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V. Nabokov.
Notes of Neotropical Plebejinæ (Lycænidæ, Lepidoptera).
Psyche 52:1-61, 1945.

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PSYCHE
VOL. 52 MAR.-JUNE, 1945 Nos. 1-2
NOTES ON NEOTROPICAL PLEBEJINB
(LYCkENIDB, LEPIDOPTERA)
BY V. NABOKOV
Museum of Comparative Zoology
In a recent paper2 I briefly listed the' only Plebejinoe (s.s.) found in the Nearctic region. Subsequently I decided to see whether any true Plebejince occurred in the neotropics besides the three or four species the genitalia of which I had happened to examine before. The results proved so unexpected and in- teresting that it seems worth while to publish the present paper despite its rather superficial and incomplete nature. In order to cover more ground (and, in some cases, owing to the scantiness of the material at hand) only a very small num- ber of specimens (about 120 in all) have been dissected and drawn (after a few Catochrysophios and representatives of other subfamilies had been weeded out by the same method). Some of these figures are appended. All the specimens, except a few supplied with his usual kindness by Mr. W. P. Comstock of the American Museum of Natural History, are preserved in the Museum of Comparative Zoology, Harvard. A rather drastic rearrangement of the species and groups was an inevitable consequence of this investigation. Seven new genera have been introduced; two have been revised and re- stricted. In several cases it was found that forms had been assigned by recent authors to the wrong species. Some syno- Published with the aid of a grant from the Museum of Comparative Zoology at Harvard College.
1944 [Feb. 19451 Psyche 51; 104-138, where the following errata should be corrected: line 12, p.
105, instead of "hanno Stoll" read "ceraunus Fabricius (nom. spec.)"; line 28, p. 107, instead of the misprint "calliopsis" read "calliopis"; p. Ill, in the sentence beginning "A complete sequence . . ." transpose "palearctic" and "nearctic."




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2 Psyche [March- June
nyrns have been tracked down, others are tentatively suggested but cannot be finally disposed of until the types are examined or neotypes fixed). The brief bibliographical references given are merely intended to indicate the identity of the forms dis- cussed. Beyond the inclusion of some random notes on certain phases of pattern, macroscopical characters are not discussed, and no attempt has been made to revise in this respect the (fortunately rather few) races that have received names. In spite of the work accomplished since 1909, by Tutt and Chapman in England and by Stempffer in France, entomologists in this country employ the term "Plebefind' simply as a eu- phemism for the "Lycosna" of German authors, or blue^,"^ and 'Plebejus7' is used for a number of heterogeneous Nearctic species only one of which (saspioZus Boisduval) belongs struc- turally to the genus of which the Palearctic Plebejzts argus Linnse-us is the type. In a way the initial blunder was Swinhoe's who while correctly giving a subfamilial ending to the group which Tutt7s intuition and Chapman's science had recognized ("tribe" Plebeidi which exactly corresponds to the Plebejk of Stempffer ) as different from other "tribes" (Lea, subfamilies) within the Lyccenidos, failed to live up to the generic diagnoses which he simply copied from Chapman's notes in Tutt and tried to combine genitalic data he had not verified or did not under- stand with the obsolete "naked v. hairy eyes" system (which at Butler's hands had resulted in probably the most ludicrous assembly of species ever concocted, see for example Butler 1900, Entom. 33: 124), so that in the case of several Indian forms which Chapman had not diagnosed, Swinhoe placed intra- generically allied species in different subfamilies and species belonging to different Tuttian "tribes" in the same subfamily. In reality the subfamily Plebejince is extremely well differen- Thus McDunnough uses "Pkbekm" in his "Check List" of Nearctic Lepi- doptera (1938 Mem. S. California Acad. Sci. 15'61, and thus Comstock uses "PIfbejhiK" in his work on Rhopalocera of Porto Rico and the Virgin Islands (1944, in Miner, Scient. Survey P. R. and V. Isls. 12:492), but the two references the latter author appends (Swinhoe 19x0, Lep. Indica 8:10 and Hampson 1918, Novit. 2001. Tring 25:38S) are most misleading: the first, because Syrttarttciis Butler, a Fenus structurally indistinguishable from Leptotes Scudder (which is one of the two genera assigned by Comstock to "Pk!ie.$na: Swinhoc") is placed by Swinhoe in a different subfamily, namely Lawpidim (now known as Cato- chrysofnce), and the second, because Hampson's (perfectly invalid) use of $'Plebejtfs" and "Plebefinai' refers to a section of a different family, namely Erycinidos (now known as Riodimda) .




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19451 Notes on Neotropical Plebejince 3
tiated in all its genitalic elements (the sedeagus and its append- ages, the tegumen, cingula, fakes, uncus lobes and valves of the male, and the cervix bursse and vaginal armature of the female) from the Catochrysopince (containing the holotropical Leptotes Scudder and a huge array of palseotropical species in several genera), the Glaucopsychince (containing, among others, the three holarctic genera Glaucopsyche Scudder, Scolitantides Hiibner [to which Phcedrotes Scudder and "Shijimia Matsu- mura" fall as synonyms] and Philotes Scudder), the Everim with the holarctic Everes, the Lyccenopsince with the holarctic Celastrina Tutt ( = Cyaniris Scudder, nec Dalman) , etc. The arrangement proposed in the present paper needs to be prefaced by a few words on taxonomic units. The strictly bio- logical meaning forcibly attached by some modern zoologists to the specific concept has crippled the latter by removing the morphological moment to a secondary or still more negligible position, while employing terms, e.g., "potential interbreeding," that might make sense only if an initial morphological approach were presupposed. What I term species, in my department, can be defined as a phase of evolutional structure, male and female, traversed more or less simultaneously by a number of, conse- quently, more or less similar organisms morphologically shading into each other in various individual or racial ways, interbreed- ing in a given area and separated there from sympatric repre- sentatives of any other such phase by a structural hiatus with absence of interbreeding between the two sets. In other words: 1. any two structurally indistinguishable individuals belong to the same species regardless of biological, physiological, geo- graphical or any other factors; 2. structurally distinguishable sympatric non-interbreeding sets represent different species re- gardless of all other considerations; 3. structurally distinguish- able sympatric individuals belong to the same species when they occur within an interbreeding set; 4. structurally distinguish- able allopatric sets belong to the same species if the hiatus between their structures is completely bridged by intermediate structures in other, not necessarily intermediate, areas; 5. ob- viously allied but structurally distinguishable allopatric sets not linked by such intergrades can be said to belong to different or the same species only by analogy, i.e., by analysing the struc- tural gaps between sympatric species, or individuals possessing the same general type of structure. Conditions 2 and 4 do not



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4 Psyche [March- June
exclude each other and so it may happen that two structurally distinguishable local forms belong to one species allopatrically because they racially intergrade, but at the same time belong to different species sympatrically because in some other region their structural counterparts occur side by side without inter- breeding (this incidentally is the position in Lyceides) . In such cases one should give precedence to the all important sympatric moment and find somewhere in the spirals of racial intergrada- tion a point at which the whole system can be elegantly, in the mathematical sense (for we are dealing with measurable structures), divided into two parts, i.e., two species, using some combination of trinomials to designate this or that interspecific form (e.g., Lycceides scudderi doei Roe trans ad melissa roei Doe). This state of affairs is not a flaw in the concept of "spe- cies" but an indirect result of its dual nature ("structure" plus "reproduction," "male" plus "female" etc.) and should be ac- cepted by the taxonomist with perfect equanimity.l The impact on the eye of a combination of characters in the whole structure or in an element of it, results in the perception of certain structural types. Structures of the same type imply phylogenetic affinities unless it can be proved, as in some cases it is easy to do, that the resemblance is "false" ie., attained by essentially different means. Such false resemblances are ex- tremely rare and the number of characters involved is small, and this is as it should be, since such "convergence" depends upon the mathematics of chance. False dissimilarities also occur (and are also rare), i.e., the striking difference between one type and another is seen, when analysed, to be due to a simple and brief process of evolution in an unusual direction. Unless we believe that certain structural resemblances and dissimilarities are not due to chance or to gross adaptional modifications, but can be classified according to their phylo- genetic sense, all horizontal genera are artificial groupings - of some practical use to collectors (e.g., the convenient lumping of all small blue butterflies with rounded hindwings and dotted undersides in one "genus") but of no scientific value. This brings us to the question as to whether a classification on the 'Subspecies" (on which I hold rather special views which I shall discuss else- where) may be briefly defined as a locally constant phase of specific alar char- acters with or without a local fixation of some stage within the graded variational range of the specific genitalic structure. The days are quite gone when easy-going describers could give names to these things without a detailed study of genitalic and pattern characters throughout the polytypic species or genus involved.



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19451 Notes on Neotropical Plebejince 5
basis of genitalia reflects natural relationships better than do other principles. I think the answer is "yes." A "polytypic genus" is determined by structural characters which are common to all the species it includes and the par- ticular combination of which, more than the presence of some particular detail, no matter how striking, distinguishes the group from any other. A "monotypic genus" (i.e., a structurally iso- lated species which does not fit into any known generic group) obviously lacks the first feature while the number of characters entering the distinctive combination is vastly increased by prac- tically coinciding with the whole array of specific characters, so that the only "reality" a monotypic genus has, lies in the im- plication that the only species it contains is the only one 'known" and that if others were "known," a common denomi- nator now "hidden" in the monotypic genus would be revealed. Among polytypic genera, a "natural genus7' is one which reflects the flickering, as it were, of a strongly differentiated type of combinational structure within limits as narrow per se as, say, the range of continuous variation within a structurally highly polytypic species, and thus consists of specific structures re- sembling each other more than they do any other species. If hi, ha, h3, h4 denote the interspecific hiatuses, and Hi, Ha, etc. the intergeneric ones, then the lesser the h's and the larger the H's, the more '(natural" the genus is - and the more liable it is to be transformed into a polytypic species by the next reviser with more material at his disposal.
A certain harmony, as yet rather obscure, seems to exist be- tween a particular type of male armature and a particular fe- male one; this has been taken into account in founding the genera discussed below. The impression I have formed so far that with "natural genera" specific differentiation in these or- gans is more marked (or at least easier to observe) in the male may be due to insufficient investigation, but anyway I cannot find any exact correlation between female lock and male key. In what manner and to what extent the sclerotized parts of the sexes in Plebejince fit each other during copulation is not clear, but I doubt whether the valves, the termination of which is evolutionally the most vulnerable part, come into any direct contact with such structures in the female organ that might lead to some intersexual adaptation.'
'Lorkoviz states (1938, Mitt. Miinchner Ent. Ges. 28:231) in an admirable paper on the European representatives of Everes (Ever&) that in that genus



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6 Psyche [March- June
Adaptation to surroundings, to climate, altitude etc., and hence "natural selection" in its simplest sense, certainly had no direct action whatever on the moulding of the genital armature, and we know nothing of the physiological processes of which that elaborate sculpture is the structural overflow. While ac- cepting evolution as a modal formula, I am not satisfied with any of the hypotheses advanced in regard to the way it works; on the other hand, I am quite certain that repetitions of struc- ture, on the Siberian tundra and on the paramos of the Andes, on a mountain in India and on an island in the Caribbean Sea, cannot be treated as a result of haphazard "convergence" since the number of coincident characters in one element, let alone the coincidence of that coincident number with a set of char- acters in another element, exceeds anything that might be pro- duced by "chance." Hence the conviction that there is some phylogenetic link where there is a recurrence of similar genitalic characters and that certain groupings - the new genera to which we now must turn - may be so devised as to reflect the natural affiliations of the species.
Plebejinae
Stempffer, 193 7-1938, Bull. Soc. ent. France 42: 2 11-2 18, 296-300; Nabokov, 1944, Psyche 51: 104-105; = Plebeiidi, sensu Tutt [et Chapman], 1909, British Butt. 3: 150-159; Chapman, 1910, Ent. Rec. 22: 101-103; 1916 Trans. Ent. Soc. London 1916: 157-180; = "Plebeius + Polyommatus" s. Be- thune Baker, 19 14, Ent. Rec. 26: 164 ; Polyommatince, Forster 1938, Mitt. Munchner ent. Ges. 38: 11 1-1 16. Parachilades n.g .
(fig. 1, figs. TIT, pi. 2,7)
Type and only known species Lyccena titicaca Weymer 1890 (in Reiss et Stubel, Reisen in Sud-America, Lepidoptera : 122- 12 3 "Titicaca Lake ; Sajama, Bolivia," pi. 4, fig. 6 [very poor] ; Itylos [s.l.] titicaca? Draudt, 1921, in Seitz, Macrolep. World, 5 : 122, pi. 144, m [coarse copy of original fig.] ; Cupido speciosa the median uncal projection (a structure not found in Plebejince and wrongly, in my opinion, regarded as being formed by the fusion of the uncus lobes) fits exactly the vaginal plate of the female, both varying together according to the species. See also Chapman 1916, Trans. Ent. Soc. London 1916:170.



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1945 1 Notes on Neotropical Plebejince. 7 Staudinger, 1894, Iris 7 :
77-78? "Huallatani and Quebrada
Malaga, Bolivia"; Lymna speciosa, ibid., pi. 2, fig. 8 8 ; Itylos [s.l.] speciosa, Draudt, 192 1, I.c., pi. 144, n, figs. 8 9 ) . Five males and one female investigated: prep. 610, "Titicaca FIG. 1. Parachilades titicaca, left hindwing underside x7 [Lake] , Bolivia," ex coll. Huntington [ex coll. Staudinger-Bang Haas], Amer. Mus. Nat. Hist.; prep. 483, 488, 589, 620, 2 590, "Sicasica, Bolivia, I .X. 1899" ex coll. Weeks, Mus. Comp. Zool. adeagus thickishJ1 about 1 mm. long, the suprazonal portion subequal to the subzonal one. In general type fairly close to In all genera examined the subzonal portion of the asdeagus appears in cross- section as a dorso-ventrad directed oval, the lengthening of which produces the appearance of "thickness" in the organ when the latter is viewed from the side.



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8 Psyche [March-June
Chilades (see pi. 2, CON I), still more curved, however, with a pronounced bulging of the outline (in lateral view) dorsally at the zone (above the zone and less conspicuous in Chilades) and a somewhat different structure of the suprazonal portion. Su- prazonal sheath terminating on the ventral side in a point (which is not notched as it is in Chilades) with two filament-like lateral portions (structurally similar to the spine-like single medial process described by Chapman in other genera and represented in Chilades I) diverging from it and rimming the vesica, the erected (everted) frothy membrane of which they seem to prop. Vesical opening (on the dorsal side) beginning just above the zone (thus at a more proximal point than in Chilades). Vesica very simple and weak as in Chilades, Frey&,'1 Lymides, etc. Alulse considerably more developed than in Chilades, forming two petals almost 0.3 long and resembling (or representing) rudiments of the peculiar element (sagurn) that exists at various degrees of development in several other neotropical genera where, however, it is well differentiated from the aIulse (except in Hemtargus). Furca considerably smaller in relation to the ~edeagus than in Ckilades, singularly thick, pincers-like, con- nected at its tips with the petals of the alulse. The whole dorsum (falx + uncus lobe + tegumen) remarkably similar in type to Chilades, which type is characterised by the breadth of the '
robust and long forearm exceeding that of the long finger-shaped uncns lobe," by the humerulus appearing to be produced (owing to the exiguity of the lobe) not from the base of the lobe but from the teamen proper, and by the latter being smaller by comparison to the fdx and the lobe than in other Piebejince. Differing from Chilades in the greater size of the falx and uncus lobe in relation to the rest of the armature and to the size of the One wonders whether this medial process in ChHades is not, perhaps, merely a lesser stage of development of the pointed part of the sheath of Parachiladn, while the lateral processes in the latter represent a lesser stage of development in comparison to the latero-ventral pointed sheath portions of Chffades. I am not fully satisfied with my observations in regard to the iedeagus of these two genera. 'I fail to find in either of the two species of Freyeria (trochilw and putii') the cornuti mentioned in the case of i-rochik by Stempffer (1937, Bull. Soc. ent. France 42:215).
In fact Fruhstorfer, the only German writing author of his time who made any attempt to follow the British authors in the study of Lycaenid genitalia for systematic purposes, in an enthusiastic, but amateurish, and poorly illustrated paper on Chiladm (1916 2001. Meded. Leiden, 290-95) mistook the uncus lobes of lams and cleotas for an additional pair of fakes (besides confusing generic characters with specific ones).




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19451 Notes on Neotropical Plebejinae. 9 win& Falx very big, long and thick, fatter than in Chilades, and not distinctly separated to the eye into its components (humerulus, elbow, etc.) owing (1) to its not bunching at the shoulder as it does in Chilades (in ventral view) ; (2) to the
unusual (unique in Plebe jince, typical in Catochrysopinae.) slant in the part that corresponds to the, very upright, forearm of Chilades, with a consequently wide and weak falcal arch; and (3) to its even breadth from basal point to almost three-quarters of its length; thus of a limacine appearance increased by the fact (again unique in this subfamily, but frequent in Catochry- sopinae.) that in ventral view the point of the oblique falx seems twisted away from the lobe instead of curving hookwise toward the latter as it does in Chilades (or other genera) where it at- tains the tip of the lobe. Uncus lobe narrow and long, exceeding the length of the tegumen (from base of falx to beginning of cingula) which is not the case in Chilades nor indeed in any other genus of the subfamily; tapering above the humerulus to form a finger-shaped projection of even breadth throughout; slightly excurved (in contrast to the straight "gothic" projection in Chilades} and at least 1% narrower than the forearm. Valve exceedingly small and squat, about half the aedeagus and about equal to the falx in length, the first proportion only approached in one other species of Plebejinae. (Hemiargus ramon Dognin) and the second unique in the subfamily (but common in other Lycaenids) ; of a peculiar stunted appearance, shaped like an elephant, about one and two-fifths as long as broad, thus strik- ingly different from the elongated shape of Chilades and all Old World members of the subfamily; with a strongly and evenly curved processus superior ending in a thickish gradually tapering rostellum (about a third of the valve in length), which continues the even curve of the whole upper margin and comes to rest upon the well-developed, strongly jutting mentum, the tip of which may assume a fluted appearance in situ. Female: fibula of ostium bursae strongly developed, of the Chilades type, with the upper lamella conspicuously long (about 0.3 mm.). Papillae anales about 0.45 mm. broad and very large in relation to the short looking rods (about 0.6). Measurements (in mm. ) : aedeagus 0.9-1, suprazonal portion 'The titicaca lobe length is only attained in Chilades by one species (cleotas, pi. 2, CLE 3) in which the whole alar surface is 5.4 times greater and the fore- wing 2.5 times longer than in titicaca, while in galba forms (e.g. pi. 2, CON 3)



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10 Psyche
[March- June
0.44-0.52 (mean 0.491, subzonal 0.44-0.54 (mean 0.49) ; breadth (in lateral view) at zone 0.16, proximad 0,12 ; penis mean 0.85. Furca 0.37. Falx 0.5 by 0.07 to 0.55 by 0.1 (mean 0.52 by 0.08) ; uncus lobe 0.5 by 0.045 to 0.55 by 0.055 (mean 0.52 by 0.05). Valve 0.54 by 0.39 to 0.55 by 0.4 (mean 0.54 by 0.4).
It is possible that individuals or broods or racially constant forms of titicaca with a complete underside forewing set of (seven) II macules and (seven) split I macules exist somewhere in the Andes. The general tendency, however, is to complete obsolescence (Staudinger selected for his figure of "Lycma speciosa" an individual with still visible I RM and Mill and MJI; Weymer's only fresh specimen had none). The narrow and pointed (almost tineoid) wing-shape is found elsewhere among Plebejiwe of great altitudes (e.g. in a Himalayan form of Albdina orbitulus Prun.). In the hindwing (see fig. I), the termen strongly recedes (below vein Ma) from scale line 85 to SO (at vein 2A). The I macules are obsolescent, except the CUi pr~etennind mark which is distinctly pigmented in some speci- mens. The I1 made is weakly pigmented (except marginally, especially along the outer edge in most specimens) from Sc to cell Ma (between, roughly, scale lines 3040, 35-50, 50-60, 50-60)) fairly strongly (with very strong edges) from Mg to cell Cu 2 (35-60, 30-40, 30-35,) and is very weak in 1A and 2A. The III made is weakly pigmented (except the proximal edge) in Sc (0-20) and fairly strongly (with still stronger edges) in Cua (10-20). The I discoidal RM (30-35) is very weak while the I1 one (R 12-25 +M 5-2 5) is fairly strong (with still stronger edges). All the macules except the anal ones and the Cu, prieterminal mark (60-65) fill the transverse breadth of the interspace (forming, if viewed from the termen a capital omega in the case of the I1 series, and a somewhat similar design in the case of Sc III, II RM, CU; 111) and are squarish, or of ;roughly triangular shape if extending to 15 or more scale lines (along the upper vein of the cell as Sc III and Cui 11 do, or along the lower one as II M and Rg I1 do). I give these scanty notes and a figure, since no intelligible description of the species exists? where dwarfs from Cyprus approach my largest titicaca (length of fore wing 8.5 mrn.) in wing span (though of course the wings remain always much fuller than in Paruchihdes), the lobe is at least twice smaller than in the latter. ' For a full discussion of the terminology employed see my paper (1944 op. rit.) on the pattern of Lycmnidee as expressed in Lywides.



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19451 Notes on Neot~opical Plebejin~ 11
Pseudothecla n.g.
(figs. FAG) PI. 2)
Type and only known species: Thecla faga Dognin 1895, Ann. SOC. ent. Belgique 39: 105-106 "Loja, Ecuador" ( = ? excisicosta Dyar 19 13, Proc. United States Natnl. Mus. 45: 63 7-63 8 "Cotahuasi ; Chuquibamba, Peru") .l One male investigated: prep. 6 1 1, "Peru," ex coll. Hunting- ton, Am. Mus. Nat. Hist. (with a somewhat more weakly marked underside than Dognin's description suggests). Bdeagus two-thirds of a millimeter in length) very slightly incurved distally, rather thickset, not unlike certain Plebejus species in type, the suprazonal portion hardly more than half the subzonal one in length, the vesical opening at 0)8 mm. from the zone, the vesica plain, rather weakly defined, thickly shielded ventrally by the suprazonal sheath; alul~ and tabs small. Furca resembling Pa~achilades, the branches still thicker) conspicuously curved, equal in length to the subzonal portion of the zdeagus. Traces of a thin membrane (? rudi- ments of sagum) between the latter and the furca. Falx bear- ing a general resemblance to certain Plebejus and Vacciniina species, its outline, however, more evenly rounded throughout. Forearm slim, incurved, tapering to a sharp point, subequal to the suprazonal portion of the zdeagus) humerulus thickish with a weak shoulder. Uncus lobe small, shorter than the forearm, rather narrow and blunt. Valve of the normal (fishlike) sub- familial shape) but exceedingly small, subequal in length to the zdeagus, about twice as long as broad, nicely tapering basad. Processus superior strongly scooped out at the rostellum which thus seems to be produced from a point lower than the upper margin of the valve and is curiously shaped: anteriorly forming a sharp point, posteriorly producing a kind of small heel at about half of the length of its inner margin. Measurements (in mm,) : zedeagus 0.67, suprazonal portion 0.24, subzonal 0.43 with breadth (lateral view) 0.1 1; penis 0.64. Furca 0.44. Ver tical/Horizontal extension of uncus : fore- arm 0.27/0.033, humerulus 0.07/0.14, shoulder 0.11/0.05) lobe


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