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89 « ISSN 0024-0966
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Journal of the
Lepidopterists Society
Published quarterly by The Lepidopterists’ Society
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VOLUME 60, NUMBER 3
Journal of the Lepidopterists’ Society
60(3), 2006, 121-137
FIVE NEW SPECIES OF PAUCIVENA DAVIS, 1975 (LEPIDOPTERA: TINEO{DEA:
PSYCHIDAE) FROM CUBA
RAYNER NUNEZ AGUILA
Division de Colecciones Zoolégicas y Sistematica, Instituto de Ecologia y Sistematica, Carretera deVarona km els ARIE
Capdevila, Boyeros, C. de La Habana, Cuba. AP 8029. CP 10800, Cuba.
E-mail: rayner@ecologia.cu
ABSTRACT. Five new species of Paucivena Davis Sau a Tineoidea: Psychidae), P. ferruginea, P. pinarensis, P. fusca, P. cubana and P.
orientalis are described from Cuba and compared with relatives. The females of P ferruginea and P. orientalis are described being the first fe-
males known within genus; their characteristics confirm the intermediate position of the genus among the American psychids. Notes on natural
history of new species (e.g. hosts, habitat) are given as available. Keys for identification of known stages of all Paucivena species are provided.
Additional key words: Tineoidea, Psychidae, bagworm, natural history, West Indies.
The Neotropical region has the richest Lepidoptera
diversity but one of the least studied psychid fauna with
only 61 known species (Heppner, 1998; Davis, 2000).
This species number is extremely low compared with
that of other faunal regions. The Paleartic, for example,
has more than 300 decenbed species (Heppner, 1998).
Members of this family were last reviewed by Davis
(1975) who described two genera and five species from
the West Indies. However, none of these taxa was
known from Cuba. Recent collections from Cuba,
chiefly on the three main mountain chains, have
resulted in the discovery of new records including
several new species. In this work, five new species
belonging to Paucivena Davis, 1975 are described. Keys
for identification of known stages of all species and
information on natural history are also provided.
MATERIALS AND METHODS
Individuals of all species, except one whose
representatives were taken flying at day, were collected
as larvae and pupae in the field and reared in the
laboratory. Lab-reared larvae were provided with field
collected hosts until pupation. All type material is
deposited at Instituto de Ecologia y Sisteméatica
(CZACC).
Diagnostic morphological characters employed
follows Davis (1964, 197: 5) and Henderickx (1982). Setal
maps of larvae follow Hinton (1946) and Stehr (1987).
Measurements were taken with an ocular micrometer in
a Carl Zeiss Stemi 2000 stereoscopic microscope.
Interocular index of head was calculated as a ratio
between the vertical diameter of the compound eye and
interocular distance measured at a point across the frons
midway between the base of antennal sockets and the
anterior tentorial pits (Davis, 1975):
Interocular index= vertical eye diameter/ interocular
distance
Characters of the two previously known species,
Paucivena reticulata Davis, 1975 and Paucivena
hispaniolae Davis, 1975, were taken from original
descriptions and illustrations. Additionally, one P.
reticulata specimen placed at CZACC, was examined.
Abbreviations: x- mean, SD- standard deviation, CV-
coefficient of variation.
RESULTS
Paucivena Davis, 1975
This genus is known only from the western part of the
cic: Davis (1975) described P. hispaniolae from
Dominican Republic, Hispaniola, and P. reticulata from
Puerto Rico and Jamaica. In the same work, this author
mentioned the possible presence of Paucivena on Cuba.
This was confirmed recently by Nuifiez (2004) based on
unknown species from Topes de Collantes, in the
Cuban central mountains, which are described here.
Characters that best define Paucivena males are labial
palpi with a single segment not fused, origin of antennal
rami at base of each antennal segment, two pairs of
tibial spurs on mid and hindlegs, reduced wing venation
and the abbreviated genitalia. Females may be
recognized by the possession of compound eyes and
functional legs, and the lack of antennae and wings.
Larvae feed on several hosts including mosses and
lichens growing on rocks and bark, and detritus.
Paucivena ferruginea Niiiez, new species
(Figs. 1, 6, 11, 16, 17, 22, 24, 26, 28-35, 41, 4546, 52)
Diagnosis: Paucivena ferruginea male differs from
all other Paucivena by its brown coloration with slight
ferruginous iridescence. Other diagnostics characters
are the acute and heavily sclerotized sacculus and the
bifid saccus of its genitalia.
Male (Figs. 1, 6, ‘1, 16, 17). Head: brown. Antennae with 23
segments; lateral pectinations about 2—2.5 times length of supporting
segment. Vertical diameter of eye 0.8 the interocular distance. Thorax
(Figs. 6, 11): anterior half dark brown, posterior half brown with slight
ferruginous iridescence; underside pale brown except inner surface of
JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY
Fics. 1-5. Paucivena spp. adult males. 1 P. ferruginea, n. sp.;
orientalis, n. sp. Scale= 3 mm,
coxa and femur of forelegs which are dark brown. Vestiture dense,
scales hairlike. Wings brown with slight ferruginous iridescence; basal
two thirds of costa on FW dark brown. Tibial spurs approximately 0.35
the length of basal tarsal segments (Fig. 6). Scales at discal cell
variable in shape: oblanceolated, ovobated, with rounded or acute
apices, or hairlike. FW with 9 veins, all veins separated (Fig. 11);
accessory cell present; CuP not reaching inner margin; one anal vein.
HW with 7 veins, all veins separated except M, and CuA, which have
a unique origin; cross vein between Sc and Rs “absent; two anal veins.
Wing expanse: 11 mm. Abdomen: brown with slight ferruginous
celeccence: underside pale brown. Vestiture dense, scales heaeeleet
Genitalia (Figs. 16-17): tegumen broad, with a pair of sparsely setose
apical lobes. Valvae with pulvilli setose; apex of sacculus acute and
heavily sclerotized, armed with three spines; cucullus apically rounded
and sparsely setose. Saccus bifid, apices blunt; approximately 0.1 the
length of main body. Aedeagus simple, cylindrical, 0.7 times the
length of valvae.
Female (Figs. 22, 24, 26). Length: 7.5 mm. Vermiform.
Stramineous, with two longitudinal bands of brown spots on dorsum.
Head (Fig. 22): stramineous, eyes black. Slightly sclerotized. Shape
near ovoid (ventral view); eyes compound, well developed, subventral.
Labial palpi 1-segmented, 100% fused; antennae absent. Thorax (Fig.
24): patterned as above; body wall slightly sclerotized. Legs functional,
armed with numerous tiny spines; tarsi 1-segmented with a pair of
claws at distal end (Fig. 24); wings absent. Abdomen: color pattern
disappearing at A2-A3; membranous and naked except for a ring of
dense brownish ochre hairlike scales around A7. External genitalia
reduced (Fig. 26), largely membranous. Two pairs of apophyses
present; anterior pair elongated, free except bifid base fused with
tegument; posterior pair straight and free.
Larva (Figs. 28-34). Length of largest larva 11 mm, maximum
width of head capsule 1.2 mm. Head and thorax whitish with dark
2 P. pinarensis, n. sp.; 3 P. fusca, n. sp.; 4 P. cubana, n. sp.; 5 P.
fuscous longitudinal bands continued on thorax forming a striated
pattern; spiracle on TI as large as spiracle on A8, both larger than
spiracles on Al—A7. Head (Figs. 28-32): patterned as above, lateral
area with five elongated bands; labrum ochre; an elongated band on
adfrontal sclerite and frons extending from C1 to slightly beyond AF 1;
AF2 and P2 absent (Fig. 28). Six stemmata present; five arranged i in
an inverted semicircle, a sixth more distant and ventrad, immediately
anterior to $3 (Fig. 29). Labrum (Figs. 30, 31) with LA3 isolated.
Mandibles with four acute teeth and a fifth, blunt tooth (Fig. 32).
Thorax (Fig. 33): patterned as above, three longitudinal bands
between body axis and lateral margin of shield, interrupted on
metathorax. TI with shield bearing D, SD, XD and L groups; XD-
group and L2 in vertical line near anterior margin of shield, SD-group
slightly posterior; XD1 about equal in length to SD1, about 2 times
longer than XD2 and D2; D1 dorsoposterior to XD1, about 1/4 its
length; SD2 above SD1, slightly posterior and about 1/4 its length; L-
group trisetose, Ll about 3 times longer than L2 and _ L3,
posteroventral to L2; L3- slightly longer than L2; spiracle
dorsoposterior to L-group, diagonal; SV-group in horizontal line on
elongated pinnaculum, SV2 about 3/5 length of SV1; MV2 on same
pinnaculum, anterior to SV2; V1 about equal in length to SV2,
posteroventral to SV1. TH-TIII: D and SD groups in a vertical line on
same pinnaculum; D2 about 2-21/2 times longer than D1; SD1 about
3 times longer than SD2; L2 separated from pinnaculum bearing L1
and L3, about 1/2 length of L3; L1 3/5 length of L2, dorsoposterior to
L3; SV group on same pinnaculum; SV1 about twice length of SV2; V1
slightly shorter than SV1. Abdomen (Figs. 33, 34): integument dark
brown, pinnacula brownish ochre. Al: D-group on separated
pinnacula, D1 dorsoposterior to D2 and about 3 1/2 times longer; SD1
above and slightly anterior to spiracle, slightly shorter than D2; SD2
minute, anterodorsal to spiracle; L-group trisetose, on separated
pinnacula; L1 posterior to L2 and about twice its length; L3 below,
VOLUME 60, NUMBER 3
midway between L1 and L2, equal in length to L1; SV-group bisetose
and on same pinnaculum, SV2 anterodorsal to SV1 and about 1/3 its
length; V1 anteroventral to SV1 and about half its length (Fig. 33). A2
(not shown) equal to Al except SV- group trisetose, SV3 on
pinnaculum bearing V1, below SV2 and about equal its length. A3—A6
with four pairs of. prolegs, crochets (22-24) seroneiall uniserial,
arranged in a lateral penellipse; setae as above except SV-group on
pinnaculum containing proleg. A7 (not shown) as above except SV-
group bisetose. AS with setae as above except L-group arranged in a
more or less vertical line, L1 on same pinnaculum bearing SD-group
and spiracle; SV-group unisetose. A9 with all setae arranged in a more
or less vertical line; SD1 and D2 on same pinnaculum, SD1 about 5
times longer than D2 and about equal in length to D1. A10 (Fig. 34):
anal plate with SD1 slightly longer than D1, about 2-2 1/2 times
longer than D2; prolegs bearing 24 uniordinal crochets, uniserial,
arranged in a lateral penellipse; anterior margin of shield irregular.
ee case (Fig. 41). Dimensions: length of main body: ¢ 12
mm (x=12, SD=0, CV=0, n=2), total length of projecting fragments:
17- ii mm (X=17.5, SD=0.71, CV=4%, n=2): 9 2 13mm, total length
of projecting fragments: 22 mm; maximum diameter: 6 2.6 mm
(x=2.6, SD=0, CV=0, n=2), ° 3.3 mm. Almost cylindrical in its entire
length; soft. External cover formed by a basal layer of tiny vegetal
fragments covered by large fragments of thin herbaceous stems,
lengthwise arranged and parallel, various projecting backward from
case.
Male pupa (Figs. 45-46). Length 5.8 mm. Uniform yellowish
ochre. Frontal ridge absent, frons rounded. Antennal sclerites
extending slightly beyond apex of prothoracic legs (Fig. 45). Wing
sheaths extending midway along A3. Sclerites of metathoracic legs
extending to anterior margin of A4. Cremaster reduced, consisting in
a pair of small spines, ventrally curved, arising form a broad conical
base; anal groove Y-shaped. Dorsum of A3-A7 with 2-3 irregular rows
of spines ‘directed caudad on anterior margin, both end of rows
thickened (Fig. 46); A8 with spines grouped in an elliptical patch:
areas surrounding rows covered by hundreds of tiny spines arranged
in 2-5 series or solitaire. Dorsum of A5—A7 with single posterior row
of slender spines oriented caudad. Tabulation of spines shown in Table
1.
Female pupa (Figs. 50-51). Length 7.5 mm. Uniform yellowish
ochre. Head with eyes and labial palpi distinct (Fig. 50). Thorax with
leg sclerites distinct; wings absent. Cremaster vestigial, reduced to a
coarse and rough area around anal groove; anal groove Y-shaped.
Dorsum of A6-A7 with 2—4 irregular rows of small spines directed
caudad on anterior margin, AS with spines grouped in an elliptical
patch. A4—A6 with a single posterior row of slender spines cephalad
oriented (Fig. 51). Tabulation of spines shown in Table 2.
Types. Holotype 6 (with associated larval case and pupal exuvium),
reared from larva (emerged 22 June 2003), CUBA: Sancti Spiritus
province, Topes de Collantes, Pico Potrerillo, 973 m, 18 May 2003 (R.
Niifiez), slides RNA 014, 038, 046, 077, 078. Paratypes,2 (with
associated larval case and pupal exuvium), reared from pupa (emerged
20 May 2003), CUBA: Sancti Spiritus province, Topes de Collantes,
Parque Codina, 800 m, (R. Niiiiez), slide RNA 042; 2 larvae (with
associated larval case), same data as holotype; 3 larval cases, same data
as holotype; 1 larval case, same data as holotype except 17 June 2004.
aA
NA
Fics. 6-10. Paucivena spp. male legs. 6 P. ferruginea, n. sp.; 7 P. pinarensis, n. sp.; 8 P. fusca, n. sp.; 9 P. cubana, n. sp.; 10 P. ori-
entalis, n. sp. Top- prothoracic leg, middle- mesothoracic leg, bottom-metathoracic leg. Scale= 2 mm.
Natural history observations. Larvae were found
feeding on mosses, Orthostichidium guyanense (Mont.)
V.F. Brotherus (Pterobryaceae) and another
unidentified species, on bark of an unidentified bush.
Two adults were reared from larvae: a male emerged
from the pupa after a month; a female emerged and was
observed hanging from distal end of the case.
The species was found in two localities with very
different vegetation and climatic conditions. Parque
Codina is characterized by a secondary and very humid
evergreen forest with the lower strata protected by a
dense canopy. A dry scrub, included in the mogote
vegetational complex, g grows on top of Pico Potrerillo, an
enviroment very epaunenl to wind and solar radiation.
Distribution (Fig. 52). Known only from two
localities at Topes de Collantes region, Trinidad
Mountains, central Cuba.
Etymology. The species name is derived from the
slight ferruginous iridiscence of male wings.
Remarks. The female may be separated from that of
P. orientalis, n. sp., by its greater size (length 7.5 versus
5 mm), its more elongated genitalia, the presence of
JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY
only two longitudinal spot bands on tegument (six in the
other species) and its unswollen tibiae. Larvae may be
recognized by the loss of AF2 on head and the isolation
of LA3 on labrum. The larval case is also diagnostic for
this species within the genus. Davis (1964, 1975)
reported similar cases from Haiti and Trinidad. Davis
noted the of these with
constructed by species of Epichnopteryx Hiibner and
Psyche Schrank, two Old World genera, considering
them as a probable introduction from the Old W forld.
Larval cases of P. ferruginea prove that this construction
pattern is not exclusive to Old World species.
similarity cases those
Paucivena pinarensis Nunez, new species
(Figs. 2, 7, 12, 18, 42, 47, 52)
Diagnosis: Paucivena pinarensis males possess a
color pattern similar to that of P. cubana, sp. n., P.
orientalis, sp. n., and P. reticulata. However, the
reticulated pattern is weaker in P. pinarensis due to its
more obscure background color. Compared to other
Paucivena within this group, P. pinarensis males exhibit
several diagnostic characters: absence of saccus in its
M1
M243
3A CuA1
1A Cup CuA2
+
11
2a
14
Fics. 11-15. Paucivena spp. wing venation. 11 P. ferruginea, n. sp.;
15 P. orientalis, n. sp. Scale= 3 mm.
15
12 P. pinarensis, n. sp.; 13 P. fusca, n. sp.; 14 P. cubana, n. sp.;
Ul
VOLUME 60, NUMBER 3 Ip?
TABLE 1. Rows and spines numbers per rows on dorsum of abdominal segments of Cuban Paucivena male pupae.
Abdominal segments
Rows
I II Ill IV Vv VI VII VIII
Paucivena ferruginea anterior 0 0 269 135 140 176 122 48
n=1
posterior 0 0 0) 0 42 44 4S 0
Paucivena pinarensis anterior 0 0 57 63 64 60 49 49
n=1
posterior 0 0 1 ri 38 38 14 0
Paucivena cubana anterior 0 0 0 52-70 47-58 42-55 29-45 22-24
n=3
posterior 0 0) 31 35-44 37-46 36-42 32-36 0
Paucivena orientalis anterior 0 (0) 0 102-136 96-125 95-111 90-96 25-32
n=3
posterior 0 0 0 27-38 23-39 24-32 23-50 0
TABLE 2. Rows and spines numbers per rows on dorsum of abdominal segments of Cuban Paucivena female pupae.
Abdominal segments
Rows
I II Il IV Vv VI VIL VIII
Paucivena ferruginea anterior 0 0) 0) 0 (0) 61 90 21
n=]
posterior 0 0 0 7 58 iss) (0) 0
Paucivena cubana anterior 0 0 0 3 14 20 21 0
ne)!
posterior 0 0 6 56 78 qAAl 0 0
Paucivena orientalis anterior 0 0 0 5-10 9-10 29-30 92-96 18—22
n=3
posterior 0 0 0 16-17 23-27 21-22 0 0
126 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY
tegumen
cucullus sacculus aedeagus
pulvilli
SaCCus
Fics. 16-21. Paucivena spp. male genitalia, ventral view. 16 P. ferruginea, n. sp., main body; 17 P. ferruginea, n. sp., aedeagus; 18
P. pinarensis, n. sp.; 19 P. fusca, n. sp.; 20 P. cwbana, n. sp.; 21 P. orientalis, n. sp. Scale= 0.5 mm.
Fics. 22-23. Paucivena spp. female head, ventral view. 22 P. ferruginea, n. sp.; 23 P. orientalis, n. sp. Scale= 1 mm.
VOLUME 60, NUMBER 3
25
Fics. 24-25. Paucivena spp. female legs. 24 P. ferruginea, n. sp., scale= 0.5mm; 25- P. orientalis, n. sp., scale= 0.25 mm. Top-
prothoracic leg, middle- mesothoracic leg, bottom-metathoracic leg.
anterior
apophyses
27
26
Fics. 26-27. Paucivena spp. female genitalia, ventral view. 26 P. ferruginea, n. sp.; 27 P. orientalis, n. sp. Scale= 0.5 mm
128 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY
SD1 py 34
Fics. 28-34. Chaetotaxy of Paucivena ferruginea, n. sp., largest larva (last instar?). 28 Head, dorsal view; 29 Stemmata of left side,
lateral view; 30 Labrum, dorsal view; 31 Labrum, ventral view; 32 Right mandible, ventral view; 33 Prothorax, mesothorax and ab-
dominal segments 1, 3-6, 8 and 9, lateral view; 34 Anal shield, dorsal view. Scale, 28—29= 0.5 mm; 30-32= 0.2 mm.
VOLUME 60, NUMBER 3
129
Fics. 35-40. Chaetotaxy of Paucivena cubana, n. sp., largest larva (last instar?). 35 Head, dorsal view; 36 Stemmata of left side,
lateral view (scale= 0.5 mm); 37 Labrum, dorsal view; 38 Labrum, ventral view; 39 Right mandible, ventral view (scale= 0.2 mm);
40 Prothorax and mesothorax, lateral view (abdomen damaged). Scale, 35-36= 0.5 mm; 37—39= 0.2 mm.
genitalia; the relative length of its tibial spurs, 0.3 versus
0.15 (PB. cubana, P. orientalis) and 0.5 (P. reticulata); and
interocular index 0.7 versus 1.1 (P. reticulata) and 1.5 (P.
cubana).
Male (Fig. 2, 7, 12, 18).: Head: pale brown, labial palpi dark
brown. Antennal tips broken; lateral pectinations 3 about times length
of supporting segment. Moe diameter of eye 0.7 the eroculae
distance. Thorax ( (Figs. 7, 12): dark brown; underside pale brown,
inner surface of fore and midlegs dark brown, joints pale brown.
Vestiture dense, scales hairlike. Tibial spurs approximately 0.3 the
length of basal tarsal segments (Fig.
costa dark brown: ground brown streaked with dark brown forming a
faint reticulated pattern; fringe with various tones of brown. Scales at
discal cell variable in shape: oblanceolated and ovobated, with
rounded or acute apices, with scattered hairlike scales. Venation (Fig.
12) as in P. ferruginea. HW uniform brown; fringe with various tones
of brown. Venation as in P. ferruginea except M,,, and CuA, which
arise separate from cell, M,_, CUCL from CuA, and M,, only one
anal vein present. Wing expanse: 9.2 mm. Abdomen: pale brown.
Vestiture dense, scales hairlike. Genitalia (Fig. 18): tegumen broad,
with a pair of sparsely setose apical lobes. Valv. ae with pulvilli setose;
apex of sacculus armed with five strong spines; cucullus apically
rounded and sparsely setose. Saccus absent. Aedeagus simple,
cylindrical, 0.6 times length of valvae.
7). FW with basal two thirds of
Female. Unknown.
Larva. Unknown.
Larval case (Fig. 42). Dimensions: length: ¢ 9.8 mm; maximum
diameter: ¢ 3.4 mm. Fusiform in outline; soft. Exterior heavily
covered with vegetal fragments, leaves and short stems, and mosses
various shaped and oriented lengthwise. Cases were found hanging
from silk filaments (3.5 mm in ler ngth) attached to rocks and tree
trunks.
Male pupa (Fig. 47). Length 5.1 mm. Ochre, wing sheaths reddish
brown. Frontal ridge absent, frons rounded. Antennal sclerites
extending slightly beyond apex of prothoracic legs. Wing sheaths
extending to posterior margin of A3. Sclerites of metathoracic legs
extending to posterior margin of A4. rae consisting in a pair of
strong spines ventrally curved; anal groove Y-shaped. Dorsum of
A3-A8 with an irregular row of spines on anterior margin; areas
surrounding rows covered by hundreds of tiny, solitaire spines.
Dorsum ae A3-A7 with single posterior row of slender spines.
Tabulation of spines shown in Table 1.
Female pupa. Unknown.
Types. Holotype, ¢ (with associated larval case and pupal
exuvium), reared from larvae (emerged 28 February 2004), CUBA:
Pinar del Rio province, Sierra del Rosario, Taco Taco River shore 1 km
northeast from Jardin de Aspiro, 200 m, 28 November 2003 (R.
Niifiez), slides RNA 054, 079, 080. Paratypes: 4 larval cases, CUBA:
Pinar del Rio province, Sierra del Rosario, Jardin de Aspiro, 150 m,
130
28-29 November 2003, (R. Nuimez).
Natural history observations. The single larva was
found on limestone rock near the Taco Taco River shore
and its food source can not be accurately identified. In
the lab, the larva was fed with several crustose lichens
and mosses collected on its substrate. Other larval cases
were found on rocks and tree trunks in an old
abandoned Botanical Garden (Jardin de Aspiro). The
unique adult emerged after a month; emergence took
place between 0900 and 1130h.
At the Taco Taco River shore secondary remnants of
gallery forest are present whereas at Jardin de Aspiro
several introduced and native trees grow forming groves
separated by cleared areas occupied by camping
installations.
Distribution (Fig. 52). Known only from two close
JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY
localities at Sierra del Rosario, Pinar del Rio province.
eee The species name is derived from the
name of the Cuban province where the type locality,
Pinar del Rio, is located.
Remarks. The larval case is identical to that of P.
cubana. The male pupa may be easily identified by the
unique arrangement of a rows on the dorsum of
abdominal segments (Table 1
Paucivena fusca Nunez, new species
Paucivena sp. n. 1: Nunez, 2004: 155
Figs. 3, 8, 13, 19, 52
Diagnosis. Males of P. fusca may be recognized by
their uniform dark brown coloration. Within the genus
only P. hispaniolae exhibits a similar coloration but it has
Key to the adult males of Paucivena
1. Wings dark brown with a ferruginous shine; genitalia with an acute and heavily sclerotized sacculus
and a bifid saccus (Fig. 16)
P. ferruginea
- Wings with a different color pattern; genitalia with sacculus not acute and weakly sclerotized, saccus
not bifid 2
2. Dorsum of wings and body entirely dark brown, almost black 3
- Dorsum of wings and body with light color pattern 4
3. Body whitish grey ventrally; eyes of medium size (vertical diameter of eye 1.1 the interocular
distance); genitalia with the margins of apex of sacculus and the apical lobes of tegumen smooth
P. hispaniolae
- Body dark brown ventrally; eyes small (vertical diameter of eye 0.8 the interocular distance);
genitalia with the margins of apex of sacculus and the apical lobes of tegumen serrulated (Fig. 19)
P. fusca
4. Tibial spurs much reduced, approximately 0.15 the length of basal tarsal segment 5
- Tibial spurs less reduced, approximately 0.3 the length of basal tarsal segment or longer 6
5. Eyes very large (vertical diameter of eye 1.5 the interocular distance): wing expanse: 12 mm
P. cubana
- Eyes very small (vertical diameter of eye 0.7 the interocular distance); wing expanse: §.2-9.1 mm
P. orientalis
6. Eyes of medium size (vertical diameter of eye 1.1 the interocular distance); tibial spurs large,
approximately 0.5 the length of basal tarsal segment; FW reticulated pattern distinct
P. reticulata
- Eyes very small Geticall diameter of eye 0.7 the interocular distance); tibial spurs reduced,
approximately 0.3 the length of basal tarsal segment; FW reticulated pattern weak, indistinct
P. pinarensis
Key to the known larvae of Paucivena (excludes P. reticulata, P. pinarensis, P. fusca and P. orientalis,
which are unknown)
1. Head and thorax whitish to light tan with irregular patches of dark fuscous; meso and metathorax
with an extra seta (SD 1a?)
P. hispaniolae
- Head and thorax whitish with longitudinal dark fuscous bands arranged in a striated pattern; meso
and metathorax without an extra seta
9)
4
2. Head with AF2 absent (Fig. 28); LA3 on labrum isolated from the rest (Fig. 30); abdominal
integument dark brown
- Head with AF2 present (Fig.
white
35); LA3 on labrum not isolated (Fig. 37
P. ferruginea
); abdominal integument dirty
P. cubana
VOLUME 60, NUMBER 3
Key to the known male pupae of Paucivena (excludes P. reticulata and P. fusca, which are unknown)
1. Anterior margin of A3 without spines
9)
- Anterior margin of A3 with at least one row of spines 3
2. Spines absent from posterior margin of A3; rows at anterior margin of A4-A7 with 90 or more spines
(Table 1); length 3.8-4.5 mm
P. orientalis
- Spines present on posterior margin of A3; rows at anterior margin of A4-A7 with 70 or fewer spines
(Table 1); length 5.0-5.4 mm
icy)
P. cubana
. Rows on anterior margin of A3-A7 with more than 100 spines (Table 1); length 5.8 mm
P. ferruginea
- Rows on anterior margin of A3-A7 with less than 100 spines (Table 1); length 5.0-5.1 mm 4
4. Spines present, although reduced, on posterior margin of A3-A4; cremaster consisting in a pair of
strong spines ventrally curved
P. pinarensis
- Spines completely absent from posterior margin of A3-A4; cremaster consisting in a pair of small
spines ventrally curved
P. hispaniolae
Key to the female pupae of Paucivena (excludes P. reticulata, P. pinarensis and P. fusca, which are
unknown)
1. Anterior rows of spines absent from dorsum of A3-A4
- Anterior rows of spines present on dorsum of A3-A4
Posterior row of spines reduced but present on dorsum of A4; cremaster vestigial, reduced to a
bo
coarse and rough area around anal groove; length 7.5 mm
w we
P. ferruginea
- Posterior row of spines absent from dorsum of A4; cremaster relatively well developed, consisting in
pair of short acute spines; length 10-11 mm
oy)
mm
- Anterior margin of AS covered only by hundreds of tiny spines; length 8.0 mm
the underside of body whitish grey. Other useful
characters are FW shape (more rounded in the Cuban
species), eye size (interocular index 0.8 in P. fusca and
l.linP hispaniolae) and the serrated margins at apex of
sacculus and apical lobes of tegumen in the genitalia of
P. fusca, both smooth in P. Pecapiallacy
Male (Figs. 3, 8, 13, 19). Head: dark brown. Antennae with 21-22
segments; lateral pectinations 2 times length of segment. Vertical
diameter of eye 0.8 the interocular distance. Thorax (Figs. 8, 13):
uniform dark brown. Vestiture dense, scales hairlike. Tibial spurs
approximately 0.25 the length of basal tarsal segments (Fig. 8). Scales
at discal cell of FW oblanceolated and ovobated with rounded or
acute apices. Venation (Fig. 13) as in P. ferruginea. HW venation as in
P. pinarensis, except the origin of M,_, which is closer to CuA, than to
M,. Wing expanse: 10-11 mm (x= 10.2, SD=0.23, CV= 4%, n=9).
Abdomen: dark brown. Vestiture dense, scales hairlike. Genitalia (Fig.
19): tegumen broad with apical cleft, lobes minutely serrated and
sparsely setose. Valvae with pulvilli setose; apex of sacculus strongly
serrated; cucullus rounded, apex sparsely setose. Saccus reduced,
approximately 0.2 the length of main body. Aedeagus simple,
cylindrical, 0.6 times the length of valvae.
Female. Unknown.
Inmature stages. Unknown.
Larval case. Unknown.
Types. Holotype, ¢ CUBA: Sancti Spiritus province, Topes de
Collantes, Pico Potrerillo, 973 m, 6 May 2002 (R. Nuifiez), slides RNA
011, 015, 039. Paratypes, 3 3, same data as holotype, slides RNA 010,
022. 5d, same data as holotype except 17 June 2004, slides RNA 026,
029, 041.
Natural history observations. All individuals were
found flying at noon on the top of Pico Potrerillo except
P. hispaniolae
Anterior margin of AS with a row of spines surrounded by hundreds of tiny spines; length 4.9-5.1
P. orientalis
P. cubana
a single specimen seen flying between rocky walls at
peak access. This species shares its habitat, dry scrub on
the top of Pico Potrerillo, with P ferruginea and P.
cubana.
Distribution (Fig. 52). Known only from
Potrerillo at Trinidad Mountains, central Cuba.
Etymology. The species name is derived from its
uniform dark brown color.
Pico
Paucivena cubana Ninez, new species
Paucivena sp. n. 2: Nunez, 2004: 155
Figs. 4, 9, 14, 20, 35-40, 43, 48, 52
Diagnosis. Males of P. cubana may be separated
from other Paucivena with reticulated wing pattern by
the following characters: 12 mm of wing expanse (the
largest eine the genus), elongated legs with tiny tibial
spurs (approximately 0.15 the length of basal tarsal
segment) and large eyes (interocular index 1.5, the
largest within the genus).
Male ( Figs. 4, 9, 14, 20). Head: pale yellowish ochre. Antennae
with 18 segments; lateral pectinations 1.5-2 times the length of
segment. Eyes large, vertical diameter of eye 1.5 the interocular
distance. Thorax (Fig. 9, 14): pale yellowish ochre with scattered dark
brown scales. Vestiture dense, scales _hairlike. Tibial
approximately 0.15 the length of basal tarsal segments (Fig. 9). Wings
thinly scaled. FW (faded) yellowish ochre with scattered dark brown
spurs
JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY
Fics. 41-44. Paucivena spp. larval cases. 41 P. ferruginea, n. sp.;
sp. Scale= 3 mm.
scales; dark brown scales concentrated at basal half of anterior and
posterior margins, forming a faint reticulated pattern on basal two
thirds; fringe ochre. Scales at discal cell variable in shape:
oblanceolated and ovobated, with rounded or acute apices, or hairlike.
Venation as in P. ferruginea, except accessory cell which is wider,
closing below origin of R,,, (Fig. 14). HW pale y yellowish ochre with
scattered dark brown scales: paler than FW; fringe pale yellowish
ochre. Venation as in P. fusca. Wing expanse: 12 mm. Abdomen: pale
yellowish ochre with scattered dark brown scales. Vestiture dense,
scales hairlike. Genitalia (Fig. 20): tegumen broad, apex damaged.
Valvae with pulvilli sparsely setose; apex of sacculus armed with three
spines; cucullus rounded with apex sparsely setose. Saccus reduced,
approximately 0.2 the length of main body. Aedeagus simple,
cylindrical, 0.7 times the length of valvae.
42 P. pinarensis, n. sp.; 43 P. cubana, n. sp.; 44 P. orientalis, n.
Female. Unknown.
Larva (Figs. 35-40). Length of longest larva 6.9 mm, maximum
width of head capsule 1.0 mm. Head and thorax whitish with dark
fuscous longitudinal bands continued on thorax forming a striated
pattern. Head (Figs. 35-39): as in P. ferruginea except, AF2 present;
adfrontal sclerite with elongated spot on upper third covering origin of
AF2; frons with spot covering origin of Fl and C2 (Fig. 35); "AFA
closer to AF2 than to AFI. Sixth stemma immediately anterior to $2
and $3, midway between them (Fig. 36). Labrum (Figs. 37, 38) with
setae approximately mesad except, LA] and M2 distinctly closer to
border. Mandibles with four acute teeth and a fifth, blunt tooth (Fig.
39). Thorax (Fig. 40): as in P. ferruginea with the following exceptions.
TI with XDI about equal in length to XD2 and SD1, and about 1 1/2
times longer than D2; MV2 separated from pinnaculum bearing SV-
VOLUME 60, NUMBER 3
group. TII-TII: D2 about 3 times longer than D1; SD1 about 4 times
longer than SD2. Abdomen damaged, integument dirty white.
Larval case (Fig. 43). Dimensions, length: ¢ 10-11 mm (x=10.5,
SD=0.71, CV=7%, n=2), 2 18 mm; maximum diameter: 3 4.0 mm
(x=4.0, SD=0, CV=0, n=2), 2 6.5 mm. Fusiform in outline, soft.
Exterior densely covered by fragments of leaves, small herbaceous
stems and mosses of different shape and lengthwise oriented. Cases
were found attached to rocky walls at Pico Potrerillo and banana trees,
Musa paradisiaca L. (Musaceae), at Mogote Mi Retiro, hanging from
silk threads (length: ¢ 3.5 mm, 2 5 mm).
Male pupa (Fig. 48). Length 5.0-5.4 mm (x=5.2, SD=0.23,
CV=4%, n=2). Uniform brownish ochre. Frontal ridge absent, frons
rounded. Antennal sclerites extending slightly beyond apex of
prothoracic legs. Wing sheaths extending to anterior margin of A4.
Sclerites of metathoracic legs extending midway along A5 or its
posterior margin. Cremaster consisting in a pair of strong and very
close spines, ventrally curved and abruptly tapered at apex; anal
groove Y-shaped. Dorsum of A4—A8 with 1-2 irregular rows of spines
on anterior margin; anterior margin A3 and areas surrounding spine
rows on A4—AS covered by hundreds of tiny, solitaire spines. Dorsum
of A3—A7 with single posterior row of slender spines. Tabulation of
spines shown in Table 1.
Female pupa. Length 8 mm. Uniform ochre. Head with eyes and
labial palpi distinct. Thorax with leg sclerites distinct; wings absent.
foreleg
midleg
antenna
forewing
Al
A2
A3
hindleg
A4
A10
45
labial palpi
Cremaster vestigial, reduced to a small pair of blunt, widely separated
spines; anal groove Y-shaped, with a pair of small rounded tubercles
on either side. Dorsum of A4 with single anterior row of reduced,
widely spaced spines; A5—A7 with 2-3 irregular rows of spines on
anterior margin; anterior margin of A3, areas surrounding rows on
A4-A8 and anterior margin of A8 covered by hundreds of tiny spines,
solitaire or in 2-4 series. A3—A6 with single posterior row of slender
spines. Tabulation of spines shown in Table 2.
Types. Holotype, ¢ (with associated larval case and pupal
exuvium), reared from larva (emerged June 2002), CUBA: Sancti
Spiritus province, Topes de Collantes, Caburni River depression, 500
m, 30 April 2002 (R. Niifiez), slides RNA 013, 016, 023, 040, 045.
Paratypes, 1 larva, same data as holotype, slides RNA 065, 066, 070: 6
larval cases (some with associated pupal exuvium), CUBA: Sancti
Spiritus province, Topes de Collantes, mogote Mi Retiro northern
base, 800 m, 17 May 2003 (R. Nujiez); 4 larval cases (some with
associated pupal exuvium), CUBA: Sancti Spiritus province, Topes de
Collantes, southern side of rocky outcrop at Pico Potrerillo access, $50
m, 18 May 2003 (R. Nuiiez), slides RNA 075, 076; 3 larval cases (one
with associated pupal exuvium), same data as preceding except 17
June 2004; 2 larval cases, CUBA: Sancti Spiritus province, Topes de
Collantes, Parque Codina, 800 m, 20 May 2003 (R. Niifiez).
Natural history observations. Larvae fed on
Plagiochila sp. (Plagiochilaceae), a hepatic growing on
Hebel Hl ail Te
VERN & lang
TT ae
ne
46
Fics. 45-46. Paucivena spp. male pupa. 45 P. ferruginea, n. sp., ventral view; 46 P. ferruginea, n. sp., dorsal view
134 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY
Fics. 47-49. Paucivena spp. male pupa. 47 P. pinarensis, n. sp., ventral view; 48 P. cubana, n. sp., ventral view; 49 P. orientalis,
n. sp. Scale= 0.25 mm.
Fics. 50-51. Paucivena ferruginea, n. sp., female pupa. 50 Ventral view; 51 Dorsal view. Scale= 0.25 mm.
VOLUME 60, NUMBER 3
rocks on the Caburnf River shore. However, larvae
probably use other hosts since cases were found on
different substrates at other localities. At the Pico
Potrerillo access cases were found attached to rocky
walls covered by crustose lichens, at mogote Mi Retiro
base they were located on banana plants whereas at
Parque Codina they live on trunks of native trees. The
single adult emerged after a month as pupa.
Bruen cubana inhabits localities with very
different vegetation and climatic conditions. At Parque
Codina it was found in the lower strata of secondary
evergreen forest, a very humid habitat, and at the
mogote Mi Retiro base in cultivated land, banana and
cotfee (Coffea arabica L., Rubiaceae), close to the
mogote rocky wall. On the other hand, Caburni River
shores are covered by gallery forest remnants, today
dominated by an introduced tree (Syzigium jambos L..,
Myrtaceae), whereas at the Pico Potrerillo access cases
were attached to rocky walls surrounded by evergreen
forest.
Distribution (Fig. 52). Known from four localities at
Topes de Collantes, Trinidad Mountains, central Cuba.
Etymology. The species name is derived from the
name of the Cuban island.
Remarks. The larval color pattern is identical to that
of P. ferruginea; however, differences in chaetotaxy and
abdominal coloration are present. Male and female
pupae are easily distinguished by the arrangement of
abdominal dorsal spines (Tables 1, 2).
Paucivena orientalis Ninez, new species
Figs. 5, 10, 15, 21, 23, 25, 27, 44, 49, 52
Diagnosis. Adult males of P. orientalis possess a
color pattern similar to that of P. pinarensis, P. cubana,
and P. reticulata. P. orientalis may be separated from P.
pinarensis by its smaller tibial spurs (0.15 versus 0.3 the
length of basal tarsal segment), the presence of a saccus
in its genitalia and its more distinct FW reticulated
pattern. From P. reticulata, it may be distinguish by its
smaller eyes (interocular index 0.7 versus 1.1) and tibial
spurs (0.15 versus 0.5 the length of basal tarsal
segment). From P. cubana it differs by its smaller size
(8.29.1 versus 12 mm of wing expanse) and eye size
(interocular index 0.7 versus 1.5).
Male (Figs. 5, 10, 15, 21). Head: pale g ereyish brown. Anten-
nae with 16 segments; lateral pectinations 22.5 times the length
of segment. Eyes small, vertical diameter of eye 0.7 the interoc-
ular distance. Thorax (Figs. 10, 15): dorsum dark brown; under-
side pale greyish brown, inner surface of legs dark brown. Vesti-
ture dense, scales hairlike. Tibial spurs approximately 0.15 the
length of basal tarsal segments (Fig. 10). Wings thinly scaled.
FW variously rounded; pale grayish brown streaked with dark
brown scales to form a reticulated pattern; fringe with various
tones of brown. Scales at discal cell mostly Sblanceolited and
ovobated, with rounded, or rarely acute apices. Venation (Fig.
15) as in P. ferruginea. HW uniform pale greyish brown; fringe
with various tones of brown; venation as in P. fusca. Wing ex-
panse: 8.2-9.1 mm (x=8.8, SD=0.49 CV=6%, n=3). Abdomen:
dorsum dark brown at both ends, remainder pale greyish brown.
Vestiture dense, scales hairlike. Genitalia (Fig. 21): tegumen
broad, with a pair of sparsely setose apical lobes! Valvae with
pulvilli sparsely setose; apex of sacculus armed with three to four
spines; cucullus with apex rounded and sparsely setose. Saccus
reduced, approximately 0.1 the length of main body. Aedeagus
simple, cylindrical, 0.6 times the length of valvae.
Female (Figs. 23, 25, 27). Length 5.0 mm. Vermiform.
Stramineous with six longitudinal bands of brown spots on dorsum
and sides of body. Head (Fig. 23): stramineous, eyes black. Slightly
sclerotized. Shape near ovoid (ventral view); eyes compound, well
developed, subventral. Labial palpi 1-segmented, almost 100% fused;
antennae absent. Thorax (Fig. 25): patterned as above; body wall
slightly sclerotized. Legs functional, armed with numerous tiny spines;
Ghise ‘swollen; tarsi 1- segmented with a pair of claws at distal end (Fi ig.
25); wings absent. iNoaloniaoe color pattern disappearing at A2-A3;
membranous and naked except for a ring of dense brownish ochre
hairlike scales around A7. External genitalia reduced (Fig. 27), largely
membranous. Two pairs of apophyses present; anterior pair elongated,
free except, bifid base fused with tegument; posterior pair straight and
free.
Larval case (Fig.
44). Dimensions, 6.5-7.1 mm
length: ¢
n=3), 2
(x=6.7, SD=0.32, CV=5%, 9-12 mm
(x=10.8, SD=1.10, CV=10%, n=5); maximum diameter: di 1.5-1.7 mm
(x=1.6, SD=0.12, CV=8%, n=3), 2 1.8-2.7 mm (x=2.2, SD=0.34,
CV=15%, n=5). Fusiform in outline, soft. Exterior ne covered by
elongated and divergent fragments of leaves and stems of bryophytes
and herbaceous plants, occasionally hair fragments are added;
material is arranged lengthwise.
Male pupa. Length 3.8-4.8 mm (x=4.3, SD=0.71, CV=17%,
n=2). Uniform brownish ochre. Frontal ridge absent, frons rounded.
Antennal sclerites usually extending beyond apex of prothoracic legs
(Fig. 49). Wing sheaths extending midway along A3. Sclerites of
metathoracic legs usually extending to A4 posterior. Cremaster
consisting in a pair of strong and widely separated, ventrally curved
spines; anal groove Y-shaped. Dorsum of A4—AS with 2-3 irregular
rows of spines on anterior margin; anterior margin of A3 and areas
surrounding rows at A4-A5 covered by hundreds of tiny spines,
solitaire or in 2-5 series. Dorsum of A4-A7 with single posterior row
of slender spines. Tabulation of spines shown in Table 1.
Female pupa. Length 4.9-5.1 mm (x=5.0, SD=0.1, CV=2%.
n=3). Uniform ochre. Head with eyes and labial palpi distinct. Thorax
with legs sclerites present; wings absent. Cremaster vestigial, reduced
to a coarse and rough area around anal groove; anal groove Y-shaped.
Dorsum of A4-A6 and AS with single row of spines on anterior
margin, A7 with two rows; areas surrounding rows at A4-A6 and
anterior margin of AS covered by hundreds of tiny spines, solitaire or
in 2-5 series. A4-A6 with single posterior row of slender spines.
Tabulation of spines shown in Table 2.
Types. Holotype, ¢ (with associated larval case and pupal
exuvium), reared from pupa (emerged 28 April 2004), CUBA:
Santiago de Cuba province, La Gran Piedra, Estacién Meteorolégica
La Gran Piedra, 1100 m, 23 April 2004 (R. Nuinez), slides RNA 024.
031, 035, 036. Paratypes, 3 larval cases (some with associated pupal
exuvium), CUBA: Granma province, La Bayamesa, abandoned coffe
plantation at Nuevo Mundo stream shore, 1600 m, 21 April 2004 (R.
Niifiez); ¢ (with associated larval case and pupal exuvium), reared
from pupa (emerged 2 May 2004), same data as holotype; ¢ (with
associated larval case and pupal exuvium), reared from pupa (emerged
9 May 2004), same data as holotype; ° (with associated larval case and
pupal exuvium), reared from pupa (emerged 24 May 2004), same data
as holotype, slides RNA 030, 034; 1 larva with its larval case, same data
as holotype; 9 larval cases (some with associated pupal exuvium), same
data as holotype.
Natural history observations. Larvae were
collected on substrates covered by several lichens and
mosses and on external walls of edifications: thus, this
species probably also feeds on detritus. The adult reared
from a larva, a female, emerged after three weeks. One
case was collected with an egg cluster and 53 larvae
hatched 1 May 2004.
At Nuevo Mundo larval cases were found on trunks of
old isolated Mangifera indica L. (Anacardiaceae) trees,
in an abandoned coffee plantation. At Gran Piedra all
individuals were located on edification walls.
Distribution. Known only from two
separated localities at Sierra Maestra, southeastern
Cuba (Fig. 52).
Etymology. The species name is referred to its
distribution range, restricted to the oriental Cuban
region.
Remarks. The larval case is diagnostic for this
species. Although constructed with small dry vegetal
fragments like those of P. pinarensis and P. cubana,
material is always cut in slender pieces and arranged
lengthwise but in a divergent way. Pupae of both sexes
are easily distinguished by the arrangement of
abdominal dorsal spines (Tables 1, 2).
widely
DISCUSSION
All new species described here fit the genus
Rio Taco Taco
Jardin de
Aspir
Topes de
, Collantes
JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY
description. However, interespecific variation in the
interocular index was noted with values between 0.7 and
1.5. This is the only character that shows significant
deviation. Measurement given by Davis (1975) was 1.1.
Deviation in Cuban representatives may be due to
differences in daily activity between species. Powell
(1973) also used an “eye index” in his study on New
World Ethmia Hiibner (Oecophoridae: Ethmiinae).
Index values were between 0.9 and 1.2 in moths known
or presumed to be nocturnal whereas diurnal species
exhibited ratios from 0.7 to 0.8. All adult males of
Paucivena species described here were lab-reared,
except those of P. fusca that always were collected flying
during the day. Data on daily activity of species
described by Davis (1975) are unavailable so this matter
will only be clarified with future work.
The female, described here for the first time, shows
characters that confirm Paucivena intermediate position
among the American psychids proposed by Davis (1975)
based on male characters. Female primitive features are
the presence of well developed compound eyes,
functional legs and the behaviour of leaving the larval
case and climbing on it, as occurs in some primitive Old
Word forms (Davis, 1964). Specialization evidences are
O Paucivena ferruginea
@ Paucivena pinarensis
@ Paucivena fusca
%* Paucivena cubana
© Paucivena orientalis
52
FIG. 52. Distribution of Cuban Paucivena spp. Scale bar in kilometers.
VOLUME 60, NUMBER 3
the reduced genitalia, the complete loss of antennae and
wings and the slightly sclerotized body wall.
In the larvae, the color pattern of the head and thorax
apparently lack diagnostic value, at least in Cuban
species. In P. ferruginea, P. cubana, P. pinarensis and P.
orientalis (the last two not described but observed
during rearing), the pattern is white to greyish white
with dark fuscous longitudinal bands. However, the
abdominal integument is differently colored in P.
ferruginea (dare brown) and P. hispaniolae and P.
cubana (dirty white). Differences in chaetotaxy may be
also used for species recognition.
Davis (1975) mentions that P. hispaniolae larvae feed
on crustose lichens growing on the bark of an
unidentified tree. Larvae of Cuban Paucivena feed on a
wide variety of hosts including mosses, hepatics, lichens
and detritus. All these food preferences have been
observed before in the Psychidae (Davis, 1964;
Hattenschwiler, 1985; Davis & Robinson, 1998) and
perhaps explain in part genus diversification together
with geographic isolation.
Paucivena appears to be well expanded on Cuba
compared to other Antillean islands such as Hispaniola,
Jamaica or Puerto Rico. However, this may due to lack
of sampling on these islands. In Cuba, more collect
effort on Psychidae is also needed. The Nipe-Sagua-
Baracoa Mountains, in the northeast part of the island,
and other habitats like coastal forests, ultramafic scrub,
and white sand savannahs are yet unexplored.
ACKNOWLEDGEMENTS
I wish to thank Adriana Lozada, Arturo Avila, Carlos Sanchez
and Ledis Regalado for their collaboration in the expeditions
and to executive and administrative personnel of Facultad
Agropecuaria de Montafia del Escambray (FAME), at Topes de
Collantes, for support during surveys. Also, I wish to thank
Ricardo Herrera and Rigel Fernandez for the photographs and
Luis Manuel Diaz for his help with the Corel Draw computer
program. Luis F. De Armas, Emily V. Saarinen and Alejandro
Barro read the manuscript suggesting valuable changes and help
with literature.
LITERATURE CITED
Davis, D. R. 1964. Bagworm Moths of the Western Hemisphere. Bull.
Unit. St.. Nat. Mus. 244:1-385.
——. 1975. A review of the West Indian moths of the family Psychi-
dae with Descriptions of New Species. Smithson. Contr. Zool.
188:1—66
. 2000. Brachygyna incae, a new genus and species of Psychidae
from Perd with atypical larval biology (Lepidoptera: Tineoidea).
Trop. Lepid. 10(2): 51-58.
Davis, D.R. & G.S. ROBINSON. 1998. The Tineoidea and Gracillari-
oidea, pp 91-117. In N.P. Kristensen (ed.), Lepidoptera, Moths
and Butterflies. Handbuch der Zoologie IV (Arthropoda): Insects
35. Walter de Gruyter and Co., Berlin and New York.
HATTENSCHWILER, P. 1985. Psychidae, pp 128-151. In J. Heath (ed.),
The Moths and Butterflies of Great Britain and Ireland. Vol. 2
Cossidae-Heliodinidae. Blackwell Scientific Ltd, Oxford, and The
Curwen Press, London.
HENDERICKS, H. 1982. Possible determination characters in psychid
females. SHILAP Rev. Lepid. 10(39): 174.
HEPPNER, J.B. 1998. Classification of Lepidoptera. Part 1 Introduc-
tion. Hol. Lepid. 5 (Suppl. 1): 1-148.
HINTON, H.E. 1946. On the homology and nomenclatura of the
setae of lepidopterous larvae, with some notes on the phylogeny
of the Lepidoptera. Trans. Roy. Entomol. Soc. Lond. 97: 1-37.
NUNEz, R. 2004. Lepidoptera (Insecta) de Topes de Collantes, Sancti
Spiritus, Cuba. Bol. S.E.A. 34: 151-159.
POWELL, J. A. 1973. A Systematic Monograph of New World Ethmiid
Moths (Lepidoptera: Gelechioidea). Smiths. Contrib. Zool.
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(ed.), Immature Insects. Kendall/Hunt Publishing Co., Dubuque.
Received for publication 3 May 2005; revised and accepted 30 May
2006
138
Journal of the Lepidopterists’ Society
60(3), 2006, 138-142
JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY
A REMARKABLE NEW RIODINID SPECIES, STALACHTIS HALLOWEENI (RIODINIDAE:
STALACHTINI), FROM MOUNT AYANGANNA, GUYANA
JASON P. W. HALL
Department of Entomology, National Museum of Natural History,Smithsonian Institution, Washington, DC 20560-0127, USA
Email: hallja@si.edu
ABSTRACT. A new riodinid species, Stalachtis halloweeni Hall n. sp. (Stalachtini), is described from Mount Ayanganna, a tepui in western
Guyana. A preliminary hypothesis of phylogenetic inter-relationships within the small genus Stalachtis Hiibner is suggested, based on an
informal study of external morphology and male genitalia. Three species groups are proposed, the phlegia, calliope and eute rpe groups, and
S. halloweeni is hypothesized to be sister to the re maining me embers of the eute rpe group.
Additional key words: endemism, montane forest, morphology, South America
Situated in the Pakaraima Mountain
the easternmost tepuis in the Guiana Shield. Like the
other tepuis in western Guyana and eastern Venezuela,
Mount Ayanganna consists of eroded sandstone
remnants of the Roraima Formation (MacCulloch &
Lathrop 2001), and its isolated high-elevation habitats
harbor significant numbers of endemic plants and
animals. During the last five years alone, several new
endemic species of frogs, snakes and lizards have been
described from Mount Ayanganna (MacCulloch &
Lathrop 2001, 2002, 2004).
During the last ten years, there have been numerous
Lepidoptera expeditions to the upland regions of
western and southern Guyana (Fratello 1996, 2001,
2003, 2005). In 1999, accompanied by several US and
2
Fics. 1-2. Stalachtis halloweeni adults (dorsal surface on
left, rac surface on right). 1. Holotype male, Mount Ayan-
ganna, Guyana ( (USNM). 2. Paratype female, Mount Ayan-
ganna, Guyana (USNM).
Range of
western Guyana, Mount Ayanganna (2042m) is one of
Guyanan colleagues, S. Fratello led the first
Lepidoptera collecting expedition to the upper slopes of
Mount Ayanganna. Among the many taxa of
butterflies collected there were several new species of
Riodinidae, including a new species of Stalachtis
Hiibner, 1818 (Stalachtini), although most of these
species were represented by only a small number of
female specimens. Fortunately, a second expedition to
Mount Ayanganna in 2002, by a different team that
included one of the 1999 Guyanan expedition members
(R. Williams), produced additional — Stalachtis
specimens, including males.
new
This new Stalachtis species is remarkable in several
respects. It is the first new species to be described in
this small, essentially South American genus of
aposematic species for over 150 years, since Westwood
(1851) described S. magdalena; it is the first known
Stalachtis species to apparently occur exclusively in
montane habitats; and its wing pattern differs
substantially from that of its congeners. I herein
describe this new Stalachtis species and attempt to
establish its phylogenetic position within the genus by
informally constructing a preliminary hypothesis of
phylogenetic relationships for Stalachtis and proposing
a new species-group classification.
Stalachtis halloweeni Hall, new species
(Figs. 1-2; 3: 4)
Description: MALE: Forewing length 29.5 mm. Forewing
elongate, costal and distal margins approximately straight, four
forewing radial veins, discal cell elongate; hindwing rounded and
slightly elongate, hindwing veins Rs and M, stalked. Dorsal surface:
Forewing round color black, discal cell orange with some black
scaling medially, broad orange rays in cells 2A and Cu, extending from
(Cu,) or near (2A) wing bags. to join submarginal band, orange
rectangle at base of cell Cu,, small orange spot at base of cell M,,
orange streaks in cells M, to R,., and R, immediately distal to atiseal
cell end, decreasing in size ‘from cell M, to cell R,, broad submarginal
orange band extending from costa to tornus, with an enlarged
rec tangular orange patch nearly encircling a black spot in cells M, atl
Cu,, distal fringe black; hindwing ground color black, pale or: ee ‘spot
at wing base, a broad orange streak through discal cell and cells Cu, to
M,, and along anal margin, those in cells Cu, and M, shorter ‘bara
VOLUME 60, NUMBER 3
others and that in cell Cu, joining submarginal band, small distal
orange spot in cell Sc+R,, broad and uneven submarginal orange band
extending from apex ey tornus, distal fringe black. Ventral sides
Forewing differs from dorsal surface by having a small orange spot at
costal wing base, only distal portion of orange streak present in cell
2A, orange streak in cell M. , joining submar. ginal band, orange spot in
cell R, replaced by a sparse scattering of “whitish scales; hindwing
differs by having a very narrow line of orange scaling at middle of
costal margin, with a sparse scattering of whitish scales distally, a
larger orange spot in cell Sc+R,, a much larger and darker or ange spot
at wing base, and a sparse scattering of Renin scales along anal
margin.
Head: First segment of labial palpi a mixture of black and white
scaling, segment two black with a broad white lateral band and some
white se: aling ventrally, and third segment black; eyes bare and black,
with a mixture of black and white scaling at margins; frons black, with
white scaling laterally; antennae 60% of forewing length, segments
black with w ahite Nee aling at ventral base and narrow seh | region
along inner ventral margin, clubs black.
Body: Dorsal surface of thorax black, tegula black with dark orange
scaling at base, ventral surface of thorax black with a white band
between legs and a large orange patch near base of forewing;
abdomen black dorsally, with a broad orange band laterally, and white
ventrally, with narrow black lines on either side ( (patterning virtually
indiscernible in Fig. 1 due to a covering of mold), all legs black.
Genitalia (Fig. 3): Uncus in lateral view rectangular and vertically
elongate, lateral “window” anterior to uncus very narrow, tegumen
very narrow, with a triangular ventral margin; falces extremely long
and ventrally directed, with a weakly bent “elbow”; vinculum narrow,
anteriorly bowed, broadest medially and slightly posteriorly indented
near ventral tip, with a posteriorly directed triangular section of
sclerotized tissue at ventral margin and no anterior saccus; aedeagus
narrow, convex and of medium length, gradually tapering to a slightly
upturned and finely pointed tip, vesica exits along ventral margin of
posterior third of aedeagus, cormutal patch consists of a “short
sclerotized rod with about six prominently curved and anteriorly
directed spines densely positioned along all but its anterior tip,
cornutal patch positioned about one third distance from posterior to
anterior tip of aedeagus on uneverted vesica; pedicel in lateral view
broad basally, becoming narrower in angular posterior section, pedicel
joins aedeagus about one third distance from anterior to posterior tip;
valvae in lateral view consist of a large rectangular basal section, a
narrow, posteriorly elongate and aan -tipped lower process, a slightly
broader, more rectangular and posteriorly elongate upper process,
with a posteriorly awa upwardly curved, finely pointed terminal
projection, a very broad and rounded inner process, slightly shorter
than lower process, and a pointed transtilla of medium length
extending posteriorly between pair of inner processes and across top
of aedeagus: narrow tuft of long, posteriorly directed, pale brown
setae around outer margin of genital capsule; eighth abdominal tergite
and sternite rectangular.
FEMALE: Differs from male in following ways: Forewing length
28 mm. Distal forewing margin convex. Wings: Orange on both wings
very slightly paler, medial black scaling in forewing discal cell more
prominent, several forewing postdiscal orange spots do not extend as
far proximally to cell bases, orange ray in ventral forewing cell M, does
not distally join submarginal Band
Head: Second pal Ipal segment slightly more elongate, third
segment about twice as long; nudum region on antennal segments
slightly larger.
\Genitalia (Fig. 4): Corpus bursae somewhat narrow and elongate,
with a pair of eal sclerotized, invaginated spine-like signa; ductus
bursae consists of a large, creased, hardened swelling immediately
posterior to corpus Barwa containing about four pieces of rec tangular
sclerotization, a short membranous section posteriorly, then a long and
twisted lightly sclerotized section, followed by a short, concave ventral
section ai sclerotization; membranous ductus seminalis exits ductus
bursae dorsally immediately anterior to ventral section of
sclerotization; ostium bursae in dorsal view consists of a small, round,
sclerotized entrance hole, with a broad and prominently convex band
139
Fic. 3. Male genitalia of Stalachtis halloweeni holotype in lat-
eral view. Scale bar = 1 mm.
of sclerotization dorsally curving anteriorly into an invaginated pouch
below papillae anales that is membranous except for a broad,
triangular, horizontal band of sclerotization along dorsal “roof” of
pouch (perhaps a protected resting place for extremely long male
genital falces during copulation); papillae anales proportionately small
and round; very broad, semicircular tuft of long, posteriorly directed,
pale brown setae around posterior margin of eighth tergite.
Types: Holotype ¢, GUYANA: Cuyuni- Mazaruni, Mount
Ayanganna, 1120 m, 5°22.22'N 59°57.34'W, 12-16 Oct 2002 (R.
Williams) (National Museum of Natural History, Smithsonian
Institution, Washington, DC, USA [USNM)]).
Paratypes: 1d, 12, same data as holotype. 3°, GUYANA: Cuyuni-
Mazaruni, Mount Ayanganna, 4500-5500 ft, 5°24.1'N 59°57.4'W,
13-18 Apr 1999 (S. Fratello et al.) (USNM).
No additional specimens have been located in the major museums
of Europe and North America (as listed in Hall 1999, 2005).
Etymology: The name is derived from the middle
English word halloween, in reference to the fact that
the wing pattern is composed of the traditional orange
and black colors of Halloween, and is reminiscent of a
carved pumpkin.
Systematic placement and diagnosis: Stalachtis (=
Nerias Boisduval, 1836) is the sole genus currently
treated in the tribe Stalachtini. The family-group name
was proposed by Bates (1861), as a subfamily, for an
undefined number of genera whose species possessed a
“pupa not flattened beneath, secured rigidly by the tail
in an inclined position, without girdle”, a set of
characters quickly discovered by Bates (1868: 36S)
himself not to be phylogenetically informative in the
context of the ee classification of the Riodinidae.
Stichel (1910-11) used the Stalachtini as a tribal name
for the first time (as the Stalachtidi), and followed Bates
(1868) in including only Stalachtis within it. Most
recently, Harvey (1987) defined the monotypic
Stalachtini by the presence of a tuft of long setae around
140
Fic. 4. Female genitalia of Stalachtis halloweeni paratype in
dorsal view. Scale bar = 1 mm
JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY
the posterior margin of abdominal segment eight in
males and, to a greater extent, females, and mentioned
that its species are unusual in the Riodinidae in having
hindwing veins Rs and M, stalked rather than arising
separ ately from the discal cell end (Bates 1868: Stichel
1910-11). Additional unique characters that are
universal within the Stalachtini include extremely long
and weakly bent (i.e. ventrally directed) falces, a
complex arrangement of inner valve processes, and a
long straight comb of prominently curved, anteriorly
directed spines on the aedeagal vesica in the male
genitalia (Hall unpubl. data). The new Stalachtis
species, S. halloween, all of the above
characters.
Traditionally, Stalachtis has been treated as
eight species (e.g. Stichel 1910-11,
1930-31), although some authors have listed as many as
ten species (D'Abrera 1994; Bridges 1994). However,
the genus is now justifiably recognized to contain only
six species (Callaghan & Lamas 2004). Hemming (1964,
1967) selected the type species of Hiibner's (1818)
Stalachtis to be S. phaedusa (Hiibner, [1813]). It is
worth mentioning that the name Stalachtis funereus
albulus Lathy, 1958, which occasionally appears in the
riodinid literature (e. g. Rebillard 1958; D'Abrera 1994;
Bridges 1994), seeually refers to a pericopine moth
possesses
containing
5
(Arctiidae) (Hall unpubl. data). Stichel (1910-11,
1930-31) divided Stalachtis into two sections, the
“Adiorati” for S. calliope (Linnaeus, 1758) and S.
magdalena Westwood, 1851, in one ee and S.
phlegia (Cramer, 1779) (+ S. suwsanna (Fabricius, 1787))
and S. euterpe (Linnaeus, 1758) in another subgroup,
and the “Diaphanes” for S. phaedusa (+ S. zephyritis
(Dalman, 1823)) and S. lineata (Guérin-Méneville,
[1844]). Based on a study of wing pattern and male
genitalia characters in all six Stalachtis species, an
alternative species-group classification and preliminary
hypothesis of phylogenetic relationships for the genus is
proposed here.
As S. calliope, 8. magdalena, S. euterpe, S. phaedusa
and S. lineata all share a pair of large, rounded, inner
valve processes, with an elongate, pointed transtilla
extending posteriorly between them and across the top
of the aedeagus, S. phlegia is hypothesized to be sister
to the remaining species in the genus. Stalachtis phlegia
has a much smaller pair of inner valve processes,
without the intervening transtilla, and has the least
derived wing pattern, with a full complement of white
basal, postdiscal and submarginal spots. Stalachtis
calliope and S. magdalena, like S. phlegia, but unlike
any other Stalachtis species, have a complete row of
submarginal white spots on the forewing, an orange
patch at the base of the dorsal forewing and hindwing,
VOLUME 60, NUMBER 3
and an entirely checkered black and white hindwing
fringe. Their similar wing g patterns, with mottled orange
markings at the base of the forewing and parallel orange
bands on the hindwing, shared possession of an upper
valve process that is broadest medially (instead of
basally in S. phlegia and distally in the remaining three
species), and parapatric geographic ranges strongly
suggest that they are sister species. Stalachtis euterpe, S.
phaedusa and S. lineata appear to form a monophyletic
group, as all three species share similarly positioned
white wing markings that are consistently formed into
rays reread of spots, the absence of an orange patch at
the base of the dorsal forewing and hindwing, the
absence of a complete row of submarginal white spots
on the forewing, largely black wing fringes, and an
upper valve process that is broadest distally. As
Stalachtis phaedusa and S. lineata both have elongate
hyaline rays on both wings they are probably sister
species.
Stalachtis halloweeni appears to exhibit external
pattern characters that are somewhat intermediate
between those of members in the calliope and euterpe
groups. It shares with the two calliope group species the
presence of orange markings at the base of the dorsal
forewing, and wath S. magdalena the absence of a lateral
white line above as well as below the lateral orange band
on the abdomen. It shares with the three euterpe group
species a similar pattern of rays at the base of the
forewing and particularly the hindwing, even if these are
orange instead of white, the absence of a complete row
of submarginal white spots on the forewing, and black
wing fringes. Based on the above characters, and the
fact that S. halloweeni has the full complement of inner
valve processes and an upper valve process that is
broadest distally, this new species is tentatively
suggested to be the most basal member of the euterpe
group. The male genitalia of Stalachtis species are
rather homogeneous, with the most significant
interspecific variation exhibited by the upper and, to a
lesser extent, lower valve processes. Although S.
halloweeni appears to be most closely related to S.
euterpe, its male genital valvae are probably most
similar to those of S. phaedusa, but its upper valve
process does not have such a prominent ventral
protrusion at the posterior tip, and the posterior margin
extends at about a 45° angle instead of vertically.
Elsewhere in the euterpe group, S. euterpe can be
characterized by its dorsal as well as ventral swelling to
the distal tip of the upper valve process and atypically
small and straight terminal projection, and S. lineata can
be characterized by its broadly triangular instead of
narrower rod-shaped lower valve process.
In conclusion, Stalachtis seems to be best divided
14]
into three species groups, the phlegia group for S.
phlegia, the calliope group for S. calliope and S.
magdalena, and the euterpe group for S. halloweeni, S.
euterpe, S. phaedusa and S. lineata. Thus, only two of
Stichel's (1910-11, 1930-31) three proposed species
groups for Stalachtis appear to be monophyletic.
Biology: This new species appears to be restricted to
lower montane forest habitats, where it is currently
known from between about 1100 and 1700 m.
Specifically, the type series was collected in wet, low
(canopy approximately 10-15 m), evergreen high-tepui
forest, a vegetation type that was described and
illustrated by Maguire (1970), Huber et al. (1995) and
MacCulloch & Lathrop (2001).
Steve Fratello (pers. comm.) observed approximately
ten individuals of S. halloweeni on Mount Ayanganna,
most of which were probably females, judging by the
fact that all individuals captured were of that sex. Within
the forest, these individuals consistently flew at about 5
to 7 m above the ground, although two individuals were
observed flying only 2 to 3 m above the ground over a
patch of low tepui scrub at 1700 m. Individuals were
seen flying over a wide area from mid-morning to mid-
afternoon, with a rather slow, steady flight, and were not
observed alighting or resting. No other Stalachtis
species were seen flying in the same habitats as S.
halloweeni, but S. phaedusa, S. calliope and S. euterpe
have been commonly collected in neighboring lowland
areas of Guyana (Fratello pers. comm.).
Stalachtis is one of the most well known groups of
aposematic riodinids (Seitz 1916-20; D'Abrera 1994).
Given that the known caterpillars are gregarious and
aposematic (Callaghan 1986), and members of at least
some of the known foodplant families (e.g.
Simaroubaceae) contain toxic phytochemicals (e.g.
Moretti et al. 1982; Polonsky et al. 1984), it seems likely
that some or all of the Stalachtis life stages are to some
extent distasteful to predators, and pauls may thus be
predominantly Mullerian rather than Batesian mimics.
However, I am not aware of any sympatric butterflies or
moths that specifically closely resemble S. halloweeni.
Distribution: Stalachtis halloweeni is curr ently
known only from the middle slopes of Mount
Ayanganna, in the uplands of western Guyana.
However, the geographic range of this species probably
extends to neighboring highland areas in Guyana and
extreme eastern Venezuela. There continues to be
debate about whether most Guiana highland endemics
are relicts of a widespread pantepui fauna or
descendants of lowland ancestors (e.g. Myers &
Donnelly 1996; MacCulloch & Lathrop 2001). Given
that all six described Stalachtis species have entirely
lowland distributions, the ancestor of S. halloweeni
presumably colonized Mount Ayanganna from the
surrounding lowlands.
ACKNOWLEDGEMENTS
I thank The National Science Foundation (DEB # 0103746)
for financial support; G. Lamas and S. Fratello for carefully re-
viewing the manuscript; S. Fratello and R. Williams for collect-
ing and donating Stalachtis specimens to the USNM; and S.
Fratello for generously providing notes on the ecology of Sta-
lachtis halloweeni from his arduous and highly productive 1999
Mount Ayanganna expedition, which was funded in part by the
Smithsonian Biological Diversity of the Guianas Program.
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Received for publication 9 August 2005; revised and accepted 2 June
2006
VOLUME 60, NUMBER 3
ournal of the Lepidopterists’ Society
} piaop y
60(3), 2006, 143-148
A NEW SPECIES OF AURATONOTA (LEPIDOPTERA: TORTRICIDAE: CHLIDANOTINAE)
FORMERLY CONFUSED WITH A. HYDROGRAMMA (MEYRICK)
JOHN W. Brown
Systematic Entomology Laboratory, P.S.I., ‘Agricultural Research Service, U.S. Department of Agriculture
National Museum of Natural History, P.O. Box 37012, W fashington, DC 20013-7012
Email: jbrown@sel .bare.usda.gov
ABSTRACT. Auratonota pharata, new species, is described and illustrated from Costa Rica, Panama, Venezuela, and French Guiana. The
species is most similar to A. hydrogramma (Meyrick), with which it formerly was confused. It can be distinguished superficially from the latter
by the absence of the narrow pale curved band beyond the distal end of the discal cell of the forewing. The male genitalia of the new species
differ by a slightly expanded, dorsally convex, and ventrally flattened distal portion of the uncus. The female genitalia possess numerous short
curved bands of 5-6 microtrichia around a larger single seta set in a shallow pit on the surface of the middle of the papillae anales compared
with the more semicircular bands of microtrichia nearly surrounding the seta in A. hydrogramma. A survey of wing coupling in numerous gen-
era of Chlidanotini and Hilarographini revealed that the female frenulum consists of two bristles usually separated throughout their length in
all representatives examined in these two tribes; three bristles are present in females of most other Tortricidae. This character state represents
an additional putative synapomorphy uniting those two tribes.
RESUMEN. Auratonota pharata, especie nueva, es descrita e ilustrada de Costa Rica, Panama, Venezuela y Guyana Francesa. Este especie
es mas similar a-A. hydrogramma (Meyrick), con la cual ha sido con tundida. Puede ser identificada de una manera ease de esta ultima
por la ausencia de una banda curva fina clara detras de la portedistal de la celda discal de las alas anteriores. La genitalia del macho de la nueva
especie puede ser distinguida por la presencia en la partediscal del uncus de una area ligeramente expandida, convexa dorsalmente y plana ven-
tralmente. La genitalia de la hembra puede ser distinguida por la presencia de numerosas bandas curvas cortas de 5-6 espinas pequenas arriba
de una sola seta en media superficie de los papillae anales en comparaci6n con bandas mas semicirculares de espinas pequenas casi rodeando la
seta en A. hydrogramma.
Additional key words: Systematics, genitalia, morphology, Costa Rica, Central America, Venezuela, French Guiana, inventory, Chlidan-
otini, frenulum
Auratonota Razowski is the largest and most diverse
genus in Chlidanotini (Tortricidae: Chlidanotinae) with
30 described species (Razowski & Becker 2000, Brown
2005) and numerous undescribed species present in
collections. The genus is restricted to the New World
tropics, ranging from Mexico and the Caribbean (Cuba,
Dominica) south through Brazil.
A previously undescribed species of Awratonota has
been concealed in entomological collections for many
years under the name A. hydrogramma (Meyrick). The
similarity of the new species to A. hydrogramma in size,
forewing pattern, and genitalia, along with their
geographic sympatry, have combined to inhibit their
recognition. The two species can be separated by a
subtle feature of the forewing pattern, but recently
discovered features of the male and female genitalia
provide convincing evidence that they are indeed
distinct and diagnosable. The purpose of this paper is to
name, describe, and illustrate the new species. This
work was stimulated, in part, by the desire to associate
scientific names with morpho-species collected during
the NSF-funded ALAS (Arthropods of La Selva, Costa
Rica) project in order to more easily discuss differences
and similarities among the tortricid faunas of transect
sites (at different elevations) sampled over the course of
the project (1993-2005).
Dissection methodology followed that presented in
Brown and Powell (1991). Images of adults and genitalia
were captured using a Microptics digital camera system
and enhanced using Adobe Photoshop and Illustrator
software. Terminology for genital structures follows
Horak (1984). The following Hreuntionsl abbreviations
are used for the denotes of specimens examined:
AMNH = American Museum of Natural History, New
York, New York, U.S.A.; BMNH = The Natural History
Museum, London, United Kingdom; INBio = Instituto
Nacional de Biodiversidad, Santo Domingo de Heredia,
Costa Rica; UCB = Essig Museum of Entomology,
University of California, Berkeley, USA; and USNM =
National Museum of Natural History, Washington,
DAG WESFA.
Auratonota pera Brown, new species
(Figs. 2, 4, 5)
Diagnosis. Among Hered species of Auratonota,
A. pharata is most similar to A. hydrogramma in
forewing pattern, size, and genitalia. However, the latter
is superficially distinguishable from all described
congeners, including A. pharata, by the presence of a
slender, pale, arched fascia in the subterminal region of
the forewing that roughly parallels the apical half of the
termen, intersecting the costa subapically (Fig. 1); the
fascia is lacking in A. pharata (Fig. 2). The male
genitalia of A. pharata can be diganeaished from those
of A. hydrogramma by the shape of the distal one-
fourth of the uncus: attenuate and apically pointed in A.
144
hydrogramma (Fig. 3); slightly broadened, convex
dorsally, and flattened ventrally in A. pharata (Fig. 4).
The female genitalia are extremely similar in both but
can be distinguished by a subtle feature of the papillae
anales. In A. pharata microtrichia on the internal,
middle portion of the papillae anales are arranged in
short, slightly curved bands of 5-6 immediately dorso-
posterad of a small, pale-colored pit from which arises a
single larger seta (Fig. 5). In A. hydrogramma
microtrichia are arranged in circular or semicircular
bands of 8-9 nearly surrounding a more rounded pit
(Fig. 6). Awratonota pharata also is similar superficially
to A. aporema (Dognin), described from Colombia, but
has a considerably smaller forewing length (mean 11.9
mm for pharata vs. 15.9 for aporema) and lacks the
yellow-gold scaling a the forewing pattern elements of
A. aporema; the latter feature is more characteristic of
members of Psewdocomotis Brown (Chlidanotini), with
which A. aporema is more similar superficially.
Description. Head: Vertex rough scaled, mostly pale cream with a
few pale cream-brown scales; frons smooth scaled, pale cream white;
labial paplus relatively slender, short, length (all segments combined)
ca. 1.2 times horizontal diameter of compound eye, brown externally,
pale cream on inner surface; antenna thickened, with setae extremely
short, inconspicuous (typical of Chlidanotini). Thorax: Dorsum
clothed in reddish-brown scales, anterior and posterior regions with
considerable scattered white and pale brown scales; tegula pale
brown, with variably expanded patch of long scales posteriorly,
frequently expressed as an erect scale patch in both sexes. Legs
unmodified; no hairpencil or secondary scale patches in tl
Forewing ee 11.5-12.9 mm (&% = 11.9; n = 10) in male, 12.2-14.1
mm (x = 13.1; n = 2) in female; costa nearly straight, apex obtuse,
termen slightly convex, rather oblique; ground color ferruginous, with
light silvery-gray reticulations formed by irregular interrupted streaks
on veins anal a series of indistinct transverse fascia crossing them,
scattered with iridescent green scales in interspaces (the green scales
inconspicuous on flight worn specimens); basal portion from near base
to ca. 0.66 disenes t to apex largely suffused with blackish brown
between the reticulations; a small, ill-defined, irregularly-shaped,
ferruginous spot near distal end of discal cell bordered basally by a
narrow bluish silvery-gray line; termen uniform red-brown; fringe
olive-ferruginous [lacking i in most specimens examined]. Hindwing
uniform dark brown, fringe concolorous; female frenulum with two
bristles separated throughout their length. Abdomen: Dark brown.
Male genitalia (Fig. 4; image of JWB slide 806, Costa Rica; n = 6) with
uncus strong, long, mostly rod-shaped, slightly broader at base, curved
near middle, slightly expanded in distal 0.25 with dorsum convex and
venter flattened or weakly concave; socius moderately short, broad,
subrectangular, clothed in long, fine scales; hami long, ca. 0.75 times
length of uncus, weakly attenuate from base to tip, distal 0.1 bent
dorsally; gnathos extremely reduced, lateral arms membranous, mesal
portion inconspicuous; valva large, broad, expanding distally, distal
0.75 covered with fine, long setae, costa sclerotized; transtilla a simple,
narrow band; juxta a broad, shield-like plate; saccus well developed,
narrow, attenuate distally; aedeagus short, stout, mostly straight,
slightly curved at phi allobase, a tiny scobinate patch of small setae on
each side near distal end, vesica with small linear patch of
sclerotization. Female genitalia (Fig. 5; image of USNM slide 95264,
Panama; n = 3) with papillae anclieg large, bearing numerous tiny,
weakly curved lateral bands of spines in adele portion, which are
situated immediately dorso-posterad of a small pale-colored pit from
which a single seta arises; perimeter and basal portions of papillae
anales with much larger setae arising from elongate, warty bases;
sterigma simple, w eakly sclerotized, ventral posterior edge of ostium
JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY
with narrow row of 5-6 long setae on each side; ductus bursae
broadest at ostium, narrowed at about 0.2 distance from ostium to
corpus bursae, then nearly uniform in width to junction with corpus
bursae; corpus bursae, large, pear-shaped sac, junction with ductus
bursae slightly anterior of posteriormost end, signum a patch of 30-35
slender, slightly curved spines originating from sclerotized plate near
middle of corpus; ductus se vantacils from posteriormost end of corpus
bursae near junction of corpus bursae and ductus bursae; a frail
accessory bursae from a long, slender ductus originating just anterad
of signum. ;
Holotype. Male: Costa Rica, Estacion Biologica La Selva, Puerto
Viejo de Sarapiqui, 50-150 m, 10°26'N, 84°01W, 7 Feb 2002, Wagner,
Rota & Kawahara (INBio).
Paratypes (24d, 3°). BRITISH GUIANA: Potaro River,
Anundubaru, 2000', Jan 1928 (1d) (AMNH). COSTA RICA: Heredia
Province: pace Biologica La Selva, 10°26'N, 84°01'W, 50-150 m,
Jan 1998 (12), INBio- OET, J. Powell (UCB), 8-25 Mar 1999 (1¢),
22-31 Mar at (12), Wagner & Rota, 7 Feb 2002 (1¢), Wagner, Rota
& Kawahara (INBio); Ciebo, 11 km ESE La Virgen, 250-350 m,
10°21'N, 84°01'W, 18 Mar 2004 (1d) (INBio); 10 tan SE La MESES
450-550 m, LO 20N, 84 05W, 17 Mar 2003 (2¢), 19 Mar 2003 (1d ) 29
Mar 2003 (1d), INBio-OET-ALAS transect (IN Bio). ae See
Cerro Tortuguero, ae Tortuguero, 0-120 m, Oct 1989 (1<), J. Solano
(INBio), Jul 1991 (1¢), J. Solano (INBio), Jul 1993 (1¢) Rh Delgado
(INBio); Sector aes Cocori, Finca de E. Rojas, 150 m, Aug 1991
(2d), E. Rojas (INBio). Unknown Province: Carchi [possibly Sarchi,
Alajuela Province], [no date] (1¢), Wm. Schaus (USNM). FRENCH
GUIANA: St. Jean, Maroni, [no date] (2¢), Wm. Schaus (USNM); Rio
Maroni, [no date] (1d), Le M[oult] (USNM). PANAMA: Canal Zone,
Barro Colorado Island, 17 Sep 1941 (19), J. Zetek (USNM).
VENEZUELA: Aragua: Rancho Grande, 1100 m, 16-23 Oct 1966
(3d), 24-31 Oct 1966 (1d), 1-5 Nov 1966 (1d), S. S. & W. D.
Duckworth (USNM), 15-21 Jun 1967 (1d), 22-31 Aug 1967 (1¢), R.
W. Poole (USNM).
Distribution and Biology. Awratonota pharata is
recorded from Costa Rica, Panama, Venezuela, British
Guiana, and French Guiana. Although it appears to be a
species of the lowlands (i.e., below 500 m), it has been
recorded on several occasions at Rancho Grande,
Venezuela, at 1100 m and once in British Guiana from
660 m. During a multi-year survey in Costa Rica along
an elevation transect, specimens were collected at
50-150 m (n = 4), 250-350 m (n = 1), and 450-550 m (n
= 4), with none recorded from 1000, 1500, or 2000 m.
Adults have been recorded in January through March,
and June through November.
A female collected by J. Powell in January 1998 at La
Selva, Costa Rica was confined in a plastic vial where it
laid several huge, bulky eggs, 3.3 x 2.8 mm, ca. 10 times
the size of comparable-sized tortricine females (J.
Powell, pers. comm.). First instars likewise were large,
2.83.0 mm in length. Various leaves, synthetic diet, and
raw carrot were offered to the larva; the last has been
used successfully with other “borers” such as
Hepialidae. The only feeding was by one larva on the
carrot, and it ceased to feed before reaching second
instar.
Etymology. The specific epithet is from the word
“pharate,” meaning cloaked or hidden.
Auratonota hydrogramma (Meyrick, 1912)
VOLUME 60, NUMBER 3
(Figs. 1, 3, 6)
Cnephasia hydrogramma Meyrick, 1912: 683.
Eulia hydrogramma: Clarke, 1958: 128.
Auratonota hydrogramma: Razowski, 1987: 62; Brown,
1990: 156: Powell et al. 1995: 151; Razowski &
Becker, 2000: 1151; Brown, 2005: 144.
Auratonota hydrogramma was described from a
single specimen from Dutch Guiana (= Surinam)
(BMNH). The adult and male genitalia of the holotype
are figured by Clarke (1958); Razowski & Becker (2000)
provided a color illustration of an adult and line
drawings of the male and female genitalia. Based on
material in several museum collections (AMNH,
BMNH, INBio, USNM), it has been recorded from
Costa Rica, Panama, Colombia, French Guiana, British
Guiana, and Ecuador and at many of the same localities
as A. pharata (e.g., Costa Rica, Estacion Biologica La
Selva; Panama, Barro Colorado Island; French Guiana,
St. Jean, Rio Maroni). Razowski & Becker (2000)
reported it from Brazil. As is the case in A. pharata, A.
hydrogramma appears to be a species of the lowlands,
rarely collected above 600 m. During the multiple-year
transect surveys of the ALAS project in Costa Rica
(1993-2005), A. hydrogramma was collected only at the
50-150 m elevation site.
Auratonota hydrogramma can be distinguished
superficially from all other congeners by the presence of
a narrow, curved, pale fascia in the apical region of the
forewing. The male genitalia (Fig. 3, image of USNM
slide 84889, Panama; n = 6) are slightly smaller and
have a slightly broader base of the saccus than those of
A. pharata; but the most conspicuous feature that
distinguishes the two species is the pointed tip of the
uncus of A. hydrogramma. The papillae anales (Fig. 6,
image of JWB slide 745, Costa Rica; n = 2) of A.
Iu jd Onno also have the tiny curved bands of
microtrichia described above for A. pharata, but in A.
hydrogramma the bands are more semicircular and
nearly surround a rounded, pale-colored pit. Even
though only two females of each species were examined,
these differences appear to be consistent.
Holotype ¢, Dutch Guiana [Surinam], Paramaribo, Aug 1892
(BMNH).
Additional specimens examined: BRITISH GUIANA: Bartica
District, Kartabo, 2 Apr 1926 (1d) (AMNH). Seon Rk
Valle, Anchicaya, 600 m, 76° 53'W, 3° 33'N, 20-24 Jan 1992 (1d), J. B.
Sullivan (USNM). COSTA RICA: Cartago Province: le ie =Il7/
Mar 1965 (12), S. S. & W. D. Duckworth (USNM). Heredia Province:
Finca La Selva, Puerto Viejo de Sarapiqui, 50 m, 6-9 Mar 1985 (1¢),
D. Janzen & W. Hallwachs (USNM); La Selva Biological Station,
10°26'N, 84°01'W, Jan 1998, INBio-OET, J. Powell (UCB),18 Feb
2003 (12), 27 Feb 2003 (1d), 28 Feb 2003 (1d), D. Wagner (INBio);
Estacion Magassay, P.N. Braulio Carrillo, 200 m, Feb 1991 (19), M.
Barrelier. Lim6n Province: Rio Sardinas, R.N.F.S. Barra del Colorado,
18-29 Feb 1993 (1d), F. Araya; 30 km N Cariari, Sector Cocori, 100
m, Dec 1993 (1¢), Nov 1993 (1d), E. Rojas; Finca de E. Rojas, Sector
Cerro Cocori, 150 m, Sep 1993 (1d), Apr 1991 (1d), Aug 1991 (1¢), E.
Rojas; Cerro Tortuguero, P.N. Tortuguero, 100 m, Apr 1959 (1 5), R.
Aguilar & J. Solano, Jan 1993 (1d), R. Delgado. Puntarenas Province:
Estacion Esquinas, Peninsula de Oso, 0-200 a Feb 1993 (42), Sep
1993 (2d), Oct 1993 (1d), Aug 1993, ee, May 1993 (1d), J.
Quesada, Feb 1993 (20), Mar 1994 (1¢ AS fesen 1d) M. Segura,
Jan 1993 (16), G. Fonseca; Est. sae Peninsula de Osa, 200 m,
Aug 1993 (1d), J. Quesada (INBio); pe Esquinas, Peninsula de
Oso, 200 m, Jan 1993 (1d), Apr 1993 (2¢), J. Quesada, Mar 1994 (1d),
M. Segura; Albergue Cerro de Oro, 150 m, 30 Aug 1995 (1d), L.
Angulo; Saat Sirena, P.N. Corcovado, i" Aw m, Jan 1993 (1¢),
Nov 1989 ( ), Jul 1991 (1d), Jun 1990 ( oa 1993 (1d), G.
Fonseca, 1— : Aug 1980 (1c), 10-12 Aug 1980 (2d), 5-11 Jan 1981
(1d), D. Janzen & W. Hallwachs, Aug 1991 (2 23), J (G ana Golfito,
R.V.S. Golfito, Sector El Tajo, 15 May a (1d), M. Moraga; eee
Quemado, Peninsula de Oso, 200 m, Oct 1991 (1<), Oct 1991 (
Nov 1990 (1¢), F. Quesada; Cerro de Oro, 200 m, 26-30 May Tape
(1d), a ane (all INBio). Unknown Province: Sixola River, [no
date] (1d) (USNM). ECUADOR: Pichincha, Tinalandia, 16 km E
Santo oan de los Colorados, 600 m, 5-11 May 1990 (1c), R.
Leuschner (USNM). FRENCH GUIANA: St. Jean, Maton [no date]
(43), Wm. Schaus (USNM), [no month] 1926 ( Aree)
LeM[oult] (USNM); Piste de la Montagne des Singes, km 10, 5°05'
52°42'W, 8 ue 1985 (1¢), J.-F. Landry (USNM): iendehere cea
[no date] (1), Collection Le Moult (USNM); Saint- -Jean du Maroni,
[no date] ao Janvier (USNM). PANAMA: Canal Zone: Barro
Colorado Island, 1-9 May 1964 (5<), ie i Mar 1965 (23), W.D. & S.
S. Duckworth (USNM), 11 Mar 1941 (1¢), J. Zetek, [no date] (<), J.
Zetek (USNM), 21 Mar 1933 (1¢) ane 19-22 Jul 1951 (1s), R.
M. Laughlin (AMNH), 14 Feb 1936 (1c) (AMNH); Navy Res. nr.
Gamboa, 29 Mar 1965 (1d), S. S. & W. D. Duckworth (USNM).
DISCUSSION
In most tortricids, the setae of the papillae anales
arise from variably sized, papillose protuberances. In A.
hydrogramma and A. pharata these protuberances are
present only around the perimeter of the papillae
anales, with most of the papillae anales bearing short,
thin setae from weakly depressed pits bordered by a
straight or curved row of microtrichia. This unusual
arrangement of setae also is present in A. petalocrossa
and is suspected to occur in A. aporema—these four
species are all similar in size, forewing markings, and
structures of the male genitalia. These features are
easily observed using a dissecting microscope because
of the large size of the moths. In Auratonota dominica
Brown there is a similar configuration of setae but at a
much smaller scale, requiring observation using a
compound scope. Similar arrangements of setae appear
to be lacking in other Chlidanotini genera examined,
leading to the possibility that it is a feature unique to
Auratonota.
The structure of the frenulum in the female of A.
pharata, with two distinct bristles clearly separated at
their base, is somewhat unusual in Tortricidae where
the female frenulum typically consists of three bristles,
usually coalesced basally. A two-bristled frenulum was
hypothesized by Komai (1999) to represent a
syanpomorphy for Shee ra Herrich-Schaffer and
146 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY
2.0 mm
Fics. 1-2. Adults of Auratonota. 1, A. hydrogramma, 2, A. pharata.
Fics. 3-4. Male genitalia of Awratonota, with valve spread and aedeagus remove (inset of uncus). 3, A. hydrogramma, 4, A. pharata.
VOLUME 60, NUMBER 3 147
2.0 mm
Fics. 5-6. Female genitalia of Auratonota, with inset of details of papillae anales. 5, A. pharata, 6, A. hydrogramma.
148
Andrioplecta Obraztsov (Grapholitini), and more
recently Brown and Baixeras (2006) discussed _ its
distribution among species of several genera of
Grapholitini. However, to my knowledge it previously
has not been reported in Chlidanotinae. A survey of
various (but not all) genera within that subfamily
revealed that all Hilarographini and Chlidanotini
examined have a female frenulum that consists of two
bristles, potentially representing an additional
synapomorphy for that putative sister-group pair. Its
distribution is less consistent within Polyorthini where
females of Polyortha Dognin, Ardeutica Meyrick,
Lopharcha Diakonoff, and Cnephasitis Razowski have
two bristles and those of Isotrias Meyrick, Olindia
Guenée, Chlorortha Razowski, Ebodina Diakonoff, and
Lypothora Razowski have three; it is variable among
females of Histura Razowski.
ACKNOWLEDGEMENTS
I thank the following for allowing me to examine material in
their care: Eugenie Phillips (formerly INBio), Jerry Powell
(UCB), Kevin Tuck (BMNH), and Randall Schuh (AMNH). I
thank Marie Metz, USDA, Systematic Entomology Laboratory,
Washington, D.C., for preparing the illustrations and plates. The
following provided helpful reviews of the manuscript: Ronald
Ochoa, USDA, Systematic Entomology Laboratory, Beltsville,
Laryland; Thomas Henry, USDA, Systematic Entomology Lab-
oratory, National Museum of Natural History, Washington,
D.C., USA; Jézef Razowski, Polish Academy of Sciences, Insti-
tute of Systematic Zoology, Krakow, Poland; Richard Brown,
Mississippi State University, Mississippi State, USA; and Jerry
Powell, University of California, Berkeley, California, USA.
Field work in Costa Rica was supported by NSF grant
LTLSI/ALAS IV, a long-term, large-scale inventory of rainforest
arthropods, through John Longino, Evergreen State University,
Olympia, Washington.
JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY
LITERATURE CITED
Brown, J. W. 1990. New species and first U.S. record of Auratonota
(Lepidoptera: Tortricidae). Florida Entomologist 73: 153-157.
Brown, J. W. 2005. World Catalogue of Insects 5. Lepidoptera: Tortri-
cidae. Apollo Books. 741 pp.
Brown, J. W. & J. Baixeras. 2006. Macrocydia divergens, a new genus
and species of Grapholitini (Lepidoptera: Tortricidae: Olethreuti-
nae) from Central America. Zootaxa 1200: 1-6.
Brown, J. W. & J. A. Powell. 1991. Systematics of the Chrysoxena
group of genera (Lepidoptera: Tortricidae: Euliini). University of
California Publication Entomology 111. 87 pp. + figs.
Clarke, J. F. G. 1958. Catalogue of the Type Specimens of Microlepi-
doptera in the British Museum (Natural History) Described by
Edward Meyrick, Volume 3. Trustees of the British Museum,
London. 600 pp.
Horak, M. 1984. Assessment of taxonomically significant structures
in Tortricinae (Lep., Tortricidae). Mitteilungen der Schweiz-
erischen Entomologischen Gesellschaft 57: 3 64.
Komai, F. 1999. A taxonomic review of the genus Grapholita and
allied genera (Lepidoptera: Tortricidae) in the Palaearctic region.
Entomologica Scandinavica Supplement 55: 1-226.
Meyrick, E. 1912. Descriptions of South American Microlepidoptera.
Transactions of the Entomological Society of London 1911:
673-718. ;
Powell, J. A., J. Razowski, and J. W. Brown. 1995. Tortricidae: Tortric-
inae, Chlidanotinae, pp. 138-151. In: Heppner, J. B. (ed.), Atlas
of Neotropical Lepidoptera, Checklist Part I: Hyblaeoidea —
Pyraloidea — Tortricoidea. Association for Tropical Lepidoptera,
Scientific Publishers, Gainesville, Florida.
Razowski, J. 1987. Neotropical Chlidanotini (Lepidoptera: Tortrici-
dae). Bulletin of the Polish Academy of Sciences, Biological Sci-
ences 35: 61-71.
Razowski, J. and V. O. Becker. 2000. A review of the New World
Chlidanotini (Lepidoptera: Tortricidae). Revista Brasileira de Zo-
ologia 16 (1999): 1149-1182.
Received for publication 8 September 2005; revised and accepted 13
April 2006
VOLUME 60, NUMBER 3
Journal of the Lepidopterists’ Society
60(3), 2006, 149-155
149
A REVISION OF PSOLOPTERA BUTLER, INCLUDING A REDESCRIPTION OF ITS KNOWN SPECIES
(ARCTIIDAE: ARCTIINAE: EUCHROMIINI)
REBECCA B. SIMMONS
Dept. of Biology, University of North Dakota, P. O. Box 9019, Grand Forks, ND 58202, rebecca.simmons@und.nodak.edu
ABSTRACT. Psoloptera Butler, a genus of three species within Euchromiini, was previously described based on wing venation and overall
appearance, resulting in a polyphyletic assemblage. Here, species of Psoloptera are redescribed and illustrated. One species, P. aurifera (Her-
rich-Schaffer)
, is transferred to Calonotos Hiibner as a new combination. Comparisons of male and female genitalia of Psoloptera with other
historically associated genera refute previous hypotheses on euchromiine relationships.
Additional key words: Neotropical fauna, taxonomy, mimicry
The tribe Euchromiini (Lepidoptera: Arctiidae:
Arctiinae) consists of highly specialized mimics of
Hymenoptera, Diptera, and Coleoptera. This
specialized mimicry has confounded efforts to classify
these animals; distantly related species may mimic the
same model, resulting in a convergence in appearance.
Historically, classifications were formed based on overall
wing venation and similar coloration. This practice often
results in polyphyletic genera. One such genus is
Psoloptera Butler (Figs. 1-3).
Butler (1876) erected Psoloptera, and originally
placed two species in the genus, P. thoracia (Walker)
and P. leucosticta (Hiibner). Later, Schaus (1894)
described P. basifulva, citing that it was allied to P.
thoracia (Walker), the type of Psoloptera. Hampson
(1898) placed P aurifera (Herrich-Schiffer) in
Psoloptera because of similar overall appearance and
wing venation.
Butler (1876) placed Psoloptera in the subfamily
Euchromiinae (= Euchromiini of Jacobson & Weller
2002), and stated that it was closely allied to Calonotos
Hiibner and Amycles Herrich-Schiffer (= Pompiliodes
Hampson 1898; = Sphecosoma Simmons & Weller in
press), but distinct because of its plumose antennae and
different hindwing venation. Hampson (1898), in his
phylogeny of the Syntominae, placed Psoloptera as
ancestral to Metaloba Hampson, and sister to Calonotos,
Chrysocale Walker, Micragyrta Butler, Mystrocneme
Herrich-Schaffer, Orcynia Walker, Paramya Druce (=
Methysia Butler; Simmons & Weller in press), Sawrita
Herrich-Schaffer, and Scena Walker. Forbes (1939a)
allied Psoloptera to Saurita based on wing venation, and
noted that the genera differed in the beanehine of the
radial veins. Though a consensus of these views
indicates a close relationship with Calonotos, this
assertion has never been examined using any characters
other than external features.
This paper redescribes Psoloptera and revises its
composition by transferring one species to Calonotos.
The individual species are diagnosed and figured. The
genitalia of both sexes of all Psoloptera species are
figured for the first time here as well. Relationships of
Psoloptera and other euchromiines are discussed.
3
Fic. 1. Male P. basifulva Schaus, type specimen (USNM).
Fic. 2. Adult P leucosticta (Hiibner), lectotype specimen
(USNM). Fic. 3. Male P pei ia (Walker), type specimen
(BMNB).
MATERIALS AND METHODS
Standard genitalic and whole-body dissections were
performed (Winter 2000). Bodies were softened in
warm 10% KOH for 5-15 minutes and then cleaned
(scales and viscera removed) in several rinses of 30-40%
150
ethanol. Structures were stained with chlorazole black E
(Sigma, St. Louis, MO) dissolved in distilled water
(saturated). Specimens were viewed in 30-40% ethanol.
Wings were bleached and then neutralized with dilute
acetic acid. They were then rinsed in distilled water, and
stained overnight with Eosin Y (1% in distilled water;
Fisher Scientific, Pittsburgh, PA).
mounts of wings, abdominal pelts, appendages, genitalia
Permanent slide
and thoraces were made with Euparol (Bioquip, Rancho
Dominguez, CA).
Male-female pairs of each species of Psoloptera
(sensu Zerny 1912) were dissected to describe genitalia.
These species included P. thoracia (Walker), P. awrifera
(Herrich-Schiffer), P basifulvua Schaus, and P.
leucosticta Hiibner. Camera lucida drawings were made
from these preparations. Forewing measurements were
made from specimens representing each species of
Psoloptera. Type specimens for P. aurifera (BMNH), P.
basifuloa (USNM), and P. thoracia (BMNH) were
examined to verify species identifications.
Genera previously associated with Psoloptera were
examined to identify putative sister taxa. Male and
female genitalia were examined for Calonotos phlegmon
(Came: r), Chrysocale principalis (Walker), Metaloba
argante (Druce), Methysia notabilis (Walker), Orcynia
calcarata Walker, Saurita cassandra (L.), Scena styx
(Walker), and Sphecosoma aliena (Walker)
Terminology for abdominal and genital morphology
follows Klots (1970) and Forbes (1939b). Collections
consulted include BMNH, the Natural History
Museum, London (M. Scoble), and NMNH, National
Museum of Natural History, Smithsonian Institution,
Washington, D.C. (D. Harvey).
RESULTS AND DISCUSSION
Psoloptera basifulva, P.
and P. thoracia are medium-sized black
moths with lateral white spots on the first abdominal
Species
leucosticta,
composition.
segment, which may mimic a wasp waist (Weller et al.
2000). Upon examination of Psoloptera species sensu
Hampson, I found that P. basifulva, P. leucosticta, and P.
thoracia can be distinguished from P. aurifera by both
genitalic and nongenitalic characters. Psoloptera
aurifera is colored metallic green and is larger than the
other species of Psoloptera. Further, P. aurifera males
have an enlarged juxta bearing large spines, bilobed
valves, and a trilobed uncus (Fig. 4). These characters
are not shared with the other members of Psoloptera
(see following species descriptions), but are diagnostic
for Calonotos males. Based on these observations, I
propose the following change:
P. aurifera (Herrich-Schiffer), 1854 = Calonotos
aurifera (Herrich-Schiaffer), new combination
JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY
Phylogenetic placement. In Euchromiini, male
genitalia are useful for assigning species to genera,
grouping genera, and for verification of tribal affiliation.
Psoloptera shares genital features with some other
euchromiines: projections on the tegumen, slightly
bilobed valves, and enlarged juxta. C alonotos phlegmon,
Chrysocale principalis, and Metaloba argante share an
enlarged juxta with spines, as in Fig. 4. This character is
also present in Macrocneme Hiibner and certainly
represents a synapomorphy for this group of genera
(Deitz 1994). Methysia notabilis appears to have highly
specialized genitalia with trilobed valves and a spirelike
juxta (figured i in Simmons & Weller in press). Males of
Saurita cassandra, Scena styx, and Sphecosoma aliena
do not have genitalia similar to Psoloptera males (not
figured). The valves of Orcynia are bilobed, while those
of Psoloptera are unilobed; however, Orcynia calcarata
has projections on the tegumen similar to those of
Psoloptera and its juxta is slightly butterfly-shaped.
Fic. 4. Calonotos aurifera (Herrich-Schiiffer), male genitalia.
J = juxta, U = uncus, V = valve.
As in other Lepidoptera, the female genitalia of
Euchromiini tend to be more conserved than those of
the males. Psoloptera females have genitalia with two
signa placed opposite each other on the corpus bursae
and an accessory bursa originating from the ductus
bursae (Figs. 8, 11, 14). These traits are shared with
other euchromiines examined here including Calonotos
phlegmon, Chrysocale principalis, Metaloba argante,
and Saurita cassandra. Based on these observations,
Butler's (1876) placement of Psoloptera in Euchromiini
VOLUME 60, NUMBER 3
is justified. Metaloba seems to be related to Calonotos,
Chrysocale, and Macrocneme because of the presence
of a knob-shaped projection on the antevagellar plate,
which is not found in Psoloptera. The sister taxon of
Psoloptera is probably not Saurita, as proposed by
Forbes (1939a) or Calonotos (Butler 1876, Hampson
1898). Female Orcynia possess a ridged bursae, like
that of P basifulva (Fig. 8), but the signa differ.
Although the male and female genitalia are not identical
to those of Psoloptera, Orcynia seems to be the most
likely candidate for the sister taxon. This relationship
remains open to further exploration with other data,
such as molecular or larval characters.
Taxonomy. Here, I redescribe Psoloptera and
provide illustrations of the habitus, wing venation, and
male and female genitalia of its three species.
Psoloptera Butler, 1876
Psoloptera Butler, 1876: 369.
Type: Euchromia thoracia Walker
original designation.
1854: 243, by
Diagnosis. This genus superficially resembles many
other euchromiine and ctenuchine genera (Figs. 1-3).
The male genitalia of Psoloptera, especially the
ornamented, V- or butterfly-shaped juxta is distinctive
within Ctenuchini and Euchromiini. The highly
sclerotized, curved uncus is also unique to oleate:
Adult habitus (Figs. 1-3). Wings entirel ty black or with scarlet at
the base. Male forewing length 12-17 mm (av erage = 14.5 mm, SD =
1.2mm, n = 20); female forewing length 12-17 mm (average = 14.9
mm, SD = 1.5 mm, n = 11). Antennae black. Ground color of head
and thorax black or red; abdomen black with paired lateral spots on
first abdominal segment.
Head and thorax. Antennae biserrate and ciliate in males; filiform
and ciliate in females; ocelli present with melanized outer ring.
Proboscis longer than head. Epiphyses and tibial spurs short and
smooth. Tarsal claws simple.
Wings. Forewing venation (Fig. 5A): Sc sinuous, extending 3/4
of the costa. R, and R, branched, arising from discal cell. R, < stalked
with R, aciisiagg closer to cell than R,_, M, arising from the cell: M,
arising from the cell medially ewe M, aia M.,,. M, arising from ihe
base of the cell. CuA, and Cua, widely separated: A, present.
Hindwing venation (Fig. 5B): Se + R, absent. Rs and M, connate.
Discal cell cross vein asyrnmetrical, V- -shaped. M,,, fused. CuA, and
CuA, stalked, branching close to wing margin. A, Ait A, present.
Abdomen. In both sexes cous stemite with shorts stubby
apodemes. Structural modifeatians for wasp waist absent (Weller et
al. 2000). Male lacking androconia.
Genitalia. Males (Figs. 6, 7, 9, 10, 12, 13): Tegumen bearing
curved projections (TP) (Figs. 6, 9, 12), one on either side of uncus;
uncus strongly sclerotized, hooked, setose. Valve (V)_ setose,
spindlelike or slightly bilobed. Juxta (J) enlarged, V- or butterfly-
shaped. Aedagus with sclerotized vesicular region (SP) (Fig. 7),
cornuti (C) or both (Fig. 10), depending on the species.
Females (Figs. 8, 11, 14): Papillae anales (PA) laterally flattened;
membrane surrounding ovipore folded; posterior apophyses (PP) long
and narrow; anterior apophyses (AP) long and narrow or reduced and
thickened; dorsal pheromone glands reduced. Seventh sternite (VII)
unmodified; antevagellar plate unmodified or bearing crescent-like,
heavily sclerotized lip: ostium bursae symmetrical or assymetrical.
Ductus bursae (DB) short, membranous. Corpus bursae (CB) with
two patches of spinose signa (S) located opposite one another.
5
Fic. 5. Wing venation of Psoloptera Butler. A. Forewing, B.
Hindwing. A = anal vein, CuA = cubital vein, M = medial vein,
R = radial vein, Rs = radial sector, Sc = subcosta, 1-5 = vein
number.
Appendix bursa (AB) from ductus bursae, irregular in shape, lacking
ornamentation. Ductus seminalis (DS) from appendix bursa.
Discussion. Species of Psoloptera are easily
distinguished from each other by habitus and male and
female genitalia. The larval host plants and stages are
not known. Tymbal organs are present, indicating that
ultrasound is utilized for either defense or intraspecific
communication; they do not appear to be sexually
dimorphic.
Psoloptera basifulva Schaus. 1894
(Figs. 1, 6, 7, 8)
Psoloptera basifulva Schaus, 1894. Proc. Zool. Soc.
Lond. 1894: 225.
Psoloptera thoracia Druce, 1884. Biol. Centr.-Am..
Het. 1: 46, preoccupied by Psoloptera thoracia Walker
(1854).
Diagnosis. Though P._ basifulva resembles P.
thoracia, P. basifulva has orange-red patches at the base
of the fore- and hindwings (Fig. 1). These patches are
lacking in P. thoracia (Fig. 3). Psoloptera basifulva also
has an orange-red thorax and head; the head and thorax
are black in P. leucosticta (Fig. 2).
Desc p gon! Medium-sized, black moths that have red heads and
thoraces (Fig. 1).
Male. Head Red; antenna black, biserrate; frons and labial palpus
black.
Thorax. Patagium, meso-, metathorax orange-red; fore, mid, and
hindlegs black with white spots on coxae and femur; tarsi black.
Wings. Forewing. Length = 14-16 mm (average = 15 mm, SD = 1
mm, n = 3). Black with basal scarlet area. Hindwing. Black, costal area
slightly hyaline, basal scarlet area present.
Abdomen. Black with white spots on the first abdominal segment.
Genitalia (Figs. 6 & 7). Tegumen heavily sclerotized, bearing
spirelike projections (TP), one on each side of uncus; uncus curved,
setose; saccus square-shaped; valve unilobed, spatulate, setose; juxta
enlarged, butterfly-shaped; base of phallus rounded; vesica with
sclerotized area.
Female. As in male, except antennae filiform and ciliate; forewing
length = 12-17 mm (average = 14.4 mm, SD = 1.7, n = 7),
Genitalia (Fig. 8). Papillae anales, posterior apophyses unmodified;
anterior apophyses greatly reduced, nublike; S7 unmodified;
antevagellar plate U-shaped, highly sclerotized; asymmetrical; ductus
bursae sclerotized tube; corpus bursae membranous, slightly ridged,
bearing two patches of signa near accessory bursa; ductus seminalis
from middle of accessory bursa.
Type material. Psoloptera basifulva Schaus. The holotype male
(USNM) is labeled: Peru; 520; not [? text unreadable]; Collection Wm
Schaus. Type locality: Peru (Fig. 1).
Psoloptera thoracia Druce. The holotype male (BMNH) is labeled:
Panama: Chiriqui; Godman-Salvin Coll. Type locality: Panama.
Specimens examined. BRAZIL: Amazones: Villa Franca
(BMNH: 2¢, 22); BRITISH HONDURAS: Punta Gorda (BMNH:
2); No Data (BMNH: 1°); COSTA RICA: Guapiles (USNM:
12); San Mateo (BMNH: 1°), Tuis (USNM: 1d); GUATEMALA:
Cayuga (USNM: 8¢, 69; BMNH: 2<, 2°); HONDURAS: Cambre
74,
JOURNAL OF THE LEPIDOPTERISTS SOCIETY
(BMNH: 3¢, 1°); MEXICO: Tabasco (BMNH: I¢, 2°): Teapa
(USNM: 1°); PANAMA: Bugaba (BMNH: 2¢); Chiriqui (BMNH:
13); NICARAGUA: No Data: Mobile 752 on a ship (USNM: 1°);
PERU: No Data (USNM: 1°); VENEZUELA: Palma Sola
(BMNH: Id); NO DATA: (USNM: 2¢).
Psoloptera leucosticta (Hiibner, 1827)
(Figs. 2,9, 10, 11)
Glaucopis leucosticta Hiibner, 1827. Samml. Exot.
Schmett. 1: t. 162.
Psoloptera leucosticta (Hiibner) Hampson 1898. Cat.
Lep. Phal. 1: 285.
Diagnosis. Psoloptera leucosticta, unlike _ its
congeners, does not have any red markings on its head
or thorax (Fig. 2).
Description. Medium-sized, black to purplish-black moths (Fig.
2).
Male. Head. Black; antenna black, biserrate and ciliate; palpus
black; vertex and frons with white spots.
Thorax. Patagium black with lateral white spot; mesothorax black;
metathorax black; fore, mid, and hind legs black with white spots on
the coxa.
Wings. Forewing. Length = 13-16 mm (average = 14.4 mm, SD =
1.1 mm, n = 8). Black, with two white spots at base. Hindwing.
7
8 See
Fic. 6. Psoloptera basifulva Schaus, male genitalia, ventral view. J = juxta, S = saccus, TP = tegumenal process, U = uncus, V =
valve. Fic. 7. Psoloptera basifulva Schaus, aedagus, lateral view. PB = phallic base, SP = sclerotized plate, V = vesica. Fic. 8.
Psoloptera basifulva Schaus, female genitalia, ventral view. VII = seventh sternite, AB = accessory bursa, AP = anterior apophysis,
AVP = antevagellar plate, CB = corpus bursae, DB = ductus bursae, DS = ductus seminalis, O = ostium, PA = papillae anales, PP
= posterior apophysis, S = signa.
VOLUME 60, NUMBER 3
Ground color black; lighter shade of black in costal area.
Abdomen. Black. Dorsal lateral white spots on first abdominal
segment. Medial white, faint patch on venter of first three abdominal
segments.
Genitalia (Figs. 9 & 10). Tegumen heavily sclerotized, bearing
spirelike projections, one on each side of uncus; uncus curved, setose;
saccus rounded; valve terminating in blunt projection, setose; juxta
elongate and narrow, V-shaped; phallus blunt at base; vesica bearing
single patch of large cornuti apically.
Female. As in male, except antennae filiform and ciliate; forewing
length = 15-16 mm (average = 15.5 mm, SD = 0.7 mm, n = 2).
Genitalia (Fig. 11). Papillae anales, posterior and anterior
apophyses unmodified; SS unmodified; antevagellar _ plate
asymmetrical, without omamentation; ductus bursae membranous;
corpus bursae membranous, bearing two patches of signa; accessory
bursa membranous, from ductus bursae; ductus seminalis from
accessory bursa.
Type material. The type of Glaucopis leucosticta Hiibner is
apparently lost. Type locality: Venezuela. The lectotype male (USNM,
here designated) is labeled: 60 m. up Maroni River, Psoloptera
leucosticta Hbn. from BM, Collection Wm Schaus. The lectotype is
designated to ensure nomenclatural stability in this genus.
Specimens examined. BRAZIL: Cayenne (USNM: 1°);
BRITISH GUIANA: Rio Potaro, Tumatumari (USNM: 42):
MEXICO: No Data (USNM: 2¢, 12). VENEZEULA: Maroni
River (USNM: 1d, 12). NO DATA (USNM: 2¢, 62).
11
Psoloptera thoracica (Walker, 1854)
(Figs. Bhs JPA AB 14)
Euchromia thoracica Walker, 1854. List Lep. Ins. Br.
Mus. 1: 243.
Psoloptera thoracica Butler, 1876. J. Linn. Soc. Lond.
Zool 12: 369.
Diagnosis. Bright scarlet coloration is restricted to
the head and thorax in P. thoracia (Fig. 3); in P.
basifulva, the base of the fore and hind wings are
orange-red, as well as the thorax and portions of the
head (Fig. 1). Psoloptera leucosticta lacks red coloration
altogether (Fig. 2).
Description. Medium-sized, black moths with red heads and
thoraces (Fig. 3).
Male. Head. Vertex black with pair of white spots; galae scarlet;
antenna black, biserrate and ciliate; palpus black.
Thorax. Patagium scarlet; mesothorax scarlet with scattered, long
black scales; metathorax scarlet; fore, mid and hind legs black with
white spots on coxae.
Wings. Forewing. Length = 12-17 mm (average = 14.6 mm, SD =
1.3 mm, n = 9). Black. Hindwing. Costal region gray, grading to black
at CuA, and CuA,,.
Abdomen. Pair of white lateral spots present on first abdominal
segment; remainder of abdomen black dorsally; medial white spot
Fic. 9. Psoloptera leucosticta (Hiibner), male genitalia, ventral view. J = juxta, TP = tegumenal process, U = uncus, V = valve.
Fic. 10. Psoloptera leucosticta (Hiibner), aedagus, lateral view. C = cornuti, PB = phallic base, V = vesica. FG. 11. Psoloptera leu-
costicta (Hiibner), female genitalia, ventral view. VII = seventh sternite, AB = accessory bursa, AP = anterior apophysis, AVP = an-
tevagellar plate, CB = corpus bursae, DB = ductus bursae, DS = ductus seminalis, PA = papillae anales, PP = posterior apophysis.
S = signa.
154
JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY
Fic. 12. Psoloptera thoracia (Walker)
13. Psoloptera thoracia (Walker),
Psoloptera thoracia (Walker)
apophysis, S = signa.
present on venter of first three abdominal segments; remainder black
ventrally.
Genitalia (Figs. 12 & 13). Tegumen rounded with elongate
spindlelike projection on either side of uncus; uncus strongly hooked,
setose; saccus rounded; valve apically narrowed, ending in rounded
projections, setose; juxta with butterfly-shaped structure, strongly
sclerotized; vesica with sclerotized plate i and medial patch of me >dium-
sized cornuti.
Female. As in male, except antennae filiform and ciliate; forewing
length = 16 mm (average = 16 mm, SD = 0, n = 2). ;
Genitalia (Fig. 14). Papillae anales, posterior and anterior
apophyses unmodified; sternite 7 unmodified; antevagellar plate with
heavily sclerotized lip, symmetrical; ductus pene membranous;
corpus bursae membranous, bearing two patches of signa; accessory
bursa large, irregular in shape, from ductus bursae; ductus seminalis
from base of accessory bursa.
Type material. The holotype male of Euchromia thoracia Walker
(BMNH) is labeled: Ega, Bates, 51-43. Type locality: Amazones
[Brazil].
Specimens examined. BRAZIL: Amazones: Fonte Boa
(BMNH: 54, 32), Pegas (BMNH: 1<), Rio Ucayla (BMNH: 2d), S. de
Villa Franca (BMNH: 12¢), S. Paulo (BMNH: 1°); Cundinmarca:
Cananche (BMNH: 1¢); Ega: (BMNH: 4¢); Humayta (BMNH: 5¢
2°); Lower Amazon and R. Madeira (BMNH: 1c); Rio Maderia:
Allianca below S. Antonio (BMNH: 1¢); San Juan: Solimoens
(BMNH: I<): 8. Paulo de Olivenca (BMNH: 4¢, 5°; USNM: 4¢):
Sao Paulo de Amazones (USNM: 2¢, 1°); Teffe (BMNH: 6¢;
USNM: 1°). COLOMBIA: Caqueta: Rio Orteguaza nr. Rio Peneya
(USNé: 1c, 12); Chiriguana District: Lake Sapatoza Region
(BMNH: 1¢); Llanos of Rio Meta: S. Martin (BMNH: 1<):
, male genitalia, ventral view. J = juxta, TP =
aedagus, lateral view. C = cornuti, PB = phallic base, SP = sclerotized plate, V = vesica. Fic. 14.
. female genitalia, ventral view. VII = seventh sternite, AB =
AVP = antevagellar plate, CB = corpus bursae, DB = ductus bursae, DS = ductus seminalis, PA = papillae anales, PP =
tegumenal process, U = uncus, V = valve. Fic.
accessory bursa, AP = anterior apophysis,
posterior
Magdalena Valley (BMNH: 1°); Ort.: Medina (USNM: 19); R.
Cantinere: Muzo (BMNH: 3°); Rio Negro (BMNH: 1¢); Villaricua
(USNM: 12). COSTA RICA: Guanacaste: Santa Rosa National Park
(BMNH: l¢, 12); Turrialba (USNM: 2c, 12). ECUADOR: Napo
Provy.: Yasuni Research Station, Rios Tivacuno & Tiputini, 76° 36' W,
0° 38' S, 250 m (USNM: 2°); Sarayacu (BMNH: 1d, 19).
Rth Or a No Data (BMNH: 19); PANAMA: Alhajuelo
(USNM: 12): Barro Colorado es (USNM: 52); Canal Zone
(USNM: 1¢): Corozal (USNM: 2c); Canno Saddle (USNM: 22); La
Chorrera (BMNH: 2°); Matachin (BMNH: 12); PortoBello
(USNM: 1d, 22); No Data (BMNH: 1¢; USNM: 1¢). PERU:
Amazones: Cavallo-Cocho (BMNH: 2°); Pebas: Loreto (BMNH:
12); Rio Udayali: Contamama (BMNH: 12); Tarapoto (BMNH: I<);
No Data (BMNH: 1d). VENEZUELA: Aroa (USNM: 6¢, 52); La
Cruces Colon. (BMNH: 22); Las Quigas: San Coe Valley (8d;
32); Las Quigas nr. San et BMNH: 3d, 22); Palma Sol
(BMNE: 1d); San Esteban (BMNH: 11¢ aleni (BMNH:
1d); Valera (USN M: 1c); No Data (BMNH: 2 25, 12). NO DATA:
(BMNH: 1d; USNM: 1d, 19).
ACKNOWLEDGEMENTS
Revisionary work is impossible without cooperation from col-
lections and associated curators. I would like to thank Dr. Don-
ald Harvey (USNM), Mr. Martin Honey (BMNH) and Dr. Mal-
colm Scoble (BMNH) for access and loans of specimens, as well
as hospitality during museum visits. I would like to thank Drs.
William Miller and Siem Weller and Ms. Michelle DaCosta for
helpful discussion and suggestions during the formation of this
manuscript. Ms. DaCosta also provided editorial aid. Funding
VOLUME 60, NUMBER 3
for associated travel was provided by ND-EPSCor and Univer-
sity of North Dakota, Dept. of Biology.
LITERATURE CITED
BuTLER, A. G. 1876. Notes on the Lepidoptera of the family Zy-
gaenidae, with descriptions of new genera and species. J. Linn.
Soc. Lond. Zool. 12: 342-407.
Dietz, R. E. 1994. Systematics and biology of the genus Macrocneme
Hiibner (Lepidoptera: Ctenuchidae). University of California
Press, Berkeley.
Forbes, W. T. M. 1939a. The Lepidoptera of Barro Colorado Island,
Panama. Bulletin of the Museum of Comparative Zoology at Har-
vard College. 85: vii + 97-322, 8 plates.
Forses, W. T. M. 1939b. The muscles of the lepidopterous male gen-
italia. Ann. Entomol. Soc. Amer. 32: 1-10.
Hampson, G. F. 1898. Syntomidae. Catalogue Lepid. Phalaenae in the
British Museum. Vol. I. Trustees of the British Museum, London.
Jacogson, N. J. & S. J. WELLER. 2002. A cladistic study of the tiger
moth family Arctiidae (Noctuoidea) based on larval and adult
morphology. Thomas Say Monograph Series, Entomol. Soc.
America.
Kxots, A. B. 1970. Lepidoptera, pp. 115-130. In Tuxen, S. L. (ed.),
Taxonomist's glossary of genitalia in insects, 2nd ed. Munksgaard.
Copenhagen.
ScHaus, W. 1894. On new species of Heterocera from Tropical
America. Proc. Zool. Soc. Lond.: 225-243.
Simmons, R. B. & S. J. WELLER. 2006. Review of the Sphecosoma
genus group using adult morphology (Lepidoptera: Arctiidae).
Thomas Say Monograph Series. Entom. Soc. America, 108 pp..
108 figs.
WALKER, F. 1854. Lepidoptera Heterocera, Part 1. List of the speci-
mens of lepidopterous insects in the collection of the British Mu-
seum. 278 pp.
WELLER, S. J., R. B. Simmons, R. Boapa, & W. E. CONNER. 2000. Ab-
dominal modifications occurring in wasp mimics of the Ctenu-
chine-Euchromiine clade (Lepidoptera: Arctiidae). Ann. Ento-
mol. Soc. Am. 93: 920-928.
Winter, W. D. 2000. Basic techniques for observing and studying
moths and butterflies, Lepidopterists' Society.
Received for publication 25 October 2005; revised and accepted 7 July
2006
156
Journal of the Lepidopterists’ Society
60(3), 2006, 156-160
JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY
FOREST TENT CATERPILLAR: MATING, OVIPOSITION, AND ADULT CONGREGATION AT TOWN
LIGHTS DURING A NORTHERN MINNESOTA OUTBREAK
WILLIAM E. MILLER
Department of Entomology, University of Minnesota, St. Paul, Minnesota 55108 email: milleOl4@umn.edu
ABSTRACT. Mating and oviposition of the forest tent caterpillar, Malacosoma disstria (Hiibner) (Lasiocampidae), were studied in two sites
~2 ha each at levels 1-5 m above the ground, and adult congregation at lights was studied in brightly lit town districts. The observations were
made near or at Ely, St. Louis Co., Minnesota, in the latter years of an outbreak peaking during 1999-2003. Descent of the nomadic larvae from
the forest canopy to less defoliated vegetative strata for feeding and cocooning later e abled mating to be observed conveniently from the
ground. Adult activity began or intensified at ~5:30 pm CDT, when males vigorously sought mates in foliage harboring cocoons. Mating pairs
were captured and caged in jars containing a host branchlet for oviposition; mean duration of copulation was 202 min. Captive mated females
always oviposited the morning after mating; mean duration of oviposition was 88 min. Dispersal behavior always preceded oviposition. Five new
oviposition hosts were recorded. Naturally deposited egg rings were found most often on branchlets oriented nearer horizontal than vertical.
Females congregating in brightly lit districts were usually gravid, but seldom oviposited.
Additional key words: Malacosoma disstria, Lasiocampidae, defoliation.
In northern Minnesota and elsewhere, the forest tent
caterpillar, Malacosoma disstria (Hiibner)
(Lasiocampidae), builds up periodically to outbreak
densities (Witter 1979). During Minnesota outbreaks,
quaking Populus tremuloides — Michx.
(Salicaceae), and most other broad-leaved trees and
shrubs except red maple, Acer rubrum L. (Aceraceae),
are severely defoliated. Red maple contains an
antifeedant (Nicol et al. 1997), but the nomadic larvae
readily spin cocoons among its leaves. The forest tent
caterpillar is univoltine, and the larvae hatch in spring at
host bud-break (Batzer and Morris 1971).
Many thousands of hectares of forest, primarily
quaking aspen, were defoliated in the outbreak that
peaked during 1999-2003 in northern Minnesota when
this study was done (Minnesota Dept. of Nat. Res. For.
Ins. Disease Newsletter 1999-2004). Although
defoliation seldom kills trees, it reduces their wood-
volume growth, and the larvae are considered a
nuisance by landowners and outdoor recreationists
(Duncan et al. 1956). The eastern tent caterpillar,
Malacosoma americanum (Fabricius), also occurs in
Minnesota (Stehr and Cook 1968), but mostly in the
eastern part of the State, and it was not seen in the
outbreak area. Fitzgerald (1995) provides an exhaustive
compendium of information for all Malacosoma.
At low, nonoutbreak densities, forest tent caterpillar
adults mate high above the ground in the forest canopy;
during outbreaks they mate on lower, understory
vegetation. Mating nearer the ground occurs because
larvae descend to less defoliated strata to find food and
cocooning sites (Batzer et al. 1995), and males
concentrate their search for mates where female
cocoons occur. The larvae require angled surfaces for
aspen,
cocooning, and they commonly create cocoon structures
by drawing and spinning leaves together (illustrated by
Batzer and Morris 1971 and Fitzgerald 1995, p. 60). In
this study, many cocoons were spun on herbaceous
plants of the ground stratum as well as shrubs. The
downward shifting later enabled mating to be
conveniently observed from the ground.
Previous reports touch on topics treated here.
Bieman and Witter (1983) described field mating
behavior at low and high levels of mate competition,
reflecting low and high population densities. Stehr and
Cook (1968) Bored that forest tent caterpillar
oviposition proceeds helically and in a layer one egg
deep. Shepherd (1979) reported daily rhythms of male
and female activity. Hodson (1941) noted adult
congregation at town lights. As elaborated in the
discussion section, the present study supplies more or
different details about these and related topics.
MATERIALS AND METHODS
This study was conducted near and in the town of Ely,
St. Louis Co., Minnesota. Mating and oviposition were
observed in two multilayered stands =40 yr old.
Overstories in these stands were dominated by quaking
aspen, but scattered examples of other tree species were
present, in descending order of abundance, balsam
poplar, Populus balsamifera L. (Salicaceae); red pine,
Pinus resinosa Ait.; jack pine, P. banksiana Lamb.;
eastern white pine, P. strobus L. (Pinaceae); white birch,
Betula papyrifera Marsh. (Betulaceae); and balsam fir,
Abies balsamea (L.) Mill. (Pinaceae). The understories
were dominated by willow, Salix sp. (Salicaceae), but
contained scattered examples of other species, in
descending order of abundance, red-osier dogwood,
VOLUME 60, NUMBER 3
Cornus stolonifera Michx. (Cornaceae); speckled alder,
Alnus rugosa (Du Roi) Spreng (Betulaceae); pin cherry,
Prunus pennsylvanica L. (Rosaceae); beaked hazel,
Corylus cornuta (Marsh.) (Betulaceae); and American
cranberrybush, Viburnum trilobum Marsh.
(Caprifoliaceae). The vertical range of observation was
1-5 m above ground, and the sites were each =2 ha.
Copulating pairs captured in late afternoon were
caged in wide-mouthed, 1-liter glass jars with gauze-
covered tops containing an aspen or willow branchlet
3-6 mm in diameter. Cages were moved in the evening
to a darkened basement with temperatures constantly at
17-19°C, which approximated cool, outdoor evening
temperatures. Cages were returned in the morning to
shaded outdoor spaces where temperatures ranged
19-26°C. Observations of copulation and oviposition
durations were made at <30-min intervals.
Gender identification of adults was based on antennal
rami, which are =2x longer in males than in females.
Males also are noticeably smaller than females (sample
measurements given later). Early in the study, active
males were netted and released after their gender was
confirmed (n> 20). Females were dissected for
verification of their gravid or spent condition. Egg
fertility was determined by the presence of embryos.
Egg numbers/ring were estimated as described by
Witter and Kulman (1969), and spumaline—the frothy
substance that covers egg rings—was removed as these
authors recommended by brushing with a discarded
toothbrush. Forewings were removed for accurate
length measurement. Angles of egg-bearing branchlets
in the field were referenced to horizontal and were
estimated with a large protractor. All clock hours refer
to Central Daylight Time. Statistics were computed
with SYSTAT ( 1992) software, except for G,,, which was
computed according to Sokal and Rohlf “C1981). The
abbreviation SD refers to standard deviation.
Mean daily temperatures during the late larval stage
were computed from daily maxima and minima
recorded at the Winton meteorological station <5 km
from Ely (Minnesota State Climatology Office 2005).
RESULTS
Mating. The first adults of the flight season were
males, as the species is protandrous. Throughout daily
adult activity, starting or intensifying ppieally= ~5:30 pm,
males visible by the hundreds in any directional view
flew vigorously and constantly in vegetative strata
containing cocoons. They flew mainly in a zone within
<0.5 m ae tree and shrub canopies, circling , Zigzagging,
alighting and crawling along branchlets before taking
flight again if they did not find an opportunity to
copulate. The number of active, searching males around
foliage that contained cocoons was ~4x that around
foliage that contained no cocoons. On windy evenings,
males confined their activity closer to canopies. During
two cool, rainy, late afternoons with ambient
temperatures <15° C, only a small fraction of the males
known to be present were active.
Many female pupae appeared to emit their calling
pheromone (Struble 1970) before they completed
eclosion, as up to 6 males often concentrated activ ity
around a single cocoon until the aia eclosed. Males
copulated with such females within 1-2 sec after the tip
of the female abdomen cleared the cocoon. The wings
of such females inflated during copulation. Most mating
pairs were found within 10 cm of a cocoon structure
presumed to be that of the female, and meconium was
often seen on such structures. Some copulating females
walked several cm as they were being observed before
capture, and because of their larger size pulled the
smaller males along. None of the females observed in
this study attempted to fly during copulation. Females
did not always begin calling before completing eclosion.
The locations of two such females found resting quietly
on cocoon structures were marked for continuing
observation during the evening. At first, captured males
released within centimeters of these females flew away.
However, by onset of darkness both females were in
copula, apparently having eventually emitted their
calling pheromone. The sexes were always positioned
end to end during copulation.
A total of 69 pairs were seen in copula during this
study, of which 48 were successfully caged for further
observation. Some copulating pairs separated on
capture, presumably because copulation had just begun
or was nearly completed. Copulating pairs were seen in
the field as soon as daily male activity commenced and
pairs known to be newly copulating were captured as
late as 9:30 pm. Neither active males nor mating pairs
were seen during early dawn hours, presumably
because of too cool temperatures. Caged pairs remained
in copula following capture for 150-255 min, averaging
202 min (n = 10), but it should be noted that beginning
and ending of copulation was not always precisely
timed, and that some durations are subject to an error of
+30 min. Copulating pairs occurred on all the trees and
shrubs named earlier, including the conifers. None of
the conifers had been fed on, but they provided
cocooning sites under conditions of extensive foodplant
defoliation. Based on forewing lengths—a proxy for
body size—there was a statistically significant relation
between gender body sizes of naturally mating pairs:
forewing length av eraged 16.8 + SD 1.42 mm for
and 12.4 + SD 1.05 mm for their
0.49, df = 23, P = 0.013).
femaleck mates
(Pearson's r =
158
Oviposition. As in other species of Malacosoma, the
forest tent caterpillar deposits its eggs in one batch
during one oviposition event unless disturbed during
the process. Of the 48 captive mated pairs, 88% of the
females began ovipositing the day after copulation, most
starting in early morning. The earliest an oviposition was
completed was 7:05 am. Other females began
ovipositing later, the latest at noon. If not ovipositing the
day after copulation, captive females failed to do so
entirely. Duration of individual ov iposition in captiv ity
monitored during daily hours of observation, which
began at 7:00 am, need 65-134 min, averaging 88 min
(n = 11). Shorter and longer oviposition durations were
associated with smaller and larger egg rings, which
imply smaller and larger females and cooler and warmer
ambient temperatures.
Just before beginning to oviposit, captive females
always exhibited dispersal behavior. For 5-15 min they
flew upward and around inside their cages. Further,
naturally deposited egg rings were never observed near
cocoon structures (n = 74). After attempting to disperse,
females became quiet, positioning themselves more or
less diagonally on the caged branchlet head up with
wings partly spread, and extended abdomenal tips
around the branchlet nearly 180° before starting to
oviposit. As they oviposited, they gradually moved
around and down the branchlet, coating eggs and the
egg ring surface with spumaline. The second tur of the
egg ring was aligned with the first so that the top margin
aries seldom haa gaps (Fig. 1d). By the sila of
had been
deposited and the females were standing on the egg ring
surface.
branchlets during oviposition were clockwise in 12 cases
(60%) and counterclockwise in 8 (40%),
reversals during the process,
oviposition, several turns of the egg ring
Directions of female progression around
Q
with only 3
2 from clockwise to
JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY
counterclockwise, and 1 the opposite. Within minutes of
completing oviposition females flew vigorously again as
before oviposition.
Whether termed 'mass' or 'ring', eggs were deposited
as a continuous helical band each turn of which was
ae fitted with the previous turn, the width of the
). When
egg ring deposition was observed closely in et
band generally diminishing downward (Fig. 1
seams in the spumaline covering between turns of the
band were usually faintly ev ident as s light depressions,
sometimes as ridges. Depending on female size,
number of eggs, and branchlet diameter, rings consisted
of up to five turns (Fig. la, c). After spumaline removal,
margins between turns were not evident. After two days
of rainy, windy weather, an increase in irregularly
deposited eggs was evident in field egg rings
presumably because oviposition had been interrupted
by such weather (Fig. le). Spumaline was more or less
clear when dispensed, but darkened after =24 h.
For egg rings deposited in captivity, number of
eggs/ring av eraged 285 + SD 112 and for those
deposited naturally, 246 + SD 66, and the difference
was not significant (Student's t = 1.20, df = 13.8,
Based on 5-39 eggs/ring
separate variances, P = 0.25). B ggs
dissected in late July
contained fertile eggs, with an average of 94% of
dissected eggs/ring fertile (n = 14). Corresponding
values for eggs collected in the field were 100 and 98%
39). These differences between captive and field
eggs were close and inferred to be nonsignificant
statistically.
Species on which naturally deposited egg rings were
found included quaking aspen (76%), willow (12%),
with the remainder (12%) on red osier dogwood, balsam
poplar, beaked hazel, speckled alder, pin cherry, and
American cranberrybush (n = 72).
and early August, 94% of rings
(n =
Fic. 1. Delineated margins of the band or bands of eggs in egg rings of the forest tent caterpillar on aspen branchlets. Delineation
is based on seams in corresponding spumaline deposition. In the absence of female disturbance, oviposition proceeds in a down-
ward helical manner.
VOLUME 60, NUMBER 3
Branchlets on which egg rings were found in the field
ranged 2-6 mm in diam. Orientation of egg-bearing
branchlets in the field tended to be nearer horizontal
than vertical. All 5 samples of 28-74 branchlets
containing naturally deposited egg rings in late July and
early August were distributed similarly in each sample
among three equally progressing angle classes. When
pooled, 50% were on branchlets angled 0—30° relative to
horizontal, 18% on those 30-60°, and 32% on those
60-90° (n = 291), with departure from a no- erence
distribution highly significant (G,,, = 42.9, df = 2, P <
0.001). The ciecrinntion aes 0-90° ane was
backward-J-shaped.
Adult congregation at lights. Like many moths, forest
tent caterpillar adults are photopositive. During flight
periods, thousands of adults congregated near
streetlights and on brightly lit exterior walls of buildings
in towns in the outlines area. In Ely, a few c congregated
males had frayed wings, indicating much flight history,
but the females showed little or no wing wear. The
source was almost certainly infested quaking aspen that
were <l km distant. Once congregated, the adults
remained notably inactive, and seemed to be present
night and day until dying or becoming prey of birds and
bats. Both genders seemed equally represented, but
copulating pairs were seldom seen. Eggs were
sometimes seen on building walls, but were scarce
relative to the numbers of females present.
Congregated females collected at various times of day
for close examination came from the 400-600 and 900
blocks of Sheridan Street in downtown Ely. Their yearly
forewing lengths during 2000-2003 averaged 20.0, 18.9,
19.1, and 18.9 mm, respectively, Sade appeared to
indicate temporally decreasing body size (F = 13.5, df =
3, 166; P < 0.001). Corresponding percentages that
were fully gravid were 96, 90, 97 and 58.
DISCUSSION
Mating and oviposition. The present study indicates
strongly that the vegetative stratum in which most
mating occurs is the one containing the most cocoons.
The high frequency with which males were observed to
copulate with females at the moment of female eclosion
is more likely to occur at high than at low population
densities, and was observed earlier in Minnesota
(Hodson 1941, Bieman and Witter 1983). Female
calling pheromone doubtless stimulated such intense
male activity. Bieman and Witter (1983) also reported
that some males lingered near empty or parasitized
cocoons, as well as brown objects, which suggests they
also use visual cues.
Typically, fernales eclosed and copulated in late
afternoon and evening, and oviposited early the next
day. The absence of early morning flight activity in this
study is not surprising for northern forest tent
caterpillar populations. Shepherd (1979) also reported
little or no male activity near dawn in Alberta and
British Columbia, and found that males ceased flying at
<11°C. Such temperatures were common near dawn in
the present study. Daily patterns of male activity, female
eclosion, and evening female calling closely match those
reported by Shepherd.
Shepherd (1979) and Bieman and Witter (1983) also
reported attempted dispersal by newly mated captive
females. Lack of proximity between naturally deposited
egg rings and cocoon structures is further evidence of
preoviposition dispersal.
Positive and significant correlation between gender
body sizes of mating pairs indicates that the sexes do not
mate randomly with respect to size. Reasons for this are
unknown, but one might speculate that larger males
out-compete smaller ones to mate with larger females.
The mean duration of copulation observed in captivity,
202 min, is near the mean duration of 197 min extracted
from Table 1 in Bieman and Witter (1983) for their high
population density.
Highly variable numbers of eggs/ring averaging 285
in captivity did not seem to differ significantly from the
similarly variable mean of 246 observed in the field, nor
did the apparent fertility means of 94% in captivity
differ significantly from the corresponding 100 and 98%
Obeenednnithe Gelder higher and lower fertilities
were recorded earlier in Minnesota (Witter and Kulman
1972). After close observation confirmed the helical
pattern of oviposition reported by Stehr and Cook
(1968) and revealed corresponding seams in spumaline
deposition, it became possible to reconstruct the course
of oviposition in detail (Fig. 1).
Of naturally deposited egg rings found at study sites,
88% were on quaking aspen ena Rvillow. The remaining
12% were on 6 lesser utilized oviposition species, of
which 5—red-osier dogwood, speckled alder, balsam
poplar, beaked hazel, andi American cranberrybush—
were absent from Fitzgerald's (1995) foodplant
tabulation for the forest tent caterpillar and other
Malacosoma. (Flowering dogwood, Cornus florida L..
an occasional foodplant of southern populations [Goyer
et al. 1987], was presumably omitted accidentally from
the tabulation.) The new oviposition host records
reported here involved no more than 2 egg rings/host,
and they are probably explainable by the presence of
these plants in areas of high forest tent caterpillar
population density.
The backward-J- shaped distribution of angles of
branchlets bearing naturally deposited egg rings 1-5 m
above the gr ound is reported here for the first time. Its
160
significance, if any, is unclear. One possibility is that
branchlets nearer horizontal than vertical—where most
egg rings were deposited—may be more abundant in
midcrowns especially of the overstory and thus provide
larvae easier access to the larger foliage volumes at
midcrown.
Adult congregation at lights. Hodson (1941) earlier
reported adult congregation at town lights where the
nearest source infestations were several kilometers
distant. In the present study, sources were sl km
distant. Such short-range attraction to lights at high
population densities does not seem surprising. Female
movement to lit areas probably occurred during the
evening after late-afternoon eclosion and mating. Most
congregated females were gravid, and while their status
with respect to mating is unknown, they seem likely to
have already mated because of the generally assiduous
male pursuit. Congregation results in little reproduction
because females do not appear to return to foodplants.
The paucity of eggs at lights may result from lack of
suitable oviposition sites (branchlets). The decreasing
yearly body sizes of congregated females during
2000-2003 may be explainable by either or both of two
effects: decreasing diet quality with repeated foodplant
defoliation, and the generally inverse relation between
body size and direction of temperature difference
during late larval development (Miller 2005). During
late larval development, 10-19 June, in 2000, when
female forewing length was greatest, daily mean
temperature averaged a cool 12.4°C, whereas in
2001-2003, when female forewing length decreased,
daily mean temperature was higher by an average of
4.4°C. Less common than adult congregation at lights is
convective transport of adults in the turbulent air of
cold fronts for hundreds of kilometers in a few hours as
Brown (1965) reported in Alberta and elsewhere. The
population dynamics significance of convective
transport is unclear as the gravid and mated statuses of
transported females have not been reported.
LITERATURE CITED
Batzer, H. O. AND R. C. Morris. 1971. Forest tent caterpillar. U. S.
Dept. Agric. For. Serv. For. Pest Leaflet 9, 5 pp.
Batzer, H. O., M. P. Martin, W. J. MaTTsON AND W. E. MILLER.
1995. The forest tent caterpillar in aspen stands: distribution and
density estimation of four life stages in four vegetation strata. For.
Sci. 41; 99-121.
BIEMAN, D.N. AND J. A. WITTER. 1983. Mating behavior of Malaco-
soma disstria at two levels of mate competition. Fla. Entomol. 66:
JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY
272-279.
Brown, C, E. 1965. Mass transport of forest tent caterpillar moths,
Malacosoma disstria Hiibner, by a cold front. Can. Entomol. 97:
1073-1075.
Duncan, D. P., A. C. Hopson, A. E. SCHNEIDER, H. BaTzeER, R.
FROELICH, D. MEYER AND C-J. SHIUE. 1956. Influence of the for-
est tent caterpillar (Malacosoma disstria Hbn.) upon the aspen
forests of Minnesota. Office of Iron Range Resources and Reha-
bilitation, St. Paul, Minnesota, 45 pp.
FITZGERALD, T. D. 1995. The tent caterpillars. Comstock, Ithaca, New
York. 303 pp.
Goyer, R. A., G. J. LENHARD, J. D. SMITH AND R. A. May. 1987. Esti-
mating the number of eggs per egg mass of the forest tent cater-
pillar, Malacosoma disstria, on three tree species in the southem
U.S. J. Entomol. Sci. 22: 188-191.
Hopson, A. C. 1941. An ecological study of the forest tent caterpillar,
Malacosoma disstria Hbn., in northern Minnesota. Univ. Minn.
Agric. Expt. Stn. Tech. Bull. 148, 55 pp.
MILLER, W. E. 2005. Extrinsic effects on fecundity-maternal weight
relations in capital-breeding Lepidoptera. J. Lepid. Soc. 59:
143-160.
MINNESOTA DEPARTMENT OF NATURAL RESOURCES. Forest Insect and
Disease Newsletter. 1999-2004. Available online at
http:/Avww.dnr.state.mn.us/fid/.
MINNESOTA STATE CLIMATOLOGY OFFICE. 2005. Closest station
climate data retrieval. Available online at
http://climate.umn.edu/HIDradius/radius.asp.
NIcoL, R. W,, J. T. ARNASON, B. HELSON AND M. M. ABou-ZaID. 1997.
Effect of host and nonhost trees on the growth and development
of the forest tent caterpillar, Malacosoma disstria (Lepidoptera:
Lasiocampidae). Can. Entomol, 129: 991-999.
SHEPHERD, R. F. 1979. Comparison of the daily cycle of adult behav-
ior of five forest Lepidoptera from western Canada, and their re-
sponse to pheromone traps. Mitt. Schweiz. Entomol. Ges. 52:
157-168.
SOKAL, R. R AND F. J. ROHLF. 1981. Biometry, ed. 2. Freeman, New
York. 859 pp.
STEHR, F. W. AND E. F. Cook. 1968. A revision of the genus Malaco-
soma Hiibner in North America (Lepidoptera: Lasiocampidae):
systematics, biology, immatures, and parasites. Smiths. Inst. Bull.
276, 321 pp.
STRUBLE, D. L. 1970. A sex pheromone in the forest tent caterpillar. J.
Econ. Entomol. 63: 295-296.
SYSTAT: statistics, version 5.2 ed. 1992. SYSTAT Inc., Evanston, Illi-
nois.
Witter, J. A. 1979. The forest tent caterpillar (Lepidoptera: Lasio-
campidae) in Minnesota: a case history review. Great Lakes En-
tomol. 12: 191-197.
Wirter, J. A. AND H. M. KuLMaN. 1969. Estimating the number of
eggs per egg mass of the forest tent caterpillar, Malacosoma diss-
tria (Lepidoptera: Lasiocampidae). Mich. Entomol. 2: 63-71.
Witter, J. A. AND H. M. KuLMAN. 1972. Mortality factors affecting
eggs of the forest tent caterpillar, Malacosoma disstria (Lepi-
doptera: Lasiocampidae). Can. Entomol. 104: 705-710.
Received for publication 20 May 2005; revised and accepted 18 July
2006
VOLUME 60, NUMBER 3
Journal of the Lepidopterists’ Society
60(3), 2006, 161-164
161
A NEW SPECIES OF EUCOSMA HUBNER (TORTRICIDAE: OLETHREUTINAE) FROM THE TALL
GRASS PRAIRIE REGION OF MIDWESTERN NORTH AMERICA
DONALD J. WRIGHT
3349 Morrison Ave., Cincinnati, Ohio 45220-1430, USA email: wrightdj@fuse.net
ABSTRACT. Eucosma haydenae, new species, is described from Iowa and Illinois. This species, which seems to be a tall grass prairie
obligate, is similar in appearance to E. rusticana (Kearfott) but is much smaller. A review of rusticana is included, and illustrations are provided
for the adults and genitalia of both species.
Additional key words: Eucosmini.
The once pervasive tall grass prairie of the North
American Midwest is now reduced to a scattered
assortment of small disjunct patches, but those
remnants still harbor insects that are rarely encountered
elsewhere. This paper proposes a name for one such
insect, a small brown moth described below as Eucosma
haydenae, new species. It was discovered during faunal
surveys in five prairie preserves, one in northeast Iowa,
four in the vicinity of Chicago, Illinois. In general
appearance it is most similar to Eucosma rusticana
Kearfott, but the two species are separated easily by
their marked difference in size.
Eucosma rusticana is distributed widely in eastern
North America but is rather poorly represented in
collections. Kearfott (1905) described the species based
on six specimens. Klots (1942) reported two syntypes in
the American Museum of Natural History (AMNH),
including one labeled LECTOTYPE, a designation he
attributed to Heinrich (1923). I examined the lectotype.
The second syntype in the AMNH, reported by Kearfott
(1905) from Algonquin, Illinois, was not found. I also
examined four syntypes at the United States National
Museum (USNM).
MATERIALS AND METHODS
This study is based on an examination of 97 adult
specimens and 10 genitalia preparations from the
following collections: AMNH, Loran D. Gibson, Todd
Gilligan (TG), Mississippi Entomological Museum
(MEM), USNM, Ron Panzer, and Donald J. Wright
(DJW). Forewing length (FWL), defined as distance
from base to apex (including fringe), was measured to
the nearest one tenth of a millimeter. A rough indication
of forewing geometry is provided by the aspect ratio
(AR), defined as FWL divided by medial forewing
width, the later measurement taken perpendicular to
the dorsal margin. Costal fold ratio (CFR) is defined as
costal fold length divided by FWL. Reported values of
AR and CFR are averages, rounded to two decimal
places, of the corresponding values calculated for a
small sample of specimens. The number of
measurements or observations supporting a particular
statement is indicated by n. Forewing pattern
terminology follows Brown & Powell (1991) and
Baixeras (2002).
SPECIES ACCOUNTS
Eucosma rusticana Kearfott
(Figs. 2, 4, 7, 8)
Eucosma rusticana Kearfott 1905:358: Barnes and
McDunnough 1917:170; Heinrich 1923:125, Fig. 162:
McDunnough 1939:47; Powell 1983:35
Types. Lectotype designated by Heinrich (1923): 3, Kerrville, Tex.,
AMNH. Paralectotypes. NORTH CAROLINA: Tryon, Fiske, 13 May
1903 (1 6, USNM), 28 May 1904 (1 3, USNM), 1 August 1904 (1 3,
USNM, genitalia slide 70461), no date (1 9, USNM).
Remarks. Eucosma rusticana is identified by the
following forewing characteristics: dorsal surface (Fig.
2) divided longitudinally into a blackish-brown anterior
region and a brownish-tan dorsoterminal region, the
line of separation running roughly along the cubital vein
from base to ocellus ee fron, there obliquely outward
to costa just short of apex: region between said line and
dorsal margin crossed longitudinally by three or four
brown streaks; ocellus with pale-tan central field,
bordered basally and distally by indistinct, transverse,
silvery-gray bars and crossed longitudinally by two,
variably expressed, dark-brown dashes: ninth costal
strigula white and conspicuous, other costal strigulae
gray and obscure. In melanic specimens the anterior
region of the forewing is nearly all black, with black
suffusion extending to dorsum, but the streaked
appearance of the dorsal region is still apparent.
Forewing statistics: 6 FWL 9-12 mm (mean = 10.3, n =
25), AR = 2.57, CFR = 0.47,2 FWL 9.9-12.2 mm (mean
= 11.2,n =5), AR = 2.43.
Male genitalia (Fig. 4) (n = 2): Uncus convex and moderately
developed, socii long and setose, dorsolateral shoulders of tegumen
well developed, gnathos a narrow band; vesica with ca. 22 deciduous
cornuti, valva with dorsal margin strongly concave, apex rounded,
SOCIETY
JOURNAL OF THE LEPIDOPTERISTS’
Fics 1-4: Adults and male genitalia. 1, haydenae, holotype. 2, rusticana, Rowan Co., Kentucky. 3, haydenae, Howard Co., Iowa,
slide DJW 556. 4, rusticana, Wy. andot Co., Ohio, slide DJW 167. Scale bars = 0.5 mm
ventral two thirds of distal margin convex, dorsal one third mildly
inset, producing narrowing of apical one third of cucullus, anal angle
rounded, neck with scooped out invagination of ventrolateral margin
(indicated by dashed line in Fig. 4), corner of sacculus rounded and
nearly right-angled, margin of basal opening with weakly developed
medial projection supporting a small patch of spines. Female
genitalia (Fig. 8) (n = 2): papillae anales small, facing laterally and
sparsely setose; sterigma (Fig. 7) semirectangular, length ca. 1.5x
width, with shallow trough from center of posterior margin to ostium,
lamella antevaginalis ringlike and very weakly sclerotized, lamella
postvaginalis with lateral and posterior surfaces densely
microtrichiate; posterior margin of sternum VII invaginated to three
fourths length of sterigma and closely approximate ‘thereto; ductus
bursae short, of nearly > aa arna width, with variably sclerotized ring
posterior to juncture with ductus seminalis; corpus bursae with two,
large, fin-shaped signa, inner surface of membrane microtrichiate.
Biology and distribution. I examined 52 specimens
(46 ¢, 6°) from Arkansas, Illinois, Indiana, Kentucky,
Missouri, Mississippi, North Carolina, Ohio, Tennessee,
Texas and Wisconsin. The flight period extends from
mid April to mid August, the earliest records coming
from Mississippi. Midwest records are predominantly
from June and July. No larval host has been reported,
but other members of the genus are known to be stem
and root borers of Asteraceae.
Eucosma haydenae new species
(Figs. 1, 3, 5, 6)
Diagnosis. Size and forewing maculation distinguish
haydenae from other eastern North American species of
Eucosma. Mean FWL of superficially similar rusticana
is ca. 3 mm longer than that of haydenae. Distinctive
male genitalic characters include the scooped out
invagination of the medioventral surface of the valval
neck and the rounded anteroventral projections of the
anellus. The sclerotized twist in the female ductus
bursae is prominent but not unique to this species.
Description. Head: Lower frons creamy white, upper frons
creamy white to light brown, vertex brown, scales adjacent to eye
lighter; labial palpus with basal segment white, second segment with
medial surface and dorsal margin white, lateral surface gray brown
with white medial mark, third segment brown, often with blackish-
brown apex; antenna with dorsal surface brown, posterior surface
white. Thorax: Dorsal surface orange brown, scales at posterior
extremity of tegula shading to tan, ventral surface creamy white, fore
and midlegs with anterior surfaces pale gray brown, posterior surfaces
white, hindlegs white to tan, midtibia with white, oblique, medial
mark on anterior surface, fore and mid tarsi with pale white
annulations. Forewing (Fig. 1): ¢ FWL 6-8.2 mm (mean = 7.3, n =
29), AR = 2.82, CFR = 0.48.2 FWL 7.5-8.2 mm (mean = 7.9, n = 2),
AR = 2.71; costa weakly convex, apex nearly right-angled, termen
weakly convex; dorsal surface with blackish- brown region bounded
anteriorly by basal one half of costa, posteriorly by cubital vein and
distally by end of discal cell, region between cubital vein and dorsum
white to tan, with orange-brown suffusion near base and along basal
margin of ocellus, a narrow, sometimes interrupted, brown streak
along dorsal margin; ocellus bordered basally and distally by silvery-
gray transverse bars, central field light tan, crossed longitudinally by
up to four, variably expressed, thin, black dashes: orange- -brown
scaling along distal one half of costa, crossed by five, paired, white to
gray strigulae and their associated silvery-gray stria; ninth costal
strigula white; fringe gray brown anteriorly, becoming paler toward
VOLUME 60, NUMBER 3
tornus. Male genitalia (Fig. 3) (n = 4): Uncus a rounded, dorsally
setose lobe, divided medially by shallow indentation; socii long and
densely setose; gnathos a narrow band, aedeagus tapered distally,
vesica with ca. 22 deciduous cornuti; anellus with small rounded
projections at anteroventral extremities; valva with costal margin
strongly concave, apex evenly rounded, distal margin convex, ventral
angle narrowly rounded, neck with strongly scooped out invagination
of medioventral margin, cucullus with medial surface densely setose
and with ca. 10 stout setae evenly distributed along distal margin,
sacculus moderately setose, margin of basal opening with narrow
raised pulvinus, the latter connected to neck by weakly developed
ridge. Female genitalia (Fig. 5) (n = 2): papillae anales small, facing
ventrally and moderately setose; sterigma (Fig. 6) long and
semirectangular, length more than 2x width, lamella antevaginalis
ringlike and weakly sclerotized, lamella postvaginalis with lateral
ridges bordering shallow medial trough, surface microtrichiate;
sternum VII with posterior margin deeply and narrowly invaginated to
length of sterigma, closely approximate to sterigma; ductus bursae
with sclerotized twist posterior to juncture with ductus seminalis;
corpus bursae with two fin shaped signa posterior to mid bursa, inner
surface microtrichiate.
Holotype. ¢, IOWA, Howard Co., Hayden Prairie, 23 June 1997,
D. J. Wright, deposited in USNM. Type locality at 43°26' 35" N,
92°29! 58" W.
163
Paratypes. ILLINOIS: Dupage Co., W. Chicago Prairie, R.
Panzer, 23 May 2004 (5 d, 2°, 6 genitalia slide DJW1291,2 genitalia
slides DJW1290 & 1296), 6 June 2004 (1 ¢); Lee Co., Green River E.,
6 July 2002 (1 ¢). IOWA: Same locality as holotoype, D. J. Wright, 21
June 2000 (1 ¢), 23 June 1997 (8 ¢, genitalia slide DJW 556), 28 June
1995 (2 ¢, genitalia slides DJW 131 & 206), T. Gilligan (2 2). Paratype
depositories: AMNH, TG, MEM, USNM, DJW.
Etymology. Dr. Ada Hayden (1884-1950) was a
botanist at Iowa State College (now Iowa State
University) who devoted much of her professional life to
the study of the native Iowa prairie (Isely, 1989). Her
advocacy in the 1940's for conservation of prairie habitat
was largely responsible for the preservation of the 240
acre tract in northeast Iowa that now bears her name
and serves as the type locality for the moth described
here. It's a pleasure to name this insect after Dr.
Hayden.
Distribution and biology. I examined 45 specimens
(43 d, 22), documenting a flight period from late May to
Fics. 5-8: Female genitalia. 5-6, haydenae, Dupage Co., Illinois, slide DJW 1290. 7-8, rusticana, Cook Co., Hlinois, slide DJW
1289. Scale bars = 0.5 mm.
164
the beginning of July. All were collected in remnant tall
grass prairie habitat in Howard County, Iowa, and in
Cook, Dupage, and Lee Counties in Illinois. The larval
host is not known but, as with rusticana, is probably a
species of Asteraceae.
Remark. The haydenae specimens from the Chicago
area have a generally blacker appearance than those
from Iowa.
ACKNOWLEDGEMENTS
I thank J. W. Brown, R. L. Brown, and R. T. Schuh for the
loan of specimens under their care. Karl Gnaedinger generously
supplied me with specimens from the Chicago area, and D.
Howell, Iowa Department of Natural Resources, helped with
permits to sample the Hayden Prairie fauna and with biograph-
ical information on Ada Hayden. Two anonymous reviewers of-
fered helpful comments on the manuscript.
LITERATURE CITED
BAIxeERAS, J. 2002. An overview of genus level taxonomic problems
surrounding Argyroploce Hiibner (Lepidoptera: Tortricidae),
with description of a new species. Ann. Entomol. Soc, Am.
95(4):422-431.
Barnes, W. & J. MCDUNNOUGH. 1917. Checklist of the Lepidoptera
of Boreal America. Herald Press, Decatur, Illinois. 392 pp.
JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY
Brown, R. L. & J. A. POWELL. 1991. Description of a new species of
Epiblema (Lepidoptera: Tortricidae: Olethreutinae) from coastal
redwood forests in California with an analysis of the forewing
patter. Pan-Pacific Entomol. 67:107-114.
Hetnricu, C. 1923. Revision of the North American moths of the
subfamily Eucosminae of the family Olethreutidae. U.S. Nat.
Mus Bull. 123:1-298.
IseLy, D. 1989. Ada Hayden: A Tribute. Jour. Iowa Acad. Sci. 96(1):1-
5.
Kearrortr, W. D. 1905. Descriptions of new species of Tortricid
moths from North Carolina, with notes. Proc. U.S. Nat. Mus.,
28:349-364.
Kiots, A. B. 1942. Type material of North American microlepi-
doptera other than Aegeriidae in the American Museum of Nat-
ural History. Bull. Amer. Mus. of Nat. Hist. 79:391-424.
McDunnouc3, J. 1939. Check List of the Lepidoptera of Canada and
the United States of America. Part II Microlepidoptera. Mem.
South. Calif. Acad. Sci. 2:3-171.
POWELL, J. A. 1983. Tortricidae, pp. 31-41. In Hodges, R. W. et al.
(eds.), Check list of the Lepidoptera of America north of Mexico.
E. W. Classey & Wedge Entomol. Res. Foundation. London,
England.
Received for publication 6 January 2006; revised and accepted 19 June
2006
VOLUME 60, NUMBER 3
Journal of the Lepidopterists’ Society
60(3), 2006, 165-170
165
MONARCH (DANAUS PLEXIPPUS L. NYMPHALIDAE) MIGRATION, NECTAR RESOURCES AND
FIRE REGIMES IN THE OUACHITA MOUNTAINS OF ARKANSAS
D. CratG RUDOLPH, CHARLES A. ELy, RICHARD R. SCHAEFER, J. HOWARD WILLIAMSON, AND RONALD E. THILL
Wildlife Habitat and Silviculture Laboratory (maintained in cooperation with the Arthur Temple College of Forestry,
Stephen F. Austin State University), Souther Research Station, USDA Forest Service,
506 Hayter Street, Nacogdoches, Texas 75965 USA Email: crudolph01@fs.fed.us
ABSTRACT. Monarchs (Danaus plexippus) pass through the Ouachita Mountains in large numbers in September and October on their
annual migration to overwintering sites in the Transv olcanic Belt of central Mexico. Monarchs are dependent on nectar resources to fuel their
migratory movements. In the Ouachita Mountains of west-central Arkansas migrating monarchs obtain nectar from a variety of plant species.
especially Bidens aristosa and other composites. Fire suppression has greatly alteredit the structure of forest communities with major implica-
tions for ecological relationships. Sites that are undergoing restoration to a shortleaf pine-bluestem grass community following thinning and fre-
quent prescribed fire, and thought to closely resemble. pre-European conditions, support ineroneed abundances of nectar resources and
migrating monarchs compared to mintreated controls. These results suggest that widespread fire-suppression since the early 1900s has substan-
tially reduced nectar production for migrating monarchs in the Ouachita Mountains Physiographic Region.
Additional key words: Interior Highlands, surveys, restoration
The eastern North American population of the
monarch butterfly, Danaus plexippus L., undertakes one
of the most remarkable migrations of any lepidopteran
(Urquhart 1976, Brower & Malcolm 1991). During the
fall most individuals of this population migrate to
extremely restricted sites in the Transvolcanic Belt of
central Mexico (Urquhart 1976, Calvert & Brower
1986). Concern has been expressed about the
continued health of this population and the persistence
of the massive migration phenomenon (Wells et al.
1983, Brower & Malcolm 1991). Changes in abundance
and quality of larval hosts (Zalucki & Brower 1992), loss
of critical overwintering sites due to logging and fire
(Brower 1996, Brower et al. 2002), vehicle mortality
(McKenna et al. 2001), pesticides (Oberhauser 2004),
introduced species (Calvert 2004), and transgenic Bt
modified crops (Losey et al. 1999, Jesse and Obrycki
2004) have been identified as actual or potential threats.
Less attention has been directed to landscape-level
changes in nectar availability which ultimately fuels the
extended fall migration to central Mexico (Garcia &
Equihau-Zamora 1997, Brower & Pyle 2004).
Land use changes and management protocols in
more natural habitats have dr: eeeally altered essentially
all the land base that constitutes the breeding range and
migration corridors of D. plexippus in eastern North
America. The Ouachita Mountains Physiographic
Region of west-central Arkansas and southeastern
Oklahoma, encompassing 3,237,600 ha, remains
primarily forested (Bukenhofer and Hedrick 1997).
However, logging, fire suppression, and silvicultural
management have altered vegetation structure and
composition throughout the region (Foti and Glenn
1991, Masters et al. 1995). Fire-maintained shortleaf
pine (Pinus echinata) forests were widespread in the
Ouachita Mountains until the early 20th century (Foti &
Glenn 1991). Since the original harvest of these pine
forests, most forested sites have been altered using
intensive short-rotation pine production or remain as
more natural forests, but have been subjected to fire
suppression for several decades (Bukenhofer and
Hedrick 1997). In either case, the abundance and
quality of nectar resources available to Lepidoptera has
been drastically altered (Thill et al. 2004).
The managers of the Ouachita National Forest have
initiated a landscape scale restoration of the fire-
maintained shortleaf pine-bluestem (Schizachrium spp..
Andropogon spp.) ecosystem on 48,706 ha (U. S. Forest
Service 1996). Restoration involves thinning the
overstory, reduction of midstory vegetation, and
prescribed burning on a three-year return interval. This
restoration was undertaken to restore habitat for the
endangered red-cockaded woodpecker (Picoides
borealis) and to restore what is thought to be the pre-
European structure and composition of the vegetation
(Foti and Glenn 1991, Bukenhofer and Hedrick 1997).
A number of authors have examined the effect of these
restoration efforts on a diversity of taxa (see Thill e¢ al.
2004).
As part of ongoing studies of the effects of restoration
of fire-maintained shortleaf pine-bluestem habitat on
lepidopteran communities, butterfly and nectar
resource surveys were conducted in restored and
untreated control plots. This paper reports results for D.
plexippus in relation to the fire regime and suggests
implications for fall migration and over-winter survival.
MATERIALS AND METHODS
This study was conducted on the Poteau Ranger
District (34°45'N, 34°15'W) of the Ouachita National
166
Forest in west-central Arkansas. Topography in this
region consists of east-west trending ridges and valleys
of 150-820 m. Mixed
hardwood forests dominate north-facing slopes and pine
and mixed pine-hardwood forests dominate south-
with an elevational range
facing slopes. Prior to the initiation of fire-suppression
activities, much of the landscape, especially the more
xeric pine communities on south- and west- facing
slopes, burned on a regular basis (Foti and Glenn 1991,
Masters et al. 1995). These were primarily low intensity
ground fires ignited by lightning, Native Americans, and
more recently by European colonists. The resulting
forest structure was characterized by pine- -dominated
overstories, sparse midstories, and diverse herbaceous
understories (du Pratz 1975, Featherstonhaugh 1844,
Nuttall 1980, Foti and Glenn 1991).
The Ouachita Mountains are still predominately
forested; however, logging and fire-suppression have
dramatically altered vegetation structure (Bukenhofer
and Hedrick 1997).
conditions, existing forests are typically characterized by
Comps ured to pre-European
a younger and denser canopy, a dense woody midstory,
anda very suppressed herbaceous understory (F enwood
et al. 1984, Masters 1991, Sparks 1996). These changes
have profoundly altered the original biodiversity of the
Ouachita Mountains (Neal and Montague 1991, Smith
and Neal 1991, Wilson et al. 1995, Sparks et al. 1998,
Thill et al. 2004).
Landscape scale restoration was initiated in 1979,
formally incorporated into the Ouachita National Forest
Plan in 1996, and currently projects the restoration of
48,706 ha (7.3% of the Forest) to a shortleaf pine-
bluestem condition (U. S. Forest 1996,
Bukenhofer and Hedrick 1997).
Restoration consists of thinning of the
Service
overstory,
removal of most midstory vegetation, and prescribed
burning on approximately a 3-year rotation (U.S. Forest
Service 1996). Ultimately, regeneration of canopy trees
will be accomplished primarily through _ the
implementation of irregular shelterwood and seed-tree
harvests with a portion of the overstory retained
indefinitely (U. S. Forest Service 1996). In addition,
rotation age will be lengthened, primarily to provide
sufficient older trees to support red-cockaded
woodpecker recovery (Rudolph and Conner 1991). A
the initiation of our studies, approximately 9,071 ha had
been restored in a 42,148 ha landscape on the Poteau
Ranger District (W. G. Montague pers. com.). We use
“restored” in a relative sense and recognize that stands
are on a trajectory toward an ecological state that
resembles pre-European conditions.
Je JURNAL OF THE LEPIDOPTERISTS’ SOCIETY
MATERIALS AND METHODS
We censused butterflies and nectar
annually in
resources
and three
from 10.5 to
All treated sites had received a minimum of
\ nine restored (treatment) sites
un-restored (control) sites. Sites varied
42.1 ha.
four prescribed burns prior to the initiation of the study.
Restored sites were included as portions of larger areas
(range 65 to 2226 ha) bummed on the same day. A total
of three restored sites were prescribe- -burned each
spring. Consequently, in any given year, first, second,
and third growing seasons post-fire were each
Additional details
concerning treatment and control sites can be found in
Thill et al. (2004).
Adult butterflies (only D. plexippus data reported
here) were censused using a time-constrained walking
along 500-m triangular transects centrally
located within each site (Pollard 1977, Pollard and Yates
1993). Individual transects were censused by slowly
walking the length of transects for approximately 20
Time involved in counting butterfly aggregations,
netting individuals for identification, or waiting for sun
or wind conditions to conform to set variables was not
included in the 20-min period. During 2000-2003,
census counts were conducted four times each year
(first week of April, June, August, October). Only
October data for D. plexippus are reported here.
Census counts were replicated three times each month,
represented by three _ sites.
census
min.
each replicate conducted by a different observer on
different days during the survey week. Individual
censuses were conducted between 0900 and 1330 hrs
CST when temperatures were between 18—36° C and
wind velocity beneath the canopy was not too high to
suppress butterfly flight (Beaufort Scale <4). Censusing
was further restr jcted to periods when sunlight was
sufficient to cast discernable shadows. The response of
butterflies to light, wind, temperature and cloud cover
seasonally and daily in complex ways.
Consequently, observer judgement further constrained
varies
censusing to those periods when butterfly activity
appeared to be substantial. Additional details
concerning treatment and control sites can be found in
Thill et al. (2004).
In addition to census counts of butterflies, we
recorded all observations of nectaring and other feeding
activities by butterflies observed during this and other
studies in the Ouachita Mountains hemveen 1999 and
2004. Voucher specimens were deposited in the
herbarium of Stephen F. Austin State University. Plant
nomenclature uses the nomenclature found in Smith
(1994).
Nectar resources were quantified by counting flowers
VOLUME 60, NUMBER 3
in three 1x100-m plots at each study site. Plots were
located parallel to each side of the triangular 500-m
butterfly census transect. Within these plots all
potential nectar resources were counted each week that
butterfly censuses were conducted. For most plant
species, individual flowers or composite heads (capitula)
were enumerated. Inflorescences, or portions thereof,
were counted for a few species with small and/or dense
aggregations of flowers (e.g. Ceanothus americanus,
Allium spp., Solidago spp., Apiaceae). These
enumeration decisions were based on estimating the
structure that most closely approximated a separate
landing site for a typical butterfly.
TaBLE 1. List of nectar plants, inclusive nectaring dates by
plant species, and number of nectaring observations for
monarchs (Danaus plexippus) in the Ouachita Mountains,
Arkansas during 1999-2005.
Plant Species Dates # Observations
Bidens aristosa 9/7-10/5 1890
Eupatorium serotinum 8/20-10/5 164
Solidago spp. 8/1-10/5 101
Vernonia baldwinii 6/30-10/2 93
Cunila origanoides 9/9-10/2 69
Solidago petiolaris 9/27-10/2 60
Liatris elegans 7/2-10/4 44
Helenium amarum 7/2-10/2 40
Aster spp. 9/8—10/5 39
Aster ericoides 9/27-10/4 28
Aster anomalus 9/27-10/2 22,
Senecio obovatus 4/54/23 21
Solidago rugosa 10/1-10/5 19
Asclepias tuberosa 5/17-8/2 14
Polygonum pensylvanicum — 10/2-10/5 13
Eupatorium spp. 8/20-10/5 10
35 other species 74
RESULTS
In the Ouachita Mountains D. plexippus adults were
rarely observed in spring and summer. However, during
the fall migration in September and October, D.
plexippus were common to abundant. We obtained a
total of 2701 D. plexippus nectaring observations from
1999 to 2004. All but 101 of these observations were
made during September and October (Table 1).
Danaus plexippus nectared most frequently on Bidens
aristosa (1890, 70.0%), Solidago spp. (180, 7.0%),
Eupatorium serotinum (164, 6.1%), Aster spp. (98,
167
3.6%), Vernonia baldwinii (93, 3.4%), Cunila
origanoides (69, 2.6%), Liatris spp. (59, 2.2%), and
occasionally on an additional 31 species. The 1890
nectaring observations on Bidens aristosa were during
the fall migration, and were concentrated primarily on
the verges of the extensive road system within the
region.
Due to low regional abundance we counted only 15
D. plexippus during April, June, and August censuses.
During October, generally the peak of migration, a total
of 1019 D. plexippus were detected during transect
surveys. These observations occurred most often on
restored treatments, especially during the first growing
season post-fire, rather than on controls (Table 2).
Significant differences were detected across treatments
(x2 = 1637.6, df = 3, P < 0.001).
Nectar resources were also more abundant on
restored treatments than on controls (Table 3). Within
restored treatments, nectar resources were more
abundant on first year post-burn treatments and least
abundant in third year post-burn treatments during the
October surveys. Significant differences were
frequently detected across these four treatments (Table
3).
TABLE 2. Number of Danaus plexippus, summed across plots
and observers, detected during October surveys on restored
sites and control sites on the Ouachita National Forest during
2000-2002.
Year B-l+ B-2 B-3 Control
2000 55 29 53 9
2001 746 101 13 0
2002 8 3 1 1
Total S09 133 67 10
“ B-1, B-2, and B-3 correspond to 1, 2"¢, and 3™ growing
seasons post-burn.
DISCUSSION
Restored sites in the Ouachita Mountains consisting
of a fire-maintained shortleaf pine-bluestem community
supported a higher abundance of D. plexippus during
migration than un-restored, fire-suppressed controls.
This pattern was most noticeable during the first
October post-fire. Both aspects of this pattern were
similar to the abundance of nectar sources. Numbers
of D. plexippus detected varied considerably across
years. This may have been due to timing of the
migration relative to our survey times, or changes in D.
plexippus numbers in the eastern North American
168
TABLE 3.
JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY
Response of nectar resources to pine-bluestem restoration on the Ouachita National Forest during October 2000-2002.
Mean number of nectar resources per 300 m? plots in restored and control stands. Means in the same row sharing the same letter
were not significantly different in a 1-way ANOVA with REGWQ at P < 0.1 (SAS Inst. Inc. 1988:598).
Because abundance values
varied eee within treatments, data were rank transformed prior to analysis.
VaWeGiet
B-l* B-2 B-3 Control
Year x SE x SE x SE x SE P
2000 746.3A 66.4 84.3B 45.9 59.3B 21.9 (EBS 5.0 0.0010
2001 2745.0A 604.9 3219.0A 1605.5 515.3B 106.2 183.7B 130.4 0.0047
2002 4587.3A 2987.2 1746.7A 689.1 719.3AB 355.0 35.0B 8.3 0.0181
* B-1, B-2, and B-3 correspond to 1*, 2"*, and 3" growing seasons post-burn
population. The very low numbers in 2002 followed of flowers located in disturbed sites, mainly the
catastrophic winter storm mortality at the Mexican over-
wintering sites the previous January (Brower et al.
2004).
In the absence of frequent fire, nectar resources in
the forested portions of the Ouachita Mountains
Physiographic Region are generally low. Improved
pastures and intensively managed pine plantations 2-3
years after planting, the only other significant land uses
in the region, also typically support a low abundance of
nectar resources. Limited areas of increased nectar
abundance occur in disturbed sites, i.e. road verges,
utility rights-of-way, fence rows, and areas of recent
timber harvest.
primarily forested area, nectar resources may be
Thus, at the regional level, in this
limiting for lepidopteran species that require them
(Thill et al. 2004, Rudolph e¢ al. In prep.). Similar
results were found for fire-maintained pine
communities in eastern Texas (Rudolph and Ely 2000).
Due to the very patchy distribution of nectar sources
in relation to roads, rights-of-way, and forest
management we were unable to assess the relative
abundance of nectar
landscape. The relative numbers of nectaring
observations in Table 1 are probably biased toward
those occurring along road_ verges. However,
butterflies, including D. plexippus, were frequently
observed nectaring along the transect surveys, especially
in the restored treatments.
Data presented here indicate that fire suppression
has resulted in a landscape that is currently depauperate
in nectar availability. We suggest that reduced nectar
availability, compared to probable pre-European
conditions, limits use of the widespread fire-suppressed
habitats by monarchs and other lepidopteran species
(Thill et al. 2004). The large numbers of monarchs
passing through the Ouachita Mountains each fall
obtain nectar resources primarily from concentrations
sources across the entire
abundant Bidens aristosa growing on road verges. How
this current availability of nectar resources compares
with the pattern of nectar resources dispersed across the
forest landscape in the pre-European fire-maintained
shortleaf pine forests is unknown, but this pattern
suggests that fire suppression has greatly limited the
availability of nectar resources across most of the
forested landscape.
Lipid-loading by D. plexippus during the fall
migration, both to fuel the migration and sustain winter
survival, is crucial (Brower 1985, Alonso- Mejia et al.
1997). Brower and Pyle (2004) suggest that lack of
nectar resources might bea limiting eiaon for migrating
D. plexippus. In addition, a significant proportion of the
monarchs inhabiting the wintering sites in the
Transvolcanic Belt lack sufficient lipids to survive until
spring (Brower & Pyle 2004). A detailed understanding
of the current quality and availability of nectar resources
along the migration corridor, and effects of historical
changes, mould allow an improved understanding of the
remarkable migration of D. plexippus. Our results
suggest that historical changes have been substantial,
even in a landscape still dominated by “natural” plant
communities. These changes may have potential
consequences for migrating monarchs. Habitat and
critical ecological processes, fire in this instance, may
both be important in maintaining a major biological
phenomenon on a continental scale.
ACKNOWLEDGMENTS
We thank R. Buford, A Carter, C. Collins, and R. Perry for as-
sistance in various aspects of fieldwork. N. Koerth provided as-
sistance with data analysis and T. Trees maintained databases.
Partial funding for this work was provided by the Poteau District
of the Ouachita National Forest and the Ouachita Mountains
Ecosystem Management Research Project. We thank R. M.
Pyle, P. A. Opler, W. G. Montague, L D. Hedrick and two anony-
mous reviewers for comments on an earlier draft of this manu-
script.
VOLUME 60, NUMBER 3
LITERATURE CITED
ALONSO-MEJIA, A., E. RENDON-SALINAS, E. MONTESINOS-PATINO, AND
L. P. BRowER. 1997. Use of lipid reserves by monarch butterflies
overwintering in Mexico: Implications for conservation. Ecol.
App. 7:934-947.
Brower, L. P. 1985. New perspectives on the migration biology of
the monarch butterfly, Danaus plexippus L. Pp. 748-785 in M.
Am. Rankin (ed.), Migration: Mechanisms and adaptative signifi-
cance. University of Texas Press, Austin. 876 pp.
Brower, L. P. 1996. Forest thinning increases monarch butterfly
mortality by altering the microclimate of the overwintering sites
in Mexico. Pp. 33-44 in S.A. Ae, T. Hirowatari, M. Ishii & L. P.
Brower (eds.), Decline and conservation of butterflies in Japan
III. Proceedings of the International Symposium on Butterfly
Conservation, Osaka, Japan. 217 pp.
BROWER, L. P., G. CASTELLEJA, A. PERALTA, J. LOPEZ-Garcta, L. Bo-
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JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY
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Received for publication 16 December 2005; revised and accepted 28
June 2006
GENERAL NOTES
Journal of the Lepidopterists’ Society
60(3), 2006, 171-174
OBSERVATIONS OF KRICOGONIA LYSIDE (PIERIDAE) IN THE FLORIDA KEYS
Additional key words: West Indies, migration, seasonal dispersal, Florida Keys
Kricogonia lyside (Godart), (Fig. 1) a butterfly native
to the West Indies, the southwestern United States, and
Central and South America, has historically occurred on
an irregular basis on several of the middle and upper
Florida Keys, as well as on the southeastern Florida
peninsula (Young 1938, Minno and Emmel 1993, Smith
et al. 1994, Glassberg et al. 2000, Minno et al, 2005). A
highly migratory species, K. lyside often travels en masse
within and between the islands of the Caribbean
Fic 1. Kricogonia lyside on Vaca Key,
Credit: H. L. Salvato).
23 July 2005 (Photo
(Wolcott 1927, Williams 1930, Smith et al. 1994, Miyata
2000). The species also demonstrates similar mass
movements in the southwestern United States with
large influxes entering southern Texas from Mexico
(Clench 1965, Gilbert 1985). Dispersal of K. lyside into
and within southern Texas appears to be triggered by
reductions in new hostplant growth (Gilbert 1985).
However, whether or not similar ecological cues
encourage K. lyside migrations, both within the
Caribbean and to southern Florida, remains unknown.
Luis R. Hernandez (pers. comm.) suggests tropical
storm activity in the Caribbean may play a large role in
the dispersal of K. lyside within the West Indies, and
perhaps to southern Florida. Smith et. al. (1994)
witnessed several K. lyside making landfall on Upper
Matecumbe Key via strong, although not storm-related,
easterly winds, indicating the species is capable of
dispersal into the Keys during typical seasonal
conditions. Young (1938), based on examination of fresh
specimens collected near Biscayne Bay, suggested that
the species reproduces within Souther Florida and the
Keys. However, neither oviposition nor larval activity
has ever been observed for K. lyside in the region. A
known hostplant of this species, lignum vitae, Guaiacum
sanctum L., (Zygophyllaceae) occurs commonly within
hardwood hammocks throughout the Keys and is also
widely used in the region as an or namental species in
landscaping.
Following an apparent decade-long absence in
Florida K. lyside was observed locally from June
through September 2002 on Plantation Key (Salvato
and Salvato 2002) and Key West (MHS unpublished
data) in the Florida Keys. After these initial
observations of K. lyside re-occurrence in the Keys
MHS and HLS continued to survey for this species as
part of a larger ongoing long-term study to determine
the status and distribution of butterflies throughout the
Keys. Surveys were conducted on warm, clear days
under conditions that were considered sufficient for
butterflies to be flying. Each sampling date included
approximately S-9 hours of field time (between
0800-1700 h) in which two surveyors (MHS and HLS)
walked a standard route at survey sites within the study
area to visually record K. lyside activity. On each
sampling date a selected span of the Keys was
monitored in either the Upper (Key Largo to Upper
Matecumbe Key), Middle (Lower Matecumbe Key to
Vaca Key) or Lower Keys (Bahia Honda Key to Key
West). Survey sites on each island were visited monthly
during May 2002 to December 2005, with the exception
of Lignumvitae Key, which due to its inaccessibility, was
only surveyed during June of each survey year. The
amount of time spent surveying for K. lyside on each
Key varied based on island size and number of survey
sites. Overall, a total of 25 survey sites, of varying sizes,
were monitored monthly during this study. These
locations included State and County parks, National
Wildlife Refuges and roadsides. Guaiacum sanctum
was observed either directly within or adjacent to the
majority of our study areas.
Although there were unconfirmed reports of K. lyside
during ¢ 2003, we did not observe the species again in the
Keys until 13 June 2004, after which K. lyside was
frequently encountered throughout the Florida Keys, as
well as southern Miami- Dade County, marking the first
known occurrences for the species on mainland Florida
in several years. Kricogonia lyside was absent from the
majority of the islands we surv eyed by mid-August 2004;
however, local occurrences remained on Bahia Honda
within the lower Keys into November of that year.
In 2005, we began to re-encounter K. lyside during
the early summer months with our first observations
occurring on Lignumvitae Key on 11 June. Despite an
abundance of G. sanctum on Lignumvitae Key, K. lyside
had never been reported from this island (Minno and
Emmel 1993, Smith et al. 1994). To our knowledge
these observations of K. lyside on Lignumvitae Key
represent the first reports from this island. From June
through August of 2005 K. lyside was observed on every
island we surveyed within the Keys (Fig. 2).
Throughout this time frame the species was witnessed
actively dispersing towards the east on both the
northern and southern coastlines of the Keys, as well as
on outer islands such as Lignumvitae Key, indicating a
wider migratory swath than was noted earlier in either
the 2002 or 2004 observations. As was observed in
JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY
2004, K. lyside remained on Bahia Honda until later in
the season (at least through 24 September 2005) than
elsewhere in the Keys. Table 1 provides an overview of
K. lyside observations made on various islands within
the Florida Keys during this survey. As with
Lignumvitae Key, several of the K. lyside observations
listed in Table 1 appear to be the first documented
occurrences of the species on select islands, particularly
within the Lower Keys (Minno and Emmel 1993).
We observed an estimated 10, 74 and 238 individuals
of K. lyside during monthly surveys of the Keys in 2002,
2004 and 2005, respectiv ely. In all years the majority of
individuals were found on Key West, Bahia Honda, Vaca
Key and Plantation Key with varying numbers of
individuals occurring on 15 other islands in the Keys.
Although the sexes are similar, males from western
Cuba and the Bahamas are distinctive from other
known K. lyside populations in that these individuals
generally lack a black bar near the apex of the upper
hindwing (Riley 1975, Minno and Emmel 1993,
Hemandez 2004, J. Y. Miller, pers. comm.).
Examination of photographs (n = 25), collected
specimens (n = 2) and field observations (n = 322)
indicate that the K. lyside males encountered during
Gulf of Mexico
Windley Key
| Lignumvitae Key | Key
Lower Matecumbe Kayo Sam
Cudjoe Key
| Sugarloaf Key
Big Pine Key
Grassy Key
Upper
Matecumbe
Key
2 >
_e
7B
Long Key
Saddlebunch Keys
West Summerland Key
Stock Island |
Atlantic Ocean
Miles
Fic 2. Islands in the Florida Keys on which Kricogonia lyside was observed during the survey period.
VOLUME 60, NUMBER 3
TaBLE 1. Kricogonia lyside observations from the Florida
Keys during June through September of 2002, 2004 and 2005.
Island 2002 2004 2005
Key Largo 3
Plantation Key 7 12 10
Upper Matecumbe Key 2 1
Windley Key 1 2
Lignumvitae Key 2
Lower Matecumbe Key 2 1
Grassy Key i
Long Key if 3
Vaca Key 3 40
Knights Key 12
West Summerland Key 3
Bahia Honda Key S25 94
Big Pine Key 28
No Name Key 2
Cudjoe Key 4
Sugarloaf Key 2
Saddlebunch Key 1 6
Stock Island 2 4
Key West 3 12 20
° Indicates observations continued until November
these studies were morphologically similar to the
western Cuban and Bahamian populations, suggesting
that the seasonal occurrences of K. lyside in the Keys
and southern Florida observed in this study were likely
of West Indian origin.
During June to mid-August of 2004 and 2005
Kricogonia lyside was consistently observed in active
fren! traveling in either an easterly or northeasterly
direction between islands in search of, and in frequent
interaction with, G. sanctum. Conversely, K. lyside
observed later in the season during 2004 and 2005,
specifically those remaining on Bahia Honda, occurred
only locally and took nectar from any available sources,
not just those in the immediate proximity of the
hostplant.
Despite extensive searches throughout the study
period for signs that K. lyside had reproduced or
underwent diapause in southern Florida it was not until
9 July 2006 that MHS, HLS and Dennis J. Olle
observed the species mating and ovipositing on the fruit
and fresh growth of G. sanctum on Stock Island and Key
West. Additionally on Key West, K. lyside was observed
actively ovipositing on the leaves of Maracaibo lignum
vitae, Bulnesia arborea ( (Jacq.)Engl., (Zygophyllaceae) a
173
non-native species that occurs as an ornamental in the
Keys and was not common in our study areas. To our
knowledge these observations of oviposition provided
the first accounts of K. lyside reproduction in southern
Florida. However, K. lyside larval activity was not
observed during these studies. Larvae of K. lt lyside feed
nocturnally (Riley 1975, Hernandez 2004) and early
instars are similar in coloration to that of the new G.
sanctum growth on which they feed (Allen et al. 2005).
The mature larvae are darker green with white and
brown stripes (Minno et al. 2005). The cryptic
coloration and nocturnal feeding habits of K. lyside
larvae makes the species difficult to find on the
hostplant (Luis R. Hernandez, pers. comm.) and may
explain our inability to locate them. Additional studies
are needed to better determine the occurrence and
natural history of immature K. lyside in southern
Florida. Furthermore, there were no signs of adult K
lyside activity across the Keys by mid-fall of 2002, 2004
and 2005, suggesting the species may disappear from
Florida and then sporadically re- -colonize, probably
from Cuba. The tropical storm seasons in 2004 and, for
the Keys particularly 2005, were extremely active.
However, the influxes and northeastern movements of
K. lyside across the Keys and into southeastern Florida
preceded any significant storms occurring during these
study years.
Our observations indicate that K h yside periodically
disperses to a greater array of islands in the Florida Keys
than previously noted. We have documented that
during some years large numbers of K. lyside adults
occur in the Keys. Additionally, K. lyside appears to
breed in southern Florida, but to what extent the
species colonizes within the state requires further
investigation.
The authors thank Luis R. Hermandez for sharing his field ob-
servations and insights on the biology of K. lyside in the West In-
dies. Jacqueline Y. Miller examined and presented the authors
with her observations of K. lyside specimens within the collec-
tion of the McGuire Center for Lepidoptera and Biodiversity,
Florida Museum of Natural History, Gainesville, Florida. The
authors also extend thanks to Tim Adams, Don Stillwaugh Jr.
Lyn and Brooks Atherton, Byrum and Linda Cooper, Dennis J.
Olle and David L. Lysinger for sharing their field observations of
K. lyside during the 2004-05 migrations. We also thank an
anonymous reviewer for comments that helped improve the
manuscript. Finally, the authors thank Barry Wood for creating
and editing Figure 2.
LITERATURE CITED
ALLEN, J. T., P. Brock & J. J. GLassBERG. 2005. Caterpillars in the
Field and Garden; A Field Guide to the Butterfly Caterpillars of
North America. Oxford University Press, New York. 232 pp.
CLENCH, H. K. 1965. A migration of Libytheana and Kricogonia in
southern Texas. J. Lepid. Soc. 19:223-224.
GILBERT, L. E. 1985. Ecological factors which influence migratory
behavior in two butterflies of the semi-arid shrublands of south
Texas. Contrib. Mar. Sci. 27: no. suppl.
GxassBERG, J., M.C. MINNO & J. V. CALHOUN. 2000. Butterflies
through binoculars. Oxford University Press, New York. 242 pp.
HERNANDEZ, L. R. 2004. Field guide of Cuban-West Indies Butter-
flies. Ediluz, Maracaibo. 269 pp.
KIMBALL, C. P. 1965. The Lepidoptera of Florida — an annotated
checklist. State of Florida Department of Agriculture, Division
of Plant Industry. Gainesville, Florida. 363 pp.
MINNO,, M.C. & T.C. EMMEL. 1993. Butterflies of the Florida Keys.
Scientific Publishers, Gainesville, Florida. 168 pp.
MINNO, M. C., J. F. Butter, & D. W. HAL. 2005. Florida Butterfly
Caterpillars and Their Host Plants. University Press of Florida,
Gainesville. 341 pp.
Miyata, A. 2000. Mass migration of Kricogonia lyside (Lepidoptera,
Pieridae) in Santa Domingo, Dominican Republic, in 1995.
Trans. Lepid. Soc. Japan. 51: 281-286.
Ritey, N. D. 1975. A field guide to the butterflies of the West Indies.
Collins. London, England. 224 pp.
SatvaTo, M. H. & H. L. Satvato,. 2002. A new host record for
Automeris io and the re-occurrence of Kricogonia lyside in
Florida. News Lepid. Soc. 44:128.
Journal of the Lepidopterists’ Society
60(3), 2006, 174-176
JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY
Situ, D.S., L. D. MILLER & J. Y. MILLER. 1994. The Butterflies of
the West Indies and South Florida. Oxford University Press, New
York. 264 pp. 32 pl
Wo corr, G. N. 1927. Notes on the pierid butterfly, Kricogonia
castalia Fab, (Lepid.), Entomol. News. 38; 97-100.
WILLIAMS, C. B. 1930. The Migration of Butterflies. Oliver and Boyd,
London. p.133.
YounNG, F. N. 1938. Some interesting butterfly records for south
Florida. Entomol. News. 49:115.
Mark H. Satvato, 1765 17th Avenue SW. Vero
Beach, Florida, 32962, USA: Email:
anaea_99@yahoo.com, JOHN V. CALHOUN, 977 Wicks
Drive, Palm Harbor, Florida, 34684, USA: Email:
bretcall @verizon.net and HOLLy L. SatvaTo, 1765 17th
Avenue SW, Vero Beach, Florida, 32962, USA
Received for publication 29 December 2005, revised and accepted 21
June 2006.
A PRECAUTIONARY TALE ABOUT RARITY: ON THE LARVA AND LIFE HISTORY
OF LITHOPHANE JOANNIS (LEPIDOPTERA: NOCTUIDAE)
Additional key words: shelter-forming, Aesculus flava, Lithophane innominata, Lithophane patefacta
This note is about rarity, and how species that are
regarded as scarce may be anything but, once aspects of
their life history are better understood. Lithophane
joannis Covell and Metzler was not described until
1992. Prior to the authors' distribution of paratypes
there were no specimens of L. joannis in any major
eastern institution, i.e., the Smithsonian, American
Museum, and Carnegie Museum. Not William Forbes;
nor Jack Franclemont, Doug Ferguson, Michael Pogue,
Eric Quinter, or Tim McCabe has collected the moth.
Dale Schweitzer wrote his dissertation on the tribe—he
has yet to see the moth alive. Despite year-round
surveys in Great Smoky Mountains National Park
(GSMNP)—and especially over the last five years
during which time the Park has been the focus of
intensive surveys as part of its “All Taxon Biodiversity
Inventory —the moth escaped detection. Yet
Lithophane joannis is among the Park's most common
lepidopterans in middle elevation cove forests.
On 19 May 2001 I collected two Lithophane larvae
crawling up the trunk of a small yellow buckeye tree
(Aesculus flava Ait.) (Hippocastanaceae) while
collecting moths at a sheet and mercury light (with
Doug Ferguson), above the Chimneys Campground
(1000m) in Great Smoky Mountains National Park,
Sevier County, Tennessee. The caterpillars looked
similar to those of the innominata group (e.g., L.
hemina Grote, L. patefacta (Walker), L. petulca Grote
and L. innominata (Small), and others), but different
enough to raise doubt. Based on the host association
and phenotype, Dale Schweitzer guessed that the larvae
were those of Lithophane joannis. Return trips to the
same pullout along Newfound Gap Road in 2002, 2003,
and 2004, yielded additional examples of the
Lithophane. Typically, only one or two caterpillars were
collected each year. Unfortunately, I failed repeatedly
to rear examples through to the adult stage-
inappropriate foliage was offered or larvae were lost
during the obligatory, four-month prepupal diapause
common to Lithophane and other xylenines. In 2005,
while light trapping at the same site above the
Chimneys Picnic area, I thoroughly searched the same
4m yellow buckeye tree that had yielded caterpillars in
every year previous. Nine Lithophane caterpillars were
found in 20 minutes of searching (by flashlight). The
larvae were feeding, perched on He underside of leaves,
or observed walking along the trunk, with the exception
of two larvae that were recovered from within leaf
shelters. Both of these latter individuals were in the
process of molting.
Returning to the same area two days later (20 May,
2005), I happened upon a buckeye tree with numerous
leaf shelters. Upon opening the first, I found a last instar
Lithophane. Searching this same tree I counted more
than 20 additional Lithophane joannis caterpillars in less
than 10 minutes by opening other leaf shelters. Nearly
every shelter had a caterpillar and some two (few if any
of these were in the process of a molt). No additional
VOLUME 60, NUMBER 3
caterpillars were obtained by beating limbs of the same
tree over a large queen- sized bed sheet. In late
September, a series of Lithophane joannis issued from
this collection (Fig. 1).
The larva of Lithophane joannis is strongly mottled
(Fig. 2). There is often a straw to yellow tint where
adlnoca segments overlap and/or a tan to straw flush to
the middorsal and lateral stripes. The white dorsal
pinacula (D1 and D2 setae) are edged with black; both
dorsal pinacula are often embedded in a diffuse dark
patch that is best developed over the eighth abdominal
segment. The well-differentiated prothoracic shield is
heavily blackened above the subdorsal stripe. Below the
lateral stripe the subventer and venter are pale a
largely unmarked. The head bears a dark coronal bar, a
black spot within the frons (frontal triangle), and a black
bar above each antenna. Fully mature last instars are
about 4 cm in length. The middle and penultimate
instars are lime green, translucent, with a strong,
somewhat creamy spiracular stripe and a broken, w hite
middorsal stripe; the body bears numerous minute
white spots over the trunk (Fig. 3). In appearance the
last instars those of L. hemina and L.
innominata and may not be separable from them,
although most individuals will be rec sognizable by their
pale g arid color, especially those individuals that have
a pale green, yellowish, or steely blue cast. In most
instances, larvae of L. joannis will be identifiable by
their host association (with buckeye).
No other eastern Lithophane is known to consistently
resemble
take up residence in leaf shelters. Other members of
the innominata complex typically rest in bark crevices
by day (Wagner 2005) Trunks of Aesculus flava—
particularly on the understory trees where one can
expect to find larvae in numbers—are often smooth and
without fissures in larvae could conceal
It is not clear if L. joannis ever spins its
own shelters or only uses those of other leps. In May
which
themselves.
2006 a collection of 13 additional larvae was made from
the Chimneys area of the Park—all came from leaf
shelters of microlepidopterans. Eight were collected
from abandoned and pupal shelters made by
Choristoneura fractivittana (Clemens) (Tortricidae) and
the remainder from prepupal and pupal shelters spun
by Yponomeuta multipunctella Clemens!
(¥: ponomeutidae).
Several Lithophane are known to be both predatory
and cannibalistic, including the apparently closely-
related L. patefacta (Schweitzer 1979; Wagner, 2005).
While there would seem to be clear advantages to
‘While Yponomeuta multipunctella larvae normally feed on
Euonymus, since 2004 I have increasingly noted larvae on other
hosts in the Smokies.
Fics 1-3. Lithophane joannis: all from Chimneys area of
Great Smoky Mountains National Park, Sevier County, Ten-
nessee. 1, Reared adult. 2, Lithophane joannis last instar. 3,
Penultimate instar.
having the ability to take over previously spun shelters,
cannibalism was not observed in five pint rearing
containers that housed 3-5 middle and late instars. And
as noted above, I occasionally found shelters with two
larvae. Similarly, 10 of 13 larvae collected in 2006 came
from microlepidopteran — shelters
above)—no evidence of predation was noted in these
occupied (see
(or any of the other shelters opened on the day of the
initial collection in the Park).
In the middle elevation cove forests of GSMNP
caterpillars of Lithophane
discovered, the insect is among the most abundant
where joannis were
noctuid caterpillars—on 20 May, 2005, L. joannis was
arguably the most common noctuid caterpillar present
in the C himneys area. At the type locality in Ohio, L.
joannis outnumbered all other members of the genus
Lithophane at bait (Covell and Metzler 1992).
Interestingly, adults ignored the light traps that were
run at the same location (Eric Metzler pers. comm.). L.
joannis provides a noteworthy case of apparent rarity—
176
if one were to depend on standard light trapping
methods one would conclude that the moth is among
the rarest lepidopterans in eastern North America.
However if one employs bait or searches for caterpillars
one could conclude just the opposite, that L. joannis is
among the most common noctuids in Appalachian
forests where its foodplant, Aesculus flava, grows in
abundance.
Identification of the adults was confirmed by Eric
Metzler. Vouchers of both larvae and adults have been
deposited at the University of Connecticut; adults have
also been deposited at the United States National
Museum.
James Adams, Dale Schweitzer, and Bo Sullivan offered
suggestions on an earlier draft of the paper and Rene Twarkins
assisted with the larval images.
Journal of the Lepidopterists’ Society
60(3), 2006, 176-178
JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY
LITERATURE CITED
COVELL, JR., C. V. AND E. H. METZLER. 1992. Two new species of
moths ieee ie: Acronictinae, Cuculliinae) from midland
United States. J. Lepid. Soc. 46; 220-232.
SCHWEITZER, D.F., 1979. Predatory behavior in Lithophane quer-
quera and other spring cate rpillars. J. Lepid. Soc. 33: 129-134.
WacNeER, D. L. 2005. Caterpillars of Eastern North America, Prince-
ton University Press.
Davip L. WAGNER Department of Ecology and
Evolutionary Biology, of Connecticut,
Storrs, Connecticut USA, E-mail:
david.wagner@uconn.edu
University
06269,
Received for publication 7 November 2005; revised and accepted 9
June 2006
DIURNAL HERBIVORY DOCUMENTED FOR SPEYERIA IDALIA (NYMPHALIDAE) LARVAE ON
VIOLA SAGITTATA (VIOLACEAE) IN PENNSYLVANIA
Additional key words: regal fritillary, violet, foraging
Only two extant populations of the regal fritillary,
Speyeria idalia Drury (Nymphalidae), are documented
east of Indiana (Barton 1996 for Pennsylvania, Hobson
1999 and Chazal 2002 for Virginia). The larger of the
two populations occurs inside National Guard Training
Center-Fort Indiantown Gap (NGTC-FIG), an
approximately 6,925-ha military base located in south-
central Pennsylvania. Comprehensive descriptions of
the old-field successional habitats occupied by S. idalia
at NGTC-FIG are presented in Barton (1996) and TNC
(2001). Morphologic and genetic evidence indicates that
eastern populations may deserve specific or subspecific
status and designation as an evolutionary significant unit
(Williams 2001a, 2001b, 2002). In light of the
conservation status of S. idalia, research is warranted on
its life history.
Nocturnal foraging on Viola species has been
reported or referenced for S. idalia larvae by Holland
(1898), Ferris & Brown (1981), Opler & Krizek (1984),
Schull (1987), Royer (1988), Iftner et al. (1992), Royer
& Marrone (1992), and West (1998). However, Kopper
et al. (2001) documented diurmal feeding on V.
pedatifida G. Don (Violaceae) in three out of 12 S.
idalia larvae observed in Kansas. At NGTC-FIG, Barton
(1995) reported diurnal movements of S. idalia larvae
and noted the predominance of V. sagittata Aiton
relative to the presence of other Viola species but did
not describe larval foraging behavior.
Because S. idalia larvae have been challenging to
locate in the field across the species' range (Scudder
1889 for New England, TNC 2001 for Pennsylvania,
Kopper et al. 2001 for Kansas, Debinski pers. com. for
Iowa), behavioral observations of larvae have been
difficult to obtain (Kopper et al. 2001). A combination of
factors such as low population density (Barton 1995),
small body size, solitary distribution, cryptic coloration
and behavior (Stamp & Wilkens 1993), high mortality
rates (Mattoon et al. 1971, Wagner et al. 1997), and
concealing vegetation may par tially explain the modest
numbers of field-documented larvae. Previous surveys
conducted at NGTC-FIG to detect larvae have resulted
in very small sample sizes (n = 9; Barton 1995) or failure
(n = 0; TNC 2000, 2001).
On May 14, 2001, one S. idalia larva
unintentionally discovered at the Pennsylvania site
during a vegetation study. Shortly thereafter, a
qualitative survey of selected er asslands, known to be
inhabited by S. idalia adults alanine previous years, was
performed in an attempt to detect more larvae.
Typically conducted between 0900 and 1600 hrs, the
survey followed a generalized protocol: searching for
individuals and groups of V. sagittata (including arrow-
and ovate-leaved varieties), inspecting violets for
evidence of strip-feeding herbivory (typical of S. idalia),
and visually scanning violets and the surrounding area
for larvae. Images of S. idalia larvae in Allen (199 7) and
Richard & Heitzman (1987) assisted with positive
species identification.
Was
VOLUME 60, NUMBER 3
177
TABLE 1. Spatio-temporal and behavioral data for S. idalia larvae observed at NGTC-FIG, Pennsylvania, May 2001. Spatial
locations have been normalized by subtracting the coordinate
values of the first sighting.
Spatial location
Diurnal feeding on
Behavior at time of discovery 8
“leaves of V. sagittata
badividual Date Time (UTM meters)
larvae
Northing Easting
1 5/14 1200 0.0 0.0
2 5/18 0945 2617.3 5049.6
3 5/18 1102 2618.4 5047.5
4 5/22 1200 361.8 TPs)
5 5/24 1110 BOOED 6057.0
6 5/25 1440 2705.6 5041.1
motionless; on bare ground® observed in field
motionless; < 30 cm from Viola observed in captivity
motionless; on Viola not observed
motionless; on Viola observed in captivity
feeding on Viola observed in field
feeding on Viola observed in field
°distance to nearest Viola was unrecorded for larva 1
Five additional S. idalia larvae were opportunistically
discovered after more than 30 observer-hours of search
effort (Table 1). Digital photographs were taken of each
larva encountered and of the habitat in the immediate
vicinity of each sighting. Spatial coordinates of larvae
were determined through Global Positioning System
(GPS) technology (hardware: Pro XR Trimble receiver
unit and TSC1 Asset Surveyor Trimble datalogger;
software: Pathfinder Office version 2.80) and expres
in Universal Transverse Mercator (UTM) meters.
Larvae were not marked so as to avoid potential
handling effects. Because newly and_ previously
identified larvae were indistinguishable, duplication was
possible but unlikely, due to the relatively long distances
between sightings on preceding days [i.e., ~80
(minimum), ~7000 m (maximum)]|. The two larvae
observed simultaneously on May 18 were separated by
Fic. 1. Images of two S. idalia larvae diumally foraging on V. sagittata in the field (a)
Pennsylvania, May 2001.
approximately two meters.
Diurnal foraging on V. sagittata was documented in
five of the six larvae observed (Table 1 and Figure 1), a
strong trend despite the extremely small sample size.
Similar to those reported in Kopper et al. (2001),
feeding bouts were short-lived. S. idalia larvae were
Tied to consume only leaves of V. sagittata, either
partially or completely. In addition to foliar herbiv ory,
Beattie & Lyons (1975) and Kopper et al. (2001)
reported floral consumption of Viola spp. by the larvae
of Argynnis spp. and S. idalia, respectively. Such
behaviors, as well as nocturnal foraging bouts, may have
been exhibited by larvae at the NGTC-FIG site but
were not observed. Nonetheless, given the observations
presented here for Pennsylvania, in conjunction with
those for Kansas by Kopper et al. (2001), diurmal
foraging by S. idalia larvae may be more prevalent than
atari
and in controlled conditions (b) at NGTC-FIG,
178
previously described in the scientific literature.
Drs. Brian Kopper and Barry Williams (University of Wis-
consin), Dr. Roger Latham (Continental Conservation), and
Joseph Hovis (Pennsylvania Department of Military and Veter-
ans Affairs or DMVA) offered insightful comments on a preced-
ing draft. John Emmett, NGTC-FIG GIS Analyst, provided the
Trimble GPS receiver and datalogger for field use and assisted
with geospatial data management. Special thanks go to the
Pennsylvania Army National Guard (PAARNG) for granting ac-
cess to field sites and vehicles for on-base transportation. This
project was sponsored by the PAARNG (Cooperative Agree-
ment # DAHA36-01-2-9001), and funding was provided by the
Pennsylvania DMVA. The content of the information presented
does not reflect the position or policy of the U.S. Government,
and no official endorsement should be inferred.
LITERATURE CITED
ALLEN, T. 1997. The butterflies of West Virginia and their caterpillars.
University of Pittsburgh Press, Pittsburgh. 385 pp.
Barton, B. 1995. Report on the life history of the regal fritillary
(Speyeria idalia) and interspecific competition with other
Speyeria species. Unpublished report to the U.S. Department of
Defense. 34 pp.
1996. Final report on the regal fritillary, 1992-1995, Fort
Indiantown, Annville, Pennsylvania. Unpublished report to the
U.S. Department of Defense.
Beattig, A., & N. Lyons. 1975. Seed dispersal in Viola (Violaceae):
adaptations and strategies. Amer. Jour. Bot. 62(7): 714-722.
CuazaL, A, 2002. Status survey of the regal fritillary (Speyeria idalia)
in 2002 on the Radford Army Ammunition Plant. Natural Her-
itage Technical Report 02-20. Virginia Department of Conserva-
tion and Recreation, Division of Natural Heritage, Richmond. 20
oF
Ferris, C., & F. Brown. 1981. Butterflies of the Rocky Mountain
States. University of Oklahoma Press, Norman. 442 pp.
Hopson, C. 1999. Conservation status assessment for the regal
fritillary (Speyeria idalia) in Virginia. Natural Heritage Technical
Report 99-25. Virginia Department of Conservation and
Recreation, Division of Natural Heritage, Richmond. 23 pp.
HoLLanp, W. 1898. The butterfly book. Ist edition. Doubleday and
McClure. New York. 382 pp.
IrtNeR, D., SHUEY, J., & J. CALHOUN. 1992. Butterflies and skippers of
Ohio. Bulletin of the Ohio Biological Survey 9(1). The Ohio lepi-
dopterists research report No. 3. Ohio State University, Colum-
bus. 212 pp.
Kopper, B., Marcoutes, D., & R. CHARLTON. 2001. Notes on the
behavior of Speyeria idalia (Drury) (Nymphalidae) larvae with
implications that they are diumal foragers. J. Lep. Soc. 54(3):
96-97.
Mattoon, S., Davis, R., & O. SPENCER. 1971. Rearing techniques for
species of Speyeria (Nymphalidae). J. Lep. Soc. 25(4): 247-256.
OPLER, P., & G. KriZeEk. 1984. Butterflies east of the Great Plains: an
illustrated natural history. Johns Hopkins University Press, Balti-
more. 294 pp.
RICHARD, J., & J. HEITZMAN. 1987. Butterflies and moths of Missouri.
Missouri Department of Conservation, Jefferson City. 385 pp.
Royer, R. 1988. Butterflies of North Dakota: an atlas and guide.
Minot State University Science Monograph No. 1. 192 pp.
Royer, R., & G. MARRONE. 1992. Conservation status of the regal frit-
illary (Speyeria idalia) in North and South Dakota, a report to
United States Department of the Interior, Fish and Wildlife Serv-
ice, Denver. 51 pp.
SCHULL, E. 1987. The butterflies of Indiana. Indiana Academy of Sci-
ence, Indiana University Press, Bloomington/Indianapolis. 179
1212
SCUDDER, S. 1889. Butterflies of the eastern United States and
Canada with special reference to New England. Published by the
author. Cambridge. Vol. 1, pp. 1-776; vol. 2, pp. 777-1774; vol. 3,
pp. 1775-1958.
JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY
Stamp, N., & R. WILKENS. 1993. On the cryptic side of life: being un-
apparent to enemies and the consequences for foraging and
growth of caterpillars, pgs. 283-330. In Stamp, N., & T. Casey
(Eds.), Caterpillars: ecological and evolutionary constraints on
foraging. Chapman & Hall, New York.
[TNC] THE NaTurRE ConsERVANCY. 2000. Population monitoring and
life history studies of the regal fritillary (Speyeria idalia) at Fort
Indiantown Gap Military Reservation, Annville, Pennsylvania: ac-
tivity summary and report of findings (January — December
1999). Prepared by The Nature Conservancy. Unpublished re-
port to the Pennsylvania Department of Military and Veterans Af-
fairs.
. 2001. Population monitoring and life history studies of the regal
fritillary (Speyeria idalia) at Fort Indiantown Gap National
Guard Training Center, Annville, Pennsylvania: activity summary
and report of findings (January — December 2000). Prepared by
The Nature Conservancy. Unpublished report to the Pennsylva-
nia Department of Military and Veterans Affairs.
WAGNER, D., WALLACE, M., BOETTNER, J., & J. ELKINTON. 1997. Sta-
tus update and life history studies on the regal fritillary (Lepi-
doptera: Nymphalidae), pgs. 261-275. In Vickery, P., Dunwiddie,
P., & C. Griffin (Eds.), The ecology and conservation of grass-
lands and heathlands in northeastern North America. Massachu-
setts Audubon, Lincoln.
West, P. 1998. Establishing long-term monitoring of the regal fritillary
(Speyeria idalia Drury) in Wisconsin. RJ/KOSE Report, WIFO:
regal fritillary monitoring. 12 pp.
WILLIAMS, B. 2001a. Patterns of morphological variation in Speyeria
idalia (Lepidoptera: Nymphalidae) with implications for taxon-
omy and conservation, Ann. Ent. Soc. Amer. 94(2): 239-243.
. 2001b. Recognition of western populations of Speyeria idalia
(Nymphalidae) as a new subspecies. J. Lep. Soc. 55(4): 144-149.
. 2002. Conservation genetics, extinction, and taxonomic status: a
case history of the regal fritillary. Cons. Biol. 16(1): 148-157.
P. MooreEsIDE!, D. ZERCHER?, AND P. MCELHENNY.
The Nature Conservancy, Fort Indiantown Gap Office,
Department of Military and Veterans Affairs,
Environmental’ Unit, Building 11-19, Annwille, PA
17003.
Received for publication 28 March 2003; revised and accepted 26
June 2006
‘Current address: American Association for the Advancement of
Science, 1200 New York Avenue, NW, Washington, DC 20005. E-
mail: pmooresi@aaas.org
*Current address: The Nature Conservancy, Peoria Office, 301 SW
Adams Street, Suite 1007, Peoria, IL 61602.
Book REVIEW
Journal of the Lepidopterists’ Society
60(2), 2006, 179-180
THE GEOMETRID MOTHS OF EUROPE (A
Hausmann, ed.), Volume 2. Sterrhinae. Axel Hausmann.
2004. Apollo Books, Stenstrup, Denmark. 600pp.
Hardback ISBN 87-88757-37-4. Priced at 140 Euros on
http:/Avww.pensoft.net/notes/12090.stm.
This volume represents another _ significant
contribution to a series on European Geometridae that
will be seen in retrospect as perhaps the most significant
milestone in our understanding of that fauna. Not only
does it provide a comprehensive review of our
knowledge to date, coupled to an immense amount of
fresh research based on exhaustive examination of
copious material, but it probably comes at a turning
point in the way taxonomic information is presented,
with the emphasis moving more towards the electronic
media, away from print on paper. The editorial preface
indicates the w ay the volume is embedded within the
context of existing or imminent initiatives in electronic
informatics for Geometridae. But, as someone who
sometimes wonders whether the rush to embrace the
electronic age may be more headlong than judicious,
given our lack of experience of the security/vulnerability
of the internet to all sorts of factors, I am reassured to
know that there are hard-copy products such as this to
fall back on under any worst case scenario.
The Sterrhinae, after the Larentiinae, are probably
the most successful subfamily of the Geometridae in
their proportional representation at temperate latitudes,
though the Ennominae may have higher numerical
representation. They also contain, after Ewpithecia in
the Larentiinae (covered in Vol 4 of this series), the two
next most species-rich geometrid genera: Scopula and
Idaea. All three genera pose emcee difficulties in
identification of their species, so publication of these
two volumes has removed this impediment for the
European fauna.
The preface notes the strong (particularly western)
Mediterranean focus of the subfamily in a European
context, relative to the more even distribution of the
Larentiinae, but this aspect is not explored to any great
extent in the main text, though all the data are there to
enable interested readers to analyse it for themselves.
For example, there is endemism at a generic level,
albeit from genera that are monotypic such as
Anthometra (eastern Mediterranean) and Emmiltis
(central Mediterranean), or Oar, if regarded as distinct
from Scopula, with just two species in the
Mediterranean. The five species of Cleta are also
essentially Mediterranean, and also the genus
Glossostrophia. Brachyglossina includes major N.
African and Levantine species groups, with one species
in Spain considered to be an outlier of the latter group.
Apart from these smaller genera, the Mediterranean
focus is seen to an extreme in Idaea, where over two-
thirds of the species are Mediterranean, 73% of these
restricted to the western part. In Scopula, as in the
larentiine Eupithecia, only one third are Mediterranean,
and those are more evenly distributed across that
region.
The book reviews extensively the recent taxonomic
advances made for the subfamily, but there is still not a
full consensus on the generic classification, as indicated
by Sihvonen (2005, Nota lepid., 28: 70-71) in his review.
There is also discussion of relationships of the
Sterrhinae within the family as a whole, where there is
growing consensus that the Sterrhinae and Larentiinae
are the most basal groups, though relationships between
the two lineages of Sterrhinae and the Larentiinae still
need further study.
All species are copiously illustrated by color plates
showing the range of variation, often accompanied by
diagnostic half-tone figures in the body of the text.
There are clear line-drawings of male and female
genitalia, and some stereoscans of critical features of
antennae and legs. As with other volumes, the
description of the adult is headed as a diagnosis, and the
true diagnosis is found as an account of differences of
similar species. The text for each species includes
extensive reviews of available biological and ecological
information, drawing on the breadth of literature and
current expertise on the fauna. Most species have larvae
that are herbaceous feeders, those of Idaea tending to
prefer dried or withered foliage. However, the larvae of
the genus Cyclophora are ar boreal defoliators.
The systematic checklist at the end of the book also
lists, with asterisks, species from neighboring areas, a
particularly useful addition given the Mediterranean
nature of the group, enabling the reader to appreciate
how the diversity extends to N. Africa, Turkey or the
Levant. Species from Madeira and the Canaries are also
listed. The distributions of all species are illustrated by
maps with shaded areas indicating the general range
within which are placed black circles or other sy mbols
indicating more precise localities from which material
180 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY
has been examined. JEREMY HOLLOway Entomology Department, Natural
The book brings together an immense and diverse History Musem, Cromwell Road, London, SW7 5BD,
amount of information in a clear and cohesive manner — UK Email: j.holloway@nhm.ac.uk
that will retain its value for many years to come. It is
therefore an investment well worth making.
COVER PHOTOS SOLICITED
The Journal solicits high-quality color photographs for consideration
as cover illustrations. The photographs may illustrate any aspect of
Lepidopteran biology.
Digital image files saved to a CD are preferred. Submit digital files
ain a erties orientation of approximately 1770 pixels by 2200 pixels
at a resolution of at least 120 pixels/cm in °.tif format. If you wish to
submit printed photographs, please ensure that they are vertical in
orientation (approximately 5.5 inches wide X 6. 75 inches long
or 14 cm wide x 17 cm long;).
Date of Issue (Vol. 60, No. 3): 2 October 2006
EDITORIAL STAFF OF THE JOURNAL
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|
|
wi
Five New Species or Paucivena Davis, 1975 (Lepiporrera: TINEOIDEA: Psycuipar) From Cupa
Rayner Nunez Aguila ----------------------------------------------------------------------------------
A Remarkas_e New Riopinip Species, STALACHTIS HALLOWEEN! (RIODINIDAE: STALACHTINI), FROM
Mount Ayancanna, Guyana Jason P.W. Hall ---------------------------------------------------------
A New Species or Auraronota (LEPIDOPTERA: TORTRICIDAE: CHLIDANOTINAE) FORMERLY CONFUSED
with A. Hyprocramma (Meyrick) John W. Brown----------------------------------------------------
A Revision or Psotorrera ButLer, INCLUDING A REDESCRIPTION OF Its KNowN Species (ARCTIIDAE:
ARCTINAE: EUCHROMINI) Rebecca B. Simmons------------------------=------2=-===—-==5-5= see
Forest Tent CatTerpittaR: Martinc, OvireosiTion, AND ADULT CONGREGATION AT Town Licuts
A New Species or Evcosma Hisner (TorrricipazE: OLETHREUTINAE) FRoM THE TaLL Grass PRAIRIE
Recion or Mipwestern Nortu America. Donald J. Wright ---------------------------------------
Monarcu (Danaus Prexippus L. NyMPHALIDAE) Micration, Nectar REsourCcES AND FIRE REGIMES IN
THE Ouacuita Mountats or Arkansas. D. Craig Rudolph, Charles A. Ely, Richard R.
Schaefer, J. Howard Williamson, and Ronald E. Thill --------------------------------------------
GENERAL Notes
OpsERVATIONS OF KricoconiA LysiDE (PIERIDAE) IN THE FLoripa Krys Mark H. Salvato, John
V. Calhoun, and Holly L. Salvato ------------------------------------------ 2-00-00 nonnnn
A Precautionary Tae apouT Rarity: On THE Larva AND Lire History or LirHopHaNe JOANNIS
(Lepmoprtera: NocturpaE) David L. Wagner -----------------------------------------------=-=------
DiurnaL Heresivory DocuMENTED For SpeyerIA IDALIA (NYMPHALIDAE) LARVAE ON VIOLA
SAGITTATA (VIOLACEAE) IN PENNSYLVANIA P. Mooreside, D. Zercher, and P. McElhenny ---------
Book Review
Tue Geometrip Morus or Europe Jeremy Holloway ------------------------------------------------------
@ This paper meets the requirements of ANSI/NISO Z39.48-1992 (Permanance of Paper).
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