Nota lepid. 34 (1): 11 – 28
Larval head microsculpture in Palaearctic Notodontidae
(Noctuoidea) and its signiﬁ cance for the systematics
of the family
Irina V. Dolinskaya
Schmalhausen Institute of Zoology, National Academy of Sciences of Ukraine, B. Chmielnicki St.15,
Kiev, 01601 Ukraine; firstname.lastname@example.org
Abstract. 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.
First studies of head microsculpture in notodontid larvae date back to the last century
(Bell 1935, Gardner 1943). A more detailed study of the cranial surface of notodontid
larvae was undertaken by Miller (1991). He studied 48 species of notodontid cater-
pillars that occur in the Palaearctic and the Americas and also examined 13 species
from other groups (Doidae; Erebidae: Arctiinae and Lymantriinae; Noctuidae; and
Oenosandridae). Miller (1996, 2009a, 2009b) later described the head surface of the
Neotropical notodontid caterpillars in the subfamily Dioptinae. Unfortunately, these
studies only looked at the ﬁ nal larval instar, making it impossible to draw any conclu-
sions regarding the microsculpture of the larval head as it changes among the different
Materials and Methods
This research is based on material collected in Ukraine and Far East of Russia (Pri-
morskii krai). Eggs were obtained from females captured at light. Hatched larvae were
reared to pupae. The epicrania left by caterpillars after moulting, as well as fresh mate-
rial preserved in alcohol, were studied. The epicranium was examined with a scanning
electron microscope (SEM) and a binocular light microscope (MBS 9). The micro-
sculpture of the head of 1st through 5th larval instars belonging to 66 no to dontid species
from the following genera were studied: Euhampsonia Dyar, Cerura Schrank, Furcula
Lamarck, Uropyia Staudinger, Dicranura Reichenbach, Harpyia Ochsenheimer, Stau-
ropus Germar, Cnethodonta Staudinger, Fentonia Butler, Neopheosia Matsumura,
Drymonia Hübner, Notodonta Ochsenheimer, Peridea Stephens, Nerice Walker, Phe o sia
Nota lepidopterologica, 21.10.2011, ISSN 0342-7536
12 Dolinskaya: Larval head microsculpture in Palaearctic Notodontidae
Hübner, Leucodonta Staudinger, Lophocosma Staudinger, Ellida Grote, Phe o siop sis
Bryk, Shaka Matsumura, Pterostoma Germar, Ptilodon Hübner, Lophontosia Stau-
dinger, Hagapteryx Matsumura, Togepteryx Matsumura, Semidonta Staudinger, Allo-
donta Staudinger, Epodonta Matsumura, Phalera Hübner, Spatalia Hübner, Glu phi-
sia Boisduval, Gonoclostera Butler, Pygaera Ochsenheimer, Clostera Samouelle, and
Micromelalopha Nagano. The taxonomic arrangement of these genera follows Schintl-
In order to clarify the character states and polarity within Notodontidae, representa-
tives of related families belonging to Lasiocampoidea, as well as other members of
Noctuoidea (Minet 1994; Kuznetzov & Stekolnikov 2001), were used as outgroup
taxa. The following species were studied: Euthrix potatoria Linnaeus, Gastropacha
quercifolia Linnaeus (Lasiocampidae), Teia dubia Tau scher, Arctornis l-nigrum Mül-
ler (Erebidae: Lymantriinae), Rhyparioides amurensis Bremer, Chio narc tia nivea
Mėnė triės (Erebidae: Arctiinae), Calocasia coryli Linnaeus, and Egira conspicillaris
It should be noted that the degree to which microsculpture can be examined and
described depends on the microscopic magniﬁ cation. In this study the term “smooth
microsculpture” is used only when microsculpture is not visible with magniﬁ cations
Comparative Morphology of Larval Head Microsculpture
The taxa and characters examined are listed in Table 1. The microsculpture varies from
instar to instar. First instar larvae are mostly without distinct microsculpture, with the
head surface being smooth (Ptilodon, Pterostoma, Spatalia and others; Fig. 1) or slight-
ly wrinkled (Harpyia, Cnethodonta; Fig. 2). In some genera, microsculpture is visible
in the apical part of the head and partly laterally where it is expressed as slight wrinkles
(Gonoclostera, Fentonia; Fig. 3) or pits (Gluphisia). In contrast, in Pygaera, Cerura,
and Furcula, the surface bears homogeneous, small, densely situated tubercles. In
Pygaera these structures are almost indistinct, smooth, and oval. In Cerura and Furcula
these tubercles are very distinct and visible even with a light microscope (Fig. 4).
In 2nd instar, the microsculpture of many genera is still smooth (Ptilodon, Allodonta).
However, sometimes the surface has small, homogeneous, and occasional tubercles
arranged on a background of implicated ﬁ brae (Epodonta, Euhampsonia, Pheosia;
Fig. 5). In some genera (Cerura, Furcula, Uropyia, Harpyia, Fentonia) the head shows
a background of heterogeneous small tubercles with occasional large tubercles distin-
guished by plicated edges. The latter are either scattered among smaller ones (Fig. 6)
or arranged as more or less distinct groups (Fig. 7).
In 3rd instar some genera have a microsculpture identical to that of preceding in-
stars. It is either smooth (Phalera; Fig. 8), slightly wrinkled with almost indistinct
Nota lepid. 34 (1): 11 – 28
Figs 1 – 6. Larval head surface of Notodontidae 1. 1st instar Ptilodon saturate hoegei. 2. 1st instar Harpyia
umbrosa. 3. 1st instar Gonoclostera timoniorum. 4. 1st instar Furcula bicuspis. 5. 2nd instar Epodonta
lineata. 6. 2nd instar Fentonia ocypete. Scale bar 1 – 4 (100 μ); 5, 6 (10 μ).
14 Dolinskaya: Larval head microsculpture in Palaearctic Notodontidae
Table 1. Character states of the larval head surface of Palaearctic Notodontidae. FH: microsculpture of
heterogeneous tubercles sparsely situated on a background of densely implicated ﬁ brae; FO: microsculp-
ture of tubercles sparsely situated on a background of densely implicated ﬁ brae; H: heterogeneously tu-
bercled microsculpture; M: microsculpture with microtrichiae; O: homogeneous microsculpture; P: pitted
microsculpture; РО: microsculpture with tubercles in crateriform depressions; S: smooth microsculpture;
SH: microsculpture with heterogeneous tubercles sparsely situated on smooth background; SO: micro-
sculpture with homogeneous tubercles sparsely situated on a smooth background; SP: pits developed only
on part of head; SW: head surface weakly wrinkled. WO: wrinkled microsculpture with homogeneous tu-
bercles. Abbreviations: cone-shaped protuberances (co); corrugated microsculpture (cor); large tubercles
concentrated in distinct groups (g).
instar 1 instar 2 instar 3 instars 4 – 5
Euhampsonia cristata (Butler) S O H, g H, g
Euhampsonia splendida (Oberthür) S O H, g H, g
Cerura erminea (Esper) O cor Н, cor H, cor H, cor,WO1
Furcula furcula (Clerck) O cor H, cor H, cor H, cor
Furcula bicuspis (Borkhausen) O cor H, cor H, cor H, cor
Furcula biﬁ da (Brahm) O H, cor H,g,cor H, cor
Uropyia meticulodina (Oberthür) S H, g, сo H,g, сo H,g, сo
Dicranura ulmi (Denis & Schiffermüller) S O O O
Harpyia milhauseri (Fabricius) S H, сo H, сo H, сo
Harpyia umbrosa (Staudinger) S H, сo H, сo H, сo
Stauropus fagi (Linnaeus) S O O H, co
Stauropus basalis Moore S S – –
Cnethodonta grisescens Staudinger S O H, cor H, cor
Fentonia ocypete (Bremer) S H, cor H,g,cor H, g, cor
Neopheosia mandschurica (Oberthür) S – – –
Drymonia dodonaea (Denis & Schiffermüller) S ОHH
Notodonta torva (Hübner) S O H, g H, g
Notodonta dromedarius (Linnaeus) S O H, g H, g
Notodonta dembowskii Oberthür S O H, g H, g
Notodonta tritophus phoebe (Siebert) S O H, g H, g
Notodonta ziczac (Linnaeus) S O H, g H, g
Peridea anceps (Goeze) S O H H
Peridea lativitta (Wileman) S O H H
Peridea elzet Kiriakoff S O H H
Peridea graeseri (Staudinger) S O H, co H, co
Peridea gigantea (Butler) S O H H
Peridea oberthueri (Staudinger) S O H H
Peridea moltrechti (Oberthür) S O H H
Nerice leechi Staudinger S O H, g H, g
Nerice davidi Oberthür S O H, g H, g
Pheosia tremula (Clerck) S O O H
Pheosia grummi (Christoph) S – – –
Pheosia gnoma (Fabricius) S O O H
Pheosia rimosa Packard S O O H
Leucodonta bicoloria (Denis & Schiffermüller) S S SW SW
Lophocosma atriplaga Staudinger S O H H
Ellida branickii (Oberthür) S – – –
Pheosiopsis cinerea (Butler) S O H, g H, g
Shaka atrovittatus (Bremer) S O H, g H, g
Pterostoma palpina (Clerck) S O H H, g
Pterostoma sinica Moore S S O H, g
Pterostoma griseum (Bremer) S S O H, g
Ptilodon capucina (Linnaeus) S S O SO
Ptilodon saturate hoegei (Graeser) S S O SO
Nota lepid. 34 (1): 11 – 28
cells (Micromelalopha, Clostera, Leucodonta; Figs 9 – 11), pitted (Gluphisia; Figs
12, 13), or homogeneously tubercled (Pheosia, Lophontosia; Fig. 14). Most species
have a consistent microsculpture similar to that of the last instar, but less distinct. In
Gonoclostera the surface shows clearly visible small tubercles on a background of
densely interlaced ﬁ brae (Fig. 15). In Ptilodon species more or less distinctive tuber-
cles can be observed on a smooth or interlaced ﬁ brae background (Fig. 16). In most
genera, the microsculpture becomes heterogeneously tubercled with large tubercles on
a background of small homogeneous tubercles. It must be noted that the latter are ar-
ranged either randomly among more small tubercles (Lophocosma, Pheosia, Spatalia;
Fig. 17) or are concentrated as separate groups (Notodonta, Euhampsonia, Epodonta;
Table 1. Continuation.
instar 1 instar 2 instar 3 instars 4 – 5
Ptilodon cucullina (Denis & Schiffermüller) S S O FO
Ptilodon ladislai (Oberthür) S S O SO, g
Lophontosia cuculus (Staudinger) S – FH FH, g
Hagapteryx admirabilis (Staudinger) S O H, g H, g
Togepteryx velutina (Oberthür) – – – H, g
Semidonta biloba (Oberthür) S S SH H
Allodonta plebeja (Oberthür) S S O H
Allodonta leucodera (Staudinger) S S O H
Epodonta lineata (Oberthür) S FO FO H, g
Phalera bucephala (Linnaeus) S S S S
Spatalia argentina (Denis & Schiffermüller) S S H H
Spatalia doerriesi Graeser S O H H
Spatalia plusiotis Oberthür S – – –
Spatalia dives Oberthür S S H H
Gluphisia crenata (Esper) S, SP SP P P
Gonoclostera timoniorum (Bremer) SW SW O РО
Pygaera timon (Hübner) O O O O
Clostera albosigma curtuloides (Erschoff) S S SW SW
Clostera pigra (Hufnagel) S S SW SW
Clostera anachoreta (Denis & Schiffermüller) S S SW SW
Clostera anastomosis (Linnaeus) S S SW SW
Micromelalopha troglodyta (Graeser) S S SW SW
Euthrix potatoria (Linnaeus) S M M M
Gastropacha quercifolia (Linnaeus) S M M M
Teia dubia (Tauscher) S S S S
Arctornis l-nigrum (Müller) S S – M
Rhyparioides amurensis (Bremer) S S – S
Chionarctia nivea (Mėnėtriės) S S S SW
Egira conspicillaris (Linnaeus) S S S SW, O (partly)2
Calocasia coryli (Linnaeus) S S S SW
1 Cerura erminea: ‘Н, cor’ refers to head sculpture of 4th instar and ‘WO’ to that of 5th instar.
2 in 4th instar head microsculpture is smooth.
16 Dolinskaya: Larval head microsculpture in Palaearctic Notodontidae
Fig. 18) that are more or less distinct. In some genera (Furcula, Fentonia, Notodonta,
Spatalia) the microsculpture is more complex. In these groups sparsely distributed,
large tubercles are found on a background of small tubercles. Sometimes the large tu-
bercles are situated as separate groups (Fig. 19). Sometimes the tubercles are modiﬁ ed
to form conical protrusions (Harpyia, Stauropus, Peridea graeseri). In Harpyia, coni-
cal protrusions become larger towards the apical and lateral sides of the head (Fig. 20).
In 4th and 5th instars the microsculpture is usually the same as in the preceding in-
star, but is expressed much more distinctly. In some genera the microsculpture changes
from homogeneously tubercled to heterogeneously tubercled (Pheosia, Lophontosia,
Allodonta, Pterostoma sinica, P. griseum). In Gonoclostera the microsculpture chang-
es in 4th instar. In the background, which is the same as in 3rd instar (Fig. 15), there are
large tubercles located in crateriform depressions (Fig. 21). Sometimes in 5th instar
a simpliﬁ cation of the microsculpture takes place. For example, the head surface of
Cerura in 1st instar is homogeneously knobby, in 2nd to 4th instars it changes to hetero-
geneously knobby. In 5th instar it becomes deeply wrinkled in the area of the epicranial
suture, whereas towards the lateral surface it appears as homogeneously oval tubercles
concentrated in somewhat separated groups.
In general, the larval head microsculpture in Notodontidae may be more or less dis-
tinct. In the majority of genera it is clearly visible (Notodonta, Pheosia, Pheosiopsis). In
Harpyia, Uropyia, and Furcula it is most prominent. In other notodontids (Pygaerinae)
it is less prominent. Overall the most prominent microsculpture appears in 4th and 5th
instars, while in 1st to 3rd instars it is usually poorly developed.
The head microsculpture is not completely homogeneous on the surface of the head
capsule. In most cases it is smoother in the frontal area and around the epicranial su-
ture (Fig. 22). In the vicinity of the stemmata and genae the microsculpture is ﬁ ner,
consisting of homogeneous, densely arranged tubercles (Fig. 23) and it may be more
(Stauropus) or less expressed than elsewhere. Usually the microsculpture in 3rd to 5th
instars consists of the microsculpture characteristic for the stemmatal and genal areas
on which larger tubercles appear (Fig. 24).
Main Transformation Types of Head Microsculpture During Larval Development
There are 12 types of head microsculpture that occur in different instars of noto dontids:
1. Head surface smooth, unmodiﬁ ed in all instars (Phalera).
2. Head surface smooth in 1st and 2nd instars, becoming weakly wrinkled with weak-
ly developed cells in 3rd to 5th instars (Clostera, Micromelalopha).
3. Head surface smooth in 1st and 2nd instars, developing weak cells in 3rd to 5th in-
4. Main head surface smooth, but with pits in the apical part of the head and along
the sides in 1st and 2nd instars, becoming pitted in 3rd to 5th instars (Gluphisia).
5. Head surface mostly smooth but wrinkled in the apical area and laterally in 1st and
nd instars, developing somewhat expressed tubercles on a background of
densely interlaced ﬁ brae in 3rd instar, producing tubercles in crateriform depres-
sions on a background of dense ﬁ brae in 4th and 5th instars (Gonoclostera).
Nota lepid. 34 (1): 11 – 28
Figs 7 – 12. Larval head surface of Notodontidae. 7. 2nd instar Furcula biﬁ da. 8. 3rd instar Phalera
bucephala. 9. 3rd instar Micromelalopha troglodyta. 10. 3rd instar Clostera anachoreta. 11. 3rd instar
Leucodonta bicoloria. 12. 3rd instar Gluphisia crenata. Scale bar 9, 11, 12 (100 μ); 7, 8, 10 (10 μ).
18 Dolinskaya: Larval head microsculpture in Palaearctic Notodontidae
6a. Head surface smooth in 1st and 2nd instars, with large tubercles expressed only
аlong the head margins and sparsely distributed on a smooth background in 3rd
to 5th instars (Ptilodon).
6b. Head surface smooth in 1st and 2nd instars, with large tubercles expressed only
аlong the head margins and sparsely distributed on a background of small tu-
bercles in 3rd to 5th instars (Allodonta).
6c. Head surface smooth in 1st and 2nd instars, with large tubercles sparsely distributed
on a background of densely implicated ﬁ brae in 3rd to 5th instars (Lophontosia).
7. Head surface smooth in 1st and 2nd instars, with homogeneous microsculpture of
small tubercles in 3rd instar, and larger tubercles on a homogeneous back-
ground of small tubercles in 4th and 5th instars (Pterostoma sinica, P. griseum).
In P. palpina knobby microsculpture appears in 2nd instar.
8. Head surface smooth in 1st instar, with homogeneous microsculpture of small tu-
bercles in 2nd instar, and larger tubercles on a background of homogeneous
small tubercles in 3rd to 5th instars (Euhampsonia, Drymonia, Notodonta, Pe ri -
dea, Nerice, Lophocosma, Pheosiopsis, Shaka, Lophontosia, Hagapteryx, Epo-
donta, Cnethodonta, Stauropus, Pheosia). There are some variations on this
pattern. In Pheosia and Stauropus 3rd instar microsculpture is homogeneously
tubercled, becoming heterogeneously tubercled only in 4th instar.
9а. Head surface smooth in 1st instar, with larger tubercles on a background of homo-
geneous small tubercles in 2nd to 5th instars (Uropyia, Harpyia, Fentonia).
9b. Head surface smooth in 1st and 2nd instars, with larger tubercles on a background
of homogeneous small tubercles in 3rd to 5th instars (Semidonta, Spatalia). In
Semidonta 3rd instar the microsculpture is only weakly expressed, becoming
more distinctly visible only in 4th and 5th instars. In Spatalia the microsculpture
appears only in 3rd instar in two species, S. argentina and S. dives, while in
S. doerriesi it is homogeneously tubercled in 2nd instar and becomes heteroge-
neously tubercled in 3rd instar.
10. Head surface smooth in 1st instar, with very small, weakly expressed, homogene-
ously dispersed tubercles in 2nd to 5th instars (Dicranura).
11. Weakly expressed homogeneous microsculpture of small tubercles without trans-
formation in all instars (Pygaera).
12. Distinctly expressed homogeneous microsculpture of small tubercles in 1st instar,
with larger tubercles on a background of homogeneous small tubercles in 2nd
to 5th instars (Cerura, Furcula).
Comparative Morphology of Larval Head Microsculpture in Other Families
It should be noted that head microsculpture is also variable in the studied outgroups,
as is the case in Notodontidae. It is also modiﬁ ed depending on the instar. In La sio-
campidae that were examined, the head is smooth in 1st instar (Euthrix potatoria,
Gastropacha quercifolia) while later (2nd to 5th instars) it develops microtrichiae (Fig.
25). One species of Lymantriinae has a smooth head in all instars (Teia dubia) while
another (Arctornis l-nigrum) has microtrichia in 4th and 5th instars as in Lasiocampidae.
Nota lepid. 34 (1): 11 – 28
Figs 13 – 18. Larval head surface of 3rd instar Notodontidae. 13. Gluphisia crenata. 14. Pheosia gnoma.
15. Gonoclostera timoniorum. 16. Ptilodon saturate hoegei. 17. Lophocosma atriplaga. 18. Notodonta
dembowskii. Scale bar 17, 18 (100 μ); 13 – 16 (10 μ).
20 Dolinskaya: Larval head microsculpture in Palaearctic Notodontidae
In examined Arctiinae the microsculpture is either smooth in all instars (Rhyparioides
amurensis) or slightly wrinkled in 4th and 5th instars (Chionarctia nivea). The head
surface of the examined Noctuidae is smooth in 1st through 4th instars. In 5th instar it is
weakly wrinkled in Calocasia coryli and Egira conspicillaris, with a few tubercles in
E. conspicillaris (Fig. 26).
Transformation of Head Microsculpture in Different Instars and Comparison
with Representatives from Outgroups
Miller (1991) deﬁ ned six character states relating to the head surface in Notodontidae:
“head surface mostly smooth, with ﬁ ne creases” (0); “head surface rugose with ru-
gosities in clusters” (1); “head surface covered with pits” (2); “rugosities extremely
small” (3); “head surface smooth, glassy” (4); and “head surface spiculate” (5). Miller
interpretated these characters as multistate nonadditive, where deﬁ nite numbers (in
brackets) were assigned to each state. He recognised (0) as plesiomorphic and (5) as
autapomorphic for Notodontidae.
In many cases I have used the same character states as Miller, but sometimes it
was necessary to redeﬁ ne them. Thus, Miller’s “head surface mostly smooth, with ﬁ ne
creases” is here described as weakly wrinkled sculpture, and “head surface rugose
with rugosities in clusters” is divided into recognisable types of tuberculous micro-
sculptures: large tubercles situated separately and randomly on a background of small
tubercles, or concentrated into separate groups.
According to Miller’s interpretation “head surface mostly smooth, with ﬁ ne creas-
es” is the plesiomorphic state for the family. Pitted sculpture is a derivation of the
granulate head type. “Head surface smooth, glassy” in his rank of transformations pre-
cedes “head surface spiculate”, which is the apomorphic state for the family. In his
paper on Dioptinae, Miller (2009) noted that “a smooth, almost glassy head surface” is
“a derived condition” compared with the sculpture of the so-called “pebblelike projec-
Following Miller (1991) I also consider that a spiculate sculpture is the apomorphic
state for the family. However, I suggest that a weakly wrinkled microsculpture is in-
termediate between smooth, treated here as plesiomorphic, and tuberculous, which is
secondary and more specialized in the morphological series of transformations. I also
consider that a pitted sculpture is a derivation of the smooth head type. These notions
are developed below.
Smooth head microsculpture in 1st instar was found in the majority of notodontid
subfamilies that were examined. Since this character is also found in the examined
outgroups (Erebidae: Lymantriinae and Arctiinae; Noctuidae; and Lasiocampidae),
this state (smooth head) is considered plesiomorphic relative to other states. An excep-
tion is Phalera, which differs by having a smooth head microsculpture in each larval
instar (Fig. 8), as is the case in some representatives of the outgroup (Lymantriinae,
Nota lepid. 34 (1): 11 – 28
Figs 19 – 24. Larval head surface of Notodontidae. 19. 3rd instar Furcula biﬁ da. 20. 3rd instar Harpyia mil-
hauseri. 21. 4th – 5th instars Gonoclostera timoniorum. 22. 4th – 5th instars Spatalia argentina. 23. 4th – 5th
instars Furcula bicuspis. 24. 4th – 5th instars Pterostoma griseum. Scale bar 19, 21 – 24 (100 μ); 20 (10 μ).
22 Dolinskaya: Larval head microsculpture in Palaearctic Notodontidae
Arctiinae, Noctuidae) which have a smooth head from 1st to 3rd instar or sometimes
until the last instar.
Weakly wrinkled head microsculpture in Notodontidae is present in larvae of four
genera only. In Clostera, Micromelalopha, and Leucodonta such microsculpture ap-
pears in larvae of 3rd to 5th instars, while their larvae have a smooth head in 1st and
2nd instars. Since the representatives of the outgroups (Erebidae: Arctiinae; Noctuidae)
have this kind of sculpture only in 4th and/or 5th instars, possessing smooth larval head
in 1st to 3rd or 4th instars, the wrinkled microsculpture is considered a derived state
(apomorphic). In Gonoclostera this type of microsculpture is present only in 1st and 2nd
instars (Fig. 3) and is displaced by tubercles in 3rd to 5th instars. Wrinkled microsculp-
ture is considered plesiomorphic relative to tuberculous microsculpture.
Tuberculous head microsculpture was found in the majority of the examined noto-
dontid genera in 2nd to 5th instars, with them having a smooth head in 1st instar. In
Pygaera such microsculpture is present in all instars. Tuberculous microsculpture is
found in the examined outgroups only in 5th instar in Egira conspicillaris (Noctuidae),
where it is present only in the apical part of the head, whereas in 1st to 4th instars the
microsculpture is smooth. Since the majority of notodontid genera have this kind of
micro sculpture in 2nd to 5th instars, possessing smooth larval head in 1st instar, and
since this character is absent in the outgroups (except for Egira conspicillaris), the
tu ber culous microsculpture is considered a derived state. There are several kinds of
tu berculous head microsculptures in the studied larvae. The microsculpture with al-
most indistinct, very small, homogeneous tubercles (Pygaera, Dicranura; Fig. 28)
is considered plesiomorphic. The microsculpture with large, uniform tubercles on a
smooth or ﬁ brous surface (Ptilodon; Fig. 16) either with tubercles situated randomly
on a background of small tubercles (Spatalia, Lophocosma; Fig. 17), or concentrated
in separate groups (Notodonta, Euhampsonia, Epodonta, and some others; Fig. 18) is
presumably derived. A further derived state would be a microsculpture with conical
projections in groups (Harpyia, Uropyia, Stauropus; Fig. 20).
Pitted head microsculpture in 1st instar is found only in Gluphisia. It is also present
in 2nd instar and is found in the apical part of the head only, while the rest of the head
capsule remains smooth. In later instars the pitted microsculpture extends across the
remaining head surface (Figs 12, 13). Based on these observations it appears that such
sculpture is secondary relative to the smooth one, and, probably, derived from the
latter. The pitted microsculpture has not been observed in any other notodontids or
outgroups. However, according to Miller (1991) such sculpture is characteristic for
Datana ministra (Drury) of the American genus Datana Walker, Therefore, pitted mi-
crosculpture in Gluphisia, on the basis of this study, can be considered as a synapo-
morphy for these two genera.
Summarizing, the smooth head microsculpture in larvae of different noctuoid fami-
lies could be treated as a plesiomorphic state. According to a comparative morphologi-
cal study, the general tendency in its evolutionary transformation within Notodontidae
(same as in other families of Noctuoidea) is changing towards a sculptured surface, at
ﬁ rst with small-sized and ﬁ nally large-sized sculptural elements.
Nota lepid. 34 (1): 11 – 28
Figs 25 – 29. Larval head surface. 25. 4th – 5th instars Euthrix potatoria. 26. 5th instar Egira conspicil-
laris. 27. 4th – 5th instars Uropyia meticulodina. 28. Dicranura ulmi. 29. Oral surface of left mandible of
2nd instar Leucodonta bicoloria. Scale bar 26 (100 μ); 25, 27 – 29 (10 μ).
24 Dolinskaya: Larval head microsculpture in Palaearctic Notodontidae
Phylogenetic Implications Within Genera
Head microsculpture can be also used for resolving phylogenetic relationships with-
in genera. This is important as there has been insufﬁ cient work on this problem in
Notodontidae. In Peridea the most complex microsculpture is recorded in P. graeseri,
where tubercles are modiﬁ ed into conical projections. In P. anceps, P. lativitta, P. elzet,
P. gigantea, P. oberthueri, and P. moltrechti the microsculpture appears as clearly
visible oval tubercles. Based on these results it appears that P. graeseri possesses the
most derived state of this character. The peculiarities of the head pattern (Dolinskaya
2009) corroborate this hypothesis.
In Pterostoma, in addition to the morphological similarity of the various spe-
cies, the transformation of the microsculpture changes from a simple to a complex
one. Thus, in 2nd instars of P. palpina the microsculpture is already homogeneously
tubercled, whereas in P. griseum and P. sinicum it is smooth. In 3rd instar, the micro-
sculpture of P. palpina becomes heterogeneously tubercled and distinct. In P. gri-
seum and P. sinicum the microsculpture changes only to homogeneously tubercled.
In P. griseum the sculpture is very well expressed and distinctive while in P. sinicum
it is poorly expressed. In 4th and 5th instars the microsculpture of P. palpina and
P. griseum becomes similar: large tubercles concentrated in distinct groups on a
background of small ones, while in P. sinicum these groups are poorly expressed.
Based on these results it appears that P. palpina possesses the more derived states
of these characters, while P. sinicum has the less derived ones. P. griseum has an
intermediate position between these two species. It should be noted that the mor-
phological characters of the pupa support this hypothesis. The sculpture of the cre-
master in P. griseum and P. palpina is very similar, and it is different in P. sinicum
(Dolinskaya 1984, 1989).
In Ptilodon in addition to the morphological similarity of some species, a transfor-
mation of the microsculpture from simple to conical takes place. In P. capucina and
P. saturate hoegei weakly expressed tubercles are randomly located on a smooth sur-
face, while in P. cucullina these tubercles are placed on a background of densely im-
plicated ﬁ brae. In P. ladislai the latter are concentrated in weakly expressed groups.
Based on these observations it appears that P. ladislai possesses a more derived states
of this character, and P. capucina and P. saturate hoegei the less derived one, while
P. cucullina appears to be intermediate.
In Spatalia, a transformation of microsculpture from simple to complex takes
place. In 2nd instar of S. doerriesi the microsculpture appears weakly expressed and
homogeneously tubercled while in S. argentina and S. dives it remains smooth. In
3rd to 5th instars the microsculpture of S. argentina appears as small and large oval
tubercles arranged on a smooth background. In S. dives and S. doerriesi the micro-
sculpture is more complex: on a background of small tubercles there are randomly
located medium-sized and large tubercles, the latter with plicated edges and some-
times concentrated in poorly deﬁ ned groups. Based on these observations, S. doer-
riesi appears to possess a more derived state of this character than S. argentina, with
S. dives being intermediate.
Nota lepid. 34 (1): 11 – 28
The Importance of Larval Head Microsculpture for the Classiﬁ cation
At present the classiﬁ cation of Notodontidae is in need of improvement. In the sys-
tems proposed by different authors the number of subfamilies within the family,
as well as the number and generic composition of subfamilies remains uncertain.
The most recent classiﬁ cations are those of Tikhomirov (1981), Miller (1991), and
Schintlmeister (2008). Below I discuss the implications of the results presented here-
in for the classiﬁ cation of the family.
Slightly visible head microsculpture in each larval instar is present in only eight
genera. It is either smooth (Phalera), weakly wrinkled, looking like weakly expressed
cells (Clostera, Micromelalopha, Leucodonta), very small, densely located homo-
geneous tubercles (Dicranura, Pygaera), tubercles placed in crateriform depres-
sions (Gonoclostera) or pitted (Gluphisia). Concerning the genera Clostera, Micro-
melalopha, Gonoclostera, and Pygaera these data coincide with the classiﬁ cations of
the three above-mentioned authors, placing them into the subfamily Pygaerinae. This
hypothesis is also supported by an examination of the larval mandible (Dolinskaya
In Pygaerinae a gradual development of the microsculpture takes place from
weakly wrinkled to small-tubercled. In Micromelalopha and Clostera the head mi-
crosculpture remains weakly formed. In 1st and 2nd instars it is smooth, while in 3rd to
5th instars it is already weakly wrinkled or appears as indistinct homogeneous cells
(Figs 9, 10). In Gonoclostera in 1st and 2nd instars the microsculpture is also smooth;
however, apically and also partly laterally it becomes wrinkled (Fig. 3). In 3rd instar
the surface appears as small, densely implicated ﬁ brae on a background of weak-
ly expressed, very small tubercles (Fig. 15). In 4th and 5th instars large tubercles are
formed, arranged in crateriform depressions on a background of smaller tubercles
(Fig. 21). This is likely a transitional state from wrinkled to tubercled microsculpture.
In Pygaera the microsculpture appears as very small, weakly expressed, densely situ-
ated homogeneous tubercles in all instars. This character is considered apomorphic
relative to the above-mentioned representatives of subfamily Pygaerinae. Based on
these results, as well as an examination of the outgroups, it would appear that the
most primitive microsculpture is found in Clostera and Micromelalopha. Head mi-
crosculpture of Pygaera and Gonoclostera is more derived.
In Dicranura in 1st instar the head microsculpture is smooth, and in 2nd to 5th instars
it appears as very small, weakly expressed, densely situated homogeneous tubercles
as in Pygaera (Fig. 28). This character along with mandibular structure (Dolinskaya
2008) supports the close afﬁ nity of Dicranura to Pygaerinae. Schintlmeister (2008)
placed Dicranura in Dicranurinae together with Stauropus, Cnethodonta, and
Harpyia. However, later in a personal communication he agreed with the conclusions
published by Dolinskaya (2008) and acknowledged the need to clarify the taxonomic
status of Dicranura.
Leucodonta needs additional examination. Tikhomirov (1981) and Schintlmeister
(2008) placed it within Notodontinae. The results of this study show that the head
26 Dolinskaya: Larval head microsculpture in Palaearctic Notodontidae
microsculpture of Leucodonta is similar to that of Clostera and Micromelalopha. It
shares the same character states as the above genera, namely having smooth micro-
sculpture in 1st and 2nd instars, while in 3rd to 5th instars it appears as almost indistinct
cells (Fig. 11). These observations support the placement of Leucodonta in Pygaerinae.
This hypothesis is augmented by the peculiarities of mandibular structures. A detailed
examination of the mandibles using SEM showed the presence of a denticulated man-
dibular edge and a large retinaculum in 2nd instar (Fig. 29) as is found in Clostera,
Gonoclostera, and Micromelalopha (Dolinskaya 2008).
The position of the genus Gluphisia within Notodontidae is anomalous. Tikhomirov
(1981) placed it within Notodontinae. Miller (1991) also included Gluphisia within this
subfamily, placing it in Notodontini along with Cerura and Furcula. Schintlmeister
(2008) assigned Gluphisia to Pygaerinae. The peculiarities of the pupa and larval
mandibles (Dolinskaya 1986, 1989, 2008) corroborate the opinion of Schintlmeister
(2008). However, the characters of the larval head surface are not concordant with this
hypothesis. Head microsculpture in Gluphisia is represented as distinct pits (Figs 12,
13). Such microsculpture is not found in either the ingroup or the examined outgroups.
According to Miller (1991), pitted head microsculpture is characteristic for Datana
(Phalerinae). Therefore it is necessary to carry out a more detailed examination of oth-
er taxa to clarify the systematic position of this genus.
The uncertainty of the systematic position of Phalera remains. Tikhomirov (1981)
included this genus in Notodontinae. Miller (1991) placed Phalera in Phalerinae along
with Datana, Peridea, and Euhampsonia. Schintlmeister (2008) also included this ge-
nus in Phalerinae together with Phalerodonta. Unfortunately I was not able to study
the larval head microsculpture of Phalerodonta. The results of my studies show that in
Phalera the head microsculpture is smooth in each instar (Fig. 8). The same type of mi-
crosculpture in 4th and 5th instars is absent in the rest of the notodontids examined. On
the other hand, Miller (1991) noted the presence of ‘extremely smooth, almost glassy,
head surface’ in some Josia Hübner and Cyanotricha Prout (Dioptinae), and Didugua
Druce (Nystaleinae). At a later time Miller (2009) noted that this head surface of the ﬁ -
nal instar is unique for Josiini (Dioptinae). In addition to that Miller (1991), following
Gardner (1943), noted that Phalera larvae have a pitted surface. However, my results
do not support this hypothesis and hence the taxonomic position of this genus should
be investigated further.
The majority of the genera in the family have a clearly visible and mainly heteroge-
nously tubercled head microsculpture. Such microsculpture is present in 26 genera,
namely Euhampsonia, Cerura, Furcula, Uropyia, Harpyia, Stauropus, Cnethodonta,
Fen tonia, Neopheosia, Drymonia, Notodonta, Peridea, Nerice, Pheosia, Lophocosma,
Pheo siopsis, Shaka, Pterostoma, Ptilodon, Lophontosia, Allodonta, Hagapteryx, To ge-
pteryx, Semidonta, Epodonta, and Spatalia. Ptilodon and Allodonta differ from these
genera by having the most smoothened microsculpture. The head capsule is smooth me-
dially, with microsculpture expressed only аlong the head margins. These data partially
support the hypothesis of Schintlmeister (2008), who placed Ptilodon and Allo donta
within Ptilodontinae together with Pterostoma, Semidonta, Lophontosia, and Epo donta.
In these genera there is a tendency towards smooth head microsculpture (Table 1).
Nota lepid. 34 (1): 11 – 28
In Cerura and Furcula the head microsculpture is distinctive, being tubercled in
1st instar (Fig. 4). I treat this character state as apomorphic because most representa-
tives of the ingroup as well as all of the examined outgroups have smooth micro-
sculpture in 1st instar. These ﬁ ndings coincide with the systems of Tikhomirov (1981)
and Schintlmeister (2008), who separated these genera into the subfamily Cerurinae.
Miller (1991) included these genera within Notodontinae together with Gluphisia,
Pheosia, and Notodonta. This arrangement is not supported by the results of this
In Cerura, Furcula, Uropyia, Harpyia, and Fentonia the head microsculpture
is heterogeneously granulated in 2nd instar, with large tubercles on a background
of small tubercles (Figs 6, 27). These ﬁ ndings partly coincide with the point of
view of Schintlmeister (2008) in including Uropyia, Harpyia, and Fentonia within
Larval head microsculpture can provide useful phylogenetic information within and
between genera. Similar types of larval head surface structures support monophyly of
a number of groups of genera. Comparative morphological examinations of the larval
head microsculpture can be used to determine the direction of morphological transfor-
mations. More derived taxa have a more complex microsculpture. For more general-
ized groups smooth or similar types of the larval head capsule surface are characteris-
tic. Among these characters I have identiﬁ ed notodontid apomorphies (tubercled head
microsculpture), as well as apomorphies that are characteristic for various other taxa
in the family: 1) presence of tubercled microsculpture in 1st instar (Cerura, Furcula,
Pygaera); 2) presence of heterogeneous microsculpture in 2nd instar (Cerura, Furcula,
Uropyia, Harpyia, and Fentonia); and 3) development of conical projections on the
head (Harpyia, Uropyia, Stauropus).
It is worth noting that it is possible to determine the larval instar according to head
microsculpture. This additional diagnostic character can be used for genera and spe-
cies together with such characters as the width of the head capsule and the larval head
pattern (Dolinskaya 2009). On the basis of peculiarities of the head microsculpture it
is also possible to estimate the degree of morphological similarity of species within the
I am deeply indebted to Dr. M. G. Ponomarenko (Institute of Biology and Soil Science, Vladivostok,
Russia) for critical reading and useful remarks on this paper. I am very grateful to Dr. J. S. Miller (American
Museum of Natural History, New York City) and Dr. S. Passoa (Ohio State University, United States) for
their help with literature. I am obliged to Dr. Paul Sokoloff (Kent, UK) and Dr. Bernard Landry (Geneva)
for their correction of the English language. The author also thanks Mr. Yu. Geryak (State Natural History
Museum, Lvov, Ukraine) and Mr. A.V Zhakov (ZUCTKUM, Zaporosh’e, Ukraine) for sending live
females of Dicranura ulmi. The study was partly ﬁ nancially supported by grant of the State Fund for
Fundamental Reseaches, Ukraine (SFFR No. F40.4/043).
28 Dolinskaya: Larval head microsculpture in Palaearctic Notodontidae
Bell, T. R. 1935. A description of the notodontid moth Dudusa nobilis Walker and its early stages. – Jour-
nal of the Bombay Natural History Society 38: 134 – 136.
Dolinskaya, I. V. 1984. Chrysalid morphology of certain notodontid moth species (Lepidoptera, Noto don-
tidae). – Vestnik zoologii 4: 54 – 60. [In Russian]
Dolinskaya, I. V. 1986. Chrysalid morphology of certain notodontid moth species (Lepidoptera, Noto-
don tidae) of the fauna of the USSR. – Vestnik zoologii 2: 59 – 66. [In Russian]
Dolinskaya, I. V. 1989. Morphology of the pupae of the Notodontidae (Lepidoptera) from Russian Far
East. – Vestnik zoologii 5: 60 – 67. [In Russian]
Dolinskaya, I. V. 2008. Taxonomic variation in larval mandibular structure in Palaearctic Notodontidae
(Noctuoidea). – Nota lepidopterologica 31: 165 – 177.
Dolinskaya, I. V. 2009. Formation of pattern and diagnostic instar features of the head in caterpillars from
genus Peridea (Lepidoptera, Notodontidae). Vestnik zoologii 1: 15 – 24.
Gardner, J. C. M. 1943. Immature stages of Indian Lepidoptera. – Indian Journal of Entomology 5: 89 –
Kuznetzov, V. I. & A. A. Stekolnikov 2001. New approaches to the system of Lepidoptera of world fauna
(on the basis of the functional morphology of the abdomen). – Trudy Zoologicheskogo Instituta, St.
Petersburg, Nauka 282: 1 – 462. [In Russian]
Miller, J. S. 1991. Cladistics and classiﬁ cation of the Notodontidae (Lepidoptera: Noctuoidea) based on
larval and adult morphology. – Bulletin of the American Museum of Natural History 204: 1 – 230.
Miller, J. S. 1996. Phylogeny of the Neotropical moth tribe Josiini (Notodontidae: Dioptinae): a hidden
case of Müllerian mimicry. Zoological Journal of the Linnean Society 118: 1 – 45.
Miller, J. S. 2009a. Generic revision of the Dioptinae (Lepidoptera: Noctuoidea: Notodontidae). Part 1:
Dioptini. – Bulletin of the American Museum of Natural History 321: 1 – 674.
Miller, J. S. 2009b. Generic revision of the Dioptinae (Lepidoptera: Noctuoidea: Notodontidae). Part 2:
Josiini. – Bulletin of the American Museum of Natural History 321: 675 – 1022.
Minet, J. 1994. The Bombycoidea: Phylogeny and higher classiﬁ cation (Lepidoptera: Glossata). – Ento-
mo logica scandinavica 25: 63 – 88.
Schintlmeister, A. 2008. Palaearctic Macrolepidoptera. Vol. 1: Notodontidae. – Apollo Books, Stenstrup.
Tikhomirov, A. M. 1981. Taxonomic structure of the family Notodontidae and its position in the system
of Lepidoptera in regard to functional morphology of genitalia of species from the Far East. – Trudy
Zoo lo gicheskogo Instituta AN SSSR (Leningrad) 103: 62 – 72. [In Russian]
Zahiri, R., Kitching, I. J., Lafontaine, J. D., Mutanen, M., Kaila, L., Holloway, J. D. & Wahlberg,
N. 2011. A new molecular phylogeny offers hope for a stable family level classiﬁ cation of the
Noctuoidea (Lepidoptera). – Zoologica Scripta 40: 158 – 173.