Wallabicoris waitzii, Schuh & P. Pedraza, 2010

Schuh, R. T. & P. Pedraza, 2010, Wallabicoris, New Genus (Hemiptera: Miridae: Phylinae: Phylini) From Australia, With The Description Of 37 New Species And An Analysis Of Host Associations, Bulletin of the American Museum of Natural History 2010 (338), pp. 1-118 : 87-101

Wallabicoris waitzii , new species Figures 47, 48; map 3; plate 5

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. sandstonensis in the elongate left paramere strongly projecting beyond the margin of the pygophore, but coloration of dorsum in W. sandstonensis not so uniformly and intensely yellow and the body not nearly as long and slender. Distinguished from W. chrysocephali , W. maralinga , and W. rutidosi by 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. waitzii not 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

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,

(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, Waitzia Wendl. ( Asteraceae : Gnaphalieae ).

HOSTS: Recorded only from Waitzia acuminata var. acuminata Steetz (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 acuminata includes much of the drier areas of interior Australia south of about 22 ° south latitude. The known occurrence of W. waitzii at 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 acuminata var. acuminata Steetz (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. acuminata Steetz (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 acuminata Steetz (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 acuminata var. acuminata Steetz (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 Wallabicoris spp. and seven outgroup taxa

0 5 101520253035404550

| | | | | | | | | | |

Leucophoropterini sp. 00010041234303330-201230320200-0200000010000000000140

Hypseloecus sp. 0000- 001234302230022404132011 -0---2020010000000000120

Pilophorus discretus -00-00002350-3330-22413231011-1---201100---0001000110

Tuxedo cruralis 10010040234403330-20213031010--0200010010000101000110

Plagiognathus chrysanthemi 21020001200304120100023120020200301000010000110100010

near Xiphoides ‘‘furvus’’ 2000-041134010101220223200200321100100012000110000130

near Xiphoides ‘‘pallidus’’ 11020000010000301220114110200321100100012000110000130

Wallabicoris baldersoni 01020001000000100221021021110101120010111011110111111

Wallabicoris cassisi 21020000000000110111013202100001110010120111111111111

Wallabicoris chrysocephali 02020001202101100221223121100101111010110001011010121

Wallabicoris commoni 01030010100000110221130020110101120010130011111110021

Wallabicoris coolabah 020300112041021002200231211101 -1110010111001110120111

Wallabicoris craspedii 01030011204202320110012121110201120010120101110111101

Wallabicoris cuneotinctus 24152231202101120011222010100311131010130101010220121

Wallabicoris dicrastyli 21020001000000120121222220110101120010130112110100021

Wallabicoris ellae 14150020000100120111121011110201100011110111010220121

Wallabicoris gingera 01020001204202320110021021100211111011120101111011111

Wallabicoris halganii 13340021111102120011123121110101111010132001111120111

Wallabicoris helichrysii 11030000101000120221431221100111120010122010111221121

Wallabicoris lachnostachyos 31020000000000110121114201100121110010130102111101101

Wallabicoris maralinga 12030010202010100221031021100221120011121011111020111

Wallabicoris newcastelii 21020001100012120121214111100111120011130112110100021

Wallabicoris norsemanius 02030001204000120211121021100121110011131002111120111

Wallabicoris olearii 11020000000010110221322020100201120010132012101001101

Wallabicoris ozothamni 11030000100000110-21221021100121120010120101110000111

Wallabicoris paradicrastyli 31030001100000110121214210100121111010111011111220121

Wallabicoris pimelei 21030001102101120111213000100221100010110001111110011

Wallabicoris pinocchii 01020000000000100200131020100311120011121211011220111

Wallabicoris pityrodiellus 11040020201000120121123021110111120010130211111111111

Wallabicoris pityrodii 11030011201101120111022110100101100011131022110110111

Wallabicoris pomaderri 220300111011011102214220201002210-0010122101110121111

Wallabicoris prostantheri 01020000001101100211223121100221111011122101010121021

Wallabicoris pultenaei 22051331121101120111014211100121131010111011111110101

Wallabicoris rhamnicola 241523312000001201012121201001110-1010110000110110111

Wallabicoris rutidosi 12030010100010100221121021100111140011120001111010111

Wallabicoris sandstonensis 12030011202101100220013120110211120010132001110110011

Wallabicoris schwartzi 01020000100101110221221020100211120011122011111101111

Wallabicoris spyridiellus 24152221200000120111323100100211131011130201010220011

Wallabicoris spyridii 242512012031011201112131101002210-1010111001110120111

Wallabicoris tasmanensis 12030001201101120100223121100211020011132002111101111

Wallabicoris thomasii 24152331103101220011124200100221131010131111110220011

Wallabicoris trymalii 24152331200000120101313001100211130010120201010120121

Wallabicoris uptoni 02030011204101300220021121100101120011122002111120111

Wallabicoris waitzii 02020001102101100220031021100211010011122101111210111

acuminata acuminata Steetz (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 Wallabicoris spp. and seven outgroup taxa were coded for 53 morphological characters. Two of the outgroup taxa, species from southern Australia related to the New Zealand genus 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 AT and 16S R CTC CGG TTT GAA CTC AGA TCA are 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 Wallabicoris and 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 Wallabicoris with 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 Wallabicoris spp , resulted in a single most parsimonious tree, 1223 steps long (fig. 51).

All three analyses recognize a monophyletic Wallabicoris , with the new genus near Xiphoides as 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 dicrastyli clade. 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. cassisi is excluded from the dicrastyli clade and placed instead within the cuneotinctus clade.

2. The cuneotinctus clade. 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 ozothamni clade: This large clade, including the ellae and rutidosi clades 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 ellae clade: The species pair W. ellae + W. pinocchii is 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 ellae clade is included within the ozothmni group, this

TABLE 3

Description of morphological characters for Wallabicoris spp. 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 –4.714 5 3; barely projecting, ratio wh/lh 5 4.884 –5.172 5 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 –0.416 5 1; moderate, ratio iod/wh 5 0.424 –0.457 5 2; relatively small, ratio iod/wh 5 0.476 –0.509 5 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

association is not recovered in absence of morphological data ( Fig. 49).

5. The rutidosi clade: The species pair W. rutidosi + W. helichrysi is recovered in all analyses (also including W. maralinga in 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 Wallabicoris spp.

HOST-PLANT ASSOCIATIONS

The evolution of host-plant associations in

Wallabicoris offers a tantalizing subject for exploration. The known host associations of Wallabicoris spp. 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. spyridii in association with Eutaxia microphylla J. Black represent 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.

As pointed out in the introduction to this paper, the host associations of Wallabicoris are 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 Sterculiaceae genera on which Wallabicoris spp. have been collected (table 5) are placed within Malvaceae by 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 Wallabicoris spp.

The nonrandom nature of host associations and restricted nature of diversification in Wallabicoris receives 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 Wallabicoris belong 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 Lamiaceae have iridoids, as the compound is lacking in Asteraceae and 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 dicrastyli clade (fig. 52, clade 1) is restricted to members of Lamiaceae (6 genera) and the host genus Dicrastylis J. 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 Australia with a single widespread species, W. paradicrastyli , which also occurs in Central Australia (map 1). Wallabicoris paradicrastyli has three host species, but all in the genus Dicrastylis . Although more restricted in geographical distribution, W. lachnostachyos and W. newcasteli have hosts from two lamiaceaous genera, while W. cassisii was found breeding on three genera of Lamiaceae . These are not the only Lamiaceae feeders 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 Lamiaceae probably evolved multiple times independently within Wallabicoris .

The cuneotinctus clade (fig. 52, clade 2) includes W. pimele i, 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. spyridii and W. trymalii , but the morphological data strongly support the placement of these species within the group. Rhamnaceae as a host-plant family appears to have evolved independently on more than one occasion, as W. pomaderri , not included in the cuneotinctus clade (based on morphological data only), also feeds on the Rhamnaceae .

All species of Wallabicoris known to feed on the Asteraceae are in the remaining grouping that we refer to as the ozothamni clade (fig. 52, clade 3), which is broadly distributed in Australia (map 3). Not all species placed in this group feed on the Asteraceae and eight of the species have unknown host associations. Nonetheless, it is likely that all species with unknown host associations are Asteraceae feeders. In terms of Asteraceae host 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 Olearia Moench. , which belongs to the Astereae . Exceptions to the rule of Asteraceae feeding 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 pinocchii has four host spp. all of them within Pimelea Banks ex Gaertn. (table 5).

Further evidence for the nonrandom nature of host associations in Wallabicoris can be deduced from the very large faunas of phyline Miridae that we have collected from other plant taxa that show great diversity in Australia, but from which no specimens of Wallabicoris have ever been recovered. Notable among these are the families Casuar-

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Fig. 51. Relationships of 37 Wallabicoris spp. 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

Wallabicoris Insect Plant Family Plant Species Species Specs. Events State Status

Asteraceae Chrysocephalum apiculatum Steetz W. chrysocephali 5 1 NT breeding Asteraceae Chrysocephalum apiculatum / W. chrysocephali 24 1 NT breeding semipapposum complex

Asteraceae Craspedia sp. W. craspedii 3 1 NSW breeding Asteraceae Helichrysum sp. W. helichrysi 49 1 NSW breeding Asteraceae Olearia axillaris (DC.) Benth. W. olearii 5 1 WA breeding Asteraceae Ozothamnus argophyllus W. schwartzi 4 1 Tasmania breeding (A. Cunn. ex DC) Anderb.

Asteraceae Ozothamnus diosmifolius DC. W. ozothamni 342 5 NSW breeding Asteraceae Ozothamnus ferrugineus Labill. Sweet W. schwartzi 29 2 Tasmania breeding Asteraceae Ozothamnus ferrugineus W. tasmanensis 27 1 Tasmania breeding Asteraceae Ozothamnus hookeri Sond. W. tasmanensis 54 2 Tasmania breeding Asteraceae Ozothamnus rosmarinifolius DC. W. schwartzi 77 1 Tasmania breeding Asteraceae Ozothamnus rosmarinifolius W. tasmanensis 1 1 Tasmania breeding Asteraceae Rutidosis helichrysoides DC. W. rutidosi 392 5 NT breeding Asteraceae Waitzia acuminata var. acuminata W. waitzii 130 1 WA breeding Asteraceae Waitzia acuminata var. acuminata W. waitzii 491 1 WA breeding Boraginaceae Halgania viscosa S. Moore W. halganii 28 1 WA breeding Fabaceae Eutaxia microphylla J. Black W. spyridii 3 1 SA sitting/ mislabeled

Fabaceae Pultenaea tenuifolia R. Br. W. pultenaei 292 2 Victoria breeding Lamiaceae Cyanostegia angustifolia Turcz. W. pityrodii 43 1 WA breeding Lamiaceae Dicrastylis beveridgei F. Muell. W. paradicrastyli 14 1 NT breeding Lamiaceae Dicrastylis beveridgei var. beveridgei W. paradicrastyli 17 1 NT breeding Lamiaceae Dicrastylis flexuosa (W.R. Price) W. dicrastyli 185 1 WA breeding C.A. Gardner

Lamiaceae Dicrastylis fulva fo. fulva W. cassisi 5 1 WA breeding Lamiaceae Dicrastylis fulva fo. fulva W. lachnostachyos 8 1 WA breeding Lamiaceae Dicrastylis fulva fo. fulva W. newcastelii 75 1 WA breeding Lamiaceae Dicrastylis gilesii var. gilesii W. paradicrastyli 13 2 NT breeding Lamiaceae Dicrastylis morrisonii Munir W. paradicrastyli 9 1 WA breeding Lamiaceae Dicrastylis parvifolia F. Muell. W. newcastelii 9 1 WA breeding Lamiaceae Lachnostachys coolgardiensis W. cassisi 1 1 WA breeding S. Moore

Lamiaceae Lachnostachys coolgardiensis W. lachnostachyos 5 1 WA breeding Lamiaceae Lachnostachys eriobotrya (F. Muell.) W. lachnostachyos 297 3 WA breeding Druce

Lamiaceae Lachnostachys eriobotrya W. lachnostachyos 29 1 WA breeding Lamiaceae Newcastelia insignis E. Pritz. W. cassisi 104 1 WA breeding Lamiaceae Newcastelia insignis W. newcastelii 76 1 WA breeding Lamiaceae Newcastelia insignis W. pityrodii 1 1 WA breeding Lamiaceae Newcastelia viscida E. Pritz. W. newcastelii 31 1 WA breeding Lamiaceae Pityrodia cuneata (Gaudich.) Benth. W. pityrodiellus 66 1 WA breeding Lamiaceae Pityrodia terminalis A.S. George W. pityrodii 353 3 WA breeding Lamiaceae Prostanthera campbellii F. Muell. W. prostantheri 18 1 WA breeding Lamiaceae Prostanthera sp. W. pityrodiellus 4 1 WA breeding Lauraceae Cassytha racemosa Nees W. pinocchii 48 1 WA mislabeled Loranthaceae Amyema preissii Tiegh. W. rutidosi 2 1 NT sitting/ mislabeled

Myrtaceae Calytrix tetragona Labill. W. spyridii 1 1 SA sitting/ mislabeled

Rhamnaceae Pomaderris apetala Labill. W. cuneotinctus 6 1 Tasmania breeding

TABLE 5 (Continued)

inaceae, Fabaceae (Mimosoideae) , Scrophulariaceae (5 Myoporaceae ), Myrtaceae , and Proteaceae .

DISTRIBUTIONAL PATTERNS

As currently known, most—although not all— Wallabicoris spp. 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 Wallabicoris species on the basis of their host associations. These distributions can be seen in maps 1–4 (pp. 16–17). The Lamiaceae feeders 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 Lamiaceae feeders belong to a monophyletic group, but the Lamiaceae-feeding di- crastyli group is nonetheless retrieved as monophyletic in most analyses.

The Rhamnaceae-feeding species, including all members of the cuneotinctus group, 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 type of 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 Miridae in Australia.

Species of Wallabicoris known to feed on the Asteraceae (map 3) show limited diversity in Western Australia. Even including some of those species that we predict to be Asteraceae feeders ( W. baldersoni , W. maralinga , W. norsemanius , W. sandstonensis , W. uptoni ), diversity in far southwestern Australia does not approach the diversity seen in the Lamiaceae- and Rhamnaceae-feeding clades. Seen from a somewhat different perspective, diversity in the interior of Australia is skewed toward the Asteraceae-feeding species, with only W. paradicrastyli (of the dicrastyli clade) found in central Australia.