Joffrey Becker

Samuel Bianchini
email: samuel.bianchini@ensad.fr

Hugo Scurto
email: hugo.scurto@ensad.fr

Elena Tosi

Elena Tosi Brandi
email: elena.tosibrandi@orange.com

Picking Up Good Vibrations: Designing an Experimental Setup to Assess the Role of Vibrations in Human-Robot Interaction

Focusing on behavioral objects (considered as art and design objects which are not figurative nor utilitarian), this article describes a first step in the design of a research and creation project called The Appprentices. Conceived as an environment containing three behavioral objects which movements are amplified, The Appprentices constitutes an experimental platform which aims to contribute to a better understanding of the modalities by which humans and objects are in a situation of nonverbal communication. The project aims to contribute to the study of animism in human-robot interaction (HRI) but also to the study of analogical forms of communication generally prevailing in multi-species interaction. It therefore implies to pay a particular attention to the design of a shared space. Reviewing several works in the field of art and design where vibration plays a central role, we address a dimension of animism that is still poorly studied in HRI anthropology, based not on the idea that objects are considered as persons but rather that their collective activities can be considered by analogy with social life. We then identify vibration as a relevant modality for relating human and nonhuman collectives, which allows a frame for interaction within which vibratory signals can be exchanged and embodied. The article concludes with a description of the architecture of the shared vibratory space designed for the project.

I. INTRODUCTION

In the context of contemporary design, interaction with artifacts is evolving thanks to the richness of human-machine interaction and ubiquitous computing. The rise of artificial intelligence and the democratization of robotics impose new modalities of relation between humans and the artificial world. Communicating and connected objects capable of sharing data with their environment and with each other, sensory interfaces, social robots, conversational agents constitute a whole new typology of artifacts which, by their expressiveness and their spontaneous or programmed behaviors, express intentions and therefore call for new forms of interaction with their audiences and their environments.

Since 2012, a multidisciplinary group of artists, designers, engineers, and researchers initiated a research project in the field of art and design to study and experiment with what we call Behavioral Objects. These non-anthropomorphic, non-zoomorphic objects and collective of objects are endowed with capacities for expressive movement, action and reactions and are also able to elicit observers' behavioral interpretations (intentional and emotional attributions) [START_REF] Samuel | The misbehavior of animated object[END_REF], [START_REF] Samuel | Mis)behavioral Objects[END_REF], [START_REF]Behavioral Objects I, A Case Study: Céleste Boursier-Mougenot[END_REF], [START_REF] Florent | Behavioral Objects: The Rise of the Evocative Machines[END_REF].

Our ongoing research, The Appprentices, considers the agency of a collective of robotic objects. It pleads for a better awareness of the complex psycho-social dynamics involved when humans interact with a group of robotic agents, specifically by paying attention to the way vibration helps compose a shareable environment and thus gives a frame for communication between humans and robots. Indeed, vibrations, sounds and even rhythmic structures are perceived and produced by many different species. Vibration also plays an important role in situations where analog multi-species communication constitutes a common frame when humans exchange signals with other nonhuman types of existence across the globe. As we are going to see, this justifies to conceive a shared, embodied and composed vibratory space as a preliminary step in our work.

To which extent such a space leads humans to consider the animacy of a robot group, that is, on the one hand, the attribution/abduction of agency [START_REF] Alfred | Art and Agency, An Anthropological Theory[END_REF] and on the other the psychosocial effects resulting from the coordinated activity of a group of objects? Our paper questions how vibrations can constitute expressive means which give a context for spectators to elicit collective dynamics and social qualities from the objects' activity. What is at stake here is the conception of a shared vibratory space which enables new modalities for human-robot interaction. Leaning firstly on art and design practice and anthropological researches conducted within the fields of robotics and artificial intelligence, this paper describes the conception of a shared vibratory ecosystem in order to address the questions raised by embodied and ambient forms of communication with behavioral objects.

II. INSIGHT FROM ART AND DESIGN PRACTICE

Some behavioral objects conceived in the field of arts and design address the role of sound in human-machine interaction. For instance, by displaying a metal container deforming noisily under what seems to be blows coming from the inside, Arcangelo Sassolino's Piccolo Animismo (2011) confronts the spectators with a radical otherness that leads us to suspect that this box contains an eerie and threatening entity. In Rebecca Horn's Ballet of the Woodpeckers (1986), motorized hammers hit the surface of mirrors while the spectators can see how their own movements create waves on the surface of two cones filled with water. These works of art are not only staging how important sound is in humanrobot encounters. They sometimes use sound and vibration as a means to highlight a more collective dynamic.

Conducted at the EnsadLab, the works of Ianis Lallemand and Olivain Porry provide a first insight on the approach we adopt today on vibration. In Tous ensemble, chacun pour soi ("All together, each one for itself"), Olivian Porry explores the effects of synchrony and de-synchrony in a small swarm of robotic agents, showing the particular relations occurring between them (see Fig. 1). When one enters the room of the installation the first thing she perceives is small nasal whispers, soft and short that are repeated with regularity. Perched on springs and covered with a plate that reflects, in the same way a mirror does, the space around them, the robots shake and turn slightly with the movements of their rods. Scattered carelessly in the exhibition space, the robots appear to be very active and if the movements of the rods are first synchronized, they change after a few minutes. Indeed, one robot in particular stands out from the group. It shakes its rod in a fast motion and then stops. The five other robots then reproduce this same movement and thus draw a choreography which is guided by the first one. The robots vibrate in a loud noise and move on the ground for a few seconds, thus adopting new positions. The loop then resumes at its start and the choreography of the rods begins again.

In a cooperative work with Olivier Dauchot and the Collective Effects & Soft Matter team at ESPCI-Paris, Ianis Lallemand also worked on vibration but adopted a differ- ent perspective by conceiving a shared vibratory space (a vibrating surface) which animate the objects it contains (see Fig. 2). In Manoeuvres (2016), Lallemand conceived an experimental setup which allows the spectators to create intricate light images, by contributing to the self-organization of a physical multi-agent system moved by a vibration plate converting energy into movements that are then translated into visual patterns thanks to light projection. Using vibration is a well known method employed to make robots move. It is a core feature used in machines such as Hexbugs or Kilobots. The difference here, however, is that Ianis Lallemand's objects are moved by their very own ecosystem. The origin of their movement is thus external, embodying vibration in a given and composed situation which directly works the perception we have of their collective dynamics.

These behavioral objects invite us to wonder whether their perceived behavior is the result of a collective organized pattern or not. They put us in a situation where we, as spectators, are facing a society made of behavioral objects, working the notion of animism in a particular way. Indeed, anthropologists have shown that animism is not only about feeling that an object is behaving as a person would, but also about considering the activity of nonhuman groups as the index of a human-like social life [START_REF] Descola | Par delà nature et culture[END_REF], [START_REF] De | Exchanging Perspectives : The Transformation of Object into Subjects in Amerindian Ontologies[END_REF], [START_REF] De | Perspectivisme et multinaturalisme en Amérique indigène[END_REF]. How does vibration contribute to create a shared space where human and nonhuman agents can exchange information or even meaningful signs?

III. AN ANTHROPOLOGICAL ISSUE

While "face-to-face" interactions with robotic agents are holding the attention of many researchers in the field of human-robot interaction, the issue of interacting with multiple robots has gained interest only recently. A growing number of researches address today the many issues raised by human multi-robot interaction, whether they consist of experimenting communication based on language, gestures and social cues, interfaces which enable mediation or a combination of all these [START_REF] Gordon | Multi-modal Belief Updates in Multi-Robot Human-Robot Dialogue Interactions[END_REF], [START_REF] Stoica Adrian | Towards Human-Friendly Efficient Control of Multi-Robot Teams[END_REF], [START_REF] Gerard | Gesture based human multi-robot interaction[END_REF], [START_REF] Boris | Wearable multi-modal interface for human multi-robot interaction[END_REF]. Still, little is known about how we perceive the collective behavior of a swarm of robots [START_REF] St-Onge | Collective expression: How robotic swarms convey information with group motion[END_REF]. Yet pioneering experiments, such as the work of Fritz Heider and Marianne Simmel on the perceived social features of moving objects [START_REF] Fritz | An Experiment Study of Apparent Behavior[END_REF], or other works about the role of motor coordination on the emergence of a feeling of identity conducted in the field of humanities and social sciences [START_REF] Mark | La danse comme texte: Idéologies du corps baroque[END_REF], [START_REF] Georges | Le corps travaillé, Gymnastes et sportifs au XIXe siècle[END_REF], [START_REF] William | Keeping Together in Time, Dance and Drill in Human History[END_REF], remind us how important such a subject is.

HRI experiments occasionally investigated the means by which robots are able to communicate with each other, focusing on the modes of communication which could appear as the more natural according to human. For instance, Williams et al. showed that, in task-oriented human-robots experimental setups, verbal communication between robots is preferable to silence [START_REF] Tom | Covert Robot-Robot Communication: Human Perceptions and Implications for Human-Robot Interaction[END_REF]. Language is a good way to avoid the creepiness felt by human users when they have to deal with silent robots. Our goal is to help enrich these researches by addressing a nonsymbolic type of communication based on vibration. As we shall see, anthropomorphism (whether it concerns forms or language) is perhaps not the most suited way to engage humans to consider robots as potential social partners.

Anthropologists such as Emmanuel Grimaud or Denis Vidal have shown that robots are good candidates for studying the cognitive processes involved in anthropomorphic thinking and the perceived agency of these objects [START_REF] Vidal | Anthropomorphism of Sub-Anthropomorphism? An Anthropological Approach to Gods and Robots[END_REF], [START_REF] Emmanuel | Le jour où les robots mangeront des pommes, conversations avec un geminoïd[END_REF], [START_REF] Denis | Robots étrangement humains[END_REF], [START_REF] Becker | Humanoïdes[END_REF], [START_REF] Vidal | Aux frontières de l'humain, Dieux: figures de cire, robots et autres artefacts[END_REF]. Experimenting human-machine interactions with the Berenson robot at the Quai Branly Museum, Vidal has shown that the main issue when encountering an anthropomorphic robot is about assessing its autonomy and limitations. This measure aims to verify that the machine is indeed one. However, once the mechanical nature of the robot is confirmed during the interaction, this does not prevent humans from behaving as if it was a "person" [START_REF] Vidal | Aux frontières de l'humain, Dieux: figures de cire, robots et autres artefacts[END_REF]. As Vidal suggests, by establishing continuities between human and machine inner processes and simultaneously noting that a discontinuity exists between our appearance and theirs, do human-robot interactions reveal that a renewed form of animism is at work? Philippe Descola has clearly shown that animism implies such a relationship of continuity and discontinuity [START_REF] Descola | Par delà nature et culture[END_REF]. In animist societies, if the whole of the existing nonhumans is thought to be crossed by an interiority identical to that of humans, it is mainly on the basis of a morphological difference that the relations of identity and otherness will be established. One can imagine that relations with robots could easily be compared to those occurring in animist worlds. In fact, anthropologists do not address another, yet fundamental, aspect of the notion of animism.

In animist worlds, people don't only consider nonhuman beings as persons. They also consider the sociability which is established between members of the same species, of different species, or between living beings and artifacts [START_REF] De | Exchanging Perspectives : The Transformation of Object into Subjects in Amerindian Ontologies[END_REF], [START_REF] De | Perspectivisme et multinaturalisme en Amérique indigène[END_REF], [START_REF] Kohn | How Forests Think: Toward an Anthropology Beyond the Human[END_REF], [START_REF] Fernando | The Occult Life of Things, Native Amazonian Theories of Materiality and Personhood[END_REF]. While it is easy to think that machine have a perceptual world of their own [START_REF] Becker | Humanoïdes[END_REF], [START_REF] Becker Joffrey | Qualcom : une expérience sur la qualification des comportements d'une lampe robotique[END_REF], an umwelt [START_REF] Uexküll | A Foray into the Worlds of Animals and Humans: With A Theory of Meaning[END_REF], experiments conducted in the field of human-robot interaction show that attributing the quality of a subject to a machine does not necessarily imply that we infer that robots have, between themselves, social relations analogous to ours, as this would be the case in the type of animist ontological inference described by Philippe Descola [START_REF] Descola | Par delà nature et culture[END_REF]. Would we spontaneously infer social qualities about groups of objects acting synchronously or collectively imitating certain human actions? Far from only considering the faceto-face interactions between a human and a machine that looks like him (as it is still too often the case in human-robot interaction experiments), the coordinated actions of groups of robots invite us to address the question of animism not only for the imputation of the quality of subject that is associated to it, but also through the inferred "social life" associated to behavioral objects. This dimension incites us to explore other modalities of communication which are weakly considered by researchers as they don't contribute to the conception of universal relational interfaces whose functioning would be thought as a kind of transfer of the human communication behaviors into machines.

Indeed, experiments made by anthropologists on social robots [START_REF] Vidal | Anthropomorphism of Sub-Anthropomorphism? An Anthropological Approach to Gods and Robots[END_REF], [START_REF] Emmanuel | Le jour où les robots mangeront des pommes, conversations avec un geminoïd[END_REF], [START_REF] Denis | Robots étrangement humains[END_REF], [START_REF] Becker | Humanoïdes[END_REF], [START_REF] Vidal | Aux frontières de l'humain, Dieux: figures de cire, robots et autres artefacts[END_REF] show that interacting with a robot almost systematically implies that the human interactant mobilizes resources which are different from those daily used to interact with members of our species [START_REF] Emmanuel | Le jour où les robots mangeront des pommes, conversations avec un geminoïd[END_REF], [START_REF] Becker | Humanoïdes[END_REF]. With robots, the principles of an analog type of communication are at work [START_REF] Gregory | Steps to an Ecology of Mind[END_REF], which can for instance be compared to that we establish with our pets. What prevails in such cases is the establishment of a framework which does not focus on the messages contents but whose function is to open and maintain a channel for communication.

Where digital communication depends on the content of the messages exchanged, analog communication is about the very possibility of communicating; the creation of a shared, jointly habitable and meaningful world, despite the extremely rudimentary nature of the means used to communicate.

IV. VIBRATIONS: A CONTEXT FOR SOCIAL BONDING

Our work follows the perspective opened by this form of communication in HRI. It aims to explore nonverbal forms of communication based on the way vibrations are the cause of movements, sound and rhythm and thus appear as a first source which give access to more complex sensory dimensions. Analog communication can be seen in situations that are very distant both geographically or culturally and is addressed in a growing number of studies conducted in the field of anthropology [START_REF] Charles | Communication rituelle et communication interspécifique[END_REF], [START_REF] Véronique | La visite au zoo et l'apprentissage de la distinction humaine[END_REF], [START_REF] Radhika | The goat that died for family: Animal sacrifice and interspecies kinship in India's Central Himalayas[END_REF], [START_REF] Lindsay | Ethnography after humanism, Power, politics and methods in multi-species research[END_REF].

However, when it comes to interspecific communication, few anthropologists focus on direct information exchange. The ethnography of the modes of intersubjectivity between species remain poor though the subject gained progressively interest in anthropology during the last decade. Although their work adopts an interspecific perspective, many anthropologists consider the human-animal relationship to better grasp the networks and dynamics of human social agency [33], [START_REF] Kohn | How Forests Think: Toward an Anthropology Beyond the Human[END_REF], thus still centering the work on humans and their institutions.

The shared world between humans and nonhumans, what it is made of and crossed by, is not always considered for the questions of communication it raises, for instance by addressing what element could be relevant when species meet and exchange information [START_REF] Sperber Dan | Relevance: Communication and Cognition[END_REF]. Interspecific modalities of communication in HRI and the common world at the interface of which signals and signs can potentially be exchanged (the semiosphere, as called by Yuri Lotman, for example) should receive special attention.

Anthropologists such as Stéphane Rennesson, Nicolas Césard, and Emmanuel Grimaud have noted, for example, that rhinoceros beetle fights in Thailand [START_REF] Stéphane | Insect Magnetism: The communication circuit of Rhinoceros beetle fighting in Thailand[END_REF] depend on a technique that involves establishing common ground between the human player and the insect relying primarily on vibrations and tactile means. Although human players do not share any cognitive or perceptual abilities with the insect, producing vibrations on the bamboo trunk used as a battle area is an attempt to affect the beetle's behavior. The vibratory space appears as an interactional space where energy is transferred from a body to another at a preverbal level in a shared material environment. Vibration is thus a mediation means which can have deep consequences on the relation one can have with a nonhuman agent. This is even more the case when sounds are or seems organized and structured into rhythm. The notion of rhythm is, however, fraught with a certain ambiguity as it confuses the rhythms of life (calendar cycles, synchronization of activities, the question of repetition in relation to learning practices over a long period of time, etc.) and a more musical and situated definition which anthropology does not always manage to grasp, especially in ritual situations. In such a situation, rhythm is not a trigger for anything specific, even if it appears as a context accompanying practices [START_REF] Gilbert | La musique et la transe, Esquisse d'une théorie générale des relations de la musique et de la posssession[END_REF]. Nevertheless, despite this ambiguity, what is at stake is that -whether they are of a musical kind or not -vibrations appear as a particular context which gives a frame for social activities and bonding.

Our approach aims at exploring this last modality without necessarily trying to transpose the universe of signs specific to verbal language to an interactional situation based firstly on vibration, then sound and the imitation of rhythmic patterns. It is thus different from the work on vibratory communication conducted in the field of human-computer interaction, which seeks mainly to design techniques in order to convey strong meaning for human users [START_REF] Brill | Tactile Technology for Covert Communications[END_REF], [START_REF] Brown | Feel who's talking : using tactons for mobile phone alerts[END_REF], [START_REF] Stephen | Tactile Feedback for Mobile Interactions[END_REF], [START_REF] Daniel | Toward a Tactile Language for Human-Robot Interactions : Two Studies of Tacton Learning and Performance[END_REF]. The experimental perspective we adopt is different from such an iconic approach. It relies on vibration as an embodied feature for communication in order to create an experimental context whose purpose is to assess how humans think about the activity of a collective of robotic agents by analogy with our own social activities.

V. DESIGNING A COMMON VIBRATORY ECOSYSTEM:

THE APPPRENTICES How to compose a shared vibratory space? While in Tous ensemble, chacun pour soi, vibration emerges through the sound coming from the physical movements of the objects, including their motorization, Manoeuvres rely on a vibration plate to animate the set of agents. In both cases the vibratory space naturally transposes material activities to our auditory perception, thus reinforcing our attention. Why not have recourse to such material actions-to this vibratory space-to interact with these objects, for example by inviting an audience who have become users to knock on the common support of these objects? How can we design a material environment for these objects that is not simply a stage, but also a kind of instrument, an environment calling for communication with these objects through a percussive gesture as a source of vibration? How to compose the whole set of this vibratory space of interaction which requires to prevent or, on the contrary, to foster the propagation of the vibrations through the various "materials" implied (metal, wood, plastics, rubber, human body, air, etc.)? How to think the continuities and the discontinuities of this space of communication thus thought as a sensitive embodied space? Thought to experience a trio of behavioral objects within a vibratory space to be shared, The Appprentices will remind an orientation table (see Fig. 3). It will consist of a large circular anthracite base, which stands in the middle of an interior space. Three objects will move irregularly on its surface. All of the same shape, these objects look like pebbles whose gray base merges with the scene while their lighter top offers enough contrast to distinguish them well.

Our project leads us to design an "instrument-stage" which consists in a vibratory space which can be shared by human and objects. This common human-robot apparatus should make it possible to stage this collective of objects while stimulating interactions by means of an instrumental form favoring affordances, such as a large drum that has become a stage, and paving the way to interactions with a collective of objects by a collective of humans. Just like the beetle fighting studied by S. Rennesson, E. Grimaud and N. Césard involves an unusual form of partnership between humans and insects that shed a new light on the notion of communication [START_REF] Stéphane | Insect Magnetism: The communication circuit of Rhinoceros beetle fighting in Thailand[END_REF], we conceived a vibratory space as a privileged environment for interspecific cooperation between humans and objects.

Beyond aesthetics, the whole apparatus-including robotic objects and stage-was designed to compose a vibration space (seen as both a space for communication and a shared material space) (see Fig. 4). Structure-borne vibrations made by robots' movements are converted into electronic signals by piezoelectric sensors placed under a metal plate. We chose metal as material for the plate due to its high impedance, which enables to soothe air-borne vibrations produced by a speaking audience, while strengthening structure-borne vibrations produced by robots' movements over the stage.

After going through some audio processing, the robots' vibratory signals are converted into structure-borne vibrations and diffused air-borne using piezoelectric speakers placed on a wooden box. As such, the audience can hear vibrations produced by robots through sounds diffused by the vibrating box, but also touch it, through haptic feedback produced by the box.

Crucially, we placed a rubber insulator between the metal plate and the wooden box, as well as an acoustic foam inside the wooden box, to respectively avoid structure-and airborne vibratory feedback between piezoelectric sensors and speakers. Similarly, we placed a rubber insulator between the wooden box and the ground to soothe structure-borne vibrations produced by a walking audience and other entities moving around the exhibition space. Lastly, we equipped the stage's edge with piezoelectric sensors to sense the audience's structure-borne tappings, thus designing distinct communication channels between robots and the audience within a shared vibratory ecosystem.

VI. CONCLUSION

Our current research and creation project aims to question the behaviors of objects from the artificial world and, on the other hand, to identify methods and tools in order to define an approach in designing the agency of artifacts and thus creating experimental condition to receive them, materialize and assess them [START_REF] Mel | Social Perception Detection and Interpretation of Animacy, Agency, and Intention[END_REF], [START_REF] Brandi | 08 De l'expressivité à l'animacy des artefacts[END_REF]. Inspired by several works involving Behavioral Objects collectives, our ongoing efforts plead for extending the research on the expressiveness of machines but also on the communication uncertainties and the inferences emerging from the interaction.

Recognizing that vibration is a fundamental component in animal behavior and communication processes [START_REF] Denis | Contribution à l'étude du ronronnement chez le chat domestique (Felis catus L.) et chez le chat sauvage (Felis silvestris S.) : Aspects morpho-fonctionnels acoustiques et éthologiques[END_REF], [START_REF] Howard | Sensory Exotica, A World beyond Human Experience[END_REF], [START_REF] Fernando | Mechanical phase shifters for coherent acoustic radiation in the stridulating wings of crickets: the plectrum mechanism[END_REF], [START_REF] Michael | Frequency information in the vibration-cued escape hatching of red-eye treefrogs[END_REF], [START_REF] Christian | Complex vibratory patterns in an elephant larynx[END_REF], [START_REF] Stéphane | Insect Magnetism: The communication circuit of Rhinoceros beetle fighting in Thailand[END_REF], The Appprentices aim to experiment the semiosphere opened by the shared vibratory space described above. In contrast to a merely symbolic approach to vibratory communication, The Appprentices aims to contribute to the study of human-robot interaction by relying on an analogical modality of communication [START_REF] Gregory | Steps to an Ecology of Mind[END_REF].

This particular form of communication is at work in multispecies relations in very different human societies around the world [START_REF] Charles | Communication rituelle et communication interspécifique[END_REF], [START_REF] Véronique | La visite au zoo et l'apprentissage de la distinction humaine[END_REF], [START_REF] Stéphane | Insect Magnetism: The communication circuit of Rhinoceros beetle fighting in Thailand[END_REF], [START_REF] Radhika | The goat that died for family: Animal sacrifice and interspecies kinship in India's Central Himalayas[END_REF], [START_REF] Lindsay | Ethnography after humanism, Power, politics and methods in multi-species research[END_REF]. Our goal is to better understand the role of vibration, sound and rhythm in the perception of animacy in a group of robotic agents and thus to tackle the links between robotics and animism from a new perspective.

Fig. 1 .

 1 Fig. 1. Tous ensemble chacun pour soi. Olivain Porry, 2018. A project developed as part of the ICOC project (Collective Interface for Connected Objects) of the Reflective Interaction research group of EnsadLab conducted in partnership with the CHArt laboratory with the support of the Carnot Cognition Institute.

Fig. 2 .

 2 Fig. 2. Manoeuvres, Ianis Lallemand, 2016. A project developed as part of EnsadLab's Reflective Interaction research group in collaboration with Olivier Dauchot and the Collective Effects & Soft Matter team of the École supérieure de physique et de chimie industrielles de la ville de Paris (ESPCI) with the support of PSL University.

Fig. 3 .

 3 Fig. 3. Preview images of the project The Appprentices, conceived by Samuel Bianchini, Elena Tosi Brandi and Hugo Scurto with the collaboration of Joffrey Becker and Franck Weens as part of a research partnership between Orange's Art Direction Nods Team (Xdlab) and EnsadLab's Reflective Interaction research group, February 2020.

Fig. 4 .

 4 Fig. 4. Description of the vibratory space designed for The Appprentices.

ACKNOWLEDGEMENTS

The authors of the present paper would like to thank Florent Levillain, Olivain Porry, Ianis Lallemand, Victor Audouze, Franck Weens, Emanuele Quinz, Corentin Loubet, and Joséphine Mas.

This research was supported by Orange's Art Direction Nods Team (Xdlab) and EnsadLab's Reflective Interaction research group, the research laboratory of the École nationale supérieure des Arts Décoratifs.


Marie Pégard

Julien Leuenberger

Bernadette Julier

Philippe Barre

Genomic prediction for phenology, yield and forage quality traits on a large diversity of alfalfa

ou non, émanant des établissements d'enseignement et de recherche français ou étrangers, des laboratoires publics ou privés.

Genomic prediction for phenology, yield and forage quality traits on a large diversity of alfalfa

Marie Pégard 1 , Julien Leuenberger 1 , Bernadette Julier 1 and Philippe Barre 1 1 INRAE P3F, 86600 Lusignan, France

China and Europe dependency on imported proteins is a problem that could be solved by growing more legumes. This requires productive and good quality cultivars. The current breeding programmes could be improved by using a large genetic diversity combined with a genomic evaluation such as in many crop or animal species, including forage species. Some studies have already been published on genomic evaluation of alfalfa, they have used between 75 and 274 accessions or populations, with a number of SNP from 8 K to 44 K. They have reported variable predictive accuracies (0-0.65) for yield and quality traits.

We investigated the ability of genomic prediction to predict various genetic material on a large number of traits. We used a set of 395 alfalfa accessions mostly coming from Europe, North and South America and China with a fall dormancy between 3 to 7. This material was genotyped by genotyping-by-sequencing (GBS) method at the accession level. After filtering, 228 568 markers (SNP) covering the genome were obtained. These genomic data were used to identify genetic clusters of accessions with the Discriminant Analysis of Principal Components (DAPC) method. From 2018 to 2020, the accessions were phenotyped in four locations (Lusignan in France, Novi Sad in Serbia, Aberystwyth in England, Store Heddinge in Denmark) for three categories of traits: Forage yield, Forage quality, and, Phenology (winter dormancy and flowering date), ending to 187 traits representing the combination of a trait in a location and a year or integrated traits (combined between year and/or locations). The phenotypes were adjusted for the micro-environmental variation inside each trial, the year and the location effects. The QTL were detected by Multi-Locus Mixed Model (mlmm) and retained if they pass the Bonferonni threshold. Then, we assessed the quality of the genomic prediction for each trait. A cross-validation was used to assess the prediction quality with a test of several models (GBLUP, Ridge-regression (RR), Bayesian lasso). We also tested the integration of QTL in the model as fixed effects (QGBLUP) compared to the classical GBLUP model.

A genetic structure with seven clusters, mostly related to the geographical origin of the accessions, was found. Two groups were clearly separated from the other accessions: one with only Chinese accessions and the other with some subsp. falcata accessions. The other five groups, respectively composed by the accessions of (1) France and Northern Europe, (2) Southern and Eastern Europe, (3) a mix from Europe, North and South America, (4) USA and 1 of China, and (5) North-American only, formed a continuum. Many QTLs were found: 438 for forage yield, 79 for phenology, and 564 for quality. Each QTL explained on average 0.17, 0.09, 0.13 and 0.07 of the phenotype for forage yield, phenology, quality and seed yield, respectively. The evaluation of the genomic prediction reveal that, we reached high predicting abilities. The infinitesimals methods showed higher quality of prediction (GBLUP : 0.42 ± 0.16, and RR : 0.44 ± 0.16) than the bayesian lasso (lasso : 0.38 ± 0.16) or the QGBLUP (QGBLUP : 0.27 ± 0.15). Although less efficient, the QGBLUP showed that with a few QTLs, it is possible to predict between 40% and 80% of the prediction quality obtained GBLUP. The traits linked to phenology tended to be better-predicted (0.62 ± 0.17) than forage yield (0.45 ± 0.19) or quality (0.43 ± 0.14). Our results showed that it was possible to achieve


Jurij Rekelj
email: jurij.rekelj@gmail.com

Željko Predovnik
email: predovnik1@gmail.com

Peter Huemer
email: p.huemer@tiroler-landesmuseen.at

Carlos Lopez-Vaamonde
email: carlos.lopezvaamonde@inrae.fr

Systematics of Slovenian Dahlica Enderlein, 1912, subgenus Brevantennia Sieder, 1953 (Lepidoptera, Psychidae)

The subgenus Brevantennia Sieder, 1953 of the genus Dahlica Enderlein, 1912 (Psychidae: Dahlicini) is represented by a group of 10 bagworm moth species, distributed in south-west and south-east Europe northwards to the Alps and Carpathians. This study is a revision of the subgenus Brevantennia species occurring in Slovenia based on our own comprehensive field studies, DNA barcoding and morphological analysis. Dahlica (B.) santicensis (Sieder, 1957) is established as bona spec. and D. (B.) gorskikotarica Weidlich, 2015, syn. nov. is synonymised with D. (B.) santicensis. Both D. (B.

Introduction

The European Lepidoptera fauna with ca. 10,700 species (Lepiforum 2021a) is the best-known continental insect fauna in the world. However, some genera are notoriously difficult to identify due to the lack of reliable morphological characters. Amongst those "difficult" genera lies the Dahlica/Siederia complex of species (Psychidae), whose taxonomy needs revision. The difficulty of identifying specimens accurately has led to numerous misidentifications in the literature and in public genetic databases like BOLD (https://www.boldsystems.org/). Indeed, the Dahlica/Siederia species complex was excluded from a general analysis of DNA barcodes of European Lepidoptera due to inaccurate identifications [START_REF] Elzinga | Pre-and post-mating reproductive barriers drive divergence of five sympatric species of Naryciinae moths (Lepidoptera: Psychidae)[END_REF]. In this paper we use both DNA barcodes and detailed morphological analyses to try to identify a small group of species of the subgenus Brevantennia [START_REF] Sieder | Vorarbeit zu einer Monographie über die Gattung Solenobia (Lepidoptera Psychidae Talaeporiinae)[END_REF] focusing on the Slovenian fauna.

The subgenus Brevantennia includes 10 species of small psychids distributed in south-west and south-east Europe northwards to the Alps and Carpathians [START_REF] Arnscheid | Psychidae[END_REF]. Three of them, D. (B.) adriatica [START_REF] Rebel | Zur Kenntnis palaearktischer Talaeporiiden[END_REF], D. (B.) triglavensis [START_REF] Rebel | Zur Kenntnis palaearktischer Talaeporiiden[END_REF] and D. (B.) styriaca [START_REF] Meier | Ein neues Subgenus und drei neue Arten der Gattung Solenobia Dup[END_REF], have been recorded from Slovenia [START_REF] Lesar | Check list of Slovenian Microlepidoptera[END_REF][START_REF] Weidlich | Ein Beitrag zur Verbreitung und Ökologie von Brevantennia adriatica (Rebel, 1919) (Lepidoptera: Psychidae)[END_REF][START_REF] Arnscheid | Psychidae[END_REF].

In this study we revise the Slovenian Dahlica species of the subgenus Brevantennia, presenting new data based on our field collections across the country, morphological and DNA barcoding analyses and review of published records. We also describe the bionomics of each species based on our own observations in the field.

Materials and methods

Sampling

Slovenian specimens of the subgenus Brevantennia were collected in four regions in Slovenia at Gorenjska, Štajerska, Notranjska and Dolenjska, at 35 locations. In addition, we also considered material collected at primary type localities in neighbouring countries to compare and clarify the identity of some species, including D. styriaca -Austria, Steiermark, Häuslberg bei Leoben; D. santicensis [START_REF] Sieder | Fünfte Vorarbeit zu einer Monographie über die Gattung Solenobia (Lep., Psychidae -Talaeporiinae). I. Sol. Brevant. saxtalis war ein Irrtum[END_REF]) -Austria, Kärnten, Warmbad-Villach; D. siederi -Schweiz, Ticino, Monte Generoso; D. reliqua [START_REF] Sieder | Vorarbeit zu einer Monographie über die Gattung Solenobia (Lepidoptera Psychidae Talaeporiinae)[END_REF]) -Austria, Kärnten, Launsdorf; D. gorskikotarica Weidlich, 2015 -Croatia, Gorski Kotar, Mrzla Vodica;D. ilonae Weidlich, 2014 -Slovakia, Nizke Tatry, Malužinská Dolina Malužiná;D. herrmanni Weidlich, 1996 -Hungary, Mátra, Matraháza;and D. pinkeri Sieder, 1964 -Spain, Massis de l'Albera, Colera. Adult specimens examined in this study were obtained mainly by collecting larval cases with mature larvae or pupae in nature. Some adults of D. triglavensis and D. adriatica were also observed and collected by net in the early morning in their natural habitat. Collected pupae (larval cases) were later laid on filter paper and stored in plastic 250 ml containers with a perforated cover until the adults emerged. Containers were kept under laboratory conditions at room temperature of 22±1 °C, and the light / dark 18:6 h photoperiod. Humidity was regulated by gently spraying with water on a daily basis.

DNA barcoding

For the DNA barcoding analyses, one or two legs were removed from each dried specimen or from females that were stored in ethanol and then transferred to lysis plates. DNA extraction, amplification and Sanger sequencing of the DNA barcode region of the mitochondrial cytochrome c oxidase I (COI) gene (658 base pairs) were carried out at the Canadian Centre for DNA Barcoding, Guelph, Ontario, Canada (CCDB), using standard high throughput protocols [START_REF] Ivanova | An inexpensive, automation-friendly protocol for recovering high-quality DNA[END_REF].

Sequence divergences within and between species were calculated using the Kimura 2-parameter model as implemented in the Barcode of Life Data System (BOLD v4, www.barcodinglife. org) [START_REF] Ratnasingham | bold: The Barcode of Life Data System (http[END_REF]. We analyzed the COI sequences using the Neighbor-joining algorithm in PAUP* 4.0a169 (http://phylosolutions.com/paup-test/) [START_REF] Swofford | PAUP*: Phylogenetic analysis using parsimony (*and other methods)[END_REF]. The level of support for individual nodes was evaluated by nonparametric bootstrapping with 1000 replicates in using PAUP*. DNA sequences, along with the voucher data, images, and trace files are deposited in BOLD and the sequences were deposited in GenBank. Barcode Index Numbers (BINs) (Ratnasingham and Hebert 2013) were obtained from BOLD. All data are available in BOLD through the public dataset: DS-BREVANTE. (dx.doi.org/10.5883/DS-BREVANTE).

Morphology

Descriptions of morphological structures for the subgenus Brevantennia follow the current terminology by [START_REF] Sauter | Morphologie und Systematik der schweizerischen Solenobia-Arten (Lep. Psychidae)[END_REF], [START_REF] Meier | Der taxonomische Wert der Hinterflügel-Aderung bei den Arten der Gattungen Brevantennia Sieder und Solenobia Duponchel (Lep., Psych.)[END_REF] and [START_REF] Hättenschwiler | Neue Merkmale als Bestimmungshilfe bei Psychiden und Beschreibung von drei neuen Solenobia Dup. -Arten[END_REF]. We used three methods for distinguishing males. The first method distinguishes the shape of cloaking scales following [START_REF] Sauter | Morphologie und Systematik der schweizerischen Solenobia-Arten (Lep. Psychidae)[END_REF]. The scales are classified into six classes, characterised by their relative widths and the number of tines (Fig. 1a). The second, method focuses on the origin of the veins m2 and m3 from the discoidal cell in the hind wings [START_REF] Meier | Der taxonomische Wert der Hinterflügel-Aderung bei den Arten der Gattungen Brevantennia Sieder und Solenobia Duponchel (Lep., Psych.)[END_REF]. This method distinguishes six different patterns of branching, i.e.: -(I) veins are connate; -(II) veins are fused more than 2/3 of their lengths; -(III) veins are fused from 2/3 to 1/3 of their lengths; -(IV) veins are fused for a short distance; -(V) the veins originate from a single point; -(VI) veins arise separately. The third method, representing maybe the most important morphometric characteristic, is used for the descriptions of the species of this genus. A ratio is calculated by measuring the length of the phallus and dividing this value by the length of the valva (Fig. 1b). The quotient resulting from this division is converted into an index, known as the "genital index'' [START_REF] Sauter | Morphologie und Systematik der schweizerischen Solenobia-Arten (Lep. Psychidae)[END_REF].

The taxonomic key for the family follows Sauter andHättenschwiler (1991, 1999) and the nomenclature follows [START_REF] Arnscheid | Psychidae[END_REF].

For all photographs we used a Canon EOS 40D digital camera. High resolution images were taken through Olympus SZ 51 stereomicroscope and later edited using ADOBE PHOTOSHOP CS5 MASTER COLLECTION software. Measurements of genitalia were done with Olympus SZ 51 and SOFT IMAGING SYSTEM ANALYSIS software. All drawings and photographs are the work of the first author.

Specimen depositories and private collections

The following abbreviations indicate the depository of the specimens investigated:

Results

Key to the Slovenian Dahlica, subgenus Brevantennia species

Subgenus Brevantennia can be easely separated from the other Dahlicini by unique female characteristics. The most reliable and important are two: antenna noticeable short, with one to 11 segments (in Slovenian species three to seven), female exuviae slightly shrunk and curved. All of these characteristics are evident in Fig. 6b.

Analysis of the Slovenian specimens, both morphological and genetic, confirms the existence of three species in the subgenus Brevantennia. Their determination is based on male morphological characteristics (see Figs 1,9,10 Distribution. D. adriatica is distributed in the area of the Dinaric Alps. Known in Slovenia, Croatia, and Bosnia and Herzegovina [START_REF] Weidlich | Ein Beitrag zur Verbreitung und Ökologie von Brevantennia adriatica (Rebel, 1919) (Lepidoptera: Psychidae)[END_REF]: 16, 2015: 65. Arnscheid and Weidlich 2017: 74).

Slovenia is the northern border of its distribution range, with presence in Brkini and surrounding hills: Snežnik in the east, Vremščica in the north and Slavnik in the west. All Slovenian localities are ilustrated in Fig. 11.

Biology. We found this species well-represented in various biotopes with a mainly humid character. The lowest locality, where we found larval cases was a south-west facing slope of the cold valley of Potok Mrzlek (420 m), where the species inhabits sunny, mossy and lightly overgrown rocks consisting of Eocene flysch sandstone, just 10 m above the creek. Most other localities were found in south to south-west facing slopes in Illyric Ostryo-Quercetum zonal forest community (Ostryo-Carpinion orientalis) from 600-700 m elevation. The subsoil here consists of limestone. Larval cases were fixed on the south side of rocks, retaining walls and bark of trees. We noticed that the larvae are likely to avoid places with high light intensity; micro-localities shadowed with vegetation are more populated. The most typical population of this zone is Črni Vrh, Vremščica (680 m) (Fig. 2c). It is interesting that the species is also represented on south-facing meadows at higher elevations, usually just above the edge of the Illyrian Fagus sylvatica forests zone (Aremonio-Fagion) from 940-1010 m. Due to the higher elevation and consequently the colder climate, the larval cases were found on the warmest places: on the south side of rocks, usually where rocks were lying in larger groups -Devin, Župnica (1010 m).

The species is univoltine. The adults are on the wing, depending on the location from late March to early May. Remarks. The first record of the presence of D. adriatica in Slovenia dates back to the year 1919. Rebel (1919: 110) reports one male specimen caught in Istrien (H. Kozina,leg. Naufock,10. Mai '08). By the place name "H. Kozina" he probably considered the villages Hrpelje and Kozina in Slovenia, or most likely the Kozina railway station there. Travelling by train was most popular at that time. The single specimen showed slightly wider and darker wings than specimens from type locality Senj ("Zengg"), Croatia, and therefore the author defines it as "belongs probably also to adriatica". This uncertainty is likely the reason why D. adriatica was not subsequently placed on the list of the Slovenian fauna, despite the fact that that is has been confirmed as present many times by Slovenian entomologists (Mojmir Lasan, personal communication 2012). The species was officially included in the list in 2012 (Weidlich 2012: 16).

The genital index of males slightly deviates from the known values, which were 0.61-0.64 [START_REF] Weidlich | Eine neue Psychide aus Nordungarn -Brevantennia herrmanni n. sp. (Lep., Psychidae)[END_REF](Weidlich : 167, 2012: 15): 15). The new index is slightly higher, from 0.61-0.68 (Fig. 9). Dahlica (Brevantennia) santicensis [START_REF] Sieder | Fünfte Vorarbeit zu einer Monographie über die Gattung Solenobia (Lep., Psychidae -Talaeporiinae). I. Sol. Brevant. saxtalis war ein Irrtum[END_REF], stat. rev. Figs 3a,b,5,8a,b,[9][10][11] Solenobia (Brevantennia) santicensis [START_REF] Sieder | Fünfte Vorarbeit zu einer Monographie über die Gattung Solenobia (Lep., Psychidae -Talaeporiinae). I. Sol. Brevant. saxtalis war ein Irrtum[END_REF] 108. = Brevantennia gorskikotarica Weidlich, 2015, syn. nov. Type locality: Mrzla Vodica, Crni Lug, Gorski Kotar, Croatia.

Material examined. Lectotype. Labelled as follows: 1. White label bordered in red: Det. L. Sieder 1957, Sol. Brevant. santicensis Sied. Typus ♂; 2. White label: 18.5.56, Warmb.-Villach, Kärnten, leg. L. Sieder; 3. Red label: LECTOTYPUS, Dahlica (Brevantennia) santicensis, [START_REF] Sieder | Fünfte Vorarbeit zu einer Monographie über die Gattung Solenobia (Lep., Psychidae -Talaeporiinae). I. Sol. Brevant. saxtalis war ein Irrtum[END_REF], designated by Rekelj J. et al. 2022 (Fig. 3a).

The lectotype is deposited in the NHMW [Hauptsammlung -Psychidae].

The total number of syntypes is unknown (in the original description Sieder unfortunately does not specify the number of specimens of the type material), but it must be larger than those we found in the museum (NHMW) in Vienna: Austria • Kärnten, Warmbad-Villach;1♂, 11.v.1956;3♂♂, 16.v.1956;6♂♂, 17.v.1956;5♂♂, 18.v.1956;1♂, 19.v.1956;1♂, 26.v.1956;1♂, 29.v.1956;2♂♂, 26.iv.1957;1♂, 25.iv.1959;1♂, 1.v.1959;1♂, 8.v.1959;1 larval case, 18.v.1956 [labeled as a "Sack-Typus ♂" -Fig. 3b]; 6 larval cases, (iv/v)1956; 1 larval case, 28.iv.1957; 1 larval case, 28.v.1957; all Gorenjska, Gozd Martuljek;740 m;7-9.v.2002;[START_REF] Leg | coll. PCML • Several old larval cases; Karavanke, Belca[END_REF]genit. prep. №: 313, Rekelj; coll. PCML • 20♂♂, 5♀♀, with larval cases; same locality; 710 m; 28.iv.2013, (e.p. 1-5.v.2013); leg. J. Rekelj; coll. PCJR • 26♂♂, 3♀♀, with larval cases; same locality; 10.iv.2014 (e.p. 17-22.iv.2014); leg. J. Rekelj; 5♂♂ genit. prep. №: 214-218; Rekelj coll. PCJR • 1♂, with larval case; Gorenjska, Rateče;850 m;10.iv.2014(e.p. 22.iv.2014); leg. J. Rekelj; coll. PCJR • 1♂, with larval case; Julijske Alpe, Mala Mojstrovka;1600 m;17.iv.2011(e.p. 1.v.2011) 1-9.iv.2019); leg. J. Rekelj; coll. PCJR • 1♂, 3♀♀, with larval cases, several empty larval cases; Naklo, Gradišče;380 m;16.iv.2016(e.p. 17.iv.2016); leg. J. Rekelj; coll. PCJR • 9♂♂, 7♀♀, with larval cases; Kamniško Savinjske Alpe, Črnivec pod Plešivcem; 900 m; 14.iv.2013 (e.p. 20., 25.iv.2013) ; 12.iv.2014 (e.p. 2., 13., 14., 15., 18.iv.2013) D. triglavensis is distinguished by smaller class of cloaking scales (I-II), more unclear and less noticeable pattern and much lighter ground colour of forewings. Fringes are lighter creamy silver, the apex of the forewing in the area of stigma is creamy and noticeably brighter.

D. santicensis is most similar to the D. siederi by the genital index and class of cloaking scales, but siederi is distinguished by lighter (yellowish white) pattern on fore wings, noticeably brighter (creamy silver) apex of the forewing in the stigma area and by the brighter (whitish silver) fringes on forewing.

Redescription. Male (Figs 3a,4a). Exp. 10 to 13 mm, forewing length 4.5-6 mm. Specimens very variable in size and colouring. Head: Vertex and frons (Fig. 8a,b) covered with yellowish grey to grey dense hairs, labial palpi small, covered with grey hairs. Eyes small, black, ocelli lacking. Antennae relatively short, in most specimens not exceeding half length of the costal margin of the forewings, with 27-34 segments (n = 25), including scapus and pedicellus, filiform (in original description Sieder states 30-34 segments). Ventral to the first third, usually covered with broad, light creamy scales, with three to four short tines, especially when the vertex hairs are light creamy. If the hairs are darker, yellowish grey, then the antennae scales are (in first third) mixed with darker scales. In the second third, the scales become darker and gradually to the apex narrower and sparser. Forewings: (Figs 3a, 4a, 10) length 2.3 times exceeding its width (n = 4). The ground colour of forewings greyish brown. Pattern (bright spots) light grey to greyish white, distinct and well expressed, more strongly than at the other narrow scales Dahlica subgenus Brevantennia species. Discal and anal spot less well expressed or no recognizable. Cloaking scales with two to four, mostly with three short tines, classified into class II and III. Fringes silvery greyish, coloured strongly at the base, long, with two to four tines. Forewing venation with nine veins, originates from the discoidal cell. Vein pcu less well expressed, accessory cell present (n = 5). Hindwings: Colour greyish brown. Fringes medium length, with two to three tines, brightly coloured than the basic colour of hindwings. Venation with six veins originates from the discoidal cell. Veins m2 and m3 very variable, separated or approximated at the base, sometimes also fused for a short distance -class IV, V or VI. All three classes of venation differ greatly, depending on the location, or even the year of collecting specimens. Thorax: Colour greyish brown, in brighter specimens grading into creamy brown. Legs greyish brown, foretibia without epiphysis. Midtibia with one pair of spurs, hindtibia with two pairs. All legs with five tarsal segments. Abdomen: Dark brown colour, becoming lighter towards the genitalia. Terminal segments covered with creamy or pale yellow hairs. Genitalia: Typical for Dahlicini. Tegumen relatively broad, with a shallow groove at the front. Valva narrow and long. Dorsal margin of valva towards the top slightly curved, apex slightly extended, without any special characteristics. Clavus long, pointed and curved at the end. Phallus quite short, curved, without specifics. Genital index 0.64-0.80, average 0.72 (n = 52). The index varies depending on location, as expressed in Fig. 5.

Female (Fig. 4b). Wingless, ground colour pale yellow, with broad, light brown plates dorsally. Freshly hatched specimens having a slight greenish appearance. Dimensions: three mm long and less than one mm wide. Eyes small, black, ocelli lacking. Antennae short with three to seven segments (including scapus and pedicellus). Tibiae of forelegs without epiphysis, all legs usually with three-segmented tarsi, fused tarsi in two segments rare but present. 7 th abdominal segment densely covered with creamy white hairs.

Female exuvia pale brown colour, curved -typical for the subgenus Brevantennia. Larval cases. Typical for Dahlicini, cases are small, more or less round and slightly rounded on the edges. They are composed of finely chewed dark brown particles of bark, small pieces of soil and brighter pieces of sand. There is an inevitable slight deviation in composition of larval cases as a result of the different surfaces available in the natural habitat. Male larval cases are 5.5-6.5 mm long and 2 mm wide; female cases are smaller, 4.5-5 mm long and 1.5 mm wide.

Morphological variation. We noticed considerable variation between populations and also between specimens from the same population, reflected in the colour of the dense vertex hairs, intensity of wing colouration and the genital index. Northern populations (Warmbad-Villach, Strmec na Predelu, Gozd Martuljek) have mostly yellowish-grey coloured vertex hairs, but among those we also found about 10% grey-coloured specimens. Their abundance increases towards the south and in southern localities (Retje, Trava) only grey-coloured specimens can be found. Differences between dense vertex hair colours are shown in Fig. 8a,b.

Intensity of wing colouration is also quite variable, but it seems that is not location dependent. Within one population (Gozd Martuljek locality) we found specimens with intensive colouration alongside more pale specimens.

Differences related to the geography of the locality are also noticeable in the genital index. Higher values were found in moths from northern locations of a more alpine character. Southern locations produce lower values, but they do not fall linearly proportionate to the latitude and the values are overlapping. Comparison of genital indexes with locations is expressed in Fig. 5.

Morphologically, the nearest species to D. santicensis is D. siederi [START_REF] Weidlich | Brevantennia gorskikotarica n. sp. aus Kroatien und ein weiterer Beitrag zu den Vorkommen von Brevantennia adriatica (Rebel, 1919) auf dem Balkan (Lepidoptera, Psychidae)[END_REF], which has the same class of cloaking scales (II-III) and shows almost the same genital index (Fig. 9). Synonymy. [START_REF] Weidlich | Brevantennia gorskikotarica n. sp. aus Kroatien und ein weiterer Beitrag zu den Vorkommen von Brevantennia adriatica (Rebel, 1919) auf dem Balkan (Lepidoptera, Psychidae)[END_REF] described D. gorskikotarica from Croatia -Gorski Kotar. We have examined the following material from the type locality: Croatia • 8 larvae, 1 old exuviae ♀, Gorski Kotar, Crni Lug, Mrzla Vodica; 880 m; 18.x.2015; leg. J. Rekelj; coll. PCJR • 14♂♂, 8♀♀, with larval cases, same locality; 2.iv.2017 (e.p. 6-10.iv.2017); leg. J. Rekelj, 5♂♂ genit. prep. №: 370-374, Rekelj; coll. PCJR • 19♂♂, 6♀♀, with larval cases; Gorski Kotar, Crni Lug, Vela Voda; 747 m; 2.iv.2017 (e.p. 6-10.iv.2017); leg. J. Rekelj; 5♂♂ genit. prep. №: 375-379, Rekelj; coll. PCJR.

In addition, we have DNA barcoded one specimen from the type series (DNA barcode sample: LEATC004-13) from Mrzla Vodica. This specimen falls within the same BIN (BOLD:AAQ1227) as D. santicensis (Suppl. material 1: Table S1). In addition, we found no diagnostic morphological character (forewing colouration and pattern, class of cloaking scales and genital index) to distinguish D. gorskikotarica from Slovenian populations of D. santicensis. Both molecular and morphological data show that D. gorskikotarica is a synonym of D. santicensis: Brevantennia gorskikotarica Weidlich, 2015, syn. nov.

= Dahlica (Brevantennia) santicensis [START_REF] Sieder | Fünfte Vorarbeit zu einer Monographie über die Gattung Solenobia (Lep., Psychidae -Talaeporiinae). I. Sol. Brevant. saxtalis war ein Irrtum[END_REF].

Distribution. The species is distributed in Austria, Slovenia and northern Croatia. In Austria by [START_REF] Sieder | Fünfte Vorarbeit zu einer Monographie über die Gattung Solenobia (Lep., Psychidae -Talaeporiinae). I. Sol. Brevant. saxtalis war ein Irrtum[END_REF] in Kärnten: Warmbad-Villach (type locality), Arnoldstein -Gailtal, Rattendorfer Alm -Karnischen Alpen and Spitzegels by Hermagor. Confirmed localities are also Faaker Zee -Kärnten and Lengberg -Osttirol [START_REF] Deutsch | Beitrag zur Lepidopterenfauna Osttirols, Österreich -Teil V (Insecta: Lepidoptera)[END_REF]: 10 -as B. styriaca, Lepiforum 2021b -as B. styriaca). In Croatia the species is known from Gorski Kotar (Weidlich 2015 -as B. gorskikotarica).

In Slovenia we found D. santicensis in Gorenjska, Notranjska, Dolenjska and Štajerska regions, in total 22 localities, which indicates a wide distribution across the country. All Slovenian localities are illustrated in Fig. 11.

Etymology. Santicum, Roman Castrum near Warmbad-Villach. Biology. At the type locality (Warmbad-Villach, Austria), larval cases can be found on the northern slopes of mixed forest, more often in shady areas. The subsoil consists of limestone. Lar- val cases are present in particular on conifers such as Larix decidua Mill., Pinus sylvestris L. and Picea abies L. and also on rocks. They are mainly found in shady parts of tree trunks, which are covered with lichens and algae and at a height of up to two metres above the ground.

Recent investigations in Slovenia show that the species is more widespread than previously known, more adaptable and occurring in different habitats in cold areas. In lowlands, where the temperatures are higher on average, we found larval cases in northern exposed slopes, in shady humid areas of mixed forest dominated by conifers, mostly Pinus sylvestris and Picea abies and in cold and damp shady valleys of creeks. Sometimes the river or lake allows cooler microclimate conditions, so animals there can also be found on the sunny, south exposed positions (location, Gozd Martuljek -Fig. 4c). The highest locality where the species has been observed is on the subalpine southern slopes of Mala Mojstrovka, where we found larval cases on the cliffs and rocks just a little above the tree line -the Larix decidua and Pinus mugo Turra zone. At all locations, the subsoil consists of limestone and larval cases were mostly found on the shady trunks of coniferous trees, wooden fences, and rocks. Only exceptionally, at Črnivec and Cezlak-Lukanja, the substrate is of volcanic origin. Larval cases were found there, mainly on the tuffite boulders and stones which are, unlike limestone, covered with algae and mosses in abundance. They were found also on the walls of the old wooden buildings, such as an old arbour in the forest and at bus stations. The lowest elevation was in the valley of the stream Gračnica is 392 m and the highest was at 1600 m on Mala Mojstrovka mountain.

Research in Slovenia shows that the species is univoltine. Pupation occurs between late March and early April. The flight period of adults lasts from the beginning of April to early May, depending on the location and year. Males hatch in the evening and become active in the early morning, after occurrence of females, between 05:30-07:00.

D. santicensis in Slovenia cohabits with the following species of bagworms: Dahlica triquetrella f. parth. (Hübner, 1813); Dahlica lichenella (Linnaeus, 1761) f. bisex.; Dahlica croatica Weidlich, 2016;Dahlica sp.;Taleporia tubulosa (Retzius, 1783); Proutia comitella (Bruand, 1853); Psyche casta (Pallas, 1767) and Bacotia claustrella (Bruand, 1845). An exception is the locality Mala Mojstrovka, where the species cohabits only with Dahlica (Siederia) meierella [START_REF] Sieder | Vierte Vorarbeit zu einer Monographie über die Gattung Solenobia[END_REF] and Dahlica triquetrella f. parth.

Remarks. Historical overview: The discovery and description of Dahlica (Brevantennia) santicensis has an interesting and informative history and is maybe among the most interesting for species in this group. For this reason, it was mentioned several times in different publications (Arnscheid, 1988(Arnscheid, , 1993)). The story began in the year 1952, when Mr. Herbert Hölzl from Klagenfurt found small larval cases while climbing to Rattendorfer Alm in the Carnic Alps. The following year Mr. Thurner, also from Klagenfurt picked up a lot of specimens. Those specimens were later on described as a new species Dahlica (Brevantennia) saxatilis [START_REF] Sieder | Zweite Vorarbeit über die Gattung Solenobia (Lepidopt., Psychidae, Talaeporiinae)[END_REF]. Two years after the description, Sieder himself visited that locality, one week after another specialist, H. Meier. Together, they found, that two species of Dahlicini occur syntopically -Dahlica (Siederia) alpicolella [START_REF] Rebel | Zur Kenntnis palaearktischer Talaeporiiden[END_REF] and the newly described D. saxatilis. This is nothing special, but we now know that Sieder used two different species for his description: males of D. alpicolella and females of his new species D. saxatilis. Later, he corrected his mistake in a new article "Fünfte Vorarbeit über die Gattung Solenobia", synonymised D. saxatilis with D. alpicolella and added a new and corrected description of the new species D. santicensis, this time with matched males [START_REF] Sieder | Fünfte Vorarbeit zu einer Monographie über die Gattung Solenobia (Lep., Psychidae -Talaeporiinae). I. Sol. Brevant. saxtalis war ein Irrtum[END_REF]. For the new type locality, he chose the forest of Warmbad-Villach (the first specimens from that locality he got in the year 1955 from Trasischker), but considering the imprecisely assembled earlier material, he visited the place and collected the type material himself. Approximately one month previously, Meier's description of Dahlica (Brevantennia) styriaca was published. In spite of the fact that he had material from Rattendorfer Alm, Meier neither compared his new species with Sieder's D. saxatilis, nor did he mention this taxon in his work. However, both D. styriaca and D. santicensis were known as separate species (bona species) for 31 years, until Arnscheid's publication in 1988, where he established the species D. santicensis as synonym of D. styriaca. Here we give his reasons: "During a personal conversation in September 1976, Sieder told the author that both taxa are most likely identical. Investigations to date of numerous specimens, males and females of both species, actually showed no significant differences which would exceed the normal range of variations, that are specific to a particular species". Unfortunately, the authors did not use the already well-known and generally confirmed methods of determination [START_REF] Sauter | Morphologie und Systematik der schweizerischen Solenobia-Arten (Lep. Psychidae)[END_REF], especially the genital index. The fact is that this species remained unnoticed until today. Only recent comparisons of Slovenian populations with already known species (according to established morphological and newer genetic methods), have shown that they are clearly separated from Styrian populations and that they are identical to those from the forest of Warmbad-Villach. (see below) which helps populations to persist in high alpine areas where the weather can be poor towards the end of the season. We found on the same stone old empty larval cases from previous years and new fresh cases with pupae from the current season as well as young larvae of different sizes. This behaviour at this elevation indicates that the species is semivoltine, likely with a two-year cycle (Sieder 1953: 122). Adults emerge from the beginning of June to mid-July, about three weeks after snow melt. They are active in the morning, when the habitat is illuminated by the first sunlight. D. triglavensis in Kanin locality cohabits with the following species of bagworms: Dahlica klimeschi [START_REF] Sieder | Vorarbeit zu einer Monographie über die Gattung Solenobia (Lepidoptera Psychidae Talaeporiinae)[END_REF]; Dahlica (Siederia) meierella; Proutia raiblensis (Mann, 1870) and Leptopterix hirsutella (Denis & Schiffermüller, 1775).

Dahlica (Brevantennia) styriaca (Meier, 1957)

On the Karavanke mountains we found this species on two localities at very different elevations. The first locality Jurčkov vrh, Belca lies on the south-eastern steep, rocky slope of entry Belca creek in a Genisto januensis -Pinetum zonal forest community, from 840-907 m. The vegetation is dominated by Pinus sylvestris L., Fraxinus ornus L., and Ostrya carpinifolia Scop. The larval cases were found on the underside of stones or gravel, also on the low areas of larger rocks or cliffs. The larvae are quite hidden throughout the season and feed on mosses, decomposing plants and lichens. At this elevation the species is univoltine. This behaviour has already been observed for D. triglavensis by Sieder (1953: 122). Larvae begin pupation in mid March to early April. The flight period of adults is in April and depends on the duration of the winter and the amount of snow cover.

At the Jurčkov vrh locality, D. triglavensis cohabits with the following bagworms: Dahlica triquetrella f. parth; Dahlica (Siederia) meierella; Taleporia tubulosa; Typhonia sp.; Psyche casta; Bijugis bombycella (Denis & Schiffermüller, 1775); Rebelia sp.; Canephora hirsuta (Poda, 1761); Leptopterix plumistrella (Hübner, 1793); Megalophanes viciella (Denis & Schiffermüller, 1775) and Sterrhopterix sp.

The second Karavanke locality lies on the rocky slope of eastern ridge of Mt. Stol, just below the summit at an elevation of 2140 m in the alpine zone. The subsoil consists of limestone. The larval cases were found there in a microhabitat where rocks are in contact with soil, on the underside of stones or gravel and on the lower parts of larger rocks. Among the collected larval cases were also found a few active young larvae which indicate that the species is semivoltine, likely with a two-year cycle. At this locality we observed activity of adults as early as the end of June.

In Stol this species cohabits with the following species of bagworms: Dahlica goltella Rekelj & Predovnik, 2014; Dahlica klimeschi [START_REF] Sieder | Vorarbeit zu einer Monographie über die Gattung Solenobia (Lepidoptera Psychidae Talaeporiinae)[END_REF]; Proutia raiblensis and Epichnopterix ardua Mann, 1867.

Remarks. Subendemic, type species of the subgenus Brevantennia (Sieder, 1953: 120), with type locality in Slovenia: Julijske Alpe, Triglav, 2400 m (Rebel 1919: 111).

Our DNA barcoding study revealed two distinct genetic lineages within the D. triglavensis group, representing different BINs (Suppl. material 1: Table S1). The first lineage consists of specimens from Julijske Alpe (BOLD:ABU8640), and a second one from Karavanke mountains (BOLD:ABU8641). Morphologically, these lineages are remarkably similar and despite precise studies of adults and immature stages of many specimens, no tangible morphological differences could be found that would exceed the normal range of variation of each population.

However, the morphology-based taxonomy and the distribution boundaries of these groups remain poorly defined, so we have decided to keep the same name for both populations.

DNA barcoding analysis

We DNA barcoded 52 specimens belonging to eight named Dahlica species of the subgenus Brevantennia (Table 1). The sequences formed nine different BINs seven of which were new to BOLD (Table 1) with a genetic intraspecific variability ranging from 0-2.51% with a mean of 0.91% (Table 1). The minimum nearest-neighbour distance varies from 2.4-4.97% (Table 1). Out of the eight named species barcoded, five had their own unique BIN (Table 1; Fig. 12), since D. triglavensis shows two different BINs which are 2.4% divergent (Table 1) and D. santicensis and D. siederi share a BIN. Indeed, D. santicensis shows a high level of intraspecific variability with a maximum intraspecific genetic distance of 2.57% (Table 1). According to BOLD, the nearest neighbour of the latter is a Dahlica species at 4.97% (p-distance) (Table 1). All 19 specimens of D. santicensis share the same BIN (BOLD:AAQ1227) (Fig. 12, Table 1). In addition, we DNA barcoded an Italian specimen of D. siederi (STG 216) which shares the same BIN (BOLD:AAQ1227) as D. santicensis (Table 1, Fig. 12 , Suppl. material 1: Table S1).

One specimen from Crimea (TLMF Lep 07217) forms its own BIN (BOLD:ACM5956) and shows a high genetic divergence of 4.33% to a Dahlica species (Suppl. material 1: Table S1). We need more specimens to assess its status and it is therefore treated here as an unconfirmed candidate species (sensu Lopez-Vaamonde et al. 2021).

Discussion

Here we studied the Dahlicini, a very challenging group in terms of species delimitation and identification. We focused primarily on the species from Slovenia, but we briefly touched also on problems of related species from neighbouring countries. Due to its rich geological development, Slovenia has a remarkable species diversity in a relatively small geographical area, and this is reflected in the subgenus Brevantennia. Accurate identification of Dahlica species is challenging and can only be successful done based on all listed external features in combination with the genital indices (Arnscheid and Weidlich 2017: 39). In our study group, the genital index method proved to be the most useful approach which, in combination with the distinguishing shape of cloaking scales, allowed us to identify all Slovenian species. Here we updated the known values of these indices as a result of our analysis of a larger number of specimens from different localities. We found that wing venation is an unreliable diagnostic character at species level. Our results have confirmed the presence of both D. adriatica and D. triglavensis in Slovenia. D. triglavensis shows two different BINS separated geographically by the Sava valley: BOLD:A-BU8640 occurs in Julijske Alpe and BOLD:ABU8641 occurs in Karavanke mountains. We have not found any morphological differentiation between both allopatric populations representing another European Lepidoptera species with multiple BINs [START_REF] Lopez-Vaamonde | Evaluating DNA Barcoding for Species Identification and Discovery in European Gracillariid Moths[END_REF]. It would be interesting to see whether there are any areas where representatives of both BINS enter into contact.

Both morphological and DNA barcoding results confirmed the differentiation between D. styriaca and D. santicensis. Indeed, we have obtained DNA barcode data of D. styriaca from the type locality in Austria and DNA barcodes of D. santicensis from populations distributed from Austrian Carinthia, East Tyrol, Slovenia, Romania and Croatia. Our morphological analysis confirmed that D. santicensis is a good species and the recently described D. gorskikotarica is a new synonym of this species.

D. styriaca does not occur in Slovenia, but further field work is needed in the less studied areas of Slovenia, such as the mountain range of Kozjak, located north of the Drava River in the border with Austria, an area which lies on the Eurasian plate.

Two species, D. siederi and D. santicensis, share the same BIN. These two species are morphologically very similar having almost the same genital index and type of cloaking scales [START_REF] Weidlich | Brevantennia gorskikotarica n. sp. aus Kroatien und ein weiterer Beitrag zu den Vorkommen von Brevantennia adriatica (Rebel, 1919) auf dem Balkan (Lepidoptera, Psychidae)[END_REF]. However, these two species can be separated clearly by the ground colour of the forewings which is much lighter in D. siederi than in D. santicensis. Furthermore, the forewing pattern is creamy white, unlike that of D. santicensis, which is light grey. The forewing apex in the area of the stigma is noticeably lighter in D. siederi, creamy silver in ground colour, unlike D. santicensis which has this area silvery grey, the same colour as the ground colour or fringes. There are also strong differences in the choice of biotope. D. siederi prefers sunny subalpine slopes above the tree line, where larval cases are found on cliffs, walls and rocks at elevations from 1600-1700 m, whereas D. santicensis is a highly eurytopic species. The latter inhabits shaded humid areas of mixed forest dominated by conifers, mostly Pinus sylvestris and Picea abies, from the lowlands to the subalpine zone, between 400-1600 m. Unfortunately, we only DNA barcoded one specimen of D. siederi from a single locality (Italy, Monte Baldo). More specimens of D. siederi are needed for DNA barcoding analysis, in particular from the type locality (Schweiz, Monte Generoso) to test the hypothesis that D. santicensis and D. siederi are species each with multiple BINs.

Figure 1 .

 1 Figure 1. Important morphological structures. a. Six classes of forewing-cloaking scales after Sauter (1956); b. Male genitalia of subgenus Brevantennia sp. in lateral view. Measured distances: PL= Phallus length (black dotted); VL= Valve length, after Sauter (1956) -modified; c. Six classes of the origin of the veins m2 and m3 from the discoidal cell in the hind wings after Meier (1958).

Figure 2 .

 2 Figure 2. D. adriatica. a. Adult male resting on rock, Potok Mrzlek; b. Fresh adult female in assembling mode, Potok Mrzlek; c. Illyric Ostryo-Quercetum zonal forest community, natural habitat of D. adriatica, Črni Vrh, Vremščica, 680 m.

  ; leg. J. Rekelj; DNA barcode sample: TIPSY125-12; 1♂ genit. prep. №: 233, Rekelj; coll. PCJR • 1♂, with larval case; same locality; 9.v.2020 (e.p. 11.v.2020); leg. J. Rekelj; genit. prep. №: 380, Rekelj; coll. PCJR • 2♀♀, with larval cases, several empty larval cases; Julijske Alpe, Zgornja Trenta, Mlinarica; 1100 m; 9.v.2020, (e.p. 10.v.2020); leg. J. Rekelj; coll. PCJR • 8♂♂, 12♀♀, with larval cases; Julijske Alpe, Strmec na Predelu; 1015 m; 21.v.2017; (e.p. 25-30.v.2017); leg. J. Rekelj; coll. PCJR • 27♂♂, 2♀♀, with larval cases; same locality; 9.v.2020, (e.p. 9-15.v.2020); leg. J. Rekelj; 5♂♂ genit. prep. №: 284-287, Rekelj; coll. PCJR • 3 larval cases with ♂ exuvia, 1 larval case with ♀ exuvia; Cerkno, Kladje; 780m; 21.vi.2017; leg. J. Rekelj; coll. PCJR • 2♂♂, with larval cases, several old cases; Zgornja Sorica [road to Soriška Planina]; 940 m; 23.iii.2019, (e.p.

Figure 3 .

 3 Figure 3. D. santicensis. a. Lectotype male with original labels in NHMW; b. Syntype male (mislabelled as neotype), larval case with original labels in NHMW.

  Diagnosis. (Figs 9, 10) D. santicensis can easily be distinguished from the wide-cloaking scaled species: D. pinkeri (class III-IV), D. adriatica (class IV-V), D. herrmanni (class IV-V) and D. estrela (class V-VI) by the narrower cloaking scales (class II-III), and also by the different genital index. D. santicensis (0.64-0.80) differs from D. reliqua (0.94-0.956), D. styriaca (0.99-1.14) and D. ilonae (1.19-1.24), by a much lower genital index and more distinct pattern on the forewings.

Figure 4 .

 4 Figure 4. D. santicensis. a. Fresh adult male resting on rock, Gozd Martuljek; b. Fresh adult female transmitting pheromones, Gozd Martuljek; c. Mixed forest with Scots pine, natural habitat of D. santicensis, Gozd Martuljek, 740 m.

Figure 5 .

 5 Figure 5. Graphical representation of genital indexes on different localities of D. santicensis.

Figure 6 .

 6 Figure 6. D. triglavensis. a. Adult male resting on rock, Kanin; b. Female on the larval case in attracting mode, Kanin; c. Rocky slope with alpine vegetation, natural habitat of D. triglavensis, Kanin, 2250 m.

FigsFigure 7 .

 7 Figs 8f, 9, 10, Solenobia (Brevantennia) styriacaMeier, 1957: 59.

Figure 8 .Figure 9 .

 89 Figure 8. Heads, colouration of vertex and frons dense hairs. a. D. santicensis, Gozd Martuljek; b. D. santicensis, Retje, Ribnica; c. D. triglavensis, Jurčkov vrh, Belca; d. D. triglavensis, Kanin; e. D. adriatica, Potok Mrzlek; f. D. styriaca, Häuslberg bei Leoben.

Figure 10 .

 10 Figure 10. Comparison of patterns and cloaking scales of male forewings: Slovenian and surrounding subgenus Brevantennia species.

Figure 11 .

 11 Figure 11. Distribution map of subgenus Brevantennia species in Slovenia. 1. Prelovce, Vremščica; 2. Potok Mrzlek, Ribnica; 3. Žlebovi, Koritnice; 4. Devin, Župnica; 5. Slavnik; 6. Misliče, Brkini; 7. Pregarje, Huje; 8. Obrov, Brkini (Weidlich 2012); 9. Gozd Martuljek; 10. Rateče, Gorenjska; 11. Mala Mojstrovka; 12. Mlinarica, Zg. Trenta; 13. Strmec na Predelu; 14. Kladje, Cerkno; 15. Zgornja Sorica; 16. Gradišče, Naklo; 17. Črnivec pod Plešivcem; 18. Strmec, Raduha; 19. Cezlak, Oplotnica; 20. Tolsti vrh, Mislinja; 21. Paški Kozjak; 22. Mrzlo Polje, Gračnica; 23. Laniše, Hrušica; 24. Nanos; 25. Kurešček, Želimlje; 26. Mala Slevica, Velike Lašče; 27. Ravni vrh, Sodražica; 28. Retje, Ribnica; 29. Breg pri Ribnici na Dolenjskem; 30. Trava, Podplanina; 31. Triglav, Julijske Alpe; 32. Kanin, Julijske Alpe; 33. Mangart, Julijske Alpe; 34. Stol, Karavanke; 35. Belca, Karavanke.

Figure 12 .

 12 Figure 12. Neighbor-Joining tree (uncorrected p-distance) of subgenus Brevantennia COI (DNA barcode) sequences. Each specimen is identified by its ProcessID code (see Suppl. material 1: Table S1). Values above branches are bootstrap supports above 70. Collecting data of the corresponding specimens, from top to bottom as follows: D. pinkeri -Spain, Portbou, Colera; D. santicensis -Slovenia, Gozd Martuljek; D. siederi -Schweiz, Monte Generoso; D. sp. -Ukraine, Crimea, Sample ID: TLMF Lep 07217, BIN: BOLD:ACM5956; D. triglavensis -Slovenia, Kanin; D. ilonae -Slovakia, Malužiná; D. styriaca -Austria, Häuslberg bei Leoben; D. adriatica -Slovenia, Vremščica; D. reliqua -Austria, Launsdorf.

Dahlica (Brevantennia) adriatica (Rebel, 1919)

  ). 1 Cloaking scales wide, class IV-V. Genital index low, 0.61-0.68 ....................................... ..............................................................

	-	Cloaking scales narrow, class I-III. Genital index higher 0.64-0.86 ...............................2

Dahlica (Brevantennia) adriatica (Rebel, 1919)

	Figs 2a-c, 8e, 9-11
	Solenobia adriatica Rebel, 1919: 110

  Austria • 1♂; Kärnten, Warmbad-Villach; 16.v.1956, leg. L. Sieder; coll. PCMP • 9♂♂, 10♀♀, with larval cases; Kärnten, Warmbad-Villach; 540 m; 28.iv.2013 (e.p. 28.-30.iv.2014); leg. J. Rekelj; 5♂♂ genit. prep. №:184-188, Rekelj; coll. PCJR • 12♂♂, 2♀♀, with larval cases; Kärnten, Faaker Zee; 540 m; 21.iv.2013 (e.p. 25-30. iv.2014); leg. J. Rekelj; coll. PCJR • 1♂, 1♀, with larval cases; Kärnten, Loibltal; 700 m; 17.iii.2012 (e.p. 20.iii.2012); leg. J. Rekelj; DNA barcode sample: 1♂ TIPSY620-12; coll. PCJR • 2♂♂, with larval cases; Osttirol, Lengberg; 800m, 4.iv.2010 e.l.; leg. H. Deutsch; DNA barcode sample: 1♂ TIPSY155-12; 1♂ genit. prep. №: 234, Rekelj; coll. PCJR. Slovenia • 1♂, with larval case;

leg. L. Sieder; coll. NHMW. Other material.

  ; DNA barcode sample: 1♂ TIPSY672-15; 5♂♂ genit. prep. №: 228-2232, Rekelj; leg. J. Rekelj; coll. PCJR • 6♂♂, with larval cases, same locality; 9.iii.2014 (e.p. 20-22.iii.2014); leg. J. Rekelj; coll. PCJR • 1♂, 5♀♀, with larval cases; Kamniško Savinjske Alpe, Raduha, Strmec [Pečovnik]; 730 m; 19.iv.2014 (e.p. 20-24.iv.2014); leg. Ž. Predovnik; DNA barcode sample: 1♂ TIPSY736-15, 1♀ TIPSY734-15; coll. PCŽP • 1♀, with larval case, 3 adult larvae; same locality; 4.iv.2015 (e.l.-e.p. 14.iv.2015); leg. Ž. Predovnik; coll. PCŽP • 3♀, with larval

	cases; Pohorje, Oplotnica, Cezlak-Lukanja [Oplotnica creek]; 789 m; 29.iii.2014 (e.p. 4-6.iv.2014); leg. Ž. Predovnik;
	DNA barcode sample: 1♀ TIPSY733-15; coll. PCŽP • 1♀, with larval cases; same locality; 28.iii.2015 (e.l. 22.4.2014);
	leg. Ž. Predovnik; coll. PCŽP • 1♀, with larval cases; same locality; 11.iv.2015 (e.l. 21.iv.2014); leg Ž. Predovnik; coll.
	PCŽP • 1♀, with larval case; Mrzlo polje [Gračnica creek]; 392 m; 28.iii.2012 (e.p. 5.iv.2012); leg. Ž. Predovnik; DNA

barcode sample: 1♀, TIPSY718-15; coll. PCŽP • 1♀, with larval case; same locality;

13.iii.2014 (e.p. 18.iii.2014)

; leg Ž. Predovnik; coll. PCŽP • 2♂, with larval cases; Kalce [Logatec], Hrušica, Laniše; 640 m; 23.iii.2012 (e.p. 2.iv., 7.iv.2012); leg. Ž. Predovnik; DNA barcode sample: 1♂ TIPSY628-12; coll. PCŽP • 1♂, with larval case; Notranjska, Nanos; 1250 m; 31.v.2004, leg J. Skyva; genit. prep. №: 367, Rekelj; coll. PCJR • 27 old larval cases, Želimlje, Kurešček [Stara žaga]; 680 m; 14.i.2012; leg. Ž. Predovnik; coll. PCŽP • 2♂♂, with larval cases; same locality; 23.iii.2012 (e.l.-e.p. 2.iv.2012);

  leg. Ž. Predovnik; DNA barcode sample: 1♂ TIPSY629-12; 1 ♂ genit. prep. №:118, Rekelj; coll. PCŽP • 2♂♂, with larval cases; same locality; 5.iv.2013 (e.p. 15-16.iv.2013); leg. Ž. Predovnik; 1♂ genit. prep. №:122, Rekelj; coll. PCŽP • 6♂♂, 4♀♀, with larval cases; same locality

  ; leg. Ž. Predovnik; 4♂♂ genit. prep. №:119-122 Rekelj; coll. PCŽP • 7♂, 1♀, with 61 larval cases;Ribnica, Retje; 843 m; (15.iv.2013 (e.p. 25-27.iv.2013); leg. Predovnik; coll. PCŽP and PCJR • 15 larval cases, exuviae ♂♂ and ♀♀, Podplanina, Trava; 660 m; 7.iv.2012; leg. Ž. Predovnik; coll. PCŽP • 2♀, with larval cases; same locality; 12.iv.2014 (e.p. 16.iv., 21.iv.2014), [ca.10 minutes long copula: 1♂ Retje x 1♀ Trava]; leg. Ž. Predovnik; DNA barcode sample: 1♀ TIPSY741-15; coll. PCŽP • Several old cases, exuviae ♂ and ♀; Paški Kozjak [Mountain Lodge]; 1028 m; 17.ii.2019; leg. Ž. Predovnik; coll. PCŽP • 2♂, 7♀, with larval cases; 13.iv.2019 (e.p. 14.iv-22.iv. 2019); leg. Ž. Predovnik; coll. PCŽP • 17 old larval cases; Mislinja, Tolsti vrh [Polenica creek]; 805 m; 23.xii.2018; leg. Ž. Predovnik; coll. PCŽP • 4♂, 7♀, with larval cases; same locality; 13.iv.2019 (ex.p. 16.iv-22.iv.2019); leg. Ž. Predovnik; coll. PCŽP • A few old larvae cases; Sodražica, Zg. Globel, Ravni vrh; 728 m; 30.iii.2015; leg. Ž. Predovnik; coll. PCŽP • 3 old larvae cases; Velike Lašče, Mala Slevica; 608 m; 15.iii.2015; leg. Ž. Predovnik; coll. PCŽP • 2 old larvae cases; Ribnica, Breg pri Ribnici na Dolenjskem; 494 m; 30. iii.2015; leg. Ž.

	Ž. Predovnik; 4♂♂, genit. prep. №:103, 115-117, Rekelj; coll. PCŽP • 44♂♂, 58♀♀, with 150 larval cases, fresh exuviae
	♂, 4 adult larvae; same locality; 12.iv.2014 (e.l.-e.p. 12-22.iv.2014); leg. Ž. Predovnik; DNA barcode sample: 1♂ TIP-
	SY742-15; 6♂, genit. prep. №:104-107, 113,114, Rekelj; coll. PCŽP • 2♂♂, with 8 larval cases; same locality; 30.iii.2015
	(e.p. 9-10.iv.2015); leg. Ž. Predovnik; coll. PCŽP.

Table 1 .

 1 Summary statistics for specimens included in this study. Dmin_NN heterospecific = minimum distance to nearest neighbour; NN= nearest neighbour; BIN Dmax= maximum intraspecific distance within a BIN.

	Species	No. in	BIN name	BIN geography	BIN	BIN Dmax	Dmin_NN	NN	NN
		dataset			Average	(P-distance)	heterospecific	heterospecific	heterospecific
					(P-distance)		(P-distance)	BIN	Identification
	Dahlica (Brevantennia)	8	BOLD:ACX7360 Slovenia, Austria,	0.35	1.22	4.33	BOLD:ACS2709	Siederia
	adriatica (Rebel, 1919)			Bosnia and
				Herzegovina
	Dahlica (Brevantennia)	7	BOLD:ACJ3876	Slovakia,	0.84	1.44	4.01	BOLD:ACC7077 Brevantennia
	ilonae Weidlich, 2014			Hungary,					styriaca
				Romania,
				Ukraine
	Dahlica (Brevantennia)	1	BOLD:ACX7360	Spain	NA	NA	4.51	BOLD:ADD5884	Dahlica
	pinkeri Sieder, 1964								nickerlii
	Dahlica (Brevantennia)	2	BOLD:ACC7257	Austria	0	0	4.81	BOLD:ADQ8354	Dahlica
	reliqua (Sieder, 1953)								exulans
	Dahlica (Brevantennia)	19	BOLD:AAQ1227 Austria, Slovenia,	1.25	2.57	4.97	BOLD:AED7194 Dahlica sp.
	santicensis (Sieder, 1957)			Croatia, Romania
	Dahlica (Brevantennia)	1	BOLD:AAQ1227	Italy
	siederi (Sauter, 1954)
	Dahlica	1	BOLD:ACM5956	Crimea	NA	NA	4.33	BOLD:ADQ8354	Dahlica
	(Brevantennia) sp.								exulans
	Dahlica (Brevantennia)	2	BOLD:ACC7077	Austria	0	0	4.01	BOLD:ACJ3876	Dahlica
	styriaca (Meier, 1957)								triquetrella
	Dahlica (Brevantennia)	3	BOLD:ABU8640	Slovenia	0	0	2.4	BOLD:ABU8641 Brevantennia
	triglavensis (Rebel, 1919)								triglavensis
	Dahlica (Brevantennia)	8	BOLD:ABU8641	Slovenia	0.23	0.92	2.4	BOLD:ABU8640 Brevantennia
	triglavensis								triglavensis
	(Rebel, 1919)
	TOTAL	52

Acknowledgements

We would like to express our cordial thanks to Sabine Gaal (Austria) for giving access to study the curated collection in NHMW. We also express our sincere gratitude to Mojmir Lasan (Slovenia). His knowledge about Psychidae helped us to make the first steps in studying this interesting family. Our special thanks go to František Fiala (Czech Republic) for the possibility of study a collection of late Miloslav Petrů, which is now under his care. We are grateful to Stanislav Gomboc (Slovenia) for helpful comments on the manuscript and assistance in processing of DNA samples. Michael Weidlich (Germany) provided valuable material for DNA barcoding. Our sincere thanks also to the editor David Lees and both reviewers Christof Zeller and Thomas Sobczyk for insightful comments that helped to improve the manuscript. We thank Marijan Govedič (Slovenia) for providing valuable literature and Colin W. Plant, (UK), for linguistic improvements and some comments on the paper. DNA barcoding was funded by FEDER InFoBioS project (EX011185) to CLV, and by grants of the Promotion of Educational Policies, University and Research Department of the Autonomous Province of Bolzano -South Tyrol and the Austrian Federal Ministry of Science, Research and Economics to PH. The authors are grateful to the staff at the Centre for Biodiversity Genomics at Guelph for their assistance in particular Claudia Steinke, Ramya Manjunath, Miduna Rahulan, Evgeny V. Zakharov, Jeremy Deeward and Vanessa Bouwer.

Abbreviations

Localities are written in the local language style, except for the names of countries for which we used the English naming. International abbreviations for the countries (Eurostat 2021) used in figures are: Dahlica (Brevantennia) triglavensis [START_REF] Rebel | Zur Kenntnis palaearktischer Talaeporiiden[END_REF] Figs 6 a-c, 7, 8c, d, 9-11 Solenobia triglavensis Rebel, 1919: 111.

Supplementary material 1