Research Article |
Corresponding author: Andrew D. Liston ( andrew.liston@senckenberg.de ) Academic editor: Dominique Zimmermann
© 2015 Andrew D. Liston, Hans-Joachim Jacobs, Marko Prous.
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:
Liston A, Jacobs H, Prous M (2015) The Sawflies of Crete (Hymenoptera, Symphyta). Deutsche Entomologische Zeitschrift 62(1): 65-79. https://doi.org/10.3897/dez.62.4737
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Forty-two sawfly species are now known from Crete, including twelve species here recorded for the first time, and excluding earlier published records of Allantus didymus (Klug, 1818) based on misidentifications. Allantus nigrolinearis (Zirngiebl, 1937) is treated as a distinct species within the laticinctus-didymus group. Aneugmenus oertzeni (Konow, 1887) is allied to the Sardo-Corsican A. bibolinii Zombori, 1979 and belongs to Aneugmenus Hartig, 1837 s. str., not Atoposelandria Enslin, 1913. Pristiphora nievesi Haris, 2002 is a new synonym of P. tetrica (Zaddach, 1883). Of the newly recorded species, Chevinia mediterranea Lacourt, 2003, Dolerus puncticollis Thomson, 1871 and Empria archangelskii Dovnar-Zapolskij, 1929 are simultaneously new for the country of Greece. Four species (Allantus nigrolinearis, Aneugmenus oertzeni, Periclista cretica (Schedl, 1981) and Pristiphora sp. [subbifida group]) have not been recorded outside Crete, and may be endemic. It is not clear whether the morphologically and genetically distinctive Cretan specimens of Strongylogaster multifasciata (Geoffroy, 1785) should be regarded as a Cretan endemic species, for which the name S. cretensis Konow, 1887 is available, or as an isolated population of S. multifasciata. This requires further study, as also the taxonomic status of Heterarthrus imbrosensis Schedl, 1981 (only known from Crete) and H. wuestneii (Konow, 1905) (widespread in the West Palaearctic). The sympatric occurrence of three related Pristiphora species on Acer sempervirens, two of which differ in their choice of host individuals that are at different stages of vegetative development, is remarkable. A checklist of the Symphyta of Crete is presented.
Mediterranean, Tenthredinidae , taxonomy, distribution, phenology, endemism
The Greek island of Crete, in the southern Aegean Sea, is well known for its richness in endemic plant and animal species (
A visit to Crete by Liston alone at the end of March was followed by a visit by all three authors at the end of April. The first visit was centered on Iraklion, with day trips mainly into central Crete. Excursions during the second visit, based at Daratsos near Chania, were mainly in western Crete and focussed on localities in and around the Lefka Ori (White Mountains). Here we paid particular attention to endemic plant species such as Zelkova abelicea: a Tertiary relict, endemic Cretan member of the Ulmaceae (Kozlowski, 2013). The two excursions allowed us to do field work at a wide range of localities spread throughout most of the island except for the most easterly part. We collected using aerial nets with extendable handles, mainly by sweeping. Most material, including vouchers of all species, is deposited in the collection of the Senckenberg Deutsches Entomologisches Institut, Müncheberg, Germany (SDEI). Additional specimens are in the private collection of H.-J. Jacobs, and some duplicates in the US National Museum, Washington DC. Results of DNA barcoding (sequencing of part of the COI mitochondrial gene) conducted as part of the Barcode of Life Project (BOLD) (
The approximate coordinates and altitudes of the collection localities, listed alphabetically under their prefecture (from West to East), are:
Chania. Agia, 35.477°N, 23.930°E, 30 m. Agia Irini, 35.328°N, 23.837°E, 580 m. Askifou, 35.288°N, 24.191°E, 690 m. Daratsos, 35.506°N, 23.974°E, 30 m. Drakona, 35.406°N, 24.020°E, 510 m. Drakona 2, 35.416°N, 24.032°E, 320 m. Fournes, 35.418°N, 23.940°E, 200 m. Fournes-Meskla, 35.417°N, 23.950°E, 320 m. Kakopetros, 35.417°N, 23.750°E, 470 m. Kandanos, 35.333°N, 23.733°E, 420 m. Komitades, 35.208°N, 24.173°E, 170 m. Kournas and Georgiopolis, 35.347°N, 24.273°E, 25 m. Lakki, 35.371°N, 23.896°E, 900 m. Lousakies, 35.458°N, 23.629°E, 260 m. Meskla, 35.401°N, 23.955°E, 210 m. Moni, 35.285°N, 23.822°E, 110 m. Nea Roumata, 35.383°N, 23.850°E, 410 m. Omalos, 35.322°N, 23.913°E, 1100 m. Omalos 2, 35.325°N, 23.891°E, 1060 m. Platanias, 35.500°N, 23.900°E, 20 m. Samonas, 35.407°N, 24.108°E, 310 m. Sempronas, 35.376°N, 23.824°E, 660 m. Strovles, 35.369°N, 23.670°E, 430 m. Theriso-Drakona, 35.400°N, 23.983°E, 600 m. Topolia, 35.424°N, 23.687°E, 270 m.
Rethimnon. Anogeia, 35.289°N, 24.882°E, 750 m. Armeni, 35.295°N, 24.453°E, 400 m. Livadia, 35.303°N, 24.808°E, 395 m. Moni Kato Preveli, 35.171°N, 24.466°E, 170 m.
Iraklion. Krasi, 35.234°N, 25.467°E, 600 m. Marathos, 35.347°N, 24.972°E, 400 m. Mesa Potami, 35.213°N, 25.521°E, 720 m. Potamies, 35.256°N, 25.387°E, 170 m.
Lasithi. Katharo Plateau, Kopraki, 35.147°N, 25.565°E, 1150 m. Mesa Lasithi, 35.181°N, 25.512°E, 820 m. Pinakiano, 35.197°N, 25.464°E, 820 m.
Names of species not previously recorded in Crete are preceded by an asterisk (*) and those for which sequences of COI barcode region of Cretan specimens are available, are indicated by a dagger (†). Unless otherwise stated, specimens were collected by the authors.
We found no evidence that any sawfly species uses Zelkova abelicea as a host. However, we only encountered trees and bushes of this species during two visits to Omalos, and further fieldwork at this and other sites should be undertaken.
Crete; 1♂, Drakona 2, 22.iv.2013. 1♀, Moni, 28.iv.2013. 2♂♂, Agia Irini, 28.iv.2013.
Crete; 1♂, Potamies, 27.iii.2013, swept from inflorescences of Papaver rhoeas: a known host (
The name C. krueperi is used instead of C. similis (Mocsáry, 1880) for this taxon, as also in
= Emphytus balteatus var. nigrolinearis Zirngiebl, 1937: 646; ♀, holotype [not examined]. Type locality: Kristallenia (Kreta) [= Panagia Kristalenia, Lasithi Plateau, East Crete].
= Allantus didymus var. nigrolinearis (Zirngiebl, 1937);
Crete; 2♀♀, 1♂, Agia Irini, 26.iv.2013. 1♀, 1♂, Agia Irini, 28.iv.2013.
The barcodes of two sequenced A. nigrolinearis specimens (DEIGISHym20641, 20642) are identical and amongst 34 sequenced specimens of A. didymus and A. laticinctus sensu lato, are most similar to those of A. ariadne Liston and Jacobs, 2012 from Cyprus (DEIGISHym11089, 11093), with divergence of about 3.9%. Minimal divergence of A. nigrolinearis from A. laticinctus (France BC ZSM 01149 and Italy, Sicily DEIGISHym11042) is 6.4%. Noteworthy is that divergence of 5.3% between A. nigrolinearis and A. didymus (Klug, 1818) (Italy DEIGISHym18775, and Germany; 10 specimens) is somewhat less than from A. laticinctus. Several other segregates cluster around the four named taxa in the barcode similarity tree, indicating that the species group requires revision and may contain more taxa than previously thought.
As noted by
Female A. nigrolinearis differ from those of A. laticinctus and A. ariadne in having an uninterrupted, broad, median, black vitta through the otherwise red sterna of the abdomen (Fig.
A. laticinctus has only definitely been reared from Rosa, in central and western Europe (
Crete; 1♀, 6♂♂, Daratsos, on roses (garden cultivar) in hotel garden, 21–25.iv.2013. 1♂, Meskla, 22.iv.2013, near roses (garden cultivar). Rosa spp. are the known hosts:
The garden habitat and association of the Cretan specimens with horticultural varieties of rose, suggest that A. viennensis may have been introduced to the island. A. viennensis, probably native to southern and central Europe, is also an introduced and established species in the eastern USA (
Crete; 1♀, Strovles, 24.iv.2013.
One of the most cosmopolitan of sawflies, introduced to North America, South America and Australia from its (presumed) native range in the Palaearctic (
Crete; 1♀ “Ajos Joánnis Ep. Ajos Wássilis” 21.iii.1925, leg. A. Schulz, Museum für Naturkunde, Berlin. 1♀, Agios Nicolaos, Almyros (Lasithi), 11.iv.1976, leg. Matti Viitasaari, Viitasaari Collection, Helsinki. 1♀, Mesa Potami, 28.iii.2013, swept from Crataegus monogyna. 1♀, Anogeia, 29.iii.2013, swept from flowering Acer sempervirens. 1♀, 1♂, Katharo Plateau, Kopraki, 30.iii.2013, swept from low vegetation on riverbank. 1♀, Agia Irini, 26.iv.2013, swept from low vegetation on a dry slope.
Based on the serrulae of the lancet,
The closest species to E. archangelskii is E. excisa (Thomson, 1871). E. archangelskii can be easily distinguished by the red-yellow pale colour of its legs, with femora mainly pale (black and whitish in E. excisa, with femora mainly black). There appear also to be slight differences between lancets and penis valves: serrulae are more protruding in E. archangelskii (Figs
The hostplant(s) of E. archangelskii are unknown. However, the larvae of Empria species which are morphologically and genetically closest to E. archangelskii all feed on non-woody Rosaceae. The plants from which the specimens were collected and were growing near the collecting sites did not offer clues as to the identity of a possible host. The only Rosaceae obviously present at all Cretan collecting sites was Crataegus monogyna, but presumably the widespread Sanguisorba minor, Sarcopoterium spinosum and Rubus sanctus were also present somewhere not too distant.
Crete; 1♂, Agia, 29.iv.2013, netted from Veronica sp. growing in wet bed of a stream. Veronica spp. are known hosts (
Crete; 1♂, Potamies, 27.iii.2013. 4♀♀, Mesa Potami, 28.iii.2013. 1♀, Krasi, 28.iii.2013. 1♂, Daratsos, pool at hotel, 21.iv.2013. 1♀, 6♂♂, Omalos, 21.iv.2013. 1♀, Sempronas, 21.iv.2013. 1♂, Drakona, 22.iv.2013. 1♀, 8♂♂, Thériso-Drakona, 22.iv.2013. 1♂, Askifou, 23.iv.2013. 3♂♂, Kakopetros, 24.iv.2013. 1♀, 1♂ Strovles, 24.iv.2013. 3♀♀, 14♂♂, Omalos 2, 25.iv.2013. 14♀♀, 25♂♂, Agia Irini, 26.iv.2013. 1♂, Samonas, 27.iv.2013. 7♂♂, Moni, 28.iv.2013. 2♀♀, 3♂♂, Agia Irini, 28.iv.2013. 4♂♂, Agia, 29.iv.2013.
We did not collect all male specimens that were seen. During both visits to Agia Irini, males were very abundant in the uppermost part of the gorge.
Crete; 1♂, Strovles, 24.iv.2013.
Crete; 1♀, Drakona, 22.iv.2013, swept from Quercus sp. (either Q. ilex or Q. coccifera).
C. mediterranea was described from localities on the Mediterranean coast of mainland France and Corsica (
Crete; 1♀, Daratsos, pool at hotel, 22.iv.2013. 1♀, Meskla, 22.iv.2013. 2♀♀, Fournes-Meskla, 22.iv.2013. 1♀, Drakona, 22.iv.2013. 4♀♀, Agia Irini, 26.iv.2013. 1♀, Samonas, 27.iv.2013. 1♀, Moni, 28.iv.2013.
Crete; 5♀♀, 7♂♂, Omalos, 25.iv.2013, swept from Acer sempervirens: the host (
the barcode of one specimen collected from Acer sempervirens from the Peloponnese and determined as H. wuestneii (Konow, 1905) (DEIGISHym11102), diverges by 0.7% from two Lower Austrian specimens of H. wuestneii (BC ZSM HYM 04121, 04122). The minimum barcode divergence of the two barcoded Cretan H. imbrosensis (DEIGISHym20655, 20656) from the Peleponnesian H. wuestneii specimen is 1.8%.
H. imbrosensis closely resembles H. wuestneii. Specimens from Crete are darker than Central European (German and Austrian) H. wuestneii. The face may be entirely black in Cretan females, but usually there are at least small white patches on the inner orbits, whilst the tegula varies from entirely black to black with the posterior edge white. The H. wuestneii specimens from the Peloponnese are intermediate in coloration. According to
Crete; 3♀♀, 32♂♂, Krasi, 27–28.iii.2013. 3♂♂, Livadia, 29.iii.2013. 2 larvae (Fig.
The buds of the bushes and small trees of Quercus coccifera from which these were collected were swollen, but had not burst. Males were obtained mostly by netting them as they flew around the twigs in the upper crowns (at Krasi reached by standing on the many large boulders), and less so by sweeping around all accessible lower parts of the oaks. Sweeping yielded the only few females that we were able to obtain. The identification of the larvae allows us to state that Q. coccifera is the host plant. P. cretica was previously only known from the female holotype and a single male paratype (
The taxonomy of this species will be dealt with in detail in a revision of West Palaearctic Periclista (A. Mol, in preparation).
Crete; 1♀, Krasi, 28.iii.2013. 1♀, Agia Irini, 26.iv.2013. Swept from low vegetation containing Sanguisorba minor, which is possibly a host plant (
Crete; 1♀, 6♂♂, Topolia, 29.iv.2013, on Rosa sp.: the main hosts are Rosa spp. (
Crete; 2♀♀, 4♂♂, Moni Kato Preveli, 25.iii.2013, swept from Salix alba, one of the main hosts:
Crete; 3 galls in leaves of Salix alba, Meskla, 22.iv.2013. Galls very abundant (hundreds) on Salix alba, Agia, 29.iv.2013.
No adults were found by sweeping foliage of the three large trees at Agia. The majority of larvae were very small when about 200 galls were collected. Nevertheless, about 30 individuals had spun cocoons by 7.v.2013. Three females emerged in June 2013. Some others seemed to have entered prolonged diapause, but no adults were reared.
Crete; 1♀, Marathos, 24.iii.2013. 2♀♀, Pinakiano, 27.iii.2013. 2♀♀, Mesa Potami, 28.iii.2013. 1♀, Anogeia, 29.iii.2013. 2♀♀, Omalos, 25.iv.2013. Specimens mostly collected from inflorescences of Pyrus cultivars: the normal hosts are Pyrus spp. (
Crete; 4♀♀, Marathos, 24.iii.2013. 9♀♀, Pinakiano, 27.iii.2013. 7♀♀, Mesa Potami, 28.iii.2013. 1♀, Krasi, 28.iii.2013. 10♀♀, Anogeia, 29.iii.2013. 1♀, Omalos, 21.iv.2013. Most specimens were swept from Crataegus monogyna with unopened inflorescences, and some others from inflorescences of cultivated Pyrus growing near Crataegus.
At none of the localities were Prunus spp. seen, although these are usually considered to be the larval hosts of H. chrysorrhoea in northern parts of Central Europe (e.g.
Crete; 1♂, roadside between Kournas and Georgiopolis, 26.iii.2013. 1♀, Livadia, 29.iii.2013. Both specimens swept from freshly flushed Quercus pubescens: Quercus spp. are the known hosts (
The depressed medial areas of the apical abdominal terga of the male are partly pale, unlike the completely dark terga of males so far examined from mainland Greece (see
Crete; 4 larvae on cultivated Pyrus sp., Fournes, 25.iv.2013. One larva, larger than the others, had spun a cocoon by 7.v.2013.
Crete; 7♂♂, Pinakiano, 27.iii.2013. 5♂♂, Mesa Lasithi, 28.iii.2013. 1♀, 8♂♂, Katharo Plateau, Kopraki, 30.iii.2013. 6♀♀, 8♂♂, Omalos, 21.iv.2013. 11♀♀, 19♂♂, Askifou, 23.iv.2013. 7♀♀, 16♂♂, Omalos, 25.iv.2013. 2♀♀, Agia Irini, 26.iv.2013. All specimens swept from Acer sempervirens: because all species of the Pristiphora subbifida group use Acer spp. as hosts (
Minimum divergence of three Cretan P. cretica (DEIGISHym20657–20659) from two northern Greek P. cretica (DEIGISHym11052, 19646) is approximately 1.2%. Interspecific divergence from the next nearest species, P. schedli (Cyprus: DEIGISHym10980, 10981) is about 3.8%.
In Crete Acer sempervirens displays considerable infraspecific variability in the timing of bud burst. Although Pristiphora cretica occurred at some localities together with P. tetrica (Zaddach, 1883), it was observed that these species have strongly differing preferences for trees at different stages of leaf and flower development. P. cretica is found exclusively on trees whose buds have fully opened and which are already flowering, whereas P. tetrica is found on trees whose buds have not yet burst, or have just opened. The preference is so strong, that this phenomenon could be observed on trees standing beside each other which were at different stages of flushing (Fig.
The Cretan specimens listed above closely resemble the female holotype and male paratype (examined:
– | Metafemur at least with black stripes on anterior and posterior edges of dorsal face. Female with clypeus and labrum pale marked; male with at least labrum pale (brown). Abdominal terga largely black, rarely yellow (Peloponnesian ♀♀) and if so then tergum 1 also largely yellow. Female length 5–6 mm; male 4–5 mm | P. cretica |
– | Metafemur entirely yellow, except for black area on extreme base. Clypeus and labrum entirely black. Abdominal tergum 1 black; other terga yellow, except at most for small medial flecks on terga 2–3. Female length 4.0–4.5 mm; male 3.5–4.0 mm | P. tetrica |
Crete; 1♀, Daratsos, pool at hotel, 21.iv.2013. 1♀, Theriso-Drakona, 22.iv.2013.
These specimens are of the colour form with a very pale abdomen, which has sometimes been treated as a separate species, P. denudata Konow, 1902.
Crete; 1♀, Krasi, 27.iii.2013. 1♀, Livadia, 29.iii.2013. Both specimens swept from Quercus coccifera.
Crete; 1♂, Omalos, 21.iv.2013. Swept from Acer sempervirens, which is probably a host plant, because this is the only Acer sp. occurring in Crete and all species of the Pristiphora subbifida group use Acer spp. as hosts (
Barcode data: the sequence of this specimen (DEIGISHym20661) diverges by about 4.8% from its nearest neighbour, P. tetrica (Zaddach, 1883) from Sicily (DEIGISHym10972). Divergence from two Cretan P. tetrica (see below) is approximately 5.3% and from P. schedli Liston and Späth, 2008 (Cyprus) approx. 8.6%.
Externally, the specimen differs from Cretan Pristiphora tetrica males only in the apex of the metatibia and most of metabasitarsomere being pale: Fig.
= Nematus tetricus Zaddach in Brischke, 1883: 148–149; ♀, holotype. Type locality: Gumperda, Thüringen. Type probably destroyed (
= Nematus velatus Zaddach in Brischke, 1883: 149; ♀, holotype. Type locality: Baiern [Bavaria]. Type should be in the Zoologische Staatssammlung, Munich, but was not located (
= Pristiphora tetrica:
= Pristiphora nievesi Haris, 2004: 164–165; ♀, holotype and paratypes [Museo Nacional de Ciencias Naturales, Madrid; examined]. Type locality: El Ventorillo, Madrid. New synonym.
Crete; 7♀♀, 3♂♂, Pinakiano, 27.iii.2013. 2♀♀, 11♂♂, Mesa Lasithi, 28.iii.2013. 1♂, Livadia, 29.iii.2013. 1♀, 4♂♂, Anogeia, 29.iii.2013. 5♀♀, 5♂♂, Katharo Plateau, Kopraki, 30.iii.2013. 1♀, 1♂, Omalos, 21.iv.2013. 2♀♀, Omalos, 25.iv.2013. All specimens swept from Acer sempervirens: because all species of the Pristiphora subbifida group use Acer spp. as hosts (
Barcode data: sequences have been obtained for five specimens that were previously identified as P. nievesi: one specimen each from France, DEIGISHym19640; Italy, Sicily, DEIGISHym10972; mainland Greece, DEIGISHym10965, and two from Crete (DEIGISHym20660, 20662). Nested within this cluster in the barcode similarity tree are three specimens of P. tetrica from Germany (BC ZSM HYM 09418, BC ZSM HYM 20222, DEIGISHym18864). Although the minimum sequence divergence of the Cretan specimens from the German ones is about 1.9%, the Sicilian specimen differed from the German ones by only 0.6%. This is a comparatively small barcode difference (
P. tetrica and P. nievesi have been recognized as being morphologically the most similar Pristiphora species within the Pristiphora subbifida group (
– | At least metafemur mainly yellow. Abdomen predominantly yellow; black are only tergum 1 and at most small medial flecks on terga 2–3. Wing membrane subhyaline. Female length 4.0–4.5 mm; male 3.5–4.0 mm | P. nievesi |
– | Hind legs completely black. Abdomen usually with numerous terga medially and contiguously black. Wing membrane fuscous. Female length 5.0–6.5 mm; male 4.0–5.0 mm | P. tetrica |
Occasional Central European specimens of P. tetrica occur that are unusually pale. They are larger than the types of P. nievesi but similarly coloured, apart from having black metafemora. One such individual is the type of Nematus velatus, described as having an almost completely yellow abdomen and less fuscous wings than typical P. tetrica. Specimens formerly identified as P. nievesi differ from each other in the colour pattern of the legs. The metatibia of the type specimens and the French specimen (
See comments on phenology and distinction of P. tetrica from P. cretica, under P. cretica (above). Colour variability in P. tetrica from Crete is in the extent of pale on the outer orbits, tegulae and posterior edges of the pronotum. These parts are usually partly pale, but are entirely black in the darkest individuals. Males tend to be darker than females. Occasionally in females, the mesepisternum is partly pale medially.
Crete; 2♀♀, 2♂♂, Marathos, 24.iii.2013. 20♀♀, 23♂♂, Armeni, 25.iii.2013. 2♀♀, Lakki, 21.iv.2013. 7♀♀, 19♂♂, Sempronas, 21.iv.2013. 2♀♀, 5♂♂, Drakona, 22.iv.2013. 5♀♀, 4♂♂, Theriso-Drakona, 22.iv.2013. 7♀♀, 3♂♂, Strovles, 24.iv.2013. 7♀♀, 11♂♂, Agia Irini, 26.iv.2013. 2♀♀, 12♂♂, Topolia, 29.iv.2013. 1♀, 16♂♂, Agia, 29.iv.2013. All swept from, or netted near, Pteridium aquilinum, the only known host (
The barcodes of four sequenced Cretan specimens (DEIGISHym20637–206340: infraspecific variability about 0.7%) diverge by a minimum of 6.1% from the nearest neighbour A. bibolinii Zombori, 1979, represented by one male each from Corsica (first record from France: Ascu, 02.05.2004, leg. Liston, SDEI (DEIGISHym19725) and Sardinia (DEIGISHym19684): infraspecific variability about 0.7%. Minimum divergence of A. oertzeni to the next nearest species A. padi is 9.2% (many specimens from central and southern Europe, and Iran).
According to
Crete; 1♀, Katharo Plateau, Kopraki, 30.iii.2013.
As well as being the first record of a Dolerus species from Crete, this also seems to be the first published record of D. puncticollis from Greece. This is the most widespread Dolerus (Poodolerus) species in the Mediterranean Region, and the only member of the subgenus known in North Africa (
Crete; 6♂♂, Marathos, 24.iii.2013. 2♀♀, 4♂♂, Armeni, 25.iii.2013. 1♂, Lakki, 21.iv.2013. 3♂♂, 1♀, Sempronas, 21.iv.2013. 1♀, 13♂♂, Thériso-Drakona, 22.iv.2013. 2♀♀, 3♂♂, Drakona, 22.iv.2013. 9♀♀, 6♂♂, Strovles, 24.iv.2013. 3♀♀, 2♂♂, Agia Irini, 26.iv.2013. 3♀♀, 2♂♂, Agia Irini, 28.iv.2013. 1♂, Topolia, 29.iv.2013. 1♂, Agia, 29.iv.2013. All swept from, or netted near Pteridium aquilinum, which is the only known host in Crete:
The barcodes of three sequenced Cretan specimens (DEIGISHym20648–20650) are identical and diverge by a minimum of 3.2% from 24 specimens of S. multifasciata collected throughout Europe (from Scotland to the Peloponnese) and Iran (infraspecific variability about 0.5%).
The Cretan specimens all belong to the form described as S. cretensis Konow, 1887, which was regarded as an allopatric local population of S. multifasciata by
– | Postocellar area with sculpture similar to that of adjoining parts of the temples: punctures on postocellar area about as large as those on temples, with shiny unsculptured interspaces (Fig. |
S. cretensis |
– | Postocellar area with denser sculpture than on adjoining parts of temples: average puncture diameter on postocellar area smaller than on temples, with interspaces matt (Fig. |
S. multifasciata |
Further studies, including sequencing of nuclear genes, are required to clarify the status of the Cretan taxon.
Crete; 7♀♀, 1♂, Nea Roumata, 21.iv.2013. 1♂, Fournes-Meskla, 22.iv.2013. 1♂, Drakona, 22.iv.2013. 1♂, Drakona 2, 22.iv.2013. 1♀, Askifou, 23.iv.2013. 1♀, Komitades, 23.iv.2013. 1♀, 3♂♂, Vrises, 23.iv.2013. 1♀, Kakopetros, 24.iv.2013. 1♂, Kandanos, 24.iv.2013. 2♂♂, Platanias, 24.iv.2013. 1♀, Strovles, 24.iv.2013. 1♀, Fournes, 25.iv.2013. 2♂♂, Samonas, 27.iv.2013. 4♂♂, Agia Irini, 28.iv.2013. 1♂, Topolia, 29.iv.2013. 1♀, Lousakies, 29.iv.2013.
Crete; 1♂, Krasi, 28.iii.2013. 1♀, Samonas, 27.iv.2013. 2♂♂, Lousakies, 29.iv.2013. 1♂, Topolia, 29.iv.2013.
Crete; 1♀, Fournes-Meskla, 22.iv.2013.
A remarkable species because of its thelytoky, unusual in the Cephidae, and its mainly southern European distribution which extends north along the Atlantic coast to the French Department of the Manche (
Crete; 7♀♀, 1♂, Moni, 28.iv.2013.
In contrast to the examples mentioned above, in rather more cases sawfly species of the same genera which feed on the same or closely related hosts belong to different species on each island. Examples are: on Quercus in Crete Mesoneura opaca, Pristiphora parnasia, Periclista cretica and possibly Chevinia mediterranea / on Quercus in Cyprus Mesoneura lanigera Benson, 1954, Pristiphora calliprina Liston & Jacobs, 2012 and Periclista rufiventris Zombori, 1979. On Acer in Crete Heterarthrus imbrosensis, Pristiphora cretica, Pristiphora sp. [subbifida group] and P. tetrica / on Acer in Cyprus Heterarthrus cypricus Schedl, 2005 and Pristiphora schedli Liston & Späth, 2008. We consider that these groups of species are assemblages which are naturally present on each island and that they illustrate a profound dissimilarity of their sawfly faunas, when these are compared at species level. On the other hand, six of the eight species mentioned above from Crete also occur in mainland Greece.
Both Crete and Cyprus have been separated from continental landmasses for at least 5 million years, and their degree of isolation since has sometimes been stated to be similar to that of today (
Several other more recent events have at times been considered to be linked to a high rate of extinction in the Cretan flora and fauna, including for example the establishment of Neolithic culture 8400–7500 BP and the eruption of Santorini at around 3600 BP (
In many cases it is still impossible to state categorically whether a particular sawfly species is endemic to a Mediterranean island. This problem was discussed by
Three species of Pristiphora belonging to the same species group occur in Crete on the same host (Acer sempervirens) and partly at the same localities. It is remarkable that adults of two of these species were found on individual trees at different stages of vegetative development: P. tetrica on plants at a less advanced stage than those frequented by P. cretica. It would be interesting to know if the third species, Pristiphora sp. [subbifida group], also differs in this habit. Although a few examples of differing phenology between closely related species on the same host are known in other Tenthredinidae (e.g. Parna apicalis (Brischke, 1888) and P. tenella (Klug, 1816): leaf-miners on Tilia), P. cretica and P. tetrica are exceptional in that their adults can be found at the same locality on the same day, but on different trees. This is made possible by the very large variability in vegetative phenology of the host. Although the larva of P. cretica remains unknown, larvae of other species in the subbifida group are similar in appearance and behaviour: cryptically coloured, solitary, leaf-edge feeders. The rather low abundance of adults at any site, the small size of individuals, the [probably] solitary habits of the larvae and the abundance of the food resource (leaves) lead us to suppose that competition between the larvae of these species is negligible. It would be very interesting to know more about the apparently differing behavioural strategies of the Cretan Acer-feeding Pristiphora.
The quantitative similarity between the sawfly faunas of Crete and Cyprus identified by
Although the Cretan sawfly fauna is species poor, it is valuable and worthy of conservation not only because of its proportionately high number of endemic species and genetically distinctive populations, but also because of those which are elsewhere known only from the southern Balkan peninsula (Pristiphora cretica), or are isolated populations on the western edge of the species range (Empria archangelskii, Mocsarya syriaca).
Dr Manfred Kraus, Dr Stefan Schmidt, Dr Olga Schmidt (Zoologische Staatssammlung, Munich), Dr Stephan M. Blank, Dr Andreas Taeger (SDEI) and the staff of the Barcoding of Life Data System (University of Guelph, Ontario, Canada) helped us to obtain barcode sequences of some specimens. Dr Frank Koch (Museum für Naturkunde, Berlin), Mr Henri Savina (Toulouse), Prof. Dr Wolfgang Schedl (Innsbruck), Mrs Lisa Standfuss, Prof. Dr Klaus Standfuss (Dortmund), Mr Matti Viitasaari (Helsinki) and Prof. Dr Meicai Wei (Changsha) made specimens available for examination. Mrs Gillian Galway drew our attention to literature which mentions the transport of plant material in the ancient eastern Mediterranean. Dr Akihiko Shinohara (Tokyo) and Dr Veli Vikberg (Turenki) reviewed the manuscript and suggested improvements. We are grateful to the Museum für Naturkunde, Berlin, for enabling open access publication of this work.
Checklist of the Symphyta of Crete
Xyelidae
Xyela graeca (J. Stein, 1876)
Cephidae
Cephus pygmeus (Linnaeus, 1767)
Calameuta haemorrhoidalis (Fabricius, 1781)
Calameuta idolon (Rossi, 1794)
Phylloecus faunus Newman, 1838
Trachelus tabidus (Fabricius, 1775)
Trachelus troglodyta (Fabricius, 1787)
Argidae
Arge melanochra (Gmelin, 1790)
Arge ochropus (Gmelin, 1790)
Arge scita (Mocsáry, 1880)
Cimbicidae
Corynis krueperi (J. Stein, 1876)
Tenthredinidae
Allantinae
Allantus nigrolinearis (Zirngiebl, 1937)
Allantus viennensis (Schrank, 1781)
Ametastegia glabrata (Fallén, 1808)
Athalia ancilla Serville, 1823
Athalia circularis (Klug, 1815)
Athalia cordata Serville, 1823
Athalia rosae (Linnaeus, 1758)
Empria archangelskii Dovnar-Zapolskij, 1929
Blennocampinae
Chevinia mediterranea Lacourt, 2003
Heterarthrus imbrosensis Schedl, 1981
Periclista cretica (Schedl, 1981)
Silliana lhommei (Hering, 1934)
Halidamia affinis (Fallén, 1807)
Nematinae
Cladius ordubadensis Konow, 1892
Cladius pectinicornis (Geoffroy, 1785)
Hoplocampa brevis (Klug, 1816)
Hoplocampa chrysorrhoea (Klug, 1816)
Mesoneura opaca (Fabricius, 1775)
Euura atra (Jurine, 1807)
Euura proxima (Serville, 1823)
Pristiphora abbreviata (Hartig, 1837)
Pristiphora cretica Schedl, 1981
Pristiphora pallidiventris (Fallén, 1808)
Pristiphora parnasia Konow, 1902
Pristiphora sp. [subbifida group]
Pristiphora tetrica (Zaddach, 1883)
Selandriinae
Aneugmenus oertzeni (Konow, 1887)
Dolerus puncticollis Thomson, 1871
Strongylogaster multifasciata (Geoffroy, 1785)
Orussidae
Mocsarya syriaca Benson, 1936
Orussus morio Guiglia, 1954