Morphological variability of Cordulegaster trinacriae in Italy 1st December 2019175

Morphological variability of Cordulegaster trinacriae in Italy (Odonata: Cordulegastridae)

Andrea Corso

MISC – Via Camastra, 10, 96100 Siracusa, Italy;

Received 15th July 2019; revised and accepted 10th September 2019

Abstract. This paper describes the most helpful features for field identification C.of trinacri­ ae and the relevant morphological variability encountered. In C. trinacriae the yellow frons was without a dark horizontal marking in about 70 % of the sample or showed a barely pat- terned frons, while the remaining 30 % showed a more or less defined dark smudge, though always narrower and less bold than in C. boltonii. The percentage of unmarkedversus marked frons was similar throughout the distributional range of C. trinacriae except in Sicily, where the unmarked yellow frons was much more common (87 %) and in Campania, where in- dividuals with a dark mark on the frons were slightly more common than unmarked ones. In C. trinacriae the occipital triangle was almost always yellow with no or almost no dark markings, cleaner, and purer yellow than in C. boltonii from central Italy. The appendages of C. trinacriae always showed the characteristic form except for a few individuals, which might be hybrids. In C. boltonii they were found to be rather variable, especially on individuals from the southern part of its distributional range in central Italy, where intermediates and hybrids occur. Here, the appendages were sometimes similar (but not identical) to C. trinacriae, and therefore, this character is considered of limited use in the field. Individuals of C. trinacriae can be distinguished in areas of sympatry from intermediate C. boltonii and from hybrids when they show the following characters: i) unmarked yellow frons; ii) bright yellow, almost unmarked occipital triangle; ii) typical deeply notched lower appendages and long, sinuous upper appendages. Further key words. Dragonfly, Anisoptera, frons pattern, occipital triangle, field identifica- tion

Introduction Cordulegaster trinacriae Waterston, 1976, is endemic to the southern half of Italy from Sicily to the regions of Lazio and the Abruzzi. The type local- ity is in Sicily, which is reflected by the scientific name; the Greek word trin­acria means “three pointed” and is the earliest known name of the island of Sicily, formerly a Greek possession, so named for its triangular

Odonatologica 48(3/4) 2019: 175-201Odonatologica – DOI:10.5281/zenodo.3539730 48(3/4) 2019: 175-201 176 A. Corso shape (Boudot et al. 2009; Riservato et al. 2014; Boudot & Kalkman 2015). The species is well differentiated, both morphologically (Boudot & Jacque­min 1995; Boudot 2001; Dijkstra & Lewington 2006; Dijkstra & Kalkman 2012; Solano et al. 2018) as well as genetically (Froufe et al. 2014; Solano et al. 2018). Its morphological variability was described, albeit only briefly, by Balestrazzi et al. (1982, 1983), Galletti & Pave- si (1985) and Ottolenghi (1991). On the other hand, Balestrazzi et al. (1983), Galletti & Pavesi (1985), and more recently Terzani (2015, 2016), discussed the morphological variability of Italian populations of Cordulegaster boltonii (Donovan, 1807), with the latter author stressing the need for larger study samples from Lazio and the Abruzzi and especially from the overlapping hybridization zone of the two species (Solano et al. 2018). Concerning other European populations of C. boltonii, the variabil- ity and the validity of the various proposed subspecies have been extensive- ly studied, sometimes erroneously with respect to current knowledge and systematics, by, e.g., St. Quentin (1952), Jurzitza (1965), Waterston (1976), Theischinger (1979), Ocharan (1989), Lohmann (1992), Bou- dot & Jacquemin (1995), Boudot (2001), and Froufe et al. (2014). The morphological variability of the appendages in male C. trinacriae, C. bolto­ nii and their hybrids was recently described by Solano et al. (2018), who did however not discuss any other field identification features. As C. tri­ nacriae is listed in the Habitats Directive (Annex II and IV) and the Bern Convention (Annex II and Revised Annex I), reliable identification in the field is a priority when planning conservation actions and for any monitor- ing projects and research. Given the variability of morphological characters in both species and their similarity, it is important to analyse the reliability of the different characters to assign individuals to a respective species and to identify the features most helpful for field identification of C. trinacriae. The objective of the present paper is to analyse the morphological traits that are most helpful to distinguish C. trinacriae from C. boltonii and their hybrids in the field. The following characters are considered: the colour pattern of the frons, the colour of the occipital triangle, the pattern of the abdomen, and the form of the appendages. For comparison, data on the variability of C. boltonii from Lazio and Tuscany − the populations near- est to populations of C. trinacriae − are provided, with a brief overview of

Odonatologica 48(3/4) 2019: 175-201 Morphological variability of Cordulegaster trinacriae in Italy 177 specimens from northern Italy as well. Also, considerations are given on the hybridization between the two species, which covers an extensive area (Solano et al. 2018).

Material and methods In order to study the morphological variability of Cordulegaster trinacriae, 355 specimens were studied: 317 males and 38 females (Fig. 1; Tables 1, 2). The specimens were assorted according to their regions of origin; 121 were from Sicily (102♂, 19♀), 88 from the Abruzzi (78♂, 10♀), 35 from Molise (33♂, 2♀), 45 (41♂, 4♀) from Campania, and 66 (63♂, 3♀) from Calabria (Fig. 1; Tables 1, 2). Specimens of pure or putative C. trinacriae from Lazio were not considered for the data-set analysis in order to avoid being misled by the many hybrids found in this area (Solano et al. 2018). All study sites were geo-referenced in decimal degrees WGS84. Date of sampling, region, provinces, locality name, and GPS coordinates are listed in Appendix 1. For comparison, a total of 186 specimens of C. boltonii were studied: 60 males and 19 females from Lazio, 45 males and 17 males from Tuscany, including Elba Island, and 47 males from Lombardy and Piedmont in northern Italy (Fig. 1). Additionally, a number of specimens from central-southern Italy (28♂ and 7♀ C. boltonii and 9♂ and 3♀ C. trinacriae) held in the entomo- logical collections of the following institutes were also compared: Museo Civico di Zoologia di Roma, Italy (MCZR); Istituto di Zoologia Università di Roma “La Sapienza”; Collezioni Dipartimento di Biologia e Biotecnolo­ gie Charles Darwin, Roma. As post-mortem changes could influence the ap- pearance of the frontal area and the occipital triangle (Balestrazzi et al. 1982), the museum specimens were only used to test the reliability of these characters during museum studies but are not included in the data-set. Most individuals were caught while patrolling along rivers and torrents with large entomological hand nets. Dragonflies were photographed in the field and immediately released. The study was conducted during specific monitor- ing projects for priority species included in the Habitats Directive commis- sioned to the author, who obtained permission from the administrations of the Aspromonte, Majella, and Nebrodi National Parks and several Nature Reserves. No specimens were collected. The individuals were only handled for photographing and were released immediately. Photographs were always

Odonatologica 48(3/4) 2019: 175-201 178 A. Corso taken from standard positions: lateral view, upper- and underside of the ab- domen, head and thorax from above, and head in frontal view. They were taken with a Canon EOS with an 18–50 mm zoom macro lens. The study fo- cused on the morphological variation of the males; in fact, female Cor­dule­ gaster are more difficult to find than males, and females C. trinacriaeof and C. boltonii are so similar (Consiglio 1958; Balestrazzi et al. 1982) that reliable identification is not always possible. Therefore, only a small sam- ple of females was analysed and the variability of female characters is only briefly discussed.

Regarding the frontal area, a code was assigned to each of the four different patterns encountered: FA − complete absence of dark transversal marking, frons completely yel- low (rarely a tiny darkish spot in the middle); FB − a dark smudge, mostly brownish, across the frons; FC − a well-marked and visible dark transversal bar, though narrow; usu- ally black/blackish; FD − well-defined and bold dark bar, rather long and wide; black (Figs 3−5, 7; Table 2). For the occipital triangle, the following codes were used: OA − completely yellow occipital triangle, only with well-defined and nar- row dark lateral margins; OB − completely yellow occipital triangle with a darkish tinge at the top and often wider dark margins and base; OC − a mostly or partially yellow/yellowish triangle, with a dark line in the middle all the way from the base to the top, usually wide dark lateral margins; OD − fully dark or almost completely dark occipital triangle, sometimes with two tiny lateral yellow spots more or less visible – often only by using a magnifying lens (Figs 3−5, 7; Tables 1, 2). The same codes were used and assigned to the studied C. boltonii.

The form of the appendages, which is considered highly species-specific, is only briefly discussed here as it was already the target of recent research (Solano et al. 2018). They were photographed from below and above and

Odonatologica 48(3/4) 2019: 175-201 Morphological variability of Cordulegaster trinacriae in Italy 179

Figure 1. Map of Italy showing the study sites. Red dots represent records of Cor­ dulegaster trinacriae and green dots records of Cordulegaster boltonii. Map by courtesy of Michele Viganò.

Odonatologica 48(3/4) 2019: 175-201 180 A. Corso their form was examined using a 10× and 20× Leica hand-held lens. The pattern (shape, boldness, number) of the yellow rings along the abdomen was also checked dorsally and ventrally.

Figure 2. Map of Italy showing regions were specimens of Cordulegaster trinacriae were studied. Composition of frons pattern type (F) and occipital triangle pattern type (O) is shown (types A, B, C, D; see text and Tables 2, 3). Map by courtesy of Michele Viganò.

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Table 1. Number of male specimens of Cordulegaster trinacria studied per region according to the different types (FA, FB, FC, FD) of frons pattern, where FA indi- cates an unmarked yellow frons, and FC to FD an increasingly darker and more distinguishable (wider/longer) dark bar across it (see text).

Region n ♂ FA FB FC FD Total Total Total FC FA FB + FC FA + FB Sicilia 102 89 11 2 0 87.2 % 12.8 % Calabria 63 38 12 13 0 (1) 60.3 % 39.7 % Campania 41 20 10 11 0 (5) 48.8 % 51.2 % Molise 33 21 10 2 0 63.6 % 36.4 % Abruzzo 78 51 10 17 0 65.4 % 34.6 % Total 317 219 53 45 0 (6) Frequency [%] 69.1 16.7 14.2 0 (1.9) 69.1 30.9 85.8 14.2

Table 2. Number of male specimens of Cordulegaster trinacria studied per Italian region according to the different types (OA, OB, OC, OD) of occipital triangle pat- tern (see text), where OA indicates an unmarked yellow occipital, and OB to OD an increasingly darker occipital.

Region n ♂ OA OB OC OD Sicilia 102 96 6 0 0 Calabria 63 58 4 1 1 Campania 41 30 5 4 2 Molise 33 31 2 0 0 Abruzzo 78 65 9 4 0 Total 317 280 26 9 3 Frequency [%] 88.3 8.2 2.8 0.7

Results Frons pattern Of the 317 Cordulegaster trinacriae males studied, 219 (69.1 %) had a com- pletely yellow frons with no frontal bar (type FA) (Fig. 3), and 53 males (16.7 %) showed a more or less marked brownish smudge (type FB) (Fig. 4;

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Table 1). The remaining 45 males (14.2 %) showed a quite well marked black/blackish bar (type FC) (Fig. 5), always narrower, usually shorter and less conspicuous than in typical C. boltonii (type FD) from central Italy (Fig. 6). Since some FB-type individuals with a brownish marking over the frons were very similar to FC-types, and the two types were not always truly separable, we clustered them together and obtained 219 male speci- mens lacking the dark frontal bar (type FA: 69.1 %) and 98 (types FB + FC: 30.9 %) showing a more or less marked pattern (Table 2). Under field condi- tions, since many FB-types could be considered as showing a barely marked dark bar and difficult to tell apart from FA-types, we could actually cluster types FA and FB together and obtain a total of 85.8 % with a more or less unmarked or hardly marked yellow frons compared to a set of 14.2 % with a well-marked frons (Table 1). Only five males from northern Campania and one from southern Calabria (Reggio Calabria, Aspromonte National Park) showed a well-defined FD-type pattern (Fig. 7). These individuals had a more striking dark frontal bar, comparable to that of average C. bol­ tonii. However, it appeared less thick and conspicuous. Such individuals were therefore designated as extreme examples of type FC. The patterns obtained in the various regions for male C. trinacriae are given in Figure 2 and Table 1. As for C. boltonii, 89 males (84.8 %) of the studied sample from central Italy showed a rather solid and well-marked dark frontal transversal bar (type FD), rather variable in length but not in width, and always better de- fined and more striking than in C. trinacriae (Fig. 6). A total of 12 males from southern and central Lazio and two from Tuscany (13.3 %) showed a shorter and less marked dark bar, comparable to type FC of C. trinacriae. Fi- nally, two males from Lazio (1.9 %) showed a type FB frons pattern (Fig. 8), less marked than some strongly marked C. trinacriae (Fig. 7). The control sample from northern Italy showed a more variable dark bar over the frons, with 18 males (38.3 %) displaying a FB-type bar as in C. trinacriae. Concerning females of C. trinacriae, individuals from the entire distribu- tion range had a dark frontal bar, always longer and wider than in males, similarly black and evident as in C. boltonii with only a few individuals showing a less marked and defined bar, sometimes shorter and narrower than usual (Fig. 9). On average, the dark markings appeared slightly nar-

Odonatologica 48(3/4) 2019: 175-201 Morphological variability of Cordulegaster trinacriae in Italy 183 rower and less conspicuous than in average female C. boltonii from central Italy (Fig. 10). However, these differences are barely visible in the field and are best discerned in series of specimens.

Figure 3. Cordulegaster trinacriae. Figure 4. Cordulegaster trinacriae. Male from Messina, Sicily. Typical un- Male from Chieti, the Abruzzi. Type FB, marked yellow frons pattern type FA OA. Photo: AC (vii--2017) and fully yellow occipital triangle type OA. Photo: AC (vii-2017)

Figure 5. Cordulegaster trinacriae. Male Figure 6. Cordulegaster boltonii. Male from Fiume Lente, Benevento, Campa- from Rome, Lazio. Frons pattern type nia. Type FC, OB. Photo: AC (vii-2017) FD. Occipital triangle type OD, almost completely dark, typical of central Ital- ian populations. Photo: AC (vi-2017)

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Occipital triangle The colour patterns of the occipital triangle from the individuals collected in the various regions are presented in Table 2 and Figure 2. In C. trinacriae, the occipital triangle was usually pure or almost pure yellow (type OA), more or less bright in colour, and uniformly yellow in front as much as behind (Figs 3, 4, 11). Only a small proportion showed darker markings (type OB and OC: 35♂, 11 %), whereby one male from Calabria and two from Cam- pania had almost completely dark occipital triangles and were classed as OD, approaching the colour that is typically found in C. boltonii from central Italy (Fig. 7; Table 2). Types OB and OC of C. trinacriae showed wider dark “frames” and top compared to type OA (Fig. 5). However, in the hand, the colour of the triangle appeared cleaner and purer yellow than in most C. bol­ tonii studied from central Italy, including C. boltonii from the overlapping areas. In fact, 40 male C. boltonii (66.7 %) from Lazio showed an almost com- pletely dark-looking occipital triangle of the OD type (Fig. 6), often with two minute yellow spots. Nine males were type OC, and only five (8.4 %) showed a (dull) yellow occipital triangle of type OB that was never as bright as in C. trinacriae. In Tuscany, 28 males of the samples were type OD (64.5 %),

Figure 7. Cordulegaster trinacriae. Male Figure 8. Cordulegaster boltonii. Male from Reggio Calabria, Calabria. The from Roma, Lazio. A very atypical most distinctly marked frons bar, type C. boltonii for the central Italian popu- FC/FD, and the darkest occipital trian- lation with frons type FB and occipital gle (OC/OD) recorded in C. trinacriae. type OA/OB. Very similar to C. trin­ Collected far from the documented hy- acriae, and could be a hybrid. Photo: AC bridization area. Photo: AC (vii-2018) (vii-2018)

Odonatologica 48(3/4) 2019: 175-201 Morphological variability of Cordulegaster trinacriae in Italy 185 eight were OC (17.8 %), and seven were type OB (15.6 %). Altogether, only eight males (7.6 %) of the whole central Italian group matched type OA, al- though their occipital triangle appeared duller than in typical C. trinacriae. Of these eight, six males were from central-southern Lazio (10 %) (probably hybrids) and two (4.4 %) from the island of Elba. The rear part of the occipi- tal triangle was always brighter and cleaner yellow and less patterned than the frontal part, giving a bicoloured appearance, while in all C. trinacriae the occipital triangle was more uniformly yellow with similar colour in the frontal and rear parts (Figs 11, 12). Concerning the C. boltonii sample from northern Italy taken for comparison, only 11 males (24.5 %) showed a very dark type OD occipital triangle, identical to that found in most central Ital- ian specimens, with the rest showing type OB/OC or even OA as in C. tri­ nacriae. No specimens from Germany, England or Sweden showed the dark occipital triangle found in central Italy, always having instead a rather clean bright yellow occiput. Concerning females, C. trinacriae usually has a clean- er and brighter yellow occipital triangle that is more uniform in appearance (Fig. 9) than female C. boltonii from central Italy. Most females of C. boltonii show a duller, more patterned and more bicoloured looking triangle (cleaner

Figure 9. Cordulegaster trinacriae. Fe- Figure 10. Cordulegaster boltonii. Fe- male from Reggio Calabria, Calabria. male from the island of Elba, Tuscany. Females always have a well-marked and Rather long and thick dark marking over thick dark bar on frons. Note the yellow the frons. Note the much darker occipi- occipital triangle. Photo: AC (vii-2018) tal triangle compared to the female of C. trinacriae (Fig. 9). Photo: Leonardo Forbicioni (vii-2016)

Odonatologica 48(3/4) 2019: 175-201 186 A. Corso and brighter yellow on the rear side) than in C. trinacriae (Fig. 10), though the pattern and colour was sometimes overlapping. Usually, the pale area on the rear part of the eyes was smaller in C. trinacriae compared to the average C. boltonii, especially those from northern Italy.

Appendages Almost all male C. trinacriae had very well notched lower appendages that sometimes appeared as two separated lobes (Fig. 13). Only a few individuals showed a slightly less clearly defined notch, with the lower appendages also appearing rather small and not as wide as usual (2.7 % of the studied sam- ples). These were all from northern Campania (Fig. 1), possibly falling into the hybridization area recently identified to the south of Rome Solano( et al. 2018). The typical lower appendages in C. trinacriae appeared wider and larger than in C. boltonii with straighter, more parallel sides. The upper ap-

Figure 11. Cordulegaster trinacriae. Figure 12. Cordulegaster boltonii. Male Male from Sortino, Siracusa, Sicily. The from Roma, Lazio. Note the strongly bi- occipital triangle looks homogenously coloured occipital triangle. Photo: AC yellow. Photo: AC (vii-2017) (vi-2018)

Odonatologica 48(3/4) 2019: 175-201 Morphological variability of Cordulegaster trinacriae in Italy 187 pendages of C. trinacriae were always longer and more sinuous than in typi- cal C. boltonii and were often also narrower. In some intermediate specimens of C. boltonii, their form was similar to those typically seen in C. trinacriae. Concerning C. boltonii, the form of the lower and upper appendages dis- played high variability. The majority (80♂, 76.2 %) had thicker, shorter, and less sinuous upper appendages with slightly or barely notched lower append- ages (Fig. 14). A total of ten male specimens ascribed to C. boltonii from central-southern Lazio (16.7 % of the sample from this region, and 9.5 % of the total) had a well-notched hind margin (Fig. 15) very similar to that of C. trinacriae, though they were smaller and narrower, with less parallel sides than in the latter. The remaining specimens appeared intermediate (15♂,

Figure 13. Cordulegaster trinacriae. Figure 14. Cordulegaster boltonii. Male from Cassaro, Siracusa, Sicily. Typ- Male from the island of Elba, Tuscany. ical lower appendages, deeply notched Typical lower appendages, very slightly and wide. Photo: AC (vi-2012) notched, narrow and with sides that are not parallel as in C. trinacriae. Pho- to: Leonardo Forbicioni (vii-2016)

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14.3 %). Several specimens of C. boltonii (almost 40 %) showed longer, nar- rower and less robust upper appendages than are illustrated in most works, similar to the typical form found in C. trinacriae. Most of these intermedi- ates were from central-southern Lazio (87 %) and could be hybrids.

Yellow abdominal pattern Differences in the abdominal pattern in C. trinacriae and C. boltonii, dor- sally, ventrally, and laterally, are often reported as helpful for identifica- tion. Usually, C. trinacriae has more restricted yellow rings than the aver- age C. boltonii, most obviously on S6–S8 (S9), with S6–S7 showing a single yellow ring (the wider basal one). Normally, C. boltonii shows two rings in these segments (a wider basal and narrower apical one in the shape of two adjacent dots − lunulae) and wider yellow rings on S6–S8 (S9). Me- dian abdominal segments have yellow markings narrowing dorso-ventrally in C. trin­acriae, while they are of same width dorso-ventrally in C. boltonii, or only slightly narrowing. During this study, a large variability of these pat-

Figure 15. Cordulegaster boltonii. Male from Roma, Lazio. Same specimen as in Figure 8. Note that lower appendages are more notched than in typical C. bol­ tonii, and also appear to be wider with more paral- lel sides. Thus, a hybrid with C. trin­acriae cannot be excluded. Photo: AC (vii--2018)

Figure 16. Cordulegaster trinacriae. Male from Caserta, Campania. Note the typi- cally restricted yellow single rings on S6-S9 with no apical lunulae and the very long and sinuous upper appendages. Photo: AC (vii-2017)

Odonatologica 48(3/4) 2019: 175-201 Morphological variability of Cordulegaster trinacriae in Italy 189 terns was observed in C. boltonii and much less in C. trinacriae. In particu- lar, a similar pattern to that of some C. trinacriae was documented in central Italian populations of C. boltonii. Many specimens of C. boltonii from this area had much-restricted yellow rings, often missing the apical lunulae on S7 and sometimes also that on S6 (S5), therefore showing a pattern match- ing or being very similar to that of C. trinacriae (Fig. 18). Some specimens from northern Italy were also missing the apical lunulae on S6, although more rarely than in the population from central Italy (Fig. 19). The pattern of S6 was found to be variable in C. trinacriae, with most specimens missing any trace of apical lunulae (Fig. 16). However, a significant percentage (35 %) of specimens studied showed more or less well visible and marked lunulae (Figs 17, 18). Ventrally and laterally, S7 usually had wider and straighter markings in C. boltonii; these markings on S7 appeared narrower and more C-shaped in C. trinacriae. However, several specimens of C. boltonii showed a pattern comparable to that of C. trinacriae and vice versa.

Figure 17. Cordulegaster trinacriae. Male from Benevento, Campania, with vague- ly marked pale apical lunulae on S6 and less sinuous upper appendages although still very long and narrow. Photo: AC (vii-2017)

Figure 18. Cordulegaster trinacriae. Male from Caserta, Campania, showing better developed lunulae on S6. Photo: AC (vi-2012)

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Discussion The data collected here on the morphological variability of Cordulegaster trin­acriae and C. boltonii reveal a complex situation. Previously, the mor- phological variability of C. trinacriae had been discussed only by Bale­ strazzi et al. (1982, 1983) and Galletti & Pavesi (1985), based on a limit- ed number of dead specimens from Sicily and Calabria. Solano et al. (2018) discussed the genetic variability found within C. trinacriae and C. boltonii in Italy, confirming the validity of the two species as already advocated by Froufe et al. (2014). Solano et al. (2018) found hybrids as far north as the Marches region, but these were apparently morphologically identical to C. boltonii, while all other hybrids appeared intermediate but much more similar to C. boltonii and never resembling C. trincariae. However, the study

Figure 19. Cordulegaster boltonii. Male from the island of Elba, Tuscany. Narrow yel- low apical lunulae on S6 but missing on S7 as in some C. trin­acriae. Note however the shorter and thicker upper appendages. Photo: Leonardo Forbicioni (vi-2016)

Figure 20. Cordulegaster boltonii. Male from Varese, Lombardia, with narrow and small apical lunulae also visible in this specimen from northern Italy. Note the rather long and sinuous upper appendages of this specimen. Photo: Michele Vi- ganò (vii-2018)

Odonatologica 48(3/4) 2019: 175-201 Morphological variability of Cordulegaster trinacriae in Italy 191 by Solano et al. (2018) only analysed the form of male appendages and did not consider colour patterns of the head or the abdomen. However, knowledge on the variability of the colour patterns of the two species is useful in the field for species identification, particularly because in the past, many specimens were misidentified and the problem of hybrids was not yet known. Furthermore, reliable field identification is essential. Considering that C. trinacriae is a protected species listed in the Habitats Directive, its study and monitoring should preferably be conducted on liv- ing individuals only. The variability of the main phenotypical identification characters is sum- marised below.

Frons pattern In the present study, with a larger sample than ever before, most male speci- mens (>70 %) of C. trinacriae showed a clean yellow frons, and only about 14 % of the studied specimens had a well-marked bar. In fact, historically, Fraser (1929) erroneously ascribed the Sicilian Cordulegaster to C. annu­ latus ssp. immaculifrons. The frons bar observed in C. trinacriae was never as well-defined, thick, and conspicuous as in typical C. boltonii from the central Italian populations. Only six male C. trinacriae from Campania and Calabria had a well-defined dark bar (Fig. 2; Table 2). In this species, the presence of a dark bar, almost absent in Sicily, became more common from south to north-west. Interestingly, the highest percentage of a more conspic- uous dark marking was found in Campania (51.2 %), particularly in north- ern Campania. Here, several Cordulegaster specimens appeared phenotypi- cally odd, bearing more resemblance to C. trinacriae. When comparing typical C. trinacriae with the ‘immaculifrons’ form of C. boltonii, formerly considered a valid subspecies but recently not accepted even if based only on genetic data (Froufe et al. 2014), the face pattern, i.e., the frons marking and occipital triangle, is very similar and thus useless as an identification feature.Ocharan (1989) as well as Boudot & Jacquemin (1995) report extensive variability in all Iberian and French populations of C. boltonii with specimens displaying a range from completely unmarked to very well marked yellow frontes, a much greater variability than in cen- tral Italian C. boltonii. Balestrazzi et al. (1983) previously reported simi-

Odonatologica 48(3/4) 2019: 175-201 192 A. Corso lar variability in a population from Liguria in north-western Italy that they identified as immaculifrons» «. However, C. boltonii of the ‘immaculifrons’ type does not occur in central-southern Italy and Sicily, and there is no risk of confusion with C. trinacriae.

Occipital triangle The occipital triangle appears entirely bright yellow in C. trinacriae on the front and rear side, and its colour is only barely variable as already reported by Balestrazzi et al. (1982, 1983) and Galletti & Pavesi (1985). In the populations of C. boltonii from central Italy, the occipital triangle − although of variable colour, as already shown by Terzani (2016) − is usually darker and more patterned than in both C. trinacriae and C. boltonii from other northern Italian and other European areas and usually appears bicoloured with the rear part paler than the front side, allowing easy identification. The average dark occipital triangle in C. boltonii agrees with considerations giv- en by Boudot (2001), who regarded the central Italian C. boltonii as »atypi- cal«. Froufe et al. (2014) found that specimens from Tuscany were geneti- cally distinct and form a separate cluster from other European C. boltonii. Therefore, some phenotypic differences in morphology and colour patterns are not surprising.

Appendages In C. trinacriae the appendages appear rather uniform in shape except for very few specimens from the north-western part of its range. However, only a very limited number of specimens from northern Campania was studied, and no specimens from southern Lazio were analysed (Table 1; Fig. 1). In contrast, the shape of the appendage of C. boltonii appeared rather variable with many specimens having long and narrow upper appendages and several showing also a more notched hind margin of the lower appendages than in any illustration found in the literature (e.g., Waterston 1976; Balestraz- zi et al. 1983; Galletti & Pavesi 1985; d’Aguilar et al. 1985; Boudot & Jacquemin 1995; Boudot 2001; Askew 2004; Dijkstra & Lewington 2006; Lehmann & Nüss 2015). More recently, Terzani (2016) illustrated for Italian C. boltonii certain variability in the shape of the appendages and Solano et al. (2018) found that upper appendages and lower appendages

Odonatologica 48(3/4) 2019: 175-201 Morphological variability of Cordulegaster trinacriae in Italy 193 generally permitted species identification, but in either case, a number of specimens were intermediate. In particular, hybrids were morphologically always similar to C. boltonii. Thus the shape of the appendages needs to be applied with caution in the field and is best to be used in museum specimens when direct comparison between the two species and the various specimens are possible. However, a specimen with appendages typical for C. trinacriae allows for a reliable identification, also considering hybrids (see below).

Abdomen pattern Like the shape of the upper appendages, the colour pattern of the abdominal segments, widely reported in literature to be diagnostic (e.g., Balestrazzi et al. 1982, 1983; Galletti & Pavesi 1985; Boudot 2001; Dijkstra & Lew- ington 2006), was also found to be unreliable because of the variability ob- served in C. boltonii. In fact, in many C. boltonii from central Italy, as well as in some from northern Italy, the yellow rings were narrower than the aver- age, and most individuals lacked the usual apical lunulae on S7, which were also sometimes missing on S6 and occasionally even on S5 (Terzani 2016; AC unpubl.). This is the same colour pattern typically found in C. trinacri­ ae. Ocharan (1989) and Boudot & Jacquemin (1995) also found similar exceptions in Spanish and French populations of C. boltonii. In contrast, a significant percentage ofC. trinacriae was found with apical pale lunulae on S6, a feature not previously reported in the literature and considered char- acteristic of C. boltonii. Of course, when the median yellow rings are broad and extensive, with S6–S7 showing well marked and broad lunulae, this is reliably diagnostic for C. boltonii, both during field studies or in good qual- ity photographs.

Females In central Italy, females of the two species appear extremely similar, and species identification is often not possible. On average, female C. trinacriae show a narrower dark bar on the frons and a cleaner, more uniform, and brighter yellow occipital triangle, which is usually not bicoloured as in most C. boltonii females, where it is darker in front and yellower behind. In most individuals of C. trinacriae, a less extensive pale area is found in the rear part of the eye. These differences are not valid when comparing C. boltonii to

Odonatologica 48(3/4) 2019: 175-201 194 A. Corso other European populations. For example, Ocharan (1989) found within females of the Iberian population of nominate C. boltonii that 77 % of the specimens had a well-marked dark bar and 23 % had the bar reduced and less conspicuous, while it was reduced or almost absent in ‘immaculifrons’- type females.

Hybrids and intermediates Solano et al. (2018) found an extended hybrid area between C. trinacriae and C. boltonii in southern Lazio with its centre south of Rome. The inves- tigated morphological characteristics need to be considered in this context. For example, the geographical distribution pattern of the frons marking could be related to a higher number of hybrids and a general genetic intro- gression in central-western Italy. Considering the form of the male append- ages, Solano et al. (2018) report that »the hybrid individuals fall exclusive- ly within the range of variability of C. boltonii«. During the present study, some of the specimens genetically identified as hybrids were examined, all of which look very much like C. boltonii but with a poorly marked frons (FC or even FB) and intermediate occipital triangle or one similar to that of C. trinacriae. In contrast, most of the well-marked type FC of C. trinacriae (including the most FD types) found during my study displayed occipital triangles and appendages that are rather typical of C. trinacriae. Only very few specimens also possessed slightly atypical appendages (2.7 %). In future investigations, specimens identified genetically as hybrids should be exam- ined for their colour patterns of the frons and occipital triangle in addition to the form of the appendages. The question remains whether the specimens of C. trinacriae with a well- marked frons (types FC/FD) are hybrids or not. They might simply repre- sent the extremes of the variability of this species, which is documented here for the first time. The presence of a rather well-marked dark bar, some type FC and one almost type FD, found as far south as Calabria, strongly favours this hypothesis. Indeed, Solano et al. (2018) did not find mixed genes in the samples analysed from Molise, Campania, and Calabria, nor were any real intermediate specimens found in these areas. Samples from the Abruzzi were missing completely, as C. trinacriae was only recently found in this re- gion; future research in this region should also include genetic study.

Odonatologica 48(3/4) 2019: 175-201 Morphological variability of Cordulegaster trinacriae in Italy 195

Conclusions In conclusion, Cordulegaster trinacriae from the southern part of its range and in Sicily do not pose any identification problems. Some problems arise when dealing with museum specimens of C. boltonii from northern Italy, northern Europe, and of the so-called ‘immaculifrons’ type populations, though the form of the appendages, and to a lesser extent the pattern of the abdomen, will always permit correct identification. However, what is most important is the distinction between C. trinacriae and C. boltonii in their contact zone in central Italy. In fact, when planning conservation actions and monitoring projects focusing on C. trinacriae in central Italy, it is essen- tial to be certain of the species’ identification. I suggest to always use a com- bination of characters (Table 4). Cordulegaster trinacriae should be identi- fied by: i) observing first the frons and occipital triangle – the frons should be unmarked yellow and the occipital triangle wholly bright yellow (except for a dark frame); ii) then examining carefully the form of the appendages – the lower appendages must be deeply and evidently notched, rather wide, with parallel sides. The upper appendages are typically long, sinuous and divergent; iii) checking the yellow markings on the median abdominal seg- ments – these should be narrow and straight, both from above and from the side, missing apical pale lunulae on S7 and showing no lunulae or very nar- row ones on S6. If all these characters agree with the typical form of C. trin­ acriae as here described, the specimen can be identified with certainty. Any individual with a more marked frons and darker occipital triangle must be extremely carefully checked for the form of the appendage and left un-iden- tified if these appear odd. So far, all hybrids identified genetically appeared phenotypically more similar to C. boltonii than to C. trinacriae. Acknowledgements For all their kind assistance and help in various ways, I warmly thank my fol- lowing colleagues and friends: Antonio Antonucci, Alessandro Biscaccian- ti, Jean-Pierre Boudot, Marco Carafa, Roberto Casalini, Enzo Colonnelli, Lorenzo De Lisio, Alex Festi, Raimondo Finati, Leonardo Forbicioni, Carlo Fracasso, Ottavio Janni, Sönke Hardersen, Otakar Holuša, Federico Landi, Calogero Muscarella, Maurizio Pavesi, Verena Penna, Lorenzo Starnini, Tiziano Terzani, Carlo Utzeri and Michele Viganò, as well as the whole team

Odonatologica 48(3/4) 2019: 175-201 196 A. Corso of Odonata.it. Leica Optics Ltd, especially Nanette Roland and Francesco Corrà, is thanked for the provided optical instruments. The Majella National Park Authority (Abruzzo) − in particular Dr Pino Marcantonio, Dr Antonio Antonucci and Dr Marco Carafa − and the Aspromonte National Park Au- thority (Reggio Calabria) − in particular Dr Sergio Tralongo and Dr Antoni­ no Siclari − are sincerely thanked for funding the researches in their respec- tive parks as part of studies to monitor and protect odonate species in the Habitats Directive. In the same way, the whole staff of the Nebrodi National park (Messina), Madonie National Park (Palermo) and Valle dell’Anapo Na- ture Reserve are thanked. But foremost, I wish to thank my beloved parents, Filippo Corso and Vittoria Filice; without their continuous support since I was a child, and still now, none of my work could have been possible.

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Odonatologica 48(3/4) 2019: 175-201 Morphological variability of Cordulegaster trinacriae in Italy 197 nata): a review. Organisms, Diversity & Riservato E., Festi A., Fabbri R., Grieco C., Evolution 12: 209-227 Hardersen S., Laporta G., Landi F., Siesa M.E. & Utzeri C. 2014. Odonata. Atlante Dijkstra K.-D.B. & Lewington R. 2006. Field delle libellule italiane – preliminare. Socie- guide to the dragonflies of Britain and Eu- tà Italiana per lo Studio e la Conservazione rope. British Wildlife Publishing, Gillingham delle Libellule. Edizioni Belvedere, Latina Fraser F.C. 1929. A revision of the Fissila- St. Quentin D. 1952. Der Rassenkreis der bioidea (Cordulegasteridae, Petaliidae and Cordulegaster boltoni (Donovan) (Odo- Petaluridae) (Order Odonata). Part 1. Cor- nata). Entomologisches Nachrichtenblatt dulegasteridae. Memoirs of the Indian Mu­ Öster­reichischer und Schweizer Entomolo­ seum 9: 69-167 gen 4: 73-75 Froufe E., Ferreira S., Boudot J.-P., Alves Solano E., Hardersen S., Audisio P., Amo- P.C. & Harris D.J. 2014. Molecular phylo­ rosi V., Senczuk G. & Anonini G. 2018. geny of the Western Palaearctic Cordule­ Asymmetric hybridization in Cordulegaster gaster taxa (Odonata: Anisoptera: Cordule- (Odonata: Cordulegastridae): secondary gastridae). Biological Journal of the Linnean postglacial contact and the possible role of Society 111: 49-57 mechanical constraints. Ecology and Evolu­ Galletti P.A. & Pavesi M. 1985. Ulteriori tion 15: 1-15 considerazioni sui Cordulegaster italiani Theischinger G. 1979. Cordulegaster heros (Odonata Cordulegasteridae). Giornale sp. nov. und Cordulegaster heros pelionen­ italia­no di Entomologia 2: 307-326 sis ssp. nov., zwei neue Taxa des Cordule­ Jurzitza G. 1965. Gedanken zu einigen Pro- gaster boltoni (Donovan) – Komplexes aus blemen des „Rassenkreises Cordulegaster Europa (Anisoptera: Cordulegasteridae). boltonii (Donovan)“ (Odonata, Anisoptera). Odonatologica 8: 23-38 Nachrichtenblatt der bayerischen Entomo­ Terzani F. 2015. Ricerche odonatologiche in logen 14: 4-8 Toscana. XIII. Il Cordulegaster boltonii (Do- Lehmann A.W. & Nüss J.H. 2015. Libellen. novan, 1807): nuovi dati (Odonata: Cordu- Deutscher Jugendbund für Naturbeobach- legastridae). Onychium 11: 67-71 tung, Göttingen Terzani F. 2016. Note su Cordulegaster bol­ Lohmann H. 1992. Revision der Cordule- tonii (Donovan, 1807) in Italia centroset- gastridae. 1. Entwurf einer neuen Klassifi- tentrionale (Odonata: Cordulegastridae). zierung der Familie (Odonata: Anisoptera). Onychium 12: 23-39 Opuscula zoologica fluminensia 96: 1-18 Waterston A.R. 1976. On the Genus Cor­ Ocharan F.J. 1989. Variabilidad en pobla- dulegaster Leach, 1815 (Odonata) with ciones ibericas de Cordulegaster boltoni special reference to the Sicilian species. (Donovan, 1807) (Odonata: Cordulegastri- Transactions of the Royal Society of Edin­ dea). Revista de Biología de la Universidad burgh 69: 457-466 de Oviedo 7: 109-121 Ottolenghi C. 1991. Osservazioni tasso- nomiche su alcuni odonati italiani. Notulae odonatologicae 3 (7): 97-112

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Appendix 1. List of recording sites of Cordulegaster trinacriae analysed during this study. Sampling date, region, province, municipality and/or locality name are given. GPS coordinates are in decimal degrees WGS84.

Date Region Province Locality Long E Lat N 08-vi-2009 Sicilia Messina Cesarò 14.720 37.948 12-vi-2009 Sicilia Siracusa Sortino – Valle 15.045 37.141 dell’Anapo 14-vi-2009 Sicilia Ragusa Ragusa 14.764 36.950 14-vi-2009 Sicilia Ragusa Casa Nobile 14.921 36.930 17-vi-2009 Sicilia Siracusa Ferla-Pantalica 15.031 37.140 26-vi-2010 Campania Caserta Prata Sannita 14.195 41.420 26-vi-2010 Campania Caserta Prata Sannita 14.195 41.420 08-vii-2010 Sicilia Siracusa Cassaro 14.957 37.102 11-vii-2010 Sicilia Siracusa Cassaro 14.957 37.102 17-vii-2010 Sicilia Siracusa Sortino 15.030 37.140 20-vii-2010 Campania Salerno Morigerati 15.550 40.145 20-vii-2010 Sicilia Siracusa Sortino 15.030 37.140 24-vii-2010 Sicilia Messina Cesarò 14.681 37.926 24-vii-2010 Sicilia Messina Cesarò 14.681 37.926 24-vii-2010 Sicilia Messina Cesarò 14.692 37.953 25-vii-2010 Sicilia Messina Roccella Valdemone 15.008 37.948 20-vii- 2010 Sicilia Messina Roccella Valdemone 15.008 37.948 16-viii-2010 Sicilia Siracusa Sortino 15.030 37.140 05-ix-2010 Sicilia Siracusa Sortino 15.030 37.140 14-ix-2010 Sicilia Siracusa Sortino 15.030 37.140 23-v-2011 Sicilia Siracusa Sortino 15.047 37.143 22-vi-2011 Molise Isernia Pescolanciano 14.343 41.686 22-vi-2011 Molise Isernia Carovilli 14.299 41.711 24-vi-2011 Basilicata Potenza Montemurro 16.023 40.274 25-vi-2011 Molise Campobasso Civitacampomarano 14.697 41.778 26-vi-2011 Molise Campobasso Campobasso 14.664 41.556 01-vii-2011 Sicilia Siracusa Ferla 14.956 37.101 27-vii-2011 Sicilia Siracusa Sortino 15.047 37.143 29-vii-2011 Sicilia Siracusa Sortino 15.030 37.140 30-vii-2011 Sicilia Siracusa Rosolini 14.886 36.845 03-viii-2011 Sicilia Siracusa Sortino 15.030 37.140

Odonatologica 48(3/4) 2019: 175-201 Morphological variability of Cordulegaster trinacriae in Italy 199

Date Region Province Locality Long E Lat N 10-viii-2011 Sicilia Messina Cesarò 14.650 37.945 10-viii-2011 Sicilia Messina Cesarò 14.645 37.954 10-viii-2011 Sicilia Messina Galati Mamertino 14.806 37.997 10-viii-2011 Sicilia Messina Cesarò 14.648 37.949 11-viii-2011 Sicilia Messina Novara di Sicilia 15.114 38.013 11-viii-2011 Sicilia Messina Roccella Valdemone 15.008 37.948 11-viii-2011 Sicilia Messina Roccella Valdemone 15.008 37.948 20-viii-2011 Sicilia Catania Licodia Eubea 14.703 37.152 20-viii-2011 Sicilia Ragusa Chiaramonte Gulfi 14.780 37.018 21-viii-2011 Sicilia Ragusa Ragusa 14.751 36.975 21-viii-2011 Sicilia Ragusa Ragusa 14.766 36.954 21-viii-2011 Sicilia Ragusa Modica 14.728 36.832 29-viii-2011 Sicilia Ragusa/Siracusa Rosolini 14.728 36.832 10-ix-2011 Sicilia Siracusa Sortino 15.030 37.140 20-ix-2011 Sicilia Siracusa Cassaro 14.728 36.832 20-ix-2011 Sicilia Siracusa Ferla 14.728 36.832 24-iv-2012 Sicilia Siracusa Sortino 15.040 37.137 13-vi-2012 Sicilia Siracusa Sortino 15.030 37.140 19-vi-2012 Sicilia Siracusa Noto 14.944 37.042 19-vi-2012 Sicilia Siracusa Noto 14.944 37.042 19-vi-2012 Sicilia Siracusa Canicattini Bagni 15.077 37.034 19-vi-2012 Sicilia Siracusa Canicattini Bagni 15.077 37.034 20-vi-2012 Sicilia Siracusa Noto 15.050 37.081 27-vi-2012 Campania Caserta Alife 14.376 41.298 28-vi-2012 Campania Caserta Sessa Aurunca 14.376 41.298 30-vi-2012 Molise Campobasso Civitacampomarano 14.223 41.720 26-vii-2013 Abruzzo Chieti Palena 14.134 41.978 27-vii-2013 Abruzzo Chieti Pretoro 14.143 42.214 30-vii-2013 Abruzzo Chieti Pizzoferrato/ 14.172 41.934 Gamberale 10-vii-2014 Campania Salerno Ceraso 15.263 40.175 12-vii-2015 Molise Campobasso Guardiaregia 14.539 41.434 17-viii-2016 Sicilia Messina Montalbano Elicona 15.028 37.994 19-viii-2016 Sicilia Messina Galati Mamertino 14.796 38.004 13-vi-2017 Abruzzo Chieti Pretoro 14.147 42.217

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Date Region Province Locality Long E Lat N 03-vii-2017 Campania Caserta Sessa Aurunca 14.376 41.298 04-vii-2017 Campania Caserta Pratella 14.181 41.405 04-vii-2017 Campania Caserta Teano/Tuoro/Furnolo 14.041 41.270 di Teano 05-vii-2017 Campania Benevento Pontelandolfo 14.668 41.282 11-vii-2017 Abruzzo Chieti Palena 14.120 41.963 13-vii-2017 Abruzzo Pescara Roccacaramanico 14.006 42.103 14-vii-2017 Abruzzo Chieti Pennapiedimonte 14.196 42.147 17-vii-2017 Abruzzo Pescara Salle vecchio 13.956 42.166 17-vii-2017 Abruzzo Pescara Tocco da Casauria 13.911 42.196 23-vi-2018 Calabria Reggio Calabria Straorino 15.753 38.132 23-vi-2018 Calabria Reggio Calabria Straorino 15.793 38.150 23-vi-2018 Calabria Reggio Calabria Straorino 15.789 38.152 23-vi-2018 Calabria Reggio Calabria Torrente Listì 15.838 38.143 23-vi-2018 Calabria Reggio Calabria San Lorenzo 15.853 38.034 24-vi-2018 Calabria Reggio Calabria S. Stefano in 15.782 38.165 Aspromonte 24-vi-2018 Calabria Reggio Calabria Masella-Montebello 15.759 37.972 Jonico 24-vi-2018 Calabria Reggio Calabria Melito Piani di Lopa 15.815 38.039 27-vi-2018 Calabria Reggio Calabria Santa Cristina 15.986 38.250 d’Aspromonte 22-vii-2018 Calabria Reggio Calabria Griffone 16.139 38.442 22-vii-2018 Calabria Reggio Calabria Canolo 16.205 38.312 22-vii-2018 Calabria Reggio Calabria Casalnuovo 15.977 38.075 22-vii-2018 Calabria Reggio Calabria Antonimina 16.140 38.273 22-vii-2018 Calabria Reggio Calabria Ciminà 16.114 38.255 23-vii-2018 Calabria Reggio Calabria Bagaladi 15.837 38.071 22-vii-2018 Calabria Reggio Calabria Griffone 16.168 38.419 23-vii-2018 Calabria Reggio Calabria Oppido Mamertina 15.986 38.034 25-vii-2018 Calabria Reggio Calabria Molochio 16.054 38.293 25-vii-2018 Calabria Reggio Calabria Oppido Mamertina 16.019 38.264 25-vii-2018 Calabria Reggio Calabria Santa Cristina 15.984 38.226 d’Aspromonte 25-vii-2018 Calabria Reggio Calabria Santa Cristina 15.992 38.226 d’Aspromonte

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Date Region Province Locality Long E Lat N 25-vii-2018 Calabria Reggio Calabria Santa Cristina 15.981 38.232 d’Aspromonte 26-vii-2018 Calabria Reggio Calabria Cinquefrondi 16.175 38.392 26-vii-2018 Calabria Reggio Calabria Cinquefrondi 16.127 38.405 26-vii-2018 Calabria Reggio Calabria Cinquefrondi 16.137 38.401 26-vii-2018 Calabria Reggio Calabria San Giorgio Morgeto 16.103 38.373 27-vii-2018 Calabria Reggio Calabria Cittanova 16.118 38.339 27-vii-2018 Calabria Reggio Calabria Cosoleto 15.884 38.207

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