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.

Unravelling the diversity of the genus Afronurus Lestage, 1924 (Ephemeroptera, Heptageniidae) in Thailand.

The genus Afronurus in Thailand is investigated using an integrative approach (morphology based, ootaxonomy and molecular data) for species delimitation. A total of four species of Afronurus was identified; A.cervina (Braasch & Soldán, 1984), A.gilliesiana (Braasch, 1990), A.rainulfiana (Braasch, 1990), and A.rubromaculata (You et al., 1981). The subimago of A.gilliesiana is described for the first time based on reared specimens. The egg structure of all four species is also described for the first time. Morphological and molecular data strongly support their species delimitation. The egg chorionic structure of the genus Afronurus, together with other morphological characters, is useful for species identification. A key to mature nymphs of the known species is provided.

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.

What Ecological Factors Favor Parthenogenesis over Sexual Reproduction? A Study on the Facultatively Parthenogenetic Mayfly Alainites muticus in Natural Populations.

AbstractDifferent reproductive modes are characterized by costs and benefits that depend on ecological contexts. For example, sex can provide benefits under complex biotic interactions, while its costs increase under mate limitation. Furthermore, ecological contexts often vary along abiotic gradients. Here, we study how these factors simultaneously influence the frequency of sex in the facultatively parthenogenetic mayfly Alainites muticus. We first verified that parthenogenesis translates into female-biased population sex ratios. We then measured the density of individuals (a proxy for mate limitation) and community diversity (biotic interaction complexity) for 159 A. muticus populations covering a broad altitudinal gradient and used structural equation modeling to investigate their direct and indirect influences on sex ratios. We found no effect of community diversity or altitude on sex ratios. Furthermore, even when females can reproduce parthenogenetically, they generally reproduce sexually, indicating that the benefits of sex exceed its costs in most situations. Sex ratios become female-biased only under low population densities, as expected if mate limitation was the main factor selecting for parthenogenesis. Mate limitation might be widespread in mayflies because of their short adult life span and limited dispersal, which can generate strong selection for reproductive assurance and may provide a stepping stone toward obligate parthenogenesis.

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.

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.

Cincticostellaebura sp. nov., a new species of mayfly (Ephemeroptera, Ephemerellidae) from Thailand.

A new species of ephemerellid mayfly, Cincticostellaebura sp. nov., is described based on larvae collected in a stream from Nan Province, Thailand. This new species is classified in the nigra complex of the genus Cincticostella based on morphological and COI phylogeny evidence. The new species is closely related to C.nigra (Uéno, 1928) and C.funki Martynov, Selvakumar, Palatov & Vasanth, 2021 based on body colour pattern. Investigation of the chorionic structure of the new species showed similar details to those of other species within this species complex. The phylogeny also placed this species into a monophyletic group with C.nigra (Uéno, 1928), C.elongatula (McLachlan, 1875) and C.fusca Kang & Yang, 1995.

A new species of the genus Prosopistoma Latreille, 1833 (Ephemeroptera, Prosopistomatidae) from Morocco.

We describe a new species of Prosopistoma collected in the High Atlas Mountains of Morocco. Prosopistomamaroccanum sp. nov. appears to be morphologically more similar to the European highly endangered P.pennigerum (Müller, 1785) than to the other Maghrebian species, P.alaini Bojkova & Soldán, 2015. A gene tree including the few available barcode sequences of Palearctic Prosopistoma specimens is provided. Possible affinities with West African species are also discussed.

Contribution to the knowledge of Teloganodidae (Ephemeroptera, Ephemerelloidea) of India.

Two new species of Dudgeodes Sartori, 2008 and a new species of Teloganodes Eaton, 1882 are described from India; they are Dudgeodesselvakumari Martynov & Palatov, sp. nov. from Himalayan region (Uttarakhand), Dudgeodesmolinerii Sivaruban, Martynov, Srinivasan, Barathy & Isack, sp. nov., and Teloganodesbarathyae Sivaruban, Martynov, Srinivasan & Isack, sp. nov. from the Tamil Nadu part of the Western Ghats. Thus, for now, the Teloganodidae fauna of India includes 11 species. Dudgeodesselvakumari sp. nov. appears to be significantly extend northward the known distribution of Dudgeodes. Partial COI sequences were used as an initial clustering method to show the relationships of D.selvakumari sp. nov. with other sequenced operational taxonomic units (OTU) of the genus.

Behningiidae and Potamanthidae (Insecta, Ephemeroptera) in Thailand.

The aim of this study is to review the family Behningiidae and Potamanthidae in Thailand. Two genera and three species of Behningiidae are recognised: Protobehningiamerga Peters & Gillies, 1991, Behningiabaei McCafferty & Jacobus, 2006, and Behningianujiangensis Zhou & Bisset, 2019, which is newly reported from Thailand. The egg structure of B.nujiangensis is described for the first time using scanning electron microscopy. The larva of P.merga is redescribed and its distribution is expended northward with a new record from Chiang Mai province. Two genera and five species of Potamanthidae are identified: Potamanthusformosus Eaton, 1892, Rhoenanthusmagnificus Ulmer, 1920 (new record for Thailand), Rhoenanthusobscurus Navás, 1922, Rhoenanthusdistafurcus Bae & McCafferty, 1991, and Rhoenanthusspeciosus Eaton, 1881. Our morphological evidence is supported by COI data for the family Potamanthidae. Diagnostic characters, distributions, and keys are presented for the larvae of all known species of Thai behningiid and potamanthid mayflies.

A new species of Neohagenulus Traver, 1938 from Hispaniola (Ephemeroptera, Leptophlebiidae, Hagenulinae, Hagenulini).

Here, I report a new species of the genus Neohagenulus Traver, 1938 from the Dominican Republic. The genus was believed to be endemic to Puerto Rico until now. Neohagenulushodeceki sp. nov. is described at the nymphal stage. Some discussion on the tribe Hagenulini is also provided.

Contribution to knowledge of the Trichoptera of northwestern Algeria: New species records for the Algerian fauna and taxonomic remarks for the Maghreb fauna.

The caddisfly fauna of Algeria is far from complete. In this study, we present data from Trichoptera sampling in northwestern Algeria (Tafna and Ghazouana River basins) during 20142019, enhancing knowledge of this group in the country. We were able to identify 30 species and 17 genera in the study area. Among the listed taxa, five species (Hydroptila fortunata, Ecnomus deceptor, Tinodes waeneri, Limnephilus lunatus, Setodes argentipunctellus,) are new records for Algeria. An updated species checklist of Algerian Trichoptera includes 63 species. We also provide an update of the Maghreb checklist with comments on species distributions. The Maghrebian caddisfly checklist, which includes 44 Maghrebian endemic species, is still very poor in relation to that of neighboring Mediterranean geographical areas and countries, such as the Iberian Peninsula (Spain and Portugal), France, or Italy.

First record of Paegniodes Eaton, 1881 (Ephemeroptera, Heptageniidae) from Thailand with description of a new species.

A new species of Heptageniidae, Paegniodes sapanensis sp. nov., is described based on larvae, subimagos, eggs, and COI data. The mayfly genus Paegniodes Eaton, 1881 is reported for the first time from Thailand. The larva of the new species can be distinguished from other known Paegniodes species by i) lamellae of gill I ca 1/4 of fibrilliform portion and ii) mandibles and basal segment of maxillary palp without dense setae on margin. The subimago characters useful to distinguish this new species from previously known species are i) the median stripes on abdominal terga and ii) shape of the female subgenital and subanal plates. The genetic distance between the new species and P. cupulatus (Eaton, 1871) was approximately 11%. The morphological characters of the new species are discussed and compared to other known species.

Larval Taxonomy and Distribution of Genus Hydropsyche (Trichoptera: Hydropsychidae) in Northwestern Algeria.

The main purpose of our paper is to document genus Hydropsyche (Trichoptera: Hydropsychidae) in northwestern Algeria and to provide the larval descriptions of the species set. Larvae, pupae, and imagines were collected from 14 sampling sites over a five-year period (2014-2019). Eight Hydropsyche species have been identified, with Hydropsyche siltalai being a new record for Algeria and North Africa, and Hydropsyche pellucidula a new record for Algeria. A comparison of our checklist with those of the Moroccan Rif and Europe is provided, as well as the geographical distribution of each species. This Algerian Hydropsyche species revision provides precise and reliable taxonomic characters for distinguishing larvae of the species, and a taxonomic key is proposed for their identification. In addition, information regarding their distribution is included.

Extremely Widespread Parthenogenesis and a Trade-Off Between Alternative Forms of Reproduction in Mayflies (Ephemeroptera).

Studying alternative forms of reproduction in natural populations is of fundamental importance for understanding the costs and benefits of sex. Mayflies are one of the few animal groups where sexual reproduction co-occurs with different types of parthenogenesis, providing ideal conditions for identifying benefits of sex in natural populations. Here, we establish a catalog of all known mayfly species capable of reproducing by parthenogenesis, as well as species unable to do so. Overall, 1.8% of the described species reproduce parthenogenetically, which is an order of magnitude higher than reported in other animal groups. This frequency even reaches 47.8% if estimates are based on the number of studied rather than described mayfly species, as reproductive modes have thus far been studied in only 17 out of 42 families. We find that sex is a more successful strategy than parthenogenesis (associated with a higher hatching success of eggs), with a trade-off between the hatching success of parthenogenetic and sexual eggs. This means that improving the capacity for parthenogenesis may come at a cost for sexual reproduction. Such a trade-off can help explain why facultative parthenogenesis is extremely rare among animals despite its potential to combine the benefits of sexual and parthenogenetic reproduction. We argue that parthenogenesis is frequently selected in mayflies in spite of this probable trade-off because their typically low dispersal ability and short and fragile adult life may frequently generate situations of mate limitation in females. Mayflies are currently clearly underappreciated for understanding the benefits of sex under natural conditions.

A reassessment of the genus Oligoneuriopsis Crass, 1947 (Ephemeroptera, Oligoneuriidae, Oligoneuriellini).

The distinction between the two closely related genera Oligoneuriella Ulmer, 1924 and Oligoneuriopsis Crass, 1947 has been much debated. First described from South Africa, Oligoneuriopsis seemed to be a clearly defined genus. However, as the known distribution of the genus widened and knowledge on it expanded, species delimitation based on morphology became less clear due to overlap in several apparently defining morphological characters, especially in the nymphs. This work attempts to reassess Oligoneuriopsis morphology in the context of all currently known species. The type species, Oligoneuriopsis lawrencei Crass, 1947 is redescribed at the imaginal and nymphal stages and a neotype is designated. The putative nymph of Oligoneuriopsis dobbsi (Eaton, 1912) is described based on material collected around Mt Elgon (Kenya). The adults of Oligoneuriella orontensis Koch, 1980 are described for the first time and the species is transferred to the genus Oligoneuriopsis (Oligoneuriopsis orontensis comb. nov.). Egg structure is also described for the first time for the species Oligoneuriopsis skhounate and O. orontensis. Some biogeographical considerations are also given. It is likely that more species will still be discovered, especially in Africa.

Stylogomphus thongphaphumensis (Odonata: Anisoptera: Gomphidae), a new gomphid dragonfly and the first record of S. malayanus Sasamoto, 2001 from Thailand.

Stylogomphus thongphaphumensis sp. nov. is described from a type series of specimens reared from larvae (holotype ♂, Huai Khayeng, Thong Pha Phum district, Kanchanaburi Province; 14°36'20″N 98°34'38″E, 206 m a.s.l., larva collected on 14.XII.2014; adult emerged on 30.IV.2015). All larvae were collected from the same locality in western Thailand. Description of the larva (based on preserved exuviae) is provided as well. The adult of this species can be distinguished from other Stylogomphus Fraser, 1922, by the morphology of the male anal appendages, pterothoracic pattern, abdominal pattern, male genitalia and female valvula vulvae (detail provided in the differential diagnosis below). This is the first species of Stylogomphus to be described from Thailand. This study also reports the first record of Stylogomphus malayanus Sasamoto, 2001, from Thailand.

Reports of Baetidae (Ephemeroptera) species from Tafna Basin, Algeria and biogeographic affinities revealed by DNA barcoding.

BACKGROUND: The Mediterranean basin is known to be the cradle of many endemic species. Within mayflies (Insecta, Ephemeroptera), North African species belonging to the family Baetidae remain poorly known and, traditionally, affinities to European fauna were proposed. Recent studies, based on molecular reconstructions, showed closer relationships to Mediterranean islands fauna. NEW INFORMATION: Baetidae were sampled from North-West Algerian wadis (Tafna basin) and involved in COI barcoding reconstructions. Seven species were identified. The subgenus Rhodobaetis is represented by Baetis atlanticus known previously from Macaronesian islands, Europe and Morocco and the Maghrebian endemic Baetis sinespinosus. Specimens, previously identified as Cloeon cf. dipterum, correspond to Cloeon peregrinator and, until now, only reported from Macaronesia. Besides the confirmation of endemicity of some species, such as Procloen stagnicola and B. sinespinosus, our molecular study showed quite original results for relationships between European, insular and Algerian species. Baetis maurus stood out as a North African endemic sister clade to an Iberian clade. Furthermore, we found clear interspecific distances between Algerian and European clades for A. cf. sinaica and B. cf. pavidus, suggesting the presence of cryptic species in Algeria. However, additional studies are needed, as, for the moment, no clear morphological characters were found to separate the different clades and support them as valid species.

An unexpected new species of Habrophlebia from Algeria (Ephemeroptera, Leptophlebiidae).

We describe a new species of Habrophlebia, H. djurdjurensis sp. nov., based on nymphal, imaginal, and egg stages obtained by sampling from the Great Kabylia watershed, north-central Algeria. The new species was previously identified as H. cf. fusca by Lounaci et al. 2000. Habrophlebia djurdjurensis is in fact more related to H. vaillantorum Thomas, 1996 but can be separated by characters on the nymphs and male imago. This is the fourth species of Habrophlebia reported from North Africa.

A New Species of Vietnamella Tshernova 1972 (Ephemeroptera: Vietnamellidae) from Thailand.

The larva, male subimago, female imago, and eggs of V. nanensis sp. n. are described based on specimens from Mae Hong Son and Nan provinces, Thailand. The female subimago is described based on a photograph of a specimen reared to the imago stage. The species previously was distinguished only by DNA barcode data and designated as Vietnamella sp. C. Based on morphology, the larva of the new species can be distinguished with the following combination of characteristics: (i) pattern of serration on the ventral margin of the forefemur, (ii) posterolateral margins of abdominal terga with pairs of acute tubercles, especially terga VI and VII, (iii) a well-developed pair of median ridge projections on tergum X, (iv) the second segment of the maxillary palp being about 1.3× the length of the third segment, and (v) females containing eggs with prominent protuberances on the chorionic surface. A key to larvae of all known species in the genus is provided.