Systematics and host plant associations of a new genus of Acacia-inhabiting plant bugs from arid Australia (Insecta : Hemiptera : Heteroptera : Miridae : Orthotylinae)

Acaciacapsus, gen. nov. is described as a new plant bug genus, with eight new included species: A. amadeus, sp. nov.; A. appha, sp. nov.; A. aureolus, sp. nov.; A. bournda, sp. nov.; A. emeraldensis, sp. nov.; A. lolworthensis, sp. nov.; A. millstreamensis, sp. nov.; and A. woodwardi, sp. nov. Differential diagnoses and descriptions are given for all species, including salient characters, and the male and female genitalia. An identification key is provided to species. Male genitalia are illustrated, and a habitus photograph is provided for each species. Female genitalia are illustrated for two species. The genus is putatively an Acacia specialist, and has cryptozoic yellowish colouration. The species are primarily found in arid and semiarid regions of non-monsoonal regions of Australia. The collection events are digitised and their distributions mapped. A phylogeny of species is given. Modifications are given to male genitalic homologies and are discussed in reference to other Australian orthotylines.

Keywords: host relationships; phylogeny; species delineation; taxonomy.

The Australian Orthotylinae is a hyperdiverse subfamily of the plant bug family Miridae (Insecta: Hemiptera: Heteroptera), which includes many hundreds of undescribed species based on recent sampling (Cassis et al. 2007). Prior to this work, the nominotypical tribe was represented in Australia by 76 described species (Appendix 1; also see Cassis and Gross 1995; Cassis et al. 2012; Schuh 1995 for respective Australian and world catalogues; Schuh 2002-2013, online world catalogue). The subfamily is represented in Australia by four of the six recognised tribes (see Cassis and Schuh 2012 classification), with only the Nichomachini (Afrotropical and Palearctic regions) and Ceratocapsini (Western Hemisphere) not represented. The Halticini (Tatarnic 2009) and Coridromiini (Tatarnic and Cassis 2008,2013) have been recently revised, with each of these tribes represented by only a handful of species (five and four species respectively). The most species-rich group at present is the Austromirini, which is currently represented by fifteen genera and 38 species, although many new taxa await description (Cassis 2008; Cassis and Gross 1995; Cassis and Moulds 2002; Cassis and Symonds 2008; Cassis et al. 2007; Symonds and Cassis 2009, 2010). The Orthotylini are the most poorly documented group with only six genera and 12 species described, but with many hundreds of new species requiring description.

The Orthotylini is a cosmopolitan tribe that comprises 259 genera (Cassis and Schuh 2012) and hundreds of species (Schuh 1995, 2002-2013) worldwide. Previous to 2010, the tribe was represented in Australia by only six described species (Cassis and Gross 1995; Schuh 1995).Namyatova et al. (2011) described Witchelinamiris Namyatova, Elias and Cassis and two new species from South Australia. Cassis et al. (2010) described a monotypic genus, Harveycapsus dimorpha Cassis, Symonds and Tatarnic from Western Australia. Cheng et al.(2012a) described a new genus, Myrtlemiris, for nine new species, from the southwest corner of Western Australia, and a single species from the western slopes of New South Wales.

In 2009 the Australian Government implemented a new program of species discovery across new additions to the national reserve system -- the Bush Blitz species discovery program (www.bushblitz.gov.au). Taxa belonging to the new genus were found during Bush Blitz sampling and here they are studied along with other material subsequently identified in other collections. The aim of this work is to describe the taxa belonging to the new orthotyline genus Acaciacapsus gen. nov. and determine their relationships.

Specimens examined in this study belong to the following institutions: Australian Museum (AM), Queensland Museum (QM), Queensland Department of Agriculture, Fisheries and Forestry (QDPI), Australian National Insect Collection (ANIC), South Australian Museum (SAMA), Western Australian Museum (WAMP), and University of New South Wales (UNSW). All specimens have been labelled with unique specimen identifier (USI) codes (listed with specimen information in species descriptions), and entered into the Planetary Biodiversity Inventory (PBI) locality database (www.amnh.org.au/pbi). Specimen information can be viewed online through the Discover Life website (www.discoverlife.org). Genitalic dissections of males were made for most populations of all species, and females of two species.

Comparative morphological investigations, including the preparation of male and female genitalic dissections, were undertaken using Leica MZ16 and MZ205 stereomicroscopes. The genitalia were illustrated using a Leica DM5000B compound microscope with an attached camera lucida. Habitus photographs were taken for all Acaciacapsus species using a Canon 40D digital SLR attached to an Infinity K2 lens and the Visionary Digital microphotography system (www.visionarydigital.com). Multiple frames at different focal lengths were stacked using the Helicon Focus software (Kozub et al. 2008). A Hitachi TM3000 desktop scanning electron microscope was used to obtain micrographs of key diagnostic characters of Acaciacapsus. The Simple Mappr (Shorthouse 2010) online software was used to map Acaciacapsus distributions based on the digitised specimen records in the PBI locality database (research.amnh.org/pbi/locality).

Morphological character measurements were made using a stage-mounted digital micrometer. Five male and five female specimens were measured for each species, where possible; smaller sample sizes are indicated where appropriate.

A phylogenetic analysis was undertaken, including all species of Acaciacapsus. Fifty-six morphological characters (Table 1) were coded and character states are given in the data matrix (Table 2), with multistate characters unordered. Half the characters are of the male genitalic structures, which are the characters for distinguishing species, including more widely distributed species where external colouration varies. Many characters, both external and genitalic, are also diagnostic for the genus Acaciacapsus and have been included with the idea of building on this character matrix to include further Australian Orthotylini genera as they are described into the future. Four outgroup taxa from the tribe Orthotylini were used (Fig. 1, Table 2). The phylogeny was rooted with Orthotylus AU sp25. Three additional orthotylines were included in this analysis: Myrtlemiris yalgoo Cheng, Mututantri and Cassis, and two undescribed taxa (morphospecies codes: PBI Orthotylini msp 1.004 and COEX msp013). The selection of outgroup taxa was based on their putative close relationship with Acaciacapus. The morphological dataset was analysed in TNT (Goloboff et al. 2008), using parsimony, with 1000 random stepwise addition replicates and tree bisection-reconnection (TBR) branch swapping (10 trees/rep), with implied weighting (K=3-7). To determine branch support, symmetric resampling values were calculated, with 1000 random addition replicates and TBR branch swapping. Synapomorphies and contradicted apomorphies were viewed on the resulting implied weights phylogeny using MacClade 4.0.8 (Maddison and Maddison 2005), and listed in Table 3.

Cassis (2008) proposed a theory for endosomal spicule definition and evolution within the Orthotylinae in an attempt to recognise homologous components of the male genitalia, which are exaggerated, highly labile in structure, and of classificatory significance. Cassis' (2008) 'torsion theory of endosomal spicules' postulates that elongate, sclerotised processes, referred to as dorsal and ventral endosomal spicules, bound the secondary gonopore and extend distally, most often with marginal serrations and distal branching. Modifications of the spicules are indicative of species-level differences as well as having phylogenetic significance. This theory was proposed for the Lattinova complex of the tribe Austromrini, which have two endosomal spicules (dorsal and ventral endosomal spicules; as, DES and VES, with the DES defined as a basal keel). The spicules were designated as such by their basal orientation relative to the secondary gonopore. Cassis et al. (2010) extended the theory to the Orthotylini, for the monotypic Australian genus, Harveycapsus Cassis, Symonds & Tatarnic for taxa with three endosomal spicules, recognising two dorsal endosomal spicules (DES1 and DES2, with DES1 possessing a basal keel, and DES2 being the outer-most spicule relative to the secondary gonopore). Cheng et al. (2012a) also applied this nomenclature to the orthotyline genus Myrtlemiris Cheng, Mututantri and Cassis, although all three spicules were dorsal in orientation. This application however assigns the spicule closest to the secondary gonopore as the VES even though it is dorsal in position. In line with this, Cheng et al. (2012b) modified endosomal spicule nomenclature, and renamed the VES as the proximal endosomal spicule (PES) for Myrtlemiris, in reference to its proximity to the secondary gonopore, rather than its dorsoventral orientation. This is in keeping with our observation of other Australian orthotylines, which we have yet to describe, which also have all spicules in the dorsal position (Cassis, pers. obs.). We are applying this revised nomenclature for Acaciacapsus, within which there are only two endosomal spicules, with the PES either left lateral (A. appha, A. aureolus, A. bournda, A. emeraldensis and A. woodwardi) or left dorsolateral (A. amadeus, A. lolworthensis and A. millstreamensis). The PES is narrow at the base and the DES is broad at the base with a keel, with the latter dorsal to left dorsolateral in position.

Phylogenetic analysis resulted in an implied weights tree given in Fig. 1. Acaciacapsus (Node 1) is strongly supported as a monophyletic group by 20 synapomorphies and six contradicted apomorphies, which comprise 16 external and 10 male genitalic characters (Table 3, Fig. 1). Acaciacapsus lolworthensis is the sister-species to the remaining members of Acaciacapsus (Node 2), with the latter supported by one synapomorphy (left paramere with serrate sensory lobe, 34-1; Table 3) and one contradicted apomorphy (apex of elongate branch of PES with smooth margins, 54-0; Table 3). Node 3 includes two major subclades (Nodes 4 and 5) that are united by one synapomorphy (cuneus darker in colouration than corium, 3-1; Table 3). Node 4 comprises the sister-species relationship of A. amadeus and A. appha which are grouped on the basis of one contradicted apomorphy (tall sensory lobe, 33-1; Table 3). Node 5 comprises A. millstreamensis (A. woodwardi (A. bournda + A. emeraldensis)), which is united by one synapomorphy (small posterior lobes on pronotum, 20-0) and one contradicted apomorphy (distally tapering and weakly pointed phallotheca, 41-0; Table 3). Node 6 groups A. woodwardi (A. bournda + A. emeraldensis) by strong red cuneal colouration (2-2), and structure of the left canting branch of the dorsal endosomal spicule, with both branches distally broad and serrate (50-1) (also present in A. lolworthensis at the base of the genus group). Species of this node inhabit the same or proximate biogeographic areas in eastern Australia and all share the more reddish colouration seen in the genus. Node 7 comprises the sister-species A. bournda + A. emeraldensis) and is defined by the retracted phallotheca (38-0).

Family MIRIDAE Hahn, 1833

Subfamily ORTHOTYLINAE Van Duzee, 1916

Tribe ORTHOTYLINI Van Duzee, 1916

Genus Acaciacapsus, gen. nov.

Type species: Acaciacapsus aureolus Cassis & Symonds, sp. nov., by original designation.

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Acaciacapsus is recognised by the following combination of characters: golden ground colour, with contrasting orange, red or brownish markings (Character state 1-0); cuneus often contrasting red to reddish orange; eyes sexually dimorphic, in male strongly produced (13-1), width >interocular distance (14-1), slightly removed from posterior margin of head (15-1); eyes of both sexes extending to gula (16-1); vertex concave; lateral margins of pronotum strongly tapered towards apex; dorsum with patterned black, flattened scale-like setae (9-1), intermixed with moderately dense distribution of robust pale semi-erect simple setae on head, lateral margins of pronotum and scutellum (11-1); mesepimeron relatively large and polished; labium greatly elongate reaching between metacoxae and pygophore (18-1); femora elongate and cylindrical (25-1); ventral surface of forefemora of males with elongate bristle-like setae (12-1); hemelytral membrane clear spot most often present adjacent to tip of cuneus; membrane embrowned adjacent to veins and clear spot, membrane veins creamy yellow (6-0); anterolateral angles of pronotum narrow (21-0); humeral angles of pronotum acute (22-1); scutellum and mesoscutum raised (23-1); median flexion line short; forewings greatly elongate in males, exceedingly beyond abdomen, abdomen not extending beyond cuneal fracture; femora with dark, often red, bands apically and basally (7-1), more distinct in some species than others; pronotum often with conical projections on posterior margin; head often with contrasting red or orange vittae; ventral margin of genital opening of pygophore slanted towards left paramere (27-1); pygophore with left and right spine-like tergal processes on dorsal margin, right tergal process smaller than left tergal process (29-1); left paramere with hooked apex, elevated tapered and serrate sensory lobe; right paramere weakly c-shaped, usually serrate from medial dorsal margin to apex; aedeagus with two endosomal spicules, PES left lateral to left dorsolateral to secondary gonopore, DES dorsal to secondary gonopore, always more over the left side; PES longer than DES (52-1); PES with downturned medial process and divided apex, most distal branch, thin, tapered, ribbonlike and smooth (51-1); DES with large basal keel often with distal ornamentation; DES bifurcate near base, with one spatulate undivided branch, other branch divided distally and often serrate (49-1).

Colouration. Pale yellow to orange, often with strongly contrasting orange or red to brown markings; forewings often striped in appearance with pale yellow highlighting along veins of clavus and corium, and strongly coloured red to orange cuneus (Fig. 4).

Vestiture. Dorsum with moderately dense distribution of black, adpressed, lanceolate scale-like setae (Fig. 5d) on head, pronotum, proepisternum, scutellum, and hemelytra, intermixed with moderate to sparse distribution of erect, pale, simple setae (Figs 4, 5a, b, d). Black scalelike setae more clumped on hemelytron and medially along scutellum (Fig. 5b). Head: with moderately dense distribution of robust, erect, pale simple setae, intermixed with scale-like setae (Fig. 5a). Antennae: AI with sparse distribution of bristle-like, erect simple setae (Fig. 5a); AII-AIV with dense distribution of semi-adpressed, pale, simple setae. Pronotum and scutellum: with more densely distributed and elongate, pale simple setae along lateral margins; sometimes with paired row of black scalelike setae about midline, at last basally (Figs 5a, b, 7a). Legs: with moderate distribution of semi-adpressed simple setae; intermixed on tibiae with pale to dark brown bristle-like setae, and on metatibiae with rows of spinules; forefemora with more elongate bristle-like setae on ventral surface, sometimes greatly exaggerated and darkened (Fig. 6a). Abdomen: male abdomen including pygophore, parameres and proctiger with moderately dense distribution of pale simple semierect setae (Figs 6c, d, 7e, f).

Structure. Head: eyes greatly enlarged and bulbous, in dorsal view removed slightly from posterior margin of head (Figs 4, 5a, b, 7a, b); vertex short and often depressed; frons short, strongly vertical; thin pronotal collar present (Fig. 5a). Antennae: AI greater in length than interocular distance (Fig. 5a); AII > in length AIII; AIV very short. Labium: elongate, reaching at least just past hind coxae and as far as pygophore (Fig. 7c); LIII elongate, a little longer than LII, labial formula LIII > LII > LIV > LI. Pronotum: narrow anteriorly and lateral margins angulate (Figs 5b, 7a); pronotum with or without two posterior projections, varying in size from small adpressed lobes to greatly elevated and elongate conical projections (Figs 4, 5b, 7a-c). Scutellum: moderately swollen (Fig. 7b, c). Thoracic pleura: proepimeron short, mesepisternum large, metathoracic spiracle large, ellipsoid (Figs 6a, 7b); metathoracic scent gland with moderately small peritreme, evaporative area with small mushroom bodies and medial vertical fold posterior to peritreme, (Figs 6b, 7d). Hemelytra: greatly elongated, tip of abdomen not reaching beyond anterior margin of cuneus in lateral view (Fig. 7c); claval vein elevated; median flexion line short, ca. level of midpoint of claval commissure; cuneus weakly to moderately deflexed. Legs: slender and elongate; femora uniformly cylindrical and narrow (Fig. 4). Abdomen: small in relation to body/hemelytral length and width (Fig. 7c).

Male genitalia: pygophore (Figs 6c, d, 7e, f 8 --15a), conical, either small to moderately sized relative to abdominal size, conical, transverse, with pair of spinelike tergal processes, uniformly positioned dorsolaterally, left process larger than right; genital opening weakly to moderately concave, ventral margin slanted towards left paramere articulation (Figs 6d, 7f); parameres at rest, in line with ventral margin of pygophore, folded medially, left inside right, meeting more or less medially (Figs 6c, d, 7e, f, 8 --15a); left paramere (Figs 6c, d, 7e, f, 8-15c) L-shaped, expanded sensory lobe, distally acute, often spinose at apex, setose, apophysis broad, with hooked apex, either apically rounded or deeply excavated; right paramere (Figs 6d, 7f, 8-15b) elongate, C-shaped, dorsal margin sometimes expanded, toothed medially towards apex; phallotheca (Figs 6c, 7e, 8-15d, e) simple, lightly sclerotised, apex straight, open dorsally, retracted to weakly exposed at rest, without processes, apex tapered to broadly rounded; aedeagus (Figs 8 --15f, g) with two endosomal spicules, PES basally narrow, at base positioned left lateral or left dorsal relative to secondary gonopore, originates near base of secondary gonopore, base not broad, downturned medial process, bifurcate in distal third, with branches of uneven length, shortest branch canting to left, broad, mostly serrate, longest branch canting to right, narrow, mostly smooth, often with a short to elongate downturned process, smooth to serrate, DES shorter than PES, originating more distal to PES, about midpoint of secondary gonopore, basally broad, at base always positioned broadly from left lateral to left dorsal, always with a prominent keel (Figs 8-15f), sometimes with a distal process which is either smooth or serrate, DES otherwise deeply divided at base with two branches, subequal in length, leftmost canting branch almost always bifurcate distally (rarely trifurcate), mostly with serrate distal margins, rightmost canting branch spatulate-like, most often smooth.

Smaller and broader across pronotum than males, hemelytra more rounded; eyes smaller than males; ventral surface of forefemora without elongate bristle-like setae; hind femora slightly thicker, broaden slightly medially; abdomen larger, reaching tip of cuneus (Fig. 4). Female genitalia: opening of vestibulum strongly asymmetrical, spiculate to smooth (Fig. 16a, d); sclerotised rings strongly folded, elongate-ovoid, with broad spiculate apex (Fig. 16a, c); without distinct medial lobes or spiculate regions mediolaterally on dorsal labiate plate (Fig. 16a); posterior wall with inter-ramal sclerites subquadrate, free, angulate and smooth mediodistally, mediobasally spiculate and rounded, inter-ramal lobes broad, subquadrate, uniformly and densely spiculate aside from inner basal margin, attached adjacent to basolateral angle, attachment to inter-ramal sclerite narrow (Fig. 16b, e).

Acaciacapsus species that have known host plant associations are all found with species of Acacia (Table 4); five of the eight species have Acacia spp. records, with the remainder unknown. Acaciacapsus appha and A. aureolus are known from multiple species of Acacia species, whereas A. bournda and A. lolworthensis are restricted to a single Acacia species based on current collection data. Acaciacapsus emeraldensis was collected from an undetermined species of Acacia. Only A. bournda is found on bipinnate Acacia (Botrycephalae) and the remaining Acaciacapsus species with host plant records are found on phyllodinous Acacia species (Juliflorae and Plurinerves).

Knowledge of the host plant associations of land bugs in Australia is being documented more extensively over the past 20 years based on literature and specimen records (Cassis et al. 1999; Cassis and Moulds 2002; Cassis and Gross 2002, 1995; Cassis and Vanags 2006; Cassis et al. 2007; Weirauch 2007; Cassis 2008; Cassis and Symonds 2008,2011; Schuh and Pedraza 2010; Schuh and Weirauch 2010; Noack et al. 2011; Cheng et al. 2012a; Symonds and Cassis 2013). These works have revealed that land bug associations are common with plants belonging to the Rosidae and Asteridae. Cassis and Schuh (2012) demonstrated a similar pattern for Miridae on a worldwide basis. Acacia belongs to the order Fabales in the Rosid I clade of the Rosidae, and is a known host of many clades of Australian Heteroptera (Cassis and Gross 2002; Cassis, pers. obs.).

Acaciacapsus is broadly distributed in non-monsoonal regions of Australia, with the majority of species found in semi-arid and arid locations (Fig. 2), within multiple areas of endemism (Crisp et al. 1995) and IBRA bioregions (Table 4) (Department of the Environment, Water, Heritage and the Arts 2012).

Acaciacapsus is a distinctive genus in colouration, vestiture and structure. They are immediately recognised by the generally yellow to orange ground colour of the body with darker orange to red or brown highlights, black lanceolate scale-like setae intermixed with pale simple setae on the dorsum, large bulbous eyes in the males, posterior pronotal lobes (in most cases), long slender legs, elongate labium, and small abdomen. The male genitalic characters of significance that are visible externally include: two dorsolateral tergal processes on the pygophore and a mostly acute sensory lobe of the left paramere. The aedeagus of Acaciacapsus is synapomorphic and diagnostic for the genus with two endosomal spicules that are more dorsal in orientation relative to the secondary gonopore.

Features of both colouration (yellowish) and structure (large eyes) relate to the habit and habitat of these species. The yellowish colouration and host collection records from Acacia spp. in flower indicate they are associated with the flowers of the host plants, and are possible pollen feeders, as well as being cryptically coloured. In addition, Acaciacapsus spp. have been collected in light traps at night and these collection records along with their large eyes infer nocturnal behaviour.

After the genus Acacia in recognition of the host plant associations of this genus and the uncommon orthotyline colouration matching that of Acacia flowers. Capsus refers to mirid bug.

1. Posterior margin of pronotum rectilinear, without posterior projections (Fig. 5b) 2

Posterior margin of pronotum with posterior projections, sometimes greatly exaggerated (Fig. 7a) 3

2. Body orange yellow; hemelytral membrane veins and within cells without prominent red highlighting, veins almost always yellow; dorsal margin of genital opening weakly concave (Fig. 9a); apex of left paramere apophysis deeply excavated (Fig. 9c) A. appha (WA)

Body yellow with extensive red bright colouring on dorsum; hemelytral membrane and within cells with prominent red highlighting; dorsal margin of genital opening moderately concave (Fig. 15a); apex of left paramere apophysis weakly excavated (Fig. 15c) A. woodwardi (QLD)

3. Body mostly yellow-orange, without prominent red markings (Fig. 4); lobes of posterior margin of pronotum moderately to greatly exaggerated (Figs 4, 7a, b) 4

Body mostly yellow with prominent red highlighting, particularly on head and cuneus (Fig. 4); lobes of posterior margin of pronotum small to moderate in size (Fig. 4) 7

4. Body with overall brownish tinge to yellow-orange colouration; wing membrane without clear spot adjacent to posterior tip of cuneus (Fig. 4) A. lolworthensis (QLD)

Body strongly yellow-orange to orange; wing membrane with round clear spot adjacent to posterior tip of cuneus 5

5. Body uniformly orange in colour, without prominent yellow markings along corial and claval veins, with orange highlighting on wing membrane veins and in cells (Fig. 4) A. amadeus (NT)

Mostly yellow-orange with strong pale yellow highlights along veins of hemelytra and at cuneal fracture, wing membrane generally yellow-orange with slight diffusion of colour into membrane cells 6

6. Posterior pronotal projections slightly elevated, only moderate in size (Fig. 4) A. millstreamensis (WA)

Posterior projections greatly exaggerated, strongly elevated above pronotal disc at ca. 45°, and slightly pointed apically (Figs 4, 7a, b) A. aureolus (SA, WA)

7. Body slightly more slender and elongate, including cells of wing membrane, hemelytra slightly wider posteriorly (Fig. 4); left paramere with small sensory lobe and deeply excavated hook at apex of apophysis (Fig. 12c), right paramere dorsal margin smooth distally (Fig. 12b) A. emeraldensis (QLD)

Body slightly more robust, hemelytra parallel-sided equal width for entire length, cells of wing membrane with slightly rounded margin (Fig. 4); left paramere sensory lobe high with evenly rounded apophysis hook (Fig. 11c), rightparamere dorsal margin serrate distally (Fig. 11b) A. bournda (NSW)

Acaciacapsus amadeus, sp. nov.

(Figs 1, 2, 4, 8)

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Holotype. Australia: Northern Territory: Reedy Rockhole, Amadeus Basin, 24.33°S 131.58°E, 01 Jun 1962, P. Ranford, 1< (AMNH_PBI 00034218) (ANIC). Paratype. AUSTRALIA: Northern Territory: 14 km SW by S of Kulgera, 25.95°S 133.22°E, 21 Sep 1978, M. S. Upton, 1♀ (AMNH_PBI 00034217) (ANIC).

Acaciacapsus amadeus is recognised by the following combination of characters: generally more uniform orange dorsal colouration, without striped pale highlights on clavus and corium; head uniformly orange without contrasting vittae or vertex markings; cuneus slightly darker orange than corium and clavus; greatly elongate forewings, with abdomen not reaching near anterior margin of cuneus in lateral view; pronotum with moderately large and greatly elevated conical posterior projections.

Body length 5.14 mm.

Colouration (Fig. 4). Head: uniformly orange, without contrasting markings or vittae. Antennae: uniformly pale orange to yellow, sometimes with AI a brighter orange. Pronotum: disc orange; collar, callosite region and posterior projections sometimes lighter, pale orange; without contrasting markings. Thoracic pleura: uniformly light orange. Scutellum: mesoscutum and scutellum uniformly light orange, sometimes mesoscutum a little darker. Hemelytra: mostly uniformly orange to yellow orange, sometimes with faint pale stripes on claval and corial veins; cuneus concolorous with rest of hemelytron, sometimes slightly darker orange; membrane veins creamy yellow, with bounded membrane areas coloured orange, subcuneal clear spot present. Legs: pale orange to yellowish orange golden yellow, femora without any distinct, apical or basal banding; femora sometimes uniformly brighter orange than remainder of leg. Abdominal venter: uniformly orange to yellow orange.

Vestiture. Legs: forefemur with moderate distribution of elongate pale brown bristle-like setae on ventral surface.

Structure. Antennae: AII length < 1.5 x distance between humeral angles. Labium: reaching pygophore. Pronotum: posterior projections conical, moderate to greatly enlarged, strongly elevated. Hemelytra: forewings greatly elongated, tip of abdomen not reaching anterior margin of cuneus in lateral view, significantly shorter; cuneus moderately deflexed. Male genitalia: pygophore (Fig. 8a), dorsal margin moderately concave; left paramere (Fig. 8c) sensory lobe elongate in height, with acute spinose apex, two large spinules on lateral sensory lobe margin, apophysis apex deeply excavated; right paramere (Fig. 8b) apically weakly curved, with slight medial expansion of dorsal margin, continuously serrate along dorsal margin; phallotheca (Fig. 8d, e), apex rounded, widely open; aedeagus (Fig. 8e, f), PES position left dorsolateral at base, with downturned medial process smooth and slender, distal left canting branch smooth, distal right canting branch smooth, with small subapical smooth process, DES keel (Fig. 8f) with branched weakly serrate apical process; leftmost canting branch of DES basal split more proximal to rightmost canting branch than to keel (Fig. 8f), leftmost canting branch of DES basally bifurcate, with sub-branches subequal length and distally serrate, rightmost canting spatulate-like branch with margins smooth.

Body length 5.42 mm. Similar to male apart from sexually dimorphic variation as in generic description.

Measurements

See Table 5.

Host plant Unknown.

Distribution

From two localities in central Australia south-east of Alice Springs (Fig. 2).

Acaciacapsus amadeus is the only species of the genus recorded from the Northern Territory. In comparison to all other Acaciacapsus species it has the most uniform colouration and its orange hue is the most intense found in the genus.

After the region of central Australia in which the type locality is found.

Acaciacapsus appha, sp. nov.

(Figs 1, 2, 3a, b, 4, 5, 6, 9, 16a-c)

um:lsid:zoobank.org:act:12DBC3D9-9583-47BB-8164-BB4D1FF0B0AB

Holotype. Australia: Western Australia: Lochada, Omega track, N of Baker Well, 29.1412°S 116.5425°E, 271m, 17Sep2009,C. Symonds, Acacia sibina Maslin (Fabaceae), det. WA Herbarium, 1< (AMNH_PBI 00030572) (WAMP).

Paratypes. AUSTRALIA: Western Australia: Charles Darwin Reserve, track to White Dam, 29.5955°S 116.9427°E, 307m, 22 Sep 2009, C. Symonds, Acacia effusifolia Maslin & Buscumb (Fabaceae), det. WA Herbarium, 1< (AMNH_PBI 00030574), 1♀ (AMNH_PBI 00030584) (WAMP). Kadji Kadji Reserve, 29.2417°S 116.4709°E, 265 m, 15 Sep 2009, C. Young, Light Trap, 1< (AMNH_PBI 00400834) (UNSW). Kadji Kadji Reserve, CY2, 29.2315°S 116.4777°E, 266 m, 17 Sep 2009, C. Young, Light Trap, 1< (AMNH_PBI 00400605) (UNSW). Karara, N of Emu Fence on track to Corner Well, 29.1683°S 116.6663°E, 268 m, 20 Sep 2009, C. Symonds, Malleostemon tuberculatus (E. Pritz.) J.W. Green (Myrtaceae), det. WA Herbarium, 1♀ (AMNH_PBI 00400604) (UNSW). Karara, on Emu Fence, N of Mungada Rd, 29.1596°S 116.64°E, 339 m, 19 Sep 2009, C. Symonds, Acacia ramulosa W. Fitzg. var. ramulosa (Fabaceae), det. WA Herbarium, 1♀ (AMNH_PBI 00400603) (UNSW). Lochada, Omega track, N of Baker Well, 29.1412°S 116.5425°E, 271m, 17 Sep 2009, C. Symonds, Acacia sp. (Fabaceae), 1♀ (AMNH_PBI 00400597) Acacia ramulosa W. Fitzg. var. ramulosa (Fabaceae), det. WA Herbarium, 1< (AMNH_PBI 00400595), 2♀ (AMNH_PBI 00400598, AMNH_PBI 00400599) Acacia sibina Maslin (Fabaceae), det. WA Herbarium, 1< (AMNH_PBI 00400596), 2♀ (AMNH_PBI 00400600, AMNH_PBI 00400601) (UNSW), Acacia sibina Maslin (Fabaceae), det. WA Herbarium, 10♀ (AMNH_PBI 00030573, AMNH_PBI 00030575-AMNH_PBI 00030583) (WAMP). Lochada, Steves track, 29.2456°S 116.5435°E, 259m, 20 Sep 2009, C. Symonds, Acacia ramulosa W. Fitzg. var. ramulosa (Fabaceae), det. WA Herbarium, 1♀ (AMNH_PBI 00400602), 1< (AMNH_PBI 00400821) (UNSW).

Acaciacapsus appha is recognised by the following combination of characters: golden yellow-orange brown, with contrasting pale yellow veins of clavus and corium in unbroken stripes; cuneus darkened; calli orange, distinct from rest of pronotum; base of coxae brown; femora with indistinct apical orange band, and indistinct basal brown band; pronotum without posterior projections; male forefemora with dense distribution of strong, black bristle-like setae ventrally, concentrated at the base of the femur; labium not greatly elongate, reaching at least to metacoxae and no more than halfway along abdomen; tip of abdomen reaching just short of anterior margin of cuneus; male genitalia, left paramere with large sensory lobe, right paramere unexpanded medially, keel of DES without apical processes.

Body length 5.30-5.85 mm.

Colouration (Fig. 4). Head: golden yellow, with contrasting orange vittae and broad orange band across posterior of vertex. Antennae: uniformly golden yellow, AI sometimes diffusely orange. Pronotum: variable; mostly uniformly golden yellow; sometimes diffusely orange posteriorly; sometimes with pale yellow band medially from anterior to posterior margin; calli orange. Thoracic pleura: mostly yellow, with margins of metepisternum brown; brown band across metepisternum dorsally; mesepimeron and scent gland evaporative area bright pale yellow. Scutellum: mesoscutum orange, scutellum uniformly paler yellowy orange. Hemelytra: mostly golden orange, pale yellow stripes along claval and corial veins; claval vein stripe extending to medial margin post-claval commissure extending to tip of cuneus; cuneus mostly darker orange-brown with base and apex pale yellow; membrane veins pale yellow, with bounded membrane areas straw coloured or orange; translucent subcuneal spot present. Legs: uniformly golden yellow; femora with indistinct apical orange band, and indistinct basal brown band; base of coxae with brown band. Abdominal venter: yellow or yellow with darker orange markings; brown markings sublaterally in a band along sternites; male pygophore with indistinct brown spot anteroventrally and anterodorsally.

Vestiture. Legs: ventral foretibial spines, black, elongate, more densely distributed towards base of femur (Fig. 6a).

Structure. Antennae: AII length < 1.5 x distance between humeral angles. Labium: variable, reaching metacoxae to mid abdomen. Pronotum: without posterior projections (Fig. 5b). Hemelytra: forewings greatly elongated, tip of abdomen reaching just short of anterior margin of cuneus in lateral view; cuneus weakly to moderately deflexed. Male genitalia: pygophore (Figs 6a, b, 9a) dorsal margin shallowly concave; left paramere (Figs 6a, b, 9c) sensory lobe elongate in height, spinose at apex, apophysis apex deeply excavated; right paramere (Figs 6a, b, 9b) apically robust, without medial expansion of dorsal margin, broadly serrate; phallotheca (Fig. 9d, e), apex tapered, widely open; aedeagus (Fig. 9f-h), PES position left lateral at base, with downturned medial process smooth or serrate, distal left canting branch serrate, distal right canting branch smooth, with small subapical smooth process, DES keel (Fig. 9f, g) splayed distally, without any apical process, leftmost canting branch of DES basal split more proximal to rightmost canting branch than to keel, leftmost canting branch of DES trifurcate distally, with horizontalmost sub-branch arcuate, margins strongly serrate, remaining sub-branches, straight, smooth, rightmost canting spatulate-like branch with margins smooth.

Body length 5.06-5.29 mm. Similar to male apart from sexually dimorphic variation as in generic description. Female genitalia: mostly as in generic description; vestibular opening speculate; sclerotised rings with apex broad, spicules variable in size from small to large (Fig. 16a).

See Table 5.

Acaciacapsus appha is known from three Acacia spp., all of which are widespread taxa in semi-arid shrublands (Fig. 3a); Acacia effusifolia, A. ramulosa var. ramulosa and A. sibina (Fig. 3 b) are all shrubs to small trees, with slender arid-adapted phyllodes, and are distributed from the northern wheatbelt across into the Murchison district and Northern Goldfields, and to the edges of the Western Desert area of endemism of Western Australia (Western Australian Herbarium 1998). One specimen of A. appha was found on Malleostemon tuberculatus, which is a possible sitting record (see Table 4). This species was also collected from black light bucket traps.

Acaciacapsus appha was found at several localities across all the reserves surveyed during the Charles Darwin Bush Blitz in Western Australia in September 2009, including Charles Darwin Reserve, Lochada (Fig. 3a), Kadji Kadji and Karara (Fig. 2).

Acaciacapsus appha and A. woodwardi lack the bilobed pronotum that is characteristic of the other species of Acaciacapsus. Our phylogenetic analysis did not detect these as sister taxa, based on other external morphological characters and those of the male genitalia. Acaciacapsus appha is differentiated from all others in the genus by the ventral spines on the forefemora of males being most exaggerated compared to all others, the compact club-shaped right paramere, and simple keel of the DES, without elongate distal ornamentation. This species was discovered for the first time on the Charles Darwin 2009 Bush Blitz pilot survey.

This species is named from the Greek appha, meaning 'dear one'.

Acaciacapsus aureolus, sp. nov.

(Figs 1, 2, 3c, 4, 7, 10)

urn:lsid:zoobank.org:act:63A1AC3F-5E1D-4F14-AA2B-0030D7FD0257

Holotype. Australia: Western Australia: Pilbara region: 150.5 km N of Ripon Hills Rd on Marble Bar -- Newman Rd, 22.3533°S 119.9548°E, 500 m, 25 Aug 2005, G. Cassis, S. Lassau, S. and G. Carter, Acacia bivenosa DC. (Fabaceae), det. Field ID, 1< (AMNH_PBI 00030566) (WAMP).

Paratypes. AUSTRALIA: South Australia: Parachilna Gorge, Flinders Ranges, 31.1291°S 138.5151°E, 23 Aug 1974, P. B. McQuillan, 1< (AMNH_PBI 00039014) (SAMA). Western Australia: Pilbara Co.: 109 km N of Marble Bar, on Marble Bar -- Newman Rd, 22.0316°S 120.0515°E, 451 m, 25 Aug 2005, G. Cassis, S. Lassau, S. and G. Carter, Acacia bivenosa DC. (Fabaceae), det. Perth staff PERTH 7300123, 3< (AMNH_PBI 00020810-AMNH_PBI 00020812), 1♀ (AMNH_PBI 00020814) (AM). 150.5 km N of Ripon Hills Rd on Marble Bar -- Newman Rd, 22.3533°S 119.9548°E, 500 m, 25 Aug 2005, G. Cassis, S. Lassau, S. and G. Carter, Acacia bivenosa DC. (Fabaceae), det. Field ID, 1♀ (AMNH_PBI 00020815) (AM), Acacia bivenosa DC. (Fabaceae), det. Field ID, 2< (AMNH_PBI 00400588, AMNH_PBI 00400589), 2♀ (AMNH_PBI 00400592, AMNH_PBI 00400593) (UNSW), Acacia bivenosa DC. (Fabaceae), det. Field ID, 1< (AMNH_PBI 00030565), 2♀ (AMNH_PBI 00030569, AMNH_PBI 00030570) (WAMP). Charles Darwin Reserve, NE Gate, 29.4899°S 117.0848°E, 308m, 23 Sep 2009, C. Young, Light Trap, 1♀ (AMNH_PBI 00030571) (WAMP). Kadji Kadji Reserve, 29.2417°S 116.4709°E, 265m, 15 Sep 2009, C. Young, Light Trap, 1♀ (AMNH_PBI 00400833) (UNSW). Lochada, Kelly Well, 29.0819°S 116.5868°E, 281m, 17 Sep 2009, C. Symonds, Acacia coolgardiensis (Fabaceae), det. WA Herbarium, 1♀ (AMNH_PBI 00400594) (UNSW). Lochada, Mungada Rd, E of Boiada Camp, 29.1803°S 116.5099°E, 275 m, 17 Sep 2009, C. Symonds, Acacia ramulosa W. Fitzg. var. ramulosa (Fabaceae), det. WA Herbarium, 1♀ (AMNH_PBI 00030623) (WAMP).

Other specimens examined. AUSTRALIA: Western Australia: Pilbara Co.: 109 km N of Marble Bar, on Marble Bar -- Newman Rd, 22.0316°S 120.0515°E, 451 m, 25 Aug 2005, G. Cassis, S. Lassau, S. and G. Carter, Acacia bivenosa DC. (Fabaceae), det. Perth staff PERTH 7300123,1 juvenile (AMNH_PBI 00020813) (AM). 150.5 km N of Ripon Hills Rd on Marble Bar -- Newman Rd, 22.3533°S 119.9548°E, 500 m, 25 Aug 2005, G. Cassis, S. Lassau, S. and G. Carter, Acacia bivenosa DC. (Fabaceae), det. Field ID, 2 juveniles (AMNH_PBI 00400590, AMNH_PBI 00400591) (UNSW), Acacia bivenosa DC. (Fabaceae), det. Field ID, 2 juveniles (AMNH_PBI 00030567, AMNH_PBI 00030568) (WAMP). Lochada, Mungada Rd, E of Boiada Camp, 29.1803°S 116.5099°E, 275 m, 17 Sep 2009, C. Symonds, Acacia ramulosa W. Fitzg. var. ramulosa (Fabaceae), det. WA Herbarium, 1 juvenile (AMNH_PBI 04000666) (UNSW).

Acaciacapsus aureolus is recognised by the following combination of characters: pronounced striped dorsal colouration, golden yellow to yellow-orange, contrasted with pale creamy yellow stripes on hemelytron; cuneus concolorous with clavus and corium; membrane adjacent to tip of cuenus with translucent spot; pronotum with moderately large, elevated posterior projections; male genitalia, left paramere with very broad apophysis, aedeagus with spatulate branch of DES greatly elongate with coarse distal serrations.

Body length 5.27-5.77 mm.

Colouration (Fig. 4). Head: uniformly golden yellow, without contrasting markings or vittae. Antennae: uniformly golden yellow, sometimes with AIV dusty yellowish brown. Pronotum: majority including callosite region uniformly golden yellow to yellowish orange, collar and tips of posterior projections sometimes paler creamy yellow. Thoracic pleura: mostly golden yellow, with mesepimeron and evaporative areas of metathoracic glands bright yellow. Scutellum: mesoscutum mostly uniformly golden yellow; scutellum paler, mostly uniformly creamy yellow. Hemelytra: mostly golden yellow orange, with pale yellow stripes on claval vein, on endocorium adjacent to claval suture, and more broadly on region bounding median flexion line, claval vein stripe extending to medial margin post-claval commissure extending to tip of cuneus; cuneus mostly golden yellow orange with base and apex pale yellow; membrane veins pale yellow, with bounded membrane areas straw coloured, translucent, post-membrane cells with embrowned regions, translucent subcuneal spot present. Legs: uniformly golden yellow, femora with indistinct, apical and basal orange bands. Abdominal venter: uniformly yellow, pregenital sternites with sublateral brown markings; male pygophore with indistinct brown spot anteroventrally.

Vestiture. Legs: forefemora with moderate distribution of pale brown bristle-like setae.

Structure. Antennae: AII length < 1.5 × distance between humeral angles. Labium: reaching pygophore (Fig. 7c). Pronotum: posterior projections conical, moderately large to large sized, elevated (Fig. 7a-c). Hemelytra: forewings greatly elongated, tip of abdomen not reaching anterior margin of cuneus in lateral view; cuneus moderately deflexed. Male genitalia: pygophore (Figs 7e, f, 10a) dorsal margin moderately concave; left paramere (Figs 7e,f 10c), sensory lobe short in height, distally acuminate and spinose, apophysis apex evenly rounded; right paramere (Figs 7e, f, 10b) with medial expansion of dorsal margin, irregularly serrate from midpoint to apex; phallotheca (Figs 7e, 10d, e) visible at rest, apex broad, lateral margins at apex moderately recurved, more enclosing; aedeagus (Fig. 10/ g): PES position left lateral at base, with downturned medial process serrate, distal left canting branch strongly downturned, almost vertical, margins smooth, distal right canting branch smooth, with short subapical process, mostly smooth, with apex weakly serrate, DES keel (Fig. 10/) with an elongate straight process, weakly serrate distally, leftmost canting branch of DES basal split more proximal to keel than to base of rightmost canting branch, leftmost canting branch of DES bifurcate distally, with two vertical slender sub-branches, one smooth, and one distally serrate, rightmost canting spatulate-like branch with margins asymmetrically serrate.

Body length 4.83 -- 5.71mm. Similar to male apart from sexually dimorphic variation as in generic description.

Second instar nymph. Golden yellow colouration. Dorsum with sparse distribution of black erect bristle-like setae, and very short pale scalelike setae. Posterior pronotal processes present.

See Table 5.

Host plant and habitat

Acaciacapsus aureolus is associated with Acacia coolgardiensis (Fig. 3 c) and A. ramulosa var. ramulosa in the Northern Wheatbelt and A. bivenosa in the Pilbara. No host records are known for South Australian populations. This species has also been collected at black light bucket traps.

Acaciacapsus aureolus is the most widely distributed of all Acaciacapsus species, ranging from the Flinders region of South Australia to the Northern Wheatbelt and Pilbara (Fig. 2).

Acaciacapsus aureolus is a paler species, without any red highlighting or markings. It has enlarged pronotal processes in both sexes. Colouration varies across the range of this species, with the Pilbara population being bright golden yellow in colouration and the Western Australian wheatbelt and South Australian populations being more yellowish orange. This species was collected during the Charles Darwin 2009 Bush Blitz survey.

Named for the golden body colouration, from the Latin aureolus meaning golden.

Acaciacapsus bournda, sp. nov.

(Figs 1, 2, 3f g,4, 11)

urn:lsid:zoobank.org:act:994EC552-FAF6-4B6E-8922-919296CE933A

Holotype. Australia: New South Wales: Bournda National Park, North Wallagoot, Turingal Head, 36.7845°S 149.9568°E, 16 m, 20 Nov 2002, Cassis, Schuh, Schwartz, Silveira, Acacia mearnsii De Wild. [introduced] (Fabaceae), det. NSW staff NSW658198, 1< (AMNH_PBI 00016092) (AM).

Paratypes. AUSTRALIA: New South Wales: Bournda National Park, North Wallagoot,Turingal Head, 36.7845°S 149.9568°E, 16 m, 20 Nov 2002, Cassis, Schuh, Schwartz, Silveira, Acacia mearnsii De Wild. [introduced] (Fabaceae), det. NSW staff NSW658198, 4< (AMNH_PBI 00016093-AMNH_PBI 00016095, AMNH_PBI 00021243), 12♀ (AMNH_PBI 00016096, AMNH_PBI 00020799-AMNH_PBI 00020802, AMNH_PBI 00021244-AMNH_PBI 00021250) (AM), Acacia mearnsii De Wild. [introduced] (Fabaceae), det. NSW staff NSW658198, 3< (AMNH_PBI 00400577-AMNH_PBI 00400579), 4♀ (AMNH_PBI 00400580-AMNH_PBI 00400583) (UNSW).

Other specimens examined. AUSTRALIA: New South Wales: Bournda National Park, North Wallagoot, Turingal Head, 36.7845°S 149.9568°E, 16m, 20 Nov 2002, Cassis, Schuh, Schwartz, Silveira, Acacia mearnsii De Wild. [introduced] (Fabaceae), det. NSW staff NSW658198, 3;juveniles (AMNH_PBI 00020803-AMNH_PBI 00020805) (AM), Acacia mearnsii De Wild. [introduced] (Fabaceae), det. NSW staff NSW658198, 2;juveniles (AMNH_PBI 00400584, AMNH_PBI 00400585) (UNSW).

Acaciacapus bournda is recognised by the following characters: body with striped colouration, dorsum mostly cream to pale orange with contrasting reddish markings on pronotum and hemelytra; cuneus mostly strong reddish colour; head with constrasting red vittae on frons connected to red sub-triangular or banded marking down the vertex to the posterior margin of head; pronotum with small conical posterior projections, only very weakly elevated if at all; abdomen just reaching the cuneal fracture; male genitalia, left paramere with acute strongly tapered sensory lobe.

Body length 4.90-5.59 mm.

Colouration (Fig. 4). Head: cream to orange with prominent reddish vittae on frons and contiguous weakly triangular to banded marking on vertex reaching posterior margin of head. Antennae: uniformly creamy orange colour, sometimes apex of AI with reddish tinge. Pronotum: mostly creamy orange, sometimes with diffuse broad red patches sublaterally and posterolaterally. Thoracic pleura: uniformly creamy orange.

Scutellum: mesoscutum and scutellum mostly creamy orange, with lateral margins of mesoscutum diffusely red. Hemelytra: mostly creamy orange, with extensive reddish markings along veins and flexion line, stripe-like; margins of clavus diffusely red, remainder creamy orange; corial region adjacent to claval vein red (concolorous with latter), endocorium with red highlighting, with creamy stripe, exocorium mostly creamy orange, with reddish patch proximal to the corial fracture, embolium mostly reddish; cuneus mostly pale reddish-orange, bounded by creamy orange at extremities; membrane veins cream with adjacent red highlighting, membrane mostly fumose brown, a little darker about veins, with subcuneal clear spot. Legs: uniformly creamy orange; mid and fore femora with basal and apical red banding; metafemora with apical red band. Abdominal venter: mostly creamy yellow, with lateral red spotting on pre-genitalia sterna; male pygophore with two large brown spots anteroventrally on ventral surface, extending to SVIII, also with brown spot medially, sometimes with reddish highlighting on dorsal surface of genital opening.

Vestiture. Dorsum with moderately dense distribution of black, adpressed scale-like setae on head, pronotum scutellum, clavus, corium and cuneus, intermixed with sparse distribution of erect, pale, simple setae. Head: with moderate distribution of erect, pale simple setae. Antennae: AI with sparse distribution of bristle-like, erect simple setae; AII-AIV with dense distribution of semi-adpressed, pale, simple setae. Legs: with moderate distribution of semi-adpressed simple setae; intermixed on tibiae with pale to dark brown bristle-like setae, and on metatibiae with rows of spinules; forefemora with brown elongate bristle-like setae.

Structure. Antennae: AII length < 1.5 x distance between humeral angles. Labium: reaching metacoxae. Pronotum: small posterior projections conical, not greatly elevated. Scutellum: moderately swollen. Hemelytra: greatly elongated, tip of abdomen reaching cuneal fracture in lateral view; claval vein elevated; median flexion line short, ca. level of midpoint of claval commissure; cuneus weakly deflexed. Male genitalia: pygophore (Fig. 11a) dorsal margin moderately concave; left paramere (Fig. 11c), sensory lobe elongate in height, spinose distally, including apex, apophysis apex evenly rounded; right paramere (Fig. 7b) with medial expansion of dorsal margin, irregularly serrate from midpoint to apex; phallotheca (Fig. 7d, e) retracted at rest, apex tapered, widely open; aedeagus (Fig. 7f g), PES position left lateral at base, with downturned medial process serrate, distal left canting branch serrate, distal right canting branch smooth, with elongate subapical smooth process, DES keel (Fig. 11f splayed distally, with a short serrate apical process, leftmost canting branch of DES basal split more proximal to rightmost canting branch than to keel, leftmost canting branch of DES bifurcate distally, with sub-branches subvertical, margins strongly serrate, rightmost canting spatulate-like branch with margins smooth.

Body length 4.94-5.40mm. Similar to male apart from sexually dimorphic variation as in generic description.

See Table 5.

From Acacia mearnsii (Fig. 3f, g).

Acaciacapsus bournda is known only from the type locality, a location in coastal south-eastern Australia (Fig. 2).

Acaciacapsus bournda is slightly more ovoid in comparison to other Acaciacapsus species, which are all more elongate and parallel-sided. The apical serrate process on the DES of A. bournda keel is shorter than in other species of the genus. Also see remarks for A. emeraldensis.

Named after the type locality, Bournda National Park; noun in apposition.

Acaciacapsus emeraldensis, sp. nov.

(Figs 1, 2, 4, 12)

urn:lsid:zoobank.org:act:4A830A06-4DBE-4B2D-B1B2-FE47B7DF09B0

Holotype. Australia: Queensland: Emerald, 23.53°S 148.181°E, 28 Jul 1976, S. K. Waite, Light Trap, 1< (AMNH_PBI 00199334) (QDPI). Paratypes. AUSTRALIA: Queensland: Brisbane, 27.4679°S 153.0280°E, 15 Jun 1963, C. Speed, 1 sex unknown (lost abdomen) (AMNH_PBI 00037438) (QM). Brassall, 27.5917°S 152.7353°E, 14 Aug 1964, A. Terauda, 1 < (AMNH_PBI 00037434) (QM). Chinchilla, Charley's Ck, 26.6666°S 150.6333°E, no date provided, Grace Lithgow, 1♀ (AMNH_PBI 00037444) (QM). Gayndah, 25.629°S 151.626°E, 13 Apr 1963, H. A. Rose, 1< (AMNH_PBI 00037437) (QM). Victoria Point, Brisbane, 27.5892°S 153.2889°E, 02 Sep 1954, T. E. Woodward, Acacia sp. (Fabaceae), 1< (AMNH_PBI 00037435), 1♀ (AMNH_PBI 00037436) (QM).

Acaciacapus emeraldensis is recognised by the following characters: body with striped colouration, dorsum mostly cream to pale yellow with contrasting reddish markings on head, pronotum, scutellum and hemelytra; calli pale yellow without red highlighting; cuneus with bright red colouration; head with prominent reddish vittae on frons contiguous with strong triangular marking on vertex reaching posterior margin of head; abdomen reaching well short of the cuneus; small pronotal projections; labium reaching mid length of abdomen in male; male genitalia, right paramere with smooth distal margin.

Body length 5.11-5.68 mm.

Colouration (Fig. 4). Head: cream to pale yellow with prominent reddish vittae on frons contiguous with strongly triangular marking on vertex reaching posterior margin of head. Antennae: uniformly creamy yellow colour, sometimes apex of AI with reddish tinge. Pronotum: mostly creamy yellow, with red highlighting; calli without red. Thoracic pleura: uniformly creamy yellow. Scutellum: mesoscutum and scutellum mostly creamy yellow, with lateral margins of mesoscutum diffusely red. Hemelytra: mostly cream to pale yellow, with extensive patchy reddish markings bounding veins; cuneus mostly bright red, bounded by creamy yellow; margins of clavus diffusely red, remainder creamy yellow; membrane veins cream with adjacent red highlighting, with subcuneal clear spot. Legs: mostly uniformly creamy yellow; femora red banding basally reduced. Abdominal venter: mostly creamy yellow, with lateral red spotting on pre-genital sterna; male pygophore with two large brown spots anteroventrally on ventral surface, extending to SVIII, also with brown spot medially, sometimes with reddish highlighting on dorsal surface of genital opening.

Vestiture. Legs: forefemora with reduced dark brown elongate bristle-like setae.

Structure. Antennae: AII length > 1.5 x distance between humeral angles; Labium: reaching mid length of abdomen. Pronotum: small posterior projections conical, not greatly elevated. Hemelytra: greatly elongated, tip of abdomen not reaching anterior margin of cuneus in lateral view; cuneus weakly deflexed. Male genitalia: pygophore (Fig. 12a) dorsal margin moderately concave; left paramere (Fig. 12c), sensory lobe short in height, spinose apex, apophysis apex deeply excavated; right paramere (Fig. 12b) with medial expansion of dorsal margin, irregularly serrate medially, apex smooth; phallotheca (Fig. 12d, e) retracted at rest, apex tapered, widely open; aedeagus (Fig. 12f, g), PES position left lateral at base, with downturned medial process serrate, distal left canting branch serrate, distal right canting branch smooth, with elongate subapical process, mostly smooth, with apex serrate, DES keel (Fig. 12f) splayed distally, with an elongate arcuate smooth process, leftmost canting branch of DES basal split more proximal to rightmost canting branch than to keel, leftmost canting branch of DES bifurcate distally, with a horizontal serrate sub-branch, subvertical serrate sub-branch, rightmost canting spatulate-like branch with margins smooth.

Body length 4.92 mm. Similar to male apart from sexually dimorphic variation as in generic description.

See Table 5.

Acaciacapsus emeraldensis was recorded from an unidentified species of Acacia in Brisbane. It has also been collected at light.

Acaciacapsus emeraldensis is known from several localities in the Brisbane district and extends to the northern Brigalow Belt region (Fig. 2).

Acaciacapsus emeraldensis is one of the three eastern Australian Acaciacapsus species, and has a distinct reddish colouration. It can be differentiated from A. bournda and A. woodwardi by: slightly larger body size; distinctive triangular red marking on vertex contiguous with red vittae of frons; only weakly elevated (short) sensory lobe and deeply excavated apex of the apophysis in the left paramere and mostly smooth apex of the right paramere in the male genitalia. Acaciacapsus bournda, like A. emeraldensis, has contiguous red markings on the head and small posterior pronotal lobes, but the head markings of A. bournda are less distinctly triangulate and more like a band along the middle of the vertex. Acaciacapsus woodwardi has red markings along the posterior margin of the head and lacks posterior pronotal projections, and is thereby easily differentiated from A. emeraldensis.

Named after the type locality.

Acaciacapsus lolworthensis, sp. nov.

(Figs 1, 2, 3d, e, 4, 13, 16c --e) urn:lsid:zoobank.org:act:8009D403-F1A2-438E-A63F-613774893AEC

Holotype. Australia: Queensland: 3.2km S of Lolworth Homestead, 20.20930S 145.02480E, 647 m, 18 May 2006, Cassis, Barrow, Finlay, Symonds, Acacia excelsa Benth. (Fabaceae), det. RBG staff, 1< (AMNH_PBI 00037431) (QM). Paratypes. AUSTRALIA: Queensland: 3.2km S of Lolworth Homestead, 20.20930S 145.02480E, 647m, 18 May 2006, Cassis, Barrow, Finlay, Symonds, Acacia excelsa Benth. (Fabaceae), det. RBG staff, 1♀ (AMNH_PBI 00020806), 2< (AMNH_PBI 00020807, AMNH_PBI 00020808) (AM), Acacia excelsa Benth. (Fabaceae), det. RBG staff, 1♀ (AMNH_PBI 00037432), 1< (AMNH_PBI 00037433) (QM), Acacia excelsa Benth. (Fabaceae), det. RBG staff, 1< (AMNH_PBI 00400586), 1♀ (AMNH_PBI 00400587) (UNSW).

Other specimens examined. AUSTRALIA: Queensland: 3.2 km S of Lolworth Homestead, 20.2093°S 145.0248°E, 647m, 18 May 2006, Cassis, Barrow, Finlay, Symonds, Acacia excelsa Benth. (Fabaceae), det. RBG staff, 1 nymph (AMNH_PBI 00020809) (AM).

Acaciacapsus lolworthensis is recognised by the following combination of characters: mottled colouration, mostly yellowish to orange brown, contrasted with bright yellow dashed stripes on hemelytra; without subcuneal clear spot; dark brown stripe on membrane post-veins; frons with very pale orange vittae, vertex without contrasting markings; calli round and prominent; pronotal projections large and elevated; femora with faint apical brown band.

Body length 4.55 -- 4.81 mm.

Colouration (Fig. 4). Head: uniformly dusty yellowish to orange brown, frons with very pale reddish vittae, vertex without contrasting markings. Antennae: uniformly dusty yellowish brown, sometimes with AI with indistinct sub-basal orange band. Pronotum: collar bright yellow; remainder mottled, mostly yellowish to orange brown, with callosite region, midline and apices of posterior pronotal projections brighter yellow. Thoracic pleura: proepisternum mostly dusty yellowish brown, with ventral margins brighter yellow; mesobasisternum dusty yellowish brown; mesepimeron bright yellow; metepisternum mostly dusty yellowish brown, with evaporative areas bright yellow. Scutellum: mesoscutum and scutellum uniformly dusty yellowish brown, midline of scutellum paler yellow. Hemelytra: mostly yellowish to orange brown, with claval vein, two parallel regions on endocorium with contrasting, dashed bright yellow stripes; cuneus uniformly dusty yellowish brown; membrane dusty yellowish brown, with contrasting dark brown markings adjacent to membrane veins, with dark brown line projecting caudadally from junction of minor and major cells, veins bright yellow, without subcuneal clear spot. Legs: mostly yellowish brown, apex of femora with indistinct apical brown band, basally with indistinct brown spot on dorsal surface. Abdominal venter: mostly yellow, with sublateral band of medium brown, and brown patches sublaterally on ventral surface of pygophore.

Vestiture. Legs: forefemora bristle-like setae pale brown and moderately distributed on ventral surface.

Structure. Antennae: AII length > 1.5 x distance between humeral angles. Labium: reaching pygophore. Pronotum: large posterior projections conical, greatly elevated. Hemelytra: forewings greatly elongated, tip of abdomen almost reaching cuneal fracture in lateral view; cuneus moderately deflexed. Male genitalia: pygophore (Fig. 13a) dorsal margin moderately concave; left paramere (Fig. 13c), sensory lobe short in height, distally smooth, apophysis apex deeply excavated; right paramere (Fig. 13b) with medial expansion of dorsal margin, irregularly serrate from midpoint to apex; phallotheca (Fig. 13d, e) retracted at rest, apex broad, lateral margins at apex moderately recurved, more enclosing; aedeagus (Fig. 13f g), PES position left dorsolateral at base, with downturned medial process serrate, distal left canting branch serrate, distal right canting branch weakly serrate, with elongate subapical process serrate distally, DES keel (Fig. 13f with an elongate broad weakly arcuate serrate process, leftmost canting branch of DES basal split more proximal to keel than to base of rightmost canting branch, leftmost canting branch of DES bifurcate distally, with a vertical slender serrate sub-branch, and a horizontal distally serrate sub-branch, rightmost canting spatulate-like branch of DES with margins smooth.

Body length 4.80-5.05 mm. Similar to male apart from sexually dimorphic variation as in generic description. Female genitalia: mostly as in generic description; vestibular opening smooth (Fig. 16d); sclerotised rings with apex weakly tapered, spicules small (Fig. 16c); inter-ramal sclerite and lobe as in Fig. 16e.

Mottled colouration dusty yellow brown, with medium brown and brighter yellow markings. Pronotum, wing pads and abdominal tergites with short, black, scale-like setae. Posterior pronotal processes present.

See Table 5.

From Acacia excelsa (Fig. 3d, e).

From savannah region of inland north Queensland, west of Charters Towers (Fig. 2).

Acaciacapsus lolworthensis is one of the more distinctive species of the genus, with pronounced mottled colouration but without darker red markings or highlighting. It has pale orange vittae on the head in a few male specimens. It is the only species without a subcuneal clear spot on the wing membrane.

Named after the type locality on Lolworth Station.

Acaciacapsus millstreamensis, sp. nov.

(Figs 1, 2, 4, 14)

urn:lsid:zoobank.org:act:690358D4-4893-4CBC-BE22-8E277DBCB057

Holotype. Australia: Western Australia: Pilbara district. Millstream-Chichester National Park, 21.5834°S 117.0667°E, 02 Jun 1999, G.Cassis, R.Silveira, Light Trap, 1< (AMNH_PBI 00030564) (WAMP).

Paratypes. AUSTRALIA: Western Australia: Pilbara Dist. Millstream-Chichester National Park, 21.5834°S 117.0667°E, 02 Jun 1999, G.Cassis, R.Silveira, Light Trap, 1< (AMNH_PBI 00017424) (AM).

Acaciacapsus millstreamensis is recognised by the following characters: body with slightly mottled colouration, dorsum mostly yellowish orange with contrasting darker orange red markings on cuneus and diffusely across clavus and corium; head uniformly yellow; abdominal venter uniformly yellow; femora usually with narrow apical red band, basally without any darker banding or colour; pronotum with moderate sized posterior projections, weakly elevated; male genitalia, left paramere sensory lobe short and rounded, only weakly tapered distally, pygophore left tergal process slightly flattened.

Body length 5.17-5.28 mm.

Colouration (Fig. 4). Head: yellow without any contrasting reddish markings on frons or vertex. Antennae: uniformly creamy yellow colour. Pronotum: mostly orange to yellow, with red highlighting; anterior callosite region and tips of posterior projections paler cream; collar cream to yellow. Thoracic pleura: uniformly pale creamy yellow. Scutellum: mesoscutum and scutellum mostly creamy yellow, with lateral margins of mesoscutum orange. Hemelytra: mostly yellow, with extensive patchy reddish markings; cuneus mostly bright orange to red, bounded by creamy yellow; clavus and corium diffusely red, remainder yellow, veins elevated and creamy yellow; membrane veins cream with adjacent red highlighting, with subcuneal clear spot. Legs: mostly uniformly yellow; femora with red apical band, sometimes reduced, basally without red marking; Abdomen: uniformly yellow, without red and brown spots.

Vestiture. Legs: forefemora bristle-like setae pale brown and moderately distributed along ventral surface.

Structure. Antennae: AII length ca. 1.5 × distance between humeral angles; Labium: reaching past metacoxae onto abdomen, not quite reaching mid length of abdomen. Pronotum: moderate sized posterior projections, conical, slightly elevated. Hemelytra: greatly elongated, tip of abdomen not reaching anterior margin of cuneus in lateral view; cuneus weakly deflexed. Male genitalia: pygophore (Fig. 14a), dorsal margin moderately concave; left paramere (Fig. 14c) sensory lobe short in height, distally weakly serrate, apophysis apex weakly excavated; right paramere (Fig. 14b) with medial expansion of dorsal margin, irregularly serrate from midpoint to apex; phallotheca (Fig. 14d, e) visible at rest, apex broadly rounded, lateral margins at apex open; aedeagus (Fig. 14f, g), PES position left dorsolateral at base, with medial process short, weakly downturned and serrate, distal left canting branch smooth, distal right canting branch smooth, with elongate subapical process smooth, DES keel (Fig. 14f) with an elongate slightly twisted weakly serrate process, leftmost canting branch of DES basal split more proximal to keel than to base of rightmost canting branch, leftmost canting branch of DES bifurcate distally, with both sub-branches weakly serrate, and projected vertically, rightmost canting spatulate-like branch of DES with margins smooth.

Unknown.

See Table 5.

Host unknown, collected by light trap.

Known from the type locality only, in the Pilbara region of Western Australia (Fig. 2).

Acaciacapsus millstreamensis has moderate sized posterior pronotal processes and less strongly contrasting colour patterning on the dorsum, with a concolorous head and abdomen, and lacks basal markings on the femora. The head is uniformly yellow without darker coloured vittae and the abdominal venter is uniformly yellow, without red and brown spots or banding. The left paramere of this species is short, more rounded and less serrate, in comparison to any of the other Acaciacapsus species.

Named after the type locality in Millstream-Chichester National Park.

Acaciacapsus woodwardi Cassis & Symonds, sp. nov.

(Figs 1, 2, 4, 15)

urn:lsid:zoobank.org:act:E6DB3AAD-C67D-4ACD-9205-32CC6DC0F48B

Holotype. Australia: Queensland: Brisbane, 27.4679°S 153.0280°E,

10 Aug 1955, L. Jackson, 1< (AMNH_PBI 00037441) (QM). Paratypes. AUSTRALIA: Queensland: Brisbane, 27.4679°S 153.0280°E, 14 Aug 1956, T. E. Woodward, Light Trap, 1< (AMNH_PBI 00037439) (QM). Carnarvon Gorge, 25.0667°S 148.25°E, 29 May 1954, T. E. Woodward, 1♀ (AMNH_PBI 00037442) (QM).

Acaciacapsus woodwardi is recognised by the following characters: with striped colouration, dorsum mostly creamy yellow with bright red markings on head, pronotum, mesoscutum, hemelytra, and diffusely on wing membrane around veins; head creamy yellow with red vittae on frons, and red band across posterior margin of vertex, disjoined from red colouration on frons; pronotum with creamy yellow calli and medial band, without posterior projections; red markings prominent across dorsum, venter and on legs; male genitalia, left paramere with high, acutely tapered apex and bulbous apophysis, left tergal process on pygophore moderately elongate, slender, aedeagus with greatly elongate distal branch of PES, keel of DES with greatly elongate serrate process.

Male

Body length 4.82-5.02 mm.

Colouration (Fig. 4). Head: cream to pale yellow with red markings on clypeus and bucculae, prominent reddish vittae on frons; vertex mostly creamy yellow with red band across posterior margin of head, not contiguous with red markings on frons. Antennae: yellow orange, entire AI with reddish tinge. Pronotum: mostly orangish red, with medial cream yellow band; calli cream yellow; collar yellow dorsally, red laterally. Thoracic pleura: yellow orange with extensive red patches across mes-and metepisternum; mesepimeron and evaporative area of scent gland creamy yellow; propleuron red anteriorly with remaining majority creamy yellow. Scutellum: mostly creamy yellow, with anterolateral margins red; mesoscutum mostly orange red with cream band medially. Hemelytra: cream to pale yellow, with extensive reddish markings bounding veins; cuneus mostly bright red, bounded by creamy yellow; embolium and majority of clavus and corium diffusely red, with patchy yellow stripes demarcating slightly raised veins; membrane veins cream with adjacent red highlighting, with subcuneal clear spot. Legs: mostly yellowy orange, femora with strong red banding apically and basally; tibiae with slightly diffuse red banding basally and apically. Abdominal venter: yellow, with red banding across pre-genital sterna; male pygophore with two large brown spots anteroventrally on ventral surface, extending to SVIII, and reddish highlighting on dorsal surface of genital opening and proctiger.

Vestiture. Legs: forefemora bristle-like setae pale brown and moderately distributed on ventral surface.

Structure. Antennae: AII length = 1.5 × distance between humeral angles. Labium: reaching at least mid length of abdomen; LIII elongate, a little longer than LII. Pronotum: without posterior projections. Hemelytra: greatly elongated, tip of abdomen not reaching anterior margin of cuneus in lateral view; cuneus moderately deflexed. Male genitalia: pygophore (Fig. 15a) dorsal margin moderately concave; left paramere (Fig. 15c), sensory lobe elongate in height, acuminate and spinose distally, including apex, apophysis apex evenly rounded; right paramere (Fig. 15b) with medial expansion of dorsal margin, irregularly serrate from midpoint to apex; phallotheca (Fig. 15d, e) visible at rest, apex tapered, widely open; aedeagus (Fig. 15f g), PES position left lateral at base, with downturned medial process serrate, distal left canting branch weakly serrate, distal right canting branch elongate, smooth, without subapical process, DES keel (Fig. 15f) splayed distally, with an elongate serrate apical process, longer than DES branches, leftmost canting branch of DES basal split more proximal to rightmost canting branch than to keel, leftmost canting branch of DES bifurcate distally, with subbranches subvertical, margins strongly serrate, rightmost canting spatulate-like branch with margins smooth.

Body length 4.84 mm. Similar to male apart from sexually dimorphic variation as in generic description.

See Table 5.

No hosts have been recorded for this species; one specimen was collected in a light trap.

From southern Brisbane to the Brigalow Belt in south-east Queensland (Fig. 2).

Acaciacapsus woodwardi is a distinctive species, and is readily recognised by the lack of posterior projections on the pronotum and the extensive contrasting red colouration across the body.

The discontiguous red markings on the vertex of the head are in contrast to the colouration found in A. bournda and A. emeraldensis, which have red markings on the vertex that are contiguous with those on the frons.

Named after late Tom Woodward, renowned entomologist of The University of Queensland, who collected some of the specimens of this species.

We thank the Australian Biological Research Study for funding to attend the Charles Darwin Reserve Bush Blitz survey and to undertake taxonomic research on the Orthotylini. We thank Brooke Glasser and Annabel Wheeler (ABRS) for fieldwork support. We thank the following people for the loan of material: Dave Britton (AM), Beth Mantle and Tom Weir (ANIC), Christine Lambkin and Geoff Monteith (QM), Peter Hudson and Jan Forrest (SAMA) and Justin Bartlett (QDPI). We thank Hannah Mathews for the genitalic illustrations. The following members of the Cassis laboratory at UNSW are thanked for assistance: Michael Elias, Marina Cheng, Anna Namyatova, Anouk Mututantri, Rossana Silveira and Jacqueline Karras.

Note that Orthotylus eurynome Kirkaldy 1902, is removed from synonymy with O. ericetorum (Fallen 1807); Melanotrichus australianus Carvalho 1965 is transferred to Orthotylus Fieber; Cysteorracha Kirkaldy, Irianocoris Carvalho 1972, Porphyrodema Poppius 1915 and Woodwardiola Carvalho are transferred from the Orthotylini to the Austrormirini. The Australian Orthotylinae comprise 27 genera and 84 species, nested within the following tribes:

Characters given in italics, character states given in regular font. Numbers refer to character state

'A' indicates a multistate character (0/1)

Characters and character states: bold text = uncontradicated synapomorphies, regular text = contradicted apomorphies

MAP: Fig. 2. Distribution of Acaciacapsus species across Australia.

DIAGRAM: Fig. 1. Implied weights phylogeny of Acaciacapsus and four outgroup taxa. Symmetric resampling values given at nodes. Synapomorphies and contradicted apomorphies at each node are listed in Table 3.

PHOTO (COLOR): Fig. 3. Habitat and host plants of Acaciacapsus species: (a) Melaleuca / Acacia shrubland in sandy soil, habitat of Acaciacapsus appha, Lochada Reserve, Western Australia (29.1412°S, 116.5425°E); (b) one host of Acaciacapsus appha, Acacia sibina, at preceding locality; (c) host of Acaciacapsus aureolus, Acacia coolgardiensis, Lochada Reserve, Western Australia (29.0819°S, 116.5868°E); (d, e) host of Acaciacapsus lolworthensis, Acacia excelsa in savannah woodland, west of Townsville, north Queensland (20.2093°S, 145.0248°E); (f g) host of Acaciacapsus bournda, Acacia mearnsii in coastal dry sclerophyll forest, south coast of New South Wales (36.7845°S, 149.9568°E).

PHOTO (COLOR): Fig. 4. Dorsal and lateral habitus photographs of Acaciacapsus species. A. amadeus (<: AMNH_PBI 00034218); A. appha (<: AMNH_PBI 00030572;AMNH_PBI 00030573); A. aureolus (<(SA): AMNH_PBI 00039014 (<(WA Pilbara) AMNH_PBI 00030566;AMNH_PBI 00030571); A. bournda (<:AMNH_PBI 00016092;,: AMNH_PBI 00016096); A. emeraldensis (<: AMNH_PBI 00037434); A. lolworthensis (<: AMNH_PBI 00037431); A. millstreamensis (<: AMNH_PBI 00030564); A. woodwardi (<: AMNH_PBI 00037441). Scale bar= 1 mm.

PHOTO (BLACK & WHITE): Fig. 5. Acaciacapsus appha, male (AMNH_PBI 00400605): (a) head, dorsal view; (b) head, pronotum and scutellum, dorsal view; (c) head and pronotum, lateral view; (d) scale-like and simple setae on dorsum; scale bars = 0.1 mm unless marked otherwise.

PHOTO (BLACK & WHITE): Fig. 6. Acaciacapsus appha, male (AMNH_PBI 00400605): (a) thoracic pleura and forefemur, lateral view; (b) metathoracic scent gland; (c) pygophore, lateral view; (d) pygophore, ventral view. Abbreviations: Ae = aedeagus, EvA = evaporative area, LP = left paramere, ltp = left tergal process, MEp = mesepisternum, MtS = metathoracic spiracle, Pe = peritreme, PEp = proepimeron, Ph = phallotheca, RP = right paramere. Scale bars = 0.1 mm unless marked otherwise.

PHOTO (BLACK & WHITE): Fig. 7. Acaciacapsus aureolus, male (AMNH_PBI 00020812): (a) head and pronotum, dorsal view; (b) head, pronotum and thoracic pleura, lateral view; (c) body, lateral view; (d) metathoracic scent gland; (e) pygophore lateral view; f) pygophore, ventral view. Abbreviations: EvA = evaporative area, LP = left paramere, ltp = left tergal process, MEp = mesepisternum, MtS = metathoracic spiracle, Pe = peritreme, PEp = proepimeron, Ph = phallotheca, RP = right paramere. Scale bars = 0.1 mm unless marked otherwise.

PHOTO (BLACK & WHITE): Fig. 8. Male genitalia of Acaciacapsus amadeus (AMNH_PBI 00034218): (a) pygophore, dorsal view, with position of parameres in situ; (b) right paramere, dorsal view; (c) left paramere, dorsal view; (d) phallotheca, right lateral view; (e) phallotheca, dorsal view; f) aedeagus, right lateral view; (g) aedeagus, ventral view. Abbreviations: ap = apophysis, DES = dorsal endosomal spicule, DESk=keel of dorsal endosomal spicule, DS = ductus seminis, ltp = left tergal process, PES = proximal endosomal spicule, rtp = right tergal process, SG = secondary gonopore, sl = sensory lobe. Scale bars = 0.1 mm.

PHOTO (BLACK & WHITE): Fig. 9. Male genitalia of Acaciacapsus appha (AMNH_PBI 00400595): AMNH_PBI 00400821)): (a) pygophore, dorsal view, with position of parameres in situ; (b) right paramere, dorsal view; (c) left paramere, dorsal view; (d) phallotheca, right lateral view; (e) phallotheca, dorsal view; f) aedeagus, right lateral view; (g) aedeagus, right lateral view; (h) aedeagus, ventral view. Abbreviations: ap = apophysis, DES = dorsal endosomal spicule, DESk = keel of dorsal endosomal spicule, DS = ductus seminis, ltp = left tergal process, PES = proximal endosomal spicule, rtp = right tergal process, SG = secondary gonopore, sl = sensory lobe. Scale bars = 0.1 mm.

PHOTO (BLACK & WHITE): Fig. 10. Male genitalia of Acaciacapsus aureolus (AMNH_PBI 00400588): (a) pygophore, dorsal view, with position of parameres in situ; (b) right paramere, dorsal view; (c) left paramere, dorsal view; (d) phallotheca, right lateral view; (e) phallotheca, dorsal view; (f) aedeagus, right dorsolateral view; (g) aedeagus, ventral view. Abbreviations: ap = apophysis, DES = dorsal endosomal spicule, DESk = keel of dorsal endosomal spicule, DS = ductus seminis, ltp = left tergal process, PES = proximal endosomal spicule, rtp = right tergal process, SG = secondary gonopore, sl = sensory lobe. Scale bars = 0.1 mm.

PHOTO (BLACK & WHITE): Fig. 11. Male genitalia of Acaciacapsus bournda (AMNH_PBI 00400579): (a) pygophore, dorsal view, with position of parameres in situ; (b) right paramere, dorsal view; (c) left paramere, dorsal view; (d) phallotheca, right lateral view; (e) phallotheca, dorsal view; f) aedeagus, right lateral view; (g) aedeagus, ventral view. Abbreviations: ap = apophysis, DES = dorsal endosomal spicule, DESk = keel of dorsal endosomal spicule, DS = ductus seminis, ltp = left tergal process, PES = proximal endosomal spicule, rtp = right tergal process, SG = secondary gonopore, sl = sensory lobe. Scale bars = 0.1 mm.

PHOTO (BLACK & WHITE): Fig. 12. Male genitalia of Acaciacapsus emeraldensis (AMNH_PBI 00037435): (a) pygophore, dorsal view, with position of parameres in situ; (b) right paramere, dorsal view; (c) left paramere, dorsal view; (d) phallotheca, right lateral view; (e) phallotheca, dorsal view; f) aedeagus, right lateral view; (g) aedeagus, right ventrolateral view. Abbreviations: ap = apophysis, DES = dorsal endosomal spicule, DESk=keel of dorsal endosomal spicule, DS = ductus seminis, ltp = left tergal process, PES = proximal endosomal spicule, rtp = right tergal process, SG = secondary gonopore, sl = sensory lobe. Scale bars = 0.1 mm.

PHOTO (BLACK & WHITE): Fig. 13. Male genitalia of Acaciacapsus lolworthensis (AMNH_PBI 00400586): (a) pygophore, dorsal view, with position of parameres in situ; (b) right paramere, dorsal view; (c) left paramere, dorsal view; (d) phallotheca, right lateral view; (e) phallotheca, dorsal view; f) aedeagus, right lateral view; (g) aedeagus, ventral view. Abbreviations: ap = apophysis, DES = dorsal endosomal spicule, DESk=keel of dorsal endosomal spicule, DS = ductus seminis, ltp = left tergal process, PES = proximal endosomal spicule, rtp = right tergal process, SG = secondary gonopore, sl = sensory lobe. Scale bars = 0.1 mm.

PHOTO (BLACK & WHITE): Fig. 14. Male genitalia of Acaciacapsus millstreamensis (AMNH_PBI 00030564): (a) pygophore, dorsal view, with position of parameres in situ; (b) right paramere, dorsal view; (c) left paramere, dorsal view; (d) phallotheca, right lateral view; (e) phallotheca, dorsal view; f) aedeagus, right lateral view; (g) aedeagus, ventral view. Abbreviations: ap = apophysis, DES = dorsal endosomal spicule, DESk = keel of dorsal endosomal spicule, DS = ductus seminis, ltp = left tergal process, PES = proximal endosomal spicule, rtp = right tergal process, SG = secondary gonopore, sl = sensory lobe. Scale bars = 0.1 mm.

PHOTO (BLACK & WHITE): Fig. 15. Male genitalia of Acaciacapsus woodwardi (AMNH_PBI 00037439): (a) pygophore, dorsal view, with position of parameres in situ; (b) right paramere, dorsal view; (c) left paramere, dorsal view; (d) phallotheca, right lateral view; (e) phallotheca, dorsal view; (f) aedeagus, right lateral view; (g) aedeagus, ventral view. Abbreviations: ap = apophysis, DES = dorsal endosomal spicule, DESk = keel of dorsal endosomal spicule, DS = ductus seminis, ltp = left tergal process, PES = proximal endosomal spicule, rtp = right tergal process, SG = secondary gonopore, sl = sensory lobe. Scale bars = 0.1 mm.

PHOTO (BLACK & WHITE): Fig. 16. Female genitalic structures of Acaciacapsus species: (a) A. appha, ventral view of bursa copulatrix (AMNH_PBI 00400597); (b) A. appha, posterior wall (AMNH_PBI 00400597); (c) A. lolworthensis, sclerotised ring (AMNH_PBI 00400587); (d) A. lolworthensis, vestibulum; (e) A. lolworthensis, right half of posterior wall, showing inter-ramal sclerite and lobe (AMNH_PBI 00400587). Abbreviations: IRL = inter-ramal lobe, IRS = inter-ramal sclerite, SR = sclerotised ring, V = vestibulum. Scale bars = 0.1 mm.