Rhagovelia novacaledonica group

This species group was originally proposed by J. Polhemus & D. Polhemus (1988) to hold R. novacaledonica Lundblad from New Caledonia and R. biroi Lundblad from eastern New Guinea, with several other species from New Caledonia and the southern Philippines provisionally assigned. Based on a subsequent comparative analysis of male paramere morphology, Zettel (1994) and D. Polhemus (1995) formally proposed the placement of the following additional species in this group: R. pidaxa J. Polhemus & Herring from New Caledonia; R. werneri Hungerford & Matsuda and R. hoogstraali Hungerford & Matsuda from Mindanao; and R. thysanotos Lansbury, R. crinita Lansbury, R. aureospicata Lansbury and R. herzogensis Lansbury from New Guinea. Subsequently, D. Polhemus & J. Polhemus (2011) added another species, R. batantana, from the Raja Ampat Islands, and also provisionally assigned another new species from this archipelago, R. rajana from Batanta Island, to the group.

The R. novacaledonica group is characterized by the short pronotum in wingless forms that leaves the posterior and lateral portions of the mesonotum well exposed (Figs. 15, 35, 41, 42, 48,49, 55, 56, 63, 64, 70, 71, 77, 78, 85, 86, 99, 100, 106, 107, 109, 118, 119, 125, 126, 133, 134, 140, 141, 146, 147, 154, 155, 161, 162, 169, 170, 171, 172, 181, 182, 189, 190, 194, 195); the large, elongate male paramere with setiferation on the basal half ventrally, and the distal section often expanded and usually forming a vertical flange apically (Figs. 38, 45, 52, 59, 67, 74, 81, 89, 103, 114, 122, 129, 137, 143, 150, 158, 165, 176, 178, 185, 192, 198); the male proctiger with prominent basolateral lobes, the distolateral lobes being only incipiently developed or absent, and a large, tapering distal cone with an angular apex (Figs. 39, 46, 53, 60, 68, 75, 82, 90, 104, 115, 123, 130, 138, 144, 151, 159, 166, 177, 179, 186, 193, 199); paired dorsal abdominal carinae in winged forms that are short, reaching at most onto abdominal tergite II; and the forewing with four closed cells, the two posterior cells having apices extending into the distal third of the wing (Fig. 10).

As currently interpreted, the R. novacaledonica group occupies an elongate geographic range along the western margin of the Pacific Plate, from New Caledonia northwestward through New Guinea to the southern Philippines (D. Polhemus 1995). Based on species described at the time of their work, J. Polhemus & D. Polhemus (1988) stated that “The few members of the novacaledonica group in New Guinea are presently known only from the north coast ranges and the Papuan Peninsula…By contrast, the papuensis group and other species groups derived from it have radiated extensively in the younger and more rugged central ranges…” This interpretation has proven incorrect on several levels. First, based current knowledge derived from far more extensive collections over the intervening 30 years, the majority of the rich Rhagovelia fauna occurring on New Guinea and adjacent islands at elevations above 500 m is composed of members of the R. novacaledonica group, which is the most speciose group on the island. Second, the accreted terranes of the New Guinea central ranges are latest Cretaceous to Eocene in age, and thus significantly older than the Oligocene to Miocene terranes of the northern coastal ranges (D. Polhemus 2007).

In the EPCT, members of the R. novacaledonica group are typical inhabitats of shaded, rocky upland streams (Figs. 40, 47, 54, 61, 83, 98, 105, 116, 117, 124, 131, 139, 145, 160, 167, 180, 187, 200, 226, 233, 239, 265, 273, 280, 289, 302, 308, 329). Many species seem to be restricted to small first-order and second-order tributaries in catchment headwaters, with some species being found only in the final several hundred meters of a stream immediately below source springs. As a result, this group exhibits a marked degree of geographically circumscribed local endemism. Species of this group also have a degree of cold tolerance and occur at some of the highest localities occupied by any Gerromorpha in New Guinea, being found on streamlets up to 2600 m. elevation in the central highlands of the island.

The R. novacaledonica group is the largest and most species-rich group of Rhagovelia occurring in the EPCT. Within this assemblage a number of distinct subgroups can be recognized, which may potentially represent monophyletic clades within the larger group, although this has not yet been phylogenetically tested. The species included in each of these subgroups are as follows:

Rhagovelia tufi subgroup: R. bowutu n. sp., R. obscura n. sp., R. tufi n. sp., and R. upalai n. sp.

Rhagovelia goilala subgroup: R. goilala n. sp., R. udabe n. sp., and R. antap n. sp.

Rhagovelia dibuwa subgroup: R. awaetowa n. sp., R. dibuwa n. sp., R. kalawai n. sp., R. peninsularis n. sp., R. torrenticola n. sp., and R. watuti n. sp.

Rhagovelia auga subgroup: R. auga n. sp.

Rhagovelia peggiae subgroup: R. peggiae Kirkaldy and R. hirsuta Lansbury.

Rhagovelia kolukou subgroup: R. kolukolu n. sp. and R. mbo n. sp.

Species not assigned to a subgroup (treated as incertae sedis within the R. novacaledonica group): R. aviavi n. sp., R. tekadu n. sp., R. mimani n. sp., and R. sapoi n. sp.

In addition, R. aureospicata Lansbury is now removed from the R. novacaledonica group and considered incertae sedis in the context of the overall New Guinea Rhagovelia assemblage, based on its male paramere morphology (Fig. 335) and the unusual armature of the male middle femur (Fig. 333)

These subgroups are discussed in greater detail within the species treatments following the key, and also noted within the key in those instances where they fall out discretely within the couplet sequence.

Key to the Species of the Rhagovelia novacaledonica Species Group occurring in the EPCT

Note: This key relies on both male and female characters, so a series containing both properly associated sexes is necessary for definitive identification. In the absence of one sex or the other, the figures provided, in association with geographic provenance, will also in many cases be sufficient to obtain an identification.

1. Female connexival margins not meeting at any point along their length (although the posterolateral angles may be convergent and inwardly folded to some degree), leaving all abdominal tergites visible along their central sections to some extent when viewed from above (although lateral margins of certain tergites may be covered) (Figs. 36, 42, 49, 56, 64, 71, 78, 86, 93, 100, 107, 109, 119, 126, 170, 172, 189, 195).....................................................................2

- Female connexival margins appressed or folded inward and meeting along some portion of their length, such that one or more abdominal tergites are covered across their central sections when viewed from above (Figs. 134, 141, 147, 155, 162, 182). 20

2. Black denticles present on inner prosternum and adjacent proacetabula, sometimes also extending onto the jugae and lower propleural area (Fig. 32).................................................................................3

- Black denticles not present on the prosternum, proacetabula, jugae or lower propleurae.............................. 7

3. Smaller species, body lengths 3.0– 4.1 mm; female connexival margins with posterolateral angles often bending weakly inward over lateral portions of abdominal tergites VII or VIII, but never touching or completely covering an underlying tergite, although the hair tufts on the posterolateral angles may touch (Figs. 36, 42, 49, 56); male paramere ventrally bilobate (Figs. 38, 45, 52, 59); coloration predominantly black, with transverse orange-brown patch on anterior pronotum behind vertex discrete, bordered laterally with black and not confluent with orange-brown coloration on propleurae (Figs. 35, 36, 41, 42, 48, 49, 55, 56).................................................................................. R. tufi subgroup…4

- Larger species, body length ranging from 4.1–4.7 mm; female connexival margins straight, evenly convergent posteriorly, not inwardly folded to any degree, orientation either vertical or angling outward, leaving all abdominal tergites completely visible from above (Fig. 189); male paramere not ventrally bilobate, instead with distal arm parallel-sided and apically truncate, angled upward before apex when viewed laterally (Fig. 192); orange-brown coloration on anterior pronotum not forming a discrete transverse patch but instead broadly confluent laterally with orange-brown coloration on propleurae (Figs. 188, 189)................................................................................................. R. sapoi n. sp.

4. Male abdominal ventrite VII with a broad, shallow, longitudinally ovate depression centrally, lateral margins of this depression not strongly raised or demarcated with dense bands of setae (Figs. 16, 17); length of female antennal segment I less than 2X the length of segment II.....................................................................................5

- Male abdominal ventrite VII strongly depressed centrally, this depression bounded laterally by a pair (1+1) of raised, setiferous longitudinal carinae (Figs. 18, 19); length of female antennal segment I equal to 2X the length of segment II..............6

5 Female abdominal tergites III–V with a raised longitudinal medial carina, this carina most prominent on tergite III, becoming progressively less well expressed on tergites IV and V; male abdominal tergite VII with shiny chestnut brown patch posteromedially; male genitalia as in Figs. 38, 39; Cape Nelson area of northeastern Papuan Peninsula (Fig. 62).............................................................................................. Rhagovelia tufi n. sp .

- Female abdominal tergites III–V without a raised longitudinal medial carina; male abdominal tergite VII shining black; male genitalia as in Figs. 45, 46; Bowutu Mountains of northern Papuan Peninsula (Fig. 62)........... Rhagovelia bowutu n. sp.

6. Shining chestnut brown patches present centrally on female abdominal tergites V–VIII; male abdominal ventrite VII bearing deep, crescent-shaped depression centrally, this depressed area becoming broader and deeper posteriorly, lateral margins flanked by setiferous arcuate ridges (Fig. 18); male genitalia as in Figs. 52, 53; southern Papuan Peninsula (Fig. 62)............................................................................................. Rhagovelia obscura n. sp.

- Shining patches absent on female abdominal tergites V–VIII, these tergites uniformly dull black; male abdominal ventrite VII with longitudinally rectangular depression centrally, lateral margins flanking this depression strongly raised, these raised margins bearing dense pads of short, dark setae posteriorly (Fig. 19); male genitalia as in Figs. 59, 60; far eastern terminus of Papuan Peninsula (Fig. 62)............................................................ Rhagovelia upalai n. sp.

7. Female connexival margins strongly convergent adjacent to abdominal tergites I–III, then parallel adjacent to tergites IV and V before converging again adjacent to tergites VI–VIII (Fig. 170)............................... Rhagovelia aviavi n. sp.

- Female connexival margins straight or at most very weakly sinuate, tapering evenly toward posterior margin of abdomen, widely separated throughout their lengths and not touching, angling slightly outward; all female abdominal tergites completely visible from above (Figs. 64, 71, 78, 86, 100, 107, 109, 119, 126, 172, 195)....................................... 8

8. Ground color orange-brown (Figs. 63, 64, 70, 71, 77, 78, 109).................................................. 9

- Ground color black to dark blackish-grey (Figs. 85, 86, 92, 93, 99, 100, 106, 107, 118, 119, 125, 126, 171, 172, 194, 195).. 13

9. Female mesonotum entirely orange-brown (Figs. 63, 64); female abdominaltergite VII with width exceeding length; male paramere with deep notch on dorsal margin, apex expanded and truncate (Fig. 67); eastern Papuan Peninsula (Fig. 84)....................................................................................... Rhagovelia antap n. sp.

- Female mesonotum predominantly brown, with orange-brown coloration on central portion only, if present at all (Figs. 70, 71, 77, 78, 109); female abdominal tergite VII with length exceeding width; male paramere lacking deep notch on dorsal margin, apex not expanded and broadly truncate, instead rounded and often produced slightly upward to varying degrees (Figs. 74, 81, 115, 129)........................................................................................... 10

10. Pale coloration on anterior pronotum consisting of a broad, orange-brown band extending laterally onto pleural region (Fig. 70, 71, 77, 78); dorsum densely covered with short, appressed golden setae; length of pronotum along longitudinal midline subequal to length of head; apex of male paramere distally truncate (Figs. 74, 81); larger species than below, body lengths ranging from 4.1–4.8 mm; Owen Stanley Range above 1000 m elevation (Fig. 84)…............................... 11

- Pale coloration on anterior pronotum confined to a transverse dark yellow patch immediately behind head, this coloration not extending laterally onto pleural region (Fig. 85, 86, 92, 93, 99, 100, 106–109, 118, 119, 125, 126); dorsum bearing only scattered short golden setae; length of pronotum along longitudinal midline clearly less than length of head; male paramere slightly transversely expanded apically, with dorsal margin forming a rounded, upward-angled lobe (Fig. 89, 96, 103, 114, 122, 129); body lengths ranging from 3.2–4.2 mm; D’Entrecaseteaux Islands (Fig. 132)................................ 12

11. Female connexiva with posterolateral angles bearing prominent posteriorly-directed tufts of black setae (Fig. 71); posterior margin of female pronotum weakly anteriorly concave centrally (Fig. 71); male paramere with apex broadly rounded and truncate, ventral margin of paramere broadly concave (Fig. 74)............................... Rhagovelia udabe n. sp.

- Female connexiva with posterolateral angles lacking prominent posteriorly-directed setal tufts (Fig. 78); posterior margin of female pronotum weakly posteriorly convex centrally (Fig. 78); male paramere with apex forming a small, upwardly-directed angle, ventral margin of paramere nearly straight, only weakly concave (Fig. 81)................ Rhagovelia goilala n. sp.

12. Male proctiger with basolateral lobes very broad, width of proctiger subequal to length including basal struts (Fig. 130); large species, body length 4.0– 4.2 mm; Goodenough Island (Fig. 132)......................... Rhagovelia torrenticola n. sp.

- Male proctiger with basolateral lobes not as broad as above, width of proctiger distinctly less than length, even excluding basal struts (Fig. 115); medium-sized species, body length 3.4–3.7 mm; Normanby Island (Fig. 132)..... Rhagovelia dibuwa n. sp.

13. Male paramere with distal arm straight, narrow, parallel sided, apex truncate (Fig. 198); female abdominal tergites bearing only a single small, irregular shining patch posteromedially on tergite VII........................ Rhagovelia mimani n. sp.

- Male paramere much thicker, distal arm not slender or parallel-sided, produced into an upward-angling, rounded distal lobe (Figs. 89, 96, 103, 114, 122, 129); female abdominal tergites bearing shining patches on multiple tergites in addition to tergite VII............................................................................... R. dibuwa subgroup...14

14. Female dorsal abdomen with shining patches present on tergites V–VIII......................................... 15

- Female dorsal abdomen with shining patches less extensive, absent on tergite V, and sometimes also absent on tergite VI.. 18

15. Posterolateral angles of female connexiva lacking tapering setal tufts when viewed laterally, bearing only a few scattered setae (Fig. 21); male paramere strongly expanded distally, with a pointed, upwardly-angling apex (Fig. 96); far eastern terminus of Papuan Peninsula (Fig. 132)...................................................... Rhagovelia peninsularis n. sp.

- Posterolateral angles of female connexiva bearing tapering setal tufts to some degree when viewed laterally (Figs. 20, 22–25); male paramere only moderately expanded distally, with a broadly rounded, upwardly-angling apex (Fig. 89, 103, 114, 122, 129); D’Entrecasteaux Islands, China Strait islands, or Cloudy Mountains (Fig. 132)............................... 16

16. Female with shining patches present on abdominal tergites VII and VIII only; angled distal process on male paramere elongate, length when measured along posterior margin about 2 times the width of the central paramere shaft (Fig. 89); Cloudy Mountains (Fig. 132)........................................................................ Rhagovelia watuti n. sp.

- Female with shining patches present on abdominal tergites VI–VIII at least, sometimes also present on tergite V; distal process on male paramere shorter than above, length when measured along posterior margin distinctly less than 2 times the width of the central paramere shaft (Fig. 103, 114, 122, 129); D'Entrecastaux Islands and China Strait Islands (Fig. 132)............. 17

17. Ratio of midline length of pronotum to midline length of head in males 1.5 or greater; base of male proctiger with a pair of well-developed, raised carinae (Figs. 104, 115), these carina triangular and sharply pointed when viewed laterally (Fig. 5); posterior margin of female abdominal ventrite VII may be produced into a posteriorly-directed, triangular process medially (Fig. 29); smaller species, body length 3.7 mm or less............................................................... 18

- Ratio of midline length of pronotum to midline length of head in males 1.35 or less; base of male proctiger with a pair of weakly developed, raised carinae (Figs. 123, 130), these carina low and obtuse when viewed laterally; posterior margin of female abdominal ventrite VII not produced into a posteriorly-directed, triangular process medially, at most bearing a slightly rounded medial projection (Figs. 30, 31); larger species, body length 4.0 mm or greater.....................................19

18. Female with shining patches present on abdominal tergites VI–VIII only; posterior margin of male abdominal ventrite VII with shallow, angular indentation centrally (Fig. 28); ground coloration dark grey with limited orange-brown markings (Fig. 99, 100), forms with orange-brown ground coloration not currently known; Sideia and Basilaki Islands (Fig. 132)........................................................................................... .. Rhagovelia kalawai n. sp.

- Female with shining patches present on abdominal tergites V–VIII; posterior margin of male abdominal ventrite VII more produced and acute, lacking a shallow, angular indentation centrally (Fig. 29); ground coloration polymorphic, varying from predominantly dark grey to predominantly orange-brown (Figs. 107, 109, and see couplet 9 above); Normanby Island (Fig. 132)............................................................................. Rhagovelia dibuwa n. sp.

19. Ground coloration polymorphic, varying from predominantly dark grey (Figs. 125, 126) to predominantly orange-brown (and see couplet 9 above); male proctiger with basolateral lobes very broad, width of proctiger subequal to length including basal struts (Fig. 130); male paramere with central section of apical lobe possessing a small translucent patch (Fig. 129) resembling a window; Goodenough Island (Fig. 132).......................................... .. Rhagovelia torrenticola n. sp.

- Ground coloration dark grey with limited orange-brown markings (Figs. 118, 119), forms with orange-brown ground coloration not currently known; male proctiger with basolateral lobes not as broad as above, width of proctiger less than length, even excluding basal struts (Fig. 123); male paramere with central section of apical lobe opaque, lacking translucent patch (Fig. 122); Fergusson Island (Fig. 132)....................................................... Rhagovelia awaetowa n. sp.

20. Female connexiva strongly infolded, closely convergent or appressed over the tops of abdominal tergites V–VIII, covering all or significant portions of tergites V–VIII when viewed from above (Figs. 141, 147, 155, 162)........................ 21

- Female connexiva not as strongly infolded as above, convergent posteriorly, touching at or near posterior apices, covering portions of abdominaltergites VII or VIII (Figs. 134, 182).................................................... 22

21. Male fore tibia concave and bearing a curving brush of setae distally, appearing spatulate (Fig. 133); large species, body length 4.4–4.9 mm; coloration predominantly black with limited dark orange markings (Figs. 133, 134)......................................................................................... R. auga subgroup… Rhagovelia auga n. sp.

- Male fore tibia slender throughout, not expanded and bearing a brush of setae distally; smaller species, body length 3.2–4.0 mm; coloration dark reddish brown with extensive yellow and orange markings (Figs. 181, 182).... Rhagovelia tekadu n. sp.

22. Anterior portion of pronotum bearing a transversely ovate dark orange or yellowish patch in the area behind the head vertex (Figs. 140, 141, 146, 147), this patch bordered laterally by areas of dark coloration overlain with prunosity, which extend laterally on the propeura, intergrading to brown on the proacetabula; male paramere distally expanded to varying degrees, not slender and tapering (Figs. 143, 150); male proctiger with prominent, laterally projecting basolateral lobes (Figs. 144, 151); New Guinea (Fig. 153)............................................................. .. R. peggiae subgroup...23

- Anterior portion of pronotum with broad, dark yellow transverse band behind head (Figs. 154, 155, 161, 162) that is confluent laterally with similar yellowish coloration extending laterally and ventrally to cover the propelura and proacetabula; male paramere slender and tapering (Figs. 158, 165); male proctiger with basolateral lobes absent or at most incipiently developed (Figs. 159, 166); Louisiade Archipelago (Fig. 168)........................................ R. kolukolu subgroup...24

23. Small black denticles present on prosternum adjacent to rostrum, and on inner portions of proacetabula; male paramere distinctively produced to an upturned quadrate process distally (Fig. 143); southern Papuan Peninsula (Fig. 153)........................................................................................ Rhagovelia peggiae Kirkaldy

- Prosternum and inner proacetabula lacking small black denticles; male paramere slender, with apex acutely pointed, ventral margin gently bilobate (Fig. 150); northern central Papuan Peninsula (Fig. 153).............. Rhagovelia hirsuta Lansbury

24. Males with abdominal ventrite VI broadly and shallowly depressed; apex of male paramere hooked slightly downward (Fig. 158); posterolateral angles of female connexiva lacking tufts of stiff black setae (Fig. 155); Tagula Island (Fig. 168)....................................................................................... Rhagovelia kolukolu n. sp.

- Males with both abdominal ventrite VI and VII broadly and shallowly depressed centrally; apex of male paramere hooked slightly upward (Fig. 165); posterolateral angles of female connexiva bearing bushy tufts of stiff black setae (Fig. 162); Rossel Island (Fig. 168).................................................................... .. Rhagovelia mbo n. sp.