Melaleucoides verticordiae, Schuh & C. Weirauch, 2010

Schuh, R. T. & C. Weirauch, 2010, Myrtaceae-Feeding Phylinae (Hemiptera: Miridae) From Australia: Description And Analysis Of Phylogenetic And Host Relationships For A Monophyletic Assemblage Of Three New Genera, Bulletin of the American Museum of Natural History 2010 (344), pp. 3-94 : 67-79

Melaleucoides verticordiae , new species Figures 33, 34; map 6; plate 4

DIAGNOSIS: Recognized by the yelloworange 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. SUR- FACE AND VESTITURE (fig. 33C): Dorsum with suberect or reclining black setae and some sericeous or woolly setae. STRUC- TURE: 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. GEN- ITALIA (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 Verti- species: Verticordia chrysantha Endl. (pl. 8C, cordia ( Myrtaceae ), from which many of the D), Verticordia densiflora Lindl. , Verticordia available specimens were taken. polytricha Benth. (pl. 9B), and Pileanthus

HOSTS: The majority of available speci- peduncularis borealis Endl. [ms. name] mens were taken from the following plant (pl. 7B) ( Myrtaceae ). Some additional taxa 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, 27.44756uS 114.6867uE, 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, 33.07681uS 120.0982uE, 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], 29.9025uS 115.24321uE, 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 U (00372359–00372369) Verticordia chrysanthella E. Pritz. (Myrtaceae) , det. Perth 05095182, 3 - (00372678–0037- 2680) ( AM), Verticordia chrysantha A.S. George (Myrtaceae) , det. Field ID; Host 96- 147, 14 - (00135225–00135238), 11 U (0013- 5242–00135252) ( AMNH), Verticordia chrysantha A.S. George (Myrtaceae) , det. Field ID; Host 96-147, 3 - (00135239–00135241), 8 U (00135253–00135260) ( WAMP). 20.6 km S of Norseman-Lake King Road on Lake King- Cascades Road, 33.16284uS 120.2813uE, 400 m, 22 Nov 1999, R.T. Schuh, G. Cassis, & R. Silveira, Verticordia chrysantha Endl. (Myrtaceae) , det. Perth 05120160, 1 - (00372417), 3 U (00372418–00372420) ( AM), 10 - (0012- 9607–00129612, 00129614–00129617), 10 U (00129621, 00129623–00129629, 00129632, 00129635) ( AMNH), 3 - (00129618–0012- 9620), 5 U (00129630–00129631, 00129633– 00129634, 00129636) ( WAMP). 123 km W of Coolgardie on Great Eastern Hiway, 31.23414uS 120.1562uE, 17 Nov 1999, R.T. Schuh, G. Cassis, & R. Silveira, Verticordia chrysantha Endl. (Myrtaceae) , det. Perth 05672015, 1 - (00372421) ( AM). Eneabba on Brand Hiway, 29.80735uS 115.2699uE, 100 m, 31 Oct 1996, Schuh and Cassis, Verticordia densiflora Lindl. (Myrtaceae) , det. Perth 05120179, 48 U (00372607– 00372629, 00372645–00372669) Verticordia chrysantha Endl. (Myrtaceae) , det. Perth 05120160, 9 - (00372519–00372527), 23 U (003- 72528–00372541, 00372551–00372559) ( AM), 26 - (00135069, 00135071–00135075, 00137- 425–00137444), 33 U (00135078–00135082, 00- 135085–00135088, 00137461–00137484) ( AMNH), Verticordia densiflora Lindl. (Myrtaceae) , det. Perth 05120179, 16 U (00372630–00372637, 00372670–00372677) ( UNSW), 7 U (00372- 638–00372644) Verticordia chrysantha Endl. (Myrtaceae) , det. Perth 05120160, 17 - (001- 37445–00137460, 00372379), 41 U (001374- 85–00137502, 00137504–00137526) ( WAMP). Kalbarri National Park, Loop Road, 27.56163uS 114.4376uE, 300 m, 28 Oct 1996, Schuh and Cassis, 29 U (00372388– 00372416) Verticordia polytricha Benth. (Myrtaceae) , det. Perth 05120594, 15 - (003- 72450–00372464), 47 U (00372465–00372511) ( AM), Melaleuca megacephala F. Muell. (Myrtaceae) , det. Perth 05120616, 6 - (0013- 6709–00136713, 00136715), 6 U (00136716– 00136721) Pileanthus peduncularis borealis Endl. (Myrtaceae) , det. Perth 05120586, 10 - (00135472–00135480, 00135926), 12 U (00135481–00135492) ( AMNH), 13 - (0037- 2374–00372378, 00372380–00372387) Melaleuca megacephala F. Muell. (Myrtaceae) , det. Perth 05120616, 7 U (00136722–00136- 728) Pileanthus peduncularis borealis Endl. (Myrtaceae) , det. Perth 05120586, 8 - (00135920–00135925, 00135927–00135928), 10 U (00135929–00135938) Verticordia polytricha Benth. (Myrtaceae) , det. Perth 05120594, 7 U (00372512–00372518) ( WAMP). Kalbarri National Park, Z-Bend Road, 27.61971uS 114.3864uE, 500 m, 28 Oct 1996, Schuh and Cassis, Pileanthus peduncularis borealis Keighery ms. ( Myrtaceae ), det. Perth 05120586, 1 - (00372570), 2 U (00372571, 00372572) ( AM). NW Coastal Hiway 57 km N of Kalbarri Road, 27.44756uS 114.6867uE, 500 m, 30 Oct 1996, Schuh and Cassis, Pileanthus peduncularis borealis Endl. (Myrtaceae) , det. Perth 05120349, 3 - (00372281– 00372283), 10 U (00372284–00372293) ( AM), 14 - (00135408, 00136748–00136760), 14 U (00135410–00135416, 00136761–00136766, 00136774) ( AMNH), 11 U (00136767–0013- 6773, 00136775–00136778) ( WAMP). ca. 1 km S of Murchison House HS, Kalbarri National Park, 27.65822uS 114.2394uE, 60 m, 23 Oct 2004, Cassis, Wall, Weirauch, Symonds, Pileanthus vernicosus F. Muell. (Myrtaceae) , det. Perth6988415, 4 - (00368645–00368648), 12 U (00368649–00368660) ( AMNH).

OTHER SPECIMENS EXAMINED: AUSTRA- LIA: Western Australia: 1 km S of Lillian Stoke Rock, 33.07681uS 120.0982uE, 380 m, 21 Nov 1999, R.T. Schuh and G. Cassis, Verticordia chrysantha Endl. (Myrtaceae) , det. Perth 05672023, 4 - (00372542–003- 72545), 4 U (00372547–00372550), 1 nymph (00372546) (AM), 35 - (00368815–003688- 35, 00368840–00368853), 46 U (00368776– 00368797, 00368802–00368813, 00368854– 00368865), 4 nymphs (00368866–00368869) (AMNH), 4 - (00368836–00368839), 4 U (00368798–00368801) (UCR). 3.5 km E of Lillian Stoke Rock, 33.07679uS 120.132uE, 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 U (00368716–00368753) (AMNH). 11 km S of Eneabba, Eneabba National Park [96-50], 29.9025uS 115.24321uE, 150 m, 01 Nov 1996, Schuh and Cassis, Verticordia chrysanthella E. Pritz. (Myrtaceae) , det. Perth 05095182, 19 - (00372681–00372699), 31 U (00372700– 00372730) (AM), Calytrix glutinosa Benth. (Myrtaceae) , det. Perth 05120209, 26 - (00135979–00136004), 23 U (00136005–0013- 6027), 1 nymph (00368898) (AMNH). 20.6 km S of Norseman-Lake King Road on Lake King-Cascades Road, 33.16284uS 120.2813uE, 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, 28.01426uS 119.0474uE, 650 m, 26 Oct 1996, Schuh and Cassis, Thryptomene aspera glabra E. Pritz. (Myrtaceae) , det. Perth 05095182, 8 - (00372579–00372586), 20 U (00372587– 00372606) (AM), 3 - (00135633–00135635), 16 U (00135639–00135654) (AMNH). 54.3 km N of jct of Agana Kalbarri Rd and Brand Hiway (rest area), 27.47362uS 114.7054uE, 240 m, 24 Oct 2004, Cassis, Wall, Weirauch, Symonds, Pileanthus vernicosus F. Muell. (Myrtaceae) , det. Field ID, 9 - (00368597– 00368599, 00368870–00368875), 2 U (0036- 8876, 00368877), 20 nymphs (00368878– 00368897) Verticordia capillaris A.S. George (Myrtaceae) , det. Perth 6989861, 6 - (0036- 8673–00368678), 11 U (00368679–00368689), 2 nymphs (00368690, 00368691) (AMNH). 123 km W of Coolgardie on Great Eastern Hiway, 31.23414uS 120.1562uE, 17 Nov 1999, R.T. Schuh, G. Cassis, & R. Silveira, Verticordia chrysantha Endl. (Myrtaceae) , det. Perth 05672015, 7 - (00372422–00372428), 21 U (00372429–00372449) (AM). Brand Hiway 55.9 km S of Dongarra Road, 29.62934uS 115.2187uE, 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, 29.80735uS 115.2699uE, 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, 27.61971uS 114.3864uE, 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, 27.65822uS 114.2394uE, 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, 29.90252uS 115.2432uE, 150 m, 01 Nov 1996, Schuh and Cassis, Allocasuarina campestris (Diels) L.A.S. Johnson (Casuarinaceae) , det. Perth 05120063, 15 - (00372733–00372747), 6 U (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 equalweights 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 5 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 5 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 impliedweights analyses recover with strong support the Melaleucoides genus group and the three genera described in this paper (figs. 35–37). 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

TABLE 2 Hosts of Melaleucoides genus group species

Host genus No. of

and species Specs. Locality Miridae species

Beaufortia micrantha 39 WA: Fitzgerald River National Park, Hammersley M. beaufortiae Road

Beaufortia micrantha 25 WA: 92.5 km W of Coolgardie at east side of M. micranthae micrantha Boorabbin National Park on Great Eastern Hwy

Calytrix glutinosa 71 WA: 11 km S of Eneabba, Eneabba National Park M. verticordiae Calytrix variabilis 10 WA: Kalbarri National Park, Meanarra Hill H. calyrtrix

(pl. 5A)

Darwinia diosmoides 8 WA: Kalbarri National Park, Loop Road T. yalgoo Malleostemon 1 WA: ca. 1 km S of Murchison House HS, T. kalbarri hursthousei Kalbarri National Park

Melaleuca brevifolia 57 SA: Scorpion Springs Cons. Park M. brevifoliae Melaleuca 7 WA: 15 km E of Merredin M. leuropomae ; conothamnoides 2 WA: 15 km E of Merredin M. similis Melaleuca laetifica 30 WA: Kalbarri National Park, 37.7 km E Kalbarri M. systenae

(pl. 5A)

Melaleuca leuropoma 29 WA: Kalbarri National Park, 7 km E of Kalbarri M. leuropomae

(pl. 5C)

Melaleuca 19 WA: Kalbarri National Park, Loop Road M. verticordiae megacephala

Melaleuca 11 WA: Brand Hiway 8.2 km N of Eneabba M. rhaphiophyllae rhaphiophylla (pl. 5D) 1 WA: Brand Hiway 8.2 km N of Eneabba M. uncinatae Melaleuca sheathiana 186 WA: 11 km N of Coolgardie-Esperance Hiway on M. annae

(pl. 6A–C)) Kambalda Road

100 WA: 11 km N of Coolgardie-Esperance Hiway on M. sheathianae Kambalda Road; 33.3 km S of Norseman

Melaleuca sp. 2 WA: Moorine Rocks, 11.7 km N of Great Eastern M. rhaphiophyllae Hiway on Noongar Road

Melaleuca sp. 2 WA: Frank Hann National Park, 37 km E of Lake M. systenae

King

Melaleuca systena 22 WA: 4.5 km S of Jurien on Indian Ocean Rd; M. systenae

WA: 8.2 km E of Indian Ocean Rd on

Coorow-Greenhead Rd, Lesuer National Park

Melaleuca teuthidoides 20 WA: 91.4 km SE of Southern Cross M. sheathianae Melaleuca uncinata 69 WA: 56.6 km W of Yalgoo M. uncinatae

(pl. 6D) 204 WA: 56.6 km W of Yalgoo M. cassisi Melaleuca undulata 44 WA: Kalbarri National Park, 22.9 km E Kalbarri M. undulatae

(pl. 6E) 4 WA: 4 km W of Lillian Stoke Rock M. castanea Melaleuca urceolaris 5 WA: 11 km S of Eneabba on Brand Hiway, M. systenae Eneabba Reserve

Melaleuca viminea 4 WA: Cervantes M. leuropomae ; M. systenae Micromyrtus 11 WA: NW Coastal Hiway 58 km N of Kalbarri M. akaina hursthousei (pl. 7A) Road

Phymatocarpus 5 WA: Brand Hiway 45.9 km S of Dongarra Road M. leuropomae ; porphyrocephalus 5 WA: Brand Hiway 45.9 km S of Dongarra Road M. systenae Pileanthus 102 WA: Kalbarri National Park, Loop Road; Kalbarri M. verticordiae peduncularis National Park, Z-Bend Road; NW Coastal Hiway

borealis (pl. 7B) 57 km N of Kalbarri Road

128 WA: Kalbarri National Park, Loop RoaD; NW M. pileanthicola Coastal Hiway 57 km N of Kalbarri Road

TABLE 2

(Continued)

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 specieslevel 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).

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 .