Diagnoses and remarks on the genera of Tortricidae ( Lepidoptera ) . Part 5

Diagnoses, redescriptions, and remarks are presented on the genera that comprise the three tortricid tribes Chlidanotini, Hilarographini, and Polyorthini. Original references, type species, type localities, synonyms, and zoogeographic regions are provided.


I. INTRODUCTION
The number of genera of Tortricidae has increased dramatically over last 40 years; by 2007 there were over 1630 described genera, including synonyms.Many of the older descriptions are scattered throughout the literature, and because there are few larger synthetic treatments of the tortricids for most major biogeographic regions, this large number of taxa complicates considerably the work of taxonomists on the faunas of poorly known regions of the planet.In addition, characters that define many of the genera are not clearly articulated.The distribution of many genera is still insufficiently known, and this shortcoming frequently results in unexpected findings, e.g., the discovery of Afrotropical genera in the Neotropics.These types of discoveries may cause confusion for specialists that focus on the fauna of a single geographic region.
The literature abounds with re-descriptions and diagnoses of tortricid genera, but many are rather short, frequently lacking comparisons with similar or related taxa.Detailed comparative diagnoses are not only useful in systematic work but are required by the International Code of Zoological Nomenclature (1999) for descriptions of new taxa.
In this series of papers on the tortricid genera, diagnoses are based on features provided in the original description, augmented by comments from subsequent papers.My own diagnoses are proposed when no earlier ones are available.Other characteristics of the genera are included when necessary or relevant.
Morphological features that define many genera require revision and/or augmentation.Also, definitions of some genera require brief comments.Some original diagnoses are quoted verbatim, especially when no subsequent evaluation has been done.
The goal of this series of papers is to present a compilation of the existing data on tortricid genera and to identify what is known and where information is incomplete or lacking.
The account for each genus consists of the original reference, type-species (t.sp.) with the countries of origin (in case of large countries also with their provinces, or large islands), the number of species included originally (e.g., monotypic), and the number of species known at present, the latter often based on the catalogue by BROWN (2005).The acronyms of the zoogeographic regions are added.Synonyms are treated in a similar way; the older, well known synonyms easily found in the literature are cited in a shortened form, i.e., without references.The references refer to re-descriptions and diagnoses.The genera are arranged alphabetically which simplifies the index to include only synonyms.
The parts of this series are published in non-systematic order, depending on the sequence of completion of each group.Part 1 (RAZOWSKI 2009a)  Only the references to comments are included in the literature cited; those to original descriptions of the taxa are omitted, as well as those that represent the generic or specific names and the names of their authors and dates of publication.All of the latter can be found in monographs and/or catalogues (e.g., BROWN 2005).
In this paper the tribes are arranged traditionally.R e d e s c r i p t i o n s.COMMON (1963), DIAKONOFF (1974).D i a g n o s i s.TURNER (1916) wrote: "In general appearance the following species [Apura xanthosoma] resembles a Laspeyresia, but there is no trace of a submedian pecten."R e d e s c r i p t i o n s.DIAKONOFF (1974), RAZOWSKI (1987RAZOWSKI ( , 2008)).D i a g n o s i s.RAZOWSKI (1965) compared Cnephasitis to Acleris HÜBNER [1825] and Polyortha.
Based on some reductions in the genitalia, RAZOWSKI (1979aRAZOWSKI ( , 1979b) )  D i a g n o s i s.RAZOWSKI (1984) compared Histurodes to Histura in colouration and male genitalia.Histurodes differs from the latter by the absence of the transtilla, the presence of strong basal apodemes connected to each other by a strong membrane, and by a subsquare, scobinate plate of the anellus.

Isotrias MEYRICK, 1895
Isotrias MEYRICK, 1895, Handbook Br. Lepid.: 542;t.R e d e s c r i p t i o n s.OBRAZTSOV (1955), RAZOWSKI (1987).D i a g n o s i s.OBRAZTSOV (1955) compared Isotrias to Olindia, placing both in Cnephasiini.RAZOWSKI (1987) also compared Isotrias to Olindia; their synapomorphies are the separate arms of the gnathos, a concave costal sclerite of the inner surface of the valva, and an oblique split of that surface.Autapomorphies of Isotrias are the presence of the dorsal grove of the uncus, the apical broadening of the gnathos, the transformed setae of the sacculus, and the shape of the sterigma.RAZOWSKI (1979b) transferred Isotrias to Polyorthini.
Lopharcha DIAKONOFF, 1941Lopharcha DIAKONOFF, 1941, Treubia, 18: 424;t R e d e s c r i p t i o n.DIAKONOFF (1974).D i a g n o s i s.Originally, Lopharcha was compared to Lophoprora ("structurally allied, but of uncertain affinity").According to the original description (DUGDALE, 1966), Laciniella "shares a common genitalic facies, the indented forewing dorsum, and raised scales with the Australian genera, Apura TURNER, and Polylopha.Polylopha lacks a thoracic crest.... Finally, in the adult Laciniella, the condition of R5 (ending on the costa) is possibly foreshadowed in Palaearctic genera such as Euledereria FERNALD, Cnephasia CURTIS ... [etc]."R e m a r k s.Laciniella was described in Cnephasiini; the genitalia are characterized by the absence of hami, the possession of a gnathos, and a plate-shaped signum.
R e d e s c r i p t i o n.DIAKONOFF (1939).D i a g n o s i s.MEYRICK (1930) concluded that this genus was related to Spatalistis (Tortricini); DIAKONOFF (1939) stated: "related to the Cnephasia-group and not to Spatalistis."DIAKONOFF (1974) transferred it to Polyorthini mentioning that "the signum is small, and rod-like."R e m a r k s.Lophoprora is also mentioned under Cnephasitis.
R e d e s c r i p t i o n.BROWN & ADAMSKI (2003).D i a g n o s i s.Originally, the male genitalia of Lypothora were compared to those of Polyortha ("and other allied genera"); BROWN & ADAMSKI (2003) compared Lypothora with Histura and Histurodes.R e d e s c r i p t i o n s.COMMON (1963), DIAKONOFF (1974).D i a g n o s i s.According to COMMON (1963), Polylopha "is closely allied to Apura TURNER, with which genus it shares the curious indentation of the dorsum in the forewing and the relative position of R 5 in the forewing which terminates on the costa." Polyortha DOGNIN, 1905 Polyortha DOGNIN, 1905 D i a g n o s i s.According to the phylogenetic hypothesis of RAZOWSKI (1999), Pseuduncifera is the sister group to Biclonuncaria + Clonuncaria and differs from that clade by having eight supposed autapomorphies: pseudouncus present, reduced gnathos replaced by a lateral sclerite, lateral sclerite of the gnathos with a process, tuba analis strongly slerotized and elaborate, and four signa in the corpus bursae.
R e d e s c r i p t i o n .DIAKONOFF (1974).D i a g n o s i s.In his original description of Scytalognatha, DIAKONOFF (1956DIAKONOFF ( , cited in 1974) ) wrote: "superficially the insect suggests a Peronea (Acleris auctt.).The unarmed and simplified valva is puzzling; it prevents the attribution of the genus to either of the tribes Cnephasiini and Peroneini (Tortricini auctt., and suggests a relationship with the tribe Cacoeciini [Archipini]."R e m a r k s.Scytalognatha is also mentioned under Apura.

R e d e s c r i p t i o n. RAZOWSKI (1981) transferred it to Polyorthini.
Diagnoses of the genera of Tortricidae.Part 5 D i a g n o s i s.There is no comparative diagnosis.RAZOWSKI (1981) placed Sociosa near Histura and listed some specialized genital characters, viz., the shape of the socius, the weak gnathos, a short costa of the valva and dorsal portion of anellus.One can add the shape and position of the uncus, which is short and dorsal, and the presence of numerous pectinate scales on the socii.R e d e s c r i p t i o n s.DIAKONOFF (1979DIAKONOFF ( , 1986)), RAZOWSKI (1987).D i a g n o s i s.DIAKONOFF (1979) compared Charitographa to Thaumatographa stating "differing considerably by the female genitalia, clothed inside with a stiff layer of modified spines and devoid of signa.The male has a thick and short aedeagus, unique in the group."DIAKONOFF (1986) wrote: "This monobasic genus is a peculiar development of the Thaumatographa type, differing considerably by the peculiarities of the venation of the forewing, by the robust aedeagus of the male, but chiefly by the female genitalia."D i a g n o s i s.According to the original description, Compsocommosis "is related to the Asian Mictocommosis but differs significantly in genital characters, with the female having the sterigma split posteriorly into two plates, unique sclerotized and dentate plates at the ductus bursae-bursal border, and a single small thorn-like signum.The genitalia characters are very different from those of Mictocommosis: females have distinctive sterigma-ostium and ductus plates, both of which are simplified in Mictocommosis, and ductus plates are lacking in the latter genus.The forewing has only 2 apical silver lines (three in Nexosa).The male is unknown."

J. RAZOWSKI
D i a g n o s i s.DIAKONOFF (1977) wrote that Embolostoma has "facies rather intermediate between Thaumatographa WALSINGHAM and Nexosa gen.nov.Unusual and striking are the long coloured palpi, pointed and beak-like, entirely different from the two other genera."
D i a g n o s i s.DIAKONOFF (1977) characterized Hilarographa and Thaumatographa as follows: labial palpi "moderate, curved and ascending, rather smoothly scaled and not flattened.In Thaumatographa the labial palpi are peculiar: rather slender and long, hardly dilated basad and more or less distinctly flattened dorso-ventrally, at least towards apex ... Male genitalia differ by the uncus in Hilarographa being more or less subsclerotic at base...In Thaumatographa the uncus is always rigid, a strongly curved and porrect hook...The socii in the first genus are flattened, rather broad, ... In Thaumatographa the socii are of two types..." The original comparative diagnosis of Idiothauma is as follows: "Allied to Hilarographa, Z., but differing in vein 8 of the forewing running to the termen instead of the costa, in the much straighter costa and the conspicuous indentation between veins 5 and 6." R e m a r k s.RAZOWSKI (2009c) listed 60 species of Hilarographa and two species placed in Thaumatographa.Hilarographa is also mentioned under Thaumatographa.

Nexosa DIAKONOFF, 1977
Nexosa DIAKONOFF, 1977, Zool.Verh.Leiden, 158: 12; t. sp.: Mictopsichia marmarastra MEYRICK, 1932, Java.Four species included.OR, AU.D i a g n o s i s.Nexosa was originally compared to Cnephasitis based on the female genitalia and abdomen.DIAKONOFF (1977) mentioned that "there are, however, no obvious coremata, but the valvae are distinctly split at the posterior portion; in this respect the genus resembles Cnephasitis RAZOWSKI, that also has split of valvae but no coremata."R e m a r k s.Nexosa was described in Hilarographini but RAZOWSKI (2009c) placed it in Archipini based chiefly on the female genitalia.Re-consideration of this decision is, however, required.For the time being I am returning it to its original placement in Hilarographini.
R e d e s c r i p t i o n s.DIAKONOFF (1986), RAZOWSKI (1987).D i a g n o s i s.DIAKONOFF (1977) compared Thaumatographa to Hilarographa which differ from the latter mainly in having the labial palpus flattened dorso-ventrally at least in its distal part, and rough scaled.For other differences see the diagnosis under Hilarographa.
Diagnoses of the genera of Tortricidae.Part 5 R e m a r k s.Some remarks on Hilarographa, Mictopsichia, Charitographa and the tribe Hilarographini were presented by RAZOWSKI (2009c).

Chlidanotini
D i a g n o s i s.RAZOWSKI & BECKER (1999) compared the male genitalia of Auratonota to those of Macrochlidia and Monortha, based mostly on the similar socii and hami.

D i a g n o s i s. There is no comparative diagnosis.
Caenognosis WALSINGHAM, 1900Caenognosis WALSINGHAM, 1900 [in]  R e d e s c r i p t i o n.COMMON (1965).D i a g n o s i s.There is no original comparative diagnosis for Epirrhoeca.COMMON (1965) compared Caenognosis to Trymalitis: "the forewing is narrower than in Trymalitis and the apex much more prominently produced.The prominent indentation of the ventral margin of the valva in the male, the presence of a large cornutus spine in the aedeagus, and the shape of the uncus, serve to distinguish the genus from Trymalitis and the other two Australian genera."TUCK (1981) mentioned that Caenognosis and Chlidanota have a medially narrow transtilla with lateral, ventral extensions different than those of Electracma and Metrernis, the latter of which have a medially broad, laterally narrow transtilla.

D i a g n o s i s.
There is no original comparative diagnosis.TUCK (1981) mentioned that Caenognosis and Chlidanota have a similar transtilla, different from those of Electracma and Metrernis (see diagnosis under Caenognosis).
Daclocnema COMMON, 1965 Daclocnema COMMON, 1965, Aust. J. Zool., 13: 715;t. sp.: Daclocnema epicharis COMMON, 1965; Queensland, Australia.One species included.AU.R e d e c r i p t i o n.COMMON (1965).D i a g n o s i s.According to the original description (COMMON 1965), the venation of Daclocnema is "close to that of Archimaga MEYR.( 1905), but the male genitalia differ greatly from those of the type species A. pyractis MEYR.and of A. philomima MEYR..In addition to the wing shape, the venation, and the genitalia characters, Daclocnema may be distinguished from Trymalitis and Caeonognosis by the subacute apical segment of the labial palpus and by the smooth-scaled anterior and median tibiae." Electracma MEYRICK, 1906Electracma MEYRICK, 1906, J. Bombay Nat. Hist, Soc., 17: 413;t D i a g n o s i s.In the original description, DIAKONOFF (1976) compared Gnaphalostoma to Leurogyia: "differing from all other genera, except Leurogyia COMMON, by the long, fringed palpi, and from the last mentioned genus, by neuration, broader hind wing, and genitalia".
Heppnerographa RAZOWSKI, 1987Heppnerographa RAZOWSKI, 1987, Bull. Acad. Pol. Sci, Sci. Biol., 35: 63;t.Iconostigma TUCK, 1981Iconostigma TUCK, 1981, Syst. Entomol., 6: 339;t. sp.: Iconostigma tryphaena TUCK, 1981, New Caledonia.Two species included.AU.D i a g n o s i s.According to the original description (TUCK, 1981), "the homologies of the R veins in the forewing of Iconostigma may be demonstrated by a comparison with the Diagnoses of the genera of Tortricidae.Part 5 venation of Trymalitis and Caenognosis.Iconostigma has similar venation to Trymalitis, but the course of M1 resembles that of the male of Caenognosis...In Iconostigma the venation has been further reduced: the basal part of M1 has been lost, so that M1 is stalked with R5." Leurogyia COMMON, 1965 Leurogyia COMMON, 1965, Aust. J. Zool., 13: 720;t. sp.: Leurogyia peristictum COMMON, 1965, Queensland, Australia.One species included.AU.D i a g n o s i s.In the original paper (COMMON, 1965), Leurogyia is compared to Picroxena MEYRICK from Java from which it differs in venation.From all known genera Leurogyia differs in "the spatulate uncus, the highly developed transtilla, the strongly pistol-shaped tortricid aedeagus and the deeply concave valvae in the male, and the sclerotized ostium bursae and ductus bursae and the deeply invaginated signum in the female....The curious abdominal organ, found in the male of Picroxena, is absent in Leurogyia."Macrochlidia BROWN, 1990 Macrochlidia BROWN, 1990, J. N.Y.Entomol.Soc., 98(3): 370; t. sp.: Macrochlidia major BROWN, 1990, Colombia.Three species included.NEO.

R e d e s c r i p t i o n. RAZOWSKI & PELZ (2005).
D i a g n o s i s.BROWN (1990)

D i a g n o s i s.
There is no original comparative diagnosis of this genus.
J. RAZOWSKI COMMON (1965) compared Picroxena to Leurogyia (see the diagnosis of the latter).
Pseudocomotis BROWN, 1989 Pseudocomotis BROWN, 1989, Ent. scand., 20(4): 440; t. sp.: Orthotaenia scardiana DOGNIN, 1905, Ecuador.Seven species included.NEO.R e d e s c r i p t i o n s.RAZOWSKI & BECKER (1999), D i a g n o s i s.In the original description BROWN (1989) mentioned that superficially Pseudocomotis is most similar to Auratonota and in genitalia to Monortha "with which it shares the possession of long spines from venter of the uncus.It differs from Monortha in having the socii and hami independent (fused in Monortha) and in its absence of long, strong, spine-like setae from the socii (well developed in Monortha).Autapomorphies for Pseudocomotis include the broadly excavated distal margin of the valva dividing the valva into a distinct dorsal and ventral portions, and the patch of spines near mid-venter of the valva."R e m a r k s.BROWN (1998) discussed the phylogeny of Pseudocomotis, hypothesizing that it is probably the sister group to Monortha.RAZOWSKI & BECKER (1999) mentioned the putative autapomorphies for Pseudocomotis.
Trymalitis MEYRICK, 1905Trymalitis MEYRICK, 1905, J. Bombay Nat. Hist. Soc., 16: 590;t.R e d e s c r i p t i o n.COMMON (1965).D i a g n o s i s.MEYRICK (1905) diagnosed Trymalitis as follows: "Probably belonging to the group of Commophila [Cochylini], but very distinct by the neuration of forewings."COMMON (1965) compared Trymalitis to Caenognosis (see diagnosis of the latter).
sp.: Heppnerographa arammclaina RAZOWSKI, 1987, Costa Rica.Thirteen species included.NEO.R e d e s c r i p t i o n.RAZOWSKI & BECKER (1999).D i a g n o s i s.RAZOWSKI & BECKER (1999) compared Heppnerographa to Auratonota and Utrivalva (see diagnosis under the latter).