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Revision of the Australo-Papuan genus Macrolema Baly (Coleoptera: Chrysomelidae: Spilopyrinae), with description of a new genus PDF

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Zootaxa 2486: 1–60 (2010) ISSN 1175-5326 (print edition) Monograph www.mapress.com/zootaxa/ ZOOTAXA Copyright © 2010 · Magnolia Press ISSN1175-5334(online edition) ZOOTAXA 2486 Revision of the Australo-Papuan genus Macrolema Baly (Coleoptera: Chrysomelidae: Spilopyrinae), with description of a new genus C. A. M. REID & M. BEATSON Department of Entomology, Australian Museum, 6 College Street, Sydney, NSW 2010, Australia. Email: [email protected]; [email protected] Magnolia Press Auckland, New Zealand Accepted by G. Morse: 16 Apr. 2010; published: 27 May 2010 C. A. M. REID & M. BEATSON Revision of the Australo-Papuan genus Macrolema Baly (Coleoptera: Chrysomelidae: Spilopyrinae), with description of a new genus (Zootaxa 2486) 60 pp.; 30 cm. 27 May 2010 ISBN 978-1-86977-543-8 (paperback) ISBN 978-1-86977-544-5 (Online edition) FIRST PUBLISHED IN 2010 BY Magnolia Press P.O. Box 41-383 Auckland 1346 New Zealand e-mail: [email protected] http://www.mapress.com/zootaxa/ © 2010 Magnolia Press All rights reserved. No part of this publication may be reproduced, stored, transmitted or disseminated, in any form, or by any means, without prior written permission from the publisher, to whom all requests to reproduce copyright material should be directed in writing. This authorization does not extend to any other kind of copying, by any means, in any form, and for any purpose other than private research use. ISSN 1175-5326 (Print edition) ISSN 1175-5334 (Online edition) 2 · Zootaxa 2486 © 2010 Magnolia Press REID & BEATSON Table of contents Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .3 Methods. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4 Allsortsiagen. nov.. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .5 Allsortsia maculata (Lea 1922) comb. nov. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .7 Key to the genera of Spilopyrinae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8 Macrolema Baly 1861. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .9 Key to species of Allsortsia Reid & Beatson and Macrolema Baly . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .11 Macrolema aenescens (Bowditch 1913) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .12 Macrolema albascuticasp. nov.. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 13 Macrolema atripennis (Bowditch 1913). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .14 Macrolema dickdaviesisp. nov. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .16 Macrolema giyasp. nov.. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .17 Macrolema karimuisp. nov. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .18 Macrolema longicornis Jacoby 1895 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .19 Macrolema metallica (Lea 1922) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .20 Macrolema pulchrasp. nov.. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 22 Macrolema quadrivittata (Jacoby 1898). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 23 Macrolema submetallica (Jacoby 1894) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 24 Macrolema ventralis (Lea 1921). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .26 Macrolema vittata Baly 1861 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 27 Acknowledgments .. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .29 References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .30 Figures 1–134 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .32–59 Abstract The chrysomelid genus Macrolema Baly 1861 is revised. 13 species are described including five new: Macrolema aenescens (Bowditch 1913), M. albascuticasp. nov., M. atripennis (Bowditch 1913), M. dickdaviesisp. nov., M. giya sp. nov., M. karimui sp. nov., M. longicornis Jacoby 1895, M. metallica (Lea 1922), M. pulchra sp. nov., M. quadrivittata (Jacoby 1898), M. submetallica (Jacoby 1894), M. ventralis Lea 1921b, M. vittata Baly 1861. Macrolema vittata is a senior synonym of M. marginata Jacoby 1898 (syn. nov.). Allsortsia gen. nov., is erected for a species formerly in Macrolema, Allsortsia maculata (Lea 1922), comb. nov. Lectotypes are designated for Macrolema longicornis, M. vittata and Macrogonus quadrivittatus. Keys are provided for identification of the genera of Spilopyrinae and for the species of Macrolema. Macrolema is confined to the eastern rainforests of Australia (11 species) and highland New Guinea (2 species). Little is known of the biology of the species of Macrolema and Allsortsia, all of which are relatively rarely collected. A first-instar larva is tentatively ascribed to Macrolema. Key words: fiery leaf beetle, morphology, Australia, New Guinea, taxonomy Introduction The chrysomelid subfamily Spilopyrinae (fiery leaf beetles) includes some of the most spectacularly coloured beetles in the world. However the group is relatively poorly known, as it has only recently been recognised and is confined to relatively remote regions of the east coast of Australia, the highlands of New Guinea, New Caledonia and southern South America (Reid 2000). Macrolema Baly 1861 was erected for a single Australian species which Baly placed in the Crioceridae, at that time poorly circumscribed, including genera from several unrelated subfamilies (Chapuis 1874; Reid 2000). Baly noted some similarity to Megascelis Sturm 1826, then also placed in Criocerinae but currently in Eumolpinae (Reid 1995, 2000). Chapuis (1874) suggested the greatest similarity was with Brachydactyla Lacordaire 1845 (now Pseudocrioceris Pic 1916) which is certainly a criocerine (material examined). Macrogonus Jacoby 1894, was erected for a similar curiously structured Australian criocerine, which only differed from Macrolema by the strongly produced pronotal lateral margins (see comments in Jacoby and REVISION OF THE AUSTRALO-PAPUAN GENUS MACROLEMA Zootaxa 2486 © 2010 Magnolia Press · 3 Clavareau 1904). By 1951, Macrolema and Macrogonus, with a total of 10 species, were still retained in Criocerinae (Monrós 1951). However the beetle morphologist Crowson was at this time working on a reclassification of the Coleoptera, and was in regular correspondence with Monrós, who was revising the world Criocerinae. Crowson noted similarity between Macrolema and other genera and placed these in Eumolpinae, as the most ‘primitive’ members of that subfamily (Crowson 1953). Monrós (1958) supported this placement, from his own studies. This group of genera was elevated to subfamily based on phylogenetic analysis of chrysomelid morphology (Reid 1995, 2000). The oldest family rank stem name is Spilopyr- Chapuis 1874, made available under Article 11.7 (International Commission on Zoological Nomenclature 1999) by Lefèvre (1885). Macrogonus, not mentioned by Crowson or Monrós, became a junior synonym of Macrolema in the world checklist of Chrysomelidae (Seeno & Wilcox 1982), probably on the strength of comments by Lea (1921a), and together these two names were placed incertae sedis in Eumolpinae. In a study of the phylogeny of Chrysomelidae based on traditional morphology (Reid 1995), the four putative tribes of Eumolpinae were analysed separately: ‘Eumolpini’ (equivalent to Eumolpinae excluding the next three taxa), Megascelidini, Synetini and Spilopyrini (including Macrolema). In this analysis larval characters were lacking for Megascelidini and Spilopyrini, but it was noted that evidence for monophyly of the four taxa was weak. The analysis was repeated after discovery of larvae for all the taxa (Reid 2000) and it was clear that Spilopyrini were sister to a larger entity than Eumolpinae alone, so the group was elevated to subfamily rank as Spilopyrinae. This status is supported in recent molecular phylogenies, although the exact relationship to other families remains unclear (Gomez-Zurita, Jolivet & Vogler 2005; Gomez-Zurita, Hunt, Kopliku & Vogler 2007; Gomez-Zurita, Hunt & Vogler 2008). A brief diagnosis was provided for Macrolema in the review of Spilopyrinae (Reid 2000), with a checklist of included species. The distinctive genus Richmondia Jacoby 1898 was noted as a possible synonym of Macrolema but is retained as a separate genus here pending a phylogenetic analysis of the whole subfamily. We have examined all type material, all specimens in the Natural History Museum, London, and all major collections in Australia for Macrolema. Other collections are unlikely to have significant numbers of specimens. There are 13 species, including 5 new and one new synonym, all of which are described below, with a key to the species. As a result of this study it was discovered that both the subfamilial and generic descriptions given by Reid (2000) need to be altered. All species of Macrolema lack a kotpresse, two have a reduced tibial spur formula and two species have some movement between the normally fused first two ventrites, states which weaken these characters in the diagnosis of Spilopyrinae. However other adult characters provided in the key to subfamilies of Chrysomelidae remain valid. The larval characters appear to be the best diagnostic tool for the subfamily. Here we tentatively identify a described but previously unknown larva as Macrolema (Reid 2000). One described species differs markedly from all other Macrolema species and other spilopyrines. We provide a new genus for this species below. Most Macrolema species are easily recognised from distinctive colour patterns combined with external morphology, however we provide full illustrations of male and female genitalia to aid further work on the group. Methods This is a traditional taxonomic paper based on descriptive morphology. Genitalia were prepared for examination by: removal of abdomen, maceration for 3–5 hours in cold 10% KOH, washing in water, cleaning the residue, transfer to glycerol on a cavity slide for microscopy. The cleaned abdominal sclerites were dried and pointed and the genitalia stored in glycerol in a polythene microvial, pinned with the rest of the specimen. Lengths are given to the nearest 0.5mm, but body and segment ratios were calculated from measurements made by eyepiece micrometer. Antennal segment ratios were measured from a single specimen of each sex. These ratios are indicative of variation between species rather than definitive, as precise boundaries of 4 · Zootaxa 2486 © 2010 Magnolia Press REID & BEATSON segments vary according to preservation of the specimens; furthermore most species are represented by few specimens. All measurements and ratios should therefore be regarded as approximate. All specimens were examined for surface sculpture, which is variable in most species of this genus. Traditional comparative terms for size and density of punctures are included in the descriptions, but are only interpretable by reference to the photographs. The punctures are variable between individuals within a species but extremes are distinct; contrast size and density of punctures in Figures 1-31. Male and female genitalia of each species are illustrated, from drawing-tube outlines made from preparations in glycerol on cavity slides. Due to lack of material only one or two specimens of each sex were dissected. The male internal sclerites include a small remnant of sternite VIII, the spiculum relictum (Lawrence, Beutel, Leschen & Slipinski 2010), which significantly varies between species. The penis is a simple slightly flattened tube, presenting few diagnostic characters, therefore generally only the apex is illustrated. In some specimens the endophallus was preserved in everted state (Figs 82, 87, 89, 93). The everted portion in these specimens was membranous, lacking any significant sclerites, and lobes if present were short, therefore it was not studied further. A long sclerotised portion of the centre of the endophallus may be present, but is not or only partially everted and is poorly defined when examined through the walls of cleared penises. This sclerite or sclerotised region is probably an ejaculatory guide, holding the flagellum or apex of the ejaculatory duct in place, similar to the much more clearly defined structure in Cryptocephalinae (Karren 1966; Reid 1991). As it is difficult to examine without damaging the specimen it was not studied for this taxonomic review. An elongate median ventral sclerite is present between the gonocoxites of some Spilopyrinae (fig. 96; Reid 1992: 111). This sclerite lacks setae or clusters of sensilla and therefore may represent secondary thickening of membrane. The spermathecae and ducts show significant differences between species. Spermathecae are drawn as if in cross-section, to illustrate the junction of both gland and duct. The duct in most species is entirely tightly coiled. The exact number and pattern of coils is not diagnostically significant in these species, so for most species only the ends of the duct are illustrated, with a line between indicating the route of that portion of the similarly coiled duct which is omitted. The spermathecal gland is a long spatulate unbranched tube after clearing with KOH, without diagnostic properties and therefore only illustrated for one species (Fig. 120). Label data are given as found on specimens; labels are separated by //. Abbreviations Collections: AMS (Australian Museum, Sydney), ANIC (Australian National Insect Collection, Canberra), BMNH (Natural History Museum, London), MCZ (Museum of Comparative Zoology, Harvard), MVM (Museum of Victoria, Melbourne), QDPIM (Queensland Department of Primary Industry, Mareeba), QMB (Queensland Museum, Brisbane), SAM (South Australian Museum, Adelaide), UQB (University of Queensland, Brisbane). Localities: CQ = Central Queensland, FR = Flora Reserve, NG = New Guinea, NP = National Park, NQ = North Queensland, NNSW = North New South Wales, New South Wales, QLD = Queensland, SF = State Forest, SQ = South Queensland. Allsortsia gen. nov. Type species: Macrogonus maculata Lea 1922, this designation Diagnosis Length 12–13mm; shape elongate-ovoid; dorsum not pubescent; eyes small, gena c. 0.6x eye length; antennae c.0.5x body length; pronotum widest at middle (male laterally produced), and width at posterior angles greater than width at anterior angles; lateral margination of pronotum absent; prosternal process transverse, depressed between coxae, apex bilobed; elytra evenly convex in lateral view, non-striate, without depresssions on disc; male tibiae curved; tibial spurs 1+2+2; claw simple, with slight angulation at base of ventral margin; junction ventrites I and II fused at sides, with some movement at middle. REVISION OF THE AUSTRALO-PAPUAN GENUS MACROLEMA Zootaxa 2486 © 2010 Magnolia Press · 5 Description of Allsortsia Length 12–13mm; body (Figs 1–2) elongate-ovate, without marked constriction at junction of elytra and pronotum, but head remarkably narrow, c. 0.5–0.6x width pronotum; body moderately to shallowly convex in profile, length 2.8x height. Dorsum shining and smooth, multicoloured, with some metallic reflection, glabrous except trichobothria adjacent to eyes and on each corner of pronotum. Head (Figs 1–2, 18): fronto-clypeus smooth, triangular, anterior margin shallowly concave, not depressed between eyes and elevated towards apex; eyes ovate, height c. 1.4x width, inner margin shallowly concave; eye laterally projecting, posterior curvature contiguous with short temples, which are abruptly constricted to parallel-sided head capsule; gena 0.5–0.6x greatest eye length; antennae c. 4.5x socket diameters apart; antennae c. 0.5x body length; antennomere 1 enlarged cylindrical, 2 shortest (c. 0.6x first), 3 equal to 1, antennomeres 6–11 almost parallel-sided, 7–11 longest; antennomeres 1–5 shining and sparsely punctured, 6– 11 dull and densely microsculptured; labrum not densely setose, with 1–2 pairs of median setae and extra setae at lateral margins; mandible with single apical tooth; apical maxillary palpomere elongate fusiform, with narrow apex (both sexes); preapical palpomere triangular, shorter than apical, and of similar width; mentum transverse, width c. 3x median length, but anterior angles narrowly produced. Thorax (Figs 1–2, 18, 41, 44): pronotum transverse, width 1.25x length, broadest near middle and strongly contracted to anterior angles; disc with or without depressions; anterior edge deeply concave, posterior medially bilobed, lateral edges convex or medially lobed; anterior partly and base distinctly margined, lateral margination absent; anterior and posterior angles laterally prominent, each with single trichobothrium; posterior angle at same vertical plane as posterior margin of hypomeron; prosternum flat between coxae and head; prosternal process broad and flat, with bilobed apex; procoxal cavities closed by insertion of hypomeral lobes into prosternal process; scutellum semi-ovate; elytra broadest at base, with strongly developed humeri, without median depressions; without distinct striae; elytral epipleuron narrow, width <0.2x elytral width, entirely visible laterally but upper margin effaced before humerus, slightly sinuate, gradually contracted from base to apex, and without setae; mesoventrite entirely visible, with quadrate median process; wings fully developed, with two large subequal cells and 4 longitudinal veins in the cubito-anal region, anterior to the weakly indicated anal fold; metaventrite transverse, width c. 2x length, without femoral plates; all femora fusiform, with base and apex narrowed, without longitudinal keels or ridges; all tibiae round in section, without keels, thin, with abruptly expanded apices; short spur on protibia present, 2 on remainder; all first tarsomeres with ovate patch of modified setae in males, with dense simple setae diverging from midline in females; apex second tarsomere concave; third tarsomere deeply bilobed, lobes c. 1.5x median length; claws simple, without basal tooth. Abdomen (Figs 60, 66, 67, 81, 96, 98, 128): pygidium (tergite 7) not greatly thickened, basal half membranous, apical half densely pubescent and punctured, without median groove; tergite VIII not fused to spiracles; ventrite I with triangular intercoxal process and without femoral plates; ventrites I and II not completely fused, with some movement at middle; ventrite V smooth, not medially depressed, and truncate or convex at apex in both sexes, but shorter in male; sternite VIII of male Y-shaped; sternite IX of male Y- shaped; male tegmen present, enclosing penis, U- shaped, thinly sclerotised and with slight internal median keel; penis simple, flattened tubular, with basal foramen c. 0.4x length of penis; vas deferens with long thickened sperm pump; female tergite VIII well-developed, sternite VIII with basal apodeme; ovipositor with pair of well-developed paraprocts, enclosed at base by thin baculus, and partly enclosing basal half of palpi, pair of well-defined elongate proctigers dorsal to these; vaginal palpi 3-segmented, apical segment (gonostylus) elongate cylindrical, basal segments (gonocoxite) massive; elongate median ventral sclerite present between gonocoxites; spermatheca hook-shaped; kotpresse present in rectum, as a poorly defined band of elongate spinules. Distribution and biology Allsortsia is confined to Australia, where it occurs in the tropical rainforest of north Queensland. The single species is known from two pre-1920 specimens. The biology is unknown, but is likely to be similar to other Spilopyrinae (Reid 2000), with an external larva feeding on leaf laminae. 6 · Zootaxa 2486 © 2010 Magnolia Press REID & BEATSON Etymology Allsortsia is named from the vague resemblance of the type species to liquorice allsorts, a childhood favourite of the senior author. The gender is feminine. Notes Allsortsia differs from Macrolema by: head structure, antennal length, sexually dimorphic pronotal shape, prosternal process, elytral sculpture, upper margin of epipleura, sexually dimorphic tibiae, almost glabrous ventrites, vaginal palpi, presence of kotpresse. Allsortsia is similar to Spilopyra Baly 1860, from which it differs by: anteriorly elevated frontoclypeus, absent lateral pronotal margins, prosternal process medially depressed and apicolaterally lobed, elytra not transversely depressed, reduced spiculum relictum, 3- segmented vaginal palpi. Allsortsia has less in common with the four remaining spilopyrine genera (Reid 2000). There is a single known species, redescribed below. Allsortsia maculata (Lea 1922), comb. nov. (Figs 1, 18, 41, 44, 60, 66, 67, 81, 96, 98, 128, 130) Macrogonus maculatus Lea 1922: 283 Macrolema maculata: Reid 2000: 862 Material examined Holotype: male/ maculatus Lea type Cairns/ Macrogonus maculatus Lea Queensland I.4773 type/ (SAM). Paratype: female/ Kuranda N Queensland G. E. Bryant F. P. D[odd]. 1909/ type of female Macrogonus maculatus Lea Queensland male has large tooth on each side of prothorax cotype/ cotype/ Australia Brit. Mus. 1921-458/ (BMNH). The female lacks antennomeres 7–11. Description Length: male 12mm, female 13mm; body moderately convex in profile, length c. 3x height. Variegated in metallic blue, red, yellow and brown: (i) anterior of head, head appendages, legs, basal half of middle of pronotum, scutellum, basal third of elytral suture and 6 spots on each elytron (beside scutellum, near suture 1/ 3 from base, on humerus, behind humerus, near side 1/2 from base, near side ¾ from base), black to blackish- brown, with dark blue reflection; (ii) remainder of elytra yellow; (iii) remainder of prothorax, mesoventrite, middle of metaventrite and base of trochanters red to reddish-yellow; (iv) abdomen brown. Head (Figs 1–2, 18): relatively small; strongly and closely punctured throughout anterior half, almost impunctate posteriorly; fronotclypeal margins deeply defined but not depressed between eyes, with deep groove on midline of vertex and irregular groove with trichobothria behind eyes; eyes small, not prominent, flatter in female, separated by c. 4x eye widths (both sexes); gena long, 0.55–0.60x eye length; antennae c. 4.5x socket diameters apart; antennae short, c. 0.5x body length (male), broken in female; all antennomeres elongate: 2 shortest (c. 0.6x first), <1=3, <4=6, <5=8, <7=9=10, <11 (male), female antennomeres 1–6 similar; labrum with 4–5 pairs of prominent lateral setae; apical maxillary palpomere elongate, almost cylindrical in male, more fusiform in female; preapical palpomere shorter than apical. Thorax (Figs 1–2, 18, 41, 44): pronotum sparsely and finely punctured throughout, with larger punctures on basal dark mark, shining, with scattered distinct micropunctures between macropunctures; pronotal shape sexually dimorphic: both sexes pronotal width 1.25x length, but male with strongly developed triangular lateral lobes at middle and deeply concave anterior margin, female evenly convex at sides and shallowly concave anterior margin; pronotal basal margin narrowly excavate at midline; posterior trichobothria anterior to basal margin; pronotal disc convex and without depressions in male, with two small lateral depressions in female; anterior margination absent from middle half; hypomeron impunctate; prosternal process broad and depressed, with two oblique lobes at apex; scutellum punctate, almost semicircular; elytron without REVISION OF THE AUSTRALO-PAPUAN GENUS MACROLEMA Zootaxa 2486 © 2010 Magnolia Press · 7 depressions on basal half of disc; elytral punctures strong and close in basal half, evanescent towards apex; elytra non-striate, with interstrial punctures as large as strial and the latter not in regular lines; 1 deep depression on basal half of elytron adjacent to epipleuron; upper margin epipleuron not reaching base of elytron, obliterated before humerus; mesoventrite median process strongly arched to shallowly concave apex; metaventrite shining and sparsely and minutely punctured, anterior of middle without depression and without margination, edge pitted laterally; metepisternum microreticulate, finely punctured; 1 short spur on protibia, 2 on remainder. Abdomen (Figs 60, 66, 67, 81, 96, 98, 128): basal half pygidium (tergite 7) without pubescence and punctures; ventrites I and II not completely fused, with some movement at middle; ventrites I–V smooth and shining, not microreticulate, sparsely and minutely punctured, on apical half of I–II, and most of III–V, pubescence just visible at x40, except a pair of trichobothria on each ventrite; ventrite I with small lateral keel <0.25x ventrite length, other ventrites without keels; apex ventrite V convex in both sexes; apex of penis triangular in dorsal view, apical angle c.80°, tip slightly curved thickened in lateral view; female sternite VIII with elongate narrow basal apodeme; apex of gonocoxite setose, setae as long as stylus; spermatheca hook- shaped, with loosely coiled duct. Notes Allsortsia maculata is only known from the two type specimens, collected in 1909 at Kuranda in the Wet Tropics bioregion (Fig. 130). The label data of the types suggests that two separate collections were made, but in the description Lea (1922: 284) states that both specimens were collected in the “Cairns district” by F. P. Dodd and G. E. Bryant. We therefore believe that Lea replaced the original labels with his own, as was his practice when writing out new type labels (pers. obs. CAMR). Months of activity are unknown. Allsortsia maculata may reasonably be considered ‘Endangered’ in conservation status (International Union for Conservation of Nature 2001), given its rarity, large size and restricted distribution. Key to the genera of Spilopyrinae This key is modified from Reid (2000), in which elytra of Hornius were wrongly described as pubescent, which is only true for H. grandis (Philippi & Philippi 1864), and erroneous distributions were given for the South American genera (see Jerez 1996). 1 Lateral margins of pronotum with a distinct raised border.......................................................................................2 - Lateral margins of pronotum without raised border .................................................................................................5 2(1) Dorsum brown and non-metallic; pronotum broadest at middle, contracted anteriorly and posteriorly; length 7- 9mm (Argentina and Chile) ....................................................................................................... Hornius Fairmaire - Dorsal surface glabrous, usually partly metallic and black; pronotum shaped otherwise; length 9-15mm..............3 3(2) Dorsal surface wholly metallic, with spots and bands; body broader (length at most 1.8x width) and profile strongly raised in middle (Australia and New Guinea) ..................................................................... Spilopyra Baly - Dorsal surface brown or non-metallic green; body more elongate (length at least 2.5x width) and not arched longi- tudinally ...................................................................................................................................................................4 4(3) Prosternal process narrow and feebly curved longitudinally, broadly expanded and shallowly bilobed at apex; basal half of elytra semistriate; anterior tibia with 2 apical spurs; basal ventrites not fused (Chile).......................... ................................................................................................................................................... Stenomela Erichson - Prosternal process flat, broad, parallel-sided, posteriorly raised and deeply bilobed at apex; elytra non-striate; anterior tibia with one apical spur; ventrites 1 and 2 fused (New Caledonia) .......................Bohumiljania Monrós 5(1) Upper surface extremely dull from microreticulation, clothed with recumbent scales; pronotum with small lateral teeth, or dorsal tubercles; claws bifid (Australia) .......................................................................... Cheiloxena Baly - Upper surface shining, almost glabrous, without scale-like setae; disc of pronotum non-tuberculate, but sides fre- quently with strong angular or rounded projection; claws simple ...........................................................................6 6(5) Elytra with pair of large angular tubercles near scutellum, rugosely and irregularly punctured (Australia).............. ..................................................................................................................................................... Richmondia Jacoby - Elytra smooth, non-tuberculate and non-rugose...................................................................................................... 7 7(6) Pronotal width at posterior angles greater than width at anterior angles; prosternal process depressed, transverse with bilobed apex (Australia)...................................................................................................... Allsortsia gen. nov. 8 · Zootaxa 2486 © 2010 Magnolia Press REID & BEATSON - Pronotal width at posterior angles less than width at anterior angles; prosternal process arched, elongate, with truncate or pointed apex (Australia and New Guinea).................................................................... Macrolema Baly Macrolema Baly 1861 Type species: Macrolema vittata Baly 1861, by original designation and monotypy = Macrogonus Jacoby 1894; Seeno & Wilcox 1982 (synonymy). Type species: Macrogonus submetallicus Jacoby, by monotypy Diagnosis Length 8–15mm; shape elongate, with strongly narrowed base of pronotum compared with elytral width at humeri; dorsum not distinctly pubescent, but may have scattered minute setae; eyes large, gena c. 0.1–0.4x eye length; pronotum widest at middle, or middle and anterior, and width at posterior angles narrower than width at anterior angles; lateral margination of pronotum absent; prosternal process elongate, arched between coxae, apex truncate or slightly pointed; elytra partially striate, but striae may be obscured by large interstrial punctures; male tibiae straight; tibial spurs 1+2+2, or 0+2+2, or 0+1+1; claw simple, with slight angulation at base of ventral margin; ventrites I and II fused or rarely not entirely fused, with some movement at middle. Description Length 8–15mm, females larger than males on average (rarely overlapping); body (Figs 3–16) elongate- ovate, length 2.5–3.0x width, but with prominent elytral humeri and narrow pronotal base, moderately to shallowly convex in profile, length 2.7–3.4x height. Dorsum shining and smooth, coloured metallic green or blue, or plain brown or black, with or without areas of pale yellow or red, glabrous except trichobothria adjacent to eyes and on each corner of pronotum, or with additional short setae on vertex, pronotum and/or elytra. Head (Figs 19–37): fronto-clypeus smooth, triangular, anterior margin shallowly concave, usually depressed between eyes and without elevated apex; vertex smooth; eyes ovate, height c. 1.5x width, inner margin shallowly concave; eye laterally projecting, posterior curvature contiguous with short temples, which are constricted to parallel-sided head capsule; head capsule generally without deep grooves or ridges, but a few species with deep median groove between eyes; gena 0.1–0.4x greatest eye length; antennae 4–6x socket diameters apart; antennae c. 0.6–0.9x body length; relative sizes of antennomeres variable, but 1 always enlarged cylindrical or widest at apex, 2 shortest (c. 0.5–0.67x first), 3 equal to or shorter than 1, antennomeres 8–11 almost parallel-sided, 7, or 7 and 11, or 5, 7 and 11 longest; antennomeres 1–4 shining and sparsely punctured, 6–11 dull and densely microsculptured, 5 intermediate; labrum not densely setose, with 1– 3 pairs of prominent setae; mandible with single apical tooth and two setose membranes (pulvilli) at middle and base of inner margin; apical maxillary palpomere elongate, fusiform to cylindrical, males with broader truncate apex; preapical palpomere triangular, length equal to or shorter than apical, and of similar width; mentum transverse, width c. 4x median length, but anterior angles narrowly produced. Thorax(Figs 3–16, 37–40, 42–43, 45–59): pronotum transverse, width 1.3–1.85x length, broadest anterior to middle (or anterior angles as well) and slightly to strongly contracted to anterior and posterior angles; disc with or without pair of depressions; anterior edge truncate to concave, posterior convex or medially truncate, lateral margins slightly convex to strongly sinuate or lobed; anterior (at least at sides) and base distinctly margined (except some individuals with border absent at hind angles), lateral margination absent; anterior and posterior angles laterally prominent, with single trichobothria; posterior trichobothria anterior to vertical plane above posterior margin of hypomeron, anterior displacement most pronounced in M. albascutica, least so in M. aenescens; posterior edge of hypomeron projecting into hollowed base of elytra; prosternum flat and punctate between coxae and head; prosternal process narrow and strongly arched, with slightly pointed or truncate apex; procoxal cavities closed by insertion of hypomeral lobes into prosternal process; scutellum elongate-triangular, apex superimposed on sutural base; elytra broadest at base, with strongly developed humeri, with or without lateral or median depressions; with scutellary striole and up to 9 striae, plus series of large punctures at epipleural margin, striae often obscured; elytral epipleuron narrow, width <0.2x elytral REVISION OF THE AUSTRALO-PAPUAN GENUS MACROLEMA Zootaxa 2486 © 2010 Magnolia Press · 9 width, entirely visible laterally, slightly sinuate, gradually contracted from base to apex; mesoventrite entirely visible, well-developed, punctured, with elongate and parallel-sided median process; anterior of mesoventrite process evenly curved or elevated with anteriorly facing ridge; wings fully developed; metaventrite shining, smooth or feebly transversely wrinkled, transverse, width c. 2x length, without femoral plates; metepisternum smooth and shining or weakly microreticulate, with sparse punctures; all femora fusiform, with base and apex narrowed, without longitudinal keels or ridges, all tibiae round in section, without keels, thin, with abruptly expanded apices; short spur on protibia present or absent, 1 (M. longicornis) or 2 on remainder; all first tarsomeres with oval patch of modified setae in males, with dense simple setae diverging from midline in females; apex second tarsomere concave; third tarsomere deeply bilobed, lobes at least 0.5x median length; claws simple, without basal tooth. Abdomen (Figs 61–65, 68–80, 82–95, 97, 99–112, 114–127, 129): pygidium (tergite 7) not strongly sclerotised except at apex, basal half membranous, apical half pubescent and punctured, without median groove; ventrite I with triangular intercoxal process and without femoral plates; ventrites I and II usually completely fused; ventrite V smooth, not medially depressed, and truncate or convex at apex in both sexes, but shorter in male; dense sclerotisation of sternite VIII of male variable, fan-shaped, Y-shaped, or linear; sternite IX of male Y-shaped; male tegmen present, enclosing penis, U- shaped, thinly sclerotised and with slight internal median keel; penis simple, flattened tubular, apex sparsely microspiculate, basal foramen c. 0.4x length of penis; vas deferens with long thickened sperm pump; female tergite VIII well-developed, sternite VIII with transverse to elongate basal apodeme; ovipositor with thin bacillus wrapped around base of paraprocts, which are well-developed, partly enclosing basal half of palpi, pair of well-defined elongate proctigers dorsal to these; vaginal palpi 2-segmented, gonocoxite massive, not divided, gonostylus variable in size and shape; membranous pad between gonocoxites lacking sclerite; spermatheca generally falciform, surface microreticulate, with long spermathecal duct; kotpresse absent, rectum without band of spinules. Notes The diagnostic description for Spilopyrinae provided in Reid (2000) needs to be modified to accommodate all species of Macrolema as follows: gena 0.1–1.5x eye length; antennomeres 8–11 to 6–11 densely setose and microsculptured; anterior pronotal border complete or effaced at middle; articulated tibial spurs 0+1+1, or 0+2+2, or 1+2+2, or 2+2+2; third tarsomere shallowly or deeply bilobed; kotpresse (faecal plate organ) present or absent. The synonymy of Macrolema and Macrogonus Jacoby has been briefly dealt with (Reid 2000), but is explained in full here. Macrogonus was defined by possession of lateral lobes on the pronotum (Jacoby 1894). This feature is a grade, at its most extreme in M. submetallicus and M. ventralis, but with varying degrees of expression in between. Differences in other features such as prothoracic morphology, elytral depressions, male and female genitalia, do not conform with development of the pronotal lobes, therefore the synonymy is justified. However the synonymy was originally based on a misunderstanding by Lea, who confused males and females of a species with lateral lobes (M. atripennis) and one without (M. longicornis) (Lea 1921a). Curiously, having decided that Macrogonus and Macrolema could not be distinguished except in males, Lea described M. ventralis in Macrogonus on the mistaken assumption that the holotype was a female specimen (Lea 1921b). This action has led to considerable confusion in Australian collections, which have been dominated by material identified by Lea. Sexual dimorphism Sexual dimorphism is slight. Males are smaller on average, with larger cephalic sensory organs: apical maxillary palpomeres broader, antennae longer, eyes proportionally larger. Males also have first tarsomeres broader and ventrite 5 shorter. Distribution and biology Macrolema is confined to Australia and New Guinea; it occurs on the eastern ranges of Australia, from Barrington Tops, New South Wales, in the south, to Cape York Peninsula, Queensland, and the eastern ranges 10 · Zootaxa 2486 © 2010 Magnolia Press REID & BEATSON

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