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Checklist of the micro- and anophthalmic soil-dwelling weevils of the world (Coleoptera: Curculionidae)

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A checklist of the soil-dwelling microphthalmic and anophthalmic weevils of the world is presented. A total of 485 species in 85 genera are recognised, distributed over eight subfamilies and 26 tribes. The subfamilies are arranged in the currently accepted phylogenetic sequence, whereas the tribes, genera and species are listed in alphabetical order. For each species, the type locality and the geographical distribution by country is provided. Particularly diversified taxa are Entiminae: Otiorhynchini (125 species), Brachycerinae: Raymondionymini (90 species), Cossoninae: Dryotribini (59 species), Cryptorhynchinae: Torneumatini (58 species) and Molytinae: Lymantini (39 species). The biogeographic region with the highest diversity of soil-dwelling weevils is the Palearctic, especially the circum-Mediterranean area, mostly because of the exploration effort that it has received, followed by the Ethiopian and Neotropical regions.
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... These sampling difficulties in obtaining deep soil beetles, which are often known only by the typical series or from only the type localities widely scattered across the Globe (Morrone and Hlaváč 2017), has also limited the extend of our current study. Our target is the lineage putatively formed by the European/Mediterranean representatives of the tribe Raymondionymini. ...
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The tribe Raymondionymini has long been neglected in phylogenetic studies. The tribe is characterized by uncertain monophyly, fluctuating taxonomic status, and a composition prone to instability. All raymondionymine weevils are wingless and have eyes either completely absent or, rarely, consisting of a single ommatidium. With body lengths predominantly below three millimeters, they inhabit deep soil environments and are infrequently collected. The core of this tribe comprises nine genera distributed in Europe and around the Mediterranean region and encompassing 76 species, while six additional genera include 17 species distributed in USA (California), Mexico, Ecuador, Venezuela, Russian Far East, and Madagascar. Here, we present eight new mitogenomes, complemented by one publicly available, encompassing all but two Mediterranean genera of raymondionymine weevils. We used publicly available Curculionoidea mitogenomes to compile an all-inclusive dataset with 391 terminals and a reduced dataset with 61 terminals representing main families of Curculionoidea and subfamilies within Curculionidae. Our maximum likelihood and Bayesian phylogenetic analyses, employing both DNA and amino acids datasets under alternative partition schemes, consistently produced congruent phylogenies. Our results show that the Mediterranean raymondionymines form a strongly supported clade, and their easternmost and morphologically distinct genus Ubychia is sister to the rest of them. Most notably, our results consistently recover a sister relationship between the clade of Mediterranean raymondionymine weevils and a clade encompassing all remaining Curculionidae. Consequently, we propose a revision of weevil taxonomy: (i) Our target group is removed from the non-monophyletic subfamily Brachycerinae; (ii) this clade is resurrected to its former subfamily level within Curculionidae, as the subfamily Raymondionyminaestat. rev; (iii) the nine Mediterranean genera Alaocephala, Alaocyba, Coiffaitiella, Derosasius, Ferreria, Raymondiellus, Raymondionymus, Tarattostichus, and Ubychia compose Raymondionyminaestat. rev; (iv) and non-Mediterranean genera Alaocybites, Bordoniola, Gilbertiola, Homosomus, Neoubychia, and Schizomicrus are considered as “incertae sedis” pending further phylogenetic corroboration. We hypothesize that the remaining Brachycerinae and the non-Mediterranean representatives within Raymondionyminae constitute a series of species-poor early-diverging lineages representing currently unrecognized subfamilies of Curculionidae.
... To date, the only subterranean cossonine weevils recorded from Madagascar are Pentebathmus insularis Richard and P. ovalis Richard (Dryotribini) (Remillet 1973;Morrone and Hlavac 2017). Here we describe the third known species of Halorhynchus and the first for the African region. ...
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Halorhynchus remii Chamorro & Steiner, sp. nov. is described from Madagascar. This new species is the third known species of the genus and the first for Africa. Halorhynchus remii is compared to other psammophilous, anophthalmous onycholipine cossonines. Transoceanic dispersal between Australia and Madagascar and sand burrowing adaptation are briefly discussed. A key to the species is provided.
... It is well known for species inhabiting caves or soil that the development of characters (e.g., longer appendices and overdevelopment of some brain areas) for life in the darkness selects against traits that are not useful in these types of habitats (e.g., cuticle pigmentation, eyes and brain associated areas). Troglobiomorphism, either intermediate or extreme, is common in several subfamilies of the Curculionidae, namely, Brachycerinae, Molytinae and Cossoninae [14]. ...
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Plumolepilius Barrios-Izás & Anderson, 2016 is a leaf litter weevil genus that inhabits montane broadleaf forests from southern Mexico to northern Panama. The genus consists of 27 species, 22 distributed in the Chiapas Highlands province (Mexican Transition Zone) and 5 found in the Pacific dominion (Neotropical region) in Costa Rica and Panamá. Here, we analyze the phylogenetic relationships of the species of Plumolepilius based on 20 external body characters and 9 characters from the genitalia. The first dichotomy of the cladogram separates two species from the Pacific dominion from the remaining species of Plumolepilius from the Chiapas Highlands province and three species restricted to the Pacific dominion. We hypothesize that redundant distributions in the taxon-area cladogram of the genus may be due to dispersal events, probably during the Pleistocene glaciations.
... The benefits are usually identified as more stable conditions, access to new resources, reduced competition and predation, and possibly also decreased threats of parasites or parasitoids (e.g., Giller, 1996). Of all insects, beetles seem to predominate in such unusual habitats, with many blind, wingless, miniaturized and often depigmented species known mainly among rove beetles, ground beetles, or weevils (e.g., Faille et al., 2013;Fancello et al., 2009;Grebennikov et al., 2009;Hlav ač et al., 2006Hlav ač et al., , 2008Hlav ač et al., , 2017Jeannel, 1957;Morrone et al., 2001;Morrone & Hlav ač, 2017;Osella, 1977). ...
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The pselaphine Bergrothia saulcyi shows features seemingly linked with life in deep soil layers, such as greatly reduced and non-functional compound eyes, a sensorium of long tactile setae, long appendages, and flightlessness. However, the tiny beetles occur in forest leaf litter, together with a community of beetles with wings and well-developed eyes. We hypothesize that B. saulcyi moves into deep soil under dry conditions, and returns to upper layers when humidity increases again. Despite the evolutionary cost of a reduced dispersal capacity, this life strategy may be more efficient and less hazardous than moving to different habitats using flight and the visual sense in an environment periodically drying out. We also discuss cephalic features with potential phylogenetic relevance. Plesiomorphies of B. saulcyi include the presence of anterior tentorial arms, well-developed labral retractors, and a full set of extrinsic maxillary and premental muscles. Apomorphic cephalic features support cla-des Protopselaphinae + Pselaphinae, and Pselaphinae. A conspicuous derived condition , the clypeo-ocular carina, is a possible synapomorphy of Batrisitae and genera assigned to Goniaceritae. A complex triple set of cephalic glands found in B. saulcyi is similar to a complex identified in the strict myrmecophile Claviger testaceus (Clavigeritae). It is conceivable that glands linked with food uptake in free-living pselaphines were genetically reprogrammed in ancestors of inquilines, to enable them to appease the host ants. We suggest that behavioral studies are necessary to understand the poorly known life habits of B. saulcyi. Additional information is required to explain why a species with irreversibly reduced visual sense and other adaptations typical of endogean or cave-dwelling beetles was only collected from the upper leaf litter layer. K E Y W O R D S 3D reconstruction, micro/anophthalmy, micro-CT, musculature, rove beetles
... The benefits are usually identified as more stable conditions, access to new resources, reduced competition and predation, and possibly also decreased threats of parasites or parasitoids (e.g., Giller, 1996). Of all insects, beetles seem to predominate in such unusual habitats, with many blind, wingless, miniaturized and often depigmented species known mainly among rove beetles, ground beetles, or weevils (e.g., Faille et al., 2013;Fancello et al., 2009;Grebennikov et al., 2009;Hlav ač et al., 2006Hlav ač et al., , 2008Hlav ač et al., , 2017Jeannel, 1957;Morrone et al., 2001;Morrone & Hlav ač, 2017;Osella, 1977). ...
Article
The pselaphine Bergrothia saulcyi shows features seemingly linked with life in deep soil layers, such as greatly reduced and non-functional compound eyes, a sensorium of long tactile setae, long appendages, and flightlessness. However, the tiny beetles occur in forest leaf litter, together with a community of beetles with wings and well-developed eyes. We hypothesize that B. saulcyi moves into deep soil under dry conditions, and returns to upper layers when humidity increases again. Despite the evolutionary cost of a reduced dispersal capacity, this life strategy may be more efficient and less hazardous than moving to different habitats using flight and the visual sense in an environment periodically drying out. We also discuss cephalic features with potential phylogenetic relevance. Plesiomorphies of B. saulcyi include the presence of anterior tentorial arms, well-developed labral retractors, and a full set of extrinsic maxillary and premental muscles. Apomorphic cephalic features support cla-des Protopselaphinae + Pselaphinae, and Pselaphinae. A conspicuous derived condition , the clypeo-ocular carina, is a possible synapomorphy of Batrisitae and genera assigned to Goniaceritae. A complex triple set of cephalic glands found in B. saulcyi is similar to a complex identified in the strict myrmecophile Claviger testaceus (Clavigeritae). It is conceivable that glands linked with food uptake in free-living pselaphines were genetically reprogrammed in ancestors of inquilines, to enable them to appease the host ants. We suggest that behavioral studies are necessary to understand the poorly known life habits of B. saulcyi. Additional information is required to explain why a species with irreversibly reduced visual sense and other adaptations typical of endogean or cave-dwelling beetles was only collected from the upper leaf litter layer. K E Y W O R D S 3D reconstruction, micro/anophthalmy, micro-CT, musculature, rove beetles
... The benefits are usually identified as more stable conditions, access to new resources, reduced competition and predation, and possibly also decreased threats of parasites or parasitoids (e.g., Giller, 1996). Of all insects, beetles seem to predominate in such unusual habitats, with many blind, wingless, miniaturized and often depigmented species known mainly among rove beetles, ground beetles, or weevils (e.g., Faille et al., 2013;Fancello et al., 2009;Grebennikov et al., 2009;Hlav ač et al., 2006Hlav ač et al., , 2008Hlav ač et al., , 2017Jeannel, 1957;Morrone et al., 2001;Morrone & Hlav ač, 2017;Osella, 1977). ...
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The pselaphine Bergrothia saulcyi shows features seemingly linked with life in deep soil layers, such as greatly reduced and non‐functional compound eyes, a sensorium of long tactile setae, long appendages, and flightlessness. However, the tiny beetles occur in forest leaf litter, together with a community of beetles with wings and well‐developed eyes. We hypothesize that B. saulcyi moves into deep soil under dry conditions, and returns to upper layers when humidity increases again. Despite the evolutionary cost of a reduced dispersal capacity, this life strategy may be more efficient and less hazardous than moving to different habitats using flight and the visual sense in an environment periodically drying out. We also discuss cephalic features with potential phylogenetic relevance. Plesiomorphies of B. saulcyi include the presence of anterior tentorial arms, well‐developed labral retractors, and a full set of extrinsic maxillary and premental muscles. Apomorphic cephalic features support clades Protopselaphinae + Pselaphinae, and Pselaphinae. A conspicuous derived condition, the clypeo‐ocular carina, is a possible synapomorphy of Batrisitae and genera assigned to Goniaceritae. A complex triple set of cephalic glands found in B. saulcyi is similar to a complex identified in the strict myrmecophile Claviger testaceus (Clavigeritae). It is conceivable that glands linked with food uptake in free‐living pselaphines were genetically re‐programmed in ancestors of inquilines, to enable them to appease the host ants. We suggest that behavioral studies are necessary to understand the poorly known life habits of B. saulcyi. Additional information is required to explain why a species with irreversibly reduced visual sense and other adaptations typical of endogean or cave‐dwelling beetles was only collected from the upper leaf litter layer. This article is protected by copyright. All rights reserved.
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An annotated catalogue of Coleoptera Curculionoidea of “Monte Grappa” Biosphere Reserve is presented. The 528 species surveyed, divided into 7 families and 283 genera and subgenera, include the 265 species already known, plus the 263 are reported for the first time for the area. Faunistic, chorological and ecological information is given for all the species together with 1.015 illustrations of distinctive species, collecting techniques and selected habitats. In addition the database of 20.000 specimens collected can be found at: doi.org/10.5281/zenodo.7497691
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We describe and illustrate a new eyeless weevil, Yagder serratus gen. & sp. nov., based on a single adult female collected by sifting forest leaf litter in Mexico. A phylogenetic analysis of 39 terminals and 2679 aligned positions from three DNA fragments places the new species into the subfamily Brachycerinae (as incertae sedis) and outside the highly diversified clade of ‘higher’ true weevils. Neither Brachycerinae, nor its tribe Raymondionymini traditionally uniting most eyeless weevils, are monophyletic unless the latter is limited to a Mediterranean core group. Both these taxa are taxonomic dumping-grounds likely containing species-poor sisters of species-rich clades. When resolved, the subfamily Brachycerinae will be likely split into two or more species-poor deeply-divergent subfamilies.
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