Title: Bulletin of the Allyn Museum
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Permanent Link: http://ufdc.ufl.edu/UF00079423/00031
 Material Information
Title: Bulletin of the Allyn Museum
Series Title: Bulletin of the Allyn Museum.
Abbreviated Title: Bull. Allyn Mus.
Physical Description: v. : ill. ; 23 cm.
Language: English
Creator: University of Florida. News Bureau.
Allyn Museum of Entomology
Florida State Museum
Florida Museum of Natural History
Publisher: The Museum
Place of Publication: Sarasota Fla
 Subjects
Subject: Entomology   ( lcsh )
Genre: government publication (state, provincial, terriorial, dependent)   ( marcgt )
 Notes
Dates or Sequential Designation: Began in 1971.
Issuing Body: Vols. for <1985>- issued by the Florida State Museum; <1988>- by the Florida Museum of Natural History.
General Note: Separately cataloged in LC before no. 48.
General Note: Description based on: No. 4, published in 1972; title from caption.
General Note: Latest issue consulted: No. 123, published in 1988.
 Record Information
Bibliographic ID: UF00079423
Volume ID: VID00031
Source Institution: University of Florida
Holding Location: University of Florida
Rights Management: All rights reserved by the source institution and holding location.
Resource Identifier: oclc - 01451276
lccn - 87643372
issn - 0097-3211

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BULLETIN OF THE ALLYN MUSEUM

3621 Bayshore Rd.
Sarasota, Florida 34234

Published By
Florida Museum of Natural History
University of Florida
Gainesville, Florida 32611
Number 126 5 April 1989
ISSN-0097-3211



A NEW OLIGOCENE FOSSIL BUTTERFLY, VANESSA
tAMERINDICA (LEPIDOPTERA: NYMPHALIDAE),
FROM THE FLORISSANT FORMATION, COLORADO

Jacqueline Y. Miller
Assistant Curator, Allyn Museum of Entomology/Florida Museum
of Natural History, 3621 Bay Shore Road, Sarasota, FL 34234
and
F. Martin Brown
Research Associate, Allyn Museum of Entomology/Florida Museum of
Natural History, and 6715 S. Marksheffel Road, Colorado Springs, CO 80925
Introduction

Numerous fossil insects, including 10 butterflies, have been excavated from the
Florissant Lake Bed Shales, which date from the early Oligocene, approximately 35 million
years ago. Recently two butterfly specimens were located and upon further examination
were determined to represent a new fossil species. The first fossil is the best preserved
and was one of the fossil specimens presented as gifts to tour guides leading groups through
the area. The specimen was given to F. Martin Brown by the late Fred Bischoff and was
collected in the early 1930's from the southwestern portion of the present Florissant Fossil
Beds National Monument. The area was later excavated by E. R. Nelson, Jr. for the
Milwaukee Public Museum, and although Nelson did not find any further butterflies, he
did locate some interesting moths.
The second specimen is incomplete and was found some 3.12 km due north of the
Florissant Fossil site on the old Stoll Ranch. This area is adjacent to the much older but
now back-filled site originally excavated by George Wilson and H. F. Wickham. The
specimen was recovered in 1981 by Frederick Sanborn.

Phylogenetic Relationships

Determining the phylogenetic relationship between these two butterfly specimens and
modern taxa has proved to be interesting. The outline of the forewing apex is similar
to that of tBarbarothea florissanti Scudder (1892), a libytheid, the type of which has been
lost or misplaced. While both specimens under study are comparable in size to this taxon
(24-27 mm), the presence of a labial palpus with the distal segment approximately one-
third the length of the second segment argues that this family should not be considered











as a possible relative.
The wing veination provided clues to the taxonomic placement of these new fossil
butterflies. The apical forewing venation in the two present specimens is markedly different
from the recently described species, tOligodonta florissantensis Brown (1976). The
branching of forewing radial veins and the presence of R, effectively eliminated the family
Pieridae. The possibility that these two fossils are related to the family Hesperiidae was
discarded due to the branching of the forewing apical veins. The faint indication of the
hindwing humeral vein effectively eliminated the Lycaenidae, and with the staggered origin
of hindwing medial veins, the Riodinidae were also rejected. Since there was no evidence
of the basal swollen forewing veins nor any indication of 3A, the butterfly families
Satyridae and Papilionidae were also disregarded from further consideration.
The only remaining butterfly family is the Nymphalidae, and one of the diagnostic
features of this family, the presence of reduced forelegs, is evident on the first fossil. The
produced apex of the forewing with the characteristic lobe at M,-M, is indicative of the
genus Vanessa. In 1965, Nekrutenko described two fossil species from Miocene deposits
in the northern Caucasus, one in the genus Vanessa, Vanessa lkaraganica, a fossil that
is actually associated with the modern genus Aglais. The second species described was
Pyrameis (= Vanessa) tfossilis. The proportional size and venation of the present fossils
are comparable to that described in the genus Pyrameis (= Vanessa) by Nekrutenko;
however, the produced lobe at the forewing apex and the smooth transition along the
lateral margin posteriad to the anal angle, in addition to the distinctive wing pattern
elements, differentiates these two fossils from those taxa previously described. However,
the wing venation, produced apical area of the forewing in conjunction with certain
forewing pattern elements in these fossils is reminiscent of the Old World taxon Vanessa
indica and is herein described.

Vanessa famerindica, new species

Figs. 1-4

This description is based on examination of two fossil specimens, both preserved in
shale with the ventral surface exposed: (1) a specimen which is relatively complete with
four wings and body, and (2) a partial specimen comprised of the distal three-fourths of
two left wings. In both specimens the hindwings are superimposed over the forewings.
The principal portion of this description is based on fossil 1 (Figs. 1, 3) with comparative
wing venation verification in fossil 2 (Fig. 2, 4).
The squashed condition of the body makes the examination and study of body parts
difficult. The outline of the eyes, and in particular the inner eye ring which encircles the
ommatidia proper, are recognizable. Neither of the antennal shafts nor the antennal clubs
have been preserved. The anterior margin of the head is outlined in prominent setal tufts.
There is a single labial palpus preserved in the lateral view along the ventral prothorax.
The distal segment is approximately one-third the length of the second segment, but the
present position of the palpus gives no indication as to whether it is erect or porrect in
relation to the labrum and frons. The ventral surface of all three segments is clothed heavily
in setae and scales. There is no indication of the proboscis.
The legs have been relatively well preserved in the first fossil specimen. The forelegs
are located in the normal position and reduced in size, a characteristic of the family
Nymphalidae. The femur and tibia (in part) of the meso- and metathoracic legs can be
easily distinguished on the left half of the thorax. The outline of the femur of the meso-
and metathoracic legs on the right thoracic half is barely discernable. Posteriad along
the abdominal midline approximately at segments A3-6, the preserved tibial and tarsal
segments of the right metathoracic leg are evident. In addition, the tibial and tarsal
segments of the mesothoracic leg are located just to the left and anterior to the
metathoracic leg. The distal margins of these legs are outlined heavily in setae. However,
there are no prominent setae nor scales outlining the distal leg margins associated with
the thorax proper.










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I t









(Ayn Museum Photo no. 880308/36A).
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Fiur 1. Veta iwo ooyeVnsa"a eidcn.s.(a cl m
{Ally Musum Poto o. 8030836A)


Figure 2. Ventral view of Paratype, Vanessa tamerindica, n. sp. (Bar scale = 0.5 cm)
(Allyn Museum Photo no. 880309/10).











There are 10 distinct abdominal segments with individual scales evident on the posterior
margin of each segment, particularly on segments A7 and A8. There are no discernible
sclerotized structures preserved on segment A10, but the irregular intersegmental
development of the posterior margin of segment A7 indicates the presence of sclerotized
structures, possibly the female sterigma.
The wing venation of this new fossil taxon is illustrated in Figures 3 and 4. The left
forewing of the first specimen is the most complete and best preserved with a forewing
costal length of 27 mm. Just to the left of the head and anterior to the base of the left
forewing, there is a separate, preserved portion of either the tegula, or more likely, the
base of the forewing costa. R,-R, branch approximately 13.5 mm from the apex or about
three-fourths of the forewing length. R, and R, are well separated with the origins
respectively at 16.5 mm and 14.5 mm from the apex. The end of the forewing cell occurs
at less than one-half of the forewing length with the cell open. The origins of M, and M2
are incomplete in both fossils but the origin of M3 is apparent. The relative positions of
M,, M,, and M3 are equidistant from each other along the lateral margin. Because of the
preservation of the specimen on the ventral surface, the base of the forewing cubitus and
anal veins cannot be distinguished.
The arrangement of the veins on the hindwings is rather well preserved in both fossils.
The veins illustrated are those which are easily discerned, with those interconnecting faint
veins indicated by the broken line. The relative position of the veins has been somewhat
distorted during preservation. There is a faint indication of a humeral vein at the wing
base in fossil 1 (Fig. 3). M, arises approximately 5.8 mm from the base. The origin of
M, is near that of M, but the interconnecting portion of the medius is absent. The hindwing
cell is also open. Cu, arises approximately 9 mm from the base with M3 markedly arched.


























3

Figure 3. Wing venation of Vanessa tamerindica, Holotype, ventral view, drawn from
the original fossil. (Scale = 0.5 cm)











The relative positions of veins M,, M2, and M, are equidistant from one another along
the lateral margin on the left hindwings of both fossils. Veins 2A and 3A are present
in fossil 1 with both veins compressed along the lateral margins of the abdomen. The
lateral margins of the hindwings are incomplete in fossil 1 but relatively complete in fossil
2, with numerous scales and setae of the hindwing fringes distinct in both fossils.
One diagnostic feature is present on the lateral margin of the forewing. The whole area
of the wing from the apex is markedly angled and produced to MI-M, with the remainder
of the lateral margin slightly tapered toward the anal angle. These features are particularly
evident on the right forewing margin of fossil 1, and these, taken in conjunction with
the wing venation, are similar to the modern Palearctic taxon Vanessa indica (Figs. 5,
6). The apical portion of the forewing is absent in part on the left forewing (fossil 1) in
M,-M2 and not quite as produced in fossil 2.
With the hindwings superimposed over the forewings in both fossils, the wing pattern
elements, especially those of the hindwings, are not easily distinguished. However, there
is a reduced forewing apical band and an enlarged irregularly shaped spot at the end of
the forewing cell near the orgin of R3 evident in both fossils. These pattern elements coupled
with the produced forewing apex and lateral margin are reminiscent of the modern taxon,
Vanessa indica (Fig. 6) as compared with the more complex pattern elements of V.
virginiensis (Fig. 7) or the subtle patterns of V. cardui (Fig. 8). These latter taxa are widely
distributed in North America today. Therefore, the name amerindica, for this nearctic
fossil reflects its systematic position near Vanessa indica, which presently has a range
throughout the Palearctic and Indomalayan zoogeographic regions.


Figure 4. Wing venation of Vanessa tamerindica, Paratype, ventral view, drawn from
the original fossil. (Scale = 0.5 cm)











Holotype (Fossil 1) (Figs. 1, 3): A fossil with four wings and body compressed, preserved
in shale with the ventral surface exposed. Forewing length, 27 mm and that of the hindwing
is approximately 24.5 mm. Colorado, Florissant Fossil Beds National Monument, T13S
R71W, Section 26 and bears the following typed gummed label, "Holotype of Vanessa
amerindica, n. sp., J. Y. Miller and F. M. Brown, 1988 (UF 21999)." Paratype (Fossil 2)
(Figs. 2, 4): a specimen with the greater portion of the left wings preserved in shale and
with the ventral surface exposed. Colorado, Stoll Ranch, N. Florissant Fossil Beds National
Monument (T13S R71W NE.25, Section 3) and bears the following typed gummed label,
"Paratype, Vanessa amerindica, n. sp., J. Y. Miller and F. M. Brown, 1988. (UF 22000)."
Both the Holotype and Paratype are deposited in the invertebrate palaeontological
collections of the Florida Museum of Natural History and are on indefinite loan to the
Allyn Museum of Entomology.
In his description of two new taxa in the genera Vanessa (=Aglais) and Pyrameis
(= Vanessa), Nekrutenko (1965) based his findings not only on wing pattern and venational
elements but also on a set of comparative morphometric measurements between the origin
of hindwing veins and the hindwing cell. While these measurements provided a convenient
method with which to compare modern taxa with the potential ancestors, such
measurements are relative and vary with preservation of the fossil specimen. Such size
variation may be affected by a number of other ecological factors at the time of
preservation. Despite the imperfect preservation of these fossils, including the absence
of the interconnecting vein between M, and M2, measurements of the easily discerned


R3

R4
MI
M2
M3
Cu1

Cu2


Figure 5. Wing venation of Vanessa indica. (Scale = 0.5 cm)










veins of V amerindica are similar in position and relative proportion to those of V. indica
(+0.2-0.5 mm).

Discussion

It is difficult to determine relationships of extant butterfly taxa with any degree of
certainty through the examination of incomplete 35 million year old fossils. The usual
diagnostic features, such as the dorsal and ventral wing pattern elements and genitalia,
used in standard taxonomic treatments are either incomplete or unavailable for further
examination in fossils. In addition, our knowledge of the fossil record in Lepidoptera and
particularly that of butterflies, is very limited with only 26-28 specimens described.
Previous descriptions of fossil butterflies from the Oligocene Florissant Shales included
a number of taxa with relationships found in modern neotropical butterfly taxa, such
as tBarbarothea florissanti Scudder, Hypanartia (tProdryas) persephone Scudder (1892),
Chlorippe twillmetae Cockerell (1913), and tOligodonta florissantensis Brown (1976). Even
a castniid moth, tDominickus castnioides Tindale (1985) has been described from this
site. This new species, Vanessa famerindica is the first described taxon from the Florissant
shale deposits closely aligned with extant species from the Holarctic and Indomalayan
geographical regions. Vanessa is today a nearly cosmopolitan butterfly genus with a broad
geographical range throughout the Holarctic to New Zealand, through central and southern
Africa, to Central and South America, and east even to the small island of San Juan de
Fuca east of the Falkland Islands. Such a broad geographical range and conspecificity
is not completely unknown within the Nymphalidae (Miller and Miller, in press) and is

I I










6 I-
6






*r' :"./ .i uife




7 ^ '

Figures 6-8. Ventral view of three extant taxa within the genus Vanessa: (6) V. indica,
(7) V. virginiensis, and (8) V. cardui. (Scale = 1 cm)











found in other butterfly families (Higgins and Riley, 1970; Higgins, 1975). In addition,
species within the genus Vanessa are notably migratory, a fact which may explain in part
the presence of this fossil in the Florissant shales.
Durden and Rose (1978) described two new fossil taxa in the Papilionoidea from the
Middle Eocene deposits of the Green River Shales in Colorado, and these discoveries
markedly altered our conception of the age of butterflies. While none of these Middle
Eocene fossils were precisely comparable to extant taxa, certain morphological attributes
placed one species, tPraepapilio colorado, in an intermediate sister subfamily between
the Papilioninae and the Parnassinae and their more synapomorphic counterparts, the
Baroniinae (Durden & Rose 1978). While the taxonomic relationships of tPraepapilio
colorado have been the subject of further refinement and interpretation (Hancock, 1983;
J. S. Miller, 1987), clearly this fossil evidence supports the hypothesis that the Papilionidae
are more than 48 million years old! Most of the early records for the Lepidoptera date
from the lower Cretaceous (Whalley, 1977, 1978), the period during which the angiosperms
arose. Thus, most of the lepidopteran evolutionary history apparently took place sometime
between the early Cretaceous and middle Oligocene.
The advanced morphological attributes for the genus Vanessa (Nymphalidae) are
represented in fossils from both Miocene and these significant Florissant Oligocene
deposits. This fossil evidence renders the previously held position that the most of extant
butterfly species and genera date from the Pleistocene untenable and indicates an earlier
origin. Indirect evidence from phylogenetic studies in other groups suggests a similar
evolutionary history. This is illustrated dramatically in the recent description of a stingless
bee, Trigona prisca (Michener and Grimaldi, 1988), discovered in Cretaceous New Jersey
amber, but significantly still another species referable to a contemporaneous genus. The
evolutionary history of the Lepidoptera is far from completely known and further
investigation of the higher classification of the butterflies in relation to other Lepidoptera
is warranted.

Acknowledgments

We would like to thank Frederick Sanborn for donating the Paratype of Vanessa
tamerindica for further scientific study. Special thanks are due to Drs. John C. Downey,
Douglas Jones, and Lee D. Miller for their comments and critical review of the manuscript.

Literature Cited

Brown, F. M. 1976. Oligodonta florissantensis, gen. n., sp. nov. (Lepidoptera: Pieridae).
Bull. Allyn Museum (37):1-4.
Cockrell, T. D. A. 1913. Some fossil insects from Florissant, Colorado. Proc. U. S. National
Museum, 44: 341-346.
Durden, C. A. and H. Rose. 1978. Butterflies from the Middle Eocene: The earliest
occurrence of fossil Papilionoidea (Lepidoptera). Texas Mem. Mus., Pearce-Sellards series,
(29):1-25.
Hancock, D. L. 1983. Classification of the Papilionidae (Lepidoptera): A phylogenetic
approach. Smithersia (2): 1-48.
Higgins, L. G. 1975. The classification of European butterflies. Collins, London, 320 pp.
Higgins, L. G. and N. D. Riley. 1970. A field guide to the butterflies of Britain and Europe.
Collins, London, 380 pp.
Michener, C. D. and D. A. Grimaldi. 1988. A Trigona from late Cretaceous amber of New
Jersey (Hymenoptera: Apidae: Meliponinae). Amer. Mus. Novit., 2917: 1-10.
Miller, J. S. 1987. Phylogenetic studies in the Papilioninae (Lepidoptera: Papilionidae).
Bull. Amer. Mus. Nat. Hist., 186:365-512.
Miller, L. D. and J. Y. Miller. Nearctic Aglais and Nymphalis (Lepidoptera, Nymphalidae):
Laurasia revisited? The Entomologist (in press).
Nekrutenko, Y. P. 1965. Tertiary nymphalid butterflies and some phylogenetic aspects
of systematic lepidopterology. J. Res. Lepid. 4(3):149-158.











Scudder, S. H. 1892. Some insects of special interest from Florissant, and other points
in the Tertiaries of Colorado and Utah. U. S. Geological Survey, Bull. 93: 21-24.
Tindale, N. B. 1985. A butterfly-moth (Lepidoptera: Castniidae) from the Oligocene Shales
of Florissant, Colorado. J. Res. Lepid., 24(1): 31-40.




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