SPHINX PINASTRI Linnaeus, 1758

GB: Pine Hawkmoth, F: Sphinx du Pin, D: Kiefernschwärmer; Tannenpfeil, RU: sosnovy brazhnik, S: Tall-Svärmare, NL: Dennenpijlstaart, CZ: Lišaj borový, H: fenyöszender, E: esfinge del pino, PL: Zawisak borowiec, FI: Mäntykiitäjä, IT: sfinge del pino.

Sphinx pinastri Linnaeus, 1758, Syst. Nat. (Edn 10) 1: 492.

Type locality: Unspecified [Europe].

(Taxonomic notes. (i) Sphinx pinastri euxinus cannot be considered a distinct subspecies as the genitalic differences used to separate it from Sphinx pinastri are not marked; in these respects it differs as little from the nominate race as do 'subsp.' cenisius Jordan or 'subsp.' medialis Jordan, which are synonymized with pinastri (for the reasons given below).

(ii) Due to consistent differences between the male genitalia and every early stage of Sphinx pinastri and Sphinx maurorum (Jordan, 1931), the latter must now be considered a distinct species.

(iii) The Oriental Sphinx morio (Rothschild & Jordan, 1903), Sphinx caligineus (Butler, 1877), Sphinx yunnana Brechlin, 2015, Sphinx bhutana Brechlin, 2015 and Sphinx oberthueri (Rothschild & Jordan, 1903), previously considered subspecies of Sphinx pinastri, are most certainly separate species, with distinct differences in behaviour and genitalia (Derzhavets, 1979b; Litvinchuk, 1986; Owada & Kogi, 1992; Brechlin, 2015).)

In this work the following classification and synonymies have been adopted for the Sphinx pinastri 'complex', including those from eastern Asia:


Sphinx pinastri Linnaeus
Sphinx maurorum Jordan
Sphinx morio morio Rothschild & Jordan
Sphinx morio arestus Jordan
Sphinx morio inouei (Owada & Kogi)

Sphinx oberthueri Rothschild & Jordan

Sphinx bhutana Brechlin

Sphinx caligineus caligineus (Butler)
Sphinx caligineus brunnescens (Mell)
Sphinx caligineus sinicus (Rothschild & Jordan)

Sphinx yunnana Brechlin

The subspecies marked with an asterisk (*) appear to be the result of hybridization between several races and subspecies which evolved in separate refugia during the last ice age, a conclusion also reached by Kernbach (1958).

Sphinx morio is now recognized as a species distinct from Sphinx pinastri. Subsp. arestus (Jordan, 1931) inhabits central and eastern Russia (Zolotarenko, Petrova & Shiryaev, 1978), Mongolia, north-eastern China (Chu & Wang, 1980) and Korea (Owada & Kogi, 1992); subsp. morio occurs in central and northern Honshu, Japan; and subsp. inouei (Owada & Kogi, 1992) is confined to northern Hokkaido. (Confusingly, Sphinx morio was incorrectly reinstated as a subspecies of Sphinx pinastri by Inoue in Inoue et al. (1982: 318).). According to Litvinchuk (1986), Sphinx morio mates only during the morning. This differs from the behaviour of Sphinx pinastri and, if confirmed by further studies, may explain how these two species remain reproductively isolated. A similar mechanism keeps Hyles euphorbiae (Linnaeus, 1758) and Hyles tithymali (Boisduval, 1834) isolated.

The very similar Sphinx caligineus, Sphinx yunnana, Sphinx bhutana and Sphinx oberthueri are restricted to temperate eastern Asia. In fact, all seven species are similar in appearance, both as adults and larvae.


A plate from Nemos (ca. 1895). A plate from Donovan (1801).

BIOGEOGRAPHICAL AFFILIATION

Holarctic; western Palaearctic region. Pleistocene refuge: Polycentric -- Pontomediterranean subsection of the Mediterranean refuge, as well as the Caspian refuge.


ADULT DESCRIPTION AND VARIATION

Female Sphinx pinastri, Duben, Germany.

Wingspan: 70--96mm. Shows little resemblance to any other western Palaearctic hawkmoth, except to some extreme forms of Sphinx ligustri Linnaeus, 1758. Ground colour very variable, ranging from dark brown (f. brunnea Spuler) to cream (f. albescens Cockayne). The normal grey colour may be of almost any shade, while the dark grey, discal bands on the forewing may be absent, or so heavy as to coalesce, forming one solid band (f. semilugens Andreas).


Male Sphinx pinastri, Wiltshire, England. Photo: © Tony Pittaway

In the male genitalia, the upper sacculus branch is much longer than the lower, more or less heavily curved, almost always cylindrical in its apical half and with only rudimentary prongs (see below right). The apical apophysis and aedeagus are long. This is markedly different from that of Sphinx maurorum (see below left).


Male genitalia of Sphinx maurorum and Sphinx pinastri. Image: © Tony Pittaway.

ADULT BIOLOGY

Found in open or mixed pine forests, especially on dry heaths and wooded steppe (Izerskiy, 1999); also in mountain conifer forests up to 1600m in the Alps (Forster & Wohlfahrt, 1960), but at around 2000m in the Lebanon (Zerny, 1933; Ellison & Wiltshire, 1939).


Typical habitat of Sphinx pinastri, Oxfordshire, England. Photo: © Tony Pittaway

Rests by day on a solid surface, usually the trunk of a pine tree, where it is extremely well camouflaged. Most often, single or isolated groups of trees are selected and it is in such situations that mating pairs are often found, for females are reluctant to fly until paired and remain on the trunk near where they emerged. Although some will separate in mid-afternoon, pairs normally remain in copula until dusk, when males fly off to feed, and possibly mate a second time, while females commence egg laying. Most sweet-smelling flowers, especially Lonicera, are avidly visited, with no time preference being shown. Both sexes are regularly attracted to light.


Feeding Sphinx pinastri, Hungary. Photo: © Szabolcs Safian.

According to Izerskiy (1999), females tend to be active between 23.00h and 01.10h, males between 23.40h and 02.50h. This is in distinct contrast to the related eastern species, Sphinx morio arestus, which is active at dawn.


FLIGHT-TIME

Univoltine; in northern latitudes and Siberia, June or even July; farther south bivoltine, during May/June, and again in August as a partial to full second brood. In the southern Urals, from late May until early July (Nupponen & Fibiger, 2002). In southern Turkey (Adana Province), from late May until late June (Feza Doganlar, pers. comm.).


EARLY STAGES

OVUM: Oval and slightly dorso-ventrally flattened (2.00 x 1.5mm); shiny pale yellow at first, changing to reddish yellow. Each female lays approximately 100 eggs, usually singly in close proximity to each other, on the needles or young twigs of isolated or small groups of trees. Development takes 6--9 days at 19--22°C, 14--20 days under cooler conditions. Just before hatching the dark head of the larva becomes visible through the now transparent shell.


Egg of Sphinx pinastri, Oxfordshire, England. Photo: © Tony Pittaway. Egg of Sphinx pinastri, Oxfordshire, England. Photo: © Tony Pittaway.

LARVA: Full-fed 75--80mm. Dimorphic: predominantly green or brown.

On hatching, the larva consumes part of its eggshell. At this stage it is approximately 5mm long and dull yellow, with a disproportionately large head marked with brown, and a dark, forked horn. With feeding, the body colour changes to green and, after the first moult, six longitudinal creamy yellow lines appear. Still disproportionately large, the head is a paler green than the body, and has dark cheeks. During this stage, the larva sits lengthways on a pine needle, with which it blends very well. Apart from the longitudinal lines becoming broader and the horn and legs acquiring a reddish tint, little change takes place to its appearance over the next two moults.


Unfed first instar larva of Sphinx pinastri, Oxfordshire, England. Photo: © Tony Pittaway. Third instar larva of Sphinx pinastri, Oxfordshire, England. Photo: Mark Boddington. Fourth instar larva of Sphinx pinastri, Oxfordshire, England. Photo: © Tony Pittaway.

Fully grown, larvae may be either green with a marked brown dorsal band and three broken cream or white lateral stripes, or mainly brown, blending well with surrounding twigs. In both colour forms the whole body, which is slender and of even thickness, has dark, sunken lines, apparently dividing it up into narrow rings behind a large, prominent head. The body is smooth, but not glossy, unlike the head, underside, legs and shield. However, an oily appearance extends over the entire body as the larva darkens prior to pupation.


Resting full-grown larva of Sphinx pinastri, Oxfordshire, England. Photo: © Tony Pittaway. Feeding full-grown larva of Sphinx pinastri, Oxfordshire, England. Photo: © Tony Pittaway.

Initially, feeding consists of nibbling at the surface of a needle but, with increasing size, whole needles are consumed, being grasped between the legs and eaten from the tip to the base; older needles are preferred to younger ones. It is a sluggish diurnal feeder, never moving more than necessary, and comparatively little plant matter is consumed during this 4--8 week period. However, just before pupation, the larva becomes very restless and, after descending from the hostplant, it will wander some distance in search of a pupation site.


Typical feeding damage of Sphinx pinastri, Oxfordshire, England. Photo: © Tony Pittaway

In Finland, larvae prefer short and weathered pines (Pinus sylvestris), especially ones growing beside bogs, marshes, sandy areas or other open places. Larvae live in the top branches and feeding sites are easy to spot later in the summer as the terminal shoots are usually eaten bare (Jaakko Pohjoismaki, pers. comm.).

Feeds during July, August and September in the north, and June/July and September/October in southern regions. At times, larvae can be so abundant as to constitute a pest (Gninenko, 1998).

Major Hostplants. Various species of Pinus (especially Pinus sylvestris) and Picea (especially in the Alps). Picea obovata is the main secondary host in Siberia (Izerskiy, 1999).

Minor Hostplants. Larix decidua (northern and central Europe), Cedrus (southern Europe) and Pseudotsuga menziesii (Douglas Fir).


PUPA: 35--40mm. Very similar to that of Sphinx ligustri and usually formed under moss or the needle mat found at the base of trees. The overwintering stage; may overwinter twice.


Pupa of Sphinx pinastri, Oxfordshire, England. Photo: © Tony Pittaway.

PARASITOIDS

Ichneumonidae: Amblyjoppa proteus (Christ), Aphanistes armatus (Wesmael), Aphanistes bellicosus (Wesmael), Aphanistes megasoma Heinrich, Aphanistes ruficornis (Gravenhorst), Coelichneumon deliratorius (Linnaeus), Coelichneumon sinister (Wesmael), Cratichneumon sicarius (Wesmael), Diphyus palliatorius (Gravenhorst), Heteropelma amictum (Fabricius), Heteropelma calcator (Wesmael), Metopius dentatus (Fabricius), Protichneumon fusorius (Linnaeus), Protichneumon pisorius (Linnaeus), Therion circumflexum (Linnaeus); Pteromalidae: Erdoesina alboannulata (Ratzeburg); Encyrtidae: Ooencyrtus pityocampae (Mercet) (an egg-parasite); Tachinidae: Campylochaeta inepta (Meigen), Compsilura concinnata (Meigen), Drino inconspicua (Meigen), Exorista larvarum (Linnaeus), Nemoraea pellucida (Meigen), Phryxe erythrostoma (Hartig), Phryxe nemea (Meigen), Phryxe vulgaris (Fallén), Tachina grossa (Linnaeus), Winthemia cruentata (Rondani).

Phryxe erythrostoma is a specific parasitoid of Sphinx pinastri, with as many as eighteen examples being recorded from one host pupa.

Attacked by the fungus Cordyceps militaris in Poland (Zablocka, 1929).


DISTRIBUTION

From Europe (except Iberia, Ireland, northern Scandinavia and Arctic Russia) and western Siberia (Eversmann, 1844; Zolotarenko, Petrova & Shiryaev, 1978) south to the Crimea (Efetov & Savchuk, 2012), Daghestan (Didmanidze, Petrov & Zolotuhin, 2013), the Republic of Georgia (Didmanidze, Petrov & Zolotuhin, 2013), the Caucasus (Derzhavets, 1979b; Abdurahmanov, 1999) and southern Turkey (Daniel, 1932; Daniel, 1939; Kernbach, 1958; de Freina, 2012), Lebanon (Zerny, 1933; Ellison & Wiltshire, 1939) and northern Israel (Müller et al., 2005b). Kurgan and Gorky (Kulebyaki) appear to be the eastern-most distribution of Sphinx pinastri in Siberia, but recorded as far north as Arkhangelsk Oblast (Kozlov, Kullberg & Zverev, 2014) and Pechora (Tatarinov, Sedykh & Dolgin, 2003) in european Russia. In western Europe it is found east and north of central France (Pittaway, 1983). There are some local populations in central-southern France showing small differences in genitalia which were separated into individual subspecies by Jordan (1931), but this is unwarranted. Also found in northern and central Greece, and in the northern Aegean on the islands of Thassos (Koutsaftikis, 1970) and Samos (Fritsch et al., 2014).

Duncan (1836) regarded this species as being rare in Britain, with it occurring with certainty only around Esher, Surrey; Drury (1837) stated that Sphinx pinastri was only a very rare vagrant. With the extensive cultivation of pine plantations over the last 100 years, it has become well established and has spread over most of southern England where both commercial and ornamental pine trees occur. A good account of its early status in England is given by South (1907).

The situation in France appears to have changed over the last 20 years, with this species having been displaced by Sphinx maurorum from many southern areas. True Sphinx pinastri can now only be found north of Chateauroux, in the Alps and Var. Populations around Toulon are clearly intermediate hybrids (J. Haxaire & J.-M. Bompar, pers. comm.). The population on Corsica (Bretherton & de Worms, 1963) appears to be Sphinx maurorum (Haxaire, 2009).

Extra-limital range. None.

Sphinx pinastri has been recorded from certain parts of the Canadian Rocky Mountains, and specimens have been found in the eastern U.S.A. (Hodges, 1971). Whether the former are native or escapees is not known. It is possible that they are the last remnants of the original North American population of this species which went on to colonize the Palaearctic, but this, of course, is pure speculation.


OTHER SUBSPECIES

None.


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