New genus and two new species of Hyaliodini from the Philippines (Miridae, Deraeocorinae)

Abstract Philicoris, a new genus of the mirid subfamily Deraeocorinae, tribe Hyaliodini, is described from the Philippines. New species Philicorismayonsp. n. and Philicorispalalisp. n. from the island of Luzon are documented with photographic images of the dorsal habitus and male genital structures.


Introduction
The island nation of the Philippines is home to a unique and highly endemic fauna, and as such is one of a small number of nations recognized globally as both a conservation biodiversity hotspot (Myers et al. 2000) and a megadiverse nation (Mittermeier et al. 1997). The capital city of Manila is on Luzon, a large island in the northern Philippines formed historically by the accretion of separate paleo-islands in the geological past (Adams and Pratt 1911;Rutland 1968;Hashimoto 1981a, b;Auffenberg 1988;Hall 1996;Yumul et al. 2009;Siler et al. 2011;. It is assumed that the isolation and subsequent accretion of these precursor paleo-islands contributed to the evolutionary diversity found today (Brown et al. 1996; Brown and Dies- Philicoris does not possess the elongate anal tube in both males and females (Ferreira et al. 2015), or the stridulatory structures on the embolium (Akingbohungbe 1979), but these characters are not consistent across all genera of the tribe (e.g., Linnavuorista Akingbohungbe). No genera of Hyaliodini are recorded from the Philippines (Schuh 2013), and we are unable to key the newly collected specimens to any known genera in Akingbohungbe's (1979) world key.
In this paper, external and internal genital features are used to demonstrate that both mirid populations on Luzon Island represent distinct evolutionary lineages (Wiley 1978;de Queiroz 1998de Queiroz , 1999 and are worthy of taxonomic recognition as members of a new genus in the tribe Hyaliodini. The recognition of these taxa represents the first records of the Hyaliodini in the Philippines. A new genus and two new species are described, technical images of key diagnostic traits provided, and its natural history, ecology, and geographic distribution are discussed. This manuscript was inspired by Dr. Thomas Henry, and this Festschrift article is dedicated to him. He never left a plant unsampled in fieldwork, and described hundreds of mirids across the world.

Materials and methods
All specimens were collected by KLM in 2016 and 2017 as part of Sam Noble Oklahoma Museum of Natural History expeditions to the Philippines in conjunction with the National Science Foundation Grant (NSF IOS 1353683) to CDS. As part of the Memorandum of Agreement with the Philippines, all specimens are temporarily deposited at the Sam Noble Oklahoma Museum of Natural History, Norman, Oklahoma, United States (OMNH).
Stacked habitus photos were taken at the Sam Noble Museum Invertebrate Paleontology Stacking Photography Lab. Color photos and illustrations were edited using Adobe Photoshop CS4 and numbered in Adobe Illustrator CS4. Helicon Focus v4.2.9 software was used for all image stacking.
Scanning Electron Micrographs (SEM) were taken at the Samuel Roberts Noble Microscopy Laboratory at the University of Oklahoma. Dried specimens attached to paper points were removed from the pins, and the points used as mounts to attach to aluminum stubs with carbon conductive tape. The stubs and specimens were then coated with gold/ palladium and examined with a Zeiss NEON 40 EsB SEM or a LEO 1450VP SEM.
Terminology for male genitalia follows Cassis (2008). Genitalia were extracted by soaking the abdomen in 85% lactic acid solution for one to a few days at room temperature, and then washing with distilled water before being dissected in glycerol. Images of the male genitalia were taken using an Olympus CX31 compound microscope with an Olympus 5MP CCD Camera using Cellsens Software. Helicon Focus v4.2.9 software was used for all image stacking.
Measurements were taken using an eyepiece micrometer (10mm/100×) on an Olympus SZX2 stereomicroscope following the methodology of Schuh (1984). Measurements include the body length (clypeus to cuneus), pronotum length (anterior margin to posterior margin), second antennal segment length, head width (distance between lateral margins of the eyes), pronotum width (width at widest point along the posterior margin), width at the widest point of the hemelytra, and interocular distance. All measurements are in millimeters.
Included species. Philicoris mayon sp. n., Philicoris palali sp. n. Diagnosis. Recognized by trapezoidal and convexly rounded pronotum, shiny dorsum with punctation on pronotum and distinct rows of punctures along clavus and corium of hemelytron, wide emboliar margins, and shape of male genitalia.
Description. Male. Coloration. Head: tan to light yellowish brown, clypeus and labrum dark brown; labium basally yellow and dark brown apically; first antennal segment dark brown, basally dark brown, transitioning to tan or orange to light brown distally with dark brown apices, second antennal segment contiguously tan or yellow transitioning to reddish brown then dark brown at distal apex, third antennal segment tan or white basally transitioning to dark brown distally, fourth antennal segment tan or dark brown. Thorax: collar tan or yellowish brown, pronotum tan or light yellowish brown, calli tan with posterior margin with lighter yellow macula or orange tinge along margin, mesoscutum brown with lateral yellow macula or light yellowish brown, scutellum dark brown medially and light whitish green along majority of lateral margins or light yellowish brown, lateral sclerites of thorax same coloration as pronotum or darker, scent gland and evaporative area same color as lateral sclerites or contrastingly whitish, procoxae light yellow or orange-brown, remaining coxae light brown apically and yellowish or orange-brown distally, femora tan or light yellowish brown with two orange-red stripes on preapical distal margins, and tibiae tan or orange and light yellowish brown basally, tarsomeres tan or light yellow. Hemelytra: embolium tan or light yellowish brown with orange tinge, cuneus tan or light yellowish brown with orange tinge, lateral margins of corium tan or light yellowish brown transitioning to light brown, clavus dark brown with light tan area along anterior one-third margin or with corium completely dark brown, membrane light brown to beige with orange-brown or beige veins. Abdomen: tan with dark brown anterior and posterior surfaces or completely-orange brown, gonopore tan or orange-brown. Surface and vestiture: Head: smooth and shiny, clothed with simple setae, antennal segments covered with dense simple setae of uniform length. Thorax: collar and calli smooth, remaining surface of pronotum punctate, covered with simple setae, scutellum and lateral sclerites of thorax with simple setae. Hemelytra: simple setae covering hemelytral surface, hemelytra surface shiny with dense punctuation in regular rows on clavus and corium, dorsal surface of emboliar margins and cuneus smooth. Abdomen: clothed with simple setae. Structure: Head: wider than high, clypeus not visible in dorsal view, frons convex, vertex flat, declining posteriorly towards anterior pronotal margin, eyes relatively large, taking up most of head in lateral view, dorsal surface confluent with vertex, posterior margin removed from anterior margin of pronotum, interocular width either greater than or less than width of single eye, first antennal segment length wider than interocular distance, less than half length of second segment, second antennal segment longest, apically narrower than first segment, distally widening to width equivalent to first, antennal segments three and four half width of antennal segment one, individually nearly equidistant in length to segment one; apex of labium extending to metacoxae. Thorax: pronotal collar narrow and rounded, dorsal surface of pronotum convexly rounded, anterior and posterior portions of pronotum not demarcated, pronotum trapezoidal with nearly straight lateral margins, calli fused into single weakly protruding plate surrounded by weakly defined rows of punctures, posterior margin of pronotum straight, mesoscutum mostly hidden or not visible, scutellum tumid, metathoracic scent gland relatively large, taking up greater than half area of metathoracic sclerite (Fig. 3A), femoral length equivalent and approximately four-fifth length of tibiae, metafemora greatest in width, medial width widest medially, tibial length nearly equivalent to emboliar length, third segment of pretarsus longest, pretarsal claws with basal tooth (Fig. 3B), parallel hair-like parempodia, lacking pulvilli. Hemelytra: weakly transversely rounded, lateral margins straight or weakly concave with wide embolium, cuneus longer than wide, bent ventrally at fracture, membrane with two visible veins forming two cells, larger cell length greater than one half total length of membrane. Abdomen: shorter than half total body length, relatively narrow, width tapering to gonopore. Genitalia: Endosoma primarily membranous with two spicules apically, preapical secondary gonopore and several membranous apical lobes (Figs 4A, 5A), phallotheca thin and simple, left paramere tall and crow-bar shaped, sometimes with basal spine projecting perpendicularly to base (Figs 4C, 5C), right paramere small, leaf-shaped, sometimes with apical bifurcation (Figs 4B, 5B).
Female. Similar to males in coloration, surface and vestiture. Interocular distance greater in females, width of pronotum sometimes wider. Genitalia: not dissected.
Etymology. The generic name is derived from the country of their discovery (Philippines) and "coris," meaning bug in Latin. The gender is masculine.
Hosts. Asteraceae. Distribution. Luzon Island, Philippines. Discussion. The combination of the basally toothed claw, the relatively wide embolium along the lateral margins of the hemelytron, the shiny and punctate pronotum and hemelytron, and the membranous endosoma clearly place this genus in the Hyaliodini as it is currently defined. Both species also have two apical endosomal spicules and multiple membranous lobes that correspond to other members of Deraeocorinae and Hyaliodini (Akingbohungbe 1979).
Philicoris is similar to the Neotropical genus Antias Distant in size and lacks a demarcation between the anterior and posterior portions of the pronotum but instead is tumid, and the hemelytron is not completely hyaline lines (Carvalho 1982). The calli are also not well developed. Unlike Antias, however, Philicoris has a smooth membrane without any setae, the eyes do not cover the entire area of the head in lateral view, and Philicoris has rows of punctation on the surface of the corium, embolium and claval suture lines. Philicoris also differs from the African genera Obudua Linnavuori and Linnavuorista Akingbohungbe by the lack of a narrowed anterior margin, the former by the lack of distinctly swollen calli, and from the latter by lacking a distinct apical spur on the metafemur. The Palearctic and African genus Stethoconus Flor also differs from Philicoris; the latter lacks an anterior constriction to the pronotum, has a narrow collar, and has a relatively concolorous pronotum and hemelytron versus the highly patterned Stethoconus. For these reasons, we argue that this taxon represents a new genus. Diagnosis. Distinguished from its congener by the following combination of characters: (1) reddish orange coloration on lateral margins of hemelytron and legs; (2) extensive and defined rows of punctuation; (3) interocular distance relatively narrow; and (4) endosomal spicules weakly sclerotized.

Philicoris mayon
Description. Male. Coloration. Head: light yellowish brown, clypeus and labrum dark brown; labium yellow basally and dark brown apically; first antennal segment light orange-brown with dark brown apices, second antennal segment yellow transitioning to reddish brown then dark brown at distal apex, third antennal segment white basally, dark brown distally, fourth segment completely dark brown. Thorax: light yellowish brown, punctures orange-brown, collar lighter yellow, margins of calli more orange, mesoscutellum and scutellum light yellowish brown as pronotum, lateral sclerites of thorax same coloration as pronotum or darker brown, scent gland with same coloration as lateral sclerites, coxae and trochanters light orange-brown, femora light yellowish brown with two orange-red stripes on preapical distal margins, tibiae orange basally and light yellowish brown distally, tarsomeres light yellow. Hemelytra: light yellowish brown with orange tinge, orange tinge most predominant along emboliar margins and cuneus, clavus darker orange-brown to dark brown, corium darker yellowish brown, membrane pale brown with orange veins. Abdomen: orange-brown. Surface and vestiture: Head: smooth and shiny, clothed with dense simple setae, antennal segments covered with simple setae of uniform length. Thorax: collar and calli smooth, remaining surface of pronotum punctate, covered with dense simple setae,  scutellum and lateral sclerites of thorax with simple setae. Hemelytra: simple setae covering hemelytral surface, hemelytra surface shiny with dense punctuation in regular rows on clavus, and corium, dorsal surface of emboliar margins and cuneus smooth. Abdomen: clothed with simple setae. Structure: Head: wider than high, clypeus not visible in dorsal view, frons convex, vertex flat, declining posteriorly towards anterior pronotal margin, eyes relatively large, taking up most of head in lateral view, dorsal surface confluent with vertex, posterior margin removed from anterior margin of pronotum, interocular width narrower than width of single eye, first antennal segment length wider than interocular distance, less than half length of second segment, second antennal segment longest, apically narrower than first segment, distally widening to width equivalent to first, antennal segments three and four half width of antennal segment one, individually nearly equidistant in length to segment one; apex of labium extending to metacoxae. Thorax: pronotal collar narrow and rounded, dorsal surface of pronotum convexly rounded, anterior and posterior portions of pronotum not demarcated, pronotum trapezoidal with nearly straight lateral margins, calli fused into single weakly protruding plate surrounded by well-defined rows of punctures, posterior margin of pronotum straight, mesoscutum mostly hidden or not visible, scutel-lum tumid, metathoracic scent gland relatively large, taking up greater than half area of metathoracic sclerite (Fig. 3A), pretarsal claws with basal tooth, parallel hair-like parempodia, pulvilli absent (Fig. 3B). Hemelytra: weakly transversely rounded, lateral margins convex with relatively wide embolium, cuneus longer than wide, bent ventrally at fracture, membrane with two visible veins forming two cells, larger cell length greater than one half total length of membrane. Abdomen: shorter than half total body length, relatively narrow, width tapering to gonophore. Genitalia: Endosoma primarily membranous with two weakly sclerotized spicules and preapical secondary gonopore surrounded by several membranous apical lobes (Fig. 4A), phallotheca thin and simple, left paramere tall and scythe-shaped with basal spine projecting perpendicularly to base and apex with round "hook" (Fig. 4C), right paramere small, leaf-shaped with apex pointed (Fig. 4B).
Female. Similar to males in coloration, surface and vestiture. Pronotum wider in females, interocular distance greater. Genitalia not dissected.
Etymology. Named for Mt. Mayon, the type locality. Noun in apposition.

Hosts. Unknown.
Distribution. Luzon Island, Philippines. Discussion. This species was found on an unidentified prostrate plant in a relatively agricultural area of Mt. Mayon, around banana and palm farms. Its coloration is unique and roughly matches the pinkish red flowers of the plant it was found on. Paratype. One adult female (OMNH 65501), same information as holotype. Diagnosis. Recognized by mostly tan overall coloration, lateral pale greenish maculation on scutellum, white scent gland, tan thorax, interocular distance wider than width of eye, relatively narrower and straight emboliar margin, less prominent and defined surface punctuation, left paramere lacking basal perpendicular spine, and right paramere apically bifurcate.

Philicoris palali
Description. Male. Coloration. Head: tan, clypeus and labrum dark brown; labium basally yellow and dark brown apically; first antennal segment basally dark brown, distally transitioning to tan, remaining segments tan. Thorax: collar and pronotum tan, calli tan with posterior margin with lighter yellow macula, mesoscutum brown with lateral yellow macula when visible, scutellum dark brown medially and light whitish green along majority of lateral margins, apex yellow, lateral sclerites of thorax same coloration as anterior pronotum, scent gland and evaporative area whitish, procoxae light yellow, remaining coxae light brown apically, yellowish distally, femora and tibiae tan, tarsomeres tan. Hemelytra: embolium and cuneus tan, lateral margins of corium tan transitioning to light brown, clavus dark brown with light tan area along anterior 1/3 of margin with corium, and corium, membrane and veins beige. Abdomen: tan with dark brown anterior and posterior surfaces, gonopore tan. Surface and vestiture: Head: smooth and shiny, clothed with simple setae, antennal segments covered with dense simple setae of uniform length. Thorax: collar and calli smooth, remaining surface of pronotum punctate, covered with simple setae, scutellum and lateral sclerites of thorax with simple setae. Hemelytra: surface with simple setae, shiny, with dense punctuation in regular rows on clavus and corium, dorsal surface of emboliar margins and cuneus smooth. Abdomen: clothed with simple setae. Structure: Head: wider than high, clypeus not visible in dorsal view, frons convex, vertex flat, declining posteriorly towards anterior pronotal margin, eyes relatively large, taking up most of head in lat-eral view, dorsal surface confluent with vertex, posterior margin removed from anterior margin of pronotum, interocular width greater than width of single eye, first antennal segment length wider than interocular distance, less than half length of second segment, second antennal segment longest, apically narrower than first segment, widening distally to width equivalent to first, antennal segments three and four half width of antennal segment one, individually nearly equidistant in length to segment one; apex of labium extending to metacoxae. Thorax: pronotal collar narrow and rounded, dorsal surface of pronotum convexly rounded, anterior and posterior portions of pronotum not demarcated, pronotum trapezoidal with nearly straight lateral margins, calli fused into single weakly protruding plate surrounded by weakly defined rows of punctures, posterior margin of pronotum straight. Hemelytra: weakly transversely rounded, lateral margins straight with relatively wide embolium, cuneus longer than wide, bent ventrally at fracture, membrane with two visible veins forming two cells, larger cell length greater than one half total length of membrane. Abdomen: shorter than half total body length, relatively narrow, width tapering to gonophore. Genitalia: endosoma primarily membranous with two sclerotized spicules surrounded by several membranous apical lobes (Fig. 5A), phallotheca thin and simple, left paramere scythe-shaped without spine projecting perpendicularly to base (Fig. 5C), right paramere small, leaf-shaped with apical bifurcation (Fig. 5B).
Female. Similar to males in coloration, surface and vestiture. Interocular distance greater in females. Genitalia not dissected.
Etymology. We name the new species in reference to Mt. Palali, the type locality. Noun in apposition.
Hosts. Purple composite (Asteraceae). Distribution. The new species is known from mid-elevation habitats at the type locality on Mt. Palali, Nueva Vizcaya Province, Luzon Island, Philippines.
Discussion. This species was found by sweeping an unidentified purple composite along a trail up Mt. Palali, approximately 800 m away from an area cleared for banana and agricultural cultivation. Therefore, the host plant might be an introduced species in the regional flora. The additional sampling of local plants, including varieties introduced for agriculture, may yet yield additional new species.
ble Museum's existing Memorandum of Agreement with the BMB of the Philippines (2015-2020), Gratuitous Permits to Collect No. 247 (2016) and 260 (Renewal;2017), and Institutional Animal Care and Use Committee (IACUC) approved protocols R13-012 and R17-019. Financial support for fieldwork was provided by National Science Foundation, Division of Integrative Organismal Systems grant (NSF IOS 1353683) to CDS. Preston Larson is thanked for assistance with preparing and imaging the specimens using the SEM at the Samuel Roberts Noble Microscopy Laboratory. For access to the Sam Noble Museum Invertebrate Paleontology Stacking Photography Lab, S. Westrop and R. Burkhalter are appreciated. In the field Jason Fernandez and the Philippine Field Team were critical for field assistance during faunal surveys of Mt. Mayon and Mt. Palali on Luzon Island, and Dr. Thomas Henry and Dr. Michael Schwartz are thanked for their advice on dissections of Deraeocorinae to prepare the figures. Lastly, we thank members of the Menard and Siler labs, and Dr. Alfred Wheeler for his helpful critiques of the manuscript.