1 2 DAN A. POLHEMUS & JOHN T. POLHEMUS

1Dept. of Entomology, National Museum of Natural History, Washington DC, USA 2Colorado Entomological Institute, Englewood, CO, USA

TWO NEW GENERA AND THIRTY NEW SPECIES OF MICROVELIINAE (HETEROPTERA: VELIIDAE) FROM THE EAST PAPUA COMPOSITE TERRANE, FAR EASTERN NEW GUINEA

Polhemus, D. A. & J. T. Polhemus, 2004. Two new genera and thirty new species of Microv- eliinae (Heteroptera: Veliidae) from the East Papua Composite Terrane, far eastern New Guinea. – Tijdschrift voor Entomologie 147: 113-189, figs. 1–86. [ISSN 0040-7496]. Pub- lished 1 December 2004. Two new genera and 30 new species of Microveliinae are described from the East Papua Com- posite Terrane of far eastern New Guinea. This geologically defined region, including the Papuan Peninsula and the D’Entrecasteaux, Louisiade, and Marshall Bennett island groups, is hypothesized by geologists to have formed as a discrete microcontinent in the early Tertiary be- fore being sutured to greater New Guinea in the Miocene, and retains a highly endemic aquatic Heteroptera biota. The new taxa described from this area are as follows: Rheovelia gen. n. con- taining type species R. petrophila sp. n. from extreme southeastern New Guinea, R. robinae sp. n. from the D’Entrecasteaux Islands, R. insularis sp. n. from Tagula and Rossel islands, R. anomala sp. n. from Misima Island, R. truncata sp. n. from the Owen Stanley Range of southeastern New Guinea, R. asymmetrica sp. n. from the Owen Stanley Range of southeastern New Guinea, R. basilaki sp. n. from Basilaki Island, and R. fonticola sp. n. from the Cape Nelson Peninsula of southeastern New Guinea; Brechivelia gen. n. containing type species B. tufi sp. n. from the Cape Nelson Peninsula of southeastern New Guinea; Tanyvelia minima sp. n. from extreme south- eastern New Guinea; T. tagulana sp. n. from Tagula Island; T. papuana sp. n. from the Owen Stanley Range of southeastern New Guinea; Aegilipsicola peninsularis sp. n. from extreme south- eastern New Guinea; A. auga sp. n. from the Owen Stanley Range of southeastern New Guinea; Tarsovelia louisiadensis sp. n. from Misima and Tagula islands; Neusterensifer sagarai sp. n. from extreme southeastern New Guinea; N. femoralis sp. n. from extreme southeastern New Guinea plus Normanby, Sariba and Basilaki islands; N. hunteri sp. n. from southeastern New Guinea plus Normanby, Sariba, Sideia, and Basilaki Islands; N. kula sp. n. from Goodenough and Fer- gusson islands; N. dentrecasteaux sp. n. from Goodenough and Fergusson islands plus the Cape Nelson Peninsula of southeastern New Guinea; N. louisiadae sp. n. from Tagula and Rossel is- lands; N. yela sp. n. from Rossel Island; N. sulcata sp. n. from Tagula Island; N. misima sp. n. from Misima Island; N. muyuw sp. n. from Woodlark Island; N. aviavi sp. n. from the Lakeka- mu Basin of southeastern New Guinea; N. microrivula sp. n. from the Lakekamu Basin of south- eastern New Guinea; N. bowutu sp. n. from the Bowutu Mountains of southeastern New Guinea; N. goilala sp. n. from the Owen Stanley Range of southeastern New Guinea; and N. tufi sp. n. from the Cape Nelson Peninsula of southeastern New Guinea. Shaded dorsal habitus fig- ures are provided for Rheovelia petrophila and Neusterensifer femoralis, and illustrations of diag- nostic morphological characters are given for all new genera and species, accompanied by distri- bution maps. New distribution records are also provided for Tarsovelia alta J. Polhemus & D. Polhemus, and Neusterensifer lubu D. Polhemus & J. Polhemus. A revised key in provided for all genera of New Guinea Microveliinae, and new or revised keys to all species are provided for Rheovelia, Tanyvelia, Aegilipsicola and Tarsovelia; a regional key is also provided for the 16 species of Neusterensifer occurring in the Papuan Peninsula and adjacent island groups. Dr. Dan A. Polhemus (Corresponding author), Dept. of Entomology, MRC 105, National Museum of Natural History, PO Box 37012, Washington, DC 20013-7012. E-mail: [email protected] Dr. John T. Polhemus, 3115 S. York St., Englewood, CO 80110 USA E-mail: [email protected] Key words. – Veliidae; Microveliinae; New Guinea region; East Papua Composite Terrane; tax- onomy; new genera; new species; keys; distribution, illustrations, maps.

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Fig 1. A general delimitation of the land masses comprising the East Papua Composite Terrane as defined by Pigram & Davies (1987).

The island of New Guinea contains a remarkable pattern and timing of the EPCT’s offshore assembly and highly endemic assemblage of genera and species and subsequent integration into greater New Guinea. in the subfamily Microveliinae, many of which are still undescribed. The current paper builds on previous REGIONAL TECTONIC SETTING contributions by Polhemus & Polhemus (1994, 2000a, 2000b) which provided descriptions for six of The EPCT, as defined by Pigram & Davies (1987), the more distinctive genera within this regional biota. comprises all of New Guinea and its proximal islands Additional collecting since that time has brought fur- east of the Aure Scarp and south of the Ramu- ther new genera and new species to our attention, as Markham fault zone (Fig. 1), and represents a com- well as providing additional distributional informa- plex geological unit containing at least six discrete tec- tion for certain previously described species. tonic sub-terranes within it. These include the In particular, the present paper focuses on the mi- partially metamorphosed remains of a Late Cretaceous croveliine fauna of far eastern New Guinea and near- island arc (the Owen Stanley terrane) in the south, a by archipelagos, and is based on recent collections in broad sheet of mantle-derived ophiolite (the Bowutu Central, Oro and Milne Bay Provinces of Papua New terrane) in the north, and an extensive exposure of for- Guinea. These provinces, which include the ‘tail’ of merly submarine basalts (the Kutu terrane) in the east mainland New Guinea plus the adjacent D’Entre- (Pigram & Davies 1987). This tectonic amalgamation casteaux Islands, Marshall Bennett Islands, and now forms the eastern section of New Guinea, from Louisiade Archipelago, encompass a large portion of a approximately the Purari River basin eastward tectonically defined geological province known as the through the Louisiade Archipelago (Fig. 1). East Papua Composite Terrane, or EPCT (Pigram & The earliest vulcanism along the south facing island Davies 1987). Because the EPCT has had a geological arc that would later form the initial core of the EPCT history largely independent of the remainder of New occurred in the Late Cretaceous, approximately 70 Guinea, existing as an isolated microcontinent for a million years (Myr) ago, and continued until this arc considerable period, it has developed a distinctive collided in the Paleocene, 62-57 Myr ago, with a sliv- suite of endemic aquatic Heteroptera, including Mi- er of Australian craton rifted off the northern margin croveliinae, and still retains a unique faunal signature of Australia by the opening of the Coral Sea (Davies et in comparison to the main body of the island (Polhe- al. 1996; Davies et al. 1997; Hill & Hall 2003). This mus & Polhemus 1998). This paper is the first in a se- enlarged offshore microcontinent then converged on ries of reports documenting elements of this biota, and mainland New Guinea from the northeast, eventually attempting to use the resultant biogeographic infor- suturing to the main island in the Late Oligocene to mation to resolve lingering uncertainties regarding the Early Miocene, 28-22 Myr ago (see Davies et al. 1997

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for a series of diagrams illustrating these events). Dur- sures of this metamorphic bedrock. These waterfall ing this same period extensive exposures of Middle zones have proven to support rich microveliine biotas, Eocene to Late Miocene submarine volcanics and sed- both on the wet rock faces bordering the falls them- iments were also uplifted along the southern and east- selves, and in the bedrock-rimmed plunge pools be- ern margins of the progressively enlarging EPCT, form- low. Many of the new species documented in the cur- ing the far eastern tip of the modern Papuan rent paper were collected in association with such Peninsula. habitats. Throughout this same time span, the Solomon Sea and Woodlark Basin were the site of periodic subduc- TAXONOMIC ANALYSIS tion and island arc formation from the Late Creta- ceous or Early Paleocene, 70-65 Myr ago, onward In this paper we describe two new genera and 30 and the islands created there appear to contain frag- new species of Microveliinae from the EPCT. As with ments of many of the terranes more broadly exposed previous papers in this series, the speciose and cos- in the modern Papuan Peninsula. Following the su- mopolitan genus Microvelia is not considered; species turing of the onshore sections of the EPCT to greater in that genus, which has several distinct lineages New Guinea, the offshore eastern section of the EPCT throughout New Guinea, will be dealt with in subse- began to split open along the Woodlark Rift. This quent publications. dismembered the old proto-Louisiades island arc, The initial results from our work on the EPCT Mi- shifting portions of it, such as Woodlark Island, hun- croveliinae clearly support certain aspects of the geo- dreds of kilometers northward toward the Solomons logical history proposed above: (Hamilton 1979, Abers et al. 2002). In very recent 1.) Within the genus Neusterensifer, a group of geological time, during the last 5 Myr from the species referred to herein as the “femoralis group” is Pliocene onward, the D’Entrecasteaux Islands have confined to the greater EPCT. Members of this group risen as exhumed metamorphic core complexes share various combinations of the following synapo- bouyed by mantle compensation along the axis of the morphic character states unique within the genus: or- Woodlark rift at its western end (Abers et al. 2002). ange brown coloration; a setiferous tumescence on the The northward crustal motion produced by the posteromedial margin of the male hind femur; posteri- rapidly opening Woodlark Rift has been accomodat- orly convergent female connexiva with tufts of stiff ed by southward subduction of the Solomon Sea mi- black setae arising from their inner faces and project- croplate beneath the Woodlark Basin via the Tro- ing vertically between the appressed connexival mar- briand Trench, resulting in the recent volcanism seen gins; and narrowing and prolongation of female ab- in the D’Entrecasteaux and Amphlett islands, and in dominal tergite VII into a finger-like posterior the Tufi region of the northeastern Papuan Peninsula projection. Representatives of the femoralis group are at the western apex of the rift. found throughout the EPCT, even on such widely dis- The EPCT thus represents a relatively old geological persed fragments as Woodlark Island, the Louisiade unit within the Melanesian region, and one that ex- Archipelago, and the D’Entrecasteaux Islands; such a isted as a separate island for at least 25 million years, distribution is congruent with the concept of an ini- accounting for its pronounced faunal signature and tially intact land mass harboring a distinct insular bio- significant degree of biotic endemism. Given its tec- ta that was subsequently subdivided by tectonic events. tonically complex history, it is thus likely to contain 2.) The hygropetric genus Rheovelia is endemic to biotic elements that arrived initially by overwater dis- the EPCT and shares a distribution similar to that of the persal during its offshore phase and underwent insu- femoralis group of Neusterensifer, with the exception of lar diversification, and other elements that subse- Woodlark Island. By contrast, another hygropetric quently arrived by overland dispersal following its microveliine genus, Aegilipsicola, which exhibits near- collision with proto-New Guinea; this mixed biota ly identical ecological preferences, occurs in similar was then subjected to vicariance by the dislocation of hygropetric habitats throughout greater New Guinea the islands along the EPCT’s eastern flank. As such, it and on the Raja Ampat Islands to the west, but does provides a challenging area for deciphering faunal his- not occur on the offshore island groups derived from tories and their relationship to regional tectonics. the EPCT. This suggests that Aegilipsicola evolved in The modern topography of the EPCT, including the western or central New Guinea and dispersed east- offshore D’Entrecasteaux Islands and Louisiade Arch- ward into what are now the onshore sections of the ipelago, is generally mountainous, with numerous EPCT following their accretion to New Guinea in the swift, high gradient streams dropping directly toward Late Tertiary, while Rheovelia evolved in the east, on the sea. The surface petrology is predominantly meta- the EPCT itself, and has never managed to disperse be- morphic, and the streams typically cascade over high yond the EPCT terranes it originally occupied. Both waterfalls in their middle reaches, formed by expo- genera may now be found sympatrically on seeps in

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the Owen Stanley Range, although in such cases ...... Phoreticovelia Polhemus & Polhemus Aegilipsicola is invariably the numerically dominant - Body not dorsoventrally flattened, not strongly taxon. A third hygropetric genus, Brechyvelia, is also sexually dimorphic, males only slightly smaller endemic to the EPCT, but known to date only from the than females; dorsum of female thorax and basal Tufi area of the Cape Nelson peninsula, where it oc- abdomen not depressed, without patches of waxy curs sympatrically with Rheovelia. secretion; claws and arolia may or may not be 3.) In contrast to the preceding patterns, the genera modified ...... 2 Tanyvelia and Tarsovelia reach their greatest diversity 2. Large species, length of males 3.25-3.50 mm, of on New Guinea itself, but also have disjunct en- females 3.80-4.10 mm.; middle and hind tarsi demics on the old islands of the Louisiade Archipel- with both arolia and both claws modified into ago. Neither genus is known from the geologically leaf-like blades (Polhemus & Polhemus 2000a, younger islands of the D’Entrecasteaux group, and fig. 3); males with abdominal sternites II-V tumid their current distributions are compatable with the medially, produced ventrally, sternite VIII with a hypothesis that the Louisiades represent isolated rem- large dagger-like structure directed anteriorly, nants of a formerly more extensive Owen Stanley ter- acuminate distally (Polhemus & Polhemus rane, that extended as a continuous range of moun- 2000a, figs. 2, 4) ...... tains far to the southeast of the current eastern ...... Papuavelia Polhemus & Polhemus terminus of New Guinea. - Smaller species, length of males not exceeding 3.0 mm, females not exceeding 3.2 mm.; tarsi with Cladistic analyses of all the above groups will even- arolia and claws not modified into leaf-like blades; tually provide more insight into the processes under- male abdominal sternites sometimes modified, lying these distributional patterns. but not as above ...... 3 3. Middle tarsi subequal in length to middle tibia (Polhemus & Polhemus 1994, fig. 9); light mark- MATERIALS AND METHODS ings on hemelytra restricted to basal angles; fore Collections of Microveliinae were made by visual femur of male at least slightly modified for searching, hand netting, and localized pyrethrin fog- phoresy ...... Tarsovelia Polhemus & Polhemus ging of riparian logs and hygropetric habitats. Speci- - Middle tarsi distinctly shorter than middle tibia; mens obtained were preserved in 80% ethanol, then light markings on hemelytra may be restricted to transported to the Bishop Museum in Honolulu and basal angles, or may occur also on distal portions; subsequently to the Smithsonian Institution in fore femur may or may not be modified for Washington, DC for preparation. Descriptions of phoresy ...... 4 color, structure and setiferation were taken from dry, 4. Second antennal segment clearly longer (1.1/1) point-mounted specimens. All measurements are giv- than first segment; male abdominal segment VIII en in millimeters, and were taken using an ocular mi- strongly asymmetrical (fig. 17) ...... crometer on a Leica M 10 dissecting microscope. CL ...... Brechivelia gen. n. numbers following collection locality data refer to a - Second antennal segment subequal to or shorter coding system used by the authors to cross-reference than first segment; male abdominal segment VIII specimens with photographs and metadata. symmetrical, or nearly so ...... 5 The holotypes of the new species described herein 5. Claws extremely long (Polhemus & Polhemus are deposited in the National Museum of Natural 2000a, fig. 18), clearly much longer than first History, Smithsonian Institution (USNM); paratypes tarsal segment; light hemelytral markings of are held in that collection, and in the Bishop Muse- macropters or micropters entirely bright greyish um, Honolulu (BPBM), and the J. T. Polhemus collec- to light bluish pruinose ...... 6 tion, Englewood, Colorado (JTPC). - Claws relatively short, subequal to or shorter than first tarsal segment; light hemelytral markings not bright pruinose ...... 7 Revised key to genera of New Guinea Microveliinae 6. Head without highly contrasting markings; ros- 1. Body dorsoventrally flattened, strongly sexually di- tral cavity demarcated by strongly raised carina; morphic, males much smaller than females; dor- only macropterous and micropterous morphs sum of female thorax and basal abdomen de- known (Polhemus & Polhemus 1994, fig. 5; Pol- pressed, often with patches of waxy secretion hemus & Polhemus 2000b, fig. 13), with the for- laterally on thorax (Polhemus & Polhemus 2000a, mer predominant ...... fig. 8); middle claws and downcurving arolium ...... Aegilipsicola Polhemus & Polhemus modified into thin narrow blades (Polhemus & - Head with highly contrasting longitudinal light Polhemus 2000a, fig. 10) ...... stripes (fig. 2); rostral cavity not demarcated by

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strongly raised carina; apterous and micropterous mostly yellowish. Wings in macropterous forms dark forms predominant ...... Rheovelia gen. n. brown to black with pale pruinose lavender markings 7. Antennae long, ¾ or more of body length (Pol- consisting of one or two elongate streaks laterally past hemus & Polhemus 1994, fig. 1); comb on fore middle in basal cells, followed by 3-5 small to large ir- tibia ¾ the length of the tibia; pale markings of regular spots on distal ¾ in distal cells. forewing tinged with bluish pruinose, occurring Structural characters. – Micropterous, submacro- only in cells on basal half of wing ...... pterous and macropterous forms known; shape nar- ...... Tanyvelia Polhemus & Polhemus row-elongate (figs. 2, 12). Eyes globose, exserted but - Antennae shorter, clearly less than ¾ of body not beyond anterolateral pronotal angles, separated by length; comb on fore tibia less than ¾ length of slightly more than twice the width of an eye, slightly tibia (except in Neusterensifer compacta), if comb removed from anterior pronotal margin, with moder- on fore tibia ¾ length of tibia then male proctiger ate length ocular setae. Head declivant anteriorly, pos- bearing elongate process and female abdominal terior margin sloping caudo-dorsally, with usual three tergites VII and VIII forming an anal plate (see pairs of facial trichobothria plus numerous short setae; below); pale markings of forewing white, variable vertex with prominent median sulcus; gular region in distribution ...... 8 short, plainly visible, rostral cavity closed posteriorly. 8. Male genitalia rotated 90° sinistrally and highly Rostrum reaching to middle of mesosternum (fig. 12), modified, bearing a long sinuate anteriorly direct- segment I short, enclosed in rostral cavity, segment IV ed sword-like process arising from the proctiger about twice the length of I, almost four times longer (figs. 39-44); female abdominal tergites VII-VIII than II, segment III more than 5 times as long as II. deflected ventrad forming an anal plate; forewing Antennae slender, segments III and IV very long, total frequently with elongate pale streaks in basal cells, length about ¾ of body length. sometimes accompanied by much smaller pale Pronotum very slightly raised medially, with weak spots in distal cells ...... median carina anteriorly; collar weakly formed, absent ...... Neusterensifer Polhemus & Polhemus medially, evident behind eyes laterally; anterior and - Male genitalia usually in normal orientation, in posterior lobes set off by a transverse row of obscure some cases rotated sinistrally to a slight degree, foveae, evanescent laterally; entire dorsum densely set but less than 90°, and always without a long with short to moderate length decumbent golden setae; sword-like process arising from the proctiger; fe- with large triangular patches of silvery setae behind male abdominal tergites VII-VIII not deflected eyes; posterior lobe with numerous small foveae, ventrad, not forming an anal plate; forewing lack- humeri not prominent, broadly rounded posteriorly, ing pale areas basally, but frequently with multi- not modified, covering metanotum. Thoracic venter ple prominent pale spots distally ...... not diagnostic, with weakly formed tubercles on either ...... Microvelia Westwood side of mesosternal midline on posterior margin op- posing an unmodified metasternum. Metasternal scent gland opening (omphalium) small but visible, marked Rheovelia gen. n. by a small tubercle; scent channels prominent, curving (figs. 2-15) slightly anterad to base of metacetabulae. Wing pads Type-species: Rheovelia petrophila Polhemus & Polhemus present but barely visible in some micropters, not visi- sp. n. ble in others, reaching tergite IV in brachypters; wings in macropterous forms reaching past tip of abdomen, Description bearing 4-5 closed cells, consisting of two elongate cells Size. – Micropterous form, length of males 2.00- basally, two more elongate cells centrally, and occa- 2.50 mm., females 2.20-2.50 mm; submacropterous sionally another single smaller cell distally. form, length of males 2.39-2.50 mm., females 2.55- Abdomen broad, several species with two (1+1) 2.61 mm; macropterous form, length of males 2.20 foveae on either side of midline between tergites I and mm., females 2.50-290 mm.; general body character- II appearing silvery; without longitudinal carinae on istics and size not sexually dimorphic, males and fe- tergites; tergites II-VI subequal in length, I shorter, males similar. VII longer. Abdominal sternites set off from lateros- Colour. – Ground colour brown, with yellowish ternites by hair-free glabrous oval lacunae; male ven- markings; head with two prominent yellowish longi- trites VII-VIII variously modified. tudinal stripes (fig. 2); pronotum anteriorly with a Legs slender, long, hind legs much longer than oth- yellowish transverse stripe behind vertex of head, in- ers; anterior femur set beneath with short light setae, terrupted medially; connexiva mostly yellowish, dark unmodified in males; anterior tibia of male with a at intersegmental sutures; female abdominal tergites comb of minute black setae occupying ⅔ of tibial VI-VIII yellowish, much darker in males; venter length; middle and hind femora set ventrally with long

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Fig 2. Rheovelia petrophila, micropterous female, dorsal habitus. Young Sohn illustration. slender setae; all tarsi long (fig. 2), claws extremely trad, concealing gonocoxae and genital opening; anal long, slender; both up- and down curving arolia long, cone often visible, sometimes sharp, projecting. slender (fig. 3). Male ventrite VII variously modified (figs. 4-11), Etymology genital segments small to prominent, ventrite VIII The generic name Rheovelia is derived from rheos variously modified, bent sharply ventrad in several (Gr.), flow, stream, referring to the wet vertical rock species; pygophore unmodified; parameres small, tri- habitat, and Velia, the nominate genus of the family. angular, symmetrical, hidden; proctiger not modified; Gender feminine. tergite IX rounded, protruding. Female with tergite VIII on same plane as VII, sometimes truncate poste- Comparative notes riorly; proctiger forming an anal plate or lid, bent ven- Rheovelia differs from all other microveliine genera

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bedrock exposures typical of high gradient streams in eastern New Guinea and proximal islands. They typi- cally prefer open bedrock substrates without a covering of moss or algae, generally in shaded situations, and have never been observed skating on pools or other standing water surfaces, even when such pools imme- diately adjoin their moist bedrock habitats. In this re- gard their ecological preferences are nearly identical to those of the genus Aegilipsicola, which occurs through- out the main island of New Guinea and is occasionally sympatric with Rheovelia in the Owen Stanley Range. For additional notes see discussions under individ- ual species below.

Distribution Far eastern New Guinea and adjacent islands (Basi- laki, Normanby, Fergusson, Misima, Tagula, Rossel) (figs. 13-15). The genus Rheovelia as currently understood is confined to land masses derived from the East Papua Composite Terrane, including the Papuan Peninsula, Basilaki Island, the D’Entrecasteaux Islands, and the Louisiade Archipelago. Despite intensive recent searches in suitable habitats on the isolated EPCT frag- ment of Woodlark Island, the genus has yet to be col- lected there.

Key to the species of Rheovelia Fig. 3. Rheovelia petrophila, male middle tarsus. Note: the following key is primarily applicable to males, but will work for females of certain species. For of the region by possessing very long slender claws (fig. many species, particularly those occurring on particu- 3) coupled with a short gula and rostral cavity not de- lar islands, geographical provinence will indicate the marcated by a strongly raised carina; it is known from species involved; in other cases, females will need to the micropterous, submacropterous and macropterous be indentified by association with males. morphs, with the former predominant in most popula- tions sampled. The closest Papuan genus in general ap- 1. Male abdominal ventrite VIII largely withdrawn pearance and ecology, Aegilipsicola J. Polhemus & D. into ventrite VII, directed ventrally, bearing a Polhemus, has similarly long slender claws (shared with large anteriorly directed process extending anteri- Aegilipsovelia J. Polhemus of the New World tropics), orly over at least the posterior margin of ventrite but with a long gula and rostral cavity clearly demar- VII (figs. 4-7) ...... 2 cated by a strongly raised carina, and is not known only - Male abdominal ventrite VIII not withdrawn from micropterous morphs, with macropterous into ventrite VII, not directed ventrally, without morphs predominating instead. The hemelytral mark- an obvious anteriorly directed process, instead ings of both genera, when visible wings are present, are with a small raised ridge or process on posterior bright pruinose greyish to light bluish or lavender. margin (figs. 8-11) ...... 5 A related genus, Brechivelia (described subsequently 2. Anteriorly directed process of male ventrite VIII in this paper), has long but somewhat thickened claws short, triangular, barely reaching over posterior (fig. 18) and a similar rheophilic ecology. It is easily margin of ventrite VII (fig 7); female with distinguished by its asymmetrical male abdominal seg- proctiger directed ventrally, not visible in lateral ment VIII, and extremely long antennae (figs. 16, 17). view; Basilaki Island ...... R. basilaki sp. n. - Anteriorly directed process of male ventrite VIII Biological notes long, reaching at least middle of ventrite VII (figs. Species of Rheovelia are obligatorily hygropetric, oc- 4-6); female with proctiger directed posteriorly, curring on the sloping to vertical seeping rheocrenes visible in lateral view ...... 3 that form adjacent to the waterfalls and riparian 3. Anteriorly directed process of male ventrite VIII

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4 5

6 7

8 9

10 11

Figs. 4–11. Rheovelia species, ventral view of male terminal abdomen, with detail of processes on ventrite VIII. – 4, R. petrophila; 5, R. asymmetrica; 6, R. truncata; 7, R. basilaki; 8, R. insularis; 9, R. robinae; 10, R. fonticola; 11, R. anomala.

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truncate to slightly concave distally (fig. 6); west- micropterous females, Upalai River at Haluwia, 13.7 km. E. of Alotau on East Cape road, midreach and waterfall approx. ern Owen Stanley Range ...... R. truncata sp. n. 2.5 km. above mouth, 45-75 m. [150-250 ft.], 10°19’39”S, - Anteriorly directed process of male ventrite VIII 150°34’36”E, water temp. 24.5 °C., 5 April 2002, 09:45- in the shape of a broad-headed dagger, distally 16:30 hrs., CL 7163, D. A. & J. T. Polhemus (USNM). triangular, slightly to strongly asymmetrical (figs. 4, 5) ...... 4 4. Anteriorly directed process of male ventrite VIII Description long, almost reaching anterior margin of ventrite Size. – Micropterous male, length 2.15-2.20 mm (x- VII (fig. 4); ventrite VII simple, not tumid, with = 2.19, n = 5); width 0.85-0.90 mm (x– = 0.86, n = 5). paired (1+1) depressions laterally (fig. 4); female Micropterous female, length 2.30-2.35 mm (x– = 2.33, connexiva distally raised, acute, projecting (figs. n = 5); width 0.90-1.00 mm, (x–) = 0.95, n = 5). 2, 12); extreme eastern Papuan Peninsula ...... Colour. – Micropterous male: ground colour dark ...... R. petrophila sp. n. brown, head, anterior pronotum, connexiva, abdomi- - Anteriorly directed process of male ventrite VIII nal tergites VI -VIII with yellowish brown to orange shorter, reaching about half way to anterior mar- brown markings; venter mostly yellowish to leucine. gin of ventrite VII (fig. 5); ventrite VII with Head with two broad longitudinal stripes on vertex, paired (1+1) tumescences anteriorly, set with long lower part yellowish brown. Pronotum dark blackish semi-erect setae; female connexiva distally flat, not brown; laterally narrowly lighter brown; anterior lobe raised, blunt, not projecting; western Owen Stan- transversely yellowish brown between eyes, interrupt- ley Range ...... R. asymmetrica sp. n. ed medially; large triangular patches of silvery hairs 5. Posterior process of male ventrite VIII simple, a present anterolaterally, extending behind vertex. Wing ventrally directed trucate triangular ridge with a pads tiny, barely visible. Abdomen brown; paired distinct deep notch; ventrite VII tricarinate (Fig. foveae (1+1) between tergites I and II with silvery 9); D’Entrecasteaux Islands ...... R. robinae sp. n. hairs; connexiva broadly orange brown, irregularly - Posterior process of male ventrite VIII with one darker at sutures; tergites VI-VIII extensively marked or two obvious notches, resulting in two or three with orange brown. Legs leucine to yellow, upper sur- protuberances (figs. 8, 10, 11); ventrite VII at faces marked with brown distally on femora, becom- most bicarinate ...... 6 ing more uniform on tibiae and tarsi. 6. Posterior process of male ventrite VIII with two Structural characters. – Micropterous male: head notches, resulting in a rounded median protuber- of moderate length, declivant anteriorly; length 0.52, ance flanked by two smaller protuberances (Fig. width 0.60; width of eye/interocular space, 10); Cape Nelson Peninsula ..... R. fonticola sp. n. 0.12/0.37. Pronotum long, posterior margin evenly - Posterior process of male ventrite VIII with only curving, not angulate, humeri obscure; length (mid- a median notch (figs. 8, 11) ...... 7 line)/width, 0.45/0.85. 7. Posterior process of male ventrite VIII with two Abdominal tergites with II-V subequal, tergite VI triangular protuberances (Fig. 8); Tagula and longest, tergite VII retracted and not visible (0.08, Rossel Islands ...... R. insularis sp. n. 0.07, 0.06, 0.10, 0.40, hidden, respectively), abdomi- - Posterior process of male ventrite VIII with two nal venter bearing short, fine, pale setae; ventrites VI- sharp pointed protuberances (Fig. 11); Misima VII slightly flattened and raised; ventrite VII excavated Island ...... R. anomala sp. n posteriorly to accomodate anteriorly directed process of segment VIII; first genital segment (segment VIII) curved downward into a vertical orientation, pos- Rheovelia petrophila sp. n. teroventral margin bearing an asymmetrical anteriorly- (figs. 2-4, 12, 13) directed, glabrous, angulate dagger-like projection (fig. Material examined. – Holotype, micropterous male: 4), clothed with long setae laterally, this projection PAPUA NEW GUINEA, Milne Bay Prov., headwater reach of conforming to the medial concavity of ventrite VII. Goilayoli River above bridge crossing on road from Watu- nou to Huhuna, 18.5 km. ENE of Alotau, 275 m. [900 ft.], Legs clothed with numerous short, appressed, pale 10°18’43”S, 150°37’16”E, water temp. 24.5 °C., 6 April setae, intermixed on ventral surface of fore femur with 2002, 10:00-13:00 hrs., CL 7161, D. A. & J. T. Polhemus longer, erect, pale, pilose setae; all femora with silvery (USNM, JTPC). – Paratypes: PAPUA NEW GUINEA, Milne Bay setae on basal half, best seen in light from behind in- Prov.: 13 micropterous males, 8 micropterous females, same sect; all legs unarmed, lacking setal tufts or other mod- data as holotype, D. A. & J. T. Polhemus (USNM); 10 mi- ifications. Antennae thickly clothed with short to long cropterous males, 17 micropterous females, Cloudy Moun- tains, headwater tributary to upper Watuti River, S. of Gele- setae, segment I also bearing 3-4 long, spine-like black malaia, 715 m. [2350 ft.], 10°29’50”S, 150°13’58”E, water setae. temp. 22 °C., 10 April 2002, 16:00-17:30 hrs., CL 7175, D. Antennal formula I : II : III : IV; 0.30 : 0.15 : 0.55 A. Polhemus (USNM, JTPC, BPBM); 1 micropterous male, 3 : 0.65.

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Fig. 12. Rheovelia petrophila, micropterous female, lateral habitus. Young Sohn illustration.

Proportions of legs as follows: Femur, tibia, tarsal nel, and several small, steeply dropping tributaries en- 1, tarsal 2 of fore leg, 0.62 : 0.52 : 0.28 : 0.0; of mid- tered from the surrounding rain forest hills, with stair- dle leg, 0.90 : 0.75 : 0.05 : 0.12; of hind leg, 1.10 : step bed profiles of alternating vertical seeps and small 1.50 : 0.10 : 0.40. pools. The type series of R. petrophila was taken from Paramere small, triangular, hidden; proctiger not the vertical seeps along both the main channel and the modified. tributaries, with the insects being particularly abun- Micropterous female: Similar to male in general dant next to small, trickling waterfalls on the tribu- structure and coloration, but larger. Connexiva raised taries. The insects appeared to be obligatorily hy- to about 45°, broadly separated posteriorly, evenly ta- gropetric, and were never observed skating on pools. pering toward tip of abdomen, posterolateral angles acute, produced postriorly; abdominal venter unmodi- Rheovelia robinae sp. n. fied. (figs. 9, 13) Etymology Material examined. – Holotype, micropterous The name petrophila describes the affinity of this male: PAPUA NEW GUINEA, Milne Bay Prov., Fergusson species for damp vertical rock surfaces. Island, east coast, cascading tributary to Mebulibuli Creek, S. of Basima, 0.9 km. upstream of mouth, 15 Distribution m. [50 ft.], water temp. 24.5 °C., 24 August 2002, Eastern New Guinea (fig. 3). 13:30-14:30 hrs., 9°30’41.7”S, 150°52’06.0”E, CL 7181, D. A. & J. T. Polhemus (USNM). – Paratypes: Comparative notes PAPUA NEW GUINEA, Milne Bay Prov., D’Entre- Rheovelia petrophila is easily separated from the casteaux Islands: 17 micropterous males, 18 micro- other species of the genus by the ventrally directed ab- pterous females, same data as holotype (USNM, JTPC, dominal segment VIII of the male, bearing a promi- BPBM); 6 micropterous males, 7 micropterous females, nent dagger-like projection, and female with distinct- D’Entrecasteaux Islands, Fergusson Island, east coast, ly protruding posterior connexival angles. Yaya River, N. of Mebulibuli Point, 0-60 m. [0-200 ft.], water temp. 25.8 °C., 23 August 2002, 10:00- Biological notes 16:30 hrs., 9°30’34.3”S, 150°52’51.1”E, CL 7178, D. The Goilayoli River at the type locality was a clear, A. & J. T. Polhemus (USNM); 1 micropterous male, 1 cool upland stream set in a narrow, channel partially micropterous female, Fergusson Island, east coast, up- incised in bedrock, and heavily shaded by primary up- per Awaetowa River and rocky tribs., W. of Basima, land rain forest. The bed profile of the main stream 575-650 m. [1885-2130 ft.], water temp. 22 °C., 27 consisted of pools 0.5-1.3 m. deep alternating with January 2003, 10:00-14:00 hrs., 9°30’34.3”S, 150° small cascades at transverse bedrock sills and shallow 48’17.3”E, CL 7225, D. A. & J. T. Polhemus (USNM); riffles of cobble and gravel. Numerous seeping bedrock 1 micropterous female, Normanby Island, south faces were present along the margins of the main chan- coast, upper Apatabuia River and rocky tribs., above

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Bunama, 1.38 km. above mouth, 60-90 m. [195-295 Antennal formula I : II : III : IV; 0.30 : 0.20 : 0.55 ft.], water temp. 23.5 °C., 29 January 2003, 09:00- : 0.63. 01:00 hrs., 10°07’04.0”S, 151°09’07.2”E, CL 7228, Proportions of legs as follows: Femur, tibia, tarsal D. A. & J. T. Polhemus (USNM); 12 micropterous 1, tarsal 2 of fore leg, 0.70 : 0.50 : 0.28 : 0.0; of mid- males, 16 micropterous females, Normanby Island, dle leg, 1.00 : 0.90 : 0.05 : 0.30; of hind leg, 1.20 : east coast, Golupona Stream above Yeluyelua, 90 m. 2.06 : 0.08 : 0.40. [295 ft.], water temp. 23 °C., 30 January 2003, 08:00- Paramere small, triangular, hidden; proctiger not 13:00 hrs, 10°02’53.3”S, 151°15’35.3”E, CL 7232, J. modified. T. Polhemus (USNM). Micropterous female: Similar to male in general structure and coloration, but larger. Connexiva raised Description to about 45°, broadly separated posteriorly, evenly ta- Size. – Micropterous male, length 2.33-2.40 mm pering toward tip of abdomen, posterolateral angles (x– = 2.37, n = 4); width 0.70-0.75 mm (x– = 0.74, n = truncate, not produced posteriorly; abdominal venter 4). Micropterous female, length 2.40-2.50 mm (x– = unmodified. 2.45, n = 4); width 0.80-0.90 mm, (x– = 0.85, n = 4). Colour. – Micropterous male: ground colour dark Etymology brown, head, anterior pronotum, connexiva, abdom- The name robinae is in honor of Robin Polhemus, inal tergites VI-VIII with yellowish brown to orange the wife of the first author. brown markings; venter mostly yellowish to leucine. Head with two broad longitudinal stripes on vertex, Distribution lower part yellowish brown. Pronotum dark blackish Eastern New Guinea, D’Entrecasteaux Islands (fig. brown; laterally narrowly lighter brown; anterior lobe 13). transversely yellowish brown between eyes, interrupt- ed medially; large triangular patches of silvery hairs Comparative notes present anterolaterally, extending behind vertex. Rheovelia robinae may be separated from the other Wing pads not visible. Abdomen brown; paired species of the genus by the tricarinate abdominal ven- foveae (1+1) between tergites I and II without silvery trite VII of the male, male abdominal ventrite VIII hairs; connexiva broadly orange brown, irregularly bearing a small truncate projection without a notch, darker at sutures; tergites VI-VIII extensively marked and the widely separated truncate posterior connexi- with orange brown. Legs leucine to yellow, upper sur- val angles of the female. faces marked with brown distally on femora, becom- ing more uniform on tibiae and tarsi. Biological notes Structural characters. – Micropterous male: head The type locality was a small, splashing cascade of moderate length, declivant anteriorly; length 0.55, partially shaded by lightly disturbed lowland rain for- width 0.60; width of eye/interocular space, est, and dropping 1.5 m. over a vertical exposure of 0.12/0.40. Pronotum long, posterior margin evenly dark bedrock. This cascade lay just above a riverside curving, not angulate, humeri obscure; length (mid- trail on a tributary to Mebulibuli Creek, a much larg- line)/width, 0.50/0.80. er, rocky stream draining the hills of eastern Fergus- Abdominal tergites with II-V subequal, tergite VI son Island. The insects were found in abundance on longest, tergite VII retracted and not visible (0.20, the wet bedrock exposures adjacent to the cascade it- 0.12, 0.15, 0.15, 0.30, hidden, respectively), abdom- self, in company with Ochterus and Hebrus species. At inal venter bearing short, fine, pale setae; ventrites IV- the nearby Yaya River, R. robinae was taken in a sim- VI medially depressed; ventrite VII tricarinate, poste- ilar situation at a small cascade flanked by seeping riorly transversely carinate and slightly excavated (fig. rock faces on the main channel of a clear, 0.3-1.0 m. 9); first genital segment (segment VIII) extending deep creek surrounded by primary rain forest. posteriorly, posteroventral margin bearing a ventrally directed truncate triangular protuberance without a Rheovelia insularis sp. n. notch (fig. 9). (figs. 8, 13) Legs clothed with numerous short, appressed, pale setae, intermixed on ventral surface of fore femur Material examined. – Holotype, micropterous with longer, erect, pale, pilose setae; all femora with male: PAPUA NEW GUINEA, Milne Bay Prov., Louisiade silvery setae on basal half, best seen in light from be- Archipelago, Rossel [Yela] Island, Woa River and hind insect; all legs unarmed, lacking setal tufts or rocky tributaries, from head of estuary to 4 km. up- other modifications. Antennae thickly clothed with stream, 0-150 m. [0-500 ft.], 11°20’36.7”S, short to long setae, segment I also bearing 3-4 long, 154°07’06.4”E, water temp. 23 C., 31 August-1 Sep- spine-like black setae. tember 2002, CL 7196, D. A. & J. T. Polhemus

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(USNM). – Paratypes: PAPUA NEW GUINEA, Milne Bay a ventrally directed truncate triangular protuberance, Prov., Louisiade Archipelago: 23 micropterous males, with a notch (fig. 8). 21 micropterous females, Rossel Island, same data as Legs clothed with numerous short, appressed, pale holotype (USNM, JTPC, BPBM); 14 micropterous males, setae, intermixed on ventral surface of fore femur 16 micropterous females, Tagula [Sudest] Island, with longer, erect, pale, pilose setae; all femora with Kolukolu Creek, near Araeda village, 0-105 m. [0- some silvery setae on basal half, best seen in light from 350 ft.], 11°26’44.7”S, 153°25’56.8”E, water temp. behind insect; all legs unarmed, lacking setal tufts or 23 °C., 30 August 2002, 11:00-16:00 hrs., CL 7194, other modifications. Antennae thickly clothed with D. A. & J. T. Polhemus (USNM, BPBM); 16 micropter- short to long setae, segment I also bearing 3-4 long, ous males, 5 micropterous females, Tagula [Sudest] spine-like black setae. Island, Kalitau Creek, bedrock face on small trib. ap- Antennal formula I : II : III : IV; 0.35 : 0.20 : 0.62 prox. 1 km. upstream from head of estuary, 30 m. : 0.70. [100 ft.], 11°21’51.7”S, 153°14’26.0”E, water temp. Proportions of legs as follows: Femur, tibia, tarsal 24.4 °C., 29 August 2002, 09:30-14:30 hrs., CL 1, tarsal 2 of fore leg, 0.70 : 0.55 : 0.27 : 0.0; of mid- 7190, D. A. & J. T. Polhemus (USNM). dle leg, 0.90 : 0.80 : 0.05 : 0.32; of hind leg, 1.13 : 1:50 : 0.07 : 0.42. Description Paramere small, triangular, hidden; proctiger not Size. – Micropterous male, length 2.15-2.25 mm modified. (x– = 2.20, n = 4); width 0.60-0.70 mm (x– = 0.64, n = Micropterous female: Similar to male in general 4). Micropterous female, length 2.35-2.40 mm (x– = structure and coloration, but larger. Connexiva raised 2.39, n = 4); width 0.70-0.80 mm, (x– = 0.76, n = 4). to about 30° to 45°, narrowly separated posteriorly, Colour. – Rossel Island specimens: Micropterous most widely separated at tergite IV, then evenly ta- male: ground colour dark brown, head, anterior pering toward tip of abdomen, posterolateral angles pronotum, connexiva, abdominal tergites with yel- acute, curved inward; abdominal venter unmodified. lowish brown to orange brown markings; venter mostly yellowish to leucine. Head with two broad Etymology longitudinal stripes on vertex, lower part yellowish The name insularis is in recognition of the occur- brown. Pronotum dark blackish brown; laterally nar- rence of this species on the most remote islands of the rowly lighter brown; anterior lobe transversely yel- Louisiade Archipelago. lowish brown between eyes, interrupted medially; large triangular patches of silvery hairs present antero- Distribution laterally, extending behind vertex. Wing pads not vis- Tagula and Rossel Islands, Louisiade Archipelago ible. Abdomen brown; paired foveae (1+1) between (fig. 13). tergites I and II without silvery hairs; connexiva broadly yellowish, infused with brown, with an irreg- Comparative notes ular median longitudinal stripe; tergite I with two Rheovelia insularis may be separated from the other (1+1) quadrate yellowish spots; tergites II-VIII exten- micropterous species of the genus by the triangular sively marked with orange brown, becoming lighter carina on ventrite VII of the male, segment VIII bear- distally. Legs leucine to yellow, upper surfaces ing a small truncate projection with a notch, and fe- marked with brown distally on femora, becoming male with weakly convergent and acute protruding more uniform on tibiae and tarsi. posterior connexival angles. It is known only from the Structural characters. – Micropterous male: head micropterous form, separating it from R. anomala. of moderate length, declivant anteriorly; length 0.50, Although there are slight differences between the width 0.57; width of eye/interocular space, populations of this species from Rossel and Tagula is- 0.15/0.40. Pronotum long, posterior margin evenly lands, we have not considered this variation significant curving, not angulate, humeri obscure; length (mid- enough to warrent separate species or subspecies sta- line)/width, 0.48/0.70. tus. In the Tagula specimens the ground coloration is Abdominal tergites with II-V subequal, tergite VI much darker in comparison to those from Rossel, and longest, tergite VII retracted and not visible (0.15, the light markings are yellowish brown to orange 0.10, 0.12, 0.12, 0.27, hidden, respectively); connex- brown. The general pattern of the pale markings on iva most widely separated at tergite IV, then evenly the head, pronotum and abdomen of the Tagula converging toward tip of abdomen but not meeting; forms is similar to that of the Rossel forms, but these abdominal venter bearing short, fine, pale setae; ven- markings are less extensive and barely evident on ab- trite VII with an anteriorly directed triangular carina, dominal tergites II-IV and on the inner part of the excavate behind; first genital segment (segment VIII) connexiva. In Tagula males, the abdominal connexiva extending posteriorly, posteroventral margin bearing converge evenly toward the posterior apex of the ab-

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150° 151° 152° 153° 154°

KIRIWINA IS. TROBRIAND ISLANDS

° 9° WOODLARK IS. 9

GOODENOUGH IS.

FERGUSSON IS. COLLINGWOOD BAY

D'ENTRECASTEAUX ISLANDS EGUM ATOLL

NORMANBY IS. NEW GUINEA GOODENOUGH BAY Rheovelia robinae SOLOMON SEA 10°

MILNE BAY ORANGERIE BAY Rheovelia ENGINEER GROUP DEBOYNE IS. MISIMA IS. insularis

LOUISIADE ARCHIPELAGO CONFLICT GROUP 11° 11° Rheovelia ROSSEL IS. petrophila

TAGULA IS. CORAL SEA

150° 151° 152° 153° 154°

Fig. 13. Distribution of Rheovelia species in the far eastern New Guinea region.

domen, whereas in the Rossel males they bow outward ly dry waterfall face with only remnant seeping flow and are most widely separated adjacent to tergite IV. rose above a narrow plunge pool. Another series was Despite these minor differences, all other external taken further east on Tagula from vertical faces of wet, characteristics, including the pregenital structures, are unshaded metamorphic bedrock at Kolukolu Creek, the same between the two populations. where this stream cascaded over a 20 m. high waterfall into the head of a mangrove estuary. In all these cases Biological notes the specimens of Rheovelia were found only on open The Woa River at the type locality was a broad, bedrock faces with a thin, nearly laminar flow of wa- clear river in a cobble bed, surrounded by undisturbed ter, further illustrating the hygropetric ecology of this primary rain forest and flowing northeastward from genus. the slopes of Mt. Mbo on western Rossel Island. In its upper midreach the river channel became set with Rheovelia anomala sp. n. scattered large boulders and confined between (figs. 11, 15) bedrock walls, with the bed profile consisting of alter- nating small rapids and deep pools. The type series of Material examined. – Holotype, submacropterous R. insularis was taken from seeps along the bedrock male: PAPUA NEW GUINEA, Milne Bay Prov., Louisiade walls bordering the channel in this reach, and from Archipelago, Misima Island, north coast, Guwana sloping to vertical bedrock seeps on steeply dropping Creek, near Nulia village, 15-30 m. [50-100 ft.], wa- tributaries that joined the main river at various points. ter temp. 22 °C., 3 September 2002, 10:00-12:00 On Tagula Island, a series of R. insularis was taken hrs., 10°39’22.8”S, 152°41’16.5”E, CL 7201, D. A. from a heavily shaded, steeply dropping tributary to & J. T. Polhemus (USNM). – Paratypes: PAPUA NEW Kalitau Creek, a small, clear stream in a very rugged GUINEA, Milne Bay Prov., Louisiade Archipelago, bed of metamorphic bedrock and cobbles. The insects Misima Island: 82 submacropterous males, 88 sub- were found along the tributary at a point where a near- macropterous females, same data as holotype (USNM,

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JTPC, BPBM); 6 submacropterous males, 6 sub- longer, erect, pale, pilose setae; all femora with silvery macropterous females, south coast, wet bedrock faces setae on basal half, best seen in light from behind in- on headwater trib. of Keyana Creek, above Bwagab- sect; all legs unarmed, lacking setal tufts or other mod- waga village, 305-335 m. [1000-1100 ft.], water ifications. Antennae thickly clothed with short to long temp. 22.5 °C., 4 September 2002, 10:00-12:00 hrs., setae, segment I also bearing 3-4 long, spine-like black 10°40’33.8”S, 152°40’44.9”E, CL 7206, D. A. & J. setae. T. Polhemus (USNM); 1 macropterous male, 1 Antennal formula I : II : III : IV; 0.37 : 0.22 : 0.60 macropterous female, 25 submacropterous males, 32 : 0.65. submacropterous females, south coast, small waterfall Proportions of legs as follows: Femur, tibia, tarsal at Awamuluwa, W. of Bwagabwaga village, 15 m. [50 1, tarsal 2 of fore leg, 0.75 : 0.62 : 0.32 : 0.0; of mid- ft.], water temp. 24.6 °C., 4 September 2002, 10:00- dle leg, 0.95 : 0.90 : 0.06 : 0.35; of hind leg, 1.25 : 10:15 hrs., 10°41’07.8”S, 152°40’12.1”E, CL 7208, 1.65 : 0.08 : 0.40. D. A. & J. T. Polhemus (USNM). Paramere small, triangular, hidden; proctiger not modified. Description Submacropterous female: Similar to male in gener- Size. – Submacropterous male, length 2.35-2.45 al structure and coloration, but larger. Connexiva mm (x– = 2.40, n = 4); width 0.70-0.75 mm (x– = 0.74, raised to about 30° basally, infolded and convergent n = 4). Submacropterous female, length 2.70-2.80 posteriorly, evenly tapering toward tip of abdomen, mm (x– = 2.72, n = 4); width 0.80-0.90 mm, (x– = 0.86, posterolateral angles truncate, curved inward; abdom- n = 4). inal venter unmodified. Colour. – Submacropterous male: ground colour Macropterous female: Similar to submacropterous dark brown, head, anterior pronotum, connexiva, ab- female in general structure and coloration with follow- dominal tergites with yellowish brown to orange ing exceptions: length 2.90 mm.; width (across prono- brown markings; venter mostly yellowish to leucine. tal humeri) 1.10 mm.; pronotum expanded, dark Head with two prominent longitudinal stripes on ver- brown with fine punctations overlain with short re- tex, lower part yellowish brown. Pronotum dark cumbent gold setae, posterior margin angulate, blackish brown; laterally narrowly lighter brown; ante- humeri prominent, rounded. Wings extending be- rior lobe transversely yellowish brown between eyes, yond tip of abdomen, forewing bearing 4 closed cells, interrupted medially; large triangular patches of sil- general coloration brown, veins black, outer basal cell very hairs present anterolaterally, extending behind with large silvery pruinose patch filling basal angle and vertex. Wing pads extending onto tergite IV, dark, extending posteriorly half the length of the cell, outer narrowly bright bluish pruinose basally, tiny grey spot distal cell with filled with bluish pruinosity on basal at distal ⅔, distally bright white. Abdomen brown; half, inner basal cell without pale markings, posterior paired foveae (1+1) between tergites I and II not visi- half of the wing with 4 small, irregular, bright white ble; connexiva broadly yellowish, irregularly darker at patches arranged transversely, lying basal to a very sutures; tergites V-VIII extensively marked with or- large, longitudinally elongate bright white patch ex- ange brown, becoming lighter distally. Legs leucine to tending medially nearly to wing apex. yellow, upper surfaces marked with brown distally on Macropterous male: Similar to macropterous male femora, becoming more uniform on tibiae and tarsi. in general structure and colour. Structural characters. – Micropterous male: head of moderate length, declivant anteriorly; length 0.50, Etymology width 0.62; width of eye/interocular space, 0.15/0.60. The name anomala refers to the predominantly sub- Pronotum long, posterior margin forming an obtuse macropterous state of this species, in contrast to other angle, humeri prominent; length (midline)/width, Rheovelia species which are predominantly micropter- 0.75/0.80. ous. Abdominal tergites with II-V subequal, tergite VI longest, tergite VII shortest (0.15, 0.15, 0.12, 0.15, Distribution 0.30, 0.10, respectively); abdominal venter bearing Eastern New Guinea, Misima Island (fig. 15). short, fine, pale setae; ventrite VII medially flattened, margins of flat area weakly carinate, posterior margin Comparative notes V-shaped, excavate behind; first genital segment (seg- Rheovelia anomala may be separated from the other ment VIII) extending posteriorly, posteroventral mar- species of the genus by the medially flattened and pos- gin bearing a ventrally directed truncate triangular teriorly V-shaped ventrite VII of the male (fig. 11), protuberance, with a notch. segment VIII bearing a small truncate projection with Legs clothed with numerous short, appressed, pale a notch (fig. 11), and female with weakly convergent setae, intermixed on ventral surface of fore femur with and acute protruding posterior connexival angles.

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Biological notes Structural characters. – Micropterous male: head Guwana Creek at the type locality was a swift, of moderate length, declivant anteriorly; length 0.60, moderately clear stream dropping steeply out of the width 0.70; width of eye/interocular space, sheer mountains of western Misima through a bed of 0.20/0.42. Pronotum long, posterior margin evenly metamorphic boulders and bedrock. The stream pro- curving, not angulate, humeri obscure; length (mid- file consisted of waterfalls, rapids and pools, some of line)/width, 0.55/0.90. the latter up to 2.0 m. deep. The waterfalls were bor- Abdominal tergites with II longer than III-VI, ter- dered by rheocrenes and standing pools in bedrock, gite VII longest (0.25, 0.15, 0.15, 0.15, 0.20, 0.35 re- and the type series of R. anomala was taken from spectively), abdominal venter bearing short, fine, pale these open rheocrenes and spray-wetted boulders ad- setae; ventrite VII weakly depressed medially, excavate jacent to the falls, where the species was extremely behind; first genital segment (segment VIII) directed abundant. ventrally, bearing an asymmetrical truncate anteriorly directed process (fig. 6), curved in side view, extending anteriorly over much of ventrite VII. Rheovelia truncata sp. n. Legs clothed with numerous short, appressed, pale (figs. 6, 14, 85) setae, intermixed on ventral surface of fore femur with Material examined. – Holotype, micropterous longer, erect, pale, pilose setae; all femora with some male: PAPUA NEW GUINEA, Central Prov., Owen Stan- silvery setae on basal half, best seen in light from be- ley Range, trib. to Mas River, 1.9 km. SW of Fane, hind insect; all legs unarmed, lacking setal tufts or oth- 1220-1250 m. [4000-4100 ft.], wet rock face above er modifications. Antennae thickly clothed with short small side spring, 8°33’33.0”S, 147°04’12.2”E, water to long setae, segment I also bearing 3-4 long, spine- temp. 20.5 °C., 5 October 2003, 10:45-12:30 hrs., like black setae. CL 7258, D. A. Polhemus (USNM). – Paratypes: PAPUA Antennal formula I : II : III : IV; 0.35 : 0.20 : 0.60 NEW GUINEA, Central Prov.: 1 micropterous male, 1 : 0.75. micropterous female, same data as holotype, D. A. Proportions of legs as follows: Femur, tibia, tarsal Polhemus (USNM, JTPC); 1 micropterous male, Owen 1, tarsal 2 of fore leg, 0.75 : 0.55 : 0.30 : 0.0; of mid- Stanley Range, small cascade and seeps on trib. to dle leg, 1.00 : 1.00 : 0.10 : 0.35; of hind leg, 1.40 : Hane River, 1.3 km. SSE of Fane, 1280 m. [4200 ft.], 1:70 : 0.12 : 0.40. 8°33’49.2”S, 147°05’14.0”E, water temp. 20 °C., 4 Paramere small, triangular, hidden; proctiger not October 2003, 08:30-09:30 hrs., CL 7255, D. A. Pol- modified. hemus (USNM). Micropterous female: Similar to male in general structure and coloration, but larger. Connexiva raised Description to about 20°, widely separated posteriorly, evenly ta- Size. – Micropterous male, length 2.35-2.40 mm pering toward tip of abdomen, posterolateral angles (x– = 2.38, n = 3); width 0.85-0.90 mm (x– = 0.88, n = weakly acute, but not pointed, forming weak triangles, 3). Micropterous female, length 2.35 mm (n = 1); not convergent; abdominal ventrite VII weakly sinu- width 0.95 mm, (n = 1). ate posteriorly; protruding proctiger forming an acute Colour. – Micropterous male: ground colour dark angle. brown, head, anterior pronotum, connexiva, abdomi- nal tergites with yellowish brown to orange brown Etymology markings; venter mostly yellowish to leucine. Head The name truncata refers to the truncate male with two broad longitudinal stripes on vertex, lower process on ventrite VIII. part yellowish brown. Pronotum dark blackish brown; laterally narrowly lighter brown; anterior lobe trans- Distribution versely yellowish brown between eyes, width of stripe Southeastern New Guinea, Central Province (fig. equal to separation from anterior margin, interrupted 14). medially; large triangular patches of silvery hairs pre- sent anterolaterally, extending behind vertex. Wing Comparative notes pads not visible. Abdomen brown; paired foveae (1+1) Rheovelia truncata may be separated from the other between tergites I and II with silvery hairs; connexiva micropterous species of the genus by the unique broadly yellowish, infused with irregular brown marks process of the male segment VIII, and the female with at each suture; tergite I with two (1+1) weak lighter widely separated and weakly acute posterior connexi- spots; tergites V-VIII extensively marked with orange val angles, slightly more prominent than in R. robi- brown, becoming lighter distally. Legs leucine to yel- nae. It is known only from the micropterous form, low, upper surfaces marked with brown distally on separating it from R. anomala. femora, becoming more uniform on tibiae and tarsi.

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Biological notes curving, not angulate, humeri obscure; length (mid- The type series was taken from a shaded, seeping ex- line)/width, 0.50/0.87. posure of metamorphic bedrock next to a small spring Abdominal tergites with II longer than III-VI, ter- pool containing a few taro plants, on a trickling tribu- gite VII longest (0.23, 0.17, 0.13, 0.13, 0.17, 0.35 re- tary to a swift mountain stream in the Auga River val- spectively), abdominal venter bearing short, fine, pale ley of the central Owen Stanley Range. The insects setae; ventrite VII tumid, anteriorly with paired (1+1) were captured by applying a light pyrethrin fog to the tumescences set with long semi-erect setae, with rock face and picking them up as they fell into the tiny paired carina extending posteriorly to the margins en- pool below. A few additional specimens were taken closing sternite VIII, posteriorly excavated; first geni- syntopically with Aegilipsicola auga on wet bedrock tal segment (segment VIII) directed ventrally, bearing faces on a tributary to the Hane River near Fane (fig. an asymmetrical dagger-like anteriorly directed 85). The two taxa appeared to segregate habitat on the process (fig. 5), this process laterally set with long se- rheocrene, with A. auga on open rock faces, and R. tae, curved in side view, extending anteriorly over truncata in areas with a mosaic of bedrock and small about half of ventrite VII. scattered moss clumps. Legs clothed with numerous short, appressed, pale setae, intermixed on ventral surface of fore femur with longer, erect, pale, pilose setae; all femora with Rheovelia asymmetrica sp. n. some silvery setae on basal half, best seen in light from (figs. 5, 14) behind insect; all legs unarmed, lacking setal tufts or Material examined. – Holotype, micropterous male: other modifications. Antennae thickly clothed with PAPUA NEW GUINEA, Central Prov., Owen Stanley short to long setae, segment I also bearing 3-4 long, Range, hill stream trib. to upper Mimani River, 2.7 spine-like black setae. km. NE of Dorobisoro, 580-685 m. [1900-2250 ft.], Antennal formula I : II : III : IV; 0.35: 0.20 : 0.60 9°27’22.8”S, 147°57’02.8”E, water temp. 21.5 °C., 8 : 0.67. October 2003, 09:30-12:30 hrs., CL 7262, D. A. Pol- Proportions of legs as follows: Femur, tibia, tarsal hemus (USNM). – Paratypes: PAPUA NEW GUINEA, Cen- 1, tarsal 2 of fore leg, 0.70 : 0.60 : 0.30 : 0.0; of mid- tral Prov.: 1 micropterous male, 6 micropterous fe- dle leg, 1.00 : 0.98 : 0.10 : 0.30; of hind leg, 1.40 : males, same data as holotype (USNM, BPBM, JTPC). 1.70 : 0.13 : 0.40. Paramere small, triangular, hidden; proctiger not Description modified. Size. – Micropterous male, length 2.35-2.40 mm Micropterous female: Similar to male in general (x– = 2.37, n = 2); width 0.85-0.90 mm (x– = 0.87, n = structure, but larger, coloration of abdominal tergites 2). Micropterous female, length 2.50-2.70 mm (x– = lighter. Connexiva raised to about 15°, widely separat- 2.58, n = 6); width 0.95-1.10 mm, (x– = 1.02, n = 6). ed posteriorly, evenly tapering toward tip of abdomen, Colour. – Micropterous male: ground colour black- posterolateral angles short, almost right angles, not ish brown, head, anterior pronotum, connexiva, ab- raised, not curved inward; abdominal ventrite VII dominal tergites with yellowish brown to orange with a small median posterior button-like protuber- brown markings; venter mostly yellowish to leucine. ance, protruding proctiger forming an acute angle. Head with two very broad longitudinal stripes on ver- tex, lower part yellowish brown. Pronotum dark Etymology blackish brown; laterally narrowly lighter brown; ante- The name asymmetrica refers to the asymmetrical rior lobe with a transverse yellowish stripe between dagger-like process on ventrite VIII of the male. eyes, interrupted medially; large triangular patches of silvery hairs present anterolaterally, extending behind Distribution vertex. Wing pads not visible. Abdomen brown; Southeastern New Guinea, Central Province (fig. paired foveae (1+1) between tergites I and II without 14). silvery hairs; connexiva broadly yellowish, infused with brown at sutures; tergite I weakly infused with Comparative notes yellowish; tergites II-VIII weakly marked with orange Rheovelia asymmetrica may be separated from its brown, becoming lighter distally. Legs leucine to yel- closest congener, R. petrophila, by the more asymmet- low, upper surfaces marked with brown distally on rical process on male ventrite VII, which does not ex- femora, becoming more uniform on tibiae and tarsi. tend as far anteriorly over ventrite VII (extending only Structural characters. – Micropterous male: head about halfway, versus almost reaching to the anterior of moderate length, declivant anteriorly; length 0.57, margin in R. petrophila). In addition, ventrite VII of R. width 0.67; width of eye/interocular space, asymmetrica is somewhat tumid, with paired (1+1) 0.15/0.40. Pronotum long, posterior margin evenly tumescences anteriorly, on either side of depressed

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midline, set with long semi-erect setae, and with Abdomen light to dark brown; paired foveae not visi- paired (1+1) carinae extending posteriorly to the mar- ble; connexiva broadly yellowish, infused with brown, gins enclosing ventrite VIII; in R. petrophila, ventrite with an irregular median longitudinal stripe; tergites VII lacks these modifications, but has a pair (1+1) of II-VIII extensively marked with orange brown, be- depressions laterally. coming lighter distally. Legs leucine to yellow, upper The females of these two species also differ in their surfaces marked with brown distally on femora, be- abdominal terminalia. In R. asymmetrica the connexi- coming more uniform on tibiae and tarsi. va are only slightly raised, and not sharp or produced, Structural characters. – submacropterous male: whereas in R. petrophila the connexiva are raised and head of moderate length, declivant anteriorly; length distally acute, projecting beyond tergite VIII posteri- 0.40, width 0.55; width of eye/interocular space, orly, with a tuft of dark setae extending posteriorly 0.10/0.30. Pronotum long, posterior margin broadly from each. In addition, the anal cone of R. asymmetri- angulate, humeri prominent; length (midline)/width, ca forms an acute triangle extending posteriorly, 0.60/0.70. whereas in R. petrophila it is button-like. Abdominal dorsum with only tergites with V-VII visible beyond abbreviated wings (lengths = 0.10, Biological notes 0.15, 0.25 respectively); abdominal venter bearing The type series was taken from a wet exposure of short, fine, pale setae; ventrite VII on a plane with metamorphic bedrock shaded by primary rain forest, ventrite VI, excavate behind, excavation bordered on bordering a cascade on a steeply dropping hill stream each side by a carina; first genital segment (segment draining the flanks of the Mimani River valley on the VIII) extending posteriorly, anteroventral margin southern flank of the Owen Stanley Range. The in- with a thin sclerotized transverse ridge, posteroventral sects were very localized in their distribution, being margin bearing a ventrally directed truncate triangular absent on other nearby rock faces of similar appear- protuberance, with two notches, resulting in three ance, and were not abundant. joined protuberances (fig. 10). Legs clothed with numerous short, appressed, pale setae, intermixed on ventral surface of fore femur Rheovelia fonticola sp. n. with longer, erect, pale, pilose setae; all femora with (figs. 10, 14) some silvery setae on basal half, best seen in light from Material examined. – Holotype, submacropterous behind insect; all legs unarmed, lacking setal tufts or male: PAPUA NEW GUINEA, Oro Prov., Kofure River, W. other modifications. Antennae thickly clothed with of Tufi, 0-45 m. [0-150 ft.], small trib. with waterfall, short to long setae, segment I also bearing 3-4 long, 9°04’58.2”S, 149°16’38.7”E, water temp. 28 °C., 25 spine-like black setae. January 2004, 09:30-14:00 hrs., CL 7304, D. A. & J. Antennal formula I : II : III : IV; 0.23 : 0.15 : 0.55 T. Polhemus (USNM). – Paratypes: PAPUA NEW GUINEA, : 0.50. Oro Prov.: 2 macropterous males, 2 submacropterous Proportions of legs as follows: Femur, tibia, tarsal males, 1 submacropterous female, same data as holo- 1, tarsal 2 of fore leg, 0.50 : 0.50 : 0.15 : 0.0; of mid- type, D. A. & J. T. Polhemus (USNM, JTPC). dle leg, 0.70 : 0.75 : 0.05 : 0.17; of hind leg, 0.90 : 1.10 : 0.07 : 0.30. Description Paramere small, triangular, hidden; proctiger not Size. – Submacropterous male, length 2.00-2.05 modified. mm (x– = 2.02, n = 3); width 0.50-0.65 mm (x– = 0.58, Macropterous male: Similar to submacropterous n = 3). Submacropterous female, length 2.20 mm. (n male in general structure and coloration, but with fol- = 1); width 0.70 (n = 1). lowing exceptions: body length 2.20 mm; maximum Colour. – Submacropterous male: ground colour width (across humeral angles of pronotum) 0.80 mm; dark brown, head, anterior pronotum, connexiva, ab- pronotum length/width = 0.62/0.80; wings longer, dominal tergites with yellowish brown to orange reaching past tip of abdomen, bearing 5 closed cells, brown markings; venter mostly yellowish brown. consisting of two elongate cells basally, two more elon- Head with two broad longitudinal stripes on vertex, gate cells centrally, and a single smaller cell distally; lower part yellowish brown. Pronotum dark blackish pattern of pruinose lavender markings on hemlytra brown; laterally narrowly lighter brown; anterior lobe similar to brachypter basally, but with one elongate transversely yellowish brown between eyes, not inter- spot laterally past middle, followed by 4-5 small to rupted medially; large triangular patches of silvery large irregular spots on distal ¾, and a single large ir- hairs present anterolaterally, extending behind vertex. regular spot subapically. Wings reaching onto tergite VI, blackish brown, each Submacropterous female: Similar to male in general with two pruinose lavender streaks basally, two small structure and coloration, but slightly larger. Connexiva spots medially, tips pruinose, all bright bluish grey. not raised, widely separated posteriorly, evenly taper-

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Rheovelia truncata

Rheovelia fonticola

Rheovelia asymmetrica

Fig. 14. Distribution of Rheovelia species on New Guinea. ing toward tip of abdomen, posterolateral angles trun- Material examined. – Holotype, micropterous cate, not curved inward, tips each with a small sharp male: PAPUA NEW GUINEA, Milne Bay Prov., Basilaki tuft of setae; abdominal venter unmodified, proctiger Is., Kalawai River and rocky trib., 5-145 m. [20-475 deflected ventrad, anal cone button-like, protruding. ft.], 10°36’50.6”S, 151°01’14.4”E, water temp. 26.5 °C., pH 8.27, 17 January 2004, 09:00-14:00 hrs., CL Etymology 7294, D. A. & J. T. Polhemus (USNM). – Paratypes: The name fonticola drives from ‘fontis’ L., spring, PAPUA NEW GUINEA, Milne Bay Prov.: 1 macropterous and ‘cola’, dweller, inhabitant. female, 13 micropterous males, 22 micropterous fe- males, same data as holotype (USNM, BPBM, JTPC); 16 Distribution micropterous males,19 micropterous females, Basilaki Southeastern New Guinea, Cape Nelson Peninsula Is., Guiagoila River, trib. with waterfall, 0-15 m. [0-50 (fig. 14). ft.], 10°36’26.7”S, 150°59’37.8”E, water temp. 26.5 °C., 18 January 2004, 09:00-13:30 hrs., CL 7296, D. Comparative notes A. & J. T. Polhemus (USNM). Rheovelia fonticola may be separated from the other species of the genus by its small size, the two separate Description protuberances of the male ventrite VIII, and by the Size. – Micropterous male, length 2.10-2.15 mm more extensive pruinose markings on the hemelytra. (x– = 2.12, n = 5); width 0.85-0.90 mm (x– = 0.84, n = The predominance of submacropterous and macro- 5). Micropterous female, length 2.20-2.30 mm (x– = pterous morphs is unusual among Rheovelia species. 2.24, n = 5); width 0.95-1.00 mm, (x– = 0.98, n = 5). Colour. – Micropterous male: ground colour black- Biological notes ish brown, head, anterior pronotum, connexiva, ab- The type locality was a small, cascading tributary to dominal tergites with yellowish brown to orange the Kofure River, a moderately large, rocky stream brown markings; venter mostly yellowish to leucine. cutting a deep gorge above the head of the Maclaren Head with two very broad longitudinal stripes on ver- Harbor fjord near Tufi. The type series of R. fonticola tex, posterior part except medially, lower part yellow- was taken from a wet, vertical exposure of volcanic ish brown. Pronotum dark blackish brown; laterally bedrock adjacent to a small cascade and plunge pool, narrowly lighter brown; anterior lobe with a transverse at a point just above the confluence of the tributary yellowish stripe between eyes, interrupted medially; with the main river. large triangular patches of silvery hairs present antero- laterally, extending behind vertex. Wing pads white, present as tiny buds, barely visible. Abdomen brown; Rheovelia basilaki sp. n. paired foveae (1+1) between tergites I and II with sil- (figs. 7, 15, 84) very hairs; connexiva broadly yellowish, infused with

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brown at sutures; tergite I weakly infused with yellow- ly circular white patch, outer distal cell bearing an ir- ish brown; tergites V-VIII weakly marked with orange regular, partly interrupted white streak, apex of wing brown, becoming lighter distally. Legs leucine basally, beyond distal cells with a narrow longitudinal white upper surfaces marked with brown distally on femora, streak medially. becoming more uniform on tibiae and tarsi. Macropterous male. Unknown. Structural characters. – Micropterous male: head of moderate length, declivant anteriorly; length 0.45, Etymology width 0.63; width of eye/interocular space, The name basilaki is a noun in apposition and 0.12/0.40. Pronotum long, posterior margin evenly refers to the type locality, Basilaki Island. curving, not angulate, humeri obscure; length (mid- line)/width, 0.43/0.85. Distribution Abdominal tergites with II longer than III-VI, ter- Basilaki Island (fig. 15). gite VII longest (0.20, 0.15, 0.12, 0.12, 0.15, 0.30 re- spectively); abdominal venter bearing short, fine, pale Comparative notes setae; ventrite VII tumid, anteriorly with paired (1+1) Rheovelia basilaki may be separated from its closest tumescences flanked by angled depressions, with congeners (those species in which males bear a promi- paired carina extending posteriorly to the margins en- nent anteriorly directed process on ventrite VIII) by closing sternite VIII, posteriorly and anteriorly exca- the shorter triangular shape of the process (fig. 7). The vated; first genital segment (segment VIII) directed females are unique in the genus in having the the ventrally, bearing a triangular anteriorly directed proctiger strongly deflected ventrad and withdrawn, process (fig. 7), extending anteriorly over posterior and the anal cone not visible in lateral view. margin of ventrite VII. Legs clothed with numerous short, appressed, pale Biological notes setae, intermixed on ventral surface of fore femur with The Kalawai River at the type locality was a moder- longer, erect, pale, pilose setae; all femora with some ately large, swift, clear stream descending from a rain silvery setae on basal half, best seen in light from be- forested catchement on northern Basilaki Island. The hind insect; all legs unarmed, lacking setal tufts or oth- type series of R. basilaki was taken from vertical wet er modifications. Antennae thickly clothed with short exposures of metamorphic bedrock adjacent to a large to long setae, segment I also bearing 3-4 long, spine- waterfall approximately 20 m. in height along the like black setae. midreach of the main river channel; additional speci- Antennal formula I : II : III : IV; 0.30 : 0.16 : 0.55 mens were taken from similar seeping bedrock faces : 0.63. that formed small rheocrenes along tributary stream- Proportions of legs as follows: Femur, tibia, tarsal lets above the main fall. Another series of R. basilaki 1, tarsal 2 of fore leg, 0.62 : 0.50 : 0.25 : 0.0; of mid- was taken in a similar situation on a tributary to the dle leg, 0.77 : 0.75 : 0.10 : 0.25; of hind leg, 1.10 : Guiagoila River, where a swift, rocky stream shaded 1.37 : 0.10 : 0.35. by primary rain forest cascaded over a sloping expo- Paramere small, triangular, hidden; proctiger not sure of metamorphic bedrock with flanking rheocre- modified. nes (fig. 84). In all these cases, as is typical for species Micropterous female: Similar to male in general of Rheovelia, the insects were found on open rock sur- structure, but larger, coloration of abdominal tergites faces lightly wetted by seeping flow. lighter. Connexiva raised to about 30°, widely sepa- rated posteriorly, evenly tapering toward tip of ab- Zoogeographic notes domen, posterolateral angles short, acute, very slight- The remarkable presence of an apparently endemic ly raised, and curved inward; abdominal ventrite VII species of Rheovelia on Basilaki indicates that this is- unmodified, proctiger deflected ventrad, withdrawn, land, although not particularly large or tall in compar- anal cone not visible. ison to others in the region, may be of considerable Macropterous female: Similar to submacropterous age, and may also have been isolated east of the China male in general structure and coloration, but with fol- Strait for a prolonged period of time. Although nar- lowing exceptions: body length 2.50 mm; maximum row, each of the straits in the sequential series separat- width (across humeral angles of pronotum) 1.10 mm; ing the eastern terminus of New Guinea from Sariba, pronotum length/width = 0.70/1.10; wings longer, Sideia, and Basilaki islands in turn is over 60 m. deep, reaching to tip of abdomen, bearing 4 closed cells, and each would have represented a persistant water consisting of two elongate cells basally, plus two more barrier thoughout all but the lowest sea level stands of elongate cells centrally; wings predominantly black, the Pleistocene (Voris 2000, Dickinson 2001). This outer basal cell mostly filled by an elongate pale bluish degree of isolation appears to have been sufficient to pruinose streak, inner distal cell bearing a large, rough- produce speciation in certain elements of the island’s

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150° 151° 152° 153° 154°

KIRIWINA IS. Brechivelia TROBRIAND ISLANDS tufi

° 9° WOODLARK IS. 9

GOODENOUGH IS.

FERGUSSON IS. COLLINGWOOD BAY

D'ENTRECASTEAUX ISLANDS EGUM ATOLL

NORMANBY IS. NEW GUINEA GOODENOUGH BAY SOLOMON SEA 10° Rheovelia

MILNE BAY anomala ORANGERIE BAY ENGINEER GROUP DEBOYNE IS. MISIMA IS.

LOUISIADE ARCHIPELAGO CONFLICT GROUP 11° 11° Rheovelia ROSSEL IS. basilaki

TAGULA IS. CORAL SEA

150° 151° 152° 153° 154°

Fig. 15. Distribution of Rheovelia and Brechivelia species in the far eastern New Guinea region. microveliine biota, such as Rheovelia, but not among mental sutures; abdominal tergites with a median pru- others, such as Neusterensifer, where common species inose line, thin basally, wider and more prominent are shared with the Cloudy Mountains of nearby distally; venter mostly yellowish, darker laterally and mainland New Guinea. The presence of an endemic on metasternum. Rheovelia, coupled with the island’s underlying meta- Structural characters. – Only micropterous form morphic geology, indicates that it may be part of the known, shape narrow-elongate (fig 16). Eyes globose, same old and fragmented mountain chain that appears exserted but not beyond anterolateral pronotal angles, again further to the southeast as the Louisiadae Islands. separated by more than twice the width of an eye, slightly removed from anterior pronotal margin, with moderate length ocular setae. Head declivant anterior- Brechivelia gen. n. ly, posterior margin sloping caudo-dorsally, with usu- (figs. 15-18) al three pairs of facial trichobothria plus numerous Type-species: Brechivelia tufi Polhemus & Polhemus sp. n. short setae; vertex with prominent median sulcus; gu- lar region short, plainly visible, rostral cavity closed Description posteriorly, weakly carinate. Rostrum reaching to Size. – Micropterous form, length of males 2.25- middle of mesosternum, segment I short, enclosed in 2.35 mm., females 2.50-2.70 mm; general body char- rostral cavity, segment IV about twice the length of I, acteristics and size slightly sexually dimorphic, males almost four times longer than II, segment III more more slender and slightly smaller than females. than 7 times as long as II. Antennae very long and Colour. – Ground colour blackish brown, with yel- slender, segments I to IV subequal in length, very lowish brown markings; head weakly marked with yel- long, total length subequal to body length. lowish brown, without clearly defined markings; Pronotum very slightly raised medially, with weak pronotum anteriorly with a yellowish brown trans- median carina; collar weakly formed, absent medially, verse stripe behind vertex of head, weakly interrupted evident behind eyes laterally; anterior and posterior medially; connexiva mostly brown, darker at interseg- lobes set off by a transverse row of obscure foveae,

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evanescent laterally; entire dorsum sparsely set with male proctiger (fig. 17); in addition, it is known only short decumbent setae; with large triangular patches of from the micropterous morph. The most closely relat- silvery setae behind eyes; posterior lobe with numer- ed Papuan genera, Aegilipsicola Polhemus & Polhe- ous small foveae, humeri not prominent, broadly mus and Rheovelia gen. n., have similarly long but rounded posteriorly, not modified, mostly covering quite slender claws (shared with Aegilipsovelia of the metanotum. Metanotum laterally pruinose. Thoracic New world tropics), but have the male proctiger un- venter not diagnostic, with tubercles on either side of modified. Aegilipsicola has a long gula and rostral cav- mesosternal midline on posterior margin opposing an ity clearly demarcated by a strongly raised carina. unmodified metasternum. Metasternal scent gland Rheovelia shares with Brechivelia the short gula, but opening (omphalium) visible, marked by a small tu- the male abdominal segment VIII is essentially sym- bercle; scent channels prominent, curving slightly an- metrical, and the antennal ratios of the two genera are terad to base of metacetabulae. Wing pads not visible greatly different. The very long slender antennae are in micropters. shared with Tanyvelia, but in the latter segment two is Abdomen broad, with two (1+1) glabrous foveae shorter than one, and neither of them are as long as on either side of midline between tergites I and II; three or four. without longitudinal carinae on tergites; tergites I- VII subequal in length, VIII slightly longer. Abdomi- Biological notes nal sternites set off from laterosternites by hair-free As with the previously described Rheovelia, with glabrous oval lacunae; male ventrites VII-VIII vari- which it is occasionally sympatric, the single known ously modified. species of Brechivelia is obligatorily hygropetric, oc- Legs slender, long, hind legs much longer than oth- curring on vertical seeping rheocrenes emerging from ers, hind tibia almost as long as body; anterior femur bedrock exposures adjacent to small cascades. In com- set beneath with short light setae, unmodified in bination with Rheovelia and Aegilipsicola, it forms males; anterior tibia of male with a comb of minute part of a remarkable local biota of rheophilic Microv- black setae occupying 4/5 of tibial length; middle and eliinae endemic to the New Guinea region. hind femora set ventrally with short slender setae; all tarsi long (fig. 16), claws extremely long, somewhat Zoogeographic notes thickened; both up- and down curving arolia long, As currently understood, Brechivelia is confined to slender (fig. 18). rheocrene habitats in the deep, shady canyons drain- Male ventrite II carinate, II to VII unmodified, ing the Nelson Range, an isolated set of Quaternary genital segments prominent, modified, segment VIII volcanoes less than 2 million years old (Bain et al. expanded on right side to accomodate large, back- 1972). These volcanoes appear to have formed in con- wardly curving process arising from proctiger (fig. junction with tectonic extension along the Woodlark 16); parameres absent or vestigial; proctiger with Rift, creating an island-like enclave of mountain habi- large, backwardly curving process on right side (fig. tat well removed from the much older main cordillera 17) and smaller, angulate process on left side (fig. 17). of medial New Guinea. Although the areas lying be- Female with tergite VIII on almost the same plane as tween the Nelson Range and the nearest high massifs VII, slightly sloping, rounded posteriorly; proctiger of the central mountains, such as Mt. Suckling and prominent, bent slightly ventrad, gonocoxae visible. Mt. Dayman, are imperfectly surveyed, the fact re- mains that Brechivelia has yet to be found elsewhere Etymology on New Guinea or nearby islands. It therefore seems The generic name Brechivelia is derived from Bre- possible that Brechivelia may represent a locally en- cho (Gr.), wet, steep, referring to the wet vertical rock demic genus that evolved on the Nelson Range itself habitat, and Velia, the nominate genus of the family. and has yet to spread into the remainder of the EPCT or Gender feminine. beyond.

Distribution Brechivelia tufi sp. n. New Guinea, Cape Nelson Peninsula, Tufi area (figs. 15-18) (fig. 15). Material examined. – Holotype, micropterous Comparative notes male: PAPUA NEW GUINEA, Oro Prov., Kofure River Comparative notes. – Brechivelia differs from all and small tribs. with waterfalls, W. of Tufi, 0-45 m. other microveliine genera of the region by possessing [0-150 ft.], 9°04’58.2”S, 149°16’38.7”E, water temp. very long but rather stout claws (fig. 18), coupled with 28 °C., 25 January 2004, 09:30-14:00 hrs., CL 7304, a short gula and rostral cavity demarcated by a weakly D. A. & J. T. Polhemus (USNM). – Paratypes: PAPUA raised carina, and a highly modified and asymmetrical NEW GUINEA, Oro Prov.: 9 micropterous males, 13

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posterior margin evenly curving, not angulate, humeri obscure; length (midline)/width, 0.55/0.75. Abdominal tergites with II longer than III-VI, ter- gite VII longest (0.15, 0.11, 0.11, 0.12, 0.12, 0.25 re- spectively); abdominal venter bearing short, fine, pale setae; ventrites II to VII unmodified; first genital seg- ment (segment VIII) modified, expanded on right side and slightly twisted to enclose the asymmetrical proctiger; ventrite VIII depressed on right side anteri- orly, then curving ventrad creating a sheath for arm on right side of the proctiger (fig. 16). Legs clothed with numerous short, appressed, pale setae, intermixed on ventral surface of fore femur with slightly longer, erect, pale, pilose setae; fore tibia with grasping comb over 4/5 of length; all legs un- armed, lacking setal tufts or other modifications. An- tennae very long, slender, thickly clothed with short setae, segment I also bearing several long, spine-like black setae. Antennal formula I : II : III : IV; 0.55 : 0.60 : 0.65 : 0.70. Proportions of legs as follows: Femur, tibia, tarsal Fig. 16. Brechivelia tufi, dorsal habitus. 1, tarsal 2 of fore leg, 0.90 : 0.75 : 0.25 : 0.0; of mid- dle leg, 1.07 : 1.10 : 0.10 : 0.25; of hind leg, 1.40 : 2.00 : 0.20 : 0.30. Parameres absent or vestigial; proctiger with large, micropterous females, same data as holotype, D. A. & backwardly curving process on right side (fig. 17) and J. T. Polhemus (USNM, JTPC). smaller, angulate process on left side (fig. 17). Micropterous female: Similar to male in general Description structure, but larger, broader, coloration of abdominal Size. – Micropterous male, length 2.25-2.35 mm tergites lighter, tergites I-III broadly silvery pruinose (x– = 2.30, n = 4); width 0.60-0.70 mm (x– = 0.66, n = laterally. Connexiva raised, almost vertical, separated 4). Micropterous female, length 2.50-2.75 mm (x– = posteriorly, evenly tapering toward tip of abdomen, 2.61, n = 6); width 0.80-0.90 mm, (x– = 0.86, n = 6). posterolateral angles short, almost right angles, not Colour. – Micropterous male (see generic descrip- raised, slightly curved inward, set with medially di- tion): ground colour blackish brown, head, anterior rected long black setae; abdominal ventrite VII un- pronotum, connexiva, abdominal tergites with yellow- modified; tergite VII shelf-like, extending beyond ish brown to orange brown markings; venter mostly connexival angles, protruding proctiger quadrate, anal yellowish. Head without prominent markings, lower cone angular, gonocoxae visible. part yellowish brown. Pronotum dark blackish brown; laterally narrowly lighter brown; anterior lobe with a Etymology transverse yellowish stripe between eyes, weakly inter- The name tufi, a noun in apposition, refers to the rupted medially; large triangular patches of silvery pru- type locality in the Tufi area of Cape Nelson on the inose present anterolaterally, extending behind vertex. north coast of New Guinea. Wing pads not visible. Abdomen brown; paired foveae (1+1) between tergites I and II glabrous; connexiva Distribution broadly yellowish, infused with brown at sutures; ter- Southeastern New Guinea, Cape Nelson Peninsula gite I weakly infused with yellowish; tergites I and II (fig. 15). silvery pruinose laterally, II-VII with a median pru- inose line, more pronounced distally. Legs leucine to Comparative notes yellow, upper surfaces marked with brown distally on Brechivelia tufi is easily separated from other femora, becoming more uniform on tibiae and tarsi. Papuan Microveliinae by the extremely long slender Structural characters (see generic description). – legs and antennae, the latter with all segments sube- Micropterous male: head of moderate length, decli- qual in length (fig. 16), and by the unique modifica- vant anteriorly; length 0.40, width 0.55; width of tions of the male proctiger (fig. 17). eye/interocular space, 0.13/0.30. Pronotum long,

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Biological notes Polhemus 2000a) subsequently described a second Most specimens were taken from a huge mossy species, T. bosavi, taken from the remote upper eleva- boulder wetted by spray near the bottom of a high (c. tions of Mt. Bosavi, an isolated Pliocene volcano lying 30 m) waterfall with modest flow. This locality was in south of the central ranges in southern New Guinea. a slot canyon in dark volcanic rock that did not re- Three more new species are described herein from the ceive sunlight for most of the day. Additional speci- East Papua Composite Terrane. To date, all members mens were taken from a wet, vertical volcanic bedrock of this genus have been taken from localities in Papua face adjacent to a trickling cascade on another tribu- New Guinea, and none are currently known from In- tary further upstream, also in a shaded situation. donesian western New Guinea, despite intensive col- lecting of Microveliinae in that latter area during re- Tanyvelia Polhemus & Polhemus cent years. This may indicate that the genus evolved on the EPCT, since four of the five species are confined Tanyvelia Polhemus & Polhemus 1994: 63 (type species to that tectonic unit. Tanyvelia missim Polhemus & Polhemus 1994, monoba- sic). Species of Tanyvelia are characterized by having very long, slender antennae; an elongate male Discussion. – The genus Tanyvelia was described by proctiger (Polhemus & Polhemus 1994: fig. 4); and a Polhemus & Polhemus (1994) to hold Tanyvelia mis- slender, acuminate male paramere (Polhemus & Pol- sim from the Wau area in the Owen Stanley Range of hemus 1994: fig. 3). Male genitalic structures are not eastern New Guinea; these same authors (Polhemus & particularly informative for species separation in Tanyvelia, with the shapes of the proctiger and para- mere being similar throughout all species. By contrast, the modifications of the female connexiva and male Figs. 17, 18. Brechivelia tufi, male, structural details.– 17. pregenital ventral abdomen are diagnostic. Proctiger; 18. Hind tarsus. For further discussion of structural characters in this genus see Polhemus & Polhemus (1994, 2000a).

Revised key to the species of Tanyvelia [Note: In the revised key given by D. & J. Polhe- mus (2000a:102) ventrite VIII was inadvertantly giv- en as ventrite VII.]

1. Body length less than 2.00 mm; male abdominal ventrite VIII unmodified, lacking medial carina or sulcus, or modified setal patches ...... 2 - Body length exceeding 2.20 mm; male abdomi- nal ventrite VIII bearing medial carina or sulcus, and may have modified setal patches; basal ab- dominal tergites pruinose or black ...... 3 17 2. Yellowish brown coloration on anterior margin of pronotum extending across entire width of pronotum and onto anterolateral areas; mi- cropterous form with apex of pronotum angulate (Fig. 20), extending posteriorly across center of metanotum to base of abdominal tergite II; east- ern New Guinea ...... T. minima sp. n. - Yellowish brown coloration on anterior prono- tum confined to a narrow transverse patch on central portion (Fig. 21), yellowish brown col- oration not extending onto anterolateral sections; micropterous form with apex of pronotum rounded (Fig. 21), not extending across central section of metasternum or reaching base of ab- 18 dominal tergite II; Tagula Island ...... T. tagulana sp. n.

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Polhemus 1994: fig. 13), flanked by small patch- es of semi-erect setae; male fore tibial comb 4/5 the length of tibia; female connexiva folded in- ward over tergites V-VII ...... T. bosavi Polhemus & Polhemus - Male abdominal ventrite VIII bearing a narrow longitudinal medial shining sulcus; male fore tib- ial comb ½ the length of tibia; female connexiva almost upright along tergites V-VII (fig. 19) ...... T. papuana sp. n.

Tanyvelia minima sp. n. (figs. 20, 22) Material examined. – Holotype, micropterous male: PAPUA NEW GUINEA, Milne Bay Prov., Cloudy Mountains, rocky stream 1 km. above Gadowalai vil- 19 lage, S. of Gelemalaia, 135 m. [450 ft.], water temp. 24.5 °C., 12 April 2002, 10:00-10:30 hrs., 10°28’57”S, 150°14’27”E, CL 7176, D. A. Polhemus (USNM). – Paratypes: PAPUA NEW GUINEA, Milne Bay Prov.: 2 macropterous males, 2 macropterous females, 3 micropterous males, 1 micropterous female, same data as holotype (USNM); 2 macropterous males, 2 macropterous females, 5 micropterous males, 2 mi- cropterous females, Cloudy Mountains, headwater tributary to upper Watuti River, S. of Gelemalaia, 715 m. [2350 ft.], 10°29’50”S, 150°13’58”E, water temp. 20 21 22 °C., 10 April 2002, 16:00-17:30 hrs., CL 7175, D. A. Polhemus (USNM, JTPC, BPBM); 7 micropterous Figs. 19-21. Tanyvelia species. – 19, T. papuana, micropter- males, 4 micropterous females, headwater reach of ous female, dorsal habitus, appendages omitted; 20-21, Goilayoli River above crossing on road from Watunou Pronotum of micropterous form. – 20, T. minima; 21, T. tagulana. to Huhuna, 18.5 km. ENE of Alotau, 275 m. [900 ft.], 10°18’43”S, 150°37’16”E, water temp. 24.5 °C., 6 April 2002, 10:00-13:00 hrs., CL 7161, D. A. & J. T. Polhemus (USNM); 1 micropterous male, Upalai 3. Male abdominal ventrite VIII bearing an anterior- River at Haluwia, 13.7 km. E. of Alotau on East Cape ly directed triangular medial carina set with an road, midreach and waterfall approx. 2.5 km. above erect tuft of stiff dark setae (Polhemus & Polhe- mouth, 45-75 m. [150-250 ft.], 10°19’39”S, mus 1994: fig. 14); male abdominal ventrite VII 150°34’36”E, water temp. 24.5 °C., 5 April 2002, depressed centrally to form an anteriorly directed 09:45-16:30 hrs., CL 7163, D. A. & J. T. Polhemus triangular sulcus; male abdominal ventrites III-V (USNM); 2 micropterous males, 1 micropterous female, broadly depressed centrally, forming a broad con- Basilaki Is., Kalawai River and rocky trib., 5-145 m. cavity on underside of abdomen; basal abdominal [20-475 ft.], 10°36’50.6”S, 151°01’14.4”E, water tergites of both males and females pruinose ...... temp. 26.5 °C., pH 8.27, 17 January 2004, 09:00- ...... T. missim Polhemus & Polhemus 14:00 hrs., CL 7294, D. A. & J. T. Polhemus (USNM). - Male abdominal ventrite VIII lacking an anterior- ly directed triangular medial carina; male abdom- Description inal ventrite VII lacking a triangular sulcus medi- Size. – Micropterous male, length 1.95-2.08 mm (x– ally, but bearing a weakly depressed longitudinal = 2.01, n = 5); width 0.60-0.70 mm (x– = 0.65, n = 5). median line; male abdominal ventrites III-V not Micropterous female, length 2.15-2.25 mm (x– = 2.20, depressed medially; basal abdominal tergites of n = 2); width 0.80-0.85 mm, (x– = 0.82, n = 2). both males and females black, lacking pruinosity Macropterous male, length 2.20-2.22 mm (x– = 2.21, n ...... 4 = 2); width 0.85-0.90 mm, (x– = 0.87, n = 2). 4. Male abdominal ventrite VIII bearing a narrow Macropterous female, length 2.35-2.40 mm (x– = 2.37, longitudinal medial impression (Polhemus & n = 2); width 1.00-1.05 mm, (x– = 1.02, n = 2).

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Colour. – Micropterous male: ground colour dark Wings extending to tip of abdomen, black, bearing 4 brown to black, venter and connexiva tinged with or- closed cells in basal half of wing, basal cell lying in- ange brown. Head dark dorsally, orange brown ven- ward from subcosta with basal half white overlain by trally; rostrum yellowish brown on basal three seg- faint lavender tint, 3 remaining cells all with small ments, piceous distally; eyes dark red to black. white spots internally, distal half of wing tinged with Pronotum with anterior lobe entirely pale yellowish golden pruinosity on distal half except for veins black, brown across entire width, this coloration extending distal membrane lacking pale markings; erect pale se- laterally onto extreme upper section of propleura; disc tae present basally on subcostal, cubital, medial, radial and collar dark brown. Metanotum exposed laterally, and anal veins. black, overlain with faint silvery pruinosity. Micropter- Macropterous female: Similar to macropterous ous wing pads black, apices sometimes embrowned. male in general structure and coloration, with follow- Abdomen dark brown, orange brown medioventrally ing exceptions: Pronotum with short erect setae on and narrowly along connexival margins, tergites uni- posterior lobe near apex (most evident when viewed formly dark brown to black and lacking pruinosity in laterally). Connexiva erect, folded slightly inward both males and females; a narrow longitudinal brown over distal portions of wings. line present on tergites III-VI. Antennae with segments I and II yellowish brown, segments III and IV medium Etymology brown. Legs pale yellowish basally and on tarsi, all tib- The name “minima” refers to the size of this iae and distal tips of femora embrowned. species, the smallest known in the genus. Structural characters. – Micropterous male: head of moderate length, declivant anteriorly; length 0.35, Distribution width 0.51; width of eye/interocular space, 0.15/0.28. Far eastern New Guinea and adjacent Basilaki Is- Pronotum length/width, 0.55/0.70; posterior lobe land (fig. 22). with apex angulate (fig. 20), covering central section of metanotum, humeri moderately produced; entire Comparative notes pronotum set with very short pale recumbent setae. Tanyvelia minima sp. n. and the superficially simi- Abdominal tergites not shining, thickly set with short lar T. tagulana sp. n. (see below) are separable from recumbent pale setae, plus a few longer setae at pos- the previously described Tanyvelia missim (Polhemus terolateral angles of connexiva, connexiva angled & Polhemus 1994) and Tanyvelia bosavi (Polhemus & broadly outward; tergites II-VII progressively greater Polhemus 2000a) plus Tanyvelia papuana sp. n. (de- in length (0.10, 0.10, 0.10, 0.10, 0.12, 0.18 respec- scribed subsequently herein) by their small size (body tively). Abdominal venter set with short appressed length less than 2.0 mm), the unmodified abdominal pale setae; ventrites III-VI not depressed, bearing a ventrites VI and VII in males, the absence of a medial narrow, glabrous, longitudinal median line; ventrite depression on male abdominal ventrites III-V, and the VII lacking longitudinal medial depression or modi- orange brown abdominal venter in both sexes. fied setal tufts. Legs, antennae thickly clothed with Both T. missim and T. bosavi exceed 2.25 mm in short to moderate length semi-erect pale setae, with a body length, T. papuana is 2.20 mm in body length, few scattered longer setae on antennal segment I and and all have modifications to abdominal ventrite VII; posterior margins of middle femora. Legs unarmed, in the former it bears an anteriorly directed triangular tibial comb on fore femur extending for about ½ the medial carina set with an erect tuft of stiff dark setae, length of the femur. while the latter two bear a narrow longitudinal sulcus Antennal formula, segments I : II : III : IV; 0.35 : or a medial impression flanked by small patches of 0.27 : 0.50 : 0.57. semi-erect setae. Abdominal ventrites III-VI are also Proportions of legs as follows: Femur, tibia, tarsal broadly depressed medially in T. missim, in contrast 1, tarsal 2 of fore leg, 0.58 : 0.55 : 0.16 : 0.0; of mid- to the unmodified state seen in T. minima or T. dle leg, 0.70 : 0.70 : 0.10 : 0.18; of hind leg, 0.75 : bosavi. The coloration of the basal abdominal tergites 1.00 : 0.13 : 0.20. is also diagnostic, being orange brown in T. minima, Paramere slender and acuminate, very similar in pruinose in T. missim, and uniformly black in T. shape to that previously illustrated for T. missim (Pol- bosavi and T. papuana. hemus & Polhemus 1994: fig. 3). Tanyvelia minima may in turn be separated from Micropterous female: Similar to micropterous T. tagulana by its more extensive yellowish brown male in general structure and coloration. coloration on the anterior pronotum, and by the an- Macropterous male: Similar to micropterous male gulate posterior pronotal margin in micropterous in general structure and coloration, with following ex- forms (fig. 20), which transgresses the central portion ceptions: Pronotum enlarged, tumid centrally, apex of the metanotum and touches the base of abdominal broadly angulate, humeri prominent, sub-quadrate. tergite II (the first abdominal tergite visible).

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The currently known species of Tanyvelia form two with apex broadly rounded (fig. 21), not covering cen- obvious groups, with the three larger species that ex- tral section of metanotum, humeri not produced or an- hibit abdominal modifications occurring in south cen- gulate; entire pronotum set with very short pale re- tral and central New Guinea, while the two smaller cumbent setae. Metanotum with lateral sections species that lack abdominal modifications occur on the slightly raised, central section slightly depressed. Ab- Papuan Peninsula and in the Louisiade Archipelago. dominal tergites not shining, thickly set with very short recumbent pale setae, connexiva angled broadly out- Biological notes ward; length of metanotum 0.11; tergites II-VII pro- The type series of T. minima was taken from a gressively greater in length (0.09, 0.09, 0.09, 0.10, clear, cascading stream descending through a bed of 0.10, 0.20 respectively). Abdominal venter set with rounded, mossy boulders, and heavily shaded by dis- short appressed pale setae; ventrites I-VI unmodified, turbed lowland rain forest. The insects were taken bearing a narrow, glabrous, longitudinal median line; from just above the water surface on the wet edges of ventrite VII slightly depressed posteromedially, lacking large boulders bordering pools below small cascades. modified setal tufts. Legs and antennae thickly clothed with short to moderate length semi-erect pale setae, with a few scattered longer setae on antennal segments Tanyvelia tagulana sp. n. I and II. Legs unarmed, tibial comb on fore femur ex- (figs. 21, 22) tending for about 0.40 the length of the femur. Material examined. – Holotype, micropterous Antennal formula, segments I : II : III : IV; 0.30 : male: PAPUA NEW GUINEA, Milne Bay Prov., Louisiade 0.25 : 0.50 : 0.58. Archipelago, Sudest [Tagula] Island, Kolukolu Creek, Proportions of legs as follows: Femur, tibia, tarsal near Araeda village, 0-105 m. [0-350 ft.], 11°26’ 1, tarsal 2 of fore leg, 0.55 : 0.52 : 0.22 : 0.0; of mid- 44.7”S, 153°25’56.8”E, water temp. 23 °C., 30 Au- dle leg, 0.65 : 0.75 : 0.10 : 0.15; of hind leg, 0.75 : gust 2002, CL 7194, D. A. & J. T. Polhemus (USNM). 0.95 : 0.12 : 0.20. – Paratypes: PAPUA NEW GUINEA, Milne Bay Prov., Paramere slender and acuminate, very similar in Louisiade Archipelago, Tagula [Sudest] Island: 5 mi- shape to that previously illustrated for T. missim (Pol- cropterous males,12 micropterous females, same data hemus & Polhemus 1994: Fig. 3). as holotype, D. A. & J. T. Polhemus (USNM, JTPC, Micropterous female: Similar to male in general BPBM). structure and coloration, with following exceptions: abdominal tergites blackish, connexival margins Description broadly orange brown, silvery pruinosity present on Size. – Micropterous male, length 1.90-2.10 mm lateral sections of abdominal tergite II, extreme poste- (x– = 2.02, n = 3); width 0.65-0.68 mm (x– = 0.66, n = rior margin of tergite V, central sections of abdominal 3). Micropterous female, length 2.05-2.15 mm (x– = tergites VI and VII; connexival margins outwardly re- 2.12, n = 3); width 0.80-0.85 mm, (x– = 0.82, n = 3). flexed adjacent to abdominal tergites II-VI, folded in- Colour. – Micropterous male: ground colour dark ward over lateral portions of tergites VII and VIII, brown, venter and connexiva tinged with orange connexival margins on infolded sections bearing thick brown. Head dark dorsally, orange brown ventrally; fringes of moderately long, stiff black setae; lateral por- rostrum pale yellowish on basal three segments, tions of abdominal tergites II and III and infolded piceous distally; eyes dark red to black. Pronotum portions of connexiva reddish brown. with anterior lobe bearing a transverse yellowish Macropterous form: Unknown. brown patch centrally, this yellowish brown col- oration not extending laterally to margins of propleu- Etymology ra; disc and collar dark brown. Metanotum exposed The name ‘tagulana’ refers to the Tagula Island across entire width, dark brown laterally, orange type locality. brown centrally. Micropterous wing pads dark brown. Abdomen dark brown, orange brown medioventrally Distribution and faintly along connexival margins, tergites dark Louisiade Archipelago (fig.22). brown, tergite II with faint pruinosity laterally. Anten- nae with segments I and II yellowish brown, segments Comparative notes III and IV medium brown. Legs pale yellowish brown, Tanyvelia tagulana is a small species that is similar apices of all femora medium brown. in many respects to T. minima from far eastern New Structural characters. – Micropterous male: head of Guinea, but may be separated from that species by the moderate length, declivant anteriorly; length 0.35, less extensive yellowish brown coloration on the ante- width 0.55; width of eye/interocular space, 0.12/0.30. rior pronotum, which is confined to single transverse Pronotum length/width, 0.45/0.70; posterior lobe spot centrally (fig. 21), and the rounded posterior

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150° 151° 152° 153° 154°

KIRIWINA IS. TROBRIAND ISLANDS

° 9° WOODLARK IS. 9

GOODENOUGH IS.

FERGUSSON IS. COLLINGWOOD BAY

D'ENTRECASTEAUX ISLANDS EGUM ATOLL

NORMANBY IS. NEW GUINEA GOODENOUGH BAY SOLOMON SEA 10°

MILNE BAY ORANGERIE BAY Tanyvelia ENGINEER GROUP DEBOYNE IS. MISIMA IS. tagulana LOUISIADE ARCHIPELAGO CONFLICT GROUP 11° 11° Tanyvelia ROSSEL IS. minima

TAGULA IS. CORAL SEA

150° 151° 152° 153° 154°

Fig. 22. Distribution of Tanyvelia species in the far eastern New Guinea region.

pronotal margin in micropterous forms (fig. 21), males, same data as holotype (USNM, JTPC); 8 micro- which does not transgress the central portion of the pterous males, 8 micropterous females, Owen Stanley metanotum. Range, trib. to upper Mimani River, 2.0 km. NE of Dorobisoro, 565 m. [1850 ft.], 9°27’20.8”S, 147° Biological notes 56’25.5”E, water temp. 22.5 °C., 7 October 2003, The type series was taken from a small, shallow, 13:40-14:20 hrs., CL 7261, D. A. Polhemus (USNM, heavily shaded tributary to Kolukolu Creek on north- BPBM); 1 micropterous female, Owen Stanley Range, western Tagula Island. Along this stream, the insects trib. to upper Mimani River, 2.9 km. NE of Doro- were found in only one reach where a low, damp sill bisoro, 640-670 m. [2100-2200 ft.], 9°27’ 18.7”S, of bedrock bordered a clear, shallow pool. The insects 147°56’52.9”E, water temp. 22.5 °C., 8 October were dislodged by splashing this bedrock exposure, 2003, 12:45-13:30 hrs., CL 7263, D. A. Polhemus and did not appear to skate on the pool itself. (USNM).

Tanyvelia papuana sp. n. Description (figs. 19, 23) Size. – Micropterous male, length 2.25-2.30 mm (x– = 2.25, n = 5); width 0.70-0.75 mm (x– = 0.71, n = Material examined. – Holotype, micropterous 5). Micropterous female, length 2.45-2.70 mm (x– = male: PAPUA NEW GUINEA, Central Prov., Owen Stan- 2.54, n = 9); width 0.80-0.95 mm, (x– = 0.86, n = 9). ley Range, trib. to upper Mimani River, 2.7 km. NE Colour. – Micropterous male: ground colour dark of Dorobisoro, 580-685 m. [1900-2250 ft.], brown to black, venter, abdominal tergites and con- 9°27’22.8”S, 147°57’02.8”E, water temp. 21.5 °C., 8 nexiva tinged with orange brown. Head dark dorsally, October 2003, 09:30-12:30 hrs., CL 7262, D. A. Pol- orange brown ventrally; rostrum yellowish to leucine hemus (USNM). – Paratypes: PAPUA NEW GUINEA, Cen- on basal three segments, piceous distally; eyes dark red tral Prov.: 5 micropterous males, 5 micropterous fe- to black. Pronotum with anterior lobe entirely pale

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Tanyvelia missim

Tanyvelia bosavi Tanyvelia papuana

Fig. 23. Distribution of Tanyvelia species on New Guinea. yellowish orange across entire width, this coloration Legs unarmed, tibial comb on fore femur extending extending laterally onto extreme upper section of pro- for about ½ the length of the femur. pleura; disc and collar dark brown. Metanotum ex- Antennal formula, segments I : II : III : IV; 0.50 : posed laterally, black, overlain with faint silvery pru- 0.35 : 0.57 : 0.75. inosity. Micropterous wing pads not visible. Abdomen Proportions of legs as follows: Femur, tibia, tarsal dark brown, orange brown medioventrally and nar- 1, tarsal 2 of fore leg, 0.70 : 0.65 : 0.25 : 0.0; of mid- rowly along connexival margins, tergites uniformly dle leg, 0.90 : 0.95 : 0.12 : 0.25; of hind leg, 1.05 : dark brown to black and lacking pruinosity in both 1.20 : 0.20 : 0.30. males and females; a narrow longitudinal orange Paramere slender and acuminate, very similar in brown line present on tergites V-VII. Antennae with shape to that previously illustrated for T. missim (Pol- segments I and II yellowish brown, segments III and hemus & Polhemus 1994: fig. 3). IV medium brown. Legs pale yellowish basally and on Micropterous female: Similar to micropterous tarsi, all tibiae and distal tips of femora embrowned. male in general structure and coloration, with follow- Structural characters. – Micropterous male: head of ing exceptions: pronotal humeri evident, connexiva moderate length, declivant anteriorly; length 0.47, almost vertical basally, becoming weakly reflexed dis- width 0.60; width of eye/interocular space, 0.15/0.30. tally (fig. 19). Pronotum length/width, 0.55/0.72; posterior lobe Macropterous male and female: Unknown. with apex broadly rounded, covering central section of metanotum, humeri slightly produced; entire prono- Etymology tum set with very short pale recumbent setae. Abdom- The name papuana refers to the Papuan Peninsula, inal tergites not shining, thickly set with short recum- to which this species is endemic. bent pale setae, plus a few longer setae at posterolateral angles of connexiva, connexiva angled broadly out- Distribution ward; abdominal tergites II-VI subequal in length, ter- Owen Stanley Range of southeastern New Guinea gite VII longer (0.18, 0.13, 0.13, 0.15, 0.15, 0.25 re- (fig. 23). spectively). Abdominal venter set with short appressed pale setae; ventrites III-VI not depressed, bearing a Comparative notes narrow, glabrous, longitudinal median line; ventrite Tanyvelia papuana sp. n. is separable from the pre- VII with a small V-shaped excavation posteriorly, and viously described Tanyvelia missim (Polhemus & Pol- a small V-shaped notch. Legs, antennae thickly hemus 1994) and Tanyvelia bosavi (Polhemus & Pol- clothed with short to moderate length semi-erect pale hemus 2000a) in having the male ventrite VII with a setae, with a few scattered longer setae on antennal sharp triangular notch in the posterior margin, and segment I and posterior margins of middle femora. ventrite VIII with a narrow glabrous median sulcus. In

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addition, T. papuana may be separated from the close- ther gently angulate (fig. 26) or rounded (fig. 24), ly related T. bosavi by the length of the male fore tib- if angulate then not folded inward over tips of ial comb (½ as long as the tibia in T. papuana, 4/5 as wings ...... 3 long in T. bosavi), and the almost upright female con- 3. Male fore femur unmodified, lacking a compact nexiva (fig. 19), as compared to state in T. bosavi in tuft of dark setae ventrally (Polhemus & Polhe- which the connexiva which are folded over and ap- mus 2000a: fig. 17); legs mostly brown above; fe- pressed onto tergites V-VII. Tanyvelia tagulana and T. male connexiva with posterolateral angles round- minima differ from T. papuana by their small sizes ed, folded inward over lateral portions of (body length less than 2.0 mm), and the unmodified abdominal tergites VI and VII (Polhemus & Pol- abdominal ventrites VII and VIII in males. See notes hemus 2000a: fig. 17) ...... under T. minima for further discussion...... A. robinae Polhemus & Polhemus - Male fore femur with small, compact tuft of dark Biological notes setae ventrally on distal third (Polhemus & Pol- The type locality was a clear, rocky, heavily shaded hemus 2000a: fig. 16); legs dark yellowish above; creek in primary premontane rain forest, descending female connexiva with posterolateral angles gen- steeply through a bed of mixed rock types including tly angulate, upright, not (or rarely) folded in- volcanics, quartz and greenschist. The insects were ward over lateral portions of abdominal tergites found along the margins of small pools set amid cob- VI and VII ...... 4 bles and boulders. 4. Venter of first genital segment (ventrite VIII) with two prominent tubercles along lateral mar- gins of depression, plus a smaller tubercle posteri- Aegilipsicola Polhemus & Polhemus orly (fig. 28); raised posterior part of female con- Aegilipsicola Polhemus & Polhemus 1994: 60 (type species nexiva smoothly curving dorsally, angulate Aegilipsicola rapida Polhemus & Polhemus 1994, distally (fig. 26) ...... monobasic)...... A. rapida Polhemus & Polhemus Discussion. – For diagnostic structural characters - Venter of first genital segment (ventrite VIII) of this genus see discussion in Polhemus & Polhemus with three small tubercles posteriorly, the middle (1994). tubercle largest (fig. 29); raised posterior part of female connexiva with a small dorsally directed angle at distal ¾, smoothly rounded distally (fig. Revised key to the species of Aegilipsicola 24) ...... A. auga sp. n. occurring on New Guinea 1. Venter of male first genital segment deeply exca- Aegilipsicola peninsularis sp. n. vate basally, the posterior margin of this depres- (figs. 25, 27, 30) sion bordered by 2-3 small sclerotized tubercles (figs. 27-29); posterolateral apices of female con- Material examined. – Holotype, macropterous nexiva variable, may be produced and sharply an- male: PAPUA NEW GUINEA, Milne Bay Prov., Upalai gulate (fig. 25), rounded or gently angulate (figs. River at Haluwia, 13.7 km. E. of Alotau on East Cape 24, 26), or unmodified ...... 2 road, midreach and waterfall approx. 2.5 km. above - Venter of male first genital segment deeply exca- mouth, 45-75 m. [150-250 ft.], 10°19’39”S, 150°34’ vate basally, the posterior margin of this depress- 36”E, water temp. 24.5 °C., 5 April 2002, 09:45- sion lacking sclerotized tubercles, but bounded 16:30 hrs., CL 7163, D. A. & J. T. Polhemus (USNM). by a narrow, upraised, lip-like carina (Polhemus – Paratypes: PAPUA NEW GUINEA, Milne Bay Prov.: 25 & Polhemus 2000a: fig. 19); posterolateral apices macropterous males, 30 macropterous females, same of female connexiva produced and sharply angu- data as holotype, D. A. & J. T. Polhemus (USNM). late, folded inward over wing tips (Polhemus & Polhemus 2000a: fig. 21) ...... Description ...... A. iriana Polhemus & Polhemus Size. – Micropterous male, length 2.90 (x– = 2.90, n 2. Male fore femur with a broad, dark tubercle near = 5); width 1.00-1.02 mm (x– = 1.00, n = 5). Mi- middle of ventral margin; posterolateral angles of cropterous female, length 3.15-3.20 mm (x– = 3.17, n female connexiva produced and strongly angulate = 5); width 1.05-1.10 mm, (x– = 1.09, n = 5). (fig. 25), folded inward over tips of wings ...... Colour. – Macropterous male: ground colour black ...... A. peninsularis sp. n. dorsally, marked with yellowish brown ventrally. - Male fore femur with either a small, compact tuft Head black, ventral surface yellowish brown, eyes red, of dark setae ventrally on distal third, or unmod- antennae dark brown; rostrum golden yellow, termi- ified; female connexiva not sharply angulate, ei- nal joint piceous. Pronotum with anterior lobe nar-

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24

25

26

27 28 29

Figs. 24-29. Aegilipsicola species, structural details. – 24-26, Lateral view of female terminal abdomen: 24, A. auga, with de- tail of posterolateral angle of connexivum; 25, A. peninsularis; 26, A. rapida Polhemus & Polhemus, with detail of postero- lateral angle of connexivum; 27-29, Ventral view of male terminal abdomen: 27, A. peninsularis; 28, A. rapida Polhemus & Polhemus, with detail of process on ventrite VIII; 29, A. auga, with detail of process on ventrite VIII.

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rowly, transversely orange brown on either side of small detached streak; distal one third of the wing midline, this coloration extending laterally to inner bearing 2-3 elongate lavender streaks. Connexiva with eye margins; lateral portions black but thickly covered posterolateral apices sharply produced and angulate with silvery pubescence; posterior lobe black. Tho- (fig. 25), folded slightly inward over distal portion of racic and abdominal venter greyish brown, acetabulae wings but not touching. and central portion of abdomen yellowish brown. Legs predominantly dark brown above, with all coxae, Etymology trochanters, basal femora and ventral surfaces yellow- The name ‘peninsularis’ refers to the slender penin- ish brown. Hemelytra black, each forewing bearing 2 sula formed by the eastern terminus of the Owen elongate bluish-grey pruinose streaks confined within Stanley Range along the northern side of Milne Bay, cells at basal angles, plus 1-4 additional small pruinose the area from which the type series was collected. streaks on distal third. Structural characters. – Macropterous male: head Distribution of moderate length, declivant anteriorly; length 0.35; Eastern New Guinea (fig. 30). width of eye/interocular space, 0.13/0.33. Pronotum long, posterior lobe broadly tumescent, apex broadly Comparative notes angulate, humeri evident but not raised; Easily separated from previously described species length/width, 0.93/1.02. Hemelytra long, extending in the genus by the tumescence on the male fore fe- slightly beyond tip of abdomen. mur coupled with the sharply angulate posterolateral Head, posterior lobe of pronotum, and veins on angles of the female connexiva (fig. 25) and the sculp- basal half of hemelytra bearing numerous short, curl- turing of the male pregenital abdomen (fig. 27). ing golden setae; lateral sections of anterior lobe of pronotum thickly covered with long, recumbent, sil- Biological notes very setae; legs and antennae thickly covered with re- The Upalai River at the type locality was a clear, cumbent pale setae, intermixed with slightly longer, rocky stream descending steeply through a bed of semi-erect pale setae on tibiae and antennal segments; stream-rounded rocks and boulders in a narrow, shady abdominal venter set with very short appressed fine se- valley retaining an original cover of undisturbed pri- tae. Abdominal ventrites III-VI with a glabrous longi- marly lowland to premontane rain forest. The type se- tudinal midline stripe; ventrite VI with a roughly ries of A. peninsularis was taken from wet bedrock ex- trapezoidal raised area centrally, with its broad base di- posures surrounding a deep plunge pool at the base of rected anterad, flanked by a pair (1+1) of shallow, a 20 m high waterfall along the river’s upper midreach. rounded depressions; ventrite VII with a pair (1+1) of As with all Aegilipsicola species, which are obligatorily strong depressions laterally on basal half to either side hygropetric, the insects ran swiftly over the vertical to of midline, these depressions contiguous with those sloping wet bedrock faces bordering the water, but did on ventrite VI, flanking a broad longitudinal medial not venture onto the surface of the plunge pool itself. carina centrally; first genital segment (ventrite VIII) deeply excavated anteromedially, the posterior margin Aegilipsicola auga sp. n. of this concavity constricted to create a smaller sec- (figs. 24, 29. 30, 85) ondary depression, the point of constriction marked by a pair (1+1) of small angulate processes with dark Material examined. – Holotype, macropterous male: apices (fig. 27). Fore femora with a broad, dark tumes- PAPUA NEW GUINEA, Central Prov., Owen Stanley cence on ventral margin just basal to midpoint. Range, small cascade and seeps on trib. to Hane River, Antennal formula I : II : III : IV; 0.50 : 0.30 : 0.65 1.3 km. SSE of Fane, 1280 m. [4200 ft.], 8°33’49.2”S, : 0.60. 147°05’14.0”E, water temp. 20 °C., 4 October 2003, Proportions of legs as follows: Femur, tibia, tarsal 08:30-09:30 hrs., CL 7255, D. A. Polhemus (USNM). – 1, tarsal 2 of fore leg, 0.85 : 0.67 : 0.33 : 0.0; of mid- Paratypes: PAPUA NEW GUINEA, Central Prov.: 66 dle leg, 1.08 : 1.10 : 0.10 : 0.40; of hind leg, 1.60 : macropterous males, 51 macropterous females, same 2.05 : 0.10 : 0.47. data as holotype (USNM, BPBM, JTPC); 2 macropterous Paramere small and thumb-shaped, very similar in males, 5 macropterous females, Owen Stanley Range, shape to that previously illustrated for A. rapida (Pol- seeps along upper Mimani River, 1.70 km. NE of hemus & Polhemus 1994: Fig. 8). Dorobisoro, 535 m. [1750 ft.], 9°27’25.1”S, 147°56’ Macropterous female: Similar to male in general 15.2”E, water temp. 23.5 °C., 7 October 2003, 13:00- structure and coloration, but somewhat larger; fore fe- 15:00 hrs., CL 7260, D. A. Polhemus (USNM). mur and abdominal venter unmodified. Forewing with the outer of the two large bluish basal streaks in- Description terrupted near posterior terminus to produce a third Size. – Macropterous male, length 2.60-2.75 mm

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Aegilipsicola peninsularis

Aegilipsicola auga

Fig. 30. Distribution of Aegilipsicola species on New Guinea.

(x– = 2.67, n = 7); width 0.80-0.90 mm (x– = 0.85, n = trally, the posterior margin of this concavity bounded 7). Macropterous female, length 2.95-3.10 mm (x– = by a thin, raised, plate-like trilobed process (fig. 29). 3.03, n = 6); width 0.90-1.05 mm, (x– = 0.98, n = 6). Legs and antennae thickly clothed with short incon- Colour. – Macropterous male: ground colour black, spicuous setae, without longer setae. Middle and pos- tinged with yellowish brown ventrally on prothorax terior legs unarmed, fore femur with a raised setal tuft. and abdomen. Head black, set with appressed golden Antennal formula I : II : III : IV; 0.35 : 0.25 : 0.65 setae; posterior margin and two (1+1) longitudinal re- : 0.60. gions lying between eyes and medial furrow brown. Proportions of legs as follows: Femur, tibia, tarsal Pronotum with anterior lobe narrowly, transversely 1, tarsal 2 of fore leg, 0.62 : 0.55 : 0.25 : 0.0; of mid- orange brown on either side of midline, this coloration dle leg, 0.95 : 0.90 : 0.10 : 0.30; of hind leg, 1.20 : extending laterally to inner eye margins; lateral por- 1.50 : 0.12 : 0.37. tions black but thickly covered with silvery pubes- Paramere small and thumb-shaped, very similar in cence; posterior lobe black, set with appressed golden shape to that previously illustrated for A. rapida (Pol- setae. Abdomen greyish black ventrally, central por- hemus & Polhemus 1994: fig. 8). tion broadly brown, acetabulae and first genital seg- Macropterous female: Similar to male in general ment brownish yellow. Antennae brown, first segment structure and coloration, but somewhat larger. lighter. Legs brown above, brownish yellow below. Forewing set with scattered long dark setae, bearing Hemelytra black, bearing four (2+2) elongate bluish- five small pruinose spots on distal third. Connexiva grey pruinose streaks at basal angles, confined within usually vertical, strongly raised and plate-like along the basal cells, plus four additional small pruinose tergite VII, dorsally with a small sclerotized triangular spots on distal third of each. projection, distally rounded (fig. 24), extending onto Structural characters. – Macropterous male: head tergite VIII; rarely folded inward and touching over of moderate length, declivant anteriorly; length 0.35, distal portion of wings, tergite VIII plate-like, extend- width 0.60; width of eye/interocular space, ing over tergite IX. 0.12/0.30. Pronotum long, humeri evident but not raised; length/width, 0.80/0.90. Hemelytra long, ex- Etymology tending beyond tip of abdomen. The name ‘auga’ refers to the type locality, a tribu- Abdominal tergites covered by wings; abdominal tary to the Auga River near Fane, in the Owen Stan- venter set with very short appressed fine setae; ventrite ley Range. The spelling ‘Auga’ is that adopted by the V and VI slightly flattened, sloping dorsad; posterior local people of the region, although the river appears half of ventrite VII broadly depressed medially; first as the ‘Aura’ on many maps. genital segment (ventrite VIII) deeply excavated ven-

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Distribution Revised key to the species of Tarsovelia Owen Stanley Range of southeastern New Guinea (fig. 30). Discussion. – This revised key builds upon the ver- sion presented in Polhemus & Polhemus (2000a), and Comparative notes is most effectively used in combination with the illus- Superficially similar to Aegilipsicola rapida (Polhe- trations of the male fore femora and parameres pro- mus & Polhemus 1994), which has a similar armature vided in Polhemus & Polhemus (1994, 2000a and of the male foreleg (Polhemus & Polhemus 2000a: fig. 2000b). As noted in these previous works, females of 16), but separable on the basis of the ventral male ab- Tarsovelia are often confusingly similar, and for the dominal sculpturing (compare figs. 28, 29), and the most part lack good specific characters. For this rea- structure and details of the female connexival apices son, the key below treats males only; females are best (compare figs. 24, 26). In A. rapida the ventral excava- determined by association with sympatrically collected tion of the male first genital segment is flanked poste- males. riorly by several tubercles laterally plus one on the pos- terior margin (fig. 28), while in A. auga there are three Males tubercles grouped together on the posterior margin 1. Fore femur with either a patch of short, stiff, pale (fig. 29). On male abdominal ventrite VII, the anteri- hairs or a patch of short, dense, dark hairs on ven- or half is unmodified in A. auga, while in A. rapida tral surface ...... 2 there are paired depressions anteriorly. The female - Fore femur lacking patch of short, stiff hairs or connexival apices of both A. auga and A. rapida are short, dense, dark hairs on ventral surface ...... 6 plate-like, usually lying widely separated and in a verti- 2. Fore femur with a patch of short, dense, dark cal orientation; A. auga has a small triangular dorsal hairs near central portion of ventral margin; Mt. projection at distal ¾ of the plate-like raised portion Bosavi area of southeastern New Guinea ...... which is smoothly rounded posteriorly (fig. 24), ...... T. bosavi Polhemus & Polhemus whereas A. rapida has the raised portion smoothly - Fore femur with a patch of pale, short, stiff hairs curving dorsally but angulate posteriorly (fig. 26). on ventral surface ...... 3 3. Fore femur slender basally, slightly expanded dis- Biological notes tally, bearing a patch of short, stiff, pale hairs in The type series was taken from a vertical face of wet the central section of the ventral face ...... 4 metamorphic bedrock adjacent to a cascading tribu- - Fore femur slightly expanded basally, slender and tary of the Hane River (a tributary to the Auga) at a gently arcuate distally, bearing a patch of short, point where this tributary crossed a contour trail run- stiff, pale hairs basally on ventral face; abdominal ning along the side of the Hane River gorge (fig. 85). venter orange brown; central ranges of western The surrounding slopes were covered with disturbed New Guinea ...... T. dani Polhemus & Polhemus upland rain forest, and due to their aspect the wet 4. Fore femur with patch of short, stiff, pale hairs bedrock faces on which A. auga occurred remained in extending inward from the middle of the ventral shade for much of the day. Individuals of A. auga were face to the beginning of the narrowed basal sec- found abundantly in this habitat, but were not present tion; Vogelkop Peninsula ...... on the wettest parts of the rheocrene, being absent in ...... T. arfak Polhemus & Polhemus areas with visible laminar flow. - Fore femur with patch of short, stiff, pale hairs Another short series of this species was taken fur- extending distally from middle of ventral face to- ther east in the Owen Stanley Range in the riparian ward tip of femur ...... 5 zone of the Mimani River, which drains the southern 5. Hair patch on ventral face of fore femur with flanks of the range near Mt. Obree. Here the insects short setae, these setae not becoming longer dis- were again found on a riverside exposure of wet bed- tally; abdominal venter reddish to yellowish rock wetted by seepage, and partially shaded by over- brown; south central highlands of Papua New hanging vegetation. Guinea ...... T. kikori Polhemus & Polhemus - Hair patch on ventral face of fore femur with se- tae becoming longer distally; abdominal venter Tarsovelia Polhemus & Polhemus black; northern central ranges of western New Tarsovelia Polhemus & Polhemus 1994: 63 (type species Guinea ...... T. ziwa Polhemus & Polhemus Tarsovelia alta Polhemus & Polhemus 1994, by original 6. Fore femur constricted to greater or lesser extent designation). when viewed laterally, due to concavity of ventral Discussion. – For diagnostic structural characters of surface; ventral surface of body dark brown to this genus see discussion in Polhemus & Polhemus black ...... 7 (1994). - Fore femur not constricted basally when viewed

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relatively even width throughout, apex slightly expanded and bulb-like; south central mountains of western New Guinea ...... T. reclusa Polhemus & Polhemus - Anterior lobe of pronotum in macropterous forms entirely orange brown, this coloration ex- tending laterally onto propleurae; fore femur lacking small, carinate ridge on central section of ventral face; paramere tapering throughout its 31 length, apex very slightly expanded but not bulb- like; central mountains of Papua New Guinea ...... T. alta Polhemus & Polhemus

Tarsovelia louisiadensis sp. n. (figs. 31-34) Material examined. – Holotype, macropterous male: PAPUA NEW GUINEA, Milne Bay Prov., Louisiade Archipelago, Sudest [Tagula] Island, Kolukolu Creek, near Araeda village, 0-105 m. [0-350 ft.], 11°26’44.7”S, 153°25’56.8”E, water temp. 23 °C., 30 August 2002, CL 7194, D. A. & J. T. Polhemus (USNM). – Paratypes: PAPUA NEW GUINEA, Milne Bay Prov., Louisiade Archipelago: 4 macropterous males, 11 macropterous females, Tagula [Sudest] Island, same data as holotype, D. A. & J. T. Polhemus (USNM, BPBM); 11 macropterous males, 8 macropterous fe- males, Misima Island, north coast, Guwana Creek, near Nulia village, 15-30 m. [50-100 ft.], 32 33 10°39’22.8”S, 152°41’16.5”E, water temp. 22 °C., 3 September 2002, 10:00-12:00 hrs., CL 7201, D. A. & J. T. Polhemus (USNM); 65 macropterous males, 46 Figs. 31-33. Tarsovelia louisadensis, male, structural details. macropterous females, Misima Island, south coast, – 31, Foreleg; 32, Proctiger. 33, Paramere. Keyana Creek, above Bwagabwaga village, 305-335 m. [1000-1100 ft.], 10°40’33.8”S, 152°40’44.9”E, water temp. 22.5 °C., 4 September 2002, 10:00-12:00 laterally, lacking a basal concavity on ventral sur- hrs., CL 7206, D. A. & J. T. Polhemus (USNM, JTPC). face; ventral surface of body orange brown ...... 8 7. Ventral concavity on fore femur long and shal- Description low, extending along basal half of femur (fig. 31); Size (Tagula Island population). – Macropterous fore femur when viewed laterally attaining its male, length 2.50-2.70 mm (x– = 2.50, n = 4); width greatest thickness on distal half (fig. 31); middle 1.00-1.02 mm (x– = 1.00, n = 4). Macropterous fe- tibia twice as long as basal tarsal segment; Louisi- male, length 2.85-3.05 mm (x– = 2.92, n = 5); width ade Archipelago ...... T. louisiadensis sp. n. 1.05-1.12 mm, (x– = 1.08, n = 5). Micropterous forms - Ventral concavity on fore femur short, occupying unknown. extreme basal section of femur; fore femur when Size (Misima Island population). – Macropterous viewed laterally attaining its greatest thickness on male, length 2.40-2.60 mm (x– = 2.53, n = 7); width basal half; middle tibia only 1.75X the length of 1.00-1.05 mm (x– = 1.02, n = 7). Macropterous female, basal tarsal segment; Raja Ampat Islands ...... length 2.90-3.05 mm (x– = 2.97, n = 5); width 1.05-1.15 ...... T. rajana Polhemus & Polhemus mm, (x– = 1.11, n = 5). Micropterous forms unknown. 8. Anterior lobe of pronotum in macropterous Colour. – Macropterous male: ground colour black forms with orange brown coloration limited to a to dark brownish black, venter slightly lighter. Head transverse spot at center, this coloration separated dark brownish black; eyes dark red; rostrum yellowish from propleurae by dark bands along lateral brown basally, piceous distally. Pronotum with a pronotal margins; fore femur with a small, cari- transverse orange brown patch centrally, this orange nate ridge centrally on ventral face; paramere of brown coloration not extending onto lateral margins

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of pronotum or reaching propleurae; disc and humeri Antennal formula I : II : III : IV; 0.25 : 0.28 : 0.35 blackish brown. Wings dark brown, basal angles yel- : 0.40. lowish white, this white coloration extending posteri- Proportions of legs as follows: Femur, tibia, tarsal orly past the apex of the pronotum; interior section of 1, tarsal 2 of fore leg, 0.62 : 0.60 : 0.25 : 0.0; of mid- central cell and raised patch on distal subcosta black. dle leg, 0.95 : 0.90 : 0.45 : 0.35; of hind leg, 0.75 : Abdomen black, embrowned ventrally. Antennae dark 0.95 : 0.20 : 0.25. brown. Coxae, trochanters, and basal 4/5 of fore femur, Paramere slender, tip rounded, slightly expanded bases of middle and hind femora dark yellow to yel- (fig. 33); proctiger broadly rounded, apex very slight- lowish brown, remainder of legs dark brown. ly angulate (fig. 32). Structural characters. – Macropterous male: head Macropterous female: Similar to male in general of moderate length, declivant anteriorly; length 0.50, structure and coloration, but somewhat larger; anteri- width 0.58; width of eye/interocular space, 0.17 or lobe of pronotum with two (1+1) tufts of stiff erect /0.37. Pronotum long, apex rounded, length/ width, black setae to either side of midline, similar setae also 0.90/1.00; humeri, prominent, raised. Abdominal present on propleura; wing costal margin slightly con- tergites not visible. cave on posterior 0.20 of its length, bearing numerous Abdominal venter not modified, set with short ap- short, curling, dark setae; fore femur slender, tapering pressed setae. evenly, not expanded as in male. Legs and antennae thickly clothed with short ap- Micropterous form: Unknown. pressed gold setae, intermixed with scattered slender erect dark setae; fore femur lacking patches of dark se- Etymology tae ventrally, but ventral surface thickly set with nu- The name ‘louisiadensis’ refers to the Louisiade Ar- merous short, stiff, erect, pale setae. Legs unarmed; chipelago, to which this species is endemic. fore femur slightly incrassate on distal half (fig. 31).

150° 151° 152° 153° 154°

KIRIWINA IS. TROBRIAND ISLANDS

° 9° WOODLARK IS. 9

GOODENOUGH IS.

FERGUSSON IS. COLLINGWOOD BAY

D'ENTRECASTEAUX ISLANDS EGUM ATOLL

NORMANBY IS. NEW GUINEA GOODENOUGH BAY SOLOMON SEA 10°

MILNE BAY MISIMA IS. ORANGERIE BAY Tarsovelia ENGINEER GROUP DEBOYNE IS. louisiadensis

LOUISIADE ARCHIPELAGO CONFLICT GROUP 11° 11°

ROSSEL IS.

TAGULA IS. CORAL SEA

150° 151° 152° 153° 154°

Fig. 34. Distribution of Tarsovelia louisiadensis.

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Fig, 35. Neusterensifer femoralis, male, dorsal habitus. Young Sohn illustration.

Distribution rior pronotum, which is interrupted laterally by a dark Louisiade Archipelago (fig. 34). band along the pleural region, rather than extending continuously across the pleural region to join with the Comparative notes orange-brown coloration of the thoracic venter as in Keys to T. alta in Polhemus & Polhemus (2000b), T. alta; the differently shaped male fore femur, which but easily separated from that species by the generally is slightly constricted on the basal half rather than dark brown rather than orange-brown coloration; the thickened basally as in T. alta (compare fig. 31 in this more restricted orange-brown coloration on the ante- paper to fig. 13 in Polhemus & Polhemus 1994); and

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the differently shaped male paramere (fig. 33). Biological notes Although occupying different high islands separat- This species was abundant at the Hane River, a ed by a significant water gap, the populations on both swift, clear mountain stream in a bed of boulders and Tagula and Misima islands are morphologically simi- cobbles, flowing out of a catchment covered in nearly lar, and are treated as conspecific herein. undisturbed montane rain forest. Tarsovelia alta oc- curred here in sheltered eddies and side pools along Biological notes the main channel, out of the full force of the current. Tarsovelia louisiadensis was abundant along the margins of rocky upland streams on Tagula and Misi- Neusterensifer Polhemus & Polhemus ma Islands, generally preferring eddies sheltered by overhanging rocks in areas of otherwise swift flow. At Neusterensifer Polhemus & Polhemus 1994: 67 (type species the Kolukolu Creek type locality the insects were Neusterensifer compacta Polhemus & Polhemus 1994, by found immediately above a large waterfall, along the original designation). margins of the main channel in an area of relatively Discussion. – For diagnostic structural characters of smooth flow. For further description of this locality this genus see discussion in Polhemus & Polhemus see the discussion under Rheovelia insularis. (1994). In using the descriptions and keys below, it is very Zoogeographic notes important to bear in mind that in Neusterensifer the The presence of this Tarsovelia species in the Louisi- more commonly encountered ‘wingless’ forms are ac- ade Archipelago is surprising, given that the genus is tually micropterous, so that the first abdominal tergite otherwise known only from mainland New Guinea to is covered by the metanotum. The first visible tergite the west of the Kokoda Gap in the Owen Stanley dorsally in both males and females is therefore tergite Range, and on Batanta Island in the Raja Ampat II. In males, the best way to ascertain tergite number- group. It is also notable that recent surveys of suitable ing is count anteriorly from the cylindrical first genital habitats at elevations in excess of 600 m. in both the segment, which represents tergite VIII. Similarly, in Cloudy Mountains of far southeastern mainland New females one may count anteriorly from the proctiger, Guinea, and on the D’Entrecasteaux Islands, failed to which represents tergite IX. produce collections of this genus. The occurrence of Ventrally, abdominal ventrite I is barely visible as a T. louisiadensis on Tagula and Misima therefore lends vertical plate lying between the metasternum and well- support to the hypothesis of Pigram & Davies (1987) developed abdominal ventrite II. By working back- that the Louisiades may be isolated remnants of a for- ward from this it is possible to see that the final con- merly larger and more extensive Owen Stanley tecton- nexival segment is formed by the laterotergites of ic terrane that formed in the Eocene, and implies that abdominal segment VII, thus as in the abdominal ter- Tarsovelia may be a relatively old component of the gites, the first visible connexivum anteriorly is segment New Guinea microveliine biota. II. Similarly, in males the first genital segment is ven- trite VIII, and the ventrite immediately anterior to it, which is often highly modified with sulci, tumescences Tarsovelia alta J. Polhemus & D. Polhemus and setal tufts is ventrite VII (for similar considera- Tarsovelia alta J. Polhemus & D. Polhemus 1994: 64. tions in regard to Rhagovelia, and illustrations of seg- ment numbering, see Polhemus & Polhemus 1988). Material examined. – PAPUA NEW GUINEA, Central Prov.: 1 macropterous male, 3 macropterous females, Owen Stanley Range, Om Creek, trib. to upper Udabe River, 1.85 km. SSE Key to species of Neusterensifer occurring on the of Woitape, 1585 m. [5200 ft.], 8°33’27.7”S, 147°15’ Papuan Peninsula and nearby island groups 21.8”E, water temp. 19 °C., 2 October 2003, 12:00-12:30 hrs., CL 7251, D. A. Polhemus (USNM); 43 macropterous Discussion. – The genus Neusterensifer is large and males, 19 macropterous females, Owen Stanley Range, tribu- complex, and because further species are in the process tary streamlet to Hane River, 1.0 km. SSE of Fane, 1310- 1370 m. [4300-4500 ft.], 8°33’32.7”S, 147°05’23.7”E, wa- of description by the authors, a revised key to the en- ter temp. 20 °C., 4 October 2003, 11:00-12:30 hrs., CL tire genus is not presented at this time. Instead, the 7256, D. A. Polhemus (USNM). key below treats all species known to occur on the land masses comprising the EPCT, including the Papuan Distributional notes Peninsula east of 146° E longitude (ie., roughly south- This species was originally described from Mt. east of a line drawn through the base of the peninsula Kaindi, above Wau, and the new records from the from Lae to Kerema), the D’Entrecasteaux Islands, the Woitape and Fane areas serve to extend its range fur- Marshall Bennett Islands, the Louisiade Archipelago, ther to the southeast along the upland axis of the and the islands immediately east of the China Strait. Owen Stanley Range. No species from the geographical region thus defined

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Figs. 36-38. Neusterensifer species, male hind femur. – 36, N. goilala; 37, N. muyuw,; 38, N. tufi.

36

37

38 is known to range more widely into the remainder of 2. Paired patches of dark setae present on ventrite greater New Guinea except for N. lubu, which occurs VII only (figs. 39, 40, 41, 44) ...... 3 westward as far as the Kikori River catchment, and - Paired patches of dark setae present on ventrites reaches the eastern limits of its range in the Lakekamu other than ventrite VII, or on other ventrites in River catchment, thus barely straying into the bounds addition to ventrite VII (fig. 43) ...... 9 of the EPCT region as defined in this paper. 3. Hind femur with a prominent tumescence on central section of hind margin, this tumescence Males set with dense dark setae, distal section of femur 1. Ventral abdomen with dense, prominent, rough- beyond tumescence often slightly concave on ly ovate to circular paired patches of short dark hind margin (figs. 35-37) ...... 4 setae to either side of midline on one or more Hind femur lacking a prominent tumescence on ventrites (most typically ventrite VII), often sepa- central section of hind margin, femur either un- rated by a deep depression or sulcus running modified or with hind margin bearing only an in- along the longitudinal midline of the ventrite cipient indication of a tumescence or slightly (figs. 39, 40, 41, 43, 44) ...... 2 more dense setiferation (fig. 38) ...... 7 - Ventral abdomen lacking any paired patches of 4. Abdominal ventrite II strongly produced into a dense dark setae to either side of midline, al- sharp, angulate, posteroventrally projecting spur; though such patches may be present singly on process on male proctiger multisinuate and various ventrites along the longitudinal midline acuminate (fig. 62); eastern Papuan Peninsula of the abdomen (fig. 42); more diffuse patches of plus Sariba, Basilaki, and Normanby islands (fig. longer golden to golden brown setae may be pre- 70) ...... N. femoralis sp. n. sent laterally on the abdominal ventrites, but - Abdominal ventrite II not strongly produced into these patches never dense or roughly circular to a sharp spur; process on male proctiger not mul- ovate in shape ...... 14 tisinuate and acuminate, either tapering or ex-

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panded at apex (figs. 68, 69, 76) ...... 5 prominent patches of dark setae that progressive- 5. Abdominal ventrite VI with medial section pro- ly widen and enlarge on each succeeding ventrite duced anteriorly as a narrow, slightly rising carina going posteriorly (fig. 39); apex of process on with a small patch of dark setae at apex, this cari- proctiger acuminate and highly produced beyond na nearly bisecting ventrites III-V (fig. 44); large the expanded head (fig. 73); northern Papuan species for genus, overall body length equal to or Peninsula (fig. 74) ...... N. tufi sp. n. exceeding 2.9 mm; process on male proctiger with 9. Paired patches of dark setae present on ventrite apex expanded and blunt when viewed ventrally VI only; southern Papuan Peninsula ...... 10 (fig. 76); occurring above 1000 m. in the Owen - Paired patches of dark setae present on either ven- Stanley Range (fig. 82) ...... N. goilala sp. n. trites V and VI (fig. 43), or VI and VII ...... 11 - Abdominal ventrite VI not produced anteriorly 10. Abdominal ventrite VI sharply raised posterome- as a narrow carina that bisects ventrites III-V dially to form a pair of angulate tumescences (figs. 40, 41); smaller species, overall body length slightly separated by a shallow sulcus along the less than or equal to 2.6 mm; process on male longitudinal midline, apices of these tumescences proctiger tapering to acute tip, not expanded at black and bearing sparse tufts of long gold setae; apex when viewed ventrally (figs. 68, 69) ...... 6 ventrite VII with a pair of low, longitudinally 6. Abdominal ventrites VI and VII with sides of elongate tumescences to either side of midline, central sulcus parallel, the flanking tufts of dark central portion of posterior margin not modified setae similarly parallel and not convergent anteri- or angulate ...... N. aviavi sp. n. orly (fig. 40); ventrite VI with a single small patch - Abdominal ventrite VI lacking a pair of angulate of dense dark setae centrally; Tagula Island (fig. tumescences, instead bearing two very large, trans- 75) ...... N. sulcata sp. n. versely ovate patches of short dark setae; abdomi- - Abdominal ventrites VI and VII with central sul- nal ventrite VII lacking elongate tumescences, cus V-shaped, with flanking tufts of dark setae central portion of posterior margin angulate and converging inward toward each other anteriorly projecting posteriorly over base of ventrite VIII .... (fig. 41); ventrite VI lacking a small patch of ...... N. lubu Polhemus & Polhemus dense dark setae centrally; Woodlark Island (fig. 11. Paired patches of dark setae present on ventrites 70) ...... N. muyuw sp. n. V and VI (fig. 43); Goodenough and Fergusson 7. Ventral abdomen with a broad and relatively Islands plus eastern Papuan Peninsula (Fig. 75) .. shallow longitudinal sulcus on ventrites IV-VII, ...... N. dentrecasteaux sp. n. this sulcus divided along midline by a narrow - Paired patches of dark setae present on ventrites setiferous carina traversing ventrites III-VI; VI and VII ...... 12 process on proctiger rounded and thumb-shaped 12. Patches of dark setae on ventrite VI located close in ventral view with apex not expanded (fig. 63b), to longitudinal midline of segment and nearly but cup-shaped in lateral view after dissection touching each other; process on male proctiger (fig. 63a); eastern mainland of New Guinea plus short, apex blunt (fig. 65); Rossel Island (fig. 74).. Sariba, Sideia, Basilaki and Normanby islands ...... N. yela sp. n. (fig. 74) ...... N. hunteri sp. n. - Patches of dark setae on ventrite VI widely sepa- - Ventral abdomen with deep longitudinal sulcus rated from each other by longitudinal median sul- along midline of ventrite VII at least; ventrites III- cus; process on male proctiger tapering, apex an- VI with prominent patches of dark setae centrally gulate (figs. 77, 80); Papuan Peninsula ...... 13 (fig. 39); process on proctiger expanded distally 13. Process on male proctiger slender, gently curved, when viewed ventrally, extreme apex produced to apex acute (fig. 77); Bowutu Mountains, northern varying degrees (figs. 72. 73); Goodenough and Papuan Peninsula (fig. 78) ...... N. bowutu sp. n. Fergusson Islands or northern Papuan Peninsula . - Process on male proctiger thicker, apex with ...... 8 small indentation (fig. 80); southern Papuan 8. Ventrites VI and VII both longitudinally sulcate Peninsula (fig. 82) ...... N. microrivula sp. n. medially; ventrites III-V with a medial carina 14. Ventral abdomen with a single patch of dark setae bearing prominent patches of dark setae that pro- centrally on ventrite VI (fig. 42); eastern Papuan gressively narrow and diminish on each succeed- Peninsula (fig. 64) ...... N. sagarai sp. n. ing ventrite going posteriorly; apex of process on - Ventral abdomen without a patch of dark setae proctiger only slightly produced beyond the ex- centrally on ventrite VI ...... 15 panded head (fig. 72); Goodenough and Fergus- 15. Ventral abdomen with ventrite VII sulcate along son Islands (fig. 64) ...... N. kula sp. n. longitudinal midline, this sulcus V-shaped when - Only ventrite VII longitudinally sulcate medially; viewed from above, widening posteriorly; lateral ventrites III-VI with a medial carina bearing margins of medial sulcus bearing long gold setae,

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Figs. 39-44. Neusterensifer species, male terminal abdomen, ventral view. – 39, N. tufi; 40, N. sul- cata; 41, N. muyuw; 42, N. sagarai; 43, N. dentrecasteaux; 44, N. goilala.

39 40

41 42

43 44

these setae becoming slightly denser and darker to Females form a pair of diffuse lateral patches on ventrite 1. Connexival margins convergent and touching to- VII; process on proctiger short, blunt, not distally gether to greater or lesser degrees at posterior expanded (fig. 66); Misima Island (fig. 75) ...... apices ...... 2 ...... N. misima sp. n. - Connexival margins separated from each other for - Ventral abdomen with ventrite VII broadly de- entire lengths, not touching at any point ...... 13 pressed along longitudinal midline, but not 2. Connexival margins strongly convergent and deeply sulcate; lateral margins of medial sulcus closely appressed along entirety of segment VII, bearing long gold setae, but these setae not form- and sometimes segments V and VI as well ...... 3 ing a pair of slightly denser and darker patches on - Connexival margins touching only at extreme ventrite VII; process on proctiger short, strongly posterior apices, not appressed along entirety of expanded distally (fig. 67); Rossel and Tagula is- segment VII, and sometimes segments V and VI lands (fig. 70) ...... N. louisiadae sp. n. as well ...... 10 3. Abdominal tergite VII produced into a finger-

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like, posteriorly directed, setiferous projection laterally (fig. 55); Rossel and Tagula islands (fig. that extends beyond the posterior margins of 70) ...... N. louisiadae sp. n. connexival segment VII (figs. 45, 46, 49-52) ... 4 - Overall coloration orange brown; connexiva - Abdominal tergite not produced into a finger- tightly appressed on segment VII only, leaving like, posteriorly directed projection extending be- tergites V visible from above; posterolateral angle yond posterior margins of connexival segment of connexivum with acuminate tuft of stiff black VII (fig. 47, 48, 53-60) ...... 6 setae when viewed laterally (fig. 47); Misima Is- 4. Posterior projection on abdominal tergite VII land (fig. 75) ...... N. misima sp. n. short and thumb-like, barely extending beyond 9. Connexival margins closely appressed only along posterior margins of connexival segment VII (fig. segments VI and VII; connexival segments VI and 52); posterolateral angles of connexival segment V VII lacking acuminate tufts of stiff black setae lacking tufts of stiff black setae on inner faces; projecting upward from between appressed mar- southern Papuan Peninsula (fig. 82) ...... gins, but outer margins of segment VII fringed ...... N. microrivula sp. n. with numerous long, stiff black setae (fig. 56); - Posterior projection on abdominal tergite VII margins of connexival segment VII sloping long, prominent and finger-like, extending well strongly downward posteriorly when viewed later- beyond posterior margins of connexival segment ally (fig. 56); occurring above 1000 m. in the VII (figs. 46, 51) ...... 5 Owen Stanley Range (fig. 82) .... N. goilala sp. n. 5. Posterolateral angles of connexival segment V - Connexival margins closely appressed along seg- with tufts of stiff black setae on inner faces, so ments V-VII; connexival segments VI and VII that abdomen possesses two acuminate, upright bearing acuminate tufts of stiff black setae origi- tufts of black setae when viewed laterally (fig. 46); nating on their central inner faces and projecting Tagula Island (fig. 75) ...... N. sulcata sp. n. upward between the appressed margins, appearing - Posterolateral angles of connexival segment V as a pair of acuminate tufts when viewed laterally lacking tufts of stiff black setae on inner faces, so (fig. 48); margins of connexival segment VII not that abdomen possesses only a single acuminate sloping strongly downward when viewed laterally tuft of black setae when viewed laterally (fig. 51); (fig. 48); Rossel Island (fig. 74) ...... N. yela sp. n. Bowutu Mountains, northern Papuan Peninsula 10. Abdominal tergite VII produced into a finger- (fig. 78) ...... N. bowutu sp. n. like posteriorly-directed, setiferous projection 6. Connexival margins closely convergent and tight- that extends beyond the posterior margins of con- ly appressed along segment VII only ...... 7 nexival segment VII (fig. 45, 49, 50) ...... 11 - Connexival margins closely convergent and tightly - Abdominal tergite not produced into a finger-like appressed along segments VI and VII or V-VII ..... posteriorly-directed projection extending beyond ...... 9 posterior margins of connexival segment VII (fig. 7. Hind femur lacking a tumescence; connexival 54); eastern mainland of New Guinea plus Sari- segment VII bearing an acuminate tuft of stiff ba, Sideia, Basilaki and Normanby islands (fig. black setae originating on the central inner faces 74) ...... N. hunteri sp. n. of the segment and projecting upward between 11. Connexival segment VII in lateral view with a the appressed margins (figs. 47, 55); Louisiade small, acuminate, upwardly-directed tuft of stiff Archipelago ...... 8 black setae lying immediately anterior to the large - Hind femur bearing a small tumescence along tuft of stiff black setae on the projecting process posterior margin; outer margins of connexival seg- of abdominal tergite VII (figs. 45, 49); Woodlark ment VII bearing stiff, erect black setae along en- Island and D’Entrecasteaux Islands ...... 12 tire length, with these setae forming a promiment, - Connexival segment VII in lateral view lacking a acuminate, posteriorly directed tuft at the pos- small, acuminate, upwardly-directed tuft of stiff terolateral angles (fig. 53), but lacking an acumi- black setae immediately anterior to the large tuft nate tuft of stiff black setae originating on central of stiff black setae on the projecting process of ab- inner faces of segment and projecting upward be- dominal tergite VII (fig. 50); northern Papuan tween appressed margins; Papuan Peninsula plus Peninsula (fig. 74) ...... N. tufi sp. n. Sariba, Basilaki, and Normanby islands (fig. 70) .. 12. Finger-like process on tergite VII when viewed ...... N. femoralis sp. n. laterally projecting to or beyond the posterior 8. Overall coloration dark brown; connexiva tightly margin of the abdomen (fig. 45); inner faces of appressed on segments VI and VII, completely connexival segments VI set with patches of long, covering underlying tergites, tergite V not visible stiff black setae posteriorly; small tuft of stiff from above; posterolateral angle of connexivum black setae arising from inner faces of connexival with a brushy tuft of stiff black setae when viewed segments VII near their middle and projecting

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45 49

46 50

47 51

48 52

Figs. 45-52. Neusterensifer species, female terminal abdomen, lateral view. – 45, N. kula; 46, N. sulcata; 47, N. misima; 48, N. yela; 49, N. muyuw; 50, N. tufi; 51, N. bowutu; 52, N. microrivula.

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upward and between the appressed connexival Description margins (fig. 45); Goodenough and Fergusson is- Size. – Micropterous male, length = 2.05-2.20 mm lands (fig. 64) ...... N. kula sp. n. (x– = 2.12, n = 5); width 0.85-1.00 mm (x– = 0.92, n = - Finger-like process on tergite VII when viewed 5). Micropterous female, length 2.25-2.45 mm (x– = laterally not projecting beyond the posterior mar- 2.34, n = 5); width 0.90-1.00 mm, (x– = 0.96, n = 5). gin of the abdomen (fig. 49); inner faces of con- Colour. – Micropterous male: Ground colour dark nexival segments VI lacking patches of long, stiff brown, marked with orange brown on anterior prono- black setae; posterolateral angle of connexival seg- tum and connexiva; patches of short, shining, silvery ment VII bearing small acuminate tuft of stiff pubescence present laterally on metanotum and ab- black setae (fig. 49); Woodlark Island (fig. 70) .... dominal tergites II-V, along anterior margin of prono- ...... N. muyuw sp. n. tum, and on sides of thorax and abdomen. Head dark 13. Posterolateral angles of connexival segment VII brown; rostrum pale whitish yellow, fuscous medially, produced into angulate projections, which may be piceous distally. Pronotum dark brown, becoming or- either upright or infolded over tergite VII (figs. ange brown anterolaterally, bearing a pair (1+1) of 59, 60); southern Papuan Peninsula ...... 14 elongate transverse yellowish brown spots to either - Posterolateral angles of connexival segment VII side of midline behind anterior margin. Metanotum not produced into angulate projections (figs. 57, and abdominal tergites dark brown, laterotergites 58); eastern Papuan Peninsula or D’Entre- blackish below connexiva; connexiva orange brown on casteaux Islands ...... 15 outer halves and ventrally. Antennae medium brown. 14. Angulate projections at posterolateral angles of Legs generally medium brown, with coxae, trochan- connexival segment VII upright, angling back- ters, basal femora and all ventral surfaces pale yellow- ward when viewed laterally (fig. 60) ...... ish to yellowish brown...... N. lubu Polhemus & Polhemus Structural characters. – Micropterous male: head - Angulate projections at posterolateral angles of of moderate length, declivant anteriorly, with weak connexival segment VII folded inward over lateral impressed median line; length 0.45, width 0.57; sections of tergite VII (fig. 59) ..... N. aviavi sp. n. width of eye/interocular space, 0.12/0.35. Pronotum 15. Inner margins of connexival segments V-VII with long, covering mesonotum, thickly set with obscure a thick fringe of dark setae (fig. 58); Goodenough foveae, humeri depressed; length/width, 0.38/0.85. and Fergusson Islands plus eastern Papuan Metanotum length 0.17. Abdominal tergites dull, Peninsula (fig. 75) ...... N. dentrecasteaux sp. n. without shining areas; lengths of tergites II-VII, re- - Inner margins of connexival segments V-VII lack- spectively: 0.10: 0.08 : 0.07 : 0.07 : 0.10 : 0.25. ing a thick fringe of dark setae (fig. 57); eastern Entire dorsum and laterotergites covered with fine Papuan Peninsula (fig. 64) ...... N. sagarai sp. n. appressed golden pubescence, intermixed with numer- ous short, erect black setae; legs and antennae thickly Neusterensifer sagarai sp. n. clothed with short pale setae, scattered long erect black (figs. 42, 57, 61, 64) setae present along anterior margins of antennal seg- ments I-III, anterior margins of fore and middle tibi- Material examined. – Holotype, micropterous ae, posterior margins of middle and hind femora; fore male: PAPUA NEW GUINEA, Milne Bay Prov., Upalai femur ventral margin straight, not modified, bearing River at Haluwia, 13.7 km. E. of Alotau on East Cape numerous short, erect pale setae along entire length, road, terminal reach, 0-20 m. [0-65 ft.], 10°20’28”S, intermixed with evenly spaced comb of 10 long, erect, 150°34’23”E, water temp. 26-26.5 °C., 4 April 2002, pilose golden setae; middle tibia bearing a row of even- 09:30-12:00 hrs., CL 7162, D. A. & J. T. Polhemus ly spaced, long pilose black setae of progressively de- (USNM). – Paratypes: PAPUA NEW GUINEA, Milne Bay creasing length, longest and most basal of these setae Prov.: 2 micropterous males, 12 micropterous fe- with length approximately equal to ⅓ that of tibia; males, same data as holotype (USNM, BPBM); 2 mi- hind femur with numerous short, erect pale setae cropterous males, 2 micropterous females, clear rocky along posterior margin, lacking tumescences or spe- stream at Tautili, SW of Alotau, 30 m. [100 ft.], cialized setal tufts. 10°29’05”S, 150°06’33”E, water temp. 27 °C., 31 Abdominal venter set with short appressed gold se- January 2002, 10:30 hrs., CL 7159, D. A. Polhemus tae, and scattered longer dark setae; central section of (USNM); 7 micropterous males, 15 micropterous fe- ventrite VII depressed, forming a broad concavity on males, Gumini River at road bridge, SW of Alotau, 15 underside of terminal abdomen, lateral sections of this m. [50 ft.], 10°19’08.0” S, 150°14’06.8” E, water concavity extending forward onto ventrite VI, isolat- temp. 30 °C. (main channel), 28 °C. (shaded side ing a small medial tumescence bearing a dense tuft of pools), 21 January 2004, 10:30-14:00 hrs., CL 7174, short black setae (fig. 42); ventrite V also slightly D. A. & J. T. Polhemus (USNM, JTPC, BPBM). tumescent medially, bearing a few short black setae

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53 57

54 58

55 59

56 60

Figs. 53-60. Neusterensifer species, female terminal abdomen, lateral view. – 53, N. femoralis; 54, N. hunteri; 55, N. louisiadae; 56, N. goilala; 57, N. sagarai; 58, N. dentrecasteaux; 59, N. aviavi; 60, N. lubu.

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along posterior margin; ventrites I-IV unmodified. Biological notes Proctiger produced anteriorly into a moderately long, This species occurs in small pockets of protected broad, blade-like process with a widened apex (fig. water amid cobbles along the margins of lower stream 61). Parameres vestigial or absent. midreaches in both the Milne Bay and Sagarai River Antennal formula I : II : III : IV; 0.30 : 0.24 : 0.28 catchments. It is notably absent from higher gradient : 0.50. upper midreach and headwater areas with greater cur- Proportions of legs as follows: Femur, tibia, tarsal rent velocities lying above 50 m. elevation. 1, tarsal 2 of fore leg, 0.57 : 0.50 : 0.21 : 0.0; of mid- At the Upalai River type locality this species was dle leg, 0.70 : 0.68 : 0.13 : 0.25; of hind leg, 0.75 : found only along the lower midreach of the river where 0.95 : 0.10 : 0.25. it exited from a narrow valley and made a transition Micropterous female: Similar to micropterous male from a high gradient bed of rocks and boulders to a low in general structure and colour, with following excep- gradient bed of gravel and cobbles. Along the higher tions: metanotum and abdominal tergites II-IV with gradient upper midreach of the river, by contrast, N. small patches of silvery setae laterally, abdominal ter- sagarai dropped out and was replaced by N. femoralis gite V covered with silvery setae for its entire width on and N. hunteri; for further habitat notes on this locali- posterior half, abdominal tergite VI covered with sil- ty see the discussion under Aegilipsicola peninsularis. very setae across its entire width on posterior ⅔, ab- dominal tergites VII and VIII bearing a few scattered Neusterensifer femoralis sp. n. silvery setae on posterior halves, tergite VII also with a (figs. 35, 53, 62, 70) transverse tuft of long, posteriorly-directed black setae along posterior margin, tergite VIII and proctiger lying Material examined. – Holotype, micropterous male: in a nearly vertical orientation; connexiva broadly sep- PAPUA NEW GUINEA, Milne Bay Prov., headwater reach arated, bowing outward, margins gradually convergent of Goilayoli River above crossing on road from Watu- posteriorly, posterolateral connexival angles simple, nou to Huhuna, 18.5 km. ENE of Alotau, 275 m. rounded, not produced, lacking tufts of black setae (al- [900 ft.], 10°18’43”S, 150°37’16”E, water temp. 24.5 though tuft of setae on posterior margin of tergite VII °C., 6 April 2002, 10:00-13:00 hrs., CL 7161, D. A. & can give the appearance of a connexival tuft when in- J. T. Polhemus (USNM). – Paratypes: PAPUA NEW sect is viewed laterally) (fig. 57); lateral sections of ab- GUINEA, Milne Bay Prov.: 6 macropterous males, 1 dominal ventrites curving upward to bases of connexi- macropterous female, 20 micropterous males, 17 mi- val segments, set with numerous long erect black setae, cropterous females, same data as holotype (USNM, JTPC, extreme lateral margins adjoining connexival bases BPBM); 1 macropterous female, 6 micropterous males, with patches of silvery setae; abdominal venter, fore fe- 14 micropterous females, Upalai River at Haluwia, mur, hind femur unmodified; anterior margins of all 13.7 km. E. of Alotau on East Cape road, midreach tibiae and posterior margins of middle and hind femo- and waterfall approx. 2.5 km. above mouth, 45-75 m. ra bearing scattered very long, slender, erect black setae. [150-250 ft.], 10°19’39”S, 150°34’36”E, water temp. Macropterous form: Unknown. 24.5 °C., 5 April 2002, 09:45-16:30 hrs., CL 7163, D. A. & J. T. Polhemus (USNM); 2 macropterous males, 3 Etymology macropterous females, streamlet and roadside seeps The name ‘sagarai’ is a noun in apposition and above Huhuna on road to Watunou, 11.2 mi. ENE of refers to the Sagarai River basin of far southeastern Alotau, 260 m. [850 ft.], 10°18’19”S, 150°36’56”E, New Guinea, where this species was first collected. water temp. 28 °C., 6 April 2002, 14:00-15:00 hrs., CL 7164, D. A. & J. T. Polhemus (USNM); 6 mi- Distribution cropterous males, 14 micropterous females, Cloudy Far eastern New Guinea (fig. 64). Mountains, headwater tributary to upper Watuti Riv- er, S. of Gelemalaia, 715 m. [2350 ft.], 10°29’50”S, Comparative notes 150°13’58”E, water temp. 22 °C., 10 April 2002, Neusterensifer sagarai is a small, brown species with 16:00-17:30 hrs., CL 7175, D. A. Polhemus (USNM, relatively few modifications. Males may be recognized BPBM); 2 micropterous males, 1 micropterous female, by the single black setal tuft medially on abdominal Sariba Is., Padi Stream, 0-105 m. [0-350 ft.], ventrite VI (fig. 42) and the shape of the process on 10°35’37.1”S, 150°42’02.5”E, water temp. 26.5-28.5 the proctiger (fig. 61). Females may be separated from °C., pH 7.7-8.0, 14 January 2004, 09:45-14:00 hrs., others occurring in far eastern New Guinea by their CL 7285, D. A. & J. T. Polhemus (USNM); 1 mi- open connexiva which lack tufts of black setae, and by cropterous male, 1 micropterous female, Basilaki Is., the transverse patch of long, posteriorly-directed black Kalawai River and rocky trib., 5-145 m. [20-475 ft.], setae along the posterior margin of abdominal tergite 10°36’50.6”S, 151°01’14.4”E, water temp. 26.5 °C., VII (fig. 57). pH 8.27, 17 January 2004, 09:00-14:00 hrs., CL

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Figs. 61-63. Neusterensifer species, process arising from male proctiger. – 61, N. sagarai, ventral view; 62, N. femoralis, ventral view; 63 a, b, N. hunteri: a, lateral view, b, ventral view.

61 62

63a 63b

7294, D. A. & J. T. Polhemus (USNM); 6 macropterous 6). Micropterous female, length 2.33-2.55 mm (x– = males, 4 macropterous females, 41 micropterous 2.45, n = 6); width 0.95-1.05 mm, (x– = 0.99, n = 6). males, 55 micropterous females, D’Entrecasteaux Is- Macropterous male, length = 2.40 mm (n = 1); width lands, Normanby Island, south coast, upper Apatabuia 1.10 mm (n = 1). Macropterous male, length 2.40- River and rocky tribs., above Bunama, 60-90 m. [195- 2.55 mm (x– = 2.49, n = 4); width 1.10-1.15 mm (x– = 295 ft.], 10°07’04.0”S, 151°09’ 07.2”E, water temp. 1.11, n = 4). Macropterous female, length 2.50-2.70 23.5 °C., 29 January 2003, 09:00-01:00 hrs., CL mm (x– = 2.62, n = 4); width 1.10-1.20 mm, (x– = 1.16, 7228, D. A. & J. T. Polhemus (USNM, JTPC, BPBM); 3 n = 4). micropterous males, 4 micropterous females, D’Entre- Colour. – Micropterous male: Ground colour or- casteaux Islands, Normanby Island, east coast, upper ange brown, marked with yellowish brown on anteri- Dibuwa River and rocky tribs., W. of Yeluyelua, 245 or pronotum and legs; patches of short, shining, sil- m. [800 ft.], 10°02’46.2”S, 151°14’53.0”E, water very pubescence present laterally on metanotum and temp. 24 °C., 30 January 2003, 09:30-12:30 hrs, CL abdominal tergites II-VII, along anterior margin of 7231, D. A. Polhemus (USNM). pronotum, and on sides of thorax and abdomen (fig. 35). Head orange brown; rostrum pale whitish yel- Description low, fuscous medially, piceous distally. Pronotum or- Size. – Micropterous male, length = 2.15-2.25 mm ange brown, darker posteromedially, bearing a pair (x– = 2.19, n = 6); width 0.95-1.00 mm (x– = 0.97, n = (1+1) of elongate transverse yellowish brown spots to

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either side of midline behind anterior margin. Metan- in general structure and colour, with following excep- otum and abdomen orange brown dorsally, extreme tions: metanotum and abdominal tergites II and III lateral and posterior margins of metanotum and ab- with small patches of silvery setae laterally, abdominal dominal tergites black, laterotergites blackish below tergites IV and V covered with silvery setae for their connexiva, connexiva entirely orange brown, some- entire widths on posterior halves, abdominal tergites what lighter along dorsolateral margins. Antennae VI and VII lacking silvery setae, abdominal tergite dark brown. Legs generally medium brown, with cox- VIII with scattered silvery setae centrally; connexiva ae, trochanters, basal femora and all ventral surfaces convergent posteriorly, meeting over and covering ab- pale yellowish white. dominal tergites VI, VII and basal margin of tergite Structural characters. – Micropterous male: head VIII, connexival segments VI and VII with brush of of moderate length, declivant anteriorly, with weak erect black setae along dorsal margins, posterolateral impressed median line; length 0.35, width 0.60; connexival angle of segment VII simple, rounded, not width of eye/interocular space, 0.17/0.40. Pronotum produced, but bearing a slender tuft of elongate, pos- long, covering mesonotum, thickly set with obscure teriorly directed black setae (fig. 53); lateral sections of foveae, humeri depressed; length/width, 0.45/0.90. abdominal ventrites curving upward and inward to Metanotum length 0.20. Abdominal tergites dull, bases of connexival segments, set with numerous long without shining areas; lengths of tergites II-VII, re- erect black setae, extreme lateral margins adjoining spectively: 0.12: 0.10 : 0.07 : 0.10 : 0.10 : 0.30. connexival bases with patches of silvery setae; abdom- Entire dorsum and laterotergites covered with fine inal venter unmodified; fore femur not modified; hind appressed golden pubescence, intermixed with numer- femur with small tumescence near middle of posterior ous long, erect black setae; legs and antennae thickly margin, this tumescence thickly covered with very clothed with short gold setae, scattered long erect short dark setae; anterior margin of middle tibia and black setae present along anterior margins of antennal posterior margins of middle and hind femora bearing segments I-III, anterior margins of fore and middle scattered very long, slender, erect black setae. tibiae, basal posterior margin of middle femur; fore fe- Macropterous male: Similar to micropterous male mur ventral margin sinuate, weakly biconcave, bearing in general structure and colour, with following excep- numerous short, stiff black setae along entire length, tions: posterior lobe of pronotum highly expanded, intermixed with evenly spaced comb of 10 long, erect, brown, set with fine dark punctations, humeri weakly pilose black setae; middle tibia bearing a row of even- tumescent, apex rounded; wings complete, matte ly spaced, long pilose black setae of progressively de- black, extending past tip of abdomen, bearing 4 closed creasing length, longest and most basal of these setae cells, inner basal cell with white streak at basal angle, with length approximately equal to ⅓ that of tibia; remaining 3 cells with small elongate white spots near hind femur with bulging tumescence centrally on pos- posterior apices, costal margin bearing numerous terior margin (fig. 35), this tumescence thickly set long, stiff, erect black setae. with short, stiff black setae. Macropterous female: Similar to micropterous male Abdominal venter set with short appressed gold se- in general structure and colour, with following excep- tae, and scattered longer dark setae; central sections of tions: posterior lobe of pronotum highly expanded, ventrites III-VI depressed, forming a deep, narrow brown, set with fine dark punctations, apex rounded, longtudinal concavity on underside of abdomen; ven- humeri weakly tumescent; wings complete, matte trite II strongly produced medially into an acute, ven- dark brown, extending past tip of abdomen, forewing trally projecting process, posteromedial margin of this bearing 4 closed cells, inner cells with white streaks process forming a low, sharp carina continuing poste- filling basal angles, 2 distal cells with small elongate riorly from base of process down center of medial ab- white spots near posterior apices, costal margin bear- dominal depression and traversing ventrites III-V; ing numerous long, stiff, erect black setae. ventrite VII bearing two (1+1) raised ovate tumes- cences to either side of midline covered with patches Etymology of dense dark setae. Proctiger produced anteriorly into The name ‘femoralis’ refers to the distinctively a relatively long, multi-sinuate, blade-like process with modified male hind femur. an acute apex (fig. 62). Parameres vestigial or absent. Antennal formula I : II : III : IV; 0.35 : 0.30 : 0.40 Distribution : 0.50. Far eastern New Guinea and adjacent Sariba, Basi- Proportions of legs as follows: Femur, tibia, tarsal laki and Normanby islands (fig. 70). 1, tarsal 2 of fore leg, 0.60 : 0.55 : 0.30 : 0.0; of mid- dle leg, 0.80 : 0.75 : 0.12 : 0.25; of hind leg, 0.90 : Comparative notes 1.00 : 0.12 : 0.28. Neusterensifer femoralis is a distinctive species with a Micropterous female: Similar to micropterous male suite of unusual modifications. Males are immediately

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recognizable by the tumescent hind femur (fig. 35), the 1 macropterous male, 4 micropterous males, 2 mi- bisinuate ventral surface of the fore femur, the acumi- cropterous females, Upalai River at Haluwia, 13.7 km. nate projection on abdominal ventrite II, the setiferous E. of Alotau on East Cape road, terminal reach, 0-20 tumescences on abdominal ventrite VII, and the sinu- m. [0-65 ft.], 10°20’28”S, 150°34’23”E, water temp. ate process on the proctiger (fig. 62). Females are easi- 26-26.5 °C., 1 February 2003, CL 7162, J. T. Polhe- ly recognized by the small tumescence on the hind fe- mus (USNM); 6 micropterous males, 3 micropterous mur, the posteriorly convergent connexiva, and the females, streamlet and roadside seeps above Huhuna acuminate tufts of stiff dark setae at the posterior angles on road to Watunou, 11.2 mi. ENE of Alotau, 260 m. of the connexiva (fig. 53). No other species of Neus- [850 ft.], 10°18’19”S, 150°36’56”E, water temp. 28 terensifer currently known has the highly modified °C., 6 April 2002, 14:00-15:00 hrs., CL 7164, D. A. male abdominal ventrite II seen in N. femoralis, al- & J. T. Polhemus (USNM); 1 macropterous female, 3 though the structure is reminiscent of the character micropterous males, 6 micropterous females, Cloudy state seen in the genus Papuavelia from western New Mountains, headwater tributary to upper Watuti Riv- Guinea. The incrassate male hind femur and longitu- er, S. of Gelemalaia, 715 m. [2350 ft.], 10°29’50”S, dinally sulcate male ventral abdomen are unusual char- 150°13’58”E, water temp. 22 °C., 10 April 2002, acter states shared with other Neusterensifer species oc- 16:00-17:30 hrs., CL 7175, D. A. Polhemus (USNM, curring in the D’Entrecasteaux, Marshall Bennett and BPBM); 1 macropterous female, 3 micropterous males, Louisiade island groups, and in the Papuan Peninsula. 4 micropterous females, Cloudy Mountains, rocky Females of N. femoralis can be confused with those stream 0.6 mi. above Gadowalai village, S. of Gele- of N. hunteri, with which this species is often syntopic, malaia, 135 m. [450 ft.], water temp. 24.5 °C., 12 but that latter species lacks the tail-like tuft of setae April 2002, 10:00-10:30 hrs., 10°28’57”S, 150°14’ projecting posteriorly from the connexival angles 27”E, CL 7176, D. A. Polhemus (USNM); 12 mi- (compare figs. 53, 54). Similar to the male characters cropterous males, 9 micropterous females, Sariba Is., discussed above, the posteriorly convergent female Padi Stream, 0-105 m. [0-350 ft.], 10°35’37.1”S, connexiva with the apices bearing projecting acumi- 150°42’02.5”E, water temp. 26.5-28.5 °C., pH 7.7- nate tufts of black setae are a character state also shared 8.0, 14 January 2004, 09:45-14:00 hrs., CL 7285, D. with other Neusterensifer species found on the offshore A. & J. T. Polhemus (USNM); 1 micropterous male, 2 islands east of New Guinea proper. micropterous females, Sideia Is., hill streamlet in south central interior, 35-40 m. [110-130 ft.], 10°36’ Biological notes 16.8”S, 150°50’55.6”E, water temp. 26.5 °C., 15 Jan- This species occurs on small, standing pools in uary 2004, 10:00-11:00 hrs., CL 7287, D. A. & J. T. bedrock adjacent to rocky streams, and on the pools Polhemus (USNM); 1 micropterous male, 2 micropter- of small headwater streamlets. A habitat description ous females, Sideia Is., river in south central interior, of the Goilayoli River type locality may be found in 10°35’59.3”S, 150°50’53.7”E, 20 m. [65 ft.], water the discussion under Rheovelia petrophila. temp. 29 °C., 15 January 2004, 11:30-14:00 hrs., CL 7288, D. A. & J. T. Polhemus (USNM); 1 micropter- Neusterensifer hunteri sp. n. ous male, Sideia Is., Kwabunamoa River, terminal (figs. 54, 63a, 63b, 74, 84) reach, sea level, 10°35’04.5”S, 150°50’51.2”E, water temp. 27 °C.,16 January 2004, 11:00-13:00 hrs., CL Material examined. – Holotype, micropterous 7292, D. A. & J. T. Polhemus (USNM); 7 micropter- male: PAPUA NEW GUINEA, Milne Bay Prov., headwater ous males, 4 micropterous females, Basilaki Is., reach of Goilayoli River above bridge crossing on road Kalawai River and rocky trib., 5-145 m. [20-475 ft.], from Watunou to Huhuna, 18.5 km. ENE of Alotau, 10°36’50.6”S, 151°01’14.4”E, water temp. 26.5 °C., 275 m. [900 ft.], 10°18’43”S, 150°37’16”E, water pH 8.27, 17 January 2004, 09:00-14:00 hrs., CL temp. 24.5 °C., 6 April 2002, 10:00-13:00 hrs., CL 7294, D. A. & J. T. Polhemus (USNM); 2 micropter- 7161, D. A. & J. T. Polhemus (USNM). – Paratypes: ous males, 2 micropterous females, Basilaki Is., Guia- PAPUA NEW GUINEA, Milne Bay Prov.: 1 macropterous goila River and trib. with waterfall, 0-15 m. [0-50 ft.], female, 42 micropterous males, 35 micropterous fe- 10°36’26.7”S, 150°59’37.8”E, water temp. 26.5 °C., males, same data as holotype (USNM, JTPC, BPBM); 1 18 January 2004, 09:00-13:30 hrs., CL 7296, D. A. & macropterous male, 1 macropterous female, 13 mi- J. T. Polhemus (USNM); 12 micropterous males, 23 cropterous males, 18 micropterous females, Upalai micropterous females, D’Entrecasteaux Islands, Nor- River at Haluwia, 13.7 km. E. of Alotau on East Cape manby Island, south coast, lower Apatabuia River and road, midreach and waterfall approx. 2.5 km. above nearby coastal springs at Bunama, 0-10 m. [0-35 ft.], mouth, 45-75 m. [150-250 ft.], 10°19’39”S, 150°34’ 10°07’51.7”S, 151°08’47.7”E, water temp. 26 °C., 28 36”E, water temp. 24.5 °C., 5 April 2002, 09:45- January 2003, 16:00-18:00 hrs., CL 7226, D. A. & J. 16:30 hrs., CL 7163, D. A. & J. T. Polhemus (USNM); T. Polhemus (USNM, JTPC). PAPUA NEW GUINEA, Cen-

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tral Prov.: 6 micropterous males, 9 micropterous fe- ous long, slender, erect black setae; legs and antennae males, Owen Stanley Range, trib. to upper Mimani thickly clothed with short gold setae, scattered long River, 2.0 km. NE of Dorobisoro, 565 m. [1850 ft.], erect black setae present along anterior margins of an- 9°27’20.8”S, 147°56’ 25.5”E, water temp. 22.5 °C., 7 tennal segments I-III, anterior margins of all tibiae, October 2003, 13:40-14:20 hrs., CL 7261, D. A. Pol- posterior margin of hind femur; fore femur with ven- hemus (USNM); 8 micropterous males, 9 micropterous tral margin very weakly sinuate and biconcave, bearing females, Owen Stanley Range, trib. to upper Mimani numerous short, stiff pale setae along entire length, River, 2.7 km. NE of Dorobisoro, 580-685 m. [1900- these setae thicker near base of femur, intermixed with 2250 ft.], 9°27’22.8”S, 147°57’02.8”E, water temp. evenly spaced comb of 10 long, erect, pilose black se- 21.5 °C., 8 October 2003, 09:30-12:30 hrs., CL tae; middle tibia with posterior margin bearing a row 7262, D. A. Polhemus (USNM); 2 micropterous males, of evenly spaced, long pilose black setae of progres- 10 micropterous females, Owen Stanley Range, trib. sively decreasing length, longest and most basal of to upper Mimani River, 2.9 km. NE of Dorobisoro, these setae with length approximately equal to ⅓ that 640-670 m. [2100-2200 ft.], 9°27’18.7”S, 147°56’ of tibia; hind femur slightly incrassate centrally along 52.9”E, water temp. 22.5 °C., 8 October 2003, posterior margin, this incipient tumescence thickly set 12:45-13:30 hrs., CL 7263, D. A. Polhemus (USNM). with numerous short, stiff pale setae. Abdominal venter set with short semi-recumbent Description gold setae; central sections of ventrites IV-VII de- Size. – Micropterous male, length = 2.15-2.20 mm pressed, forming a broad longtudinal concavity on (x– = 2.15, n = 6); width 0.90-0.95 mm (x– = 0.91, n = underside of abdomen; ventrite II tumescent central- 6). Micropterous female, length 2.30-2.60 mm (x– = ly, posteromedial extension of this tumescence form- 2.41, n = 6); width 0.95-1.00 mm, (x– = 0.97, n = 5). ing a prominent, sharp carina continuing posteriorly Macropterous male, length 2.30 mm (x– = 2.30, n = 2); from posterior margin of ventrite II, across ventrite width 1.10-1.15 mm (x– = 1.12, n = 2). Macropterous III, and down center of medial abdominal depression, female, length 2.60-2.70 mm (x– = 2.65, n = 2); width traversing ventrites IV-VI, this carina bearing numer- 1.20 mm, (x– = 1.20, n = 2). ous short gold setae; ventrite VII bearing two (1+1) Colour. – Micropterous male: Ground colour small, raised, ovate tumescences to either side of mid- medium brown, marked with orange brown on con- line covered with patches of dense dark setae. nexiva and anterior pronotum, legs yellowish brown; Proctiger produced anteriorly into a relatively short, patches of short, shining, silvery pubescence present broad, thumb-like process with a small angular pro- laterally on metanotum and abdominal tergites II-VII, jection at apex (fig. 63b), this process cup-shaped along anterior margin of pronotum, and on sides of when dissected and viewed laterally (fig. 63a). Para- thorax and abdomen. Head medium brown; eyes red; meres vestigial or absent. rostrum pale whitish yellow, fuscous medially, piceous Antennal formula I : II : III : IV; 0.35 : 0.25 : 0.40 distally. Pronotum medium brown, becoming orange : 0.50. brown anterolaterally, bearing a pair (1+1) of elongate Proportions of legs as follows: Femur, tibia, tarsal transverse yellowish brown spots to either side of mid- 1, tarsal 2 of fore leg, 0.58 : 0.50 : 0.24 : 0.0; of mid- line behind anterior margin. Metanotum and abdom- dle leg, 0.75 : 0.70 : 0.10 : 0.18; of hind leg, 0.85 : inal tergites I and II medium brown, somewhat lighter 0.95 : 0.10 : 0.25. centrally orange brown dorsally, abdominal tergites Micropterous female: Similar to micropterous male III-VII darker brown, laterotergites blackish below in general structure and colour, with following excep- connexiva, connexiva entirely orange brown. Anten- tions: metanotum and abdominal tergites II-IV with nae dark brown. Legs with coxae, trochanters, basal 4/5 small patches of silvery setae laterally, abdominal ter- of femora yellowish brown, apices of femora, tibiae gite V entirely covered with silvery setae for its entire and tarsi medium brown. width, abdominal tergites VI and VII blackish and Structural characters. – Micropterous male: head faintly pruinose but lacking silvery setae, tergite VIII of moderate length, declivant anteriorly, with weak with scattered silvery setae across entire width on pos- impressed median line; length 0.43, width 0.57; terior ⅔; connexiva convergent posteriorly, meeting width of eye/interocular space, 0.12/0.40. Pronotum over and covering posterior half of abdominal tergite long, covering mesonotum, thickly set with obscure VII and anterior half of abdominal tergite VIII, pos- foveae, humeri depressed; length/width, 0.35/0.85. terolateral connexival angles simple, rounded, not pro- Metanotum length 0.20. Abdominal tergites dull, duced, but bearing a slender, posteriorly directed tuft without shining areas; lengths of tergites II-VII, re- of elongate black setae (fig. 54); lateral sections of ab- spectively: 0.10: 0.10 : 0.08 : 0.06 : 0.08 : 0.30. dominal ventrites curving upward and inward to bases Entire dorsum and laterotergites covered with fine of connexival segments, set with numerous long erect appressed golden pubescence, intermixed with numer- black setae, extreme lateral margins adjoining connexi-

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150° 151° 152° 153° 154°

KIRIWINA IS. Neusterensifer TROBRIAND ISLANDS kula

° 9° WOODLARK IS. 9

GOODENOUGH IS.

FERGUSSON IS. COLLINGWOOD BAY

D'ENTRECASTEAUX ISLANDS EGUM ATOLL SOLOMON SEA NORMANBY IS. NEW GUINEA GOODENOUGH BAY Neusterensifer 10° sagarai

ORANGERIE BAY ENGINEER GROUP DEBOYNE IS. MISIMA IS.

LOUISIADE ARCHIPELAGO CONFLICT GROUP 11° 11°

ROSSEL IS.

TAGULA IS. CORAL SEA

150° 151° 152° 153° 154°

Fig. 64. Distribution of Neusterensifer species in the far eastern New Guinea region.

val bases with patches of silvery setae; abdominal ven- Distribution ter unmodified; fore femur not modified; hind femur Eastern New Guinea and adjacent Sariba, Sideia, very weakly swollen centrally on posterior margin, this Basilaki and Normanby islands (fig. 74). area thickly covered with very short, pale, upright setae; anterior margins of all tibiae and posterior margins of Comparative notes middle tibia hind femur bearing scattered very long, Neusterensifer hunteri is relatively small, orange- slender, erect black setae. brown species that exhibits an incipient development Macropterous male: Similar to micropterous male of the character states seen N. femoralis, with which it is in general structure and colour, with following excep- often syntopic on the hill streams of far eastern New tions: posterior lobe of pronotum highly expanded, Guinea and nearby islands. The overall coloration brown, set with fine dark punctations, apex rounded, trends more toward medium brown with orange humeri weakly tumescent; wings complete, matte brown accents, rather than being generally orange black, extending past tip of abdomen, forewing bear- brown with yellowish brown highlights as in N. femo- ing 4 closed cells, outer basal cell with broad white ralis. The male has a broader longitudinal sulcus on the streak filling basal angle, 2 distal cells lightly maculate ventral abdomen bisected by a much more prominent, with white centrally, costal margin bearing numerous setiferous medial carina; lacks the large, pointed, poste- long, stiff, erect black setae. riorly-directed process on ventrite II typical of N. Macropterous female: Similar to macropterous femoralis; has the hind femur only incipiently swollen male in general structure and colour, but with pale along the posterior margin (similar to fig. 38); and has maculation in distal cells of forewing more obscure. a much shorter and differently shaped process on the male proctiger (compare figs. 62, 63). Females of N. Etymology hunteri have only a single small tuft of black setae at the The name ‘hunteri’ refers to the first author’s son, posterior angles of the connexiva (fig. 54), rather than Hunter Polhemus. a brushy tuft basal to the angle with a second more acuminate tuft below as in N. femoralis (fig. 53).

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Biological notes male, length 2.25-2.35 mm (x– = 2.29, n = 5); width At the type locality this species was taken from shal- 0.90-1.00 mm (x– = 0.95, n = 5). Micropterous fe- low pools at the bases of small cascades that descended male, length 2.75-2.95 mm (x– = 2.85, n = 5); width in a step-like series down a bedrock chute on a narrow 1.00-1.10 mm, (x– = 1.05, n = 5). tributary gully to the Goilayoli River, in the mountains Colour. – Micropterous male: Ground colour dark east of Alotau. This area was heavily shaded by prima- brown, marked with orange brown on connexiva; ry upland rain forest. For further description of this lo- patches of short, shining, silvery pubescence present cality see the discussion under Rheovelia petrophila. along anterior margin of pronotum, on sides of thorax In general, N. hunteri seems to be an adaptable and abdomen, laterally on metanotum and abdominal species that is capable of exploiting diverse habitats tergites II-V, across entire posterior half of abdominal ranging from coastal springs to rocky streams (fig. 84) tergite VI, and covering entire abdominal tergite VII. and small first order upland streamlets, perhaps ac- Head dark brown; rostrum yellowish brown, fuscous counting for its broad distributional range in eastern medially, piceous distally. Pronotum dark brown, be- Papuan Peninsula and nearby islands (fig. 74). It is coming orange brown anterolaterally, bearing a pair most commonly encountered at spring heads or along (1+1) of elongate transverse yellowish brown spots to first order streams, and rarely along the margins of either side of midline behind anterior margin. Metan- larger lowland rivers, where in the Milne Bay area one otum and abdomen generally dark brown dorsally, instead finds N. sagarai. central sections of metanotum and abdominal tergite I becoming orange brown, extreme lateral and posterior Neusterensifer louisiadae sp. n. margins of metanotum and abdominal tergites black- (figs. 55, 67, 70) ish, connexiva entirely orange brown, laterotergites below connexiva blackish. Antennae dark brown. Legs Material examined. – Holotype, micropterous generally medium brown, with coxae, trochanters, male: PAPUA NEW GUINEA, Milne Bay Prov., Louisiade and basal femora pale yellowish white. Archipelago, Rossel [Yela] Island, Woa River and Structural characters. – Micropterous male: head rocky tributaries, approx. 4 km. upstream from estu- of moderate length, declivant anteriorly, with weak ary, 0-150 m. [0-500 ft.], 11°20’36.7”S, 154°07’ impressed median line; length 0.40, width 0.62; 06.4”E, water temp. 23 °C., 31 August-1 September width of eye/interocular space, 0.12/0.37. Pronotum 2002, CL 7196, D. A. & J. T. Polhemus (USNM). – long, covering mesonotum, thickly set with obscure Paratypes: PAPUA NEW GUINEA, Milne Bay Prov., foveae, humeri depressed; length/width, 0.40/0.80. Louisiade Archipelago: 21 micropterous males, 21 mi- Metanotum length 0.20. Abdominal tergites dull, cropterous females, Rossel [Yela] Island, same data as without shining areas; lengths of tergites II-VII, re- holotype, D. A. & J. T. Polhemus (USNM, JTPC, spectively: 0.14: 0.10 : 0.08 : 0.10 : 0.10 : 0.32. BPBM); 15 micropterous males, 27 micropterous fe- Entire dorsum and laterotergites covered with fine males, Tagula [Sudest] Island, Kalitau Creek, 1 km. appressed golden pubescence, intermixed with numer- upstream from head of estuary, 0-30 m. [0-100 ft.], ous long, erect golden setae; legs and antennae thickly 11°21’51.7”S, 153°14’26.0”E, water temp. 24.4 °C., clothed with short gold setae, scattered long erect 29 August 2002, 09:30-14:30 hrs., CL 7190, D. A. & black setae present along anterior margins of antennal J. T. Polhemus (USNM); 2 micropterous males, 1 mi- segments I-III, anterior margins of fore and middle cropterous female, Tagula Island, small spring fed tibiae, basal posterior margin of middle femur; fore fe- stream near Kalitau Creek, 45 m. [150 ft.], 11°21’ mur with ventral margin very weakly sinuate, bearing 29.7”S, 153°14’39.9”E, water temp. 26 °C., 29 Au- moderately short, erect, brushy black setae along en- gust 2002, 10:00-12:00 hrs., CL 7192, D. A. & J. T. tire length, these setae more numerous basally and in- Polhemus (USNM, BPBM); 1 micropterous female, Tag- termixed with evenly spaced comb of about 14 long, ula Island, Kolukolu Creek, near Araeda village, 0-105 erect, pilose golden setae; middle tibia bearing a row of m. [0-350 ft.], 11°26’44.7”S, 153°25’56.8”E, water evenly spaced, long pilose black setae of progressively temp. 23 °C., 30 August 2002, 11:00-16:00 hrs., CL decreasing length along posterior margin, longest and 7194, D. A. & J. T. Polhemus (USNM, BPBM). most basal of these setae with length approximately equal to ⅓ that of tibia; hind femur simple and un- Description modified, lacking setal tufts. Size (Rossel Island population). – Micropterous Abdominal venter set with numerous erect, moder- male, length 2.20-2.30 mm (x– = 2.25, n = 5); width ately long gold setae; central sections of ventrites IV- 0.90-0.95 mm (x– = 0.91, n = 5). Micropterous fe- VII depressed, forming a deep longtudinal sulcus on male, length 2.50-2.85 mm (x– = 2.66, n = 5); width underside of abdomen; ventrites II and III slightly 0.95-1.05 mm, (x– = 1.00, n = 5). raised centrally, densely setiferous. Proctiger produced Size (Tagula Island population). – Micropterous anteriorly into a relatively short, broad, blade-like

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process with expanded, palmate apex (fig. 67). Para- cent to the upper Woa River, occurring sympatrically meres vestigial or absent. with N. yela. On Tagula Island N. louisiadae was tak- Antennal formula I : II : III : IV; 0.40 : 0.30 : 0.45 en from similar standing pools in rockholes next to a : 0.52. waterfall cascading over a sloping face of metamor- Proportions of legs as follows: Femur, tibia, tarsal phic bedrock, in company with N. sulcata. 1, tarsal 2 of fore leg, 0.60 : 0.57 : 0.30 : 0.0; of mid- Further habitat description of the Woa River type dle leg, 0.90 : 0.85 : 0.13 : 0.25; of hind leg, 0.90 : locality may be found in the discussion under 1.10 : 0.20 : 0.30. Rheovelia insularis. Micropterous female: Similar to micropterous male in general structure and colour, with following excep- Neusterensifer yela sp. n. tions: metanotum and abdominal tergites II and III (figs. 48, 65, 74) with small patches of silvery setae laterally, abdominal tergite IV lacking silvery setae, abdominal tergite V Material examined. – Holotype, micropterous entirely covered with silvery setae, abdominal tergites male: PAPUA NEW GUINEA, Milne Bay Prov., Louisiade VI and VII hidden; connexiva convergent posteriorly, Archipelago, Rossel [Yela] Island, Woa River and meeting over and covering abdominal tergites VI-VII rocky tributaries, approx. 4 km. upstream from estu- plus base of tergite VIII, posterolateral connexival an- ary, 0-150 m. [0-500 ft.], 11°20’36.7”S, 154°07’ gles simple, obtusely angulate, bearing a brushy tuft of 06.4”E, water temp. 23 °C., 31 August-1 September stiff black setae, the posterior portion of this brushy 2002, CL 7196, D. A. & J. T. Polhemus (USNM). – tuft denser and somewhat acuminate (fig. 55), lateral Paratypes: PAPUA NEW GUINEA, Milne Bay Prov., sections of abdominal ventrites black, curving upward Louisiade Archipelago, Rossel [Yela] Island: 19 micro- and inward to bases of connexival segments, extreme pterous males, 15 micropterous females, same data as lateral margins adjoining connexival bases with broad holotype, D. A. & J. T. Polhemus (USNM, JTPC, patches of silvery setae; abdominal venter and legs un- BPBM). modified. Macropterous form: Unknown. Description Size. – Micropterous male, length 2.40-2.60 mm Etymology (x– = 2.50, n = 5); width 0.90-1.05 mm (x– = 0.99, n = The name ‘louisiadae’ refers to the Louisiade Is- 5). Micropterous female, length 2.85-3.00 mm (x– = lands, to which this species is endemic. 2.92, n = 5); width 1.00-1.10 mm, (x– = 1.04, n = 5). Colour. – Micropterous male: Ground colour light Distribution orange brown, marked with yellowish brown on ante- Louisiade Archipelago (fig. 70). rior pronotum and legs; very small patches of short, shining, silvery pubescence present laterally on metan- Comparative notes otum and abdominal tergites II-VII, and along anteri- Neusterensifer louisiadae occurs on both Rossel and or margin of pronotum. Head light orange brown; Tagula islands, and is easily separated from the other rostrum pale whitish yellow, fuscous medially, piceous Neusterensifer species in the Louisiade Archipelago by distally. Pronotum light orange brown, bearing a pair its dark brown coloration, male abdominal structures (1+1) of faintly indicated elongate transverse yellowish (fig. 67), and female connexival modifications (fig. spots to either side of midline behind anterior margin. 55). The venter of the male abdomen has a broad lon- Metanotum and abdomen light orange brown dorsal- gitidinal sulcus centrally that is not nearly as deep or ly, lateral margins of metanotum and all abdominal narrow as that seen in N. sulcata from Tagula or N. tergites plus extreme posterior margins of metanotum yela from Rossel, and unlike those two species has no and abdominal tergites II and III narrowly black, lat- specialized setal patches. The apex of the process on erotergites below connexiva narrowly black along su- the proctiger is also different in shape from that in N. tures and adjoining spiracles; connexiva entirely light sulcata and N. yela, being palmate rather than elongate orange brown. Antennae orange brown, segment IV (compare figs. 65, 67, 68). Females of N. louisiadae are darker. Legs generally yellowish brown with tarsi dark- easily separated from the two above species by the er, coxae, trochanters, basal femora and all ventral sur- structure of the abdomen, having no posteromedial faces pale yellowish white. projection, and connexiva with only a single setal tuft, Structural characters. – Micropterous male: head at the posterior angles (compare figs. 46, 48, 55). of moderate length, declivant anteriorly, with weak impressed median line; length 0.55, width 0.65; Biological notes width of eye/interocular space, 0.20/0.40. Pronotum On Rossel Island his species was taken from stand- long, covering mesonotum, thickly set with obscure ing pools in exposures of metamorphic bedrock adja- foveae, humeri depressed; length/width, 0.48/0.92.

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Metanotum length 0.22. Abdominal tergites dull, connexival apices of segment VII slightly produced without shining areas; lengths of tergites II-VII, re- and angulate, another acuminate tuft of stiff black se- spectively: 0.15: 0.10 : 0.08 : 0.08 : 0.12 : 0.42. tae arising on the inner margins of connexival seg- Entire dorsum and laterotergites covered with fine ments VII and projecting upward between the ap- appressed golden pubescence, intermixed with numer- pressed dorsal margins of the segments (fig. 48); lateral ous long, slender, erect black setae; legs and antennae sections of abdominal ventrites curving upward and thickly clothed with short gold setae, scattered long inward to bases of connexival segments, set with scat- erect black setae present along anterior margins of an- tered moderately long erect black setae, extreme later- tennal segments I-III, anterior margins of fore and al margins adjoining connexival bases black, overlain middle tibiae, basal posterior margin of middle femur; with patches of silvery setae, black coloration also nar- fore femur ventral margin sinuate, weakly biconcave, rowly present along all abdominal sutures; abdominal bearing numerous short, stiff black setae along entire venter, fore femur, and hind femur unmodified. length, intermixed with evenly spaced comb of 10 Macropterous form: Unknown. long, erect, pilose black setae; middle tibia bearing a row of evenly spaced, long pilose black setae of pro- Etymology gressively decreasing length, longest and most basal of The name ‘yela’ is a noun in apposition, and refers these setae with length approximately equal to ⅓ that to the traditional Melanesian name for Rossel Island. of tibia; hind femur with incipient tumescence cen- trally on posterior margin, this area moderately set Distribution with very short, stiff black setae. Louisiade Archipelago (fig. 74). Abdominal venter set with short appressed gold se- tae, and scattered longer erect gold setae; abdominal Comparative notes ventrite II forming a small conical tumescence central- Neusterensifer yela is an orange-colored species from ly, ventrite III unmodified, central sections of ventrites Rossel Island, and is easily distinguished from the syn- IV-VII strongly depressed, forming a deep, narrow, topic N. louisiadae by the highly incised posteroventral longitudinal sulcus on underside of abdomen; ventrite male abdomen, bearing large longitudinally ovate hair VI with dense patch of short, stiff black setae central- patches flanking the medial sulcus on ventrite VII, ly,these setae lying on either side of medial sulcus and and by the elongate, blade-like process of the male directed inward toward its longitudinal axis, lateral proctiger, which comes to a rounded rather than sections of ventrite VII to either side of sulcus forming palmate apex (compare figs. 65, 67). The male hind a pair (1+1) of strongly raised, longitudinally ovate femur shows an incipient development of the swelling tumescences bearing thick tufts of short, stiff black se- and dark hair patches on the central portion of the tae along along their central axes. Proctiger produced posterior margin (similar to the character state shown anteriorly into a moderately long, relatively slender, in fig. 38) that is seen in a much more extreme form in thin, transparent, blade-like process with a rounded N. femoralis from far eastern New Guinea and proxi- apex (fig. 65). Parameres vestigial or absent. mate islands (fig. 35). The female is easily distin- Antennal formula I : II : III : IV; 0.45 : 0.35 : 0.48 guished by the strongly infolded connexiva and the : 0.50. pair of acuminate black setal tufts near the posterior Proportions of legs as follows: Femur, tibia, tarsal apex of the abdomen (fig. 48), best seen in lateral view. 1, tarsal 2 of fore leg, 0.80 : 0.60 : 0.33 : 0.0; of mid- dle leg, 1.00 : 0.95 : 0.15 : 0.35; of hind leg, 1.10 : Biological notes 1.10 : 0.15 : 0.35. This species was taken from small, cut off overflow Micropterous female: Similar to micropterous male pools near the mouth of a tributary to the Woa River, in general structure and colour, with following excep- and in standing pools in exposures of metamorphic tions: metanotum and abdominal tergite II with very bedrock adjacent to the upper reaches of this same limited patches of silvery setae laterally, abdominal river; in the latter situation it occurred sympatrically tergite IV bearing silvery setae on posterior half, ab- with N. louisiadae. A more extensive habitat descrip- dominal tergite VIII with numerous silvery setae, ter- tion of the Woa River type locality may be found in gite VIII and proctiger lying in nearly vertical orienta- the discussion under Rheovelia insularis. tion; connexiva strongly and suddenly convergent posteriorly, meeting over and covering abdominal ter- Neusterensifer sulcata sp. n. gites V-VII, connexival segments VI and VII with (figs. 46, 68, 75) scattered moderately long erect black setae along dor- sal margins, these setae becoming more dense at pos- Material examined. – Holotype, micropterous terior angle of segment VI, forming a small, acumi- male: PAPUA NEW GUINEA, Milne Bay Prov., Louisiade nate, posteriorly-directed tuft (fig. 48), posterolateral Archipelago, Sudest [Tagula] Island, Kolukolu Creek,

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near Araeda village, 0-105 m. [0-350 ft.], 11°26’ thickly clothed with short gold setae, scattered long 44.7”S, 153°25’56.8”E, water temp. 23 °C., 30 Au- erect black setae present along anterior margins of an- gust 2002, CL 7194, D. A. & J. T. Polhemus (USNM). tennal segments I-III, anterior margins of all tibiae, – Paratypes: PAPUA NEW GUINEA, Milne Bay Prov., posterior margin of middle femur; fore femur ventral Louisiade Archipelago, Tagula [Sudest] Island: 56 mi- margin sinuate, weakly biconcave, bearing numerous cropterous males, 73 micropterous females, same data short, stiff black setae along entire length, intermixed as holotype, D. A. & J. T. Polhemus (USNM, JTPC, with evenly spaced comb of 10 long, erect, pilose black BPBM); 8 micropterous males, 19 micropterous fe- setae; middle tibia bearing a row of evenly spaced, long males, Kalitau Creek, 1 km. upstream from head of es- pilose black setae of progressively decreasing length, tuary, 0-30 m. [0-100 ft.], 11°21’51.7”S, 153°14’ longest and most basal of these setae with length ap- 26.0”E, water temp. 24.4 °C., 29 August 2002, proximately equal to ⅓ that of tibia; hind femur with a 09:30-14:30 hrs., CL 7190, D. A. & J. T. Polhemus tumescence centrally on posterior margin, this tumes- (USNM); 6 micropterous males, 9 micropterous fe- cence thickly set with very short, stiff black setae. males, small spring fed stream near Kalitau Creek, 45 Abdominal venter set with short appressed gold se- m. [150 ft.], 11°21’29.7”S, 153°14’39.9”E, water tae, and scattered much longer erect gold setae; ab- temp. 26 °C., 29 August 2002, 10:00-12:00 hrs., CL dominal ventrite II forming a small tumescence cen- 7192, D. A. & J. T. Polhemus (USNM). trally, this tumescence contiguous with a low carina continuing posteriorly on midline of ventrites III-V, Description ventrite VI forming a small tumescence anteromedial- Size. – Micropterous male, length 2.05-2.20 mm ly bearing a tuft of fine gold setae, depressed postero- (x– = 2.13, n = 5); width 0.87-1.00 mm (x– = 0.95, n = medially to form a small sulcus, ventrite VII strongly 5). Micropterous female, length 2.45-2.60 mm (x– = depressed along longitudinal midline, forming a deep, 2.53, n = 5); width 1.00-1.05 mm, (x– = 1.06, n = 5). narrow sulcus, anterior end of this sulcus contiguous Colour. – Micropterous male: Ground colour light with medial sulcus on posterior section of ventrite VI, orange brown, marked with darker brown on dorsal lateral sections of ventrite VII to either side of sulcus thorax and abdomen, legs yellowish brown; small bearing a pair (1+1) of thick, longitudinally ovate tufts patches of short, shining, silvery pubescence present of short, stiff black setae (fig. 40). Proctiger produced laterally on metanotum and abdominal tergites II- anteriorly into a moderately long, acuminate, blade- VIII, and along anterior margin of pronotum. Head like process with a sharply angulate apex (fig. 68). orange brown, darker along midline; eyes silvery red; Parameres vestigial or absent. rostrum pale whitish yellow, fuscous medially, piceous Antennal formula I : II : III : IV; 0.37 : 0.30 : 0.40 distally. Pronotum orange brown, darker centrally and : 0.47. at punctations on posterior lobe, bearing a pair (1+1) Proportions of legs as follows: Femur, tibia, tarsal of faintly indicated elongate transverse yellowish spots 1, tarsal 2 of fore leg, 0.62 : 0.53 : 0.27 : 0.0; of mid- to either side of midline behind anterior margin. dle leg, 0.80 : 0.75 : 0.08 : 0.23; of hind leg, 0.85 : Metanotum and abdomen orange brown dorsally, lat- 0.95 : 0.07 : 0.20. eral and extreme posterior margins of metanotum and Micropterous female: Similar to micropterous male all abdominal tergites narrowly black, posterior por- in general structure, but with following exceptions: tion of tergite VI plus all of tergites VII and VIII overall coloration darker orange brown to rusty medium to dark brown, laterotergites below connexi- brown, metanotum and abdominal tergites II-IV with va narrowly black along sutures and adjoining spira- patches of silvery setae laterally, abdominal tergite V cles; connexiva entirely light orange brown. Antennae bearing silvery setae on posterior half, abdominal ter- medium brown. Legs generally yellowish brown with gite VI hidden beneath infolded connexiva, abdomi- tarsi darker. nal tergite VII produced into posteriorly-directed, fin- Structural characters. – Micropterous male: head ger-like medial process, apex of this process projecting of moderate length, declivant anteriorly, with weak under and beyond posterior angles of connexival seg- impressed median line; length 0.35, width 0.60; ment VII and over base of tergite VIII, abdominal ter- width of eye/interocular space, 0.10/0.35. Pronotum gite VIII with a few scattered silvery setae, this tergite long, covering mesonotum, thickly set with obscure plus proctiger lying in nearly vertical orientation; con- foveae, humeri depressed; length/width, 0.37/0.90. nexiva strongly and suddenly convergent posteriorly, Metanotum length 0.20. Abdominal tergites dull, meeting over and covering abdominal tergites VI and without shining areas; lengths of tergites II-VII, re- VII, connexival segments V-VII with a few moderate- spectively: 0.12: 0.10 : 0.08 : 0.08 : 0.07 : 0.25. ly long erect black setae along dorsal margins, these se- Entire dorsum and laterotergites covered with fine tae forming a small, diffuse tuft at posterior angle of appressed golden pubescence, intermixed with numer- segment V, another acuminate tuft of stiff black setae ous long, slender, erect golden setae; legs and antennae arising on the inner margins of connexival segments

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Figs. 65-69. Neusterensifer species, process arising from male proctiger. – 65, N. yela, ventral view; 66, N. misima, ventral view; 67, N. louisiadae, ventral view; 68, N. sulcata, ventral view; 69, N. muyuw, ventral view.

65 66 67

68 69

VII and projecting upward between the appressed Comparative notes dorsal margins of the segments, posterolateral connex- Neusterensifer sulcata is an orange-brown species en- ival angle of segment VII obtusely angulate, not pro- demic to Tagula (or Sudest) Island, the largest island in duced (fig. 46); lateral sections of abdominal ventrites the Louisiade Archipelago. It is superficially similar in curving upward and inward to bases of connexival seg- structure and coloration to N. yela from nearby Rossel ments, set with scattered moderately long erect black Island, but may be easily separated from that species by setae, extreme lateral margins adjoining connexival the following characters: in N. sulcata the ground bases broadly black, overlain with a few silvery setae, colour is darker orange brown with more extensive black coloration also narrowly present along all ab- overlying dark markings; in males of N. sulcata the me- dominal sutures; abdominal venter and fore femur un- dial sulcus on the ventral abdomen transgresses only modified, and hind femur slightly tumescent and ventrite VII and the base of ventrite VI (fig. 40), rather setiferous centrally on posterior margin. than occurring on ventrites IV-VII as in N. yela, the Macropterous form: Unknown. apex of the process on the male proctiger is acuminate, not rounded as in N. yela (compare figs. 65, 68), and Etymology the male hind femur is tumescent on its posterior mar- The name ‘sulcata’ refers to the deep sulcus on the gin, reminiscent of the character state in N. femoralis abdominal venter of the male. (fig. 35), rather than incipiently tumescent as in N. yela; and females of N. sulcata possess a finger-like pro- Distribution jection with a setiferous apex on abdominal tergite VII Louisiade Archipelago (fig. 75). (fig. 46), and have the connexiva covering only tergites VI and VII, rather than tergites V-VII as in N. yela.

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150° 151° 152° 153° 154°

KIRIWINA IS. TROBRIAND ISLANDS

° 9° WOODLARK IS. 9

GOODENOUGH IS.

FERGUSSON IS. COLLINGWOOD BAY D'ENTRECASTEAUX ISLANDS Neusterensifer EGUM ATOLL SOLOMON SEA muyuw NORMANBY IS. NEW GUINEA GOODENOUGH BAY Neusterensifer 10° femoralis

ORANGERIE BAY Neusterensifer ENGINEER GROUP DEBOYNE IS. MISIMA IS. louisiadae

LOUISIADE ARCHIPELAGO CONFLICT GROUP 11° 11°

ROSSEL IS.

TAGULA IS. CORAL SEA

150° 151° 152° 153° 154°

Fig. 70. Distribution of Neusterensifer species in the far eastern New Guinea region.

Biological notes lage, 305-335 m. [1000-1100 ft.], 10°40’33.8”S, This species was taken in great abundance from 152°40’44.9”E, water temp. 22.5 °C., 4 September standing pools set amid exposures of metamorphic 2002, 10:00-12:00 hrs., CL 7206, D. A. & J. T. Pol- bedrock adjacent to waterfalls on the midreach of hemus (USNM). Kolukolu Creek, a relatively large, swift stream in a partially shaded bed draining the northern slopes of Description Tagula Island. Further notes on this locality may be Size. – Micropterous male, length 2.05-2.10 mm found in the discussion under Rheovelia insularis. (x– = 2.09, n = 5); width 0.80-0.90 mm (x– = 0.85, n = 5). Micropterous female, length 2.30-2.45 mm (x– = 2.35, n = 5); width 0.95-1.00 mm, (x– = 0.97, n = 5). Neusterensifer misima sp. n. – (figs. 47, 66, 75) Macropterous female, length 2.60-2.70 mm (x = 2.65, n = 2); width 1.10-1.15 mm (x– = 1.12, n = 2). Material examined. – Holotype, micropterous Colour. – Micropterous male: Ground colour or- male: PAPUA NEW GUINEA, Milne Bay Prov., Louisiade ange brown, marked with yellowish brown on prono- Archipelago, Misima Island, north coast, Guwana tum and legs; small patches of short, shining, silvery Creek, near Nulia village, 15-30 m. [50-100 ft.], pubescence present laterally on metanotum and ab- 10°39’22.8”S, 152°41’16.5”E, water temp. 22 °C., 3 dominal tergites II and III plus V-VII, along anterior September 2002, 10:00-12:00 hrs., CL 7201, D. A. & margin of pronotum, and laterally on thorax and ab- J. T. Polhemus (USNM). – Paratypes: PAPUA NEW domen. Head orange brown; eyes blackish red; ros- GUINEA, Milne Bay Prov., Louisiade Archipelago, trum pale whitish yellow, fuscous medially, piceous Misima Island: 2 macropterous females, 29 micropter- distally. Pronotum orange brown, bearing a pair (1+1) ous males, 68 micropterous females, same data as of elongate transverse yellowish spots to either side of holotype, D. A. & J. T. Polhemus (USNM, JTPC, midline behind anterior margin. Metanotum and ab- BPBM); 3 micropterous males, 6 micropterous females, domen orange brown dorsally, lateral and extreme south coast, Keyana Creek, above Bwagabwaga vil- posterior margins of metanotum and abdominal ter-

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gites I and II narrowly black, laterotergites below con- metanotum and abdominal tergites II and III with nexiva narrowly black along sutures and adjoining patches of silvery setae laterally, abdominal tergite IV spiracles; connexiva entirely light orange brown. An- lacking silvery setae, abdominal tergite V bearing sil- tennae medium brown. Legs with coxae, trochanters very setae on posterior half, abdominal tergite VII hid- and basal femora yellowish brown, distal femora, all den beneath infolded connexiva, abdominal tergite tibiae and all tarsi medium brown. VIII with scattered silvery setae centrally, this tergite Structural characters. – Micropterous male: head plus proctiger lying in nearly vertical orientation; con- of moderate length, declivant anteriorly, with weak nexiva strongly and suddenly convergent posteriorly, impressed median line; length 0.45, width 0.55; meeting over and covering posterior portion of ab- width of eye/interocular space, 0.10/0.35. Pronotum dominal tergite VI, all of tergite VII, base of tergite long, covering mesonotum, thickly set with obscure VIII, connexival segments VII with a few moderately foveae, humeri depressed; length:width, 0.30 : 0.80. short erect black setae along dorsal margins plus an Metanotum length 0.22. Abdominal tergites dull, acuminate tuft of stiff black setae arising on the inner without shining areas; lengths of tergites II-VII, re- margins and projecting upward between the appressed spectively: 0.10: 0.09 : 0.07 : 0.07 : 0.07 : 0.25. dorsal margins of these segments, posterolateral con- Entire dorsum and laterotergites covered with fine nexival angle of segment VII sharply angulate, not appressed golden pubescence, intermixed with numer- produced (fig. 47); lateral sections of abdominal ven- ous long, slender, erect black setae; legs and antennae trites curving upward and inward to bases of connexi- thickly clothed with short gold setae, scattered long val segments (fig. 47), set with scattered moderately erect black setae present along anterior margins of an- short erect black setae, extreme lateral margins adjoin- tennal segments I-III, anterior margins of all tibiae, ing connexival bases broadly black, overlain with sil- posterior margin of middle femur; fore femur ventral very setae, black coloration also narrowly present margin straight, unmodified, bearing numerous short, along all abdominal sutures; abdominal venter, fore fe- stiff pale setae along entire length, intermixed with mur, and hind femur unmodified. evenly spaced comb of 10 long, erect, pilose black se- Macropterous female: Similar to micropterous fe- tae; middle tibia bearing a row of evenly spaced, long male in general structure and colour, with following pilose black setae of progressively decreasing length, exceptions: posterior lobe of pronotum highly ex- longest and most basal of these setae with length ap- panded, orange brown, set with fine dark puncta- proximately equal to ⅓ that of tibia; hind femur un- tions, apex rounded, humeri weakly tumescent; wings modified. complete, matte dark brown, extending past tip of ab- Abdominal venter set with short appressed gold se- domen, forewing bearing 4 closed cells, both basal tae, and scattered longer erect golden brown setae; ab- cells with white streaks at basal angles, 2 distal cells dominal ventrite II forming a small tumescence cen- lacking pale markings, costal margin bearing numer- trally bearing erect golden setae, this tumescence ous long, stiff, erect black setae. contiguous with a low carina continuing posteriorly Macropterous male: Unknown. on midline of ventrites III-VI, areas immediately later- al to this carina slightly depressed to form a pair (1+1) Etymology of shallow longitudinal sulci, ventrite VII strongly de- The name ‘misima’ is a noun in apposition and pressed along longitudinal midline, forming a V- refers to the island of Misima, to which this species is shaped sulcus with apex directed anteriorly, lateral sec- endemic. tions of ventrites V-VII to either side of sulcus/carina modifications bearing incipient tufts of moderately Distribution long, posteriorly-slanting golden setae. Proctiger pro- Louisiade Archipelago (fig. 75). duced anteriorly into a short, blunt process with a very small angle near the apex (fig. 66). Parameres vestigial Comparative notes or absent. Neusterensifer misima is an orange-brown species Antennal formula I : II : III : IV; 0.40 : 0.32 : 0.42 that is superficially similar to N. yela from Rossel Is- : 0.45. land. It differs from that species in having a much dif- Proportions of legs as follows: Femur, tibia, tarsal ferently shaped process on the male proctiger (com- 1, tarsal 2 of fore leg, 0.60 : 0.62 : 0.25 : 0.0; of mid- pare figs. 65, 66), and by the structure of the female dle leg, 0.70 : 0.75 : 0.10 : 0.25; of hind leg, 0.85 : connexiva, which meet posteriorly to cover only por- 1.05 : 0.12 : 0.30. tions of abdominal tergites VI-VIII, rather than cover- Micropterous female: Similar to micropterous male ing tergites V-VII as in N. yela, and have only a single in general structure, but with following exceptions: acuminate setal tuft in lateral view, rather than two overall coloration darker, dorsal head, thorax and ab- (compare figs. 47, 48). The absence of silvery setae on dominal tergites suffused with medium brown, abdominal tergite IV in both sexes is also a useful di-

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Figs. 71-73. Neusterensifer species, process arising from male proctiger. – 71a, b, N. dentrecasteaux: a, lateral view, b, ventral view; 72, N. kula, ventral view; 73, N. tufi, ventral view.

71b

71a 72

73

Neusterensifer muyuw sp. n. agnostic spot character. Females of N. misima do share (figs. 37, 41, 49, 69, 70, 83) with N. yela and N. sulcata the presence of an acumi- nate tuft of black setae arising from the inner margins Material examined. – Holotype, micropterous of the VII connexival segments and projecting posteri- male: PAPUA NEW GUINEA, Milne Bay Prov., Woodlark orly between the dorsal margins of the infolded VII Island, Suloga Peninsula, stream entering W. side of connexival segments (figs. 46-48), a putative synapo- Suloga Harbor, 0-180 m. [0-590 ft.], 9°12’21.6”S, morphy indicating that all three of these orange- 152°42’29.2”E, water temp. 27-28 °C., 23-24 Janu- brown species endemic to separate islands in the ary 2003, CL 7217, D. A. and J. T. Polhemus (USNM). Louisiades form a closely related group. – Paratypes: PAPUA NEW GUINEA, Milne Bay Prov.: 3 macropterous males, 4 macropterous females, 71 mi- Biological notes cropterous males, 80 micropterous females, same data This species was taken from standing pools amid as holotype, D. A. and J. T. Polhemus (USNM, JTPC, boulders and metamorphic bedrock exposures adja- BPBM). cent to cascades along the midreach of Guwana Creek, a small, clear, steeply-dropping stream de- Description scending from the mountains of western Misima. Size. – Micropterous male, length 2.15-2.20 mm Further habitat notes on this locality may be found in (x– = 2.17, n = 3); width 0.90-1.00 mm (x– = 0.93, n = the discussion under Rheovelia anomala. 3). Micropterous female, length 2.40-2.50 mm (x– = 2.45, n = 3); width 0.90-1.00 mm, (x– = 0.93, n = 3).

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Macropterous male, length 2.50 (n = 1); width 1.10 trally, ventrite VI depressed along longitudinal midline mm (n = 1). Macropterous female, length 2.50-2.70 to form a small sulcus, ventrite VII strongly depressed mm (x– = 2.57, n = 3); width 1.10-1.20 mm, (x– = 1.15, centrally, forming a deep, V-shaped sulcus with apex n = 3). directed anterad and contiguous with narrow medial Colour. – Micropterous male: Ground colour light sulcus on posterior section of ventrite VI, lateral sec- orange brown, sparingly marked with darker brown tions of ventrite VII to either side of sulcus bearing a on dorsal thorax and abdomen, legs yellowish brown; pair (1+1) of thick, ovate tufts of short, stiff black setae small patches of short, shining, silvery pubescence pre- (fig. 41). Proctiger produced anteriorly into a moder- sent laterally on metanotum and abdominal tergites ately long, acuminate, blade-like process with a sharply II-VII, and along anterior margin of pronotum, and angulate apex (fig. 69). Parameres vestigial or absent. on sides of thorax and abdomen. Head orange brown, Antennal formula I : II : III : IV; 0.40 : 0.30 : 0.42 darker along midline; eyes dark red; rostrum pale : 0.50. whitish yellow, fuscous medially, piceous distally. Proportions of legs as follows: Femur, tibia, tarsal Pronotum orange brown, darker anteromedially and 1, tarsal 2 of fore leg, 0.60 : 0.55 : 0.28 : 0.0; of mid- at punctations on posterior lobe, bearing a pair (1+1) dle leg, 0.75 : 0.75 : 0.08 : 0.25; of hind leg, 0.95 : of elongate transverse yellowish spots to either side of 0.98 : 0.10 : 0.20. midline behind anterior margin. Metanotum and ab- Micropterous female: Similar to micropterous male domen orange brown dorsally, lateral and extreme in general structure and coloration, but with following posterior margins of metanotum and all abdominal exceptions: metanotum and abdominal tergites II and tergites narrowly dark brown to blackish, posterior III with patches of silvery setae laterally, abdominal portion of tergite VII plus all of tergite VIII medium tergite IV with small patches of silvery setae laterally to dark brown, laterotergites below connexiva narrow- and along longitudinal midline, abdominal tergites V ly black along sutures and adjoining spiracles; connex- and VI bearing silvery setae on posterior halves, ab- iva entirely light orange brown. Antennae medium dominal tergite VII produced into posteriorly-direct- brown. Legs generally yellowish brown with tibiae and ed, finger-like medial process, apex of this process pro- tarsi darker. jecting under and beyond posterior angles of Structural characters. – Micropterous male: head connexival segment VII and over base of tergite VIII, of moderate length, declivant anteriorly, with weak apex bearing an acuminate tuft of stiff black setae (fig. impressed median line; length 0.50, width 0.55; 49), abdominal tergites VIII and IX with a few scat- width of eye/interocular space, 0.12/0.20. Pronotum tered silvery setae, these tergites lying in nearly vertical long, covering mesonotum, thickly set with obscure orientation; connexiva strongly and suddenly conver- foveae, humeri depressed; length/width, 0.35/0.85. gent posteriorly, meeting over and covering abdomi- Metanotum length 0.20. Abdominal tergites dull, nal tergite VII and base of tergite VIIII, connexival without shining areas; lengths of tergites II-VII, re- segments II-VII with very sparse, moderately long spectively: 0.15: 0.15 : 0.10 : 0.10 : 0.10 : 0.22. erect black setae along dorsal margins, posterolateral Entire dorsum and laterotergites covered with fine connexival angle of segment VII sharply angulate, appressed golden pubescence, intermixed with nu- bearing an acuminate, posteriorly-directed tuft of stiff merous long, slender, erect brown setae; legs and an- black setae (fig. 49); lateral sections of abdominal ven- tennae thickly clothed with short gold setae, scattered trites curving upward and inward to bases of connexi- long erect black setae present along anterior margins val segments, set with scattered moderately long erect of antennal segments I-III, anterior margins of all tib- black setae, extreme lateral margins adjoining connex- iae, posterior margin of middle femur, distal ⅓ of ival bases broadly black, overlain with a few silvery se- posterior margin of hind femur; fore femur ventral tae, black coloration also narrowly present along all margin sinuate, weakly biconcave, bearing numerous abdominal sutures; abdominal venter and fore femur short, stiff black setae along entire length, intermixed unmodified, and hind femur slightly incrassate, incip- with evenly spaced comb of about 10 long, erect, pi- iently swollen on central section of posterior margin. lose black setae; middle tibia bearing a row of evenly Macropterous male: Similar to micropterous male spaced, long pilose black setae of progressively de- in general structure and colour, with following excep- creasing length, longest and most basal of these setae tions: posterior lobe of pronotum highly expanded, with length approximately equal to ⅓ that of tibia; orange brown, set with fine dark punctations, apex hind femur with a tumescence centrally on posterior rounded, humeri weakly tumescent; wings complete, margin (fig. 37), this tumescence thickly set with very matte dark brown, extending past tip of abdomen, short, stiff black setae. forewing bearing 4 closed cells, inner basal cell with Abdominal venter set with short appressed gold se- white streak at basal angle, remaining 3 cells lacking tae, and scattered much longer erect gold setae; ab- pale markings, costal margin bearing numerous long, dominal ventrite II forming a small tumescence cen- stiff, erect black setae.

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Macropterous female: Similar to macropterous J. T. Polhemus (USNM). – Paratypes: PAPUA NEW male in general structure and colour, but with pale GUINEA, Milne Bay Prov., D’Entrecasteaux Islands: 1 streak in inner basal cell obscure; abdominal tergite macropterous male, 7 micropterous males, 27 micro- VII not forming a setiferous posterior projection as in pterous females, Fergusson Island, same data as holo- micropterous form. type, D. A. & J. T. Polhemus (USNM, JTPC, BPBM); 5 macropterous males, 2 macropterous females, 42 mi- Etymology cropterous males, 61 micropterous females, same data The name ‘muyuw’ refers to the traditional Papuan as holotype except 26 January 2003, D. A. & J. T. name for Woodlark Island. Polhemus (USNM); 1 micropterous male, Fergusson Is- land, east coast, Yaya River, N. of Mebulibuli Point, Distribution 0-60 m. [0-200 ft.], 9°30’34.3”S, 150°52’51.1”E, wa- Woodlark Island (fig. 70). ter temp. 25.8 °C., 23 August 2002, 10:00-16:30 hrs., CL 7178, D. A. & J. T. Polhemus (USNM); 3 micro- Comparative notes pterous males, 1 micropterous female, Fergusson Is- Neusterensifer muyuw is most similar morphologi- land, east coast, Mebulibuli Creek and rocky tribu- cally to N. sulcata from Tagula Island, with both tary, 1.3 km. upstream of mouth, S. of Basima, 15-75 species having deep sulci on male abdominal ventrite m. [50-250 ft.], 9°30’54.4”S, 150°52’04.2”E, water VII (compare figs. 40, 41) and acuminate processes on temp. 24 °C., 24 August 2002, 09:00-13:00 hrs., CL the male proctiger (Figs. 68, 69). In N. muyuw, how- 7180, D. A. & J. T. Polhemus (USNM); 2 micropter- ever, the central sulcus on ventrite VII is shaped like a ous males, Goodenough Island, Tuabeda River at posteriorly-opening V when viewed from below, and Kalauna village, 335 m. [1100 ft.], 9°23’01.5”S, the flanking tufts of black setae angle inward toward 150°19’32.0”E, water temp. 22.5 °C., 26 August each other anteriorly (fig. 41), while in N. sulcata the 2002, 10:00-12:00 hrs., CL 7184, D. A. & J. T. Pol- central suclus on ventrite VII follows the longitudinal hemus (USNM); 3 micropterous males, Gumini River midline, and the flanking black setal tufts are parallel at road bridge, SW of Alotau, 15 m. [50 ft.], 10°19’ (fig. 40). The shapes of the processes on the male 08.0” S, 150°14’06.8” E, water temp. 28 °C. (shaded proctiger, though similar, also differ between the two side pools), 21 January 2004, 10:30-14:00 hrs., CL species (compare Figs. 68, 69). 7174, D. A. & J. T. Polhemus (USNM). PAPUA NEW Females of N. muyuw have abdominal tergite VII GUINEA, Oro Prov.: 45 micropterous males, 33 mi- prolonged into a posteriorly-directed, finger-like cropterous females, Kofure River, W. of Tufi, 0-45 m. process with a long tuft of black setae at the apex (fig. [0-150 ft.], 9°04’58.2”S, 149°16’38.7”E, water temp. 49), a character state once again similar that seen in 28 °C., 25 January 2004, 09:30-14:00 hrs., CL 7304, N. sulcata (fig. 46). Females of N. muyuw may be sep- D. A. & J. T. Polhemus (USNM, JTPC). arated from those of N. sulcata by their lighter overall coloration, the presence of a small patch of silvery se- Description tae centrally as well as laterally on abdominal tergite Size. – Micropterous male, length 2.15-2.30 mm IV, and a band of silvery setae running transversely (x– = 2.23, n = 7); width 0.85-1.00 mm (x– = 0.95, n = across the posterior margin of abdominal tergite V. 7). Micropterous female, length 2.50-2.85 mm (x– = 2.74, n = 6); width 1.00-1.10 mm, (x– = 1.04, n = 6). Biological notes Macropterous male, length 2.50-2.70 mm (x– = 2.57, The type locality was a clear, rocky, heavily shaded n = 4); width 1.15-1.20 mm (x– = 1.19, n = 4). stream descending from the forested hills of the Sulo- Macropterous female, length 2.90 (n = 1); width 1.25 ga Peninsula on southwestern Woodlark Island (fig. mm, (n=1). 83). Nesuterensifer muyuw was abundant at this site, Colour. – Micropterous male: Ground colour dark occurring in small, sheltered pockets of water along brown, marked with orange brown on connexiva, yel- the stream margins. lowish brown on basal legs; patches of short, shining, silvery pubescence present along anterior margin of Neusterensifer dentrecasteaux sp. n. pronotum, on sides of thorax and abdomen, laterally (figs. 43, 58, 71, 75, 86) on metanotum and abdominal tergites II-VII. Head dark brown; rostrum yellowish brown, fuscous medi- Material examined. – Holotype, micropterous ally, piceous distally. Pronotum dark brown, becom- male: PAPUA NEW GUINEA, Milne Bay Prov., D’Entre- ing lighter reddish brown anterolaterally, bearing a casteaux Islands, Fergusson Island, east coast, Awae- pair (1+1) of elongate transverse yellowish brown towa River, lower midreach, E. of Basima, 15 m. [50 spots to either side of midline behind anterior margin. ft.], 9°27’34.0”S, 150°49’37.5”E, water temp. 23 °C., Metanotum and abdomen dark brown dorsally, ex- 25 August 2002, 09:45-11:15 hrs., CL 7183, D. A. & treme lateral and posterior margins of metanotum and

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abdominal tergites blackish, connexiva orange brown longitudinal midline, abdominal tergite VIII with nu- on outer halves, laterotergites below connexiva black- merous diffusely scattered silvery setae, this tergite ish. Antennae dark brown. Legs generally medium plus proctiger lying in nearly vertical orientation; con- brown, with coxae, trochanters, and basal femora pale nexiva open and not touching for their entire length, yellowish brown. gradually convergent posteriorly, posterolateral angles Structural characters. – Micropterous male: head simple, gently curving, not produced (fig. 58), margin of moderate length, declivant anteriorly, with weak of connexival segment VII narrowly flattened posteri- impressed median line; length 0.37, width 0.62; orly to form a small, pad-like structure, all connexival width of eye/interocular space, 0.12/0.35. Pronotum segments bearing short, dark, erect setae, these setae long, covering mesonotum, thickly set with obscure becoming longer and denser on inner faces of seg- foveae, humeri depressed; length/width, 0.45/0.92. ments V-VII; lateral sections of abdominal ventrites Metanotum length 0.25. Abdominal tergites dull, black, curving upward into vertical orientation at con- without shining areas; lengths of tergites II-VII, re- nexival bases, margins adjoining connexival bases spectively: 0.15: 0.12 : 0.10 : 0.12 : 0.10 : 0.30. black with broad patches of silvery setae; abdominal Entire dorsum and laterotergites covered with fine venter and legs unmodified. appressed golden pubescence, intermixed with numer- Macropterous male: Similar to micropterous male ous long, erect brown setae; legs and antennae thickly in general structure and colour, with following excep- clothed with short gold setae, scattered long erect tions: posterior lobe of pronotum highly expanded, black setae present along anterior margins of antennal dark blackish brown, set with fine dark punctations, segments I-III, anterior margins of all tibiae, posterior apex rounded, humeri weakly tumescent; wings com- margins of middle and hind femora; fore femur with plete, matte black, extending past tip of abdomen, ventral margin nearly straight, bearing a few short, forewing bearing 4 closed cells, both basal cells with stiff, semi-erect black setae basally plus short pale white streaks at basal angles, 2 distal cells with small, along entire length, these latter setae intermixed with faint, elongate white spots near posterior apices, costal evenly spaced comb of about 10 long, erect, pilose margin bearing long, stiff, erect black setae. golden setae; middle tibia bearing a row of evenly Macropterous female: Similar to macropterous spaced, long pilose golden setae of progressively de- male in general structure and colour. creasing length along posterior margin, longest and most basal of these setae with length approximately Etymology equal to ⅓ that of tibia; hind femur simple and un- The name ‘dentrecasteaux’ is a noun in apposition, modified, lacking setal tufts. and refers to the D’Entrecasteaux Islands, to which Abdominal venter set with numerous erect, moder- this species is endemic. ately long gold setae; central sections of ventrites V and VI narrowly depressed medially to form a contin- Distribution uous, shallow longitudinal sulcus, both ventrites with Goodenough and Fergusson Islands, and the east- a pair (1+1) of dense tufts of short, black setae border- ern Papuan Peninsula (fig. 75). ing sulcus to either side, ventrite VII broadly de- pressed centrally, forming concavity on underside of Comparative notes terminal abdomen (fig. 43). Proctiger produced ante- Neusterensifer dentrecasteaux is a dark colored taxon riorly into a moderately long, broad, blade-like process that is easily separated from the other Neusterensifer with an sharply angulate apex when viewed dorsally, species occurring in far eastern New Guinea by the this apex multi-pronged when viewed ventrolaterally complete covering of silvery setae on abdominal ter- (figs. 71a, b). Parameres vestigial or absent. gites V and VI in females; by the widely separated fe- Antennal formula I : II : III : IV; 0.42 : 0.30 : 0.47 male connexiva that leave all abdominal tergites visi- : 0.50. ble; by the dense fringe of black setae along the inner Proportions of legs as follows: Femur, tibia, tarsal faces of female connexival segments V-VII; by the 1, tarsal 2 of fore leg, 0.62 : 0.60 : 0.35 : 0.0; of mid- male abdominal venter that is only shallowly sulcate dle leg, 0.90 : 0.80 : 0.17 : 0.25; of hind leg, 1.00 : medially but bears dense paired tufts of black setae on 1.10 : 0.17 : 0.30. both ventrites V and VI (fig. 43); and by the distinc- Micropterous female: Similar to micropterous male tively multifurcate apex of the process arising from the in general structure and colour, with following excep- male proctiger (figs. 71a, b). tions: connexiva more broadly marked with orange brown; metanotum and abdominal tergites II-IV with Biological notes small patches of silvery setae laterally, abdominal ter- Neusterensifer dentrecasteaux is found along the mar- gites V and VI entirely covered with silvery setae, ab- gins of lowland streams and rivers, often next to shad- dominal tergite VII with a very few silvery setae along ed banks along slowly flowing or sheltered side pools.

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150° 151° 152° 153° 154°

KIRIWINA IS. Neusterensifer TROBRIAND ISLANDS tufi

° 9° WOODLARK IS. 9

GOODENOUGH IS.

FERGUSSON IS. COLLINGWOOD BAY

D'ENTRECASTEAUX ISLANDS EGUM ATOLL SOLOMON SEA NORMANBY IS. NEW GUINEA GOODENOUGH BAY Neusterensifer 10° hunteri

ORANGERIE BAY Neusterensifer ENGINEER GROUP DEBOYNE IS. MISIMA IS. yela

LOUISIADE ARCHIPELAGO CONFLICT GROUP 11° 11°

ROSSEL IS.

TAGULA IS. CORAL SEA

150° 151° 152° 153° 154°

Fig. 74. Distribution of Neusterensifer species in the far eastern New Guinea region.

The type locality was a swift, clear river in a mostly 04.2”E, water temp. 24 °C., 24 August 2002, 09:00- unshaded bed of water-rounded rocks and cobbles, 13:00 hrs., CL 7180, D. A. & J. T. Polhemus (USNM); surrounded by gardens and disturbed lowland rain 2 macropterous females, 3 micropterous males, 1 mi- forest (fig. 86). The insects were found in sheltered cropterous female, Fergusson Island, east coast, upper side pools and water pockets amid cobbles along the Awaetowa River and rocky tribs., W. of Basima, 575- stream margins. 650 m. [1885-2130 ft.], 9°30’34.3”S, 150°48’17.3”E, water temp. 22 °C., 27 January 2003, 10:00-14:00 Neusterensifer kula sp. n. hrs., CL 7225, D. A. Polhemus (USNM); 8 micropter- (figs. 45, 64, 72) ous males, 7 micropterous females, Goodenough Is- land, Waidala Creek, 3 km. W. of Bolubolu on road Material examined. – Holotype, micropterous to Vivigani, 15 m. [50 ft.], 9°22’04.9”S, 150°21’ male: PAPUA NEW GUINEA, Milne Bay Prov., D’Entre- 14.3”E, water temp. 25 °C., 27 August 2002, 08:30- casteaux Islands, Fergusson Island, east coast, Yaya 10:00 hrs., CL 7188. D. A. & J. T. Polhemus (USNM); River, N. of Mebulibuli Point, 0-60 m. [0-200 ft.], 1 micropterous male, Goodenough Island, Kuliana 9°30’34.3”S, 150°52’51.1”E, water temp. 26 °C., 23 Creek at Ewaoiana, near Kalauna village, 290 m. [950 August 2002, 10:00-16:30 hrs., CL 7178, D. A. & J. ft.], 9°22’53.8”S, 150°19’29.6”E, water temp. 24.5 T. Polhemus (USNM). – Paratypes: PAPUA NEW °C., 26 August 2002, 10:00-12:00 hrs., CL 7185, D. GUINEA, Milne Bay Prov., D’Entrecasteaux Islands: 2 A. & J. T. Polhemus (USNM). macropterous females, 20 micropterous males, 28 mi- cropterous females, Fergusson Island, same data as Description holotype, D. A. & J. T. Polhemus (USNM, JTPC, Size. – Micropterous male, length 2.25-2.35 mm BPBM); 2 micropterous males, 1 micropterous female, (x– = 2.30, n = 6); width 0.90-1.05 mm (x– = 0.98, n = Fergusson Island, east coast, Mebulibuli Creek and 6). Micropterous female, length 2.75-2.80 mm (x– = rocky tributary, 1.3 km. upstream of mouth, S. of 2.79, n = 5); width 1.00-1.05 mm, (x– = 1.04, n = 5). Basima, 15-75 m. [50-250 ft.], 9°30’54.4”S, 150°52’ Macropterous female, length 2.75-2.80 mm (x– =

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2.78, n = 2); width 1.20 mm, (x– = 1.20, n = 2). tae gold in colour on ventrite II but becoming pro- Colour. – Micropterous male: Ground colour or- gressively darker and eventually blackish on ventrites ange brown, marked with darker brown on dorsal tho- III-VI, lateral sections of ventrites IV-VI to either side rax and abdomen, legs yellowish brown; small patches of medial sulcus bearing numerous moderately long, of short, shining, silvery pubescence present laterally inwardly-directed gold setae, lateral portions of ven- on metanotum and abdominal tergites II-VII, along trite VII to either side of sulcus not strongly tumes- anterior margin of pronotum, on lateral thorax and cent, but bearing a pair (1+1) of thick, roughly circu- abdomen. Head orange brown, darker along midline; lar tufts of short, stiff, inwardly-directed dark brown eyes dark red; rostrum pale whitish yellow, fuscous to black setae. Proctiger produced anteriorly into a medially, piceous distally. Pronotum orange brown, moderately long blade-like process with an expanded darker centrally and at punctations on posterior lobe, apex bearing a small terminal point (fig. 72). Para- bearing a pair (1+1) of faintly indicated elongate trans- meres vestigial or absent. verse yellowish spots to either side of midline behind Antennal formula I : II : III : IV; 0.40 : 0.30 : 0.42 anterior margin. Metanotum and abdomen orange : 0.60. brown dorsally, lateral and extreme posterior margins Proportions of legs as follows: Femur, tibia, tarsal of metanotum and abdominal tergites II-VII darker, 1, tarsal 2 of fore leg, 0.65 : 0.55 : 0.30 : 0.0; of mid- laterotergites below connexiva narrowly black along dle leg, 0.85 : 0.90 : 0.12 : 0.30; of hind leg, 1.00 : sutures and adjoining spiracles; connexiva entirely or- 1.10 : 0.17 : 0.30. ange brown. Antennae dark brown. Legs with coxae, Micropterous female: Similar to micropterous male trochanters and femora except at apices yellowish in general structure, but with following exceptions: brown, apical femora, tibiae and tarsi medium brown. overall coloration darker orange brown to rusty Structural characters. – Micropterous male: head brown, metanotum and abdominal tergites II-IV with of moderate length, declivant anteriorly, with weak patches of silvery setae laterally, abdominal tergites V impressed median line; length 0.50, width 0.60; and VI bearing silvery setae on posterior ⅔, abdomi- width of eye/interocular space, 0.12/0.40. Pronotum nal tergite VII produced into posteriorly-directed, fin- long, covering mesonotum, thickly set with obscure ger-like medial process, apex of this process projecting foveae, humeri depressed; length/width, 0.40/0.92. under and beyond posterior angles of connexival seg- Metanotum length 0.23. Abdominal tergites dull, ment VII and over base of tergite VIII and bearing a without shining areas; lengths of tergites II-VII, re- long, acuminate tuft of stiff black setae (fig. 45), ab- spectively: 0.10: 0.10 : 0.07 : 0.07 : 0.10 : 0.35. dominal tergite VIII with a few scattered silvery setae, Entire dorsum and laterotergites covered with fine this tergite plus proctiger lying in nearly vertical orien- appressed golden pubescence, intermixed with numer- tation; connexiva strongly and suddenly convergent ous long, slender, erect golden brown setae; legs and posteriorly, meeting over and covering base of abdom- antennae thickly clothed with short gold setae, scat- inal tergite VI plus all of tergite VII, connexival seg- tered long erect black setae present along anterior ment V with a few moderately long erect black setae margins of antennal segments I-III, anterior margins along dorsal margin, acuminate tufts of stiff black se- of all tibiae, posterior margin of middle femur; fore fe- tae arising on the inner margins of connexival seg- mur ventral margin weakly sinuate and biconcave, ments VII and projecting upward between the ap- bearing a small patch of stiff black setae basally plus pressed dorsal margins of these segments (fig. 45), numerous short, stiff, pale setae on distal ¾ , inter- posterolateral connexival angles of segment VII ob- mixed with evenly spaced comb of 10 long, erect, pi- tusely angulate, not produced; lateral sections of ab- lose golden setae; middle tibia bearing a row of evenly dominal ventrites curving upward and inward to bases spaced, long pilose golden setae of progressively de- of connexival segments, set with scattered moderately creasing length, longest and most basal of these setae long erect black setae, extreme lateral margins adjoin- with length approximately equal to ⅓ that of tibia; ing connexival bases narrowly black, overlain with a hind femur slightly incrassate, posterior margin bear- few silvery setae, black coloration also narrowly pre- ing numerous very short, stiff gold setae centrally. sent along all abdominal sutures; abdominal venter, Abdominal venter set with short appressed gold se- fore femur and hind femur unmodified. tae, and scattered much longer erect golden brown se- Macropterous female: Similar to micropterous fe- tae; abdominal ventrite II forming a small tumescence male in general structure and colour, with following centrally, this tumescence contiguous with a low cari- exceptions: posterior lobe of pronotum highly expand- na continuing posteriorly on midline of ventrite III, ed, brown, set with fine dark punctations, apex round- ventrites IV-VII depressed along longitudinal midline, ed, humeri weakly tumescent; wings complete, matte forming a narrow sulcus that becomes progressively dark brown, extending past tip of abdomen, forewing deeper posteriorly, ventrites II-VI with numerous bearing 4 closed cells, both basal cells with white long, inwardly- to posteriorly-directed setae, these se- streaks at basal angles, 2 distal cells lacking pale mark-

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150° 151° 152° 153° 154°

KIRIWINA IS. Neusterensifer TROBRIAND ISLANDS dentrecasteaux

° 9° WOODLARK IS. 9

GOODENOUGH IS.

FERGUSSON IS. COLLINGWOOD BAY D'ENTRECASTEAUX ISLANDS EGUM ATOLL

NORMANBY IS. SOLOMON SEA NEW GUINEA Neusterensifer ° misima 10

ORANGERIE BAY Neusterensifer ENGINEER GROUP DEBOYNE IS. MISIMA IS. sulcata

LOUISIADE ARCHIPELAGO CONFLICT GROUP 11° 11°

ROSSEL IS.

TAGULA IS. CORAL SEA

150° 151° 152° 153° 154°

Fig. 75. Distribution of Neusterensifer species in the far eastern New Guinea region. ings, costal margin bearing long, stiff, erect black setae faces of connexiva VII and projecting between the basally; abdominal tergite VII not forming a setiferous dorsal margins of the infolded connexiva themselves posterior projection as in micropterous form. (fig. 45). Females of N. kula may be distinguished Macropterous male: Unknown. from those of N. sulcata by the degree of posterior in- folding on the connexiva, which cover only the ex- Etymology treme posterior portion of abdominal tergite VI plus The name ‘kula’ refers to the ancient kula trading all of tergite VII, rather than the entirety of tergites VI ring that links the offshore islands of eastern New and VII as in N. sulcata, and by the absence of a small, Guinea. diffuse tuft of upright black setae at the posterior angle of connexival segment V (compare figs. 45, 46). Fe- Distribution males of N. kula may also be distinguished from those D’Entrecasteaux Islands (fig. 64). of N. tufi by possessing two separate upright, acumi- nate tufts of stiff black setae arising from the abdomen Comparative notes in lateral view, rather than a single tuft arising from Neusterensifer kula is an orange-brown species from the posterior prolongation of tergite VII, as in N. tufi Fergusson and Goodenough islands in the D’Entre- (compare figs. 45, 49). All three of these species under casteaux group that is the apparent sister species to N. discussion also share with N. muyuw, N, bowutu, and tufi from the Cape Nelson peninsula of mainland N. microrivula the common synapomorphy of a poste- New Guinea (see comparative notes under that riorly prolonged, finger-like projection on the posteri- species), and is also very similar in many respects to N. or margin of abdominal tergite VII, indicating that sulcata from Tagula Island in the Louisiades. As in N. they are all members of a single clade distributed on sulcata, females have a setiferous posteromedial projec- various sectors of the EPCT. tion on abdominal tergite VII that projects backward Males of N. kula may be distinguished from those below the apices of the infolded connexiva (fig. 45), of N. sulcata in having a longitudinal sulcus medially and a secondary acuminate tuft arising from the inner on abdominal ventrites IV-VII, rather than on ven-

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trites VI and VII; in having the pair of dark setae tufts pterous females, same data as holotype (USNM, JTPC, on abdominal ventrite VII roughly circular rather than BPBM). elongate-ovate; and in having the apex of the process on the proctiger expanded to form a “bird head” shape Description (fig. 72), rather than being tapering and acuminate. Size. – Micropterous male, length 2.30-2.35 mm The shape of the male proctiger process is instead very (x– = 2.33, n = 5); width 0.90-1.10 mm (x– = 1.01, n = similar to the state seen in N. tufi, but has a sharter 5). Micropterous female, length 2.50-2.60 mm (x– = apical prolongation beyond the expanded distal head, 2.55, n = 5); width 0.90-1.10 mm, (x– = 1.02, n = 5). and lacks small angulations on posterior margin of the Colour. – Micropterous male: Ground colour or- head (compare figs. 72, 73). ange brown, marked with darker brown on dorsal tho- In addition, the male fore femur of N. kula is weak- rax and abdomen, legs yellowish brown; small patches ly sinuate along its posterior margin and bears a patch of short, shining, silvery pubescence present laterally of stiff, slightly darkened setae basally. Abdominal on metanotum and abdominal tergites II-VII, along ventrites II-V also possess a medial patch of long hairs anterior margin of pronotum, on lateral thorax and that runs from the tumescent basal segments down abdomen. Head orange brown, darker along midline; into the anterior end of the medial sulcus; these hairs eyes dark red; rostrum pale whitish yellow, fuscous become darker as they enter the sulcus on the central medially, piceous distally. Pronotum orange brown, portions of ventrites IV and V. The male hind femur darker centrally and at punctations on posterior lobe, and tibia are not modified. In contrast to males of N. bearing a pair (1+1) of faintly indicated elongate trans- sulcata and N. femoralis, which have a similar general verse yellowish spots to either side of midline behind orange-brown appearance, the male hind femur in N. anterior margin. Metanotum and abdomen orange kula is not tumescent or otherwise modified. brown dorsally, lateral and extreme posterior margins of metanotum and abdominal tergites II-VII darker, Biological notes laterotergites below connexiva narrowly black along The type series was taken from small pools along sutures and adjoining spiracles; connexiva entirely or- the margins of rocky creeks draining the hilly eastern ange brown. Antennae dark brown. Legs with coxae, shore of Fergusson Island. For a further description of trochanters and femora except at apices yellowish the Yaya River type locality see the discussion under brown, apical femora, tibiae and tarsi medium brown. Rheovelia robinae. Structural characters. – Micropterous male: head of moderate length, declivant anteriorly, with weak Zoogeographic notes impressed median line; length 0.50, width 0.62; As currently understood, N. kula is confined to width of eye/interocular space, 0.12/0.37. Pronotum Goodenough and Fergusson islands in the D’Entre- long, covering mesonotum, thickly set with obscure casteaux group; these islands were formed by a group foveae, humeri depressed; length/width, 0.40/0.90. of metamorphic core complexes that emerged along Metanotum length 0.20. Abdominal tergites dull, the axis of the Woodlark Rift during the last 5 Myr without shining areas; lengths of tergites II-VII, re- (Abers et al. 2002). To date, N. kula has not been col- spectively: 0.17: 0.12 : 0.10 : 0.10 : 0.10 : 0.40. lected on Normanby Island, the third major island in Entire dorsum and laterotergites covered with fine the D’Entrecasteaux group; instead, this latter island appressed golden pubescence, intermixed with numer- harbors taxa such as N. hunteri and N. femoralis that are ous long, slender, erect dark brown setae; legs and an- also found at the eastern end of the adjacent Papuan tennae thickly clothed with short gold setae, scattered Peninsula. This pattern of distribution implies that long erect black setae present along anterior margins Normanby Island has had a separate faunal history of antennal segments I-III, anterior margins of all tib- from the remainder of the D’Entrecasteaux group, and iae, posterior margins of middle and hind femora; fore may have had an independent geological origin. femur ventral margin weakly sinuate and biconcave, bearing a small, dense patch of stiff black setae basally ¾ Neusterensifer tufi sp. n. plus numerous shorter, stiff, paler setae on distal , (figs. 38, 39, 50, 73, 74) intermixed with evenly spaced comb of 10 long, erect, pilose golden setae; middle tibia bearing a row of even- Material examined. – Holotype, micropterous ly spaced, long pilose golden setae of progressively de- male: PAPUA NEW GUINEA, Oro Prov., Kofure River creasing length, longest and most basal of these setae and cascading tribs., W. of Tufi, 0-45 m. [0-150 ft.], with length approximately equal to ⅓ that of tibia; 9°04’58.2”S, 149°16’38.7”E, water temp. 28 °C., 25 hind femur very slightly incrassate (fig. 38), posterior January 2004, 09:30-14:00 hrs., CL 7304, D. A. & J. margin bearing numerous very short, stiff gold setae T. Polhemus (USNM). – Paratypes: PAPUA NEW centrally. GUINEA, Oro Prov.: 19 micropterous males, 18 micro- Abdominal venter set with short appressed gold se-

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Figs. 76-77. Neusterensifer species, process arising from male proctiger. – 76 a-c, N. goilala: a, left later- al view, b, ventral view, c, right lateral view. 77 a, b, N. bowutu: a, ventral view, b, right lateral view.

76a 76b 76c

77b

77a

tae; abdominal ventrite II forming a prominent Proportions of legs as follows: Femur, tibia, tarsal domed tumescence centrally, this tumescence con- 1, tarsal 2 of fore leg, 0.62 : 0.60 : 0.28 : 0.0; of mid- tiguous with a well developed setiferous carina contin- dle leg, 0.80 : 0.80 : 0.15 : 0.30; of hind leg, 0.90 : uing posteriorly on midline of ventrites III-VI (fig. 1.10 : 0.15 : 0.30. 39), lateral portions of ventrites V and VI flanking this Micropterous female: Similar to micropterous male tumescence slightly depressed, forming a pair (1+1) of in general structure, but with following exceptions: shallow longitudinal sulci flanking the carina; ventrite overall coloration darker orange brown to rusty VII strongly depressed along longitudinal midline, brown, metanotum and abdominal tergites II-IV with forming a narrow sulcus that becomes progressively patches of silvery setae laterally, abdominal tergite V deeper posteriorly, lateral portions of ventrite VII to with silvery setae on posterior half, abdominal tergite either side of sulcus not strongly tumescent, but bear- VI bearing silvery setae on posterior ⅔, abdominal ter- ing a pair (1+1) of dense, roughly circular tufts of gite VII mostly hidden beneath infolded connexival short, stiff, posteriorly-directed dark brown to black apices, posterior margin produced medially into a pos- setae. Proctiger produced anteriorly into a moderately teriorly-directed, finger-like medial process bearing an long blade-like process with an expanded apex bearing acuminate tuft of long black setae at its tip (fig. 50), an extended terminal point (fig. 73). Parameres vesti- apex of this process projecting under and beyond pos- gial or absent. terior angles of connexival segment VII and over base Antennal formula I : II : III : IV; 0.38 : 0.27 : 0.40 of tergite VIII; abdominal tergite VIII with covered : 0.52. with numerous silvery setae except at extreme base,

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this tergite plus proctiger lying in nearly vertical orien- Nelson peninsula, indicated by similarities in the male tation; connexiva strongly convergent posteriorly, proctiger process and ventral abdominal structures, is meeting over and covering much of abdominal tergite of zoogeographic interest, because the presence of VII, connexival segments V-VII with a few moderate- these closely related species along the western exten- ly long erect black setae along outer margins but lack- sion of the Woodlark Rift supports the current geo- ing specialized hair tufts, posterolateral connexival an- logical hypothesis that the D’Entrecasteaux Islands gles of segment VII obtusely angulate, not produced; and the Tufi area have shared a common geological lateral sections of abdominal ventrites curving upward history linked by this structure (Abers et al. 2002). and inward to bases of connexival segments, set with scattered moderately long erect black setae, extreme Biological notes lateral margins adjoining connexival bases narrowly The type series was taken from a standing pool in black, overlain with silvery setae, black coloration also bedrock next to a cascade in a narrow, rock walled narrowly present along all abdominal sutures; abdom- gorge. inal venter, fore femur and hind femur unmodified. Macropterous form: Unknown. Neusterensifer bowutu sp. n. (figs. 51, 77a, 77b, 78) Etymology The name ‘tufi’ is a noun an apposition and refers Material examined. – Holotype, micropterous to the town of Tufi near the type locality. male: PAPUA NEW GUINEA, Morobe Prov., ‘Kerosine Creek’ and rocky trib., middle Aleater [= Alewiri] Riv- Distribution er basin, W. of Lababia village, 30 m. [100 ft.], Eastern New Guinea (fig. 74). 7°18’46.2”S, 147°06’30.1”E, water temp. 25 °C., 5 May 2003, 10:30-12:30 hrs., CL 7245, D. A. Polhe- Comparative notes mus (USNM). – Paratypes: PAPUA NEW GUINEA, Mo- Neusterensifer tufi is similar in many respects to N. robe Prov.: 14 micropterous males, 8 micropterous fe- kula, from which it can be distinguished by the more males, same data as holotype, D. A. Polhemus (USNM, highly developed and setiferous medial tumescence on JTPC, BPBM); 5 micropterous males, 6 micropterous fe- male abdominal ventrites II-VI (fig. 39), and the males, rocky hill stream behind Kamiali guesthouse, S. somewhat different shape of the process on the male of Lababia village, nr. Cape Dinga, 30 m. [100 ft.], proctiger (compare figs. 72, 73). In both species the 7°18’02.0”S, 147°08’01.4”E, CL 7243, water temp. proctiger process has an expanded apex of similar 25.5 °C., 4 May 2003, 14:00-15:00 hrs., D. A. Polhe- overall shape, but in N. tufi the extreme tip is more mus (USNM); 3 micropterous females, trib to lower elongate, and posterodorsal margin of the expanded Aleater [= Alewiri] River, 5 m. [15 ft.], 7°19’14.7”S, section is multidentate (Fig. 73). The female differs 147°07’34.9”E, 27 °C., 3 May 2003, 13:00-15:00 from that of N. kula in having the connexival margins hrs., CL 7241 D. A. Polhemus (USNM). not as closely appressed posteriorly, covering only the posterior half of abdominal tergite VII when viewed Description from above, rather than the posterior half of tergite VI Size. – Micropterous male, length 1.80-2.10 mm and all of tergite VII as in N. kula. The connexiva of (x– = 1.93, n = 3); width 0.75-0.80 mm (x– = 0.78, n = N. tufi also lack tufts of stiff black setae on the inner 3). Micropterous female, length 2.20-2.25 mm (x– = faces of the seventh connexival segments that project 2.23, n = 2); width 0.80-0.85 mm, (x– = 0.83, n = 2). upward between the appressed margins of the seg- Colour. – Micropterous male: Ground colour dark ments (fig. 50); such tufts are present in N. kula (fig. orange brown, marked with yellowish brown on ante- 45). By contrast, both N. tufi and N. kula do possess a rior pronotum and basal legs; patches of short, shin- posterior elongation of abdominal tergite VII into a ing, silvery pubescence present laterally on metan- finger-like structure with a tuft of long black setae at otum, abdominal tergites II, III, and V-VII, along its apex that projects posteriorly beneath the infolded anterior margin of pronotum, and on sides of thorax posterolateral angles of the connexiva and beyond ter- and abdomen. Head dark orange brown; eyes silvery gite VIII (figs. 45, 50); this character state allies N. tufi red; rostrum pale whitish yellow, fuscous medially, with other members of the ‘kula group’ of Neusteren- piceous distally. Pronotum orange brown, slightly sifer distributed throughout the EPCT. darker anteromedially, bearing a pair (1+1) of elongate transverse yellowish spots to either side of midline be- Zoogeographic notes hind anterior margin. Metanotum and abdomen The apparent sister species relationship of N. kula brown dorsally, laterotergites below connexiva black from Goodenough and Fergusson islands in the along sutures and adjoining spiracles; connexiva light D’Entrecasteaux group and N. tufi from the Cape orange brown. Antennae dark brown, becoming

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Neusterensifer aviavi

Neusterensifer bowutu

Neusterensifer hunteri

Fig. 78. Distribution of Neusterensifer species in the far eastern New Guinea region. blackish distally. Legs generally medium brown with longitudinal midline, this carina declining in height in tarsi darker, coxae, trochanters, basal femora and all posterior direction, flanked to either side by deep, nar- ventral surfaces pale yellowish white. row parallel sulci, these parallel sulci merging to form Structural characters. – Micropterous male: head of a single deep sulcus on central sections of ventrites VI moderate length, declivant anteriorly, with weak im- and VII, these two ventrites each with a pair (1+1) of pressed median line; length 0.40, width 0.60; width of dense tufts of short, stiff black setae lying to either side eye/interocular space, 0.15/0.30. Pronotum moder- of the medial sulcus. Proctiger produced anteriorly ately long, covering mesonotum, thickly set with ob- into a moderately long, tapering blade-like projection scure foveae, humeri depressed; length/width, with an acuminate apex (figs. 77a, b). Parameres vesti- 0.27/0.80. Metanotum length 0.20. Abdominal ter- gial or absent. gites dull, without shining areas; lengths of tergites II- Antennal formula I : II : III : IV; 0.30 : 0.25 : 0.35 VII, respectively: 0.15: 0.12 : 0.08 : 0.10 : 0.08 : 0.27. : 0.50. Entire dorsum and laterotergites covered with fine Proportions of legs as follows: Femur, tibia, tarsal appressed golden pubescence, intermixed with numer- 1, tarsal 2 of fore leg, 0.55 : 0.50 : 0.27 : 0.0; of mid- ous long, fine, slender, erect dark setae; legs and an- dle leg, 0.75 : 0.70 : 0.09 : 0.25; of hind leg, 0.75 : tennae thickly clothed with short gold setae, scattered 0.95 : 0.12 : 0.25. long erect black setae present along anterior margins Micropterous female: Similar to micropterous male of antennal segments I-III, anterior margins of all tib- in general structure, with following exceptions: overall iae, posterior margins of middle and hind femora; fore coloration somewhat darker, rusty orange brown, femur with ventral margin flattened and weakly con- strongly contrasting with pale orange brown connexi- cave on basal half, this area bearing numerous short, va; metanotum and abdominal tergites II and III with stiff gold setae, posterior margin also with evenly small patches of silvery setae laterally, abdominal ter- spaced comb of 6-8 long, erect, pilose golden setae; gites IV and V bearing silvery setae across entire middle tibia bearing a row of evenly spaced, long pi- widths on posterior ⅔, abdominal tergite VI hidden lose golden setae of progressively decreasing length, beneath infolded connexival margins, abdominal ter- longest and most basal of these setae with length ap- gite VII produced posteromedially to form a small, proximately equal to ⅓ that of tibia; hind femur very acuminate process extending beneath and slightly be- slightly incrassate, lacking tumescences or specialized yond the infolded posterior margins of the connexiva, setal patches. apex of this process bearing a posteriorly directed tuft Abdominal venter set with short appressed gold se- of long, stiff, black setae (fig. 51); abdominal tergite tae, and scattered longer erect golden setae; abdominal VIII with numerous silvery setae across entire width, ventrite II forming a small tumescence centrally, ven- this tergite and proctiger lying in nearly vertical orien- trites III-V forming a prominent median carina along tation; connexiva strongly and suddenly convergent

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posteriorly, meeting over and covering abdominal ter- were found along the margins of the small pools, but gite VI and most of VII except for tip of posteromedi- were not abundant. Another series was taken from a al projection, all connexival segments with a few mod- similar small, shaded, high gradient stream draining erately long erect black setae along dorsal margins, directly to the sea near Lababia, while a few addition- inner margins of connexival segments VII with acumi- al individuals were captured along the shaded, under- nate tuft of stiff black setae that projects upward and cut margins of a swift but much lower gradient low- between the infolded connexival margins, posterolat- land stream entering the terminal reach of the Aleater eral connexival apices of segment VII forming small River at the point where it exited the mountains near obtuse angles; lateral sections of abdominal ventrites its mouth at Sachsen Bay. curving upward and inward to bases of connexival segments, set with scattered moderately long erect Neusterensifer goilala sp. n. black setae, extreme lateral margins adjoining connex- (figs. 36, 44, 56, 76a-c, 82) ival bases diffusely blackish, overlain with patches of silvery setae, black coloration also narrowly present Material examined. – Holotype, micropterous along all abdominal sutures; abdominal venter, fore fe- male: PAPUA NEW GUINEA, Central Prov., Owen Stan- mur, and hind femur unmodified. ley Range, small hill streamlet, 1.3 km. SSE of Macropterous form: Unknown. Woitape, 1600 m. [5250 ft.], 8°32’59.8”S, 147°15’ 34.3”E, water temp. 17.5 °C., 2 October 2003, Etymology 08:30-09:15 hrs., CL 7248, D. A. Polhemus (USNM). The name ‘bowutu’ is a noun in apposition and – Paratypes: PAPUA NEW GUINEA, Central Prov.: 1 mi- refers to the Bowutu Mountains, from which this cropterous male, 7 micropterous females, same data as species was collected. holotype (BPBM); 1 micropterous male, Owen Stanley Range, Om Creek, trib. to upper Udabe River, 1.85 Distribution km. SSE of Woitape, 1585 m. [5200 ft.], 8°33’ Eastern New Guinea (fig. 78). 27.7”S, 147°15’21.8”E, water temp. 19 °C., 2 Octo- ber 2003, 12:00-12:30 hrs., CL 7251, D. A. Polhe- Comparative notes mus (USNM); 4 micropterous males, Owen Stanley Neusterensifer bowutu is a small, rusty brown species Range, tributary streamlet to Hane River, 1.0 km. SSE occurring in the Bowutu Mountains of the northern of Fane, 1310-1370 m. [4300-4500 ft.], 8°33’32.7”S, Papuan Peninsula (fig. 78). Females share with those of 147°05’23.7”E, water temp. 20 °C., 4 October 2003, N. yela from Rossel Island, N. hunteri from the Milne 11:00-12:30 hrs., CL 7256, D. A. Polhemus (USNM); Bay area, and N. microrivula from the Lakekamu Basin 1 macropterous female, 8 micropterous males, 9 mi- a tuft of stiff black setae on the inner margin of female cropterous females, Owen Stanley Range, trib. to Mas connexival segment VII that projects upward between River, 1.9 km. SW of Fane, 1220-1250 m. [4000- the infolded connexival margins. Females of N. bowu- 4100 ft.], 8°33’33.0”S, 147°04’12.2”E, water temp. tu also possess the posteromedial prolongation of ab- 20.5 °C., 5 October 2003, 10:45-12:30 hrs., CL dominal tergite VII seen in these and several other 7258, D. A. Polhemus (USNM, JTPC, BPBM). species, with this tergite forming a small, setiferous posterior tumescence that extends slightly beyond the Description posterior apices of the infolded connexiva (fig. 51). Size. – Micropterous male, length 2.85-3.00 mm The males are similar to those of N. yela and N. sulcata (x– = 2.92, n = 6); width 1.00-1.20 mm (x– = 1.08, n = from the Louisiades and N. microrivula in having a 6). Micropterous female, length 3.10-3.25 mm (x– = deep, V-shaped sulcus along the midline of the ventral 3.18, n = 6); width 1.10-1.20 mm, (x– = 1.15, n = 6). abdomen, but may be easily separated by the shape of Macropterous female, length 3.25 (n = 1), maximum the process on the proctiger (figs. 77a, b). As with N. width (across humeral angles of pronotum) 2.35 mm microrivula, these character states indicate that N. (n = 1). bowutu is yet another member of a monophyletic Neus- Colour. – Micropterous male: Ground colour dark terensifer lineage scattered across the various sectors of orange brown, marked with lighter orange brown on the East Papua Composite Terrane. anterior pronotum and connexiva; patches of short, shining, silvery pubescence present laterally on metan- Biological notes otum, abdominal tergites II, III, VI and VII, along an- The type locality was a small, high gradient stream terior margin of pronotum, and on sides of thorax and descending out hills covered by dense primary rain abdomen. Head dark orange brown; eyes silvery red; forest, with a bed profile consisting of alternating rostrum pale whitish yellow, fuscous medially, piceous rocky riffles, small shallow pools, and small cascades distally. Pronotum orange brown, slightly darker me- over dark volcanic bedrock. Individuals of N. bowutu dially, bearing a pair (1+1) of elongate transverse yel-

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Figs. 79-81. Neusterensifer species, process arising from male proctiger. – 79, N. lubu Polhemus & Pol- hemus, ventral view; 80, N. microrivula, ventral view; 81, N. aviavi, ventral view.

79 81

80

lowish spots to either side of midline behind anterior VII, respectively: 0.22: 0.20 : 0.15 : 0.15 : 0.15 : 0.47. margin. Metanotum and abdomen brown dorsally, Entire dorsum and laterotergites covered with fine laterotergites below connexiva black along sutures and appressed golden pubescence, intermixed with scat- adjoining spiracles; connexiva medium orange brown. tered medium length, semi-erect golden setae; legs and Antennae dark orange brown, paler ventrally. Legs antennae thickly clothed with short gold setae, scat- generally pale brown with tarsi darker; coxae, tered long erect golden setae present along anterior trochanters, basal femora and all ventral surfaces pale margins of antennal segments I-III, anterior margins of yellowish; metasternum matte black, ventral head and all tibiae, posterior margins of middle and hind femo- abdomen prodominantly orange brown. ra; fore femur with ventral margin weakly concave cen- Structural characters. – Micropterous male: head of trally, this area bearing numerous short, stiff dark gold- moderate length, declivant anteriorly, with weak im- en setae, posterior margin also with evenly spaced pressed median line; length 0.37, width 0.67; width of comb of 6-8 long, erect, pilose golden setae; middle eye/interocular space, 0.15/0.45. Pronotum moder- tibia bearing a row of evenly spaced, long pilose golden ately long, covering mesonotum, thickly set with ob- setae of progressively decreasing length, longest and scure foveae, humeri depressed; length/width, most basal of these setae with length approximately 0.50/1.00. Metanotum length 0.30. Abdominal ter- equal to ⅓ that of tibia; hind femur tumescent on cen- gites dull, without shining areas; lengths of tergites II- tral portion of posterior margin, this tumescence bear-

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ing numerous short, stiff, dark setae (fig. 36). mus & Polhemus, 1988, fig. 9), followed by two more Venter with metasternum broadly angulate on pos- larger and elongate cells in filling most of the central terior margin; abdominal venter set with short ap- portion of the wing lying laterally to the Cu vein, then pressed gold setae, and scattered longer erect golden se- two very small cells distal to the outermost of the two tae; abdominal ventrite II forming a small tumescence middle cells; wing coloration dull black, with elon- centrally with small tuft of erect, posteriorly directed gate, creamy white streak in outer basal cell and very gold setae at apex, ventrites III-V forming a prominent small creamy white patches centrally in each of the median carina along longitudinal midline (fig. 44), this two middle cells. carina descending from tumescence on ventrite II and Macropterous male: Unknown. declining in height in posterior direction, flanked to ei- ther side by deep, narrow parallel sulci, these parallel Etymology sulci merging to form a single deep, nearly hair-free The name ‘goilala’ is a noun an apposition and sulcus on central sections of ventrites VI and VII, ven- refers to the highland people who inhabit the region trite VII bearing a pair (1+1) of large, strongly raised surrounding the type locality. tumescences lying on either side of the medial sulcus, these tumescences bearing dense tufts of short, stiff Distribution black setae. Proctiger produced anteriorly into a mod- Eastern New Guinea (fig. 82). erately long, projection with a blunt, hooked tip (figs. 76a-c). Parameres vestigial or absent. Comparative notes Antennal formula I : II : III : IV; 0.50 : 0.30 : 0.45 Neusterensifer goilala is one of the largest known : 0.55. species in the genus, with body lengths in both males Proportions of legs as follows: Femur, tibia, tarsal and females exceeding 2.90 mm, and occurs at eleva- 1, tarsal 2 of fore leg, 0.75 : 0.70 : 0.35 : 0.0; of mid- tions above 1200 m in the Owen Stanley Range of the dle leg, 1.10 : 1.00 : 0.20 : 0.30; of hind leg, 1.25 : Papuan Peninsula (fig. 82). Females may be recog- 1.40 : 0.20 : 0.30. nized by the strongly convergent and appressed con- Micropterous female: Similar to micropterous male nexival margins, which cover part or all of tergites VI- in general structure and coloration, with following ex- VIII, forming a keel-like structure with a broadly ceptions: metanotum and abdominal tergites II and obtuse posterior angle when viewed laterally with its III with small patches of silvery setae laterally, abdom- apex at the suture between segments VI and VII (fig. inal tergite IV with narrow band of silvery setae across 56). Males may be easily recognized by the structure of entire widths on posterior margin, abdominal tergite the process on the proctiger, with its blunt, hooked tip V forming an elongate triangle set between strongly (figs. 76a-c); the pair of convergent sulci on abdomi- converging connexiva, abdominal tergites VIII hidden nal ventrites III-V that merge into a single deep sulcus beneath appressed connexival margins, proctiger lying on ventrites VI and VII, flanked by prominent setifer- in nearly vertical orientation, thickly covered with ous tumescences on the lateral sections of ventrite VII short silvery setae; connexiva strongly and suddenly (fig. 44); by the slightly rising medial carina on ab- convergent posteriorly, meeting over and covering ab- dominal ventrite VI with a small patch of dark setae at dominal tergites VI-VIII, connexival segment VII the apex, which extends anteriorly so as to nearly bi- bearing a fringe of stiff, black, posteriorly-directed se- sect ventrites III-V (fig. 44); and by the broad ventral tae along posterior margin when viewed laterally (fig. tumescence on the central section of the hind femur 56), posterolateral connexival apices of segments VI (fig. 36). and VII forming a broad, keel-like obtuse angle when The ventral abdominal modifications of the male viewed laterally (fig. 56); lateral sections of abdominal of N. goilala are similar in many respects to those seen ventrites curving upward and inward to bases of con- in N. bowutu from the northern coast of the Papuan nexival segments, predominantly black, overlain with Peninsula, except that in the latter species there are extensive patches of silvery setae; abdominal venter, setiferous tumescences flanking the central longitudi- fore femur, and hind femur unmodified. nal sulcus on both ventrites VI and VII, rather than Macropterous female: Similar to micropterous form only on VII as seen in N. goilala (fig. 44), and there is in general structure and coloration but with pronotum no anteriorly produced medial carina on ventrite VI. enlarged, length/width = 1.10/1.30, humeral angles Neusterensifer bowutu also lacks the tumescent male well developed, prominent, posterior lobe well pro- hind femur found in N. goilala (fig. 36); this tumes- duced, almost entirely covering scutellum, apex form- cent state is instead similar to that seen in other ing a rounded angle; wings fully developed, bearing 6 species found further east in the Papuan Peninsula closed cells, consisting of two long, narrow cells basal- and on adjacent archipelagos, including N. femoralis ly between the Sc and An veins and separated by the (fig. 35), N. sulcata, and N. muyuw (fig. 37). combined R+M+Cu vein (terminology follows Polhe-

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Biological notes long, covering mesonotum, thickly set with obscure This species was taken on small, shallow, first order foveae, humeri depressed; length/width, 0.30/0.80. spring-fed streamlets shaded by primary montane rain Metanotum length 0.20. Abdominal tergites dull, forest at elevations above 1200 m. in the central Owen without shining areas; lengths of tergites II-VII, re- Stanley Range. The orange-brown coloration of the spectively: 0.10: 0.10 : 0.08 : 0.08 : 0.07 : 0.25. insects closely matches the laterite substrate of these Entire dorsum and laterotergites covered with fine clear streamlets, rendering the insects cryptic on the appressed golden pubescence, intermixed with numer- water surface. ous long, fine, slender, erect golden setae; legs and an- tennae thickly clothed with short gold setae, scattered Neusterensifer microrivula sp. n. long erect black setae present along anterior margins (figs. 52, 80, 82) of antennal segments I-III, anterior margins of all tib- iae, posterior margins of middle and hind femora; fore Material examined. – Holotype, micropterous femur ventral margin very weakly sinuate and bicon- male: PAPUA NEW GUINEA, Gulf Prov., rocky tributary cave, bearing a small patch of short, stiff black setae to Sapoi River, upstream of Ivimka Research Station, near base, posterior margin also with evenly spaced 120-150 m. [395-490 ft.], 7°43’36”S, 146°29’59”E, comb of 6-8 long, erect, pilose golden setae; middle water temp. 24.5 °C., 16 January 2001, 07:00-13:00 tibia bearing a row of evenly spaced, long pilose gold- hrs., CL 7148, D. A. Polhemus (USNM). – Paratypes: en setae of progressively decreasing length, longest and PAPUA NEW GUINEA: 9 micropterous males, 12 mi- most basal of these setae with length approximately cropterous females, Gulf Prov., same data as holotype, equal to ⅓ that of tibia; hind femur unmodified, lack- D. A. Polhemus (USNM, JTPC, BPBM); 1 micropterous ing tumescences or specialized setal patches. male, 2 micropterous females, Central Prov., small Abdominal venter set with short appressed gold se- tributary streamlet to Yaniwe River at Tekadu, 300 m. tae, and scattered longer erect golden setae; abdominal [985 ft.], 7°40’36”S, 146°33’05”E, water temp. 24 ventrite II forming a small conical tumescence central- °C., 21 January 2001, 12:00-16:30 hrs., CL 7157, D. ly, ventrite III with a low median carina along longi- A. Polhemus (USNM, BPBM). tudinal midline flanked to either side by shallow par- allel sulci, central sections of ventrites IV-VII strongly Description depressed, forming a deep longitudinal sulcus on un- Size. – Micropterous male, length 1.95-2.05 mm derside of abdomen; ventrites VI and VII each with a (x– = 2.02, n = 6); width 0.75-0.85 mm (x– = 0.82, n = pair (1+1) of dense tufts of short, stiff black setae to ei- 6). Micropterous female, length 2.35-2.40 mm (x– = ther side of medial sulcus. Proctiger produced anteri- 2.41, n = 5); width 0.90-0.95 mm, (x– = 0.92, n = 5). orly into a moderately short, stout, tapering blade-like Colour. – Micropterous male: Ground colour or- projection with a weakly bifurcate apex (fig. 80). Para- ange brown, marked with yellowish brown on anteri- meres vestigial or absent. or pronotum and basal legs; patches of short, shining, Antennal formula I : II : III : IV; 0.40 : 0.25 : 0.42 silvery pubescence present laterally on metanotum, : 0.49. abdominal tergites II, III, and V-VII, along anterior Proportions of legs as follows: Femur, tibia, tarsal margin of pronotum, and on sides of thorax and ab- 1, tarsal 2 of fore leg, 0.60 : 0.50 : 0.30 : 0.0; of mid- domen. Head orange brown; eyes dark red; rostrum dle leg, 0.70 : 0.70 : 0.12 : 0.25; of hind leg, 0.90 : pale whitish yellow, fuscous medially, piceous distally. 1.00 : 0.13 : 0.25. Pronotum orange brown, slightly darker anteromedi- Micropterous female: Similar to micropterous male ally, bearing a pair (1+1) of elongate transverse yellow- in general structure, with following exceptions: overall ish spots to either side of midline behind anterior mar- coloration somewhat darker, rusty orange brown, gin. Metanotum and abdomen orange brown dorsally, strongly contrasting with pale orange brown connexi- lateral and extreme posterior margins of metanotum va; metanotum and abdominal tergites II-IV with and all abdominal tergites narrowly dark brown to small patches of silvery setae laterally, abdominal ter- blackish, laterotergites below connexiva black along gite V bearing silvery setae across entire width on pos- sutures and adjoining spiracles; connexiva entirely terior ⅔, abdominal tergite VII produced medially into light orange brown. Antennae medium brown. Legs a small, thumb-like projection bearing a tuft of elon- generally medium brown with tarsi darker, coxae, gate, stiff black setae that projects posteriorly below the trochanters, basal femora and all ventral surfaces pale infolded margins of connexival segment VII (fig. 52), yellowish white. abdominal tergite VIII with numerous scattered silvery Structural characters. – Micropterous male: head setae across entire width, tergite VIII and proctiger ly- of moderate length, declivant anteriorly, with weak ing in nearly vertical orientation; connexiva strongly impressed median line; length 0.40, width 0.57; and suddenly convergent posteriorly, meeting over and width of eye/interocular space, 0.16/0.35. Pronotum covering abdominal tergites VI-VII plus basal portion

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of VIII, all connexival segments with a few moderately small, shallow pool adjacent to the main channel, and long erect black setae along dorsal margins, a brushy were otherwise uncommon along the remainder of the tuft of stiff black setae arising on the inner margins of stream. Another series was taken from a small headwa- connexival segments VII and projecting upward be- ter streamlet at Tekadu, nearly 300 m. higher in eleva- tween the appressed dorsal margins of the segments, tion in the same Aviavi River catchment. At the latter posterolateral connexival apices of segment VII form- locality, the insects occurred along pool margins in a ing small right angles; lateral sections of abdominal small, shaded channel draining hill slopes covered ventrites curving upward and inward to bases of con- with lightly disturbed premontane rain forest. nexival segments, set with scattered moderately long erect black setae, extreme lateral margins adjoining Neusterensifer aviavi sp. n. connexival bases diffusely blackish, overlain with (figs. 59, 78, 81) patches of silvery setae, black coloration also narrowly present along all abdominal sutures; abdominal venter, Material examined. – Holotype, micropterous fore femur, and hind femur unmodified. male: PAPUA NEW GUINEA, Gulf Prov., rocky tributary Macropterous form: Unknown. to Sapoi River, upstream of Ivimka Research Station, 120-150 m. [395-490 ft.], 7°43’36”S, 146°29’59”E, Etymology water temp. 24.5 °C., 16 January 2001, 07:00-13:00 The name ‘microrivula’ comes from the Latin ‘mi- hrs., CL 7148, D. A. Polhemus (USNM). – Paratypes: cros’, or very small, and ‘rivulo’, or small stream, re- PAPUA NEW GUINEA, Gulf Prov.: 5 micropterous males, ferring to this species’ preference for small headwater 2 micropterous females, same data as holotype, D. A. streams in the New Guinea mountains. Polhemus (USNM, JTPC, BPBM).

Distribution Description Eastern New Guinea (fig. 82). Size. – Micropterous male, length 2.15-2.30 mm (x– = 2.22, n = 5); width 0.85-0.90 mm (x– = 0.86, n = Comparative notes 5). Micropterous female, length 2.30-2.50 mm (x– = Neusterensifer microrivula is a small, orange-brown 2.40, n = 3); width 1.00-1.05 mm, (x– = 1.03, n = 3). species occurring along the southern flanks of the far Colour. – Micropterous male: Ground colour dark western Owen Stanley Range, in the upper Lakekamu brown, marked with orange brown on anterior prono- River basin. Females are similar to those of N. yela tum and connexiva; patches of short, shining, silvery from Rossel Island and N. kula from the D’Entre- pubescence present laterally on metanotum and ab- casteaux Islands in possessing a tuft of stiff black setae dominal tergites II-IV and VII, on posterior halves of on the inner connexival margins of segment VII that tergites V and VI, broadly across central portion of projects posteriorly between the infolded margins of tergite VIII, along anterior margin of pronotum, and the segment (fig. 52), although in the case of N. mi- on sides of thorax and abdomen. Head dark brown; crorivula this tuft is brushy rather than acuminate. It is eyes blackish red; rostrum pale whitish yellow, fuscous also similar to the foregoing species in having a pos- medially, piceous distally. Pronotum dark brown, be- teromedial prolongation of female abdominal tergite coming orange brown anterolaterally, bearing a pair VII (fig. 52), but in N. microrivula this projection is (1+1) of elongate transverse orange brown spots to ei- incipient and shorter than that seen in other similarly ther side of midline behind anterior margin. Metan- modified Neusterensifer species. Males are similar to otum and abdominal tergites dark brown, lateroter- those of N. yela and N. sulcata, the latter from Tagula gites broadly blackish below connexiva, connexiva Island, in having the male ventral abdomen deeply orange brown on outer halves and ventrally. Antennae sulcate along the longitudinal midline, but may be medium brown. Legs generally medium brown, with easily separated by the shape of the process on the coxae, trochanters, basal femora and all ventral sur- male proctiger (fig. 80). On the basis of these charac- faces pale yellowish to yellowish brown. ter states, N. microrivula is hypothesized to be a mem- Structural characters. – Micropterous male: head ber of the same clade of orange-brown species that is of moderate length, declivant anteriorly, with weak widely spread across the various fragments of the East impressed median line; length 0.40, width 0.60; Papua Composite Terrane. width of eye/interocular space, 0.15/0.37. Pronotum long, covering mesonotum, thickly set with obscure Biological notes foveae, humeri depressed; length/width, 0.40/0.85. The type locality was a small, swift, rocky tributary Metanotum length 0.17. Abdominal tergites dull, to the Aviavi River, flowing through a bed of rocks without shining areas; lengths of tergites II-VII, re- and cobbles in a catchment shaded by primary pre- spectively: 0.12: 0.10 : 0.10 : 0.10 : 0.08 : 0.30. montane rain forest. This insects were found on a Entire dorsum and laterotergites covered with fine

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Neusterensifer microrivula

Neusterensifer goilala

Neusterensifer lubu

Fig. 82. Distribution of Neusterensifer species in the far eastern New Guinea region. appressed golden pubescence, intermixed with numer- Proportions of legs as follows: Femur, tibia, tarsal ous short, erect golden setae; legs and antennae thick- 1, tarsal 2 of fore leg, 0.60 : 0.52 : 0.22 : 0.0; of mid- ly clothed with short pale setae, scattered long erect dle leg, 0.75 : 0.75 : 0.17 : 0.27; of hind leg, 0.90 : black setae present along anterior margins of antennal 1.07 : 0.20 : 0.30. segments I-III, anterior margins of all tibiae, posterior Micropterous female: Similar to micropterous male margins of middle and hind femora; fore femur ven- in general structure and colour, with following excep- tral margin straight, not modified, bearing numerous tions: metanotum and abdominal tergites II and III short, erect pale setae along entire length, intermixed with small patches of silvery setae laterally, tergite IV with evenly spaced comb of 6-8 long, erect, pilose pale covered with silvery setae for its entire width on poste- setae; middle tibia bearing a row of evenly spaced, long rior half, tergites V and VI almost entirely covered pilose pale setae of progressively decreasing length, with silvery setae across their entire widths, tergite VII longest and most basal of these setae with length ap- lacking silvery setae, abdominal tergite VIII bearing a proximately equal to ⅓ that of tibia; hind femur with few scattered silvery setae centrally, this tergite plus numerous short, erect pale setae along posterior mar- proctiger lying 45° from vertical when viewed lateral- gin, lacking tumescences or specialized setal tufts. ly; connexiva broadly separated, bowing outward, Abdominal venter set with short appressed gold se- margins gradually convergent posteriorly, posterolat- tae, and scattered longer pale setae; abdominal ven- eral angles of segments VII produced into stout, angu- trites I-IV unmodified; ventrite V shallowly sulcate late projections (fig. 59) with apices angling inward centrally along longitudinal midline; ventrite VI over lateral sections of abdominal tergite VII, margins sharply raised posteromedially to form a pair (1+1) of of all connexival segments bearing scattered long, angulate tumescences slightly separated by a shallow erect, black setae but lacking specialized setal tufts; lat- sulcus along longitudinal midline, apices of these eral sections of abdominal ventrites curving upward to tumescences black, bearing sparse tufts of moderately bases of connexival segments, set with numerous long long, fine, gold setae; ventrite VII broadly and trans- erect black setae, lateral margins adjoining connexival versely flattened across central section, bearing a pair bases broadly blackish overlain with silvery setae; ab- (1+1) of low longitudinal tumescences separated by a dominal venter, fore femur, hind femur unmodified. shallow sulcus along longitudinal midline, these Macropterous form: Unknown. tumescences contiguous with bases of paired angulate tumescences on ventrite VI. Proctiger produced ante- Etymology riorly into a moderately long, acuminate process (fig. The name ‘aviavi’ is a noun in apposition and refers 81). Parameres vestigial or absent. to the Aviavi River, to which the Sapoi River at the Antennal formula I : II : III : IV; 0.37 : 0.25 : 0.42 type locality is a tributary. : 0.56.

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Distribution ACKNOWLEDGEMENTS Eastern New Guinea (fig. 78). This project would not have been possible without Comparative notes the help of numerous individuals in Papua New Neusterensifer aviavi may be recognized by its gener- Guinea. We wish in particular to thank Gai Kula, ally dark coloration, the presence of paired angulate David Mitchell, Bena Sena and Neal Lima of Conser- processes centrally on male abdominal ventrite VI, the vation International’s Melanesia Program for facilitat- moderately long, slender and acuminate process on ing surveys in Milne Bay Province, which led to the the male proctiger (fig. 81), and the angulate, infolded discovery of many of the new species described herein. projections of the female connexival apices (fig. 59). We also thank Andy Mack of the Wildlife Conserva- tion Society for his generous invitation for the first au- Biological notes thor to visit the Ivimka Field Station in the Lakekamu This species occurred on small pockets of water Basin and his wonderful hospitality during that trip; amid cobbles bordering a rocky stream in the hills Davidson Yariyari of Kofure village, who coordinated surrounding the Lakekamu Basin. For further de- our field work in the Tufi area; John Sengo and Karol scription of this locality see the discussion under N. Kisokau of the Village Development Trust, who made microrivula. possible the field work in the Kamiali area of the Bowutu Mountains; and Jim Amiato and Bulisa Ino- Neusterensifer lubu D. Polhemus & J. Polhemus va of the Papua New Guinea National Museum and (figs. 60, 79, 82) Gallery, who provided field assistance and village lia- sons in the Owen Stanley Range. Neusterensifer lubu D. Polhemus & J. Polhemus 2000a: 115. In addition, we wish to express our deepest appreci- ation to Wayne and Lee Thompson and their crew on Material examined. – PAPUA NEW GUINEA, Gulf the research vessel Marlin 1, which served as our float- Prov.: 1 micropterous male, 2 micropterous females, ing laboratory during the Kula Ring 1, Kula Ring 2, pools along Sapoi River, upstream of Ivimka Research and Kula Ring 3 expeditions. Without their excellent Station, 120-150 m. [395-490 ft.], 7°43’36”S, seamanship and knowledge of local conditions we 146°29’59”E, water temp. 24.5 °C., 16 January 2001, could never have reached many of the remote islands 07:00-13:00 hrs., CL 7148, D. A. Polhemus (USNM). lying east of New Guinea that are treated in this re- port. Similarly, we wish to thank Gerry Allen of Con- Distributional notes servation International and Ron Englund of the Bish- This species was originally described from the Lubu op Museum for their tireless assistance and excellent River at Omo, in the premontane foreland of the companionship in the field as part of the Melanesia Kikori River basin. The new records from the Lakeka- freshwater survey program. mu River basin extend the species range considerably This research was conducted in association with Bi- to the east along the southern flanks of the New ological Surveys and Inventories grant DEB-0103794 Guinea central mountains, and indicate that N. lubu from the National Science Foundation, Washington, will also be found in the Purari River catchment as D.C. to the Bishop Museum, with field work in Papua well. This same distribution pattern is shared by the New Guinea supported by the Smithsonian Institu- trepobatine gerrid Stygiobates mubi J. Polhemus & D. tion’s Drake Fund and by a grant for Melanesian fresh- Polhemus (Polhemus & Polhemus 2002). water surveys from Conservation International. In ad- dition, JTP conducted this research as an adjunct Biological notes faculty member in the Dept. of Entomology at Colora- The Lakekamu series was taken from shaded, par- do State University. We thank all these organizations tially vegetated vertical banks bordering a long, quiet for their continued support of our research into the sys- side pool 1.0-1.5 m. deep along the midreach of the tematics and zoogeography of aquatic Heteroptera. Aviavi River, a large, very swift, rocky stream in an open, mostly unshaded channel draining the south REFERENCES flanks of the western Owen Stanley Range. This local- ity was very near the small, more heavily shaded tribu- Abers, G. A., A. Ferris, M. Craig, H. Davies, A. L. Lerner- tary stream from which the type series of N. aviavi and Lam, J. C. Mutter & B. Taylor, 2002. Mantle compensa- tion of active metamorphic core complexes at Woodlark N. microrivula were taken, but the three species ap- rift in Papua New Guinea. – Nature 418: 862-865. peared to adopt a strict habitat segregation, with the Bain, J. H. C., H. L. Davies, P. D. Hohnen, R. J. Ryburn, I. former two only occurring along the tributary hill E. Smith, R. Grainger, R. J. Tingey & M. R. Moffat. stream, and N. lubu occurring only along the main 1972. Geology of Papua New Guinea. .- Bureau of Min- river channel. eral Resources Australia, Canberra, Geology and Geo-

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83 84

85 86 Figs. 83-86. Habitats of Microveliinae in eastern New Guinea and nearby islands. – 83, Woodlark Island, rocky stream drain- ing to Suloga Harbor; type locality for Neusterensifer muyuw. 84, Basilaki Island, waterfall on a tributary to the Guiagoila Riv- er. Wet faces next to the falls are habitat for Rheovelia basilaki, while pool margins support Neusterensifer hunteri; 85, Seeps and small cascades on a small tributary to the Hane River, near Fane, Owen Stanley Range; type locality for Aegilipsicola auga and paratype locality for Rheovelia truncata. 86, Midreach of Awaetowa River near Basima, Fergusson Island, D’Entrecasteaux group; sheltered pocket water along the river margins is habitat for Neusterensifer dentrecasteaux.

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