El último y extraordinario tomo de la serie Noctuidae Europaeae (vol. 13, Entomological Press), editado por Thomas Witt y László Ronkay, llegó ayer a mis manos. En él se incluye un capítulo de revisión sobre filogenia de noctuidos (sensu lato, como es necesario decir ahora, porque tal como han sido considerados habitualmente hasta ahora representaban un grupo parafilético). Puesto que soy el autor principal del texto, allá va tal cual ha aparecido, una vez corregidos dos errores menores detectados. Ni que decir tiene que he dedicado mucho tiempo y esfuerzo a escribirlo, aunque se trate de un trabajo despreciado en este país enfermo de impactolatría aguda crónica; mis contribuciones a Noctuidae Europaeae han sido vilipendiadas recurrentemente por los miembros las comisiones de valoración de los méritos investigadores, lo cual no deja de ser una excelsa muestra más del país de semianalfabetos funcionales en el que vivimos, abducidos hasta los tuétanos por la filosofía mercantilista.
Phylogenetic overview of Noctuidae sensu lato
José Luis Yela and Reza Zahiri
in cooperation with
Niklas Wahlberg, László Ronkay and Alberto Zilli
We present in this chapter a brief rationale of the history of the classification of noctuid moths, synthesising the available phylogenetic information in order to underpin the updated list of European Noctuidae sensu lato with a solid basis. We are perfectly aware of the difficulties in doing this. Translating the results of analytical phylogenetic procedures into cladograms is a well-established scientific routine. Translating them into formal classifications is less immediate, not least because classifying organisms is trying to categorise biological entities that are the results of a continuous, although admittedly not uniform, process, and thus classifications are to some degree subjective agreements. For this reason, we have tried to be as eclectic as possible, and have adopted the most recently published results on noctuid phylogeny although they may be partly in contradiction with our previous views, as expressed throughout this series. Also, we argue for the inclusion of lymantriine and arctiine moths into the noctuid (with quadrifid forewing venation) lineage and, hence, in this volume.
Latreille (1809) recognised for the first time noctuid moths as a taxonomic unit and proposed the family name. Guenée (1837-1841, 1852, 1854) did a thorough review of the available information (that in e.g. Ochsenheimer 1816, Treitschke 1825-1835 or Duponchel 1822-1838) and arranged noctuid moths in a large series of groups, defined mostly on the basis of adult external features, and acknowledged the existence of two main patterns of hind wing venation (trifine and quadrifine). The influential arrangement of Guenée was followed, in its streamlines, by e.g. Herrich-Schäffer (1843-1956) or Duponchel (1844-1846). Around 40 years later, Grote (1882) proposed a more elaborated system, which was rearranged again 20 to 40 years later by Hampson (1898-1920) in his monumental catalogue of the moths of the formerBritishMuseum. The renowned Hampsonian system was still mostly based on superficial characters of limited phylogenetic significance (Kitching, 1984; Yela & Kitching, 2000; Lafontaine & Fibiger, 2006); nevertheless, it was accepted as the main framework during more than half a century (e.g. Warren 1907-1914, Corti 1931-1932, Draudt 1932-1938, Boursin 1964). During that time (1920-1984), in spite of the relative stability of the system, almost every published classification of noctuid moths differed to some degree from the others, mainly in nomenclature and taxonomic order. Zimmerman (1958) stated, for example, that “the classification [of the family Noctuidae] rests in a state of great confusion, and few authors appear to hold similar views regarding the suprageneric taxonomy. I have concluded after spending much time (perhaps I should say wasting time) on the problem, that it is impossible to present …a correct suprageneric classification”; Nye (1975) wrote that “it is exceptional to find any two authors who use the same combination of subfamily names within the Noctuidae”. In 1984, moreover, a first cladistic approach to noctuid phylogeny was undertaken (Kitching, 1984), in a paper that represents the inflexion point towards an understanding of the evolutionary relationships of the noctuid lineages. Cladistic procedures, which group all descendants of a common ancestor to form monophyletic entities, provide the basic framework for hypothesis-driven phylogenetic studies (Hennig, 1966; Kitching et al., 2002; Williams & Knapp, 2010; see Fibiger & Hacker, 2007). Therefore, all subsequent attempts to reconstruct the phylogeny have been grounded on cladistic principles.
However, in order to perform an appropriate phylogenetic analysis, the group under discussion has to be adequately characterized and the relevant features of representatives of all lineages have to be well known. This is not the case of noctuids (Noctuidae sensu lato, as they have been regarded until recently), at least based on external morphological features. They comprise an evolutionarily relatively young, highly diversified group (Kitching & Rawlins 1998, Yela & Kitching 2000), the most speciose among Lepidoptera (around 35,000 describes species and many undescribed yet; Poole 1989; Kitching & Rawlins 1998; van Nieukerken et al., 2011), of which we still have, as a general rule and at a global scale, a rather superficial knowledge. In addition, within a general background of remarkable structural homogeneity, they show a noteworthy disparity, which is explained by each of the recognised phyletic lines differing only very slightly from the rest (in one or very few characters). The interpretation of this morphological disparity has been traditionally very difficult, mainly due to a high degree of homoplasy (similarity among phylogenetically unrelated lineages due to convergent evolution or non-adaptive reversal of character states) affecting most character systems (Poole 1995, Kitching & Rawlins 1998, Yela & Kitching 2000, Zahiri et al. 2012), for example the larval secondary hairiness on verrucae, the larval 2+2+2 chaetotactic condition of the thoracic subventral group of setae or the possession of raduloid on the larval hypopharynx (for details see Kitching & Rawlins 1998, Beck 1999, Yela & Kitching 2000 or Lafontaine & Fibiger 2006). Consequently, inasmuch as classification consists of the translation of the phylogenetic background to a hierarchically ordered register, a great instability took over noctuid classification following the seminal paper of Kitching (1984), as subsequent attempts to re-analyze or re-synthesize noctuid phylogeny incorporated much more morphological information from many more species of the whole taxonomic spectrum, from all stages of the biological cycle (Beck 1960, 1991, 1992; Lafontaine & Poole 1991, Poole 1995) and from all over the world (e. g., Lafontaine & Poole 1991, Poole 1995, Speidel & Naumann 1995, Speidel et al. 1996b, Kitching & Rawlins 1998, Yela 1998, Yela & Kitching 2000, Fibiger & Lafontaine 2005, Lafontaine & Fibiger 2006). Contrasting interpretations of noctuid phylogeny result from contrasting interpretations of the meaning of the distribution of character states across the noctuid tree.
The period of phylogenetic turmoil has corresponded fully with the production of the Noctuidae Europaeae series. The commotion is reflected in the changing views that appear in the different volumes about the phylogeny of noctuids. The first five volumes published (Fibiger 1990, 1993, 1997; Ronkay & Ronkay 1994, 1995) yet reflect the classical conception of Hampson and the traditional systematic order used by Boursin (1964), although right in the first volume there is a warning about the major changes that were beginning to take place. From the sixth on (Ronkay et al. 2001), these changes are noticeable in the very way the text is structured.
Given the difficulties in interpreting the distribution of morphological character states due to high rates of morphological homoplasy, one way to shed more light on our understanding of noctuid evolutionary relationships is switching from the phenotypic orientation to the genotypic one, which means using molecular techniques. The ultimate rationale is that if the phenotype (external appearance) is a result of the effects of genes (genotype) plus environmental determinants during development (plus interactions of both terms), if we are able to overcome the environmental effects by directly analysing the genetic background, we would get a more reliable picture of evolutionary relationships. This is largely true, but the limited number of character states in DNA leads also to large amounts of homoplasy that must be taken into account in molecular analyses (Mitchell et al. 2006, Regier et al. 2009, Zahiri et al. 2012). Fortunately, this is a property of DNA that has been intensively studied over several decades, and currently there are sophisticated statistical models that are able to estimate robust phylogenetic hypotheses despite the homoplastic nature of DNA data. Thus, molecular approaches have been successfully undertaken during the last decades (Weller et al. 1994, Mitchell et al. 1997, 2000, 2006, Fang et al. 2000, Zahiri et al. 2011, 2012), which have significantly helped to arrive at a reliable noctuid phylogeny and thus to provide a stable phylogenetic classification. This has been achieved not only by the examination of increasingly more and more genes, particularly nuclear ones, and by the study of an increasingly large number of species of most noctuid lineages (see Zahiri et al. 2011, 2012); it has also been owed to the mentioned refinement of analytical tools, which among other advantages routinely incorporate model-based (Bayesian inference and maximum likelihood) evolutionary methods (Mitchell et al. 2006, Zahiri et al. 2011, 2012), together with classical parsimony.
Fortunately, molecular taxonomists have paid special attention to the difficult and extremely speciose “quadrifine” noctuids (as traditionally defined, i.e. noctuids with quadrifid hind wing configuration), which have been extensively surveyed during the last years. The main conclusions and corroborations of these investigations, and the main subsequent re-interpretations of the topology of morphological evolutionary changes are as follows:
1. The superfamily Noctuoidea consists of two groups as generally defined by the forewing venation, the trifid and the quadrifid ones (see Lafontaine & Fibiger, 2006 or Zahiri et al. 2012 for definitions and figures). Trifid Noctuoidea are Oenosandridae and Notodontidae, and quadrifid Noctuoidea are a monophyletic assemblage of four evolutionary lineages that can be treated either as constituents of a comprehensive family (Noctuidae as interpreted by Lafontaine & Fibiger, 2006; monofamiliar scenario) or as four different families (Nolidae, Erebidae, Euteliinae and Noctuidae as interpreted by Lafontaine & Schmidt 2010 and Zahiri et al. 2011, 2012; quadrifamiliar scenario, followed here). The relationships between these four families are not resolved yet, and at present we may only tentatively hope to guess which is sister to the other (Zahiri et al. 2012). It may also be that the four families have diverged so rapidly from each other, that it will not be possible to ever establish their exact relationships.
2. Noctuidae sensu lato (in the traditional view) form a monophyletic group only if arctiine and lymantriine moths are included among them (Weller et al. 1994, Mitchell et al. 1997, 2000, 2006; Zahiri et al. 2011, 2012). Accordingly, Noctuidae sensu lato can be characterised by the quadrifid forewing venation (which is still a controversial feature, because there are a few known exceptions and there are members of the trifid Noctuoidea showing quadrifid forewing configurations; Schintlmeister pers. comm. in Fibiger et al. 2010) but also by the structure of the thoracic tympanum, which is directed posteriorly towards a counter-tympanal hood in the anterior part of the abdomen (Eggers 1919, 1925, Richards 1932, Brock 1971; Speidel et al. 1996b, Lafontaine & Fibiger 2006), and probably also by the presence of adenosma (larval ventral cervical gland, arranged “vertically”; Gardner 1941, Godfrey 1987), if we admit that it has been secondarily lost in Rivulinae, Arctiinae and some Hypeninae for the European fauna (Speidel et al. 1996b, Kitching & Rawlins 1998, Yela & Kitching 2000, Fibiger & Lafontaine 2005).
3. Consequently, arctiine and lymantriine moths have to be seen as highly modified Noctuidae sensu lato – in particular, Erebidae (the LAQ clade of Mitchell et al. 2006).
4. Nolidae, best characterised by the two-walled, boat-shaped cocoon (Gardner1948), may be sister to the other three families (Zahiri et al. 2012). Tentatively, it comprises for the European fauna the subfamilies Nolinae and Chloephorinae (with Chloephorini, Sarrothripini and Eariadini; Zahiri et al. in prep.). Nolidae lack cylindrical galeal lobes on the larval maxillary complex (Grimes & Neunzig 1986, Kitching & Rawlins 1998), which are present in Erebidae, Eutelidae and Noctuidae but arguably secondarily lost in Pantheinae and in Acronictinae (Kitching & Rawlins 1998, Yela & Kitching 2000).
5. Erebidae, tentatively characterised by the dorsally sclerotised scaphium in the male genitalia (Fibiger & Lafontaine 2005) and the silk pore of the larval spinneret concealed by an apical depression characteristically edged laterally by four flaps (in contrast to that of euteliines and others, with many, structurally different flaps) (Crumb 1956, Beck 2000), comprises eleven subfamilies with European representatives: Scoliopteryginae, Rivulinae, Hypeninae, Lymantriinae, Herminiinae, Arctiinae (with Arctiini, Lithosiini and Syntomini), Calpinae, Toxocampinae, Hypenodinae (with Hypenodini and Micronoctuini), Boletobiinae (with Boletobini, Phytometrini, Aventiini, Araeopteronini and Eublemmini; the genus Colobochyla Hübner, , assigned by Fibiger & Lafontaine (2005) and Fibiger et al. (2010) to Phytometrinae, remains unassociated yet) and Erebinae (with Catephiini, Pandesmini, Pericymini, Melipotini, Erebini, Catocalini, Euclidiini and Ophiusini) (Zahiri et al. 2012). However, it must be noted that few relationships are fully resolved yet. One of the more robust association has been found between (Herminiinae+Aganainae) + Arctiinae, which all share a prespiracular, and therefore homologous, counter-tympanal hood (Zahiri et al. 2012; see Kitching & Rawlins 1998 and Yela & Kitching 2000 for alternative interpretations).
6. Euteliidae is best characterised by the double counter-tympanal hood (Richards 1933, Kitching & Rawlins 1998), the female frenulum reduced to a single seta and the pair of lenticular flanges at basal position in the A2 sternal sclerite (Holloway 1985). It has two subfamilies globally, Euteliinae (the only one with European species) and Stictopterinae.
Lafontaine & Fibiger (2006), in another key paper, went another step further. They undertook the most comprehensive noctuid phylogenetic analysis primarily based on morphological characters, but discussed the molecular information accumulated up to then (mostly Weller et al. 1994 and Mitchell et al. 2000, 2006) and incorporated it when appropriate. Since then, no relevant additional information has been made available regarding the traditionally called “trifine noctuids”, or Noctuidae sensu stricto (in which vein M2 in the hind wing is usually vestigial or absent), since Zahiri et al (2011, 2012) included reduced sets of “trifine” species in their analyses and their results render only weak support to most of the trifine clades. For our phylogenetic interpretation of Noctuidae sensu stricto we therefore rely in Lafontaine & Fibiger (2006), as modified by Lafontaine & Schmidt (2010). In short, the earlier known as “trifine noctuids”, i.e. Noctuidae sensu stricto, can be characterized by the medially fused tympanal bullae (Richards 1933, Kitching & Rawlins 1998), by the presence of a single muscle (called M2) between tegumen and transtilla in the male genitalia (Tikhomirov 1979, Yela 1998) and by the possession of a subapical plate in the posterior apophyses of the female genitalia (Fibiger & Lafontaine 2005). Following subfamilies are represented inEurope: Plusiinae, Bagisarinae, Eustrotiinae, Acontiinae (with Acontiini, Hypercalymniini, Armadini and Aediini), Cuculliinae, Oncocnemidinae, Amphipyrinae, Psaphidinae, Pantheinae, Dilobinae, Raphiinae, Acronictinae, Metoponiinae, Heliothinae, Condicinae, Eriopinae, Bryophilinae, Xyleninae, Hadeninae and Noctuinae (Fibiger & Lafontaine 2005, Lafontaine & Fibiger 2006). The relationships among each other are far from being well established in many instances, as is the rank of some of them. For example, several authors subsume Oncocnemidinae in Cuculliinae (Kitching & Rawlins 1998), Psaphidinae in Amphipyrinae (Lafontaine & Schmidt 2010), Pantheinae in Acronictinae (Beck 1992) or Raphiinae in Dilobinae (Fibiger et al. 2009), whereas others consider Xylenini, Hadenini and Noctuini as tribes of a homogeneous Noctuinae (Beck 1960, 1999; Poole 1995; Yela & Kitching 2000; Lafontaine & Schmidt 2010). We prefer to adopt an eclectic position and adhere here to the most extended opinion until the entire lineage/family has been submitted to molecular scrutiny (Zahiri et al. in prep.).
The diagnostic morphological characters of the different taxa have been thoroughly described and discussed in several papers (Kitching 1984, 1987, Lafontaine & Poole 1991, Matthews 1991, Poole 1995, Speidel et al. 1996b, Kitching & Rawlins 1998, Benkhelil 1999, Yela & Kitching 2000, Jacobson & Weller 2002, Goater et al. 2003, DaCosta & Weller 2005, Fibiger & Lafontaine 2005, Lafontaine & Fibiger 2006), and in the volumes of Noctuidae Europaeae. As a summary, the putative morphological apomorphic character states of the European subfamilies are given in Table 1.
Table 1. Updated subfamiliar sequence, with putative apomorphic character states (when known).
Nolidae Bruand, 1846
Nolinae Bruand, 1846
Scaphium with lateral setose patches (Holloway 1998)
Chloephorinae Stainton, 1859 (including Chloephorini Stainton, 1859, Sarrothripini Hampson, 1894 and Eariadini Hampson, 1912)
The group, as a whole, is uncharacterized yet by unequivocal apomorphies. Chloephorini + Sarrothripini have tymbal organs at the male A2 sternite (Holloway 1998, 2003), secondarily lost in a number of species, whereas Eariadini show larval setae D2 and L2 on conical elevations, particularly visible in T2 and T3 (Beck 1960, 1996), and the L3 group of larval setae consists of a single seta in A3 to A6 (Beck 1960, 1996).
Erebidae Leach, 
Scoliopteryginae Herrich-Schäffer,  (for usage of the name see Kühne & Speidel 2004; Speidel & Naumann 2005 and Holloway 2005)
Larval seta MD1 on T1 moved to a very posterior position, on the caudal margin of the segment (Beck 1960, 1992)
Rivulinae Grote, 1895
SV3 larval seta on A1 and A2 displaced dorso-laterally, meeting L3 in a common pinaculum (Beck 1999)
Hypeninae Herrich-Schäffer, 
SV3 larval seta on A1 and A2 equidistant both from SV1 and L3 or even closer to L3, resulting in SV groups of setae that appears to have two rather than three setae (Beck 1999)
Lymantriinae Hampson, 
Single, middorsal glands at the centre of larval A6 and A7 (Godfrey 1987; Holloway et al. 1987; Benkhelil 1999)
Herminiinae Leach, 
Elongations of the tibial sclerites (operculi sensu Berio 1991) covering conspicuous brush organs located mostly on the tibia or on the basitarsi (Smith 1895; Owada 1987; Berio 1991)
Muscle M4 in the male genitalia divided, so that the two parts originate from the apex and from the margin of the pleurite respectively (Tikhomirov 1979)
Arctiinae Leach,  (including Arctiini Leach, , Lithosiini Billberg, 1820 and Syntomini Herrich-Schäffer, )
Tymbal organ in the metepisternum (Minet 1986; Kitching & Rawlins 1998)
Paired eversible pheromone glands at the anal papillae of females (Holloway 1988; Bendib & Minet 1998; Kitching & Rawlins 1998)
Calpinae Boisduval, 1840
Anal margin of forewings sinuate (Kitching & Rawlins 1998)
Starting of the postmedial line of the forewing towards the apex (Kitching & Rawlins 1998)
Toxocampinae Guenée, 1852
Postvaginal plate (female genitalia) half-way divided medially (Goater et al. 2003)
Hypenodinae Forbes, 1954 (including Hypenodini Forbes, 1954 and Micronoctuini Fibiger, 2005)
Larval SV group of setae on A2 consisting of two setae (Beck 1999)
Loss of one SV seta on larval A2 (so that the SV group in the two first segments has a unique 3:2 count), and
First four larval abdominal segments swollen, the first of which is as wide as the thoracic segments (Beck, 1999)
Boletobiinae Guenée,  (with Boletobini Guenée, , Phytometrini Hampson, 1913, Aventiini Tutt, 1896, Araeopteronini Fibiger, 2005 and Eublemmini Forbes, 1954)
The group as a whole is yet uncharacterised by apomorphies. The tympanal system is relatively simple and distinctive (Richards 1933), and will probably render relevant information. For morphological definitions of the tribes see Fibiger et al. (2010)
Erebinae Leach,  (including Catephiini Guenée, 1852, Pandesmini Wiltshire 1990, Pericymini Wiltshire, 1976, Melipotini Grote, 1895, Erebini Leach, , Catocalini Boisduval, , Euclidiini Guenée, 1852 and Ophiusini Guenée, 1837) (for usage of the subfamiliar name see Fibiger & Lafontaine 2005, and Zahiri et al. 2011a)
Styloconic sensilla of the tip of the proboscis situated dorsally (Speidel et al. 2007)
A7 sternite in adult females with a central notch, with ostium bursae moved forwards into the notch (Speidel et al. 1995; Goater et al. 2003)
Euteliidae Grote, 1882
Euteliinae Grote, 1882
Basal abdominal sternite with two internal longitudinal flanges arising from the bases of the anterior apophyses and directed towards midline (Kitching 1987; Holloway 1985)
Noctuidae Latreille, 1809
Plusiinae Boisduval, 
Hair pencil on modified sternite of adult A8 (Kitching 1987)
Laterally protruding raduloid on the larval hypopharynx (Eichlin & Cunningham 1978; Lafontaine & Poole 1991)
Pupal subdorsal grooves between tergites A1-A4 and intersegmental transverse ridges anterior to each moveable abdominal segment (Nakamura 1974; Lafontaine & Poole 1991)
Bagisarinae Crumb, 1956
Bases of the valves of the male genitalia fused together (Ferguson 1997; Fibiger & Lafontaine 2005; Holloway 2009)
Eustrotiinae Grote, 1882
Anellus plate on the male genitalia characteristically lined with spicules or spines (Fibiger et al. 2009)
Antrum (female genitalia) with lateral extensions into postvaginal plate (Fibiger et al. 2009)
Acontiinae Guenée, 1841 (including Acontiini Guenée, 1841, Hypercalymniini Fibiger & Lafontaine, 2005, Armadini Wiltshire, 1961 and Aediini Beck, 1960)
Tympanal organ with a large alula forming a flap over the tympanic opening and with the hood reduced or absent (Richards 1932; Speidel et al. 1996b)
Dorsal crest on the sacculus (Fibiger & Lafontaine, 2005)
(From here on, all subfamilies bear a nodular sclerite or epaulette on the tympanum; Eggers 1919, Speidel et al. 1996b, Yela 1998)
Pantheinae Smith, 1898
Counter-tympanal hood almost or completely reduced (Richards 1932; Kitching 1984)
Dilobinae Aurivillius, 1889
Secondary setae on the basal part of the larval prolegs on A3 to A6 (Merzheevskaya, 1989)
Highly modified basiconic sensillae on the adult proboscis, which surface is densely covered with irregular cuticular dents (Speidel et al. 1996a)
Raphiinae Boisduval, 
Larval seta D1 in T1 and T2 on a conical process; D1 in T1 is moved forward and laterally regarding the standard pattern (Beck 1996; Yela 1998)
Enlarged prolegs with large number of crochets (Crumb 1956; Beck 1996)
Acronictinae Heinemann, 1859
Larval A7 in the first instar larva thin and pale (Beck 1960)
(From here on, adults show basal abdominal brushes and pockets – Lafontaine & Poole 1991; Poole 1995; Speidel & Naumann 1995; Lafontaine & Fibiger 2006 – and corona at the tip of the valva – Lafontaine & Poole 1991; Poole 1995; Speidel et al. 1996b; Lafontaine & Fibiger 2006 – Cuculliinae to Psaphidinae share two synapomorphies: the clasper or harpe is situated close to the ventral edge of the valva – Lafontaine & Poole 1991; Kitching & Rawlins 1998 – and the lateral line of the larva borders the anal plate, see Beck 1960, 1996).
Cuculliinae Herrich-Schäffer,  + Oncocnemidinae Forbes & Franclemont, 1954
Monk-hood disposition of the patagia (Ronkay & Ronkay 1994, 1995)
Long and tubular vesica with an elongated spinose field or one-three cornuti (appearing as irregularly shaped sclerotised plates in one clade)
No unambiguous apomorphy is available at present to characterise either group
Amphipyrinae Guenée, 1837
Heavily sclerotised scaphium as two lateral rods, with subscaphium only sclerotised posteriorly; rest of the male genitalia simple (Fibiger & Lafontaine 2005)
Psaphidinae Grote, 1896
Row of pits on the anterior margin of pupal A10 (Kitching & Rawlins 1998)
Metoponiinae Herrich-Schäffer, 
Species of this subfamily share with Stiriinae the greatly reduced, scale-like spinneret (Fibiger & Lafontaine 2006), but it is at present yet uncharacterised by apomorphies. Metoponiinae species resemble Stiriinae without apical foretibial claw
Heliothinae Boisduval, 
Prothoracic larval setae L1 and L2 arranged transversally, being L2 slightly anterior to L1 (Hardwick 1970; Matthews 1991; Mitter et al 1993)
Condicinae Poole, 1995
Peculiar structure of larval SV3 seta on A1, when present (“Makroborste”; Beck 2000)
Short and semi-scale – like larval spinneret (Crumb 1956; Kitching & Rawlins 1998)
Eriopinae Herrich-Schäffer, 
Streak-like sclerotisation of the scaphium (Fibiger & Lafontaine 2005)
Bryophilinae Guenée, 1852
Plate-like free pleurite between tegumen and vinculum in the male genitalia (Fibiger et al. 2009)
(From here on, all subfamilies show three synapomorphies: larval spinneret flattened, larval seta SD1 on A9 hair-like and clasper or harpe in the centre of the valve connected by rod to ventral margin of sacculus; Lafontaine & Poole 1991, Poole 1995, Speidel et al. 1996b, Lafontaine & Fibiger 2006)
Xyleninae Guenée, 1837
Not yet characterised by apomorphies. Anal edges of the forewings touching each other when adults are resting and double helix-like free pleurite between tegumen and vinculum in the male genitalia point to a very close relationship with Hadeninae, although the surface of the adult eyes is not covered by hairs (Hampson 1903, 1905) in Xyleninae
Hadeninae Guenée, 1837
Not yet characterised by apomorphies. Anal edges of the forewings touch each other when adults are resting; a double helix-like free pleurite between tegumen and vinculum in the male genitalia. Surface of the adult eyes covered by hairs (Hampson 1903, 1905) (diagnostic, but not apomorphic)
Noctuinae Latreille, 1809
Pterothoracic basal sclerites modified, so that wings hold flat over the abdomen in an approximately horizontal plane when adults are resting and costae remain almost parallel to the longitudinal axis of the body (Holloway 1989)
The subfamiliar sequence expressed in Table 1 entails a conception of the phylogenetic relationships among noctuid lineages as in Text Figure 1.
Fig. Fig. 1. Dendrogram showing the conception used in this book about the phylogenetic relationships among the noctuid lineages present inEurope, according to the quadrifamiliar scenario. Nolidae, Erebidae and Eutelidae (subfamilies Nolinae to Eutelinae) are arranged following Zahiri et al. (2012) and Noctuidae (sensu stricto; Plusiinae to Noctuinae) according to our own modification of Mitchell et al. (2006), as interpreted by Lafontaine & Fibiger (2006) and Lafontaine & Schmidt (2010)
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