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This thesis presents three dynamic stochastic general equilibrium models to answer three macroeconomic questions. In each model, I impose one or more frictions or constraints and analyse how these frictions affect macroeconomic dynamics. Chapter 2 studies the coordination of fiscal and monetary policies under optimal commitment and discretion policies under a New Keynesian framework. The chapter shows that when there is indexation in price setting which depends on the lagged output gap as in Steinsson (2003), under the optimal commitment policy, both fiscal and monetary policies have active roles in inflation stabilisation, even although debt follows a unit-root process. Under the optimal discretion policy, both fiscal and monetary policies have active roles in inflation stabilisation to drive debt back to the pre-shock level, consistent with Leith and Wren-Lewis (2008). Extending the model to include capital accumulation does not alter these results. Chapter 3 presents a microfounded two-country model of global imbalances and debt deleveraging. During global imbalances a sustained rise in saving in one country can lead to a worldwide fall in the interest rates and an accumulation of debt in the other country. When an ensuing deleveraging shock occurs as a result of the global financial crisis, the interest rates are forced further down. I show that in the presence of a liquidity trap the deleveraging country may face a combination of a large fall in output, deflation and real exchange rate appreciation, as a result of debt deflation. Chapter 4 adds a highly-leveraged financial sector to the Ramsey model and shows that this augments the macroeconomic effects of aggregate productivity shocks. My model is built on the financial-accelerator approach of Bernanke, Gertler and Gilchrist (BGG), in which leveraged goods-producers borrow from a competitive financial sector. In this chapter, by contrast, financial institutions are leveraged and subject to idiosyncratic productivity shocks. They obtain funds by paying an interest rate above the risk free rate, and this risk premium is anti-cyclical, and so amplifies the shocks. My parameterisation, based on US data, is one in which the leverage of the financial sector is two and a half times that of the goods-producers in the BGG model. This causes a much more significant augmentation of aggregate productivity shocks than that found in the BGG model.

Howickia nudistylus Marshall sp. nov. (Fig. 45) Description. Colour: Head pale reddish except for silvery frontal strips. General colour dull pale brown. Tarsi pale yellow to white, fore and hind tarsi paler than mid tarsi. Face entirely pale microtrichose. Head: Interfrontal bristles in 2 pairs, lower pair minute. One large orbital bristle, similar in size to outer vertical bristle. Ocelli small but distinct. Eye 2.5X genal height. Vibrissa strong; subvibrissa weak or absent. Arista with long hairs, hairs twice as long as width of first aristomere. Thorax: Scutellum microtrichose, 3.0X as wide as long; basal marginal bristles short, apical bristles broken off unique type. Wing rudiment flat, slightly longer than scutellum, microtrichose, anterior half ringed by a loop vein. Halter fully developed, pale. Male abdomen: Sternite 5 subequal to sternite 4, slightly paler medially but otherwise unmodified; sternite 3 as long as sternites 4 and 5 combined. Subanal plate damaged on unique type. Surstylus narrowly triangular, tapering posteriorly and with irregular ventral surface and only small setae. Distiphallus narrow, tubular. Postgonites strongly bent posteriorly, narrow in distal portion as in H. mercurialis. Female unknown. Type material. Holotype (male, NZAC, debu00127402): North Island, Te Paki, forest litter, 6.Dec. 1984, N.O. Mitchell. The internal genitalia plus the subepandrial sclerite and surstyli have been detached from the epandrium on this specimen so some characters (cerci, subanal plate) are obscured. Etymology. The specific name refers to the characteristically shaped surstylus, which lacks the stout ventral bristle found in closely related species. Comments. The strongly bent postgonite is similar to that of H. mercurialis and H. bicolor. These three species are certainly closely related and also share similar wing rudiments and a fused, capsule-like tergites 1���3. Howickia nudistylus is described on the basis of a single somewhat damaged specimen, but several characters, especially the distinctive surstylus, distinguish it from the other two species in the clade., Published as part of Marshall, S. A., Luk, S. P. L. & Dong, H., 2014, A revision of the New Zealand species of Howickia Richards, pp. 1-36 in Zootaxa 3887 (1) on page 19, DOI: 10.11646/zootaxa.3887.1.1, http://zenodo.org/record/287460

Howickia trilineata (Hutton) (Figs. 9, 10) Apterina trilineata Hutton, 1901: 93. Howickia trilineata, Richards, 1951: 839, 848; Harrison, 1959: 253 ���254; Richards, 1973: 389, Marshall 1989: 603; Roh��ček et al., 2001, 149. Description. Colour: Thorax and abdomen subshining black, sparsely microtrichose. Head orange with three narrow vertical silver strips on the frons: one running from ocelli to anterior margin and one at each edge of interfrontal area. Face entirely bare and shining, gena bare and shining posteriorly, vibrissal angle and anterior gena pruinose. Legs mostly brown with orange knees, fore tarsi white, other tarsi reddish brown. Head: Interfrontal bristles in 3 small equal pairs. Orbital bristles minute or absent; orbital setulae present. Ocelli very small (absent in males according to Richards (1973)). Eye 2.0X genal height. Anterior genal bristle not enlarged, similar in size to genal setulae. Face tuberculate between antennae, concave at middle. Scape with a single conspicuously long bristle (longer than pedicel). Pedicel with several bristles along distal margin and with a large inclinate dorsal bristle. Arista with relatively long hairs, hairs at least twice as long as width of first aristomere. Thorax: Scutum with single postpronotal, notopleural, supraalar, postalar and prescutellar dorsocentral bristles; postalar, dorsocentral and apical scutellar bristles subequal, basal scutellars shorter. Acrostichal setulae uniformly small. Scutellum microtrichose, 3.0X as wide as long. Anterior dorsal katepisternal bristle minute or absent, posterior dorsal bristle large. Fore femur swollen, twice as thick as mid femur. Mid tibia with strong, subequal paired proximal and distal dorsal bristles and with a smaller anterior bristle above large distal dorsal bristle; ventrally with only a long apical bristle. Hind tarsus with first two tarsomeres strongly swollen, together longer than distal three tarsomeres. Wing entirely absent; halter well developed, stem white, knob rounded, dark and uniformly microtrichose. Abdomen capsule-like, syntergite 1 + 2 entirely separate from and slightly longer than tergite 3; tergites 1���5 dark, microtrichose and sparsely setulose. Sternites 1���5 similar in colour and dimension to tergites. Female terminalia narrow, pale and telescoped into segment 6. The ovipositor was retracted on all specimens examined, and no specimens were dissected. Type material and other material examined. Paratypes (2 females, one male, BMNH): New Zealand. Auckland, Howick (no date), Brown. The holotype female (Canterbury Museum, Christchurch, New Zealand) was not examined. Additional material examined. New Zealand. Auckland, Lynfield, 15.May. 1978, W. Kuschel (7 females, NZAC); Waipuna Reserve, near Waitomo, 28.Oct. 1977, litter, B.M. May (1 female, NZAC), Lynfield, Tropicana Drive, 16.Aug. 1975, in mixed mat of Oplismenus (a perennial grass), W. Kuschel (1 female, in alcohol, NZAC). No specimens were suitable for dissection. Comments. Three paratypes (from Howick) were examined (by the first author) in the BMNH in 1989; one of these was a specimen found in the bottom of the unit tray and re-attached to an empty point bearing the type label, and one had no legs, head or bristles. The remounted specimen is the only male we have seen, and the details of its terminalia are obscured under dirt. The above description is based mostly on a series of females collected in Lynfield, not far from the type locality. Both Lynfield and Howick are suburbs in the city of Auckland. This is the only Howickia species with a halter and a bare face., Published as part of Marshall, S. A., Luk, S. P. L. & Dong, H., 2014, A revision of the New Zealand species of Howickia Richards, pp. 1-36 in Zootaxa 3887 (1) on page 7, DOI: 10.11646/zootaxa.3887.1.1, http://zenodo.org/record/287460, {"references":["Hutton, F. W. (1901) Synopsis of the Diptera Brachycera of New Zealand. Transactions of the New Zealand Institute, 33, 1 - 95.","Richards, O. W. (1951) Brachypterous Sphaeroceridae. British Museum (Natural History) Ruwenzori Expedition, 2 (8), 829 - 851. [1934 - 1935]","Harrison, R. A. (1959) Acalyptrate Diptera of New Zealand. Bulletin, New Zealand Department of Scientific and Industrial Research, 128, 1 - 382.","Richards, O. W. (1973) The Sphaeroceridae (= Borboridae or Cypselidae; Diptera Cyclorrhapha) of the Australian Region. Australian Journal of Zoology, 22, 297 - 401. [supplementary series] http: // dx. doi. org / 10.1071 / AJZS 022","Marshall, S. A. (1989) 96. Family Sphaeroceridae. In: Evenhuis, N. L. (Ed.), Catalog of Diptera of the Australasian and Oceanian Regions. Bishop Museum Press and E. J. Brill, Honolulu, pp. 601 - 607. [1155 pp]","Rohacek, J., Marshall, S. A., Norrbom, A. L., Buck, M., Quiros, D. I. & Smith, I. (2001) World catalog of Sphaeroceridae (Diptera). Slezske zemske Museum, Opava, 414 pp."]}

Howickia wahaika Marshall sp. nov. (Figs. 70���74) Description. Head: Eye 2.8���3.0X genal height. Thorax: Scutellum 2.0X as wide as long; basal and apical marginal bristles subequal in length. Male abdomen: Sternite 5 as long as sternite 4 medially, posteromedial area pale with a very pale central strip flanked by about 7 inclinate, short, stout bristles on each side; posteromedial margin with a straight comb of about 20 closely appressed flat bristles. Subanal plate complete, unusually high, fringed with short bristles on each side; ventral part of cercus projecting well below subanal plate, with one long and several shorter bristles. Surstylus broadly ovate, with a single long, narrow anteroventrally directed lobe bristle behind a posteriorly directed fingerlike lobe; fingerlike lobe with a smaller pale apical bristle. Distiphallus very narrow, tubular, tapered. Postgonite long and narrow, curved in apical third. Basiphallus frame-like, projecting posteriorly beyond distiphallus base for a distance almost equal to width of distiphallus base. Female. No females with everted ovipositors were available for study. Type material. Holotype (male, NZAC): "Upper Maitai, 3.5. 50, E.S. Gourlay" (near Nelson, South Island). Paratypes: Same as holotype (2 males, one headless, NZAC); South Island, Lewis River Valley, Duggan���s Creek, fungus trap # 3, beech forest, S. Hodge, 1998, debu00104550 (male, DEBU); Titi Island, Cook Straight, in litter at entrance of petrel burrow, J.G. Stahl, 17.Jan. 2012 (9 specimens in alcohol; NMNZ; 2 males with abdomens removed). Etymology. The specific epithet is from the Maori name for a short war club, and refers to the short and broad surstylus of this species., Published as part of Marshall, S. A., Luk, S. P. L. & Dong, H., 2014, A revision of the New Zealand species of Howickia Richards, pp. 1-36 in Zootaxa 3887 (1) on page 35, DOI: 10.11646/zootaxa.3887.1.1, http://zenodo.org/record/287460

Howickia oliveri Marshall sp. nov. (Figs. 2���4, 46��� 48) Description. Colour: Thorax and abdomen dull brown, densely microtrichose. Head brown to reddish brown. Face and gena microtrichose except for a bare anterodorsal patch on the gena. Legs mostly brown with orange knees, tarsi dirty white to yellowish. Head: Interfrontal bristles in 3 pairs, upper 2 large and slightly cruciate. Two orbital bristles, upper orbital bristle large, lower small. Ocelli large, ocellar triangle raised and distinct. Eye 2.5���2.8X genal height. Vibrissa strong, subvibrissa small, anterior genal bristle large. Arista with relatively long hairs, hairs at least twice as long as width of first aristomere. Thorax: Scutum with single postpronotal, notopleural, supraalar, postalar and prescutellar dorsocentral bristles; postalar and dorsocentral bristles subequal; basal scutellars shorter, half as long as apical scutellar bristles. Acrostichal setulae uniformly small. Scutellum microtrichose, 2.0X as wide as long. Wing and halter well developed; wing almost uniformly infuscated, with pale areas along hind margin and along a distinct longitudinal fold running up the middle of cell dm. Vein R 2 + 3 very short, running almost directly to costa, slightly sinuate; R 4 + 5 gently turned up to costa, costa bypassing apex of R 4 + 5 by several vein widths. Halter stem white, knob rounded and dark anterodorsally, white posteroventrally, uniformly microtrichose. Abdomen: Syntergite 1 + 2 separate laterally, entirely fused medially and equal in length to tergites 3 and 4. Male abdomen: Sternite 5 as long as sternite 4, posteromedial area with a pale concavity margined with 4 prominent black teeth on posterior edge. Sternite 7 (unlike all congeners except H. lepidostylus) prominently forked at ventral apex. Subanal plate broad; lower/outer part of cercus projecting ventrally as small point. Surstylus with a short, subquadrate posterior lobe and a much larger, ventrally projecting anterior lobe with an anteriorly bent, bifid apex. Distiphallus narrow, tubular. Postgonite narrow, almost straight, slightly tapered distally, posterior surface flat. Basiphallus frame-like, projecting posteriorly at right angle to distiphallus base for a distance equal to distiphallus base. Female abdomen: Tergite 8 divided into 3 parts anteriorly but connected posteriorly, dorsal (middle) part shining bare, lateral parts setulose and setose posteriorly. Epiproct very small, 0.25X as long as tergite 8, pale medially and bare except for 2 setae. Cerci small, entirely setulose and setose, similar in size to epiproct. Spermathecae spherical, ringed with weak transverse striae, paired spermathecae with duct section from junction to bulb slightly longer than bulb. Sternite 8 darker than other sclerites, triangular, tapered anteriorly and setulose except for dark, tab-like posterior margin. Hypoproct divided into a bare anterior plate and a setulose posterior plate. Type material. Holotype male (NZAC): New Zealand, Moehau, Okahutahi Stream, liver bait trap, 21���25.Apr. 1984, H. Oliver. Paratypes (140 males) same locality as holotype, but taken in water traps and Malaise traps (DEBU, NZAC). Other material examined. About 150 females, same locality as paratypes (DEBU). Etymology. This species is named for friend and colleague Hugh Oliver, who provided a rich collection of Howickia, including the entire long type series of H. oliveri. Comments. Although this species is known from a large collection of about 300 specimens taken during one week in 1984, it is unknown from any other collections. A second winged species was taken in much smaller numbers at the same time. That other species, H. lepidostylus, is easily distinguished by the abdominal characters given in the key but is externally identical to H. oliveri. Females of these two species were associated by relative abundance (the rare females were assumed to be associated with the rare male and were both treated as H. lepidostylus). No females were treated as type specimens., Published as part of Marshall, S. A., Luk, S. P. L. & Dong, H., 2014, A revision of the New Zealand species of Howickia Richards, pp. 1-36 in Zootaxa 3887 (1) on page 20, DOI: 10.11646/zootaxa.3887.1.1, http://zenodo.org/record/287460

Howickia zonula Marshall sp. nov. (Figs. 75���78) Description. Head: Eye small, 1.5���1.8X genal height. Thorax: Scutellum 2.0X as wide as long; basal and apical marginal bristles subequal in length. Male abdomen: Sternite 5 slightly shorter than sternite 4 medially, posteromedial area pale with only small setulae; posteromedial margin with two straight combs, the anterior one with about 15 bristles and the posterior one with about 30 much smaller bristles. Subanal plate with ventral 2 / 3 widely split medially, upper third complete. Cercus broadly triangular, tapered ventral part projecting well below subanal plate, with several short bristles. Elongate-oval with ventral margin strongly concave, posteromedial margin with a stout, medially directed dark spur and 2 flat, pale bristles projecting over base of spur (one dorsal, one lateral). Distiphallus simple, with a saddle-like sclerotized middle section and an expanded membranous apex. Postgonite long, straight and narrow. Basiphallus frame-like, projecting posteriorly beyond distiphallus base for a distance almost equal to width of distiphallus base. Female. No females with everted ovipositors were available for study. Type material. Holotype (male, NZAC): Big South Cape Island, 60���90 m, ex. litter, 8.Nov. 1988, J. McBurney. Paratypes: Same as holotype (7 males, 1 female, NZAC; three of the males bleached white from alcohol storage). Etymology. The specific epithet is from the Latin diminutive for a girdle or belt, referring to the double-rowed belt of posteromedial bristles on the male fifth sternite., Published as part of Marshall, S. A., Luk, S. P. L. & Dong, H., 2014, A revision of the New Zealand species of Howickia Richards, pp. 1-36 in Zootaxa 3887 (1) on page 35, DOI: 10.11646/zootaxa.3887.1.1, http://zenodo.org/record/287460

Key to the macropterous Howickia of New Zealand (Macropterous Howickia differ from other fully winged Sphaeroceridae in New Zealand by the three narrow silver strips on the frons, and by the extremely short, sinuate R 2 + 3). 1 Male sternite 5 with stout teeth along posterior margin; sternite 6 with a simple ventral lobe; sternite 7 with a strongly bifurcate posteroventral lobe (Fig. 46). Female tergite 7 broadly concave posteriorly; tergite 8 with three long anterior lobes but narrowly complete posteriorly (Fig. 48). Apex of female abdomen without conspicuous internal sclerite (Fig. 48)........................................................................................... H. oliveri Marshall sp. nov. - Male sternite 5 with a small posteromedial comb of flat bristles, posterior margin of sternite 5 without prominent teeth (Fig. 31) but ventral lobe of sternite 6 with prominent posterior processes, posteroventral lobe of sternite 7 simple. Female tergite 7 narrowly split posteriorly and tergite 8 split into three parts, small middle part entirely separate from darker and larger lateral parts (Fig. 33). Apex of female abdomen with a large, dark U-shaped internal sclerite, with arms facing posteriorly (Fig. 33)........................................................................... H. lepidostylus Marshall sp. nov., Published as part of Marshall, S. A., Luk, S. P. L. & Dong, H., 2014, A revision of the New Zealand species of Howickia Richards, pp. 1-36 in Zootaxa 3887 (1) on page 3, DOI: 10.11646/zootaxa.3887.1.1, http://zenodo.org/record/287460

Howickia palmai Marshall sp. nov. (Figs. 62���65) Description. Colour: Body and most of head black with palpus, antenna (except pedicel), trochanters, knees and tarsi yellow to orange; halter brown to black on a yellow stem. Head: Eye height 2.4X genal height. Thorax: Scutellum microtrichose, 3.0X as wide as long; apical scutellar bristles as long as basal pair. Male abdomen: Sternite 5 subequal in length to sternite 4 laterally but shallowly excised posteromedially with a pale concavity; pale area with a cluster of short, stout bristles on each half. Subanal plate broad and complete, each half subquadrate, not strongly differentiated from broadly rounded lower/outer part of cercus. Surstylus small, subquadrate, with a very large posteroventral spur on a narrow process. Postgonite uniformly broad over most of length, straight, slightly tapered apically. Basiphallus frame-like, projecting beyond distiphallus base for a distance equal to one half of width of distiphallus base. Female terminalia: Not observed, the only available females have the ovipositor totally retracted. Type material. Holotype male and 4 paratypes (2 females, 2 males, 1 with abdomen removed, paratypes in alcohol, NMNZ): New Zealand: Mt. Kaukau, Wellington, R 27 / 58954, 20.Dec. 2012, F.M. Climo, under divaricating shrubs. Etymology. The species name is a patronym honoring Ricardo Palma at the NMNZ. Comments. Howickia palmai is externally very similar to H. myersi, but the postgonites differ. Furthermore, the surstyli of H. palmai are symmetrical and lack the prominent anterior lobe of the latter species., Published as part of Marshall, S. A., Luk, S. P. L. & Dong, H., 2014, A revision of the New Zealand species of Howickia Richards, pp. 1-36 in Zootaxa 3887 (1) on page 28, DOI: 10.11646/zootaxa.3887.1.1, http://zenodo.org/record/287460

Key to the apterous or micropterous Sphaeroceridae of New Zealand (all in the genus Howickia) 1 Front of head with three distinct, complete, narrow, parallel-sided silver-white pruinose strips (Figs. 1, 3): one in the middle of the interfrontal area and one on each side of the interfrontal area. Dark part of frons forming parallel bands, not an M-shape (Figs. 1, 3, 7, 10, 12, 14). Halter often absent, face often at least partly bare and shining (Fig. 12). (H. trilineata group)... 2 - Front of head with interfrontal bristles on converging dull darker strips, middle of interfrontal area forming a triangular, indistinctly silvery area similar to the dorsally incomplete, indistinctly silvery orbital strips (Figs. 16, 17). Dark part of frons forming an M-shape (Figs. 16, 17). Halter always present, face always entirely microtrichose. (H. myersi group, species identification requires male specimens)................................................................... 11 2 Tergites 1–3 fused into a large syntergite covering most of the abdomen (Figs. 5, 8). Wing rudiment forming a flat, brown flap with a loop of venation (Figs. 5, 6, 8). Postgonite strongly bent anteriorly, with an almost right-angled bend at middle (Fig. 37).................................................................................................... 3 - Syntergite 1 + 2 separate from tergite 3 (Figs. 1, 9, 11, 13). Wing variable, often entirely absent, if present then usually without venation (Figs. 1, 9, 11, 13). Postgonite gently curved or straight, without a right-angled bend at middle (Figs, 22, 28, 43, 52, 65)................................................................................................. 5 3 Scutum and scutellum similarly dull, densely microtrichose. Orbital bristle very large, similar in size to outer vertical bristle. Face mostly microtrichose, dull. Surstylus without elongate posteroventral lobe or stout ventral bristles (Fig. 45). Halter present........................................................................ H. nudistylus Marshall sp. nov. - Scutellum shining, bare (Figs. 5, 8). Orbital bristles variable (Figs. 5, 7). Face all or mostly shining (Fig. 7). Surstylus with a stout bristle at apex of posterior lobe (Figs. 35, 39). Halter absent (Figs. 5, 6, 8)................................... 4 4 Upper orbital and outer vertical bristles well developed (Fig. 7). Portion of wing flap distal to loop of venation subequal to portion enclosed by vein (Fig. 8). Scutum entirely microtrichose, dull, contrasting with the bare scutellum (Fig. 8)......................................................................................... H. omamari Marshall sp. nov. - Upper orbital and outer vertical bristles minute or absent (Fig. 5). Portion of wing beyond loop of venation less than half as large as portion enclosed by vein (Fig. 6). Scutum and scutellum shining, without microtrichia except for narrow anterior and posterior fringes (Fig. 6)...................................................... H. mercurialis Marshall sp. nov. 5 Halter and wing rudiments absent (Figs. 1, 13). Scutum orange to reddish brown, often contrasting with darker abdomen (Figs. 1, 13). Upper orbital bristle strong, as long as outer vertical bristle (Figs. 1, 14). Basal scutellar bristle very long, as long as or longer than apical bristle (Figs. 1, 13)..................................................................... 6 - Halter present or absent, if absent then wing present as a distinct, veinless flap (Figs. 9, 11). Scutum as dark as abdomen (Figs. 9, 11). Upper orbital bristle minute or absent, at most half as long as outer vertical (Figs. 10, 12). Basal scutellar bristle shorter than apical scutellar bristle (Figs. 9, 11).................................................................... 8 6 Thorax and abdomen similarly reddish brown. Surstylus trilobed, with an elongate posterior lobe ending in a terminal spur, a triangular ventral lobe, and a spatulate anterior lobe (Figs. 49, 50). Sternite 5 of male with prominent pale posteromedial area armed with a cluster of very short, broad bristles (Fig. 51)............................... H. regalis Marshall sp. nov. - Thorax reddish brown, abdomen much darker. Surstylus bilobed. Sternite 5 of male with posteromedial bristles similar in length to other bristles on sternite....................................................................... 7 7 Scutum reddish but lower pleuron, including katepisternum, dark brown (Fig. 13). Hind tibia dorsally dark on distal quarter. Male postabdomen dark, similar to preabdomen. Male sternite 4 with enlarged posteromedial bristles near middle..................................................................................... H. nigriventer Marshall sp. nov. - Thorax orange except for dark katatergite; katatergite sharply contrasting in colour with balance of thorax (Fig. 1). Hind tibia entirely yellow-orange (Fig. 1). Male postabdomen orange, contrasting with dark abdominal segments 1–5 (Fig. 1). Male sternite 4 without enlarged bristles..................................................... H. bicolor Marshall sp. nov. 8 Halter absent (Fig. 11). Wing rudiment present as a dark, broadly oval, veinless flap several times larger than a normal halter knob (Fig. 11). Ocelli reduced, sometimes apparently absent (Fig. 12)................. H. nigrilegula Marshall sp. nov. - Halter present (Fig. 9). Wing rudiment small or absent, if present then no larger than a normal halter knob and usually stub-like (Fig. 9). At least posterior ocelli distinct (Fig. 10)........................................................... 9 9 Face entirely shining and bare (Fig. 10). Apex of halter knob truncate, whitish microtrichose (Fig. 9). Orbital bristles absent (Fig. 10)............................................................................ H. trilineata (Hutton) - Lower margin of face silvery pruinose. Apex of halter knob rounded, dark-microtrichose. Orbital bristles present or absent..................................................................................................... 10 10 Upper orbital bristle absent. Scutellum shining, sparsely setulose. Wing present as a minute, flat, veinless flap. Male sternite 5 with a small posteromedial comb row (Fig. 24)..................................... H. exasperata Marshall sp. nov. - Upper orbital bristle present. Scutellum densely and evenly microtrichose, dull. Wing a minute stub. Sternite 5 without a posteromedial comb (Fig. 27)....................................................... H. harrisoni Marshall sp. nov. 11 Sternite 5 of male without a posteromedial comb, but sometimes flanked by clusters of short setae (Figs. 55, 64)........ 12 - Male sternite 5 with a posteromedial comb (Figs. 59, 68, 72, 77)............................................... 13 12 Male sternite 5 concave posteromedially, deeper than medial length of sternite; with scattered setae but without a comb or thickened bristles (Fig. 55). Surstyli asymmetrical; right surstylus shorter than left, with a conspicuously blunter and broader terminal spur on posterior lobe (Figs. 53, 54). Both surstyli deeply U-shaped, with anterior and posterior lobes (Fig. 54). Postgonite with beak-like preapical lobe on anterior surface (Fig. 56). Basal scutellar bristle shorter than apical scutellar bristles (Fig. 18). Eye less than 2.0X genal height (Fig. 18)................................. H. myersi (Richards) comb. nov. - Male sternite 5 with posteromedial concavity shallower than medial length of sternite; posteromedial setae enlarged (Fig. 64). Surstyli symmetrical, with main body subquadrate, without prominent anterior lobe, posterior lobe smaller than large terminal spur (Figs. 62, 63). Postgonite without beak-like preapical lobe on anterior surface (Fig. 65). Basal and apical scutellar bristles equal. Eye more than 2.0X genal height............................................. H. palmai Marshall sp. nov. 13 Posteromedial comb of male sternite 5 double, with a prominent, straight row of about 15 short, broad bristles followed by a row of about 30 much smaller bristles (Fig. 77). Surstylus simple, broadly concave ventrally with posterior spur strongly inclinate (Figs. 75, 76)............................................................... H. zonula Marshall sp. nov. - Posteromedial comb of male sternite 5 single (Figs. 59, 68, 72). Surstylus differently shaped, either subquadrate or with spur directed ventrally................................................................................... 14 14 Posteromedial comb of male sternite 5 straight with teeth fused over most of their length, area immediately anterior to comb with two groups of 8 short stout bristles (Fig. 72). Scutellum 2.0X as wide as long........... H. wahaika Marshall sp. nov. - Area immediately anterior to posteromedial comb without stout bristles, if comb straight then teeth separated by at least half of their length (Figs. 59, 68). Scutellum 3.0X as wide as long................................................... 15 15 Male sternite 5 with straight posteromedial comb and a cluster of short stout bristles midway between comb and anterior margin (Fig. 68). Surstylus with anterior and posterior ventral lobes each with long spurs, posterior part of surstylus with multiple stout bristles (Figs. 66, 67)........................................................ H. tangata Marshall sp. nov. - Posteromedial comb triangular or heart-shaped, middle part of sternite 5 without additional bristles (Fig. 59). Surstylus with a single long posteroventral lobe bearing two long spurs (Figs. 57, 58)....................... H. cordata Marshall sp. nov.

Howickia regalis Marshall sp. nov. (Figs. 49���52) Description. Colour: Thorax and abdomen reddish brown except for dark posterior margins of tergites. Face and antenna pale. Tarsi pale yellow to white, fore and hind tarsi paler than mid tarsi. Head: Face entirely microtrichose. Interfrontal bristles in 3 strong equal pairs. One large orbital bristle (lower orbital absent). Ocelli small, anterior ocellus sometimes indistinct. Eye 2.5���3.0X genal height. Anterior genal bristle strong. Arista with short hairs basally and longer hairs distally, hairs on distal half twice width of first aristomere. Thorax: Scutellum uniformly microtrichose, 3.0X as wide as long; basal and apical marginal bristles subequal in length or basal slightly longer than apical. Anterior dorsal katepisternal bristle minute, posterior dorsal bristle large, reaching more than halfway to wing base. Wing and halter absent. Male abdomen: Sternite 5 with sclerotized part about half as long as sternite 4 medially, posteromedial area desclerotized and divided into 2 broad lobes each with a cluster of about 20 short, broad, flattened, bristles; those on posterior margin largest. Subanal plate incomplete, with halves broadly separated to expose a well-developed subepandrial sclerite; lower/outer part of cercus broadly rounded and inner part of cercus forming a distinct quadrate ventral lobe. Surstylus highly characteristic, trilobed with a long narrow posterior lobe terminating in a short spur, a broadly triangular ventral lobe, and a spatulate anterior lobe. Hypandrium broadly but asymmetrically fused with anteroventral corners of epandrium, fused area of right side shorter and twice as wide as fused area on left. Ventral lobe of hypandrium prominent, anterior surface of ventral part conspicuously denticulate. Posterodorsal apodeme of hypandrium strongly developed, reaching anterior surface of distiphallus near base. Distiphallus tubular and sclerotized over basal 2 / 3, distal third bulbous and membranous, anterior surface finely denticulate near middle. Postgonites broad on basal part, gradually tapering distally with anterior surface serrate on distal half. Basiphallus frame-like, projecting posteriorly at right angle to distiphallus base for a distance half as long as distiphallus base. Female abdomen: Tergite 8 divided into 3 parts, lateral parts setose and setulose along posterior margin. Epiproct and cercus greatly reduced, pale, essentially indistinguishable from surrounding membrane. Sternite 8 pale, small, setulose, with 2 small bristles along posterior margin. Type material. Holotype (male, NZAC, debu00370490) and 8 paratypes (3 males, 5 females, NZAC): Three Kings Island, 275 m, ex. litter, 23.Nov. 1970, G. Kuschel. Etymology. The species name is from the Latin for "kingly", and refers the type locality, Three Kings Island. Comments. This species is externally similar to H. nigriventer, which also has a strikingly long and narrow posterior surstylar lobe. These species otherwise differ markedly in surstylar shape and armature of male sternite 5., Published as part of Marshall, S. A., Luk, S. P. L. & Dong, H., 2014, A revision of the New Zealand species of Howickia Richards, pp. 1-36 in Zootaxa 3887 (1) on pages 22-24, DOI: 10.11646/zootaxa.3887.1.1, http://zenodo.org/record/287460

Howickia lepidostylus Marshall sp. nov. (Figs. 29���33) Description. Colour: Face and gena microtrichose except for a bare anterodorsal patch on the gena. Legs mostly brown with orange knees; tarsi dirty white to yellowish. Head: Interfrontal bristles in 3 pairs, upper 2 large and slightly cruciate. Two orbital bristles, upper orbital bristle large, lower small. Ocelli large, ocellar triangle raised and distinct. Eye 2.0���2.7X genal height. Anterior genal bristle large. Arista with relatively long hairs, hairs at least twice as long as width of first aristomere. Thorax: Scutum with single postpronotal, notopleural, supraalar, postalar and prescutellar dorsocentral bristles; postalar and prescutellar dorsocentral bristles subequal, basal scutellars shorter, half as long as apical scutellar bristles. Acrostichal setulae uniformly small; most specimens with a small anterior (postsutural) dorsocentral bristle slightly larger than surrounding acrostichals. Scutellum microtrichose, 2.0X as wide as long. Wing and halter well developed; wing almost uniformly infuscated, with pale areas along hind margin and along a distinct longitudinal fold running up the at middle of cell dm. Vein R 2 + 3 very short, running almost directly to costa, slightly sinuate; R 4 + 5 distally turned up to costa, costa bypassing apex of R 4 + 5 by several vein widths. Halter stem white, knob rounded and dark anterodorsally, white posteroventrally, uniformly microtrichose. Abdomen: Syntergite 1 + 2 separate laterally, entirely fused medially and equal in length to tergites 3 and 4. Male abdomen: Sternite 5 1.5X as long as sternite 4, posteromedial area with small median comb of 8 flat bristles preceded by a pale medial patch flanked by 2 rows of 3���4 stout, short bristles on each side. Synsternite 6 + 7 strongly modified, sternite 6 with a broad anterior apodeme projecting over sternite 5 and a bifid, toothed posteroventral lobe, sternite 7 with a bifid ventral lobe (like H. oliveri but unlike other congeners). Subanal plate broad but contiguous only in upper half, cerci widely separated below; lower/outer part of cercus projecting ventrally as small point. Surstylus with a short, subquadrate anterior lobe and a much larger, ventrally projecting posterior lobe with dark, broad, flat, scale-like bristle at apex and 3���4 flat, scale like bristles along posterior margin. Distiphallus simple, looped on basal half, expanded distally. Postgonite narrow, almost straight, slightly tapered distally, posterior surface flat. Basiphallus frame-like, projecting posteriorly at right angle to distiphallus base for a distance of about half the width of the distiphallus base. Female abdomen (see also comments below): Tergite 7 narrowly notched posteriorly; tergite 8 split into 3 parts, small middle part entirely separate from darker and larger lateral parts. Female sternite 8 small, pale and oval anteriorly, very dark and straight along posterior margin. Interior of segment 8 with a large, heavily sclerotized Ushaped internal sclerite, with arms facing posteriorly (homology uncertain, no obvious homologue present in closely related species but this sclerite is probably homologous with the smaller U-shaped anterior part of sternite 8 in wingless species). Hypoproct with an anteromedial bare portion and a transverse microtrichose portion. Large but indistinct, thin, ring-like internal vaginal sclerites (spectacles-shaped sclerites) present. Type material. Holotype (male, NZAC) and 18 male paratypes (NZAC, DEBU): New Zealand, Moehau Mt., Coromandel Range, 21���25.Apr. 1984, bush on upper Okahutahi Stream, Malaise and water tray, H. Oliver. Other paratypes: Kauaeranga Valley, near Thames, H. A. Oliver (1 male, NZAC); Moehau, Okahutahi Stream, liver bait trap, 21 ���25.1984, H. Oliver (2 males, NZAC); S.E. National Park near Tongariro River, meat-baited pitfall trap, 4���26.Jan. 1971, H.A. Oliver and J. Gillespie (3 females, 1 male, in alcohol, NZAC). Other material examined: Three females, same locality as holotype. Three females and one headless male, in alcohol: meat-baited pitfall trap near Whangaiterenga Stream, near Thames, 18.Jan. 1971 ��� 18.Apr. 1971, H.A. Oliver. Etymology. Howickia lepidostylus is named for the distinctive scale like bristles on the surstylus. Comments. Although this species is very distinctive in many characteristics of both male and female terminalia, there seem to be no non-abdominal characters by which to reliably distinguish H. lepidostylus from the much more common H. oliveri. Almost all of the males here designated as types of H. lepidostylus were collected along with much longer series of H. oliveri; males and females of these species were associated by their proportions in the collections (it was assumed that the rare females corresponded to the rare males)., Published as part of Marshall, S. A., Luk, S. P. L. & Dong, H., 2014, A revision of the New Zealand species of Howickia Richards, pp. 1-36 in Zootaxa 3887 (1) on pages 13-15, DOI: 10.11646/zootaxa.3887.1.1, http://zenodo.org/record/287460

Howickia harrisoni Marshall sp. nov. (Figs. 25���28) Description. Head: Lower half of face bare and shining except for a white pruinose ventral fringe. Upper orbital bristle large, equal to outer vertical bristle; orbital strip otherwise with only minute orbital setulae. Ocelli distinct. Anterior genal bristle enlarged. Thorax: Postalar, dorsocentral, basal scutellar and apical scutellar bristles subequal. Scutellum microtrichose, 2.0X as wide as long. Wing rudiment short and narrow, similar in dimensions to the halter stem; halter fully developed. Male abdomen: Sternite 5 slightly longer than sternite 4, slightly paler and concave medially and with posteromedial area flanked by a loose cluster of slightly enlarged bristles; sternite 4 unmodified. Subanal plate complete in upper third only; lower/outer parts of cerci broadly rounded and widely separated. Surstylus deeply bilobed, with a short stout spur at apex of posteroventral lobe, posteroventral lobe narrower and longer than anterior lobe. Distiphallus with a relatively broad but simple tubular distal part overlapped posterodorsally by a narrow lobe of basal part. Postgonite narrow, tapered, gently curved anteriorly and with small setulae on anterior surface. Basiphallus frame-like, projecting posteriorly at right angle to distiphallus base for a distance equal to distiphallus base. Female abdomen: Hypoproct U-shaped, complete along posterior margin only. Sternite 8 small, setulose on posterior half in a continuous band with the posterior setulosity of the lateral portions of tergite 8. Type material. Holotype (male, NZAC, debu00134829) and 4 paratypes (2 males, 2 females, DEBU): North Island, Coromandel Peninsula, 10 km E Thames, pan traps, 22���29.Jan. 1999, S.A. Marshall. Other paratypes: North Island, Kauaeranga Valley, Whangaiterenga River, 1.Dec. 1970 ��� 18.Jan. 1971, H.A. Oliver (13 males, 2 females, NZAC); Tiki Tiki Stream, Horohoro State Forest, Mamaku Plateau, 24.Jul. 1976, J.S. Dugdale, forest litter 76 / 60 (male, NZAC); Coromandel Peninsula, Kopu Rd., leaf litter, 20 March 1967, R.A. Comber (female, NZAC); Mangakara Stream, Pirongia Mt., meat baited pit traps, 26.Sep. 1971, H.A. Oliver (male, female, NZAC). Etymology. Howickia harrisoni is named after the great New Zealand dipterist Roy Harrison. Comments. This species is easily recognized by its stubby, narrow wing rudiment in combination with the dense white fringe of microtrichia along the lower margin of the mostly bare face. It is clearly defined on the basis of the distinctive surstylus and other male postabdominal features., Published as part of Marshall, S. A., Luk, S. P. L. & Dong, H., 2014, A revision of the New Zealand species of Howickia Richards, pp. 1-36 in Zootaxa 3887 (1) on pages 11-13, DOI: 10.11646/zootaxa.3887.1.1, http://zenodo.org/record/287460

Howickia myersi (Richards) comb. nov. (Figs. 17, 18, 53���56) Leptocera (Biroina) myersi Richards, 1973: 349 Biroina meyersi, Marshall, 1989: 602 Biroina myersi, Roh��ček et al., 2001: 125 Description. Colour: Brown to black, head reddish brown to reddish yellow; knees paler; antenna, palpus and tarsi white; halter brown to black on a yellow stem; anepimeron darker. Interfrontal area with a broad, silvery microtrichose triangle tapering from ocelli to anterior margin of frons, apex of triangle flanked by bare V-shaped area. Interfrontal bristles inserted on inner margin of silvery microtrichose interfrontal strips; orbital bristles on silvery microtrichose orbital strips; orbital strips usually much darker than interfrontal strips (indistinctly so on very pale or very dark specimens); orbital and interfrontal strips separated by a very narrow bare strip. Head: Interfrontal bristles in 3 strong equal pairs. Two orbital bristles, lower smaller, upper slightly shorter than outer vertical bristle and much shorter than inner vertical and ocellar bristles. Orbital and interfrontal setulae well developed. Ocelli well developed. Eye distinctly notched at level of antennae, eye relatively small, 1.5���1.8X genal height. Gena mostly microtrichose, with a small anterodorsal shining area. Vibrissa and anterior genal bristle strong. Face tuberculate between antennae, concave at middle. Pedicel with several long bristles along distal margin and with 2 large inclinate dorsal bristles. Arista with long hairs, hairs as long as first aristomere. Thorax: Scutum with single postpronotal, notopleural, supraalar, postalar and prescutellar dorsocentral bristles; postalar, dorsocentral and apical scutellar bristles subequal. Acrostichal setulae strong but prescutellar acrostichals not enlarged. Scutellum microtrichose, 3.0X as wide as long; apical scutellar bristles twice as long as basal pair. Anterior dorsal katepisternal bristle minute, posterior dorsal bristle small, reaching half way to wing base. Mid tibia with strong, subequal paired proximal and distal dorsal bristles and an additional small anterior bristle above large distal dorsal bristle; ventrally with only a long apical bristle. Hind tarsus with first 2 tarsomeres strongly swollen, together longer than distal 3 tarsomeres. Wing absent, halter well developed. Abdomen: Capsule-like, syntergite 1 + 2 entirely separate from and slightly longer than tergite 3; all tergites and sternites dark, microtrichose and sparsely setulose. Male abdomen: Sternite 5 subequal in length to sternite 4 laterally but deeply excised posteromedially with a pale concavity leaving the sclerotized part of sternite 5 less than half as long as sternite 4 at middle. Subanal plate broad and complete, each half subquadrate, not strongly differentiated from lower/outer part of cercus. Surstylus U-shaped, broad anteroventral lobe with only small bristles, narrow posteroventral lobe terminating in a very large apical spur; spur of left surstylus conspicuously longer and narrower than spur of right surstylus. Hypandrium broadly but asymmetrically fused with anteroventral corners of epandrium, fused area of right side shorter and twice as wide as fused area on left. Ventral lobe of hypandrium prominent. Posterodorsal apodeme of hypandrium strongly developed, reaching anterior surface of distiphallus near base. Distiphallus narrow, tubular. Postgonite uniformly broad, slightly bent anteriorly beyond middle, apex rounded with a beak-like anterior process. Basiphallus frame-like, projecting beyond distiphallus base for a distance equal to one third of width of distiphallus base. Female terminalia: Tergite 8 divided into 3 sclerites, dorsal sclerite greatly reduced, bare and pale medially; lateral sclerites large, pale and shining except along posterior quarter. Epiproct small, medially pale, with 2 small setae. Hypoproct divided into anterior and posterior parts: posterior part bare and shining laterally, pale medially, setulose along posterior margin; anterior part entirely bare and shining. Sternite 8 small, but distinct and darkly pigmented, posterior portion transverse, broad, microtrichose and setose; anterior portion narrow and bare. Spermathecae broadly oval, with transverse grooves. Type material. Holotype: The unique holotype (BMNH; examined and illustrated by the first author in 1989) is a female from Day's Bridge ("in leaf mould, 12.Jun. 1922, J.G. Myers"). Although there is no further locality information on the label, and no current locality is called "Day's Bridge", there is a "Days Bay" in Wellington, North Island. According to Dr. Trevor Crosby (pers. comm.), John Golding Myers was based in Wellington in 1922 and it is therefore "without question that the locality is Days Bay -- at the bridge [New Zealand, WN]". Other material examined (Approximately 100 specimens: DEBU, NZAC): North Island: Mt. Ruepehu, Ohakune, 700 m, Rimu Track, 18.Jan. 1999, S.A. Marshall; Egmont National Park, Stratford Plateau, 1200 m, pan trap, 8���12.Jan. 1999, S.A. Marshall; Mt. Taranaki, Wilkies Pools, 3000 m, moss, 16.Jul. 1965, J.I. Townsend; Mt. Pirongia, 800 m, TK 2 to Wharauroa, 9.Jul. 1977, B.M. May. South Island and nearby small islands: Stephens Island, 14���28.Jan. 1933, E.S. Gourlay; Stephen's Island, ex. litter, Feb. 1971, G.W. Ramsay; Westland National Park, Franz Josef, 100���250 m, 14���19.Feb. 1999, S.A. Marshall; Takaka, Pupu Springs, 60 m, 6.Feb. 1999, dung traps, S.A. Marshall; Reefton, 12.Apr. 1977, J.A. Wightman; Abel Tasman National Park, 30 km NW Motueka, 19.May. 1982, S. Peck; Nelson, Canaan, ex. litter 17.Sep. 1964, L.P. Marchant. Comments. The type specimen was examined and sketched during a visit to BMNH in 1989, providing some details of head pigmentation and chaetotaxy absent from the original description but mostly supporting the interpretation of the species followed here. Richards��� description differs from all known New Zealand Howickia in one important detail. He clearly states that the mid tibia has "no proximal bristles but a pair of preapical dorsals", while all Howickia species known to us have a strong pair of proximal dorsal bristles. The type has a proximal anterodorsal bristle and an apparent socket where a posterodorsal bristle has probably been broken off. The most significant difference between the type and the specimens described above is the anterior orbital bristle, which is equal in size to the posterior orbital in the type but smaller in the specimens here treated as H. myersi. This species has the widest distribution and the widest range of colour forms of any New Zealand Howickia, and is the only species known from both the North Island and South Island. The genitalia are distinctive (asymmetrical surstyli, uniquely shaped postgonites) and consistent across the entire range, as are the diagnostic details of head pruinosity., Published as part of Marshall, S. A., Luk, S. P. L. & Dong, H., 2014, A revision of the New Zealand species of Howickia Richards, pp. 1-36 in Zootaxa 3887 (1) on pages 24-26, DOI: 10.11646/zootaxa.3887.1.1, http://zenodo.org/record/287460, {"references":["Richards, O. W. (1973) The Sphaeroceridae (= Borboridae or Cypselidae; Diptera Cyclorrhapha) of the Australian Region. Australian Journal of Zoology, 22, 297 - 401. [supplementary series] http: // dx. doi. org / 10.1071 / AJZS 022","Marshall, S. A. (1989) 96. Family Sphaeroceridae. In: Evenhuis, N. L. (Ed.), Catalog of Diptera of the Australasian and Oceanian Regions. Bishop Museum Press and E. J. Brill, Honolulu, pp. 601 - 607. [1155 pp]","Rohacek, J., Marshall, S. A., Norrbom, A. L., Buck, M., Quiros, D. I. & Smith, I. (2001) World catalog of Sphaeroceridae (Diptera). Slezske zemske Museum, Opava, 414 pp."]}

Howickia bicolor Marshall sp. nov. (Figs. 1, 19���22) Description. Colour: Head, most of the thorax and terminalia yellow; abdominal tergites 1���5, katatergite dark Head: Orbital bristles in a single large pair, subequal to outer vertical, smaller than ocellar and inner vertical bristles. Ocelli small but distinct. Eye 3.0X genal height. Anterior genal bristle strong. Arista with very short hairs, hairs no longer than width of first aristomere. Thorax: Scutellum with basal marginal bristles slightly longer than apical pair. Wing and halter entirely absent. Male abdomen: Sternite 5 narrow, dark except for a weakly spinulose posteromedial patch. Synsternite 6 + 7, sternite 8 and epandrial complex pale orange in contrast with preabdomen. Cercus divided into two parts, upper/ inner part flat and tapered medially to form an incomplete subanal plate, lower/outer part larger, bulging, rounded, and setose. Surstylus broadly U-shaped, posterior lobe narrower and longer than anterior lobe, without spurs or bristles. Hypandrium broadly but asymmetrically fused with anteroventral corners of epandrium, fused area of right side twice as wide as fused area on left. Ventral lobe of hypandrium very prominent, with median anterior part finely serrate on anterior surface. Posterodorsal apodeme of hypandrium strongly developed, reaching anterior surface of distiphallus near base. Distiphallus narrow, tubular. Postgonites narrow, tapered to tip. Basiphallus frame-like, projecting posteriorly at right angle to distiphallus base. Female abdomen: Ovipositor (segment 6 and posterior) very small and lightly sclerotized, normally retracted into preabdomen, apex compact and difficult to interpret in dissected specimens. Tergite 8 divided into 3 parts, dorsal part bare and shining, lateral parts setose and setulose along posterior margin. Epiproct with two small bristles, posteromedially pale, dark posterolaterally. Cercus very short, as wide as long, setose. Sternite 8 simple, microtrichose, with a few small setae along posterior margin. Hypoproct U-shaped, with arms extending into abdomen as part of a bare plate, microtrichose only at posterior extreme. Spermathecae spherical, ringed with weak transverse striae, paired spermathecae with duct section from junction to bulb as long as bulb. Type material. Holotype (pointed male, NZAC) and 7 pointed paratypes (5 males, 2 females, NZAC): New Zealand, North Island, Coppermine Island, Hen and Chicken Islands, 24���31.Oct. 1968, litter, J.C. Watt leg. Other paratypes: New Zealand, North Island, Lady Alice Island, Hen and Chicken Islands, 18.Oct. 1971, litter, R.H. May (1 female in alcohol, NZAC); Chickens Islands, Whatupuke Island, 28.Oct. 1968 (2 males, NZAC). Comments. The distinctive and contrasting colours, with the head, most of the thorax and male postabdomen yellow in contrast with the dark preabdomen and katatergite, render this species unmistakable. Howickia bicolor is known only from the Hen and Chicken Islands, east of the North Auckland Peninsula, 40 kilometers southeast of Whangarei. Only one female was dissected, and the apical abdominal sclerites are minute, pale and compact, with individual sclerites difficult to discern., Published as part of Marshall, S. A., Luk, S. P. L. & Dong, H., 2014, A revision of the New Zealand species of Howickia Richards, pp. 1-36 in Zootaxa 3887 (1) on pages 9-10, DOI: 10.11646/zootaxa.3887.1.1, http://zenodo.org/record/287460

Howickia exasperata Marshall sp. nov. (Figs. 23, 24) Description. Colour: Head reddish brown; thorax shining black; preabdomen dark brown-black. Tarsi pale yellow to white, fore and hind tarsi paler than mid tarsi. Face white pruinose along upper and lower margins, shining at middle. Upper white pruinose area of face extending to frons, broken by antennae but reappearing in eye notch. Head: Interfrontal bristles in 2 equal pairs. Orbital bristles absent. Ocelli small but distinct. Eye 2.5X genal height. Anterior genal bristle strong. Aristae broken off from unique type. Thorax: Scutellum shining, sparsely microtrichose (possibly rubbed off on unique type), 2.0X as wide as long; basal and marginal bristles conspicuously finer and shorter than apical bristles. Wing present as a minute, oval, shining brown, veinless flap; halter well developed. Male abdomen (only available male cleared but not dissected): Sternite 5 subequal to sternite 4, medial area slightly paler and flanked by 6���7 stout bristles and with a weak posteromedial comb row of 12 small, flat bristles; sternite 4 simple. Subanal plate incomplete; cercus indistinct. Surstylus distinctive, with bilobed anterior part and a narrow, elongate posteroventral lobe ending in a short, stout bristle. Distiphallus heavily sclerotized and tubular near base but expanded and largely membranous distally. Postgonites broadest apically, with a weak beak-like anteroventral point. Female. Unknown. Type material. Holotype (male, NZAC): New Zealand, Moehau, Okahutahi Stream, liver bait trap, 21���25.Apr. 1984, H. Oliver. Etymology. The species name reflects the way most taxonomists feel when they find yet one more singleton representing yet another unnamed species. Comments. This is the only species in the H. trilineata group with a posteromedial comb. The surstylus of this species is distinctive, and leaves no doubt about its validity although it is known from only a single specimen. The terminalia suggest that this species is more closely related to the H. myersi group than the H. trilineata group, but it is grouped with the latter because of the distinctively striped head., Published as part of Marshall, S. A., Luk, S. P. L. & Dong, H., 2014, A revision of the New Zealand species of Howickia Richards, pp. 1-36 in Zootaxa 3887 (1) on pages 10-11, DOI: 10.11646/zootaxa.3887.1.1, http://zenodo.org/record/287460

Marshall, S. A., Luk, S. P. L., Dong, H. (2014): A revision of the New Zealand species of Howickia Richards. Zootaxa 3887 (1): 1-36, DOI: http://dx.doi.org/10.11646/zootaxa.3887.1.1

Howickia tangata Marshall sp. nov. (Figs. 15, 16, 66���69) Description. Head: Eye 3.0X genal height. Thorax: Scutellum 3.0X as wide as long; basal and apical marginal bristles subequal in length. Male abdomen: Sternite 5 twice as long as sternite 4, with a pale, strongly spinulose medial patch and a straight posteromedial comb of 13 basally contiguous but distally widely spaced flat bristles. Subanal plate broad and complete with a straight ventral margin, not strongly differentiated from lower/outer part of cercus; lower parts of cerci widely separated and projecting below subanal plate. Surstylus with 3 lobes: a broad anterior lobe without large bristles; a small, pedunculate ventral lobe with a stout posteromedially curved bristle; and a large posterior lobe with several stout bristles including a very large ventral one. Distiphallus very narrow, tubular. Postgonites broad on basal part, abruptly constricted to a narrow and weakly clubbed distal part. Basiphallus frame-like, projecting posteriorly at right angle to distiphallus base for a distance equal to distiphallus base. Female abdomen (described from undissected abdomen with everted ovipositor): Tergite 8 divided into narrow bare dorsal sclerite and larger posteriorly setulose and setose lateral sclerites. Epiproct longitudinally divided into two sclerites, each with 1 small bristle and several small setulae; epiproct separate from the strongly sclerotized, elongate, narrow, setose cerci. Sternite 8 small and bare. Hypoproct large, mostly bare, setulose and setose only along posterior margin. Type material. Holotype (male, NZAC, debu00134615): New Zealand, North Island, Putara Valley, 10 km W Eketahuna, 400 m, pan traps, 14���16.Jan. 1999, S.A. Marshall. Paratypes: same as holotype (1 male, DEBU, debu00134616); same as holotype but 15.Jan. 1999 (1 male, DEBU); North Island, Tongariro National Park, southeast corner, Nothofagus fusca bush, baited pit traps, 4���26.Jan. 1971, H.A. Oliver (4 males, 3 females, NZAC); Kaimanewa Road end, NW Kaimanawa Ranges meat-baited pit traps, 23���31.Jan. 1971, H.A. Oliver (1 male, 1 female, NZAC). Etymology. The specific epithet is from the name for the ceremonial Maori war club called the Toki Pou Tangata. It is applied to this species because of its elaborate and well-armed surstylus. Comments. The male genitalia of this species are unmistakable, but Howickia tangata resembles H. myersi in external features., Published as part of Marshall, S. A., Luk, S. P. L. & Dong, H., 2014, A revision of the New Zealand species of Howickia Richards, pp. 1-36 in Zootaxa 3887 (1) on page 28, DOI: 10.11646/zootaxa.3887.1.1, http://zenodo.org/record/287460

Howickia mercurialis Marshall sp. nov. (Figs. 5, 6, 34���38) Description. Colour: Face bare and shining at middle only, upper and lower margins white pruinose; gena mostly bare with vibrissal angle and upper margin white pruinose. Legs mostly brown with orange knees; first tarsomere of fore and mid leg reddish brown, other tarsomeres pale brown to white. Head: Upper orbital bristle minute, outer vertical similarly reduced. Ocelli small but distinct. Arista with hairs 1.5X as long as width of first aristomere. Thorax: Scutum shining, without microtrichia except for narrow anterior and posterior fringes. Postalar and dorsocentral twice as long as scutellar bristles. Scutellum 4 X as wide as long, bare and shining except for microtrichose margin. Wing present as a short, oval, transluscent flap with a single loop of venation. Portion of wing beyond loop of venation less than half as large as portion enclosed by vein. Abdomen: Tergites 1���3 fused into a large syntergite covering most of the abdomen. Male abdomen: Synsternite 2 + 3 large, longer than sternites 4���5 combined. Sternite 5 short, subequal to sternite 4, slightly paler medially and with medial area flanked by small setae; sternite 4 unmodified. Subanal plate complete, narrow; lower/outer part of cercus broadly rounded. Surstylus with stout posteroventral spur and quadrate anteroventral process, ventral surface otherwise smooth. Hypandrium broadly but asymmetrically fused with anteroventral corners of epandrium, fused area of right side shorter and twice as wide as fused area on left. Ventral lobe of hypandrium prominent, U-shaped. Posterodorsal apodeme of hypandrium strongly developed, reaching anterior surface of distiphallus near base. Distiphallus relatively broad and short, with a broad spinulose apex. Postgonites narrow, slightly broader basally and bent anteriorly at right angles near middle, distal part parallel-sided. Basiphallus frame-like, projecting posteriorly but deflexed ventrally and distally almost parallel to distiphallus base. Female abdomen: As described for H. bicolor, but cerci and epiproct greatly reduced and apparently fused into a small, bare sclerite. Type material. (NZAC, all point-mounted): Holotype male (debu00127423) and 16 paratypes (10 males, 6 females): New Zealand, Mercury Islands, Green Island, pitfall trap under Coprosma, 11���23.Dec. 1987, G. Hall. An additional 3 males and 7 females have the same data except "under Melicytus ramiflorus ". Other paratypes: Mercury Islands, Middle Island, litter 83 / 110, 20.Oct. 1983, Island Southey (30 males, 9 females); same data but collected from litter 83 / 111 (17 males, 13 females); Cuvier Island, Old Radar Station, pitfall trap, 25.Feb��� 2.Mar. 1982, G. Hall (2 males, 1 female, see notes below); Ohena Islands, ex. litter, 26.Nov. 1972, G.W. Ramsay (1 female, NZAC). Other material examined (in alcohol, not designated as paratypes; in addition to several further vials from the Mercury Islands): Ohena Islands, Keruenga Island, 27.Nov. 1972, G.W. Ramsay (4 specimens, NZAC). Etymology. Howickia mercurialis is named for the type locality. Comments. Howickia mercurialis is closely related to H. omamari, with which it shares several unequivocal synapomorphies including the capsule-like abdomen dominated by a hugely expanded syntergite 1���3. These species have distinctly different surstyli and can be easily separated by the relative pruinosity of the scutum (bare and shining in H. mercurialis, microtrichose and dull in H. omamari). All specimens are from the Mercury Islands except the three specimens from Cuvier Island, about 15 km north of Mercury Island. Those three specimens have a more reddish thorax than any of the specimens from the Mercury Islands, but they seem identical in male genitalic features., Published as part of Marshall, S. A., Luk, S. P. L. & Dong, H., 2014, A revision of the New Zealand species of Howickia Richards, pp. 1-36 in Zootaxa 3887 (1) on page 15, DOI: 10.11646/zootaxa.3887.1.1, http://zenodo.org/record/287460

© 2014 Macmillan Publishers Limited. All rights reserved. Tumour-stromal interactions are a determining factor in cancer progression. In vivo, the interaction interface is associated with spatially resolved distributions of cancer and stromal phenotypes. Here, we establish a micropatterned tumour-stromal assay (Î 1/4TSA) with laser capture microdissection to control the location of co-cultured cells and analyse bulk and interfacial tumour-stromal signalling in driving cancer progression. Î 1/4TSA reveals a spatial distribution of phenotypes in concordance with human oestrogen receptor-positive (ER+) breast cancer samples, and heterogeneous drug activity relative to the tumour-stroma interface. Specifically, an unknown mechanism of reversine is shown in targeting tumour-stromal interfacial interactions using ER+ MCF-7 breast cancer and bone marrow-derived stromal cells. Reversine suppresses MCF-7 tumour growth and bone metastasis in vivo by reducing tumour stromalization including collagen deposition and recruitment of activated stromal cells. This study advocates Î 1/4TSA as a platform for studying tumour microenvironmental interactions and cancer field effects with applications in drug discovery and development.