-FEEDING PHYLINAE (HEMIPTERA: MIRIDAE) FROM AUSTRALIA: DESCRIPTION AND ANALYSIS OF PHYLOGENETIC AND HOST RELATIONSHIPS FOR A MONOPHYLETIC ASSEMBLAGE OF THREE NEW GENERA

RANDALL T. SCHUH George Willett Curator, Division of Invertebrate Zoology American Museum of Natural History ([email protected])

CHRISTIANE WEIRAUCH Associate Professor, Department of Entomology University of California at Riverside ([email protected])

BULLETIN OF THE AMERICAN MUSEUM OF NATURAL HISTORY Number 344, 95 pp., 37 figures, 9 plates, 2 tables Issued December 8, 2010

Copyright E American Museum of Natural History 2010 ISSN 0003-0090 CONTENTS

Abstract...... 5 Introduction...... 5 TaxonomicMaterialsandMethods...... 5 Key to Melaleucoides genusgroupgenera...... 12 Harpagophylus,newgenus...... 12 Key to species of Harpagophylus ...... 13 Harpgophylus agnew, new species ...... 14 Harpgophylus , new species...... 15 Harpgophylus scholtizii,newspecies...... 16 Harpgophylus thryptomeni, new species ...... 17 Harpgophylus verticordii,newspecies...... 18 Thryptomenomiris,newgenus...... 21 Thryptomenomiris kalbarri, new species...... 22 Thryptomenomiris yalgoo,newspecies...... 23 Melaleucoides,newgenus...... 26 Key to species of Melaleucoides ...... 27 Melaleucoides akaina,newspecies...... 30 Melaleucoides annae, new species ...... 32 Melaleucoides beaufortiae, new species ...... 35 Melaleucoides brevifoliae,newspecies...... 38 Melaleucoides cassisi,newspecies...... 40 Melaleucoides castanea,newspecies...... 43 Melaleucoides grossi,newspecies...... 44 Melaleucoides leuropomae,newspecies...... 45 Melaleucoides micranthae,newspecies...... 48 Melaleucoides ozzii,newspecies...... 50 Melaleucoides pileanthicola, new species ...... 52 Melaleucoides rhaphiophyllae,newspecies...... 55 Melaleucoides sheathianae, new species ...... 57 Melaleucoides similis,newspecies...... 59 Melaleucoides systenae, new species ...... 60 Melaleucoides uncinatae, new species ...... 63 Melaleucoides undulatae,newspecies...... 65 Melaleucoides verticordiae, new species ...... 67 Phylogenetic Analysis ...... 71 Host Relationships ...... 78 DistributionalPatterns...... 78 Acknowledgments...... 79 References...... 79 Platesfollowingpage...... 85

3 ABSTRACT

Three new genera and 25 new species of Myrtaceae-feeding Phylinae are described from Australia. A cladistic analysis of a broader cross section of Australian Phylinae indicates that these taxa all belong to a monophyletic group, on the basis of pretarsal and male genitalic structures. Line drawings are used to present information of male and female genitalic structures; scanning electron micrographs illustrate pretarsal, genitalic, and other morphology. Color digital habitus images are presented for the males and females of all species. Digital images from the field and of pressed specimens are presented for a broad cross section of host . The newly described taxa represent a portion of the species of Australian Phylinae known to feed on the Myrtaceae; the remaining Myrtaceae-feeding species belong to other lineages and are therefore excluded from this monograph. Twenty-three of the species are recorded only from southwestern ; two species are recorded from southeastern South Australia. All species breed on members of the subfamily Myrtoideae, tribes and Melaleuceae.

INTRODUCTION endosomal types within certain Australian Phylinae. Further study revealed that not This paper represents part of an ongoing only was the endosoma distinctive, but that effort to document the fauna of Miridae, or the parempodia and left paramere were also bugs, of Australia, including distribu- novel, permitting the recognition of a mono- tional patterns and associations with host phyletic taxon that we here refer to as the plants. Although in the past this fauna has Melaleucoides group and that is received only limited study (Carvalho and restricted to host plants within two tribes of Gross, 1982; Malipatil, 1992), the nature of the family Myrtaceae: Myrtoideae. its actual diversity is beginning to take shape. Within the Phylini (Phylinae), a group with TAXONOMIC MATERIALS its greatest diversity in the Mediterranean- AND METHODS climate regions of the world, recent papers have documented well-defined associations Unique specimen identifiers: During the with a variety of host lineages within course of this project matrix code labels were Australia, the occurrence of both Indo- affixed to about 2350 specimens examined as Australian (Schuh and Menard, in prep.) a way to uniquely identify them; these codes and Transantarctic distributional patterns are therefore referred to as ‘‘unique specimen (Weirauch and Schuh, in press), and radia- identifiers’’ (USIs). The USI codes, e.g., tion of Miridae in association with extreme AMNH_PBI 00094810, are composed of an floristic diversity in southwestern Australia institution and project code (AMNH_PBI) (Weirauch, 2007; Soto and Weirauch, 2009; and a unique number (00094810). USI codes Schuh and Pedraza, 2010). are included in locality data, figures, and The present study details the phylogenetic captions. To reduce the amount of space species-level relationships, classification, and occupied by the USIs in the material host-plant associations for a monophyletic examined sections the prefix (AMNH_PBI) group of Myrtaceae-feeding taxa that are is omitted except for the holotypes. part of a larger lineage within Phylini whose All latitude-longitude data presented in the members are restricted to Australia. The specimens examined sections are in degrees genesis of the paper began with dissections and decimal parts thereof; the vast majority of male genitalia of a comprehensive sample of georeferences were obtained with a GPS of Australian Phylinae (, 250 morphospe- device in the field. Altitude data are treated cies) by coauthor Weirauch during her tenure as metric. as a postdoctoral fellow funded by a NSF Comments on descriptions: The descrip- Planetary Biodiversity Inventories project for tions in the present paper were generated the study of Miridae. Those efforts allowed from a matrix of character data using us to recognize the existence of unique WinClada (Nixon, 2000) and then underwent

5 6 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 344

TABLE 1 Measurements of Melaleucoides genus group species

Length Width Body CunClyp Head Pron Scut Cun Head Pron Scut InterOc AntSeg2

Harpagophylus agnew M(N = 3) Mean 2.49 1.62 0.13 0.29 0.31 0.40 0.57 0.79 0.42 0.30 0.74 St Dev 0.11 0.08 0.03 0.05 0.02 0.03 0.01 0.02 0.01 0.01 0.11 Range 0.21 0.16 0.07 0.08 0.04 0.06 0.02 0.04 0.02 0.03 0.21 Minimum 2.36 1.54 0.10 0.24 0.29 0.37 0.56 0.77 0.40 0.28 0.62 Maximum 2.57 1.70 0.16 0.32 0.33 0.43 0.58 0.82 0.42 0.31 0.83 F(N = 3) Mean 2.41 1.61 0.17 0.31 0.30 0.36 0.54 0.80 0.40 0.30 0.90 St Dev 0.08 0.01 0.03 0.02 0.02 0.03 0.03 0.04 0.03 0.02 0.11 Range 0.17 0.02 0.06 0.03 0.05 0.06 0.06 0.08 0.06 0.03 0.21 Minimum 2.33 1.60 0.14 0.31 0.28 0.33 0.51 0.77 0.37 0.28 0.79 Maximum 2.50 1.62 0.20 0.33 0.32 0.39 0.57 0.85 0.43 0.31 1.00 Harpagophylus calytrix M(N = 6) Mean 2.51 1.70 0.14 0.36 0.32 0.39 0.62 0.91 0.46 0.33 0.75 St Dev 0.10 0.09 0.02 0.02 0.02 0.02 0.02 0.05 0.02 0.01 0.06 Range 0.23 0.23 0.07 0.05 0.06 0.06 0.04 0.14 0.06 0.03 0.17 Minimum 2.40 1.59 0.10 0.34 0.29 0.35 0.60 0.82 0.43 0.32 0.65 Maximum 2.63 1.82 0.16 0.39 0.35 0.41 0.64 0.96 0.48 0.35 0.82 F(N = 4) Mean 2.52 1.79 0.18 0.36 0.35 0.36 0.62 0.96 0.48 0.35 0.80 St Dev 0.07 0.05 0.02 0.02 0.01 0.02 0.02 0.04 0.01 0.01 0.02 Range 0.13 0.11 0.05 0.04 0.01 0.05 0.05 0.09 0.03 0.03 0.05 Minimum 2.45 1.73 0.15 0.34 0.34 0.33 0.60 0.91 0.47 0.33 0.77 Maximum 2.58 1.83 0.20 0.37 0.36 0.38 0.64 1.00 0.50 0.36 0.82 Harpagophylus scholtzii M(N = 4) Mean 2.17 1.47 0.15 0.30 0.28 0.35 0.51 0.75 0.36 0.29 0.66 St Dev 0.09 0.07 0.05 0.01 0.01 0.02 0.03 0.04 0.01 0.01 0.02 Range 0.20 0.14 0.10 0.03 0.01 0.05 0.07 0.09 0.02 0.03 0.04 Minimum 2.10 1.43 0.12 0.29 0.27 0.32 0.47 0.71 0.35 0.28 0.64 Maximum 2.30 1.57 0.22 0.32 0.29 0.37 0.54 0.80 0.37 0.31 0.68 F(N = 4) Mean 2.26 1.57 0.09 0.33 0.31 0.36 0.54 0.83 0.40 0.32 0.76 St Dev 0.16 0.13 0.03 0.03 0.03 0.02 0.01 0.03 0.02 0.02 0.05 Range 0.38 0.32 0.08 0.08 0.06 0.05 0.01 0.08 0.05 0.04 0.12 Minimum 2.06 1.40 0.06 0.30 0.27 0.34 0.53 0.78 0.37 0.29 0.71 Maximum 2.43 1.72 0.13 0.37 0.33 0.39 0.54 0.86 0.42 0.34 0.83 Harpagophylus thryptomeni M(N = 3) Mean 2.23 1.47 0.14 0.29 0.26 0.36 0.51 0.73 0.37 0.29 0.69 St Dev 0.12 0.06 0.03 0.01 0.01 0.01 0.01 0.03 0.02 0.01 0.04 Range 0.23 0.12 0.07 0.02 0.02 0.02 0.02 0.05 0.03 0.02 0.09 Minimum 2.11 1.41 0.10 0.28 0.25 0.36 0.50 0.70 0.35 0.28 0.64 Maximum 2.34 1.53 0.17 0.30 0.27 0.38 0.52 0.76 0.39 0.30 0.73 F(N = 5) Mean 2.36 1.60 0.18 0.30 0.28 0.38 0.54 0.77 0.37 0.31 0.75 St Dev 0.18 0.12 0.05 0.03 0.02 0.02 0.03 0.03 0.02 0.02 0.05 Range 0.48 0.29 0.14 0.08 0.04 0.05 0.07 0.06 0.05 0.05 0.12 Minimum 2.14 1.43 0.11 0.25 0.25 0.35 0.51 0.74 0.34 0.29 0.68 Maximum 2.61 1.72 0.25 0.33 0.29 0.40 0.58 0.80 0.39 0.34 0.80 Harpagophylus verticordii M(N = 5) Mean 2.43 1.63 0.18 0.32 0.30 0.38 0.56 0.81 0.38 0.32 0.76 St Dev 0.09 0.09 0.06 0.02 0.02 0.02 0.01 0.04 0.03 0.01 0.05 Range 0.20 0.24 0.16 0.06 0.04 0.07 0.04 0.09 0.07 0.04 0.12 Minimum 2.34 1.53 0.13 0.29 0.28 0.35 0.54 0.77 0.34 0.30 0.71 Maximum 2.54 1.77 0.29 0.34 0.32 0.41 0.58 0.86 0.42 0.34 0.83 2010 SCHUH AND WEIRAUCH: MYRTACEAE-FEEDING PHYLINAE 7

TABLE 1 (Continued)

Length Width Body CunClyp Head Pron Scut Cun Head Pron Scut InterOc AntSeg2 F(N = 5) Mean 2.54 1.72 0.17 0.35 0.31 0.40 0.58 0.86 0.43 0.33 0.86 St Dev 0.11 0.07 0.03 0.01 0.04 0.03 0.02 0.04 0.03 0.01 0.04 Range 0.31 0.19 0.06 0.02 0.10 0.07 0.07 0.12 0.07 0.03 0.11 Minimum 2.40 1.63 0.13 0.34 0.27 0.37 0.55 0.81 0.39 0.32 0.80 Maximum 2.70 1.83 0.19 0.36 0.36 0.44 0.62 0.93 0.46 0.35 0.91 Thryptomenomiris kalbarri M(N = 4) Mean 2.48 1.72 0.16 0.35 0.32 0.38 0.57 0.82 0.40 0.30 0.82 St Dev 0.05 0.03 0.03 0.01 0.03 0.06 0.02 0.03 0.02 0.01 0.02 Range 0.12 0.07 0.08 0.03 0.06 0.14 0.05 0.08 0.05 0.03 0.03 Minimum 2.44 1.69 0.12 0.33 0.30 0.30 0.55 0.77 0.37 0.28 0.80 Maximum 2.55 1.76 0.20 0.36 0.36 0.44 0.60 0.85 0.42 0.32 0.83 F(N = 4) Mean 2.66 1.81 0.21 0.34 0.31 0.42 0.59 0.84 0.42 0.33 0.97 St Dev 0.11 0.09 0.08 0.02 0.02 0.03 0.01 0.03 0.02 0.01 0.01 Range 0.27 0.21 0.19 0.06 0.04 0.05 0.03 0.06 0.05 0.02 0.03 Minimum 2.52 1.72 0.13 0.31 0.29 0.39 0.59 0.80 0.39 0.32 0.95 Maximum 2.79 1.93 0.32 0.37 0.33 0.44 0.61 0.86 0.44 0.33 0.98 Thryptomenomiris yalgoo M(N = 5) Mean 2.65 1.82 0.17 0.37 0.33 0.40 0.59 0.88 0.45 0.30 0.83 St Dev 0.08 0.05 0.02 0.03 0.02 0.03 0.01 0.02 0.04 0.00 0.01 Range 0.21 0.10 0.03 0.05 0.05 0.06 0.03 0.06 0.10 0.01 0.02 Minimum 2.58 1.77 0.15 0.36 0.31 0.37 0.57 0.86 0.40 0.30 0.83 Maximum 2.79 1.87 0.19 0.41 0.36 0.43 0.60 0.92 0.50 0.31 0.85 F(N = 5) Mean 2.77 1.91 0.20 0.39 0.34 0.40 0.61 0.90 0.45 0.33 0.95 St Dev 0.04 0.02 0.01 0.02 0.01 0.02 0.02 0.03 0.02 0.02 0.03 Range 0.10 0.04 0.03 0.05 0.03 0.05 0.04 0.07 0.05 0.04 0.08 Minimum 2.71 1.89 0.18 0.37 0.33 0.38 0.58 0.87 0.42 0.32 0.91 Maximum 2.82 1.94 0.21 0.42 0.36 0.43 0.62 0.93 0.47 0.35 0.99 Melaleucoides akaina M(N = 4) Mean 2.36 1.63 0.18 0.35 0.31 0.32 0.54 0.79 0.38 0.27 0.75 St Dev 0.04 0.03 0.04 0.01 0.00 0.03 0.01 0.01 0.01 0.00 0.02 Range 0.10 0.06 0.10 0.03 0.01 0.07 0.04 0.01 0.02 0.01 0.04 Minimum 2.33 1.61 0.14 0.33 0.30 0.29 0.52 0.78 0.38 0.27 0.73 Maximum 2.43 1.67 0.24 0.36 0.31 0.36 0.56 0.79 0.39 0.28 0.77 F(N = 4) Mean 2.61 1.79 0.18 0.37 0.32 0.40 0.55 0.85 0.41 0.31 0.90 St Dev 0.09 0.09 0.04 0.03 0.01 0.06 0.00 0.02 0.02 0.01 0.03 Range 0.17 0.18 0.09 0.07 0.03 0.12 0.01 0.04 0.06 0.02 0.05 Minimum 2.52 1.68 0.15 0.34 0.31 0.32 0.54 0.82 0.38 0.30 0.87 Maximum 2.68 1.87 0.24 0.40 0.33 0.44 0.56 0.86 0.44 0.32 0.93 Melaleucoides annae M(N = 5) Mean 3.46 2.40 0.19 0.45 0.49 0.48 0.80 1.29 0.66 0.38 0.80 St Dev 0.08 0.05 0.03 0.02 0.01 0.06 0.01 0.06 0.02 0.01 0.05 Range 0.21 0.11 0.07 0.06 0.04 0.14 0.03 0.14 0.05 0.03 0.13 Minimum 3.35 2.34 0.15 0.42 0.48 0.41 0.79 1.20 0.64 0.36 0.72 Maximum 3.56 2.46 0.22 0.48 0.52 0.56 0.82 1.34 0.69 0.39 0.85 F(N = 5) Mean 3.51 2.54 0.23 0.48 0.51 0.48 0.83 1.36 0.70 0.41 0.97 St Dev 0.14 0.06 0.02 0.03 0.03 0.08 0.03 0.01 0.02 0.04 0.16 Range 0.33 0.14 0.05 0.07 0.08 0.20 0.07 0.03 0.05 0.10 0.40 Minimum 3.33 2.47 0.20 0.44 0.46 0.35 0.79 1.33 0.69 0.37 0.68 Maximum 3.67 2.61 0.25 0.51 0.53 0.55 0.86 1.37 0.73 0.47 1.09 8 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 344

TABLE 1 (Continued)

Length Width Body CunClyp Head Pron Scut Cun Head Pron Scut InterOc AntSeg2

Melaleucoides beaufortiae M(N = 5) Mean 2.98 2.08 0.19 0.39 0.43 0.37 0.80 1.10 0.54 0.39 0.82 St Dev 0.09 0.09 0.06 0.05 0.03 0.03 0.04 0.03 0.02 0.03 0.07 Range 0.26 0.25 0.14 0.11 0.07 0.09 0.10 0.09 0.04 0.09 0.15 Minimum 2.85 1.94 0.12 0.34 0.40 0.32 0.75 1.05 0.51 0.34 0.75 Maximum 3.11 2.19 0.27 0.45 0.48 0.41 0.86 1.13 0.55 0.44 0.90 F(N = 3) Mean 3.06 2.14 0.22 0.40 0.44 0.41 0.82 1.13 0.56 0.42 0.92 St Dev 0.07 0.15 0.04 0.05 0.03 0.07 0.04 0.05 0.03 0.02 0.05 Range 0.16 0.38 0.09 0.11 0.06 0.18 0.10 0.14 0.07 0.04 0.13 Minimum 2.96 1.99 0.18 0.36 0.42 0.30 0.77 1.06 0.54 0.39 0.86 Maximum 3.13 2.37 0.27 0.47 0.48 0.48 0.87 1.20 0.61 0.43 1.00 Melaleucoides brevifoliae M(N = 5) Mean 3.41 2.42 0.20 0.46 0.46 0.48 0.87 1.29 0.65 0.44 0.97 St Dev 0.13 0.07 0.02 0.03 0.06 0.06 0.04 0.07 0.06 0.05 0.05 Range 0.32 0.20 0.05 0.09 0.15 0.14 0.09 0.17 0.17 0.11 0.12 Minimum 3.19 2.31 0.17 0.41 0.39 0.42 0.84 1.21 0.57 0.39 0.92 Maximum 3.52 2.51 0.22 0.49 0.54 0.56 0.93 1.38 0.74 0.50 1.04 F(N = 5) Mean 3.70 2.57 0.23 0.48 0.52 0.53 0.92 1.36 0.69 0.51 1.07 St Dev 0.13 0.11 0.05 0.05 0.05 0.03 0.05 0.03 0.01 0.02 0.06 Range 0.29 0.26 0.12 0.13 0.12 0.08 0.11 0.06 0.03 0.04 0.15 Minimum 3.58 2.46 0.16 0.40 0.48 0.51 0.86 1.32 0.67 0.50 1.02 Maximum 3.87 2.72 0.29 0.53 0.60 0.59 0.97 1.38 0.70 0.54 1.16 Melaleucoides cassisi M(N = 5) Mean 3.95 2.79 0.28 0.54 0.52 0.53 0.85 1.38 0.68 0.38 1.20 St Dev 0.11 0.10 0.03 0.02 0.03 0.06 0.03 0.04 0.04 0.02 0.09 Range 0.26 0.27 0.07 0.04 0.07 0.15 0.07 0.12 0.08 0.05 0.23 Minimum 3.78 2.63 0.25 0.52 0.48 0.43 0.82 1.32 0.65 0.35 1.04 Maximum 4.05 2.90 0.32 0.56 0.55 0.58 0.89 1.44 0.73 0.41 1.28 F(N = 5) Mean 4.12 2.96 0.25 0.56 0.54 0.54 0.90 1.53 0.76 0.45 1.22 St Dev 0.17 0.05 0.04 0.02 0.08 0.08 0.02 0.02 0.02 0.03 0.11 Range 0.39 0.13 0.11 0.06 0.20 0.21 0.05 0.05 0.06 0.08 0.27 Minimum 3.86 2.89 0.21 0.52 0.41 0.42 0.88 1.50 0.73 0.41 1.05 Maximum 4.26 3.02 0.32 0.58 0.61 0.63 0.93 1.55 0.79 0.49 1.33 Melaleucoides castanea M(N =1) Mean 3.61 2.39 0.17 0.47 0.54 0.62 1.03 1.44 0.75 0.58 1.00 F(N = 3) Mean 3.84 2.69 0.22 0.52 0.59 0.53 1.12 1.55 0.82 0.60 0.92 St Dev 0.07 0.08 0.05 0.02 0.01 0.07 0.05 0.04 0.02 0.01 0.05 Range 0.12 0.15 0.10 0.03 0.01 0.14 0.11 0.08 0.04 0.02 0.11 Minimum 3.80 2.60 0.18 0.51 0.58 0.46 1.06 1.51 0.80 0.59 0.86 Maximum 3.92 2.75 0.28 0.54 0.60 0.59 1.17 1.59 0.84 0.61 0.97 Melaleucoides grossi M(N = 4) Mean 3.34 2.46 0.17 0.43 0.51 0.50 0.92 1.38 0.70 0.41 0.94 St Dev 0.21 0.19 0.06 0.11 0.03 0.07 0.02 0.03 0.03 0.02 0.13 Range 0.40 0.45 0.15 0.26 0.08 0.14 0.05 0.05 0.06 0.04 0.28 Minimum 3.13 2.22 0.08 0.32 0.48 0.43 0.90 1.35 0.66 0.40 0.78 Maximum 3.52 2.67 0.24 0.58 0.55 0.57 0.96 1.40 0.72 0.43 1.06 F(N = 4) Mean 3.75 2.74 0.24 0.54 0.52 0.54 0.92 1.46 0.72 0.45 1.03 St Dev 0.10 0.10 0.08 0.08 0.07 0.07 0.03 0.04 0.02 0.04 0.03 Range 0.21 0.23 0.18 0.18 0.16 0.14 0.07 0.09 0.04 0.10 0.07 Minimum 3.63 2.63 0.14 0.43 0.43 0.49 0.89 1.42 0.70 0.39 1.00 Maximum 3.84 2.85 0.32 0.61 0.59 0.63 0.96 1.51 0.74 0.49 1.08 2010 SCHUH AND WEIRAUCH: MYRTACEAE-FEEDING PHYLINAE 9

TABLE 1 (Continued)

Length Width Body CunClyp Head Pron Scut Cun Head Pron Scut InterOc AntSeg2

Melaleucoides leuropomae M(N = 5) Mean 3.25 2.29 0.21 0.45 0.46 0.41 0.89 1.25 0.60 0.46 0.85 St Dev 0.10 0.06 0.02 0.03 0.02 0.04 0.03 0.03 0.02 0.02 0.05 Range 0.23 0.15 0.06 0.07 0.06 0.08 0.08 0.09 0.06 0.06 0.14 Minimum 3.11 2.22 0.19 0.42 0.42 0.37 0.85 1.20 0.57 0.43 0.79 Maximum 3.34 2.36 0.25 0.48 0.48 0.45 0.92 1.29 0.63 0.49 0.93 F(N = 5) Mean 3.46 2.43 0.19 0.48 0.52 0.47 0.93 1.32 0.65 0.51 0.93 St Dev 0.09 0.08 0.02 0.02 0.03 0.03 0.06 0.03 0.04 0.04 0.05 Range 0.25 0.21 0.06 0.06 0.07 0.07 0.15 0.08 0.10 0.08 0.13 Minimum 3.35 2.34 0.15 0.44 0.50 0.44 0.85 1.30 0.60 0.47 0.88 Maximum 3.60 2.55 0.21 0.50 0.57 0.50 1.01 1.38 0.70 0.54 1.01 Melaleucoides micranthae M(N = 5) Mean 3.14 2.24 0.21 0.39 0.46 0.44 0.82 1.22 0.63 0.42 0.91 St Dev 0.14 0.09 0.03 0.02 0.03 0.02 0.02 0.04 0.05 0.04 0.10 Range 0.37 0.24 0.09 0.06 0.07 0.05 0.04 0.10 0.12 0.09 0.25 Minimum 2.94 2.14 0.16 0.36 0.43 0.43 0.80 1.18 0.55 0.37 0.78 Maximum 3.31 2.37 0.25 0.42 0.51 0.47 0.84 1.28 0.67 0.46 1.03 F(N = 5) Mean 3.53 2.45 0.21 0.46 0.50 0.48 0.94 1.16 0.56 0.50 1.12 St Dev 0.15 0.17 0.04 0.05 0.02 0.04 0.10 0.40 0.25 0.09 0.12 Range 0.33 0.39 0.11 0.14 0.04 0.12 0.25 0.93 0.59 0.23 0.28 Minimum 3.38 2.25 0.16 0.38 0.47 0.41 0.86 0.45 0.12 0.44 1.00 Maximum 3.71 2.64 0.27 0.53 0.52 0.53 1.12 1.38 0.71 0.67 1.27 Melaleucoides ozzii M(N = 5) Mean 2.90 1.93 0.20 0.33 0.36 0.45 0.64 1.02 0.49 0.39 0.94 St Dev 0.10 0.08 0.03 0.03 0.02 0.04 0.03 0.03 0.02 0.05 0.03 Range 0.27 0.20 0.07 0.09 0.05 0.09 0.07 0.06 0.06 0.14 0.07 Minimum 2.74 1.86 0.17 0.30 0.34 0.41 0.61 1.00 0.46 0.30 0.91 Maximum 3.01 2.06 0.24 0.38 0.38 0.50 0.67 1.06 0.52 0.44 0.98 F(N = 5) Mean 3.04 2.04 0.21 0.31 0.43 0.45 0.65 1.04 0.54 0.42 0.79 St Dev 0.12 0.07 0.05 0.02 0.06 0.05 0.01 0.06 0.05 0.02 0.18 Range 0.30 0.17 0.12 0.04 0.15 0.13 0.03 0.17 0.13 0.04 0.43 Minimum 2.90 1.95 0.15 0.29 0.36 0.41 0.64 0.93 0.47 0.40 0.57 Maximum 3.20 2.12 0.27 0.33 0.51 0.54 0.66 1.10 0.60 0.44 1.00 Melaleucoides pileanthicola M(N = 5) Mean 2.72 1.90 0.20 0.41 0.35 0.38 0.62 0.97 0.48 0.33 0.90 St Dev 0.09 0.07 0.03 0.02 0.01 0.04 0.01 0.05 0.01 0.02 0.05 Range 0.21 0.16 0.08 0.04 0.03 0.10 0.04 0.15 0.04 0.04 0.12 Minimum 2.61 1.79 0.16 0.39 0.33 0.34 0.60 0.89 0.46 0.31 0.86 Maximum 2.82 1.95 0.24 0.43 0.37 0.44 0.63 1.04 0.50 0.35 0.98 F(N = 5) Mean 2.94 2.03 0.20 0.43 0.37 0.41 0.63 1.01 0.50 0.35 0.97 St Dev 0.07 0.05 0.04 0.01 0.01 0.04 0.01 0.02 0.01 0.01 0.04 Range 0.19 0.12 0.10 0.03 0.03 0.08 0.02 0.05 0.02 0.02 0.11 Minimum 2.87 1.97 0.16 0.42 0.36 0.38 0.62 0.99 0.49 0.34 0.91 Maximum 3.07 2.09 0.26 0.45 0.39 0.46 0.64 1.05 0.51 0.37 1.03 Melaleucoides rhaphiophyllae M(N = 3) Mean 2.87 2.13 0.17 0.40 0.46 0.40 0.84 1.22 0.61 0.44 0.99 St Dev 0.16 0.04 0.06 0.05 0.02 0.04 0.03 0.06 0.01 0.03 0.06 Range 0.31 0.07 0.12 0.10 0.03 0.07 0.06 0.12 0.02 0.07 0.11 Minimum 2.69 2.11 0.11 0.35 0.45 0.36 0.81 1.17 0.60 0.41 0.95 Maximum 3.00 2.18 0.23 0.45 0.48 0.43 0.87 1.29 0.61 0.48 1.06 10 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 344

TABLE 1 (Continued)

Length Width Body CunClyp Head Pron Scut Cun Head Pron Scut InterOc AntSeg2 F(N = 2) Mean 2.98 1.96 0.12 0.34 0.43 0.51 0.83 1.20 0.63 0.48 1.05 St Dev 0.15 0.12 0.02 0.04 0.04 0.02 0.03 0.05 0.03 0.04 0.07 Range 0.21 0.17 0.03 0.06 0.05 0.03 0.05 0.07 0.04 0.05 0.09 Minimum 2.87 1.87 0.10 0.31 0.40 0.50 0.80 1.17 0.60 0.45 1.00 Maximum 3.08 2.04 0.13 0.37 0.45 0.53 0.85 1.24 0.65 0.50 1.10 Melaleucoides sheathianae M(N = 5) Mean 3.67 2.59 0.20 0.49 0.55 0.52 0.87 1.37 0.68 0.39 1.01 St Dev 0.15 0.16 0.05 0.05 0.01 0.05 0.03 0.04 0.04 0.03 0.10 Range 0.31 0.35 0.12 0.12 0.03 0.14 0.06 0.11 0.09 0.07 0.27 Minimum 3.50 2.39 0.14 0.43 0.53 0.45 0.85 1.31 0.64 0.35 0.89 Maximum 3.80 2.74 0.27 0.55 0.56 0.59 0.90 1.42 0.73 0.42 1.16 F(N = 5) Mean 3.82 2.73 0.22 0.51 0.57 0.50 0.89 1.43 0.72 0.43 1.11 St Dev 0.28 0.09 0.03 0.04 0.03 0.09 0.03 0.09 0.03 0.02 0.05 Range 0.71 0.24 0.07 0.09 0.08 0.22 0.07 0.23 0.09 0.06 0.11 Minimum 3.49 2.63 0.18 0.48 0.53 0.39 0.86 1.31 0.68 0.40 1.05 Maximum 4.21 2.86 0.25 0.57 0.62 0.61 0.94 1.53 0.77 0.46 1.16 Melaleucoides similis M(N = 4) Mean 3.38 2.24 0.23 0.34 0.40 0.59 0.69 1.11 0.53 0.35 0.88 St Dev 0.16 0.12 0.06 0.03 0.03 0.04 0.02 0.05 0.02 0.09 0.21 Range 0.39 0.26 0.14 0.07 0.08 0.09 0.05 0.11 0.05 0.20 0.50 Minimum 3.19 2.16 0.15 0.30 0.36 0.54 0.66 1.06 0.51 0.21 0.67 Maximum 3.59 2.42 0.29 0.37 0.44 0.63 0.71 1.17 0.55 0.41 1.17 F(N = 1) Mean 3.11 2.10 0.13 0.37 0.43 0.46 0.66 1.08 0.52 0.38 0.94 Melaleucoides systenae M(N = 5) Mean 3.23 2.34 0.24 0.47 0.48 0.42 0.84 1.22 0.70 0.44 0.84 St Dev 0.21 0.11 0.06 0.04 0.04 0.08 0.04 0.06 0.06 0.01 0.05 Range 0.52 0.24 0.16 0.09 0.11 0.20 0.10 0.13 0.16 0.02 0.13 Minimum 2.93 2.20 0.13 0.44 0.44 0.30 0.79 1.17 0.63 0.43 0.77 Maximum 3.45 2.44 0.29 0.53 0.55 0.49 0.89 1.29 0.79 0.45 0.90 F(N = 5) Mean 3.51 2.48 0.25 0.47 0.47 0.47 0.80 1.32 0.71 0.44 0.85 St Dev 0.07 0.06 0.04 0.02 0.05 0.03 0.08 0.05 0.09 0.04 0.09 Range 0.18 0.17 0.10 0.06 0.12 0.07 0.16 0.15 0.22 0.09 0.24 Minimum 3.41 2.40 0.22 0.45 0.43 0.43 0.73 1.24 0.61 0.39 0.73 Maximum 3.59 2.57 0.31 0.51 0.55 0.50 0.90 1.39 0.84 0.48 0.97 Melaleucoides uncinatae M(N = 5) Mean 2.99 2.25 0.19 0.41 0.52 0.35 0.76 1.25 0.69 0.40 1.07 St Dev 0.14 0.05 0.03 0.02 0.01 0.03 0.02 0.01 0.05 0.01 0.12 Range 0.34 0.13 0.07 0.05 0.02 0.09 0.04 0.03 0.13 0.03 0.27 Minimum 2.89 2.17 0.15 0.39 0.51 0.29 0.74 1.24 0.63 0.38 0.94 Maximum 3.23 2.29 0.22 0.44 0.53 0.38 0.78 1.27 0.76 0.42 1.21 F(N = 5) Mean 3.26 2.39 0.21 0.46 0.53 0.39 0.80 1.34 0.70 0.45 1.12 St Dev 0.09 0.05 0.04 0.04 0.02 0.05 0.03 0.03 0.10 0.04 0.10 Range 0.22 0.10 0.11 0.07 0.06 0.11 0.08 0.06 0.27 0.10 0.25 Minimum 3.15 2.34 0.15 0.42 0.49 0.35 0.75 1.32 0.54 0.40 1.00 Maximum 3.37 2.44 0.26 0.49 0.55 0.46 0.83 1.38 0.81 0.50 1.25 Melaleucoides undulatae M(N = 5) Mean 3.03 2.22 0.20 0.41 0.43 0.36 0.77 1.17 0.55 0.39 1.04 St Dev 0.16 0.10 0.03 0.03 0.04 0.07 0.02 0.04 0.03 0.02 0.07 Range 0.40 0.23 0.07 0.07 0.08 0.20 0.05 0.10 0.08 0.06 0.16 Minimum 2.81 2.11 0.17 0.39 0.39 0.24 0.73 1.13 0.50 0.36 0.96 Maximum 3.21 2.34 0.24 0.46 0.47 0.44 0.78 1.23 0.58 0.42 1.12 2010 SCHUH AND WEIRAUCH: MYRTACEAE-FEEDING PHYLINAE 11

TABLE 1 (Continued)

Length Width Body CunClyp Head Pron Scut Cun Head Pron Scut InterOc AntSeg2 F(N = 5) Mean 3.27 2.26 0.19 0.42 0.48 0.50 0.80 1.25 0.61 0.46 1.09 St Dev 0.07 0.07 0.04 0.04 0.04 0.02 0.01 0.03 0.05 0.03 0.05 Range 0.18 0.15 0.10 0.09 0.09 0.06 0.03 0.06 0.15 0.07 0.14 Minimum 3.19 2.18 0.14 0.39 0.44 0.47 0.79 1.23 0.55 0.43 1.02 Maximum 3.37 2.33 0.24 0.48 0.54 0.52 0.82 1.29 0.70 0.50 1.16 Melaleucoides verticordiae M(N = 5) Mean 3.49 2.39 0.26 0.43 0.48 0.51 0.74 1.19 0.72 0.39 1.13 St Dev 0.22 0.21 0.01 0.05 0.05 0.07 0.05 0.10 0.11 0.02 0.19 Range 0.59 0.46 0.03 0.14 0.12 0.18 0.11 0.22 0.27 0.06 0.46 Minimum 3.17 2.10 0.25 0.36 0.42 0.41 0.69 1.06 0.55 0.37 0.84 Maximum 3.76 2.56 0.28 0.50 0.54 0.59 0.80 1.28 0.82 0.42 1.30 F(N = 5) Mean 3.59 2.53 0.30 0.43 0.46 0.48 0.75 1.24 0.63 0.46 1.28 St Dev 0.37 0.28 0.06 0.07 0.06 0.08 0.05 0.10 0.04 0.04 0.23 Range 0.79 0.54 0.17 0.16 0.15 0.21 0.12 0.23 0.10 0.10 0.55 Minimum 3.26 2.31 0.22 0.36 0.37 0.40 0.68 1.12 0.57 0.41 1.07 Maximum 4.05 2.85 0.39 0.52 0.52 0.61 0.81 1.35 0.68 0.51 1.62 additional editing. Where size serves to help meter output being written directly to a diagnose a taxon, we have included relevant spreadsheet. measurement data in the diagnoses and Designation of paratypes: We designate as descriptions. Detailed measurements for all paratypes specimens that represent a confi- species are presented in table 1. dent association of material with the concept Host-plant identifications: Hosts recorded for the species and which were examined as in this study were pressed in the field and part of the descriptive process. submitted as vouchers to the Western Aus- Deposition of specimens and abbreviations tralian Herbarium or the Royal Botanic for institutional depositories: Within Australia Gardens, Sydney, for identification by bo- the collecting permitting process is reserved tanical specialists. Thus, the identifications to the states. Following the stipulations of are authoritative and independent of our own our permits (see Acknowledgments) holo- predilections about the identities of the plant types are deposited in the states of origin. groups on which we were collecting. Host Depending on the conditions of the collecting voucher numbers and/or locations are listed permits, a subset of paratypes may also be in the specimens examined sections. deposited in the state or origin. The prepon- Terminology for female genitalia:Wehave derance of the remaining material is lodged in used the terminology interramal sclerites and the institutions that supported the fieldwork interramal lobes for structures of the posteri- (Australian Museum, American Museum of or wall in the female (figs. 6, 7, 11, 18, 25, Natural History), with a representative sam- 34), following the practice established for ple of specimens deposited in institutions Orthotylinae by Schaffner and Schwartz participating in the Plant Bug Planetary (2008) and Forero (2008). Our use of both Biodiversity Inventory Project or other insti- of these terms in Phylinae should be con- tutions of record. strued as descriptive in nature, because these Institutional depositories for specimens structures may not be homologous with those utilized during the course of this study and in Orthotylinae. their acronyms as used in this paper are as Measurements: Measurements for all taxa follows: are presented in table 1. Means for total length and pronotal width are presented in AM Australian Museum, Sydney the descriptions. Measurements were made AMNH American Museum of Natural using a micrometer driven stage, with micro- History, New York 12 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 344

ANIC Australian National Insect Col- paramere not greatly elongate in dorsoventral lection, CSIRO, Canberra perspective, covering about half of height of CNC Canadian National Collection of pygophore in lateral view (fig. 5D), with Insects, Ottawa posterior process very long and curving SAMA South Australian Museum, Ade- ventrally (figs. 1–3); endosoma always with 2 appendages in region of secondary gono- laide pore...... Harpagophylus UCR University of California, River- – Right paramere short, broadly lanceolate, not side traversing entire width of pygophore in UNSW University of New South Wales, repose (figs. 8F, 9, 12F, 17F, 19); posterior Sydney process of left paramere relatively short, USNM United States National Museum never strongly curving toward venter . . . 2 of Natural History, Smithsonian 2. Left paramere elongate in dorsoventral per- Institution, Washington, DC spective, flattened, spoon shaped, and erect WAMP Western Australian Museum, (figs. 12D, E, 13, 17E, 19, 23, 24E); apex of left paramere sometimes elongate ...... ZISP Zoological Institute, Russian ...... Melaleucoides Academy of Sciences, St. Peters- – Left paramere more quadrate, not erect and spoon shaped as above, always with apex burg elongate (figs. 7, 8E, F, 9), reaching across pygophore in repose (fig. 8F) ...... Thryptomenomiris Melaleucoides genus group

DIAGNOSIS: Recognized by fleshy lyriform Harpagophylus, new genus parempodia (fig. 5C; character 23-1) and Figures 1–6, 35–37; map 1; plate 1 clawlike decurved apex of the right paramere TYPE SPECIES: Harpagophylus verticordii, (fig. 1; 61-0); other attributes found in the new species. group include the dark spots at the bases of DIAGNOSIS: Recognized within the Mela- the tibial spines and the female genitalia with leucoides genus group by the unique, greatly interramal sclerites and interramal lobes elongate right paramere with a bulbous base (fig. 6). The group is also diagnosed by the (60-2) and the long, decurved posterior weakly arcuate and dorsally curving primary process of the left paramere (58-3) (figs. 1– endosomal strap of the male genitalia (fig. 1; 6); size small, total length 2.10–2.63. Poten- character 30-1) and the presence of a tially confused with Thryptomenomiris, and spinelike elongate process near the gonopore small species of Melaleucoides on the basis of on the ventral surface of the endosoma that is small size and association with the Myrta- recurved toward the base of the endosoma ceae. The structure of the parameres in (fig. 1; 37-2), although these attributes do not occur in all species. Harpagophylus precludes any confusion with these other groups, however. DISCUSSION: Lyriform parempodia occur in all species in the Melaleucoides genus DESCRIPTION: Male: Body weakly elon- group, whereas other Phylini in Australia gate, weakly ovoid; small, total length 2.10– and elsewhere possess setiform parempodia. 2.63, width pronotum 0.70–0.96. COLORA- Most species in the group possess the TION (pl. 1): Pale, with or without reddish clawlike apex of the right paramere, but areas on dorsum. SURFACE AND VESTI- based on our observations and cladistic TURE (fig. 5B): Dorsum with reclining analyses, this character is secondarily modi- simple setae matching background coloration fied in several species within the group. and with some sericeous or woolly setae. STRUCTURE: Head (fig. 5A): Somewhat projecting; interocular space relatively large; KEY TO MELALEUCOIDES GENUS eyes leaving gena moderately exposed in GROUP GENERA lateral view. Thorax: Pretarsus (fig. 5C): 1. Right paramere greatly elongate, reaching Parempodia fleshy, recurved, lyriform; pul- across entire width of pygophore in repose villi absent. GENITALIA (figs. 1–6): Endo- (fig. 5F), with apex in the shape of a claw; left soma: Base short, curving, C- or J-shaped; 2010 SCHUH AND WEIRAUCH: MYRTACEAE-FEEDING PHYLINAE 13 body without torsion, primary strap ventral to secondary gonopore, apically greatly elongate, nearly straight, angled relative to body of endosoma, without ornamentation; spinelike, elongate process present near gonopore on dorsal surface, erect, unorna- mented; spinelike, elongate process present near gonopore on ventral surface, recurved toward base of endosoma; secondary gono- pore seen laterally in lateral view of endo- soma. Phallotheca: Nearly erect, apical region of variable structure. Left paramere (fig. 5E): Horizontal; shaft at right angles to body; in dorsal perspective just exceeding margin of pygophore; in lateral perspective covering limited extent of lateral surface of pygophore; body narrowed toward apex; apex medially angulate, posterolaterally simple; anterior process in lateral view varying from triangu- lar to undeveloped; posterior process very long, curving 90u toward base of paramere; base of posterior process without conspicu- ous shoulder. Right Paramere (fig. 5F): Body greatly elongate, bulbous at base; apex short, Map 1. Localities for species of Harpagophylus. clawlike, decurved. Female: Slightly more ovoid than male; new genus. The structure of the left and right small, total length 2.06–2.70, width prono- parameres clearly distinguishes them from tum 0.74–1.00. COLORATION (pl. 1): As in other members in the Melaleucoides genus male. GENITALIA (fig. 6): Posterior wall group. laterally with distinct crescent-shaped inter- ramal sclerites; posterolaterally with a dis- KEYTOSPECIESOFHARPAGOPHYLUS tinct swelling covered with microtrichia; longitudinal fold on either side of midline 1. Dorsum with conspicuous red areas, particu- along part of length; interramal lobes pres- larly posterior half of endocorium (pl. 1); apex ent, roughly symmetrical, erect, and not of phallotheca complex, body of endosoma ornamented with spicules or with very few; broad (fig. 2)...... Harpagophylus calytrix vestibulum with medial plates weakly sclero- – Dorsum lacking conspicuous red areas, most- tized or apparently absent, with caplike ly yellow (pl. 1); apex of phallotheca without such complex ornamentation and body of sclerotized guide with internal differentiation. endosoma relatively slender ...... 2 ETYMOLOGY: Named for the hooklike 2. Left side of pygophoral opening bearing a form of the right paramere; from the Latin distinct clawlike appendage (fig. 4) . . . . . harpago, ‘‘grappling hook,’’ in combination ...... Harpagophylus thryptomeni with Phylus; gender masculine. – Left side of pygophoral opening without a DISCUSSION: The five new species de- distinct clawlike appendage ...... 3 scribed below as belonging to the genus 3. Endosoma with both processes in region of Harpagophylus are well supported as a secondary gonopore very long and recurved monophyletic group in both the equal toward its base (fig. 6); apex of right paramere verylong...... Harpagophylus verticordii weights and the implied weights analyses – Endosoma with at least one process near (figs. 35, 36). Based on overall appearance secondary gonopore erect, perpendicular to and their association with Myrtaceae in the axis (figs. 1, 3); apex of right paramere not so tribe Chamelaucieae, they are easily confused long...... 4 with species of Thryptomenomiris, new genus, 4. Dorsal process of endosoma near secondary and the small species within Melaleucoides, gonopore longer than ventral process (fig. 1); 14 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 344

Fig. 1. Harpagophylus agnew, male genitalia (AMNH_PBI 00373245).

ventral process with a serrate margin DESCRIPTION: Male: Body weakly elon- (fig.1)...... Harpagophylus agnew gate, weakly ovoid; small, mean total length – Dorsal process of endosoma near secondary 2.49, mean width pronotum 0.79. COLOR- gonopore shorter than ventral process ATION (pl. 1): Head: Uniformly pale, yel- (fig. 1); ventral process with a simple margin low; scapus and pedicellus unicolorous pale; (fig.3)...... Harpagophylus scholtzii labium pale with segment 4 heavily infuscate. Harpagophylus agnew, new species Thorax: Pronotum scutellum, and hemely- Figure 1, map 1, plate 1 tron unicolorous pale, yellow; markings on cuneus absent; membrane very weakly fu- DIAGNOSIS: Recognized by the pale, yel- mose, veins; hind femur unicolorous pale, lowish coloration, the endosomal processes without black spots; hind tibial spines dark, both long, straight, and erect, the simple apex without dark spots at bases. Abdomen: of the phallotheca, the long, relatively slender Venter unicolorous pale or mostly so. SUR- right paramere, and the denticulate anterior FACE AND VESTITURE: Dorsum with process on the left paramere. Most easily reclining simple setae matching background confused with H. scholtzii, H. thryptomeni, coloration and some sericeous or woolly and H. verticordii on the basis of pale setae. STRUCTURE: Head: Barely project- coloration, but those species all differing in ing; interocular space moderate; eyes leaving the structure of the anterior process of the gena only very slightly exposed in lateral left paramere, the conformation of the right view. Antenna: Segment 2 of equal diameter paramere, the structure of the apex of the over entire length; antennal fossa with ventral phallotheca, and the conformation of the margin at ventral margin of eye. Labium: dorsal and ventral endosomal processes. Reaching to about anterior margin of pygo- 2010 SCHUH AND WEIRAUCH: MYRTACEAE-FEEDING PHYLINAE 15

Fig. 2. Harpagophylus calytrix, male genitalia (AMNH_PBI 00134821). phore. GENITALIA (fig. 1): Phallotheca: PARATYPES: AUSTRALIA: Western Aus- More or less right angulate; dorsal surface tralia: 31.7 km W of Agnew toward Sand- without a fingerlike projection, and without a stone, 27.96227uS 120.4277uE, 800 m, 26 Oct platelike projection; anterior surface without 1996, Schuh and Cassis, aspera a keel; ventral surface without a projecting glabra E. Pritz. (Myrtaceae), det. Perth keel; apex simple. 05095190, 3- (00373242–00373244), 5U Female: Slightly more ovoid than male; (00373246–00373250) (AM), 3- (00136499, small, mean total length 2.41, mean width 00136501, 00373245), 4U (00136502, 00136504, pronotum 0.80. COLORATION (pl. 1): As 00136506–00136507) (AMNH), 4U (00373251– in male. GENITALIA: Not examined. 00373254) (WAMP). ETYMOLOGY: Named for the town of Agnew, Western Australia, near the type locality; a noun in apposition. Harpagophylus calytrix, new species HOSTS (table 2): Thryptomene aspera gla- Figure 2, map 1, plate 1 bra E. Pritz. (Myrtaceae). DISTRIBUTION (map 1): Known only from DIAGNOSIS: Recognized by the heavily the type locality in the Goldfields region of reddish coloration on much of the dorsum, Western Australia. the absence of a developed anterior process HOLOTYPE: AUSTRALIA: Western Aus- on the left paramere, the complex structure of tralia: 31.7 km W of Agnew toward Sandstone, apex of phallotheca, and the broad body of 27.96227uS 120.4277uE, 800 m, 26 Oct 1996, the endosoma. All other species of Harpago- Schuh and Cassis, Thryptomene aspera glabra phylus are pale in coloration, with differing E. Pritz. (Myrtaceae), PERTH 05095190, 1- structures on the apex of the phallotheca, and (AMNH_PBI 00136500) (WAMP). with the body of the endosoma not so robust. 16 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 344

Fig. 3. Harpagophylus scholtzii, male genitalia (AMNH_PBI 00327137).

DESCRIPTION: Male: Body moderately ETYMOLOGY: Named for the host genus, elongate, parallel sided; small, mean total Calytrix (Myrtaceae); a noun in apposition. length 2.51, mean width pronotum 0.91. HOSTS: Calytrix variabilis Lindl. (Myrta- COLORATION (pl. 1): Head: Uniformly ceae) (pl. 5A). dark, reddish; scapus unicolorous dark; ped- DISTRIBUTION (map 1): Known only from icellus black at extreme base, remainder pale; the type locality in Western Australia. labium generally infuscate, heavily so apical- HOLOTYPE: AUSTRALIA: Western Aus- ly. Thorax: Pronotum and scutellum unicol- tralia: , Meanarra orous red to brown; hemelytron partially pale Hill, 27.69228uS 114.2104uE, 200 m, 29 Oct with red or carmine areas, especially on clavus 1996, Schuh and Cassis, Calytrix variabilis along scutellum and on endocorium; cuneus Lindl. (Myrtaceae), det. Perth 05120470, 1- transparent, without markings; membrane (AMNH_PBI 0134818) (WAMP). and veins strongly fumose; hind femur nearly PARATYPES: AUSTRALIA: Western Aus- unicolorous dark; hind tibial spines dark, tralia: Kalbarri National Park, Meanarra without dark spots at bases. Abdomen: Venter Hill, 27.69228uS 114.2104uE, 200 m, 29 Oct unicolorous dark. SURFACE AND VESTI- 1996, Schuh and Cassis, Calytrix variabilis Lindl. (Myrtaceae), det. Perth 05120470, 2- TURE: Dorsum with suberect or reclining U black setae and some sericeous or woolly (00134820, 00134821), 1 (00134826) (AM), 2- (00134822, 00134823), 2U (00134824, setae. STRUCTURE: Head: Somewhat pro- - U jecting; interocular space relatively large; eyes 00134825) (AMNH), 1 (00134819), 1 leaving gena only very slightly exposed in (00134827) (WAMP). lateral view. Antenna: Segment 2 of equal diameter over entire length; antennal fossa Harpagophylus scholtzii, new species with ventral margin at ventral margin of eye. Figure 3, map 1, plate 1 Labium: Reaching to about anterior margin DIAGNOSIS: Recognized by the pale, yellow- of pygophore. GENITALIA (fig. 2): Phal- ish coloration, the endosomal processes both lotheca: Nearly erect; apex complex. straight and erect, the dorsal process short, the Female: Slightly more ovoid than male; ventral process much longer, the relatively short small, mean total length 2.51, mean width right paramere, and the triangular anterior pronotum 0.96. COLORATION (pl. 1): As process on the left paramere. Most easily in male. GENITALIA: Not examined. confused with H. agnew, H. thryptomeni, and 2010 SCHUH AND WEIRAUCH: MYRTACEAE-FEEDING PHYLINAE 17

H. verticordii on the basis of the pale color- Cassis, Wall, Weirauch, Tatarnic, Symonds, ation, but those species differing from H. leptantha Benth. (Myrtaceae), det. scholtzii in the structure and conformation of Perth 6988687, 1- (00327135), 1U (0032- the dorsal and ventral endosomal processes. 7140) (AM), 2- (00327136, 00327137), 2U DESCRIPTION: Male: Body weakly elon- (00327138, 00327139) (AMNH), 1U (0032- gate, weakly ovoid; small, mean total length 7141) (WAMP). 2.17, mean width pronotum 0.75. COLOR- ATION (pl. 1): Head: Uniformly pale yellow; Harpagophylus thryptomeni, new species scapus and pedicellus unicolorous pale; Figure 4, map 1, plate 1 labium pale with segment 4 heavily infuscate. Thorax: Pronotum, scutellum, and hemely- DIAGNOSIS: Recognized by the pale yel- tron unicolorous pale yellow; markings on lowish coloration, and uniquely by the cuneus absent; membrane and veins weakly clawlike appendage on the left side of fumose; hind femur unicolorous pale, with- pygophoral opening (fig. 4) and the project- out black spots; hind tibial spines dark, ing hook on the apex of the phallotheca. without dark spots at bases. Abdomen: Most easily confused with H. agnew, H. Venter unicolorous pale or mostly so. SUR- scholtzii,andH. verticordii on the basis of FACE AND VESTITURE: Dorsum with pale coloration, but those species all lacking reclining simple setae matching background the hook on the pygophoral opening and also coloration and some sericeous or woolly showing differences in the structure of the setae. STRUCTURE: Head: Weakly project- anterior process of the left paramere, the ing; interocular space relatively large; eyes conformation of the right paramere, and the leaving gena only very slightly exposed in conformation of the dorsal and ventral lateral view. Antenna: Segment 2 of equal endosomal processes. diameter over entire length; antennal fossa DESCRIPTION: Male: Body weakly elon- with ventral margin at ventral margin of eye. gate, weakly ovoid; small, mean total length Labium: Reaching to about anterior margin 2.23, mean width pronotum 0.73. COLOR- of pygophore. GENITALIA (fig. 3): Phal- ATION (pl. 1): Head: Uniformly yellow- lotheca: Nearly erect; dorsal surface without orange; scapus and pedicellus unicolorous a fingerlike projection; apex complex. pale; labium pale with segment 4 heavily Female: Slightly more ovoid than male; infuscate. Thorax: Pronotum, scutellum, and small, mean total length 2.26, mean width hemelytron unicolorous yellow-orange; pronotum 0.83. COLORATION (pl. 1): As markings on cuneus absent; membrane and in male. GENITALIA: Not examined. veins weakly fumose; hind femur unicolorous ETYMOLOGY: Named for the host genus, pale, without black spots; hind tibial spines Scholtzia (Myrtaceae); a noun in apposition. dark, without dark spots at bases. Abdomen: HOSTS: Scholtzia leptantha Benth. (Myrta- Venter unicolorous pale or mostly so. SUR- ceae). FACE AND VESTITURE: Dorsum with DISTRIBUTION (map 1): Known from one reclining simple setae matching background locality in the Carnarvon region of Western coloration and some sericeous or woolly Australia. setae. STRUCTURE: Head: Weakly project- HOLOTYPE: AUSTRALIA: Western Aus- ing; interocular space moderate; eyes leaving tralia: 4.5 km NW of jct of Blowholes Rd and gena moderately exposed in lateral view. North West Coastal Hiway, N of Carnarvon, Antenna: Segment 2 of equal diameter over 24.72267uS 113.7158uE, 28 m, 27 Oct 2004, entire length; antennal fossa with ventral Cassis, Wall, Weirauch, Tatarnic, Symonds, margin at ventral margin of eye. Labium: Just Scholtzia leptantha Benth. (Myrtaceae), det. reaching onto abdomen. GENITALIA Perth6988687, 1- (AMNH_PBI 00327134) (fig. 4): Phallotheca: Nearly erect; dorsal (WAMP). surface without ornamentation; anterior and PARATYPES: AUSTRALIA: Western Aus- ventral surfaces without a keel; apex com- tralia: 4.5 km NW of jct of Blowholes Rd and plex. North West Coastal Hiway, N of Carnarvon, Female: Slightly more ovoid than male; 24.72267uS 113.7158uE, 28 m, 27 Oct 2004, small, mean total length 2.36, mean width 18 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 344

Fig. 4. Harpagophylus thryptomeni, male genitalia (AMNH_PBI 00373228). pronotum 0.77. COLORATION (pl. 1): As 05095093, 1- (00373227), 8U (00373233– in male. GENITALIA: As in generic descrip- 00373240) (AM), 3- (00372760–00372761, tion. 00373228), 6U (00372762–00372766, 00373241) ETYMOLOGY: Named for the host genus, (AMNH), 4U (00373229–00373232) (WAMP). Thryptomene (Myrtaceae); a noun in apposi- tion. HOSTS: Thryptomene aspera glabra E. Harpagophylus verticordii, new species Pritz. (Myrtaceae). Figures 5, 6; map 1; plate 1 DISTRIBUTION (map 1): Known from one DIAGNOSIS: Recognized by the pale, yel- locality in the Goldfields-Esperance region in lowish, coloration, and the conformation of Western Australia. the endosomal processes, both long, curving, HOLOTYPE: AUSTRALIA: Western Aus- and recurved toward the base of the endo- tralia: 28 km S of Menzies (3.5 km E of soma. Most easily confused with H. agnew, Hiway), 29.91917uS 121.1514uE, 500 m, 25 H. scholtzii,andH. thryptomeni on the basis Oct 1996, Schuh and Cassis, Thryptomene of pale coloration, but H. verticordii differing aspera aspera E. Pritz. (Myrtaceae), det. from all of those species in the structure and Perth 05095093, 1- (AMNH_PBI 00372759) conformation of the dorsal and ventral (WAMP). endosomal processes. PARATYPES: AUSTRALIA: Western Aus- DESCRIPTION: Male: Body weakly elon- tralia: 28 km S of Menzies (3.5 km E of gate, weakly ovoid; small, mean total length Hiway), 29.91917uS 121.1514uE, 500 m, 25 2.43, mean width pronotum 0.81. COLOR- Oct 1996, Schuh and Cassis, Thryptomene ATION (pl. 1): Head: Uniformly pale; sca- aspera aspera E.Pritz. (Myrtaceae), det. Perth pus unicolorous pale; pedicellus unicolorous 2010 SCHUH AND WEIRAUCH: MYRTACEAE-FEEDING PHYLINAE 19

Fig. 5. Harpagophylus verticordii, male. Scanning electron micrographs. A. Head and prothorax, lateral view. B. Setae on hemelytra. C. Pretarsus, dorsofrontal view. D. Abdomen, lateral view. E. Parameres, lateral view. F. Parameres, dorsal view. lp 5 left paramere; pe 5 parempodium; rp 5 right paramere (AMNH_PBI 00372775). pale; labium pale with segment 4 heavily veins weakly fumose; hind femur unicolorous infuscate. Thorax: Pronotum unicolorous pale, without black spots; hind tibial spines pale; scutellum unicolorous with remainder dark, without dark spots at bases. Abdomen: of dorsum; hemelytron unicolorous pale; Venter unicolorous pale or mostly so. SUR- markings on cuneus absent; membrane and FACE AND VESTITURE (fig. 5B): Dor- 20 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 344

Fig. 6. Harpagophylus verticordii, male and female genitalia (AMNH_PBI 00373222, 00373204). 2010 SCHUH AND WEIRAUCH: MYRTACEAE-FEEDING PHYLINAE 21 sum with reclining simple setae matching det. PERTH staff, 1- (00373201) background coloration and some sericeous or polytricha Benth. (Myrtaceae), det. Perth woolly setae. STRUCTURE: Head (fig. 5A): 05120594, 1U (00373209) (UCR), Verticordia Somewhat projecting; interocular space rela- monadelpha Turcz. (Myrtaceae), det. PERTH tively large; eyes leaving gena moderately staff, 1- (00373202) exposed in lateral view. Antenna: Segment 2 Benth. (Myrtaceae), det. Perth 05120594, 1U weakly tapering, more slender at base; (00373210) (UNSW), Verticordia monadelpha antennal fossa with ventral margin 1 diam- Turcz. (Myrtaceae), det. PERTH staff, 1- eter above ventral margin of eye. Labium: (00373185) Verticordia polytricha Benth. Reaching to about anterior margin of pygo- (Myrtaceae), det. Perth 05120594, 3- (0037- phore. GENITALIA (figs. 5D–F, 6): Phal- 3182–00373184), 4U (00373211–00373214) lotheca: Nearly erect; dorsal surface without (WAMP). North West Coast Hiway 72 km a fingerlike projection, and without a plate- NE of jct with Blowholes Rd, 24.18336uS like projection; anterior and ventral surfaces 114.0381uE, 34 m, 28 Oct 2004, Cassis, Wall, without a keel; apex complex. Weirauch, Tatarnic, Symonds, Verticordia Female: Slightly more ovoid than male; forrestii F.Muell. (Myrtaceae), det. Perth- small, mean total length 2.54, mean width 6989004, 1- (00373215), 2U (00373216, pronotum 0.86. COLORATION (pl. 1): As 00373217) (AM), Verticordia forrestii F. in male. GENITALIA (fig. 6): As in generic Muell. (Myrtaceae), det. Perth6989004, 2U description. (00322681, 00322682) (AMNH). ETYMOLOGY: Named for the host genus, OTHER SPECIMENS EXAMINED: AUSTRA- Verticordia (Myrtaceae). LIA: Western Australia: Kalbarri National HOSTS: diosmoides (DC.) Benth., Park, Loop Road, 27.56163uS 114.4376uE, Verticordia forrestii F. Muell., Verticordia 300 m, 28 Oct 1996, Schuh and Cassis, Dar- monadelpha Turcz (pl. 9A), V. polytricha winia diosmoides (DC.) Benth. (Myrtaceae), Benth. (pl. 9B) (Myrtaceae). det. Perth 05120551, 5- (00373218–00373- DISTRIBUTION (map 1): Known from 222), 3U (00373223–00373225) (AM). Western Australia, from Kalbarri National Park in the south to the Carnarvon region in Thryptomenomiris, new genus the north. Figures 7–9, 35–37; map 2; plate 1 HOLOTYPE: AUSTRALIA: Western Aus- tralia: Kalbarri National Park, Loop Road, TYPE SPECIES: Thryptomenomiris kalbarri, 27.56163uS 114.4376uE, 300 m, 28 Oct 1996, new species. Schuh and Cassis, Verticordia polytricha DIAGNOSIS: Recognized within the Mela- Benth. (Myrtaceae), det. Perth 05120594, leucoides genus group by the small size, the 1- (AMNH_PBI 00370995) (WAMP). apparent absence of a secondary endosomal PARATYPES: AUSTRALIA: Western Aus- strap (fig. 7; 31-3), the unique conformation tralia: Kalbarri National Park, Loop Road, of the left paramere, with a greatly elongate, 27.56163uS 114.4376uE, 300 m, 28 Oct 1996, medial, apical process (fig. 7; 56-2), and the Schuh and Cassis, Verticordia monadelpha female vestibulum with large, nearly symmet- Turcz. (Myrtaceae), det. PERTH staff, 6- rical medial plates (fig. 7; 74-1). Also recog- (00087221, 00373196–00373200), 5U (00373- nized within the Melaleucoides genus group 192, 00373205–00373208) Verticordia poly- by possessing only the ventral endosomal tricha Benth. (Myrtaceae), det. Perth 0512- process. Easily confused with Harpagophylus 0594, 9- (00373176–00373181, 00373193– calytrix, Melaleucoides akaina,andM. pi- 00373195), 5U (00373187–00373191) (AM), leanthicola on the basis of small size and Verticordia monadelpha Turcz. (Myrtaceae), reddish markings on a pale background. det. PERTH staff, 5- (00087511, 00087513, Easily distinguished from Melaleucoides aki- 00370998–00370999, 00373186), 5U (00371000– ana by the absence of the ventral endosomal 00371002, 00373203–00373204) Verticordia process in that species, from Harpagophylus polytricha Benth. (Myrtaceae), det. Perth 05- spp. by their possession of both ventral and 120594, 2- (00370996, 00370997) (AMNH), dorsal endosomal processes, and from all of Verticordia monadelpha Turcz. (Myrtaceae), those taxa by the presence of a secondary 22 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 344

ary gonopore seen laterally in lateral view of endosoma. Phallotheca: Smoothly curving or nearly straight on dorsal margin; ventral surface with or without a projecting keel; apex simple. Left Paramere: Vertical; shaft at right angles to body; in dorsal perspective (fig. 8F), not exceeding margin of pygophore; in lateral perspective (fig. 8E) covering limit- ed extent of lateral surface of pygophore; body more or less quadrate; apex medially broadly rounded to weakly quadrate; apex posterolaterally with a greatly elongate pro- jection (figs. 7, 8F, 9); anterior process slender and cylindrical in lateral view; poste- rior process in the form of a short, straight, fingerlike projection; base of posterior pro- cess without conspicuous shoulder. Right Paramere: Body short and broad, lanceolate; apex short, clawlike, decurved. Female: Slightly more ovoid than male; small, mean total length 2.52–2.82, mean width pronotum 0.80–0.93. COLORATION (pl. 1): As in male. STRUCTURE: Head Map 2. Localities for species of Thryptomeno- (fig. 8B). GENITALIA (fig. 7): Posterior miris. wall laterally with distinct crescent-shaped interramal sclerites; posterolaterally with a distinct swelling covered with microtrichia; endosomal strap and the distinctive structure longitudinal fold on either side of midline of the left paramere in Thryptomenomiris. along part of length; interramal lobes roughly DESCRIPTION: Male: Body moderately symmetrical, erect, and not ornamented with elongate, parallel sided; small, mean total spicules or with a very few; vestibulum with length 2.44–2.79, mean width pronotum medial plates sclerotized and readily visible, 0.77–0.92. COLORATION (pl. 1): Pale with large, nearly symmetrical, with sclerotized some brownish or reddish areas. SURFACE guide structure present as caplike structure AND VESTITURE (fig. 8C): Dorsum with with internal differentiation. suberect or reclining black setae and some ETYMOLOGY: Named for the host genus, sericeous or woolly setae. STRUCTURE: Thryptomene (Myrtaceae), in combination Head (fig. 8A): Somewhat projecting; intero- with Miris; gender masculine. cular space moderate; eyes leaving gena moderately exposed in lateral view. Antenna: Thryptomenomiris kalbarri, new species Segment 2 weakly tapering, more slender at Figure 7, map 2, plate 1 base; antennal fossa with ventral margin 1 diameter above ventral margin of eye. DIAGNOSIS: In addition to the character- Labium: Just reaching onto abdomen. Tho- istics in the generic diagnosis, recognized by rax: Pretarsus (fig. 8D): Parempodia fleshy, the attenuated apex of the apical spine of the recurved, lyriform; pulvilli absent. GENITA- left paramere (fig. 7), in contrast to the blunt LIA (figs. 7, 8E, F, 9): Endosoma: Base short, spine in T. yalgoo (fig. 9). curving, C- or J-shaped; body without DESCRIPTION: Male: Body weakly to torsion, primary strap ventral to secondary distinctly ovoid; small, mean total length gonopore, apically greatly elongate, without 2.48, mean width pronotum 0.82. COLOR- ornamentation; spinelike, elongate process ATION (pl. 1): Head: Uniformly pale; sca- near gonopore on ventral surface present, pus and pedicellus unicolorous pale; labium recurved toward base of endosoma; second- pale with segment 4 heavily infuscate. Tho- 2010 SCHUH AND WEIRAUCH: MYRTACEAE-FEEDING PHYLINAE 23 rax: Pronotum unicolorous pale; scutellum 00373257, 00373275–00373276, 00373284), 17U unicolorous with remainder of dorsum; (00373258–00373265, 00373277–00373283, 00- hemelytron pale with endocorium somewhat 373287, 00373292) (AM), 4- (00087217, darker than remainder; markings on cuneus 00372769–00372770, 00373291), 7U (000872- absent, cuneus transparent; membrane and 18, 00372772–00372774, 00373272–00373274) veins weakly fumose; hind femur unicolorous (AMNH), 1- (00373285), 2U (00373288, pale, without black spots; hind tibial spines 00373289) (UCR), 2- (00373255, 00373290), dark, without dark spots at bases. Abdomen: 1U (00373286) (UNSW), 6U (00373266– Venter unicolorous pale or mostly so. SUR- 00373271) (WAMP). FACE AND VESTITURE: Dorsum with OTHER SPECIMENS EXAMINED: AUSTRA- suberect or reclining black setae and some LIA: Western Australia: 25 km S of Gasco- sericeous or woolly setae. STRUCTURE: yne Junction, on Towrana Homestead Rd, Head: Somewhat projecting; interocular 25.39014uS 115.1506uE, 232 m, 04 Nov 2004, space moderate; eyes leaving gena moderate- Cassis, Weirauch, Tatarnic, Symonds, Sola- ly exposed in lateral view. Antenna: Segment num sp. (Solanaceae), det. Field ID, 1- 2 weakly tapering, more slender at base; (00373293), 1U (00373294) (AM). ca. 1 km S antennal fossa with ventral margin 1 diam- of Murchison House HS, Kalbarri National eter above ventral margin of eye. Labium: Park, 27.65822uS 114.2394uE, 60 m, 23 Oct Just reaching onto abdomen. GENITALIA 2004, Cassis, Wall, Weirauch, Symonds, (fig. 7): Phallotheca: Smoothly curving on hursthousei (W.Fitzg.) J.W.- dorsal margin; dorsal surface without a Green (Myrtaceae), det. Perth6988423, 1- fingerlike or platelike projection; anterior (00373295) (AM). surface without a keel; ventral surface with a projecting keel; apex simple. Thryptomenomiris yalgoo, new species Female: Slightly more ovoid than male; Figures 8, 9; map 2; plate 1 small, mean total length 2.66, mean width pronotum 0.84. COLORATION (pl. 1): As DIAGNOSIS: In addition to the character- in male. GENITALIA (fig. 7): See generic istics in the generic diagnosis, recognized by description. the blunt apex of the apical spine of the left ETYMOLOGY: Named for the region of the paramere (fig. 9), in contrast to the attenu- type locality, Kalbarri, Western Australia; a ated spine in T. kalbarri (fig. 7). noun in apposition. DESCRIPTION: Male: Body weakly elon- HOSTS: Thryptomene sp. (Myrtaceae). The gate, weakly ovoid; small, mean total length record from Malleostemon hursthousei may 2.65, mean width pronotum 0.88. COLOR- well represent a genuine host, but would ATION (pl. 1): Head: Uniformly pale; sca- benefit from additional documentation. The pus and pedicellus unicolorous pale; labium record from Solanum probably results from pale with segment 4 heavily infuscate. Tho- specimens being commingled in the field and rax: Pronotum unicolorous pale; scutellum does not represent a breeding host. entirely red or orange; hemelytron with red DISTRIBUTION (map 2): Known from or carmine spots, botches, or solid areas; Kalbarri National Park and the Gascoyne markings on cuneus absent; membrane and region in Western Australia. veins weakly fumose; hind femur unicolorous HOLOTYPE: AUSTRALIA: Western Aus- pale, without black spots; hind tibial spines tralia: Kalbarri National Park, Loop Road, dark, without dark spots at bases. Abdomen: 27.56163uS 114.4376uE, 300 m, 28 Oct 1996, Venter unicolorous pale or mostly so. SUR- Schuh and Cassis, Thryptomene sp. (Myrta- FACE AND VESTITURE (fig. 8C): Dor- ceae), det. Perth 05120543, 1- (AMNH_PBI sum with reclining simple setae matching 00372768) (WAMP). background coloration and some sericeous or PARATYPES: AUSTRALIA: Western Aus- woolly setae. STRUCTURE: Head (fig. 8A): tralia: Kalbarri National Park, Loop Road, Somewhat projecting; interocular space rela- 27.56163uS 114.4376uE, 300 m, 28 Oct 1996, tively large; eyes leaving gena moderately Schuh and Cassis, Thryptomene sp. (Myrta- exposed in lateral view. Antenna: Segment 2 ceae), det. Perth 05120543, 5- (00373256– weakly tapering, more slender at base; 24 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 344

Fig. 7. Thryptomenomiris kalbarri, male and female genitalia (AMNH_PBI 00373257, 00373274). 2010 SCHUH AND WEIRAUCH: MYRTACEAE-FEEDING PHYLINAE 25

Fig. 8. Thryptomenomiris yalgoo. Scanning electron micrographs. A. Head and prothorax, lateral view, male. B. Head and prothorax, lateral view, female. C. Setae on hemelytra. D. Pretarsus, laterofrontal view. E. Pygophore, lateral view. F. Parameres, dorsal view. lp 5 left paramere; pe 5 pulvillus; rp 5 right paramere (AMNH_PBI 00135313). antennal fossa with ventral margin at ventral GENITALIA (figs. 8E, F, 9). Phallotheca: margin of eye. Labium: Reaching to about Smoothly curving on dorsal margin; dorsal anterior margin of pygophore. Thorax: Pre- surface without a fingerlike or platelike tarsus (fig. 8D): As in generic description. projection; anterior surface with a short to 26 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 344

Fig. 9. Thryptomenomiris yalgoo, male genitalia (AMNH_PBI 00373296). elongate keel; ventral surface with a project- (00372767), 3U (00087220, 00373297–003732- ing keel; apex simple. 98) (AM), 1- (00373296) (AMNH). 46.5 km Female: Slightly more ovoid than male; W of Yalgoo, 28.41302uS 116.2151uE, 600 m, small, mean total length 2.77, mean width 27 Oct 1996, Schuh and Cassis, Thryptomene pronotum 0.90. COLORATION (pl. 1): As aspera glabra E. Pritz. (Myrtaceae), det. Perth in male. STRUCTURE: Head (fig. 8B). 05120683, 2- (00135304, 00135312), 1U (001- GENITALIA: Not examined. 35317) (AM), 9- (00135305–00135306, 0013- ETYMOLOGY: Named for the small town 5308–00135311, 00135314–00135316), 10U of Yalgoo in the region of the type locality, (00135318–00135320, 00135323, 00135325– Western Australia; a noun in apposition. 00135330) (AMNH), 2U (00135321, 001353- HOSTS: Thryptomene aspera glabra E. 22) (WAMP). Pritz. (Myrtaceae). DISTRIBUTION (map 2): Known from Melaleucoides, new genus Yalgoo and Sandstone (Mid West region) Figures 10–37; maps 3–6; plates 1–4 in Western Australia. HOLOTYPE: AUSTRALIA: Western Aus- TYPE SPECIES: Melaleucoides cassisi, new tralia: 46.5 km W of Yalgoo, 28.41302uS species. 116.2151uE, 600 m, 27 Oct 1996, Schuh and DIAGNOSIS: Recognized within the Mela- Cassis, Thryptomene aspera glabra E. Pritz. leucoides genus group by the unique structure (Myrtaceae), det. Perth 05120683, 1- of the left paramere, with its flattened, spoon- (AMNH_PBI 00135307) (WAMP). shaped form and erect position, covering the PARATYPES: AUSTRALIA: Western Aus- dorsal margin of the pygophore in lateral tralia: 24 km W of Sandstone, 28.01426uS view (fig. 12D; 53-1). Most easily confused 119.0474uE, 650 m, 26 Oct 1996, Schuh and with Harpagophylus and Thryptomenomiris Cassis, Thryptomene aspera glabra E. Pritz. on the basis of unique parempodial structure (Myrtaceae), det. Perth 05095182, 1- possessed by all three genera, but easily 2010 SCHUH AND WEIRAUCH: MYRTACEAE-FEEDING PHYLINAE 27 separated from both on the structure of the sal margin of pygophore; body quadrate or left paramere. spoon shaped; apex medially broadly round- DESCRIPTION: Male: Body weakly elon- ed to weakly quadrate, sometimes elongated gate, weakly ovoid; mean total length 2.33– and or recurved; apex posterolaterally simple 4.05, mean width pronotum 0.78–1.55. COL- or in the form of a prolongation; anterior ORATION (pls. 1–4): Variable, ranging process triangular or elongate and cylindrical from completely castaneous to almost com- in lateral view; posterior process in the form pletely pale, and sometimes with spots. of a short to long, fingerlike projection; base SURFACE AND VESTITURE (figs. 12B, of posterior process with or without conspic- 17B): Dorsum with suberect or reclining uous shoulder. Right Paramere: Body short black setae, without sericeous or woolly and broad, lanceolate; apex short, clawlike, setae. STRUCTURE: Head (figs. 12A, 14A, decurved. 17A, 33A): Barely projecting; interocular Female: Slightly more ovoid than male; space moderate; eyes leaving gena moderate- small, mean total length 2.52–4.26, mean ly exposed in lateral view. Antenna: Segment width pronotum 0.82–1.59. COLORATION 2 weakly tapering, more slender at base; (pls. 1–4): As in male. GENITALIA antennal fossa with ventral margin at ventral (figs. 11, 18, 25, 34): Posterior wall laterally margin of eye. Labium: Just reaching onto with distinct crescent-shaped interramal abdomen. Thorax: Pretarsus (figs. 12C, 17C, sclerites; posterolaterally with a distinct 33D): Parempodia fleshy, recurved, lyriform; swelling covered with microtrichia; longitu- pulvilli present, flaplike, covering about one dinal fold on either side of midline along part third of ventral claw surface. GENITALIA of length; interramal lobes present and (e.g., figs. 11, 12E, F, 13, 18, 33F, G, 34): asymmetrical, reclining, overlapping, and Endosoma: Base short; curving, C- or J- heavily ornamented with spicules; vestibulum shaped; body without torsion; primary strap with medial plates sclerotized and readily ventral to secondary gonopore, apically observed; small, nearly symmetrical, triangu- greatly elongate, straight, weakly curving, lar; with sclerotized guide structure present as or angulate relative to body of endosoma, caplike structure with internal differentiation. without ornamentation; secondary endoso- ETYMOLOGY: Named after the host genus, mal strap fused with primary strap proximal (Myrtaceae), in combination with to secondary gonopore, usually reaching well the Latin suffix -oides, a diminutive; gender beyond gonopore, often fused with primary feminine. strap, sometimes with denticles or comblike ornamentation; sometimes with elongate spinelike process arising near gonopore on KEYTOSPECIESOFMELALEUCOIDES dorsal surface; sometimes with spinelike 1. Dorsum largely castaneous, including entire elongate process near gonopore on ventral clavus(pls.2–4)...... 2 surface, ornamented with denticles or not; – Dorsum not so heavily castaneous, at least sometimes with bladderlike process distad of clavus and anterior half of endocorium pale, secondary gonopore; secondary gonopore red, or with dark spots on pale background sclerite absent; secondary gonopore usually ...... 4 seen frontally (facing up) in lateral view of .2. Dorsum uniformly castaneous, including endosoma. Phallotheca: Smoothly curving on cuneus and apex of scutellum (pl. 2); male dorsal margin, without a fingerlike projec- genitalia as in figure 19 ...... tion, and with a curving, posteriorly directed, ...... Melaleucoides castanea platelike projection; anterior surface without – At least clavus and apex of scutellum pale . . . 3 3. Male genitalia as in figure 26, phallotheca a keel; ventral surface without a projecting with a short fingerlike projection on dorsal keel; posterior surface lacking transparent margin; known from Melaleuca rhaphiophyl- window; apex simple. Left Paramere: Verti- la...... Melaleucoides rhaphiophyllae cal; shaft at right angles to or in same axis as – Male genitalia as in figure 31, phallotheca body; in dorsal perspective (fig. 12F) not with a long fingerlike projection on dorsal exceeding margin of pygophore; in lateral margin; known from Melaleuca uncina- perspective (fig. 12E) extending beyond dor- ta...... Melaleucoides uncinatae 28 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 344

Maps 3 and 4. Localities for species of Melaleucoides: M. akaina–M. castanea (top; map 3) and M. grossi–M. rhapiophyllae (bottom; map 4). 2010 SCHUH AND WEIRAUCH: MYRTACEAE-FEEDING PHYLINAE 29

Map 5. Localities for species of Melaleucoides: Map 6. Localities for species of Melaleucoides: M. sheathianae–M. systenae. M. uncinatae–M. verticordiae.

4. Dorsum with conspicuous small dark spots – Calli not castaneous, although sometimes and a pale background ...... 5 weakly red, not strongly contrasting with – Dorsum without small dark spots on a pale surrounding pronotum...... 11 background...... 8 10. Red markings on hemelytron broad, includ- 5. All spots on dorsum small (pl. 2); apex of ing most of clavus and endocorium, but not scutellum dark; male genitalia as in figure 16; including claval and radiomedial veins (pl. 1); known from Melaleuca brevifolia ...... male genitalia as in figure 13, known from Me- ...... Melaleucoides brevifoliae laleuca sheathiana .... Melaleucoides annae – Spots on dorsum larger (pls. 2, 3); apex of – Red markings on hemelytron narrow, running scutellum pale as remainder of scutellum. 6 along claval and radiomedial veins (pl. 4); male 6. Spots round, never coalescent (pls. 2, 3). . . 7 genitalia as in figure 32, known from Melaleu- – Many spots of irregular shape, often coalescent, ca undulata ...... Melaleucoides undulatae particularly on hemelytron (pl. 2); male geni- 11. Cuneus almost entirely pale; broad marking talia as in figure 15; known from on endocorium intensely red (pls. 1, 3) . . 12 micrantha ...... Melaleucoides beaufortiae – Apical half of cuneus red, in contrast with 7. Male genitalia as in figure 21; known from remainder (pl. 3); male genitalia as in figure 25; Melaleuca leuropomae ...... known from Thryptomene kochii ...... Melaleucoides leuropomae ...... Melaleucoides ozzii – Male genitalia as in figure 22; known from 12. Male genitalia as in figure 25, endosoma with Melaleuca micrantha ...... two distinct, elongate, apical spines and a ...... Melaleucoides micranthae ventral process adjacent to the secondary 8. Hemelytron with conspicuous longitudinal gonopore; known from spp. redareas...... 9 ...... Melaleucoides pileanthicola – Hemelytron without conspicuous longitudi- – Male genitalia as in figure 11, endosoma with nalredareas...... 13 a single, short, apical spine and no processes 9. Calli castaneous, much darker than sur- associated with the secondary gonopore; rounding pronotum and strongly contrasting known from hursthousei and withit...... 10 Scholtzia leptantha... Melaleucoides akaina 30 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 344

13. Coloration of dorsum nearly uniform, either of dorsum; hemelytron mostly pale, endocor- faded green, greenish yellow, or yellow. . . 14 ium somewhat darker than remainder; mark- – Coloration of dorsum not uniform, posterior ings on cuneus absent; membrane, including half of endocorium and scutellum dark, in veins, pale; hind femur unicolorous pale, contrast with much of remainder of hemely- without black spots; hind tibial spines dark, tron and pronotum (pl. 4); male genitalia as in figure 29; known from Melaleuca systena without dark spots at bases. Abdomen: ...... Melaleucoides systenae Venter unicolorous pale or mostly so. SUR- 14. Large, robust species, total length at least FACE AND VESTITURE (fig. 10C): Vesti- 3.50, width pronotum at least 1.31; coloration ture of dorsum consisting of simple, recum- of dorsum faded green or yellow green (pls. 2, bent setae with coloration of dorsum. 3)...... 15 STRUCTURE: Head: Somewhat projecting; – More elongate species, total length no more interocular space relatively large; eyes leaving than 3.76, width pronotum no more than gena only very slightly exposed in lateral 1.28; coloration of dorsum bright yellow view. Antenna: Segment 2 weakly tapering, orange(pls.3,4)...... 16 more slender at base; antennal fossa with 15. Scutellum white or nearly so (pl. 2); male ventral margin at ventral margin of eye. genitalia as in figure 18; known from Mela- leuca uncinata ...... Melaleucoides cassisi Labium: Reaching to about anterior margin – Scutellum orange (pl. 3); male genitalia as in of pygophore. Thorax: Pretarsus (fig. 10D): figure 18; known from Melaleuca sheathia- Parempodia fleshy, recurved, lyriform; pul- na ...... Melaleucoides sheathianae villi absent. GENITALIA (fig. 11): Endo- 16. Male genitalia as in figure 28; known from soma: Base short, curving, C- or J-shaped; multiple genera of Myrtaceae ...... body with torsion, primary strap dorsal to ...... Melaleucoides similis secondary gonopore, apically short, ratio – Male genitalia as in figure 34; known from length apex strap/length secondary gonopore Verticordia spp. and Pileanthus spp., . . . . 1.33–1.50, at least weakly arcuate and ...... Melaleucoides verticoridiae curving dorsally, no ornamentation; second- ary endosomal strap fused with primary Melaleucoides akaina, new species strap, reaching just beyond secondary gono- Figures 10, 11; map 3; plate 1 pore as fingerlike extension with one tip, and DIAGNOSIS: Unequivocally recognized devoid of ornamentation; secondary gono- among Melaleucoides species by the uniquely pore seen laterally in lateral view of endo- short, stout endosoma with a short apical soma, with a denticulate ridge on right side of projection, no spinelike processes associated endosoma. Phallotheca: Smoothly curving on with the region of the secondary gonopore, dorsal margin; apex simple. Left Paramere: and a row of denticles on the margin of the Vertical; shaft at right angles to body; in secondary gonopore, as well as the long, dorsal perspective not exceeding margin of recurved, tapered, apical spine on the apex of pygophore; in lateral perspective covering the left paramere. Easily confused with limited extent of lateral surface of pygophore; Harpagophylus calytrix, Thryptomenomiris body spoon shaped; apex medially with an spp., and Melaleucoides pileanthicola on the elongate, acuminate, recurved, medial spine; basis its small size and reddish coloration on anterior process triangular in lateral view; a pale background. All of those taxa with posterior process in the form of a long, ventral/and or dorsal processes on the straight, slender, fingerlike projection; base endosoma and with a long apical endosomal of posterior process without conspicuous spine. shoulder. Right Paramere: Body short and DESCRIPTION: Male: Body moderately broad, lanceolate; apex short, clawlike, de- elongate, parallel sided; small, mean total curved. length 2.36, mean width pronotum 0.79. Female: Slightly more ovoid than male; COLORATION (pl. 1): Head: Uniformly mean total length 2.61, mean width prono- pale; scapus and pedicellus yellow to dirty tum 0.85. COLORATION (pl. 1): Paler than yellow; labium pale with segment 4 heavily male. STRUCTURE: Head (fig. 10A). GEN- infuscate. Thorax: Pronotum unicolorous ITALIA (fig. 11): Posterior wall laterally pale; scutellum unicolorous with remainder with distinct crescent-shaped interramal scler- 2010 SCHUH AND WEIRAUCH: MYRTACEAE-FEEDING PHYLINAE 31

Fig. 10. Melaleucoides akaina, female. Scanning electron micrographs. A. Head and prothorax, lateral view. B. Metathoracic scent-gland evaporatory and mesothoracic spiracle. C. Setae on hemelytra. D. Pretarsus, dorsofrontal view. eva 5 evaporatory area; mssp 5 mesothoracic spiracle; pe 5 parempodium (AMNH_PBI 00137503).

ites; posteriorly without a sclerotized trans- DISTRIBUTION (map 3): Known from two verse band; posterolaterally with a distinct localities north of Kalbarri National Park swelling covered with microtrichia; longitudi- and the Carnarvon region of Western Aus- nal fold on either side of midline along part of tralia. length; posterior margin of posterior wall DISCUSSION: Melaleucoides akaina is without spicules, and not reflexed dorsally; unique among members of the genus in having interramal lobes roughly symmetrical, erect, the endosoma with no processes associated and heavily ornamented with spicules; vestib- with the region of the secondary gonopore. ulum with medial plates sclerotized and Nonetheless, other aspects of morphology, readily observed, large, nearly symmetrical, especially the novel structure of the parempo- with sclerotized guide structure present as dia and left paramere agree closely with the short heavily sclerotized tube on left side. remaining species we place in Melaleucoides. ETYMOLOGY: Named for the spearlike On the basis of our phylogenetic analyses, we apex of the left paramere, from the Greek conclude that the relatively simple endosoma akaina, ‘‘thorn, spine.’’ in M. akaina is the result of reduction. HOSTS: Micromyrtus hursthousei W. Fitzg. HOLOTYPE: AUSTRALIA: Western Aus- (pl. 7A) and Scholtzia leptantha Benth. tralia: NW Coastal Hiway 58 km N of Kalbarri (Myrtaceae). Road, 27.43701uS 114.6768uE, 500 m, 30 Oct 32 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 344

Fig. 11. Melaleucoides akaina, male and female genitalia (AMNH_PBI 00087543, 00373174).

1996, Schuh and Cassis, Micromyrtus hurst- Weirauch, Tatarnic, Symonds, Scholtzia housei W. Fitzgerald (Myrtaceae), det. Perth leptantha Benth. (Myrtaceae), det. Perth- 05120241, 1-(AMNH_PBI 00372340) (WAMP). 6988687, 3- (00373164–00373166), 3U PARATYPES: AUSTRALIA: Western Aus- (00373170–00373172) (AM), 3- (003731- tralia: NW Coastal Hiway 58 km N of Kal- 67–00373169), 2U (00373173, 00373174) barri Road, 27.43701uS 114.6768uE, 500 m, 30 (AMNH). Oct 1996, Schuh and Cassis, Micromyrtus hursthousei W. Fitzgerald (Myrtaceae), det. - Melaleucoides annae, new species Perth 05120241, 4 (00372342, 00372343, Figures 12, 13; map 3; plate 1 00087543, 00087346), 3U (00372345–00087347) (AM), 1-(00372344), 1U(00372347) (AMNH), DIAGNOSIS: Recognized by the castaneous 1- (00372341), 1U (00372348) (WAMP). calli, the elongate red markings on the OTHER SPECIMENS EXAMINED: AUSTRA- hemelytron (pl. 1), and the endosoma with LIA: Western Australia: 4.5 km NW of jct denticles on the secondary strap distad of the of Blowholes Rd and North West Coas- secondary gonopore and a recurved ventral tal Hiway, N of Carnarvon, 24.72267uS process associated with the secondary gono- 113.7158uE, 28 m, 27 Oct 2004, Cassis, Wall, pore (fig. 13). Most easily confused with M. 2010 SCHUH AND WEIRAUCH: MYRTACEAE-FEEDING PHYLINAE 33 undulatae on the basis of coloration, but the directed, platelike projection; anterior surface red markings on the hemelytron more with a short to elongate keel; apex simple. strongly linear in that species (pl. 4) and the Left Paramere: Shaft at right angles to body; lack of denticles distad of the secondary body spoon shaped; apex medially broadly gonopore, a dorsal rather than a ventral rounded; anterior process triangular in later- process, and a bladderlike process just distad al view; posterior process in the form of a of the gonopore (fig. 32). short, straight, fingerlike projection; base of DESCRIPTION: Male: Body weakly elon- posterior process without conspicuous shoul- gate, weakly ovoid; mean total length 3.46, der. Right Paramere: Body short and broad, mean width pronotum 0.1.29. COLORA- lanceolate; apex short, clawlike, decurved. TION (pl. 1): Head: Uniformly pale; scapus Female: Slightly more ovoid than male; yellow to dirty yellow; pedicellus dirty yellow mean total length 3.51, mean width prono- proximally, weakly to heavily infuscate dis- tum 0.1.36. COLORATION (pl. 1): As in tally; labium generally infuscate, heavily so male. GENITALIA: Posterior wall laterally apically. Thorax: Pronotum pale with casta- with distinct crescent-shaped interramal neous calli and sometimes humeri; scutellum sclerites; posteriorly without a sclerotized white; hemelytron with red or carmine areas transverse band; posterolaterally with a along mesial margin of clavus and on distinct swelling covered with microtrichia; endocorium; markings on cuneus present as longitudinal fold on either side of midline a contrasting white basal fascia, remainder along part of length; posterior margin of red; membrane weakly fumose, veins pale; posterior wall without spicules and not hind femur with a few brown or black spots; reflexed dorsally; interramal lobes present hind tibial spines black with very small dark and asymmetrical, reclining, overlapping, bases. Abdomen: Venter unicolorous pale or and heavily ornamented with spicules; ves- mostly so. SURFACE AND VESTITURE tibulum with medial plates sclerotized and (fig. 12B): Dorsum with suberect or reclining readily observed, small, nearly symmetrical, black setae and some sericeous or woolly triangular; with sclerotized guide present as setae. STRUCTURE: Head (fig. 12A): Bare- caplike structure with internal differentiation. ly projecting; interocular space moderate; ETYMOLOGY: Named for Anna Massie, eyes leaving gena moderately exposed in with thanks for her encouragement and lateral view. Antenna: Segment 2 weakly interest in our work on the Australian tapering, more slender at base; antennal fossa Miridae fauna. with ventral margin at ventral margin of eye. HOSTS: Melaleuca sheathiana W. Fitz. Labium: Just reaching onto abdomen. Tho- (Myrtaceae) (pl. 6A–C). rax: Pretarsus (fig. 12C): Pulvilli present, DISTRIBUTION (map 3): Known from one flaplike, covering about one third of ventral locality in the Goldfields region of Western claw surface. GENITALIA (figs. 12E, F, 13): Australia. Endosoma: Primary strap apically greatly HOLOTYPE: AUSTRALIA: Western Aus- elongate, angled dorsally relative to body of tralia: 11 km N of Coolgardie-Esperance endosoma, no ornamentation; secondary Hiway on Kambalda Road, 31.25231uS endosomal strap fused with primary strap 121.5899uE, 320 m, 18 Nov 1999, R.T. Schuh, proximal to secondary gonopore, reaching G. Cassis, & R. Silveira, Melaleuca sheathiana well beyond gonopore, fused with primary W. Fitzg. (Myrtaceae), det. Perth 05671396, strap, ornamented with some short denticles; 1- (AMNH_PBI 00129550) (WAMP). spinelike, elongate process arising near gon- PARATYPES: AUSTRALIA: Western Aus- opore on dorsal surface absent; spinelike, tralia: 11 km N of Coolgardie-Esperance elongate process near gonopore on ventral Hiway on Kambalda Road, 31.25231uS surface present, strongly recurved toward 121.5899uE, 320 m, 18 Nov 1999, R.T. Schuh, base of endosoma; bladderlike process distad G. Cassis, & R. Silveira, Melaleuca sheathiana of secondary gonopore absent; secondary W. Fitzg. (Myrtaceae), det. Perth 05671396, gonopore seen frontally (facing up) in lateral 23- (00371206, 00371250–00371261, 0037- view of endosoma. Phallotheca: Nearly erect; 1265, 00371288–00371296), 35U (00371266– dorsal surface with a curving, posteriorly 00371270, 00371272–00371277, 00371280– 34 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 344

Fig. 12. Melaleucoides annae, male. Scanning electron micrographs. A. Lateral view. B. Setae on hemelytra. C. Pretarsus, frontal view. D. Abdomen, lateral view. E. Pygophore and left paramere, lateral view. F. Pygophore and parameres, dorsal view. lp 5 left paramere; pe 5 parempodium; rp 5 right paramere (AMNH_PBI 00129566).

00371285, 00371287, 00371297–00371312, 00129582–00129583, 00129601–00129604, 00- 00372109), 1 nymph (00371313) (AM), 31- 371278) (AMNH), 3- (00372092–00372094), (00087299, 00087498, 00129544–00129547, 2U (00372113, 00372114) (ANIC), 1- (0037- 00129551–00129569, 00129595–00129600), 17U 1264), 1U (00371286) (CNC), 2- (00129570, (00087300, 00129548, 00129572–00129579, 00129571), 2U (00129580, 00129581) (UCR), 2010 SCHUH AND WEIRAUCH: MYRTACEAE-FEEDING PHYLINAE 35

Fig. 13. Melaleucoides annae, male genitalia (AMNH_PBI 00129556).

10- (00372082–00372091), 4U (00372095, Melaleucoides beaufortiae, new species 00372110–00372112) (UNSW), 1- (00371- Figures 14, 15; map 3; plate 2 262), 1U (00371271) (USNM), 9- (00129- 531–00129533, 00371207–00371212), 16U (00- DIAGNOSIS: Unique among Melaleucoides 129534–00129543, 00371223–00371228) (WAMP), spp. for the coalescent small dark spots on 1- (00371263), 1U (00371279) (ZISP). the dorsum (pl. 2), the compact body form, OTHER SPECIMENS EXAMINED: AUSTRA- and the structure of the male genitalia, the LIA: Western Australia: 11 km N of Cool- left paramere with a moderately elongate, gardie-Esperance Hiway on Kambalda Road, blunt, fingerlike, recurved, medial apical 31.25231uS 121.5899uE, 320 m, 18 Nov 1999, process, and the endosoma with some denti- R.T. Schuh, G. Cassis, & R. Silveira, Mela- cles on the postgonoporal region of the leuca sheathiana W.Fitzg. (Myrtaceae), det. secondary strap but lacking both the dorsal Perth 05671396, 14- (00371192–00371205), and ventral processes frequently seen in the 10U (00371213–00371222) (AM). genus (fig. 15). Most easily confused with M. 36 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 344

Fig. 14. Melaleucoides beaufortiae, male. Scanning electron micrographs. A. Head and thorax, lateral view. B. Pretarsus, frontal view. C. Pygophore, lateral view. D. Pygophore, dorsal view. lp 5 left paramere; pe 5 parempodium; pv 5 pulvillus (AMNH_PBI 00371072). brevifoliae, M. leuropomae, and M. mi- many brown or black spots; hind tibial spines cranthae, but the spots never coalescent in dark with conspicuous dark spots at bases. those species and the apical medial process of Abdomen: Venter unicolorous pale or mostly the left paramere much less elongate and not so. SURFACE AND VESTITURE: Dorsum recurved (figs. 16, 21, 22, respectively). with suberect or reclining black setae, with- DESCRIPTION: Male: Body weakly elon- out sericeous or woolly setae. STRUC- gate, weakly ovoid; mean total length 2.98, TURE: Head (fig. 14A): Somewhat project- mean width pronotum 1.10. COLORATION ing; interocular space moderate; eyes leaving (pl. 2): Head: Pale, clypeus dark and large, gena broadly exposed in lateral view. Anten- dark, linear, bilateral markings on frons; na: Segment 2 weakly tapering, more slender scapus pale with a dark base; pedicellus dirty at base; antennal fossa with dorsal margin yellow proximally, weakly to heavily infus- somewhat below ventral margin of eye. cate distally; labium generally infuscate, Labium: Reaching to about anterior margin heavily so apically. Thorax: Pronotum pale of pygophore. Thorax: Pretarsus (fig. 14B): with brown spots of varying size; scutellum Pulvilli present, flaplike, covering about one pale with brown spots; hemelytron unicolor- third of ventral claw surface. GENITALIA ous pale with brown spots of varying size, (fig. 14C, D, 15): Endosoma: Primary strap some coalescent; markings on cuneus present apically greatly elongate, curving dorsally as a red to castaneous apex; membrane near apex, without ornamentation; secondary weakly fumose, veins pale; hind femur with endosomal strap fused with primary strap 2010 SCHUH AND WEIRAUCH: MYRTACEAE-FEEDING PHYLINAE 37

Fig. 15. Melaleucoides beaufortiae, male genitalia (AMNH_PBI 00371078). proximal to secondary gonopore, reaching of a short curved fingerlike projection; base well beyond gonopore, ornamented with of posterior process without conspicuous short denticles just distad of gonopore; shoulder. Right Paramere: Body short and spinelike, elongate process arising near gon- broad, lanceolate; apex short, clawlike, de- opore on dorsal surface absent; spinelike, curved. elongate process near gonopore on ventral Female: Slightly more ovoid than male; surface absent; bladderlike process distad of mean total length 3.06, mean width prono- secondary gonopore absent; secondary gon- tum 1.13. COLORATION (pl. 2): As in opore seen frontally (facing up) in lateral male. GENITALIA: Posterior wall laterally view of endosoma. Phallotheca: Nearly erect; with distinct crescent-shaped interramal apex simple. Left Paramere: Shaft at right sclerites; posteriorly without a sclerotized angles to body; body spoon shaped; apex transverse band; posterolaterally with a medially drawn into a blunt, recurved, distinct swelling covered with microtrichia; fingerlike process; anterior process triangular longitudinal fold on either side of midline in lateral view; posterior process in the form along part of length; posterior margin of 38 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 344 posterior wall without spicules, and not the male genitalia, the left paramere with a reflexed dorsally; interramal lobes present somewhat elongate apex positioned asym- and asymmetrical, reclining, overlapping, metrically on the anterior side of the struc- and heavily ornamented with spicules; ves- ture; endosoma unknown. Most easily con- tibulum with medial plates sclerotized and fused with M. beaufortiae, M. leuropomae, readily observed, small, nearly symmetrical, and M. micranthae, but the spots in those triangular; with sclerotized guide present as species always larger and the apical medial short heavily sclerotized tube on left side. process much longer in M. beaufortiae and ETYMOLOGY: Named for the host genus, placed medially in all three species (figs. 15, Beaufortia R. Br. (Myrtaceae). 21, 22, respectively). HOSTS: Beaufortia micrantha Schauer DESCRIPTION: Male: Body weakly to (Myrtaceae). distinctly ovoid; mean total length 3.41, mean DISTRIBUTION (map 3): Known from one width pronotum 1.10. COLORATION locality in Fitzgerald River National Park in (pl. 2): Head: Pale or pale with some small Southwestern Australia. dark dots; scapus and pedicellus unicolorous HOLOTYPE: AUSTRALIA: Western Aus- pale; labium generally infuscate, heavily so tralia: Fitzgerald River National Park, Ham- apically. Thorax: Pronotum pale with tiny mersley Road, 33.81038uS 119.787uE, 215 m, brown spots; scutellum pale with brown 06 Dec 1997, Schuh, Cassis, Brailovsky, spots, apex dark; hemelytron unicolorous Asquith, Beaufortia micrantha Schauer (Myr- pale with small brown spots; markings on taceae), det. Perth 05055482, 1- (AMNH_ cuneus present as small dark dots; membrane PBI 00371069) (WAMP). veins white or pale; hind femur with many PARATYPES: AUSTRALIA: Western Aus- brown or black spots; hind tibial spines black tralia: Fitzgerald River National Park, Ham- with conspicuous dark spots at bases. Abdo- mersley Road, 33.81038uS 119.787uE, 215 m, men: Venter unicolorous pale or mostly so. 06 Dec 1997, Schuh, Cassis, Brailovsky, SURFACE AND VESTITURE: Dorsum Asquith, Beaufortia micrantha Schauer (Myr- with suberect or reclining black setae and taceae), det. Perth 05055482, 10- (00371063, some sericeous or woolly setae. STRUC- 00371065–00371068, 00371072–00371075, 00- TURE: Head: Somewhat projecting; intero- 371077), 5U (00371082, 00371084–00371085, cular space moderate; eyes leaving gena 00371087, 00371095) (AM), 7- (00087473, moderately exposed in lateral view. Antenna: 00371064, 00371070–00371071, 00371076, Segment 2 weakly tapering, more slender at 00371078–00371079), 6U (00371083, 00371086, base; antennal fossa with ventral margin at 00371088–00371089, 00371094, 00371097) ventral margin of eye. Labium: Just reaching (AMNH), 1- (00371080), 1U (00371098) onto abdomen. Thorax: Pretarsus: Pulvilli (UNSW), 3- (00087247–00087248, 00371081), present, flaplike, covering about one third of 4U (00371090, 00371092–00371093, 00371096) ventral claw surface. GENITALIA (fig. 16): (WAMP). Endosoma: All available specimens teneral OTHER SPECIMENS EXAMINED: AUSTRA- and therefore not observed. Phallotheca: LIA: Western Australia: Fitzgerald River Nearly erect; dorsal surface with a curving, National Park, Hammersley Road, posteriorly directed, platelike projection; 33.81038uS 119.787uE, 215 m, 06 Dec 1997, apex simple. Left Paramere: Shaft at right Schuh, Cassis, Brailovsky, Asquith, Beaufor- angles to body; body broad basally, apex tia micrantha Schauer (Myrtaceae), det. Perth medially drawn into a blunt fingerlike 05055482, 1 nymph (00371091) (AM). process; anterior process triangular in lateral view; posterior process in the form of a long, straight, fingerlike projection; base of poste- Melaleucoides brevifoliae, new species rior process without conspicuous shoulder. Figure 16, map 3, plate 2 Right Paramere: Body short and broad, DIAGNOSIS: Recognized by the presence of lanceolate; apex short, clawlike, decurved. numerous tiny dark spots on the weakly Female: Slightly more ovoid than male; greenish dorsum (pl. 2), and the structure of mean total length 3.70, mean width prono- 2010 SCHUH AND WEIRAUCH: MYRTACEAE-FEEDING PHYLINAE 39

Fig. 16. Melaleucoides brevifoliae, male genitalia (AMNH_PBI 00130428).

tum 1.36. COLORATION (pl. 2): As in (Myrtaceae), det. Det: Royal Bot. male. GENITALIA: Not examined. Gard. NSW NSW427362, 1- (00130421) ETYMOLOGY: Named for the host species, (SAMA). Melaleuca brevifolia (Myrtaceae). PARATYPES: AUSTRALIA: South Austra- HOSTS: Melaleuca brevifolia Turcz. (Myr- lia: Scorpion Springs Cons. Park, 35.60421uS taceae). 140.8646uE, 125 m, 10 Nov 1998, Schuh, DISTRIBUTION (map 3): Recorded from Cassis, Silveira, Melaleuca brevifolia Turcz. one locality on the eastern border of southern (Myrtaceae), det. Det: Royal Bot. Gard. South Australia. Melaleucoides brevifoliae is NSW NSW427362, 9- (00087236, 00371099– one of only two species in the Melaleucoides 00371102, 00371121–00371124), 6U (00371118– genus group that is known to occur outside 00371120, 00371130–00371132) (AM), 12- Western Australia. (00087506, 00130422–00130432), 9U (00087- DISCUSSION: All available male specimens 237, 00130433–00130440) (AMNH), 1- for this species were apparently partially (00371103), 1U (00371117) (SAMA). teneral, because after several dissections we OTHER SPECIMENS EXAMINED: AUSTRA- were unable to find a specimen in which the LIA: South Australia: Scorpion Springs Cons. endosoma was suitably sclerotized for obser- Park, 35.60421uS 140.8646uE, 125 m, 10 Nov vation and illustration. 1998, Schuh, Cassis, Silveira, Melaleuca HOLOTYPE: AUSTRALIA: South Austra- brevifolia Turcz. (Myrtaceae), det. Det: Royal lia: Scorpion Springs Cons. Park, 35.60421uS Bot. Gard. NSW NSW427362, 18 nymphs 140.8646uE, 125 m, 10 Nov 1998, Schuh, (00371104–00371116, 00371125–00371129) Cassis, Silveira, Melaleuca brevifolia Turcz. (AM). 40 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 344

Melaleucoides cassisi, new species strap distal to gonopore ornamented with Figures 17, 18; map 3; plate 2 denticles at about midpoint of apical region of primary strap; spinelike, elongate process DIAGNOSIS: Recognized among Melaleu- arising near gonopore on dorsal surface coides spp. by the relatively large size (mean absent; spinelike, elongate process near gon- total length 3.95), the monotonous pale opore on ventral surface present, strongly coloration of the dorsum, and the structure recurved toward base of endosoma; bladder- of the male genitalia, notably the clublike like process distad of secondary gonopore aggregation of denticles distad of the second- absent; secondary gonopore seen frontally ary gonopore and their position ventral to (facing up) in lateral view of endosoma. the primary endosomal strap. Most similar in Phallotheca: More or less right angulate; size and coloration to M. grossi and M. dorsal surface with a curving, posteriorly sheathiana. Distinguished from the former by directed, platelike projection; apex simple. the arrangement of the endosomal denticles Left Paramere: Shaft at right angles to body; on an elongate, conical, free spine ventral to body spoon shaped; apex medially drawn the primary endosomal spine, and from the into a blunt, recurved, fingerlike process; latter by the strongly orange scutellum and anterior process triangular in lateral view; the dorsal position of the endosomal spicules posterior process in the form of a short, and the fusion of the primary and secondary straight, fingerlike projection; base of poste- endosomal straps distad of the secondary rior process without conspicuous shoulder. gonopore in that species. Right Paramere: Body short and broad, DESCRIPTION: Male: Body weakly elon- lanceolate; apex short, clawlike, decurved. gate, weakly ovoid; mean total length 3.95, Female: Distinctly more ovoid than mean width pronotum 1.38. COLORATION male; mean total length 4.12, mean width (pl. 2): Head: Uniformly pale, faded yellow; pronotum 1.53. COLORATION (pl. 2): scapus and pedicellus yellow to dirty yellow; As in male. GENITALIA (fig. 18): Poste- labium pale with segment 4 heavily infuscate. rior wall laterally with distinct crescent- Thorax: Pronotum unicolorous, faded yel- shaped interramal sclerites; posteriorly low; scutellum white; hemelytron unicolor- without a sclerotized transverse band; ous, faded yellow; cuneus mostly white, posterolaterally with a distinct swelling markings absent; membrane and veins weak- covered with microtrichia; longitudinal ly fumose; hind femur unicolorous pale, fold on either side of midline along part without black spots; hind tibial spines black, of length; posterior margin of posterior without dark spots at bases. Abdomen: wall without spicules and not reflexed Venter unicolorous pale or mostly so. SUR- dorsally; interramal lobes present and FACE AND VESTITURE (fig. 17B): Dor- asymmetrical, reclining, overlapping, and sum with suberect or reclining black setae heavily ornamented with spicules; vestibu- and some sericeous or woolly setae. STRUC- lum with medial plates sclerotized and TURE: Head (fig. 17A): Barely projecting; readily observed, small, nearly symmetri- interocular space relatively large; eyes leaving cal, triangular; with sclerotized guide gena moderately exposed in lateral view. present as short, heavily sclerotized tube Antenna: Segment 2 of equal diameter over on left side. entire length; antennal fossa with ventral ETYMOLOGY: Named for Gerasimos Cas- margin at ventral margin of eye. Labium: Just sis, who collected most of the known reaching onto abdomen. Thorax: Pretarsus specimens, in recognition of his ground- (fig. 17C, D): Pulvilli present, flaplike, cov- breaking efforts in field documentation and ering about one third of ventral claw surface. study of Australian Heteroptera. GENITALIA (figs. 17E, F, 18): Endosoma: HOSTS: Melaleuca uncinata R. Br. (Myrta- Primary strap apically greatly elongate, ceae). weakly arcuate and curving ventrally, with- DISTRIBUTION (map 3): Known from two out ornamentation; secondary endosomal localities west and north of Yalgoo and in the strap fused with primary strap proximal to northeastern Wheatbelt region of Western secondary gonopore, fused with primary Australia. 2010 SCHUH AND WEIRAUCH: MYRTACEAE-FEEDING PHYLINAE 41

Fig. 17. Melaleucoides cassisi, male. Scanning electron micrographs. A. Head and thorax, lateral view. B. Setae on hemelytra. C. Pretarsus, frontal view. D. Pretarsus, frontal view, detail of rudimentary dorsal arolium. E. Pygophore, lateral view. F. Pygophore, dorsal view. da 5 dorsal arolium; lp 5 left paramere; pe 5 parempodium; ph 5 phallotheca; pv 5 pulvillus; rp 5 right paramere (AMNH_PBI 00371141).

HOLOTYPE: AUSTRALIA: Western Aus- ceae), det. Perth 05120640, 1- (AMNH_PBI tralia: 56.6 km W of Yalgoo, 28.42397uS 00371607) (WAMP). 116.1233uE, 600 m, 27 Oct 1996, Schuh and PARATYPES: AUSTRALIA: Western Aus- Cassis, Melaleuca uncinata R. Br. (Myrta- tralia: 56.6 km W of Yalgoo, 28.42397uS 42 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 344

Fig. 18. Melaleucoides cassisi, male and female genitalia (AMNH_PBI 00371147, 00371699). 2010 SCHUH AND WEIRAUCH: MYRTACEAE-FEEDING PHYLINAE 43

116.1233uE, 600 m, 27 Oct 1996, Schuh and strongly fumose; hind femur unicolorous Cassis, Melaleuca uncinata R. Br. (Myrta- pale, without black spots; hind tibial spines ceae), det. Perth 05120640, 10- (00087172, black, without dark spots at bases. Abdomen: 00371142–00371150), 33U (00087173, 003- Venter unicolorous dark. SURFACE AND 71151–00371182) (AM), 24- (00087452, 003- VESTITURE: Dorsum with suberect or 71608–00371610, 00371612–00371630, 0037- reclining black setae, without sericeous or 1639), 31U (00371565–00371593, 0037- woolly setae. STRUCTURE: Head: Weakly 1611, 00371699) (AMNH), 1- (00371631) projecting; interocular space relatively large; (UCR), 1- (00371056), 6U (00371057– eyes leaving gena broadly exposed in lateral 00371062) (UNSW), 6- (00371633– view. Antenna: Segment 2 weakly tapering, 00371638), 21U (00371183–00371191, more slender at base; antennal fossa with 00371594–00371605) (WAMP). ventral margin at ventral margin of eye. OTHER SPECIMENS EXAMINED: AUSTRA- Labium: Just reaching onto abdomen. Tho- LIA: Western Australia: 13.5 km W of rax: Pretarsus: Pulvilli present, flaplike, Nungarin on Rt 50, 31.11547uS 117.945uE, covering about one third of ventral claw 300 m, 16 Nov 1999, R.T. Schuh, G. Cassis, surface. GENITALIA (fig. 19): Endosoma: & R. Silveira, 1- (00129374) (UNSW). Primary strap apically greatly elongate, 56.6 km W of Yalgoo, 28.42397uS nearly straight, apex spadelike, otherwise 116.1233uE, 600 m, 27 Oct 1996, Schuh and without ornamentation; secondary endoso- Cassis, Melaleuca uncinata R. Br. (Myrta- mal strap fused with primary strap proximal ceae), det. Perth 05120640, 9- (00371133– to secondary gonopore, reaching well beyond 00371141) (AM), 41U (00371641–00371661, gonopore, fused with primary strap, orna- 00371663–00371682), 1 nymph (00371662) mented with denticles; spinelike, elongate (AMNH), 1- (00371632), 4U (00371687– process arising near gonopore on dorsal 00371690) (CNC), 4U (00371683–00371686) surface absent; spinelike, elongate process (UCR), 1- (00371640), 4U (00371695–003- near gonopore on ventral surface present and 71698) (USNM), 1- (00371606), 4U (003- erect, parallel to primary endosomal strap; 71691–00371694) (ZISP). bladderlike process distad of secondary gonopore absent; secondary gonopore seen Melaleucoides castanea, new species frontally (facing up) in lateral view of Figure 19, map 3, plate 2 endosoma. Phallotheca: Nearly erect; dorsal surface with a curving, posteriorly directed, DIAGNOSIS: Recognized uniquely among platelike projection; ventral surface with a Melaleucoides spp. by the entirely castaneous projecting keel; apex simple. Left Paramere: coloration, the large, broad body (pl. 2), and Shaft at right angles to body; body spoon the structure of the male genitalia, with the shaped; apex medially elongate and broadly left paramere elongate but broad apically, rounded; anterior process triangular in later- and the endosoma with a long, apically al view; posterior process in the form of a oriented, ventral process associated with the short, straight, fingerlike projection; base of secondary gonopore. Coloration most simi- posterior process without conspicuous shoul- lar to M. rhaphiophyllae and M. uncinatae, der. Right Paramere: Body short and broad, but both of those species with the apex of lanceolate; apex with short fingerlike process. the scutellum and the base of the cuneus Female: Body shape as in male; mean total pale. length 3.84, mean width pronotum 1.55. DESCRIPTION: Male: Body broadly ovoid; COLORATION (pl. 2): As in male. GENI- mean total length 3.61, mean width prono- TALIA: Not examined. tum 1.44. COLORATION (pl. 2): Head: ETYMOLOGY: Named for the dark color- Uniformly castaneous; scapus unicolorous ation, from the Latin castanea, ‘‘chestnut.’’ dark; pedicellus yellow to dirty yellow; HOSTS: Melaleuca undulata Benth. (Myr- labium pale with segment 4 heavily infuscate. taceae). Thorax: Pronotum, scutellum, and hemely- DISTRIBUTION (map 3): Recorded from tron unicolorous castaneous; markings on one locality in southwestern Western Aus- cuneus absent; membrane and veins very tralia. 44 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 344

Fig. 19. Melaleucoides castanea, male genitalia (AMNH_PBI 00087372).

HOLOTYPE: AUSTRALIA: Western Aus- 1U (00372754) (AM), 1U (00327133) tralia: 4 km W of Lillian Stoke Rock, (AMNH), 1U (00087373) (WAMP). 33.079uS 120.0669uE, 370 m, 21 Nov 1999, R.T. Schuh and G. Cassis, Melaleuca un- Melaleucoides grossi, new species dulata Benth. (Myrtaceae), det. Perth Figure 20, map 4, plate 2 - 05671930, 1 (AMNH_PBI 00087372) DIAGNOSIS: Recognized among Melaleu- (WAMP). coides spp. by the relatively large size (mean PARATYPES: AUSTRALIA: Western Aus- total length 3.34), the monotonous pale tralia: 4 km W of Lillian Stoke Rock, coloration of the dorsum, and the structure 33.079uS 120.0669uE, 370 m, 21 Nov 1999, of the male genitalia, notably the secondary R.T. Schuh and G. Cassis, Melaleuca undu- endosomal strap dorsal to the primary strap, lata Benth. (Myrtaceae), det. Perth 05671930, free, elongate, conical, and uniformly covered 2010 SCHUH AND WEIRAUCH: MYRTACEAE-FEEDING PHYLINAE 45 with short denticles. Most similar in size and a curving, posteriorly directed, platelike coloration to M. cassisi and M. sheathiana. projection; anterior surface with a short to Distinguished from the former by the ar- elongate keel; apex simple. Left Paramere: rangement of the endosomal denticles in a Shaft at right angles to body; body spoon clublike fashion ventral to the primary shaped; apex medially broadly rounded; endosomal strap, and from the latter by the anterior process triangular in lateral view; strongly orange scutellum and the fusion of posterior process in the form of a long, the primary and secondary endosomal straps straight, fingerlike projection; base of poste- distad of the secondary gonopore with the rior process without conspicuous shoulder. spicules placed in somewhat irregular ar- Right Paramere: Body short and broad, rangement. lanceolate; apex with short fingerlike process. DESCRIPTION: Male: Body weakly to Female: Body shape as in male; mean total distinctly ovoid; mean total length 3.34, mean length 3.75, mean width pronotum 1.46. width pronotum 1.38. COLORATION COLORATION (pl. 2): As in male. GENI- (pl. 2): Head: Uniformly pale, weakly yellow; TALIA: Not examined. scapus and pedicellus yellow to dirty yellow; ETYMOLOGY: Named for the collector of labium pale with segment 4 heavily infuscate. all known specimens, the late Gordon Gross, Thorax: Pronotum unicolorous, weakly yel- in recognition of his contributions to Aus- low; scutellum white to weakly yellow; tralian heteropterology. hemelytron, including cuneus, unicolorous HOSTS: Melaleuca sp. (Myrtaceae). weakly yellow; markings on cuneus absent; DISTRIBUTION (map 4): Known from membrane weakly fumose, veins entirely red; Wallaroo in southeastern South Australia. hind femur unicolorous pale, without black Melaleucoides grossi is one of only two spots; hind tibial spines black, without dark species in the Melaleucoides genus group that spots at bases. Abdomen: Venter unicolorous occur outside Western Australia. pale or mostly so. SURFACE AND VESTI- HOLOTYPE: AUSTRALIA: South Austra- TURE: Dorsum with suberect or reclining lia: North Beach, Wallaroo, 33.906uS black setae and some sericeous or woolly 137.63uE, 12 Feb 1964, G.F. Gross, 1- setae. STRUCTURE: Head: Barely project- (AMNH_PBI 00169227) (SAMA). ing; interocular space moderate; eyes leaving PARATYPES: AUSTRALIA: South Austra- gena moderately exposed in lateral view. lia: North Beach, Wallaroo, 33.906uS Antenna: Segment 2 of equal diameter over 137.63uE, 12 Feb 1964, G.F. Gross, 2- entire length; antennal fossa with ventral (00169239, 00169240), 3U (00169256–0016- margin at ventral margin of eye. Labium: Just 9258) (AM), 4- (00169219–00169220, 0016- reaching onto abdomen. Thorax: Pretarsus: 9225, 00169238), 5U (00169215, 00169217, Pulvilli present, flaplike, covering about one 00169246, 00169249, 00169253) (AMNH), third of ventral claw surface. GENITALIA 15- (00169221–00169224, 00169226, 001- (fig. 20): Endosoma: Primary strap apically 69228–00169237), 17U (00169209–00169212, 00- greatly elongate, weakly angled dorsally 169216, 00169218, 00169241, 00169243–001- relative to body of endosoma, no ornamen- 69245, 00169247–00169248, 00169250–001- tation; secondary endosomal strap fused with 69252, 00169254–00169255) (SAMA). primary strap proximal to secondary gono- pore, free from primary strap distal to Melaleucoides leuropomae, new species secondary gonopore, ornamented with den- Figure 21, map 4, plate 2 ticles; spinelike, elongate process arising near gonopore on dorsal surface absent; spinelike, DIAGNOSIS: Recognized by the presence of elongate process near gonopore on ventral discrete small dark spots on the dorsum surface present, recurved toward base of (pl. 2), and the structure of the male genita- endosoma; bladderlike process distad of lia, the left paramere with a somewhat secondary gonopore absent; secondary gon- elongate, apex positioned more or less opore seen frontally (facing up) in lateral symmetrically and the posterolateral shoul- view of endosoma. Phallotheca: Smoothly der with a low projection; endosoma with a curving on dorsal margin; dorsal surface with long row of heavy, elongate denticles on the 46 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 344

Fig. 20. Melaleucoides grossi, male genitalia (AMNH_PBI 00169233).

postgonoporal portion of the secondary strap DESCRIPTION: Male: Body weakly elon- and recurved ventral process associated with gate, weakly ovoid; mean total length 3.25, the secondary gonopore. Most easily con- mean width pronotum 1.25. COLORATION fused with M. micranthae on the form of the (pl. 2): Head: Pale with heavy dark dots; spots and the structure of the male genitalia, scapus unicolorous pale; pedicellus dirty but the spots in that species sometimes larger, yellow proximally, weakly to heavily infus- the apical medial process of the left paramere cate distally; labium pale with segment 4 more nearly symmetrically placed, and the heavily infuscate. Thorax: Pronotum pale row of denticles on the secondary endosomal with heavy dark spots; scutellum pale with strap shorter a and in the form of a more brown spots; hemelytron unicolorous pale condense grouping. Spots on the dorsum of with scattered dark spots; markings on M. beaufortiae always somewhat coalescent cuneus present as weakly darkened apex; and those in M. brevifoliae tiny and much membrane weakly fumose, veins pale; hind more numerous. femur with many brown or black spots; hind 2010 SCHUH AND WEIRAUCH: MYRTACEAE-FEEDING PHYLINAE 47

Fig. 21. Melaleucoides leuropomae, male genitalia (AMNH_PBI 00137076).

tibial spines black with conspicuous dark surface. GENITALIA (fig. 21): Endosoma: spots at bases. Abdomen: Venter unicolorous Primary strap apically greatly elongate, pale or mostly so. SURFACE AND VESTI- without ornamentation, extreme apex curv- TURE: Dorsum with suberect or reclining ing dorsally; secondary endosomal strap black setae, without sericeous or woolly fused with primary strap proximal to sec- setae. STRUCTURE: Head: Weakly project- ondary gonopore, reaching well beyond ing; interocular space relatively large; eyes gonopore, with a comblike edge; spinelike, leaving gena broadly exposed in lateral view. elongate process arising near gonopore on Antenna: Segment 2 weakly tapering, more dorsal surface absent; spinelike, elongate slender at base; antennal fossa with dorsal process near gonopore on ventral surface margin somewhat below ventral margin of present, strongly recurved toward base of eye. Labium: Just reaching onto abdomen. endosoma; bladderlike process distad of Thorax: Pretarsus: Pulvilli present, flaplike, secondary gonopore absent; secondary gon- covering about one third of ventral claw opore seen frontally (facing up) in later- 48 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 344

al view of endosoma. Phallotheca: Nearly PARATYPES: AUSTRALIA: Western Aus- erect; anterior surface with a short to tralia: 15 km E of Merredin, 31.37749uS elongate keel; apex simple. Left Paramere: 118.6933uE, 330 m, 16 Nov 1999, R.T. Schuh Shaft at right angles to body; body more or and G. Cassis, Melaleuca conothamnoides less quadrate; apex medially drawn into a C.A.Gardner (Myrtaceae), det. PERTH staff blunt fingerlike process; apex posterolaterally PERTH 05670624, 3;m (AMNH_PBI 000- with a low shoulder; anterior process trian- 87325, AMNH_PBI 00087534, AMNH_ gular in lateral view; posterior process in the PBI 00372261), 4;f (AMNH_PBI 00087326, form of a short, straight, fingerlike projec- AMNH_PBI 00372262 - AMNH_PBI 003- tion; base of posterior process without 72264) (AM). Brand Hiway 45.9 km S of conspicuous shoulder. Right Paramere: Body Dongarra Road, 29.57703uS 115.1348uE, short and broad, lanceolate; apex short, claw- 100 m, 31 Oct 1996, Schuh and Cassis, Phy- like, decurved. matocarpus porphyrocephalus F. Muell. (Myr- Female: Slightly more ovoid than male; taceae), det. PERTH staff PERTH 05879264, mean total length 3.46, mean width prono- 1;m (AMNH_PBI 00087192), 1;f (AMNH_ tum 1.32. COLORATION (pl. 2): As in PBI 00087193) (AM), 2;m (AMNH_PBI male. GENITALIA: Posterior wall laterally 00130199, AMNH_PBI 00130200), 1;f with distinct crescent-shaped interramal (AMNH_PBI 00130201) (AMNH). Cer- sclerites; posteriorly without a sclerotized vantes, 30.49902uS 115.0684uE, 3 m, 10 Dec transverse band; posterolaterally with a 1997, Schuh, Brailovsky, Melaleuca vimi- distinct swelling covered with microtrichia; nea Lindl. (Myrtaceae), det. PERTH staff longitudinal fold on either side of midline PERTH 05879205, 1;f (AMNH_PBI 00181- along part of length; posterior margin of 857) (AMNH), 3;f (AMNH_PBI 001305- posterior wall without spicules and not 52, AMNH_PBI 00181855 - AMNH_PBI reflexed dorsally; interramal lobes present 00181856) (WAMP). Kalbarri National and asymmetrical, reclining, overlapping, Park, 7 km E of Kalbarri, 27.68008uS and heavily ornamented with spicules; ves- 114.2386uE,400m,29Oct1996,Schuh tibulum with medial plates sclerotized and and Cassis, Melaleuca leuropoma L. A. readily observed, small, nearly symmetrical, Craven (Myrtaceae), det. PERTH staff triangular, with sclerotized guide present as PERTH 05879175, 8;m (AMNH_PBI 001- short heavily sclerotized tube on left side. 37073 - AMNH_PBI 00137074, AMNH_PBI ETYMOLOGY: Named for the host species, 00137076 - AMNH_PBI 00137081), 16;f Melaleuca leuropoma (Myrtaceae). (AMNH_PBI 00137082 - AMNH_PBI 001- HOSTS: Most known specimens were 37097) (AMNH), 1;m (AMNH_PBI 001- taken on Melaleuca leuropoma L.A. Craven 30202), 3;f (AMNH_PBI 00137098, AMNH_ (Myrtaceae) (pl. 5C). Additional speci- PBI 00137320 - AMNH_PBI 00137321) mens are recorded from the following species (WAMP). Kalbarri National Park, 22.9 km of Myrtaceae: Melaleuca conothamnoides E Kalbarri, 27.75408uS 114.3711uE, 500 m, 29 C.A. Gardner, Melaleuca viminea Lindl., Oct 1996, Schuh and Cassis, 3;m (AMNH_ and Phymatocarpus porphyrocephalus F. PBI 00135363 - AMNH_PBI 00135365), 1;f Muell. (AMNH_PBI 00135366) (WAMP). DISTRIBUTION (map 4): Known from sev- eral localities in Western Australia, including Melaleucoides micranthae, new species one locality in the Wheatbelt region and Figure 22, map 4, plate 3 coastal areas north to Kalbarri. HOLOTYPE: AUSTRALIA: Western Aus- DIAGNOSIS: Recognized by the presence of tralia: Kalbarri National Park, 7 km E of discrete small dark spots on the dorsum Kalbarri, 27.68008uS 114.2386uE, 400 m, 29 (pl. 3), and the structure of the male genita- Oct 1996, Schuh and Cassis, Melaleuca lia, the left paramere with a somewhat leuropoma L.A. Craven (Myrtaceae), det. elongate, apex positioned somewhat asym- Perth 05879175, 1- (AMNH_PBI 00137075) metrically and the posterolateral shoulder (WAMP). with a low projection; endosoma with a 2010 SCHUH AND WEIRAUCH: MYRTACEAE-FEEDING PHYLINAE 49

Fig. 22. Melaleucoides micranthae, male genitalia (AMNH_PBI 00372253). weakly elongate, condensed grouping of always somewhat coalescent and those in heavy denticles on the postgonoporal portion M. brevifoliae tiny and much more numer- of the secondary strap and recurved ventral ous. process associated with the secondary gono- DESCRIPTION: Male: Body weakly to pore. Most easily confused with M. leuropo- distinctly ovoid; mean total length 3.14, mean mae on the form of the spots and the width pronotum 1.22. COLORATION structure of the male genitalia, but the spots (pl. 3): Head: Pale with a few dark dots; in that species usually smaller, the apical scapus yellow to dirty yellow; pedicellus medial process of the left paramere more yellow to dirty yellow; labium uniformly strongly asymmetrical, and the row of pale. Thorax: Pronotum pale, yellowish, with denticles on the secondary endosomal strap a few brown spots; scutellum pale, yellowish; longer, with the denticles more evenly spaced. hemelytron unicolorous pale, yellowish, with Spots on the dorsum of M. beaufortiae small a few scattered dark spots; markings on 50 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 344

cuneus present as a weakly castaneous apex; DISTRIBUTION (map 4): Known from two membrane weakly fumose, veins yellow; hind localities in the Boorabbin and Wheatbelt femur with many brown or black spots; hind regions in Western Australia. tibial spines dark with very small dark bases. HOLOTYPE: AUSTRALIA: Western Aus- Abdomen: Venter unicolorous pale or mostly tralia: 32 km SE of Paynes Find toward so. SURFACE AND VESTITURE: Dorsum Beacon, 29.48558uS 117.7836uE, 250 m, 12 with suberect or reclining black setae, with- Dec 1997, Schuh, Brailovsky, 1- (AMNH_ out sericeous or woolly setae. STRUC- PBI 00131081) (WAMP). TURE: Head: Barely projecting; interocular PARATYPES: AUSTRALIA: Western Aus- space moderate; eyes leaving gena broadly tralia: 32 km SE of Paynes Find toward exposed in lateral view. Antenna: Segment 2 Beacon, 29.48558uS 117.7836uE, 250 m, 12 weakly tapering, more slender at base; Dec 1997, Schuh, Brailovsky, 1- (00131082), antennal fossa with dorsal margin somewhat 5U (00131083–00131087) (AMNH). 92.5 km below ventral margin of eye. Labium: Just W of Coolgardie at east side of Boorab- reaching onto abdomen. Thorax: Pretarsus: bin National Park on Great Eastern Hwy, Pulvilli present, flaplike, covering about one 31.21233uS 120.31uE, 445 m, 17 Nov 1999, third of ventral claw surface. GENITALIA R.T. Schuh and G. Cassis, Beaufortia mi- (fig. 22): Endosoma: Primary strap apically crantha micrantha Schauer (Myrtaceae), det. greatly elongate, curving dorsally near apex, Perth 05671957, 5- (00087320, 00372252, no ornamentation; secondary endosomal 00372255–00372257), 5U (00087321, 003722- strap fused with primary strap proximal to 65–00372268) (AM), 1- (00087530) (AMNH), secondary gonopore, reaching well beyond 2- (00372253, 00372254), 2U (00372269, gonopore, fused with primary strap, and with 00372270) (WAMP). a few heavy denticles; spinelike, elongate OTHER SPECIMENS EXAMINED: AUSTRA- process arising near gonopore on dorsal LIA: Western Australia: 92.5 km W of Cool- surface absent; spinelike, elongate process gardie at east side of Boorabbin National near gonopore on ventral surface present, Park on Great Eastern Hwy, 31.21233uS strongly recurved toward base of endosoma; 120.31uE, 445 m, 17 Nov 1999, R.T. Schuh bladderlike process distad of secondary and G. Cassis, Beaufortia micrantha mi- gonopore absent; secondary gonopore seen crantha Schauer (Myrtaceae), det. Perth frontally (facing up) in lateral view of 05671957, 10 nymphs (00372271–00372280) endosoma. Phallotheca: Smoothly curving (AM). on dorsal margin; anterior surface with a short to elongate keel; apex simple. Left Melaleucoides ozzii, new species Paramere: Shaft at right angles to body; body Figure 23, map 4, plate 3 spoon shaped, apex medially drawn into a blunt fingerlike process; apex posterolaterally DIAGNOSIS: Similar in size and coloration with a low shoulder; anterior process slender to Melaleucoides pileanthicola, the total and cylindrical in lateral view; posterior length not exceeding 3.01, and the strongly process in the form of a short, curved, reddish clavus and posterior portion of the fingerlike projection; base of posterior pro- endocorium. Also similar to M. akaina in size cess without conspicuous shoulder. Right and general coloration. Distinguished from Paramere: Body short and broad, lanceolate; M. akaina and M. pileanthicola by the pale apex short, clawlike, decurved. coloration of the cuneus in both of those Female: Slightly more ovoid than male; species; distinguished from M. pileanthicola mean total length 3.53, mean width prono- by the strongly reddish clavus and posterior tum 1.16. COLORATION (pl. 3): As in portion of the endocorium in that species as male. GENITALIA: Not examined. well as the structure of the endosoma. ETYMOLOGY: Named for the host species, Further distinguished from M. akaina by Beaufortia micrantha (Myrtaceae). short apical endosomal spine and the com- HOSTS: Beaufortia micrantha micrantha plete absence of processes associated with the Schauer (Myrtaceae). secondary gonopore in that species. 2010 SCHUH AND WEIRAUCH: MYRTACEAE-FEEDING PHYLINAE 51

Fig. 23. Melaleucoides ozzii, male genitalia (AMNH_PBI 00371237).

DESCRIPTION: Male: Body moderately absent. GENITALIA (fig. 23): Endosoma: elongate, parallel sided; mean total length Primary strap relatively short, straight, with 2.90, mean width pronotum 1.02. COLOR- a bifid apex; secondary endosomal strap free ATION (pl. 3): Head: Uniformly pale; sca- of primary strap and much longer, orna- pus and pedicellus unicolorous pale; labium mented with denticles near apex; spinelike, pale with segment 4 heavily infuscate. Tho- elongate process arising near gonopore on rax: Pronotum pale with red or carmine dorsal surface present, erect, unornamented; blotches; scutellum white; hemelytron pale secondary gonopore seen laterally in lateral yellow with red suffusion; markings on view of endosoma. Phallotheca: Nearly erect; cuneus present as a contrasting white basal dorsal surface without a fingerlike projec- fascia, remainder red; membrane weakly tion, and without a platelike projection; fumose, veins white or pale; hind femur anterior surface with a short to elongate keel; unicolorous pale, without black spots; tibial ventral surface with a projecting keel; apex spines with very small dark bases. Abdomen: simple. Left Paramere: Shaft at right angles Venter unicolorous pale or mostly so. SUR- to body; body spoon shaped; apex medially FACE AND VESTITURE: Dorsum with broadly rounded to weakly quadrate; anteri- reclining simple setae matching background or process slender and cylindrical in lateral coloration, without sericeous or woolly setae. view; posterior process in the form of a short, STRUCTURE: Head: Moderately project- slender, straight, fingerlike projection; base ing; interocular space large; eyes leaving gena of posterior process without conspicuous moderately exposed in lateral view. Antenna: shoulder. Right Paramere: Body short and Segment 2 weakly tapering, more slender at broad, lanceolate; apex short, clawlike, de- base; antennal fossa with ventral margin at curved. ventral margin of eye. Labium: Just reaching Female: Slightly more ovoid than male; onto abdomen. Thorax: Pretarsus: Pulvilli mean total length 3.04, mean width prono- 52 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 344 tum 1.04. COLORATION (pl. 3): As in orange; scutellum entirely red to brown; male. GENITALIA: Not examined. hemelytron with most of endocorium car- ETYMOLOGY: Named for the late Ozzie mine; cuneus pale; membrane and veins Massie, with thanks for his encouragement strongly fumose; hind femur unicolorous and interest in our work on the Australian pale, without black spots; hind tibial spines Miridae fauna. black, without dark spots at bases. Abdomen: HOSTS: Thryptomene kochii E. Prinz. Venter unicolorous pale or mostly so. SUR- (Myrtaceae). FACE AND VESTITURE (fig. 24C): DISTRIBUTION (map 4): Known from one Dorsum with suberect or reclining black locality in the Goldfields region of Western setae and some sericeous or woolly setae. Australia. STRUCTURE: Head (fig. 24A): Moderately HOLOTYPE: AUSTRALIA: Western Aus- projecting; interocular space large; eyes tralia: 135 km W of Coolgardie on Great leaving gena only very slightly exposed in Eastern Hiway, 31.27202uS 120.0059uE, lateral view. Antenna: Segment 2 weakly 489 m, 17 Nov 1999, R.T. Schuh, G. Cassis, tapering, more slender at base; antennal fossa & R. Silveira, Thryptomene kochii E. Pritz. with ventral margin at ventral margin of eye. (Myrtaceae), det. Perth 05671302, 1- Labium: Just reaching onto abdomen. Tho- (AMNH_PBI 00371239) (WAMP). rax: Pretarsus (fig. 24D): Pulvilli absent. PARATYPES: AUSTRALIA: Western Aus- GENITALIA (figs. 24E, F, 25): Endosoma: tralia: 135 km W of Coolgardie on Great Primary strap apically greatly elongate, Eastern Hiway, 31.27202uS 120.0059uE, 489 m, weakly arcuate and curving dorsally, no 17 Nov 1999, R.T. Schuh, G. Cassis, & R. ornamentation; secondary endosomal strap Silveira, Thryptomene kochii E. Pritz. (Myrta- fused with primary strap proximal to sec- ceae), det. Perth 05671302, 4- (00371230, ondary gonopore, reaching well beyond 00371233, 00371236, 00371238), 5U (003- gonopore, devoid of ornamentation, free 71242, 00371245, 00371247–00371249) (AM), from primary endosomal strap; spinelike, 3- (00371231, 00371234, 00371237), 3U elongate process arising near gonopore on (00371241, 00371243–00371244) (AMNH), dorsal surface absent; spinelike, elongate 3- (00371229, 00371232, 00371235), 2U process near gonopore on ventral surface (00371240, 00371246) (WAMP). present and erect, oriented toward apex of endosoma; bladderlike process distad of secondary gonopore absent; secondary gon- Melaleucoides pileanthicola, new species opore seen frontally (facing up) in lateral Figures 24, 25; map 4; plate 3 view of endosoma. Phallotheca: Nearly erect; DIAGNOSIS: Similar in size and coloration dorsal surface with a curving, posteriorly to M. ozzii, the total length not exceeding directed, platelike projection; apex simple. 3.01 and the strongly reddish clavus and Left Paramere: Shaft at right angles to body; posterior portion of the endocorium. Also body spoon shaped; apex medially broadly; similar to M. akaina in size and general anterior process triangular in lateral view; coloration, but distinguished from that spe- posterior process in the form of a long, cies by the short apical endosomal spine and straight, fingerlike projection; base of poste- the absence of processes associated with the rior process with slightly projecting shoulder. secondary gonopore and from M. ozzii by the Right Paramere: Body short and broad, distinctive apical endosomal spine and red lanceolate; apex short, clawlike, decurved. cuneus in that species. Female: Body shape as in male; mean total DESCRIPTION: Male: Body weakly to length 2.94, mean width pronotum 1.01. distinctly ovoid; mean total length 2.72, mean COLORATION (pl. 3): As in male. GENI- width pronotum 0.97. COLORATION TALIA (fig. 25): Posterior wall laterally with (pl. 3): Head: Uniformly pale, yellow orange; distinct crescent-shaped interramal sclerites; scapus and pedicellus unicolorous pale; posteriorly without a sclerotized transverse labium pale with segment 4 heavily infuscate. band; posterolaterally with a distinct swelling Thorax: Pronotum unicolorous yellow covered with microtrichia; longitudinal fold 2010 SCHUH AND WEIRAUCH: MYRTACEAE-FEEDING PHYLINAE 53

Fig. 24. Melaleucoides pileanthicola, male. Scanning electron micrographs. A. Head, pro-, and mesothorax, lateral view. B. Meso- and metathorax, lateral view. C. Setae on hemelytra. D. Pretarsus, frontal view. E. Pygophore, lateral view. F. Pygophore, dorsal view. eva 5 evaporatory area; lp 5 left paramere; mssp 5 mesothoracic spiracle; pe 5 parempodium; ph 5 phallotheca; rp 5 right paramere (AMNH_PBI 00137348). on either side of midline along part of length; ornamented with spicules or with a very posterior margin of posterior wall without few; vestibulum with medial plates sclerotized spicules and not reflexed dorsally; interramal and readily observed, small, nearly symmet- lobes roughly symmetrical, erect, and not rical, triangular with sclerotized guide pre- 54 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 344

Fig. 25. Melaleucoides pileanthicola, male and female genitalia (AMNH_PBI 00137358, 00137396). sent as caplike structure with internal differ- mens are recorded from the following species entiation. of Myrtaceae: subsp. ETYMOLOGY: Named for the host genus, pilifer Keighery (pl. 7D), Pileanthus vernico- Pileanthus (Myrtaceae), in combination with sus F. Muell., Scholtzia leptantha Benth., the Latin suffix -cola, ‘‘inhabitant.’’ Thryptomene aspera aspera E. Pritz. (pl. 8B). HOSTS: The majority of known specimens DISTRIBUTION (map 4): Widespread in were taken on Pileanthus peduncularis bor- Western Australia, from the outback Gold- ealis Endl. (Myrtaceae). Additional speci- fields to the coastal Carnarvon region. 2010 SCHUH AND WEIRAUCH: MYRTACEAE-FEEDING PHYLINAE 55

HOLOTYPE: AUSTRALIA: Western Aus- Cassis, Wall, Weirauch, Symonds, Pileanthus tralia: NW Coastal Hiway 57 km N of Kal- vernicosus F. Muell. (Myrtaceae), det. Field barri Road, 27.44756uS 114.6867uE, 500 m, ID, 3- (00373116–00373118) (AM), 1- 30 Oct 1996, Schuh and Cassis, Pileanthus (00373299) (AMNH). 66.2 km E of North peduncularis borealis Endl. (Myrtaceae), det. West Coastal Hiway on Mardathuna Rd, Perth 05120349, 1- (AMNH_PBI 00137356) 24.45443uS 114.5233uE, 103 m, 01 Nov 2004, (WAMP). Cassis, Wall, Weirauch, Tatarnic, Symonds, PARATYPES: AUSTRALIA: Western Aus- Pileanthus peduncularis subsp. pilifer Keigh- tralia: 28 km S of Menzies (3.5 km E of ery (Myrtaceae), det. Perth 6989985, 2- Hiway), 29.91917uS 121.1514uE, 500 m, 25 (00373110, 00373111), 1U (00373112) (AM). Oct 1996, Schuh and Cassis, Thryptomene aspera aspera E. Pritz. (Myrtaceae), det. Melaleucoides rhaphiophyllae, new species Perth 05095093, 4- (00087219, 00087510, Figure 26, map 4, plate 3 00373108–00373109), 2U (00373106, 00373107) (AM). 54.3 km N of jct of Agana DIAGNOSIS: Most similar to M. uncinatae Kalbarri Rd and Brand Hiway (rest area), in the heavily castaneous coloration (pls. 3, 27.47362uS 114.7054uE, 240 m, 24 Oct 2004, 4) of the dorsum, but with the apex of the Cassis, Wall, Weirauch, Symonds, Pileanthus scutellum and the base of the cuneus pale. vernicosus F. Muell. (Myrtaceae), det. Field Distinguished from that species by the ID, 4U (00373119–00373122) (AM). Kalbarri different form of the phallotheca and the left National Park, Loop Road, 27.56163uS paramere (figs. 26, 31), the endosoma of M. 114.4376uE, 300 m, 28 Oct 1996, Schuh and rhaphiophyllae being unknown. Cassis, Pileanthus peduncularis borealis Endl. DESCRIPTION: Male: Body moderately (Myrtaceae), det. Perth 05120586, 7- (0037- elongate, parallel sided; mean total length 3147–00373153), 8U (00373156–00373163) 2.87, mean width pronotum 1.22. COLOR- (AM), 2- (00373154, 00373155) (WAMP). ATION (pl. 3): Head: Uniformly dark; sca- NW Coastal Hiway 57 km N of Kalbarri pus brown; pedicellus unicolorous black or Road, 27.44756uS 114.6867uE, 500 m, 30 Oct castaneous; labium generally infuscate, heavi- 1996, Schuh and Cassis, Pileanthus peduncu- ly so apically. Thorax: Pronotum unicolorous laris borealis Endl. (Myrtaceae), det. Perth deep red to black; scutellum red or dark 05120349, 11- (00087215, 00373123–00373- brown with white or pale tip; corium and 132), 9U (00087216, 00373136–00373143) clavus mostly castaneous, markings on cune- (AM), 29- (00087468, 00137322–00137324, us present as a contrasting white basal fascia, 00137329–00137353), 32U (00137359–001- remainder castaneous to black; membrane 37369, 00137373–00137376, 00137379– fumose, veins white; hind femur with many 00137380, 00137383–00137397) (AMNH), brown or black spots; hind tibial spines with 2- (00137354, 00137355), 2U (00137377, conspicuous dark spots at bases. Abdomen: 00137378) (CNC), 3- (00373133–00373135), Venter heavily red laterally. SURFACE 3U (00373144–00373146) (UNSW), 2- (00- AND VESTITURE: Dorsum with golden 137357, 00137358), 2U (00137381, 0013- reclining setae; dorsum without sericeous or 7382) (USNM), 4- (00137325–00137328), woolly setae. STRUCTURE: Head: Weakly 11U (00137370–00137372, 00137398–001- projecting; interocular space moderate; eyes 37405) (WAMP). leaving gena broadly exposed in lateral view. OTHER SPECIMENS EXAMINED: AUSTRA- Antenna: Segment 2 weakly tapering, more LIA: Western Australia: 4.5 km NW of jct of slender at base; antennal fossa with dorsal Blowholes Rd and North West Coastal Hi- margin somewhat below ventral margin of way, N of Carnarvon, 24.72267uS113.7158uE, eye. Labium: Just reaching onto abdomen. 28 m, 27 Oct 2004, Cassis, Wall, Weirauch, Thorax: Pretarsus: Pulvilli present, flaplike, Tatarnic, Symonds, Scholtzia leptantha covering about one third of ventral claw Benth. (Myrtaceae), det. Perth6988687, 1- surface. GENITALIA (fig. 26): Endosoma: (00373175) (AM). 54.3 km N of jct of Agana All available specimens teneral and therefore Kalbarri Rd and Brand Hiway (rest area), not observed. Phallotheca: Nearly erect; 27.47362uS 114.7054uE, 240 m, 24 Oct 2004, dorsal surface with a fingerlike projection; 56 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 344

Fig. 26. Melaleucoides rhaphiophyllae, male genitalia (AMNH_PBI 00372295). apex simple. Left Paramere: Shaft in same teneral, because after several dissections we axis as body; body spoon shaped; apex were unable to find a specimen in which the medially angulate; anterior process triangular endosoma was suitably sclerotized for obser- in lateral view; posterior process in the form vation and illustration. of a short, curved, fingerlike projection; base HOLOTYPE: AUSTRALIA: Western Aus- of posterior process with distinct shoulder. tralia: Brand Hiway 8.2 km N of Eneabba, Right Paramere: Body short and broad, 29.7462uS 115.254uE, 100 m, 31 Oct 1996, lanceolate; apex short, clawlike, decurved. Schuh and Cassis, Melaleuca rhaphiophylla Female: Slightly more ovoid than male; Schauer (Myrtaceae), det. Perth 05120195, mean total length 2.98, mean width prono- 1- (AMNH_PBI 00128826) (WAMP). tum 1.20. COLORATION (pl. 3): Paler than PARATYPES: AUSTRALIA: Western Aus- male. GENITALIA: Not examined. tralia: Brand Hiway 8.2 km N of Eneabba, ETYMOLOGY: Named for the host species, 29.7462uS 115.254uE, 100 m, 31 Oct 1996, Melaleuca rhaphiophylla (Myrtaceae). Schuh and Cassis, Melaleuca rhaphiophylla HOSTS: Melaleuca rhaphiophylla Schauer Schauer (Myrtaceae), det. Perth 05120195, (Myrtaceae) (pl. 5D); Melaleuca sp. 4- (00087265–00087266, 00128827, 00372296), DISTRIBUTION (map 4): Known from two 2U (00128828, 00372298) (AM), 2- (00372294, localities on the Carnamah Shire Coast 00372297) (AMNH), 1- (00372295), 1U (obscured on the map) and the eastern (00128829) (WAMP). agricultural region in Western Australia. OTHER SPECIMENS EXAMINED: AUSTRA- DISCUSSION: All available male specimens LIA: Western Australia: Moorine Rocks, for this species were apparently partially 11.7 km N of Great Eastern Hiway on 2010 SCHUH AND WEIRAUCH: MYRTACEAE-FEEDING PHYLINAE 57

Fig. 27. Melaleucoides sheathianae, male genitalia (AMNH_PBI 00371561).

Noongar Road, 31.22843uS 118.979uE, irregular arrangement. Most similar in size 345 m, 04 Dec 1997, Schuh, Cassis, Brai- and coloration to M. cassisi and M. grossi. lovsky, Asquith, Melaleuca sp. (Myrtaceae), Distinguished from the former by the ar- det. field ID; host 97-06, 1- (00371776), 1U rangement of the endosomal denticles in a (00371777) (AMNH). clublike fashion ventral to the primary endosoma strap, and from the latter by the secondary endosomal strap dorsal to the Melaleucoides sheathianae, new species primary strap, free, elongate, conical, and Figure 27, map 5, plate 3 uniformly covered with short denticles. DIAGNOSIS: Recognized among Melaleu- DESCRIPTION: Male: Body weakly elon- coides spp. by the relatively large size (mean gate, weakly ovoid; mean total length 3.67, total length 3.67), the monotonous pale mean width pronotum 1.37. COLORATION coloration of the dorsum in conjunction with (pl. 3): Head: Uniformly dirty yellow; scapus a conspicuously orange scutellum, and the unicolorous dark; pedicellus yellow to dirty structure of the male genitalia, with the yellow; labium pale with segment 4 heavily fusion of the primary and secondary endo- infuscate. Thorax: Pronotum unicolorous somal straps distad of the secondary gono- dirty yellow-green; scutellum entirely orange; pore and the spicules placed in somewhat hemelytron unicolorous yellow-green; mark- 58 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 344 ings on cuneus absent; membrane and veins reflexed dorsally; interramal lobes present weakly fumose; hind femur unicolorous pale, and asymmetrical, reclining, overlapping, without black spots; hind tibial spines black and heavily ornamented with spicules; ves- without dark spots at bases. Abdomen: tibulum with medial plates sclerotized and Venter unicolorous pale or mostly so. SUR- readily observed, small, nearly symmetrical, FACE AND VESTITURE: Dorsum with triangular, with sclerotized guide present as suberect or reclining black setae, without caplike structure with internal differentiation. sericeous or woolly setae. STRUCTURE: ETYMOLOGY: Named for the host species, Head: Barely projecting; interocular space Melaleuca sheathiana (Myrtaceae). moderate; eyes leaving gena moderately HOSTS: Melaleuca sheathiana W. Fitz., exposed in lateral view. Antenna: Segment 2 Melaleuca teuthidoides Barlow (Myrtaceae). weakly tapering, more slender at base; DISTRIBUTION (map 5): Recorded from antennal fossa with ventral margin at ventral several localities in the Goldfields region of margin of eye. Labium: Just reaching onto Western Australia. abdomen. Thorax: Pretarsus: Pulvilli present, HOLOTYPE: AUSTRALIA: Western Aus- flaplike, covering about one third of ventral tralia: 11 km N of Coolgardie-Esperance claw surface. GENITALIA (fig. 27): Endo- Hiway on Kambalda Road, 31.25231uS soma: Primary strap apically greatly elongate, 121.5899uE, 320 m, 18 Nov 1999, R.T. weakly arcuate and curving dorsally, no Schuh, G. Cassis, & R. Silveira, Melale- ornamentation; secondary endosomal strap uca sheathiana W. Fitzg. (Myrtaceae), det. fused with primary strap proximal to sec- Perth 05671396, 1- (AMNH_PBI 00371554) ondary gonopore, reaching well beyond (WAMP). gonopore, ornamented with denticles; spine- PARATYPES: AUSTRALIA: Western Aus- like, elongate process arising near gonopore tralia: 11 km N of Coolgardie-Esperance on dorsal surface absent; spinelike, elongate Hiway on Kambalda Road, 31.25231uS process near gonopore on ventral surface 121.5899uE, 320 m, 18 Nov 1999, R.T. present, recurved toward base of endosoma; Schuh, G. Cassis, & R. Silveira, Melaleuca bladderlike process distad of secondary sheathiana W.Fitzg. (Myrtaceae), det. Perth gonopore absent; secondary gonopore seen 05671396, 8- (00087301, 00087499, 0037- frontally (facing up) in lateral view of U endosoma. Phallotheca: Smoothly curving 1970–00371973, 00371975), 29 (00087302, on dorsal margin; dorsal surface with a 00371976, 00371978, 00371980–00371984, 00372031–00372035, 00372066–00372081) curving, posteriorly directed, platelike pro- - jection; apex simple. Left Paramere: Shaft at (AM), 17 (00129423–00129427, 00129504, 00129584–00129585, 00371552, 00371555– right angles to body; body spoon shaped; U apex medially broadly rounded; anterior 00371562), 36 (00129428–00129444, 00129- 505–00129512, 00129587–00129594, 003715- process triangular in lateral view; posterior U process in the form of a short, straight, 53, 00371563–00371564) (AMNH), 3 (00- fingerlike projection; base of posterior pro- 371985–00371987) (UNSW). 91.4 km SE of cess without conspicuous shoulder. Right Southern Cross, 31.97145uS 119.287uE, 375 m, Paramere: Body short and broad, lanceolate; 04 Dec 1997, Schuh, Cassis, Brailovsky, apex short, clawlike, decurved. Asquith, Melaleuca teuthidoides Barlow Female: Slightly more ovoid than male; (Myrtaceae), det. Perth 05054834, 6- (0013- mean total length 3.82, mean width prono- 0795–00130800), 6U (00130801–00130806) tum 1.43. COLORATION (pl. 3): As in (WAMP). male. GENITALIA: Posterior wall laterally OTHER SPECIMENS EXAMINED: AUSTRA- with distinct crescent-shaped interramal LIA: Western Australia: 11 km N of Cool- sclerites; posteriorly without a sclerotized gardie-Esperance Hiway on Kambalda Road, transverse band; posterolaterally with a 31.25231uS 121.5899uE, 320 m, 18 Nov 1999, distinct swelling covered with microtrichia; R.T. Schuh, G. Cassis, & R. Silveira, Mela- longitudinal fold on either side of midline leuca sheathiana W. Fitzg. (Myrtaceae), along part of length; posterior margin of det. Perth 05671396, 1- (00371314) (AM). posterior wall without spicules and not 33.3 km S of Norseman, 32.46461uS 2010 SCHUH AND WEIRAUCH: MYRTACEAE-FEEDING PHYLINAE 59

Fig. 28. Melaleucoides similis, male genitalia (AMNH_PBI 00087307).

121.6778uE, 300 m, 19 Nov 1999, R.T. Schuh, ages, the elongate slender form of the body in G. Cassis, & R. Silveira, 2- (00371315, most specimens, and the form of the male 00089117), 1- (00371966) (AM). 91.4 km genitalia with the elongate, fingerlike apical SE of Southern Cross, 31.97145uS 119.287uE, projection on the apex of the left paramere. 375 m, 04 Dec 1997, Schuh, Cassis, Brai- Most similar in size and coloration to M. lovsky, Asquith, Melaleuca teuthidoides Bar- verticordiae, but easily distinguished by the low (Myrtaceae), det. Perth 05054834, 1- comblike fringe on the apical spine of the (00087308) (AM), 7U (00373099–00373105) endosoma in that species. (AMNH). DESCRIPTION: Male: Body greatly elon- gate, parallel sided; mean total length 3.38, Melaleucoides similis, new species mean width pronotum 1.11. COLORATION Figure 28, map 5, plate 3 (pl. 3): Head: Uniformly yellow; scapus and pedicellus yellow to dirty yellow; labium pale DIAGNOSIS: Recognized by the yellow- with segment 4 heavily infuscate. Thorax: orange coloration of the body and append- Pronotum unicolorous yellow; scutellum 60 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 344 unicolorous yellow; hemelytron, including mondii Benth., and Thryptomene aspera cuneus, unicolorous yellow; markings on glabra E. Pritz. (Myrtaceae). The very limited cuneus absent; membrane weakly fumose, numbers of available specimens preclude veins yellow; hind femur unicolorous pale, conclusions regarding host specificity in this without black spots; hind tibial spines black, taxon. without dark spots at bases. Abdomen: DISTRIBUTION (map 5): Known form the Venter unicolorous pale or mostly so. SUR- Carnamah Shire Coast near Eneabba. FACE AND VESTITURE: Dorsum with HOLOTYPE: AUSTRALIA: Western Aus- suberect or reclining black setae, without tralia: 15 km E of Merredin, 31.37749uS sericeous or woolly setae. STRUCTURE: 118.6933uE, 330 m, 16 Nov 1999, R.T. Schuh Head: Somewhat projecting; interocular and G. Cassis, Melaleuca conothamnoides space relatively large; eyes leaving gena C.A. Gardner (Myrtaceae), det. Perth 0567- moderately exposed in lateral view. Antenna: 0624, 1- (AMNH_PBI 00370994) (WAMP). Segment 2 of equal diameter over entire PARATYPES: AUSTRALIA: Western Aus- length; antennal fossa with ventral margin at tralia: 11 km S of Eneabba, Eneabba ventral margin of eye. Labium: Reaching to National Park [96-50], 29.9025uS 115.24321uE, about anterior margin of pygophore. Thorax: 150 m, 01 Nov 1996, Schuh and Cassis, Scholt- Pretarsus: Pulvilli absent. GENITALIA zia drummondii Benth. (Myrtaceae), det. (fig. 28): Endosoma: Primary strap apically Perth 05120209, 1- (00087307) (AMNH). greatly elongate, angled dorsally relative to 15 km E of Merredin, 31.37749uS 118.6933uE, body of endosoma, denticulate on apical half 330 m, 16 Nov 1999, R.T. Schuh and G. of ventral margin; secondary endosomal Cassis, Melaleuca conothamnoides C.A.Gard- strap fused with primary strap primal to ner (Myrtaceae), det. Perth 05670624, 1U secondary gonopore, fused with primary (00370993) (AM). 24 km W of Sandstone, strap distad of gonopore; spinelike, elongate 28.01426uS 119.0474uE, 650 m, 26 Oct 1996, process arising near gonopore on dorsal Schuh and Cassis, Thryptomene aspera glabra surface present, erect, unornamented; spine- E. Pritz. (Myrtaceae), det. Perth 05095182, like, elongate process near gonopore on 1- (00135637) (AM), Thryptomene aspera ventral surface present, sharply recurved glabra E. Pritz. (Myrtaceae), det. Perth toward base of endosoma; bladderlike pro- 05095182, 1- (00135638) (AMNH). cess distad of secondary gonopore present, balloonlike; secondary gonopore seen fron- Melaleucoides systenae, new species tally (facing up) in lateral view of endosoma. Figure 29, map 5, plate 4 Phallotheca: Nearly erect; apex simple. Left Paramere: Shaft at right angles to body; body DIAGNOSIS: Males recognized in most more or less quadrate; apex medially broadly specimens by the castaneous pronotum ante- rounded; apex posterolaterally with a greatly rior lobe, scutellum, and posterior half of the elongate projection (fig. 28); anterior process endocorium; phallotheca with a very broad triangular in lateral view; posterior process in reflexed flange on the dorsal margin; left the form of a short, straight, fingerlike paramere very broad apically; endosoma projection; base of posterior process without with a ventral process associated with the conspicuous shoulder. Right Paramere: Body secondary gonopore and no denticles distad short and broad, lanceolate; apex short, of the gonopore. Except for rather variable clawlike, decurved. coloration with the taxon, M. systenae is not Female: Distinctly more ovoid than male; easily confused with any other known species mean total length 3.11, mean width prono- of Melaleucoides. tum 1.08. COLORATION (pl. 3): As in DESCRIPTION: Male: Body weakly to male. GENITALIA: Not examined. distinctly ovoid; mean total length 3.23, mean ETYMOLOGY: Named for the remarkable width pronotum 1.22. COLORATION similarity of appearance with Melaleucoides (pl. 4): Head: Mostly dark with pale gula verticordiae, new species. and posterior head margin; scapus unicolor- HOSTS: Recorded from Melaleuca con- ous dark; pedicellus black at extreme base, othamnoides C.A. Gardner, Scholtzia drum- remainder pale; labium generally infuscate, 2010 SCHUH AND WEIRAUCH: MYRTACEAE-FEEDING PHYLINAE 61

Fig. 29. Melaleucoides systenae, male genitalia (AMNH_PBI 00368601). heavily so apically. Thorax: Pronotum usu- mal strap fused with primary strap proximal ally with dark anterior lobe and contrasting to secondary gonopore, reaching only to level lighter-colored posterior lobe; scutellum en- of secondary gonopore; spinelike elongate tirely dark; clavus pale, corium pale on process arising near gonopore on dorsal anterior half, dark posteriorly; markings on surface absent; spinelike, elongate process cuneus present as a contrasting white basal near gonopore on ventral surface present, fascia; membrane fumose, veins white or relatively short, recurved toward base of pale; hind femur nearly unicolorous dark; endosoma; bladderlike process distad of hind tibial spines dark with very small dark secondary gonopore present, elongate, and bases. Abdomen: Venter unicolorous dark. paralleling primary endosomal strap; second- SURFACE AND VESTITURE: Dorsum ary gonopore seen frontally (facing up) in with suberect or reclining black setae, with- lateral view of endosoma. Phallotheca: Near- out sericeous or woolly setae. STRUC- ly erect; dorsal surface with a curving, TURE: Head: Weakly projecting; interocular posteriorly directed, platelike projection; space relatively large; eyes leaving gena ventral surface with a projecting keel; apex broadly exposed in lateral view. Antenna: simple. Left Paramere: Shaft at right angles Segment 2 weakly tapering, more slender at to body; body spoon shaped; apex quadrate; base; antennal fossa with ventral margin at anterior process triangular in lateral view; ventral margin of eye. Labium: Just reaching posterior process in the form of a short, onto abdomen. Thorax: Pretarsus: Pulvilli straight, fingerlike projection; base of poste- present, flaplike, covering about one third of rior process without conspicuous shoulder. ventral claw surface. GENITALIA (fig. 29): Right Paramere: Body short and broad, Endosoma: Primary strap apically elongate, lanceolate; apex with short fingerlike process. angled dorsal relative to body of endosoma, Female: Slightly more ovoid than male; without ornamentation; secondary endoso- mean total length 3.51, mean width prono- 62 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 344 tum 1.32. COLORATION (pl. 4): Paler than 05879264, 2- (00130029, 00130030), 3U male, more strongly reddish, without the (00130031–00130033) (AMNH). Cervantes, nearly castaneous quality of most males. 30.49902uS 115.0684uE, 3 m, 10 Dec 1997, GENITALIA: Posterior wall laterally with Schuh, Brailovsky, Melaleuca viminea Lindl. distinct crescent-shaped interramal sclerites; (Myrtaceae), det. Perth 05879205, 1- (001- posteriorly without a sclerotized transverse 30601), 4U (00130602–00130605) (AMNH), band; posterolaterally with a distinct swelling 3U (00130606–00130608) (WAMP). Frank covered with microtrichia; longitudinal fold Hann National Park, 37 km E of Lake King, on either side of midline along part of length; 33.07753uS 120.0918uE, 400 m, 05 Nov 1996, posterior margin of posterior wall without Schuh and Cassis, Melaleuca sp. (Myrta- spicules and not reflexed dorsally; interramal ceae), det. Perth 05236908, 1- (00087405), lobes present and asymmetrical, reclining, 1U (00087406) (AM). Kalbarri National overlapping, and heavily ornamented with Park, 22.9 km E Kalbarri, 27.75408uS spicules; vestibulum with medial plates scler- 114.3711uE, 500 m, 29 Oct 1996, Schuh and otized and readily observed, small, nearly Cassis, 6U (00135367–00135372) (WAMP). symmetrical, triangular, with sclerotized Kalbarri National Park, 37.7 km E Kalbarri, guide present as short heavily sclerotized 27.8482uS 114.4746uE, 500 m, 29 Oct 1996, tube on left side. Schuh and Cassis, Melaleuca laetifica Craven ETYMOLOGY: Named for the host species, ms (Myrtaceae), det. Perth 05054540, 2- Melaleuca systena (Myrtaceae). (00135064, 00135065), 5U (00368631–0036- HOSTS: Melaleuca laetifica Craven [ms. 8632, 00368635–00368636, 00368638) (AM), name] (pl. 5B), Melaleuca systena Craven, 6- (00135059–00135063, 00135066), 11U Melaleuca urceolaris Benth., Melaleuca vimi- (00368629–00368630, 00368633–00368634, 00- nea Lindl., and Phymatocarpus porphyroce- 368637, 00368639–00368644) (AMNH), 2- phalus F. Muell. (Myrtaceae). (00135067, 00135068), 4U (00368625–0036- DISTRIBUTION (map 5): Relatively wide- 8628) (WAMP). spread in Western Australia, with localities OTHER SPECIMENS EXAMINED: AUSTRA- ranging from the Nullabor Plain in the LIA: Western Australia: 1 km S of Lillian southeast to Kalbarri National Park in the Stoke Rock, 33.07681uS 120.0982uE, 380 m, north. 21 Nov 1999, R.T. Schuh and G. Cassis, HOLOTYPE: AUSTRALIA: Western Aus- Verticordia chrysantha Endl. (Myrtaceae), tralia: 8.2 km E of Indian Ocean Rd on det. Perth 05672023, 1U (00371784) (AM). Coorow-Greenhead Rd, Lesuer National 4.5 km S of Jurien on Indian Ocean Rd, Park, 30.04767uS 115.0551uE, 30 m, 06 Nov 30.33667uS 115.069uE, 18 m, 06 Nov 2004, 2004, Cassis, Weirauch, Tatarnic, Symonds, Cassis, Weirauch, Tatarnic, Symonds, Mela- Melaleuca systena Craven (Myrtaceae), det. leuca systena Craven (Myrtaceae), det. Perth 6990401, 1- (AMNH_PBI 00368603) Perth6987567, 1U (00371783) (AM). 8.2 km (WAMP). E of Indian Ocean Rd on Coorow-Green- PARATYPES: AUSTRALIA: Western Aus- head Rd, Lesuer National Park, 30.04767uS tralia: 8.2 km E of Indian Ocean Rd on 115.0551uE, 30 m, 06 Nov 2004, Cassis, Coorow-Greenhead Rd, Lesuer National Weirauch, Tatarnic, Symonds, Melaleuca sys- Park, 30.04767uS 115.0551uE, 30 m, 06 Nov tena Craven (Myrtaceae), det. Perth 6990401, 2004, Cassis, Weirauch, Tatarnic, Symonds, 4 nymphs (00368619, 00368621, 00368623– Melaleuca systena Craven (Myrtaceae), det. 00368624) (AM). 11 km S of Eneabba on Perth6990401, 2- (00368602, 00368604), 3U Brand Hiway, Eneabba Reserve, 29.91094uS (00368606–00368608) (AM), 2- (00368600, 115.1175uE, 100 m, 21 Oct 2004, Cassis, Wall, 00368601), 3U (00368609–00368610, 0036- Weirauch, Symonds, Melaleuca urceolaris 8613) (AMNH) 2U (00368617, 00368618) Benth. (Myrtaceae), det. Perth 6986919, 5U (UNSW), 3U (00368614–00368616) (WAMP). (00371778–00371782) (AM). Cervantes, Brand Hiway 45.9 km S of Dongarra Road, 30.49902uS 115.0684uE, 3 m, 10 Dec 1997, 29.57703uS 115.1348uE, 100 m, 31 Oct 1996, Schuh, Brailovsky, Melaleuca viminea Lindl. Schuh and Cassis, Phymatocarpus porphyro- (Myrtaceae), det. Perth 05879205, 1 nymph cephalus F. Muell. (Myrtaceae), det. Perth (00372247) (AM). 2010 SCHUH AND WEIRAUCH: MYRTACEAE-FEEDING PHYLINAE 63

Fig. 30. Melaleucoides uncinatae, male. Scanning electron micrographs. A. Head and thorax, lateral view. B. Setae on hemelytra. C. Pretarsus, ventral view. D. Abdomen, lateral view. E. Pygophore, lateral view. F. Pygophore, dorsal view. lp 5 left paramere; pe 5 parempodium; pv 5 pulvillus; rp 5 right paramere (AMNH_PBI 00368327).

Melaleucoides uncinatae, new species coloration (pls. 3, 4) of the dorsum, with Figures 30, 31; map 6; plate 4 the apex of the scutellum and the base of the cuneus pale. Distinguished from that DIAGNOSIS: Most similar to M. species by the different form of the phal- rhaphiophyllae in the heavily castaneous lotheca and the left paramere (figs. 26, 31), 64 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 344

Fig. 31. Melaleucoides uncinatae, male genitalia (AMNH_PBI 00368328). the endosoma of M. rhaphiophyllae being space moderate; eyes leaving gena moderate- unknown. ly exposed in lateral view. Antenna: Segment DESCRIPTION: Male: Body moderately 2 weakly tapering, more slender at base; elongate, parallel sided; mean total length antennal fossa with ventral margin at ventral 2.99, mean width pronotum 1.25. COLOR- margin of eye. Labium: Just reaching onto ATION (pl. 4): Head: Uniformly dark; sca- abdomen. Thorax: Pretarsus (fig. 30C): Pul- pus brown; pedicellus unicolorous black or villi present, flaplike, covering about one castaneous; labium generally infuscate, heavi- third of ventral claw surface. GENITALIA ly so apically. Thorax: Pronotum unicolorous (figs. 30D–F, 31): Endosoma: Primary strap deep red to black; scutellum red or dark apically weakly elongate, weakly curving, brown with white or pale tip; corium and without ornamentation; secondary endoso- clavus largely castaneous, markings on cune- mal strap fused with primary strap proximal us present as a contrasting white basal fascia, to level of secondary gonopore, and not remainder castaneous; membrane strongly projecting beyond gonopore; spinelike, elon- fumose, veins white; hind femur nearly gate process arising near gonopore on dorsal unicolorous dark; hind tibial spines with surface present, erect, ornamented with conspicuous dark spots at bases. Abdomen: denticles apically; spinelike, elongate process Venter heavily red laterally. SURFACE near gonopore on ventral surface absent; AND VESTITURE: Dorsum with golden bladderlike process distad of secondary reclining setae; dorsum without sericeous gonopore present; secondary gonopore seen or woolly setae. STRUCTURE: Head laterally in lateral view of endosoma. Phal- (fig. 30A): Barely projecting; interocular lotheca: More or less right angulate; dorsal 2010 SCHUH AND WEIRAUCH: MYRTACEAE-FEEDING PHYLINAE 65 surface with a long fingerlike projection; apex 8.2 km N of Eneabba, 29.7462uS 115.254uE, simple. Left Paramere: Shaft in same axis as 100 m, 31 Oct 1996, Schuh and Cassis, body; body spoon shaped; apex medially Melaleuca rhaphiophylla Schauer (Myrta- broadly rounded; anterior process triangular ceae), det. Perth 05120195, 1U (00008726) in lateral view; posterior process in the form (AM). of a short, curved, fingerlike projection; base OTHER SPECIMENS EXAMINED: AUSTRA- of posterior process with distinct shoulder. LIA: Western Australia: 56.6 km W of Right Paramere: Body short and broad, Yalgoo, 28.42397uS 116.1233uE, 600 m, 27 lanceolate; apex short, clawlike, decurved. Oct 1996, Schuh and Cassis, Melaleuca unci- Female: Distinctly more ovoid than male; nata R. Br. (Myrtaceae), det. Perth 05120640, mean total length 3.26, mean width prono- 8U (00368345–00368352) (AMNH). tum 1.34. COLORATION (pl. 4): Paler than male. GENITALIA: Posterior wall laterally with distinct crescent-shaped interramal Melaleucoides undulatae, new species sclerites; posteriorly without a sclerotized Figure 32, map 6, plate 4 transverse band; posterolaterally with a distinct swelling covered with microtrichia; DIAGNOSIS: Recognized by the castaneous longitudinal fold on either side of midline calli, the elongate red markings on the along part of length; posterior margin of hemelytron (pl. 4), and the endosoma lacking posterior wall without spicules and not denticles distad of the secondary gonopore, a reflexed dorsally; interramal lobes roughly dorsal process, and a bladderlike process just symmetrical, erect, and heavily ornamented distad of the gonopore (fig. 32). Most easily with spicules; vestibulum with medial plates confused with M. annae on the basis of sclerotized and readily observed, small, near- coloration, but the red markings on the ly symmetrical, triangular, with sclerotized hemelytron less strongly linear in that species guide present as short heavily sclerotized tube (pl. 1) and the endosoma with denticles on on left side. the secondary strap distad of the secondary ETYMOLOGY: Named for the host species, gonopore, a recurved ventral process associ- Melaleuca uncinata (Myrtaceae). ated with the secondary gonopore, and no HOSTS: Nearly all available specimens are bladderlike process (fig. 13). recorded from Melaleuca uncinata R. Br. DESCRIPTION: Male: Body weakly elon- (pl. 6D); a single specimen is recorded from gate, weakly ovoid; mean total length 3.03, Melaleuca rhaphiophylla Schauer (Myrta- mean width pronotum 1.17. COLORATION ceae). (pl. 4): Head: Pale, clypeus dark, frons with DISTRIBUTION (map 6): Known from large dark bilateral markings; scapus uni- Yalgoo and Eneabba in Western Australia. colorous pale; pedicellus unicolorous pale; HOLOTYPE: AUSTRALIA: Western Aus- labium pale with segment 4 heavily infuscate. tralia: 56.6 km W of Yalgoo, 28.42397uS Thorax: Pronotum pale with castaneous calli 116.1233uE, 600 m, 27 Oct 1996, Schuh and and humeri; scutellum pale; hemelytron pale Cassis, Melaleuca uncinata R.Br. (Myrta- with longitudinal castaneous stripes along ceae), det. Perth 05120640, 1- (AMNH_PBI veins; markings on cuneus present as a red or 00368315) (WAMP). castaneous apex, otherwise pale; membrane PARATYPES: AUSTRALIA: Western Aus- weakly fumose, veins white; hind femur with tralia: 56.6 km W of Yalgoo, 28.42397uS many brown or black spots; hind tibial spines 116.1233uE, 600 m, 27 Oct 1996, Schuh and with conspicuous dark spots at bases. Abdo- Cassis, Melaleuca uncinata R. Br. (Myrta- men: Venter unicolorous pale or mostly so. ceae), det. Perth 05120640, 3- (00136545– SURFACE AND VESTITURE: Dorsum 00136546, 00368325), 12U (00136547–001- with golden reclining setae; dorsum without 36558) (AM), 14- (00368313–00368314, 003- sericeous or woolly setae. STRUCTURE: 68316–00368324, 00368327–00368328, 0036- Head: Barely projecting; interocular space 8364), 16U (00368329–00368344) (AMNH), moderate; eyes leaving gena moderately 15- (00368326, 00368353–00368363, 00368- exposed in lateral view. Antenna: Segment 2 365–00368367) (WAMP). Brand Hiway weakly tapering, more slender at base; 66 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 344

Fig. 32. Melaleucoides undulatae, male genitalia (AMNH_PBI 00087483). antennal fossa with ventral margin at ventral apex medially angulate, bluntly acuminate; margin of eye. Labium: Just reaching onto anterior process slender and cylindrical in abdomen. Thorax: Pretarsus: Pulvilli present, lateral view; posterior process in the form of flaplike, covering about one third of ventral a short, straight, fingerlike projection; base of claw surface. GENITALIA (fig. 32): Endo- posterior process with distinct shoulder. soma: Primary strap apically weakly elon- Right Paramere: Body short and broad, gate, weakly arcuate and curving dorsally, lanceolate; apex short, clawlike, decurved. without ornamentation; secondary endoso- Female: Slightly more ovoid than male; mal strap fused with primary strap proximal mean total length 3.27, mean width prono- to secondary gonopore, reaching only to level tum 1.25. COLORATION (pl. 4): As in of secondary gonopore; elongate process male. GENITALIA: Not examined. arising near gonopore on dorsal surface ETYMOLOGY: Named for the host species, present, erect, curving apically, unornament- Melaleuca undulata (Myrtaceae). ed; spinelike, elongate process near gonopore HOSTS: Melaleuca undulata Benth. (Myr- on ventral surface absent; bladderlike process taceae) (pl. 6E). distad of secondary gonopore present; sec- DISTRIBUTION (map 6): Recorded only ondary gonopore seen laterally in lateral view from Kalbarri National Park in Western of endosoma. Phallotheca: Nearly erect; Australia. dorsal surface with a fingerlike projection HOLOTYPE: AUSTRALIA: Western Aus- and a curving, posteriorly directed, platelike tralia: Kalbarri National Park, 22.9 km E projection; apex simple. Left Paramere: Shaft Kalbarri, 27.75408uS 114.3711uE, 500 m, 29 in same axis as body; body spoon shaped; Oct 1996, Schuh and Cassis, Melaleuca undu- 2010 SCHUH AND WEIRAUCH: MYRTACEAE-FEEDING PHYLINAE 67 lata Benth. (Myrtaceae), det. Perth 05120403, and some sericeous or woolly setae. STRUC- 1- (AMNH_PBI 00372306) (WAMP). TURE: Head (fig. 33A): Moderately project- PARATYPES: AUSTRALIA: Western Aus- ing; interocular space large; eyes leaving gena tralia: Kalbarri National Park, 22.9 km E moderately exposed in lateral view. Antenna: Kalbarri, 27.75408uS 114.3711uE, 500 m, 29 Segment 2 of equal diameter over entire Oct 1996, Schuh and Cassis, Melaleuca undu- length; antennal fossa with ventral margin at lata Benth. (Myrtaceae), det. Perth 05120403, ventral margin of eye. Labium: Reaching to 9- (00372300–00372305, 00372307, 00372310– about anterior margin of pygophore. Thorax: 00372311), 6U (00372319–00372320, 00372323, Pretarsus (fig. 33D, E): Pulvilli absent. GEN- 00372326–00372328) (AM), 6- (00087483, 00- ITALIA (figs. 33F, G, 34): Endosoma: Pri- 372299, 00372308–00372309, 00372312–0037- mary strap apically greatly elongate and 2313), 5U (00372321–00372322, 00372324– near straight, with a comblike ventral mar- 00372325, 00372331) (AMNH), 6- (00087- gin; secondary endosomal strap fused with 275, 00372314–00372318), 5U (00087276, 0037- primary strap proximal to secondary gono- 2329–00372330, 00372332–00372333) (WAMP). pore, fused with primary strap distad of OTHER SPECIMENS EXAMINED: AUSTRA- secondary gonopore; spinelike, elongate LIA: Western Australia: Kalbarri National process arising near gonopore on dorsal Park, 22.9 km E Kalbarri, 27.75408uS surface present, erect, slender, relatively 114.3711uE, 500 m, 29 Oct 1996, Schuh and short, unornamented; spinelike, elongate Cassis, Melaleuca undulata Benth. (Myrta- process near gonopore on ventral surface ceae), det. Perth 05120403, 6 nymphs present, recurved toward base of endosoma; (00372334–00372339) (AM). bladderlike process distad of secondary gonopore absent; secondary gonopore seen laterally in lateral view of endosoma. Phal- Melaleucoides verticordiae, new species lotheca: Smoothly curving on dorsal margin; Figures 33, 34; map 6; plate 4 ventral surface with a projecting, fingerlike keel; apex simple. Left Paramere: Shaft at DIAGNOSIS: Recognized by the yellow- right angles to body; body spoon shaped; orange coloration of the body and append- apex medially broadly rounded; anterior ages, the elongate slender form of the body in process triangular in lateral view; posterior most specimens, and the form of the male process in the form of a short, straight, genitalia with the comblike fringe on the fingerlike projection; base of posterior pro- apical spine of the endosoma. Most similar in cess with distinct shoulder. Right Paramere: size and coloration to M. similis, but easily Body short and broad, lanceolate; apex distinguished by the elongate, fingerlike short, clawlike, decurved. apical projection on the apex of the left Female: Distinctly more ovoid than male; paramere in that species. mean total length 3.59, mean width prono- DESCRIPTION: Male: Body greatly elon- tum 1.24. COLORATION (pl. 4): As in gate, parallel sided; mean total length 3.49, male. GENITALIA (fig. 34): Posterior wall mean width pronotum 1.19. COLORATION laterally with distinct crescent-shaped inter- (pl. 4): Head: Uniformly yellow; scapus and ramal sclerites; posteriorly without a sclero- pedicellus yellow to dirty yellow; labium pale tized transverse band; posterolaterally with a with segment 4 heavily infuscate. Thorax: distinct swelling covered with microtrichia; Pronotum unicolorous yellow; scutellum longitudinal fold on either side of midline unicolorous yellow; hemelytron, including along part of length; posterior margin of cuneus, unicolorous yellow; markings on posterior wall without spicules and not cuneus absent; membrane weakly fumose, reflexed dorsally; interramal lobes roughly veins yellow; hind femur unicolorous pale, symmetrical, erect, and heavily ornamented without black spots; hind tibial spines black, with spicules; vestibulum with medial plates without dark spots at bases. Abdomen: sclerotized and readily observed; small, near- Venter unicolorous pale or mostly so. SUR- ly symmetrical, triangular; with sclerotized FACE AND VESTITURE (fig. 33C): Dor- guide structure present as caplike structure sum with suberect or reclining black setae with internal differentiation. 68 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 344

Fig. 33. Melaleucoides verticordiae, male. Scanning electron micrographs. A. Head and thorax, lateral view. B. Abdomen, lateral view. C. Setae on hemelytra. D. Pretarsus, frontal view. E. Pretarsus, frontal view, detail of rudimentary dorsal arolium. F. Pygophore, lateral view. G. Pygophore, dorsal view. da 5 dorsal arolium; lp 5 left paramere; pe 5 parempodium; ph 5 phallotheca; rp 5 right paramere (AMNH_PBI 00129613). ETYMOLOGY: Named for the genus Verti- species: Verticordia chrysantha Endl. (pl. 8C, cordia (Myrtaceae), from which many of the D), Lindl., Verticordia available specimens were taken. polytricha Benth. (pl. 9B), and Pileanthus HOSTS: The majority of available speci- peduncularis borealis Endl. [ms. name] mens were taken from the following plant (pl. 7B) (Myrtaceae). Some additional taxa 2010 SCHUH AND WEIRAUCH: MYRTACEAE-FEEDING PHYLINAE 69

Fig. 34. Melaleucoides verticordiae, male and female genitalia (AMNH_PBI 00135071, 00135416). 70 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 344 of Myrtaceae are also recorded as hosts for of Coolgardie on Great Eastern Hiway, much smaller numbers of specimens. The 31.23414uS 120.1562uE, 17 Nov 1999, R.T. record of 21 specimens from Allocasuarina Schuh, G. Cassis, & R. Silveira, Verticordia (Casuarinaceae) does not represent a breed- chrysantha Endl. (Myrtaceae), det. Perth ing host in our view, in the face of the 05672015, 1- (00372421) (AM). Eneabba hundreds of specimens known only from the on Brand Hiway, 29.80735uS 115.2699uE, Myrtaceae; this is probably the result of 100 m, 31 Oct 1996, Schuh and Cassis, mislabeling or commingling of specimens in Verticordia densiflora Lindl. (Myrtaceae), the field. det. Perth 05120179, 48U (00372607– DISTRIBUTION (map 6): Relatively wide- 00372629, 00372645–00372669) Verticordia spread in Western Australia, ranging from chrysantha Endl. (Myrtaceae), det. Perth south of the Goldfields in the east to Kalbarri 05120160, 9- (00372519–00372527), 23U (003- National Park in the north. 72528–00372541, 00372551–00372559) (AM), HOLOTYPE: AUSTRALIA: Western Aus- 26- (00135069, 00135071–00135075, 00137- tralia: NW Coastal Hiway 57 km N of Kal- 425–00137444), 33U (00135078–00135082, 00- barri Road, 27.44756uS 114.6867uE, 500 m, 135085–00135088, 00137461–00137484) (AMNH), 30 Oct 1996, Schuh and Cassis, Pileanthus Verticordia densiflora Lindl. (Myrtaceae), det. peduncularis borealis Endl. (Myrtaceae), det. Perth 05120179, 16U (00372630–00372637, Perth 05120349, 1- (AMNH_PBI 00135409) 00372670–00372677) (UNSW), 7U (00372- (WAMP). 638–00372644) Verticordia chrysantha Endl. PARATYPES: AUSTRALIA: Western Aus- (Myrtaceae), det. Perth 05120160, 17- (001- tralia: 1 km S of Lillian Stoke Rock, 37445–00137460, 00372379), 41U (001374- 33.07681uS 120.0982uE, 380 m, 21 Nov 85–00137502, 00137504–00137526) (WAMP). 1999, R.T. Schuh and G. Cassis, Verticordia Kalbarri National Park, Loop Road, chrysantha Endl. (Myrtaceae), det. Perth 27.56163uS 114.4376uE, 300 m, 28 Oct 05672023, 1- (00368814) (AMNH). 11 km 1996, Schuh and Cassis, 29U (00372388– S of Eneabba, Eneabba National Park [96- 00372416) Verticordia polytricha Benth. 50], 29.9025uS 115.24321uE, 150 m, 01 Nov (Myrtaceae), det. Perth 05120594, 15- (003- 1996, Schuh and Cassis, Allocasuarina cam- 72450–00372464), 47U (00372465–00372511) pestris E. Pritz. (Casuarinaceae), det. Perth (AM), Melaleuca megacephala F. Muell. 05095182, 15- (00372733–00372747) Caly- (Myrtaceae), det. Perth 05120616, 6- (0013- trix glutinosa (Myrtaceae), 10- (00372349– 6709–00136713, 00136715), 6U (00136716– 00372358), 11U (00372359–00372369) Verti- 00136721) Pileanthus peduncularis borealis cordia chrysanthella E. Pritz. (Myrtaceae), Endl. (Myrtaceae), det. Perth 05120586, det. Perth 05095182, 3- (00372678–0037- 10- (00135472–00135480, 00135926), 12U 2680) (AM), Verticordia chrysantha A.S. (00135481–00135492) (AMNH), 13- (0037- George (Myrtaceae), det. Field ID; Host 96- 2374–00372378, 00372380–00372387) Mela- 147, 14- (00135225–00135238), 11U (0013- leuca megacephala F. Muell. (Myrtaceae), 5242–00135252) (AMNH), Verticordia chry- det. Perth 05120616, 7U (00136722–00136- santha A.S. George (Myrtaceae), det. Field 728) Pileanthus peduncularis borealis Endl. ID; Host 96-147, 3- (00135239–00135241), (Myrtaceae), det. Perth 05120586, 8- 8U (00135253–00135260) (WAMP). 20.6 km S (00135920–00135925, 00135927–00135928), of Norseman-Lake King Road on Lake King- 10U (00135929–00135938) Verticordia poly- Cascades Road, 33.16284uS 120.2813uE, 400 m, tricha Benth. (Myrtaceae), det. Perth 22 Nov 1999, R.T. Schuh, G. Cassis, & R. 05120594, 7U (00372512–00372518) (WAMP). Silveira, Verticordia chrysantha Endl. (Myrta- Kalbarri National Park, Z-Bend Road, ceae), det. Perth 05120160, 1- (00372417), 27.61971uS 114.3864uE,500m,28Oct1996, 3U (00372418–00372420) (AM), 10- (0012- Schuh and Cassis, Pileanthus peduncularis 9607–00129612, 00129614–00129617), 10U borealis Keighery ms. (Myrtaceae), det. Perth (00129621, 00129623–00129629, 00129632, 05120586, 1- (00372570), 2U (00372571, 00129635) (AMNH), 3- (00129618–0012- 00372572) (AM). NW Coastal Hiway 57 km 9620), 5U (00129630–00129631, 00129633– N of Kalbarri Road, 27.44756uS 114.6867uE, 00129634, 00129636) (WAMP). 123 km W 500 m, 30 Oct 1996, Schuh and Cassis, 2010 SCHUH AND WEIRAUCH: MYRTACEAE-FEEDING PHYLINAE 71

Pileanthus peduncularis borealis Endl. (Myr- 00372606) (AM), 3- (00135633–00135635), taceae), det. Perth 05120349, 3- (00372281– 16U (00135639–00135654) (AMNH). 54.3 km 00372283), 10U (00372284–00372293) (AM), N of jct of Agana Kalbarri Rd and Brand 14- (00135408, 00136748–00136760), 14U Hiway (rest area), 27.47362uS 114.7054uE, (00135410–00135416, 00136761–00136766, 240 m, 24 Oct 2004, Cassis, Wall, Weirauch, 00136774) (AMNH), 11U (00136767–0013- Symonds, Pileanthus vernicosus F. Muell. 6773, 00136775–00136778) (WAMP). ca. 1 km (Myrtaceae), det. Field ID, 9- (00368597– S of Murchison House HS, Kalbarri National 00368599, 00368870–00368875), 2U (0036- Park, 27.65822uS 114.2394uE, 60 m, 23 Oct 8876, 00368877), 20 nymphs (00368878– 2004, Cassis, Wall, Weirauch, Symonds, 00368897) Verticordia capillaris A.S. George Pileanthus vernicosus F. Muell. (Myrtaceae), (Myrtaceae), det. Perth 6989861, 6- (0036- det. Perth6988415, 4- (00368645–00368648), U U 8673–00368678), 11 (00368679–00368689), 12 (00368649–00368660) (AMNH). 2 nymphs (00368690, 00368691) (AMNH). OTHER SPECIMENS EXAMINED: AUSTRA- 123 km W of Coolgardie on Great Eastern LIA: Western Australia: 1 km S of Lillian Hiway, 31.23414uS 120.1562uE, 17 Nov 1999, u u Stoke Rock, 33.07681 S 120.0982 E, 380 m, R.T. Schuh, G. Cassis, & R. Silveira, Verti- 21 Nov 1999, R.T. Schuh and G. Cassis, cordia chrysantha Endl. (Myrtaceae), det. Verticordia chrysantha Endl. (Myrtaceae), Perth 05672015, 7- (00372422–00372428), det. Perth 05672023, 4- (00372542–003- U U 21 (00372429–00372449) (AM). Brand Hi- 72545), 4 (00372547–00372550), 1 nymph way 55.9 km S of Dongarra Road, 29.62934uS (00372546) (AM), 35- (00368815–003688- U 115.2187uE, 100 m, 31 Oct 1996, Schuh and 35, 00368840–00368853), 46 (00368776– Cassis, Scholtzia drummondii Benth. (Myrta- 00368797, 00368802–00368813, 00368854– ceae), det. Perth 05120209, 2- (00088976, 00368865), 4 nymphs (00368866–00368869) 00088979) (AM), 3- (00135450, 00135453– (AMNH), 4- (00368836–00368839), 4U 00135454) (AMNH). Eneabba on Brand (00368798–00368801) (UCR). 3.5 km E of Hiway, 29.80735uS 115.2699uE, 100 m, 31 Lillian Stoke Rock, 33.07679uS 120.132uE, Oct 1996, Schuh and Cassis, Verticordia 360 m, 21 Nov 1999, R.T. Schuh, G. Cassis, chrysantha Endl. (Myrtaceae), det. Perth & R. Silveira, Verticordia roei (Myrtaceae), 2- (00372370, 00372371), 2 nymphs (00372372, 05120160, 10 nymphs (00372560–00372569) 00372373) (AM), 22- (00368754–00368775), (AM), 1 nymph (00135077) (AMNH). Kal- 38U (00368716–00368753) (AMNH). 11 km S barri National Park, Z-Bend Road, u u of Eneabba, Eneabba National Park [96-50], 27.61971 S 114.3864 E, 500 m, 28 Oct 1996, 29.9025uS 115.24321uE, 150 m, 01 Nov 1996, Schuh and Cassis, Pileanthus peduncularis Schuh and Cassis, Verticordia chrysanthella borealis Keighery ms. (Myrtaceae), det. Perth E. Pritz. (Myrtaceae), det. Perth 05095182, 05120586, 6 nymphs (00372573–00372578) 19- (00372681–00372699), 31U (00372700– (AM). ca. 1 km S of Murchison House HS, 00372730) (AM), Calytrix glutinosa Benth. Kalbarri National Park, 27.65822uS (Myrtaceae), det. Perth 05120209, 26- 114.2394uE, 60 m, 23 Oct 2004, Cassis, Wall, (00135979–00136004), 23U (00136005–0013- Weirauch, Symonds, Pileanthus vernicosus F. 6027), 1 nymph (00368898) (AMNH). 20.6 km Muell. (Myrtaceae), det. Perth 6988415, 12 S of Norseman-Lake King Road on Lake nymphs (00368661–00368672) (AMNH). ca. King-Cascades Road, 33.16284uS 120.2813uE, 11 km S of Eneabba, Eneabba National Park, 400 m, 22 Nov 1999, R.T. Schuh, G. Cassis, & 29.90252uS 115.2432uE, 150 m, 01 Nov 1996, R. Silveira, Verticordia chrysantha Endl. Schuh and Cassis, Allocasuarina campestris (Myrtaceae), det. Perth 05672023, 1 nymph (Diels) L.A.S. Johnson (Casuarinaceae), det. (00129622) (AMNH); 22 Nov 1999, R.T. Perth 05120063, 15- (00372733–00372747), Schuh and G. Cassis, 1- (00129613) 6U (00372748–00372753) (AM). (AMNH). 24 km W of Sandstone, u u 28.01426 S 119.0474 E, 650 m, 26 Oct 1996, PHYLOGENETIC ANALYSIS Schuh and Cassis, Thryptomene aspera glabra E. Pritz. (Myrtaceae), det. Perth 05095182, We prepared the matrix shown in appen- 8- (00372579–00372586), 20U (00372587– dix 1 as a way of creating a classification 72 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 344 based on synapomorphy for the taxa treated Our concepts of female genitalic homology in this paper. It contains 25 in-group taxa were informed by the studies of Weirauch and 15 outgroups, mostly Australian, with a (2007), Schuh and Pedraza (2010), and species of Leucophoropterini used as the Weirauch and Schuh (2010). Genera included root. The selection of outgroups among in those studies and members of the Melaleu- Australian Phylini was informed by the coides group of genera possess morphological studies of Weirauch (2007) on the Polyozus features of the posterior wall that are not seen Eyles and Schuh group of genera, Soto and outside of the Australian fauna. Nonetheless, Weirauch (2009) on Jiwarli, Weirauch and the correlation between those features and the Schuh (2010) on the Xiphoidellus and related novel structures found in the males is genera, and Schuh and Pedraza (2010) on obviously in need of additional study. Wallabicoris. Outgroups include Capecapsus Our equal-weights analysis used ratchet tradouwensis Schuh from South Africa and and drift in addition to the default sectorial the Australian taxa Wallabicoris ozothamni search with fusing. Default parameters Schuh and Pedraza, W. pityrodii Schuh and for the driven search were used with the Pedraza, W. spyridii Schuh and Pedraza, minimum length tree set to 10. The equal- Xiphoides myersi (Woodward), Araucanophy- weights analysis produced a total of 10 most lus pacificus Carvalho, Xiphoidellus aureus parsimonious with a length (L) of 506, Weirauch and Schuh, X. furvus Weirauch consistency index (CI) of 35, and a retention and Schuh, X. pallidus Weirauch and Schuh, index (RI) of 62. The strict consensus of Jiwarli heliotropium Soto and Weirauch, those trees is shown in figure 35. Polyozus galbanus Eyles and Schuh, Exocar- Our implied weights analysis used the pocoris tantulus Weirauch, and the manu- same settings as above with a constant of script taxon Protemiris conospermi. concavity of K 5 3. Stronger constants Seventy-five characters are included, the produced essentially the same results. The majority related to morphology of the male single resultant tree, with a fit value of and female genitalia, but characters derived 36.982, is shown in figure 36. from somatic morphology and color are As a way of indicating clade support we also included. We have treated as additive chose the jackknife as a measure. These those multistate characters for which we values are plotted at the nodes in figures 35 could hypothesize some credible transforma- and 36 for percentages over 50. They were tion; all other characters are treated as obtained by running the jackknife resampling nonadditive. Additivities are indicated in option in TNT with resampling probability appendix 1 in the data matrix and the 5 36, 1000 replications and otherwise iden- character descriptions. We used TNT (Go- tical parameters as in the initial searches. loboff et al., 2008, 2009) for our analyses, Characters and character states were mapped applying both equal weights and implied on the resulting trees (figs. 35, 36) using weights (Goloboff, 1993) approaches to WinClada and show unambiguous changes. character optimization. Homoplasy settings indicate any extra step as Our concepts for male genitalic homology rendering a character homoplastic. used the study of Weirauch (2007) as a Both the equal-weights and the implied- baseline, because the genera Polyozus and weights analyses recover with strong support Exocarpocoris Weirauch possess features in the Melaleucoides genus group and the three common with the Melaleucoides group of genera described in this paper (figs. 35–37). genera. Nonetheless, material examined for This grouping is supported by a number of the present study showed structural variation characters, among which the fleshy structure that was not always easily interpreted under of the parempodia (23-1) is easily recognized Weirauch’s (2007) homology theories. There- and apparently unique within the Australian fore, in arriving at the matrix presented in Phylinae fauna. Harpgophylus is recognized this paper, we employed a variety of alterna- as the sister group of the remaining members tive codings for the endosomal spines found of the Melaleucoides genus group. Under our apically and in association with the second- character codings none of the characters ary gonopore. diagnostic for Harpgophylus is unique, but Fig. 35. Phylogenetic relationships within the Melaleucoides genus group based on equal weight analysis using TNT and maximum parsimony. L 5 506, CI 5 35, RI 5 62. Fig. 36. Phylogenetic relationships within the Melaleucoides genus group based on implied weights analysis using TNT and a constant of concavity of 3. Fit 5 35.982. 2010 SCHUH AND WEIRAUCH: MYRTACEAE-FEEDING PHYLINAE 75

Fig. 37. Host relationships of Melaleucoides group plotted on implied-weights phylogenetic analysis in figure 36. 76 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 344

TABLE 2 Hosts of Melaleucoides genus group species

Host genus No. of and species Specs. Locality Miridae species

Beaufortia micrantha 39 WA: Fitzgerald River National Park, Hammersley M. beaufortiae Road Beaufortia micrantha 25 WA: 92.5 km W of Coolgardie at east side of M. micranthae micrantha on Great Eastern Hwy Calytrix glutinosa 71 WA: 11 km S of Eneabba, Eneabba National Park M. verticordiae Calytrix variabilis 10 WA: Kalbarri National Park, Meanarra Hill H. calyrtrix (pl. 5A) Darwinia diosmoides 8 WA: Kalbarri National Park, Loop Road T. yalgoo Malleostemon 1 WA: ca. 1 km S of Murchison House HS, T. kalbarri hursthousei Kalbarri National Park Melaleuca brevifolia 57 SA: Scorpion Springs Cons. Park M. brevifoliae Melaleuca 7 WA: 15 km E of Merredin M. leuropomae; conothamnoides 2 WA: 15 km E of Merredin M. similis Melaleuca laetifica 30 WA: Kalbarri National Park, 37.7 km E Kalbarri M. systenae (pl. 5A) Melaleuca leuropoma 29 WA: Kalbarri National Park, 7 km E of Kalbarri M. leuropomae (pl. 5C) Melaleuca 19 WA: Kalbarri National Park, Loop Road M. verticordiae megacephala Melaleuca 11 WA: Brand Hiway 8.2 km N of Eneabba M. rhaphiophyllae rhaphiophylla (pl. 5D) 1 WA: Brand Hiway 8.2 km N of Eneabba M. uncinatae Melaleuca sheathiana 186 WA: 11 km N of Coolgardie-Esperance Hiway on M. annae (pl. 6A–C)) Kambalda Road 100 WA: 11 km N of Coolgardie-Esperance Hiway on M. sheathianae Kambalda Road; 33.3 km S of Norseman Melaleuca sp. 2 WA: Moorine Rocks, 11.7 km N of Great Eastern M. rhaphiophyllae Hiway on Noongar Road Melaleuca sp. 2 WA: , 37 km E of Lake M. systenae King Melaleuca systena 22 WA: 4.5 km S of Jurien on Indian Ocean Rd; M. systenae WA: 8.2 km E of Indian Ocean Rd on Coorow-Greenhead Rd, Lesuer National Park Melaleuca teuthidoides 20 WA: 91.4 km SE of Southern Cross M. sheathianae Melaleuca uncinata 69 WA: 56.6 km W of Yalgoo M. uncinatae (pl. 6D) 204 WA: 56.6 km W of Yalgoo M. cassisi Melaleuca undulata 44 WA: Kalbarri National Park, 22.9 km E Kalbarri M. undulatae (pl. 6E) 4 WA: 4 km W of Lillian Stoke Rock M. castanea Melaleuca urceolaris 5 WA: 11 km S of Eneabba on Brand Hiway, M. systenae Eneabba Reserve Melaleuca viminea 4 WA: Cervantes M. leuropomae; M. systenae Micromyrtus 11 WA: NW Coastal Hiway 58 km N of Kalbarri M. akaina hursthousei (pl. 7A) Road Phymatocarpus 5 WA: Brand Hiway 45.9 km S of Dongarra Road M. leuropomae; porphyrocephalus 5 WA: Brand Hiway 45.9 km S of Dongarra Road M. systenae Pileanthus 102 WA: Kalbarri National Park, Loop Road; Kalbarri M. verticordiae peduncularis National Park, Z-Bend Road; NW Coastal Hiway borealis (pl. 7B) 57 km N of Kalbarri Road 128 WA: Kalbarri National Park, Loop RoaD; NW M. pileanthicola Coastal Hiway 57 km N of Kalbarri Road 2010 SCHUH AND WEIRAUCH: MYRTACEAE-FEEDING PHYLINAE 77

TABLE 2 (Continued)

Host genus No. of and species Specs. Locality Miridae species Pileanthus 3 WA: 66.2 km E of North West Coastal Hiway on M. pileanthicola peduncularis subsp. Mardathuna Rd pilifer (pl. 7D) Pileanthus vernicosus 59 WA: 54.3 km N of jct of Agana Kalbarri Rd and M. verticordiae Brand Hiway (rest area); WA: ca. 1 km S of Murchison House HS, Kalbarri National Park WA: 54.3 km N of jct of Agana Kalbarri Rd and M. pileanthicola Brand Hiway (rest area) Scholtzia drummondii 5 WA: Brand Hiway 55.9 km S of Dongarra Road M. verticordiae (pl. 8A) 1 WA: 11 km S of Eneabba, Eneabba National Park M. similis Scholtzia leptantha 11 WA: 4.5 km NW of jct of Blowholes Rd and M. akaina North West Coastal Hiway, N of Carnarvon 1 WA: 4.5 km NW of jct of Blowholes Rd and M. pileanthicola North West Coastal Hiway, N of Carnarvon 8 WA: 4.5 km NW of jct of Blowholes Rd and H. scholtzii North West Coastal Hiway, N of Carnarvon Thryptomene aspera 6 WA: 28 km S of Menzies (3.5 km E of Hiway) M. pileanthicola aspera (pl. 8B) 23 WA: 28 km S of Menzies (3.5 km E of Hiway) H. thryptomeni Thryptomene aspera 47 WA: 24 km W of Sandstone M. verticordiae glabra 2 WA: 24 km W of Sandstone M. similis 30 WA: 24 km W of Sandstone; WA: 46.5 km W T. yalgoo of Yalgoo 20 WA: 31.7 km W of Agnew toward Sandstone H. agnew Thryptomene kochii 21 WA: 135 km W of Coolgardie on Great Eastern Melaleucoidea ozzii Hiway Thryptomene sp. 46 WA: Kalbarri National Park, Loop Road T. kalbarri Verticordia capillaris 19 WA: 54.3 km N of jct of Agana Kalbarri Rd and M. verticordiae Brand Hiway (rest area) Verticordia 421 WA: 1 km S of Lillian Stoke Rock; 11 km S of M. verticordiae chrysantha Eneabba, Eneabba National Park; 123 km W of (pl. 8C, D) Coolgardie on Great Eastern Hiway; 20.6 km S of Norseman-Lake King Road on Lake King-Cascades Road; 3.5 km E of Lillian Stoke Rock; Eneabba on Brand Hiway 1 WA: 1 km S of Lillian Stoke Rock M. systenae Verticordia 53 WA: 11 km S of Eneabba, Eneabba National Park M. verticordiae chrysanthella Verticordia densiflora 71 WA: Eneabba on Brand Hiway M. verticordiae Verticordia forrestii 5 WA: North West Coast Hiway 72 km NE of jct H. verticordii with Blowholes Rd Verticordia 24 WA: Kalbarri National Park, Loop Road H. verticordii monadelpha (pl. 9A) Verticordia polytricha 69 WA: Kalbarri National Park, Loop Road M. verticordiae (pl. 9B) 26 WA: Kalbarri National Park, Loop Road H. verticordii Verticordia roei 4 WA: 3.5 km E of Lillian Stoke Rock M. verticordiae

the monophyly of the group is nonetheless Thryptomenomiris and Melaleucoides is diag- strongly supported in our analyses; the form nosed by three characters, the most obvious of the right paramere (60-2) is easily recog- of which is the vertical orientation of the left nized. The monophyletic group combining paramere (50-1). Under our character codings 78 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 344 none of the characters diagnostic for Thryp- Myrtaceae. According to Wilson et al. (2005), tomenomiris is unique, but the monophyly of 15 tribes are currently recognized in this the group is nonetheless strongly supported in subfamily including the Melaleuceae (e.g., our analyses; the form of the left paramere Melaleuca, ), Leptospermeae (56-2) is the most easily recognized. The (e.g., Kunzea, Agonis), and Chamelaucieae monophyly of Melaleucoides is recognized (e.g., Verticordia, Thryptomene), all of which by the left paramere extending vertically to are speciose in Australia. The phylogenetic the dorsal margin of the pygophore (53-1), a analyses by Wilson et al. (2001, 2005) feature seen nowhere else in the Phylinae. demonstrated that Chamelaucieae and Lep- Although the form of the left paramere in tospermeae are sister taxa and that Melaleu- Melaleucoides shows great consistency across ceae are more distantly related to that clade; the group, the structure of the endosoma is species in the Melaleucoides genus group rather variable. Most species of Melaleu- occur on species in the Chamelaucieae and coides have two long apical spines and at Melaleuceae. Harpagophylus spp. and Thryp- least one additional spine associated with the tomenomiris spp. are restricted to Chamelau- secondary gonopore. Novel within the group cieae and Melaleucoides spp. are known to be is M. akaina, which has a single, short, apical associated with both tribes. We mapped host spine and no processes associated with the genera on the implied weights tree and the secondary gonopore. Nonetheless, all of our following pattern appears: the association phylogenetic analyses place M. akaina within with species in the Chamelaucieae is the Melaleucoides because of the structure of the ancestral host association for the genus group parempodia and the left paramere. and is seen in Harpagophylus, Thrpytome- The tree topologies resulting from our nomris, and the basal taxa within Melaleu- analyses differ with respect to certain species- coides (fig. 37) representing five species. level relationships within Harpagophylus and Within the genus Melaleucoides a host switch Melaleucoides. Among the clades within occurred to the Melaleuceae, specifically Melaleucoides recovered in both analyses species of Melaleuca and Beaufortia, and are, e.g., a clade that comprises M. undulatae, the 12 remaining species of Melaleucoides for M. rhaphiophyllae,andM. uncinatae and a which host plants are known are restricted to clade that consists of M. beaufortiae, M. these two genera. leuropomae, and M. micranthae. DISTRIBUTIONAL PATTERNS HOST RELATIONSHIPS Twenty-three of the 25 species placed in All species in the Melaleucoides genus the Melaleucoides genus group are known group are associated with plant species in only from a limited region in southwestern the family Myrtaceae, subfamily Myrtoideae Western Australia; these distributions are sensu Wilson et al. (2005) (table 2; fig. 37). shown in maps 1–6. The other two are known The majority of plant bug species were from the southeastern costal region of South recorded from one or very few host plant Australia, as shown in maps 3 and 4. One species (table 2). A noteworthy exception is might conclude that this distribution is the M. verticordiae, the most commonly collected result of sampling bias. On the contrary, G. species in the genus group (976 specimens), Cassis, R.T. Schuh, and their colleagues have which was recorded from 13 plant species collected over a wide area of Australia, using comprising a wide range of Myrtaceae. techniques similar to those applied in the Several plant species harbored more than capture of members of the Melalecoides genus one species of plant bug in the Melaleucoides group. And they have captured many other genus group, among them Melaleuca rha- species of Phylinae breeding on species of phiophylla, Melaleuca viminea, and Verticor- Myrtoideae as part of those efforts. Thus, we dia polytricha (table 2). conclude that these restrictions to the western The Melaleucoides genus group appears to part of the continent are not an artifact, but have a relatively clear-cut evolutionary pat- represent a real aspect of the distribution of tern of host associations within the family this lineage. 2010 SCHUH AND WEIRAUCH: MYRTACEAE-FEEDING PHYLINAE 79

Because we do not have a broader Cassis for the study of the Miridae subfam- sampling of phylogenetic relationships for ilies Orthotylinae and Phylinae. Australian Phylinae, it is not possible to Funding for fieldwork was provided by the specify the distribution of the sister group of Australian Museum (Western Australia, the Melaleucoides genus group. Nonetheless, 1996, 1999; South Australia, 1998), the the distribution of the Melaleucoides group American Museum of Natural History species has great similarity with distributions (Western Australia, 1996, 1999; South Aus- seen in monophyletic species groups within tralia, 1998), the National Geographic Soci- the recently described genus Wallabicoris ety (Western Australia, 1997), and the NSF (Schuh and Pedraza, 2010). No meaningful PBI award (Western Australia, 2004). biogeographic statements can be made on the All material collected by R.T. Schuh, G. basis of two presumed areas of endemism as Cassis, and their collaborators was acquired seen in the Melalecoides genus group (SW, under permits issued by the Department of SE). Correspondence with broader biogeo- Environment and Heritage in South Austra- graphic theories will have to await improved lia and Department of Conservation and knowledge of phylogenetic relationships Land Management in Western Australia within the Australian Phylinae. (1996, License No. SF001992; 1997, License No. SF002354; 1999, License No. NE002337). ACKNOWLEDGMENTS We thank these agencies for their coopera- tion. We offer special thanks to Gerasimos For the loan of specimens we thank the Cassis, University of New South Wales, for following institutions and their respective his contribution to the existing collections in collections management personnel: David the Melaleucoides group of genera. His Britton (Australian Museum, Sydney) and efforts in the field are to a very great degree Jan Forrest (South Australian Museum, responsible for bringing together the material Adelaide). studied during the course of this project. We thank the Willi Hennig Society for Other persons whose contributions to this supporting the phylogenetics program project require individual acknowledgment T.N.T. and for making that software freely because of their direct bearing on its com- available for our use in the preparation of pletion for publication include: Steve Thur- this paper. Our thanks also to the authors, ston (AMNH), who prepared the habitus P.A. Goloboff, J.S. Farris, and K.C. Nixon, images, inked and digitized the genitalic for their continuing contributions to the field illustrations, and assembled all of the art- of phylogenetic analysis. work in its final form; Kate Carmody, who For comments on the manuscript we thank entered specimen data and prepared most of Thomas J. Henry, Michael D. Schwartz, and the measurements; Christine A. Johnson, Alfred G. Wheeler, Jr. American Museum of Natural History (AMNH), who prepared some of the speci- REFERENCES men measurements; Matthew Frankel of the AMNH Microscopy and Imaging Facil- Australia’s Virtual Herbarium, Commonwealth of ity, who assisted with scanning electron Australia and others. Internet resource (http:// microscopy; and the collections staffs at www.anbg.gov.au/avh/), accessed August 31, the Western Australian Herbarium, Depart- 2010. ment of Conservation and Land Manage- Carvalho, J.C.M., and G.F. Gross. 1982. Austra- ment, Perth, and the Royal Botanic Gar- lian ant-mimetic Miridae (Hemiptera: Heterop- dens, Sydney, who provided authoritative tera). I. The Leucophoroptera group of the subfamily Phylinae. Australian Journal of Zo- host identifications. ology, suppl. ser., no. 86: 1–75. This project represents a contribution to, Cassis, G. 2008. The Lattinova complex of and was partially supported by, the National austromirine plant bugs (Hemiptera: Heterop- Science Foundation (NSF) Planetary Biodi- tera: Miridae: Orthotylinae). Proceedings of the versity Inventories (PBI) award (DEB- Entomological Society of Washington 110: 0316495) to Randall T. Schuh and Gerasimos 845–939. 80 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 344

Cassis, G., and G.F. Gross. 1995. Zoological the vulvar area and the vestibulum in Orthoty- catalogue of Australia. 27.3A. Hemiptera: linae and Phylinae (Heteroptera: Miridae). Heteroptera (Coleorrhyncha to Cimicomorpha. Denisia 19: 557–570. Australian Biological Resources Study. Mel- Schaffner, J.C., and M.D. Schwartz. 2008. Revi- bourne: CSIRO, 506 pp. sion of the Mexican genera Ficinus Distant and Cassis, G., M.A. Wall, and R.T. Schuh. 2007. Jornandes Distant with the description of 21 Insect biodiversity and industrializing the taxo- new species (Heteroptera: Miridae: Orthotyli- nomic process: a case study with the Miridae nae: Orthotylini). Bulletin of the American (Heteroptera). In Trevor. Hodkinson and John. Museum of Natural History 309: 1–87. Parnell (editors), Towards the tree of life: Schuh, R.T., and K. Menard. In prep.. Lorantha- and systematics of large and species ceae-feeding plant bugs from Australia and South rich clades: 193–212. Boca Raton, FL: CRC Africa (Heteroptera: Miridae: Phylini): ten new Press. species and their placement in the Pilophorini. Eyles, A.C., and R.T. Schuh. 2003. Revision of Schuh, R.T., and P. Pedraza. 2010. Wallabicoris, New Zealand Bryocorinae and Phylinae (Insec- new genus (Hemiptera: Miridae: Phylinae: ta: Hemiptera: Miridae). New Zealand Journal Phylini), from Australia, with the description of Zoology 30: 263–325. of 37 new species and an analysis of host Forero, D. 2008. Revision and phylogenetic associations. Bulletin of the American Museum analysis of the Hadronema group (Miridae: of Natural History 338: 1–118. Orthotylinae: Orthotylini), with descriptions of Soto, D., and C. Weirauch. 2009. Description of new genera and species, and comments on the the Australian plant bug genus Jiwarli,n.gen. Neotropical genus Tupimiris. Bulletin of the (Heteroptera: Miridae: Phylinae). American American Museum of Natural History 312: Museum Novitates 3653: 1–14. 1–172. Weirauch, C. 2007. Revision and cladistic analysis Goloboff, P.A. 1993. Estimating character weights of the Polyozus group of Australian Phylini during tree search. Cladistics 9: 83–91. (Heteroptera: Miridae: Phylini). American Mu- Goloboff, P.A., J.A. Farris, and K.C. Nixon. 2008. seum Novitates 3590: 1–60. TNT, a free program for phylogenetic analysis. Weirauch, C., and R.T. Schuh. In press. Southern Cladistics 24: 774–786. hemisphere patterns in plant bugs: Xiphoidellus, Goloboff, P.A., J.A. Farris, and K.C. Nixon. 2009. new genus (Hemiptera: Miridae: Phylinae) from T.N.T. Tree analysis using new technology. Australia and Ampimpacoris, new genus from Version 1.1. Willi Hennig Society Edtion. Argentina show transatlantic relationships. In- Available from the authors. vertebrate Systematics. Malipatil, M. 1992. Revision of Australian Cam- Wilson, P.G., M.M. O’Brien, P.A. Gadek, and pylomma Reuter (Hemiptera: Miridae: Phyli- C.J. Quinn. 2001. Myrtaceae revisited: A nae). Journal of the Australian Entomological reassessment of infrafamilial groups. American Society 31: 357–368. Journal of 88: 2013–2025. Nixon, K.C. 2000. WinClada, version 0.9.99.60 Wilson, P.G., M.M. O’Brien, M.M. Heslewood, (program). Ithaca, NY: Published by the and C.J. Quinn. 2005. Relationships within author. Myrtaceae sensu lato based on a matK phylog- Pluot-Sigwalt, D., and A. Matocq. 2006. On some eny. Plant Systematics and Evolution 251: particular sclerotized structures associated with 3–19. 2010 SCHUH AND WEIRAUCH: MYRTACEAE-FEEDING PHYLINAE 81

APPENDIX 1 DATA MATRIX AND CHARACTER DESCRIPTIONS

0 5 10 15 20 25 30 35 40 45 50 55 60 65 70 75 | | | || | || |||| | || |

Leucophoropterini sp. 0033300122330-4201300200102000-02--0000000-000000000100000000102010000000001 Coatonocapsus sp. 0243330450020061124102001002030--000000102-100000000100001010102010000000004 Capecapsus tradouwensis 0243330450020061124102001002030--000000102-100000000100001010102010000000004 Wallabicoris ozothamni 1300110301010-0212100210100101011--0000000-1000010002000011101020101200111-4 Wallabicoris pityrodii 131111031202011110210210100101011--0000000-100001000300001111102010120011024 Wallabicoris spyridii 231114250202013112310110100102010--0000100-1000010001000011111000101200111-4 Xyphoides myersi 204320000233116312310310110203012--0000100-110000000100000200311211001000004 Araucanophylus pacificus 2443200002330161123203101102020----0000100-000000000100001200102001000000004 Xyphoidellus aureus 250215364124120402320220110203010--10000011000000000100000010312001001100024 Xyphoidellus furvus 202210304134124402320220120203010--10000010000000000100000120312101001100024 Xyphoidellus pallidus 130011020010120402420120110203010--10000010000000000100000010311001001100024 Jiwarli heliotropium 230233041101000301310120102002021--0000100-0000000001000020001000101100111-3 Polyozus galbanus 161131070112000111300310102003023100020000-000000000200000200102310110011002 Exocarpocoris tantulus 3611310801020001134201?0202003023100010000-000000000200000110102010110011020 Protemiris conospermi,ms 214013040031000212421000212010001000000000-000000000300100111201010121110005 Harpagophylus agnew 260011020200020214201121002013121-00110000-1000000001000013020010------Harpagophylus calytrix 104432056253026012301121002013121-00110000-2----01001000033021010------Harpagophylus scholtzii 260011020250020213301121002013121-00110000-20---01001000013020010------Harpagophylus thryptomeni 260012020250020213211111002013121-00110000-20000010010000330 1010110010002 Harpagophylus verticordii 260011020250020212310221002013121-00120000-2000001001000013020010101100101-2 Thryptomenomiris kalbarri 36001102625002001221021100201313--00020000-000010010002120000001010110010012 Thryptomenomiris yalgoo 26001125025002021231012100201313--00020000-000110010002120200001010110010012 Melaleucoides akaina 161111030250020--2300121002000122000000010-000100010011301100001310110011013 Melaleucoides annae 261236452201010014210111102013123010020100-201100010011101000001010110021002 Melaleucoides beaufortiae 273233644202000002220021102013123010000100-200000010011201200001010110021003 Melaleucoides brevifoliae 33003364520200001221011110------2010000100112011000010------Melaleucoides cassisi 261112420250020014311111102013123010020100-101000010011201000002010110021003 Melaleucoides castanea 404110000250026003320111102013123010010100-2010100100111010002000------Melaleucoides grossi 361112420020020014211111102013123010020100-0011000100111011002000------Melaleucoides leuropomae 230213644202000003320011102013123020020100-200100010012211000001010110021003 Melaleucoides micranthae 331103044212010004220011102013123010020100-0001000100112102000010------Melaleucoides ozzii 160014462200010201410111002012123010100100-2001100100111000000010------Melaleucoides pileanthicola 360011250250020011400111002013123000010100-201000010011101110000010110010002 Melaleucoides rhaphiophyllae 10233031220200350322001110------2100000110110012200013------Melaleucoides sheathianae 264112220250020004210111102013123010020100-001000010011101000001010110021002 Melaleucoides similis 061112020210020002311121002013221000121100-2000000100121210000020------Melaleucoides systenae 3244375-2203016003320111102013121--0021100-201010010011101000201310110021003 Melaleucoides uncinatae 102330312203003504210111102011121--0201000-110000011011101220002310110011003 Melaleucoides undulatae 270016494102000504210111102011121--0101000-2110000110110000200010------Melaleucoides verticordiae 061112020210020011411121002013321000120000-000010010011101020002010110011002 82 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 344

CHARACTER DESCRIPTIONS white, cuneus white 5 8; pale with longitu- dinal castaneous stripes 5 9. [nonadditive] 8. Markings on cuneus: Absent 5 0; present as 0. Body: Greatly elongate, parallel sided 5 0; partial or complete but weak infuscation 5 moderately elongate, parallel sided 5 1; 1; present as a contrasting white basal fascia weakly elongate, weakly ovoid 5 2; weakly 5 2; present as pale narrow band, red over to distinctly ovoid 5 3; broadly ovoid 5 4. remaining area 5 3; present as a red or [additive] castaneous apex 5 4; present as small dark dots 5 5; absent, cuneus transparent 5 6. COLORATION [nonadditive] 5 5 9. Membrane: Unicolorous pale 0; partially 1. Head: Uniformly dark 0; pale with dark fumose 5 1; weakly to strongly fumose over 5 clypeus 1; mostly dark with pale gula and entire area 5 2. [nonadditive] 5 posterior head margin 2; pale or pale with 10. Membrane veins: White or pale 5 0; yellow 5 5 red or dark dots 3; dark with pale mark at 1; entirely red 5 2; dark or dark red 5 3; 5 medial eye margin 4; Pale with red pale with distal portion of anterior cell red 5 5 suffusion on gena 5; Uniformly pale 5 4; unicolorous with membrane 5 5. 6; Pale, clypeus dark and large dark bilateral [nonadditive] 5 markings on frons 7. [nonadditive] 11. Hind femur: Unicolorous pale, without black 5 2. Scapus: Unicolorous pale 0; yellow to dirty spots 5 0; with a few brown or black spots 5 5 yellow 1; brown 2; pale with a dark base 5 1; with many brown or black spots 5 2; 5 5 3; unicolorous dark 4. [nonadditive] nearly unicolorous dark 5 3; pale with distal 5 3. Pedicellus: Unicolorous pale 0; yellow to red suffusion 5 4. [nonadditive] 5 dirty yellow 1; dirty yellow proximally, 12. Hind tibial spines:Dark5 0; pale 5 1. 5 weakly to heavily infuscate distally 2; 13. Hind tibial spines (bases): With conspicuous 5 unicolorous black or castaneous 3; black dark spots at bases 5 0; with very small 5 at extreme base, remainder pale 4. dark bases 5 1; without dark spots at bases [nonadditive] 5 2. [nonadditive] 5 4. Labium: Uniformly pale 0; pale with 14. Abdominal venter: Unicolorous pale or mostly 5 segment 4 heavily infuscate 1; unicolorous so 5 0; light green 5 1; light yellow 5 2; red 5 2; generally infuscate, heavily so heavily red laterally 5 3; heavily infuscate, apically 5 3. [nonadditive] at least on mesopleuron and abdomen 5 4; 5. Thorax: Pronotum: Unicolorous deep red to with thorax pale and abdomen dark 5 5; black 5 0; unicolorous pale 5 1; unicolor- unicolorous dark 5 6. [nonadditive] ous (yellow, brown, or green) 5 2; pale with brown spots 5 3; pale with red or carmine blotches 5 4; pale with orange tinge, SURFACE AND VESTITURE anterior lobe cream with dark marks 5 5; 15. Dorsal common setae: With erect, suberect, or pale with castaneous cali and humeri 5 6; reclining black setae 5 0; with black setae usually with dark anterior lobe and con- intermixed with pale setae on hemelytra 5 1; trasting posterior lobe 5 7. [nonadditive] with reclining simple setae matching back- 6. Scutellum: Unicolorous with remainder of ground coloration 5 2; with subadpressed dorsum 5 0; with some red or carmine flattened and erect simple pale setae 5 3; spots or blotches on pale background 5 1; with simple pale suberect setae 5 4; with entirely red or orange 5 2; red or dark golden reclining setae 5 5. [nonadditive] brown with white or pale tip 5 3; white 5 4; 16. Dorsal woolly setae: Without sericeous or entirely dark 5 5; pale with brown spots 5 woolly setae 5 0; with some sericeous or 6. [nonadditive] woolly setae 5 1. 7. Hemelytron: Unicolorous, deep red to black 5 0; castaneous to black with contrasting STRUCTURE white markings 5 1; unicolorous [pale, white, or yellow] 5 2; mostly pale, or 17. Head: Elongate, projecting anteriorly 5 0; yellow, endocorium with dirty or golden moderately projecting 5 1; somewhat pro- areas 5 3; unicolorous pale with small jecting 5 2; weakly projecting 5 3; barely brown spots 5 4; with red or carmine spots, projecting, 5 4. [nonadditive] botches, or solid areas 5 5; pale yellow with 18. Interocular space: Relatively small 5 1; mod- red suffusion 5 6; brown with white cuneus erate 5 2; relatively large 5 3; large 5 4. 5 7; with pattern of yellow, brown, and [additive] 2010 SCHUH AND WEIRAUCH: MYRTACEAE-FEEDING PHYLINAE 83

19. Eyes: Leaving gena only very slightly exposed to gonopore 5 1; fused with primary strap in lateral view 5 0; leaving gena moderately 5 2; absent 5 3. [additive] exposed in lateral view 5 1; leaving gena 32. Secondary endosoma strap (2): Reaching mid- broadly exposed in lateral view 5 2. way to gonopore from major bend in vesica [additive] 5 0; reaching to level of secondary gono- 20. Antenna: Segment 2: weakly tapering, more pore 5 1; reaching just beyond secondary slender at base 5 0; of equal diameter over gonopore as fingerlike extension 5 2; entire length 5 1. reaching well beyond gonopore 5 3. [non- 21. Antennal fossa: With dorsal margin somewhat additive] below ventral margin of eye 5 0; with 33. Secondary endosomal strap (3): With one tip 5 ventral margin at ventral margin of eye 5 1; 0; anchor shaped 5 1. with ventral margin 1 diameter above 34. Secondary endosomal strap (4): Devoid of ventral margin of eye 5 2; with ventral ornamentation 5 0; with denticles 5 1; with margin 2 diameters above ventral margin of a comblike edge 5 2. [additive] eye 5 3. [nonadditive] 35. Fingerlike protuberance at distal margin of 22. Labium: Reaching to posterior margin of secondary gonopore: Absent 5 0; present 5 mesosternum 5 0; just reaching onto 1. abdomen 5 1; reaching to about anterior 36. Spinelike, elongate process arising near gono- margin of pygophore 5 2. [additive] pore on dorsal surface: Absent 5 0; present, 23. Paremopodia: Setiform 5 0; fleshy, recurved. erect, unornamented 5 1; present, erect, lyriform 5 1. ornamented with denticles 5 2. [additive] 24. Pulvilli: Absent 5 0; present, flaplike, covering 37. Spinelike, elongate process near gonopore on about one third of ventral claw surface 5 1; ventral surface: Absent 5 0; present and present, flaplike, covering most of ventral erect 5 1; present, recurved toward base of surface of claw 5 2. [additive] endosoma 5 2. [nonadditive] 25. Pygophore: Without row or tuft of setae on left 38. Bladderlike process distad of secondary gono- side 5 0; with dense tuft of setae on left side pore: Absent 5 0; present 5 1. 5 1; with double row of setae on left side 5 39. Secondary gonopore: Seen laterally in lateral 2. [nonadditive] view of vesica 5 0; seen frontally (facing up) 26. Endosoma base: Very long 5 0; moderately in lateral view of vesica 5 1. long 5 1; short 5 2. [nonadditive] 40. Secondary gonopore (2): Without a denticulate 27. Endosoma shape: Curving, often C- or J- ridge on right-hand surface of endosoma 5 shaped 5 0; with a U-shaped bend 5 1; 0; with a denticulate ridge on right-hand forming a single complete coil 5 2. [addi- surface of endosoma 5 1. tive] 41. Secondary gonopore sclerite (1): Absent 5 0; 28. Endosoma body: With torsion, primary strap present 5 1; absent, but short, slender dorsal to secondary gonopore 5 0; without sclerite close to gonopore 5 2. [nonadditive] torsion, primary strap ventral to secondary 42. Secondary gonopore sclerite (2): Separated gonopore 5 1. from secondary endosomal strap at apex 5 29. Primary endosomal strap apically: Short, ratio 0; apparently fused to secondary endosomal length apex strap/length secondary gono- strap at apex 5 1. pore 1.33–1.50 5 0; weakly elongate, length 43. Phallotheca: Smoothly curving on dorsal apex strap/length secondary gonopore 1.67– margin 5 0; more or less right angulate 5 2.00 5 1; elongate, ratio length apex strap/ 1; nearly erect 5 2. [nonadditive] length secondary gonopore 2.15–2.83 5 2; 44. Dorsal surface of phallotheca: Without a greatly elongate, length apex strap/length fingerlike projection 5 0; with a fingerlike secondary gonopore 3.00–8.6 5 3. [additive] projection 5 1. 30. Primary endosoma strap apically (2): At least 45. Dorsal surface of phallotheca (2): Without a weakly arcuate and curving ventrally 5 0; at platelike projection 5 0; with a curving, least weakly arcuate and curving dorsally, posteriorly directed, platelike projection 5 1. no ornamentation 5 1; at least weakly 46. Anterior surface of phallotheca: Without a keel arcuate and curving dorsally, denticulate 5 0; with a short to elongate keel 5 1. margin 5 2; at least weakly arcuate and 47. Ventral surface of phallotheca: Without a curving dorsally, comblike margin 5 3. projecting keel 5 0; with a projecting keel [additive] 5 1. 31. Secondary endosomal strap: Broad, about 48. Posterior surface of phallotheca: Lacking equal in width to primary strap 5 0; very transparent window 5 0; with conspicuous slender, of uniform width from vesical bend transparent window 5 1. 84 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 344

49. Apex of phallotheca: Simple 5 0; complex 5 1. FEMALE 50. Left Paramere: Horizontal 5 0; vertical 5 1. 5 51. Left paramere shaft: At right angles to body 5 63. Body shape:Asinmale 0; slightly more 5 0; in same axis as body 5 1. ovoid than male 1; distinctly more ovoid 5 52. Left paramere in dorsal perspective:Not than male 2. [additive] exceeding margin of pygophore 5 0; just 64. Coloration: As in male 5 0; more vividly red exceeding margin of pygophore 5 1; some- than male 5 1; as in male but with cuneus what exceeding pygophore margin 5 2; bright red 5 2; paler than male 5 3. greatly exceeding margin of pygophore 5 [nonadditive] 3. [nonadditive] 65. Posterior wall laterally: Without interramal 53. Left paramere in lateral perspective: Covering sclerites 5 0; with distinct crescent-shaped limited extent of lateral surface of pygo- interramal sclerites 5 1. phore 5 0; erect and reaching dorsal margin 66. Posterior wall posteriorly: Without a sclero- of pygophore or beyond 5 1. tized transverse band 5 0; with a sclerotized 54. Left paramere body: Narrowed toward apex 5 transverse band 5 1. 0; spoon shaped 5 1; more or less quadrate 67. Posterior wall posterolaterally: Without a 5 2. [nonadditive] distinct swelling covered with microtrichia 55. Left paramere apex medially: Angulate 5 0; 5 0; with a distinct swelling covered with broadly rounded to weakly quadrate 5 1; microtrichia 5 1. drawn into a blunt fingerlike process 5 2; 68. Posterior wall longitudinal midline: Undiffer- with an elongate, acuminate, recurved me- entiated 5 0; fold on either side of midline dial process 5 3. [nonadditive] along part of length 5 1; fold on either side 56. Left paramere apex posterolaterally: Simple 5 of midline along most of length 5 2. 0; with a low shoulder 5 1; with a greatly [additive] elongate projection 5 2. [additive] 69. Posterior wall posterior margin: Without spic- 57. Left paramere anterior process: Slender and ules 5 0; ornamented with spicules 5 1. cylindrical in lateral view 5 0; triangular in 70. Posterial wall posterior margin (2): Not re- lateral view 5 1; broad and elongate in flexed dorsally 5 0; reflexed dorsally 5 1. lateral view 5 2; undeveloped 5 3. [nonad- 71. Posterior wall interramal lobes: Absent 5 0; ditive] present and roughly symmetrical, erect 5 1; 58. Left paramere posterior process: In the form of present and asymmetrical, reclining, over- a short, straight, fingerlike projection 5 0; lapping 5 2. [additive] in the form of a long, straight, fingerlike 72. Posterior wall interramal lobes (2): Not orna- projection 5 1; in the form of a short curved mented with spicules or with a very few 5 0; fingerlike projection 5 2; very long, curving heavily ornamented with spicules 5 1. 90u toward base of paramere 5 3. [nonad- 73. Vestibulum medial plates (1): Sclerotized and ditive] readily observed 5 0; weakly sclerotized or 59. Left paramere base of posterior process:With- apparently absent 5 1. out conspicuous shoulder 5 0; with slightly 74. Vestibulum medial plates (2): Small, nearly projecting shoulder 5 1; with distinct symmetrical, triangular 5 0; large, nearly shoulder 5 2. [nonadditive] symmetrical 5 1; right plate larger, more 60. Right paramere body: Short and broad, lance- strongly elongate than left 5 2. [nonaddi- olate 5 0; moderately elongate 5 1; greatly tive] elongate, bulbous at base 5 2. [nonadditive] 75. Vestibulum medial plates sclerotized guide 61. Right paramere apex: Short, clawlike, de- structure: Absent 5 0; present as short curved 5 0; tapered and acuminate 5 1; weakly sclerotized medial tube 5 1; present with short fingerlike process 5 2; with a as caplike structure with internal differenti- very long fingerlike process 5 3. [nonaddi- ation 5 2; present as short, heavily sclero- tive] tized tube on left side 5 3; present as long 62. Right paramere posterior margin of apex: tube on left side doubling back 5 4; present Smooth 5 0; with distinct serration 5 1. as lateral sclerotizations 5 5. [nonadditive] PLATES 2010 SCHUH AND WEIRAUCH: MYRTACEAE-FEEDING PHYLINAE 87

Plate 1. Habitus images of Harpagophylus spp., Thryptomenomiris spp., and Melaleucoides akaina,and M. annae. 88 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 344

Plate 2. Habitus images of Melaleucoides beaufortiae–Melaleucoides leuropomae. 2010 SCHUH AND WEIRAUCH: MYRTACEAE-FEEDING PHYLINAE 89

Plate 3. Habitus images of Melaleucoides micranthae–Melaleucoides similis. 90 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 344

Plate 4. Habitus images of Melaleucoides systenae–Melaleucoides verticordiae. 2010 SCHUH AND WEIRAUCH: MYRTACEAE-FEEDING PHYLINAE 91

Plate 5. Host plants of Melaleucoides genus group taxa. A. Calytrix variabilis Lindl. B. Melaleuca laetifica L.A. Carven. C. Melaleuca leuropomae L.A. Craven. D. Melaleuca rhaphiophylla Schauer. 92 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 344

Plate 6. Host plants of Melaleucoides genus group taxa. A.–C. Melaleuca sheathiana W. Fitz. D. Melaleuca uncinata R. Br. E. Melaleuca undulata Benth. 2010 SCHUH AND WEIRAUCH: MYRTACEAE-FEEDING PHYLINAE 93

Plate 7. Host plants of Melaleucoides genus group taxa. A. Micromyrtus hursthousei W. Fitz. B. Pileanthus peduncularis borealis Endl. C. Pileanthus spp. (foreground). D. Pileanthus peduncularis pilifer Keighery. 94 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 344

Plate 8. Host plants of Melaleucoides genus group taxa. A. Scholtzia drummondii Benth. B. Thryptomene aspera aspera E. Pritz. C–D. Verticordia chrysantha Endl. 2010 SCHUH AND WEIRAUCH: MYRTACEAE-FEEDING PHYLINAE 95

Plate 9. Host plants of Melaleucoides genus group taxa. A. Verticordia monadelpha Turcz. B. Verticordia polytrichia Benth.