Life Sciences Contributions 79
Royal Ontario Museum

A Review of the
North American Hawk Moth

Genus Lapara
(Lepidoptera: Sphingidae)

J. C. E. Riotte

RO M

—

LIFE SCIENCES CONTRIBUTIONS
ROYAL ONTARIO MUSEUM

NUMBER 79

J.C.E. RIOTTE A Review of the North American
Hawk Moth Genus Lapara

(Lepidoptera: Sphingidae)

Publication date: 2 February 1972

Suggested citation: Life Sci. Contr., R. Ont. Mus.

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CONTENTS

Abstract, 1
Introduction, 1
Methods, 2

Lapara Walker, 3

Generic Diagnosis, 3
Original Description, 3
Summary of Comparative Morphology, 3
Systematic Position of Lapara, 5
Phylogenetic Relationships of Species, 6
Biology, 7
Key to Adults Based on External Features, 8
Key to Adults Based on Genitalic Features, 8
Key to Mature Larvae, 9
Lapara halicarnie (Strecker), 10
Lapara coniferarum (J.E. Smith), 13
Lapara bombycoides Walker, 17
Incerta sedis: Lapara pinea (Lintner), 23
Discussion, 25
Acknowledgments, 26

Literature Cited, 27

Figures, 31

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A Review of the North American
Hawk Moth Genus Lapara
(Lepidoptera: Sphingidae)

Abstract

The status of Lapara species has long been in doubt. Examination
of new material has produced additional evidence from the
(adult) genitalia, head, epiphysis and pretarsus, as well as from
the larval and pupal stages, confirming the validity of the three
traditional species, L. halicarnie (Strecker), L. coniferarum (J. E.
Smith) and L. bombycoides Walker.

The status of a fourth taxon, L. pinea (Lintner) remains un-
certain. Described from two specimens with characteristics inter-
mediate between L. coniferarum and L. bombycoides, it has not
been recorded since its original discovery in 1859.

Introduction

Moths of the North American sphingid genus Lapara occur over much of the
eastern half of the continent, from Nova Scotia south to Florida and west to
British Columbia and Nebraska. In their classical revision of the sphingids, Roths-
child and Jordan (1903) assigned four species to Lapara: coniferarum, pineum
[sic], bombycoides, and halicarniae |sic|. The status of these forms has been
controversial among students of the Sphingidae for many years. The most
directly opposing view is that of Forbes (1948), who suggested that, apart from
L. halicarnie, the other taxa should be regarded only as races of a second species.
Clearly, then, additional evidence to resolve these conflicting views has been
needed.

My examination of specimens indicates that certain characters in this genus
were not fully appreciated by previous workers. Enlarging upon these, I made a
comparative study of specimens from many parts of the range of the genus and
examined all available associated larval and pupal material. The evidence ob-
tained adds strength to the view that there are three valid species of Lapara
apparently sympatric over wide areas of their range. The status of a fourth
taxon, L. pinea, is doubtful.

After the present paper had been submitted for publication, Hodges’ (1971)
monograph of the Sphingoidea appeared in print. His treatment of Lapara
(ibid.: 72—76) is an improvement over Forbes (1948), especially in according
full species status to both L. bombycoides and L. coniferarum. However, he
placed L. halicarnie as a synonym of L. coniferarum, a conclusion not supported
by my own findings. The present study also demonstrated that L. pinea is not
clearly a synonym of L. bombycoides as Hodges indicates (ibid.: 75).

Methods

For each species, the genitalia of a number of specimens from different popula-
tions representing widely separated portions of the range were dissected. Male
genitalia were dissected in detail to separate the left valve so that it could be
photographed and measured in a horizontal position. Each adult specimen dis-
sected was photographed with Kodak infra-red-sensitive film (IR 135) producing
a clearer definition of the wing pattern and other features, such as the clusters of
white hairs at the tegulae of L. bombycoides, than does standard film. The head
was denuded to examine the head structure and the labrum, and anterior legs
were removed from some specimens and cleaned for examination of the epiphy-
sis. In some the labial palpi were also removed, cleaned, and studied.

Localities and flight periods for specimens examined are cited under each
species account. Full specimen data have been deposited in the library of the
Royal Ontario Museum and are accessible there for reference. The abbreviations
for sites of specimen deposition are as follows:

AMNH The American Museum of Natural History
BM British Museum (Natural History)

CLU Clemson University, South Carolina

CM Carnegie Museum

CNC Canadian National Collection of Insects
FMNH Field Museum of Natural History

FSCA Florida State Collection of Arthropods
JGF J. G. Franclemont, Cornell University
LEM Lyman Entomological Museum

MCZ Museum of Comparative Zoology

MSU Michigan State University

NYSM New York State Museum

OSM Ohio State Museum

OSU Ohio State University

ROM Royal Ontario Museum

USNM United States National Museum

WES W. E. Sieker, Madison, Wisconsin

YPM or DCF Yale Peabody Museum of Natural History, or
D. C. Ferguson

Lapara Walker

Lapara Walker, 1856, p. 232 (type: bombycoides); Clemens, 1859, p. 187;
Beutenmiiller, 1895, p. 309; Rothschild and Jordan, 1903, p. 150; Holland,
1903% p.*533) Rethschild™and Jordan, 1907) p. 30; ‘Wagner; 1913). p.: 70;
Draudt, 1931, p. 860; Forbes, 1948, p. 192.

Ellema Clemens, 1859, p. 187 (type: harrisii); Fernald, 1886, p. 82; J. B. Smith,
1888; p.192;:206.

Exedrium Grote, 1882, p. 11 (type: halicarnie); J. B. Smith, 1888, p. 152.

Compared with other groups of Lepidoptera, the genus Lapara has enjoyed a
relatively uncomplicated taxonomic and nomenclatorial history. Although the
name was first used by Walker (1856) in association with the species bomby-
coides, the three species now included in Lapara were placed in other genera
during the second half of the 19th century. There is no doubt, however, that
Lapara has priority.

Rothschild and Jordan (1903: 150), in their monograph of the world sphin-
gids, and Wagner in the Lepidopterorum Catalogus (1913: 70) gave complete
bibliographies of earlier publications concerned with this group.

GENERIC DIAGNOSIS

Adult with antennae slender, subclavate, slightly longer than thorax, attenu-
ate, and weakly hooked distally; proboscis short, the labial palpus small; pulvil-
lus, paronychium with one lobe, and empodium present. Larva without caudal
horn, with eight or nine pairs of labral setae. Pupa with tongue case concealed.

ORIGINAL DESCRIPTION

“Body rather slender; head small, short; proboscis moderately long; palpi very
short; antennae slender, subclavate, hardly longer than the thorax, and attenu-
ated but weakly hooked towards the tips; abdomen linear, fully twice the length
of the thorax; legs slender; hind tibiae with four moderately long spurs; wings
rather narrow, not long; fore wings almost straight in front, slightly rounded at
the tips, straight and very oblique along the exterior border; interior border
straight; second interior vein far nearer to the first than to the third; fourth very
remote; hind wings rounded at tips” (Walker, 1856: 232).

SUMMARY OF COMPARATIVE MORPHOLOGY

Adult (Figs. 1—5, 10—20). The anterior wing has 11 veins and an oblique, evenly
rounded outer margin; the posterior wing shows a typical sphingid venation, the
outer margin is entire and rounded, but slightly excavated between 2d A and
Cus’.

The head is small, retracted; the vestiture forms an indistinct tuft between the
antennae, which are slender, fusiform, widest beyond the middle, and terminate
in a bent, ciliated seta; antennae biciliate in males, simple in females. The head
(Fig. la—c) is trapezoidal in shape. The eyes are moderately lashed with
blackish-brown bristles. The laterofrontal sutures are obscured by the convexity
of the sides of the frons. The labrum is roughly triangular, and its shape is
species specific (Fig. 2).

The proboscis is short, much shorter than in other members of the tribe
Sphingini, and the labial palpus is smaller; the pilifer is covered with short, stiff
bristles. The small third segment of the labial palpus appears as a little knob with
an apical pit containing sensory hairs (Fig. 3a—c). This third segment is identical
in the male and the female except in females of L. halicarnie, which have an
elongated third segment (Fig. 3a), a character that distinguishes them from large
coniferarum females occurring in the southern-most part of Mississippi.

The thorax is short and stout, rounded in front, and scarcely produced be-
yond the base of the anterior wings; vestiture thin but close.

The spines of the legs are not prominent; anterior tibia with one strongly
developed and elongate apical spine; middle tibia with a pair of spurs, one short
and one long; posterior tibia bearing a pair of subapical and a pair of apical
spurs, one apical spur considerably longer than the others, especially in L. hali-
carnie; secondary spines may be present or absent on spurs. The tarsus of the
middle leg lacks a comb, length of first segment approximately that of the other
four segments combined.

The epiphysis is a distinguishing specific character in sphingids (Mooser,
1940: 426), and this structure on the anterior leg differs strikingly in the three
species of Lapara (see Fig. 4).

The structure of the pretarsus is important for grouping sphingid genera and
is also important for grouping the species of Lapara (Fig. 5):

L halicarnie with well-developed pulvillus; paronychium with
one strong lateral lobe; empodium one tubercle
bearing a single seta (Fig. 5a);

L. coniferarum with pulvillus much reduced; paronychium with
one lateral lobe; empodium one tubercle bearing a
single seta;

L. bombycoides with pulvillus greatly reduced; paronychium rudi-
mentary; empodium of anterior pretarsi of males
two tubercles, each with one seta (Fig. 5b);
empodium of other male and all female pretarsi,
one tubercle bearing a single seta.

The abdomen is cylindrical and tapering, untufted; posterior edges of seg-
ments weakly spinulose.

In the male genitalia (Fig. 10) tegumen and vinculum are fused but visibly
distinct; uncus pyramidal, blunt, with small rounded projection; gnathos bifid at
tip but fused, usually blunt but more pointed in some specimens. Valve large,
sacculus with prominent process. Aedoeagus extended caudally into a pointed
process.

In the female genitalia (Figs. 15—18) signum absent on bursa copulatrix;
ostium bursae oval.

Larva (Figs. 6—8, 21). The larvae of the three species resemble one another in
lacking a caudal horn and in having a roughly triangular shaped head. Differences
in colour and the number of labral setae, however, clearly separate them into
two groups: L. halicarnie, of dull brown colour, with three pairs of additional
labral setae (Figs. 21a—b, 6a, 7a); L. coniferarum and L. bombycoides, of green

4

colour, with only two pairs of additional labral setae (Figs. 21c—d, 6b—c, 7b—c).
Forbes (1923: 31) discussed the setae on the labrum of lepidopterous larvae:
“There are a considerable number of setae which are constant in number and
position in any given species.”’ Although there are only two sets of setae on the
labrum of the larva in many moth species, L, —L,; and M, —M; (sensu Heinrich),
some sphingid larvae may have additional setae; e.g., the additional pairs illus-
trated by Bell and Scott (1937: 8, fig. 2) for a species of Polyptychus. Addi-
tional pairs of setae (designated X,—X,) are therefore a significant character in
demonstrating the relationships of the three species of Lapara.

Pupa (Fig. 9). The pupa is cylindrical. The tongue-case is probably concealed in
all three species of Lapara, a character which separates the group from moths of
the genus Sphinx. Apparently the only valid descriptions of the pupae of Lapara
in the literature are those of Beutenmiuller (1895) for L. coniferarum and L.
bombycoides (L. harrisii). The pupa of halicarnie has not been described. Mosher
(1918) worked specifically on sphingid pupae, but her conclusions regarding
the pupae of Lapara are questionable. She studied material that was purchased
and therefore of unconfirmed identification, a circumstance that may explain
why she did not comment upon the very different shape of the cremaster in L.
coniferarum and L. bombycoides.

SYSTEMATIC POSITION OF LAPARA

In colouring and size, the species of Lapara are similar to the east Asiatic species
of the Sphingulini, a tribe created by Rothschild and Jordan (1903) for several
genera including Kentrochrysalis and Sphingulus. Some of the anatomical fea-
tures discussed above for the adults as well as the larvae and pupae also place
Lapara near the Sphingulini.

When Rothschild and Jordan (1903: 154) separated the Sphingulini from the
Sphingini, they distinguished the two tribes primarily on differences of the
structure of the pretarsus and of the antenna in the adults, and the similarity of
larvae and pupae of the former to those of the Ambulicinae (i.e., Smerinthini).

The structure of the pretarsus is important in the taxonomy of these moths.
In their definition of the tribe Sphingini, Rothschild and Jordan (1903: 27)
referred to this structure as follows: ‘“‘Pulvillus and paronychium present or
absent, the pulvillus disappearing before the paronychium, there being no species
with pulvillus and without paronychium, the order in which these organs be-
come obliterated being this: ventral lobes of paronychium, pulvillus, lateral lobes
of paronychium.” For the tribe Sphingulini, however, they noted (1903: 154)
that all species have a pulvillus and that the paronychium is either with two
lobes or absent. In addition they stated (1903: LX VIII) that “the paronychium
of Sphingidae possesses in its most generalized state two slender lobes on each
side, of which the upper (or lateral) lobe is generally the longer, very rarely the
shorter one. The reduction of the paronychium begins with the ventral lobe; this
is always the first to disappear, there being no hawk moths which possess the
ventral lobe and are devoid of the lateral one. The remaining lobe becomes also
gradually shortened, and disappears too.”

I found all three species of Lapara to have a pulvillus (greatly reduced in L.
bombycoides) and a paronychium with only one (lateral) lobe (rudimentary in
L. bombycoides, Fig. 5). The empodium is well-developed in all three species,
especially in L. bombycoides, a surprising observation in light of Rothschild and
Jordan’s (1903: LXVIII) statement: ““The empodium is in Lepidoptera a small
tubercle above the pad [pulvillus] between the claws, bearing one bristle, seldom
two. It is widely distributed among the pad-bearing species, but is seldom found
in Sphingidae (Sphingulus; Hopliocnema).”’

In the discussion of certain genera of their Sphingulini, Rothschild and
Jordan (1903: LXIX) said: “...two closely allied genera Kentrochrysalis and
Sphingulus. The former possesses a complete paronychium and a pulvillus; the
latter genus has only a pulvillus. Here the paronychium has disappeared before
the pulvillus. The intermediate stage between the two genera is not known; in
this the paronychium would have one lobe.” This latter description fits Lapara
exactly, as would undoubtedly have been discovered by Rothschild and Jordan
had they had sufficient material for the genus, especially L. halicarnie.

The structure of the third segment of the labial palpus of Lapara species
indicates their closer relationship to the species of the tribe Sphingulini than to
the species of the tribe Sphingini. Structure of the male genitalia also shows
Lapara to have closer affinity with Kentrochrysalis and Sphingulus than with the
Sphingini (particularly the genus Sphinx). Early authors (Butler, 1877; Martyn,
1797; J. E. Smith in Abbot and Smith, 1797; Strecker, 1880) placed Lapara
species in Sphinx. But the clear distinction between the two genera is reinforced
by the lack of a pulvillus in the latter. The larvae of Lapara are referred to by
Koebele (1881) as ““Smerinthus-like”, an expression that Rothschild and Jordan
(1903: 154) also used when they established the tribe Sphingulini. The pupae of
Lapara described to date also correspond to the known pupae of the Sphingulini.
However, the apical segment of the antenna in Lapara is similar to that in all
genera belonging to the tribe Sphingini, where it was placed by Rothschild and
Jordan (1903). The importance of this structure was noted by Bell and Scott
(1937: 21), who remarked: “The end-segment is of particular taxcnomic value.
The length and shape, and the clothing with scales and bristles of this segment,
vary very much and offer good distinguishing characters of genera and even
tribes.”

In the light of available evidence Lapara may be considered to be an evolu-
tionary line intermediate between the Sphingini and Sphingulini, in many re-
spects even nearer to the latter than to the former. However, because of their
antennal structure, they should be left where they are, at the end of the Sphin-
gini.

PHYLOGENETIC RELATIONSHIPS OF SPECIES

Two evolutionary lines can be distinguished in Lapara from larval and adult
characters. One line is represented by L. halicarnie in which larvae have three
pairs of X-setae on the labrum, and adults have the pulvillus well-developed and
the paronychium with a single well-developed lateral lobe. Another distinct line
is represented by L. coniferarum and L. bombycoides, in which larvae possess
only two pairs of X-setae (the derivative condition) and adults have the pulvillus

6

poorly developed; in addition, the paronychium is short in coniferarum and
minute in bombycoides. Thus, L. halicarnie would seem to be the most gener-
alized and L. coniferarum and L. bombycoides the more specialized species of
Lapara .

BIOLOGY

Although the species of Lapara are not rare, except perhaps L. halicarnie, they
are nowhere abundant, and their natural history is poorly known. At northern
latitudes populations are univoltine, especially in Canada. Farther south, perhaps
south of New Jersey, a second generation, or at least a partial one, is likely.
Koebele (1881: 21) reported “at least 2 broods each year” for L. coniferarum in
the southern United States. The size of populations of L. bombycoides, the only
species for which data are available, appears to be stable with little fluctuation at
localities in Ontario where the species has been collected for several years.

Adults fly at night from about 10:00 p.m. to approximately 4:00 a.m. in
southern Ontario, but the pattern of activity probably varies in different parts of
the range. In southern Ontario, L. bombycoides deposits eggs in mid-July. In
L. bombycoides and L. coniferarum the eggs hatch in seven to ten days (Koebele,
loc.cit.), and larval development takes about six weeks in both species. Larvae of
all species normally feed on coniferous trees. An exception is a record of a
caterpillar of L. coniferarum taken from Citrus sp. in Dade City, Florida
(FSCA). To date no damage to economically important trees has been attributed
to members of this genus.

Koebele (loc.cit.) recorded a characteristic behaviour for the protectively
coloured larva of L. coniferarum. In the third and fourth instars when the head
is most triangular (Fig. 8), if disturbed, the caterpillar thrusts its head ventrally
so that it lies horizontally in a straight line with the body. Ordinarily the apex is
carried erect. Larvae of L. bombycoides reared in our laboratory exhibited the
same behavioural pattern when disturbed. The camouflage pattern of larvae of
these two species is illustrated in Fig. 21d.

The larva pupates in the earth in a loose web and remains in the pupal state
for a month (Koebele, loc.cit.) or more.

Key to Adults Based on External Features

. Apex of tegula with a prominent cluster of white hairs
usually extending to the wing bases (Figs. 19c, 20c);
basal area of fore wing heavily marked; single trans-
verse brown band on underside of both wings usually
well defined, particularly in males (Fig. 19d); length
of fore wing in males 21-26 mm (52 specimens; 28
mm in one); length of fore wing in females 25—27
mim-(SiX Specimens) 1) 5 Sia SOR OLE ee

Apex of tegula without white hairs (Figs. 19a—b,
20a—b); no heavy markings in basal area of fore wing

. Ground colour tawny; hind wing uniformly light tawny;
longitudinal markings on fore wing often weak or
absent (Figs. 19a, 20a); submarginal line indistinct,
only suggested; no transverse bands on underside of
wings; length of fore wing in males 29—36 mm (13
specimens); length of fore wing in females 32—40
AMM SHC SPECHMIENS reese s gehen on, iene ee ees

Ground colour brownish-ashy-grey; hind wing uni-
formly of this colour; postmedian longitudinal
markings on fore wing usually well-developed (Figs.
19b, 20b); a thin, dark submarginal line connecting
basal furcations of apical veins; transverse brown
bands on underside of wings absent or, if present,
only weakly developed; length of fore wing in males
24—30 mm (46 specimens), length of fore wing in
females 27—36 mm (eight specimens)...........

brags logs L. halicarnie

... L. coniferarum

Key to Adults Based on Genitalic Features

. Process of sacculus slender and somewhat sinuate in ven-

tralyie Ww: (Ries. Whe. A 2C\s os a: cna cee oe age are e
Process of sacculus short and curved, not sinuate, in
ve minaleviews(hiOw SC) cues: vee eeuapaee acer os co es aos eee

. Mesal margin of process of sacculus strongly sinuate in
ventral view, the base of the process averaging 0.37
mimawide: (hig aiklc) miei. amyl, Abate sonee. tes ave

Mesal margin of process of sacculus weakly sinuate in
ventral view, the base of the process narrower, aver-
acine-0:24 mmeawide (Pig s12 6\ier. Pens a ee

4. Anterolateral angles of vaginal plate extending into a
deep concavity on each side of the lip between the
seventh and eighth sternites, the surface of the con-
cavity somewhat sclerotized and pigmented (Fig.

GES: ROE ge ee ea a er rd ty te eer an L. bombycoides
Anterolateral angles of vaginal plate extending into the
lip between the seventh and eighth sternites, but
without the prominent concavity described above

GP igsnshoon dean! ¢ EE ake srs log were: geek trinsurishion ha quae ation es 5
5. Ovipositor valves small; vaginal plate fairly evenly
founded. throughout (Fig. 16a) ade: a:. ceatleraucny. act: be L. coniferarum
Ovipositor valves large; lateral margin of vaginal plate
Withea distinct ancle (Hie ajc he. Sa\cieas -f « tewce - eae L. halicarnie

Key to Mature Larvae

1. Body reddish-brown, with a regular, blackish, checkered
pattern (Figs. 21a—b); labrum with three pairs of

agGilional setae, X,—X5—_X5 (Pig. 7a)... . a1 .aaens wees: L. halicarnie
Body green, labrum with two pairs of additional setae,
WAP Sd Dae ied’ het warncer, alyaewicas he da qa isakas Bhs 2

2. Body light yellowish-green, with an unbroken cherry-
reddish median dorsal stripe between the dorsal lines
from thoracic segment II (occasionally III) to abdom-
inal segment eight (Fig. 21c); ventral surface broadly
suffused with a cherry-reddish coloration; head cap-
sule rounded dorsolaterally in frontal view (Fig. 6b)... L. coniferarum
Body pine-needle green, with either an unbroken red-
dish-brown median dorsal stripe from thoracic seg-
ment III to abdominal segment eight, with only
splashes of colour on the most posterior segments, or
with colouring absent (Fig. 21d); ventral coloration
condensed into a pinkish stripe, commencing at the
third pair of legs, widening as it proceeds caudally,
and embracing the prolegs; head capsule less rounded
dorsolaterally, tending to be angular in frontal view
CRs Ged berriserentt ities sid taxing bo exe care oe weet Lager srs L. bombycoides

9

Lapara halicarnie (Strecker)

Sphinx halicarnie Strecker, 1880, p. 35, fig.; Strecker (1872 ss.) 1900, p. 26.

Exedrium halicarnie: Grote, 1882, p. 11; J.B. Smith, 1888, p. 211.

Exedrium halicarniae: J. B. Smith, 1888, p. 240; Kirby, 1892, p. 696.

Lapara halicarniae: Rothschild & Jordan, 1903, p. 153; Wagner, 1913, p. 73;
B..P..Clark, 1919, p.102, pl. xi, iis. Lol Xin, me. 1: Draudt. 1931p. 660,
plo 5.a-

The orthography of the specific name as used in the few instances where the
species has been mentioned in the literature since 1888 is incorrect. J. B. Smith
(1888) introduced the misspelling in the list of the Sphingidae of temperate
North America at the end of his systematic monograph. As a Latin word “‘hali-
carnia’” does not exist. A genitive ending in “ae” cannot be formed. Thus,
another explanation of Strecker’s “halicarnie” is needed. A perusal of other
names Strecker applied to species reveals his preference for feminine proper
names. This is especially true in the paper containing the original description of
“halicarnie.”’ {n this original description Strecker said: “This Sphinx, of which I
append a figure in order to exhibit the great breadth of primaries, is without
doubt the most remarkable species yet discovered in this country.”’ No wonder,
then, that he chose the female proper name “‘Halicarnie’”’, probably derived (with
some linguistic acrobatics) from Halicarnasse, one of the seven wonders of the
ancient world.

For a considerable period of time, it was thought that Strecker’s type, a
hypertrophic female now in the collection of the Field Museum of Natural
History, was the only existing representative of the species. Although Rothschild
and Jordan (1903: 153) acknowledged the existence of a single specimen, their
description of coniferarum, as well as that by J. B. Smith (1888: 210) and others
for this species, suggests that they had before them a mixed series that also
included halicarnie (see p. 13).

Knowledge of the species halicarnie was greatly enhanced when Clark (1919),
an avid collector of world sphingids, recognized that the type specimen was an
abnormal female of that species. His material, six males and eight females, is
preserved in his collection, which has been deposited in the Carnegie Museum.
Now at least a few representatives of this species are in collections in many of
the larger North American museums.

Adult. Clark (1919) compared halicarnie (Figs. 19a 6, 20a 2) with coniferarum
and stated: “It [i.e. halicarnie| is a much larger insect. The markings of
the fore-wing are fainter, and often so vestigial that the wing is unicolourous,
lacking even the two longitudinal postcellular dashes. These differences become
very evident as one compares series of the two forms |[i.e., species]. The genitalia
are entirely different.”” To this description now may be added the following:
fringes of the wings exhibiting almost no evidence of a checkered pattern; under-
side of wings without transverse brown bands. Representatives of the species are
shown in Figs. 19a and 20a. Coloured reproductions of the hypertrophic female
type and that of a normal female can be found in a publication by B. P. Clark

10

(1919, pl. xIll, fig. 1; pl. XI, fig. 1). The figure given by Draudt (1931, pl. 95d)
is a replica of Clark’s original illustration.

The third segment of the male labial palpus (Fig. 3a) is short and broad, with
both sides convexly rounded; in the female it is long and slender, with almost
straight sides. As mentioned before, this kind of sexual dimorphism occurs only
in L. halicarnie.

The epiphysis (Fig. 4a) is massive, elongate, with a wide furrow, and nearly
closed at the tip.

Male genitalia (Fig. 1la—d): valve large and triangular, the apex attenuate.
Process of sacculus (Figs. 11b, 11c) slender and sigmoidal; a thorn-like projec-
tion near the middle of the process; apical third covered with small, thorn-like
spines; average width of process at its base 0.37 mm. Aedoeagus (Fig. 11d)
drawn out caudally into a bent, pointed process; vesica with two somewhat
cone-shaped projections; uncus-gnathos chelate in lateral aspect, not heavily
sclerotized.

Female genitalia (Fig. 15a—b): ovipositor valves large, rounded (Fig. 15b,
lateral view). Vaginal plate chevron-shaped, with lateral margins distinctly angu-
late. A narrow rectangular sclerotized plate situated at about the level of the
ostium bursae in the membranous band that goes around the ostial area from
end to end of the vaginal plate. Ratio of postapophyses: antapophyses, 1: 0.84.
Ductus bursae tube-like, turning dorsally at three-quarters of its length, with
thick walls imparting a doubled appearance. Ductus seminalis arising at a point
approximately one-quarter of length from distal end. Corpus bursae globular or
oblong, surface heavily marked with grooves.

Larva (Figs. 6a, 7a, 21a—b). A brown, dorsally checkered caterpillar was de-
scribed and figured by J. E. Smith in Abbot and Smith (1797, plate XLII) under
the specific name coniferarum. But the identity of the specimen illustrated and
its relationship to the adult figured have been doubted. Boisduval (1874: 105)
noted the dissimilarity between the specimen of this figure and other illustra-
tions of presumed coniferarum that he received from Abbot, the delineator of
the illustrations in Abbot and Smith (1797). Because he questioned the validity
of the caterpillar described in Abbot and Smith, Boisduval described another
caterpillar as coniferarum, based on Abbot’s additional drawings.

Further mention is made of the checkered caterpillar (sensu J. E. Smith in
Abbot and Smith) by Strecker (1874: 93), Fernald (1886: 85), J. B. Smith
(1888: 211), and Packard (1890: 768), the last three in reference to reports by
Koebele (1881) of a parasitized caterpillar taken in Tallahassee, Florida. Beu-
tenmuller (1895) and others subsequently demonstrated that the immature stage
described by Boisduval was indeed that of coniferarum. However, the identity of
J. E. Smith’s caterpillar remained to be determined. Forbes (1948: 192) stated
that the colour pattern, “dull brown and checkered with unusual regularity”,
was diagnostic of the species halicarnie, and his description agrees with the figure
published by Abbot and Smith (1797). Thus, the source of confusion in the
earlier literature was the incorrect association of the halicarnie caterpillar with
an adult coniferarum by J. E. Smith.

The figure of the caterpillar published in the work of Abbot and Smith
(1797) as the larva of coniferarum agrees with characters of two inflated cater-

vy

pillars without locality data in the collection of the Carnegie Museum. The
description of these caterpillars follows: head moderately triangular (Fig. 6a);
frons reddish-brown, bordered by a blackish marking having the form of a leaf of
the waterplant Sagittaria; anterior ends of marking extending about 5/6 of dis-
tance from vertex to lower edge of head capsule. This marking is bordered by
yellowish stripes with a reddish-brown tinge; these stripes again bordered later-
ally by blackish stripes terminating shortly above the eyes; rest of head capsule
reddish-brown, heavily sprinkled with blackish and yellowish granules; labrum
(Fig. 7a) with setae L, —L;, M,—M3, and X, —X3.

Body reddish-brown “checkered with unusual regularity” (Forbes, 1948:
192) (Figs. 2la—b); entire body sprinkled with conspicuous small brownish
warts on yellowish dots. Pronotum patterned differently from rest of body; as
on the head capsule, heavily granulated with blackish tubercles; marked with a
narrow, yellowish mid-dorsal line and with a pair of sub-median and a pair of
dorso-lateral blackish lines. Remainder of body segments with a complex
marking composed of a continuous, narrow, yellowish mid-dorsal line, flanked
on either side by several yellowish sub-median and dorso-lateral longitudinal
bands of various widths; these bands regularly interrupted by blackish patches of
various intensities, which are staggered in alternate bands creating the unique
checker-board effect. In the inflated specimen, illustrated in Fig. 21a, some of
the integument has stretched and colours have faded so that the over-all pattern
has been partially obliterated. However, if Fig. 21a is compared with a copy
(Fig. 21b) of the illustration in Abbot and Smith (1797), the elements of the
pattern can be distinguished. Anal segment with only remnants of the different
lines and patches; without caudal horn.

Pupa. Pupae of this species could not be obtained for study, and this stage has
not been described in the literature.

Type material. The type of L. halicarnie, a female in good condition, is in
Strecker’s collection in the Field Museum of Natural History, Chicago. The
specimen was taken in Florida in the summer of 1878 by the Reverend George
D. Hulst.

Distribution. L. halicarnie is restricted to the south-eastern portion of the United
States, with records from Florida, Georgia, South Carolina, and North Carolina.

Material Examined. Specimens examined were from the following localities and
collections: UNITED STATES. America Borealis: no other data (CM). Florida:
no other data (FMNH—type, AMNH, CM, FMNH, ROM); central Florida (CM);
Archbold Biological Station (DCF, JGF, YPM); Cassadaga (FSCA); Charlotte
Harbor (AMNH, CM); Coral Gables (WES); De Land (FMNH); Enterprise (CM);
Florida City (AMNH); Ft. Meade (USNM not seen); Gainesville (FSCA); Miami
(AMNH, CM, WES); Port Sewell (AMNH); St. Petersburg (USNM not seen).
Georgia: Nashville (FMNH); Waycross (FMNH). North Carolina: Southern Pines
(CM). South Carolina: Charleston (USNM not seen).

12

The material consisted of 25 males and 23 females of which five males and
six females were dissected. There are probably two generations, with the first
flight period from early January to late April and the second from mid-June to
mid-August.

Lapara coniferarum (J. E. Smith)

Sphinx coniferarum: J. E. Smith in Abbot and Smith, 1797, p. 83, t. 42; Harris,
1839-09297, a. 4lO- Clemens, 859, p. 174; Strecker, 1874, p. 93, idem,
1876p. 1 6, pl. X11, fig. 1.5)(d).

Hyloicus coniferarum: Hubner, 1822, p. 139.

Anceryx coniferarum: Walker, 1856, p. 224.

Ellema coniferarum: Grote, 1873, p. 27; Butler, 1877, p. 626; Packard, 1890, p.
768; Fernald, 1886, p. 84; J. B. Smith, 1888, p. 210.

Lapara coniferarum: Grote, 1875, p. 228, n. 72; Beutenmuller, 1895, p. 310, pl.
Vil, fie.-1; Rothschild and Jordan, 1903, p.. 150; Holland, 1903, p. 53, pl.
lly fe l6(2); Waener, 1913, p11; Draudt, 11931, op..860, pl. 95c,F orbes,
1948, p. 192.

Sphinx cana Martyn, 1797, pl. XIX, fig. 46—synonymy established Butler, 1877,
p. 626 (Ellema).

The descriptions of L. coniferar'um by Rothschild and Jordan (1903), Fernald
(1886), J. B. Smith (1888), and others were not based only on this species but
apparently were derived from mixed series of L. halicarnie and coniferarum.
Certain descriptive passages raise doubts as to the homogeneity of the series
studied by these authors. For example: “Variable in the distinctness of the
markings of the forewing and in size. The transverse lines disappear sometimes,
the discal streaks become faint in many specimens and the second is often
absent” (Rothschild and Jordan, 1903: 151). A re-evaluation of the material,
based on the morphological and anatomical features used here, indicates that
intra-specific variation is not as great as was previously assumed. Moreover, the
markings of the male often have a tendency towards greater expression, whereas
in females there is a tendency towards less accentuated markings.

Adult. Ground colour of adults (Figs. 19b 6, 20b 2) brownish-ashy-grey, the
postmedian dashes usually well-developed. Veins of anterior wing only slightly
suffused with brownish-grey and terminating apically as inconspicuous dots;
fringes of wings thus appearing only slightly checkered. Transverse brown bands
on underside of wings absent, or occasionally weakly developed. Tegula without
a cluster of white hairs at apex. Good colour reproductions are found in the
works of Strecker (1876, pl. XIII, fig. 15) and Draudt (1931, pl. 95c).

The third segment of the labial palpus (Fig. 3b) has almost straight sides and
narrows considerably towards the articulation with the second segment.

The epiphysis (Fig. 4b) is slender and elongate, with only a narrow furrow.

Male genitalia (Figs. 12a—d): valve (Fig. 12a) small and typically ovate,
occasionally somewhat triangular. Process of sacculus (Fig. 12b, c) strongly
sinuate in ventral view; usually a small thorn-like projection near the base of the

13

process, sometimes more strongly developed (Fig. 12, inset); apical third with
small thorn-like teeth laterally, rarely large (see Fig. 12, inset). The figure of the
process of the sacculus by Rothschild and Jordan (1903, pl. XXXIXx, fig. 12)
illustrates the characteristics of a normal specimen of L. coniferarum. Average
width of process of sacculus at its base is 0.24 mm. Aedoeagus (Fig. 12d) similar
to that of L. halicarnie; some filaments midway along length; cornuti absent;
coecum oval. Uncus—gnathos as in L. halicarnie but not as heavily sclerotized and
slightly more rounded.

Female genitalia (Figs. 16a, b): ovipositor valves small and somewhat pointed
(Fig. 16b, lateral view). Sterigmal area heavily sclerotized but not so heavy as in
L. halicarnie. Vaginal plate fairly evenly rounded throughout. As in L. halicarnie,
a rectangular, lightly sclerotized plate occurs opposite the ostium bursae in the
membranous band joining the ends of the vaginal plate around the ostium
bursae. Ratio of postapophysis:antapophysis, 1:1.05. Ductus bursae tubular, the
walls moderately thick. Ductus seminalis arising at end of posterior third of the
ductus bursae. Corpus bursae oblong, marked with shallow grooves.

Larva (Figs. 6b, 7b, 8, 21c). Boisduval (1875: 105) who noted the discrepancies
between the figure published by Abbot and Smith (1797, t. 42) and those of
other illustrations prepared by Abbot for Boisduval as the larva of L. conifera-
rum, subsequently described a non-checkered caterpillar as that of coniferarum,
an association that has since been verified by many authors (see discussion under
Larva of L. halicarnie). As was discussed previously, the checkered caterpillar
pictured by Abbot and Smith (1797) and mentioned by Koebele (1881) and
others, is that of L. halicarnie, not of coniferarum. In addition, Forbes (1948:
192) stated: “Larva of coniferarum tending to have more red [than bomby-
coides|, often with a continuous red dorsal area and heavy shading both sides of
the white substigmal stripes.”

Koebele (1881), who reared L. coniferarum, described the immature stages as
follows: “. . .light yellowish-green in color with three white lines on each side—
one just below the dorsal line—a second stigmatal—and the third half way be-
tween these. The back stigmatal spaces and the under part of the body are
strongly marked with red. The body is cylindrical, hardly varying in size from
one end to the other. There is no caudal horn through all its history. The head is
of medium size, light yellowish-green, edged along the collar with a blue line.
There is a black line running from each corner of the mouth to the summit of
the head and there they meet one another. The head is rounded, somewhat
conical, flattened in front. The length of the full grown larva is 2-3/4 to 3 inches
[10.8—11.8 cm] . —The egg is very dark green and hatches in 8 days... .The
pink color occurs in the larva only after the third moult. —The most re-
markable part of the history of the insect is the extraordinary change which
takes place in the shape of the head of the larva at different periods of its
growth. Immediately after birth it is round. With the first moult it becomes
angular and Smerinthus-like. This is very much increased with the second and
third moults so that in these it is fully 4 or 5 times its width running up to a
sharp point at the summit. When disturbed at this age the larva thrusts down the
extremity of its head so it lies straight in a line with the body. Ordinarily it
carries the point erect.’ The head of the immature caterpillar is shown in Fig. 8.

14

The following additions to the description of the caterpillar of L. coniferarum
are based on four inflated caterpillars and four preserved in alcohol. The inflated
specimens, without locality data, are in the collections of the American Museum
of Natural History and the Carnegie Museum. Those preserved in alcohol are
from the collection of the Florida State Collection of Arthropods—two from
Perry, one from Dade City, and one from Sanford, Florida; the foodplants were
given as Pinus longifolia, Pinus taeda and Citrus sp., respectively. Abbot and
Smith (1797) also mentioned cypress as a foodplant.

Head (Fig. 6b) of the mature larva slightly triangular; frons cherry-reddish,
bordered by black stripes fading out anteriorly; light yellow stripes between the
latter and the yellowish-green (body colour) remaining areas of the head capsule;
frontal plate somewhat shorter and wider than in bombycoides; labrum with
setae L,—L,, M,—M, and only X, and X, (Fig. 7b) forming a concave line;
pronotum and head capsule bearing yellowish granules; on thoracic segment II
(sometimes also III) a cherry-reddish spot in place of the spiracle; spiracles on
the abdominal segments have anteriorly and posteriorly a cherry-reddish smear
increasing in size caudally, filling the entire segmental space above the creamy-
white unbroken infraspiracular line in abdominal segment 8; the paired dorsal
and lateral stripes creamy-yellowish in colour; thoracic segment II (or III),
abdominal segments 1—8, and anal segment with a cherry-reddish patch between
the dorsal stripes; anal plate covered with small spinules, unicolourous, greenish-
or cherry-reddish, bordered by yellow; cherry-reddish splashes ventral to the
infraspiracular line beginning on thoracic segment II (or posteriorly) to abdomi-
nal segment 8; abdomen broadly suffused ventrally with cherry-reddish col-
ouring that also covers the semicoronate prolegs and the anal segment with the
anal legs; the entire body of the larva sprinkled with yellowish dots, more
distinct on the ventral surface, and with small spinules, especially dorsally. No
caudal horn.

Pupa. Pupae were not available for this study. Beutenmiiller (1895), however,
described the pupa as: “Cylindrical, pitchy black. Head, thorax, and anterior
margin of wing-covers rugosely punctate as are also the fore margins of the
segments. The four posterior segments are rugosely punctate nearly over their
entire surface. Anal spine pointed. Tongue-case concealed. Length, 32 mm.” The
applicability of Mosher’s (1918) description of the pupa of this species is doubt-
ful, because she used purchased material whose authenticity she suspected
herself.

Type material. It has not been possible to locate type specimens of either L.
coniferarum or L. cana. Clemens (1859: 174) based his description on “Abbot &
Smith’s figure”. Butler (1877: 626) did not mention any type of coniferarum or
cana in the British Museum (Natural History), although he treated all types kept
in that institution. A. H. Hayes, British Museum (Natural History), stated (in
litt.) that no specimens of Lapara that could have served as the basis for illustra-
tions of coniferarum or cana could be located in the British Museum collection.

The type material is therefore considered to be lost; only the illustrations are
available.

|e)

It seemed, however, to be neither necessary nor opportune to assign a neo-
type in this case.

Distribution. L. coniferarum is sympatric with L. bombycoides over much of its
range and is also sympatric with L. halicarnie wherever the latter occurs. The
distribution of this species includes the following states and provinces: Florida,
Alabama, Mississippi, Tennessee, Georgia, South Carolina, North Carolina,
Virginia, Maryland, New Jersey, Pennsylvania, New York, Rhode Island, and
possibly Nova Scotia.

In the Carnegie Museum collection are two specimens of L. coniferarum with
the locality given as “Nova Scotia’’. If valid, these are the northernmost and also
the only known records for this species from Canada. D. C. Ferguson (now of
the United States Department of Agriculture, Washington, D.C.), who worked
for many years in Nova Scotia, remarked (in litt.): ““Coniferarum seems to be
associated exclusively with the so-called hard pines, species such as P. rigida and
virginia, and not with White Pine. It is possible that the Nova Scotian coni-
ferarum that you spoke of were taken in an area of Jack Pine, which is closely
related to Pinus rigida. Jack Pine is limited mostly to Cumberland and Colchester
Counties in Nova Scotia, and I never managed to collect any Lapara in these
areas.” The specimens in question belong to B. P. Clark’s collection and were
obtained by him from a dealer in Brooklyn, New York.

Material Examined. Specimens examined were from the following localities and
collections: CANADA. Nova Scotia: no other data (CM). UNITED STATES.
Alabama: Cottondale (ROM); Loxley (AMNH); Shoal Creek Ford, 10 km NNW
Edwardsville (MSU); Wilmer (WES). Connecticut: no other data (CM). Florida:
no other data (AMNH, CM, FMNH, MSU); central Florida (CM); Alachua Co.
(FSCA); Altamonte (MCZ); Archbold Biological Station (DCF, YPM); Cassadaga
(FSCA); Coral Gables (CM); Dade City (larva FSCA); Fernandina (AMNH);
Florida City (JGF); Ft. Walton (FMNH); Gainesville (AMNH, FSCA); Hilliard
(AMNH); Levy Co. (FSCA); Ocean City (FSCA); Orlando (WES); Perry (larva
FSCA); Quincy (AMNH); St. Petersburg (AMNH); Sanford (larva FSCA);
Shalimar (FSCA); Tallahassee (AMNH, FSCA); Tampa (CM); Titusville (CM).
Georgia: Baldwin (ROM); Emory Univ. (FSCA); Thomasville (CM). Maryland:
Baltimore (CM, FMNH, ROM); Chevy Chase (MCZ). Massachusetts: Weston
(MCZ). Mississippi: Biloxi (AMNH, YPM, ROM, WES); Brooklyn (FSCA); Camp
Shelby (AMNH); Clear Spring Recreation Area nr. Meadville (MSU); Carnes
(WES); Clinton (ROM); Hattiesburg (AMNH, ROM); Jackson (AMNH, ROM,
WES); nr. Meadville (MSU); Pearl (ROM); Tape Co. (ROM). New Jersey:
Clementon (CM); Edgewater Hts. (AMNH); Lakehurst (AMNH, CM, CNC, DCF,
MCZ, ROM, WES); Manahawkin (AMNH, ROM); Ocean Co. (AMNH); Wrangle
Brook Rd. (AMNH, DCF, ROM). New York: no other data (CM); Bedford
(ROM); Coram, L. I. (CM); Hudson (CM); Leeds (CM); New Windsor (CM); New
York City and vicinity (AMNH); Sull Co. (CM); Yaphank, L.I. (CNC). North
Carolina: Charlotte (AMNH); Cherry Point (YPM); Fontana (CM); Sylva
(FMNH); Wilmington (YPM). Ohio: Hocking Co. (OSM). Pennsylvania: Lititz
(FMNH); Pittsburgh (WES); Williamsport (CNC). Rhode Island: Lonsdale

16

(FMNH); Rockland (ROM). South Carolina: Clemson (CLU); Coosawhatchie
(AMNH); Florence (CLU); McClellanville (ROM); Myrtle Beach (WES); Ridge-
land (ROM). Tennessee: Great Smoky Mt. Nat. Park, Gatlinburg (MSU—the only
melanic specimen); Johnson City (AMNH). Virginia: Amherst (CM); Arlington
(JGF); Clarksville (WES).

The material consisted of 192 males and 49 females, of which 41 males and
13 females were dissected. In the southern part of the range (Florida, Mississippi,
Carolinas, Georgia), there are two distinct flight periods, one from late March to
early May and another from early June to mid-September; in the central area of
the range (New Jersey, Maryland, Massachusetts) there is a single period, from
the beginning of May to mid-August (probably representing two overlapping
generations); and in the northern part of the range (New York, Connecticut,
Pennsylvania) a single flight period occurs from mid-June to September, probab-
ly representing one primary and a partial second generation.

Lapara bombycoides Walker

Papara bombycoides: Walker, 1856, p. 233; Clemens, 1859, p. 187; Butler,
1377, p. 626; Strecker, 1877, p. 127, pl. XIV, fis. 7 @ipara!); Rothschild &
Jordan, 1903, p, 152, pl. xXXVMI,1ig. 21, pl. XX XIX, fie 13; Holland, 1903,
p35) pl ie aig. J. Waener, 1913.>p. 72, Draudit, 1931. p:- 8600. pl. 95d°
Forbes, 1948, p. 192.

Ellema bombycoides: Fernald, 1886, p. 82; J.B. Smith, 1888, p. 208.

Ellema harrisii: Clemens, 1859, p. 188; Walker, 1864, p. 37 (Eilema!); Bethune,
isoo.p lo: Lintner. lo72.p. 170, pl. 8, ties. &, 9, 10, 1; Boisduval, 1875,
p. 106 (Sphinx); Strecker, 1876, p. 116, pl. XIII, fig. 16 (Sphinx); Butler,
1877, p. 626; ibid. p. 642 (Hyloicus); Packard, 1890, p. 768, figs. 264 a, b,
265 a, b, pl. XI, fig. 5, pl. XXXIV, fig. 1 a—j (larval details); Fernald, 1886, p.
83 (bombycoides var.); J.B. Smith, 1888, p. 207; Beutenmuller, 1895, p. 310,
pl. vu, fig. 2 (Lapara bombycoides var.) —synonymy established Grote,
1875, p. 228, no. 73, and others, latest Rothschild & Jordan, 1903, p. 152.

Adult (Fig. 19c, d 6, 20c ?). Most with distinct brown ante- and postmedian
transverse bands and postcellular dashes, giving a warm-brown, somewhat
checkered appearance; the light, serrate, postmedian transverse band occluded
apically by a brown arcuate band; a brown patch on the inner margin of the fore
wing; the veins of the fore wing suffused with brown, terminating at the wing
margin in distinct brown dots; fringes strongly checkered.

Geographic variation in colour is pronounced in this species. In the New
Jersey population and in the more southern populations, the predominantly
brown colour is muted by a lead-grey tone. Throughout the entire range, except
for southern areas, specimens are found that individually tend to be suffused
with brown, often to the extent of obliterating almost all markings. In the
central portion of the range, specimens occur which exhibit more accentuated
markings and a somewhat lighter shade of brown. These latter were described by
Clemens (1859) as a distinct species (sui iuris), under the name of Ellema
harrisii, with the type locality given as Brunswick, Maine. Grote (1875) synony-

If

mized harrisii without further explanation. Rothschild and Jordan (1903: 152),
who compared the type of bombycoides in Oxford with specimens of harrisii,
found no difference between the taxa and synonymized them. The concordance
of the genitalic structures, the shape of the head and the structure of the epiphy-
sis of the legs, as well as similarities of the caterpillars (see Larva), are further
evidence in support of the argument that arrisii is a synonym of bombycoides.

Despite the variation in coloration, bombycoides can usually be distinguished
from coniferarum and halicarnie by the presence of prominent transverse bands
on the underside of the wings (Fig. 19d) and by the unique cluster of white hairs
at the apex of the mesothoracic tegulae, which are particularly conspicuous in
infra-red photographs.

The third segment of the labial palpus (Fig. 3c) is distinctly convex on one
side, on the other more nearly straight, and not narrowed basally.

The epiphysis (Fig. 4c) is characterized by one uniformly convex side, with
the other almost entirely straight; the tip is not as pointed as in the other
species, the furrow more extended.

The anterior pretarsus of the male is unique for the genus in having an
empodium of two tubercles (Fig. 5b), each bearing a seta; tubercles much
reduced and difficult to locate, setae very fine.

Male genitalia (Figs. 13a—d): valve small and oval (Fig. 13a). Process of
sacculus (Figs. 13b, c) short and angled dorsally and mesally, not sinuate,
consisting of two angulate rods fused near the apex, which is adorned with small
teeth. Small, thorn-like process (frequently unilateral), often found at elbow
formed by junction of arm and terminal process of sacculus. The figure pub-
lished by Rothschild and Jordan (1903, pl. XX XIX, fig. 13) also illustrates the
fusion of the rod-like structures in marked contrast with those of the other two
species. Average width of process of sacculus at its base, 0.315 mm.

Aedoeagus (Fig. 13d) similar to that of L. halicarnie and L. coniferarum but
with slight differences in the vesica (cf. Figs. 11d, 12d); coecum slightly shorter
and oblong to oval in shape; uncus-gnathos similar to that of L. halicarnie but
more massive and slightly shorter when viewed laterally.

Female genitalia (Figs. 17a—b): structures remarkably different from those of
L. halicarnie and L. coniferarum. Valves of ovipositor large and rounded (Fig.
17b, lateral view). Vaginal plate shaped like an arrowhead; lateral margins
strongly rounded, anterolateral angles extending into a deep concavity on either
side of the lip between the seventh and eighth sternites; surface of this concavity
somewhat sclerotized and pigmented. Ratio of postapophysis: antapophysis,
1:0.77. Ductus bursae with posterior half swollen into a goblet-shaped structure,
anterior half tubular, walls thick. Ductus seminalis arising near middle of ductus
bursae, at base of posterior swelling; corpus bursae partially invaginated at end
of ductus bursae.

Larva (Figs. 6c, 7c, 21d). Forbes (1948: 192) described the caterpillar of
bombycoides: “Larva mostly green and white, at most with a red face and small
red dorsal spots.” This account is similar to the description presented for harrisii
(Clemens), by Boisduval (1874: 106). His description included the following
statements: “La chenille, comme celle du coniferarum, est dépourvue de corne

18

sur le onziéme segment. Elle est verte, un peu amincie aux extrémités avec un
raie dorsale un peu interrompue en arriére, d’un brun-rougeatre, et avec une raie
latérale jaune et une bande sous-stigmale blanche; outre cela, ses pattes écail-
leuses sont roses ainsi qu’une raie ventrale.” Lintner (1864), who was most
familiar with Lapara caterpillars, described the caterpillar of “harrisii’’ as
follows: “2 in. long [i.e. 7.8 cm], .23 in. broad [i.e. 0.9 cm]. Subcylindrical,
tapering slightly anteriorly, and the last two segments quite tapering. Head, size
of first segment, granulated, flattened anteriorly, sub-triangular, with an im-
pressed medial line, and straight yellow lateral lines terminating at the apex in
two black granulations, and bordered interiorly above with black. Body grass-
green. Subdorsal and lateral bands yellow. Substigmatal stripe bordering the
stigmata, white, enlarged on the central portions of the segments. Between the
subdorsal and substigmatal stripes,—on the fourth and fifth segments ventrally ,—
and exteriorly to the legs and prolegs, dotted on the annulations with paler green
or yellow. On the vascular line, a series of crimson spots on the anterior of the
segment, commencing usually on the fourth,—the first small, sometimes double—
the anterior ones triangular or lozenge-shaped, regularly increasing in size and
extending over more of the segment—the posterior ones quadrangular, and
uniting on the last two segments in a stripe. A ventral stripe of rose-colour,
commencing at the third pair of legs, widening as it proceeds, and embracing the
prolegs. No caudal horn. Caudal shield granulated, and edged with white.
Stigmata oval.” (This description is re-printed here to give additional evidence
from the caterpillar that “‘harrisii’’ is correctly a synonym of bombycoides.)
Detailed illustrations of the caterpillar of bombycoides are found in the work of
Packard (1890), p. 769, pl. XI, fig. 5, and pl. XXXIV (mistakenly listed under
coniferarum on text p. 768).

The following description of the immature stages of bombycoides is based on
living material reared from eggs of two females taken at Kendal, Ontario, 11 July
1967 and on two inflated caterpillars and one preserved in alcohol. The inflated
caterpillars (from the collection of AMNH) are without locality data. The one
preserved in alcohol, from Perry, Florida, is in the Florida State Collection of
Arthropods. Also used was a Kodachrome transparency taken by A. B. Klots
(AMNH) of a mature bombycoides caterpillar from Putnam, Connecticut.

The egg is elliptical, of a greenish-yellow pastel colour with the surface finely
chiselled. About 9 days are required for it to hatch.

Caterpillar in last instar about 50 mm long; body pine-needle green, well-
camouflaged by the presence of equally spaced creamy-white, unbroken dorsal
and lateral stripes that are comparable to the white stripes of the pine needles.
Head (Fig. 6c) of the mature caterpillar more triangular and pointed than in
coniferarum, frons light purplish-pinkish, bordered by dark brownish-red stripes
that fade out anteriorly; these bordered by bright yellow stripes; rest of head
capsule of the same green colour as the body; frontal plate more elongate and
narrower than in coniferarum; labrum with setae L;—L3, M,—M3 and only X,
and X, (Fig. 7c); X-setae forming a convex line; head capsule more granulate
than in coniferarum; pronotum as the head capsule, densely covered with fine
yellow granulations. Spiracles on abdominal segments surrounded by a slight
tinge of purplish-pink; infraspiracular line pure white (unbroken on thoracic

19

segment I to abdominal segment 1, interrupted at intersegmental boundaries,
rounded on abdominal segments 2 to 8, and absent on anal segment), wider than
other longitudinal lines. On the anal segment occur only lateral lines. In the
reared caterpillars (Fig. 21d) the space between the dorsal lines is completely
filled with purplish reddish-brown from thoracic segment III to the anal seg-
ment, whereas thoracic segment II shows only a little colouring. The specimen
photographed by A.B. Klots has some colouring but only on abdominal seg-
ments 6 to 8 and the anal segment. The inflated caterpillars have an intermediate
colouring. The abdomen ventrally with not more than a purplish-pink, relatively
broad median line, with lateral connections to the semicoronate prolegs and anal
legs; the entire body of the caterpillar sprinkled with yellowish dots and with
minute spinules, but not as heavily as in coniferarum and spinules smaller. The
first signs of dorsal colouring appear in some caterpillars by the fourth instar.

Known foodplants (see McGugan, 1958) are Pinus banksiana, P. resinosa, P.
rigida, P. strobus (in the eastern provinces and states and in southern Ontario),
and Larix laricina. P. taeda is the foodplant given for the specimen from Perry,
Florida.

Pupa (Fig. 9, pupa of Fig. 21d). Beutenmuller (1895) wrote: “Chestnut brown,
with a rough, not produced head-case. Tongue-case buried. Posterior segment
tapering. Terminal spine black, contracted at base, minutely bifid at tip. Length,
...[25—27 mm].” This description agrees with a pupa (Fig. 9) from the ROM
collection reared from the eggs of females taken at Kendal, Ontario, in 1967. It
should be noted that the bifurcation of the terminal spine is so fine that it
appears as a “tuning fork”’ only in the very recently formed pupa. Usually only
the two basal stumps remain in older pupae. Lintner (1864) described the pupa
in the same way as Beutenmuller.

Type material. According to Strecker (1877), Fernald (1886), and Rothschild
and Jordan (1903), the type of L. bombycoides is in the Museum at Oxford,
England, specifically in the Hope collection of Oxford University (Strecker,
1877). Strecker (1877) published “‘an accurate coloured figure of the type” that
was made for him by Prof. Westwood in 1875. It appears to be a female.

In his description Walker (1856) gave the type locality as ““Canada. In Mr.
Saunder’s collection.” As Strecker (1877) said, the type was then unique.

Where the type of harrisii is deposited is unknown. Clemens (1859) says after
his description: “From the collection of Mr. A. S. Packard Jr., Brunswick, Me.”
In his special study of Clemens’ types, Darlington (1947) said: “Lapara harrisii
(=bombycoides Walker), p. 188. Type not located. Described from a specimen in
A. S. Packard’s collection.”

In connection with these remarks about type material of bombycoides, the
question of the creation of a synonym by Harris must be resolved. In 1839
Harris published his catalogue of the North American Sphinx and listed on p.
297, under no. 10, ““S. Coniferarum. Smith-Abbot.” His description agrees well
with that of the insect originally described by J. E. Smith in Abbot and Smith in
1797. It would appear that Harris did not intend to describe any new and

20

hitherto unknown insect, for he placed the authors’ names after the specific
name and as well referred to the caterpillar pictured by Abbot and Smith.

On the other hand, it seems equally clear from the second paragraph of his
note (p. 297), that Harris’ own specimen, the specimen “in the Cabinet of
T.W.H.”, probably was not L. coniferarum, because this species, as far as is
known, does not occur in Vermont. The small size of the specimen is probably
not taxonomically important, because it was reared. Nevertheless, Harris men-
tioned that the specimen, taken in North Carolina and in the collection of the
Boston Society of Natural History, was larger. This specimen is undoubtedly
referable to L. coniferarum (J. E. Smith), for Harris’ last words in his note were:
“the bands on the wings in the latter are less distinct than in my specimen.”’ The
description clearly applies to this species.

Harris undoubtedly believed that the two specimens of Lapara represented a
single taxon, when actually they were representatives of two distinct species. His
own specimen was L. bombycoides, a new taxon that was described 16 years
later by Walker (1856). Thus, the distinctiveness of his specimen was not
detected by Harris and he consequently misidentified it (Bethune, 1868: 18).
Even so, a misidentification is not a cause for synonymizing a name, and the
synonymy of “Sphinx coniferar'um Harris (non Smith and Abbot, 1797)
(partim)” under bombycoides or harrisii, respectively, as was given by Clemens
(1859), Morris (1862), J. B. Smith (1888), Rothschild and Jordan (1903), and
Wagner (1913) is incorrect.

A final comment on the synonymy under L. bombycoides. In 1862 (p. 382)
Harris published on the caterpillar of S. coniferarum, and Wagner (1913, pars 12,
p. 72) included Harris’ short statement in his synonymy of bombycoides.
Wagner’s action was not only incorrect but also inappropriate, for Harris clearly
referred to the caterpillar pictured by Abbot and Smith (1797) on plate XLII.

Distribution. L. bombycoides, the most common of the three species of Lapara,
is widely distributed in Canada and the United States from the Atlantic Coast
south to Florida, and west to Saskatchewan, Nebraska, and British Columbia. In
the central and southern portions of its range, it is sympatric with L. halicarnie,
L. coniferarum, or both.

Records (including those from the literature) have been obtained for the
following states and provinces: Florida, Georgia, North Carolina, Tennessee,
Indiana, West Virginia, Maryland, Pennsylvania, New Jersey, New York, Connec-
ticut, Rhode Island, Massachusetts, Vermont, New Hampshire, Maine, Nova
Scotia, New Brunswick, Quebec, Ontario, Manitoba, Saskatchewan, British Col-
umbia, Nebraska, Minnesota, Wisconsin, Michigan.

Kimball (1965: 61) questioned the occurrence of L. bombycoides in Florida
and stated: “Certainly there is no clear-cut record of it here; in fact, the only
one is ‘Florida’ (Rothschild and Jordan, 1903: 152). . .”’. The specimens referred
to by Rothschild and Jordan are now housed in the collection of the British
Museum (Natural History) and were kindly sent on loan for this investigation.
Both are males, with one bearing the label “Florida” and the other “Palm Beach,
Florida.” Neither specimen differs in any aspect from Ontario specimens, and
the slight differences in the structure of the genitalia mentioned by Rothschild
and Jordan are within the normal range of individual variation for this species.

21

Moreover, there is a caterpillar of this species, taken 30 April 1959 at Perry,
Florida, in the Florida State Collection of Arthropods in Gainesville. L. bhomby-
coides obviously does occur in Florida, if only rarely.

Material Examined. Specimens examined were from the following localities and
collections:

CANADA. Canada: no other data (AMNH). Manitoba: Red Rock Lake,
Whiteshell Park (AMNH). New Brunswick: Bathurst (DCF); Caraquet (DCF);
Fredericton (CNC). Nova Scotia: no other data (CM); Armdale (CNC, DCF);
Aylesford (DCF); Baddeck (DCF); Brierly Brook (DCF); Lake Kejimkujik
(AMNH, DCF); Petite Riviére (DCF); S. Milford (CNC); White Point Beach
(AMNH, CNC). Ontario: no other data (CNC); Algonquin Prov. Park (ROM);
Ancaster (CNC); Bainsville (LEM); Bancroft (CNC); Bertram I., Lake Nipissing
(ROM); Biscotasing (CNC); Bracebridge (AMNH); Camden East (CNC); Camp
Borden (ROM); Chaffeys Locks (ROM); Chapleau (ROM); Constance Bay
(CNC); Dunnville (ROM); Franklin I., Georgian Bay (ROM); Geraldton (AMNH,
ROM); Grand Bend (CNC); Gull Lake, Muskoka (AMNH, CNC); Kendal (ROM);
Marmora (CNC); Morris I., Muskoka (ROM); Muskoka (ROM); Normandale
(CNC); One Sided Lake (AMNH, CNC, ROM); Ottawa (CNC); Port Credit
(ROM); Rouge Hill (CNC); Rouge River (CNC); Sudbury (AMNH, CNC, ROM,
WES); 1000 Islands (cM); Toronto (ROM); Trenton (CNC, ROM). Québec:
Alcove (CNC); Amos (CM); Forestville (CNC); Hudson (LEM); Kazabazua
(CNC); Kingsmere (CNC); Meach Lake (CNC); Montréal (CM, LEM ); Murray Bay
(CM, LEM); Norway Bay (CNC); Oka (cM); Ste. Thérése I. (cM), UNITED
STATES. Connecticut: Putnam (AMNH). Florida: no other data (BM); Palm
Beach (BM); Perry (larva FSCA). Georgia: Oglethorpe University Campus (not
seen, mentioned by Knudson, 1954). /ndiana: Dune State Park (FMNH); Miller
(FMNH); Mineral Springs (FMNH). Maine: no other data (CM); Augusta (FMNH,
WES); Bar Harbor (AMNH, FMNH); Biddeford (LEM, WES); Dixfield (AMNH);
Enfield (AMNH); Harrison (AMNH); Hope (AMNH, WES); Jackman (WES);
Kittery Point (MCZ); Lincoln (AMNH, DCF); Mattawamkeag (AMNH); Mt.
Desert (AMNH); Norway (AMNH); Old Orchard (CM); Oquossoc (WES); Paris
(CM); Passadumkeag (JGF); Tim Pond (WES). Massachusetts: no other data
(CM, FMNH); Amherst (MSU); Barnstable (AMNH, FSCA); Beverly Farms
(MCZ); Cohasset (AMNH); Dorchester (MCZ); Forest Hills (CM); Groton
(AMNH); Lenox (AMNH); Marlborough (YPM); Martha’s Vineyard (YPM);
Middlesex Co. (CNC); Neville (MCZ); Tyngsbro (MCZ); Weston (MCZ); Winchen-
don (CM); Wollaston (MCZ). Michigan: Allegan (MSU); Cheboygan Co. (WES);
Chippewa Co. (MSU); Colombianville (WES); East Lansing (MSU); Keweenan
Park (MSU); Otsego Co. (ROM); Schoolcraft Co. (AMNH, MSU, WES); Tecon
Lake (MSU). Minnesota: Itasca State Park (OSU). Nebraska: no other data (CM).
New Hampshire: Franconia (AMNH); Hampton (WES); Jefferson (CM); Jeffer-
son Notch (DCF); Portsmouth (yPM); Squaw Lake (MCZ); Walpole (cm). New
Jersey: Lakehurst (AMNH, CNC, WES); Manahawkin (AMNH, ROM); Morgan
(CM); Ocean Co. (AMNH, CM); Paul Smith’s (LEM); Sussex Co. (AMNH);
Wrangle Brook Rd. (AMNH). New York: no other data (AMNH, CM, FMNH,
MCZ); Northern N.Y. (CM); Adirondacks (CM, FMNH); Albany (AMNH, CM,
MSU); Albany Co. (CM); Ausable (LEM); Bear Mt. (AMNH); Bedford (AMNH);

22

Bluff I. (CM); Clayton (Picton I.) (AMNH); Cornwall (CM); Dryden, Ringwood
Reserve (JGF); Green Co. (FMNH); Hudson (CM); Ithaca (AMNH, CM, JGF);
Jamestown (CM, FMNH); Leeds (CM); Orange Co. (AMNH); Promised Land, L.I.
(CM); Saratoga (CM); Taughannock (Ithaca) (AMNH); Ulster Co. (CNC);
Valcour (LEM); Wading River, L.I. (CM). North Carolina: Brevard (FMNH);
Doughton Park (MSU); Fontana (CM); L. Junaluska (FSCA); Tryon (CM); Tsali
Recreation Area, 10 km. NE Stecoah (Msu). Pennsylvania: Clearfield (CM);
Shawville (CM). Rhode Island: Cranston (CM); N. Scituate (ROM). Tennessee:
Crossville (ROM); Great Smoky Mt. Nat. Park, Gatlinburg (MSU). Vermont:
Pomfret (CM). West Virginia: no other data (CM). Wisconsin: Argonne (WES);
Bailey’s Harbor (WES); Chippewa Falls (WES); Cumberland (CM); Hancock
(WES); Hazelhurst (AMNH); L. Katherine (WES); Lost Lake (Vilas Co.)
(AMNH); North Bay (WES); Sauk City (WES); Vilas Co., nr. Sayner (AMNH);
Waussa (AMNH).

The material consists of 605 males and 44 females, of which 18 males and 3
females were dissected. A single flight period occurs from early June to early
August. One record from September (Clearfield, Pennsylvania) suggests a partial
second generation.

Incertae sedis: Lapara pinea (Lintner)

Fliema pineunr Winther, 1872, p. 169, pl. 8, Tigs. 12, 13; ibid: p. 192; Butler,
1e7 7, p. O26; Packard, 1890) p. 770, fig. 2664, -b; J. B. Smith, 18838, p.-209.

Sphinx pinea: Boisduval, 1875, p. 107.

Lapara pineum ('): Grote, 1875, p. 228; Rothschild and Jordan, 1903, p. 151;
Holland, 1903, p. 53; Wagner, 1913, p. 71; Draudt, 1931, p: 860 (conijer-
arum ssp.); Forbes, 1948, p. 192 (bombycoides var.).

Lapara bombycoides pinetum ('): Dyar, 1902, p. 70.

The adjectival form used with Lapara, a noun of female gender, must be pinea;
Ellema, the genus to which it was originally assigned, is neuter.

The status of this insect has always been questioned. Rothschild and Jordan
(1903) considered L. pinea tentatively as a species but reserved judgement as to
its definitive rank by the provision that “the two specimens should be
re-examined.” Dyar (1902) considered it to be a subspecies of bombycoides, and
Draudt (1931), a subspecies of coniferarum. For this review the morphology of
the genitalia and other structures was examined.

Adult. The type specimens, a male and a female, were pictured by Lintner (1872)
in the original description beside a male and a female of “‘Fllema harrisii”’
(=bombycoides). Lintner (1872) stated that: “The species is readily distin-
guished from FE. harrisii (Plate 8, figs. 10, 6, 11, 2) by the darker ground of its
wings, the absence of the grey shades, and its much less distinct markings.” L.
pinea is known only from the type specimens, and no additional specimens
referable to this taxon were found among the material that I examined. Two
specimens in the Clark collection of sphingids in the Carnegie Museum, however,
are labelled “Ellema pineum.” One (Clark #5697, locality given only as New

Pde

York, collected by Bruce) bears the notation “fide J. B. Smith”. The other
(Clark #5564) is from Baltimore, Maryland, taken in June, the year and the
collector unknown.

The process of the sacculus of the male genitalia of both Clark specimens is
identical to that of L. coniferarum indicating that they are nothing other than
small specimens of this species. The base of the process in both is 0.22 mm wide.
The process is in one piece, and there are no fused rods. Also, the epiphysis is
that of a small L. coniferarum. The colour of both specimens is the ashy-greyish-
brown colour characteristic of L. coniferarum. The absence of distinct markings
on the wings is a result of the worn condition of the specimens. The shape of the
head and third joint of the labial palpi furthermore identify both as L.
coniferarum. With a wing measurement of 24 mm, Clark 5697 is within the usual
range for L. coniferarum, Clark 5564, with the wing only 23 mm long, is rather
small but agrees with L. coniferarum in all other characters.

The two Lintner specimens (Fig. 20d, 2? type) are among the smaller Lapara
specimens examined (d type measures 21 mm from right fore wing apex to base;
? 25 mm); fore wings not heavily scaled, umber coloured, a little greyish dusting
at the base and near the apex, along the terminal margin, and on the principal
veins and their branches; within the cell is a subquadrangular dark-brown heavy
spot (in the 2 4 x 1.25 mm) parallel to the costal margin, a feature which all
species of Lapara lack; a shade of the same dark-brown colour at the middle of
the costal margin and at the apex; distally, a lighter brown semicircular area from
the middle of the costal margin to the middle of the inner margin; hind wings of
the same umber colour but only about a third as dark as the fore wings, paler
still at the base; fringes distinctly checkered; colour beneath uniformly medium
umber and without lines as in L. coniferarum; tegulae of the same umber colour,
with some greyish hairs laterally and fine (arcuate) white lines on mesal margins
as in L. coniferarum; abdomen of the same general umber colouring. Epiphysis
very small, its shape intermediate between that of L. coniferarum and L.
bombycoides. Labrum as in L. coniferarum. Third joint of labial palpi inter-
mediate between that of L. coniferarum and L. bombycoides.

Valve of male genitalia (Fig. 14a) similar to L. bombycoides; process of
sacculus short, not sinuate, as in L. bombycoides, abruptly bent dorsally and
bearing thorns and hooks suggestive of L. halicarnie or L. coniferarum; structure
similar to that of L. bombycoides, with two short rod-like components fused
distally at the apex; width at base 0.18 mm; aedoeagus (Fig. 14b) similar to the
other three species of Lapara but thinner.

The female genitalia (Fig. 18) of L. pinea are in most respects similar to those
of L. bombycoides. Ovipositor valves small and somewhat pointed, as in L.
coniferarum; shape of the vaginal plate as in L. halicarnie; anterior lateral
margins somewhat angulate, more so on one side than on the other; the
anterolateral angles of the vaginal plate extending into a deep concavity on each
side of the lip as in L. bombycoides. Ductus bursae tube-like, posterior half
lightly sclerotized, turning anterolaterally to the right shortly before its
termination; bursa copulatrix connected dorsally and obliquely to the left;
ductus seminalis arising as a rather strong tube shortly before middle of ductus
bursae. Corpus bursae similar to L. bombycoides but not invaginated, connected
as in L. coniferarum; not heavily marked with grooves; signum absent.

24

Larva. The type specimens were reared from caterpillars by Lintner, who found
them in September of the years 1858 and 1859 in Schoharie, New York. Lintner
succinctly stated the principal difference between the pinea and “‘harrisii”
(=bombycoides) caterpillars. The former exhibits as “its characteristic feature” a
dorsal row of interrupted red squares, which, as we know now, are also found in
some bombycoides caterpillars. In 1860 Lintner observed that “in those taken
this year the dorsal squares are not visible”, later, however, he discovered that
the caterpillars taken in 1860 were those of “harrisii’’. He stated further that he
was unable to find additional caterpillars of L. pinea.

Lintner’s statement was somewhat misunderstood by Boisduval (1875), who
said: “La chenille a une si grande ressemblance avec la chenille de Harrisii, que
Mr. Lintner a confondu ensemble les premiéres qu’il a trouvées. Les deux espéces
vivent a méme époque sur le Pinus strobus, dans Etat de New-York.” This
statement gives the impression that Lintner had confused his first and only pinea
caterpillars with others and did not know from which batch he obtained pinea
and from which one ‘“‘harrisii”’. This was not so, however, as the two groups of
caterpillars were collected in different years and were recognized by Lintner as
separate entities.

Lintner (1872) described the caterpillar of L. pinea as follows: “Length two
inches [i.e. 7.9 cm]. Colour, grass-green. Head subtriangular, green, bordered
with bright yellow, within which at the apex is a /\ of black. Body
subcylindrical, tapering at the extremities, and without caudal horn. Dorsally, a
reddish-brown line interrupted on the hind-portion of each segment by a square
of green transversed by diagonal lines; a subdorsal stripe white, interrupted at
the sutures by light green; ventral stripe and prolegs, rose-red. Feeds on White
Pine, and matures about the middle of September, when it enters the ground and
forms a cell for pupation.”

In the Carnegie Museum, there is one inflated caterpillar from the Holland
collection, bearing only the label “Sphinx pineum Lintner’’, which is not distinct
from L. bombycoides caterpillars (on the basis of head capsule morphology).

Pupa. Unknown.

Distribution. Butler (1877) and J.B. Smith (1888) reported LZ. pinea from
Canada, but specimens are not available to verify their records.

Discussion

The two known specimens of this taxon are morphologically intermediate be-
tween L. coniferarum and L. bombycoides, but whether they represent a fourth
species of Lapara cannot readily be decided until more material is available for
study. Lintner’s two specimens are more than 110 years old, and he apparently
did not collect others at Schoharie. The country west of the Hudson River and
north of the Catskill Mts. was and is frequently visited by collectors of Lepidop-
tera, and none ever reported L. pinea. | know of no sphingids with such a limited
distribution, but even so, they would be expected to occur at least at some
localities in the vicinity of the type locality.

rs)

For the time being at least, Lintner’s two specimens cannot be assigned
rightly to any known species of Lapara, although they certainly belong to this
genus. As they show characteristics of both bombycoides and coniferarum, |
strongly recommend that crossbreeding experiments with these two species be
undertaken to determine if hybridization is possible. Perhaps Lintner’s speci-
mens are nothing more than hybrids and the taxon “pinea”’ is an invalid one.

Acknowledgments

The assistance of the following institutions and individuals in making material
available for this study is gratefully acknowledged: H.K. Clench, Carnegie
Museum; B. Colborne, Ohio State Museum; G. W. Dekle, Florida State Collec-
tion of Arthropods; H. E. Evans, Museum of Comparative Zoology; D. C. Fer-
guson, Yale Peabody Museum of Natural History; W. D. Field, United States
National Museum; R. L. Fischer, Michigan State University; J. G. Franclemont,
Cornell University; D. F. Hardwick, Canadian National Collection of Insects;
A. H. Hayes, British Museum (Natural History); J. R. Holman, Clemson Universi-
ty, South Carolina; F. W. Mead, Florida State Collection of Arthropods; F. H.
Rindge, The American Museum of Natural History; W. E. Sieker, Madison,
Wisconsin; C. A. Triplehorn, Ohio State University; V.R. Vickery, Lyman
Entomological Museum; R. L. Wenzel, Field Museum of Natural History; G. B.
Wiggins, Royal Ontario Museum; J. A. Wilcox, New York State Museum, types
of L. pinea; A. K. Wyatt, Field Museum of Natural History.

Special thanks are extended to F.H. Rindge, AMNH, who made at the
author’s request the first dissections of Lapara specimens and so aided in the
initiation of this project, and to B. Mather, Jackson, Mississippi, who collected
with undiminished enthusiasm over the years many of the Mississippi specimens
employed in this study. Thanks are also due to C. F. dos Passos for expert advice
concerning the etymology of the name L. /halicarnie.

Gratefully acknowledged is the help of A. B. Klots and F. H. Rindge, AMNH,
D. F. Hardwick, CNC, G. B. Wiggins, ROM, in critically reading this paper, of T.
Yamamoto, ROM, and D. Barr, ROM, in editing the manuscripts. All figures
were prepared by A. Odum, scientific illustrator in the Department of Ento-
mology and Invertebrate Zoology, ROM. The photographs were taken by L.
Warren, Head Photographer, Department of Photography, ROM. Miss A. Hillmer
typed the final manuscript.

26

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iA

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29

Figs. 1—5
Fig. 1

Fig. 2

Fig, 3

Fig. 4

Fig. 5

30

Adult structures.

Frontal view of denuded head (diagrammatic): a) L. halicarnie. b)
L. coniferarum;c)L. bombycoides.

Ventral view of denuded head (diagrammatic): a) L. halicarnie; b)
L. coniferarum;c)L. bombycoides.

Labial palpus, second and third segments: a) L. halicarnie (AMNH
11,353 6, Sewall, Florida, 6 February 1948; ROM 9-180 9,
Florida, no date); b) L. coniferarum (ROM 9—037 6, Pearl,
Mississippi, 11 June 1960); c) L. bombycoides (AMNH 11,373 6,
L. Kejimkujik, Nova Scotia, 19 June 1957).

Epiphysis of anterior leg: a) L. halicarnie (AMNH 11,353 6, Sewall,
Florida, 6 February 1948); b) L. coniferarum (ROM 9—039 a,
Jackson, Mississippi, 11 June 1960); c) L. bombycoides (AMNH
11,373 6, L. Kejimkujik, Nova Scotia, 19 June 1957).

Pretarsus of anterior leg: a) L. halicarnie (AMNH 11,353 6, Sewall,
Florida, 6 February 1948); b) L. bombycoides (ROM 9-159 6,
White Point Beach, Nova Scotia, 24 June 1953).

J]

Figs. 6—9
Fig. 6

Fig. 7
Fig. 8

Fig. 9

Larval and pupal structures.

Larval headcapsules. a) L. halicarnie (CM, inflated larva, no data);
b) L. coniferarum (FSCA, larva in alcohol, Perry, Florida, 30
September 1958, on Pinus taeda);c) L. bombycoides (FSCA, larva
in alcohol, Perry, Florida, 30 April 1959, on Pinus taeda).

Setal groups on larval labrum of the three specimens of Fig. 6.
L. coniferarum (FSCA, larva in alcohol, fourth instar, Sanford,
Florida, 10 March 1961, on Pinus longifolia).

L. bombycoides (ROM, cremaster of pupa in alcohol, ex ovo,
Kendal, Ontario, 10 September 1967).

a5

Figs. 10—14 Male genitalia.

Fig. 10

Pig.11

Fig. 12

Pig. ob

Fig. 14

34

Entire structure of L. bombycoides (ROM 9—157 6, Chaffeys
Locks, Ontario, 3 July 1964), ventral view.

L. halicarnie (AMNH 11,353 6, Sewall, Florida, 6 February 1948):
a) entire left valve, medial view; b) process of sacculus, lateral view;
c) process of sacculus, ventral view; d) aedoeagus.

L. coniferar'um (ROM 9—037 6, Pearl, Mississippi, 11 June 1960):
a) left valve, medial view; b) process of sacculus, lateral view; c)
process of sacculus, ventral view with two outline drawings of same
view illustrating variation; d) aedoeagus.

L. bombycoides (AMNH 11,373 6, L. Kejimkujik, Nova Scotia, 19
June 1957): a) left valve, medial view; b) process of sacculus,
lateral view; c) process of sacculus, ventral view; d) aedoeagus.

‘“T. pinea’’ (NYSM 6 type, no further data): a) left valve, medial
view; b) aedoeagus.

IS

ee
= eee

iucwtts (ROM F-15528
7
= Aes 1 ea ny

5b

Figs. 15—18 Female genitalia.
Fig. 15

Fig. 16
Fig. 17

Fig. 18

L. halicarnie (ROM 9—180 9, Florida, no date): a) ventral view; b)
Ovipositor valve, lateral view.

L. coniferarum (ROM 9—179 9, Baltimore, Maryland, 25 June): a)
ventral view; b) ovipositor valve, lateral view.

L. bombycoides (ROM 9—177 9, One Sided Lake, Ontario, 5 July
1962): a) ventral view; b) ovipositor valve, lateral view.

“TL. pinea” (NYSM @ type, no further data), ventral view.

I

19b

Fig. 19 Adult moths. a) L. halicarnie (AMNH 11,353 6, Sewall, Florida, 6
February 1948); b) L. coniferarum (AMNH 11,3674 , Lakehurst,
new Jersey, 21—31 May); c) L. bombycoides (AMNH 11,373 6, L.
Kejimkujik, Nova Scotia, 19 June 1957), dorsal view; d) same,
ventral view.

3S

Fig. 20 Adult moths. a) L. halicarnie (ROM 9—180 9, Florida, no date); 5)
L. coniferar'um (ROM 9—124Q, Florida, no date); c) L. bomby-
coides (ROM 9—210 9, Chaffeys Locks, Ontario, 13 July 1963);

d) ‘‘L. pinea’”’ (NYSM & type, no further data).

39

Fig. 21 Larvae. a) L. halicarnie (CM inflated larva, no data); Db) L.
halicarnie (copy of Abbot’s drawing (1797: pl. XLII)); c) L.
coniferarum (CM inflated larva, no data);d) LZ. bombycoides (ROM
in alcohol, live photograph by J.C.E. Riotte, Kendal, Ontario, 3
September 1967).

40