HEMARIS TITYUS (Linnaeus, 1758)

GB: Narrow-bordered Bee Hawkmoth, F: Sphinx Bombyliforme, D: Skabiosenschwärmer, RU: Shmelevidka skabiozovaya, S: Svävflugelika Dagsvärmare, NL: Glasvlerkje, CZ: Dlouhozobka zimolezová, H: pöszörszender, PL: Fruczak trutniowiec, FI: Ruusuruohokiitäjä, Ruusuruohonpäiväkiitäjä, IT: sfinge della scabiosa.

Sphinx tityus Linnaeus, 1758, Syst. Nat. (Edn 10) 1: 493.

Type locality: Unspecified [Europe].

(Taxonomic notes. (i) Derzhavets (1984) considered 'subsp.' alaiana Rothschild & Jordan, 1903, from the Alai range (Alayskiy Khrebat) of the western Tian Shan (Tajikistan), to be merely a form of Hemaris tityus, at best a subspecies. However, Eitschberger & Churkin (2002) quite rightly treat Hemaris alaiana as a distinct species. Comparison of both male and female genitalia indicate that Hemaris alaiana has evolved from an isolated population of the eastern Palaearctic Hemaris radians (Walker, 1856).

(ii) Recent DNA barcoding studies indicate that Hemaris galunae (Eitschberger, Müller & Kravchenko, 2005) from the Levant is a good species closely related to Hemaris tityus. It appears to be the eastern remnant of a species which once occurred right across a cooler, greener and wetter North Africa during the last ice age. The western remnant is Hemaris aksana (Le Cerf, 1923) from the mountains of Morocco.)


A plate from Kirby (1907). Part of a plate from Curtis (1823-1840).

BIOGEOGRAPHICAL AFFILIATION

Holarctic; Palaearctic (both eastern and western). Pleistocene refuge: Polycentric -- Adriatomediterranean and Pontomediterranean subsections of the Mediterranean refuge, and the Caspian, Iranian, Turkestan, Nepali and Manchurian refugia.


ADULT DESCRIPTION AND VARIATION

Wingspan: 40--50mm. Distinguished from Hemaris fuciformis (Linnaeus, 1758) by the narrow band of scaling along the outer margin of each wing and the presence of an undivided forewing cell. Although some confusion may arise with freshly-emerged examples which have their entire wings covered with loose, grey scales, these are lost after the first flight, after which identification is made easier by a near absence of variation.

In the male genitalia, uncus and gnathos as in Hemaris fuciformis. Juxta rough with granules only apically, produced into a slender lobe. Valves asymmetrical; left valve broader than in Hemaris fuciformis, not spatulate, right valve narrowed in apical third. Left harpe represented by an obvious setose hump, evenly and broadly convex or flat; right harpe almost straight, apically clubbed and apically armed with spines. Aedeagus process longer than in Hemaris fuciformis, acutely pointed. In the female genitalia, antrum quite small, approximately square with rounded corners. Ductus bursae distinct, separating the antrum from the corpus bursae.


Pairing adults of Hemaris tityus, Hungary. Photo: © Szabolcs Safian. Adult of Hemaris tityus, Catalonia, Spain. Photo: © Tony Pittaway. Adult of Hemaris tityus (pre-flight), Catalonia, Spain. Photo: © Tony Pittaway.

ADULT BIOLOGY

Diurnal. An insect of meadows, roadside verges and woodland glades rich in flowers, especially on chalk and sand where Ajuga (bugle) thrives. In such localities, vast numbers can be found in some years but very few in others, though it is never totally absent. Where this species occurs, Scabiosa (scabious) is nearly always present; however, many areas rich in this plant are devoid of Hemaris tityus. Up to 2000m in the Alps (Vorbrodt & Müller-Rutz, 1911; Forster & Wohlfahrt, 1960), even higher (3300-3600m) in Kyrgyzstan (Eitschberger & Churkin, 2002).


Typical habitat of Hemaris tityus, Catalonia, Spain. Photo: © Ben Trott. Typical habitat of Hemaris tityus, Joutseno, SE Finland. Photo: © Jaakko Pohjoismäki.

Unlike most other sphingids, this species is active during the day, generally between 10.00 and 15.00 hours, when it can be seen flitting rapidly from flower to flower of its favourite plants -- Ajuga, Glechoma, Lychnis, Viscaria, Salvia, Pulmonaria and Knautia (Bergmann, 1953). Although mimicking a bumble bee in coloration, its flight is much more rapid and agile. Its skill is superbly demonstrated during courtship when pairs chase each other low over the ground as well as spiralling upwards like a whirlwind. Copulation generally follows, with pairs remaining in copula for up to two hours.


Pairing adults of Hemaris tityus, Joutseno, SE Finland. Photo: © Jaakko Pohjoismäki.

FLIGHT-TIME

Mainly univoltine in northern Europe, mid-May to mid-June (sometimes early July) with an irregular partial second brood in August. Fully bivoltine farther south, during late April or early May, and again in August. Around Saratov, Russia, this species flies in June and July (Kumakov, 1977).

Late June (29-30.vi) in Kyrgyzstan at 3300-3600m (Eitschberger & Churkin, 2002), but mid May (13.v) in western Mongolia at 2000m (Yakovlev & Doroshkin, 2005). July only in the Altai Mountains (Kondratiev coll., BM(NH)). However, in the mountains of Central Asia the flight-time varies with altitude.


EARLY STAGES

OVUM: Small (1.1 x 1.0mm), almost spherical, with a small depression on top; pale glossy green. Laid singly on the underside of a leaf of the hostplant, with up to six per plant or group of plants not uncommon.


Egg of Hemaris tityus, Catalonia, Spain. Photo: © Ben Trott.

LARVA: Full-fed 50mm.


First instar larva of Hemaris tityus, Catalonia, Spain. Photo: © Ben Trott. Second instar larva of Hemaris tityus, Catalonia, Spain. Photo: © Ben Trott. Third instar larva of Hemaris tityus, Catalonia, Spain. Photo: © Ben Trott. Fourth instar larva of Hemaris tityus, Catalonia, Spain. Photo: © Ben Trott.

The newly-hatched, white larva is about 3mm long and covered with small black tubercles bearing forked, black hairs. The caudal horn is small with two black hairs at its apex. With each moult and feeding the body and tubercles gradually become green, the horn redder, and pale dorso-lateral and ventro-lateral lines appear along the side. Fully grown, most are whitish green with a dull red horn, purple-red ventral surface and with purple-red surrounds to the spiracles. The body is very rough to the touch. Many are heavily spotted with additional reddish blotches, especially those feeding on purple-spotted scabious leaves. When fully grown, such blotches are masked as the larva turns a rich plum colour prior to pupation.


Full-grown green form larva of Hemaris tityus, Finland. Photo: © Tony Pittaway Heavily blotched full-grown green form larva of Hemaris tityus, Catalonia, Spain. Photo: © Ben Trott. Final instar blotched green formlarva of Hemaris tityus, Catalonia, Spain. Photo: © Ben Trott.

Throughout its life, the larva generally remains beneath a leaf, nibbling holes on either side of the midrib on which it rests. Most active at night; if disturbed during the day, it will drop to the ground, many even taking shelter there when not feeding. A noteworthy feature of this species is the variation in growth-rate of larvae of the same brood; however, all tend to mature much more rapidly than those of other sphingids.


Final instar larva of Hemaris tityus at rest, Catalonia, Spain. Photo: © Ben Trott. Young larvae of Hemaris tityus showing typical feeding damage, Catalonia, Spain. Photo: © Ben Trott. Final instar larva of Hemaris tityus at rest, Catalonia, Spain. Photo: © Ben Trott. Final instar larva of Hemaris tityus at rest, Catalonia, Spain. Photo: © Ben Trott. Part of a plate from Curtis (1823-1840).

In Finland and other parts of northern Europe usually found in June-July; in the warmest parts of its range, from May to September.

Major Hostplants. In northern and central Europe, Succisa pratensis and Knautia arvensis.

Minor Hostplants. Species of Scabiosa, Galium, Lonicera, Symphoricarpos, Dipsacus and Lychnis. Reports of Centaurea (Marttila et al., 1996) require confirmation, but seem unlikely.

PUPA: 24--27mm. Typically Hemaris in shape but blackish brown, tinted with reddish brown at the juncture of each segment. Body tapers at either end, with two sharp head tubercles and a flat triangular cremaster. Formed in a rather strong but coarse cocoon among grass tussocks, or slightly into the soil. The overwintering stage.


Pupa of Hemaris tityus (ventral view), Barcelona, Spain. Image: © Tony Pittaway. Pupa of Hemaris tityus (dorsal view), Barcelona, Spain. Image: © Tony Pittaway.

PARASITOIDS

Tachinidae: Tachina praeceps (Meigen).


DISTRIBUTION

From Ireland (Birchall, 1870; Lavery, 1991) across temperate Europe (including Spain, where it occurs in the north (Galicia (Pino Pérez et al., 2009)), in a few central mountain refugia (Pérez De-Gregorio et al., 2001), and the Sierra Nevada of Almeria Province (Muñoz Sariot, 2014)) to the Urals (Eversmann, 1844; Kumakov, 1977), western Siberia, the Altai Mountains (Zolotarenko, Petrova & Shiryaev, 1978; Izerskiy, 1999), the Tian Shan of both western China and eastern Kazakhstan (Alpheraky, 1882; Eitschberger & Churkin, 2002), and Kyrgyzstan (Eitschberger & Churkin, 2002). Also, from Turkey (de Freina, 1979) to northern Iran (Brandt, 1938; Sutton, 1963), the Republic of Georgia, Armenia and Azerbaijan (Didmanidze, Petrov & Zolotuhin, 2013). Absent from northern Scandinavia, most of arctic Russia, Portugal and the Netherlands, where it is now extinct (Meerman, 1987). Recorded as far north as Pomozdino in european Russia (Tatarinov, Sedykh & Dolgin, 2003), as well as from Sicily and most of the rest of Italy (Nardi & Spada, 2013).

Its extinction in the Netherlands coincides with the extinction of the butterfly Eurodryas aurinia (Rottemburg, 1775), which tends to share the same type of habitat (van Swaay, 1990). It has also declined in eastern Finland due to the loss of suitable dry habitats (Marttila et al., 1996).

The records from Almeria Province, Spain, (Muñoz Sariot, 2014) require further study as the population involved could be Hemaris aksana (Le Cerf, 1923).

Extra-limital range. From Novosibirsk and the Altai (Kondratiev coll., BM(NH); Izerskiy, 1999) eastward across the Tuva A.S.S.R. (Eitschberger et al., 2005) and the rest of southern Siberia to north-eastern China (Derzhavets, 1984) and southwards to Xizang Province/Tibet (Chu & Wang, 1980b), including Qinghai Province (Lake Qinghai), China.

This species is very sparodic and rare east of Novosibirsk.


OTHER SUBSPECIES

None.


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