SYSTEMATICS


INTRODUCTION

The various systems used to classify the Lepidoptera, and in particular the Sphingidae, are reviewed briefly, as is the morphology of the ovum, larva, pupa and imago. This provides a basic understanding of those features used to classify the species and subspecies covered by this work, and hence a sound scientific basis for analysing the ecology and biogeography of certain taxa. The species concept is discussed, but only in so far as it helps to explain why certain taxonomic and biogeographical conclusions were arrived at.

The synonymy of each taxa is fully revised and those features characteristic of each family, subfamily, tribe and genus is updated and expanded. The information for each species and subspecies has been enhanced with new observations, particularly with regard to hostplants, distribution an seasonal abundance. Field observations, the rearing of many species and subspecies, coupled with a species concept based on population biology provides the basic data for several taxonomic changes at the specific or subspecific level. These conclusions refute or corroborate earlier taxonomic decisions resulting from the study of limited samples in museums. Ecological and biological observations are extremely useful in helping resolve taxonomic problems in the formally confused and confusing Hyles euphorbiae complex.

However, the other question that needs to be addressed here is of what relevance to sphingid biogeography is the study of their anatomy, morphology and taxonomy? To quote Hengeveld (1990), "Although the immediate aim of dynamic biogeography is to describe and explain spatial patterns and processes of taxa, its ultimate goal is to aid our understanding of evolutionary processes. We must integrate information of a great number of properties of individuals, populations and the species as a whole. Thus, it will sometimes be necessary to collect information about an individual's physiology, anatomy or morphology, whereas at other times information is required about the spatial distribution of genetic traits, behavioural characteristics, or population dynamic statistics." He further states that "Looking from the viewpoint of the central issue of biology, namely evolution, distribution ranges and the processes occurring within them should not be considered merely as a taxonomic character, but as the outcome of the interaction of a specie's biological properties with each other and with the properties of the environment. Characteristics such as shape, size, structure, location and dynamics of species ranges integrate and reflect all of these interactions, and can show their relative importance at a particular moment or through evolutionary time."


CLASSIFICATION

From Linnaeus until the present century, the criteria on which entomologists based the systematic division of Lepidoptera were numerous. Classification systems differed one from another and are difficult to compare with those in use today as they were based mainly on superficial characters, such as wing pattern and shape.

As his criteria, Linnaeus used wing positions of adults at rest. Latreille based his classification on the times of adult activity (i.e. Nocturni, Diurni) whereas Boisduval used antennal shape -- the Rhopalocera (butterflies) having 'clubbed horns' (i.e. clubbed antennae); the Heterocera (moths) having 'horns otherwise' (i.e. antennae without clubs). None of these distinctions is valid, for even though Boisduval's Rhopalocera do form a monophyletic group, his Heterocera do not.

Another loose form of classification still in use today -- Microlepidoptera and Macrolepidoptera -- is based on adult size, although some 'Micros' have wingspans up to 200mm. Four modern suborders of Lepidoptera represented in the western Palaearctic (Zeugloptera, Dacnonypha, Exoporia and Monotrysia) contain only families of Microlepidoptera, but a fifth, the Ditrysia, includes families of both 'Micros' and 'Macros', among which is the Sphingidae.

Many other classifications have been proposed in addition to those referred to above, based on different morphological features (Comstock (1893), Packard (1895), Karsch (1898), Tillyard (1918, 1919), Börner (1925), Hinton (1946) and Common (1970)); none of these has contributed much to the present systematic arrangement of the sphingids themselves, but since 1900 three systems have been of sufficient merit to have had a significant impact on sphingid classification, namely Rothschild and Jordan (1903), Carcasson (1968) and Hodges (1971).

The position of the Sphingidae within the Lepidoptera is well reviewed by Scoble (1991) in his valuable summary of the classification of the Lepidoptera at higher taxonomic levels. He points out that this classification is in a state of flux, especially between superfamily and suborder levels, but he follows Minet (1986) and Holloway et al. (1987) in keeping the Sphingidae within the Bombycoidea rather than treating them as a separate superfamily, the Sphingoidea, as do Common (1970), Hodges (1971) and Kuznetsov & Stekolnikov (1985) although all these authors have recognized their close relationship to the Bombycoidea.

In this work the following classification is adopted:



In their monumental work, Rothschild & Jordan (1903) recognized 772 species. Prior to this, the Sphingidae had been classified on purely superficial characters with the result that many unrelated species had been lumped together and many closely related species had been separated. However, Rothschild & Jordan admitted that they were very much hampered in their attempts by a lack of material and knowledge regarding the early stages. Only a few of the more common species had been bred or reared and the published descriptions of these were inadequate. Nevertheless, they were the first authors to adopt a natural, phylogenetic classification, using characters such as the structure of antennae, palpi, pilifer and feet, spination of the legs and abdomen, and dry genitalia preparations of a large number of species. The family Sphingidae was split into two divisions (Asemanophorae and Semanophorae), five subfamilies and seven tribes.

Carcasson (1968), in his excellent work on the African Sphingidae, proposed a revision to the classification of the Sphingidae of Rothschild & Jordan based on the genital armatures of both sexes and on the early stages. Because they were using dry genitalia preparations, Rothschild & Jordan were not enabled'....to take these structures into sufficient account when defining genera and following up their relationships. In the Philampelini and Choerocampini this did not matter much as the genitalial structure of these insects is extremely uniform, but it did lead to a number of misconceptions in the more advanced Ambulicini'. The names proposed by Rothschild & Jordan for supra-generic taxa were adopted by Carcasson, although several older and more appropriate names were substituted. The terms Sesiinae and Sesiicae were rejected because the type genus of these taxa is not a sphingid. The other groups were given the terminations recommended in the International Code on Zoological Nomenclature (1961, 1985). The term 'subfamily' was substituted for 'division', which has no nomenclatural status; all subsequent taxa above the rank of genus were demoted by one step. The 'tribes' of Rothschild & Jordan became 'subtribes', a useful rank which was not recognized in the International Code until 1985.

Hodges (1971), in his authoritative work on the Sphingidae of America north of Mexico, drew up a further revised system of classification, commenting that Rothschild & Jordan based their's '....on a very large number of characters, mainly of the adults, but unfortunately they studied the male and female genitalia in a dry condition, so they were unable to discern characters of the female genitalia, and they did not figure (and thus probably did not take into full account) the full array of male genital characters. These two character systems in combination provide what seems to be a cohesive framework for a supergeneric [sic] classification within the Sphingidae. Many additional characters of the larvae and pupae fall into line if the associations of the genital systems are followed rather than those of the spination of the pilifer or of certain palpal characters'. Hodges further added that '....Rothschild & Jordan pointed out that the subfamilies and tribes within the Semanophorae were not equal and that several species and genera within each had characters which would tend to put them either one way or the other with the system. The higher categories could not be defined on the basis of just a few characters, but, perhaps in combination, several characters would suffice for determination'. Hodges agreed with Carcasson in redefining the Semanophorae and Asemanophorae as subfamilies but points out that Carcasson incorrectly used these names; the International Code of Zoological Nomenclature states that family-group names must be based on a valid or available generic name. The subfamilies Sphinginae and Macroglossinae were proposed as substitutes, respectively.

Minet (1994) raised the point that since Hodges' Sphinginae were most likely paraphyletic, with the Smerinthini and Sphingini only showing a few symplesiomorphies, the two tribes should be raised to subfamily status. Kitching, I.J. (pers. comm.) is in agreement with this. Thus the subfamilies and tribes as used by Kitching & Cadiou (2000) have been adopted here. The supraspecific categories for the Sphingidae of the western Palaearctic region described in this work are shown in the species list.

Recently, some of the above findings have been questioned. Placement of the Sphingidae within the Bombycoidea is still supported by Kawahara et al. (2009) in their large-scale molecular phylogenetic analysis of the Sphingidae, where they analysed 6793 bp of sequence from five protein-coding nuclear genes from 131 species representing all subfamilies, tribes and subtribes. However, some of their results challenge aspects of the current lower taxonomy based on morphology, and provides a foundation for a new classification [to come].


INDIVIDUAL TAXA

The categories used by Kitching & Cadiou (2000) have been adopted as the classification system throughout this work for the various western Palaearctic genera.


FAMILY SPHINGIDAE Latreille, [1802]

GB: Hawkmoths, F: Sphingides, D: Schwärmer, RU: Brazhniki, S: Svärmarnas, NL: Pijlstaarten, CZ: Lišajovití, H: Szenderek, E: Esfíngidos

Sphingides Latreille, [1802], in Sonnini, Hist. nat. gen. particuliere Crustaces Insectes 3: 400.

Characterized by the combination of vein R1 of the hindwing crossing to vein Sc from the middle of the cell, the absence of the base of vein M1 and loss of vein 1A.



SUBFAMILY SMERINTHINAE Grote & Robinson, 1865

Smerinthini Grote & Robinson, 1865

Type genus: Smerinthus Latreille, [1802].

Distributed worldwide and represented by almost XXX species.

IMAGO:

Genitalia:

LARVA:

PUPA:


TRIBE SMERINTHINI Grote & Robinson, 1865

Smerinthini Grote & Robinson, 1865

Type genus: Smerinthus Latreille, [1802].

A tribe of more than 200 species distributed throughout the Old World, but with 10 species in the Holarctic region north of Mexico.

IMAGO: Forewing primarily yellow-brown, greyish lilac or green, with a few crossbands and spots. Hindwing usually with a brownish or reddish brown field; crossbands reduced to a pale or ocellate spot at anal angle. Proboscis often atrophied. Spines on caudal margins of abdominal terga generally diffuse.

Genitalia: Male with uncus variable, with a flat, entire or split tip; gnathos absent, bilobate or entire; sacculus with two simple or bilobate processes and pectens; saccus not well developed; aedeagus relatively short and often with a stout apical cornutus. Valva often with pouches on inner surface distally. Female with ductus bursa usually broad and short.

LARVA: Horn developed, except in Pachysphinx; cuticle rough with fine tubercles; oblique lateral stripes present, sometimes in combination with longitudinal lines. Body usually tapering forward from abdominal segment 5.

PUPA: Proboscis fused to body. Labrum displaced ventrally. Abdominal segments 5--7 with 1--4 interrupted, punctate furrows over each spiracle.


TRIBE SPHINGULINI Rothschild & Jordan, 1903

Sphingulicae Rothschild & Jordan, 1903

Type genus: Staudinger, 1887.

A tribe of XXX species with a very disjunct distribution. Pentateucha and Dolbina occur mostly in eastern and southeast Asia, with two species penetrating westwards. The second group is confined to Australia. This may reflect Gondwanaland origins.

IMAGO: Antenna of male with well-developed setae, those of the female weakly ciliate; terminal segment short. Proboscis shorter than thorax. Posterior margin of abdominal segments with weak spines, which are sometimes lacking on the sternites.

Genitalia:

LARVA: Head large and triangular, tapering dorsally. Cuticle rough, covered with minute tubercles in final instar. Horn straight. Several oblique lateral stripes present on abdominal segments.

PUPA: Known only for Dolbina. Smerinthine, similar to Marumba, with no dorsal sculpturing on thoracic segment 3 but with indications of antespiracular ridges on abdominal segments 5--7.


TRIBE AMBULYCINI Butler, 1876

Ambulicinae Butler, 1876

Type genus: Ambulyx Walker, 1856 (=Protambulyx Rothschild & Jordan, 1903)

Distributed worldwide throughout the tropics and subtropics, and represented by almost XXX species.

IMAGO: Proboscis sometimes reaching abdomen but usually short and non-functional. Terminal antennal segment long or short. Forewing with extended or falcate tip.

Genitalia:

LARVA: Head large, usually rounded. Body granulose with several oblique lateral stripes.

PUPA: Anterior end blunt.



SUBFAMILY SPHINGINAE Latreille, [1802]

Sphingides Latreille, [1802], in Sonnini, Hist. nat. gen. particuliere Crustaces Insectes 3: 400.

Type genus: Sphinx Linnaeus, 1758.

Distributed worldwide and represented by almost 400 species.

IMAGO: First segment of labial palpus medially without sensory hairs on the naked area; proboscis developed or vestigial; frenulum and retinaculum sometimes absent; eighth sternum evenly sclerotized; midtibia with one pair of spurs distally or with two pairs of spurs.

Genitalia: Symmetrical in the male, with the well-developed sacculus bearing complex excrescences; saccus also well developed. Both uncus and gnathos usually present with single process. Female with both ante- and postvaginal lamellae; base of ductus bursae heavily sclerotized.

LARVA: Head large and round or tapering dorsally. Horn usually present, as are oblique side stripes.

PUPA: Proboscis free or fused with body.


TRIBE SPHINGINI Latreille, [1802]

Sphingides Latreille, [1802], in Sonnini, Hist. nat. gen. particuliere Crustaces Insectes 3: 400.

Type genus: Sphinx Linnaeus, 1758.

A tribe comprising more than 150 species distributed throughout the Old and New World.

IMAGO: Forewing ground colour brown or grey, with numerous undulating crossbands. Hindwing usually with dark submarginal bands; central area yellow, brown or pink. Inner surface of second segment of labial palpus depressed with a naked area, or scaled and lacking this depression. Proboscis normally well developed and functional. Abdominal spines of variable length and arranged in several overlapping rows.

Genitalia: Male with uncus broad basally and with uncinate extension or a flat tip; gnathos absent, but if present, bilobate or entire, then both uncus and gnathos heavily sclerotized apically; sacculus variable, but with two pointed, straight or curved processes, or with extended, pectinate apical processes. Female with short or long ductus bursae, which is evenly broadened.

LARVA: Usually with several oblique lateral stripes, and smooth in the final instar. Horn usually present (except in Lapara).

PUPA: In those species with adults bearing an elongate proboscis, proboscis free from body. In species where the adult has a short proboscis, this is fused to body. Labrum displaced ventrally. Abdominal segments 5--7 bearing one or two spiracular furrows.


TRIBE ACHERONTIINI Boisduval, [1875]

Achérontides Boisduval, [1875]

Type genus: Acherontia [Laspeyres], 1809.

Distributed worldwide and represented by almost XXX species.

IMAGO: Segment 2 of labial palpus with a deep cavity on the inner side; terminal antennal segment long, filiform and covered with scales and setae; proboscis functional; posterior margin of abdominal segments with a few irregular rows of spinules.

Genitalia:

LARVA: Head rounded; cuticle smooth in final instar. With several pronounced oblique lateral stripes.

PUPA: Glossy, with or without free proboscis. Metathorax with transverse carina.



SUBFAMILY MACROGLOSSINAE Harris, 1839

Macroglossiadae Harris, 1839, Am. J. Sci. Arts 36: 287.

Type genus: Macroglossum Scopoli, 1777.

Distributed worldwide and comprising more than 600 species.

IMAGO: First segment of labial palpus on mesal side with short sensory hairs on the naked area; proboscis long; frenulum and retinaculum always well developed; spines on caudal margins of abdominal terga either strong or weak, sometimes diffuse; midtibia with a single pair of spurs, hindtibia with two pairs.

Genitalia: In male, symmetrical or slightly to moderately asymmetrical, with gnathos and uncus divided or undivided; saccus sometimes developed; sacculus with a single apical process; eighth sternum usually modified by being sclerotized medially, laterally and basally. In female, lamella postvaginalis developed, lamella antevaginalis absent.

LARVA: Variable. Head large or small, but always rounded. Horn may be absent and replaced by a 'button' in the final instar. Tapering anteriorly from thoracic segment 3 or abdominal segment 1 to head. Eye-spots may be present on these segments.

PUPA: Proboscis fused with body, but may project into a ridge (carinate).


TRIBE DILOPHONOTINI Burmeister, 1878

Dilophonotidae Burmeister, 1878

Type genus: Dilophonota Burmeister, 1856 (=Erinnyis Hübner, [1819]).

A tribe of nearly 150 species distributed throughout the tropical and temperate regions of the Old and New Worlds, but with its stronghold in the Neotropical Region.

IMAGO: Pattern of forewing variable.

Genitalia: Very diverse and often asymmetrical in both sexes. In male, uncus with beak-like extension, entire or divided, but with a marked furrow on the medial line; gnathos spatulate (entire or split), but sometimes absent; sacculus with complex apical process armed with spines; saccus well developed. In female, lamella postvaginalis broad and developed; sclerotized part of ductus bursae short but the two slender bands on the posterior margin always well defined.

LARVA: Diverse; the horn may be very slender and movable. Many have tubercles or excrescences on the thoracic segments and/or anal segments.

PUPA: Proboscis fused with body. Labrum displaced ventrally.


TRIBE MACROGLOSSINI Harris, 1839

Macroglossiadae Harris, 1839, Am. J. Sci. Arts 36: 287.

Type genus: Macroglossum Scopoli, 1777.

A cosmopolitan tribe of more than 400 species.

IMAGO: Pattern of forewing consisting of transverse (Macroglossina) or oblique (Choerocampina) bands; basal tone brown-grey, olive-green or brown-purple; hindwing with contrasting median area and dark submarginal crossbands.

Genitalia. In male, uncus and gnathos with rostrate extensions directed caudad so as to resemble a crab's claw; the apex of each heavily sclerotized and tapering to a sharp or blunt point. Tegumen extended caudad into a broad tube; sacculus with apical process, accessory armature weakly developed or absent; saccus in the form of a narrow fold of vinculum. In female, lamella postvaginalis short.

LARVA: Head small or large, rounded; horn developed or replaced by a tubercle in the final instar; cuticle smooth in all instars; body with longitudinal stripes or faint oblique lateral stripes; several ocellate spots may be present along the body, that on the third thoracic or first abdominal segment being well developed in some species.

PUPA: Proboscis fused with body; may be raised into a protruding ridge; in Pergesa, proboscis free of body.


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