Sunday 13 January 2019

An updated European species list

For several years Fauna Europaea (FE) offered a stabe taxonomical list for European fauna, including butterflies. Funding however for the FE project had dried up and several attempts to keep the project running seemed to fail. Meanwhile new developments in taxonomical research couldn't be implemented in the FE species lists. This led to a situation where the FE species list for butterflies became outdated and as such less suitable as a widely accepted tool in butterfly research and conservation. For this reason the species specialists for butterflies on FE took the initiative to write a paper summarizing the most important updates to the European butterfluy species list. 
This paper can be found here:

But as not everyone likes to read these kind of articles or is well aware of the old taxonomy, I will try to summarize the most important changes in this blogpost.
The most important changes can be grouped into a few categories.

- First of all there are a bunch of new taxa in the list, these can be divided in two groups:

Newly discovered species
As the European butterfly fauna is already well known only few species fit in this category. 
In a few cases this involves names that where already in use for a longer time, although the species discoveries are really new and in some cases unexpected.

Spialia rosae Hernández-Roldán, Dapporto, Dincă, Vicente & Vila, 2016
This taxon was split from S. sertorius based on constant differences in mtDNA, chemical profiles and ecology. At this point only known from Spain, although presence in mountainous neighbouring regions in Portugal or France is possible. Larval foodplant is not Sanguisorba species like in most related species but Rosa species.   

Spialia rosae egg on Rosa species, at the LT in the Sierra Nevada.

Zerynthia cassandra (Geyer, 1828)
This taxon was split from Z. polyxena based on differences in mtDNA and male genitalia. It is an endemic from the Italian peninsula and Sicily, where this pic was made.
For this taxon an older name has been digged up as this was the oldes available description for this taxon (LT in Tuscany). This name has been used to describe specimens from southern France as well but now this name has come inavailable to describe the French specimens.

Zerynthia cassandra in Sicily

Leptidea juvernica Williams, 1946
For some time now we know that the Leptidea sinapis complex comprises of 3 species. Leptidea reali seems to be comprised to the western Mediterranean area while Leptidea juvernica is present - often sympatric with L. sinapis - in large parts of Europe. 

Polyommatus timfristos Lukhtanov, Vishnevskaya & Shapoval, 2016
Based on differences in karyotype & mtDNA another species was raised in the Agrodiaetus subgenus. Determination in the field with Polyommatus ripartii seems hazardous... For now on only confirmed from the very south of mainland Greece.

Polyommatus celina (Austaut, 1879)

The difference between the group of realy new species and subspecies raised to species level is sometimes fine. The knowledge that specimens of P. icarus in S Spain, N-Africa & some Mediterranean islands were a bit different for example, was already long known but these differences were long considered to be very small. Wiemers & Fiedler (2007) were one of the first to notice the large genetic differences between these populations and the ones in the rest of Europe but on the external characters even write: "while in the latter species phenotypic differences have never been noted."
More research confirmed Polyommatus celina to be sufficiently different of P. icarus to be considered a species on its own.

A female Polyommatus celina on Fuerteventura

Taxa raised from subspecies to species level

Long known subspecies but now evaluated as considerably distant from the nominate.

Iphiclides feisthamelii (Duponchel, 1832)
Species rank is based on the differences in genitalia and nucleair DNA (see Fig. 6 here). The taxon replaces I. podalirius on the Iberian peninsula & Northern Africa. More information on the border region of the two taxa in Southern France can be found here.

Iphiclides feitshamelii from Barcelona

Iolana debilitata (Schultz, 1905)
The Iberian & Northern African Iolana populations seem to differ in morphology and genetics.

Pseudochazara tisiphone Brown, 1981

The genus Pseudochazara is a brain-racking genus with several very closely related taxa. Above that determination is hindered by large local variation (for example), apparently often induced by the local rock colour. A first step to unravel the complex via genetic research was set by Takáts & Mølgaard. Where this article was focussing on the Balkanic taxa, a next article by Verovnik & Wiemers soon followed focussing on a wider scope. Out of that research it became clear that the above taxon was not really related to the Turkish P. mniszechiiI.

Pseudochazara tisiphone from Albania

Pseudochazara amalthea (Frivaldszky, 1845)
Out of the same Pseudochazara research it became clear that P. amalthea from the Balkanic mainland was distinct from the nominate P. anthelea from the Greek Islands and Turkey.

Pseudochazara amalthea from Albania

Erebia arvernensis Oberthür, 1908 & Erebia neleus (Freyer, 1832)
Erebia cassioides s.l. seems to consist of three closely related but molecularly different taxa. Erebia arvernensis from the Western Alps to Spain, Erebia cassioides in the Eastern Alps and the Italian peninsula and Erebia neleus in the Balkan. The main base for this split can be found in this article.

Erebia arvernensis in the French Alps

Melitaea celadussa 
Fruhstorfer, 1910 

Apart from small differences in the genitalia the southwestern representatives from the Melitaea athalia complex seem to be genetically different as well, see following article.

Melitaea celadussa (Drôme, France)

Melitaea ornata Christoph, 1893

The populations with red headed larvar in the M. phoebe complex seem to consist out of multiple species, the one in Europe is now called M. ornata while M. telona is confined to the Middle East

Melitaea ornata in Greece

- Name changes: these changes can involve 3 types of changes

Re-evaluation of the species name

Especially in older names there can always be some discussion on what name was published first.

Pyrgus foulquieri (Oberthür, 1910)

As descriptions of both Syrichtus alveus f. foulquieri and Syrichtus alveus f. bellieri were published simultaneously (Oberthür, 1910), the name used by the first reviser (i. e. Rebel 1914), Pyrgus foulquieri, should therefore be used.

Proterebia phegea (Borkhausen, 1788)

Another long known enigmatic species whose nomenclature has been subject for discussion since long time. The name Papilio afer was already in use for an African species of Nymphalid when Esper named this taxon as such. So the next name that pops up is Papilio phegea Borkhausen, 1788.

Pseudochazara mercurius (Staudinger, 1887)

This taxon only reaches Europe in the very southeast of European Russia on the slopes of the southern Ural. It was long known as Pseudochazara hippolyte and the Iberian P. williamsi was long considered a subspecies of it but was already considered full species in the previous FE checklist. P. hippolyte seems to be an unvalid name so the next name that pops up to describe this taxon is Pseudochazara mercurius.

Re-evalution of the genus/subgenus
Especially in the Lycaenidae several former subgenera have been reinstated as genus, especially in the large genera Plebejus and Polyommatus. This descision is based on this research.
Following species of Lycaenids are concerned:

Chilades galba                    ---> Luthrodes galba
Chilades trochylus              ---> Freyeria trochylus
Plebejus orbitulus               ---> Agriades orbitulus
Plebejus optilete                 ---> Agriades optilete
Plebejus pyrenaicus           ---> Agriades pyrenaicus
Plebejus dardanus             ---> Agriades dardanus
Plebejus zullichi                 ---> Agriades zullichi
Plebejus glandon               ---> Agriades glandon
Plebejus aquilo                  ---> Agriades aquilo
Plebejus loewii                   ---> Plebejidea loewii
Aricia eumedon                  ---> Eumedonia eumedon
Plebejus psylorite               ---> Kretania psylorita
Plebejus hespercia             ---> Kretania hesperica
Plebejus eurypilus              ---> Kretania eurypilus
Plebejus trappi                   ---> Kretania trappi
Plebejus sephirus               ---> Kretania sephirus
Plebejus pylaon                  ---> Kretania pylaon
Polyommatus cyane           ---> Glabroculus cyane
Polyommatus coelestinus   ---> Neolysandra coelestina
Polyommatus hispana         ---> Lysandra hispana
Polyommatus corydonius    ---> Lysandra corydonius  
Polyommatus bellargus       ---> Lysandra bellargus 
Polyommatus coridon          ---> Lysandra coridon
Polyommatus caelestissima ---> Lysandra caelestissima
Polyommatus albicans         ---> Lysandra albicans

Luthrodes galba

But also in the genus Argynnis there are some changes. Mainly based on following article.

Argynnis aglaja       ---> Speyeria aglaja

Argynnis elisa         ---> Fabriciana elisa
Argynnis niobe        ---> Fabriciana niobe
Argynnis adippe      ---> Fabriciana adippe

Speyeria aglaja seems to be more related to the N-American genus Speyeria than to Argynnis s.l.

Some small changes in the gender of species names

Several species have got a different gender to their species names. 
For example Colias croceus has become Colias crocea. For a full list I refer to the original article.

- Taxa no longer considered valid 

Polyommatus eleniae Coutsis & De Prins, 2005

In 2005 2 species from the brown Agrodiaetus subgenus complex were described within a few months time, one from NE Greece but the first one was found not that far across the border in S Bulgaria, Polyommatus orphicus Kolev, 2005. Recent genetic research (see earlier with Polyommatus timfristos) confirms these taxa to be one species.

Polyommatus eleniae orphicus, found in NE Greece

- Some remarks on possible changes that didn't make it in the article

This new checklist offers butterfly researchers & conservationists a stable list again to go on for a few years so they don't need to stay stuck to the knowledge from 20 years ago and butterfly research and conservation . 

But this ofcourse doesn't mean that butterfly taxonomy is settled and there isn't anything left we can learn. There are still some older taxa with unclear status remaining in the checklist such as Pieris balcana & other taxa in the P. napi complex or Erebia sthennyo (see for example). On the other side of the discussion the last few years saw lots of new research arise discovering more cryptic diversity in European butterflies but in lots of cases the exact relationships between genetic lineages still have to be unentangled. So when time's ready, be prepared for more taxonomical finetuning of European butterfly fauna

Pyrgus armoricanus is one of the taxa that possibly is in fact a complex of 2 species

Friday 13 April 2018

Early spring in S France

Last week I spent some time in Southern France. Unfortunately weather wasn't very favorable for butterfly watching so I had to cancel some of my plans due to predicted torrential rainfall. I have missed out on some targets because of that but still have enough to share.

Zerynthia polyxena

Two of my primary targets were the two festoon species present in S France. I had seen Zerynthia polyxena sensu lato a few years earlier in Sicily but recently it became clear that the populations in Italy south of the river Po are different enough - genetically as well as in the male genitalia - to be threated as a seperate species, Zerynthia cassandra (see this article and see here for pictures). 
So it was time to see Z. polyxena ss. and I was lucky enough to find some individuals in the low plains of the Camargue. Second picture shows the local suspected larval foodplant Aristolochia rotunda.

Zerynthia rumina

This species seems to favor more hilly and rocky terrain than the previous and I found some individuals N of Perpignan. Its distribution spreads from northern Africa to S France. Last picture again shows the larval foodplant, Aristolochia rotunda.

Euchloe tagis

This as well is a species with a distribution from Tuscany and S France to N Africa with some regional variation. This variation has lead to the description of several subspecies, in S France we are talking about the well patterned subspecies bellezina. Unlike its common look-a-like Euchloe crameri it is local and only to be found on calcareous, rocky locations with the larval foodplant Iberis species in abundance.

Iberis species observed on this location

Anthocharis euphenoides
I had seen this species before in Spain but was then unable to make pictures of it. Present from Iberia to S Italy.

Callophrys avis

This is another western Mediterranean specialist, local in S France. Although the more common Callophrys rubi can certainly be seen on it the trick to find this species is definetly to search on or nearby the larval foodplant, Arbutus unedo.

This last picture shows Callophrys rubi on an Arbutus unedo plant, check however the white 
bordering the eyes. This is white in C. rubi but reddish brown in C. avis.

Pseudophilotes baton
This one gets an entry on this page because of these nice individuals from the taxon 'praepanoptes' that form a transition between P. baton & P. panoptes in the Pyrenées Orientales. 

Wednesday 30 August 2017

Romania - Eastern Transylvania & the foothills of the eastern Carpathians

After visiting Dobrogea we headed back NW to visit some grassland habitats at the foothills of the Eastern Carpathians (800-1000m). Along the way we stopped here & there to have a quick scan of the butterfly fauna.

Lycaena dispar, males
In a lower valley (250m) still on the east side of the Carpathians.

Lycaena virgaureae, male
Higher up in the Carpathians (950m) this fiery beauty flew around. 

Nymphalis antiopa
Several seen and all very fresh early august.

Coenonympha glycerion
On lower grasslands, E of the Carpathians and in central Transylvania. All relatively fresh, I don't know if these are belated ones (as in the W part of its distribution this species flies in June - early July) or if this species has a second generation here.

Minois dryas
On the nicer grasslands one of the most common butterflies

Colias myrmidone, female

Colias myrmidone, male in flight showing the absence of pale veins in the dark band on the frontwing and showing the very slender dark band on the hindwing. The colour on the upperside is much brighter than in C. croceus giving it an almost fluorescent appearance in the field.

Colias myrmidone, male

Colias myrmidone, female

Colias myrmidone, female (L) & male (R) in flight. 
The large pale spots in the marginal band of the female are obvious and the male is showing absence of pale veins, slender black edging to hindwing with the impression of pale spots on the innerside of the black edging and fluorescent orange ground color.

My number one target for this trip was to see Colias myrmidone. This species has got a tremendous backfall during the last decades. Until the late nineties the distribution reached into the eastern part of Germany, the last observation there however was made in 2000. The species has disapeared as well out of Czech Republic, Austria, Slovenia, Hungary and probably Bulgaria. The situation in Slowakia is unclear to me, could still be there but I have not seen any recent information. Also in Poland the situation is unclear, possibly still present in the east of the country. Romania seems to be the stronghold for the species in the EU and because of the big decrease in such a short period the species is listed as "Critically Endangered" on the EU27 red list. The reasons for the decrease are still a bit unclear but most probably changes in land management have a big hand in this. The species is dependent on thriving growth of the larval foodplant Chamaecytisus ratisbonensis and this plant grows mainly on grasslands on a specific point in succession in the beginning of afforestation. Intensification under the form of large scale mowing has a negative effect on the foodplant growth and abandonement leads to too much afforestation making other plants overgrowing the larval foodplant. Traditional mowing practices leaving parts of grasslands unmowed and extensive grazing practices are still common in some parts of Romania giving chances to C. myrmidone. And with the large decrease of the species in Europe some of the Romanian grasslands are now protected areas so lets hope this is enough to stop the further decrease of the species.

Habitat at the foothills of the Eastern Carpathians (900-1000m)

Tuesday 29 August 2017

Romania - Dobrogea: plains, wetlands & oak forests

Dobrogea is the Romanian coastal region stretching from the Bulgarian to the Ukranian border. it is organised in a southern county Constanta & a northern county Tulcea. Both are named after their respective main city.
We visited some coastal plains, salt marshes and lower hills from the north of the county Constanta until just south of the Danube delta; we visited some old open oak forests and ajacent grasslands near Babadag & in the foothills of the Macin Mountains (<500m) and we visited the Danube delta so to make it easy I will divide the pictures in 3 parts: (1) plains, (2) oak forests & (3) Danube delta.


Papilio machaon, some hilltops near Enisala held nice amounts of hilltopping males of this species

Colias erate
A rather common species near the coast. Most of them were clearly lemon yellow (like these two, a fresh and an abraded male). Interestingly up to 20% of them (rough estimate) however held a variable amount of an orangeish tinge, fainter than in C. croceus. I caught one real orange one as well, to check in the hand, and that male looked phenotypically just like W-European C. croceus (including a clear androconial patch on the hindwing). There is some discussion whether these orangeish individuals are hybrids/intergrades between C. erate & C. croceus with some "non-believers" saying all can be explained by interspecific variation of both species (and indeed, on the Azores for example you have the whole range of C. croceus from pure orange to lemon yellow without any C. erate being in the next 5000km). The fact however that both species share COI mitochondrial DNA suggests that there is or has been at least some introgression between these two taxa. Also the amount of orangeish colored C. erate seems to be considerably higher in the western part of the distribution where the distribution overlaps with C. croceus, again suggesting hybridisation at some level. Hybridisation is not such a rare event in butterflies and has been described in a lot of species (up to 16% of European butterfly fauna is known to hybridise to some extent!), sometimes anekdotical but in some cases also on a higher, stable level (fe. in Papilio machaon/P. hospiton).

Lycaena dispar in coastal marshes

Oak forests near Babadag and the foothills of Macin Mountains

Habitat at the foothills of Macin mountains

Argynnis pandora, male

Hipparchia syriaca
A check of the Juliën organ in the field with 20x magnification showed this male having 7 broad & stout lamella confirming the ID.

Kirinia roxelana
A pleasant surprise, easily 10+ spotted on one morning in Macin Mountains. In the literature I checked beforehand this species was mentioned for N Dobrogea but the mentions were very few and only for Babadag forest. Aparently recently it appears to be a bit more widespread in Babadag & Macin Mountains.

Lycaena thersamon, underside

Lycaena thersamon, female

Lycaena thersamon, male
Widespread in Macin mountains with easily 50+ seen

Polyommatus admetus, female, recently only known from N Dobrogea in Romania

Danube delta

We mainly visited this area for birdwatching & to enjoy the amazing landscapes but still got some nice butterfly sightings.

Apatura metis 
I have been searching for this species a few years ago without succes (see this older post) but this time I could see and enjoy the real deal!

Heteropterus morpheus
Remarkably fresh early august, I only know the Dutch populations and these start to fly end of june - early july. In the Danube delta, more or less at sea level, it seems like this species is on the wing a month later than in the Netherlands!

More on the eastern Carpathians in a next thread.