The system of the Catocalinae — a historical survey

The present paper reviews the family-group names of the Catocalinae subfamily within the Noctuidae (Lepidoptera). The underlying concepts and their historical evolution to date are described. All known family group names which were assigned within the Catocalinae are listed. New status: Anumet[ini] Wilt­ shire, 1976 stat. rev., new synonyms: Lagopter[idae] Kirby, 1897 syn. no?, of Dvsgoni[idae] Moore, [1885] 1884-7; Lvgephilfini] Wiltshire, 1976 syn. nov. of Toxocamp[idae] Guenee, 1852; Mocis[ini] Berio, 1992 syn. nov. of Remigi[idae] Guenee, 1852; Pangrapt[inae] Grote, 1882 syn. nov. of Focillfidae] Guenee, 1852; Phaeocvmfini] Grote, 1890a syn. nov. of Omopter[idae] Grote, 1895.

Introduction "All natural bodies share with one another a greater or a lesser affinity; the sum of all these affinities forms the epitome of a natural system in abstracto. The proposal of a system in this sense is impossible for the human spirit, in that man is not able to fully investigate the origin and constitutuion of all natural bodies in all their relationships to one another. A list of the natural bodies in accordance with the manner in which they appear to, and are rec ognised by, the human spirit forms a system in concreto. In abstracto there can be only one system of nature; in concreto there can be as many as the human spirit can judge and interpret. All systems formed in this way are therefore only attempts to investigate the one, unalterable law of nature" (Herrich-Schaeffer 1843).
["Alle Naturkörper stehen zu einander in grösserer oder geringerer Verwandschaft; die Summe aller dieser Verwandschaften bildet den Inbegriff eines Systemes der Natur in Abstracto. Die Aufstellung eines Systemes in diesem Sinne ist für den menschlichen Geist unmöglich, indem er nicht befähigt ist, alle Naturkörper in allen Beziehungen zueinander, alle in ihrem Würden und Seyn vollständig zu erforschen. Eine Zusammenstellung der Naturkörper gemäss der Art und Weise, wie sie dem menschlichen Geiste erscheinen und von ihm erkannt werden, gibt ein System in Concreto. In Abstracto kann es also nur ein System der Natur geben; in Concreto so viele, als der menschliche Geist die Natur auf verschiedene Weise auffassen und beurtheilen kann. Alle auf diese Art gebildete Systeme sind also nur Versuch, das eine und unabänderliche Gesetz der Natur zu erforschen" (Herrich-Schäffer, 1843).] The lepidopterists working systematically were always eager to represent the system in his "natural" arrangement even if they did not know, or could not formulate, "the one, unalterable law of nature", i.e. mutation recombination and selection as driving forces of evolution. The term 'affinity' was at this time abstract. It was based on similarity alone and not on descendence. For this reason earlier systems could not claim to be constructed exclusively from monophyletic groups. Wing pattern, wing form and position initially stood in the foreground as characters (Borkhausen 1792). Subsequently morphological and anatomical features were increasingly adopted as arguments for taxa and their classification. The importance of the praeimaginal stages was recognised and used at an early stage (see e.g. Denis & Schiffermüller 1775). The diversity of knowledge and the different opinions of several lepidopterists resulted in different systematic models which are still, in part, not monophyletic and which are thus impossible to falsify. Kitching (1984) vividly demonstrates the historical evolution of schemes for the family Noctuidae. To date around 35,000 species have been described within the family Noctuidae (Yela & Kitching 1999), arranged in about 30 subfamilies (Nye 1975, Poole 1989, Fibiger 1990, Heppner 1991, Speidel & al. 1996a). About 7,000 species with approximately 10,000 described taxa are currently assigned to the subfamily Catocalinae. The systematics and phylogeny of this subfamily represents an unresolved problem (e.g. Kitching 1984; Mitchell & al. 2000). The principal reasons for this are the historical evolution of the systematics of this subfamily, the enormous species abundance, the insufficiently known morphological variety, the predominant distribution of the genera and species in the tropical and subtropical region as well as the lack of specialists able to survey this group on a world-wide basis. It is safe to assume that the Catocalinae are not a monophylum in their current delimitation. The most practical approach for a long-term solution of this problem is to distinguish uniform (i.e. monophyletic) groups within the Catocalinae -including all taxa known world wide-and to examine these separately (Kitching 1984, Berio 1992, Speidel & Naumann 1995. Only after this can hypotheses be developed for the kinship of these groups in relation to each other and/or a systematic scheme for the Catocalinae. For the delimitation of uniform groups numerous older and younger models exist which need to be taken into account. In this paper the changing historical evolution is presented and the availability of all known family group names is checked.

M eth o do lo gy
The term Catocalinae is used here in the current sense which includes the so-called Ophiderinae (Noctuinae sensu Hampson). All taxa at the family-group rank as well as the mentioned genera for the Catocalinae were extracted from die relevant literature. The original spelling and original grouping was adopted. As a consequence of the successive development of the systematics with synonyms and changing interpretations it is difficult to compare all opinions concerning the arrangement of the genus groups by various authors directly. Therefore the current systematic position of the mentioned species in all publications has been checked. The work of Poole (1989) and Nye (1975) formed the basis for this. If the present species assignment differs from that in the original work, then the current one is added in square brackets [ ]. In case of assignment to another genus, subfamily or family, this is indicated by round brackets ( ). The few species of the Catocalinae in the Holarctic region represent only a small section of this world-wide group. Therefore the many travelogues, faunal lists, etc. which include a grouping of the species into suprageneric units from this region without justification were not analysed in so far as no new family-group names are proposed here.

H istory
The Catocalinae species placed initially by Linné in the genus Noctua were arranged by Hübner (1816 ff.) in his list into numerous genera. This list forms the basis of our present systematics on a world-wide basis. He consistently employed the taxonomic units genus '(Coitus/Verein), tribus (Familia) and subfamily (Stirps/Stamm) (cf. Hemming 1937: 16). The species of the Catocalinae were arranged by Hübner into 9 subfamilies (table 1). On the other hand Boisduval (1829Boisduval ( , 1833 distinguished only two major subunits, the Catocalides (Ophideres, Ophiusa) and the Homopterides (Polydesma, Cyligramma, Erebus). The basic idea of Hübner (1816), to consolidate similar genera into groups was consistently continued by Guenée in the following years. Guenée (1837: 321) initially used the tribus names Catocalidi, Ophiusidi, Noctuoidi and Noctuo-Phaloenidi in his survey of the European Noctuidae. However, he later (Guenée 1841) added further tribus names and their definitions (table 2). In his following comprehensive study of all known Lepidoptera he significantly revised the system of the Catocalinae (Guenée 1852a(Guenée ,b, 1854 and arranged the known genera into numerous groups. Walker (1857-1858) adopted this system in its entirety. He added the newer species only (in part with new generic terms) and revised the approach at the genus level (table 3).
The systematic studies of Duponchel (1844), Herrich-Schäffer (1843, 1845, 1847, Packard (1869), Meyrick (1895), Tutt (1896), Haworth (1803, Lederer (1857) and Smith (1893) predominantly considered the species of Europe and/or North America. The species of the Catocalinae were mostly put only into one family-group because of their small number of species occurring in these areas. As mentioned already, this geographically limited consideration is not suitable for carrying out systematic groupings. This "nordic" viewpoint -to put the Catocalinae into a single subfamily -obviously domi nated the systematic thinking and strongly influenced the subsequent development of ideas. A.R. Grote formed in North America an exception to this general tendency. He endeav oured to distinguish family groups and name them. His opinions were of course subject to changes over time. Since his family-group names are today frequently misquoted, we express here the relevant contents of his papers; publications which are often difficult to come by. Fundamentally he subdivided the Noctuae (as a family) into Noctuelitae fasciatae (includes mostly the quadrifine Noctuidae) and Noctuelitae nonfasciatae (all trifine Noctuidae as well as the quadrifine groups mentioned in the table) (Grote 1882, table 4).      In his following papers Grote increasingly refined and modified his system (Grote 1883(Grote ,1889(Grote ,1890a(Grote , 1890b(Grote , 1895(Grote ,1896a. (Grote 1890a, (1897) a few years later. At the time of these publications nobody presumably suspected that they would remain the last studies containing detailed subdivisions of the Catocalinae for almost eight decades. Kirby pub lished in 1897 a well-illustrated survey still using Guenee's system, in which he drew a small number of typical representatives for every genus. He initially separated out the Ophiderinae from the quadrifine noctuids and subdivided them strongly (table 10). Why he failed to continue this concept further in his later study (Kirby 1907) remains unex plained. His reasons are unknown and must be a subject for speculation. About at the same time in the German-speaking region -then a centre of Palaearctic lepidopterology -there was a struggle over guiding principals with respect to systematic work. Rebel (1898) pointed out the necessity' of phylogentic systematics as the basis for a modern system of the Lepidoptera. In his opinion the selection of the morphological features used was decisive and the forming of relationships based on mere similarities was impossible. He hoped this principle would take precedence over the work of Staudinger during the preparation of their joint catalogue (Staudinger & Rebel 1901). Apparently in the end he only succeeded in part, since the system of the preceding cata logue (Staudinger, in Staudinger & Wocke 1871) was retained to a large extent. In this widely distributed catalogue an already out of date system was used in particular for the subfamily Catocalinae and was thus presumably cemented as a standard.   Catocalinae-species remained in the great block of the quadrifine noctuids. He missed a great opportunity to refine die previous systems during his preparation of the catalogue. He put properly separated taxa -for example die Calpidae -back into the Quadiifinae. Since Hampson was regarded as an audiority and a special expert on die Lepidoptera these simplifications, although already understood to be steps backward, were generally accepted. Leech (1900) basically followed Hampson and still distinguished only die Quadiifinae and Focillinae.
In his studies of die fauna of Africa Hampson (1902) evolved his system and separated die present Catocalinae into two subfamilies. The first subfamily -the Catocalinae sensu strictohe first designated as Homopterinae (Nyctipao, Homoptera, Aiiclea ... Ophiusa), then later (Hampson 1910) as Catocalinae (Cyligramma, CaUiodes, Ophiusa, Ctemtsa, Grammodes, Attatha, Remigid). He arranged the "remains" into the group of his so-called Noctuinae (= Ophiderinae). He characterised his subfamily Catocalinae by a fully-developed vein 5 (M2) in the hindwing which originates from the proximity of the lower discoidal cell and the spined tibia. He consolidated this view for the Catocalinae in his subsequent world-wide account of the Catocalinae (Hampson 1913). He separated the Noctuinae (= Ophiderinae) by the main distinguishing character "without spined middle tibia" from the Catocalinae "with spined middle tibiae". Prout (1921) and Richards (1933) later showed that this feature leads to an artificial group and cannot be employed. Prout considered allied genera, for example CoQ'todes/Arete or Achaea/Mimophisma, as closely related on the basis of several features, although according to the "spined tibia" character of Hampson they would have had to use the different subfamilies of Catocalinae and/or Noctuinae (= Ophiderinae). The output of Hampson with respect to the svstematics of the Noctuidae is surely undisputed, but for the Catocalinae Hampson cannot be regarded as the precursor of the subject (Hitching 1984). Rather he welded together two monolithic, probably polyphvletic groups or 'grades'; heterogeneous groups whose separation soon proved to be untenable. His methods have been of great advantages for identification, however the main groups he created are not natural ones.
In spite of some criticisms of the features employed for Hampson's groupings -for instance by Forbes (1914) who criticised the presence/absence of vein M2 in the hindwing since intermediate forms exist with an already reduced M2 or by Prout (1921), Gaede (1938) and Draudt (1939) who stated "delimitation of the Catocalinae is unsure, [and the] feature of spined middle tibia insignificant" -his divisions were largely followed (Dyar 1902, Meyrick 1912, Janse 1917, Mosher 1916, Barnes & Benjamin 1923, Lhomme 1923-35, Bang-Haas 1926. They were particularly carried on in the famous publication of Seitz (Warren 1913, Gaede 1936, Draudt 1935, Draudt & Gaede 1944). This division is mostly employed even now in summarised systematic or faunistic works (McDunnough 1938, Zimmerman 1958, Common 1968, Pinhey 1975, Leraut 1980, Inoue & al. 1982, Franclement & Todd in Hodges 1983, Poole 1989, Hacker 1990. Here allied groups are often separated by means of the feature "tibiae with or without spines" and thus put into either the Catocalinae or Ophiderinae. Berio ( , 1992, Hitching (1984), Speidel & al. (1996a), Hitching & Rawlins (1999, and Yela & Hitching (1999) explicitly showed that this procedure does not form any sort of suitable basis for the formulation of phylogenetic systematic hypotheses for the Catocalinae. Some authors reject completely this insupportable division and take all groups together as the Catocalinae (Hodges & al. 1983, Common 1990, Sugi 1992, Kobes 1985, Kononenko 1990. Hitching & Rawlins (1999) also provisionally reject any subdivision of the polyphvletic Catocalinae since the delimitation, composition and monophvly of most of their constituent groups is uncertain. They removed only the Calpinae (= Ophiderinae sensu stricto of other authors) from which they recognised the tribes Calpini, Gonopterini and Anomiini. Yela & Kitching (1999) regarded the Gonopterinae (= Anomiinae) as a further independent subfamily. The widespread use of Hampson's classification hinders the development of a con sistently phylogenetic system. A main reason for holding on to the old system is the complexity within the highly diverse Catocalinae, such that every attempt to resolve the systematics of this subfamily on a phylogenetic basis appears hopeless.

The new way
In spite of this dilemma there were repeated attempts to achieve progress in the sys tematics of the Catocalinae through the investigation of different features (in so far as the quadrifine noctuids were included). The inclusion of the tympanal-organs (Forbes 1916, Eggers 1919, 1925, Richards 1933 or larval features (Gardner 1947, Forbes 1954, Crumb 1956, Beck 1960 brought out different, but sometimes similar, results through which groups could be constructed. These results depended on the examined material and/or the investigated area.
On the basis of different features, illustrated by Boerner (1939), he later distinguished in his fauna of Germany (Boerner 1944(Boerner ,1949 again two subfamilies (table 12). Richards' (1933) investigations of the tympanal organ brought out 6 groups in the "Erebinae -Catocalinae complex" (table 11). Forbes (1954) also supported the need for a new model and determined a systematic division of the Catocalinae (table 13). As a result of this development the 150 year old ideas about the systematics of the Catocalinae were picked up again and pushed into a promising direction. One can already speak of a Renaissance when Berio ( , 1965 published a list of phylogenetic units within the Catocalinae. This important step inspired other researchers.  supported these opinions and he separated several tribes in a consistent continuation of this idea and designated them unfortunately only in part according to the rules of the ICZN. Kitching (1984) picked up this approach to the problem and proposed as an initial solution a first step of forming uniform (= monophyletic) genus-groups (= tribes) within the Catocalinae (incl. Ophiderinae) and a second step of examining these groups sepa rately and later integrating them into a coherent (= consistent-phylogenetic) system.          (table 14) and Berio (1992) (table 15) continued this further and ar ranged the related genera into tribes. Unfortunately Wiltshire neglected, in part, to give a definition and/or description for his new family-group names according to the ICZN which results that these proposed new family group names remain nomenclatorially unavailable. In order to avoid unnecessary confusion, we have judged the family-group names of Berio (1992) as available described by bibliographical reference to his corresponding "Phyla" in his former publications (Berio , 1965 in accordance with article 13.1.2 ICZN. The serious disadvantage of both these studies is, however, that for every newly described tribe no autapomorphy was designated and thus for these no reasonable grounds for their monophyly exist. But the system outlined in Berio seems to be a good basis which can be built upon due to the partlv world-wide consideration and because of the use of features which can be interpreted, in part, as apomorphies. Despite the problems, some of Berio's named tribes have begun to be used in faunistic studies (Speidel & Hassler 1989, Yela 1997

Perspectives
It is safe to assume that the taxa which are currently united in the Catocalinae (Kitching 1984) are polyphyletic. The adherence to the traditional systematics prevents further progress. Fundamentally necessary is a new way of thinking which rejects the traditional system of the Catocalinae/Ophiderinae in order to exclude "system-constrained" inter pretations of results. All available models must be subjected to critical analysis with regard to the features employed. Of course the currently known and employed features are not sufficient to develop a phylogenetic system at the present time. Additional apomorphic features must be sought which can establish supraspecific taxa. New molecular analyses suggest that the Catocalinae can be placed differently within the Noctuoidea (Fang & al. 2000, Mitchell & al. 2000. However, the main species block probably forms a monophyletic unit (Speidel & al. 1996a) and these authors proposed to summarise this block within the tribe Catocalini based on the autapomorphy of "proboscis tip with dorsal sensilla" and the putative autapomorphy of "existence of a corema at the middle tibia" (Barth 1957). The relationships between the individual tribes of the Catocalinae and an answer to the ques tion whether ultimately all tribes (or those except the Catocalini) will be able to remain in this subfamily must to be subject to further investigations.
Investigations which exclusively based themselves on the model of Hampson (1913) and/ or the following treatments in Seitz' encyclopedia (Warren 1913, Gaede 1936, Gaede 1938, Draudt 1935 or else consider only species for phylogenetic analysis already regarded as closely-related are unsuitable for making progress; as are those considering only the species from a specific area like, for example, . Furthermore it has proven necessary that systematic revisions of the respective group as a first step are nec essary in order to exclude the use of synonyms in the phylogenetic analysis. Furthermore, comparative morphological studies of all taxa of the Catocalinae are neces sary. In particular the highly diverse fauna of the tropical and subtropical regions must be included into these investigations. This extensive work can be achieved only by several scientists. An "atomization" into countless further tribes should be avoided completely, so long as the degree of affinity of the existing tribes and their possible synonymy remain undeter mined. Monophyla characterised by autapomorphies should not be designated until in tense investigation has been carried out, in order to avoid nomenclatural confusion in the long run (cf. Speidel & Hassler 1989).

Catocalinae/Erebinae -Ophiderinae/Othreinae/Noctuinae?
The diverse designations in the past for the same major group -whose monophyletic status has yet to be established -creates confusion. Most widely used is the name Catocalinae (Catocalidi Boisduval, 1829). Therefore we retain this name here as a nomen protectum. In fact the name Erebinae (Erebidae Leach, 1815) would have priority, but we choose the name Catocalinae as the designated one in accordance with article 23.9.1. of the ICZN to keep the stability (reversal of precedence), since, firstly, no author used Erebinae in priority as name for the family group instead of Catocalinae since 1899 and, second, the younger synonym is in general use. A distinction between Catocalini and Erebini (Nye 1975:11, Forbes 1954 also appears unnecessary since both groups agree closely in the structure of the proboscis. For the Ophiderinae -which were formerly separated from the Catocalinae because of their lack of spines on the middle tibia-three names are available: Ophiderinae (Ophideridae Guenee, 1852), Othreinae Berio, 1955 andNoctuinae (sensu Hampson 1910). The name Ophiderinae unambiguously has priority here and is also in general use. However, in the famous work of Seitz (Draudt 1935, Draudt & Gaede 1944) the name Noctuinae is generally employed for this group. Berio and Fletcher (1958) pro posed Othreinae (Ophiderinae = Noctuinae). This was based on their former opinion for the validity of the generic terms (cf. svstematics section). Wiltshire (1971) also still used the name Othreinae. The use of the name Noctuinae for this group is based on Hampson. He derived the family group names from the oldest generic name in the respective group. The type spe cies of the genera were always the ones which were listed first in the original description of the author (Hampson 1918). For instance Linnaeus (1758) first listed the species strix within the genus Noctua, such that Hampson (1918: 384) understood this as the type spe cies of Noctua and consequently of the Noctuinae. The description of the Phalaena strixby Linnaeus is based, however, on a mixture of two big moths: a south-American Noctuidae ( Thysaiiia agrippina Cramer, 1776) and a south-east Asian Cossidae (Xjleutes strix Linnaeus, 1758) (Fletcher &N ye 1982:170). Apparently Hampson interpreted Linnes description in the sense of the Noctuidae, not in the sense of the Cossidae as became later generally accepted. This procedure of Hampson's stands in contrast to present (and previous?) valid rules of nomenclature. The type species of Noctua is pronuba (official list of the generic names in zoology: name 1057). Therefore the name is employed correctly for the Agrotinae in the sense of Hampson, but this case is not considered here.

Systematics
In the following section all names of the family-group of the Catocalinae in a wider sense are listed and their taxonomic status is indicated. Names were taken into consideration in the list even if at least one author put them into the Catocalinae, although they undoubt edly represent independent subfamilies. This is valid for example for the Stictopterinae and Euteliinae which are included within the Catocalinae in some older studies. The main aim is to produce a sound basis for future investigations of the Catocalinae and for the determination of monophyletic groups. It was not possible to check all already existing names of the family group for their systematic status here. This would require an underlying, wrell-founded hypothesis as to the phylogenv of the entire subfamily and their environment. Only when this hypothesis has been developed can the higher systematics be corrected. There are likelv to be different concurrent models in which the supraspecific taxa will be not different in their components but the rank of these supraspecific taxa can be quite different and can only be determined due to historical continuity. In accordance with article 11.7. ICZN the name of a family group is formed based on the stem word of an available genus name. Article 29. ICZN regulates the determination of the stem word of the type genus and the use of suffixes. In the following list the correct spelling of the stem word stands at the beginning of line. Wrong or invalid spelling is indicated afterwards as "initial/original spelling". The priority status designating suffix of the respective author is separated from the stem word of the family group name by square brackets. An incorrect appendix of the suffix onto the stem word (e.g. i/ii) is corrected without comment. The name giving type genus ol the family group stands afterwards in square brackets.
The ending employed by the authors corresponds to their respective opinion as to the rank within the family group and is in accordance with article 29.2. with respect to the stem word. In this respect the different opinions of several authors to the rank of a taxa are reflected in the different endings. Taxa that are considered to be used in the same sense, but with another rank, are listed with a prefixed equals symbol (=) and the reference in which the another rank is assigned to the taxa is quoted with one prefixed " in ". Inspected references in which the taxa was employed in the same rank are stated in round brackets and -where necessary -explanatory remarks are added. Unavailable names (for example nomina nuda, homonyms) are characterised with a prefixed asterisk (*). Nomenclatorical changes carried out in this work are expressed in bold type. According to the ICZN (1999: 111) a nomen nudum is a name "that if published before 1931, fails to conform to Article 12; or, if published after 1930, fails to conform to Article 13." All new descriptions which were published after 1930 must satisfy the provisions of Article 13 of the ICZN and must be accompanied by a description or a definition in words or a bibliographic reference to such a published statement, or they must be proposed as a new replacement name for an available name. Designations assigned before 1930 are not subject to these orders. Here it satisfies in the case of the family group names that the formation of a family-group name is from an available generic name (indication). In accordance with article 11.7.1.1. of the ICZN the use of the stem is meant, then the author used the generic name as valid unless there is evidence to the contrary. These designations are valid without further description. All these orders are valid also for family group names, which some authors unfortunately did not consider (for example Wiltshire). Important for family group names is the fact that in accordance with article 50 a nomen nudum can be made available by a corresponding nomenclatorical action (for example a definition). In such cases, however, this active person is regarded then as the author with the date of his publication. Furthermore it should be noted that Berio (1992) and ) apparently simplv name family groups without giving descriptions, but definitions were given in part, however, in the former publications of Berio (Berio ,1965. We evaluate these cases as a correct nomenclatorical action (description by reference to a source) and consider the names as valid; the reference is referred to correspondingly.  (Berio 1992, Hacker 1999) Achaefini] : 160 [Acbaea Hübner, [1823], definition of the "Phylum di Achaea" in Berio 1960: 321, synonym of Catocalini (Speidel, Fänger & Naumann 1997), (Speidel & Hassler 1989, Berio 1992, Hacker 1999 Hübner [1823]], independent subfamily (Beck 1960, Speidel & al 1996a (Grote 1890b, Forbes 1954, Berio 1992, Hacker 1999, Hitching & Rawlins 1999, Hollowav & al. 2001 Speidel & Hassler (1989), , definition of the "Phylum di Anua" in Berio 1960: 315 Anumet[ini] :161 [■ \mundu Walker, 1858, stat. rev., previously treated as synonym of Svnedini by Speidel & Hassler 1989, (Berio 1992) Anydrophil[ini] : 161, [A/ydmpb/Ia J o h n , 1909, , Hacker 1999 Naumann 1995) Apopest [ini] Beck, 1996: 15 [Apopestes Hübner, [1823, The systematic position of this tribe is unclear, it is erroneously placed in the Cuculliinae by , whereas it is regarded as synonym of Toxocampini by    Staudinger, 1884], ,1979, Berio 1992, Hacker 1999 : 224 [Calesia Guenee, 1852, nomen nudum according to article 13.1 ICZN, (Hacker 1999 Guenee, 1852(Speidel & Naumann 1995, , Speidel & Hassler 1989, Hacker 1999) Tachosfini] Berio, 1992: 297, 300 [Tmvosa Walter, 1869 , Speidel & Hassler 1989, Hacker 1999 This list includes all family group names of Catocalinae s. 1. published until December, 2002, with the references to the first valid use of every name which we could find. There are no published catalogues containing these references, and so we may not have indicated the oldest reference in evert' case. We hope that minor errors are excusable in this first attempt and welcome every correction.