BULLETIN OF THE ALLYN MUSEUM
THE ALLYN MUSEUM OF ENTOMOLOGY
Number 50 6 December 1978
REVISION OF THE EUPTYCHIINI (SATYRIDAE)
4. Pindis R. Felder
Lee D. Miller
Curator, Allyn Museum of Entomology
This paper is the fourth in a series of generic reviews (L. Miller, 1972, 1974, 1976)
and another for which sufficient material exists in North American collections. The
coverage of this genus at this time is governed more by convenience than by systematic
relationship it is hoped that the final paper in this series will be a compilation of the
available data toward a systematically correct checklist of the species in the tribe and a
phylogenetic interpretation of their interrelationships.
The present study is based upon 356 specimens from a cross section of the range of
the single species that apparently belongs to Pindis. Additionally I have examined the
type specimens or photographs thereof of the three names applied to that species. The
sources of material are abbreviated in the same manner as in previous parts, and these
acronyms are as given in L. Miller (1972: 2 and 1974: 1). Only newly cited references are
given in the appended biliography, but if the references cannot be found there, they
may be in one of the previous parts of the revision.
There is only one species in Pindis, and it is rather variable. The possibility that all
of the available names could be applied was examined in light of the plasticity of
phenotypes, but the genitalia, which also vary slightly, do so without regard to
phenotypic variations. The entire picture, then, is one of a single, variable species
strongly affected by environmental conditions.
One other euptychiine has been placed in Pindis by Lewis (1973: 235; pl. 63, fig. 16),
pellonia Godman and Salvin, but the male genitalia of pellonia have nothing in common
with those of Pindis squamistriga, the type of the genus. In the case of "Euptychia
pellonia the gnathos arms are attached, as in Paramacera and Cyllopsis (L. Miller, 1972
and 1974, respectively), whereas those of Pindis are free. The proper placement of "E."
pellonia must await a later portion of the revision, and it will not be discussed further
The discussion that follows here will be longer than usual because of the degree of
variability displayed by the single species involved.
Pindis R. Felder, 1869
Pindis R. Felder, 1869: 475. Type-species: Pindis squamistriga R. Felder, 1869: 475;
The distinctness of Pindis from other euptychiines is attested to by its being one of
the half-dozen genera into which Weymer (1910-1911) placed all members of the tribe.
This distinctiveness has not always been accepted, however, because Godman and
Salvin (1879-1901) did not separate P. squamistriga from other "Euptychia", despite
their usual predilection for generic splitting in other groups.
Pindis has no readily apparent close relatives in the tribe. Superficially it may be
confused only with Cyllopsis clinas (Godman and Salvin), as indicated by L. Miller
(1974: 95-96) in an earlier part of the revision. This resemblance is not a clue to relation-
ship, however; perhaps a better case might be made with P. squamistriga and
"Taygetis" kerea Butler, but even this relationship is not very close (see "Phylogenetic
and Zoogeographic Considerations" section).
Since it displays the free gnathos arms, Pindis may be considered to be a "main
line" euptychiine, but this in no way should be construed to mean that it is "typical".
The wing shape, pattern and male genitalia (exclusive of the gnathos) are quite atypical
in the group, and it stands as an aberrant member of the tribe, perhaps as much so as
Paramacera or Cyllopsis. This genus, then, is another of the "old Mexican" ones and
stands virtually alone within the Euptychiini.
It may be characterized as follows:
Eyes naked. Antenna about one-third length of forewing costa; club indistinct, oc-
cupying terminal third of antenna and inflated less than twice width of shaft. Palpus
(Fig. 3) about half again length of head, semiporrect with tips slightly divergent; ter-
minal segment just under half length of second; hairs of second segment long, greater
than three times width of segment.
Thorax densely clothed with hairs dorsally and ventrally. 6 foreleg (Fig. 4) slender,
tibia slightly longer than femur and tarsus nearly as long as either, slender and pointed.
9 foreleg (Fig.5) slender and miniaturized (nearly as much so as that of 6), with slender
pentamerous tarsus spined on the second, third and fourth segments. Mid- and
hindlegs short and slender, weakly spined on tibia and with well-developed tibial spurs.
Wing venation (Fig. 1) of a typical euptychiine pattern, but the wing shape is not
(see discussion below). Forewing radial veins arising from cell in three branches; veins
Rs and M, well separated at their origins; Sc and cubital stem noticeably, 2A only
slightly, inflated at their bases, as shown; cells of both wings about half lengths of
wings to ends of M,; Cu, of both wings somewhat nearer Ms than Cu,; hindwing humeral
vein strongly developed and distinctly curved distad toward apex.
There is a large androconial patch (Figl) occupying the area in the cell of the fore-
wing from the base of Cu2 to the origin of Rs, thence continuing outside the cell to
M,-Cu, (the most distal area included) and down to 2A, occupying the middle third of
that cell. The individual scales are rather like those of the last three genera covered in
the revision (L. Miller, 1972, 1974, 1976), differing from them chiefly in the rib structure
6 genitalia (Figs. 12-14) as illustrated and characterized by the free gnathos arms,
the heavy shieldlike tegumen and especially the heavy, downturned uncus. The valvae
are relatively unornamented, and the penis is rather longer than that of most Eup-
tychiini covered to date.
9 genitalia (Fig. 15) also as illustrated, simple, lightly sclerotized (except in the
sterigmal region) and only barely diagnostic.
To my knowledge nothing has been written concerning the early stages of Pindis.
Such information would be of great interest, especially for determination of relation-
ships with other genera within the tribe, relationships that are presently obscure at
The single species is highly polymorphic with respect to wing shape and color pat-
tern, but the terminalia are far more constant. The wing shape varies between two ex-
tremes: The "normal", or truncate-winged morph (Figs. 1, 16-20, 28-29), is a basically
smaller, more dull-colored insect with more uniformly and completely developed pat-
terns of spots and lines; the acute-winged morph (Figs. 2, 22, 24-26, 32-39) is a redder in-
sect with greater pattern above and a reduced, frequently heavily obscured pattern
below, and it is a larger butterfly. Two of the available names were based upon the
truncate-winged morph, squamistriga R. Felder and obnixa Draudt, whereas zabdi
(Butler) was based on a specimen approaching the acute-winged form. It is interesting
to note with the acute-winged morph that the production of the forewing takes place in
interspace Mi-M,, rather than along either of the veins (Fig. 2).
Figures 1-5: Pindis squamistriga R. Felder. 1, 5 venation; truncate-winged morph.
2, outline of outer margin of wing of acute-winged morph. 3, palpus. 4, & foreleg. 5, 9
foreleg. Unless otherwise indicated, all specimens illustrated from Allyn Museum col-
Pindis squamistriga R. Felder, 1869
Pindis squamistriga R. Felder, 1869: 475 ("Huahuapam" [perhaps Huajaapan de
Leon, Oaxaca], Mexico). Holotype in BM.
Euptychia zabdi Butler, 1869 [1869-1874]: 11 ("Chaetum" recte Choctum), Guatemala.
Holotype in BM.
Pindis obnixa Draudt, 1931: 104 (Guerrero, Mexico; here designated). Lectotype in
BM, here designated (see below).
In parts of its range P. squamistriga appears to be a highly variable species.
Elsewhere the populations are remarkably monomorphic. This variability suggested
that more than one species might be involved, and the male genitalia of the first few of
each extreme morph examined appeared to be somewhat different than those of super-
ficially unlike ones, but dissection of a series demonstrated that the genitalia, like the
faces, varied slightly between individuals without apparent regard to other characters.
In fact, even geographic subspeciation has not taken place in P. squamistriga, and all of
the observed variation must be due to seasonal and/or climatic conditions. The
redescription that follows, therefore, will be perhaps a bit longer than for previous eup-
Variations in wing shape: The specimen used to illustrate the venation of the genus
(Fig. 1) represents the most frequently encountered morph, what I earlier called the
"truncate-winged morph". Other specimens range from this wing shape to the one il-
lustrated in Fig. 2, the "acute-winged morph", in which the following modifications of
the wing shape may be noted: 1) the forewing is produced, especially in M,-M2, with
smaller projections as far costad as on R,; 2) the crenulations at th ends of hindwing
veins Cu,, Cu, and 2A are far more pronounced than in the truncate-winged form; and 3)
the forewing costa is somewhat more arched. These characters are by no means
diagnostic, though, since they intergrade more or less independently from one extreme
to the other. Some specimens are clearly intermediate, and these may well be flying
with either or (rarely) both of the extremes. The shape as well as the size of the
specimens is usually correlated with the brightness or dullness of the pattern and with
development or reduction in the size and number of the hindwing ventral ocelli. The
acute-winged morph is more frequent in specimens before me from southern Mexico,
and at first I thought they might represent a distinct subspecies, but additional
material from Guatemala and El Salvador proved to be indistinguishable from
truncate-winged specimens from central Mexico. Apparently, then, these observed
phenotypic differences are due to environmental vicissitudes and are not genetic in
Male: Head, thorax and abdomen thickly clothed with warm brown dorsal and
gray-brown ventral hairs. Antenna dark brown above, reddish-brown below; club
reddish-brown on both surfaces, slightly darker at the tip. Palpus with dorsal brown
hairs, elsewhere grayish-brown, slightly darker along ventral margin. Legs clothed with
dense grayish hairs.
Wings above (truncate-winged morph) uniformly dull, dark brown; acute-winged
morph slightly paler, lightest in distal third of forewing, this area delimited from inner
two-thirds by a dark line roughly paralleling the end of the cell, thence produced basally
and continued to inner margin; a second submarginal dark brown line parallel to outer
margin. Fringes tan, slightly reddened in acute-winged morph.
Under surface of forewing (truncate-winged morph) generally warm, dark brown
with two darker marginal lines, a darker, slightly thickened submarginal line and a
poorly developed extradiscal line from the costa to M, paralleling the end of the cell,
thence slightly inset and continued to 2A. In more acute-winged males the ground color
is significantly reddened; the extradiscal band better developed, dark brown and exten-
ding to the inner margin; the marginal line missing altogether in some specimens,
especially those that are heavily overscaled with gray distad; and in these specimens
the submarginal line is obscured toward the costa. All intergradations may be noted
between the extremes in long series.
Hindwing below (truncate-winged morph) usually gray-brown (occasionally slightly
laved with reddish outside cell) with a narrow transcellular, slightly curved dark brown
line, an irregular dark brown extradiscal band or line, produced distad in M,-M,, two
marginal and one submarginal dark brown lines roughly following the outline of the
wing margin and black ocelli from Rs-M, to Cu,-2A between the extradiscal and sub-
marginal lines (the ocellus in Rs-M, often absent, and any of the ocelli may be pupilled in
white, most frequently those in M,-M, and Cu,-Cu,); the entire surface heavily scrawled
with gray-brown lines. The acute-winged morph is often much redder, the extradiscal
and transcellular bands much more thickened, the marginal lines frequently absent (the
submarginal one only faintly and irregularly indicated) and the ocelli are often absent
(more commonly indicated by small black points of a few scales each); occasionally the
ocellus in Mi-M, is indicated only by a small white patch with the other ocelli absent or
very poorly developed. The fringes of both the fore- and hindwings below are tan-gray,
somewhat redder in the acute-winged morph.
Figures 6-11: Type specimens. 6-7, Pindis squamistriga R. Felder, Holotype 6 up-
per (6) and under (7) surfaces; MEXICO: "Huahauapam" (see text) (Allyn Museum
photos 100477-C-1/2). 8-9, Euptychia zabdi Butler, Holotype d upper (8) and under (9)
surfaces; GUATEMALA: Choctum (Allyn Museum photos 091677-B-20/21). 10-11, Pin-
dis obnixa Draudt, Lectotype 6 (see text for designation) upper (10) and under (11) sur-
faces; MEXICO: GUERRERO (Allyn Museum photos 091677-B-26/27). All specimens
in BM collection.
The males examined had forewing lengths of between 19.7 and 25.5 mm., averaging
22.4 mm. The truncate-winged males measured 19.7 to 24.9 mm., averaging 21.76 mm.,
whereas the most acute winged ones measured 21.8 to 25.5 mm., averaging 24.1 mm.
The measurements of some representative populations are summarized in Table I.
The 8 genitalia are as figured (Figs. 12-14) for the holotypes of squamistriga and
zabdi and the lectotype of obnixa, respectively.
Female: The truncate-winged form varies little from its a, differing chiefly in the
following: upper surface slightly laved with red in a few specimens; upper forewing with
more pronounced extradiscal and submarginal bands delimiting a somewhat paler area
between them; under surface just the same, but forewing slightly more likely to have a
faint reddish shading in distal half. The acute-winged form is just the same as in the 8,
but the reddish color on both surfaces is more pronounced and the confused pattern of
the hindwing below is even more so in this sex.
Lengths of forewings of the females at hand range from 20.8 to 28.3 mm., averaging
24.6 mm.; the acute-winged females range from 20.9 to 28.3 mm., averaging 25.7 mm.,
and the forewings of the truncate-winged ones range from 20.8 to 26.1 mm, averaging
24.3 mm. The forewing length measurements for females are summarized in Table I.
9 genitalia as illustrated (Fig. 15): the sterigma is the only part that is strongly
sclerotized, and the genitalia themselves are not especially characteristic.
Figures 12-14: & genitalia of type specimens of synonyms of Pindis squamistriga
R. Felder. Specimens are the same ones figured in Figs. 6-11, and the same data apply.
12, P. squamistriga; slide M-1172 (Lee D. Miller). 13, E. zabdi; slide 13,167 (BMNH). 14,
P. obnixa; slide 13,168 (BMNH). All specimens and slides in BM collection.
I have seen 356 specimens, 205 males and 151 females, from the following localities
and collections. Not all of these have been used in the analyses that follow.
MEXICO: SINALOA: Loberas Summit, 5 mi. NE Potrerillos, 1820 m., viii, 19 (A).
JALISCO: viii, 2 19 (A); 8.7 mi. W Magdalena, 1380 m., viii, 1 (A); Guadalajara,
2 39 (AMNH); La Cumbre de Autlan, 3200-4200 ft., 6 79 (AMNH).
MICHOACAN: vic. Pozos de Ixtlan, 1520 m., viii, 1 (A); El Sabino, nr. Uruapan, vii,
1 (A)' Santa Lucrecia, ix, 2 3 (A). MEXICO: Tenancingo, iii, vi, 3 3 1 9 (A); Malinalco,
viii, 63 29 (A). DISTRITO FEDERAL: Mexico, viii, 19 (A). HIDALGO: Jacala,
4500', vi, 29 (A). MORELOS: Cuernavaca, xi, 3 (A, BM, including "9" Paralec-
totype of Pindis obnixa); TepoxtlAn, 1650 m., vii-x, 78 3 85 9 (A); 10 mi. S Cuernavaca,
1100 m., viii, 1 3 9 (A); 5 mi. S Amacuzac, 900 m., viii, 2 6 4 9 (A); 1 mi. S Xochicalco
ruins, 1400 m., viii, 1 2 9 (A); Yautepec, vii-viii, 5 3 1 9 (A); Taxco, ix, 3 3 (JBS); Te-
jalpa, ix, 1 19 (JBS). GUERRERO: "Guerrero", 1 (Lectotype of Pindis obnixa);
Omilteme, x, xi, 2 S (A); Acahuizotla, vi, viii, 6 6 79 (A); Chilpancingo, vi, ix, 2 3 19
(AMNH, JBS); Huacapa Canyon, S of Petaquillas, 1020 m., viii, 19 (A); Tierra Col-
orada, viii, ix, 27 S 3 9 (A); 2 mi. W Colotlipa, 1020 m., viii, 3 C 1 9 (A); 2 mi. N of El
Treinte, 220 m., ix, 3 d (A); 5 mi. N of El Playon, ix, 2 9 (A). PUEBLA: Izucar de
Matomoros, ix, 2 6 (A); "Matomoros" (same as preceding?), ix, 1 9 (AMNH); Caltepec,
x, 13 (AMNH). VERACRUZ: Jalapa, 2 (CM); Presidio, vi, 16 (A). OAXACA: Hua-
japan de Leon, ix, 39 (A); Chimalapa, v, 19 (A); San Jose Pacifico, Mpio. Rio Hondo,
2400 m., xi, 14 6 14 9 (A). NOT LOCATED: Almoloya, vi, 18 (A); "Huahuapam" (?=
Huajapan de Leon, Oaxaca), 1 & (Holotype of Pindis squamistriga; BM); "Mexico", 2
GUATEMALA: "Guatemala", 1 (CM); Choctum (corrected from "Chaetum"),
1 6 (Holotype of Euptychia zabdi; BM).
EL SALVADOR: Tamanique, 1000 m., ix, vii, x, xi, 14 29 (A).
NO LOCALITY: 1 29 (A).
It was fortunate that when my wife was in England last autumn she was able to
locate and photograph the type specimens of all three names presently carried in the
synonymy of the present species. These specimens are figured in Figs. 6-11, and a brief
discussion of them is here in order.
Pindis squamistriga (Figures 6, 7, Holotype 3; 12 3 genitalia of Holotype) was
described by R. Felder from a single specimen captured by Hedemann at
"Huahuapam". This locality is not on any of the maps I have consulted, but I suspect
that it might have been a misspelling of Huajapan de Leon in Oaxaca, a locality from
whence I have seen additional material. Such errors are common, even in modern
Figure 15, Pindis squamistriga R. Felder, 9 genitalia; MEXICO: MEXICO: Tenan-
cingo; slide M-3728 (Lee D. Miller).
material. The specimen, along with most of the Felders' material, passed to the British
Museum (Natural History) via the Rothschild bequest. The Felder collection contained
one other specimen of squamistriga, a 6 taken considerably later at Cuernavaca,
Morelos. Both specimens are of the truncate-winged morph and are fairly well ocellated
below, as shown in the figures.
Euptychia zabdi (Figures 8,9, Holotype 6; 13, a genitalia of Holotype) was based
on a single a in the Godman and Salvin collection from Choctum, Guatemala. In-
advertently Butler gave "Chaetum" as the type-locality, an error that was rectified by
Godman and Salvin (1880 [1879-1901]: 81). As those authors state, squamistriga and
zabdi were proposed within a few weeks of one another, and the only means by which we
may be certain that the former name has precedence is that Butler (1869-1874: Correc-
tions) states that Felder's name should be given precedence over his own. The Holotype
6 is a large, reddish insect that approaches, but is not as extreme as, the "acute-winged
morph" alluded to here. The hindwing maculation on the ventral surface is very dif-
Figures 16-27: variations in under surfaces of selected males of Pindis squamistriga
R. Felder. 16, MEXICO: GUERRERO: Tierra Colorada. 17, MEXICO: MORELOS:
Tepoztlan. 18-19, EL SALVADOR: Tamanique. 20-24, MEXICO: MORELOS:
Tepoztlin. 25, MEXICO: MEXICO: Tenancingo. 26, MEXICO: MEXICO: Malinalco.
27, MEXICO: MORELOS: TepoztlAn. Allyn Museum photos 072178-2 through
072178-13, respectively. Figures 16-39 all taken at same magnification, so size com-
parisons are as shown on plates.
ferent from squamistriga, but the & genitalia, as shown, are almost identical.
Pindis obnixa (Figures 10, 11, Lectotype 6; 14 6 genitalia of Lectotype) was
described from "a pair" of specimens, both of which are now in the British Museum
(Natural History) collection. Draudt's 9 type is actually a somewhat worn 6 with
the androconial patch obscured. It is from Cuernavaca, Morelos. Draudt's 6 type is in-
To i. 3, J A
.a' -" .. '. .,-'* & .. '
Figures 28-39: variations in under surfaces of selected females of Pindis
squamistriga R. Felder. 28, MEXICO: JALISCO: Ameca. 29, MEXICO: MORELOS:
Tepoztlfi. 30, MEXICO: OAXACA: San Jose Pacifico. 31, MEXICO: MORELOS:
TepoztlAn. 32-33, MEXICO: OAXACA: San Jose Pacifico. 34, MEXICO: MORELOS:
Tepoztlan. 35, MEXICO: MEXICO: Tenancingo. 36, MEXICO: MORELOS:
Tepoztlin. 37, MEXICO: OAXACA: San Jos6 Pacfico. 38-39, MEXICO: MORELOS:
Tepoztlan. Allyn Museum photos 072178-13 through 072178-19; 072178-A-1 through
072178-A-6, respectively. Figures 16-39 all taken at same magnification, so size com-
parisons are as shown on plates.
deed that and is from "Guerrero". 1 should have preferred to have made the Cuer-
navaca specimen the Lectotype, all other things being equal. But they are not equal:
the Cuernavaca specimen was mislabelled as a 9; it is in rather poor condition and the
old "Cuernavaca" label could cover a multitude of places in which the insect flies. Ac-
cordingly, then, I am designating the other specimen (Draudt's original & type) as the
Lectotype of Pindis obnixa. It bears the follwoing labels: 1) a round, red bordered BM
"Type" label; 2) a label with the printed number "54"; 3) a black bordered label stating
"Mexico/ GUERRERO / ix.12" (Italicized entries = handwritten); 4) a partly printed,
partly handwritten pinkish label, "Type/ Pindis/ obnixa/ S Draudt" and 5) a printed
label, "Brit. Mus/ 1934-239" referring to the BM accession. To this I have added a red
label hand printed in black, "Lectotype 5/Pindis obnixa /Draudt/ designated by /Lee
D. Miller, 1978". The other syntype is a Paralectotype and will be labelled accordingly.
The types of obnixa are very little different superficially from the type of squamistriga
(perhaps a bit more heavily ocellated), both belonging to the truncate-winged morph.
The androconial scales (Figs. 40-42) are not appreciably different from those of some
other Euptychiini, but the ribbing structure is quite different. Just what taxonomic
significance to attach to these scales must await further studies in the tribe: the con-
figurations are presented here simply for comparison purposes.
P. squamistriga is a species of mesic woodland environments in the Mexican
localities in which I have encountered it. It behaves very much like other open woods
euptychiines, but perhaps its flight is a bit more erratic than others. The adults will
land on the ground with the wings about half open and bask in dappled sunlight. When
approached, however, they will snap the wings together and heel over slightly to one
side, thus effectively cutting down on their visibility by minimizing shadows. When
disturbed the flight is rather rapid (for one of the Euptychiini) and very jerky; their
habit of darting in and out of the brush makes them difficult to capture.
Variation: A series of specimens of either sex will yield specimens that vary greatly
in under surface facies (Figs. 16-39). The specimens that are like the type of
squamistriga are heavily ocellated on the hindwing, and the ground color is a rather
uniform gray-brown. From that extreme one may observe specimens with an almost un-
marked grayish ventral hindwing (Fig. 31, for example), others with strong reddish
flush and either patterned (Fig. 34) or unpatterned (Fig. 36). The ocelli on these forms
are usually reduced or absent, but a few specimens (Fig. 20) will show development of
the ocelli rivalling that of the truncate-winged morph.
All or some of the ocelli may be pupilled in white (Fig. 28), as in the type of
squamistriga (Fig. 7), but in other specimens all ocelli are blind (Fig. 29, for example). In
a very few specimens the ocelli, especially the anterior ones, may be entirely
represented by white blotches with no black rides (Fig. 27 is an extreme example).
The dorsal surface is relatively invariable, individuals differing chiefly in the
amount or lack of reddish suffusion. The submarginal and extradiscal lines of the under
40 / 4 1~4
Figures 40-42: androconial scale of Pindis squamistriga R. Felder. 40, base of scale,
1500x (Allyn Museum SEM 0432). 41, detail of tip of scale, 7500x (Allyn Museum SEM
0430). 42, detail of ribbing near base of scale, 5000x (Allyn Museum SEM 0429).
forewing are usually better developed in the zabdi-type morph, except where the costa
is heavily overscaled with gray.
The wing shape differences between the various morphs have been discussed
earlier, and it suffices to say that intergrades between the extremes are not uncommon-
ly found flying with examples of other morphs. Similarly, size differentials exist be-
tween individuals, and as shown in Table I, a case could be made for separating some
populations from one another. Taking into consideration the presence of several wing-
shape and pattern morphs within a single population, however, it was determined that
the observed size differences were more likely to be environmentally controlled
(foodplant quality, etc.) than genetically related. Hence, the decision not to subdivide P.
squamistriga into geographic races.
I do not expect P. squamistriga to be taken much further south than the central El
Salvador locality recorded herein. There is a possibility that it might be discovered in
northwestern Honduras, but the fact that it has not been collected in southern
Guatemala or southeastern El Salvador suggests that the Tamanique locality is about
the furthest possible southern extent of this species. Furthermore, it is unlikely that
squamistriga will be found in southwestern New Mexico or southeastern Arizona: both
of these areas have been collected rather extensively without uncovering the present in-
sect. The fact that it was taken on Loberas Summit, Sinaloa, in company of such
Arizona residents as Neophasia terlooti (Behr) and Pyrrhopyge araxes arizonae God-
man and Salvin at least suggests the possibility (no matter how remote) that
squamistriga might occur in the mountains of southern Arizona.
PYLOGENETIC AND ZOOGEOGRAPHIC CONSIDERATIONS
No other euptychiine closely corresponds with P. squamistriga superficially or in
genitalic configuration. "Taygetis" kerea and Megeuptychia antonoe (Westwood) have
somewhat similar genitalia, but even here the relationship is not very close. It appears,
therefore, that squamistriga is another of the "old Mexican" species that stands apart
from its relatives, thus signifying long isolation. Historical geography gives us
evidence that the western part of Mexico was an island through much of the Tertiary,
and this isolation at about the right time gives a plausible explanation for such a
divergent genus as Pindis. Again, this faunal area has produced many strangely evolv-
ed butterflies, Baronia brevicornis Salvin (Papilionidae), Prestonia clarki Schaus
(Pieridae) and Phanus rilma Evans (Hesperiidae) being only examples.
It appears then that the ancestor of Pindis reached this Guerrero-Oaxaca
"refugium" long ago and developed in situ. This progenitor of Pindis was not the same
as the ancestor of Cyllopsis or Paramacera, since both of those genera have gnathos im-
Table I: forewing length measurements in millimeters for selected populations of
Pindis squamistriga R. Felder. S. D. = standard deviation; S. E. = standard error; N =
number of individuals sampled.
N Range Mean S. D. S. E.
all 6 147 19.7-25.5 22.41 1.301 0.107
Mexican & 130 19.7-25.5 22.35 1.368 0.120
Guerrero 15 19.7-23.9 21.83 1.275 0.329
Jalisco 6 11 20.0-25.1 23.35 1.470 0.443
Oaxaca 6 16 21.0-25.5 24.08 0.995 0.249
Morelos 6 67 20.0-23.9 22.34 0.910 0.111
Mexico (st.) d 9 21.7-25.1 23.48 1.618 0.540
El Salvador S 14 21.3-24.7 22.96 1.050 0.281
all 9 112 20.8-28.3 24.62 1.506 0.142
Mexican 9 110 20.8-28.3 24.62 1.513 0.144
Morelos 9 59 22.1-26.1 24.32 0.937 0.122
Oaxaca 9 18 20.9-28.0 25.68 1.956 0.461
El Salvador 9 2 23.6-25.7 24.65
movably placed, rather than free as in the present genus. The western Mexican area
must have supported a moderately well developed butterfly fauna during this time,
much of which has survived (doubtless in modified form) to the present. Pindis spread
from this ancestral home further than did many other members of the "old Mexican"
fauna, but it did not move as far north as did Paramacera, concentrating its dispersal in
areas further to the south.
While little of substance can be said about the relationship between Pindis and
other Euptychiini, it is hoped that these will become clearer as the study progresses.
I am grateful to the owners and curators of the various collections in which Pindis
were found for access to these specimens. Special thanks go to Messrs. R. I. Vane-
Wright and P. R. Ackery of the British Museum (Natural History) for allowing and
assisting my wife's photography of the types of the various names and for allowing me
to dissect the genitalia of these specimens.
During her stay at the British Museum (Natural History) my wife and colleague,
Jacqueline, took time from her busy schedule of the study of the Castniidae to locate
and photograph the type specimens; she also read and commented upon this
manuscript. I owe her a great debt of gratitude for these and other chores cheerfully
performed on my behalf. Thanks also are due A. C. Allyn, Director of this institution,
for photographing additional specimens, reading and commenting on the manuscript
and for other favors not otherwise acknowledged.
If a paper is not listed herein, reference to it will be found in L. Miller (1972, 1974,
Draudt, M., 1931. Neue Mexicanische Tagfalterformen. Ent. Rundschau, 48: 103-104.
Lewis, H. L., 1973. Butterflies of the world. Chicago, Follett: xvi + 312 pp.; 208 pls.
Miller, L. D., 1976. Resvision of the Euptychiini. 3. Megisto Hiibner. Bull. Allyn
Mus., (33): 1-23; ill.
`-r' Y -^
v v ,.,^ \ ;
Figure 43: distribution of Pindis squamistriga R. Felder.