Diversity of the Larval Cranial Setae in Palaearctic Notodontidae (Noctuoidea) and and their Taxonomic Distribution
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), Sphin...
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irk-123456789-1096672016-12-07T03:02:09Z Diversity of the Larval Cranial Setae in Palaearctic Notodontidae (Noctuoidea) and and their Taxonomic Distribution Dolinskaya, I.V. Фауна и систематика 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. С помощью сканирующего электронного микроскопа изучены щетинки головы гусениц всех возрастов 66 видов из 35 родов палеарктических хохлаток из Украины и Приморского края России. Проведено сравнение с представителями внешней группы — Lasiocampoidea (Lasiocampidae), Sphingoidea (Sphingidae) и (Erebidae: Lymantriinae, Arctiinae; Noctuidae). Рассмотрены вероятные направления преобразований в волосяном покрове головы у гусениц различных возрастов. Обсуждаются возможные апоморфные и плезиоморфные состояния признаков различных таксонов. 2013 Article Diversity of the Larval Cranial Setae in Palaearctic Notodontidae (Noctuoidea) and and their Taxonomic Distribution / I.V. Dolinskaya // Вестник зоологии. — 2013. — Т. 47, № 1. — С. 27–37. — Бібліогр.: 20 назв. — англ. 0084-5604 DOI 10.2478/vzoo-2013-0002 http://dspace.nbuv.gov.ua/handle/123456789/109667 595.786:591.342 en Вестник зоологии Інститут зоології ім. І.І. Шмальгаузена НАН України |
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Фауна и систематика Фауна и систематика Dolinskaya, I.V. Diversity of the Larval Cranial Setae in Palaearctic Notodontidae (Noctuoidea) and and their Taxonomic Distribution Вестник зоологии |
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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. |
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Dolinskaya, I.V. |
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Dolinskaya, I.V. |
| title |
Diversity of the Larval Cranial Setae in Palaearctic Notodontidae (Noctuoidea) and and their Taxonomic Distribution |
| title_short |
Diversity of the Larval Cranial Setae in Palaearctic Notodontidae (Noctuoidea) and and their Taxonomic Distribution |
| title_full |
Diversity of the Larval Cranial Setae in Palaearctic Notodontidae (Noctuoidea) and and their Taxonomic Distribution |
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Diversity of the Larval Cranial Setae in Palaearctic Notodontidae (Noctuoidea) and and their Taxonomic Distribution |
| title_full_unstemmed |
Diversity of the Larval Cranial Setae in Palaearctic Notodontidae (Noctuoidea) and and their Taxonomic Distribution |
| title_sort |
diversity of the larval cranial setae in palaearctic notodontidae (noctuoidea) and and their taxonomic distribution |
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Інститут зоології ім. І.І. Шмальгаузена НАН України |
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2013 |
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Фауна и систематика |
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| citation_txt |
Diversity of the Larval Cranial Setae in Palaearctic Notodontidae (Noctuoidea) and and their Taxonomic Distribution / I.V. Dolinskaya // Вестник зоологии. — 2013. — Т. 47, № 1. — С. 27–37. — Бібліогр.: 20 назв. — англ. |
| series |
Вестник зоологии |
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AT dolinskayaiv diversityofthelarvalcranialsetaeinpalaearcticnotodontidaenoctuoideaandandtheirtaxonomicdistribution |
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2025-07-07T23:27:54Z |
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2025-07-07T23:27:54Z |
| _version_ |
1837032669513252864 |
| fulltext |
UDC 595.786:591.342
DIVERSITY OF THE LARVAL CRANIAL SETAE
IN PALEARCTIC NOTODONTIDAE (NOCTUOIDEA)
AND THEIR TAXONOMIC DISTRIBUTION
I. V. Dolinskaya
Schmalhausen Institute of Zoology, NAS of Ukraine,
vul. B. Khmelnytskogo, 15, Kyiv, 01601 Ukraine
E-mail: dd8v @mail.ru
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.
Ke y wo 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. Îáñóæäàþòñÿ âîçìîæíûå àïîìîðôíûå è ïëåçèî-
ìîðôíûå ñîñòîÿíèÿ ïðèçíàêîâ ðàçëè÷íûõ òàêñîíîâ.
Êëþ÷åâûå ñ ëîâ à: Notodontidae, Lepidoptera, ãóñåíèöû, ìîðôîëîãèÿ, ùåòèíêè ãîëîâû, êëàñ-
ñèôèêàöèÿ, ñêàíèðóþùèé ýëåêòðîííûé ìèêðîñêîï.
Introduction
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, 2011). While
studying these structures we noted the great variety of setae on the larval head. Taking as examples the notodon-
tid caterpillars we intended to show the changes in the shape, disposition and other characters of the head setae
during the larval development.
Hinton (1946), Beck (1960), Merzheevskaya (1967) and Stehr (1987) characterized the chaetotaxy of
the lepidopteran caterpillars in details. Miller (1991) described chaetotaxy of some notodontid caterpillars of
the final instars. Earlier we analized the chaetotaxy of the notodontid first instar larval head (Dolinskaya, Pljushch,
2003). Thus, we did not incorporate the chaetotaxy of the larval head in this paper.
Material and methods
This research is based on material collected in Ukraine and Far East of Russia (Primorskii Krai). Eggs
were obtained from females captured at light. Hatched larvae were reared to pupae. The epicrania left by cater-
pillars after moulting, as well as fresh material preserved in alcohol, were studied. The epicranium was exa-
mined with a scanning electron microscope (SEM) and a binocular light microscope (MBS 9). The microsculp-
ture of the head of 1st through 5th larval instars belonging to 66 notodontid species from the following genera
were studied: Euhampsonia, Cerura, Furcula, Uropyia, Dicranura, Harpyia, Stauropus, Cnethodonta, Fentonia,
Neopheosia, Drymonia, Notodonta, Peridea, Nerice, Pheosia, Leucodonta, Lophocosma, Ellida, Pheosiopsis, Shaka,
Pterostoma, Ptilodon, Lophontosia, Hagapteryx, Togepteryx, Semidonta, Allodonta, Epodonta, Phalera, Spatalia,
Vestnik zoologii, 47(1): e-24—e-34, 2013
DOI 10.2478/vzoo-2013-0002
Unauthenticated
Download Date | 12/5/16 8:28 PM
Gluphisia, Gonoclostera, Pygaera, Clostera, and Micromelalopha. Terminology for larval structures follows Hinton
(1946) and Stehr (1987). The taxonomic arrangement of these genera follows Schintlmeister (2008).
In order to clarify the character states and polarity within Notodontidae, representatives of related fam-
ilies belonging to Lasiocampoidea, Sphingoidea, as well as other members of Noctuoidea (Kuznetzov,
Stekolnikov, 2001; Zahiri and other, 2011, 2012), arre used as outgroup taxa. The following species were stud-
ied: Euthrix potatoria Linnaeus, Gastropacha quercifolia Linnaeus (Lasiocampidae), Agrius convolvuli Linnaeus,
Smerinthus planus Walker (Sphingidae), Teia dubia Tauscher, Arctornis l-nigrum Müller (Erebidae: Lymantriinae),
Rhyparioides amurensis Bremer, Chionarctia nivea Mėnėtriės, Phragmatobia amurensis Seitz (Erebidae: Arctiinae)
and Calocasia coryli Linnaeus (Noctuidae).
Results
Comparative Morphology of Larval Head Setae
The taxa and characters examined are listed in table 1. Covering of the head setae
varies from instar to instar. Head setae of the notodontid larvae in the 1st to 2nd instars
are most distinct.
First instar of notodontid caterpillars have mostly or only primary setae. These setae
are typical for many species. Primary setae are simple, mainly firm, with smooth sculp-
ture, thinned towards their apices (Fentonia, Furcula, Gluphisia and others, fig. 1). In the
genera Stauropus, Cnethodonta and Harpyia, the setae have small mace-shaped bulge in
the apical part (figs 2, 3). In the genus Phalera the setae are plumose (fig. 4). In the genus
Uropyia, sculpture of the primary setae looks like well developed apically rounded spines
(figs 5—7).
Primary setae are mainly short, their length not expanded of the half of width of head
capsule (figs 1, 2). Species of some genera have very long, hairlike setae many times longer
than the width of head capsule (Uropyia, Ptilodon, Hexafrenum, Allodonta, fig. 8).
The 1st instar caterpillars posess numerous, small, secondary setae only in one genus
Harpyia (figs 9, 10).
Many genera have setae of the 2nd instar identical to that of the preceding instar.
However, sometimes the larval head of some genera is covered with numerous se-
condary setae (Stauropus, Dicranura, Phalera, Pygaera and Clostera). These setae are either
very short, slightly visible (Dicranura, Stauropus, figs 11, 12, 15), moderately short
(Pygaera, fig. 13) or very long (Phalera, Clostera, figs 4, 14). Primary setae are longer
and stronger than secondary setae and more distinct.
In the 3rd to 5th instars no significant transformations are observed, except species
with short setae which become shorter, thinner and less visible on the head (Peridea,
Lophontosia, Lophocosma and others); at the same time in the species with long setae,
they are very distinct (Phalera, Clostera and others). In the representatives of some gen-
era (Stauropus, Harpyia), where the sculpture looks conical the setae replaced to the tuber-
cles (fig. 16).
The setae are mostly located on the distinct pinacula. It must be noted that the pri-
mary setae have larger pinacula than secondary setae (fig. 17). In some genera the setae
are located on the chalaza (Uropyia, Pygaera, fig. 6).
Main types of larval setae
The setae occuring in different instars of notodontids can be grouped in 13 types as
follows.
1. Primary setae in all instars (Euhampsonia, Cerura, Furcula, Uropyia, Cnethodonta,
Fentonia, Drymonia, Notodonta, Peridea, Pheosia, Nerice, Leucodonta, Lophocosma,
Ellida, Pheosiopsis, Shaka, Pterostoma, Ptilodon, Lophontosia, Hagapteryx, Togepteryx,
Semidonta, Allodonta, Epodonta, Spatalia, Gluphisia, Gonoclostera, Micromelalopha).
2. Primary setae in 1st instar, secondary setae appearing in 2nd to 5th instars
(Stauropus, Dicranura, Phalera, Pygaera and Clostera).
e-25 I. V. Dolinskaya
Unauthenticated
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e-26Diversity of the Larval Cranial Setae Palaearctic Notodontidae (Noctuoidea)...
Ta b l e 1. Character states of the larval cranial setae of the Palaearctic Notodontidae
Ò à á ëèö à 1. Ïðèçíàêè ùåòèíîê ãîëîâû ãóñåíèö ïàëåàðêòè÷åñêèõ õîõëàòîê
Notodontidae
Euhampsonia cristata (Butler) P, À, H P, À, H P, À, H
Euhampsonia splendida (Oberthür) P, A, H P, A, H P, A, H
Cerura erminea (Esper) P, A, H P, A, H P, A, H
Furcula furcula (Clerck) P, A, H P, A, H P, A, H
Furcula bicuspis (Borkhausen) P, A, H P, A, H P, A, H
Furcula bifida (Brahm) P, A, H P, A, H P, A, H
Uropyia meticulodina (Oberthür) P, A, L P, A, L P, A, L
Dicranura ulmi (Denis et Schiffermüller) P, A, VH S, A, VH S, A, VH
Harpyia milhauseri (Fabricius) S, PCL, VH S, PCL, VH S, A, VH
Harpyia umbrosa (Staudinger) S, PCL, VH S, PCL, VH S, A, VH
Stauropus fagi (Linnaeus) P, PCL, VH S, PCL, VH S, A, VH
Stauropus basalis Moore P, PCL, VH S, PCL, VH S, A, VH
Cnethodonta grisescens Staudinger PCL, VH PCL, VH P, A, VH
Fentonia ocypete (Bremer) P, A, H P, A, H P, A, H
Neopheosia mandschurica (Oberthür) P, A, H - -
Drymonia dodonaea (Denis et Schiffermüller) P, A, H P, A, H P, A, H
Notodonta torva (Hübner) P, A, H P, A, H P, A, H
Notodonta dromedarius (Linnaeus) P, A, H P, A, H P, A, H
Notodonta dembowskii Oberthür P, A, H P, A, H P, A, H
Notodonta tritophus phoebe (Siebert) P, A, H P, A, H P, A, H
Notodonta ziczac (Linnaeus) P, A, H P, A, H P, A, H
Peridea anceps (Goeze) P, A, H P, A, H P, A, H
Peridea lativitta (Wileman) P, A, H P, A, H P, A, H
Peridea elzet Kiriakoff P, A, H P, A, H P, A, H
Peridea graeseri (Staudinger) P, A, H P, A, H P, A, H
Peridea gigantea (Butler) P, A, H P, A, H P, A, H
Peridea oberthueri (Staudinger) P, A, H P, A, H P, A, H
Peridea moltrechti (Oberthür) P, A, H P, A, H P, A, H
Nerice leechi Staudinger P, A, H P, A, H P, A, H
Nerice davidi Oberthür P, A, H P, A, H P, A, H
Pheosia tremula (Clerck) P, A, H P, A, H P, A, H
Pheosia grummi (Christoph) P, A, H - -
Pheosia gnoma (Fabricius) P, A, H P, A, H P, A, H
Pheosia rimosa Packard P, A, H P, A, H P, A, H
Leucodonta bicoloria (Denis et Schiffermüller) P, A, H P, A, H P, A, H
Lophocosma atriplaga Staudinger P, A, H P, A, H P, A, H
Ellida branickii (Oberthür) P, A, H - -
Pheosiopsis cinerea (Butler) P, A, H P, A, H P, A, H
Shaka atrovittatus (Bremer) P, A, H P, A, H P, A, H
Pterostoma palpina (Clerck) P, A, H P, A, H P, A, H
Pterostoma gigantina Staudinger P, A, H P, A, H P, A, H
Pterostoma griseum (Bremer) P, A, H P, A, H P, A, H
Ptilodon capucina (Linnaeus) P, A, L P, A, L P, A, L
Ptilodon saturate hoegei (Graeser) P, A, L P, A, L P, A, L
Ptilodon cucullina (Denis et Schiffermüller) P, A, L P, A, L P, A, L
Ptilodon ladislai (Oberthür) P, A, L P, A, L P, A, L
Lophontosia cuculus (Staudinger) P, A, H P, A, H P, A, H
Hagapteryx admirabilis (Staudinger) P, A, H P, A, H P, A, H
Togepteryx velutina (Oberthür) P, A, H P, A, H P, A, H
Semidonta biloba (Oberthür) P, A, H P, A, H P, A, H
Allodonta plebeja (Oberthür) P, A, L P, A, L P, A, L
Allodonta leucodera (Staudinger) P, A, L P, A, L P, A, L
Epodonta lineata (Oberthür) P, A, H P, A, H P, A, H
Phalera bucephala (Linnaeus) P, B, L S, B, L S, B, L
Spatalia argentina (Denis et Schiffermüller) P, A, H P, A, H P, A, H
Spatalia doerriesi Graeser P, A, H P, A, H P, A, H
Species Instar 1 Instar 2 Instars 3—5
Unauthenticated
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3. Primary setae in 1st and 2nd instars with small mace-shaped bulge in the apical
part (Stauropus, Cnethodonta, Harpyia).
4. Numerous secondary setae in all instars (Harpyia).
5. Secondary setae long (Phalera, Clostera).
6. Secondary setae very short (Harpyia, Stauropus, Dicranura, Pygaera). In Pygaera,
setae become very short in 4—5th instars.
7. Simple setae in all instars (Euhampsonia, Cerura, Furcula, Fentonia, Drymonia,
Notodonta, Peridea, Pheosia, Nerice, Leucodonta, Lophocosma, Ellida, Pheosiopsis, Shaka,
Pterostoma, Ptilodon, Lophontosia, Hagapteryx, Togepteryx, Semidonta, Allodonta, Epodonta,
Spatalia, Gluphisia, Gonoclostera, Micromelalopha).
8. Plumose setae in all instars (Phalera).
9. Sculpture of the setae in all instars with well developed spines rounded apically
(Uropyia).
10. Setae moderately short in 1st and 2nd instars, becoming smaller and thinner in
3rd to 5th instars (Euhampsonia, Cerura, Furcula, Fentonia, Drymonia, Notodonta,
Peridea, Nerice, Pheosia, Leucodonta, Lophocosma, Ellida, Lophontosia, Hagapteryx,
Togepteryx, Semidonta, Epodonta, Spatalia, Gluphisia, Gonoclostera, Pygaera,
Micromelalopha).
11. In all instars, primary and secondary setae very long, many times longer than
width of head capsule (Ptilodon, Hexafrenum, Allodonta, Phalera, Clostera).
12. In all instars, primary and secondary setae very short, poorly visible (Harpyia,
Stauropus, Cnethodonta, Dicranura).
13. In different instars setae are located on chalaza (Uropyia, Pygaera).
e-27 I. V. Dolinskaya
P – primary setae; S – secondary setae; À – simple setae; B – plumose setae; L – long setae; H –
short setae; VH – very short setae; PCL – primary setae with mace-shaped bulge in the apical part, T – tre-
elike setae; C – hairlike setae.
Table 1 continuation
Îêîí÷àíèå òàáë. 1
Species Instar 1 Instar 2 Instars 3—5
Spatalia plusiotis Oberthür P, A, H - -
Spatalia dives Oberthür P, A, H P, A, H P, A, H
Gluphisia crenata (Esper) P, A, H P, A, H P, A, VH
Gonoclostera timoniorum (Bremer) P, A, H P, A, H P, A, H
Pygaera timon (Hübner) P, A, H S, A, H S, A, VH
Clostera albosigma curtuloides (Erschoff) P, C, L S, C, L S, C, L
Clostera pigra (Hufnagel) P, C, L S, C, L S, C, L
Clostera anachoreta (Denis et Schiffermüller) P, C, L S, C, L S, C, L
Clostera anastomosis (Linnaeus) P, C, L S, C, L S, C, L
Micromelalopha troglodyta (Graeser) P, A, H P, A, H P, A, H
Lasiocampidae
Euthrix potatoria (Linnaeus) S, C, L S, C, L S, C, L
Gastropacha quercifolia (Linnaeus) S, C, L S, C, L S, C, L
Sphingidae
Agrius convolvuli (Linnaeus) - - S, A, H
Smerinthus planus Walker - S, T, H S, A, H
Erebidae: Lymantrinae
Teia dubia (Tauscher) - - S, B, L
Arctornis l-nigrum (Müller) - S, B, L S, B, L
Erebidae: Arctiinae
Rhyparioides amurensis (Bremer) S, B, L S, B, L S, B, L
Chionarctia nivea (Mėnėtriės) - S, C, L S, C, L
Phragmatobia amurensis Seitz P, B, L - S, B, L
Noctuidae
Calocasia coryli (Linnaeus) P, B, H P, B, H S, B, H
Unauthenticated
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e-28Diversity of the Larval Cranial Setae Palaearctic Notodontidae (Noctuoidea)...
Fig. 1—6. Larval head of Notodontidae: 1 – head capsule of 1st instar Fentonia ocypete; 2 – head capsule of
1st instar Stauropus fagi; 3 – setae with small mace-shaped bulge of 1st instar Cnethodonta grisescens; 4 – plumose
setae of 5th instar Phalera bucephala; 5 – head capsule of 2nd instar Uropyia meticulodina; 6 – cranial seta
of 2nd instar U. meticulodina. Scale bars: 1, 2, 4, 5, 6 – 100 μ; 3 – 10 μ.
Ðèñ. 1—6. Ãîëîâà ãóñåíèö Notodontidae: 1 – ãîëîâíàÿ êàïñóëà 1-ãî âîçðàñòà Fentonia ocypete; 2 – ãîëîâ-
íàÿ êàïñóëà 1-ãî âîçðàñòà Stauropus fagi; 3 – ùåòèíêà ñ íåáîëüøèì áóëàâîâèäíûì óòîëùåíèåì ó ãóñå-
íèö 1-ãî âîçðàñòà Cnethodonta grisescens; 4 – ïåðèñòûå ùåòèíêè ó ãóñåíèö 5-ãî âîçðàñòà Phalera bucep-
hala; 5 – ãîëîâíàÿ êàïñóëà 2-ãî âîçðàñòà Uropyia meticulodina; 6 – êðàíèàëüíàÿ ùåòèíêà ãóñåíèö 2-ãî
âîçðàñòà U. meticulodina. Ìàñøòàáíûå ëèíåéêè: 1, 2, 4, 5, 6 – 100 ìêì; 3 – 10 ìêì.
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e-29 I. V. Dolinskaya
Fig. 7—12. Larval head of Notodontidae: 7 – fragment of seta of 2nd instar Uropyia meticulodina; 8 – head
capsule of 1st instar Ptilodon saturate hoegei; 9 – head capsule of 1st instar Harpyia umbrosa; 10 – setae with
small mace-shaped bulge of 1st instar Harpyia umbrosa; 11 – head capsule of 2nd instar Dicranura ulmi; 12 –
head capsule of 4th- instar Dicranura ulmi. Scale bars: 11 – 1000 μ; 8, 9, 10, 12 – 100 μ; 7 – 10 μ.
Ðèñ. 7—12. Ãîëîâà ãóñåíèö Notodontidae: 7 – ôðàãìåíò ùåòèíêè 2-ãî âîçðàñòà Uropyia meticulodina; 8 –
ãîëîâíàÿ êàïñóëà 1-ãî âîçðàñòà Ptilodon saturate hoegei; 9 – ãîëîâíàÿ êàïñóëà 1-ãî âîçðàñòà Harpyia umbrosa;
10 – ùåòèíêè ñ íåáîëüøèì áóëàâîâèäíûì óòîëùåíèåì ó ãóñåíèö 1-ãî âîçðàñòà Harpyia umbrosa; 11 –
ãîëîâíàÿ êàïñóëà 2-ãî âîçðàñòà Dicranura ulmi;12 – ãîëîâíàÿ êàïñóëà 4-ãî âîçðàñòà Dicranura ulmi.
Ìàñøòàáíûå ëèíåéêè: 11 – 1000 ìêì; 8, 9, 10, 12 – 100 ìêì; 7 – 10 ìêì.
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e-30Diversity of the Larval Cranial Setae Palaearctic Notodontidae (Noctuoidea)...
Fig. 13—18. Larval head: 13 – head capsule of 2nd instar Pygaera timon; 14 – head capsule of 2nd instar Clostera
anastomosis; 15 – head capsule of 2nd instar Stauropus basalis; 16 – head capsule of 4th- instar Stauropus fagi;
17 – pinacula on the setae on the head capsule of 4th- instar Clostera pigra; 18 – head capsule of 2nd instar
Euthrix potatoria (Lasiocampidae). Scale bars: 13, 15, 18 – 100 μ; 14, 16, 17 – 10 μ.
Ðèñ. 13—18. Ãîëîâà ãóñåíèö: 13 – ãîëîâíàÿ êàïñóëà 2-ãî âîçðàñòà Pygaera timon; 14 – ãîëîâíàÿ êàïñó-
ëà 2-ãî âîçðàñòà Clostera anastomosis; 15 – ãîëîâíàÿ êàïñóëà 2-ãî âîçðàñòà Stauropus basalis; 16 – ãîëîâ-
íàÿ êàïñóëà 4-ãî âîçðàñòà Stauropus fagi; 17 – òåêè ùåòèíîê íà ãîëîâíîé êàïñóëå 4-ãî âîçðàñòà Clostera
pigra; 18 – ãîëîâíàÿ êàïñóëà 2-ãî âîçðàñòà Euthrix potatoria (Lasiocampidae). Ìàñøòàáíûå ëèíåéêè: 13,
15, 18 – 100 ìêì; 14, 16, 17 – 10 ìêì.
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e-31 I. V. Dolinskaya
Fig. 19—24. Larval head: 19—22 – Sphingidae; 23—24 – Erebidae: Lymantrinae; 19 – head capsule of 5th
instar Agrius convolvuli; 20 – head capsule of 5th instar Smerinthus planus; 21 – head capsule of 2nd instar
Smerinthus planus; 22 – setae of 2nd instar Smerinthus planus; 23 – plumose setae of 4th instar Teia dubia;
24 – plumose setae of 2nd instar Arctornis l-nigrum. Scale bars: 19—21 – 100 μ); 22—24 – 10 μ.
Ðèñ. 19—24. Ãîëîâà ãóñåíèö: 19—22 – Sphingidae; 23—24 – Erebidae: Lymantrinae; 19 – ãîëîâíàÿ êàï-
ñóëà 5-ãî âîçðàñòà Agrius convolvuli; 20 – ãîëîâíàÿ êàïñóëà 5-ãî âîçðàñòà Smerinthus planus; 21 – ãîëîâ-
íàÿ êàïñóëà 2-ãî âîçðàñòà Smerinthus planus; 22 – ùåòèíêè 2-ãî âîçðàñòà Smerinthus planus; 23 – ïåðè-
ñòûå ùåòèíêè 4-ãî âîçðàñòà Teia dubia; 24 – ïåðèñòûå ùåòèíêè 2-ãî âîçðàñòà Arctornis l-nigrum.
Ìàñøòàáíûå ëèíåéêè: 19—21 – 100 ìêì; 22—24 – 10 ìêì.
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e-32Diversity of the Larval Cranial Setae Palaearctic Notodontidae (Noctuoidea)...
Fig. 25—29. Larval head: 25 – head capsule of 4th instar Arctornis l-nigrum (Erebidae: Lymantrinae); 26 –
plumose setae of 1st instar Rhyparioides amurensis (Erebidae: Arctiinae); 27—29 – Calocasia coryli (Noctuidae);
27 – plumose setae of 1st instar; 28 – plumose setae of 2nd instar; 29 – head capsule of 4th instar. Scale
bars: 29 – 1000 μ; 25 – 100 μ; 26—28 – 10 μ.
Ðèñ. 25—29. Ãîëîâà ãóñåíèö: 25 – ãîëîâíàÿ êàïñóëà 4-ãî âîçðàñòà Arctornis l-nigrum (Erebidae: Lymantrinae);
26 – ïåðèñòûå ùåòèíêè 1-ãî âîçðàñòà Rhyparioides amurensis (Erebidae: Arctiinae); 27—29 – Calocasia coryli
(Noctuidae); 27 – ïåðèñòûå ùåòèíêè 1-ãî âîçðàñòà; 28 – ïåðèñòûå ùåòèíêè 2-ãî âîçðàñòà; 29 – ãîëîâ-
íàÿ êàïñóëà 4-ãî âîçðàñòà. Ìàñøòàáíûå ëèíåéêè: 29 – 1000 ìêì; 25 – 100 ìêì; 26—28 – 10 ìêì.
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Comparative Morphology of Larval Head Setae in Other Families (table 1)
In Lasiocampidae that were examined, the secondary setae are present in all instars,
however in 2nd instar they become more numerous (Euthrix potatoria, Gastropacha
quercifolia). Secondary setae are hairlike and long (fig. 18).
One species of Sphingidae (Agrius convolvuli) has the numerous, short and simple
secondary setae in 5th instar (fig. 19). Another species (Smerinthus planus) has treelike
secondary setae in 2nd instar and simple secondary setae in 5th instar (fig. 20—22).
Two species of Erebidae: Lymantriinae (Teia dubia – 4th to 5th instars and Arctornis
l-nigrum – 2nd to 5th instars) have numerous, secondary, plumose, long setae (fig. 23—25).
In examined Erebidae: Arctiinae primary and secondary setae are plumose
(Rhyparioides amurensis, Phragmatobia amurensis) or hairlike (Chionarctia nivea). In
Rhyparioides amurensis secondary setae there are from 1st instar (fig. 26).
The setae of the examined Erebidae: Noctuidae (Calocasia coryli) are primary, plumose
and short in 1st and 2nd instars. In 3rd to 5th instarsthere are a few secondary plumose
setae (fig. 27—29).
Discussion
The numerous secondary setae are widespread among the lepidopteran caterpillars
(Packard, 1895; Kozhanchikov, 1950; Beck, 1960; Merzheevskaya, 1988; Singh, 1991;
Danner et al., 1998; Lemaire, Minet, 1999; Kitching, Rawlins, 1999 and others). From
this, Packard (1895) and then Miller (1991) consider the presence the secondary setae
to be a primitive character of notodontids.
Among all of studied genera of notodontids the numerous secondary setae are typ-
ical for six genera – Clostera, Pygaera, Dicranura, Phalera, Stauropus and Harpyia. In
addition, the Phalera and Clostera have long setae and at the others have short setae.
In general, for many notodontids are typical primary simple setae in all instars. Only
the species of some genera have their own peculiarities.
Genera Harpyia, Stauropus and Cnethodonta have the primary setae in 1st and 2nd
instars with the mace-shaped bulge. This form of setae is absent in the rest of the notodon-
tids and outgroup species examined. Therefore, Such a character is considered derived
state in relation to the typical primary simple setae most genera notodontids. Miller (1991)
included only these three genera into the tribe Stauropini of the subfamily Heterocampinae.
In the same time the presence of numerous secondary setae of Harpyia and Stauropus
and their absence at Cnethodonta can be testify as derived states of this character for
Cnethodonta.
Genus Phalera is unique in the family. The plumose setae are found only in this genus.
The same setae are found in the outgroup species. Therefore, such a character is con-
sidered to be a generalized state in relation to the typical primary simple setae most
notodontid genera . In the 2nd to 5th instars Phalera have numerous secondary setae and
this character is considered generalized state for the genus. These data are also support-
ed by examination of the larval head microsculpture (Dolinskaya, 2011), where the smooth
head microsculpture is recorded only in this genus of the Notodontidae, and this state
is considered plesiomorphic relative to other states.
There are no disagreements among researchers as to taxonomic composition of the
subfamily Pygaerinae. Tikhomirov (1981) and Schintlmeister (2008) included the gen-
era Clostera, Pygaera, Gonoclostera and Micromelalopha in here. The mandibular char-
acters and head sculpture of the caterpillars (Dolinskaya, 2008, 2011) also support this
placement. However, the characters of the cranial larval setae show rather puzzle distri-
bution pattern. Of all these genera, the secondary setae appear only in Clostera and
Pygaera L. Species of the genus Clostera have long setae, whereas in Pygaera the setae
are short. As the numerous secondary setae are typical for the outgroup species, we con-
sider this character state to be plesiomorphic for these genera.
e-33 I. V. Dolinskaya
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Genus Dicranura has numerous, simply, short secondary setae. Same setae are typ-
ical for Pygaera. Our previous studies (Dolinskaya, 2008, 2011) show strong similarity
of Dicranura to the Pygaerinae and especially with the genus Pygaera.
Summarizing these data, I consider the larval cranial setae to provide useful phylo-
genetic information about relationships within and between genera.
I am very grateful to Dr. Z. S. Gershenzon (I. I. Schmalhausen Institute of Zoology, Kyiv, Ukraine) for
helpful comments on the manuscript. This work is a part of project supported by the State Fund for
Fundamental Researches, Ukraine (project SFFR No F40.4/043) and the Russian Foundation for Basic Research
(project No 11—04—09454).
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Received 16 November 2012
Accepted 21 November 2012
e-34Diversity of the Larval Cranial Setae Palaearctic Notodontidae (Noctuoidea)...
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