A review of the genus Larrisson Menke , 1967 , and description of the new genus Larrissa ( Hymenoptera , Crabronidae )

Larrisson menkei Pagliano, 1995, is transferred to Clitemnestra comb. n. and twelve new species of Larrisson are described: armatus, carinatus, latifrons, niger, orbitalis, punctatus, quintus, spinosus, sulcatus, tegularis, tibialis, and variegatus, all from Australia. Based on a cladistic analysis of the genus, Larrisson nedymus Menke is transferred to Larrissa gen. n. Additional locality records are provided for Larrisson abnormis Turner, azyx Menke, rieki Menke, and for Larrissa nedyma (Menke). An updated key to the species of Larrisson and Larrissa is provided.


Introduction
Larrisson is a little known, rarely collected, strictly Australian genus of the solitary wasp family Crabronidae.It was established for Sericophorus abnormis Turner, 1914 by Menke (1967) who in 1979 added three new species, revised the genus, and provided a key to their identification.Pagliano (1995) described another species, Larrisson menkei, from a single specimen, that he later generously donated to the California Academy of Sciences.A study of the holotype revealed that it is actually a member of Clitemnestra, a conclusion confirmed by Michael Ohl (Museum für Naturkunde der Humboldt-Universität, Berlin, Germany), who has also examined the specimen.The justification for the new combination is provided below.
Little is known about the habits of Larrisson.I collected them in open habitats, exposed to the sun, on light, mainly sandy soils (as in Fig. 15).Certainly they nest in the ground, and the only prey record is a mirid.

Materials and methods
During my recent expeditions to Australia (Northern Territory, 3 March -27 April 2008;Western Australia, 14 October -13 December 2008; New South Wales, 1 December 2009 -15 January 2010; South Australia, 1 December 2010 -4 February 2011), I collected six new species of Larrisson, and visits to the Western Australian Museum, Perth, Western Australia (11-12 December 2008), Australian National Insect Collection, Canberra, Australian Capital Territory (20-24 April 2009), and South Australian Museum, Adelaide, South Australia (1-3 February 2011) revealed additional new species.Descriptions of the undescribed species are given below, as are additional locality records for four previously known species.I have examined specimens of all species of Larrisson and provide an updated key to species.I have also performed a cladistic analysis of the species of Larrisson, which demonstrates that L. nedymus should be placed in a separate genus that I describe below under the name of Larrissa.As a result, Larrisson now totals 15 species, up from previously known three.
Abbreviations in the text include: AMS Australian Museum, Sydney, New South Wales, Australia.

ANIC
Australian National Insect Collection (CSIRO), Canberra, Australian Capital Territory, Australia.BMNH The Natural History Museum (formerly British Museum Natural History), London, United Kingdom.

CAS
California Academy of Sciences, San Francisco, California, USA.
Recognition.In his key to world genera of Miscophini, Menke (1977) differentiated Larrisson from Sericophorus by the length of the occipital carina, which ends before reaching the hypostomal carina in the former and reaches the hypostomal carina in the latter.This character, however, no longer holds, as the occipital carina effaces before reaching the hypostomal carina in three Sericophorus described in Lomholdt and Pulawski (2010): S. centralis Pulawski, S. genalis Pulawski, and S. politus Lomholdt.
A useful, although not universal recognition character, is the shape of the posterior propodeal surface: it has no median carina in Larrisson, whereas in the vast majority of Sericophorus there is a well-defined carina below the median sulcus (also present in Lyroda and Sphodrotes).The carina, however, is only one quarter of the posterior surface long in the holotype of Sericophorus centralis, and absent in the paratype.The essential difference between the two genera is in two male characters: Larrisson have 11 flagellomeres and a well defined volsella, whereas in Sericophorus the antenna has 10 flagellomeres and the volsella is absent.Menke (1979) recognized two species groups in Larrisson: the monotypic nedymus group and the abnormis group that included abnormis Turner, azyx Menke, and rieki Menke (the latter two known from the male sex only).The twelve new species (armatus, carinatus, latifrons, niger, orbitalis, punctatus, quintus, spinosus, sulcatus, tegularis, tibialis, and variegatus) generally agree well with his diagnosis of the abnormis group except the inner mandibular margin has a tooth near midlength in tibialis (and also in an additional specimen of rieki), the metanotum has no spine or tubercle in armatus, carinatus, niger, punctatus, and tegularis (as in nedymus), mesothoracic venter is conspicuously depressed in punctatus (as in nedymus), the propodeal side is unridged mesally in armatus and most quintus (as in nedymus), male sternum VIII is emarginate apically in armatus, orbitalis, quintus, tegularis, and variegatus (as in nedymus), and gonocoxite has no accessory lobes and no elaborate setal fringe in tegularis (as in nedymus).The twelve species fit well the abnormis group in all other characters: scapal basin present, impunctate or sparsely to densely punctate (frons uniformly punctate in nedymus), scape longer than flagellomeres I-III combined (shorter than that in nedymus), inner mandibular margin with preapical tooth, with small incision at about midlength in female (without preapical tooth, with two teeth near midlength in nedymus), mesopleuron rounded anteriorly (abruptly angular below pronotal lobe in nedymus), forewing vein M diverging from M+Cu distad of cu-a or interstitial with cu-a (diverging basad of cu-a in nedymus), glabrous area of propodeal dorsum covering entire enclosure except limited to slightly more than median sulcus in tegularis and variegatus (limited to median sulcus in nedymus), male forecoxa and foretrochanter simple (forecoxa with apical spine and foretrochanter emarginate basally in nedymus), basolateral carina of tergum I not expanded into lamella (lamella present in nedymus), apical tergum of female without a narrow, impunctate and asetose marginal lamella (lamella present in nedymus), volsella ending near apex of penis valve or exceeding it (ending at half length of penis valve in nedymus); and head of penis valve not dentate (dentate in nedymus).Menke (1979) also claimed that the stipes and prementum are shortened in the abnormis group, the prementum being less than twice as long as wide, but not shortened in nedymus, with the prementum little more than twice as long as wide.I cannot confirm this difference: the length of these mouthparts is practically identical in azyx, nedymus, and rieki, although definitely smaller in abnormis.
Prey of Larrisson was unknown until now, but the holotype female of niger is pinned with her prey, an adult mirid 3.6 mm long.The specimen has an additional label that reads "Wasp grasped bug on terete-leaved Acacia", indicating that the mirid was not placed on the same pin accidentally.Randall T. Schuh (American Museum of Natural History, New York, New York) kindly identified it as a male of the tribe Orthotylini belonging to an undescribed genus and species.
Phylogenetic analysis.Lomholdt (1985) placed Larrisson, together with Sericophorus, in the tribe Sericophorini Dalla Torre, that he characterized by a second submarginal cell conspicuously narrowed anteriorly, a synapomorphy.In the following analysis, I have used as outgroups representatives of three other miscophine genera with posteroventrally emarginate mandible that occur in Australia (Lyroda, Sericophorus, and Sphodrotes) and that appear to be more closely related to Sericophorus than the non-Australian genera.Sericophorus was used below as a sister taxon of Larrisson, and S. relucens Rayment, a relatively unspecialized species of the genus, was selected as the first outgroup.Sphodrotes punctuosa Kohl was selected as the second outgroup, and Lyroda venusta Bingham as the root taxon.Since females are known for only nine species of Larrisson (out of the total of 15), whereas the males are known for 13 species, the analysis was based entirely on male characters, and L. latifrons, niger, and punctatus (known from the female only) were excluded.Autapomorphies were included.The following is the list of characters.
The following data matrix (Table 1) was constructed using Winclada version 1.00.08 (Nixon 2002).Multiple character states were treated as additive.
The above data matrix was analyzed using the Willi Hennig Society edition of TNT (Goloboff et al. 2008) with 1,000 replications and 1,000 trees to be held.An equal weight analysis resulted in 18 equally parsimonious trees, each of 73 steps, consistency index = 0.575, and retention index = 0.613.An implied weight analysis (k = 3) produced three trees, and the strict consensus tree (Fig. 1) had 74 steps, consistency index = 0.568 , and retention index = 0.600.Critical in the analysis was the position of Larrisson nedymus that appeared as the sister species of Sericophorus relucens, and not a part of the clade encompassing the remaining Larrisson.The same position was retained in another analysis, in which Aha ha Menke was added as another outgroup.This result demonstrates that L. nedymus is not congeneric with the remaining Larrisson and that it either should be transferred to Sericophorus or that it belongs to a genus of its own.Given the degree of differences between L. nedymus and Sericophorus, I believe it should be treated as a separate genus.A new genus, Larrissa is established below for the species.Name-derivation.Armatus is a Latin masculine adjective meaning armed, with reference to the lateral spines on male tergum VII.
Recognition.Larrisson armatus is unique in having the impunctate, medioventral area of the clypeus narrow and elongate, extending dorsally to the clypeal midlength of more.The male is also unique in having tergum I with a gap between the basolateral carina and the lateral ridge that delimits the basal concavity (Fig. 2c) and an anterolateral spine on tergum VII (Fig. 2d).Subsidiary recognition features are: orbital fovea well defined (in female about as wide as 0.8 × ocellocular distance), mesothoracic venter densely punctate throughout, gaster all red, and male sternum VIII emarginate apically, with glabrous basal platform.
Description.Width of face across clypeus and vertex in female = 60:46-50, least interocular distance 44-45; in male 60:48, and 42, respectively.Orbital fovea well defined, in female about as wide as 0.8 × ocellocular distance, in male not quite half ocellocular distance.Clypeal lobe only slightly prominent, its free margin arcuate, slightly angulate laterally; medioventral asetose area narrow, extending to about clypeal midlength in female (Fig. 2a) and slightly higher in male.Scapal basin impunctate, glabrous.Length of scape (excluding radicle) 2.3 × width in female, 2.5 × in male, length equal to flagellomeres I-III combined in female, to flagellomeres I-III + half IV combined in male.Mesopleural with small tooth at top of precoxal declivity in female, with well-defined tooth in male; mesothoracic venter uniformly densely punctate and setose (punctures about one diameter apart, setae concealing integument).Metanotum without median tooth.Propodeal dorsum without spine or tubercle behind spiracle; side punctate, minutely ridged below spiracle; posterior surface unsculptured both mesodorsally and mesoventrally.Outer surface of hindtibia impunctate between spines.
Female.Forebasitarsus with four rake spines, apical spine of foretarsomere III about equal to apical basitarsal width.Pygidial plate with punctures that are about one diameter apart.Length 5.4-5.5 mm.
Geographic distribution (Fig. 3).Name-derivation.Carinatus is a Latin masculine adjective derived from carina, with reference to the longitudinal carina on male sternum II.
Recognition.The male of carinatus is unique in having a sharply pointed apically median carina in the basal half of sternum II (Fig. 4c).It can also be recognized by the combination of a largely red gaster, metanotum without a spine or tubercle, propodeum without a spine or tubercle behind the spiracle, presence of a spine in front of the midcoxa, and concave ventral surface of sternum VIII.The female is unknown.
Setae all silvery, appressed on head, thorax, propodeum, and legs, nearly completely concealing integument on clypeus (except glabrous ventral portion of median lobe) and frons (except for glabrous scapal basin), not concealing integument on mesopleuron or mesothoracic venter, forming apical fasciae on terga I-IV.Hindfemoral venter and inner (= posterior) surface asetose.Head, thorax, and propodeum black except the following are pale yellow: scape, pedicel, mandible (except apically), and pronotal lobe, whereas glabrous portion of clypeal lobe brown, and flagellum yellowish brown (darkened dorsally on at least basal half).Forefemur reddish brown dorsally, yellow ventrally and apically; midfemur reddish brown except yellow apically and ventrally in distal half; hindfemur reddish brown except yellow apically; foretibia yellow on outer side, reddish brown on inner side; mid-and hindtibiae varying from mostly pale yellow to mostly reddish brown; foretarsus reddish brown; mid-and hindtarsi pale yellow except apical tarsomere dark brown.Gaster reddish brown except terga III-VI or IV-V with black basal spots that may be interrupted mesally.
Posterior mandibular margin not expanded between base and notch, inner margin without tooth near midlength (Fig. 4a).Flagellum cylindrical; dorsal length of flagellomere I 1.1 × apical width.Propodeum without spine behind spiracle.Legs unmodified except hindfemur slightly expanded posteroventrally at apex, not concave ventrally and not carinate between ventral and posterior (= inner) surfaces; forebasitarsus with five rake spines; apical spine of forebasitarsus III 1.1 × as long as apical basitarsal width.Tergum VII punctate throughout, rounded apically (Fig. 4d).Sternum II elevated in basal half, concave in distal half, with obtuse median carina in basal half (Fig. 4c), carina pointed apically and projecting over concave portion.Sterna with long, erect setae at bases of apical depressions, sterna VI and VII also with numerous erect setae that  Name-derivation.Latifrons derives from two Latin words, latus¸ broad, and frons, the forehead; a noun in apposition to the generic name.
Recognition.The female of latifrons (the male is unknown) has an all black gaster (without red markings or yellow fasciae), and the setae appressed on the head, thorax, propodeum and legs.Larrisson niger is similar, but unlike that species the hindfemoral apex of latifrons is simple, not broadened (Fig. 7b), the scutum and mesopleuron are dull, with interspaces between punctures linear, and the setae of the pygidial plate do not conceal the integument.
Forebasitarsus with four rake spines, apical spine of foretarsomere III about equal to apical basitarsal width.Pygidial plate with punctures that are more than one diameter basally, less than one diameter apart apically.Length 4.8-6.1 mm.
Male.Unknown.Geographic distribution (Fig. 6).Known from one locality in northern Queensland.Recognition.Like latifrons, the female of niger has an all black gaster, without red markings or yellow apical fasciae on segments, and the setae appressed on the head, thorax, propodeum and legs.Unlike latifrons, however, the hindfemoral apex of niger is broadened (Fig. 8b) rather than simple, scutal and mesopleural punctures average about one diameter apart and the interspaces are shiny (scutum and mesopleuron dull in latifrons, with linear interspaces), and the setae of the pygidial plate are dense, largely concealing the integument (rather than sparse, not concealing integument).The male is unknown.Description (based on holotype only).Female.Width of face across clypeus and vertex = 60:50, least interocular distance 37. Orbital fovea well defined, slightly wider than half ocellocular distance.Clypeal lobe only slightly prominent, its free margin arcuate, not angulate laterally.Scapal basin sparsely punctate (punctures averaging several diameters apart).Length of scape (excluding radicle) 2.8 × width, length equal to flagellomeres I-V combined.Scutal and mesopleural punctures averaging about one diameter apart (Fig. 8a).Mesopleuron with ill-defined transverse crest in front of midcoxa; mesothoracic venter densely punctate throughout (punctures about one diameter apart).Metanotum with obtuse median carina.Propodeum with minimal, obtuse tubercle behind spiracle; side ridged; posterior surface almost unsculptured mesodorsally, unsculptured mesoventrally.Hindfemoral apex broadened (Fig. 8b).Outer surface of hindtibia with small setigerous punctures, including dorsal half (punctures sparse in distal half or so of dorsal half ).
Forebasitarsus with five rake spines, apical spine of foretarsomere III minimally shorter than apical basitarsal width.Pygidial plate with punctures that are about one diameter apart basally, less than one diameter apart apically.Length 4.5 mm.
Recognition.The female of orbitalis has a red gastral base and a densely punctate mesothoracic venter, with the integument totally concealed by vestiture.Three other species, armatus, quintus and variegatus, share these characters, but orbitalis differs from quintus in lacking dense, small punctures on the outer surface of the hindtibia (between the spines); unlike armatus, the unsculptured medioventral area of the clypeus does not extend to clypeal midlength; and unlike variegatus, the inner eye margins of orbitalis are markedly bowed toward the frons midline (Fig. 9a), rather than nearly parallel, and the first recurrent vein is received by the first submarginal cell, rather than the second.
The male of orbitalis shares with quintus the posterior mandibular margin that is angulate between base and notch (Fig. 9c).Unlike that species, the legs of orbitalis are unspecialized: the forefemoral venter is not expanded subbasally and not concave anterobasally, the inner margin of the forebasitarsus is straight, the foretarsomeres II-IV are not expanded on the inner side, and the hindbasitarsus is not convex on outer margin.The emarginate apically sternum VIII, with a glabrous basal platform, is a subsidiary recognition feature of the male orbitalis.
Description.Width of face across clypeus and vertex = 60:52, least interocular distance 50 in female, in male, respectively, 60:58 and 40.Orbital fovea ill defined, about as wide as half ocellocular distance in female, less than that in male.Inner eye margin markedly bowed out toward frons midline (Figs 9a, b).Clypeal lobe only slightly prominent, its free margin arcuate, rounded laterally.Scapal basin punctate along margins in female, impunctate in male.Length of scape (excluding radicle) 2.5 × width, length equal to flagellomeres I-IV + half V combined in female, to flagellomeres I-III + half IV combined in male.Scutal punctures less than one diameter apart.Mesopleuron with ill-defined tubercle at top of precoxal declivity in female, with obtuse, transverse carina in front of midcoxa in male; mesothoracic venter densely punctate throughout (punctures less than one diameter apart).Metanotum with low median carina in female, with well-defined median tooth in male.Propodeum with tubercle behind spiracle (tubercle ill defined in female); side conspicuously ridged; posterior surface ridged both mesodorsally and mesoventrally in female, with irregular sculpture mesodorsally and not ridged medioventrally in male.Outer surface of hindtibia impunctate and asetose between spines in dorsal half.Tergum I concave basally.
Setae all silvery, appressed on head, thorax, propodeum, and legs, nearly completely concealing integument on clypeus (except glabrous ventral portion of median lobe), on frons lateroventrally (on scapal basin markedly shorter than on remaining frons and not concealing integument in female, absent in male), on mesopleuron, and in female on mesothoracic venter (not so in male); in female setae of pygidial plate light brown, not concealing integument.Hindfemoral venter asetose, inner (= posterior) face setose except asetose along dorsal margin in male.
Head, thorax, and propodeum black except the following are pale yellow: scape (only ventrally in male), mandible basally (black apically), and pronotal lobe; flagellum black dorsally, brown ventrally in female, brown dorsally and yellow ventrally in male (dark brown basally, light brown apically, flagellomeres X-XIII all light brown).Color of legs and gaster: see below.
Female.Forebasitarsus with four rake spines, apical spine of foretarsomere III about equal to 0.7 × apical basitarsal width.Pygidial plate with punctures that are more than one diameter basally, less than one diameter apart apically.Length 7.4 mm.Forefemur black basally, yellow apically, mid-and hindfemora reddish brown, yellow apically; foretibia reddish brown on inner surface, yellow on outer surface; midtibia reddish brown except yellow apically; hindtibia reddish brown; forebasitarsus yellowish brown, foretarsomeres II-V brown; mid-and hindtarsi reddish brown.Gastral terga I and VI reddish brown, tergum II reddish brown with black basomedian spot, terga III and V black except reddish brown apically, tergum IV black Male.Posterior mandibular margin angulate between base and notch, concave adjacent to notch (Fig. 9c).Flagellum cylindrical; dorsal length of flagellomere I 1.4 × apical width.Tergum VII rounded apically.Sternum II with transverse, glabrous swelling behind midlength (swelling higher laterally than mesally).Length 6.3 mm.Fore-and midfemora black basally and dorsally (except at apex), pale yellow ventrally (except near base) and apically; hindfemur black except pale yellow near apex; tibiae and tarsi pale yellow.Gaster black except tergum I and large median part of tergum II reddish brown and except apical depression conspicuously yellowish (inconspicuously so on tergum I).
Geographic distribution (Fig. 6).Known from two localities in western New South Wales.Recognition.The female of punctatus differs from all its congeners by the markedly depressed mesothoracic venter and larger gastral punctures; in particular, the punctures of the basal concavity of tergum I are almost as large as those on the scutum (Fig. 10a, b).In all other Larrisson the mesothoracic venter is only slightly depressed and the basal concavity of tergum I is either impunctate, or has minute, inconspicuous punctures, or (niger) the punctures are visibly smaller than those on the scutum.Also, like tegularis and unlike other Larrisson, the scapal basin is densely punctate (punctures less than one diameter apart), rather than impunctate or with punctures that average several diameters apart.As in tegularis, most of the propodeal enclosure is covered with  setae, only the median sulcus and the adjacent area being glabrous; in all other Larrisson, the propodeal enclosure is glabrous (all or nearly so).The absence of a metanotal tubercle or crest is a subsidiary recognition feature.The male is unknown Description.Female.Width of face across clypeus and vertex = 60:55-58, least interocular distance 28.Orbital fovea narrow, about one quarter width of ocellocular distance.Clypeal lobe only slightly prominent, its free margin arcuate, angulate laterally.Scapal basin punctate, punctures less than one diameter apart.Length of scape (excluding radicle) 2.3-2.4 × width, length equal to flagellomeres I-IV combined.Mesopleuron without transverse crest or tubercle in front of midcoxa; mesothoracic venter conspicuously depressed, densely punctate mesally (punctures about one diameter apart, up to several diameters apart sublaterally).Metanotum without median tubercle or crest.Propodeum without spine or tubercle behind spiracle; side ridged; posterior surface sculptured mesodorsally and mesoventrally.Outer surface of hindtibia with small setiferous punctures, glabrous adjacent to dorsal margin (except in basal half ).Tergal punctures (Fig. 10a, b) larger than in other Larrisson, those of basal concavity of tergum I (which is well defined) almost as large as scutal punctures.
Head (including flagellum), thorax, propodeum, and gaster black except scape and pronotal lobe pale yellow.Femora either black, brown at very apex, or largely reddish brown; tibiae either light brown, yellow at very apex, or all pale yellow; tarsi yellowish brown.Gaster black in specimens from Calperum Station, with the following reddish brown: basal concavity of tergum I largely, narrow preapical stripes on terga, and tergum VI; in specimen from Grevillea camp terga I and II reddish brown, terga III and IV reddish brown with black basomedian spot, tergum V black basally, and tergum VI yellow, becoming brownish apically; apical depressions of terga yellowish brown in all three specimens examined.
Forebasitarsus with four rake spines, apical spine of foretarsomere III equal to 1.25 × of apical basitarsal width.Pygidial plate with punctures that are more than one diameter apart except less than one diameter apart near apex.Length 5.3-5.5 mm.
Male.Unknown.Geographic distribution (Fig. 11).Known from two localities in South Australia.Specimens examined.Holotype: E, AUSTRALIA: South Australia: Calperum Station 32 km N Renmark at 33°53'S, 140°44'E, 9 Nov -12 Dec 1995, K.R. Pullen (ANIC).Paratypes: AUSTRALIA: South Australia: same data as holotype (1 ♀, CAS); Grevillea WAT camp at 27°01'30"S, 129°52'32"E, 18-20 Oct 1996, Pitjantjatjara Land Survey (1 ♀, SAM).outer surface of the hindtibia.In addition, the mesothoracic venter is densely punctate and setose (punctures about one diameter apart, setae concealing integument), at least tergum I is reddish brown, and in the female most setae of the pygidial plate do not conceal the integument.In most other Larrisson, the outer surface of the hindtibia is impunctate at least in dorsal half or has a few, sparse punctures, although it is punctate and setose in punctatus, spinosus, niger, and tegularis; in the first two species, the punctures of the mesothoracic venter are 2-3 diameters apart on each side of the median zone, and the setae do not conceal the integument; in niger, the gaster is all black and most setae of the female pygidial plate conceal the integument (the male is unknown); in tegularis, gastral terga have apical yellow fasciae and the tegular inner margin is concave (rather than evenly rounded.

Larrisson quintus
The male of Larrisson quintus differs from all its congeners in having the legs markedly modified: the forefemoral venter is roundly expanded subbasally (Fig. 13b), concave anterobasally, the forebasitarsus concave on the inner margin (Fig. 13c), foretarsomeres II-IV are expanded on the inner side (Fig. 13c), midtarsomeres II-IV wider than long (Fig. 13e), and the hindbasitarsus is convex on the outer margin (Fig. 13f ); as in orbitalis, the posterior mandibular margin is angulate between base and notch (Fig. 13a); as in sulcatus and tibialis, the hindfemur is concave ventrally (Fig. 13d).
Description.Width of face across clypeus and vertex in female = 60:50-52, least interocular distance 47-48; in male 60:50, and 45, respectively.Orbital fovea well defined, in female more than half ocellocular distance, in male not quite half ocellocular distance.Clypeal lobe only slightly prominent, its free margin arcuate, rounded laterally in female (Fig. 12a), slightly angulate in male (Fig. 12b).Scapal basin impunctate, glabrous.Length of scape (excluding radicle) 2.2-2.3 × width in female, 2.8-2.9 × in male, length equal to flagellomeres I-IV + half V combined.Precoxal mesopleural declivity simple, not expanded into spine or tubercle; mesothoracic venter uniformly densely punctate and setose (punctures about one diameter apart, setae concealing integument).Metanotum with median tooth that is vestigial in females but well defined (up to about 1.3 × midocellar width) in males.Propodeal dorsum without spine or tubercle behind spiracle in female, with conspicuous tubercle or obtuse spine in male; side minutely ridged adjacent to metapleural sulcus (ridges larger under spiracle), with several punctures near middle, punctate posteriorly and also on posterior surface laterally; posterior surface ridged both mesodorsally and mesoventrally.Outer surface of hindtibia (except in one male from Ellery Creek Big Hole, WMNP) with many small setiferous punctures between spines, including dorsal half.
Head, thorax, and propodeum black in most specimens with the following exceptions: clypeus reddish brown ventrally (narrowly so on lateral lobes); mandible yellow basally, dark brown apically; scape yellow (black dorsally), all black in female from Heathlands, Queensland; flagellum reddish brown at least ventrally (all reddish brown in specimens from Victoria River Roadhouse); thorax and propodeum reddish brown to varying degree in several specimens from WMNP (only scutum black in one female).Forefemur in most females black basally, yellow apically, in most males yellow, with black spot in basal three quarter of length on posterior surface (black replaced by reddish brown in most females and single male from WMNP, forefemur all reddish brown in one female from there); midfemur in most females reddish brown anteriorly and black posteriorly, except yellow apically and ventrally in distal half or third, in most males yellow anteriorly and ventrally, reddish brown posteriorly, dark brown dorsally (midfemur all reddish brown in specimens from WMNP, all black in female from Heathlands); hindfemur in most females reddish brown except black dorsally and yellow at very apex, in most males reddish brown except yellow apically and dark brown on posterior (= inner) surface in distal half, also dorsally in some specimens (hindfemur all reddish brown in specimens from WMNP except yellow apically in single male, all black in female from Heathlands); tibiae yellow (foretibia dark brown on ventral surface, mid-and hindtibiae dark brown on posterior surface); tarsi yellow in most specimens (apical tarsomeres reddish brown, brown in specimen from Heathlands), reddish brown in females from WMNP.Gaster all reddish brown in specimens from WMNP, but terga II and III largely black mesally and terga IV and V (IV-VI in male) black except laterally in those from Gregory National Park and Maud River, and only tergum I reddish brown in specimen from Heathlands; apical depressions of terga II-V (II-VI in male) reddish brown.
Female.Pygidial plate with punctures that are less than one diameter apart in apical half or third (Fig. 12c).Forebasitarsus with four or five rake spines; apical spine of foretarsomere III about 1.3 × as long as apical basitarsal width.Length 5.1-6.5 mm.
Geographic variation.In specimens from Queensland and from Gregory National Park and Maud River, Northern Territory, the basolateral carina of tergum I is continued mesad by a short, oblique carina, in the female the glabrous, apicomedian portion of the clypeus is convex, the femora are darker, while the male forebasitarsus has a lamellar, translucent expansion on the outer margin in the distal half.In speci-mens from WMNP, the additional tergal carina is absent, the apicomedian portion of the female clypeus is concave, the legs are more reddish brown, and the male forebasitarsus is not expanded on the outer margin.In females from Western Australia, the additional basolateral carina on tergum I is absent, the glabrous, apicomedian portion of the clypeus is flat, and the femora are more reddish brown in the specimen from Pardoo Roadhouse area.
Geographic distribution (Fig. 11).Northern Australia.Specimens examined.Holotype: ♂, AUSTRALIA: Northern Territory: Gregory National Park: Victoria River bank near Timber Creek at 15°37.8'S, 130°28.6'E(Fig. 15 Name-derivation.Spinosus is a Latin masculine adjective meaning spiny; with respect to the spine on the metanotum and another behind the propodeal spiracle, the structures that differentiate this species from carinatus. Recognition.The male of spinosus resembles carinatus and quintus in having gastral terga I and II reddish brown rather than black combined with nonemarginate apically sternum VIII.It differs from these species in having a more prominent middle clypeal lobe (Fig. 16a).Unlike quintus, the legs are unmodified in spinosus (see quintus for details), and unlike carinatus the mesopleuron has a sharp median tooth in front of the midcoxa (rather than a transverse crest), the metanotum has a sharp middle spine (Fig. 16b, spine absent in carinatus), the propodeum has a spine behind the spiracle (spine absent in carinatus), and sternum II has a transverse swelling (rather than a median, pointed carina).The female is unknown.
Setae all silvery, appressed on head, thorax, propodeum, and legs, partly concealing integument on clypeus and pronotal collar, concealing integument from most angles on lower frons except for glabrous scapal basin, concealing or not concealing on mesopleuron, not concealing on mesothoracic venter, forming ill-defined apical fasciae on terga I-III.Hindfemoral ventral surface asetose, inner (= posterior) surface setose except ventrally in holotype and specimen from Calperum Station.
Head, thorax, and propodeum black except the following are pale yellow: scape, pedicel, mandible (except apically), and pronotal lobe, whereas flagellum is light brown ventrally and light brown to black dorsally.Forefemur black basally (reddish brown in specimen from Cocata Conservation Park), pale yellow apically and in apical half ventrally; midfemur reddish brown basally, pale yellow apically and in apical third ventrally; hindfemur reddish brown, pale yellow apically; tibiae pale yellow, partly reddish brown; forebasitarsus pale yellow (reddish brown on inner surface), remaining article yellowish brown; mid-and hindtarsi pale yellow except apical tarsomeres light brown.Gastral terga I, II, and VII reddish brown, remaining terga reddish brown laterally and on apical depressions, otherwise black.
Posterior mandibular margin not expanded between base and notch, inner margin with obtuse tooth near midlength (Fig. 16a).Flagellum cylindrical; dorsal length of flagellomere I about equal to apical width.Metanotum with conspicuous spine (Fig. 16b).Propodeum with conspicuous spine behind spiracle (spine longer than that on metanotum).Legs unmodified except hindfemur concave ventrally, slightly expanded ventrad at apex, carinate between ventral and posterior (= inner) surfaces; forebasitarsus with four rake spines; apical spine of foretarsomere III equal to apical width of basitarsus.Tergum VII punctate throughout, rounded apically.Sternum II with transverse swelling behind midlength, swelling glabrous, similar to that of quintus.Sterna III-VII with long, erect setae at bases of apical depressions, otherwise practically asetose.Sternum VIII flat, glabrous, rounded apically.Length 8.4-9.6 mm.Genitalia similar to those of quintus.
Female.Unknown.Geographic distribution (Fig. 11).South Australia and Western Australia.Recognition.The male of sulcatus differs from all other species of Larrisson by the presence of a pair of longitudinal sulci on the posterior propodeal surface that are convergent ventrad (Fig. 17a), and a triangular rather than open anteriorly second submarginal cell (Fig. 17b).The female is unknown.
Description (based on holotype only).Male.Width of face across clypeus and vertex = 60:58, least interocular distance 44.Orbital fovea ill defined, narrower than half ocellocular distance.Clypeal lobe only slightly prominent, its free margin arcuate, obtusely angulate laterally.Scapal basin impunctate.Length of scape (excluding radicle) 2.2 × width, length equal to flagellomeres I-IV + half V combined.Mesopleural punctures less than one diameter apart; impunctate, low tubercle present in front of midcoxa; mesothoracic venter densely punctate throughout (punctures less than one diameter apart).Metanotum with median tubercle.Propodeal side with well-defined ridges; posterior surface ridged mesodorsally, finely rugose mesoventrally, with pair of longitudinal sulci that are converging ventrad (Fig. 17a).Forewing vein M diverging from M+Cu distad of cu-a by about 0.5 length of cu-a; second submarginal cell triangular (Fig. 17b).Outer surface of hindtibia impunctate and asetose between spines along dorsal margin.Tergum I concave basally, concavity with well-defined median line.Setae all silvery except golden beneath midocellus, appressed on head, thorax, propodeum, and legs, nearly completely concealing integument on clypeus (except glabrous ventral portion of median lobe) and on ventral half of frons laterally (except for glabrous scapal basin), not concealing integument on mesopleuron or mesothoracic venter, not forming well-defined apical fasciae on terga.Hindfemoral venter and inner (= posterior) surface setose.
Head, thorax, and propodeum black except the following are pale yellow: scape, part of pedicel, mandible basally (apex dark brown), pronotal lobe, and humeral plate of wing base; flagellum black dorsally, brown ventrally (two apical flagellomeres all brown).Femora black basally, yellow apically; tibiae and tarsi yellow.
Posterior mandibular margin not expanded between base and notch, inner margin with small tooth near midlength.Flagellum cylindrical; dorsal length of flagellomere I about equal to apical width.Propodeum without spine or tubercle behind spiracle.Legs unmodified; forebasitarsus with four rake spines; apical spine of forebasitarsus III as long as apical basitarsal width.Sternum II without transverse swelling, but with low, transverse convexity behind midlength.Sterna III-VII with long, erect setae at bases of apical depressions, sterna V-VII also with erect setae on remaining surface.Sternum VIII punctate and setose along margin, rounded apically.Volsella ending shortly before apex of penis valve.Length 6.4 mm.
Female.Unknown.Geographic distribution (Fig. 18).Known from one locality in northwestern Northern Territory, Australia.
Recognition.Larrisson tegularis has a reddish brown gaster with yellow apical bands on terga, a unique such coloration (yellow bands somewhat obscured by vestiture).It also has a unique tegula: elongate, with concave inner margin (Fig. 19b).In the female, the punctures of the pygidial plate are sparser than in all the congeners, averaging more than one diameter apart mesally, and the setae do not conceal the integument (Fig. 19c).In the male, flagellomere I is shorter than in the congeners (dorsal length equal to 0.8 × apical width, rather than at least equal) and sternum VIII is unique: it is emarginate apicomesally (Fig. 19d), but less so than in armatus, orbitalis, quintus, and variegatus, and unlike these species the ventral surface is all flat, punctate throughout (rather than with a glabrous, raised platform basomedially).
Female.Punctures of pygidial plate averaging more than one diameter apart mesally, setae not concealing integument.Forebasitarsus with three or four rake spines; apical spine of foretarsomere III about 1.4 × as long as apical basitarsal width.Length 4.2-4.9mm.
Male.Posterior mandibular margin not expanded between base and notch.Flagellum cylindrical; dorsal length of flagellomere I 0.8 × apical width.Legs unmodified; forebasitarsus with four rake spines; apical spine of forebasitarsus III as long as apical basitarsal width.Sternum II without transverse swelling; apex of sternum IV and all sterna V-VIII with dense, erect setae; sternum VIII flat, all punctate, shallowly emarginate apically (Fig. 17d).Genitalia unlike those of other Larrisson, with broad, rounded gonocoxite and setae invisible from above (Fig. 20).Length 4.2 mm.
Geographic distribution (Fig. 3).Known from one locality in southwestern New South Wales.Name-derivation.Tibialis is a Latin adjective derived from tibia; with reference to the unusual male hindtibia of this species.
Recognition.The male of tibialis (the female is unknown) has uniquely modified hindtibia, thickened on the outer side at about one third length (the thickening is best seen in dorsal view, Fig. 21a), and the lateral surface has three unevenly spaced spines: two at the thickening and one near the apex (Fig. 21a, b).In the other Larrisson, the hindtibia is not thickened, and the spines on the lateral surface are evenly spaced.Subsidiary recognition characters of tibialis are: propodeal dorsum with spine behind spiracle and sternum II with arcuate swelling.
Description (based on holotype only).Male.Width of face across clypeus and vertex = 60:58, least interocular distance 35.Orbital fovea well defined, narrower than half ocellocular distance.Clypeal lobe only slightly prominent, its free margin arcuate, not angulate laterally.Scapal basin impunctate except punctate along inner margin.Length of scape (excluding radicle) 2.8 × width, length equal to flagellomeres I-V combined.Flagellomeres cylindrical.Mesopleuron with convexity in front of midcoxa; mesothoracic venter sparsely punctate on each side of median zone (punctures several diameters apart).Metanotum with small median tooth.Propodeal side ridged; posterior surface ridged both mesodorsally and medioventrally.Outer surface of hindtibia sparsely punctate and setose between spines.Tergum I concave basally.
Posterior mandibular margin not expanded between base and notch, inner margin with small tooth near midlength.Flagellum cylindrical; dorsal length of flagellomere I about equal apical width.Propodeal dorsum with spine behind spiracle.Legs unmodified except midtibia slightly curved near basis, hindfemur with small emargination near apex of externoventral (= anteroventral) margin, carinate between ventral and posterior (= inner) surfaces, venter concave; hindtibia thickened on the outer side at about one third length (thickening best seen in dorsal view, Fig. 21a), with densely punctate area on lateral surface between thickening and apex that is gradually enlarging toward apex, and lateral surface with three unevenly spaced spines: two at thickening and one near apex (Fig. 21a, b); forebasitarsus with four rake spines; apical spine of forebasitarsus III as long as apical basitarsal width.Sternum II with arcuate swelling, anterad of swelling with dense, erect setae that are shorter than midocellus width; sterna III-VII with long, erect setae at bases of apical depressions, sterna VI and VII also with numerous erect setae that are about as long as midocellar width.Sternum VIII punctate and setose along margin, rounded apically.Genitalia similar to those of quintus except volsella ending shortly before apex of penis valve.Length 9.5 mm.
Recognition.Unlike all other Larrisson, the first recurrent vein of variegatus is received by the second submarginal cell (Fig. 23) rather than the first.The species is further characterized by an all or largely reddish brown gaster and a densely punctate mesothoracic venter, with the integument totally concealed by vestiture, two features shared with orbitalis and quintus.Unlike quintus, variegatus lacks small, dense punctures and setae between spines on the outer surface of the hindtibia (except ventrally).Unlike orbitalis, the inner eye margin are almost parallel above the level of the antennal socket (Fig. 22a) rather than markedly bowing toward the frons midline.The male has three unique characters: 1. multicolored antenna (Fig. 22b, c), with flagellomeres I-VI convex ventrally (Fig. 22c); 2. an unusually large basal concavity of tergum I that extends to both lateral margins, is largely asetose, and is bordered laterally by the basolateral tergal carina up to its dorsal end (Fig. 22d, e); and 3. sternum II with a transverse swelling that is markedly curved posterad (Fig. 22f ).The emarginate apically male sternum VIII is a subsidiary recognition feature, shared with armatus, orbitalis, quintus, tegularis, and also with Larrissa nedyma.
Description.Width of face across clypeus and vertex in female = 60:49-51, least interocular distance 34-35; in male 60:48-51, and 32-33, respectively.Orbital fovea in female well defined but narrow, less than half ocellocular distance, in male varying from well defined (as narrow as in female) to nearly absent.Clypeal lobe only slightly prominent, its free margin arcuate, not angulate laterally.Length of scape (excluding radicle) 2.7 × width in female, 2.8-2.9 × in male, length equal to flagellomeres I-IV combined.Mesopleuron with obtuse, transverse crest in front of midcoxa; mesothoracic venter uniformly densely punctate, punctures about one diameter apart.Metanotum with rudimentary median carina.Propodeal dorsum without spine or tubercle behind spiracle; side finely, densely ridged; posterior surface unsculptured both mesodorsally and mesoventrally.First recurrent vein received by second submarginal cell (Fig. 23).Outer surface of hindtibia impunctate and asetose between spines (except basally and ventrally).
Setae all silvery, appressed on head, thorax, propodeum, and legs, concealing integument on clypeus, large part of frons in female and most of frons in male (except for glabrous scapal basin), largely concealing integument on mesopleuron and mesothoracic venter in female, totally so in male.Hindfemoral venter asetose, inner (= posterior) surface asetose except setose dorsally.
Head, thorax, and propodeum black except scapal venter and pronotal lobe yellow, also mandibular base in female (yellowish in male).See below for color of flagellum, legs, and gaster.
Female.Inner eye orbits nearly parallel between antennal socket and midocellus level (Fig. 22a); hindfemoral venter slightly concave near apex and with a few setae.Length 5.0 mm.Flagellum brown dorsally, reddish brown ventrally; fore-and midfemora reddish brown, pale yellow ventrally (except near base), hindfemur reddish brow, pale yellow at very apex; tibiae reddish brown, pale yellow dorsally; tarsi reddish brown.Gaster all reddish brown.
Male.Flagellomeres I-VI convex ventrally (Fig. 22c), dorsal length of flagellomere I equal to apical width.Posterior mandibular margin slightly concave between base and notch.Forefemur flat ventrally.Basal concavity of tergum I unusually large, largely asetose, extending to both lateral margins (Fig. 22d), bordered laterally by basolateral tergal carina attaining concavity's dorsal end (Fig. 22e).Sternum II with transverse swelling that is markedly curved posterad (Fig. 22f ); sterna III-VII with long, erect setae at bases of apical depressions and also with shorter, erect setae on remaining surface; sternum VIII emarginate apically, with glabrous, slightly elevated platform that covers most of its surface, punctate and setose along margins outside platform.Length 7.5-8.4mm.Flagellomeres I-VI reddish brown, VII black, VIII and IX pale yellow, X and XI brown (Fig. 22b, c).Forefemur reddish brown, pale yellow ventrally, with some black areas basodorsally, midfemur black dorsally, pale yellow ventrally, with intermediate areas reddish brown, hindfemur black, pale yellow at very apex; tibiae reddish brown, pale yellow dorsally; tarsi reddish brown.Gaster largely reddish brown, but terga II-VII with lateral spots that become large toward apex (black areas larger on terga V and VI than red median zone).
Geographic distribution (Fig. 18).Known from two localities in South Australia.

Larrisson abnormis (R. Turner)
Sericophorus abnormis R. Turner, 1914: 352 In spite of these differences, I consider them to be individual or geographic variants of one species, rather than members of two different species, as they share a number of unique characters: yellow antennae (Fig. 24a), dense, directed upward frontal setae, a flattened, concave midfemoral venter, obtusely carinate along both anterior and posterior margins (Fig. 24b), and an anteroventral hindfemoral margin that is slightly expanded preapically, with a row of small setae emerging from the expansion.They also share the following, non-unique characters: metanotum with median tooth, propodeum with prominent spine behind spiracle, and sternum II with transverse swelling.
Description.As indicated under Phylogenetic Analysis above, Larrisson nedymus does not cluster with either the remaining Larrisson or Sericophorus and requires a genus of its own.
Like Larrisson, Larrissa is a member of the tribe Miscophini because of the round, not modified, hind ocellus and the simply attenuate hindfemur in combination with the emarginate posterior mandibular margin.It is characterized by the presence of two discoidal and three submarginal cells, the second submarginal not petiolate but distinctly narrowing toward the front margin, and the first recurrent vein ending on the first submarginal cell.
Larrissa appears most closely related to Larrisson and Sericophorus.Like Larrisson, it differs from Sericophorus by the following: the posterior propodeal surface has no medioventral carina (carina present in Sericophorus), the male flagellum has 11 articles (10 in Sericophorus), a volsella is present (absent in Sericophorus), and the occipital carina does not join the hypostomal carina (joins in the vast majority of Sericophorus).It also differs in having a mesopleuron abruptly angular below the pronotal lobe, the mesothoracic venter conspicuously concave mesally, and the female gena with an angular bulge near the middle.
Unlike Larrisson, Larrissa has the following: frons uniformly punctate and setose (without glabrous scapal basin, Fig. 25a); length of scape 1.1-1.3× maximum width; inner mandibular margin without preapical tooth, but with two teeth near midlength (Fig. 25a); mesopleuron abruptly angular below pronotal lobe; forewing vein M diverging from M+Cu basad of cu-a; basolateral carina of tergum I expanded into lamella; female gena with angular bulge near middle; female tergum VI with impunctate, glabrous marginal lamella (Fig. 26a); male forecoxa with apical spine and foretrochanter excavated basally, volsella ending at half length of penis valve, and head of penis valve dentate.In Larrisson, the frons has an impunctate or sparsely to densely punctate scapal basin above each antennal socket (Figs 25b-d); length of scape (excluding radicle) is 2.3-2.8 × maximum width; inner mandibular margin with preapical tooth, at most with one obtuse tooth near midlength; mesopleuron rounded anteriorly, not abruptly angulate; forewing vein M diverging from M+Cu distad of cu-a or interstitial with cu-a; basolateral carina of tergum I not expanded into lamella; female gena without angular bulge; female tergum VI without impunctate, glabrous marginal lamella; male forecoxa and foretrochanter not modified, volsella ending shortly before apex of penis valve or exceeding apex of penis valve, and head of penis valve not dentate.

Figure 2 .
Figure 2. Larrisson armatus: a female clypeus b male antenna c male tergum I in dorsolateral view (arrow indicates basolateral emargination) d apex of male gaster in dorsal view (upper arrow indicates basolateral tooth of tergum VII, lower arrow indicates sternum VIII).

Figure 7 .
Figure 7. Larrisson latifrons ♀: a head in frontal view b hindfemur and base of hindtibia in lateral view.

Figure 9 .
Figure 9. Larrisson orbitalis: a female head in frontal view b male head in frontal view c male mandible in lateral view.

Figure 10 .
Figure 10.Larrisson punctatus ♀: a tergum I in dorsal view b tergum I in oblique anterodorsal view.
Figure 12.Larrisson quintus: a female clypeus and mandible b male clypeus and mandible c female pygidial plate d male tergum VII e male sternum VIII in ventral view f male genitalia in ventral view.

Figure 15 .
Figure 15.A collecting site near Timber Creek in Gregory National Park where most specimens of Larrisson quintus were caught (including the holotype).

Figure 19 .
Figure 19.Larrisson tegularis: a female head in frontal view b tegula (arrow indicates concave inner margin) c pygidial plate of female d apical sterna of male in ventral view e female gaster in dorsal view.

Figure 22 .
Figure 22.Larrisson variegatus: a female head in frontal view b male head in dorsal view c male antenna d male tergum I in dorsal view e male tergum I in oblique lateral view f gastral base of male in lateral view.

Figure 24 .
Figure 24.Larrisson rieki, ♂ from South Australia: a head in frontal view b midfemur in ventral view.

table 1 .
Character States of Larrisson and the Outgroup

Larrisson carinatus Pulawski, sp. n.
Known from one locality in New South Wales.

table 2 .
Comparison of the two known males of Larrisson rieki