What’s Inside the Hole? A Review of European Dendrolimnetic Moth Flies (Diptera: Psychodidae: Psychodinae)

: We conducted an extensive literature review in search of records of dendrolimnetic Psychodinae, with additional ﬁeld sampling of the European species of Psychodinae associated with water-ﬁlled tree holes. After checking more than 100 publications, only 11 speciﬁc published records involving dendrolimnetic Psychodinae were found. Our results show that six genera, represented by 13 species of Psychodinae, are associated with 13 species of plant trees. As a result of our ﬁeld sampling, we report Lepiseodina latipennis (Sar à , 1953) and Telmatoscopus bartai (Ježek, 2004) comb. nov . for the ﬁrst time in Germany. Furthermore, we redescribe L. latipennis based on freshly collected material with a closer examination of the holotype. Derived from our ﬁndings, we review the genera Lepiseodina Enderlein, 1937 and Telmatoscopus Eaton, 1904, providing an identiﬁcation key for the males of both genera. In addition, we synonymyze Krivosheinoscopus Ježek, 2001 syn. nov. under Telmatoscopus, changing combination of Telmatoscopus ussuricus (Ježek, 2001) comb. nov. and Telmatoscopus bartai (Ježek, 2004), additionally, we change combination and a sononymy of Tematoscopus wagneri (Salmanna, 1982) comb. et syn. nov. under Telmatoscopus advena (Eaton, 1893). Furthermore, we describe for the ﬁrst time the female and eggs of Telmatoscopus advena . Moreover, we provide the ﬁrst published DNA barcodes (COI) for Telmatoscopus bartai , Lepiseodina latipennis (Sar à , 1953) , Lepiseodina rothschildi (Eaton, 1912), and Lepiseodina tristis (Meigen, 1830). Finally, we also discuss the taxonomy and ecology of the European dendrolimnetic species of the subfamily Psychodinae.


Introduction
Phytotelmata (singular phytotelma) derives from the Greek words "phyto", which means plant, and "telma", meaning pond, and was first proposed by Varga [1], based on his observations, to define the microhabitats product of water-filled bodies in plant surfaces [2]. Within the phytotelmata, we can find the dendrotelmata (singular dendrotelma), deriving from the Greek words "dendron", meaning tree, and "telma", which circumscribe the body waters only to water-filled tree holes. These dendrotelmata are considered an integral microhabitat inside forest ecosystems, as they provide a suitable space for development, prey, and water sources for many organisms ranging from invertebrates to vertebrates [3,4]. Species that relate to or inhabit these water-filled tree holes are known as dendrolimnetic, from the Greek words "dendron", and "limnétēs", which means relating to or inhabiting the open water of a body of fresh water.
Dendrotelmata are commonly found in old trees, dead or alive, and are a considered crucial component of ecosystems, especially with the modern forestry management where the number of mature and over-mature trees (commonly referred as old trees) has decreased [5]. The main sources of nutrients flowing in the tree holes consist of leaf litter and arthropod cadavers, while the quality and composition of these nutrients vary across species and habitats [6,7]. The fact that these microhabitats have a small size, discrete boundaries, and are naturally replicated in nature, makes them attractive for studies of community structure and functionalities [7]. Researchers have reported the usage of dendrotelmata by amphibians during development, as a water source for reptiles and small mammals, and, as bathing sites for birds and bats [4]. Nonetheless, mainly invertebrates with aquatic development have been reported developing inside these microhabitats, with some reports of other invertebrates that use them as a water source [4,8]. Among them, the most common organisms found inside are immature stages of Diptera and Coleoptera [7][8][9][10].
Moth flies (Insecta: Diptera: Psychodidae) are commonly found in water bodies, as the majority of psychodid species develop in water during larval stages, with a few exceptions that develop in soil, dung, or fungi [11][12][13]. There are six subfamilies recognized worldwide, namely Bruchomyiinae, Horaiellinae, Phlebotominae, Psychodinae, Sycoracinae, and Trichomyiinae; all of them except Horaiellinae are present in Europe [14,15]. Species in the subfamily Trichomyiinae develop inside tree holes or rotting wood and their larvae are considered xylophagous (from Greek "xylon", meaning wood, and "faguein", meaning eating) [5]. Species of Bruchomyiinae and Phlebotominae have been reported developing on the ground and leaf litter feeding on decaying organic matter [15]. The larval stages of Sycoracinae species have been found developing in aquatic mosses and leaf litter [15]. On the contrary, species of the subfamily Psychodinae are commonly reported developing in water bodies such as ponds and streams, but there are a few genera present in Europe whose larvae develop inside of tree holes, namely Clogmia, Clytocerus, Lepiseodina, Pneumia, Psychoda, and Telmatoscopus [13,[16][17][18][19][20][21][22][23][24][25]. Although the larval development and habitat for some European species are known, the moth flies associated with water-filled tree holes have been poorly studied in Europe [25].

Literature Records
Literature search was conducted by tracking references from known literature with the help of search engines (e.g., www.scholar.google.com, accessed on 5 Jaunary 2022) and scientific databases (e.g., www.jstor.org, www.scopus.com, www.webofscience.com, 5 January 2022) using the search keywords "Psychodidae, Psychodinae, dendrolimnetic, Diptera, water-filled tree holes". Literature used for the study encompasses records since the description of the species to the most recent published works until the beginning of 2022, focusing on Psychodinae and dendrolimnetic studies. More than 100 items were analyzed; however, only 11 included records of dendrolimnetic Psychodinae (as listed in Table 1).

Sampling
Specimens were collected using Malaise traps during the years 2013-2021 as part of the German Barcode of Life (GBOL) project [26] (www.bolgermany.de, accessed on 2 April 2022). Specimens were preserved in 96% ethanol and stored at −20 • C until they were dissected for DNA extraction and preparing permanent slides. All sampled specimens are stored at the Zoologisches Forschungsmuseum Alexander Koenig, Bonn, Germany (ZFMK). Further material was bred from organic matter sampled from dendrotelmata, and, occasionally collected with a Malaise trap and directly collected in Italy, during 2010-2022 (AM).
Additional examined material is deposited in the following natural history collections mentioned in the text using their acronyms: AM

Study of the Collected Material and Terminology
After lysis (see Genetics below), the specimens were cleared using NaOH 10%, dissected, and permanently mounted using Euparal (Waldeck GmbH & Co. KG, Division Chroma, Havixbecker Straße, Münster, Germany) as mounting medium, following the procedure detailed in Ibáñez-Bernal [28], with the modification that prior to the diaphanization process, we performed the dissection of the head, wings, and terminal abdominal segments to macerate them in NaOH 10% and continue with the procedure, while preserving the remaining tissue (thorax, legs, first abdominal segments) in ethanol for posterior DNA extraction. Additional material was macerated in KOH 10%, transferred in acetic acid 10%, and dehydrated in acetone 99%. Specimens were dissected in clove oil and parts mounted on microscope slides with Canada balm. For the additional material, some specimens were photographed to keep a quality image of the habitus.
We follow the general terminology proposed by Cumming et al. and Kvifte and Wagner [15,29], except for the male terminalia that we use the term "hypopods" for the posterior genitalic appendages, which have been treated in the literature as cercopodia or surstyli originating in the 9th abdominal segment, or 10th segment, or a combination of both, as proposed by Kvifte and Wagner [30]. For the female Terminalia we follow Kotrba [31]. Egg terminology follows De Almeida et al. [32].
In the examined material of each species, a code is provided for each examined specimen (e.g., ZFMK-DIP-000852020), and all the label information is found as Supplementary Material (Table S1).

Genetics
Specimens were processed at the ZFMK, where lysis and PCR were performed following the protocol and primers by [33,34]. After the PCR, samples were sent to Beijing Genomics Institute (BGI) for bidirectional sequencing. Raw data were curated manually using Geneious (v. 7.1.9). Final COI sequences were 658 bp long. All sequences will be publicly available at www.bolgermany.de. BOLD and genebank accession IDs can be found in the Supplementary Material.

Results
Based on our literature review, we report 13 species of Psychodinae associated with dendrotelmata belonging to six genera (Table 1)

Key to the Males of European Psychodinae Genera Found in Dendrotelmata
Differential diagnosis. Adults of the subfamily Psychodinae can be easily differentiated from the adults of the exclusively xylophagous subfamily Trichomyiinae, which can also be found in tree holes, by the presence of an eye bridge in Psychodinae (absent in Trichomyiinae) and wing vein R with five branches in Psychodinae, with two longitudinal veins between radial and medial forks (vein R with four branches in Trichomyiinae, with one longitudinal vein between radial and medial forks).

1.
Antenna with at least flagellomeres 2-10 nodiform, divided into a basal nod and a distal neck ( Figure 1B

2.
Cornicula (everted sac-shaped structures on the back surface of the head (presumed scent organs); also known as patagia) usually present; antennal scape more than three times longer than pedicel; flagellomere 1 with a distal brush of wavy setae ( Figure 1C

Systematic Assessment
Diagnosis: Cornicula (everted sac-shaped structures on the back surface of the head) are usually present; eye bridge with 3-6 facet rows, separated by 1-3 facet diameters; antennal scape elongate, three-six times longer than wide; flagellomere I and II fused to form a compound flagellomere, with a distal brush of wavy setae [14].
Examined material. None. Distribution. France [43]. Remarks. Vaillant [37] mentions under the description of C. xylophilus that the larvae of Clytocerus pulvereus Vaillant, 1983 can be found in the same substrate as C. xylophilus; however, it is referring to the "black humus" found near a river (same substrate mentioned by Krek [49]), and later Vaillant [37] clarifies that only C. xylophilus is known as a dendrolimnetic species.
Genus Diagnosis. Frons and clypeus separated and not protruding over eye margin; scape short, less than 2 times length of pedicel; flagellomeres symmetrically nodiform in dorsal view, with a pair of digitiform sinuous ascoids; flagellomere 14 with elongated apiculus; wing vein R 2+3 not connected to R 4 ; apex of R 5 ending on, or below wing apex; gonocoxites short, with dorsal projection undeveloped; ejaculatory apodeme narrow in dorsal view; aedeagal complex asymmetric, with a parameral structure more or less sclerotized, connecting aedeagus to the gonocoxal apodemes; hypopods with indistinctly fringed tenacula; epandrium with a single foramen.
Differential diagnosis. This species is closely related to Lepiseodina tristis, but it can be distinguished by the combination of the following characters: hypandrium broad along its entire length with convexity in the basal margin in L. latipennis (hypandrium narrow with a medial projection in the basal margin in L. tristis); gonocoxal alveoli distributed in the entire gonocoxal surface in L. latipennis (gonocoxal alveoli restricted to the apical margin in two or three irregular rows in L. tristis); gonostyli without alveoli on the base in L. latipennis (gonostyli with alveoli in the base in L. tristis); epandrium not divided in the apical margin, with a kidney-shaped foramen in L. latipennis (epandrium divided in the apical margin with a rounded foramen in L. tristis); aedeagus curved in L. latipennis (aedeagus straight in L. tristis); parameres not strongly sclerotized and almost not visible in L. latipennis (parameres strongly sclerotized in L. tristis).
Lepiseodina latipennis can be differentiated from Lepiseodina rothschildi by the gonostyli twice the length of gonocoxites in L. latipennis (gonostyli less than half of the length of gonocoxite in L. rothschildi), apical margin of hypandrium convex in L. latipennis (apical margin of hypandrium concave in L. rothschildi), and parameres not strongly sclerotized and almost not visible in L. latipennis (paramere strongly sclerotized in L. rothschildi).
Redescription Male. Head: eye bridge with rows of four facets (rarely 3 facets); frontal patch undivided, with an irregular row of alveoli extending towards the interocular suture, the whole frontal patch together with the irregular row resemble the shape of a handbell; interocular suture as an inverted "v". No alveoli of supra ocular setae are present. Labella bulbous, longer than wide, with 10-12 small setae. Antenna with scape cylindrical, 1.34 times longer than its width, 1.43 (1.2-1.4) times the length of pedicel; pedicel spherical; 14 nodiform flagellomeres with basal bulb and distal neck, apical flagellomere with long apiculus, apiculus almost half the length of the flagellomere ( Figure 4A); antennal ascoids slightly flattened, digitiform, sinous (S-shaped), total length about 1.88 the length of flagellomeres; flagellomeres with a sensilla (as shown in Figure 6A). Palpal segment proportions 1.0:2.63:2.37:3.03.
Thorax with no particular characters.
Wing: Length about 2.5 (2-2.5) times its width, variable in shape, with anterior margin more or less curved; membrane bare except on veins; hyaline with a slight infuscation on costal cell; Sc straight, ending at level of the origin of R 2+3 ; Origin of R 2+3 not joining R 4 , a little distal to the origin of M 1+2 ; origin of M 1+2 broad and rounded; forks of R 2+3 and M 1+2 almost at the same level, the fork of R 2+3 being slightly distal to M 1+2 ; R 5 ending at wing apex; CuA ending in wing margin at the level of R 2+3 fork.
Abdomen. Without any particular characters. Genitalia. Hypandrium plate-like, broad, basal margin with medial projection widely rounded, distal margin almost straight; gonocoxites longer than wide, covered in alveoli on almost all the surface; gonostyli about 1.5 times the length of gonocoxites, tapering towards apex, almost straight; gonocoxal apodemes poorly distinguishable in dorsal view, in ventral view plate-like, strongly sinuous, fused and narrow at livel of the midline; parameres forming a single slight sclerotized structure, pyriform, resembling the upper half of a bowling pin connecting like a bridge aedeagus to gonocoxal apodemes (as in Figures 4D, 5C, 6B, 9C and 10A); epandrium subrectangular, about twice wider than long with a single kidney-shaped foramen; hypopods slightly out curved, apex rounded, with 9-13 tenacula (as in Figures 4B, 5C and 9C); epiproct short, triangular and covered in pilosity; hypoproct broad and tongue-shaped, covered in pilosity, extending towards mid of hypopods. Aedeagus with ejaculatory apodeme digitiform, narrow 7.5 times longer than its width, about 1.5 times the length of gonocoxites; distiphallus about the same length of ejaculatory apodeme, incurved, extending towards the apex of gonostyli, both ejaculatory apodeme and distiphallus form a single complex, jointed with parameral structure and encircles by a membranous parameral sheath.
Female. Unknown. Remarks. The holotype (slide mounted) is quite dark, making the observation of structures difficult. The head is dissected, and quite difficult to see clearly, one antenna is dissected and the other antenna is missing, one complete palpus is dissected, the other palpus is missing. The hypopods are dissected and placed apart from each other, one gonocoxites and gonostylus are dissected and placed apart from the genitalia, the remaining parts of the genitalia (aedeagal complex, one gonocoxites and gonostylus) are found together. One wing is dissected and separated from the thorax. On the original description Sarà (1953) mentions the shape of the hypoproct being trilobed (Figure 2), after examination of the holotype in the preparation the hypoproct looks indeed trilobed; however, we consider this a malformation on the slide itself, and not the natural shape of the hypoproct, all other examined material present a broad tongue-shaped hypoproct. On the original description Sarà abstained to illustrate or describe the aedeagal complex; however, after examination we are sure all our specimens belong to the same species.
Biology. Some specimens (No. 0190-0193) emerged from organic matter sampled from an Acer sp. dendrotelma (Figure 14), in a mixed submontan forest, additionally, the specimen used for the SEM pictures was collected in a Malaise trap next to dendrotelmata of an oak (Quercus sp.) tree, suggesting that also L. latipennis is a dendrolimnetic species. In Germany altitude ranges from 250-280 m a.s.l. (meters above sea level), while in Italy it has a range from 40-800 m a.s.l.
Examined material.    Diagnosis. Male hypandrium narrow, with a small abrupt median projection; gonocoxal alveoli restricted to two irregular lines on the apical surface; gonocoxites with a patch of alveoli at base; two parameres strongly sclerotized. Distribution. Algeria, Austria, Belgium, Croatia, Czech Republic, France (incl. Corsica), Germany, Ireland, Italy, Lithuania, Slovakia, and United Kingdom [25,36,43,50,53,59,60]. In Italy, the species is known only from two old and uncertain records for the northern and peninsular region [53,61]. The specimens here reported confirm the occurrence of this species in the Italian peninsula.
Key to the European adult males of Lepiseodina
Examined material. Diagnosis. Eye bridge with 7 facet rows; frons not extending beyond first palpal segment; all pal segments of similar width; fore tibia straight and not engrossed; wing vein fork R 2+3 without a backward projection; hypopods with 7 or fewer tenacula; ejaculatory apodeme straight, not wider at base; aedeagal complex shorter than gonocoxites, not tapering towards apex, distal part of aedeagus smooth without wrinkles. Diagnosis. Species with short vertex; labellum flattened and carrying digitiform setae; eyebridge without interocular suture; antenna with 12-14 flagellomeres, those beyond 11th always reduced in size and showing different types of fusion; flagellomeres nodiform with a neck (except apical flagellomeres); ascoids usually with three branches, two anterior and one posterior branch, Y-shaped or with four branches, shaped like a plus sign (+); aedeagus often asymmetrical; hypopods with a single apical tenaculum.
Examined material.  [83]). Frons and clypeus separated and not protruding over eye margin; flagellomeres asymmetrically nodiform, with paired leaf-shaped to digitiform ascoids; flagellomere 14 with elongated apiculus; wing veins R 2+3 not connected to R4; apex of R5 ending at wing apex; ejaculatory apodeme narrow in dorsal view and distally ending in two short branches with membranous connection to aedeagal complex; aedeagal complex symmetrical; aedeagal complex encapsulated in a parameral sheath; hypopods with indistinctly fringed tenacula; epandrium with a single foramen.
In the table, seven morphological characters were provided to differentiate Krivosheinoscopus from Sciria (=Telmatoscopus), they are numbered and discussed below.

2.
The ascoids are very long, thin and coiled in Krivosheinoscopus (large, flat, leaf or hood-shaped in Sciria). This character is contradicted by Telmatoscopus laurencei [38], which has digitiform ascoids and coiled, and these character states are polymorphic in Vaillantodes Wagner, 2001 and Panimerus Eaton, 1913. Intermediary forms also occur in some genera of Telmatoscopoids. Due to widespread polymorphism of this character we do not consider this a reliable genus-level character unless supported by other, independent lines of evidence. 3.
The first palpal segment very short and keg-shaped in Krivosheinoscopus (long and cylindrical in Sciria). This character is variable inside the genus Telmatoscopus (e.g., long in Telmatosocpus advena and shorter in Telmatoscopus thuringicus Beran, Doczkal, Pfister & Wagner, 2010), therefore it is here considered as intrageneric variability, and thus not a diagnostic character.

4.
The position of the radial and medial forks of wing venation, at the same distance in Krivosheinoscopus (medial fork distad to radial fork in Sciria). This is another character presenting as variable within the genus Telmatoscopus (e.g., medial fork distal in T. advena, medial fork basal in T. thuringicus, forks at the same level in Telmatoscopus bartai).

5.
Two pairs of protuberances in the aedeagal complex in Krivosheinoscopus (one pair in Sciria). According to the homologization of [83], the absence of protruding parameres in Telmatoscopus are not due to absence of parameres, they are present as transverse sclerites within the aedeagal-parameral complex. The difference between T. advena, T. bartai and T. ussuricus in the parameres is a question of degree of development rather than a clear-cut presence/absence question, and intermediate cases occur in Nearctic species (e.g., T. patibulus Quate).

6.
The hypoproct has a terminal projection, long and narrow in Krivosheinoscopus (without a terminal projection, triangular in Sciria). This character is contradicted by Telmatoscopus bartai comb. nov. where the hypoproct is triangular, and not thin and elongated.
With three shared characters between Krivosheinoscopus and Telmatoscopus (=Sciria sensu Ježek) including the wing vein R 5 ending at wing apex, vein Cu ending distal to M 1+2 and, the ejaculatory apodeme laterally compressed (=basal apodeme of the aedeagal complex in Ježek) [84].
2. First palpal segment short and keg-shaped in Krivosheinoscopus (long and cylindrical in Seoda). This character is variable inside Telmatoscopus, and is thus not a diagnostic character.
3. Vein Cu ending distal to M 1+2 in Krivosheinoscopus (samel level or distad to M 1+2 in Seoda) yet again, a variable character; 4. Wing vein R 5 ending at wing apex in Krivosheinoscopus (ending below wing apex in Seoda), this character is diagnostic between Seoda and Telmatoscopus as pointed out in [83].
Diagnostic morphological characters to separate Telmatoscopus from Seoda presented in [83] include single pair of digitiform to filiform ascoids (an additional ring of small setiform ascoids + the main pair of digitiform ascoids in Seoda); frons clearly separated from clypeus not protruding over the mesal margin of eyes in Telmatoscopus (frons fused with clypeaus and protruding over the mesal margin of eyes in Seoda); parameral sclerites not fused in Telmatoscopus (parameral sclerites fused in Seoda).
Our currently reformulated conscription of Telmatoscopus includes 10 species globally, namely T. advena (Eaton, 1893), Palaearctic; T. bartai (Ježek, 2004)  Notes. Previously, Kvifte, [83] included Panimerus wagneri Salamanna, 1982 as Seoda, however the inclusion of T. wagneri (=T. advena) comb. nov. et syn. nov. inside of Telmatoscopus is based on the male genital morphology presented in the original description, therefore the new combination from Seoda to Telmatoscopus. Additionally, the characters separating T. wagneri (=T. advena) syn. nov. and, T. advena are quite inconspicuous, however, we did not examine the holotype of T. wagneri (=T. advena) syn. nov., nonetheless, based on the original description and figures, we did not find any morphological characters that support them as separates species, and we treat T. wagneri as a new synonym of T. advena.
Additional characters and closer examination of the species T. pappi, T. frondeus, and, T. tanegashimensis is desirable as they share some characters with the genus Lepiseodina, however, these species are outside the geographical scope of this work, and therefore are included as Telmatoscopus following [83] until a broader revision of the genus is available.
Female description (Figures 15 and 16) based on the description of the male in [83] the female is similar to the male except: Head slightly longer than wide (0.5 mm wide, 0.6 mm length). Eye bridge with four facet rows, separated by 1.5 facet diameter; only first palpal segment present in examined material; apical antennal flagellomeres absent in examined material, ascoids s-shaped, coiled, thin (not broad leaf-shaped as in male). Wing length, wing width, length x times its width.   Wing 2.7 times longer than wide (1 mm wide, 2.7 mm long), Sc long, ending at junction of R 4 + R 2+3 . Infuscated in area between C and R1.  Diagnosis. Male. Ascoids are broad, leaf-shaped, tapering towards the apex and coiled; phallomeres out curved, less than half the length of ejaculatory apodeme; hypopods with a cluster of approximately 30 tenacula at apex, and more tenacula scattered along the entire surface of hypopods.
Examined material. None. Distribution. Only known from Germany [5]. Key to the European adult males of Telmatoscopus

Discussion
All of the above-mentioned species (also in Table 1) are recorded to be associated with dendrotelmata to a certain degree, whether they are specialized to complete their life cycle in this ecosystem or they are only opportunistic is a different matter. As mentioned by Oboňa and Ježek [25] the extreme variation of environmental conditions such as pH, temperature, frequent water loss with rapid flooding and, oxygen deficit can cause the death of non-specialized species, especially in larval stages that are using dendrotelmata as an irregular breeding/developing site, while specialized species can endure these harsh 822 environmental variations thriving through all their life cycle.
The occurrence of Clogmia albipunctata, C. xylophilus, Pneumia canescens, P. trivialis, Psychoda alternata, P. cinerea, and P. minuta in dendrotelmata is rather incidental or highly understudied [23,25]. In other words, the presence of these species could be an extension of their regular development sites (e.g., small water bodies, streams, etc.) and adults happen to find water-filled tree holes that are a potential development site for their offspring, thus, they could develop in Dendrotelmata and survive, but their long-term usage of this site is not well documented. Further studies in different habitat could provide key information to better understand the ecological relationship between water-filled tree holes and the species of Psychodinae that develop in them. To the date, only a handful of studies specifically targeted moth flies in dendrotelmata.
Clogmia albipunctata is broadly distributed both in Europe and worldwide and is the most synanthropic species inside the Psychodidae fauna, it can be commonly found inside buildings, and sewage treatment plants in almost every city. The high synanthropy is shared with some species of the genus Psychoda, which are also often found in cities, including P. alternata and P. cinerea, this species can certainly adapt and develop in a wide range of habitats, including dendrotelmata, therefore, these species could be classified as generalists when it comes to sites for larval development and not bound to develop in water-filled tree holes. Some further Psychoda species may be found in dendrotelmata, but the development of most species is still unknown.
Lepiseodina tristis, L. rothschildi, Telmatoscopus advena and T. laurencei are well-documented species that develop inside dendrotelmata (Table 1) with multiple records of both adults and larvae. On the contrary, Telmatoscopus thuringicus and T. bartai comb. nov., are only assumed to be dendrolimnetic based on observations of the closely related species T. advena [5]. In the case of the herein described species Lepiseodina latipennis-some specimens collected in Italy were reared from decaying organic matter collected in a dendrotelmata from a maple tree (Acer sp.), further specimens collected in Germany came from a Malaise Trap placed next to a water-filled tree whole in an Oak tree, thus proving that this species develop inside dendrotelmata as congeneric species.

Conclusions
In Europe, only 13 species of Psychodinae are known to develop inside water-filled tree holes, after our extensive record search we report that the Psychodinae species develop in 13 different tree species, including two new tree species in which no previous record of larval development was reported. We redescribed Lepiseodina latipennis through holotype and new material examination, and we report it for the first time in Germany. We also report Telmatoscopus bartai comb. nov. for the first time in Germany, and we provide a generic discussion of Telmatoscopus. Water-filled tree holes (dendrotelmata) are usually associated with old trees which are commonly endangered through the forest management strategies applied in European countries, and they are becoming rare to find inside forests. To summarize, there is a gap in the knowledge of the ecological interactions inside the Psychodinae and their environment, further studies can potentially provide new information, new records and new interactions that would be beneficial to better understand the ecosystems. Furthermore, old tree individuals remain a key component in the forests, as they harbor high biodiversity that is still understudied and they should remain untouched until the natural decomposition takes place.