An integrative taxonomic and phylogenetic approach reveals a new genus of Australasian Cycas-pollinating weevils (Coleoptera: Curculionidae: Cossoninae) Hsiao, Yun Oberprieler, Rolf G. Zoological Journal of the Linnean Society 2024 Zool. J. Linn. Soc. 2024-01-04 202 2 1 22 Hsiao & Oberprieler 2024 [1008,1210,1529,1556] Insecta Curculionidae Zimmiodes Animalia Coleoptera 6 7 Arthropoda species australis sp. nov.   ( Figs 8A, B, 9A, B, 10A, C, 11A, C, E, 12A–C, 13A, B, 14)  LSID: urn:lsid:zoobank.org:act: 0BCB4DF9-22DF-4C1D-8926- 029FB03D52BA   Type locality  Mulgrave River Forestry Road, Wooroonooran, Queensland, Australia.  Type material   Holotype, ♂:  2. XI. 2019/ AU: QLD: 17°12'37.0"S/ 145°44'58.9"E/ Mulgrave River Rd., / Wooroonooran / Y. Hsiao, Z. Liu& H. Escalona/ leg. // Ex ♂cone of  Cycas media/ banksii// HOLOTYPE/  Zimmiodes australis/ Hsiao& Oberprieler, 2024 ( ANIC).   Figure 5.Terminalia of  Zimmiodes. A, tergite VII, male. B, tergite VII, female. C, tergite VIII, male. D, tergite VIII, female. E, female terminalia. Scale bars: 0.20 mm.   Figure 6.Habitus of final-instar larva of  Zimmiodes australis, lateral view. Scale bar: 1.00 mm.   Paratypes(all labelled ‘PARATYPE /  Zimmiodes australis/ Hsiao & Oberprieler, 2024’): 15 ♂♂, 14 ♀♀, same as holotype ( ANIC).  Other material   Northern Territory:  30 milessouth of Berry Spring,  13 August 1964,  Cycas armstrongii, P.B. Carneleg. ( 1 ♂, 2 ♀♀, 9 exx., ANIC);   30 milessouthof Darwin,  September 1969,  Cycas armstrongii, R.H. Whartonleg. ( 1 ♂, 2 ♀♀, 16 exx., ANIC);  Channel Pointvia Stapleton Station,  25 November 1982,  Cycas canalis, C. Wilsonleg. ( 1 ♂, 7 exx., ANIC);  Rum Jungle, Darwin region,  29 June 1987,  Cycas armstrongii, A .N. Andersenleg. ( 2 exx., ANIC);  Arnhem Highway,  3.1 kmeastjunction with Wanderrie Road,  15 July 1990,  Cycas armstrongii, R. Ornduffleg. ( 1 ♂, 2 exx., ANIC); Road to Mandorah, westof Stuart Highway, southwestof Darwin,  17 July 1990,  Cycas armstrongii, R. Ornduffleg. ( 1 ♂, 1 ex., ANIC);  Acacia Hills, along Stuart Highway, southof Darwin,  18 July 1990,  Cycas armstrongii, R. Ornduffleg. ( 1 ♀, 1 ex., ANIC);  Humpty Doo,  20. August 1995,  Cycas armstrongii, J. Rislerleg. ( 2 ♂♂, 9 exx., ANIC);  12°27 ʹ14.0″S, 130°55 ʹ26.0″E, Wishart, Darwin,  8 April 2006,  Cycas armstrongii, M.F. Brabyleg. ( 1 ♂, 2 ♀♀, 23 exx., ANIC);  12°24 ʹS, 130°55 ʹE, Berrimah,  12 September 2006,  Cycas armstrongii, M.F. Brabyleg. ( 60 exx., 22 larvae, 15 pupae, ANIC);  Darwin,  15 August 2006,  Cycas armstrongii, M.F. Brabyleg. ( 222 exx., ANIC, ANIC database no. 25 077620–25 077621);  12°23 ʹ16.0″S, 136°37 ʹ08.0″E, Repeater Station, Gove Peninsula,  21 August 2007,  Cycas orientis, R.G. Oberprielerleg. ( 24 exx., ANIC, ANIC database no. 25 077624–25 077625);  Goyder River Crossing, Arnhem Land,  27 August 2007,  Cycas arnhemica, R.G. Oberprielerleg. ( 74 exx., ANIC, ANIC database no. 25 077622–25 077623);  12°28'48.0"S 130°43'48.0"E,  7 kmsouth Wagait, Cox Peninsula,  1 July 2021,  Cycas maconochiei, R.G. Oberprielerleg. ( 1 ♂, 5 larvae, 5 pupae, 413 exx., ANIC).  Queensland: same as type series ( 537 exx., ANIC, ANIC database no. 25 077599, 25 077615);  Bamaga,  Cycas semota, B.& Y. Dalzielleg.(nodate) ( 1637 exx., ANIC);  RockyRiver, EastCoast, Silver Plains, Cape York Peninsula,  18 November 1958,  Cycas media ensata, J.L. Wassellleg. ( 2 ♂♂, l larva, 15 exx., ANIC);   15 mileswestof Gordonvale,  13 December 1966, B. Cantrellleg. ( 1 ♂, 21 exx., ANIC);  Bamaga, Cape York,  15–18 June 1969,  Cycas semota, G.B. Monteithleg. ( 1 ♂, 6 exx., ANIC, ANICdatabase no. 25 077618–25 077619);  15°24'36.0"S 145°07'12.0"E, Annan R.  3 kmwest by southof Black Mountain,  27 November 1980,  Cycas media ensata, T. Weirleg. ( 1 ♂, 1 ♀, 11 exx., ANIC);  17°07'48.0"S 145°27'00.0"E, Mulgrave R. Valley,  13 kmsouth by westof Gordonvale,  15 November 1981, J. Baldersonleg. ( 1 ♂, 2 exx., ANIC);  17°09'36.0"S 145°15'36.0"E,  2 kmwest northwestof Mt. Baldy,  860 m,  29 March 1984,  Cycassp., A. Calder& T. Weirleg. ( 2 ♂♂, 95 exx., ANIC);  17°12'36.0"S 144°32'24.0"E,  23 kmeast Almaden,  500 m,  31 March 1984,  Cycas platyphylla, A. Calder& T. Weirleg. ( 2 ♂♂, 1 ♀, 33 exx., ANIC);  Atherton Rifle Range,  17 December 1986, H.& A. Howdenleg. ( 1 ♀, ANIC);   6 kmnorth Ellis Beachnear Cairns,  7 November 1988,  Cycas media banksii, R. Ornduffleg. ( 1 ♂, 1 ♀, 92 exx., ANIC);  Gillies Range Highway,  14 February 1988,  Cycas media banksii, R.G. Oberprielerleg. ( 14 pupae, 236 exx., ANIC);  17°12'36.0"S 144°31'48.0"E,  3.3 kmpast Petford on Almadenroad,  1 June 1991,  Cycas platyphylla, P.I. Forsterleg. ( 1 ♂, 1 ♀, 36 exx., ANIC);  Keppel Sands,  23 October 1991,  Cycas ophiolitica, G.W. Wilsonleg. ( 2 ♂♂, 16 exx., ANIC);  23°20 ʹS, 150°43 ʹE,  6.5 kmalong Keppel Sands Roadoff Emu Park Road, Ross Range,  4 November 1992,  Cycas ophiolitica, P.I. Forster& P.J. Machinleg. ( 229 exx., ANIC);  23°01 ʹS, 150°16 ʹE,  1 kmwestof Glen Geddes,  4 November 1992,  Cycas ophiolitica, P.I. Forster& P.J. Machinleg. ( 16 exx., ANIC);  24°26 ʹS, 151°26 ʹE, Blackman’s Gap Road,  18 kmfrom Bruce Highwaynear Miriam Vale,  4 November 1992,  Cycas megacarpa, P.I. Forster& P.J. Machinleg. ( 5 exx., ANIC);  24°31 ʹS, 151°25 ʹE, Bulburin Road,  5 kmoff Builyanto Bundaberg Road,  5 November 1992,  Cycas megacarpa, P.I. Forster& P.J. Machinleg. ( 14 exx., ANIC);  Keppel Sands,  1 October 1994,  Cycas ophiolitica, G.W. Wilsonleg. ( 4 exx., ANIC);  Tungamul,  8 January 1995,  Cycas ophiolitica, G.W. Wilsonleg. ( 3 ♂♂, 3 ♀♀, 8 exx., ANIC);  17°02 ʹS, 145°21 ʹE, Granite Gorge,  450 m,  26 November 1997,  Cycassp., Monteith, Cook& Burwellleg. ( 7 exx., QMBA);  24°04 ʹ30″S, 150°39 ʹ11″E, State Forest  618, 38 kmnorth northeastof Biloela,  22 November 2003,  Cycas megacarpa, P.I. Forsterleg. ( 14 exx., ANIC);  24°08 ʹ24.74″S, 150°44 ʹ19.52″E,  10 December 2018, probably  Cycas ophiolitica(no collector) ( 8 exx., ANIC, ANIC database no. 25 077616–25 077617);  Near Palm Cove, Captain Cook Highway,  29 October 2019,  Cycas media banksii, Y. Hsiaoleg. ( 8 exx., ANIC);  16°42 ʹ07.7″S, 145°37 ʹ18.9″E, Captain Cook Highway,  2 November 2019,  Cycas media banksii, Y. Hsiao& L. Lileg. ( 1 ♂, 3 exx., ANIC);  17°09 ʹ08.3″S, 145°42 ʹ18.0″E, Little Mulgrave Forest Reserve, Lamb Range, ~  220 m,  2 November 2019,  Cycas media banksii, Y. Hsiao& H. Escalonaleg. ( 1 ♂, 27 exx., ANIC);  16°37 ʹ46.0″S, 145°32 ʹ56.5″E, Oak Beach, Captain Cook Highway,  2 November 2019,  Cycas media banksii, Y. Hsiao& L. Lileg. ( 1 ♂, 39 exx., ANIC).   Diagnosis Adults of this species are distinguishable from  Z. papuanusby the following characters (states of  Z. papuanusin parentheses): rostrum nearly straight, slightly ventrally curved, slightly thickened at antennal insertions ( Figs 8A, B, 9A, B) (distinctly ventrally curved, distinctly thickened at antennal insertions; Figs 8C, D, 9C, D); eyes strongly convex and protruding ( Figs 8A, B, 9A, B) (slightly flatter; Figs 8C, D, 9C, D); scapes thinner and internal margin of funicle segments 1 straight ( Fig. 10A) (scapes thicker and internal margin of funicle segments 1 distinctly emarginate basally; Fig. 10B); pronotal punctures coarser ( Fig. 8A, B) (finer; Fig. 8C, D); elytral interstriae distinctly convex ( Figs 8A, B, 9A, B) (flatter; Figs 8C, D, 9C, D); profemora moderately narrowing apicad ( Fig. 10C) (distinctly obtusely protruding inwards in male; Fig. 10D); sternite VIII of male with smaller sclerotized lateral plates and larger membranous apical plate, of female with lateral margins angled apically ( Fig. 11A, C) (of male with larger lateral plates and smaller apical plate, of female with lateral margins slightly rounded apically; Fig. 11B, D); tegmen with manubrium longer ( Fig. 12A–C) (very short; Fig. 12D–F); penis short and thick, with temones ~2.0× as long as body of penis ( Fig. 12A–C) (elongate and slender, with temones ~1.2× as long as body of penis; Fig. 12D–F); endophallus short, medially with a sleeve of weakly developed denticles, basally with a pair of sclerites ( Fig. 12A–C) (long, extending below body of penis, medially with a sleeve of strongly developed denticles, apically with a pair of sclerites; Fig. 12D–F).   Description  Shape and size:Body slender and elongate ( Fig. 8A, B), length 2.3–3.2 mmin both sexes, width ~0.35–0.37× length, weakly convex in lateral view ( Fig. 9A, B).  Colour and vestiture:Body and legs reddish to blackish brown, semilustrous, glabrous.   Figure 13.Female genitalia of  Zimmiodes. A,  Zimmiodes australis, ovipositor. B,  Z. australis, spermatheca with gland. C,  Zimmiodes papuanus, ovipositor. D,  Z. papuanus, spermatheca with gland. Scale bars: 0.20 mm.   Figure 14.Geographical distribution of  Zimmiodesin Australia and Papua New Guinea.  Head:Rostrum: moderately long, as long as pronotum, robust, slightly thinner in female, nearly straight, slightly ventrally curved, dorsoventrally flattened, slightly broadened apically in dorsal view, coarsely punctate dorsally, punctures finer in distal half. Eyes: subcircular in outline, strongly convex and protruding ( Fig. 3A, B). Antennae: inserted close to base of rostrum in male and female ( Figs 8A, B, 9A, B); scapes reaching eye; funicles with segment 1 ~0.9×, 1.6×, 1.6×, and 1.5× as long as segments 2–5, respectively; clubs stout, ~1.7–1.8× as long as wide, densely and finely pubescent ( Figs 3C, 10A).  Thorax:Pronotum: roundly trapezoidal, apex ~0.6× as wide as base ( Fig. 3D); anterior and posterior margins subtruncate, lateral margins mostly rounded, distinctly narrowed anteriorly; disc weakly and evenly convex; surface densely and distinctly punctate. Scutellar shield: roundly pentagonal. Elytra: ~2.1– 2.2× as long as pronotum, jointly ~0.5× as broad as long, slightly broader base of pronotum; humeri broadly rounded, slightly protruding; surface slightly evenly convex, deeply and coarsely punctate in rows, forming distinct striae. Legs: femora ventrally without subapical tooth ( Figs 4, 10C); tibiae with premucro smaller than uncus; tarsi with claws free, divergent.   Terminalia:Sternite VIII of male subtrapezoidal, plates sclerotized, slanting, apical margin rounded, sparsely covered with short setae subapically, basal margin strongly sclerotized ( Fig. 11A), of female abruptly angled laterally, with apical margin densely covered with short setae ( Fig. 11C); spiculum gastrale strongly curved apically, base lightly sclerotized ( Fig. 11E); tegmen with complete ring, without parameroid lobes, manubrium short; penis thick (~1.6– 1.8× as long as wide), subparallel-sided, slightly widening apicad, moderately narrowed apically, dorsum strongly sclerotized, without median groove, apical margin dorsally roundly emarginate, ventrally broadly rounded, apical opening with a pair of lanceolate orificial sclerites, temones slender, ~2.0× as long as body of penis, anteriorly confluent, joined ventrally to body of penis ( Fig. 12A–C); endophallus short, membranous, medially with a sleeve of dense, weakly developed denticles, basally with a pair of slender sclerites, bent outwards; ovipositor with gonocoxites slender, elongate, apically bluntly rounded, styli short, subcylindrical, bluntly rounded and setose apically ( Fig. 13A); spermatheca evenly narrow, right-angled, gland elongate, narrowing apicad, rounded apically ( Fig. 13B).   Derivation of name The specific epithet  australismeans ‘southern’, referring to the only known distribution of the species in Australia.   Distribution This species is known from the coastal regions of the Northern Territoryto central Queensland( Fig. 14). Owing to the distributional range of  Cycasin Australia, it is probably also distributed in northern Western Australia, and it might also occur in southern Papua New Guinea, where three species of the Cycad mediagroup occur ( Hill 1994).   Natural history The species apparently has a considerably wide range of hosts, including  Cycas armstrongii,  Cycas arnhemica,  Cycas canalis,  Cycas maconochiei,  Cycas media banksii,  Cycas media ensata,  Cycas megacarpa,  Cycas ophiolitica,  Cycas orientis,  Cycas platyphyllaand  Cycas semota. The role of  Z. australisin the pollination of  Cycashas been studied by Hall and Walter (2018), who mentioned it as an undescribed weevil ( Curculionidae). Based on our field observations on  Cycas media banksiiin northern Queensland( Fig. 1A) and on  Cycas armstrongii,  Cycas arnhemicaand  Cycas maconochieiin the Northern Territory, large numbers of adults congregate among the sporophylls of microstrobili (male cones) during the coning period, with> 500 adultscollected from a single cone ( Fig. 1B, C), and smaller numbers of weevils also gather in young (closed) megastrobili, predominantly among the inner sporophylls. The larvae develop only in the male cones, tunnelling and feeding in the sporophylls and the central rachis ( Fig. 1F), where they also pupate ( Fig. 1G), not leaving the cones to pupate in surrounding substrates as do some species of  TranesSchoenherr ( Hall et al.2004)and  MiltotranesZimmerman ( Hsiao and Oberprieler 2020a). The adults eclose from their pupal chambers on the lower surface of the outer (thicker) part of the sporophylls and inside the disintegrating rachis. As reported in previous studies ( Ornduff 1991, Forster et al.1994),  Z. australiscommonly co-exists with certain other beetles in the male cones, mainly  Aethina( Circopes) unguiculataKirejtshuk, 1987( Nitidulidae),  Hapalipssp.( Erotylidae) and  Ulomoides australis(Carter, 1932)( Tenebrionidae), but the numbers of these beetles are considerably smaller than those of  Z. australis. Adults of  Z. australisalso visit young (closed) female strobili of  Cycas( Oberprieler 2004, Hall and Walter 2018) and play a major role in the pollination of their hosts ( Hall and Walter 2018). 4989270342 2019-11-02 Y. Hsiao & Z. Liu & H. Escalona Australia -17.21028 Mulgrave River 1 145.7497 Wooroonooran 6 7 1 1 Queensland holotype 4989270304 2019-11-02 ANIC Y. Hsiao & Z. Liu & H. Escalona Australia Hsiao & Oberprieler -17.21028 Mulgrave River 1 145.7497 Wooroonooran 8 9 29 14 15 Queensland paratype 4989270308 1964-08-13 ANIC P. B. Carne Berry Spring 8 9 12 2 1 Northern Territory 4989270340 1969-09 ANIC R. H. Wharton Darwin 8 9 19 2 1 Northern Territory 4989270343 1982-11-25 ANIC C. Wilson Stapleton Station Channel Point 8 9 8 1 Northern Territory 4989270303 1987-06-29 ANIC . N. Andersen Darwin region Rum Jungle 8 9 2 Northern Territory 4989270341 1990-07-15 ANIC R. Ornduff Wanderrie Road Arnhem Highway 8 9 3 1 Northern Territory 4989270324 1990-07-17 ANIC R. Ornduff Road Stuart Highway Mandorah 8 9 2 1 Northern Territory 4989270339 1990-07-18 ANIC R. Ornduff Acacia Hills Darwin Stuart Highway 8 9 2 1 Northern Territory 4989270320 1995-08-20 ANIC J. Risler Humpty Doo 8 9 11 2 Northern Territory 4989270330 2006-04-08 ANIC M. F. Braby -12.453889 Darwin 1 130.92389 Wishart 8 9 26 2 1 Northern Territory 4989270305 2006-09-12 ANIC M. F. Braby -12.4 Berrimah 1293 130.91667 8 9 97 22 15 Northern Territory 4989270327 2006-08-15 ANIC M. F. Braby Darwin 8 9 25 077620-25 077621 222 Northern Territory 4989270326 2007-08-21 ANIC R. G. Oberprieler -12.387777 Gove Peninsula 1 136.6189 Repeater Station 8 9 25 077624-25 077625 24 Northern Territory 4989270306 2007-08-27 ANIC R. G. Oberprieler Arnhem Land Goyder River Crossing 8 9 25 077622-25 077623 74 Northern Territory 4989270311 2021-07-01 ANIC R. G. Oberprieler -12.48 Cox Peninsula 1 130.73 Wagait 8 9 424 5 1 5 Northern Territory 4989270325 ANIC Queensland 8 9 25 077599, 25 077615 537 Queensland 4989270316 ANIC B. & Y. Dalzielleg. Bamaga 8 9 1637 Queensland 4989270314 1958-11-18 ANIC J. L. Wassell RockyRiver EastCoast EastCoast 8 9 1 2 Queensland 4989270337 1966-12-13 ANIC B. Cantrell Gordonvale 8 9 22 1 Cape York 4989270321 1969-06-15 1969-06-18 1969-06-15 ANIC G. B. Monteith Bamaga 8 9 25 077618-25 077619 7 1 Cape York 4989270312 1980-11-27 ANIC T. Weir -15.41 Black Mountain 1 145.12 Annan R. 8 9 13 1 1 Cape York 4989270332 1981-11-15 ANIC J. Balderson Mulgrave -17.130001 Gordonvale 1 145.45 Mulgrave R. Valley 8 9 3 1 Cape York 4989270313 1984-03-29 ANIC A. Calder & T. Weir 860 -17.16 Mt. Baldy 1 145.26 8 9 97 2 Cape York 4989270328 1984-03-31 ANIC A. Calder & T. Weir 500 13 14 -17.210001 Almaden 1 144.54001 8 9 36 1 2 Cape York 4989270301 1986-12-17 ANIC H. & A. Howden Atherton Rifle Range 13 14 1 1 Cape York 4989270334 1988-11-07 ANIC R. Ornduff Cairns Ellis Beach 13 14 94 1 1 Cape York 4989270322 1988-02-14 ANIC R. G. Oberprieler Gillies Range Highway 13 14 250 14 Cape York 4989270307 1991-06-01 ANIC P. I. Forster -17.210001 Petford on Almaden 1 144.53 13 14 38 1 1 Cape York 4989270319 1991-10-23 ANIC G. W. Wilson Keppel Sands 13 14 18 2 Cape York 4989270344 1992-11-04 ANIC P. I. Forster & P. J. Machin Keppel Sands Road -23.333334 Emu Park Road 1256 150.71666 Emu Park Road 13 14 229 Cape York 4989270336 1992-11-04 ANIC P. I. Forster & P. J. Machin -23.016666 Glen Geddes 1257 150.26666 13 14 16 Cape York 4989270317 1992-11-04 ANIC P. I. Forster & P. J. Machin Blackman's Gap Road -24.433332 Bruce Highway 1251 151.43333 Bruce Highway 13 14 5 Cape York 4989270310 1992-11-05 ANIC P. I. Forster & P. J. Machin Bulburin Road -24.516666 Builyan 1251 151.41667 Builyan 13 14 14 Cape York 4989270309 1994-10-01 ANIC G. W. Wilson Keppel Sands 13 14 4 Cape York 4989270315 1995-01-08 ANIC G. W. Wilson Tungamul 13 14 14 3 3 Cape York 4989270346 1997-11-26 QMBA Monteith & Cook & Burwell 450 -17.033333 Granite Gorge 1279 145.35 13 14 7 Cape York 4989270329 2003-11-22 ANIC P. I. Forster 14 15 -24.075 Biloela 21 150.65305 State Forest 13 14 14 Cape York 4989270345 2018-12-10 ANIC -24.140205 1 150.73875 14 15 25 077616-25 077617 8 Cape York 4989270338 2019-10-29 ANIC Y. Hsiao Captain Cook Highway Near Palm Cove 14 15 8 Cape York 4989270335 2019-11-02 ANIC Y. Hsiao & L. Li -16.702139 Captain Cook Highway 1 145.62192 14 15 4 1 Cape York 4989270318 2019-11-02 ANIC Y. Hsiao & H. Escalona 220 -17.152306 Lamb Range 1 145.705 Little Mulgrave Forest Reserve 14 15 28 1 Cape York 4989270302 2019-11-02 ANIC Y. Hsiao & L. Li -16.629444 Captain Cook Highway 1 145.54904 Oak Beach 14 15 40 1 Cape York