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Article ; Online: The thoracic morphology of the troglobiontic cholevine species Troglocharinus ferreri (Coleoptera, Leiodidae).

Luo, Xiao-Zhu / Antunes-Carvalho, Caio / Ribera, Ignacio / Beutel, Rolf Georg

2019 Volume 53, Page(s) 100900

Abstract: The thoracic morphology of the troglobiontic leiodid species Troglocharinus ferreri (Cholevinae, Leptodirini) is described and documented in detail. The features are mainly discussed with respect to modifications linked with subterranean habits. ... ...

Abstract	The thoracic morphology of the troglobiontic leiodid species Troglocharinus ferreri (Cholevinae, Leptodirini) is described and documented in detail. The features are mainly discussed with respect to modifications linked with subterranean habits. Troglocharinus is assigned to the moderately modified pholeuonoid morphotype. The body is elongated and slender compared to epigean leiodids and also cave-dwelling species of Ptomaphagini. The legs are elongated, especially the hindlegs, though to a lesser degree than in the most advanced troglobiontic species. The prothorax is moderately elongated but otherwise largely unmodified. Its muscular system is strongly developed, with more muscle bundles that in free-living staphylinoid or hydrophiloid species. The pterothorax is greatly modified, especially the metathoracic flight apparatus. The meso- and metathoracic elements of the elytral locking device are well-developed, whereas the other notal parts are largely reduced. The mesonotum is simplified, with the triangular scutellar shield as the only distinctly developed part. The mesothoracic musculature is strongly reduced, with only 6 muscles compared to 12 or 13 in free-living staphylinoid or hydrophiloid species. The metanotum is greatly reduced, without a recognizable subdivision into prescutum scutum and scutellum. It is strongly narrowing laterally and lacks notal wing processes and other wing-related elements, but well-developed alacristae are present. The wings are reduced to small membranous flap-like structures inserted at the posterior end of the metanotum. A metapostnotum is not developed. Like in the case of the head, cave dwelling species of the related Ptomaphagini and Leptodirini show different trends of adaptations, with a compact ovoid or navicular body shape in the former, and a distinct trend towards elongation of the body and appendages in the latter tribe. Structural affinities of the thoraces of T. ferreri and the troglobiontic trechine carabid Sinaphaenops wangorum are mainly due to the reduced flight apparatus. The degree of muscle reduction in the pterothorax is very similar in both species.
MeSH term(s)	Animals ; Caves ; Coleoptera/anatomy & histology ; Coleoptera/ultrastructure ; Female ; Male ; Microscopy ; Microscopy, Electron, Scanning ; Wings, Animal/anatomy & histology
Language	English
Publishing date	2019-11-25
Publishing country	England
Document type	Journal Article
ZDB-ID	2001498-3
ISSN	1873-5495 ; 1467-8039
ISSN (online)	1873-5495
ISSN	1467-8039
DOI	10.1016/j.asd.2019.100900
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Article ; Online: Pretarsal structures in Leiodidae and Agyrtidae (Coleoptera, Staphylinoidea).

Gnaspini, Pedro / Antunes-Carvalho, Caio / Leschen, Richard A B

Journal of morphology

2017 Volume 278, Issue 10, Page(s) 1354–1379

Abstract: We analysed pretarsal characters of 87 species of Leiodidae (including 10 cholevines and representatives of all tribes and ca. 60% of the genera of non-cholevines), five species of Agyrtidae, and nine representatives of outgroup taxa (Hydraenidae, ... ...

Abstract	We analysed pretarsal characters of 87 species of Leiodidae (including 10 cholevines and representatives of all tribes and ca. 60% of the genera of non-cholevines), five species of Agyrtidae, and nine representatives of outgroup taxa (Hydraenidae, Staphylinidae, Hydrophilidae, and Histeridae) using scanning electron microscopy. We focused our observations on the architecture of the empodium (including the sclerites and associated setae), the shape and composition of the medial projection of the distal margin of the terminal tarsomere, and the armature of the claws, which were considered a promising source of information for delimiting supraspecific taxa in our previous study. We identified several diagnostic features and recognize potential synapomorphies at the tribal, subtribal and generic levels. The internal systematic arrangement and/or even the monophyletic status of most of the subfamilies of Leiodidae (Camiarinae, Catopocerinae, Leiodinae, and Platypsyllinae) are challenged. We identified potential synapomorphies for Camiarinae (Camiarini and Agyrtodini) and Leiodinae. The non-monophyly of Cholevinae is possible because part of the tribe (Anemadini, Eucatopini, and Oritocatopini) shares potentially apomorphic features with Leiodinae (e.g., a triangular medial projection with a diagonal row of conical spines), whereas another part (Leptodirini and Ptomaphagini) shares a potentially apomorphic feature with Coloninae and Platypsyllinae (a typical medial projection with two distinct triangular projections).
Language	English
Publishing date	2017-10
Publishing country	United States
Document type	Journal Article
ZDB-ID	3084-3
ISSN	1097-4687 ; 0022-2887 ; 0362-2525
ISSN (online)	1097-4687
ISSN	0022-2887 ; 0362-2525
DOI	10.1002/jmor.20717
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Article ; Online: Morphology‐based phylogenetic reconstruction of Cholevinae (Coleoptera: Leiodidae): a new view on higher‐level relationships

Antunes‐Carvalho, Caio / Ribera, Ignacio / Beutel, Rolf / Gnaspini, Pedro

Cladistics. 2019 Feb., v. 35, no. 1 p.1-41

2019

Abstract: The current state of knowledge of the suprageneric relationships in Cholevinae is either derived from informal evaluations of putative synapomorphies or based on molecular studies with limited taxonomic sampling. Here we assessed the higher‐level ... ...

Abstract	The current state of knowledge of the suprageneric relationships in Cholevinae is either derived from informal evaluations of putative synapomorphies or based on molecular studies with limited taxonomic sampling. Here we assessed the higher‐level relationships in this subfamily based on a phylogenetic analysis of 97 morphological characters scored for 93 terminals, representing all tribes. Both parsimony and Bayesian analyses were used. The monophyletic origin of Cholevinae was corroborated, except for the unexpected inclusion of Leptinus in the implied weighting analysis. Eucatopini + Oritocatopini were retrieved as basal branches in the evolution of Cholevinae. The monophyletic origin of all remaining Cholevinae was confirmed, which is consistent with molecular evidence. Anemadini was non‐monophyletic, in accordance with earlier hypotheses. Cholevini was rendered non‐monophyletic by the uncertain inclusion of Prionochaeta and the consistent exclusion of Cholevinus. A close affinity of Ptomaphagini to Sciaphyini and Leptodirini was suggested, although the position of Sciaphyes remains uncertain. The phylogenetic hypothesis of Cholevinae provided here is the most comprehensive presently available. The list of characters shows that a substantial part of the data was obtained from the ventral side. This is a strong argument for a detailed pictorial documentation of the ventral body parts in taxonomic descriptions, in contrast to the common practice of only illustrating the dorsal habitus of the beetles.
Keywords	Leiodidae ; monophyly ; synapomorphy ; taxon descriptions
Language	English
Dates of publication	2019-02
Size	p. 1-41.
Publishing place	John Wiley & Sons, Ltd
Document type	Article ; Online
Note	JOURNAL ARTICLE
ZDB-ID	1462608-1
ISSN	1096-0031 ; 0748-3007
ISSN (online)	1096-0031
ISSN	0748-3007
DOI	10.1111/cla.12230
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Article ; Online: Two new synonymies at the generic level in Neotropical Ptomaphagini (Coleoptera, Leiodidae, Cholevinae).

Gnaspini, Pedro / Antunes-Carvalho, Caio / Peck, Stewart B

Zootaxa

2016 Volume 4132, Issue 2, Page(s) 272–274

Abstract: We here recognize the genus Excelsiorella (s. str.) Salgado, 2008 as a junior synonym of the genus Adelopsis Portevin, 1907, and Viruana Salgado, 2013, a subgenus of Excelsiorella, as a junior synonym of the genus Parapaulipalpina Gnaspini, 1996. As a ... ...

Abstract	We here recognize the genus Excelsiorella (s. str.) Salgado, 2008 as a junior synonym of the genus Adelopsis Portevin, 1907, and Viruana Salgado, 2013, a subgenus of Excelsiorella, as a junior synonym of the genus Parapaulipalpina Gnaspini, 1996. As a consequence, we propose the following new combinations: Adelopsis latissima (Salgado, 2008) n. comb. (from Excelsiorella (s. str.)) and Parapaulipalpina tambopata (Salgado, 2013) n. comb. (from Excelsiorella (Viruana)).
MeSH term(s)	Animal Structures/anatomy & histology ; Animal Structures/growth & development ; Animals ; Body Size ; Coleoptera/anatomy & histology ; Coleoptera/classification ; Coleoptera/growth & development ; Male ; Organ Size
Language	English
Publishing date	2016-06-29
Publishing country	New Zealand
Document type	Journal Article
ISSN	1175-5334
ISSN (online)	1175-5334
DOI	10.11646/zootaxa.4132.2.7
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Article ; Online: The cephalic morphology of the troglobiontic cholevine species Troglocharinus ferreri (Coleoptera, Leiodidae).

Luo, Xiao-Zhu / Antunes-Carvalho, Caio / Wipfler, Benjamin / Ribera, Ignacio / Beutel, Rolf G

Journal of morphology

2019 Volume 280, Issue 8, Page(s) 1207–1221

Abstract: Leiodidae are the second largest subterranean radiation of beetles at family rank. To explore morphological trends linked with troglobiontic habits and characters with potential phylogenetic significance, the head of the cave-dwelling species ... ...

Abstract	Leiodidae are the second largest subterranean radiation of beetles at family rank. To explore morphological trends linked with troglobiontic habits and characters with potential phylogenetic significance, the head of the cave-dwelling species Troglocharinus ferreri (Cholevinae, Leptodirini) was examined in detail. Overall, the general pattern is similar to what is found in Catops ventricosus (Cholevini). Shared apomorphic features include a fully exposed anterolateral concavity containing the antennal socket, a distinct bead above this depression, a bilobed lip-like structure anterad the labrum, a flat elevated portion of the ventral mandibular surface, and a ventral process at the proximomesal edge of this mandibular area. The tentorial structures are well-developed as in C. ventricosus, with a large laminatentorium and somewhat shortened dorsal arms. The mouthparts are largely unmodified, with the exception of unusually well-developed extrinsic maxillary muscles. Features of T. ferreri obviously linked with subterranean habits are the complete lack of compound eyes, circumocular ridges, and optic lobes. A series of characters is similar to conditions found in other genera of Leptodirini: the head capsule completely lacks a protruding ocular region, a distinct neck is missing, the transverse occipital crest is indistinct, and the antennae are elongate and lack a distinct club. Two different trends of cephalic transformations occur in troglobiontic Leptodirini, with some genera like Troglocharinus and Leptodirus having elongated head capsules and antennae, and others having broadened, more transverse heads. In contrast, the modifications are more uniform in the closely related Ptomaphagini, with a pattern distinctly differing from Leptodirini: the head is transverse, with a distinctly protruding ocular region, a distinct transverse occipital crest, and a very narrow neck region.
MeSH term(s)	Animals ; Bone and Bones/anatomy & histology ; Bone and Bones/ultrastructure ; Caves ; Coleoptera/anatomy & histology ; Head/anatomy & histology ; Imaging, Three-Dimensional ; Phylogeny
Language	English
Publishing date	2019-06-20
Publishing country	United States
Document type	Journal Article ; Research Support, Non-U.S. Gov't
ZDB-ID	3084-3
ISSN	1097-4687 ; 0022-2887 ; 0362-2525
ISSN (online)	1097-4687
ISSN	0022-2887 ; 0362-2525
DOI	10.1002/jmor.21025
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Article ; Online: Morphology-based phylogenetic reconstruction of Cholevinae (Coleoptera: Leiodidae): a new view on higher-level relationships.

Antunes-Carvalho, Caio / Ribera, Ignacio / Beutel, Rolf G / Gnaspini, Pedro

Cladistics : the international journal of the Willi Hennig Society

2017 Volume 35, Issue 1, Page(s) 1–41

Abstract	The current state of knowledge of the suprageneric relationships in Cholevinae is either derived from informal evaluations of putative synapomorphies or based on molecular studies with limited taxonomic sampling. Here we assessed the higher-level relationships in this subfamily based on a phylogenetic analysis of 97 morphological characters scored for 93 terminals, representing all tribes. Both parsimony and Bayesian analyses were used. The monophyletic origin of Cholevinae was corroborated, except for the unexpected inclusion of Leptinus in the implied weighting analysis. Eucatopini + Oritocatopini were retrieved as basal branches in the evolution of Cholevinae. The monophyletic origin of all remaining Cholevinae was confirmed, which is consistent with molecular evidence. Anemadini was non-monophyletic, in accordance with earlier hypotheses. Cholevini was rendered non-monophyletic by the uncertain inclusion of Prionochaeta and the consistent exclusion of Cholevinus. A close affinity of Ptomaphagini to Sciaphyini and Leptodirini was suggested, although the position of Sciaphyes remains uncertain. The phylogenetic hypothesis of Cholevinae provided here is the most comprehensive presently available. The list of characters shows that a substantial part of the data was obtained from the ventral side. This is a strong argument for a detailed pictorial documentation of the ventral body parts in taxonomic descriptions, in contrast to the common practice of only illustrating the dorsal habitus of the beetles.
Language	English
Publishing date	2017-12-18
Publishing country	United States
Document type	Journal Article
ZDB-ID	1462608-1
ISSN	1096-0031 ; 0748-3007
ISSN (online)	1096-0031
ISSN	0748-3007
DOI	10.1111/cla.12230
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Article ; Online: Show me your tenent setae and I tell you who you are - Telling the story of a neglected character complex with phylogenetic signals using Leiodidae (Coleoptera) as a case study.

Gnaspini, Pedro / Antunes-Carvalho, Caio / Newton, Alfred F / Leschen, Richard A B

Arthropod structure & development

2017 Volume 46, Issue 4, Page(s) 662–685

Abstract: The tarsal setae in 97 species of Leiodidae and eight outgroups were examined using SEM imaging and dissections. Modified adhesive setae present in males are referred to as "male tenent setae" (MTS). In most cases, dilated tarsomeres were associated with ...

Abstract	The tarsal setae in 97 species of Leiodidae and eight outgroups were examined using SEM imaging and dissections. Modified adhesive setae present in males are referred to as "male tenent setae" (MTS). In most cases, dilated tarsomeres were associated with MTS, which were always present on the protarsi and sometimes the mesotarsi. MTS are reported for the first time on the mesotarsi of Leptodirini and on the metatarsi in two genera of Sogdini. Contrary to reports in the literature, the reduction in the number of the MTS bearing mesotarsomeres is considered a derived condition. Both sexes of Leptinus (Platypsyllinae) have modified setae (referred to as tenent setae in the literature), probably related to their specialised association with mammals, and a patch of MTS was recognized for the first time among those modified setae among males. Four main types of MTS are recognised: (1) a plesiomorphic discoidal type that has a shaft with a round cross-section and maintains a similar diameter throughout its length until forming the expanded discoidal terminal plate; (2) a minidiscoidal type, similar to discoidal but with a relatively small terminal plate, found in Cholevinae; (3) a conical type, present in Leiodinae (excluding Estadiini) where the shaft increases in diameter until forming the terminal plate; and (4) a spatulate type, where an even wider terminal plate has a lateral projection, derived from the conical form and synapomorphic for the leiodine tribes Pseudoliodini, Scotocryptini, and possibly Agathidiini.
MeSH term(s)	Animals ; Coleoptera/classification ; Coleoptera/ultrastructure ; Female ; Male ; Microscopy, Electron, Scanning ; Phylogeny ; Sensilla/ultrastructure
Language	English
Publishing date	2017-07
Publishing country	England
Document type	Journal Article
ZDB-ID	2001498-3
ISSN	1873-5495 ; 1467-8039
ISSN (online)	1873-5495
ISSN	1467-8039
DOI	10.1016/j.asd.2017.06.004
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Article ; Online: Two invaders instead of one: the true identity of species under the name Ceracis cucullatus (Coleoptera: Ciidae).

Antunes-Carvalho, Caio / Lopes-Andrade, Cristiano

PloS one

2013 Volume 8, Issue 8, Page(s) e72319

Abstract: The Neotropical obligate fungivorous beetle Ceracis cucullatus (Mellié) has attracted attention of coleopterists due to the increasing number of records of populations in Africa. Although its disjunct populations have been interpreted as a cohesive ... ...

Abstract	The Neotropical obligate fungivorous beetle Ceracis cucullatus (Mellié) has attracted attention of coleopterists due to the increasing number of records of populations in Africa. Although its disjunct populations have been interpreted as a cohesive taxonomic unity, previous comparisons between African and Neotropical specimens revealed differences in their external morphology, causing uncertainty about the true unity of the species. Here, we compare the external morphology of specimens named Cer. cucullatus from several localities of the Neotropical, Palearctic, Afrotropical, Afrotemperate and Oriental regions. As results, we reverse three previous junior synonymies of Cer. cucullatus, proposing Cer. lamellatus (Pic) and Cer. tabellifer (Mellié), both reinstated status and new combinations, as separate species. We also propose Enn. bilamellatum Pic as a new synonym of Cer. tabellifer. In face of these taxonomic changes, we identify Cer. tabellifer as the actual invasive species on African lands, instead of Cer. cucullatus as was previously accepted. Then, through historical records gathered from scientific collections and literature, and through examination of recently collected specimens from South Africa and Brazil, we provide data on host fungi and geographic distribution of Cer. tabellifer. Based on these data, we discuss possible explanations to the successful invasion of Cer. tabellifer in Africa and elsewhere and its potential threat to native faunas of ciids. This study helps to fulfil an old gap in the literature on biological invasions, with considerably more studies on predatory species, disease vectors or potential pests of agricultural crops, than on non-pest fungivorous organisms.
MeSH term(s)	Africa ; Animal Distribution ; Animals ; Coleoptera/anatomy & histology ; Coleoptera/classification ; Female ; Introduced Species ; Male ; South America
Language	English
Publishing date	2013-08-28
Publishing country	United States
Document type	Journal Article ; Research Support, Non-U.S. Gov't
ZDB-ID	2267670-3
ISSN	1932-6203 ; 1932-6203
ISSN (online)	1932-6203
ISSN	1932-6203
DOI	10.1371/journal.pone.0072319
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Article ; Online: Cephalic anatomy and three-dimensional reconstruction of the head of Catops ventricosus (Weise, 1877) (Coleoptera: Leiodidae: Cholevinae)

Antunes-Carvalho, Caio / Yavorskaya, Margarita / Gnaspini, Pedro / Ribera, Ignacio / Hammel, Jörg U. / Beutel, Rolf

Organisms, diversity, & evolution. 2017 Mar., v. 17, no. 1 p.199-212

2017

Abstract: Adult head structures are well known in the coleopteran suborders Archostemata and Adephaga, whereas the available information is very fragmentary in the megadiverse Polyphaga, including the successful superfamily Staphylinoidea. In the present study, ... ...

Abstract	Adult head structures are well known in the coleopteran suborders Archostemata and Adephaga, whereas the available information is very fragmentary in the megadiverse Polyphaga, including the successful superfamily Staphylinoidea. In the present study, the cephalic morphology of the cholevine species Catops ventricosus is described in detail and documented. The results were compared to conditions occurring in other polyphagan lineages, especially staphylinoid and scarabaeoid representatives. Specific external features documented in Catops and potential autapomorphies of Leiodidae include a five-segmented antennal club with a reduced eighth antennomere and the presence of periarticular grooves filled with sensilla on antennomeres 7, 9, and 10. The firm connection of the head and pronotum is possibly an apomorphy of Cholevinae. The monophyly of Cholevinae excluding Eucatopini and Oritocatopini is supported by the apical maxillary palpomere as long as or shorter than the subapical one, and the presence of cryptic pore plates on the surface of these palpomeres—a feature described and documented here for the first time. The internal cephalic structures of Catops are mostly plesiomorphic, as for instance the complete tentorium. The pattern of the muscles is similar to what is found in other staphylinoid taxa. The unusual maxillary muscle “Mx” is likely a groundplan apomorphy of the clade Staphyliniformia + Scarabaeoidea. M. hypopharyngomandibularis (M13) was identified in Catops and is ancestral for Coleoptera, even though it is often missing. The same applies to M. tentoriohypopharyngalis (M42).
Keywords	Leiodidae ; adults ; head ; monophyly ; muscles ; sensilla
Language	English
Dates of publication	2017-03
Size	p. 199-212.
Publishing place	Springer Berlin Heidelberg
Document type	Article ; Online
ZDB-ID	2040179-6
ISSN	1618-1077 ; 1439-6092
ISSN (online)	1618-1077
ISSN	1439-6092
DOI	10.1007/s13127-016-0305-3
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Article ; Online: Two new Neotropical species of Ceracis Mellié (Coleoptera, Ciidae) and redefinition of the cucullatus group.

Antunes-Carvalho, Caio / Lopes-Andrade, Cristiano

ZooKeys

2011 , Issue 132, Page(s) 51–64

Abstract: Two new Neotropical species of Ceracis Mellié are described: Ceracis cassumbensis Antunes-Carvalho & Lopes-Andrade, sp. n. from a single locality in northeastern Brazil and Ceracis navarretei Antunes-Carvalho & Lopes-Andrade, sp. n. from a single ... ...

Abstract	Two new Neotropical species of Ceracis Mellié are described: Ceracis cassumbensis Antunes-Carvalho & Lopes-Andrade, sp. n. from a single locality in northeastern Brazil and Ceracis navarretei Antunes-Carvalho & Lopes-Andrade, sp. n. from a single locality in southern Mexico. Scanning Electron Microscope images of adults and photographs of holotypes and male terminalia are provided for both species, their similarities and differences with other Ceracis are briefly discussed, and the cucullatus species-group is redefined for including the new species described herein.
Language	English
Publishing date	2011-10-03
Publishing country	Bulgaria
Document type	Journal Article
ZDB-ID	2445640-8
ISSN	1313-2970 ; 1313-2989
ISSN (online)	1313-2970
ISSN	1313-2989
DOI	10.3897/zookeys.66.1570
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