Catalogue of the Diptera (Insecta) of Morocco- an annotated checklist, with distributions and a bibliography.

The faunistic knowledge of the Diptera of Morocco recorded from 1787 to 2021 is summarized and updated in this first catalogue of Moroccan Diptera species. A total of 3057 species, classified into 948 genera and 93 families (21 Nematocera and 72 Brachycera), are listed. Taxa (superfamily, family, genus and species) have been updated according to current interpretations, based on reviews in the literature, the expertise of authors and contributors, and recently conducted fieldwork. Data to compile this catalogue were primarily gathered from the literature. In total, 1225 references were consulted and some information was also obtained from online databases. Each family was reviewed and the checklist updated by the respective taxon expert(s), including the number of species that can be expected for that family in Morocco. For each valid species, synonyms known to have been used for published records from Morocco are listed under the currently accepted name. Where available, distribution within Morocco is also included. One new combination is proposed: Assuaniamelanoleuca (Séguy, 1941), comb. nov. (Chloropidae).

Local Insect Availability Partly Explains Geographical Differences in Floral Visitor Assemblages of Arum maculatum L. (Araceae).

Geographical variation in abundance and composition of pollinator assemblages may result in variable selection pressures among plant populations and drive plant diversification. However, there is limited knowledge on whether differences in local visitor and pollinator assemblages are the result of site-specific strategies of plants to interact with their pollinators and/or merely reflect the pollinator availability at a given locality. To address this question, we compared locally available insect communities obtained by light-trapping with assemblages of floral visitors in populations of Arum maculatum (Araceae) from north vs. south of the Alps. We further investigated whether and how the abundance of different visitors affects plants' female reproductive success and examined the pollen loads of abundant visitors. Local insect availability explained inter-regional differences in total visitor abundance, but only partly the composition of visitor assemblages. Northern populations predominantly attracted females of Psychoda phalaenoides (Psychodidae, Diptera), reflecting the high availability of this moth fly in this region. More generalized visitor assemblages, including other psychodid and non-psychodid groups, were observed in the south, where the availability of P. phalaenoides/Psychodidae was limited. Fruit set was higher in the north than in the south but correlated positively in both regions with the abundance of total visitors and psychodids; in the north, however, this relationship disappeared when visitor abundances were too high. High pollen loads were recorded on both psychodids and other Diptera. We demonstrate for the first time that the quantitative assessment of floral visitor assemblages in relation to locally available insect communities is helpful to understand patterns of geographical variation in plant-pollinator interactions. This combined approach revealed that geographical differences in floral visitors of A. maculatum are only partly shaped by the local insect availability. Potential other factors that may contribute to the geographical pattern of visitor assemblages include the region-specific attractiveness of this plant species to flower visitors and the population-specific behavior of pollinators.

Antennae of psychodid and sphaerocerid flies respond to a high variety of floral scent compounds of deceptive Arum maculatum L.

Insect-pollinated plants often release complex mixtures of floral scents to attract their pollinators. Yet scent compounds eliciting physiological or behavioural responses in pollinators have only been identified in few plant species. The sapromyiophilous aroid Arum maculatum releases a highly diverse dung-like scent with overall more than 300 different compounds recorded so far to attract its psychodid and other fly pollinators. The volatiles' role in pollinator attraction is mostly unknown. To identify potential behaviourally active compounds, we recorded electroantennographic responses of four Psychodidae and one Sphaeroceridae species to (1) inflorescence scents of A. maculatum and (2) the scents released by cow dung, likely imitated by the plant species. Here we show that these flies are sensitive to 78 floral volatiles of various chemical classes, 18 of which were also found in cow dung. Our study, which for the first time determined physiologically active compounds in the antennae of Psychoda spp. and Sphaeroceridae, identified various volatiles not known to be biologically active in any floral visitors so far. The obtained results help deciphering the chemical basis that enables A. maculatum and other plants, pollinated by psychodids and sphaerocerids, to attract and deceive their pollinators.

New synonyms and new species of European aquatic dance flies (Diptera, Empididae).

Knowledge of the European aquatic dance flies (Clinocerinae, Hemerodromiinae) is updated. The following new synonyms are proposed: Chelifera pallida Vaillant, 1981 with Chelifera diversicauda Collin, 1927, syn. nov.; Chelifera giraudae Vaillant, 1981 and Chelifera berdeni Vaillant, 1978 (nomen nudum) with Chelifera subangusta Collin, 1961, syn. nov.; Hemerodromia slovenica Horvat & Wagner, 1989 with Hemerodromia laudatoria Collin, 1927, syn. nov.; Hemerodromia maculata Vaillant, 1968 with Hemerodromia unilineata Zetterstedt, 1842, syn. nov.; Wiedemannia aerea Vaillant, 1967 with Wiedemannia hastata (Mik, 1880), syn. nov.; Wiedemannia brevilamellata Wagner, 1985 with Wiedemannia debilis Collin, 1961, syn. nov.; Wiedemannia bilobata Oldenberg, 1910 with Wiedemannia oxystoma (Bezzi, 1905), syn. nov. Lectotypes are designated for Clinocera (Wiedemannia) oxystoma Bezzi, 1905, Wiedemannia aerea Vaillant, 1967, Wiedemannia bilobata Oldenberg, 1910, Philolutra erminea Mik, 1887, Wiedemannia hastata (Mik, 1880), Wiedemannia mirousei Vaillant, 1956 and Phaeobalia pokornyi Mik, 1886. A neotype is designated for Wiedemannia alpina Vaillant, 1953. Wiedemannia rudolfi Wagner & Ivkovic, sp. nov. and Phaeobalia remschakae Wagner sp. nov. are described and figures of male terminalia are provided.

Moth flies (Diptera, Psychodidae) living in the dark of caves in the Dinaric Karst.

Seoda cavernicola sp. nov. and Psychoda glamocensis sp. nov., are new species and cave dwellers from Bosnia and Hercegovina, and Croatia. Adults of S. cavernicola are pale and small; the eye bridge is reduced, ommatidia irregularly arranged, epandrium with a pair of setose excrescences. The eye bridge of P. glamocensis is likewise reduced with 2 or 3 irregularly ordered facet rows, palpus segments of some individuals are malformed; its closest relative is Psychoda alticola Vaillant based on the morphology of male and female genitalia as well as on COI barcodes.

The oldest of the rare: A new species of the family Nymphomyiidae (Diptera) from Burmese amber.

Nymphomyia allissae sp. nov. from the approximately 100 million years old Burmese amber is described and figured. The species is distinguished from other known taxa by the scimitar shaped wing, and the well sclerotized female terminalia with large cerci.

The West-Palearctic species of the genus Tonnoiriella Vaillant, 1971 (Diptera: Psychodidae, Psychodinae).

Tonnoiriella Vaillant, 1971 was one of the genera not treated by F. Vaillant for his revision in Die Fliegen der paläartktischen Region. Here we provide a genus diagnosis, re-describe known species based mainly on type material, and provide descriptions and figures of the West-Palearctic species. New species described are T. aurasica sp. nov., T. italiae sp. nov., T. ikariae sp. nov., T. andradei sp. nov., T. rhodesica sp. nov., and T. goncalvesi sp. nov.; T. androsica Vaillant, 1978, T. orientalis Vaillant, 1978, and T. pallidipenis Vaillant, 1978 are nomina nuda; finally 21 species are known to date from Europe and North Africa. The ejaculatory apodeme (basiphallus) is asymmetric and bilobed, lobes are stacked; one or both lobes are connected with asymmetric, cross positioned (distiphallus) sclerites. Operating of the open-close mechanism of the aedeagus is explained. Information on larval ecology is provided and the systematic position of the genus is discussed.

Complex and nonlinear climate-driven changes in freshwater insect communities over 42 years.

The ongoing biodiversity crisis becomes evident in the widely observed decline in abundance and diversity of species, profound changes in community structure, and shifts in species' phenology. Insects are among the most affected groups, with documented decreases in abundance up to 76% in the last 25-30 years in some terrestrial ecosystems. Identifying the underlying drivers is a major obstacle as most ecosystems are affected by multiple stressors simultaneously and in situ measurements of environmental variables are often missing. In our study, we investigated a headwater stream belonging to the most common stream type in Germany located in a nature reserve with no major anthropogenic impacts except climate change. We used the most comprehensive quantitative long-term data set on aquatic insects available, which includes weekly measurements of species-level insect abundance, daily water temperature and stream discharge as well as measurements of additional physicochemical variables for a 42-year period (1969-2010). Overall, water temperature increased by 1.88 °C and discharge patterns changed significantly. These changes were accompanied by an 81.6% decline in insect abundance, but an increase in richness (+8.5%), Shannon diversity (+22.7%), evenness (+22.4%), and interannual turnover (+34%). Moreover, the community's trophic structure and phenology changed: the duration of emergence increased by 15.2 days, whereas the peak of emergence moved 13.4 days earlier. Additionally, we observed short-term fluctuations (<5 years) in almost all metrics as well as complex and nonlinear responses of the community toward climate change that would have been missed by simply using snapshot data or shorter time series. Our results indicate that climate change has already altered biotic communities severely even in protected areas, where no other interacting stressors (pollution, habitat fragmentation, etc.) are present. This is a striking example of the scientific value of comprehensive long-term data in capturing the complex responses of communities toward climate change.

Cambios Complejos y No Lineales Causados por el Clima en Comunidades de Insectos de Agua Dulce durante 42 Años Resumen La continua crisis de la biodiversidad se vuelve evidente en la ampliamente documentada declinación en la abundancia y diversidad de especies, cambios profundos en la estructura de las comunidades y modificaciones en la fenología de las especies. Los insectos se encuentran entre los grupos más afectados; se han documentado reducciones en la abundancia de hasta el 76% en los últimos 25-30 años en algunos ecosistemas terrestres. La identificación de los causantes subyacentes es un gran obstáculo porque la mayoría de los ecosistemas están afectados por varios factores estresantes simultáneamente y con frecuencia faltan las medidas in situ de las variables ambientales. Investigamos un flujo naciente perteneciente al tipo de arroyo más común en Alemania ubicado dentro de una reserva natural sin impactos antropogénicos importantes a excepción del cambio climático. Usamos el conjunto más completo disponible de datos cuantitativos de largo plazo para insectos acuáticos que incluye las medidas semanales de abundancia de insectos a nivel especie, las medidas diarias de la temperatura del agua y la descarga del flujo y las medidas de variables físico-químicas adicionales durante 42 años (1969 - 2010). En general, la temperatura del agua incrementó 1.88°C y los patrones de descarga cambiaron significativamente. Estos cambios estuvieron acompañados por una declinación del 81.6% en la abundancia de insectos, pero también de un incremento en la riqueza (+8.5%), la diversidad Shannon (+22.7%), la uniformidad (+22.4%) y la renovación interanual (+34%). Además, la estructura trófica y la fenología de la comunidad cambiaron: la duración del surgimiento incrementó en 15.2 días y el pico del surgimiento ocurrió con 13.4 días de anticipación. Observamos fluctuaciones a corto plazo (<5 años) en casi todas las medidas, así como respuestas complejas y no lineales de la comunidad hacia el cambio climático que podrían haber sido ignoradas si sólo se usaran datos instantáneos o series de tiempo más cortas. Nuestros resultados indican que el cambio climático ya ha alterado seriamente a las comunidades bióticas, incluso en áreas protegidas en las que no están presentes otros factores estresantes en interacción (contaminación, fragmentación del hábitat, etc.). Ésto es un ejemplo notable del valor científico que tienen los datos completos de escalas de tiempo a largo plazo para la captura de las respuestas complejas de las comunidades ante el cambio climático.

Unnoticed arrival of two dipteran species in Austria: the synanthropic moth fly Clogmia albipunctata (Williston, 1893) and the parasitic bird louse fly Ornithoica turdi (Olivier in Latreille, 1811).

In the framework of a mosquito-monitoring program conducted from 2014 to 2018, non-culicid dipteran bycatch was identified to species-level with a focus on Diptera of medical and veterinary importance as part of a biodiversity initiative and barcoding project ("Austrian Barcode of Life"). Two species hitherto not known from Austria, the regularly sampled synanthropic moth fly Clogmia albipunctata (Psychodidae) and a single specimen of the louse fly Ornithoica turdi (Hippoboscidae), were collected in Vienna and Lower Austria. We confirmed identification results using a barcoding approach and provide the first reference sequence for O. turdi.

Elevated temperatures translate into reduced dispersal abilities in a natural population of an aquatic insect.

Rising global temperatures force many species to shift their distribution ranges. However, whether or not (and how fast) such range shifts occur depends on species' dispersal capacities. In most ecological studies, dispersal-related traits (such as the wing size or wing loading in insects) are treated as fixed, species-specific characteristics, ignoring the important role of phenotypic plasticity during insect development. We tested the hypothesis that dispersal-related traits themselves vary in dependence of ambient environmental conditions (temperature regimes, discharge patterns and biotic interactions during individual development). We collected data over 8 years from a natural population of the crane fly Tipula maxima in central Germany. Using linear mixed-effect models, we analysed how phenotypic traits, phenological characteristics and population densities are affected by environmental conditions during the preceding 3, 6 and 12 months. We found a moderate (5.6%) increase in wing length per 1°C increase in mean annual temperatures during the previous year. At the same time, body weight increased by as much as 17.8% in females and 26.9% in males per 1°C, likely driven by increased habitat productivity, which resulted in a 16.4% (female) and 19.3% (male) increased wing loading. We further found a shorter, more synchronized emergence period (i.e. a narrower time frame for dispersal) with increasing temperatures. Altogether, our results suggest that dispersal abilities of T. maxima were negatively affected by elevated temperatures, and we discuss how similar patterns might affect the persistence of populations of other aquatic insects, especially stenoecious taxa with narrow distribution ranges. Our study calls for integration of information on temperature-induced phenotypic plasticity of dispersal-related traits into models forecasting range shifts in the face of climate change. Furthermore, the patterns reported here are likely to affect metapopulation dynamics of aquatic insects under climate change conditions and may contribute to the ongoing decline of insect biomass and diversity.

Remarkable new Australian mothflies (Diptera, Psychodidae).

Queenslandomyia cannula gen. nov. et sp. nov. from Queensland, and Theischingerus imbricatus gen. nov. et sp. nov. (Diptera, Psychodidae) from New South Wales, are described and figured. Queenslandomyia is characterized by the shape of the basal 4 flagellomeres with digitate lateral extensions, and an afferent sclerotized thin channel to flagellomere 4; parameres connate to a large central sclerite with apical setae. Theischingerus is conspicuous by the unusual furcation of the Radius sector vein in R2 and R3+4 so that between radial and medial forks only a single vein (R5) occurs. Rotundopteryx notata (Duckhouse, 1966) is redescribed and the genus diagnosis of Rotundopteryx Duckhouse 1990 is expanded, based on a remarkable connection of the basiphallus, with the distiphallus and the parameres, and of the subepandrial plate with the surstyli and the hypoproct. Records of other 6 species are provided.

Cross-realm assessment of climate change impacts on species' abundance trends.

Climate change, land-use change, pollution and exploitation are among the main drivers of species' population trends; however, their relative importance is much debated. We used a unique collection of over 1,000 local population time series in 22 communities across terrestrial, freshwater and marine realms within central Europe to compare the impacts of long-term temperature change and other environmental drivers from 1980 onwards. To disentangle different drivers, we related species' population trends to species- and driver-specific attributes, such as temperature and habitat preference or pollution tolerance. We found a consistent impact of temperature change on the local abundances of terrestrial species. Populations of warm-dwelling species increased more than those of cold-dwelling species. In contrast, impacts of temperature change on aquatic species' abundances were variable. Effects of temperature preference were more consistent in terrestrial communities than effects of habitat preference, suggesting that the impacts of temperature change have become widespread for recent changes in abundance within many terrestrial communities of central Europe.

Cladistic analysis of Subfamily Bruchomyiinae (Diptera: Psychodidae).

Subfamily Bruchomyiinae is comprised of 60 species and has been referred to as the most primitive within the Psychodidae. The assumed sister-group relationship with Phlebotominae is based on ecological constraints of their environment. A cladistics analysis based on 29 characters and 52 species revealed the distinction of an Old World clade characterized by males with elongate, narrow vasa deferentia, and a New World clade with males having shorter and basally widened vasa deferentia. The Old World clade consists of the genera Nemopalpus Macquart (9 species), and Eutonnoiria Alexander (1 species). The New World clade includes Bruchomyia Alexander (10 species), Boreofairchildia genus nov. (13 species), Laurenceomyia genus nov. (5 species), and Notofairchildia genus nov. (15 species). Parsimony and Bayesian analyses resulted in trees that generally support this generic classification; however, with some species groups less resolved. Diagnostic features for genera are provided. In contrast to the other New World genera, Notofairchildia is paraphyletic with the provisional inclusion of at least the Australasian taxa.

Review of the Psychodinae from Mallorca, Spain, with description of Pericoma unipennata, sp. n. (Diptera, Psychodidae).

We review the Psychodinae of Mallorca, recognising fifteen species based on recent collections and available literature. Previously unpublished data is presented for eleven species, of which Neoarisemus ibericus Wagner, 1978, Mormia tenebricosa (Vaillant, 1954), Clogmia albipunctata (Williston, 1893), Lepiseodina rothschildi (Eaton, 1913), Paramormia ustulata (Walker, 1856), Philosepedon pyrenaicus Vaillant, 1974 and Psychoda (Psycha) grisescens Tonnoir, 1922 are first records for Mallorca. An old record of Pericoma trifasciata (Meigen, 1804) is considered doubtful. Pericoma unipennata sp. n is described and illustrated based on a male collected at Deía. Distributional data are reviewed for all newly recorded species. Based on the Psychodinae fauna, the zoogeographical affinities of Mallorca are briefly discussed.

Datziinae as a new subfamily name for the unavailable name Protopsychodinae Stebner et al., 2015, (Diptera: Psychodidae).

In a recent paper a new subfamily of Psychodidae was inadequately named Protopsychodinae. This nomenclatural act cannot be considered as a valid name under ICZN regulations because the subfamily name is not based on the type genus Datzia Stebner et al., 2015, and furthermore the fossil genus Protopsychoda Azar et al., 1999 was originally described under the subfamily Psychodinae. Therefore, the new family-group name Datziinae is herein proposed.

Thaumaleidae (Diptera) collected by the late Dr. W. Joost in the Caucasus Mountains.

The aquatic insect collection of the late Dr. W. Joost contained two new species of Thaumaleidae (Diptera) from the Caucasus Mountains: Thaumalea monikae sp. n. and Thaumalea biacuminata sp. n. These two new species are herein described, and the most abundant species in Dr. Joost's collection, Thaumalea martinovskyi Joost, 1979, is redescribed based on the type material. Figures of male and female genitalia for all species are provided. All three species show morphological similarities to taxa from the Eastern Mediterranean area. Thaumalea monikae is related to the European T. bezzii-species group, T. biacuminata to the T. serrata-group, and T. martinovskyi to T. kyladica Wagner, 1981 and T. malickyi Theischinger, 1979 from the Eastern Mediterranean area.

Moth flies and sand flies (Diptera: Psychodidae) in Cretaceous Burmese amber.

One new subfamily, four new genera and 10 new species of Psychodidae are described from Burmese amber which significantly increases our knowledge about this group in the Cretaceous. Protopsychodinae n. subfam. probably represents the oldest known ancestor of modern Psychodinae and includes three species within two genera: Datzia setosa gen. et sp. n., Datzia bispina gen. et sp. n., and Mandalayia beumersorum gen. et sp. n. Sycoracinae and Phlebotominae are represented by two genera each in the studied material, Palaeoparasycorax globosus gen. et sp. n., Palaeoparasycorax suppus gen. et sp. n., Parasycorax simplex sp. n., and Phlebotomites aphoe sp. n. and Phlebotomus vetus sp. n., respectively. Bruchomyiinae is represented by Nemopalpus quadrispiculatus sp. n. Furthermore, one genus of an incertae sedis subfamily, Bamara groehni gen. et sp. n., is described. The systematic positions of the new taxa are discussed.

Assessing the phylogeographic history of the montane caddisfly Thremma gallicum using mitochondrial and restriction-site-associated DNA (RAD) markers.

Repeated Quaternary glaciations have significantly shaped the present distribution and diversity of several European species in aquatic and terrestrial habitats. To study the phylogeography of freshwater invertebrates, patterns of intraspecific variation have been examined primarily using mitochondrial DNA markers that may yield results unrepresentative of the true species history. Here, population genetic parameters were inferred for a montane aquatic caddisfly, Thremma gallicum, by sequencing a 658-bp fragment of the mitochondrial CO1 gene, and 12,514 nuclear RAD loci. T. gallicum has a highly disjunct distribution in southern and central Europe, with known populations in the Cantabrian Mountains, Pyrenees, Massif Central, and Black Forest. Both datasets represented rangewide sampling of T. gallicum. For the CO1 dataset, this included 352 specimens from 26 populations, and for the RAD dataset, 17 specimens from eight populations. We tested 20 competing phylogeographic scenarios using approximate Bayesian computation (ABC) and estimated genetic diversity patterns. Support for phylogeographic scenarios and diversity estimates differed between datasets with the RAD data favouring a southern origin of extant populations and indicating the Cantabrian Mountains and Massif Central populations to represent highly diverse populations as compared with the Pyrenees and Black Forest populations. The CO1 data supported a vicariance scenario (north-south) and yielded inconsistent diversity estimates. Permutation tests suggest that a few hundred polymorphic RAD SNPs are necessary for reliable parameter estimates. Our results highlight the potential of RAD and ABC-based hypothesis testing to complement phylogeographic studies on non-model species.