Research Article |
Corresponding author: Thomas Kaltenbach ( thomas.kaltenbach@bluewin.ch ) Academic editor: Lyndall Pereira-da-Conceicoa
© 2020 Thomas Kaltenbach, Jhoana M. Garces, 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, Garces J, Gattolliat J-L (2020) The success story of Labiobaetis Novikova & Kluge in the Philippines (Ephemeroptera, Baetidae), with description of 18 new species. ZooKeys 1002: 1-114. https://doi.org/10.3897/zookeys.1002.58017
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Material collected between 1994 and 2020 in the Philippines, covering most main islands like Luzon, Mindoro, Palawan, Negros, Cebu, Leyte, and Mindanao and some smaller islands, substantially increased our knowledge of Labiobaetis Novikova & Kluge in this archipelago. Only three species were previously reported: L. molawinensis (Müller-Liebenau, 1982) and L. sumigarensis (Müller-Liebenau, 1982) from larvae and L. boettgeri (Ulmer, 1924) from adults. Eighteen new species have been identified using a combination of morphology and genetic distance (COI, Kimura 2-parameter). They are described and illustrated based on their larvae and a key to all species in the Philippines is provided. The total number of Labiobaetis in the Philippines has increased to 21 species. Additional diversity of Labiobaetis based on molecular evidence only is presented as Molecular Operational Taxonomic Units (MOTUs) without description. The interspecific K2P distances in the Philippines are between 15% and 27%, the intraspecific distances are usually between 0% and 3%. The total number of Labiobaetis species worldwide is augmented to 144.
COI, genetic distance, integrated taxonomy, Molecular Operational Taxonomic Unit (MOTU), Southeast Asia
The family Baetidae has the highest species diversity among mayflies, comprising 1,070 species in 110 genera (
The genus Labiobaetis Novikova & Kluge (
Recently, integrative taxonomy was done on collections from the diverse and poorly explored Southeast Asia and New Guinea regions where 47 species were described and named (
The megadiversity of the Philippines is partly attributed to the complex biogeographic history and isolation of the archipelago. The discussion of the biogeographic history of the Philippine archipelago includes landmass movements, collisions between landmasses of different origin in Miocene, and temporary Pleistocene land bridges which were possible colonization pathways of species. Originally part of the Eurasian continent, the oldest landmasses of the current Philippines are parts of Palawan, Mindoro, Romblon and Panay. Whether these landmasses were entirely submerged during the drift is still a matter of debate (
The diversity of Labiobaetis in the Philippines was poorly known, as only two species were previously reported from larvae (L. molawinensis and L. sumigarensis by
All specimens were collected between 1994 and 2020 by Dr. Hendrik Freitag and his team (Ateneo de Manila University) and preserved in 70%–96% ethanol.
The dissection of larvae was done in Cellosolve (2-Ethoxyethanol) with subsequent mounting on slides with Euparal liquid, using an Olympus SZX7 stereomicroscope.
The DNA of part of the specimens was extracted using non-destructive methods allowing subsequent morphological analysis (see
The GenBank accession numbers are given in Table
Species | Species group | Locality | Specimens catalog # | GenBank # | GenSeq |
---|---|---|---|---|---|
(COI) | Nomenclature | ||||
L. dalisay sp. nov. | dendrisetis gr. | Philippines: Luzon | GBIFCH 00763649 | MT830940 | genseq-2 COI |
L. acei sp. nov. | numeratus gr. | Philippines: Luzon | GBIFCH 00763643 | MT830941 | genseq-1 COI |
GBIFCH 00763645 | MT830942 | genseq-2 COI | |||
GBIFCH 00763651 | MT830943 | genseq-2 COI | |||
L. aldabae sp. nov. | numeratus gr. | Philippines: Luzon | GBIFCH 00654913 | MT830944 | genseq-1 COI |
GBIFCH 00654908 | MT830945 | genseq-2 COI | |||
GBIFCH 00763646 | MT830946 | genseq-2 COI | |||
GBIFCH 00763648 | MT830947 | genseq-2 COI | |||
Philippines: Negros | GBIFCH 00654889 | MT830948 | genseq-2 COI | ||
L. camiguinensis sp. nov. | numeratus gr. | Philippines: Camiguin | GBIFCH 00654915 | MT830949 | genseq-1 COI |
L. lachicae sp. nov. | numeratus gr. | Philippines: Mindanao | GBIFCH 00654891 | MT830950 | genseq-1 COI |
L. palawano sp. nov. | numeratus gr. | Philippines: Busuanga | GBIFCH 00763688 | MT830987 | genseq-1 COI |
Philippines: Palawan | GBIFCH 00763679 | MT830988 | genseq-2 COI | ||
L. sabordoi sp. nov. | numeratus gr. | Philippines: Negros | GBIFCH 00654878 | MT830951 | genseq-2 COI |
Philippines: Romblon | GBIFCH 00763674 | MT830952 | genseq-2 COI | ||
L. gamay sp. nov. | operosus gr. | Philippines: Mindoro | GBIFCH 00654922 | MT830953 | genseq-2 COI |
GBIFCH 00763637 | MT830954 | genseq-2 COI | |||
GBIFCH 00763639 | MT830955 | genseq-2 COI | |||
Philippines: Luzon | GBIFCH 00763655 | MT830956 | genseq-2 COI | ||
GBIFCH 00763657 | MT830957 | genseq-2 COI | |||
GBIFCH 00763658 | MT830958 | genseq-2 COI | |||
L. pangantihoni sp. nov. | operosus gr. | Philippines: Palawan | GBIFCH 00763684 | MT830959 | genseq-2 COI |
L. tagbanwa sp. nov. | operosus gr. | Philippines: Palawan | GBIFCH 00654885 | MT830960 | genseq-2 COI |
GBIFCH 00763681 | MT830961 | genseq-2 COI | |||
GBIFCH 00763680 | MT830962 | genseq-2 COI | |||
L. valdezorum sp. nov. | operosus gr. | Philippines: Negros | GBIFCH 00654888 | MT830963 | genseq-1 COI |
GBIFCH 00654882 | MT830964 | genseq-2 COI | |||
GBIFCH 00654879 | MT830965 | genseq-2 COI | |||
GBIFCH 00654880 | MT830966 | genseq-2 COI | |||
Philippines: Cebu | GBIFCH 00763671 | MT830967 | genseq-2 COI | ||
L. wantzeni sp. nov. | operosus gr. | Philippines: Camiguin | GBIFCH 00654898 | MT830968 | genseq-1 COI |
GBIFCH 00654897 | MT830969 | genseq-2 COI | |||
GBIFCH 00763641 | MT830970 | genseq-2 COI | |||
GBIFCH 00763642 | MT830971 | genseq-2 COI | |||
GBIFCH 00654896 | MT830972 | genseq-2 COI | |||
GBIFCH 00654900 | MT830973 | genseq-2 COI | |||
L. baganii sp. nov. | sumigarensis gr. | Philippines: Mindanao | GBIFCH 00654895 | MT830974 | genseq-1 COI |
Philippines: Camiguin | GBIFCH 00654899 | MT830975 | genseq-2 COI | ||
L. delocadoi sp. nov. | sumigarensis gr. | Philippines: Cebu | GBIFCH 00654886 | MT830976 | genseq-1 COI |
Philippines: Leyte | GBIFCH 00763668 | MT830977 | genseq-2 COI | ||
L. freitagi sp. nov. | sumigarensis gr. | Philippines: Palawan | GBIFCH 00763677 | MT830978 | genseq-2 COI |
GBIFCH 00763678 | MT830979 | genseq-2 COI | |||
GBIFCH 00763682 | MT830980 | genseq-2 COI | |||
GBIFCH 00763683 | MT830981 | genseq-2 COI | |||
L. pelingeni sp. nov. | sumigarensis gr. | Philippines: Negros | GBIFCH 00654901 | MT830982 | genseq-2 COI |
Philippines: Cebu | GBIFCH 00763672 | MT830983 | genseq-2 COI | ||
L. giselae sp. nov. | vallus gr. | Philippines: Luzon | GBIFCH 00654911 | MT830984 | genseq-2 COI |
L. mendozai sp. nov. | vallus gr. | Philippines: Mindanao | GBIFCH 00654894 | MT830985 | genseq-2 COI |
The nomenclature used for Molecular Operational Taxonomic Units (MOTUs) is somewhat different as the one proposed by
Drawings were made using an Olympus BX43 microscope. To facilitate the determination of species and the comparison of important structures, we partly used a combination of dorsal and ventral aspects in one drawing. Explanations are given in Fig.
Photographs of larvae were taken using a Canon EOS 6D camera and the Visionary Digital Passport imaging system (http://www.duninc.com) and processed with 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 maps were generated with SimpleMappr (https://simplemappr.net,
The taxonomic descriptions were generated with a DELTA (
The dichotomous key was elaborated with the support of DKey version 1.3.0 (http://drawwing.org/dkey,
The terminology follows
Labiobaetis, character states of selected characters a–e setae of the submarginal arc on the dorsal surface of the labrum: a simple b feathered c clavate d dendritic e lanceolate f–h distolateral process at scape of antenna: f absent g poorly developed h well developed i–m labial palp, distomedial protuberance of segment II: i thumb-like j broad thumb-like k slender thumb-like l hook-like m small thumb-like n–p distolateral excavation at maxillary palp segment II: n well developed o poorly developed p absent q–t hypopharynx, medial tuft of stout setae: q well developed, long r well developed, average length s well developed, short t poorly developed u–x hind protoptera: u absent v minute w small x well developed y, z paraproct: y distally not expanded z distally expanded.
Abbreviations:
AdMU Ateneo de Manila University, Quezon City (Philippines)
MZL Musée de Zoologie Lausanne (Switzerland)
PCSD Palawan Council for Sustainable Development, Puerto Princesa, Palawan (Philippines)
dendrisetis group
1. L. dalisay sp. nov.
numeratus group
2. L. acei sp. nov.
3. L. aldabae sp. nov.
4. L. camiguinensis sp. nov.
5. L. lachicae sp. nov.
6. L. palawano sp. nov.
7. L. sabordoi sp. nov.
operosus group
8. L. gamay sp. nov.
9. L. pangantihoni sp. nov.
10. L. tagbanwa sp. nov.
11. L. valdezorum sp. nov.
12. L. wantzeni sp. nov.
sumigarensis group
13. L. molawinensis (Müller-Liebenau, 1982)
14. L. sumigarensis (Müller-Liebenau, 1982)
15. L. baganii sp. nov.
16. L. delocadoi sp. nov.
17. L. freitagi sp. nov.
18. L. pelingeni sp. nov.
vallus group
19. L. giselae sp. nov.
20. L. mendozai sp. nov.
Not assigned to a group
21. L. boettgeri (Ulmer, 1924), no further treatment in this study.
Following combination of characters: A) dorsal surface of labrum with submarginal arc of dendritic setae; B) labial palp segment II slender or small thumb-like; C) labial palp segment III wide; D) maxillary palp shorter than galea-lacinia, rather thick; E) seven pairs of gills.
Larva. Following combination of characters: A) dorsal surface of labrum with 6–8 long, dendritic setae; B) labial palp segment II with a slender thumb-like distomedial protuberance, segment III subrectangular; C) right mandible without row of thin setae at inner margin of innermost denticle; D) fore femur length 3.4 × maximum width, dorsal margin with 10–19 curved, spine-like setae; E) hind protoptera well developed; F) paraproct distally not expanded, with ca. 15 stout, marginal spines.
Larva (Figs
Colouration. Head, thorax, and abdomen dorsally brown with bright pattern as in Fig.
Antenna
(Fig.
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 and superlinguae
(Fig.
Maxilla
(Fig.
Labium
(Fig.
Hind protoptera
(Fig.
Foreleg
(Fig.
Terga
(Fig.
Gills
(Fig.
Paraproct
(Fig.
Named after the Filipino word dalisay meaning pristine, which describes the localities where the species was collected.
Philippines: Luzon (Fig.
The specimens were collected at altitudes from 60 m to 400 m, mainly in pristine areas.
Holotype. Philippines • larva; Luzon, Maria Aurora, Wenceslao, Bingwangan River; 15°45'48"N, 121°25'21"E; 60 m; 05.II.1998; leg. Mendoza; on slide; GBIFCH 00592279;
Following combination of characters: A) dorsal surface of labrum with submarginal arc of simple setae, 1st and 2nd setae after submedian seta close together; B) labial palp segment II with thumb-like distomedial protuberance; C) glossae with robust setae at inner margin; D) paraglossae with setae at anterolateral and lateral outer margin; E) right mandible with a pronounced hump between prostheca and mola, thin setae at base of mola; F) left mandible with convex margin between prostheca and mola; G) maxillary palp segment II much longer vs. segment I, bent; H) superlinguae sclerotized along margins (Fig.
Larva. Following combination of characters: A) dorsal surface of labrum with submarginal arc of six or seven long, simple setae; B) labial palp segment II with a thumb-like distomedial protuberance, segment III conical; C) left mandible with a comb-shaped structure at base of mola; D) fore femur rather broad, length ca. 3 × maximum width, dorsal margin with 9–12 curved, spine-like setae and a partial second row near margin; E) tergum IV with rounded, partly fused spines at posterior margin, surface with irregular, dense rows of U-shaped scale bases; F) paraproct distally not expanded, with 29–34 stout, marginal spines.
Larva (Figs
Colouration. Head, thorax, and abdomen dorsally brown, with pattern as in Fig.
Antenna
(Fig.
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 and superlinguae
(Fig.
Maxilla
(Fig.
Labium
(Fig.
Hind protoptera
(Fig.
Foreleg
(Fig.
Terga
(Fig.
Gills
(Fig.
Paraproct
(Fig.
Dedicated to Mr. Ace Kevin Amarga (Philippines/Taiwan), an outstanding collector and entomologist, for loaning some precious material to one of the authors (JG).
Philippines: Luzon (Fig.
The specimens were collected at altitudes of 1700 m and 1820 m, either on rock surface or bottom gravel in riffles or runs, or in root packs or grass bunches in riffles or runs.
Holotype. Philippines • larva; Luzon, Mountain Province, Bauko, mineral rich mountain creek; 17°03'53"N, 121°05'10"E; 1820 m; XI.1997; leg. Mey; on slide; GenBank: MT830941; GBIFCH 00763643;
Larva. Following combination of characters: A) dorsal surface of labrum with submarginal arc of 4–6 long, simple setae; B) labial palp segment II with a thumb-like distomedial protuberance, segment III conical; C) left mandible with a comb-shaped structure at base of mola; D) fore femur rather broad, length 3.4 × maximum width, with ca. nine curved, spine-like setae and a partial second row near margin; E) claw with 12–15 denticles; F) paraproct distally not expanded, with 18–21 stout, marginal spines.
Larva (Figs
Colouration. Head dorsally light brown, thorax and abdomen dorsally brown with bright pattern as in Fig.
Antenna
(Fig.
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 and superlinguae
(Fig.
Maxilla
(Fig.
Labium
(Fig.
Hind protoptera
(Fig.
Foreleg
(Fig.
Terga
(Fig.
Gills
(Fig.
Paraproct
(Fig.
Dedicated to Ms. Kyra Mari Dominique Aldaba (Philippines), member of the AdMU Biodiversity Laboratory.
Philippines: Luzon and Negros (Fig.
The specimens were collected at altitudes from 50 m to 1400 m, mainly in riverside pools, but also in root packs or grass bunches in the runs.
Holotype. Philippines • larva; Luzon, Laguna, Samil River; 14°08'N, 121°31'E; 370 m; 27.VI.2018; leg. BIO-PHIL exped.; on slide; GenBank: MT830944; GBIFCH 00654913;
Philippines • 27 larvae; Luzon, Ifugao, Tinoc; 16°40'58"N, 120°56'59"E; 1400 m; XI.1997; leg. Mey; in alcohol; GBIFCH 00515444; AdMU • 4 larvae; Luzon, Ilocos Sur, Suyo Municipality, Tagudin-Cervantes-Sabangan Road, Besang Pass Area; 16°57'17"N, 120°38'52"E; 1200 m; 15.IV.2019; leg. Freitag, Garces and Pangantihon; in alcohol; GBIFCH 00515442;
Larva. Following combination of characters: A) dorsal surface of labrum with submarginal arc of one plus four or five long, simple setae; B) labial palp segment II with thumb-like distomedial protuberance; C) fore femur rather broad, length ca. 3 × maximum width, dorsal margin with ca. ten curved, spine-like setae and a partial second row near margin; D) claw with 14 or 15 denticles; E) paraproct distally not expanded, with 15–17 stout, marginal spines.
Larva (Figs
Colouration. Head dorsally light brown, thorax and abdomen dorsally brown with bright pattern as in Fig.
Antenna
(Fig.
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 and superlinguae
(Figs
Maxilla
(Fig.
Labium
(Fig.
Hind protoptera
(Fig.
Foreleg
(Fig.
Terga
(Fig.
Gills
(Fig.
Paraproct
(Fig.
Refers to the island Camiguin, where the specimens were collected.
Philippines: Camiguin (Fig.
The specimens were collected at altitudes of 30 m and 900 m, mainly on bottom gravel.
Holotype. Philippines • larva; Camiguin, Sagay, Bonbon, lower Binangawan River; 09°06'39"N 124°43'45"E; 30 m; 09.XII.2018; leg. Freitag and Wantzen; on slide; GenBank: MT830949; GBIFCH 00654915;
Larva. Following combination of characters: A) dorsal surface of labrum with submarginal arc of one plus 5–7 long, simple setae; B) labial palp segment II with a thumb-like distomedial protuberance, segment III conical; C) left mandible without setae at apex of mola; D) fore femur rather broad, length ca. 3 × maximum width, dorsal margin with 10–12 curved, spine-like setae and a partial second row near margin; E) claw with 15–17 denticles; F) paraproct distally not expanded, with ca. 13 stout, marginal spines.
Larva (Figs
Colouration. Head dorsally light brown, thorax and abdomen dorsally brown, with bright pattern as in Fig.
Antenna
(Fig.
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 and superlinguae
(Fig.
Maxilla
(Fig.
Labium
(Fig.
Hind protoptera
(Fig.
Foreleg
(Fig.
Terga
(Fig.
Gills
(Fig.
Paraproct
(Fig.
Dedicated to Ms. Maria Kenosis Lachica (Philippines/Japan), friend of the AdMU Biodiversity Laboratory.
Philippines: Mindanao (Fig.
The specimens were collected at altitudes from sea level to 120 m, in root packs, grass bunches or submerged wood in riffles or runs.
Holotype. Philippines • larva; Mindanao, Agusan del Sur, San Francisco, Bayogan, Tagkunayan Creek; 08°28'N, 125°59'E; 120 m; 05.II.1998; leg. Mendoza; on slide; GenBank: MT830950; GBIFCH 00654891;
Larva. Following combination of characters: A) dorsal surface of labrum with submarginal arc of one plus seven long, simple setae; B) labial palp segment II with a thumb-like distomedial protuberance, segment III conical; C) mola of right mandible proximally beginning with a double hump; D) hypopharynx with medial tuft of stout setae poorly developed; E) fore femur rather broad, length ca. 3 × maximum width, dorsal margin with 10–13 curved, spine-like setae and a partial second row near margin; F) claw with 15–17 denticles; G) posterior margin of tergum IV with rounded spines, wider than long.
Larva (Figs
Colouration. Head, thorax, and abdomen dorsally brown, with dark brown markings as in Fig.
Antenna
(Fig.
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 and superlinguae
(Fig.
Maxilla
(Fig.
Labium
(Fig.
Hind protoptera
(Fig.
Foreleg
(Fig.
Terga
(Fig.
Gills
(Fig.
Paraproct
(Fig.
Dedicated to the indigenous Palawano people of Palawan.
Philippines: Palawan (incl. Busuanga) (Fig.
The specimens were collected at altitudes below 100 m, partly on rock surface in riffles or runs.
Holotype. Philippines • larva; Palawan, P. Princesa, Bindujan, Talabigan River; 10°01'26"N, 119°04'37"E; 10 m; 27.VII.2019; leg. Freitag and Molls; on slide; GenBank: MT830987; GBIFCH 00763688;
Larva. Following combination of characters: A) dorsal surface of labrum with submarginal arc of one plus five or six long, simple setae; B) labial palp segment II with a thumb-like distomedial protuberance, segment III conical; C) mola of right mandible proximally beginning with a double hump; D) hypopharynx with medial tuft of stout setae poorly developed; E) fore femur rather broad, length ca. 3 × maximum width, dorsal margin with 11–14 curved, spine-like setae and a partial second row near margin; F) claw with 16–18 denticles; G) posterior margin of tergum IV with triangular spines, wider than long.
Larva (Figs
Colouration. Head dorsally light brown with brown markings, thorax and abdomen dorsally brown with bright pattern as in Fig.
Antenna
(Fig.
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 and superlinguae
(Fig.
Maxilla
(Fig.
Labium
(Fig.
Hind protoptera
(Fig.
Foreleg
(Fig.
Terga
(Fig.
Gills
(Fig.
Paraproct
(Fig.
Dedicated to Mr. Marc Ryan Sabordo (Philippines), collector and project assistant of the AdMU Biodiversity Laboratory.
Philippines: Negros, Sibuyan and Tablas (Fig.
The specimens were collected at altitudes from sea level to 480 m, partly in leaf litter.
Holotype. Philippines • larva; Negros Oriental, Valencia, Casaroro River downstream; 09°18'N, 123°14'E; 150 m; 01.IX.2019; leg. Garces and Pelingen; on slide; GBIFCH 00592270;
Following combination of characters: A) dorsal surface of labrum with submarginal arc of feathered setae; B) labial palp segment II with thumb-like or lobed distomedial protuberance; C) seven pairs of gills; D) hind protoptera well developed; E) distolateral process at scape well developed; F) fore tarsus with thin setae at ventrodistal margin.
Larva. Following combination of characters: A) dorsal surface of labrum with submarginal arc of one plus 7–9 feathered setae with strongly reduced feathers; B) labial palp segment II with a broad, thumb-like distomedial protuberance; segment III conical; C) fore femur rather broad, length ca. 3 × maximum width, dorsal margin with 10–14 curved, spine-like setae; D) hind protoptera well developed; E) paraproct distally not expanded, with 19–22 stout marginal spines.
Larva (Figs
Colouration. Head, thorax, and abdomen dorsally grey-brown, with bright pattern as in Fig.
Antenna
(Fig.
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 and superlinguae
(Fig.
Maxilla
(Fig.
Labium
(Fig.
Hind protoptera
(Fig.
Foreleg
(Fig.
Terga
(Fig.
Gills
(Fig.
Paraproct
(Fig.
Named after the Cebuano word meaning reduced, referring to the strongly reduced feathers of the submarginal setae on the dorsal labrum surface.
Philippines: Luzon and Mindoro (Fig.
The specimens were collected at altitudes from sea level to 140 m, partly on hygropetric rocks or rock surface in riffles or runs.
Holotype. Philippines • larva; Oriental Mindoro, Roxas, Brgy. San Vicente, lower reach of Taugad River; 12°37.30'N, 121°22.97'E; 140 m; 2016–2019; leg. Freitag and Garces; on slide; GBIFCH 00592274;
Larva. Following combination of characters: A) dorsal surface of labrum with submarginal arc of one plus nine or ten long, feathered setae; B) labial palp segment II with a rather slender, thumb-like distomedial protuberance, segment III slightly pentagonal; C) fore femur rather broad, length 3.4 × maximum width, dorsal margin with eleven or 12 curved, spine-like setae; D) claw with 10–13 denticles; E) paraproct distally not expanded, with ca. 16 stout marginal spines.
Larva (Figs
Colouration. Head, thorax, and abdomen dorsally brown, with bright pattern as in Fig.
Antenna
(Fig.
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 and superlinguae
(Fig.
Maxilla
(Fig.
Labium
(Fig.
Hind protoptera
(Fig.
Foreleg
(Fig.
Terga
(Fig.
Gills
(Fig.
Paraproct
(Fig.
Dedicated to Mr. Clister V. Pangantihon, outstanding collector, entomologist and project assistant of the AdMU Biodiversity Laboratory.
Philippines: Palawan (Fig.
The specimens were collected at altitudes from sea level to 180 m, partly on submerged wood in runs or riffles.
Holotype. Philippines • larva; Palawan, Balabac, Danglis River near the road; 07°57'39"N, 117°02'59"E; 170 m; 2019; leg. Pelingen and Pangantihon; on slide; GBIFCH 00592336;
Larva. Following combination of characters: A) dorsal surface of labrum with submarginal arc of one plus 8–10 long, feathered setae with reduced feathers; B) labial palp segment II with a broad thumb-like distomedial protuberance, segment III conical; C) fore femur rather broad, length ca. 3 × maximum width, dorsal margin with ca. 13 curved, spine-like setae; D) hind protoptera medium developed; E) paraproct distally not expanded, with ca. 18 stout, marginal spines.
Larva (Figs
Colouration.
Head, thorax, and abdomen dorsally brown, with bright markings as in Fig.
Antenna
(Fig.
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 and superlinguae
(Fig.
Maxilla
(Fig.
Labium
(Fig.
Hind protoptera
(Fig.
Foreleg
(Fig.
Terga
(Fig.
Gills
(Fig.
Paraproct
(Fig.
Dedicated to the indigenous Tagbanwa people of Palawan.
Philippines: Palawan (Fig.
The specimens were collected from sea level to 100 m on rock surface or submerged wood in riffles or runs.
Holotype. Philippines • larva; Palawan, Quezon, Aramaywan River, cogon grass; 09°22'33"N, 118°08'41"E; 15 m; 2019; leg. Pangantihon and Pelingen; on slide; GBIFCH 00592324;
Larva. Following combination of characters: A) dorsal surface of labrum with submarginal arc of one plus 9–12 feathered setae; B) labial palp segment II with broad, thumb-like distomedial protuberance; C) fore femur rather broad, length ca. 3 × maximum width, dorsal margin with 12–15 curved, spine-like setae, fore tarsus broad with dorsal and ventral margins slightly convex; D) claw with eleven or twelve denticles; E) paraproct distally not expanded, with 23–32 stout marginal spines and some additional submarginal spines.
Larva (Figs
Colouration. Head, thorax, and abdomen dorsally light brown, with darker pattern as in Fig.
Antenna
(Fig.
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 and superlinguae
(Fig.
Maxilla
(Fig.
Labium
(Fig.
Hind protoptera
(Fig.
Foreleg
(Fig.
Terga
(Fig.
Gills
(Fig.
Paraproct
(Fig.
Dedicated to Dr. Emma Aguada Valdez, Mr. Rolando Valdez and Mr. Francis Paolo Valdez for their generous help and support in the scientific projects of the AdMU Biodiversity Laboratory.
Philippines: Negros and Cebu (Fig.
The specimens were collected at altitudes from 150 m to 480 m, on bottom gravel or submerged wood in runs or riffles.
Holotype. Philippines • larva; Negros Oriental, Valencia, Casaroro River downstream; 09°18'N, 123°14'E; 150 m; 01.IX.2019; leg. Garces and Pelingen; on slide; GenBank. MT830963; GBIFCH 00654888;
Larva. Following combination of characters: A) dorsal surface of labrum with submarginal arc of one plus 7–10 long, feathered setae; B) labial palp segment II with a broad, thumb-like distomedial protuberance, segment III conical; C) fore femur rather broad, length ca. 3 × maximum width, dorsal margin with a row of ca. 18 curved, spine-like setae; D) claw with 10–13 denticles; E) hind protoptera medium developed; F) paraproct distally not expanded, with ca. 30 stout marginal spines.
Larva (Figs
Colouration. Head dorsally light brown, thorax and abdomen dorsally mainly dark brown with pattern as in Fig.
Antenna
(Fig.
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 and superlinguae
(Fig.
Maxilla
(Fig.
Labium
(Fig.
Hind protoptera
(Fig.
Foreleg
(Fig.
Terga
(Fig.
Gills
(Fig.
Paraproct
(Fig.
Dedicated to Prof. Karl Matthias Wantzen (France), collector of some material, for his outstanding contribution to freshwater ecological research and conservation.
Philippines: Camiguin and Mindanao (Fig.
The specimens were collected at altitudes from sea level to 900 m, on bottom gravel, submerged wood, leaf packs or grass bunches in runs or riffles.
Holotype. Philippines • larva; Camiguin, Sagay, Bonbon, lower Binangawan River; 09°06'39"N 124°43'45"E; 30 m; 09.XII.2018; leg. Freitag and Wantzen; on slide; GenBank: MT830968; GBIFCH 00654898;
Following combination of characters: A) dorsal surface of labrum with submarginal arc of clavate, apically smooth setae; B) labial palp segment II with large, lobed or thumb-like distomedial protuberance, outer margin of protuberance predominantly concave (sometimes with hook-like modification of the protuberance); C) left mandible without setae at apex of mola, with minute denticles between prostheca and mola; D) six pairs of gills; E) hind protoptera absent; F) distolateral process at scape poorly developed or absent; G) patellotibial suture short, on basal 1/3 area of tibia; H) colour of larvae dorsally uniform brown.
Larva. Following combination of characters: A) dorsal surface of labrum with submarginal arc of ca. 15 clavate setae; B) labial palp segment II with a large, lobed distomedial protuberance, segment III slightly pentagonal, apically slightly truncate; C) left mandible without setae at apex of mola; D) fore femur rather slender, length 3.6 × maximum width, dorsal margin with a row of ca. ten curved, spine-like setae; E) tarsal claw with ca. eleven denticles; F) paraproct distally not expanded, with > 40 stout marginal spines.
Paratype. Philippines • 1 larva; rapids, Molawin Creek, college, Laguna; 28.VII.1977; leg. C.R. Realon; Coll. Pescador; on slide;
Philippines • 1 larva; Luzon, Laguna, Los Banos, UP Campus, Molawin River; 14°10'05"N, 121°11'44"E; 29.IX.1998; leg. Mendoza; on slide; GBIFCH 00654910; AdMU • 1 larva; Luzon, Laguna, Samil River; 14°08'N, 121°31'E; 27.VI.2018; 370 m; leg. Freitag et al.; on slide; GBIFCH 00654912;
Larva. Following combination of characters: A) dorsal surface of labrum with submarginal arc of ca. 26 clavate setae; B) labial palp segment II with a hook-like distomedial protuberance, segment III slightly pentagonal, apically slightly pointed; C) left mandible without setae at apex of mola; D) fore femur rather broad, length 3.4 × maximum width, dorsal margin with ca. 15 curved, spine-like setae; E) tarsal claw with ca. ten denticles; F) paraproct slightly expanded, with 35–39 stout marginal spines, some with split tips.
Holotype. Philippines • 1 larva; Mountain Prov., Sumigar Stream, Sumigar, Banaue; 03.X.1967; leg. Pescador; on slide;
Larva. Following combination of characters: A) dorsal surface of labrum with submarginal arc of 17–21 long, clavate setae; B) labial palp segment II with a thumb-like distomedial protuberance, segment III slightly pentagonal; C) left mandible without setae at apex of mola; D) fore femur rather slender, length ca. 4 × maximum width, dorsal margin with 8–11 curved, spine-like setae; E) paraproct distally expanded, with 39–43 stout, marginal spines.
Larva (Figs
Colouration. Head, thorax, and abdomen dorsally brown, thorax with pattern as in Fig.
Antenna
(Fig.
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 and superlinguae
(Fig.
Maxilla
(Fig.
Labium
(Fig.
Hind protoptera absent.
Foreleg
(Fig.
Terga
(Fig.
Gills
(Fig.
Paraproct
(Fig.
Dedicated to Mr. Bagani Sularte (Philippines), outstanding illustrator and friend of one of the authors (JG), for support in her dissertation work.
Philippines: Mindanao and Camiguin (Fig.
The specimens were collected at altitudes from sea level to 660 m, mainly in submerged wood or leaf packs in the runs.
Holotype. Philippines • larva; Mindanao, Agusan N, Cabadbaran River; 09°10'15"N, 125°40'55"E; 240 m.; 03.VI.2018; leg. Freitag and Pangantihon; on slide; GenBank: MT830974; GBIFCH 00654895;
Larva. Following combinations of characters: A) dorsal surface of labrum with submarginal arc of 16–19 long, clavate setae; B) labial palp segment II with a hook-like distomedial protuberance, segment III slightly pentagonal; C) left mandible without setae at apex of mola; D) fore femur rather slender, length ca. 4 × maximum width, dorsal margin with ca. eleven curved, spine-like setae; E) paraproct distally expanded, with 36–39 stout, marginal spines.
Larva (Figs
Colouration. Head, thorax, and abdomen dorsally light brown, fore protoptera light brown with bright striation. Head, thorax, and abdomen ventrally light brown. Legs light brown, femur with a distomedial brown spot and a brown streak at dorsal margin, apex brown, tibia darker in distomedial area. Caudalii light brown, with a brown band at ca. 1/2 of cerci length.
Antenna
(Fig.
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 and superlinguae
(Fig.
Maxilla
(Fig.
Labium
(Fig.
Hind protoptera absent.
Foreleg
(Fig.
Terga
(Fig.
Gills
(Fig.
Paraproct
(Fig.
Dedicated to Mr. Emmanuel Delocado (Philippines), entomologist and member of the AdMU Biodiversity Laboratory.
Philippines: Cebu and Leyte (Fig.
The specimens were collected at altitudes of 50 m and 700 m, partly on rock surface in riffles or runs.
Holotype. Philippines • larva; Cebu, Cantipla; 10°24'55"N, 123°49'05"E; 750 m; 30.X.1995; leg. Mendoza; on slide; GenBank: MT830976; GBIFCH 00654886;
Larva. Following combination of characters: A) dorsal surface of labrum with submarginal arc of ca. 17 long, clavate setae; B) labial palp segment II with a thumb-like distomedial protuberance, segment III sub-rectangular; C) left mandible without setae at apex of mola; D) fore femur rather slender, length 3.6 × maximum width, dorsal margin with ca. ten curved, spine-like setae; E) paraproct distally expanded, with 39–46 stout, marginal spines.
Larva (Figs
Colouration. Head, thorax, and abdomen dorsally light brown, fore protoptera light brown with bright striation. Head, thorax, and abdomen ventrally light brown, genae brown. Legs light brown, femur with a distomedial brown spot, dorsal and ventral margin and apex brown. Caudalii light brown.
Antenna
(Fig.
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 and superlinguae
(Fig.
Maxilla
(Fig.
Labium
(Fig.
Hind protoptera absent.
Foreleg
(Fig.
Terga
(Fig.
Gills
(Fig.
Paraproct
(Fig.
Dedicated to Prof. Hendrik Freitag (Philippines/Germany), collector of some material, for his outstanding contribution to freshwater biodiversity research in Southeast Asia.
Philippines: Palawan (Fig.
The specimens were collected at altitudes from sea level to 150 m, in leaf packs or submerged wood in runs or riffles.
Holotype. Philippines • larva; Palawan, Brooke’s Point, Mainit 7 Falls; 08°51'48"N, 117°47'45"E; 150 m; 2019; leg. Pelingen and Pangantihon; on slide; GBIFCH 00592322;
Larva. Following combination of characters: A) dorsal surface of labrum with submarginal arc of 15–17 long, clavate setae; B) labial palp segment II with a thumb-like distomedial protuberance, segment III slightly pentagonal; C) left mandible without setae at apex of mola; D) fore femur rather broad, length ca. 3 × maximum width, dorsal margin with 11–14 curved, spine-like setae; E) paraproct distally expanded, with ca. 35 stout, marginal spines.
Larva (Figs
Colouration. Head, thorax, and abdomen dorsally light brown, fore protoptera light brown. Head, thorax, and abdomen ventrally light brown. Legs light brown, femur with distomedial brown spot and brown apex. Caudalii light brown.
Antenna
(Fig.
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 and superlinguae
(Fig.
Maxilla
(Fig.
Labium
(Fig.
Hind protoptera absent.
Foreleg
(Fig.
Terga
(Fig.
Gills
(Fig.
Paraproct
(Fig.
Dedicated to Mr. Arthien Lovell Pelingen (Philippines), entomologist and former project assistant of the AdMU Biodiversity Laboratory.
Philippines: Negros and Cebu (Fig.
The specimens were collected at altitudes of 50 m and 480 m, on submerged wood or in riverside pools.
Holotype. Philippines • larva; Negros Oriental, Valencia, Malaunay, small tributary; 09°18'17"N, 123°10'07"E; 480 m; 01.IX.2019; leg. Garces and Pelingen; on slide; GBIFCH 00592315;
Following combination of characters: A) dorsal surface of labrum with submarginal arc of lanceolate setae; B) labial palp segment II with rather small, slender, thumb-like protuberance, distally bent upwards; C) hypopharynx with medial tuft of stout setae consisting of setae with minute apical serration; D) six pairs of gills; E) hind protoptera absent; F) no distolateral process at scape; G) paracercus short, ca. ¼ of cerci length.
Larva. Following combination of characters: A) dorsal surface of labrum with submarginal arc of three long, lanceolate setae; B) labial palp segment II with a hook-like distomedial protuberance, segment III conical; C) hypopharynx with well-developed medial tuft of long, stout setae, setae apically with minute serration; D) fore femur rather broad, length 2.6 × maximum width, dorsal margin with ca. 12 long, curved, spine-like setae and a second row of spine-like setae near margin; E) paraproct distally expanded with ca. six stout, marginal spines; F) paracercus short, ca. ¼ of cerci length.
Larva (Figs
Colouration. Head and pronotum dorsally brown, meso- and metanotum light brown, fore protoptera light brown with brown apex. Abdominal segments II–VI dorsally dark brown, segments I and VII–X light brown, segments VII and X with brown markings as in Fig.
Antenna
(Fig.
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 and superlinguae
(Fig.
Maxilla
(Fig.
Labium
(Fig.
Hind protoptera absent.
Foreleg
(Fig.
Terga
(Fig.
Gills
(Fig.
Paraproct
(Fig.
Dedicated to the late Mrs. Gisela Horzel (Germany), mother of one of the authors (TK).
Philippines: Luzon (Fig.
The specimens were collected at an altitude of 240 m on bottom gravel in runs or riffles.
Holotype. Philippines • larva; Luzon, Nueva Ecija, Pantabangan, Candaclan River; 15°46'48"N, 121°13'17"E; 240 m; 05.II.1998, leg. Mendoza; on slide; GBIFCH 00592280;
Larva. Following combination of characters: A) dorsal surface of labrum with submarginal arc of one plus 5–8 long, lanceolate setae; B) labial palp segment II with a slender, thumb-like protuberance, segment III slightly pentagonal with small apical projection; C) hypopharynx with well-developed medial tuft of long, stout setae, setae apically with minute serration; D) fore femur length 3.4 × maximum width, dorsal margin with 8–11 curved, spine-like setae and at least a partial row of spine-like setae near margin; E) paraproct distally expanded, with ca. nine stout, marginal spines; F) paracercus short, ca. 1/4 of cerci length.
Larva (Figs
Colouration. Head, thorax, and abdomen dorsally brown with bright pattern as in Fig.
Antenna
(Fig.
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 and superlinguae
(Fig.
Maxilla
(Fig.
Labium
(Fig.
Hind protoptera absent.
Foreleg
(Fig.
Terga
(Fig.
Gills
(Fig.
Paraproct
(Fig.
Dedicated to the late Mr. Joseph Mendoza (Philippines), outstanding insect collector.
Philippines: Mindanao (Fig.
The specimens were collected at altitudes from sea level to 120 m, on bottom gravel or submerged wood, or in leaf litter in side pools.
Holotype. Philippines • larva; Mindanao, Surigao del Sur, Tandag, Pangi, Pangi River; 09°06'18"N, 126°08'53"E; 10 m; 30.XI.2018, leg. Pangantihon; on slide; GBIFCH 00592272;
1 | Dorsal surface of labrum with submarginal arc of simple setae (Fig. |
(numeratus group) 2 |
– | Dorsal surface of labrum with submarginal arc of other types of setae | 7 |
2 | Setae at apex of mola of left mandible absent (Fig. |
L. lachicae sp. nov. |
– | Setae at apex of mola of left mandible present | 3 |
3 |
Hypopharynx with medial tuft of stout setae poorly developed (Fig. |
4 |
– |
Hypopharynx with medial tuft of stout setae well developed, short (Fig. |
5 |
4 | Posterior margin of tergum IV with triangular spines (Fig. |
L. sabordoi sp. nov. |
– | Posterior margin of tergum IV with rounded spines (Fig. |
L. palawano sp. nov. |
5 | Left mandible with comb-shaped structure at base of mola (Fig. |
6 |
– | Left mandible without comb-shaped structure at base of mola (Fig. |
L. camiguinensis sp. nov. |
6 | Paraproct with 18–21 marginal spines | L. aldabae sp. nov. |
– | Paraproct with 29–34 marginal spines plus a few submarginal spines | L. acei sp. nov. |
7 | Dorsal surface of labrum with submarginal arc of clavate setae (Fig. |
(sumigarensis group) 8 |
– | Dorsal surface of labrum with submarginal arc of other types of setae | 13 |
8 | Labial palp segment II enlargement pronounced hook-like with inner margin straight, segment III oblong and apically slightly truncate (Fig. |
L. sumigarensis |
– | Labial palp different | 9 |
9 | Labial palp segment II elongate thumb-like with distal margin strongly concave and inner margin concave (Figs |
10 |
– | Labial palp segment II broad thumb-like with distal margin not or only slightly concave (Fig. |
11 |
10 | Labial palp segment III apically slightly pointed (Fig. |
L. delocadoi sp. nov. |
– | Labial palp segment III apically rounded (Fig. |
L. freitagi sp. nov. |
11 | Paraproct distally not expanded (Fig. |
L. molawinensis |
– | Paraproct distally expanded (Fig. |
12 |
12 | Maxillary palp with distolateral excavation (Fig. |
L. baganii sp. nov. |
– | Maxillary palp with slight distolateral excavation (Fig. |
L. pelingeni sp. nov. |
13 | Dorsal surface of labrum with submarginal arc of lanceolate setae (Fig. |
(vallus group) 14 |
– | Dorsal surface of labrum with submarginal arc of other types of setae | 15 |
14 | Dorsal surface of labrum with submarginal arc of ca. three lanceolate setae; row of fine setae at innermost denticle of kinetodontium of right mandible absent (Fig. |
L. giselae sp. nov. |
– | Dorsal surface of labrum with submarginal arc of one plus ca. eight lanceolate setae; row of fine setae at innermost denticle of kinetodontium of right mandible present (Fig. |
L. mendozai sp. nov. |
15 | Dorsal surface of labrum with submarginal arc of feathered setae (Fig. |
(operosus group) 16 |
– | Dorsal surface of labrum with submarginal arc of dendritic setae (Fig. |
(dendrisetis group) L. dalisay sp. nov. |
16 | Dorsal surface of labrum with submarginal arc of setae with reduced or strongly reduced feathers (Figs |
17 |
– | Dorsal surface of labrum with submarginal arc of feathered setae, feathers not reduced (Fig. |
18 |
17 | Feathered setae of submarginal arc on dorsal surface of labrum strongly reduced, with few lateral branches only (Fig. |
L. gamay sp. nov. |
– | Feathered setae of submarginal arc on dorsal surface of labrum moderately reduced (Fig. |
L. tagbanwa sp. nov. |
18 | Labial palp segment II enlargement narrow thumb-like (Fig. |
L. pangantihoni sp. nov. |
– | Labial palp segment II enlargement broad thumb-like (Fig. |
19 |
19 | Fore tarsus dorsoventrally concave (broadened) (Fig. |
L. valdezorum sp. nov. |
– | Fore tarsus dorsoventrally parallel sided (Fig. |
L. wantzeni sp. nov. |
The material treated in this study was collected in many different locations across the Philippine archipelago, including most of the big islands as well as some smaller islands (Figs
Species | Species group | Locality | GPS coordinates |
---|---|---|---|
L. dalisay sp. nov. | dendrisetis gr. | Philippines: Luzon | 15°45'48''N, 121°25'21''E |
15°46'48''N, 121°13'17''E | |||
16°21'33"N, 120°30'31"E | |||
L. acei sp. nov. | numeratus gr. | Philippines: Luzon | 17°03'53"N, 121°05'10"E |
16°59'37"N, 121°02'51"E | |||
L. aldabae sp. nov. | numeratus gr. | Philippines: Luzon | 14°08'N, 121°31'E |
15°46'48''N, 121°13'17''E | |||
14°32'47''N, 121°13'42''E | |||
16°57'17''N, 120°38'52' E | |||
16°34'11"N, 120°50'12"E | |||
16°21'33"N, 120°30'31"E | |||
16°40'58"N, 120°56'59"E | |||
Philippines: Negros | 09°18'17''N, 123°10'07''E | ||
L. camiguinensis sp. nov. | numeratus gr. | Philippines: Camiguin | 09°06'39''N, 124°43'45''E |
09°09'25''N, 124°43'57''E | |||
L. lachicae sp. nov. | numeratus gr. | Philippines: Mindanao | 08°28'N, 125°59'E |
09°03'33''N, 126°05'57''E | |||
09°06'18''N, 126°08'53''E | |||
L. palawano sp. nov. | numeratus gr. | Philippines: Busuanga | 12°03'46''N, 120°13'25''E |
12°01'45''N, 120°12'19''E | |||
Philippines: Palawan | 10°09'47''N, 118°50'37''E | ||
10°01'26''N, 119°04'37''E | |||
L. sabordoi sp. nov. | numeratus gr. | Philippines: Negros | 09°18'N, 123°14'E |
Philippines: Romblon | 12°33'38''N, 122°07'19''E | ||
12°20'40''N, 122°40'37''E | |||
L. gamay sp. nov. | operosus gr. | Philippines: Mindoro | 12°37'06''N, 121°23'49''E |
12°37'18"N, 121°22'58"E | |||
Philippines: Luzon | 16°54'38"N, 120°28'40"E | ||
16°59'32"N, 120°32'21"E | |||
16°39'27"N, 120°25'55"E | |||
L. pangantihoni sp. nov. | operosus gr. | Philippines: Palawan | 07°57'39''N, 117°02'59''E |
07°57'39''N, 117°02'59''E | |||
09°21'07''N, 118°08'26''E | |||
09°18'25''N, 118°07'42''E | |||
L. tagbanwa sp. nov. | operosus gr. | Philippines: Palawan | 09°41'20''N, 118°37'29''E |
09°26'55''N, 118°26'44''E | |||
09°22'33''N, 118°08'41''E | |||
L. valdezorum sp. nov. | operosus gr. | Philippines: Negros | 09°18'N, 123°14'E |
09°17'N, 123°13'E | |||
Philippines: Cebu | 10°28'13"N, 123°52'26"E | ||
L. wantzeni sp. nov. | operosus gr. | Philippines: Camiguin | 09°06'39''N, 124°43'45''E |
09°12'N, 124°41'E | |||
09°09'25''N, 124°43'57''E | |||
Philippines: Mindanao | 08°09'42''N, 124°42'28''E | ||
L. molawinensis | sumigarensis gr. | Philippines: Luzon | 14°09'53''N, 121°14'48''E |
14°10'05''N, 121°11'44''E | |||
14°08'N, 121°31'E | |||
15°45'21''N, 121°34'46''E | |||
L. sumigarensis | sumigarensis gr. | Philippines: Luzon | 16°59'37''N, 121°02'51''E |
L. baganii sp. nov. | sumigarensis gr. | Philippines: Mindanao | 09°10'15''N, 125°40'55''E |
09°20'40''N, 125°30'50''E | |||
08°28'N, 125°59'E | |||
09°11'34''N, 125°36'34''E | |||
Philippines: Camiguin | 09°06'39''N, 124°43'45''E | ||
L. delocadoi sp. nov. | sumigarensis gr. | Philippines: Cebu | 10°24'55''N, 123°49'05''E |
Philippines: Leyte | 10°01'07''N, 125°12'35'E | ||
L. freitagi sp. nov. | sumigarensis gr. | Philippines: Palawan | 08°51'48''N, 117°47'45''E |
10°09'47''N, 118°50'37''E | |||
09°21'07''N, 118°08'26''E | |||
07°57'01''N, 117°04'29''E | |||
L. pelingeni sp. nov. | sumigarensis gr. | Philippines: Negros | 09°18'17''N, 123°10'07''E |
Philippines: Cebu | 10°45'32"N, 123°59'49"E | ||
L. giselae sp. nov. | vallus gr. | Philippines: Luzon | 15°46'48''N, 121°13'17''E |
L. mendozai sp. nov. | vallus gr. | Philippines: Mindanao | 09°08'18''N, 126°08'45''E |
08°28'N, 125°59'E | |||
09°06'18''N, 126°08'53''E |
COI sequences were obtained from all 18 new species (Table
Intraspecific (bold) and interspecific genetic distances of the new species (COI; Kimura 2-parameter; %; mean; minimum-maximum).
1 | 2 | 3 | 4 | 5 | 6 | 7 | 8 | 9 | 10 | 11 | 12 | 13 | 14 | 15 | 16 | 17 | 18 | ||
---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|
1 | L. dalisay sp. nov. | 0 | |||||||||||||||||
2 | L. acei sp. nov. | 18 | 0 | ||||||||||||||||
3 | L. aldabae sp. nov. | 20 | 16 | 3 | |||||||||||||||
20–22 | 15–16 | 0–6 | |||||||||||||||||
4 | L. camiguinensis sp. nov. | 23 | 19 | 16 | 0 | ||||||||||||||
5 | L. lachicae sp. nov. | 22 | 18 | 18 | 6 | 0 | |||||||||||||
6 | L. palawano sp. nov. | 22 | 17 | 18 | 15 | 16 | 2 | ||||||||||||
21–22 | 16–17 | 17–18 | 14–15 | 15–16 | |||||||||||||||
7 | L. sabordoi sp. nov. | 22 | 17 | 19 | 20 | 18 | 16 | 1 | |||||||||||
21–22 | 18–19 | 17–18 | 15–16 | ||||||||||||||||
8 | L. gamay sp. nov. | 25 | 21 | 20 | 20 | 21 | 20 | 22 | 0 | ||||||||||
24–25 | 21–22 | 19–21 | 19–20 | 22–23 | 0–1 | ||||||||||||||
9 | L. pangantihoni sp. nov. | 22 | 15 | 17 | 17 | 18 | 17 | 17 | 20 | 0 | |||||||||
16–17 | 16–18 | 20–21 | |||||||||||||||||
10 | L. tagbanwa sp. nov. | 22 | 20 | 17 | 19 | 19 | 20 | 19 | 13 | 20 | 1 | ||||||||
17–18 | 19–20 | 18–19 | 0–1 | ||||||||||||||||
11 | L. valdezorum sp. nov. | 22 | 22 | 20 | 21 | 22 | 22 | 23 | 18 | 23 | 17 | 0 | |||||||
19–21 | 21–22 | 22–23 | 18–19 | 17–18 | |||||||||||||||
12 | L. wantzeni sp. nov. | 24 | 19 | 18 | 21 | 23 | 20 | 21 | 21 | 19 | 21 | 20 | 1 | ||||||
17–19 | 19–21 | 20–21 | 0–2 | ||||||||||||||||
13 | L. baganii sp. nov. | 22 | 18 | 18 | 20 | 21 | 18 | 19 | 21 | 20 | 19 | 24 | 21 | 2 | |||||
17–19 | 19–20 | 20–21 | 20–21 | 19–20 | 18–20 | ||||||||||||||
14 | L. delocadoi sp. nov. | 24 | 24 | 23 | 23 | 23 | 21 | 23 | 23 | 22 | 23 | 24 | 23 | 24 | 2 | ||||
23–24 | 23–24 | 21–25 | 22–23 | 22–23 | 20–21 | 22–24 | 22–24 | 21–22 | 22–23 | 23–24 | |||||||||
15 | L. freitagi sp. nov. | 26 | 25 | 22 | 23 | 23 | 23 | 23 | 26 | 23 | 26 | 27 | 26 | 22 | 23 | 0 | |||
21–22 | 22–23 | 23–24 | 23–24 | 22–23 | 26–27 | 22–23 | 25–26 | 26–27 | 22–24 | 0–1 | |||||||||
16 | L. pelingeni sp. nov. | 24 | 19 | 19 | 22 | 19 | 20 | 22 | 24 | 21 | 22 | 23 | 24 | 21 | 23 | 23 | 3 | ||
23–24 | 18–19 | 18–21 | 21–22 | 19–20 | 21–22 | 23–24 | 21–22 | 23–24 | 22–24 | 20–21 | 22–23 | 22–24 | |||||||
17 | L. giselae sp. nov. | 21 | 18 | 18 | 21 | 20 | 23 | 20 | 22 | 19 | 20 | 20 | 17 | 21 | 21 | 23 | 22 | 0 | |
17–19 | 22–23 | 19–20 | 20–21 | 17–18 | 20–21 | 20–21 | 22–23 | 21–22 | |||||||||||
18 | L. mendozai sp. nov. | 23 | 21 | 19 | 20 | 20 | 19 | 22 | 22 | 23 | 19 | 20 | 24 | 22 | 24 | 27 | 21 | 21 | 0 |
18–19 | 18–19 | 20–21 | 23–25 | 23–24 | 26–27 |
For the assignment of the new species to Labiobaetis we refer to
From the 16 species of Labiobaetis (or previously assigned to Pseudocloeon) only known at the imaginal stage, one was described from the Philippines (Mindanao; Pseudocloeon boettgeri Ulmer, 1924). As the identification of the imaginal stage of Labiobaetis is generally very difficult, we consider it unrealistic to safely associate the larval stage with old type material at the imaginal stage. In this case, rearing material will provide little help. Furthermore, the generic assignment of the species remains questionable. Therefore, we did not take this species into account in our study and wait for an eventual clarification of its status in the future by using ancient DNA methods.
The 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 genus. The inclusion of nuclear gene sequences may prove that some are natural groups. Most of the species in the Philippines belong to three different groups only, six to the numeratus group, five to the operosus group, and six to the sumigarensis group. These groups are widespread and highly diversified in Asia. Species of the numeratus group are also known from Sri Lanka, Malaysia, Indonesia and China, species of the operosus group from Malaysia and Indonesia and species of the sumigarensis group from India, Sri Lanka, Malaysia, Indonesia, Brunei, China and Taiwan (
Interestingly, a few of these newly described species share characters with the fauna of New Guinea. Labiobaetis dalisay sp. nov. shares the dendritic setae on the dorsal surface of the labrum with L. dendrisetis Kaltenbach & Gattoliat from New Guinea; it also has a similar labial palp and maxillary palp and seven pairs of gills (
Labiobaetis giselae sp. nov. and L. mendozai sp. nov. share the lanceolate setae on the dorsal surface of the labrum with L. vallus Kaltenbach & Gattolliat from New Guinea, have a similar labial palp and also no scape process, no hind protoptera and six pairs of gills (
In general, the genetic distances between the different species of Labiobaetis in the Philippines are rather high, between 15% and 27% (K2P, Table
In addition to the species described in this paper, we obtained nine COI sequences with clearly interspecific genetic distance to other specimens with the same morphology. To give a more complete view on the diversity, including this cryptic diversity, we are presenting them as Molecular Operational Taxonomic Units (MOTUs) based on genetic evidence only (COI; Table
Molecular Operational Taxonomic Units (MOTUs) of Labiobaetis in the Philippines.
MOTU denomination | Species group | Locality | Specimens catalog # | GenBank # | GenSeq |
---|---|---|---|---|---|
(COI) | Nomenclature | ||||
L. cf. lachicae sp. nov. I | numeratus group | Philippines: Mindanao | GBIFCH 00654893 | MT830932 | genseq-4 COI |
L. cf. aldabae sp. nov. I | numeratus group | Philippines: Luzon | GBIFCH 00654905 | MT830934 | genseq-4 COI |
L. cf. wantzeni sp. nov. I | operosus group | Philippines: Mindanao | GBIFCH 00654876 | MT830931 | genseq-4 COI |
L. cf. wantzeni sp. nov. II | operosus group | Philippines: Mindanao | GBIFCH 00763665 | MT830937 | genseq-4 COI |
L. cf. wantzeni sp. nov. III | operosus group | Philippines: Mindanao | GBIFCH 00763666 | MT830938 | genseq-4 COI |
L. cf. baganii sp. nov. I | sumigarensis group | Philippines: Camiguin | GBIFCH 00654887 | MT830933 | genseq-4 COI |
L. cf. delocadoi sp. nov. I | sumigarensis group | Philippines: Negros | GBIFCH 00654890 | MT830935 | genseq-4 COI |
L. cf. pelingeni sp. nov. I | sumigarensis group | Philippines: Luzon | GBIFCH 00763654 | MT830936 | genseq-4 COI |
L. cf. molawinensis I | sumigarensis group | Philippines: Mindanao | GBIFCH 00763673 | MT830939 | genseq-4 COI |
As mentioned above, of the 21 species of Labiobaetis known from the Philippines, six belong to the numeratus group, five to the operosus group and six to the sumigarensis group. All these groups are widespread and highly diversified in Southeast Asia. Members of sumigarensis and operosus group are also known from Borneo (
Based on the present data, all the Philippine Labiobaetis species are endemic to the Philippines. Moreover, most species (14) are restricted to one island. Although partly due to localities missing in this study, island endemics are quite common among aquatic insect species of the Philippines (
In a few cases, we have more than one species of the same species group occuring on the same island: L. acei sp. nov. and L. aldabae sp. nov. from numeratus group on Luzon; L. sumigarensis and L. molawinensis from sumigarensis group on Luzon. Moreover, we have additional diversity as MOTUs partly sharing the same species groups and islands (Table
We also have cases of species, which are distributed on more than one island: L. baganii sp. nov. and L. wantzeni sp. nov. both on Mindanao and Camiguin and with a genetic distance of 2% between the islands; L. sabordoi sp. nov. on Negros and Romblon (K2P 1%); L. aldabae sp. nov. on Luzon and Negros (K2P 2%–6%); L. gamay sp. nov. on Luzon and Mindoro (K2P 0%–1%); L. pelingeni sp. nov. on Negros and Cebu (K2P 3%); L. delocadoi sp. nov. on Cebu and Leyte (K2P 2%). Based on the small genetic distances and the mostly close geographic distances, these cases suggest either current or remnant intra-archipelagic dispersal crossing sea channels. Interestingly, most of these island pairs do not belong to the same intra-Philippine biogeographic region (Greater Luzon, Greater Mindoro, Romblon, West Visayas, Camiguin, and Greater Mindanao) (
Four species in Palawan offer an interesting zoogeographic affinity based on morphology. The morphology of L. freitagi sp. nov. (sumigarensis group) is closer to L. delocadoi sp. nov. than to the species from Borneo (
The number of sampled localities and different habitats is still limited and there are regions without any collection activities so far (Figs
This study was made possible with the Gratuitous Permits (GP 0133-17 and renewals) for the collection of aquatic wildlife as kindly issued by the Bureau of Fisheries and Aquatic Resources (BFAR), Quezon City and the Palawan Council for Sustainable Development, Puerto Princesa City (No. 2019-20). Prerequisite permissions were given by the respective local government units.
We sincerely thank Hendrik Freitag, Emmanuel Delocado, Clister V. Pangantihon, Arthien Lovell Pelingen, Marc Ryan Sabordo, and Kyra Mari Dominique Aldaba (AdMU, Philippines) for the assistance in the field and collection management. Additional precious materials were contributed by Ace Kevin Amarga (Philippines/Taiwan), Joseph Mendoza (Philippines), Wolfram Mey (Germany), and Karl Matthias Wantzen (France). We are deeply thankful to Lars Hendrich (Zoologische Staatssammlung München,
Lastly, the authors are grateful to Cassandra Caedo (USA) for the corrections and improvements of the English language and to the reviewers for the recommendations and comments on the manuscript.
Expeditions and management of aquatic macroinvertebrate material in the AdMU Biodiversity Laboratory were partly supported by the German Academic Exchange Service (DAAD project BIO-PHIL 57393541) and School of Science and Engineering Industry 4.0 Research Fund (SI4-013).