Research Article |
Corresponding author: John K. Moulton ( johnkmoulton@gmail.com ) Academic editor: Art Borkent
© 2021 Robert J. Pivar, Bradley J. Sinclair, John K. Moulton.
This is an open access article distributed under the terms of the CC0 Public Domain Dedication.
Citation:
Pivar RJ, Sinclair BJ, Moulton JK (2021) Revision of the genus Niphta (Diptera, Thaumaleidae) Theischinger of South America, with descriptions of nine new species and a new immature morphotype. ZooKeys 1063: 49-104. https://doi.org/10.3897/zookeys.1063.71180
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The Niphta Theischinger fauna of South America is revised to include 11 species, nine of which are described as new to science (N. acus Pivar, sp. nov., N. bifurcata Pivar & Moulton, sp. nov., N. bispinosa Pivar & Sinclair, sp. nov., N. brunnea Pivar, sp. nov., N. courtneyi Pivar, sp. nov., N. daniellae Pivar, sp. nov., N. downesi Pivar, sp. nov., N. eurydactyla Pivar, sp. nov., N. mapuche Pivar, sp. nov.). The genus Niphta is redefined, both previously described Chilean species are redescribed, N. halteralis (Edwards) and N. nudipennis (Edwards), and females are described or redescribed where possible. The first descriptions of the immature stages of South American Niphta are provided, which represent a new larval morphotype in Thaumaleidae, as larvae and pupae possess ventral adhesive structures. Furthermore, these larvae were collected from vegetation rather than rocky substrates. Illustrations and micrographs are provided for all species, and scanning electron microscopy images are included for select immatures. A key to species, distribution maps, and discussions regarding phylogenetic affinities and habitat are also included.
Andes, Chile, diversity, madicolous, midge, seepages
Thaumaleidae, or madicolous midges, consist of nearly 200 species classified in seven genera. They have no known medical or economic importance to humans, and are considerably less studied than their presumptive sister family, Simuliidae (black flies) (
South America has eight described species of Thaumaleidae in three genera prior to this study, known from only three countries: Austrothaumalea Tonnoir (five species, Chile and Argentina), Neothaumalea Pivar, Moulton and Sinclair (one species, Brazil) and Niphta Theischinger (two species, Chile).
The focus of this study is the South American Niphta, a small genus with two described species in Chile (
Efforts were made to recollect fresh material from as near as possible to the perceived type localities. Adults were collected using an aerial net to sweep above the madicolous substrate and adjacent riparian vegetation. Sweeping for adults should be attempted first, if possible, to avoid having to sweep less accessible areas if they try to escape when searching for larvae. Immatures were collected by using forceps to pull them off the substrate, or by pouring water over the substrate and flushing immatures into a white pan (
Adult genitalia were cleared with hot 85% lactic acid. Representative adults and immatures were also cleared with the GeneJET Genomic DNA Purification Kit #K0722 (ThermoScientific, Waltham, MA) to maintain important membranous and lightly sclerotised structures, and to extract DNA for subsequent molecular study. The GeneJET lysate preparation protocol was followed and cleared voucher specimens were stored in 70% non-denatured ethanol. Positive identifications of females and immatures were made by comparing their DNA sequences to those of identified males. Specimens identified through molecular means are denoted by an asterisk (*) in the Type material and Additional material examined sections. Pinned specimens were dried using
Specimens were viewed using a Meiji Techno RZ stereomicroscope mounted with a Progres Gryphax Naos 22-megapixel camera (Jenoptik, Jena, Germany) and aided by iSolution Lite x64 software (Focus Precision Industries, Victoria, MN, USA) to take light micrographs of pinned adults and immatures in alcohol. Image stacks were created using Helicon Focus 6.7.1 (HeliconSoft, Roseau Valley, Dominica). Cleared terminalia and larval head capsules in glycerine were viewed with an Olympus BH-2 compound microscope equipped with DIC and images were taken following the same methods as above. Line drawings were first traced from stacked micrograph images captured using the compound microscope, structures were re-evaluated by closely re-examining specimens (as stacking failed to clearly differentiate critical internal structures), then line drawings were inked and digitised for publication. The left gonocoxite and gonostylus were intentionally omitted for male Niphta nudipennis group species to allow for clearer visualisation of remaining genitalic characters.
Samples were prepared for scanning electron microscopy by transferring specimens from 95% non-denatured ethanol into a 12 mm × 30 μm microporous specimen capsule (Electron Microscopy Services, Hatfield, PA). Capsules were then subjected to the following HMDS dehydration series, each step lasting 20 minutes: 100% ethanol, 1:1 ethanol to HMDS, 1:75 ethanol to HMDS, then two steps of 100% HMDS. Dried specimens were mounted on carbon tape affixed to 45°/90° aluminium stubs and sputter-coated with gold for 10 sec at 20 μA in a SPI-Module Sputter Coater (West Chester, PA). Specimens were viewed with a Hitachi TM3030 electron microscope (Tokyo, Japan) at a voltage of 15 kV.
Terms used for adult structures follow
Specimens are deposited in the following repositories: Canadian National Collection of Insects, Ottawa, Canada (
Label data for primary types are presented exactly as they appear. Data are listed from the top downward on the staging pin, with data from each label enclosed in quotation marks; lines are delimited by a forward slash mark. Repository is given in parentheses and
A key to adults of South American genera of Thaumaleidae is provided in
Niphta Theischinger, 1986: 314. Type-species: Niphta bickeli Theischinger (original designation).
Niphta is characterised as follows: presence of a distinct antealar ridge; proepisternal setae absent; microtrichia of R1(+R2+3) confined to base near humeral crossvein; R2+3 crossvein situated closer to apex of R1(+R2+3) than to origin of R4+5; R1 and R1(+R2+3) with three weakenings or depigmented gaps; R4+5 often with arch not strongly produced; basal spur of CuA absent; gonocoxites broad, not much longer than wide; hypandrium absent; gonocoxal plate extended posterodorsally forming a medial process; parameres fused medially, emerging from gonocoxal plate complex.
Adult. Eye bridge broad, comprising more than five facets. Scutum clothed in both short and long setae; scutellum with row of long, black marginal setae. Supra-alar region produced into distinct antealar ridge (Fig.
Chile and Australia.
Prior to this study, only five described species of Niphta were known from all regions and few phylogenetic affinities had been discussed.
The N. bickeli group: This group is characterised by the following features: broad gonocoxites extending to the posterior epandrial margin and lacking projections; long gonostyli; parameres fused medially, then separating into two arms that do not project anteriorly; a pair of apodemes from base of parameres extend to posterior margin of epandrium, on either side of the anus; cerci inconspicuous, thinly sclerotised and unpigmented; females lack projection at base of hypogynial valves. This group is greyish black in colouration. Additionally, immatures of N. collessi Theischinger lack ventral adhesive structures and protuberances on the larval head capsule (
The N. halteralis group: This group is characterised by the following features: gonocoxites extending to midpoint of epandrium, lacking pointed projections; short and narrow gonostyli; parameres fused throughout; prominent cerci projecting anteriorly, extending well beyond posterior epandrial margin; females lack projection at base of hypogynial valves. This group is darkly coloured, mostly black and grey. Immature stages have ventral adhesive structures, are collected from rocky substrates and larval head capsules have many protuberances. The N. halteralis group is known from Chile and includes the following species: N. acus sp. nov., N. downesi sp. nov., N. halteralis (Edwards), and N. mapuche sp. nov.
The N. nudipennis group: This group is characterised by the following features: gonocoxites extending, at most, to midpoint of epandrium, and bearing pointed projections; broad, short gonostyli cheliform or finger-like; parameres fused medially. separated into two arms that project anteriorly and may be flexed or extended; cerci small, projecting anteroventrally; females possess distinct blunt or pointed projection at base of hypogynial valves; sternite 8 highly modified (genital fork and lateral arms); Female sternite 9 greatly expanded and heavily sclerotised, presumably reflecting the highly modified male genitalia. This group tends to be lighter in colouration. Immature stages have ventral adhesive structure, are collected from vegetation in splash zones, and larval head capsules have many protuberances. The N. nudipennis group is known from Chile and includes: N. bifurcata sp. nov., N. bispinosa sp. nov., N. brunnea sp. nov., N. courtneyi sp. nov., N. daniellae sp. nov., N. eurydactyla sp. nov. and N. nudipennis.
1 | Gonocoxite subquadrate or conical, with posteromedial projection broad, rounded, projecting posteriorly (Fig. |
2 (N. halteralis group) |
– | Gonocoxite oblong, with posteromedial projection narrow, pointed, projecting medially (Figs |
5 (N. nudipennis group) |
2 | Paramere with hooked apex. Gonostylus straight (Figs |
N. halteralis (Edwards) |
– | Paramere without hooked apex. Gonostylus arched outwards | 3 |
3 | Gonostylus bifurcate apically. Paramere with abrupt, strongly tapered, off-centred needle-like apex (Figs |
N. acus Pivar, sp. nov. |
– | Gonostylus tapered to single apex. Paramere evenly tapered throughout | 4 |
4 | Paramere, in lateral view, divided into two filaments; dorsal filament, at most, nearly reaching posterior margin of epandrium; ventral filament not extended beyond apex of gonostylus, not easily visible (Figs |
N. mapuche Pivar, sp. nov. |
– | Paramere, in lateral view, divided into three filaments; dorsal filament extended beyond posterior margin of epandrium; paired ventral filaments extended beyond apex of gonostylus, easily visible (Figs |
N. downesi Pivar, sp. nov. |
5 | Gonostyli cheliform or bearing a finger-like projection (Figs |
6 |
– | Gonostyli not cheliform, without projections; broad at base, tapered to pointed apex (Figs |
N. daniellae Pivar, sp. nov. |
6 | Gonostyli with finger-like projection (Fig. |
7 |
– | Gonostyli cheliform, resembling crab claw (Fig. |
8 |
7 | Gonostyli with projection broad, tapered slightly at apex, without bend (Figs |
N. eurydactyla Pivar, sp. nov. |
– | Gonostyli with projection narrow throughout, bent at midpoint (Figs |
N. nudipennis (Edwards) |
8 | Gonostyli with posterior apex bifurcate (Fig. |
9 |
– | Gonostyli with posterior apex bearing single apex (Fig. |
10 |
9 | Gonocoxites with two projections, anterior one bifurcate (Figs |
N. bifurcata Pivar & Moulton, sp. nov. |
– | Gonocoxites with three separate projections (Figs |
N. courtneyi Pivar , sp. nov. |
10 | Gonocoxites with two projections; anterior one long, bifurcate; posterior one small, tooth-like (Figs |
N. bispinosa Pivar & Sinclair, sp. nov. |
– | Gonocoxites with three projections, two anterior (one small and inconspicuous at base of large one), one posterior; posterior projection slender, about as long as larger anterior projection (Figs |
N. brunnea Pivar, sp. nov. |
1 | Sternite 8 with distinct projection between hypogynial valves (Figs |
N. nudipennis group |
– | Sternite 8 without distinct projection between hypogynial valves (Fig. |
N. halteralis group |
The N. halteralis group
Holotype: ♂, glued to point with abdomen in glycerine microvial pinned beneath, labelled: “Chile: Region VIII (Bío Bío)/ Rte. Q-61, Estero Agua/ Blanca, 8.xii.2016/ 37°46'30.8"S 71°42'03.9"W/ elev. 552 m, vegetation near/ splash zones, J.K. Moulton &/ R.J. Pivar”; “HOLOTYPE/ Niphta/ acus/ Pivar [red label]” (
Ventral views of male Niphta halteralis group terminalia A N. acus sp. nov. B N. downesi sp. nov. C N. halteralis D N. mapuche sp. nov. Abbreviations: cerc, cercus; epand, epandrium; gcx, gonocoxite; gcx pl, gonocoxal plate; gcx pl fl, gonocoxal plate filament; gst, gonostylus; pm, paramere. Scale bars : 0.1 mm.
Lateral views of male Niphta halteralis group terminalia A N. acus sp. nov. B N. downesi sp. nov. C N. halteralis D N. mapuche sp. nov. Abbreviations: cerc, cercus; epand, epandrium; gcx, gonocoxite; gcx pl, gonocoxal plate; gcx pl fl, gonocoxal plate filament; gst, gonostylus; pm, paramere. Scale bars: 0.1 mm.
Niphta acus is recognised by the bifurcated apex of the gonostylus and the strongly apically tapered parameres, giving the appearance of a needle-like tip.
Male. n = 71.
Length 1.6–2.5 mm.
Colouration (Fig.
Head. Eyes above antennae broadly joined, with small triangular frons visible above antennae; frons with three to five strong setae. Flagellomeres 1–3 subquadrate, with flagellomere 1 expanded, 3 × as wide as next segment, equal to lengths of 2 and 3 combined; flagellomeres 4–10 cylindrical, becoming progressively thinner and elongate. Vertex with black setae of uniform length, with longer, black orbital setae.
Thorax. Mesoscutum with prominent antealar ridge, bearing thee pronounced setae. Scutum clothed dorsally in short, black setulae; notopleural, supra-alar and postsutural setae long, black. Pteropleuron bare. All legs with tarsi simple.
Wing. Wing length: 1.8–2.5 mm. Dark, infuscate throughout, apex somewhat narrowed; C fringed in small setulae, with a few microtrichia scattered throughout; posterior wing margin with closely spaced fringe of microtrichia; Sc incomplete; R1 and R1(+R2+3) with three weakenings or depigmented gaps, first slightly beyond R2+3, second and third closely approximated, near C; microtrichia of R1(+R2+3) confined to base near humeral crossvein, remaining veins bare; R flexed into cell br; R2+3 distinct, situated in basal third of R1(+R2+3); bend in R4+5 gentle; R4+5 and M1 running parallel toward margin; M1 straight; M2 with gentle bend in apical third; M4 with slight bend.
Abdomen. Abdominal sternite 1 narrow, spectacle-shaped; sternite 2 reduced to slender median sclerite, a few setae restricted to laterad on posterior third and medially beneath sclerite; sternites 3–7 rectangular, lacking distinct sclerites, setae restricted to posterior two-thirds; sternite 8 strongly reduced, anterior margin well sclerotised, arched slightly into preceding segment, lacking setae.
Terminalia (Figs
Female. n = 15.
Similar to male except as follows: Terminalia (Fig.
Ventral views of male Niphta nudipennis group terminalia A N. daniellae sp. nov. B N. eurydactyla sp. nov. C N. nudipennis with parameres retracted D N. nudipennis with parameres extended. Arrows indicate range of motion of parameres. Abbreviations: aed gd, aedeagal guide; cerc, cercus; epand, epandrium; gcx, gonocoxite; gcx pl, gonocoxal plate; gst, gonostylus; pm, paramere. Scale bars: 0.1 mm.
Pupa. n = 8 (Figs
Length 3.0–4.0 mm.
Colouration. Brown; with black spot above eye in developing adult.
Head. Maxillary sheath short, posteromedially directed; gently tapered toward truncate apex; apices of palpi separated medially. Three short, slender setae above black spot over eye.
Ventral views of male Niphta nudipennis group terminalia A N. bifurcata sp. nov. B N. courtneyi sp. nov. C N. bispinosa sp. nov. D N. brunnea sp. nov. Abbreviations: aed gd, aedeagal guide; cerc, cercus; epand, epandrium; gcx, gonocoxite; gcx pl, gonocoxal plate; gst, gonostylus; pm, paramere. Scale bars: 0.1 mm.
Thorax. Width nearly subequal to abdomen at widest point. Foreleg sheath projecting straight, slightly longer than wing sheaths, reaching posterior margin of sternite 2; anterior half of midleg visible anterior to wing sheath, then hidden behind foreleg, slightly shorter than foreleg; hind leg concealed behind wing sheath, only apex visible between apex of foreleg and wing sheath, shorter than foreleg. Wing sheaths not reaching posterior margin of abdominal sternite 2; large tubercle at base bearing pair of short, slender setae. Respiratory organ short and squat, much shorter than maxillary sheath, broadest subapically; bulbous; spiracular openings encircling apex; stalk wide, emerging from small tubercle. Tubercle situated posterodorsally to respiratory organ, rounded, projected slightly laterally; apex nearly touching or touching respiratory organ. Tubercle situated posterolaterally to respiratory organ crenulate, projected slightly anteriorly. Ridge located anteroventrally to respiratory organ with single, thin midlateral seta; mesothorax with group of four short, slender dorsocentral setae near ridge; single seta on humeral lobe.
Lateral views of male Niphta nudipennis group terminalia A N. daniellae sp. nov. B N. eurydactyla sp. nov. C N. nudipennis with parameres retracted D N. nudipennis with parameres extended. Arrows indicate range of motion of parameres. Abbreviations: aed gd, aedeagal guide; cerc, cercus; epand, epandrium; gcx, gonocoxite; gcx pl, gonocoxal plate; gst, gonostylus; pm, paramere. Scale bars: 0.1 mm.
Lateral views of male Niphta nudipennis group terminalia A N. bifurcata sp. nov. B N. courtneyi sp. nov. C N. bispinosa sp. nov. D N. brunnea sp. nov. Abbreviations: aed gd, aedeagal guide; cerc, cercus; epand, epandrium; gcx, gonocoxite; gcx pl, gonocoxal plate; gst, gonostylus; pm, paramere. Scale bars: 0.1 mm.
Abdomen. Subcylindrical, strongly tapered at caudal segment. Spiracles weakly developed, not projected or distinctly visible. Tergites 1–8 rectangular, without ridges; bearing pair of slender lateral setae above lateral margins and pair of slender dorsolateral setae. Tergite 9 rounded, posterior with dorsolateral ridges bearing pair of lateral setae and hind margin emarginated; projection directed posteriorly in lateral view. Sternites 3–8 rectangular, without row of faint setulae along anterior margin; lateral margins crenulate, bearing a few thin, short setae. Sternites 3 and 4 bearing pair of small lateral adhesive structures, sternite 5 bearing pair of large lateral adhesive structures on lateral margin. Caudal sternite subquadrate, with pair of posteriorly projected medial lobes; posterior margin with pair of medial ridges, curved dorsally forming small, dorsally projected tubercle in lateral view; without distinct caudal hooks.
Larva. n = 8 (Figs
Length of final instar 6.2–6.7 mm.
Colouration. Head capsule usually black or dark brown, sometimes black with light brown markings. Body mottled with various shades of grey and brown.
Head capsule (Fig.
Thorax. Prothorax with pair of anterodorsolateral protuberances bare; anterolateral protuberances with one long and two short setae; spiracular protuberance bearing two protuberances, inner protuberance with pair of setae, outer with single seta; pair of midlateral setae below anterolateral protuberance; three closely approximated setae near base of prothoracic leg (Keilin’s organ). Mesothorax and metathorax with pair of dorsolateral protuberances bearing pair of closely approximated setae, one thickened, one slender; mesothorax with additional seta beneath protuberance; lateral protuberance on both segments bearing four setae; one long seta slightly ventral to lateral protuberance; three mid-ventrolateral setae directed ventrally. Prothorax bearing proleg, posterior half with rectangular adhesive structure; meso- and metathoracic sternites with rectangular adhesive structures.
Abdomen. Sternites 1–7 modified into circular, suction cup-like adhesive structures; sternite 8 with quadrate adhesive structure, extending over sternite 9; sternite 9 smooth, bearing anal proleg. Tergites 1–7 with single anterolateral protuberance on each side with single seta, and pair of posterodorsolateral protuberances, each bearing two closely approximated short, thin setae; lateral adhesive structure swelling bearing four setae, two lateral, two basal; additional single seta located anterior to lateral swelling. Segment 8 with dorsolateral protuberance on either side of posterior spiracular plate, each bearing pair of small setae; lateral protuberance with three setae; single short ventrolateral seta; ventral surface bearing pair of setae. Posterior spiracular plate with sclerite encircling procerci; procercus shorter than length of spiracular plate, bearing four setae, two thick, two slender; without cone-like protuberance on either side of procerci. Terminal segment with anterior dorsal protuberance bare; pair of posterior lateral protuberances with pair of setae; five lateral setae; two pairs of long setae on posterior margin, above pair of anal papillae; ventral sternite bearing single pair of setae.
Known only from the type series.
Known from the south-central Andes of Chile (Fig.
The species name is from the Latin acu (needle, pin) in allusion to the needle-like tip of the paramere.
This species was collected at higher elevations than any other species in this publication. Larvae and pupae have ventral adhesive structures and were collected only from rocky substrates (Fig.
Holotype: ♂, glued to point with abdomen in glycerine microvial pinned beneath, labelled: “Chile: Region XIV (Los Ríos)/ Rte. T-85, 13.xii.2016/ 40°19'58.6"S 72°16'56.1"W/ elev. 95 m, roadcut seep, J.K./ Moulton & R.J. Pivar”; “HOLOTYPE/ Niphta/ downesi/ Pivar [red label]” (
This species is recognised by both filaments of the paramere and the aedeagal guide being easily visible in lateral view, giving it a tri-filamentous appearance. Also, the bend in the gonostylus is slightly stronger than that of N. mapuche.
The description of N. downesi differs from that of N. acus in the following regards:
Male. n = 3.
Length 1.9–2.4 mm.
Colouration (Fig.
Head. Frons with two strong setae. Flagellomeres 1–3 subquadrate, expanded, 1.5 × as wide as next segment, as long as 2 and 3 combined.
Thorax. Antealar ridge bearing single pronounced, medial seta flanked by two smaller setae.
Wing. Wing length: 2.2–2.4 mm. C and posterior wing margin with fringe of microtrichia.
Abdomen. Sternites 3–7 with setae restricted to posterior two-thirds and laterad; sternite 8 strongly arched into preceding segment, lacking setae.
Terminalia (Figs
Female. Unknown.
Unknown.
Known only from the type series.
Known from the foothills of the southern Andes in Chile (Fig.
Niphta downesi is named in honour of veterinary and medical entomologist J.A. Downes, who collected the first specimen of this species in 1985.
Austrothaumalea halteralis Edwards, 1930: 114. Stuardo, 1946: 42 (catalogue); Stone, 1966: 1 (catalogue); Arnaud, 1977: 284 (distribution).
Niphta halteris
(Edwards):
Holotype: ♂, minuten pinned with abdomen mounted in resin, labelled: “Casa Pangue./ 4–10.xii.1926.”; “Austrothaumalea/ halteralis Edw./ F.W. Edwards/ det. 1930.”; “S. Chile:/ Llanquihue prov./ F. & M. Edwards./ B.M. 1927 – 63.”; “HOLO-/ TYPE [white label with red margin]”; “NHMUK010210689”.
This species is recognised by its distinct hook-tipped paramere.
The redescriptions of N. halteralis differ from that of N. acus in the following regards:
Male. n = 44.
Length 1.3–2.0 mm.
Colouration (Fig.
Head. Frons with two strong setae. Flagellomeres 1–3 subquadrate, slightly expanded, 0.25 × as wide as next segment, slightly shorter than 2 and 3 combined. Vertex with yellow setae of uniform length, with longer, black orbital setae.
Thorax. Antealar ridge bearing single pronounced medial seta flanked by two smaller setae.
Wing. Wing length: 2.0–2.9 mm. C and posterior wing margin with fringe of microtrichia; R2+3 distinct, situated slightly before middle of R1(+R2+3); M1 and M2 straight.
Abdomen. Sternites 3–7 rectangular, lacking distinct sclerites, setae restricted to posterior half; sternite 8 strongly arched into preceding segment, lacking setae.
Terminalia (Figs
Female. n = 6.
Similar to male except as follows: Abdomen. Tergite 9 noticeably more sclerotised than preceding tergites; sternite 8 well sclerotised.
Terminalia (Fig.
Unknown.
Chile: Region X (Los Lagos): Camino de Penetracion @ Hwy. 7 sign, 16.xii.2013, 42°07'57.5"S 72°27'45.3"W, seep, sweeping veg., G.R. Curler (1♂, 1♀*); Camino de Penetracion @ km post 125.600, 16.xii.2013, 42°03'33.3"S 72°27'07.4"W, rock seep, G.R. Curler (1♂); Casa Pangue, Llanquihue, 12.1926, R&E Shannon, USNMENT01115811 (1♂,
Known from both the Andes and Chilean Coastal Range in southern Chile (Fig.
This species appears restricted predominantly to low elevations in the Valdivian temperate rain forest.
Holotype: ♂, glued to point with abdomen in glycerine microvial pinned beneath, labelled: “Chile: Region IX (Araucanía)/ Rte. S-365, 14.xii.2016/ 38°46'27.0"S 71°36'51.0"W/ elev. 809 m, creek/small falls/ J.K. Moulton & R.J. Pivar”; “HOLOTYPE/ Niphta/ mapuche/ Pivar [red label]” (
This species is recognised by the paramere being mostly hidden within the epandrium in lateral view, giving the paramere a two-filament appearance. The gonostylus is less recurved than that of N. downesi.
The descriptions of N. mapuche differ from that of N. acus in the following regards:
Male. n = 10.
Length 1.4–2.5 mm.
Colouration (Fig.
Head. Flagellomeres 1–3 subquadrate, 1 expanded, 2 × as wide as next segment, as long as 2 and 3 combined.
Abdomen. Abdominal sternite 2 with few setae restricted to laterad on posterior third; sternites 3–7 with setae restricted to lateral margins and middle third; sternite 8 with three or fewer setae medially and on lateral margins.
Terminalia (Figs
Female. n = 10.
Similar to male except as follows: Terminalia (Fig.
Pupa. n = 4 (not figured due to condition of specimens).
Length 3.5–4.0 mm.
Head. Setae not observed.
Thorax. Hindleg concealed behind wing sheath, only apex visible between apex of foreleg and wing sheath, slightly shorter than foreleg, but longer than wing sheath. Wing sheaths with large tubercle at base, setae not visible.
Abdomen. Setae not visible on tergites 1–8.
Larva. n = 17.
Length of final instar 6.3–6.7 mm.
Colouration. Head capsule variable, ranging from light brown to black. Body mottled brown and grey, possibly pale brown to creamy; cream coloured ventrally.
Head capsule (Fig.
Abdomen. Tergites 1–7 with lateral adhesive structure swelling bearing five setae, two lateral, three basal.
Known only from the type series.
Known from the southern Andes of Chile (Fig.
This species is named after the Mapuche (mapu, land, che, people) indigenous peoples, who since ~ 500 B.C., have inhabited the regions of southern Chile, where N. mapuche is known.
This is a mid-elevation species. Adults were observed flying around and resting on leaf tips, roughly two meters from the nearest splash zones. Larvae and pupae have ventral adhesive structures and were collected only from rocky substrates at the margin of a waterfall.
Holotype: ♂, glued to point with abdomen in glycerine microvial pinned beneath, labelled: “Chile: Region XIV (Los Ríos)/ Antilhue, Rte. T-35, 9.xii.2016/ 39°49'09.8"S 72°56'22.6"W/ elev. 40 m, roadside creek,/ J.K. Moulton & R.J. Pivar”; “HOLOTYPE/ Niphta/ bifurcata/ Pivar & Moulton [red label]” (
This species is recognised by the bifurcate, posterior apex of the cheliform gonostylus and the bifurcate anterior projection of the gonocoxite. It is darker in colouration compared to the closely related N. courtneyi.
Male. n = 3.
Length 1.7–1.9 mm.
Colouration (Figs
Head. Eyes above antennae broadly joined, with small triangular frons visible above antennae; frons with two strong setae. Flagellomeres 1–3 subquadrate, 1 expanded, twice as wide as next segment, shorter in length than 2 and 3 combined; flagellomeres 4–10 cylindrical, becoming progressively thinner and elongate. Vertex with black setae of uniform length, with longer, black orbital setae.
Thorax. Mesoscutum with prominent antealar ridge, bearing three setae, middle seta most pronounced. Scutum clothed dorsally in short, black setulae; notopleural, supra-alar and postsutural setae long, black. Pteropleuron bare. All legs with tarsi simple.
Adult lateral habitus micrographs of the Niphta nudipennis group A N. bifurcata sp. nov. (♂) B N. brunnea sp. nov. (♂) C N. courtneyi sp. nov. (♀), abdomen dissected D N. daniellae sp. nov. (♂) E N. eurydactyla sp. nov. (♂) F N. bispinosa sp. nov. (♂), abdomen dissected G N. nudipennis (♂) H N. courtneyi sp. nov. (♂), inset with arrow indicating antealar ridge. Scale bars: 1.0 mm.
Wing. Wing length: 2.1–2.4 mm. Infuscate throughout, apex somewhat narrowed; C fringed in small setulae, with widely spaced microtrichia; posterior wing margin with closely spaced fringe of microtrichia; Sc incomplete; R1 and R1(+R2+3) with three weakenings or depigmented gaps, first slightly beyond R2+3, second and third closely approximated, near C; microtrichia of R1(+R2+3) confined to base near humeral crossvein, remaining veins bare; R flexed into cell br; R2+3 distinct, situated in basal third of R1(+R2+3); bend in R4+5 strong; R4+5 and M1 running parallel toward margin; M1 straight; M2 with gentle bend in apical third; M4 with slight bend.
Adult dorsal habitus micrographs of the Niphta nudipennis group A N. bifurcata sp. nov. (♂) B N. brunnea sp. nov. (♂) C N. courtneyi sp. nov. (♀), abdomen dissected D N. daniellae sp. nov. (♂) E N. eurydactyla sp. nov. (♂) F N. bispinosa sp. nov. (♂), abdomen dissected G N. nudipennis (♂) Scale bars: 1.0 mm.
Abdomen. Abdominal sternite 1 narrow, spectacle-shaped; sternite 2 reduced to slender median sclerite, a few setae restricted to posterior third; sternites 3–7 rectangular, lightly sclerotised, setae restricted to posterior half; sternite 8 strongly reduced, anterior margin well sclerotised, arched slightly into preceding segment, a few setae restricted to laterad.
Terminalia (Figs
Ventral views of female Niphta nudipennis group terminalia A N. bifurcata sp. nov. B N. courtneyi sp. nov. C N. bispinosa sp. nov. D N. nudipennis. Abbreviations: cerc, cercus; gen fk, genital fork; hyp pr, hypogynial protuberance; hyp vlv, hypogynial valve; hypct, hypoproct; lat arm crcl, lateral arm circle; lat arm cmplx, lateral arm complex; spthc p, spermathecal pump. Scale bars: 0.1 mm.
Lateral views of female Niphta nudipennis group terminalia A N. bifurcata sp. nov. B N. courtneyi sp. nov. C N. bispinosa sp. nov. D N. nudipennis. Abbreviations: cerc, cercus; gen fk, genital fork; hyp pr, hypogynial protuberance; hyp vlv, hypogynial valve; hypct, hypoproct; lat arm crcl, lateral arm circle; lat arm cmplx, lateral arm complex; S, sternite; spthc p, spermathecal pump; T, tergite. Scale bars: 0.1 mm.
Female. n = 1.
Similar to male except as follows: Abdomen. Tergite 9 noticeably more sclerotised than preceding tergites; sternite 8 well sclerotised, with distinct blunt projection at base of hypogynial valve. Terminalia (Figs
Unknown.
Known only from the type series.
Known only from the type locality in the Chilean Coastal Range (Fig.
Niphta bifurcata is named in reference to the posterior apex of the gonostylus and the anterior projection of the gonocoxite, both of which are bifurcate.
Holotype: ♂, glued to point with abdomen in glycerine microvial pinned beneath, labelled: “Chile: Region VII (Maule)/ Los Queñes, Rte. J-25,/ 6.xii.2016, 34°59'46.7"S 70°49'19.2"W/ elev. 679 m, cascading creek,/ J.K. Moulton & R.J. Pivar”; “HOLOTYPE/ Niphta/ bispinosa/ Pivar & Sinclair [red label]” (
This species is recognised by the cheliform gonostylus with non-bifurcate apices and the gonocoxite with two projections, the anterior one long and bifurcate, the posterior one small, tooth-like. It is lighter in colouration than the closely related N. brunnea.
Female terminalia of the Niphta halteralis group A lateral, N. acus sp. nov. B ventral, N. acus sp. nov. C lateral, N. halteralis D ventral, N. halteralis E lateral, N. mapuche sp. nov. F ventral, N. mapuche sp. nov. Abbreviations: cerc, cercus; gen fk, genital fork; hyp vlv, hypogynial valve; hypct, hypoproct; lat arm, lateral arm; S, sternite; spthc p, spermathecal pump; T, tergite; v scl, ventral sclerite. Scale bars: 0.1 mm.
The descriptions of N. bispinosa differ from that of N. bifurcata in the following regards:
Male. n = 1.
Length 1.9–2.4 mm.
Colouration (Figs
Head. Frons with three strong setae. Flagellomere 1 expanded, 1.5 × as wide as next segment, equal in length to 2 and 3 combined.
Wing. Wing length: 2.0–2.4 mm. Lightly infuscate throughout; bend in R4+5 gentle.
Terminalia (Figs
Female. n = 2.
Similar to male except as follows: Terminalia (Figs
Unknown.
Known only from the type series.
Known only from the type locality in central Chile (Fig.
Niphta bispinosa is named in reference to the two projections from the gonocoxite.
Holotype: ♂, glued to point with abdomen in glycerine microvial pinned beneath, labelled: “Chile: Region IX (Araucanía)/ Rte. 71, 15.xii.2016/ 38°14'20.6"S 71°53'46.6"W,/ elev. 953 m, roadside seeps,/ J.K. Moulton & R.J. Pivar”; “HOLOTYPE/ Niphta/ brunnea/ Pivar [red label]” (
This species is recognised by the cheliform gonostylus bearing non-bifurcate apices and the gonocoxite with three projections: two that are nearly equal in size and the third, much smaller and inconspicuous, situated at the base of the anterior one. It is darker in colouration compared to the closely related N. bispinosa, and the darkest of the N. nudipennis group.
The descriptions of N. brunnea differ from that of N. bifurcata in the following regards:
Male. n = 5.
Length 1.6–1.9 mm.
Colouration (Figs
Head. Flagellomere 1 expanded, 1.5 × as wide as next segment, shorter in length than 2 and 3 combined.
Wing. Wing length: 1.9–2.2 mm.
Terminalia (Figs
Female. Unknown.
Pupa. n = 6 (Figs
Length 2.7–2.9 mm.
Colouration. Brown; sometimes with black spot above eyes in developing adult.
Head. Maxillary sheath short, posteromedially directed, gently tapered toward truncate apex, apices of palpi separated medially; devoid of tubercles and setae.
Thorax. 1.5 × wider than abdomen at widest point. Foreleg sheath projected straight and slightly beyond wing sheaths, reaching hind margin of sternite 2; anterior half of midleg visible anterior to wing sheath, then hidden behind foreleg, not projected beyond wing sheath; hindleg concealed beneath wing sheath, only small triangular apex visible between apex of foreleg and wing sheath, not extended beyond wing sheath. Wing sheath extended to posterior margin of abdominal sternite 2. Respiratory organ slightly shorter than maxillary sheath, broadest subapically; ovate, slightly arched medially, tapered toward apex; spiracular openings encircling apex; stalk thin, emerging from small tubercle. Tubercle situated posterodorsally to respiratory organ, rounded, projected laterally; apex nearly touching or touching respiratory organ. Thorax devoid of setae.
Abdomen. Subcylindrical, evenly tapered toward caudal segment. Spiracles well developed, distinct on segments 3–7, projected anterodorsolaterally; all bearing minute spine-like setulae. Segment 8 with short lateral projection, less than half length of preceding spiracles, projected dorsolaterally. Tergites 1–8 quadrate, devoid of setae, with pair of dorsolateral ridges and faint medial transverse groove, groove not meeting dorsolateral ridges. Tergite 9 rounded, posterior margin emarginated, laterally compressed compared to preceding segments; small, rounded projection pointing posteriorly in lateral view. Sternites 3–8 rectangular, with row of faint setulae along anterior margin, not connecting to lateral margin; lateral margins crenulate, lacking setae. Sternites 3 and 4 bearing pair of small lateral adhesive structures, sternite 5 bearing pair of large lateral adhesive structures on lateral margin. Caudal sternite triangular, with medial sclerotised groove and pair of medial rounded, posteromedially projected lobes; posterior margin with medial longitudinal ridge; without distinct caudal hooks.
Dorsal views of Niphta larvae A N. brunnea sp. nov. B N. nudipennis C N. acus sp. nov. Abbreviations: A, abdominal segment; Hc, head capsule; Mst, mesothorax; Mtt, metathorax; Pt, prothorax; pro spr, prothoracic spiracle; prtb, protuberance; post spr pl, posterior spiracular plate. Scale bars: 1.0 mm.
Larva. n = 6 (Figs
Length of final instar 4.8–5.1 mm.
Colouration. Head capsule pale brown, anterolateral margin of ecdysial line black. Body creamy brown.
Head capsule (Fig. 28A, B). Two large, circular eye spots, elevated on tubercle; antenna with three finger-like processes, elevated on largest tubercle; with five pairs of smaller tubercles outside of ecdysial lines (not including antennal and ocular tubercle); single tubercle between ecdysial lines, about same size as ocular tubercle; 15 pairs of unbranched setae; six sensory pits (13, 14, 18, 19, 20, 21), sensory pit 13 above antennal tubercle.
Thorax. Prothorax with single pair of protuberances bearing single spiracle; spiracular protuberance bearing one pair of dorsal setae anterior to spiracle and single dorsolateral seta; three mid-lateral setae, two long, one short and fine; two closely approximated setae near base of prothoracic leg (Keilin’s organ). Mesothorax and metathorax with pair of small dorsolateral protuberances and pair of large lateral protuberances; mesothoracic dorsolateral protuberance bearing single thickened seta, metathoracic lateral protuberance bearing pair of closely approximated setae, anterior seta thickened and longer than posterior seta; lateral protuberance on both segments bearing three setae, two short, one long; single long seta slightly ventral to lateral protuberance; one pair of mid-ventrolateral setae. Prothorax bearing proleg, posterior half with rectangular adhesive structure; meso- and metathoracic sternites with rectangular adhesive structure.
Abdomen. Sternites 1–7 modified into circular, suction cup-like adhesive structure; sternite 8 with quadrate adhesive structure; sternite 9 smooth, bearing anal proleg. Segments 1–7 lacking distinct protuberances, at most, pair of dorsolateral swellings bearing single or paired short, thin setae; single seta situated anterolaterally; lateral adhesive structure swelling bearing numerous setae, two anterolateral, two midlateral, four basalateral. Segment 8 with small dorsolateral protuberance on either side of posterior spiracular plate, each bearing pair of small setae; lateral protuberance with single seta; single short ventrolateral seta; ventral sternite bearing pair of setae. Posterior spiracular plate with sclerite encircling procerci; procercus shorter than length of spiracular plate, bearing four setae, two thick, two slender; without cone-like protuberance on either side of procerci. Terminal segment with pair of protuberances, no setae; pair of dorsolateral setae; five lateral setae; two pairs of long setae on posterior margin, above pair of anal papillae; ventral sternite lacking setae.
Known only from the type series.
Known only from two localities in the Andes of southern Chile (Fig.
Niphta brunnea is from the Latin brunneus (brown) in allusion to its brown colouration, the darkest of the N. nudipennis group.
The larvae and pupae both possess the ventral adhesive structures seen in other known immature stages of South American Niphta. Immatures were collected from wetted vegetation in the splash zones, never from rocks (Fig.
Scanning electron micrographs of ventral view of larvae of Niphta brunnea sp. nov. A habitus B head and thoracic segments 1 & 2 C thoracic segments 2 & 3 and abdominal segment 1 D abdominal segments 2 & 3 E abdominal segments 6, 7, 8 & 9. Abbreviations: A, abdominal segment; T, thoracic segment. Scale bars: 1.0 mm.
Holotype: ♂, glued to point with abdomen in glycerine microvial pinned beneath, labelled: “Chile, Region X (Los Lagos)/ East side Lago Llanquihue/ small falls on road (nr PN/ VPR) 41°08.47'S 72°35.28'W/ ≈ 100 m 2.xii.2008 GW/ Courtney (CH08–30)”; “HOLOTYPE/ Niphta/ courtneyi/ Pivar [red label]” (
This species is recognised by the bifurcate posterior apex of the cheliform gonostylus and the presence of three gonocoxal projections. It is lighter in colouration than the closely related N. bifurcata.
The descriptions of N. courtneyi differ from that of N. bifurcata in the following regards:
Male. n = 1.
Length 2.1–2.3 mm.
Colouration (Figs
Head. Frons with three strong setae. Flagellomere 1 expanded, 1.5 × as wide as next segment, shorter in length than 2 and 3 combined.
Wing. Wing length: 1.9–2.2 mm. Lightly infuscate throughout.
Terminalia (Figs
Female. n = 2.
Similar to male except as follows: Terminalia (Figs
Unknown.
Known only from the type series.
Known only from the type locality, the East side of Lago Llanquihue in Southern Chile (Fig.
Niphta courtneyi is named in honour of its collector, Gregory W. Courtney (Iowa State University). Courtney collected three new species of Thaumaleidae from Chile (A. fredericki Pivar, N. courtneyi, and N. mapuche), as well as immature material, prompting us to further investigate the Chilean fauna.
Holotype: ♂, glued to point with abdomen in glycerine microvial pinned beneath, labelled: “Chile: Region IX (Araucanía)/ Rte. 71, 15.xii.2016/ 38°14'20.6"S 71°53'46.6"W/ elev. 953 m, roadside seeps/ J.K. Moulton & R.J. Pivar”; “HOLOTYPE/ Niphta/ daniellae/ Pivar [red label]” (
Habitat and larvae of Niphta acus sp. nov. (36°55'02.7"S 71°25'49.6"W) A Moulton shown next to the falls for scale B close up of falls with box indicating where immatures were captured (Note: they were not found in high flow zones) C lateral view of larva, with adhesive structures visibly in contact with substrate D dorsal view of larva illustrating camouflage E larva and pupal exuviae on rock face.
This species is recognised by the sharply pointed, tapered gonostylus with no projections and not cheliform, unlike all remaining species in the N. nudipennis group. The gonocoxal plate also has a pair of lateral arms projected anteriorly.
The description of N. daniellae differs from that of N. bifurcata in the following regards:
Male. n = 2.
Length 1.7–2.0 mm.
Colouration (Figs
Habitat and immatures of members of the Niphta nudipennis group A type locality of N. brunnea sp. nov. and N. daniellae sp. nov., box indicating where plant stem in images B, C was taken from (38°14'20.6"S 71°53'46.6"W) B larvae of N. brunnea sp. nov. on plant stem from splash zone C close up of N. brunnea sp. nov. larva, adhesive structure visibly in contact with substrate D habitat of N. nudipennis, box indicating foliage in splash zone where immatures were found E pupa of N. nudipennis affixed to leaf with final instar larval exuviae visible.
Head. Frons with two to three strong setae. Flagellomere 1 expanded, 1.5 × as wide as next segment, subequal in length to 2 and 3 combined.
Wing. Wing length: 2.2–2.3 mm. Lightly infuscate throughout; bend in R4+5 gentle; M4 with slight apical bend.
Abdomen. Abdominal sternite 2 reduced to slender median sclerite, lacking setae; sternites 3–7 rectangular, setae restricted to posterior two-thirds; sternite 8 strongly reduced, lacking setae.
Terminalia (Figs
Female. Unknown.
Unknown.
Known only from the type series.
Known from two localities in the Andes of south-central Chile (Fig.
This species is named in honour of RJP’s wife, Danielle Lombardi, for her support during Pivar’s graduate research and entomological endeavours, and for playing an important role in organising the Chilean expedition. Raised in northern Chile (Arica), Danielle’s Spanish skills were critical for translating all communications with government and national park contacts, as well as translating our requests for collecting permits.
Holotype: ♂, glued to point with abdomen in glycerine microvial pinned beneath, labelled: “Chile: Region X (Los Lagos)/Rte. U-99, 10.xii.2016/ 41°08'28.2"S 72°35'16.8"W/ elev. 101 m, roadside seeps/ and creek, J.K. Moulton & R.J./ Pivar”; “HOLOTYPE/ Niphta/ eurydactyla/ Pivar [red label]” (
Examples of thaumaleid habitats in Chile A waterfalls where Niphta downesi sp. nov., N. eurydactyla sp. nov. and N. nudipennis were collected by sweeping adjacent foliage (40°19'58.7"S 72°16'54.8"W) B collection site of N. bispinosa sp. nov. (34°59'48.8"S 70°48'37.0"W) (after Pivar et al. 2020) C type locality of N. acus sp. nov., specimens were mainly collected from plants on left (37°46'30.8"S 71°42'03.9"W) D rock face seep, type locality of N. bispinosa sp. nov. (34°59'46.7"S 70°49'19.2"W) E Chiloé, N. nudipennis was collected at a small waterfall next to the larger Cascada Tocoihue, Pivar is shown for scale (42°18'20.3"S 73°26'08.9"W); F, Puente El Salto, N. halteralis was collected above the falls, Pivar is shown above the falls for scale (41°31'29.2"S 72°17'14.6"W).
This species is recognised by a broad, straight, finger-like projection on the gonostylus.
The description of N. eurydactyla differs from that of N. bifurcata in the following regards:
Male. n = 17.
Length 1.5–1.8 mm.
Colouration (Figs
Thorax. Antealar ridge bearing three to four setae, middle seta most pronounced.
Wing. Wing length: 1.8–2.2 mm.
Abdomen. Abdominal sternites with setae restricted to posterior two-thirds.
Terminalia (Figs
Female. Unknown.
Unknown.
Known only from the type series.
Known from both the Chilean Coastal Range and Andes of southern Chile (Fig.
The specific name is from the Greek eury (broad, wide) and daktylos (finger), in allusion to the broad, finger-like projection on the gonostylus.
Austrothaumalea nudipennis Edwards, 1930: 113. Stuardo, 1946: 42 (catalogue); Stone, 1966: 1 (catalogue); Arnaud, 1977: 284 (distribution).
Niphta nudipennis
(Edwards):
Holotype: ♂, minuten pinned with abdomen mounted in resin, labelled: “Ancud./ 17–19.xii.1926.”; “Austrothaumalea/ nudipennis Edw./ F.W. Edwards/ det. 1930.”; “S. Chile:/ Chiloe I./ F. & M. Edwards./ B.M. 1927 – 63.”; “HOLO-/ TYPE [white label with red margin]”; “NHMUK010210690”. Allotype: ♀, same label data as holotype (NHMUK). Paratypes: same data as holotype (4♂, 4♀, NHMUK).
This species is recognised by a long, narrow, finger-like projection on the gonostylus that has a medial bend.
The adult and immature descriptions of N. nudipennis differ from those of N. bifurcata and N. brunnea, respectively in the following regards:
Male. n = 22.
Length 1.5–2.3 mm.
Colouration (Figs
Head. Flagellomere 1 subequal in length to 2 and 3 combined.
Wing (Fig.
Abdomen. Abdominal sternite 2 with a few setae restricted to laterad on posterior third; sternites 3–7 with setae restricted to posterior two-thirds; sternite 8 lacking setae.
Terminalia (Figs
Female. n = 6.
Similar to male except as follows: Abdomen. Tergite 9 noticeably more sclerotised than preceding tergites; sternite 8 well sclerotised, with distinct blunt projection at base of hypogynial valve.
Terminalia (Figs
Pupa. n = 4 (Figs
Length 3.0–3.1 mm.
Colouration. Light brown; with black spot above eyes in developing adult.
Thorax. 1.25 × wider than abdomen at widest point. Foreleg sheath projected straight, slightly shorter than wing sheaths; anterior half of midleg visible anterior to wing sheath, then hidden behind foreleg, apices visible, slightly longer than foreleg; hindleg concealed beneath wing sheath, only apex visible between apex of foreleg and wing sheath, longer than foreleg, extended slightly beyond wing sheath but not reaching hind margin of sternite 2. Wing sheaths not reaching posterior margin of abdominal sternite 2. Respiratory organ slightly longer than maxillary sheath, broadest subapically. Tubercle situated posterodorsally to respiratory organ, rounded, projected slightly posterolaterally; apex well separated from respiratory organ. Thorax devoid of setae.
Abdomen. Sternite 8 with small, indistinct lateral projection, directed slightly anterolaterally. Tergites 1–8 quadrate, devoid of setae, with pair of dorsolateral ridges (indistinct on tergites 1–6). Caudal sternite subquadrate, lacking medial lobes; posterior margin with medial longitudinal groove; without distinct caudal hooks.
Larva. n = 27 (Figs
Length of final instar 4.7–5.4 mm.
Colouration. Head capsule pale brown (sometimes dark brown). Body creamy brown.
Head capsule (Fig.
Thorax. Spiracular protuberance bearing one pair of dorsal setae anterior to spiracle and single lateral seta.
Abdomen. Segments 1–7 with lateral adhesive structure inflatted bearing four setae, two midlateral, two basalateral. Terminal segment with pair of protuberances bearing pair of setae; four lateral setae, two long, two short and fine.
Chile: Region X (Los Lagos): Chiloé, Cascada Tocoihue, 10.xii.2016, 42°18'20.3"S 73°26'08.9"W, elev. 32 m, smaller falls, J.K. Moulton & R.J. Pivar (2♂, 4♀*); Ensenada, nr. Baños de Petrohué, 12.i.1985, J.A. Downes (13♂, 2♀,
Known from both the Chilean Coastal Range and the Andes of southern Chile (Fig.
Niphta nudipennis is a low-elevation species inhabiting the Valdivian temperate rainforest. Adults were collected mainly from foliage around splash zones (Figs
Each group’s morphology is well adapted to their microhabitat. On a rock face, immatures have to contend with debris being washed down from the substrate above. Pupae of the N. halteralis group were collected from exposed microhabitats on the rock face, with no protection from flowing debris. Their stout and stocky body shape, hidden spiracles and small respiratory organs that do not extend far from the body, likely help in withstanding any potential debris impact and reduce breakage of exposed appendages. Both pupae and larvae are mottled and much darker in colouration than the pupae of the N. nudipennis group, offering greater camouflage from predators on the exposed rock face. Larvae have the added vestiture of dorsal tubercles, which help to break up the outline of the body. Immatures of the N. halteralis group also bear more setae than species of the N. nudipennis group, perhaps for sensing debris or predators that may be nearby.
Immatures of the N. nudipennis group were all collected from plant material, either living or dead, but always directly in the splash zone. They were collected from both smooth leaf surfaces and more textured vegetation, such as ferns and herbaceous plant stems. They were found on both upper and lower leaf surfaces, depending on which surface was in the splash zone. Contrary to a rock face, the amount of debris flowing down a leaf is likely minimal. Pupae of the N. nudipennis group are conoid, with a broad head and thorax and a narrow abdomen, protruding spiracles, and large, laterally projecting respiratory organs. All of these features are indicative of life in a habitat where debris does not pose a problem of displacement, breakage, or blockage of the respiratory organ. The leaf habitats were generally shaded and hidden from direct view, and coupled with constant splashing, individuals may experience less predation than on a rock face, thereby reducing the need for the dark, mottled colouration and tubercles. Eggs on vegetation were not observed, nor was oviposition by the adults, but they are presumably laid on the vegetation surface.
The presence of the adhesive structures is associated with behavioural traits that differ from those of other Thaumaleidae. The typical thaumaleid larva will exhibit a characteristic quick, sidewinding motion to evade predation (
Despite the discovery of this new larval morphotype, specimens were never collected in areas of extremely high flow or aggressive splash zones; rather, they were collected in typical thaumaleid habitat consisting of slow flowing, thin films of water. Perhaps antecedents originated in environments with higher water flow, such as in a river, and the extant species are a result of having adapted to the more familiar recent habitats. Alternatively, these slow flowing zones may be subject to torrential flooding after rainfall. The continued presence of the adhesive structures suggests a continuing evolutionary advantage in their present-day habitat.
Immatures of the Australian N. collessi (
The pupa of N. collessi has caudal hooks, much like in Neothaumalea atlantica; however, the Chilean species lack caudal hooks or any other projections. There is also a significant reduction in setae, both in number and length. Niphta collessi has numerous setae, many of which are long, while members of the N. halteralis group have very few, short setae. Pupae of the N. nudipennis group lack abdominal setae altogether. The respiratory organs of N. collessi are similar in appearance to those of the N. nudipennis group, where they are broadest subapically and project laterally, whereas those in the N. halteralis group are shorter and barely project laterally. The spiracles of N. collessi and the N. nudipennis group are also similar; well developed on sternites 3–7 and mounted on long, narrow, lateral tubercles. The spiracles are barely visible in the N. halteralis group.
The larvae also exhibit some significant differences and similarities. The most apparent difference, aside from the adhesive structures, is in the sculpture of the head capsule. All described Chilean Niphta have protuberances and large antennal tubercles, similar to those of Neothaumalea atlantica; Niphta collessi lacks protuberances and the antennae are on short tubercles. Also, much as in the pupae, Chilean Niphta larvae exhibit a reduction in setae compared to N. collessi. Another significant difference is the presence of cauliflower-like protuberances on both thoracic and abdominal tergites. These protuberances are especially prominent in the N. halteralis group, and are reduced but still present in Ni. nudipennis group. Based on
Larvae with ventral adhesive structures have been collected in Australia. Though they have not been reared, nor identification confirmed via DNA fingerprinting (attempts to fingerprint were made, but with no success), morphology indicates these larvae are likely a member of Niphta. They appear very similar to members of the N. halteralis group: presence of distinct thoracic and abdominal protuberances, darker colouration, long abdominal setae (longer and more abundant than N. halteralis group) and pronounced head-capsule protuberances (not as large as N. halteralis group, but larger than N. nudipennis group). The adhesive structures appear very similar between all species: thoracic segments rectangular, adhesive structures felt-like; abdominal segments 1–7 circular, margins felt-like with smooth interior; abdominal segment 8 quadrate, felt-like (Fig.
South American thaumaleid diversity has now increased to 17 described species. Additionally, there is the likely new species to be discovered in Ecuador, plus three more new species from Chile that were collected by the authors, but not yet described because they are represented by females only. Since males possess readily recognised genitalic features, the authors have decided to wait until further material is collected before describing these species. Both morphological and molecular data (Pivar, unpublished data) support the presence of these new species. Of the described South American species, only Neothaumalea atlantica is not found in the Andes Range and is recorded from the Atlantic Forest of southeastern Brazil, in the Serra Geral mountains (
In Chile, thaumaleid diversity increases as one moves south. Beginning in central Chile and progressing south, the following are the known diversity of species per region (including the three undescribed females): Valparaíso (1 sp.), Santiago (2 spp.), O’Higgins (1 sp.), Maule (2 spp.), Bío Bío (5 spp.), Araucanía (7 spp.), Los Ríos (6 spp.), Los Lagos (8 spp.) and Aysén (1 sp.). These numbers are reflective of both the amount of time spent collecting in certain regions and regions sampled; some regions have likely never been sampled (particularly in northern Chile). Available habitat is also a large contributor to diversity. From Bío Bío to Los Lagos, specimens were collected in Valdivian temperate rainforests, characterised by high rainfall and cooler temperatures. Vegetation types include southern beech, laurel and broadleaved forest, bamboo and ferns. With an abundance of mountain streams and waterfalls, similar to the Nearctic Pacific Northwest (
Most Chilean species are found in temperate rainforest regions, but some are found in both wet and dry climates (A. chilensis Edwards, N. acus). The type of madicolous habitat (i.e., creek, rock face seep, stream, waterfall, etc.) (Fig.
Future collections should focus on all areas of the Andes, with an emphasis on the northern sections to provide insight into the northern limits of the family and genera. Currently, the southern-most Nearctic species is Androprosopa zempoala Sinclair and Huerta from central Mexico (
The authors would like to thank colleagues who provided fresh material for this study: Gregory Courtney, Greg Curler and Isai Madriz. Additionally, Duncan Sivell (NHMUK), Torsten Dikow (