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Systematics and phylogeny of the hatchet head plant bug genus myrmecoroides gross (insecta: heteroptera: miridae: orthotylinae).

Abstract--Myrmecoroides Gross is revised and a new diagnosis is provided for the genus* The two previously recognized species, M. carinatus Gross and M. grossi Carvalho, are redescribed. Three new species are described: M. cassowary, M. monteithi, and M. rufescens. The male and female genitalia are documented and illustrated where specimens of each gender were available. Phylogenetic analysis of the species is given. The common name, Hatchet Head Plant Bug, is introduced for the genus, in recognition of the greatly bicompressed head.

Key words: Heteroptera, Miridae, Orthotylinae, Austromirini, ant mimicry, myrmecomorphy, new species, Australia.

INTRODUCTION

Myrmecomorphy is one of the more exaggerated phenotypes in arthropods. Batesian myrmecomorphs are well-known in the true bug suborder Heteroptera, probably more so than in any other higher taxon, rivaled only by the Arachnida (McIver and Stonedahl, 1993). Independent evolution of myrmecomorphy in true bugs is inferred by its occurrence in numerous true bug families (Enicocephalomorpha: Enicocephalidae; Leptopodomorpha: Saldidae; Cimicomorpha: Nabidae and Miridae; Pentatomomorpha: Geocoridae, Oxycarenidae, Rhyparochrornidae, Alydidae and Pyrrhocoridae--see McIver and Stonedahl, 1993).

Within the Heteroptera, myrmecomorphy is most pronounced in the Miridae, where it has presumably evolved independently within six of the eight plant bug subfamilies (Cylapinae, Bryocorinae, Deraeocorinae, Mirinae, Orthotylinae and Phylinae). This independent derivation of myrmecomorphy within plant bugs is consistent with Schuh's (1974, 1975, 1976) argument that ant mimicry alone is not safely regarded as an indicator of phylogenetic relationships.

Myrmecomorphy in plant bugs is most prevalent in Orthotylinae and Phylinae. Within the Orthotylinae, ant mimicry is characteristic of many species within the tribes Austromirini Carvalho and Ceratocapsini Knight (=Sericophanes group sensu Schuh, 1974). The Austromirini are a monophyletic tribe of Orthotylinae (Insecta: Heteroptera: Miridae) (Cassis, 2006, 2008), comprising non-myrmecomorphic and ant mimetic clades (Cassis, 2006). Among austromirine myrmecomorphic genera, species of the Australian endemic genus Myrmecoroides Gross are remarkable in terms of their antlike facies, with significant pronotal, forewing and abdominal modifications. In addition, Myrmecoroides has a highly autapomorphic head, which shows no obvious convergence toward any putative "ant model." The head (frons + clypeus) is exaggeratedly bicompressed, giving a hatchetlike appearance, hence the introduction here of the common name, Hatchet Head Plant Bug.

Myrmecoroides was first described by Gross (1964) for a single species, M. carinatus Gross, based on a small series of micropterous females from Victoria. He placed the genus within the Orthotylinae: Pilophorini. Schuh (1974) sub sequently placed Myrmecoroides, as incertae sedis, within the Orthotylinae, as part of his redefinition of suprageneric groupings within the Miridae, which included the placement of the Pilophorini within the Phylinae. Carvalho (1973) described a second species, M. grossi Carvalho, from New South Wales, but did not place the genus within the Orthotylinae. Cassis and Gross (1995) first placed Myrmecoroides in the orthotyline tribe Austromirini.

This work includes the description of three new species of Myrmecoroides from eastern Australia. In addition, a phylogenetic analysis of Myrmecoroides is undertaken and their biology is discussed.

MATERIALS AND METHODS

Materials

This study was based on an examination of 65 specimens from eastern Australia. Specimens were borrowed from the following collections: Australian Museum (AM), Australian National Insect Collection (ANIC), Entomological Collection of the University of Queensland (EUQ), Museum Victoria (MV), Queensland Museum (QM) and South Australian Museum (SAMA). Additional specimens are housed at the University of New South Wales Insect Collection (UNSWIC).

Morphometrics

All measurements are in millimeters and refer to maximal lengths. Abbreviations for morphometric characters are: BL = body length; HL = head length; HW = head width; IOD = interocular distance; AI = first antennal segment length; AII= second antennal segment length; AIII = third antennal segment length; PL = pronotal length; PW = width of posterior margin of pronotum; and, LL = labial length. The range, mean and standard deviation for each of these characters is given (Table 1).

Morphology

Morphological terminology follows that given by Cassis (1995) for external features and Cassis et al. (2003) for female genitalia. The bicompressed head, with the enlarged and flattened vertex and frons, is referred to as the frontal plate in this work.

Terminology for the male genitalia follows that given by Cassis (2008), in that the male phallus is composed of a basal articulatory apparatus (=phallobase), a distal membraneous inner tubule (=endosoma) bounding the ductus seminis, and the endosoma encased by an outer tubule (=phallotheca), which in turn is connected to the phallobase.

Cassis et al. (2003), Cassis and Eyles (2006) and Cassis (2008) rejected the homologies drawn by Kerzhner and Konstantinov (1999), particularly their division of the endosoma into a proximal conjunctiva (attached to the phallobase) and distal vesica. Cassis (2008) promoted the hypothesis that the inner tubule of the aedeagus be referred to as the endosoma, without further subdivision, as the notion of the vesica in the Miridae is non-homologous. The single spicule in Myrmecoroides is considered to be the dorsal endosomal spicule (=DES; sensu Cassis 2008) because of its more dorsal position relative to the secondary gonopore.

Microscopy

Light microscope observations were made using a Leica MZ16 stereomicroscope and a Leica DMB compound microscope. Scanning electron micrographs were prepared using techniques outlined in Bolte (1996) and Swearingen et al. (1997). Images were prepared using a Cambridge electron microscope.

Phylogeny

The data were analyzed using TNT software (Goloboff et al., 2008). All characters are binary and unweighted. The data were run using the "Traditional search" routine, with 500 replications. Three undescribed species of ant mimetic Austromirini (taxon codes: TASM001, PSEUD001 and AUSTRO001) were used as outgroups.

BIOLOGY

Little is known of the biology of the species of Myrmecoroides. We have recently discovered that M. grossi is found on grasses (Fig. 1). Both sexes and a nymph of this species were found on grass leaves, and the presence of the active life-stages provides evidence for grass association. Other specimens of this species, from independent collection events, have also been collected on grasses.

Grass specialization occurs independently in the Heteroptera a number of times (e.g., Alydidae: Leptocorisinae, Blissidae, Pachygronthidae, Pentatomidae: Aeptini; see Cassis and Gross, 2002), and within the Miridae, notably in the Stenodemini (Schwartz 1987), and many Orthotylinae: Halticini and the genus Lopus Hahn (Phylini). One of us [GC] has recently recorded a new nonmyrmecomorphic austromirine genus from Australia, Watarrkamiris Cassis, from grasses (Cassis, 2008). The fact that nearly all other species of Austromirini are found on dicot angiosperms (mostly rosid and asterid flowering plants) suggests that grass affiliation is likely a host switching event. The generality of grass associations for Myrmecoroides, is yet to be established. However, collection data suggest that such an hypothesis requires further exploration. For example, M. rufescens n.sp. has also been collected from (unspecified) grasses. Also, M. cassowary n.sp., was found in pitfall traps among a range of perennials, including grasses.

[FIGURE 1 OMITTED]

Habitat information for Myrmecoroides is very limited, with species found on woodland edges (M. grossi), among dune vegetation (M. cassowary n.sp.), and in cool temperate forests (M. monteithi n.sp.). Exploration of possible gender habitat specialization is required, as females are most commonly found on the ground, collected in pitfall or flight-intercept traps, and nearly all males are collected by sweeping vegetation. Only in a handful of collection events, and only for M. grossi and M. monteithi, have both sexes been found, both on the ground and on vegetation. In such cases, however, sex ratios are heavily biased towards the above putative gender habitat preferences. It is also noteworthy that all known females of the genus are micropterous, whereas all males are macropterous, which provides ecomorphological support for an hypothesis of gender-specific habitat specialization.

TAXONOMY

Myrmecoroides Gross, 1964

Myrmecoroides Gross, 1964:8 (nov. gen.); Carvalho, 1973:9 (diagnosis); Schuh, 1974:319 (taxonomic position); Cassis and Gross, 1995: 184 (transferred to Austromirini); Schuh, 1995: 146 (catalog)

TYPE SPECIES: Myrmecoroides carinatus Gross, by original designation.

DIAGNOSIS: Myrmecoroides is recognized by the following combination of characters: males macropterous (Figs. 8, 13); females micropterous (Figs. 7, 10); frons and clypeus strongly bicompressed, platelike, with hatchetlike appearance (Figs. 2a, b, 4); bucculae short, arcuate (Fig. 2b); labrum flat, triangular, not modified (Fig. 2c); labium reduced, slender (Fig. 2b), reaching base of mesocoxae; antennae and tibiae bicoloured; antennae slender, AI bottle-shaped; collar present; pronotum campanulate in males, callosite region restricted, margins parallel-sided, weakly hourglass shaped in females (Figs 7, 10); propleural suture anterodorsally directed; proepisternum rounded; proepimeron short; external efferent system of metathoracic glands moderately developed (Fig. 2e), reaching mesepimeron, peritreme short, tumid (Fig. 2e); metathoracic spiracle visible, bounded by evaporative bodies (Fig. 2e); costal margins of male forewings strongly concave; pretarsal parempodia lamellate, broad, recurved apically (Fig. 2d); pulvilli prominent (Fig. 2d); pretarsal claws weakly angulate apically (Fig. 2d); genital opening of pygophore small (Fig. 2g, h); parameres short, clublike (Figs. l lb, c, 13b, c, 15b, c); aedeagus simple, with one enlarged, elongate basal sclerite (Figs l ld, 13d, 15d), secondary gonopore simple (Figs l ld, 13d, 15d); female pregenital abdomen strongly constricted, basally, TII and TIII elongate, bilaterally compressed; laterotergites prominent; sclerotized rings elaborate, folded (Figs. 5, 12), and spiculate; inter-ramal lobes prominent, distally free, obovate (Fig. 5c, d) to claw-shaped (Figs. 5a, b, 12).

DISTRIBUTION: Myrmecoroides is distributed from South Australia, along the coastal margins of southeastern Victoria and New South Wales, and as far north as southeast Queensland. Two of the species, M. carinatus and M. cassowary, are only known from the type localities. M. grossi is the most widely distributed species, and is found from Victoria, to the Carnarvon Range in southcentral Queensland.

REMARKS: Myrmecoroides is highly autapomorphic on the basis of the hatchetlike head and the reduced genitalia of the included species. It is one of the most recognizable genera of the Austromirini, and certainly one of the most remarkable ant mimics in the Heteroptera. Myrmecoroides is related to other ant mimetic genera in the Austromirini with simple genitalia. This clade is distinct in relation to the clade of ant mimetic Austromirini that possess a bicompressed labrum and triangular head, such as the genus Myrmecoridea Poppius.
KEY TO SPECIES OF MYRMECOROIDES

1. Females                                                           2
   Males                                                             5

2. Clypeus greatly enlarged, extending below anterior
   margin of the labrum (Figs 4a-d, f, g)                            3

   --Clypeus moderate in size, reaching anterior
   margin of labrum (Fig. 4b)
                                            M. cassowary, new species

3. Forelegs yellow, mid and hind legs red; frontal
   plate posteriorly rounded (Fig. 4d)
                                                   M. carinatus Gross

   --Forelegs concolorous with mid and hind legs;
   frontal plate posteriorly angulate (Fig. 4d, f)                   4

4. Body mostly black, with orange to yellow
   markings; sclerotized rings broad (Fig. 4c)
                                                   M. grossi Carvalho

   --Body more orange brown; sclerotized rings
   narrow (Fig. 4e)                       M. monteithi, new species

5. Frontal plate of head strongly tapered posteriorly
   (Fig. 4g); right paramere T-shaped (Fig. 15c);
   pygophore without dorsosinistral tumescence
   (Fig. 15a); genital opening of pygophore with
   elongate, arcuate tubercle (Fig. 15a)
                                            M. rufescens, new species

   --Frontal plate of head not tapered posteriorly
   (Fig. 4a, c); right paramere club-shaped, with a
   subperpendicular shaft (Figs 8d, 11c, 13c);
   genital opening of pygophore without tubercles
   (Figs. 8a, b, 11a, 13a) or with two short
   tubercles (Figs 11a, 13a)                                         6

6. Frontal plate of head greatly reduced (Fig. 4a);
   dorsosinistral tubercle of pygophore greatly
   enlarged (Fig. 8a, b)                    M. cassowary, new species

   --Frontal plate of head greatly enlarged (Figs 4c,
   d); dorsosinistral tubercle of pygophore moderately
   enlarged (Figs 11a, 13a)                                          7

7. Head uniformly black; basal sclerite of endosoma
   serrate apically (Fig. 11d) grossi Carvalho

   --Head orange, gula black; basal sclerite of
   endosoma smooth apically (Fig. 13d)      M. monteithi, new species


[FIGURE 2 OMITTED]

[FIGURE 3 OMITTED]

[FIGURE 4 OMITTED]

Myrmecoroides carinatus Gross, 1964

Figures 3, 4h, 5, 6, and 16

Myrmecoroides carinatus Gross, 1964:8 (n.sp.); Carvalho, 1973:9 (diagnosis); Cassis and Gross, 1995:184 (catalog); Schuh, 1995:146 (catalog).

[FIGURE 5 OMITTED]

HOLOTYPE: VICTORIA: [female], Donnybrook, 29 November 1959, J. Ozols (MV).

Paratypes: 2[female] [female]), same data as for holotype (MV; SAMA). Types seen.

DIAGNOSIS: This species is only known from females and is recognized by the following combination of characters: most of head and most of forewings orange brown, with apices of forewings whitish, pronotum and most of abdomen dark brown to fuscous; abdominal connexiva yellowish or white with red highlighting; clypeus large, rounded, extending below bucculae (Fig. 4h); frons evenly arcuate, posteriorly arcuate, extending to posterior margin of eyes (Fig. 4h); AI short; forelegs mostly yellowish brown, contrasting with mostly reddish brown mid and hind legs; forewings with vestige of claval vein; sclerotized rings thickened, strongly folded (Fig. 5a, b); inter-ramal lobes obovate, narrowed distally, with apex truncate (Fig. 5a, b).

[FIGURE 6 OMITTED]

M. carinatus is only known from the three females of the type series. This species has the following autapomorphies: posteriorly rounded frontal plate; short labium; forelegs color contrasting with mid and hind legs; and, vestiges of median flexion line and claval vein visible. Females are micropterous, with the forewing subdivisions still present, despite the extent of wing-shortening. These attributes though subtle, are sufficient to differentiate it from M. grossi and M. monteithi. In addition, the posterior margin of the frontal plate is more truncate in the latter two species (cf. Fig. 4a, f, w)

DESCRIPTION: Female only. COLORATION: Head, forewings, and forelegs mostly yellowish brown, contrasting with dark reddish brown to fuscous thorax and abdomen. Head: Mostly yellowish brown, with gula fuscous, outer margins of clypeus and frons more reddened, vertex obscurely embrowned; mandibular and maxillary plates red. Antennae: AI yellowish brown; AII banded, medially yellowish brown, remainder reddish brown; AIII and AIV red brown. Labium: Mostly yellowish brown, LIV embrowned distally. Thorax: Pronotum uniformly dark reddish brown, pleural regions more reddish brown; scutellum red-fuscous; forelegs mostly yellow brown, with ventral margins of femora more embrowned; mid and hind legs mostly reddish brown, with bases and apices of tibiae yellow; tarsi embrowned distally. Abdomen: Mostly dark red-fuscous, mediosternites mostly white to yellow, with red highlighting. TEXTURE: Body impunctate, polished, callosite region of pronotum, corium and abdominal terga weakly shagreened. VESTITURE: Body with scattered simple, pale, semierect setae. Head: Leading edge of frons with sparse distribution of short, decumbent setae; vertex and gula with sparse distribution of more elongate setae. Antennae: AII-AIV with dense distribution of short, decumbent, simple setae. Thorax: Pronotum, forewings, and abdomen with sparse distribution of elongate setae, most dense on abdomen; femora with sparse distribution of semierect simple setae; tibiae with decumbent simple setae, more dense distally, without bristle-like, semierect setae. STRUCTURE: Robust body. Head: Clypeus large, rounded distally, extending well below bucculae; frons strongly flattened, leading edge of frontal plate evenly arcuate, posteriorly truncate (Fig. 4h), extending anteriorly beyond fronto-clypeal suture, extending posteriorly beyond eye. Antennae: Not greatly elongate, AI short, barely passing anterior margin of clypeus, AII length more than twice breadth of posterior margin of pronotum, AIII shorter than AII, AI and AIV subequal in length. Labium: Short, barely reaching base of mesocoxae, LI much shorter than posterior margin of head, just surpassing bucculae. Pronotum: Hour-glass shaped (Fig. 3), posterior margin of pronotum weakly rectilinear; disk shorter than callosite region; collar not distinct. Forewings: Obovate, median flexion line present, claval vein discernible. Pregenital abdomen: urites II and III strongly constricted, elongate. Female genitalia: (Fig. 5a, b): Sclerotized rings strongly folded basally, outer margins thickened, distally compressed; dorsal labiate plate spiculate on outer margins of sclerotized rings; ventral 1ablate plate spiculate more broadly across sclerotized rings; opening of common oviduct moderately sclerotized; inter-ramal sclerites suboval, rounded apically, spiculate basally; inter-ramal lobes claw-shaped, narrowed distally, apex rounded, weakly spiculate, mostly on apex; medial portion of the inter-ramal sclerite short (Fig. 5a, b).

[FIGURE 7 OMITTED]

[FIGURE 8 OMITTED]

Males and larvae unknown.

MEASUREMENTS: See Table 1.

DISTRIBUTION: This species is known only from type locality, in Victoria (Fig. 6).

HOST ASSOCIATIONS AND HABITATS: Nothing is known of the biology of this species. The microptery of the females suggests that, as with other species of Myrmecoroides, they are likely to be ground dwelling.

[FIGURE 9 OMITTED]

Myrmecoroides cassowary n.sp.

Figures 4a, b, 5c, d, 6, 7, 16

ETYMOLOGY: This species is named for the crest on the head that is reminiscent of the spectacular ratite of tropical Queensland and Papua New Guinea, the cassowary.

HOLOTYPE: SOUTH AUSTRALIA: [male], 25.8 km N Miandaria, 27[degrees]03'23"S 137[degrees]32'32"E, 9-14 September 1998, Sandy Desert Survey KAL040, ex Hakea leucophara shrubland over Zygochloa paradoxa open hummock grassland in broad sandy swale (SAMA). Paratypes: 10[female][female] (2 badly damaged), 13.6 km WNW of Beal Hill, 26[degrees]34'25"S 137[degrees]42'54"E, 14-18 September 1998, Dune crest dominated by Zygochloa paradoxa, Acacia ramulosa, and ephemeral herbs and grasses, Sandy Des[ert] Sur[vey] PEE0301, ex pitfalls (SAMA; UNSWIC).

[FIGURE 10 OMITTED]

DIAGNOSIS: This species is only known from females and is recognized by the following combination of characters: uniformly dark reddish brown, apices of forewings narrowly whitish, abdominal connexiva reddish; clypeus shortened, not extending below bucculae; frons explanate and raised posteriorly, crestlike in female (Fig. 4c); antennae uniformly yellowish brown; AI and AIV short (Fig. 7); AII and AIII banded (Fig. 7); forewings coriaceous; sclerotized rings not greatly thickened (Fig. 5c, d); inter-ramal lobes obovate, densely spiculate and uniformly sclerotized (Fig. 5c, d); base of medial portion of the interramal sclerite thickened (Fig. 5c, d).

This species is the most distinct of all the species of Myrmecoroides, with its highly autapomorphic head (relatively shortened clypeus, and the crestlike frontal plate, Fig. 4b). In addition, the female genitalia of this species are clearly differentiated from its congeners, with thin sclerotized rings (Fig. 5c, d), obovate and heavily sclerotized interramal lobes (Fig. 5c, d), and elaborate medial portion of the inter-ramal sclerite (Fig. 5c, d). It shares some characters with carinatus, which is most proximate to in, the latter found in eastern Victoria; both have short first antennal segment and pale foretibiae. However, they differ markedly in the shape of the frontal plate (cf. Fig. 4a, b).

[FIGURE 11 OMITTED]

DESCRIPTION. COLORATION: Mostly dark reddish brown, occasionally head and pronotum a little paler. Antennae: Stramineous to yellowish brown, AII and AIII sometimes banded apically. Labium: Sometimes more fuscous distally. Forewings: Reddish fuscous, apex with whitish spot, costal margin sometimes paler. Legs: Tarsi fuscous, contrasting with paler tibiae. Abdomen: Connexiva stramineous to yellowish brown with red highlighting.

TEXTURE: Impunctate, polished, not shagreened; forewings coriaceous, more so along inner claval margin. VESTITURE: With scattered simple, pale, semierect setae. Head: Leading edge of frons with sparse distribution of short setae; vertex and gula with sparse distribution of more elongate setae. Thorax: Pronotum, forewings, and abdomen with most dense distribution of elongate setae. Legs: Femora with sparse distribution of semierect simple setae; tibiae with decumbent simple setae, more dense distally, intermixed with bristlelike, semierect setae. STRUCTURE: Robust body. Head: Clypeus truncate distally, not extending below bucculae, much shorter than frons; frons posteriorly explanate, and greatly elevated, crestlike (Fig. 4e), not extending anteriorly beyond fronto-clypeal suture. Antennae: Not greatly elongate, AI short, barely passing anterior margin of clypeus, AII length about twice breadth of posterior margin of pronotum, AIII shorter than AII, AIV very short, shorter than AI. Labium: Reaching metacoxae, LI almost reaching posterior margin of head. Pronotum: Hourglass shaped (Fig. 7), posterior margin weakly convex; outline of collar discernable, without posterior margin; forewings obovate. Metafemora unknown. Male genitalia: Pygophore with sinistrodorsal tumescence (Fig. 8a, b), left paramere claw-shaped, apex bifid (Fig. 8c); right paramere clubshaped (Fig. 8a), basal spicula of endosoma elongate, bifid distally (Fig. 8e). Female genitalia: (Fig. 5c, d) Sclerotized rings strongly folded basally, margins narrow, distally compressed; dorsal labiate plate spiculate distad to sclerotized rings; ventral labiate plate more broadly spiculate across sclerotized rings; opening of common oviduct moderately sclerotized; inter-ramal sclerites subtriangular, tapered apically, spiculate ventrally along basal margin; inter-ramal lobes obovate, narrowed distally, apex rounded, densely spiculate and strongly sclerotized; medial portion of the inter-ramal sclerite elongate, thickened basally and spiculate, lateral margins sinuate.

MEASUREMENTS: Table 1.

Males and larvae unknown.

DISTRIBUTION: This species is known only from two localities in southeastern South Australia (Fig. 6).

HOST ASSOCIATIONS AND HABITATS: This species is known from a sand dune habitat, which was dominated by Zygochloa paradoxa (R.Br.)S.T.Blake, Acacia ramulosa W.Fitzg., and ephemeral herbs and grasses. Its collection in pitfall traps indicates that the females of this species are likely to be epigaeic.

Myrmecoroides grossi Carvalho, 1973

Figures 2, 4c, d, 6, 8, 9, 10, 11, 12, 16

Myrmecoroides grossi Carvalho, 1973:7 (n.sp.); Cassis and Gross, 1995:184 (catalog); Schuh, 1995:146 (catalog).

[FIGURE 12 OMITTED]

HOLOTYPE: [male], Otford, New South Wales, D. McAlpine (AM; very badly damaged).

SPECIMENS EXAMINED: QUEENSLAND: 2[male][male], Bunya Mountains, 26 March 1957 (QM); [male], 1 larva, Mahogany Forest, Mt Moffatt National Park, 24[degrees]55'S 148[degrees]04'E, 1,200 m, 26 September 26 November 1995, GB Monteith, intercept (QM); 1[male], same data as previous, CJ Burwell and S Evans (QM); NEW SOUTH WALES: 8[male][male], 19, Marengo State Forest, jct. Chalundi and Top Roads 30[degrees]05'51.3"S 152[degrees]22'23.8"E, G Cassis and R. Silveira, 28 February 2002, (AM); 1[male], Yadboro State Forest, start of Castle Walking Track, 20 January 1994, G Cassis, ex grass sweep (AM; 2[male][male], damaged) Cabbage Tree Creek, Clyde Mountain, 22 February 1965 DH Colless (ANIC); 4[male][male], Cabbage Tree Creek, foot of Clyde Mountain, 9 November 1981 (ANIC; 1[male], damaged), The Creel, Kosciusko, 8 November 1961, EF Reik (ANIC); AUSTRALIAN CAPITAL TERRITORY: 1[male], Australian Capital Territory: 35[degrees]22'S 148[degrees]50'E, February 1987, DH Colless, malaise trap (ANIC). Paratypes. 1[male], 1[female], same data as holotype (AM; ANIC); 1[male], 599, Australian Capital Territory: Blundells Creek, near Piccadilly Circus, February 1984, T Weir, JF Lawrence, ML Johnson, flight intercept (AM; ANIC); 1[male], Blundell's, 18 February 1931, AL Tonnoir (ANIC); VICTORIA: 1[male], Inverloch, foreshore veg[etation], 6 December 1977, TR New (SAMA); 1[male], Mitta River, Dartmouth Survey, location 546493, 28 November-3 December, 1973 (MV).

[FIGURE 13 OMITTED]

DIAGNOSIS: This species is recognized by the following combination of characters: slender species (Figs 9, 10), dorsum mostly fuscous, with orange and white markings; clypeus elliptoid, distally rounded (Figs 2b, 4c, f); frontal plate evenly arcuate, obtusely tapered posteriorly (Figs 2a, b, 4c, d); first antennal segment elongate, bottle-shaped (Figs 2b, 9, 10); second antennal segment cylindrical (Figs 9, 10); labium short; hind tibiae linear, with elongate, pale bristlelike setae; genital opening of pygophore without tubercles (Figs 2g, h, 11a); left paramere L-shaped (Fig. 10b); right paramere club-shaped (Fig. 11c); and, apex of basal sclerite of endosoma serrate (Fig. 11d).

DESCRIPTION: COLORATION: Mostly fuscous, occasionally head and pronotum, and markings on forewings paler, mostly orangish brown. Antennae: AI yellowish brown to dark reddish brown; AII mostly reddish brown, with a submedial yellow band proximal to midpoint; AIII and AIV red brown. Labium: Mostly yellowish brown, with LIV mostly fuscous. Pronotum: Fuscous to callosite region fuscous with disk more orangish brown. Forewings: Mostly fuscous, with corium orangish brown, sometimes medial aspects enbrowned; apex of corium and base of cuneus narrowly whitish; membrane fumose. Legs: Femora reddish brown; tibiae mostly reddish brown, with apices yellow; tarsi dark reddish brown to fuscous. Abdomen: Mostly dark reddish brown to fuscous; terga sometimes with red highlighting. TEXTURE: Impunctate, polished, callosite region of pronotum and corium anteriad to orangish brown marking, shagreened. VESTITURE: With scattered simple, pale, semierect setae; pronotum, forewings, and abdomen with moderate distribution of semierect, pale, elongate setae. Head: Leading edge of frontal plate with sparse distribution of short setae; clypeus with sparse distribution of dark, more robust setae; vertex and gula with moderate distribution of more elongate setae; Legs: Femora with moderate distribution of semierect simple setae; tibiae with decumbent simple setae, more dense distally, inter-mixed with bristlelike, semierect setae, latter longer than tibial width. STRUCTURE: Head: Clypeus elliptoid, extending below bucculae (Figs 2b, 4c, d); frontal plate evenly arcuate, obtusely tapered posteriorly (Figs 2b, 4c, d), extending anteriorly beyond fronto-clypeal suture, sometimes with a toothlike appearance (Figs 2b, 4c, d). Antennae: Moderately elongate, AI bottle-shaped, elongate, well surpassing leading edge of clypeus (Figs 2a, 4c, d, 9, 10); AII length about twice breadth of posterior margin of pronotum (Figs 9, 10); AIII shorter than AII; AIV subequal in length to AI. Labium: Reaching apices of mesocoxae; LI surpassing bucculae by half its length. Pronotum: Male strongly campanulate (Fig. 9); female hourglass shaped (Fig. 10); collar obsolete. Pterothoracic pleura: Propleural suture anterodorsally directed; proepisternum rounded; proepimeron short. Forewings: Female obovate. Thorax: External efferent system of metathoracic glands: peritreme strongly tumose (Fig. 2e); opening of metathoracic spiracle, small, oval, visible, with evaporative areas (Fig. 2f). Legs: Metafemora very weakly arcuate. Male genitalia: Pygophore without tergal processes, with sinistrolateral tumescence (Figs 2g, h, 11a); basal sclerite of endosoma serrate apically, on inner margin (Fig. 11d); left paramere club-shaped, apically rounded (Fig. 10b); right paramere L-shaped, distally scalloped (Fig. 11c). Female genitalia: (Fig. 12a, b) Sclerotized rings strongly folded basally, margins moderately broad, distally compressed; dorsal labiate plate spiculate on outer margin of sclerotized rings, and adjacent to the medial margin of sclerotized rings; ventral labiate plate more broadly spiculate across sclerotized rings; opening of common oviduct moderately sclerotized; inter-ramal sclerites subtriangular, weakly spiculate ventrally along basal margin; inter-ramal lobes claw-shaped, with thickened base, strongly narrowed distally, weakly spiculate and weakly sclerotized; medial portion of the interramal sclerite elongate, not thickened basally and not spiculate, lateral margins rectilinear.

[FIGURE 14 OMITTED]

[FIGURE 15 OMITTED]

[FIGURE 16 OMITTED]

MEASUREMENTS: Table 1.

DISTRIBUTION: This species is known from numerous localities in eastern Australia (Fig. 6), from Victoria to north of Brisbane, and as far west as the Australian Capital Territory, and two localities in the gorge country of south-central Queensland.

HOST ASSOCIATIONS AND HABITATS: This species has been collected from two localities in New South Wales (Yadboro and Marengo State Forest), sweeping grasses.

REMARKS: Myrmecoroides grossi is the most widely distributed species and exhibits the most morphological variation. The species was originally described from a single specimen, from the southern reaches of Sydney (Otford), probably in more open habitat. Populations at Cabbage Tree Creek, further south of Yadboro, exhibit some differences from the former populations in color, being generally a little lighter, particularly on the frontal plate of the head, pronotal disk and clavus, all of which are more orangish brown. However, the male genitalia are nearly identical between all of these populations, and support the decision that they are conspecific. There may also be some preservation factors that explain color differences; the latter specimens were collected in ethanol, from which they were dry mounted, which likely resulted in color fading.

Males from the most southern (Victoria) and northern (Queensland) reaches of the distribution of M. grossi have also been dissected, again indicating conspecificity.

Myrmecoroides monteithi n.sp.

Figures 4e, f, 6, 12c, d, 13 and 16

ETYMOLOLOGY: This species is named after Dr Geoff Monteith, who collected this species.

HOLOTYPE: QUEENSLAND: [male], Bald Mountain area, 3,500-4,000 ft, via Emu Vale, 27-31 January 1972, BK Cantrell (EUQ). Paratypes: 9, same data as for holotype (EUQ); 3[male][male][female], same location as for holotype, GB Monteith (EUQ); Bunya Mountains, 2 March 1957, EF Riek (ANIC)

DIAGNOSIS: This species is known from male and female specimens and is recognized by the following combination of characters: body mostly orangish brown, with fuscous pronotum and abdomen; clypeus enlarged; male frontal plate posteriorly arcuate, terminating past posterior margin of eyes (Fig. 4e); female frontal plate terminating posteriorly in line with posterior margin of eyes, posteriorly truncate (Fig. 4f); AI elongate, extending beyond leading edge of frontal plate (Fig. 4e, f); female micropterous forewing orangish brown, without apical white band; male macropterous forewing mostly orangish brown, sometimes base of clavus darker; region bounding corial fracture white; pygophore with dorso-sinistral tumescence, genital opening without tubercles (Fig. 13a); left paramere clubshaped (Fig. 13b); right paramere L-shaped, distally scalloped (Fig. 13c); basal sclerite of endosoma smooth (without serrations) (Fig. 13d); inter-ramal lobes claw-shaped (Fig. 12c, d); sclerotized rings folded basally, not strongly pinched distally (Fig. 12c, d); medial portion of the interramal sclerite elongate (Fig. 12c, d).

This species is very similar to M. grossi, from which it is difficult to distinguish on the basis of external characters. M. monteithi is a more robust species that is generally paler in color than M. grossi. In addition, the clypeus of both sexes of M. monteithi is larger, and in the male of this species the posterior termination of the frontal plate is closer to the posterior margin of the eye than in M. grossi (cf. Figs 4c, d, e, f).

DESCRIPTION: COLORATION: Mostly orangish brown, with dark brown and pale markings. Head: Mostly orangish brown, sometimes with outer margins of clypeus and frontal plate, and mandibular and maxillary plates with red highlighting; gula dark brown to fuscous. Antennae: AI yellow to yellowish brown; AII mostly brown with broad, sub-basal pale band; AIII-AIV brown. Labium: mostly yellow, with apex of LIV embrowned. Thorax: Pronotum uniformly fuscous, sometimes a little paler on posterior margin of disk; mesoscuturn reddish brown; scutellum fuscous. Legs: Mostly reddish brown; tibiae with bases and apices with short yellow annulations, foretibiae sometimes more broadly yellow distally; tarsi brown. Female forewing: Mostly orangish brown, sometimes a little paler distally. Male forewing: Mostly orangish brown, sometimes with base of clavus darker brown; apex of corium at fracture with white fasciae; cuneus and membrane orangish brown. Abdomen: Mostly dark brown with reddish highlighting; lateral aspect of female sternites whitish. TEXTURE: Impunctate, polished, pronotum and corium shagreened. VESTITURE: With scattered simple, pale, elongate, semierect setae. Head: Leading edge of frontal plate with moderate distribution of short to elongate setae; vertex and gula with sparse distribution of more elongate setae. Pronotum, forewings, and abdomen with most dense distribution of elongate setae. Legs: Moderate distribution of elongate, semierect setae; tibiae with elongate setae more bristlelike, intermixed with shorter more decumbent setae. STRUCTURE: Slender species; males macropterous; females micropterous. Head: Clypeus enlarged, outer margin arcuate (Fig. 4e, f), extending well below bucculae; male frontal plate not tapered, evenly arcuate, posterior margin rounded, extending beyond posterior margins of eyes (Fig. 40; female frontal plate enlarged, more truncate posteriorly, terminating at posterior margins of eyes (Fig. 4f). Antennae: AI elongate, extending beyond leading edge of frontal plate (Fig. 4e, f); AII elongate, about twice breadth of posterior margin of pronotum; AIII shorter than AII; AI and AIV subequal in length. Labium: Reaching just beyond forecoxae in males, and apices of mesocoxae in females. LI almost reaching posterior margin of head in females. Pronotum: Campanulate in males; hour-glass shaped in females; collar and posterior margin of disk weakly convex in both sexes. Forewings: Females obovate. Legs: Metatibiae arcuate in both sexes. Abdomen: Strongly constricted, basally, TII and Till elongate, bilaterally compressed. Male genitalia: Pygophore with dorsosinistral tumescence (Fig. 13a); genital opening without tubercles (Fig. 13a); left paramere club-shaped, distally rounded, without basal processes (Fig. 13b); right paramere angulate, with apex scalloped (Fig. 13c); endosoma with straplike basal sclerite, tapered distally, smooth (without serrations) (Fig. 13d). Female genitalia: (Fig. 12c, d) Sclerotized rings strongly folded basally, margins relatively narrow, not distally compressed; dorsal labiate plate spiculate on outer margin of sclerotized rings; ventral labiate plate more broadly spiculate across sclerotized rings; opening of common oviduct weakly sclerotized; inter-ramal sclerites subtriangular, weakly spiculate along medial margin; inter-ramal lobes clawshaped, with angulate base, strongly narrowed distally, weakly spiculate and weakly sclerotized; medial portion of the inter-ramal sclerite elongate, not thickened basally and not spiculate, lateral margins rectilinear.

Larvae unknown.

MEASUREMENTS: See Table 1.

DISTRIBUTION: This species is known from two locations in southeast Queensland (Fig. 6).

HOST ASSOCIATIONS AND HABITATS: There is no biological information for this species. Monteith (pers. comm.) indicates that the Bald Mountain locality is characterized by cool temperate forest that contains numerous southern elements.

Myrmecoroides rufescens n.sp.

Figures 4g, 6, 14, 15, 16

ETYMOLOGY: This species is named for the reddish coloration of the dorsum.

HOLOTYPE: NEW SOUTH WALES: [male], Warrah, WW Froggatt, 6-11-1900 (SAMA). Paratypes: 2[male][male], same data as for holotype (SAMA; badly damaged); 1[male], Maclean, IC Yeo, 13 December 1964, sweeping grasses (QM). QUEENSLAND: 1[male], Beewah, 2 October 1966, JC Cardale (AM); 1[male], Whetstone, IC Yeo, 27 April 1959 (EUQ).

DIAGNOSIS: This species is only known from males and is recognized by the following combination of characters: robust species (Fig. 14); dorsum mostly fuscous, with reddish, orange and white markings; clypeus elliptoid, distally rounded (Fig. 4g); frontal plate strongly tapered posteriorly (Fig. 4g); first antennal segment elongate, bottle-shaped (Figs 4g, 14); second antennal segment fusiform; labium short; hind tibiae arcuate, with elongate, pale bristlelike setae; genital opening of pygophore with digitiform tubercle (Fig. 15a); left paramere T-shaped (Fig. 15b); right paramere claw-shaped (Fig. 15c); and, apex of basal sclerite of endosoma serrate (Fig. 15d).

This species is the most distinct in the genus, with its unique frontal plate, which is strongly tapered posteriorly (Figs4g, 13). The male genitalia are particularly distinct, with the genital opening of the pygophore with an enlarged tergal process (Fig. 15a), and the uniquely shaped parameres (Fig. 15b, c).

DESCRIPTION (males only): COLORATION: Pronotum, mesoscutum and scutellum and cuneus dark-red brown, head and wings mostly orangish brown. Head: Orange to reddish brown, darker on vertex and gula. Antennae: Orangish brown, with base of AII yellow. Labium: Mostly orangish brown, with LIV darker, almost fuscous. Pronotum: Fuscous to dark reddish brown, a little lighter on disk, latter concolorous with mesoscutum and scutellum. Forewings: Clavus mostly orangish brown, darker at extremities; basal corium brown, then yellowish brown, with narrow apical white stripe contiguous with whitish basal area of cuneus, remainder of cuneus and membrane dark brown. Legs: Mostly reddish fuscous, with apices of tibiae lighter, yellow to orangish brown. Abdomen: Uniformly dark reddish brown to fuscous, with posterior border of SII yellow, sometimes with red highlighting. TEXTURE: Impunctate, polished, callosite region of pronotum and basal half of corium shagreened. VESTITURE: Body with relatively dense distribution of simple, pale, erect setae, intermixed on tibiae with elongate bristlelike setae Head: Leading edge of clypeus and frons, vertex and gula with particularly dense distribution of simple, erect setae. Antennae: AII-AIV with dense distribution of short, simple, decumbent setae, AII with row of trichobothrialike setae. STRUCTURE: robust body. Head: Clypeus flattened, elliptoid, distally rounded (Figs 4g, 14); distally, extending below bucculae (Figs 4g, 14); frontal plate extending below dorsal margin of clypeus, with a beaklike appearance, strongly tapered posteriorly (Figs. 4g, 14). Antennae: AI elongate, bottle-shaped (Figs. 4g, 14); AII incrassate distally, about 1.8 longer than posterior width of pronotum; AIII and AIV narrow, AIII 1/2 length of AIII, AIV short, less than width across eyes. Labium: Reaching apices of mesocoxae, LI extending beyond bucculae by 1/2 its length. Pronotum: Campanulate; collar discernable, without posterior margin; posterior margin of pronotum weakly concave. Forewings: costal margin strongly excavate; cuneus elongate (Fig. 14); metafemora and metatibiae bowed (Fig. 14). Male genitalia: Genital opening of pygophore with dorsal, digitiform tubercle (Fig. 15a); left paramere T-shaped (Fig. 15b); right paramere claw-shaped (Fig. 15c); aedeagus with basal sclerite serrate distally (Fig. 15d).

Female and larvae unknown.

MEASUREMENTS: See Table 1.

DISTRIBUTION: This species is known from four localities in northeast New South Wales and southeast Queensland (Fig. 6). Its distribution crosses the geographic barrier of the Macleay-McPherson Overlap.

Host ASSOCIATIONS AND HABITATS: This species has been collected by sweeping grasses from two localities in New South Wales (Maclean and Warrah).

PHYLOGENY

The characters and character state lists are given in Table 2. The data matrix is given in Table 3. A traditional search was undertaken with 500 replications. A single tree was found of 23 steps, with a consistency index of CI: 0.9565 and a retention index of 0.9714 (Fig. 16). The following discussion refers to character state optimized on the above tree.

Clade Myrmecoroides. Myrmecoroides is strongly supported by synapomorphies of the forewings, head and pretarsus, and 100 bootstrap value. As opposed to the outgroups, females are micropterous in all species of Myrmecoroides (character 1-1). This condition is not exclusive to Myrmecoroides, and occurs in other genera of Austromirini, but not in the subclade of genera of Austromirini as identified by Cassis et al. (in preparation). The most compelling synapomorphies of Myrmecoroides are those of the highly modified head, with the strongly bicompressed clypeus (3-1) and the frons and vertex modified along the midline into a crestlike frontal plate (5-1). These character states are not known to occur in any other species of Miridae. The pretarsus is also distinctive, with the distally angulate claws (11-1), greatly expanded lamellate parempodia (12-1), and greatly enlarged pulvilli (13-1). The latter character is expressed elsewhere in the Orthotylinae, and is probably not as significant as the other two characters of the pretarsus.

Clade M. carinatus (M. rufescens (M. grossi + M. monteithi). This clade is supported by a bootstrap value of 100. The most significant differences exist in the development of the clypeus, where in M. cassowary it is bicompressed but not greatly enlarged as in the remaining species (2-1), where it extends beyond the clypeus. Similarly, the frons does not extend beyond the clypeal suture in M. cassowary but in all other species, projects well onto the surface of the clypeus (4-1). In addition, the first antennal segment of cassowary is not modified into a distinctly bottle-shaped outline (6-1) for M. carinatus + (M. rufescens (M. grossi + M. monteithi)). Less significant characters for this subclade include the tomentose dorsum (2-1) and the basal brown annulation of the second antennal segment (9-1) for the latter subclade.

Clade M. rufescens (M. grossi + M. monteithi). This clade is the least supported with a single synapomorphy (7-1: posterior margin of female frontal plate angulate) and a bootstrap value of 65. Unlike in M. carinatus and M. cassowary, the frontal plate of the female is truncate posteriorly.

Clade M. grossi + M. monteithi. This clade is supported by two synapomorphies (15-1: pygophore of male with a sinistrodorsal tumescence; 16-1: club-shaped left paramere) and two homoplasies (14-1: genital opening of male with a tergal process; 17-1: right paramere L-shaped), and a bootstrap value of 64.

DISCUSSION

The hatchet head bugs are one of the most enigmatic genera of Miridae found in Australia. Like other austromirine myrmecomorphs, there is the characteristic "petiolelike" bilateral compression of the basal abdominal urites (particularly segments II and III), elongation of the legs, the forewing transverse white markings in males (suggestive of an abdominal petiole), and microptery of females (also correlated with pronotal shortening). However, unlike other myrmecomorphs, hatch head bugs have a remarkable development of the frons and vertex into a crest shaped frontal plate (contiguous with bicompressed clypeus). These structures represent some of the most exaggerated morphologies seen in the Heteroptera, whose adaptiveness is unknown. We know of no ants or ant morphology that would form the basis of a model for such head specialization.

Myrmecoroides is nested within a clade of the Orthotylinae: Austromirini, where the labrum is flattened, and not bilaterally compressed as in Myrmecoridea Poppius (Cassis et al. in preparation). The former genus is also characterized by the very small male genitalia (Fig. 2g, h).

As with many other mirid species, the males are obligatively macropterous and the female micropterous. Such wing dimorphism appears to be correlated to grass-inhabiting and epigaeic habits respectively. In all likelihood, the females are also to be found upon plants as well, at least facultatively, probably at the base of grass sheaths, where oviposition may take place. One hypothesis for such wing dimorphism in mirids, is that males act as "gene-dispersers" enabling gene flow within metapopulations, and flightless females are more closely tied to their host plants to ensure "faithful" oviposition so that the next generation has predetermined host selection (Wheeler, 2001). There is no empirical work to support this hypothesis, and it is unknown whether such an hypothesis is pertinent to Myrmecoroides. This is likely a fruitful area of investigation.

The distribution of Myrmecoroides is consistent with other supraspecific groups in the Heteroptera. Many other true bug taxa show a Torresian distribution (Cassis and Gross, 1995, 2002). However, unlike a number of true bug taxa, which have a broad distribution in temperate Australia, from east to west (e.g., Cassis and Silveira, 2001: Nerthra alaticollis species-group [Gelastocoridae: Nerthrinae]; Cassis and Moulds 2002: Kirkaldyella Reuter [Miridae: Orthotylinae]), Myrmecoroides is not known from Western Australia, with its western limit in the Adelaide area of endemism, bounding the Southern Gulfs drainage basin. Within Kirkaldyella, there are repeated east-west area relationships, with eastern Australia basal to the west (Cassis and Moulds 2002). In Myrmecoroides, there is a reversed relationship, with the most western area of endemism (Adelaide) basal to eastern Australia. However, the cryptic habits of the hatchet head bugs are such that additional species and/or more extensive distribution of known species are possible, pending more litter and grass tufting collections in the southwest of Western Australia.

ACKNOWLEDGMENTS

We thank the late Dr. Gordon Gross and Jan Forrest (SAMA) for the loan of material, and the use of the habitus illustrations of Myrmecoroides carinatus, M. grossi and M. rufescens. The scientific illustrators Jenni Thurmer and Hannah Finlay are thanked for the habitus illustrations. Hannah Finlay drafted the genital illustrations. Dr. Geoff Monteith (QM), Tom Weir (ANIC) and Ken Walker (MV) are thanked for the loan of material. Rossana Silveira is thanked for assistance in fieldwork. Other members of the Cassis lab, Nikolai Tatarnic and Celia Symonds are thanked for assistance with preparation of the manuscript. We thank the anonymous referees for critical reviews of the paper.

This work is in part supported by the Australian Biological Resources Study and the U.S. National Science Foundation, Planetary Biodiversity Inventory Program, DEB-0316459.

Received 22 March 2010; accepted 9 October 2010

LITERATURE CITED

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Cassis, G., Schwartz, M. D. and Molds, T. 2003. Systematics and new taxa of the Vannius complex (Hemiptera: Miridae: Cylapinae) from the Australian Region. Memoirs of the Queensland Museum 49(1): 123-151.

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Heteroptera: Miridae), pp. 193-212 in Hodkinson, T. R., Parnell, J. and Waldren, S. (eds.), Taxonomy and Systematics of Species Rich Taxa: Towards the Tree of Life. CRC, Boca Raton.

Davis, N. T. 1955. Morphology of the female organs of reproduction in the Miridae (Hemiptera). Annals of the Entomological Society of America 48: 132-150.

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Gross, G. F. 1964. A new ant-mimicking mirid bug (Hemiptera: Heteroptera) from Victoria. Memoirs of the National Museum of Victoria 26: 7-9.

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Schuh, R. T. 1995. Plant Bugs of the World (Insecta: Heteroptera: Miridae). Systematic Catalog, Distributions, Host List, and Bibliography. The New York Entomological Society, New York. xii + 1329 PP.

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GERASIMOS CASSIS (1,3) AND MICHAEL A. WALL (2)

(1) Evolution & Ecology Research Center, School of Biological, Earth and Environmental Sciences, University of New South Wales, Sydney 2052, Australia

(2) Department of Entomology, San Diego Natural History Museum, P.O. Box 121390, San Diego, California 92112-1390, USA

(3) Email address for correspondence: gcassis@unsw. edu.au
Table 1. Morphometric data for species of Myrmecoroides.

                                           Length

                       Body   Cun-Clyp   Head   Pron   Scut     Cun

M. carinatus
  Females (N = 2)
    Mean                5.49       9.73   1.36   0.77   0.19    5.45
    SD                  0.04       6.04   0.16   0.03   0.05    7.7
    Range               0.06       8.54   0.23   0.04   0.07   10.89
    Minimum             5.46       5.46   1.24   0.75   0.16    0
    Maximum             5.53      14      1.47   0.79   0.23   10.89

M. cassowary
  Females (N = 5)
    Mean                5.23      18.13   1.11   0.86   0.2        0
    SD                  0.1        4.66   0.04   0.03   0.05       0
    Range               0.25      11.46   0.09   0.09   0.13       0
    Minimum             5.11      12.36   1.06   0.83   0.14       0
    Maximum             5.37      23.82   1.16   0.92   0.27       0
M. grossi
  Males (N = 5)
    Mean                3.1        2.31   0.65   0.74   0.52    0.6
    SD                  1.79       1.55   0.57   0.57   0.26    0.03
    Range               3.38       3.17   1.16   1.59   0.62    0.07
    Minimum             1.11       0.34   0      0      0.35    0.56
    Maximum             4.49       3.51   1.16   1.59   0.96    0.64
  Females (N = 3)
    Mean                4.63       5.93   1.16   0.71   0.15    3.31
    SD                  0.14       2.26   0.05   0.04   0.07    3.28
    Range               0.29       4.05   0.11   0.07   0.13    6.57
    Minimum             4.49       4.49    1.1   0.68   0.11    0
    Maximum             4.77       8.54   1.21   0.75   0.24    6.57
M. monteithi
  Males (N = 2)
    Mean                3.91       2.76   0.52   0.77   0.28    0.8
    SD                  0.18       0.71   0.73   0.23   0.01    0.29
    Range               0.26       1.01   1.03   0.32   0.02    0.41
    Minimum             3.78       2.25   0      0.61   0.27    0.59
    Maximum             4.04       3.26   1.03   0.93   0.29    1
  Females (N = 1)
                       25.61      25.61   1.28   0.78   0.2      0
M. rufescens
  Males (N = 5)
    Mean                5.53       4.19   1.3    0.99   0.47    0.6
    SD                  0.59       0.47   0.11   0.16   0.08    0.09
    Range               1.41       1.09   0.26   0.44   0.2     0.23
    Minimum             4.64       3.42   1.14   0.73   0.33    0.45
    Maximum             6.05       4.52   1.41   1.17   0.54    0.68

                                        Width

                      Head    Pron   Scut   Inter0c   AntSeg2

M. carinatus
  Females (N = 2)
    Mean               0.96    0.75   0.26      0.53      1.75
    SD                 0.02    0.04   0.06      0.01      0.02
    Range              0.02    0.06   0.09      0.02      0.03
    Minimum            0.95    0.72   0.22      0.52      1.74
    Maximum            0.97    0.78    0.3      0.54      1.77

M. cassowary
  Females (N = 5)
    Mean               0.91    0.98   0.24      0.42      2.01
    SD                 0.03    0.03   0.04      0.03      0.08
    Range              0.07    0.09    0.1      0.08      0.17
    Minimum            0.87    0.94   0.21      0.36      1.93
    Maximum            0.94    1.03   0.31      0.44       2.1
M. grossi
  Males (N = 5)
    Mean               1.15    0.86   1.36      1.14      0.78
    SD                 0.64     0.1   1.35      1.19      1.02
    Range              1.23    0.24   2.51      2.27      1.99
    Minimum            0.67    0.72   0.34      0.25      0
    Maximum            1.9     0.96   2.84      2.52      1.99
  Females (N = 3)
    Mean               0.89    0.7    0.15      0.48      1.83
    SD                 0.07    0.11   0.04      0.05      0.08
    Range              0.13    0.21   0.08      0.1       0.16
    Minimum            0.83    0.61   0.11      0.42      1.74
    Maximum            0.96    0.82   0.19      0.53      1.91
M. monteithi
  Males (N = 2)
    Mean               0.69    1.9    0.19      0.35      1.85
    SD                 0.02    1.36   0.14      0.12      0.1
    Range              0.03    1.92   0.2       0.18      0.14
    Minimum            0.67    0.94   0.09      0.26      1.78
    Maximum            0.7     2.86   0.29      0.44      1.92
  Females (N = 1)
                       0.95    0.77   0.28      0.55      1.93
M. rufescens
  Males (N = 5)
    Mean               0.91    1.29   0.54      0.44      2.15
    SD                 0.16    0.24   0.1       0.13      0.32
    Range              0.34    0.63   0.25      0.32      0.56
    Minimum            0.69    0.92   0.4       0.27      1.96
    Maximum            1.03    1.55   0.65      0.59      2.52

Table 2. Characters and character states for outgroups
and species of Myrmecoroides. Outgroup taxon
codes are: TASM001, PSEUD001 and AUSTRO001.

1. Female forewing condition: macropterous = 0;
submacropterous = 1.

2. Body texture: not tomentose = 0; tomentose = 1.

3. Clypeus profile: rounded = 0; strongly bilaterally
compressed = 1.

4. Shape of apical margin of clypeus: truncate = 0;
rounded = 1.

5. Frons + vertex profile: evenly arcuate = 0; frontal
plate = 1.

6. Apex of frons termination: at clypeal margin = 0;
below clypeal margin = 1.

7. Female frontal plate posterior margin: rounded = 0;
angulate = 1.

8. First antennal segment shape: cylindrical = 0;
vasiform = 1.

9. AII coloration: basally not banded = 0; basally
banded = 1.

10. Hind tibiae cross section: flattened = 0; round = 1.

11. Tarsal claw shape: arcuate = 0; angulate = 1.

12. Parempodia width: narrow = 0; broad = 1.

13. Pulvilli size: small = 0; large = 1.

14. Genital opening of male: without tergal process =
0; with tergal process = 1.

15. Male pygophore profile: without sinistrodorsal
tumescence = 0; with sinistrodorsal tumescence = 1.

16. Left paramere shape: claw shaped = 0; club shaped = 1.

17. Right paramere: multifurcate = 0; L-shaped = 1.

Table 3. Data Matrix for species of Myrmecoroides and outgroups
(TASM001, PSEUD001 and AUSTR0001). All 17 characters are binary.
? = missing data or inapplicable data. Note that the first
character is labeled as character 0.

            0   0   0   0   0   0   0   0   0   1   1   1   1   1   1
            1   2   3   4   5   6   7   8   9   0   1   2   3   4   5

TASM001     0   0   0   0   0   0   ?   0   0   1   0   0   0   0   0
PSEUD001    0   0   0   0   0   0   ?   0   0   1   0   0   0   0   0
AUSTR0001   0   0   0   0   0   0   ?   0   0   1   0   0   0   1   0
carinatus   1   1   1   1   1   1   0   1   1   1   1   1   1   ?   ?
cassowary   1   0   1   0   1   0   0   0   0   ?   1   1   1   ?   ?
grossi      1   1   1   1   1   1   1   1   1   1   1   1   1   1   1
monteithi   1   1   1   1   1   1   1   1   1   1   1   1   1   1   1
rufescens   1   1   1   1   1   1   1   1   1   0   1   1   1   0   0

            1   1
            6   7

TASM001     0   0
PSEUD001    0   1
AUSTR0001   0   0
carinatus   ?   ?
cassowary   ?   ?
grossi      1   1
monteithi   1   1
rufescens   0   0
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Article Details
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Author:Cassis, Gerasimos; Wall, Michael A.
Publication:Entomologica Americana
Article Type:Report
Geographic Code:1USA
Date:Jul 1, 2010
Words:8920
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