Research Article |
Corresponding author: Alice Wells ( alice.wells@environment.gov.au ) Academic editor: Ralph Holzenthal
© 2015 Alice Wells, Kjell Arne Johanson.
This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Citation:
Wells A, Johanson KA (2015) Review of New Caledonian species of Oxyethira Eaton, with description of 17 new species, and new records for Hydroptila Dalman and Hellyethira Neboiss (Trichoptera, Hydroptilidae). ZooKeys 530: 37-90. https://doi.org/10.3897/zookeys.530.6047
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New Caledonian representation of the cosmopolitan genus Oxyethira Eaton is reviewed, with the description of new species bringing to 26 the total for the genus on the island. The species are referred to three subgenera: Trichoglene Neboiss (11 species), Pacificotrichia Kelley (13 species) and Dampfitrichia Ulmer (one species) and one species is unplaced to subgenus. A key is provided to Oxyethira species of New Caledonia. In addition, new records are given for two otherwise Australian species, Hydroptila losida Mosely and Hellyethira malleoforma Wells. Points marked on a series of small maps of New Caledonia indicate the site or sites at which the species were collected. This final paper in a series of generic revisions brings the hydroptilid fauna of the island of New Caledonia to 60 species, distributed in six genera.
Spicipalpia, Hydroptilidae , New Caledonia, endemic, key, new species
The cosmopolitan genus Oxyethira Eaton exhibits a diverse array of male genital structures and arrangements. The species show some variability in female terminalia, but exhibit extreme conservatism in larval and case morphology. Representatives of the genus found in New Caledonia appear to be no exception. This is apparent upon consideration of the variability among 26 species recorded here, and the similarity of the considerable number of unassociated final instar larvae collected at many sites by
This final paper in a series of genus-level reviews (
Oxyethira is well represented in New Caledonia, at genus level only surpassed in known species diversity among the island Trichoptera by the helicopsychid genus Helicopsyche (
Most of the 17 species newly described here from New Caledonian can be referred to the above three subgenera with a degree of confidence. One species, however, cannot be placed at present: O. macropennis sp. n. shares the diagnostic features of Oxyethira as defined by
Among females in the samples at least two general morphological forms can be recognised in abdominal terminalia: a short oviscapt of the form illustrated by
Apparent distributions of species are difficult to interpret (see Figs
Most of the material this study is based upon was collected in light traps and Malaise traps situated near or across running water. Specimens were prepared for study as Canada balsam slide mounts following maceration in KOH and clearing in clove oil. Male genitalia are illustrated in line drawings, traced from draft figures using Adobe Illustrator CS5, for species for which suitable slides are available.
An identification key and descriptions of New Caledonian Oxyethira species are provided, as well as brief diagnoses of previously described Oxyethira species and new illustrations of their male genitalia, drawn from types and/or newly collected non-type specimens. Species descriptions are based on male genital features, although identification of homologies among these is often difficult, especially for some of the more aberrant species. Usually diagnostic features are indicated on figures. Terms applied to genital structures follow the recommendations of
Development of a useful key for easy identification of species was difficult as observation of most readily diagnostic features requires preparation of slide mounts of specimens and examination under a compound microscope.
Collection sites for species were plotted on a series of maps (Figs
MNHP
The chief diagnostic characteristics that
In discussions of subgenus Trichoglene (
Three species groups are recognised among these New Caledonian members of subgenus Trichoglene. A set of species, the spinifera-group, with abdominal segment IX subquadrate comprises O. spinifera sp. n., O. tiwaka sp. n., O. perignonica sp. n., and O. abbreviata sp. n. A second set, the caledoniensis-group, with venter of abdominal segment IX in ventral view produced anteriorly, proximally either rounded or tapered and somewhat triangular, includes O. caledoniensis Kelley, O. incurvata sp. n., O. arok Oláh & Johanson, O. amieu sp. n., and O. houailou sp. n. The distinctions between these two groups of species, however, are not clear cut. The venter of O. caledoniensis Kelley is only slightly produced anteriorly. A third group, the insularis-group, characterised by the prominently Y-shaped gonopods has only two members: O. insularis sp. n. and O. parinsularis sp. n.
Males closely resemble those of O. (T.) tiwaka sp. n. and O. (T.) perignonica sp. n. in having abdominal segment IX subquadrate in ventral view, with the apico-ventral margin truncate but both those species have recognisable gonopods, albeit strongly reduced, whereas as in O. (T.) arok, O. (T.) amieu and O. (T.) spinifera gonopods are so severely reduced that no gonopods can be identified. The rods of ventral processes of O. (T.) spinifera are slender and spiny, rather sharper than in O. (T.) tiwaka sp. n. but in both species the rods are almost parallel whereas in O. (T.) perignonica sp. n. they are sharply pointed and convergent.
Male antennae with 23–25 flagellomeres, flagellomeres rectangular in profile, without sensilla placodea, terminal 3 flagellomeres pale, rest dark; anterior wing length 1.3–1.5 mm (n=8); tibial spurs 0,3,4. Female antennae with 18 flagellomeres, flagellomeres all dark; anterior wing length 1.3–1.6 mm (n=4); tibial spurs 0,3,4. Abdominal sternite VII with sharp median spur.
Male, genitalia (Figs
Oxyethira species male genitalia. 1–3 O. spinifera sp. n., ventral and lateral views, and phallic apparatus 4–6 O. tiwaka sp. n., ventral view, phallic apparatus and lateral view 7–9 O. perignonica sp. n., ventral view, phallic apparatus and lateral view 10–12 O. abbreviata sp. n., ventral view, phallic apparatus and lateral view. Abbreviations: gon = gonopod; r sp = rod of subgenital process; st lb = setose lobe of subgenital process; VIII, IX = abdominal segments VIII and IX.
Holotype. Male (on slide), New Caledonia, small fall ~10 km SW Houailou on Houailou-Bourail road, 26.xii.1998, leg. A. Wells (MNHP).
Paratypes. 2 males (on slides), small stream ~15 km SW Houailou on Houailou–Bourail road, 26.xii.1998, leg. A. Wells (
12 males, 6 females, Province Sud, Sarraméa, 2907 m, stony forest stream, loc 13, 21°37.097'S 165°49.351'E, Malaise trap, 18–21.xi.2001, leg. K.A. Johanson, T. Pape & B. Viklund (
Name spinifera is descriptive of the sharp spines of the subgenital processes.
This species appears to be quite widespread from the far south towards the north of the island (Fig.
Readily recognised by the short, blunt, darkly sclerotised peg-like gonopods but in other respects showing very close resemblance to O. (T.) spinifera which has gonopods reduced so severely that they can be recognised as only small convexities on the apico-ventral margin of segment IX. Also similar to O. (T.) perignonica sp. n. in having abdominal segment IX subquadrate, but that species has the gonopods situated laterally, and curved mesally, and the subgenital processes in the form of convergent, rather than parallel, spines.
Male antennae with 18–19 flagellomeres, flagellomeres all dark, without sensilla placodea, each flagellomere about as long as wide; anterior wing length 1.4–1.7 mm (n=8); tibial spurs 0,3,4; abdominal sternite VII with small sharp median spur.
Male, genitalia (Figs
Holotype. Male, (on slide), New Caledonia, Province Nord, Bouérabate Stream, S Mont Ninndo, along road Barabache–Boulagoma, 20°17.409'S, 164°11.242'E, 60 m, 19.xii.2003–7.i.2004, Malaise trap, loc#089, leg. K.A. Johanson (MNHP).
Paratypes. 1 male (on slide), data as for holotype, (
Named for the river beside which one of the specimens was collected.
Oxyethira tiwaka was collected quite commonly in the southern region, but at only two disjunct localities in the north (Fig.
Male is distinguished from O. incurvata sp. n. which also has the gonopods situated laterally and mesally directed although in O. perignonica they are more spur-like, and the rods of the ventral processes are sharply pointed and convergent.
Male antennae with 17–18 flagellomeres, flagellomeres without sensilla placodea, each flagellomere about 1.5 X longer than wide; anterior wing length 1.4–1.7 mm (n=2); tibial spurs 0,2,4; abdominal sternite VII without median spur.
Male, genitalia (Figs
Holotype. Male (on slide), New Caledonia, Province Sud, stream draining to Marais de la Rivière Blanche, 5 km SW Pont Pérignon, 22°09.513'S, 166°39.942'E, 180 m, 6–16.xi.2003, Malaise trap, loc#011, leg. K.A. Johanson (MNHP).
Paratypes. 2 males (1 on slide), Province Sud, W part of Plaine des lacs, 150 m downstream bridge at La Capture, 22°15.967'S, 166°49.493'E, 261 m, 04–22.xi.2003, Malaise trap, loc#007, leg. K.A. Johanson (
Named for the bridge on the river near where the holotype was collected.
Taken only at several sites in the south of the island (Fig.
Most closely similar to O. perignonica and O. tiwaka, all 3 having males with abdominal segment IX quadrate in ventral view. But O. abbreviata sp. n. is distinguished by having the ventral processes distally rounded and with a short sharp spine angled proximally compared with elongate convergent spines of O. perignonica, and elongate parallel spines of O. tiwaka.
Male antennae with 18–19 flagellomeres, flagellomeres rectangular in profile, sensilla placodea absent; anterior wing length 1.4–1.7 mm (n=4); tibial spurs 0,3,4; abdominal sternite VII with small sharp medial spine on distal margin.
Male, genitalia (Figs
Holotype. Male (on slide), New Caledonia, Province Sud, Monts des Koghis, ca 800 m S Koghi Restaurant, 22.18447°S, 166.50315°E, 400 m, 11–26.xi.2003, Malaise trap, loc. 23, leg. K.A. Johanson (MNHP).
Paratypes. 2 males, Province Nord, Mt Aoupinié, 354 m, stream, loc. 17, light trap, 24.xi.2001, leg. K.A. Johanson, T. Pape & B. Viklund (
abbreviata, named for the very abbreviated male genital structures.
This species was collected from several quite central sites (Fig.
Males resemble superficially those of O. perignonica both having mesally directed, laterally situated gonopods, but in ventral view these are more slender than those of O. perignonica and the rods of the subgenital processes are aligned in parallel with the distal margins of sternite IX in contrast to O. perignonica in which the they form sharp spines angled obliquely.
Male antennae with 20–25 flagellomeres, flagellomeres with few sensilla placodea, each rectangular in profile and 1.5–2× longer than wide; anterior wing length 1.2–1.5 mm (n=8); tibial spurs 0,3,4; abdominal sternite VII without median spur.
Male, genitalia (Figs
Oxyethira species male genitalia. 13–15 O. incurvata sp. n., ventral view, phallic apparatus and lateral view. 16–18 O. caledoniensis Kelley: 16 ventral view of holotype specimen drawn from holotype 17, 18 ventral and later al views of variant form 19–22 O. arok Oláh & Johanson: 19 ventral view drawn from paratype specimen 20–22 ventral and lateral views and phallic apparatus of variant form. Abbreviations: gon = gonopod; r sp = rod of subgenital process; st lb = setose lobe of subgenital process; VIII, IX = abdominal segments VIII and IX.
Holotype. Male, New Caledonia, Province Nord, Mt Panié, 20.57306°S, 164.77139°E, 902 m, 9.xii.2003, Malaise trap, loc#075, leg. K.A. Johanson (MNHP).
Paratypes. 4 males (2 on slides), Province Sud., Rivière Bleue, 282 m, stony river, loc 4, 22°05.705'S, 166°38.225'E, Malaise trap, 13–16.xi.2001, leg. K.A. Johanson, T. Pape & B. Viklund (
The name incurvata is descriptive of the orientation of the gonopods.
From the collecting records the distribution of this species appears to be highly disjunct (Fig.
Oxyethira caledoniensis Kelley, 1989: 196, figs 33, 42, 56.
A typical member of subgenus Trichoglene, with males superficially difficult to distinguish from other closely similar species such as O. abbreviata, O. arok and O. spinifera. Neither O. arok nor O. spinifera sp. n. has recognisable gonopods whereas both O. caledoniensis and O. abbreviata sp. n. have widely separated, shallowly dome-shaped gonopods; O. abbreviata sp. n. has very short, sharply pointed subgenital processes that are directed anteriorly and abdominal segment IX quadrate in ventral view, compared with the rod-shaped dorsal processes of O. caledoniensis and sub-triangular to shield-shaped abdominal segment IX.
Antennae: male with 19–24 flagellomeres, all dark; flagellomeres rectangular in profile, without sensilla placodea; female with 15 flagellomeres. Anterior wing length: male 1.4–1.9 mm (n=12); female 1.4–1.7 mm (n= 8). Tibial spurs 0,3,4. Abdominal segment VII with mid-ventral sharp spine.
Holotype. Male, New Caledonia, Plum, (
Delineation of this species among the large collection before us proved difficult, with only a very few specimens conforming closely to the holotype. In describing the species,
Oxyethira arok Oláh & Johanson, 2010a: 91, figs 42–44.
In general appearance of male genitalia O. arok is closest to O. caledoniensis but appears to have gonopods reduced completely, which feature it shares with O. amieu sp. n. and O. spinifera; however, O. arok has abdominal segment IX almost parallel-sided in distal half, rather than tapered as in O. amieu sp. n., and O. spinifera has the rods of subgenital processes tapered to acute apices compared with the blunt apices of O. arok.
Antennae: male with 20–21 flagellomeres; flagellomeres quadrate to slightly rectangular in profile; female with 17 flagellomeres, terminal 2 pale, rest dark. Anterior wing length: male 1.2–1.8 mm (n=10); female 1.4–1.6 mm (n =10). Tibial spurs 0,3,4. Abdominal sternite VII with small median spur offset from distal margin. Female with length of abdominal segment IX almost twice width, distal margin, with a sclerotised margin, mesally produced distally. Segment X stouter at base than apex, gradually tapered distally, truncate apically.
Paratype. 1 male, New Caledonia, Province Sud, Monts Kwa Ne Mwa, on road between Nouméa andYaté, 2.0 km E Pic Mouirange, 22°12.356'S, 166°40.798'E, 220 m, 16–30.xi.2003, Malaise trap, loc#014, leg. K.A. Johanson. Other material. 5 males, Province Nord, Mt Aoupinié, 354 m, stream, loc 17, light trap, 24.xi.2001, leg. K.A. Johanson, T. Pape & B. Viklund (
Specimens here identified as O. arok show some variability in proportions of abdominal segment IX and in male genital structures as apparent in Figures
Oxyethira arok has been collected from disjunct localities in the far south and central part of the island (Fig.
Males are similar to O. arok and O. houailou sp. n. in the shape of abdominal segment IX, which in ventral view is strongly tapered and more or less triangular proximally, but both O. amieu sp. n. and O. houailou sp. n. also taper distally, while O. arok is more or less parallel-sided in distal half; in O. amieu sp. n. and O. arok gonopods are so reduced they cannot be identified clearly whereas in ventral view they are subquadrate in O. houailou sp. n.
Male antennae with 22 flagellomeres; flagellomeres rectangular in profile, without sensilla placodea; anterior wing length 1.4 mm (n=1); tibial spurs 0,3,4; abdominal sternite VII with a short slender apico-mesal spur.
Male, genitalia (Figs
Oxyethira species male genitalia. 23, 24 O. amieu sp. n., ventral and lateral views 25–27 O. houailu sp. n., ventral view, phallic apparatus and lateral view 28–30 O. insularis Kelley, ventral view, spine and setose lobe of left subgenital process and phallic apparatus 31–34 O. parinsularis sp. n., dorsal, ventral and lateral views and phallic apparatus. Abbreviations: gon = gonopod; r sp = rod of subgenital process; st lb = setose lobe of subgenital process; VIII, IX = abdominal segments VIII and IX.
Holotype. Male, New Caledonia, Chute, ~15 km N Col d’Amieu on La Foa–Canala Rd, xii.1998, A. Wells, (MNHP).
Named for the Col d’Amieu.
Known only from the type locality, a waterfall towards the top of the massif (Fig.
Males of O. houailou sp. n. resemble those of O. amieu and O. arok, but are distinguished by having gonopods quadrate in ventral view and separated by mid-ventral V-shaped excision, by having mesally directed apices on ventral processes, and spine on the phallic apparatus subapical and short compared with longer spines of the other two species
Male antennae with 21–27 flagellomeres; flagellomeres subquadrate in profile, without sensilla placodea, but with dense sensilla auricillica; anterior wing length, male 1.4–1.9 mm (n=10); tibial spurs 0,3,4; abdominal sternite VII with a short sharp mesal spur.
Male, genitalia (Figs
Female unknown.
Holotype. Male (on slide), New Caledonia, Province Nord, small fall ~10 km SW Houaïlou, on Bourail road, 16.xii.1998, A. Wells (MNHP).
Paratypes. New Caledonia: 1 male (on slide), data as for holotype, (
Named for the settlement of Houaïlou, near the type locality.
This species was collected from widespread localities along the length of the island (Fig.
Oxyethira insularis Kelley, 1989: 196, figs 34, 43, 57.
This species shares with O. parinsularis sp. n. the feature of gonopods, in ventral view, in the form of a prominent, sclerotised, Y-shaped structure, arising at around the distal third of abdominal segment IX, but differs from O. parinsularis sp. n. by having the fused basal section over 2× length of the divergent distal arms, not forming a shallow sclerotised band, and the bifid distal arms about 1/3 length of basal stem, not greatly exceeding length of base as in O. parinsularis sp. n.
Male antennae with 18 flagellomeres; flagellomeres rectangular in profile, without sensilla placodea, with abundant sensilla auricillica. Anterior wing length 1.5–1.8 mm (n=3). Tibial spurs 0,3,4. Venter of abdominal sternite VII with median spine.
Holotype. Male, New Caledonia, mountain stream up Boulari River, (
Among the extensive collection of New Caledonian Oxyethira at hand, only three specimens of this species were identified, all from the south of the island (Fig.
Males are distinguished from those of the closely similar O. insularis by the shape of gonopods which have longer, more elongate and slender divergent arms and very short, fused basal portion.
Male antennae with 18–19 flagellomeres, flagellomeres rectangular in profile, without sensilla placodea, with numerous sensilla auricillica; anterior wing length 1.5–1.8 mm (n=7); tibial spurs 0,3,4; abdominal sternite VII with median spine.
Male, genitalia (Figs
Holotype. Male (on slide), New Caledonia, Province Sud, Mt Dzumac, source stream of Ouinne River, near crosspoint to mountain track, 22°02.073'S, 166°28.460'E, 810 m, 18.xi–4.xii.2003, Malaise trap, loc#030, leg. K.A. Johanson, (MNHP).
Paratypes. 12 males (2 on slides), data as for holotype, (
The close similarity between this species and O. insularis is worrying, especially since both were taken at one site, yet the differences are clear and consistent. The species was taken only in the southern province (Fig.
Subgenus Pacificotrichia, based on the type species, Oxyethira oropedion Kelley, was diagnosed by
Seven species were referred by
The following New Caledonian species are referred to this subgenus: O. oropedion (including O. derek Oláh & Johanson (syn. n.)); O. quadrata sp. n.; O. dorsennus Kelley; O. rougensis sp. n.; O. indorsennus Kelley (including O. tompa Oláh & Johanson (syn. n.); O. melasma Kelley; O. nehoue sp. n.; O. ouenghi sp. n.; O. mouirange sp. n.; O. enigmatica sp. n.; O. scutica Kelley, O. spicula sp. n. and O. digitata sp. n.
Oxyethira oropedion Kelley, 1989: 198, figs 38, 46, 50, 51, 60.
Oxyethira derek Oláh & Johanson, 2010a: 95, figs 49–51. syn. n.
Males are readily recognised by the short, setose mid-apicoventral lobes situated between the gonopods [present in the holotype, although not illustrated clearly by
Antennae: male 22–23 flagellomeres, banded with terminal 3 flagellomeres pale, then 5 dark, 1 light, 2 dark 4 light, then dark to base, flagellomeres bearing sensilla placodea; female antennae with 19 flagellomeres, with terminal 3 flagellomeres pale, 5 dark, 5 light, then dark to base. Wing length male 1.6–1.9 mm; female 1.4–1.9 mm. Tibial spurs 0,3,4. Abdominal sternite VII with median spur on distal margin.
Oxyethira species male genitalia. 35–37 O. oropedion Kelley, phallic apparatus, ventral and lateral views 38 O. quadrata sp. n. ventral view 39, 40 O. dorsennus Kelley, lateral and ventral views 41–43 O. indorsennus Kelley, lateral view, phallic apparatus and ventral view. Abbreviations: gon = gonopod; sp = subgenital process; st lb = setose lobe of subgenital process; VIII, IX = abdominal segments VIII and IX.
Holotype. Male, New Caledonia, Plateau de Dogny, (
Oxyethira oropedion is one of the more commonly collected New Caledonian Oxyethira species, occurring throughout the island (Fig.
This species is distinguished in the male from the similar species O. oropedion by the broadly truncate subgenital process rather than triangular to rounded, flanges on the phallic apparatus; and by the significantly longer antennae of 32 flagellomeres compared to 22–24 flagellomeres.
Male antennae with 32 flagellomeres, flagellomeres rectangular in profile, without sensilla placodea; anterior wing length 1.8 mm (n=1); tibial spurs 0,3,4; posterior spurs on hind leg longer than anterior spurs; abdominal sternite VII with sharp spur sub-apico-medially.
Male, genitalia (Fig.
Holotype. Male (on slide), New Caledonia, Province Sud, Mt Dzumac, source stream of Ouinne River, near crosspoint to mountain track, 22°02.073'S, 166°28.460'E, 810 m, 18.xi–4.xii.2003, Malaise trap, loc#030, leg. K.A. Johanson (MNHP).
The name quadrata is descriptive of the shape of the subgenital processes.
Only one specimen of this species was identified, from the far south of the island (Fig.
Oxyethira dorsennus Kelley, 1989: 199, figs 35, 44, 58.
Males are distinguished from the closely similar O. indorsennus by having spur formula 0,2,4, apical margins of gonopods truncate and subgenital process triangular rather than rounded as in O. indorsennus. These features also distinguish O. dorsennus from O. oropedion, which has apically well-separated gonopods with a pair of small setose lobes midventrally.
Male, antennae with 24 flagellomeres, flagellomeres rectangular in profile, without sensilla placodea. Anterior wing length, 1.5–2.1 mm (n=2). Tibial spurs 0,2,4. Abdominal sternite VII with short strong spur medially, offset from distal margin.
Holotype. Male, New Caledonia, mountain stream up Boulari River, (
Only three further specimens of this species have been identified, all from the south of the island (Fig.
Oxyethira indorsennus Kelley, 1989: 199, fig. 36.
Oxyethira tompa Oláh & Johanson, 2010a: 98, figs 56–58. Syn. n.
In ventral view males of O. indorsennus are distinguished from those of similar species, such as O. dorsennus, O. oropedion and O. quadrata by the rounded apical margins of the gonopods and apically rounded subgenital process, which contrast with the truncate apical margins of gonopods and triangular subgenital process of O. dorsennus; the absence of a pair of small median setose lobes between gonopods and clearly rounded subgenital process distinguish them from O. oropedion; and closely abutting gonopods, rather than widely separated, and rounded subgenital process rather than quadrate separates O. indorsennus from O. quadrata.
Male, antennae with 23–24 flagellomeres, flagellomeres rectangular in profile, without sensilla placodea. Anterior wing length, 1.3–2.1 mm (n=10). Tibial spurs 0,3,4. Abdominal sternite VII with sharp median spine, offset from distal margin.
Oxyethira indorsennus Kelley, Holotype. male, New Caledonia, mountain stream up Boulari River, (
1 Male, Province Sud., Rivière Bleue, 282 m, stony river, loc 4, 22°05.705'S, 166°38.225'E, Malaise trap, 13–16.xi.2001, leg. K.A. Johanson, T. Pape & B. Viklund(
This species shows a general resemblance to O. smolpela Wells, from New Guinea, but that species has a distinctive titillator on the phallic apparatus which is lacking in O. indorsennus.
Oxyethira rougensis sp. n. groups with O. oropedion, O. quadrata, O. dorsennus, and O. indorsennus, but unlike any of those species, males have venter of abdominal segment VIII deeply and narrowly excised apico-medially and subgenital process sculpted latero-distally.
Male antennae: with 23–24 flagellomeres, flagellomeres rectangular in profile; anterior wing length 1.1–1.8 mm (n=3); tibial spurs 0,3,4; abdominal sternite VII with short sharp median spine on distal margin.
Male, genitalia. Abdominal segment VIII conical, distally about half width of proximal quarter. Segment IX in ventral view almond shaped, in lateral view triangular proximally, mid-dorsally less than half length of venter. Gonopods fused basally, discrete and rounded distally, with slender mid-ventral apodeme reaching to proximal margin of segment IX; subgenital processes fused, forming a stout plate, rounded distally in ventral view, angular in lateral view, with subapical sculpturing towards apex on each side, and small notch mid apically; setose lobes about 2/3 length of plate. Phallic apparatus slender, straight, with short apical spine.
Oxyethira species male genitalia. 44–46 O. rougensis sp. n., ventral and lateral views and phallic apparatus 47–49 O. mouirange sp. n., ventral view, phallic apparatus and lateral view 50–52 O. ouenghica sp. n., phallic apparatus, ventral and lateral views. Abbreviations: gon = gonopod; sp = subgenital process; st lb = setose lobe of subgenital process; VIII, IX = abdominal segments VIII and IX.
Holotype. Male (on slide), New Caledonia, Province Nord, Plaine des Gaïacs, Rivière Rouge, 14.2 km NW summit of Mt Rouge, 50 m upstream road RT1 Noumea–Koné, 21°31.573'S, 164°46.690'E, 23 m, 2.i.2004, light trap, loc#104, leg. K.A. Johanson (
Paratypes. 2 males (on slides), collection data as for holotype.
Oxyethira rougensis was taken only at the type locality, in the north-west of the island (Fig.
A member of Pacificotrichia subgenus, but distinguished from other species by males with discrete, elongate, sub-triangular gonopods.
male. Antennae with 24–25 flagellomeres; flagellomeres without sensilla placodea, rectangular in profile; anterior wing length 1.6–1.7 mm (n=2); tibial spurs 0,3,4; abdominal sternite VII with small sharp median spur.
Male, genitalia. In ventral view with abdominal segment VIII ovoid, apical margin with wide V-shaped excavation medially; without midventral apodeme; in lateral view, with apical margins broadly rounded; abdominal segment IX almond shaped. Gonopods sharply triangular; subgenital processes fused, forming subtriangular plate that is sharply triangular in lateral view; setose lobes less than half length of plate. Phallic apparatus extremely long, apically bifid, split into two equal-length sections.
Holotype. Male (on slide), New Caledonia, Province Sud, Monts Kwa Ne Mwa, on road between Noumea and Yaté, 2.0 km E Pic Mouirange, 22°12.356'S, 166°40.798'E, 220 m, 7–16.xi.2003, Malaise trap, loc#014, leg. K.A. Johanson (MNHP).
Paratypes. 2 males (on slides, one headless), collection data as for holotype (
Named for Mt Mouirange near which the type was collected.
Taken only at two well-separated sites in the south of the island (Fig.
Males resemble O. nehoue sp. n., O. melasma, and O. spicula sp. n. in having more or less triangular median ventral processes in the male genitalia. However, males of O. ouenghi differ from other New Caledonian species, having gonopods in form of slender curved spines adjacent to the narrow midventral process.
male. Antennae with 23–25 flagellomeres, each flagellomere length greater than width, without sensilla placodea; Anterior wing length 1.7–1.8 mm (n=2); tibial spurs 0,3,4; abdominal sternite VII with mid apical spine.
Male, genitalia (Figs
Holotype. Male (on slide), New Caledonia, Province Nord, Bouérabate Stream, S Mont Ninndo, along road Barabache–Boulagoma, 20°17.409'S, 164°11.242'E, 60 m, 19.xii.2003–7.i.2004, Malaise trap, loc#089, leg. K.A. Johanson (MNHP).
Paratypes. 8 males, same data as for holotype; 1 male (on slide), Ouenghi River, nr Boulouparis, 14.xii.1983, A Wells (
Named for the Ouenghi River beside which the first specimen was collected.
The apparent distribution of this very distinctive species is very odd, with one sample being taken in the extreme north of the island, the rest at sites in the south-west (Fig.
In having an elongate apical spine on the phallic apparatus, males of this species resemble those of O. scutica, but in O. enigmatica sp. n. the spine is shorter and strap-like, not thread-like as in O. scutica; abdominal segment VIII tapers and gradually increases in width distally, and distal margin of venter is more widely and shallowly excavated than in O. scutica in which it is deeply and narrowly excised.
Male antennae with 23–24 flagellomeres, flagellomeres without sensilla placodea, in profile almost subquadrate, but wider apically than proximally; anterior wing length, 1.5–1.6 mm (n=10); tibial spurs 0,2,4; abdominal segment VII with a sharp spur medially on distal margin.
Male, genitalia (Figs
Oxyethira species male genitalia. 53, 54 O. enigmatica sp. n., ventral and lateral views 55–57 O. melasma Kelley, ventral and lateral views, and phallic apparatus, drawn from holotype 58–60 O. nehoue sp. n., ventral view, phallic apparatus and lateral view. Abbreviations: gon = gonopod; sp = subgenital process; st lb = setose lobe of subgenital process; VIII, IX = abdominal segments VIII and IX.
Holotype. Male (on slide), Province Sud, W part of Plaine des lacs, 150 m downstream bridge at La Capture, 22°15.967'S, 166°49.493'E, 261 m, 4–22.xi.2003, Malaise trap, loc#007, leg. K.A. Johanson (MNHP).
Paratypes. 54 males (9 on slides), data as for holotype.
Enigmatica: enigmatic in presenting a puzzle, being so similar to O. (P.) scutica in some respects, yet distinctive.
This species is known only from the large sample taken at the type locality in the extreme south of the island (Fig.
Oxyethira melasma Kelley, 1989: 200, figs 37, 45, 59.
A mismatch between the published description and
Oxyethira melasma Kelley is here redescribed and figured from the holotype specimen. We can only suggest that Kelley had several specimens at hand, macerated one and labelled it “holotype”, but described and illustrated a specimen other than that labelled “Type”. The type has to be the name-bearing specimen and is redescribed here.
Males resemble O. nehoue sp. n., O. spicula sp. n., and O. ouenghi in having have a more or less triangular median ventral processes in the male genitalia. However, they most closely resemble O. nehoue sp. n. from which they are distinguished by spur count 0,2,4, gonopods fused, in ventral view sharply tapered proximally, narrowly parallel-sided distally, rounded apically, apex of phallic apparatus rounded without apical spine, in contrast to O. nehoue sp. n. in which the fused gonopods appear triangular in ventral view, and acuminate apically, and ventral process in lateral view sclerotised and arched ventrally.
male. Antennae with 19–26 flagellomeres, apical 3 flagellomeres pale, next 5 dark, then 13 pale and basal flagellomeres dark; anterior wing length 1.4–1.9 mm (n=10); tibial spurs 0,2,4; abdominal sternite VII with small sharp median spine, offset from margin.
Male, genitalia (Figs
Holotype. Male, Mountain stream up Boulari River (
The species was taken at a number of sites in the southern province of the island (Fig.
This species resembles O. melasma and was illustrated and described as that species by
Male antennae with 25–27 flagellomeres; anterior wing length 1.3–1.7 mm (n=7); tibial spurs 0,3,4; abdominal sternite VII with sharp apical spine.
Genitalia (Figs
Female unknown.
Holotype. Male (on slide), Province Nord, Rivière Néhoué, camp Amenage de Néhoué, 20°25.037'S, 164°13.222'E, 12 m, 19.xii.2003, light trap, loc#090, leg. K.A. Johanson (MNHP).
Paratypes. 5 males (on slides), data as for holotype (
Named for the Rivière Néhoué where it was collected.
The distribution of this species is similar to that of O. ouenghi, mainly collected from a cluster of southern sites, but with one site in the far north (Fig.
Oxyethira scutica Kelley, 1989: 200, figs 39, 47, 52, 53, 61.
Males superficially resemble those of O. enigmatica, having a long apical process on the phallic apparatus, antennae with 23–25 flagellomeres, and spur formula of 0,2,4, but are distinguished by apical process on phallic apparatus longer and thread-or whip-like in appearance, rather than strap-like as in O. enigmatica, and abdominal segment VIII with distal margin of venter of deeply and narrowly excised, compared with shallowly excavated margin of O. enigmatica. Females resemble those of O. oropedion (
Antennae: male with 21–24 flagellomeres, flagellomeres subquadrate in profile, without sensilla placodea; female with 18 flagellomeres. Fore wing length: male 1.1–1.5 mm (n=8), female 1.4 mm (n=2). Spurs 0,2,4. Abdominal sternite VII with sharp median spur.
Oxyethira species male genitalia. 61–63 O. scutica Kelley, ventral view and enlarged gonopods and subgenital process, and lateral view 64–66 O. spicula sp. n., ventral view and enlarged gonopods and subgenital process, and lateral view. Abbreviations: gon = gonopod; sp = subgenital process; st lb = setose lobe of subgenital process; VIII, IX = abdominal segments VIII and IX.
Holotype. Male, New Caledonia, mountain stream up Boulari River, (
This appears to be another southern species (Fig.
Males are most similar to O. melasma, O. nehoue and O. ouenghi all of which have more or less triangular median ventral processes in the male genitalia, but can be recognised by the expanded apex of the phallic apparatus with a prominent acute spine, the very long, proximally rounded, abdominal segment VIII that tapers distally and completely obscures segment IX, and the shape of the plate formed from fused gonopods and subgenital processes.
Male antennae damaged, at least with 19 flagellomeres, flagellomeres rectangular in profile, without sensilla placodea; anterior wing length 1.9 mm (n=1); tibial spurs 0,3,4; abdominal sternite VII with a coarse spur medially.
Male, genitalia (Figs
Female unknown, although a single unknown female of an Oxyethira species was collected with the holotype and could be of this species. The terminalia of this female are slender and elongate, of the form seen in O. oropedion and O. scutica, with a V-shaped marking ventrally on abdominal segment VIII.
Holotype. Male (slide), New Caledonia, Province Sud, Rivière des Lacs, 1.1 km NW Lac en Huit, 4.9 km NW summit of Pic du Grand Kaori, 22°15.195'S, 166°52.178'E, 10.xii.2003, light trap, loc#078, leg. K.A. Johanson (MNHP).
Name being descriptive of the spicule-like spine on the phallic apparatus.
This species is known only from the type specimen, collected in the far south (Fig.
One of the New Caledonian species of Oxyethira with genitalia retracted within the very rounded abdominal segment VIII but differs from other species with this feature such as O. incana and O. spicula by well-defined clearly branched gonopods and the phallic apparatus a simple rod, sharply bifid apically.
Male antennae with 25–29 flagellomeres; flagellomeres slender rectangular in profile, without sensilla placodea, terminal 5 flagellomeres pale, followed by 3 dark, 10 pale, rest dark; anterior wing. Length 3.0–3.7 mm (n=4); tibial spurs 0,3,4; abdominal segment VII short, sharp mid ventral spur.
Male, genitalia (Figs
Oxyethira species male genitalia. 67–69 O. digitata sp. n., phallic apparatus, ventral and lateral views 70, 71 O. incana (Ulmer), ventral view and phallic apparatus 72–74 O. macropennis sp. n., ventral and lateral views and phallic apparatus. Abbreviations: gon = gonopod; sp = subgenital process; st lb = setose lobe of subgenital process; VIII, IX = abdominal segments VIII and IX.
Holotype. Male (on slide), Province Sud, side stream to Rivière Blanche, 10.75 km SW Pont Pérignon, 22°10.073'S, 166°39.903'E, 180 m, 6–16.xi.2003, Malaise trap, loc#012, leg. K.A. Johanson (MNHP).
Paratypes. 6 males (on slides), same data as holotype (
Named for the finger-like lobes of the gonopods in male.
Oxyethira digitata shares a southern distribution (Fig.
Type localities of Oxyethira species. 75 O. tiwaka sp. n. and O. ouenghi sp. n. (collected together with the hydroptilid species Oxyethira caledonensis, O. oropedion, O. indorsennus, O. incana, Hydroptila losida, Hellyethira malleoforma, Acritoptila disjuncta Kelley, 1989, A. crinita Kelley, 1989, A. glossocercus Kelley, 1989 and A. amphapsis Kelley, 1989) 76 O. perignonica sp. n. (collected together with the hydroptilid species Oxyethira oropedion) 77 O. abbreviata sp. n. (no other Hydroptilidae species collected at this site) 78 O. incurvata (collected together with the hydroptilid species Oxyethira caledonensis) 79 O. parinsularis sp. n. and O. quadrata sp. n. (collected together with the hydroptilid species Oxyethira incurvata sp. n., O. caledonensis, O. houailou sp. n., O. insularis, O. oropedion, O. melasma, O. digitata sp. n., Acritoptila disjuncta, A. crinita, A. ouenghica Wells, 1995, Caledonotrichia illiesi Sykora, 1967, C. minuta Wells, Johanson & Mary-Sasal, 2013, C. ouinnica Wells, Johanson & Mary-Sasal, 2013, C. nyurga Oláh & Johanson, 2010, Paroxyethira atypica Wells & Johanson, 2012 and P. dzumac Wells & Johanson, 2012) 80 O. dorsennus sp. n. (collected together with the hydroptilid species Hydroptila losida and Paroxyethira dumagnes Kelley, 1984).
Type localities of Oxyethira species. 81 O. rougensis sp. n. (collected together with the hydroptilid species Oxyethira oropedion, Hydroptila losida, Hellyethira malleoforma, Acritoptila crinita, A. macrospina Wells & Johanson, 2014 and Paroxyethira opposita Wells & Johanson, 2012) 82 O. enigmatica sp. n. (collected together with the hydroptilid species O. perignonica sp. n., O. melasma and Acritoptila disjuncta) 83 O. nehoue sp. n. (collected together with the hydroptilid species Oxyethira oropedion, O. incana, Hydroptila losida, Hellyethira malleoforma and Acritoptila disjuncta) 84 O. digitata sp. n. (collected together with the hydroptilid species Oxyethira incurvata sp. n., O. indorsennus, O. scutica, Hydroptila losida, Caledonotrichia illiesi and C. minuta) 85 O. macropennis sp. n. (no other Hydroptilidae species collected at this site).
Erected at genus level by
Oxyethira incana Ulmer, 1906: 102 (see
Males of this species are distinguished from others in the New Caledonian fauna by abdominal segment VIII with disto-lateral angles spiny, and venter produced and rounded distally not excised apico-ventrally, forming a shield over other genital structures which are strongly fused; and by phallic apparatus curiously stout and medially curved, lacking a titillator. Female terminalia longer and terminally more slender than those of caledoniensis group species, but not as slender as in members of subgenus Pacificotrichia, with sternite X bearing a jet black quadrate plate.
Antennae: males 25–28 flagellomeres, flagellomeres about twice as long as wide; female 20–21 flagellomeres, flagellomeres subquadrate in profile. Anterior wing length: males 1.8–2.2 mm (n=10); females 1.8–2.4 mm (n=10). Spurs 0,3,4. Abdominal sternite VII without median spine.
Numerous males, females, Province Nord, Amoa River, 23 m, loc 20, 12 km W Poindimié, 22°58.092'S, 165°11.804'E, light trap, 26.xi.2001, leg. K.A. Johanson, T. Pape & B. Viklund (
In New Caledonia this species was taken only in the northern province (Fig.
One highly aberrant species, O. macropennis sp. n., is here unplaced to subgenus. Males share the diagnostic features of species of Oxyethira, including wing shape and venation, but have unusual male genital features, possibly aligning the species more with species of Paroxyethira. For the present we assign it to Oxyethira, albeit tentatively.
Immediately recognised by the remarkable form of the phallic apparatus, which has a contorted twist towards the stout seta-bearing apex, and the brush of shorter stout setae medially on abdominal sternite IX. By these features it is distinguished clearly from all other New Caledonian species.
Male antennae with 22–24 flagellomeres; flagellomeres urn-shaped, few sensilla placodea subapically, dense sensilla auricillica; anterior wing length 1.7–2.1 mm (n=5); tibial spurs 0,3,4; abdominal sternite VII without medial spur.
Male, genitalia (Figs
Oxyethira macropennis is quite unlike all other New Caledonian species, however it conforms with the diagnostic features of members of the Oxyethira and thus is assigned to this genus, albeit somewhat tentatively.
Holotype. Male (on slide), New Caledonia, Province Sud, south of Plaine des Lacs, 4.0 km N Prony, 22°16.906'S, 166°49.402'E, 9–22.xi.2003, Malaise trap, loc#017, leg. K.A. Johanson (MNHP).
Paratypes. 6 males (2 on slides), data as for holotype (NMHR); 1 male, Province Sud, Sarraméa, 220 m, forest stream, loc 10, 21°37.883'S 165°51.958'E, Malaise trap, 18–21.xi.2001, leg. K.A. Johanson, T. Pape & B. Viklund (
Name descriptive of the relatively large phallic apparatus.
Oxyethira macropennis was taken only at two well-separated sites in the south of the island. A photograph of the type locality with the trap is rendered in Fig.
Hydroptila losida Mosely, 1953: 505;
The only Hydroptila species among New Caledonian micro-caddisflies, H. losida is recognised by the absence of ocelli on the dorsal head, and tibial spur count of 0,2,4; and in the male by gonopods well developed, in ventral view elongate divergent rods, each with a pair of dark spurs distally, in lateral view, club-shaped; and phallic apparatus slender, distally comprising a slender, tapered spine adpressed to the section containing the ejaculatory duct, or in some specimens these two parts are separated and divergent; and in the female by the triangular shape of sternite VIII with the two small triangular sclerotised areas laterally at about half length of the sternite.
Numerous males, females, Province Nord, Amoa River, 23 m, loc 20, 12 km W Poindimié, 22°58.092'S, 165°11.804'E, light trap, 26.xi.2001, leg. K.A. Johanson, T. Pape & B. Viklund(
In New Caledonia H. losida is abundant and widespread across the island (Fig.
Hellyethira malleoforma Wells, 1979: figs 41–45;
Males of this species are distinguished by their complex asymmetrical genital structures, including multilobed gonopods, and females by the sclerotised annulus formed by abdominal segment VIII (see
2 males, females, Province Sud, Sarraméa, 220 m, forest stream, loc 10, 21°37.883'S 165°51.958'E, Malaise trap, 18–21.xi.2001, leg. K.A. Johanson, T. Pape & B. Viklund(
Hellyethira malleoforma is the only representative in New Caledonia of this diverse Australian genus that also occurs more broadly but less commonly in SE Asia and New Guinea. This species is widespread and often abundant in New Caledonia (Fig.
1 | Tibial spur formula 0,2,4 | 2 |
– | Tibial spur formula 0,3,4 | 6 |
2(1) | Phallic apparatus tipped by fine whip-like flagellum (Fig. |
O. scutica |
– | Phallic apparatus with strap-like distal process (Figs |
3 |
3(2) | Phallic apparatus with strap-like apical or subapical process (Figs |
4 |
– | Phallic apparatus without apical process (Figs |
5 |
4(3) | Phallic apparatus with elongate strap-like apical process (Fig. |
O. enigmatica |
– | Phallic apparatus with short strap-like subapical process (Fig. |
O. perignonica |
5(3) | Gonopods fused at base, separate distally, apices truncate in ventral view (Fig. |
O. dorsennus |
– | Gonopods fused throughout, narrowly truncate apically, a pair of small setose lobes at mid length (Fig. |
O. melasma |
6(1) | Phallic apparatus twisted and irregular in shape distally, bearing a single stout seta (Fig. |
O. macropennis |
– | Phallic apparatus without seta, with strap-like flange or process (e.g. Figs |
7 |
7(6) | Gonopods inserted midway or proximally on venter of abdominal segment IX, in ventral view in form of Y-shaped structure or pair of widely separated ‘horns’ (Figs |
8 |
– | Gonopods situated distally on venter of abdominal segment IX (e.g. Figs |
9 |
8(7) | Gonopods in ventral view distinctly Y-shaped (Fig. |
O. insularis |
– | Gonopods in ventral view in form of pair of widely separated spines joined basally by short sclerotised strip (Fig. |
O. parinsularis |
9(7) | Abdominal segment VIII extended disto-laterally as pair of sclerotised spines (Fig. |
O. incana |
– | Abdominal segment VIII without apico-lateral sclerotised spines (Figs |
10 |
10(9) | Ventral processes in form of pair of laterally situated rods or spines (Figs |
11 |
– | Ventral processes fused, in ventral view forming median plate (Figs |
18 |
11(10) | Abdominal segment IX in ventral view subquadrate (Figs |
12 |
– | Abdominal segment IX in ventral view rounded, conical or triangular proximally (Figs |
14 |
12(11) | Gonopods forming sclerotised cones at apico-lateral angles of abdominal segment IX (Fig. |
O. abbreviata |
– | Gonopods reduced completely or in form of short blunt tabs, widely separated on apical margin of abdominal segment IX (Figs |
13 |
13(12) | Gonopods reduced completely (Fig. |
O. spinifera |
– | Gonopods in ventral view in form of blunt sclerotised tabs scarcely longer than wide, well separated on apical margin of abdominal segment IX (Fig. |
O. tiwaka |
14(10) | Gonopods apparently reduced completely or possibly present as marginal sclerotisations on distal margin of abdominal segment IX (Figs |
15 |
– | Gonopods recognisable as sclerotised prominences or processes on distal margin of abdominal segment IX (Figs |
16 |
15(14) | Abdominal segment IX in ventral view tapered distally (Fig. |
O. amieu |
– | Abdominal segment IX in ventral view parallel-sided in distal half (Figs |
O. arok |
16.(14) | Gonopods in form of short domes situated slightly laterally on distal margin of abdominal segment IX (Figs |
O. caledoniensis |
– | Gonopods in ventral view broad, stoutly quadrate and separated by narrow v-shaped cleft (Fig. |
17 |
17(16) | Gonopods in ventral view stoutly quadrate (Fig. |
O. houailu |
– | Gonopods in ventral view in form of laterally situated finger-like mesally curved processes (Fig. |
O. incurvata |
18(10) | Phallic apparatus dilated distally, with a sharp, sclerotised straight apical spine (Fig. |
O. spicula |
– | Phallic apparatus not as above; abdominal segment VIII not completely obscuring gonopods and other genital processes (e.g. Figs |
19 |
19(18) | Gonopods completely fused, in ventral view in form of triangular plate; ventral process in lateral view a stoutly sclerotised arch (Fig. |
O. nehoue |
– | Gonopods either not fused or only fused basally (e.g. Figs |
20 |
20(19) | Gonopods in ventral view branched, digitiform (Fig. |
O. digitata |
– | Gonopods unbranched (e.g. Figs |
21 |
21(20) | Gonopods elongate triangular, acute apically (Figs |
22 |
– | Gonopods in ventral view rounded apically (Figs |
23 |
22(21) | Plate formed by fusion of subgenital processes subtriangular, slender in distal half, with paired short setae subapically (Fig. |
O. ouenghica |
– | Plate formed by fusion of subgenital processes broadly triangular, rounded apically and without setae (Fig. |
O. mouirange |
23(22) | Abdominal segment VIII, in ventral view, with medial cleft on distal margin narrow, deeper than wide (Fig. |
O. rougensis |
– | Abdominal segment VIII, in ventral view, with wide U- or V-shaped excavation on distal margin, width greatly exceeding depth (Figs |
24 |
24(23) | Fused subgenital processes in ventral view in form of subquadrate plate (Fig. |
O. quadrata |
– | Fused subgenital processes in ventral view tapered or rounded distally (Figs |
25 |
25(24) | Gonopods in ventral view fused basally, widely separated distally pair of small membranous lobes in mid ventral position, each bearing a pale stout seta (Fig. |
O. oropedion |
– | Gonopods in ventral view fused basally, free but closely abutting, with only sharp median cleft separation; without pair of median setal lobes (Fig. |
O. indorsennus |
As part of the broader study on the Trichoptera of New Caledonia by researchers from the Swedish Museum of Natural History, this work was supported by the Swedish Research Council (grant #2005-4834) and National Geographic Committee for Research and Exploration (grant #7546-03). Advice on localities and help with field work were contributed by Dr. Christina Pöllabauer (Etudes et Recherches Biologiques, New Caledonia), and Dr. Nathalie Mary-Sasal (at the time in New Caledonia). We are grateful to the authorities at Direction des Ressources Naturelles (Nouméa, New Caledonia) and the authorities at the Environment Division, Department of Economic Development and Environment, Province Nord (Koné, New Caledonia) for supporting the project with collecting and export permits. Dr. Christian Mille (Institut Agronomique néo-Calédonien, Station de Recherches Fruitières de Pocquereux, Laboratoire d’entomologie, La Foa, New Caledonia) was always enthusiastically helpful during the collecting on New Caledonia. Access for A. Wells to laboratory facilities at the Australian National Insect Collection, Canberra, is gratefully acknowledged, and for computer facilities AW thanks Australian Biological Resources Studies. The Bishop Museum, Honolulu, and particularly Shepherd Meyers, kindly hosted A. Wells’ visit in 2013 and facilitated access to the Kelley types. Two anonymous referees are thanked for their very constructive comments on the manuscript.