Towards global insect biomonitoring with frugal methods.

None of the global targets for protecting nature are currently met, although humanity is critically dependent on biodiversity. A significant issue is the lack of data for most biodiverse regions of the planet where the use of frugal methods for biomonitoring would be particularly important because the available funding for monitoring is insufficient, especially in low-income countries. We here discuss how three approaches to insect biomonitoring (computer vision, lidar, DNA sequences) could be made more frugal and urge that all biomonitoring techniques should be evaluated for global suitability before becoming the default in high-income countries. This requires that techniques popular in high-income countries should undergo a phase of 'innovation through simplification' before they are implemented more broadly. We predict that techniques that acquire raw data at low cost and are suitable for analysis with AI (e.g. images, lidar-signals) will be particularly suitable for global biomonitoring, while techniques that rely heavily on patented technologies may be less promising (e.g. DNA sequences). We conclude the opinion piece by pointing out that the widespread use of AI for data analysis will require a global strategy for providing the necessary computational resources and training. This article is part of the theme issue 'Towards a toolkit for global insect biodiversity monitoring'.

Lidar as a potential tool for monitoring migratory insects.

The seasonal migrations of insects involve a substantial displacement of biomass with significant ecological and economic consequences for regions of departure and arrival. Remote sensors have played a pivotal role in revealing the magnitude and general direction of bioflows above 150 m. Nevertheless, the takeoff and descent activity of insects below this height is poorly understood. Our lidar observations elucidate the low-height dusk movements and detailed information of insects in southern Sweden from May to July, during the yearly northward migration period. Importantly, by filtering out moths from other insects based on optical information and wingbeat frequency, we have introduced a promising new method to monitor the flight activities of nocturnal moths near the ground, many of which participate in migration through the area. Lidar thus holds the potential to enhance the scientific understanding of insect migratory behavior and improve pest control strategies.

Discrimination of Hover Fly Species and Sexes by Wing Interference Signals.

Remote automated surveillance of insect abundance and diversity is poised to revolutionize insect decline studies. The study reveals spectral analysis of thin-film wing interference signals (WISs) can discriminate free-flying insects beyond what can be accomplished by machine vision. Detectable by photonic sensors, WISs are robust indicators enabling species and sex identification. The first quantitative survey of insect wing thickness and modulation through shortwave-infrared hyperspectral imaging of 600 wings from 30 hover fly species is presented. Fringy spectral reflectance of WIS can be explained by four optical parameters, including membrane thickness. Using a Naïve Bayes Classifier with five parameters that can be retrieved remotely, 91% is achieved accuracy in identification of species and sexes. WIS-based surveillance is therefore a potent tool for remote insect identification and surveillance.

Spatial monitoring of flying insects over a Swedish lake using a continuous-wave lidar system.

We have used a continuous-wave bi-static lidar system based on the Scheimpflug principle in measurements on flying insects above, and in the vicinity of, a small lake located in a forested area in Southern Sweden. The system, which operates on triangulation principles, has a high spatial resolution at close distance, followed by a subsequent decline in resolution further from the sensor, related to the compact system design with a separation of transmitter and receiver by only 0.81 m. Our study showed a strong increase in insect abundance especially at dusk, but also at dawn. Insect numbers decreased over water compared to over land, and larger insects were over-represented over water. Further, the average size of the insects increased at night compared to day time.

Dual-Band Infrared Scheimpflug Lidar Reveals Insect Activity in a Tropical Cloud Forest.

We describe an entomological dual-band 808 and 980 nm lidar system which has been implemented in a tropical cloud forest (Ecuador). The system was successfully tested at a sample rate of 5 kHz in a cloud forest during challenging foggy conditions (extinction coefficients up to 20 km-1). At times, the backscattered signal could be retrieved from a distance of 2.929 km. We present insect and bat observations up to 200 m during a single night with an emphasis on fog aspects, potentials, and benefits of such dual-band systems. We demonstrate that the modulation contrast between insects and fog is high in the frequency domain compared to intensity in the time domain, thus allowing for better identification and quantification in misty forests. Oscillatory lidar extinction effects are shown in this work for the first time, caused by the combination of dense fog and large moths partially obstructing the beam. We demonstrate here an interesting case of a moth where left- and right-wing movements induced oscillations in both intensity and pixel spread. In addition, we were able to identify the dorsal and ventral sides of the wings by estimating the corresponding melanization with the dual-band lidar. We demonstrate that the wing beat trajectories in the dual-band parameter space are complementary rather than covarying or redundant, thus a dual-band entomological lidar approach to biodiversity studies is feasible in situ and endows species specificity differentiation. Future improvements are discussed. The introduction of these methodologies opens the door to a wealth of possible experiments to monitor, understand, and safeguard the biological resources of one of the most biodiverse countries on Earth.

Remote Nanoscopy with Infrared Elastic Hyperspectral Lidar.

Monitoring insects of different species to understand the factors affecting their diversity and decline is a major challenge. Laser remote sensing and spectroscopy offer promising novel solutions to this. Coherent scattering from thin wing membranes also known as wing interference patterns (WIPs) have recently been demonstrated to be species specific. The colors of WIPs arise due to unique fringy spectra, which can be retrieved over long distances. To demonstrate this, a new concept of infrared (950-1650 nm) hyperspectral lidar with 64 spectral bands based on a supercontinuum light source using ray-tracing and 3D printing is developed. A lidar with an unprecedented number of spectral channels, high signal-to-noise ratio, and spatio-temporal resolution enabling detection of free-flying insects and their wingbeats. As proof of principle, coherent scatter from a damselfly wing at 87 m distance without averaging (4 ms recording) is retrieved. The fringed signal properties are used to determine an effective wing membrane thickness of 1412 nm with ±4 nm precision matching laboratory recordings of the same wing. Similar signals from free flying insects (2 ms recording) are later recorded. The accuracy and the method's potential are discussed to discriminate species by capturing coherent features from free-flying insects.

A biophotonic platform for quantitative analysis in the spatial, spectral, polarimetric, and goniometric domains.

Advanced instrumentation and versatile setups are needed for understanding light interaction with biological targets. Such instruments include (1) microscopes and 3D scanners for detailed spatial analysis, (2) spectral instruments for deducing molecular composition, (3) polarimeters for assessing structural properties, and (4) goniometers probing the scattering phase function of, e.g., tissue slabs. While a large selection of commercial biophotonic instruments and laboratory equipment are available, they are often bulky and expensive. Therefore, they remain inaccessible for secondary education, hobbyists, and research groups in low-income countries. This lack of equipment impedes hands-on proficiency with basic biophotonic principles and the ability to solve local problems with applied physics. We have designed, prototyped, and evaluated the low-cost Biophotonics, Imaging, Optical, Spectral, Polarimetric, Angular, and Compact Equipment (BIOSPACE) for high-quality quantitative analysis. BIOSPACE uses multiplexed light-emitting diodes with emission wavelengths from ultraviolet to near-infrared, captured by a synchronized camera. The angles of the light source, the target, and the polarization filters are automated by low-cost mechanics and a microcomputer. This enables multi-dimensional scatter analysis of centimeter-sized biological targets. We present the construction, calibration, and evaluation of BIOSPACE. The diverse functions of BIOSPACE include small animal spectral imaging, measuring the nanometer thickness of a bark-beetle wing, acquiring the scattering phase function of a blood smear and estimating the anisotropic scattering and the extinction coefficients, and contrasting muscle fibers using polarization. We provide blueprints, component list, and software for replication by enthusiasts and educators to simplify the hands-on investigation of fundamental optical properties in biological samples.

Weak population genetic structure in Eurasian spruce bark beetle over large regional scales in Sweden.

The Eurasian spruce bark beetle, Ips typographus, is a major pest, capable of killing spruce forests during large population outbreaks. Recorded dispersal distances of individual beetles are typically within hundreds of meters or a few kilometers. However, the connectivity between populations at larger distances and longer time spans and how this is affected by the habitat is less studied, despite its importance for understanding at which distances local outbreaks may spread. Previous population genetic studies in I. typographus typically used low resolution markers. Here, we use genome-wide data to assess population structure and connectivity of I. typographus in Sweden. We used 152 individuals from 19 population samples, distributed over 830 km from Strömsund (63° 46' 8″ N) in the north to Nyteboda (56° 8' 50″ N) in the south, to capture processes at a large regional scale, and a transect sampling design adjacent to a recent outbreak to capture processes at a smaller scale (76 km). Using restriction site-associated DNA sequencing (RADseq) markers capturing 1409-1997 SNPs throughout the genome, we document a weak genetic structure over the large scale, potentially indicative of high connectivity with extensive gene flow. No differentiation was detected at the smaller scale. We find indications of isolation-by-distance both for relative (F ST) and absolute divergence (Dxy). The two northernmost populations are most differentiated from the remaining populations, and diverge in parallel to the southern populations for a set of outlier loci. In conclusion, the population structure of I. typographus in Sweden is weak, suggesting a high capacity to disperse and establish outbreak populations in new territories.

Potential for identification of wild night-flying moths by remote infrared microscopy.

There are hundreds of thousands of moth species with crucial ecological roles that are often obscured by their nocturnal lifestyles. The pigmentation and appearance of moths are dominated by cryptic diffuse shades of brown. In this study, 82 specimens representing 26 moth species were analysed using infrared polarimetric hyperspectral imaging in the range of 0.95-2.5 µm. Contrary to previous studies, we demonstrate that since infrared light does not resolve the surface roughness, wings appear glossy and specular at longer wavelengths. Such properties provide unique reflectance spectra between species. The reflectance of the majority of our species could be explained by comprehensive models, and a complete parametrization of the spectral, polarimetric and angular optical properties was reduced to just 11 parameters with physical units. These parameters are complementary and, compared with the within-species variation, were significantly distinct between species. Counterintuitively to the aperture-limited resolution criterion, we could deduce microscopic features along the surface from their infrared properties. These features were confirmed by electron microscopy. Finally, we show how our findings could greatly enhance opportunities for remote identification of free-flying moth species, and we hypothesize that such flat specular wing targets could be expected to be sensed over considerable distances.

Automating insect monitoring using unsupervised near-infrared sensors.

Insect monitoring is critical to improve our understanding and ability to preserve and restore biodiversity, sustainably produce crops, and reduce vectors of human and livestock disease. Conventional monitoring methods of trapping and identification are time consuming and thus expensive. Automation would significantly improve the state of the art. Here, we present a network of distributed wireless sensors that moves the field towards automation by recording backscattered near-infrared modulation signatures from insects. The instrument is a compact sensor based on dual-wavelength infrared light emitting diodes and is capable of unsupervised, autonomous long-term insect monitoring over weather and seasons. The sensor records the backscattered light at kHz pace from each insect transiting the measurement volume. Insect observations are automatically extracted and transmitted with environmental metadata over cellular connection to a cloud-based database. The recorded features include wing beat harmonics, melanisation and flight direction. To validate the sensor's capabilities, we tested the correlation between daily insect counts from an oil seed rape field measured with six yellow water traps and six sensors during a 4-week period. A comparison of the methods found a Spearman's rank correlation coefficient of 0.61 and a p-value = 0.0065, with the sensors recording approximately 19 times more insect observations and demonstrating a larger temporal dynamic than conventional yellow water trap monitoring.

Real-time dispersal of malaria vectors in rural Africa monitored with lidar.

Lack of tools for detailed, real-time observation of mosquito behavior with high spatio-temporal resolution limits progress towards improved malaria vector control. We deployed a high-resolution entomological lidar to monitor a half-kilometer static transect positioned over rice fields outside a Tanzanian village. A quarter of a million in situ insect observations were classified, and several insect taxa were identified based on their modulation signatures. We observed distinct range distributions of male and female mosquitoes in relation to the village periphery, and spatio-temporal behavioral features, such as swarming. Furthermore, we observed that the spatial distributions of males and females change independently of each other during the day, and were able to estimate the daily dispersal of mosquitoes towards and away from the village. The findings of this study demonstrate how lidar-based monitoring could dramatically improve our understanding of malaria vector ecology and control options.

Advances in automatic identification of flying insects using optical sensors and machine learning.

Worldwide, farmers use insecticides to prevent crop damage caused by insect pests, while they also rely on insect pollinators to enhance crop yield and other insect as natural enemies of pests. In order to target pesticides to pests only, farmers must know exactly where and when pests and beneficial insects are present in the field. A promising solution to this problem could be optical sensors combined with machine learning. We obtained around 10,000 records of flying insects found in oilseed rape (Brassica napus) crops, using an optical remote sensor and evaluated three different classification methods for the obtained signals, reaching over 80% accuracy. We demonstrate that it is possible to classify insects in flight, making it possible to optimize the application of insecticides in space and time. This will enable a technological leap in precision agriculture, where focus on prudent and environmentally-sensitive use of pesticides is a top priority.

Bark beetles as lidar targets and prospects of photonic surveillance.

Forestry is raising concern about the outbreaks of European spruce bark beetle, Ips typographus, causing extensive damage to the spruce forest and timber values. Precise monitoring of these beetles is a necessary step towards preventing outbreaks. Current commercial monitoring methods are catch-based and lack in both temporal and spatial resolution. In this work, light scattering from beetles is characterized, and the feasibility of entomological lidar as a tool for long-term monitoring of bark beetles is explored. Laboratory optical properties, wing thickness, and wingbeat frequency of bark beetles are reported, and these parameters can infer target identity in lidar data. Lidar results from a Swedish forest with controlled bark beetle release event are presented. The capability of lidar to simultaneously monitor both insects and a pheromone plume mixed with chemical smoke governing the dispersal of many insects is demonstrated. In conclusion, entomological lidar is a promising tool for monitoring bark beetles.

Lidar reveals activity anomaly of malaria vectors during pan-African eclipse.

Yearly, a quarter billion people are infected and a half a million killed by the mosquito-borne disease malaria. Lack of real-time observational tools for continuously assessing the unperturbed mosquito flight activity in situ limits progress toward improved vector control. We deployed a high-resolution entomological lidar to monitor a half-kilometer static transect adjacent to a Tanzanian village. We evaluated one-third million insect observations during five nights, four days, and one annular solar eclipse. We demonstrate in situ lidar classification of several insect families and their sexes based on their modulation signatures. We were able to compare the fine-scale spatiotemporal activity patterns of malaria vectors during ordinary days and an eclipse to disentangle phototactic activity patterns from the circadian mechanism. We observed an increased insect activity during the eclipse attributable to mosquitoes. These unprecedented findings demonstrate how lidar-based monitoring of distinct mosquito activities could advance our understanding of vector ecology.

Atmospheric CO2 sensing using Scheimpflug-lidar based on a 1.57-µm fiber source.

A molecular laser-radar system, based on the Scheimpflug principle, has been constructed and demonstrated for remote sensing of atmospheric CO2 concentrations using Differential Absorption Lidar (DIAL) in the (30012←00001) absorption band. The laser source is a Continues Wave (CW) Distributed-FeedBack (DFB) diode laser seeding an Erbium-doped fiber amplifier, emitting narrowband (3 MHz) tunable radiation with an output power of 1.3 W at 1.57 µm. The laser beam is expanded and transmitted to the atmosphere. The atmospheric backscattered signal is collected with a Newtonian telescope and detected with a linear InGaAs array detector satisfying the Scheimpflug condition. We present range-resolved measurements of atmospheric CO2 concentration from a test range of 2 km located in the city of Lund, Sweden. We discuss and provide scalable results for CO2 profiling with the Scheimpflug-lidar method.

1.57 µm fiber source for atmospheric CO2 continuous-wave differential absorption lidar.

We present an efficient fiber source designed for continuous-wave differential absorption light detection and ranging (CW DIAL) of atmospheric CO2-concentration. It has a linewidth of 3 MHz, a tuning range of 2 nm over the CO2 absorption peaks at 1.572 µm, and an output power of 1.3 W limited by available pump power. Results from the initial CW DIAL testing are also presented and discussed.

Scheimpflug Lidar for combustion diagnostics.

A portable Lidar system developed for large-scale (~1-20 m) combustion diagnostics is described and demonstrated. The system is able to perform remote backscattering measurements with range and temporal resolution. The range resolution is obtained by sharply imaging a part of the laser beam onto a CMOS-array or ICCD detector. The large focal depth required to do this is attained by placing the laser beam, the collection optics and the detector in a so-called Scheimpflug configuration. Results from simulations of the range capabilities and range resolution of the system are presented and its temporal resolution is also discussed. Various applications, important for combustion diagnostics, are also demonstrated, including Rayleigh scattering thermometry, aerosol detection and laser-induced fluorescence measurements. These measurements have been carried out using various continuous-wave GaN diode lasers, emitting in the violet-blue (405 - 450 nm) wavelength regime. It is anticipated that Scheimpflug Lidar will provide a useful and versatile diagnostic tool for combustion research, not only for fundamental studies, but in particular for applications at industrial sites.

Can the narrow red bands of dragonflies be used to perceive wing interference patterns?

Despite numerous studies of selection on position and number of spectral vision bands, explanations to the function of narrow spectral bands are lacking. We investigate dragonflies (Odonata), which have the narrowest spectral bands reported, in order to investigate what features these narrow spectral bands may be used to perceive. We address whether it is likely that narrow red bands can be used to identify conspecifics by the optical signature from wing interference patterns (WIPs). We investigate the optical signatures of Odonata wings using hyperspectral imaging, laser profiling, ellipsometry, polarimetric modulation spectroscopy, and laser radar experiments. Based on results, we estimate the prospects for Odonata perception of WIPs to identify conspecifics in the spectral, spatial, intensity, polarization, angular, and temporal domains. We find six lines of evidence consistent with an ability to perceive WIPs. First, the wing membrane thickness of the studied Odonata is 2.3 µm, coinciding with the maximal thickness perceivable by the reported bandwidth. Second, flat wings imply that WIPs persist from whole wings, which can be seen at a distance. Third, WIPs constitute a major brightness in the visual environment only second after the solar disk. Fourth, WIPs exhibit high degree of polarization and polarization vision coincides with frontal narrow red bands in Odonata. Fifth, the angular light incidence on the Odonata composite eye provides all prerequisites for direct assessment of the refractive index which is associated with age. Sixth, WIPs from conspecifics in flight make a significant contribution even to the fundamental wingbeat frequency within the flicker fusion bandwidth of Odonata vision. We conclude that it is likely that WIPs can be perceived by the narrow red bands found in some Odonata species and propose future behavioral and electrophysiological tests of this hypothesis.

The bat-bird-bug battle: daily flight activity of insects and their predators over a rice field revealed by high-resolution Scheimpflug Lidar.

We present the results of, to our knowledge, the first Lidar study applied to continuous and simultaneous monitoring of aerial insects, bats and birds. It illustrates how common patterns of flight activity, e.g. insect swarming around twilight, depend on predation risk and other constraints acting on the faunal components. Flight activity was monitored over a rice field in China during one week in July 2016, using a high-resolution Scheimpflug Lidar system. The monitored Lidar transect was about 520 m long and covered approximately 2.5 m3. The observed biomass spectrum was bimodal, and targets were separated into insects and vertebrates in a categorization supported by visual observations. Peak flight activity occurred at dusk and dawn, with a 37 min time difference between the bat and insect peaks. Hence, bats started to feed in declining insect activity after dusk and stopped before the rise in activity before dawn. A similar time difference between insects and birds may have occurred, but it was not obvious, perhaps because birds were relatively scarce. Our observations are consistent with the hypothesis that flight activity of bats is constrained by predation in bright light, and that crepuscular insects exploit this constraint by swarming near to sunset/sunrise to minimize predation from bats.

Multiband modulation spectroscopy for the determination of sex and species of mosquitoes in flight.

We present a dual-wavelength polarimetric measurement method to distinguish species and sexes of disease transmitting mosquitoes in flight. By measuring co- and de-polarized backscattered light at 808 and 1550 nm, the degree of linear polarization, wingbeat frequency, reflectance, spectral ratio and glossiness of mosquitoes can be retrieved. Body and wing contributions to these signals can be separated. Whereas the optical cross section is sensitive to the aspect of observation, thus the heading direction of the insect in flight, we demonstrate that polarimetric- and spectral-band ratios are largely invariant to the aspect of observation. We show that wing glossiness, as well as wing- and body-spectral ratios are particularly efficient in distinguishing Anopheles coluzzii and Anopheles arabiensis, 2 closely related species of malaria vectors. Spectral and polarimetric ratios relate to microstructural and melanization features of the wing and body of these species. We conclude that multiband modulation spectroscopy is a useful expansion of the parameter space that can be used to improve the specificity of entomological lidars.