New genus and new species of spittlebugs (Hemiptera: Cercopidae) from the Philippines

The following new taxa are described from the Philippines: Mioscarta nubisa Crispolon & Soulier-Perkins sp. nov., M. translucida Crispolon & Yap sp. nov. and Trigonosch e ma Crispolon & Soulier-Perkins gen. nov. with three new species: T. manoborum Crispolon & Soulier-Perkins sp. nov. (as type species), T. negrosensis Crispolon & Yap sp. nov and T. rubercella Crispolon & Guilbert sp. nov. Trigonoschema pallida (Lallemand, 1927) comb. nov. is transferred from Mioscarta Breddin, 1901. Descriptions of male genitalia are illustrated and keys to species of Philippine Mioscarta and Trigonoschema gen. nov. are provided. Although phylogenetic results confi rm the monophyly of all genera and Trigonoschema being a distinct genus from Mioscarta , relationships between genera remain uncertain. A checklist of the genera and species of Cercopidae found in the Philippines is included.


Molecular study
Taxon and data sampling A phylogenetic analysis based on molecular characters is performed in order to check if its results support our decision to describe Trigonoschema gen. nov. as a new genus distinct from Mioscarta . This study includes 16 exemplars of Cercopidae as ingroup representing genera Eoscarta Breddin, 1902 (3), Mioscarta (3), Trigonosch e ma (3), Poeciloterpa (5), Jacobsoniella Melichar, 1914 (1) and Wawi Soulier-Perkins & Le Cesne, 2016 (1) and 14 terminals as outgroup selected from Aphrophoridae, Clastopteridae and Machaerotidae. Genera included in the ingroup are supposed to be phylogenetically as close as possible to Mioscarta and Trigonoschema gen. nov. in order to test our hypothesis . We have selected genera that belong to the same tribe, Rhinaulacini: Poeciloterpa , Eoscarta from the Philippines and Wawi from Papua New Guinea. Jacobsoniella is found in the Philippines but in the actual classifi cation it is incertae sedis concerning its tribe placement. However, the male terminalia for this genus present the same characteristic sterno-lateral plates observed in most Rhinaulacini genera. For this reason, we used it in our analysis, included in the ingroup.
Most specimens sampled were stored into 95-100% ethanol otherwise dry. DNA extractions were conducted using standard protocols for QIAmp DNA microkit (Qiagen) from disarticulated legs. Intact voucher specimens from which the disarticulated legs were extracted were mounted on pins and are deposited in the MNHN. PCR reactions, with negative controls included to detect contamination, were conducted in 25 μl volume using Taq DNA Polymerase, from Taq Core kit (Qiagen) under standard thermocycler protocols (for Histone 3, 18S and 28S: initial denaturation for 3 minutes at 94°C, then 35 cycles of 94°C for 30 seconds, 49-56°C for 40 seconds and 72°C for 1 minute, then a fi nal extension step at 72°C for 10 minutes, and fi nally held at 10°C before being removed from the cycler. For CO1, the thermal cycling protocol are the following: initial denaturation for 3 minutes at 94°C; fi ve cycles of 30 seconds at 94°C, 40 seconds at 47°C, and 1 minute at 72°C; 30 cycles of 30 seconds at 94°C, 40 seconds at 52°C, and 1 minute at 72°C; 10 minutes at 72°C; and fi nally held at 10°C) and using Oligonucleotide primers (Table 3) targeting four loci: CO1, Histone 3, 18S, and 28S.
Amplifi ed DNA was visualized using 1-2% agarose gel electrophoresis with midori green staining. Sequence fragments were imported into codoncode aligner V.5.1.4. (CodonCode Corporation, Dedham, Massachusetts, USA) and trimmed to remove primer sequence. After sequence inspection, contiguous sequences were assembled and edited based on chromatograms to ensure the accuracy of base calls. In addition, insertions/deletions and checking of contaminations were confi rmed for accuracy by using a reference sequence for comparison in BLAST ( https://blast.ncbi.nlm.nih.gov/Blast.cgi ). All sequence data are accessioned into GenBank (Table 4).
For the combined analyses all sequence alignments were concatenated into a single data set using Phylosuite v1.2.2 (Zhang et al. 2020). The resulting data matrix consisted of 3575 bp of DNA nucleotide sequence data for 30 Cercopiodea specimens used as terminals. Phylogenetic reconstructions were conducted using maximum likelihood (ML) criteria (Guindon et al. 2010) and Bayesian inference (BI). Under all reconstructions method, gaps were treated as missing data. Partitioned analyses were conducted with Partition Finder (Lanfear et al. 2012(Lanfear et al. , 2016. Best fi tting model was searched using partition fi nder with following confi guration: branchlengths = linked, models = all, model_selection AICc and the search = greedy. Codon code mode was activated for CO1 before running the analysis. Results obtained with the corrected Akaike Information Criterion (AICc) (Guindon et al. 2010;Lanfear et al. 2012Lanfear et al. , 2016 indicated that GTR+G was best fi tting model for Histone 3; TRNEF+I+G for 18S; GTR+I+G, TRN+I and HKY+I+G for each codon base for CO1 and GTR+I+G for 28S. For ML analysis the results were then imported to IQ tree (Guindon et al. 2010;Minh et al. 2013;Nguyen et al. 2015;Hoang et al. 2018) for fast and effective stochastic algorithm to reconstruct phylogenetic trees. Partition mode was selected,"Models" argument will be ignored and models and thread were automatically set to auto. Maximum likelihood phylogeny were inferred under edge-linked partition model for 1000 standard bootstraps, as well as the Shimodaira-Hasegawa-like approximate likelihoodratio test (SH_aLRT branch test) with 1000 replicates (Guindon et al. 2010;Minh et al. 2013;Nguyen et al. 2015;Hoang et al. 2018).
BI analysis was conducted with MrBayes version 3.1.2 (Huelsenbeck & Ronquist 2001;Ronquist & Huelsenbeck 2003) as implemented in CIPRES (Miller et al. 2010). Because MrBayes only allows a relatively small collection of models, models in the analyses were approximated using GTR+I+G (nst = 6) for DNA, or 'wag' for Protein (Guindon et al. 2010;Lanfear et al. 2012Lanfear et al. , 2016. The analysis coupled with Metropolis Coupled Markov Chain Monte Carlo (MCMCMC) was implemented. These analyses were conducted with six-gene partition implemented across two independent runs each with four chains run (one cold and three heated) for 20 million generations with 1000 sampling frequency. Burn in fraction value was set to 0.25 in which the initial 25% of sampled data were discarded as burn-in. The fi nal posterior probability tree was calculated as a 50% majority-rule consensus tree (Huelsenbeck & Imennov 2002;. Clade support was evaluated by their posterior probabilities (BPP). The IQ Tree analysis tree and the Bayesian inference analysis tree were visualized with FIGTREE v1.1.3 (Rambaut 2016 Class Insecta Linnaeus, 1758Order Hemiptera Linnaeus, 1758Suborder Auchenorrhyncha Duméril, 1806Infraorder Cicadomorpha Evans, 1946Family Cercopidae Leach, 1815Subfamily Cercopinae Oshanin, 1916Tribe Rhinaulacini Kirkaldy 1906Subtribe Poeciloterpina Schmidt, 1920Genus Mioscarta Breddin, 1901Mioscarta Breddin, 1901, 183 (Zoogeography).

Diagnosis
The genus can be identifi ed by the following combination of characters: Habitus general shape dorso-ventrally fl attened, in lateral view total length nearly 4 times height (Figs 2, 4-11A). Pronotum angle not more than 25° (Figs 2,. Distance between ocellus and compound eye 2 times ocellus diameter (Fig. 1A). Ocelli large, distance between eyes less than 8 times ocellus diameter (Fig. 1A). Apical reticulation of the tegmen generally developed and reduced in few cases (Figs 4-11A-C). Widest part of postclypeus in frontal view is at midheight (Fig. 1A). Apical curve of tegmen visible in dorsal view . Widest part of habitus in dorsal view at midlength of tegmen . Male subgenital plates is at least 1.5 times longer than pygofer height. Male subgenital plates appendage always present, longer than main plate (Figs 5,(7)(8).
Among the Rhinaulacini, Mioscarta closely resembles Trigonoschema Crispolon & Soulier-Perkins gen. nov . with respect to the distance between ocelli, postclypeus shape in frontal view, postclypeus longitudinal groove, apical cells of the tegmen, Rp posterior wing, absence of r-m crossvein, and presence of sterno-lateral plate between subgenital plate and pygofer while Peociloterpa with respect to postclypeus longitudinal groove, absence of r-m crossvein, presence of sterno-lateral plate between subgenital plate and pygofer and paramere general shape, but they differ by the following characters presented in Table 2 below.

Remarks
The description provided by Jacobi (1927) was based on four females. Here we provide photographs of a female syntype and its labels. Male remains unknown. (Walker, 1851)   6 Size of ocelli evaluated according to the distance between eyes using ocellus diameter large, distance between eyes less than 8 times ocellus diameter ( Fig. 1A) very small, distance between eyes 16 times ocellus diameter ( Fig. 1C) small, distance between eyes 9-10.5 times ocellus diameter ( 14 Subgenital plates length at least 1.5 times longer than pygofer height (Figs 5, 7-8, 10-11F).

Mioscarta ferruginea
slightly longer than pygofer height clearly shorter than 1.5 times longer than pygofer height (

Description
Male Terminalia In lateral view, posterior margin of pygofer slightly undulating (Fig. 5E). Subgenital plate (Fig. 5F) very long relative to height of pygofer with fi ne tapering appendage slightly longer than main plate, dorsal margin of plate largely rounded, sterno-lateral plate present, largely concave subapically on dorsal and ventral margins. Intermediate plate present, rectangular, linking internal side of lateral and subgenital plates. Paramere (Fig. 5G) not globose, dorsal margin rounded and regularly curving, apex with a sharp spine pointing dorsally, ventral margin regularly rounded with subapical part bearing a spiniform process pointing ventrally. Aedeagus ( Fig. 5H) with basal third of dorsal margin angled and sharply bent in acute angle and last 2 ⁄ 3 vertical and roughly C-shaped, slightly humped in apical part of dorsal margin before apical extension, apical extension pointing postero-dorsally, posterior protrusion thin and hook-shaped, postero-dorsal protrusion absent.

Remarks
Walker (1851) based his description on two male specimens "a" and "b" from the "Philippine Islands" coming from the collections of Mr Cuming and Mr Wood. Both specimens are currently kept in the NHM. According to Walker' description and Lallemand' key (1949), we identifi ed specimens, for which we provide photographs and male terminalia description and drawings. Schmidt, 1925 Fig. 6 Mioscarta lutea Schmidt, 1925: 36.

Distribution
Philippines: Luzon Island. Schmidt (1925) described this species based on a single female specimen. Therefore, the description does not contain any information on the male terminalia. Here, we provide photographs of the female holotype and labels. Male remains unknown.

Mioscarta nubisa
Crispolon & Soulier-Perkins sp. nov. urn:lsid:zoobank.org:act:17979C2C-D071-44EA-9D16-974A742808A1 Fig. 7 Diagnosis General shape of M. nubisa is similar to M . obscuripennis but are distinctly different in color. M. nubisa in dorsal view presents a brownish and opaque tegmen, darker toward apex, pronotum brownish and yellowish or brown legs while M . obscuripennis has a dark brown or black tegmen, brown or orange pronotum and orange legs.

Etymology
Species name refers to the light to darker coloration forming a cloudy pattern on the tegmen and is the female superlative of the latin word "nubis" which means cloudy. HEAD (Figs 1A, 7B). In dorsal view, large ocelli, distance between eyes less than 8 times ocellus diameter, distance between ocelli equals one ocellus diameter, distance between ocellus and compound eye 2 times ocellus diameter, ocelli closer to each other than from compound eyes. Eyes not prominent, length 1.44 times than wide. Vertex and frons longitudinal median carina absent. Vertex slightly longer than wide with 3 times ocellus diameter in between two vertex grooves outside ocelli and 3.5 times ocellus diameter between anterior and posterior vertex margins. Postclypeus with longitudinal furrow, slightly swollen and ovoid shape in frontal view, widest part at mid height (Figs 1A, 7B), not receding and prior to anteclypeus where it bends forming obtuse angle in lateral view (Fig. 7A). Rostrum long, reaching but not surpassing mesocoxae. Thorax (Figs 2A, 7A-C). In dorsal view, pronotum with anterior concavities on each side, anterior margin of pronotum as wide as posterior margin of head including eyes, anterolateral margin curved, posterior margin grooved, postero-lateral margin slightly concave, longer than anterolateral margin, humeral angle rounded. In lateral view, pronotum curving not more than 25º (Figs 2, 7A). Scutellum as long as wide with large median dimple (Fig. 7C). Tegmen ( Fig. 7A-C). R bifurcates on apical half, M bifurcate on basal third, apical reticulation well developed without concave apical cells. Posterior wing (Fig. 3A), Rp separating from SC+Ra nearly at midlength, M reaches ambient vein, Cua and Cup fused at base and m-cu links M to Cua before Cua bifurcation, common base for Cup and Cua originate at base of wing, posterior wing with 7 longitudinal veins and 5 apical cells between SC+Ra and Cup, angular protrusion of costal margin near its base present. Metafemur with apical spine in inner margin, metatibiae bearing 1 lateral spine. MALE TERMINALIA. In lateral view, posterior margin of pygofer largely convex in middle with slight curved on last third (Fig. 7E). Subgenital plates (Fig. 7F) very long relative to height of pygofer with fi ne tapering appendage longer than main plate, dorsal and ventral margin of main plate straight, sterno-lateral plate present, slightly elongated. Intermediate plate present, elongated, roughly oblong shaped, linking internal sides of lateral and subgenital plates. Paramere (Fig. 7G) not globose, dorsal margin convex and regularly curving fi nishing by a sharp process pointing dorsally, ventral margin convex and largely angled subapically, apex with spiniform process pointing postero-ventrally. Aedeagus (Fig. 7H) with basal third of dorsal margin regularly bent without angle before the bent part, last 2 ⁄ 3 vertical and S-shaped, ventral margin regularly curved, apical extension pointing posteriorly, posterior protrusion axe-shaped with edge prolonged ventrally by a straight, long and thin extension, postero-dorsal protrusion absent. COLOR. Head, antennae, pronotum, abdomen, legs scutellum light brown, rostrum brownish highlighted with white. Tegmen opaque light brown to brown.

Mioscarta obscuripennis
Mioscarta obscuripennis Schmidt, 1920: 47.      Description MALE TERMINALIA. In lateral view, posterior margin of pygofer hump shaped in middle, last third abruptly truncated (Fig. 8E). Subgenital plates (Fig. 8F) very long relative to height of pygofer with fi ne tapering appendage longer than main plate, dorsal margin regularly convex and slightly curving on basal part, ventral margin roughly straight. Sterno-lateral plate present and roughly bean-shaped. Intermediate plate present, square shaped with a corner truncated, linking internal sides of lateral plate and subgenital plate. Paramere (Fig. 8G) dorsal margin rounded then regularly curving up to apex. Lateral margin with spine located subapically pointing dorsally, ventral margin rounded then sharply angled subapically pointing ventrally, apex without any spines. Aedeagus (Fig. 8H) with basal third of dorsal margin regularly bent, basal half enlarged then largely narrowed in the middle and gradually enlarging up to apex, roughly C-shaped, ventral margin regularly curved, apical extension pointing posteriorly, posterior protrusion axe-shaped with long thin extension prolonging the edge ventrally and slightly curving, postero-dorsal protrusion absent.

Distribution
Philippines: Luzon Island, Baguio, Quezon province, Mountain province; Visayas, Negros Island. Schmidt (1920) described this species from a single male specimen with few details on the male terminalia. Here, we provide, a detailed illustration and description of male terminalia from identifi ed specimens using Schmidt description (Fig. 8), along with photographs of the male holotype and its labels (Fig. 9). The coloration of the specimen we identifi ed and the holotype are different but the specimen from which we illustrated the male terminalia (Fig. 8) has a brighter coloration and is the closest to the description given by Schmidt while the holotype has a faded coloration. Jacobi, 1905 Fig. 10 Mioscarta semperi Jacobi, 1905: 21.

Mioscarta semperi
Mioscarta semperi -Schmidt 1909: 239-240. -Lallemand 19121949: 84-85. -Jacobi 1927 549. Description MALE TERMINALIA. In lateral view, posterior margin of pygofer regularly convex last third slightly curved (Fig. 10E). Subgenital plates (Fig. 10F) very long relative to height of pygofer with fi ne tapering appendage slightly longer than main plate, dorsal margin of plate regularly rounded, ventral margin straight. Sterno-lateral plate present, slightly elongated. Intermediate plate present, cylindrical, linking internal side of lateral and subgenital plates. Paramere (Fig. 10G) not globose, dorsal margin rounded and regularly curving, apex with a sharp spine pointing antero-dorsally, ventral margin regularly rounded then subapical part largely rounded, protruding and fi nishing with a sharp spine pointing ventrally. Aedeagus ( Fig. 10H) with basal third of dorsal margin angled and sharply bent in acute angle and last 2 ⁄ 3 vertical and roughly C-shaped, slightly humped in apical part of dorsal margin before apical extension, apical extension pointing posteriorly, posterior protrusion thin very short and spine-shaped, postero-dorsal protrusion absent.

Type Locality
Philippines: Philippine Islands.

Distribution
Philippines: Luzon, Visayas and Mindanao Islands.

Remarks
According to what Jacobi (1905) mentioned at the end of the description "Mus. Berol. Nr. 9107: Semper coll., 2 Ex.". He examined two specimens and described this species from atleast one female without a description of the male terminalia. We illustrate and describe them here based on a specimen identifi ed using Lallemand's (1949) key.

Etymology
The species name refers to the translucid tegmen and is based on the latin word "translucidus" which means allowing light to pass through. HEAD (Figs 1A, 11B). In dorsal view, large ocelli, distance between eyes less than 8 times ocellus diameter, distance between ocelli equals one ocellus diameter, distance between ocellus and the compound eye 2 times ocellus diameter, ocelli closer to each other than from compound eyes. Eyes not prominent, length 1.44 times than wide. Vertex and frons longitudinal median carina absent. Vertex as long as wide with 3 times ocellus diameter in between two vertex grooves outside ocelli and 3 times ocellus diameter between anterior and posterior vertex margins. Postclypeus with longitudinal furrow, slightly swollen and ovoid shape in frontal view, widest part at mid height (Figs 1A, 11B), not receding and prior to anteclypeus where it bends forming right angle in lateral view (Fig. 11A). Rostrum long, surpassing mesocoxae. Thorax (Figs 2A, 11A-C). In dorsal view, pronotum with anterior concavities on each side, anterior margin as wide as posterior margin of head including eyes, anterolateral margins curved, posterior margin grooved, postero-lateral margins slightly concave, longer than anterolateral margins, humeral angle rounded. In lateral view, pronotum curving not more than 25º (Figs 2, 11A) Scutellum as long as wide with large median dimple (Fig. 11C). Tegmen (Fig. 11A-C). R bifurcates on apical half, M bifurcate on basal third, apical reticulation not well developed without concave apical cells. Posterior wing (Fig. 3A). Rp separating from SC+Ra nearly at midlength, M reaches ambient vein, Cua and Cup fused at base and m-cu links M to Cua before Cua bifurcation, common base for Cup and Cua originate at base of wing, 7 longitudinal veins and 5 apical cells between SC+Ra and Cup, angular protrusion of costal margin near its base. Metafemur with apical spine in inner margin, metatibiae bearing 1 lateral spine. MALE TERMINALIA. In lateral view, posterior margin of pygofer slightly undulating in the middle with slight curved on the last third (Fig. 11E). Subgenital plates (Fig. 11F) dorsal and ventral margin of main plate roughly straight, sterno-lateral plate present, slightly elongated, intermediate plate present, elongated slightly triangular shaped, linking internal sides of lateral and subgenital plates. Paramere (Fig. 11G) not globose, dorsal margin convex and regularly curving fi nishing with rounded apex with very minute groove, ventral margin convex, apex with spiniform process pointing antero-ventrally. Dorsal and ventral margins of aedeagus undulating. Aedeagus (Fig. 11H) with basal third of dorsal margin regularly bent without angle before the bent part and last 2 ⁄ 3 vertical and S-shaped, ventral margin regularly curved then slightly concaved before the base, apical extension sharp pointing postero-ventrally, posterior protrusion sharp at the apex hook-shaped, postero-dorsal protrusion absent.

Holotype
COLOR. Head and pronotum brown with yellow patches, rostrum yellowish white, pedicel of antenna brown, legs yellowish and abdomen light brown. Tegmen partially translucid, opaque parts being yellowish with darker patches.

Type species
Trigonosch e ma manoborum sp. nov. here designated.

Diagnosis
In dorsal view, teardrop shaped and apex of tegmen not visible due to folding. In lateral view, very steeply declivous pronotum and crown of the head, nearly form a right angle with the rest of the dorsum in profi le.

Etymology
When observed in dorsal view, the general shape of the habitus fi ts in a triangle even if a closer observation would lead to describe it as teardrop shaped. The name is built using two greek words "trigonos" for triangular and "schema" for shape. It is neutral.
MALE TERMINALIA. Subgenital plates without fi ne appendage (Fig. 12F), if present shorter than main plate, not forming an acute angle shape with the main plate (Figs 14, 16F). Paramere globose, lateral margin of paramere with protrusion, on the ventral margin on the sub apex with angular protrusion ornamented with set of setae (Figs 12, 14, 16G). Pp of aedeagus thick and beak-like shape (Figs 12, 14, 16H).

Distribution
Philippines.
Key to the species of Philippine Trigonoschema gen. nov. (Figs 12-13) (Figs 14-16) ..... 2 2. Red band running across base of tegmen and scutellum (Fig. 16) ( Lallemand, 1927) comb. nov. HEAD (Figs 1, 12-13B). In dorsal view, ocelli small, distance between eyes 10.5 times ocellus diameter (Figs 1, 12-13B). Distance between ocellus and compound eye less than 3.5 times ocellus diameter ( (Figs 1, 12-13B). Distance between ocelli 1.5 ocellus diameter (Figs 1, 12-13B). Eyes not prominent, length 1.15 times than wide. Vertex with longitudinal short median carina, slightly longer than wide. Frons without any carina. Postclypeus with dimple below margin of frons and a longitudinal furrow clearly marked in male, swollen laterally and slightly ovoid shape in frontal view, widest part close to frons, not receding and prior to anteclypeus where it bends rounded in lateral view (Figs 1B, 12-13A). Rostrum long, reaching but not surpassing mesocoxae. Thorax (Figs 2B, 12-13A-C). In dorsal view, pronotum with anterior deep concavities on each side, much clearly marked in female, longitudinal median carina absent. Anterior margin of pronotum as wide as posterior margin of head including eyes, anterolateral margins curved, posterior margin grooved, postero-lateral margins slightly concave, slightly longer than anterolateral margins, humeral angle rounded. In lateral view, pronotum angle around 45º (Figs 2B, 12-13A), scutellum as long as wide with large median dimple (Figs 12-13C). Tegmen (Figs 12-13A-C). R bifurcates on apical half, M bifurcate on basal third, apical reticulation not well developed without concave apical cells. Posterior wing (Fig. 13E). 2 ⁄ 3 of M length alone then fuses with Rp before reaching ambient vein, 6 longitudinal veins on posterior wing reaching apex, four apical cells between SC+Ra and Cup, angular projection near base of costal margin present. Legs. Metafemur with apical spine in inner margin, metatibiae bearing 1 lateral spine. MALE TERMINALIA. In lateral view, posterior margin of pygofer (Fig. 12E) largely convex in middle getting concave in last third and presenting a bump in fi rst third. Subgenital plates (Fig. 12F) with equal length relative to height of pygofer without any appendage, regularly tapering towards apex, dorsal margin straight then curving up towards apex, ventral margin straight half its length then regularly curving dorsally, minute denticuli covering last quarter toward apex. Sterno-lateral plate present, slightly elongated. Intermediate plates present, elongated and linked with internal sides of lateral plate and subgenital plate. Paramere (Fig. 12G) globose, dorsal margin convex then curving regularly and fi nishing with a sharp spine pointed antero dorsally, lateral margin with process in middle part pointing postero-dorsally, ventral margin regularly rounded with subapical part largely protruding posteriorly then fi nishing with a sharp spine pointing ventrally. Aedeagus (Fig. 12H) with dorsal margin slightly humped at base then concave most its length, curving regularly half its length vertically and fi nishing ¼ its length convex in a C-shaped, apical extension pointing posteriorly, posterior protrusion thick, duck head shaped with minute denticuli along posterior margin, in posterior view basal half lobular (Fig. 12I), postero-dorsal protrusion absent .

Trigonosch e ma manoborum
COLOR. Head, and pronotum white-yellowish with orange markings, postclypeus, rostrum, abdomen and legs orange, pedicel of antennae yellowish brown, scutellum orange with yellowish white markings medially. Tegmen, translucid yellowish white, white in basal ¼ with orange marking in between white and on the basal half of the margin of the clavus, last 3 ⁄ 4 yellowish getting yellowish brown in apical fourth.

Diagnosis
In general shape, T. negrosensis is similar to T. rubercella but their coloration is distinctly different. T. negrosensis in dorsal view presents a dirty yellow pronotum followed by a light orange scutellum framed by yellow irregular patches on clavus of tegmen while T. rubercella presents a bright yellow pronotum followed by a red band running on basal parts of tegmen and across scutellum.

Etymology
The species is named after the island where it was found: Negros. HEAD (Figs 1, 14B). In dorsal view, ocelli small, distance between eyes 9.5 times ocellus diameter (Figs 1, 14B). Distance between ocellus and compound eye less than 3 times ocellus diameter (Figs 1,14B). Distance between ocelli 1.5 ocellus diameter (Figs 1, 14B). Eyes not prominent, length 1.33 times than wide. Vertex with longitudinal median carina, as long as wide. Frons without median carina. Postclypeus with dimple below margin of frons and a longitudinal furrow clearly marked in males, swollen laterally and slightly ovoid shape in frontal view, widest part close to frons, not receding and prior to anteclypeus where it bends rounded in lateral view (Figs 1B, 14A). Rostrum long, reaching but not surpassing mesocoxae. Thorax (Figs 2B,. In dorsal view, pronotum with anterior deep concavities on each side, much clearly marked in female, longitudinal median carina absent. Anterior margin of pronotum as wide as posterior margin of head including eyes, anterolateral margins curved, posterior margin grooved, postero-lateral margins slightly concave, slightly longer than anterolateral margins, humeral angle rounded. In lateral view, pronotum angle around 45º (Figs 2B, 14A). Scutellum as long as wide with large median dimple (Fig. 14C). Tegmen (Fig. 14A, C). R bifurcates on apical half, M bifurcate on basal third, apical reticulation not well developed without concave apical cells. Posterior wing (Fig. 3A). Rp separating from SC+Ra nearly at midlength, M reaches ambient vein, Cua and Cup fused at base and m-cu links M to Cua before Cua bifurcation, 7 longitudinal veins and 5 apical cells between SC+Ra and Cup, angular protrusion of costal margin near its base. Metafemur with apical spine in inner margin, metatibiae bearing 1 lateral spine. MALE TERMINALIA. In lateral view, posterior margin of pygofer (Fig. 14E) convex in middle and clearly concave on last third. Subgenital plates (Fig. 14F) slightly longer relative to height of pygofer with fi ne appendage shorter forming acute angle with main plate, dorsal and ventral margin of main plate roughly straight. Sterno-lateral plate present and slightly elongated. Intermediate plates present, boomerangshaped, linking internal sides of lateral plate and subgenital plate. Paramere (Fig. 14G) globose, dorsal margin convex then curving regularly and fi nishing with a sharp spine pointed antero dorsally, lateral margin with process in middle part pointing dorsally, ventral margin regularly rounded with subapical part largely protruding posteriorly then fi nishing with two sharp spines pointing postero-ventrally. Aedeagus (Fig. 14H) with basal fourth almost straight then bending vertically and slightly wave-shape up to postero-dorsal protrusion, apical extension pointing dorsally, posterior protrusion thick, beakshaped, in posterior view, very slim at the top and petal shaped and foliaceous toward the base (Fig. 14I), postero-dorsal protrusion present.

Holotype
COLOR. Head including postclypeus and rostrum, legs and abdomen orange, pronotum orange white anterior half, greenish on apical part, scutellum greenish white with orange coloration, antennal scape and pedicel orange, fl agellum with brown coloration. Tegmen translucid yellow, basal third with white coloration in costal margin and clavus and with orange coloration in between white coloration, last 2 ⁄ 3 yellowish brown.

Distribution
Philippines: Philippine Islands.

Remarks
Lallemand in 1927 described this species based on the pattern of coloration and placed it in Mioscarta . He mentioned only a single male specimen (holotype), however, in the collection it is labelled as female (Fig. 15D). This species does not conform to Breddin's defi nition of Mioscarta but possesses characters of Trigonoschema gen. nov. For this reason we transfer it to this new genus. We could not borrow the holotype, however some photographs of the habitus and labels were kindly provided by Mick Webb (NHM). They are integrated here (Fig. 15) and are suffi cient to see the most obvious characters of Trigonoschema gen. nov.

Diagnosis
In general shape, T. rubercella is similar to T. negrosensis but is distinctly different in color. T. rubercella presents a bright yellow pronotum followed by red band running on the basal parts of the tegmen and scutellum when T. negrosensis in dorsal view presents a dirty yellow pronotum followed by a light orange scutelum framed by yellow patches sprawling on the clavus of the tegmen.

Etymology
This species has a distinct red coloration in the network of cells at the apex of tegmen. The species name is a combination of two latin words "ruber" and "cella" respectively meaning red and cell. The name is placed in apposition. HEAD (Figs 1, 16B). In dorsal view, ocelli small, distance between eyes 9 times ocellus diameter (Figs 1, 16B). Distance between ocellus and compound eye less than 4 times ocellus diameter (Figs 1, 16B). Distance between ocelli 1.5 ocellus diameter (Figs 1, 16B). Eyes not prominent, length 1.33 times than wide. Vertex slightly longer than wide. bearing a median longitudinal carina. Frons without carina. Postclypeus with dimple below margin of frons and a longitudinal furrow, swollen laterally and slightly ovoid shape in frontal view, widest part close to frons, not receding, prior to anteclypeus where it bends forming obtuse angle in lateral view (Figs 1B, 16A). Rostrum long, reaching but not surpassing mesocoxae.

Holotype
THORAX (Figs 2B, 16A-C). In dorsal view, pronotum with anterior deep concavities on each side, much clearly marked in female, longitudinal median carina absent. Anterior margin of pronotum as wide as posterior margin of head including eyes, anterolateral margins curved, posterior margin grooved, postero-lateral margins slightly concave, slightly longer than anterolateral margins, humeral angle rounded. In lateral view, pronotum angle around 45º (Figs 2B, 16A). Scutellum as long as wide with large median dimple (Fig. 16C). TEGMEN ( Fig. 16A-C). R bifurcates on apical half, M bifurcate on basal third, apical reticulation not well developed without concave apical cells. Posterior wing (Fig. 3A). Rp separating from SC+Ra nearly at midlength, M reaches ambient vein, Cua and Cup fused at base and m-cu links M to Cua before Cua bifurcation, 7 longitudinal veins and 5 apical cells between SC+Ra and Cup, angular protrusion of costal margin near its base present. Metafemur with apical spine in inner margin and metatibiae bearing 1 lateral spine. MALE TERMINALIA. In lateral view, posterior margin of pygofer (Fig. 16E) convex in middle with slight concavity toward ventral margin. Subgenital plates (Fig. 16F) with equal length relative to height of pygofer with fi ne appendage shorter than main plate directed posteriorly not forming acute angle with main plate, dorsal and ventral margin of main plate regularly curved. Sterno-lateral plates present, triangular shaped. Intermediate plates present, roughly boomerang-shaped linking internal sides of lateral plate and subgenital plate. Paramere (Fig. 16G) globose, dorsal margin convex then curving regularly and fi nishing with a sharp spine pointed dorsally, lateral margin with slightly and rounded, ventral margin roughly straight with subapical part angled largely protruding posteriorly then fi nishing with two sharp spines pointing postero-ventrally. Aedeagus (Fig. 16H) with dorsal margin making a right angle at its base, straight on a small portion before curving up regularly, apical part bending posteriorly and straight, apical extension pointing dorsally, posterior protrusion thick and beak-shaped, postero-dorsal protrusion absent. COLOR. Head, legs, abdomen, scutellum and anterior part of pronotum red, rest of pronotum yellow, rostrum red, antennal scape and pedicel reddish orange, fl agellum yellowish. Tegmen translucid yellow, basal and apical third including network of cells red.

Molecular Phylogeny
Results of the Bayesian 50% consensus tree and ML analyses with likelihood score of −13870.493 are shown in Figures 17 and 18 respectively. The resulting topologies are not similar with respect to the placement of Jacobsoniella and Wawi . In the Bayesian topology (Fig. 17), Jacobsoniella is recovered within a clade containing Eoscarta , Mioscarta , Poeciloterpa and Trigonoschema gen. nov. clades, each Fig. 17. Bayesian 50% consensus tree based on partitioned analysis of combined sequences Histone 3 + CO1 + 18S + 28S for Eoscarta Breddin, 1902, Jacobsoniella Melichar, 1914, Mioscarta Breddin, 1901, Poeciloterpa Stål, 1870 supported by a probability value (PPv) of 100%. This clade ( Eoscarta + Mioscarta + Poeciloterpa + Trigonoschema + Jacobsoniella ) is supported PPv of 82.61% and Wawi appears as sister group to it. However, the clade containing Wawi plus the fi ve other genera is supported with a high PPv of 100%. In the ML analysis (Fig. 18), Poeciloterpa and Trigonoschema gen. nov. are sister groups with a ML bootstrap value (MLBv) of 85.2/21% (SH-aLRT support/ standard bootstrap support). Then Mioscarta appears as sister group of ( Eoscarta + Wawi + ( Poeciloterpa + Trigonoschema )). Jacobsoniella is recovered basally as sister of the other ingroup taxa and all together the ingroup clade is supported with a MLBv of 100/100% branch support.

Distribution, Biology and Ecology
Eight species currently belonging to the genus Mioscarta (Fig. 19) and four species of the new genus Trigonosch e ma (Fig. 20) are known from the Philippines. Of the four species of Trigonoschema gen. nov., only T. pallida cannot be placed precisely since the locality provided by Lallemand (1927) was "Philippine Islands". Some specimens were collected using a light trap, thus very little information on their natural history is available to date. Such positive phototaxy was also reported by Soulier-Perkins & Kunz (2012). Their host-plants remain unknown, even if, some species of both genera were observed alighting on the leaves of different plants of secondary and primary forests. This observation is not enough to conclude the insect's hostplant. Direct observations therefore remain diffi cult. With the development of next generation sequencing it is possible now to identify plants using their barcoding even when in small quantity. It is most likely that in a near future, when analysing the content of the insect guts we should fi nd enough genetic material belonging to the ingested host plants that would allow their identifi cation.

Placement of Trigonoschema gen. nov. in the classifi cation
According to Liang & Webb (2002) the tribe Rhinaulacini is characterized by a relatively broad head with non-globose eyes and a concave posterior margin of pronotum. It is also usually mentioned that a central longitudinal concave postclypeus and a hind tibia bearing a single strong lateral spine (Lallemand 1949, Liang & Webb 2002. However, the most obvious character that can separate Rhinaulacini from other Cercopidae is the presence of a sterno-lateral plate between the pygofer and subgenital plate (Liang & Webb 2002). This sterno-lateral plate is observed in all new described taxa belonging to Eoscarta (Liang & Webb 2002), Amberana and Bourgoinrana (Soulier-Perkins & Kunz, 2012), Euryaulax (Liang et al. 2012), Wawi (Soulier-Perkins & Le Cesne 2016), Poeciloterpa (Crispolon et al. 2019), and Jacobsoniella (based on a specimen we identifi ed), and here in all new Mioscarta and Trigonoschema gen. nov. We are therefore certain in our placement of the new genus Trigonoschema in Rhinaulacini. Liang & Webb (2002) formally placed the Eoscartini as a junior synonym of Rhinaulacini without placing the genera in any of the existing subtribes due to the need of further studies on their male terminalia. They emphasized the diffi culties in identifying genera and species in this group, separations mainly based on small differences in male genitalia. Trigonoschema gen. nov. possesses characters that allow its clear identifi cation from the other Rhinaulacini genera found in the Philippines, specifi cally Mioscarta and Poeciloterpa . It appears as well close to those genera, which belong to the Poeciloterpina subtribe. But we have decided for now to follow Liang and Webb's example and leave Trigonoschema gen. nov. as incertae sedis in the Rhinaulacini. The phylogeny shows that Trigonoschema gen. nov. is a distinct clade from Mioscarta . Although the sample is reduced to only six genera with a partial representation for each of them, both resulting topologies (ML and BI) show a well-supported monophyly of Eoscarta , Mioscarta , Poeciloterpa, and Trigonoschema gen. nov., revealing Mioscarta and Trigonoschema gen. nov. as two distinct genera. This justifi es the description of the new genus Trigonoschema gen. nov . It is also clear from the differences in topologies between ML and BI, it would be unwise to make assumptions on genera relationships here. All the ingroup genera selected, except Jacobsoniella , belong to the Rhinaulacini and the ingroup support in both analyses is strong but no hasty conclusion should be made on placing Jacobsoniella in the Rhinaulacini or even believing this tribe to be monophyletic. Such hypotheses could only be tested with a larger sampling of Cercopinae genera.
Using the 50% majority rule topology highlighted the need of a wider analysis including the rest of Cercopidae to reconsider the family's relationships and to build up more robust phylogenetic hypotheses for the family. It is also congruent with the ML analysis, as the relationships between the targeted genera are not well established.