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 plants. 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 Western Australia; two species are recorded from southeastern South Australia. All species breed on members of the subfamily Myrtoideae, tribes Chamelaucieae and Melaleuceae.
INTRODUCTION
This paper represents part of an ongoing effort to document the fauna of Miridae, or plant bugs, of Australia, including distributional patterns and associations with host plants. Although in the past this fauna has received only limited study (Carvalho and Gross, 1982; Malipatil, 1992), the nature of its actual diversity is beginning to take shape. Within the Phylini (Phylinae), a group with its greatest diversity in the Mediterranean-climate regions of the world, recent papers have documented well-defined associations with a variety of host lineages within Australia, the occurrence of both Indo-Australian (Schuh and Menard, in prep.) and Transantarctic distributional patterns (Weirauch and Schuh, in press), and radiation of Miridae in association with extreme floristic diversity in southwestern Australia (Weirauch, 2007; Soto and Weirauch, 2009; Schuh and Pedraza, 2010).
The present study details the phylogenetic species-level relationships, classification, and host-plant associations for a monophyletic group of Myrtaceae-feeding taxa that are part of a larger lineage within Phylini whose members are restricted to Australia. The genesis of the paper began with dissections of male genitalia of a comprehensive sample of Australian Phylinae (∼ 250 morphospecies) by coauthor Weirauch during her tenure as a postdoctoral fellow funded by a NSF Planetary Biodiversity Inventories project for the study of Miridae. Those efforts allowed us to recognize the existence of unique endosomal types within certain Australian Phylinae. Further study revealed that not only was the endosoma distinctive, but that the parempodia and left paramere were also novel, permitting the recognition of a monophyletic taxon that we here refer to as the Melaleucoides genus group and that is restricted to host plants within two tribes of the family Myrtaceae: Myrtoideae.
TAXONOMIC MATERIALS AND METHODS
Unique specimen identifiers: During the course of this project matrix code labels were affixed to about 2350 specimens examined as a way to uniquely identify them; these codes are therefore referred to as “unique specimen identifiers” (USIs). The USI codes, e.g., AMNH_PBI 00094810, are composed of an institution and project code (AMNH_PBI) and a unique number (00094810). USI codes are included in locality data, figures, and captions. To reduce the amount of space occupied by the USIs in the material examined sections the prefix (AMNH_PBI) is omitted except for the holotypes.
All latitude-longitude data presented in the specimens examined sections are in degrees and decimal parts thereof; the vast majority of georeferences were obtained with a GPS device in the field. Altitude data are treated as metric.
Comments on descriptions: The descriptions in the present paper were generated from a matrix of character data using WinClada (Nixon, 2000) and then underwent additional editing. Where size serves to help diagnose a taxon, we have included relevant measurement data in the diagnoses and descriptions. Detailed measurements for all species are presented in table 1.
TABLE 1
Measurements of Melaleucoides genus group species
Host-plant identifications: Hosts recorded in this study were pressed in the field and submitted as vouchers to the Western Australian Herbarium or the Royal Botanic Gardens, Sydney, for identification by botanical specialists. Thus, the identifications are authoritative and independent of our own predilections about the identities of the plant groups on which we were collecting. Host voucher numbers and/or locations are listed in the specimens examined sections.
Terminology for female genitalia: We have used the terminology interramal sclerites and interramal lobes for structures of the posterior wall in the female (figs. 6, 7, 11, 18, 25, 34), following the practice established for Orthotylinae by Schaffner and Schwartz (2008) and Forero (2008). Our use of both of these terms in Phylinae should be construed as descriptive in nature, because these structures may not be homologous with those in Orthotylinae.
Measurements: Measurements for all taxa are presented in table 1. Means for total length and pronotal width are presented in the descriptions. Measurements were made using a micrometer driven stage, with micrometer output being written directly to a spreadsheet.
Designation of paratypes: We designate as paratypes specimens that represent a confident association of material with the concept for the species and which were examined as part of the descriptive process.
Deposition of specimens and abbreviations for institutional depositories: Within Australia the collecting permitting process is reserved to the states. Following the stipulations of our permits (see Acknowledgments) holotypes are deposited in the states of origin. Depending on the conditions of the collecting permits, a subset of paratypes may also be deposited in the state or origin. The preponderance of the remaining material is lodged in the institutions that supported the fieldwork (Australian Museum, American Museum of Natural History), with a representative sample of specimens deposited in institutions participating in the Plant Bug Planetary Biodiversity Inventory Project or other institutions of record.
Institutional depositories for specimens utilized during the course of this study and their acronyms as used in this paper are as follows:
AM
Australian Museum, Sydney
AMNH
American Museum of Natural History, New York
ANIC
Australian National Insect Collection, CSIRO, Canberra
CNC
Canadian National Collection of Insects, Ottawa
SAMA
South Australian Museum, Adelaide
UCR
University of California, Riverside
UNSW
University of New South Wales, Sydney
USNM
United States National Museum of Natural History, Smithsonian Institution, Washington, DC
WAMP
Western Australian Museum, Perth
ZISP
Zoological Institute, Russian Academy of Sciences, St. Petersburg
Melaleucoides genus group
Diagnosis
Recognized by fleshy lyriform parempodia (fig. 5C; character 23-1) and clawlike decurved apex of the right paramere (fig. 1; 61-0); other attributes found in the group include the dark spots at the bases of the tibial spines and the female genitalia with interramal sclerites and interramal lobes (fig. 6). The group is also diagnosed by the weakly arcuate and dorsally curving primary endosomal strap of the male genitalia (fig. 1; character 30-1) and the presence of a spinelike elongate process near the gonopore on the ventral surface of the endosoma that is recurved toward the base of the endosoma (fig. 1; 37-2), although these attributes do not occur in all species.
Discussion
Lyriform parempodia occur in all species in the Melaleucoides genus group, whereas other Phylini in Australia and elsewhere possess setiform parempodia. Most species in the group possess the clawlike apex of the right paramere, but based on our observations and cladistic analyses, this character is secondarily modified in several species within the group.
Key to Melaleucoides genus group genera
1. Right paramere greatly elongate, reaching across entire width of pygophore in repose (fig. 5F), with apex in the shape of a claw; left paramere not greatly elongate in dorsoventral perspective, covering about half of height of pygophore in lateral view (fig. 5D), with posterior process very long and curving ventrally (figs. 1–Fig. 2.3); endosoma always with 2 appendages in region of secondary gonoporeHarpagophylus
– Right paramere short, broadly lanceolate, not traversing entire width of pygophore in repose (figs. 8F, 9, 12F, 17F, 19); posterior process of left paramere relatively short, never strongly curving toward venter2
2. Left paramere elongate in dorsoventral perspective, flattened, spoon shaped, and erect (figs. 12D, E, 13, 17E, 19, 23, 24E); apex of left paramere sometimes elongateMelaleucoides
– Left paramere more quadrate, not erect and spoon shaped as above, always with apex elongate (figs. 7, 8E, F, 9), reaching across pygophore in repose (fig. 8F)Thryptomenomiris
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 = left paramere; pe = parempodium; rp = right paramere (AMNH_PBI 00372775).
Diagnosis
Recognized within the Melaleucoides genus group by the unique, greatly elongate right paramere with a bulbous base (60-2) and the long, decurved posterior process of the left paramere (58-3) (figs. 1–Fig. 2.Fig. 3.Fig. 4.Fig. 5.6); size small, total length 2.10–2.63. Potentially confused with Thryptomenomiris, and small species of Melaleucoides on the basis of small size and association with the Myrtaceae. The structure of the parameres in Harpagophylus precludes any confusion with these other groups, however.
Description
Male: Body weakly elongate, weakly ovoid; small, total length 2.10–2.63, width pronotum 0.70–0.96. COLORATION (pl. 1): Pale, with or without reddish areas on dorsum. SURFACE AND VESTITURE (fig. 5B): Dorsum with reclining simple setae matching background coloration and with some sericeous or woolly setae. STRUCTURE: Head (fig. 5A): Somewhat projecting; interocular space relatively large; eyes leaving gena moderately exposed in lateral view. Thorax: Pretarsus (fig. 5C): Parempodia fleshy, recurved, lyriform; pulvilli absent. GENITALIA (figs. 1–Fig. 2.Fig. 3.Fig. 4.Fig. 5.6): Endosoma: Base short, curving, C- or J-shaped; 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, unornamented; spinelike, elongate process present near gonopore on ventral surface, recurved toward base of endosoma; secondary gonopore seen laterally in lateral view of endosoma. 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 triangular to undeveloped; posterior process very long, curving 90° toward base of paramere; base of posterior process without conspicuous shoulder. Right Paramere (fig. 5F): Body greatly elongate, bulbous at base; apex short, clawlike, decurved.
Female: Slightly more ovoid than male; small, total length 2.06–2.70, width pronotum 0.74–1.00. COLORATION (pl. 1): As in male. GENITALIA (fig. 6): Posterior wall laterally with distinct crescent-shaped interramal sclerites; posterolaterally with a distinct swelling covered with microtrichia; longitudinal fold on either side of midline along part of length; interramal lobes present, roughly symmetrical, erect, and not ornamented with spicules or with very few; vestibulum with medial plates weakly sclerotized or apparently absent, with caplike sclerotized guide with internal differentiation.
Etymology
Named for the hooklike form of the right paramere; from the Latin harpago, “grappling hook,” in combination with Phylus; gender masculine.
Discussion
The five new species described below as belonging to the genus Harpagophylus are well supported as a monophyletic group in both the equal weights and the implied weights analyses (figs. 35, 36). Based on overall appearance and their association with Myrtaceae in the tribe Chamelaucieae, they are easily confused with species of Thryptomenomiris, new genus, and the small species within Melaleucoides, new genus. The structure of the left and right parameres clearly distinguishes them from other members in the Melaleucoides genus group.
Key to species of Harpagophylus
1. Dorsum with conspicuous red areas, particularly posterior half of endocorium (pl. 1); apex of phallotheca complex, body of endosoma broad (fig. 2)Harpagophylus calytrix
– Dorsum lacking conspicuous red areas, mostly yellow (pl. 1); apex of phallotheca without such complex ornamentation and body of endosoma relatively slender2
2. Left side of pygophoral opening bearing a distinct clawlike appendage (fig. 4)Harpagophylus thryptomeni
– Left side of pygophoral opening without a distinct clawlike appendage3
3. Endosoma with both processes in region of secondary gonopore very long and recurved toward its base (fig. 6); apex of right paramere very longHarpagophylus verticordii
– Endosoma with at least one process near secondary gonopore erect, perpendicular to axis (figs. 1, 3); apex of right paramere not so long4
4. Dorsal process of endosoma near secondary gonopore longer than ventral process (fig. 1); ventral process with a serrate margin (fig. 1)Harpagophylus agnew
– Dorsal process of endosoma near secondary gonopore shorter than ventral process (fig. 1); ventral process with a simple margin (fig. 3)Harpagophylus scholtzii
Harpagophylus agnew, new species
Diagnosis
Recognized by the pale, yellowish coloration, the endosomal processes both long, straight, and erect, the simple apex of the phallotheca, the long, relatively slender right paramere, and the denticulate anterior process on the left paramere. Most easily confused with H. scholtzii, H. thryptomeni, and H. verticordii on the basis of pale coloration, but those species all differing in the structure of the anterior process of the left paramere, the conformation of the right paramere, the structure of the apex of the phallotheca, and the conformation of the dorsal and ventral endosomal processes.
Description
Male: Body weakly elongate, weakly ovoid; small, mean total length 2.49, mean width pronotum 0.79. COLORATION (pl. 1): Head: Uniformly pale, yellow; scapus and pedicellus unicolorous pale; labium pale with segment 4 heavily infuscate. Thorax: Pronotum scutellum, and hemelytron unicolorous pale, yellow; markings on cuneus absent; membrane very weakly fumose, veins; hind femur unicolorous pale, without black spots; hind tibial spines dark, without dark spots at bases. Abdomen: Venter unicolorous pale or mostly so. SURFACE AND VESTITURE: Dorsum with reclining simple setae matching background coloration and some sericeous or woolly setae. STRUCTURE: Head: Barely projecting; interocular space moderate; eyes leaving gena only very slightly exposed in lateral view. Antenna: Segment 2 of equal diameter over entire length; antennal fossa with ventral margin at ventral margin of eye. Labium: Reaching to about anterior margin of pygophore. GENITALIA (fig. 1): Phallotheca: More or less right angulate; dorsal surface without a fingerlike projection, and without a platelike projection; anterior surface without a keel; ventral surface without a projecting keel; apex simple.
Female: Slightly more ovoid than male; small, mean total length 2.41, mean width pronotum 0.80. COLORATION (pl. 1): As in male. GENITALIA: Not examined.
Etymology
Named for the town of Agnew, Western Australia, near the type locality; a noun in apposition.
Distribution (map 1)
Known only from the type locality in the Goldfields region of Western Australia.
Holotype
AUSTRALIA: Western Australia: 31.7 km W of Agnew toward Sandstone, 27.96227°S 120.4277°E, 800 m, 26 Oct 1996, Schuh and Cassis, Thryptomene aspera glabra E. Pritz. (Myrtaceae), PERTH 05095190, 1♂ (AMNH_PBI 00136500) (WAMP).
Paratypes
AUSTRALIA: Western Australia: 31.7 km W of Agnew toward Sandstone, 27.96227°S 120.4277°E, 800 m, 26 Oct 1996, Schuh and Cassis, Thryptomene aspera glabra E. Pritz. (Myrtaceae), det. Perth 05095190, 3♂ (00373242–00373244), 5♀ (00373246–00373250) (AM), 3♂ (00136499, 00136501, 00373245), 4♀ (00136502, 00136504, 00136506–00136507) (AMNH), 4♀ (00373251–00373254) (WAMP).
Diagnosis
Recognized by the heavily reddish coloration on much of the dorsum, the absence of a developed anterior process on the left paramere, the complex structure of apex of phallotheca, and the broad body of the endosoma. All other species of Harpagophylus are pale in coloration, with differing structures on the apex of the phallotheca, and with the body of the endosoma not so robust.
Description
Male: Body moderately elongate, parallel sided; small, mean total length 2.51, mean width pronotum 0.91. COLORATION (pl. 1): Head: Uniformly dark, reddish; scapus unicolorous dark; pedicellus black at extreme base, remainder pale; labium generally infuscate, heavily so apically. Thorax: Pronotum and scutellum unicolorous red to brown; hemelytron partially pale with red or carmine areas, especially on clavus along scutellum and on endocorium; cuneus transparent, without markings; membrane and veins strongly fumose; hind femur nearly unicolorous dark; hind tibial spines dark, without dark spots at bases. Abdomen: Venter unicolorous dark. SURFACE AND VESTITURE: Dorsum with suberect or reclining black setae and some sericeous or woolly setae. STRUCTURE: Head: Somewhat projecting; interocular space relatively large; eyes leaving gena only very slightly exposed in lateral view. Antenna: Segment 2 of equal diameter over entire length; antennal fossa with ventral margin at ventral margin of eye. Labium: Reaching to about anterior margin of pygophore. GENITALIA (fig. 2): Phallotheca: Nearly erect; apex complex.
Female: Slightly more ovoid than male; small, mean total length 2.51, mean width pronotum 0.96. COLORATION (pl. 1): As in male. GENITALIA: Not examined.
Holotype
AUSTRALIA: Western Australia: Kalbarri National Park, Meanarra Hill, 27.69228°S 114.2104°E, 200 m, 29 Oct 1996, Schuh and Cassis, Calytrix variabilis Lindl. (Myrtaceae), det. Perth 05120470, 1♂ (AMNH_PBI 0134818) (WAMP).
Paratypes
AUSTRALIA: Western Australia: Kalbarri National Park, Meanarra Hill, 27.69228°S 114.2104°E, 200 m, 29 Oct 1996, Schuh and Cassis, Calytrix variabilis Lindl. (Myrtaceae), det. Perth 05120470, 2♂ (00134820, 00134821), 1♀ (00134826) (AM), 2♂ (00134822, 00134823), 2♀ (00134824, 00134825) (AMNH), 1♂ (00134819), 1♀ (00134827) (WAMP).
Diagnosis
Recognized by the pale, yellowish coloration, the endosomal processes both straight and erect, the dorsal process short, the ventral process much longer, the relatively short right paramere, and the triangular anterior process on the left paramere. Most easily confused with H. agnew, H. thryptomeni, and H. verticordii on the basis of the pale coloration, but those species differing from H. scholtzii in the structure and conformation of the dorsal and ventral endosomal processes.
Description
Male: Body weakly elongate, weakly ovoid; small, mean total length 2.17, mean width pronotum 0.75. COLORATION (pl. 1): Head: Uniformly pale yellow; scapus and pedicellus unicolorous pale; labium pale with segment 4 heavily infuscate. Thorax: Pronotum, scutellum, and hemelytron unicolorous pale yellow; markings on cuneus absent; membrane and veins weakly fumose; hind femur unicolorous pale, without black spots; hind tibial spines dark, without dark spots at bases. Abdomen: Venter unicolorous pale or mostly so. SURFACE AND VESTITURE: Dorsum with reclining simple setae matching background coloration and some sericeous or woolly setae. STRUCTURE: Head: Weakly projecting; interocular space relatively large; eyes leaving gena only very slightly exposed in lateral view. Antenna: Segment 2 of equal diameter over entire length; antennal fossa with ventral margin at ventral margin of eye. Labium: Reaching to about anterior margin of pygophore. GENITALIA (fig. 3): Phallotheca: Nearly erect; dorsal surface without a fingerlike projection; apex complex.
Female: Slightly more ovoid than male; small, mean total length 2.26, mean width pronotum 0.83. COLORATION (pl. 1): As in male. GENITALIA: Not examined.
Holotype
AUSTRALIA: Western Australia: 4.5 km NW of jct of Blowholes Rd and North West Coastal Hiway, N of Carnarvon, 24.72267°S 113.7158°E, 28 m, 27 Oct 2004, Cassis, Wall, Weirauch, Tatarnic, Symonds, Scholtzia leptantha Benth. (Myrtaceae), det. Perth6988687, 1♂ (AMNH_PBI 00327134) (WAMP).
Paratypes
AUSTRALIA: Western Australia: 4.5 km NW of jct of Blowholes Rd and North West Coastal Hiway, N of Carnarvon, 24.72267°S 113.7158°E, 28 m, 27 Oct 2004, Cassis, Wall, Weirauch, Tatarnic, Symonds, Scholtzia leptantha Benth. (Myrtaceae), det. Perth 6988687, 1♂ (00327135), 1♀ (00327140) (AM), 2♂ (00327136, 00327137), 2♀ (00327138, 00327139) (AMNH), 1♀ (00327141) (WAMP).
Diagnosis
Recognized by the pale yellowish coloration, and uniquely by the clawlike appendage on the left side of pygophoral opening (fig. 4) and the projecting hook on the apex of the phallotheca. Most easily confused with H. agnew, H. scholtzii, and H. verticordii on the basis of pale coloration, but those species all lacking the hook on the pygophoral opening and also showing differences in the structure of the anterior process of the left paramere, the conformation of the right paramere, and the conformation of the dorsal and ventral endosomal processes.
Description
Male: Body weakly elongate, weakly ovoid; small, mean total length 2.23, mean width pronotum 0.73. COLORATION (pl. 1): Head: Uniformly yellow-orange; scapus and pedicellus unicolorous pale; labium pale with segment 4 heavily infuscate. Thorax: Pronotum, scutellum, and hemelytron unicolorous yellow-orange; markings on cuneus absent; membrane and veins weakly fumose; hind femur unicolorous pale, without black spots; hind tibial spines dark, without dark spots at bases. Abdomen: Venter unicolorous pale or mostly so. SURFACE AND VESTITURE: Dorsum with reclining simple setae matching background coloration and some sericeous or woolly setae. STRUCTURE: Head: Weakly projecting; interocular space moderate; eyes leaving gena moderately exposed in lateral view. Antenna: Segment 2 of equal diameter over entire length; antennal fossa with ventral margin at ventral margin of eye. Labium: Just reaching onto abdomen. GENITALIA (fig. 4): Phallotheca: Nearly erect; dorsal surface without ornamentation; anterior and ventral surfaces without a keel; apex complex.
Female: Slightly more ovoid than male; small, mean total length 2.36, mean width pronotum 0.77. COLORATION (pl. 1): As in male. GENITALIA: As in generic description.
Distribution (map 1)
Known from one locality in the Goldfields-Esperance region in Western Australia.
Holotype
AUSTRALIA: Western Australia: 28 km S of Menzies (3.5 km E of Hiway), 29.91917°S 121.1514°E, 500 m, 25 Oct 1996, Schuh and Cassis, Thryptomene aspera aspera E. Pritz. (Myrtaceae), det. Perth 05095093, 1♂ (AMNH_PBI 00372759) (WAMP).
Paratypes
AUSTRALIA: Western Australia: 28 km S of Menzies (3.5 km E of Hiway), 29.91917°S 121.1514°E, 500 m, 25 Oct 1996, Schuh and Cassis, Thryptomene aspera aspera E.Pritz. (Myrtaceae), det. Perth 05095093, 1♂ (00373227), 8♀ (00373233–00373240) (AM), 3♂ (00372760–00372761, 00373228), 6♀ (00372762–00372766, 00373241) (AMNH), 4♀ (00373229–00373232) (WAMP).
Diagnosis
Recognized by the pale, yellowish, coloration, and the conformation of the endosomal processes, both long, curving, and recurved toward the base of the endosoma. Most easily confused with H. agnew, H. scholtzii, and H. thryptomeni on the basis of pale coloration, but H. verticordii differing from all of those species in the structure and conformation of the dorsal and ventral endosomal processes.
Description
Male: Body weakly elongate, weakly ovoid; small, mean total length 2.43, mean width pronotum 0.81. COLORATION (pl. 1): Head: Uniformly pale; scapus unicolorous pale; pedicellus unicolorous pale; labium pale with segment 4 heavily infuscate. Thorax: Pronotum unicolorous pale; scutellum unicolorous with remainder of dorsum; hemelytron unicolorous pale; markings on cuneus absent; membrane and veins weakly fumose; hind femur unicolorous pale, without black spots; hind tibial spines dark, without dark spots at bases. Abdomen: Venter unicolorous pale or mostly so. SURFACE AND VESTITURE (fig. 5B): Dorsum with reclining simple setae matching background coloration and some sericeous or woolly setae. STRUCTURE: Head (fig. 5A): Somewhat projecting; interocular space relatively large; eyes leaving gena moderately exposed in lateral view. Antenna: Segment 2 weakly tapering, more slender at base; antennal fossa with ventral margin 1 diameter above ventral margin of eye. Labium: Reaching to about anterior margin of pygophore. GENITALIA (figs. 5D–F, 6): Phallotheca: Nearly erect; dorsal surface without a fingerlike projection, and without a platelike projection; anterior and ventral surfaces without a keel; apex complex.
Female: Slightly more ovoid than male; small, mean total length 2.54, mean width pronotum 0.86. COLORATION (pl. 1): As in male. GENITALIA (fig. 6): As in generic description.
Hosts
Darwinia diosmoides (DC.) Benth., Verticordia forrestii F. Muell., Verticordia monadelpha Turcz (pl. 9A), V. polytricha Benth. (pl. 9B) (Myrtaceae).
Distribution (map 1)
Known from Western Australia, from Kalbarri National Park in the south to the Carnarvon region in the north.
Holotype
AUSTRALIA: Western Australia: Kalbarri National Park, Loop Road, 27.56163°S 114.4376°E, 300 m, 28 Oct 1996, Schuh and Cassis, Verticordia polytricha Benth. (Myrtaceae), det. Perth 05120594, 1♂ (AMNH_PBI 00370995) (WAMP).
Paratypes
AUSTRALIA: Western Australia: Kalbarri National Park, Loop Road, 27.56163°S 114.4376°E, 300 m, 28 Oct 1996, Schuh and Cassis, Verticordia monadelpha Turcz. (Myrtaceae), det. PERTH staff, 6♂ (00087221, 00373196–00373200), 5♀ (00373192, 00373205–00373208) Verticordia polytricha Benth. (Myrtaceae), det. Perth 05120594, 9♂ (00373176–00373181, 00373193–00373195), 5♀ (00373187–00373191) (AM), Verticordia monadelpha Turcz. (Myrtaceae), det. PERTH staff, 5♂ (00087511, 00087513, 00370998–00370999, 00373186), 5♀ (00371000–00371002, 00373203–00373204) Verticordia polytricha Benth. (Myrtaceae), det. Perth 05120594, 2♂ (00370996, 00370997) (AMNH), Verticordia monadelpha Turcz. (Myrtaceae), det. PERTH staff, 1♂ (00373201) Verticordia polytricha Benth. (Myrtaceae), det. Perth 05120594, 1♀ (00373209) (UCR), Verticordia monadelpha Turcz. (Myrtaceae), det. PERTH staff, 1♂ (00373202) Verticordia polytricha Benth. (Myrtaceae), det. Perth 05120594, 1♀ (00373210) (UNSW), Verticordia monadelpha Turcz. (Myrtaceae), det. PERTH staff, 1♂ (00373185) Verticordia polytricha Benth. (Myrtaceae), det. Perth 05120594, 3♂ (00373182–00373184), 4♀ (00373211–00373214) (WAMP). North West Coast Hiway 72 km NE of jct with Blowholes Rd, 24.18336°S 114.0381°E, 34 m, 28 Oct 2004, Cassis, Wall, Weirauch, Tatarnic, Symonds, Verticordia forrestii F.Muell. (Myrtaceae), det. Perth6989004, 1♂ (00373215), 2♀ (00373216, 00373217) (AM), Verticordia forrestii F. Muell. (Myrtaceae), det. Perth6989004, 2♀ (00322681, 00322682) (AMNH).
Other Specimens Examined
AUSTRALIA: Western Australia: Kalbarri National Park, Loop Road, 27.56163°S 114.4376°E, 300 m, 28 Oct 1996, Schuh and Cassis, Darwinia diosmoides (DC.) Benth. (Myrtaceae), det. Perth 05120551, 5♂ (00373218–00373222), 3♀ (00373223–00373225) (AM).
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 = left paramere; pe = pulvillus; rp = right paramere (AMNH_PBI 00135313).
Diagnosis
Recognized within the Melaleucoides genus group by the small size, the apparent absence of a secondary endosomal strap (fig. 7; 31-3), the unique conformation of the left paramere, with a greatly elongate, medial, apical process (fig. 7; 56-2), and the female vestibulum with large, nearly symmetrical medial plates (fig. 7; 74-1). Also recognized within the Melaleucoides genus group by possessing only the ventral endosomal process. Easily confused with Harpagophylus calytrix, Melaleucoides akaina, and M. pileanthicola on the basis of small size and reddish markings on a pale background. Easily distinguished from Melaleucoides akiana by the absence of the ventral endosomal process in that species, from Harpagophylus spp. by their possession of both ventral and dorsal endosomal processes, and from all of those taxa by the presence of a secondary endosomal strap and the distinctive structure of the left paramere in Thryptomenomiris.
Description
Male: Body moderately elongate, parallel sided; small, mean total length 2.44–2.79, mean width pronotum 0.77–0.92. COLORATION (pl. 1): Pale with some brownish or reddish areas. SURFACE AND VESTITURE (fig. 8C): Dorsum with suberect or reclining black setae and some sericeous or woolly setae. STRUCTURE: Head (fig. 8A): Somewhat projecting; interocular space moderate; eyes leaving gena moderately exposed in lateral view. Antenna: Segment 2 weakly tapering, more slender at base; antennal fossa with ventral margin 1 diameter above ventral margin of eye. Labium: Just reaching onto abdomen. Thorax: Pretarsus (fig. 8D): Parempodia fleshy, recurved, lyriform; pulvilli absent. GENITALIA (figs. 7, 8E, F, 9): Endosoma: Base short, curving, C- or J-shaped; body without torsion, primary strap ventral to secondary gonopore, apically greatly elongate, without ornamentation; spinelike, elongate process near gonopore on ventral surface present, recurved toward base of endosoma; secondary 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 limited extent of lateral surface of pygophore; body more or less quadrate; apex medially broadly rounded to weakly quadrate; apex posterolaterally with a greatly elongate projection (figs. 7, 8F, 9); anterior process slender and cylindrical in lateral view; posterior process in the form of a short, straight, fingerlike projection; base of posterior process 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 (fig. 8B). GENITALIA (fig. 7): Posterior wall laterally with distinct crescent-shaped interramal sclerites; posterolaterally with a distinct swelling covered with microtrichia; longitudinal fold on either side of midline along part of length; interramal lobes roughly symmetrical, erect, and not ornamented with spicules or with a very few; vestibulum with medial plates sclerotized and readily visible, large, nearly symmetrical, with sclerotized guide structure present as caplike structure with internal differentiation.
Etymology
Named for the host genus, Thryptomene (Myrtaceae), in combination with Miris; gender masculine.
Diagnosis
In addition to the characteristics in the generic diagnosis, recognized by the attenuated apex of the apical spine of the left paramere (fig. 7), in contrast to the blunt spine in T. yalgoo (fig. 9).
Description
Male: Body weakly to distinctly ovoid; small, mean total length 2.48, mean width pronotum 0.82. COLORATION (pl. 1): Head: Uniformly pale; scapus and pedicellus unicolorous pale; labium pale with segment 4 heavily infuscate. Thorax: Pronotum unicolorous pale; scutellum unicolorous with remainder of dorsum; hemelytron pale with endocorium somewhat darker than remainder; markings on cuneus absent, cuneus transparent; membrane and veins weakly fumose; hind femur unicolorous pale, without black spots; hind tibial spines dark, without dark spots at bases. Abdomen: Venter unicolorous pale or mostly so. SURFACE AND VESTITURE: Dorsum with suberect or reclining black setae and some sericeous or woolly setae. STRUCTURE: Head: Somewhat projecting; interocular space moderate; eyes leaving gena moderately exposed in lateral view. Antenna: Segment 2 weakly tapering, more slender at base; antennal fossa with ventral margin 1 diameter above ventral margin of eye. Labium: Just reaching onto abdomen. GENITALIA (fig. 7): Phallotheca: Smoothly curving on dorsal margin; dorsal surface without a fingerlike or platelike projection; anterior surface without a keel; ventral surface with a projecting keel; apex simple.
Female: Slightly more ovoid than male; small, mean total length 2.66, mean width pronotum 0.84. COLORATION (pl. 1): As in male. GENITALIA (fig. 7): See generic description.
Etymology
Named for the region of the type locality, Kalbarri, Western Australia; a noun in apposition.
Hosts
Thryptomene sp. (Myrtaceae). The record from Malleostemon hursthousei may well represent a genuine host, but would benefit from additional documentation. The record from Solanum probably results from specimens being commingled in the field and does not represent a breeding host.
Holotype
AUSTRALIA: Western Australia: Kalbarri National Park, Loop Road, 27.56163°S 114.4376°E, 300 m, 28 Oct 1996, Schuh and Cassis, Thryptomene sp. (Myrtaceae), det. Perth 05120543, 1♂ (AMNH_PBI 00372768) (WAMP).
Paratypes
AUSTRALIA: Western Australia: Kalbarri National Park, Loop Road, 27.56163°S 114.4376°E, 300 m, 28 Oct 1996, Schuh and Cassis, Thryptomene sp. (Myrtaceae), det. Perth 05120543, 5♂ (00373256–00373257, 00373275–00373276, 00373284), 17♀ (00373258–00373265, 00373277–00373283, 00373287, 00373292) (AM), 4♂ (00087217, 00372769–00372770, 00373291), 7♀ (00087218, 00372772–00372774, 00373272–00373274) (AMNH), 1♂ (00373285), 2♀ (00373288, 00373289) (UCR), 2♂ (00373255, 00373290), 1♀ (00373286) (UNSW), 6♀ (00373266–00373271) (WAMP).
Other Specimens Examined
AUSTRALIA: Western Australia: 25 km S of Gascoyne Junction, on Towrana Homestead Rd, 25.39014°S 115.1506°E, 232 m, 04 Nov 2004, Cassis, Weirauch, Tatarnic, Symonds, Solanum sp. (Solanaceae), det. Field ID, 1♂ (00373293), 1♀ (00373294) (AM). ca. 1 km S of Murchison House HS, Kalbarri National Park, 27.65822°S 114.2394°E, 60 m, 23 Oct 2004, Cassis, Wall, Weirauch, Symonds, Malleostemon hursthousei (W.Fitzg.) J.W.Green (Myrtaceae), det. Perth6988423, 1♂ (00373295) (AM).
Diagnosis
In addition to the characteristics in the generic diagnosis, recognized by the blunt apex of the apical spine of the left paramere (fig. 9), in contrast to the attenuated spine in T. kalbarri (fig. 7).
Description
Male: Body weakly elongate, weakly ovoid; small, mean total length 2.65, mean width pronotum 0.88. COLORATION (pl. 1): Head: Uniformly pale; scapus and pedicellus unicolorous pale; labium pale with segment 4 heavily infuscate. Thorax: Pronotum unicolorous pale; scutellum entirely red or orange; hemelytron with red or carmine spots, botches, or solid areas; markings on cuneus absent; membrane and veins weakly fumose; hind femur unicolorous pale, without black spots; hind tibial spines dark, without dark spots at bases. Abdomen: Venter unicolorous pale or mostly so. SURFACE AND VESTITURE (fig. 8C): Dorsum with reclining simple setae matching background coloration and some sericeous or woolly setae. STRUCTURE: Head (fig. 8A): Somewhat projecting; interocular space relatively large; eyes leaving gena moderately exposed in lateral view. Antenna: Segment 2 weakly tapering, more slender at base; antennal fossa with ventral margin at ventral margin of eye. Labium: Reaching to about anterior margin of pygophore. Thorax: Pretarsus (fig. 8D): As in generic description. GENITALIA (figs. 8E, F, 9). Phallotheca: Smoothly curving on dorsal margin; dorsal surface without a fingerlike or platelike projection; anterior surface with a short to elongate keel; ventral surface with a projecting keel; apex simple.
Female: Slightly more ovoid than male; small, mean total length 2.77, mean width pronotum 0.90. COLORATION (pl. 1): As in male. STRUCTURE: Head (fig. 8B). GENITALIA: Not examined.
Etymology
Named for the small town of Yalgoo in the region of the type locality, Western Australia; a noun in apposition.
Holotype
AUSTRALIA: Western Australia: 46.5 km W of Yalgoo, 28.41302°S 116.2151°E, 600 m, 27 Oct 1996, Schuh and Cassis, Thryptomene aspera glabra E. Pritz. (Myrtaceae), det. Perth 05120683, 1♂ (AMNH_PBI 00135307) (WAMP).
Paratypes
AUSTRALIA: Western Australia: 24 km W of Sandstone, 28.01426°S 119.0474°E, 650 m, 26 Oct 1996, Schuh and Cassis, Thryptomene aspera glabra E. Pritz. (Myrtaceae), det. Perth 05095182, 1♂ (00372767), 3♀ (00087220, 00373297–00373298) (AM), 1♂ (00373296) (AMNH). 46.5 km W of Yalgoo, 28.41302°S 116.2151°E, 600 m, 27 Oct 1996, Schuh and Cassis, Thryptomene aspera glabra E. Pritz. (Myrtaceae), det. Perth 05120683, 2♂ (00135304, 00135312), 1♀ (00135317) (AM), 9♂ (00135305–00135306, 00135308–00135311, 00135314–00135316), 10♀ (00135318–00135320, 00135323, 00135325–00135330) (AMNH), 2♀ (00135321, 00135322) (WAMP).
Maps 3 and 4
Localities for species of Melaleucoides: M. akaina–M. castanea (top; map 3) and M. grossi–M. rhapiophyllae (bottom; map 4).
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 = evaporatory area; mssp = mesothoracic spiracle; pe = parempodium (AMNH_PBI 00137503).
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 = left paramere; pe = parempodium; rp = right paramere (AMNH_PBI 00129566).
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 = left paramere; pe = parempodium; pv = pulvillus (AMNH_PBI 00371072).
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 = dorsal arolium; lp = left paramere; pe = parempodium; ph = phallotheca; pv = pulvillus; rp = right paramere (AMNH_PBI 00371141).
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 = evaporatory area; lp = left paramere; mssp = mesothoracic spiracle; pe = parempodium; ph = phallotheca; rp = right paramere (AMNH_PBI 00137348).
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 = left paramere; pe = parempodium; pv = pulvillus; rp = right paramere (AMNH_PBI 00368327).
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 = dorsal arolium; lp = left paramere; pe = parempodium; ph = phallotheca; rp = right paramere (AMNH_PBI 00129613).
Fig. 35.
Phylogenetic relationships within the Melaleucoides genus group based on equal weight analysis using TNT and maximum parsimony. L = 506, CI = 35, RI = 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 = 35.982.
Fig. 37.
Host relationships of Melaleucoides group plotted on implied-weights phylogenetic analysis in figure 36.
Diagnosis
Recognized within the Melaleucoides genus group by the unique structure of the left paramere, with its flattened, spoon-shaped form and erect position, covering the dorsal margin of the pygophore in lateral view (fig. 12D; 53-1). Most easily confused with Harpagophylus and Thryptomenomiris on the basis of unique parempodial structure possessed by all three genera, but easily separated from both on the structure of the left paramere.
Description
Male: Body weakly elongate, weakly ovoid; mean total length 2.33–4.05, mean width pronotum 0.78–1.55. COLORATION (pls. 1–Plate 2Plate 34): Variable, ranging from completely castaneous to almost completely pale, and sometimes with spots. SURFACE AND VESTITURE (figs. 12B, 17B): Dorsum with suberect or reclining black setae, without sericeous or woolly setae. STRUCTURE: Head (figs. 12A, 14A, 17A, 33A): Barely projecting; interocular space moderate; eyes leaving gena moderately exposed in lateral view. Antenna: Segment 2 weakly tapering, more slender at base; antennal fossa with ventral margin at ventral margin of eye. Labium: Just reaching onto abdomen. Thorax: Pretarsus (figs. 12C, 17C, 33D): Parempodia fleshy, recurved, lyriform; pulvilli present, flaplike, covering about one third of ventral claw surface. GENITALIA (e.g., figs. 11, 12E, F, 13, 18, 33F, G, 34): Endosoma: Base short; curving, C- or J-shaped; body without torsion; primary strap ventral to secondary gonopore, apically greatly elongate, straight, weakly curving, or angulate relative to body of endosoma, without ornamentation; secondary endosomal strap fused with primary strap proximal to secondary gonopore, usually reaching well beyond gonopore, often fused with primary strap, sometimes with denticles or comblike ornamentation; sometimes with elongate spinelike process arising near gonopore on dorsal surface; sometimes with spinelike elongate process near gonopore on ventral surface, ornamented with denticles or not; sometimes with bladderlike process distad of secondary gonopore; secondary gonopore sclerite absent; secondary gonopore usually seen frontally (facing up) in lateral view of endosoma. Phallotheca: Smoothly curving on dorsal margin, without a fingerlike projection, and with a curving, posteriorly directed, platelike projection; anterior surface without a keel; ventral surface without a projecting keel; posterior surface lacking transparent window; apex simple. Left Paramere: Vertical; shaft at right angles to or in same axis as body; in dorsal perspective (fig. 12F) not exceeding margin of pygophore; in lateral perspective (fig. 12E) extending beyond dorsal margin of pygophore; body quadrate or spoon shaped; apex medially broadly rounded to weakly quadrate, sometimes elongated and or recurved; apex posterolaterally simple or in the form of a prolongation; anterior process triangular or elongate and cylindrical in lateral view; posterior process in the form of a short to long, fingerlike projection; base of posterior process with or without conspicuous shoulder. Right Paramere: Body short and broad, lanceolate; apex short, clawlike, decurved.
Female: Slightly more ovoid than male; small, mean total length 2.52–4.26, mean width pronotum 0.82–1.59. COLORATION (pls. 1–4): As in male. GENITALIA (figs. 11, 18, 25, 34): Posterior wall laterally with distinct crescent-shaped interramal sclerites; posterolaterally with a distinct swelling covered with microtrichia; longitudinal fold on either side of midline along part of length; interramal lobes present and asymmetrical, reclining, overlapping, and heavily ornamented with spicules; vestibulum with medial plates sclerotized and readily observed; small, nearly symmetrical, triangular; with sclerotized guide structure present as caplike structure with internal differentiation.
Etymology
Named after the host genus, Melaleuca (Myrtaceae), in combination with the Latin suffix -oides, a diminutive; gender feminine.
Key to species of Melaleucoides
1. Dorsum largely castaneous, including entire clavus (pls. 2–Plate 34)2
– Dorsum not so heavily castaneous, at least clavus and anterior half of endocorium pale, red, or with dark spots on pale background4
.2. Dorsum uniformly castaneous, including cuneus and apex of scutellum (pl. 2); male genitalia as in figure 19Melaleucoides castanea
– At least clavus and apex of scutellum pale3
3. Male genitalia as in figure 26, phallotheca with a short fingerlike projection on dorsal margin; known from Melaleuca rhaphiophyllaMelaleucoides rhaphiophyllae
– Male genitalia as in figure 31, phallotheca with a long fingerlike projection on dorsal margin; known from Melaleuca uncinataMelaleucoides uncinatae
4. Dorsum with conspicuous small dark spots and a pale background5
– Dorsum without small dark spots on a pale background8
5. All spots on dorsum small (pl. 2); apex of scutellum dark; male genitalia as in figure 16; known from Melaleuca brevifoliaMelaleucoides brevifoliae
– Spots on dorsum larger (pls. 2, 3); apex of scutellum pale as remainder of scutellum6
6. Spots round, never coalescent (pls. 2, 3)7
– Many spots of irregular shape, often coalescent, particularly on hemelytron (pl. 2); male genitalia as in figure 15; known from Beaufortia micranthaMelaleucoides beaufortiae
7. Male genitalia as in figure 21; known from Melaleuca leuropomaeMelaleucoides leuropomae
– Male genitalia as in figure 22; known from Melaleuca micranthaMelaleucoides micranthae
8. Hemelytron with conspicuous longitudinal red areas9
– Hemelytron without conspicuous longitudinal red areas13
9. Calli castaneous, much darker than surrounding pronotum and strongly contrasting with it10
– Calli not castaneous, although sometimes weakly red, not strongly contrasting with surrounding pronotum11
10. Red markings on hemelytron broad, including most of clavus and endocorium, but not including claval and radiomedial veins (pl. 1); male genitalia as in figure 13, known from Melaleuca sheathianaMelaleucoides annae
– Red markings on hemelytron narrow, running along claval and radiomedial veins (pl. 4); male genitalia as in figure 32, known from Melaleuca undulataMelaleucoides undulatae
11. Cuneus almost entirely pale; broad marking on endocorium intensely red (pls. 1, 3)12
– Apical half of cuneus red, in contrast with remainder (pl. 3); male genitalia as in figure 25; known from Thryptomene kochiiMelaleucoides ozzii
12. Male genitalia as in figure 25, endosoma with two distinct, elongate, apical spines and a ventral process adjacent to the secondary gonopore; known from Pileanthus spp.Melaleucoides pileanthicola
– Male genitalia as in figure 11, endosoma with a single, short, apical spine and no processes associated with the secondary gonopore; known from Micromyrtus hursthousei and Scholtzia leptanthaMelaleucoides akaina
13. Coloration of dorsum nearly uniform, either faded green, greenish yellow, or yellow14
– Coloration of dorsum not uniform, posterior half of endocorium and scutellum dark, in contrast with much of remainder of hemelytron and pronotum (pl. 4); male genitalia as in figure 29; known from Melaleuca systenaMelaleucoides systenae
14. Large, robust species, total length at least 3.50, width pronotum at least 1.31; coloration of dorsum faded green or yellow green (pls. 2, 3)15
– More elongate species, total length no more than 3.76, width pronotum no more than 1.28; coloration of dorsum bright yellow orange (pls. 3, 4)16
15. Scutellum white or nearly so (pl. 2); male genitalia as in figure 18; known from Melaleuca uncinataMelaleucoides cassisi
– Scutellum orange (pl. 3); male genitalia as in figure 18; known from Melaleuca sheathianaMelaleucoides sheathianae
16. Male genitalia as in figure 28; known from multiple genera of MyrtaceaeMelaleucoides similis
– Male genitalia as in figure 34; known from Verticordia spp. and Pileanthus spp.,Melaleucoides verticoridiae
Melaleucoides akaina, new species
Figures 10, 11; map 3; plate 1
Diagnosis
Unequivocally recognized among Melaleucoides species by the uniquely short, stout endosoma with a short apical projection, no spinelike processes associated with the region of the secondary gonopore, and a row of denticles on the margin of the secondary gonopore, as well as the long, recurved, tapered, apical spine on the apex of the left paramere. Easily confused with Harpagophylus calytrix, Thryptomenomiris spp., and Melaleucoides pileanthicola on the basis its small size and reddish coloration on a pale background. All of those taxa with ventral/and or dorsal processes on the endosoma and with a long apical endosomal spine.
Description
Male: Body moderately elongate, parallel sided; small, mean total length 2.36, mean width pronotum 0.79. COLORATION (pl. 1): Head: Uniformly pale; scapus and pedicellus yellow to dirty yellow; labium pale with segment 4 heavily infuscate. Thorax: Pronotum unicolorous pale; scutellum unicolorous with remainder of dorsum; hemelytron mostly pale, endocorium somewhat darker than remainder; markings on cuneus absent; membrane, including veins, pale; hind femur unicolorous pale, without black spots; hind tibial spines dark, without dark spots at bases. Abdomen: Venter unicolorous pale or mostly so. SURFACE AND VESTITURE (fig. 10C): Vestiture of dorsum consisting of simple, recumbent setae with coloration of dorsum. STRUCTURE: Head: Somewhat projecting; interocular space relatively large; eyes leaving gena only very slightly exposed in lateral view. Antenna: Segment 2 weakly tapering, more slender at base; antennal fossa with ventral margin at ventral margin of eye. Labium: Reaching to about anterior margin of pygophore. Thorax: Pretarsus (fig. 10D): Parempodia fleshy, recurved, lyriform; pulvilli absent. GENITALIA (fig. 11): Endosoma: Base short, curving, C- or J-shaped; body with torsion, primary strap dorsal to secondary gonopore, apically short, ratio length apex strap/length secondary gonopore 1.33–1.50, at least weakly arcuate and curving dorsally, no ornamentation; secondary endosomal strap fused with primary strap, reaching just beyond secondary gonopore as fingerlike extension with one tip, and devoid of ornamentation; secondary gonopore seen laterally in lateral view of endosoma, with a denticulate ridge on right side of endosoma. Phallotheca: Smoothly curving on dorsal margin; apex simple. Left Paramere: Vertical; shaft at right angles to body; in dorsal perspective not exceeding margin of pygophore; in lateral perspective covering limited extent of lateral surface of pygophore; body spoon shaped; apex medially with an elongate, acuminate, recurved, medial spine; anterior process triangular in lateral view; posterior process in the form of a long, straight, slender, fingerlike projection; base of posterior process without conspicuous shoulder. Right Paramere: Body short and broad, lanceolate; apex short, clawlike, decurved.
Female: Slightly more ovoid than male; mean total length 2.61, mean width pronotum 0.85. COLORATION (pl. 1): Paler than male. STRUCTURE: Head (fig. 10A). GENITALIA (fig. 11): Posterior wall laterally with distinct crescent-shaped interramal sclerites; posteriorly without a sclerotized transverse band; posterolaterally with a distinct swelling covered with microtrichia; longitudinal fold on either side of midline along part of length; posterior margin of posterior wall without spicules, and not reflexed dorsally; interramal lobes roughly symmetrical, erect, and heavily ornamented with spicules; vestibulum with medial plates sclerotized and readily observed, large, nearly symmetrical, with sclerotized guide structure present as short heavily sclerotized tube on left side.
Distribution (map 3)
Known from two localities north of Kalbarri National Park and the Carnarvon region of Western Australia.
Discussion
Melaleucoides akaina is unique among members of the genus in having the endosoma with no processes associated with the region of the secondary gonopore. Nonetheless, other aspects of morphology, especially the novel structure of the parempodia and left paramere agree closely with the remaining species we place in Melaleucoides. On the basis of our phylogenetic analyses, we conclude that the relatively simple endosoma in M. akaina is the result of reduction.
Holotype
AUSTRALIA: Western Australia: NW Coastal Hiway 58 km N of Kalbarri Road, 27.43701°S 114.6768°E, 500 m, 30 Oct 1996, Schuh and Cassis, Micromyrtus hursthousei W. Fitzgerald (Myrtaceae), det. Perth 05120241, 1♂ (AMNH_PBI 00372340) (WAMP).
Paratypes
AUSTRALIA: Western Australia: NW Coastal Hiway 58 km N of Kalbarri Road, 27.43701°S 114.6768°E, 500 m, 30 Oct 1996, Schuh and Cassis, Micromyrtus hursthousei W. Fitzgerald (Myrtaceae), det. Perth 05120241, 4♂ (00372342, 00372343, 00087543, 00087346), 3♀ (00372345–00087347) (AM), 1♂ (00372344), 1♀ (00372347) (AMNH), 1♂ (00372341), 1♀ (00372348) (WAMP).
Other Specimens Examined
AUSTRALIA: Western Australia: 4.5 km NW of jct of Blowholes Rd and North West Coastal Hiway, N of Carnarvon, 24.72267°S 113.7158°E, 28 m, 27 Oct 2004, Cassis, Wall, Weirauch, Tatarnic, Symonds, Scholtzia leptantha Benth. (Myrtaceae), det. Perth6988687, 3♂ (00373164–00373166), 3♀ (00373170–00373172) (AM), 3♂ (00373167–00373169), 2♀ (00373173, 00373174) (AMNH).
Melaleucoides annae, new species
Figures 12, 13; map 3; plate 1
Diagnosis
Recognized by the castaneous calli, the elongate red markings on the hemelytron (pl. 1), and the endosoma with denticles on the secondary strap distad of the secondary gonopore and a recurved ventral process associated with the secondary gonopore (fig. 13). Most easily confused with M. undulatae on the basis of coloration, but the red markings on the hemelytron more strongly linear in that species (pl. 4) and the lack of denticles distad of the secondary gonopore, a dorsal rather than a ventral process, and a bladderlike process just distad of the gonopore (fig. 32).
Description
Male: Body weakly elongate, weakly ovoid; mean total length 3.46, mean width pronotum 0.1.29. COLORATION (pl. 1): Head: Uniformly pale; scapus yellow to dirty yellow; pedicellus dirty yellow proximally, weakly to heavily infuscate distally; labium generally infuscate, heavily so apically. Thorax: Pronotum pale with castaneous calli and sometimes humeri; scutellum white; hemelytron with red or carmine areas along mesial margin of clavus and on endocorium; markings on cuneus present as a contrasting white basal fascia, remainder red; membrane weakly fumose, veins pale; hind femur with a few brown or black spots; hind tibial spines black with very small dark bases. Abdomen: Venter unicolorous pale or mostly so. SURFACE AND VESTITURE (fig. 12B): Dorsum with suberect or reclining black setae and some sericeous or woolly setae. STRUCTURE: Head (fig. 12A): Barely projecting; interocular space moderate; eyes leaving gena moderately exposed in lateral view. Antenna: Segment 2 weakly tapering, more slender at base; antennal fossa with ventral margin at ventral margin of eye. Labium: Just reaching onto abdomen. Thorax: Pretarsus (fig. 12C): Pulvilli present, flaplike, covering about one third of ventral claw surface. GENITALIA (figs. 12E, F, 13): Endosoma: Primary strap apically greatly elongate, angled dorsally relative to body of endosoma, no ornamentation; secondary endosomal strap fused with primary strap proximal to secondary gonopore, reaching well beyond gonopore, fused with primary strap, ornamented with some short denticles; spinelike, elongate process arising near gonopore on dorsal surface absent; spinelike, elongate process near gonopore on ventral surface present, strongly recurved toward base of endosoma; bladderlike process distad of secondary gonopore absent; secondary gonopore seen frontally (facing up) in lateral view of endosoma. Phallotheca: Nearly erect; dorsal surface with a curving, posteriorly directed, platelike projection; anterior surface with a short to elongate keel; apex simple. Left Paramere: Shaft at right angles to body; body spoon shaped; apex medially broadly rounded; anterior process triangular in lateral view; posterior process in the form of a short, straight, fingerlike projection; base of posterior process without conspicuous shoulder. Right Paramere: Body short and broad, lanceolate; apex short, clawlike, decurved.
Female: Slightly more ovoid than male; mean total length 3.51, mean width pronotum 0.1.36. COLORATION (pl. 1): As in male. GENITALIA: Posterior wall laterally with distinct crescent-shaped interramal sclerites; posteriorly without a sclerotized transverse band; posterolaterally with a distinct swelling covered with microtrichia; longitudinal fold on either side of midline along part of length; posterior margin of posterior wall without spicules and not reflexed dorsally; interramal lobes present and asymmetrical, reclining, overlapping, and heavily ornamented with spicules; vestibulum with medial plates sclerotized and readily observed, small, nearly symmetrical, triangular; with sclerotized guide present as caplike structure with internal differentiation.
Etymology
Named for Anna Massie, with thanks for her encouragement and interest in our work on the Australian Miridae fauna.
Holotype
AUSTRALIA: Western Australia: 11 km N of Coolgardie-Esperance Hiway on Kambalda Road, 31.25231°S 121.5899°E, 320 m, 18 Nov 1999, R.T. Schuh, G. Cassis, & R. Silveira, Melaleuca sheathiana W. Fitzg. (Myrtaceae), det. Perth 05671396, 1♂ (AMNH_PBI 00129550) (WAMP).
Paratypes
AUSTRALIA: Western Australia: 11 km N of Coolgardie-Esperance Hiway on Kambalda Road, 31.25231°S 121.5899°E, 320 m, 18 Nov 1999, R.T. Schuh, G. Cassis, & R. Silveira, Melaleuca sheathiana W. Fitzg. (Myrtaceae), det. Perth 05671396, 23♂ (00371206, 00371250–00371261, 00371265, 00371288–00371296), 35♀ (00371266–00371270, 00371272–00371277, 00371280–00371285, 00371287, 00371297–00371312, 00372109), 1 nymph (00371313) (AM), 31♂ (00087299, 00087498, 00129544–00129547, 00129551–00129569, 00129595–00129600), 17♀ (00087300, 00129548, 00129572–00129579, 00129582–00129583, 00129601–00129604, 00371278) (AMNH), 3♂ (00372092–00372094), 2♀ (00372113, 00372114) (ANIC), 1♂ (00371264), 1♀ (00371286) (CNC), 2♂ (00129570, 00129571), 2♀ (00129580, 00129581) (UCR), 10♂ (00372082–00372091), 4♀ (00372095, 00372110–00372112) (UNSW), 1♂ (00371262), 1♀ (00371271) (USNM), 9♂ (00129531–00129533, 00371207–00371212), 16♀ (00129534–00129543, 00371223–00371228) (WAMP), 1♂ (00371263), 1♀ (00371279) (ZISP).
Other Specimens Examined
AUSTRALIA: Western Australia: 11 km N of Coolgardie-Esperance Hiway on Kambalda Road, 31.25231°S 121.5899°E, 320 m, 18 Nov 1999, R.T. Schuh, G. Cassis, & R. Silveira, Melaleuca sheathiana W.Fitzg. (Myrtaceae), det. Perth 05671396, 14♂ (00371192–00371205), 10♀ (00371213–00371222) (AM).
Melaleucoides beaufortiae, new species
Figures 14, 15; map 3; plate 2
Diagnosis
Unique among Melaleucoides spp. for the coalescent small dark spots on the dorsum (pl. 2), the compact body form, and the structure of the male genitalia, the left paramere with a moderately elongate, blunt, fingerlike, recurved, medial apical process, and the endosoma with some denticles on the postgonoporal region of the secondary strap but lacking both the dorsal and ventral processes frequently seen in the genus (fig. 15). Most easily confused with M. brevifoliae, M. leuropomae, and M. micranthae, but the spots never coalescent in those species and the apical medial process of the left paramere much less elongate and not recurved (figs. 16, 21, 22, respectively).
Description
Male: Body weakly elongate, weakly ovoid; mean total length 2.98, mean width pronotum 1.10. COLORATION (pl. 2): Head: Pale, clypeus dark and large, dark, linear, bilateral markings on frons; scapus pale with a dark base; pedicellus dirty yellow proximally, weakly to heavily infuscate distally; labium generally infuscate, heavily so apically. Thorax: Pronotum pale with brown spots of varying size; scutellum pale with brown spots; hemelytron unicolorous pale with brown spots of varying size, some coalescent; markings on cuneus present as a red to castaneous apex; membrane weakly fumose, veins pale; hind femur with many brown or black spots; hind tibial spines dark with conspicuous dark spots at bases. Abdomen: Venter unicolorous pale or mostly so. SURFACE AND VESTITURE: Dorsum with suberect or reclining black setae, without sericeous or woolly setae. STRUCTURE: Head (fig. 14A): Somewhat projecting; interocular space moderate; eyes leaving gena broadly exposed in lateral view. Antenna: Segment 2 weakly tapering, more slender at base; antennal fossa with dorsal margin somewhat below ventral margin of eye. Labium: Reaching to about anterior margin of pygophore. Thorax: Pretarsus (fig. 14B): Pulvilli present, flaplike, covering about one third of ventral claw surface. GENITALIA (fig. 14C, D, 15): Endosoma: Primary strap apically greatly elongate, curving dorsally near apex, without ornamentation; secondary endosomal strap fused with primary strap proximal to secondary gonopore, reaching well beyond gonopore, ornamented with short denticles just distad of gonopore; spinelike, elongate process arising near gonopore on dorsal surface absent; spinelike, elongate process near gonopore on ventral surface absent; bladderlike process distad of secondary gonopore absent; secondary gonopore seen frontally (facing up) in lateral view of endosoma. Phallotheca: Nearly erect; apex simple. Left Paramere: Shaft at right angles to body; body spoon shaped; apex medially drawn into a blunt, recurved, fingerlike process; anterior process triangular in lateral view; posterior process in the form of a short curved fingerlike projection; base of posterior process without conspicuous shoulder. Right Paramere: Body short and broad, lanceolate; apex short, clawlike, decurved.
Female: Slightly more ovoid than male; mean total length 3.06, mean width pronotum 1.13. COLORATION (pl. 2): As in male. GENITALIA: Posterior wall laterally with distinct crescent-shaped interramal sclerites; posteriorly without a sclerotized transverse band; posterolaterally with a distinct swelling covered with microtrichia; longitudinal fold on either side of midline along part of length; posterior margin of posterior wall without spicules, and not reflexed dorsally; interramal lobes present and asymmetrical, reclining, overlapping, and heavily ornamented with spicules; vestibulum with medial plates sclerotized and readily observed, small, nearly symmetrical, triangular; with sclerotized guide present as short heavily sclerotized tube on left side.
Distribution (map 3)
Known from one locality in Fitzgerald River National Park in Southwestern Australia.
Holotype
AUSTRALIA: Western Australia: Fitzgerald River National Park, Hammersley Road, 33.81038°S 119.787°E, 215 m, 06 Dec 1997, Schuh, Cassis, Brailovsky, Asquith, Beaufortia micrantha Schauer (Myrtaceae), det. Perth 05055482, 1♂ (AMNH_PBI 00371069) (WAMP).
Paratypes
AUSTRALIA: Western Australia: Fitzgerald River National Park, Hammersley Road, 33.81038°S 119.787°E, 215 m, 06 Dec 1997, Schuh, Cassis, Brailovsky, Asquith, Beaufortia micrantha Schauer (Myrtaceae), det. Perth 05055482, 10♂ (00371063, 00371065–00371068, 00371072–00371075, 00371077), 5♀ (00371082, 00371084–00371085, 00371087, 00371095) (AM), 7♂ (00087473, 00371064, 00371070–00371071, 00371076, 00371078–00371079), 6♀ (00371083, 00371086, 00371088–00371089, 00371094, 00371097) (AMNH), 1♂ (00371080), 1♀ (00371098) (UNSW), 3♂ (00087247–00087248, 00371081), 4♀ (00371090, 00371092–00371093, 00371096) (WAMP).
Other Specimens Examined
AUSTRALIA: Western Australia: Fitzgerald River National Park, Hammersley Road, 33.81038°S 119.787°E, 215 m, 06 Dec 1997, Schuh, Cassis, Brailovsky, Asquith, Beaufortia micrantha Schauer (Myrtaceae), det. Perth 05055482, 1 nymph (00371091) (AM).
Diagnosis
Recognized by the presence of numerous tiny dark spots on the weakly greenish dorsum (pl. 2), and the structure of the male genitalia, the left paramere with a somewhat elongate apex positioned asymmetrically on the anterior side of the structure; endosoma unknown. Most easily confused with M. beaufortiae, M. leuropomae, and M. micranthae, but the spots in those species always larger and the apical medial process much longer in M. beaufortiae and placed medially in all three species (figs. 15, 21, 22, respectively).
Description
Male: Body weakly to distinctly ovoid; mean total length 3.41, mean width pronotum 1.10. COLORATION (pl. 2): Head: Pale or pale with some small dark dots; scapus and pedicellus unicolorous pale; labium generally infuscate, heavily so apically. Thorax: Pronotum pale with tiny brown spots; scutellum pale with brown spots, apex dark; hemelytron unicolorous pale with small brown spots; markings on cuneus present as small dark dots; membrane veins white or pale; hind femur with many brown or black spots; hind tibial spines black with conspicuous dark spots at bases. Abdomen: Venter unicolorous pale or mostly so. SURFACE AND VESTITURE: Dorsum with suberect or reclining black setae and some sericeous or woolly setae. STRUCTURE: Head: Somewhat projecting; interocular space moderate; eyes leaving gena moderately exposed in lateral view. Antenna: Segment 2 weakly tapering, more slender at base; antennal fossa with ventral margin at ventral margin of eye. Labium: Just reaching onto abdomen. Thorax: Pretarsus: Pulvilli present, flaplike, covering about one third of ventral claw surface. GENITALIA (fig. 16): Endosoma: All available specimens teneral and therefore not observed. Phallotheca: Nearly erect; dorsal surface with a curving, posteriorly directed, platelike projection; apex simple. Left Paramere: Shaft at right angles to body; body broad basally, apex medially drawn into a blunt fingerlike process; anterior process triangular in lateral view; posterior process in the form of a long, straight, fingerlike projection; base of posterior process without conspicuous shoulder. Right Paramere: Body short and broad, lanceolate; apex short, clawlike, decurved.
Female: Slightly more ovoid than male; mean total length 3.70, mean width pronotum 1.36. COLORATION (pl. 2): As in male. GENITALIA: Not examined.
Distribution (map 3)
Recorded from one locality on the eastern border of southern South Australia. Melaleucoides brevifoliae is one of only two species in the Melaleucoides genus group that is known to occur outside Western Australia.
Discussion
All available male specimens for this species were apparently partially teneral, because after several dissections we were unable to find a specimen in which the endosoma was suitably sclerotized for observation and illustration.
Holotype
AUSTRALIA: South Australia: Scorpion Springs Cons. Park, 35.60421°S 140.8646°E, 125 m, 10 Nov 1998, Schuh, Cassis, Silveira, Melaleuca brevifolia Turcz. (Myrtaceae), det. Det: Royal Bot. Gard. NSW NSW427362, 1♂ (00130421) (SAMA).
Paratypes
AUSTRALIA: South Australia: Scorpion Springs Cons. Park, 35.60421°S 140.8646°E, 125 m, 10 Nov 1998, Schuh, Cassis, Silveira, Melaleuca brevifolia Turcz. (Myrtaceae), det. Det: Royal Bot. Gard. NSW NSW427362, 9♂ (00087236, 00371099–00371102, 00371121–00371124), 6♀ (00371118–00371120, 00371130–00371132) (AM), 12♂ (00087506, 00130422–00130432), 9♀ (00087237, 00130433–00130440) (AMNH), 1♂ (00371103), 1♀ (00371117) (SAMA).
Other Specimens Examined
AUSTRALIA: South Australia: Scorpion Springs Cons. Park, 35.60421°S 140.8646°E, 125 m, 10 Nov 1998, Schuh, Cassis, Silveira, Melaleuca brevifolia Turcz. (Myrtaceae), det. Det: Royal Bot. Gard. NSW NSW427362, 18 nymphs (00371104–00371116, 00371125–00371129) (AM).
Melaleucoides cassisi, new species
Figures 17, 18; map 3; plate 2
Diagnosis
Recognized among Melaleucoides spp. by the relatively large size (mean total length 3.95), the monotonous pale coloration of the dorsum, and the structure of the male genitalia, notably the clublike aggregation of denticles distad of the secondary gonopore and their position ventral to the primary endosomal strap. Most similar in size and coloration to M. grossi and M. sheathiana. Distinguished from the former by the arrangement of the endosomal denticles on an elongate, conical, free spine ventral to the primary endosomal spine, and from the latter by the strongly orange scutellum and the dorsal position of the endosomal spicules and the fusion of the primary and secondary endosomal straps distad of the secondary gonopore in that species.
Description
Male: Body weakly elongate, weakly ovoid; mean total length 3.95, mean width pronotum 1.38. COLORATION (pl. 2): Head: Uniformly pale, faded yellow; scapus and pedicellus yellow to dirty yellow; labium pale with segment 4 heavily infuscate. Thorax: Pronotum unicolorous, faded yellow; scutellum white; hemelytron unicolorous, faded yellow; cuneus mostly white, markings absent; membrane and veins weakly fumose; hind femur unicolorous pale, without black spots; hind tibial spines black, without dark spots at bases. Abdomen: Venter unicolorous pale or mostly so. SURFACE AND VESTITURE (fig. 17B): Dorsum with suberect or reclining black setae and some sericeous or woolly setae. STRUCTURE: Head (fig. 17A): Barely projecting; interocular space relatively large; eyes leaving gena moderately exposed in lateral view. Antenna: Segment 2 of equal diameter over entire length; antennal fossa with ventral margin at ventral margin of eye. Labium: Just reaching onto abdomen. Thorax: Pretarsus (fig. 17C, D): Pulvilli present, flaplike, covering about one third of ventral claw surface. GENITALIA (figs. 17E, F, 18): Endosoma: Primary strap apically greatly elongate, weakly arcuate and curving ventrally, without ornamentation; secondary endosomal strap fused with primary strap proximal to secondary gonopore, fused with primary strap distal to gonopore ornamented with denticles at about midpoint of apical region of primary strap; spinelike, elongate process arising near gonopore on dorsal surface absent; spinelike, elongate process near gonopore on ventral surface present, strongly recurved toward base of endosoma; bladderlike process distad of secondary gonopore absent; secondary gonopore seen frontally (facing up) in lateral view of endosoma. Phallotheca: More or less right angulate; dorsal surface with a curving, posteriorly directed, platelike projection; apex simple. Left Paramere: Shaft at right angles to body; body spoon shaped; apex medially drawn into a blunt, recurved, fingerlike process; anterior process triangular in lateral view; posterior process in the form of a short, straight, fingerlike projection; base of posterior process without conspicuous shoulder. Right Paramere: Body short and broad, lanceolate; apex short, clawlike, decurved.
Female: Distinctly more ovoid than male; mean total length 4.12, mean width pronotum 1.53. COLORATION (pl. 2): As in male. GENITALIA (fig. 18): Posterior wall laterally with distinct crescent-shaped interramal sclerites; posteriorly without a sclerotized transverse band; posterolaterally with a distinct swelling covered with microtrichia; longitudinal fold on either side of midline along part of length; posterior margin of posterior wall without spicules and not reflexed dorsally; interramal lobes present and asymmetrical, reclining, overlapping, and heavily ornamented with spicules; vestibulum with medial plates sclerotized and readily observed, small, nearly symmetrical, triangular; with sclerotized guide present as short, heavily sclerotized tube on left side.
Etymology
Named for Gerasimos Cassis, who collected most of the known specimens, in recognition of his groundbreaking efforts in field documentation and study of Australian Heteroptera.
Distribution (map 3)
Known from two localities west and north of Yalgoo and in the northeastern Wheatbelt region of Western Australia.
Holotype
AUSTRALIA: Western Australia: 56.6 km W of Yalgoo, 28.42397°S 116.1233°E, 600 m, 27 Oct 1996, Schuh and Cassis, Melaleuca uncinata R. Br. (Myrtaceae), det. Perth 05120640, 1♂ (AMNH_PBI 00371607) (WAMP).
Paratypes
AUSTRALIA: Western Australia: 56.6 km W of Yalgoo, 28.42397°S 116.1233°E, 600 m, 27 Oct 1996, Schuh and Cassis, Melaleuca uncinata R. Br. (Myrtaceae), det. Perth 05120640, 10♂ (00087172, 00371142–00371150), 33♀ (00087173, 00371151–00371182) (AM), 24♂ (00087452, 00371608–00371610, 00371612–00371630, 00371639), 31♀ (00371565–00371593, 00371611, 00371699) (AMNH), 1♂ (00371631) (UCR), 1♂ (00371056), 6♀ (00371057–00371062) (UNSW), 6♂ (00371633–00371638), 21♀ (00371183–00371191, 00371594–00371605) (WAMP).
Other Specimens Examined
AUSTRALIA: Western Australia: 13.5 km W of Nungarin on Rt 50, 31.11547°S 117.945°E, 300 m, 16 Nov 1999, R.T. Schuh, G. Cassis, & R. Silveira, 1♂ (00129374) (UNSW). 56.6 km W of Yalgoo, 28.42397°S 116.1233°E, 600 m, 27 Oct 1996, Schuh and Cassis, Melaleuca uncinata R. Br. (Myrtaceae), det. Perth 05120640, 9♂ (00371133–00371141) (AM), 41♀ (00371641–00371661, 00371663–00371682), 1 nymph (00371662) (AMNH), 1♂ (00371632), 4♀ (00371687–00371690) (CNC), 4♀ (00371683–00371686) (UCR), 1♂ (00371640), 4♀ (00371695–00371698) (USNM), 1♂ (00371606), 4♀ (00371691–00371694) (ZISP).
Diagnosis
Recognized uniquely among Melaleucoides spp. by the entirely castaneous coloration, the large, broad body (pl. 2), and the structure of the male genitalia, with the left paramere elongate but broad apically, and the endosoma with a long, apically oriented, ventral process associated with the secondary gonopore. Coloration most similar to M. rhaphiophyllae and M. uncinatae, but both of those species with the apex of the scutellum and the base of the cuneus pale.
Description
Male: Body broadly ovoid; mean total length 3.61, mean width pronotum 1.44. COLORATION (pl. 2): Head: Uniformly castaneous; scapus unicolorous dark; pedicellus yellow to dirty yellow; labium pale with segment 4 heavily infuscate. Thorax: Pronotum, scutellum, and hemelytron unicolorous castaneous; markings on cuneus absent; membrane and veins very strongly fumose; hind femur unicolorous pale, without black spots; hind tibial spines black, without dark spots at bases. Abdomen: Venter unicolorous dark. SURFACE AND VESTITURE: Dorsum with suberect or reclining black setae, without sericeous or woolly setae. STRUCTURE: Head: Weakly projecting; interocular space relatively large; eyes leaving gena broadly exposed in lateral view. Antenna: Segment 2 weakly tapering, more slender at base; antennal fossa with ventral margin at ventral margin of eye. Labium: Just reaching onto abdomen. Thorax: Pretarsus: Pulvilli present, flaplike, covering about one third of ventral claw surface. GENITALIA (fig. 19): Endosoma: Primary strap apically greatly elongate, nearly straight, apex spadelike, otherwise without ornamentation; secondary endosomal strap fused with primary strap proximal to secondary gonopore, reaching well beyond gonopore, fused with primary strap, ornamented with denticles; spinelike, elongate process arising near gonopore on dorsal surface absent; spinelike, elongate process near gonopore on ventral surface present and erect, parallel to primary endosomal strap; bladderlike process distad of secondary gonopore absent; secondary gonopore seen frontally (facing up) in lateral view of endosoma. Phallotheca: Nearly erect; dorsal surface with a curving, posteriorly directed, platelike projection; ventral surface with a projecting keel; apex simple. Left Paramere: Shaft at right angles to body; body spoon shaped; apex medially elongate and broadly rounded; anterior process triangular in lateral view; posterior process in the form of a short, straight, fingerlike projection; base of posterior process without conspicuous shoulder. Right Paramere: Body short and broad, lanceolate; apex with short fingerlike process.
Female: Body shape as in male; mean total length 3.84, mean width pronotum 1.55. COLORATION (pl. 2): As in male. GENITALIA: Not examined.
Holotype
AUSTRALIA: Western Australia: 4 km W of Lillian Stoke Rock, 33.079°S 120.0669°E, 370 m, 21 Nov 1999, R.T. Schuh and G. Cassis, Melaleuca undulata Benth. (Myrtaceae), det. Perth 05671930, 1♂ (AMNH_PBI 00087372) (WAMP).
Paratypes
AUSTRALIA: Western Australia: 4 km W of Lillian Stoke Rock, 33.079°S 120.0669°E, 370 m, 21 Nov 1999, R.T. Schuh and G. Cassis, Melaleuca undulata Benth. (Myrtaceae), det. Perth 05671930, 1♀ (00372754) (AM), 1♀ (00327133) (AMNH), 1♀ (00087373) (WAMP).
Diagnosis
Recognized among Melaleucoides spp. by the relatively large size (mean total length 3.34), the monotonous pale coloration of the dorsum, and the structure of the male genitalia, notably the secondary endosomal strap dorsal to the primary strap, free, elongate, conical, and uniformly covered with short denticles. Most similar in size and coloration to M. cassisi and M. sheathiana. Distinguished from the former by the arrangement of the endosomal denticles in a clublike fashion ventral to the primary endosomal strap, and from the latter by the strongly orange scutellum and the fusion of the primary and secondary endosomal straps distad of the secondary gonopore with the spicules placed in somewhat irregular arrangement.
Description
Male: Body weakly to distinctly ovoid; mean total length 3.34, mean width pronotum 1.38. COLORATION (pl. 2): Head: Uniformly pale, weakly yellow; scapus and pedicellus yellow to dirty yellow; labium pale with segment 4 heavily infuscate. Thorax: Pronotum unicolorous, weakly yellow; scutellum white to weakly yellow; hemelytron, including cuneus, unicolorous weakly yellow; markings on cuneus absent; membrane weakly fumose, veins entirely red; hind femur unicolorous pale, without black spots; hind tibial spines black, without dark spots at bases. Abdomen: Venter unicolorous pale or mostly so. SURFACE AND VESTITURE: Dorsum with suberect or reclining black setae and some sericeous or woolly setae. STRUCTURE: Head: Barely projecting; interocular space moderate; eyes leaving gena moderately exposed in lateral view. Antenna: Segment 2 of equal diameter over entire length; antennal fossa with ventral margin at ventral margin of eye. Labium: Just reaching onto abdomen. Thorax: Pretarsus: Pulvilli present, flaplike, covering about one third of ventral claw surface. GENITALIA (fig. 20): Endosoma: Primary strap apically greatly elongate, weakly angled dorsally relative to body of endosoma, no ornamentation; secondary endosomal strap fused with primary strap proximal to secondary gonopore, free from primary strap distal to secondary gonopore, ornamented with denticles; spinelike, elongate process arising near gonopore on dorsal surface absent; spinelike, elongate process near gonopore on ventral surface present, recurved toward base of endosoma; bladderlike process distad of secondary gonopore absent; secondary gonopore seen frontally (facing up) in lateral view of endosoma. Phallotheca: Smoothly curving on dorsal margin; dorsal surface with a curving, posteriorly directed, platelike projection; anterior surface with a short to elongate keel; apex simple. Left Paramere: Shaft at right angles to body; body spoon shaped; apex medially broadly rounded; anterior process triangular in lateral view; posterior process in the form of a long, straight, fingerlike projection; base of posterior process without conspicuous shoulder. Right Paramere: Body short and broad, lanceolate; apex with short fingerlike process.
Female: Body shape as in male; mean total length 3.75, mean width pronotum 1.46. COLORATION (pl. 2): As in male. GENITALIA: Not examined.
Etymology
Named for the collector of all known specimens, the late Gordon Gross, in recognition of his contributions to Australian heteropterology.
Distribution (map 4)
Known from Wallaroo in southeastern South Australia. Melaleucoides grossi is one of only two species in the Melaleucoides genus group that occur outside Western Australia.
Holotype
AUSTRALIA: South Australia: North Beach, Wallaroo, 33.906°S 137.63°E, 12 Feb 1964, G.F. Gross, 1♂ (AMNH_PBI 00169227) (SAMA).
Paratypes
AUSTRALIA: South Australia: North Beach, Wallaroo, 33.906°S 137.63°E, 12 Feb 1964, G.F. Gross, 2♂ (00169239, 00169240), 3♀ (00169256–00169258) (AM), 4♂ (00169219–00169220, 00169225, 00169238), 5♀ (00169215, 00169217, 00169246, 00169249, 00169253) (AMNH), 15♂ (00169221–00169224, 00169226, 00169228–00169237), 17♀ (00169209–00169212, 00169216, 00169218, 00169241, 00169243–00169245, 00169247–00169248, 00169250–00169252, 00169254–00169255) (SAMA).
Diagnosis
Recognized by the presence of discrete small dark spots on the dorsum (pl. 2), and the structure of the male genitalia, the left paramere with a somewhat elongate, apex positioned more or less symmetrically and the posterolateral shoulder with a low projection; endosoma with a long row of heavy, elongate denticles on the postgonoporal portion of the secondary strap and recurved ventral process associated with the secondary gonopore. Most easily confused with M. micranthae on the form of the spots and the structure of the male genitalia, but the spots in that species sometimes larger, the apical medial process of the left paramere more nearly symmetrically placed, and the row of denticles on the secondary endosomal strap shorter a and in the form of a more condense grouping. Spots on the dorsum of M. beaufortiae always somewhat coalescent and those in M. brevifoliae tiny and much more numerous.
Description
Male: Body weakly elongate, weakly ovoid; mean total length 3.25, mean width pronotum 1.25. COLORATION (pl. 2): Head: Pale with heavy dark dots; scapus unicolorous pale; pedicellus dirty yellow proximally, weakly to heavily infuscate distally; labium pale with segment 4 heavily infuscate. Thorax: Pronotum pale with heavy dark spots; scutellum pale with brown spots; hemelytron unicolorous pale with scattered dark spots; markings on cuneus present as weakly darkened apex; membrane weakly fumose, veins pale; hind femur with many brown or black spots; hind tibial spines black with conspicuous dark spots at bases. Abdomen: Venter unicolorous pale or mostly so. SURFACE AND VESTITURE: Dorsum with suberect or reclining black setae, without sericeous or woolly setae. STRUCTURE: Head: Weakly projecting; interocular space relatively large; eyes leaving gena broadly exposed in lateral view. Antenna: Segment 2 weakly tapering, more slender at base; antennal fossa with dorsal margin somewhat below ventral margin of eye. Labium: Just reaching onto abdomen. Thorax: Pretarsus: Pulvilli present, flaplike, covering about one third of ventral claw surface. GENITALIA (fig. 21): Endosoma: Primary strap apically greatly elongate, without ornamentation, extreme apex curving dorsally; secondary endosomal strap fused with primary strap proximal to secondary gonopore, reaching well beyond gonopore, with a comblike edge; spinelike, elongate process arising near gonopore on dorsal surface absent; spinelike, elongate process near gonopore on ventral surface present, strongly recurved toward base of endosoma; bladderlike process distad of secondary gonopore absent; secondary gonopore seen frontally (facing up) in lateral view of endosoma. Phallotheca: Nearly erect; anterior surface with a short to elongate keel; apex simple. Left Paramere: Shaft at right angles to body; body more or less quadrate; apex medially drawn into a blunt fingerlike process; apex posterolaterally with a low shoulder; anterior process triangular in lateral view; posterior process in the form of a short, straight, fingerlike projection; base of posterior process without conspicuous shoulder. Right Paramere: Body short and broad, lanceolate; apex short, clawlike, decurved.
Female: Slightly more ovoid than male; mean total length 3.46, mean width pronotum 1.32. COLORATION (pl. 2): As in male. GENITALIA: Posterior wall laterally with distinct crescent-shaped interramal sclerites; posteriorly without a sclerotized transverse band; posterolaterally with a distinct swelling covered with microtrichia; longitudinal fold on either side of midline along part of length; posterior margin of posterior wall without spicules and not reflexed dorsally; interramal lobes present and asymmetrical, reclining, overlapping, and heavily ornamented with spicules; vestibulum with medial plates sclerotized and readily observed, small, nearly symmetrical, triangular, with sclerotized guide present as short heavily sclerotized tube on left side.
Hosts
Most known specimens were taken on Melaleuca leuropoma L.A. Craven (Myrtaceae) (pl. 5C). Additional specimens are recorded from the following species of Myrtaceae: Melaleuca conothamnoides C.A. Gardner, Melaleuca viminea Lindl., and Phymatocarpus porphyrocephalus F. Muell.
Distribution (map 4)
Known from several localities in Western Australia, including one locality in the Wheatbelt region and coastal areas north to Kalbarri.
Holotype
AUSTRALIA: Western Australia: Kalbarri National Park, 7 km E of Kalbarri, 27.68008°S 114.2386°E, 400 m, 29 Oct 1996, Schuh and Cassis, Melaleuca leuropoma L.A. Craven (Myrtaceae), det. Perth 05879175, 1♂ (AMNH_PBI 00137075) (WAMP).
Paratypes
AUSTRALIA: Western Australia: 15 km E of Merredin, 31.37749°S 118.6933°E, 330 m, 16 Nov 1999, R.T. Schuh and G. Cassis, Melaleuca conothamnoides C.A.Gardner (Myrtaceae), det. PERTH staff PERTH 05670624, 3;m (AMNH_PBI 00087325, AMNH_PBI 00087534, AMNH_PBI 00372261), 4;f (AMNH_PBI 00087326, AMNH_PBI 00372262 - AMNH_PBI 00372264) (AM). Brand Hiway 45.9 km S of Dongarra Road, 29.57703°S 115.1348°E, 100 m, 31 Oct 1996, Schuh and Cassis, Phymatocarpus porphyrocephalus F. Muell. (Myrtaceae), det. PERTH staff PERTH 05879264, 1;m (AMNH_PBI 00087192), 1;f (AMNH_PBI 00087193) (AM), 2;m (AMNH_PBI 00130199, AMNH_PBI 00130200), 1;f (AMNH_PBI 00130201) (AMNH). Cervantes, 30.49902°S 115.0684°E, 3 m, 10 Dec 1997, Schuh, Brailovsky, Melaleuca viminea Lindl. (Myrtaceae), det. PERTH staff PERTH 05879205, 1;f (AMNH_PBI 00181857) (AMNH), 3;f (AMNH_PBI 00130552, AMNH_PBI 00181855 - AMNH_PBI 00181856) (WAMP). Kalbarri National Park, 7 km E of Kalbarri, 27.68008°S 114.2386°E, 400 m, 29 Oct 1996, Schuh and Cassis, Melaleuca leuropoma L. A. Craven (Myrtaceae), det. PERTH staff PERTH 05879175, 8;m (AMNH_PBI 00137073 - AMNH_PBI 00137074, AMNH_PBI 00137076 - AMNH_PBI 00137081), 16;f (AMNH_PBI 00137082 - AMNH_PBI 00137097) (AMNH), 1;m (AMNH_PBI 00130202), 3;f (AMNH_PBI 00137098, AMNH_PBI 00137320 - AMNH_PBI 00137321) (WAMP). Kalbarri National Park, 22.9 km E Kalbarri, 27.75408°S 114.3711°E, 500 m, 29 Oct 1996, Schuh and Cassis, 3;m (AMNH_PBI 00135363 - AMNH_PBI 00135365), 1;f (AMNH_PBI 00135366) (WAMP).
Diagnosis
Recognized by the presence of discrete small dark spots on the dorsum (pl. 3), and the structure of the male genitalia, the left paramere with a somewhat elongate, apex positioned somewhat asymmetrically and the posterolateral shoulder with a low projection; endosoma with a weakly elongate, condensed grouping of heavy denticles on the postgonoporal portion of the secondary strap and recurved ventral process associated with the secondary gonopore. Most easily confused with M. leuropomae on the form of the spots and the structure of the male genitalia, but the spots in that species usually smaller, the apical medial process of the left paramere more strongly asymmetrical, and the row of denticles on the secondary endosomal strap longer, with the denticles more evenly spaced. Spots on the dorsum of M. beaufortiae always somewhat coalescent and those in M. brevifoliae tiny and much more numerous.
Description
Male: Body weakly to distinctly ovoid; mean total length 3.14, mean width pronotum 1.22. COLORATION (pl. 3): Head: Pale with a few dark dots; scapus yellow to dirty yellow; pedicellus yellow to dirty yellow; labium uniformly pale. Thorax: Pronotum pale, yellowish, with a few brown spots; scutellum pale, yellowish; hemelytron unicolorous pale, yellowish, with small a few scattered dark spots; markings on cuneus present as a weakly castaneous apex; membrane weakly fumose, veins yellow; hind femur with many brown or black spots; hind tibial spines dark with very small dark bases. Abdomen: Venter unicolorous pale or mostly so. SURFACE AND VESTITURE: Dorsum with suberect or reclining black setae, without sericeous or woolly setae. STRUCTURE: Head: Barely projecting; interocular space moderate; eyes leaving gena broadly exposed in lateral view. Antenna: Segment 2 weakly tapering, more slender at base; antennal fossa with dorsal margin somewhat below ventral margin of eye. Labium: Just reaching onto abdomen. Thorax: Pretarsus: Pulvilli present, flaplike, covering about one third of ventral claw surface. GENITALIA (fig. 22): Endosoma: Primary strap apically greatly elongate, curving dorsally near apex, no ornamentation; secondary endosomal strap fused with primary strap proximal to secondary gonopore, reaching well beyond gonopore, fused with primary strap, and with a few heavy denticles; spinelike, elongate process arising near gonopore on dorsal surface absent; spinelike, elongate process near gonopore on ventral surface present, strongly recurved toward base of endosoma; bladderlike process distad of secondary gonopore absent; secondary gonopore seen frontally (facing up) in lateral view of endosoma. Phallotheca: Smoothly curving on dorsal margin; anterior surface with a short to elongate keel; apex simple. Left Paramere: Shaft at right angles to body; body spoon shaped, apex medially drawn into a blunt fingerlike process; apex posterolaterally with a low shoulder; anterior process slender and cylindrical in lateral view; posterior process in the form of a short, curved, fingerlike projection; base of posterior process without conspicuous shoulder. Right Paramere: Body short and broad, lanceolate; apex short, clawlike, decurved.
Female: Slightly more ovoid than male; mean total length 3.53, mean width pronotum 1.16. COLORATION (pl. 3): As in male. GENITALIA: Not examined.
Distribution (map 4)
Known from two localities in the Boorabbin and Wheatbelt regions in Western Australia.
Holotype
AUSTRALIA: Western Australia: 32 km SE of Paynes Find toward Beacon, 29.48558°S 117.7836°E, 250 m, 12 Dec 1997, Schuh, Brailovsky, 1♂ (AMNH_PBI 00131081) (WAMP).
Paratypes
AUSTRALIA: Western Australia: 32 km SE of Paynes Find toward Beacon, 29.48558°S 117.7836°E, 250 m, 12 Dec 1997, Schuh, Brailovsky, 1♂ (00131082), 5♀ (00131083–00131087) (AMNH). 92.5 km W of Coolgardie at east side of Boorabbin National Park on Great Eastern Hwy, 31.21233°S 120.31°E, 445 m, 17 Nov 1999, R.T. Schuh and G. Cassis, Beaufortia micrantha micrantha Schauer (Myrtaceae), det. Perth 05671957, 5♂ (00087320, 00372252, 00372255–00372257), 5♀ (00087321, 00372265–00372268) (AM), 1♂ (00087530) (AMNH), 2♂ (00372253, 00372254), 2♀ (00372269, 00372270) (WAMP).
Other Specimens Examined
AUSTRALIA: Western Australia: 92.5 km W of Coolgardie at east side of Boorabbin National Park on Great Eastern Hwy, 31.21233°S 120.31°E, 445 m, 17 Nov 1999, R.T. Schuh and G. Cassis, Beaufortia micrantha micrantha Schauer (Myrtaceae), det. Perth 05671957, 10 nymphs (00372271–00372280) (AM).
Diagnosis
Similar in size and coloration to Melaleucoides pileanthicola, the total length not exceeding 3.01, and the strongly reddish clavus and posterior portion of the endocorium. Also similar to M. akaina in size and general coloration. Distinguished from M. akaina and M. pileanthicola by the pale coloration of the cuneus in both of those species; distinguished from M. pileanthicola by the strongly reddish clavus and posterior portion of the endocorium in that species as well as the structure of the endosoma. Further distinguished from M. akaina by short apical endosomal spine and the complete absence of processes associated with the secondary gonopore in that species.
Description
Male: Body moderately elongate, parallel sided; mean total length 2.90, mean width pronotum 1.02. COLORATION (pl. 3): Head: Uniformly pale; scapus and pedicellus unicolorous pale; labium pale with segment 4 heavily infuscate. Thorax: Pronotum pale with red or carmine blotches; scutellum white; hemelytron pale yellow with red suffusion; markings on cuneus present as a contrasting white basal fascia, remainder red; membrane weakly fumose, veins white or pale; hind femur unicolorous pale, without black spots; tibial spines with very small dark bases. Abdomen: Venter unicolorous pale or mostly so. SURFACE AND VESTITURE: Dorsum with reclining simple setae matching background coloration, without sericeous or woolly setae. STRUCTURE: Head: Moderately projecting; interocular space large; eyes leaving gena moderately exposed in lateral view. Antenna: Segment 2 weakly tapering, more slender at base; antennal fossa with ventral margin at ventral margin of eye. Labium: Just reaching onto abdomen. Thorax: Pretarsus: Pulvilli absent. GENITALIA (fig. 23): Endosoma: Primary strap relatively short, straight, with a bifid apex; secondary endosomal strap free of primary strap and much longer, ornamented with denticles near apex; spinelike, elongate process arising near gonopore on dorsal surface present, erect, unornamented; secondary gonopore seen laterally in lateral view of endosoma. Phallotheca: Nearly erect; dorsal surface without a fingerlike projection, and without a platelike projection; anterior surface with a short to elongate keel; ventral surface with a projecting keel; apex simple. Left Paramere: Shaft at right angles to body; body spoon shaped; apex medially broadly rounded to weakly quadrate; anterior process slender and cylindrical in lateral view; posterior process in the form of a short, slender, straight, fingerlike projection; base of posterior process without conspicuous shoulder. Right Paramere: Body short and broad, lanceolate; apex short, clawlike, decurved.
Female: Slightly more ovoid than male; mean total length 3.04, mean width pronotum 1.04. COLORATION (pl. 3): As in male. GENITALIA: Not examined.
Etymology
Named for the late Ozzie Massie, with thanks for his encouragement and interest in our work on the Australian Miridae fauna.
Holotype
AUSTRALIA: Western Australia: 135 km W of Coolgardie on Great Eastern Hiway, 31.27202°S 120.0059°E, 489 m, 17 Nov 1999, R.T. Schuh, G. Cassis, & R. Silveira, Thryptomene kochii E. Pritz. (Myrtaceae), det. Perth 05671302, 1♂ (AMNH_PBI 00371239) (WAMP).
Paratypes
AUSTRALIA: Western Australia: 135 km W of Coolgardie on Great Eastern Hiway, 31.27202°S 120.0059°E, 489 m, 17 Nov 1999, R.T. Schuh, G. Cassis, & R. Silveira, Thryptomene kochii E. Pritz. (Myrtaceae), det. Perth 05671302, 4♂ (00371230, 00371233, 00371236, 00371238), 5♀ (00371242, 00371245, 00371247–00371249) (AM), 3♂ (00371231, 00371234, 00371237), 3♀ (00371241, 00371243–00371244) (AMNH), 3♂ (00371229, 00371232, 00371235), 2♀ (00371240, 00371246) (WAMP).
Melaleucoides pileanthicola, new species
Figures 24, 25; map 4; plate 3
Diagnosis
Similar in size and coloration to M. ozzii, the total length not exceeding 3.01 and the strongly reddish clavus and posterior portion of the endocorium. Also similar to M. akaina in size and general coloration, but distinguished from that species by the short apical endosomal spine and the absence of processes associated with the secondary gonopore and from M. ozzii by the distinctive apical endosomal spine and red cuneus in that species.
Description
Male: Body weakly to distinctly ovoid; mean total length 2.72, mean width pronotum 0.97. COLORATION (pl. 3): Head: Uniformly pale, yellow orange; scapus and pedicellus unicolorous pale; labium pale with segment 4 heavily infuscate. Thorax: Pronotum unicolorous yellow orange; scutellum entirely red to brown; hemelytron with most of endocorium carmine; cuneus pale; membrane and veins strongly fumose; hind femur unicolorous pale, without black spots; hind tibial spines black, without dark spots at bases. Abdomen: Venter unicolorous pale or mostly so. SURFACE AND VESTITURE (fig. 24C): Dorsum with suberect or reclining black setae and some sericeous or woolly setae. STRUCTURE: Head (fig. 24A): Moderately projecting; interocular space large; eyes leaving gena only very slightly exposed in lateral view. Antenna: Segment 2 weakly tapering, more slender at base; antennal fossa with ventral margin at ventral margin of eye. Labium: Just reaching onto abdomen. Thorax: Pretarsus (fig. 24D): Pulvilli absent. GENITALIA (figs. 24E, F, 25): Endosoma: Primary strap apically greatly elongate, weakly arcuate and curving dorsally, no ornamentation; secondary endosomal strap fused with primary strap proximal to secondary gonopore, reaching well beyond gonopore, devoid of ornamentation, free from primary endosomal strap; spinelike, elongate process arising near gonopore on dorsal surface absent; spinelike, elongate process near gonopore on ventral surface present and erect, oriented toward apex of endosoma; bladderlike process distad of secondary gonopore absent; secondary gonopore seen frontally (facing up) in lateral view of endosoma. Phallotheca: Nearly erect; dorsal surface with a curving, posteriorly directed, platelike projection; apex simple. Left Paramere: Shaft at right angles to body; body spoon shaped; apex medially broadly; anterior process triangular in lateral view; posterior process in the form of a long, straight, fingerlike projection; base of posterior process with slightly projecting shoulder. Right Paramere: Body short and broad, lanceolate; apex short, clawlike, decurved.
Female: Body shape as in male; mean total length 2.94, mean width pronotum 1.01. COLORATION (pl. 3): As in male. GENITALIA (fig. 25): Posterior wall laterally with distinct crescent-shaped interramal sclerites; posteriorly without a sclerotized transverse band; posterolaterally with a distinct swelling covered with microtrichia; longitudinal fold on either side of midline along part of length; posterior margin of posterior wall without spicules and not reflexed dorsally; interramal lobes roughly symmetrical, erect, and not ornamented with spicules or with a very few; vestibulum with medial plates sclerotized and readily observed, small, nearly symmetrical, triangular with sclerotized guide present as caplike structure with internal differentiation.
Etymology
Named for the host genus, Pileanthus (Myrtaceae), in combination with the Latin suffix -cola, “inhabitant.”
Hosts
The majority of known specimens were taken on Pileanthus peduncularis borealis Endl. (Myrtaceae). Additional specimens are recorded from the following species of Myrtaceae: Pileanthus peduncularis subsp. pilifer Keighery (pl. 7D), Pileanthus vernicosus F. Muell., Scholtzia leptantha Benth., Thryptomene aspera aspera E. Pritz. (pl. 8B).
Distribution (map 4)
Widespread in Western Australia, from the outback Goldfields to the coastal Carnarvon region.
Holotype
AUSTRALIA: Western Australia: NW Coastal Hiway 57 km N of Kalbarri Road, 27.44756°S 114.6867°E, 500 m, 30 Oct 1996, Schuh and Cassis, Pileanthus peduncularis borealis Endl. (Myrtaceae), det. Perth 05120349, 1♂ (AMNH_PBI 00137356) (WAMP).
Paratypes
AUSTRALIA: Western Australia: 28 km S of Menzies (3.5 km E of Hiway), 29.91917°S 121.1514°E, 500 m, 25 Oct 1996, Schuh and Cassis, Thryptomene aspera aspera E. Pritz. (Myrtaceae), det. Perth 05095093, 4♂ (00087219, 00087510, 00373108–00373109), 2♀ (00373106, 00373107) (AM). 54.3 km N of jct of Agana Kalbarri Rd and Brand Hiway (rest area), 27.47362°S 114.7054°E, 240 m, 24 Oct 2004, Cassis, Wall, Weirauch, Symonds, Pileanthus vernicosus F. Muell. (Myrtaceae), det. Field ID, 4♀ (00373119–00373122) (AM). Kalbarri National Park, Loop Road, 27.56163°S 114.4376°E, 300 m, 28 Oct 1996, Schuh and Cassis, Pileanthus peduncularis borealis Endl. (Myrtaceae), det. Perth 05120586, 7♂ (00373147–00373153), 8♀ (00373156–00373163) (AM), 2♂ (00373154, 00373155) (WAMP). NW Coastal Hiway 57 km N of Kalbarri Road, 27.44756°S 114.6867°E, 500 m, 30 Oct 1996, Schuh and Cassis, Pileanthus peduncularis borealis Endl. (Myrtaceae), det. Perth 05120349, 11♂ (00087215, 00373123–00373132), 9♀ (00087216, 00373136–00373143) (AM), 29♂ (00087468, 00137322–00137324, 00137329–00137353), 32♀ (00137359–00137369, 00137373–00137376, 00137379–00137380, 00137383–00137397) (AMNH), 2♂ (00137354, 00137355), 2♀ (00137377, 00137378) (CNC), 3♂ (00373133–00373135), 3♀ (00373144–00373146) (UNSW), 2♂ (00137357, 00137358), 2♀ (00137381, 00137382) (USNM), 4♂ (00137325–00137328), 11♀ (00137370–00137372, 00137398–00137405) (WAMP).
Other Specimens Examined
AUSTRALIA: Western Australia: 4.5 km NW of jct of Blowholes Rd and North West Coastal Hiway, N of Carnarvon, 24.72267°S 113.7158°E, 28 m, 27 Oct 2004, Cassis, Wall, Weirauch, Tatarnic, Symonds, Scholtzia leptantha Benth. (Myrtaceae), det. Perth6988687, 1♂ (00373175) (AM). 54.3 km N of jct of Agana Kalbarri Rd and Brand Hiway (rest area), 27.47362°S 114.7054°E, 240 m, 24 Oct 2004, Cassis, Wall, Weirauch, Symonds, Pileanthus vernicosus F. Muell. (Myrtaceae), det. Field ID, 3♂ (00373116–00373118) (AM), 1♂ (00373299) (AMNH). 66.2 km E of North West Coastal Hiway on Mardathuna Rd, 24.45443°S 114.5233°E, 103 m, 01 Nov 2004, Cassis, Wall, Weirauch, Tatarnic, Symonds, Pileanthus peduncularis subsp. pilifer Keighery (Myrtaceae), det. Perth 6989985, 2♂ (00373110, 00373111), 1♀ (00373112) (AM).
Diagnosis
Most similar to M. uncinatae in the heavily castaneous coloration (pls. 3, 4) of the dorsum, but with the apex of the scutellum and the base of the cuneus pale. Distinguished from that species by the different form of the phallotheca and the left paramere (figs. 26, 31), the endosoma of M. rhaphiophyllae being unknown.
Description
Male: Body moderately elongate, parallel sided; mean total length 2.87, mean width pronotum 1.22. COLORATION (pl. 3): Head: Uniformly dark; scapus brown; pedicellus unicolorous black or castaneous; labium generally infuscate, heavily so apically. Thorax: Pronotum unicolorous deep red to black; scutellum red or dark brown with white or pale tip; corium and clavus mostly castaneous, markings on cuneus present as a contrasting white basal fascia, remainder castaneous to black; membrane fumose, veins white; hind femur with many brown or black spots; hind tibial spines with conspicuous dark spots at bases. Abdomen: Venter heavily red laterally. SURFACE AND VESTITURE: Dorsum with golden reclining setae; dorsum without sericeous or woolly setae. STRUCTURE: Head: Weakly projecting; interocular space moderate; eyes leaving gena broadly exposed in lateral view. Antenna: Segment 2 weakly tapering, more slender at base; antennal fossa with dorsal margin somewhat below ventral margin of eye. Labium: Just reaching onto abdomen. Thorax: Pretarsus: Pulvilli present, flaplike, covering about one third of ventral claw surface. GENITALIA (fig. 26): Endosoma: All available specimens teneral and therefore not observed. Phallotheca: Nearly erect; dorsal surface with a fingerlike projection; apex simple. Left Paramere: Shaft in same axis as body; body spoon shaped; apex medially angulate; anterior process triangular in lateral view; posterior process in the form of a short, curved, fingerlike projection; base of posterior process with distinct shoulder. Right Paramere: Body short and broad, lanceolate; apex short, clawlike, decurved.
Female: Slightly more ovoid than male; mean total length 2.98, mean width pronotum 1.20. COLORATION (pl. 3): Paler than male. GENITALIA: Not examined.
Distribution (map 4)
Known from two localities on the Carnamah Shire Coast (obscured on the map) and the eastern agricultural region in Western Australia.
Discussion
All available male specimens for this species were apparently partially teneral, because after several dissections we were unable to find a specimen in which the endosoma was suitably sclerotized for observation and illustration.
Holotype
AUSTRALIA: Western Australia: Brand Hiway 8.2 km N of Eneabba, 29.7462°S 115.254°E, 100 m, 31 Oct 1996, Schuh and Cassis, Melaleuca rhaphiophylla Schauer (Myrtaceae), det. Perth 05120195, 1♂ (AMNH_PBI 00128826) (WAMP).
Paratypes
AUSTRALIA: Western Australia: Brand Hiway 8.2 km N of Eneabba, 29.7462°S 115.254°E, 100 m, 31 Oct 1996, Schuh and Cassis, Melaleuca rhaphiophylla Schauer (Myrtaceae), det. Perth 05120195, 4♂ (00087265–00087266, 00128827, 00372296), 2♀ (00128828, 00372298) (AM), 2♂ (00372294, 00372297) (AMNH), 1♂ (00372295), 1♀ (00128829) (WAMP).
Other Specimens Examined
AUSTRALIA: Western Australia: Moorine Rocks, 11.7 km N of Great Eastern Hiway on Noongar Road, 31.22843°S 118.979°E, 345 m, 04 Dec 1997, Schuh, Cassis, Brailovsky, Asquith, Melaleuca sp. (Myrtaceae), det. field ID; host 97-06, 1♂ (00371776), 1♀ (00371777) (AMNH).
Diagnosis
Recognized among Melaleucoides spp. by the relatively large size (mean total length 3.67), the monotonous pale coloration of the dorsum in conjunction with a conspicuously orange scutellum, and the structure of the male genitalia, with the fusion of the primary and secondary endosomal straps distad of the secondary gonopore and the spicules placed in somewhat irregular arrangement. Most similar in size and coloration to M. cassisi and M. grossi. Distinguished from the former by the arrangement of the endosomal denticles in a clublike fashion ventral to the primary endosoma strap, and from the latter by the secondary endosomal strap dorsal to the primary strap, free, elongate, conical, and uniformly covered with short denticles.
Description
Male: Body weakly elongate, weakly ovoid; mean total length 3.67, mean width pronotum 1.37. COLORATION (pl. 3): Head: Uniformly dirty yellow; scapus unicolorous dark; pedicellus yellow to dirty yellow; labium pale with segment 4 heavily infuscate. Thorax: Pronotum unicolorous dirty yellow-green; scutellum entirely orange; hemelytron unicolorous yellow-green; markings on cuneus absent; membrane and veins weakly fumose; hind femur unicolorous pale, without black spots; hind tibial spines black without dark spots at bases. Abdomen: Venter unicolorous pale or mostly so. SURFACE AND VESTITURE: Dorsum with suberect or reclining black setae, without sericeous or woolly setae. STRUCTURE: Head: Barely projecting; interocular space moderate; eyes leaving gena moderately exposed in lateral view. Antenna: Segment 2 weakly tapering, more slender at base; antennal fossa with ventral margin at ventral margin of eye. Labium: Just reaching onto abdomen. Thorax: Pretarsus: Pulvilli present, flaplike, covering about one third of ventral claw surface. GENITALIA (fig. 27): Endosoma: Primary strap apically greatly elongate, weakly arcuate and curving dorsally, no ornamentation; secondary endosomal strap fused with primary strap proximal to secondary gonopore, reaching well beyond gonopore, ornamented with denticles; spinelike, elongate process arising near gonopore on dorsal surface absent; spinelike, elongate process near gonopore on ventral surface present, recurved toward base of endosoma; bladderlike process distad of secondary gonopore absent; secondary gonopore seen frontally (facing up) in lateral view of endosoma. Phallotheca: Smoothly curving on dorsal margin; dorsal surface with a curving, posteriorly directed, platelike projection; apex simple. Left Paramere: Shaft at right angles to body; body spoon shaped; apex medially broadly rounded; anterior process triangular in lateral view; posterior process in the form of a short, straight, fingerlike projection; base of posterior process without conspicuous shoulder. Right Paramere: Body short and broad, lanceolate; apex short, clawlike, decurved.
Female: Slightly more ovoid than male; mean total length 3.82, mean width pronotum 1.43. COLORATION (pl. 3): As in male. GENITALIA: Posterior wall laterally with distinct crescent-shaped interramal sclerites; posteriorly without a sclerotized transverse band; posterolaterally with a distinct swelling covered with microtrichia; longitudinal fold on either side of midline along part of length; posterior margin of posterior wall without spicules and not reflexed dorsally; interramal lobes present and asymmetrical, reclining, overlapping, and heavily ornamented with spicules; vestibulum with medial plates sclerotized and readily observed, small, nearly symmetrical, triangular, with sclerotized guide present as caplike structure with internal differentiation.
Holotype
AUSTRALIA: Western Australia: 11 km N of Coolgardie-Esperance Hiway on Kambalda Road, 31.25231°S 121.5899°E, 320 m, 18 Nov 1999, R.T. Schuh, G. Cassis, & R. Silveira, Melaleuca sheathiana W. Fitzg. (Myrtaceae), det. Perth 05671396, 1♂ (AMNH_PBI 00371554) (WAMP).
Paratypes
AUSTRALIA: Western Australia: 11 km N of Coolgardie-Esperance Hiway on Kambalda Road, 31.25231°S 121.5899°E, 320 m, 18 Nov 1999, R.T. Schuh, G. Cassis, & R. Silveira, Melaleuca sheathiana W.Fitzg. (Myrtaceae), det. Perth 05671396, 8♂ (00087301, 00087499, 00371970–00371973, 00371975), 29♀ (00087302, 00371976, 00371978, 00371980–00371984, 00372031–00372035, 00372066–00372081) (AM), 17♂ (00129423–00129427, 00129504, 00129584–00129585, 00371552, 00371555–00371562), 36♀ (00129428–00129444, 00129505–00129512, 00129587–00129594, 00371553, 00371563–00371564) (AMNH), 3♀ (00371985–00371987) (UNSW). 91.4 km SE of Southern Cross, 31.97145°S 119.287°E, 375 m, 04 Dec 1997, Schuh, Cassis, Brailovsky, Asquith, Melaleuca teuthidoides Barlow (Myrtaceae), det. Perth 05054834, 6♂ (00130795–00130800), 6♀ (00130801–00130806) (WAMP).
Other Specimens Examined
AUSTRALIA: Western Australia: 11 km N of Coolgardie-Esperance Hiway on Kambalda Road, 31.25231°S 121.5899°E, 320 m, 18 Nov 1999, R.T. Schuh, G. Cassis, & R. Silveira, Melaleuca sheathiana W. Fitzg. (Myrtaceae), det. Perth 05671396, 1♂ (00371314) (AM). 33.3 km S of Norseman, 32.46461°S 121.6778°E, 300 m, 19 Nov 1999, R.T. Schuh, G. Cassis, & R. Silveira, 2♂ (00371315, 00089117), 1♂ (00371966) (AM). 91.4 km SE of Southern Cross, 31.97145°S 119.287°E, 375 m, 04 Dec 1997, Schuh, Cassis, Brailovsky, Asquith, Melaleuca teuthidoides Barlow (Myrtaceae), det. Perth 05054834, 1♂ (00087308) (AM), 7♀ (00373099–00373105) (AMNH).
Diagnosis
Recognized by the yellow-orange coloration of the body and appendages, the elongate slender form of the body in most specimens, and the form of the male genitalia with the elongate, fingerlike apical projection on the apex of the left paramere. Most similar in size and coloration to M. verticordiae, but easily distinguished by the comblike fringe on the apical spine of the endosoma in that species.
Description
Male: Body greatly elongate, parallel sided; mean total length 3.38, mean width pronotum 1.11. COLORATION (pl. 3): Head: Uniformly yellow; scapus and pedicellus yellow to dirty yellow; labium pale with segment 4 heavily infuscate. Thorax: Pronotum unicolorous yellow; scutellum unicolorous yellow; hemelytron, including cuneus, unicolorous yellow; markings on cuneus absent; membrane weakly fumose, veins yellow; hind femur unicolorous pale, without black spots; hind tibial spines black, without dark spots at bases. Abdomen: Venter unicolorous pale or mostly so. SURFACE AND VESTITURE: Dorsum with suberect or reclining black setae, without sericeous or woolly setae. STRUCTURE: Head: Somewhat projecting; interocular space relatively large; eyes leaving gena moderately exposed in lateral view. Antenna: Segment 2 of equal diameter over entire length; antennal fossa with ventral margin at ventral margin of eye. Labium: Reaching to about anterior margin of pygophore. Thorax: Pretarsus: Pulvilli absent. GENITALIA (fig. 28): Endosoma: Primary strap apically greatly elongate, angled dorsally relative to body of endosoma, denticulate on apical half of ventral margin; secondary endosomal strap fused with primary strap primal to secondary gonopore, fused with primary strap distad of gonopore; spinelike, elongate process arising near gonopore on dorsal surface present, erect, unornamented; spinelike, elongate process near gonopore on ventral surface present, sharply recurved toward base of endosoma; bladderlike process distad of secondary gonopore present, balloonlike; secondary gonopore seen frontally (facing up) in lateral view of endosoma. Phallotheca: Nearly erect; apex simple. Left Paramere: Shaft at right angles to body; body more or less quadrate; apex medially broadly rounded; apex posterolaterally with a greatly elongate projection (fig. 28); anterior process triangular in lateral view; posterior process in the form of a short, straight, fingerlike projection; base of posterior process without conspicuous shoulder. Right Paramere: Body short and broad, lanceolate; apex short, clawlike, decurved.
Female: Distinctly more ovoid than male; mean total length 3.11, mean width pronotum 1.08. COLORATION (pl. 3): As in male. GENITALIA: Not examined.
Etymology
Named for the remarkable similarity of appearance with Melaleucoides verticordiae, new species.
Hosts
Recorded from Melaleuca conothamnoides C.A. Gardner, Scholtzia drummondii Benth., and Thryptomene aspera glabra E. Pritz. (Myrtaceae). The very limited numbers of available specimens preclude conclusions regarding host specificity in this taxon.
Holotype
AUSTRALIA: Western Australia: 15 km E of Merredin, 31.37749°S 118.6933°E, 330 m, 16 Nov 1999, R.T. Schuh and G. Cassis, Melaleuca conothamnoides C.A. Gardner (Myrtaceae), det. Perth 05670624, 1♂ (AMNH_PBI 00370994) (WAMP).
Paratypes
AUSTRALIA: Western Australia: 11 km S of Eneabba, Eneabba National Park [96-50], 29.9025°S 115.24321°E, 150 m, 01 Nov 1996, Schuh and Cassis, Scholtzia drummondii Benth. (Myrtaceae), det. Perth 05120209, 1♂ (00087307) (AMNH). 15 km E of Merredin, 31.37749°S 118.6933°E, 330 m, 16 Nov 1999, R.T. Schuh and G. Cassis, Melaleuca conothamnoides C.A.Gardner (Myrtaceae), det. Perth 05670624, 1♀ (00370993) (AM). 24 km W of Sandstone, 28.01426°S 119.0474°E, 650 m, 26 Oct 1996, Schuh and Cassis, Thryptomene aspera glabra E. Pritz. (Myrtaceae), det. Perth 05095182, 1♂ (00135637) (AM), Thryptomene aspera glabra E. Pritz. (Myrtaceae), det. Perth 05095182, 1♂ (00135638) (AMNH).
Diagnosis
Males recognized in most specimens by the castaneous pronotum anterior lobe, scutellum, and posterior half of the endocorium; phallotheca with a very broad reflexed flange on the dorsal margin; left paramere very broad apically; endosoma with a ventral process associated with the secondary gonopore and no denticles distad of the gonopore. Except for rather variable coloration with the taxon, M. systenae is not easily confused with any other known species of Melaleucoides.
Description
Male: Body weakly to distinctly ovoid; mean total length 3.23, mean width pronotum 1.22. COLORATION (pl. 4): Head: Mostly dark with pale gula and posterior head margin; scapus unicolorous dark; pedicellus black at extreme base, remainder pale; labium generally infuscate, heavily so apically. Thorax: Pronotum usually with dark anterior lobe and contrasting lighter-colored posterior lobe; scutellum entirely dark; clavus pale, corium pale on anterior half, dark posteriorly; markings on cuneus present as a contrasting white basal fascia; membrane fumose, veins white or pale; hind femur nearly unicolorous dark; hind tibial spines dark with very small dark bases. Abdomen: Venter unicolorous dark. SURFACE AND VESTITURE: Dorsum with suberect or reclining black setae, without sericeous or woolly setae. STRUCTURE: Head: Weakly projecting; interocular space relatively large; eyes leaving gena broadly exposed in lateral view. Antenna: Segment 2 weakly tapering, more slender at base; antennal fossa with ventral margin at ventral margin of eye. Labium: Just reaching onto abdomen. Thorax: Pretarsus: Pulvilli present, flaplike, covering about one third of ventral claw surface. GENITALIA (fig. 29): Endosoma: Primary strap apically elongate, angled dorsal relative to body of endosoma, without ornamentation; secondary endosomal strap fused with primary strap proximal to secondary gonopore, reaching only to level of secondary gonopore; spinelike elongate process arising near gonopore on dorsal surface absent; spinelike, elongate process near gonopore on ventral surface present, relatively short, recurved toward base of endosoma; bladderlike process distad of secondary gonopore present, elongate, and paralleling primary endosomal strap; secondary gonopore seen frontally (facing up) in lateral view of endosoma. Phallotheca: Nearly erect; dorsal surface with a curving, posteriorly directed, platelike projection; ventral surface with a projecting keel; apex simple. Left Paramere: Shaft at right angles to body; body spoon shaped; apex quadrate; anterior process triangular in lateral view; posterior process in the form of a short, straight, fingerlike projection; base of posterior process without conspicuous shoulder. Right Paramere: Body short and broad, lanceolate; apex with short fingerlike process.
Female: Slightly more ovoid than male; mean total length 3.51, mean width pronotum 1.32. COLORATION (pl. 4): Paler than male, more strongly reddish, without the nearly castaneous quality of most males. GENITALIA: Posterior wall laterally with distinct crescent-shaped interramal sclerites; posteriorly without a sclerotized transverse band; posterolaterally with a distinct swelling covered with microtrichia; longitudinal fold on either side of midline along part of length; posterior margin of posterior wall without spicules and not reflexed dorsally; interramal lobes present and asymmetrical, reclining, overlapping, and heavily ornamented with spicules; vestibulum with medial plates sclerotized and readily observed, small, nearly symmetrical, triangular, with sclerotized guide present as short heavily sclerotized tube on left side.
Hosts
Melaleuca laetifica Craven [ms. name] (pl. 5B), Melaleuca systena Craven, Melaleuca urceolaris Benth., Melaleuca viminea Lindl., and Phymatocarpus porphyrocephalus F. Muell. (Myrtaceae).
Distribution (map 5)
Relatively widespread in Western Australia, with localities ranging from the Nullabor Plain in the southeast to Kalbarri National Park in the north.
Holotype
AUSTRALIA: Western Australia: 8.2 km E of Indian Ocean Rd on Coorow-Greenhead Rd, Lesuer National Park, 30.04767°S 115.0551°E, 30 m, 06 Nov 2004, Cassis, Weirauch, Tatarnic, Symonds, Melaleuca systena Craven (Myrtaceae), det. Perth 6990401, 1♂ (AMNH_PBI 00368603) (WAMP).
Paratypes
AUSTRALIA: Western Australia: 8.2 km E of Indian Ocean Rd on Coorow-Greenhead Rd, Lesuer National Park, 30.04767°S 115.0551°E, 30 m, 06 Nov 2004, Cassis, Weirauch, Tatarnic, Symonds, Melaleuca systena Craven (Myrtaceae), det. Perth6990401, 2♂ (00368602, 00368604), 3♀ (00368606–00368608) (AM), 2♂ (00368600, 00368601), 3♀ (00368609–00368610, 00368613) (AMNH) 2♀ (00368617, 00368618) (UNSW), 3♀ (00368614–00368616) (WAMP). Brand Hiway 45.9 km S of Dongarra Road, 29.57703°S 115.1348°E, 100 m, 31 Oct 1996, Schuh and Cassis, Phymatocarpus porphyrocephalus F. Muell. (Myrtaceae), det. Perth 05879264, 2♂ (00130029, 00130030), 3♀ (00130031–00130033) (AMNH). Cervantes, 30.49902°S 115.0684°E, 3 m, 10 Dec 1997, Schuh, Brailovsky, Melaleuca viminea Lindl. (Myrtaceae), det. Perth 05879205, 1♂ (00130601), 4♀ (00130602–00130605) (AMNH), 3♀ (00130606–00130608) (WAMP). Frank Hann National Park, 37 km E of Lake King, 33.07753°S 120.0918°E, 400 m, 05 Nov 1996, Schuh and Cassis, Melaleuca sp. (Myrtaceae), det. Perth 05236908, 1♂ (00087405), 1♀ (00087406) (AM). Kalbarri National Park, 22.9 km E Kalbarri, 27.75408°S 114.3711°E, 500 m, 29 Oct 1996, Schuh and Cassis, 6♀ (00135367–00135372) (WAMP). Kalbarri National Park, 37.7 km E Kalbarri, 27.8482°S 114.4746°E, 500 m, 29 Oct 1996, Schuh and Cassis, Melaleuca laetifica Craven ms (Myrtaceae), det. Perth 05054540, 2♂ (00135064, 00135065), 5♀ (00368631–00368632, 00368635–00368636, 00368638) (AM), 6♂ (00135059–00135063, 00135066), 11♀ (00368629–00368630, 00368633–00368634, 00368637, 00368639–00368644) (AMNH), 2♂ (00135067, 00135068), 4♀ (00368625–00368628) (WAMP).
Other Specimens Examined
AUSTRALIA: Western Australia: 1 km S of Lillian Stoke Rock, 33.07681°S 120.0982°E, 380 m, 21 Nov 1999, R.T. Schuh and G. Cassis, Verticordia chrysantha Endl. (Myrtaceae), det. Perth 05672023, 1♀ (00371784) (AM). 4.5 km S of Jurien on Indian Ocean Rd, 30.33667°S 115.069°E, 18 m, 06 Nov 2004, Cassis, Weirauch, Tatarnic, Symonds, Melaleuca systena Craven (Myrtaceae), det. Perth6987567, 1♀ (00371783) (AM). 8.2 km E of Indian Ocean Rd on Coorow-Greenhead Rd, Lesuer National Park, 30.04767°S 115.0551°E, 30 m, 06 Nov 2004, Cassis, Weirauch, Tatarnic, Symonds, Melaleuca systena Craven (Myrtaceae), det. Perth 6990401, 4 nymphs (00368619, 00368621, 00368623–00368624) (AM). 11 km S of Eneabba on Brand Hiway, Eneabba Reserve, 29.91094°S 115.1175°E, 100 m, 21 Oct 2004, Cassis, Wall, Weirauch, Symonds, Melaleuca urceolaris Benth. (Myrtaceae), det. Perth 6986919, 5♀ (00371778–00371782) (AM). Cervantes, 30.49902°S 115.0684°E, 3 m, 10 Dec 1997, Schuh, Brailovsky, Melaleuca viminea Lindl. (Myrtaceae), det. Perth 05879205, 1 nymph (00372247) (AM).
Melaleucoides uncinatae, new species
Figures 30, 31; map 6; plate 4
Diagnosis
Most similar to M. rhaphiophyllae in the heavily castaneous coloration (pls. 3, 4) of the dorsum, with the apex of the scutellum and the base of the cuneus pale. Distinguished from that species by the different form of the phallotheca and the left paramere (figs. 26, 31), the endosoma of M. rhaphiophyllae being unknown.
Description
Male: Body moderately elongate, parallel sided; mean total length 2.99, mean width pronotum 1.25. COLORATION (pl. 4): Head: Uniformly dark; scapus brown; pedicellus unicolorous black or castaneous; labium generally infuscate, heavily so apically. Thorax: Pronotum unicolorous deep red to black; scutellum red or dark brown with white or pale tip; corium and clavus largely castaneous, markings on cuneus present as a contrasting white basal fascia, remainder castaneous; membrane strongly fumose, veins white; hind femur nearly unicolorous dark; hind tibial spines with conspicuous dark spots at bases. Abdomen: Venter heavily red laterally. SURFACE AND VESTITURE: Dorsum with golden reclining setae; dorsum without sericeous or woolly setae. STRUCTURE: Head (fig. 30A): Barely projecting; interocular space moderate; eyes leaving gena moderately exposed in lateral view. Antenna: Segment 2 weakly tapering, more slender at base; antennal fossa with ventral margin at ventral margin of eye. Labium: Just reaching onto abdomen. Thorax: Pretarsus (fig. 30C): Pulvilli present, flaplike, covering about one third of ventral claw surface. GENITALIA (figs. 30D–F, 31): Endosoma: Primary strap apically weakly elongate, weakly curving, without ornamentation; secondary endosomal strap fused with primary strap proximal to level of secondary gonopore, and not projecting beyond gonopore; spinelike, elongate process arising near gonopore on dorsal surface present, erect, ornamented with denticles apically; spinelike, elongate process near gonopore on ventral surface absent; bladderlike process distad of secondary gonopore present; secondary gonopore seen laterally in lateral view of endosoma. Phallotheca: More or less right angulate; dorsal surface with a long fingerlike projection; apex simple. Left Paramere: Shaft in same axis as body; body spoon shaped; apex medially broadly rounded; anterior process triangular in lateral view; posterior process in the form of a short, curved, fingerlike projection; base of posterior process with distinct shoulder. Right Paramere: Body short and broad, lanceolate; apex short, clawlike, decurved.
Female: Distinctly more ovoid than male; mean total length 3.26, mean width pronotum 1.34. COLORATION (pl. 4): Paler than male. GENITALIA: Posterior wall laterally with distinct crescent-shaped interramal sclerites; posteriorly without a sclerotized transverse band; posterolaterally with a distinct swelling covered with microtrichia; longitudinal fold on either side of midline along part of length; posterior margin of posterior wall without spicules and not reflexed dorsally; interramal lobes roughly symmetrical, erect, and heavily ornamented with spicules; vestibulum with medial plates sclerotized and readily observed, small, nearly symmetrical, triangular, with sclerotized guide present as short heavily sclerotized tube on left side.
Hosts
Nearly all available specimens are recorded from Melaleuca uncinata R. Br. (pl. 6D); a single specimen is recorded from Melaleuca rhaphiophylla Schauer (Myrtaceae).
Holotype
AUSTRALIA: Western Australia: 56.6 km W of Yalgoo, 28.42397°S 116.1233°E, 600 m, 27 Oct 1996, Schuh and Cassis, Melaleuca uncinata R.Br. (Myrtaceae), det. Perth 05120640, 1♂ (AMNH_PBI 00368315) (WAMP).
Paratypes
AUSTRALIA: Western Australia: 56.6 km W of Yalgoo, 28.42397°S 116.1233°E, 600 m, 27 Oct 1996, Schuh and Cassis, Melaleuca uncinata R. Br. (Myrtaceae), det. Perth 05120640, 3♂ (00136545–00136546, 00368325), 12♀ (00136547–00136558) (AM), 14♂ (00368313–00368314, 00368316–00368324, 00368327–00368328, 00368364), 16♀ (00368329–00368344) (AMNH), 15♂ (00368326, 00368353–00368363, 00368365–00368367) (WAMP). Brand Hiway 8.2 km N of Eneabba, 29.7462°S 115.254°E, 100 m, 31 Oct 1996, Schuh and Cassis, Melaleuca rhaphiophylla Schauer (Myrtaceae), det. Perth 05120195, 1♀ (00008726) (AM).
Other Specimens Examined
AUSTRALIA: Western Australia: 56.6 km W of Yalgoo, 28.42397°S 116.1233°E, 600 m, 27 Oct 1996, Schuh and Cassis, Melaleuca uncinata R. Br. (Myrtaceae), det. Perth 05120640, 8♀ (00368345–00368352) (AMNH).
Diagnosis
Recognized by the castaneous calli, the elongate red markings on the hemelytron (pl. 4), and the endosoma lacking denticles distad of the secondary gonopore, a dorsal process, and a bladderlike process just distad of the gonopore (fig. 32). Most easily confused with M. annae on the basis of coloration, but the red markings on the hemelytron less strongly linear in that species (pl. 1) and the endosoma with denticles on the secondary strap distad of the secondary gonopore, a recurved ventral process associated with the secondary gonopore, and no bladderlike process (fig. 13).
Description
Male: Body weakly elongate, weakly ovoid; mean total length 3.03, mean width pronotum 1.17. COLORATION (pl. 4): Head: Pale, clypeus dark, frons with large dark bilateral markings; scapus unicolorous pale; pedicellus unicolorous pale; labium pale with segment 4 heavily infuscate. Thorax: Pronotum pale with castaneous calli and humeri; scutellum pale; hemelytron pale with longitudinal castaneous stripes along veins; markings on cuneus present as a red or castaneous apex, otherwise pale; membrane weakly fumose, veins white; hind femur with many brown or black spots; hind tibial spines with conspicuous dark spots at bases. Abdomen: Venter unicolorous pale or mostly so. SURFACE AND VESTITURE: Dorsum with golden reclining setae; dorsum without sericeous or woolly setae. STRUCTURE: Head: Barely projecting; interocular space moderate; eyes leaving gena moderately exposed in lateral view. Antenna: Segment 2 weakly tapering, more slender at base; antennal fossa with ventral margin at ventral margin of eye. Labium: Just reaching onto abdomen. Thorax: Pretarsus: Pulvilli present, flaplike, covering about one third of ventral claw surface. GENITALIA (fig. 32): Endosoma: Primary strap apically weakly elongate, weakly arcuate and curving dorsally, without ornamentation; secondary endosomal strap fused with primary strap proximal to secondary gonopore, reaching only to level of secondary gonopore; elongate process arising near gonopore on dorsal surface present, erect, curving apically, unornamented; spinelike, elongate process near gonopore on ventral surface absent; bladderlike process distad of secondary gonopore present; secondary gonopore seen laterally in lateral view of endosoma. Phallotheca: Nearly erect; dorsal surface with a fingerlike projection and a curving, posteriorly directed, platelike projection; apex simple. Left Paramere: Shaft in same axis as body; body spoon shaped; apex medially angulate, bluntly acuminate; anterior process slender and cylindrical in lateral view; posterior process in the form of a short, straight, fingerlike projection; base of posterior process with distinct shoulder. Right Paramere: Body short and broad, lanceolate; apex short, clawlike, decurved.
Female: Slightly more ovoid than male; mean total length 3.27, mean width pronotum 1.25. COLORATION (pl. 4): As in male. GENITALIA: Not examined.
Holotype
AUSTRALIA: Western Australia: Kalbarri National Park, 22.9 km E Kalbarri, 27.75408°S 114.3711°E, 500 m, 29 Oct 1996, Schuh and Cassis, Melaleuca undulata Benth. (Myrtaceae), det. Perth 05120403, 1♂ (AMNH_PBI 00372306) (WAMP).
Paratypes
AUSTRALIA: Western Australia: Kalbarri National Park, 22.9 km E Kalbarri, 27.75408°S 114.3711°E, 500 m, 29 Oct 1996, Schuh and Cassis, Melaleuca undulata Benth. (Myrtaceae), det. Perth 05120403, 9♂ (00372300–00372305, 00372307, 00372310–00372311), 6♀ (00372319–00372320, 00372323, 00372326–00372328) (AM), 6♂ (00087483, 00372299, 00372308–00372309, 00372312–00372313), 5♀ (00372321–00372322, 00372324–00372325, 00372331) (AMNH), 6♂ (00087275, 00372314–00372318), 5♀ (00087276, 00372329–00372330, 00372332–00372333) (WAMP).
Other Specimens Examined
AUSTRALIA: Western Australia: Kalbarri National Park, 22.9 km E Kalbarri, 27.75408°S 114.3711°E, 500 m, 29 Oct 1996, Schuh and Cassis, Melaleuca undulata Benth. (Myrtaceae), det. Perth 05120403, 6 nymphs (00372334–00372339) (AM).
Melaleucoides verticordiae, new species
Figures 33, 34; map 6; plate 4
Diagnosis
Recognized by the yellow-orange coloration of the body and appendages, the elongate slender form of the body in most specimens, and the form of the male genitalia with the comblike fringe on the apical spine of the endosoma. Most similar in size and coloration to M. similis, but easily distinguished by the elongate, fingerlike apical projection on the apex of the left paramere in that species.
Description
Male: Body greatly elongate, parallel sided; mean total length 3.49, mean width pronotum 1.19. COLORATION (pl. 4): Head: Uniformly yellow; scapus and pedicellus yellow to dirty yellow; labium pale with segment 4 heavily infuscate. Thorax: Pronotum unicolorous yellow; scutellum unicolorous yellow; hemelytron, including cuneus, unicolorous yellow; markings on cuneus absent; membrane weakly fumose, veins yellow; hind femur unicolorous pale, without black spots; hind tibial spines black, without dark spots at bases. Abdomen: Venter unicolorous pale or mostly so. SURFACE AND VESTITURE (fig. 33C): Dorsum with suberect or reclining black setae and some sericeous or woolly setae. STRUCTURE: Head (fig. 33A): Moderately projecting; interocular space large; eyes leaving gena moderately exposed in lateral view. Antenna: Segment 2 of equal diameter over entire length; antennal fossa with ventral margin at ventral margin of eye. Labium: Reaching to about anterior margin of pygophore. Thorax: Pretarsus (fig. 33D, E): Pulvilli absent. GENITALIA (figs. 33F, G, 34): Endosoma: Primary strap apically greatly elongate and near straight, with a comblike ventral margin; secondary endosomal strap fused with primary strap proximal to secondary gonopore, fused with primary strap distad of secondary gonopore; spinelike, elongate process arising near gonopore on dorsal surface present, erect, slender, relatively short, unornamented; spinelike, elongate process near gonopore on ventral surface present, recurved toward base of endosoma; bladderlike process distad of secondary gonopore absent; secondary gonopore seen laterally in lateral view of endosoma. Phallotheca: Smoothly curving on dorsal margin; ventral surface with a projecting, fingerlike keel; apex simple. Left Paramere: Shaft at right angles to body; body spoon shaped; apex medially broadly rounded; anterior process triangular in lateral view; posterior process in the form of a short, straight, fingerlike projection; base of posterior process with distinct shoulder. Right Paramere: Body short and broad, lanceolate; apex short, clawlike, decurved.
Female: Distinctly more ovoid than male; mean total length 3.59, mean width pronotum 1.24. COLORATION (pl. 4): As in male. GENITALIA (fig. 34): Posterior wall laterally with distinct crescent-shaped interramal sclerites; posteriorly without a sclerotized transverse band; posterolaterally with a distinct swelling covered with microtrichia; longitudinal fold on either side of midline along part of length; posterior margin of posterior wall without spicules and not reflexed dorsally; interramal lobes roughly symmetrical, erect, and heavily ornamented with spicules; vestibulum with medial plates sclerotized and readily observed; small, nearly symmetrical, triangular; with sclerotized guide structure present as caplike structure with internal differentiation.
Etymology
Named for the genus Verticordia (Myrtaceae), from which many of the available specimens were taken.
Hosts
The majority of available specimens were taken from the following plant species: Verticordia chrysantha Endl. (pl. 8C, D), Verticordia densiflora Lindl., Verticordia polytricha Benth. (pl. 9B), and Pileanthus peduncularis borealis Endl. [ms. name] (pl. 7B) (Myrtaceae). Some additional taxa of Myrtaceae are also recorded as hosts for much smaller numbers of specimens. The record of 21 specimens from Allocasuarina (Casuarinaceae) does not represent a breeding host in our view, in the face of the hundreds of specimens known only from the Myrtaceae; this is probably the result of mislabeling or commingling of specimens in the field.
Distribution (map 6)
Relatively widespread in Western Australia, ranging from south of the Goldfields in the east to Kalbarri National Park in the north.
Holotype
AUSTRALIA: Western Australia: NW Coastal Hiway 57 km N of Kalbarri Road, 27.44756°S 114.6867°E, 500 m, 30 Oct 1996, Schuh and Cassis, Pileanthus peduncularis borealis Endl. (Myrtaceae), det. Perth 05120349, 1♂ (AMNH_PBI 00135409) (WAMP).
Paratypes
AUSTRALIA: Western Australia: 1 km S of Lillian Stoke Rock, 33.07681°S 120.0982°E, 380 m, 21 Nov 1999, R.T. Schuh and G. Cassis, Verticordia chrysantha Endl. (Myrtaceae), det. Perth 05672023, 1♂ (00368814) (AMNH). 11 km S of Eneabba, Eneabba National Park [96-50], 29.9025°S 115.24321°E, 150 m, 01 Nov 1996, Schuh and Cassis, Allocasuarina campestris E. Pritz. (Casuarinaceae), det. Perth 05095182, 15♂ (00372733–00372747) Calytrix glutinosa (Myrtaceae), 10♂ (00372349–00372358), 11♀ (00372359–00372369) Verticordia chrysanthella E. Pritz. (Myrtaceae), det. Perth 05095182, 3♂ (00372678–00372680) (AM), Verticordia chrysantha A.S. George (Myrtaceae), det. Field ID; Host 96-147, 14♂ (00135225–00135238), 11♀ (00135242–00135252) (AMNH), Verticordia chrysantha A.S. George (Myrtaceae), det. Field ID; Host 96-147, 3♂ (00135239–00135241), 8♀ (00135253–00135260) (WAMP). 20.6 km S of Norseman-Lake King Road on Lake King-Cascades Road, 33.16284°S 120.2813°E, 400 m, 22 Nov 1999, R.T. Schuh, G. Cassis, & R. Silveira, Verticordia chrysantha Endl. (Myrtaceae), det. Perth 05120160, 1♂ (00372417), 3♀ (00372418–00372420) (AM), 10♂ (00129607–00129612, 00129614–00129617), 10♀ (00129621, 00129623–00129629, 00129632, 00129635) (AMNH), 3♂ (00129618–00129620), 5♀ (00129630–00129631, 00129633–00129634, 00129636) (WAMP). 123 km W of Coolgardie on Great Eastern Hiway, 31.23414°S 120.1562°E, 17 Nov 1999, R.T. Schuh, G. Cassis, & R. Silveira, Verticordia chrysantha Endl. (Myrtaceae), det. Perth 05672015, 1♂ (00372421) (AM). Eneabba on Brand Hiway, 29.80735°S 115.2699°E, 100 m, 31 Oct 1996, Schuh and Cassis, Verticordia densiflora Lindl. (Myrtaceae), det. Perth 05120179, 48♀ (00372607–00372629, 00372645–00372669) Verticordia chrysantha Endl. (Myrtaceae), det. Perth 05120160, 9♂ (00372519–00372527), 23♀ (00372528–00372541, 00372551–00372559) (AM), 26♂ (00135069, 00135071–00135075, 00137425–00137444), 33♀ (00135078–00135082, 00135085–00135088, 00137461–00137484) (AMNH), Verticordia densiflora Lindl. (Myrtaceae), det. Perth 05120179, 16♀ (00372630–00372637, 00372670–00372677) (UNSW), 7♀ (00372638–00372644) Verticordia chrysantha Endl. (Myrtaceae), det. Perth 05120160, 17♂ (00137445–00137460, 00372379), 41♀ (00137485–00137502, 00137504–00137526) (WAMP). Kalbarri National Park, Loop Road, 27.56163°S 114.4376°E, 300 m, 28 Oct 1996, Schuh and Cassis, 29♀ (00372388–00372416) Verticordia polytricha Benth. (Myrtaceae), det. Perth 05120594, 15♂ (00372450–00372464), 47♀ (00372465–00372511) (AM), Melaleuca megacephala F. Muell. (Myrtaceae), det. Perth 05120616, 6♂ (00136709–00136713, 00136715), 6♀ (00136716–00136721) Pileanthus peduncularis borealis Endl. (Myrtaceae), det. Perth 05120586, 10♂ (00135472–00135480, 00135926), 12♀ (00135481–00135492) (AMNH), 13♂ (00372374–00372378, 00372380–00372387) Melaleuca megacephala F. Muell. (Myrtaceae), det. Perth 05120616, 7♀ (00136722–00136728) Pileanthus peduncularis borealis Endl. (Myrtaceae), det. Perth 05120586, 8♂ (00135920–00135925, 00135927–00135928), 10♀ (00135929–00135938) Verticordia polytricha Benth. (Myrtaceae), det. Perth 05120594, 7♀ (00372512–00372518) (WAMP). Kalbarri National Park, Z-Bend Road, 27.61971°S 114.3864°E, 500 m, 28 Oct 1996, Schuh and Cassis, Pileanthus peduncularis borealis Keighery ms. (Myrtaceae), det. Perth 05120586, 1♂ (00372570), 2♀ (00372571, 00372572) (AM). NW Coastal Hiway 57 km N of Kalbarri Road, 27.44756°S 114.6867°E, 500 m, 30 Oct 1996, Schuh and Cassis, Pileanthus peduncularis borealis Endl. (Myrtaceae), det. Perth 05120349, 3♂ (00372281–00372283), 10♀ (00372284–00372293) (AM), 14♂ (00135408, 00136748–00136760), 14♀ (00135410–00135416, 00136761–00136766, 00136774) (AMNH), 11♀ (00136767–00136773, 00136775–00136778) (WAMP). ca. 1 km S of Murchison House HS, Kalbarri National Park, 27.65822°S 114.2394°E, 60 m, 23 Oct 2004, Cassis, Wall, Weirauch, Symonds, Pileanthus vernicosus F. Muell. (Myrtaceae), det. Perth6988415, 4♂ (00368645–00368648), 12♀ (00368649–00368660) (AMNH).
Other Specimens Examined
AUSTRALIA: Western Australia: 1 km S of Lillian Stoke Rock, 33.07681°S 120.0982°E, 380 m, 21 Nov 1999, R.T. Schuh and G. Cassis, Verticordia chrysantha Endl. (Myrtaceae), det. Perth 05672023, 4♂ (00372542–00372545), 4♀ (00372547–00372550), 1 nymph (00372546) (AM), 35♂ (00368815–00368835, 00368840–00368853), 46♀ (00368776–00368797, 00368802–00368813, 00368854–00368865), 4 nymphs (00368866–00368869) (AMNH), 4♂ (00368836–00368839), 4♀ (00368798–00368801) (UCR). 3.5 km E of Lillian Stoke Rock, 33.07679°S 120.132°E, 360 m, 21 Nov 1999, R.T. Schuh, G. Cassis, & R. Silveira, Verticordia roei (Myrtaceae), 2♂ (00372370, 00372371), 2 nymphs (00372372, 00372373) (AM), 22♂ (00368754–00368775), 38♀ (00368716–00368753) (AMNH). 11 km S of Eneabba, Eneabba National Park [96-50], 29.9025°S 115.24321°E, 150 m, 01 Nov 1996, Schuh and Cassis, Verticordia chrysanthella E. Pritz. (Myrtaceae), det. Perth 05095182, 19♂ (00372681–00372699), 31♀ (00372700–00372730) (AM), Calytrix glutinosa Benth. (Myrtaceae), det. Perth 05120209, 26♂ (00135979–00136004), 23♀ (00136005–00136027), 1 nymph (00368898) (AMNH). 20.6 km S of Norseman-Lake King Road on Lake King-Cascades Road, 33.16284°S 120.2813°E, 400 m, 22 Nov 1999, R.T. Schuh, G. Cassis, & R. Silveira, Verticordia chrysantha Endl. (Myrtaceae), det. Perth 05672023, 1 nymph (00129622) (AMNH); 22 Nov 1999, R.T. Schuh and G. Cassis, 1♂ (00129613) (AMNH). 24 km W of Sandstone, 28.01426°S 119.0474°E, 650 m, 26 Oct 1996, Schuh and Cassis, Thryptomene aspera glabra E. Pritz. (Myrtaceae), det. Perth 05095182, 8♂ (00372579–00372586), 20♀ (00372587–00372606) (AM), 3♂ (00135633–00135635), 16♀ (00135639–00135654) (AMNH). 54.3 km N of jct of Agana Kalbarri Rd and Brand Hiway (rest area), 27.47362°S 114.7054°E, 240 m, 24 Oct 2004, Cassis, Wall, Weirauch, Symonds, Pileanthus vernicosus F. Muell. (Myrtaceae), det. Field ID, 9♂ (00368597–00368599, 00368870–00368875), 2♀ (00368876, 00368877), 20 nymphs (00368878–00368897) Verticordia capillaris A.S. George (Myrtaceae), det. Perth 6989861, 6♂ (00368673–00368678), 11♀ (00368679–00368689), 2 nymphs (00368690, 00368691) (AMNH). 123 km W of Coolgardie on Great Eastern Hiway, 31.23414°S 120.1562°E, 17 Nov 1999, R.T. Schuh, G. Cassis, & R. Silveira, Verticordia chrysantha Endl. (Myrtaceae), det. Perth 05672015, 7♂ (00372422–00372428), 21♀ (00372429–00372449) (AM). Brand Hiway 55.9 km S of Dongarra Road, 29.62934°S 115.2187°E, 100 m, 31 Oct 1996, Schuh and Cassis, Scholtzia drummondii Benth. (Myrtaceae), det. Perth 05120209, 2♂ (00088976, 00088979) (AM), 3♂ (00135450, 00135453–00135454) (AMNH). Eneabba on Brand Hiway, 29.80735°S 115.2699°E, 100 m, 31 Oct 1996, Schuh and Cassis, Verticordia chrysantha Endl. (Myrtaceae), det. Perth 05120160, 10 nymphs (00372560–00372569) (AM), 1 nymph (00135077) (AMNH). Kalbarri National Park, Z-Bend Road, 27.61971°S 114.3864°E, 500 m, 28 Oct 1996, Schuh and Cassis, Pileanthus peduncularis borealis Keighery ms. (Myrtaceae), det. Perth 05120586, 6 nymphs (00372573–00372578) (AM). ca. 1 km S of Murchison House HS, Kalbarri National Park, 27.65822°S 114.2394°E, 60 m, 23 Oct 2004, Cassis, Wall, Weirauch, Symonds, Pileanthus vernicosus F. Muell. (Myrtaceae), det. Perth 6988415, 12 nymphs (00368661–00368672) (AMNH). ca. 11 km S of Eneabba, Eneabba National Park, 29.90252°S 115.2432°E, 150 m, 01 Nov 1996, Schuh and Cassis, Allocasuarina campestris (Diels) L.A.S. Johnson (Casuarinaceae), det. Perth 05120063, 15♂ (00372733–00372747), 6♀ (00372748–00372753) (AM).
PHYLOGENETIC ANALYSIS
We prepared the matrix shown in appendix 1 as a way of creating a classification based on synapomorphy for the taxa treated in this paper. It contains 25 in-group taxa and 15 outgroups, mostly Australian, with a species of Leucophoropterini used as the root. The selection of outgroups among Australian Phylini was informed by the studies of Weirauch (2007) on the Polyozus Eyles and Schuh group of genera, Soto and Weirauch (2009) on Jiwarli, Weirauch and Schuh (2010) on the Xiphoidellus and related genera, and Schuh and Pedraza (2010) on Wallabicoris. Outgroups include Capecapsus tradouwensis Schuh from South Africa and the Australian taxa Wallabicoris ozothamni Schuh and Pedraza, W. pityrodii Schuh and Pedraza, W. spyridii Schuh and Pedraza, Xiphoides myersi (Woodward), Araucanophylus pacificus Carvalho, Xiphoidellus aureus Weirauch and Schuh, X. furvus Weirauch and Schuh, X. pallidus Weirauch and Schuh, Jiwarli heliotropium Soto and Weirauch, Polyozus galbanus Eyles and Schuh, Exocarpocoris tantulus Weirauch, and the manuscript taxon Protemiris conospermi.
Seventy-five characters are included, the majority related to morphology of the male and female genitalia, but characters derived from somatic morphology and color are also included. We have treated as additive those multistate characters for which we could hypothesize some credible transformation; all other characters are treated as nonadditive. Additivities are indicated in appendix 1 in the data matrix and the character descriptions. We used TNT (Goloboff et al., 2008, 2009) for our analyses, applying both equal weights and implied weights (Goloboff, 1993) approaches to character optimization.
Our concepts for male genitalic homology used the study of Weirauch (2007) as a baseline, because the genera Polyozus and Exocarpocoris Weirauch possess features in common with the Melaleucoides group of genera. Nonetheless, material examined for the present study showed structural variation that was not always easily interpreted under Weirauch's (2007) homology theories. Therefore, in arriving at the matrix presented in this paper, we employed a variety of alternative codings for the endosomal spines found apically and in association with the secondary gonopore.
Our concepts of female genitalic homology were informed by the studies of Weirauch (2007), Schuh and Pedraza (2010), and Weirauch and Schuh (2010). Genera included in those studies and members of the Melaleucoides group of genera possess morphological features of the posterior wall that are not seen outside of the Australian fauna. Nonetheless, the correlation between those features and the novel structures found in the males is obviously in need of additional study.
Our equal-weights analysis used ratchet and drift in addition to the default sectorial search with tree fusing. Default parameters for the driven search were used with the minimum length tree set to 10. The equal-weights analysis produced a total of 10 most parsimonious trees with a length (L) of 506, consistency index (CI) of 35, and a retention index (RI) of 62. The strict consensus of those trees is shown in figure 35.
Our implied weights analysis used the same settings as above with a constant of concavity of K = 3. Stronger constants produced essentially the same results. The single resultant tree, with a fit value of 36.982, is shown in figure 36.
As a way of indicating clade support we chose the jackknife as a measure. These values are plotted at the nodes in figures 35 and 36 for percentages over 50. They were obtained by running the jackknife resampling option in TNT with resampling probability = 36, 1000 replications and otherwise identical parameters as in the initial searches. Characters and character states were mapped on the resulting trees (figs. 35, 36) using WinClada and show unambiguous changes. Homoplasy settings indicate any extra step as rendering a character homoplastic.
Both the equal-weights and the implied-weights analyses recover with strong support the Melaleucoides genus group and the three genera described in this paper (figs. 35–Fig. 36.37). This grouping is supported by a number of characters, among which the fleshy structure of the parempodia (23-1) is easily recognized and apparently unique within the Australian Phylinae fauna. Harpgophylus is recognized as the sister group of the remaining members of the Melaleucoides genus group. Under our character codings none of the characters diagnostic for Harpgophylus is unique, but the monophyly of the group is nonetheless strongly supported in our analyses; the form of the right paramere (60-2) is easily recognized. The monophyletic group combining Thryptomenomiris and Melaleucoides is diagnosed by three characters, the most obvious of which is the vertical orientation of the left paramere (50-1). Under our character codings none of the characters diagnostic for Thryptomenomiris is unique, but the monophyly of the group is nonetheless strongly supported in our analyses; the form of the left paramere (56-2) is the most easily recognized. The monophyly of Melaleucoides is recognized by the left paramere extending vertically to the dorsal margin of the pygophore (53-1), a feature seen nowhere else in the Phylinae.
Although the form of the left paramere in Melaleucoides shows great consistency across the group, the structure of the endosoma is rather variable. Most species of Melaleucoides have two long apical spines and at least one additional spine associated with the secondary gonopore. Novel within the group is M. akaina, which has a single, short, apical spine and no processes associated with the secondary gonopore. Nonetheless, all of our phylogenetic analyses place M. akaina within Melaleucoides because of the structure of the parempodia and the left paramere.
The tree topologies resulting from our analyses differ with respect to certain species-level relationships within Harpagophylus and Melaleucoides. Among the clades within Melaleucoides recovered in both analyses are, e.g., a clade that comprises M. undulatae, M. rhaphiophyllae, and M. uncinatae and a clade that consists of M. beaufortiae, M. leuropomae, and M. micranthae.
HOST RELATIONSHIPS
All species in the Melaleucoides genus group are associated with plant species in the family Myrtaceae, subfamily Myrtoideae sensu Wilson et al. (2005) (table 2; fig. 37). The majority of plant bug species were recorded from one or very few host plant species (table 2). A noteworthy exception is M. verticordiae, the most commonly collected species in the genus group (976 specimens), which was recorded from 13 plant species comprising a wide range of Myrtaceae. Several plant species harbored more than one species of plant bug in the Melaleucoides genus group, among them Melaleuca rhaphiophylla, Melaleuca viminea, and Verticordia polytricha (table 2).
TABLE 2
Hosts of Melaleucoides genus group species
The Melaleucoides genus group appears to have a relatively clear-cut evolutionary pattern of host associations within the family Myrtaceae. According to Wilson et al. (2005), 15 tribes are currently recognized in this subfamily including the Melaleuceae (e.g., Melaleuca, Callistemon), Leptospermeae (e.g., Kunzea, Agonis), and Chamelaucieae (e.g., Verticordia, Thryptomene), all of which are speciose in Australia. The phylogenetic analyses by Wilson et al. (2001, 2005) demonstrated that Chamelaucieae and Leptospermeae are sister taxa and that Melaleuceae are more distantly related to that clade; species in the Melaleucoides genus group occur on species in the Chamelaucieae and Melaleuceae. Harpagophylus spp. and Thryptomenomiris spp. are restricted to Chamelaucieae and Melaleucoides spp. are known to be associated with both tribes. We mapped host genera on the implied weights tree and the following pattern appears: the association with species in the Chamelaucieae is the ancestral host association for the genus group and is seen in Harpagophylus, Thrpytomenomris, and the basal taxa within Melaleucoides (fig. 37) representing five species. Within the genus Melaleucoides a host switch occurred to the Melaleuceae, specifically species of Melaleuca and Beaufortia, and the 12 remaining species of Melaleucoides for which host plants are known are restricted to these two genera.
Distributional Patterns
Twenty-three of the 25 species placed in the Melaleucoides genus group are known only from a limited region in southwestern Western Australia; these distributions are shown in maps 1–6. The other two are known from the southeastern costal region of South Australia, as shown in maps 3 and 4. One might conclude that this distribution is the result of sampling bias. On the contrary, G. Cassis, R.T. Schuh, and their colleagues have collected over a wide area of Australia, using techniques similar to those applied in the capture of members of the Melalecoides genus group. And they have captured many other species of Phylinae breeding on species of Myrtoideae as part of those efforts. Thus, we conclude that these restrictions to the western part of the continent are not an artifact, but represent a real aspect of the distribution of this lineage.
Because we do not have a broader sampling of phylogenetic relationships for Australian Phylinae, it is not possible to specify the distribution of the sister group of the Melaleucoides genus group. Nonetheless, the distribution of the Melaleucoides group species has great similarity with distributions seen in monophyletic species groups within the recently described genus Wallabicoris (Schuh and Pedraza, 2010). No meaningful biogeographic statements can be made on the basis of two presumed areas of endemism as seen in the Melalecoides genus group (SW, SE). Correspondence with broader biogeographic theories will have to await improved knowledge of phylogenetic relationships within the Australian Phylinae.
Acknowledgments
We offer special thanks to Gerasimos Cassis, University of New South Wales, for his contribution to the existing collections in the Melaleucoides group of genera. His efforts in the field are to a very great degree responsible for bringing together the material studied during the course of this project. Other persons whose contributions to this project require individual acknowledgment because of their direct bearing on its completion for publication include: Steve Thurston (AMNH), who prepared the habitus images, inked and digitized the genitalic illustrations, and assembled all of the artwork in its final form; Kate Carmody, who entered specimen data and prepared most of the measurements; Christine A. Johnson, American Museum of Natural History (AMNH), who prepared some of the specimen measurements; Matthew Frankel of the AMNH Microscopy and Imaging Facility, who assisted with scanning electron microscopy; and the collections staffs at the Western Australian Herbarium, Department of Conservation and Land Management, Perth, and the Royal Botanic Gardens, Sydney, who provided authoritative host identifications.
This project represents a contribution to, and was partially supported by, the National Science Foundation (NSF) Planetary Biodiversity Inventories (PBI) award (DEB-0316495) to Randall T. Schuh and Gerasimos Cassis for the study of the Miridae subfamilies Orthotylinae and Phylinae.
Funding for fieldwork was provided by the Australian Museum (Western Australia, 1996, 1999; South Australia, 1998), the American Museum of Natural History (Western Australia, 1996, 1999; South Australia, 1998), the National Geographic Society (Western Australia, 1997), and the NSF PBI award (Western Australia, 2004).
All material collected by R.T. Schuh, G. Cassis, and their collaborators was acquired under permits issued by the Department of Environment and Heritage in South Australia and Department of Conservation and Land Management in Western Australia (1996, License No. SF001992; 1997, License No. SF002354; 1999, License No. NE002337). We thank these agencies for their cooperation.
For the loan of specimens we thank the following institutions and their respective collections management personnel: David Britton (Australian Museum, Sydney) and Jan Forrest (South Australian Museum, Adelaide).
We thank the Willi Hennig Society for supporting the phylogenetics program T.N.T. and for making that software freely available for our use in the preparation of this paper. Our thanks also to the authors, P.A. Goloboff, J.S. Farris, and K.C. Nixon, for their continuing contributions to the field of phylogenetic analysis.
For comments on the manuscript we thank Thomas J. Henry, Michael D. Schwartz, and Alfred G. Wheeler, Jr.
REFERENCES
Appendices
APPENDIX 1
Character Descriptions
0. Body: Greatly elongate, parallel sided = 0; moderately elongate, parallel sided = 1; weakly elongate, weakly ovoid = 2; weakly to distinctly ovoid = 3; broadly ovoid = 4. [additive]
COLORATION
1. Head: Uniformly dark = 0; pale with dark clypeus = 1; mostly dark with pale gula and posterior head margin = 2; pale or pale with red or dark dots = 3; dark with pale mark at medial eye margin = 4; Pale with red suffusion on gena = 5; Uniformly pale = 6; Pale, clypeus dark and large dark bilateral markings on frons = 7. [nonadditive]
2. Scapus: Unicolorous pale = 0; yellow to dirty yellow = 1; brown = 2; pale with a dark base = 3; unicolorous dark = 4. [nonadditive]
3. Pedicellus: Unicolorous pale = 0; yellow to dirty yellow = 1; dirty yellow proximally, weakly to heavily infuscate distally = 2; unicolorous black or castaneous = 3; black at extreme base, remainder pale = 4. [nonadditive]
4. Labium: Uniformly pale = 0; pale with segment 4 heavily infuscate = 1; unicolorous red = 2; generally infuscate, heavily so apically = 3. [nonadditive]
5. Thorax: Pronotum: Unicolorous deep red to black = 0; unicolorous pale = 1; unicolorous (yellow, brown, or green) = 2; pale with brown spots = 3; pale with red or carmine blotches = 4; pale with orange tinge, anterior lobe cream with dark marks = 5; pale with castaneous cali and humeri = 6; usually with dark anterior lobe and contrasting posterior lobe = 7. [nonadditive]
6. Scutellum: Unicolorous with remainder of dorsum = 0; with some red or carmine spots or blotches on pale background = 1; entirely red or orange = 2; red or dark brown with white or pale tip = 3; white = 4; entirely dark = 5; pale with brown spots = 6. [nonadditive]
7. Hemelytron: Unicolorous, deep red to black = 0; castaneous to black with contrasting white markings = 1; unicolorous [pale, white, or yellow] = 2; mostly pale, or yellow, endocorium with dirty or golden areas = 3; unicolorous pale with small brown spots = 4; with red or carmine spots, botches, or solid areas = 5; pale yellow with red suffusion = 6; brown with white cuneus = 7; with pattern of yellow, brown, and white, cuneus white = 8; pale with longitudinal castaneous stripes = 9. [nonadditive]
8. Markings on cuneus: Absent = 0; present as partial or complete but weak infuscation = 1; present as a contrasting white basal fascia = 2; present as pale narrow band, red over remaining area = 3; present as a red or castaneous apex = 4; present as small dark dots = 5; absent, cuneus transparent = 6. [nonadditive]
9. Membrane: Unicolorous pale = 0; partially fumose = 1; weakly to strongly fumose over entire area = 2. [nonadditive]
10. Membrane veins: White or pale = 0; yellow = 1; entirely red = 2; dark or dark red = 3; pale with distal portion of anterior cell red = 4; unicolorous with membrane = 5. [nonadditive]
11. Hind femur: Unicolorous pale, without black spots = 0; with a few brown or black spots = 1; with many brown or black spots = 2; nearly unicolorous dark = 3; pale with distal red suffusion = 4. [nonadditive]
12. Hind tibial spines: Dark = 0; pale = 1.
13. Hind tibial spines (bases): With conspicuous dark spots at bases = 0; with very small dark bases = 1; without dark spots at bases = 2. [nonadditive]
14. Abdominal venter: Unicolorous pale or mostly so = 0; light green = 1; light yellow = 2; heavily red laterally = 3; heavily infuscate, at least on mesopleuron and abdomen = 4; with thorax pale and abdomen dark = 5; unicolorous dark = 6. [nonadditive]
SURFACE AND VESTITURE
15. Dorsal common setae: With erect, suberect, or reclining black setae = 0; with black setae intermixed with pale setae on hemelytra = 1; with reclining simple setae matching background coloration = 2; with subadpressed flattened and erect simple pale setae = 3; with simple pale suberect setae = 4; with golden reclining setae = 5. [nonadditive]
16. Dorsal woolly setae: Without sericeous or woolly setae = 0; with some sericeous or woolly setae = 1.
STRUCTURE
17. Head: Elongate, projecting anteriorly = 0; moderately projecting = 1; somewhat projecting = 2; weakly projecting = 3; barely projecting, = 4. [nonadditive]
18. Interocular space: Relatively small = 1; moderate = 2; relatively large = 3; large = 4. [additive]
19. Eyes: Leaving gena only very slightly exposed in lateral view = 0; leaving gena moderately exposed in lateral view = 1; leaving gena broadly exposed in lateral view = 2. [additive]
20. Antenna: Segment 2: weakly tapering, more slender at base = 0; of equal diameter over entire length = 1.
21. Antennal fossa: With dorsal margin somewhat below ventral margin of eye = 0; with ventral margin at ventral margin of eye = 1; with ventral margin 1 diameter above ventral margin of eye = 2; with ventral margin 2 diameters above ventral margin of eye = 3. [nonadditive]
22. Labium: Reaching to posterior margin of mesosternum = 0; just reaching onto abdomen = 1; reaching to about anterior margin of pygophore = 2. [additive]
23. Paremopodia: Setiform = 0; fleshy, recurved. lyriform = 1.
24. Pulvilli: Absent = 0; present, flaplike, covering about one third of ventral claw surface = 1; present, flaplike, covering most of ventral surface of claw = 2. [additive]
25. Pygophore: Without row or tuft of setae on left side = 0; with dense tuft of setae on left side = 1; with double row of setae on left side = 2. [nonadditive]
26. Endosoma base: Very long = 0; moderately long = 1; short = 2. [nonadditive]
27. Endosoma shape: Curving, often C- or J-shaped = 0; with a U-shaped bend = 1; forming a single complete coil = 2. [additive]
28. Endosoma body: With torsion, primary strap dorsal to secondary gonopore = 0; without torsion, primary strap ventral to secondary gonopore = 1.
29. Primary endosomal strap apically: Short, ratio length apex strap/length secondary gonopore 1.33–1.50 = 0; weakly elongate, length apex strap/length secondary gonopore 1.67–2.00 = 1; elongate, ratio length apex strap/length secondary gonopore 2.15–2.83 = 2; greatly elongate, length apex strap/length secondary gonopore 3.00–8.6 = 3. [additive]
30. Primary endosoma strap apically (2): At least weakly arcuate and curving ventrally = 0; at least weakly arcuate and curving dorsally, no ornamentation = 1; at least weakly arcuate and curving dorsally, denticulate margin = 2; at least weakly arcuate and curving dorsally, comblike margin = 3. [additive]
31. Secondary endosomal strap: Broad, about equal in width to primary strap = 0; very slender, of uniform width from vesical bend to gonopore = 1; fused with primary strap = 2; absent = 3. [additive]
32. Secondary endosoma strap (2): Reaching midway to gonopore from major bend in vesica = 0; reaching to level of secondary gonopore = 1; reaching just beyond secondary gonopore as fingerlike extension = 2; reaching well beyond gonopore = 3. [nonadditive]
33. Secondary endosomal strap (3): With one tip = 0; anchor shaped = 1.
34. Secondary endosomal strap (4): Devoid of ornamentation = 0; with denticles = 1; with a comblike edge = 2. [additive]
35. Fingerlike protuberance at distal margin of secondary gonopore: Absent = 0; present = 1.
36. Spinelike, elongate process arising near gonopore on dorsal surface: Absent = 0; present, erect, unornamented = 1; present, erect, ornamented with denticles = 2. [additive]
37. Spinelike, elongate process near gonopore on ventral surface: Absent = 0; present and erect = 1; present, recurved toward base of endosoma = 2. [nonadditive]
38. Bladderlike process distad of secondary gonopore: Absent = 0; present = 1.
39. Secondary gonopore: Seen laterally in lateral view of vesica = 0; seen frontally (facing up) in lateral view of vesica = 1.
40. Secondary gonopore (2): Without a denticulate ridge on right-hand surface of endosoma = 0; with a denticulate ridge on right-hand surface of endosoma = 1.
41. Secondary gonopore sclerite (1): Absent = 0; present = 1; absent, but short, slender sclerite close to gonopore = 2. [nonadditive]
42. Secondary gonopore sclerite (2): Separated from secondary endosomal strap at apex = 0; apparently fused to secondary endosomal strap at apex = 1.
43. Phallotheca: Smoothly curving on dorsal margin = 0; more or less right angulate = 1; nearly erect = 2. [nonadditive]
44. Dorsal surface of phallotheca: Without a fingerlike projection = 0; with a fingerlike projection = 1.
45. Dorsal surface of phallotheca (2): Without a platelike projection = 0; with a curving, posteriorly directed, platelike projection = 1.
46. Anterior surface of phallotheca: Without a keel = 0; with a short to elongate keel = 1.
47. Ventral surface of phallotheca: Without a projecting keel = 0; with a projecting keel = 1.
48. Posterior surface of phallotheca: Lacking transparent window = 0; with conspicuous transparent window = 1.
49. Apex of phallotheca: Simple = 0; complex = 1.
50. Left Paramere: Horizontal = 0; vertical = 1.
51. Left paramere shaft: At right angles to body = 0; in same axis as body = 1.
52. Left paramere in dorsal perspective: Not exceeding margin of pygophore = 0; just exceeding margin of pygophore = 1; somewhat exceeding pygophore margin = 2; greatly exceeding margin of pygophore = 3. [nonadditive]
53. Left paramere in lateral perspective: Covering limited extent of lateral surface of pygophore = 0; erect and reaching dorsal margin of pygophore or beyond = 1.
54. Left paramere body: Narrowed toward apex = 0; spoon shaped = 1; more or less quadrate = 2. [nonadditive]
55. Left paramere apex medially: Angulate = 0; broadly rounded to weakly quadrate = 1; drawn into a blunt fingerlike process = 2; with an elongate, acuminate, recurved medial process = 3. [nonadditive]
56. Left paramere apex posterolaterally: Simple = 0; with a low shoulder = 1; with a greatly elongate projection = 2. [additive]
57. Left paramere anterior process: Slender and cylindrical in lateral view = 0; triangular in lateral view = 1; broad and elongate in lateral view = 2; undeveloped = 3. [nonadditive]
58. Left paramere posterior process: In the form of a short, straight, fingerlike projection = 0; in the form of a long, straight, fingerlike projection = 1; in the form of a short curved fingerlike projection = 2; very long, curving 90° toward base of paramere = 3. [nonadditive]
59. Left paramere base of posterior process: Without conspicuous shoulder = 0; with slightly projecting shoulder = 1; with distinct shoulder = 2. [nonadditive]
60. Right paramere body: Short and broad, lanceolate = 0; moderately elongate = 1; greatly elongate, bulbous at base = 2. [nonadditive]
61. Right paramere apex: Short, clawlike, decurved = 0; tapered and acuminate = 1; with short fingerlike process = 2; with a very long fingerlike process = 3. [nonadditive]
62. Right paramere posterior margin of apex: Smooth = 0; with distinct serration = 1.
FEMALE
63. Body shape: As in male = 0; slightly more ovoid than male = 1; distinctly more ovoid than male = 2. [additive]
64. Coloration: As in male = 0; more vividly red than male = 1; as in male but with cuneus bright red = 2; paler than male = 3. [nonadditive]
65. Posterior wall laterally: Without interramal sclerites = 0; with distinct crescent-shaped interramal sclerites = 1.
66. Posterior wall posteriorly: Without a sclerotized transverse band = 0; with a sclerotized transverse band = 1.
67. Posterior wall posterolaterally: Without a distinct swelling covered with microtrichia = 0; with a distinct swelling covered with microtrichia = 1.
68. Posterior wall longitudinal midline: Undifferentiated = 0; fold on either side of midline along part of length = 1; fold on either side of midline along most of length = 2. [additive]
69. Posterior wall posterior margin: Without spicules = 0; ornamented with spicules = 1.
70. Posterial wall posterior margin (2): Not reflexed dorsally = 0; reflexed dorsally = 1.
71. Posterior wall interramal lobes: Absent = 0; present and roughly symmetrical, erect = 1; present and asymmetrical, reclining, overlapping = 2. [additive]
72. Posterior wall interramal lobes (2): Not ornamented with spicules or with a very few = 0; heavily ornamented with spicules = 1.
73. Vestibulum medial plates (1): Sclerotized and readily observed = 0; weakly sclerotized or apparently absent = 1.
74. Vestibulum medial plates (2): Small, nearly symmetrical, triangular = 0; large, nearly symmetrical = 1; right plate larger, more strongly elongate than left = 2. [nonadditive]
75. Vestibulum medial plates sclerotized guide structure: Absent = 0; present as short weakly sclerotized medial tube = 1; present as caplike structure with internal differentiation = 2; present as short, heavily sclerotized tube on left side = 3; present as long tube on left side doubling back = 4; present as lateral sclerotizations = 5. [nonadditive]
Plates
Plate 1
Habitus images of Harpagophylus spp., Thryptomenomiris spp., and Melaleucoides akaina, and M. annae.
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.
Plate 6
Host plants of Melaleucoides genus group taxa. A.–C. Melaleuca sheathiana W. Fitz. D. Melaleuca uncinata R. Br. E. Melaleuca undulata Benth.
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.
