Laboulbeniales are a highly specialized group of fungi living only on arthropods. They have high ... more Laboulbeniales are a highly specialized group of fungi living only on arthropods. They have high host specificity and spend their entire life-cycle on an arthropod host. Taxonomic characters of Laboulbeniales are based on the architecture of the cells of the parenchymal thallus, i.e. the visible part of the fungus outside the host. The extent of the fungus spreading inside the host-the haustorium-remains largely unknown. The attachment to the arthropod host is fundamental to understand the fungus-animal interaction, but how this truly occurs is unclear. Recent evidences question the strictly parasitic lifestyle of Laboulbeniales. We used micro-computed tomography (µCT) and 3D reconstructions to visualize, for the first time, the complete structure of Laboulbeniales species in situ on their hosts. We compared the haustoriate species, Arthrorhynchus nycteribiae on an insect host to the non-haustoriate species, Rickia gigas on a millipede host. Our results confirm that some Laboulbeniales species are ectoparasitic and have a haustorial structure that penetrates the host's cuticle, while others are ectobionts and are only firmly attached to the host's cuticle without penetrating it. The presence and the morphology of the haustorium are important traits for Laboulbeniales evolution, and key factors for future understanding of host dependence and specificity.
Millipedes play an important role in the decomposition of organic matter in the subterranean envi... more Millipedes play an important role in the decomposition of organic matter in the subterranean environment. Despite the existence of several cave-adapted species of millipedes in adjacent geographic areas, their study has been largely ignored in Portugal. Over the last decade, intense fieldwork in caves of the mainland and the island of Madeira has provided new data about the distribution and diversity of millipedes. A review of millipedes from caves of Portugal is presented, listing fourteen species belonging to eight families, among which six species are considered troglobionts. The distribution of millipedes in caves of Portugal is discussed and compared with the troglobiont biodiversity in the overall Iberian Peninsula and the Macaronesian archipelagos.
Laboulbeniales are highly specialized arthropod-associated fungi. The majority of the almost 2200... more Laboulbeniales are highly specialized arthropod-associated fungi. The majority of the almost 2200 known species live on insects, although they also occur on other arthropod hosts. Recently, the number of Laboulbeniales associated with millipedes has increased considerably. Here we describe the first species of a Laboulbeniales fungus, Troglomyces twitteri sp. nov., from an American millipede. The new species was initially discovered on a photo of Cambala annulata (Say, 1821) from Ohio, USA, which had been shared on Twitter. A subsequent microscopic study of Cambala millipedes in museum collections in Denmark and France confirmed the discovery.
Two new millipede species of the genus Coxobolellus Pimvichai, Enghoff, Panha & Backeljau, 2020 f... more Two new millipede species of the genus Coxobolellus Pimvichai, Enghoff, Panha & Backeljau, 2020 from Thailand are described: Coxobolellus saratanisp. nov. from Loei Province and Coxobolellus serratoligulatussp. nov. from Uttaradit Province. The descriptions are based on gonopod morphology and two mitochondrial gene fragments (COI and 16S rRNA). The phylogenetic mtDNA analysis assigned the two new species unequivocally to the consistently well-supported Coxobolellus clade, in which they form a fifth subclade that was well supported by maximum likelihood analysis of 16S rRNA, though neither by Bayesian inference nor by COI. The two new Coxobolellus species share four conspicuous gonopodal synapomorphies of the genus: (1) the protruding process on the coxae of the 3rd (and sometimes 4th) pair of male legs, (2) a large, triangular coxae on the 4th–5th pair of legs, (3) a short process of the preanal ring protruding as far as, or slightly beyond, the anal valves, and (4) the posterior go...
The mesovoid shallow substratum (MSS) can act as a climatic refuge for invertebrates, as a biogeo... more The mesovoid shallow substratum (MSS) can act as a climatic refuge for invertebrates, as a biogeographic corridor to deeper substrates or as a permanent habitat for some species. This study characterizes the seasonal invertebrate diversity and abundance of MSS ecosystems in central Portugal focusing on Diplopoda, Diplura, Orthoptera and Coleoptera during one year. Sampling was performed with standard MSS pitfalls in scree slopes (colluvial MSS) of karst areas and environmental parameters (temperature, pH, conductivity, water content, organic carbon, nitrate, phosphate and ammonium) were quantified. Our results show that winter was the season with the highest arthropod abundance and that the MSS acts as a permanent habitat for chordeumatidan millipedes and as a climatic refuge for orthopterans and most beetles. All Diplura collected belong to a single species known previously from surface habitats in the Iberian Peninsula, which does not seem to use the Portuguese MSS as a refuge. MSS habitats in central Portugal, classified as western Mediterranean and thermophile deposits protected by the Natura 2000 network based on plant communities and geology, revealed an abundant and diverse invertebrate community that urges characterization and protection.
Species-level taxonomy and phylogeny of two genera of South-East (SE) Asian pachybolid millipedes... more Species-level taxonomy and phylogeny of two genera of South-East (SE) Asian pachybolid millipedes are analysed with a combination of morphological characters and DNA sequences (two mitochondrial gene fragments: COI and 16S rRNA). Strong support is found for the genera Litostrophus Chamberlin, 1921 and Atopochetus Attems, 1953 and for a clade consisting of Litostrophus + Atopochetus. Four species of Litostrophus are recognised and (re)described: L. segregatus Chamberlin, 1921 (type species of the genus), L. scaber (Verhoeff, 1938), comb. nov., L. chamaeleon, sp. nov. and L. saraburensis, sp. nov. The genus Tonkinbolus Verhoeff, 1938 (type species T. scaber Verhoeff, 1938) is synonymised under Litostrophus. Atopochetus (type species A. rubropunctatus Attems, 1953), hitherto considered a dubious synonym of Aulacobolus Pocock, 1903, is re-instated for several species until now placed in Tonkinbolus and seven new species. All in all, 10 species of Atopochetus are (re)described: A. dollfu...
A catalogue of type specimens of basal hexapods (Collembola, Protura, Diplura, Archaeognatha, Zyg... more A catalogue of type specimens of basal hexapods (Collembola, Protura, Diplura, Archaeognatha, Zygentoma) in the collection of the Natural History Museum of Denmark, University of Copenhagen is presented. The NHMD collection harbours type material of 175 species, and it is composed of 132 holotypes, 4 neotypes, 3 syntypes, 32 lectotypes, and 39 paratypes. For Diplura one neotype is newly designated. The order Protura is represented by 5 families, Collembola by 7 families, Diplura by 6 families, Archaeognatha by 1 family and Zygentoma by 2 families.
A new millipede species, Ommatoiulus longicornis n. sp. is described from the subterranean habita... more A new millipede species, Ommatoiulus longicornis n. sp. is described from the subterranean habitat in eastern Spain, and the elongation of its antennae is discussed in relation to the habitat where it dwells. We take the occasion to discuss the elongation of appendages as a troglobiomorphic character in millipedes and shed the light on the various, sometimes controversial, hypotheses that could have led to these modifications in millipedes and other subterranean arthropods.
The troglobiont millipede, Cantabrodesmus lorioli Mauriès, 1971, inhabits a few caves in a small ... more The troglobiont millipede, Cantabrodesmus lorioli Mauriès, 1971, inhabits a few caves in a small region of Northern Spain. It is considered a relict species because of its geographical isolation compared to others in its family, and its to some extent uncertain systematic position. Cantabrodesmus lorioli has been placed in three different families, and now is considered a member of the Chelodesmidae, a family with only one other known native European representative. There are few records of this species, and almost nothing is known about its biology. The only available information on this species is the description of the male and a list of caves where it has been captured. In the present work, we provide the first description of the female morphology, illustrated by scanning electron microscopy, and a series of new records from caves in a region where it had never been captured before, including the deepest known pit in Spain, Torca del Porrón. The female morphology does not show many differences from that of the male, but several characters are discussed. All of the known information on the morphology of the species is used to discuss the implications for its biology and systematic position.
The troglobiont millipede Typhlopsychrosoma baeticaense (Mauriès, 2013) is recorded in the Mesovo... more The troglobiont millipede Typhlopsychrosoma baeticaense (Mauriès, 2013) is recorded in the Mesovoid Shallow Substratum (MSS) of two screes in the Aitana and Bernia mountains (Eastern Iberian Peninsula), far away from its known distribution area. A detailed Scanning Electron Microscopy iconography provides additional information on gonopod morphology, as well as other details such as the inconspicuous evaginations of the cuticle at the place of the eyes in the anophthalmous specimens. We present an updated distribution map and a key to species of the genus, with illustrations of the gonopods of all species described so far. The implications of the appearance of this hypogean species in an MSS are discussed.
The chordeumatidan fauna of the Iberian Peninsula is far from being well known, but recent effort... more The chordeumatidan fauna of the Iberian Peninsula is far from being well known, but recent efforts are improving that knowledge. Samplings carried out in the Milieu Souterrain Superficiel (also known as the Mesovoid Shallow Substratum) on several screes of the Sierra de Guadarrama National Park (central Spain) have led to the discovery of a new species of chordeumatidan millipede. This new species belongs to the family Haplobainosomatidae, which is mainly known from the northern Iberian Peninsula, and thus this is the first record of the family in central Spain. However, the extremely simple gonopods of this species are so different from all other known species of the family that it must be placed in a new genus. In this work, a complete description of the species, named as Guadarramasoma ramosae gen. & sp. nov., with a detailed iconography based on scanning electron microscopy images is provided together with a distribution map and a brief discussion of the implications of this new finding.
New records are given for secondary capilliconidia of Basidiobolus from several species of millip... more New records are given for secondary capilliconidia of Basidiobolus from several species of millipedes (Diplopoda) belonging to three different orders. The anamorph 'Thaxteriola" stage of Rhynchomyces is recorded (with doubt) from a Brazilian millipede belonging to the order Spirostreptida. An enigmatic fungus showing characteristics of Coreomycetopsis, Hormiscioideus and Antennopsis is recorded from two species of Danish millipedes of the order Julida. Peculiar structures, tentatively referred to fungi are recorded from several millipede orders where they occur between micro-scutes of the external cuticle.
A large, newly collected material of the genus Aquattuor from the Udzungwa Mts, Tanzania, has bee... more A large, newly collected material of the genus Aquattuor from the Udzungwa Mts, Tanzania, has been studied. Two new species are described, viz., A. mollilobus sp. nov. from the Udzungwa Mts and A. nguruensis sp. nov. from the Nguru Mts, Tanzania. Aquattuor claudiahempae Enghoff & Frederiksen, 2015, is recorded as new for the Udzungwa fauna. Characters from the first pair of male legs are illustrated for all Aquattuor species and are shown to be useful for species discrimination. A new gonopodal difference between A. submajor Enghoff, 2015 and A. udzungwensis Enghoff, 2015 is described and illustrated. Intrageneric relationships and distribution patterns are discussed. A distribution map and a key to all Aquattuor species are presented
The millipede Blaniulus dollfusi (Brölemann, 1894) is recorded as new for the Belgian fauna, base... more The millipede Blaniulus dollfusi (Brölemann, 1894) is recorded as new for the Belgian fauna, based on specimens collected in Tervuren. The species is native to S. France, N. Spain and N. Portugal and is probably introduced in Belgium. Samenvatting. Een 700 km disjuncte populatie van de duizendpoot Blaniulus dollfusi in België (Diplopoda, Julida: Blaniuludae) De duizendpoot Blaniulus dollfusi (Brölemann, 1894) wordt hier als nieuw voor de Belgische fauna vermeld, gebaseerd op exemplaren verzameld te Tervuren. De soort komt voor in Zuid-Frankrijk, Noord-Spanje en Noord-Portugal en is waarschijnlijk geïmporteerd in België. Résumé. Une population disjoncte de 700 km du millipède Blaniulus dollfusi en Belgique (Diplopoda, Julida: Blaniulidae) Le millipède Blaniulus dollfusi (Brölemann, 1894) est mentionné ici comme nouveau pour la faune belge d'après des exemplaires collectés à Tervuren. L'espèce vit en France méridionale, en Espagne septentrional et au Portugal septentrional et elle est probablement importée en Belgique.
Estes co-autores contribuíram com dados originais relativos à distribuição de espécies e são apre... more Estes co-autores contribuíram com dados originais relativos à distribuição de espécies e são apresentados por ordem alfabética do último nome (These co-authors contributed with original data on species distribution and we rank them by alphabetical order).
Laboulbeniales are a highly specialized group of fungi living only on arthropods. They have high ... more Laboulbeniales are a highly specialized group of fungi living only on arthropods. They have high host specificity and spend their entire life-cycle on an arthropod host. Taxonomic characters of Laboulbeniales are based on the architecture of the cells of the parenchymal thallus, i.e. the visible part of the fungus outside the host. The extent of the fungus spreading inside the host-the haustorium-remains largely unknown. The attachment to the arthropod host is fundamental to understand the fungus-animal interaction, but how this truly occurs is unclear. Recent evidences question the strictly parasitic lifestyle of Laboulbeniales. We used micro-computed tomography (µCT) and 3D reconstructions to visualize, for the first time, the complete structure of Laboulbeniales species in situ on their hosts. We compared the haustoriate species, Arthrorhynchus nycteribiae on an insect host to the non-haustoriate species, Rickia gigas on a millipede host. Our results confirm that some Laboulbeniales species are ectoparasitic and have a haustorial structure that penetrates the host's cuticle, while others are ectobionts and are only firmly attached to the host's cuticle without penetrating it. The presence and the morphology of the haustorium are important traits for Laboulbeniales evolution, and key factors for future understanding of host dependence and specificity.
Millipedes play an important role in the decomposition of organic matter in the subterranean envi... more Millipedes play an important role in the decomposition of organic matter in the subterranean environment. Despite the existence of several cave-adapted species of millipedes in adjacent geographic areas, their study has been largely ignored in Portugal. Over the last decade, intense fieldwork in caves of the mainland and the island of Madeira has provided new data about the distribution and diversity of millipedes. A review of millipedes from caves of Portugal is presented, listing fourteen species belonging to eight families, among which six species are considered troglobionts. The distribution of millipedes in caves of Portugal is discussed and compared with the troglobiont biodiversity in the overall Iberian Peninsula and the Macaronesian archipelagos.
Laboulbeniales are highly specialized arthropod-associated fungi. The majority of the almost 2200... more Laboulbeniales are highly specialized arthropod-associated fungi. The majority of the almost 2200 known species live on insects, although they also occur on other arthropod hosts. Recently, the number of Laboulbeniales associated with millipedes has increased considerably. Here we describe the first species of a Laboulbeniales fungus, Troglomyces twitteri sp. nov., from an American millipede. The new species was initially discovered on a photo of Cambala annulata (Say, 1821) from Ohio, USA, which had been shared on Twitter. A subsequent microscopic study of Cambala millipedes in museum collections in Denmark and France confirmed the discovery.
Two new millipede species of the genus Coxobolellus Pimvichai, Enghoff, Panha & Backeljau, 2020 f... more Two new millipede species of the genus Coxobolellus Pimvichai, Enghoff, Panha & Backeljau, 2020 from Thailand are described: Coxobolellus saratanisp. nov. from Loei Province and Coxobolellus serratoligulatussp. nov. from Uttaradit Province. The descriptions are based on gonopod morphology and two mitochondrial gene fragments (COI and 16S rRNA). The phylogenetic mtDNA analysis assigned the two new species unequivocally to the consistently well-supported Coxobolellus clade, in which they form a fifth subclade that was well supported by maximum likelihood analysis of 16S rRNA, though neither by Bayesian inference nor by COI. The two new Coxobolellus species share four conspicuous gonopodal synapomorphies of the genus: (1) the protruding process on the coxae of the 3rd (and sometimes 4th) pair of male legs, (2) a large, triangular coxae on the 4th–5th pair of legs, (3) a short process of the preanal ring protruding as far as, or slightly beyond, the anal valves, and (4) the posterior go...
The mesovoid shallow substratum (MSS) can act as a climatic refuge for invertebrates, as a biogeo... more The mesovoid shallow substratum (MSS) can act as a climatic refuge for invertebrates, as a biogeographic corridor to deeper substrates or as a permanent habitat for some species. This study characterizes the seasonal invertebrate diversity and abundance of MSS ecosystems in central Portugal focusing on Diplopoda, Diplura, Orthoptera and Coleoptera during one year. Sampling was performed with standard MSS pitfalls in scree slopes (colluvial MSS) of karst areas and environmental parameters (temperature, pH, conductivity, water content, organic carbon, nitrate, phosphate and ammonium) were quantified. Our results show that winter was the season with the highest arthropod abundance and that the MSS acts as a permanent habitat for chordeumatidan millipedes and as a climatic refuge for orthopterans and most beetles. All Diplura collected belong to a single species known previously from surface habitats in the Iberian Peninsula, which does not seem to use the Portuguese MSS as a refuge. MSS habitats in central Portugal, classified as western Mediterranean and thermophile deposits protected by the Natura 2000 network based on plant communities and geology, revealed an abundant and diverse invertebrate community that urges characterization and protection.
Species-level taxonomy and phylogeny of two genera of South-East (SE) Asian pachybolid millipedes... more Species-level taxonomy and phylogeny of two genera of South-East (SE) Asian pachybolid millipedes are analysed with a combination of morphological characters and DNA sequences (two mitochondrial gene fragments: COI and 16S rRNA). Strong support is found for the genera Litostrophus Chamberlin, 1921 and Atopochetus Attems, 1953 and for a clade consisting of Litostrophus + Atopochetus. Four species of Litostrophus are recognised and (re)described: L. segregatus Chamberlin, 1921 (type species of the genus), L. scaber (Verhoeff, 1938), comb. nov., L. chamaeleon, sp. nov. and L. saraburensis, sp. nov. The genus Tonkinbolus Verhoeff, 1938 (type species T. scaber Verhoeff, 1938) is synonymised under Litostrophus. Atopochetus (type species A. rubropunctatus Attems, 1953), hitherto considered a dubious synonym of Aulacobolus Pocock, 1903, is re-instated for several species until now placed in Tonkinbolus and seven new species. All in all, 10 species of Atopochetus are (re)described: A. dollfu...
A catalogue of type specimens of basal hexapods (Collembola, Protura, Diplura, Archaeognatha, Zyg... more A catalogue of type specimens of basal hexapods (Collembola, Protura, Diplura, Archaeognatha, Zygentoma) in the collection of the Natural History Museum of Denmark, University of Copenhagen is presented. The NHMD collection harbours type material of 175 species, and it is composed of 132 holotypes, 4 neotypes, 3 syntypes, 32 lectotypes, and 39 paratypes. For Diplura one neotype is newly designated. The order Protura is represented by 5 families, Collembola by 7 families, Diplura by 6 families, Archaeognatha by 1 family and Zygentoma by 2 families.
A new millipede species, Ommatoiulus longicornis n. sp. is described from the subterranean habita... more A new millipede species, Ommatoiulus longicornis n. sp. is described from the subterranean habitat in eastern Spain, and the elongation of its antennae is discussed in relation to the habitat where it dwells. We take the occasion to discuss the elongation of appendages as a troglobiomorphic character in millipedes and shed the light on the various, sometimes controversial, hypotheses that could have led to these modifications in millipedes and other subterranean arthropods.
The troglobiont millipede, Cantabrodesmus lorioli Mauriès, 1971, inhabits a few caves in a small ... more The troglobiont millipede, Cantabrodesmus lorioli Mauriès, 1971, inhabits a few caves in a small region of Northern Spain. It is considered a relict species because of its geographical isolation compared to others in its family, and its to some extent uncertain systematic position. Cantabrodesmus lorioli has been placed in three different families, and now is considered a member of the Chelodesmidae, a family with only one other known native European representative. There are few records of this species, and almost nothing is known about its biology. The only available information on this species is the description of the male and a list of caves where it has been captured. In the present work, we provide the first description of the female morphology, illustrated by scanning electron microscopy, and a series of new records from caves in a region where it had never been captured before, including the deepest known pit in Spain, Torca del Porrón. The female morphology does not show many differences from that of the male, but several characters are discussed. All of the known information on the morphology of the species is used to discuss the implications for its biology and systematic position.
The troglobiont millipede Typhlopsychrosoma baeticaense (Mauriès, 2013) is recorded in the Mesovo... more The troglobiont millipede Typhlopsychrosoma baeticaense (Mauriès, 2013) is recorded in the Mesovoid Shallow Substratum (MSS) of two screes in the Aitana and Bernia mountains (Eastern Iberian Peninsula), far away from its known distribution area. A detailed Scanning Electron Microscopy iconography provides additional information on gonopod morphology, as well as other details such as the inconspicuous evaginations of the cuticle at the place of the eyes in the anophthalmous specimens. We present an updated distribution map and a key to species of the genus, with illustrations of the gonopods of all species described so far. The implications of the appearance of this hypogean species in an MSS are discussed.
The chordeumatidan fauna of the Iberian Peninsula is far from being well known, but recent effort... more The chordeumatidan fauna of the Iberian Peninsula is far from being well known, but recent efforts are improving that knowledge. Samplings carried out in the Milieu Souterrain Superficiel (also known as the Mesovoid Shallow Substratum) on several screes of the Sierra de Guadarrama National Park (central Spain) have led to the discovery of a new species of chordeumatidan millipede. This new species belongs to the family Haplobainosomatidae, which is mainly known from the northern Iberian Peninsula, and thus this is the first record of the family in central Spain. However, the extremely simple gonopods of this species are so different from all other known species of the family that it must be placed in a new genus. In this work, a complete description of the species, named as Guadarramasoma ramosae gen. & sp. nov., with a detailed iconography based on scanning electron microscopy images is provided together with a distribution map and a brief discussion of the implications of this new finding.
New records are given for secondary capilliconidia of Basidiobolus from several species of millip... more New records are given for secondary capilliconidia of Basidiobolus from several species of millipedes (Diplopoda) belonging to three different orders. The anamorph 'Thaxteriola" stage of Rhynchomyces is recorded (with doubt) from a Brazilian millipede belonging to the order Spirostreptida. An enigmatic fungus showing characteristics of Coreomycetopsis, Hormiscioideus and Antennopsis is recorded from two species of Danish millipedes of the order Julida. Peculiar structures, tentatively referred to fungi are recorded from several millipede orders where they occur between micro-scutes of the external cuticle.
A large, newly collected material of the genus Aquattuor from the Udzungwa Mts, Tanzania, has bee... more A large, newly collected material of the genus Aquattuor from the Udzungwa Mts, Tanzania, has been studied. Two new species are described, viz., A. mollilobus sp. nov. from the Udzungwa Mts and A. nguruensis sp. nov. from the Nguru Mts, Tanzania. Aquattuor claudiahempae Enghoff & Frederiksen, 2015, is recorded as new for the Udzungwa fauna. Characters from the first pair of male legs are illustrated for all Aquattuor species and are shown to be useful for species discrimination. A new gonopodal difference between A. submajor Enghoff, 2015 and A. udzungwensis Enghoff, 2015 is described and illustrated. Intrageneric relationships and distribution patterns are discussed. A distribution map and a key to all Aquattuor species are presented
The millipede Blaniulus dollfusi (Brölemann, 1894) is recorded as new for the Belgian fauna, base... more The millipede Blaniulus dollfusi (Brölemann, 1894) is recorded as new for the Belgian fauna, based on specimens collected in Tervuren. The species is native to S. France, N. Spain and N. Portugal and is probably introduced in Belgium. Samenvatting. Een 700 km disjuncte populatie van de duizendpoot Blaniulus dollfusi in België (Diplopoda, Julida: Blaniuludae) De duizendpoot Blaniulus dollfusi (Brölemann, 1894) wordt hier als nieuw voor de Belgische fauna vermeld, gebaseerd op exemplaren verzameld te Tervuren. De soort komt voor in Zuid-Frankrijk, Noord-Spanje en Noord-Portugal en is waarschijnlijk geïmporteerd in België. Résumé. Une population disjoncte de 700 km du millipède Blaniulus dollfusi en Belgique (Diplopoda, Julida: Blaniulidae) Le millipède Blaniulus dollfusi (Brölemann, 1894) est mentionné ici comme nouveau pour la faune belge d'après des exemplaires collectés à Tervuren. L'espèce vit en France méridionale, en Espagne septentrional et au Portugal septentrional et elle est probablement importée en Belgique.
Estes co-autores contribuíram com dados originais relativos à distribuição de espécies e são apre... more Estes co-autores contribuíram com dados originais relativos à distribuição de espécies e são apresentados por ordem alfabética do último nome (These co-authors contributed with original data on species distribution and we rank them by alphabetical order).
Uploads
Papers by Henrik Enghoff