Wallabicoris, New Genus (Hemiptera: Miridae: Phylinae: Phylini) From Australia, With The Description Of 37 New Species And An Analysis Of Host Associations
Schuh, R. T.
P. Pedraza
Bulletin of the American Museum of Natural History
2010
2010-06-03
2010
338
1
118
7G6NN
Schuh & P. Pedraza, 2010
Schuh & P. Pedraza
2010
[201,429,1246,1269]
Insecta
Miridae
Wallabicoris
Animalia
Hemiptera
86
87
Arthropoda
species
waitzii
sp. nov.
DIAGNOSIS: Recognized among those species with heavily yellow coloration by the intensely and uniformly yellow dorsum (pl. 5), by the elongate, slender body, mean total length 5.59, mean ratio length/width 3.80, the head conspicuously projecting beyond the anterior margin of the eyes, ratio head length/head width 2.70, the left paramere greatly elongate and strongly projecting beyond margin of pygophore (fig. 47E, F), and the secondary endosomal strap with a short, shallow submedial undulation (fig. 48). Breeds on Waitzia acuminata(Asteraceae). Similar to W. sandstonensisin the elongate left paramere strongly projecting beyond the margin of the pygophore, but coloration of dorsum in W. sandstonensisnot so uniformly and intensely yellow and the body not nearly as long and slender. Distinguished from W. chrysocephali, W. maralinga, and W. rutidosiby the much shorter, more deep-bodied left paramere in those species only moderately projecting beyond the margin of the pygophore. Potentially confused with W. commoni, but yellowish coloration in that species much less intense than in W. waitzii, secondary endosomal strap with a short medial undulation rather than the broad subbasal undulation seen in W. commoni, and left paramere in W. waitziinot nearly so deep bodied as in W. commoni.
DESCRIPTION: Male: Body greatly elongate, parallel sided, total length 5.27–5.85, l/w 5 3.80. COLORATION (pl. 5): Pronotum unicolorous yellow, calli weakly brown; hemelytron, including cuneus, unicolorous yellow; membrane with elongate, contrasting, fumose marking at extreme base, remainder partially fumose, especially within cells, veins white; venter light yellow; antennal segment 1 yellow to dirty yellow, with one or two black medial setae; antennal segment 2 yellow to dirty yellow; labium pale with segment 4 heavily infuscate; hind femora unicolorous pale, without black spots; hind tibial spines dark without dark spots at bases. SURFACE AND VESTITURE: Dorsum with reclining simple setae matching background coloration, without sericeous or woolly setae. STRUCTURE: Head:Elongate, projecting anteriorly, ratio wh/lh 5 2.70 (fig. 47A); antennal fossa with ventral margin 2 diameters above ventral margin of eye; interocular space moderately large, ratio iod/wh 5 0.38; eyes occupying entire height of head in lateral view (hg0–1/he20); labium just reaching to midpoint of abdomen. GENITALIA (fig. 47, E, F, 48): Endosoma:Base moderately long, with an open U-shaped bend; distal half of shaft smoothly curving; primary endosomal strap elongate apically, ratio lae/lsg 2.50; apex of primary endosomal strap weakly arcuate; secondary endosomal strap very slender, of uniform width from endosomal bend to gonopore, reaching midway to gonopore from major bend in endosoma, with a short submedial undulation; secondary gonopore seen laterally in lateral view of Fig. 47. Wallabicoris waitzii, male, scanning electron micrographs. A.Head in lateral view. B.Setae on corium adjacent to claval suture. C.Mesothoracic spiracle and metathoracic scent-efferent system. D.Pretarsus in medial view. E.Pygophore in lateral view. F.Detail of left paramere and apex of pygophore, dorsal view (AMNH_PBI 00136324). endosoma. Phallotheca:More or less right angulate, dorsal surface without a hump; dorsal surface with a conspicuous keel. Left Paramere:Body elongate, somewhat exceeding pygophore margin; in dorsal perspective closed over about three-fourths length; body tapered toward apex in lateral perspective; anterior process arising at posterior margin of shaft; anterior process angled posterodorsally; posterior process appearing tubular, Fig. 48. Wallabicoris waitzii, male genitalia (AMNH_PBI 00194143). fingerlike; posterior process with strongly projecting shoulder at base; base of posterior process at about level of paramere body or very weakly elevated. Right Paramere:Body elongate and slender, ratio lrp/wrp 4.12; body tapered from base to apex; posterior margin without distinct protuberance subapically; body elevated at juncture with base; apex with short fingerlike process.
ETYMOLOGY: Named for the host genus, WaitziaWendl.( Asteraceae: Gnaphalieae).
HOSTS: Recorded only from Waitzia acuminatavar. acuminataSteetz (Asteraceae)(pl. 7D, E).
DISTRIBUTION (map 3): Known only from Frank Hann National Park, Lilian Stoke Rock, southern Western Australia.
DISCUSSION: The actual distribution of Waitzia acuminataincludes much of the drier areas of interior Australiasouth of about 22 °southlatitude. The known occurrence of W. waitziiat a single site may be an artifact of sampling, but only further collecting on annual composites will provide the answer to this question.
HOLOTYPE: AUSTRALIA: Western Australia:Lillian Stoke Rock, 33.06784 °S 120.0964 °E, 400 m, 21 Nov 1999, R.T. Schuh and G. Cassis, Waitzia acuminatavar. acuminataSteetz (Asteraceae), det. PERTH staff PERTH 05670101, 13 (AMNH_PBI 00088680) ( WAMP). PARATYPES: AUSTRALIA: Western Australia:Frank Hann National Park, Lillian Stoke Rock, 33.06773 °S 120.0971 °E, 400 m, 05 Nov 1996, Schuh and Cassis, Waitzia acuminata var. acuminataSteetz (Asteraceae), det. PERTH staff PERTH 05236819, 643 (00087154–00087155, 00087443, 00087559, 00088503–00088548, 00088583, 00088657– 00088669), 48♀(00088584–00088592, 00088651– 00088652, 00089993–00090025, 00090039– 00090042) ( AM), 933 (00130397, 00131- 970–00131988, 00136252, 00136287–00136- 299, 00136300–00136318–00136330, 001727- 86–00172808, 00194143–00194147) 86♀(0013- 0398–00130401, 00131990–00132012, 001363- 32–00136360, 00172749–00172753–00172778) ( AMNH), 43 (00088631–00088634), 4♀(00088575–00088578) ( ANIC), 43 (00136279– 00136282), 4♀(00136400–00136403) ( CNC), 203 (00088639–00088650, 00088797–00088800, 00089987–00089990), 16♀(00088653–00088656, 00088670–00088674, 00088801–00088807) ( UNSW), 43 (00088635–00088638), 4♀(00- 088579–00088582) ( USNM), 533 (00088- 604–00088630, 00136253–00136278), 65♀(00- 088549–00088574, 00136361–00136399) ( WAMP), 43 (00136283–00136286), 4♀(00136404–0013- 6407) ( ZISP). Lillian Stoke Rock, 33.06784 °S 120.0964 °E, 400 m, 21 Nov 1999, R.T. Schuh and G. Cassis, Waitzia acuminata acuminataSteetz (Asteraceae), det. PERTH staff PERTH 05670101, 323 (00088676–00088679, 0008- 8681, 00088683–00088688, 00088796, 0008- 9936–00089955), 79♀(00088689–00088705, 00088707–00088712, 00088740–00088795) ( AM), 93 (00088682, 00172809–00172816), 8♀(00088706, 00172779–00172785) ( AMNH).
OTHER SPECIMENS EXAMINED: AUSTRA- LIA: Western Australia:Frank Hann National Park, Lillian Stoke Rock, 33.06773 °S 120.0971 °E, 400 m, 05 Nov 1996, Schuh and Cassis, Waitzia acuminatavar. acuminataSteetz (Asteraceae), det. PERTH staff PERTH 05236819, 12 nymphs (00088593– 00088603, 00089991) (AM), 2 nymphs (00131989, 00136331) (AMNH). Lillian Stoke Rock, 33.06784 °S 120.0964 °E, 400 m, 21 Nov 1999, R.T. Schuh and G. Cassis, Waitzia TABLE 2 Matrix of 53 morphological characters for 37 Wallabicorisspp.and seven outgroup taxa 0 5 101520253035404550 | | | | | | | | | | | Leucophoropterinisp. 00010041234303330-201230320200-0200000010000000000140 Hypseloecussp.0000- 001234302230022404132011-0---2020010000000000120 Pilophorusdiscretus-00-00002350-3330-22413231011-1---201100---0001000110 Tuxedocruralis10010040234403330-20213031010--0200010010000101000110 Plagiognathuschrysanthemi21020001200304120100023120020200301000010000110100010 near Xiphoides‘‘furvus’’ 2000-041134010101220223200200321100100012000110000130 near Xiphoides‘‘pallidus’’ 11020000010000301220114110200321100100012000110000130 Wallabicoris baldersoni01020001000000100221021021110101120010111011110111111 Wallabicoris cassisi21020000000000110111013202100001110010120111111111111 Wallabicoris chrysocephali02020001202101100221223121100101111010110001011010121 Wallabicoris commoni01030010100000110221130020110101120010130011111110021 Wallabicoris coolabah 020300112041021002200231211101-1110010111001110120111 Wallabicoris craspedii01030011204202320110012121110201120010120101110111101 Wallabicoris cuneotinctus24152231202101120011222010100311131010130101010220121 Wallabicoris dicrastyli21020001000000120121222220110101120010130112110100021 Wallabicoris ellae14150020000100120111121011110201100011110111010220121 Wallabicoris gingera01020001204202320110021021100211111011120101111011111 Wallabicoris halganii13340021111102120011123121110101111010132001111120111 Wallabicoris helichrysii11030000101000120221431221100111120010122010111221121 Wallabicoris lachnostachyos31020000000000110121114201100121110010130102111101101 Wallabicoris maralinga12030010202010100221031021100221120011121011111020111 Wallabicoris newcastelii21020001100012120121214111100111120011130112110100021 Wallabicoris norsemanius02030001204000120211121021100121110011131002111120111 Wallabicoris olearii11020000000010110221322020100201120010132012101001101 Wallabicoris ozothamni11030000100000110-21221021100121120010120101110000111 Wallabicoris paradicrastyli31030001100000110121214210100121111010111011111220121 Wallabicoris pimelei21030001102101120111213000100221100010110001111110011 Wallabicoris pinocchii01020000000000100200131020100311120011121211011220111 Wallabicoris pityrodiellus11040020201000120121123021110111120010130211111111111 Wallabicoris pityrodii11030011201101120111022110100101100011131022110110111 Wallabicoris pomaderri220300111011011102214220201002210-0010122101110121111 Wallabicoris prostantheri01020000001101100211223121100221111011122101010121021 Wallabicoris pultenaei22051331121101120111014211100121131010111011111110101 Wallabicoris rhamnicola241523312000001201012121201001110-1010110000110110111 Wallabicoris rutidosi12030010100010100221121021100111140011120001111010111 Wallabicoris sandstonensis12030011202101100220013120110211120010132001110110011 Wallabicoris schwartzi01020000100101110221221020100211120011122011111101111 Wallabicoris spyridiellus24152221200000120111323100100211131011130201010220011 Wallabicoris spyridii242512012031011201112131101002210-1010111001110120111 Wallabicoris tasmanensis12030001201101120100223121100211020011132002111101111 Wallabicoris thomasii24152331103101220011124200100221131010131111110220011 Wallabicoris trymalii24152331200000120101313001100211130010120201010120121 Wallabicoris uptoni02030011204101300220021121100101120011122002111120111 Wallabicoris waitzii02020001102101100220031021100211010011122101111210111 acuminata acuminataSteetz (Asteraceae), det. PERTH staff PERTH 05670101, 1 nymph (00088739) (AM). PHYLOGENETIC ANALYSES In order to test the monophyly of the group and to evaluate host and biogeographical relationships, we performed a phylogenetic analysis for Wallabicoris. Thirty-seven Wallabicorisspp.and seven outgroup taxa were coded for 53 morphological characters. Two of the outgroup taxa, species from southern Australiarelated to the New Zealandgenus Xiphoides, are being described by Weirauch and Schuh (in press). The morphological character matrix is shown in table 2. States for the morphometric characters were determined by the identification of breaks in the ratio ranges. Character descriptions are given in table 3. In an effort to produce a more robust character set for the group, we also sequenced,470 bases of 16S mtDNA for 25 taxa. Total genomic DNA was extracted from dried specimens using QIAGEN products. The large mitochondrial ribosomal subunit (16S rRNA) was amplified using the Illustra PuRe Taq Ready-to-Go PCR Beads. The primer pairs: 16S F CGC CTG TTT ATC AAA AAC ATand 16S R CTC CGG TTT GAA CTC AGA TCAare from Colgan et al. (1998). The annealing temperature of the PCR conditions was 48 °C. The PCR purification and cycle-sequencing were carried out with a Biomek NX Laboratory Automation Workstation and using the Gencourtẹ AM- Pureẹ and CleanSEQẹ systems. The reactions were sequenced using an automated Applied Biosystem 3730 DNA analyzer and the sequences were edited with Sequencher 4.8 (Gene Codes Corporation). The specimen voucher numbers and GenBank accession numbers for the 16S sequence data are given in table 4. Molecular and total evidence analyses were performed under the parsimony criterion using direct optimization ( Wheeler, 1996), as implemented in the program POY 4.1.1 ( Varón et al., 2010), to calculate dynamic homologies. The search strategy used in POY 4 is as follows, in sequential order: 1000 random addition sequences, SPR+TBR keeping one tree per search, select all. All characters were equally weighted, with gap opening 0. To assess nodal support, jackknife values were calculated in POY 4 using 1000 replicates, 1 random taxon entry, SPR+TBR, and saving 10 trees per replicate. Analyzing the molecular data alone for the 25 species of Wallabicorisand seven outgroups under the criteria specified above resulted in a single most parsimonious tree, 697 steps long (fig. 49). Combined analysis of 25 species of Wallabicoriswith corresponding molecular and morphological data produced two most parsimonious trees (L 5 1117); strict consensus tree shown in figure 50. A second total-evidence analysis including all 37 Wallabicorisspp, resulted in a single most parsimonious tree, 1223 steps long (fig. 51). All three analyses recognize a monophyletic Wallabicoris, with the new genus near Xiphoidesas its sister group, this result being recovered with 98% or greater jackknife values in each analysis. In addition to a monophyletic Wallabicoris, several additional clades were recovered in most analyses. These, identified by numbered nodes on figures 49–51, include: 1. The dicrastyliclade. This clade, comprising 5 species, is supported by characters 22-1 and 42-1 (fig. 51, clade 1), with a low to moderate jackknife value of 45%–73%. In the molecular only analysis W. cassisiis excluded from the dicrastyliclade and placed instead within the cuneotinctusclade. 2. The cuneotinctusclade. This clade comprises 7 species (fig. 51, node 2), with a jackknife value of 56%; a subset of six of those species, is supported by characters 1-4, 3-5, 5-2, 8-2, 34-1, and 48-2 with a jackknife value of 63%. 3. The ozothamniclade: This large clade, including the ellaeand rutidosiclades mentioned below, is of less consistent composition and topology across all analyses, but shows very little commingling with species from the two clades discussed above. 4. The ellaeclade: The species pair W. ellae+ W. pinocchiiis recovered in all analyses and it is supported by morphological characters 37-1, 44-0, 47-2, and 48-2 (fig. 51, clade 4), with a jackknife value of 54%. Although in the total evidence analyses (figs. 50, 51) the ellaeclade is included within the ozothmnigroup, this TABLE 3 Description of morphological characters for Wallabicorisspp.and outgroup taxa COLORATION 0. Body shape: greatly elongate, parallel sided (l/w 5 3.436 –3.961) 5 0; moderately elongate, parallel sided (l/w 5 3.115 –3.391) 5 1; weakly elongate, weakly ovoid (l/w 5 2.865 –3.091) 5 2; weakly to distinctly ovoid (l/w 5 2.727 –2.746) 5 3. 1. Coloration of pronotum: unicolorous, deep red to black 5 0; unicolorous pale 5 1; unicolorous [yellow, brown, or green] 5 2; pale with brown spots 5 3; pale with red or carmine botches 5 4. 2. Coloration of scutellum: unicolorous with remainder of dorsum 5 0; with some red or carmine spots or blotches on pale background 5 1; entirely red 5 2; pale with some brown spots 5 3. 3. Coloration of hemelytron: unicolorous, deep red to black 5 0; castaneous to black with contrasting white markings 5 1; unicolorous [pale, white, or yellow] 5 2; mostly pale, or yellow, endocorium with dirty or golden areas 5 3; unicolorous pale with small brown spots 5 4; with red or carmine spots, botches, or solid areas 5 5. 4. Red fascia on corium: absent 5 0; present on endocorium just anterior to cuneal fracture 5 1; present on exo- and endocorium just anterior to cuneal fracture 5 2. 5. Red markings on clavus: absent 5 0; present as spots only 5 1; present as some spots and a solid area at level apex scutellum 5 2; solid over most of area 5 3. 6. Markings on cuneus: absent 5 0; present as partial or complete but weak infuscation 5 1; present as individual or coalesced red or brown spots 5 2; pale basally, solid orange or red over most of remaining area 5 3; present as a contrasting white basal fascia 5 4. 7. Coloration of membrane base: with most of extreme base unicolorous with remainder 5 0; with elongate, contrasting, fumose marking at extreme base 5 1. 8. Coloration of membrane: unicolorous pale 5 0; partially fumose 5 1; weakly to strongly fumose over entire area 5 2. 9. Coloration of membrane veins: white 5 0; yellow 5 1; red 5 2; dark 5 3. 10. Coloration of venter: unicolorous pale or mostly so 5 0; light green 5 1; light yellow 5 2; heavily red laterally 5 3; heavily infuscate, at least on mesopleuron and abdomen 5 4; with thorax pale and abdomen dark 5 5. 11. Coloration of antennal segment 1: unicolorous pale 5 0; yellow to dirty yellow 5 1; dirty yellow, with strong infuscation at base and apex 5 2; pale with a dark base 5 3; black 5 4. 12. Antennal segment 1 coloration of medial seta: with one or two black medial setae 5 0; with pale medial seta 5 1. 13. Coloration of antennal segment 2: unicolorous pale 5 0; yellow to dirty yellow 5 1; dirty yellow proximally, weakly to heavily infuscate distally 5 2; black or castaneous 5 3; black at extreme base, remainder pale 5 4. 14. Coloration of labium:pale with segment 4 heavily infuscate 5 1; unicolorous red 5 2; generally infuscate, heavily so apically 5 3. 15. Coloration of hind femur: unicolorous pale, without black spots 5 0; with a few brown or black spots 5 1; with many brown or black spots 5 2; nearly unicolorous dark 5 3. 16. Coloration of hind tibial spines: dark 5 0; pale 5 1. 17. Coloration of bases of tibial spines: with conspicuous dark spots at bases 5 0; with very small dark bases 5 1; without dark spots at bases 5 2. SURFACE AND VESTITURE 18. Vestiture of dorsum(1): with erect or suberect black setae 5 0; with black setae intermixed with pale setae on hemelytra 5 1; with reclining simple setae matching background coloration 5 2. 19. Vestiture of dorsum(2): without sericeous or woolly setae 5 0; with some sericeous or woolly setae 5 1; present, lanceolate, and appressed, sometimes in rows 5 2. STRUCTURE 20. Head structure: elongate, projecting anteriorly, ratio wh/lh 5 2.47–2.77 5 0; moderately projecting, ratio wh/lh 5 3.00–3.335 5 1; somewhat projecting, ratio wh/lh 5 3.44–3.92 5 2; weakly projecting, ratio wh/lh 5 4.264 –4.7145 3; barely projecting, ratio wh/lh 5 4.884 –5.1725 4. 21. Position of antennal fossa: with dorsal margin somewhat below ventral margin of eye 5 0; with ventral margin of fossa at ventral margin of eye 5 1; with ventral margin 1 diameter above ventral margin of eye 5 2; with ventral margin 2 diameters above ventral margin of eye 5 3. 22. Interocular space: large, ratio iod/wh 5 0.321 5 0; moderately large, ratio iod/wh 5 0.358 –0.4165 1; moderate, ratio iod/wh 5 0.424 –0.4575 2; relatively small, ratio iod/wh 5 0.476 –0.5095 3; small, ratio iod/wh 5 0.52–0.56 5 4. 23. Eyes: occupying entire height of head in lateral view (hg0–1/he20) 5 0; leaving gena moderately exposed in lateral view (hg3/he20) 5 1; leaving gena broadly exposed in lateral view (hg5–14/he20) 5 2. 24. Labial length: very long, reaching well onto pygophore 5 0; reaching from abdominal midpoint to margin of pygophore 5 1; just reaching onto abdomen up to midpoint 5 2; reaching between fore- and middle trochanters 5 3. TABLE 3. (Continued) MALE GENITALIA 25. Base of endosoma length: base very long 5 0; base moderately long 5 1; base short 5 2. 26. Base of endosoma shape: weakly curving 5 0; with a tight U-shaped bend 5 1; forming a single complete coil 5 2. 27. Distal half of endosomal shaft: smoothly curving 5 0; more or less straight and erect 5 1; sinuous 5 2. 28. Straps of endosoma: with two straps 5 0; with a single unified strap, U-shaped in cross section 5 1. 29. Primary endosomal strap: short, ratio length apex strap/length sec gono 1.40–1.50 5 0; weakly elongate, ratio length apex strap/length sec gono 1.67–2.00 5 1; elongate, ratio length apex strap/length sec gono 2.15–2.83 5 2; greatly elongate, ratio length apex strap/length sec gono 3.00–5.5 5 3. 30. Apex of primary endosomal strap: nearly straight 5 0; weakly arcuate 5 1; distinctly curving 5 2. 31. Secondary endosomal strap width: broad, about equal in width to primary strap 5 0; very slender, of uniform width from endosomal bend to gonopore 5 1. 32. Secondary endosomal strap extent: reaching midway to gonopore from major bend in endosoma 5 0; reaching to level of secondary gonopore 5 1; reaching just beyond secondary gonopore as fingerlike extension 5 2; reaching well beyond gonopore as bladelike extension 5 3. 33. Secondary endosomal strap curvature: smoothly curving over entire length 5 0; with a short submedial undulation 5 1; with a broad submedial undulation 5 2; bifurcating at about midpoint 5 3; broken at about midpoint 5 4. 34. Secondary gonopore orientation: seen laterally in lateral view of endosoma 5 0; seen frontally in lateral view of endosoma 5 1; unsclerotized, orientation unclear 5 2. 35. Fingerlike protuberance at distal margin of secondary gonopore: absent 5 0; present 5 1. 36. Phallotheca shape: smoothly curving on dorsal margin 5 0; more or less right angulate 5 1; nearly erect 5 2. 37. Dorsal surface of phallotheca: without a conspicuous keel 5 0; with a conspicuous keel 5 1. 38. Posterior surface of phallotheca: lacking transparent window 5 0; with conspicuous transparent window 5 1. 39. Left paramere shape: body flattened, processes facing in opposite directions 5 0; body relatively short, just exceeding margin of pygophore 5 1; body elongate, somewhat exceeding pygophore margin 5 2; body very elongate, greatly exceeding margin of pygophore 5 3. 40. Left paramere in dorsal perspective: open over nearly entire length 5 0; closed over about one-half length 5 1; closed over about three-fourths length 5 2. 41. Left paramere body: not distinctly tapering toward apex 5 0; tapered toward apex in lateral perspective 5 1; tapered toward apex, apex elongate nipplelike 5 2. 42. Position of anterior process of left paramere: arising at posterior margin of shaft 5 0; arising slightly forward of posterior margin of shaft 5 1; arising near middle of anterodorsal margin of paramere5 2. 43. Direction of anterior process of left paramere: directed posteriorly 5 0; angled posterodorsally 5 1; directed vertically 5 2. 44. Posterior process of left paramere: flattened, broadest at angulate apex 5 0; appearing tubular, fingerlike 5 1. 45. Posterior process shoulder of left paramere: without strong shoulder at base 5 0; with strongly projecting shoulder at base 5 1. 46. Base of posterior process of left paramere: at least somewhat elevated above level of paramere body 5 0; at about level of paramere body or very weakly elevated 5 1. 47. Right paramere length: short and broad, ratio length/width 2.3–2.83 5 0; moderately elongate, ratio length/width 2.93–3.93 5 1; elongate and slender, ratio length/width 4.00–4.67 5 2. 48. Right paramere body shape: rounded laterally 5 0; tapered from base to apex 5 1; more or less parallel sided 5 2. 49. Right paramere posterior margin: without distinct protuberance subapically 5 0; with a distinct protuberance subapically 5 1. 50. Right paramere body elevation: more or less confluent with base 5 0; elevated at juncture with base 5 1. 51. Right paramere apex: tapered 5 0; with short fingerlike process 5 1; with a weakly elongate fingerlike process 5 2; with a very long fingerlike process 5 3; blunt 5 4. FEMALE GENITALIA 52. Posterior wall: simple 5 0; with strongly elevated longitudinal folding 5 1. TABLE 4 AMNH voucher and NCBI GenBank acession numbers for 16S mtDNA sequences Taxon AMNH voucher GenBank Accession Hypseloecussp. (Australia) AMNH_PBI 0023220 AY252897 Pilophorus discretusVan Duzee AMNH_PBI 00024105 AY252838 Leucophoropterinisp. (Australia) AMNH_PBI 00023260 HM142166 Tuxedo cruralis(Van Duzee) NA AY252823 Plagiognathus chrysanthemi AMNH_PBI 00023224 AY252709 near Xiphoides‘‘furvus’’ (Australia) AMNH_PBI 00131244 HM217005 near Xiphoides‘‘pallidus’’ (Australia) AMNH_PBI 00131056 HM142165 Wallabicoris cassisi AMNH_PBI 00133142 HM142141 Wallabicoris chrysocephali AMNH_PBI 00098811 HM142149 Wallabicoris cuneotinctus AMNH_PBI 00194224 HM142150 Wallabicoris dicrastyli AMNH_PBI 00135721 HM142151 Wallabicoris ellae AMNH_PBI 00133297 HM142152 Wallabicoris halganii AMNH_PBI 00136998 HM142153 Wallabicoris helichrysii AMNH_PBI 00089256 HM142154 Wallabicoris lachnostachyos AMNH_PBI 00136125 HM142155 Wallabicoris newcastelii AMNH_PBI 00090083 HM142143 Wallabicoris ozothamni AMNH_PBI 00090450 HM142144 Wallabicoris paradicrastyli AMNH_PBI 00097948 HM142146 Wallabicoris pimelei AMNH_PBI 00194159 HM142148 Wallabicoris pinocchii AMNH_PBI 00090539 HM142147 Wallabicoris pityrodii AMNH_PBI 00136974 HM142156 Wallabicoris pityrodiellus AMNH_PBI 00302770 HM142145 Wallabicoris prostantheri AMNH_PBI 00090063 HM142157 Wallabicoris pultenaei AMNH_PBI 00089440 HM142158 Wallabicoris rhamnicola AMNH_PBI 00172658 HM142159 Wallabicoris rudidosi AMNH_PBI 00097284 HM142160 Wallabicoris sandstonensis AMNH_PBI 00090049 HM142161 Wallabicoris schwartzi AMNH_PBI 00102026 HM142162 Wallabicoris spyridiellus AMNH_PBI 00087906 HM142163 Wallabicoris tasmanensis AMNH_PBI 00194082 HM142164 Wallabicoris thomasii AMNH_PBI 00128570 HM142142 Wallabicoris waitzii AMNH_PBI 00088703 HM217004 association is not recovered in absence of morphological data ( Fig. 49). 5. The rutidosiclade: The species pair W. rutidosi+ W. helichrysiis recovered in all analyses (also including W. maralingain the 37-taxon analysis; fig. 51, node 5), with jackknife values near 100% in the 25-taxon analyses (figs. 42, 50). We base the remainder of our discussion on figure 51, the total-evidence analysis of all 37 Wallabicorisspp. HOST-PLANT ASSOCIATIONS The evolution of host-plant associations in Wallabicorisoffers a tantalizing subject for exploration. The known host associations of Wallabicorisspp.are listed in the specimensexamined sections of the taxonomic treatments and are summarized for each species under the heading Hosts. Table 5 offers an alternative view of host associations, listing known hosts by family, genus and species, their associated bug predators, and the number of collecting events per bug species, and the number of bug specimens collected. The records for the plant families Rutaceae, Scrophularaceae, Loranthaceae, and Lauraceae, and for W. spyridiiin association with Eutaxia microphyllaJ. Blackrepresent apparent commingling of specimens in the field rather than actual host records; they involve singular collecting events and miniscule numbers of insect specimens. Thus, they are discarded from the analysis. Fig. 49. Relationships of 25 Wallabicorisspp.and seven outgroups based on analysis of 16S mDNA data using POY 4 with a 1:1:1:1:0 cost ratio (morphological change:transitions:transversions:gap extension:gap opening). Single tree of cost 697. Jackknife values above 50% shown. As pointed out in the introduction to this paper, the host associations of Wallabicorisare far from random, but rather appear to be restricted to just seven recognized plant families: Asteraceae, Boraginaceae, Faba- ceae, Lamiaceae, Rhamnaceae, Sterculiaceae, and Thymeleaceae. The Sterculiaceaegenera on which Wallabicorisspp.have been collected (table 5) are placed within Malvaceaeby the Angiosperm Phylogeny Group ( Stevens, 2001). Host associations can be appreciated by examination of figure 52 where host-plant families are plotted on the results of the total-evidence analysis of all 37 Wallabicorisspp. Fig. 50. Relationships of 25 Wallabicorisspp.and seven outgroups based on analysis of 16S mDNA data and 53 morphological characters using POY 4 with a 1:1:1:1:0 cost ratio, (morphological change:transitions:transversions:gap extension:gap opening). Consensus of two trees of cost 1117). Jackknife values above 50% shown. The nonrandom nature of host associations and restricted nature of diversification in Wallabicorisreceives substantial support from our phylogenetic analyses, as can be seen in figure 52 and from the following discussion. Although spread across seven eudicot families, all the known hosts of Wallabicorisbelong to two major clades, the Asterids ( Asteraceae, Lamiaceae, and Boraginaceae) and Rosids ( Rhamnaceae, Thymelaceae, Fabaceae, and Sterculiaceae). Iridoids are cyclopentanoid monoterpenes highly correlated with asterids, although not exclusively so ( Albach et al., 2001). These compounds have been implicated in herbivore preferences, detering some and attracting others. The evolution of iridoid compounds may, therefore, have had a selective advantage for the plants able to synthesize them; likewise, the same may be inferred from hervibores that feed from these plants despite the iridoids. Iridoids have a bitter taste and cause nausea in vertebrates, thus some herviborous insects sequester them to use in defense against predators ( Bowers 1980, 1988). However, among the asterids that are hosts of Wallabicoris, only those in the Lamiaceaehave iridoids, as the compound is lacking in Asteraceaeand Boraginaceae. Rosids are a large and heterogeneous group. Although the monophyly of rosids was corroborated by recent molecular studies, no clear nonmolecular apomorphies unite the rosid clade ( Angiosperm Phylogeny Group, 2003; Simpson, 2006). The rosid clade also contains novel biochemical pathways such as the machinery necessary for symbiosis with nitrogen-fixing bacteria (nitrogen-fixing clade) and defense mechanisms such as glucosinolate production (Brassicales) and cyanogenic glycosides (e.g., Fabaceae) (Solits et al., 2005). The dicrastyliclade (fig. 52, clade 1) is restricted to members of Lamiaceae(6 genera) and the host genus DicrastylisJ. Drumm. ex Harv.is common to the entire clade (table 5). The clade is composed of W. cassisi, W. dicrastyli, W. lachnostachyos, W. newcastelii, and W. paradicrastyli; it is found primarily in Western Australiawith a single widespread species, W. paradicrastyli, which also occurs in Central Australia(map 1). Wallabicoris paradicrastylihas three host species, but all in the genus Dicrastylis. Although more restricted in geographical distribution, W. lachnostachyosand W. newcastelihave hosts from two lamiaceaous genera, while W. cassisiiwas found breeding on three genera of Lamiaceae. These are not the only Lamiaceaefeeders within Wallabicoris, the others being W. pityrodii, W. pityrodiellus, and W. prostantheri. These last three species, however, do not group in our cladistic analysis (figs. 49–51); thus, the preference for Lamiaceaeprobably evolved multiple times independently within Wallabicoris. The cuneotinctusclade (fig. 52, clade 2) includes W. pimelei, W. cuneotinctus, W. rhamnicola, W. spyridiellus, W. spyridii, W. thomasii, and W. trymalii. All are restricted to the south coastal regions of western and eastern Australia(Map 2). They feed on members of Rhamnaceae, Sterculiaceae, and Thymeleaceae, all rosids. We did not have sequence data for W. spyridiiand W. trymalii, but the morphological data strongly support the placement of these species within the group. Rhamnaceaeas a host-plant family appears to have evolved independently on more than one occasion, as W. pomaderri, not included in the cuneotinctusclade (based on morphological data only), also feeds on the Rhamnaceae. All species of Wallabicorisknown to feed on the Asteraceaeare in the remaining grouping that we refer to as the ozothamniclade (fig. 52, clade 3), which is broadly distributed in Australia(map 3). Not all species placed in this group feed on the Asteraceaeand eight of the species have unknown host associations. Nonetheless, it is likely that all species with unknown host associations are Asteraceaefeeders. In terms of Asteraceaehost diversity, the number of genera is only slightly higher (7 genera) when compared with Lamiaceae, the most frequent host family of Wallabicoris. Six of those seven genera belong to the tribe Gnaphalieae, the only exception being OleariaMoench., which belongs to the Astereae. Exceptions to the rule of Asteraceaefeeding in this clade are W. halganii(Boraginaceae), W. pityrodiellus(Lamiaceae), W. ellae(Sterculiaceae), and W. pinocchii(Thymeliaceae). The last species is the most widely distributed of all known species of Wallabicoris, occurring along the eastern and western coasts of Australia, but with no known collections in the center of the continent (map. 4); Wallabicoris pinocchiihas four host spp. all of them within PimeleaBanks ex Gaertn.(table 5). Further evidence for the nonrandom nature of host associations in Wallabicoriscan be deduced from the very large faunas of phyline Miridaethat we have collected from other plant taxa that show great diversity in Australia, but from which no specimens of Wallabicorishave ever been recovered. Notable among these are the families Casuar- Fig. 52. Host associations of Wallabicorisspp.superimposed on phylogenetic relationships from figure 51. + Fig. 51. Relationships of 37 Wallabicorisspp.and seven outgroups based on analysis of 16S mDNA data and 53 morphological characters using POY 4 with a 1:1:1:1:0 cost ratio, (morphological change:transitions:transversions:gap extension:gap opening) Single tree of cost 1223. Jackknife values above 50% are shown. TABLE 5 Host plants of Wallabicoris WallabicorisInsect PlantFamily PlantSpecies Species Specs. Events State Status Asteraceae Chrysocephalum apiculatumSteetz W. chrysocephali5 1 NT breeding Asteraceae Chrysocephalum apiculatum/ W. chrysocephali24 1 NT breeding semipapposum complex Asteraceae Craspediasp. W. craspedii3 1 NSW breeding Asteraceae Helichrysumsp. W. helichrysi49 1 NSW breeding Asteraceae Olearia axillaris(DC.) Benth. W. olearii5 1 WA breeding Asteraceae Ozothamnus argophyllus W. schwartzi4 1 Tasmaniabreeding (A. Cunn. ex DC) Anderb. Asteraceae Ozothamnus diosmifoliusDC. W. ozothamni342 5 NSW breeding Asteraceae Ozothamnus ferrugineusLabill. Sweet W. schwartzi29 2 Tasmaniabreeding Asteraceae Ozothamnus ferrugineus W. tasmanensis27 1 Tasmaniabreeding Asteraceae Ozothamnus hookeriSond. W. tasmanensis54 2 Tasmaniabreeding Asteraceae Ozothamnus rosmarinifoliusDC. W. schwartzi77 1 Tasmaniabreeding Asteraceae Ozothamnus rosmarinifolius W. tasmanensis1 1 Tasmaniabreeding Asteraceae Rutidosis helichrysoidesDC. W. rutidosi392 5 NT breeding Asteraceae Waitzia acuminatavar. acuminata W. waitzii130 1 WA breeding Asteraceae Waitzia acuminatavar. acuminata W. waitzii491 1 WA breeding Boraginaceae Halgania viscosaS. Moore W. halganii28 1 WA breeding Fabaceae Eutaxia microphyllaJ. Black W. spyridii3 1 SA sitting/ mislabeled Fabaceae Pultenaea tenuifoliaR. Br. W. pultenaei292 2 Victoriabreeding Lamiaceae Cyanostegia angustifoliaTurcz. W. pityrodii43 1 WA breeding Lamiaceae Dicrastylis beveridgeiF. Muell. W. paradicrastyli14 1 NT breeding Lamiaceae Dicrastylis beveridgei var. beveridgei W. paradicrastyli17 1 NT breeding Lamiaceae Dicrastylis flexuosa(W.R. Price) W. dicrastyli185 1 WA breeding C.A. Gardner Lamiaceae Dicrastylis fulvafo. fulva W. cassisi5 1 WA breeding Lamiaceae Dicrastylis fulvafo. fulva W. lachnostachyos8 1 WA breeding Lamiaceae Dicrastylis fulvafo. fulva W. newcastelii75 1 WA breeding Lamiaceae Dicrastylis gilesiivar. gilesii W. paradicrastyli13 2 NT breeding Lamiaceae Dicrastylis morrisoniiMunir W. paradicrastyli9 1 WA breeding Lamiaceae Dicrastylis parvifoliaF. Muell. W. newcastelii9 1 WA breeding Lamiaceae Lachnostachys coolgardiensis W. cassisi1 1 WA breeding S. Moore Lamiaceae Lachnostachys coolgardiensis W. lachnostachyos5 1 WA breeding Lamiaceae Lachnostachys eriobotrya(F. Muell.) W. lachnostachyos297 3 WA breeding Druce Lamiaceae Lachnostachys eriobotrya W. lachnostachyos29 1 WA breeding Lamiaceae Newcastelia insignisE. Pritz. W. cassisi104 1 WA breeding Lamiaceae Newcastelia insignis W. newcastelii76 1 WA breeding Lamiaceae Newcastelia insignis W. pityrodii1 1 WA breeding Lamiaceae Newcastelia viscidaE. Pritz. W. newcastelii31 1 WA breeding Lamiaceae Pityrodia cuneata(Gaudich.) Benth. W. pityrodiellus66 1 WA breeding Lamiaceae Pityrodia terminalisA.S. George W. pityrodii353 3 WA breeding Lamiaceae Prostanthera campbelliiF. Muell. W. prostantheri18 1 WA breeding Lamiaceae Prostantherasp. W. pityrodiellus4 1 WA breeding Lauraceae Cassytha racemosaNees W. pinocchii48 1 WA mislabeled Loranthaceae Amyema preissiiTiegh. W. rutidosi2 1 NT sitting/ mislabeled Myrtaceae Calytrix tetragonaLabill. W. spyridii1 1 SA sitting/ mislabeled Rhamnaceae Pomaderris apetalaLabill. W. cuneotinctus6 1 Tasmaniabreeding TABLE 5 (Continued) Wallabicoris Insect Plant Family PlantSpecies Species Specs. Events State Status Rhamnaceae Pomaderris orariasubsp. calcicola W. pomaderri 26 1 Victoria breeding Rhamnaceae Pomaderris orariasubsp. oraria W. cuneotinctus 46 1 Victoria breeding Rhamnaceae Spyridium globulosumBenth. W. spyridiellus 98 4 WA breeding Rhamnaceae Spyridium globulosum W. thomasii 2 1 WA breeding Rhamnaceae Spyridiumsp. W. spyridii 221 1 SA breeding Rhamnaceae Spyridium vexilliferum(Hook.) W. rhamnicola 67 1 Victoria breeding Reissek Rhamnaceae Spyridium vexilliferum W. spyridii 1 1 Victoria breeding Rhamnaceae Trymalium odoratissimum Lindl. W. trymalii 4 1 WA breeding (5 T. floribundumSteud.fo. fulva) Rhamnaceae Trymalium odoratissimum Lindl. W. trymalii 5 1 WA breeding (5 T. floribundumSteud.subsp. trifidum) Rutaceae Chorilaena quercifoliaEndl. W. thomasii 4 1 WA sitting/ mislabeled Scrophulariaceae Eremophila gilesiiF. Muell. W. rutidosi 1 1 NT sitting/ mislabeled Sterculiaceae Keraudrenia integrifoliaHook. W. ellae 108 2 WA breeding Sterculiaceae Lasiopetalum floribundum W. thomasii 21 2 WA breeding Sterculiaceae Thomasiams name heterophylla W. thomasii 38 1 WA breeding Thymelaeaceae Pimelea glaucaR. Br. W. pinocchii 1 1 NSW breeding Thymelaeaceae Pimelea linifoliasubsp. collina W. pinocchii 18 1 NSW breeding Thymelaeaceae Pimelea longiflorasubsp. longiflora W. pinocchii 91 1 WA breeding Thymelaeaceae Pimelea sericeaR. Br. W. pimelei 25 1 Tasmania breeding Thymelaeaceae Pimeleasp. W. pinocchii 2 1 NSW breeding Thymelaeaceae Pimelea sylvestrisR. Br. W. pinocchii 79 3 WA breeding State Abbreviations:New South Wales 5NSW;Northern Territory 5 NT; South Australia 5 SA; Western Australia 5 WA. inaceae, Fabaceae (Mimosoideae), Scrophulariaceae(5 Myoporaceae), Myrtaceae, and Proteaceae. DISTRIBUTIONAL PATTERNS As currently known, most—although not all— Wallabicorisspp.show restricted patterns of distribution, many known from only a single collecting event or from a few geographically proximal localities. We have plotted the distributions of Wallabicorisspecieson the basis of their host associations. These distributions can be seen in maps 1–4 (pp. 16–17). The Lamiaceaefeeders are almost totally restricted to Western Australia, and more particularly to the Goldfields-Kalbarri region (map 1). Our phylogenetic analyses do not suggest that all Lamiaceaefeeders belong to a monophyletic group, but the Lamiaceae-feeding di- crastyligroup is nonetheless retrieved as monophyletic in most analyses. The Rhamnaceae-feeding species, including all members of the cuneotinctusgroup, are restricted to the most southerly coastal regions of Australia, and show an amphicontinental pattern (map 2), with the greatest diversity in Western Australia. This typeof disjunction and skewed level of diversity has been documented by Weirauch and Schuh (in press) and Schuh and Weirauch (submitted) in other genus-level groupings of phyline Miridaein Australia. Species of Wallabicorisknown to feed on the Asteraceae(map 3) show limited diversity in Western Australia. Even including some of those species that we predict to be Asteraceaefeeders ( W. baldersoni, W. maralinga, W. norsemanius, W. sandstonensis, W. uptoni), diversity in far southwestern Australiadoes not approach the diversity seen in the Lamiaceae- and Rhamnaceae-feeding clades. Seen from a somewhat different perspective, diversity in the interior of Australiais skewed toward the Asteraceae-feeding species, with only W. paradicrastyli(of the dicrastyliclade) found in central Australia.