Introduction and general advices This part deals with the life histories and bionomics of all currently recognized Zygaena species (checklist vide 1/52–57), when these are known. For all information we have tried to provide an exact re­ference, while accepting that this might interrupt the flow of the discursive text and that we risk receiving criticism from the reader for such inconvenience. In addition to so much data based on our own field observations and cultures, authentic information received from colleagues has been included as well as every important reference that we could find in the published literature. The main goal of this part is to describe, demonstrate and where possible to analyse infraspecific diversity in a wide sense, not only what is mani­fested in the morphological structures of adults and pre-imaginal stages, but also in the spectrum of their specific and infraspecific habitats and ecological adaptations. The plates of infraspecific differentiation Each species chapter starts with a double page, the geographical differentiation (preferably the valid subspecies) figured by sets of three (or four) typical specimens in one line on the left page, so that the reader of the text on the right page can easily see what the described infraspecific characters figured on the left page. Often the phenotypic spectrum of a subspecies is wider than can be shown with a few specimens. We have reduced the selection to 'population typical specimens' accepting that aberrations - even regular aberrations - are often not represented. Additional information on subspecific descriptions and further figures of specimens of western Palaearctic Zygaena species are to be found in de Freina & Witt (2002). For an extented phenotypic comparison we recommend the virtual collection of the Museum Witt Munich (http://www.insecta-web.org/MWM/). Unless if mentioned otherwise all figured specimens are stored in the collection of the junior author (AH). In the legends to the plates with triplets of set speci­mens valid subspecies are cited in bold, framed by a cartouche with cast shadow. When specimens of synonymic taxa or of unclear status (valid or not ?) or of populations are figured, then the nominal taxon is cited as originally described. That does not necessarily mean that we agree with this position or even rectify/revise them here, but want give the reader a chance of forming his own opinion about such available, subjective taxa. Only very obvious invalid taxa are placed in synonymy (distinguishable by citation with 'syn.' and in normal italic instead of bold italic). Unnamed but slightly to mediumly distinct populations are occasionally highlighted as 'subsp. ?'; in no way does this imply that nomenclatoric naming is urgently needed, but shows that even distinct populations can remain unnamed. The selection on the plates allows the reader to get a quick overview about distinct (or not distinct) charac­ters and geographical distances can be seen on the distribution map. In this monograph taxono­my has taken a back seat and only some relevant, recent taxonomic changes are discussed. Data cited here (e.g. altitude) are original citations from the specimen pin-labels. When citing taxa in the legends of the pictures of landscapes and bio­topes the authors and dates of publication are omitted; this information is provided in the running text beside. The species introductions The text starts under the species heading with the reference to the original description of the taxon, as cited by the author but rationalised to a certain extent. This is followed by a short introductory section providing brief biographical details of the author or authors of the nominal taxon and the field workers who originally discovered the material, together with the derivation of the name when either is known. The geographical range, the vertical distribution and the most significant variation within subareals of each species are briefly described and all noteworthy subspecies are characterised and figured. Here each nominal taxon (normally subspecies) is fully cited and in bold when first mentioned, in the subsequent text and in the captions the authors(s) and the date of publication are omitted. The subspecific taxonomy is following Hofmann & Tremewan (1996) where further synonyma can be found. We have abstained from comprehensive descriptions and detailed discussions of subspecific characters. Only the most remarkable characters in note form are mentioned. Otherwise it would lead to endless paragraphs with subsequent debates about subspecific criteria, subjective decisions and transition zones. All of which is not the goal of this monograph, as these but can be found in standard works (Reiss in Seitz, 1930-1933; Naumann, Tarmann & Tremewan, 1999; de Freina & Witt, 2002). For every species a selection of pictures of characteristic landscapes and habitats are provided to give the reader the opportunity to imagine the species specific ecological plasticity. Here our aim and working question was "Could one gets the specie's name just from seeing these pictures of its natural habitats?" The distribution maps The specific distribution with subspecific type-localities of each species are shown on simple maps with few colours while exact sites are figured on separate maps with distinct coloured altitudinal zones. Based on georeferenced maps from U.S. Geological Supply we succeeded in transforming these data into maps of altitudinal zones for the whole palaearctic region. On such maps not only the distribution is exactly shown but also information on ecological limitations are provided. For every species a distribution map is generated which is based on authentic records that could be georeferenced and incorporated in ESRI ArcView and QGis projects (the latter an open source software). These maps in their present completeness could not be worked out only based on personal data. They are the product of intensive cooperation. Originally based on literature analysis (vide References, p. 1022–1039) and on a complete data gathering of the collection of A. Hofmann it was subsequently extended by targeted data from missing regions (especially United Kingdom) from the collection W. G. Tremewan and selected data from the Museum Witt (München), Zoologische Staatssammlung (München), The Na­tural History Museum (London), Musée d'Histoire Naturelle (Paris), Museum Alexander Koenig (Bonn) and Staatliches Museum für Naturkunde (Karlsruhe). These collections contain the famous collections of F. Daniel, F. Dujardin, K.-H. Wiegel, H. & G. Reiss (all in Munich) and C. M. Naumann (in Bonn). More­over, an essential extension was the cooperation with Laszlo Rakosy (Cluj, Romania), Igor Kostyuk (Kiev, Ukraine) and Tiroler Landesmuseum Ferdinandeum (Innsbruck). G. M. Tarmann has gathered data on Palaearctic Zygaenindae since 2005 with main focus on the Apennnine Peninsula, the Alps and the Balkans (s.l.). These data of which the full sets are stored in the biooffice (Innsbruck) perfectly completed the under-represented sets of eastern and south-eastern Europe. A very essential contribution for occurrences in France was provided by the 'Groupe d'Information de Recherche et d'Animation sur les Zygaenidae' (GIRAZ). Nevertheless a regionally more or less equal presentation of distribution dots was not possible as the availablity of easily incorporated data sets was not given for many areas. Although well investigated and with a large amount of data being stored in the collections for the Iberian Peninsula, central, northern and eastern Europe, continuous digital data were lacking and we contented ourselves with the incorporation of the most important cornerstone data sets, being aware that much more data was available in the collections. In contrast, for the more peripheral regions (e.g. North Africa, Iran, Afghanistan), which were much less well explored, the majority of collections and recorded data could here be incorporated. Every dot on the map is based on at least one veri­fied set of data. The red coloured, framed areas represent ranges of potential occurrences, i.e. within these areas further records are known or to be expected and in no way do such areas indicate that the species is omnipresent here. When dots remain outside the specific range in red it means that such records are either based on what are now most probably extinct populations (last records more than 50 years ago), or they are unclear or doubtful and therefore require confirmation. For some species with very spare records (e. g. Z. sedi, Z. nevadensis) or for some subareals with very few records, we understandably refrained from using framed ranges. Life history Under the heading ‘Life history’, the known ontogenetic stages (ovum, larva, pupa and cocoon) and the infraspecific variability of each individual species are described. The description of the egg and the terms used for their position (i.e. whether they are deposited horizontally or vertically), the position of the transparent pole within the egg and the formation of the egg batch follow Hofmann & Kia-Hofmann (2011: 39–41, fig. 5). It is obvious that when eggs or cocoons are found in the wild, their age is unknown. Therefore the duration of the egg phase and especially that of the cocoon phase is usually based on cultures reared under European conditions and these may vary in nature, depending on the ambient environmental temperatures. Based on a few observations in the wild the duration can be shortened by 3–4 days, especially in arid regions such as those found in Iran or Morocco. Whenever we cite data based on observations in the wild, this is emphasized. A list of plant species on which the larvae are known to feed is provided under the subheading ‘Larval host-plants’; however, it should be noted that records of plants erroneously cited as larval host-plants in the literature have been ignored. Under the heading ‘Bionomics’, observations on the ecology and ethology of the adult (and of the larva, when available) are described and, where necessary, opinions on genetic or geographical phenomena are expressed. Consistent characters that are present in the ova and larvae are not repeated in the description of each species provided below. For example, when first laid, the ovoid ovum usually has one pole that is transparent, depending on its deposition, but as the embryo develops and expands within the egg, the whitish to yellow coloration of the yolk extends throughout the egg; a day or so prior to hatching the coloration of the egg changes to dark grey or even brown. Such development is general and occurs in every species. The eggs of the majority of species are placed flat on the substrate in a horizontal position and have a characteristic depression that is caused by the female when prodding her papillae anales against the egg to provide it with stronger attachment to the substrate (Hofmann & Kia-Hofmann, 2011). In contrast, nine species are known to deposit their eggs in a vertical position, viz. Z. sengana, Z. escalerai, Z. formosa, Z. rosinae, Z. sedi, Z. olivieri, Z. haberhaueri, Z. chirazica and Z. naumanni. The full description of the larval development with details of every stage (L1–L3 or L4, diapause instar, L5–L7 or more) are omitted in order to prevent endless repetition. However, when extraordinary phenotypic changes occur during the larval phase, these are mentioned. Each larval description has been made from a dorsolateral aspect and includes the length of the fully-grown larva that is still feeding, measured when it is at rest. In almost all species the head and thoracic legs are black and, as a consequence, they are not mentioned in the descriptions unless they are coloured otherwise. It should be noted that nearly all larvae, from the first to the final instar, have two pairs of small dark shiny ‘spots’, one pair situated on the first thoracic segment, the second on the tenth abdominal segment; these are of unknown function and should not be confused with the black anterior and posterior dorsal pigment spots when these are present on each segment from the second thoracic to the eighth/ninth abdominal segment. The term ‘cocoon phase’ denotes the period from the completion of the cocoon to the emergence of the imago and includes the time during which the larva is a prepupa. When the measurements of the larvae and cocoons are based on less that 20 examples, the number used for descriptive purposes is provided in parentheses. Every attempt has been made to base the descriptions on living material; however, in a few cases a description is based on a transparency or transparencies and digital photographs provided by colleagues. When this has been done a reference has been provided, e.g. ‘C. M. Naumann, transparency archive’, otherwise the transparencies were photographed by the present authors. The transparency collection of the late C. M. Naumann is now in the possession of A. Hofmann. Bionomics Under ‘Bionomics’ we record our own observations, data received from colleagues and published information about habitat preferences, ecology, biology and behaviour, which could not be easily incorporated in one of the other sections. Discovered by Fred H. Brandt (1908–1994) and described by Hugo Reiss (1890–1974) who dedicated the species to the famous German lepidopterist Adalbert Seitz (1860–1938). Mainly represented by monomorphic populations, this Iranian species also exhibits some dimorphic populations (Hofmann & Tremewan, 2010: 123, 124; Keil, 2014: 156–157, figs 1–4). It is endemic to the Kuhhā-ye Zāgros, where its range extends from the Zardkuh region in the central part to Dasht-e Arzhan in the province of Fārs. No records are known further south or east and it is obviously absent in the Kuhhā-ye Qohrud. A single record from north of Qāderābād, 20.v.1995 (F. Carbonell) requires confirmation. The species is strictly allopatric with Z. tamara. The great majority of populations are monomorphic and consist of moths that have red collare, and red abdominal cingulum, yellow forewing spots and red hindwings (Z. seitzi seitzi Reiss, 1938b; ③); within such monomorphic populations, the phenotypic variation is slight. However, there are a few locations known where there are dimorphic populations, a situation comparable to that found in the closely related Z. tamara. Zygaena seitzi tenhageni Hofmann & Tremewan, 2003, (②) from the vicinity of Semirom (Esfahān) is such a polychromatic taxon that has two main morphs, red and yellow, together with an orange morph that occurs rarely; it is represented by populations in which the dominant red phenotype can comprise ca 90%. A mixed copula found in the wild (Hofmann & Tremewan, 2003: 19, pl. 3, fig. 18) and ab ovo cultures have verified that both morphs belong to one biospecies. Moreover, the Mendelian gene­tics of the red and yellow morphs confirm this (Tremewan, 2006c: 177–178). Another dimorphic population comprising ca 80% yellow and 20% red individuals was recently discovered at Kuh-e Kalār (Lordegān NW.) (Hofmann & Tremewan, 2010: 124). A successful ab ovo culture reared in 2009–2010 from eggs oviposited by a yellow female taken in the wild produced both morphs, so there can be no doubt that in this locality both morphs are also conspecific (A. Hofmann, pers. obs.). The most northern and most western populations exhibit reduced spot 6 and orange or yellow instead of red collare (Z. seitzi mesopotamica Hofmann & Naderi, 2017; ①). At Dezh Kord (Fārs), a pure yellow male, i.e. with yellow fore- and hindwings and with yellow collare and a yellow abdominal cingulum, was found, a morph that is probably controlled by a single recessive allele, as known for the flava forms occurring in Z. filipendulae, Z. trifolii and Z. carniolica. Life history Ovum. Ivory to pale yellow; deposited on the underside of a leaf of the larval host-plant, usually near the central rib (Hofmann, 2000a: pl. 1, fig. 2; Hofmann & Kia-Hofmann, 2011: 58–59, figs 70, 76; Keil, 2014: 158–159, fig. 5), occasionally in small batches consisting of less than 20 eggs, but often comprising 40–70 eggs arranged in a single or even more frequently in a double layer. Of three batches from Z. seitzi seitzi found at Dasht-e Arzhan (Fārs, Iran), one was laid in three layers, the two lower layers regular, the uppermost partly irregular (125 eggs), the second was regular and deposited in a double layer (84 eggs) and the third was in a regular, single layer consisting of 34 eggs (A. Hofmann, pers. obs.). Egg batches of Z. seitzi tenhageni are often bigger than those of Z. seitzi seitzi (Hofmann & Tremewan, 2003: 19, pl. 3, fig. 13; Hofmann & Kia-Hofmann, 2011: 58, fig. 70) and are deposited in several layers consisting of one to three or four hundred eggs. During and after the main appearance of the moths, egg batches can often be found in numbers on the underside of Eryngium leaves. In captivity, an egg batch of F2 (Ry × Ry) consisted of six layers. Immediately after separating from a copula, another female of Z. seitzi tenhageni (Semirom, Esfahān) deposited a large batch consisting of more than 400 eggs arranged in seven layers. On average the first layer consists of ca 30 eggs; the largest batch of eggs found in the wild consisted of four layers and comprised more than 200 eggs (description based on eggs observed in the field at Semirom, Dasht-e Arzhan and Dorāhun). Egg parasitoids were noted in an egg batch from Dorāhun; the coloration of all the eggs in one of two dozen batches found in the wild changed to black. Egg-phase 11 days (Dezh Kord); 12 days (Semirom and Pol-e Dokhtar); 11 days were also noted for Z. seitzi tenhageni in captivity in the autumn (A. Hofmann, pers. obs.). Larva. Aposematic/predominantly cryptic. Length: 17–22 mm (n = 6). Body pale green, of a similar colour to the underside of an Eryngium leaf and becoming greener towards the end of the final instar, verrucae sometimes pale grey or dove-grey, a variably broad, primrose-yellow mediodorsal line, venter and abdominal and anal prolegs yellowish green, an irregularly shaped, black anterior dorsal spot present on second thoracic to ninth abdominal segment, those in anterior and posterior regions of body sometimes fragmented, posterior black dorsal spots vestigial or absent, on second thoracic to eighth abdominal segment when present, dorso-subdorsal spots indistinct, pale yellow, present on second thoracic to eighth abdominal segment, those on second thoracic and eighth abdominal segment vestigial, peritremes of spiracles black, white setae fairly long (2 mm), black setae shorter, bristly, on thoracic region longer, pointing caudad (nominotypical subspecies – Fārs: Dasht-e Arzhan (Hofmann, 2000a: pl. 1, figs 3, 4)). Fully grown larvae of Z. seitzi seitzi and Z. seitzi tenhageni sometimes remain inactive for three weeks, hardly ever moving or feeding, before spinning their cocoons and pupating, behaviour that is typical for Z. tamara. On the Kuh-e Cheri, five fully-grown larvae were found in the early afternoon sitting inside Astragalus bushes close to Eryngium plants, while none was observed on the larval host-plant. Larvae of Z. seitzi tenhageni have the body yellowish green with verrucae tinged yellow or darker, almost canary yellow, light bluish green intersegmentally, anterior black dorsal spots elongate triangular, that on ninth segment smaller, posterior black dorsal spots smaller, linear, vestigial on thoracic region, or even absent, venter and abdominal and anal prolegs greenish yellow (Esfahān: vic. Semirom, Kūh-e Behrouz, 2600 m) (Hofmann & Tremewan, 2003: 19, pl. 3, figs 10–12; Keil, 2014: 158–159, figs 6–8). Wiltshire (1952: 176, fig. 5) was the first to describe and illustrate the larva from the vicinity of Shirāz, an example of which he found feeding on a plant that was determined as Cachrys ferulacea (L.) Calestani. Larval host-plants. Cachrys ferulacea (L.) Calestani (Prangos ferulacea (L.) Lindley), Eryngium billardieri Delar. (Hofmann & Tremewan, 1996: 41). The main host-plant is undoubtedly Eryngium billardieri; at many sites in the wild, cocoons, larvae or egg-batches were found on this plant. In captivity the larvae will feed on various species of Eryngium, on Pastinaca sativa L. and on Seseli libanotis (L.) W. Koch. Pupa and cocoon. Pupa: length 12 mm (n = 1); rather stout, head, thorax, wings and appendages shiny, brown, or mahogany brown to dark brown, wings lighter, abdomen light green, caudal segments tinged brown, anterior dorsal spots showing through the cuticle as dull, greyish marks. Pupal phase: 15–17 days. Cocoon: length 16–21 mm, breadth 6–9 mm, height 6 mm (n = 10); subfusiform, surface irregularly ribbed or wrinkled, with two prominent ribs dorsolaterally and smaller ribs below, with a weak sheen, white to off-white and cream-white, seldom brownish, occasionally irregularly marked with pale yellowish (Hofmann, 2000a: 188, pl. 1, fig. 5; Hofmann & Tremewan, 2003: 19, pl. 3, figs 16, 17). Spun on the underside (seldom upperside) of a leaf of the larval host-plant, either on an upper leaf or amongst the basal leaves. Cocoon phase: 15–28 days, the majority around 20 days in captivity; e.g. 21 and 22 days for a female and male from Dasht-e Arzhan; but only 15–18 days were noted for moths of Z. seitzi tenhageni, those from Dezh Kord (Zāgros) emerged in Iran after 15–16 days. Zygaena seitzi mesopotamica needed 17 days (A. Hofmann, pers. obs.). Wiltshire (1952: 176) also recorded 16 days. The pupal exuviae remain protruding from the cocoon after eclosion of the imago. Bionomics Zygaena seitzi is restricted to the Zāgros range where it is found in the southern Iranian provinces of Fārs, Kohgiluyeh-va-Buyer Ahmad, Çahārmahāl-va-Bakhtiyāri and Esfahān. Of special interest is its occurrence in the Zardkuh region (Çahārmahāl-va-Bakhtiyāri) at a site that is within the distributional range of Z. nocturna. It inhabits dry, sparsely or profusely vegetated steppe and hillsides, or areas adjacent to or even in flowery clearings in open oak forest or shrub, preferably at altitudes between 1,900–2,900 m. Its southernmost occurrences in the province of Fārs are in clearings in light Quercus brantiii Lindl. woodland and shrub. Near Dorāhun (1,850 m) it occurs in semi-ruderal locations and uncultivated areas within fruit orchards. In the Zardkuh region it reaches the thorny ‘Polster’ zone (3,100 m) where it is sympatric with Z. fredi and Z. bakhtiyari. All checks for night activity in its habitats near Samsami, Dezh Kord, Semirom and Dasht-e Arzhan were negative, showing a clear separation of niches between Z. seitzi and Z. nocturna. Moreover, in captivity males that were collected in the wild behaved like typical diurnal Lepidoptera, i.e. after sunset they sat without showing any activity (A. Hofmann, pers. obs.). Nevertheless males were observed visiting flowers of Eryngium sp. and Echinops sp. and searching for females until 18.00 h. At Dasht-e Arzhan it was noted that the phase of highest activity occurred between 16–18 h; later in the evening the moths were inactive and preferably sat on the underside of Eryngium leaves. Dead moths were regularly noted in spiders’ webs at this site and a Mantis sp. was observed sitting inside an Astragalus polster waiting to catch moths, which it finally succeeded in doing. Subsequent checks of Astragalus polsters have shown that often a praying mantis sat inside, indicating a targeted strategy of predation. In lower biotopes (1,900–2,200 m) in the province of Fārs the adults have already emerged by the end of May, but in those situated at the highest altitudes near Samsami (2,800–3,100 m), the flight period begins nearly two months later (end of June, first decade of July). At Dorāhun, egg batches, L1, L2 and L3 larvae as well as freshly emerged moths were observed on the same date (13.vii.2005), thus indicating an extended flight period of 3–4 weeks (A. Hofmann, pers. obs.). The adults often spend much of their time resting under the basal leaves of the larval host-plant, Eryngium billardieri; however, they can be very wary and will often fly swiftly away when approached. The underside of the leaves of the larval host-plant is one of the preferred locations for copulae, where they are well concealed and protected, and only the antennae are occasionally protruded. Copulae were observed from early afternoon (14.30 h) onward. Both sexes fly strongly, the flight period ceasing at 17.30–18.30 h. Individuals of the yellow morph of the dichromatic population on Kuh-e Kalār were observed to have a preference for nectaring at the yellow-flowered Phlomis olivieri Benth.; from a distance they were difficult to spot when sitting on the yellow flowers. The males are readily attracted to Sesiidae artificial pheromones and approach the bait as near as one metre, most of them then flying away. It should be noted that, in contrast to Z. nocturna, Z. seitzi is strictly diurnal and no moths were ever attracted to actinic lights that were worked by the authors on several occasions in habitats where the species was active by day. Moreover, at Dorāhun (Borujen S.), both species were found to be syntopic (Hofmann & Tremewan, 2003: 12). However, the syntopic situation in this locality requires further investigation. Zygaena seitzi is usually accompanied by other burnet moth species, e.g. Z. escalerai, Z. chirazica and Z. cambysea. It is of interest to note that it has not been found syntopically with Z. turkmenica, which occurs in the same region, while it appears to be strictly allopatric with Z. tamara. On the Gardaneh-ye Cheri (Samsami vic., Zardkuh region), Z. seitzi seitzi is syntopic with the closely related Z. fredi escaleraiana Holik, 1958, while the dimorphic Z. seitzi tenhageni at Semirom is found together with Z. fredi syntopica. While the moths have only been observed to nectar infrequently at flowers, they occasionally visit those of the yellow-flowered Phlomis olivieri Benth. (Semirom, Kuh-e Kalār); furthermore, Stachys inflata Benth., Echinops sp., Cirsium sp. and a pinkish Fabaceae (Dasht-e Arzhan) have also been noted as nectar plants. At the type-locality (Semirom) of Z. seitzi tenhageni, the males were observed mate-searching or nectaring from 16.00–18.30 h; only rarely are they seen flying during the morning and early afternoon and then they are usually found resting on herbage. The adults rest amongst the leaves of Eryngium billardieri Delar. or within Astragalus bushes where they are afforded some protection by the spines of these plants. In its treeless, not strongly overgrazed, herb-rich habitats, Z. seitzi tenhageni is syntopic with Z. mirzayansi, Z. fredi syntopica, Z. haematina and Z. chirazica. However, in this locality the flight period of Z. chirazica and Z. fredi overlaps with that of Z. seitzi for only a few days, the last-mentioned species occurring from the beginning of June to the beginning of July. Moreover, Z. seitzi tenhageni is concentrated in an area at the base of a very steep scree slope on which Z. chirazica, Z. haematina and Z. fredi are found. After successfully copulating, females of Z. seitzi tenhageni sometimes sit for several days before starting to oviposit, behaviour that is also known for Z. tamara (A. Hofmann, pers. obs.). Discovered by Günter Ebert (1935–) and Heinz Falkner (1927–) in 1972 southeast of Yasuj; the former, when describing it, named it after its predominantly nocturnal behaviour that is unusual for a Zygaena species. Zygaena nocturna is endemic to the central Kuhhā-ye Zāgros (Iran) where it is more northerly and westerly distributed than the closely related Z. seitzi. Compared with the latter species it is larger and exhibits greater infraspecific variation. The nominotypical Z. nocturna nocturna Ebert, 1974 (③) (Keil, 2014: 160–161, fig. 1), occurs in the southern part of the range of the species (Esfahān, Fereydūn Shahr region; Çahārmahāl-va-Bakhtiyāri; Kohgiluyeh-va-Buyer Ahmad). The most northern populations (Z. nocturna eberti Hofmann, 2005, (①) with predominantly connected spots 5+6 (Keil, 2014: 160–161, fig. 2) that are reminiscent of those of Z. cuvieri) occur in the Oshtorān-Kuh (Lorestān). Individuals with yellow and yellowish hindwings are found in the Darreh-ye Kamarān where the moths are syntopic with Z. cambysea kamarana, a subspecies represented by red, orange and yellow individuals. In a series of some dozen specimens from this site, one male exhibits a three-segmented, red abdominal belt, a character that is otherwise found in the manlia-group only in Z. tamara. Populations occurring further south around the Gardaneh Meymand, Yasuj and Ardekān have spots 5 and 6 of the forewing separated and are thus reminiscent of Z. seitzi. However, the strictly nocturnal activity of Z. nocturna and its large size are unmistakeable diagnostic characters. The most easterly populations near Eqlid (Z. nocturna meinekei Hofmann & Tremewan, 2003; ②) are characterized by having the forewing spots varying from yellow to pale red; this subspecies comes into close geographical contact with Z. kermanensis whose nearest records are from Surmaq and Abadeh, two localities that are only ca 25 km apart as the crow flies. Life history Ovum. Pale yellow; in the wild often deposited on the underside of a leaf of the larval host-plant, usually in a large pyramidal batch consisting of four to five layers ( 150 eggs); in captivity occasionally deposited in smaller batches of one to two layers consisting of from 6–10 up to 50–120 eggs (Fārs). Egg parasitoids were noted in an egg batch from Chenār Mahmoudi; all eggs in a batch found in the wild changed their coloration to black. The egg development, from oviposition to hatching of the larvae, usually lasts from 9–10 days in late spring (May) (Hofmann & Kia-Hofmann, 2011: 65), but can be extended under ‘normal’ conditions for up to 12 days. Larva. Predominantly or entirely cryptic. Length: 19–21 mm. Body light green or yellowish green, venter light green, abdominal and anal prolegs yellowish, black dorsal, subdorsal and lateral spots absent, or black anterior dorsal spots vestigial, present on second thoracic to eighth abdominal segment, dorso-subdorsal spots inconspicuous, pale greenish yellow, present on second thoracic to eighth abdominal segment, those on second thoracic and eighth abdominal segment vestigial, peritremes of spiracles black, white setae fairly long (2 mm) (Hofmann, 2000a: pl. 1, figs 8, 9; Hofmann & Tremewan, 2003: 15, pl. 1, figs 11–14). The black anterior dorsal spots are always present throughout the different instars, from L1 to the penultimate instars, but become vestigial or absent in the final instar (Hofmann & Tremewan, 2003: 15, pl. 1, figs 11–15; Keil, 2014: 162–163, figs 3–5). Larvae of Z. nocturna meinekei Hofmann & Tremewan, 2003, from Eqlid (Esfahān) have the black anterior dorsal spots stronger and the black posterior spots are present, thus providing them with a weak aposematic pattern, while those larvae from the most north-westerly population (Oshtorān-Kuh, Lorestān) have these elements reduced or even absent, leaving only the greenish ground coloration and thus making them strongly cryptic. The larvae of populations geographically situated in between (Chenār Mahmoudi, Çahārmahāl-va-Bakhtiyāri) are intermediate in pattern. Larval host-plant. Eryngium billardieri Delar.; in captivity the larvae will feed readily on various Eryngium species. Keil (2014: 162) records Echinophora cinerea (Boiss.) Hedge & Lamond as a larval host-plant, but this probably refers to Ferulago carduchorum Boiss. & Hausskn., a plant that has not been confirmed as a foodplant by the present authors. Pupa and cocoon. Pupa: length 14–17 mm (n = 4); thorax, wings and appendages dark brown to black, thorax sometimes mottled lighter, abdomen green to yellowish green, lighter on venter, sometimes marked brownish laterally, anterior dorsal spots discernible under the surface of the cuticle, caudal segments brownish. Pupal phase: 18–19 days. Cocoon: length 19–21 mm, breadth 7–8 mm, height 7–8 mm (n = 4); subfusiform, surface weakly and irregularly ribbed, mat or with a weak sheen, white to pale beige (Hofmann, 2000a: pl. 1, fig. 10). Spun on the upper side or, more frequently, on the underside of a leaf of the larval host-plant, or deep inside large Astragalus bushes (Hofmann & Tremewan, 2003: 15, pl. 1, fig. 18). Cocoon phase: usually 18–22 or even 24 days; for Z. nocturna meinekei 14 (female) and 18 days (male) were noted, while a duration of 17 days was observed for individuals from Chenār Mahmoudi and Gardaneh Meymand (Z. nocturna nocturna); 17–22 days for Z. nocturna eberti from Oshtorān-Kuh (A. Hofmann, pers. obs.). The pupal exuviae remain protruding from the cocoon after eclosion of the imago. A phenomenon that is almost typical for Z. tamara has also been observed for Z. nocturna nocturna and Z. nocturna meinekei. Fully grown larvae sometimes sit relatively inactively for two or more weeks without feeding before starting to spin their cocoons (A. Hofmann, pers. obs.). Bionomics The most southerly record is the type-locality north-west of Ardekān (Fārs) where the species is parapatric with the closely related Z. seitzi. Moreover, both species are parapatric on the north-east side of the Kuh-e Denā range. The species inhabits sparsely vegetated, dry slopes and treeless mountain steppe as well as open oak forest at altitudes of 1,950–3,050 m; this vertical distribution is almost identical to that of Z. seitzi (1,850–3,100 m), but the majority of Z. nocturna sites are at higher altitudes than those of Z. seitzi. It is known to be syntopic with this species in a habitat near Dorāhun (Çahārmahāl-va-Bakhtiyāri) (Hofmann & Tremewan, 2003: 12), where Z. seitzi was observed flying during the afternoon. As a male of Z. nocturna had been captured flying strongly at 17.30 h nearby, two actinic light traps were located in the habitat after sunset, one in the flat area where Z. seitzi had been taken in the afternoon, the other on a ridge some 60 m or so above but out of sight of the lower trap. Only two worn males of Z. nocturna came to the latter site, but in the Z. seitzi habitat, 15 males of Z. nocturna came to the light between 21.00–23.00 h. It is of interest to note that the males of Z. nocturna in this locality are smaller than those from other populations. In many of its habitats, Z. nocturna is often syntopic with Z. escalerai, Z. cambysea and Z. haematina, but the distribution of the last-mentioned species usually extends to higher altitudes; occasionally it is also syntopic with Z. fredi and Z. chirazica. It is noteworthy that in the Oshtorān-Kuh region it is sympatric with Z. tamara while no syntopic occurrences with Z. turkmenica or any other nocturnal or semi-nocturnal Zygaena species are known. Zygaena nocturna is predominantly nocturnal, the males coming to actinic light from one hour after sunset to 23.30 h. In the Darreh-ye Kamarān at 2,700–3,050 m on 10 July 2003, the flight period began just after sunset (21.00 h) but it was very short and ceased again at 22.00 h; 35 males were noted at three light traps. A full moon was shining so that it is possible that this affected the activity of the males. Females are not attracted to light and no activity of the males was observed after this time, in spite of lights being used in its habitats to trap nocturnal Lepidoptera. When individuals come to a light source they very quickly settle down on the sheet. Very occasionally, males can be observed in the early morning nectaring at the blue flowers of Echinops or at a white-flowered species of Scabiosa. During the day both sexes rest preferably under the basal leaves of the larval host-plant with just the tips of their antennae protruding from beneath. Occasionally moths were found under the leaves of flowering plants such as Phlomis species. While they are rather lethargic in the morning, during the heat of the afternoon they are wary and will quickly fly away when one approaches their resting site. Artificial Sesiidae pheromones that have been used to attract the males of Z. seitzi by day have been placed near to where a light source was used at night; in contrast to Z. seitzi, the males of Z. nocturna showed no interest in the pheromones whereas two males were attracted to a light near where no pheromones had been placed (Hofmann, 2000a: 190). A predawn flight has been recorded, as several individuals were observed flying just before sunrise (06.00 h), indicating another period of activity for possible nectaring, mate searching or copulation. This might explain the occasional observation of a solitary male nectaring later in the morning (ca 09.30 h). In captivity, several males and one female were put into a cage at 12.30 h to observe any possible activity that might occur during the day, which was sunny with a temperature of 25°C and perfect for copula trials; moths of other Zygaena species were flying actively and nectaring from late mor­ning onward, while the males of Z. nocturna only showed some activity for 10–15 minutes after 12.00 h and after that remained inactive for the rest of the afternoon and the evening until sunset. At 22.00 h a copula was found, while on the same day Z. tamara copulated at 17.00 h. It should be noted that it is incorrect to regard Z. nocturna as completely inactive during the day! Females of nominotypical Z. nocturna and Z. nocturna meinekei have been observed in captivity releasing sexual pheromones after sunset (Hofmann & Tremewan, 2003: 15, pl. 1, fig. 10; Hofmann & Kia-Hofmann, 2010: 85, fig. 5). Moreover, several copulae were obtained in culture experiments undertaken in Spain and in Germany. With the exception of one, all of these copulae took place after sunset and in the dark, from 21.00–23.00 h (Hofmann & Tremewan, 2003: 10), and lasted until the next morning when the individuals of each pair separated, as they do in the diurnal Zygaena species. In contrast to the diurnal species, the transfer of the spermatozoa must have occurred during the hours of darkness (Hofmann & Kia-Hofmann, 2010: 89). Like the females of all other Zygaena species, those of Z. nocturna lay their eggs exclusively during the warmth of the day, usually during the afternoon. Moreover, as with the majority of females of Z. tamara, some females of Z. nocturna were not observed to oviposit on the same day that the copula had separated – in fact they sat for three or more days before ovipositing began. In an open Quercus forest at Gardaneh Meymand, prediapause larvae have been found in numbers on preferably weak plants of Eryngium billardieri. Characteristic feeding marks were noted on the upper side of the leaves, but during the day all larvae were sitting on the underside of the leaves. They very quickly fall down with the slightest disturbance. Zygaena kermanensis was originally described as a subspecies of Z. manlia by Walter Gerald Tremewan (1931–2016) who named it after the province of Kermān (Iran) where the original series of specimens was taken. Zygaena kermanensis is a relatively uniform species, but exhibits geographical variation in the size of the red spots that also vary in the presence/absence, or in the intensity of the lighter, whitish scales, especially in the populations occurring in the vicinity of Qāderābād, Neyriz and Hasanābād. Some individuals even exhibit a vestigial white edging to the forewing spots (Hofmann, 2000b: 228, fig. 11), reminiscent of species that belong to the subgenus Agrumenia. Such characters are lacking in specimens from Kermān and Yazd (Hofmann, 2000b: 230, figs 18, 19); the taxonomic position of the strongly isolated populations from the Shir-Kuh region (Yazd) remains unclear. The species is endemic to Iran, where it is restricted to the eremic mountain ranges east of the Kuhhā-ye Zāgros, from between Eqlid and Qāderābād to south of the Kermān region; there it inhabits semi-desert biotopes at altitudes of 1,900–2,770 m. Surprisingly, it has not been recorded from further east, from Sistān-va-Baluchestān, but it is found in the vicinity of Dārāb (Fig. 128) and in the isolated Khabr-Kuh (Fig. 127) ca 120 km south of Bāft (28°54′15″ N 56°24′42″ E, 2,370–2,500 m, 13.v.2005, A. Hofmann); both sites belong to the most southerly known locations for any Zygaena species. Populations from the eastern foothills of the Zāgros (Z. kermanensis qashqai Tremewan, 1775; ①) are variable and characterized by orange coloration of spots and hindwings and a broad hindwing border. The syntopic occurrence of Z. kermanensis at Qadr Abad Pass (described as subsp. askarii Tremewan, 1975; ②) (slightly smaller than subsp. qashqai with more pinkish coloration and often with traces of whitish surrounding the forewing spots; Keil, 2014: 164–165, figs 1–3; 167, fig. 5) with Z. turkmenica isfahanica and with Z. fredi near Eqlid, Dārāb and Estabanat shows quite clearly that neither can be conspecific with Z. kermanensis. The nominotypical Z. kermanensis kermanensis Tremewan, 1975, (③) which is widely distributed in the mountainous regions around Kermān (Khabr-Kuh, Kuh-e Madvar, Kuh-e Hezar, Kuh-e Segoch, Kuh-e Masahim, Gardaneh Khorāsāni, Kotale Khāne Sorkh), is characterised by its large size, less pinkish spots with only traces of lighter surroundings and disconnected spots 5 and 6 (Keil, 2014: 164–165, fig. 4). Populations on the Kuh-e Segoch exhibit a slight tendency toward melanism (spot 6 reduced, broad hindwing border). Life history Ovum. Pale yellow or yellowish cream, changing to orange-brown and finally brown before hatching; deposited in small batches of 6–40 eggs, each batch consisting of a single layer (Fārs, Gardaneh-ye Kowli Kosh) (Tremewan, 1975: 237; Keil, 2014: 165, fig. 1; 166), on the underside of a basal leaf of the host-plant, the latter apparently always in the rosette stage. It remains unclear whether less than two dozen eggs in one particular batch are not an artefact because of disturbance of the female during ovipositing. In captivity the first batches always consist of more than 50 eggs. Hofmann (2000b: 228, fig. 3; 231) observed eggs that were laid in two (Gardaneh-ye Khorāsāni) or three (Kotale Khāne Sorkh) layers consisting of more than 60 eggs. Larva. Aposematic. Length: 20–22 mm (n = 3). Body varying from light grey or dove-grey to grey, a narrow, whitish mediodorsal line, a subquadrate or subtriangular, black anterior dorsal spot present on second thoracic to ninth abdominal segment, that on ninth abdominal segment smaller, black posterior dorsal spots smaller, linear, present on second thoracic to ninth abdominal segment, dorso-subdorsal spots pale yellow, present on second thoracic to eighth abdominal segment, venter and abdominal and anal prolegs grey, dorsal and subdorsal verrucae dark grey, setae comparatively long, giving the larva a hairy appearance, white, some of those in thoracic region black, setal rings black, peritremes of spiracles black (Fārs: Gardaneh-ye Kowli Kosh, Z. kermanensis qashqai). Larvae of Z. kermanensis askarii have the body light grey, venter grey or dirty grey, abdominal and anal prolegs cream, the black anterior dorsal spots subtriangular, those on second thoracic and ninth abdominal segments smaller, verrucae light grey, dorso-subdorsal spots bright yellow, present on second thoracic to eighth abdominal segment, that on eighth abdominal segment slightly smaller, setae short, white (Fārs: Gardaneh-ye Qāder Ăbād). Larvae from Gardaneh-ye Hasanābād (Fārs) have the dorsum, verrucae and the area anterior to each dorso-subdorsal spot sometimes tinged with greenish yellow. Larvae from the province of Kermān (Gardaneh-ye Khorāsāni and Kotale Khāne Sorkh – Z. kermanensis kermanensis) are darker with a broad, whitish mediodorsal line, the yellow dorso-subdorsal spots are more conspicuous, abdominal and ventral prolegs, intersegmental regions and dorsum off-white to light grey; black anterior dorsal spots triangular, black posterior dorsal spots small, linear (Hofmann, 2000b: 228, figs 5–8; Keil, 2014: 166–167, fig. 6). Larvae from Khabr Kuh (Kermān) exhibit a more white or whitish-grey venter with the black spots being slightly smaller, thus giving the larvae a lighter phenotype. Larvae in all cultures (e.g. from Khabr Kuh, Qāderābād, Neyriz) fractionized and some diapaused several times, up to 4 years, and in the last diapause period they were fairly large in size (L12D) and often more than 12 mm in length. Even these larvae moulted two times in the instars after diapause before pupating (A. Hofmann, pers. obs.). Larval host-plant. Eryngium billardieri Delar. (E. nigromontanum Boiss. & Buhse); in captivity the species has been reared on E. campestre L., E. giganteum Bieb., E. planum L. and E. variifolium Coss. (Hofmann & Tremewan, 1996: 41). Keil (2014: 166) records Echinophora cinerea (Boiss.) Hedge & Lamond as a larval host-plant, but this probably refers to Ferulago carduchorum Boiss. & Hausskn., a plant that has not been confirmed as a foodplant by the present authors. Pupa and cocoon. Pupa: shiny, head, thorax, wings and appendages brown or blackish brown, abdomen lighter. Cocoon: length 15–22 mm, breadth 7–10 mm, height 7–8 mm (n = 8), subfusiform, broad at base, parchment-like surface irregularly wrinkled, with a weak sheen, varying from white or cream to light brown (Keil, 2014: 166–167, fig. 7). Usually spun concealed amongst plant litter or in the soil, but occasionally exposed and attached to a stem of the host-plant or other vegetation. Cocoon phase: often 19–24 days, relatively long, usually one or two days longer than that of Z. turkmenica when reared at the same time. The pupal exuviae remain protruding from the cocoon after eclosion of the imago. A detailed account of the early stages and rearing this species ab ovo is provided by Tremewan (1975: 237–239) under ‘Z. manlia qashqai’, and by Hofmann (2000b: 231) under ‘Z. rubricollis kermanensis’. Bionomics Inhabits plain and hilly Artemisia steppe, dry rivers embankments and mountain slopes that are mostly grazed by sheep and goats and where the larval host-plant often grows in great abundance, at elevations of ca 1,900–2,770 m. The flight period is from the middle of May to the beginning of June. At Sir-Kuh (Yazd) the species occurs in a habitat characterized by Artemisia bushes and Stipa grass (Hofmann, 2000b: 228, fig. 2). The flight of the adult is strong, fast and direct; in comparison, that of Z. turkmenica isfahanica is slow and buzzing, a good character for distinguishing the two species in the field. In contrast, the larva of Z. kermanensis is sluggish compared with that of Z. turkmenica, which is very active and will quickly hide under the leaves of the host-plant in order to avoid bright sunshine! The moths visit the flowers of a scabious and those of an Acantholimon sp., but very often they were observed sitting concealed and inactive on the underside of leaves of the host-plant or amongst dry vegetation during the day (Hofmann, 2000b: 228, fig. 11). At Khabr-Kuh the moths were common in mid May 2005 and immediately took flight when disturbed, but not a single individual was observed nectaring between 12.00–15.00 h. From 15.00 h onward, several males were observed flying to and targeting Eryngium bushes, even when no females were sitting inside. At the same time of the day the first copulae were found (A. Hofmann, pers. obs.). Although closely related to Z. turkmenica, Z. aisha and Z. nocturna, all species that exhibit nocturnal behaviour, Z. kermanensis is strictly diurnal and not a single male has ever been observed at light traps in spite of several experiments that were undertaken in an attempt to attract them at Qāderābād, Hasanābād and in the Kermān region (Gardaneh-ye Khorāsāni, Kotale Khāne Sorkh, Kuh-e Segoch). In many habitats, Z. kermanensis is the only burnet moth species to be found. However, in the vicinity of Sirjān it can be found flying syntopically and synchronously with Z. rosinae, while at Qāderābād, Z. kermanensis askarii and Z. turkmenica isfahanica are syntopic. In the Kermān region, Z. kermanensis kermanensis seems to be parapatric with Z. ginnereissi, Z. aisha and Z. fredi, as no syntopy has been observed. From the vicinity of Estabanat (Shirāz ESE.), W. Eckweiler collected a small series of burnet moths consisting of Z. fredi and Z. kermanensis. First described as a subspecies of Zygaena manlia by Hugo Reiss (1890–1974), who named the species after the Turkmen or Turkmenistan, the region from where the original type series was collected; the taxon was subsequently recognized as a valid biospecies by Hofmann & Tremewan (2010: 125). However, species delimitation still remains critical. Zygaena turkmenica is actually one of the biggest burnet moths known, as set specimens from Damghan (Semnān, Iran) can have a wingspan of more than 40 mm when measured from apex to apex of the forewings. The exact distributional range of this species is still unclear because the reference of several records to Z. manlia or Z. cuvieri, and even to Z. turkmenica itself, remains undetermined. Populations from western Afghanistan (Prov. Shindand, Alaqadari, Fārsi, Dehkhak, 10–20.vii.1977, M. Casimir), from eastern Iran (Prov. Khorasān) close to the Afghan border (Farimān, Birjand, Kuh-e Mirza Arab) to the Kuh-e Sorkh, the Iranian Kopet Dagh region, Quchān, Dargaz, from Golestān (Shāh-Kuh region, Golestān National Park) and from northern parts of the province of Semnān (Gardaneh Khoshyeylagh, Shahrud E.) are all referable to the nominotypical subspecies (Z. turkmenica turkmenica Reiss, 1933) (③) (Keil, 2014: 168–170, figs 1, 2), which was described from Turkmenistan (Kopet Dagh, Jablonowka). They are characterised by the smaller, mostly isolated spot-pairs (1+2, 3+4, 5+6), with spot 6 often reduced and not connected to spot 5 in the upper part, and the more translucent, cold red coloration. Adjacent populations from the semi-desert areas around (and within) the great inner Iranian deserts (Dasht-e Kavir, Dasht-e Lut) can still be referred to this taxon even though they are larger in size with enlarged spots. Those from the central Alborz (Damāvand region) and from the vicinity of Karaj (Kuh-e Jaroo) have a warmer red coloration and lead to a complex of populations consisting of individuals intermediate with Z. cuvieri as well as populations from Zanjān and Hamadān with different frequencies of phenotypes (cf. Z. cuvieri piti Hofmann, 2000). Due to these variable or even dimorphic populations, taxonomic problems arise because their distributional range is very close to or even parapatric with the type-locality of Z. cuvieri. A similar situation exists in the central Zāgros range and adjacent areas (Nahāvand, Oshtorān-Kuh, Aligudarz, Khonsār, Afous, Sibak, Dāmaneh) where the populations also exhibit characters in both the adult and preimaginal stages which may indicate hybrid populations/belts between Z. turkmenica and Z. cuvieri (Hofmann & Tremewan, 2010: 125). Zygaena turkmenica isfahanica Tremewan, 1975, (②) was described from the vicinity of Meymeh (province of Esfahān, Iran). Populations of the Kuhhā-ye Qohrud (Kashān), northern Fārs and Esfahān are referable to this taxon, which is characterised by the large forewing spots, especially spots 4 and 6, and the warm red coloration (Keil, 2014: 168–169, fig. 3 [aberrant morph]). While there is obviously a belt of hybridisation in the central Zāgros and western Alborz, Z. turkmenica and Z. cuvieri are very distinct in some areas, being sympatric in eastern Turkey (Van and Hakkâri regions) (Z. turkmenica pjotri Hofmann, 1983; ①) where, north-east of Hakkâri (2,000 m), they were observed in numbers by day and were absolutely syntopic and synchronous (Hofmann, 1983: 116, 117). Zygaena turkmenica pjotri is pure ‘turkmenica-like’, without intermediate individuals, with colder red coloration, typically spot 6 detached from 5 in the upper part and with spot 4 smaller than that in Z. turkmenica isfahanica. The reddest forms are found in the Zangezur Mountains of Armenia, in the vicinity of Zaritap where the spots are enlarged and confluent forms are not infrequent. Zygaena excellens Reiss, 1940 (④), was described from a single female specimen from central Afghanistan (Badghis/Ghowr, Firoz-koh, 2,800–3,000 m). The poor information available does not allow us to make a final evaluation of the validity of this description; its great similarity possibly suggests (with reservations) that it is a synonym of the nominotypical subspecies of Z. turkmenica. Life history Ovum. Greenish white or pale green; deposited in a regularly shaped batch, consisting of a double or triple layer, or several layers, on the underside of a basal leaf of the host-plant when the latter is in the rosette stage (Tremewan, 1975: 243, pl. 6, fig. 6; Hofmann & Kia-Hofmann, 2011: 58, figs 71, 72; Keil, 2014: 170–171, fig. 4 [cuvieri?]). Three large batches of eggs that were found in this position at Meimeh on 10.vi.2009 were pyramidal and consisted of 5–7 layers each of which comprised more than 150 eggs. Published under ‘Z. manlia pjotri’, an egg phase of 9 days was recorded (Hofmann, 1983: 116). Larva. Aposematic. Length 16–22 mm (n = 3). Body dark bluish grey or bluish black, inter-segmental areas bluish grey, a narrow, whitish grey mediodorsal line, abdominal and anal prolegs yellowish to brownish yellow, an inconspicuous, black anterior dorsal spot present from second thoracic to ninth abdominal segment, black posterior dorsal spots even more inconspicuous, present on second thoracic to ninth abdominal segment, a conspicuous, yellow dorso-subdorsal spot present on second thoracic to eighth abdominal segment, peritremes of spiracles black, short setae black, longer setae white (Esfahān: Meymeh) (Tremewan, 1975: 244, pl. 6, figs 7, 8). Also figured by Keil (2014: 170–171, figs 5–7; 173, figs 8–10 [hybrid populations with Z. cuvieri]). Even more contrasting, with darker coloration, larger dorsal spots and a narrow mediodorsal line, are the larvae from Tshuli in the Kopet Dagh (Turkmenistan; C. M. Naumann, transparency archive). Larvae from Van (Turkey) have the body very dark bluish grey with the black spots enlarged, giving the larva a very dark, almost black appearance; the prominent dorso-subdorsal spots are bright dark yellow and contrast strongly with the surrounding coloration (Hofmann & Tremewan, 2003: 17, pl. 2, figs 8, 9). Larvae from Hakkâri (Turkey) have the ground colour dirty white, with the black spots greatly enlarged to leave a whitish mediodorsal band irrorate with black (C. M. Naumann, transparency archive). Larval host-plants. Eryngium billardieri Delar. (E. nigromontanum Boiss. & Buhse), Ferula latisecta Rech. f. & Aell. (F. undulata M. Pimen. & Baran.), Ferula sp. (Kuznetsov, 1960: 19); in captivity the species has been reared on Eryngium campestre L., E. maritimum L. and E. variifolium Coss. (Hofmann & Tremewan, 1996: 43), the larvae also accepting Seseli libanotis (L.) Koch. Quite extraordinary is a record by R. Trusch & T. Keil (pers. comm.) who, in 2007, found several larvae feeding (!) on Artemisia cf. herba-alba Asso (Figs 191, 192) north-east of Birjand (Hofmann & Tremewan, 2010: 125; Keil, 2014: 172). Pupa and cocoon. Pupa: length 15–16 mm (n = 3); shiny, dark or blackish brown. Cocoon: length 19–23 mm, breadth 9–12 mm, height 6–8 mm (n = 10); subfusiform, broad at base, parchment-like surface irregularly wrinkled, with a weak sheen, varying from white to cream (Esfahān: Meimeh) (Tremewan, 1975: 244–245, pl. 6, fig. 9); the same description applies to those from Turkmenistan (Kopet Dagh). Spun concealed amongst vegetation on or under the surface of the ground, or near the midrib on the underside of a leaf of the larval host-plant. At Meimeh and Ashān, several cocoons were observed in different years spun on the underside of Eryngium leaves rather than amongst the surrounding dry vegetation, while one cocoon was spun in an Artemisia bush (A. Hofmann, pers. obs.). Cocoon phase: 14–21 days (Iran), for Z. turkmenica isfahanica 15 days were counted; 15–17 days were noted for cultures from Armenia and Sibak (Iran) (A. Hofmann, pers. obs.); Kokanova (2000: 18; 2003: 81) recorded 10–12 days. The pupal exuviae are detached from, or remain protruding from, the cocoon after eclosion of the imago. A good character for separating the cocoons of Z. turkmenica from those of Z. cuvieri in areas (Iran, Turkey) where the two species are sympatric is that the former species produces a white or cream cocoon, whereas the latter produces a yellow cocoon. A detailed account of the early stages and rearing this species ab ovo is provided by Tremewan (1975: 243–245), under the name ‘Z. excellens Reiss, 1940’. Bionomics Inhabits Artemisia and Stipa steppe and mountain slopes at elevations of 500–2,800 m, often with Phlomis spp. and Astragalus cushions; the main occurrence in its centre of distribution (Iran) seems to be between 1,400–2,100 m. In Turkmenistan it has been recorded at 500–600 m in Artemisia steppe where its larval host-plant is possibly an Eryngium sp. (G. M. Tarmann, pers. comm.). However, Kuznetsov (1960: 19) and Kokanova (2000: 18; 2003: 80) found larvae feeding on a Ferula sp. in the western Kopet Dagh (Igdejik Gorge and Chuli Gorge respectively). Both sexes of Z. turkmenica can be observed by day flying actively and nectaring at flowers, but at night the males are attracted to light and sometimes come from considerable distances to the light source (e.g. several observations at the camping site at Gevaş (Van, Turkey) (P. Kautt, pers. obs.). More than 40 males, mostly worn, were attracted in one night to a light trap near Sibak (Fereydun Shahr, Iran). Near Ashān (Dāmaneh N., Iran), 36 males came to the light trap within one hour between 21.30 and 22.30 h. Two copulae were found next morning close to the light. In the mountains south-west of Kāshān (Esfahān), two dozen individuals came to actinic light from 21.45–23.30 h, but the following morning, in spite of intensive searching, none were found up to 13.00 h at which time we left the habitat. At the type-locality near Meymeh (Esfahān) males of Z. turkmenica isfahanica came to the light trap between 21.00–22.00 h with no further observation after 22.00 h (A. & K. Hofmann, pers. obs.). It was interesting to note that many males displayed courtship behaviour and were seen to buzz around in circles on the ground with the valvae extended, suggesting that in this population copulation takes place at night. In captivity copulation did not start before dusk, mostly after 21.30 h; the couples usually remain in this position until the following morning, but occasionally they can even be observed until the early afternoon. Males were inactive during the day, but started their activities after 18.00 h and began flying in indoors after 20.00 h. In nature during the day and in the late afternoon and evening, adults can be observed at rest on the underside of Eryngium leaves with only the antennae protruding from beneath; they also rest on the flower heads of Stipa and on other plants. They preferably nectar at the pinkish flowers of Acantholimon sp., Centaurea candolleana Boiss. and Stachys inflata Benth., but were also observed at the yellow flowers of Serratula latifolia Boiss., a Melilotus sp., several Cephalaria species and a purple-flowering, non-thorny Astragalus sp. Some moths were found sitting upside down in a Stachys sp. In the Hakkâri region (SE. Turkey) moths were strongly attracted by the bluish flowers of a Salvia sp. Exposed plants were mostly visited by several individuals, often accompanied by Z. cambysea, the latter much shyer with a stronger flight. The flight of the moth is slow and heavy compared with that of Z. kermanensis; moreover, as mentioned above, the behaviour of the larvae differs between the two species. Prediapause larvae were observed in the wild in the vicinity of Meymeh (Iran) sitting mostly on the underside of the Eryngium leaves that had characteristic feeding marks. It was remarkable that when disturbed they immediately fell to the ground (A. Hofmann, pers. obs.). Zygaena cacuminum was first discovered in 1873 by Hugo Theodor Christoph (1831–1894) and was not recorded for more than 100 years until Bernard Jean Mollet (1945–) rediscovered it in the type-locality on 17–18 July 1998. The name is derived from the Latin ‘cacumen’, meaning the ‘peak or utmost point’. Endemic to Iran, this monotypic species is only known from two high mountain ranges in the eastern Alborz (Shāh-Kuh, Kuh-e Shāhvār) where it inhabits altitudes above the tree line, from 2,900 m to over 3,800 m (Kuh-e Shāhvār-e Bozorg). From the closely related Z. speciosa that is restricted to the central Alborz, it is easily distinguished by the form and position of spots 3+4, which do not form a line but are connected to form an angle. There is a gap of 250 km between the ranges of both species. The species and its biology has been figured by Hofmann (2000d: 342–347, figs 50–59, 1–14) and Keil (2014: 174–175, figs 1–3). Life history Ovum. Comparatively large, whitish cream; deposited in a double-layered batch on the underside of a stone (Hofmann, 2000d: pl. 3, figs 1, 2; Hofmann & Kia-Hofmann, 2011: 52, fig. 38), behaviour that is characteristic of high-mountain species; the depression on the upper side, which is found in nearly all Zygaena eggs when freshly laid, is particularly prominent. An egg batch found in the wild consisted of two layers, the basal with 20, the upper with 13 eggs. Egg phase 9–10 days. Keil (2014: 174, 177, fig. 4) also figures the egg batches. Larva. Aposematic and disruptively cryptic. Length: 23 mm. Black; body bluish-greyish to dirty whitish tinged with pink, venter and abdominal and anal prolegs concolorous with ground colour, black anterior and posterior dorsal spots and black anterior and posterior subdorsal spots of each segment coalescing to form a very large blotch that is less strongly pigmented on the dorsal and subdorsal verrucae (the fused anterior and posterior dorsal spots are so expanded dorsally that they almost contact those on the opposite side of the body, thus leaving a thin, zigzag mediodorsal line of ground colour (because of the intersegmental divisions), dorso-subdorsal spots extraordinarily large, ovoid, bright yellow, present on second thoracic to eighth abdominal segment, that on eighth segment smaller, peritremes of spiracles black, setae black (short, bristly), or white (longer) (Hofmann, 2000d: pl. 3, figs 11, 12; Keil, 2014: 174, 177, figs 5, 6) (Māzandarān/Semnān: Reshteh-ye Shāh-Kuh). Larval host-plant. Trachydium eriocarpum Bornm. & Gauba (Hofmann, 2000d: pl. 3, figs 12, 13); in captivity larvae accepted Levisticum officinale W. D. J. Koch and fed also occasionally on Eryngium planum L. (A. Hofmann, pers. obs.; Keil, 2014: 176). A fully-grown larva was found high up on the stem of the host-plant that had the inflorescenses characteristically eaten (A. Hofmann, pers. obs.). Pupa and cocoon. Pupa: mid brown (described from exuviae). Cocoon: length 20–22 mm, breadth 11 mm, height 10 mm (n = 2); subfusiform, surface finely sculptured, dirty brownish white to white with a pronounced sheen; the cocoons are well adapted to the substrate and often of the same coloration as the stones on which they are spun (Hofmann, 2000d: pl. 3, fig. 10). The pupal exuviae usually remain protruding from the cocoon after eclosion of the imago. Hofmann (2000d) provides a detailed account of the biology and ecology of Z. cacuminum. Bionomics The original discovery of Z. cacuminum by Christoph and its rediscovery by Bernard Mollet are described by Naumann (2000b: 206) and Hofmann (2000d: 342) respectively. The species is endemic to northern Iran where, in the eastern Alborz, it is restricted to the northern side of the Shāh-Kuh range, the habitat being situated above the village of Shāh-Kuh-e-Pāiin at an elevation of 2,850–3,000 m. Based on information provided by the Iranian botanist Hossein Akhani, B. Mollet and A. Hofmann succeeded in finding the species (initially recognised as unilocal) in 2005 at two new localities in the Kuh-e Shāhvār range ca 25 km ENE. from the type-locality, as the crow flies (Hofmann & Kia-Hofmann, 2008: 50). On 25–26 July, a few specimens were collected at Kuh-e Shāhvār-e Kuçek on south-facing slopes close to and on the ridge at altitudes of 3,300–3,500 m. On 27 July a single male was discovered on Kuh-e Shāhvār-e Bozorg in a saddle at 3,720 m close to a snowfield. Here in 2008, B. Mollet (pers. comm.) found the species up to 3,850 m. In both localities it was flying together with a few individuals of Z. ecki and Z. carniolica. As a strong wind was blowing for three daysweiterlesen