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.

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.

Bioturbation enhances the aerobic respiration of lake sediments in warming lakes.

While lakes occupy less than 2% of the total surface of the Earth, they play a substantial role in global biogeochemical cycles. For instance, shallow lakes are important sites of carbon metabolism. Aerobic respiration is one of the important drivers of the carbon metabolism in lakes. In this context, bioturbation impacts of benthic animals (biological reworking of sediment matrix and ventilation of the sediment) on sediment aerobic respiration have previously been underestimated. Biological activity is likely to change over the course of a year due to seasonal changes of water temperatures. This study uses microcosm experiments to investigate how the impact of bioturbation (by Diptera, Chironomidae larvae) on lake sediment respiration changes when temperatures increase. While at 5°C, respiration in sediments with and without chironomids did not differ, at 30°C sediment respiration in microcosms with 2000 chironomids per m(2) was 4.9 times higher than in uninhabited sediments. Our results indicate that lake water temperature increases could significantly enhance lake sediment respiration, which allows us to better understand seasonal changes in lake respiration and carbon metabolism as well as the potential impacts of global warming.

A new genus of Podonominae (Diptera: Chironomidae) in Late Eocene Rovno amber from Ukraine.

The genus Palaeoboreochlus Baranov et Andersen, n. gen. is erected based on P. inornatus Baranov et Andersen, n. sp. described from a male found in Late Eocene Rovno amber. The new genus groups with Boreochlus Edwards in the tribe "Boreochlini".

New records of immature aquatic Diptera from the Foulden Maar Fossil-Lagerstätte, New Zealand, and their biogeographic implications.

Background: The biogeographical and ecological history of true flies (Diptera) in New Zealand is little known due to a scarcity of fossil specimens. Here, we report a fauna of immature aquatic dipterans from freshwater diatomites of the early Miocene Foulden Maar Fossil-Lagerstätte in southern New Zealand. Methods: We document 30 specimens of immature dipterans, mostly pupae, and compare their external morphology to extant aquatic Diptera. Based on the reconstructed paleoenvironment of Foulden Maar, we discuss taxonomic, ecological and taphonomic implications of this early Miocene fauna. Results: Among Chironomidae, one pupal morphotype is attributed to Tanypodinae, one pupal morphotype and one larval morphotype are placed into Chironomus (Chironominae) and a further morphotype into Chironominae incertae sedis. Chaoboridae are represented by a pupal morphotype congeneric or very close to the extant Chaoborus, today globally distributed except for New Zealand. Additional immature specimens are likely larvae and puparia of brachyceran flies but cannot be identified to a narrower range. These finds document an aquatic dipteran fauna in New Zealand in the earliest Miocene and highlight Neogene extinction as a factor in shaping the extant Diptera fauna in New Zealand. Immature aquatic dipterans were a common and likely ecologically important component of the early Miocene Foulden Maar lake. Preservation of larvae and pupae may have been promoted by diatomaceous microbial mats and the light colour of the diatomite likely facilitated spotting of these minute fossils in the field.

Long-Term Chironomid Emergence at a Karst Tufa Barrier in Plitvice Lakes National Park, Croatia.

Chironomids are found in all types of freshwater habitats; they are a ubiquitous and highly diverse group of aquatic insects. Plitvice Lakes National Park is the oldest and largest national park in Croatia and consists of numerous and diverse freshwater habitats, making the area an ideal location for long-term research into the chironomid emergence patterns and phenology. The main objectives of this study were to identify the composition of the chironomid community, determine the phenology of the identified species, and assess the main factors influencing their emergence in Plitvice Lakes. During 14 years of research, more than 13,000 chironomids belonging to more than 80 species were recorded. The most abundant species was found to be Parametriocnemus stylatus. The highest abundance of chironomids was recorded in lotic habitats with faster water current over substrates of moss and algae and pebbles. Water temperature and the availability of organic matter were found to be the main factors that drive chironomid emergence at the tufa barrier studied. In the last years of this study, a prolonged flight period was observed. Although this is not statistically significant (at this stage of the study), it could be due to a higher water temperature in winter.

Poxyaibamberus Andersen & Dantas, gen. nov. (Diptera, Chironomidae, Orthocladiinae) from Brazil.

Poxyaibamberus Andersen & Dantas, gen. nov. is erected based on the males of two species, P.jamanximensis Andersen & Dantas, sp. nov. from Jamanxim National Park, Pará State, Brazil, and P.ubajarensis Andersen & Dantas, sp. nov. from Ubajara National Park, Ceará State, Brazil. Both species have a comparatively short and wide head, with large eyes and short, five-segmented palps; a strong subapical seta on the ultimate flagellomere; scalpellate acrostichals; no setae on the wing veins except for one seta on the brachiolum; a long costal extension; and a large triangular anal point and a very long heel on the gonostylus. The systematic position of the new genus is briefly discussed.

Expanding the Mesozoic Record of Early Brachyceran Fly Larvae, including New Larval Forms with Chimera-Type Morphologies.

Diptera are one of the four megadiverse groups of holometabolan insects. Flies perform numerous ecological functions, especially in their larval stages. We can assume that this was already the case in the past; however, fly larvae remain rare in most deposits. Here we report new dipteran larvae preserved in Cretaceous (about 99 Ma) Kachin amber from Myanmar and, even older, Jurassic (about 165 Ma) compression fossils from China. Through light microscopy and micro-CT scanning we explore their peculiar morphology and discuss their possible phylogenetic affinities. Several larvae seem to represent the lineage of Stratiomyomorpha. A few others present characters unique to Xylophagidae (awl-flies), as well as to Athericidae (water sniper-flies), resulting in a chimeric morphology. Understanding the exact relationships of most of these specimens with a particular lineage remains challenging, since they differ considerably from any other known dipteran larvae and present some unique traits. Additionally, we report new specimens of Qiyia jurassica Chen et al., 2014, supposedly parasitic larvae, most likely representatives of Athericidae. These new findings offer valuable insights into the evolution of the early diversification of the brachyceran flies and underscore the importance of immature stages in understanding the evolutionary history and ecology of flies.

Beneath the surface: Decoding the impact of Chironomus riparius bioturbation on microplastic dispersion in sedimentary matrix.

A detailed understanding of microplastics (MPs) behaviour in freshwater ecosystems is crucial for a proper ecological assessment. This includes the identification of significant transport pathways and net accumulation zones, considering their inherent, and already proven influence on aquatic ecosystems. Bioavailability of toxic agents is significantly influenced by macroinvertebrates' behaviour, such as bioturbation and burrowing, and their prior exposure history. This study investigates the effect of bioturbation activity of Chironomus riparius Meigen, 1804 on the vertical transfer of polyethylene MPs ex-situ. The experimental setup exposes larvae to a scenario of 10× the environmentally relevant high concentration of MPs (80 g m-2). Bioturbation activity was estimated using sediment profile imaging with luminophore tracers. This study demonstrated that spherical MPs are vertically transferred in the sediment due to the bioturbation activity of C. riparius larvae and that their presence influences the intensity of the bioturbation activity over time. The present findings provide a noteworthy contribution to the understanding of the relationship between ecosystem engineers and the dispersion and accumulation of MPs within freshwater ecosystems.

Scratching the tip of the iceberg: integrative taxonomy reveals 30 new species records of Microgastrinae (Braconidae) parasitoid wasps for Germany, including new Holarctic distributions.

Substantial parts of the European and German insect fauna still remain largely unexplored, the so-called "dark taxa". In particular, midges (Diptera) and parasitoid wasps (Hymenoptera) are abundant and species-rich throughout Europe, yet are often neglected in biodiversity research. One such dark taxon is Microgastrinae wasps (Hymenoptera: Braconidae), a group of parasitoids of lepidopteran caterpillars with 252 species reported in Germany so far. As part of the German Barcode of Life Project GBOL III: Dark Taxa, reverse DNA barcoding and integrative taxonomic approaches were used to shed some light on the German Fauna of Microgastrinae wasps. In our workflow, DNA barcoding was used for molecular clustering of our specimens in a first step, morphological examination of the voucher specimens in a second step, and host data compared in a third step. Here, 30 species are reported for the first time in Germany, adding more than 10% to the known German fauna. Information for four species is provided in a new Holarctic context, reporting them for the Nearctic or, respectively, Palaearctic region, and 26 additional country records are added from sequenced material available in the collections accessible to us. Molecular clusters that show signs of discrepancies are discussed. Results show that we are just scratching the tip of the iceberg of the unexplored Microgastrinae diversity in Germany.

Diagnostic clues for identification of selected species of the Micropsectra atrofasciata group, with description of M. uva sp. nov. from Croatia (Diptera: Chironomidae: Tanytarsini).

Micropsectra uva sp. nov. is described from the Plitvice Lakes National Park (Croatia), and placed in the Micropsectra atrofasciata systematic species group. Morphological key structures/characters for adult males of the new species and several closest Micropsectra Kieffer are illustrated in detail and evaluated.

Facing the infinity: tackling large samples of challenging Chironomidae (Diptera) with an integrative approach.

Background: Integrative taxonomy is becoming ever more significant in biodiversity research as scientists are tackling increasingly taxonomically challenging groups. Implementing a combined approach not only guarantees more accurate species identification, but also helps overcome limitations that each method presents when applied on its own. In this study, we present one application of integrative taxonomy for the highly abundant and particularly diverse fly taxon Chironomidae (Diptera). Although non-biting midges are key organisms in merolimnic systems, they are often cast aside in ecological surveys because they are very challenging to identify and extremely abundant. Methods: Here, we demonstrate one way of applying integrative methods to tackle this highly diverse taxon. We present a three-level subsampling method to drastically reduce the workload of bulk sample processing, then apply morphological and molecular identification methods in parallel to evaluate species diversity and to examine inconsistencies across methods. Results: Our results suggest that using our subsampling approach, identifying less than 10% of a sample's contents can reliably detect >90% of its diversity. However, despite reducing the processing workload drastically, the performance of our taxonomist was affected by mistakes, caused by large amounts of material. We conducted misidentifications for 9% of vouchers, which may not have been recovered had we not applied a second identification method. On the other hand, we were able to provide species information in cases where molecular methods could not, which was the case for 14% of vouchers. Therefore, we conclude that when wanting to implement non-biting midges into ecological frameworks, it is imperative to use an integrative approach.

Extreme freshwater discharge events exacerbated by climate change influence the structure and functional response of the chironomid community in a biodiversity hotspot.

Global climate change is expected to exacerbate extreme discharge events in freshwater ecosystems as a consequence of changes in precipitation volume and snow cover duration. Chironomid midges were used as a model organism in this study because of their small size and short life cycles, which enable fast colonization of new habitats and great resilience. This resilience is often expressed in easy recolonization after an extreme event. Chironomid samples together with physico-chemical water measurements were collected for 14 years, between 2007 and 2020, in a karst tufa barrier that is part of the Plitvice Lakes National Park in Croatia. More than 13,000 individuals belonging to >90 taxa were collected. Mean annual water temperature increased by 0.1 °C during this period. Multiple change-point analysis revealed three main periods by discharge patterns: the first one from January 2007 to June 2010, the second from July 2010 to March 2013, characterised by extreme low discharge, and the third from April 2013 to December 2020, characterised by an increase in extreme peak discharge values. Based on multilevel pattern analysis, indicator species of the first and the third discharge period were detected. The ecological preferences of these species indicate an environmental change related to the changes in discharge. Along with species composition, functional composition has changed with the abundance of passive filtrators, shredders and predators increasing over time. Species richness and abundance did not change over the period of observation, thus emphasizing the importance of species-level identification in detecting the earliest community response to change that would otherwise be overlooked.

New species and records of Pseudochironomini Sæther, 1977 (Diptera, Chironomidae) from the Dominican Republic.

Background: Pseudochironomini is a relatively small and poorly-studied tribe of subfamily Chironominae (Diptera, Chironomidae). New information: Pseudochironomusruthae Andersen & Baranov sp. nov. is described and figured, based on a single male collected in a light trap at Matadero, Dominican Republic. The species can be separated from its congeners by the combination of the following characters: wing without dark bands, dorsocentrals in partly double row and apex of superior volsella rounded. The species is the first Pseudochironomus species to be formally recorded and described from the Caribbean. In addition, a new record of Manoapahayokeensis Jacobsen & Perry, 2002 from the Dominican Republic is given. One specimen was DNA-barcoded and the barcode is given.

The Morphological Diversity of Dragon Lacewing Larvae (Nevrorthidae, Neuroptera) Changed More over Geological Time Scales Than Anticipated.

Nevrorthidae, the group of dragon lacewings, has often been considered a relic group. Today, dragon lacewings show a scattered distribution, with some species occurring in southern Europe, Japan, Australia, and one in China. The idea that this distribution is only a remnant of an originally larger distribution is further supported by fossils of the group preserved in ambers from the Baltic region (Eocene, ca. 35-40 MaBP) and Myanmar (Kachin amber, Cretaceous, ca. 100 MaBP). Larvae of the group are slender and elongated and live mostly in water. Yet, larvae are in fact very rare. So far, only slightly more than 30 larval specimens, counting all extant and fossil larvae, have been depicted in the literature. Here, we report numerous additional specimens, including extant larvae, but also fossil ones from Baltic and Kachin amber. Together with the already known ones, this sums up to over 100 specimens. We analysed quantitative aspects of the morphology of these larvae and compared them over time to identify changes in the diversity. Despite the enriched sample size, the data set is still unbalanced, with, for example, newly hatched larvae (several dozen specimens) only known from the Eocene. We expected little change in larval morphology over geological time, as indicated by earlier studies. However, on the contrary, we recognised morphologies present in fossils that are now extinct. This result is similar to those for other groups of lacewings which have a relic distribution today, as these have also suffered a loss in diversity in larval forms.

The first fossil immature of Elmidae: an unusual riffle beetle larva preserved in Baltic amber.

Elmidae, riffle beetles, have both adult and immature stages that show specializations for water environments. Fossils of adults of Elmidae are already known from amber, however a record of immatures was so far lacking. We report here the first fossil larva of Elmidae, preserved in Baltic amber. To be able to access details of the body hidden by inclusions and "Verlumung" we conducted, in addition to optical documentation methods, micro-CT and synchrotron documentation methods. The larva is characterised by prominent dorso-lateral and lateral processes and a plate-like ventral operculum at the end of the abdomen. The new fossil has similarities in the general body shape and the prominent characters with some modern larvae of Elmidae. The posterior protrusions on the trunk end possibly represent gills, which would imply that fossil larvae of Elmidae also led a water-related life style similar to modern representatives.

Peering into the Darkness: DNA Barcoding Reveals Surprisingly High Diversity of Unknown Species of Diptera (Insecta) in Germany.

Determining the size of the German insect fauna requires better knowledge of several megadiverse families of Diptera and Hymenoptera that are taxonomically challenging. This study takes the first step in assessing these "dark taxa" families and provides species estimates for four challenging groups of Diptera (Cecidomyiidae, Chironomidae, Phoridae, and Sciaridae). These estimates are based on more than 48,000 DNA barcodes (COI) from Diptera collected by Malaise traps that were deployed in southern Germany. We assessed the fraction of German species belonging to 11 fly families with well-studied taxonomy in these samples. The resultant ratios were then used to estimate the species richness of the four "dark taxa" families (DT families hereafter). Our results suggest a surprisingly high proportion of undetected biodiversity in a supposedly well-investigated country: at least 1800-2200 species await discovery in Germany in these four families. As this estimate is based on collections from one region of Germany, the species count will likely increase with expanded geographic sampling.

A Lagerstätte from Australia provides insight into the nature of Miocene mesic ecosystems.

Reduced precipitation in the Miocene triggered the geographic contraction of rainforest ecosystems around the world. In Australia, this change was particularly pronounced; mesic rainforest ecosystems that once dominated the landscape transformed into the shrublands, grasslands, and deserts of today. A lack of well-preserved fossils has made it difficult to understand the nature of Australian ecosystems before the aridification. Here, we report on an exceptionally well-preserved rainforest biota from New South Wales, Australia. This Konservat-Lagerstätte hosts a rich diversity of microfossils, plants, insects, spiders, and vertebrate remains preserved in goethite. We document evidence for several species interactions including predation, parasitism, and pollination. The fossils are indicative of an oxbow lake in a mesic rainforest and suggest that rainforest distributions have shifted since the Miocene. The variety of fossils preserved, together with high fidelity of preservation, allows for unprecedented insights into the mesic ecosystems that dominated Australia during the Miocene.

New extreme morphologies as exemplified by 100 million-year-old lacewing larvae.

Larvae of the group Holometabola (beetles, wasps, flies, moths and others) differ significantly in their morphology from their corresponding adults. In most larvae, appendages and other structures protruding from the body (antennae, palps, legs, trunk processes) appear less elongate than in their corresponding adults, providing the impression that these larvae are restricted to a certain degree in developing more elongate structures. We provide here numerous counterexamples of larvae of lacewings (Neuroptera). These include different forms of elongated antennae, mandibles, maxillae, labial palps, legs, trunk processes and neck regions. Most of these examples are larvae preserved in different types of 100 million-year-old amber. The longest neck region was found in an extant specimen. All these examples demonstrate that certain branches of Neuroptera indeed had larval forms that possessed strongly elongated structures. Hence there is no principal constraint that hinders holometabolan larvae to develop such structures.

Global Size Pattern in a Group of Important Ecological Indicators (Diptera, Chironomidae) Is Driven by Latitudinal Temperature Gradients.

Size is one of the most outwardly obvious characteristics of animals, determined by multiple phylogenetic and environmental variables. Numerous hypotheses have been suggested to explain the relationship between the body size of animals and their geographic latitude. Bergmann's Rule, describing a positive relationship between the body size of endothermic animals and their geographic latitude, is especially well known. Whether or not insects exhibit a similar pattern has long been a subject for debate. We hypothesize that latitudinal size gradients are coupled to temperature variation affecting the metabolic rate of these merolimnic insects. We showcase a strong latitudinal size gradient in non-biting midges (Diptera: Chironomidae), based on the examination of 4309 specimens of these midges from around the world. Although phylogenetic position was a key predictor of wing length, we also found that wing length decreases by 32.4 µm per every 1 °C of mean annual temperature increase. This pattern was found across different taxa and could be detected in 20 of 24 genera studied. We discuss the reasons for this pattern origin and its palaeoecological implications.