Academic literature on the topic 'Hipparchia alcyone'

This paper discusses three problems concerning the Woodland Grayling, Hipparchia fagi Scopoli, 1763, with respect to the identity and application of the junior name Papilio hermione Linnaeus, 1764. In 1977, the late Otakar Kudrna designated a specimen of the Rock Grayling, Hipparchia alcyone [Denis & Schiffermüller], 1775, to become the lectotype of Papilio hermione – as a result of which hermione supplanted alcyone as the senior epithet for this species. Because P. hermione is the nominal type species of Hipparchia Fabricius, 1807, Kudrna’s action rendered this a genus based on a misidentified species. Third, while a majority of lepidopterists have ignored Kudrna’s action and continue to apply the name H. alcyone to the Rock Grayling, and still regard P. hermione as a junior subjective synonym of H. fagi, the formal nomenclature for the Rock Grayling has become unstable because a large minority have nonetheless accepted Kudrna’s lectotype designation and all that follows from it. It is demonstrated here that no syntypes of Papilio hermione (or Papilio fagi) have survived; consequently, Kudrna’s lectotype designation for P. hermione is invalid. By designation of a single specimen of the Woodland Grayling as neotype for both P. fagi and P. hermione, the two names are rendered objectively synonymous, thereby restoring stability to the species name for the Rock Grayling (as Hipparchia alcyone), and to the application of Papilio hermione (= Hipparchia fagi) as nominal type species of the generic name Hipparchia.

In 1977, Otakar Kudrna (*1939-†2021, obituary see Balletto and Leigheb, 2021) published his “Revision of the Genus Hipparchia”, where he classified all the known species and forms of this genus according to characters of wings, androconia, male genital armatures and further subjective criteria. Until today, Kudrna’s study is considered as the guideline of systematics of the genus Hipparchia. He selected there a lectotype specimen of a Rock Grayling male in the Linnaean collection. “Hipparchia hermione Linnaeus, 1764” is therefore, at the moment, the technically correct name to identify the species. The “International Commission of Zoological Nomenclature” (ICZN) has nothing to add at this point; it only comments on cases submitted to it in the Bulletin of Zoological Nomenclature. Within the meaning of the present study and in accordance with Verity (1913), this damaged specimen without abdomen represents the same species as Ignaz Schiffermüller – allegedly the only author of the Vienna directory (see Kudrna and B., 2005, p. 5) – has described under H. alcyone from the Vienna region by referring to a coloured copper engraving published by Rösel von Rosenhof (1755). Kudrna’s “Revision” became the starting point of an extended scientific research activity during my free time containing, as a matter of priority, the examination of problem cases of systematics by checking the preimaginal characters of many rearing series from different sites. It turned out that a number of classifications proposed by Kudrna (1977) had to be reassessed as soon as characters of the pre-imaginal stages were available. The most complex case I have verified concerned the third European Grayling species which Leraut (1990) introduced under the name of Hipparchia genava (Fruhstorfer, 1908). Kudrna (1977) failed to separate this species from H. alcyone (D. & S., 1775). Throughout his life, he never agreed with Leraut’s opinion. In recent years, Kudrna had hoped that genetic examinations would make redundant every rearing attempt by amateur lepidopterists and furnish the proof that his opinion was the correct one. Since he never undertook any rearing experiments, his systematics were based only on prepared imagines being housed in museum collections. He saw himself as a person with the competency to decide within a few minutes upon complex questions of taxonomy and ignored completely the assessments of others. For verifying the effective rank of H. genava, I had to examine also the two most closely related species: Hipparchia fagi and H. alcyone by rearing them all ex ovo with material from several widely spaced sites. Already on finalising my rearing work of this group, it became apparent that Leraut (1990) had been on the absolutely right path by accepting a third Grayling species, within this group. For the first time, I published the results of my rearing attempts between 2002 and 2006 in several articles in the quarterly bulletin Linneana Belgica and I readily provided information on this case to interested colleagues. Over time only, I realized that the Rock Grayling I knew from the volume on butterflies (Diurna) by Forster and Wohlfahrt (1955) as from the guides by Higgins and Riley (1970-84) under the name of H. alcyone had become H. hermione, because of the lectotype designation by Kudrna (1977). A stony path was in front of me to substantiate the factual correctness of the view taken by Leraut (1990). Complex clarifications by Peter Russell furnished well-founded arguments on the complex scientific issue why the use of the name “hermione Linnaeus, 1764” should be rejected for the Rock Grayling previously known as H. alcyone.

25 years ago, my friend Guido Volpe (Castel Volturno, Campania) asked me for help to identify the Grayling species occurring in his region. In summer 1998, I started several rearing experiments with batches of ova from Italy to check their identity with the help of the larval stages. Guido Volpe procured the females for oviposition on different sites in the mountains of Lazio, Campania and Calabria and sent the obtained ova by mail to Switzerland. The question arose soon, which one of the two smaller Grayling species was indigenous to this region, Hipparchia genava or alcyone. In his study “Contribution à l’étude des Satyrinae de France”, Patrice Leraut (1990) elevated to species rank the taxon genava Fruhstorfer (1908) occurring in the Valais, claiming that its distribution area extends across the whole Italy to eastern Sicily. I hoped to confirm this assertion, being based mainly on the rods of the Jullien organ, also in the larval stages. For this purpose, I was forced to examine rearing series from the entire area inhabited by populations of the two smaller Grayling species. During the period 1998 to 2006, I investigated them with series from 8 European countries in total (see Jutzeler et al. 2005, map p. 152), by rearing them always from the ovum. In fact, the two smaller species alcyone and genava produced caterpillars of different appearance. The imagines that emerged under rearing conditions were retained to search them for characters being typical of each of the two species. The results from all these rearing experiments were recorded in the study on H. fagi and genava (Volpe and Jutzeler, 2001 – genava still named there alcyone) and in 3 studies being dedicated to alcyone and genava (Jutzeler et al., 2002, 2005, 2006). Already in those days, I didn’t use the name of “hermione Linnaeus, 1764” favoured by Kudrna (1977) to rename H. alcyone D. & S. In my eyes, it referred to two different species. Moreover, Kudrna’s lectotype designation did not conform to the rules of ICZN according to Higgins and Riley (1978) and I had detected during my investigations that this name was applied to designate the taxon fagi since 1775 until the mid-20th century and that it was often used in reality to designate unknowingly also the similar species genava. The synonymy of Papilio hermione Linnaeus, 1764 with P. fagi Scopoli, 1763 has been irrevocably established recently by Russell and Vane-Wright (2022). In the first part of “Doubts about the validity”, I posited the recognition of the specific rank of Hipparchia genava. Since the beginning of my research program on the Grayling complex, I tackled this question under various aspects. Of central importance was the determination of the characteristics of wings using reared imagines of the three Grayling species resulting from the numerous rearing series of Hipparchia alcyone, genava and fagi. The most important variations found among caterpillars and imagines are presented once again in this part of the study. Furthermore, the evaluation of preparations housed in the collections of the Zoological Museum Amsterdam (since 2011 housed in the Naturalis Biodiversity Centre Leiden) and other museums being figured on the 9 synoptical plates together with their genital armatures, was of primary importance for the present article. Thereby, the accuracy of Leraut’s (1990) diagnosis could be confirmed in the main, but with the limitation that the number of rods of the Jullien organ of alcyone and genava can also be beyond the range indicated by Leraut. Thus, the rods do not compellingly lead to a correct identification of every individual. As a new character of male genitalia, the strongly curved inferior edge of the male genital armature of H. alcyone should be noted. Accompanied by Peter Russell, I verified also Fruhstorfer’s (1908a) type material of H. genava housed at the British Museum of Natural History in London (BMNH). Contrary to all of the ever-voiced findings by Kudrna, all the individuals of the type series in London accorded perfectly with genava from the Valais using the wing characteristics. The variability of the wing design of H. genava (and fagi) is additionally illustrated with the figures reproduced from plate 73 of Verity’s 5th volume “Satyridae” (1953) of his work “Farfalle Diurne d’Italia”. The occurrence of Grayling species in some selected actual and historical distributional areas was furthermore checked by using material in collections. Accordingly, the Palatinate region was populated only by alcyone. Proofs that genava existed in this region do not seem to exist. Of the two small Grayling species in Alsace, Hipparchia genava still occurs in the Alsatian Jura (Haut-Rhin) whereas H. alcyone formerly populated, until the 1960s, some sites in the northern part of the Vosges (Bas-Rhin). In the Jura Mountains of Basel and Solothurn, only H. genava occurs, whereas alcyone and fagi are completely absent from those areas. In Liguria, H. fagi and genava are widespread whereas only 4 specimens representing alcyone could be detected from this region being collected in 1973 near Celle Ligure (Savona). I supported the inquiries in the field by Tristan Lafranchis and colleagues in the region of the contact areas of H. genava and alcyone in south-eastern France. Accordingly, Duponchel (1832-35) diagnosed correctly the occurrence of H. alcyone in the surroundings of Marseille. Individuals of H. alcyone from this region were described by Fruhstorfer as ssp. sogdiana, whereas Varin (1962) referred ssp. sogdiana erroneously to the “altitudinal form” corresponding with H. genava, considering the true alcyone from mount Faron near Toulon (Var) as “under-race faronica” of ssp. sogdiana (sensu genava).

The critically endangered grayling (Hipparchia hermione, syn.: alcyone) is one of the rapidly declining diurnal butterfly species which occupies only few remaining localities in the Czech Republic. Currently, its remaining local populations can be found in the central Povltavi area where they inhabit mostly sparse light oak forests with low cover of the herb layer. As a diploma thesis, the study was conducted during the season 2015. The populations of H. alcyone were researched around the Orlik water reservoir. The occurrence was confirmed at six localities out of which two had been unknown until then. Within four dense populations have been using the capture-recapture method evaluated the mobility and dispersal abilities of the species. The populations differed in their dispersal abilities. Various average long distances across localities and sexes were detected. The average long distances varied (males 142to300 m, females 78to261 m) across all locations. The flight probability were ascertained with two methods: the inverse power function (IPF) and the negative exponential function (NEF). The NEF method fitted better the flight probability at all localities. The interchanges of individuals between localities were noted only in a case of two closest populations. These one-way interchanges (three males and two females) were always directed from dense to smaller population. The adults of H. alcyone were typical by very low dispersions between separate populations, probably due to lower densities of populations and innapropriate structure of migration paths. Three overflights of males and two overflights of females to the neighbouring location were recorded. With respect to the size of local population and the structure of migration routes, the individuals expand with difficulties. In order to maintain and support habitats of new H. alcyone biotopes, the more open canopies and creation of a larger number of small clearings in the neighbourhood of the H. alcyone localities is necessary.

Il genere Hipparchia Fabricius è uno dei generi tassonomicamente più difficile dei satiridi paleartici. Esso comprende gruppi di taxa con caratteristiche esterne simili, ma con profonde differenze nei genitali maschili e negli androconi. Studi pregressi sul comportamento hanno evidenziato in alcune specie l’esistenza di un corteggiamento articolato per l'accoppiamento. La maggior parte delle volte, le sequenze di corteggiamento rilevate sono comparabili tra specie, ma possono presentare differenze nell’esecuzione e nell’ordine dei moduli comportamentali comuni; in altri casi, invece, sono caratterizzati da moduli esclusivi che presentano solo determinate specie. L’isolamento riproduttivo fra due specie è il risultato di un gran numero di meccanismi, più o meno indipendenti gli uni dagli altri, e l’importanza relativa dei vari fattori può cambiare da un gruppo di specie all’altro. La riproduzione tra specie animali, soprattutto simpatriche e strettamente affini, è comunemente impedita da una serie di fattori ecologici, comportamentali e citogenetici. Tra i meccanismi di isolamento riproduttivo precopula, quelli di tipo etologico costituiscono la classe più importante. Il funzionamento di tali meccanismi si basa essenzialmente sull’incompatibilità comportamentale dei segnali scambiati tra gli individui eterospecifici. In generale, in condizioni di allopatria , specie strettamente affini possono permettersi di utilizzare segnali di corteggiamento più generici (i.e., aspecifici), meno costosi e variabili, mentre in condizioni di simpatria la selezione naturale può favorire la diversificazione di atti di corteggiamento e di esibizioni sessuali articolate, mediate dalla scelta femminile, anche laddove non vi sia alcuna ibridazione eterospecifica. Questo lavoro di tesi ha l’obiettivo principale di approfondire lo studio del corteggiamento nelle due specie criptiche, Hipparchia fagi ed H. alcyone, al fine di comprendere quali stimoli, visivo-comportamentali ed, in misura preliminare, chimici, caratterizzano i due taxa ed assicurano l’accoppiamento tra individui conspecifici. Queste due specie sono simpatriche e sintopiche ad altitudini comprese tra 800 - 1000 m; gli adulti mostrano un periodo di volo sovrapposto (luglio -agosto) e frequentano habitat simili tra bosco ed aree ecotonali prato-bosco; inoltre sono molto simili nel pattern alare e di difficile determinazione, senza un attento esame delle strutture genitali maschili e femminili. Lo studio del comportamento sessuale in queste due specie è stato eseguito in natura su popolazioni localizzate nei pressi della Pineta (930 m) situata tra le località di Vallemare e Borbona (RI) nel Lazio. Alle osservazioni in natura sono state affiancate osservazioni all’interno di una voliera sia su individui selvatici, che su individui di allevamento, provenienti dalla stessa area di studio, presso i Laboratori di Ecologia Sperimentale ed Acquacoltura dell’Università di Roma Tor Vergata. Questo studio ha rivelato nelle due specie lo stesso sistema nuziale, una poliginia con monandria delle femmine, ed un pattern comportamentale sessuale molto simile: il maschio attende la femmina, posato sui tronchi d’albero come sulle pareti della voliera; al passaggio della femmina, prende volo e la insegue fino alla posa; segue quindi un rituale articolato dalla struttura temporale e dai meccanismi di interazione maschiofemmina analoghi nelle due specie. La sequenza di corteggiamento appare costituita dai sei moduli comportamentali: Fanning, Circling, Bowing, Antenna orientation, Copulation attempt e Clasping, eseguiti nell’ordine dato in entrambe le specie. Il maschio può ottenere l’accoppiamento dopo aver eseguito una sola volta la sequenza di corteggiamento, ma nella maggior parte dei casi, ciò avviene dopo la successione di più sequenze (o di frammenti di essa), probabilmente in risposta alla disponibilità ad accoppiarsi della femmina. Il comportamento della femmina sembra infatti influenzare l’esecuzione delle azioni del maschio, determinando la frammentazione della sequenza e la ripetizione dei singoli moduli o dell’intera sequenza. Nei rituali più articolati, il maschio tende a ricominciare la sequenza di corteggiamento dal Bowing o da moduli precedenti e, solo dopo un apparente consenso della femmina, può procedere nella sequenza fino alla copula. Le due specie, inoltre, mostrano un corteggiamento molto simile nelle tempistiche dei sei moduli comportamentali componenti la sequenza. In entrambe, infatti, i moduli iniziali (Fanning e Circling) e finali (Copulation attempt e Clasping) mostrano durate comparabili e sono eseguiti in media una sola volta nella sequenza di corteggiamento. I moduli centrali della sequenza (Bowing ed Antenna orientation) sono invece replicati molte volte, sempre in associazione, e caratterizzati da durate molto brevi. L’unico carattere comportamentale che distingue le due specie è la frequenza con cui il maschio effettua il Bowing: in H. alcyone sempre maggiore di quella misurata in H. fagi. Questo modulo sembra essere la fase cruciale del corteggiamento, sia per il numero di repliche sia per la sua perfomance. Nel Bowing infatti il maschio afferra e trattiene le antenne della femmina tra le ali anteriori facendole scivolare sulle sue squame androconiali, secretrici di feromoni. Moduli fortemente stereotipati, come il Bowing, probabilmente incrementano l’intensità del segnale ormonale rilasciato, assumendo un ruolo importante nella stimolazione della partner ad acconsentire alla copula. Queste osservazioni suggeriscono il coinvolgimento di segnali chimici, oltre che visivo-comportamentali, nell’interazione tra i sessi. Le indagini preliminari sul rilascio dei feromoni mediante l’applicazione del Naso elettronico hanno rilevato un simile pattern chimico dei composti volatili emessi dai maschi nelle due specie. Specie affini possono avere una simile composizione chimica del bouquet dei composti rilasciati e differire per la concentrazione relativa degli stessi composti. Non è facile determinare in quale misura le differenze rilevate nelle specie esaminate possano contribuire all’isolamento riproduttivo tra i due taxa, ma differenze quantitative comportamentali sembrano essere sufficienti ad impedire una perfetta sincronizzazione della disponibilità all’accoppiamento in molte specie animali e, soprattutto, in specie affini.<br>Species differences in characters involved in reproductive isolation are candidates for factors that caused speciation. It is not easy to understand whether the species differences that we can detect have been actually involved in speciation, because most species pairs have been accumulating genetic differences since the speciation process was completed often developing multiple reproductive isolating mechanisms. In Hipparchia species and in many taxa of Satyrine butterflies, both sexes have evolved complex genitalia that can mechanically impede the copulation between different species while males have scent located in patches or dispersed on their wings, probably pheromone releaser. Surprisingly, in spite of a growing amount of information on morphology and rearing methods of several species, only the works of Tinbergen on H. semele and some recent research specifically deal with mating behaviour of the genus Hipparchia. Courtship is an important character linked to fitness and can evolve in response to female mate choice, to ensure sex identity and quality of a potential mate. It can also be a major pre-mating isolating mechanism in butterflies and many other species. The aim of this work was to study in detail the reproductive behaviour of two sibling species, Hipparchia fagi and H. alcyone in order to understand which species -specific, visual, behavioural or chemical stimuli are involved in courtship. These two species are sympatric and syntopic at 800 - 1000 m of altitude; adult flight periods overlap in July and August and take place in similar habitats; the two butterflies have very similar wing patterns and sometimes it is very difficult to recognise the species without the exam of male and female genitalia structures. The behavioural study was conducted daily, both in the wild and in captivity, during the reproductive seasons in some areas near Vallemare (1,100 m a.s.l.) in the Central Appennines (Rieti, Italy). The observations were carried out with captive and reared individuals inside a large outdoor “flight cage” built in the same locality and in the garden of the Laboratori di Ecologia Sperimentale ed Acquacoltura (University of Roma Tor Vergata). In this study, different aspects of the butterfly reproductive behaviour are described. First, the ethogram and the sequential structure of courtship are showed; second, various parameters of each courtship step are measured; third, the results in H. fagi are compared with the behaviour of the species H. alcyone; fourth, the mating system of each species is described. Moreover, the male chemical patterns were detected by the application of the Electronic Nose and the results in H. fagi were compared with those of H.alcyone. The study reveals general patterns of sexual behaviour in H. fagi and in H. alcyone, that are mainly identical in both species: perching strategy, flight pursuit and courtship. Likewise, the two species show a similar polygynous mating system, with female monandry. In both species, the results revealed a highly stereotypic courtship that consists of six steps (Fanning, Circling, Bowing, Antenna orientation, Copulation attempt e Clasping) leading to the copulation. Courtship most likely follows up to the end, once the male has started with the sequence. However, the development of the courtship seems to respond to the female behaviour, reflecting the complexity of the male -female interaction during the whole sequence. In both species courtship sequence seems to mostly be the outcome of the female’s reaction: her interruptions can produce fragmented sequences and, more interestingly, can induce the repetition of the sequence around a specific point, with the male persisting in courting the female. Male can mate after displaying the courtship sequence once, but mostly repeats the whole sequence or part of it more than once, restarting from Bowing or previous steps, likely waiting for female signals. The behavioural sequential analysis also showed that a male displays the initial and the final phases of the sequence only once, while he can repeat Bowing and Antenna orientation a lot of times in the same sequence. The comparison between H. fagi and H. alcyone showed quantitative interesting differences. The frequency of Bowing is the unique behavioural character that allows to tell apart the two study species , as H. alcyone shows a greater frequency than H. fagi. In satyrine butterflies, male androconia provide the necessary cue, probably of chemical nature, for a successful courtship, and the display of Bowing is likely of primary importance for the production of male pheromones. During Bowing the male repeatedly bashes or strokes the female’s antennae against the androconial scales on his forewing dorsal surface. Bowing could convey chemotactic information to the female, and its repetition within a sequence could represent a mechanical stimulation for her, with the secretion from androconial patches on the male’s forewings. This phase could be of great importance, as a persistent male could drive a female at first unreceptive to eventually mate. Scents and visual stimuli could have an essential role in mating communication: they could operate synergistically and help the female in her decision of acceptance or rejection of a specific male; they could function as sexspecific signals facilitating sexual identification during courtship. The detection of the butterfly smell by Electronic Nose revealed a similar male chemical pattern in both species. In insects and especially in butterflies, pheromones are chemical signals whose composition varies enormously between species and they typically comprise more than one active component. Frequently sympatric species show the same pheromone components, but in different combinations or ratios. In absence of experimental crossability tests between species, it is not possible to be investigate if different visual (i.e., different performance of a male action) or chemical stimuli would change the female response and impede crossmating. However, the behavioural differences observed in H. fagi and H. alcyone could act as important discordant cues, in case of encounters with heterospecific females, and could reflect the presence of different chemical cues (not yet identified) between the two species.