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Taxonomic variation in larval mandibular structure in Palaearctic Notodontidae (Noctuoidea)

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The mandibles of the 1 st to 5 th instar larvae of 61 species belonging to 32 genera of Palaeartic Notodontidae from Ukraine and Primorskii krai (Far East of Russia) were examined with the use of a scan-ning electron microscope. The characters of the mandibular margin, retinaculum and mandibular carina are discussed. A comparative morphological analysis is provided for all these structures. Apomorphic and plesiomorphic states of the different characters are argumented in relation to the different taxa. The results are compared to the recent classifi cations of Notodontidae. Zusammenfassung. Die Mandibeln der ersten bis fünften Larvenstadien von 61 Arten und 32 Gattungen palärktischer Notodontidae aus der Ukraine und dem Primorski Krai in Russland sind rasterelektronen-mirkoskopisch untersucht. Merkmale des Mandibelrandes, des Retinaculums und der Mandibelcarina werden diskutiert. Eine vergleichende morphologische Analyse wird für diese Merkmale gegeben. Die apomorphen und plesiomorphen Zustände der einzelnen Merkmale werden in Beziehung zu den einzelnen Taxa diskutiert. Die Ergebnisse werden mit rezenten Klassifi kationen der Notodontidae verglichen. Résumé. Les mandibules des chenilles de plusieurs Notodontidés paléarctiques ont été étudiés à l'aide du microscope électronique à balayage. L'étude est fondée sur les chenilles des stades I-V appartenant à 61 espèces de 32 genres provenant de l'Ukraine et de la Région d'Ussuri (au sud de l'Extrême est de la Russie). La morphologie des caractères tels que la marge mandibulaire, le retinaculum et la carène man-dibulaire a été comparée. Une argumentation concernant les états de ces caractères en tant qu'apomorphies et plésiomorphies est fournie. Les récentes classifi cations des Notodontidés sont comparées en regard des résultats obtenus.
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Nota lepid. 31 (2): 165 177
Taxonomic variation in larval mandibular structure in
Palaearctic Notodontidae (Noctuoidea)
IRINA V. D OLINSKAYA
Schmalhausen Institute of Zoology, National Academy of Sciences of Ukraine, UA-01601 Kiev MSP,
Ukraine; e-mail: idolinskaya@izan.kiev.ua
Abstract. The mandibles of the 1st to 5th instar larvae of 61 species belonging to 32 genera of Palaeartic
Notodontidae from Ukraine and Primorskii krai (Far East of Russia) were examined with the use of a scan-
ning electron microscope. The characters of the mandibular margin, retinaculum and mandibular carina
are discussed. A comparative morphological analysis is provided for all these structures. Apomorphic and
plesiomorphic states of the different characters are argumented in relation to the different taxa. The results
are compared to the recent classi cations of Notodontidae.
Zusammenfassung. Die Mandibeln der ersten bis fünften Larvenstadien von 61 Arten und 32 Gattungen
palärktischer Notodontidae aus der Ukraine und dem Primorski Krai in Russland sind rasterelektronen-
mirkoskopisch untersucht. Merkmale des Mandibelrandes, des Retinaculums und der Mandibelcarina
werden diskutiert. Eine vergleichende morphologische Analyse wird für diese Merkmale gegeben. Die
apomorphen und plesiomorphen Zustände der einzelnen Merkmale werden in Beziehung zu den einzelnen
Taxa diskutiert. Die Ergebnisse werden mit rezenten Klassi kationen der Notodontidae verglichen.
Résumé. Les mandibules des chenilles de plusieurs Notodontidés paléarctiques ont été étudiés à l‘aide
du microscope électronique à balayage. L‘étude est fondée sur les chenilles des stades I-V appartenant à
61 espèces de 32 genres provenant de l‘Ukraine et de la Région d‘Ussuri (au sud de l‘Extrême est de la
Russie). La morphologie des caractères tels que la marge mandibulaire, le retinaculum et la carène man-
dibulaire a été comparée. Une argumentation concernant les états de ces caractères en tant qu’apomorphies
et plésiomorphies est fournie. Les récentes classi cations des Notodontidés sont comparées en regard des
résultats obtenus.
Introduction
The mandibles of nal instar notodontid larvae have been described in certain publica-
tions (Godfrey 1984; Godfrey et al. 1989; Miller 1991), but our knowledge concerning
other instars, particularly the morphological peculiarities of the rst instar, is insuf-
cient. Faunal coverage is also inadequate; most information treats notodontid species
from North America (Weller 1987; Dockter 1993). This study is an attempt to, at least
partly, close this gap.
Materials and Methods
This research is based on material that I have collected in Ukraine and Primorskii krai
(Far East of Russia). Eggs were obtained from females captured in the eld. Hatched
larvae were reared to pupae. The epicrania left by caterpillars after moulting as well
as fresh material stored in alcohol were studied. Mandibles were separated from the
epicranium and then examined with a scanning electron microscope (SEM) as well
as a binocular light microscope (MBS-9). The mandibular structures of 1st through
5th larval instars belonging to 61 notodontid species from the following genera were
studied: Euhampsonia Dyar, Furcula Lamarck, Uropyia Staudinger, Stauropus Ger-
mar, Cnethodonta Staudinger, Harpyia Ochsenheimer, Dicranura Reichenbach, Fen-
tonia Butler, Drymonia Hübner, Notodonta Ochsenheimer, Peridea Stephens, Nerice
Nota lepidopterologica, 15.11.2008, ISSN 0342-7536
166 DOLINSKAYA: Larval mandibles of Palaearctic Notodontidae
Walker, Semidonta Staudinger, Pheosia Hübner, Leucodonta Staudinger, Lopho cosma
Staudinger, Pheosiopsis Bryk, Shaka Matsumura, Pterostoma Germar, Ptilodon Hübner,
Lophontosia Staudinger, Hagapteryx Matsumura, Allodonta Staudinger, Hexafrenum
Matsumura, Epodonta Matsumura, Phalera Hübner, Spatalia Hübner, Gluphisia Bois-
duval, Pygaera Ochsenheimer, Gonoclostera Butler, Clostera Samouelle and Micro-
melalopha Nagano. The taxonomic arrangement of these genera follows Schintl meister
(1989) (Table 1).
Results
Table 1 gives a summary of the distribution of the character states found during my
investigations of the mandibular margin, retinaculum, and mandibular carina of the
investigated Palaearctic genera of Notodontidae.
Mandibular margin. The presence of mandibles with a serrate cutting edge in the
rst larval instar, which then becomes smooth in the last instar, has been previously
recorded (Weller 1987; Godfrey et al. 1989). Miller (1991) considered this smooth
mandibular margin of Notodontidae to be an apomorphic character of the family.
The 1st instar mandibular margin in most species possesses 6 acutely-angled denticles.
These are either narrow or broad and they vary in length. The middle denticles are the
largest, while the 6th is usually indistinct (Fig. 1).
First instar larvae show taxonomic differences in mandibular edge structure. First of
all, there is a trend toward smoothing out of the cutting edge. Thus, in most species
the rst instar mandibular edge shows 6 acutely-angled narrow denticles (Hagapteryx,
Phesiopsis, Furcula, and others). Some species of Notodontinae have smooth, broad
and more rounded denticles (e.g., Pheosia, Phalera, Fig. 2). In some taxa, the number
of denticles is reduced to 4, the denticles are broad (perhaps due to fusion of separate
denticles) and attened. Thus, in genus Cnethodonta there are 4 large broad denti-
cles (Fig. 3), while the denticles in species of Stauropus are almost indistinct (Fig. 4).
Weller (1987: 189, g. 2) recorded only 3 nger-like denticles for Litodonta hydromeli
Harvey. Since the third denticle is broad, it is assumed to represent a fusion of two
denticles.
My studies show that the mandibular margin of Notodontidae becomes smooth in the
2nd through 5th instars (Fig. 5). However, in some genera serrate mandibles do not com-
pletely disappear by the 2nd instar, but instead gradually disappear during subsequent
instars, becoming absent by the nal instar. For example, Clostera, Gonoclostera,
Pygaera, Micromelalopha, and Gluphisia have 6 denticles in the 1st instar (Figs 6, 7)
and retain 5 or 6 of them in the 2nd instar, though they become broader (Fig. 8). These
genera have a smooth mandibular margin beginning in the 3rd instar.
In Nerice and Dicranura, 6 denticles persist in the 2nd instar. In Nerice they are short
and rounded (Figs 9, 10) in the 2nd instar, but are expressed as 6 fused but distinct den-
ticles in subsequent instars (Fig. 11). In Dicranura, the 2nd instar shows 5 or 6 indistinct
denticles (Fig. 12). The mandibular margin in the 3rd instar is expressed as 5 fused less
distinct denticles (Fig. 13), while the 4th instar mandibular edge is wavy (Fig. 14).
167
Nota lepid. 31 (2): 165 177
Tab. 1. Character states of the larval mandibles of Palaearctic Notodontidae.
Species
Mandibular margin Retinaculum Mandibular
carina
1st instar 2nd instar 3–5th instars
Euhampsonia cristata (Butler) 6A S S 2 i, L E
Euhampsonia splendida (Oberthür) 6A S S 2 i, L E
Furcula furcula (Clerck) 6A S S 2 i, L E
Furcula bicuspis (Borkhausen) 6A S S 2 i, L E, M
Furcula biÀ da (Brahm) 6A S S 2 i, L E
Uropyia meticulodina (Oberthür) 6A S S 2 i, H E
Stauropus fagi (Linnaeus) 4B S S 1 i, L E, M
Stauropus basalis Moore 4B S S 1 i, L E
Cnethodonta grisescens Staudinger 4B S S 1 i, L E, M
Harpyia milhauseri (Fabricius) 6A S S 2 i, L E
Harpyia umbrosa (Staudinger) 6A S S 2 i, L E
Dicranura ulmi (Denis & Schiffermüller) 6A T W** 3 i, H M
Fentonia ocypete (Bremer) 6A S S 2 i, L E
Drymonia dodonaea [Denis & Schiffermüller] 6A S S 2 i, L E
Notodonta torva (Hübner) 6A S S 2 i, L E
Notodonta dromedarius (Linnaeus) 6A S S 2 i, L E
Notodonta dembowskii Oberthür 6A S SW 2 i, L E
Notodonta tritophus phoebe (Siebert) 6A S T S 2 i, L E
Notodonta ziczac (Linnaeus) 6A S S 2 i, L E
Peridea anceps (Goeze) 6A S S 2 i, L E
Peridea lativitta (Wileman) 6A S S 2 i, L E
Peridea elzet Kiriakoff 6A S S 2 i, L E
Peridea graeseri (Staudinger) 6A S S 2 i, L E
168 DOLINSKAYA: Larval mandibles of Palaearctic Notodontidae
Tab. 1. Continuation.
Species
Mandibular margin Retinaculum Mandibular
carina
1st instar 2nd instar 3–5th instars
Peridea moltrechti (Oberthür) 6A S S 2 i, L E
Peridea oberthueri (Staudinger) 6A T S 2 i, L E
Nerice davidi Oberthür 6A T T 2 i, H M
Nerice leechi Staudinger 6A S T 2 i, H M
Semidonta biloba(Oberthür) 6A S S 2 i, L E
Pheosia tremula (Clerck) 6B S T S 2 i, L E
Pheosia gnoma (Fabricius) 6B S S 2 i, L E, M
Pheosia rimosa Packard 6B S S 2 i, L E
Leucodonta bicoloria (Den. & Schiffermüller) 6A S S 2 i,L E
Lophocosma atriplaga Staudinger 6A S S 2 i, L E
Pheosiopsis cinerea (Butler) 6A S S 2 i, L E
Shaka atrovittatus (Bremer) 6A S T S 2 i, L E
Pterostoma palpina (Clerck) 6A S S 2 i, L E
Pterostoma sinicum Moore 6A S S 2 i, L E
Pterostoma griseum (Bremer) 6A S S 2 i, L E
Ptilodon capucina (Linnaeus) 6A S S 2 i, L E, M
Ptilodon hoegei (Graeser) 6A S S 2 i, L E
Ptilodon cucullina (Denis & Schiffermüller) 6A S S 2 i, L E
Ptilodon ladislai (Oberthür) 6A S S 2 i, L E
Lophontosia cuculus (Staudinger) 6A S S 2 i, L E
Hagapteryx admirabilis (Staudinger) 6A S S 2 i, L E
Allodonta plebeja (Oberthür) 6A S S 2 i, L E, M
Hexafrenum leucodera (Staudinger) 6A S T S 2 i, L E, M
S
S
T
T
S
S
ST
S
S
S
S
S
ST
S
S
S
S
S
S
S
S
S
S
169
Nota lepid. 31 (2): 165 177
Tab. 1. Continuation.
Species
Mandibular margin Retinaculum Mandibular
carina
1st instar 2nd instar 3–5th instars
Epodonta lineata (Oberthür) 6A S SW 2 i, L E
Phalera bucephala (Linnaeus) 6B S S 2 i, L H M
Spatalia argentina (Denis & Schiffermüller) 6A S S 2 i,L E
Spatalia doerriesi Graeser 6A S S 2 i, L E
Spatalia plusiotis (Oberthür) 6A S S 2 i, L E
Spatalia dives Oberthür 6A T S 2 i, L E, M
Gluphisia crenata (Esper)*** 6A T S 3–4 i, L E
Pygaera timon (Hübner) 6A T S 4 i, L M
Gonoclostera timoniorum (Bremer) 6A T S 2 i, L E
Clostera curtula (Linnaeus) 6A T S 2 i, L M
Clostera albosigma curtuloides (Erschoff) 6A T S 2 i, L M
Clostera pigra (Hufnagel) 6A T S 2 i, L M
Clostera anachoreta [Denis & Schiffermüller] 6A T S 2 i, L M
Clostera anastomosis (Linnaeus) 6A T S 2 i, L M
Micromelalopha troglodyta (Graeser) 6A T S 2 i, L M
Mandibular margins with 6 acutely-angled narrow denticles (6A), with 6 broad, Á attened denticles (6B), with 4 broad, Á attened denticle (4B), smooth (S),
toothed (T), or wavy (W).
Retinaculum Á attened or with small projection (L) or with large, high comb (H). The presence of a retinaculum in the 1st instar (1i), in the 2nd instar (2i), in the
3rd instar (3i), in the 4th instar (4i).
Mandibular carina weakly expressed (E) or clearly expressed (M). Average situation between two extremes (ST, SW, EM).
** The mandibular margin in the 3rd instar is expressed as 5 fused, less distinct denticles, while the 4th instar mandibular edge is wavy.
*** did not have the 3rd instar larva and I do not know its characteristics.
ST
S
170 DOLINSKAYA: Larval mandibles of Palaearctic Notodontidae
Dockter (1993: 37, 38, Fig. 8) pointed out that in 1st instar larvae of Heterocampa gut-
tivitta Walker the “third and fourth teeth have Á anges on the bases of their ventrolateral
edges”. My studies of 1st instar larvae show that most species in which the mandibles
have acutely-angled narrow denticles, also have a large medial comb on the inner sur-
face. These imitate “Á anges” on separate denticles (Figs 9, 15, 16).
Retinaculum. The mandibles of Notodontidae have a retinaculum located on the oral
surface. My investigations show that the retinaculum is not yet formed in the 1st instar,
except in the highly specialized genera Stauropus and Cnethodonta, where it is distinct
but not sclerotized (Figs 3, 4).
Dockter (1993) noted the presence of a retinaculum in the 2nd instar for two species of
Heterocampa Doubleday from North America. The Palaearctic species I studied have a
Figs 1–4. Oral surface of mandibles of 1st instar Notodontidae larvae. 1. Ptilodon hoegei (left mandible,
× 600). 2. Phalera bucephala (left mandible, × 720). 3. Cnethodonta grisescens (right mandible, × 600).
4. Stauropus fagi (right mandible, x480).
Figs 1–4. Oral surface of mandibles of 1st instar Notodontidae larvae. 1. Ptilodon hoegei (left mandible,
600 ×). 2. Phalera bucephala (left mandible, 720 ×). 3. Cnethodonta grisescens (right mandible, 600 ×).
4. Stauropus fagi (right mandible, 480 ×).
171
Nota lepid. 31 (2): 165 177
well sclerotized retinaculum in the 2nd instar which becomes more heavily sclerotized
in the 4th and 5th instars, thus providing additional strength for the mandibular cutting
edge. Rarely the retinaculum appears not in the 2nd but in the 3rd instar (Dicranura) and
even in the 4th (Pygaera).
The retinaculum may be Á attened (Pterostona, Micromelalopha,Gonoclostera, and
others, Fig. 5), with a small projection (Semidonta, Euhampsonia and others), or with
a large, high comb (Uropyia, Dicranura, Nerice, Figs 11, 14), which increases in the
3rd and 4th instars.
Mandibular carina. The presence of a mandibular carina and its disposition on the
ventrolateral mandibular surface of some species of Notodontidae has been noted by
Miller (1991: 124, 125, Figs 395–397). My studies show that this structure looks like a
Figs 5–8. Oral surface of right mandible of Notodontidae larvae. 5. 4th instar Gonoclostera timoniorum
(× 220). 6. 1st instar Clostera anachoreta (× 480). 7. 1st instar Pygaera timon (× 600). 8. 2nd instar Pygaera
timon (× 430).
Figs 5–8. Oral surface of right mandible of Notodontidae larvae. 5. 4th instar Gonoclostera timoniorum
(220 ×). 6. 1st instar Clostera anachoreta (480 ×). 7. 1st instar Pygaera timon (600 ×). 8. 2nd instar Pygaera
timon (430 ×).
172 DOLINSKAYA: Larval mandibles of Palaearctic Notodontidae
Figs 9–14. Oral surface of mandibles of Notodontidae larvae. 9–11. Nerice davidi (right mandible).
9. 1st instar (× 660). 10. 2nd instar (× 440). 11. 3rd instar (× 160). 12–14. Dicranura ulmi.12. 2nd instar
(right mandible, × 450). 13. 3rd instar (left mandible, × 230). 14. 4th instar (right mandible, × 110).
Figs 9–14. Oral surface of mandibles of Notodontidae larvae. 9–11. Nerice davidi (right mandible). 9.
1st instar (660 ×). 10. 2nd instar (440 ×). 11. 3rd instar (160 ×). 12–14. Dicranura ulmi. 12. 2nd instar (right
mandible, 450 ×). 13. 3rd instar (left mandible, 230 ×). 14. 4th instar (right mandible, 110 ×).
173
Nota lepid. 31 (2): 165 177
Figs 15–20. Surface of mandibles of Notodontidae larvae. 15. Oral surface of right mandible of 1st instar
Clostera pigra (660 ×). 16. Lateral surface of left mandible of 1st instar Furcula bi da (600 ×). 17. Lateral
surface of left mandible of 1st instar Notodonta torva (780 ×). 18. Part of mandibular carina (above side)
of right mandible of 2nd instar Spatalia dives (440 ×). 19. Part of mandibular carina (above side) of right
mandible of 4th instar Uropyia meticulodina. 20. Ventrolateral surface (area limited by mandibular carina)
of left mandible of 4th instar Dicranura ulmi (140 ×).
174 DOLINSKAYA: Larval mandibles of Palaearctic Notodontidae
comb, limited to the ventrolateral surface of the mandible (see Fig. 20). Most genera of
Notodontidae have a more or less developed carina (Figs 17–20) and only some have
a clearly expressed carina (Dicranura,Phalera, Nerice,Clostera, and others). This
character is difÀ cult for taxonomic interpretation.
Mandibles with secondary setae. Gardner (1943) and Miller (1991) noted the pres-
ence of secondary setae on the mandibles of Thaumetopoeinae. I found secondary setae
on the mandibles in 4th and 5th instar larvae of Dicranura (Fig. 20), but nowhere else
among the taxa studied.
Discussion
The classiÀ cation of the Notodontidae is in dire need of improvement. In the systems
proposed by different authors an uncertainty about the number of subfamilies as well
as the classiÀ cation of their genera remains until now. The recent classiÀ cations of the
Notodontidae (Tikhomirov 1981; Schintlmeister 1985; Miller 1991) have the best ar-
gumentations. Now I want to show how the results of my original investigations agree
or disagree with the above-mentioned classiÀ cations.
The larval mandibles of the majority of the investigated genera of Notodontidae are
uniform. Their mandibular margin possesses 6 acutely-angled denticles in the À rst in-
star, which then becomes smooth in the 2nd through 5th instars. Such structure is present
in 21 genera, namely Euhampsonia, Furcula, Uropyia, Harpyia, Fentonia, Drymonia,
Notodonta, Peridea, Semidonta, Leucodonta, Lophocosma, Pheosiopsis, Shaka,
Pterostoma, Ptilodon, Lophontosia, Hagapteryx, Allodonta, Hexafrenum, Epodonta,
and Spatalia. In this case my data are mostly coordinated with the system of Tikhomirov
(1981) where these genera, except Furcula are included into subfamily Notodontinae.
Genus Furcula in the classiÀ cations of Tikhomirov (1981) and Schintlmeister (1985) is
placed in the derived subfamily Cerurinae. On the other hand, Miller (1991) transferred
this genus to subfamily Notodontinae, in tribe Dicranurini. My data of the larval man-
dible structure do not contradict this opinion.
The placement of genus Harpyia in the Notodontinae by Tikhomirov (1981) is sup-
ported by characters of the structure of the larval mandibles whereas other peculiarities
of the larva, egg, and pupa characterize this genus as highly specialized (Dolinskaya
1986, 1987 a, c; Dolinskaya & Plushch 2003), which is reÁ ected in the classiÀ cations
proposed by Schintlmeister (1985) and Miller (1991). The À rst author placed this ge-
nus into subfamily Stauropinae while the second included it into the Heterocampinae,
Stauropini.
Genera Pheosia and Phalera as well as the above-mentioned 21 genera are character-
ized by mandibles with 6 acutely-angled denticles in the À rst instar; however, they are
not narrow and acute, but broad and Á lattened.Tikhomirov (1981) included both gen-
era into subfamily Notodontinae and noted that Pheosia is characterized by the highest
specialization among the Notodontinae. The genus is characterized also by a rather spe-
cialized sculpture of the egg chorion (Dolinskaya 1987 b, c). The taxonomic position of
the genus is intended to be investigated further. There are two opinions concerning the
position of genus Phalera in the notodontid classiÀ cation. Tikhomirov (1981) consid-
175
Nota lepid. 31 (2): 165 177
ers that it must be included within the Notodontinae while the other authors suggested
that it belongs to the Phalerinae (Schintlmeister 1985; Miller 1991).
Genera Stauropus and Cnethodonta have Á attened, broad mandibles with 4 denticles
in the 1st instar. I consider this state to be apomorphic because most representatives of
the outgroup (Noctuidae, Lymantriidae, Arctiidae) have the mandibular edge with 5–6
acutely-angled, narrow denticles in the 1st instar. Besides, these genera have a retinacu-
lum in the 1st instar. In this case my conclusions are consistant with those of the other
authors. Miller (1991) included these two genera into the Heterocampinae, Stauropini,
while Schintlmeister (1985) and Tikhomirov (1981) placed only genus Stauropus
into their Stauropinae. Tikhomirov (1981), based on characters of the functional mor-
phology of the genitalia retains Cnethodonta in the less advanced Notodontinae, i.e.
Cnethodonta is regarded by him as the less advanced genus relative to Stauropus, and
this fact is also corroborated here (the denticles of the 1st instar larvae are less Á at-
tened).
In genera Clostera, Gonoclostera, Pygaera, Micromelalopha, Gluphisia, Dicranura,
and Nerice, as opposed to the other notodontid larvae, the serrate mandibles do not dis-
appear in the 2nd instar and there is a smooth process of their transformation to the last
instar. I consider such state to be plesiomorphic because of the presence of mandibular
denticles in 1–5th instars in most representatives of the outgroup. The mandibular margin
becomes smooth in the 3rd instar in Clostera, Gonoclostera, Pygaera,Micromelalopha,
and Gluphisia. In the 4th instar of Dicranura the mandibular edge becomes sinuous. In
Nerice the mandibular edge looks like 6 fused distinct denticles until the 5th instar. The
above-mentioned genera have common characters as well as speciÀ c peculiarities.
For example, Clostera, Pygaera, Micromelalopha, Dicranura, and Nerice have a well
expressed carina. In Gonoclostera and Gluphisia the mandibular carina is weakly
expressed. Pygaera,Gluphisia, and Dicranura may be united by the absence of the
retinaculum in the 2nd instar, whereas in the rest of the genera this structure is present.
Besides that, Clostera, Pygaera,and Micromelalopha have the retinaculum with only
a small projection while Dicranura and Nerice have a large, high comb. In addition,
Dicranura have secondary setae on the mandibles in the 4th and 5th instars.
Concerning genera Clostera, Gonoclostera, Pygaera, and Micromelalopha my data
coincide with the classiÀ cations of the three above-mentioned authors. In the classiÀ ca-
tions of Tikhomirov(1981) and Schintlmeister (1985) these genera are included into the
primitive subfamily Pygaerinae. Miller (1991) placed into this subfamily only genus
Clostera and said that following further investigation genera Micromelalopha, Pygaera,
and Gonoclostera perhaps will be included into Pygaerinae also. It must be noted that
the mandibular denticles of Micromelalopha in the 2nd instar are rather broadened, Á at-
tened, and rounded in comparison with those of Clostera, Gonoclostera,and Pygaera.
I consider this state of the denticles to represent an apomorphy for this genus.
Gluphisia requires additional research. The peculiarities of the mandibles and pupa
(Dolinskaya 1986, 1989) show similarities with genus Clostera, Gonoclostera,
Pygaera, and Micromelalopha. The data that I obtained do not match with any of the
published classiÀ cations. Packard (1895) placed Gluphisia into the separate subfamily
Gluphisiinae; Tikhomirov (1981) and Miller (1991) into subfamily Notodontinae.
176 DOLINSKAYA: Larval mandibles of Palaearctic Notodontidae
Schintlmeister (1985) placed Dicranura into the derived Stauropinae together with
Harpyia and Stauropus. The results of my studies are not concordant with this point
of view. I consider that Dicranura has a complex of primitive attributes that are char-
acterictic of the Pygaerinae and Thaumetopoeinae, as well as a complex of specialized
features that are characteristic only to the genus. Thus, it is necessary to undertake more
detailed investigations.
The position of genus Nerice in the Notodontidae remains unclear. Among all known
classi cations its status is discussed only by Packard (1895) and Tikhomirov (1981)
who include it into the Notodontinae.
Summarizing the above-discussed characters of the Notodontidae it can be concluded
that the morphology of the larval mandibles allows to unravel related groups within the
family. The genera with more specialized morphology, as a rule, possess apomorphic
states of the mandibular characters. However, the reconstruction of the Notodontidae
phylogeny is not the main goal of the present paper, and a well-argumented classi ca-
tion should be based on a phylogenetic analysis involving a complex of characters,
including larval, but also characters of the eggs, pupae, and imagos.
Acknowledgements
I am very grateful to Z. A. Panina (Department of Electronic Microscopy, N. G. Kholodny Institute of
Botany, National Academy of Sciences of Ukraine, Kiev) for her help with the SEM and to V. V. Zhuravlev
(Schmalhausen Institute of Zoology, National Academy of Sciences of Ukraine, Kiew) for the help ar-
ranging the illustrations. I am obliged to Mr I. Kostyuk (Zoological Museum of the Shevchenko National
University in Kiev), to Dr A. Schintlmeister (Germany), and to Dr G. L. Godfrey (Center for Biodiversity,
Illinois Natural History Survey, USA) for their assistance with literature.
References
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H. subrotata (Notodontidae). – Journal of the Lepidopterists’ Society 47 (1): 32–48.
Dolinskaya, I. V. 1986. Chrysalid morphology of certain Notodontid moth species (Lepidoptera,
Notodontidae) of the fauna of the USSR. – Vestnik zoologii 2: 59–66 [in Russian].
Dolinskaya, I. V. 1987a. Morphology of the eggs of the Notodontidae (Lepidoptera) of the fauna of the
USSR. Communication 1. – Vestnik zoologii 1: 65–74 [in Russian].
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Entomological Society 97 (4): 455–470.
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Supplementary resource (1)

... Schintlmeister (2008) included Spatalia in Pygaerinae together with Gluphisia, Gonoclostera, Pygaera, Clostera and Micromelalopha. The characters of the larvae (Dolinskaya, 2008(Dolinskaya, , 2011a(Dolinskaya, , 2013Dolinskaya & Pljushch, 2003) are mostly coordinated with Tikhomirov's system (1981). However, the pupae show a common character in sculpture of А10 for Spatalia, Fentonia, Wilemanus (Dolinskaya, 1986(Dolinskaya, , 1989(Dolinskaya, , 1993 and some species from Heterocampinae (Miller, 1992). ...
... These genera have also several additional unique characters-pitted egg chorion sculpture and transition region. Their larvae and pupae are also characterized by a complex of unique characters (Dolinskaya, 1989(Dolinskaya, , 2008(Dolinskaya, , 2013Dolinskaya & Pljushch, 2003). Some authors consider Cnethodonta as less advanced genus than Stauropus and even place these genera in different subfamilies (Matsunura, 1929;Tikhomirov, 1981). ...
... Matsumura (1929), Tikhomirov (1981) and Schintlmeister (2008) placed Cerura and Furcula in subfamily Cerurinae. Miller (1991), and after him Nakamura (2007) The previously studied larval characters of the mandibles, the head microsculpture and the cranial setae (Dolinskaya, 2008(Dolinskaya, , 2011(Dolinskaya, a, 2013 showed that Clostera, Pygaera, Dicranura, Leucodonta, Phalera unites with the genera Gonoclostera, Micromelalopha and Gluphisia only a complex of plesiomorphic characters. These studies confirm the point of view of Tikhomirov (1981) which included Clostera, Pygaera, Gonoclostera and Micromelalopha into the primitive subfamily Pygaerinae. ...
Article
On the basis of comparative-morphological analysis of 43 genera and 92 species of Palaearctic Notodontidae, as well as the study of the eggs of outgroup species, complexes of characters that are diagnostic, taxonomic or phylogenetic are singled out. It is shown that the egg characteristics are of great taxonomic value at species and generic levels. Some characters are useful for grouping genera. In general, a complex of characters should be used, because different species or genera often share the same characters. Possible apomorphic and plesiomorphic states of the different characters are discussed in relation to the different taxa. The results of this study are discussed with reference to recently published classifications of Notodontidae. As a result of the studies, the keys for identification to the eggs of 43 genera and 92 species of notodontid moths from the Palaearctic region are presented. Reliable diagnostic characters that do not disappear with the injury of eggs or with eggs preserved in alcohol were used. Characters including egg shape, egg and chorion colour, the shape of gnawed holes in eggs when caterpillars hatched, chorionic sculpture, the type of oviposition, foodplants, and geographic distribution of the genera and species were applied. Occasionally, characters that are typical for live eggs, which vary during development, were used. These are characters of egg colour and pattern. The keys are illustrated with photographs made using a digital camera and a scanning electron microscope.
... The larval head microsculpture and morphology of the larval mandibles allows unraveling related groups within the family that can be used to accomplish classification of the family (Dolinskaya, 2008(Dolinskaya, , 2011. While studying these structures we noted the great variety of setae on the larval head. ...
... Tikhomirov (1981) and Schintlmeister (2008) included the genera Clostera, Pygaera, Gonoclostera and Micromelalopha in here. The mandibular characters and head sculpture of the caterpillars (Dolinskaya, 2008(Dolinskaya, , 2011 also support this placement. However, the characters of the cranial larval setae show rather puzzle distribution pattern. ...
... Same setae are typical for Pygaera. Our previous studies (Dolinskaya, 2008(Dolinskaya, , 2011 show strong similarity of Dicranura to the Pygaerinae and especially with the genus Pygaera. ...
Article
Full-text available
Larval cranial setae of each larval instar of 66 species belonging to 35 genera of Palaeartic Notodontid moths from Ukraine and Far East of Russia (Primorskii krai) was examined with the use of a scanning electron microscope. A comparison with outgroup species - Lasiocampoidea (Lasiocampidae), Sphingoidea (Sphingidae) and Noctuoidea (Erebidae: Lymantriinae, Arctiinae; Noctuidae) is conducted. Main kinds of setae during larval development and their transformation are discussed. Possible apomorphic and plesiomorphic states of the different characters are discussed in relation to the different taxa.
... The larval head microsculpture and morphology of the larval mandibles allows unraveling related groups within the family that can be used to accomplish classification of the family (Dolinskaya, 2008(Dolinskaya, , 2011. While studying these structures we noted the great variety of setae on the larval head. ...
... Tikhomirov (1981) and Schintlmeister (2008) included the genera Clostera, Pygaera, Gonoclostera and Micromelalopha in here. The mandibular characters and head sculpture of the caterpillars (Dolinskaya, 2008(Dolinskaya, , 2011 also support this placement. However, the characters of the cranial larval setae show rather puzzle distribution pattern. ...
... Same setae are typical for Pygaera. Our previous studies (Dolinskaya, 2008(Dolinskaya, , 2011 show strong similarity of Dicranura to the Pygaerinae and especially with the genus Pygaera. ...
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Mandibular incisor lobes are important for insect feeding behavior, living habits and niche. However, the molecular regulation of insect incisor lobe development remains unknown. In this study, we found that two maize pests, oriental armyworm Mythimna separata and fall armyworm Spodoptera frugiperda, have different feeding patterns in maize, which are closely associated with the different development patterns of their incisor lobes. Different from first to sixth instar S. frugiperda, which feed on leaf tissues and whorls with sharp incisor lobes, older instars of M. separata feed from leaf margins with no incisor lobes. Hox gene Deformed (Dfd) is important for head appendages, but its function in incisor lobe development is not clear. Here, Dfds were identified from two armyworm species, and both were expressed highly in heads and eggs. Interestingly, the expression levels of MsDfd were relatively high in larval mandibles and decreased dramatically from fourth-instar mandibles in M. separata. Knockdown of MsDfd resulted in malformed mandibles with no incisor lobe in M. separata, making the larvae unable to perform window-feeding. However, RNAi of SfDfd did not affect the mandibles and window-feeding pattern of S. frugiperda, indicating the different roles of Dfd in these two species. Moreover, the mortality of new first instar M. separata increased after feeding dsMsDfd but did not for S. frugiperda feeding dsSfDfd. These findings revealed that Dfd mediated the larval mandibular incisor lobe morphology, affecting its feeding pattern in M. separata, broadening the knowledge of Dfd functions in insect mandibles and feeding behavior.
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In Indonesia, lobster moth (Neostauropus alternus) is called by ulat kuda, uler jaran or hileud kuda. The larva has 4 + 1 pair prolegs; 4 pairs prolegs located on A3-6, while one pair proleg is on A10 which is modified as like as a pair of antenna or spines. The A10 prolegs lifted upwards when the larva is resting so it looks like pegs (peglike structure), while the larva adheres on tree by using 4 pairs of prolegs A3-6. Crochet on the A10 almost not found. Furthermore, Notodontidae have secondary setae or additional setae on prolegs A3-6
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The larval head microsculpture of each instar of 66 species belonging to 35 genera of Palaeartic notodontids from Ukraine and Far East of Russia (Primorskii krai) was examined with the use of a scan-ning electron microscope. A comparison with representatives from Lasiocampoidea (Lasiocampidae) and Noctuoidea (Erebidae: Lymantriinae, Arctiinae; Noctuidae) is conducted. Differences in head microsculp-ture and the transformation during development of different larval instars are discussed. Apomorphic and plesiomorphic states of these characters are also discussed. The results of this study are discussed with reference to recently published classifi cations of Notodontidae.
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Diversity of the Larval Cranial Setae in Palaearctic Notodontidae (Noctuoidea) and and their Taxonomic Distribution. Dolinskaya I. V. – Larval cranial setae of each larval instar of 66 species belonging to 35 genera of Palaeartic Notodontid moths from Ukraine and Far East of Russia (Primorskii krai) was exam-ined with the use of a scanning electron microscope. A comparison with outgroup species – Lasiocampoidea (Lasiocampidae), Sphingoidea (Sphingidae) and Noctuoidea (Erebidae: Lymantriinae, Arctiinae; Noctuidae) is conducted. Main kinds of setae during larval development and their transformation are dis-cussed. Possible apomorphic and plesiomorphic states of the different characters are discussed in rela-tion to the different taxa. K e y w o r d s: Notodontidae, Lepidoptera, larvae, morphology, larval cranial setae, classification, scan-ning electron microscopy. Ðàçíîîáðàçèå ùåòèíîê ãîëîâû ïàëåàðêòè÷åñêèõ õîõëàòîê (Notodontidae, Noctuoidea) è èõ òàêñîíî-ìè÷åñêîå ðàñïðåäåëåíèå. Äîëèíñêàÿ È. Â. – Ñ ïîìîùüþ ñêàíèðóþùåãî ýëåêòðîííîãî ìèêðîñêî-ïà èçó÷åíû ùåòèíêè ãîëîâû ãóñåíèö âñåõ âîçðàñòîâ 66 âèäîâ èç 35 ðîäîâ ïàëåàðêòè÷åñêèõ õîõ-ëàòîê èç Óêðàèíû è Ïðèìîðñêîãî êðàÿ Ðîññèè. Ïðîâåäåíî ñðàâíåíèå ñ ïðåäñòàâèòåëÿìè âíåø-íåé ãðóïïû – Lasiocampoidea (Lasiocampidae), Sphingoidea (Sphingidae) è (Erebidae: Lymantriinae, Arctiinae; Noctuidae). Ðàññìîòðåíû âåðîÿòíûå íàïðàâëåíèÿ ïðåîáðàçîâàíèé â âîëîñÿíîì ïîêðî-âå ãîëîâû ó ãóñåíèö ðàçëè÷íûõ âîçðàñòîâ. Îáñóaeäàþòñÿ âîçìîaeíûå àïîìîðôíûå è ïëåçèî-ìîðôíûå ñîñòîÿíèÿ ïðèçíàêîâ ðàçëè÷íûõ òàêñîíîâ. Ê ë þ ÷ å â û å ñ ë î â à: Notodontidae, Lepidoptera, ãóñåíèöû, ìîðôîëîãèÿ, ùåòèíêè ãîëîâû, êëàñ-ñèôèêàöèÿ, ñêàíèðóþùèé ýëåêòðîííûé ìèêðîñêîï.
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The simple matching coefficient (per cent similarity) and the coefficients of taxonomic distance are calculated for 47 european Notodontidae basing upon 50 characters. For this a calculator program in basic is given. The obtained dendrograms are used for discussing classification and systematics of the european Notodontidae. Clearly the ranks of genera and subfamilies are shown in the dendrograms.
Notes on the larva of Cargida pyrrha (Notodontidae)
  • G L Godfrey
Godfrey, G. L. 1984. Notes on the larva of Cargida pyrrha (Notodontidae). -Journal of the Lepidopterists' Society 38 (2): 88-91.
Two mouthpart modifi cations in larval Notodontidae (Lepidoptera): their taxonomic distributions and putative functions
  • G L Godfrey
  • J S Miller
  • D J Carter
Godfrey, G. L., J. S. Miller & D. J. Carter 1989. Two mouthpart modifi cations in larval Notodontidae (Lepidoptera): their taxonomic distributions and putative functions. – Journal of the New York Entomological Society 97 (4): 455–470.