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Taxonomic History of the Xyloryctidae
Taxonomic History of the Xyloryctidae
Cryptophasidae, Swainson, W., and Shuckard, W.E., 1840, On the History and Natural Arrangement of Insects. Lardner's Cabinet Cyclopedia. Longman, Orme, Brown, Green, & Longmans, London, 1-406 (106-107).
Cryptophasidae. Newman, 1841, Analytical Notice of the 129th Volume of Lardner's Cabinet Cyclopcedia, entitled ‘On the History and Natural Arrangement of Insects:’ By William Swainson, A.C.G., F.R.S. & L.S., Hon. F.C.P.S., and of several Foreign Societies ; and W. E. Shuckard, Lib. R.S., &c., The Entomologist, III, 33-48 (38-41).
Cryptolechiidae Meyrick 1883, On the classification of some families of the Tineina. Transactions of the Royal Entomological Society of London 1883: 119-131 (124-125).
Cryptolechiadae Meyrick, 1887, Descriptions of new Lepidoptera. Proc. Linn. Soc. N.S.W. 2-n.s. 1(4): 1037-1048 [1040].
Cryptolechiadae. Meyrick, 1890, Descriptions of Australian Lepidoptera. Part I. Xyloryctidae. TRSSA 13: 23–81 (205).
Xyloryctidae Meyrick, 1890, Descriptions of Australian Lepidoptera. Part I. Xyloryctidae. TRSSA 13: 23–81 (205).
Cryptolechidae. Walsingham, 1891, African Micro-Lepidoptera, Transactions of the Entomological Society of London, 39, 63-133, (100-102).
Xyloryctidae. Walsingham, 1891, African Micro-Lepidoptera, Transactions of the Entomological Society of London, 39, 63-133, (100-102).
Cryptophasidae. Kirby, W.F., 1897. A Handbook to the Order Lepidoptera, Lloyd’s Natural History, Vol V, part III. Edward Lloyd, London, 1-332 [303].
Xyloryctidae. Meyrick, 1897, Descriptions of new Lepidoptera from Australia and New Zealand. Transactions of the Entomological Society of London 1897: 367–390 (382).
Xyloryctinae, Gelechiidae. Blandford, W.H.F., [report on an exhibition of Xyloryctinae presented by Lord Walsingham], Entomological Society of London, 3 March 1898. Entomologists’ Monthly Magazine, 34, 91-92, [91].
Xyloryctidae. Turner, 1898. The Xyloryctidae of Queensland. Ann. Qd Mus. 4: 1–32 (3).
Xyloryctidae. Turner, 1900, New Micro-lepidoptera -- mostly from Queensland. TRSSA 24: 6-23 (6).
Xylorictidae. Dyar, H.G. 1902 [1903]. A list of the North American Lepidoptera and key to the literature of this order of insects. Bulletin of the United States National Museum 52: i-xix, 1-723 (518).
Xyloryctidae. Meyrick, 1904, Descriptions of Australian Micro-lepidoptera. XVIII. Gelechiadae. Proc. Linn. Soc. N.S.W. 29: 255-441 (256).
Xyloryctidae Meyrick, 1890. Meyrick, 1906, Descriptions of Australian Tineina. TRSSA 30: 33–66 (50).
Stenomatidae, Walsingham, 1907, Descriptions of New North American Tineid Moths, with a Generic Table of the Family Blastobasidae, No. 1567. October 29, 1907, Proceedings of the United States National Museum, 33, 197-228 [214].
Xyloryctidae. Meyrick, 1914, Descriptions of New Zealand Lepidoptera, Transactions and Proceedings of the New Zealand Institute, XLVII, 205-244 (221).
Uzuchidae, Hampson, G.F. 1918. Some small families of the Lepidoptera which are not included in the key to the families in the Catalogue of Lepidoptera Phalanae, a list of the families and subfamilies of the Lepidoptera with their types, and a key to the families. Novitates Zoologicae 25: 366-394 (386).
Crytophasidae. Fletcher, 1929, A list of generic names used for Microlepidoptera.Memoirs of the Department of Agriculture of India, Entomology 11: 1-244
Xyloryctidae. Diakonoff, 1954, Microlepidoptera of New Guinea. Results of the third Archbold Expedition (American-Netherlands Indian Expedition 1938-1939. Part 4. Verhandelingen der Koninklijke Akademie van Wetenschappen [Verh. Akad. Amsterdam] 2 ser. 50 (1): 1-191
Xyloryctidae. Gates Clark, J. F., 1955. Catalogue of the Type Specimens of Microlepidoptera in the British Museum (Natural History) Described by Edward Meyrick. Vol. 1.
Xyloryctidae. Duckworth, 1964, North American Stenomidae (Lepidoptera: Gelechioidea), Proceedings of the United States National Museum, Smithsonian Institution, Washington, D.C., 116, No. 3493, 23-72 (25-26).
Xyloryctidae. Common, 1970: Lepidoptera (Moths and Butterflies), The Insects of Australia, Melbourne University Press, 765-866 (822-824).
Xyloryctidae. Duckworth, 1973, The Old World Stenomidae: A Preliminary Survey of the Fauna, Notes on Relationships, and Revision of the Genus Eriogenes (Lepidoptera: Gelechioidea), Smithsonian Contributions to Zoology No. 147, 1-21.
Xyloryctinae (Gelechiidae). Zimmerman, 1978, Microlepidoptera, Gelechioidea, Insects of Hawaii, A Manual of the Insects of the Hawaiian Islands, including an Enumeration of the Species and Notes on their Origin, Distribution, Hosts, Parasites, etc., The University Press of Hawaii, Honolulu, 9, ii 1-1903, (928 – 931).
Xyloryctinae (Oecophoridae). Common, 1990, Moths of Australia, Melbourne University Press, 1-535 (227-230).
Xyloryctinae (Oecophoridae). Scoble, 1992, The Lepidoptera: Form, Function and Diversity, (243-244).
Xyloryctinae (Oecophoridae). Robinson, Tuck, & Shaffer, 1994, A Field Guide to the Smaller Moths of South-East Asia, Natural History Museum, London, 1-308 ((61).
Xyloryctinae (Oecophoridae). Hodges, 1998, The Gelechioidea, in Kristensen, 1999, Handbook of Zoology, Volume IV, Arthropoda: Insecta, Part 35, Lepidoptera, Moths and Butterflies Vol, 1,
Xyloryctinae (Oecophoridae). Meyrick, 1890. Common, in Nielsen, Edwards, Rangsi, 1996, Checklist of the Lepidoptera of Australia. Monogr. Aust. Lepid 4: i-xiv, 1-529 & CD-ROM [87].
Xyloryctidae. Holloway, 2001, The families of Malesian moths and butterflies, Fauna Malesiana handbooks, (205).
Xyloryctinae, Xyloryctidae. Kaila, 2004, Phylogeny of the superfamily Gelechioidea (Lepidoptera: Ditrysia): an exemplar approach, Cladistics 20 (2004) 303–340.
Xyloryctinae, Xyloryctidae. Hoare, 2005, Hierodoris (Insecta: Lepidoptera: Gelechioidea: Oecophoridae), and overview of Oecophoridae, Fauna of New Zealand, Ko te Aitanga Pepeke o Aotearoa, 54 (1-102) [13-25].
Xyloryctidae, Xyloryctinae. Richard Brown, Sibyl Bucheli, and SangMi Lee, 2006, Gelechioidea, A Global Framework (web page). http://mississippientomologicalmuseum.org.msstate.edu/Researchtaxapages/Lepidoptera/Xyloryctidae/Xyloryctidaehome.html
Xyloryctidae. Zborowski and Edwards, 2007, A Guide to Australian Moths, CSIRO, 1- 214.
The natural divisions of the Bombycides are completely unknown; but as, for the purpose of reference, and the more easy detection of species, it is essential to make some rude assessment of the genera, we shall place them under the following heads, until a better acquaintance with the whole has been obtained by analysis: — Hepialidae, or ghost moths; 2. Bombycidae, or silk spinners; 3. Arctiadae, or tiger moths; 4. Lithosiadae, or day-moths; 5. Cryptophasidae, or hermit moths.
These divisions may be thus slightly characterised.— ...
... the fifth and last division seems closely allied, both to the Tortricidae in the next family, and to the Hepialidae in this; they are, however, distinguished by an economy so remarkable, that we shall view them, for the present, as a separate group. The larva, in shape, much resembles that of the ghost moth; the head is large, and on the body are a few scattered hairs: the chrysalis is not enclosed by any web, but lies within the chamber, or habitation, previously made by the caterpillar in the solid trunk of the tree. The perfect insect differs from all the families we have here noticed, in having the palpi curved up before the eyes, and diverging: the antennae, also, are very long; and the wings possess a particular glossiness. We are indebted to the late J.W. Lewin for a knowledge of these extraordinary moths, hitherto found only in New Holland, and which he has admirably illustrated in his “Prodromus” of Australian Entomology. It is, perhaps, owing to the rarity of this book, that modern systematists appear to be but little acquainted with these singular insects.
[Swainson provided the system of classification used in this book, parts of which were written by Shuckard. The full classification is Order Lepidoptera, Phalaenae, Bombycides, Cryptophasidae; this follows in principle Lewin’s original classification, Fam. Bombyx, Sect. Cryptophasa.
Cryptophasidae, strictly speaking, has precedence over Xyloryctidae. Later authors were either unaware of this reference or take Kirby's 1897 use of Cryptophasidae as the primary source. However, Xyloryctidae is a term hallowed by much usage.
The term Cryptophasidae is still current in India, viz: Jayanth, K.P.; Nagarkatti, S. (1981) Feasibility of rearing Nephantis serinopa Meyr. (Lepidoptera: Cryptophasidae) on an artificial diet. Journal of Entomology Research 5(2) 43-74.]
(Shuckard, in Swainson, 1840).
Cryptophasidae
[Newman’s sole reference to the family. In this work, Newman mercilessly criticises Swainson’s systematics, mocking his association with the apparently discredited ‘MacLeayian or quinarian’ system of classification.]
(Newman, 1841).
Cryptolechidae
In general this family [Oecophoridae] maybe distinguished ... from the Cryptolechidae by the parallel veins 6 and 7 of the hindwings, ...
[This paper also includes an explanation of Meyrick’s numbering system for antennae and cilia.]
In the following descriptions, a number placed after the description of the antennal ciliations indicates the length of the ciliations in terms of the breadth of the stalk of the antennae e.g., 3-5 signifies that the ciliations are from three to five times as long as the breadth of the antennal stalk. Similarly, a number after the cilia of the hindwings denotes the length of the cilia in terms of the breadth of the hindwings.
(Meyrick 1883, 417).
Cryptolechiadae
[Used to head the description of Cryptophasa leucadelpha and C. [Thysiarcha] eccleciastis.]
(Meyrick, 1887).
Perhaps some exotic forms described under the name of Cryptolechia are to be referred here; but I am indebted to Lord Walsingham for pointing out that the original type of Zeller’s Cryptolechia belongs in fact to the Oecophoridae. It is therefore impossible to employ for this family the name Cryptolechiadae, which I formerly used for it, and I have renamed it accordingly.
(Meyrick, 1890).
XYLORYCTIDAE.
Head smooth or with more or less loosely appressed hairs; ocelli absent; tongue developed. Antennae 2/3 – ¾, in male pectinated, ciliated, or simple, basal joint without pecten. Labial palpi recurved, terminal joint pointed. Maxillary palpi very short, more or less appressed to tongue. Abdomen in male with uncus developed, variable in length. Forewings with vein 1 furcate towards base, 7 and 8 stalked or rarely separate or coincident, 11 from middle of cell. Hindwings as broad or generally broader than forewings, trapezoidal to ovate, 1b clothed with long hairs above towards base, shortly furcate at base, 3 and 4 from a point or stalked, 6 and 7 stalked or approximated towards base, 8 connected with upper margin of cell by a short bar.
Most related to the Oecophoridae; probably the two families are parallel developments from a common source; they are analogous in many respects, but are easily separated by the neuration of the hindwings. None of the Xyloryctidae possess the basal pecten of the antennae, which is so common in the Oecophoridae. Whether this family is represented to any extent outside Australia, I am not at present able definitely to say. One species alone is found in New Zealand; it is an Australian insect, which has perhaps made its way thither within recent times. Perhaps some exotic forms described under the name of Cryptolechia are to be referred here; but I am indebted to Lord Walsingham for pointing out that the original type of Zeller’s Cryptolechia belongs in fact to the Oecophoridae. It is therefore impossible to employ for this family the name Cryptolechiadae, which I formerly used for it, and I have renamed it accordingly. It consists of a group of Australian genera which are intimately connected together; and even if it should be found hereafter that many South American and African forms are capable of being placed with them, they would probably not interrupt the close connection of the Australian genera, and any systematic change that might be necessary would perhaps be rather in the direction of a widening of the family characters.
The structure of the head is essentially identical with that of the Oecophoridae. The neuration of the forewings is also identical in the typical forms, except that vein 2 is commonly much more widely remote from the angle of cell; but there is a wider range of structure, since there can be no question that the forms in which veins 7 and 8 are separate are rightly included. The hindwings are almost always relatively broader, and the neuration as described contains the essential points of distinction of the family; but I may say that the connecting bar between vein 8 and the cell, often very short when these are close together, is by no means so conspicuous a structure as might be supposed, and may very readily be passed over, especially when near the base; I have however satisfied myself that it is invariably present.
[Describes the Xyloryctidae. Meyrick’s description includes the Stenomatidae and the Autostichinae; Meyrick did not establish the Stenomidae until 1906 (Stenomatidae Walsingham, 1907) and the Autostichinae were not established until Le Marchand, 1947, Les Tineina, Revue Fr. Lépidopt., 145-163 (153). A single species of Depressariinae is included: Meyrick had already established the Depressariidae in 1883.]
(Meyrick, 1890).
If the name Cryptolechia is to be retained for the species originally described as the type of that genus, and I fail to see how the rule can be departed from in this instance, any family founded upon an alliance with that genus must at least retain its essential characters, and cannot be established to include the forms in which veins 6 and 7 of the hind wings are not separated, this wide difference in neuration being admitted by all authorities to be of the utmost importance in systematic classification. It follows that Zeller's genus Cryptolechia falls into the family Oecophoridae of Meyrick, and annihilates Meyrick' s family Cryptolechidae, which was not founded on the typical form. Mr. Meyrick, recognising this, has since recharacterised his family Cryptolechidae under the name Xyloryctidae (Tr. Roy. Soc. South Australia, 1890, 234). ...
In pointing out these inconsistencies I have had the great advantage of possessing nearly the whole series of Zeller's generic types, in many cases the actual specimens used by him in writing his descriptions, and where these are not available, specimens of each species from his own collection, named in his handwriting.
It is not surprising that Mr. Meyrick, without the guidance of such valuable material, should have apparently failed to identify the precise form of neuration characteristic of the original genus Cryptolechia, which is as follows :
Fore wings 12 veins; 2 from near lower angle of cell; 7 and 8 from a common stem, the fork enclosing the apex; the rest separate. Hind wings 8 veins; 3 and 4 from a point at lower angle of cell; 5 bent over at its origin and somewhat approximate to 4 ; 6 and 7 separate, almost parallel ; 1b furcate at base ; 8 joined to upper edge of cell by a cross vein. [PI. VII., fig. 86.]
Cryptolechia straminella, Z., ; a head, b fore wing, c hind wing.
(Walsingham, 1891).
GENUS CRYPTOPHASA. (Cryptophasidae)
[The only reference in Kirby’s works to Swainson’s family Cryptophasidae.]
(Kirby, 1897).
This genus [Eschatura] belongs to the group formerly maintained as a distinct family under the name Xyloryctidae, and is intermediate between Uzucha and Pilostibes.
[Somewhere between 1890 and 1897 Meyrick has repudiated the Xyloryctidae and returned them to the Gelechiadae. However he is still using the term in 1898, TLSNSW 22.]
(Meyrick, 1897).
Lord Walsingham exhibited a series of larger and more striking species of Xyloryctinae, a subfamily of the Gelechiidae, charachteristic of the Australian fauna. The series illustrated the life-histories and the great disparity in colour and form between the sexes of many species. He also gave an account of the family, chiefly from notes by Mr Dodd, of Queensland, with special reference to the habits of the larvae, which live in holes in tree-trunks, to which they drag leaves in the night for the next day’s consumption.
(Blandford, 1898).
The family Xyloryctidae was instituted by Mr. E , Meyrick, B.A., F.E.S., to receive a large and important section of the Tineina. His monograph, published in the transactions of the Royal Society of South Australia, 1890, page 23, laid the foundation of all our knowledge of this group, which he defines as follows:
“Head smooth or with more or less loosely appressed hairs; ocelli absent; tongue developed. Antennae 2/3 – ¾, in male pectinated, ciliated, or simple, basal joint without pecten. Labial palpi recurved, terminal joint pointed. Maxillary palpi very short, more or less appressed to tongue. Abdomen in male with uncus developed, variable in length. Forewings with vein 1 furcate towards base, 7 and 8 stalked or rarely separate or coincident, 11 from middle of cell. Hindwings as broad or generally broader than forewings, trapezoidal to ovate, 1b clothed with long hairs above towards base, shortly furcate at base, 3 and 4 from a point or stalked, 6 and 7 stalked or approximated towards base, 8 connected with upper margin of cell by a short bar.”
The family is most nearly related to the Oecophoridae, and many species of both families present such a close general resemblance that care is necessary to avoid confusing them. This can always be done by observing the neuration of the hind wings, and there are also usually other points of distinction which are however not quite absolute. For the classification the genera we are indebted to Meyrick's paper quoted above, which will be repeatedly referred to in the following pages. In the few instances in which I have ventured to differ from Mr. Meyrick it has been in the endeavour to apply his methods to the more ample material at my disposal.
A number of species have been more recently described by Dr. T.P. Lucas, of Brisbane, and by Mr. Oswald Lower, of South Australia. Of these, I have noticed all those of the former author, having had the good fortune to obtain access in all but one instance to the original types. Those of Lower’s species which occur in Queensland are also referred to.
This paper owes its value mainly to the splendid collection placed at my disposal by Mr. R. Illidge, whose assiduous labours in the discovery and rearing of the larvae have resulted in a rich harvest of specimens of new or previously little known species. Mr. F.P. Dodd has obtained a small but highly interesting collection of specimens reared from larvae found in the neighbourhood of Charters Towers; these are now in Mr. Illidge’s collection. Our previous knowledge of this family in Queensland was mainly due to specimens obtained by the late Mr. G. Barnard, of Duaringa.
Yet there remain however many new species, more particularly in the less conspicuous genera, to be discovered in the locality of Brisbane, and our knowledge of those in the more distant parts of the colony is still extremely fragmentary.
(Turner, 1898).
This family is now merged by Mr. Meyrick in the Gelechiadae, but as the Australian Gelechiadae, with the exception of this section, have not yet received systematic treatment, it is convenient to retain the above designation [Xyloryctidae] for the present. The present contribution is supplementary to my paper on the Queensland Xyloryctidae in the Annals of the Queensland Museum, No. 4, 1897.
(Turner, 1900).
Family Xyloryctidae
[Includes the two North American Stenomatine genera Stenoma Zeller 1854 and Ide Chambers 1879].
(Dyar, 1902).
Assuming that the Xyloryctidae are maintained as a distinct family (which still appears to me to be convenient, though I think ultimately it must be reduced to a group of the Gelechiadae), I rely for distinction mainly on the character of vein 2 of the forewings, which in that group rises widely remote from 3 (generally disproportionately so); considering this in combination with other characters, I have not hitherto found any species as to which I had the least doubt. From other families the Gelechiadae are most reliably distinguished by the connection of 8 of hindwings with cell by a more or less evident bar; this is not always easy to observe, but the sinuation or emargination of termen is usually perceptible, and where this fails, the greater width of hindwings relatively to forewings, or the approximation of veins 6 and 7 at base are frequent characters which help to distinguish from the Oecophoridae, in which family they never occur.
(Meyrick, 1904).
STENOMIDAE.
I propose to constitute this a distinct family. It agrees in the main characters with the Xyloryctidae, but differs in having veins 7 and 8 of the forewings separate. To this family I refer the genus Agriophara, now containing about twenty species; this is the only Australian genus at present known to me, but the New Zealand genus Hypeuryntis also belongs here. The family is very extensively represented in South America, which appears to be its home.
(Meyrick, 1906).
Family STENOMATIDAE.
= Xyloryctidae Dyar, Bull. U. S. Nat. Mus., No. 52, 1902, pp. 518-9.
Allied to Xyloryctidae Meyrick, but differing in having veins 7 and 8 of the forewings separate.
This family is characteristic of tropical America, but would include Agriophara Rosenstock [Ann. Mag. Nat. Hist. (5), XVI, 1885, p. 439.] (the only Australian genus with veins 7 and 8 of the forewings separate referred by Meyrick to the Xyloryctidae) and a few Indian forms.
The species belonging to various genera of the Stenomatidse have been generally erroneously described as “Cryptolechia” (Oecophoridae) which genus differs in having 7 and 8 of the forewings stalked, and 6 and 7 of the hindwings separate and parallel.
(Walsingham, 1907).
Xyloryctidae.
Head with loosely appressed scales. Labial palpi long, recurved, acute. Maxillary palpi very short, appressed. Forewings with 2 remote from angle, 7 and 8 stalked or separate. Hindwings broadly trapezoidal, apex obtuse, termen faintly sinuate; 3 and 4 connate, 5 rather approximated, 6 and 7 approximated or stalked.
A large family, chiefly found in the Southern Hemisphere and Indian regions; most numerous in South America.
[Here Meyrick conflates the Stenomidae with the Xyloryctidae once more].
(Meyrick, 1914).
LIST OF THE FAMILIES AND SUBFAMILIES OF THE LEPIDOPTERA. The types of the genera are the first species in the author's original list, when he does not cite the type, which agrees with his generic description. The Family and Subfamily names are derived from the oldest generic name in the respective groups. The names from Hübner's Verzeichniss should strictly be excluded as not binomial ; his stirps are the genera and the subdivisions merely colour and pattern groups. The names in brackets are those used by : * G. F. Hampson, Catalogue of Moths and other works, † L. W. Rothschild and K. Jordan, Revision of the Sphingidae. ‡ D. Sharp, Cambridge Natural History. § J. H. Durrant, Biologia Centrali-Americana and other works, or tabulated from other authors.
[Here Hampson attempts to rationalise the formation of family-group names by relating them to the first described genus in each family. Flying in the face of the rules of priority, this system was never widely adopted, but the family Uzuchidae survived for some time in some American publications].
Family
|
Genus
|
Type
|
UZUCHIDAE2 § (Xyloryctidae)
|
Uzucha Wlk 1864
|
humeralis
|
2 Cryptophag(s)a McLeay 1805, type irrorata, (nec Cryptophagus Herbs., Col 1792) is the oldest name in the family. (Hampson, 1918).
[Hampson gives no reason for abandoning Cryptophasa as the nominative genus of the Uzuchidae, other than suggesting that Lewin’s claim to be the author is insecure. McLeay’s claim to be the author is controversial, possibly being a nomen nudum. Meyrick’s subsequent emendation had been long abandoned by 1918. The earliest Xyloryctine generic name after Cryptophasa is in fact Boydia Newman, 1856.]
[Hampson also includes here a key to Lepidopteran families, complied by J.H. Durrant, from which the following information on Xyloryctine characteristics can be extracted]:
Antennae not clubbed or dilated or frenulum present when clubbed or dilated. Hindwing with vein 1.c present. (* Sometimes absent by asthenogenesis in some genera of the Eucoamidae and the leaf-mining Microlepidoptera.) Wings not divided into plumes. (* Except in Cemloba and Oxychirola.) Hindwing with vein 8 remote from 7. Hindwing with vein 8 free, or connected with the cell by a bar. Middle spurs of hind tibiae, or at least one, well developed. Palpi with the 1st joint much shorter than the 2nd joint. Antennae not bipectinate in both sexes, or if bipectinate in female the forewing with veins 7 to 10 separate. Hindwing with vein 8 not closely approximated to the cell and vein 7 throughout. Hind tibiae without whorls of bristles or scales at origin of spurs, the tarsi without bristles at the apex of the joints. Palpi long, upcurved, the terminal joint acuminate at tip, usually acute (rudimentary in some Blastobasidae). Forewing with veins 7, 8 stalked or coincident. Hindwing with vein 8 connected with the cell by a bar. Hindwing with vein 5 approximated to 4. Hindwing with vein 6 present ; veins 6, 7 generally approximated or stalked, the termen usually sinuate or excised below apex. Forewing with vein 2 remote from 3.
(Durrant, in Hampson, 1918).
XYLORYCTIDAE
This extensive family is widely distributed also outside Australia, but Cryptophasa Mc Leay, and allies, comprising moths of conspicuously large and even gigantic size, are as characteristic for the fauna of that region as is Kanguru. Not less than 37 species of Cryptophasa occur in New Guinea and the Bismark Islands, but they are all endemic, except one (C. mesotoma Meyrick which is also recorded from the Moluccas) and are not known from the Australian continent.
Certain species, as C. pseudogramma Meyrick and C. curialis Meyrick are locally common and come in large numbers to lamp traps. Dr Toxopeus informed us that when caught these large insects behave in the way of a clothes moth and try to escape by quickly crawling around. When taken in the hand they scratch the collector with their strong thorny legs.
Nothing is known about the biology of the Papuan species, but most likely it will appear to 'be the same as in the allied Australian ones. The larvae of certain species in that country are known to bore tunnels in living trees, in which they bring leaves of their food plant and where they take refuge in daytime; after having closed the entrance of the tunnel with a barricade of silk they consume the leaves in peace.
The genus owes its name to this remarkable biology χρυπτος = concealed, ϕασος = eating), “Cryptophasa” being a misprint for ‘Cryptophaga”. The name has been emendated by Meyrick in 1890, but the emendation has been abandoned again, the generic name Cryptophaga being preoccupied for a genus in the Coleoptera.
Several genera of the Xyloryctidae, viz. Cryptophasa, Paralecta, and others, seem to be rather arbitrary; they show considerable variation as to the neuration, the structure of the male antennae and the length of the terminal segment of the labial palpi, all characteristics, which, otherwise, are of great taxonomic importance. This makes the discrimination of the genera very difficult at times, as can be seen from the following key, which is compiled to our best knowledge, but appears to us rather unsatisfactory. It is probable, that future study will reveal these variations in more genera, and consequently will enable us to re-diagnose them on a better and more natural basis. Our present knowledge obliges us to maintain not less than some 112 genera, many of which are based on a single species only.
The abdominal tergites in the two sexes are clothed with moderate or fine bristles which are arranged in transverse rows over posterior half of the tergites or along their posterior margins, and are directed caudad. These bristles are orange-coppery coloured, show through scale clothing and appear like more or less distinct transverse coppery or orange bands. Outside the Xyloryctidae such bristles are richly developed in the Schreckensteiniidae, and are also known in the Oecophoridae (Clarke), the Gelechiidae and the Cosmopterigidae. They are entirely absent in the Stenomidae.
The male genitalia are of a simple and uniform type, with only slight differences between the genera. Tegumen rather short, triangular. Uncus and gnathos both triangular, strong, porrect, together forming a pair of tongs, gnathos often embossed towards the apex above. Small lateral haired knobs below the base of the uncus represent the socii. Valva simple, elongate-ovate or pointed, thinly bristled, harpe mostly developed as a sclerotized continuation of the median part of the edge of the sacculus, running longitudinally over the disc of the valva and often forming a strong blunt hook; sacculus broad, folded over the valva. Vinculum strong, pointed or ovate. Juxta sclerotized, under a blunt angle to this fits a very strong anellus, which is often shaped as a large and bulbate sheet, rolled around the aedoeagus and enclosing this distally altogether. Aedoeagus is a strong, slender, narrow tube slightly and gradually tapering apically, telescoping within the juxta. A cornutus is seldom present.
The female genitalia are of the Oecophorid type. Ovipositor rather broad, erect, lobes simple ; it is extensile over the length of the eighth segment, which is mostly shorter than the ovipositor itself; postapophyses long. Ostium wide, seldom modified, limen mostly a broad weak transverse plate, emarginate in the middle, dorsal wall of ostium formed by one, two or three broad, densely haired lobes. Ductus bursae simple or with a moderate tubular colliculum in its proximal half, distal half sometimes spiraled. Bursa copulatrix ovoid, mostly simple.
Key to the Papuan genera of the Xyloryctidae
1. Fore wing with
a costal scale-projection before and beyond
middle, and appearing
to be excavate between
these projections.
Fore wing with veins 2 and 3 separate, 8 absent
. . . . . Acria STEPHENS
(Diakonoff, 1954)
MEYRICK'S classification [Meyrick, E., Proc. Linnean Soc. N.S. Wales, 5: 205, 1880] is based primarily on the venation of the wings of Lepidoptera supplemented by such other characters [Meyrick, E., Trans. Ent. Soc. London, 1883, pp. 119-31] as he considered important. He even went so far as to lay down principles [Meyrick, E., Ent. Mon. Mag., 25: 175-8, 1889] of classification based solely on venational characters and ridiculed the value of other structural features, particularly those of the genitalia.
At the outset Meyrick began with a false premise and as late as 1928 [Meyrick, E., A Revised Handbook of British Lepidoptera, p. 13, 1908] he made the following statement: “In my classification the standard aimed at is an average of about 10 species to the genus and 50 genera to the family in the world fauna; which corresponds pretty nearly to the numbers found in the genera and natural orders adopted by botanists for flowering plants, whose history is bound up with that of the Lepidoptera. Some monotypic genera are of course necessary; the largest existing genus is Stenoma, which contains at present over 600 species ....” (!) .
This was an attempt by Meyrick to employ a purely artificial hypothesis, based on a mathematical formula, to provide a convenient system to which he could fit his classification and one not based on natural phenomena and evolutionary development of the insects he proposed to study. As will be pointed out in the following pages his “genus” Stenoma is one of the largest conglomerations of unrelated species ever brought under one name in the Lepidoptera. ...
His almost fanatical reliance on venation as a means of classification, although admitting considerable variation, coupled with an equally determined refusal to consider the genitalic organs as a means of classification not only of species but also of higher categories, led Meyrick astray in many cases. In this connection he stated, [Meyrick, E., A Handbook of British Lepidoptera, p . 11, 1895.]
“Thus colour and outline, the hairs of the larvae, and the genital organs of the imagos are likely to be of slight importance in the definition of groups; whilst neuration occupies a high position, except when directly influenced by an alteration in form of wing, which is seldom the case."
Although Meyrick disdained the use of genitalia in classification he condescended to mention them on several occasions [Exotic Microlepidoptera, 3: 164, 1925; Trans. Ent. Soc. London, 1913, pp. 192-9, 1913; Trans. Ent. Soc. London, 76: 521, 1929] but then he gave only a superficial treatment of a few features which could be seen by denuding the uncus and ends of the harpes,and he never gave an illustration of one.
Regarding genera he says further, [Meyrick, E ., in litt., to August Busck: Thornhanger, 18.1.15.] “Genera which (according to Durrant) would be identical ‘under a lens’ but distinct ‘under a microscope’, are too fine for me. As one cannot convert all one's specimens into microscope slides . ... ”
This persistent refusal to examine the genitalia, in conjunction witth other structures, not only led Meyrick into error in delimiting families and genera but accounted in large part for his misdetermination of the sexes of many of his species. ...
The family XYLORYCTIDAE was proposed by Meyrick in 1890 [Trans. R. Soc. S. Australia, 13: 23, 1890] for a group of Australian genera and species characterised by a short cross-vein between the cell and vein 8 of the hindwing which the author, at that time, claimed to be “ ... invariably present.” As time went on, Meyrick broadened his concept of the family and included in it genera and species from many tropical and temperate regions of the world, and not only assigned many of his own genera to this family but included those of other authors. The thirty Meyrick genera treated in this work, assigned by him to the XYLORYCTIDAE, comprise a large assortment of types and representatives of several families. Of these genera only four, Amorbaea, Epichostis, Linoclostis and Nephantis, possess the crossvein between the cell and vein 8 on which Meyrick laid so much stress. Two other genera, Neospastis and Synchalara, have vein 8 almost anastomosing with the cell, indicating a tendency to the formation of a cross-vein, and in the genus Phthonerodes vein 8 fuses along the cell for more than one-third its length.
Amorbaea, Epichostis, Linoclostis and Nephantis form a group of closely allied genera in which there is no blind-sac in the aedeagus and the abdomen is spined. Epichostis stelota, however, is misplaced and is hereby transferred to the OECOPHORIDAE. In the hindwing of Nephantis veins 6 and 7 are stalked but in the other three genera veins 6 and 7 are well separated and divergent. In Phthonerodes the blind-sac of the aedeagus is absent but it is present in Synchalara and Neospastis.
The following genera have veins 6 and 7 of the hindwing stalked and have no crossvein between vein 8 and the cell: Aeolanthes, Amphitrias (synonym of Odites), Antithyra, Antolaea, Epimactis, Myriopleura (synonym of Odites), Procometis, Prothamnodes, Rhizosthenes, Thymiatris, Trichernis (synonym of Odites) and Trypherantis.
Aside from the two features mentioned above, there is little to recommend the close association of these genera and much less to support their relationship to XYLORYCTIDAE (s. str.). In the genus Antolaea the gnathos is divided into two spined knobs and the aedeagus has a blind-sac. The character of the divided spined gnathos is shared by the Meyrick species placed in the genus Acria Stephens, by the two species Durrantia montivola and D. flaccescens, and Odites navigatrix, but none of them has the blind-sac of the aedeagus. The spined gnathos is a common character in the OECOPHORIDAE and these species appear to be a development of that family. We cannot place them in the OECOPHORIDAE at present because of the stalking of veins 6 and 7 of the hindwing, and because we have no supporting evidence from larvae for such an association.
The genus Aeolanthes presents a peculiar but highly characteristic type of male genitalia in which the aedeagus is disproportionately large and lacks the blind-sac, the harpes are variously shaped and ornamented and always reduced, and the uncus assumes various shapes, but it is always present. The females generally have double signa of dentate plates but at least one species (sagulata) possesses no signum at all.
The species referred to the genus Odites or its synonyms conform very much to type in ·form of genitalia: the transtilla usually exhibits strong development of the lateral lobes. The females are diverse and, at present, do not give much help in delimiting the genus. The true species of Odites possess a peculiar type of sharply beaked gnathos, which is directed anterad, and a long narrow tegumen. In this genus Meyrick has placed such atypical species as brachyclista, concreta, continua, hermetica, mistharma, sphenodontis and spoliatrix which must ultimately be transferred elsewhere. Rhizosthenes possesses the same type of gnathos as Odites.
The genus Epimactis is extremely close to Odites, and indeed may be a synonym but, as veins 3 and 4 of the hindwing are united, I am maintaining its identity for the time being. Thymiatris and Trypherantis are very closely related, having symmetrical male genitalia with a semi-tubular anellus, broadly attached harpe with clasper, aedeagus without blind-sac and gnathos and uncus present.
Another group of genera, Deloryctis, Metathrinca and Ptochoryctis lack the crossvein between vein 8 and the cell in the hindwing, veins 6 and 7 are well separated and divergent and the abdomens are spined. These genera are probably nearer the OECOPHORIDAE than the. XYLORYCTIDAE.
(Gates Clark, 1955) [footnotes have been incorporated into the text].
The relationship between Stenomidae and Xyloryctidae has presented a difficult problem; Busck (1921a) transferred the genus Setiostoma Zeller from Glyphipterygidae to Stenomidae. Forbes (1923) evidently felt the two families were not distinct because he listed them as subfamilies under the family Xyloryctidae. Clarke (1955a) recognized them as separate families: the Xyloryctidae being confined principally to the Old World and the Stenomidae to the New World.
(Duckworth, 1964, p.25).
Family Stenomidae Meyrick
Stenomidae Meyrick (in part), 1906, Trans. Roy. Soc. South Australia, vol. 30, p. 50; 1909, Trans. Ent. Soc. London, p. 28; 1912, Trans. Ent. Soc. Loudon, p. 706; 1931, Anal. Mus. Nac. Hist. Nat. Buenos Aires, vol. 36, p. 378. — Walsingham, 1912, Lepidoptera-Heterocera, vol. 4 (vol. 42 in Godman and Salvin, Biologia Centrali-Americana), pp. 153-187; 1913, Lepidoptera-Heterocera, vol. 4 (vol. 42 in Godman and Salvin, Biologia Centrali-Americana), pp. 188-190.— Barnes and Busck, 1920, Contrib. Nat. Hist. Lepidop. North America, vol. 4, p. 236.
Stenomatidae Walsingham, 1907, Proc. U.S. Nat. Mus., vol 33, p. 214. Cryptolechiidae Meyrick (in part), 1883, Trans. Ent. Soc. London, p. 124.
Xyloryctidae Meyrick (in part), 1925-1934, Exotic Microlepidoptera, vols. 1-4.
—Forbes, 1923, Cornell Agric. Exp. Sta. Mem., vol. 68, p. 250.
Cryptophasidae Fletcher (in part), 1929, Mem. Dept. Agric. India, Ent. Ser., vol. 11, pp. 1-244.
Stenomides Meyrick, 1930, Ann. Naturhist. Mus. Wien, vol. 44, p. 233.
Stenominae Janse, 1932, The moths of South Africa, vol. 1, p. 61.
(Duckworth, 1964, p.25).
Fig 36.33C: Xylorycta, Xyloryctidae
Fig 36.31D: Xylorycta, Xyloryctidae
Xyloryctidae. Small to medium-sized; head (Fig. 36.33C) smooth-scaled; ocelli usually absent; antennae in male simple, ciliated, or pectinated, scape without pecten; maxillary palpi 3- or 4-segmented, usually folded over base of haustellum; labial palpi recurved; hind tibiae with long hair-scales; fore wing (fig. 36.31D) with R4 rarely to termen, CuA2 arising well before lower angle of discal cell, CuP present; hind wing often broader than fore wing, Sc+R1 usually separate from and diverging from Rs well before upper angle of discal cell, R1 sometimes present, Rs and M1 usually approximated, connate, or stalked, CuP present; abdomen usually with dorsal spining. Larva with crochets biordinal, in circle, or ellipse; feeding on lichens, tying leaves, feeding in shelter beneath bark, tunnelling in bark or stems of trees, often dragging leaves to the entrance for food.
The family is well developed in Australia, where species of up to 75mm wing expanse occur. The adults are seldom seen during the day, but come to light at night. Many are distinctively marked, and some are sexually dimorphic.
The maxillary palpi have 4 segments in most genera, including Lichenaula (56 spp.), Procometis (20 spp.), Telecrates (4 spp.), Scieropepla (17 spp.), Catoryctis (20 spp.), and Uzucha (2 spp.). The relationships of these genera are not yet understood. The larvae of some, such as L. choriodes Meyr. (Fig 36.32C) and L. lichenea Meyr., feed upon lichens growing on fences and rocks, sheltering in a gallery of silk and refuse particles. Procometis bears a pencil of long hair-scales on the costa of the hind wing in the male. The larvae of P. bisulcata Meyr. form a vertical tunnel in the soil, emerging at night through a flexible, silken, soil-encrusted tube to feed on terrestrial lichens, pieces of which are stored in a rounded chamber just beneath the soil surface. Scieropepla larvae tunnel in flower spikes, usually on Banksia, but those of S. typhicola Meyr. burrow amongst the seeds of Typha. Banksia flower spikes are also tunnelled by larvae of Chalarotona intabescens Meyr. Telecrates laetiorella (Walk.) (Plate 7, H) larvae form a webbing gallery and feed on the inner bark of Eucalyptus. A conspicuous gallery of silk and bark particles is constructed by the larva of Uzucha humeralis Walk. (Fig 36.32D) which feeds on the surface of the bark of Angophora and smooth-barked Eucalyptus. The adult has the base of the costa in the fore wing strongly arched, very short labial palpi, and smoothly scaled hind tibiae.
In Neodrepta (7 spp.) and Xylorycta (93 spp.) the maxillary palpi are 3-segmented, and the male antennae are ciliated. Many of the species are shining white. The larva of N. luteotactella (Walk.) lives either in a webbing shelter amongst twigs and leaves or in a short tunnel in a twig or the woody fruits of Proteaceae, including Banksia and Hakea, and is a pest of Macadamia. X. strigata (Lew.) is white with a broad fuscous longitudinal stripe on the fore wing. The larva tunnels in the branches of Banksia serrata and B. integrifolia, and of Lambertia formosa, feeding on leaves which it drags to the entrance of the tunnel. This habit is common in Cryptophasa (19 spp.), which contains the giants of the family, with maxillary palpi of 3-4 segments and male antennae usually bipectinate. C. rubescens Lew. (Fig 36.32E) tunnels in the stems of Acacia, covering the entrance with a web of silk, and feeding on leaves it drags to the tunnel. By contrast, C. melanostigma (Walk) feeds on the bark of many native and exotic trees, often ring-barking them. Its main native host is Acacia, but it attacks citrus, stone and pome fruits, figs, and ornamentals.
(Common, 1970).
Although both families [Xyloryctidae and Stenomatidae] were originally proposed by Edward Meyrick, his concepts concerning them fluctuated through the years primarily due to his steadfast refusal to utilize characters other than wing venation for his higher classification. This dependence on a single character source led to numerous misconceptions, particularly with the higher categories. As the venational characters he originally used to separate the xyloryctids and stenomids began to break down with the continuing discovery and description of new species, Meyrick (1915) merged the two families under the older name Xyloryctidae. From this point he proceeded during the remainder of his lifetime to describe hundreds of species from both the Old and New World tropics under the family Xyloryctidae.
Other workers, principally August Busck, continued to utilize the original two-family concept placing the New World species and genera in Stenomidae. This geographical distinction was further emphasized by Busck (1934) when in his Stenomidae portion of the Lepidopterorum Catalogus series he only included genera and species from the New World. Thus, through omission, the genera and species from areas other than North and South America remained largely unknown and, until the present, uncataloged. In addition, the Xyloryctidae (sensu stricto) have never been cataloged and remain essentially unstudied.
In recent years a number of papers have reflected the distinctiveness of the two families; however, due to the nature of the studies (faunistic, type catalog) they do not provide a sufficiently clear picture of the two families to serve as a basis for future studies. Clarke (1955a) provides a thorough account of Meyrick's treatment of xyloryctids and stenomids in his introductory material to the catalog of Meyrick types; however, his detailed treatment (1955b) of the type specimens does not include species from Australia where the majority of xyloryctids and Old World stenomids occur.
Diakonoff (1954) provided a complete and accurate definition of the two families based on his study of the Microlepidoptera of New Guinea. His descriptions coupled with those of Common (1970) based on the Australian fauna provide the most comprehensive characterization of the two families yet published. Both studies, however, are restricted in scope and the latter provides only fragmentary information below family level.
(Duckworth, 1974).
Subfamily XYLORYCTINAE (Meyrick), new status.
Xyloryctidae Meyrick, 1890a: 23.
Xylorictidae: Dyar, 1903a(l902) :518. Forbes, 1923:250. Brues and Melander,
1932,1945:228.
Cryptolechiidae Meyrick, 1883b: 124.
Cryptolechidae: Walsingham, 1891 : 100.
Cryptophasidae Kirby, 1897: 303.
Uzuchidae Hampson, 1918: 336.
Diakonoff, 1954b:89; key to New Guinea genera, p. 90.
This group is usually given family rank by lepidopterists, but it can hardly be considered more than a subfamily of the Gelechiidae.
The name Cryptophasidae has been used by those who unfortunately believe that the family name should be based upon the oldest generic name within the family instead of on the oldest family-group name. It is based upon Cryptophasa Lewin, 1805. Hampson, 1918, proposed the synonymous name Uzuchidae for a similar reason. Xylorycta Meyrick, l890a:57, is an Australian genus.
Walsingham, 1891:100, has explained how “Zeller's genus Cryptolechia falls into the family Oecophoridae of Meyrick, and annihilates Meyrick's family Cryptolechidae, which was not founded on the typical form. Mr. Meyrick, recognising this, has since recharacterised his family Cryptolechidae under the name Xyloryctidae . ... The error, for which Mr. Meyrick cannot rightly be held responsible, has evidently arisen through confusion which Zeller created by his attempts to expand and amplify his original work.” Additional details of importance will be found in Walsingham's discussion.
All the members of the Xyloryctinae in Hawaii are endemic. Walsingham divided them into five endemic “genera”: Thyrocopa, Catamempsis, Psychra, Ptychothrix, and Hodegia. Catamempsis, Psychra, and Ptychothrix were separated from Thyrocopa largely on the basis of sexual characters of the male antennae. I cannot agree that such characters of one sex can be used to maintain genera. Hodegia was erected to receive a single female specimen with reduced wings. We now know that both sexes are flightless, but the species is otherwise a typical Thyrocopa. I have examined all of the Hawaiian species, including their wing venations and genitalia, and I can find nothing to indicate that more than one genus in the process of rapid specific radiation is involved. I am, therefore, reducing all ofWalsingham's “generic” names to new synonyms of Thyrocopa Meyrick l883a.
Although my late friend August Busck had not examined the Hawaiian Xyloryctinae at the time, he had the following pertinent remarks to make in his 1908: 137-138 review of Walsingham's Fauna Hawaiiensis monograph.
“The writer regrets one single feature in this masterful work, namely, the erection of genera (Ptychothrix, Catamempsis) on secondary sexual characters alone, and this in spite of His Lordship's own statement in his remarks (page 738- 9), that such characters are of very doubtful value, and especially so in the Hawaiian fauna, where the most embarrassing plasticity of such characters prevails. Undoubtedly, other sounder structural characters, common to both sexes, could have been found , or if not, the genera are, in the writer's judgment, not justified. To him it seems essential, for a sound appreciation of the natural grouping of the Microlepidoptera, that we get away altogether from these superficial characters, however tempting, and rely solely on the more ... dependable ... modifications. ...
The greatest known development of the Xyloryctinae is in Australia where there are more than 400 species, but in nearby New Zealand the group is hardly represented. The subfamily is confined mostly to the Southern Hemisphere: Australia, South America, and southern Africa. There are none recorded from North America.
The abdominal tergites in this group have characteristic transverse bands of spines, as in figures 668 and 690. Except for Endrosis, Oecia, and Blastobasis no other group now known in Hawaii has transverse bands of spines, and the character is almost diagnostic of the subfamily in Hawaii. Often the spines are easily seen on dried specimens where they may appear golden in colour. Batrachedra also has spinose abdominal tergites, but its spines are placed in longitudinal rows. Vein 1c is preserved near the forewing margin in Hawaiian Xyloryctinae except evidently in the highly modified “Hodegia” apatela as it is in the Hawaiian representatives of the Ethmiinae and Endrosis in the Oecophorinae.
In his detailed work on the Australian gelechiids Meyrick 1904c:256 said: “Assuming that the Xyloryctidae are maintained as a distinct family (which still appears to me to be convenient, though I think ultimately it must be reduced to a group of the Gelechiadae), I rely for distinction mainly on the character of vein 2 of the forewings, which in that group rises widely remote from 3 (generally disproportionately so ...”
(Zimmerman, 1978).
Xyloryctinae Although this subfamily occurs in most regions, the species and genera are most numerous in Australia, where about 250 described species in 46 genera are recognized. In addition many undescribed species are known. However, unlike the Australian Oecophorinae in which there is a close association with Eucalyptus (Myrtaceae), especially with dead eucalypt leaves, the food plants of Xyloryctinae include many plant families, and only two species are known with larvae dependent on dead eucalypt leaves. The habits of the larvae also vary; many live between joined leaves, some constructing portable cases, others tunnel in the bark of living trees or tunnel in the stems, feeding either on the regrowth of bark around the edges of a vestibule excavated at the entrance to a tunnel, as in some Hepialidae, or use the tunnel as a shelter cutting off green leaves and attaching them at the tunnel entrance to be used as food. Most species feed on woody shrubs and trees, but a few feed on lichens. In Australia the Xyloryctinae vary enormously in size and include the largest gelechioids known, with a wingspan of over 70 mm.
The adults usually have 4-segmented maxillary palpi, which fold over the base of the scaled proboscis, but in a few species the number of segments may be reduced to three or two. The labial palpi are strongly recurved and usually long and slender, but occasionally extremely short. The antennae in males are frequently bipectinate, and the scape never has a pecten. In the fore wing, veins R4 and R5 are nearly always stalked, with R5 usually running to the termen but occasionally to the costa or apex. CuA2 usually arises well before the lower angle of the cell, and CuP is present. The hind wing is as broad as the fore wing or broader, with Rs and M1 approximated, connate or stalked from the upper angle of the cell; CuP is present. The abdomen bears areas of spines on terga 2 to 6, and often has a conspicuous area of orange scales dorsally on segment 2 and sometimes on other segments. The male genitalia of most species are characteristic, with a short, thick, heavily sclerotized uncus, joined laterally to a short, thick, upturned, heavily sclerotized gnathos beneath it. The valva is long and narrow with a curved apical process of the sacculus.
Many small species come to lights at night, but the early stages of most of them are unknown.
[Here Common estalishes a mistaken notion about the structure of the Xyloryctine clasper, which develops not from the apex of the sacculus but from the inner edge of the sacculus, sometimes from the vinculum, but is distinct from the apex of the sacculus, which sometimes has its own developments.]
(Common, 1990).
Minet (1990) derived 32 characters from larval, pupal and adult morphology, as well as larval behavior, to revise the family classification of Gelechioidea. The characters were not subjected to any rigorous analysis. These observations were largely used to redefine families, and inter-relationships of the families were only considered to a lesser extent. Minet (1990) somewhat stressed characters derived from immature stages, 20 of his 32 characters being from larvae and pupae. He recognized 17 families within Gelechioidea.
(Minet, 1990, described in Kaila, 2004).
Autostichinae (e.g., fig, 223), a small group, are Indo-Australian and include genera that had been placed mainly in Gelechiidae or Xyloryctinae (Hodges, 1978). The presence of spines arranged in transverse rows at the posterior edge of the abdominal terga of both autostichines and xyloryctine led Minet (1986) to treat Autoshichinae as Xyloryctinae.
Xyloryctinae are fairly widely distributed but occur mostly in Indo-Australia, where there are 250 described species (Common, 1990). Includes small to large moths, (e.g. Fig 223) External ocelli are absent, the antennae of males are ciliated or pectinate, and the scape lacks a pecten. Vein CuP is present in both wings, and Sc+R1 diverges from Rs well before the end of the cell in the hindwing. Tergal spines are typically present in transverse bands on the posterior region of each abdominal tergum. The crochets on the prolegs of the larva are biordinal and arranged in circle or an ellipse.
The larvae live concealed lives, but may emerge from a hidden position to feed on lichens. Galleries may be made of silk and debris, or they may be formed of silk and soil. Some species tie leaves or bore into bark or the wood of branches. Whilst some species actually feed on bark, others drag leaves to their burrows. Xyloryctinae feed on several families of plants, and a few are lichenivorous.
(Scoble, 1992).
Sinev (1993) promoted the use of morphofunctional character complexes to understand the evolutionary trends within the superfamily. The classification of Sinev (1993) was based on three morphofunctional complexes: the wing apparatus, the genital apparatus, and the ecdysial complex. This approach has its merits because certain characters tend to correlate with each other, and they probably functionally co-operate. Such character complexes, however, have appeared hard to operationalize in the identification of phylogenetic relationships because observations from a wide array of taxa do not unequivocally support the existence of fixed character complexes. In fact, Sinev (1993) himself listed numerous exceptions to the general patterns. He promoted the elevation of Gelechioidea as the infraorder Coleophoromorpha with six superfamilies. His Oecophoroidea contained the families Oecophoridae s.s., Xyloryctidae, Symmocidae, Chimabachidae and Autostichidae; Coleophoroidea contained Epimarptidae, Blastobasidae, Ashinagidae, Stathmopodidae, Batrachedridae, Momphidae and Coleophoridae; Elachistoidea contained Stenomatidae, Ethmiidae, Depressariidae, Elachistidae, Peleopodidae, Agonoxenidae and Blastodacnidae; Gelechioidea contained Lecithoceridae, Gelechiidae, Scythrididae and Metachandidae; Chrysopeleioidea consisted of Chrysopeleiidae, and Cosmopterigoidea of Scaesophidae, Diplosaridae and Cosmopterigidae. A total of 26 families with 45 subfamilies were recognized.
(Sinev, 1993, summarised in Kaila, 2004).
Oecophoridae (Xyloryctinae)
Small to large moths, 10-50 mm, which fold their wings tent-like, more or less rolled around the body, and rest parallel to the surface. Head shortened, scales smooth or semi-erect on top of head, face smooth-scaled; antenna 0.4 – 0.7 length of forewing, filiform, pectinate. or with short dense ventral cilia, usually without pecten; ocelli absent; chaetosemata absent; labial palpi long, slender and recurved, second segment rarely with brush of elongate scales; maxillary palpi folded over or lying along base of proboscis, or vestigial. Proboscis long and coiled, scaled basally, rarely reduced. Wings broad, fringes shorter than width of wing; forewing with vein R5 to termen, in many SE Asian species predominantly glossy white, but in some species dull-coloured. Legs with long hair-like semi-erect scales on dorsal surface of hind tibia. Abdomen with patches of dorsal spines.
Compare Oecophorinae (often with antennal pecten; narrower-winged; R5 to costa; abdomen usually without dorsal spines.
Larva with a varied biology, as Oecophorinae. Pupa not protruded prior to emergence of adult.
110+ species known from SE Asia: all areas. Lowland to montane. Attracted to light.
(Robinson, Tuck, & Shaffer, 1994).
Xyloryctidae. Sister-group to clades 11-36. Defined by 3 apomorphies: 10 larval segments A1-8 with pinaculum ring around SD1; 2) a pore posterad/ventroposterad of SD1; 3) segments 3-7 with secondary SV setae. Worldwide: 86 genera, 1200 + species.
Xyloryctinae (fig. 9.2.I). Within Xyloryctidae possess autapomorphy, abdomen with band of spiniform setae on posterior part of terga 2-6. ♂ antenna simple, ciliate, bipectinate. Larva: head with bordered submental plate (a submental pit in some Cryptophasa), subgenal sclerite triangular, short, stemmata with gap between ½, 4/5, 5/6; A1 with 3 SV setae; A3-6 with 5-7 SV setae. Pupa: abdominal terga often with crenulate raised ridge near anterior margins, becoming spined in some species; metathoracic legs exposed distally. Wings reduced in Thyracopa apatela. Larval hosts 21 plant families (47% Protaceae + Myrtaceae), some on lichens. Subsaharan Africa, Indoaustralia, Polynesia: 60 + genera, 500 + species (Cryptophasa, Lichenaula, Metathrinca, Ommatothelxis, Pansepta, Phthonerodes, Scieropepla, Telecrates, Uzucha, Xylorycta).
Common (1900) Duckworth (1973), Hodges (1978), Moriuti (1982), Powell (1980), Zimmerman (1978).
(Hodges, 1998).
Xyloryctidae, Meyrick 1890: 23, revised status, new concept
Xyloryctinae Meyrick 1890, 23
Cryptophasidae Kirby 1897, 303
Uzuchidae Hampson 1918, 386
Scythridinae Rebel 1901: 179, new synonymy, revised status
Butalidae Heinemann & Wocke [1876]: 436 (nomenclaturally invalid)
[Scythrididae becomes a subfamily of the Xyloryctidae.]
(Hodges, 1988)
Hodges (1998) implemented a parsimony analysis with 38 characters coded for a hypothetized ancestor and 37 suprageneric terminal gelechioid taxa. A majority of the characters were derived from adults. Of these, 11 characters were derived from male genitalia and 14 from wing characteristics, especially from venation. Gelechioidea comprised 15 families with 37 constituent subfamilies. This work is the only one to follow the principles of phylogenetic systematics and consider the whole Gelechioidea. The methodology adopted by Hodges (1998) is, however, problematic. First, the use of a hypothetized ancestor for rooting the cladogram is questionable as there is little knowledge of the monophyly or sister groups of the group under study. This practice artificially dictates the result of the study by forcing the target group to be monophyletic, and the polarities to be a priori ‘‘known’’, leaving them untested. Erroneous assumptions of monophyly can be detrimental to the resultant cladograms (Bininda-Emonds et al., 1998; Wiens, 1998). Second, the monophyly of the (sub)families used as terminals seems to have been taken for granted by Hodges (1998). The work of Hodges (1998) may also suffer from clustering independent characters as artificial character complexes that do not always reflect true observations. Such character clusters are, e.g., the gnathos (character 4 of Hodges, 1998); ankylosation of aedeagus (character 8 of Hodges, 1998) and the orientation and level of fusion of the forewing veins CuA1 and CuA2 (character 16 of Hodges, 1998).
(Hodges, 1998, summarised in Kaila, 2004).
Xyloryctidae
The family is defined on the basis of three larval autapomorphies, and brings together two groups that have previously (e.g. Common, 1990; Scoble, 1992; Robinson et al, 1994) been treated as a subfamily of the Oecophoridae (Xyloryctinae) and a distinct family (Scythrididae).
The Xyloryctinae have a band of spine-like setae on the posterior margins of most abdominal tergites, but no antennal pecten. Almost half the host records are from Protaceae and Myrtaceae according to Hodges (1998), but Common (1990) described a very diverse additional range of hosts, including lichens. The subfamily appears to be diverse in Malesia, especially in New Guinea, and more so in Australia. Many species have broad forewings.
The Scythridinae lack tergal spining but have a pecten on the antenna (though not according to Robinson et al, 1994) who discussed two species of Eretmocera).
The higher classification of the subfamily (as a family) is reviewed by Landry (1991) but is in need of further revision, especially the many species in ‘Scythris’. Eretmocera species have narrow wings, the forewings generally dark, occasionally with yellow spots, and there may be yellow bands or tufts in the abdomen. The larvae web flowers and leaves, or are leaf miners, exploiting a wide range of plant famlies.
(Holloway, 2001).
... the oecophorid lineage constituting the ‘‘autostichid’’ family assemblage (including taxa formerly assigned to Autostichinae, Holcopogoninae, Symmocinae, Glyphidoceridae and Lecithoceridae), Xyloryctidae s.l. (including a paraphyletic Xyloryctidae of authors, some oecophorids of authors, Deuterogoniinae and Blastobasinae), Oecophoridae s.s., Amphisbatidae s.s., Carcinidae, Stenomati[n/d]ae, Chimabachidae and Elachistidae (including Depressariinae s.s., Telechrysis, Ethmiinae, Hypertrophinae s.l., miscellaneous ‘‘amphisbatids’’ sensu authors, Aeolanthinae, Parametriotinae, Agonoxeninae and Elachistinae).”
[Xyloryctidae seen as a paraphyletic group incorporating some Oecophorids, Deuterogoniinae and Blastobasinae; Scythrididae removed from Xyloryctidae.]
Gelechioidea is a cosmopolitan, megadiverse radiation of Lepidoptera, belonging to ‘‘Microlepidopteran’’ Ditrysia. The superfamily comprises over 16,000 described species (Hodges, 1998) and innumerable undescribed ones. For example, according to Hodges (1998) only 10–40% of species are presently named in several species-rich gelechioidean families in the Nearctic region. The ratio is probably similar, or even worse, in other regions of the World, except in Europe and Russian Asia. For example, in the Subsaharan Africa, only the Gelechiidae
and Lecithoceridae of South Africa have been more thoroughly studied (Janse, 1949–54; Janse, 1958–63). Gelechioidea may be the least known superfamily of Lepidoptera (Hodges, 1998) and it may eventually be among the three largest lepidopteran radiations together with Pyraloidea and Noctuoidea.
The spectrum of lifestyles among gelechioid larvae is diverse. Among the taxa there are predators of other insects, scavengers, detritus and fungus feeders, although feeding on living plant material is prevailing. Species dependent on living plant tissue can be external feeders, borers or miners of any plant tissue: roots, stems, leaves, flowers or seeds (Powell et al., 1998). Larvae in several gelechioid lineages bear portable cases. Many of the external feeders conceal themselves within silky tents or webs, or by tying leaves. Some shelter in silky galleries in the soil from which they consume plant roots or attack leaf rosettes or other above-ground parts of plants.
A thorough entry to the history of the classification of Gelechioidea is provided by Hodges (1978). In modern literature, the superfamily Gelechioidea has been defined as moths having ascending recurved labial palpi, basally scaled haustellum and tineid-type thoraco-abdominal articulation (Common, 1970; Hodges, 1978; Common, 1990; Hodges, 1998). None of these characters is, however, unique to Gelechioidea, and only the basally scaled haustellum is universal within the superfamily. This character is, however, also present in Tischerioidea, Pyraloidea and Choreutoidea. Minet (1988) emphasized the significance of the distal invagination of pupal mesothoracic legs as defining the superfamily. This condition is prevailing although not universal within Gelechioidea. Therefore, morphological support for the monophyly of the Gelechioidea is rather weak, and remains to be tested by a rigorous analysis.
The family level classification of Gelechioidea has been hampered by the scarcity of obvious synapomorphies for any lineages. Narrow character sampling based on adult (Forbes, 1923; Bradley, 1972; Kuznetsov and Stekolnikov, 1979, 1984), or pupal (Mosher, 1916) characters, often combined with a regional approach, has led to widely differing hypotheses of inter-relationships and unstable classifications. The first, more thorough attempts to combine characters from the morphology of adult and at least to some extent from immature stages over the whole superfamily, on a world-wide basis were made by Minet (1990), Sinev (1993), and Hodges (1998).
Minet (1990) derived 32 characters from larval, pupal and adult morphology, as well as larval behavior, to revise the family classification of Gelechioidea. The characters were not subjected to any rigorous analysis. These observations were largely used to redefine families, and inter-relationships of the families were only considered to a lesser extent. Minet (1990) somewhat stressed characters derived from immature stages, 20 of his 32 characters being from larvae and pupae. He recognized 17 families within Gelechioidea.
Sinev (1993) promoted the use of ‘morphofunctional character complexes’ to understand the evolutionary trends within the superfamily. The classification of Sinev (1993) was based on three morphofunctional complexes: the wing apparatus, the genital apparatus, and the ecdysial complex. This approach has its merits because certain characters tend to correlate with each other, and they probably functionally co-operate. Such character complexes, however, have appeared hard to operationalize in the identification of phylogenetic relationships because observations from a wide array of taxa do not unequivocally support the existence of fixed character complexes. In fact, Sinev (1993) himself listed numerous exceptions to the general patterns. He promoted the elevation of Gelechioidea as the infraorder Coleophoromorpha
with six superfamilies. His Oecophoroidea contained the families Oecophoridae s.s., Xyloryctidae, Symmocidae, Chimabachidae and Autostichidae; Coleophoroidea contained Epimarptidae, Blastobasidae, Ashinagidae, Stathmopodidae, Batrachedridae, Momphidae and Coleophoridae; Elachistoidea contained Stenomatidae, Ethmiidae, Depressariidae, Elachistidae, Peleopodidae, Agonoxenidae and Blastodacnidae; Gelechioidea contained Lecithoceridae, Gelechiidae, Scythrididae and Metachandidae; Chrysopeleioidea consisted of Chrysopeleiidae, and Cosmopterigoidea of Scaesophidae, Diplosaridae and Cosmopterigidae. A total of 26 families with 45 subfamilies were recognized.
Hodges (1998) implemented a parsimony analysis with 38 characters coded for a hypothetized ancestor and 37 suprageneric terminal gelechioid taxa. A majority of the characters were derived from adults. Of these, 11 characters were derived from male genitalia and 14 from wing characteristics, especially from venation. Gelechioidea comprised 15 families with 37 constituent subfamilies. This work is the only one to follow the principles of phylogenetic systematics and consider the whole Gelechioidea. The methodology adopted by Hodges (1998) is, however, problematic. First, the use of a hypothetized ancestor for rooting the cladogram is questionable as there is little knowledge of the monophyly or sister groups of the group under study. This practice artificially dictates the result of the study by forcing the target group to be monophyletic, and the polarities to be a priori ‘‘known’’, leaving them untested. Erroneous assumptions of monophyly can be detrimental to the resultant cladograms (Bininda-Emonds et al., 1998; Wiens, 1998). Second, the monophyly of the (sub)families used as terminals seems to have been taken for granted by Hodges (1998). The work of Hodges (1998) may also suffer from clustering independent characters as artificial character complexes that do not always reflect true observations. Such character clusters are, e.g., the gnathos (character 4 of Hodges, 1998); ankylosation of aedeagus (character 8 of Hodges, 1998) and the orientation and level of fusion of the forewing veins CuA1 and CuA2 (character 16 of Hodges, 1998).
The aims of this investigation were: (1) to resolve relationships among the major clades within the superfamily Gelechioidea, and (2) to test the monophyly of the superfamily and its constituent (sub)families by implementing the ‘‘exemplar’’ approach (Yeates, 1995), and dividing character clusters into their units. In this way the above-mentioned pitfalls of the earlier published results were attempted to be avoided.
(Kaila, 2004).
In an outstanding recent contribution to the systematics of Gelechioidea, Kaila (2004) presented a cladistic analysis based on 187 morphological and 5 behavioural characters scored for 143 species representing almost the full range of recognised gelechioid lineages. Given his extensive use of characters from the immature stages as well as formerly neglected character-complexes from adults (e.g., internal thoracic structure), Kaila’s attempt to resolve relationships in the group must be regarded as the most thorough and objective to date. His results support a basal division of the superfamily into two major lineages, the ‘gelechiid lineage’ and the ‘oecophorid lineage’. The low consistency index for Kaila’s most parsimonious trees indicates the high incidence of convergent evolution amongst Gelechioidea, and whilst there is support for the monophyly of many of the lineages recognised, e.g., by Hodges (1998), a large number of groups lack unique defining apomorphies and are recovered only on the basis of homoplasious characters. Also, as pointed out by Kaila, some of the basal clades, especially in his ‘oecophorid lineage’, have relatively weak character support, and there is a lack of data for immature stages of some taxa in these groups. Because of these problems, Kaila did not recommend the use of his cladogram (Kaila 2004: fig. 1) to construct a new classification of Gelechioidea, but indicated how Hodges’ (1998) classification could be minimally rearranged to accommodate the revised hypothesis of relationships.
Kaila’s study suggests that Stathmopoda and related genera are closely allied to genera currently assigned to Batrachedridae and Coleophoridae in the ‘gelechiid lineage’. The support for this relationship is strong: four synapomorphies, two of them unique, are shared by Stathmopoda with Idioglossa, Batrachedra, Goniodoma, and Coleophora (Kaila 2004). I accept this here as sufficient evidence for the removal of Stathmopodinae from Oecophoridae.
Taxa formerly assigned to Oecophorinae in New Zealand appear to belong to two distinct groups. On the basis of Kaila’s cladogram (Kaila 2004: fig. 1), Tingena (and related genera: see below) would be referable to Oecophoridae sensu stricto, whilst Hierodoris, Izatha, and Phaeosaces (and related genera: see below) would be assigned to the ‘xyloryctid assemblage’, with Hierodoris occupying a basal position as sister to all remaining taxa in this clade. However, as Kaila emphasises, character support at the base of the ‘oecophorid lineage’ is weak, and the inclusion in the analysis of the Russian ‘oecophorine’ Martyringa ussuriella Lvovsky collapses the basal resolution in this large clade. Likewise, one of the apomorphies supporting monophyly of the sister-group of the ‘xyloryctid assemblage’ in Kaila’s cladogram (i.e. Oecophoridae s.s. + Amphisbatidae s.s. + Carcinidae + Stenomatidae + Chimabachidae + Elachistidae s.l.) is ‘mesal part of gnathos scobinate with small thorns’; this characterstate also occurs in Hierodoris, but not in the species sampled by Kaila. So although there is evidence for assigning Hierodoris and its relatives to an expanded ‘Xyloryctidae’, the phylogenetic position of these genera remains potentially unstable and sensitive to taxon sampling. Therefore, for the present I have adopted a conservative approach, retaining in Oecophoridae all New Zealand taxa assigned to Oecophorinae by Dugdale (1988) except for Compsistis bifaciella Walker, which is placed in Lecithoceridae (Dugdale 1996). However, following the approach of Common (1994; 1997; 2000), I here assign the New Zealand Oecophoridae to informal genus groups and erect the ‘Hierodoris group’ (see below for definition) to accommodate the basal genera of Kaila’s ‘xyloryctid assemblage’. All New Zealand taxa not assigned to the Hierodoris group are collectively placed in ‘Oecophoridae sensu stricto’, and divided between Common’s named groups. The assembly of these groups into a stable higher classification should await further phylogenetic analysis of the basal taxa in the ‘oecophorid lineage’ (cf. Kaila 2004: 324).
The following Australian genera are tentatively assigned to the Hierodoris group: Scieropepla (14 described species, including 1, S. typhicola Meyrick, shared with New Zealand), Nemotyla (1 Tasmanian alpine species, N. oribates Nielsen, McQuillan & Common) and Athrotaxivora (1 Tasmanian species, A. tasmanica McQuillan). The inclusion of Scieropepla and Nemotyla is based on their position in Kaila’s (2004) cladogram, where together with Izatha + Phaeosaces they form a monophyletic group basal to the core Xyloryctinae + Blastobasinae. Athrotaxivora was provisionally associated with Xyloryctinae by McQuillan (1998), who noted however that it lacked characters of the core Xyloryctinae. In characters illustrated by McQuillan (loc. cit.) it matches the diagnosis of the Hierodoris group given here.
No claim is made here for the monophyly of the Hierodoris group, which is intended as a convenient informal association of genera that fall outside the core Oecophoridae and Xyloryctidae, whilst sharing some characters with each of these groups. Indeed, Kaila’s (2004) cladogram would indicate that the Hierodoris group is not monophyletic. However, it is possible that denser taxon sampling in this region of the phylogeny, and a better coverage of immature stages, might recover a monophyletic Hierodoris group.
Relationships of the Hierodoris group
As indicated above, the Hierodoris group appears to be related to taxa traditionally assigned to Xyloryctidae / Xyloryctinae (Kaila 2004). Probably the most convincing character supporting this relationship is the presence of paired sclerotised slits in the larval mentum: outside the Hierodoris group and Xyloryctinae, these slits are known only from a few Lecithoceridae. However, because some more derived taxa in Kaila’s ‘xyloryctid assemblage’ (Uzucha humeralis Walker and Blastobasinae) lack these slits, and because there are no data for the larvae of other taxa, their presence is not recovered as an unambiguous apomorphy supporting the monophyly of the assemblage (L. Kaila, pers. comm.). The second character linking the Hierodoris group to the xyloryctid assemblage in Kaila’s phylogeny is the presence of a pinacular ring around the abdominal SD setae of larvae. This is again paralleled in Lecithoceridae, and in some species of Scythris and Stathmopoda; Hodges (1998) also listed it as an apomorphy of Autostichinae (Autostichidae).
(Hoare, 2005).
Xyloryctinae
This subfamily can be defined by the autapomorphy of the abdomen having a band of spiniform setae on the posterior areas of terga 2-6. In addition, the antenna lacks a pecten and in males, is ciliate or pectinate, a frenulum is absent, the gnathos is fused with the tegumen laterally, the juxta is present, and the forewing has CuP present. The pupa has abdominal terga with a crenulate ridge near the anterior margins, becoming spined in some species. Two species of Thyrocopa, endemic to Hawaii, are wingless and have a jumping behavior (Medeiros, 2008).
Xyloryctinae includes more than 500 species in 60+ genera, with the highest diversity in the Indo-Australian Region, but with species also occurring in sub-saharan Africa and Polynesia (Scoble, 1992; Hodges, 1998).
Larvae feed on hosts in 21 plant families, including lichens, but almost half of the known hosts are species in Proteaceae and Myrtaceae (Hodges, 1998). Larvae live in concealed shelters or galleries made of silk and debris or soil. Larvae of some species tie leaves or bore into bark or wood of branches, and others drag leaves into their burrows (Scoble, 1992).
References: Common (1990), Duckworth (1973), Hodges (1978), Medeiros (2008), Moriuti (1982), Powell (1980), Zimmerman (1978).
(Brown, Bucheli, and Lee, 2006).
Xyloryctidae • small to large • usually smooth head • wings held roof-wise • antennae simple or pectinate (feathery) in male, held back along leading edge of wing • palpi upturned, sickle-shaped, sharp-pointed • hindwing broadly lanceolate
This is a diverse and widely distributed family in Australia. Most are small moths, but many are giants with wing-spans up to 75 mm. The larvae of many species tunnel in stems, and some form silken tunnels in lichens or in clusters of leaves. The genus Cryptophasa contains many large species that typically bore in the trunks of trees. They leave their tunnel to cut leaves from the tree, which they drag back and tie with silk to the entrance of the tunnel and then feed for the next few days. There are about 350 species in Australia, and 1200 worldwide.
(Zborowski and Edwards, 2007).