ZOOTAXAISSN 1175-5326 (print edition)

ISSN 1175-5334 (online edition)Copyright 2013 Magnolia Press

Zootaxa 3693 (1): 049063 www.mapress.com/zootaxa/ Article

http://dx.doi.org/10.11646/zootaxa.3693.1.3http://zoobank.org/urn:lsid:zoobank.org:pub:9C7822BB-05A7-4844-8964-C7E556967527

Systematics of cryptic species of Lebinthus crickets in Mount Makiling (Grylloidea, Eneopterinae)

TONY ROBILLARD1, SHERYL YAP2 & MARK V. YNGENTE31Musum national dHistoire naturelle, Dpartement Systmatique et Evolution, UMR 7205 CNRS-OSEB, CP 50 (Entomologie), 75231 Paris Cedex 05, France2Crop Protection Cluster, College of Agriculture and UPLB Museum of Natural History, University of the Philippines Los Baos, Laguna, Philippines 3Institute of Biology, College of Science, University of the Philippines Diliman, Quezon City, Philippines

Abstract

In the present study, we address Lebinthus cricket species from Luzon (Philippines) being sympatric in Mount Makiling. We describe Lebinthus puyos new species and redescribe the species Lebinthus sanchezi Bolvar, 1889 and select a neo-type series. The species Lebinthus makilingus Otte, 2007 is synonymized under Lebinthus sanchezi. For each species we provide complete descriptions of morphology, including male and female genitalia and forewing venation and we describe calling song for L. sanchezi.

Key words: Lebinthus, new species, redescription, neotype, Philippines, sympatry

Introduction

The crickets of the subfamily Eneopterinae (sensu Robillard & Desutter-Grandcolas 2008) show original traits relative to acoustic communication (e.g., Robillard & Desutter-Grandcolas 2004a, b; Robillard & Desutter-Grancolas 2011; Robillard et al. 2007, 2013). As observed in several cricket groups, they tend to speciate quickly, especially in islands (Shaw 1999; Otte 2007a; Oneal et al. 2010). As demonstrated by Nattier et al. (2012) in the eneopterine genus Agnotecous, diversification often takes place through allopatric speciation. However, allopatric speciation can be followed by range expansion in available habitats, which may cause secondary contacts between closely related species, resulting in sympatric distribution between endemic species.

We document here a case of multiple sympatries in Mount Makiling (Luzon, Philippines). Three eneopterine genera are present in the area: the apterous long-legged Paranisitra, represented by the species P. longipes Chopard, 1925 (review in Gorochov 2009), the long-winged Cardiodactylus, represented by C. kondoi Otte, 2007, a species widespread in the Philippines (Robillard et al. in prep.), and the brachypterous genus Lebinthus. Three species of this genus are currently known in Luzon island: L. bitaeniatus Bolvar, 1889, redescribed in Robillard & Tan 2013), Lebinthus sanchezi Bolvar, 1889 and Lebinthus makilingus Otte, 2007, all being described from the same geographical area, Mount Makiling and its surroundings.

Field work in Mount Makiling and in the University of the Philippines Laguna Land Grant, east of Mt. Makiling, followed by morphological and taxonomical studies, revealed that there are indeed three species of Lebinthus in this area corresponding to the yellow striped L. bitaeniatus and to two brown species. However, careful examination of original descriptions, type localities, newly collected material and specimens from natural history collections show that L. makilingus should be synonymized under L. sanchezi while a new species occurs in the lower part of Mt. Makiling. L. sanchezis type has not been found in Madrid, where Bolvars collection is located (Mercedes Paris pers. com.; Paris 1993), but the original description and type locality allow relating L. sanchezi to our series of collected specimens found both in Mt. Makiling and in UP Land Grant. We redescribe L. sanchezi and define a neotype series for this species, deposited in Los Banos, Madrid and Paris. Accepted by D. Rentz: 1 Jul. 2013; published: 26 Jul. 2013 49

The other brown species is a rather rare species, new to sciences, found in Mount Makiling (only a few specimens collected in 2011 and none found in 2012 at the same location) and living in sympatry with L. sancheziand L. bitaeniatus. This new species, Lebinthus puyos n. sp., differs from L. sanchezi by pattern of coloration including a whitish face and absence of orange band at apex of fastigium, and by male genitalia, which structures are closer to Lebinthus villemantae Robillard, 2010 from Sulawesi and Lebinthus truncatipennis Chopard, 1929 from Sumatra (Robillard 2010).

Material and methods

Material. Field collections and observations were made in several localities in Mount Makiling and UP Land Grant, Pakil, Laguna (Philippines, Luzon) in July 2011 and June 2012 (TR & MY). Specimens were collected by sight only, by night and day, in order to observe their habitat and period of activity. Newly collected specimens are deposited in the collections of the Musum national dHistoire naturelle, Paris (MNHN), in the Museum of Natural History of Los Baos (UPLB-MNH) or in Museo Nacional de Ciencias Naturales, Madrid (MNCN). Square brackets in lists of specimens are used for additional information not mentioned on specimen labels.

Observations and morphological analysis. Direct observations and dissections have been made using a binocular microscope Leica MZ16 at magnifications up to 160, equipped with a camera lucida for the line drawings. SEM observations were performed at the Plateforme de Microscopie lectronique of the MNHN, using a JEOL-JSM 840 electronic microscope (7kV), after a 60 s gold-coating. Male tegminal veins and cells follow terminology by Desutter-Grandcolas (2003) and Robillard & Desutter-Grandcolas (2004a). Male and female genitalia have been dissected in softened specimens, by cutting the membranes between the paraprocts and the subgenital plate, or between the ovipositor and the subgenital plate respectively, and then kept in glycerine in vials pinned under specimens. Photographs of male genitalia have been made with a binocular microscope Leica MZ12 and the montage software Leica Application Suite ver. 2.8.1 (Leica Microsystems). Male genitalia are named according to Desutter (1987), modified in Desutter-Grandcolas (2003) and Robillard & Desutter-Grandcolas (2004a). Dotted parts in figures correspond to membranous areas. Abbreviations: see below.

Acoustic data. The basic cricket song terminology follows Ragge & Reynolds (1998). One song unit is called a syllable and corresponds to one opening-closure cycle of the male forewings.

L. sanchezi has been recorded in the field and/or in the laboratory from specimens collected in the field as juveniles or sub-adults. The recordings were made with a modified Condenser Microphone Capsule CM16 (Avisoft Bioacoustics, Berlin), with a relatively flat frequency response from 3 to 150 kHz (R. Specht pers. comm.), connected to a Fostex FR-2LE digital field recorder (96 kHz sampling frequency, 16 bit) in the field, or using or using Avisoft Triggering Harddisk Recorder version 2.97 and a 8-Pre MOTU sound card at a sampling frequency of 96 k-samples s-1 (16 bit). Acoustic analyses were performed using the computer software Avisoft-SASLab Pro version 4.40 (Specht 2009) by measuring the song features with automatic commands. All recording files are deposited in the Sound Library of the Musum national dHistoire naturelle, Paris.

Abbreviations

InstitutionsMNHN Musum national d'Histoire naturelle, Paris, France.UPLB-MNH Museum of Natural History, University of the Philippines Los Baos. MNCN Museo Nacional de Ciencias Naturales, Madrid, Spain.

General morphology:I, II, III: front, median, hind respectively (femora, legs, tibiae);F: femora;FW: forewing;Tarsomere III-1: basal segment of hind leg tarsomere;T: tibiae.ROBILLARD ET AL.50 Zootaxa 3693 (1) 2013 Magnolia Press

Male genitalia:ec arc: ectophallic arc;ecap: ectophallic apodeme;enap: endophallic apodeme;en s: endophallic sclerite;ps p: pseudepiphallic paramere;r: rami.

Tegminal venation:1A4A: first to fourth anal veins;CuA: anterior cubitus; CuA1, CuA2, ...: first, second, ... bifurcations of CuA;CuP: posterior cubitus;MA, MP: anterior, posterior media veins;R: radial vein;d1 cell (mirror): first cell(s) of D alignment;

Measurements:FIIIL: length of hind femora;FIIIW: width of hind femora;FWL: forewing length;FWW: forewing width (at the level of maximal width);Ias: inner spines on TIII dorsal side, above the spurs;Ibs: inner spines on TIII dorsal side, between the spurs;Oas: outer spines on TIII dorsal side, above the spurs;Obs: outer spines on TIII dorsal side, between the spurs;OL: ovipositor length;PronL: pronotum length;PronW: pronotum width;TIIIL: length of hind tibiae;TaIIIs: spines on outer edge of third hind tarsomere, not including the apical spine.

Taxonomy

Subfamily Eneopterinae Saussure, 1874

Tribe Lebinthini Robillard, 2004

Genus Lebinthus Stl, 1877

Synonym Paraeneopterus Saussure, 1878Type species: Lebinthus bitaeniatus Stl, 1877

Diagnosis. Among Lebinthini genera, Lebinthus is closely related to Agnotecous Saussure, 1878, to which it resembles by microptery and FW venation. It is characterized by its rather smaller size, microptery in both sexes (FW short and hind wings absent), and male FW venation with mirror almost not differentiated from apical field, dorsal field as long or longer than lateral field (it is shorter in Agnotecous), median fold short, triangular and located on dorsum. Zootaxa 3693 (1) 2013 Magnolia Press 51LEBINTHUS CRICKETS FROM MOUNT MAKILING

Lebinthus sanchezi Bolvar, 1889(Figs 1AD; 2AC; 3A,B,E; 4AC; 5AD; 6)

Lebinthus sanchezi Bolvar, 1889: 425; Chopard 1968: 354; Otte 1994: 67; Eades et al. 2013.Synonym namesLebinthus makilingus Otte, 2007 new synonymy. Otte 2007a: 356, not validly published; 2007b: 33. This study: similar to L. sanchezi according to the description and photographs of male genitalia.

Systematic discussion.The types of Bolvar (1889) supposedly located in Madrid (Otte 1994; Eades et al. 2013) could not be located in the collection (Paris 1993; M. Paris, pers. comm.) and are considered lost. Due to the necessity for a clear taxonomic reference for the species of the diverse genus Lebinthus upon which to base future systematic work, we designate a neotype from the material examined in the present study.

The original description of L. sanchezi is very brief (Lebinthus without the yellow lateral stripes as in L. bitaeniatus) and may match both L. makilingus and L. puyos n. sp., consequently we choose the neotype specimens according to geographical proximity with the original type locality. The type locality of Bolvars types being Jala Jala (Distrito of Morong), a place very close to our sampled locality Pakil, University of the Phillipines Laguna Land Grant, we consider the only Lebinthus specimens found in this area as the neotype series of L. sanchezi. L. sanchezi has in fact a wider distribution, as it is found both in Land Grant and in Mount Makiling, while L. puyos n. sp. was only found in Mount Makiling.

According to photographs of male genitalia in Ottes (2007a) description of L. makilingus, this species is the same as the Lebinthus species found both in Land Grant (original type locality of L. sanchezi) and in Mount Makiling. L. makilingus should thus be synonymised under L. sanchezi.

Type material. Male neotype: Philippines. Luzon, Paete [Pakil], University of the Philippines Laguna Land Grant, secondary forest at 500 m from station, 1423'56.9"N 12132'47.2"E, 376 m (GPS LG2), 4.vii.2011, day, leaf litter, coll. T. Robillard (UPLB-MNH). Paraneotypes (6 males, 3 females). Philippines. Luzon, Paete [Pakil], University of the Philippines Laguna Land Grant station, secondary forest, reared specimens F1 generation (F0 collected on vii.2011, day, leaf litter), coll. T. Robillard: 2 males, 1 female (UPLB-MNH); 3 males, 1 female (MNHN); 1 male, 1 female (MNCN).

Other material examined. Philippines: Luzon, Laguna, Los Baos, Mount Makiling, base, secondary forest on campus, 1409'12.9"N 12114'05.0"E, 168 m (GPS Maki1), 1 female, TR282b,coll. T. Robillard (MNHN). Philippines: Luzon, Laguna, Los Baos, Mount Makiling, Flat Rock, W of Mulawin Creek, secondary forest, 1408'50.2"N 12113'41.5"E, 244 m (GPS Maki2), 28.vi.2011, coll. T. Robillard, night, leaf litter: 1 female, TR180 (UPLB-MNH); 4 males, TR311, TR179 (UPLB-MNH); TR283, TR178, on plant (h = 10 cm) (MNHN); 3 females, TR645 (UPLB-MNH); TR344, TR345 (MNHN).

Type locality. Philippines, Luzon, Jala Jala, Morong District. Neotype locality: Philippines, Luzon, Pakil, University of the Philippines Laguna Land Grant.

Distribution. Philippines, Luzon Island, west of Laguna Bay.Diagnosis. Species of small size, brown color, differing from sympatric L. puyos n. sp. by black mask on face,

fastigium with an orange band apically, vertex with faint longitudinal bands, characteristic color pattern on lateral lobes of pronotum, harp of males with a rectangular false mirror, diagonal vein absent, male genitalia with little differentiated triangular pseudephiphallic lophi. L. sanchezi appears as an intermediate form between the yellow banded species of Lebinthus (e.g. L. bitaeniatus and L. luae) in terms of genitalic structure, and the brown species with a false mirror (e.g. L. truncatipennis, L. villemantae and L. puyos) in terms of external morphology and coloration.

Redescription. Size small. Coloration brown with contrasted light and dark patterns (Fig. 1AD). Head dorsum (Fig.2A) with 4 wide but faint brown longitudinal band, each band containing a thin black line; area posterior to eyes yellow. Eyes dark brown, with a paler dorsal longitudinal band. Fastigium wider than long, setose, dark brown, with a contrasted vivid orange band apically; upper facial part yellow. Scapes whitish with faint darker patterns; antennae yellow at base then progressively dark brown. Face (Fig. 2B) with a black mask extended bellow antennae and eyes. Mouthparts variably mottled with dark brown and yellow brown; maxillary palpi yellow brown with dark brown longitudinal patterns, darker apically. Lateral part of head (Fig.2C) yellow, with a black band posterior to eyes prolonging the dark coloration of pronotum. Pronotum: Dorsal disk slightly trapezoidal, straight posteriorly; yellow brown mottled with dark brown, lateralROBILLARD ET AL.52 Zootaxa 3693 (1) 2013 Magnolia Press

FIGURE 1. .Lebinthus sanchezi (AD), male in dorsal (A) and lateral (B) views; female in dorsal (C) and lateral (D) views. Lebinthus puyos (EH), male in dorsal (E) and lateral (F) views; female in dorsal (G) and lateral (H) views. Photographs of Lebinthus bitaeniatus male (I) and female (J) are represented for comparison purpose (modified from Robillard & Tan 2013). Scale bar = 1 cm. Zootaxa 3693 (1) 2013 Magnolia Press 53LEBINTHUS CRICKETS FROM MOUNT MAKILING

FIGURE 2. Lebinthus sanchezi (AC), head in dorsal (A), facial (B) and lateral (C) views. Lebinthus puyos (DF), head in dorsal (D), facial (E) and lateral (F) views. Scale bar = 1 mm.

edges always whitish, posterior area mostly dark brown. Lateral lobes black dorsally, ventral margin yellow with a median black patch sometimes linked to the anterior margin and/or to the dorsal dark coloration (Fig. 2C). Legs: Fore and mid legs light brown, femora with dark brown spots, tibiae with dark rings. Hind femora brown, with strong striated dark patterns on outer faces, knees dark brown with a pale inner preapical area; hind tibiae with dark rings. Tarsomeres I/III-1 yellow brown, apices dark brown. Tarsomeres III-1 with 2 spines on dorsal outer edges (n = 10) and 02 (m = 1, n=10) on outer faces. Abdomen homogeneously brown dorsally, yellowish brown ventrally. Cerci yellowish brown, with dark rings near apex.

Male: FWs not reaching abdomen midlength. FW coloration (Fig. 3A): Cells dark brown, not translucent; veins dark brown except anterior parts of anal veins orange brown; MA and MP orange brown, intermedian area whitish, without transverse veins; R dark brown, area between MA and R dark brown, rest of lateral field progressively lighter toward ventral margin, with dark brown longitudinal veins. FW venation (Fig. 3A): 1A widely curved (angle>120); stridulatory file with 240 teeth (n = 1) (Fig. 3E), located on transverse part of 1A only. CuP not visible. Diagonal vein absent or very faint, preplaced by a convex fold underlying harp posterior margin. Harp wide, occupying most of dorsal field surface, with a strong transverse harp vein, polyfurcated anteriorly and delimiting a false mirror barely rounded, i.e. a distinctive area located on harp posterior corner (not homologous to the mirror of other cricket species: cell d1, Robillard & Desutter-Grandcolas 2004a). CuA curved innerly near apex, its distal part weak, surrounding the median fold, small, triangular, and located dorsally. Longitudinal veins of dorsal field strong apically, transverse veins very weak. Mirror (d1) not differentiated. Apical field absent, with no bifurcation of CuA posterior to diagonal vein. Lateral field with 5 strong longitudinal veins including MA, R and 3 more ventral veins; latero-dorsal angle made by MP; R without bifurcating veins. Subgenital plate elongate, clog-shaped.

Male genitalia (Fig. 4AC): Intermediate between L. bitaeniatus and L. villemantae. Pseudepiphallic sclerite trapezoidal, slightly convex dorsally; posterior apex with short triangular lophi, setose, wider than long and ROBILLARD ET AL.54 Zootaxa 3693 (1) 2013 Magnolia Press

separated by a V-shaped indentation; anterior margin slightly indented, its lateral margins slightly raised dorsally. Rami short. Pseudepiphallic parameres F-shaped. Ectophallic arc complete almost straight, its base with short ventro-posterior expansions, sclerotized basally. Ectophallic fold with a wide X-shaped preapical sclerite. Ectophallic apodemes very long, thin and divergent, exceeding anterior margin of pseudepiphallus. Endophallic sclerite wide and long, exceeding anterior margin of pseudepiphallus, its posterior apex with a median triangular expansion and short lateral arms; endophallic apodeme made of wide lateral lamellas but without a median crest.

Female: FWs short (Fig. 3B), rounded, going slightly beyond posterior margin of second tergite, not overlapping but close together; dorsal field light brown with 6 strong orange brown longitudinal veins. Lateral field dark brown, with 2 strong dark brown longitudinal veins. Ovipositor shorter than hind femora; apex lanceolate, slightly denticulate on dorsal edge.

FIGURE 3. Forewing venation in dorsal view: Lebinthus sanchezi, male (A), female (B); Lebinthus puyos, male (C), female (D); scale bar = 1 mm. SEM photograph of male stridulatory file of L. sanchezi in ventral view (E); scale bar = 100 m. Zootaxa 3693 (1) 2013 Magnolia Press 55LEBINTHUS CRICKETS FROM MOUNT MAKILING

FIGURE 4. Lebinthus sanchezi (AC), male genitalia in dorsal (A), ventral (B) and lateral (C) views. Lebinthus puyos (DF), male genitalia in dorsal (D), ventral (E) and lateral (F) views. Scale bars = 1 mm.

Female genitalia: flat, little sclerotized, with a basal sclerotized ring, apex rounded. Juvenile. Dark brown mottled with light brown (Fig. 5B).Measurements. See Table 1.Habitat and life history traits. L. sanchezi lives in the leaf litter of secondary forested areas (Fig. 5AD). It is

found on top of leaves in small bushes and sometimes in the leaf litter. Males have been observed singing on top of leaf litter or low plants during day and night.

Behavior. Calling song (Fig. 6): In the field the calling song of L. sanchezi consists of short trills comprising 5813 syllables. Each echeme lasts for 0.990.19 s with a period of 28.368.89 s. Syllables are very short (duration =6.41.9 ms; period = 17.06.3 ms) and show a homogeneous amplitude profile suggesting a regular wing closing movement during wing stridulation. The spectrum shows a clear dominant peak at 24.41.2 kHz with one peak harmonically related at about 48 kHz. Mating behavior: Observations in the field and in the laboratory showed multiple matings, as described in another Lebinthus species (Narvaez & Robillard 2012).ROBILLARD ET AL.56 Zootaxa 3693 (1) 2013 Magnolia Press

TABLE 1. Measurements of Lebinthus sanchezi.

TABLE 1. (Continued)

Lebinthus puyos Robillard, n. sp.(Figs. 1EH; 2DF; 3CD; 4DF; 5E)

Type material. Holotype male: Philippines. Luzon, Laguna, Los Baos, Mount Makiling, base, secondary forest on campus, 1409'12.9"N 12114'05.0"E, 168 m (GPS Maki1), 27.vi.2011, night, leaf litter, TR652, coll. T. Robillard (UPLB-MNH). Allotype female: same information as HT, TR653 (UPLB-MNH). Paratypes (5 males): Philippines. Luzon, Laguna, Los Baos, Mount Makiling, coll. T. Robillard: 4 males, same information as HT, TR644 (UPLB-MNH); TR651 (MNHN-ENSIF); TR280, on plant (h = 20 cm), TR393, mort en levage (MNHN-ENSIF). Luzon, Laguna, Los Baos, Mount Makiling, Flat Rock, W of Mulawin Creek, adult in captivity, 1408'50.2"N 12113'41.5"E, 244 m (GPS Maki2), 28.vi.2011, night, leaf litter, 1 male, TR281 (MNHN).

Other material examined. Philippines. Same information as HT, 1 juvenile, TR281b, coll. T. Robillard (MNHN).

Type locality. Philippines, Luzon, Laguna, Los Baos, Mount Makiling,Distribution. Philippines, Luzon Island, base of Mount Makiling.Etymology. Named after its cryptic and jumpy attitude, based on a Latinized version of the French word

pou, which refers to small and jumpy animals.Diagnosis. Species of small size, brown color, differing from sympatric L. sanchezi by whitish face, fastigium

not orange apically, and vertex without clear longitudinal bands, characteristic color pattern on lateral lobes of pronotum, harp of males with a well-rounded false mirror, diagonal vein present anteriorly. According to structure of male genitalia with well-differentiated pseudepiphallic lophi, L. puyos seems closer to L. villemantae than to L. sanchezi, but differs from L. villamentae by more contrasted coloration and details of male genitalia.

Description. Size small. Coloration contrasted, light brown with darker patterns (Fig. 1EH). Head dorsum (Fig. 2D) almost homogeneously dark brown posteriorly, lighter toward fastigium, without distinct longitudinal bands; area posterior to eyes yellow. Eyes light brown dorsally, dark brown ventrally. Fastigium wider than long, setose, light brown, without a contrasted yellow band apically; upper facial part whitish, black laterally, sometimes with 2 faint median brown spots. Scapes whitish with faint darker patterns; antennae yellow at base then progressively dark brown. Face (Fig. 2E) mostly whitish, with dark brown patches below antennae and a thin vertical black line bellow eyes. Mouthparts mostly light brown; maxillary palpi yellow brown, darker apically. Lateral part of head (Fig. 2F) whitish with two thin black lines. Pronotum: Dorsal disk slightly trapezoidal, straight posteriorly; yellow brown to light brown, with black spots. Lateral lobes black dorsally, ventral margin with two yellow areas, an anterior one and a ventral one, separated by a horizontal dark line fused to the dorsal dark area (Fig. 2F). Legs: Legs I and II light brown, femora with dark brown spots, tibiae with dark rings. Hind femora brown, darker dorsally than ventrally, with faint striated dark patterns on outer faces, knees dark brown; hind tibiae with dark rings. Tarsomeres I/III-1 yellow brown, apices dark brown. Tarsomeres III-1 with 23 spines on dorsal

PronL PronW FWL FWW FIIIL FIIIW

Male neotype 1.9 3.2 2.7 2.3 8.7 3.3

Males (n=5) 1.92.1 3.23.3 2.63.4 22.5 8.79.5 3.23.3

(mean) (2.0) (3.2) (2.9) (2.3) (9.2) (3.3)

Females (n=5) 22.1 3.23.4 0.61.1 - 9.710 3.33.7

TIIIL TIII OL

Ias Ibs Oas Obs

Male neotype 6.9 4 3 7 3 -

Males (n=5) 6.97.7 46 35 710 35 -

(mean) (7.3) (5) (4) (8) (4) -

Females (n=5)(mean)

7.38.2(7.8)

45(5)

35(4)

510(8)

45(4)

7.37.4(7.3) Zootaxa 3693 (1) 2013 Magnolia Press 57LEBINTHUS CRICKETS FROM MOUNT MAKILING

outer edges (m = 2, n = 6) and 01 (m = 0.3, n= 6) on outer faces. Abdomen homogeneously brown dorsally, yellowish brown ventrally. Cerci yellowish brown, with faint dark rings near apex.

FIGURE 5. Lebinthus sanchezi (AD): female on leaf litter during day in Mount Makiling (Luzon, Philippines) (A), juveniles on plant in Land Grant during day (B); male courting at night on low plant (C), view of habitat in Mount Makiling (near Flat Rock). L. puyos (E): view of secondary habitat on campus in Mount Makiling, near UPLB MNH. Photographs by TR, except (B) courtesy of J. Baroga (UPLB).ROBILLARD ET AL.58 Zootaxa 3693 (1) 2013 Magnolia Press

FIGURE 6. Calling song of Lebinthus sanchezi. Oscillogram of 3 echemes (A); oscillogram (B) and sonogram (C) of 1 echeme; oscillogram (D) and linear power spectrum (E) of 1 syllable.

Male. FWs not reaching abdomen midlength. FW coloration: Cells brown, not translucent; veins dark brown or orange brown; intermedian area whitish, sometimes with dark brown veins; lateral field dark brown dorsally, Zootaxa 3693 (1) 2013 Magnolia Press 59LEBINTHUS CRICKETS FROM MOUNT MAKILING

progressively lighter toward ventral margin; lateral field with strong orange brown longitudinal veins and faint black transverse veins. FW venation (Fig. 3C): 1A curved, without a clear angle; stridulatory file with ca. 250 teeth (n = 1) located on transverse part of 1A only. CuP not visible. Diagonal vein clearly visible on anterior half, faint posteriorly. Harp wide, occupying most of dorsal field surface, with a strong semi-circular harp vein, polyfurcated anteriorly and delimiting a rounded false mirror with the curved area of CuA, i.e. a distinctive rounded area located on harp posterior corner (not homologous to the mirror (cell d1) of other cricket species). CuA curved innerly near apex, it distal part weak posterior to curved area, surrounding the median fold, small and located on abdomen dorsum. Posterior part of diagonal vein not visible. Longitudinal veins strong apically, transverse veins very weak. Mirror (d1) not differentiated. Apical field absent, with no bifurcation of CuA posterior to diagonal vein. Lateral field with 56 strong longitudinal veins including MA, R and 34 more ventral veins; latero-dorsal angle made by MP; R without bifurcating veins. Subgenital plate elongate, clog-shaped.

Male genitalia (Fig. 4DF): Close to male genitalia of L. villemantae. Pseudepiphallic sclerite elongate, convex dorsally; more or less rectangular, narrowed posteriorly near base of short individualized lophi, setose, parallel and separated by a V-shaped indentation; anterior margin slightly indented, its lateral margins slightly raised dorsally. Rami short. Pseudepiphallic parameres C-shaped, trilobate, the posterior lobe dorsal, the two other lobes ventral. Ectophallic arc complete, deeply curved posteriorly, its base with long ventro-posterior expansions, sclerotized basally. Ectophallic fold with a wide preapical sclerite including a rounded part and two antero-lateral expansions. Ectophallic apodemes parallel and very long, exceeding anterior margin of pseudepiphallus. Endophallic sclerite very long, exceeding anterior margin of pseudepiphallus, its posterior apex with a median triangular expansion and with short lateral arms; endophallic apodeme made of lateral lamellas but without a median crest.

Female. FWs short (Fig. 3D), not reaching posterior margin of second tergite, not overlapping but close together; dorsal field light brown with 5 strong brown longitudinal veins. Lateral field light brown, with 4 strong brown longitudinal veins, intermedian area without a distinctive coloration. Ovipositor slightly shorter than hind femora; apex lanceolate, almost not denticulate on dorsal edge.

Female genitalia: not observed. Juvenile. Dark brown mottled with whitish; vertex whitish with faint longitudinal bands, posterior area black.Measurements. see Table 2. Habitat and life history traits. L. puyos is found in low densities in the leaf litter of secondary forested areas

(Fig. 5E). It is found on top of leaves in small bushes and sometimes in the leaf litter. Behavior. Unknown.

TABLE 2. Measurements of Lebinthus puyos.

TABLE 2. (Continued)

PronL PronW FWL FWW FIIIL FIIIW

Male holotype 2.1 3.3 3 2.4 10.4 3.3

Males (n=5) 22.1 33.3 33.3 2.22.4 9.610.4 3.23.5

(mean) (2.1) (3.2) (3.1) (2.3) (10.1) (3.3)

Females (n=1) 1.9 3.1 1.2 - 8.7 3.1

TIIIL TIII OL

Ias Ibs Oas Obs

Male holotype 9 9 5 11 6 -

Males (n=5) 7.99 79 5 1011 6 -

(mean) (8.6) (8) (5) (11) (6) -

Females (n=1) 7.4 6 5 8 6 7.4ROBILLARD ET AL.60 Zootaxa 3693 (1) 2013 Magnolia Press

Key to the Lebinthus species in Luzon

1. Body with lateral yellow or whitish bands running dorsally from head to apex of abdomen, male FW without a false mirror in the harp. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Lebinthus bitaeniatus Bolvar, 1889

- Body mostly brown without lateral band on abdomen, male FW with a more or less rounded false mirror in the posterior corner of the harp . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2.

2. Face with a black mask, apex of fastigium with a contrasted orange or yellow band, male genitalia with short and wide triangu-lar lophi, pseudepiphallic parameres F-shaped . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Lebinthus sanchezi Bolvar, 1889

- Face mostly whitish, apex of fastigium without contrasted orange and, male genitalia with longer parallel lophi, pseudepiphal-lic parameres C-shaped . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .Lebinthus puyos Robillard, n. sp.

Discussion

Documented cases of cricket diversification in islands involve processes of allopatric speciation through geographic isolation (Oneal et al. 2010; Nattier et al. 2012). However clear examples of sympatry amongst closely related species suggest possible cases of sympatric speciation, as speciation can also take place even in absence of geographic barrier, usually by exploitation of new niches involving divergence in factors such as food resources, microhabitats, different communication channels or other life history traits (e.g. Via 2001).

In the present case of multiple sympatries, could the three Lebinthus species found in Mount Makiling originate from sympatric speciation or do their features suggest multiple colonization events? Independent origins of L. bitaeniatus and L. sanchezi + L. puyos is obvious, as these species clearly belong to different species groups within the genus (Fig. 1), according to external morphology (male and female venation, absence of false mirror in L. bitaeniatus), coloration (yellow band along the whole body in L bitaeniatus), male genitalia and acoustic features (hearable calling song in L. bitaeniatus made of two distinct parts vs. ultrasounds only in L. sanchezi, and likely in L. puyos).

The case of L. sanchezi and L. puyos is a better candidate for sympatric speciation, as both species belong to the same group of small brown species showing a false mirror in the posterior part of the harp (Fig. 3). This feature is however present in other Lebinthus species such as L. truncatipennis and L. villemantae, respectively from Sumatra and South Sulawesi (Robillard 2010), and in L. yaeyamensis Oshiro, 1996 from Ryu Kyu islands and Taiwan. Also, close examination of male genitalia, and especially the shape of the pseudepiphallus and pseudepiphallic lophi reveal that L. puyos more resembles L. villemantae from Sulawesi than L. sanchezi. Under the hypothesis of sympatric speciation, this suggests either that this genitalic pattern would be a plesiomorphy shared by L. villemantae and L. puyos, and that L. sanchezi would have developed a novel pattern. The alternate hypothesis is that L. sanchezi and L. puyos originated independently and that observed sympatry in Mount Makiling is a recent secondary contact, as demonstrated in other eneopterine crickets from New Caledonia for example (Nattier et al. 2012).

The hypothesis that sympatries between Lebinthus species in Mount Makiling originate from multiple secondary contacts after allopatric speciation is supported by morphology, but will have to be tested by reconstructing the phylogeograhy of Lebinthus in the Philippines. This will however necessitate more sampling efforts in this region, as the cricket faunas from major islands are yet unknown or insufficiently studied to address such questions with adequate methods.

Acknowledgements

The field work in Luzon (20112012) was organized by TR (MNHN) and SY (UPLB), funded by grants from the ATM Biodiversit actuelle et fossile, MNHN (Stphane Peign & Philippe Janvier) and ATM Formes possibles, formes ralises, MNHN (Vincent Bels & Pierre-Henri Gouyon). We thank Pablo V. Quilao (UPLB) and Jessica. Baroga (UPLB) for helping collecting and photographing crickets. We also thank Simon Poulain (CNRS) and Guy Lecorvec (MNHN) for their help in specimen preparation and pictures and Laure Desutter-Grandcolas for the SEM photograph of the file of L. sanchezi. Zootaxa 3693 (1) 2013 Magnolia Press 61LEBINTHUS CRICKETS FROM MOUNT MAKILING

References

Bolvar, I. (1889) Enumeracion de Grilidos de Filipinas. Anales de la Sociedad espanola de Historia natural, 18, 415431.Chopard, L. (1968) In: Pars 12. Fam Gryllidae: Subfam. Mogoplistinae, Myrmecophilinae, Scleropterinae, Cachoplistinae,

Pteroplistinae, Pentacentrinae, Phalangopsinae, Trigonidiinae, Eneopterinae. Fam. Oecanthidae, Gryllotalpidae. Dr. W. Junk N.V.'s Gravenhage, pp. 215500.

De Saussure, H. (1874) Mission scientifique au Mexique et dans l'Amrique centrale. 6me partie: tudes sur les Myriapodes et les Insectes. Imprimerie impriale, Paris, 531 pp.

Desutter, L. (1987) Structure et volution du complexe phallique des Gryllidea (Orthoptera) et classification des genres notropicaux de Grylloidea.1re partie. Annales de la Socit Entomologique de France (N.S.), 23, 213239.

Desutter-Grandcolas, L. (2003) Phylogeny and the evolution of acoustic communication in extant Ensifera (Insecta, Orthoptera). Zoologica Scripta, 32, 525561. http://dx.doi.org/10.1046/j.1463-6409.2003.00142.x

Eades, D.C., Otte, D., Cigliano M.M. & Braun, H. (2013) Orthoptera Species File. Version 5.0/5.0. Available from: http://Orthoptera.SpeciesFile.org (Accessed 26 May 2013)

Gorochov, A.V. (2009) A review of the genus Paranisitra Chopard (Orthoptera: Gryllidae: Eneopterinae). Proceedings of the Zoological Institute RAS, 313(4), 435439.

Narvaez, A. & Robillard, T. (2012) The reproductive behaviour of the cricket Lebinthus santoensis Robillard, 2009 (Grylloidea, Eneopterinae, Lebinthini). Zoosystema, 34 (2), 279286. http://dx.doi.org/10.5252/z2012n2a6

Nattier, R., Grandcolas, P., Elias, M., Desutter-Grandcolas, L., Jourdan, H., Couloux, A. & Robillard, T. (2012). Secondary Sympatry Caused by Range Expansion Informs on the Dynamics of Microendemism in a Biodiversity Hotspot. PLoS ONE, 7(11), e48047. http://dx.doi.org/10.1371/journal.pone.0048047

Oneal, E., Otte, D. & Knowles, L.L. (2010) Testing for biogeographic mechanisms promoting divergence in Caribbean crickets (genus Amphiacusta). Journal of Biogeography, 37, 530540. http://dx.doi.org/10.1111/j.1365-2699.2009.02231.x

Otte, D. (1994) Orthoptera species file. 1. Crickets (Grylloidea). Orthopterists's Society and the ANSP, Philadelphia, Pennsylvania, 120 pp.

Otte, D. (2007a). New species of Cardiodactylus from the western Pacific region (Gryllidae: Eneopterinae). Proceedings of the Academy of Natural Sciences of Philadelphia, 156, 341400. http://dx.doi.org/10.1635/0097-3157(2007)156[341:nsocft]2.0.co;2

Otte, D. (2007b) New cricket genera and species (Orthoptera: Grylloidea) from the Pacific Region deposited in the Bishop Museum, Honolulu. In: Evenhuis, N.L. & Bickel, D.J. (Eds.), Fiji Arthropods IX, Vol. 94, pp. 2134.

Paris, M. (1993) Catalogo de tipos de Orthopteroides (insecta) de Ignocio Bolivar, I: Blattaria, Mantodea, Phasmoptera Y Orthoptera (Stenopelmatoidea, Rhaphidophoroidea, Tettigonioidea, Grylloidea, Tetrigoidea). Eos, 69, 143264.

Ragge, D.R. & Reynolds, W.J. (1998) The songs of the grasshoppers and crickets of Western Europe. Colchester, England: Harley Books, 612 pp.

Robillard, T. (2010) New species of the genus Lebinthus (Orthoptera, Grylloidea, Eneopterinae, Lebinthini) from Indonesia and the Solomon Islands. Zootaxa, 2386, 2548.

Robillard, T. & Desutter-Grandcolas, L. (2004a) Phylogeny and the modalities of acoustic diversification in extant Eneopterinae (Insecta, Orthoptera, Grylloidea, Eneopteridae). Cladistics, 20(3), 271293. http://dx.doi.org/10.1111/j.1096-0031.2004.00025.x

Robillard, T. & Desutter-Grandcolas, L. (2004b) High-frequency calling in Eneopterinae crickets (Orthoptera, Grylloidea, Eneopteridae): an adaptive radiation revealed by phylogenetic analysis. Biological Journal of the Linnean Society, 83, 577584. http://dx.doi.org/10.1111/j.1095-8312.2004.00417.x

Robillard, T. & Desutter-Grandcolas, L. (2008) Clarification of the taxonomy of extant crickets of the subfamily Eneopterinae (Orthoptera: Grylloidea; Gryllidae). Zootaxa, 1789, 6668. http://dx.doi.org/10.1163/000579511x572044

Robillard, T. & Desutter-Grandcolas, L. (2011) Evolution of calling songs as multicomponent signals in crickets (Orthoptera: Grylloidea: Eneopterinae). Behaviour, 148(56), 627672. http://dx.doi.org/10.1163/000579511x572044

Robillard, T., Grandcolas, P. & Desutter-Grandcolas, L. (2007) A shift toward harmonics for high-frequency calling shown with phylogenetic study of frequency spectra in Eneopterinae crickets (Orthoptera, Grylloidea, Eneopteridae). Canadian Journal of Zoology, 85(12), 12641275.http://dx.doi.org/10.1139/z07-106

Robillard, T., Montealegre-Z, F., Desutter-Grandcolas, L., Grandcolas, P. & Robert, D. (2013) Mechanisms of high frequency song generation in brachypterous crickets and the role of ghost frequencies. Journal of Experimental Biology, 216(11), 20012011. http://dx.doi.org/10.1242/jeb.083964ROBILLARD ET AL.62 Zootaxa 3693 (1) 2013 Magnolia Press

http://dx.doi.org/10.1635/0097-3157(2007)156[341:nsocft]2.0.co;2http://dx.doi.org/10.1635/0097-3157(2007)156[341:nsocft]2.0.co;2

Robillard, T. & Tan, M.K. (2013) A taxonomic review of common but little known crickets from Singapore and the Philippines (Insecta: Orthoptera: Eneopterinae). Raffles Bulletin of Zoology, in press.

Shaw, K.C. (1999) Conflict between nuclear and mitochondrial DNA phylogenies of a recent species radiation: what mtDNA reveals and conceals about modes of speciation in Hawaiian crickets. Proceedings of the National Academy of Sciences, 99(25), 1612216127. http://dx.doi.org/10.1073/pnas.242585899

Specht, R. (2009) Avisoft-SASLab Pro - Sound Analysis and Synthesis Laboratory. Avisoft Bioacoustics, Berlin.Via, S. (2001) Sympatric speciation in animals: the ugly duckling grows up. Trends in Ecology and Evolution, 16(7), 381390.

http://dx.doi.org/10.1016/s0169-5347(01)02188-7 Zootaxa 3693 (1) 2013 Magnolia Press 63LEBINTHUS CRICKETS FROM MOUNT MAKILING

AbstractIntroductionMaterial and methodsTaxonomySubfamily Eneopterinae Saussure, 1874Tribe Lebinthini Robillard, 2004Genus Lebinthus Stl, 1877Lebinthus sanchezi Bolvar, 1889Lebinthus puyos Robillard, n. sp.Key to the Lebinthus species in LuzonDiscussionAcknowledgementsReferences

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