Source imaging method based on diagonal covariance bases and its applications to OPM-MEG.

Magnetoencephalography (MEG) is a noninvasive imaging technique used in neuroscience and clinical research. The source estimation of MEG involves solving a highly underdetermined inverse problem, which requires additional constraints to restrict the solution space. Traditional methods tend to obscure the extent of the sources. However, an accurate estimation of the source extent is important for studying brain activity or preoperatively estimating pathogenic regions. To improve the estimation accuracy of the extended source extent, the spatial constraint of sources is employed in the Bayesian framework. For example, the source is decomposed into a linear combination of validated spatial basis functions, which is proved to improve the source imaging accuracy. In this work, we further construct the spatial properties of the source using the diagonal covariance bases (DCB), which we summarize as the source imaging method SI-DCB. In this approach, specifically, the covariance matrix of the spatial coefficients is modeled as a weighted combination of diagonal covariance basis functions. The convex analysis is used to estimate noise and model parameters under the Bayesian framework. Extensive numerical simulations showed that SI-DCB outperformed five benchmark methods in accurately estimating the location and extent of patch sources. The effectiveness of SI-DCB was verified through somatosensory stimulation experiments performed on a 31-channel OPM-MEG system. The SI-DCB correctly identified the source area where each brain response occurred. The superior performance of SI-DCB suggests that it can provide a template approach for improving the accuracy of source extent estimations under a sparse Bayesian framework.

Magnetoencephalography for Epilepsy Presurgical Evaluation.

PURPOSE OF THE REVIEW: Magnetoencephalography (MEG) is a functional neuroimaging technique that records neurophysiology data with millisecond temporal resolution and localizes it with subcentimeter accuracy. Its capability to provide high resolution in both of these domains makes it a powerful tool both in basic neuroscience as well as clinical applications. In neurology, it has proven useful in its ability to record and localize epileptiform activity. Epilepsy workup typically begins with scalp electroencephalography (EEG), but in many situations, EEG-based localization of the epileptogenic zone is inadequate. The complementary sensitivity of MEG can be crucial in such cases, and MEG has been adopted at many centers as an important resource in building a surgical hypothesis. In this paper, we review recent work evaluating the extent of MEG influence of presurgical evaluations, novel analyses of MEG data employed in surgical workup, and new MEG instrumentation that will likely affect the field of clinical MEG. RECENT FINDINGS: MEG consistently contributes to presurgical evaluation and these contributions often change the plan for epilepsy surgery. Extensive work has been done to develop new analytic methods for localizing the source of epileptiform activity with MEG. Systems using optically pumped magnetometry (OPM) have been successfully deployed to record and localize epileptiform activity. MEG remains an important noninvasive tool for epilepsy presurgical evaluation. Continued improvements in analytic methodology will likely increase the diagnostic yield of the test. Novel instrumentation with OPM may contribute to this as well, and may increase accessibility of MEG by decreasing cost.

Validation of On-Head OPM MEG for Language Laterality Assessment.

Pre-surgical localization of language function in the brain is critical for patients with medically intractable epilepsy. MEG has emerged as a valuable clinical tool for localizing language areas in clinical populations, however, it is limited for widespread application due to the low availability of the system. Recent advances in optically pumped magnetometer (OPM) systems account for some of the limitations of traditional MEG and have been shown to have a similar signal-to-noise ratio. However, the novelty of these systems means that they have only been tested for limited sensory and motor applications. In this work, we aim to validate a novel on-head OPM MEG procedure for lateralizing language processes. OPM recordings, using a soft cap with flexible sensor placement, were collected from 19 healthy, right-handed controls during an auditory word recognition task. The resulting evoked fields were assessed for hemispheric laterality of the response. Principal component analysis (PCA) of the grand average language response indicated that the first two principal components were lateralized to the left hemisphere. The PCA also revealed that all participants had evoked topographies that closely resembled the average left-lateralized response. Left-lateralized responses were consistent with what is expected for a group of healthy right-handed individuals. These findings demonstrate that language-related evoked fields can be elucidated from on-head OPM MEG recordings in a group of healthy adult participants. In the future, on-head OPM MEG and the associated lateralization methods should be validated in patient populations as they may have utility in the pre-surgical mapping of language functions in patients with epilepsy.

Sensor array design of optically pumped magnetometers for accurately estimating source currents.

An optically pumped magnetometer (OPM) is a new generation of magnetoencephalography (MEG) devices that is small, light, and works at room temperature. Due to these characteristics, OPMs enable flexible and wearable MEG systems. On the other hand, if we have a limited number of OPM sensors, we need to carefully design their sensor arrays depending on our purposes and regions of interests (ROIs). In this study, we propose a method that designs OPM sensor arrays for accurately estimating the cortical currents at the ROIs. Based on the resolution matrix of minimum norm estimate (MNE), our method sequentially determines the position of each sensor to optimize its inverse filter pointing to the ROIs and suppressing the signal leakage from the other areas. We call this method the Sensor array Optimization based on Resolution Matrix (SORM). We conducted simple and realistic simulation tests to evaluate its characteristics and efficacy for real OPM-MEG data. SORM designed the sensor arrays so that their leadfield matrices had high effective ranks as well as high sensitivities to ROIs. Although SORM is based on MNE, the sensor arrays designed by SORM were effective not only when we estimated the cortical currents by MNE but also when we did so by other methods. With real OPM-MEG data we confirmed its validity for real data. These analyses suggest that SORM is especially useful when we want to accurately estimate ROIs' activities with a limited number of OPM sensors, such as brain-machine interfaces and diagnosing brain diseases.

A New Generation of OPM for High Dynamic and Large Bandwidth MEG: The 4He OPMs-First Applications in Healthy Volunteers.

MagnetoEncephaloGraphy (MEG) provides a measure of electrical activity in the brain at a millisecond time scale. From these signals, one can non-invasively derive the dynamics of brain activity. Conventional MEG systems (SQUID-MEG) use very low temperatures to achieve the necessary sensitivity. This leads to severe experimental and economical limitations. A new generation of MEG sensors is emerging: the optically pumped magnetometers (OPM). In OPM, an atomic gas enclosed in a glass cell is traversed by a laser beam whose modulation depends on the local magnetic field. MAG4Health is developing OPMs using Helium gas (4He-OPM). They operate at room temperature with a large dynamic range and a large frequency bandwidth and output natively a 3D vectorial measure of the magnetic field. In this study, five 4He-OPMs were compared to a classical SQUID-MEG system in a group of 18 volunteers to evaluate their experimental performances. Considering that the 4He-OPMs operate at real room temperature and can be placed directly on the head, our assumption was that 4He-OPMs would provide a reliable recording of physiological magnetic brain activity. Indeed, the results showed that the 4He-OPMs showed very similar results to the classical SQUID-MEG system by taking advantage of a shorter distance to the brain, despite having a lower sensitivity.

Fostering engineering and science students' and teachers' systems thinking and conceptual modeling skills.

As science and technology create an ecosystem that is becoming increasingly more knowledge-intensive, complex, and interconnected, the next generation science standards include systems thinking and systems modeling among 21st skills that should be fostered. We examined the effect of an online cross-disciplinary learning process on the development of systems thinking and modeling skills among engineering students and engineering and science teachers. The study, which used quantitative and qualitative tools, included 55 participants who performed four food-related learning assignments and created conceptual models in Object-Process Methodology. Their responses to online assignments were analyzed along with their perceptions, captured via a reflection questionnaire. The online learning process in this study effectively enhanced systems thinking and modeling skills of all learners, including those with no relevant background. One main conclusion that extends beyond the online learning was that imparting the basics of systems thinking and conceptual modeling skills can be achieved even within a short period of time-less than one semester. The contribution of the study is the formation of theoretical and practical frameworks for the integration of an cross-disciplinary model-based systems engineering online assignments into engineering and science curricula.

Cross-Axis projection error in optically pumped magnetometers and its implication for magnetoencephalography systems.

Optically pumped magnetometers (OPMs) developed for magnetoencephalography (MEG) typically operate in the spin-exchange-relaxation-free (SERF) regime and measure a magnetic field component perpendicular to the propagation axis of the optical-pumping photons. The most common type of OPM for MEG employs alkali atoms, e.g. 87Rb, as the sensing element and one or more lasers for preparation and interrogation of the magnetically sensitive states of the alkali atoms ensemble. The sensitivity of the OPM can be greatly enhanced by operating it in the SERF regime, where the alkali atoms' spin exchange rate is much faster than the Larmor precession frequency. The SERF regime accommodates remnant static magnetic fields up to ±5 nT. However, in the presented work, through simulation and experiment, we demonstrate that multi-axis magnetic signals in the presence of small remnant static magnetic fields, not violating the SERF criteria, can introduce significant error terms in OPM's output signal. We call these deterministic errors cross-axis projection errors (CAPE), where magnetic field components of the MEG signal perpendicular to the nominal sensing axis contribute to the OPM signal giving rise to substantial amplitude and phase errors. Furthermore, through simulation, we have discovered that CAPE can degrade localization and calibration accuracy of OPM-based magnetoencephalography (OPM-MEG) systems.

Interference suppression techniques for OPM-based MEG: Opportunities and challenges.

One of the primary technical challenges facing magnetoencephalography (MEG) is that the magnitude of neuromagnetic fields is several orders of magnitude lower than interfering signals. Recently, a new type of sensor has been developed - the optically pumped magnetometer (OPM). These sensors can be placed directly on the scalp and move with the head during participant movement, making them wearable. This opens up a range of exciting experimental and clinical opportunities for OPM-based MEG experiments, including paediatric studies, and the incorporation of naturalistic movements into neuroimaging paradigms. However, OPMs face some unique challenges in terms of interference suppression, especially in situations involving mobile participants, and when OPMs are integrated with electrical equipment required for naturalistic paradigms, such as motion capture systems. Here we briefly review various hardware solutions for OPM interference suppression. We then outline several signal processing strategies aimed at increasing the signal from neuromagnetic sources. These include regression-based strategies, temporal filtering and spatial filtering approaches. The focus is on the practical application of these signal processing algorithms to OPM data. In a similar vein, we include two worked-through experiments using OPM data collected from a whole-head sensor array. These tutorial-style examples illustrate how the steps for suppressing external interference can be implemented, including the associated data and code so that researchers can try the pipelines for themselves. With the popularity of OPM-based MEG rising, there will be an increasing need to deal with interference suppression. We hope this practical paper provides a resource for OPM-based MEG researchers to build upon.

Wearable OPM-MEG: A changing landscape for epilepsy.

Magnetoencephalography with optically pumped magnometers (OPM-MEG) is an emerging and novel, cost-effective wearable system that can simultaneously record neuronal activity with high temporal resolution ("when" neuronal activity occurs) and spatial resolution ("where" neuronal activity occurs). This paper will first outline recent methodological advances in OPM-MEG compared to conventional superconducting quantum interference device (SQUID)-MEG before discussing how OPM-MEG can become a valuable and noninvasive clinical support tool in epilepsy surgery evaluation. Although OPM-MEG and SQUID-MEG share similar data features, OPM-MEG is a wearable design that fits children and adults, and it is also robust to head motion within a magnetically shielded room. This means that OPM-MEG can potentially extend the application of MEG into the neurobiology of severe childhood epilepsies with intellectual disabilities (e.g., epileptic encephalopathies) without sedation. It is worth noting that most OPM-MEG sensors are heated, which may become an issue with large OPM sensor arrays (OPM-MEG currently has fewer sensors than SQUID-MEG). Future implementation of triaxial sensors may alleviate the need for large OPM sensor arrays. OPM-MEG designs allowing both awake and sleep recording are essential for potential long-term epilepsy monitoring.

Practical real-time MEG-based neural interfacing with optically pumped magnetometers.

BACKGROUND: Brain-computer interfaces decode intentions directly from the human brain with the aim to restore lost functionality, control external devices or augment daily experiences. To combine optimal performance with wide applicability, high-quality brain signals should be captured non-invasively. Magnetoencephalography (MEG) is a potent candidate but currently requires costly and confining recording hardware. The recently developed optically pumped magnetometers (OPMs) promise to overcome this limitation, but are currently untested in the context of neural interfacing. RESULTS: In this work, we show that OPM-MEG allows robust single-trial analysis which we exploited in a real-time 'mind-spelling' application yielding an average accuracy of 97.7%. CONCLUSIONS: This shows that OPM-MEG can be used to exploit neuro-magnetic brain responses in a practical and flexible manner, and opens up new avenues for a wide range of new neural interface applications in the future.

Modeling with Real-Time Informative Feedback: Implementing and Evaluating a New Massive Open Online Course Component.

As part of the design, development, and deployment of a massive open online course (MOOC) on model-based systems engineering, we introduced MORTIF-Modeling with Real-Time Informative Feedback, a new learning-by-doing feature that enables the learner to model, receive detailed feedback, and resubmit improved solutions. We examined the pedagogical usability of MORTIF by investigating characteristics of participants working with it, and their perceived contribution, preferred question type, and learning style. The research included 295 participants and applied the mixed-methods approach, using MOOC server data and online questionnaires. Analyzing 12,095 submissions, we found increasing frequency of using the model resubmitting option. Students ranked MORTIF as the highest of six question types in terms of preference and perceived contribution level. Nine learning style categories were identified and classified based on students' verbal explanations regarding their preference of MORTIF over the other question types. MORTIF has been effective in promoting meaningful learning, supporting our hypothesis that the combination of active learning with real-time informative feedback is a learning mode that students eagerly embrace and benefit from. The benefits we identified for using MORTIF include active learning, provision of meaningful immediate feedback to the learner, the option to use the feedback on the spot and resubmitting an improved model, and its suitability for a variety of learning styles.

Contactless measurements of retinal activity using optically pumped magnetometers.

Optically pumped magnetometers (OPMs) have been adopted for the measurement of brain activity. Without the need to be cooled to cryogenic temperatures, an array of these sensors can be placed more flexibly, which allows for the recording of neuronal structures other than neocortex. Here we use eight OPM sensors to record human retinal activity following flash stimulation. We compare this magnetoretinographic (MRG) activity to the simultaneously recorded electroretinogram of the eight participants. The MRG shows the familiar flash-evoked potentials (a-wave and b-wave) and shares a highly significant amount of information with the electroretinogram (both in a simultaneous and separate measurement). We conclude that OPM sensors have the potential to become a contactless alternative to fiber electrodes for the measurement of retinal activity. Such a contactless solution can benefit both clinical and neuroscientific settings.

Mouth magnetoencephalography: A unique perspective on the human hippocampus.

Traditional magnetoencephalographic (MEG) brain imaging scanners consist of a rigid sensor array surrounding the head; this means that they are maximally sensitive to superficial brain structures. New technology based on optical pumping means that we can now consider more flexible and creative sensor placement. Here we explored the magnetic fields generated by a model of the human hippocampus not only across scalp but also at the roof of the mouth. We found that simulated hippocampal sources gave rise to dipolar field patterns with one scalp surface field extremum at the temporal lobe and a corresponding maximum or minimum at the roof of the mouth. We then constructed a fitted dental mould to accommodate an Optically Pumped Magnetometer (OPM). We collected data using a previously validated hippocampal-dependant task to test the empirical utility of a mouth-based sensor, with an accompanying array of left and right temporal lobe OPMs. We found that the mouth sensor showed the greatest task-related theta power change. We found that this sensor had a mild effect on the reconstructed power in the hippocampus (~10% change) but that coherence images between the mouth sensor and reconstructed source images showed a global maximum in the right hippocampus. We conclude that augmenting a scalp-based MEG array with sensors in the mouth shows unique promise for both basic scientists and clinicians interested in interrogating the hippocampus.

Measuring functional connectivity with wearable MEG.

Optically-pumped magnetometers (OPMs) offer the potential for a step change in magnetoencephalography (MEG) enabling wearable systems that provide improved data quality, accommodate any subject group, allow data capture during movement and potentially reduce cost. However, OPM-MEG is a nascent technology and, to realise its potential, it must be shown to facilitate key neuroscientific measurements, such as the characterisation of brain networks. Networks, and the connectivities that underlie them, have become a core area of neuroscientific investigation, and their importance is underscored by many demonstrations of their disruption in brain disorders. Consequently, a demonstration of network measurements using OPM-MEG would be a significant step forward. Here, we aimed to show that a wearable 50-channel OPM-MEG system enables characterisation of the electrophysiological connectome. To this end, we measured connectivity in the resting state and during a visuo-motor task, using both OPM-MEG and a state-of-the-art 275-channel cryogenic MEG device. Our results show that resting-state connectome matrices from OPM and cryogenic systems exhibit a high degree of similarity, with correlation values >70%. In addition, in task data, similar differences in connectivity between individuals (scanned multiple times) were observed in cryogenic and OPM-MEG data, again demonstrating the fidelity of the OPM-MEG device. This is the first demonstration of network connectivity measured using OPM-MEG, and results add weight to the argument that OPMs will ultimately supersede cryogenic sensors for MEG measurement.

Theoretical advantages of a triaxial optically pumped magnetometer magnetoencephalography system.

The optically pumped magnetometer (OPM) is a viable means to detect magnetic fields generated by human brain activity. Compared to conventional detectors (superconducting quantum interference devices) OPMs are small, lightweight, flexible, and operate without cryogenics. This has led to a step change in instrumentation for magnetoencephalography (MEG), enabling a "wearable" scanner platform, adaptable to fit any head size, able to acquire data whilst subjects move, and offering improved data quality. Although many studies have shown the efficacy of 'OPM-MEG', one relatively untapped advantage relates to improved array design. Specifically, OPMs enable the simultaneous measurement of magnetic field components along multiple axes (distinct from a single radial orientation, as used in most conventional MEG systems). This enables characterisation of the magnetic field vector at all sensors, affording extra information which has the potential to improve source reconstruction. Here, we conduct a theoretical analysis of the critical parameters that should be optimised for effective source reconstruction. We show that these parameters can be optimised by judicious array design incorporating triaxial MEG measurements. Using simulations, we demonstrate how a triaxial array offers a dramatic improvement on our ability to differentiate real brain activity from sources of magnetic interference (external to the brain). Further, a triaxial system is shown to offer a marked improvement in the elimination of artefact caused by head movement. Theoretical results are supplemented by an experimental recording demonstrating improved interference reduction. These findings offer new insights into how future OPM-MEG arrays can be designed with improved performance.

Precision magnetic field modelling and control for wearable magnetoencephalography.

Optically-pumped magnetometers (OPMs) are highly sensitive, compact magnetic field sensors, which offer a viable alternative to cryogenic sensors (superconducting quantum interference devices - SQUIDs) for magnetoencephalography (MEG). With the promise of a wearable system that offers lifespan compliance, enables movement during scanning, and provides higher quality data, OPMs could drive a step change in MEG instrumentation. However, this potential can only be realised if background magnetic fields are appropriately controlled, via a combination of optimised passive magnetic screening (i.e. enclosing the system in layers of high-permeability materials), and electromagnetic coils to further null the remnant magnetic field. In this work, we show that even in an OPM-optimised passive shield with extremely low (<2 nT) remnant magnetic field, head movement generates significant artefacts in MEG data that manifest as low-frequency interference. To counter this effect we introduce a magnetic field mapping technique, in which the participant moves their head to sample the background magnetic field using a wearable sensor array; resulting data are compared to a model to derive coefficients representing three uniform magnetic field components and five magnetic field gradient components inside the passive shield. We show that this technique accurately reconstructs the magnitude of known magnetic fields. Moreover, by feeding the obtained coefficients into a bi-planar electromagnetic coil system, we were able to reduce the uniform magnetic field experienced by the array from a magnitude of 1.3±0.3 nT to 0.29±0.07 nT. Most importantly, we show that this field compensation generates a five-fold reduction in motion artefact at 0‒2 Hz, in a visual steady-state evoked response experiment using 6 Hz stimulation. We suggest that this technique could be used in future OPM-MEG experiments to improve the quality of data, especially in paradigms seeking to measure low-frequency oscillations, or in experiments where head movement is encouraged.

Multi-channel whole-head OPM-MEG: Helmet design and a comparison with a conventional system.

Magnetoencephalography (MEG) is a powerful technique for functional neuroimaging, offering a non-invasive window on brain electrophysiology. MEG systems have traditionally been based on cryogenic sensors which detect the small extracranial magnetic fields generated by synchronised current in neuronal assemblies, however, such systems have fundamental limitations. In recent years, non-cryogenic quantum-enabled sensors, called optically-pumped magnetometers (OPMs), in combination with novel techniques for accurate background magnetic field control, have promised to lift those restrictions offering an adaptable, motion-robust MEG system, with improved data quality, at reduced cost. However, OPM-MEG remains a nascent technology, and whilst viable systems exist, most employ small numbers of sensors sited above targeted brain regions. Here, building on previous work, we construct a wearable OPM-MEG system with 'whole-head' coverage based upon commercially available OPMs, and test its capabilities to measure alpha, beta and gamma oscillations. We design two methods for OPM mounting; a flexible (EEG-like) cap and rigid (additively-manufactured) helmet. Whilst both designs allow for high quality data to be collected, we argue that the rigid helmet offers a more robust option with significant advantages for reconstruction of field data into 3D images of changes in neuronal current. Using repeat measurements in two participants, we show signal detection for our device to be highly robust. Moreover, via application of source-space modelling, we show that, despite having 5 times fewer sensors, our system exhibits comparable performance to an established cryogenic MEG device. While significant challenges still remain, these developments provide further evidence that OPM-MEG is likely to facilitate a step change for functional neuroimaging.

Obscuration Threshold Database Construction of Smoke Detectors for Various Combustibles.

The obscuration thresholds for various smoke detectors and combustibles, required as an input parameter in fire simulation, were measured to predict the accurate activation time of detectors. One ionization detector and nine photoelectric detectors were selected. A fire detector evaluator, which can uniformly control the velocity and smoke concentration, was utilized. Filter paper, liquid fuels, and polymer pellets were employed as smoke-generation combustibles. The nominal obscuration thresholds of the considered detectors were 15 %/m, but the ionization detectors activated at approximately 40 %/m and 16 %/m, respectively, on applying filter paper and kerosene. In contrast, the reverse obscuration thresholds were found quantitatively according to the combustibles in the photoelectric detector. This phenomenon was caused by differences in the color of the smoke particles according to the combustibles, which is explained by single-scattering albedo (ratio of light scattering to light extinction). The obscuration thresholds for liquid fuels (kerosene, heptane and toluene) as well as fire types of polymer plastic pellets were also measured for several photoelectric detectors. A database of obscuration thresholds was thereby established according to the detector and combustible types, and it is expected to provide useful information for predicting more accurate detector activation time and required safe egress time (REST).

Magnetic Source Imaging Using a Pulsed Optically Pumped Magnetometer Array.

We have developed a pulsed optically pumped magnetometer (OPM) array for detecting magnetic field maps originated from an arbitrary current distribution. The presented magnetic source imaging (MSI) system features 24 OPM channels, has a data rate of 500 S/s, a sensitivity of 0.8 p T / H z , and a dynamic range of 72 dB. We have employed our pulsed- OPM MSI system for measuring the magnetic field map of a test coil structure. The coils are moved across the array in an indexed fashion to measure the magnetic field over an area larger than the array. The captured magnetic field maps show excellent agreement with the simulation results. Assuming a 2D current distribution, we have solved the inverse problem, using the measured magnetic field maps, and the reconstructed current distribution image is compared to that of the simulation.

Two's company, three's a crowd: co-occurring pollinators and parasite species in Breynia oblongifolia (Phyllanthaceae).

BACKGROUND: Obligate pollination mutualisms (OPMs) are specialized interactions in which female pollinators transport pollen between the male and female flowers of a single plant species and then lay eggs into those same flowers. The pollinator offspring hatch and feed upon some or all of the developing ovules pollinated by their mothers. Strong trait matching between plants and their pollinators in OPMs is expected to result in reciprocal partner specificity i.e., a single pollinator species using a single plant species and vice versa, and strict co-speciation. These issues have been studied extensively in figs and fig wasps, but little in the more recently discovered co-diversification of Epicephala moths and their Phyllanthaceae hosts. OPMs involving Epicephala moths are believed occur in approximately 500 species of Phyllanthaceae, making it the second largest OPM group after the Ficus radiation (> 750 species). In this study, we used a mixture of DNA barcoding, genital morphology and behavioral observations to determine the number of Epicephala moth species inhabiting the fruits of Breynia oblongifolia, their geographic distribution, pollinating behavior and phylogenetic relationships. RESULTS: We found that B. oblongifolia hosts two species of pollinator that co-occurred at all study sites, violating the assumption of reciprocal specificity. Male and female genital morphologies both differed considerably between the two moth species. In particular, females differed in the shape of their ovipositors, eggs and oviposition sites. Phylogenetic analyses indicated that the two Epicephala spp. on B. oblongifolia likely co-exist due to a host switch. In addition, we discovered that Breynia fruits are also often inhabited by a third moth, an undescribed species of Herpystis, which is a non-pollinating seed parasite. CONCLUSIONS: Our study reveals new complexity in interactions between Phyllantheae and Epicephala pollinators and highlights that host switching, co-speciation and non-pollinating seed parasites can shape species interactions in OPMs. Our finding that co-occurring Epicephala species have contrasting oviposition modes parallels other studies and suggests that such traits are important in Epicephala species coexistence.