Tabanidae (Diptera) of New Zealand By I. M. Mackerras Queensland Institute of Medical Research, Brisbane [Communicated to the Nelson Institute by Dr. D. Miller; received by the Editor March 12, 1956.] Abstract The relationships of the New Zealand Tabanidae lie with Australia and South America; there is no evidence that any of their ancestors entered the country from the north. There is evidence of recent speciation, and possibly hybridization, which makes precise definition of the species difficult. Three tribes are represented, each by a single subgenus, namely: Pangoniini—Ectenopsis, subgen. Paranopsis (1 species). Scionini—Scaptia, subgen. Pseudoscione (6 species). Diachlorini—Dasybasis, subgen. Protodasyommia (9 species). There may be three or four additional species of Protodasyommia, but the material available is inadequate for description. Two new names are proposed: Scaptia (Pseudoscione) milleri, sp. nov., and Dasybasis (Protodasyommia) thereviformis, nom. nov. for viridis Krober nec Hudson. The following synonymy is established or confirmed: hirticeps Nowicki = S. adrel (Walker); brevipalpis var. palpalis Kröber = S. ricardoae (Hutton); impar Walker and gravis Hutton = D. sarpa (Walker); pseudobratrankii Kröber = D. bratrankii (Nowicki). Therioplectes difficilis Kröber has not been recognized, and Tabanus sordidus Walker is deleted from the New Zealand list. The New Zealand tabanid fauna is isolated and interesting. It was last reviewed by Kröber (1931), who followed the complex and often unnatural classification that had been developed by Enderlein. He recognized eighteen species and two varieties, divided among seven genera, of which two were Australian, one Palaearctic, and four endemic. Enderlein (1935) proposed a fifth endemic genus (Glaridapha) for the species that had been included in the Palaearctic genus. A further review seems to be desirable now, first, because the classification of the family has been considerably changed since 1931, and secondly, because there are inconsistencies which make some of Kröber's descriptions difficult to follow. The present work was made possible by the kindness of Dr. David Miller, of the Cawthron Institute, Nelson, who sent me the whole of his considerable collection, and also arranged for the collections of the Canterbury Museum, Christchurch, and the Dominion Museum, Wellington, to be sent to me. These included Hudson's and Hutton's types, and Mr. H. Oldroyd, of the British Museum (Natural History), has compared material sent to him with the types in that Institution, so that nearly all of the described species have been satisfactorily authenticated. I am most grateful to Dr. Miller, Mr. R. R. Forster, of the Canterbury Museum, Miss Beverley A. Holloway, of the Dominion Museum, and Mr. Oldroyd for their help. I would also express my thanks to the Editor of the Australian Journal of Zoology for permission to re-publish Text-fig. 1 and most of Text-fig. 2; to my wife for help with the drawings; and to Miss Barbara McGilvery for help in many ways in the preparation of the paper. Composition of the Fauna The present arrangement of the Tabanidae (Mackerras, 1954, 1955) is shown in Table I, from which it is apparent that the New Zealand fauna is a restricted one, and that its relationships have to be examined at the subgeneric level.
Table I.—Composition of New Zealand tabanid fauna. Known from New Zealand Subfamilies and Tribes Genera and Subgenera No. of Species Pangoniinae Pangoniini Ectenopsis (Paranopsis) 1 Scionini Scaptia (Pseudoscione) 6 Philolichini None Scepsidinae None Chrysopinae Bouvieromyiini None Chrysopini None Rhinomyzini None Tabaninae Diachlorini Dasybasis (Protodasyommia) 9 Haematopotini None Tabanini None Ectenopsis has been recognized so far only in Australia (but not Tasmania) and New Zealand. It has near relatives in South and North America, but none in Africa or the Oriental region. The New Zealand species is sufficiently divergent to require a subgenus to itself, but it nevertheless clearly belongs to a fairly compact Australian-South American complex. Pseudoscione is the most widely distributed subgenus of Scaptia, having been found in South America, Australia (including Tasmania), New Guinea, and New Zealand. The New Guinea element is clearly derived from the Australian; but the New Zealand species show more points of resemblance to forms that occur in South America than to Australian members of the subgenus. Dasybasis is limited to South America, southern Africa, Australia (including Tasmania and a small northern extension), and New Zealand. The generalized Diachlorini, to which it belongs, are extremely difficult to classify at the generic and subgeneric levels, and all that can be said of the New Zealand species is that their relationships lie with Dasybasis rather than Cydistomyia, and that they are sufficiently distinctive to require separate recognition within the genus. There is no evidence that any of these genera entered New Zealand from the north. Ectenopsis (subgenus Ectenopsis) reaches its northern limit on Moa Island in Torres Strait; Scaptia (concolor group of Pseudoscione) extends into the mountains of New Guinea; and Dasybasis is represented by three coastal species, one in Aru Islands and New Guinea, one in Manus and Bougainville, and one in New Caledonia. As with Scaptia, the relationships of the last three lie with Australia rather than New Zealand, and no annectant species have been found. By contrast, the few northern Tabanidae that have invaded the Pacific pass clearly above New Zealand. The Australian genus Cydistomyia is strongly developed in New Guinea, with 35 rather characteristic species. I know at present of three from the Solomon Islands, six from New Caledonia, one from Loyalty Islands, and three from Fiji, and all show evident affinity with the Papuan element. “Neobolbodimyia” veitchi Bezzi from Fiji is not closely related to the true Neobolbodimyia nigra Ric. of Aru Islands and New Guinea, but is in all probability a separate offshoot from Cydistomyia
Table II.—Generalized Members of Southern Tribes of Tabanidae. South America Southern Africa Australia Tasmania New Zealand Pangoniini + – ++ – + Scionini (Scaptia) +++ – ++++ + + Bouvieromyiini + +++ +++ – – Diachlorini ++++ ++ ++++ ++ ++ stock. The genus Tabanus forms a well defined “Oriental” element in the Australasian fauna. It is moderately represented by 32 species in New Guinea and 20 in Australia, but tails off with four in the Solomon Islands, one in New Hebrides, none in New Caledonia* A curious, contrasting feature of the New Caledonian fauna is the occurrence of two species of the Ethiopian-Oriental pangoniine genus Philoliche. or Loyalty Islands, one in Fiji, and one in Samoa. Neither Tabanus nor Cydistomyia, nor anything obviously derived from them, is known from New Zealand. The inference from all this is that the whole of the New Zealand tabanid fauna was derived from an extensive “Antarctic” radiation, which has influenced different southern countries to different degrees. These differences are indicated in Table II by a series of plus signs, as listing the numbers of known species might give an exaggerated impression of precision. I do not believe that these differences should be given too much weight in palaeogeographical speculation. So far as New Zealand is concerned, they may simply reflect a limited range of environments into which the original tabanid fauna could radiate, and limited opportunities for it to escape the rigours of the Pleistocene glaciation. Tasmania presents an interesting parallel, with its small endemic fauna of Scaptia and Dasybasis, evidence of recent invasion, or possibly re-invasion, from the mainland, and absence of Bouvieromyiini. which reach their southern known limit on Flinders Island, in Bass Strait. New Zealand has been too isolated since the Pleistocene for recolonization to have occurred, but there is enough evidence of recent speciation to suggest rehabilitation of a previously depressed fauna. Systematic Review The subfamilies and tribes of Tabanidae have been defined in an earlier paper (Mackerras, 1954). It is unnecessary to repeat the details here, because the subgenera that occur in New Zealand may be readily distinguished by the following simple key, and essential tribal characters can be incorporated in their definitions. Key to New Zealand Genera of Tabanidae 1. Hind tibiae with spurs; ocelli large and prominent; vein sc bare. (Pangoniinae) 2 Hind tibiae without spurs; ocelli, when present, small and inconspicuous; vein sc setulose, at least below (Tabaninae) Dasybasis (Protodasyommia) 2. Eyes bare; palpi longer than shaft of proboscis; smooth, parallel-sided species Ectenopsis (Paranopsis) Eyes hairy; palpi much shorter than shaft of proboscis; more hairy, rotund species Scaptia (Pseudoscione) While the genera are easy to distinguish, the species present a much more difficult problem. There are pairs or groups of closely related forms, separated by quite minor, but apparently fairly constant, differences in colour or pattern, width of frons. shape of antennae or palpi, and so on, and there is also, at least in some species, an appreciable degree of general variability; so that one is often at a loss to know whether one.
is dealing with species, races, or mere variants. This phenomenon is common in the Tabanidae, but it is more than usually evident in New Zealand. The species are still evolving and differentiating vigorously in this small fauna. It is possible, too, that some of the apparently better differentiated forms may still be able to interbreed, and that some of the difficult and anomalous specimens seen in collections may be hybrids between them. There are most interesting problems here for the local entomologist, who can study fresh material and observe the behaviour of the species in the field. An immediate, practical consequence, however, is that it is extremely difficult to determine exactly what forms older authors had before them when they wrote their descriptions. The prime purpose of the present paper is to attempt to provide this basic information, so the criteria of identification, material examined, and variations encountered, will be stated clearly under each species. For the rest, generic synonymy will be noted only insofar as it concerns New Zealand species; specific synonymy and location of types will be listed as completely as possible; and, to conserve space, citation of strictly taxonomic references will be limited to papers not already catalogued by Kröber (1931) and Miller (1950). The following species is deleted from the New Zealand list. Tabanus sordidus Walker, 1854. Mr. Oldroyd has informed me that the type female “is not from New Zealand, in spite of its label. It is a Hybomitra, and seems to me to be the North American Hybomitra affinis (Kirby), which… has priority”. Subfamily Pangoniinae Genus Ectenopsis Macquart, 1838 Genotype: Originally monotypic for Chrysops vulpecula Wiedemann, 1828, from unknown country (now known to be Australia). Subgenus Paranopsis Mackerras, 1955 Subgenotype: Monotypic for Apatolestes lutulentus Hutton, 1901, New Zealand. ♀. Smooth, long-bodied species. Eyes bare. Ocellar tubercle raised and prominent, with large ocelli. Frons wide, tomentose, without callus. Face not unusually projecting. Third antennal segment with basal four annuli widened and more or less completely fused, apical four clearly defined and forming a well differentiated style. Palpi longer than shaft of proboscis, curved, tapering, somewhat flattened, without lateral bare area or concavity. Proboscis shorter than head height; labella large and soft. Hind tibial spurs unusually large and powerful. Wing with vein sc bare; R4 with strong appendix (an almost universal character in the tribe, but rare in the New Zealand fauna); cell R5 open. Genitalia with ninth tergite a continuous transverse bar; cerci truncate apically; caudal ends of spermathecal ducts slender, moderately chitinized, without mushroom-like expansions. ♂. Unknown. It may be expected to have an undivided, shield-like ninth tergite and a bifid hypopygial style. Characters of the Australian subgenera Ectenopsis and Parasilvius are combined in a most interesting way in this subgenus, but it has a somewhat distinctive habitus of its own, and is further differentiated by its very large hind tibial spurs. Hutton was not wildly wrong in allotting lutulentus to Apatolestes, a primitive Nearctic genus of the same tribe; but Enderlein and Kröber misinterpreted Pseudotabanus, which is a chrysopine subgenus with somewhat similar antennal characters. Ectenopsis (Paranopsis) lutulenta (Hutton). Text-fig. 1. Apatolestes lutulentus Hutton, 1901. Type ♀, from? Christchurch, New Zealand, in the Canterbury Museum, Christchurch. Hutton doubted the locality, but there is a more recent specimen from Christchurch also in the Museum, so it may be correct.
Fig. 1.—Ectenopsis (Paranopsis) lutulenta (Hutt.), ♀; head; antenna of type; hind tibial spur; eighth sternite; terminal segments (dorsal view). From Mackerras (1955), by permission of the Editor, Australian Journal of Zoology. Pseudotabanus lutulentus (Hutton). Kröber, 1931, p. 70; Miller, 1950, p. 70. Ectenopsis (Paranopsis) lutulenta (Hutton). Mackerras, 1955, p. 473. Material Examined: The type (which is in excellent condition) and three other females. A long-bodied, almost uniformly grey species, with wide frons, blackish brown third antennal segment, brown legs, and faintly greyish wings with yellowish brown veins. Length, 15–16 mm. ♀. Head: Eyes “bare” (actually with very short, fine hairs, which are just visible at 35 X). Frons wide (index 2.* The frons index is defined as: Length of frons from vertex to junction with subcallus/Width of frons at its mid-length), slightly diverging, pale creamy grey, with short black hairs; ocellar tubercle grey. Subcallus short, pale creamy grey, with a few short dark hairs in mid-line just below frons. Parafacials and face creamy grey; parafacials with cream hairs; face with short mixed dark and cream hairs on each side below antennae. Antennae: First and second segments light yellowish brown, with brown hairs; third deep to blackish brown, paler at extreme base, with a rounded dorsal angle. Palpi grey basally, more yellowish apically, with short, fairly dense cream hairs, mixed with some brown ones apically. Beard white. Thorax: Scutum and scutellum almost uniformly grey, a little paler anteriorly and on side margins, Hairs on disc semi-appressed, greyish cream; darker erect hairs few and inconspicuous; marginal hairs light yellowish brown in front of wing root, creamy white above and behind it, and mixed creamy white and black on scutellum. Pleurae grey, with creamy white to dull yellowish cream hairs. Legs: Femora dark brown, with predominantly cream to creamy white hairs; tibiae brighter brown, with predominantly dark brown to black hairs; tarsi darker than tibiae, with brown hairs, except for a paler zone ventro-laterally; hind tibial spurs brown, conspicuous. Wings: Uniformly faintly greyish, costal cell not darkened; stigma yellowish brown; veins light yellowish brown, darkening somewhat apically. Abdomen: Almost uniformly grey, and covered with creamy white hairs. There is a tendency to development of large, slightly paler, median triangles in the type, and the cream hairs are mixed with black to a variable degree in the sublateral parts of the second and subsequent tergites. Marginal hairs short, cream to greyish white. Venter grey, with apices of the sternites paler; hairs creamy white, except for a few black ones near mid-line on fifth and subsequent sternites. Distribution. South Island: Christchurch (Clarke), also a female collected 8.3.1928.? North Island: Hutton thought his specimen might have come from Taranaki.
Two females without locality label in col. Miller differ from the type in being slightly darker; having the basal four annuli of the third antennal segment almost completely amalgamated; hairs on first and second antennal segments and palpi dark brown; legs darker; the lateral fringe on hind tibiae entirely dark; wings more brownish; hairs on dorsum of abdomen predominantly black, except on pale median triangles and hind margins of the second to the fourth tergites. They agree in other respects, and I think merely represent a variation. Fig. 2.—Scaptia (Pseudoscione) lerda (Walk.), ♀ ♂: head and frons of ♀; proboscis and palp of ♂; ninth tergite and hypopygium of ♂; eighth sternite and terminal segments of ♀, antenna of ♀; hind tibial spur of ♀. A, pupal aster of ♂ S. adrel (Walk.) at same magnification as antenna and tibial spur. Mostly from Mackerras (1955), by permission of the Editor, Australian Journal of Zoology. Genus Scaptia Walker, 1850 Genotype: Pangonia aurata Macquart, 1838, Australia, by designation of Coquillett, 1910. Subgenus Pseudoscione Lutz, Araujo and Fonseca, 1918. Subgenotype: Diatomineura longipennis Ricardo, 1902, Brazil, by designation of Fairchild, 1950. ♀. Stoutly built, usually tomentose and hairy flies. Hairs on eyes usually dense and conspicuous. Ocellar tubercle well defined, but not conspicuously raised; ocelli well developed. Frons slightly to moderately diverging, relatively narrow (index about 3 to 5), tomentose, without callus. Face bulging forward, but not produced into a snout. Third antennal segment subulate, 8-annulate. Palpi much shorter than the shaft of the proboscis, sometimes very short and wide, with a well marked, lateral bare area or concavity.* The palp is often rotated, so that the bare area appears to be dorsal. Proboscis about one and a quarter times head height, with well developed labella. Hind tibial spurs well developed, slender. Wing with vein sc bare; R4 usually curved, occasionally angulate and with short or rudimentary appendix; cell R5 open or closed. Genitalia: ninth tergite an undivided, transverse bar; cerci obliquely truncate apically; caudal ends of spermathecal ducts lightly chitinized, without mushroom-like expansions. ♂ Usually darker and more hairy than female. Eyes meeting in mid-line; upper facets not enlarged. Palpi slender and pointed, or rod-like, with a subapical dorsolateral bare area. Genitalia: ninth tergite large, undivided, forming a prominent shield over the other parts; aedeagus and coxites smooth; style simple, finger-like, rounded at tip.
This definition is based on New Zealand species. Australian members of the subgenus have a wider, more diverging frons, usually less protuberant face, longer proboscis, and differently shaped palpi in both sexes; those from South America tend to have a more projecting face, more slender proboscis, and smaller labella. The New Zealand species fall into two groups. Most are tomentose, with indications of pale scutal lines, and an abdominal pattern of white to golden hairs; these resemble their relatives from other regions. S. montana is quite different. It is a small, plump, dark, semi-metallic species, which looks superficially as if it should be placed in the Australian subgenus Myioscaptia. The style of the male hypopygium is, however, rounded at the tip, and it must be regarded simply as an aberrant member of the New Zealand complex. Key to New Zealand Species of Scaptia Females 1. Small (9–10 mm), rotund, semi-metallic blackish species, with short, wide palpi (Text-fig. 3, L); legs almost uniformly brown montana (Hutt.) Larger species, with at least scutum tomentose and an evident adornment of pigment or pale hairs on abdomen 2 2. Black species, with post-alar tuft and median spots on abdominal tergites cream; palpi and legs deep brown to black; hind tibiae with entirely black hairs posteriorly and laterally: palpi long (Text-fig. 3, G) adrel (Walk.) Brown species, with brown palpi and legs, and at least some reddish brown hairs posteriorly and laterally on hind tibiae 3 3. Palpi long (Text-fig. 3, H); hairs on hind tibiae entirely reddish to yellowish brown; larger species (11–15 mm) 4 Palpi short and wide, leaf-like (Text-fig. 3, I, J); hairs on hind tibiae usually partly black; smaller species (9–12 mm) 5 4. Frons narrow (index more than 4); post-alar tufts and paler abdominal hairs rich golden lerda (Walk.) Frons wider (index 3–3.5); post-alar tuft and paler abdominal hairs dull cream ricardoae (Hutt.) 5. Wing pattern strongly marked; frons and face light creamy fawn; dorsocentral vittae on scutum well defined to posterior margin; venter with darker median zone and fairly numerous black hairs on at least the third and subsequent sternites milleri, sp. nov. Wing pattern more diffuse; frons and face darker brownish cream; pale scutal vittae poorly defined posteriorly; venter without median darker zone or black hairs on third and subsequent sternites brevipalpis Krob. Males 1. A small (8–11 mm), uniformly semi-metallic blackish species, with uniformly brown legs montana (Hutt.) Larger, black or brown species, with well defined abdominal pattern 2 2. Posterior and lateral hairs on hind tibiae predominantly reddish to yellowish brown; brown species, with reddish to yellowish brown tibiae 3 Hairs on hind tibiae predominantly or entirely black; black or brown species, with black or reddish brown tibiae 4 3. First and second antennal segments light brown; abdominal hair tufts rich golden; femora usually brown lerda (Walk.) First and second antennal segments grey; abdominal hair tufts dull cream; femora usually blackish ricardoae (Hutt.) 4. Larger (12–16 mm), black species, with cream hair tufts; hind tibiae dark brown to black adrel (Walk.) Smaller (10–12 mm), more brownish species, with dull cream to golden hair tufts; hind tibiae reddish brown 5 5. Wing pattern strongly marked; apical pale fringe on second abdominal tergite full width of segment; venter with indefinite darker median stripe bearing black hairs on at least the third and subsequent sternites milleri, sp. nov. Wing pattern more diffuse; apical pale fringe on second abdominal tergite limited to median zone; venter without darker median stripe, and with at most a few scattered black hairs on third and subsequent sternites brevipalpis Kröb.
Scaptia (Pseudoscione) adrel (Walker). Text-figs. 2, A; 3, A, G. Pangonia adrel (White ms.) Walker, 1850. Type ♂ ♀, from New Zealand, in the British Museum (Natural History). Scaptia adrel (Walker). Kröber, 1931, p. 61; Miller, 1950, p. 70; Mackerras, 1954, Fig. 7d. Pangonia hirticeps Nowicki, 1875. Type ♂ ♀, from New Zealand, in the Zoological Museum, Cracow University, Poland (stated by Miller, 1950, to have been destroyed during war). Kröber (1931) considered it to be merely a lighter variation of adrel. I can find nothing in his notes nor transcript of Nowicki's description to separate the two. Material Examined: 47 ♂ ♂, 13 ♀ ♀, including specimens returned by Mr. Oldroyd as agreeing with the types in the British Museum. A large to medium sized, black species, with cream thoracic and abdominal hair tufts, dark brown to black legs, black hairs on hind tibiae, and the wings more or less darkened, especially anteriorly. Length, 12–16 mm. ♀. Head: Eyes with long, dense, brown hairs, darker above, paler below. Frons relatively narrow (index 4.5 to 6), yellowish fawn above, more creamy fawn below, sometimes irregularly suffused with brown, and with fine brown hairs; ocellar tubercle Fig. 3.—Scaptia. A-F, frons of ♀ of: A, adrel (Walk.); B, lerda (Walk.); C, ricardoae (Hutt.); D, brevipalpis Kröb.; E, milleri, sp. nov.; F, montana (Hutt.). G-J, palp of ♀ (above) and ♀ (below) of: G, adrel; H, lerda; I, brevipalpis; J, milleri (normal ♀, variant ♂). K-M, montana: antenna of ♀, palp of ♀, palp of ♂. N, palp of ♀ variant of ricardoae. grey to brown, with a tuft of black hairs. Subcallus, parafacials and face creamy fawn; parafacials with creamy hairs; face with mixed yellowish cream and black hairs. Antennae: First and second segments with grey to dark brown tomentum and black hairs above, some yellowish cream hairs below; third dark brown to black, paler at extreme base. Palpi: First segment black, with cream hairs; second (Text-fig. 3, G) relatively slender, brown to blackish brown, with a large, orange-brown bare area surrounded by short black hairs. Beard cream to yellowish cream. Thorax: Scutum dark brown, with some greyish overlay on anterior margin, and indications of paler median and dorsocentral lines and suture. Hairs mixed black erect, and brownish yellow appressed; marginal hairs black in front of wing root, cream above and behind it. Scutellum deep brown, with mainly dark hairs, except at lateral corners. Pleurae light greyish fawn, with cream to yellowish cream hairs. Legs: Femora black, mid and hind with some greyish overlay; with black hairs anteriorly, yellowish cream posteriorly on fore femora, mainly yellowish cream on mid and hind. Fore tibiae and tarsi brown, somewhat yellowish basally, darker dis-
tally; mid darker; hind deep brown to black; hairs dark brown to black, except for a reddish to yellowish brown ventral zone. Wings: Suffused with brown to a variable extent; sometimes almost completely, but tending to leave the discal cell clearer; sometimes with the brown tint more or less restricted to the anterior half of the wing, with some darkening leading back from the dark brown stigma obliquely across the apices of the basal cells. Veins brown; R4 often angulate but with at most a rudimentary appendix; cell R5 usually narrowed, sometimes closed, occasionally widely open. Abdomen: Black, sometimes with more or less reddish to yellowish brown suffusion laterally on the first and second tergites. Hairs black, except for cream tufts in midline on first four tergites, on apical lateral corner of second, and lateral margins of fourth to sixth, leaving a dense black lateral tuft on the third. Venter brown, more or less banded with black, and with the apical edges of the sternites paler; hairs yellowish cream. ♂. Similar to female; but the brown colour at the side of the abdomen more variable, and often so extensive as to reduce the black on the first two or three tergites to a broad median vitta; pale apical hairs on second abdominal tergite usually restricted to a median spot, but sometimes extending as a fringe across the whole width of the segment. Palpi (Text-fig. 3, G) long and slender, brownish black, and with a narrow, elongate, brown dorso-lateral bare area. Distribution.* The general arrangement of localities in this paper is from north to south in each Island. Distribution will be listed from Hutton (1901) or Kröber (1931), only when the identity of the species is evident, and the locality is not represented in the material examined. North Island: Whangarei, January; Parakai (tidal flats), December; Brown's Bay, near Auckland, December (on flowers); Rangitoto Island, near Auckland, December, January; Auckland, February; Mt. Tauhara, December; New Plymouth, August; Okato; Ohakune, January; Aramoho, Wanganui, December; Levin; Silverstream, Hutt Valley, January; Wellington, December, January; Gollan's Valley. South Island: Trio Island, January; Nelson, December, January; Third House, near Nelson, December (reared); Dun Mt., near Nelson, 3,000ft, January, February; Tapawera, near Nelson, January; Waiho Gorge, South Westland, December; Upper Hororata, January; Otago. Scaptia (Pseudoscione) lerda (Walker). Text-figs. 2; 3, B, H. Pangonia lerda (White ms.) Walker, 1850. Type ♂ ♀, from New Zealand, in the British Museum (Natural History). Scaptia lerda (Walker). Kröber, 1931, p. 66; Miller, 1950, p. 70. Scaptia (Pseudoscione) lerda (Walker). Mackerras, 1955, Fig. 28, C, F. Material Examined: 13 ♂ ♂, 12 ♀ ♀, including specimens returned by Mr. Oldroyd as agreeing with the types in the British Museum. A large to medium sized, brown species, distinguished by its elongate palpi, reddish brown legs, entirely reddish brown hairs on hind tibiae, and bright golden thoracic and abdominal hair tufts. Length, 12–15 mm. ♀. Head: Eyes with rich brown hairs above, light cream below. Frons relatively narrow (index 4 to 5.5), bright brown, with a darker patch at junction of lower and middle third, and another around the brown to blackish ocellar tubercle; hairs dark brown, relatively sparse. Subcallus, parafacials and face yellowish to brownish fawn, with yellowish cream hairs laterally, brown ones on most of face, central area bare. Antennae: Basal segments light brownish yellow, with black hairs above and some pale ones below; third reddish brown, paler at extreme base, and darkening almost to blackish brown on apical four annuli. Palpi long and narrow; brown with an orange tint; basal segment with long creamy yellow hairs; second with an extensive lateral bare area, which is often turned upwards, and with short, dark brown hairs around its margins. Beard creamy yellow.
Thorax: Scutum dark brown, with indications of paler median and dorsocentral lines, suture, and lateral margins. Hairs on disc dark brown erect and dull yellowish brown appressed; marginal hairs mixed black and yellowish brown in front of wing, creamy gold above and behind it. Scutellum dark brown, with brown hairs on disc and mainly dull golden ones marginally. Pleurae greyish fawn, with creamy yellow hairs. Legs: Bright to yellowish brown, the femora often somewhat darker than the other segments; hairs mixed brown and yellowish cream on the femora, more definitely golden to reddish brown on the remaining segments. Wings: Similarly marked to S. adrel, but the veins are often more yellowish brown; cell R5 usually narrowed, sometimes widely open. Abdomen: Bright to mahogany brown, irregularly darkened in the median part of the first and second tergites and across most of the width of the remainder. Hairs black, except for median tuft and apical fringe of rich gold on the first to the fifth tergites, more or less interrupted sublaterally by dark hairs on the first, third and fifth; apical lateral corners of tergites with bright golden tufts, except for the completely black zone on the third. Venter bright mahogany brown, irregularly darkened, and with creamy golden hairs, which tend to form pale apical fringes to the sternites. ♂. Darker than female, and more like male of S. adrel. The median dark markings on the basal abdominal tergites tend to be reduced to spots, and the hair tufts are rich golden in colour; palpi similar in form to S. adrel, but usually not so dark; legs brown, hairs on hind tibiae entirely reddish brown. Distribution. North Island: Brown's Bay, near Auckland, November, December (on flowers); Henderson, December; Manganamu, Lake Taupo, January; Whakapapa, Ruapehu, 4,000ft, January; Wanganui; Korokoro, February; York Bay, near Wellington, February; Wellington, December; Muritai Track, January. S. adrel and S. lerda are distinctive species, which should not be difficult to recognize in the field. In using these notes for identification, it is necessary to remember, however, that fresh material is always more richly coloured and better marked than relatively old museum specimens, on which the descriptions have necessarily been based. Browns, in particular, tend to become yellowish with age, and reddish black to end as a dull red. Old specimens may be freshened, as well as greasy ones cleaned, by immersion over-night in amyl acetate (a method for which I am indebted to Dr. C. B. Philip), but their colours are rarely fully restored. These flies have apparently not been observed to suck blood, although Dr. Miller has reports of stock being disturbed in the presence of S. adrel. Their habits were vividly described by Hudson more than sixty years ago under the general name of “Comptosia bicolor”. He wrote: “This conspicuous species is very abundant in glades throughout the summer, flying with great rapidity, and delighting to suck honey from the numerous shrubs which are in blossom at that time of year. It is a social species, and is usually found in companies of fifteen to twenty individuals, which engage in endless dances, two insects often seizing one another on the wing and then revolving together like a wheel in rapid motion. Their manoeuvres in avoiding the strong gusty wind, so often prevalent in early summer, are also interesting; the insects play upon the wing whilst the air is quiet, but if a breeze springs up they instantly settle on the nearest bush, rising to renew their sports when it is again calm.” Tillyard (1926, p. 359) noted that S. adrel frequented the blossoms of Leptospermum, a habit which is common in Australian species also. Nothing is recorded of their life histories, but there is a bred male of S. adrel, with the pupal skin attached, in Dr. Miller's collection. The aster (Text-fig. 2, A) is remarkable, in that all six lobes are clearly represented, although the dorsal and ventral pairs are greatly reduced. This is the most primitive condition so far encountered in the tribe, the Australian species of Scaptia described (subgenera Scaptia, Pseudoscione and Myioscaptia) and the Nearctic Goniops having “bilobed” asters,
with the dorsal and ventral lobes reduced to low, unarmed swellings, and the lateral lobes usually larger than in S. adrel. Scaptia (Pseudoscione) ricardoae (Hutton). Text-fig. 3, C, N. Comptosia bicolor Hudson, 1892. Preoccupied in Scaptia by Pangonia bicolor Macquart, 1845. No type designated, and Miss Holloway has informed me that there are no specimens in the Hudson collection in the Dominion Museum, nor are any mentioned in the catalogue of its contents. Pangonia ricardoi Hutton, 1901. No type designated. Mr. Forster found specimens identified by Hutton in the Canterbury Museum, but none labelled as type, although Hutton's three other types of Tabanidae are present in the collection. The name is now corrected to ricardoae to comply with the “Copenhagen Decisions,” 1953, paras. 86, 87. Scaptia brevipalpis var. palpalis Kröber, 1931, p. 69. Type ♀, from New Zealand, in the British Museum (Natural History). Females returned by Mr. Oldroyd as agreeing with Kröber's type included one determined by Hutton as Pangonia ricardoi, and agree with his description of that species. Scaptia palpalis Krober. Miller, 1950, p. 70. Material Examined: 22 ♂ ♂ (including three det. Hutton), 9 ♀ ♀ (including those noted above). The nomenclature of this species is difficult. Hudson's description embraced a complex of species, for which the name he used* It was not followed by “n.s.”, and there is no direct evidence that he regarded it as new. is not available. Hutton followed his name by “sp. nov.”, gave a recognizable description of a species he had before him, and listed its distribution; but he recorded “bicolor Hudson… not of Macquart” as a synonym, apparently did not mark a type, and labelled his specimens “Pangonia ricardoi Huds.” He seems to have been confused between two different procedures: describing a misidentified species as new, and proposing a new name for a junior homonym. Nevertheless, I believe that his name is valid, and should have priority over Kröber's. In order to identify this species precisely, I now select, as lectotype of Scaptia (Pseudoscione) ricardoae (Hutton, 1901), a female in the Canterbury Museum, Christchurch, without locality label, but bearing a label “Pangonia ricardoi Huds., det. F. W. Hutton”, to which has now been added a further label, stating that it had been seen by Mr. H. Oldroyd of the British Museum (Natural History), and returned by him as agreeing with the type female of Scaptia brevipalpis var. palpalis Kröber, 1931. ♀. Like S. lerda, with similarly shaped palpi, and entirely red-brown hairs on hind tibiae; but distinguished by the wider, paler frons (index 3 to 3.5); creamy fawn subcallus, parafacials and face; darker third antennal segment, which is quite black in some specimens; duller general colour; and dull cream (instead of bright golden) thoracic and abdominal hair tufts. ♂ Darker than female, and at first I thought that it was a light form of S. adrel, distinguished by the yellowish brown tibiae and more extensive lighter markings on the abdominal tergites. When I came to ally the males with the females, I found from his labels that Hutton had already done so. The male has the hairs on hind tibiae reddish brown, as in the female, and is distinguished from the male of S. lerda by its darker general colour, grey first and second antennal segments, darker palpi, deep brown to blackish femora, and dull cream thoracic and abdominal hair tufts. Distribution. North Island: Recorded from “Wellington (Hudson)” by Hutton (1901); I have seen no specimens that correspond with this record. South Island: Nelson, December; Upper Maitai, near Nelson, December; Grey River, January; Lake Moana, December; Mt. Grey, January; White Rock, Canterbury, December; Upper Hororata, January; Christchurch, December; Peel Forest, December; Ashburton (Hutton, 1901); Dunedin, October; Wallacetown, Otago, January. Several males have scattered black hairs among the reddish brown on the hind tibiae, and four have relatively short palpi, like that illustrated for the male variant
of S. milleri (Text-fig. 3, J). One male (Lake Moana, Tonnoir) in the Canterbury Museum has brown femora, and the basal abdominal tergites almost entirely black. Two females have unusually wide palpi (Text-fig. 3, N), and one of them (Wallacetown, col. Miller) also has relatively prominent post-alar tufts and a distinct hint of gold in the hairs at the sides of the fourth to sixth abdominal tergites; they agree in other respects with ricardoae rather than brevipalpis. Scaptia (Pseudoscione) brevipalpis Kröber. Text-fig. 3, D, I. Scaptia brevipalpis Kröber, 1931, p. 67; Miller, 1950, p. 70. Type ♂, from Mt Earnslaw, ♀ from Ohakune, New Zealand, stated to be in the British Museum (Natural History). Mr. Oldroyd has written of these: “There are no types here, nor even any specimens det. Kröber, but there are specimens labelled by him as lerda which clearly belong to brevipalpis—as if he had changed his mind, but forgotten to alter the labels. There is a male from Mt. Earnslaw and several females from Ohakune, which might be types.” Material Examined: 8 ♂ ♂, 17 ♀ ♀, including specimens returned by Mr. Oldroyd as agreeing with the above. A medium sized, brown species, resembling S. lerda in general coloration, but distinguished from all except S. milleri by the short, wide palpi of the female. Length, 10–12 mm. ♀. Head: Eyes with brown hairs above, paler ones below. Frons medium (index 3.5 to 4), bright yellowish brown, with bright brown hairs; ocellar tubercle deep brown. Subcallus fawn-cream. Parafacials and face light fawn, with fine cream hairs on parafacials, mainly brown ones on face. Antennae: First and second segments creamy yellow, with golden and black hairs; third brown, darkening distally. Palpi (Text-fig. 3, I) short and broad; bright orange-brown, with black marginal hairs, and a few scattered ones on the otherwise bare area. Beard cream. Thorax: Scutum olive-brown, with quite evident greyish fawn median and dorsocentral vittae, which fade behind suture, and more brownish fawn lateral areas, with a brown spot above wing root. Hairs erect dark brown, and quite dense, somewhat appressed light to yellowish brown; marginal hairs mixed dark brown and orange in front of wing root, yellowish cream above and behind it; post-alar tuft conspicuous. Scutellum brown, with yellowish brown to bright brown hairs, including the apical margin. Pleurae light greyish fawn, with mainly cream hairs, which are mixed with dull yellowish ones on the upper mesopleural convexity and in the hypopleural tuft. Legs: Almost uniformly light brown, darkening somewhat on hind tibiae and all tarsi; hairs cream to yellowish on the femora, yellowish cream and brown on fore and mid tibiae, usually mixed reddish to yellowish brown and black on hind tibiae, darker on the tarsi. Wings: Similar to S. lerda and S. ricardoae, but the brown colour tends to suffuse the anterior half of the wing more uniformly; veins brown; R4 angulate, and often with a short appendix. Abdomen: Bright mahogany brown, with an interrupted median black vitta on first three tergites, and more extensive darkening more posteriorly. Hairs black on disc; long and golden on median triangles, apical edges, and lateral margins of all tergites, except for a black patch at basal lateral corners of third. Venter creamy fawn, with cream hairs, particularly at edges of sternites, and with few or no dark hairs. ♂ Darker than female, and with less evident scutal vittae. Palpi relatively short, more or less rounded at tip (Text-fig. 3, I); lateral and posterior surfaces of hind tibiae with black hairs which are longer and less dense than those of S. adrel; abdomen with median apical golden tufts on second and subsequent tergites; there is no continuous pale fringe on second, as in S. milleri; lateral margin with apical golden tufts at corners of tergites, except the third, which has entirely black fringe; venter as in female.
Distribution. North Island: Whakapapa, Ruapehu, 4,000ft, January; Ohakune, January; Mt. Holdsworth (lower slopes), January; Wainui-o-mata, near Wellington, December (Kröber, 1931); Wellington. South Island: Dun Mt., near Nelson, 2,500 to 3,000ft, January, February; Lake Rotoroa, January; Gowan, Murchison, January; Lake Moana, December; Arthur's Pass, 3,000ft, January; Waiho Gorge, December; Mt. Earnslaw, 3,500ft, January (Kröber, 1931); Wallacetown, Otago., December. Most of the males have a few red-brown hairs among the black ones on the hind tibiae; they are fairly numerous in one specimen, leading, in this respect, towards the males of ricardoae with some black hairs in this position. Six females have entirely red-brown hairs on hind tibiae; in one female the hairs on first and second antennal segments are almost entirely black, and the third segment is very dark. Scaptia (Pseudoscione) milleri, sp. nov. Text-fig. 3, E, J. Scaptia bicolor Krober, 1931, p. 65, ♂, nec Hudson, 1892. Krober's description fits the ♂ of this species very well. Types: Holotype ♀, from Dun Mt., Nelson, 3,000ft, 27.1.1921, A Philpot, and allotype ♂, from Woodend, Otago.,—12.1909, D. Miller (No. 470), in the Cawthron Institute, Nelson. Material Examined: 5 ♂ ♂, 5 ♀ ♀. I hesitate to describe a new species in this variable group, but the present form is constantly distinct from S. brevipalpis in the short series before me. Length, 9–12 mm. ♀. Distinguished from S. brevipalpis by more uniformly light creamy fawn frons; darker antennae, with entirely black hairs on first and second segments; somewhat larger, wider palpi (Text-fig. 3, J); more conspicuous dorsocentral scutal vittae, which are continuous to scutellum; less conspicuous, more whitish post-alar tufts; creamy white abdominal hair tufts; darker venter, with a vague dark vitta and quite numerous dark brown hairs in the median area of the third and subsequent sternites; and by the markings of the wings. The costal cell is not darkened, the brown colour forming an irregular pattern extending obliquely backwards from the stigma, across the apices of the basal cells, enclosing the discal cell, and narrowly along the veins in the distal and posterior part of the wings. The general appearance is somewhat mottled, reminiscent of that characteristic of the subgenus Plinthina in Australia. ♂ Recognizable primarily by the wing markings and dark hairs on the venter, both of which are similar to those of the female. The pale apical fringe on the second abdominal tergite is continuous and well defined. The palpi are short in three of the males, as in S. brevipalpis (Text-fig. 3, I) but darker; they are more slender (Text-fig. 3, J) in the other two. Distribution. North Island: Ohakune, January, February (in nest of crabronid wasp identified as Rhopalum carbonicolor);* Dr. Miller doubts the reliability of this identification. Wellington, January. South Island: Dun Mt., near Nelson, 3,000ft, January, February; Waiho Gorge, South Westland, February; Upper Hororata, January; Woodend, Otago, December. Scaptia (Pseudoscione) montana (Hutton). Text-fig. 3, F, K, L, M. Pangonia (Corizoneura) montana Hutton, 1901. Type ♀, from Mt. Arthur, Nelson. New Zealand, col. Broun, in the Canterbury Museum, Auckland. Although described as bare, the eyes of the type are distinctly hairy. Corizoneura montana Hutton. Kröber, 1931, p. 70; Miller, 1950, p. 69. Scaptia (Pseudoscione) montana (Hutton). Mackerras, 1955, p. 495. Material Examined: 9 ♂ ♂, 5 ♀ ♀, including the type. A small, compact, distinctive, semi-metallic, black species, with wide frons, short, wide palpi, and uniformly brown legs. Length 8–11 mm. ♀. Head: Eyes with long, but not dense, light brown hairs. Frons relatively wide (index 2.5 to 3), uniformly reddish brown, with yellowish brown hairs; in some specimens there is a rubbed area above the subcallus suggestive of an incipient callus;
ocellar tubercle dark brown. Subcallus, parafacials and face brownish orange, the parafacials and face with creamy yellow and brown hairs. Antennae relatively short and thick (Text-fig. 3, K); basal segments brownish orange, with black hairs; third reddish brown on the basal half, darkening to blackish brown apically. Palpi short and wide (Text-fig. 3, L), bright brownish orange, with short brown marginal hairs. Beard yellowish cream. Thorax: Scutum and scutellum blackish grey, lighter grey near anterior margin on each side; lateral margins yellowish brown, contrasting with the dark colour of the disc. Hairs erect black, and rather sparse appressed dull cream; marginal hairs short, mixed black and yellowish in front of wing root, creamy yellow above and behind it, rather dull and straggling on the edge of the scutellum. Pleurae light yellowish brown, greyish on lower part of mesopleural area, with yellowish cream hairs, except for a dull yellowish group on mesopleural convexity. Legs: Almost uniformly bright, somewhat yellowish brown, darkening slightly on hind tibiae and tarsi; hairs mainly yellowish cream on femora, brown elsewhere; hind tibial fringes inconspicuous. Wings: Lightly suffused with yellowish brown; veins light brownish yellow to brown; R4 gently curved, without angle or appendix; cell R5 widely open or slightly narrowed. Abdomen: Semi-metallic black, duller on first tergite owing to overlying greyish tomentum, and somewhat suffused with brown at lateral edges of first two or three tergites. Hairs brown or black, except for rather indefinite creamy white at apical lateral corners of fourth and fifth tergites; those at lateral edge of third entirely dark. Venter bright brown, darkened somewhat across middle of sternites, and paler on their apical edges; some dark hairs in median zone, but mostly cream, especially along apical edges of sternites. ♂ Darker and more shining than female, and with almost entirely black hairs, except for those on the eyes, the mixed creamy white and black beard, and white on apical lateral corners of some abdominal tergites and variably on venter; legs darker than in female, with entirely black hairs, except ventrally on tibiae and tarsi; palpi (Text-fig. 3, M) short, brown, and with oval, orange-brown bare area. Distribution. North Island: Ohakune, January, February (in nest of crabronid wasp identified as Rhopalum carbonicolor); Kaitoke, December; Hutt Valley, January; Wellington, January; Gollan's Valley, December. South Island: Balloon Hut, Mt. Arthur Tableland, February: Mt Arthur (Hutton, 1901). Fig. 4.—Diagram indicating possible evolution of New Zealand species of Scaptia. Evolution of the Species Judged solely by the ease with which they can be placed in the cabinet, there are four well defined “species” in the material examined, namely, adrel, lerda, brevipalpis and montana. Their relationships are indicated diagrammatically by heavy lines in Text-fig. 4. Structurally, they are quite homogeneous, including the characters of the genitalia, although there are minor variations in the shape of the ninth tergite of
the male and the eighth sternite of the female. S. montana stands apart in appearance, but it resembles S. brevipalpis structurally, and I believe that the whole group had a common ancestry, which had diverged somewhat from the direct ancestry of the Australian species on the one side and the South American species on the other. S. milleri is a more recent derivative from brevipalpis stock, and this also is indicated by heavy lines in the diagram. It has been taken in company with S. brevipalpis, no annectant forms have been seen, and it has therefore been given specific rank. On the other hand, the males do vary towards S. adrel. It seems possible that milleri may be reproductively isolated from brevipalpis, but not completely so from the more divergent adrel. One cannot, in the present state of our knowledge, undertake laboratory breeding experiments with Tabanidae, but useful evidence might nevertheless be obtainable by careful collecting and observation in the field. S. ricardoae presents a somewhat different problem. It shows evidence of affinity in three directions—with brevipalpis, adrel, and to a lesser degree with lerda. Such variation as has been noted is chiefly towards brevipalpis. It could have originated as an adrel × brevipalpis hybrid, but appears to have become fairly well stabilized, so the existence of some isolating mechanism would seem to be indicated. This possible relationship is shown by broken lines in the diagram. I do not think that ricardoae is a subspecies of lerda, in the ordinary sense of the term, but the almost complete separation of their territories, if confirmed, might explain the absence of intermediates. These speculations are offered purely as a possible basis for field studies. There is no doubt that hybridization does occur in nature. Dobrotworsky (1955) has reported an interesting recent case in Australian mosquitoes. Writing of the Culex pipiens group, he notes that globocoxitus and molestus are as a rule ecologically isolated from one another, “but in 1953 and 1954 a permanent breeding place of globocoxitus was found to harbour also molestus in the winter; during the period of joint occupancy, hybrids were very common; they were absent during the summer.” The same sort of thing may be occurring in Tabanidae, and New Zealand, with its limited fauna of closely related forms, may offer special opportunities to throw light on this problem. Subfamily Tabaninae Genus Dasybasis Macquart, 1847 Genotype: Originally Monotypic For Dasybasis Appendiculata Macquart, 1847, Australia. Subgenus Protodasyommia Enderlein, 1925 Subgenotype: Originally Monotypic For Protodasyommia Loewi Enderlein, 1925, New Zealand. Aphopeas Enderlein, 1930. Originally monotypic for Aphopeas priscum Enderlein, 1930 (= Tabanus truncatus Walker, 1850), New Zealand. Discussed below. Leptotabanus Kröber, 1931, p. 75. Monotypic for Tabanus transversus Walker, 1854, New Zealand. Distinguished from Aphopeas only by the male having the upper facets of the eyes markedly enlarged. Thereviotabanus Kröber, 1931, p. 77. Monotypic for Thereviotabanus viridis Kröber, 1931, New Zealand. Distinguished from Aphopeas only by its unusual colour. Therioplectes Kröber, 1931, p. 79, nec Zeller, 1842* The true Therioplectes is a Palaearctic genus of the tribe Tabanini. The New Zealand species included here are distinguished from Protodasyommia only by the males lacking ocelli; this character is variable; P. loewi is difficult to separate specifically from “Therioplectes” sarpa. Glaridapha Enderlein, 1935. Type Tabanus bratnankii Nowicki, 1875 (lapsus for bratrankii), by original designation. Proposed for the New Zealand species erroneously included by Kröber in Therioplectes. Neoleptotabanus Miller, 1945. Nom. nov. for Leptotabanus Krober, 1931, nec Lutz, 1914. Fairchild (1950) has pointed out that Leptotabanus Lutz is a nomen nudem, and therefore does not invalidate Kröber's use of the name.
♀. Eyes varying from unequivocally hairy to practically bare; brown or green, unbanded, in relaxed specimens. Frons usually of medium width (indices usually 3 to 4, but ranging from 2 to 5), diverging, tomentose; ocellar tubercle not clearly differentiated, but ocelli often represented by one to three small spots or punctures; callus usually small and pale, sometimes relatively large and dark, but always narrower than frons and restricted to its lower half. Subcallus tomentose, with or without hairs. Face and parafacials normal. Antennae of normal tabanine form; third seg- Fig. 5.—Dasybasis (Protodasyommia) sarpa (Walk.), ♀ ♂: head of ♀; ninth tergite and hypopygium of ♂; eighth sternite and terminal segments of ♀; antenna and palp of ♀ (left). ♂ (right). ment with the basal four annuli fused and widened to form a “basal plate”, which is sometimes rounded above, or sometimes with a dorsal angle or hump; apical four annuli distinct and forming a clearly differentiated style. Palpi usually slender, somewhat compressed, tapering and rounded at tip. Proboscis short and stout; labella large and soft. Thoracic spiracles widely open, with small lips. Hind tibiae without apical spurs. Wings: basicosta (subepaulet) without setulae; vein sc setulose, at least below; R4 usually smoothly curved, sometimes more or less angulate and with an incipient appendix; cell R5 widely open. Genitalia: Terminal segments dorsoventrally compressed; ninth tergite divided into two widely separated triangular plates; caudal ends of spermathecal ducts with characteristic mushroom-shaped expansions (Mackerras, 1955, Fig. 2, I). ♂ Eyes contiguous; upper facets usually small, but somewhat enlarged in one species and markedly so in another. Ocellar tubercle usually clearly visible at vertex, sometimes distinctly raised, and often with more or less clear indications of three ocelli. Palpi short, usually more or less acorn-shaped. Genitalia: Ninth tergite clearly divided in mid-line; style truncate apically in dorsal view. As at present recognised, the subgenus is restricted to New Zealand, but there is a small group of Australian species which may possibly belong to it. The species fall into two groups, which I would be inclined to treat as separate subgenera, if the distinguishing characters were more reliable in related, larger subgenera in other countries. They are not completely satisfactory even here, and I believe that the groups had a common origin, so amalgamation or separation does not affect the zoogeographical discussion at the beginning of this paper. They are: truncata group: Eyes appearing bare or faintly hairy at 15 X; frons of female relatively wide (index usually less than 3); first antennal segment small, not expanded dorsally; basal plate of third rounded or with obtuse dorsal angle (Text-fig. 6, K-M); wings tapering apically, and with relatively narrow cell Cu2 and anal lobe; vein sc bare or irregularly setulose above (always setulose below in both groups). North and South Islands. sarpa group: Eyes unequivocally hairy at 15 X; frons of female relatively narrow (index 3 to 5); first antennal segment larger, usually more or less hood-like dorsally;
basal plate of third with sub-basal dorsal angle or hump (Text-fig. 6, N-Q); wings more rounded apically, and with wider cell Cu2 and anal lobe; vein sc setulose on most of length above. North Island. If subgeneric distinction is considered desirable, the name Aphopeas End. (syns. Leptotabanus Kröb., Thereviotabanus Kröb,. Neoleptotabanus Miller) is available for the truncata group, and Protodasyommia End. (syns. Therioplectes Kröb. nec Zell., Glaridapha End.) for the sarpa group. Both groups present problems of speciation similar to those discussed above under Scaptia, although variation within populations appears to be greater, and definition of the species is correspondingly more difficult. I have found no accounts of their behaviour, other than Tillyard's (1926, p. 359), statement that Tabaninae “are found resting on rocks in river-beds; they seldom appear to bite”, and Hudson's (1892) note that D. viridis may be taken in similar situations to the Scaptias. Dr. Miller (personal communication) has confirmed that they are not pests of any note, and he knows of none attacking man. Hudson recorded D. viridis as breeding in moss on tree-trunks in the forest, but his account is unconvincing, and Dr. Miller has informed me that he has observed this species ovipositing in damp sand on the margin of a river. Key to Species of truncata Group 1. Wings with brown shading at apices of basal cells, apex of discal cell, and fork of R4+3; a light brown species, with pale grey median abdominal vitta or triangles; male with upper facets of eyes markedly enlarged, and ocellar tubercle deeply sunken transversa (Walk.) Wings more or less infuscate anteriorly, but without discrete brown markings; colour and pattern otherwise; male with upper facets of eyes not enlarged 2 2. Medium sized (11–13 mm), grey to greenish grey, almost concolorous species; eyes with short but perceptible hairs; ocellar tubercle of male deeply sunken thereviformis, n. nov. Small (8–11 mm), dark grey to blackish species, with pale abdominal pattern; eyes bare; ocellar tubercle of male prominent 3 3. Female with wide frons (index about 2), brown femora, and entirely pale pleural hairs; male with frontal hairs projecting between the eyes truncata (Walk.) Female with narrower frons (index about 3), deep brown to blackish femora, and at least some of mesopleural hairs dark; male without projecting frontal hairs nigripes (Kröb.) Note.—Some males with frontal hairs have dark legs and mesopleural hairs, like nigripes, but they are treated here as variants of truncata. Dasybasis (Protodasyommia) transversa (Walker). Text-fig. 6, A, K. Tabanus transversus Walker, 1854. Type ♂ ♀, from New Zealand, in the British Museum (Natural History). Leptotabanus transversus (Walker). Kröber, 1931, p. 76. Neoleptotabanus transversus (Walker). Miller, 1945, p. 72; 1950, p. 69. Material Examined: 4 ♂ ♂, 5 ♀ ♀, including specimens returned by Mr. Oldroyd as agreeing with the types. A small, easily recognised, brown to greyish brown species, with unusually large stigma, characteristic pattern of brown spots on the wing, and median pale stripe or row of triangles on abdominal tergites. Length, 9–10 mm. ♀. Head: Eyes (relaxed) brown; bare. Frons slightly diverging, index 2.5 to 3, fawn-cream, irregularly suffused with brown, and with brown hairs; vertexal triangle inconspicuous; position of anterior ocellus indicated by a small brown spot; callus small, pale brown, about one-third width of frons at base, more or less triangular, and with a short median extension Subcallus, parafacials and face greyish to yellowish cream; subcallus sometimes with a few fine cream hairs, parafacials and face with cream and some brown hairs. Antennae: First and second segments light yellowish fawn, with some greyish overlay, and with brown to black hairs; basal
Fig. 6.—Dasybasis: A-E and K-M, truncata group; F-J and N-R, sarpa group. A-J, frons of ♀ of: A, transversa (Walk.); B, C, the two possible females of thereviformis, n. nov.; D, truncata (Walk.); E, nigripes (Kröb.); F, sarpa (Walk.); G, loewi End.; H, viridis (Huds.); I, opla (Walk.); J, bratrankii (Now.). K-N, antenna of ♀ (except L, which is of ♂):, palp of ♀ and palp of ♂ of: K, transversa; L, thereviformis; M, truncata; N, loewi. O-R, antenna and palp of ♀ of: O, viridis; P, opla; Q, bratrankii; R, sp. A nr. bratrankii. plate of third bright yellowish brown, style brown. Palpi fawn-cream, with relatively long cream and brown hairs. Beard cream. Thorax: Scutum with ground colour considerably reduced, the appearance being of a grey to greyish fawn scutum, bearing a pair of broad admedian brown stripes, fading behind suture, a pair of broad sublateral brown stripes, interrupted by suture, and a brown patch above wing root. Hairs erect dark brown, and appressed dull cream; marginal hairs predominantly brown. Scutellum grey, paler apically and laterally, with brown and dull cream hairs. Pleurae pale fawn-grey, with cream hairs. Legs: Light brown, darkening a little on tarsi, especially on the fore pair; hairs predominantly cream to yellowish cream on femora and tibiae, predominantly brown on apical parts of tibiae and all tarsi; hind tibial fringes poorly developed, predominantly brown. Wings: Longer than usual, projecting well beyond tip of abdomen; lightly suffused with brown; stigma large, brown. There are distinct brown clouds across apices of basal cells, apex of discal cell, and at fork of R4+5. In some specimens the centres of the cells are a little paler than the general ground colour. Abdomen: Brown; median area and lateral margin of first tergite pale grey; second and subsequent tergites with a variable median pale grey stripe, sometimes reduced to a row of median triangles, and with narrow pale grey apical margins, which widen to form apical lateral triangles. Hairs brown on the darker parts, cream on the paler areas. Venter creamy fawn, with paler apical margins to the sternites, and pale cream hairs. ♂ Similar to female in coloration and markings. Upper facets of eyes markedly enlarged, reddish brown, contrasting with the darker, small, lower and lateral facets; ocellar tubercle small, deeply sunken. Palpi short and relatively broad, yellow, with long cream and some brown hairs.
Distribution. North Island: Kaitaia; Taranga Is. (Kröber, 1931); Auckland; Tangoio Falls, December; Ohakune, February, March, April; Lake Horowhenua, February (Kröber, 1931); Mangatiriri R., Tararua Range; Wellington, February. D. transversa is the only species in New Zealand which looks superficially as if it might have been derived from northern Cydistomyia stock. However, it has so many features in common with the other members of the truncata group, particularly with D. thereviformis, that its relationship to them can scarcely be doubted. Dasybasis (Protodasyommia) thereviformis, nom. nov. Text-fig. 6, B, C, L. Thereviotabanus viridis Kröber, 1931, p. 78; Miller, 1950, p. 70; nec Comptosia virida Hudson, 1892 (q.v.). Type ♂, from New Zealand, in the British Museum (Natural History). Material Examined: 18 ♂ ♂, 2 ♀ ♀, including a male returned by Mr. Oldroyd as agreeing with the type in the British Museum. A grey, almost concolorous species, with short but distinct hairs on the eyes, deeply buried ocellar tubercle in the male, brown legs, and predominantly pale hairs on the abdominal tergites. The male will be described first, as there is some doubt about the associated females. Length: ♂, 10–12 mm; ♀, 11, 13 mm. ♂ Head: Eyes green when relaxed; with short, fine, pale hairs, which are visible at 15 X; upper facets not enlarged. Ocellar tubercle small and sunken, difficult to see. No long frontal hairs between eyes. Frontal triangle and subcallus creamy grey, with only an occasional pale hair. Parafacials and face creamy grey, with mixed dark brown and greyish cream hairs laterally, entirely greyish cream medially. Antennae: Basal segments brown, with a grey overlay and black hairs; third dark brown, paler at base, basal plate often with a more definite dorsal angle than in other members of the group. Palpi light greyish fawn, with long yellowish cream and a few dark hairs. Beard greyish to yellowish cream. Thorax: Scutum almost uniformly olive-grey, paler anteriorly and laterally, but with only faint indications of paler vittae. Hairs erect dark brown and long semiappressed dull cream; marginal hairs predominantly brown in front of wing root, cream above and behind it. Scutellum grey, with some dark hairs on disc and cream fringe. Pleurae pale grey, with greyish white to cream hairs. Legs: Light brown; femora darkened basally, with a greyish overlay in some specimens; fore tarsi also somewhat darker. Hairs creamy white to cream on femora and tibiae, more predominantly brown apically on fore tibiae and on all tarsi; hind tibial fringes mixed cream and brown. Wings: Faintly greyish, costal cell barely, if at all, darkened; stigma brown, inconspicuous; veins light brown. Abdomen: Grey, with the apical edges of the tergites narrowly pale, and with mixed long fine cream and dark brown hairs, the former predominating, especially on apical and lateral edges of tergites. Venter grey, with narrow apical pale margins to the sternites, and long fine creamy white hairs. ♀. The females are both similar to the males, but more definitely grey in general colour, and with a fairly evident median paler triangle on the second abdominal tergite. Hairs on eyes very fine, but visible at 15 ×. Antennae missing. One (Wellington, N.I., Hudson, in the Canterbury Museum) is relatively longbodied (length 13 mm), and has a wide (index 1.8), diverging frons, with a well developed brown callus (Text-fig. 6, B); its palpi (Text-fig. 6, L) are short, and covered with long cream hairs. The other (Waiho, S.I., 26.1.1922, Tonnoir, in the Canterbury Museum) is shorter (length 11 mm), more thick-set, and has a narrower (index 3), nearly parallel frons, with a very small, inconspicuous callus, which is almost entirely covered by tomentum (Text-fig. 6, C); hairs on palpi predominantly dark brown.
In the absence of more material, it is impossible to say what may be the relationship between these females. I cannot separate the males into two series, which might correspond with them. Distribution. North Island: Tiritea R., January; Wellington, January, March. South Island: White Rock, Canterbury, December; Waiho, January; Alexandra (Kröber, 1931). It seems likely that two species may be recognizable in life by having bright green eyes, thereviformis, which is grey in general colour, and viridis, which is more olive and has different antennae (Text-fig. 6, 0). Hudson apparently saw both, although his type series included only the latter. Dasybasis (Protodasyommia) truncata (Walker). Text-fig. 6, D, M. Tabanus truncatus Walker, 1850. Type ♀, from New Zealand, in the British Museum (Natural History). Mesomyia maoriorum Bigot, 1892. Type ♂, from New Zealand, in the British Museum (Natural History). Synonymy by Ricardo, 1915, who noted the presence of black hairs on the subcallus and between the eyes. Aphopeas priscum Enderlein, 1930. Type ♀, from New Zealand, in the Zoological Museum, Berlin. Synonymy by Krober, 1931, on authority of F. W. Edwards. Aphopeas truncatus (Walker). Krober, 1931, p. 72; Miller, 1950, p. 69. Material Examined: 10 ♂ ♂, 9 ♀ ♀, including specimens returned by Mr. Oldroyd as agreeing with the type. A small, dark, brown or grey to blackish species, with wide frons, brown legs, and variable pale markings on abdominal tergites; male with long frontal hairs Length, 8–10 mm. ♀. Head: Eyes dark brown (relaxed); bare. Frons diverging, index 1.8 to 2.2, light grey, often irregularly patterned with brown, and with relatively long and dense dark brown hairs; vertexal triangle indefinite; ocellar tubercle sometimes indicated by a brownish mark, but ocellar spots usually absent; callus dark brown, raised in lateral view, about one-third width of frons, irregularly triangular or diamond-shaped, and with a short median extension. Subcallus projecting, pale to creamy grey, with numerous brown and yellowish cream hairs. Parafacials and face pale to creamy grey, with yellowish cream and brown hairs. Antennae: First and second segments light brown, the first with some greyish overlay, both with brown to black hairs; third dark brown, paler at base. Palpi variable in length and thickness, light greyish to brownish fawn, with relatively long cream and brown hairs. Beard greyish cream. Thorax: Scutum and scutellum brown, sometimes dark, with greyish overlay in sublateral areas anteriorly and laterally in front of wing root, and with more or less well developed pale dorsocentral vittae. Hairs erect black and appressed dull yellowish cream; marginal hairs predominantly brown in front of wing root, mixed brown and dull cream behind it and on scutellum. Pleurae grey, variably marked with brown, and with entirely cream to yellowish cream hairs. Legs: Brown, the mid and hind tibiae yellowish; hairs mainly yellowish cream on basal segments, brown apically on tibae and on all tarsi. Wings: Lightly suffused with greyish brown; stigma dark brown; veins brown. Abdomen: Dark to blackish brown, with a well developed median pale grey triangle and apical margin on the second tergite, and variable, usually less conspicuous, triangles and pale margins on the remainder. Hairs dark brown to black on most of the abdomen, yellowish cream on the pale triangles, irregularly on the apical margins, and more definitely on the apical lateral corners of the tergites. Venter grey, the sternites with irregular basal brown bands; hairs yellowish cream, most conspicuous on the apical edges of the sternites, and more or less mixed with dark brown in the median zone. ♂ Generally darker than female. Upper facets of eyes not enlarged; ocellar tubercle prominent, reaching level of eyes. There is a row of strong black hairs projecting between the eyes on the full length of the frons, which is characteristic
of this species;* A male from Karori, in the Canterbury Museum, lacks frontal hairs, but is otherwise typical. Examination suggests that the hairs may have been broken off, but it could be a variant.; subcallus with conspicuous dark hairs. Palpi small, slender, yellowish brown, with long yellowish cream and black hairs. Distribution. North Island: East Lake Taupo, March; Crow's Nest, February; Silverstream, Hutt Valley, March; Korokoro, February; Makara Bush, near Wellington, November (Kröber, 1931); Karori, near Wellington, autumn; Wainui-o-mata, near Wellington, February; Wilton's Bush, near Wellington, February. South Island: Gouland Downs, near Collingwood, February; Moutere Inlet, near Nelson, December; Nelson, January. Three males, from nest of crabronid wasp identified as Rhopalum carbonicolor, Ohakune, N.I., January, are darker, with brown palpi, blackish brown legs, and brown hairs on the posterior part of the mesopleural convexity. They resemble nigripes rather than truncata in appearance, but have the characteristic frontal hairs of the latter, and are retained here provisionally as variants. It seems possible that they may be hybrids. Dasybasis (Protodasyommia) nigripes Kröber. Text-fig. 6, E. Aphopeas nigripes Kröber, 1931, p. 74; Miller, 1950, p. 69. Type ♂, from Ohakune, New Zealand, in the British Museum (Natural History). Material Examined: 1 ♂, 1 ♀, the male returned by Mr. Oldroyd as agreeing with the type in the British Museum. A small (8–9 mm) blackish brown species, which is distinguished from D. truncata by the following characters. ♀. (Described from a specimen in fair condition, from Porirua, Wellington, N.I., 23.1.1937, in the Dominion Museum). Frons narrower (index 3.3), darker, and bearing short, inconspicuous hairs; palpi longer and more slender, with short dark brown hairs; hairs on posterior margin of mesopleural convexity largely dark brown; legs dark brown, with mainly black hairs; wing with more prominent stigma. ♂. Immediately distinguished by having no long frontal hairs projecting between the eyes, the few that can be detected being short and appressed; upper facets slightly enlarged and a more coppery brown than the lower and lateral facets; antennae and palpi darker, the hairs on the second palpal segment entirely dark; legs darker than in female, the femora being almost black; pleural hairs, wings and abdomen as in female. Distribution. North Island: Ohakune; Porirua, near Wellington, January. One male without locality label in Dr. Miller's collection is entirely black, except for brown tibiae. It has no pale tomentose triangles or apical bands on the abdominal tergites, and all its hairs are brown or black, including those on parafacials, palpi, legs and venter. It may represent a distinct species. Key to Species of sarpa Group Females 1. Dark species, with prominent brownish black callus† Rarely absent as an aberration.; abdomen predominantly black, with well defined pale median triangles on at least some of the tergites 2 Reddish brown to fawn species, with pale, inconspicuous callus; with or without a darker median vitta on abdomen, but pale triangles, when developed, on a brown rather than black ground 3 2. Smaller species (12 mm), with narrow frons (index 5), and pale median triangles on abdomen small, that on second tergite the largest loewi End. Larger species (13–16 mm); frons not so narrow (index 3.5 to 4); pale median triangles on abdomen usually more conspicuously developed on second to fifth or sixth tergites sarpa (Walk)
3. Abdomen with a narrow, interrupted, dark brown median vitta, contrasting with the bright yellowish to reddish brown ground colour opla (Walk.) Abdomen sometimes vaguely darkened in median area, but without strongly contrasting vitta 4 4. Abdomen bright yellowish to reddish brown, with bright pale golden hairs, which sometimes tend to form incipient median triangle opla (Walk.) var. Abdomen dull to fawn brown, with or without cream to greyish median triangles 5 5. Scutum with narrow, dark, median and dorsocentral lines; abdominal tergites with only indications of pale median triangles viridis (Huds) Scutum with pale dorsocentral vittae and lateral margins; pale triangles on abdominal tergites well developed bratrankii (Now.) Males 1. Abdomen dark brown to black; the first three tergites variably yellowish to reddish brown laterally, sometimes reducing the black to a broad median vitta enclosing the pale median triangles on the tergites 2 Abdomen predominantly dull to yellowish or greenish brown; if with a narrow median black vitta, then without conspicuous pale triangles 3 2. A smaller species (10–11 mm); ocellar tubercle large, ocelli differentiated; at least some mesopleural hairs dark; lateral areas of abdominal tergites black, except for the apical lateral triangles loewi End. A larger species (13–15 mm); ocellar tubercle smaller and narrower, ocelli usually not detectable; pleural hairs entirely pale; lateral parts of first two or three abdominal tergites often more or less suffused with yellowish to reddish brown sarpa (Walk. 3. Abdomen bright yellowish brown, with a narrow interrupted median black vitta opla (Walk.) Abdomen dull to paler yellowish brown or fawn; median dark vitta, if present, greenish grey 4. Abdomen generally with greenish grey to olive vitta, indefinite median triangles, and predominantly long pale hairs viridis (Huds) Abdomen uniformly dull brown, with better defined pale median apical triangles on tergites, and sublateral areas predominantly dark haired bratrankii (Now.) Dasybasis (Protodasyommia) sarpa (Walker). Text-figs. 5; 6, F. Tabanus sarpa (White ms.) Walker, 1850. Type ♀, from New Zealand, in the British Museum (Natural History). Therioplectes sarpa (Walker). Kröber, 1931, p. 83; Miller, 1950, p. 71. Glaridapha sarpa (Walker). Enderlein, 1935, p. 235. Tabanus impar Walker, 1850. Type ♂, from New Zealand, in the British Museum (Natural History). Synonymy by Ricardo, 1915; confirmed by Oldroyd (personal communication). Tabanus gravis Hutton, 1901. Type ♀, from Auckland, Broun, in the Canterbury Museum, Christchurch. Synonymy by Kröber, 1931; there are points of difference that are worth further study. Material Examined: 15 ♂ ♂, 7 ♀ ♀, including the type of gravis and specimens returned by Mr. Oldroyd as agreeing with the type of sarpa. A relatively large, dark, variable species, distinguished by possessing a conspicuous dark callus, long slender palpi, and a prominent row of pale median abdominal triangles on a black ground. Length 13–16 mm. The typical form will be described first. ♀. Head: Eyes (relaxed) dark brown; with dense brown hairs, paler below. Frons bulging, diverging, index 3.5 to 4, irregularly brownish lateral to callus, and with dark brown hairs; vertexal triangle moderately defined, with black hairs; ocelli not defined; callus shining brownish black, about two-thirds width of frons at base, elongate-triangular. Subcallus bulging, greyish cream, with numerous cream hairs. Parafacials and face greyish cream, with cream hairs. Antennae: First and second.
segments yellowish brown, with black hairs, third entirely brownish black. Palpi light greyish fawn, with short brown and cream hairs. Beard cream. Thorax: Scutum brownish black, with wide, indefinite, grey dorsocentral vittae, which are interrupted behind suture; lateral margins light brown, with a dark patch above wing root. Hairs erect dark brown, and appressed dull cream; marginal hairs predominantly brown to black in front of wing root, creamy white above and behind it. Scutellum deep brown, grey on apical lateral margins; hairs dark brown on disc, dull cream marginally. Pleurae grey, except for brownish yellow mesopleural convexity; hairs entirely cream. Legs: Femora dark brown, with grey overlay; hairs on fore pair mainly black, on mid and hind mainly cream; tibiae yellowish brown, darkening apically, and with cream and black hairs; hind tibial fringes predominantly pale; tarsi blackish brown, with black hairs. Wings: Slightly greyish, almost clear; stigma dark brown; veins brown. Abdomen: Brownish black; first and second tergites extensively suffused with light brown laterally; second and subsequent tergites with narrow, pale apical margins, and large, clearly defined, pale grey median triangles, which almost reach their anterior margins; seventh entirely dark. Hairs dark brown on the darker parts, cream on the median triangles and apical lateral corners of the second to sixth tergites, and the basal lateral parts of the first and second. Venter dark brown, the apical margins of the sternites cream; hairs cream, mixed with black in the median zone. ♂. Upper facets of eyes slightly enlarged, but not differentiated from lower; ocellar tubercle well developed, usually narrow, and usually without indications of ocelli; parafacials and face darker than in female, the parafacials with predominantly brown hairs; palpi short and wide (Text-fig. 5), brownish yellow, with mixed black and pale golden hairs; beard creamy yellow. Pale hairs on thorax and abdomen creamy yellow to golden. Abdomen with the bright yellowish brown colour more extensive on second and third tergites than in female, reducing the black to a broad median stripe enclosing the pale triangles; fourth and subsequent tergites almost entirely dark, except for the pale median triangles. Distribution. North Island: Auckland; Te Aroha, March; Manganamu, Taupo, January; Okato; Ohakune, March; Rangataua, January; Wanganui; Mangatiriri R., Tararua Range, January; Lower Hutt, February; Day's Bay, near Wellington, February; Muritai Track, February; Gollan's Valley, February. There is considerable variation in shape of callus, hairiness of subcallus, proportion of brown hairs on parafacials and palpi, amount of grey tomentose overlay on first antennal segment, and the degree to which the dark colour of the abdomen is invaded laterally by yellowish brown on the first three tergites. The following forms can be recognized in the material studied: sarpa form: Described above. Distinguishing features: light yellowish brown first and second antennal segments; more slender third antennal segment, with smaller, sharper tooth; paler palpi, with longer cream and brown hairs; tomentum on parafacials and pale hairs on thorax and abdomen more creamy yellow to golden, at least in male; hind tibial fringes predominantly pale; lateral yellowish brown areas on first three abdominal tergites extensive, reducing the black to a broad median vitta. gravis form: First and second antennal segments dark brown, with grey tomentum; third with large, blunter tooth; palpi darker, with entirely dark hairs; tomentum on parafacials and pale hairs on thorax and abdomen generally cream; hind tibial fringes entirely dark; yellowish brown on abdomen reduced to small patches at sides of first and second tergites. var.?: A single female (Ohakune, N.I., January, H.H.) differs from the gravis form in lacking a callus, and in having the pleurae predominantly fawn and the femora brown. It may be merely an aberration.
No single character is reliable, but it does seem generally possible to distinguish the sarpa and gravis forms when their characters are considered collectively. There are no apparent distributional differences between them, so they cannot be treated as subspecies on present evidence. Dasybasis (Protodasyommia) loewi Enderlein. Text-fig. 6, G, N. Protodasyommia loewi Enderlein, 1925; Kröber, 1931, p. 79; Miller, 1950, p. 69. Type ♂, from New Zealand, in the Zoological Museum, Berlin. Material Examined: 3 ♂ ♂, 1 ♀, including a male returned by Mr. Oldroyd as agreeing with a specimen in the British Museum det. Kröber as loewi End. I do not think that there is any doubt of the identity of this species. The males agree very well with Enderlein's and Kröber's descriptions of the type, and the female, allowing for normal sexual differences, agrees with the male. This species is very close to D. sarpa, of which it may be only a variant. It is distinguished as follows: ♀. (Described from a specimen in fair condition, from Titirangi, N.I., 15.4.1943, M. W. Carter, in the collection of Dr. C. B. Philip, Montana, U.S.A.). Smaller (12 mm); frons narrower, index 5; a small brown spot representing the median ocellus; callus smaller; subcallus with short but darker hairs; first antennal segment very dark; palpi (Text-fig. 6, N) much shorter. Abdomen almost entirely dark, and the median triangles relatively small, except on the second tergite. The pleural hairs are all creamy white, and the hind tibial fringes entirely dark. ♂. Like the gravis form of D. sarpa, but distinguished by smaller size (10–11 mm), large, prominent ocellar tubercle, bearing definite indications of three ocelli, darker pleurae, and (in contrast with female) dark brown mesopleural hairs. Distribution. North Island: Titirangi, April; Ohakune; Lake Horowhenua, February (Kröber, 1931). Dasybasis (Protodasyommia) opla (Walker). Text-fig. 6, I, P. Tabanus oplus (White ms.) Walker, 1850. Type ♂, from New Zealand, in the British Museum (Natural History). Therioplectes oplus (Walker). Kröber, 1931, p. 82; Miller, 1950, p. 71. Glaridapha oplus (Walker). Enderlein, 1935, p. 235. Miss Ricardo (1915) was in error in treating gravis Hutton as a synonym of oplus Walk. Material Examined: 3 ♂ ♂, 2 ♀ ♀, including a male returned by Mr. Oldroyd as agreeing with Walker's type. The females agree well with the males, and I therefore designate one, from Auckland, N.I., without further data, but with an identification label “Tabanus oplus Wlk. F. W. Hutton det.”, in the Canterbury Museum, Christchurch, as the allotype female of Dasybasis (Protodasyommia) opla (Walker, 1850). A brown species, with parallel frons, pale callus, yellow-brown legs, and bright yellowish brown abdomen, on which there is typically a narrow, blackish brown median vitta; pale median triangles indefinite or absent. Length, 12–13 mm. ♀. Head: Eyes (relaxed) greenish to brownish black; with short, rather inconspicuous brown hairs. Frons nearly parallel, index 3.5 to 4, brown, with short brown hairs; vertexal triangle indefinite; at least one brown ocellar spot present; callus yellowish brown, one-third width of frons, tapering above to a short extension. Subcallus yellowish brown, with only a few short inconspicuous hairs. Parafacials and face creamy fawn, with short, fine, cream hairs on the parafacials, somewhat darker on the face. Antennae: First and second segment bright brownish yellow, with black hairs above and some pale ones below; third reddish brown at base, darkening to blackish brown; dorsal tooth small but sharp. Palpi bright fawn-yellow, with short brown and cream hairs. Beard cream. Thorax: Scutum and scutellum dark olive-brown, somewhat discoloured; lateral margins and a dorsocentral patch in front of scutellum fawn-brown. Hairs sparse
fine black erect and denser appressed dull yellowish cream; marginal hairs yellowish brown and black in front of wing root, cream above and behind it, and dull yellowish cream on scutellum. Pleurae grey, with fawn patches on mesopleural area; hairs cream to yellowish cream. Legs: Light brownish yellow, the femora darkened somewhat basally, and the tarsi a little darkened distally; hairs bright creamy yellow, merging into brown on the tarsi, and with a brown lateral fringe on the hind tibiae. Wings: Slightly greyish, almost clear; costal cell, stigma, and a trace of colour along Rs brown; veins bright to yellowish brown anteriorly, somewhat darker distally and posteriorly. Abdomen: Bright brownish yellow basally, more definitely brown on the third and subsequent tergites, and with a row of deep brown oblong blocks which form a narrow median vitta, interrupted by the paler apical margins of the tergites, and merging on fifth and subsequent tergites into the brown ground colour. Hairs short, black, rather sparse on discs of second and subsequent tergites; bright creamy yellow on most of first tergite, apical margins of first to sixth, and tending to form vague median triangles on second to fourth; lateral margins with almost entirely cream to yellowish cream hairs. Venter light brownish yellow, the apical margins of the sternites paler, and the hairs yellowish cream, except for a few dark ones near the mid-line, and more extensively on the seventh sternite. ♂. Darker than female. Eyes with dense brown hairs; upper inner facets somewhat enlarged but merging into the others; ocellar tubercle small, but well defined and reaching to level of eyes; subcallus and parafacials more yellowish than in female; palpi yellowish cream, with long yellowish cream and a few dark hairs; femora darker than in female, and with some greyish overlay. Distribution. North Island: Whangaroa, January; Brown's Bay, near Auckland, February; Auckland; Mayor Island, March. Four females (Okato, N.I.; Pokaka, N.I., February; Wanganui, N.I.) differ from the above in having more definitely diverging frons, ocellar spots varying from none to prominent, palpi variably shorter and with greater mixture of pale hairs, and the abdomen without a median vitta, or with only some darkening in this position; the tendency of the pale hairs on the abdomen to form vague median triangles is more marked than in the typical form. These agree with Kröber's (1931) description of the specimens which he treated tentatively as females of opla. The typical form of opla is easy to separate from D. viridis, and will probably prove to be further distinguishable in life by having darker eyes. The variant noted above, however, may provide a link between these two apparently well defined “species”, unless the eye colour proves to be a constant distinguishing feature. Dasybasis (Protodasyommia) viridis (Hudson). Text-fig. 6, H, O. Comptosia virida Hudson, 1892. Type ♂, from Bush Hill, Karori, near Wellington, New Zealand, 10.2.1883, in the Dominion Museum, Wellington. Tabanus viridis (Hudson). Hutton, 1901, p. 13. This emendation appears to be in accord with the “Copenhagen Decisions”, 1953. Therioplectes viridis (Hudson). Kröber, 1931, p. 81; Miller, 1950, p. 71. Glaridapha viridis (Hudson). Enderlein, 1935, p. 235. Material Examined: 19 ♂ ♂, 4 ♀ ♀, including the type series, comprising two males (one marked with a red label) and a female, all three collected at Bush Hill on the same day. A dull olive-brown species. Female distinguished by diverging frons, transverse callus, narrow dark median and dorsocentral lines on the scutum, abdomen without dark vitta, and the pale median triangles ill-defined or absent. The male has a diffuse, dark olive, median vitta and long greyish cream hairs on the abdomen. Length: ♂, 9–13 mm; ♀, 12–14 mm. ♀. Head: Eyes (relaxed) dark green; with inconspicuous light brown hairs. Frons diverging, index 3 to 3.5, dull yellowish brown, with short, inconspicuous brown
hairs; vertexal triangle indefinite; one or more brown spots in position of ocelli; callus light brown, transverse, nearly full width of frons, and with short median extension. Subcallus bulging, yellowish brown, without hairs. Parafacials and face cream, the parafacials more yellowish adjacent to subcallus, both with inconspicuous cream hairs. Antennae: First and second segments brown, the first covered with cream tomentum, both with short black hairs; third brownish black, slender, but with strong dorsal angle. Palpi yellowish, with creamy overlay above, and with yellowish cream and a few brown hairs. Beard cream. Thorax: Scutum and scutellum dull olive-brown with a greyish overlay, with narrow dark brown median line, slightly wider and less definite brown dorsocentral lines, and fawn-brown lateral margins. Hairs sparse erect brown and appressed and semi-appressed cream; marginal hairs dark brown in front of wing root, greyish cream above and behind it, and on scutellum. Pleurae grey and pale fawn, with cream hairs. Legs: Light yellowish brown, slightly darkened basally on femora and apically on tarsi; hairs predominantly cream on femora, light yellow on tibiae, and becoming brown on tarsi; hind tibial fringes indefinite, mixed yellowish cream and some dark brown. Wings: Faintly greyish, costal cell slightly darkened; stigma brown; veins brown. Abdomen: Dull brown, with apices of the tergites fawn. Hairs dark brown on most of tergites; yellowish cream in median part of their apical edges, with a tendency to form indefinite median triangles; mixed dark and yellowish cream sublaterally on apical edges; and brighter cream on apical lateral corners and lateral margins. Venter dull brown, apical margins of sternites pale; hairs black in median zone, light yellowish cream laterally and on apical edges of sternites, except for the seventh, on which the hairs are almost entirely dark brown. ♂. The male differs appreciably from the female, and specimens are much more numerous, so it is advisable to describe it separately. Head: Eyes (relaxed) bright green; with long, light brown hairs, paler below; upper facets not enlarged nor differentiated from lower. Ocellar tubercle reaching level of eyes, sometimes with indications of ocelli. Frontal triangle and subcallus dull to yellowish cream, the latter with a few fine cream hairs lateral to antennae. Parafacials bulging, yellowish cream, with some brown hairs above, entirely cream below; face hollow, greyish cream, with cream hairs. Antennae similar to female, but third segment more slender. Palpi creamy yellow, with cream and a few brown hairs. Beard cream. Thorax: Similar to female, but with longer, denser, greyish cream hairs, and less evident longitudinal lines. Legs: Femora brown, with a grey overlay; tibiae with predominantly dark brown to black hairs; hind tibial fringes yellowish cream, more or less extensively mixed with dark brown laterally; otherwise as in female. Wings: As in female. Abdomen: Light yellowish brown, with a broad, diffuse, dark olive, median vitta, and the apical tergites more or less extensively darkened; apical margins of tergites narrowly cream. Hairs long and conspicuous, greyish cream, mixed with a few black ones in sublateral areas. Venter similar to female, but the hairs long, conspicuous, and almost entirely pale. Distribution. North Island: Kaitaia, February; Titahi, February; Silverstream, Hutt Valley, March; Bush Hill, Karori, February; Wellington, January, February, March. Three males (not included in the series listed above) differ appreciably from the typical form. One (Wellington, N.I., January) is paler, and completely lacks the dark abdominal stripe; it may have been immature. Another (without locality) also lacks the abdominal stripe, but it is darker, and has some dark sublateral hairs on the
abdominal tergites. The third (Okato, N.I.) has the dark abdominal stripe more or less divided by incipient median greyish olive triangles covered by relatively conspicuous greyish cream hairs, the sublateral areas on the tergites with predominantly black hairs, and the femora largely dark grey. One female (Wilton's Bush, Wellington, N.I., February) is difficult to place. The frons, thorax, legs and abdomen are like viridis; but the ocellar spots are well developed, the callus is elongate, and the antennae and palpi are more like those of the variant of opla noted above. It could, with almost equal propriety, be placed in either. Dasybasis (Protodasyommia) bratrankii (Nowicki). Text-fig, 6, J, Q. Tabanus bratrankii Nowicki, 1875. Type ♂, from “Queensland” (but really from New Zealand), in the Zoological Museum, Cracow University, Poland (stated by Miller, 1950, to have been destroyed during the war). Therioplectes bratrankii (Nowicki), Kröber, 1931, p. 87; Miller, 1950, p. 71. Glaridapha bratnankii (Nowicki). Enderlein, 1935, p. 235 (lapsus). Therioplectes pseudobratrankii Kröber, 1931, p. 88. Type ♂, from Ohakune, in the British Museum (Natural History). Synonymy by Oldroyd (personal communication), who says: “I do not see what the difference is between pseudobratrankii Kröber and bratrankii Now. The former may be a little smaller, and with a slightly narrower frons, but a specimen determined by Kröber himself as bratrankii agrees perfectly with his type of the former”. Material Examined: 1 ♂, 4 ♀ ♀, two of which were returned by Mr. Oldroyd without comment on the provisional identification. These specimens agree with Kröber's re-description, but there is a complex of closely related forms in this group, and the precise identity of Nowicki's species will probably never be known. A medium sized, rather dull brown species, with diverging frons, narrow light brown callus, indications of pale scutal vittae, well developed pale triangles on abdomen, and light yellowish brown legs. Length, 11–14 mm. ♀ Head: Eyes (relaxed) greenish to brownish black; with brown hairs, paler below. Frons diverging, index 3 to 4, yellowish brown, with dark brown hairs; vertexal triangle greyish, not strongly defined; one to three brown ocellar spots usually present; callus light brown, less than one-third width of frons, elongate-triangular, with a short extension. Subcallus yellowish fawn, with some dark brown hairs laterally. Parafacials light yellowish fawn, with brown hairs; face paler and more greyish, with brown hairs. Antennae: First and second segments bright brownish yellow, with black hairs; third brownish black, paler at base, slender, and with small but sharp tooth. Palpi yellowish fawn, with yellowish cream and scattered dark brown hairs. Beard cream. Thorax: Median and sublateral areas of scutum dark brown, with an olive tint, more greyish near anterior margin; dorsocentral vittae and lateral areas lighter brown with a yellowish tint, the dorsocentral vittae widening considerably medially in front of scutellum, and the lateral areas with a variable brown patch above wing root. Hairs erect dark brown and appressed greyish cream; marginal hairs dark brown in front of wing root, creamy white above and behind it. Scutellum light reddish brown, with some dark hairs, and quite dense yellowish cream to creamy white ones, especially on margins. Pleurae pale fawn with a greyish suffusion, and with cream hairs. Legs: Light yellowish brown, darkening to deep brown on apical tarsi; hairs predominantly cream to pale yellow on femora and most of tibiae, darkening to brown apically on tibiae and on tarsi; hind tibial fringes dull golden mixed with a few black hairs. Wings: Lightly suffused with greyish brown, costal cell slightly darkened; stigma brown; veins yellowish brown anteriorly, darkening somewhat distally and posteriorly. Abdomen: Brown, with a tendency to a yellowish hue laterally on the first and second tergites, and with large, well defined, creamy grey median triangles on the second to the fifth or sixth tergites. Hairs dark brown on the darker parts, cream on
most of the triangles, and more yellowish sublaterally on the apical edges of the tergites; apical lateral corners paler, and with bright cream hairs. Venter light brown, vaguely and irregularly darkened on some of the sternites, and with their apical margins pale; hairs creamy white laterally and on apical edges, black in the median zone, and entirely on the seventh sternite. ♂. More yellowish brown than female, and with pale hairs more golden in colour. Eyes with upper facets not appreciably enlarged; ocellar tubercle elongate, reaching level of eyes; parafacials darker than in female; palpi creamy yellow, with pale golden and dark brown hairs; scutum not evidently vittate, but somewhat paler on lateral margins, median triangles on abdominal tergites smaller than in female, and with bright yellow hairs. Distribution. North Island: Great Barrier Island; Waitakerei Hills, January; Taumaranui, February (Kröber, 1931, as pseudobratrankii); Ohakune, February, March; Korokoro, February (Kröber, 1931); Silverstream, Hutt Valley, February (Kröber, 1931). The chief variations in the females are in the development of the ocellar spots, and in the shape of the palpi, which are unusually long and slender in the specimen illustrated (Text-fig. 6, Q). Dasybasis (Protodasyommia) spp. Some specimens which belong to the viridis-bratrankii complex cannot be placed precisely on the material available. A. Two females (Ohakune, N.I., March; Wilton's Bush, Wellington, N.I., January) resemble bratrankii; but are smaller (length 11–12 mm), the first antennal segment is less expanded, the third has only a blunt tooth, and the palpi are shorter and wider (Text-fig. 6, R). They are particularly interesting in providing a link with the truncata group. B. Two females (Auckland, N.I.; York Bay, Wellington, N.I., December) are rather uniformly dull yellowish brown, without scutal vittae or abdominal triangles, and with light golden hairs on thorax and abdomen; the antennae are normal, and they may be variants of D. viridis. C. A. male (Great Barrier Island, N.I.) may be a variant of viridis; but the subcallus and parafacials are a darker yellowish fawn, hairs on parafacials mostly dark brown; first and second antennal segments darker; palpi with predominantly dark brown hairs; hairs on thorax more creamy yellow; legs darker, the femora dark grey with creamy yellow hairs, hind tibial fringes entirely black; hairs on the sublateral parts of the abdominal tergites and in the median ventral zone black. D. A male (Waitakerei Hills, N.I., January) is similar to the preceding; but the first antennal segment is covered with creamy grey tomentum; parafacial hairs are all yellowish cream; the pleural hairs are light golden anteriorly, greyish white on the metapleural convexity; and the abdomen is dull olive, with somewhat paler median triangles, and conspicuous black sublateral hairs on the second and subsequent tergites. Except for the black ones in the median zone, the hairs on the venter are ashy white, like those on the posterior part of the pleurae, contrasting with the golden hairs elsewhere. The following species appears to belong to the same complex, but cannot be identified in the material before me. Dasybasis (Protodasyommia) difficilis (Kröber) Therioplectes difficilis Kröber, 1931, p. 86; Miller, 1950, p. 71. Type ♂, from New Zealand, in the Zoological Museum, Berlin. Glaridapha difficilis (Kröber). Enderlein, 1935, p. 235. Kröber's describtion, translated, is: “♂. Length, 12 mm. “Is very similar to Th. bratrankii, but the tergites have only light hairy spots and no white centre triangles; also somewhat similar to Th. viridis, but the abdomen has
a broad, black, median stria (over one-third width of segment). Probably this is not the case with the male of viridis, since the female merely shows a stripe of a sort produced by the black hairy covering. Femora black as far as the knee. Thorax without striae, with three very poorly recognizable darkish lines, similar to the female of viridis. “Eyes perfectly contiguous, covered with short light hairs. Ocellar tubercle deeply embedded, grey. Frontal triangle and face ochre-yellow, slightly hairy. First and second antennal segments and the base of the third brownish. Hairy covering black and relatively short. First segment grey tomentose; third and style black. Third with a small distinct tooth near the base. Palpi yellow-brown, terminal segment elongate-oval, approximately two and a half times longer than wide, covered with relatively short yellow hairs. Occiput yellow, with hairs of the same colour, at the margin of the eyes with long, delicate, black hairs inclining forward. “Thorax and scutellum greenish grey on account of the tomentum, covered with clinging yellowish hairs and erect, very delicate black hairs, with indication of three fine brown lines. A black hairy stria is in front of the wing base. “Anterior coxae and femora black-grey, appearing yellowish through tomentum and hairy covering. Knees and tibiae light yellowish, with hairs of the same colour. Apex of fore tibiae brownish, hind tibiae wholly brown on the outside, covered with black hairs. Tarsi black. “Squames and halteres yellow, stem dark brown. Wings slightly brownish, stigma long, black-brown. “Abdomen black. First to fourth tergites yellow-brown at the sides, tending in parts toward red. The extreme sides of fifth and sixth are similar, also the posterior margins of first to fifth (the sixth shows only traces). Centre triangles with yellow hairs on a black ground. The whole hairy covering is black, excepting the margins and the posterior corners of second to sixth tergites, where it is golden yellow. Seventh segment black on both sides, covered with black hairs. Venter red-yellow, with blackish central stripe; the whole densely covered with yellow hairs.” The type was apparently unique, and no other locality was given. References Dobrotworsky, N. V., 1955. The Culex pipiens group in south-eastern Australia. IV. Cross-breeding experiments within the Culex pipiens group. Proc. Linn. Soc. N.S.W., 80: 33–43. Enderlein, G., 1935. Dipterologica III. S. B. Ges. naturf. Fr., 1935: 235–246. Fairchild, G. B., 1950. The generic names for Tabanidae (Diptera) proposed by Adolfo Lutz. Psyche, 57: 117–127. Hudson, G. V., 1892. “An Elementary Manual of New Zealand Entomology.” London. ——, F. W., 1901. Synopsis of the Diptera Brachycera of New Zealand. Trans. N.Z. Inst., 33: 1–95. Kröber, O., 1931. Die Tabaniden Neuseelands. Stettin. ent. Zeit., 92: 58–89. Mackerras, I. M., 1954. The classification and distribution of Tabanidae (Diptera). I. Aust. J. Zool., 2: 431–454. Mackerras, I. M., 1955. The classification and distribution of Tabanidae (Diptera). II. Aust. J. Zool., 3: 439–511. Miller, D., 1950. Catalogue of the Diptera of the New Zealand sub-region. Dept. Sci. Ind. Res., New Zealand, Bull. 100: 194 pp. Tillyard, R. J., 1926. “The Insects of Australia and New Zealand.” Sydney: Angus and Robertson. Appendix Difficulty was experienced in finding some of the localities on available maps, and Dr. Miller was put to considerable trouble to place them for me. All the localities recorded in this paper are therefore listed below, in sequence from north to south in each island, with their approximate south latitude, so that other workers may be able to find their position without difficulty.
North Island Whangaroa 35° 03′ Ohakune 39° 25′ Kaitaia 35° 07′ Rangataua 39° 25′ Whangarei 35° 43′ Aramoho, Wanganui 39° 55′ Taranga Island 35° 57′ Tiritea R. 40° 26′ Great Barrier Island 36° 10′ Lake Horowhenua 40° 37′ Parakai (tidal flats) 36° 35′ Levin 40° 48′ Brown's Bay, near Auckland 36° 40′ Mt. Holdsworth 40° 53′ Rangitoto Island, near Auckland 36° 47′ Mangatiriri R., Tararua Range 40° 55′ Auckland 36° 50′ Kaitoke 41° 05′ Henderson 36° 52′ Titahi 41° 06′ Waitakerei Hills, W. of Auckland 36° 55′ Hutt Valley 41° 10′ Titirangi, near Auckland 36° 58′ Silverstream, Hutt Valley 41° 10′ Mayor Island 37° 17′ Porirua 41° 10′ Te Aroha 37° 33′ Lower Hutt 41° 12′ Mt. Tauhara 38° 48′ Korokoro 41° 12′ Taumaranui 38° 52′ Ngaio Bush 41° 12′ Lake Taupo (East) 38° 55′ Makara Bush, Wellington 41° 14′ Manganamu, Taup 38° 59′ Bush Hill, Karori, Wellington 41° 15′ New Plymouth 39° 03′ Wainui-o-mata, Wellington 41° 16′ Crow's Nest 39° 09′ York Bay, Wellington 41° 16′ Okato 39° 12′ Day's Bay, Wellington 41° 16′ Taranaki 39–40° Wellington 41° 17′ Whakapapa, Ruapehu 39° 13′ Wilton's Bush, Wellington 41° 17′ Pokaka 39° 18′ Muritai Track 41° 18′ Tangoio Falls 39° 19′ Gollan's Valley 41° 20′ South Island Gouland Downs, near Collingwood 40° 53′ Lake Moana 42° 35′ Trio Island 40° 54′ Arthur's Pass 42° 54′ Moutere Inlet, Nelson 41° 08′ Mt. Grey, Canterbury 43° 08′ Balloon Hut, Mt. Arthur Tableland 41° 12′ White Rock, Canterbury 43° 11′ Mt. Arthur 41° 13′ Waiho Gorge, S. Westland 43° 22′ Nelson 41° 17′ Upper Hororata 43° 32′ Third House, near Dun Mt. 41° 18′ Christchurch 43° 33′ Dun Mt., Nelson 41° 18′ Peel Forest 43° 52′ Upper Maitai, Nelson 41° 18′ Ashburton 43° 55′ Tapawera, near Nelson 41° 23′ Mt. Earnslaw 44° 37′ Lake Rotoroa 41° 55′ Alexandra 45° 15′ Murchison 41° 55′ Dunedin 45° 53′ Gowan 42° Wallacetown 46° 20′ Grey River 42° 30′ Woodend 46° 28′ Dr. I. M. Mackerras, Queensland Institute of Medical Research, Herston Road, Herston N9, Brisbane.
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Transactions and Proceedings of the Royal Society of New Zealand, Volume 84, 1956-57, Page 581
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16,281Tabanidae (Diptera) of New Zealand Transactions and Proceedings of the Royal Society of New Zealand, Volume 84, 1956-57, Page 581
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