of the six macro-areas. The asterisk preceding the species name indicates that the presence of the species must be confirmed, as it has not been found since the catches that allowed it to be described/reported.
# | Family | Species | Species inquirenda | Italy | Endemic | Lowland and hilly areas of the Po basin | the Alps | Northern Apennines | Central-southern Apennines | Sicily | Sardinia |
---|---|---|---|---|---|---|---|---|---|---|---|
1 | Siphlonuridae | Siphlonurus lacustris Eaton, 1870 | x | x | x | x | x | x | |||
2 | Ametropodidae | Ametropus fragilis Albarda, 1878 | x | x | |||||||
3 | Baetidae | Acentrella sinaica Bogoescu, 1931 | x | x | x | x | x | ||||
4 | Alainites muticus (Linnaeus, 1758) | x | x | x | x | x | x | x | |||
5 | Alainites bengunn Yanai & Gattolliat, 2022 | x | x | x | |||||||
6 | Baetis alpinus (F. J. Pictet, 1843) | x | x | x | x | x | x | ||||
7 | Baetis buceratus Eaton, 1870 | x | x | x | x | x | x | ||||
8 | Baetis cyrneus Thomas & Gazagnes, 1984 | x | x | x | x | ||||||
9 | Baetis fuscatus (Linnaeus, 1761) | x | x | x | x | x | x | x | |||
10 | Baetis ingridae Thomas & Soldán, 1987 | x | x | x | |||||||
11 | Baetis liebenauae Keffermüller, 1974 | x | x | ||||||||
12 | Baetis lutheri Müller-Liebenau, 1967 | x | x | x | x | x | |||||
13 | Baetis melanonyx (F. J. Pictet, 1843) | x | x | x | x | x | x | ||||
14 | Baetis nubecularis Eaton, 1898 | x | x | ||||||||
15 | Baetis pavidus Grandi, 1951 | x | x | x | x | x | |||||
16 | Baetis rhodani (F. J. Pictet, 1843) | x | x | x | x | x | x | x | |||
17 | Baetis vardarensis Ikonomov, 1962 | x | x | x | x | ||||||
18 | Baetis vernus Curtis, 1834 | x | x | x | x | x | |||||
19 | Centroptilum luteolum (O. F. Müller, 1776) | x | x | x | x | x | x | ||||
20 | Cloeon dipterum (Linnaeus, 1761) | x | x | x | x | x | x | x | |||
- | Cloeon languidum Grandi, 1959 | x | (x) | (x) | (x) | ||||||
- | Cloeon praetextum Bengtsson, 1914 | x | (x) | (x) | (x) | ||||||
21 | Cloeon simile Eaton, 1870 | x | x | x | x | x | x | ||||
22 | Nigrobaetis digitatus (Bengtsson, 1912) | x | x | x | x | ||||||
23 | Nigrobaetis niger (Linnaeus, 1761) | x | x | ||||||||
24 | Procloeon bifidum (Bengtsson, 1912) | x | x | x | x | x | |||||
25 | *Procloeon calabrum (Belfiore & D’Antonio, 1990) | x | x | x | |||||||
- | Procloeon forlivense Grandi, 1964 | x | (x) | (x) | (x) | ||||||
- | Procloeon lacustre (Eaton, 1885) | x | (x) | (x) | (x) | ||||||
- | Procloeon nemorale (Eaton, 1885) | x | (x) | (x) | (x) | ||||||
26 | Procloeon pennulatum (Eaton, 1870) | x | x | x | x | ||||||
27 | Procloeon pulchrum (Eaton, 1885) | x | x | x | x | x | |||||
28 | Oligoneuriidae | Oligoneuriella rhenana (Imhoff, 1852) | x | x | x | x | |||||
29 | Heptageniidae | Anapos zebratus (Hagen, 1864) | x | x | x | ||||||
30 | Ecdyonurus alpinus Hefti, Tomka & Zurwerra, 1987 | x | x | ||||||||
31 | Ecdyonurus belfiorei Haybach & Thomas, 2001 | x | x | x | x | x | x | ||||
32 | Ecdyonurus bellieri (Hagen, 1860) | x | x | x | x | ||||||
33 | Ecdyonurus corsicus Esben-Petersen, 1912 | x | x | x | |||||||
34 | Ecdyonurus helveticus Eaton, 1883 | x | x | x | x | x | x | ||||
35 | Ecdyonurus macani Thomas & Sowa, 1970 | x | |||||||||
36 | Ecdyonurus picteti (Meyer-Dür, 1864) | x | x | ||||||||
37 | Ecdyonurus ruffii Grandi, 1953 | x | x | x | |||||||
38 | Ecdyonurus venosus (Fabricius, 1775) | x | x | x | x | x | x | ||||
39 | Ecdyonurus zelleri Eaton, 1885 | x | x | ||||||||
40 | Electrogena brulini Wagner, 2017 | x | x | x | |||||||
41 | Heptageniidae | Electrogena calabra Belfiore, 1995 | x | x | x | ||||||
42 | Electrogena fallax (Hagen, 1864) | x | x | x | |||||||
43 | Electrogena grandiae (Belfiore, 1981) | x | x | x | x | ||||||
44 | Electrogena gridellii (Grandi, 1953) | x | x | ||||||||
45 | Electrogena hyblaea Belfiore, 1994 | x | x | x | |||||||
46 | Electrogena lateralis (Curtis, 1834) | x | x | x | x | x | x | ||||
47 | Electrogena lunaris Belfiore & Scillitani, 1997 | x | x | x | |||||||
48 | Electrogena ujhelyii (Sowa, 1981) | x | x | ||||||||
49 | Epeorus alpicola (Eaton, 1871) | x | x | ||||||||
50 | Epeorus assimilis Eaton, 1885 | x | x | x | x | x | |||||
51 | Epeorus yougoslavicus (Šamal, 1935) | x | x | x | |||||||
52 | Heptagenia coerulans Rostock, 1878 | x | x | x | |||||||
53 | Heptagenia longicauda (Stephens, 1836) | x | x | x | x | ||||||
54 | Heptagenia sulphurea (O. F. Müller, 1776) | x | x | ||||||||
55 | Rhithrogena adrianae Belfiore, 1983 | x | x | x | |||||||
56 | Rhithrogena alpestris Eaton, 1885 | x | x | ||||||||
57 | Rhithrogena degrangei Sowa, 1969 | x | x | ||||||||
58 | Rhithrogena dorieri Sowa, 1971 | x | x | ||||||||
59 | Rhithrogena fiorii Grandi, 1953 | x | x | x | x | ||||||
60 | Rhithrogena hybrida Eaton, 1885 | x | x | x | x | ||||||
61 | Rhithrogena johannis Belfiore, 1990 | x | x | x | x | ||||||
62 | Rhithrogena loyolaea Navás, 1922 | x | x | x | x | ||||||
63 | Rhithrogena nivata (Eaton, 1871) | x | x | ||||||||
64 | Rhithrogena nuragica Belfiore, 1987 | x | x | x | |||||||
65 | Rhithrogena reatina Sowa & Belfiore, 1984 | x | x | x | |||||||
66 | Rhithrogena savoiensis Alba-Tercedor & Sowa, 1987 | x | x | ||||||||
67 | Rhithrogena semicolorata (Curtis, 1834) | x | x | x | x | x | x | ||||
68 | *Rhithrogena siciliana Braasch, 1989 | x | x | x | x | ||||||
69 | Leptophlebiidae | Choroterpes borbonica Belfiore, 1988 | x | x | x | x | |||||
70 | Choroterpes picteti (Eaton, 1871) | x | x | x | x | ||||||
71 | Euthraulus balcanicus Ikonomov, 1961 | x | x | x | |||||||
72 | Habroleptoides auberti Biancheri, 1954 | x | x | ||||||||
73 | Habroleptoides confusa Sartori & Jacob, 1986 | x | x | x | x | x | |||||
74 | Habroleptoides modesta (Hagen, 1864) | x | x | x | |||||||
75 | Habroleptoides pauliana (Grandi, 1959) | x | x | x | x | ||||||
76 | Habroleptoides umbratilis (Eaton, 1884) | x | x | x | x | ||||||
77 | Habrophlebia consiglioi Biancheri, 1959 | x | x | x | |||||||
78 | Habrophlebia eldae Jacob & Sartori, 1984 | x | x | x | x | x | x | ||||
79 | Habrophlebia fusca (Curtis, 1834) | x | x | x | |||||||
80 | Habrophlebia lauta Eaton, 1884 | x | x | x | |||||||
81 | Paraleptophlebia ruffoi Biancheri, 1956 | x | x | x | x | ||||||
82 | Paraleptophlebia submarginata (Stephens, 1836) | x | x | x | x | ||||||
83 | Thraulus bellus Eaton, 1881 | x | x | x | |||||||
84 | Ephemeridae | Ephemera danica Müller, 1764 | x | x | x | x | x | ||||
85 | Ephemera glaucops Pictet, 1843 | x | x | x | x | ||||||
86 | Ephemera vulgata Linnaeus, 1758 | x | x | ||||||||
87 | Ephemera zettana Kimmins, 1937 | x | x | ||||||||
88 | Polymitarcyidae | Ephoron virgo (Olivier, 1791) | x | x | x | x | |||||
89 | Potamanthidae | Potamanthus luteus (Linnaeus, 1767) | x | x | x | ||||||
90 | Ephemerellidae | Ephemerella ignita (Poda, 1761) | x | x | x | x | x | x | x | ||
91 | Ephemerella mucronata (Bengtsson, 1909) | x | x | ||||||||
92 | Serratella ikonomovi (Puthz, 1971) | x | x | x | |||||||
93 | Torleya major (Klapálek, 1905) | x | x | x | |||||||
94 | Caenidae | Brachycercus harrisellus Curtis, 1834 | x | x | x | ||||||
95 | Caenis belfiorei Malzacher, 1986 | x | x | x | |||||||
96 | Caenis beskidensis Sowa, 1973 | x | x | x | |||||||
97 | Caenis horaria (Linnaeus, 1758) | x | x | x | x | ||||||
98 | Caenis lactea (Burmeister, 1839) | x | x | ||||||||
99 | Caenis luctuosa (Burmeister, 1839) | x | x | x | x | x | x | ||||
100 | Caenis macrura Stephens, 1836 | x | x | ||||||||
101 | Caenis martae Belfiore, 1984 | x | x | x | x | x | x | ||||
102 | Caenis pseudorivulorum Keffermüller, 1960 | x | x | x | x | ||||||
103 | Caenis pusilla Navás, 1913 | x | x | x | x | x | |||||
104 | Caenis robusta Eaton, 1884 | x | x | x | |||||||
105 | *Caenis valentinae Grandi, 1951 | x | x | x | |||||||
106 | Prosopistomatidae | *Prosopistoma pennigerum (O. F. Müller, 1785) | x | x? | |||||||
Number of species | 106 | 29 | 63 | 29 | 55 | 56 | 33 | 23 | |||
Number of endemic species | 5 | 0 | 6 | 13 | 6 | 10 | |||||
% endemic species (Italy and Med) | 27.4 | 7.9 | 0.0 | 10.9 | 23.2 | 18.2 | 43.5 | ||||
No. endemic species present only in one area | 3 | 0 | 1 | 5 | 3 | 8 | |||||
% endemic species present only in one area | 4.8 | 0.0 | 1.8 | 8.9 | 9.1 | 34.8 |
Given the importance of establishing verifiable references for the identification of species present in Italy, a specimen vouchering process supported by COI barcoding (AB, unpublished) was initiated as part of the ITINERIS project, with best-practice procedures for archiving of selected Ephemeroptera based on GBIF and Darwin Core standards.
At least two primary needs can be identified when drafting species distributions. The first is to provide highly detailed information about the specific areas and locations where species have been found. The second is to identify and summarise broader distribution patterns, providing a general overview and allowing comparison and correlation of data across species. Due to the absence of a centralised repository for Ephemeroptera distribution data and the resulting gaps in detailed knowledge for many areas, the second approach has traditionally been preferred. In
For this work, we opted for an intermediate scale, dividing the Italian territory into six main areas. These are: the Alps, the lowland and hilly areas of the Po basin (≈ < 300 m, including the lowlands of Friuli-Venezia Giulia and the lower Adige catchment), the northern Apennines, the central-southern Apennines, Sardinia, and Sicily (Fig.
The distinction between the Alps/Apennines and the lowland/hilly areas of the Po basin is based on ecological factors that influence species distribution. More specifically, to define the boundary between the lowland/hilly areas and the mountain areas, we used the boundaries already defined for hydro-ecoregions (
Table
In total, we report the presence of 106 species of Ephemeroptera in Italy. Of these, 29 species are endemic, accounting for approximately 27% of the total. Among the six macro-areas, the Po Valley hosts the highest number of species, with 63 recorded, followed by the central-southern and northern Apennines, reporting 56 and 55 species, respectively. Approximately 30 species are reported for both Sicily and the Alpine region, while Sardinia has the lowest total, with just 23 species.
An examination of the number of species endemic to Italy or southern Europe by macro-area reveals interesting patterns. In the Po Valley, only five endemic species are recorded, likely reflecting the overlap of species common to other European regions. The northern Apennines show ~ 11% endemic species. Endemism rises significantly in Sicily, where it exceeds 18%, and in the central-southern Apennines, where it reaches 23%. Sardinia, despite hosting the fewest species overall, exhibits a remarkably high level of endemism, with > 40% of its species classified as endemic.
Focusing on endemic species that are unique to a single area (i.e., present in only one of the six macro-areas adopted here to present species distributions) provides further insights. Overall, as many as 20 of the 29 endemic species believed to be present in Italy are found in only one of the six macro-areas. In the Po Valley and the northern Apennines, the proportion of species endemic to these regions remains < 5%. This figure increases to nearly 10% in the central-southern Apennines and Sicily. Sardinia stands out with nearly 35% of its species being endemic solely to the region (including Corsica), making it the richest biodiversity hotspot for endemic mayflies in Italy. The high levels of endemism observed in areas such as Sardinia and Sicily suggest the potential for undiscovered species, particularly in Sicily, where additional endemic Ephemeroptera may still await description. In fact, to date not all families/genera have been studied with the same emphasis. These findings underscore the importance of targeted biodiversity studies to further refine our understanding of Italian mayfly diversity and endemism.
According to
Since the most recent publication providing synoptic information on Italian Ephemeroptera (
The following section provides commentary on some of the mayfly species found in Italy, aiming to highlight potential issues or critical situations of various kinds. For the four most represented families—Heptageniidae, Baetidae, Leptophlebiidae, and Caenidae—the species-specific comments are grouped within the same paragraph for clarity and coherence. With a few exceptions, we have not made a comparison with the distribution of species in neighbouring countries. Since the Ephemeroptera fauna of these countries is usually better known than that of Italy, we do not believe that such a comparison would have been particularly valuable, especially at this stage of taxonomic revision in Italy. Moreover, such an activity would have implied a different approach to the work, which is not in line with the proposed objectives.
The situation of Ametropus fragilis is discussed in greater detail than other species because it highlights some interesting and potentially generalisable challenges regarding the detection and actual presence of rare taxa in Italy.
In the recent history of Ephemeroptera research, Ametropus fragilis was first collected in the lower stretch of the Adige River during the winter of 1993–1994 (
The fragmented history of Ametropus fragilis carries important implications. For certain species, extinction may be only apparent, and populations may persist undetected for decades in specific conditions. For example, nymphs of A. fragilis were recorded for the first time in the Ipeľ (Ipoly) river on the Slovak-Hungarian border (
The taxonomic status within the family Baetidae is both complex and dynamic. Molecular methods have revealed the existence of numerous putative and/or undescribed species across various genera and species groups (
Baetis pavidus, was once prevalent and relatively abundant in the Po Valley. However, during the past decade, its distribution in northern Italy, including its type locality, has declined sharply—likely as a consequence of climate change—with no recent specimens available from this region. The species has also nearly vanished from central Italy. In contrast, some specimens collected in Sicily have been identified as presumably belonging to this species.
Procloeon lacustre (Eaton, 1885) and Procloeon nemorale (Eaton, 1885), belonging to the subgenus Pseudocentroptilum Bogoescu, are not included in the list of Italian mayflies because their status is essentially unclear and we have never directly examined material of these species. Both species were described by Eaton in 1885 and have not been reported since. Similarly, Procloeon forlivense (Grandi, 1964) and Cloeon languidum Grandi, 1959, although described from Italy by an Italian author, are not included in the list. These four species, together with Cloeon praetextum Bengtsson, 1914, are considered as species inquirenda by
Alongside the genus Anapos Yanai & Sartori, 2017, represented by a single species in Sardinia, the only other genus in this family with a reasonably accurate understanding of its species composition is Electrogena Zurwerra & Tomka, 1985. This genus has been the focus of extensive research over the years (e.g.,
As far as the taxa of the genus Ecdyonurus present in Italy are concerned, a few species are easily recognised in both adult and larval stages and have a clear status i.e., E. alpinus, E. belfiorei, and E. corsicus, while for all other species taxonomic refinements with the selection of new diagnostic characters would be appropriate. In many cases, a careful analysis of the distributional ranges of different taxa is likely to be an effective aid in interpreting the taxa actually present, making it possible to highlight any discontinuities or gradients in intraspecific morphological variability.
As far as the Rhithrogena genus, we report here the presence of R. savoiensis, which is new for Italy (upstream reaches of river Marecchia, 22.6.2019, C. Belfiore leg. det.). In general, a few species of the genus have a clear status i.e., R. adrianae, R. johannis, R. nivata, R. nuragica, R. reatina, and R. savoiensis. For all the other species, the specific name and attribution should be verified. Consequently, all information available to date on the distribution and ecology of the various species of the genus reported for Italy must be considered with great caution, given the inherent degree of uncertainty of the specific identification itself. According to
With regard to this family, while being aware that identification at the larval stage is sometimes based on characters that should be reviewed, the situation- compared to other families- seems clearer. For the status of two species, Habroleptoides modesta (Hagen, 1864) and Habroleptoides umbratilis (Eaton, 1884), clarification would be appropriate in general, not only for Italy. Paraleptophlebia ruffoi Biancheri, 1956 and Habroleptoides auberti (Biancheri, 1954) would require further investigation regarding their distribution. The taxon Euthraulus balcanicus Ikonomov, 1961, is clearly distinct from the related species found in Italy, although it is considered a species inquirenda (
In general, there are probably more species in the macrura group than those currently listed. However, the overall situation and the potential for describing new species is complicated by the high morphological similarity between the taxa, especially in the larval stages, which makes it difficult to highlight the species really present in Italy. A similar argument to that made for the Baetidae, i.e., that some species would be more appropriately considered as groups of species, probably applies to some species of Caenidae. For example, it is very likely that Caenis martae Belfiore, 1984 comprises several cryptic species occurring in different parts of Italy, sometimes even in sympatry.
Caenis belfiorei Malzacher, 1986 is widespread in southern Italy, where it is sympatric with C. pseudorivulorum Keffermüller, 1960. Caenis belfiorei was originally described as a subspecies of C. pseudorivulorum but, according to
Across the different families of Ephemeroptera, we have a number of easily identified species whose presence in Italian watercourses seems to have decreased considerably in the last two decades. Although this indication results mainly from the direct activity of the authors not expressly aimed at assessing the range of these species and from occasional analyses of biomonitoring data, it unfortunately seems that the presence of the species is rarer than in the past, although some of them may locally reach high densities where present. Among these species, mention should be made of Ephoron virgo (Olivier, 1791), Oligoneuriella rhenana (Imhoff, 1852), Acentrella sinaica Bogoescu, 1931, Heptagenia coerulans Rostock, 1878 and Nigrobaetis digitatus (Bengtsson, 1912).
Then there are other, relatively rare species whose distribution in Italy has never been really known in detail and which would require specific surveys to confirm their presence and verify the conservation status of their habitats. These include Ephemera vulgata Linnaeus, 1758, Ephemera glaucops Pictet, 1843, Nigrobaetis niger (Linnaeus, 1761), Ecdyonurus zelleri (Eaton, 1885), Ecdyonurus ruffii Grandi, 1953, Rhithrogena nivata (Eaton, 1871), Caenis beskidensis Sowa, 1973, and Caenis lactea (Burmeister, 1839).
Finally, we have five species no longer found after the first collection, which, in four cases, supported the species description. A very interesting and peculiar species, both for its morphology and for the environments it colonises (
Although we have not directly analysed specimens of the two species, Ecdyonurus bellieri (Hagen, 1860) and Rhithrogena siciliana Braasch, 1989, which were no longer collected after their description, they are presented in the taxa list. In this case, although they are both species inquirenda, they were described on material collected in Sicily, a land rich in endemism, and we have, therefore, cautiously preferred to record them in the taxa list.
Caenis valentinae Grandi, 1951 and Procloeon calabrum (Belfiore and D’Antonio, 1990), Italian endemics, have not been caught since the collections that allowed their description. Dedicated sampling campaigns are planned in the near future in the hope of confirming the presence of these two species and thus their non-extinction.
The general picture of the Italian Ephemeroptera, briefly described above, allows us to highlight some relevant aspects to guide future research on the taxonomy of this order. A first element, at once of great stimulus for research and of concern, is that several species described on material collected in Italy have not been caught since the initial collection. In some cases, type material appears to have been lost or, where present, is not usable for genetic analysis. The collection of new material from type localities is therefore urgent. Some species are known from only one or a few close localities. In this case, new collections are indispensable, although there is no guarantee of finding the species again. When a species has been described from specimens collected from different areas in Italy, it will be easier to make new collections, but it will be advisable to check whether all specimens belong to a single species, possibly with the aid of genetic techniques, for the possible presence of cryptic species. Consequently, such verification should also be done for possible lectotypes or syntypes, whenever possible.
As mentioned above, there are currently insufficient data for the compilation of a Red List of Italian Ephemeroptera, although this is a matter of extreme urgency. At present, there is no repository for information on the collection of Ephemeroptera species and, in fact, data are archived, in a more or less systematic and organised manner, to manage the authors’ physical collections. At the national level, there are repositories of information at genus or family level. In general, apart from the activities of the authors, few attempts exist in Italy to arrive at a specific identification of the Ephemeroptera.
The focus on Ephemeroptera has traditionally been on running water environments, which are home to many more species than lentic environments. This has therefore led to a lack of knowledge about the species present in ponds and lakes. In any case, the greatest urgency for further knowledge seems to be related to lotic environments, by virtue of their tendency to become temporary (
The advent of new genetic techniques and standards for comparing different taxa and defining the boundary between different species can facilitate taxonomic investigation (
The primary reason for the limited focus on taxonomic studies of Ephemeroptera appears to be the lack of dedicated funding (reason 1). The scarcity of resources for basic research, combined with a broader shift towards more applied scientific topics (reason 2), has rendered taxonomic studies increasingly unattractive, uninteresting, or simply unsustainable for newer generations of researchers in Italy (see
Although accurate and reproducible taxonomic identification is the cornerstone of biology, the level of validation of taxonomic identification in entomological studies is often poor (
Comparative tables and the quantification of diagnostic characteristics for species identification and divergence—along with the species descriptions and reviews compiled by the authors—are likely outdated. These are often based on a limited selection of species present in Italy (and Europe), as recent biomolecular analyses increasingly indicate. This suggests that interspecific differences may have been conflated with intraspecific variability, or that inappropriate diagnostic characteristics were selected. For example, Ecdyonurus austriacus was not recorded in Italy when the review of Italian species in the helveticus group was compiled (
This paper aims to contribute meaningfully to the study of Italian Ephemeroptera, acknowledging both the highly dynamic nature of species concepts and the importance of incremental advances in scientific understanding. We argue that morphospecies (i.e., an approximate typological interpretation of Linnaean species) hold a crucial role in establishing ‘stable’ reference points (
In closing, we wish to highlight one of the apparent paradoxes of biodiversity. In a world where extinction rates are alarmingly high, the paradox lies in the fact that the number of recorded species may appear to increase simply because these species- which in reality are likely to have existed for millennia- are defined and described by the academic community (
This work would not have been possible without the contributions of all our colleagues who, over the years, have alerted us to the presence of interesting species in the area, sent us samples and assisted us in various stages of the study of Ephemeroptera. Without naming names- they would be too many- we would like to thank them all here. More specifically, for the present work, we would like to thank Marcello Cazzola (CNR-IRSA) for the production of Fig.
The authors have declared that no competing interests exist.
No ethical statement was reported.
NBFC - Project funded under the National Recovery and Resilience Plan (NRRP), Mission 4 Component 2 Investment 1.4 - Call for tender No. 3138 of 16 December 2021, rectified by Decree n.3175 of 18 December 2021 of Italian Ministry of University and Research funded by the European Union – NextGenerationEU; Project code CN_00000033, Concession Decree No. 1034 of 17 June 2022 adopted by the Italian Ministry of University and Research, CUP B83C22002930006, Project title “National Biodiversity Future Center - NBFC”.
ITINERIS - IR0000032 – ITINERIS, Italian Integrated Environmental Research Infrastructures System (D.D. n. 130/2022 - CUP B53C22002150006) Funded by EU - Next Generation EU PNRR- Mission 4 “Education and Research” - Component 2: “From research to business” - Investment 3.1: “Fund for the realisation of an integrated system of research and innovation infrastructures” CUP B53C220021500006).
Both authors contributed to the study conception and design. The first draft of the manuscript was written by Andrea Buffagni and both authors commented on previous versions of the manuscript. Both authors read and approved the final manuscript.
Andrea Buffagni https://orcid.org/0000-0002-3944-1861
Carlo Belfiore https://orcid.org/0000-0002-0061-8658
All of the data that support the findings of this study are available in the main text or Supplementary Information.
Map of the Italian Hydro-ecoregions
Data type: pdf