The Australian endemic ant-mimetic seed bug genus Daerlac is revised. This paper provides a redescription of the genus Daerlac and four species: D. apicalis, D. cephalotes, D. nigricans and D. picturatus. Daerlac tricolor is synonomised with D. cephalotes. A taxonomic key to species is provided. Known distributions of D. apicalis, D. nigricans and D. picturatus are each extended beyond previously known ranges. Daerlac species are found predominantly in temperate open forest and woodlands in association with ants and eucalypts. All species are broadly distributed and there is a high degree of overlap in distributions. They are seed predators found on the ground, in leaf litter, under bark or on trunks of eucalypts, and putatively forage on post-dispersed seeds. Cladistic analysis of morphological characters finds that the four species of Daerlac form two well-supported sister-groups (D. apicalis + D. picturatus, and D. cephalotes + D. nigricans). A discussion of the distribution, biology and myrmecomorphy of the genus is provided, and the tribal placement of Daerlac and its relationship to Laryngodus are discussed.
Additional keywords: synonomy, distribution, ant mimicry, Australia.
G. Cassis and C. Symonds
Systematics of Daerlac
Myrmecomorphy is common in land bugs (Insecta : Heteroptera : Geocorisae), where it has putatively evolved independently at least 20 times in 11 different families (McIver and Stonedahl 1993; Grimaldi and Engel 2005; Cassis and Wall 2010). Its occurrence in the seed bug family Rhyparochromidae (Insecta : Heteroptera : Lygaeoidea) has received little attention, but is likely to be a common attribute in many of the tribes of the nominotypical subfamily. In the tribe Udeocorini, several genera have ant-mimetic facies for both nymphs and adults, including the Australian genera Cryptocoris Gross, Daerlac Signoret and Fontejus Stål. The genus Daerlac is the most striking of these, with species having the head strongly triangular and flattened, the costal margins of the forewing concave, and deceptive colour markings on the forewings. Tillyard (1926) proposed that Daerlac species are ant-mimetic, in both structure and movement. Aside from this observation, nothing of the biology of this group has been published to date.
Daerlac and its included species were described in the late 19th and early 20th centuries, but these taxa have received almost no attention since their description (Slater 1964; Cassis and Gross 2002). Daerlac was described by Signoret (1881) as a monotypic genus for the species D. tricolor Signoret. Kirkaldy (1908) described the new genus Vulturnia, for the inclusion of a single species, V. albonotata Kirkaldy. Bergroth (1916) synonymised it with Daerlac. The identity and species composition of Daerlac have been the subject of alternative viewpoints. Distant (1901) described Daerlac affinis Distant, but it was synonomised with D. tricolor by Scudder (1967). Dallas (1852), in his description of Rhyparochromus Dallas, included species that were subsequently transferred to four udeocorine genera - Daerlac, Euander Stål, Fontejus and Telocoris Gross. Scudder (1967) also synonymised Rhyparochromus cephalotes Dallas and R. nigripes Dallas and transferred them to Daerlac. Scudder (1962) also transferred Daerlac apicalis (Distant) and D. picturatus (Distant) from Pamera Distant to Daerlac.
This paper provides a redescription of the genus Daerlac and four species considered valid in this work, as well as erection of a new species synonymy. We also provide information on comparative morphology, distribution patterns and habitat information, and discuss myrmecomorphy in the genus. This work is part of a review of seed bug taxa of Australia, funded by the Australian Biological Resources Study (
One hundred and seventy-two specimens were examined from the collections of the Australian Museum (AM), South Australian Museum (SAMA), Queensland Museum (QM), Queensland Department of Primary Industries (QDPI), Australian National Insect Collection (ANIC), Western Australian Museum (WAM), American Museum of Natural History (AMNH) and United States National Museum (USNM). All specimens were labelled with unique specimen identifier (USI) codes, and the data entered into the Planetary Biodiversity Inventory (PBI) locality database. Specimen information can be viewed online through the Discover Life website (www.discoverlife.org, verified March 2012).
Cassis and Gross (2002) gave locality data for D. cephalotes outside the distributional ranges of the specimens examined in this study. Some of these specimens were not available for re-examination for this work and we have not included them in the distribution records or maps, although we do refer to these records in the species treatment.
Specimens were examined using Leica MZ16 microscopes. Genital illustrations were prepared on a Leica DM5000B compound microscope with camera lucida. Colour photographic images were taken using a Canon 40D digital SLR coupled with an Infinity K2 lens integrated with the Visionary Digital microphotography system (www.visionarydigital.com, verified March 2012). Multiple images of each subject were composited using Helicon Focus software (Kozub et al. 2008). A Hitachi TM3000 was used to take the scanning electron micrographs. The distribution maps were prepared using ArcMap 9.3.1 (ESRI 2009). The major ecoregions of Australia identified by the World Wide Fund for Nature have been imposed on the distribution maps for Daerlac; the ecoregions are based on Interim Biogeographic Regionalisation of Australia (IBRA) bioregions (DEWHA 2010). The 300 mm isohyet depicting average rainfall data was also imposed on the distribution maps and obtained from BIOCLIM data (Busby 1991).
Eight morphological character measurements were made using a digital micrometer, focusing on characters with perceived interspecific variation. The characters measured were body length, head width, interocular distance, pronotal length, anterior pronotal lobe length and width, and posterior pronotal lobe length and width. A maximum of five male and five female specimens were measured for each species, where sufficient specimens were available.
The phylogenetic relationships of all four Daerlac species were analysed using morphological data. A total of 28 morphological characters (Table 1) were coded and are given in the data matrix (Table 2). These comprised 13 external structural, 10 colouration and five male genitalic characters. One outgroup taxon from the tribe Myodochini, Remaudieriana inornata (Walker), was used to root the tree. Five outgroup taxa from the tribe Udeocorini were also used: Fontejus westraliensis Gross, Udeocoris rolandi (Distant), Laryngodus australiae Herrich-Schaeffer, Laryngodus cervantes Slater, Schuh, Cassis, Jonhson & Pedraza-Peñalosa and Euander lacertosus (Erichson). No male specimens of D. picturatus or F. westraliensis were available for study and as a consequence the five male genitalic characters are coded as unknown for these taxa. The morphological dataset was analysed in TNT (Goloboff et al. 2003) using maximum parsimony, with 10 000 random stepwise addition replicates and tree bisection-reconnection (TBR) branch swapping. To determine branch support, bootstrap values were calculated, also using TNT, with 10 000 random addition replicates and TBR branch swapping. All characters were unweighted and multistate characters were unordered. Unambiguous synapomorphies are shown on the resulting phylogeny using MacClade 4.0.8 (Maddison and Maddison 2005).
No. Character and states 1. Forewing colouration - corium orange in part: (0) absent; (
Taxa Characters 0 0 0 0 0 0 0 0 0 1 1 1 1 1 1 1 1 1 1 2 2 2 2 2 2 2 2 2 1 2 3 4 5 6 7 8 9 0 1 2 3 4 5 6 7 8 9 0 1 2 3 4 5 6 7 8 Outgroups Euander lacertosus 0 0 1 0 0 0 0 - 0 0 0 0 0 1 0 2 0 1 0 0 0 0 - 0 2 2 0 Fotentejus westraliensis 0 0 0 - - - 0 - 0 0 0 0 0 3 0 1 0 0 2 ? ? ? ? ? 0 2 2 0 Laryngodus australiae 0 0 2 0 0 0 0 - 0 0 0 2 0 3 0 2 1 1 1 2 0 0 - - 1 1 0 1 Laryngodus cervantes 0 0 2 0 0 0 0 - 0 0 0 2 0 3 0 2 1 1 1 2 0 0 - - 1 1 0 1 Remaudieriana inornata 0 0 2 0 0 0 0 - 0 0 0 0 0 1 1 1 1 2 0 1 1 0 - - 1 2 2 1 Udeocoris rolandi 0 0 2 0 0 0 0 - 0 0 0 0 0 0 0 1 0 0 0 0 2 0 - 0 1 2 0 Daerlac D. apicalis 0 1 1 1 1 0 1 1 1 0 1 1 1 1 1 1 1 0 2 0 1 1 0 1 3 0 1 1 D. cephalotes 1 1 0 1 0 ? 1 0 0 2 1 1 1 2 1 1 1 0 2 1 1 1 1 0 2 1 1 1 D. nigricans 0 1 0 1 0 1 1 0 0 2 1 1 1 2 1 1 1 0 2 1 1 1 1 0 2 1 1 1 D. picturatus 1 1 1 1 1 0 1 1 1 0 1 1 1 1 1 1 1 0 2 ? ? ? ? ? 3 0 ? 1
In this work we recognise that colour characters may have phylogenetic signal, which has been recently addressed in insects more generally as worthy of investigation (Areekul and Quicke 2006). Colour characters have been used elsewhere in heteropteran systematics (Cassis and Silveira 2001, 2002; Gelastocoridae: Nerthra Say).
The following abbreviations are used in this work: first to fourth antennal segments, AI, AII, AIII and AIV; first to fourth labial segments, LI, LII, LIII and LIV; anterior pronotal lobe, APL; posterior pronotal lobe, PPL; and third to eighth abdominal sternites, SIII-SVIII.
Phylogenetic analysis resulted in one most parsimonious tree, 49 steps long (CI = 0.82, RI = 0.857), with strong bootstrap supports (over 85%) at all nodes (Fig. 1). Daerlac forms a monophyletic group with bootstrap support of 99%, and all examined Udeocorini taxa are grouped together relative to the myodochine outgroup taxon, with 100% bootstrap support. The tree shows two clades of Daerlac: D. apicalis + D. picturatus and D. cephalotes + D. nigricans. Multiple synapomorphic characters were found to support all clades, with nine supporting the genus and two and three each supporting the two ingroup clades of sister species (Fig. 1).
The eight nonhomoplasious synapomorphies for Daerlac are: cuneus-like apex of the exocorium (2-1), forewing membrane with blocked colouration (rather than striate or patchy) (4-1), pale colour banding on the female abdominal sternite IV (7-1), dense silvery setae sub-laterally on the abdominal sternite IV (males and females) (11-1), head shape being sub-quadrate, as wide as long (13-1), explanate post-antenniferous area (14-1), anteriorly angled but not strongly curved SIV-SV abdominal suture (20-1) and base of parameres with a dorsal and ventral flange on mesial surface (with rounded margins), separated by a narrow groove (24-1). The one homoplasious synapomorphy for Daerlac is a weak ridge or midline on the ventral surface of the abdomen along SIII to SV (21-2).
The D. apicalis + D. picturatus clade is supported by 92% bootstrap value and three nonhomoplasious synapomorphies: an apical yellow spot on the forewing membrane (5-1), pale colouration of the lateral margin of abdominal sternite IV (9-1) and relatively short ovipositor (28-0).
The D. cephalotes + D. nigricans clade is supported by 87% bootstrap value and two nonhomoplasious synapomorphies: pale colouration of the anterior half or entire lateral margin of abdominal sternite V (10-2), and the longer anterior pronotal lobe (approximately twice length of posterior pronotal lobe) (15-2).
Daerlac Signoret, 1881: clviii (gen. nov.; type species: Daerlac tricolor Signoret, 1881 by monotypy). - Lethierry & Severin, 1894: 196 (catalogue); Slater, 1964: 1058-1059 (catalogue); Slater & O'Donnell, 1995: 141 (catalogue); Cassis & Gross, 2002: 344 (catalogue). Vulturnia Kirkaldy, 1908: 776 (gen. nov.; type species: Vulturnia albonotata Kirkaldy, 1908). - Bergroth, 1916: 221 (synonymy).
Daerlac is characterised by the following combination of characters: ant-mimetic; moderately sized, male body length 5.56-7.44 mm, female body length 6.14-8.17 mm; head subquadrate, porrect, post-antenniferous region explanate; antennae elongate, AI short; labium reaching at least to mesosternum, LII equal to LIII; pronotum strongly divided, anterior lobe globose, sides rounded, 1.5× to 2× length of posterior lobe; metapleuron anterodorsally convex; forecoxae with small spine on lateral surface; forefemur with two rows of spines, row on anteroventral surface only very slightly larger than those on posteroventral surface; tarsal segment I longer than segments II and III combined; hemelytra with fully developed membrane; abdominal SIV/SV suture incomplete, angled anteriorly, without strongly curved dorsal end; dense setiferous punctations on head, pronotum and hemelytra; vestiture mixed, dense distribution of short pale setae interspersed with elongate erect setae; posterior half of posterior pronotal lobe with matte surface texture, glabrous; abdominal sternite IV with denser, paler, longer, simple setae; colouration mostly dark brown to black, sometimes with orange and red-brown legs and patterning on clavus and corium, apex or subapex of corium with cream-coloured wedge, wing membrane uniformly dark brown or bicoloured, cream and dark brown; and female abdominal sternite IV with pale band of colour.
Colouration (Figs 2, 3). Generally dark brown to black or reddish brown; head, most of pronotum, thoracic pleura and sterna, and abdomen fuscous; hemelytra bi- or tricoloured, with cream wedge-shaped marking in apical portion of corium, membrane unicolourous dark brown or bicoloured with cream patches basally or distally; legs and antennae concolourous with fuscous body or lighter red to orange-brown (at least in part).
Surface and vestiture. Body polished and shiny; posterior half of posterior pronotal lobe and often hemelytra, with matte, velvet-like texture (Fig. 2); head and anterior pronotal lobe densely rugopunctate (Fig. 5A, B); anterior half of posterior pronotal lobe densely rugopunctate, posterior half of posterior pronotal lobe with moderate distribution of setiferous punctures (Fig. 5B); scutellum mostly rugopunctate and posterior third more granular; clavus with three rows of deep setiferous punctures; endocorium with two rows of setiferous punctures; exocorium with deep, setiferous punctures, evenly distributed, not patterned into rows; ventral surface of head and thoracic pleura and sterna moderately rugopunctate; abdominal venter mostly smooth, shiny, polished; body mostly densely setose, with dense distribution of short, adpressed, weakly flattened, pale, simple setae interspersed with elongate, erect, bristle-like, pale to medium brown simple setae (Figs 2, 3, 5, 6); short simple setae absent from posterior half of posterior pronotal lobe and hemelytra, except for setiferous punctures; elongate, erect simple setae more densely distributed on head, pronotum, scutellum and legs, less densely distributed on abdomen and absent from hemelytra; antennae with dense distribution of very short, fine, adpressed, pale simple setae, interspersed with sparse distribution of moderately short, bristle-like, light brown simple setae (Fig. 7A-C); legs with ventral margin of foretibiae and tarsi with dense distribution of moderately short, light brown spine-like setae; distal half of mesotibiae and distal two-thirds of metatibiae with widely separated row of moderately elongate, semi-erect, medium to dark brown-coloured spines; abdominal venter as rest of body, also with slightly denser patterned areas of paler silvery setae sublaterally on abdominal SIV (Fig. 3).
Structure. Ant-mimetic; body robust, parallel sided; submacropterous. Head: subquadrate, triangular anteriorly, porrect, somewhat flattened (Fig. 5A, C, D); eyes moderately large, removed from anterior margin of pronotum by width of second antennal segment; vertex plus frons weakly convex; post-antenniferous area of frons explanate (Fig. 5C); jugal region conspicuous, explanate; clypeus flat; bucculae minute, oval, explanate; gula pronounced, weakly convex. Antennae: two-thirds length of body; AII weakly arcuate; AIV weakly fusiform; AII ≥ AIV AIII AI; rarely oligomeric (Figs 2, 7A-C). Labium: moderately elongate, at least reaching mesocoxae; LI reaching middle of eyes; LII equal length to LIII; segment IV short; LII = LIII < LI < LIV. Pronotum: bipartite, deeply transversely, divided at two-thirds of length, division depressed (Fig. 5B, D); anterior lobe strongly rounded, globose, lateral margins rounded; posterior lobe subtrapezoidal, moderately convex, lateral margins rounded; humeral angles rounded; posterior margin weakly concave. Scutellum: triangular; slightly longer than wide; moderate size; sculpted with a depressed concavity one-third along from anterior margin, then medially tumid, with weakly defined median carina along posterior half; apex pointed. Thoracic pleura and sterna: propleuron conspicuous, convex; propleural suture obscure; proxyphus ring-like and elongate; mesopleuron subquadrate, flat; metapleuron convex anterodorsally; external efferent system of metathoracic scent gland large (Fig. 5E); evaporative areas, covering half to two-thirds of metepisternum (Fig. 5E), weakly rugulose, extending to posterior margin of mesepisternum, with narrow evaporative area reaching along entire posterior margin (Fig. 5E, F); peritreme raised, polished, tongue-like, short, terminating in line with dorsal end of suture of mesosupracoxal lobe (Fig. 5E, G). Legs: coxae globose, meso- and metacoxae excavated on mesial surface; forecoxae weakly separated, with single lateral spine (Fig. 5H); trochanters enlarged, triangular (Fig. 5H); forefemora greatly enlarged, fusiform, ventral margin with two rows of small teeth, with three larger moderately sized spines on anteroventral row and two or three larger moderate spines on posteroventral row (Figs 5H, 7D-H); foretibiae arcuate, expanded apically as part of tibial comb (Fig. 5H); mesocoxae separated by two times width of labium, globose, unarmed; mesofemora subequal in length to forefemora, not expanded, weakly fusiform; mesotibiae straight, cylindrical; metacoxae large, moderately separated, unarmed; metafemora elongate, ∼1.5× longer than mesofemora, weakly fusiform; metatibiae elongate, cylindrical, longer than metafemora; pretarsus with elongate setiform parempodia and prominent lamellate pulvilli, claws evenly rounded (Fig. 6A); first tarsomere elongate, a little longer than second and third tarsomeres combined; second tarsomere weakly lobate; third tarsomere expanded distally. Abdomen: tergites T4 and T5 posteriorly expanded, subtriangular (Fig. 8A); inner laterotergites present T3-T6 (Fig. 8A); dorsal abdominal glands absent (Fig. 8A); sternites SIII and SIV large; SIV/SV suture arcuate, directed anteriorly, incomplete (Figs 6B, D, 8B); other sutures on sternum straight and complete, except for SIII/SIV suture just incomplete (Figs 6B, C, 8B); abdominal SV and SVI with single anterolateral trichobothrium, SV-SVII with paired posterlateral trichobothria, posteriad to spiracle (Figs 6D, 8B); SIII and SIV with three medial trichobothria either side of midline; midline of abdomen from SIII to SV with weakly defined ridge/line, more pronounced on SIII. Hemelytra: lateral margins weakly excavate; claval commisure short; membrane veins obscure. Male genitalia: pygophore small, subspherical, posteroventral surface excavated before apex, posteriorly truncate (visible in lateral view), retracted within abdominal segment VII (Fig. 9A, B); genital opening dorsal, subrectangular, anterior margin width subequal to posterior margin, anterior margin concave, posterior margin concave to straight and slightly upturned (visible in lateral view), with lobe-like lateral processes anterior to paramere insertions (Fig. 9B); parameres sickle-shaped, sensory lobe small with rounded bulge on outer margin, arcuate dorsally towards base forming a curved ridge on dorsal surface, mesioproximal margin with small weakly expanded rounded flanges on dorsal and ventral surfaces, dorsal flange bilobed, narrow groove formed between two flanges, apophysis strongly curved, compressed with slight ridge along mesial margin, apex acute (Fig. 9C); endosoma with prominent helicoid process, gonoporal process with short base and five to six rings distally, ejaculatory wings directed distally away from base of endosoma, ejaculatory body strongly curved ventrally towards base at junction of ductus seminis and ejaculatory reservoir; paired, linear sclerites present, either side of the gonoporal process, originating adjacent to ejaculatory body and reaching base of coiled portion of gonoporal process (Fig. 9D, E).
As for the male with the following differences.
Colouration. Abdominal venter with cream to light brown band across abdominal SIV, extent of light coloured band varying from medial region to across entire sternite (Figs 3, 4B).
Vestiture. Patterned areas of paler, silvery setae mediolaterally on abdominal SIV, less dense than in male (Fig. 3).
Structure. Slightly larger body size (Figs 2, 3). Thorax: pronotum with anterior lobe more swollen, wider and slightly more elongate in relation to the posterior lobe than in males (Figs 2, 3; see Table 3), posterior lobe also slightly wider than head than in males. Abdomen: wider, more swollen medially than in males; abdominal venter with SIV large to greatly enlarged, one-quarter to one-third length of abdomen (Fig. 4B); SVII tapered, slightly longer than wide, or rounded, as wide as long (Fig. 4B); ovipositor moderate length, either slightly less than or greater than one-third abdomen length (Fig. 4B). Female genitalia: spermatheca moderately elongate; with weakly clavate-shaped bulb, smoothly rounded at apex; with irregular and overlapping coils; basal portion of tube moderately short, tube only slightly narrower than subapical coiled part of spermatheca (Fig. 12).
D. apicalis Males ? Mean 5.89 1.21 0.65 1.33 0.82 0.51 1.05 1.33 s.d. 0.28 0.02 0.02 0.04 0.04 0.02 0.04 0.02 Min. 5.56 1.17 0.63 1.29 0.77 0.49 1.00 1.30 Max. 6.26 1.24 0.67 1.38 0.87 0.54 1.08 1.35 n = 5 Females ? Mean 6.69 1.35 0.75 1.48 0.94 0.54 1.19 1.55 s.d. 0.07 0.02 0.01 0.06 0.07 0.06 0.04 0.05 Min. 6.62 1.32 0.73 1.37 0.86 0.48 1.15 1.47 Max. 6.80 1.37 0.77 1.52 1.04 0.62 1.24 1.60 n = 5 D. cephalotes Males ? Mean 7.15 1.44 0.82 1.60 1.10 0.50 1.25 1.49 s.d. 0.17 0.06 0.08 0.08 0.09 0.04 0.01 0.03 Min. 6.99 1.37 0.75 1.49 1.03 0.46 1.24 1.44 Max. 7.44 1.53 0.90 1.69 1.22 0.57 1.27 1.52 n = 5 Females ? Mean 7.62 1.50 0.82 1.82 1.20 0.62 1.36 1.63 s.d. 0.33 0.08 0.05 0.16 0.07 0.10 0.05 0.13 Min. 7.36 1.40 0.78 1.61 1.10 0.51 1.30 1.49 Max. 8.17 1.63 0.91 2.04 1.27 0.78 1.43 1.76 n = 5 D. nigricans Males ? Mean 6.84 1.45 0.78 1.60 1.07 0.53 1.22 1.42 s.d. 0.35 0.06 0.02 0.10 0.12 0.03 0.08 0.10 Min. 6.24 1.36 0.76 1.42 0.88 0.49 1.09 1.26 Max. 7.13 1.51 0.81 1.69 1.20 0.56 1.27 1.54 n = 5 Females ? Mean 8.08 1.66 0.93 1.78 1.20 0.57 1.44 1.71 s.d. 0.09 0.02 0.03 0.06 0.05 0.02 0.02 0.02 Min. 7.93 1.63 0.90 1.68 1.13 0.54 1.42 1.68 Max. 8.16 1.68 0.96 1.83 1.25 0.60 1.47 1.73 n = 5 D. picturatus Females ? Mean 6.50 1.25 0.68 1.41 0.86 0.56 1.12 1.44 s.d. 0.26 0.09 0.07 0.15 0.08 0.08 0.10 0.11 Min. 6.14 1.15 0.61 1.29 0.78 0.48 0.99 1.31 Max. 6.74 1.34 0.75 1.58 0.96 0.63 1.24 1.56 n = 4
Daerlac is distinguished from all other udeocorines by its strong ant-mimetic facies, broad and flattened triangular head, arcuate second antennal segment, strongly globose or convex pronotum, cream-coloured cuneus-like colour patterning of the apex of the exocorium, cream-coloured basal half of the embolium, pale colouration on the female abdomen, and dense silvery setae laterally on the abdomen. Other putative ant-mimetic features include the rugopunctate texture of the head and thorax, smooth texture of the abdomen, and the elongate erect setae. Many ants, including species of the eucalypt seed-foraging genus Rhytidoponera Mayr (Bashford 1993), possess these texture and vestiture characters.
The major differences between species of Daerlac are those of colouration and size. In particular, colour differences are the most reliable characters in diagnosing Daerlac species. Minor structural differences between species include those of body size, size and proportions of the pronotal lobes, female abdominal shape distally, and size of female abdominal SIV. In general, we found that genital characters are of little use in distinguishing between species of Daerlac, with only minor variations found. The male endosoma is indistinguishable between species, but the parameres and pygophore differ slightly between some species. In the female genitalia the length of the ovipositor varies, but the spermatheca exhibits no noticeable discontinuities.
D. apicalis (Distant, 1904) ACT, NSW, NT, QLD, VIC
D. cephalotes (Dallas, 1852) ACT, NSW, QLD, SA, TAS, VIC
D. nigricans (Distant, 1918) ACT, NSW, QLD
D. picturatus (Distant, 1904) QLD, SA, WA
1. Wing membrane with apex pale light brown (Fig. 2); female abdominal SIV with thick cream band across full width of abdomen (Fig. 4B)
Wing membrane with apex dark brown, sometimes pale cream at base of membrane (Fig. 2); female abdominal SIV with thin cream band across medial part of abdomen (Fig. 4B)
2. Clavus and corium of hemelytra mostly dark brown (Fig. 2); legs and antennae uniformly dark brown (Figs 2, 3), AI with light brown spot on outer margin at apex (Fig. 3); ACT, NSW, NT, QLD, VIC D. apicalis (Distant, 1904)
Clavus and corium of hemelytra mostly orange-brown (Fig. 2); legs mostly orange-brown, tibiae and tarsi sometimes red-brown (Fig. 3); antennae light brown basally, AI uniformly light brown (Figs 2, 3); QLD, SA, WA, D. picturatus (Distant, 1904)
3. Antennae and legs uniformly dark brown (Figs 2, 3); corium dark brown and cream (Fig. 2); wing membrane base light creamy brown, apex dark brown, diffusion between two colours (Fig. 2); ACT, NSW, QLD D. nigricans Distant, 1918
Antennae lighter basally (Figs 2, 3); legs mostly orange, forefemora medially darker red-brown (Fig. 3); corium tricoloured, orange-brown, dark brown, cream (Fig. 2); wing membrane mostly uniformly dark brown, sometimes light brown to cream at base adjacent to corium (Fig. 2); ACT, NSW, QLD, SA, TAS, VIC D. cephalotes (Dallas, 1852)
(Figs 2, 3, 4B, 7A, D, 9, 12C, 13A)
Pamera apicalis Distant, 1904: 483 (sp. nov.). Daerlac apicalis: Scudder, 1962: 766 (comb. nov.). - Slater, 1964: 1059 (catalogue); Scudder, 1967: 257 (subsequent designation); Slater & O'Donnell, 1995: 141 (catalogue); Cassis & Gross, 2002: 345 (catalogue).
Lectotype. ♂, AUSTRALIA: Queensland: Townsville [BMNH].
Other material examined. AUSTRALIA: Australian Capital Territory: Paddy's River, 35.45861°S 149.03972°E, 633 m, 23 Apr 1968, E. B. Britton, 1♀ (33793) (ANIC); Weston Creek, Cotter Rd, 35.31666°S 149.01666°E, 610 m, 06 Apr 1969, E. B. Britton, 1♂ (33790) (ANIC); New South Wales: 30 mi. W of Junee, 34.86666°S 147.13333°E, 05 Apr 1969, G. B. Monteith, 1♀ (201315) (QM); Bogan River, J. Armstrong, 3♂ (18020 - 18022), 1♀ (18023) (AM); Darling River, 1.5 km S of 'Trilby' Stn homestead, 30.65116°S 144.9335°E, 01 Dec 1999 - 21 Dec 1999, Christie, Flemons and Elliott, 1♀ (18025) (AM); Deniliquin, 35.51666°S 144.93333°E, 1926, W. W. Froggatt, 1♂ (33789) (ANIC). Northern Territory: 4 mi. W of Coolibah HS, 15.34°S 130.54°E, 17 m, 20 Jun 1968, M. Mendum, 1♂ (33791) (ANIC). Queensland: Cloncurry-Julia Creek, 20.7°S 140.5°E, 21 Aug 1963, T. E. Woodward, 1♂ (201316) (QM); Acacia Ridge, Brisbane, 27.58946°S 153.02671°E, 03 Mar 1965, E. C. Dahms, 1♀ (201313) (QM); Dunwich, North Stradbroke Island, 27.5°S 153.4°E, 02 May 1972, G. B. Monteith, 1♀ (201314) (QM); Nudgee, 27.371°S 153.09°E, 5 m, Jan 1924, H. Hacker, 1♀ (201357) (QM); 50 [mi] S Ayr, 20.3°S 147.43333°E, 11 Sep 1950, E. F. Riek, 1♂ (33792) (ANIC); Boondall Wetlands, site 1, 27.33683°S 153.0712°E, 22 Apr 2003, C. J. Burwell, 1♂ (201309), 2♀ (201307, 201308) (QM); Bulimba Ck, Carindale, 27.33666°S 153.11°E, 16 Apr 2003, C. J. Burwell, 1♂ (201312), 1♀ (201311) (QM); 03 Nov 2003, QM party, 1♀ (201310) (QM); Carnarvon Range, 25.28333°S 148.73333°E, Feb 1944, N. Geary, 1♀ (18024) (AM); Eidsvold, 25.37277°S 151.12277°E, 193 m, ca. 1900, unknown collector, 1♂ 1♀ (18027) (AM); Gayndah, 25.629°S 151.626°E, Jan 1935, unknown collector, 1♂ 1♀ (199280) (QDPI); Warwick, 200 m W Wildash Sch., 28.26666°S 152.1°E, 30 Nov 1997, R. Eastwood, A. McArthur, 1♂ (38863), 1♀ (38864) (SAMA). Victoria: Chiltern Forest, 36.12555°S 146.62333°E, 273 m, Feb 1967, R. S. McInnes, 1♂ (33795) (ANIC); Little Desert National Park, Horseshoe Bend, 36.5°S 142.0166°E, 06 Jul 1982, M. Harvey and B. Roberts, ex. Eucalyptus camaldulensis (Myrtaceae), 1♀ (33794) (ANIC); Werribee Plains, 37.9°S 144.65°E, Jul 1941, C. Le Soeuf, 1♀ (18026) (AM). Western Australia: Kalbarri, 27.71°S 114.165°E, 28 Sep 1985, R. P. McMillan, 1♂ (30364) (WAM). Palm Springs, 15 km W of Millstream, 21.55°S 116.96666°E, 16 Jun 1984, R. P. McMillan, 1♀ (30365) (WAM).
Daerlac apicalis is distinguished by the following combination of characters: moderate size, mostly dark colouration; antennae uniformly dark brown, AI with light brown spot on lateral margin at apex; most of dorsum and venter dark brown to black; corium bicoloured with cream-coloured wedge before apex, apex of corium dark brown to black; wing membrane dark brown, apex yellow, sometimes also with small yellow spot adjacent to apex of corium; abdomen with cream band along entire lateral margin of abdominal SIV; female abdominal SIV with cream band across majority of sternite, extending to lateral margins, abdominal SIII entirely dark brown; legs and, supracoxal lobes uniformly dark brown.
Colouration (Figs 2, 3). Head: black; labium dark brown. Antennae: dark brown; apex of AI on outer side, light brown or cream spot (Fig. 3). Thorax: mostly black, or very dark brown; posterior ridge of pronotum dark brown, concolourous with rest of pronotum; scutellum black, tip of apex cream; supracoxal lobes concolourous with most of thorax. Legs: generally uniformly dark brown; trochanters and tarsi often slightly lighter brown; coxae dark brown, concolourous with most of legs and thorax. Hemelytra: setiferous punctuations dark brown; clavus dark brown, often lighter brown with cream streaky highlights; corium basal half same as clavus, apical half dark brown with subapical cream wedge, apex dark brown; cream wedge on corium sometimes medially suffused with dark brown patch; embolium, basal half cream, apical half concolourous with adjacent part of corium; wing membrane bicoloured, mostly dark brown, apex yellow often with small yellow spot adjacent to apex of corium. Abdomen: abdomen mostly dark brown; abdominal SIV sometimes slightly lighter than rest of abdomen, with cream band along entire lateral margin; lateral margins of SV and SVI concolourous with rest of abdomen, lateral margins of SVII sometimes light brown.
Structure. Moderate size, body length 5.56-6.26 mm. Thorax: anterior lobe of pronotum ∼1.61× longer than posterior lobe, posterior lobe ∼1.25× wider than anterior lobe and 1.10× wider than head. Male genitalia: pygophore (Fig. 9A, B), with lobe-like lateral processes on genital opening rounded (Fig. 9B); bilobed dorsal flange at base of parameres with lobes narrowly separated, distal lobe smaller than lobe of ventral flange, basal groove between dorsal and ventral flanges moderately narrow, visible (Fig. 9C); aedeagus (Fig. 9D, E).
As for male except for the following characters.
Colouration. Female abdominal SIV with cream band across majority of sternite, extending to lateral margins (Figs 3, 4B); SIII entirely dark brown (Figs 3, 4B); lateral margins of SVII sometimes light brown and extending to apex of abdomen (Fig. 3).
Structure. Larger than male, body length 6.62-6.80 mm. Thorax: Pronotum, anterior lobe ∼1.75× longer than posterior lobe, posterior lobe ∼1.3× wider than anterior lobe and ∼1.15× wider than head. Abdomen: SIV greatly enlarged, one-third abdomen length; SVII rounded, as wide as long; ovipositor less than one-third abdomen length (Fig. 4B). Female genitalia: spermatheca (Fig. 12C).
See Table 3.
Daerlac apicalis is distinguished from all other species in the genus by the unique light brown spot on the lateral margin at the apex of the first antennal segment, and the colouration of the clavus and corium. This species is best differentiated from D. cephalotes and D. nigricans by its smaller size, light yellow spot at the apex of the wing membrane and the cream colouration along the entire lateral length of abdominal SIV. On the basis of these latter characters, Daerlac apicalis is similar to D. picturatus, but may be distinguished from D. picturatus by its slightly larger size, more robust pronotum, and uniformly dark brown legs and antennae (aside for the light brown apicolateral spot on AI), and lack of any orangey brown colouration on the pronotum and hemelytra. As in all Daerlac species there is some variation in the extent of the cream banding across the female abdomen. In D. apicalis this varies from being just a thin band across the posterior margin of abdominal SIV to sometimes extending to the anterior margin, medially on abdominal SIV. However, this cream band always extends entirely across abdominal SIV to the lateral margins.
Daerlac apicalis is predominantly distributed in eastern Australia from Townsville to Melbourne. It is found in coastal to montane areas, and also extends to semi-arid areas of central to western New South Wales and western Queensland (Fig. 13A). The species also has three disjunct records, one from tropical northern Australia and two from near the coast in central Western Australia. Although the widespread distribution of this species results in broadscale sympatry with D. cephalotes and D. nigricans, D. apicalis is largely absent from eastern New South Wales, where the other two species are more abundant and co-occur. At a local scale, D. apicalis has been found sympatrically in south-east Queensland with D. picturatus (Nudgee) and with D. nigricans (Boondall Wetlands and North Stradbroke Island). Daerlac apicalis has also been collected sympatrically with D. picturatus in the north of Western Australia, with D. cephalotes in south-eastern Australia (Weston Creek, ACT, and Chiltern Forest, VIC), and with both D. cephalotes and D. nigricans near Warwick, QLD. Prior to this work D. apicalis was known only from north-east Queensland (Cassis and Gross 2002).
(Figs 2, 3, 4, 7B, E, F, 8, 10, 12B, 13B)
Rhyparochromus cephalotes Dallas, 1852: 577 (sp. nov.). Rhyparochromus nigripes Dallas, 1852: 578 (sp. nov.). - Walker, 1872: 109 (synonymy). Gydnes cephalotes Stål, 1866: 161 (comb. nov.). Pamera cephalotes Stål, 1874: 151 (comb. nov.). - Lethierry & Severin, 1894: 192 (catalogue); Distant, 1901: 479 (taxonomy). Pamera nigripes Stål, 1874: 151 (comb. nov.). - Lethierry & Severin, 1894: 192 (catalogue). Stigmatonotum cephalotes Scudder, 1962: 772 (comb. nov.). - Slater, 1964: 1181 (catalogue). Daerlac cephalotes Scudder, 1967: 250 (comb. nov.). - Slater & O'Donnell, 1995: 141 (catalogue); Cassis & Gross, 2002: 345 (catalogue). Daerlac tricolor Signoret, 1881: clviii (sp. nov.). - Lethierry & Severin, 1894: 196 (catalogue; separate species); Tillyard, 1926: 147 (ant mimicry); Slater, 1964: 1181 (catalogue); Slater & O'Donnell, 1995: 141 (catalogue; separate species); Cassis & Gross, 2002: 346 (catalogue; separate species); syn. nov. Daerlac affinis Distant, 1901: 483 (sp. nov.). - Slater, 1964: 1059 (catalogue; separate species); Scudder, 1967: 250 (synonymy with Daerlac tricolor). Vulturnia albonotata Kirkaldy, 1908: 776 (sp. nov.). - Bergroth, 1916: 255 (synonymy with Daerlac tricolor).
Lectotype. ♂, New Holland (BMNH). Paralectotypes: 2♀, New Holland (BMNH).
Other material examined. AUSTRALIA: Australian Capital Territory: Black Mountain, 35.26387°S 149.10051°E, 30 Jan 1965, D. F. Waterhouse, 2♀ (33816, 33818) (ANIC); Blundell's, 35.367°S 148.833°E, 02 May 1950, Paramonov, 1♀ (33821) (ANIC); Blundell's, 35.29°S 149.14°E, 30 Apr 1930, I. M. Mackerras, 3♀ (33822 - 33824) (ANIC); Canberra, 35.2833°S 149.2167°E, 605 m, 07 Jul 1941, G. F. Hill, 1♀ (33815) (ANIC); 31 Jan 1957, W. J. M. Vestjens, 1♀ (33825) (ANIC); 24 Jan 1958, W. J. M. Vestjens, 1♀ (33826) (ANIC); 17 Aug 1979, J. F. Donaldson, 3♀ (199283, 199289 - 199290) (QDPI); Deakin, 35.31416°S 149.10777°E, 586 m, 10 Jul 1965, R. Angus, 1♂ (33814) (ANIC); Gibraltar Creek, 35.478°S 148.947°E, 15 Jan 1978, Z. Liepa, 1♂ (33802) (ANIC); Lee's Spring, Brindabella Ra., 35.35°S 148.8°E, 1242 m, 04 Aug 1930, G. F. Hill, 1♀ (33819) (ANIC); Piccadilly Circus, 35.22°S 148.48°E, 1240 m, Mar 1984, J. Lawrence, T. Weir, M-L. Johnson, 1♂ (33811), 1♀ (33813), 1 nymph (33812) (ANIC); Weston Creek, Cotter Rd, 35.31666°S 149.01666°E, 610 m, 06 Apr 1969, E. B. Britton, 1♂ (33801), 2♀ (33797, 33798) (ANIC). New South Wales: Albury, 36.065°S 146.901°E, 03 Jul 1927, F. E. Wilson, 1♂ (18034) (AM); Blue Mountains, 33.4985°S 150.3346°E, 1933, K. K. Spence, 1♂ (18030) (AM); Bombala, 36.91222°S 149.23833°E, 706 m, May 1929, Rev. A. J. Barrett, 1♀ (18038) (AM); Bondi Beach, 33.891°S 151.2741°E, 18 Jan 1932, K. K. Spence, 1♂ (18031) (AM); Brindabella, 35.3675°S 148.6528°E, 18 Aug 1930, J. W. Evans, 3♂ (33803 - 33805) (ANIC); Cooma, 36.23527°S 149.12527°E, 799 m, Feb 1889, Helms, 1♂ (18035) (AM); Delegate, 37.0485°S 148.9346°E, 761 m, 16 Sep 1930, Rev. A. J. Barrett, 1♀ (18037) (AM); Ebor Falls near Ebor, 30.40001°S 152.35°E, 10 Nov 1972 - 11 Nov 1972, M. Malipatil, 1♀ (201317) (QM); Greenacres estate nr Queanbeyan, 35.35555°S 149.23166°E, 584 m, 06 Nov 1993, M. S. Upton, 1♂ (33806) (ANIC); Jindabyne, 36.415°S 148.618°E, Mar 1889, Helms, 1♂ (18032), 1♀ (18033) (AM); Katoomba, 33.7°S 150.3°E, 1900, K. K. Spence, 1♂ (18028) (AM); Moonbar, Monaro, 36.5°S 148.53333°E, 1067 m, Mar 1889, Helms, 1♀1♂ (18041) (AM); Mt Wilson, Blue Mtns, 33.50361°S 150.37694°E, 1023 m, 14 Mar 1970, D. K. McAlpine, 1♂ (18036) (AM); Pilliga Scrub, via Coonabarabran, 30.4985°S 149.5012°E, 18 Dec 1974, I. Naumann, 1♂ (201323) (QM); Tin Mine Huts, Kosciuszko, 36.7°S 148.25076°E, 1250 m, 03 Feb 1945, K. C. McKeown, 1♂ (18029) (AM); Tom Groggin, Kosciuszko National Park, 36.5485°S 148.1346°E, 564 m, 12 May 1980, M. S. Harvey, 1♀ (33820) (ANIC); Tumut, 35.304°S 148.223°E, 18 Mar 1929, unknown collector, 1♀ (18039) (AM); Whiskers, 7 km WNW of Hoskinstown, 35.24°S 149.23°E, 23 Feb 1994, M. S. Upton, 1♀ (33829) (ANIC). Queensland: Acacia Ridge, Brisbane, 27.58946°S 153.02671°E, 22 Feb 1966, E. C. Dahms, Eucalyptus sp. (Myrtaceae), 2♂ (201359, 201360) (QM); Bald Mountain Area, 28.23333°S 152.41666°E, 18 May 1969, J. F. Donaldson, 2♂ (199281, 199282) (QDPI); Bald Mountain area, via Emu Vale, 28.27172°S 152.20322°E, 1219 m, 18 Jan 1973, G. Monteith, 2♀ (201319, 201322) (QM); Warwick, 200 m W Wildash Sch., 28.26666°S 152.1°E, 30 Nov 1997, R. Eastwood, A. McArthur, 1♂ (38865) (SAMA). South Australia: 2 mi. NW of Crafers, 34.97638°S 138.67944°E, 430 m, 15 Jan 1952, H. M. Cane, 1 adult (abdomen missing) (33827) (ANIC); Mount Lofty, 34.974°S 138.709°E, 31 Aug 1981, W. M. Wheeler, 1♂ (355330) (AMNH); Scott Ck CP, Little Falls Ck, North side, 35.1°S 138.68333°E, 353 m, 23 Jul 2000, T. Hands, 1♂ (38868) (SAMA); no other data, 4♀ (355326 -355329) (AMNH). Tasmania: Bust Me Gall Pass, 42.62824°S 147.62087°E, 25 May 1985, G. A. Holloway & C. N. Smithers, 1♀ (18042) (AM); Hobart, 42.9167°S 147.3333°E, 13 Jul 1913, G. H. Hardy, 1♀ (201320) (QM). Victoria: Bundoora, 37.69916°S 145.05888°E, 105 m, 12 Nov 1975, P P Stahle, 1♀ (201321) (QM); Chiltern Forest, 36.12555°S 146.62333°E, 273 m, Feb 1967, R. S. McInnes, 1♀ (33796) (ANIC); Coranderrk Reserve, [nr] Healesville, 37.68416°S 145.52166°E, 127 m, 03 Mar 1980, M. S. Harvey, 1♂ (33807), 1♀ (33808), 1 nymph (33810) (ANIC); 05 Mar 1980, M. S. Harvey, 1♀ (33809) (ANIC); Gisborne, 37.48805°S 144.59194°E, 421 m, 06 Jun 1901, Lyell, 1♂1♀ (33828) (ANIC); Mt Drummer, via Cann River, 37.55569°S 149.34486°E, 243 m, 29 Jan 1967, G. Monteith, 1♂ (201318) (QM); Strathbogie Range, 36.92986°S 145.74375°E, 496 m, 25 Oct 1961, K. L. Taylor, 1♀ (33817) (ANIC); no other data, 1 adult (abdomen missing) (18040) (AM).
Daerlac cephalotes is distinguished by the following combination of characters: moderately large size; tricoloured dorsum; antennae generally uniformly red-brown to dark brown, or light brown basally graduating to dark brown apically, inner side of AI mostly with light brown stripe; corium tricoloured, with cream-coloured wedge at apex, posterior edge of corium along wing membrane sometimes thinly dark brown; wing membrane predominantly unicolourous, dark brown, sometimes pale cream basally; abdominal SIV with cream spot at anterior end of lateral margin, entire lateral margins of abdominal SVI and SVII cream to light brown, rarely absent; female abdominal SIV with cream band medially across posterior half, not extending to lateral margins; legs mostly orange, forefemur bicoloured and medially red-brown, hind femur sometimes with longitudinal dark brown marking; supracoxal lobes cream in part.
Colouration (Figs 2, 3). Head: black; labium medium brown, LII and LIII sometimes lighter orange-brown. Antennae: AI red-brown, inner side with light brown stripe (Fig. 2); AII-AIV dark brown; rarely either all dark brown or gradually darkening from light brown AI to red-brown AIV. Thorax: mostly black, or very dark brown; posterior ridge of pronotum paler than rest of pronotum, colour varying from dark brown, to red-brown, and sometimes with orange highlights on posterolateral corners; scutellum dark brown to black, tip of apex cream; supracoxal lobes cream in part. Legs: legs bicoloured, mostly orange, forefemur bicoloured and medially red-brown to dark red-brown, hind femur sometimes with longitudinal red-brown stripe on inner side, hind tibia sometimes red-brown, tarsi slightly darkened (legs sometimes entirely red-brown), coxae orange, concolourous with leg. Hemelytra: setiferous punctuations mostly red-brown, dark brown on dark brown areas of hemelytra; clavus red-brown to dark brown, mostly with orange-brown highlights medially; corium, basal two-thirds with inner margin adjacent to clavus and at base adjacent to embolium dark brown to red-brown and medially orange with extent of orange varying, apical third dark brown with apical cream wedge, rarely with inner margin of apex adjacent to membrane dark brown; embolium, basal half cream, apical half dark brown, then cream at apex, partly concolourous with adjacent part of corium; wing membrane predominantly unicolourous, dark brown, sometimes with pale cream patches basally adjacent to inner margin of corium with extent of cream varying. Abdomen: mostly dark brown; lateral margin of SV always with cream spot anteriorly; entire lateral margins of SVI and SVII cream to light brown (rarely not) sometimes with continuous light lateral band from SV to posterior of abdomen (Fig. 3).
Structure. Moderately large size, body length 6.99-7.44 mm. Thorax: anterior lobe of pronotum ∼2.20× longer than posterior lobe, posterior lobe ∼1.19× wider than anterior lobe and ∼1.03× wider than head. Male genitalia: pygophore (see Fig. 10A, B), with lobe-like lateral processes on genital opening square (Fig. 10B); bilobed dorsal flange at base of parameres with lobes widely separated, distal lobe larger than lobe of ventral flange, basal groove very narrow, barely discernible (Fig. 10C).
As for male except for the following characters.
Colouration. Female abdominal SIV with cream band medially across posterior half of sternite, not extending to lateral margin (Figs 3, 4B).
Structure. Larger than male, body length 7.36-8.17 mm. Thorax: Pronotum, anterior lobe ∼1.94× longer than posterior lobe, posterior lobe ∼1.20× wider than anterior lobe and ∼1.09× wider than head. Abdomen: SIV large, one-quarter abdomen length; SVII tapered, slightly longer than wide; ovipositor greater than one-third abdomen length (Fig. 4B). Female genitalia: spermatheca (see Fig. 11B).
Colouration. Head, pronotum and scutellar area fuscous. Antennae either entirely dark brown or light brown basally, darkening to medium brown apically. Wing pads red-brown to dark brown with lateral triangular light patch, red to orange colour. Abdominal segments II and III light orange to red with some brown markings on lateral margins, post segment IV brown to dark brown, segment III with prominent dark brown spot medially on dorsum, sternite IV with broad cream-coloured V-shape across venter. Thoracic pleura fuscous. Thoracic sterna light pale cream to red. Legs polychromatic, femora dark brown with red patches apically, tibiae and tarsi brown, tarsal segment I sometimes lighter orange. Coxae and trochanters pale cream.
Surface and vestiture. Head, pronotum, scutellum and wing pads very finely punctate. Remainder of body mostly smooth. Body surface with dull shine. Body covered with moderate distribution of very short pale, adpressed, simple setae, intermixed with moderately elongate, erect, dark, bristle-like setae.
Structure. Body length 5.70-6.68 mm (n = 2). Head and antennae as in adult. Pronotum subquadrate, weakly convex dorsally, concave groove subposteriorly (visible in lateral view), weakly dividing a narrow and elevated posterior ridge, lateral margins carinate. Wing pads reaching anterior half of abdominal segment III. Abdominal tergal sutures reduced.
See Table 3.
Daerlac cephalotes is distinguished from all other species in the genus by a light brown stripe on the medial margin of the first antennal segment (rarely absent). Daerlac cephalotes is similar to D. nigricans in size and abdominal colouration and D. picturatus in dorsal colouration. It is distinguished from D. nigricans by being slightly smaller in size, and by colouration characters, inclusive of the posterior ridge of the pronotum (being lighter orange to red-brown), legs (bicoloured, orange and red-brown), and hemelytra (tricoloured corium and mostly dark brown wing membrane). The apex of the corium in D. cephalotes and D. nigricans differ with D. nigricans having a very small dark brown spot at the tip of the apex of the corium, whilst in D. cephalotes the tip of the apex is cream (concolourous with the rest of apex of corium) or sometimes with a thin dark brown stripe along entire edge of the posterior margin of the corium adjacent to the membrane. The light brown abdominal colour patterns in D. cephalotes are confined to lateral margins of sternites VI and VII (and also SVIII in females), and although very similar to D. nigricans, differ in the latter by the light colouration extending more medially onto abdominal SVII in males and covering the entire female terminalia (Fig. 3). Daerlac cephalotes is distinguished from D. picturatus by the larger size, more robust pronotum, antennal colouration, bicoloured legs (particular the forefemur), abdominal colouration in both sexes, mostly unicolourous wing membrane without a yellow apical spot, and the hemelytra with less coverage of orange colouration on the clavus and corium, and the cream wedge at the apex of corium rather than subapically as in D. picturatus.
Daerlac cephalotes is the most morphologically variable species in the genus in size and colouration, both within and between populations (Figs 2, 3, 4B). Colour characters in D. cephalotes exhibit significant and continuous variation, and as such accommodates that observed in the holotype and specimens identified as D. tricolor. Colour variation within D. cephalotes includes: ratio of orange versus darker red-brown colouration on the corium and clavus; the colour of the posterior margin of the corium adjacent to the wing membrane sometimes with a thin dark brown line along the edge; pale colouration on lateral margins of abdominal SV to SVII in males and SVIII in females (varies greatly from being absent, aside from a pale cream spot on abdominal SV (always present), to pale lateral margins from abdominal SVI to terminalia of abdomen, to complete pale lateral margins from abdominal SV to terminalia of abdomen; Fig. 3); and legs being either bicoloured (orange with red-brown markings) or uniformly darker red-brown. In addition, the wing membrane vein in D. cephalotes varies from being entirely dark brown to having light brown to yellow patches adjacent to the corium, which when present are confined to two discrete patches adjacent to the corium (compare with broadly light brown across the base of the membrane in D. nigricans). Dark colour morphs of D. cephalotes have been found at two localities in the ACT, which lack the orange colouration on the legs and hemelytra, but have the entire clavus and corium (except for the cream wedge), legs and antennae uniformly dark red-brown. All specimens from South Australia are smaller overall, with pale legs and antennae, but these characters are also present in individuals of east coast populations.
Daerlac cephalotes is distributed along the south-east coast of Australia from Brisbane to Tasmania (Fig. 13B). The distribution extends to the western slopes of the Great Dividing Range, with the western-most populations in South Australia (Adelaide district) disjunct from the east coast populations. The majority of collection records are from the south-east of New South Wales and the eastern half of Victoria. Daerlac cephalotes has also been recorded from north Queensland (Magnetic Island) and Western Australia (Lake Johnstone), and as D. tricolor from northern South Australia (Everard Ranges) and western NSW (Broken Hill region) (Cassis and Gross 2002). However, these latter localities have not been reconfirmed in this study. Daerlac cephalotes is sympatric with D. apicalis at Chiltern Forest, VIC and Weston Creek, ACT, with D. nigricans at Ebor Falls, NSW, and with both D. apicalis and D. nigricans near Warwick, QLD.
(Figs 2, 3, 4B, 5, 6, 7C, G, H, 11, 12A, 13C)
Daerlac nigricans Distant, 1918: 492 (sp. nov.). - Slater, 1964: 1058 (catalogue); Scudder, 1967: 274 (lectotype designation); Slater & O'Donnell, 1995: 141. - Cassis & Gross, 2002: 345 (catalogue).
Lectotype. ♂, Sydney (BMNH).
Other material examined. AUSTRALIA: Australian Capital Territory: Mt. Ainslie, Canberra, 35.267°S 149.167°E, Jun 1991, C. Reid, 1♂ (33831), 1♀ (33830) (ANIC). New South Wales: Armidale, 30.5152°S 151.6651°E, 979 m, 30 Aug 1986, D. S. Horning, Jr, 1♀ (33851) (ANIC); Broulee, 35.85666°S 150.17361°E, 5 m, 29 Jan 2007, S. O. Shattuck, 1♂ (33832) (ANIC); Ebor Falls near Ebor, 30.40001°S 152.35°E, 10-11 Nov 1972, M. Malipatil, 1♂ (201351), 2♀ (201353, 201354) (QM); Hanging Rock, 31.48333°S 151.2°E, 1051 m, 08 Jun 1992, D. S. Horning, Jr, 1♀ (33833) (ANIC); St Albans, 33.29194°S 150.97055°E, 24 m, 05 Jul 1980, D. A. Doolan, 1♀ (33834) (ANIC); Whiskers, 7 km WNW of Hoskinstown, 35.24°S 149.23°E, 12 Nov 1993, M. S. Upton, 1♀ (33835) (ANIC); Wingello State Forest, Gap Rd, 6km from Burnt Pine Rd, 34.7275°S 150.24027°E, 18 Sep 2008, C. Symonds and F. Powell, ex. Kunzea ambigua (Myrtaceae), 1♀ (18056) (UNSW). Queensland: Acacia Ridge, Brisbane, 27.58946°S 153.02671°E, 11 Mar 1965, E. C. Dahms, 1♂ (201344) (QM); 28 Mar 1965, E. C. Dahms, 1♀ (201352) (QM); 01 Apr 1965, E. C. Dahms, 3♂ (201341 -201343) (QM); 08 Jul 1965, E. C. Dahms, 1♀ (201349) (QM); 18 May 1966, E. C. Dahms, 2♂ (201347, 201348) (QM); Mar 1968, E. C. Dahms, 3♀ (201336 - 201338) (QM); 16 Jan 1973, E. C. Dahms, 1♂ (201345), 1♀ (201346) (QM); 23 Apr 1985, L. Haren, 1♂ (201339), 1♀ (201340) (QM); Dunwich, Stradbroke Island, 27.5°S 153.4°E, 02 May 1972, G. B. Monteith, 1♂ (201350) (QM); Fletcher, nr Glen Alpin, 28.77416°S 151.86527°E, 801 m, 24 Jan 1967, E. C. Dahms, 1♂ (192983), 1♀ (192984) (UQIC); One Tree Hill, Brisbane, 27.48527°S 152.95944°E, 213 m, 12 Dec 1925, A. Musgrave, 2♀ (18044, 18045) (AM); Belmont Hills Bushlands, site 1, 27.51305°S 153.11805°E, 80 m, 16 Apr 2003, C. J. Burwell, 1♀ (201329) (QM); 03 Nov 2003, QM party, 1♀ (201328) (QM); 19 Feb 2004, QM party, 1♂ (201327) (QM); Boondall Wetlands, site 1, 27.33683°S 153.0712°E, 22 Apr 2003, C. J. Burwell, 1♂ (201331), 1♀ (201330) (QM); 02 Sep 2003, G. B. Monteith, 1♂ (201335) (QM); Brisbane, 27.46785°S 153.02801°E, 19 Mar 1954, A. G. Barrie, 1♀ (201334) (QM); Brisbane, Long Pocket, 27.51°S 153°E, 06 Nov 1979, J. F. Donaldson, 2♂ (199284, 199285) (QDPI); Gold Creek Reservoir, site 1, 27.45883°S 152.872°E, 04 Nov 2003, QM party, 2♀ (201325, 201326) (QM); 23 Feb 2004, QM party, 1♂ (201324) (QM); Illaweena Street, Drewvale, 27.63983°S 153.0578°E, 18 Sep 2003, QM party, 1♂ (201332), 1♀ (201333) (QM); Kingaroy, 26.54083°S 151.83944°E, 432 m, 01 Oct 1963, unknown collector, 1♀ (201355) (QM); Kuranda, 16.81888°S 145.63638°E, 355 m, Jan 1950, J. G. Brooks, 1♀ (18043) (AM); Stanthorpe, 28.662°S 151.935°E, 18 Feb 1928, E. Sutton, 1 adult (abdomen missing) (201356) (QM); Jul 1931, H. James, 1♂ (199286), 2♀ (199287, 199288) (QDPI); Warwick, 200 m W Wildash Sch., 28.26666°S 152.1°E, 30 Nov 1997, R. Eastwood, A. McArthur, 1 adult (abdomen missing) (38869) (SAMA).
Daerlac nigricans is distinguished by the following combination of characters: moderately large body size; mostly dark brown to black colouration; antennae uniformly dark brown; most of dorsum and venter dark brown to black; corium bicoloured with apex cream, extreme apex of corium with small dark brown spot; wing membrane bicoloured, basally creamy yellow, apically brown; abdomen with cream patch anteriorly on lateral margin of abdominal SV, entire lateral margins from abdominal SVI to terminalia of abdomen cream to light brown, posterior of abdomen more broadly light brown and more so in females; female abdominal SIV with cream band medially across posterior half, not extending to lateral margins; legs uniformly dark brown to black, except for light brown to cream trochanter of forefemora; and supracoxal lobes cream in part.
Colouration (Figs 2, 3). Head: black; labium medium to dark brown. Antennae: dark brown, sometimes slightly lighter medium brown. Thorax: mostly black, or very dark brown; posterior ridge of pronotum concolourous with rest of pronotum, sometimes slightly lighter dark red-brown; scutellum dark brown to black, tip of apex cream; supracoxal lobes cream in part, not concolourous with rest of thorax. Legs: uniformly red-brown to dark brown; trochanter of forefemur cream (Fig. 3), rest of trochanter and all coxae concolourous with rest of leg. Hemelytra: setiferous punctuations dark brown on dark brown areas of hemelytra, red-brown on cream area of corium; clavus dark brown; corium, basal three quarters dark brown, apical quarter cream, tip of apex of corium dark brown; embolium, basal half cream, apical half concolourous with adjacent part of corium, dark brown then cream at apex; wing membrane bicoloured, basal half creamy yellow, diffusing to brown at apex, extent of lighter basal colouration varying. Abdomen: mostly dark brown, almost black; lateral margin of SV always with cream spot anteriorly; entire lateral margins of SVI and SVII always light brown, light colouration extending medially on abdominal sternites (Fig. 3).
Structure. Moderately large size, body length 6.24-7.13 mm. Thorax: anterior lobe of pronotum ∼2.02× longer than posterior lobe, posterior lobe ∼1.16× wider than anterior lobe and ∼0.78× wider than head. Male genitalia: pygophore (see Fig. 10A, B) with quadrate lobe-like lateral processes on genital opening (Fig. 10B); bilobed dorsal flange at base of parameres with lobes widely separated, distal lobe larger than lobe of ventral flange, basal groove very narrow, barely discernible (Fig. 10C); aedeagus (Fig. 10D, E).
As for male except for the following characters.
Colouration. Female abdominal SIV with cream band medially across posterior half, not extending to lateral margin (Figs 3, 4B); apex of abdomen light brown (Fig. 3).
Structure. Larger than male, body length 7.93-8.16 mm. Thorax: anterior lobe of pronotum ∼2.11× longer than posterior lobe, posterior lobe ∼1.18× wider than anterior lobe and ∼1.03× wider than head. Abdomen: SIV large, one-quarter abdomen length; SVII tapered, slightly longer than wide; ovipositor greater than one-third abdominal length. Female genitalia: spermatheca (see Fig. 12A).
See Table 3.
In D. nigricans the female abdomen is more extensively light brown at apex than in the male. However, in both males and females the light colouration is more extensively spread across the posterior segments of the abdomen compared to D. cephalotes, where it is restricted to the lateral margins (when present). Oligomeric antennae (antennal fusion involving the loss of one antennal segment and the elongation of the remaining three) was observed in one male D. nigricans specimen in this study (Fig. 7C). Also see remarks for D. apicalis and D. cephalotes for differential diagnoses.
Daerlac nigricans extends from south-east Queensland to the ACT, with the exception of one disjunct population recorded from far north-east Queensland (Fig. 13C). Daerlac nigricans appears to be restricted to on or east of the Great Dividing Range. It was previously known only from south-east New South Wales (Cassis and Gross 2002). Daerlac nigricans has been collected sympatrically with D. apicalis (Boondall Wetlands and Stradbroke Island, QLD) and D. cephalotes (Ebor Falls, NSW), and also with both of these species near Warwick, QLD.
(Figs 2, 3, 4B, 13D)
Pamera picturatus Distant, 1904: 267 (sp. nov.). Daerlac picturata Scudder, 1962: 766 (comb. nov.). - Slater, 1964: 1059 (catalogue); Scudder, 1967: 278 (lectotype designation). Daerlac picturatus Steyskal, 1973: 276 (spelling emendation). - Slater & O'Donnell, 1995: 141 (catalogue); Cassis & Gross, 2002: 345 (catalogue).
Lectotype. ♀, Townsville (BMNH).
Other material examined. AUSTRALIA: Queensland: Nudgee, 27.37062°S 153.08995°E, 5 m, Jan 1924, H. Hacker, 1♀(201358) (QM). South Australia: Flinders Ranges, Mawson Plateau, 30.08666°S 139.44305°E, 630 m, 06 Sep 2000, T. Hands, 1♀ (38870) (SAMA). Western Australia: Armadale, 32.13527°S 116.02972°E, 98 m, 12 May 1934, K. R. Norris, 2♀ (33837, 33838) (ANIC); Broomehill, 33.85°S 117.55°E, 12 Feb 1985, R. P. McMillan, 1♀ (30361) (WAM); Eneabba, 29.81666°S 115.26666°E, 31 May 1990, R. P. McMillan, 2♀ (30359, 30360) (WAM); Lorna Glen Station, 26.20972°S 121.4075°E, 05 Apr 2002 -14 Apr 2002, M. A. Cowan, 1♀ (30362) (WAM); Manjimup, 34.23528°S 116.1411°E, 280 m, unknown date and collector, 1♂ (30357) (WAM); Palm Springs, 15 km W of Millstream, 21.55°S 116.96666°E, 16 Jun 1984, R. P. McMillan, 1♀ (30358) (WAM); W. A. Museum, Perth, 31.94944°S 115.86194°E, 19 m, 01 Dec 1988, B. A. Jones, 1♀ (30363) (WAM).
Daerlac picturatus is distinguished by the following combination of characters: moderate size; tricoloured dorsum; antennae light brown basally, dark brown apically; clavus and corium mostly orange-brown; corium tricoloured, corium with cream-coloured wedge before apex, apex of corium dark brown, corium with small dark brown medial patch between orange and cream sections; wing membrane bicoloured, mostly dark brown, apex yellow; abdominal SIV with cream band along entire lateral margin, female abdominal SVII and SVIII sometimes light brown dorsally, female abdominal SIV with cream band across majority of sternite, extending to lateral margins, abdominal SIII lighter orange-brown medially; legs mostly orange, tibiae and tarsi usually darker red-brown at least in part, legs sometimes entirely red-brown; supracoxal lobes orange in part.
Colouration (Figs 2, 3, 4B). Head: black; labium medium brown. Antennae: mostly AI-AII light brown, AIII-AIV medium brown, sometimes mostly light brown with apical portions of AIII and AIV medium to dark brown. Thorax: mostly black, or very dark brown; posterior ridge of pronotum paler than rest of pronotum, orange-brown; scutellum dark brown to black, apex lighter orange-brown with small cream tip; supracoxal lobes orange to red-brown in part, rest concolourous with rest of thorax. Legs: uniformly orange to red-brown, trochanter and tarsi sometimes slightly lighter brown; coxae orange-brown, concolourous with leg. Hemelytra: setiferous punctuations red-brown; clavus orange; corium, basal two-thirds orange, apical third with subapical cream wedge, apex dark brown; embolium, basal half cream, apical half dark brown, then cream, dark brown, partly concolourous with adjacent part of corium; corium sometimes with dark brown spot medially at junction of orange and cream sections; wing membrane predominantly bicoloured, mostly dark brown, apex yellow. Abdomen: mostly red-brown to dark brown; female abdominal SIV with cream band across majority of sternite, extending to lateral margin (Figs 3, 4B), abdominal SIII lighter orange-brown medially, adjacent to abdominal SIV (Fig. 4B); lateral margins of abdominal SV and SVI concolourous with rest of abdomen (Fig. 3); lateral margins of abdominal SVII sometimes light brown, extending to apex of abdomen (Fig. 3).
Structure. Moderate to small size, body length 6.14-6.74 mm. Thorax: anterior lobe of pronotum ∼1.54× longer than posterior lobe, posterior lobe ∼1.28× wider than anterior lobe and ∼1.15× wider than head. Abdomen: SIV greatly enlarged, one-third abdomen length; SVII rounded, as wide as long; ovipositor less than one-third abdominal length.
See Table 3.
The above description is based primarily on female specimens. At the completion of this work a single male was obtained from the Western Australian Museum, and was confirmed as D. picturatus. It was found to be similar in most respects to the female. Daerlac picturatus is most similar to its sister species D. apicalis in size, structure and colour patterning, but also shares the orange colour patterning character on the hemelytra with D. cephalotes. Similar characters to D. apicalis include: small size, pronotum more slender and elongate anteriorly, wider posterior margin of pronotum, cream-coloured abdominal markings of female, and dark brown apex of the corium and pale apical spot on the wing membrane. However, D. picturatus may be distinguished from D. apicalis by the following characters: female abdomen slightly more swollen; head and pronotum more slender; orange colouration on clavus and corium; and, bicoloured antennae and legs. Also, see remarks for D. cephalotes for differential diagnosis. Daerlac picturatus has also been found to have uniformly dark brown legs in some populations (Manjimup, WA), and variation in the extent of orange-brown colouration on the posterior pronotal lobe, with either the entire lobe orange-brown or just the posterior half of the lobe.
Daerlac picturatus is widely distributed across Australia, from south-west, central and northern Western Australia to central-east South Australia to eastern Queensland (including north and south coasts) (Fig. 13D). Daerlac picturatus has been collected sympatrically with D. apicalis at Nudgee in Queensland and Palm Springs in Western Australia, and both species are recorded from Townsville, QLD. Daerlac picturatus was formerly known only from the type locality (Townsville) (Cassis and Gross 2002).
Species of Daerlac and specimens examined in this study have been collected in association with eucalypt species (mostly in forests and woodlands) and, on one occasion, from Melaleuca-dominated woodland in south-east Queensland. They have been found under the bark or on the trunks of eucalypts, and in leaf litter at the base of eucalypts. It is unknown whether species of Daerlac favour particular eucalypt species. To date they have been recorded in association with Eucalyptus camaldulensis Dehnh., E. sideroxylon Woolls and E. tereticornis Sm. There are no data to indicate whether they are just sheltering on eucalypts, or whether they are foraging on pre-dispersed or post-dispersed seeds of putative host plants. Although most rhyparochromids are known to be epigaeic seed feeders (Sweet 1964a; Sweet 1964b; Anderson 1985; Schuh and Slater 1995; Cassis and Gross 2002), the udeocorine genus Laryngodus has been found to be largely arboreal, feeding on pre-dispersed seeds in the cones of sheoaks (Allocasuarina spp.) (Slater et al. 2009).
Daerlac is found across continental Australia, including eastern Tasmania. However, it is concentrated in temperate eastern Australia, with a few specimen records from south-west and central to northern Western Australia, semi-arid Queensland and New South Wales, and tropical northern Australia. Limits of distribution for Daerlac are attributable to distribution of eucalypt forests and woodlands and, to a lesser extent, shrublands that dominate Australia's major ecoregions beyond the arid zone (Fig. 13). Thus, Daerlac is a predominantly Bassian taxon with some Torresian elements (CSIRO 1970). In eastern Australia, Daerlac spp. are found on the edge of the Eyrean faunal province, but limited to semi-arid regions where rainfall is above 250-300 mm per annum (see Fig. 13).
There is a surprisingly high degree of overlap in distribution of Daerlac species and all species are broadly distributed. Daerlac picturatus, while being the least collected species, is perhaps the most widely distributed, extending from south-western Western Australia to north-eastern Queensland (Fig. 13D). However, it is absent from the south-east of Australia (New South Wales and Victoria), where the other three species are found. Daerlac apicalis is also widely distributed, extending from northern Australia south to Victoria and west to Western Australia (Fig. 13A). Daerlac nigricans is the most easterly restricted species, and does not extend west of the Great Dividing Range (Fig. 13C). Daerlac cephalotes is mostly restricted to south-eastern Australia and is the only species found in the Adelaide district and in Tasmania, and the only species not found in northern Queensland (Fig. 13B).
The sister species D. apicalis and D. picturatus are the only two species recorded from Western Australia, are the most widely distributed from the east to the west of the continent, and extend into the semi-arid zone. Both have been found in northern and south-eastern Queensland and in the Pilbara region of Western Australia. Although overlapping in distribution, including sympatry in south-east Queensland and northern Western Australia, the distributions of D. apicalis and D. picturatus are divergent, with D. apicalis having a northern and eastern distribution and D. picturatus having a more south-central and south-western distribution. Daerlac apicalis is absent from the south-west of Western Australia where most specimens of D. picturatus have been found.
The sister species Daerlac cephalotes and D. nigricans are both largely restricted to eastern Australia and are largely overlapping from south-east Queensland to southern New South Wales. However, D. nigricans extends further north to far north Queensland and D. cephalotes extends further south into Victoria and Tasmania. In addition, D. cephalotes extends further west, through the western slopes of the Great Dividing Range in NSW and Victoria as far as south-eastern South Australia. In the southern reaches of its distribution, D. cephalotes overlaps with D. apicalis, with the latter species overlapping with D. nigricans beyond the northern limits of D. cephalotes. Daerlac apicalis is largely absent from eastern New South Wales where D. cephalotes and D. nigricans overlap. It is common for two species of Daerlac to occur across the generic distributional range, and on occasion D. apicalis, D. cephalotes and D. nigricans overlap, inclusive of localities in the Australian Capital Territory, Brisbane (Acacia Ridge) and near the Queensland and New South Wales state boundary (near Warwick), although collected on different dates.
Daerlac is widely distributed across continental Australia, although most specimens have been collected in temperate Australia. Despite extensive recent surveys of Heteroptera in temperate Australia (Cassis et al. 2007), there have been few new collections of Daerlac. This may be due to a lack of sampling effort. However, this may also have conservation significance, as the extent of land clearing in open woodland habitats in Australia is considered a serious threat to biodiversity, where their exploitation has been hypothesised as a reason for bird decline (Ford et al. 2001).
McIver and Stonedahl (1993) have proposed that heteropteran ant-mimics are, generally, Batesian mimics. Species of Daerlac have been recorded on the ground with ants and at the base and on trunks of eucalypts with ants, but there are no ant association records for Daerlac that would allow for hypotheses of putative ant model(s). It is noteworthy that Daerlac species differ in colour hue and patterns. For example, Daerlac nigricans is dark brown to black with a striking golden abdomen in the nymphs (Chew 2011) and cream to light brown of wings and lateral margins of abdomen in the adults, which bears resemblance to the golden-tailed species found in the ant genera Camponotus Mayr and Polyrhachis (Hagiomyrma) Smith. Species of these ant genera are known to nest on the ground and forage upon trunks of eucalypts (Shattuck 1999; CSIRO 2011). In contrast, D. cephalotes and D. picturatus have red and orange colouration, which show resemblance to some Camponotus spp., the aggressive predatory muscleman tree ant (Podomyrma gratiosa (Smith, 1858)), as well as larger predatory or seed harvesting ants found in eucalypt forests, such as Myrmecia Fabricius spp. and Rhytidoponera spp. (Bashford 1993). On the basis of colouration and structural characters we regard Daerlac species as ant-mimetic and worthy of investigation as a model system in mimicry studies.
There has been no modern treatment of generic concepts within the tribe Udeocorini. Gross (1962) published on several xerophilic species found in temperate Australia (including Cryptocoris Gross, Porander Gross, Telocoris Gross and Zygocoris Gross). Most recently, Slater et al. (2009) published a revision of the enigmatic genus Laryngodus Herrich-Schaefer, without making comments on generic limits within the tribe. These authors indicated putative problems with the tribal classification of the family Rhyparochromidae, pointedly criticising the use of characters of the abdominal sternal sutures. They noted the SIV/SV suture was complete (reaching lateral margins) and straight in Laryngodus. They also reported that the suture was complete in the South American udeocorine genus Astemmoplitus Spinola, as well as representatives of seven other rhyparochromid tribes. We have re-examined this character in Laryngodus and found the SIV/SV suture is just incomplete (not reaching dorsal sutures) (Fig. 14). In comparison, the suture in Daerlac is also just incomplete but curves anteriorly (Figs 6B, D, 8B). In other Australian Udeocorini genera, such as Euander, Fontejus and Udeocoris, this suture is more strongly curved anteriorly. Our observations in the Rhyparochrominae suggest that in the Udeocorini, even when the abdominal SIV/SV sutures curve anteriorly, they do so less than in other tribes, such as the Drymini and Lethaeini, where they terminate proximal to the abdominal SIII/SIV suture. Henry's (1997) theory of rhyparochromine monophyly is based on the incomplete abdominal SIV/SV suture. Slater et al. (2009) and Cassis and Schuh (2010) brought this into question, but on the basis of our examination of udeocorine representatives, a more detailed comparative study and phylogenetic analysis is required, with emphasis on broader taxon sampling.
The phylogenetic position of Daerlac is uncertain, pending a comprehensive analysis of all the tribal members. However, on the basis of comparative morphology, Daerlac exhibits similarities with Laryngodus in possessing a bilobed pronotum, with an enlarged anterior lobe and a short campanulate posterior lobe. The anterior lobe in Daerlac is more globose and less elongate in comparison to Laryngodus. This type of pronotal development is most exaggerated in these two genera in the Udeocorini, with most Australian representatives having a broader and flatter pronotum. In Porander Gross and Fontejus, the anterior lobe is also more elevated, but the posterior lobe is longer, compared with that in Daerlac and Laryngodus. Daerlac and Laryngodus are both moderately large and have an elongate, mostly parallel-sided body. Daerlac and Laryngodus both have a distinctive mixed vestiture, comprising short, adpressed simple setae, interspersed with elongate, erect simple setae, on the body and legs. However, the adpressed setae in Laryngodus are slightly shorter and more densely distributed than in Daerlac.
Fontejus is the other genus in the Udeocorini that possesses this mixed vestiture. However, in this genus, the short adpressed setae on the head and thorax are more sparsely distributed. Also, in Fontejus the perpendicular, densely distributed elongate setae are absent on the meta- and mesofemora and tibiae, whose presence is distinctive in Daerlac and Laryngodus.
Daerlac and Laryngodus possess slightly shortened wings, being submacropterous, with the membrane well developed. Slater (1975) identified and classified wing polymorphism in the Australian rhyparochromids, recognising in the Udeocorini several polymorphic species in seven genera, mostly from the south-west Western Australia, although brachyptery has not been observed in Laryngodus or Daerlac. Malipatil (1979) noted the low incidence of wing polymorphism in Australian rhyparochromines in south-east Queensland, and suggested the wide scope for dispersal and migration in parts of mainland Australia. The wide distributions of all Daerlac species in eastern Australia may be explained by their fully functional wings.
To date, there has been no description of male genitalia in the Udeocorini, aside from Laryngodus cervantes Slater, Schuh, Cassis, Johnson & Pedraza-Peñalosa. In the present work, the male genitalia are illustrated and described in the generic and species descriptions. In Laryngodus the gonoporal process has seven to eight coils and a well-developed helicoid process. The gonoporal process in Daerlac has five to six coils and a well-developed helicoid process, which is also the case in Euander (G. Cassis and C. Symonds, unpubl. data). All of the above taxa have a secondary gonopore with an expanded membranous flute. The parameres in the above three genera are similar, being hook-like, with a broad sensory lobe, sometimes with a short flange-like process and an arcuate apophysis. Laryngodus exhibits a unique lateral process of the genital opening of the pygophore, where they are bifid and more complex than the short triangular processes found in Daerlac and Euander.
The female spermatheca of Daerlac spp. resembles the structure of the spermatheca of Australian Myodochini genera, described in detail by Malipatil (1978). However, in Daerlac spp. the coiled part appears more irregularly arranged, more compact and possibly shorter, the basal part (before the wider ribbed part) generally shorter and not as narrow in relation to the coiled part, and the wider ribbed medial portion shorter in general.
We would like to thank Hannah Finlay for the preparation of dissections and illustrations of the male and female genitalia, and some of the photographs. We would also like to thank the following people: Alex Brown of UNSW, for help completing some of the measurements; Shawn Laffan, School of Biological Earth and Environmental Sciences, UNSW, for providing the BIOCLIM data and assistance using ArcMap GIS; Glenn Johnstone, Environmental Resources Inventory Network, Dept. of Sustainability, Environment, Water, Population & Communities, for providing the Terrestrial Ecoregions of Australia mapping data and for assistance using these files; and the Willi Hennig Society for the use of TNT software. We thank the following institutions and their research and collection management personnel for the loan of specimens: Dave Britton and Derek Smith, Australian Museum; Tom Weir, Australian National Insect Collection; Jan Forrest, South Australian Museum; Geoff Thompson and Geoff Monteith, Queensland Museum; Shaun Winterton, while at the then Queensland Department of Primary Industries; Terry Houston and Brian Hanich, Western Australian Museum; Tom Henry, United States National Museum; and Toby Schuh and Christine Johnson, American Museum of Natural History. The Australian Biological Resources Study is thanked for funding this research. We also thank the corresponding editor Lyn Cook and two reviewers whose comments have enhanced this work.
By Gerasimos Cassis, Evolution and Ecology Research Centre, University of New South Wales, Sydney, NSW 2052, Australia, gcassis@unsw.edu.au and Celia Symonds, Evolution and Ecology Research Centre, University of New South Wales, Sydney, NSW 2052, Australia.
