BULLETIN OF THE ALLYN MUSEUM
THE ALLYN MUSEUM OF ENTOMOLOGY
Number 8 6 November 1972
REVISION OF THE EUPTYCHIINI (SATYRIDAE)
1. Introduction and Paramacera Butler
Lee D. Miller
Curator, Allyn Museum of IEntomology
At the time of my revision of the higher categories of the Satyridae
(Miller, 1968) it became apparent that the Euptychiini as so defined were
badly in need of a modern comprehensive treatment. The last attempt at an
arrangement of all of the species within the tribe was that of Weymer (1910-1911),
but his work was little more than a compilation of the species which had been
described to that time and was of little more value in the understanding of the
euptychiines than various papers by Butler (1866, 1867, 1868a, 1868b). Forster
(1964) substantially altered taxonomic thought on the relationships of those
species previously included in Euptychia and Taygetis, describing many new
genera and a few species. His work, unfortunately, was by no means complete
and was concerned chiefly with the Bolivian species.
This paper begins a series which will lead eventually to a monograph of the
Euptychiini as defined by Miller (1968). The sheer volume of names available in the
the tribe (nearly a thousand species-group names have been proposed) precludes
a single volume revision, if one is to make a comprehensive study. Individual
papers will appear as they become available on a genus-by-genus basis, without
regard to their final systematic positions within the tribe. The final paper in the
series will provide a checklist for the Euptychiini and the intergeneric phylogenetic
relationships, if such become apparent. For several reasons those genera that
impinge on the North American fauna will be among the first considered,
with the wholly Neotropical genera reserved for later treatment, generally. The
primary reason for dealing with the North American species and their
Neotropical congenors first is to make the results available to my colleagues
in this country as soon as possible. There are a limited number of well-defined
euptychiines in the United States and Canada, and with one exception these
species belong to quite distinctive Mexican and Central American genera.
Furthermore, enough material is available in collections in this country to do
meaningful revisionary work on these genera.
By contrast, many groups of wholly Neotropical euptychiines are very poorly
represented in our collections, and a great deal of material from foreign collections
will have to be examined in the future. Many taxa within the tribe have been
consistently misidentified and the names misapplied in the literature, based on the
early work, so the types of these species, almost all in European collections,
must be consulted before the chaotic nomenclature can be untangled. In other
instances, hopefully fewer than I fear, the type specimens have been lost or destroyed,
and neotypes will require designation, but this can be done only when the nonexistence
of authentic type material has been ascertained.
The genera and their associated species will be described in detail and pertinent
aspects illustrated where possible. Citations of the original descriptions,
type-localities and the present location of the types will be given. Distributional
data will be given for the material examined, and where possible and desirable
these data will be plotted on maps. Each genus will have a key to its species, and
where species are comprised of more than one subspecies, keys will be provided
for the discrimination of these taxa. In the final paper in the series a key to the
genera of the Euptychiini will be given.
In the future, as additional material and information become available,
"Addenda and Corrigenda" sections will be appended to papers referring not
to the paper at hand, but to previous papers in the series. Such sections will
appear irregularly as necessary.
With the exception of the usual male (Q) and female (9) symbols, only one set
of abbreviations will apear in the text. The source of material will be
abbreviated: for the present paper the following institutional and private collections
have provided material examined and are abbreviated as:
Allyn Museum of Entomology and the associated private holdings
of A. C. Allyn -- (A).
American Museum of Natural History, New York, N. Y. -- (AMNH).
Carnegie Museum, Pittsburgh, Pa. (CM).
Private collection of Eduardo C. Willing M., Merida, Yucatan,
Private collection of Kilian Roever, Phoenix, Ariz. (KR).
Los Angeles County Museum, Los Angeles, Calif. -- (LACM).
National Museum of Natural History, Washington, D. C. -- (USNM).
The genus Paramacera, an aberrant member of the tribe known only from
Mexico and the southwestern United States, is a convenient one with which to
begin the revision of the tribe. Additionally, the nomenclatorial problems
associated with Paramacera illustrate the problems inherent in the study of the
Euptychiini as a whole: too long erroneous identifications and designations
have been passed uncritically from author to author.
Paramacera Butler, 1868
Paramacera Butler, 1868a: 194. Type-species: Neonympha xicaque Reakirt, 1867
["1866"]: 336-337, designated by Scudder, 1875: 240.
=Paramecera Butler, 1868b: 98. Type-species: Neonympha xicaque Reakirt,
1867 ["1866"]: 366-337, by monotypy.
As noted by both Scudder (1875: 240) and Hemming (1967: 340), Butler
referred to, but did not describe, a manuscript specific name for the type of
Paramacera. Nothing would be gained by resurrecting this name at this time,
inasmuch as two specific names already have priority over it. The generic name
Paramecera was proposed as new, not as an emendation of Paramacera, a few
months after the latter name's proposition, hence, Paramecera should fall as junior
objective synonym of Paramecera. Various correspondents have pointed out
that since Paramecera was compared with the genus Amecera Butler in the
original description that perhaps Paramacera was the result of a typographical error.
Furthermore, since the descriptions in Butler's Catalogue (1868b) are more detailed,
it seems likely that Butler intended that paper to be the vehicle of description,
rather than the brief paper in Entomologists' Monthly Magazine (Butler, 1868a)
which actually appeared first. These correspondents may be right, but the fact
remains that both generic names were proposed as new, and Butler actually
designated two different, though now known to be synonymous, type-species
for them. Those writers who have subsequently mentioned the genus, including
Godman and Salvin (1879-1901), Weymer (1910-1911), Holland (1931),
Hoffmann (1940), dos Passos (1964) and Miller (1968), almost without
exception have used "Paramecera", and if the spellings of the generic names in
question were markedly different, Hemming's (1967: 340) validation of
Paramacera would be open to question. Such is not the case, however; the
spellings of the two names in question are similar and the pronunciations
virtually the same, so there is no reason for not retaining the name with priority
for this genus.
Superficially Paramacera resembles a number of Palearctic Maniolini, but
structurally (especially in the form of the 9 foreleg, the venation, etc.) members
of the present genus are definitely euptychiines (Miller, 1968: 89-95). The
"Euptychia" paeon (Godart) group of Euptychiini from the southern Neotropics
resemble Paramacera superficially, but these austral insects are very
different genitalically and are certainly not congeneric with the northern ones.
The present genus is characterized as follows:
Eyes sparsely hairy. Antennae about two-fifths length of forewing costa;
club occupying distal quarter of antenna and about three times width of shaft at
widest point. Palpi (Miller, 1968: fig. 207) long and more or less erect, second
segment about three times length of third; hairs of palpi up to three times
width of second segment.
Thorax heavily clothed with long hairs above and below. a foreleg (Miller,
1968: fig. 208) reduced and slender with a monomerous, unspined tarsus.
9 foreleg also reduced and slenderer than in most Euptychiini with pentamerous
tarsus spined on first four subsegments (Miller, 1968: fig. 209). Mid-and hindlegs
short, bearing terminal tibial spurs on both.
Wing venation (Miller, 1968: fig. 206) basically like that of most
Euptychiini, but with an unusually large androconial patch outside forewing
cell, beginning in anal cell, broadest in Cu2-2A and narrowing toward M3-Cu1
(occasionally with small satellite patches in Mi-M and M2-M:); forewing
radius three-branched (two-branched in most Euptychiini). Forewing margin
produced at apex in P. chinanteca, n. sp., less so in other species; hindwing
margin scalloped, especially in P. chinanteca.
& genitalia with shield-like tegumen, long, slightly curved, undivided uncus
and paired, tapered gnathoi which are not freely articulated. Saccus short.
Valvae laterally bowed and slender, bearing teeth at distal end. Penis relatively
straight and simple.
9 genitalia relatively simple in the two species in which these structures
are known and similar to one another. Ductus bursae somewhat more sclerotized
than in many Euptychiini.
Four species are recognized in this genus, three of which are new. These
butterflies are found from eastern Arizona to southern Mexico, but the populations
are isolated and generally restricted to mesic or xeric montane habitats.
The species prefer rather open coniferous grasslands; the larvae are no doubt grass
feeders, though noting is known of the life-history to my knowledge. A key
to Paramacera follows:
KEY TO THE SPECIES OF Paramacera BUTLER
1. Upper surface of forewing without ocelli, or at most a small one in
M,-M,; Oaxaca, Mexico .................................... chinanteca, n. sp.
1'. Upper surface of forewing with ocelli in M,-M2 and M:-CuI, occasionally
with a subsidiary spot in M2-M: . ...... ............. 2.
2. Upper surface of forewing with large, black androconial patch;
under surface of hindwing with pale discal band terminating before
costa and ocelli in hindwing spaces MI-M, and M2-M3 silvered without
black irises; Oaxaca, Mexico ................................. copiosa, n. sp.
2'. Upper surface of forewing with more nearly concolorous androconial
patch; under surface of hindwing with pale discal band continuing to
costa and hindwing ocelli in Mi-M2 and M2-M3 below with black
irises ..................................................................... 3 .
3. Upper surface of hindwing with marginal and submarginal lines reddish-
brown from anal angle to at least M:3; Mexico .............. xicaque (Reakirt)
3'. Upper surface of hindwing with marginal and submarginal lines fuscous
(rarely reddish-brown only at anal angle); Arizona and Chihuahua,
M exico ..... ........................................... allyni, n. sp.
Paramacera chinanteca, new species
Figures 1, 2 (5), 3, 4 (9), 5 (3 genitalia)
Male: Head, thorax and abdomen clothed with olive-brown hairs above,
dark grayish ones below. Palpus black, white at tip. Antenna brown ringed
narrowly with buff above, buff narrowly ringed with brown below; club brown
above, buff below with tip blackish-brown. Legs clothed with long black hairs
proximad, shorter buff ones on tibia and tarsus.
Forewing above olive-brown, fuscous along margin, with androconial patch
only slightly darker than ground color and with no ocelli present.
Hindwing above olive-brown, slightly darker than that of forewing, with
fuscous marginal and submarginal lines, minute dark brown blind ocelli from
M -M, to Cu,-Cus, dark extradiscal zigzag band, shaded rust-brown basad, and
brown patches mid-costa and marginally in Sc+R'Rs.
Forewing below as figured: warm tan, darker basally, with two straight
rust-brown median bands, two dark brown marginal lines, a crenulate
submarginal brown line and very small blind blackish ocelli with narrow brown
rings from M1-M, to M,,-Cui.
Hindwing below as figured: base dark brown peppered with white scales,
median band warm tan splotched with reddish-brown, extradiscal areas
shaded purplish and marginal and submarginal lines dark brown, reddish-brown
at anal angle.
Fringes of forewing above and below broadly fuscous at tends of veins,
narrowly white in interspaces. Fringes of hindwing above and below white, rather
broadly fuscous at ends of veins.
Length of forewing of Holotype a 19.5 mm., that of the single S Paratype
5 genitalia as figured, differing from those of other members of genus
particularly as regards configuration of tip of valva.
Androconial patch less well-defined than in other species because
distribution of androconial scales far less dense within patch and those present
scattered among regular scales. Shape of scale as in other species, but shorter
than that of P. copinsa.
Female: Head, thorax, abdomen (what was remaining) and appendages as in 8.
Upper surface as in 5, but forewing extradiscal area much paler and
bearing a very small blind brown ocellus in Mj-M,; hindwing ground color
paler than in 3 with markings more prominent and brown mid-costal patch only
Under surface also as in 8, but extradiscal area of forewing much paler.
Lengths of forewings of three 9 Paratypes 21.0, 21.0 and 22.0 mm.
The females all lacked at least distal portion of abdomen, so 9 genitalia
Described from five speciments, two males and three females, from the
high mountains of Oaxaca, Mexico.
HOLOTYPE c: MEXICO: OAXACA: Muo Cu6u (Cerro Pelon), Mpio.
Yolox, 3000 m., 17.ix.1962 (E. C. Welling).
PARATYPES: Same locality as Holotype: 19 12.ix.1962, 19 17.ix.1962, 15 (8
genitalia slide no. M-2232, Lee I). Miller) 19 18.ix.1962 (all E. C. Welling).
Disposition of type material: Holotype and 6 Paratype (A), three 9
The specific name refers to the Chinantec tribe which inhabits the Sierra
Chinantla, Oaxaca, from whence the types came.
This very striking species seems to tie Paramacera to more "conventional"
Euptychiini through the basic lack of an upper surface pattern, especially the
usual Paramacera large blind forewing ocelli on the upper surface. On the
under side, however, P. chinanteca is unmistakably a member of Paramacera,
and its genitalia are remarkably similar to those of the other three species in
the genus. On the basis of the more generalized upper surface pattern and the more
diffuse androconial patch I consider P. chinanteca to be the most primitive
member of Paramacera. Both it and the somewhat primitive P. copiosa seem to be
restricted to small, relict populations in the high mountains of Oaxaca, not far
from populations (much larger ones, incidentally) of the widespread P. xicaque.
Figures 1-5, Paramacera chinanteca, new species. 1-2, Holotype 6 upper (1)
and under (2) surfaces; MEXICO: OAXACA: Muo Cu6u (A). 3-4, Paratype 9
upper (3) and under (4) surfaces; MEXICO: OAXACA: Muo Cu6u (ECW).
5, S genitalia of Paratype (slide M-2232) (A).
Paramacera copiosa, new species
Figures 6,7 (3), 8, 9 (androconial scales), 10 (5 genitalia)
Male: Head, thorax and abdomen clothed with olive-brown hairs above, dark
gray ones below. Antenna brown narrowly ringed with yellow above, more widely
ringed with yellow below; club brown above, buff below, dark brown at tip.
Palpus smoky-black, white laterally. Legs with long smoky-black hairs.
Forewing above olive-fuscous, somewhat darker in cell, with dark fuscous
margin, two coalesced dull brown blind ocelli in M1-M2 (much the larger) and
M.,-M., and an indistinct brown ocellus in M -Cu,; androconial patch dark
gray-black extending outside cell from anal cell to M.-Cu, with a relatively
straight distal margin.
Hindwing above olive-fuscous, hairs at base slightly darker, with dull
brown blind extradiscal ocelli from Sc+R,-Rs to Cur-2A (those in Rs-M, and
Cu -Cu, being the largest and most prominent, that in Cu22A being very faint
and that in M,-Cu, bearing a small buff pupil); marginal and submarginal lines
scale, approx 165 x. 9, interrib structure of same scale, approx. 10,
genitalia of otype (slide M-2338) (A).
'fSS ai "' I0
genitalia of Holotype (slide M-2338) (A).
dull brown anteriad of Cu|, shading reddish-brown to anal angle.
Forewing below as figured: reddish-tan, darker basad and gray along inner
margin, with two thick reddish-brown median lines, one across cell, the other
relatively straight from M, just outside cell to 2A near anal angle; three
white-pupilled black ocelli from MM-M, to M:-Cu, (the one in M -M, much the
larger and with thick black iris) with yellow and reddish-brown rings; two
marginal and a slightly dentate submarginal reddish-brown lines and a gray shade
between submarginal band and rings of ocelli from R.-R., to M|-M,.
Hindwing below as figured: brown peppered with gray scales at base,
tan central band (grayish-brown in cell) bounded on either side by reddish-brown
hands and not continuous to costa, area enclosing ocelli grayish-violet with
black, white-pupilled ocelli in Rs-M|, MI-M Cu,-Cu, and Cuj2A and silvery-white
ocelli in M.,-M: and M:.-Cu,, all with yellow and reddish-brown rings, and two
marginal and one submarginal reddish-brown lines meeting at anal angle.
Fringes above pale buff, dull brown between veins (more narrowly on
forewing); fringes below pales gray, brown at ends of forewing veins, dark gray
at ends of hindwing veins.
Length of forewing of Holotype $ 21.0 mm., that of single 3 Paratype 20.0 mm.
3 genitalia as figured, differing from those of otherspecies in Paramacera chiefly
in broader valva with different terminal ornamentation and having more
Androconial scales longer than those of other species, but with similar
general appearance. Androconial mass with denser concentration of androconial
scales than in P. chinanteca -- in this more closely approximating situation
in other two species.
Described from two specimens, both males, from the high mountains
of Oaxaca, Mexico.
HOLOTYPE 5: MEXICO: OAXACA: Muo Cu6u (Cerro Pelon), Mpio. Yolox,
300( m., 9.v.1962 (E. C. Welling): ( genitalia slide No. M-2338 (Lee I). Miller).
PARATYIE S: Same data as Hlolotype.
Disposition of type material: Holotype (A), ( Paratype (ECW).
P. copiosa is a more "conventional" Paramacera but can be distinguished
immediately by the silvery-white ocelli without black irises in M.,-M and M.-Cu,
on the under surface of the hindwing. The dull diffuse pattern of the upper surface
gives the species its name and is characteristic when compared with the xicaque
group. Whereas the upper surface pattern is reminiscent of the of P. xicaque,
the heavy, rather straight extradiscal band of the forewing beneath and the
general appearance of the surface on both wings suggests an affinity with
P. chinanteca. It is curious that both P. copiosa and chinanteca have been found only
at Muo Cuou to date, but while they share a common locality, the collection
dates suggest that the two species may be temporally isolated. The two species
appear to be the most primitive in the genus.
Paramacera xicaque (Reakirt), 1867 ["1866"]
This species is restricted to Mexico, having its metropolis in the Valle
de Mexico, extending northward to at least northern San Luis Potosi amd
southward to at least Oaxaca. Our experience would indicate that this butterfly is
restricted to localized populations in open woodlands, often many miles from
the next population. It is therefore not surprising that some subspeciation has
taken place, perhaps more than I am recognizing.
There is some question about the use of the name xicaque for this species,
as well be discussed under the nominate subspecies, but Reakirt's name is
maintained here in the interest of nomenclatorial stability.
Two subspecies are currently recognized within P. xicaque, but there is
evidence that at least one more may be separable when more material becomes
available for study. The females are more variable than are the males.
KEY TO THE SUBSPECIES OF Paramacera xicaque (REAKIRT)
1. Hindwing extradiscal spot in M,-M, elongate, tapering distad; all
ocelli on upper surface very much enlarged, especially in Y; Oaxaca,
M exico .................................... P xicaque rubrosuffusa, n. ssp.
1'. Hindwing extradiscal spot in M,-M, rounded or slightly oval, not
tapering distad; 9 with spots normally developed
.................. ............................. P. xicaque xicaque (Reakirt)
Paramacera xicaque xicaque (Reakirt), 1867 ["1866"]
Figures 11, 12 (6), 13, 14 (9), 15, 16, 17 (androconial scales),.18 (3 genitalia),
19 (9 genitalia)
Neonympha xicaque Reakirt, 1867 ["1866"j: 336-337. Type-locality: "nr.
Vera Cruz, Mexico". Type apparently lost: see notes below.
= Neonympha epinephele C. & R. Felder, 1867 [1864-1867]: 476. Type-locality:
"Mexico". Type in British Museum (Natural History), ex Rothschild Bequest.
A certain amount of confusion surrounds the proper name to be applied
to this species. The "1866" volume of The Proceedings of the Academy of
Natural Sciences of Philadelphia, as is so often the case, was published partly
during the year it represented and partly during the following one. The description
of Neonympha xicaque Reakirt (1867 ["1866"]: 336-337) was published entirely
in 1867 (A. N. S. P., 1913: xii), according to the only dates available, those
of receipt of the parts by the library of the U. S. National Museum, Washington, I). C.
Pages 280-336 of the Proceedings for "1866" were published sometime before 13
February 1867, and page 337 to the end of the volume were mailed sometime
before 10 July 1867. These dates splitting Reakirt's description of P. xicaque
are of significance since the Felder brothers described Neonympha epinephele in
the third part of the "Reise Novara" which was specifically dated March, 1867,
between the two parts of Reakirt's description! Articles 10-12 of the International
Code of Zoological Nomenclature imply that the name xicaque could only become
available after all the relevant criteria for availability were met: the major point
that seems in question is whether or not the part of the description on p. 336 is
sufficient to qualify the February, 1867, date as the date of publication of the
name. If the description is complete enough on p. 336, then xicaque is
legitimately the senior synonym; if the description did not provide a sufficient
"indication" until July, 1867 (p. 337), then technically xicaque should fall to the
Felders' name. The superficial character which best separates P. xicaque from
P. allyni is the presence of the reddish-brown marginal and submarginal lines
on the upper surface of the hindwing, and here too a problem develops. These
lines are mentioned in Reakirt's original description as follows: ". .[page
3361 the margin presents three continuous red-brown / [page 337] lines
obscured by a darker shade towards the apex . ." Therefore, the criterion of a
"definition" is only partly satisfied in that part of the description published in
or before February, 1867. The entire description of the under surface and the
designation of the type-locality were given on p. 337. No figure was provided.
The very controversial "Fifty-Year Rule" (Int. Code Zool. Nomen., 1961: Art. 23b)
could be applied conveniently to suppress epinephele, but there is a strong possibility
that this rule may be written out of the "Code" in the very near future.
There is some evidence that Reakirt may have distributed preprints of
his article well before the complete publication of it in July, 1867. F. M. Brown (1964,
1965) has dealt with this phenomenon of the early distribution of preprints and
accepts the date of preprint distribution as the date of publication. Ultimately the
International Commission on Zoological Nomenclature must rule definitively
on this subject. Recognition of a preprint date earlier that March, 1867, gives
xicaque clear priority over epinephele; rejection of preprint dates gives epinephele
priority under a strict interpretation of Articles 10 and 16 of the "Code"
(Int. Code Zool. Nomen., 1961). This latter course, of course, would tend to
upset the stability of nomenclature with regard to this species (such stability is
an avowed aim of the "Code"): epinephele, when recognized as a synonym of
xicaque has invariably been placed in the synonymy of xicaque. Suppression of
tne name xicaque in favor of the Felders' name would have a destabilizing
effect on the nomenclature, hence, the retention of Reakirt's name in this
Figures 11-17, Paramacera xicaque xicaque (Reakirt). 11, 12, 3 upper (11)
and under (12) surfaces; MEXICO: "Dept. Fed." (CM). 13-14, 9 upper (13) and
under (14) surfaces; MEXICO: HIIALGO: 5 mi. NE Zimapdn (A). 15,
androconial scale, approx. 230 x 16, detail at tip of same scale, approx. 2,000 x.
17, interrib structure of same scale, approx. 6650 x.
S* *- 2
: 5 y<1 A--
A redescription of nominate P. x. xicaque follows:
Male: Head, thorax and abdomen blackish-brown clothed with olive-brown
hairs above, gray-brown ones below. Palpus dark brown above, gray-brown
below. Antenna brown above, narrowly ringed with cream, cream below; club
brown above, cream below, blackish at tip. Legs clothed with grayish-brown
hairs, tan ones distally.
Forewing above olive-brown with costa and margin brown, a dull, slightly
darker androconial patch continuous outside cell from anal cell to M:-Cu,, thence
broken to vein M, and at least a dark brown ocellus in M,-M2 (usually one in
M:.-Cul, occasionally a small one in M,-M:,).
Hindwing above olive-brown, darker basally and often slightly red-tinged
in extradiscal areas, with at most six extradiscal brown spots from Sc+R,-Rs to M,
Cu, (those in Sc+R,-Rs and MI-M, often wanting) and with marginal fuscous
area resolving basad into three distinct red-brown lines from at least MM-M,,
to anal angle.
Forewing below reddish-tan, darker costad and grayish apically, with two
prominent red-brown lines, one across middle of cell, the other arising near
origin of Rs and proceeding posteriad toward anal angle, two straight marginal and
one crenulate submarginal brown lines, grading to red-brown near anal angle
(subnarginal line sometimes joining outer extradiscal band near anal angle),
and bearing a well-developed black ocellus with white pupil in M,-M, and smaller
ones in M~-M:, and M:,-Cu, surrounded by tan patch delimited by red-brown ring
(occasionally with separate ring around ocellus in M:,-Cul).
Hindwing below basally dark brown grizzled with gray, then two zigzag
red-brown bands from costa to inner margin across and outside cell enclosing
a buff central area broadly overscaled with gray, then a grayish-tan distal
area with a slight violet sheen enclosing six black ocelli with white pupils and
tan then red-brown rings from Sc+R,-Rs to Cul-Cu, (those in Rs-M, and M:.-Cu,
the largest), with a heavy irregular dentate red-brown submarginal line
connecting at anal angle with extradiscal band and two wavy red-brown
Fringes white above, checkered with brown at ends of veins, tan below
with brown checkering at ends of veins.
Figures 18-19, genitalia of Paramacera xicaque xicaque (Reakirt).
18, g genitalia (slide M-2102); MEXICO: HIDALGO: vic. El Encarnaci6n (A).
19, 9 genitalia (slide M-1981); MEXICO: "Dept. Fed." (CM).
Lengths of forewings of 6 specimens at hand range from 19 to 22 mm., and most
specimens have forewing lengths between 20 and 21 mm.
6 genitalia as figured and distinguished from those of the relatively close
P. allyni particularly by the relatively shorter uncus (less than half combined
length of uncus + tegemen) and the distally broader valva.
Androconia in a compact patch with dense concentration ofandroconial scales.
Scales not so long as those of P. copiosa and not easily separable from those
of P. chinanteca or P. allyni, although the interribs of P. allyni are broader
and those of P. chinanteca not so broad as in P. xicaque.
Female: Upper surface similar to that of 8, but with no androconial patch;
ground color somewhat paler and extradiscal spots larger than in 3. Females from
Hidalgo are extensively laved with reddish both without and within forewing
cell and extradiscally on hindwing; this color only faintly indicated in females
from other P. x. xicaque populations examined.
Under surface similar to that of $.
Lengths of forewings of specimens examined range from 17 to 23 mm., most
being between 20.5 and 22 mm.
9 genitalia as figured and differing from the very similar P. allyni genitalia
by the broader ductus bursae.
64 male and 35 female specimens have been examined representing this
subspecies from the following localities, all in Mexico:
SAN LUIS POTOSI: El Salto, viii, 18 (A).
HIDALGO: 5 m. NE Zimapin, 1980-2140 m., i, ii, 113 109 (A); vic. El
Encarnaci6n, 2400-2500 m., i, ii, 12C3 169 (A). 1P
MICHOACAN: Tancitaro, 660', vii, la (A).
DISTRITO FEDERAL: Desierto des Laones, mr. La Venta, 7800-8000', vi, 18
(AMNH)' Desierto, iii, 386 19 (AMNH); Popocatapetl Park, 9500-11,500', vi, 46
(USNM); "Dept. Fed.", v, x, xii, 143 39 (CM)
MORELOS: Barranca des Lobos, ix, 19 (A).
PUEBLA: Manzanilla, ix, 36 19 (AMNH).
VERACRUZ: Jalapa, 86 39 (AMNH, USNM); Misantla, 1(6 (USNM).
"MEXICO": ix, 26 (A).
no locality: 36 (USNM).
The type specimen of P. x. xicaque appears to be lost. I have checked the
collections in which it might be expected to be housed and have found no specimen
that might qualify as the type. The stated type-locality -- "near Vera Cruz" -
is a most unlikely one, unless one interprets "near Vera Cruz" very broadly; the
town of Veracruz (present spelling) is situated on the Gulf of Mexico at sea
level, and P. x. xicaque is a montane, or at least foothill, insect throughout its
range. Harry Clench (in litt.) informs me that he has been able to trace the
sources of some of the material that Reakirt used in his "1866" paper, and
at least some of these specimens were sent to William Henry Edwards (and
ultimately to Reakirt) by a collector who resided at Orizaba, Veracruz. In all
likelihood the type of xicaque came from this lot of specimens, and the
collector might well have labelled the specimens from the state in which he
lived, Veracruz. The type of this species, and several others named in the same
paper, probably came from the area of a triangle bounded by Jalapa and
Orizaba, Veracruz, and Puebla, Puebla, quite probably from the flanks of
Mt. Orizaba. There are no recent specimens from the Veracruz side of the
Veracruz-Puebla border with adequate data that have come to my attention: if
such specimens are extant, they are fine candidates for designation as the neotype
of this species. Until such specimens are located and a definitive type-locality
can be designated, it seems inadvisable to create a neotype of xicaque.
We have taken this butterfly in wooded montane habitats, both very dry as
northeast of Zimapan and mesic as near El Encarnaci6n, in Hidalgo. These
two localities are discussed in some detail by Miller and Miller (1970) and
The flight of these insects is erratic and near the ground, usually in or
near the underbrush. They are rather nervous and difficult to approach, but their
flight is weak enough to make capture on the wing easy. The butterflies will
"sun" themselves, especially in the early morning, by alighting on the ground
with the wings held upright and oriented so that sunlight gets maximum
intensity on one side or the other; the insects will walk around on the ground,
turning about until they achieve the optimum position with regard to the sun.
_ ~_ ~
The ruddy upper surface and very dark under surface of the Hidalgo specimens
suggest that these insects might represent another subspecies, but the material from
other localities in the Valle de Mexico is generally somewhat faded, and at least
some of the differences noted may be those-of fresh us. worn specimens. Material
from either side of the Hidalgo populations, from El Salto, San Luis Potosi, to the
northeast and from Distrito Federal to the southwest, are almost indistinguishable.
The presence in Hidalgo of a subspecies surrounded on all sides by the nominate
one, while not impossible, is somewhat less than likely.
The present species is distinguished from the more northern P. allyni,
described below, by the very prominent red-brown marginal and submarginal
lines on the hindwing above extending from at least M:-Cu, to the tornus in even
worn specimens; these lines are either totally fuscous or only faintly reddened
in the tornal area in P. allyni. There are also genitalic differences which may be seen
in comparison of the figures of the terminalia of both species (Fig. 18- xicaque;
Fig. 28 -- allyni). Additionally, the northern insect is superficially larger and
duller than is P. xicaque.
Specimens from the most southerly population thusfar examined, from
the mountains of eastern Oaxaca, are distinct and are described below.
Paramacera xicaque rubrosuffusa, new subspecies
Figures 20, 21 (a), 22, 23 (9)
Male: Upper surface differs from that of nominate xicaque in the less
olive ground color, the extradiscal area of the forewing and most of the hindwing
strongly reddened and by the hindwing extradiscal spot in M,-M, elongated
distad. Beneath the ground color of the hindwing is paler and the hindwing
extradiscal spots larger than in the nominate subspecies.
6 genitalia as in P. x. xicaque, but the uncus is straighter; androconia as
in the nominate subspecies.
Length of forewing of Holotype 6 21.0 mm., those of the 134 6 Paratypes ranging
from 18.0 to 22.5 mm., most having forewing lengths between 21.0 and 21.5 mm.
Female: Redder above than any nominate 9 xicaque with a better defined,
broader chocolate-brown forewing margin and much larger extradiscal spots
on both wings with one in hindwing space M,-M, elongated distad. Forewing beneath
ruddier than in nominate subspecies, hindwing paler and all extradiscal ocelli
larger in present subspecies.
9 genitalia indistinguishable from those of P. x. xicaque.
Lengths of forewings of the 52 9 Paratypes ranging from 21.0 to 24.5, most being
between 22 and 23 mm.
Described from 187 specimens, 135 males and 52 females, from the
mountains of Oazaca, Mexico.
HOLOTYPE 6: MEXICO: OAXACA: San Jose Pacifico, Mpio. Ri6 Hondo,
2400 m., 9.x.1967 (E. C. Welling); 6 genitalia slide No. M-2101 (Lee D. Miller).
PARATYPES: Same locality as Holotype: 1326 519, x-xi; OAXACA: Cerro
Machin, Mpio. Comaltepec, 2800 m., 16 19, xi; Agua Azul, Mpio. S. Andres
Paxtlin, 2300 m., 16 x (all E. C. Welling).
Disposition of type material: The entire type-series is now in collection (A),
but Paratypes will be distributed to various museums and to Mr. Welling at
a later date.
This subspecies is somewhat larger than is typical xicaque but is particularly
characterized by the reddish upper surface and the elongate hindwing discal
spot in M,-M.,. Thestraighter.uncus in the present species is also characteristic,
but certainly not at the specific level.
I am unable to determine to which subspecies the Oaxaca specimens mentioned
by Godman and Salvin (1879-1901: 101) belong, though from the description
given by those authors, they seem to be the nominate one. I also expected that
P. xicaque in some form would have been found in the montane areas of
Chiapas and possibly even Guatemala, but extensive collections from Chiapas by
Mr. Robert Wind and others have not located the species in that state: perhaps
Oaxaca is the southernmost limit for it.
Figures 20-23, Paramacera xicaque rubrosuffusa, new subspecies. 20-21,
Holotype 8 upper (20) and under (21) surfaces; MEXICO: OAXACA: San
Jose Pacifico (A). 22-23, Paratype 9 upper (22) and under (23) surfaces;
MEXICO: OAXACA: San Jose Pacifico (A).
Paramacera allyni, new species
Figures 24, 25 (3), 26, 27 (9), 28 (d genitalia)
Paramecera Isic.] xicaque of authors, not Reakirt (1867 ["1866"]: 336-337).
Male: Differs from that of xicaque above by the more uniform olive-
brown ground color, the duller fuscous extradiscal spots, the marginal lines
of the hindwing being uniformly fuscous or only slightly tinged with reddish
at anal angle (never bright red-brown from M2-M:i to tornus as in Mexican
xicaque) and in the androconial patch being better developed in forewing
spaces MI-M2 and M.-M,~ outside cell. Below differing from xicaque in the
paler ground color, especially outside hindwing cell where the area enclosing
the ocelli is silvery-gray.
& genitalia similar to those of xicaque, but uncus longer (greater than half
length of uncus + tegumen) and valvae more gently tapered distad to narrower
Androconia similar to those of P. xicaque, but the rib spacing is greater
Figures 24-28, Paramacera allyni, new species. 24-25, Holotype 3 upper
(24) and under (25) surfaces; ARIZONA: Cochise Co.: Barfoot Park, Chiricahua Mtns.
(A). 26-27, Paratype Y upper (26) and under (27) surfaces; ARIZONA: Cochise
Co.: Barloot Park, Chiricahua Mtns. (A). 28, 3 genitalia of Holotype (M-2199) (A).
in the present species.
Length of forewing of Holotype a 22.0 mm., those of 93 5 Paratypes range from
21.0 to 23.0 mm., averaging 22.1 mm.
Female: Differs from 9 of P. xicaque in same general characteristics as does
8, but without androconial spot, of course. Reddish tinge above, where present,
only faintly indicated and mostly on forewing.
9 genitalia similar to those of P. xicaque, but distal part of ductus bursae
not so bulbous as in that species.
Lengths of forewings of 31 9 Paratypes range from 21.5 to 24.5 mm., averaging
Described from 124 specimens, 93 males and 31 females, from the
Chiricahua Mountains, Arizona.
HOLOTYPE 5: U. S. A. : ARIZONA: Cochise Co.: Barfoot Park, 8000',
Chiricahua Mtns., 12.vii.1970 (K. Roever); a genitalia slide No. M-2199
(Lee D. Miller).
PARATYPES: Same locality as Holotype: 15( 49, vii; same locality as
Holotype, but 8600', 28 19, vii; same locality (as "Barefoot" Park), no
elevation, 29, vii; upper camp, Pinery Canyon, Chiricahua Mtsn., 15, vi; Pinery
Canyon, 35, vi; Rustler Park, Chiricahua Mtns., 55 189, vii, viii; above
Rustler Park, 8200', 4c, vi, vii; "Chiricahua Mtns.", 63( 69, vi, vii.
Disposition of type material: Holotype 8, 158 and three 9 Paratypes (A);
nine c and 219 Paratypes (AMNH); 253 Paratypes (LACM); 153 and three 9 Paratypes
(CM); 213 and three 9 Paratypes (USNM); five 3 and one 9 Paratypes (KR).
I take great pleasure in naming this species for Mr. A. C. Allyn in
recognition of his contributions to lepidopterology.
In restricting the type-locality of P. allyni to the Chiricahua Mountains, I
have excluded additional material from the type series which has, nevertheless,
been examined. These specimens, 89 in all, 75 males and 14 females, refer
to this species and bear locality labels as follows:
U. S. A.: ARIZONA: Apache (?) Co.: "White Mtns.", 8500', 25, vi (AMNH).
Cochise Co.: Miller Canyon, 6 mi. SW Rt. 92, Huachuca Mtns., 7500', 1(,
vii (KR); "Huachuca Mtns.", 353 69, vi-ix (AMNH, CM, LACM, USNM); "H.
Ariz." IHuachucas?|, 13, vi (AMNH); Ft. Huachuca, 13 19 (LACM); Paradis,
133 29, vi, vii (AMNH, USNM). Santa Cruz Co.: Nogales, 23 vi (CM, USNM).
"So. Arizona", 23 vi (AMNH, LACM). "Ariz.", 25 19, vi (AMNH, USNM).
MEXICO: CHIHUAHUA: "Chihuahua" (Harry K. Clench has determined
a more precise locality for these specimens from the notes that the collector
Townsend made, which notes are now in the archives of Carnegie Museum:
Upper Rio Piedras Verdes [300 15' N, 1080 15' W], 7100-7300'), 33 19, viii,
This species, long confused with P. x. xicaque, can be separated from that
species by its larger size, by the lack (or virtual lack) of red-brown tornal shading on
the hindwing above, by the generally duller upper surface coloration, the
silvery extradiscal part of the hindwing below and by the genitalic characteristics
cited in the description.
The Chihuahua specimens are more "washed out" than are those from
Arizona and could represent a different subspecies, but the material at hand
is not sufficient to confirm such a judgment with certainty.
P. allyni is found in the high pine woodlands in the mountains of
southeastern Arizona, and its habits are comparable with those of P. xicaque.
I have no idea what the habitat is like in Chihuahua where Townsend took the
short series in Carnegie Museum, but I assume it is similar to that in the
Arizona mountains based on the other species taken there, including the
type-series of Speyeria nokomis coerulescens (Holland). The White Mountains,
Apache (?) Co., Arizona seem a bit strange, since P. allyni has not been
recorded from intervening and equally logical areas, such as Mt. Graham or
the Santa Catalina Mountains, but suitable-looking habitat is not uncommon
in at least the southern Whites, so the record may be correct. If it is, I can see no
reason why P. allyni shouldn't be found in the mountains of southern New
Mexico and perhaps on Guadalupe Peak, Texas.
PHYLOGENETIC AND ZOOGEOGRAPHIC CONSIDERATIONS
It is rather difficult to postulate which of the euptychiine stocks gave rise
to Paramacera, but this really will be the province of the final paper in this
series. Some inferences, however, may be drawn within the genus.
Despite the close similarity in the genitalia of all four species, P. chinanteca has
a highly aberrant pattern for a Paramacera, but a more "conventional" one within
the tribe. Unfortunately, the pattern represented by P. chinanteca is not
duplicated elsewhere in Central America by any euptychiine group, but it is
represented in South America by members of the "Euptychia" paeon complex.
These latter insects are genitalically very remote from Paramacera. This
factor suggests that the basic stock from which Paramacera rose is a rather
ancient one in the tribe and that the genus may have been isolated from its
nearest relatives over a considerable period of time. Paramacera is not the only
member of this ancient Mesoamerican fauna: the primitive danaid genus
Anelia Hiibner (= Clothilda Blanchard) and the satyrid genera Cyllopsis and
Pindis have their metropolis in Mexico and/or Central America, for example.
One can only assume that the progenitor of Paramacera arrived or arose in
Mexico (possibly Central America) during the Tertiary when this area was
isolated from South America, or possibly the "proto-Paramacera"
represents one of the original invading stocks of Euptychiini from the Old
World and never made it into South America. In either event, Paramacera is
an old euptychiine genus, far older than most of the South American genera and
With the exception of P. copiosa, which seems to be of an earlier vintage,
the remaining Paramacera are closely related and seem to represent a much
more recent radiation of the genus. The bifurcation of P. xicaque and P. allyni
is likely to be of post-Pleistocene age and coincident with the drying of the
entire area from Arizona through the Valle de Mexico which effectively
separated the ancestral stocks into a northern (allyni) and southern (xicaque)
Three of the four species of Paramacera are found in the mountains of
Oaxaca, and this area appears to be a likely site of the basic evolution within the
group. The fourth species, and certainly the youngest in the genus, is found far
to the north of the other three and is an offshoot of one of the Mexican species.
Two species, P. chinanteca and copiosa, are known only from very small relict
populations high in the mountains of Oaxaca, and both species appear to be
"on the way out", a common fate for primitive, less well-adapted species.
The other species were derived from a single widespread species during
glacial maximum, at which time the pine woods environment was much more wide-
spread and continuous than is the case today. The contraction of these
woodlands during recent times probably caused the present-day fragmentation
of the populations and their subsequent evolution.
The distribution of the species of Paramacera is shown on the map in
Many people have aided in the analysis of this genus and in the final
preparation of the manuscript. Mr. William D. Field (USNM), Mr. Julian P. Donahue
(LACM) and Dr. Frederick H. Rindge (AMNH) lent material for the study and
were patient with me when the entire manuscript was rewritten after
discovery of the two species from Oaxaca, P. chinanteca and P. copiosa; I
am most grateful for their generosity and forebearance. Mr. Harry K. Clench
(CM) made the collections of that institution available to me and gave helpful
advice and information relative to several vexing points in the study. Dr. F.
Martin Brown, Colorado Springs, Colorado, was a great help in resolving some of
the problems of authorship and in the dating of relevant publications. Mr.
Kilian Roever, Phoenix, Arizona, collected the basic type-series of P. allyni at
my request and was generally most cooperative. Sr. Eduardo C. Welling M. of
M6rida, Yucatan, Mexico, collected the type-series of P. xicaque rubrosuffusa
at my request, then generously made available two species hitherto unsuspected
by me for description in this paper: to him, of course, I owe a great debt of
thanks. The photographs which illustrate this paper and the electron micrography
of the androconial scales were done by Mr. A. C. Allyn of this institution. Mr.
Allyn and my wife and colleague, Jacqueline, read and commented upon the
Figure 29, distribution of Paramacera. Closed circles, P. x. xicaque; open
circles, P. xicaque ruhrosuffusa: closed triangles, P. allyni; star, only spot
from whence P. chinanteca and P. copiosa are known.
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