Discussion: Schuh and Schwartz (2016) discussed a single female of an undescribed species of Pulvillophylus as occurring on the genus Calytrix in the large myrtaceous tribe Chaemelauciae DC. They ventured that this undescribed taxon might offer insight into the hosts of other species. The recording of Pulvillophylus ecdeiocoleae from the monocot family Ecdeiocoleaceae (Poales) suggests a radically different conclusion: that some or all members of Pulvillophylus are actually monocot feeders and that their shape and coloration help to conceal them on their hosts. As noted by Schuh and Schwartz (2016), most members of the Cremnorrhinina from outside Australia have an elongate head and projecting clypeus. Within the Australian fauna, most cremnorrhiniines are green or pale, with the obvious exception of most Pulvillophyllus species and some species of Halophylus Schuh and Schwartz, 2016. There are only a few Pulvillophylus species, including the two described in this paper, that have a long face and also offer a color resemblance to the seeds of some monocots, including members of the Ecdeiocoleaceae, Poaceae, and Restionaceae.

The Ecdeiocoleaceae, previously contained in the Restionaceae (Culter and Airy Shaw, 1965), is considered the sister taxon of the Poaceae (Givnish et al., 2010; Briggs and Tinker, 2014). The species of this drought-adapted family are endemic to a limited area of Western Australia (fig. 4A), wind pollinated and monoecious, with spikes containing separate zones of pistillate and staminate flowers developing in acropetal succession (Briggs and Tinker, 2014). Ecdeiocoleaceae contains three species placed in two genera (Ecdeiocolea monostachya, E. rigens B.G. Briggs, and Georgeantha hexandra B.G. Briggs and L.A.S. Johnson). Cassis and Gross (1995, 2002) do not list any Heteroptera species associated with Ecdeiocoleaceae. Only the ghost moth, Fraus simulans Walker, 1856 (Hepialidae), also found in several other Australian states, was reported to utilize E. monostachya (Main, 2001) as well as species of Poaceae (Hardy, 1973).

In the southern hemisphere the flora of East Africa is dominated by grasses and a large number of heteropterans are documented as feeding on the group, including within the Miridae (Collaria Provancher, 1872, Dolichomiris Reuter, 1882, Nabidomiris Poppius, 1914, and Stenotus Jakovlev, 1877). The monocot family Restionaceae is also diverse in southern Africa and has been documented as the host of several species in the family Blissidae (Slater, 1976, 1979) as well as the Miridae genus Zalmunna Distant, 1909 (R.T.S., personal obs.). Grasses also form a significant landscape element in Australia, but relatively very few species of Miridae have been documented as feeding on them (Cassis and Gross, 1995; Namayatova et al., 2013), but see Slater (1979) for information on the Blissidae and Cassis and Gross (2002) for treatments of other Australian pentatomorphan grass feeders. The Restionaceae are diverse in southwestern Australia, although they do not form such a conspicuous part of the landscape as they often do in the Cape region of South Africa. Leon and Weirauch (2015) described four species in the genus Restiophylus Leon and Weirauch, placing them in the phyline tribe Semiini, subtribe Semiina. The recorded hosts of the species are in the Restionaceae genera Hypolaena (H. exsulca R. Br., H. humilis (Gilg) B.G. Briggs and L.A.S. Johnson) and Leptocarpus (L. scariosus R. Br. and L. tenax (Labill.) R. Br.) and the Anarthriaceae genus Lyginia (L. barbata R. Br. and L. imberbis R. Br.) from southwestern Australia. These bugs have predominantly reddish coloration reminiscent of their restionaceous hosts and an elongate face, similar to that seen in some species of Pulvillophylus, although the male genitalia and pretarsus are distinct in the two groups. In the discussion of P. augustus Schuh and Schwartz, 2016, reference was made to a series of specimens from Hat Head National Park, New South Wales, taken on Leptospermum (Myrtaceae) with unknown generic placement. Schuh and Schwartz (2016: 176) suggested that these specimens were not members of the Cremnorrhinina; upon further examination of the genitalia we can now conclude that they represent a new species of Restiophylus and that perhaps the host association for these specimens is in error.

On the basis of these observations we posit that when the hosts of other members of Pulvillophylus are clarified most or all will be monocot feeders. We encourage future collectors to sample the Australian monocot fauna and document hosts in a more consistent fashion, as a way of testing this theory. To date, most specimens of the known taxa have been secured by general collectors, some at lights, but most probably through the use of sweep nets, the former method never providing host data and the latter usually applied without regard to better understanding the biology of the taxa being sampled.

Diagnosis: Recognized by the large size, somewhat projecting face, mostly uniform dark gray coloration, base of cuneus conspicuously yellowish white with narrow reddish brown adjoining margin; dorsum with mixture of obvious black and wooly sericeous setae (fig. 1A); endosoma sigmoid, ventral strap forming a relatively stout, strongly curving, hooked apical spine (figs. 1B, C, 2). Most similar in size and form of head to P. rubritinctus Schuh and Schwartz, 2016; distinguished by clavus and corium strongly reddish in that species, rather than dark gray in P. cuneomaculatus, as well as subtle differences in length of ventral strap distad to secondary gonopore and form of hook on apical endosomal spine. Endosomal structure similar to P. cuneotinctus (Schuh and Schwartz, 2016) with ventral strap terminating in small hooked spine; distinguished by longer strap extending beyond secondary gonopore, smaller eye of male and overall gray coloration with contrasting base of cuneus in P. cuneomaculatus. Overall coloration of this taxon is reminiscent of the Palearctic phyline species Europiella artemisiae (Becker, 1864).

Description: Male : Total length 5.85, pronotum width 1.35. COLORATION (fig. 1A): Uniformly dark gray, including most of appendages; vertex with faint medial mark, posterior margin of eye narrowly pale; midline of corium posteriad of clavus and basad of membrane faintly pale; base of cuneus and corium, narrowly adjacent to cuneus, conspicuously yellowish white, cuneus posteriad of white area narrowly reddish brown; antennal segment 2, except apex, much paler than segment 1; membrane including veins strongly fumose; femora narrowly pale apically, tibiae paler than femora. SURFACE AND VESTITURE (fig. 1A): Body surface with reclining, black, common setae, pronotum with some erect spinelike setae on anterior lobe, and head, pronotum, scutellum, clavus, and anteriormost portion of corium also with scattered, weakly flattened, woolly, sericeous setae. STRUCTURE (fig. 1A): Body elongate, nearly parallel sided. Head: Moderately long, eyes slightly bulging; frons faintly swollen, barely prognathous, projecting beyond anterior margin of eye by length of eye, clypeus visible from above; eye occupying about four-fifths height of head in lateral view; antenna inserted at ventral margin of eye, eye not emarginate at insertion; labium reaching just beyond apex of metacoxa. Antenna: Segment 1 elongate, exceeding apex of head by one-half length of segment, segment 2 long (1.75), 1.78 times width of head. Thorax: Pronotum weakly campanulate, lateral margins weakly sinuous, posterior lobe and calli weakly elevated, posterior margin straight; mesoscutum broadly exposed. Hemelytron: Costal margin very weakly convex, body elongate nearly parallel sided. GENITALIA (figs. 1B, C, 2): Pygophore: Elongate conical, ventral surface with faint transverse serrations. Endosoma: Sigmoid; ventral strap relatively short, subequal to length of secondary gonopore, forming sharply hooked spine with minute subtending tubercle; dorsal strap terminating subequal to distal edge of secondary gonopore. Phallotheca: Long, narrowly conical; aperture on anterior surface, long, with equal moderate width, attenuate distally; surface of basal portion strongly ribbed. Parameres: Left paramere with typical phyline structure. Right paramere of moderate size, broadly fusiform, with short terminal spine.

Female: Unknown.

Etymology: From the Latin, macula, “spot,” and cuneus, in reference to the conspicuous white base of the cuneus.

Host: Unknown.

Distribution (fig. 3): Known from the type locality, Kadji Kadji, in Western Australia.

Holotype: AUSTRALIA: Western Australia: Kadji Kadji Reserve, CY2, 29.23147°S 116.47769°E, 266 m, 17 Sep 2009, C. Young, black light bucket trap, Bush Blitz WA Sep 09, 1♂ (AMNH_PBI 00387550) (WAMP).

Diagnosis: Recognized by the moderately large size, projecting face, pale brown general coloration with black head, antenna, calli, and scutellum, reddish brown pattern on hemelytron dorsum, male with fusiform antennal segment 2 (fig. 1D); endosoma sigmoid, apical region equal to length of secondary gonopore with simple hooked terminal spine (figs. 1E, F, 2). Most similar in size, form of head, and coloration, to P. rubritinctus; distinguished by slightly smaller size, distinct black coloration on head, pronotum, scutellum, and anterior region hemelytron, fusiform antennal segment 2, and endosoma with shorter ventral strap distad of secondary gonopore.

Description: Male: Mean total length 5.00, mean pronotum width 1.35. COLORATION (fig. 1D): Background coloration tan to pale brown; dark brown to black on antennal segments, head, except medially on vertex, calli, mesoscutum, scutellum, and costal vein; reddish brown to red on pair of ray-shaped markings emanating posteriad of calli reaching posterior margin of pronotum, lateral spots on mesoscutum, clavus posteriorly, endocorium except medially, and medial strip on cuneus; membrane including veins fumose; hind femora dark reddish brown, remaining femora, tibiae, and tarsi paler reddish brown. SURFACE AND VESTITURE (fig. 1D): Body surface smooth, very weakly shining, with reclining, black, common setae, pronotum with some erect spinelike setae. STRUCTURE (fig. 1D): Body elongate ovoid. Head: Head prognathous, frons flattened, projecting beyond eye by about length of eye, clypeus projecting and visible in dorsal view; eyes large angulate along lateral margin of head in dorsal view; eye occupying 90% height of head in lateral view; antenna inserted just above ventral margin of eye; labium surpassing apex of metacoxa and reaching onto abdomen. Antenna: Segment 1 relatively short, exceeding apex of head by one-half length of segment, segment 2 fusiform, relatively short (1.43), 1.55 times width of head. Thorax: Pronotum weakly campanulate, lateral margin slightly concave, posterior lobe flattened, posterior margin excavated; mesoscutum broadly exposed. Hemelytron: Costal margin slightly convex. GENITALIA (figs. 1E, F, 2): Pygophore: Elongate somewhat parallel sided distally, ventral surface with prominent serration. Endosoma: Sigmoid; ventral strap moderately long, equal to length of secondary gonopore, forming sharply hooked spine with one accessory spine and two spicules basad of hooked apical spine; dorsal strap terminating equal to distal edge of secondary gonopore. Phallotheca: Long, narrowly conical; aperture on anterior surface, long, with equal moderate width, attenuate distally; surface of basal portion strongly ribbed. Parameres: Left paramere with typical phyline structure. Right paramere of moderate size, fusiform, with abruptly narrow terminal spine.

Female: (fig. 1G): Structure and coloration similar to male, except body more ovoid, antennal segment 2 narrower, slightly bowed; mean total length 5.90, mean pronotum width 1.56. GENITALIA: Subgenital plate of sternite 6: With posteriorly directed conical projection. Vestibular sclerites: Medium size, not attaining anterior edge of dorsal labiate plate (fig. 1H–J). First gonapophyses: Small basal blocks (fig. 1K). Ventral labiate plate: Platelike medial anteroventral extension absent (fig. 1K). Dorsal labiate plate: Medium size, short longitudinally (fig. 1H). Sclerotized rings: Moderately large, subovoid, anterior angle pointed, relatively flat, thin walled; dorsal labiate plate ventrad of rings finely spiculate (fig. 1H). Posteromedial region: Surface without conspicuous microstructure. Anterolateral region: Anterior margin extending slightly beyond anterior edge of sclerotized rings, narrow band spanning rings. Posterior wall: Intersegmental structure: Flat, broadly bilobed, platelike, apically rounded. Interramal sclerites: Strongly sclerotized, lateral sclerites widest mediad, narrowed laterally, medial sclerite bell shaped, produced posteriorly (fig. 1J, L).

Etymology: Named for the association with a plant species in the genus Ecdeiocolea F. Mull., which currently contains the only known host for any species of Pulvillophylus. A noun in genitive case.

Host: All specimens were taken on the flowering spikes of Ecdeiocolea monostachya (Ecdeiocoleaceae) (C. Symonds, personal commun.) (fig. 4).

Distribution (fig. 3): Known only from the type locality near Lochada, Western Australia.

Holotype: AUSTRALIA: Western Australia: Lochada, Omega track 1.5 km N of Mungada Rd, 29.16827°S 116.54336°E, 281 m, 18 Sep 2009, C. Symonds, Bush Blitz WA Sep 09, Ecdeiocolea monostachya F. Muell. (Ecdeiocoleaceae), det. WA Herbarium, 1♂ (AMNH_ PBI 00387551) (WAMP).

Paratypes: AUSTRALIA: Western Australia: Lochada, Omega track 1.5 km N of Mungada Rd, 29.16827°S 116.54336°E, 281 m, 18 Sep 2009, C. Symonds, Ecdeiocolea monostachya F. Muell. (Ecdeiocoleaceae), det. WA Herbarium, 1 ♀ (AMNH_PBI 00387556) (AMNH), 1♂ (AMNH_PBI 00387554), 4♀ (AMNH_PBI 00387552, AMNH_PBI 00387555, AMNH_PBI 00387557, AMNH_PBI 00387558) (UNSW), 3♀ (AMNH_PBI 00387559-AMNH_PBI 00387561) (WAMP).

Other Specimens Examined: AUSTRALIA: Western Australia: Lochada, Omega track 1.5 km N of Mungada Rd, 29.16827°S 116.54336°E, 281 m, 18 Sep 2009, C. Symonds, Bush Blitz WA Sep 09, Ecdeiocolea monostachya F. Muell. (Ecdeiocoleaceae), det. WA Herbarium, 1 nymph; (AMNH_PBI 00387562) (UNSW).

Discussion: If our prediction that Pulvillophylus species for which hosts are currently unknown are also monocot feeders, then we would also predict that the eventual hosts are in families other than the Ecdeiocoleaceae, because most of those species have been collected outside the known distributions of members of the Ecdeiocoleaceae (Briggs and Tinker, 2014). Only P. cuneomaculatus, n. sp., P. rossi Schuh and Schwartz, 2016, and an unnamed species, known from one female, appear to have been collected within the range of ecdeiocoleaceous species (see fig. 4A; collection records of the other two species of Ecdeicoleaceae are subsumed within the range of E. monostachya).