Gene regulatory dynamics during the development of a paleopteran insect, the mayfly Cloeon dipterum.

The evolution of insects has been marked by the appearance of key body plan innovations that promoted the outstanding ability of this lineage to adapt to new habitats, boosting the most successful radiation in animals. To understand the evolution of these new structures, it is essential to investigate which are the genes and gene regulatory networks participating during the embryonic development of insects. Great efforts have been made to fully understand gene expression and gene regulation during the development of holometabolous insects, in particular Drosophila melanogaster. Conversely, functional genomics resources and databases in other insect lineages are scarce. To provide a new platform to study gene regulation in insects, we generated ATAC-seq (Assay for transposase-Accessible Chromatin) for the first time during the development of the mayfly Cloeon dipterum, which belongs to Paleoptera, the sister group to all other winged insects. With these comprehensive datasets along six developmental stages, we characterised pronounced changes in accessible chromatin between early and late embryogenesis. The application of ATAC-seq in mayflies provides a fundamental resource to understand the evolution of gene regulation in insects.

Taxonomic notes on the genus Baetiella Uéno, 1931 (Ephemeroptera, Baetidae), with description of three new species from Thailand.

Based on material recently collected in northern Thailand, the present study provides an updated of the genus Baetiella, including Gratia. It comprises six species in Thailand, three of them being new species: Baetiella (Gratia) narumonae, Baetiella (Gratia) sororculaenadinae, Baetiella (Baetiella) bispinosa, Baetiella (Baetiella) baeisp. nov., Baetiella (Baetiella) lannaensissp. nov. and Baetiella (Baetiella) bibranchiasp. nov.Baetiella (Baetiella) baeisp. nov. can be distinguished from other species by the reduction of the posteromedian protuberances on abdominal tergites I-III, the asymmetrical coniform terminal segment of labial palp, the distal margin of abdominal sternites VII-X each with a row of long, spatulate setae, the dorsal margin of femur with two long, robust setae distally. Baetiella (Baetiella) lannaensissp. nov. is diagnosed by the posteromedian protuberances present on tergites I-VIII, dorsal margin of femur with a regular row of long, rounded, ciliated setae and body surface covered with numerous, dense, rounded scale-like setae. Baetiella (Baetiella) bibranchiasp. nov. can be separated from other species by coxal gills present at the base of forelegs and midlegs. The molecular study based on the mitochondrial gene COI and a larval key to species of Thai Baetiella are also provided.

Another new species of the genus Habrophlebia Eaton, 1881 (Ephemeroptera, Leptophlebiidae) from the Maghreb.

A new species belonging to the genus Habrophlebia Eaton, 1881 is described at the nymphal stage from the Rif Mountains of Morocco. This species presents unique features, such as the chorionic arrangement of the egg and the ornamentation of the posterior margin of abdominal tergites. It is compared to all west European Habrophlebia species and a table with discriminating characters is given. A phylogenetic reconstruction based on COI sequences fully supports the hypothesis of a new species in the Rif Mountains, with possible further distribution in southern Spain.

Baetidae (Insecta: Ephemeroptera) of Aurès Mountains (Algeria): A New Species of the Baetis alpinus Species Group, with Notes on Baetis Laech, 1815 Biogeography within Maghreb.

A new species, Baetis (Baetis) dihyaesp. nov., belonging to the Baetis alpinus species group, is described and illustrated based on larval material collected in the Aurès Mountains (northeastern Algeria) in 2020-2021. This new species is closely related to three European species, e.g., Baetis (B.) alpinus (Pictet, 1843); B. (B.) nubecularis Eaton, 1898; and B. (B.) pasquetorum Righetti & Thomas, 2002 by the combination of the following characteristics: (i) more than one short, stout bristle at the tip of segment II of the maxillary palp and (ii) a well-developed paracercus. However, the new species clearly differs from all congeners of the Baetis alpinus species group primarily by the (a) structure of mouthparts-with 14-18 long submarginal setae arranged in a single irregular row on the dorsal surface of the labrum; 2-6 short, stout bristles at the tip of segment II of the maxillary palp; and segment II of the labial palp without a considerably developed apico-internal lobe); (b) setation of abdominal terga, with a few triangular-shaped scales sparsely scattered near the posterior margin only; and (c) a well-developed paracercus, comprised of more than 50 segments. Primary data on the biology and distribution of this new species are provided, and molecular affinities are verified by the analysis of COI (barcode) sequences. Detailed notes on the distribution of mayfly species belonging to the Baetis alpinus species group common in Western Europe and the western part of North Africa are presented. The historical movement of Baetis representatives between Europe, North West Africa, and subsequently Algeria, with the land bridges 'Strait of Gibraltar' and 'Strait of Sicily' as colonization routes, is discussed in detail and identified in the present study as the Western Algeria colonization path and Eastern Algeria colonization path, respectively.

A new species of the genus Alainites Waltz & McCafferty, 1994 (Ephemeroptera, Baetidae) from the north of Morocco.

A new species of Alainites is described from northern of Morocco Alainitesalbaisp. nov. It can be separated from the other west Palearctic species by the gill number, the spination of the distal margin of tergites, the leg setation, and the paraproct shape and spination. This species is widespread in the study area but never abundant. It prefers small to medium streams with slow flow, and does not seem to be very sensitive to pollution and water logging activities.

A new species of the Baetisfuscatus group (Ephemeroptera, Baetidae) from Morocco.

Baetisrifensissp. nov. is the first representative of the Baetisfuscatus group to be described from the Maghreb. It was collected from streams in the Rif region of northern Morocco. All species of the B.fuscatus group are morphologically very similar, with slight differences in colour. Thus, in addition to morphological description, species delimitation based on genetic evidence was carried out. The new species was compared with other members of the B.fuscatus group from the Palaearctic region.

New species of Nigrobaetis from Southeast Asia (Ephemeroptera, Baetidae).

Based on larvae collected in the Philippines, Borneo (Kalimantan), Sumba, and Sumatra, the presence of the genus Nigrobaetis in the Philippines and Indonesia is reported for the first time. Six new species are described and illustrated, two from the Philippines and four from Indonesia. A larval key to all Nigrobaetis species of the Philippines, Indonesia and neighbouring continental Southeast Asia is provided, morphological differences to the species of Taiwan are discussed as well. The eggs of three new species are described, and the morphology of the eggs of Nigrobaetis is briefly discussed.

Big diversity in a small hotspot: two new species of Leptophlebiidae (Insecta, Ephemeroptera) from New Caledonia.

Two new species from Grande Terre Island, New Caledonia, namely Fasciamiruspetersorumsp. nov. and Simulacalararasp. nov. are described based on larval morphology and molecular data (COI sequences). Fasciamiruspetersorumsp. nov. is distributed in the southern part of the island and is characterised by a reduced third segment of the labial palps and all abdominal gills divided from the base. The species inhabits slow-flowing aquatic habitats with fine-grained substrate in forest brooks. Simulacalararasp. nov. is known from a single locality in the northern part of the island and is characterised by narrow and distinctly elongated abdominal gills 1-7. It was collected from fine substrates behind stones in riffles with slightly turbulent flow. Both species were recorded only in areas with ultramafic bedrock.

First report of the Afrotropical genus Securiops Jacobus, McCafferty & Gattolliat (Ephemeroptera, Baetidae) from Southeast Asia, with description of a new species.

Recent collections in Thailand revealed the occurrence of the genus Securiops in Asia, formerly known from the Afrotropical Realm only. A new species of Securiops is described and illustrated based on larvae and eggs. Eggs of this genus are described for the first time. Morphological differences between the new species and the species from Africa are discussed. The number of species in the genus Securiops is augmented to five.

Baetidae (Baetidae, Ephemeroptera) in the Maghreb: state of the art, key, and perspectives.

Among mayflies, Baetidae are often considered as easy to recognise at the family level, but difficult to identify at lower level. In several faunistic or ecological studies, the identification remains at the family level; Baetidae are generally considered as widespread and ubiquitous, therefore as poorly informative for ecological studies or bioassessments. Here, a straightforward identification key is offered to larvae of the ten genera of Baetidae reported from Maghreb based on easily observable and understandable characters. The diversity, ecology, and distribution of each taxonomic unit (genera or subgenera) are discussed and the main difficulties for deeper identification are pointed out. Future challenges and remaining taxonomic riddles for Maghrebian Baetidae are detailed.

The Tribe Hyrtanellini Allen, 1980 (Ephemeroptera: Ephemerellidae) of Western and Central Asia with Description of a New Species.

A new species, Serratella leonidi Martynov & Palatov, sp. nov., is described from Tajikistan based on immature stage. Based on larval material from Iran including the topotypes, Serratella elissa Jacobus, Zhou & McCafferty, 2009 is complementary described, and its generic placement is clarified. The delimitation of three genera that are members of the tribe Hyrtanellini Allen, 1980, namely Serratella Edmunds, 1959, Torleya Lestage, 1917 and Quatica Jacobus & McCafferty, 2008 is briefly discussed. The phylogenetic reconstruction of Hyrtanellini based on the COI gene showed the relations of representatives of these genera on the one hand, and distinct delimitation of Serratella leonidi sp. nov. and S. elissa on the other. A list of species from Western and Central Asia attributed to Hyrtanellini, their currently known distribution and a key for the determination of the larvae are proposed.

Hermanella complex on Northern Brazil (Ephemeroptera: Leptophlebiidae): New species and stage descriptions

ABSTRACT The male and female imago stage and a new species of the Hermanella complex are described based on material from Pará State, Northern Brazil: Hydromastodon cf. mikei (Thomas & Boutonnet, 2004) and Paramaka froehlichi sp. nov. The male imago of Hydromastodon cf. mikei can be distinguished from Hydromastodon sallesi by the following combination of characters: (1) body color pattern, especially abdominal terga translucent white with terga I to IV washed with black on posterior margin; (2) compound eyes anteriorly rounded; (3) medial projection of the styliger plate short; and (4) penis lobes close to each other and with posterolateral projection as long as ½ of its length. The male imago of Paramaka froehlichi sp.nov. can be distinguished the from the other species of the genus by the following combination of characters: (1) fore and hind wings with membrane brown; (2) legs uniformly yellowish-white, without distal band; (3) abdominal terga II to VI with marks more expanded laterally, median line absent; (4) medial projection of the styliger plate broad; (5) base of telopenis arising at distal ⅔ of penis lobe; and (6): body size of 6.1 to 7.6 mm.

A new, widespread genus of Baetidae from South Asia (Insecta, Ephemeroptera).

Material collected on different islands across South Asia revealed a new genus of Baetidae with a widespread distribution, Arcobaetisgen. nov. The larvae present important similarities with Nigrobaetis, but have paraglossae dorsally with an arc of long, spine-like setae in distal area; long, slightly feathered setae between prostheca and mola of both mandibles; and very slender legs with row of short setae at dorsal margin of femur. The male imago has an extraordinarily small 3rd (apical) segment of gonostylus, which is much narrower than the apex of the 2nd segment. The new genus includes five species: A.sumbawensissp. nov. is described from Sumbawa (Indonesia) based on larvae, A.sumatrensissp. nov. from Sumatra (Indonesia) based on larvae, A.bornensissp. nov. from Borneo (Brunei) based on larvae, and A.sripadaisp. nov. (type species) is described from Sri Lanka based on a reared male imago with its larval and subimaginal exuviae; A.gracilentus (Chang & Yang, 1994), comb. nov. from Taiwan, formerly described in Margobaetis Kang & Yang, 1994, a subgenus of Baetis Leach, 1815, and subsequently transferred to the genus Nigrobaetis Kazlauskas (in Novikova & Kluge), 1987, is transferred to the new genus. A key to the larvae of all species is provided. Morphological similarities and the relationship of the new genus to other genera of Baetidae are discussed.

First report of the genus Tenuibaetis (Ephemeroptera, Baetidae) from Thailand revealing a complex of cryptic species.

A new species of the genus Tenuibaetis Kang & Yang, 1994 is described from Thailand and the genus is reported for the first time from this country. Tenuibaetispanhai sp. nov. is easily distinguished from other known Tenuibaetis by the complete absence of hindwing pads. Molecular evidence based on COI confirmed the validity of the new species. Additional putative species of Tenuibaetis based on molecular evidence only are considered as Molecular Operational Taxonomic Units (MOTUs) without description. The morphological characters of the new species and its closely related species are discussed; a key to the Oriental species is provided.

Mercury levels in sediment, fish and macroinvertebrates of the Boroo River, northern Mongolia, under the legacy of gold mining.

Gold mining is currently one of the main anthropogenic sources of mercury in the environment. In this study, the total mercury content was measured in bottom sediments, benthic macroinvertebrates (mayfly larvae), and fish (Siberian dace) along the Boroo River in northern Mongolia. There was a gold recovery plant in the middle reaches of the river until the mid-twentieth century; an accident there in the 1950s caused a mercury spill. We found an increased content of mercury in measured ecosystem components near the plant compared to the upper reaches of the river. The mercury content in sediments varied from trace amounts in the upper Boroo to 2200 ng/g dry weight (dw) in the vicinity of the plant ruins. The mercury content in mayfly larvae ranged from 50 to 2940 ng/g dw and had a spatial pattern as sediments, with the highest concentrations near the plant. The mercury content in sediments was lower at the mouth of the Boroo River than near the plant, reflecting the lower boundary of the mercury spill. Maximum values of mercury content in fish muscle were found at the river's mouth and were several times higher than in other rivers of northern Mongolia. Median mercury content in muscles of dace from the lower Boroo in 2016 has doubled since studies in 2010-2012, which may be the result of current mercury releases from gold mining.

Ecdyonurusaurasius sp. nov. (Insecta, Ephemeroptera, Heptageniidae, Ecdyonurinae), a new micro-endemic mayfly species from Aurès Mountains (north-eastern Algeria).

Ecdyonurusaurasius sp. nov., a micro-endemic species reported from several streams within the Aurès Mountains (north-eastern Algeria), is described and illustrated at nymphal, subimaginal and imaginal stages of both sexes. Critical morphological diagnostic characters distinguishing the new species are presented, together with molecular affinities as well as notes on the biology and distribution of the species.

Megabranchiella gen. nov., a new mayfly genus (Ephemeroptera, Baetidae) from Thailand with description of two new species.

Megabranchiella gen. nov. (Ephemeroptera: Baetidae) is established as a new baetid mayfly genus from northern Thailand. Two new species, Megabranchiellascutulata sp. nov. and Megabranchiellalongusa sp. nov., are described. This genus is distinguished from other Baetidae by abdominal segment I, bearing a pair of enlarged, ventrally oriented single gills, covering abdominal sternites II-V; other gills have normal size and are dorsolaterally oriented. The two new species Megabranchiellalongusa sp. nov. and Megabranchiellascutulata sp. nov. can be differentiated by the setation of femur dorsal margin and the shape of abdominal gill I. This mayfly genus was found in flowing water with cobble microhabitats in headwater streams of northern Thailand.

Two new species of Centroptilum Eaton, 1869 from North Africa (Ephemeroptera, Baetidae).

Based on recently collected larvae from Algeria and Morocco, the species delimitation within the genus Centroptilum Eaton, 1869 in that region is validated. Two new species are described and illustrated, one from north-eastern Algeria, and one from North Morocco, using an integrated approach with morphological and molecular evidence. A table summarising the morphological differences between the new species and Centroptilumluteolum (Müller, 1776) from Central Europe is provided. Further, molecular evidence for additional undescribed species of Centroptilum in other regions of the West Palearctic is provided and discussed.

Mayfly Ephemeraglaucops (Ephemeroptera, Ephemeridae) recorded in the Czech Republic after almost a century.

Background: The mayfly Ephemera (Sinephemera) glaucops Pictet, 1843 has been considered regionally extinct in the Czech Republic, with the last occurrence dating from 1933. Its extinction was connected with the anthropogenic changes of lowland rivers, forming the original habitat of E.glaucops within the area of the Czech Republic. However, the species has been reported as spreading in man-made, often post-industrial waterbodies in Germany, The Netherlands and Austria since the 1970s. New information: We report E.glaucops from the Czech Republic, based on 27 larvae collected in the North Bohemia lignite basin in 2018. All individuals were found at one locality - a small kaolin pit in the shallow part near the shore, mostly without macrophytes. We provide details about the locality and morphological diagnostic characters of E.glaucops. This study highlights the importance of post-industrial sites for aquatic biodiversity as surrogate biotopes for degraded original habitats.

Methylmercury in caddisflies and mayflies: Influences of water and sediment chemistry.

Quantifying methylmercury (MeHg) concentrations and uptake at the base of the food web is useful for assessing mercury exposure risk to higher trophic level organisms. Higher MeHg concentrations near the base of the food web may result in more MeHg exposure and accumulation in higher trophic organisms. Here, we analyze MeHg in caddisflies, mayflies, lake water, and sediment collected from two temperate lakes and one brook in Kejimkujik National Park, Nova Scotia, Canada. Overall, caddisfly larvae MeHg (15.38-276.96 ng/g; n = 29) was not significantly correlated with water chemistry. Whereas mayfly naiads MeHg (14.28-166.82 ng/g; n = 31) was positively correlated with water MeHg (rs = 0.43), negatively correlated with pH (rs = -0.49), and positively correlated with dissolved organic carbon (DOC; rs = 0.48). Of the mercury in insect tissues, the %MeHg ranged from 56 to 75 % in caddisfly larvae and 38-47 % in mayfly naiads. MeHg bioaccumulation factors (BAF) varied greatly (water to tissue BAFs = 0.145 × 106-1.054 × 106; sediment to tissue BAFs = 0.017 × 106-0.541 × 106). This study highlights the importance of quantifying variations in MeHg bioaccumulation and BAFs of common aquatic insect bioindicators at the base of complex food webs.