Zootaxa 4446 (2): 151–202 ISSN 1175-5326 (print edition) Article ZOOTAXA http://www.mapress.com/j/zt/ Copyright © 2018 Magnolia Press ISSN 1175-5334 (online edition) https://doi.org/10.11646/zootaxa.4446.2.1 http://zoobank.org/urn:lsid:zoobank.org:pub:4B7A9974-CD3A-4BF5-9062-E48D73F3CADF Revision of the genus Lamprima Latreille, 1804 (Coleoptera: Lucanidae) CHRIS A.M. REID1, KINDI SMITH2 & MAX BEATSON1 1Entomology, Australian Museum, 1 William Street, Sydney, NSW 2010, Australia. E-mail: [email protected] 224 Milyerra Road, Kariong, NSW 2250, Australia. Table of contents Abstract . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .151 Introduction. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 152 Taxonomic history of Lamprima . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .152 Methods and abbreviations . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 153 Lamprima Latreille, 1804 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 154 Key to species of Lamprima Latreille, 1804. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 160 Lamprima adolphinae (Gestro 1875) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .161 Lamprima aenea (Fabricius, 1792). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 163 Lamprima aurata Latreille, 1817 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 171 Lamprima imberbis Carter, 1926 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 195 Lamprima insularis W.J. Macleay, 1885. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 197 Acknowledgements. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 199 References. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 199 Abstract The genus Lamprima Latreille, 1804 (Coleoptera: Lucanidae: Lampriminae: Lamprimini), is revised. Five species are rec- ognised: one in New Guinea (L. adolphinae (Gestro, 1875)), two on isolated western Pacific islands (L. aenea Fabricius, 1792: Norfolk Island; L. insularis W.J. Macleay, 1885: Lord Howe Island), one in northeastern New South Wales (L. im- berbis Carter, 1926) and a common widespread species in eastern and southern Australia, L. aurata Latreille, 1817. Lam- prima aurata varies considerably morphologically and many of the different forms encompassed by this variation have been described as species. Our study of morphology does not support this classification. Therefore, Lamprima aurata is designated a senior synonym of the following 24 names: L. cuprea Latreille, 1817; L. latreillii W.S. MacLeay, 1819 (new synonym); L. pygmaea W.S. MacLeay, 1819 (new synonym); L. fulgida Boisduval, 1835; L. micardi Reiche, 1841 (new synonym); L. rutilans Erichson, 1842; L. splendens Erichson, 1842; L. viridis Erichson, 1842; L. nigricollis Hope in West- wood, 1845 (new synonym); L. purpurascens Hope in Westwood, 1845 (new synonym); L. sumptuosa Hope in West- wood, 1845 (new synonym); L. tasmaniae Hope in Westwood, 1845 (new synonym); L. varians Burmeister, 1847 (new synonym); L. cultridens Burmeister, 1847 (new synonym); L. amplicollis Thomson, 1862 (new synonym); L. krefftii W.J. MacLeay, 1871 (new synonym); L. violacea W.J. Macleay, 1885 (new synonym); L. mandibularis W.J. Macleay, 1885 (new synonym); L. sericea W..J Macleay, 1885 (new synonym); L.nigripennis W.J. Macleay, 1885 (new synonym); L. minima W.J. Macleay, 1885 (new synonym); L. mariae Lea, 1910; L. coerulea Boileau, 1913 (new synonym); L. insularis Boileau, 1913 (new synonym). Lamprima adolphinae is a senior synonym of L. bohni (Darge & Séguy, 1953) (new syn- onym). Lamprima schreibersi Hope in Westwood, 1845, is an unnecessary nomen novum for L. aenea redescribed by Schreibers in 1802 from the same material as Fabricius, and therefore an objective synonym of L. aenea. Lamprima punc- ticollis Dejean, 1833, L. coerulea Hope in Westwood, 1845, and L. insularis Hope in Westwood, 1845, are nomina nuda, the last two names first made available by Boileau in 1913. The five Lamprima species are redescribed and recommenda- tions made for their conservation. Type specimens of the species of Lamprima described by William Sharpe MacLeay and William John Macleay are illustrated for the first time. Lectotypes are designated for Lamprima insularis, L. latreillii, L. latreillii sericea, and L. mandibularis. Key words: stag beetle, synonymy, morphology, nomenclature, polymorphism, insect trade, island endemism, distribu- tion Accepted by A.B.T. Smith: 30 Apr. 2018; published: 17 Jul. 2018 151 Licensed under a Creative Commons Attribution License http://creativecommons.org/licenses/by/3.0 Introduction The Australopapuan genus Lamprima Latreille, 1804 (Coleoptera: Lucanidae: Lampriminae: Lamprimini), includes some of the most spectacularly coloured species of stag beetles worldwide, in a family of predominantly dowdy brown or black species. For this reason Lamprima is also a popular genus with collectors and the international insect trade. Despite, or perhaps because of, its popularity, the species are not clearly defined. Thirty- six species rank names have been used in Lamprima, of which 34 are available (Table 1). Most of these have been synonymised under three names. However, even these three species concepts are poorly defined and therefore considerable confusion still exists in determining the species of Lamprima. The purpose of our paper is to resolve the poor taxonomy of Lamprima. Taxonomic history of Lamprima The complicated taxonomy of Lamprima species is summarised in Table 1. The first species of Lamprima to be described remains one of the least known. Lethrus aeneus Fabricius, 1792, was described from remote Norfolk Island in the western Pacific shortly after the island was discovered by Europeans. The genus Lamprima was created for L. aenea 12 years later (Latreille 1804a) when this was still the only known species. However, Latreille (1804b: 239) noted that the Natural History Museum (Paris) had "plusieurs de ces insectes" from the naturalists accompanying Baudin to Australia. As Baudin only visited the south and west coasts of Australia, it is likely that Latreille's generic description is based on a combination of species. 13 years after that Latreille described two species, L. aurata Latreille, 1817, and L. cuprea Latreille, 1817, from Australian mainland material. Not long afterwards, William Sharpe [‘W.S.’] MacLeay studied the classification of Lucanidae, describing two further species in the process, L. latreillii (commonly misspelt 'latreillei') W.S. MacLeay, 1819, and L. pygmaea W.S. MacLeay, 1819. Boisduval (1835) placed these species in synonymy, and also described a new species. By the 1840s, a considerable area of Australia had been visited by collectors, producing plenty of specimens for European collections. Twelve new species were described during this decade (Reiche 1841; Erichson 1842; Hope in Westwood 1845; Burmeister 1847) and northern Queensland was the only forested area not explored entomologically by this time. During the next 30 years, Thomson (1862) and William John [‘W.J.’] MacLeay (1871) added two Australian species and the single New Guinea species was discovered and described, but in its own monotypic genus, Neolamprima Gestro 1875. Hope in Westwood (1845), Parry (1864, 1870) and Harold (1868) catalogued the species of Lamprima, and each provided different combinations of synonyms. In 1884 there were 23 Australian species names in Lamprima, but without a key for their identification. W.J. Macleay (1885a) reviewed the Australian species to make sense of the taxonomy, but in the process described a further six new species and one new subspecies, recognising 15 species. One of the new species was described in Neolamprima. W.J. Macleay’s taxonomic work was partly a result of new collections from northern Queensland and Lord Howe Island. Macleay (1885c) later created the genus Phalacrognathus for Lamprima muelleri (W.J. Macleay, 1885). Arthur Lea added a subspecies from the Bass Strait islands (Lea 1910). Lea also noted that the species described by Macleay in Neolamprima, was variable, with some specimens showing strongly asymmetric mandibles combining the characteristics of Lamprima and Neolamprima (Lea 1910). Nagel (1922) synonymised these genera and Carter (1926) described one further Australian species. Nagel examined some of W.J. Macleay’s type material and identified new synonyms, reducing the number of Lamprima species to eight (Nagel 1930). From then on, no further taxonomic novelties have been described from Australia but four colour forms of the New Guinea species (L. adolphinae) have been named (Nagel 1930; Kriesche 1940; Didier & Séguy 1952; Darge & Séguy 1953), three of which were synonymised with L. adolphinae by Krajcik (2001). The male genitalia of L. aurata have been described as part of wider systematic studies of Lucanidae (Holloway 1960; d’Hotman & Scholtz 1990). Lamprima aurata has been introduced to New Zealand, and has been described in detail as part of that fauna (Holloway 2007). Benesh (1960) and Moore & Cassis (1992) closely followed Nagel in their catalogues of the world and Australian lucanids respectively, but they also listed the older synonymic names ignored by Nagel. Moore & Cassis ignored the implicit or suggested synonymy of three species in popular guidebooks by Matthews (1984) and Moore (1986). Mizunuma & Nagai (1994) suggested that L. aurata and L. latreillii might be synonymous, but did not 152 · Zootaxa 4446 (2) © 2018 Magnolia Press REID ET AL. formally make them so. The most recent catalogue of Lamprima names (Krajcik 2001) is similar to that of Benesh (1960). Popular works since then have maintained their separation (Fujita 2010; Hangay & de Keyzer 2017; Bartolozzi et al. 2017). The conflict between popular literature (now largely pursued online) and outdated taxonomic work has contributed to the continuing confusion over which Lamprima species are valid and their correct names. Approximately 1200 specimens were examined by the authors for this revision. Label data from an additional 87 specimens in the Canadian Museum for Nature are also included, through the research associate Andrew Smith (18 March 2018). We have measured and dissected males and females of Lamprima from many localities, including many size and colour forms, and reached the conclusion that there are only two mainland species of Lamprima, one of which is known from a single specimen from New South Wales. The three species described from Norfolk Island, Lord Howe Island and New Guinea remain valid. All five species are redescribed below. Methods and abbreviations Morphological nomenclature follows Lawrence & Ślipiński (2013). All measurements were made with an eyepiece graticule. Figures 1–74 and Fig. 77 were imaged using either: Visionary Digital lenses and P-51 camlift system with a Canon EO5 7D body then processed with Helicon Focus 5.0 and Adobe Photoshop; or, a Leica MZ16 A, then processed with the Leica Application Suite V3. Figures 78–85 were taken by Cate Lemann, Australian National Insect Collection, using a Dun Inc. BK Imaging-Plus Lab System; source images were then aligned and stacked in Zerene Stacker v.1.0 and processed in Adobe Photoshop CS6 to obtain a fully-sharpened image. The placement of host plant genera in named families is derived from Stevens (2017). Authors for host plant genera and species can be found in the Australian Plant Name Index (Anonymous 2018). The following collections are referred to: Australian Museum, Sydney (AMS); Australian National Insect Collection, Canberra (ANIC); Canadian Museum of Nature, Canada (CMNC); Hope Collection, Oxford University (OXUM); National Agricultural Insect Collection, Port Moresby (NAIC); Natural History Museum, London (BMNH); Queensland Museum, Brisbane (QMB); Queen Victoria Museum & Art Gallery, Launceston (QVMAG); South Australian Museum, Adelaide (SAM); Western Australian Museum, Perth (WAM). In an earlier lucanid revision it was noted that William Sharp MacLeay spelt his surname with a capital 'L' and that his nephew William John Macleay spelt his name with a lower case 'l', in later works (Reid & Beatson 2016). In that review it was unecessary to use initials for each author as they were distinguished by the orthography of their surnames. However, in Lamprima, W.J. Macleay used a capital 'L' in his first paper dealing with this genus (MacLeay 1871). We have therefore indicated the authors by adding their initials. There has been confusion over the authorship of the anonymous catalogue of the Hope collection of Lucanidae, usually cited as Hope & Westwood (1845). In the obituary of Hope, the writer notes "he also published catalogues of his collections of Lucanidae" (Anonymous 1862: 786), which suggests that Hope was the author of the catalogue, although the word published equally means 'enabled to be published'. Two contemporaries or near contemporaries, Parry (1864) and Macleay (1885a), both cite only Hope. Parry, who was a friend of Westwood, simply mentions the catalogue under Hope's name, as "Hope, Cat." (Parry 1864:7). Macleay is more specific: "described by the Rev. F. W. Hope in the year 1845, in a list of Lucanidae published by him in London, in pamphlet form" (Macleay 1885a: 135). However Boilieu, who visited OXUM and revised all of Hope and Westwood's type material, noted that several copies of the catalogue existed with 'by J. O. Westwood" handwritten by Westwood on the title page (Boilieu 1913: 213). There is no reason to doubt the sincerity of this attribution, as Westwood worked closely with Hope and owed his position at Oxford University to Hope. Therefore this definitively identifies Westwood as the author of the catalogue. Arrow's casual remark "Westwood, in his anonymously published 'Catalogue of the Lucanoid Coleoptera' (generally attributed to Hope)" (Arrow 1936) may have been due to Boilieu or to his own observations of the annotated catalogues. However, since Arrow's comment, the catalogue has generally been cited as "Hope & Westwood" (Benesh 1960; Moore & Cassis 1992; Mizunuma & Nagai 2004; Holloway 2007; Hangay & de Keyzer 2017), or Hope only (Bouchard et al. 2011). Paulsen (2005) corrected this citation and we follow him in citing Hope's species as Hope in Westwood, 1845. THE STAG BEETLE GENUS LAMPRIMA Zootaxa 4446 (2) © 2018 Magnolia Press · 153 Lamprima Latreille, 1804 Lamprima Latreille, 1804a: 150. Type species. Lethrus aeneus Fabricius, 1792, by monotypy. Neolamprima Gestro, 1875: 997; Nagel 1922: 16 (synonymy). Type species. Neolamprima adolphinae Gestro, 1875, by monotypy. Description. Length. Male 13–60 mm (usually 20–45 mm) including mandibles, mandibles 10–38% of overall length (Figs 1–14); female 13–27 mm including mandibles, mandibles 5–9% of overall length (Figs 15–20). Dorsal surface without scales or visible setae; venter setose, setae simple. Head. Male without dorsal tubercles on head, anterior of head concave or truncate (Figs 21–32); genae well- developed anterior to eyes but not laterally expanded, at most slightly anteriorly angulate in males; female head not narrowed compared with male; temples short but slightly angulate, shallowly grooved or notched to accommodate anterior angles of pronotum; head deeply inserted into pronotum which almost reaches eyes (Figs 33–44); eyes undivided, reniform, with shallowly concave anterior and posterior margins, the anteroventral margin sharp and linear, forming the outer edge of the antennal groove; antennae not geniculate, with 10 antennomeres, with 3 antennomere club, the club antennomeres entirely densely setose and often closely appressed; antennomere 7 cupuliform, with thin lateral lobe and 6 slightly asymmetric (male) or with thin lobe (female); male mandibles as long as or longer than head and usually densely internally setose in male (not L. imberbis); male mandible without basal dorsal tooth; each female mandible with only one dorsal cusp, with large strongly incurved basal ventral tooth, overlapping at tips; penultimate labial palpomere angulate on inner margin in male, not angulate in female; labium broad, approximately one third width of head; mentum solid and punctate, apex truncate; pregula flat; lateroventral grooves present between eyes and sides of buccal cavity for retention of scape. Thorax. Pronotum (Figs 1–44) convex with angulate (most males) to rounded sides, broadest slightly posterior to middle, without prominent anterior or posterior angles; anterior of pronotum broadly margined; middle of prosternal process concealed between procoxae, apex flat. Scutellum almost equilateral triangular. Elytra non- striate but may have irregular, shallow grooves, with scattered shallow simple concave punctures, surface of interspaces smooth and shiny to dull and wrinkled but always with finely microreticulate microsculpture, usually slightly granulose in males (Figs 54–57). Elytral humeri not spined. Elytral epipleurae hidden from above, transversely strigose. Hindwings fully developed. Anterior field of mesoscutum strongly and closely punctate; scutellum semicircular to heart shaped; mesanepisternum with sparse, large punctures; posterior half mesepimeron, visible parts of metanepisternum, metepimeron and sides of metaventrite with dense, small punctures (partly coalescent, interspaces less than diameters) and setae; mesoventral process almost parallel-sided to junction with metaventrite, abruptly elevated anterior to this (Figs 58–61); metanepisternum with elevated lobe at anterior angle locking into small depression on elytral epipleuron. Profemur without anterior ridge; male protibia (Figs 62–75) with expanded blade- or fan-like flat spur, except L. imberbis with narrowly elongate, triangular, curved spur; female protibial spur narrowly elongate triangular; male protibia without secondary teeth between major teeth; female protibia without subsidiary teeth between large teeth on outer edge; male metatibia without setose excavation on inner edge, usually with spines on outer edge; tarsal empodium prominent, with paired divergent thin tufts of apical setae. Abdomen. Ventrites not laterally ridged and without basal transverse grooves. Male genitalia (Figs 76–88; unique male of L. imberbis undissected). Phallobase fusiform or spindle-shaped and uniformly sclerotised, with the apical margins rounded laterally and with V-shaped excavations dorsally (shallower) and ventrally (deeper); apical half of dorsal surface with at least a few small raised spicules, which may form irregular oblique ridges. Parameres about 2/5 length phallobase, symmetrical, with preapical setal tuft on ventral surface and incurved pointed tip; ventral inner edge of parameres soft, irregularly ridged, with the surface either inflated towards the penis or collapsed laterally, depending on preservation of the specimen; penis symmetrical, in two visible parts: basal 2/3 darkly sclerotised, rigid and narrowly conical between parameres, obliquely ridged at base; apical third thinly sclerotised and usually translucent, as a thin straight cylinder; endophallus, if everted, of similar width to apex of penis but soft and flexible, apically contracting to long, thin flagellum. Female genitalia (Figs 89–94; L. imberbis unknown). Tergite IX with acute to broadly rounded translucent apex; pleurite IX and sternite IX well developed as elongate sclerites; hemisternites flat and apical halves elongate 154 · Zootaxa 4446 (2) © 2018 Magnolia Press REID ET AL. rectangular with truncate apices, gonostyli flat, inverted-trapezoidal in shape, inserted on middle of membranous apex of hemisternites; spermathecal duct coiled and convoluted, spermatheca sclerotised, variable in shape from tear-drop to hook; spermathecal gland smaller than spermatheca, glandular duct longer than spermatheca. Larval diagnosis. The following diagnosis is based on published descriptions of L. aurata larvae (Alderson 1975 [as L. varians]; Lawrence 1981). The larva is similar to the lamprimine Phalacrognathus W.J. Macleay, 1885 (Wood et al. 1996). Head: antenna with 3 antennomeres; second antennomere without setae or sensory spots, apex slightly produced beyond base of third antennomere, the latter elongate but much smaller than second. Left mandible without incisor teeth between 2–3 apical teeth and mola; epipharynx trapezoidal, with dense, long setae on apical margin and dense, short setae on sides, apex of median area with transverse row of 6 short, blunt pegs and row of 4 circular sensilla proximal to this, epitorma absent. Legs without distal claw, last 3 segments short and broad and densely setose; tibiotarsus reduced to an ovate lobe, length equal to width at base; mesocoxal stridulatory file (pars stridens) with a single line of about 50 quadrate to slightly transverse dense ridges, granulose towards base, and placed in a field of minute granules; metatrochanteral stridulatory file (plectrum) a single line of 45–60 closely placed transverse ridges or granules, increasing in size from base to apex. Anal area of abdomen with two adjacent pear-shaped and minutely but densely setose lobes, without ovate pads, their bases subtending about 90° in ventral view, separated from 10th segment by a dorsal lobe, which is triangular, glabrous and unsculptured; 10th abdominal segment not foreshortened dorsally, ventral apex triangularly excavate, raster confined to apical quarter, consisting of dense, minute, inwardly-directed setae. Ecology and natural history. The following notes mostly concern the common and widespread species L. aurata (Fearn 1996, 2015, 2016) and the Lord Howe endemic L. insularis (Reid 2004). Lamprima adults are commonly diurnal and are often found on flowers, where they may mate. Adult males snip off the apical shoots of living plant material to provide sap flows. They have been recorded feeding on many genera, both native and exotic, listed by family as follows: Asparagaceae: Lomandra; Asteraceae: Ozothamnus; Casuarinaceae: Casuarina; Fabaceae: Acacia, Virgilia; Malvaceae: Lavatera; Myrtaceae: Eucalyptus, Leptospermum, Melaleuca; Pinaceae: Pinus; Proteaceae: Banksia; Rhamnaceae: Alphitonia; Rosaceae: Photinia, Prunus; Salicaceae: Populus (Fearn 1996, 2015; Hangay & de Keyzer 2017). Feeding by the males of Lamprima with elongated mandibles (most L. adolphinae and some L. aurata) has not been described. Female mandibles are apparently non-functional, therefore females fly to the male feeding sites to lap up sap released by males. Feeding sites are used for male-male aggression and for copulation. Fearn (1996) noticed a 3:1 male to female sex ratio in the field, which is corroborated by the male biased material of most species in collections. Olliff (1889) noted that male L. insularis were much more common than females. In contrast, the material available for L. insularis has a roughly 1:1 sex ratio, but were mostly collected by breaking open logs, so does not represent typical field activity of the species. Adults will also feed on soft fruits (Hangay & de Keyzer 2017). Oviposition is usually underground, the females tunnelling into soil around partly buried decaying wood. There is a great variety of larval host genera, including exotics: Araucariaceae: Araucaria; Arecaceae: Howea; Casuarinaceae: Allocasuarina, Casuarina; Celastraceae: Elaeodendron; Fabaceae: Acacia; Lauraceae: Cryptocarya; Myrtaceae: Eucalyptus, Syzygium; Oleaceae: Olea; Salicaceae: Salix(Fearn 1996; Reid 2004; Hangay & de Keyzer 2017). Lamprima aurata larvae are usually in decaying roots and buried timber in Tasmania (Fearn 1996) but they prefer standing timber in northern Queensland rainforest (Wood et al. 1996). Lamprima insularis larvae usually inhabit fallen timber on or above ground level in the subtropical rainforests of Lord Howe Island (Reid 2004). In wetter areas Lamprima larvae may be better able to survive above ground, or less able to survive below ground, but there may be a trade-off between humidity and temperature, as L. aurata avoids cool temperate rainforests in Tasmania (Fearn 1996). In logs and stumps the larvae generally bore upwards. Pupation is in a chamber, usually just beneath the wood surface but sometimes in adjacent soil (Fearn 1996). In Tasmania the entire life cycle is at least three years but it may be 1–2 years in Queensland (Hangay & de Keyzer 2017). In captivity, the life cycle of L. adolphinae is 9–14 months (Levet 2016). There are numerous photographs and several videos of Lamprima species on the Internet (for example: Anonymous 2017a), showing: different colour varieties, mating, fighting between males, feeding, rearing methods, larvae and pupae. In copulation and precopulation the protibial spurs of the males have little function. They may scrape lightly over the pronotum of the female as the prothoracic legs are moved backwards and forwards, but this activity seems erratic and brief. In male-to-male combat, each male uses its mandibles to try to embrace the mandibles of the other, so longer mandibles provide a wider net for the embrace. Once one male has enclosed and THE STAG BEETLE GENUS LAMPRIMA Zootaxa 4446 (2) © 2018 Magnolia Press · 155 squeezed together the mandibles of its rival, it shakes the whole animal quickly to one side to unbalance it, then abruptly to the other side, letting go at the end of this second swing. The rival can be flung a few centimetres (see video by Kan 2016). The elongate mandibles of L. adolphinae allow males to grab wayward appendages of rivals rather than gripping the whole head. In fights, the protibial spurs may be used as braces against the substrate and this activity might be their primary function. The international pet trade is heavily involved in rearing Lamprima species, with goals including production of enlarged mandibles and unusual colour varieties. This may extend the range of variation for each species given here, which is based on field-collected specimens. Lamprima is widespread in Australia (Fig. 95), occupying almost the entire eastern edge of the continent from Cooktown in northern Queensland to Tasmania and most of the south coast from Mallacoota west to Perth, with an 850 km gap at the Nullarbor Plain. Lamprima occurs up to 400 km inland on the mainland. Lamprima also occurs on two oceanic islands, Norfolk and Lord Howe and a single species is widespread on mainland New Guinea (Fig. 95). In New Guinea it occurs up to 2800 m. Conservation. Conservation status and threats are discussed under each species. In general Lamprima species are extremely popular with stag beetle collectors and most species are being, or have been, reared in commercial quantities in eastern Asia and probably Europe and North America. One species, L. imberbis, is of considerable concern as it has not been collected for 100 years. Comparison with other genera of Lampriminae. There are four other genera of Lampriminae, all monotypic. The New Zealand genus Dendroblax White, 1846, is unmistakably different from Lamprima, with a dynastine-shaped body, densely punctate, non-metallic, reddish-brown upper surface, venter with long setae and minimal sexual dimorphism (Holloway 2007). The Australian endemic Homolamprima W.J. Macleay, 1885, is relatively easily distinguished from Lamprima by: mesometaventrite junction anteriorly bilobed; apex prosternal process elevated; male protibia with large narrow spines. Homolamprima is also much flatter than any Lamprima species (Macleay 1885b). The South American genus Streptocerus Fairmaire, 1850, is similar to Homolamprima but differs from it and all other Lampriminae by the antennal club having four antennomeres (Paulsen 2010). Lamprima is morphologically most similar to the northern Australian endemic Phalacrognathus Macleay, 1885, although this is not supported by an analysis of four gene regions (Kim & Farrell 2015). Most species of Lamprima can easily be distinguished from Phalacrognathus by male protibia with spur expanded as a flat blade and female protibia without subsidiary teeth between large teeth on outer margin. In Phalacrognathus, the male protibia has a simple spur (as in female) and the female protibia has small subsidiary teeth present between the large teeth on the outer margin. However, Lamprima imberbis, with unknown female, is unusual in Lamprima for its male mandibles lacking internal setae, male protibiae with narrow spurs and without a setal tuft and elytra broadly explanate. It shares these characters with Phalacrognathus, but differs from that genus by: genae prominent anterior to eyes; eyes anteriorly concave; antennomere 7 with flat lateral lobe; temples prominent and notched to accommodate anterior angles of pronotum; male mandible without basal dorsal tooth; anterior of pronotum broadly margined; middle of prosternal process concealed by procoxae; sides of male elytra not crenulate; male protibia without secondary teeth between major teeth; protibial spur on a lobe. All of these characters are common to other male Lamprima and justify placement of L. imberbis in Lamprima. Included species. The most recent peer-reviewed checklist of Lamprima species (Moore & Cassis 1992) lists seven in Australia (Table 1), of which Lamprima insularis is unique to Lord Howe Island and Lamprima aenea is unique to Norfolk Island. One non-Australian species of Lamprima is known, from New Guinea: L. adolphinae, as catalogued by Benesh (1960). The mainland Australian species listed by Moore & Cassis (1992) are L. aurata, L. imberbis, L. latreillii, L. micardi and L. varians. Lamprima micardi is supposedly endemic to Western Australia and L. varians was described from South Australia. The other species, L. aurata, L. imberbis and L. latreillii, were described from the eastern coastal region of Australia, from northern Queensland to Tasmania. 156 · Zootaxa 4446 (2) © 2018 Magnolia Press REID ET AL. e (heads of columns). Abbreviations in type = northern Queensland; NSW = New South Krajcik This work Moore & 2001 2017 Cassis 1992 - L. L. adolphinae adolphinae L. aenea L. aenea L. aenea L. aurata - L. aenea & L. aurata L. aurata L. aurata L. aenea & L. aurata - L. latreillii L. aurata L. latreillii L. latreillii L. aurata L. aurata L. aurata L. aurata - - L. adolphinae - L. L. adolphinae adolphinae nomen L. aurata nomen nudum? nudum - L. latreillii L. aurata L. varians L. micardi L. aurata L. aurata L. aurata L. aurata L. aurata L. aurata L. aurata - L. latreillii L. aurata ……continued on the next page d valid by each authority and datolland; NI = Norfolk Island; NQ Nagel 1930 Benesh 1960 L. L. adolphinae adolphinae L. aenea L. aenea - - - L. aurata - L. latreillii - L. latreillii L. aurata L. aurata - L. adolphinae - L. adolphinae - L. aurata - - - L. varians - L. aurata - L. aurata - L. latreillii was considereNH = New H Boileau 1913 - L. aenea - - - - L. aurata - - MS name L. latreillii L. latreillii - - L. aurata - me ea; TABLE 1Lamprima. History of all catalogued species rank names in Latreille, 1804. Bold indicates that the nalocality column: CQ = central Queensland; LHI = Lord Howe Island; NA = northern Australia; NG = New GuinWales; SA = South Australia; SQ = southern Queensland; Tas = Tasmania; WA = Western Australia. Harold Macleay Parry Author Date Type Hope in Lamprima 1868 1885a 1864, locality Westwood species rank 1870 1845 name L. adolphinae Gestro 1875 NG - - - L. adolphinae L. aenea Fabricius 1792 NI L. aenea L. aenea L. aenea L. aenea L. aenea Fabricius 1802 NI L. L. aenea L. aenea - sensu schreibersi Schreibers 1805 NI & - L. aurata - L. aurata L. aenea Fabricius NH sensu Donovan L. aenea Boisduval 1835 NH - L. L. latreillii L. latreillii latreillii L. amplicollis Thomson 1862 SQ - L. L. latreillii L. latreillii latreillii L. aurata Latreille 1817 NH L. aenea L. aurata L. aurata L. aurata L. bohni Darge & 1953 NG - - - - Séguy L. chalcitidis Didier & 1952 NG - - - - Séguy L. coerulea Donovan 1805 NH L. latreillii L. L. latreillii L. latreillii latreillii L. coerulea Boileau 1913 not - - - - given L. cultridens Burmeister 1847 NH - L. L. varians L. micardi cultridens L. cuprea Latreille 1817 NH L. aenea L. L. aenea L. aurata latreillii? L. fulgida Boisduval 1835 NG[!] - L. L. aurata L. aurata aurata? L. fulgida Thomson 1862 NH - L. - - splendens THE STAG BEETLE GENUS LAMPRIMA Zootaxa 4446 (2) © 2018 Magnolia Press · 157 This work 2017 L. imberbis nomen nudum L. insularis L. aurata L. aurata L. aurata L. adolphinae L. aurata L. aurata L. aurata L. aurata [not Lamprima] L. aurata L. aurata L. adolphinae nomen nudum L. aurata he next page Krajcik 2001 L. imberbis L. micardi L. insularis - L. latreillii L. latreillii L. adolphinae L. latreillii L. aurata L. micardi L. micardi [not Lamprima] L. micardi - L. adolphinae L. aurata L. micardi continued on t … Moore & Cassis 1992 L. imberbis - L. insularis - L. latreillii L. latreillii - L. latreillii L. aurata L. micardi L. varians [not Lamprima] L. micardi - - L. aurata L. micardi … Benesh 1960 L. imberbis L. micardi L. insularis - L. latreillii L. latreillii adolphinae L. latreillii L. aurata L. micardi L. varians [not Lamprima] L. micardi L. latreillii L. adolphinae L. aurata L. micardi Nagel 1930 L. imberbis - L. insularis - L. latreillii? L. latreillii - L. latreillii L. aurata L. micardi L. varians [not Lamprima] - L. latreillii L. adolphinae - - Boileau 1913 - L. micardi - L. micardi - L. latreillii - - - L. micardi - [not Lamprima] L. micardi - - L. aurata L. micardi Macleay 1885a - - L. insularis - L. krefftii L. latreillii - L. mandibularis - L. micardi L. minima L. muelleri L. micardi L. nigripennis - - L. micardi Harold 1868 - nomen nudum - - - L. latreillii - - - L. micardi - - L. nigricollis - - L. latreillii L. micardi Parry 1864, 1870 - - - - - L. latreillii - - - L. micardi - - L. micardi - - L. latreillii - Hope in Westwood 1845 - L. micardi - - - L. latreillii - - - L. micardi - - L. nigricollis - - - L. micardi y Type localit NSW WA LHI not given CQ NSW NG NQ Tas WA SA NA WA NH NG NH WA e 6 5 5a 3 1 9 0 5a 0 1 5a 5a 5 5a 0 3 5 Dat 192 184 188 191 187 181 194 188 191 184 188 188 184 188 193 183 184 nued) Author Carter Hope W.J. Macleay Boileau W.J. MacLeay W.S. Macleay Kriesche W.J. Macleay Lea Reiche W.J. Macleay W.J. Macleay Hope in Westwood W.J. Macleay Nagel Dejean Hope in Westwood nti TABLE 1. (Co Lamprima species rank name L. imberbis L. insularis L. insularis L. insularis L. krefftii L. latreillii L. lulua L. mandibularis L. mariae L. micardi L. minima L. muelleri L. nigricollis L. nigripennis L. olivacea L. puncticollis L. purpurascens 158 · Zootaxa 4446 (2) © 2018 Magnolia Press REID ET AL. This work 2017 L. aurata L. aurata L. aenea L. aurata L. aurata L. aenea L. aurata L. aurata L. aurata L. aurata L. aurata Krajcik 2001 L. latreillii L. aurata L. aurata L. latreillei L. latreillii L. aenea L. micardi L. latreillii L. varians L. latreillii L. aenea Moore & Cassis 1992 L. latreillii L. aurata L. aurata L. latreillii L. latreillii L. aenea L. micardi L. aurata L. varians L. latreillii L. aenea Benesh 1960 L. latreillii L. aurata L. aurata L. latreillii L. latreillii L. aenea L. micardi L. latreillii L. varians L. latreillii L. aurata Nagel 1930 - L. aurata - L. latreillii - - - L. varians L. latreillii L. aurata Boileau 1913 - - L. aurata - - L. aenea L. micardi L. latreillii - - - s Macleay 1885a L. latreillii L. rutilans - L. latreillii L. splenden L. aenea L. micardi L. latreillii L. varians L. violacea ? Harold 1868 L. latreillii L. rutilans L. aurata - L. splendens L. aenea L. sumptuosa L. latreillii L. micardi - L. aenea Parry 1864, 1870 L. latreillii L. aurata L. aurata - L. aurata L. aenea L. micardi L. latreillii L. micardi or L. varians - L. aenea Hope in Westwood 1845 - - L. schreibersi - - L. subrugosa L. sumptuosa L. tasmaniae - - - y Type localit AUS Tas NH NQ not given not given WA Tas SA NSW not given e 9 2 5 5a 2 5 5 5 8 5a 2 Dat 181 184 184 188 184 184 184 184 184 188 184 nued) Author W.S. MacLeay Erichson Hope in Westwood W.J. Macleay Erichson Hope in Westwood Hope in Westwood Hope in Westwood Germar W.J. Macleay Erichson nti TABLE 1. (Co Lamprima species rank name L. pygmaea L. rutilans L. schreibersi nom nov for L. aenea Schreibers L. sericea L. splendens L. subrugosa L. sumptuosa L. tasmaniae L. varians L. violacea L. viridis THE STAG BEETLE GENUS LAMPRIMA Zootaxa 4446 (2) © 2018 Magnolia Press · 159 There are good male characters for diagnosing L. aenea, L. adolphinae, L. imberbis and L. insularis. There remain the Lamprima species, excluding L. imberbis, described from mainland Australia and Tasmania. It has already been noted that Matthews (1984) treated L. varians and L. aurata as one species in South Australia (L. aurata) and Moore (1984, 1986) suggested that L. aurata and L. latreillii were variants (“overlapping subspecies”) of one species in southeastern Australia, even though these were originally separated by dorsal punctation and structure of thoracic ventrites (Macleay 1885a). Lamprima micardi in Western Australia and L. varians from South Australia were originally distinguished from eastern Australian Lamprima by their narrower male protibial spurs (Reiche 1841; Burmeister 1847) but the presence of intermediates blurs their distinction. We have examined more than 400 specimens of Lamprima from throughout southern and eastern Australia and conclude that these represent just one species, L. aurata. Key to species of Lamprima Latreille, 1804 1. Mandibles as long as or longer than head, without large ventrally directed basal tooth on inner edges (Figs 21–32); protibial spur usually flat, blade-like (Figs 62–75); apical ventrite truncate (males) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .2 - Mandibles shorter than head, each with large anteroventrally directed basal tooth on inner edge, the teeth curved inwards and overlapping in repose (Figs 33–44); protibial spur always narrowly conical; apical ventrite rounded (females; L. imberbis unknown). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .8 2(1) Mandibles with dense erect golden setae on inner surfaces (Figs 21–32), which may be abraded in old specimens; mandible apices blunt, with tips incurved and often subdivided; inner ventral edge of each mandible with prominent tooth or teeth; man- dibles in lateral view gradually contracted from base to middle and pre-apical tooth close to apex or absent (Figs 45–51, 53); protibial spur flat and blade-like, with adjacent setal tuft on a short lobe (Figs 62–63, 65–75). . . . . . . . . . . . . . . . . . . . . . . . . .3 - Mandibles glabrous or almost so on inner surfaces (Fig. 3); mandible apices sharply pointed with straight tips, not subdivided; mandibles without prominent tooth on inner ventral edge; mandibles in lateral view strongly narrowed from base to middle and pre-apical tooth distant from apex (Fig. 52); protibial spur not expanded, without adjacent setal tuft or short lobe (Fig. 64) (dark bronze-brown; length 23 mm; northeastern New South Wales) . . . . . . . . . . . . . . . . . . . . . . . . . . L. imberbis Carter, 1926 3(2) Mandibles elongate, length 27–37% body length, with short peg-like teeth on inner edge beyond ventral mandibular tooth (Figs 1, 25–26, 28, 102) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4 - Mandibles shorter, length 10–28% body length, without short peg-like teeth on inner edge beyond ventral tooth. . . . . . . . . . .5 4(3) Upper surface relatively shiny, usually bright yellowish green (Figs 5, 102); upper edge of mandible in lateral view not evenly curved to apex, but with abrupt elevation before apex (Fig. 48); mesoventrite process rounded at sides; pronotum conspicu- ously punctate, at least at midline (Fig. 25–26); penis not extending beyond parameres (Figs 78–82, 84–88) (length 30–38 mm; Australia). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . L. aurata Latreille, 1817 - Upper surface relatively dull, usually brownish-bronze or dark green (Fig. 1); upper edge of mandible in lateral view evenly curved to apex (Fig. 45); mesoventrite process usually concave at sides; pronotal punctures obscured by dense microsculpture (Fig. 21); penis extending beyond parameres (Fig. 76) (length 30–60 mm; New Guinea) . . . . . . .L. adolphinae (Gestro, 1875) 5(3) Preapical dorsal mandibular tooth present (Figs 40, 42–44); if pronotum green, head not concolourous (Figs 25–32) . . . . . . .6 - Preapical dorsal mandibular tooth absent (Fig. 53); head and pronotum concolorous green or bronze-green (Fig. 24) (pronotum and elytra smooth and shiny, not densely microsculptured, not rugose (Fig. 4); length 18–33 mm; Lord Howe Island) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . L. insularis W.J. Macleay, 1885 6(5) Elytra shallowly sculptured (Figs 5–14, 54–57); protibial spur variable, if narrow then protibiae narrow with long lateral teeth (Figs 62, 66–75); mandible apices blunt, usually subdivided (Figs 22, 27–32); preapical dorsal tooth usually large, acute in lat- eral view (Figs 46, 49–51) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .7 - Elytra deeply sculptured, with irregular but relatively dense grooves and punctures (Fig. 2); protibial spur narrow (length 2x width) in contrast to broad flat protibiae with short blunt marginal teeth (Fig. 63); mandible apices strongly incurved, not sub- divided (Fig. 23); preapical dorsal tooth small, obtuse in lateral view (Fig. 47) (length 26–30 mm; Norfolk Island). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .L. aenea (Fabricius, 1792) 7(6) Upper surface usually relatively shiny, green, coppery, bronze, blue, purple (Figs 7–14, 97–105); elytra conspicuously punctate against relatively even microsculpture (Figs 56–57); mandibles shorter, 10–23% of body length, edge smooth between ventral mandibular tooth and apex (except one specimen with a single secondary tooth); penis not extending beyond parameres (Figs 78–82, 84–88) (length 15–37 mm; Australia). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . L. aurata Latreille, 1817 (typical form) Upper surface dull, usually brownish-bronze, rarely dark green, blue or purple (Fig. 22); elytra inconspicuously punctate against rela- tively uneven microsculpture (Figs 54–55); mandibles longer, 21–27% of body length, with some short irregular projections between ventral mandibular tooth and apex; penis extending beyond parameres (Fig. 76) (length 24–30 mm; New Guinea) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . L. adolphinae (Gestro, 1875) (short mandible form) 8(1) Elytra more shallowly sculptured, without deep grooves (Figs 15, 17–20); spermathecal duct tightly coiled (Figs 89, 91–94) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .9 - Elytra with irregular deep elongate, transverse and oblique grooves (Fig. 16); spermathecal duct loosely coiled (Fig. 90) (head colour variable but not strongly contrasting with pronotum; tergite IX with rounded translucent apex; length 23–27 mm; Nor- folk Island) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .L. aenea (Fabricius, 1792) 160 · Zootaxa 4446 (2) © 2018 Magnolia Press REID ET AL.