Phylogenetics and taxonomy of the Indo-Australian genus Ulonemia sensu Drake (Hemiptera: Tingidae), with the recognition of new genera and species collected from Proteaceae in Australia Author Shofner, Ryan Evolution and Ecology Research Centre, School of Biological, Earth, and Environmental Sciences, University of New South Wales, UNSW Sydney, NSW 2052, Australia r.shofner@unsw.edu.au Author Cassis, Gerasimos Evolution and Ecology Research Centre, School of Biological, Earth, and Environmental Sciences, University of New South Wales, UNSW Sydney, NSW 2052, Australia text Zoological Journal of the Linnean Society 2024 2023-08-28 200 2 360 425 http://dx.doi.org/10.1093/zoolinnean/zlad080 journal article 10.1093/zoolinnean/zlad080 0024-4082 11240237 E024A072-8083-4D91-BC40-A420D1280BEDC Proteatingis howardi Shofner and Cassis sp. nov. ( Figs 13I, J , 14I, J , 16 , 18 ) Zoobank registration: urn:lsid:zoobank.org:act: EA872671- B47E-4B30-B448-FFE0F398C1CE . Type material Holotype : AUSTRALIA : New South Wales : Victoria Park Nature Reserve , 7.2 km SSW of Alstonville , 28.90116°S 153.41002°E , 163 m , 07 Sep 2017 , S. Cullerton and R. Shofner , Macadamia tetraphylla L.A.S. Johnson ( Proteaceae ), det. R. Shofner , 1 ♂ (UNSW_ ENT 00046544 ) (AM). Paratypes : AUSTRALIA : New South Wales : Old Byron Bay Road , 1.6 km SE Newrybar , 28.73009°S 153.54454°E , 132 m , 14 Apr 2015 , R. Shofner , G. Cassis , and W. Sherwin , Macadamia sp. ( Proteaceae ), det. G. Cassis , 5 ♂ ( UNSW _ ENT 00046620 - UNSW _ ENT 00046624 ), 5 ♀ ( UNSW _ ENT 00046630 - UNSW _ ENT 00046634 ) ( ASCU ) . Victoria Park Nature Reserve , 7.2 km SSW of Alstonville , 28.90116°S 153.41002°E , 163 m , 07 Sep 2017 , S. Cullerton and R. Shofner , Macadamia tetraphylla L.A.S. Johnson ( Proteaceae ), det. R. Shofner , 3 ♂ ( UNSW _ ENT 00046541 , UNSW _ ENT 00046552 , UNSW _ ENT 00046553 ), 2 ♀ ( UNSW _ ENT 00046572 , UNSW _ ENT 00046581 ) ( ASCU ) , Macadamia tetraphylla L.A.S. Johnson ( Proteaceae ), det. R. Shofner , 24 ♂ ( UNSW _ ENT 00046542 , UNSW _ ENT 00046543 , UNSW _ ENT 00046545 - UNSW _ ENT 00046551 , UNSW _ ENT 00046554 - UNSW _ ENT 00046568 ), 18 ♀ ( UNSW _ ENT 00046569 - UNSW _ ENT 00046571 , UNSW _ ENT 00046573 - UNSW _ ENT 00046580 , UNSW _ ENT 00046582 - UNSW _ ENT 00046588 ) ( UNSW ) . Other material examined AUSTRALIA : New South Wales : Crooks Valley Road, 1.5 km S Crystal Creek, 28.32451°S 153.32056°E , 68 m , 05 Oct 2017 , R. Shofner, Macadamia tetraphylla L.A.S. Johnson ( Proteaceae ), det. R. Shofner, 2 ♂ ( UNSW _ENT 00046616, UNSW _ENT 00046617), 2 ♀ ( UNSW _ENT 00046618, UNSW _ENT 00046619) ( UNSW ). Old Byron Bay Road, 1.6 km SE Newrybar, 28.73009°S 153.54454°E , 132 m , 14 Apr 2015 , R. Shofner, G. Cassis, and W.Sherwin, Macadamia sp. ( Proteaceae ), det. G. Cassis, 22 ♂ ( UNSW _ENT 00046620- UNSW _ ENT 00046627, UNSW _ENT 00046640- UNSW _ENT 00046653), 14 ♀ ( UNSW _ENT 00046628- UNSW _ENT 00046636, UNSW _ENT 00046654- UNSW _ENT 00046658) ( UNSW ). Old Byron Bay Road, 1.8 km SE of Newrybar, 28.73213°S 153.54451°E , 130 m , 02 Oct 2017 , R. Shofner, Macadamia sp. ( Proteaceae ), det. R. Shofner, 5 ♂ ( UNSW _ ENT 00046609- UNSW _ENT 00046612, UNSW _ENT 00046615), 2 ♀ ( UNSW _ENT 00046613, UNSW _ENT 00046614) ( UNSW ). Queensland : Australia Zoo, 2.25 km N of Beerwah, 26.83752°S 152.96146°E , 55 m , 04 Oct 2017 , R. Shofner, Macadamia sp. ( Proteaceae ), det. R. Shofner, 1 ♂ ( UNSW _ENT 00046595), 1 ♀ ( UNSW _ENT 00046596) ( UNSW ). Blackall Range, 26.7°S 152.883°E , 25 May 1966 , D. A. I., Macadamia integrifolia Maiden and Betche ( Proteaceae ), 2 ♀ ( UNSW _ENT 00046516, UNSW _ENT 00046517), 1 ♂ ( UNSW _ENT 00046518), 1 unknown sex ( UNSW _ENT 00046519) ( QDPI ). Maleny, 26.767°S 152.85°E , 29 Mar 1966 , D. A. I., Macadamia integrifolia Maiden and Betche ( Proteaceae ), 2 ♂ ( UNSW _ENT 00046520, UNSW _ENT 00046521) ( QDPI ). Mapleton, 26.6248°S 152.86662°E , 28 Aug 1966 , D. A. I., Macadamia sp. ( Proteaceae ), 1 ♀ ( UNSW _ENT 00046522) ( QDPI ). Montville, 26.69°S 152.893°E , 09 May 1967 , D. A. I., Macadamia integrifolia Maiden and Betche ( Proteaceae ), 1 ♀ ( UNSW _ENT 00046515) ( QDPI ). Diagnosis: Proteatingis howardi can be recognized by the following combination of characters: dorsal surface with prominent pattern of dark brown reticulations; costal area areolae irregular, reticulated, appearing biseriate more than half the length of costal area. Male: Macropterous; males 2.99 ± 0.05 mm ( N = 10) ( Table 3 ). COLORATION. Brown or golden-brown, each pronotal carina with dark brown line medially on pronotal disc,with dark brown reticulate pattern covering hemelytra, with alternating light and dark bands on costal area of hemelytra and paranota, with pale maculae at juncture of cubitus and R + M veins. Head : dark brown; spines dark brown to golden-brown, lightening distally; bucculae dark brown, margins straw-coloured; eyes reddish-black. Antennae : AI–AII brown; AIII golden-brown; AIV brown, to golden-brown proximally. Rostrum : brown proximally, straw-coloured medially, dark brown apex. Pronotum : disc brown medially, with three dark brown lines medially along carinae, pale on posterior projection; calli dark brown to black; collum brown; paranota with alternating light and dark bands; carinae brown, occasionally paler posteriorly. Thoracic pleura and sterna : propleura brown to straw-coloured posteriorly; proepisternum, mesepisternum, and mesepimeron brown to straw-coloured along coxae; metepisternum brown, straw-coloured posteriorly; peritreme of metathoracic gland straw-coloured; pro-, meso-, and metasternum brown; sternal ridges straw-coloured. Hemelytra : brown or golden-brown, with dark brown reticulate pattern covering costal and sutural areas; alternating light and dark bands on costal area; pale maculae at juncture of cubitus and R + M veins. Legs : coxae brown; trochanters golden-brown; femora brown medially, golden-brown proximally and distally; tibiae and tarsi golden-brown. Abdominal venter : brown medially, darkening laterally. VESTITURE. Head : sparse distribution of golden-brown aciculate setae along either side of medial spine; occipital, medial, and frontal spines with short golden-brown slightly arcuate aciculate setae; waxy globules frequently present from median and occipital spines to lorum. Antennae : AI–AII with golden-brown sparse, aciculate arcuate setae; AIII- with golden-brown aciculate decumbent setae; AIV withgolden-brownaciculatesetaeandgolden-brownhairlikesetae. Pronotum : paranotal and anterior margins with arcuate aciculate setae, continuing posteriorly on medial and lateral carinae; disc with aciculate scale-like setae and numerous punctures with five to six cilia distributed evenly around internal rim of punctures; anterior angle frequently with waxy globules similar to head. Thoracic pleura and sterna : prosternum, mesepisternum, metasternum, and mesepimeron with arcuate aciculate setae and ciliate punctures as on pronotal disc. Legs : coxae to tarsi with golden-brown aciculate setae. Hemelytra : costal margin and veins of costal area sparsely covered with arcuate aciculate setae. Abdominal venter : decumbent to arcuate aciculate setae, denser posteriorly than anteriorly. STRUCTURE. Head : spines truncate; frontal spines convergent, in contact distally, less than half length of AI; medial spine straight, stout, height equal to frontal spines, width about half of height; occipital spines straight, cylindrical, often barely protruding beyond collum. Antennae : AI and AII short, AII subequal in length to AI; AIII long, length subequal to exceeding distance between humeral angles; AIV weakly clavate. Labium : moderate length, extending to anterior margin of metasternum. Pronotum : strongly convex; collum apex weakly inflated, height to two-thirds of disc; carinae minutely elevated off disc, one areola in height, medial and lateral carinae equal thickness, lateral carinae weakly divaricating anteriorly; paranota curved, following disc, upturned obliquely, one or two areolae wide. Thoracic sterna : mesosternal and metasternal carinae elevated, one areole wide, areolae rectangular; mesosternal carinae straight, parallel, metasternal carinae curved, truncateovate. Hemelytra : areolae small and uniformly sized in discoidal and subcostal areas; sutural area areolae similar to discoidal area proximally, increasing in size to 3× distally; costal area areolae small, irregular to rectangular one areole wide; subcostal area two areolae wide, areolae rectangular; discoidal area increasing from six to eight areolae wide, sutural area ten to eleven areolae wide. Male genitalia : pygophore ( Fig. 18A, B ); parameres ( Fig. 18C–E ); dorsal plate Y-shaped ( Fig. 18F ); aedeagus with paired elongate spinulate endosomal lobal sclerites ( Fig. 18F–H ); endosomal membrane spinulate ( Fig. 18F–H ). Female: Macropterous; 3.22 ± 0.07 mm ( N = 10) ( Table 3 ). COLORATION. As in male, occasionally ground colour slightly lighter than male. VESTITURE. As in male. STRUCTURE. As in male. Plant associations: Recorded from one genus and one family. Proteaceae : Macadamia integrifolia , seven specimens ; M . tetraphylla , 52 specimens ; and Macadamia . sp., 46 specimens . Distribution: Known from the Blackall Range and Glasshouse Mountains in Queensland , and the Border Ranges and Northern Rivers region in New South Wales ( Fig. 16 ). Etymology: Named after Bob Howard, who greatly supported this research; additionally, the first specimens of this species seen during this study were collected on his property. Remarks: Proteatingis howardi is similar to Pr . mjobergi but can be distinguished by the reticulate areolae of the costal area (vs. ovorectangular in Pr . mjobergi ). Proteatingis howardi is a major pest of macadamia nut crops in the same manner as C . decoris . It has caused significant economic losses by feeding on macadamia flowers, which wilt and fail to produce fruit.