Exceptional diversity of Tischeriidae (Lepidoptera) from a single tropical forest site in Belize, Central America

We report on fourteen species and four genera of Tischeriidae recorded from Las Cuevas, a single tropical forest locality in Belize, Central America. This is the highest number of species of Tischeriidae recorded from a single locality worldwide, exceeding the species and generic diversity of the entire Tischeriidae fauna of Europe and accounting for about 9% of the documented global fauna for this family. We describe and name six new species: Astrotischeria papilloma Diškus & Stonis sp. nov., mining on Lasianthaea fruticosa (L.) K.M.Becker (Asteraceae); A. scutifera Diškus & Stonis sp. nov., mining on Sida glabra Mill. (Malvaceae); A. basilobata Remeikis & Stonis sp. nov., mining on Lasianthaea fruticosa; Paratischeria robinsoni Diškus & Stonis sp. nov., mining on Otopappus verbesinoides Benth. (Asteraceae); P. tubifex Diškus & Stonis sp. nov., mining on Lasianthaea fruticosa; and P. belizensis Remeikis & Stonis sp. nov. (host plant unknown). Additionally, we review eight previously described species from the same period of collecting at Las Cuevas in 1997–1998: A. selvica Diškus, Carvalho-Filho & Stonis, 2018, mining on Sphagneticola trilobata (L.) Pruski and Synedrella nodifl ora (L.) Gaertn. (Asteraceae); A. casila Diškus & Stonis, 2018, mining on Montanoa atriplicifolia (Pers.) Sch.Bip. (Asteraceae); A. furcata Diškus & Stonis, 2018 (host plant unknown); Paratischeria neotropicana (Diškus & Stonis, 2015), mining on Sida L. (Malvaceae), including S. rhombifolia L.; Dishkeya gouaniae (Stonis & Diškus, 2007), mining on Gouania polygama (Jacq.) Urb. (Rhamnaceae); Coptotriche pulverea (Walsingham, 1897), mining on Terminalia amazonia (J.F.Gmel.) Exell (Combretaceae); C. forsteroniae Stonis & Diškus, 2008, mining on Forsteronia myriantha Donn Sm. (Apocynaceae); and C. singularis Stonis & Diškus, 2008 (host plant unknown). All taxa, except for C. singularis, are illustrated with photographs of the adults and their genitalia. We also briefl y discuss the discovery of some novel characters for Astrotischeria Puplesis & Diškus, 2003 and Paratischeria Diškus & Stonis, 2017, Tischeriidae, and provide the fi rst photographic documentation of Coptotriche pulverea and C. forsteroniae. European Journal of Taxonomy 723: 33–76 (2020) 34


Introduction
Larvae of Tischeriidae (trumpet moths) are specialized miners of the green tissues of plants, producing mostly irregular, blotch-like, sometimes slender and sinuous or other leaf mines. The morphological and ecological characteristics of these primitive, monotrysian microlepidopterans are summarised by Braun (1972), Puplesis & Diškus (2003), Xu et al. (2017Xu et al. ( , 2018 and Stonis et al. (2017Stonis et al. ( , 2018aStonis et al. ( , 2019aStonis et al. ( , 2019b. Species of Tischeriidae are represented in a wide variety of terrestrial ecosystems from the tropics to the temperate regions but, as a rule, the family is more abundant in subtropical and tropical regions, except for cooler habitats of the tropics at elevations above 3500-4000 m (Stonis et al. 2018a).
The tiny size of the adults, the concealed mining life-style of the larvae within leaves and the lack of qualified specialists on these insects go some way towards explaining why trumpet moths are still so little studied in many regions, including the Neotropics. For example, until recently not a single species of Tischeriidae was reported from Colombia , which is regarded as one of the world's ʻmegadiverseʼ countries. Only during the last year have the first two species of Tischeriidae been described from this country (Stonis et al. 2019b). The study of the Tischeriidae fauna in the Neotropics began in the late nineteenth to early twentieth century with the descriptions of the first species (Walsingham 1897(Walsingham , 1914Meyrick 1915). However, only during the last two decades has the study of Neotropical Tischeriidae become more purposeful and active through collecting in the field and the compilation of species inventories for Central and South America (Stonis et al. 2019a). It has been suggested that Tischeriidae might make up a significant portion of the Neotropical fauna of leaf-mining insects (Stonis et al. 2019a).
This paper reports on samples of material of leaf mines collected at Las Cuevas, Belize ( Fig. 1) by Owen T. Lewis in 1997and 1998(Lewis et al. 2002. Mining larvae of tischeriids were very sparse during the dry season but more abundant during more humid periods of the year. Additional material was obtained at Las Cuevas in April 1998 by light trapping by Jonas R. Stonis (formerly R. Puplesis) and Simon R. Hill (Puplesis & Hill 1998;Puplesis & Robinson 2000). . The Chiquibul Forest was subject to selective logging for timber during much of the 20th Century. The majority of the vegetation around Las Cuevas is recovering from selective logging during the past ca 40-50 years, although a few pockets of relatively undisturbed or primary forest remain. The Chiquibul receives between 1500 and 2000 mm of rain per year. Rainfall is seasonal, with a dry season typically from February to June (approximately) and sometimes a shorter dry period of two to three weeks in August (Puplesis & Robinson 2000). Hilltops, hillsides, and valley bottoms have markedly different forests both in terms of structure and species composition. At our main collecting sites (on hillsides and in valley bottoms) the forest has a greater proportion of evergreen species, e.g., Manilkara zapota (L.) P.Royen, Calophyllum brasiliense Cambess. and Aspidosperma megalocarpon Müll.Arg., although deciduous species such as Swietenia macrophylla King and Terminalia amazonia (J.F.Gmel.) Exell were still a significant component (Puplesis & Robinson 2000). Las Cuevas is an important centre for biological or ecological research, established jointly by the Forest Department of Belize and the Natural History Museum, London in 1994 and now co-managed by the Friends for Conservation and Development and the Government of Belize.
The collected material from Las Cuevas has yielded fourteen species of Tischeriidae. Externally all species collected at Las Cuevas look very similar . External characters are not sufficient for species identification; therefore, we do not use them in the diagnoses of the species.
With one exception, Coptotriche pulverea (Walsingham, 1897), all species of Tischeriidae collected at Las Cuevas appear to be new to science. Dishkeya gouaniae (Stonis & Diškus, 2007) was described within a review of the genus Tischeria Zeller, 1839 (Stonis & Diškus 2007), and Coptotriche forsteroniae C. singularis Stonis & in the first review of Coptotriche Walsingham, 1890 in the Americas . Paratischeria neotropicana (Diškus & Stonis, 2015) was described in a separate article indicating the broadest distribution of trumpet moths (Diškus & Stonis 2015). Three other species, Astrotischeria selvica Diškus, Carvalho-Filho & Stonis, 2018, A. casila Diškus & Stonis, 2018and A. furcata Diškus & Stonis, 2018, were documented along with a recent review of three-lobed Astrotischeria Puplesis & Diškus, 2003(Stonis et al. 2018a International concern over the biodiversity crisis has underscored the urgency to inventory global biotas and to provide taxonomic reviews of diverse groups of organisms, particularly in poorly-studied yet species-rich tropical areas (Puplesis & Robinson 2000). This account of taxa recorded from a single locality is intended to illustrate the high local diversity of Tischeriidae that can occur within the tropical forests of Central America. It will provide a foundation upon which we and others can build a more detailed account of the diversity of Central American and Neotropical Tischeriidae.

Material and methods
Leaf mines containing living larvae were sampled from tropical forest vegetation from September 1997 to September 1998. Sampling occurred in all months except December, January and May. The sampling regime for larval leaf miners was described by Lewis et al. (2002) and concentrated on vegetation growing along the margins of four-wheel drive vehicle tracks within 4 km of the field station. Mines were reared individually in airtight plastic containers, lined with absorbent paper. Alternative techniques for rearing adults from mining larvae were described in detail by Stonis et al. (2018a). Adult moths emerged from the mines typically within 20 days (maximum 35 days) of collection. Overall, 2053 living mines of Tischeriidae were collected, of which 654 died during rearing, 901 produced adult moths and 498 produced Hymenoptera parasitoids (35.6% parasitism across all Tischeriidae). Interactions between parasitoid species and leaf miner hosts were reported and analysed by Lewis et al. (2002), and a manipulative experiment on the host-parasitoid food web (also comprising non-tischeriid leaf miners) was reported in Morris et al. (2004). The text below associates, for the first time, the morphospecies names used in those ecological studies with species identifications and descriptions.
Adult moths were collected by attracting them to mercury-vapour light from a lamp suspended slightly above eye-level and 5-10 cm in front of a white screen (Puplesis & Robinson 2000), rather closer than is  usual in the standard method for light collecting (described by Robinson et al. 1994) in which the lamp is about 0.5 m from the illuminated surface. A Honda EX 350 generator was used as a power source.
Moths attracted to the screen were collected in small glass tubes.
Preparation of temporary and permanent micro-mounts of genital structures was undertaken following the standard method described by Stonis et al. (2014). After maceration of the abdomen in 10% KOH and subsequent cleaning, abdominal pelts and female genitalia were stained with Chlorazol Black (Direct Black 38/Azo Black), but male genitalia were left unstained. Both male and female genitalia were mounted in Euparal. The genitalia were fixed with the ventral part facing up; occasionally, some sclerites were disassembled from the capsule; the phallus was usually removed from the capsule; the pelt or loose sclerites of the genital armature (e.g., removed phallus) were fixed under a separate cover slip, always on the same slide. Sometimes we rolled and photographed genitalia in Euparol before finally closing the mount with a cover slip; this generated images of the genitalia in lateral view, especially useful with complex structures, viewed differently from the ventral side. In our study, the genital micro-mounts were examined using a Leica DM2500 microscope and a Leica DFC420 digital camera connected to the microscope and the computer; each photograph of a genitalia slide was supplied with the slide number and species name.
The descriptive terminology of morphological structures follows Puplesis & Diškus (2003), except for the term ʻaedeagusʼ, which is referred to here as ʻphallusʼ, and the term ʻciliaʼ, which is referred to here as ʻfringeʼ.

Repositories
Descriptions of the new species treated in this paper are based on material deposited in the collection of the Natural History Museum, London (NHMUK). Additionally, some specimens of Astrotischeria papilloma sp. nov. were available for our study from Honduras, and they are deposited at the collection of the Biosystematics Research Group (former Lithuanian University of Educational Sciences, formerly abbreviated as LEU or VPU), Vilnius, Lithuania (with subsequent transfer to the Zoological Institute of the Russian Academy of Sciences, ZIN, because of the LEU closure).

Diagnosis
External characters are not sufficient for the identification of this species. In the male genitalia, the unique, highly specific, finger-like dorsal lobe of the valva (Fig. 42), the bifurcated dorsal lobe of the uncus (Fig. 35) and the angular apex of the valva (Fig. 40) distinguish Astrotischeria papilloma sp. nov. from other congeneric species. In the female genitalia, the slender band of long chetae (Fig. 49) is hypothesized to be unique to this species. This character may not remain valid for species differentiation because females of many other species of Astrotischeria remain to be discovered. The host plant, Lasianthaea fruticosa (L.) K.M.Becker (Asteraceae Bercht. & J.Presl), coincides with the host plant of another new species, Paratischeria tubifex sp. nov., described below.

Etymology
The species name is derived from the Latin ʻpapillaʼ with the prefix ʻomaʼ, i.e., the widely used medical term papilloma (an outward finger-like frond), in reference to the unique, finger-like dorsal lobe of the valva in the male genitalia.

Male
ExtErnal charactErs . Forewing length 2.3-2.5 mm; wingspan 5.2-5.5 mm (n = 6). Head: palpus, pecten and frons cream; frontal tuft comprised of lamellar scales, from yellowish ochre or ochre cream to pale grey, often with pale grey, cream-tipped or cream, brown-tipped scales medially; collar comprised of lamellar, yellowish cream scales; antenna exceeding ½ of forewing; flagellum cream, brown-annulated to entirely grey-brown. Thorax glossy, pale grey-brown medially, yellowish cream to ochre laterally; tegula pale grey-brown. Forewing variable, usually silver glossy to cream, irregularly speckled with grey-brown and some ochre scales, sometimes with a wide band of bright ochre scales along fold; fringe ochre cream to golden ochre; fringe line often irregular, indistinctive, comprised of dark brown scales; forewing underside dark grey, without spots or androconia. Hindwing and fringe grey on upper side and underside, without androconia. Legs cream, glossy to grey or dark grey; forelegs and midlegs with blackish brown scales on upper side. Abdomen grey-brown with some purple iridescence on upper side and underside, but distally ochre cream on underside; genital plates pale grey to greyish cream; anal tufts large, grey.

Biology
Host plant: Lasianthaea fruticosa (L.) K.M.Becker (Asteraceae). Larvae were recorded mining leaves throughout the year (i.e., during all months of fieldwork), with 810 records in total. The leaf mine is a pale, transluscent blotch on the upper leaf surface, located along the leaf margin. It causes the leaf margin to curl and roll upwards and inwards in a characteristic manner reminiscent of the mines of the European species Coptotriche gauacella (Duponchel, 1843). This species was listed as 'Tischeria sp. 7' by Lewis et al. (2002), a morphospecies grouping which incorrectly grouped this species with A. basilobata sp. nov., which shares the host plant. Retrospectively, records of these two species can now be distinguished on the basis of characteristics of the leaf mine.

Flight period
Based on a single specimen collected at light, adults fly in April; based on rearing data, adults are likely to occur year-round. Adults eclose for up to 33 days after the collection of mines.

Distribution
So far this species is known from a single locality in Belize, Chiquibul Forest Reserve, Las Cuevas, at an elevation of about 550 m. In the female genitalia, the strongly reduced ovipositor lobes make this species similar to the other Malvaceae-feeding species of Astrotischeria (which make up a minority of the genus); however, unlike A. omissa (Braun, 1927) or A. heliopsisella (Chambers, 1875), females of A. scutifera sp. nov. do not possess thickened tooth-like projections or curved hooks distally on the ovipositor.

Etymology
The species name is derived from the Latin ʻscutumʼ (a ʻshieldʼ) and ʻferoʼ (ʻto bearʼ, ʻto carryʼ) in reference to the valva possessing a shield-like lobe in the male genitalia.

Male
ExtErnal charactErs (Figs 9-10). Forewing length 2.5-3.0 mm; wingspan 5.4-6.5 mm (n = 5). Head: palpus cream, frons ochre cream; frontal tuft comprised of lamellar, golden cream scales; collar pale brown, comprised of lamellar, cream-tipped scales; antenna approximately ½ of forewing; flagellum pale grey. Thorax pale brownish grey, laterally sometimes ochre; tegula pale brownish grey. Forewing variable, densely speckled with dark grey scales and irregular, pale ochre to bright ochre spots or a band along fold; sometimes, instead a band or spots, ochre scales form only a line along fold; fringe grey; fringe line often irregular, indistinctive, comprised of a few dark brown scales; forewing underside with some purple iridescence, entirely covered with dark grey scales, without spots or androconia. Hindwing and fringe grey-brown on upper side and underside, without androconia. Legs glossy, with purple iridescence, pale grey or grey to yellowish cream. Abdomen dark brown on upper side and underside, with purple and blue iridescence on upper side; genital plates greyish cream to pale grey; anal tufts large, grey to yellowish grey.

Female
ExtErnal charactErs (Fig. 11). Forewing length 3.0-3.1 mm; wingspan 6.6-6.8 mm (n = 3). Similar to male but ochre scales of forewing prevail and form large ochre spots. On underside, abdomen covered with ochre cream medially and distally, without anal tufts. Otherwise, similar to male.

Biology
Host plant: Sida glabra Mill. (Malvaceae Juss.). Larvae mine leaves in September to November. The leaf mine is a white, opaque blotch; as the larva matures it causes the leaf to distort or curl. This species was listed as 'Tischeria sp. 6' by Lewis et al. (2002). Of 353 mines successfully reared, 189 produced parasitoids (53.5% parasitism).

Flight period
Based on reared specimens, adults occur throughout the year.

Distribution
So far this species is known from a single locality in Belize, Chiquibul Forest Reserve, Las Cuevas, at an elevation of about 550 m.

Etymology
The species name is derived from the Latin ʻbasisʼ (a ʻbaseʼ) and ʻlobatusʼ (ʻlobedʼ) in reference to the basally thickened, lobe-like structure of the anellus in the male genitalia.  (Figs 12-14). Forewing length 2.6-3.1 mm; wingspan 5.5-6.6 mm (n = 7). Head: palpus yellowish cream, frons yellow ochre, smoothly scaled; frontal tuft and collar comprised of golden yellow or yellow-ochre to pale grey, slender, lamellar scales; antenna exceeding ½ of forewing; flagellum yellow cream, annulated with pale brown scales. Thorax ochre-yellow, medially covered with pale brown, cream-tipped scales; tegula covered with a mixture of ochre-yellow and pale brown  scales or entirely covered with pale brown, cream-tipped scales. Forewing ochre-yellow with irregular markings of dark brown or black-brown scales; fringe pale brown, ochre-yellow apically; fringe line indistinctive or absent; forewing underside grey-brown to pale grey-brown, without spots or androconia. Hindwing pale brown; fringe brownish cream to pale brown with reddish tint, without androconia. Legs ochre cream, densely speckled or entirely covered with pale grey-brown scales on upper side. Abdomen dark brown to brown with some purple and green iridescence on upper side, yellowish ochre, annulated with dark brown scales on underside; genital plates yellowish cream; dorsal anal tuft large, comprised of ochre cream piliform scales.

Female
Unknown.

Biology
Host plant: Lasianthaea fruticosa (L.) K.M.Becker (Asteraceae). Larvae mine leaves year-round. The leaf mine is a translucent blotch, not usually at the leaf margin, but the biology of this species is otherwise unknown. This species was listed as 'Tischeria sp. 7' by Lewis et al. (2002), a morphospecies name which incorrectly grouped this species with A. papilloma sp. nov., which shares the larval host plant. Retrospectively, records of these two species can now be distinguished based on the appearance of the leaf mines.

Flight period
Based on reared specimens, adults occur throughout the year.

Distribution
So far this species is known from a single locality in Belize, Chiquibul Forest Reserve, Las Cuevas, at an elevation of about 550 m.

Flight period
Adults are known from February, April, June-July, and October-November; based on reared specimens from Las Cuevas, adults are likely to occur year-round in Belize.

Distribution
Currently known from Central America (Belize and Guatemala), the Caribbean (U.S. Virgin Islands) and equatorial Brazil (State of Pará) (Stonis et al. 2018a

Flight period
Based on reared specimens from Las Cuevas, adults occur during April and June-October.

Distribution
Currently known only from the Cayo District of Belize (Las Cuevas and San Ignacio).

Biology
Host plant: unknown.

Flight period
Based on a single specimen collected at light, adults occur in April.

Diagnosis
External characters are not sufficient for the identification of this species. In the male genitalia, the unique four-lobed uncus (Fig. 97) and the presence of pectens on the valva (Fig. 96) distinguish Paratischeria robinsoni sp. nov. from other congeneric species. In the female genitalia, the combination of a specific shape of prela (Fig. 101), large ovipositor lobes, densely covered with peg setae (Fig. 103), and a dentate thickening on the tergum (Figs 104-105) is hypothesized to be unique to this species. The Asteraceae host plant, Otopappus verbesinoides Benth., make this species unique in the Neotropical fauna.

Flight period
Based on rearing data, adults are likely to occur throughout much of the year.

Distribution
So far this species is known from a single locality in Belize, Chiquibul Forest Reserve, Las Cuevas, at an elevation of about 550 m.

Diagnosis
External characters and characters of the female genitalia are of little or no use in the identification of this species. However, in the male genitalia, the very unusual, wide valva (Fig. 108), apically covered with modified, spine-like setae (Fig. 110), the unusually large tube-like socii (Figs 113-114), and the large, deeply divided uncus (Fig. 114) instantly distinguish Paratischeria tubifex sp. nov. from other Tischeriidae. The host plant, Lasianthaea fruticosa (Asteraceae), coicides with the host plant of another new species, Astrotischeria papilloma sp. nov., described above.

Flight period
Based on a single specimen collected at light, adults fly in April; based on rearing data, adults occur in February-September.

Distribution
So far this species is known from a single locality in Belize, Chiquibul Forest Reserve, Las Cuevas, at an elevation of about 550 m.

Diagnosis
External characters are not sufficient for the identification of this species. In the male genitalia, the phallus being fused with the anellus, with two apical, pointed lobes (Figs 124, 127), and the elaborated uncus (Fig. 125) distinguish Paratischeria belizensis sp. nov. from other congeneric species.

Etymology
This species is named after Belize where it occurs.

Female
Unknown.

Biology
Host plant and biology unknown.

Flight period
Based on a single specimen collected at light, adults fly in April.

Diagnosis
External characters are not sufficient for the identification of this species. In the male genitalia, the combination of a slender valva, a long, simple uncus, a unique, distally bilobed anellus (Fig. 128) and an apically deeply divided phallus (Fig. 129) distinguish Paratischeria neotropicana from other congeneric species. The host plant genus, Sida L. (Malvaceae), coincides with the host plant of only two other species, Astrotischeria scutifera sp. nov., described above, and the Nearctic A. omissa (Braun, 1927); however, A. scutifera sp. nov. and A. omissa belong to the genus Astrotischeria and are chracterized by a very different genitalic morphology.

Flight period
Based on the rearing data from Belize, adults are likely to occur throughout the year.

Distribution
Paratischeria neotropicana is the tischeriid species with the broadest distribution range yet documented in the Neotropics, having been recorded from various localities in Belize, Mexico, Guatemala, Ecuador, Peru and Bolivia (Stonis & Solis 2020).

Diagnosis
External characters are not sufficient for the identification of this species . In the male genitalia, Dishkeya gouaniae resembles the Nearctic D. bifurcata (Braun, 1915); however, D. gouaniae is distinguishable from the latter by its widely rounded vinculum, spiny juxta and slender apical processes of the phallus.

Biology
Host plant: Gouania polygama (Jacq.) Urb. (Rhamnaceae). Larvae mine leaves in March-July and September. The leaf mine has a linear or corridor shape (not preserved by the collector with the reared specimens, and therefore not studied in detail). This species was listed as 'Tischeria sp. 12' by Lewis et al. (2002). Of 25 living mines reared successfully, 14 (56%) were parasitised.

Flight period
Based on reared specimens, adults occur during March-September.

Distribution
Currently known only from a single locality in Belize (Las Cuevas), at an elevation of about 550 m.

Description
This species was redescribed and illustrated by Stonis et al. (2008: 165, 166, figs 1, 2a-b, 3a-e). Here, on the basis of the material from Belize, we provide the first photographic documentation of the male genitalia of the species (Figs 132-134).

Biology
Host plant: Terminalia amazonia (J.F.Gmel.) Exell (Combretaceae). Larvae mine leaves in March. The leaf mine starts as a corridor, broadening into a blotch as the larva develops. The leaf mine sample was not preserved by the collector with the reared specimen and is therefore not documented in detail.

Flight period
Based on specimens collected at light, adults occur in April. Based on reared specimens in Belize and the Caribbean (lectotype), adults also occur in March.

Distribution
This species was described from the Caribbean (Virgin Is., USA) (Walsinhgam 1897) and was recently recorded from Belize ).

Diagnosis
External characters are not sufficient for the identification of this species. In the male genitalia, Coptotriche forsteroniae resembles C. pulverea (Walsingham, 1897); however, C. forsteroniae is distinguishable from the latter and all other species of Coptotriche by the rhomboid apex of the phallus (Fig. 138)

Description
The species was described and illustrated by Stonis & Diškus (2008: 104-105, fig. 5a-c). Here, on the basis of the studied material, we are the first to provide photographic documentation of the male genitalia of the species (Figs 135-142).

Biology
Host plant: Forsteronia myriantha Donn.Sm. (Apocynaceae). Larvae mine leaves in March and probably at other times of the year. The leaf mine has a blotch shape (not preserved with the reared specimens by the collector and therefore not studied in detail and undocumented). This species was listed as 'Tischeria sp. 8' by Lewis et al. (2002). Of 6 living mines reared successfully, none were parasitised.

Flight period
Based on specimens collected at light, adults occur in April. Based on reared specimens, adults also occur in March.

Distribution
So far this species is known from a single locality in Belize, Chiquibul Forest Reserve, Las Cuevas, at an elevation of about 550 m.   Coptotriche singularis : 104, figs 2, 3a-e.

Diagnosis
External characters are not sufficient for the identification of this species. In the male genitalia, the apically narrowing valva, absence of transtilla and highly specific, apically gradually widening phallus distinguish Coptotriche singularis from other congeneric species.

Biology
The larvae are leaf miners, creating a blotch-type mine, and the host plant is recorded as Cardiospermum grandiflorum Sw. (Sapindaceae). Only a single mine was reared successfully.

Flight period
Based on the single reared specimen, adults also occur in September or early October.

Distribution
So far this species is known from a single locality in Belize, Chiquibul Forest Reserve, Las Cuevas, at an elevation of about 550 m.

Discussion
One of the most unexpected results of our study of the Belize material collected in Las Cuevas was the discovery of a diverse fauna of Tischeriidae, a total of fourteen species including representatives of four of five currently known genera of the family. This represents the highest number of species ever recorded from a single locality worldwide. In total, the documented global fauna of Tischeriidae now comprises 148 described species, including all these species reported from Las Cuevas. Twelve species are known from Europe, the most investigated region with respect to Tischeriidae: eight species of Coptotriche and four species of Tischeria (Karsholt & van Nieukerken 2013). Forty-six species are known from the Nearctic and 49 from the Neotropics, including 21 from Central America. The identified fauna of Las Cuevas, a single locality in the Central American tropical forest, exceeds the European fauna and represents almost one third of the Tischeriidae fauna of the Neotropics (for the updated Neotropical data see Stonis et al. 2020), about 67% of the fauna currently described from Central America and 9% of the documented global fauna. Considering the insufficient sampling in most of Central America, it is possible that the diversity observed at Las Cuevas is just the tip of the iceberg and that the tropical forests of Central America are more widely characterized by high richness and taxonomic diversity of Tischeriidae.
It should also be noted that the Tischeriidae documented from Las Cuevas exhibit a remarkable range of morphological structures. This is probably due to the historical and ecological biogeography of the Neotropical region, which is currently dominated by warmer conditions in comparison to the Holarctic region, and also to the particularities of the members of the host tribes of Asteraceae that predominate, and probably originated in the Neotropics (Stonis et al. 2018b).
Despite their very close external resemblance (see Figs 6-34), internally, in the male genitalia, the species from Las Cuevas seem morphologically very distinct and apparently more isolated from each other than is typical of Nearctic or Palaearctic species. Species differences in the European fauna are generally slight, and problems of identification of similar species may be compounded by intraspecific variation. The fauna of Tischeriidae from Las Cuevas comprises clearly differentiated and disjunct species, with hypothesized old origin. Our study revealed novel morphological characters previously unknown for the family, including: a specialized uncus with a bifid dorsal lobe in Astrotischeria papilloma sp. nov. and A. scutifera sp. nov.; a pectinifer on the valva of Paratischeria robinsoni sp. nov. and spine-like setae on the valva of P. tubifex sp. nov.; very large socii in P. robinsoni sp. nov.; and extremely long phallus and enormously enlarged socii in P. tubifex sp. nov. We hypothesize that these features may represent derived characters and our documentation of the genital morphology of the species from Las Cuevas contributes to a better understanding of the morphology of Tischeriidae and their possible evolutionary trends.