New Zealand Ichneumonidae (Paper No. 3.) Sub-family Ophioninae. Tribe Ophionini
[Read by title before Nelson Institute, 20th April, 1953; received by Editor, 15th May, 1953.]
This is the third paper of a series on the systematics of the New Zealand Ichneumonidae, and deals with the Tribe Ophionini. A brief historical review and the status of the various species previously recorded from New Zealand, with keys to the genera and species is given.
Australophion inflatus Cushman, 1947, is considered synonymous with A. peregrinus (Smith). A detailed account of the variability of this species is given with notes on its seasonal occurrence. The genus Ophion is considered to be so far represented in New Zealand by two plastic species, each of which may be separated into two or more well defined sub-species. There are described in the present paper four sub-species of O. inutilis Smith, and two sub-species of O. occulatus, a previously undescribed species.
The Tribe Ophionini, consisting of small to medium sized red-brown Ichneumonflies, forms a significant element of the ichneumonid fauna in all the main faunal regions of the world.
Although numerically the number of species found in New Zealand is not large, individuals are frequently observed in a wide variety of situations, from sea-level to an altitude of 5,000 feet and from densely populated areas to virgin bush.
This paper is based on the collections of the Dominion Museum, Canterbury Museum, Auckland Museum, the Cawthron Institute, and the Entomological Research Station, Nelson. I wish to express my sincere thanks to Dr. R. A. Falla, Director of the Dominion Museum, Wellington, Dr. Gilbert Archey, Director of the Auckland Institute and Museum, Auckland, Dr. Roger Duff, Director of the Canterbury Museum, Christchurch, and to Dr. D. Miller, Director of the Entomological Research Station, and Assistant Director of the Cawthron Institute, Nelson, for allowing me access to the collections under their charge.
The classification of the Ophioninae has become over the years rather confusing, and the limits assigned to the various sub-groups unsatisfactory. Recently, Cushman (1947) and Townes (1944) as well as some European workers, have clarified the position to some extent.
Ashmead (1900: 84) divided the Ophioninae into twelve tribes; Townes (1944 and 1945) raised the status of the tribes Plecticini and Mesochorini to sub-family rank and transferred the tribes Banchini and Panicini to the subfamilies Lissonotinae and Tryphoninae respectively.
The Ophionini may be distinguished from all other Ophioninae, in that the 2nd recurrent vein is emitted from the cubitus distinctly before the transverse
cubitus, a feature shared only by the Nototrachini (not so far represented in New Zealand) in which the antennae are very short and each intermediate tibia is furnished with only a single tibial spur.
The following combination of characters may serve to distinguish members of this tribe:—
Mandibles when closed and in cephalic view, with lower tooth not directly posterior to upper tooth, but at a lower level and visible; eyes parallel or divergent below, never distinctly convergent (i.e., face not narrower than frons) always at least shallowly emarginate opposite the antennal foramina; occipital carina medially far below level of posterior ocelli, fading out below or joining hypostomal carina at a considerable distance from the lower articulation of mandibles, rarely entirely absent; epomia entirely lacking; propodeal spiracles lateral in position, very long and slit-like; front tibiae without a minute tooth at apex opposite calcarium; middle tibiae with two tibial spurs; claws usually long, each with many strong teeth; 2nd recurrent vein basad of intercubitus by a distance nearly or quite as great as the length of the intercubitus; intercubitus strongly oblique and nearly continuous with the basal abscissa of radius; nervellus sharply broken, discoidella distinct, rarely nervellus not broken, but interstitial at its upper end with discoidella; hind tarsus with apical two segments not especially small; ovipositor rarely longer than depth of abdomen at apex.
Three genera are at present represented in the New Zealand fauna. It is doubtful whether Australophion Morley, should be retained as it grades somewhat gradually into the genus Ophion, in which representatives are found with the scutellum, sub-alar tubercles and the metapleurae somewhat inflated but not to the remarkable degree found in Australophion. Cushman (1947:440) expressed the opinion that Australophion is hardly more than subgenerically distinct from Ophion. Although I agree with Cushman, it is convenient, in the meantime, to retain Morley's genus.
The following species of Ophionini have been described or recorded from New Zealand:—
Ophion peregrinus Smith, 1876.
Ophion inutilis Smith, 1876.
Ophion inutilis Smith, 1878.
Ophion ferrugineous (Fabr.) Smith, 1878.
Ophion insularis Kirby, 1881.
Ophion skeltoni Kirby, 1881.
Ophion luteus (Linnaeus) Hutton, 1882.
Ophion punctatus Cameron, 1898.
Ophion insulicola Dalla Torre, 1901.
Australophion inflatus Cushman, 1947.
There has been considerable confusion in the past concerning the specific determination of the species listed above. This has mainly been due to their uniformity of structure, the absence of types, the majority of which are deposited in overseas museums, and are therefore not readily available to workers in this country, and to the undue importance that has been placed on colour, which varies considerably within a species, and finally, to the inadequate descriptions of the older authors. Colour varies from light-brown to dark-brown in individuals
of the same species, and the black markings which occur in some species show variations in the extent and density of these markings. Colour may be used to differentiate species in a very general manner.
Morley (1912: 30) established the genus Australophion to accommodate Smith's species Ophion peregrinus in which the metapleurae in particular, were inflated to a most extraordinary degree. Cushman (1947:440) added another species, A. inflatus, collected at Mt. Egmont, Taranaki, North Island. Smith's Ophion ferrugineous was considered by Morley, who examined the type in the British Museum, to be synonymous with A peregrinus (Smith).
Both Kirby's species O. insularis and O. skeltoni belong to the genus Enicospilus. Smith described O. inutilis in 1876, and again two years later, in 1878. Morley (1912) considered Smith's O. inutilis 1876, and O. inutilis 1878 to be distinct species, and that O. inutilis described in 1878 is the same as Cameron's species O. punctatus described in 1898. Dalla Torre (1901) also considered the two descriptions referred to different species, and renamed Smith's O. inutilis 1878 as O. insulicola.
Based on the above synonymy both Smith's name O. inutilis 1878, and Dalla Torre's name O. insulicola, must fall as synonymous to Cameron's O. punctatus, as Dalla Torre's name post-dates Cameron's name by three years. The European species Ophion luteus Linnaeus has not been identified from New Zealand since Hutton's record in 1882, and it will not be considered further, as Kirby (1884) has stated “he probably mistook one of the indigenous New Zealand species for it,” and as the only apparent reason for its retention in the New Zealand list is that Fabricius mentioned New Zealand among the localities for this common European form. To summarise what has been said above, we have to consider six species of Ophionini referable to three genera, as follows:—
Australophion inflatus Cushman.
Australophion peregrinus (Smith)
Enicospilus insularis (Kirby)
Enicospilus skeltoni (Kirby)
Ophion inutilis Smith.
Ophion punctatus Cameron.
As the results of the present study, Australophion inflatus Cushman is considered to be synonymous with Australophion peregrinus (Smith) and Ophion punctatus Cameron (= O. inutilis Sm. 1878) as a sub-species of O. inutilis Sm. 1876.
The following is a check-list of N.Z. species and sub-species of Ophionini, with corrections and additions:—
Enicospilus insularis (Smith).
Enicospilus skeltoni (Smith).
Australophion peregrinus (Smith).
Ophion inutilis inutilis Smith.
Ophion inutilis punctatus Cameron.
Ophion inutilis vittatus N. Sub. sp.
Ophion inutilis intermedius N. Sub. sp.
Ophion inutilis nigravarium N. Sub. sp.
Ophion inutilis distinctus N. Sub. sp.
Ophion oculatus oculatus N. Sub. sp.
Ophion oculatus rugosa N. Sub. sp.
In the present paper five species are recognised and six sub-species. Although generally, I do not consider it a useful taxonomic procedure to name subspecific categories on the present state of our knowledge of these insects, but a few sub-specific names seem necessary to call attention to the relations existing within some extremely variable species. What these sub-specific divisions should be called is a matter of opinion. They may or may not be generally characteristic of geographical areas, but may occur in small numbers in widely separated regions. Perhaps the non-committal word “Form” may be considered best to designate these groups of individuals, until more is known of the reasons for their apparently sporadic occurrences. Such forms in the past have been vaguely described as species, sub-species, biotypes, races, forms, and varieties.
Key to Genera Represented in New Zealand
|1. Basal constriction of propodeum undivided, though sometimes coarsely foveolated: Nervellus with upper abscissa usually inclivous: Tergite 2 without umbo: Fenestra distinct with or without scleromes||Enicospilus Stephens|
|Basal constriction of propodeum more or less divided into a medium and two lateral fovea: Nervellus with upper abscissa perpendicular or reclivous: Tergite 2 with umbo: Fenestra absent*||2|
|2. Metapleurae, scutellum and sub-alar tubercles strongly inflated||Australophion Morley|
|Metapleurae, scutellum and sub-alar tubercles not or only weakly inflated: tergite two with lateral fovea at base||Ophion Fabricius|
Genus Enicospilus Stephens 1835
Enicospilus Stephens, Catalogue of British Insects, p. 352, 1820. (Without description or species.) Illustrations of British Entomology, Vol. 7, p. 126, Pl. 10, fig. e, 1835. Cushman, Hawaiian Entom. Soc. Proc., Vol. 12, p. 39, 1944.
Henicospilus Agassiz, Nomen. Zool. index universalis, p. 138, 1846. (Emendation of Enicospilus Stephens.)
(Type species Ophion merdarius Stephens.)
For a complete generic synonymy of this genus reference should be made to Cushman (1947:467) and Muesebeck, Krombein and Townes in their Catalogue of Hymenoptera of America (1951:403). Enicospilus is used here as defined by Cushman (1947:467).
Key to New Zealand Species of Enicospilus.
|1. Fenestra with the proximal, distal and marginal scleromes present (Fig. 1): thorax somewhat deeply punctate, especially noticeable on the basal area of propodeum and on the mesonotum.||E. insularis Sm.|
|Fenestra with the proximal, and marginal scleromes present, the distal sclerome absent (Fig. 2): thorax not strongly punctate, especially is this so, on the basal area of propodeum and the mesonotum.||E. skeltoni Sm.|
Enicospilus skeltoni (Kirby)
Ophion skeltoni Kirby, Trans. Ent. Soc. 1881, p. 46; N.Z. Journ. Sci. 1884, p. 73.
Henicospilus skeltoni (Kirby), Morley, Rev. Ich B.M. Part I, 1912, p. 53.
This species was originally described by Kirby from specimens collected at Blenheim, N.Z., in 1881. Morley (1912:53) recorded a single male of E. skeltoni
[Footnote] * A hairless area below base of stigma should not be confused with a true fenestra.
from Killopanma, in S. Australia, 100 miles east of Lake Eyre, collected by H. J. Hillier, in 1905.
This is the common Enicospilus occurring in at least the northern part of the South Island of New Zealand, and is probably widely distributed throughout both Islands.
Male: 14 mm. to 21 mm. in length.
A testaceous or less frequently a ferrugineous species, with or without the external and internal orbits flavous, rarely face entirely fulvous, claws black, mandibles with teeth dark red-brown, costa and veins brown with stigma testaceous; eyes and ocelli usually liver-brown colour.
Face and clypeus almost impunctate, occasionally minutely punctate, especially at base of clypeus; cheeks narrow, width about ½ the width of eye in profile: cheeks and ocellar area impunctate; pronotum nitidulous, very minutely and sparsely punctate; mesonotum minutely punctate, the punctures widely spaced mesopleurae finely but distinctly punctate, with obsolete longitudinal aciculations, usually more strongly developed on the lower parts of the segment; metapleurae distinctly punctate dorsally, nearly impunctate ventrally: scutellum distinctly punctate, lateral carinae not high, and continued to apex; propodeum with transverse carina usually weak, and sub-straight, occasionally it is obsolete, only indicated at the centre: basal area not strongly punctate, nitidulous: apical area finely rugose, nearly flat: abdomen slender 2nd tergite subequal to 1st in length; 3rd tergite ¾ the length of 2nd with a depth at apex of about ⅔ its own length: antennae not reaching to end of abdomen.
Forewing: 1st abscissa of radius straight, thickened along its basal ⅔: fenestra (Text-fig 1, Fig. 2) long, extending ¾ the length of the 1st abscissa of radius: proximal and marginal scleromes present: distal sclerome absent: proximal sclerome situated under basal ¼ of radius; marginal sclerome widely separated from proximal sclerome, and extending around distal border of fenestra, not quite reaching the radius: nervulus usually not interstitial with basal: discocubital vein gently bent under centre of fenestra and distal to proximal sclerome: Hindwing: abscissula slightly over three times the length of intercubitella: nervellus (Text-Fig. 1, Fig. 3) postfurcal, upper arm perpendicular and broken below centre.
Female: 16 mm. to 20 mm. in length.
Similar to the male: length of ovipositor slightly shorter than the depth of abdomen at apex.
7 males, Birdlings Flat, Canterbury, December, 1932. (S. Lindsay.)
1 male, Claverley, Conway River, Canterbury.
2 males, Tahunanui, Nelson, October, 1948.
1 female, Amberley, Canterbury, January, 1937 (S. Lindsay).
1 female, Waipara, N. Canterbury, March, 1921 (G. Archey).
1 female, Horseshow Lake, Christchurch, Canterbury, February, 1924 (J. W. Campbell).
3 females, Birdlings Flat, Canterbury, December, 1932 and January, 1933 (S. Lindsay).
1 female, Lake Rotoiti, Nelson, January, 34 (S. Lindsay).
Text-fig. 1.—Fig. 1—Enicospilus insularis Smith. Portion of forewing showing fenestra and adjacent veins. Fig. 2—Enicospilus skeltoni Smith. Portion of forewing showing fenestra and adjacent veins. Fig. 3—Enicospilus skeltoni Smith. Nervellus of hind wing Fig. 4—Enicospilus insularis Smith. Nervellus of hindwing. Fig. 5.—Australophion peregrinus Smith Dorsal view of propodeum, showing carinae. Fig 6—Australophion peregrinus Smith. Foreand hind-wings. Fig. 7—Ophion inutilis punctatus Cameron. Abscissula, Intercubitella, and Nervellus of hindwing. Fig. 8—Ophion oculatus oculatus N.Sp. Abscissula, Intercubitella, and Nervellus of hindwing.
Variation. This polymorphic species shows a wide range in colour and intensity of punctation. From the present material it is not possible to analyse these variations, although it seems that when sufficient material is available for study, E. skeltoni may fall into several distinct sub-species or varieties. In some specimens the yellowish colour pattern is reduced by lack of contrast or is entirely suppressed. In regard to the Hawaiian species of Enicospilus, Cushman (1944: 39) stated that in order to identify species with certainty it is necessary to ignore colour almost entirely, except for a few species, and depend upon structure. Cushman's statement applies equally well to this New Zealand species.
Bionomics: Members of this genus are usually parasitic on the larger caterpillars, and are nocturnal or crepuscular and are frequently attracted to light. Nothing is known of the life history or host relationships of the New Zealand species.
Enicospilus insularis (Kirby)
Ophion insularis Kirby, Trans. Ent. Soc. 1881, p. 46 N. Zeal. Sci., 1884, p. 73.
Henicospilus insularis (Kirby), Morley, Rev. Ich. B.M. Part I, 1912, p. 51.
This species was described from a female from New Zealand presented to the British Museum by Dr. A. Sinclair, R.N., in 1845. The exact locality from which the type was obtained is not known.
This would appear to be a much rarer species than E. skeltoni, as the only specimen I have seen from New Zealand is from the Great Barrier Island, off the East Coast of the northern part of the North Island. Two specimens in the Dominion Museum Collections were obtained from Sunday Island, one of the Islands of the Kermadec Group, 614 miles north of New Zealand.
Male: 16 mm. to 19 mm. in length.
A darker brown species than E. skeltoni, with the inner and outer orbits fulvous, or in one specimen the head is entirely brown. Stigma, costa, and veins dark brown; claws brown; mandibles elongate, upper tooth considerably longer than lower tooth; face and clypeus punctate, clypeus convex; cheeks and caput nitidulous, a few minute punctures scattered over the surface; pronotum closely and distinctly punctate, with somewhat larger punctures on the scutellum; mesopleurae and metapleurae closely punctate with obsolete aciculations on the mesopleurae: propodeum with the transverse basal carina not well developed and gently curved forward at the centre; basal area closely and distinctly punctate; apical area rugose; abdomen with 1st and 2nd tergites sub-equal in length; 3rd tergite ⅘ the length of 2nd, and has an apical depth of ½ its own length: except the 1st tergite, which is nitidulous and only faintly punctate in the post petiole area, the abdomen is finely punctate.
Forewing: 1st abscissa of radius sub-straight, very faintly sinuate, thickened basally to slightly beyond the middle; proximal, distal and marginal scleromes present (Text-fig 1, Fig. 1); proximal sclerome large, sinuated below basal quarter of radius, distal sclerome situated about midway along the 1st abscissa of radius; marginal sclerome well separated from proximal sclerome and extending around the distal border of fenestra, not quite reaching the radius; transverse cubitus ⅓ the length of 2nd abscissa of cubitus, the latter vein ⅓ longer than the 2nd recurrent: nervulus may be perpendicular or slightly oblique, may or may not be interstital with basal.
Hindwing: Abscissula nearly three times the length of intercubitella; nervellus (Text-Fig. 1, Fig. 4) slightly post-furcal, upper abscissa almost perpendicular and broken ⅓ from bottom.
1 male, Sunday Island, Kermadec Islands, 1908.
1 male, Sunday Island, Kermadec Islands, May, 1940 (Coll. Lukins).
1 female, Great Barrier Island. New Zealand, November, 1940 (D. Spiller).
Genus Australophion Morley 1912
Australophion Morley, Rev. Ich. B.M. Part I, 1912, p. 30.
Morley (1912) erected this genus for Smith's Ophion peregrinus described in 1876 from two males taken in Canterbury, South Island. Morley defined the genus as follows:—
“Mandibles vertical; clypeus apically truncate, hind coxae and metathorax not short: metapleurae tuberculiform, apophyses acute and emitting carinae to apex: scutellum often subpyramidal and not margined: nervellus centrally intercepted; ocelli small: discoidal vein geniculate and emitting elongate ramellus: radius of forewing both basally and apically straight, of hindwing basally curved: cubital cell with no comeous spots and but a small glabrous area: tarsal claws, very closely pectinate: body strongly nitidulous.
This genus, as previously pointed out, is very close to Ophion, differing in its greatly inflated scutellum, sub-alar tubercles and metapleurae. Morley considered it closely allied to Stenophthalmus Szepligeti, but differing in its shorter cheeks, exareolate metathorax, not usually short 2nd segment, but especially remarkable in the strongly convex metapleurae. Cushman (1947:440) correctly pointed out that this genus, despite the remarkable inflation of the metapleurae, scutellum and sub-alar tubercles, is hardly more than subgenerically distinct from Ophion.
I have about 40 specimens belonging to Australophion Morley. The majority comprising specimens on loan from the Auckland, Dominion and Canterbury Museums. These specimens have been variously determined as Ophion peregrinus, Ophion ferrugineus and Ophion inutilis. Morley, who had examined the types of peregrinus and ferrugineus in the British Museum, considered Smith's peregrinus 1876 to be the same as the one described two years later, in 1878, as ferrugineus by the same author.
It would appear from Smith's descriptions that his two species refer to a light and dark form respectively. In 1947 Cushman described another species A inflatus from a unique male taken at Egmont, N. Island, by R. J. Tillyard (Type U.S.N.M. No. 57603). Recently two males taken in Canterbury (type locality of Smith's Ophion peregrinus) were sent to Dr. Mueseback, of the U.S. Nat. Mus. for comparison with Cushman's type of A. inflatus. Through the courtesy of Dr. Mueseback, my request was handed to Dr. B. D. Burks, who is the Museum's specialist in the Ichneumonidae, and to whom I am very grateful for the following comparative comments:—
|(1) Discoidella diverging from nervellus above its middle.||(1) Discoidella diverging from nervellus below its middle.|
|(2) Propodeal costulae and areola well-marked, distinct.||(2) Propodeal costulae and areola virtually wanting only vaguely indicated.|
|(3) Spiracles of petiole slightly projecting laterally.||(3) Spiracles of petiole not projecting.|
|(4) Post-petiole almost rectangular, 1 1–3 times as long as wide.||(4) Post-petiole wider at apex than base, 1 ½ times as long as its maximum width, and almost twice as long as its minimum width.|
|(5) Legs entirely yellowish-tan with bases of femora slightly darkened.||(5) Coxae, trochanters and femora brown, tarsi yellow.|
|(6) Antennal scape and pedicel tan, flagellum very dark brown.||(6) Entire antennae entirely dark-brown (original description says it is black, but I would not call it black).|
|(7) Thorax and abdomen yellowish-tan, flagellum very dark brown.||(7) Thorax and abdomen dark brown with yellow markings.|
Dr. Burks expressed the opinion that the differences were not very great, and it may be that the two species would eventually be found to intergrade.
From the material available here I am able to confirm Dr. Burks' opinion, and have little hesitation in synonymising Cushman's A. inflatus with Smith's A. peregrinus. The evidence on which this synonymy is based is presented under the heading “Variability” of this species.
Australophion peregrinus (Smith)
Ophion peregrinus Smith, Trs. Ent. Soc. Lond. 1876, p. 487.
Ophion ferrugineus Smith, Trs. Ent. Soc. Lond. 1878, p. 2.
Ophion peregrinus Smith, Hutton, Cat. N.Z. Dipt., etc., p. 125, 1881.
Ophion ferrugineus Smith, Hutton, Cat. N.Z. Dipt. etc., p. 125, 1881.
Ophion maoricum D.T. 1902, Vol. 2, p. 573.
Australophion peregrinus (Smith) Morley, Rev. Uch. B.M. Part I, p. 30, 1912.
Australophion inflatus Cushman, Proc. U.S. Nat. Mus., Vol. 96, p. 440, 1947. (New Synon.)
Head ferrugineous, outer orbits widely inner orbits very narrowly flavous, the external flavous markings do not usually extend to dorsal margin of eyes. Mandibles except teeth are testaceous, flagellum of antennae brown to very dark brown; teeth mandibles and claws dark red brown; thorax ferrugineous except the sub-alar tubercles, tegulae, and large mark on the posterior-ventral corner of the mesopleurae, the posterior three-quarters of the metapleurae, scutellum and post-scutellum bright flavous; anterior legs lighter brown than posterior legs; abdomen infuscated, especially towards apex.
Face and clypeus minutely punctate, in some cases the clypeus is more sparsely punctate and the punctures are somewhat larger; mandibles very slightly narrowed towards apex, and punctate; malar space narrow, and malar suture distinct; teeth sub-equal or the lower tooth slightly narrower at base, and more acutely pointed, cheeks very finely and sparsely punctate, almost glabrous; moderately wide, nearly equal in width, to width of eye in profile; pronotum very minutely punctate (sparsely); mesonotum sparsely and minutely finely punctate; notaulices not deeply impressed posteriorly; scutellum without lateral carinae.
Male: A ferrugineous species with bright flavous markings on the head and except at the very base, punctate, and convex; mesopleurae punctate except for a raised area adjacent to the posterior border under base of forewing; the sub-alar tubercle inflated with punctures widely separated; the upper half of the mesopleurae is uneven, with two raised areas beneath the sub-alar tubercle; metapleurae, tuberculate and sparsely punctate; propodeum (Text-fig 1, Fig. 5) nor-
mally with a transverse basal carina, two short longitudinal and parallel carinae enclosing the areola, and a posterior semicircular carina joining a lateral cristula on each side, and two lateral longitudinal carinae from base to cristulae and continuing to apex. The above description applies to those individuals in which the carinae described above are obsolete or entirely wanting. The surface of the propodeum is punctate, very rarely finely rugose; abdomen slender, not particularly elongated, spiracles of petiole may be either slightly projecting laterally or not; postpetiole varies in shape, sometimes rectangular or somewhat wider at apex.
Forewing (Text-Fig. 1, Fig. 6): 1st abscissa of radius straight and not thickened; intercubitus a little shorter than 2nd abscissa of cubitus, the 2nd recurrent vein three-quarters as long again as the 2nd abscissa of cubitus; discocubitus with distal arm subequal with proximal arm, the former evenly curved, the latter straight; at the junction of the two arms which is angular, a distinct and usually long ramulus projects towards the base of the basal vein, nervulus not interstitial with basal.
Hindwings (Text-fig 1, Fig. 6): Abscissula, evenly curved at base, about three times as long as intercubitella; nervellus postfurcal, upper arm perpendicular, broken at, or about the middle.
20 females and 36 males were examined, from Otago, Canterbury, Nelson, Wellington and Auckland. The species appear to be generally distributed throughout both the North and the South Islands of New Zealand. It may be somewhat more plentiful in the South Island.
Variability of A. peregrinus
There is a wide variation in colour of A. peregrinus the majority of specimens are either light-brown or dark-brown, intermediate shades, however, occur. The light-form was probably the one originally described by Smith, in 1876, from Canterbury; the same author later (1878) described the dark-form from Otago as a different species, and in 1947 Cushman again described the dark-form from a unique male from Mt. Egmont, in the North Island of New Zealand, collected by the late R. J. Tillyard. A long series of specimens show a striking gradation of shades between the dark-brown and light-brown forms. In the case of the light-form the legs may be entirely light-brown, and in some individuals the darkening of the coxae, trochanters, and femora is very noticeable. The darkening of the stigma, thorax and antennae, is to some extent correlated with the darkening of the basal joints of the legs. There are two specimens in the Hudson Collection, at the Dominion Museum, with the basal ⅔ of hind femora black, also another specimen in the Dominion Museum Collections, from Dunedin with similarly coloured hind femora, but in all other respects they are similar to the typical form. The light and dark forms of A. peregrinus may eventually be correlated with seasonal, geographical distribution or host relationship of the species, but there is no useful purpose to be gained, in the present state of our knowledge of this species, to recognise sub-specific rank within the species.
The discoidella diverges from the nervellus either above, at, or below the middle. The following Table (No. 1) gives the relative occurrence of the position on the nervellus from which the discoidella diverges, in both the light and dark forms.
|Nervellus broken at||Above Middle.||Middle.||Below Middle.|
|Dark form %||25.0||37.5||37.5|
From the above Table, it is seen, that in the majority of dark forms the nervellus is broken at or below the middle, while in the light form, it is broken at or above the middle, although in both forms the nervellus may be broken, above, at or below the middle.
The propodeal costulae (carinae) show considerable variation in degree of development. Some individuals show the costulae well developed and clearly marked, the areola entire, while in others the costulae and areola are virtually wanting, except that the lateral and posterior costulae, are usually present; in the majority of individuals the costulae are more or less indicated.
The post-petiole varies in shape from rectangular to 1 ½ times as long as its minimum width. The length/maximum breadth ratio of the post-petiole area shows gradations similar to that of other characters, ranging from 1.0 (as long as wide apically) to 2.16 (or over twice as long as wide apically). The average ratio for the specimens measured is 1.54, which is about 1 ½ times as long as its apical breadth. The post-petiole is usually wider at the apex than it is at the base, but in one individual in which the post-petiole is as long as wide apically, the basal width is equal to the width at apex.
The number of joints in the flagellum of the antennae varied from 52 to 58 in the dark form and from 53–56 in the light form. The relative length of the 1st and 2nd joints of the flagellum is given by the ratio 1st joint/2nd joint. The dark-brown form gave a ratio from 1.5 to 1.6, the light form from 1.2 to 1.5. In this case the 2nd flagellum joint in the dark form was relatively shorter in relation to the 1st joint, than in the light form.
Seasonal Distribution of A. peregrinus
The relative seasonal abundance of adults of A. peregrinus was obtained from date of capture of specimens. Collections made over the past 40 years were tabulated by months and the graph constructed from the resulting frequency distribution. The total number of specimens from which data were obtained was 51 The resulting frequency distribution is shown in Text-Fig. 2. Their main flight period would appear to extend from November to February, and another during April. The number of specimens on which the graph is based is small, and so, can only serve to indicate the probably seasonal distribution of adult A. peregrinus
Genus Ophion Fabricius 1798
Three species of this genus have been described from New Zealand, O. inutilis, Smith 1876, O. inutilis Smith 1878 and O. punctatus, Cameron 1898. The widespread O. luteus Linneaus was recorded by Fabricius from New Zealand, but this species has not been recorded or specimens seen from New Zealand since. It may safely be excluded from the New Zealand list as it was probably recorded by mistake. Dalla Torre (1898) considered Smith's description of O. inutilis 1876 and O. inutilis 1878, as referred to separate species and renamed the 1878 species, O. insulicalo Reference to the synonymy of these species has been referred to in a previous section of this paper. Over 50 specimens of Ophionini referable to the
present genus have been examined. The indigenous species of Ophion are extraordinarily difficult to separate, due to what appears to be speciation. Characters which may be used with advantage to differentiate the species of closely allied genera, are of little use in this genus, because of the range of variability. A series of individuals clearly shows colour and morphological gradations which constitute differences within a single species or population, that are greater than those between separate or related species of other groups. On the present materials it is not practicable to weigh and interpret the various characters that could be selected. The New Zealand members of Ophion, clearly demonstrate a fundamental fact in systematics, that populations and not only specimens are being classified. After a considerable time has been spent in searching for constant morphological characters to separate these superficially (colour and habitus) distinct forms that occur within this genus, it is considered desirable to deal with two main groups, which are at once separated by distinct and constant characters, and to deal with the various forms as sub-species within these two groups, which are here considered as widely distributed Polymorphic species. The two groups considered here as species are:—
Ophion inutilis Smith 1876 and.
Ophion occulatus n.sp.
There are five sub-species of O. inutilis and one sub-species of O. occulatus.
When further material is available for study and with improved techniques, constant morphological characters may be found, in which case the sub-species dealt with here may be with some justification considered as distinct species, with little disturbance to the nomenclature. The present procedure is further justified when it is considered that nothing is known of the ecology, distribution and host relationship of the New Zealand Ophion.
Mr. G. J. Kerrich, of the Commonwealth Institute of Entomology, London, who kindly compared Smith's and Cameron's types in the British Museum with the three typical forms (A, B and C) I sent him from New Zealand gives the following interesting comments:—
“On the basis of structure, sculpture and colour, I feel pretty confident that your species C agrees with the female type of inutilis Smith 1878. The only difference I find is that in your specimen the abdominal segments are relatively a little shorter and stouter. The male type of punctatus Cam. may be the same species, but I am less confident of this. The head is more narrowed behind the eyes, the tentorial impressions are larger, and the genal costa meets the hypostomal distinctly further from the lower mandibular base, almost as far as the distance from the lower mandibular base to the eye. Moreover, the malar space is very decidedly shorter. The first, fourth and perhaps second differences might be due to sexual dimorphism as to whether the distance that the genal costa meets the hypostomal behind the lower mandibular base could be, I can only leave it to you to judge in accordance with your own knowledge of the genus. In the type of inutilis Smith 1878 the antennae are somewhat darkened in about apical two-fifths, much as in your species C, and are nothing like black. In the unique type of punctatus they are both broken beyond the pedicellus. Your specimen has the ocelli rather smaller and more remote from eyes than in the type of inutilis 1878, and decidedly more so than in the male type of punctatus.
“On general appearance, particularly the form of the propodeum, also the paler colour than in the foregoing, I should say that your species B agrees with inutilis Smith 1876; though the decided traces of lateral keels of the area posteromedia in your specimen are not or scarcely discernible in the type, which has also the ocelli smaller and the genal costa meeting the hypostomal just a little more acutely and a little nearer mandibular base.
“In your species B and in the type of inutilis 1876 the area pleuralis of the propodeum is bulging in a manner that possibly suggests that of Australophion, but not to anything like the same degree. In inutilis 1876 the ocelli are small: in Australophion they are much larger, the lateral ocelli separated from the eyes by about half their longer diameter; but they are decidedly not so large as in Eniscospilus antennatus Morley mss. (Queensland and Tasmania) not even as in the European ramidulus L.
“Your species A is larger, has the ocelli relatively larger than in B and C, the malar space much shorter, the dorsal furrow of the pronotum more deeply and sharply impressed, the propodeum of different general appearance, the petiolar segment more uniformly explanate, and the nervellus intercepted low down. I do not find it in our collection.”
The specimens (B and C) sent to Mr. Kerrich, were carefully selected as typical forms, and after receiving the above comments an examination of a series of specimens showed gradation in all the characters mentioned, and failure has
attended all effort to discover any constant morphological character that may be used to segregate these two forms.
The following key (except couplet No. 1) has been constructed with difficulty, and it should be fully realised that it refers to the more typical members of the respective sub-species. If due allowance be made for the wide variation in the characters used, it may serve to separate the majority of individuals into their respective sub-species.
Key to N.Z. Species and Sub-species of Ophion
|1. Ocelli small (Text-Fig. 3, Fig. 11), the posterior ocelli about their diameter or more from the eyes: width of malar space as great or greater than the basal width of mandibles||O. inutilis Smith||2|
|Ocelli large (Text-Fig. 3, Fig. 12), the posterior cell usually ¼ to ¾ their diameter from eyes: width of malar space not as great as width of mandibles at base||O. oculatus n.sp.||7|
|2. Thorax entirely brown without black markings||3|
|Thorax brown, marked sparsely or extensively with black||4|
|3. Entirely red-brown, with light stramineous costa, stigma, and veins||O. inutilis inutilis Smith|
|Dark brown, with dark brown costa and dark testaceous stigma and veins||O. inutilis punctatus Cam.|
|4. Pronotum, metapleurae and propodeum entirely brown||5|
|Pronotum, meopleura propodeum, metapleura entirely black||6|
|5. Mesonotum testaceous with stramineous vittati: basal fovea of propodeum, flagellum transverse costae extending from scutellum to base of hindwings; mesosuteus of sternum deeply infuscated: propodeum finely rugosely punctate: Posterior coxae glabrous||O. inutilis vittatus n. sub.sp.|
|Mesonotum without stramineous vittati: basal fovea of propodeum and sternum black; propodeum more coarsely rugosely punctate; posterior coxae densely punctate||O. inutilis intermedius n.sub.sp.|
|6. Mesopleurae, metapleurae and propodeum black with brown markings: posterior coxae entirely brown||O. inutilis nigravarium n.sub sp.|
|Mesopleurae, metapleurae and propodeum entirely black: posterior coxae basally black||O. inutilis distinctus n.sub sp.|
|7 Spiracles of 1st abdominal segment projecting strongly laterally, post petiolar area ⅔ as wide apically as long. Thorax rugosely punctate; stigma and costa light testaceous||O. oculatus rugosus n.sub.sp.|
|Spiracle of 1st abdominal segment not projecting laterally: post petiolar area under ¾ the width at apex as long; with width at apex and base subequal: thorax not rugosely punctate stigma, and costa dark brown||O. oculatus oculatus n. sub.sp.|
Ophion inutilis Smith 1876.
Ophion inutilis inutilis Smith 1876.
Ophion inutilis Smith. Trans. Ent. Soc. Lond., 1876. p. 478.
Female. A ferrugineous species 7–11 mm. in length.
Face and clypeus finely punctate, cheeks and caput, minutely punctate: malar space wide sub-equal to width of mandibles at base: genal costa join hypostomal costa at a distance from base of mandibles about equal to molar space; mandibles only slightly narrower at apex than at base: teeth equal in length. Cheeks wide, slightly buccate, their width not quite width of eye in profile. posterior ocelli not
quite their diameter from the eye, and about the same distance from each other, and about their diameter from occipital carina: pronotum, and pleurae very closely punctate, punctures small and in places contiguous: metapleurae not so densely punctate: propodeum rugosely punctate, evenly round dorsally, with usually a short, obsolete, transverse median carina near base. Abdomen short and apically moderately deep. Length of ovipositor about half the depth of abdomen at apex.
1 ♀ Manawatu, N. Island (Entom. Res. Stn. Coll.).
1 ♂ Wellington, January, 1924 (Dom. Mus.).
1 ♀ Whitanui, November, 1916 (Cawth. Inst. Coll.).
1 ♂ Paiaka (Ent. Res. Stn.)
1 ♀ Chatham Islands, January, 1924 (Cawth. Inst. Coll.).
Ophion inutilis punctatus Smith, 1878.
Ophion inutilis Smith, Trans. Ent Soc Lond, 1878, p. 2.
Ophion punctatus Cameron, Mem. Manch. Lit. Boc., 1898, p. 34.
Ophion insulicola Dalla Torre 1902, Catal. Hynenoptera, p. 192.
Ophion punctatus Morley, Rev Ichneum. B.M, Part I, 1912, p. 65.
Female. A dark-brown species with inner orbits widely, and mandibles fulvous; antennae dark brown throughout, darker towards apex. Legs yellowish brown, claws darker.
Face, clypeus and mandibles punctate. caput and cheeks nearly impunctate, nitidulous; internal orbits shallowly emargmate opposite internal base, parallel towards clypeus. Mandibles very slightly narrower at apex than at base, teeth equal in length, malar space wide, nearly equal width of mandibles at base: width of cheeks, usually slightly less than width of eye in profile; faintly buccate: genal costa meets the hypostomal at about ½ malar space, or nearer from base of mandibles ocelli relatively small, posterior ocelli about their diameter from eyes and a similar distance from each other, and up to twice their diameter from occipital carina: pronotum, and mesopleurae, evenly punctate, punctures on mesonotum somewhat smaller and closer together: scutellum more sparsely punctate: mesopleurae, convex, normally not quite so much so in the typical sub-species; distinctly punctate, but punctures more widely spaced than on mesopleurae posterior coxae minutely punctate: abdomen 1st tergite glabrous, ⅓ longer than 2nd, the latter subequal to 3rd tergite in length, hind and forewing venation similar to typical sub-species, nervulus usually oblique, and not interstitial with basal vein, which is gently curved before joining the medius: discocubitus and adjacent veins and cells as figured (Text-Fig. 3, Fig. 10): post nervulus broken by subdiscoideus about the middle: abscissula faintly bent, nearly three times the length of intercubitella. nervellus, slightly postfurcal, under abscissa perpendicular, and broken at lower 3rd. Length, 8–11 mm.
2 ♀ Wellington. (Cawth Instit. Coll.).
2 ♀ ♀ Aniseed Valley, Nelson, December, 1923. (A. Philpott.) (Cawth. Inst. Coll.).
3 ♀ ♀ Broad Bay, Otago (G. Howes), November, 1916. (Auck. Mus.).
Text-fig. 3—Fig 9—Ophion oculatus oculatus N. Sp Discocubital area of forewing. Fig 10—Ophion inutilis punctatus Cameron. Discocubital area of forewing. Fig. 11—Ophion inutilis inutilis Smith. Dorsal view of head showing relative size and position of ocelli Fig. 12—Ophion oculatus N.Sp Dorsal view of head showing relative size and position of ocelli.
1 ♂ Dallington, Christchurch (E. S. Gourlay). (Cawthron Inst. Coll.).
1 ♂ Opoho, Dunedin, January, 1924 (G. Howes). (Auck. Mus Coll.)
1 ♀ Arthur's Pass, January, 1943 (E. S. Gourlay). (Cawth. Inst. Coll.)
1 ♀ Canterbury (Wakefield Coll. Cant. Mus.). Ohakune, November. 1919 (J. W. Campbell). (Auck. Mus.)
1 ♂ Cape Saunders, November, 1922. (Auck. Mus.)
Ophion inutilis vittatus, n.sub.sp.
A testaceous and strammeous sub-species; with distinct testaceous vittati on mesonotum. Hand with the inner orbits and mandibles and scape testaceous: flagellum deeply infuscated, black towards apex pronotum and two wide median stripes on anterior of mesonotum, joining at centre and posteriorly testaceous Tegulae, anterior border of mesopleurae, an obscure patch on posterior border stramineous. The mesosuteus of sternum, sclerite at base of forewing, the transverse costae extending from apex of scutellum to base of hindwing and the propodial fovea infuscated, the remaining parts of thorax, and legs testaceous; the anterior tarsi lightly infuscated. Venation and other characters similar to typical form. Length, 13 mm. Length forewing, 10 mm.
1 ♀ Mt. Ida, Febr. 1922 (Auck. Mus.). 1 ♀ Mt. Maungatua, Febr. 1924.
Ophion inutilis nigra-foveatus n.sub.sp.
A ferrugineous sub-species with inner orbits flavous, a black patch on frons and at base of mandibles: width of malar space about 1 ¼ times the width of mandibles at base: thorax ferrugineous, except for the following areas, which are black: posterior and dorsal margins of pronotum; mesopleural fovea, patch under forewing, anterior and ventral margins of metanotum, scutellar fovea, lateral excavations of scutellum and post-scutellum, basal fovea of propodeum, apex of propodeum, and sternum: abdomen is ferrugineous with the ventral area black, and clouded irregularly on tergites with black: tegulae flavous, costa and stigma testaceous.
1 ♂ (Holotype) Cleughearn, November, 1919 (Cawth. Inst).
Ophion inutilis intermedius n.sub.sp.
Male. Length, 10 mm.
A rugosely punctate brown sub-species, with the following parts of thorax black: at base of forewing: transverse carinae extending from apex of scutellum to base of hindwing: propodeal fovea: mesopleural fovea and continued under base of wings: an irregular spot on ventral area of mesopleurae, and the sternum.
1 ♂ (Holotype) Otago, October, 1923.
Ophion inutilis nigravarius n.sub.sp.
Male. Length, 11 mm.
Face flavous, frons and caput black: mesonotum with a short, median longitudinal band of black on anterior slope, remainder brown infuscated lightly, centrally and laterally: pronotum with a central infuscated stripe: scutellum and post-scutellum brown, lateral excavations black, mesopleurae black, with a wide longitudinal brown stripe. Metapleurae black, with the centre brown: propodeum
black, with a large patch of brown on each side anteriorly: hind coxae entirely brown: scape and flagellum black-brown: sternum entirely black.
1 ♂ (Holotype) “The Hump”, Fabr. 1919. (Cant. Mus. Coll.)
Ophion inutilis distinctus n.sub.sp.
Male. Length, 11 mm.
This sub-species may be distinguished by the entirely black, mesopleurae, metapleurae and propodeum: mesonotum brown with a wide lateral stripe each side and a wide, short median longitudinal stripe on anterior slope: legs, stigma, costa and flagellum inner orbits and basal portion of vein of both fore and hindwings flavous.
1 ♂ Homer Tunnel, S.W. Otago, 21st Jan. 1946 (Coll. J. T. Salmon).
Ophion occulatus n.sp.
Ophion occulatus occulatus n.sub.sp.
Male and Female.
A dark brown median size species: characterised by the following combination of characters, malar space short, about equal to or shorter than ¾ the width of mandibles at base: cheeks, sub-buccate, a little less in width than width of eye in profile: ocelli relatively large, posterior pair about ½ or less their diameter from eye, and usually a little nearer to each other; and separated from the occipital carina by a space about equal to their diameter: pronotum, and mesopleurae, finely punctate; metapleurae punctate on upper half, impunctate ventrally, conspicuously convex: abdomen with 2nd tergite just under ⅗ the length of 1st tergite, and subequal with 3rd tergite: forewing with basal vein slightly sinuated, interstitial or sub-interstitial with nervulus, which is nearly vertical, seldom oblique: the discocubitus and adjacent veins see fig. (Text-Fig. 3, Fig. 9). Hindwing with abscissula, slightly bent and nearly four times the length of intercubitella (see Text-Fig. 1, fig. 8) Nervellus as figured (Text-Fig. 1, Fig. 8).
Length, 12–15 mm.
1 ♀ Auckland, December, 1911 (G. Howes). (Aust. Mus.)
2 ♀ Ohakune, December, 1919 (J W. Campbell) (D.S.I.R O)
1 specimen (damaged), Arthur's Pass, Fabr., 1920 (E. S. Gourlay). (D.S.I.R O. Coll.)
1 ♂ Auckland, December, 1911 (G. Howes). (Cawth. Inst.)
1 ♀ Jack's Pass, Hanmer, November, 1932 (S. Lindsay). (Cant. Mus.)
1 ♀ The Wilderness, January, 1946 (J. T. Salmon). (Dom. Mus.)
1 ♂ Waitaki, S. Canterbury, March, 1917. (Auck. Mus)
1 ♂ Woodhaugh, Dunedin, January, 1920 (G. Howes). (Auck. Mus.)
1 ♂ Alexandra, Otago, December, 1914 (Fenwick Coll.). (Dom. Mus.)
1 ♂ — Peal Forest, December, 1917 (S Lindsay). (Cant. Mus)
1 ♀ Leith, Dunedin, Fabr., 1920 (G. Howes). (Auck. Mus.)
Ophion oculatus rugosa n.sub.sp.
Female. Length, 15 mm.
Thorax rugosely punctate, the lateral carinae of scutellum, low and inconspicuous: scutellum rugosely punctate, not nitidulous and sparsely punctate as in typical sub-species. ♀ (Holotype) Arthur's Pass, West Coast, S. Island. Febr., 1940 (J. T. Salmon). (Dom. Mus.)