Research Article |
Corresponding author: Thomas Kaltenbach ( thomas.kaltenbach@bluewin.ch ) Academic editor: Ben Price
© 2018 Thomas Kaltenbach, Jean-Luc Gattolliat.
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:
Kaltenbach T, Gattolliat J-L (2018) The incredible diversity of Labiobaetis Novikova & Kluge in New Guinea revealed by integrative taxonomy (Ephemeroptera, Baetidae). ZooKeys 804: 1-136. https://doi.org/10.3897/zookeys.804.28988
|
Material collected between 1999 and 2011 in Papua New Guinea and the Papua Province of Indonesia unveiled the enormous diversity of Labiobaetis on this island. Twenty-six new species were identified and delimited by integrative taxonomy using genetic distance (COI, Kimura-2-parameter) and morphology. These new species are described and illustrated based on larvae, augmenting the total number of Labiobaetis species on the island of New Guinea to 32. Seven morpho-groups of species are proposed based on morphological characters and a key to all New Guinea species is provided. The generic attributes of the larvae are summarised and slightly modified based on the examinations of the new species. Results on the genetics of most species (COI) are also provided. The interspecific K2P distances are between 13% and 32%, the intraspecific distances usually between 0% and 2%. Possible reasons for the remarkable richness of this genus in New Guinea are discussed.
COI, Indonesia, morphology, new species, Papua New Guinea, species delimitation
The family Baetidae has the highest species diversity among mayflies, comprising more than 1000 species in 104 genera, which is approximately one quarter of all mayfly species worldwide (
The genus Labiobaetis Novikova & Kluge, 1987 is one of the richest genera of Baetidae with 79 described species (
Originally Labiobaetis was established as a subgenus of Baetis Leach (
Up to now there was only one study on Labiobaetis s. l. under the genus name Pseudocloeon Klapálek, 1905 from New Guinea (
The specimens were mainly collected by Michael Balke (Zoologische Staatssammlung München,
The specimens were preserved in 70%–96% ethanol. The dissection of larvae was done in Cellosolve (2-Ethoxyethanol) under Olympus SZX7 stereomicroscope and mounted on slides with Euparal liquid.
The DNA of part of the specimens was extracted by using non-destructive methods, which allows for subsequent morphological analysis (see
Drawings were made using an Olympus BX43 microscope. Photographs of larvae were taken with a Canon EOS 6D camera and the Visionary Digital Passport imaging system (http://www.duninc.com) and processed with the programs Adobe Photoshop Lightroom (http://www.adobe.com) and Helicon Focus version 5.3 (http://www.heliconsoft.com). Photographs were subsequently enhanced with Adobe Photoshop Elements 13.
The distribution map was generated with the program Simple Mapper (http://research.amnh.org/pbi/maps), the program GEOLocate (http://www.museum.tulane.edu/geolocate/web/WebGeoref.aspx) as well as Google Earth (http://www.google.com/earth/download/ge/) were used to attribute approximate GPS coordinates to sample locations of
The taxonomic descriptions and the key presented herein were generated with a DELTA (
The new species described in this study were all compared to paratypes (on slides) of the six known species from New Guinea (deposited in Purdue University, West Lafayette, Indiana, USA).
For the terminology, we are referring to
Species | Locality | Specimen catalog # | GenBank # (COI) | GenSeq Nomenclature |
---|---|---|---|---|
L. balkei sp. n. | Papua New Guinea, Central Prov. | GBIFCH 00465156 | MH619492 | genseq-1 COI |
GBIFCH 00465157 | MH619493 | genseq-2 COI | ||
L. lobatus sp. n. | Papua New Guinea, Central Prov. | GBIFCH 00508141 | MH619503 | genseq-1 COI |
L. michaeli sp. n. | Papua New Guinea, Eastern Highlands | GBIFCH 00508129 | MH619477 | genseq-1 COI |
GBIFCH 00508130 | MH619478 | genseq-2 COI | ||
GBIFCH 00508134 | MH619484 | genseq-2 COI | ||
L. claudiae sp. n. | Papua New Guinea, Madang Prov. | GBIFCH 00508144 | MH619479 | genseq-1 COI |
L. stagnum sp. n. | Indonesia, Papua Prov. | GBIFCH 00465168 | MH619491 | genseq-2 COI |
L. orientis sp. n. | Papua New Guinea, Eastern Highlands | GBIFCH 00465169 | MH619496 | genseq-1 COI |
L. papuaensis sp. n. | Indonesia, Papua Prov. | GBIFCH 00465170 | MH619502 | genseq-1 COI |
L. gladius sp. n. | Papua New Guinea, Western Highlands | GBIFCH 00465179 | MH619486 | genseq-4 COI |
L. janae sp. n. | Indonesia, Papua Prov. | GBIFCH 00465181 | MH619483 | genseq-1 COI |
GBIFCH 00465182 | MH619489 | genseq-2 COI | ||
L. branchiaesetis sp. n. | Papua New Guinea, Eastern Highlands | GBIFCH 00465183 | MH619480 | genseq-1 COI |
L. planus sp. n. | Indonesia, Papua Prov. | GBIFCH 00508149 | MH619485 | genseq-2 COI |
GBIFCH 00508150 | MH619487 | genseq-2 COI | ||
L. podolakae sp. n. | Papua New Guinea, Eastern Highlands | GBIFCH 00465194 | MH619500 | genseq-2 COI |
L. schwanderae sp. n. | Papua New Guinea, Gulf Prov. | GBIFCH 00465197 | MH619501 | genseq-1 COI |
L. altus sp. n. | Papua New Guinea, Enga Prov. | GBIFCH 00508131 | MH619481 | genseq-2 COI |
L. gindroi sp. n. | Indonesia, Papua Prov. | GBIFCH 00465203 | MH619490 | genseq-2 COI |
L. paravitilis sp. n. | Papua New Guinea, Madang Prov. | GBIFCH 00508148 | MH619482 | genseq-2 COI |
L. paravultuosus sp. n. | Papua New Guinea, Enga Prov. | GBIFCH 00465213 | MH619498 | genseq-1 COI |
GBIFCH 00465214 | MH619499 | genseq-2 COI | ||
L. centralensis sp. n. | Papua New Guinea, Central Prov. | GBIFCH 00465215 | MH619495 | genseq-1 COI |
GBIFCH 00465216 | MH619494 | genseq-2 COI | ||
L. elisae sp. n. | Papua New Guinea, Western Highlands | GBIFCH 00465219 | MH619497 | genseq-1 COI |
L. vallus sp. n. | Papua New Guinea, Madang Prov. | GBIFCH 00465226 | MH619488 | genseq-1 COI |
Abbreviations:
balkei group
1. L. balkei sp. n.
2. L. lobatus sp. n.
3. L. michaeli sp. n.
claudiae group
4. L. claudiae sp. n.
5. L. stagnum sp. n.
orientis group
6. L. orientis sp. n.
7. L. papuaensis sp. n.
petersorum group
8. L. petersorum
9. L. gladius sp. n.
10. L. janae sp. n.
tuberpalpus group
11. L. tuberpalpus
12. L. branchiaesetis sp. n.
13. L. magnovaldus sp. n.
14. L. planus sp. n.
15. L. podolakae sp. n.
16. L. rutschmannae sp. n.
17. L. schwanderae sp. n.
vitilis group
18. L. vitilis
19. L. altus sp. n.
20. L. gindroi sp. n.
21. L. paravitilis sp. n.
22. L. wilhelmensis sp. n.
vultuosus group
23. L. vultuosus
24. L. paravultuosus sp. n.
not assigned to a group
25. L. centralensis sp. n.
26. L. dendrisetis sp. n.
27. L. elisae sp. n.
28. L. inopinatus sp. n.
29. L. involutus
30. L. pindaundensis sp. n.
31. L. vallus sp. n.
32. L. xeniolus
Labiobaetis, states of selected characters: a-f setae of the labrum dorsal, submarginal arc of setae a simple b feathered c dendritic d spatulate e lanceolate f lanceolate with apical pectination; g–k labial palp, distomedial protuberance of segment II and shape of segment III, g lobed and slightly pentagonal h compact, rounded and conical i hook-like and slightly pentagonal j thumb-like elongated and about semicircular k short thumb-like and oblong; l-m left mandible, l outermost denticle blade-like m denticles unmodified; n–p maxillary palp segment II, n with distolateral excavation o with slight distolateral excavation p without distolateral excavation; q–s paraproct, q distally expanded r distally slightly expanded s distally not expanded.
The group is characterised by a large, lobed, distomedial protuberance of labial palp segment II and a dorsal, submarginal arc of setae of the labrum composed of spatulate, apically pectinate setae.
Larva. Following combination of characters: A) labrum dorsal submarginal arc of setae composed of 10–11 spatulate, apically pectinate setae; B) labial palp segment II with a large, lobed distomedial protuberance; C) fore femur length ca. 3× maximum width, dorsally with a row of ca. 17 curved, spine-like setae on margin and a few curved, spine-like setae near proximal margin ; D) tibia dorsally with stout, spatulate, apically rounded setae along margin; E) claw with a row of 11–12 denticles; F) paraproct distally expanded.
Larva (Figs
Colouration. Head, thorax and abdomen dorsally brown, thorax with brighter, faint pattern as in Fig.
Antenna with scape and pedicel sub-cylindrical, without distolateral process at scape; flagellum with lanceolate spines on apex of each segment.
Labrum (Fig.
Right mandible (Fig.
Left mandible (Fig.
Both mandibles with lateral margins almost straight. Basal half with fine, simple setae scattered over dorsal surface.
Hypopharynx (Fig.
Maxilla (Fig.
Labium (Fig.
Hind wing pads absent.
Foreleg (Fig.
Tergum (Fig.
Gills (Fig.
Paraproct (Fig.
Dedicated to Michael Balke (Zoologische Staatssammlung München,
New Guinea.
The specimens were collected at an altitude of 1400 m a.s.l.
Holotype. Nymph (on slide, GBIFCH 00465156), Papua New Guinea, Central, Kokoda Trek, 1400 m, Jan 2008, 09°01.95'S, 147°44.46'E, Posman (PNG 172). Deposited in
Larva. Following combination of characters: A) labrum dorsal, submarginal arc of setae composed of 15 spatulate, apically pectinate setae; B) labial palp segment II with a large, lobed distomedial protuberance; C) fore femur length ca. 4× maximum width, dorsal margin with a row of ca. 16 curved, spine-like setae and a few spine-like setae near margin; D) fore claw with a row of 13–14 denticles; E) paraproct distally expanded.
Larva (Figs
Colouration. Head, thorax and abdomen dorsally brown, thorax with bright, faint pattern as in Fig.
Antenna with scape and pedicel sub-cylindrical, without distolateral process at scape; flagellum with lanceolate spines and fine, simple setae on apex of each segment.
Labrum (Fig.
Right mandible (Fig.
Left mandible (Fig.
Both mandibles with lateral margins almost straight. Basal half with fine, simple setae scattered over dorsal surface.
Hypopharynx (Fig.
Maxilla (Fig.
Labium (Fig.
Hind wing pads absent.
Foreleg (Fig.
Tergum (Fig.
Gills (Fig.
Paraproct (Fig.
Refers to the large, lobed, distomedial protuberance of labial palp segment II.
New Guinea.
The specimens were collected at an altitude of 1390 m a.s.l.
Holotype. Nymph (on slide, GBIFCH 00508141), Papua New Guinea, Central, Kokoda Trek, 1390 m, Jan 2008, 09°00.34'S, 147°44.25'E, Posman (PNG 173). Deposited in
Larva. Following combination of characters: A) labrum dorsal, submarginal arc of setae composed of 14–16 spatulate, apically pectinate setae; B) labial palp segment II with a large, lobed distomedial protuberance; C) fore femur rather broad, length ca. 3× maximum width, dorsal margin with a row of ca. 19 curved, spine-like setae and a few spine-like setae near margin; D) fore claw with 11–12 denticles; E) paraproct distally slightly expanded.
Larva (Figs
Colouration. Head, thorax and abdomen dorsally brown, thorax with bright, faint pattern as in Fig.
Antenna with scape and pedicel sub-cylindrical, without distolateral process at scape; flagellum with lanceolate spines and fine, simple setae on apex of each segment.
Labrum (Fig.
Right mandible (Fig.
Left mandible (Fig.
Both mandibles with lateral margins almost straight. Basal half with fine, simple setae scattered over dorsal surface.
Hypopharynx (Fig.
Maxilla (Fig.
Labium (Fig.
Hind wing pads absent.
Foreleg (Fig.
Tergum (Fig.
Gills (Fig.
Paraproct (Fig.
Dedicated to Michael Balke (Zoologische Staatssammlung München,
New Guinea.
The specimens were collected at an altitude of 2200 m a.s.l.
Holotype. Nymph (on slide, GBIFCH 00508129), Papua New Guinea, Eastern Highlands, Akameku-Brahmin, Bismarck Range, 2200 m, 23 Nov 2006, 05°56.80'S, 145°22.24'E, Balke & Kinibel (PNG 106). Deposited in
The group is characterised by an elongated, thumb-like protuberance of labial palp segment II and a dorsal, submarginal arc of setae composed of simple setae. Additionally, the gills have both longer and shorter fine, simple setae at the margin and the femoral patch is well developed.
Larva. Following combination of characters: A) labrum dorsal submarginal arc of setae composed of one plus five long, simple setae; B) labial palp segment II with an elongated thumb-like distomedial protuberance, segment III subrectangular; C) fore femur rather broad, length ca. 3× width, dorsal margin with a row of ca. 27 curved, spine-like setae; D) gills with small denticles and both medium and long, fine simple setae on margin; E) paraproct distally not expanded, with ca. 18 marginal, stout spines.
Larva (Figs
Colouration. Head, thorax and abdomen dorsally brown, head and thorax with bright median, dorsal suture, thorax with bright pattern as in Fig.
Antenna with scape and pedicel sub-cylindrical, without distolateral process at scape; flagellum with broad, lanceolate spines and fine, simple setae on apex of each segment.
Labrum (Fig.
Right mandible (Fig.
Left mandible (Fig.
Both mandibles with lateral margins almost straight. Basal half with fine, simple setae scattered over dorsal surface.
Hypopharynx (Fig.
Maxilla (Fig.
Labium (Fig.
Hind wing pads absent.
Foreleg (Fig.
Tergum (Fig.
Gills (Fig.
Paraproct (Fig.
Dedicated to Claudia Kaltenbach, the wife of one of the authors (TK) for her constant support during the study.
New Guinea.
The specimens were collected in a ford at an altitude of 80 m a.s.l.
Holotype. Nymph (on slide, GBIFCH 00508144), Papua New Guinea, Madang, highway nr Madang, ford, 80 m, 26 Nov and 2–3 Dec 2006, 05°24.41'S, 145°38.21'E, Binatang Boys (PNG 117). Deposited in
Larva. Following combination of characters: A) labrum dorsal submarginal arc of setae composed of one plus six long, simple setae; B) maxillary palp about as long as length of galea-lacinia; C) labial palp segment II with an elongated, thumb-like distomedial protuberance; D) fore femur rather broad, length ca. 3× maximum width, dorsal margin with a row of ca. 18 curved, spine-like setae on margin and a few stout, pointed setae near margin; E) fore claw with 10–12 denticles.
Larva (Figs
Colouration. Head, thorax and abdomen dorsally brown, head and thorax with bright median, dorsal suture, head, thorax and abdomen with bright pattern as in Fig.
Antenna with scape and pedicel sub-cylindrical, without distolateral process at scape; flagellum with broad, lanceolate spines and fine, simple setae on apex of each segment.
Labrum (Fig.
Right mandible (Fig.
Left mandible (Fig.
Both mandibles with lateral margins almost straight. Basal half with fine, simple setae scattered over dorsal surface.
Hypopharynx (Fig.
Maxilla (Fig.
Labium (Fig.
Hind wing pads absent.
Foreleg (Fig.
Tergum (Fig.
Gills (Fig.
Paraproct (Fig.
Latin word for pool, refers to the appearance in stream pools.
New Guinea.
The specimens were collected in stream pools at an altitude of 115 m a.s.l.
Holotype. Nymph (on slide, GBIFCH 00465166), Indonesia, Papua, Setani-Maribu, stream pools, 115 m, 14 Oct 2011, 02°30.51'S, 140°22.83'E, Balke (PAP01). Temporary deposited in
The group is characterised by a large, lobed, distomedial protuberance of labial palp segment II and a dorsal, submarginal arc of setae of the labrum composed of feathered setae.
Larva. Following combination of characters: A) labrum dorsal submarginal arc of setae composed of 14 long, feathered setae; B) labial palp segment II with a large, lobed distomedial protuberance and with segment III slightly pentagonal and slightly pointed on apex; C) hind femur slender, length ca. 4× maximum width, dorsal margin with a row of ca. 18 curved, spine-like setae and a few stout, pointed setae near margin; D) femur and tibia of hind leg ventrally with stout, bipectinate, pointed setae; E) hind tarsal claw with 14 denticles.
Larva (Figs
Colouration. Unknown.
Antenna with scape and pedicel sub-cylindrical, without distolateral process at scape; flagellum with lanceolate spines and fine, simple setae on apex of each segment.
Labrum (Fig.
Right mandible (Fig.
Left mandible (Fig.
Both mandibles with lateral margins almost straight. Basal half with fine, simple setae scattered over dorsal surface.
Hypopharynx (Fig.
Maxilla (Fig.
Labium (Fig.
Hind wing pads absent.
Hind leg (Fig.
Tergum (Fig.
Gills (Fig.
Paraproct (Fig.
Refers to the type locality, which is in the Eastern Highlands of Papua New Guinea.
New Guinea.
The specimen was collected at an altitude of 1700–1800 m a.s.l.
Holotype. Nymph (on slide, GBIFCH 00465169), Papua New Guinea, Eastern Highlands, Marawaka, Ande, 1700–1800 m, 09 Nov 2006, nr 07°01.70'S, 145°49.81'E, Balke & Kinibel (PNG 87). Deposited in
Larva. Following combination of characters: A) labrum dorsal submarginal arc of setae composed of 14–17 feathered setae, B) labial palp segment II with a large, lobed distomedial protuberance; C) labial palp segment III slightly pentagonal, apically slightly pointed; D) fore femur very slender, length ca. 5× maximum width, dorsal margin with a row of ca. 13 robust, slightly spatulate, apically blunt setae; E) tibia dorsal margin with a row of stout, lanceolate, apically rounded setae.
Larva (Figs
Colouration. Head, thorax and abdomen dorsally brown, with bright pattern as in Fig.
Antenna with scape and pedicel sub-cylindrical, without distolateral process at scape; flagellum with lanceolate spines on apex of each segment.
Labrum (Fig.
Right mandible (Fig.
Left mandible (Fig.
Both mandibles with lateral margins almost straight. Basal half with fine, simple setae scattered over dorsal surface.
Hypopharynx. Unknown.
Maxilla (Fig.
Labium (Fig.
Hind wing pads absent.
Foreleg (Fig.
Tergum (Fig.
Gills (Fig.
Paraproct (Fig.
Refers to the type locality in the Papua Province of Indonesia.
New Guinea.
The specimens were collected at an altitude of 774 m a.s.l.
Holotype. Nymph (on slide, GBIFCH 00465170), Indonesia, Papua, Road Nabire-Enarotali KM 55, 774 m, 22 Oct 2011, 03°29.80'S, 135°43.89'E, Balke (PAP09). Temporary deposited in
The group is characterised by mandibles with an outermost blade-like enlarged denticle, a short and rounded distomedial protuberance of labial palp segment II and a dorsal, submarginal arc of setae of the labrum composed of simple setae.
Larva. Following combination of characters: A) labrum dorsal arc of setae composed of one plus 6–7 long, simple setae; B) both mandibles with outermost denticle blade-like; C) labial palp as Fig.
Paratypes. 4 nymphs (on slides, PERC 0 012 564, PERC 0 012 565, PERC 0 012 566, PERC 0 012 567), Papua New Guinea, Bulolo R, 2950 ft, E of Wau, 15 Oct 1964, W.L. and J.G. Peters.
Larva. Following combination of characters: A) labrum dorsal arc of setae composed of one plus 9–11 long, simple setae; B) both mandibles with outermost denticles blade-like; C) hypopharynx with concave distal margin of superlingua; D) maxillary palp somewhat longer than length of galea-lacinia; E) fore femur broad, length 2.5× maximum width, dorsal margin with a row of ca. 37 curved, spine-like setae and distally with some stout, pointed setae near margin; F) gills margin serrate with alternating smaller and bigger denticles intercalating long, fine, simple setae; G) spines at posterior margin of tergum IV rounded, wider than long; H) paraproct distally not expanded.
Larva (Figs
Colouration. Head, thorax and abdomen dorsally brown, head and thorax with bright median, dorsal suture, forewing pads with slightly darker striation. Head, thorax and abdomen ventrally light brown, legs light brown and with brown marks as in Fig.
Antenna with scape and pedicel sub-cylindrical, without distolateral process at scape; flagellum with broad, apically blunt spines and fine, simple setae on apex of each segment.
Labrum (Fig.
Right mandible (Fig.
Left mandible (Fig.
Both mandibles with lateral margins almost straight. Basal half with fine, simple setae scattered over dorsal surface.
Hypopharynx (Fig.
Maxilla (Fig.
Labium (Fig.
Hind wing pads absent.
Foreleg (Fig.
Tergum (Fig.
Gills (Fig.
Paraproct (Fig.
Latin word for sword, refers to the blade-like outermost denticle of the mandibles.
New Guinea.
The specimens were collected at altitudes of 1800–3210 m a.s.l.
Holotype. Nymph (on slide, GBIFCH 00465172), Papua New Guinea, Simbu Prov., 05°49.96'S, 145°06.129'E, (GPS), Mt. Wilhelm, Pindaunde Creek, 2350 m a.s.l. (7915 ft), S5 (oria.6), 18 Aug 1999, leg. L. Čížek . Deposited in
5 nymphs (1 on slide, GBIFCH 00465179, 4 in alcohol, GBIFCH 00515275, GBIFCH 00508135, deposited in
Larva. Following combination of characters: A) labrum dorsal arc of setae with one plus 6–7 long, simple setae; B) both mandibles with outermost denticle blade-like; C) hypopharynx with slightly concave distal margin of superlingua; D) maxillary palp somewhat longer than length of galea-lacinia; E) fore femur rather broad, length 2.6× maximum width, dorsal margin with a row of ca. 34 curved, spine-like setae and with some stout, pointed setae near margin; F) gills long, gill IV as long as length of segments V to VII combined, gill VII somewhat longer than length of segments VIII to X combined.
Larva (Figs
Colouration. Head, thorax and abdomen dorsally grey-brown, abdominal segments VI, IX and X brighter, head and thorax with bright median, dorsal suture, forewing pads with bright striation. Head, thorax and abdomen ventrally light grey-brown, legs colourless with brown medial spot and dorsal margin on femur, caudal filaments light brown.
Antenna with scape and pedicel sub-cylindrical, without distolateral process at scape; flagellum with lanceolate spines on apex of each segment.
Labrum (Fig.
Right mandible (Fig.
Left mandible (Fig.
Both mandibles with lateral margins almost straight. Basal half with fine, simple setae scattered over dorsal surface.
Hypopharynx (Fig.
Maxilla (Fig.
Labium (Fig.
Hind wing pads absent.
Foreleg (Fig.
Tergum (Fig.
Gills (Fig.
Paraproct (Fig.
Dedicated to Janice (“Jan”) Peters (Florida A & M University), who pioneered the collection of mayflies in New Guinea.
New Guinea.
The specimens were collected at an altitude of 3200 m a.s.l.
Holotype. Nymph (on slide, GBIFCH 00465181), Indonesia, Papua, Lake Habemma, stream, 3200 m, 19 Oct 2011, 04°07.77'S, 138°40.77'E, Balke (PAP07). Temporary deposited in
The group is characterised by a compact, rounded distomedial protuberance of labial palp segment II and a dorsal, submarginal arc of setae of the labrum composed of simple setae. In this arc of setae the first and second setae after the central, submedian seta are standing closely together.
Larva. Following combination of characters: A) labrum dorsal arc of setae composed of one plus 4–5 long, simple setae; B) maxillary palp longer than galwa-lacinia, with well-developed excavation at inner distolateral margin of segment II; C) fore femur rather broad, length 2.8× maximum width, foreleg setation as Fig.
Paratypes. 1 nymph (on slides, PERC 0 010 571), Papua New Guinea, Bulolo R, E of Wau, 2950 ft, 18 Oct 1964, W.L. and J.G. Peters; 1 nymph (on slide, PERC 0 010 573), Papua New Guinea, Morobe Prov., Clearwater Cr, nr Luau, 15 Sept 1983, J.T. and D.A. Polhemus.
Larva. Following combination of characters: A) labrum dorsal submarginal arc of setae composed of one plus eight or nine long, simple setae; B) labial palp segment II with a compact, rounded distomedial protuberance; C) fore femur broad, length ca. 2× maximum width, dorsal margin with a row of ca. 26 curved, spine-like setae and many stout, pointed setae near margin; D) gills margin serrate with small spines intercalating long, fine, simple setae, and with robust, lanceolate setae on margin; E) paraproct surface with scales or scale bases and fine, slightly lanceolate setae as well as fine, simple setae.
Larva (Figs
Colouration. Head, thorax and abdomen dorsally dark brown, abdominal segment X light brown, head and thorax with bright median, dorsal suture. Head, thorax and abdomen ventrally brown, femur dorsal and ventral margins brown, legs otherwise colourless, caudal filaments light brown.
Antenna with scape and pedicel sub-cylindrical, without distolateral process at scape; flagellum with lanceolate spines and fine, simple setae on apex of each segment.
Labrum (Fig.
Right mandible (Fig.
Left mandible (Fig.
Both mandibles with lateral margins slightly convex. Basal half with fine, simple setae scattered over dorsal surface.
Hypopharynx (Fig.
Maxilla (Fig.
Labium (Fig.
Hind wing pads absent.
Foreleg (Fig.
Tergum (Fig.
Gills (Fig.
Paraproct (Fig.
Latin words for gills and seta, refers to the robust, lanceolate setae on the margin of the gills.
New Guinea.
The specimens were collected in altitudes of 1000 m a.s.l. and 1700 m–1800 m a.s.l.
Holotype. Nymph (on slide, GBIFCH 00465183), Papua New Guinea, Eastern Highlands, Marawaka, Ande, 1700–1800 m, 09 Nov 2006, nr 07°01.70'S, 145°49.81'E, Balke & Kinibel (PNG 87). Deposited in
Larva. Following combination of characters: A) labrum dorsal submarginal arc of setae composed of one plus six or seven long, simple setae; B) maxillary palp about as long as galea-lacinia, excavation on inner distal margin of segment II poorly developed; C) labial palp segment II with a compact, rounded distomedial protuberance; D) fore femur rather broad, length ca. 3× maximum width, dorsal margin with a row of ca. 34 curved, spine-like setae and many stout, pointed setae near margin; E) fore claw with one row of 11–12 denticles; F) tracheae of gills restricted to main trunk, without pigmentation; G) paraproct surface with slightly lanceolate setae and fine, simple setae.
Larva (Figs
Colouration. Head, thorax and abdomen dorsally brown, head and thorax with bright median, dorsal suture, forewing pads with bright striation. Head, thorax and abdomen ventrally brown, legs light brown, dorsal magin of femur and tibia brown, caudal filaments brown.
Antenna with scape and pedicel sub-cylindrical, without distolateral process at scape; flagellum with lanceolate spines and fine, simple setae on apex of each segment.
Labrum (Fig.
Right mandible (Fig.
Left mandible (Fig.
Both mandibles with lateral margins almost straight. Basal half with fine, simple setae scattered over dorsal surface.
Hypopharynx (Fig.
Maxilla (Fig.
Labium (Fig.
Hind wing pads absent.
Foreleg (Fig.
Tergum (Fig.
Gills (Fig.
Paraproct (Fig.
Terminal filament 0.6× length of cerci.
Latin for “very high”, refers to the high altitude (2900 m a.s.l.) of the type locality.
New Guinea.
The specimens were collected in altitudes of 2200 m a.s.l. and 2900 m a.s.l. (in forest).
Holotype. 1 Nymph (on slide, GBIFCH 00465187), Papua New Guinea, Simbu Prov., 05°49'S, 145°04.5'E, Mt. Wilhelm, Pindaunde Creek, 2900 m a.s.l. (in forest), S3 (oria.4), 18 Aug 1999, leg. L. Čížek. Deposited in
Larva. Following combination of characters: A) labrum dorsal submarginal arc of setae composed of one plus 5–7 long, simple setae; B) maxillary palp much longer than length of galea-lacinia, apically rounded, with very poorly developed excavation at inner distolateral margin; C) labial palp segment II with a compact, rounded distomedial protuberance; D) fore femur rather broad, length ca. 3× maximum width, dorsal margin with a row of ca. 19 curved, spine-like setae and some stout, pointed setae near margin; E) fore claw with 7 – 8 denticles; F) paraproct distally expanded, surface with few slightly spatulate setae.
Larva (Figs
Colouration. Head, thorax and abdomen dorsally brown, head and thorax with bright median, dorsal suture, forewing pads with bright striation. Head, thorax and abdomen ventrally light brown. Legs colourless, femur with large, distomedial, brown spot and brown dorsal margin, caudal filaments light brown.
Antenna with scape and pedicel sub-cylindrical, without distolateral process at scape; flagellum without spines on apex of segments.
Labrum (Fig.
Right mandible (Fig.
Left mandible (Fig.
Both mandibles with lateral margins almost straight. Basal half with fine, simple setae scattered over dorsal surface.
Hypopharynx (Fig.
Maxilla (Fig.
Labium (Fig.
Hind wing pads absent.
Foreleg (Fig.
Tergum (Fig.
Gills (Fig.
Paraproct (Fig.
Refers to the low altitude (95 m a.s.l.) of the type locality.
New Guinea.
The specimens were collected at an altitude of 95 m a.s.l.
Holotype. Nymph (on slide, GBIFCH 00465191), Indonesia, Papua, Sorong, inland, 95 m, 19 Feb 2006, 00°49.35'S, 131°24.20'E, Balke & Tindige (BH 20). Temporary deposited in
Larva. Following combination of characters: A) labrum dorsal submarginal arc of setae composed of one plus six long, simple setae; B) maxillary palp 1.2× length of galea-lacinia; C) labial palp segment II with a compact, rounded distomedial protuberance; D) fore femur rather broad, length ca. 3× maximum width, dorsal margin with a row of ca. 16 curved, spine-like setae and a row of spine-like setae near margin; E) paraproct with three robust, lanceolate setae near distolateral margin.
Larva (Figs
Colouration. Head, thorax and abdomen dorsally brown, head and thorax with bright median, dorsal suture, forewing pads with bright striation. Head, thorax and abdomen ventrally light brown, femur dorsal margin brown, legs otherwise colourless, caudal filaments light brown.
Antenna with scape and pedicel sub-cylindrical, without distolateral process at scape; flagellum with broad spines on apex of each segment.
Labrum (Fig.
Right mandible (Fig.
Left mandible (Fig.
Both mandibles with lateral margins almost straight. Basal half with fine, simple setae scattered over dorsal surface.
Hypopharynx (Fig.
Maxilla (Fig.
Labium (Fig.
Hind wing pads absent.
Foreleg (Fig.
Tergum (Fig.
Gills (Fig.
Paraproct (Fig.
Dedicated to Marion Podolak (Museum for Zoology Lausanne,
New Guinea.
Holotype. Nymph (on slide, GBIFCH 00465227), Papua New Guinea, Eastern Highlands, Aiyura, 1710 m, 15 Jan 2003, 06°21.41'S, 145°54.34'E, grassland stream, K. Sagata leg. Deposited in
Larva. Following combination of characters: A) labrum dorsal submarginal arc of setae composed of one plus nine long, simple setae ; B) maxillary palp longer than length of galea-lacinia, apically rounded, without excavation at inner distolateral margin; C) labial palp segment II with a compact, rounded distomedial protuberance; D) fore femur rather broad, length ca. 3× maximum width, dorsal margin with a row of ca. 26 curved, spine-like setae and many stout, pointed setae near margin; E) paraproct distally expanded, surface with slightly spatulate setae.
Larva (Figs
Colouration. Head, thorax and abdomen dorsally brown, head and thorax with bright median, dorsal suture. Head, thorax and abdomen ventrally light brown. Femur dorsal margin light brown, legs otherwise colourless, caudal filaments basally light brown, otherwise colourless.
Antenna with scape and pedicel sub-cylindrical, without distolateral process at scape; flagellum with broad, lanceolate spines and fine, simple setae on apex of each segment.
Labrum (Fig.
Right mandible (Fig.
Left mandible (Fig.
Both mandibles with lateral margins almost straight. Basal half with fine, simple setae scattered over dorsal surface.
Hypopharynx (Fig.
Maxilla (Fig.
Labium (Fig.
Hind wing pads absent.
Foreleg (Fig.
Tergum (Fig.
Gills (Fig.
Paraproct (Fig.
Dedicated to Sereina Rutschmann (University of Vigo, Spain) for her long-lasting support of our group in genetics.
New Guinea.
The specimens were collected in altitudes of 1800 m–2000 m a.s.l.
Holotype. Nymph (on slide, GBIFCH 00465195), Papua New Guinea, Western Highlands, Simbai, 1800–2000 m, 01 Mar 2007, 05°14.28'S, 144°28.74'E, Kinibel (PNG 138). Deposited in
Larva. Following combination of characters: A) labrum dorsal arc of setae composed of one plus five long, simple setae; B) maxillary palp 1.3× as long as length of galea-lacinia; segment II apically rounded, with an excavation at inner lateral margin; C) labial palp segment II with a compact, rounded distomedial protuberance, segment III conical; D) fore femur rather broad, length 2.7× maximum width, dorsal margin with a row of ca. 21 curved, spine-like setae and many stout, pointed setae near margin; E) fore claw with a row of 8 denticles; F) spines at posterior margin of tergum IV mostly rounded, about as long as wide; G) paraproct surface with U-shaped scale bases and scattered fine, slightly spatulate setae and micropores.
Larva (Figs
Colouration. Head, thorax and abdomen dorsally brown, head and thorax with bright median, dorsal suture. Head, thorax and abdomen ventrally colourless. Legs colourless with some brown spots, femur with brown dorsal margin, caudal filaments brown.
Antenna with scape and pedicel sub-cylindrical, without distolateral process at scape; flagellum with broad, lanceolate spines and fine, simple setae on apex of each segment.
Labrum (Fig.
Right mandible (Fig.
Left mandible (Fig.
Both mandibles with lateral margins almost straight. Basal half with fine, simple setae scattered over dorsal surface.
Hypopharynx (Fig.
Maxilla (Fig.
Labium (Fig.
Hind wing pads absent.
Foreleg (Fig.
Tergum (Fig.
Gills (Fig.
Paraproct (Fig.
Dedicated to Tanja Schwander (University of Lausanne, UNIL) for her constant support during a master project of one of the authors (TK) in her lab.
New Guinea.
The specimens were collected at an altitude of 1400 m a.s.l.
Holotype. Nymph (on slide, GBIFCH 00465197), Papua New Guinea, Gulf, Marawaka, 1400 m, 11 Nov 2006, 07°05.66'S, 145°44.47'E, Balke & Kinibel (PNG 90). Deposited in
The group is characterized by a short, thumb-like protuberance of labial palp segment II and a dorsal, submarginal arc of setae composed of simple setae. Additionally, there is no distolateral excavation at maxillary palp segment II.
Larva. Following combination of characters: A) labrum dorsal arc of setae composed of one plus eight long, simple setae, 5 setae standing closely together; B) right mandible with 3+3 denticles; C) left mandible with 3+3 denticles; D) maxillary palp longer than galea-lacinia, without excavation at inner distolateral margin; E) labial palp segment II with short thumb-like protuberance, segment III slightly pentagonal; F) fore femur rather broad, length ca. 3× as long as maximum width; G) fore leg setation as Fig.
Paratype. 1 nymph (on slide, PERC 0 012 576), Papua New Guinea, Western Highlands Prov., Kaugel R, nr Alkena, 07 Sept 1983, J.T. and D.A. Polhemus.
Larva. Following combination of characters: A) labrum dorsal submarginal arc of setae composed of one plus 5–6 long, simple setae; B) maxillary palp somewhat longer as length of galea-lacinia, apically constricted, without excavation at inner distolateral margin; C) labium paraglossa apically with four rows of long, robust, apically pectinate setae; D) labial palp segment II with a short, thumb-like distomedial protuberance; E) fore femur rather broad, length ca. 3× maximum width, dorsal margin with a row of ca. 30 curved, spine-like setae and many stout, pointed setae near margin; F) fore claw with one row of 12–13 denticles.
Larva (Figs
Colouration. Head, thorax and abdomen dorsally brown, head and thorax with bright median, dorsal suture, thorax and abdomen with bright pattern as in Fig.
Antenna with scape and pedicel sub-cylindrical, without distolateral process at scape; flagellum with lanceolate spines and fine, simple setae on apex of each segment.
Labrum (Fig.
Right mandible (Fig.
Left mandible (Fig.
Both mandibles with lateral margins almost straight. Basal half with fine, simple setae scattered over dorsal surface.
Hypopharynx (Fig.
Maxilla (Fig.
Labium (Fig.
Hind wing pads absent.
Foreleg (Fig.
Tergum (Fig.
Gills (Fig.
Paraproct (Fig.
Latin word for high, refers to the altitude of the type locality (2700 m a.s.l.).
New Guinea.
The specimens were collected in altitudes of 2700 m a.s.l. and 2900 m a.s.l.
Holotype. Nymph (on slide, GBIFCH 00465199), Papua New Guinea, Enga, Kumul Lodge at foot of Mt. Hagen, 2700 m, 05 Dec 2006, 05°47.55'S, 143°58.76'E, Balke & Kinibel (PNG 124). Deposited in
Larva. Following combination of characters: A) labrum dorsal arc of submarginal setae composed of one plus 6–7 long simple setae; B) maxillary palp about as long as galea-lacinia, apically slightly pointed and without excavation at inner lateral margin; C) labial palp segment II with an elongated, thumb-like distomedial protuberance; D) fore femur rather broad, length ca. 3× maximum width, dorsal margin with a row of ca. 26 curved, spine-like setae; E) fore claw with 13–15 denticles.
Larva (Figs
Colouration. Head, thorax and abdomen dorsally brown, head and thorax with bright median, dorsal suture, forewing pads with bright striation, thorax and abdomen with bright pattern as in Fig.
Antenna with scape and pedicel sub-cylindrical, without distolateral process at scape; flagellum with lanceolate spines and fine, simple setae on apex of each segment.
Labrum (Fig.
Right mandible (Fig.
Left mandible (Fig.
Both mandibles with lateral margins almost straight. Basal half with fine, simple setae scattered over dorsal surface.
Hypopharynx (Fig.
Maxilla (Fig.
Labium (Fig.
Hind wing pads absent.
Foreleg (Fig.
Tergum (Fig.
Gills (Fig.
Paraproct (Fig.
Dedicated to the late friend of one of the authors (JLG), the biologist Cédric Gindro.
New Guinea.
The specimens were collected in altitudes of 1620 m a.s.l. and 2000 m a.s.l.
Holotype. Nymph (on slide, GBIFCH 00465202), Indonesia, Papua, Wamena, 20 mins towd Jiwika, limestone creek, 1620 m, 18 Oct 2011, 03°56.95'S, 138°54.38'E, Balke (PAP07). Temporary deposited in
5 nymphs (1 on slide, GBIFCH 00465206, 4 in alcohol, GBIFCH 00515291, deposited in
Larva. Following combination of characters: A) labrum dorsal submarginal arc of setae composed of one plus 5–6 long, simple setae; B) maxillary palp longer as length of galea-lacinia, apically rounded, without excavation at inner distolateral margin; C) labial glossae much shorter than paraglossae; D) labial palp segment II with an elongated, thumb-like distomedial protuberance, segment III conical, apically slightly truncate; E) fore femur slender, length 3.6× maximum width, dorsal margin with a row of ca. 12 curved, spine-like setae; F) fore claw with one row of eleven denticles.
Larva (Figs
Colouration. Head, thorax and abdomen dorsally brown, head and thorax with bright median, dorsal suture. Head, thorax and abdomen ventrally light brown, femur dorsal margin light brown, legs otherwise colourless, caudal filaments colourless.
Antenna with scape and pedicel sub-cylindrical, without distolateral process at scape; flagellum with lanceolate spines and fine, simple setae on apex of each segment.
Labrum (Fig.
Right mandible (Fig.
Left mandible (Fig.
Both mandibles with lateral margins almost straight. Basal half with fine, simple setae scattered over dorsal surface.
Hypopharynx (Fig.
Maxilla (Fig.
Labium (Fig.
Hind wing pads absent.
Foreleg (Fig.
Tergum (Fig.
Gills (Fig.
Paraproct (Fig.
Refers to the morphological similarity with L. vitilis.
New Guinea.
The specimens were collected at an altitude of 30 m a.s.l.
Holotype. Nymph (on slide, GBIFCH 00465207), Papua New Guinea, Madang, Trans Gogol, 30 m, 02.2008, 05°18.09'S, 145°36.45'E, BRC leg. (PNG 179). Deposited in
Larva. Following combination of characters: A) labrum dorsal submarginal arc of setae composed of one plus 7–10 long, simple setae; B) maxillary palp somewhat longer than length of galea-lacinia, segment II apically slightly pointed and without excavation at inner lateral margin; C) labial palp segment III oblong, apically slightly pointed; D) labium paraglossa apically with five rows of long, robust, apically pectinate setae; E) fore femur rather broad, length ca. 3× maximum width, dorsal margin with a row of ca. 29 curved, spine-like setae and many stout, pointed setae near margin; F) fore claw with one row of 13–15 denticles; G) tracheae of gills restricted to main trunk.
Larva (Figs
Colouration. Head, thorax and abdomen dorsally brown, head and thorax with bright median, dorsal suture. Head, thorax and abdomen ventrally light brown, legs colourless with light brown pattern as in Fig.
Antenna with scape and pedicel sub-cylindrical, without distolateral process at scape; flagellum with lanceolate spines and fine, simple setae on apex of each segment.
Labrum (Fig.
Right mandible (Fig.
Left mandible (Fig.
Both mandibles with lateral margins almost straight. Basal half with fine, simple setae scattered over dorsal surface.
Hypopharynx (Fig.
Maxilla (Fig.
Labium (Fig.
Hind wing pads absent.
Foreleg (Fig.
Tergum (Fig.
Gills (Fig.
Paraproct (Fig.
Refers to the type locality at Mt. Wilhelm.
New Guinea.
The specimens were collected in altitudes of 2900 m a.s.l. and 3210 m a.s.l.
Holotype. Nymph (on slide, GBIFCH 00465209), Papua New Guinea, Simbu Prov., 05°48.050'S, 145°04.15'E, (GPS)’, Mt. Wilhelm, Pindaunde Creek, 3210 m a.s.l., (10895 ft GPS), S32 (oria.3), 17 Aug 1999, L. Čížek leg. Deposited in
The group is characterised by a hook-like protuberance of labial palp segment II and the labrum with a dorsal, submarginal arc of setae composed of simple setae.
Larva. Following combination of characters: A) labrum dorsal submarginal arc of setae composed of one plus 8–9 long, simple setae; B) maxillary palp longer than galea-lacinia, with well-developed excavation at inner distolateral margin; C) labial palp segment II with hook-like distomedial protuberance, as Fig.
Paratype. 1 nymph (on slide, PERC 0 012 577), Papua New Guinea, Western Highlands Prov., 17 km N of Mt. Hagen, 06 Sept 1983, J.T. and D.A. Polhemus leg.
Larva. Following combination of characters: A) labrum dorsal submarginal arc of setae composed of one plus nine long, simple setae; B) maxillary palp with segment II longer than length of segment I, segment II with excavation at inner distolateral margin; C) labial palp segment II with a hook-like distomedial protuberance; D) fore femur rather broad, length ca. 3× maximum width, dorsal margin with a row of ca. 28 curved, spine-like setae; E) fore claw with 10 denticles; F) gills margin serrated with alternating bigger and smaller spines.
Larva (Figs
Colouration. Head, thorax and abdomen dorsally brown, head and thorax with bright median, dorsal suture, forewing pads with bright striation. Head, thorax and abdomen ventrally light brown, femur colourless with a medial brown spot and a brown dorsal margin, tibia and tarsus brown, caudal filaments brown.
Antenna with scape and pedicel sub-cylindrical, without distolateral process at scape; flagellum with lanceolate spines and fine, simple setae on apex of each segment.
Labrum (Fig.
Right mandible (Fig.
Left mandible (Fig.
Both mandibles with lateral margins slightly convex. Basal half with fine, simple setae scattered over dorsal surface.
Hypopharynx (Fig.
Maxilla (Fig.
Labium (Fig.
Hind wing pads absent.
Foreleg (Fig.
Tergum (Fig.
Gills (Fig.
Paraproct (Fig.
Refers to the similarity and close relationship to L. vultuosus.
New Guinea.
The specimens were collected at an altitude of 1500 m a.s.l.
Holotype. Nymph (on slide, GBIFCH 00465213), Papua New Guinea, Enga, Wapanamanda, 1500 m, 06 Dec 2006, 05°38.11'S, 143°55.34'E, Balke & Kinibel (PNG 128). Deposited in
Larva. Following combination of characters: A) labrum with dorsal submarginal arc of setae composed of one plus six long, simple setae; B) labial palp segment II with an elongated, thumb-like distomedial protuberance; C) maxillary palp slightly longer than length of galea-lacinia, segment II apically slightly pointed and without excavation at inner lateral margin; D) right mandible outer and inner sets of denticles with 4 + 4 denticles respectively plus one small intermediate denticle; E) left mandible outer and inner sets of denticles with 4 + 4 denticles respectively; F) fore femur slender, length ca. 4× maximum width, dorsal margin with a row of ca. 21 curved, spine-like setae and distally a row of robust, spine-like setae close to margin.
Larva (Figs
Colouration. Head, thorax and abdomen dorsally brown, with bright pattern as in Fig.
Antenna with scape and pedicel sub-cylindrical, without distolateral process at scape; flagellum with broad spines on apex of each segment.
Labrum (Fig.
Right mandible (Fig.
Left mandible (Fig.
Both mandibles with lateral margins almost straight. Basal half with fine, simple setae scattered over dorsal surface.
Hypopharynx (Fig.
Maxilla (Fig.
Labium (Fig.
Hind wing pads absent.
Foreleg (Fig.
Tergum (Fig.
Gills (Fig.
Paraproct (Fig.
Refers to the type locality in the Central Province of Papua New Guinea.
New Guinea.
The specimens were collected at an altitude of 590 m a.s.l.
Holotype. Nymph (on slide, GBIFCH 00465215), Papua New Guinea, Central, Kokoda Trek, 590 m, 01.2008, 09°14.34'S, 147°36.92'E, Posman (PNG170). Deposited in
Larva. Following combination of characters: A) labrum dorsal submarginal arc of setae composed of one plus 5 long, dendritic, apically pointed setae and 3 middle to long, simple setae; B) both mandibles with outermost denticles blade-like; C) maxillary palp somewhat shorter than length of galea-lacinia; D) distomedial protuberance of labial palp segment II short, thumb-like; E) fore femur rather broad, length ca. 3× maximum width, dorsal margin with a row of ca. 20 curved, spine-like setae and basally a row of robust, spine-like setae near margin; F) tarsal claw with one row of seven denticles and without striation.
Larva (Figs
Colouration. Unknown.
Antenna with scape and pedicel sub-cylindrical, without distolateral process at scape; flagellum with lanceolate spines on apex of each segment.
Labrum (Fig.
Right mandible (Fig.
Left mandible (Fig.
Both mandibles with lateral margins almost straight. Basal half with fine, simple setae scattered over dorsal surface.
Hypopharynx (Fig.
Maxilla (Fig.
Labium (Fig.
Hind wing pads absent.
Foreleg (Fig.
Tergum (Fig.
Gills (Fig.
Paraproct (Fig.
Refers to the dendritic submarginal setae on dorsal surface of labrum.
New Guinea.
The specimens were collected in forest at an altitude of 2900 m a.s.l.
Holotype. Nymph (on slide, GBIFCH 00465217), Papua New Guinea, Simbu Prov., 05°49.00'S, 145°04.50'E, Mt. Wilhelm, Pindaunde Creek, 2900 m a.s.l. (in forest), S3 (oria.4), 18 Aug 1999, leg. L. Čížek . Deposited in
Larva. Following combination of characters: A) labrum dorsal submarginal arc of setae composed of one plus 11–13 long, simple setae; B) labrum dorsally with many medium, fine, simple setae, mainly arranged in one band; C) labial palp segment III with emargination at distal margin; D) maxillary palp shorter than galea-lacinia; E) fore femur broad, length ca. 2× maximum width, dorsal margin with an irregular row of more than 40 curved, spine-like setae and some curved, spine like setae near margin.
Larva (Figs
Colouration. Head, thorax and abdomen dorsally brown, abdominal terga I and X lighter. Head and thorax with bright median, dorsal suture, forewing pads with bright striation. Thorax and abdomen ventrally light brown, femur with brown dorsal margin, legs otherwise light brown, caudal filaments light brown.
Antenna with scape and pedicel sub-cylindrical, without distolateral process at scape; flagellum with broad, apically blunt spines and fine, simple setae on apex of each segment.
Labrum (Fig.
Right mandible (Fig.
Left mandible (Fig.
Both mandibles with lateral margins slightly convex. Basal half with fine, simple setae scattered over dorsal surface.
Hypopharynx (Fig.
Maxilla (Fig.
Labium (Fig.
Hind wing pads absent.
Foreleg (Fig.
Tergum (Fig.
Gills (Fig.
Paraproct (Fig.
Dedicated to Elisa Gattolliat, the daughter of one of the authors.
New Guinea.
The specimens were collected in altitudes of 1200 m and 1400 m a.s.l.
Holotype. Nymph (on slide, GBIFCH 00465219), Papua New Guinea, Western Highlands, Kundum, 1400 m, 03 Mar 2007, 05°16.10'S, 144°27.87'E, Kinibel (PNG142). Deposited in
Larva. Following combination of characters: A) labrum dorsal submarginal arc of setae composed of 21 long, lanceolate, apically pectinate setae; B) maxillary palp much longer than length of galea-lacinia, apically rounded, with excavation at inner distolateral margin; C) labial palp segment II with a large, lobed distomedial protuberance, segment III slightly pentagonal, apically slightly pointed; D) fore femur very slender, length ca. 5× maximum width, dorsal margin with a row of ca. 12 curved, spine-like setae; E) fore claw with one row of eleven denticles.
Larva (Figs
Colouration. Unknown.
Labrum (Fig.
Right mandible (Fig.
Left mandible (Fig.
Both mandibles with lateral margins almost straight. Basal half with fine, simple setae scattered over dorsal surface.
Hypopharynx (Fig.
Maxilla (Fig.
Labium (Fig.
Hind wing pads unknown.
Foreleg (Fig.
Tergum. Unknown.
Gills. Unknown.
Paraproct. Unknown.
Latin word for unexpected, refers to the unexpected finding of this species amongst other material.
New Guinea.
Holotype. Nymph (on slide, GBIFCH 00465230), Papua New Guinea, Gulf Prov., Supa-Hala, 1032 m, 10 Nov 2002, forest stream, K. Sagata leg. Deposited in
Larva. Following combination of characters: A) labrum dorsal submarginal arc of setae composed of one plus 4–5 long, simple setae; B) labial palp segment II with elongated thumb-like distomadial protuberance, segment III about semicircular; C) maxillary palp shorter than length of galea-lacinia, without excavation at inner distolateral margin of segment II; D) fore femur rather broad, length ca. 3× maximum width; E) fore claw with a row of 10–12 denticles; F) spines at posterior margin of tergum IV triangular, pointed, longer than wide; G) paraproct distally not expanded.
Paratypes. 2 nymphs (on slides, PERC 0.012.560, PERC 0.012.561), Papua New Guinea, Bulolo River, East of Wau, 2950 ft, 15 Oct 1964, W.L. and J.G. Peters leg.
Larva. Following combinations of characters: A) labrum dorsal submarginal arc of setae composed of one plus 10–12 long, simple setae; B) maxillary palp 1.6× as long as length of galea-lacinia; C) labial palp segment III conical, slightly pointed; D) labial palp segment II with a large, lobed distomedial protuberance; E) fore femur slender, length ca. 4× maximum width, dorsal margin with a row of ca. 27 curved, spine-like setae and with stout, pointed setae near margin.
Larva (Figs
Colouration. Head, thorax and abdomen dorsally brown, head and thorax with bright median, dorsal suture, forewing pads with bright striation. Head, thorax and abdomen ventrally light brown, legs colourless, caudal filaments light brown.
Antenna with scape and pedicel sub-cylindrical, without distolateral process at scape; flagellum with lanceolate spines on apex of each segment.
Labrum (Fig.
Right mandible (Fig.
Left mandible (Fig.
Both mandibles with lateral margins almost straight. Basal half with fine, simple setae scattered over dorsal surface.
Hypopharynx (Fig.
Maxilla (Fig.
Labium (Fig.
Hind wing pads absent.
Foreleg (Fig.
Tergum (Fig.
Gills (Fig.
Paraproct (Fig.
Refers to Pindaunde Creek in Papua New Guinea, where the species has been collected.
New Guinea.
The specimens were collected at altitudes of 1700 m, 2600 m and 2900 m a.s.l., partly in forest.
Holotype. Nymph (on slide, GBIFCH 00465222), Papua New Guinea, Simbu Prov., 05°49.00'S, 145°04.50'E, (GPS), Mt. Wilhelm, Pindaunde Creek, 2900 m a.s.l. (in forest), S3 (oria.4),18 Aug 1999, leg. L. Čížek. Deposited in
10 nymphs (1 on slide, GBIFCH 00465225, 9 in alcohol, GBIFCH 00515278, deposited in
Larva. Following combination of characters: A) labrum dorsal submarginal arc of setae composed of 23 long, lanceolate, apically pointed setae; B) maxillary palp with segment II 2.4× longer than length of segment I, slender, apically pointed, without excavation at inner lateral margin; C) labial palp segment III sub-rectangular, apically slightly pointed; D) labial palp segment II with an elongated, thumb-like distomedial protuberance; E) fore femur slender, length ca. 4× maximum width, dorsal margin with a row of ca. 10 curved, spine-like setae and some stout, pointed setae near margin.
Larva (Figs
Colouration. Thorax and abdomen dorsally brown, with bright pattern as in Fig.
Antenna with scape and pedicel sub-cylindrical, without distolateral process at scape.
Labrum (Fig.
Right mandible (Fig.
Left mandible (Fig.
Both mandibles with lateral margins almost straight. Basal half with fine, simple setae scattered over dorsal surface.
Hypopharynx (Fig.
Maxilla (Fig.
Labium (Fig.
Hind wing pads absent.
Foreleg (Fig.
Tergum (Fig.
Gills (Fig.
Paraproct (Fig.
Refers to the remarkable, fence-like, dorsal, submarginal arc of setae of the labrum.
New Guinea.
The specimen was collected at an altitude of 400 m a.s.l.
Holotype. Nymph (on slide, GBIFCH 00465226), Papua New Guinea, Madang, Keki, Adalbert Mts, 400 m, 29 Nov 2006, 04°43.06'S, 145°24.44'E, Binatang Boys (PNG 119). Deposited in
Larva. Following combination of characters: A) labrum dorsal submarginal arc of setae composed of one plus 7–9 long, simple setae; B) maxillary palp shorter than length of galea-lacinia, segment II without excavation at inner distolateral margin; C) labial palp segment II with elongated thumb-like distomedial protuberance, segment III about semicircular; D) femur rather broad, length ca. 2–3× maximum width, dorsal margin with >40 curved, spine-like setae; E) fore claw with a row of 8–10 denticles; F) Gills with strongly developed, dense, pigmented tracheation; G) spines at posterior margin of tergum IV irregular, mostly triangular, apically rounded, wider than long; H) paraproct with marginal spines poorly defined.
Paratypes. 2 nymphs (on slides, PERC 0.012.578, PERC 0.012.579), Papua New Guinea, Morobe Prov., Poverty Cr., Mt. Missim, 1600 m, 18.09. Sept 1983, J.T. and D.A. Polhemus leg.
1 | Labrum submarginal dorsal arc of simple setae (Fig. |
2 |
– | Labrum submarginal dorsal arc of lanceolate setae (Fig. |
L. vallus sp. n. |
– | Labrum submarginal dorsal arc of lanceolate, apically pectinate setae (Fig. |
L. inopinatus sp. n. |
– | Labrum submarginal dorsal arc of spatulate, apically pectinate setae (Fig. |
17 |
– | Labrum submarginal dorsal arc of feathered setae (Fig. |
19 |
– | Labrum submarginal dorsal arc of dendritic setae (Fig. |
L. dendrisetis sp. n. |
2(1) | Labial palp segment II enlargement elongated, thumb-like (Fig. |
3 |
– | Labial palp segment II enlargement short, thumb-like (Fig. |
7 |
– | Labial palp segment II enlargement compact, rounded (Fig. |
12 |
– | Labial palp segment II enlargement large, lobed (Fig. |
L. pindaundensis sp. n. |
– | Labial palp segment II enlargement hook-like (Fig. |
16 |
3(2) | Labial palp segment III subrectangular (Fig. |
L. claudiae sp. n. |
– | Labial palp segment III about semicircular (Fig. |
4 |
– | Labial palp segment III conical (Fig. |
6 |
– | Labial palp segment III oblong (Fig. |
L. gindroi sp.n. |
4(3) | Mandibles with outermost denticle normally developed (Fig. |
5 |
– | Mandibles with outermost denticle blade-like (Fig. |
L. xeniolus |
5(4) | Maxillary palp shorter than length of galea-lacinia; gills margin serrate with alternating smaller and bigger denticles and with fine, simple setae; number of femur dorsal setae on margin > 40; paraproct distally not expanded ( |
L. involutus |
– | Maxillary palp longer than length of galea-lacinia; gills margin serrate with small denticles and with fine simple setae (Fig. |
L. stagnum sp. n. |
6(3) | Labial palp segment III distal margin rounded (Fig. |
L. centralensis sp. n. |
– | Labial palp segment III distal margin truncate (Fig. |
L. paravitilis sp. n. |
7(2) | Labium paraglossae distally with three rows of setae | 8 |
– | Labium paraglossae distally with four rows of setae | L. altus sp. n. |
– | Labium paraglossae distally with five rows of setae | L. wilhelmensis sp. n. |
8(7) | Mandibles with outermost denticle normally developed (Fig. |
9 |
– | Mandibles with outermost denticle blade-like (Fig. |
10 |
9(8) | Labial palp segment III conical, distal margin with emargination (Fig. |
L. elisae sp. n. |
– | Labial palp segment III slightly pentagonal, distal margin rounded (Fig. |
L. vitilis |
10(8) | Labial palp segment III distal margin slightly pointed (Fig. |
L. gladius sp. n. |
– | Labial palp segment III distal margin rounded (Fig. |
11 |
11(10) | Maxillary palp segment II longer than segment I; spines at posterior margin of abdominal terga wider than long; gills margin serrate with small denticles and with fine simple setae (Fig. |
L. janae sp. n. |
– | Maxillary palp segment II about as long or shorter than segment I; spines at posterior margin of abdominal terga longer than wide; gills margin serrate with alternating smaller and bigger denticles and with fine, simple setae ( |
L. petersorum |
12(2) | Maxillary palp about as long as length of galea-lacinia | L. magnovaldus sp. n. |
– | Maxillary palp longer (1.2–1.4×) than length of galea-lacinia | 13 |
– | Maxillary palp much longer (> 1.4×) than length of galea-lacinia | L. planus sp. n. |
13(12) |
Gills margin serrate with small denticles and with robust, lanceolate setae (Fig. |
L. branchiaesetis sp. n. |
– |
Gills margin serrate with alternating smaller and bigger denticles and with fine, simple setae (Fig. |
L. podolakae sp. n. |
– |
Gills margin serrate with small denticles and with fine simple setae (Fig. |
14 |
14(13) | Spines at posterior margin of abdominal terga about as long as wide; labrum submarginal dorsal arc of setae with 0–9 setae; maxillary palp segment II with well-developed distolateral excavation; paraproct distally not or slightly expanded (Fig. |
15 |
– | Spines at posterior margin of abdominal terga wider than long; labrum submarginal dorsal arc of setae with 10–14 setae; maxillary palp segment II without distolateral excavation; paraproct distally expanded (Fig. |
L. rutschmannae sp. n. |
15(14) | Spines at posterior margin of abdominal terga mostly rounded; paraproct with > 40 marginal spines; tibia as Fig. |
L. schwanderae sp. n. |
– | Spines at posterior margin of abdominal terga mostly triangular, pointed; paraproct with 31–40 marginal spines; tibia as Fig. |
L. tuberpalpus |
16(2) | Maxillary palp segment II longer than segment I; gills margin serrate with alternating smaller and bigger denticles and with fine, simple setae (Fig. |
L. paravultuosus sp. n. |
– | Maxillary palp segment II about as long or shorter than segment I; gills margin serrate with small denticles and with fine simple setae; fore femur ca. 4–5× as long as wide; labial palp as Fig. |
L. vultuosus |
17(1) | Maxillary palp longer (1.1–1.4×) than length of galea-lacinia; fore femur ca. 2–3× as long as wide | 18 |
– | Maxillary palp much longer (> 1.4×) than length of galea-lacinia; fore femur ca. 4–5× as long as wide | L. lobatus sp. n. |
18(17) | Maxillary palp segment II longer than segment I; spines at posterior margin of abdominal terga mostly triangular, pointed; paraproct distally expanded (Fig. |
L. balkei sp. n. |
– | Maxillary palp segment II about as long or shorter than segment I; spines at posterior margin of abdominal terga mostly rounded; paraproct distally slightly expanded (Fig. |
L. michaeli sp. n. |
19(1) | Spines at posterior margin of abdominal terga mostly rounded; femur ventral without pectinate setae | L. papuaensis sp. n. |
– | Spines at posterior margin of abdominal terga mostly triangular, pointed; femur ventral with pectinate setae (difficult to see) | L. orientis sp. n. |
The distributions of all species known from New Guinea are shown in Figures
Distribution of Labiobaetis in New Guinea: a red circle, L. balkei sp. n.; blue star, L. lobatus sp. n.; yellow circle, L. michaeli sp. n.; red square, L. claudiae sp. n.; yellow square, L. stagnum sp. n.; red triangle, L. orientis sp. n.; yellow triangle, L. papuaensis sp. n.; green star, L. elisae sp. n. b red circle, L. petersorum; yellow circle, L. gladius sp. n.; green circle, L. janae sp. n.; green star, L. vultuosus; red star, L. paravultuosus sp. n.; yellow triangle, L. involutus; red triangle, L. vallus sp. n.; green triangle, L. inopinatus sp. n.
Distribution of Labiobaetis in New Guinea: a red circle, L. tuberpalpus; yellow circle, L. branchiaesetis sp. n.; green circle, L. magnovaldus sp. n.; red triangle, L. planus sp. n.; blue circle, L. podolakae sp. n.; yellow triangle, L. rutschmannae sp. n.; green triangle, L. schwanderae sp. n.; red square, L. centralensis sp. n. b yellow circle, L. paravitilis sp. n.; red circle, L. altus sp. n.; green circle, L. gindroi sp. n.; yellow triangle, L. vitilis; green triangle, L. pindaundensis sp. n.; red square, L. xeniolus; blue star, L. wilhelmemsis sp. n. / L. dendrisetis sp. n. / L. pindaundensis sp. n.
COI sequences were obtained from 20 of the new species (Table
Species | Locality | GPS coordinates | |
---|---|---|---|
L. balkei sp. n. | Papua New Guinea, Central Prov. | 09°01.95'S, 147°44.46'E | |
L. lobatus sp. n. | Papua New Guinea, Central Prov. | 09°00.34'S, 147°44.25'E | |
L. michaeli sp. n. | Papua New Guinea, Eastern Highlands | 05°56.80'S, 145°22.24'E | |
L. claudiae sp. n. | Papua New Guinea, Madang Prov. | 05°24.41'S, 145°38.21'E | |
L. stagnum sp. n. | Indonesia, Papua Prov. | 02°30.51'S, 140°22.83'E | |
L. orientis sp. n. | Papua New Guinea, Eastern Highlands | 07°01.70'S, 145°49.81'E | |
L. papuaensis sp. n. | Indonesia, Papua Prov. | 03°29.80'S, 135°43.89'E | |
L. petersorum | Papua New Guinea, Morobe Prov. | 07°20.30'S, 146°43.72'E | |
L. gladius sp. n. | Papua New Guinea, Simbu Prov. | 05°49.96'S, 145°06.13'E | |
05°48.05'S, 145°04.15'E | |||
05°49.03'S, 145°05.27'E | |||
Papua New Guinea, Western Highlands | 05°15.87'S, 144°32.72'E | ||
Papua New Guinea, Eastern Highlands | 05°56.80'S, 145°22.23'E | ||
L. janae sp. n. | Indonesia, Papua Prov. | 04°07.77'S, 138°40.77'E | |
L. tuberpalpus | Papua New Guinea, Morobe Prov. | 07°20.30'S, 146°43.72'E | |
06°59.00'S, 146°37.00'E | |||
L. branchiaesetis sp. n. | Papua New Guinea, Eastern Highlands | 07°01.70'S, 145°49.81'E | |
Papua New Guinea, Gulf Prov. | 07°03.60'S, 145°44.38'E | ||
L. magnovaldus sp. n. | Papua New Guinea, Simbu Prov. | 05°49.00'S, 145°04.50'E | |
Papua New Guinea, Eastern Highlands | 05°56.80'S, 145°22.24'E | ||
L. planus sp. n. | Indonesia, Papua Prov. | 00°49.35'S, 131°24.20'E | |
L. podolakae sp. n. | Papua New Guinea, Eastern Highlands | 06°21.41'S, 145°54.34'E | |
L. rutschmannae sp. n. | PapuaNewGuinea,WesternHighlands | 05°14.28'S, 144°28.74'E | |
05°15.17'S, 144°32.81'E | |||
L. schwanderae sp. n. | Papua New Guinea, Gulf Prov. | 07°05.66'S, 145°44.47'E | |
L. vitilis | Papua New Guinea, Western Highlands | 05°54.91'S, 143°59.06'E | |
L. altus sp. n. | Papua New Guinea, Enga Prov. | 05°47.55'S, 143°58.76'E | |
Papua New Guinea, Simbu Prov. | 05°49.00'S, 145°04.50'E | ||
L. gindroi sp. n. | Indonesia, Papua Prov. | 03°56.95'S, 138°54.38'E | |
Papua New Guinea, Western Highlands | 05°15.17'S, 144°32.81'E | ||
L. paravitilis sp. n. | Papua New Guinea, Madang Prov. | 05°18.09'S, 145°36.45'E | |
L. wilhelmensis sp. n | Papua New Guinea, Simbu Prov. | 05°49.00'S, 145°04.50'E | |
05°48.05'S, 145°04.15'E | |||
L. vultuosus | Papua New Guinea, Western Highlands | 05°51.00'S, 144°14.72'E | |
L. paravultuosus sp. n. | Papua New Guinea, Enga Prov. | 05°38.11'S, 143°55.34'E | |
L. centralensis sp. n. | Papua New Guinea, Central Prov. | 09°14.34'S, 147°36.92'E | |
L. dendrisetis sp. n. | Papua New Guinea, Simbu Prov. | 05°49.00'S, 145°04.50'E | |
L. elisae sp. n. | Papua New Guinea, Western Highlands | 05°16.10'S, 144°27.87'E | |
Papua New Guinea, Madang Prov. | 05°13.39'S, 144°37.29'E | ||
Papua New Guinea, Eastern Highlands | 06°11.02'S, 145°26.41'E | ||
L. inopinatus sp. n. | Papua New Guinea, Gulf Prov. | 07°37'S | 146°04'E |
L. pindaundensis sp. n. | Papua New Guinea, Simbu Prov. | 05°49.00'S, 145°04.50'E | |
05°49.03'S, 145°05.27'E | |||
Papua New Guinea, Central Prov. | 08°31.29'S, 147°13.68'E | ||
L. vallus sp. n. | Papua New Guinea, Madang Prov. | 04°43.06'S, 145°24.44'E | |
L. involutus | Papua New Guinea, Morobe Prov. | 07°20.30'S, 146°43.72'E | |
L. xeniolus | Papua New Guinea, Morobe Prov. | 07°13.00'S, 146°49.00'E |
Genetic distances (COI) between sequenced specimens, using the Kimura 2-parameter.
1 | 2 | 3 | 4 | 5 | 6 | 7 | 8 | 9 | 10 | 11 | 12 | 13 | 14 | 15 | 16 | 17 | 18 | 19 | 20 | 21 | 22 | 23 | 24 | 25 | 26 | ||
---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|
1 | L. balkei sp. n. | ||||||||||||||||||||||||||
2 | L. balkei sp. n. | 0,00 | |||||||||||||||||||||||||
3 | L. michaeli sp. n. | 0,26 | 0,26 | ||||||||||||||||||||||||
4 | L. michaeli sp. n. | 0,26 | 0,26 | 0,00 | |||||||||||||||||||||||
5 | L. michaeli sp. n. | 0,28 | 0,28 | 0,05 | 0,05 | ||||||||||||||||||||||
6 | L. lobatus sp. n. | 0,20 | 0,20 | 0,28 | 0,28 | 0,28 | |||||||||||||||||||||
7 | L. claudiae sp. n. | 0,21 | 0,21 | 0,25 | 0,25 | 0,27 | 0,24 | ||||||||||||||||||||
8 | L. stagnum sp. n. | 0,22 | 0,22 | 0,27 | 0,27 | 0,26 | 0,22 | 0,21 | |||||||||||||||||||
9 | L. orientis sp. n. | 0,26 | 0,26 | 0,27 | 0,27 | 0,26 | 0,24 | 0,18 | 0,22 | ||||||||||||||||||
10 | L. papuaensis sp. n. | 0,27 | 0,27 | 0,25 | 0,25 | 0,25 | 0,24 | 0,20 | 0,19 | 0,21 | |||||||||||||||||
11 | L. gladius sp. n. | 0,22 | 0,22 | 0,25 | 0,25 | 0,27 | 0,24 | 0,19 | 0,23 | 0,24 | 0,20 | ||||||||||||||||
12 | L. janae sp. n. | 0,24 | 0,24 | 0,29 | 0,29 | 0,29 | 0,24 | 0,23 | 0,23 | 0,23 | 0,24 | 0,23 | |||||||||||||||
13 | L. janae sp. n. | 0,24 | 0,24 | 0,29 | 0,29 | 0,29 | 0,24 | 0,23 | 0,23 | 0,23 | 0,24 | 0,23 | 0,00 | ||||||||||||||
14 | L. branchiaesetis sp. n. | 0,20 | 0,20 | 0,26 | 0,26 | 0,25 | 0,24 | 0,18 | 0,21 | 0,22 | 0,23 | 0,19 | 0,19 | 0,19 | |||||||||||||
15 | L. planus sp. n. | 0,22 | 0,22 | 0,25 | 0,25 | 0,25 | 0,24 | 0,18 | 0,16 | 0,19 | 0,17 | 0,18 | 0,18 | 0,18 | 0,18 | ||||||||||||
16 | L. planus sp. n. | 0,22 | 0,22 | 0,25 | 0,25 | 0,25 | 0,24 | 0,18 | 0,16 | 0,19 | 0,17 | 0,18 | 0,18 | 0,18 | 0,18 | 0,00 | |||||||||||
17 | L. podolakae sp. n. | 0,25 | 0,25 | 0,25 | 0,25 | 0,25 | 0,24 | 0,20 | 0,19 | 0,22 | 0,20 | 0,18 | 0,20 | 0,20 | 0,14 | 0,17 | 0,17 | ||||||||||
18 | L. schwanderae sp. n. | 0,21 | 0,21 | 0,25 | 0,25 | 0,25 | 0,24 | 0,18 | 0,18 | 0,19 | 0,19 | 0,16 | 0,22 | 0,22 | 0,19 | 0,13 | 0,13 | 0,19 | |||||||||
19 | L. altus sp. n. | 0,22 | 0,22 | 0,29 | 0,29 | 0,30 | 0,24 | 0,20 | 0,18 | 0,15 | 0,20 | 0,22 | 0,24 | 0,24 | 0,22 | 0,18 | 0,18 | 0,20 | 0,20 | ||||||||
20 | L. gindroi sp. n. | 0,26 | 0,26 | 0,24 | 0,24 | 0,25 | 0,24 | 0,21 | 0,19 | 0,20 | 0,19 | 0,22 | 0,19 | 0,19 | 0,20 | 0,20 | 0,20 | 0,20 | 0,18 | 0,16 | |||||||
21 | L. paravitilis sp. n. | 0,24 | 0,24 | 0,27 | 0,27 | 0,26 | 0,24 | 0,24 | 0,20 | 0,24 | 0,21 | 0,21 | 0,22 | 0,22 | 0,17 | 0,21 | 0,21 | 0,21 | 0,20 | 0,19 | 0,20 | ||||||
22 | L. paravultuosus sp. n. | 0,22 | 0,22 | 0,24 | 0,24 | 0,24 | 0,24 | 0,20 | 0,18 | 0,21 | 0,18 | 0,21 | 0,21 | 0,21 | 0,19 | 0,15 | 0,15 | 0,19 | 0,19 | 0,19 | 0,23 | 0,20 | |||||
23 | L. paravultuosus sp. n. | 0,23 | 0,23 | 0,26 | 0,26 | 0,26 | 0,24 | 0,23 | 0,20 | 0,23 | 0,20 | 0,23 | 0,22 | 0,22 | 0,21 | 0,17 | 0,17 | 0,21 | 0,21 | 0,21 | 0,25 | 0,21 | 0,02 | ||||
24 | L. centralensis sp. n. | 0,22 | 0,22 | 0,31 | 0,31 | 0,29 | 0,24 | 0,21 | 0,22 | 0,25 | 0,23 | 0,23 | 0,20 | 0,20 | 0,24 | 0,22 | 0,22 | 0,21 | 0,22 | 0,23 | 0,22 | 0,26 | 0,26 | 0,27 | |||
25 | L. centralensis sp. n. | 0,23 | 0,23 | 0,32 | 0,32 | 0,30 | 0,24 | 0,21 | 0,23 | 0,24 | 0,23 | 0,23 | 0,20 | 0,20 | 0,24 | 0,22 | 0,22 | 0,21 | 0,21 | 0,23 | 0,22 | 0,25 | 0,26 | 0,27 | 0,01 | ||
26 | L. elisae sp. n. | 0,22 | 0,22 | 0,28 | 0,28 | 0,27 | 0,24 | 0,19 | 0,20 | 0,21 | 0,19 | 0,21 | 0,23 | 0,23 | 0,22 | 0,17 | 0,17 | 0,19 | 0,18 | 0,19 | 0,19 | 0,19 | 0,20 | 0,21 | 0,21 | 0,21 | |
27 | L. vallus sp. n. | 0,26 | 0,26 | 0,25 | 0,25 | 0,26 | 0,24 | 0,18 | 0,18 | 0,24 | 0,19 | 0,21 | 0,19 | 0,19 | 0,17 | 0,17 | 0,17 | 0,18 | 0,18 | 0,21 | 0,16 | 0,19 | 0,21 | 0,23 | 0,20 | 0,20 | 0,20 |
For the attribution of the new species to Labiobaetis we are referring to
The seven species groups proposed in this paper are mainly based on the combination of two characters: the kind of setae composing the dorsal, submarginal arc of setae on the labrum and the shape of the distomedial protuberance of labial palp segment II, sometimes together with other additional characters. An exception is the petersorum group, where the main character is the blade-like outermost denticle of the mandibles. Species sharing combinations of character states as the type of setae forming the dorsal, submarginal arc of setae or the shape of the labial palp segment II are often very similar in most other characters, which justify the formation of species groups. These morphological groups within Labiobaetis are primarily a working tool, but could also serve as a basis for future studies on the generic delimitation and phylogeny of this probably polyphyletic genus. The inclusion of nuclear gene sequences may prove that some of them may be natural groups. In other realms, especially the Oriental realm, some species share the main characters of some of the proposed species groups: L. molawinensis (Müller-Liebenau) from the Philippines and Taiwan and L. atrebatinus (Eaton) from Eurasia have spatulate, apically pectinate setae composing the dorsal, submarginal arc of setae of the labrum and a large, lobed distomedial protuberance of the labial palp segment II like the species of the balkei group. They also share other characters such as the distolateral excavation of maxillary palp segment II, the shape of labial palp segment III, which is slightly pentagonal and apically slightly pointed, and the arrangement of outer and inner sets of denticles of the mandibles, which are as usually fused, but have a small gap between them. Therefore both species should be considered as members of that group. They are both differentiated from the species in New Guinea by a distolateral process at the antennal scape and L. atrebatinus additionally by the presence of hindwing pads and seven pairs of gills (
In general, the genetic distances between the different species of Labiobaetis are rather high in New Guinea, on average 22% (K2P, Table
In addition to the 26 new species described in this paper we obtained five other different COI sequences with clearly interspecific genetic distance to other species with similar morphology: two sequences L. cf. balkei sp. n. (K2P 16%–30%), two sequences L. cf. vultuosus (K2P 16%–21%) and one sequence L. cf. tuberpalpus (K2P 13%). Based on the existing material we are not in a position to distinguish these specimens morphologically from all species within their morphological group. Therefore, they have to remain species hypotheses for now without further treatment in this paper. Additional material will be necessary to confirm their status in the future.
The Labiobaetis species distribution seems to be restricted to New Guinea, as far as we know, which is fully in line with other Baetidae genera having a species endemicity of close to 100% in the Australasian realm (Gattoliat and Nieto 2009). In Madagascar there is a similar situation with eight species, which are all restricted to this island (
Our recent knowledge shows high levels of micro-endemism restricted to smaller areas in New Guinea. This indicates that allopatry could be a major driver of diversity in the genus. Large studies about the highly diversified genus Exocelina Balke, 1998 (Coleoptera, Dytiscidae) demonstrated allopatry to be the main mechanism of diversification in New Guinea and found strong evidence that recent environmental change in the extremely structured central highlands of New Guinea with its ongoing formation of rich aquatic resources and remote valleys and mountain blocks was the primary driver of diversification in that area (
Despite covering an important part of New Guinea, the sampling effort and the number of localities and different habitats is still limited and there are huge areas without any collection activities so far. Additionally we have already five species hypotheses based on genetics only in the present material, which may be confirmed as separate species in the future. Therefore we may assume, that the number of Labiobaetis species in New Guinea will continue to increase substantially with further collections in the future.
We sincerely thank Michael Balke (Zoologische Staatssammlung München,