ACTA ENTOMOLOGICA MUSEI NATIONALIS PRAGAE www.aemnp.euISSN 1804-6487 (online) – 0374-1036 (print) R E S E A R C H P A P E R Phylogeny, diversity and biogeography of fl ightless amphi-Pacifi c lymantine weevils (Coleoptera: Curculionidae: Molytinae) Vasily V. GREBENNIKOV1) & Robert S. ANDERSON2) 1) Canadian Food Inspection Agency, 960 Carling Ave., Ottawa, ON, K1A 0Y9, Canada; e-mail: vasily.grebennikov@inspection.gc.ca 2) Beaty Centre for Species Discovery, Canadian Museum of Nature, PO Box 3443, Station D, Ottawa, ON, K1P 6P4, Canada; e-mail: randerson@nature.ca Abstract. We use DNA sequence data to generate the fi rst phylogenetic hypothesis for the weevil tribe Lymantini. These are leaf litter inhabiting beetles generally regarded as restricted to the New World and taxonomically arranged in two subtribes, 11 genera and some 150 named species. An additional genus of questionable affi nities to the tribe, Devernodes Grebennikov, 2018, has fi ve described species in Southeastern Asia. All these beetles are fl ightless and some have eyes reduced in size or absent, traits normally associated with limited dispersal capacity. We performed a phylogenetic analysis of 153 terminals (50 of them belong to Lymantini re- presenting Devernodes and all but three named genera) based on 4,174 bp alignment of one mitochondrial (cox1) and two nuclear fragments (ITS2 and 28S). We fi nd that both Lymantini subtribes Lymantina and Caecossonina are monophyletic, the latter sister to the amphi-Atlantic tribe Anchonini. The Asian genus Devernodes is deeply nested among American Lymantina. The clade of Anchonini plus Lymantini is consistently recovered outside of the CCCMS clade of “higher” weevils (Curculioninae, Conoderinae, Cossoninae, Molytinae and Scolytinae). We hypothesize that the polished head capsule of adult beetles is an apomorphy of Anchonini and Lymantini, the 8-segmented antennal funicle is an apomorphy of Anchonini plus Caecossoni- na. We attribute the origin of the currently observed amphi-Pacifi c distribution of Lymantina to normal ecological dispersal facilitated by the warmer periods of the Cenozoic such as the Eocene, and by presently submerged Arctic land bridges. Using parsimony we hypothesize a North American origin for the Anchonini plus Lymantini crown group, as well as that of Lymantina. We argue that Bronchotibia adunatus Poinar & Legalov, 2021, a Dominican amber adult weevil fossil, is not a member of Lymantini and re-classify it as Curculionidae incertae sedis. We present an image gallery of 28 Lymantini specimens to document the morphological diversity of the tribe. We hypothesize the existence of unnamed American genera of Lymantina and make public the DNA-barcode dataset of 89 Lymantini specimens. Key words. Coleoptera, Anchonini, Caecossonina, Lymantina, DNA barcode, ITS2, 28S, phylogeny, forest litter, biogeography Zoobank: http://zoobank.org/urn:lsid:zoobank.org:pub:C9E0D0F0-F4F7-4946-B2B2-83E1F9F706E2 © 2022 The Authors. This work is licensed under the Creative Commons Attribution-NonCommercial-NoDerivs 3.0 Licence. Accepted: 7th November 2022 Published online: 31st December 2022 2022 62(2): 411–442 doi: 10.37520/aemnp.2022.023 valid genera, and some 150 described species, almost 100 of these placed in the genus Theognete Champion, 1902. Adult Lymantini are fl ightless, have eyes variously reduced in size or absent, and are collected in primary forests by sifting leaf litter, or in the soil, or rarely, in caves. Imma- ture stages of Lymantini have been described only once, as larvae of an unknown genus “near Ithaura Pascoe and Dioptrophorus Faust” found in sweet potatoes in Mexico (A 1952). No reliable fossil records of the tribe are known. Assignment to the tribe of the monotypic genus Introduction Lymantini, the focal group of this paper, are small to me- dium-sized elongate weevils (Figs 1, 2) distributed between the USA and Bolivia, including the West Indies (Fig. 3). Taxonomic boundaries and placement of these beetles have always been, and remain, murky (e.g., H 1992). As most recently defi ned (A -Z & L 1999, L 2014) and excluding the subsequently discovered Asian genus Devernodes Grebennikov, 2018, extant Lymantini diversity is composed of two subtribes, 11 GREBENNIKOV & ANDERSON: Phylogeny, diversity and biogeography of Lymantini (Coleoptera: Curculionidae: Molytinae)412 Bronchotibia Poinar & Legalov, 2021 described based on a Dominican amber fossil (P & L 2021) was based on non-informative characters employed outside of the phylogenetic framework and must be re-interpreted (see Discussion). The weevil tribe Lymantini has at least three scienti- fi cally intriguing peculiarities. Firstly, the monophyly of the tribe remains untested and the sister group unknown. Secondly, the number of named Lymantini species is li- kely highly underestimated. Thirdly, although exclusively fl ightless, often with reduced eyes, and biologically linked with primary wet forests (and, therefore, presumably seve- rely restricted in their dispersal capacity), this New World taxon has been recently tentatively reported from Southeas- tern Asia (G 2018) implying an amphi-Pacifi c distribution for the tribe. This paper is our attempt to shed the fi rst evolutionary light on all these issues. The monophyly of the tribe Lymantini, although never explicitly challenged, has never been tested in a formal phylogenetic analysis. The taxonomic recognition of the tribe implicitly suggesting its monophyly was historically pivoted on the biological association of these weevils with the forest leaf litter or the soil, on their coherent North American distribution, and two potential morphological apomorphies (Fig. 4). Firstly, at least the eyed members of Lymantini are immediately recognizable among almost all weevils by having their eyes “… placed on the rostral part of the head, which is often sharply delimited from the main head capsule, sometimes by a dorsal, lateral, and even ventral groove” (L 2014; a similar condition is also found in some Cycloterini, Phrynixini and Orthorhinini; L 2014). Three Lymantini genera forming the subtri- be Caecossonina, as well two cave species of the genus Lymantes Schoenherr, 1838, although possessing the trans- verse rostral groove, are eyeless. Secondly, all Lymantini examined in this respect have the female hemisternites IX undivided by a transverse membrane. This character state was previously called “fused coxite-stylus” (H 1992) or “lack styli on the coxites” (A 2016). If indeed monophyletic, the sister group of the tribe Lyman- tini is entirely unknown, while its taxonomic assignment oscillates between Molytinae and Cossoninae (reviewed in H 1992). Remarkably, the tribe’s putative sole Asian genus Devernodes was resolved as a sister to the primarily American tribe Anchonini (G & A 2021a) although this was without the inclusi- on of data about undescribed Lymantini presented here. Even more surprising, this moderately supported clade in a molecular phylogeny was placed outside of the large and strongly supported CCCMS clade of “higher” weevils (Curculioninae, Conoderinae, Cossoninae, Molytinae and Scolytinae), to which both Lymantini and Anchonini are assigned taxonomically. Besides uncertain monophyly and phylogenetic place- ment, Lymantini are likely acutely under-sampled, under- studied, and, therefore, remain largely unknown to science. During the 30+ years of Lymantini studies, one of us (RA) accumulated specimens of multiple unnamed species and perhaps genera. The assumption of under representation was corroborated by the recent revision of the Mesoame- rican genus Theognete, which increased the number of named species from one to 94 (A 2010). If the same ratio remains true throughout the rest of the tribe, Lymantini extant diversity might rival that of amphibians (about 5,700 species) or mammals (about 5,400 species). Two recent developments triggered our study. Firstly, the newly described genus Devernodes containing fi ve new species from Southern China, Vietnam and Malaysia was tentatively assigned to the otherwise exclusively American tribe Lymantini (G 2018). Consistent with the rest of Lymantini, all species of Devernodes are wingless and found by sifting forest leaf litter. Moreover, all species of Devernodes have both putative Lymantini morphological synapomorphies: the peculiar constriction separating the eye-bearing rostrum from the head capsule, as well as the undivided female hemisternite IX (Fig. 4). At the time of the discovery of Devernodes, no Mesoamerican Lymantini were available for DNA sequencing and, therefore, assignment of this Asian genus to the tribe was made based on similarities, rather than on a formal phylogenetic analysis. Secondly, this genus of questionable relationship to Lymantini was resolved as a moderately supported sister to the re-defi ned, monophyletic, and primarily Mesoamerican fl ightless tribe Anchonini (G & A 2021a). The latter clade is morphologically supported by the antennal funicle consisting of eight (not seven or less) antennomeres. Cu- riously, the same trait is also diagnostic for the sympatric Lymantini subtribe Caecossonina uniting all eyeless mem- bers of the latter tribe (excepting two convergent eyeless cave Lymantes species) and, therefore, “bridging the gap” between both Anchonini and Lymantini. Lacking any Mesoamerican Lymantini in the analysis, and assuming Devernodes represented Lymantini, we suggested (G - & A 2021a) that Devernodes, Lymantina, Caecossonina and Anchonini might form a clade supported by at least one morphological apomorphy: the polished head capsule of adult beetles. Remarkably, so defi ned, this group corresponds to “Anchonina” by C (1902: 66, 1903). This author emphasised the same morphological character (“... recognizable by their globose, deeply inserted, almost smooth head...”) and provided two large plates of high-quali- ty Anchonini and Lymantini illustrations. In 2021, however, we lacked suffi cient DNA data and, therefore, were unable to test the monophyly of C ’ “Anchonina”. Results of G (2018) and G & A (2021a), therefore, strongly suggested a Lymantini-focused phylogenetic analysis designed to test the following predictions: 1) All Mesoamerican Lymantini family- and genus- group taxa, as defi ned in L (2014), that is the tribe, both subtribes and all non-monotypic genera, are monophyletic. 2) Monophyletic Asian Devernodes has its sister among American Lymantini and if so, then a plausible interpreta- tion might be off ered to explain the disjunct amphi-Pacifi c distribution of these low-dispersing terrestrial animals. 3) The polished head capsule is a synapomorphy of Anchonini and Lymantini (= “Anchonina” of C 1902, 1903). Acta Entomologica Musei Nationalis Pragae, volume 62, number 2, 2022 413 Fig. 1. Morphological diversity of the weevil tribe Lymantini, dorsal view. Specimen numbers refer to Table 2 and Fig. 7. GREBENNIKOV & ANDERSON: Phylogeny, diversity and biogeography of Lymantini (Coleoptera: Curculionidae: Molytinae)414 Fig. 2. Morphological diversity of the weevil tribe Lymantini, lateral view. Specimen numbers refer to Table 2 and Fig. 7. Acta Entomologica Musei Nationalis Pragae, volume 62, number 2, 2022 415 Fig. 3. Geographical distribution and hypothesized overland dispersal routes of recent Lymantina. Fig. 4. Morphological diagnostic features and possible apomorphies of Anchonini (A, B) and Lymantini (C–F). A, C, D: head, left lateral view; B: left antenna; E, F: female genitalia and apical sclerites (E: ventral, F: right dorso-lateral). A: Titilayo geiseri Cristóvão & Lyal, 2018; B: T. barclayi Cris- tóvão & Lyal, 2018; C: Lymantes scrobicollis Gyllenhal, 1838; D–F: Devernodes chthonia Grebennikov, 2018. A, B: from G & A (2021a); E, F: from G (2018). GREBENNIKOV & ANDERSON: Phylogeny, diversity and biogeography of Lymantini (Coleoptera: Curculionidae: Molytinae)416 Fig. 5. Morphological diversity of the weevil tribe Lymantini, antennae. Specimen numbers refer to Table 2 and Fig. 7. Acta Entomologica Musei Nationalis Pragae, volume 62, number 2, 2022 417 4) the 8-segmented antennal funicle is a synapomorphy of Anchonini and Caecossonina. 5) The tribe Lymantini is outside the CCCMS clade of “higher” weevils. 6) The tribe Lymantini contains unnamed genera. This paper is our attempt to test all these predictions using the newly constructed fi rst phylogenetic tree of American Lymantini and their relatives, itself based on the newly generated DNA data. Our more inclusive goal is to establish a baseline for the further research of these and related organisms by releasing their genetic, morphological and geographical data. Specifi cally, we want to document the morphological diversity of Lymantini, output an on- line DNA-barcode (H et al. 2003) dataset of these organisms, and generate their fi rst phylogenetic tree, the latter likely including representatives of unnamed genera awaiting formal description. Overall we designed this pa- per to trigger and facilitate further evolutionary research of these morphologically distinct, diverse, and acutely understudied organisms. Material and methods Sampling of Mesoamerican Lymantini. Herein newly sequenced adult specimens of Mesoamerican Lymantini were sifted from forest leaf litter using hand-held sifters, with subsequent extraction of live specimens using suspen- ded Winkler funnels. Live adults were preserved in 96% ethanol and subsequently sorted, imaged, and processed for DNA barcoding, mounted on pins, and stored in the Canadian National Collection of Insects, Arachnids and Nematodes (CNC, Ottawa) or the Canadian Museum of Nature (CMN, Ottawa). Images and geographic data of each herein reported 50 Lymantini specimens (and of all non-Lymantini outgroups used in the analysis, Table 2) are available online in the public datasets of the Barcode of Life Data System (BOLD, R & H 2007); see below. DNA sequencing. Three DNA fragments were sequenced and analyzed (Table 1, fragment abbreviations are in brack- ets): mitochondrial cytochrome c oxidase subunit I (COI, the DNA barcode fragment), nuclear internal transcribed spacer 2 (ITS2) and nuclear 28S ribosomal DNA (28S). Sequencing of DNA was made at the Canadian Centre for DNA Barcode (http://ccdb.ca/) using standard protocols. The primers are listed in Table 1 in G (2017). All details of the lab work (such as DNA extraction, ampli- fi cation, PCR protocols), as well as images of the original electropherograms for all specimens, are available online in BOLD, in two public datasets, each pertaining to one of two herein implemented analyses (see below). DNA barcode dataset of Lymantini. Our fi rst analysis was to generate and make public the DNA barcode dataset of Lymantini, following the currently accepted tribal limits (even if perhaps non-monophyletic; see Results). By doing this, we wanted to document the genetic, morphological and geographic diversity of Lymantini available to us, even if many of them represent unnamed and/or unidentifi ed ge- nera and/or species. This DNA barcode dataset is designed to facilitate future taxonomic and other research of these beetles by allowing direct online comparison between our specimens and those of others. This dataset contains 89 DNA barcodes of Lymantini, each of them longer than 350 base pairs (bp) and most of them 658 bp. The dataset includes 26 specimens of the Asian genus Devernodes (their DNA barcodes fi rst released in G 2018), plus 63 newly generated DNA barcodes of American Ly- mantini. We subjected these DNA barcodes to the standard analytical pipeline procedure implemented in the BOLD website (http://www.boldsystems.org/) by clustering them into an unrooted topology using the Neighbour Joining (=NJ) algorithm (BOLD commands “Sequence Analysis: taxon ID tree”). For this purpose, we used the default Kimura 2 model of nucleotide substitutions and selected “BOLD Aligner” for the “Align Sequences” parameter. In this analysis we used the Barcode Index Numbers (BINs, R & H 2013), to identify minimal evolutionary signifi cant units. The resulting topology with GenBank accession numbers of all 89 DNA barcodes of Lymantini are in Supplementary File. The entire dataset is available online as a BOLD public dataset at dx.doi. org/10.5883/DS-VGDS25. Selection of terminals for a phylogenetic analysis. Our second analysis was a phylogenetic one, based on a three- marker DNA dataset of Lymantini, and designed to test all six predictions made in the Introduction. The ingroup was formed by 45 newly sequenced representatives of Mesoamerican Lymantini (41 of Lymantina and four of Caecossonina) plus fi ve terminals representing all fi ve valid species of the Asian genus Devernodes. We used the DNA barcode tree as a guide to maximizing the phy- logenetic diversity of the ingroup. The nearest outgroup was formed by 17 representatives of the tribe Anchonini, as suggested by the recovery of Devernodes sister to this tribe (G & A 2021a). The more distant outgroup was formed by 60 representatives of the CCCMS clade other than Anchonini, Lymantini, or Devernodes, as well as by 14 representatives of the CEGH clade (Cyclo- minae, Entiminae, Gonipterini and Hyperinae, S et al. 2017), which forms the sister to the CCCMS clade (S et al. 2017). We specifi cally included fi ve representatives of the predominantly Australian and New Zealand tribe Phrynixini (Molytinae) because of similarly to Lymantini as at least some Phrynixini have their eye positioned at the base of the rostrum (L 2014). Since the monophyletic subfamily Dryophthorinae consistently branches outside the CCCMS plus CEGH clade, fi ve representatives of this subfamily were added as more distant outgroups. Consi- Table 1. DNA fragments used in the phylogenetic analyses of Lymantini weevils, followed by total number of sequenced terminals, minimal, maximal, and aligned length of each fragment, and the fi rst and the last position of each aligned fragment in the concatenated matrix. Fragment # min max aligned positions COI-5P 152 353 658 658 1 to 658 ITS2 133 223 763 2645 659 to 3303 28S 152 341 607 871 3304 to 4174 GREBENNIKOV & ANDERSON: Phylogeny, diversity and biogeography of Lymantini (Coleoptera: Curculionidae: Molytinae)418 Table 2. DNA fragments and their GenBank accession numbers of 153 weevil (Coleoptera) specimens used in the three marker phylogenetic analysis of the tribe Lymanini (including 137 newly sequenced fragments shown in bold: OL671058–OL671194). Voucher Subfamily Tribe Genus and/or species Country COI ITS2 28S 431 Molytinae Molytini Anchonidium unguiculare Morocco HM417678 none KY110382 434 Dryophthorinae Rhynchophorini Sphenophorus parumpunctatus Morocco HM417724 KY110320 KY110384 487 Molytinae Emphyastini Thalasselephas maximus Russia HM417677 KY110313 KY110375 703 Molytinae Pissodini Pissodes punctatus China HQ987002 none KY110366 704 Molytinae Ithyporini Ectatorhinus adamsii China HQ987003 KY110315 KY110377 861 Molytinae none Zembrus perseus China HQ987100 MG648823 MG648736 1678 Cossoninae Rhyncolini Himatium Tanzania JN265954 KY110323 KY110388 1791 Cossoninae Dryotribini Trichopentarthrum uluguricus Tanzania JN265975 KY110327 KY110392 2288 Molytinae Lymantini Devernodes alkippe China MH034387 MH034357 MH034414 2533 Molytinae Aminyopini Niphadonyx China MG648752 MG648826 MG648738 2640 Molytinae Anchonini Himalanchonus China MT889126 MT889147 MT889172 2676 Molytinae none Aater cangshanensis China MG648761 MG648835 MG648747 2723 Molytinae Aminyopini Niphades China MG648751 MG648825 MG648737 2731 Molytinae Molytini Niphadomimus maia China KJ427744 KY110324 KY110389 2735 Dryophthorinae Rhynchophorini Sitophilus zeamais China KJ672255 MG968837 MG968894 2955 Molytinae Molytini Lobosoma rausense Russia KJ427738 KY110316 KY110378 2968 Entiminae Alophini Trichalophus alternatus Canada KM538666 MW536413 MW536465 2970 Cossoninae Rhyncolini Carphonotus testaceus Canada KY110606 KY110309 KY110371 3060 Molytinae Molytini Lupangus asterius Tanzania KY034280 KY250485 KY250480 3280 Molytinae Cycloterini Prothrombosternus tarsalis Tanzania KU748541 KY110337 KY110402 3561 Dryophthorinae Dryophthorini Dryophthorus Tanzania MG968913 MG968814 MG968871 4118 Molytinae Molytini Microplinthus China MG648755 MG648829 MG648741 4337 Molytinae Lymantini Devernodes asteria Vietnam MH034376 MH034352 MH034409 4339 Molytinae Lymantini Devernodes chthonia Vietnam MH034400 MH034364 MH034421 4353 Molytinae Lithinini Seleuca Vietnam MG648754 MG648828 MG648740 4355 Molytinae none Otibazo polyphemus Vietnam KJ841732 KY110328 KY110393 4402 Dryophthorinae Stromboscerini Nephius argus Vietnam MH034380 MH034354 MH034411 4537 Molytinae Molytini Morimotodes ismene China KJ871649 KY110338 KY110403 4846 Molytinae Cycloterini Thrombosternus cucullatus Tanzania KJ445714 KY110335 KY110400 4991 Molytinae Aminyopini Niphadonothus gentilis Tanzania KX360489 KY110336 KY110401 5001 Molytinae Molytini Aparopionella elliptica Tanzania KX360455 KY110318 KY110381 5402 Entiminae Cneorhinini Catapionus mopsus China KU748534 MW536396 MW536448 5848 Molytinae Molytini Adexius scrobipennis Poland KJ445686 KY110305 KY110367 5954 Molytinae Lymantini Devernodes drimo Malaysia MH034401 MH034365 MH034422 5975 Molytinae Lymantini Devernodes methone Malaysia MH034390 MH034360 MH034417 6485 Molytinae Molytini Plinthus amplicollis Georgia KY110617 KY110331 KY110396 6552 Molytinae Molytini Aparopion costatum Georgia KJ445700 none KY110387 6608 Molytinae Molytini Leiosoma reitteri Georgia KJ445698 KY110322 KY110386 6683 Molytinae Molytini Euthycus Taiwan KJ445702 KY110325 KY110390 6858 Molytinae Lithinini Seleuca Taiwan KY110611 KY110317 KY110380 7166 Molytinae Molytini Typoderus antennarius Tanzania KY250487 KY250484 KY250479 7281 Molytinae Cycloterini Allocycloteres circellariceps Tanzania MK813366 MK813357 MK813361 7530 Cryptorhynchinae Cryptorhynchini Cryptorhynchus lapathi Russia KY110605 KY110303 KY110365 7531 Molytinae Aminyopini Niphades verrucosus Russia KY110610 KY110314 KY110376 8046 Molytinae Aminyopini Niphades Tanzania MG648748 MG648821 MG648734 8317 Molytinae Aminyopini Niphades Cameroon MG648749 MG648822 MG648735 8474 Molytinae Lepyrini Lepyrus palustris Poland KX360483 KY110332 KY110397 8480 Molytinae Molytini Leiosoma defl exum Poland KY110614 KY110326 KY110391 8484 Molytinae Trachodini Trachodes hispidus Poland KX360436 KY110307 KY110369 8489 Brachycerinae Erirhinini Notaris scirpi Poland KR736279 MW201453 MW201464 8578 Brachycerinae Erirhinini Tournotaris bimaculata Poland KR736283 MW201456 MW201467 8721 Molytinae Aminyopini Oreoscotus Ethiopia MG648760 MG648834 MG648746 8878 Molytinae Molytini Microplinthus emeishanicus China MG648757 MG648831 MG648743 8912 Entiminae Alophini Graptus weberi Czech Rep. MW536361 MW536409 MW536461 8915 Molytinae Paipalesomini Peribleptus Vietnam KY110615 KY110329 KY110394 8936 Molytinae Trachodini Acicnemis albofasciata Russia KY110609 KY110312 KY110374 9056 Entiminae Nastini Nastus Kazakhstan KY110618 KY110334 KY110399 9187 Molytinae Anchonini Aethiopacorep africanus Eq. Guinea MT889122 MT889144 MT889168 9190 Molytinae Anchonini Aethiopacorep africanus Eq. Guinea MT889120 MT889142 MT889166 9254 Molytinae Anchonini Eq. Guinea MT889123 MT889145 MT889169 9337 Molytinae none Tazarcus aeaea Tanzania MK813371 MK813359 MK813363 9542 Molytinae Anchonini Cameroon MT889109 MT889133 MT889155 9750 Hyperinae Hyperini Hypera Kazakhstan MW201362 MW201462 MW201475 9802 Molytinae Anchonini Acorep spinosus Guadeloupe MT889127 MT889148 MT889173 9804 Molytinae Anchonini Acorep piliger Guadeloupe MT889125 none MT889171 9806 Molytinae Anchonini Ixanchonus hustachei Guadeloupe MT889128 MT889149 MT889174 9807 Molytinae Anchonini Geobyrsa trossula Guadeloupe MT889117 MT889139 MT889163 9816 Molytinae Anchonini Leprosomus Colombia MT889107 none MT889153 (continues on the next page) Acta Entomologica Musei Nationalis Pragae, volume 62, number 2, 2022 419 Table 2. DNA fragments and their GenBank accession numbers of 153 weevil (Coleoptera) specimens used in the three marker phylogenetic analysis of the tribe Lymanini (including 137 newly sequenced fragments shown in bold: OL671058–OL671194). Voucher Subfamily Tribe Genus and/or species Country COI ITS2 28S 9817 Molytinae Lymantini Lymantes scrobicollis United States OL671066 OL671163 OL671115 9819 Molytinae Lymantini Epibaenus pinicola Mexico OL671067 OL671164 OL671116 9821 Molytinae Lymantini Theognete cozari Mexico OL671065 OL671162 OL671114 9828 Molytinae Lymantini Theognete chiapaneca Mexico OL671064 OL671161 OL671113 9829 Molytinae Lymantini Theognete galvezi Mexico OL671081 OL671174 OL671129 9831 Molytinae Lymantini Theognete montana Mexico OL671105 OL671193 OL671154 9832 Molytinae Anchonini Anchonus Mexico MT889111 MT889135 MT889157 9834 Molytinae Anchonini Anchonus Mexico MT889113 none MT889159 9934 Lixinae Lixini Bangasternus orientalis Tajikistan MW726818 MW726727 MW726908 9960 Molytinae Anchonini Anchonus blatchleyi Cuba MT889108 MT889132 MT889154 9968 Molytinae Cycloterini Dufauiella Cuba MT889130 MT889151 MT889176 9972 Dryophthorinae Stromboscerini Allaeotes niger Cuba MN621866 MN621859 MN621862 9985 Lixinae Cleonini Leucophyes pedestris Russia MW726742 MW726665 MW726832 9989 Lixinae Cleonini Pachycerus segnis Russia MW726753 MW726674 MW726843 10060 Molytinae Lymantini Mexico OL671072 none OL671121 10067 Molytinae Lymantini Mexico OL671076 none OL671125 10070 Molytinae Cycloterini Paranchonus Costa Rica MT889131 MT889152 MT889177 10071 Molytinae Anchonini Anchonus Costa Rica MT889114 none MT889160 10074 Molytinae Lymantini Dioptrophorus Mexico OL671099 OL671189 OL671148 10075 Molytinae Lymantini Mexico OL671068 OL671165 OL671117 10077 Molytinae Lymantini Mexico OL671074 OL671168 OL671123 10079 Molytinae Lymantini Mexico OL671061 OL671158 OL671110 10080 Molytinae Lymantini Dioptrophorus Mexico OL671096 OL671186 OL671144 10082 Molytinae Lymantini Dioptrophorus Mexico OL671070 OL671166 OL671119 10086 Molytinae Lymantini Lymantes Mexico OL671058 OL671156 OL671107 10089 Molytinae Lymantini Mexico OL671093 OL671184 OL671141 10092 Molytinae Lymantini Dioptrophorus Mexico OL671102 none OL671151 10093 Molytinae Lymantini Lymantes Mexico OL671092 none OL671140 10094 Molytinae Lymantini Dioptrophorus Mexico OL671075 OL671169 OL671124 10095 Molytinae Lymantini Dioptrophorus Mexico OL671073 OL671167 OL671122 10101 Molytinae Lymantini Dioptrophorus Mexico OL671059 OL671157 OL671108 10102 Molytinae Lymantini Dioptrophorus Mexico OL671083 OL671176 OL671131 10103 Molytinae Lymantini Epibaenus Mexico OL671091 OL671183 OL671139 10105 Molytinae Lymantini Dioptrophorus Mexico OL671098 OL671188 OL671147 10313 Molytinae Conotrachelini Conotrachelus United States MT889115 MT889137 MT889161 10315 Molytinae Lymantini Caecossonus Belize OL671085 OL671178 OL671133 10325 Entiminae Sitonini Sitona Canada MW201359 MW201459 MW201472 10326 Entiminae Alophini Lepidophorus lineaticollis Canada MW536368 MW536417 MW536469 10327 Entiminae Phyllobiini Evotus naso Canada MW536370 MW536419 MW536471 10329 Molytinae Lymantini Mexico OL671089 OL671181 OL671137 10330 Brachycerinae Raymondionymini Mexico MW201357 MW201458 MW201470 10331 Brachycerinae Raymondionymini Mexico MW201361 MW201461 MW201474 10334 Molytinae Lymantini Dioptrophorus Mexico OL671095 OL671185 OL671143 10335 Molytinae Lymantini Mexico OL671086 OL671179 OL671134 10338 Molytinae Lymantini Dioptrophorus Mexico OL671060 none OL671109 10339 Brachycerinae n/a Yagder serratus Mexico MW201355 MW201457 MW201468 10341 Molytinae Lymantini Dioptrophorus Mexico OL671080 OL671173 OL671128 10391 Molytinae Anchonini Titilayo barclayi S. Tome & Pr. MT889119 MT889141 MT889165 10393 Molytinae Anchonini Titilayo geiseri Guinea MT889112 MT889136 MT889158 10394 Molytinae Phrynixini New Zealand OL671069 none OL671118 10395 Molytinae Phrynixini New Zealand OL671100 OL671190 OL671149 10401 Molytinae Phrynixini New Zealand OL671090 OL671182 OL671138 10403 Molytinae Phrynixini New Zealand OL671087 OL671180 OL671135 10404 Molytinae Phrynixini New Zealand OL671084 OL671177 OL671132 10407 Curculioninae Geochini Geochus New Zealand MT889110 MT889134 MT889156 10443 Entiminae Ophryastini Deracanthus Mongolia MW536349 MW536391 MW536443 10546 Molytinae Lymantini Ithaura Costa Rica OL671079 OL671172 none 10548 Molytinae Lymantini Costa Rica OL671078 OL671171 OL671127 10557 Molytinae Lymantini Ithaura Costa Rica OL671097 none OL671145 10559 Molytinae Lymantini Costa Rica OL671103 OL671192 OL671152 10584 Molytinae Lymantini Dioptrophorus Mexico OL671077 OL671170 OL671126 10592 Molytinae Lymantini Epibaenus Mexico OL671094 none OL671142 10644 Lixinae Lixini Rhinocyllus conicus Ukraine MW726746 MW726668 MW726836 10708 Molytinae Lymantini Decuanellus Puerto Rico OL671101 OL671191 OL671150 10709 Molytinae Lymantini Decuanellus Puerto Rico OL671082 OL671175 OL671130 10732 Molytinae Lymantini Dioptrophorus Mexico OL671106 OL671194 OL671155 10733 Molytinae Lymantini Mexico OL671088 none OL671136 10747 Molytinae Lymantini Caecossonus Costa Rica OL671071 none OL671120 (continues on the next page) GREBENNIKOV & ANDERSON: Phylogeny, diversity and biogeography of Lymantini (Coleoptera: Curculionidae: Molytinae)420 dering that either Lymantini or Anchonini consistently emerged outside of the CCCMS clade (G 2018, G & A 2021a,b), we widened the outgroup by including seven representatives of non- -monophyletic Brachycerinae, a waste-basket taxon at least some members of which forming the twilight zone of “true weevils” (Curculionidae, S et al. 2017, G & A 2021b). To root the Curculionidae topology consistently with earlier results (S et al. 2017, G - & A 2021b), we used fi ve eyeless species of Brachycerinae, four of them belonging to the likely non-monophyletic tribe Raymondionymini (G & A 2021b). Altogether, 153 weevil terminals constituted the matrix (Table 2 and an online BOLD public dataset dx.doi.org/10.5883/DS-VGDS24). Three-marker Maximum Likelihood (ML) phylogene- tic analysis. The methodological approach of the analysis follows those of our recent works (G & A - 2021a,b) and, therefore, is only briefl y described. Alignment of all three DNA fragments was done separately using the online MAFFT Q-INS-i algorithm utilizing, when applicable, the secondary stricture information (K et al. 2017; https://maff t.cbrc.jp/alignment/server/). No internal parts of DNA fragments were removed before the analysis, even if consisting mainly of indels (insertions or deletions, particularly frequent in ITS2). Inconsistently sequenced 5′-end and ‎3′-ends of the ITS2 alignment were trimmed of 11 and 12 positions on each side, respecti- vely; 21 such positions were also trimmed at the 3′-end of the 28S alignment. Three aligned single-fragment datasets (Table 1) were concatenated using Mesquite 3.61 (M & M 2020) into a matrix of 4,174 positions. An unrooted topology was built using an ML approach, as implemented in CIPRES Science Gateway online platform (M et al. 2010; http://www.phylo. org/, tool “RAxML-HPC2 on XSEDE”) and using RAxML version 8 algorithm (S 2014) which applies the CAT approximation to the GTR+G nucleotide substitution model independently to each of the three partitions. Branch support values were generated based on 1000 bootstrap re- plicates (S et al. 2008) and categorized as strong (≥95%), moderate (<95% and ≥75%), or weak (<75%). The tree was visualized in FigTree v1.4.4. (R 2020). Specimen illustration and documentation. To document the inadequately known adult morphological diversity of the tribe Lymantini, a dedicated eff ort was made to illustra- te these beetles. For this purpose, 26 ingroup specimens (from 50 included in the ML analysis) were imaged in fi ve standard views (habitus dorsal, habitus left lateral, habitus left fronto-lateral, habitus ventral, antenna). Two additional specimens from two Lymantini genera lacking DNA data and not represented in the analysis were similarly illustrated: Gononotus angulicollis (Suff rian, 1871) in Fig. 34 and Kuschelaxius discifer Howden, 1992 in Fig. 35. All fi ve images of each of the 28 specimens, together with the specimen’s number, its geographic coordinates and the most detailed currently available taxonomic assignment, were arranged into 28 plates (Figs 8–35). The only Lyman- tini genus not herein illustrated (and not seen by us) is the monotypic Pseudocaecocossonus Osella, 1977 known only from two Cuban specimens (H 1992). An uncertain number of segments in antennal funic- le. During this study we concluded that determining the homology (and, therefore, the number) of antennomeres in a funicle (Fig. 5) of the subtribe Lymantina is far from straightforward. It appears likely that the club of at least some Lymantina (e.g., the genus Theognete) came to inc- lude the much enlarged distal (seventh) funicle antenno- mere. If so, this distal antennomere is likely misinterpreted as part of the club, giving the 7-segmented funicle the appearance of being 6-segmented. Presently we did not make an eff ort to clarify this uncertainty, but thoroughly documented antennal diversity throughout the tribe (Figs 5, 8–35). When giving the number of funicle segments in Lymantina, we use published numbers, which might, or might not be correct. Results The three-marker ML analysis of 153 terminals resulted in a phylogenetic tree depicted in Figs 6 and 7. The tribe Lymantini, the ingroup of the analysis, was rendered para- phyletic by the monophyletic tribe Anchonini. The internal Table 2. DNA fragments and their GenBank accession numbers of 153 weevil (Coleoptera) specimens used in the three marker phylogenetic analysis of the tribe Lymanini (including 137 newly sequenced fragments shown in bold: OL671058–OL671194). Voucher Subfamily Tribe Genus and/or species Country COI ITS2 28S 10780 Entiminae Cneorhinini Attactagenus albinus Ukraine MW536374 MW536424 MW536476 10785 Entiminae Tanymecini Tanymecus palliatus Ukraine MW536341 MW536383 MW536433 10788 Entiminae Otiorhynchini Otiorhynchus albidus Ukraine MW536379 MW536430 MW536482 10790 Entiminae Phyllobiini Phyllobius oblongus Ukraine MW536378 MW536429 MW536481 10801 Molytinae Mecysolobini Sternuchopsis South Africa MW726787 none MW726877 10804 Molytinae Mecysolobini Sternuchopsis Madagascar MW726745 none MW726835 10810 Molytinae Anchonini Cote d’Ivoire MT889116 MT889138 MT889162 10811 Molytinae Anchonini Cote d’Ivoire MT889129 MT889150 MT889175 10826 Brachycerinae Raymondionymini Alaocyba Italy MW201354 MW201455 MW201466 10827 Brachycerinae Raymondionymini Raymondiellus Italy MW201353 MW201454 MW201465 10835 Molytinae Lymantini Ithaura Nicaragua OL671104 none OL671153 10836 Molytinae Lymantini Nicaragua OL671063 OL671160 OL671112 10837 Molytinae Lymantini Nicaragua OL671062 OL671159 OL671111 10842 Molytinae Lymantini Pseudoalaocybites Guatemala none OL671187 OL671146 10853 Molytinae Mecysolobini Sternuchopsis Madagascar MW726750 MW726671 MW726840 11026 Lixinae Lixini Lixus fi liformis Ukraine MW726777 MW726695 MW726867 11046 Lixinae Lixini Lixus rubicundus Ukraine MW726793 MW726706 MW726883 Acta Entomologica Musei Nationalis Pragae, volume 62, number 2, 2022 421 Fig. 6. Maximum likelihood tree of true weevil relationships reconstructed by RAxML from the three-fragment concatenated matrix. Three subclades forming the clade of Anchonini plus Lymantini are collapsed. Large and small circles denote strongly and moderately supported clades, respectively. GREBENNIKOV & ANDERSON: Phylogeny, diversity and biogeography of Lymantini (Coleoptera: Curculionidae: Molytinae)422 relationships of this weakly supported and weakly resolved clade was Lymantina + (Anchonini + Caecossonina); the latter two taxa each strongly statistically supported clades and together uniquely characterized by a funicle with eight antennomeres. The monophyletic Asian genus Devernodes was placed inside the monophyletic subtribe Lymantina; the latter weakly supported if including the genus Decua- nellus Osella, 1977, or strongly supported, if without it. The Lymantina genera Dioptrophorus Faust, 1892 and Theognete were both recovered as strongly supported. Conversely, the genera Epibaenus Kuschel, 1959 and Lymantes were recovered as non-monophyletic. Outside of the Anchonini plus Lymantini clade, the re- maining 86 analysed terminals clustered into the following six groups, all weakly resolved among themselves. All fi ve terminals of eyeless Brachycerinae formed a weakly supported cluster, permitting straightforward rooting between them and the rest of the topology. The remaining fi ve clusters/clades were Dryophthorinae (moderately supported), the CCCMS clade (weakly supported and excluding the Anchonini plus Lymantini clade, as well as Phrynixini), eyed Brachycerinae (moderately suppor- ted), the CEGH clade (weakly supported) and Phrynixini (strongly supported). Discussion Reliability of phylogenetic tree. Excepting a few devi- ations discussed below, our ML topology (Figs 6, 7) is remarkably consistent with the existing ideas on weevil phylogeny based on a much larger set of DNA data (e.g., S et al. 2017; references therein). Specifi cally, we reco- vered the following well-established clades, some of them with moderate or strong statistical support: Dryophthori- nae, CEGH clade, CCCMS clade (excluding, however, Phrynixini, Anchonini and Lymantini; see below), Phry- nixini and Anchonini. This consistency between our results and those of earlier studies suggest that our topology is a credible source of phylogenetic interpretations (see below). Non-monophyletic Lymantini form a clade with mo- nophyletic Anchonini. Perhaps the most signifi cant phy- logenetic result of our analysis is that the tribe Lymantini, the ingroup of this study, emerged paraphyletic to the tribe Anchonini. Although weakly statistically supported, this result lends credence to the century-old “Anchonina” of C (1902, 1903) and appears sound in light of at least four other lines of evidence. Firstly, both tribes have been already linked into a moderately supported clade in our recent Anchonini-focused analysis (G & A 2021a). There, however, the tribe Lymantini was represented by a single species of the genus Deverno- des, then a questionable member of the latter tribe (but see below). Secondly, in the same study, we hypothesized that the clade of Anchonini plus Lymantini might have at least one morphological apomorphy, the polished head capsule of these beetles; a supposition corroborated in the present analysis. Thirdly, larvae of Anchonini and Lymantini are remarkably similar (A 1952). Fourthly, available biogeographic interpretations for amphi-Atlantic mono- phyletic Anchonini (G & A 2021a) and amphi-Pacific non-monophyletic Lymantini (see below) suggest that their most recent common ancestor (MRCA) likely inhabited the North American continent before the Eocene (see below). If Anchonini and Lymantini indeed share a MRCA, as all available data consistently suggest, its exact age, geographic localization, and the identity of its sister group are three main unknowns yet to be elucidated. Summing up, in the current absence of alter- natives, the monophyly of Anchonini and Lymantini is the only existing hypothesis that, although weakly statistically supported, agrees with all available evidence. The Mesoamerican Lymantini subtribe Caecossonina is sister to amphi-Atlantic Anchonini. The monophyly of the Lymantini subtribe Caecossonina plus the tribe An- chonini is statistically weakly supported, although likely credible. Two independent lines of evidence support this conclusion. Firstly, both analyzed genera of Caecossonina (and by extension its third and the last genus, Pseudocae- cocossonus) likely form a clade supported by at least two morphological characters: lack of eyes and small adult bodies not exceeding 3 mm in length. Secondly, all members of the subtribe Caecossonina diff er from those of the subtribe Lymantina by sharing with Anchonini a rare morphological trait: the 8-segmented antennal funicle (Figs 4, 5, 20, 33). Moreover, the MRCA of American Caecossonina and amphi-Atlantic Anchonini, if it has existed, likely inhabited the North American continent (at that time widely separated by the sea from insular South America) not later than the eastwards transatlantic dispersal of Anchonini to West Africa some 9.5–5.2 million years ago (G & A 2021a; see below). The amphi-Pacifi c subtribe Lymantina with its MRCA likely living in North America (see below) is herein considered as sister to the Caecossonina plus Anchonini clade. Summing up, in the current absence of alternatives, sister relations of the Lymantini subtribe Caecossonina and the tribe An- chonini is the only existing hypothesis which, although weakly statistically supported, is in agreement with all available evidence. The Anchonini plus Lymantini clade is outside of the CCCMS clade. The current taxonomic assignment of both Anchonini and Lymantini in the subfamily Molytinae (A -Z & L 1999, L 2014) implies that both tribes are phylogenetically nested within the CCCMS clade of “higher” weevils (e.g., S et al. 2017). Our results, however, indicate that the clade of Anchonini plus Lymantini is outside of the CCCMS clade; the latter having moderate statistical support (Fig. 6). This result is consistent with our earlier analyses which resolved these tribes outside of the CCCMS clade (e.g., G 2018, G & A 2021a). These analyses, however, used the subset of the herein analyzed dataset and the same analytical methods, which might make them similarly biased. In the present lack of other evidence, two alternatives best explain the observed discrepancy between taxonomy-based expectations and our topolo- gies. One alternative is that the taxonomic interpretation is correct, and its inconsistency with the topology is the result of analytical shortcomings, such as the scarcity Acta Entomologica Musei Nationalis Pragae, volume 62, number 2, 2022 423 Fig. 7. Maximum likelihood tree of Anchonini and Lymantini relationships reconstructed by RAxML from the three-fragment concatenated matrix. Cla- des outside of the Anchonini plus Lymantini clade are collapsed. Large and small circles denote strongly and moderately supported clades, respectively. Arrows indicate 26 specimens shown in Figs 1, 2, 8–33. Superimposed globes indicate the current distribution. GREBENNIKOV & ANDERSON: Phylogeny, diversity and biogeography of Lymantini (Coleoptera: Curculionidae: Molytinae)424 of the phylogenetic signal extracted from our dataset. Another alternative is that the subfamily Molytinae, which is known to be non-monophyletic (e.g., S et al. 2017), might artifi cially unite grossly unrelated organisms, some of them perhaps even nested outside the CCCMS clade. Phrynixini, an obscure Gondwanan tribe, is outside of the CCCMS clade. Similarly, with the Anchonini plus Lymantini clade, all fi ve herein analyzed members of the tribe Phrynixini formed a strongly supported clade placed outside of the CCCMS clade (Fig. 6). Phrynixini is a phylogenetically neglected group of some 35 genera taxonomically assigned to Molytinae (L 2014). P et al. (2014) did not assign Phrynixini to any subfamily when providing a catalog of Australian weevils but sug- gested that the tribe belongs to the CEGH clade. G et al. (2016) included two Australian genera of this tribe in a molecular phylogenetic analysis; these genera resolving in separate clades within the CEGH clade. We corrobo- rate the molecular results of L et al. (2022) who recently recovered a monophyletic Phrynixini outside of the CCCMS clade. K (1987) noted that Phrynixini have the “Gondwanan” distribution, being found in New Zealand, Australia, New Caledonia and Chile. If the plate tectonics was the factor behind the Phrynixini distribution, then the age of this clade might be comparable with the time of the Gondwana breakup and, therefore, be at least twice greater than the age of the CCCMS crown group; the later originating about 75 million years ago (S et al. 2017). High altitude inter-continental dispersal of thermo- philic Lymantina across Arctic land bridges. A new, strongly supported and evolutionary signifi cant result of our analysis is the recovery of the recently described Asian genus Devernodes nested within the otherwise exclusively American subtribe Lymantina (Fig. 7). This corroborates the earlier morphology-based assumption of Devernodes relationships with Lymantini (G 2018) made, however, without the benefi t of a formal analysis. This re- sult also means that the stenotopic, thermophilic, fl ightless and presumably low-dispersing monophyletic subtribe Lymantina is found in two widely separated unglaciated warm regions of the World: in Southeastern Asia (the genus Devernodes) and the tropical Americas (the rest of the subtribe; Fig. 3). Below we off er a biogeographic interpretation of this distribution. Sister-relationships between Asian and American extant animals is not an infrequent phenomenon. It is perhaps best known for tapirs (Tapiridae), a clade of large, herbivorous, odd-toed mammals similar in appearance to pigs with a short, prehensile trunk. Malayan tapir, Tapirus indicus Desmarest, 1819, inhabits Southeast Asia, while the re- maining three or four extant congeners are found between Mexico and Argentina (C et al. 2013). Arthropod examples are numerous (e.g., A 1983) and include, among others, Penichrolucaninae stag beetles (R 1984) and freshwater water fl eas Leydigiopsis Sars, 1901 (Cladocera: Anomopoda: Chydoridae; V D & S - 2013). To account for the intercontinental distribution of terrestrial animals, risky and low-probability long-dis- tance transoceanic dispersals are occasionally justifi ably evoked ( Q 2014). Examples include the likely out-of-America Cretaceous single dispersal event of the opilionid family Zalmoxidae, leading to their spectacular radiation in Southeast Asia and Australia (S & G - 2012). Another transoceanic example of dispersal involves anchonine weevils, the clade likely rendering Lymantini paraphyletic (Fig. 7). These fl ightless beetles have been shown to disperse overwater (C & L 2018) in the later Miocene (G & A 2021a) across the Atlantic from the Americas to West Africa. Might then the overwater dispersal be the likeliest hypothesis for the present-day amphi-Pacifi c distribution of the East Asian genus Devernodes and its American Lymantina relatives? The answer is likely “no”. Our main analytical limita- tion is the lack of Lymantina fossils to determine the past distribution of the clade and to date our topology (Fig. 7). Still, the amphi-Pacifi c distribution of this monophyletic subtribe can be plausibly explained without evoking long- distance chance dispersal. All available data consistently suggest that the current disjunct distribution of Lymantina in both Asia and the Americas is most likely a result of high latitude inter-continental normal ecological dispersal (H 2014). It implies gradual overland dispersal of thermophilic Lymantina across Arctic land bridges during the warmest periods of the Cenozoic, e.g., the Paleocene– Eocene Thermal Maximum some 56 Ma (M I & W 2011). We assume, therefore, those fl ightless lymantine weevils have dispersed overland between their current areas of distribution in North America and Asia using the currently submerged North Atlantic and/or the Beringia land bridges (e.g., the De Geer, Thulean, or Be- ringia; B 2014). This gradual dispersal likely took place before the Eocene-Oligocene boundary some 33.5 Ma, when the warm global “greenhouse” climate turned to that of an “icehouse” (E et al. 2009). This cli- matic event is considered to have triggered the decline and disappearance of the Boreotropical fl ora (W 1975). This was a belt of thermophilic vegetation in the Northern Hemisphere during the Eocene epoch reaching as far north as 80°N in which these weevils may have thrived. Last but not least, the Eocene timing appears consistent with the phylogenetic position of Devernodes nested deeply within Lymantina (Fig. 7) and suffi ciently long (as opposite to e.g., Pliocene-Pleistocene timing) to account for the sizable morphological distinctness of this Asian genus from the American rest of the subtribe. Our assumption of climate-mediated vicariance between American and Asian Lymantina is consistent with hypo- theses evoked for other similarly distributed clades of terrestrial thermophilic animals, such as lizards (S 2011), or extinct giant ants (A et al. 2011); for a review on the Eocene fl ora and vertebrate fauna see E & G (2012). At that time warm-loving non-vo- lant terrestrial animals such as stem-group tapirs (E & E 2015) and camels (R et al. 2013) inhabited what is presently Ellesmere Island, Canada’s northernmost island lying within the Arctic Archipelago. Beetle examples of such vicariance include the giant Acta Entomologica Musei Nationalis Pragae, volume 62, number 2, 2022 425 Callipogon Audinet-Serville, 1832 longhorns (K et al. 2018), Bolitogyrus Chevrolat, 1842 rove beetles (B et al. 2017) and Megasternini terrestrial water scavenger beetles (A -V et al. 2021). This assumption is also consistent with the likely relictual presence of the genus Lymantes in the northwestern USA (Fig. 3). These coastal populations of Lymantina are widely isolated from the more southwards rest of the subtribe’s Ameri- can distribution, occupy areas that were unglaciated and wetter during at least the Last Glacial Maximum (L et al. 2019), and likely represent a remnant of the former much wider Lymantina presence in the American North. Directionality of the trans-Arctic Lymantina dispersal remains unknown, although a single overland migration event of the stem Devernodes from North America to Asia appears most plausible. We conclude, therefore that the disjunct presence of Lymantina in the Americas and Asia is a result of normal ecological dispersal fi rst creating an uninterrupted Holarctic distribution of this clade, with subsequent climatic cooling obliterating these cold-into- lerant beetles between the widely disjunct areas of their recent distribution. Our hypothesis on the normal overland ecological dispersal of Lymantina across arctic land bridges implies the presence of Lymantina fossils in the presently Lymanti- na-free temperate and arctic regions along the hypothesized dispersal routes (Fig. 3). Examples of such coveted fossil discoveries for other terrestrial animals include the Eoce- ne Bolitogyrus Chevrolat, 1842 rove beetles found in the Baltic region of Europe and from Green River formation in Colorado, USA (B et al. 2017) and the early Oligocene bones of legless Dibamidae burrowing blind skinks found in the presently Dibamidae-free Mongolia (Č 2019). The discovery of such a Lymantina fossil from these intervening Lymantina-free regions would, therefore, considerably strengthen our hypothesis. North American origin of Lymantina crown group and that of Anchonini plus Lymantini. It is tempting to speculate which of the three continents currently inhabited by the crown group Lymantina, if any, i.e., Asia, North America and South America, has supported the clade’s MRCA. Numerical preponderance of recent Lymantina in Mesoamerica, and their corresponding scarcity in Asia are not informative in this respect. Clades of non-volant terrestrial animals with exceptionally well-documented fossil records are known to have their MRCA on one continent, disperse to others, and then become extinct in the continent of their origin. Examples include crown group camels originating in the Eocene of North Ameri- ca, dispersing to Eurasia across the Arctic land bridges (and then to Africa) and also to South America across the newly formed Isthmus of Panama, and becoming extinct in North America (H et al. 2015). Even though our analysis is inconclusive on this point, a North American origin and subsequent dispersal to Asia appears to be the most parsimonious explanation for Lymantina, particularly in light of the American origin of their sister group, Anchonini (G & A 2021a) plus Caecossonina (Fig. 7). Finally, assuming that (1) MRCA of Lymantini inhabited North America likely in Eocene time to permit overland dispersal of the stem Devernodes to Asia and (2) assuming existence of the Lymantina + (Caecossonina + Anchonini) clade, on which continent did the MRCA of this clade live? If restricting our choice to either North (including Central America) or South America, as it is most parsimonious options considering all available evidence, North America is by far the likeliest candidate. The choice is pivoted on a consideration that South America, being for most of its geological and biotic history widely separated from other landmasses, is highly unlikely to have any of its native organisms reaching across the sea to North America, to account for the herein hypothesized Lymantina dispersal event between North America and Asia. Formation of the Isthmus of Panama (and corresponding closure of the Cen- tral American Seaway separating both Americas), which would be needed to permit the South American origin of the Lymantina + (Caecossonina + Anchonini) clade, is a hotly debated subject (e.g., W 2010), with dates varying widely, depending on the evidence used. Either way, geological evidence suggesting the earlier date (the mid-Miocene, M et al. 2015) or biological evidence suggesting a much later date (3 Ma, O’D et al. 2016) of the Great American Biotic Interchange (GABI) both greatly postdate the time when members of Lymantina have likely made their way overland from North America to Asia. Thus, if the logic above is sound and the assumptions correct, the crown group of the Anchonini plus Lymantini clade originated in North America. It follows that the Late Miocene eastwards transatlantic overseas dispersal of Anchonini to West Africa (G & A 2021a) took place from North America, and not from South America; the latter at that time likely still surrounded by the sea and uninhabited by these beetles. It also follows, that the current presence of Anchonini and Lymantini in South America is yet another example the Great American Biotic Interchange, a fascinating phenomenon exceptio- nally well-documented for vertebrates, while with only a few examples among beetles (e.g., T et al. 2021 on a dung beetle subfossil; Ż et al. 2021 on paederine rove beetles). Unsatisfying taxonomy of the tribe Lymantini. The taxonomy of Lymantini weevils is unsatisfactory for two reasons. Firstly, if our phylogenetic interpretation of Ly- mantini is correct, the tribe is paraphyletic with respect to Anchonini (Fig. 7). To address this inconsistency, the younger name Lymantini Lacordaire, 1865 might be synonymized under Anchonini Imhoff , 1856, to return to the concept of “Anchonina” of C (1902, 1903). This will result in the larger monophyletic tribe Anchonini containing three monophyletic subtribes: Anchonina, Cae- cossonina and Lymantina. Alternatively, the Lymantini sub- tribe Caecossonina might be elevated to the tribe level, to create three monophyletic tribes: Anchonini, Caecossonini and Lymantini. As there are additional tribes in Molytinae other than these three, the latter taxonomic solution will fail to imply that they form a clade and is, therefore, less preferable. Lacking suffi cient statistical confi dence in the GREBENNIKOV & ANDERSON: Phylogeny, diversity and biogeography of Lymantini (Coleoptera: Curculionidae: Molytinae)426 Fig. 8. Sequenced Lymantini specimen 4339: Devernodes chthonia Grebennikov, 2018. Fig. 9. Sequenced Lymantini specimen 9817: Lymantes scrobicollis Gyllenhal, 1838. Acta Entomologica Musei Nationalis Pragae, volume 62, number 2, 2022 427 Fig. 10. Sequenced Lymantini specimen 9819: Epibaenus pinicola Kuschel, 1959. Fig. 11. Sequenced Lymantini specimen 9821: Theognete cozari Anderson, 2010. GREBENNIKOV & ANDERSON: Phylogeny, diversity and biogeography of Lymantini (Coleoptera: Curculionidae: Molytinae)428 Fig. 12. Sequenced Lymantini specimen 9829: Theognete galvezi Anderson, 2010. Fig. 13. Sequenced Lymantini specimen 10060: Lymantina. Acta Entomologica Musei Nationalis Pragae, volume 62, number 2, 2022 429 Fig. 14. Sequenced Lymantini specimen 10067: Lymantina. Fig. 15. Sequenced Lymantini specimen 10079: Lymantina. GREBENNIKOV & ANDERSON: Phylogeny, diversity and biogeography of Lymantini (Coleoptera: Curculionidae: Molytinae)430 Fig. 16. Sequenced Lymantini specimen 10080: Dioptrophorus sp. Fig. 17. Sequenced Lymantini specimen 10089: Lymantina. Acta Entomologica Musei Nationalis Pragae, volume 62, number 2, 2022 431 Fig. 18. Sequenced Lymantini specimen 10093: Lymantes sp. Fig. 19. Sequenced Lymantini specimen 10105: Dioptrophorus sp. GREBENNIKOV & ANDERSON: Phylogeny, diversity and biogeography of Lymantini (Coleoptera: Curculionidae: Molytinae)432 Fig. 20. Sequenced Lymantini specimen 10315: Caecossonus sp. Fig. 21. Sequenced Lymantini specimen 10334: Dioptrophorus sp. Acta Entomologica Musei Nationalis Pragae, volume 62, number 2, 2022 433 Fig. 22. Sequenced Lymantini specimen 10335: Lymantina. Fig. 23. Sequenced Lymantini specimen 10546: Ithaura sp. GREBENNIKOV & ANDERSON: Phylogeny, diversity and biogeography of Lymantini (Coleoptera: Curculionidae: Molytinae)434 Fig. 24. Sequenced Lymantini specimen 10548: Lymantina. Fig. 25. Sequenced Lymantini specimen 10557: Ithaura sp. Acta Entomologica Musei Nationalis Pragae, volume 62, number 2, 2022 435 Fig. 26. Sequenced Lymantini specimen 10559: Lymantina. Fig. 27. Sequenced Lymantini specimen 10592: Epibaenus sp. GREBENNIKOV & ANDERSON: Phylogeny, diversity and biogeography of Lymantini (Coleoptera: Curculionidae: Molytinae)436 Fig. 28. Sequenced Lymantini specimen 10708: Decuanellus sp. Fig. 29. Sequenced Lymantini specimen 10732: Dioptrophorus sp. Acta Entomologica Musei Nationalis Pragae, volume 62, number 2, 2022 437 Fig. 30. Sequenced Lymantini specimen 10733: Lymantina. Fig. 31. Sequenced Lymantini specimen 10835: Ithaura sp. GREBENNIKOV & ANDERSON: Phylogeny, diversity and biogeography of Lymantini (Coleoptera: Curculionidae: Molytinae)438 Fig. 32. Sequenced Lymantini specimen 10836: Lymantina. Fig. 33. Sequenced Lymantini specimen 10842: Pseudoalaocybites sp. Acta Entomologica Musei Nationalis Pragae, volume 62, number 2, 2022 439 Fig. 34. Not sequenced Lymantini specimen of Gononotus angulicollis (Suff rian, 1871). Fig. 35. Not sequenced Lymantini specimen of Kuschelaxius discifer Howden, 1992. GREBENNIKOV & ANDERSON: Phylogeny, diversity and biogeography of Lymantini (Coleoptera: Curculionidae: Molytinae)440 phylogenetic affi nities of these beetles precludes us from introducing these taxonomic changes. Secondly, the subtribe Lymantina likely contains non- monophyletic genera (e.g., Lymantes and Epibaenus), as well as unnamed species not currently attributable to any genus, monophyletic or not (e.g., all specimens identifi ed as “Lymantina” in Figs 1 and 2). Below we list all 12 valid genera of the tribe Lymantini and provide information on their diversity and distribution. Tribe Lymantini Lacordaire, 1865 Subtribe Caecossonina Osella, 1980 Caecossonus Gilbert, 1955: four species in Belize, Cuba, Mexico and USA (Florida). Pseudoalaocybites Osella, 1980: 16 species in Colombia, Cuba, Jamaica, Ecuador and Venezuela (G 2020). Pseudocaecocossonus Osella, 1977: monotypic, Cuba (H 1992). Subtribe Lymantina Lacordaire, 1865 Decuanellus Osella, 1977: 12 species inhabiting Caribbean islands between the Bahamas and St. Lucia (C - 2010; R & V D 2021). Devernodes Grebennikov, 2018: fi ve species in China (Sichuan), Malaysia and Vietnam (G 2018). Dioptrophorus Faust, 1892: seven species in Guatemala and Mexico (O’B & W 1982). The Cuban generic record by C (1902: 92, followed by O’B & W 1982) refers to the type species of the genus Gononotus. Epibaenus Kuschel, 1959: two species in Guatemala and Mexico (K 1959). Gononotus LeConte, 1876: monotypic, USA (Florida), Mexico, Cuba, Puerto Rico (O’B & W 1982). This genus has been recently added to the subtribe (L 2014), not available for our analysis, and its phyloge- netic position is the least known. Ithaura Pascoe, 1871: at least six species in Central Ameri- ca and the northern half of South America (R 2006). This is the only genus of the subtribe found in South America, and as far south as Bolivia and central Brazil (Fig. 3). Kuschelaxius Howden, 1992: two species in the Dominican Republic and Puerto Rico (H 1992). Lymantes Schoenherr, 1838: seven species in the USA and El Salvador (A 2016, 2022). Theognete Champion, 1902: 94 species in Mexico, Hondu- ras, Guatemala and El Salvador (A 2010). Alleged Lymantini fossil from Dominican amber. P - & L (2021) established a new extinct genus and species, Bronchotibia adunatus Poinar & Legalov, 2021, based on a Dominican amber adult weevil inclusi- on. They decisively attributed this taxon to the subfamily Molytinae and to the tribe Lymantini, and less decisively to the subtribe Lymantina, by listing morphological simi- larities. These authors pivoted their Lymantini attribution of the fossil on “... eyes ... basally located on rostral part of head ...”; a diagnostic character of non-monophyletic Lymantini. Illustrations of the fossil, however, do not corroborate this morphological interpretation. As illustra- ted on their Figure 2, the placement of the eyes is clearly on the head and dissimilar to that in Lymantini. Three ad- ditional morphological characters, namely (1) rectangular shape of the well-developed elytral shoulders suggesting presence of hind wings and capacity of active fl ight, (2) opisthognathous (rather than prognathous) orientation of the rostrum, and (3) strongly bilobed and likely adhesive tarsomeres 3 suggesting plant climbing behaviour are notably unlike anything known among the members of the Anchonini plus Lymantini clade, which are fl ightless and ground-dwelling beetles. P & L (2021) also did not mention the well-established opinion that the subfamily Molytinae is not monophyletic (e.g., S et al. 2017) and, therefore, meaningless in the phylogenetic sense. We, therefore, conclude that attribution to the extinct weevil genus Bronchotibia to either Molytinae, Lymantini, or Lymantina is unwarranted and use the criteria of C et al. (2019: 31) to re-classify this fossil as Curculionidae incertae sedis. Acknowledgements Through their work on leaf litter ants, John T. 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