Estonian Caddisflies (Insecta: Trichoptera). An annotated checklist

Achecklist of 190 species of Estonian caddisflies is compiled based on literature and voucher material in collections at Estonian University of Life Sciences. 31 species are listed first time for the country while 159 taxa were referred to in literature. Two taxa are rejected as possible misidentifications (Diplectrona felix and Molanna submarginalis). 25 species are listed in Estonian Red Data book. Geographical distribution of Trichoptera both in Estonia and along the east shore of Baltic Sea is reviewed in brief

Larentiini Duponchel 1845

Tribe <i>Larentiini</i> Duponchel, 1845 <p>(Figs 98–107) (Mesoleucini McGuffin, 1958)</p> <p> Pierce (1914) described his group “Entephriinae” as follows: valva rounded (i.e. sacculus region wholly fused to), costa thickened, plate-like; manica thickly and minutely spined; anellus dorsal lobes or calcar [present]. (Pierce grouped some <i>Colostygia</i> Hübner (Fig. 83) species with <i>Entephria</i>, therefore anellus lobes mentioned); calcar produced to a long arm is mentioned in texts for <i>Entephria</i> Stephens species; medially dilated valve costa reminds that in <i>Thera</i> Stephens and allied cidariine genera but differs in pollex -like outstanding apical process.</p> <p>Following characters enable us to delimit the tribe: male genital armature with dorsal arms of vinculum dilated triangular, tegumen much slender; a thick hair lock arising from base of valve costa; female apophyses short (shared with Xanthorhoini); vinculum projecting flap-like between the bases of sacculi.</p> <p> McGuffin (1958) separated a tribe Mesoleucini from Hydriomenini s. l. according to morphological characters of larvae; Herbulot recombined these genera into Larentiini. Krüger (2005) has described some species of <i>Entephria</i> with <i>Piercia</i> -like ornamentation to juxta. Distribution: (Neotropical, Afrotropical), Palaearctic.</p>Published as part of <i>Viidalepp, Jaan, 2011, A morphological review of tribes in Larentiinae (Lepidoptera: Geometridae), pp. 1-44 in Zootaxa 3136</i> on pages 28-29, DOI: <a href="http://zenodo.org/record/279481">10.5281/zenodo.279481</a&gt

Pyrochlora majorcula Dyar 1925

Pyrochlora majorcula Dyar, 1925 Figs. 1, 4, 5, 11, 15, 25– 30 Pyrochlora majorcula Dyar, 1925: 7. Type locality: French Guiana: Cayenne. Measurements: Wing span 25.0–28.0 mm in males (n= 21), 33.0 mm in females (n= 2). Length of antennal pectinations in males 0.51–0.52 mm (n= 10) (Form A), or 0.52–0.56 mm (n= 5) (Form B), 0.45 mm in female. Description: The species is characterized by the sack-shaped aedeagus similar to but longer than that of P. rhanis, with a short apical spine; sternite A 8 sclerotized and distally straight-edged, not emarginated posteriorly. There are differences between the populations from Kaw Mts. and NE Brazil (male antennae shorter pectinated; the dorsobasal appendix from valve costa pollex-like, straight, and valve ventral margin less angulate at its basal third; Fig. 25), and from Ecuador (antennal pectinations longer; the costobasal appendix of valve rounded and valve ventral margin angulate at sacculus: Fig. 28). The entire male armature is relatively more massive in moths from Ecuador and thinner in the Fr. Guyana population (Figs. 25–30). Moths from SE Brazil differ in wings shape, relatively shorter and broader, sharing genital characters of the Form A (except a slightly more slender saccus). Material studied: Form A: 11 ɗ, FRENCH GUIANA, Kaw Mts., 6.11. 2002 (V. Soon), (gen. 7039, 7752, 7902) (IZBE); 2 ɗ, Kaw Road, 04° 32 ’N, 52 ° 12 ’ 18 ”W, 0 7.01. 2005, A. Selin & T. Armolik leg. (CAS). 2 ɗ 1 Ψ, “ BRASILIA sept. / Amazonas / Uypiranga / V. 31 / leg. W. Hopp” (ZSM, gen. 7981). Form B: 1 ɗ, ECUADOR, Sucumbios, Panayacu riv., Jarrin Expeditiones Lodge, 0° 23 ’S, 76 ° 11 ’W, 14 – 19.10.1999 (gen. 6975) (CAL); 2 ɗ 1 Ψ, Napo, Rio Shiripuno, 26.10.02; 1 ɗ, Napo, Sahuangal, 15.10. 2002 (CAS); 1 ɗ, Morona Santiago, Santiago 370m, 03°01’ 16 ”S, 77 ° 56 ’ 52 ”W, 27.04. 2007 (CAL); 1 ɗ, Pichincha, Cotundo, 1080 m, 00° 41 ’ 28 ”S, 77 ° 43 ’ 56 ”W, 19.05. 2007 (gen. 7953) (NHMT). Biology: Confined to tropical rain forests on lowland and low mountains. Distribution: Fr. Guiana, N. Brazil, Ecuador (IZBE, ZSM, CAL, CAS). Remarks: The specimens in the British Museum collection were identified by Prout (1932) and recently rechecked by Pitkin (1996). The type specimen no. 27500 was selected and designated as lectotype for Pyrochlora majorcula Dyar by L. Pitkin in the collection of Smithsonian Institution, Washington, USA. (Fig. 1).Published as part of Viidalepp, Jaan, 2009, Revision of the genus Pyrochlora Warren, 1895 (Lepidoptera: Geometridae: Geometrinae), pp. 55-65 in Zootaxa 1989 on pages 57-58, DOI: 10.5281/zenodo.18551

Chesiadini Pierce 1914

Tribe <i>Chesiadini</i> Pierce, 1914 <p>(Figs 31–37)</p> <p> Pierce (1914), while discussing characters of five genera, found female genitalia of <i>Carsia</i> Hübner and <i>Aplocera</i> similar to each other and substantially deviating from those of other larentiines. Forbes (1948) treated <i>Carsia</i> among the tribe Lobophorini. The names Eucestiini Warren, 1894 and Chesiadini Stephens, 1850 refer to the same concept, but Eucestiini is based on a younger synonym (<i>Chesias</i> Treitschke, 1825 = <i>Eucestis</i> Hübner, [1825]). Chesiadini Stephens, 1850 was proposed when <i>Chesias</i> was synonymized in the text to its younger synonym, <i>Eucestis</i> Hübner and therefore is not valid. Pierce (1914 was the first to use <i>Chesias</i> and Chesiadinae validly.</p> <p>Chesiadini has apomorphic characters summarized as follows: strongly bulbed fore femora; short, flat and strong foretibia ending in two distal teeth (valid for subtribe Chesiadina, see below); remnants of gnathos fused to the posterior margin of tegumen (separate in some aplocerine genera, Fig. 47); tergite A8 rough and heavily sclerotized at its posterior margin (Chesiadina, Figs 31–32); venation of hind wing sexually dimorphic as vein An2 is absent from male but present in female; the contact of Sc with anterior margin of discal cell also varies between species and sexes (Aplocerina, see below); fringe of wings long and relatively strong; (Cheriadina) costal basal corners prolonged and produced; valval ornamentation usually medial and costal in Chesiadina, while costal, saccular and medial in Aplocetrina; bursa copulatrix generally scobinate, signa simple or petaloid (Chesiadina).</p> <p> Here it is obvious that the tribe is to be divided into two groups of taxa as subtribes Chesiadina and Aplocerina, subtribe nov. (Figs 38–45). The aplocerine genera <i>Aplocera</i>, and <i>Carsia</i> possess following apomorphic characters of its own: foretibia slender with a single, external-apical dentiform projection; male genital armature large (Figs 38, 39, 40, 43), or medium sized (Figs 41,42); valva with dorsal and saccular ornamentations present, often long (Figs 38–40); bursa copulatrix membranous, globular, without any signa (Fig. 45); ductus bursae heavily chitinized, tubular, ending in bursa in a pair of leaf-like appendages (Fig. 45); the last abdominal segment in female is completely fused into a cone (Fig. 45), while separated by thinner pleurites in Chesiadina.</p> <p>Distribution of the Chesiadina: Old-Mediterranean. Afrotropical, (Neotropical, Nearctic). Distribution of the Aplocerina, subtribe nov. Palaearctic (penetrating as antropochorous in Australia and America).</p>Published as part of <i>Viidalepp, Jaan, 2011, A morphological review of tribes in Larentiinae (Lepidoptera: Geometridae), pp. 1-44 in Zootaxa 3136</i> on pages 11-12, DOI: <a href="http://zenodo.org/record/279481">10.5281/zenodo.279481</a&gt

Pyrochlora vogli Viidalepp, sp.nov.

Pyrochlora vogli Viidalepp sp.nov. Figs. 8, 9, 20, 37– 39 Type series: Holotype: ɗ, “ VENEZUELA, Maracay, ges[ammelt]. P. Vogl. / April–May 1934 ” (ZSM). Paratypes: 5 ɗ, “ VENEZUELA, Maracay, leg. P. Vogl” (ZSM). Additional labels: “ Juli 1934 ” (1), “ April–May 1934 ” (1), “ August 1934 ” (1), “ November 1934 ” (1), “Gen. prep. 7982 J. Viidalepp”); 3 Ψ, “ VENEZUELA, Maracay, leg. P. Vogl” (gen. prep. 7993) (ZSM). Additional labels: “ August 1934 ” (1), “Staatssamml. München” (printed label with hand-written datum “ 17.vi ” (1); “ April–May 1934 ” and white hand-written “ majorcula Dyar?” (1). The types are deposited in the collection of Zoologisches Staatsmuseum München (ZSM). Measurements: Wing span 21.0–24.0 mm in male, 26.0–29.0 mm in female. Length of antennal pectinations in male 0.45–0.48 mm, 0.3 mm in female. Diagnosis: Forewing tornal blotch enlarged, underside with dark markings separated. Hindwing with large yellow blotches at costa and tornus. Aedeagus short sack-shaped with longer apical hook. Description: Externally similar to pale P. m a j o rc u l a, but with medial and postmedial lines fused Xshaped in anal area of forewing and broken by yellow connection of light blotches distal and proximal to it, i.e the yellow tornal blotch enlarged and fused with, or connected to yellow shading proximal to the medial line. Hindwing above with postmedial line supported distally by a dull yellow line, costal yellow blotch at postmedial area large and expanded along costa towards wing base. The postmedial field at hindwing tornus with a distinct yellowish blotch, while this area is not paler in other species. Male genitalia with aedeagus similar to that in P. rhanis and P. m a j o rc u l a, sack-like with an apical spine, but aedeagus shorter (1.6 mm) and the apical spine long (0.35 mm) (Fig. 39). Valva with basal costal projection finger-shaped as in P. m a j o rc u l a. Sternite A 8 distal sclerotization weakly heart-shaped, 0.8 mm broad and incised 0.4 mm deep. Female genitalia: antrum slenderer than in P. rhanis, not bulbous (Fig. 41). Etymology: The specific name is from the family name of the collector, P. Vogl. Gender masculine. Distribution: N. Venezuela.Published as part of Viidalepp, Jaan, 2009, Revision of the genus Pyrochlora Warren, 1895 (Lepidoptera: Geometridae: Geometrinae), pp. 55-65 in Zootaxa 1989 on pages 59-65, DOI: 10.5281/zenodo.18551

Trichopterygini Warren 1894

Tribe <i>Trichopterygini</i> Warren, 1894 <p>(Figs 22–30)</p> <p>Pierce (1914) was the first to present a definition of the group (as Lobophorinae); unfortunately he also grouped two genera of the tribe Triphosini with. The genera combined now in Trichopterygini appear to be characterized by a wide ostium, and corpus bursae ornamented with petaloid signa. The variation of wing venation and sexual differences within species are markedly large.</p> <p>The tribe is relatively well studied. Dugdale (1980) described its Australian taxa and explained the modification of male sternite A2 as a shelter for androconial pencils. Parra (e.g. 1996) studied the diversity of trichopterygine genera in Chile and Holloway (1997) that in the Indo-Australian region. Holloway (1997: 100) characterizes Trichopterygini by the reduction of the anal area of male hind wing to fold, crevice, flap or lobe(s); the 2nd sternite modified in male, having an internal, hollow sack; valva in male genitalia has both costal and saccular ornamentation; bursa copulatrix is generally scobinate (shared with Chesiadini, Eupitheciini, Melanthiini etc.), the character also occurs in Idaeini (Sterrhinae).</p> <p>Trichopterygini shares the trend of modification of hind wing venation with Dyspteridini and Chesiadini. The tribe is a composite of many deeply and differently specialized elements, maintaining also archaic characters like the presence of gnathos or gnathi in some East Asiatic genera (Xue & Zhu 1999), or the occurrence of lateral appendages to juxta in a Chilean genus. Being speciose in Neotropical and Oriental faunas, only two lineages (one with the uncus conical and valva apex produced a spine, another with the uncus distal projection long and valva indented distally) have Holarctic distribution.</p> <p> Dugdale (1980) and Xue <i>et al.</i> (2008) pay attention to the modification of male hind leg characteristic of this tribe: sometimes there is a thin tibial hair pencil (Fig. 73), and the trend of spur reduction has genus-level significance. However, there are at least three trends of modification, partly overlapping. Two genera, <i>Tatosoma</i> Butler, and <i>Tricalcaria</i> Han, have lost one of the medial spurs on the normally long hind legs. The <i>Sauris</i> Guenée group of genera has proximal spurs or all spurs on male hind tibia reduced and various hair tufts present (see Dugdale 1980). A thin and long hair pencil in some Holarctic genera, such as <i>Lobophora</i> Curtis, <i>Oulobophora</i> Staudinger a.o., attached to the base of male hind tibia and concealed in sternite A2 makes out the third trait (Fig. 73). The tribe is diverse and some interesting characters occur within subclades: hind wing costal venation often different between sexes; subapical thickening or clubbing of male antennae which are apically sickle-shaped; very long abdomen in male; modifications of vestiture and scaling of wings and abdomen (Fig. 72); occurrence of abdominal or valval coremata, usually concealed in the hollow structure of sternite A2, but also as hair pencils or scale tufts on male hind legs.</p> <p> <b>Distribution:</b> Neotropical, Indo-Australian, Afrotropical, Holarctic. The morphological diversity of genera seems to be the greatest in South America (e.g. Parra 1996) and East Asia (Yamamoto and Sota 2007; Choi 2008).</p>Published as part of <i>Viidalepp, Jaan, 2011, A morphological review of tribes in Larentiinae (Lepidoptera: Geometridae), pp. 1-44 in Zootaxa 3136</i> on pages 10-11, DOI: <a href="http://zenodo.org/record/279481">10.5281/zenodo.279481</a&gt

Cidariini Duponchel 1845

Tribe <i>Cidariini</i> Duponchel, 1845 <p>(Figs 78–80, 82–83)</p> <p> Pierce (1914) discussed the genera, now placed in the tribes Cidariini and Hydriomenini, together, listing the common characters as follows: “a rather compact group, composed of a few closely allied genera. The large simple valva, and the extended anellus lobes. Often with clavate scales, … manica finely spined. The signum … small, scobinate”. However, concerning the first included genus, <i>Lampropteryx</i> Stephens, it is added that there is a “hinge” between the ninth and the tenth segments (i.e. between tegumen and uncus), and Pierce described the labides attached to the point of connection of transtilla and tegumen. Labides of <i>Lampropteryx</i> Stephens and <i>Nebula</i> Bruand have labides bipartite, with a hinge between.</p> <p>Pierce separated the subfamily Therinae for the species with ”costa well developed, sacculus extended, aedeagus ctenoid at orifice and anellus lobes strong, with spatulate scales.” Forbes (1948) has collected an over-heterogenous assemblage of genera under his equivalent of Cidariini (Hydriomenini).</p> <p>Choi (e.g. 1997), Viidalepp and Kostjuk (2005) and Viidalepp (2003) have analyzed different sets of cidariine genera using cladistic methodology. Choi (1997) found seven strict synapomorphic characters to support the monophyly of the tribe: bulged postmedial line of the forewing; long male eighth abdominal segment; oblong or posteriorly broad male eighth sternite; absence of calcar; membranous female eighth tergite; ostium with sclerotized short cingulum, which is open dorsally; SV4 seta of larval third abdominal segment outside the L1–SV1 line; moderate and rounded paraprocts in larva.</p> <p> Choi (2007) found four characters discriminating <i>Thera</i> Stephens: valve costa with medial projection; cucullus large; cornuti, a few in number, arranged corona-like around the aedeagus mouth.</p> <p> He paid attention to characters of anellus and relations of tegumen and vinculum in male genitalia, and synonymized Therini Pierce with Cidariini, Viidalepp and Kostjuk (2005) analyzed a case of character reduction on the cidariine background. Retrograde evolution of morphological characters, if occurs, then in several sections of a group, and will be ignored by cladistic methodology. Another case study of the Mediterranean genus <i>Protothera</i> (Viidalepp 2003) demonstrated how essential it is to include more than one representative species of each group into a matrix for cladistic analysis to be sure that their synapomorphic traits will be revealed. These studies revealed the presence of four main clades within Holarctic Cidariini, characterized by synapomorphic traits: conifer-feeding genera with male genital capsule well sclerotized and valva a plate,,with sacculus tip projecting scelerotized” (<i>Thera</i> Stephens and allies); presence of specialized thick setae (hamuli) on labides (<i>Nebula</i> Bruand and allies); valva with a subcostal furrow on inner side, between costa and valvula, associated with vestiture on medial wall of valva long and springing from ring-like ornamentation; thick vestiture to labides, consisting of clavate hairscales, associated with long palpi in moths.</p> <p> The variation of putative cidariine genera in Neotropical fauna (e.g. <i>Brabirodes cerevia</i> Druce, Fig. 82) is considerably larger than in boreal fauna.</p> <p> Scarce data on <i>Erateina</i> indicate a kind of similarity to Cidariini (see also discussion under Eratenini). The genus <i>Hagnagora</i> Druce has some similarities to Cidariini as well:</p> <p> However, Sihvonen <i>et al.</i> (2011) showed <i>Hagnagora</i> clustered with <i>Hydriomena.</i> This may be correct as <i>H. vittata</i> Philippi has more setae on proleg of mature larva than typical to cidariine larvae studied (King & Parra 2011; Viidalepp 2006). <i>Hagnagora clustimena</i> Druce (Figs 149, 150) has a solid male genital armature with putative clusters of androconian hairscales at uncus base and bases of valvae. <i>H. ephestris</i> Felder and Rogenhofer (Figs 151, 152) has a pyriform bursa with well defined cingulum and with two longitudinal bands of flat signa. King & Parra (2011) described the biology and larval characters of <i>Hagnagora vittata</i> Philippi.</p> <p>Distribution: Neotropical, Holarctic.</p>Published as part of <i>Viidalepp, Jaan, 2011, A morphological review of tribes in Larentiinae (Lepidoptera: Geometridae), pp. 1-44 in Zootaxa 3136</i> on pages 23-25, DOI: <a href="http://zenodo.org/record/279481">10.5281/zenodo.279481</a&gt

Asthenini Warren 1893

Tribe <i>Asthenini</i> Warren, 1893 <p>(Figs 53–57)</p> <p>The tribe was established (as Astheninae) by Warren (1893), characterized according to genitalia by Pierce (1914) and recently revised by Xue and Scoble (2002). Pierce described its characters as follows: valva in male with sacculus projecting long; female bursa copulatrix with a long, evenly spined signum; uncus weak and attached to dorsal side of anus tube, whereas ventral side of the tube bears a sclerotized subscaphium.</p> <p>Holloway (1997: 121), discussing the relations between Eupitheciini and Asthenini, noted that labides are not entirely similar in these tribes by the absence in asthenines of the branch from basal costal projection of valva towards juxta. The absence of saccular projection of valva is stressed by Pierce as another characteristic of Eupitheciini but not in Asthenini. However, Holloway grouped the relevant Indo-Australian asthenine genera to Eupitheciini according to: the presence of labides, “Springing from the points of union of the transtilla with the costa, there may arise two long arms, each bearing a soft hairy pad, and united together by a thin membrane”. Forbes (1948) found the chaetosemata of Asthenini united by a continuous row of setae in a deep groove between scales of vertex (Figs 5, 7).</p> <p>McGuffin (1958) distinguished the larvae of Asthenini as follows: the spinneret is much longer than the labial palpi; the thoracic claws are almost straight, and the angle of the notch acute.</p> <p>Xue and Scoble (2002) diagnosed the asthenine as follows: uncus distal projection either absent or vestigial; uncus fused to the anal tube; the authors reject the idea of homology of asthenine and eupitheciine labides, and suppose that the build of signum in female (consisting of thin spines radiating from the central area) is the best characteristic for the tribe.</p> <p> Large coremata loosely attached to the base of valva in Australian <i>Poecilasthena</i> Warren (Fig. 54) may be homologous, or not to those in Eupitheciini which are more tightly fused to valva base (Fig. 48). There are some synapomorphies restricted to the subclades of Asthenini: frons high and broad, cylindrical, projecting; palpi short and weak (longer in <i>Eois</i> Hübner); colliculum flattened and re-folded in female at least in boreal genera; bursa copulatrix finely spiculose.</p> <p>Asthenini and Chesiadini share the presence of long projections from the base of costa, which are not not connected to the juxta (Fig. 56). These bear setose heads in Asthenini, but not in Chesiadini.</p> <p>Distribution: Indo-Australian, Holarctic, (Afrotropical, Neotropical).</p>Published as part of <i>Viidalepp, Jaan, 2011, A morphological review of tribes in Larentiinae (Lepidoptera: Geometridae), pp. 1-44 in Zootaxa 3136</i> on pages 16-17, DOI: <a href="http://zenodo.org/record/279481">10.5281/zenodo.279481</a&gt

Solitaneini Leraut 1980

Tribe <i>Solitaneini</i> Leraut, 1980 <p>(Fig. 81)</p> <p> Leraut (1980) separated the tribe without giving a diagnosis. The genus <i>Solitsnea</i> Djakonov has some unusual for larentiines features: female bursa copulatrix with a ring of flat spines across the equatorial part: tegumen with a pair of tapering projections with unknown function; valva with all its parts well fused and chitinized, flat, without ornamentation.</p> <p> It is noteworthy that flat signa occur in female bursa of the Erateinini too, but arranged into a longitudinal band. Sihvonen <i>et al.</i> (2011) included the Solitaneini in their tree but used <i>Baptria</i> Hübner as an example species; however, <i>Baptria</i> is just an as solitary genus in Larentiinae as <i>Solitanea</i> is.</p> <p>Distribution: Palaearctic.</p>Published as part of <i>Viidalepp, Jaan, 2011, A morphological review of tribes in Larentiinae (Lepidoptera: Geometridae), pp. 1-44 in Zootaxa 3136</i> on page 27, DOI: <a href="http://zenodo.org/record/279481">10.5281/zenodo.279481</a&gt

Hydriomenini Meyrick 1872

Tribe <i>Hydriomenini</i> Meyrick, 1872 <p>(Figs 95–97)</p> <p>The tribe is discussed here as delimited by Herbulot (1962). The tribe is composite in the treatment by Forbes (1948), and Ferguson (1983) when characterized as: front sloped and tufted or thickly scaled below; metathorax strongly tufted; the upper discocellular bent in hind wing and M2 closer to M3 than M1.</p> <p> Herbulot (1961-1963) moved <i>Mesoleuca</i> Hübner and <i>Entephria</i> Stephens from Hydriomenini s. l. to Larentiini, other genera to Cidariini, Perizomini and Rheumapterini and restricted Hydriomenini for the nominate genus. McDunnough (1954), dealing with Nearctic species of the genus <i>Hydriomena</i> Hübner listed its characters as follows: specific shape of uncus, flat, bifurcate or dilated apically; modifications on costal region of valva (tubercles, bearing single stiff, curved seta („ribbon”) and locks of specialized hairscales); large, sparsely setose plate of juxta with dorsoapical papillae in male; membranous bursa copulatrix almost without signa but with heavily sclerotized ductus bursae, and semiglobular chitinous digitabulum attached to the latter in female; hemitranstillae connections to juxta membranous; manica rugose, not spiny.</p> <p>The presence of a long costal or subcostal hair lock is shared, although less complicated, by Larentiini, Stamnodini and some Neotropical genera not assigned to tribes. The digitabulum to the female bursa seems to be characteristic to the species from the northern hemispheric (Figs in McDunnough 1954).</p> <p>Distribution: Neotropical, Nearctic, (Palaearctic).</p>Published as part of <i>Viidalepp, Jaan, 2011, A morphological review of tribes in Larentiinae (Lepidoptera: Geometridae), pp. 1-44 in Zootaxa 3136</i> on pages 27-28, DOI: <a href="http://zenodo.org/record/279481">10.5281/zenodo.279481</a&gt