ZoologtcalJoumal ofthe Ltnnean Soriety, 70: 55- 101. With 49 figures
Srptelllber 1980
An integumental anatomy for the butterfly
Glaucopsyche lygdamus (Lepidoptera:
Lycaenidae):a morphological
terminology and homology
J O H N T. SORENSEN
Department of Entomologtcal Sciences,
University of California, Berkeley, Cal$ 94720, U . S . A .
Arcrprd f o r publication February 1980
An iiiteguinentnl anatomy for the lycaenid butterfly Glaucopsyche lygdumus is presented. Comparisons
with other lepidopteran taxa are made to rectify the homology of parts and contrast anatomical
clivergencrs within the Lycaenidae. A general terminology based on Snodgrass is given, to replace
inany of the sprcialized a n d often synonymous terms restricted to the Lepidoptera. Many common
~uiato~nical
svni~nyinsare also given. Several reinterpretations of the anatomv a n d homologv of
various integumental regions are discussed. A previously unreported cuticular anon+
on
;~bdoniinalterguni 2 of male Polyommatinae (Downey’s area) is described. Thc l b l l o ~
ing 11ew o r
i i c ~ w l ycoinbineti terrns are used : postgenal-occipital area, postgenal-occipital protuberance, dorsal
teinpordl gulci, postantennal projections, pronotal projection, infraepisternal-basisternal plate,
pai-ac~)xal-~narginopIeural
sulci, dorsal epimeral sulci, ventral epimeral sulci, secondary coxal sulci,
veiitral subrostal-radial process, lateral secondary sclerite a n d Downey’s area.
K E Y W O R D S : - intrgunient - exoskeleton - anatomy - morphology - trrminology - honlologv
Lepidoptera - Pdpilionoidea - Lycaenidae- Gluucopsyche lygdamus.
-
CONTENTS
. . .
Illtl.oducrlr)n
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Mrthods arid m,iterials
. . . . . .
Material studied and taxonomic position
. . . . . . . .
Terininologv
Re\uli\ and discussion
. . . . . . .
Head
. . . . . . . . . .
Cervix
. . . . . . . . . .
Prothorn( ic notum
. . . . . .
PI-othor-acicsternopleural region
. .
Mesothorac ic norum . . . . . .
Mesi)thoracic-sternopleural region
. .
Metathoracic notum
. . . . . .
Metathoracic sternopleural region
. .
. . . . . . .
Prothor-acit legs
Mesothoi-acic legs
. . . . . .
Metathoracic legs . . . . . . .
Mesothoracic wings and wing base sclerites
MetathoracicwinRsand wingbasesclerites
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56
J. T. SORENSEN
Pregenitalic abdomen
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Genitalic abdomen and genitalia
Male
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Fernale
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Acknowledgements
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References
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INTRODUCTION
There are several complete descriptive morphologies for the integument of the
Lepidoptera, but few for the Papilionoidea and none for the Lycaenidae. This
work provides an anatomical description of the gross features of the integument
of Glaucopsyche lygdamus (Doubleday). More importantly, it attempts to establish a
consistent and general morphological terminology for the Lepidoptera, to
replace the confusing array of specialized, often synonymous, terms currently
used without regard for homology. Finally, it explores both intra- and interordinal homologies, and points out structural specializations within family,
superfamily and order, wherever possible. Extensive use is made of
morphological investigations that concentrate on specific anatomical regions,
noting their variation and evolution throughout the order.
METHODS A N D MATERIALS
Specimens for cuticular dissection were preserved in ethanol or dried, then
usually treated in hot 10% potassium hydroxide solution. Those used for
checking muscular attachments were preserved in Kahle’s fixative. Scanning
electron microscopy was used to check the surface features of Downey’s area on
abdominal tergum 2 of G. lygdamus males.
Morphological homologies were largely established by comparison of adult
structures among taxa. This method appears to have no appreciable
disadvantage compared to other methods, such as tracing innervation and
muscle insertions, or following ontogenetic development in the embryo, since
they may sometimes be of questionable reliability (Matsuda, 1976; Rempel,
1975). In some cases, where comparisons were too confusing, tracings of muscle
insertions were employed.
Material studied and taxonomic position
About 60 Glaucopsyche lygdamus of both sexes were studied for gross
integumental features. Downey’s area on abdominal tergum 2 was examined in
12 additional males.
Eliot ( 1973) provides a tentative higher classification of the Lycaenidae, and
divides the family into eight subfamilies and 33 tribes. Using his hierarchy,
Glaucopsyche is placed in the subfamily Polyommatinae, tribe Polyommatini,
section Glaucopsyche. Eliot states that the Polyommatini is very large and lacks
major intratribal groupings; he divides the tribe into 30 sections.
0ther lycaenids examined, and their intrafamily positions are :
Subfamily Polyommatinae, tribe Polyommatini :
Glaucopsyche section : Glaucopsychepiasus (Boisduval),Maculinea alcon (Denis
8c Schiffermuller), Maculinea arion (Linnaeus),Philotes sonorensis (Felder &
Felder), Scolitantides orion ( Pallas).
INTEGUMENTAL ANATOMY OF GLAUCOPSYCHE
i7
Polyornrnatus section : Plebejus icaroides (Boisduval),Hemiargus isola (Reakirt).
Eueres section: Eueres comyntus (Godart).
Lycaenopsis section : Celustrina argiolus (Linnaeus).
Subfamily Lycaeninae :
Lycaena section : Lycaena xanthoides (Boisduval), Lycaena phlaeas (Linnaeus).
Subfamily Theclinae, tribe Eumaeini :
Eumaeus section : Satyrium edwardsii (Grote 8c Robinson), Strymon melinus
Hiibner.
Sorensen ( 197 2) examined the members of the Glaucopsyche section of the
Polyommatini listed above, to establish generic morphological variation of the
integument, and implied that the group is heterogeneous.
Other lepidopterans examined for comparative purposes are :
Eriocraniidae: undetermined sp.
Hepialidae: Hepialus sp.
Incurvariidae: Adela sp.
Scythridae : ,$reniscythris brachypteris Powell
Ethmiidae: Ethmia Charybdis Powell
Pyralidae: Pyralis sp.
Geometridae: Palaeacrita sp., Ennomos sp.
Saturniidae : Eacles sp., Hyalophora cecropia (Linnaeus)
Sphingidae : Hyles lineata (Fabricius), Sphinx sp.
Noctuidae : Heliothis zea (Boddie),Catocala sp., Trichoplusia sp.
Arctiidae : Halisidota sp., Hyphantria cunea Drury, Apantesis sp.
Hesperiidae : Poanes hobomok (Harris), Hesperia comma (Linnaeus), Epargyreus
clarus (Cramer)
Papilionidae : Papilio rutulus Lucas, Papilio multicaudata Kirby, Battus philenor
(Linnaeus1
Pieridae : Colias eurytheme Boisduval, Pieris rapae (Linnaeus), Phoebis sennae
(Linnaeusj, Eurema lisa Boisduval8c LeConte, Nathalis iole Boisduval
Danaidae : Danaus plexippus (Linnaeus)
Satyridae : Oeneis nevadensis (Felder 8c Felder), Cercyonis pegala (Fabricius),
Euptychia cymela (Cramer)
Nymphalidae : Nymphalis califmica Boisduval, Speyeria atlanta (Edwards),
Euphydryas phaeton (Drury),Limenitis arthemis (Drury).
Among other butterflies, the most extensive comparisons were made with
Nymphalis calfornica, a putatively generalized nymphalid, to assess structures that
are highly modified in the Lycaenidae (e.g., those of the head). Unless otherwise
stated, all descriptions throughout this work are of male Glaucopsyche lygdamus.
Terminology
Terminology follows Snodgrass ( 1935) wherever feasible, since it is widely
accepted for other insects. Deviations are adopted for the following reasons : ( 1)
morphological interpretations for specific regions have been refined by
subsequent workers and Snodgrass’s terms are n o longer appropriate; (2)
specialized structures exist either without apparent homologues elsewhere o r
whose homology remains unclear; or (3)no Snodgrass term could be found for a
generalized structure.
58
J. T. SORENSEN
Alternative sources for terms are indicated in the sections, and synonyms and
their sources are often given. The following new or newly combined terms are
applied : postgenal-occipital area, postgenal-occipital protuberance, dorsal
temporal sulci, postantennal projections (see head); pronotal projection (see
prothoracic notum) ; infraepisternal-basisternalplate, paracoxal-marginopleural
sulci, dorsal epimeral sulci, ventral epimeral sulci (see mesothoracic
sternopleural region) ; secondary coxal sulci (see metathoracic leg); ventral
subcostal-radial process (see mesothoracic wing); lateral secondary sclerite,
Downey’s area (see pregenitalic abdomen).
Dicke 8c Howe (1978) have attempted a drastic revision of pterygote
morphological terminology, but their terms have not been employed here.
The term ‘suture’ has been erroneously used in insect morphology. Snodgrass
(1960)discusses its use, along with that of sulcus, noting that most insect ‘sutures’
are actually sulci. He states that suture should be reserved for true, primitive intersegmental fusion lines. Matsuda ( 1965) agrees, but confusingly retains suture
in broad usage. I use sulcus exclusively, since the exact nature of most true
sutures (sensu strict01 is unknown, or disputed.
Terms are printed in capital letters to facilitate location. Because the genitalia
are exceedingly important in taxonomy and also present an exceptional problem
in recognizing homologies, specialized taxonomic (morphologically
meaningless) terms are included in italics in that section and within parentheses
in the figures.
RESULTS AND DISCUSSION
Head (Figs 1-81
Some features shared among several insect orders are altered in the
Lepidoptera, obscuring homologies, due to the unique siphon mechanism.
Moreover, lycaenids sometimes exhibit fusion or atrophy of head features
beyond the norm for the order.
Snodgrass’s ( 1960)head terminology is used, with modifications after Duporte
( 1956)and Ehrlich ( 1958a)for unique lepidopteran structures.
The COMPOUND EYES are dorsally emarginate around the antenna1 bases, and
bear sparse interommatidial setae. OCELLI are absent. Internally, the compound
eyes have an OCULAR DIAPHRAGM perforated by the central OCULAR FORAMEN
(Duporte, 1959). The diaphragm separates the internal eye from the head
haemocoel while the optic nerve passes through the foramen. The diaphragm is
slightly sclerotized peripherally, forming the OCULAR DIAPHRAGM SCLERITES,
which remain intact after maceration.
Anteriorly, a large FRONTOCLYPEAL SCLERITE (Duporte, 1956) represents the
fused frons and clypeus; the FRONTOCLYPEAL (EPISTOMAL) SULCUS is absent. The
ANTERIOR TENTORIAL PITS are located laterally along the frontoclypeal sclerite.
The relative proportions of the frons and clypeus in this fused plate may vary
among papilionoid groups, and various authors have used different definitions
for their demarcation. Srivastava (1957) considers the anterior tentorial pits to
mark the frons-clypeus fusion line in Papilio, citing cibarial and pharyngeal
dilator muscle insertions as evidence. In contrast, Eassa (1963a) states that the
clypeus enlarges greatly during pupal metamorphosis in Pieris, restricting the
diminished frons to the plate’s dorsal interantennal region. Eassa ( 1963b) shows
INTEGUMENTAL ANATOMY OF GLAUCOPSYCHE
59
DORSAL TEMPORAL SULCUS
TRANSFRONTAL SULLUS
\
POSTANTENNAL PROJECTION ___zI
SCAPE
PARAOCULAR AREA
OF GENA
C I SCUMANTENNAL SULCUS
COMPOUND EYE
FRONTOCLYPEAL SCLERITE
LATEROFACIAL SULCUS
ANTERIOR
TENTORIAL P I T
LATEROFACIAL SULCLS
MANDIBULAR RUDIMENT
PLEUROSTOMAL
SUBGENAL SULCUS
SUBGENA
-----
CLYPEOLABRAL SULCUS
L A B I A L PALPUS
GALEA
1
LATEROFACIAL SULCUS
COMPOUND EYE
PARAOCULAR AREA OF GENA
PEDICEL
CI RCUMANTENNAL SULCUS
VERTEX
POSTANTENNAL PROJECTION
POSTGENAL-OCCIPITAL AREA
TEMPORAL SULCUS
TRANSFRONTAL SULCUS
DORSAL TEMPORAL SLlLCUS
POSTGENAL-OCC I P I T A L
PROTUBERANCE
2
Figures 1, 2 . Head. Fig. I . Anterior view; flagella removed, left labial palput ~-mmoved.gcilc<ie
revered. Fig. 2 . Dorsal view; flagella removed, left labial palpus removed.
pump
dilator
insertions
verifying this interpretation. A roughened
1958a1, traversing this plate ventrally, is absent in
Glaucopsyche but present in Nymphalis.
An extremely weak TRANSFRONTAL SULCUS bridges the antenna1 bases,
demarcating the frontoclypeal sclerite dorsally. In Glaucopsyche the sulcus is
usually present as a lightly sclerotized interantennal band, allowing the
frontoclypeal sclerite to merge with the head’s dorsal surface, and in some
individuals it is developed laterally, with internal ridges that are continuous with
the internal ridge of the circumantennal sulcus. Ehrlich ( 1958b) considers the
FRONTOCLYPEAL BAND (Ehrlich
J. T. SORENSEN
60
GALEA
RUDIMENTARY
MAXILLARY PALPUS-
!I
‘I
x
/
FRONTOCLYPEAL SCLERl T E
I I O R TENTORIAL PI1
LATEROFACIAL SULCUS
PLEUROSTOMAL SUBGENAL SULCUS
TRANSOCC 1P I T A L SULCUS
SUBGENA
HYPOSTOMAL SUBGENAL SU
PARALABIAL SULCUS
LABIUM
4
Figures 3, 4. Fig. 3. Excised left maxilla base, ventral view (anterior to top, midline to left); galea
severed. Fig. 4 . Head, ventral view; left labial palpus removed, right galea severed, left galea
removed at basal articulation.
TEMPORAL SULCUS
FORAMEN MAGNUM
ARTICULATION FOSSA
TRANSOCCIPITAL SULCUS
POSTGENAL-OCCIPITAL AREA
TENTORIAL BRIDGE
HYPOSTONAL SUBGENAL SULCUS
PARALABIAL SULCUS
Figure 5. Head, posterior view; left labial palpus removed, cervical membrane removed, flagella
severed.
transfrontal sulcus to be an interantennal reinforcement, noting that its strength
is positively correlated with interantennal distance in papilionoids.
A CIRCUMANTENNAL SULCUS circumscribes the base of each antenna. Its
internal ridge is formed by the inflected margins of the surface plates, and
anteriorly produces a flange-like ANTENNIFER that serves as the monocondylic
articulation of the scape. The posterior margins of the antenna1 socket are
elevated, forming POSTANTENNAL PROJECTIONS,bearing numerous long setae.
Each circumantennal sulcus is anteroventrally continuous with a
INTEGUMENTAL ANATOMY O F GLAUCOPSYCHE
FLAGELLUM
PEDICEL
POSTANTENNAL PROJECTION
SCAPE
WTUBERANCE
COMPOUND EYE
POSTGENAL-OCCIPITAL
AREA
I
FRONTOCLYPEAL S C L E R I T E
,
GALEA
/‘
L A B I A L PALPUS
C I RCUXANTENNAL
RIDGE
TENTOR! A! ARMS
HYPOPHARYNGEAL
PEDICEL’
8
FLAG~LLUM
Figures 6-8. Fig. 6. Head, right lateral view; flagella severed, galeae severed. Fig. 7 . As Fig. 6. light
compound eve removed. Fig. 8. Antenna, ventral view.
LATEROFACIAL SULCUS (Duporte, 1956). The latter demarcates the frontoclypeal
sclerite laterally, separating it from the narrow PARAOCULAR AREA OF THE GENA
(Michener, 1944a). The laterofacial sulci produce strong internal ridges
reinforcing the anterior margins of the compound eye.
Madden (1944) and Srivastava (1957) use Snodgrass’s (1935) terms ocular
sulcus (circumocular sulcus : Snodgrass, 1960) and ocular sclerite for the
laterofacial sulci and paraocular areas, respectively. These terms are avoided here
because of the doubtful homologies between these structures in papilionoids and
those defined by Snodgrass ( 1935); neither structure completely encircles the eye
in butterflies. Furthermore, Ehrlich (1960) and Miller (197 1) depict structures in
the Hesperioidea that are more probably equivalent to the circumocular sulcus
62
J. T.SORENSEN
and sclerite, and are distinct from the laterofacial sulcus and paraocular area.
The possible circumocular sulcus and sclerite may be secondary in the
Hesperioidea, and are apparently absent in the Papilionoidea.
The homology of the laterofacial sulci is unclear and requires further study.
Duporte ( 1956) believes that these represent the composite fusion of the
frontogenal and the clypeogenal sulci, but this does not appear to account for
the range of variation encountered in the Lepidoptera. At least three types of
variation occur. In type I (i.e., Manduca: Madden, 1944, Duporte, 1956; E a c h :
Michener, 1952; Danaus: Duporte, 1956, Ehrlich, 1958a),these sulci begin at the
circumantennal sulci and run parallel with the anterior margin of the eye to just
above the anterior tentorial pits, where they curve ventromedially to reach these
pits. The dorsal paraocular area is continuous with the lower gena ventrally. In
type 11 (i.e., Papilio: Srivastava, 1957; Epargyreus: Ehrlich, 1960) the sulcus
resembles type I, but branches. The ventromedial branch reaches the anterior
tentorial pits as in type I, while a ventrolateral branch continues to parallel the
eye, ending near its posteroventral corner. This separates the paraocular area
and lower gena ventrally. Type 111 h e . , Calinaga, Lycaena: Ehrlich, 1958b;
Glaucopysche, Nymphalis: present data) resembles type I1 except that the
ventromedial branch to the tentorial pits is obliterated. The paraocular area and
gena are separated as in type 11, but the frontoclypeal sclerite is continuous with
the lower gena ventrolaterally.
The CLYPEOLABRAL SULCUS separates the frontoclypeal sclerite ventrally from
the narrow transverse JABRUM. Srivastava ( 1957) considers this sulcus to be
absent in Papilio, with fusion of the labrum and frontoclypeal sclerite. In
Glaucopsyche the protuberant frontoclypeal sclerite causes both the clypeolabral
sulcus and labrum to be visible in ventral view only.
Interpretation of the clypeolabral sulcus is confused in the literature. Ehrlich
( 1958a, 1960) and Miller ( 197 1) erroneously consider this sulcus to reach the
anterior tentorial pits, defining the frontoclypeal sclerite ventrolaterally, in
Danaus and various hesperiids. A clypeolabral sulcus/anterior tentorial pit
association occurs in Manduca (Duporte, 1956) and Smerinthus (Duporte, 19461,
because the pits lie immediately lateral to the labrum. Duporte (1956),
considering Danaus, where the pits and labrum are not immediately adjacent,
believes that a ventral extension of the laterofacial (clypeogenal) sulcus serves as
the tentorial pit/clypeolabral sulcus connective. This interpretation follows
Duporte’s (1946) theory that the anterior tentorial pits lie primitively on the
frontogenal sulcus in insects. I t also incorporates Duporte & Bigelow’s (1952)
concept that the gena, among higher insects, expands ventrally to flank the
clypeus and creates a clypeogenal sulcus.
In contrast, Srivastava ( 1957 1 considers that the pleurostomal subgenal sulcus
connects dorsally with the tentorial pits, representing the clypeogenal inflection
in Papilio. This follows Snodgrass’s (1935) concept that these pits lie primitively
on the epistomal-subgenal sulcal system.
Neither Duporte’s nor Snodgrass’s concept for the primitive occurrence of the
anterior tentorial pits is substantiated, but Duporte’s theory seems more
plausible in view of the variation within the Lepidoptera.
Srivastava (1957)argues against Duporte’s (1956)use of clypeogenal sulcus for
the inflection between the tentorial pits and the corners of the labrum, stating
that this sulcus extends below the labrum. This is not in conflict with Duporte &
INTEGUMENTAL ANATOMY OF GLAUCOPSYCHE.
63
Bigelow’s ( 1952) theory of a ventral extension of the gena and, in fact, nullifies
Srivastava’s application of pleurostomal sulcus, following Snodgrass ( 1935 : 128).
The latter defines the PLEUROSTOMAL SUBGENAL SULCUS to occur proximal to the
mandible, not anterior and anteroventral to it as Srivastava implies for Papilio, and
present data indicate for Glaucopsyche and Nymphalis.
A MANDIBULAR RUDIMENT represents the atrophied mandible in papilionoids
and is fused with the gena. In Nymphalis the mandibular rudiment is protuberant,
strongly sclerotized and roughened, while in Glaucopsyche it is much weaker,
lacking contrast to the surrounding cuticle, and may be obliterated in some
individuals. The pleurostomal sulcus is obliterated in the fusion of the mandibular
rudiment with the gena in Glaucopsyche. In Nymphalis and other papilionoids where
the mandibular rudiment is more apparent, the pleurostomal sulcus may be very
weakly present anteriorly, but becomes obliterated posteriorly in the
mandibular-genal fusion.
The head’s ventral surface forms a deep, high-roofed PROBOSCIDIAL FOSSA for
storage of the coiled galeal siphon. This fossa’s lateral walls are formed by
inflection of the fused subgena and gena. A large triangular LABIUM forms the
roof of the fossa and is entirely sclerotized in Glaucopsyche, as is typical of
lycaenids, nymphalids and libytheids (Ehrlich, 1958b). The labium is fused with
the subgena anteriorly and posterolaterally, and bears an anterior flexible peg
that fits between the bases of the galeae. The labium bears 3-segmented LABIAL
PALPI that pivot via monocondylic articulations at the rear of their. sockets.
The maxillae flank the labium anterolaterally but are separated from it by
narrow membranous areas. They are fused to the subgena and labium
posteriorly. Each MAXILLA has a CARDO, STIPES, GALEA and LACINIA. The latter
has atrophied, but may be present along the distal edge of the maxilla. The stipes
has a RUDIMENTARY MAXILLARY PALPUS represented as a heavily sclerotized ridge
in Glaucopsyche and an elongated nipple-like protuberance in Nymphalis. The
galeae are elongate and interlock to form the feeding siphon. Their bases are
partially membranous and invaginated, allowing flexion of the siphon. The
functional morphology of the mouthparts is described elsewhere (Kellogg, 1897 ;
Philpott, 1927; Schmitt, 1938; Pradham & Aren, 1941; Eastham 8c Eassa, 1955;
Srivastava, 1956, 1957; Matsuda, 1965). Mouthpart evolution within the order is
treated by Kristensen ( 1968a).
The lateral surface of the head is vertically divided by a broad, shallow
inflection in Glaucopsyche. Ehrlich (1958a, 1960) and Miller (197 1 ) call this the
TRANSOCCIPITAL SULCUS o r band. This may represent an OCCIPITAL SULCUS, as
occurs in the Zeugloptera (Issiki, 1931; Hannemann, 1956) and the Dacnonypha
(Kristensen 1968b, c). In the former, Issiki (1931) states the occipital sulcus ends
ventrally anteriad of the posterior articulation of the mandible, hence complying
with Snodgrass’s ( 1935: 128) definition. This is also true in the Neopseustidae
(Kristensen, 196813: fig. 6), but in the Eriocraniidae the sulcus is ventrally
obliterated (Kristensen, 1968a). Among some higher Lepidoptera the occipital
sulcus may be totally obliterated; Madden (1944) and Michener (1952) do not
report it in sphingids o r saturniids. An inflected structure matching Ehrlich’s
( 1958a) description of Danaus or the current description, however, occurs among
some noctuids, arctiids, geometrids, hesperiids and papilionids (present data).
When present, this inflection is usually weak or partially obliterated and may be
associated ventrally with the hypostomal subgenal sulcus. I t is uncertain
64
J. T.SORENSEN
whether, among the higher Lepidoptera, this is homologous with the occipital
sulcus of primitive forms or merely represents a secondary feature.
The posterior region of the head is pierced by a FORAMEN MAGNUM divided by
a TENTORIALBRIDGE into a dorsal orifice (alaforamen: Srivastava, 19571, and a
ventral orifice (neuraforamen: Srivastava, 1957). The alimentary tract and the
central nervous system pass through these, respectively. The tentorial bridge
bears CERVICAL ARTICULATION FOSSAE laterally. The POSTERIOR TENTORIAL PITS
are located ventrolateral to these, at the junction of the tentorial bridge and the
tentorial arms.
The POSTOCCIPUTborders the foramen magnum dorsally. The POSTOCCIPITAL
SULCUS runs dorsally from the posterior tentorial pits to encompass the
postocciput, and produce a strong internal ridge. Kristensen (1968~)
believes this
sulcus extends ventrally from the pits in the Eriocraniidae, but this interpretation
disagrees with Snodgrass’s ( 1935)restriction of the postoccipital sulcus to dorsad
of the pits. The sulci below these pits should represent the HYPOSTOMAL
SUBGENAL SULCI, which demarcate the subgena of the posterior region of the
head, and run ventrally from the pits to obliteration near the posteroventral
corner of the eye in Gluucopsyche. Among other papilionoids, these sulci may be
very weak or partially obliterated, as in Nymphalis, making detection difficult.
The posterior margin of the labium produces dorsolateral extensions and
forms the ventral and ventrolateral borders of the foramen magnum. These
labial extensions are delimited from the subgena by the PARALABIAL SULCI
(Ehrlich, 1958a).
The TEMPORAL SULCI (Michener, 1952, not Ehrlich, 1958a) branch from the
ventral portion of the postoccipital sulcus and run anterodorsally to obliteration
posterolateral to the antennal sockets in Glaucopsyche. In Nymphalis, the temporal
sulci reach the antennal sockets laterally. These sulci form strong internal ridges,
and have also been called postgenal sulci (Madden, 1944),occipitopostgenal sulci
(Srivastava, 19571, and paratemporal sulci (Ehrlich, 1958a). Hannemann (1956)
considers these structures to be the postoccipital sulci in the Zeugloptera, but
Klemm (1966)and Kristensen (1968~)
dispute this.
The homology and derivation of the temporal sulci are unclear but they occur
throughout most Lepidoptera and are reported or depicted in association with
the postoccipital sulci in the Mecoptera (Issiki, 1933; Ferris, 1942: fig. 231, and
primitive Trichoptera (Klemm, 1966). It is possible that the temporal sulci, at
least among lepidopterans, may have a composite origin, with their posterior
part derived from the postoccipital sulci. Kristensen (1968~)
speculates that in the
Eriocraniidae much of the temporal sulci (and the dorsal part of his ‘postfrontal’
sulcus) may be derived from a fusion of the postinterocellar sulcus with the
dorsal section of the occipital sulcus. This is plausible since among the higher
Lepidoptera the lateral ocelli, if present, lie at the anterior end of the temporal
sulci, posterior to the antenna base (present data). In primitive forms they occur
anterior to the postinterocellar sulci (Issiki, 1931; Kristensen, 1968~).The
derivation of the latter is unknown.
The DORSAL TEMPORAL SULCI are closely associated dorsally with the
postoccipital sulci. The latter are weakened dorsally, but their distinction from
the former is usually apparent. In lycaenids the dorsal temporal sulci show two
distinct paths, in contrast to the nymphalids. In Gluucopsyche they merge dorsally,
forming a strong internal ridge, and usually become obliterated on the top of the
INTEGUMENTAL ANATOMY OF GI.AUCOPSYCHE
65
head. In some individuals, however, they separate past the usual obliteration
point and extend weakly to the posteromedial edge of the antenna1 socket. In
Nymphalis, these sulci separate and run anterolaterally, paralleling the temporal
sulci. Ehrlich (1958a) erroneously refers to these sulci as the temporal sulci in
Danaus.
The homology of the dorsal temporal sulci is uncertain. They may be restricted
to the Papilionoidea, although they are absent in some papilionids (Papilio:
Srivastava, 1957) and pierids (Colzas: present data). Their unique form in the
lycaenids compared to nymphalids and danaids (see ‘paratemporal’ sulcus of
Ehrlich, 1958a: figs 2, 4) probably results from the specialized shape of the
posterodorsal aspect of the head in lycaenids (see below).
There is disagreement in the literature concerning the topographical areas of
the dorsal, lateral and posterior regions of the head. Most authors denote such
areas apparentlv without any explicit rationale. I follow Snodgrass (19601, who
considers the posterior and lateral areas of the head to be of parietal derivation
and defines their regions in relation to adjacent sulci.
The area between the eye, postoccipital sulcus and hypostomal subgenal sulcus
is the POSTGENA. The SUBGENA is posteroventrally separated from it by the
hypostornal subgenal sulci. The posterior part of the postgena represents the
OCCIPUT, even in the absence of a demarcating occipital sulcus according to
Snodgrass ( 1960). Therefore, in the ‘macro’ lepidoptera, where the occipital
sulcus is questionable at best, the postgena and occiput intergrade with one
another along the anteroposterior axis of the head. This region represents the
POSTGENAL-OCCIPITAL AREA. The temporal sulci merely cross this area, but d o
not delimit specific areas as some authors maintain (Issiki, 1931; Madden, 1944;
Michener, 1952; Hannemann, 1956; Srivastava, 1957; Ehrlich, 1958a).
The VERTEX represents the postgenal-occipital area on the dorsal head surface,
dorsad of the transfrontal sulcus, and lacks specific boundaries. Eassa ( 1963a)
considers the vertex (‘epicraniurn’) in adult Pieris to arise anew during
metamorphosis, growing posteriorly from between the antennae. The dorsal and
posterolateral surfaces of the head may thus have separate derivations in adult
and larval Lepidoptera.
In lycaenids, the head is posterodorsally produced as a broad swelling between
the temporal sulci. This specialized area is medially divided by the dorsal
temporal sulci, and can be referred to as the POSTGENAL-OCCIPITAL
PROTUBERANCE.
The posterior region of the head has two dorsoventrally oriented arcs of semipersistent setae that flank the foramen magnum laterally. These setal areas might
be homologous with Jordan’s ( 1923) chaetosemata, or the sensory area noted by
Ehrlich (1958a).
Each A N T E N N A has a globe-like SCAPE, a smaller ring-like PEDICEL and an
elongate, clubed FLAGELLUM. The flagellar club segments show transverse rows
of minute setae flanking their anteroventral margins and are less sclerotized
dorsally. Sulci o r carinae are absent on the surface of the club.
The head is internally reinforced by a n-shaped TENTORIUM, which is
invaginated at the tentorial pits. The anteriorly extended TENTORIAL ARMS
expand before finally constricting near the anterior tentorial pits.
A tri-lobed HYPOPHARYNGEAL APODEME projects internally from the
anteroventral region of the head capsule, providing muscle anchorage for the
5
J. T. SORENSEN
66
hypopharyngeal pump dilators. Its two posterolateral lobes are machete-shaped,
while the third spade-like lobe projects posterodorsally.
Cervix (Figs 9-1 1)
A CERVICAL MEMBRANE links the postocciput and the anterior prothoracic
margin. A pair of T-shaped LATERAL CERVICAL SCLERITES (lateral cervicalia :
Weber, 1933) occur ventrolaterally. The posterior zone of the cervical sclerite’s
dorsal lobe articulates, in a rocking fashion, against an elongate fossa on the
anterodorsal margin of the propleuron, and is attached to it by a reinforced
membranous connective. The anterior tip of the cervical sclerite articulates
internally with the cervical articulation fossa of the tentorial bridge. A VENTRAL
CERVICAL SCLERITE (Srivastava, 1961) or sclerotized bridge (intercervicalia:
Weber, 1933)connecting the lateral sclerites is absent.
A CERVICAL ORGAN (Ehrlich, 1958a)is present on the posteroventral margin of
the anterior lobe of the cervical sclerite, and is pitted, bearing numerous minute
setae. The structure may be sensory, and is not as clearly demarcated in
Glaucopsyche as is shown for Danaus by Ehrlich (1958a: fig. 1 1).
Prothoracic notum (Figs 9, 1 1, 12)
The PRONOTUM consists of three plates: a DORSAL PLATE and a pair of medially
fused LATERAL PLATES. Nuesch (1953) erroneously applies the pteronotal terms
PRESCUTAL SULCUS I I
ANTERIOR NO
I
___
I
POSTERIOR NOTAL WING PROCESS I I
AXILLARY CORD I I %
-.
ANTERIOR NOTAL
W I N G PROCESS Ill-{\
I
-
V
MEDIAN SCUTAL
SULCUS
1
SCUTOSCUTELLAR
SULCUS I 1
K<
SCUTELLUM I 1
AXILLARY CORD 111’
SCUTELLUM I I I
SCUTOSCUTELLAR
SULCUS I l l
Figure 9. Thoracic notum, dorsal view; pro-, rneso- and rnetathoracic parts in common designated
1, 11 and 111, respectively; left wings and axillary region removed; right wings and axillar). region
distorted distally to display articulation with notum; right tegula removed.
IhTEGULMENTALANATOMY O F GIAMCOPSYLHF
PRESTERNUM
7
,
LATERAL.
\
67
CERVICAL SCLERITE
EPISTERNUM I
COXA GENUINA I
EFIMERON I
PLATE
‘‘1
/
PLEURAL SULCUS I I
SECONDARY STERNOPLEURAL
SULCUS
1
ANTERIOR COXA GENUINA I 1
DORSAL EPIMERAL SULCUS I I
EPIMERON 1’
I’
/,
y y
4
MAPGINOPLEURAL SULCUS I ,I I
PLEUPAI SULCUS I I I
-7
,
b
\
E P I M E R A L SULCUS I 1
POSTERIOR COXA GENUINA I 1
BAS I COSTAL SULCUS I I
MEDIAN LONGITUDINAL
SULCUS 111
10
pigui-c 10. Thoi-acic sterrlopleural region, ventral view; pro-, meso- a n d r l I ( . I d I h ~ ~ i d ( i cf ~ d1\i iii
( o i i ~ i ~ i oticwgirdted
ii
I , I1 dnd 111, respertively; abdominal swriia designarvd i i i r ~ i d l ~ i i( i ~ i i ~ ( ~ i ~I iiglii
I\.
W X ~ K i~c i r i w r d : left ri-(~thanters
removed; abdominal srei-na alrcr Ehilich w ( ’ I iy 1 I
scutum and prescuturn to these structures. Snodgrass (1935) states that the
pronoturn is derived from the primary segmental region and its divisions are not
equivalent to those of the pteronota.
Weber (1924ai believes the lateral plates (‘Spange’)are pleural in origin. noting
that they are continuous with the pleura in Cossus and some othcr Lcpidoptera,
but are separated from them by a sulcate line in Pupilio. Most other authors
(Berlese, 1909; Schultz, 1914; Madden, 1944; Michener, 1952; Niiesch, 1953;
Ehrlich, 1958a; Srivastava, 1961) consider the lateral plates to be of tergal
derivation.
The dorsal plate (postzona: Srivastava, 196 1 ) is subtriangular with secondary
lobes on its anterolateral margins. The anterior margin is fused with the
posterodorsal margin of the unified lateral plates (prozona: Srivastava, 196 11,
forming a sulcate invagination (interzonal suture: Srivastava, 1961). Srivastava
(196 1 ) calls the inedian fusion line of the lateral plates the longitudiiial prozonal
sulcus; there are no sulci homologous with his transverse prezonal sulcus o r
longitudinal postzonal sulcus. The median fusion line of the lateral plates
produces a small internal ridge which give rise to a knob-like PRONOTAL
PROJECTION (prozonal keel: Srivastava, 1961) anteroventrally. The lateral plates
are sern-fused ventrally with the pleura, articulating against thcnn tiv a
J. T. SORENSEN
68
translucent, resilient strip of cuticle. A pair of semi-sclerotized PATAGIA flank the
anterior margins of the lateral plate. PARAPATAGIA (Schultz, 1914)are absent.
Prothoracic sternopleural region (Figs 10-1 2)
There is disagreement in the literature concerning this region. Snodgrass
adult Pterygota, except the Plecoptera, show the propleuron
divided into an episternum and an epimeron by a pleural sulcus and ridge.
Madden (19441, Nuesch (1953)and Srivastava (1961)concur, but Michener (1952)
and Ehrlich ( 1958a)oversimplify the propleuron, showing no divisions.
Most of the PROPLEURON is the EPISTERNUM. The PLEURAL SULCUS extends
between the pronoto-pleural and coxo-pleural articulations and represents the
external posterior margin of the propleuron. The episternum is divided into an
( 1935: 173) states all
.
FURCASTERNUM/
,
\
DORSAL PRONOTAL PLATE
LATERAL PRONOTAL PLATE
PRONOTAL PROJECTION
COXO-PLEURAL ART I CULATI ON
12
Figures 1 1 , 12. Fig. 1 1 . Prothorax, left lateral view (anterior to left); lateral cervical sclerite slightly
enlarged to show detail, legs removed, internal components outlined by dashed lines. Fig. 12.
Excised prothorax, posterior view; legs and spinasternum removed.
I N T E G U M E N T A L A N A T O M Y O F GLAUCOPSYCHE
69
anepisternurn and an infraepisternum by an ANAPLEURAL CLEFT, but these
divisions are obscured because the clift is poorly defined. The A N E P I S T E R N U M is
represented as a marginal lobe on the anterodorsal margin of' the episternum.
The INFRAEPISTERNUM is represented by the much larger, lower portion of the
episternum. The EPIMERON is comprised of two, mostly membranous parts. The
dorsal part is present as a small sclerotized flap below the pronoto-pleural
articulation, but may be absent in some individuals. The ventral part borders the
coxal cavity posterior to the coxo-pleural articulation. The PLEURAL R I D G E is
greatly enlarged and extends posteromedially to fuse with the furca (see below).
The COXO-PLEURAL ARTICULATION is monocondylic, but bifid on the
propleuron, and the coxa rocks between these projections. This articulation
probably does not represent a fused trochantin, since the pleural sulcus is
associated with the anterior projection of the articulation on the propleuron; it
should be associated with the posterior projection on the propleuron if a fused
trochantin is involved.
The BASISTERNUM is invaginated medially forming a MEDIAN L O N G I T U D I N A L
SULCUS, and a strong internal STERNAL RIDGE which diminishes posteriorly
before fusing with the furcasternum.
A narrow, transverse and entire sclerite is present along the anteroventral
margin of the prothorax. The homology of this structure is unclear. Madden
(1944) and Ehrlich (1958a) consider it to be the PRESTERNUM but it may be a
secondary sclerotization.
The FURCASTERNUM is present externally as a transverse sclerite forming the
posterior margin of the coxal cavity, but does not form a ventral coxal articulation, as on the pterothoracic segments. Internally, the furcasternum produces a
bipronged FURCA. Weber (1924b) believes the pleural ridge serves as the furcopleural connective, a view shared here. However, Dierl (1964) and Matsuda
( 1970) believe this furco-pleural connective represents the ventral end of the
epimeron, while Ehrlich (1958a) labels this the furcal lamella.
A two-part SPINASTERNUM articulates against the posterior margin of the
furcasternum, and invaginates centrally, to form the SPINA articulation.
Mesothoracic notum (Figs 9, 13, 15, 1 7 )
The enlarged and convex mesonotum is derived by secondary segmentation
and occupies most of the pterothoracic tergal region. Its acrotergite, antecosta,
prescutum, scutum and scutellum are of mesothoracic derivation, while its
postnotum is metathoracic in origin (Snodgrass, 1935).
The ACROTERGITE traverses the anterior margin of the mesonotum and occurs
as a fine anterior marginal lip along the ANTECOSTAL SULCUS. The extreme
reduction of the acrotergite among papilionoids may have contributed to
Ehrlich's (1958a, b) failure to detect it, although Srivastava (1962)and Crombach
(1967) recognize it. I t is slightly indented anteromedially to articulate with the
posterior tip of the pronotal dorsal plate. The antecostal sulcus marks the
internal ANTECOSTA. The latter is reduced medially, but produces lateral lobes. I t
represents PHRAGMA 1, the first phragma of Snodgrass (1935).
The PRESCUTUM occurs between the antecostal sulcus and the PRESCUTAL
SULCUS (prescutoscutal sulcus: Matsuda, 1970). Its anterolateral margins extend
posteroventrally as the PREALAR ARMS. A SUBTEGULA, o r prescutal apodeme
70
J . T. SORENSEN
(Sharplin, 1963a) represents a ventral, detached portion of the prescutum,
according to Freeman (1947). Matsuda (1970: 15) agrees with this interpretation
and states the subtegula is homologous with the prealar sclerite of the other
orders.
The SCUTUM occurs between the prescutum and the SCUTOSCuTELLAR S u L C u s .
Its lateral margin has three extensions that form wing articulation points. The
anterior of these, the SURALARE, is separated from the scutum by a weak
anterolateral SCUTAL SULCUS (Matsuda, 1970) that produces a strong internal
ridge. There is no MARGINAL SULCUS (Srivastava, 19621, outlining the edge of the
suralare. The suralare forms the ANTERIOR NOTAL W I N G PROCESS. A second
scutal extension represents the MEDIAN NOTAL WING PROCESS. A small, thin
incision occurs on the margin of the scutum, immediately anterior to the base of
this process, allowing greater flexibility (Sharplin, 1963a). A third scutal
extension, the POSTERIOR NOTAL WING PROCESS, is closely associated with the
median notal wing process. A POSTEROLATERAL SCUTAL SULCUS (Matsuda, 1970)
is absent between these processes.
These processes have a variety of synonyms in the literature. The suralare is
referred to as the procondilo (Berlese, 1909). The median notal wing process is
referred to as the mesocondilo (Berlese, 19091, the adnotal (Madden, 1944;
Michener, 1952; Ehrlich, 1958a) and the lob0 scrutal (Onesto, 1959). The
posterior notal wing process is referred to as the postadanale (Madden, 1944),
the postalare (Michener, 1952)and the postalare plate (Ehrlich, 1958a).
There is also disagreement as to the delimitation of the various notal wing
processes recognized as functional units. Srivastava (1962) terms the suralare the
anterior notal wing process, and the remaining scutal extensions collectively as
the posterior notal wing process. Ehrlich (1958a) denotes the suralare plus the
medial scutal extension as the anterior notal wing process, since both articulate
with the first axillary sclerite. I follow Matsuda (1970) in terming the suralare
alone the anterior notal wing process, since the median scutal extension is well
separated from it. The latter, by articulating with the posterior part of the first
axillary sclerite, does not fit Snodgrass’s (1935: 190) definition for the anterior
notal wing process. He gives no term to the articulation point of the scutum with
the posterior part of the first axillary sclerite.
The scutum bears a MEDIAN SCUTALSULCUS (Matsuda, 1970)that arises medially
from the prescutal sulcus, but lacks an internal ridge. The scutoscutellar sulcus and
its internal ridge are obliterated medially. The posterolateral margins of the
SCUTELLUM bend ventrally to connect to the postnotum via intersegmental
membrane, while the anterolateral corners are continuous with the mesonotal
axillary cords.
The mesothoracic postnotum represents the detached metathoracic
acrotergite, antecostal sulcus and antecosta. The latter represents Snodgrass’s
(1935) second phragma. The antecosta forms a pair of LATEROPHRAGMATA 2
laterally (Sharplin, 196sb), anchoring the lateral oblique dorsal longitudinal
flight muscles. The antecosta and the semi-detached metathoracic ‘Prescutum’
(see the metathoracic notum section) medially form a greatly enlarged MEDIAN
PHRAGMA 2 for anchorage of the dorsal longitudinal flight muscles.
The postnotum has been vertically displaced below the posterior edge of the
scutellum, and bears a pair of external lateral arms, the POSTALAR BRIDGE
(Matsuda, 1970). These serve as articulated levers for the pleural region during
I N T E G U M E N T A L ANATOMY O F CLAlJCOP.SY(.HI.
l3
SCUTAL SULCUS
71
/ MEDIAN NOT4L WING
\
PROCESS
POSTERIOR NOTAL WING PROCESS
$
\
/
SCUTOSCUTEILAR SULCUC
TERGOPLEURAL AP
//\La-
TRAL EPIMERAL SULCUS
BASICOSTAL SULCUS
INFRAEP ISTERNALBASISTERNAL
INFRAEPISTERNALPLATE
BASISTERNAL
PLATE
/
SECONDARY STERNO2LEURAL SULCUS
FARACOXAI-MARGINOPIEURAL
PLEURAL WINLPROCESS
ANTERIOR COXA GENUI NA
SULCUS’
S P I NASTERNUM I
PREPECTUS
BASISTERNAL PLATE
SECONDARY STERNOPLEURAL SULCUS
MEDIAN LONGITUDINAL
SULCUS
PARACOXAL-MARGINOPLEURAL SULCUS
14
Figures 13. 11. Fig. 13. Mesothorax, left lateral view (anterior to left); wing\ dnd axiilai\ i-cgioii
I-emoved, ti-ochanter removed, tegula removed but position outlined, tergopleural npotiemc 15
internal but outlined. Fig. 14. Schematic map of mesothoracic sternopleu~-al
1-egiori(antel-ioi 1 0 r q ~ .
flight. The postalar bridge articulates with the posterior margin of the posterior
notal wing process of the scutum, posterior to the wing. I t fits against the
posterodorsal margin of the epimeron ventrally, being separated from the latter
by the epimeropostalar sulcus. The postalar bridge articulates dorsally with the
posterior scutellar margin, near the origin of the axillary cords, and against the
lateral margin of the dorsomedial portion of the postnotum. The
laterophragrnata occur at the invaginated posterior margins of the postalar
bridges. Sharplin ( 196313) indicates that the contribution of the metathoracic
prescutum to them is solely membranous in the higher Ditrysia, and is
represented only by a rudimentary detached prescutal apodeme connected to the
72
J. T. SORENSEN
posterior margin of the laterophragma by a short ligament immediately beneath
the nearby spiracles. Ehrlich (1958a)implies the postalar bridge to be epimeral in
origin (“postalare portion of the epimeron”), but an association with the
laterophragmata indicates a postnotal derivation.
The median phragma 2 reaches deep into the thorax and is ventrally bilobed
to permit passage of the gut and the dorsal blood vessel. The anterior margins of
this phragma produce a pair of internal lateral arms that extend anteriorly to
meet and flex against the inner surface of the median notal wing process of the
scutum. This allows the dorsal longitudinal muscles direct leverage on the first
axillary sclerite during wing movement (Sharplin, 1963b).The degree of anterior
extension of these arms appears to reflect phylogenetic advancement within the
Lepidoptera according to Sharplin ( 1963b). The internal contact point of these
arms with the median notal wing process is described as an articulating socket in
Danaus (Ehrlich, 1958a), but in Glaucopsyche it resembles a blade fused against the
inner scutal surface along a less sclerotized flexion line.
As Sharplin ( 1963b) notes for the Ditrysia, there is no complete membranous
invagination between the laterophragmata and the median phragma.
Mesothoracic sternopleural regton (Figs 10, 13, 14, 16- 18 )
The PLEURAL SULCUS traverses the pleural region vertically, demarcating the
episternum anteriorly, and the epimeron posteriorly. Internally, the strong
PLEURAL RIDGE is fused with the coxal ridge ventrally, and forms the PLEURAL
WING process dorsally.
The episternum is differentiated into several subdivisions. The ANEPISTERNUM
(sensu stricto) is extremely reduced and separated from the lower episternum by a
membranous ANAPLEURAL CLEFT (Matsuda, 1960). The anepisternum is hidden
at the base of the overlying basalare, but occurs internally as a small lobe
projecting distally from the anterior margin of the pleural ridge at the level of the
pleural wing process. The posterodorsal margin of the anepisternum extends to
form the dorsal half of the PLEURAL WING PROCESS, the epimeron forming the
ventral half and the pleural sulcus separating the two. In Papilio, Crombach
( 1967) implies that the anepisternum and epimeron fuse to form the pleural wing
process in the absence of the pleural sulcus, which-he illustrates as ending
ventrad to the process at the anterior margin of the anepisternum. This interpretation is erroneous since it violates Snodgrass’s (1935: 190)definition of the
posterior demarcation of the episternum by the pleural sulcus. Srivastava (1962)
correctly portrays the relationship of these parts in forming the pleural wing
process in Papilio.
The anterodorsal margin of the anepisternum extends to form an internal
TERGOPLEURAL APODEME (Sharplin, 1963a), which is distally fused to and
supports the SUBTEGULA (Sharplin, 1963a). This entire structure has been called
the tegular arm (Weber, 1924a, 1928; Ehrlich, 1958a; Crombach, 1967)and the
subtegular arm (Weber, 1933; Nuesch, 1953; Srivastava, 1962).
The tergopleural apodeme appears unique to the Lepidoptera and serves as
the third tergopleural muscle (t-p 4 : Matsuda, 1970) anchorage (Sharplin,
1963a). In Glaucopsych, as is typical of the Ditrysia (Sharplin, 1963a), the unified
subtegula and tergopleural apodeme are fused to the prealar arm.
The basalare is bipartite (Fig. 18). Sharplin ( 1963a)believes both of its sections
IKTEGLMENTAL ANATOMY OF GLAlJCOPSYCHE
15
/ SCUTAL RIDGE
PRESCUTUM
PRESCUTAL RIDGE
SUBTEGULA
TERGOPLEURAL APODEME
SURALARE
MEClAN NOTAL WING PROCESS
POSTALAR BRIDGE
EPIMEROPOSTALAR RIDGE
LATEROPHRAGMA 2
1
,PLEL
IRA1 WING PROCESS
TERGOPLEURAL AP
I NFRAEP ISTERNALBASISTERNAL PLATE
"U
DORSAL EPIMERAL
PREEPIMERA, SUL
l6
COXAL R I D G E 7
BAS I COXITE
BASICOSTAL SULCUS
ANTERIOR COXA GENUINA'
Figurea 15, l h Fig. 15. Mesothoracic notum, right lateral view of parasagittal w c t i o i i ( a n r c i i o i t o
Irk); tergoplrural apoderne severed. Fig. 16. Mesothoracic sternopleural region, I ighr l a t c ~ a l iev o t
parasagittal h e c t i o n (anterior to left); trochanter removed, prealar arm srverccl, p o s ~ t l a i hi idge
acvc1-ed.
are typically separate from the episternum. In Glaucopsyche, however, the
posterior margin of the SECOND BASALARE (posteriad) is fused with the anterior
(distal) margin of the anepisternum. The FIRST B A S A L ~ R E(anteriad) articulates
against the second, to which it is attached by a short ligament. A BASALAR
A P O D E M E arises internally from each basalar section.
The lower portion of the lepidopteran mesothoracic episternum, below the
anapleural cleft, is represented by such chaos in the literature that synonymy of
terms of different authors is difficult. Several systems for the region are available
(Weber, 1928; Shepard, 1930; Ferris, 1940; Michener, 1952; Matsuda, 1970;
Brock, 19711, but interpretations differ markedly. Although most work on the
J. T. SORENSEN
74
SCUTELLUM
MEDIAN PHRAGMA 2
AXILLARY CORD
~
POSTALAR BRIDGE
\\\
LATEROPHRAGMA 2
SECONDARY FURCAL ARM
EPIMERON
FURCA
POSTERIOR COXA GENUINA
MERON
COXAL SULCUS
ANTERIOR COXA GENUlNA
Figulc I 7 Mesothorax, posterior view, wlngs and trochanters removed, axilldry c o l d s
bC\el
ed.
18
\
\r \
TERGOPLEURAL APODEME
\\
BASALARE 2
,
/-
/PLEURAL WING PROCESS
\\
)
I
ANEPISTERNUM
PLEURAL SULCUS
BASALARE 1ANAPLEURAL CLEFT
I'
BASALAR PROCESS
-I
\ INFRAEPISTERNAL-BASISTERNAL
P,LEURAL
BASALAR PAD
I'LATE
WING PROCESS
ANEPISTERNUM
BASALARE 1
PLEURAL SULCUS
EPIMERON
ANAPLEURAL SULCUS
19
I
Figures 18, 19. Fig. 18. Mesothoracic basalare region, left lateral view (anterior to left); tergopleural
apodeme is outlined with dashed lines. Fig. 19. Metathoracic basalare region, left lateral view
(anterior to left).
INTEGU-MENTAL ANATOMY OF GLAUCOP.YYCHI
75
Papilionoidea (Ehrlich, 1958a, b ; Srivastava, 1962; Crombach, 1967) has been
based upon Shepard’s ( 1930) system, Brock (197 1), employing Matsuda’s ( 1960)
concepts, argues that Shepard incorrectly interpreted ’the precoxal sulcus,
resulting in improper episternal homologies. Thus, both Brock ( 197 1 ) and
Matsuda ( 19 70) have independently recognized serious errors in the interpretations of their predecessors, but unfortunately neither was aware of the
other’s forthcoming work (Matsuda, personal communication) s o their
terminologies do not agree. Brock, limiting his study within the order, examined
a large number of genera and proposed a rather complicated evolutionary
pattern for sternopleural development. After examining several genera
throughout the order, I have decided to employ Brock’s concepts, but use
Matsuda’s broader terminology wherever feasible, to eliminate Brock’s more
specialized terms.
The episternum below the anapleural cleft is the infraepisternum (Brock,
197 1). While the term katepisternum is used synonymously by several workers
(Shepard, 1930; Snodgrass, 1935; and others), its use should be’restricred 10 that
portion of the episternum of katapleural origin that is marked off dorsallv and
anteriorly from the anapleural region by the paracoxal sulcus (Matsuda, 19 70).
The paracoxal sulcus is in part synonymous with Snodgrass’s (1935) precoxal
sulcus (see Matsuda, 1970:35), and it is used here in preference since Matsuda
maintains that, in a primitive condition, it completely circumscribes the coxa.
The infraepisternum appears to extend ventrally to the median longitudinal
sulcus, but this plate represents a fusion product of the preepisternal and
katepisternal subdivisions of the infraepisternum with the basisternum (Brock,
197 1), through loss of the pleurosternal sulci, to create an INFRAEPISTERNALBASISTERNAL PLATE. N o trochantin is evident along this plate’s posterior margin.
An area of secondary sclerotization, the PREPECTUS o r hypopteron
(parepisternurn : Brock, 197 1), flanks the infraepisternal-basisternal plate’s
anterior margin, but a secondary demarcating sulcus (parepisternal sulcus :
Brock, 197 1 ) and accompanying internal ridge are absent. Previous workers
(Madden, 1944; Michener, 1952; Ehrlich, 1958a; Srivastava, 1962) have
erroneously considered the prepectus to represent the preepisternum.
A PARACOXAL-MARGINOPLEURALSULCUS flanks the infraepisternal-basisternal
plate’s posteroventral margin, and is continuous with the pleural sulcus latc.rally.
Brock ( 197 1) believes that the lower section of the bipartite paracoxal sulcus (his
‘precoxal’ sulcus) merges dorsally with the marginopleural sulcus in the
Papilionoidea, t o form a fused, continuous structure. Internally, the posterior
section of this composite sulcus, anteriad to the pleural ridge, has a
MARGINOPLEURAL RIDGE which expands laterally, merging with the pleural
ridge. According to Brock (197 I ) the intermediate step to this formation is
evident in some advanced hesperiids, where the lower section of the paracoxal
sulcus (sternopleural sulcus : Ehrlich, 1960) is unfused with the marginopleural
sulcus (see Brock, 1971; fig. 33a, b ; Ehrlich, 1960: figs 10, 181.
In Glaucopsyche a SECONDARY STERNOPLEURAL S U L ~ U S(Brock, 197 1) arises near
the pleural sulcus and ends abruptly in the infraepisternal-basisternalplate. Brock
states that this sulcus is restricted to the Nymphalidae and Lycaenidae, and is
complete in the former to the infraepisternal-basisternalplate’s anterior margin,
near the spinasternum. This is equivalent to Ehrlich’s ( 1958a) precoxal sulcus in
Danaus. Brock believes this sulcus arose as a posterior extension of a composite
J . T. SORENSEN
76
20
\&A,
SCUTUM \
SCUTAL SULCUS\
/ SUBALARE
SCUTOSCUTELLAR SULCUS
/ SCUTELLUM
METATHORACIC SPIRACLE
PLEURAL WING PROCESS
INFRAEPI STERNALBASISTERNAL PLAT
PLEURAL SULCUS
MARGINOPLEURAL SU
VENTRAL EPIMERAL SULCUS
RY COXAL SULCUS
BASICOSTAL SULCUS
SUBALAR APODEME
I NFRAEP I STERNAL
BASISTERNAL PLATE
MARGINOPLEURAL RIDGE
BASICOSTAL SULC
VENTRAL EPIMERAL RIDGE.
SECONDARY COXAL RIDGE
SECONDARY COXAL SULCUS
BASICOSTAL SULCUS
21
Figures 20, 21. Fig. 20. Metathorax, left lateral view (anterior to left); wings and axilla? region
removed, trochanter removed. Fig. 21. Metathoracic sternopleural region, right lateral view of
parasagittal section (anterior to left); trochanter removed.
sulcus, formed from a combination of the anterior section of his basisternal sulcus
with the ventral end of the sulcus separating the prepectus and the preepisternum
(Brock’s ‘parepisternal’ sulcus), extending to the paracoxal sulcus. It should be
stressed that the homology of this sulcus and the paracoxal-marginopleural
sulcus, as discussed by Brock (197 11, represent a dramatic evolutionary change
within the Hesperioidea and Papilionoidea, and is therefore tentative.
The MEDIAN LONGITUDINAL SULCUS (discrimen: Michener, 1952) represents
the medial section of the basisternum which has invaginated to form the median
longitudinal, or sternal, ridge. This sulcus splits anteriorly before reaching the
mesothoracic margin, near the spinasternurn. While Srivastava ( 1962) considers
INTEGUMENTAL ANATOMY O F G L A U C O P S Y C N E
77
these anterior branches to be sternopleural sulci in Papilio, the term is avoided
here due to disagreement with Brock’s concepts. These sulci may represent the
anterior margin of the basisternum, with the anteriad sclerotized triangle a
secondary development.
The STERNAL RIDGE rises as an arch, but diminishes posteriorly before
reaching the furca; a condition that Ehrlich (1958b) considers typical among
Lycaenidae. The FIJRCA itself represents the invaginated furcasternum, the base
of which joins the sternal ridge posteriorly, to form the VENTRAL COXAL
ARTICULATION. The main stem of the furca produces a vertical, circular lobe
from the posterior margin, and bears two pairs of arms dorsally. A wide,
flattened pair of SECONDARY FURCAL ARMS (lateral furcal arms) project
dorsolaterallv to fuse with the invaginated posterior margin of the epimeron.
The dorsal Inargin of these secondary arms bears a pair of rod-like PRIMARY
FURCAL ARMS (dorsal furcal arms), which project anterodorsolaterallv to fuse
with the pleural ridge, dorsad of the preepimeron.
The dorsal margin of the epimeron is invaded by a broad membranous
intrusion which reaches the plate’s centre. The posterodorsal margin is joined to
the postalar bridge of the postnotum; an EPIMEROPOSTALAR SULCUS (Srivastava,
1962) delimits their juncture. The upper section of the anterior margin of the
epimeron overrides the pleural sulcus, and provides anchorage for the muscles to
the third axillary sclerite. A small PREEPIMERON occurs along the pleural sulcus,
just ventral to the overriding epimeron, but is difficult to distinguish. The
demarcating PREEPIMERAL SULCUS is extremely faint, and often completely
obliterated.
Two sulci transect the lower half of the epimeron horizontally. Neither is
mentioned nor depicted by Madden ( 1944), Srivastava i1962) and Crombach
(1967). Michener (1952: fig. 8) and Ehrlich (1958a: fig. 9) depict the more dorsal
of these as a broad, curved inflection which traverses the epimeron, but neither
mention this structure in the text. In Glaucopysyche the more dorsal of these sulci,
the DORSAL EPIMERAL SULCUS, is continuous anteriorly with the pleural sulcus
beneath the overriding epimeron, and extends posteriorly to near the centre of
the epimeron. In some individuals the dorsal epimeral sulcus extends, extremely
faintly, posterodorsally to the posterior margin of the epimeron, and produces a
faint ridge internally. This posterior part of the sulcus is slightly more evident in
Nymphalis (present data).
The more ventral of these suki, the VENTRAL EPIMERAL SULCUS, is marginal
s
along the anterior half of the ventral border of the epimeron. I t c u n ~ dorsad
centrally along the ventral border and becomes weakened or obliteratcd, but
becomes well defined again just dorsad of its disappearance, running posteriorly
to terminate internally on the ventral margin of the secondary furcal arm. The
ventral epimeral sulcus produces an internal ridge anteriorly and posteriorly.
This posterior section of the sulcus may be mistaken for that of the dorsal
epimeral sulcus, since it appears in line with the anterior section of the latter.
The homologies of these sulci are uncertain, but both are probably secondary
reinforcements for the epimeron. The dorsal epimeral . sulcus may represent
Matsuda’s ( 1970) transepimeral sulcus, since it runs posterodorsally (see
Matsuda, 1970: 36). Probably neither represents the paracoxal sulcus on the
epimeron, since Matsuda ( 1970:35) considers that to be an archaic characteristic.
The SUBALARE is membranously encircled, dorsal to the epimeron. I t is
J. T. SORENSEN
78
attached to the posterior notal wing process by a short elastic ligament. The
dorsal margin of-the subalare forms a shelf'which extends internally as a winglike SUBALAR APODEME. The latter is externally demarcated posteriorly by a deep
asymmetrical cleft. The ventral margin of the subalare is simple, with a less
sclerotized cleft that allows flexion during pleural distortion in flight. The
MESOTHORACIC SPIRACLE is anterior to the prealare.
Metathoracic notum (Figs 9, 20, 22 and 23)
As noted, the metathoracic acrotergite and antecostal sulcus are completely
fused to the mesothorax by secondary membranization, to form that segment's
postnotum. Sharplin (1963b: 1123) erroneously and confusingly refers to the
metathoracic acrotergite as the mesothoracic acrotergite. The mesothoracic
acrotergite occurs at the anterior margin of the mesonotum.
Most workers on the Papilionoidea (Ehrlich, 1958a; Srivastava, 1962;
Crombach, 1967) consider the prescutum absent. Sharplin ( 1963b1, however,
shows that it is externally present in the Zeugloptera and has gradually
invaginated (usually becoming membranous) during evolutionary advancement
in the Lepidoptera.
/ SCUTELLUM
SCUTOSCUTELLAR
SULC2.s
22
\ /---
SECONDARY SCUTELLAR
SCLERITE
\
I I
LATEROPHRAGMA 3
23
Figures 22, 23. Metathoracic noturn. Fig. 22. Anterior view. Fig. 23. Posterior view; wings removed,
axillary cords severed.
I S T E G U % l E N T A L ANATOMY O F GLAIJCOP.CYCElF
79
I n Glaucopsyche the condition of the PRESCUTUM is unclear. I t may be
completely invaginated to form the sclerotized, laminated posterior zone of the
median phragrna 2, thus serving as the posterior part of the metathoracic
antecosta. If this is the case, the interface between the ‘prescutum’ (the apparent
posterior region of median phragma 2, or the posterior region of the
metathoracic antecosta) and the median phragma 2 itself (the apparent anterior
region of median phragma 2, o r the anterior region of the metathoracic
antecosta), represents the deeply intruding antecostal sulcus. The ‘prescutum’
remains sclerotized (perhaps secondarily) but has become semi-detached from
the scutum by membranization of the PRESCUTAL SULCUS; only at the lateral
extremes, near the anterior notal wing process, is a scutal connective evident (Fig.
2 2 ) . The ‘prescuturn’ is discernible only by carefully teasing it apart from the
median phragrna 2, beginning at the lateral scutal connective, and separating it
along the antecostal sulcus. If this scutal connective is broken, the scutum
separates, leaving the ‘prescutum’ indistinguishably fused to median phragma 2 .
This difficult,. may explain the previous failure to detect the prescutum in
papilionoids.‘For example, Ehrlich’s phragmal processes (1958a: fig. 10) would
be ‘prescutal’, and the ‘prescutum’ would be represented by the phragma
illustrated there. The problem of interpreting the prescutum may be semantic in
large part, since there is no definite boundary between the prescutum and the
posterior part of the antecosta.
The PRESCLTAL A P O D E M E is rudimentary and dissociated from the
mesothoracic laterophragma 2, but is connected to it by a short ligament. The
prescutal arm is fused to the second basalare (Fig. 19) according to Sharplin
(1963b),who states this occurs throughout the Ditrysia and refers to the resulting
composite structure as the basalar process. This concept is adopted here since the
structure is laminar in Glaucopsyche. A SUBTEGULA is absent.
The SCUTLM is U-shaped to accommodate the enlarged median phragma 2, and
the anteromedial margins each have a less sclerotized notch allowing flexion.
Laterally, the scutum has well developed ANTEROLATERAL and POSTEROLXTERAL
SCUTAL SULCI, both with strong internal ridges. The suralare, bearing the
ANTERIOR KOTAL W I N G PROCESS is reduced. The M E D I A N NOTAL W I N G PROCESS is
not developed or differentiated from the lateral margin of the scutum.
The POSTF.RIOR NOTAL W I N G PROCESS is well developed posterior to the
intersection of the posterolateral scutal sulcus with the lateral margin of the
scuturn.
The SCUTOSCUTELLAR SULCUS occurs as an inverted ‘V’ o n the posterior face of
the rnetanotuin and dorsomedially curves anteriad between the lobes of the
scutum. The SCUTELLUM is largely restricted to the posterior face of the
inetanotuin, but produces the axillary cords laterally. Brock (197 1) believes that
the lycaenid scutellum has developed SECONDARY SCUTELLAR S U L C I which
distinguish a sE:CONDARY SCUTELLAR sCLERITE, ventrolaterally. This is true in
Glaucopsyche.
A wide SUPRAPHRAGMAL SCLERITE (Brock, 197 1) occurs between the ventral
(posterior) rnargin of the scutellum and the anterior margin of the postnotum.
Brock (197 1) believes this to be a secondary sclerotization, among advanced
Ditrysia, of this normally membranous region in primitive forms. He considers
the supraphragrnal sclerite to represent fusion of a pair of mesially advancing
sclerotizations arising from the laterophragmata of lower Ditrysia.
80
J. T. SORENSEN
The metanotal postnotum represents the acrotergite and antecosta of the first
abdominal tergum which have been sequestered anteriorly to the metathorax. In
the Ditrysia this region has undergone a unique and complex evolution. In some
papilionoids the development of numerous structural secondary sulci make interpretations of regional boundaries difficult and confusing. Brock (1971) has
outlined the general development of this area throughout the Lepidoptera ; his
concepts are followed here.
The acrotergite of the postnotum is represented as a strip that is fused to the
metathorax laterally but remains separate from it dorsomedially. The antecostal
sulcus invaginates medially, forming an extremely reduced antecosta along the
posteromedial margin of the acrotergite. This reduced internal ridge represents,
in a strict sense, the medial portion of the third phragma, PHRAGMA 3, equivalent
to the enlarged, median phragma 2 of the mesothorax. This ridge serves as the
anterior anchorage for the muscles originating on the antecosta of the second
abdominal tergum (muscle set 116: Ehrlich & Davidson, 1961 and Ehrlich &
Ehrlich, 1963; muscle set 126: Srivastava, 1966). Brock (1971) refers to this
reduced posteromedial portion of phragma 3 as the tergal phragma.
The antecostal sulcus laterally invaginates forming the LATEROPHRAGMATA 3.
These may produce anteroventral lobes or stalks in some papilionoids (Ehrlich,
1958a, b). Gluucopsyche shows the latter condition.
The laterophragmata 3 are offset anteriorly from the medial, reduced
antecosta, the antecostal sulcus curving somewhat anteriad, laterally. The interlaterophragmal gap resulting from their transverse misalignment with the medial
section of the antecosta is bridged by a TRANSPHRAGMA 3 according to Brook
( 197 1), who considers such transphragmal development as characteristic of the
higher Ditrysia, and to have evolved as proximal extensions of the
laterophragmata that have fused medially.
In Gluucopsyche the extremely reduced median portion of the antecosta is
difficultto distinguish; in Nymphulis (present data) it is more evident.
Metathoracic sternopleural region (Figs 10, 19-2 1, 23)
The metathoracic sternopleural region resembles its mesothoracic
counterpart; only the differences are discussed here.
The anapleural cleft of the episternum is sulcate, and the anepisternum is
fused anteroventrally with the infraepisternum. The basalare is tripartite (Fig. 19)
and has a FIRST BASALARE (anterior), a BASALAR PROCESS (dorsal), and a BASALAR
PAD (posteroventral). The basalar process is a composite structure resulting,
according to Sharplin ( 1963b1, from fusion of the prescutal arm and the second
basalare. Sharplin also states that the first basalare is fused to the upper
episternum. In Glaucopsyche it is free (as on the mesothorax), but dorsally
connected to the basalar process by a short ligament. The basalare is fused
posteroventrally with the anepisternum, contrary to Sharplin’s ( 196313)statement
that such fusion with the episternum is restricted to the Zeugloptera,
Eriocranioidea and Hepialoidea. The basalar pad (Shepard, 1930)is present as a
sclerotized, ovoid cushion bearing numerous setae. Ehrlich ( 1958a)calls this pad
the anepisternum, while Sharplin (196313: fig. 36) illustrates it as the basalar scale
plate but does not mention it in her text. Matsuda (1970) considers the basalar
pad to correspond with the first basalare in some cases. This pad appears more
INTEGUMENTAL ANATOMY OF GLAUCOPSYCHE
81
closely associated with the ventral margin of the basalar process in Glaucopsyche,
and may represent a secondary, sensory structure. A TERGOPLEURAL APODEME is
absent; instead, Matsuda (1970) states that muscle t-p 4 (third tergopleural
muscle: Sharplin, 196313) occurs in a corresponding position in the lepidopteran
metathorax.
The INFRAEPISTERNAL-BASISTERNAL PLATE lacks a PARACOXAL SULCUS or a
PLEUROSTOMAL SULCUS, hence the existence or degree of fusion of the
preepisternurn, katepisternum and basisternum is uncertain. It is assumed that
the latter is at least partially invaginated at the MEDIAN LONGITUDINAL SULCUS,
forming the STERNAL RIDGE.A MARGINOPLEURAL SULCUS occurs along the
infraepisternal-basisternalplate’s ventral margin, between the PLEURAL SULCUS
and median longitudinal sulcus. The internal MARGINOPLEURAL RIDGE merges
with the PLEURAL RIDGE. A secondary PREPECTUS may occur along the
anteroventral infraepisternal-basisternalplate’s margin, but its demarcation is
vague.
The FURCA stem of the furcasternum is short, and produces an anterodorsal
hook and a pair ofwing-like apophyses (furcal apophyses: Brock, 197 1 ) dorsally.
The PRIMARY FURCAL ARMS resemble those of the mesothorax, but arise from the
anterior margin of the dorsal apophyses. The SECONDARY FURCAL ARMS arise
broadly along the furcal stem’s posterior margin, but narrow in their
posterolateral extension to the epimeron. The thoraco-abdominal conjunctive
membrane inserts anteriorly along the ventral margin of the secondary furcal
arm. Brock (197 1) outlines development of the metathoracic furca throughout
the Lepidoptera.
The E P I M E R O N resembles that of the mesothorax, but the PREEPIMERON,
PREEPIMERAL SULCUS and DORSAL EPIMERAL SULCUS are absent or very vague.
The VENTRAL EPIMERAL SULCUS is entire and relatively straight, with a strong
internal ridge. Fusions on the posterior part of the metathorax have clouded the
exact boundaries of the epimeral, furcal and postnotal contributions.
The SUBALARE is reduced, but produces a spatulate internal SUBALAR APODEME
anteriorly. The METATHORACIC SPIRACLES occur near the posterior margin of the
mesothoracic epimeron. Sharplin ( 1963b) notes that the short ligament
connecting the detached prescutal apodeme to mesothoracic laterophragma 2 is
immediatelv adjacent to the metathoracic spiracles, and that Nuesch ( 1953)
erroneous$ considered it to be the detached dilator muscle.
Prothoracic legs (Figs 10, 26-33)
References to orientation of all legs follows Snodgrass ( 1935: 193): the preaxial
surface is anterior, the postaxial surface posterior, the outer surface dorsal, and
the inner surface ventral. All legs consist of a basipodite, represented by the coxa
and trochanter, and a tebpodite, represented by all the distal components
(Srivastava, 1961).
The prothoracic legs are reduced compared to the pterothoracic legs. The
coxo-pleural articulation is monocondylic, allowing the coxa rotational
mobility. The coxa is simple and cylindrical. The proximal margin is attached to
the body by the COXAL CORIUM membrane h o d g r a s s , 1935), and is
circumscribed by a BASICOSTAL SULCUS. This sulcus produces an internal
basicostal ridge, the BASICOSTA, that forms an expanded apodeme anteriorly.
6
82
24
1
J. T. SORENSEN
COXO-PLEURAL ART1 CULATI 014
A B A S I C O S T A L SULCUS
-
COXA GENUINA
COXA GENUINA
COXAL SUl
TROCHANTER
MERON
"\\
-
COXAL SULCllS
COXA GE
\
..".....
TROCHANTER
'
FEMUR
\\
/
27
28
29
Figures 24-29. Fig. 24. Right metathoracic leg, lateral view (anterior to right). Fig. 25. Right
mesothoracic leg (male), lateral view (anterior to right). Fig. 26. Right prothoracic leg imale), laterdl
view (anterior to right). Fig. 27. Prothoracic tarsus (female), lateral view (anterior to right). Fig. 28.
Prothoracic tarsus (female), ventral view. Fig. 29. Prothoracic ankylosed tarsus iniale), ventral view;
note fused tarsomeres depicted by tarsal spination pattern.
The narrow coxal margin, proximal to the basicostal sulcus, represents the
BASICOXITE. The coxa distal to the basicostal sulcus is the COXA GENUINA. N o
coxal sulcus or coxal ridge is present, as described for Papilio by Srivastava ( 196 1 ) .
The TROCHANTER, FEMUR, and TIBIA follow distally, with the tibia showing an
apical TIBIAL PROCESS dorsodistally. TIBIAL SPINES may occur ventrodistally on
some individuals. The coxo-trochanteral, femoro-tibia1 and tibio-tarsal
articulations are dicondylic, with an anteroposterior axis. The trochanterofemoral articulation is monocondylic, occurring ventrally. The membrane of the
leg joints is the ARTICULAR CORIUM (Snodgrass, 1935).
The prothoracic TARSUS is sexually dimorphic (Figs 26-29). In females, five
TARSOMERES are present, with the proximal and distal ones representing the
BASITARSUS and DISTITARSUS, respectively. The tarsomeres have non-ranked
INTEGUMENTAL ANATOMY O F GLAIICOPSYCHL
UNGUITRACTOR TENDON
UNGUITRACTOR PLATE
~
BASIPULVILLUS
MACROTRACHIA
U I ST I TARSUS
SUBSIDIARY SETA
PULVl LLUS
ENDODONT LOBE
EMPOD I UM
EMPODIAL SETA
CAMERA
AROLl LIM
-SUBSIDI
33
32
Figurr\ 3&33. big. 30. Pretarsus, lateral view. Fig. 31. Prerarsus, ventral
distal viekv. Fig. 33. Prothoracic pretarsus (male), lateral view.
ARY
SETA
\im.
big. 3 2 . P1wtCiiws,
SPINES ventrally, with clusters at the ventrodistal margins. The
prothoracic tarsus of the male is ankylosed, but shows spine clustering similar to
the female’s tarsomere margins, thus delimiting the fused tarsomeres of the
male.
The PRETARSLS represents the most distal region of the leg, and also shows
sexual dimorphism on the prothoracic legs (Figs 30-33). The terminology of the
pretarsus is confused throughout the literature, due to poorly defined o r
specialized terms. Homologies between orders are poorly known, except for the
early efforts of Crampton (1923) and Holoway (1935). Dashman’s (1953) attempt
to standardize pretarsal terminology suffers from misprints and selfcontradiction, and has not been readily adopted according to Goel & Schaefer
( 1970), who attempt to establish a consistent, homologous terminology among
the orders, based on the concepts of Crampton (1923), Holoway (1935) and
Snodgrass (1935). Goel & Schaefer (1970) are followed here, except that the term
pretarsus is retained because of universal acceptance. Additions, following
TARSAL
84
J. T. SORENSEN
Clench (1955), are made where structures appear to be derived within the
Lepidoptera.
The prothoracic pretarsus of the female bears a pair of claw-like UNGUES that
articulate laterally against the ventral lobes of the distolateral margins of the
distitarsus. The inner margin of the ungues bears an ENDODONT LOBE (Clench,
1955) subequal to the ungues tip. A semi-sclerotized, pad-like BASIPULVILLUS is
produced from the ventrodistal margin of the distitarsus, and is attached
internally to the UNGUITRACTOR TENDON. The basipulvillus distolaterally
produces four membranous ~ U L V I L L Ithat bear numerous minute setae. A
membranous, pad-like AROLIUM occurs ventromedially, between the ungues,
and ventrally bears a sclerotized horseshoe-like CAMERA. Dorsally, the arolium
bears a narrow sclerotized EMPODIUM that produces a single bristle-like
EMPODIAL SETA. A pair of long, coarse MACROTRACHIA (Clench, 1955) arise
medially from the dorsodistal margin of the distitarsus, and extend beyond the
ungual tips. A pair of shorter bristle-like SUBSIDIARY SETAE (Clench, 1955) flank
the macrotrachia. Other minor auxiliary setae occur randomly on the distal
distitarsus.
The prothoracic pretarsus of the male is greatly reduced and fused with the
ankylosed tarsus. The only vestiges are a pair of rudimentary macrotrachia and
several minute subsidiary setae that project from the dorsodistal surface of the
ventrally curved tarsal tip.
Mesothoracic legs (Figs 10, 12, 15, 16, 25)
The pro- and mesothoracic legs are generally similar except for size
differences, coxal divisions, and the presence of a tibia1 brush and complete
tarsus and pretarsus in the male. The pterothoracic coxae are immobile and
ventrally reduced to accommodate the acute projection angle from the body. The
coxo-pleural articulation is fused with the pleural ridge and the ventral coxal
articulation is fused with the furcasternal base. The COXAL CORIUM is reduced,
except posteriorly where it is continuous with the intersegmental membrane.
Interpretations of the homology of coxal divisions are varied in the literature;
some terminologies confuse homology. The BASICOSTAL SULCUS is marginal,
anterior to the coxo-pleural articulation, and produces an enlarged BASICOSTA
internally (eucoxal apodeme: Ehrlich, 1958a). The BASICOXITE of this region
(epicoxal piece: Shepard, 1930) is narrow or absent. Posterior to this articulation, the coxal margin curves proximally, but is not followed by the basicostal
sulcus. Thus, a portion of the postarticular basicoxite, the MERON (Snodgrass,
19351, is expanded proximally. Ehrlich (1958a) refers to this meron and its
immediately associated basicostal sulcus, as the postcoxal sclerite and postcoxal
sulcus. He apparently considers the basicoxite to be restricted to the anterior part
of the coxo-pleural articulation, and to be equivalent to Shepard’s (1930)
epicoxal piece.
A COXAL SULCUS and internal COXAL RIDGE dorsally divide the coxa below the
basicostal sulcus into an ANTERIOR and POSTERIOR C o r n GENUINA. Shepard
(19301, Weber (1933) and Snodgrass (1935) consider the posterior region of the
lepidopteran pterothoracic coxa to represent a distally enlarged meron, and
believe that the basicostal sulcus runs distally, posteriad to the coxo-pleural
articulation. Srivastava ( 1962) considers this interpretation erroneous for the
INTEGUMENTAL ANATOMY OF GLAUCOPSYCHE
85
mesothoracic coxae, since their posterior division includes the dorsal coxotrochanteral articulation, whereas the meron cannot-according t o Snodgrass
( 1935: 196). In Glaucopsyche the ventral and dorsal coxo-trochanteral articulations
arise respectively on the anterior and posterior coxa genuina. This may represent
a primitive condition since eriocraniids also show it (present data). Matsuda
( 1970 : fig. 14 7a) erroneously depicts both coxo-trochanteral articulations as
rising from the posterior division of the coxa genuina in the Zeugloptera. This
condition should not exist and probably represents an artifact in illustration. The
coxal ridge expands distally to reinforce the dorsal coxo-trochanteral
articulation anteriorly.
Several terms exist to describe a coxa genuina divided by a true coxal sulcus,
but are unacceptable due to confusion of homology. Srivastava ( 1962) applies
coxal episternum and coxal epimeron to the anterior and posterior coxa
genuina, respectively, citing analogy with the pleural divisions. Ehrlich ( 1958a)
appears to follow Crampton ( 19231, who applies eucoxa and meron, respectively,
to these divisions. Crampton’s use of meron is not equivalent to Snodgrass’s
( 1935) definition, which is commonly recognized as the postarticular section of
the basicoxite. Crampton’s use of eucoxa, for the coxa genuina anterior to the
coxal sulcus (his ‘meral’ sulcus), is avoided because of confusion caused by the
co-definition with his misapplied term meron.
The plates of the coxa genuina are separated ventrally, and a triangular
sclerotized plate, the STERNOCOXALE, bridges their gap proximally and is fused
with them. The coxo-trochanteral articular corium extends proximally to the
sternocoxale, bridging the gap distally. This ARTICULAR CORIUM is completely
membranous posteriorly, lacking any sclerotized intrusions (coxal sclerule :
Ehrlich, 1958a).
The ventral margin of the tibia bears a TIBIAL BRUSH of persistent setae in
males ; this is used in antenna1 grooming.
Metathoracic legs (Figs 10, 20, 2 1, 24)
The meso- and metathoracic legs are similar except for the divisions of the
coxae and the absence of a tibia1 brush o n the latter. The metathoracic coxa has a
distally enlarged MERON, as described by Snodgrass (1935). The prearticular
BASICOXITEis narrow o r absent, and the BASICOSTAL SULCUS is marginal anterior
to the coxo-pleural articulation but runs distally along the same course as the
mesothoracic coxal sulcus once it reaches the coxo-pleural articulation point. I t
curves posteriorly, however, immediately before reaching the distal margin of
the coxa, so that the dorsal coxo-trochanteral articulation arises from the
anterior division of the coxa, which represents the COXA GENUINA. The basicostal
sulcus skirts this articulation proximally, then weakens and curves proximally to
obliteration on the ventral surface of the coxa. The posterior division of the
metathoracic coxa thus represents a true meron of basicostal derivation. The
BASICOSTAL RIDGE expands slightly to reinforce the dorsal coxo- trochanteral
articulation dorsally. Srivastava (1962) considers the coxal sulcus to occur
posteroventrally on this coxa in Papilio. N o such sulcus exists in Glaucopsyche, and
the COXAL SULCUS is considered lost. The proximal edge of the meron has a
secondary inflected sulcus, the SECONDARY COXAL SULCUS, which can cause
confusion by its resemblance to the postarticular basicostal sulcus of the
86
J. T. SORENSEN
mesothoracic coxa. The STERNOCOXALE is somewhat enlarged and almost
indistinguishably fused with the coxa genuina ventrally.
Mesothoracic wings and wing base sclerites (Figs 34, 36, 37)
Despite widespread use in systematics, wing venation systems among insect
orders remain difficult to homologize. Much of the problem apparently lies in
contradictions and confusions among the numerous terminologies available;
Wootton (1979) reviews the history of the major schemes. Recently, Hamilton
( 197 1 ; 1972a, b, c) has discussed the evolution of wing venation and proposed a
terminology and homology, but his introduction of new terms (‘empusal’)adds
unnecessary confusion. Wootton ( 1979)provides a relatively simple, conservative
venation terminology, with homology based in part upon the structural
morphology of the veins in relation to flexion of wing regions. His
recommendations are followed here.
The mesothoracic wing has a SUBCOSTA, a RADIUS, a 3-branched RADIAL
SECTOR, three MEDIA veins, a 2-branched ANTERIOR CUBITUS, and two ANAL
veins. It lacks a COSTA and a POSTERIOR CUBITUS. The radial sector branches
are designated 1, 2 and 3 + 4. Comstock 8c Needham (1898-1 899) and Snodgrass
( 1935) refer to the radius as radius 1, and to the radial sector branches as radius
2, 3 and 4 + 5. The three media veins represent the ANTERIOR and POSTERIOR
MEDIAwhich have become partially obliterated in the formation of the discal cell.
The surviving portion of the anterior media has fused with the radial sector
anterodistal to that cell, while the posterior media has fused with the anterior
cubitus, posterodistal to the discal cell. The media veins are designated simply as 1,
2 and 3 here, following Wootton’s (1979) recommendation, since i t is unclear
whether media 2 is derived from the anterior or posterior media. The anterior
media may be represented by media 1 and 2, and the posterior media solely by
media 3. A scar-like cross vein delimits the discal cell connecting the radial sector
and media vein systems.
The loss of the posterior cubitus (plical vein: Hamilton, 1972a, b, c;
postcubitus: Snodgrass, 1935; anal 1: Comstock 8c Needham, 1898-1899) is
incomplete and it remains detectable as a faint scar midway between anterior
cubitus 2 and anal 1. This loss is also reflected by the extra width of the cell
posterior to anterior cubitus 2 in comparison to its eers. A double eyespot
macule of the setal-scaling is also present in that cell ue to coalescence of the
eyespots of the previously separate cells.
Anal 1 (empusal vein: Hamilton, 1972a, b, c; vannal 1 : Snodgrass, 1935; anal
2: Comstock & Needham, 1898-1899) is developed, but anal 2 (anal 1 :
Hamilton, 1972a, b, c; vannal 2: Snodgrass, 1935; anal 3: Comstock &
Needham, 1898-1899) is present only as a rudiment. Such a rudimentary anal
vein is present in hesperiids, haeterine satyrids, papilionids, ithomiids and
danaids, as well as lycaenids, but is apparently absent in nymphalids, most
satyrids, pierids and riodinids.
The wings articulate by a series of complex, minute sclerites contained between
the DORSAL and VENTRAL ALARY MEMBRANES. This region is treated by a number
of authors (Snodgrass, 1909; Crampton, 1928; Jordan, 1928; Ehrlich, 1958a;
Onesto, 1959; many others), but their unique terminologies give a multitude of
confusing synonymous names for each component, often with erroneous
x
INTEGUMENTAL ANATOMY OF GLAUCOPSYCHL
RADIAL SECTOR 2.
.
RADIAL SECTOR
, , l A
RADIUS
SUBCOSTA -
-
\ ANAL 2 RUDIMENT
87
/ RADIAL SECTOR 3 + 4
G
--MEDIUS 1
-
MEDIUS 2
-MEDIUS 3
\ANAL
34
Figui-e\ 34. 55. Fig. 34. Right mesothoracic wing, dorsal view. Fig. 35. Right
dorsal vie\\
35
irirratlioi-dc
i( ~ m g ,
homologies. Sharplin ( 1963a, b) provides an excellent, broad-based treatment of
this region in the order, and gives a partial synonomy of divergent terminologies.
Her terms (following Snodgrass, 1909) are used here.
The FIRST AXILLARY SCLERITE is only visible dorsally. It represents a detached
lateromedial portion of the notum and is enlarged (along with the median notal
wing process) in advanced Lepidoptera (Sharplin, 1963a). The first axillary
associates with a BASAL PROCESS OF THE SUBCOSTA, and articulates with the
suralare anteroproximally, with the second axillary anterodistally, and with the
median notal wing process posteriorly. Sharplin (1963a) states that it has
ligament connectives to the suralare and median notal wing process.
The second axillary sclerite is uniquely doubled in the Lepidoptera and
Trichoptera (Sharplin, 1963a1, forming DORSAL and VENTRAL SECOND AXILLARY
SCLERITES, which together vertically span the thickness of the wing base. These
subcomponents articulate at a peg-in-socket structure contained between the
alary membranes. According to Sharplin the dorsal second axillary is
hornologous t o the generalized second axillary o f Snodgras\ w1iic.h
originates from the basal portion of the radial vein. The ventral second axillary is
formed from a detached ventral part of the dorsal second axillary that has fused
with a second invagination of the proximal part of the radial vein. The dorsal
second axillarv articulates with the first axillary proximally, and with thc first
median plate posterodistally. I t is associated anteriorly with a dorsal pi.oximal
process of the radius that Sharplin (1963a)calls theRADIAL PLATE. (second a x i h r v :
Ehrlich, 1958a: fig. 28). The ventral second axillary has ligament conne(.tivcs\\.itti
the first median plate, the subalare and the pleural wing process; i t articulates
against the latter ventrally.
The proximal portions of' the subcosta and radial veins jointly produce
ventrally a compound basal process which is suspended from the pleural wing
process by two ligaments, but does not articulate directly against it (Sharplin,
1963a). Sharplin (1963a) does not name this structure; Ehrlich's (19.5Xai use of
radial plate for i t is avoided here due to confusion of homology; I refer to it as
the VENTRAL SUBCOSTAL-RADIAL PROCESS.
The THIRD AXILLARY SCLERITE spans the wing base vertically, and is associated
distally with the first anal and second anal rudiment veins. It does not directly
articulate with the second axillary; instead it is fused anteriorly with the first
median plate, which in turn articulates with the second axillary. The third axillary
articulates posteriorly with an elongate extension of the posterior notal wing
J. T. SORENSEN
88
37
\PLEURAL
WING PROCESS
Figures 36, 37. Left mesothoracic axillary region. Fig. 36. Dorsal view (anterior to lelt); wing base
stretched distally to show notal articulations. Fig. 37. Lateral view (anterior to left,; wing held
upright to show ventral aspect.
process. Madden (19441, Nuesch (1953)and Ehrlich (1958a) consider this process
to be a fourth axillary sclerite. When the wings is fully depressed, the ventral
portion of the third axillary fits into the depression on the dorsal margin of the
subalare.
The median plates are separated. The FIRST MEDIAN PLATE is fused to the third
axillary and articulates with the dorsal second axillary. I t reaches the ventral
alary membrane posterior to the ventral second axillary. The SECOND MEDIAN
PLATE is softened and cowacts the radial plate proximally, but is not fused with it
(Sharplin, 1963a).The cubital vein base is associated with this plate, but does not
overlie it. Hamilton (1972b) considers the plate associated with the base of the
cubitus to be the cubital plate. The sclerotized base of the first anal overrides
both median plates slightly. Sharplin (1963a) believes that the secondary loss of
wing flexion over the tergum in the papilionoids may account for the unique
condition of median plate separation and the lack of a cubital base overlap of the
second median plate.
INTEGUMENTAL ANATOMY O F GLAUCOPSYCHf
89
The HUMERAL PLATE occurs as a semi-detached lobe on the wing’s
anteroproximal margin. The TEGULA is represented by an enlarged, sclerotized,
shoulderpad-like lobe proximal to the humeral plate. I t is semi-detached and
articulates with the wing margin by a membranous connection. The tegula bears
a posterior lobe that covers the axillary region dorsally, and a ventral lobe that
shields the basalar region. The proximal surface is concave and is supported by,
but is unattached to, the subtegula.
.Metathoracic wings and wing base sclerites (Figs 35, 38, 39)
This region is generally similar to its mesothoracic counterpart and only the
differences are discussed. The subcosta combines with the hrst radial vein to form
a composite SUBCOSTA+RADIUS while the RADIAL SECTOR is unbranched. ANAL 2
is developed. A HUMERAL vein, often present in other lepidopteran groups, is
absent.
All three axillary sclerites are present, although their shapes differ somewhat
AXILLARY SCLERITE 1
DORSAL ALARY MEMBRANE
E R I O R NOTAL
WING
PROCESS
SUBCOSTA + RADIUS
AXILLARY SCLERITE 3
VENTRAL ALARY MEMBRANE
PLEURAL SULCUS
PLEURAL WING PROCESS
Figure, 38, 39 Lett inetathoracic axillary region Fig. 38 Dorsal view (anteiior to lelti, 7411ng- base
atretched distallv to show notal articulations Flg 39 Lateral vww ianlcrwl tc) lcft), M L ~ held
I ~
upright t o s h o \ v ventral aspect
7
90
J. T. SORENSEN
from those on the mesothorax. The FIRST AXILLARY SCLERITE does not articulate
against an enlarged median notal wing process, but against the undifferentiated
lateral margin of the scutum. Sharplin (196313) believes that the metathoracic
median plates are represented by a single sclerite in the Lepidoptera, except the
Zeugloptera and the Eriocranioidea, and that this single MEDIAN PLATE is
associated with the anterior part of the third axillary, as on the mesothorax. In
Glaucopsyche this association, pr any differentiation between the two, is
undetectable and the entire structure appears uniform. N o median arm or
cubital plates, as defined by Sharplin (1963b), are evident. A BASAL PROCESS OF
THE SUBCOSTA is present, but a RADIAL PLATE homologous to that of the
mesothorax is absent. A structure analogous to the radial plate is represented by
a BASAL PROCESS OF THE CUBITUS, according to Sharplin (1963b). This process
produces a narrow lobe which reaches around the anterior margin of the wing to
the ventral surface, to merge with the ventral subcostal-radial process; Ehrlich
( 1958a)refers to this anterior lobe as the costal sclerite.
Pregenitalic abdomen (Figs 10, 40, 4 1)
The pregenitalic abdomen has seven telescopic segments, each with spiracles.
Segments 3 through 7 are simplified with a rectangular TERGUM and
subtriangular STERNUM separated by a membranous pleural region. Antecostae
are absent from these segments. Srivastava (1966) implies that they are also
absent in Papilio, but Arnold 8c Fisher (1977) report their presence in Speyeria.
Segments 1 and 2 are modified for thoraco-abdominal articulation. The
acrotergite and antecosta of tergum 1 are separated, and represent the metathoracic postnotum. The abdominal portion is membranous anterodorsally,
but becomes sclerotized posteriorly. This tergum is laterally divided, forming a
LATERAL SECONDARY SCLERITE (laterotergite: Srivastava, 1966) which bears a
TERGAL GROOVE (Ehrlich, 1958a)marking an invagination that forms an internal
TERGAL BRACE (Ehrlich, 1958a; Brock, 1971). The anterior part of the brace is
fused with metathoracic laterophragma 3, creating the dorsal thoracoabdominal articulation. A small, setous, semi-sclerotized TERGAL LOBE (Ehrlich,
1958a1, resembling the metathoracic basalar pad, occurs on the anterolateral
margin of this tergum. This lobe contacts the posterior end of the metathorax
when abdominal segment 1 is displaced laterally or dorsally, and it may be a
sensor for abdominal orientation.
Terga 1 and 2 are separated by the latter’s antecostal sulcus and well developed
antecosta. Their common margin produces an anteroventrally extending
tergopleural process, the POSTSPIRACULAR BAR (Ehrlich, 1958a). This structure
has a median sulcus that invaginates, forming an internal ridge which is dorsally
associated with the tergal brace and the lateral portion of the antecosta oftergum 2 .
The occurrence and extent of sterna 1 and 2 are uncertain. Sternum 1 may be
lost, at least partially, to the ventral thoraco-abdominal conjunctive membrane
(see Brock, 197 1). Sternum 2 has moved anteriad to partially underlie tergum 1,
and its anterolateral corners are produced anterodorsally into the pleural region
to connect with a bowed PRESPIRACULAR BAR (Ehrlich, 1958a; tergalsternal rod:
Brock, 1971). This, in turn, articulates with the tergal lobe of tergum 1. The
resultant sterno-tergal connection represents the ventral thoraco-abdominal
articulation.
INTEGUMENTAL ANATOMY OF GLAUCOPSYCHE
q\T/
TERGUM 4\
A n
4u
TERGUM
\ 3
DOWNEY'S AREA
TERGUM 2
TERGUM
91
TERGUM 5
h\q /K
\
'
I
b<
TERGUM 6
q
\
'J
77'
STERNUM 1 7
STERNUM 2
f
/ //
STERNUM
STERNUM
3' 4
STERNUM 5 '
STERNUM 6'
"---. ,..
7
/ /
LATERAL SECONDAFY SCI-ERlTE
POSTSPIRACULAR EAR
PRESPIRACULAR BAR
41
TERGUM 2
STERNUM 2
T h e anterior margin a n d anterolateral process of sternum 2 a r e
incompletelv separated, as a narrow band, from the sternum by a transverse,
medially obscured sulcus which laterally produces an internal ridge. Ehrlich
(1958a) considers this anterior sclerite (see Fig. 10) to represent a portion o f
sternum 1 which has fused with sternum 2. Brock (197 1) believes i t represents a
secondary sclerite derived from sternum 2. An equivalent anterior di\ision is
absent in some hesperiids (Brock, 197 1) and papilionoids (Srivastava, 19661,
although the sterno-tergal connective remains intact in these g.r-oupr. Biiriier
(1939) depicts sternum 1 as separate and reduced in the Zeugloptera and
Wpialidae. The narrow anterior sclerite and its demarcating sulcus and associated
internal ridge may represent, respectively, the acrosternite, antcwxtal sulcus and
antecosta of sternum 2. Ehrlich 8c Davidson (1961) and Srivastava (19663 indicate
that the muscles inserting on this area in part originate from the nictathoracic
furca. The evolution of this sternal area remains unclear arid both Ehl-lich's
(Fig. 10)and Brock's (Fig. 41) concepts are shown here.
Brock ( 197 1) believes that the pre- and postspiracular bars of papilionoids
represent secondary structures that have developed within the group. He
92
J. T. SORENSEN
considers similar structures in some of the Pyraloidea, Geometroidea and
Noctuoidea to be of separate derivation.
Glaucopsyche shows several sexual differences on abdominal segments 1 and 2.
Males show greater development of the postspiracular bar, and tergum 1 is more
protuberant, with heavier sclerotization. They have an additional unique area of
sclerotization on the posterolateral surface of tergum 2. The latter is unreported,
but J. C. Downey (personal communication) states that it occurs generally among
male ‘Plebejinae’. I call this structure DOWNEY’S AREA, after its discoverer. In
Glaucopsyche it is elliptical and may contact the tergum margin laterally, showing a
degree of development that is positively correlated with degree of sclerotization
of the male. The cuticle of Downey’s area shows prominent reticulation and
mosaic texture, yet scanning electron microscopy reveals no surface differentiation from the surrounding cuticle, from which it must be concluded that the
mosaic appearance of the area results from sub-surface cuticular structure.
Initial histological studies reveal that no specialized sub-cuticular structure is
present, but the intra-cuticular structure appears to differ from the surrounding
tergum. Further studies with improved resolution might determine the precise
structure of the area.
Genitalic abdomen and genitalia
The derivation and homology of insect genitalia are exceedingly difficult
questions. Numerous unique theories have been proposed and two camps exist:
one espousing an abdominal appendicular origin, based largely upon
interordinal comparisons of adults (Michener, 194413; Dupuis, 1950; Nielsen,
1957; Scudder, 1957, 1961; Sharov, 1966; Smith, 19691, the other advocating a
sternal integumental derivation, usually employing ontogenetic evidence
(Gustafson, 1950; Snodgrass, 1957; Matsuda, 1958, 1976); some theories
‘hybridize’ these (Stys, 1959). Unfortunately, most workers advance unique
morphological terminologies which reflect their biases. Tuxen ( 1969) and
Matsuda (1976) review some of these theories; the former is unbiased. This
treatment is neutral regarding derivation, since both philosophies have inherent
problems and are currently too unstable for employment (see Tuxen, 1969).
Comments on derivation are expressed conservatively.
Since the area is of great systematic importance, numerous taxonomic
(morphologically meaningless) terms have been applied making the
nomenclature of the region almost hopelessly entangled. Beirne ( 1942) estimates
that an average of six or more such terms exist for each structure in
lepidopteran genitalia, while Niculescu ( 1973) states that about 900 terms have
been appied for the order. Klots (1970) provides an extensive glossary of
taxonomic terms for the region, but Sibatani (1972) and Niculescu (1973)
comment critically on parts of it. Michener (1958)and Tuxen (1969) discuss the
relative merits of a morphological versus taxonomic terminology for the region.
I follow the latter’s recommendation and partially incorporate Klots’s ( 1970)
taxonomic terminology, where appropriate, in italics in the text and within
parentheses in the figures. This is done largely because of the unstable
morphological interpretations available for this region, as well as the absence of
a complete, unbiased, generalized morphological terminology. Synonymy of
INTEGUMENTAL ANATOMY OF GLAUCOPSYCHt
93
taxonomic terminology is largely avoided here since synonyms are listed by Klots
( 1970) and Niculescu (1973).
The functional and muscular morphology of lepidopteran genitalia is treated
elsewhere (Stekol’nikov, 1965, 1967a, b ; Birket-Smith, 1974a, b ; Forbes, 1939;
Hannemann, 1954a, b ; Srivastava, 1966; Arnold 8c Fisher, 1977). Chapman
( 19 16) describes functional aspects of mating and also comments on anatomy of
the genitalia in the ‘Plebeiidi’.
There is disagreement regarding which abdominal segments have genitalic
involvement in the Lepidoptera. Warren (1957) considers only segment 10 as
genitalic. Ever (1924) and Srivastava (1966) consider segments 9 and 10 as
genitalic, despite the involvement of tergum 8 in the Papilionidae to form a
superuncus (see below) and sternum 8 in certain Danaidae to form pseudovalues
(Ehrlich, 1958a). Klots (1970) considers segments 7-10 as genitalic. I consider
segments 8-10 and possibly remnants of 11 (see below) to be genitalic in the
Papilionoidea ; these segments lack spiracles.
Male (Figs 42-46)
TERGUM 8 and STERNUM 8 show no differences from
TERGUM 9 and STERNUM 9 are fused, forming a ring
the preceding segments.
that supports the more
posterior structures. Sternum 9 is wishbone-shaped and is represented by the
vinculum and saccus. Tergum 9 is represented by a roof-like tegumen.
A pair of large spatulate claspers, the vulvae, are fused with sternum 9
ventrally. Although the derivation (sternal o r appendicular) of the valvae is
unclear, thev probably pertain to segment 9. Since the vulvae vary greatly among
taxa, their taxonomic importance has prompted several attempts to subdivide
and name their various portions. Many of these attempts (Petersen, 1904; Pierce,
1909, 1914; Warren, 1926) fail to provide a satisfactory generalized format
because of their author’s restrictive group orientation, but others (Sibatani et al.,
1954; possibly Birket-Smith, 1974a, b) deal more successfully with the problem;
see Klots (19701, Sibatani (1972)and Niculescu (1973) for commentary.
SEGMENT 10 is represented by the pathos, a pair of slender curved arms fused
to the posterolateral margin of tergum 9. The pathos has also been called the
subunci, f a k e s and brachia in various groups (see Ogata et al., 1957). Mehta (1933)
believes that the pathos is derived from pupal sternum 10. The pathos varies in
different groups, and its arms may fuse below the anus; all of its variants may not
be homologous.
The posterodorsal margin of tergum 9 bears a modified bilobed structure
representing the semi-fused socii (see Ogata et al., 1957). The socii are usually
paired in non-lycaenids. The lycaenid socii are commonly referred to as the
labides.
The Lycaenidae lack an uncus, which is usually present in other non-lycaenid
papilioniods as a single, median, hooked lobe borne o n the posterior margin of
tergum 9. The Papilionidae are also an exception in lacking an uncus, but they
possess instead an analagous superuncus derived from tergum 8. A similar
development also occurs in one Calinaginae (Nymphalidae).
The relationship between the socii and uncus is unclear. Ogata rt ul. (1957)
consider both to represent a partially equivalent morphological unit and state
that these are continuous when both are present. They state that the flexor
J . T. SORENSEN
94
TERGUM 9 (TEGUMEN)
(SOCIUS) SEGMENT 10 OR 11 ?
(GNATHOS) SEGMENT 10
42
(JUXTA) SEGMENT 9 ?
STERNUM 9
(VALVA) SEGMENT 9
TERGUM 9 (TEGUMEN)
(SOCIUS) SEGMENT 10 OR 11 7’
ANUS
43
(GNATHOS) SEGMENT 10
(ANELLUS)
SEGMENT 9 ? (JUXTA)
(VINCULUM)
1
STERNUM 9
STERNUM 9 (VINCULUM)
(JUXTA) SEGMENT 9 ?
TERGUM 9 (TEGUMEN)
(ANELLUS)
(SOCIUS) SEGMENT 10 OR 11
SEGMENT 9 (VALVA)
44
SEGMENT 10 (GNATHOS)
Figures 42-44 Genitalia (male) Fig 42. Left lateral view (anterior to left), genitalia extended, intersegmental membrane 8-9 removed, phallus removed Taxonomic terms in parentheses Fig 43
Posterior view; genitalia extended, phallus removed Taxonomic terms in parentheses Fig 44
Dorsal view (anterior to top); genitalia extended, 8-9 intersegmental membrane removed, phallus
removed Taxonomic terms in parentheses.
muscles from the tegumen (tergum 9) insert on the uncus when it alone is present,
but that they insert on the socii when both the socii and uncus occur together.
Forbes ( 1939) reports a similar muscle transfer in noctuids, but notes that when
the flexor muscles from the tegumen (tergum 9) insert on the socii (his ‘pedunculi’),
the uncus is ankylosed with the tegumen. Sibatani (1972) proposes the term
sociuncus for the combined socii and uncus, which he considers a single functional
unit derived from the “10th somite”.
1 1 1 some lepidopterans, the uncus arises from paired imaginal discs, lateral to
the anus, which migrate dorsally and fuse during metamorphosis to form tergum
10 of the adult (Kosminsky, 1927a, b; Kosminsky & Golowinskaja, 1929; DuBois,
1931; Dewes, 1972). Matsuda (1976) suggests that the uncus, by replacing larval
segment 10 (embryonic segments 11 and 12) to become the adult tergum 10,
represents the supra-anal lobe of segment 12. He also suggests that the socii are
INTEGUMENTAL ANATOMY O F GLAUCOPSYLHL
95
probably homologous with the cerci of Trichoptera and represent remnants of
segment 11 which lie lateral to tergum 10. Matsuda believes that a similar
topographical relationship of these structures occurs throughout the
Holornetabola.
The 9th- 10th intersegmental membrane represents the diaphragma and seals
the abdominal haemocoel posteriorly. The phallus penetrates this membrane
through a conical extrusion termed the anellus. The manica represents the portion
of the anellus closest to the phallus. The ANUS also penetrates this membrane
forming a protruded membranous tube, the tuba analis, between the gnutho5 arms
(segment 10).The diaphragma is divided into thefultura superior andfultura inferior,
dorsal and ventral to the anellus, respectively (see Klots, 1970).
The P H A L L I ~ S , or aedeagus, has an enlarged phallobase anteriorly. The ductus
qaculatorius attaches internally between the lateral bulbous lobrc of’the, p h t r l l o h ~ ~ . ~ ~ .
Posteriorly the phallus bears an invaginated, extrudable membranous
E N D O P H A ~ ~ L I J So, r vesica, which has minute, sclerotized spines and hooks, the
conuti.
(CORNUTI)
/
ENDOPHALLUS ( V E S I C A
PHALLUS (AEDEAGUS)
(PHALLOBASE)
JJ(DUCTUS
EJACULATORIUS)
F i g u r e 4.5-47. Fig. 4 5 . Phallus, dorsal view (anterior to bottom); endophallus e x t c r i d d . Tzwiioniic.
t e r m in parentheses. Fig. 46. Phallus, left lateral view (anterior to b o t t o m ) ; endophallu\ eutriided.
Taxonomic t e r m in parentheses. Fig. 47. Lamella antevaginalis, ventral view iaritri i o r to h i t t 0 1 1 i 1 ;
plate removed from normal telescoped position within abdominal segment 7
The phallus is laterally supported by a slender V-shaped juxta that fuses
ventrally with sternum 9. I t is unclear if thejuxta is derived from sternum 9 or the
9th-10th intersegmental membrane. Snodgrass ( 1957) considers the vulvae, juxta
and aedeugus to arise from the primary phallic lobe. He applies the terms interparamere and paramere to thejuxta and vulva and states that the latter can be
homologized among the Holometabola. Matsuda (1976) and others dispute this.
Employment of these morphological terms is avoided here since their use would
have definite implications of homology and derivation.
Female (Figs 47-49)
The female genitalia are typical of the Ditrysia, with two genital openings.
They are representative of Mutuura’s ( 1972) “typical lepidopterous type”, with
J T SORENSEN
96
TERGUM 7
(APOPHYSES POSTERIORES)
(APOPHYSES ANTERIORE
TERGA 9 + 10
(PAPILLAE ANALES)
STERNUM 7
INUS VAGINALIS)
(LAMELLA ANTEVAG I N A L I S )
48
ANUS
TERGA 9 + 10
(PAPILLAE ANALES)
(SINUS VAGINALIS)
(LAMELLA ANTEVAG I NALl S )
(LAMELLA POSTVAG I NALl S)
VULVA
(OSTIUR BURSAE)
49
Figures 48, 49. Genitalia (female). Fig. 48. Left lateral view (anterior to left); teiminal abdominal
segments and genitalia extended, lamella antevaginalis extended and moved ventrally showing sinuc
vaginalis, internal components outlined with dashed lines. Taxonomic terms in parentheses. Fig. 49.
Posterior view; lamella antevaginalis extended and moved ventrally showing sinus vaginalis, vulva
and lamella postvaginalis. Taxonomic terms in parentheses.
structural modifications divergent from those of the Zeugloptera, apparently to
give a more flexible abdomen for oviposition on foodplant surfaces.
Segments 8 , 9 and 10 are represented by terga only; the sterna are lost. TERGUM
8 resembles a typical pregenitalic tergum, but is one-third longer. Its anterolateral
margins, under the overriding tergum 7 , are produced into short pointed
processes, the apophyses anteriores, which serve as retractor muscle anchorages.
Terga 9 and 10 are fused, forming a pair of lateral triangular plates, the papillae
anales. These are less sclerotized posteriorly and bear numerous long setae. Their
anterior margins produce narrow, elongate rods, the apophyses posteriores, which
also anchor retractor muscles. The papillae anales may be homologous, at least in
part, with the male uncus. They arise in some lepidopterans during metamorphosis
from the same paired imaginal discs on larval segment 10, but in females these
discs do not migrate dorsallyand fuse, as they do in males (Matsuda, 1976).
The ANUS occurs between terga 9+ 10 posteriorly, as does the OVIPORUS, or
ostium oviductus, which serves as the ovipositional aperture. A ventral membrane,
the sinus vaginalis, extends below terga 9+ 10 and 8, and invaginates to form the
insemination aperture, the VULVA or ostium bursae. In the Ditrysia, the vulva
represents a secondary invaginated aperture originating from the 7 th-8th intersegmental membrane. The oviporus in the Ditrysia has secondarily migrated
posteriorly to segment 9, from its original position on segment 8.
A large sub- triangular secondary sclerite, the lamella antevaginalis, is present
anterior and ventral to the vulva, and telescopes into segment 7 . This sclerite is
derived from the 7 th-8th intersegmental membrane, and can be protracted to
I h TE G UME NTAL ANATOMY 0 F GLA IJL OPSYCHE
97
move the vulva posteriad during mating. In lycaenids, this structure can easily
be mistaken for the absent 8th sternum, but its position anterior to the vulva
disproves such homology. A much smaller U-shaped secondary sclerite, the
lamella postuaginalis, flanks the vulva laterally and posteriorly. The lamella
anteuaginalis and postuaginalis are sometimes collectively referred to as the sterigma
o r genital plate. Chapman ( 19 16) refers to the lamella antevaginalis in the 'Plebeiidi'
as the hypostema. He refers to an eversible membranous tube, arising from the
lamella anteuaginalis to extend the vulva further during mating in some blues, as
the rein o r henia; this eversible tube is absent in Glaucopsyche.
ACKNOWLEDGEMENTS
I thank the following people for their contributions: Dr J . C. Downey,
University of Northern Iowa, for early guidance and specimens; Dr H. A. Hanian,
University of Northern Iowa, Dr R. Matsuda, Biosystematics Research
Institute-Canada Agriculture, J . Y. Miller, Allyn Museum of Entomology, and
Dr E. L. Smith, California Academy of Science, for information; Dr L. D. Miller,
Allyn Museum of Entomology, for specimens; Dr J. A. Powell, Unil-ersity of
California, Berkeley, for reviewing parts of the manuscript; and Dr R. L. Pipa,
University of California, Berkeley, for information, discussion5 and reviewing
parts of the manuscript.
I also thank M . Ramee for assistance during the early research, K. H . Sor-ensen
for manuscript assistance, and an anonymous reviewer.
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