Ring T. Cardé, Arthur M. Shapiro, and Harry K. Clench.
Sibling Species in the Eurydice Group of Lethe (Lepidoptera: Satyridae).
Psyche 77(1):70-103, 1970.

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SIBLING SPECIES IN THE EURYDICE GROUP OF LETHE ( LEPIDOPTERA : SATYRIDAE)
We have uncovered a pair of sibling species in the comparatively well-known butterfly fauna of eastern North America: the common Grass Nymph or Eyed Brown, Lethe eurydice of recent authors, is actually two species, which are extensively sympa,tric. Curiously, the distinctness of these two has been known since at least 1936, when W. D. Field discovered and characterized them as subspecies. He assigned names to them which we now know to be inapplicable. This was corrected in 1947 by R. L. Chermock, who named the presumably more southern "subspeciesn appalachia. Neither of these authors was aware that the "subspecies" are sym- patric.
The present investigation was first suggested when one of us (Clench) found both forms flying in the same area near Leesburg, Mercer Co., Pennsylvania in 1966. The conspicuous habitat differ- ence between them implied that two species might be involved. In 1968 another of us (Shapiro) found the same situation in western and central New York and (with Card;) investigated the immature stages and biology of the insects. The results of this study are partly reported elsewhere ( Shapiro and Card;, I 970). Independently of us, C. F. dos Passos and his correspondents simultaneously made the same discovery. Several of the conclusions contained in the resulting paper (dos Passos, 1969) appear erroneous. Since the taxonomic situation is very complex, we here review the whole subject, nomenclatorially, morphologically, and distributionally. In brief, we recognize two species in this group, as follows: ( la) Lethe eurydice eurydice (Johansson) , widely distributed from Labrador to Great Slave Lake and south to Delaware and Illinois, occurring in open marshes and sedge meadows. ( 1b) Lethe eurydice fumosa (Leussler) , scattered in small isolated colonies (many now extinct) in sedgy permanent marshes in the 'Department of Entomology and Lirnnology, Cornel! University, Ithaca, N.Y. 14850.
'Section of Insects and Spiders, Carnegie Museum, Pittsburgh, Pa. 15213 'Manuscript received by the editor April 24, 1970.



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19701 Cad, Shapiro, Clench - Lethe 71
Figs. 1-2. Lethe apfaiathia male, McLean Bogs Reserve, Tampkina Co., New York. Figs. 3-4. Lethe eurydice (male) neotype, Morris Arboretum, Philadelphia Co., Pennsylvania. Figs. 5-6. Lethe ewydire fumosa male, Sarpy Co., Nebraska.




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prairie regions from Minnesota and South Dakota to Indiana, Nebraska and Colorado.
(2) Lethe appalachia R. L. Chermock, widely distributed from Maine to northern Florida, westward to South Dakota and Alabama, broadly sympatric with L. e. eurydice but occurring primarily in swamp forest, shrub swamps and forest-edge ecotones. These forms have had a particularly extensive history in earlier literature, involving not just the above names but several others as well, as may be seen below.
TAXONQMY
In reviewing the taxa of the Lethe eurydice group, we have given the synonymies in strict chronological order, following the practice of some of the older authors. We hope this practice will add his- torical perspective to the discussions of these intricate synonymies. The distributional data and maps (figs. 21, 22) were based on material examined in the following institutional collections: Ameri- can Museum of Natural History, New York (AMNH) ; Academy of Natural Sciences of Philadelphia (ANSP) ; United States Na- tional Musuem, Washington, D.C. (USNM) ; Carnegie Museum, Pittsburgh (CM) ; Cornell University, Ithaca, New York (CU) ; New York State Museum, Albany (NYSM) ; and Hope Depart- ment of Entomology, Oxford University, England (Oxon. ) . Also the following private collections : Robert H. Whittaker (RHW) ; David J. Horn (DJH) ; Arthur M. Shapiro (AMS). A few reliable printed or other records, based on specimens not seen, are given separately, along with a few which are queried. The sexes of Lethe are very similar and may at times be difficult to determine in a superficial examination. We have therefore listed only the localities and dates of material examined. In the case of large series, only a range of dates may be given. The characters differentiating the three entities recognized in this paper are summarized in Table I and in the accompanying section of the text. A brief summary of the most conspicuous characters of each is given following its taxonomic history. Lethe eurydice eurydice (Johansson)
Papillo eurydice Johansson 1763, Amoen. Acad. 6 : 406 ; type locality Phila- delphia [Pennsylvania]; type formerly in the De Geer collection (Stockholm, Sweden), now lost ; neotype designated below. Papilia canthus Linnaeus 1767, Syst. Nat. (12th ed.) : 768; type locality "in America septentrionali;" no type exists (replacement name for Papilla eurydice) .




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? Papilio canthus: Fabricius 1775 (partim), Syst. Ent.: 486. Argus canthus: Scopoli 1777, Introd. Hist. Nat.: 432. Satyrus canthus: Godart 1821, Encycl. Meth. 9: 465, 493. Neonympha canthus: Westwood 1851, in Doubleday, Westwood, and Hew- itson, Gen. Diurn. Lep. 2: 375.
Neonympha cantheus (nec Godart 1821, see below) : Morris 1860, Cat. Lepid. N. Amer.: 10.
Hipparchia boisduvallii Harris 1862, Ins. Inj. Veg. (Flint ed.) : 305, fig. 128 ; type locality "this State" (Massachusetts), type now lost; no neotype designated.
Debis canthus: Herrich-Schaeffer 1865, Correspbl. 2001.-Min. Ver. Regens- burg 19: 72.
Pararge canthus: Butler 1868, Cat. Satyridae Br. Mus.: 123. Euptychia canthus: Kirby 1871, Syn. Cat. Diurn. Lep.: 55. Pararge boisdudii: Edwards 1872, Synopsis N. Amer. Butt.: 26. Argus eurydice: Scudder 1872, Syst. Rev. Amer. Butt.: 6. Satyrodes eurydice: Scudder 1875, Bull. Buffalo Soc. Nat. Sci. 2: 243. Satyrodes canthus: Smith 1884, Bull. Brooklyn Ent. Soc. 6: 119. ? Satyrodes canthus ab. boweri F. H. Chermock 1927, Bull. Brooklyn Ent. Soc. 22: 118; type locality Port Hope, Ontario; type not located, stated by dos Passes to be in Carnegie Museum, but not found. (Infrasub- specific name with no standing in nomenclature.) Satyrodes eurydice transmontana Field 1936, J. Ent. 2001. (Pomona) 28: 22; type locality Compton, Quebec; no type designated. Satyrodes eurydice transmontana f. ? rawsoni Field 1936, J. Ent. 2001. (Pomona) 28 : 22 ; type locality Bloomfield, Michigan ; type deposited in U. S. National Museum. (Infrasubspecific name with no standing in nomenclature.)
Lethe (Enodia) eurydice: R. L. Chermock 1947, Ent. News 58: 29. The descriptions of both eurydice yohansson and canthus Linnaeus are too scanty to restrict on internal evidence to either of the sympatric northeastern species, both of which occur at the type locality (Shapiro, 1g7oa). If that locality (Philadelphia) is accu- rate, there can be no doubt that Johansson's description applies only to a member of this group, even though no mention is made of eyespots on the forewing above (an objection to this usage, raised by Harris, 1862 and Edwards, 1897). No type of eurydice or canthus (which was proposed explicitly as a replacement name for eurydic~ and hence has the same type) exists in the British Museum (Natural History) or in the De Geer collection at the Naturhis- toriska Ri ksmuseum, Stockholm.
When appalachia (see below) was described as the southern sub- species of eurydice, the latter name became firmly associated with the assumed "northern" subspecies whose color and pattern were contrasted with appalachia by Chermock. It seems desirable to



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stabilize the nomenclature by preserving this usage through a neo- type designation. This removes the possibility that a specimen of Chermock's appalachia might eventually be selected as neotype of eurydice, leaving the familiar "northern" insect's name in question. The only Philadelphia specimens of eurydice auct. with full data which we have found were collected by one of us (Shapiro). Several of these were placed in the United States National Museum two years ago, and we desiginate one such specimen the neotype of Papilio eurydice Johansson.
Neotype. -A male deposited in the U.S. National Museum bearing the manuscript label "eurydice d/Morris Arboretum/Phila. Co. Pa./29 June 1967/A. M. Shapiro" (fig. I). We have added a label identifying the specimen as the neotype of PapiZio eurydice Johansson. The U.S.N.M. also contains a second specimen with the same locality and collection date.
Taxonomic History: the Euptychia names. - The taxonomy of L. eurydice is complicated by confusion with Yphfhimoides (= Euptychia) argulus (Godart). This ~roblem was not noted by dos Passos, and is reviewed here.
Fabricius (1775) reworked the description of canthufJ adding "immaculatis" to the upperside diagnosis and altering various details. The "immaculatis" may have been by inference from the lack of reference to spots in the earlier descriptions, but it seems more likely that Fabricius was working from some other insect he confused with the Linnean one.
In 1779 Cramer described and figured a
species from Surinam as Papilio argante. This name is a junior homonym of PapiZio argante Fabricius I 7 75 (now Phoebis argante, Pieridae) . Fabricius synonymized argante Cramer to canthus ( Fabricius, I 78 I ) , improperly emending it to argant he in synonymy. (Arganthle is not available as a replacement name because it was proposed in synonymy.) He repeated this usage in 1787 and 1793. His own descriptions of "canthusJ' do not fit Cramer's figure well. Godart ( I 82 I ) recognized that three species were included in the Fabrician concept "canthusJJ and attempted to end the confusion by redescribing the true canthus (translating Linnaeus), and naming two new entities, mgulus and cantheus. Godart's argulus is a re- placement name for the preoccupied argante and is the oldest valid name for this taxon. Cantheus is a renaming of the entity Fabricius first thought was canthus, theretofore without a valid name. The identity of this animal cannot be determined if, as appears, Fabrician specimens of "canthusJJ do not exist.




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Fig*. 7-8. Lethe apfalachia female, McLean Bogs Reserve, Tompkins CO., New York. Figs. 9-10. Lethe ewfdice eurydice female, Md-ean Boga Re- serve. Tompkina Co., New York. Figs. 11-12. Lethe eurydice fumosa fe- -.
male, Sarpy Co., Nebraska.




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76 Psyche [March
Butler ( I 868) described Euptychia perfuscata and subsequently ( 1869)
synonymized it to argante. We have not seen his speci- mens (which should be in the British Museum), nor any specimen definitely determinable under any of these names. The most recent discussion of argulus is by Weymer ( 1907, p. 202). The leading North American authority on Euptychia and its allies, L. D. Miller, advises us (in Hit.)
that he does not know the species, but from Cramer's description and figure would place it near Yphthirnoidies grimon. The partial synonymy of argante = argulus is thus: Papilio argante Cramer 1779 (nec Papilio argante Fabricius 1775), De Uitlandkch. Kapell. 3: 19, pi. 204; type locality Surinam; type not investigated.
$ Papilio canthus (nec Linnaeus 1767) : Fabricius 1781 (partim), Spec. Ins. 2: 64
(arganthe in synonymy) ; 1787, Mant. Ins. 2: 31; 1793, Ent. Syst. 3(l) : 157.
Satyrus argulus Godart 1821, Encyl. Meth. 9: 463, 488; type locality Surinam ; type never existed (replacement name for argante) . ? Euptychia perfuscata Butler 1868, Cat. Satyridae Br. Mus.: 18; type locality Para, Brazil; type probably in British Museum, not investigated. ? Euptychia argante: Butler 1869, Cat. Diurn. Lep. Fabr. Br. Mus. 13. Cantheus, which is the unknown animal Fabricius confounded first with canthus and then with argante, usually appears in the synonymy of eurydice = canthus, but its only proper claim there is its mis- taken use in synonymy by Morris (1860). We have removed cantheus Godart from the synonymies of the other entities and regard it as a nomen dubwm, presumably a species of Euptychia sens. lat. Its synonymy is :
$ Papilio canthus (nec Linnaeus 1767) : Fabricius 1775 (fartim), Syst. Ent.: 486; 1781, Spec. Ins. 2: 64; 1787, Mant. Ins. 2: 31; 1793, Ent. Syst. 3 (1) : 157.
Satyrus cantheus Godart 1821, Encyl. Meth. 9: 465, 493; type locality TAmerique septentrionale": type not investigated, probably never existed.
Godart's description of cantheus erroneously cites Fabricius, "Species Insectorum" for "Mantissa Insectorum." The name is misspelled "cautheus" in the heading on page 465. Taxonomic History : other names. - Gosse ( I 841 ) attributes the name Hipparchia transmontana to Say, but it does not appear in any extant work by that author. It usually appears in the synonymy of eurydice attributed to Gosse 1840. Apparently Gosse believed it was described elsewhere, since his "description" (1840, p. 247) is




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Fig. 13.
Male genitalia of Lethe appalachia.
Fig. 14. Male genitalia of Lethe eurydice eurydice.



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78 Psyche
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inadequate to associate it with any biological entity. It could apply to any of several Quebec Satyridae, which are not exhausted by the other species enumerated in the text. His statement that "this is likewise described by the American naturalists as very rare, and is found only beyond the Rocky Mountains . . ." makes no sense when applied to any species of Lethe. We regard this as a nonzen nudum and have omitted it from the synonymy.
Field ( 1936) resurrected transmontana as the northern sub- species of eurydice describing it adequately and giving as the type locality Gosse's base at Compton, Quebec. This is the oldest valid publication of the name, which should thus be credited to Field 1936. This subspecific distinction was grounded in confusion over the entities now called eurydice and appalachia. Observing differences between northern eurydice and specimens from near the type locality, Philadelphia, which he took as typical of that taxon but which were really appalachia, Field felt that a subspecific name was war- ranted. This is clear from his article, particularly the citation of Clark's ( 1932) figures of Beltsville, Maryland appalachia which Field (like Clark) calls typical eurydice. Thus transmontana be- comes a junior subjective synonym of eu.rydice. We c?n see no subspecific differences among eastern populations of eurydice as here restricted. Field's female form rawsoni is based on specimens faded in life; such specimens occur throughout the range of eurydice. The name is infrasubspecific and therefore has no formal standing. The name boisduvallii was attributed by dos Passos (1964) to Morris ( 1862), an error corrected later (dos Passos, 1969). Morris published the name in synonymy, spelled boisduvalli. The first valid publication was in the posthumous (1862) edition of Harris's "In- sects Injurious to Vegetation," edited by Flint. The editor's preface makes clear that the name should be attributed to Harris. It was emended to boisduvalii by Scudder ( I 889) in synonymy ; this spelling is used by Forbes (1960) and dos Passos ( 1964). DOS Passos (1969) has further emended it to boisduvali. Although not the preferred form, the double "i" is acceptable in taxonomy as the genitive of the Latinized name, i.e. "Boisduvalius." While Boisduval spelled his name with only one "1" and there is no orthographic reason to double it in forming the Latin genitive, the fact that the Fig. 15. Male genitalia of Lethe eurydire fumosa (valve in slightly different position than valves of figs. 13 and 14). Fig. 16. Ventral view of Lethe ap-bafachia valve. Fig. 17. Ventral view of Lethe eur~dice eury- dice valve. Fig. 18. Ventral view of Lethe eurydice fumosa valve.



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80 Psyche [March
name is spelled boisduvallii in both the text and index of Harris (1862) indicates that the double "I" was the form used in the Harris manuscript, and this is confirmed by its use in Morris's ( I 862) citation from that manuscript. Unfortunately, then, boisdu- valli is technically a "correct original spelling" and cannot be emended under the Code.
Summary of Characters. - Lethe eurydice eurydice is relatively pale, ~inkish brown above, somewhat yellowish beneath, with the postmedial line deeply indented beneath on all wings. The male valve is strongly 4-sided when viewed laterally, and the tegumen is dorsally rounded. The larva appears superficially to have dark side- stripes on the head capsule, extending from near the tip of the horns to the ocelli. These and other characters are discussed more com- ~letely under the headings "Diagnostic Characters" and "Biological Differences," after the taxonomic treatment of Lethe appalachia, below.
Distribution
(fig. 2 I ) . - Material examined :
DELAWARE : New Castle Co. : Blackbird, vii.62-65 ( AMS) ; Kent Co. : Smyrna, vii.62-65 (AMS)
PENNSYLVANIA : Philadelphia Co. : Morris Arboretum, vi.29.67 (AMS) (USNM) , Tinicum Wildlife Preserve, vi-viii.60-68 (AMS) , George's Hill, no date (P. Laurent) (ANSP) ; Montgomery Co.: Cheltenham Twp., Horsham Twp., Montgomeryville, vi-viii.58-68 ( AMS) , Pennsburg, vi.2 1-66
( AMS) ( USNM) ; Bucks Co. :
Buck-
ingham, East Rockhill, Ivyland, Chalfont, Trevose, vi-viii.58-68 (AMS) , Bristol, vi.27.67 ( AMS) (USNM) ; Chester Co. : vie. Down- ingtown, vi-viii.58-68 (AMS) ; Susquehanna Co. : vii.22-3 I. ? (CM) ; Lackawanna Co : Scranton, vii.4.05 (M. Rothke) (CM) ; Elk Co. : Medix Run, vii.16.64 (H. K. Clench) (CM) ; Beaver Co.: New Brighton, vii.?.o3 (W. C. Wood coll.) (AMNH) ; Mercer Co.: 2 mi. SE Leesburg, vii.11.66 (H. K. Clench) (cM), North Liberty, viii.3.60, viii.g.59 (J. Bauer) (CM ) ; Allegheny Co. : Nadine, vii.2g.24 (CM) ; Erie Co.: Presque Isle, no date, vii.7.26, vii. ?.40 ( CAT)
NEW JERSEY: Camden Co. : Westville, Haddon Heights, Atco, Magnolia, vi-viii.58-68 (AMS) ; Burlington Co.: Mt. Holly, Whites- bog, vi-viii.58-68 (AMS) ; Gloucester Co.: Woodbury, Glassboro, vi-viii.58-68 (AMS) ; Atlantic Co.: Da Costa, vi-viii.58-68 (AMS) ; Mercer Co. : Pennington, Washington's Crossing State Park, Dutch Neck, vi-viii.58-68 (AMS) ; Ocean Co.: Lakehurst, New Egypt, vi- viii.58-68 (AMS) ; Middlesex Co.: Jamesburg, vii.8.32 (A.S. Pin-



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19701 Curdi, Shapiro, Clench - Lethe 81
kus) (AMNH) ; Union Co.: Elizabeth, vii.13.? (0. Buchholz) (AMNH) ; Sussex Co. : Arlington, vii. I 3.18 (0. Buchholz) (AMNH), Hopatcong, no date (C. Palm) (AMNH), "Sussex Co." vii.1.43, vii.6.41 (0. Buchholz) (AMNH) . NEW YORK:
New York Co.: West Farms, no date (J. Angus) (AMNH) ; Queens Co.: Flushing, vii.27.18 (E. L. Bell) (AIMNH) ; Kings Co.:
East New York, vii.P.03 (W. C. Wood) (AMNH) ; Suffolk Co. : Calverton, v.26.25, vii.14.29 (R. Latham) (CU) , Orient, vi.2.38 (R. Latham) (cu) ; Richmond Co.: Staten Island, no date (Barnes coll.) (USNM), "S.I." no date (USNM) ; Rock- land Co.: Spring Valley, vii.20.68 (E. L. Rittershausen) (AMS) ; Orange Co. :
I mi. E Monroe, vii.21.68 (E. L. Rittershausen) (AMS) ; Westchester Co. : Somers, no date, viii.g.16, vii.3 1.26 (W. C. Wood) (AMNH), Bedford, no date (R. B. Dominick), vii.17.37 (A. C. Frederick), vii. 16-1 8.37 (all AMNH) , Lake Wacabuc, vii. 14.10 ( AMNH) ; Sullivan Co. : Lava, vi. ?. ? (Barnes coll.) (USNM) ; Albany Co. : Karner, vii.1 I .03 (J. Cook) (Oxon.), Albany, vii.24.27, vii.25.32 (A. C. Frederick) (AMNH), vii.7.28 (A. C. Frederick) (cu) ; Otsego Co.:
Cooperstown, vii.27.24 (B.
Smith) (cu) ; Cortland Co.: 2.7 mi. W Willet, viii.2.68 (AMS), McGraw, vi.8.14 (Engel coll.) (CM) ; Tompkins Co. : McLean, vie. Tompkins Co. Airport, Cayuga Inlet Valley, Michigan Hollow, Ringwood Hollow, Wilseyville, vi-ix.67-69 ( AMS) ; Schuyler Co. : Texas Hollow, vii-viii.68 ( A~MS) , Watkins Glen, vii.19-68 (AMS) ; Yates Co. : Potter Swamp, vi.14.15 (cu) ; Oswego Co.: Minetto,
vi.22.38 (W. T. M. Forbes) (cu) ; Livingston Co.: Lakeville, vii.18.27 (E. A. Maynard) (NYSM) ; Clinton Co.: Pittsburgh, vii.2.96, vii.1g.93 (G. H. Hudson) (NYSM) ; Columbia Co.: Ghent, viii.?.31 (AMNH) ; Saratoga Co.: Saratoga Lake, vii.8.28 (A. C. Frederick) ( AMNH) ; Jefferson Co. : Wellesley Island, viii. I 3.68 ( L. L. Pechuman) ( AMS) , Thousand Islands, vii. I 2.09 ( AMNH ) , Clayton, no date (J. H. Stebbins) (AMNH) ; Cattaraugus Co. : Crystal Lake, vii.6.30 (J. G. Franclemont) (CU) ; Erie Co.: Chafee, vi.18.32 (J. G. Franclemont) (cu), Buffalo, no date (C. V. Riley coll.) (USNM) ; Lewis Co. : vii.18.76 (W. W. Hill) (USNM) , vii. I 5.47 ( c. P. Kimball) ( AMNH) ; Monroe co. : vii.2.48, vii.23.46 (C. P. Kimball) (AMNH) ; Ontario Co.: Fishers, vii.30.48 (C. P. Kimball) (AMNH) ; Orleans Co.: Oak Orchard Swamp, vii. I 6.68 (AMS) ; Genesee Co. : Oak Orchard Swamp, vii. I 6.68 (AMS) ; County undetermined: "NY," no da.te (G. D. Hulst coll.) (AMNH), "vie. NYC" no date (S. L. Elliot) (AMNH)



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82 Psyche [March
CONNECTICUT: Tolland Co.: Rockville, no date (Engel coll.) (CM) ; Litchfield Co.: Litchfield, vii.1.94, vii.15.94 (L. B. Wood- ruff) (AMNH) ; Windham Co.: Putnam) vii.21.50 (A. B. Klots) (AMNH) ; County undetermined: "Ct." no date (G. D. Hulst coll.) ( AMNH)
MASSACHUSETTS : Worcester Co. : Winchendon, vii.3. ? (J. A. Gross- beck) (AMH), Princeton, no date (W. T. M. Forbes) (CU) ; Middlesex Co. :
Wayland, vii.7.2 I (cu) ; Silver Hill, vie. Lincoln, vii.7.23 (figured by Clark, 1932, pi. I, figs. 5, 6) ; County unde- termined : ('Mass." no date ( Barnes Coll.) (USNM ) NEW HAMPSHIRE: Cheshire Co.: Dublin, 1899 (A. H. Thayer) (Oxon.), West Rindge, vii.15.60, vii.10.61 (D JH) ; Coos Co. : Jefferson, vii.15-ZI.? (Engel coll.) (CM) vii.7.32 (G. & J. Sperry) (AMNH), Shelburne, vii.4-10.01 (USNM), White Mts., no date (H. Edwards) (AMNH) ; Grafton Co. : Franconia, no date (A. T. Slosson) (AMNH) ; Sullivan Co. : Claremont, 1908 (USNM) ; County undetermined: "N.H." no date (H. Edwards coll.) (AMNH)
VERMONT: Windham Co. : Stratton, vii.21.37 (H. Kahl) (CM) ; Rutland Co.: Mt. Killington, 4000', viii.17.40 (AMNH) ; County undetermined : vie. Sandgate, vii.13.49 (A. B. KIots) (AMNH) MAINE : Piscataquis Co. : Greenville, vii.2 1-29. I 9 (I?. Haimbach) (CM) , Sebec Lake vii.24-3 I.? (Barnes coll.) (USNM) ; Hancock Co.: Bar Harbor, vii.3.38 (A. E. Brower) (USNM), North Blue- hill, vii.21.23 (AIMNH), Mt. Desert, vii.?.? (W. C. Wood) (AMNH) , vii.13.33 (0. Buchholz) (AMNH) ; Kennebec Co. : Au-
gusta, vii.23.38 (A. E. Brower) (USNM), vii.22.50 (A. E. Brower) (AMNH) ; Penobscot Co. : Orono, no date (M. Fernald) (cu), Bangor, no date (Engel coll.) (OM), vii.10.89 (E. A. Smyth) ( USNM) , Passadumkeag Bog, vii.1-7.? (W. Sweadner coll.) (CM) , vii.12.34 (A. E. Brower) (AMNH), vii.2.33 (L. P. Grey) (TJSNM), South Lincoln, vii.15.50 (L. P. Grey) (AMNH), Lincoln, no date (L. P. Grey) (AMNH), (J. C. Hopfinger) (USNM), vii.io.40 (J. C. Hopfinger coll.) (USNM) ; County undetermined : "Maine," no date (E. A. Smyth) (USNM)
OHIO: Stark Co.: Waynesburg, vii.21.29, vii.4.30 (AMNH) MICHIGAN: Allegan Co.: Douglas Lake, vii.10.30 (H. C. Will) (CM) ; Livingston Co.: Pinckney, vii.9.? (cM), vii.9.39, vii.23.39 ( AMNH) , George Reserve, Pinckney, vii.23.38, vii.3 I .38 (G. W. Rawson) (USNM) , " Livingston Co." vii.9.32 (G. W. Rawson) (USNM) ; Branch Co. :
no date (B. Stroup) (cu) ; Otsego Co. :




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19701 Card;, Shapiro, Clench - Lethe 83
Sturgeon River, 5 mi. E Vanderbilt, vii.8.55 (Klots & Rindge) (AMNH), 7 mi. E Vanderbilt, vii.7.55 (F. H. Rindge) (AMNH), Lake Otsego, vii.7.55 (F. H. Rindge) (AMNH), Pigeon River, I I mi. E Vanderbilt, vii.8.55 (Klots & Rindge) (AMNH) ; Cheboygan Co.: vii.6.52, vii.19.52 (H. V. Daly) (AMNH) ; Emmett Co.: 6 mi. W Pellston, vii.9.55 (Klots & Rindge) (AMNH), Petoskey, vii.8.13, vii.8.14 (J. J. Lichter) (AMNH), Galloway Lake, North Levering, vii.9.55 (Klots & Rindge) (AMNH) ; Huron Co.: Hume Twp. Arboretum, vi.28.52 (H. V. Daly) (AMNH) ; School- craft Co.: Thompson, vii.10.55 (Klots & Rindge) (AMNH) ; Oak- land Co.: New Hudson, vi.20.27 (G. W. Rawson) (USNM), Bloomfield, viii.4.29 (G. W. Rawson) (AMNH), viii.12.28 (G. W. Rawson, paratype of rawsoni Field) (USNM), viii.4.29 (G. W. Rawson) (USNM) ; Washtenaw Co. : Willis, vii.30.39 (AMNH) , Sharon, vii.2.44 (G. W. Rawson) (USNM) ; County undetermined: 'Michigan" vii.8.90 (AMNH), Green Oak, vi.25.33 (G. W. Raw- son) (AMNH), Calvin, vii.3.90 (AMNH), "Snow I., Lake Michi- gan," no date (CM)
INDIANA : Steuben Co. : vi.16.03 ( AMNH) ; Lake Co. : Hessville, vii.4.08 (E. Beer) (USNM)
ILLINOIS: Lake Co.: "NE Lake Co." viii.24.30 (H. IVL Bower) (AMNH) ; Cook Co.: Chicago, vii.6.13 (1. D. Gunder coll.) (AMNH)
MINNESOTA: Ramsey Co.: St. Paul, no ckte (Barnes coll.) (USNM) ; County undetermined: "Minn." no date (AMNH) WISCONSIN : Douglas Co.: 2 mi. E Maple, vii.11.55 (Klots & Rindge) (AMNH) ; Milwaukee Co. : Milwaukee, vii.10.08, viii.5.17 (H. M. Bower) (AMNH) ; Waukesha Co. : Dousman, vii.14.16, vii.20.19 (H. M Bower) (AMNH) ; Dane Co.: Madison, no date (E. T. Owen coll.) (USNM) ; County undetermined: "Wis." no date (A. T. Slosson) (AMNH), (E. T. Owen coll.) (USNM) NOVA SCOTIA: Cape Breton, viii.?.4g (G. 3lacmillan) (CM) ; Cape Breton National Park, viii.?.54 (H. Dietrich) (CU) NEW BRUXSWICK : Bathurst, viii.5-6.5 I (A. B. Klots) (AMNH) QUEBEC: Dunlop Rd., Gatineau Provincial Park, vii.6.52 (F. H. Rindge) (AMNH) ; Montreal, vii.1.29 (J. C. Hopfinger coll.) (USNM) ; "Quebec" vii.3.35 (J. C. Hopfinger coll.) (USNM) ONTARIO: Ottawa, bred (W. H. Edwards) (cM), vii.23.97 (M. Holmes) (Oxon.) ; Toronto, viii.g.24 (R. N. & F. A. Dixey) (Oxon.), vii.19.15, vii.20.18 (H. V. Andrews) (CM) ; Bancroft, vii.1-7.? (W. Sweadner coll.) (CM) ; Spider Lake, Georgian Bay,



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8 4* Psyche [March
vii.22.14 (G. K. Jennings) (CM) ; Point au Bad, vii.21.35 (E. D. McDonald) (cu) ; Don Valley, Toronto, vii.8.57 (J. C. E. Riotte) (AMNH) ; Sudbury, vii.7-10.59, vii.7-10.60 (J. C. E. Riotte) (AMNH), vii.7.60 and vii.12.58 at UV lights (J. C. E. Riotte) ( AMNH) ; Gravenhurst, Muskoka Dist., iv.7.18 ( ! ) ( AMNH) ; Geraldton, Ashmore Twp., vii. I 6.55 (Klots & Rindge) ( AMNH ) ; Grand la Cloche, vi.27.41, vii. I ,41 (0. Buchholz) ( AMNH) ; Leam- ington, Essex Co. vi.?.go (E. A. Smyth coll.) (USXM) ; ('Ont." no date (Blackmore coll.) (USNM)
MANITOBA : Aweme, vii. I 5.07 (CM) , vii. I 9.08 (Barnes coll.) (USNM) ; Riding Mts., vii.11.38, vii.g.39, vii. 2-3.40 (C. S. Quelch) (AMNH), vii.17.38 (J. F. May) (AMNH), vii.11.38 (G. W. Raw- son) ( USNW) ; Transcona, vii.19.48 (C. S. Quelch) (AMNH) ; Birtle, vii.7.44, vii.1g.44 (J. Dennis) (AMNH) ; Telford, White- shell Provincial Park, vii.24.55 (Klots & Rindge) (AMNH) ALBERTA: Rivercourse, near Lloydminster vii.6.41 (R. J. Fitch) cu)
Other records : DOS Passos ( I 969) erroneously records eurydice from "south to Colorado and east of the Rocky Mountains to Georgia and Florida." The Colorado records represent L. e. fumosa (see below). At present the southernmost record of true L. e. eury- dice is northern Delaware.
Scudder (1889) records L. eurydice from Rupert's Fort, Quebec (east shore Hudson's Bay) ; Alingan, Labrador; and Great Slave Lake, NWT. All of these are plotted on the map. The western distribution of eurydice is unclear. Puckering and Post (1960) record it from Cass, Cavalier, Dickey, Grand Forks, and Pembina Cos., North Dakota. These are entered on the map. We have seen no South Dakota records. However, Leussler (1938), who was well acquainted with L. e. fumosa, reported typical eiirydice in Sioux Co., northwestern Nebraska. This record seems to require special confirmation, and has not been plotted. The "eurydice" reported from Monroe Co., Tennessee (Mather, 1961) was appalachia (W. Reinthal, pers. comm.). The latter species was found at Jackson, Tenn. by Roever (Mather and Mather, 1958).
Lethe eurydice funzosa (Leussler)
Satyrodes canthus n.v. fumosus Leussler 1916, Ent. News 27: 99, pi. iv, figs. 1, 2; type locality Sarpy County, Nebraska; type reportedly de- posited in Ohio State University, not seen. Lethe furnosus: dos Passos 1969 (partim), J. New York Ent. Soc. 77: 120.



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19701 Card;, Shapiro, Clench -Lethe 8 5
Taxonomic History. - Described from 17 males and 8 females, all labeled "Omaha" by Leussler as are various later topotypes. Al- though it was described as a "variety," the geographic nature of fumosa was clearly expressed.
The Greek noun in611 is feminine and retains its gender in the Latinized form Lethe. Both the species names in this group are also feminine. We have adopted the spelling fumosa to make this subspecies agree in gender, as provided by the Code. Summary of Characters. - Lethe eurydice fumosa resembles L.e. eurydice in most respects, but the males and some females are darker above. The four eyespots on the forewing are consistently graded in size, from the smallest on top to the largest at the bottom; this is especially obvious beneath. The male valves have far fewer setae than in L. e. eurydice. The early stages are unknown. Distribution (fig. 2 I ) . - Material examined : NEBRASKA : Sarpy Co. : "Omaha" vi.28.13, vii. I. I 5 ( R. A. Leusslei-) (cotypes) (ANSP) ; vii.12.13, vi.14.13, vi.27.14 (cotype) , vii.7.17 (R. A. Leussler) (USNM) ; vi.27.14 ("topotype") , vii.5.13, vi.28.13, vi.26.15 (cotype), vii.7.17 (R. A. Leussler) (AMNH) ; vi.27.14 (paratype), vii.1.16 (figured by Holland, pi. 63, fig. 11) (R. A.
Leussler) (CM) ; County undetermined : "Nebraska," no date (J. Angus coll.) (AMNH)
IOWA: Dickinson Co.: Lake Okoboji, vi.25.21 (R. A. Leussler) (USNM) ; Hancock Co.:
I mi. W Klemme, vii.24.60 (L. D. Miller) ( CM ) ; Powesheik Co. :
Grinnell, vii.4.8 I ( AMNH ) ; County un- determined: "la." no date (H. Skinner) (CM) WISCONSIN : Kenosha Co. : Twin Lakes, vi. i -4. i I (A. Kwiat) (USXM) ; County undetermined: "Wis." no date (E. T. Owen coll.) (USNM)
MINNESOTA : Hennepin Co. : Lake Minnetonka, no date (USXM) , St. Anthony Park, vii.15.91 (USNM)
INDIANA : County undetermined : Ti-emont, vii.24.48 (0. Buch- holz) (AMNH)
SOUTH DAKOTA: Brookings Co.: Volga, no date (P. C. Truman) CM)
COLORADO: Larimer Co.: Loveland, no date (W. H. Edwards coll.) (CM) ; County undetermined: "Cola." not date (David Bruce) (cM), "Colorado" no date (E. T. Owen coll.) (USNM), "Colorado" no date (H. S, Burnett) (CSNM)



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86 Psyche [March
Bruce collected fumosa at Estes Park, Larimer Co., Colorado (Edwards 1897, p. 197). The identities of the Indiana, Minnesota, and Wisconsin specimens noted above were not checked by the genitalia, and are somewhat uncertain.
Lethe appalachia R. L. Chermock
Lethe (Enodia) eurydice affalachia R. L. Chermock 1947, Ent. News 58: 29; type locality Conestee Falls, North Carolina; type in R. L. Chermock collection, not seen.
L&e fumosus appalachia: dos Passos 1969, J. New York Ent. Soc. 77: 121. Taxonofnic History. - Unmistakable figures of appalachia appear in three older works under other names. None of these has any taxo- nomic significance. Boisduval and Le Conte (1829) figure a male appalachia with an ambiguous female as Satyrus canthus (pi. 60). Edwards
(1897) figures a female appalachia (pi. 26, figs. 3, 4) with a normal male eurydice (figs. I, 2) as Satyrodes canthus, along with a dark male which is probably also eurydice but might be fumosa (fig. 5). Denton (1900) figures an ambiguous specimen (2. 2 I 7), an eurydice (p. 2 18). and an appalmhia (p. 2 19), all as Neonymph~~ canthus.
DOS Passos ( I 969) erred in sinking appalachia to "/uvzosus." There is no evidence for his statement that "fz~mosus and appalachia occur at opposite ends of a cline."
DOS Passos lists ab. boweri I?. H. Chermock under appalaclhia. It cannot be identified to species by the description, and the type has not been found; it is not in the Carnegie Museum, where dos Passos recorded it. We have placed boweri provisionally in the synonymy of eurydice because that species is considered more likely from the type locality, Port Hope, Ontario. However, a specimen of appalachia with no ocelli on the forewings above, labeled "Bowie, Md./ v-29-45/DDT experiment," is in the U.S. National Museum. At any rate, the name is clearly infrasubspecific and has no standing. Summary of Characters. -Lethe appalachia differs from both sub- species of eurydice in being grayish or mousy brown above (blackish when fresh) and somewhat purplish or lilac-tinged beneath; the postmedial lines rounded, with only slight indentations. The male valve is less clearly 4-sided in lateral view, and the tegumen is dorsally flattened. The larval head capsule bears side-stripes not reaching below the bases of the horns.
Distribution (fig. 22). - Material examined : FLORIDA : Jefferson Co. : Monticello, x.4.I 4 (paratype) ( AMNH)



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19701 Gar&, Shapiro, Clench -Lethe
Fig. 19.
Larval head capsule of Lethe appalachia. Fig. 20. Larval head capsule of Lethe eurydice eurydice. SOUTH CAROLINA: Jasper Co.: Coosawhatchie, vii.26.36 (R. B. Dominick) (paratype) (AMNH) ; Clarendon Co.: ix. ?.89 (E. A. Smyth) (USNM) (see Smyth, 1890)
NORTH CAROLINA: Mitchell Co.: vii.?.92 (0. Buchholz) (AMNH) ; Transylvania Co. : Conestee Falls, vii. I -7. ?, vii. I 5-2 I. ? (CM ) VIRGINIA: Amherst Co. : 1936 (J. Bauer) (CM) ; Prince George's Co.: New Bohemia Swamp, viii.22.67 (J. Bauer) (CM) ; Nanse- mond Co. : Dismal Swamp, vi.19.40 (CM) ; Giles Co.: Little Meadows, vii.25-26.40 (L. Carr) (paratypes) (USNM) ; Mont- gomery Co.: viii.20.98 (E. A. Smyth) (USNM) ; Fairfax Co.: Vienna vii.19.38 (A. H. Clark) (paratype) (USNM) ; Wythe Co.: Speedwell, viii.11.38 (A. H. Clark) (paratype) (USNM) ; Grayson Co. : Long's Gap, viii.11.38
(A. H. Clark) (paratype) (USNM) ;
County undetermined : Glen Carlyn, viii. I 2. ? (A. N. Caudell) (paratype) ( USNM)
DISTRICT OF COLUMBIA: Washington, vii.17.29 (G. W. Rawson) ( USNM) , vi.29.29 (paratypes) ( USNM) , vi.17.29 (paratype) (fig- ured by Clark, 1932, pi. I, figs. 3,4) (USNM) MARYLAND : Prince George's Co. : Hyattsville, vi.20.39 (G. W.



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88 Psyche [March
( Rawson) (USNM) , Bowie, v.29.45 (USNM) , Beltsville, vii.15.28 (paratype) (USNM) ; Calvert Co. : Mutual, vi.21.36 (G. W. Rawson) (USNM)
DELAWARE: New Castle Co.: Christina, vii.64 (AMS) PENNSYLVANIA : Crawford Co. : Hartstown, vii.4.21 (H. Kahl) CM), viii.8-14. ? (CM) ; Mercer Co. : North Liberty, viii.3.60 (J. Bauer) (cM), 2 mi. SE Leesburg, vii.11.66 (H. K. Clench) (CM) ; Butler Co. : Slippery Rock, vii.4.3 I ( W. Sweadner) (CM ) ; Fayette Co. : Dunbar, vii.6.3 I (CM) ; Westmoreland Co. : Powder- mill Nature Reserve, vi-viii.56-68 (H. K. Clench) (CM) ; Chester Co. : Exton, vii-viii.59-68 (AMS) ; Montgomery Co. : Horsham Twp., Cheltenham Twp., Enfield, vi-viii.59-68 (AM) ; Delaware Co. :
Chadd's Ford, vii-viii.62-66 ( AMS) ; Philadelphia Co. : Mt. Airy, v.28. ? (P. Laurent) (AMNH) , Tinicum Wildlife Preserve, Eastwick, vii-viii.58-68 (AMS)
NEW JERSEY: Cape May Co. : Woodbine, vi-viii.59-67 (AMS) ; Camden Co. : Westville, viii.6.92, viii. 14.92 (P. Nell) (CM) , vi. 12. ? (P. Laurent) (CM) (AMNH) ; Burlington Co.: Warren Grove, Wading River, vii-viii.64-68 (AMS) ; Gloucester Co.: Wenonah, viii.21 .IO (CM) ; Ocean Co. : Cassville, viii.17.10 ( AMNH) ; Morris Co.: Green Village, vii.30.? (C. Rummel) (cM), vii.15.? (C. Rummel) (ANSP), "Morris Co." vii.21.50 (P. Ehrlich) AMNH) , vii.9.30, vii.6.41, vii.18.50 (0. Buchholz) ( AMNH) ; Union Co.: vi.16.40 (0. Buchholz) (AMNH) ; Somerset Co.: Orange Mts., vi. 14. ?, viii.20.3 I (0. Buchholz) (AMNH) ; Bergen Co. :
Ramsey, viii. ?.I 7 (AMNH ) ; Passaic Co. : Paterson, vii. I 7. ? (J. A. Grossbeck) (AMNH) ; Sussex Co. : Springdale, vii.9.49 (P. Ehrlich) (AMNH), vii.10.49 (N. W. Gillham) (AMNH), Lake Lackawanna, vii.9.49 (P. Ehrlich) (AMNH) , Stanhope, vii.28.33 (C. Rummel) (AMNH), Hopatcong, no date (C. Palm) (AMNH), "Sussex Co." vii.9.30, vii.6.41, vii.1.43, vii.18.50 (0. Buchholz) (AMNH) ; County undetermined : "N. J." (Neumoegen coll.) (USNM), "N.J." (C, Palm) (AMNH)
NEW YORK:
New York Co.: West Farms, no date (J. Angus) (AMNH), Bronxville, vii.9.11, vii.22-23.11 (L. B. Woodruff) (AMNH) , Bronx, bred (E. Gerstenkorn) (AMNH) ; Suffolk Co. : Riverhead, vii.7.49, vii.8.49, vii. I 7.52, vii.5.53, viii.1 I .53 (R. Latham) (CU) , Orient, vi.30.41, viii.2.49, vi.17.52 (R. Latham) (cu), East Hmpton, vi.14.49 (R. Latham) (cu), Green- port, viii.1.20, ix.1.51, ix.6.28 (R. Latham) (cu), Calverton, vii.8.30 (R. Latham) (cu) , Brookhaven, vii.5-13.65 (RHW) ;



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19701 Card;, Shapiro, Clench -Lethe 8 9
Queens Co.: Flushing, vii.27.18 (E. L. Bell) (AMNH) ; West- chester Co. : Bedford, vii.7-9.34, vii.17.37 (R. B. Dominick) (AMNH), Somers, no date (W. C. Wood) (AMNH), Crugers, vii.16.12 (AMNH) ; Dutchess Co. : Fishkill, ix.7.65 (D JH) ; Sul- livan Co.: Lava, no date (Barnes coll.) (USNM) ; Albany Co.: Albany, vii.24.27 (A. C. Frederick) (AMNH), Karner, vii.7.70 (Lintner) (USNM) , vii.20.79 (W. W. Hill) (NYSM) , vii.11.03 (J. Cook) (Oxon.) ; Tompkins Co. : McLean, vii.18.91 (local col- lection, CU) , viii.7.25, vii.27.29, viii.i.25 (CU) , vii.18-21.68 (AMS), Sapsucker Woods, viii.7.68 (AMS) ; Schuyler Co.: Texas Hollow, viii.3.68 (AMS) ; Orleans Co.: Oak Orchard Swamp, vii.16.68 (AMS) ; Genesee Co. : Batavia, vii.16.87 (CU) ; Cattarau- gus Co.: Allegany State Park, vii.21.40 (A. R. Shadle) (USNM) ; Franklin Co.: Paul Smith's, vii.?.03 (A. P. Hunt) (Oxon.) ; County undetermined : "New York" (H. Edwards coll.) (AMNH) CONNECTICUT: New Haven Co.: Sound View, vii.16-21.34 (A. H. Clark) (USNM) ; Litchfield Co.: Sharon, vii.14-21.40, vii.?.41 (L. J. Sanford) (AMNH) (one figured by Klots, 1951, pi. 10) ; Fairfield Co.: Stamford, vii.22.37 (J. G. Thorndike) (AMNH) ; Hartford Co.: Avon, viii.1.03 (R. C. Williams) (cM), vii.18.22 (R. C. Williams) (ANSP)
MASSACHUSETTS: Suffolk Co.: Newton Highlands, no date (W. Barnes) (USNM) ; Hampden Co.: Wilbi-aham, viii.?.94 (cu) ; Bristol Co. : Swansea, vii.18-22.34 (E. T. Learned) (ANSP) RHODE ISLAND: Providence, vii.10-20.? (H. Engel) (cM), North Scituate, 1912 (G. H. & J. L. Sperry) (AMNH) NEW HAMPSHIRE:
Coos Co.: Jefferson, vii.7.32 (G. H. & J. L. Sperry) (AMNH), "White Mts." no date (W. H. Edwards) (CM) MAINE: Penobscot Co. : Bangor, no date (CM) QUEBEC : "Quebec," vii.3.35 (J. C. Hopfinger coll.) (USNM) ILLINOIS: Cook Co. : Chicago, vii.6.13 (J. D. Gunder coll.) (AMNH)
INDIANA : Lake Co. : Hessville, vii.4.08 (E. Beer) (USNM) ; County undetermined: "Indiana," no date (E. A. Smyth coll.) (USNM)
MICHIGAN : Huron Co. : Hume Twp. Arboretum, vi.28.52 (H.V. Daly) (AMNH) ; cass co.: Wakelee, viii.3.58 (L. J. Sanford) (AMNH) ; County undetermined: "Michigan," no date (AMNH) MINNESOTA : Hennepin Co. : Lake Minnetonka, viii. ?.86 (AMNH) WISCONSIN: County undetermined: "Wis." no date (E. T. Owen coll.) ( USNM )




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90 Psyche [March
SOUTH DAKOTA: Brookings Co.: Volga, no date (Ehrman coll.) CM)
MISSOURI: St. Louis Co: St. Louis, vii.io.? (CM) We regard the South Dakota and Missouri records of L. appa- lachia as somewhat dubious, but they are shown on the map. Other records: The following are probably accurate, although the specimens have not been seen. They are included on the map: GEORGIA: Thomas Co. : Linton Lake, viii.9.67, viii.29.67 ; Fulton Co. : Atlanta (Harris Trail), vii.29.60, viii.20-26.61 : De Kalb Co. : Avondale Estates, vi. I 6.44 ; Union Co. : Copper Creek State Park, vi.6.58, vii.16.61, vi.18.62, vii.18.62, viii.22.59; White Co.: Cleveland, vi.13.57 (all from L. Harris, unpublished MS, P. 244)
MINNESOTA : Anoka Co. : Bald Eagle Lake, 1966 (Masters, 1967) INDIANA: Steuben Co.: Hogback Lake, vii.17.42 (Price and Shull, 1969)
TENNESSEE: Madison Co.: Jackson (Mather and Mather, 1958) ALABAMA : Tuscaloosa Co. : vie. Tuscaloosa (Chermock, 1949) MICHIGAN" : Montcal~n Co. : Sidney, vii.26.50 (F. Rutkowski) PENNSYLVANIA: Fayette Co.: Markleysburg Bog, 2 mi. N Mark- leysburg (H. K. Clench)
DIAGNOSTIC CHARACTERS
Color and Pattern.- Lethe e. eurydice and L. e. fumosa differ subtly but consistently from L. appalachia. Nearly all specimens can be assigned to the correct species by color and pattern alone. The most useful characters separating the two species are the ground color above and beneath, and the waviness of the postmedial line beneath. The two subspecies of eurydice differ most consistently in the relative sizes of the forewing ocelli. All of the observed differences are given in Table I.
We have not seen a truly fresh specimen of L. e. furnosa. Leussler ( 1916) describes the ground color of fresh specimens as "a very dark smoky grey . . . even a blackish appearance." This sounds very much like the color of newly emerged appalachia. Old speci- mens of the two are very different, however: funzosa males are an even, somewhat purplish or reddish brown, while appalachia is grayish or mousy brown. Some female fumosa, particularly from Colorado, are nearly identical in color to nominate female eurydice, but the ground color of the males is nearly always distinctive. A few male eurydice from the northeast are dark purplish brown when



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19701 Card, Shapiro, Clench -Lethe 91
fresh, and fade to an even dark reddish brown. Their spot-sizes are normal and they lack the fumosa tendencies to "high angledness" of the forewing and blind and rimless ocelli above. A specimen of this dark form of nominate eurydice is probably represented by Edwards' figure 5 ( 1897, pi. 26).
Specimens of the three taxa are shown in figs. 1-12. Male Genitalia - Chermock ( I 947) and dos Passes ( I 969) re- ported no genitalic differences between L. e. eur~jdice and L. appala- chia. However, we have found that they do differ slightly but significantly. The tegumen of appalachia is flattened dorsally, while that of eurydice (both subspecies) is rounded. The valves of appa- lachia are shorter and narrower dorso-ventrally, and from the side appear less quadrilateral than those of the eurydice subspecies. The male genitalia of L. c. eurydice and L. e. fumosa are substantially similar, but differ from each other and from appalachia in the density and arrangement of setae on the valves. See Table r and figs. 13-18.
Female Genitalia. -There seem to be no useful characters here. Some minor differences in the sclerotization of the genital plate were found among all three taxa.
Early Stages. - The larvae of L. e. eurydice and L. aptalachia from central New York differ consistently in the maculation and tubercles of the head capsule. In L. c. eurydice the red side stripes become darker below the bases of the horns, extending to the ocelli. The darker part of the stripe consists of small, heavily pigmented, regu- larly arranged tubercles on a less heavily pigmented ground. In L. appalachia the stripe does not extend below the horn, and its lower end contains several large, pale, irregularly placed tubercles which contrast with the red ground (figs. 19, 20). The early stages of L. e. fumosa are completely unknown. Developmental Rate. -Larvae of L. e. eurydice and L. appalachia from AlcLean, N.Y. reared ex ovo at 24OC on late summer photo periods showed developmental differences. Eurydice larvae invariably entered diapause in the third or fourth instar. Appalachia larvae usually developed without diapause, the entire life cycle requiring about 60 days. Lethe appalachia is at least double-brooded in its southern range; apparently it has the potential to breed continuously



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92 Psyche [March
Table 1.
Differences separating taxa of the Lethe eurydice group. Color terminology follows Kornerup and Wanscher (1963) and Ridgway ( 1912).
character
I. COLOR AND PATTERN
postmedial line of forewing beneath
postmedial line of hindwing beneath
ground color beneath (fresh males)
ground color beneath
(fresh females)
ground color above (fresh males)
ground color above (fresh females)
color between ocelli and subterminal
line beneath
color marginad of postmedial line
on hindwing beneath
contrast between discal and limbal
areas on forewing above
rings around ocelli above
ocelli of forewing beneath
ocelli 4, 5 on hindwing above
apex of forewing
11. MALE GENITALIA
valve shape
valve costa
valve setae
tegumen
111. LARVAL HEAD CAPSULE
red stripe
tubercles in red stripe
IV. HABITAT
'only old specimens seen
"fades in life to Van Dyke brown
L. e. eurydice
projects marginad into teeth at Ms,
c uz
projects marginad into teeth at CUI,
cuz
red-haired (6C4) = wood brown
greyish orange (5B5) = clay color
sunburn (6D5) === snuff brown
clay (5D5) = Saccardo's umber
darker than ground, tinged with
orange
lighter than ground, yellowish
especially in Ms
moderate to strong
usually strong
variable, subequal, 4 usually largest
usually pupilled
tending to be "low angled'ld
4-sided in lateral view
inner lip larger
valve tip heavily armed;
many setae on sacculus
dorsally rounded
top of horns to ocelli
red, small, regularly arranged
open sedge marshes
(6F6) = bistre
'fades in life to brown (6E5) brownish olive



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Cardi, Shapiro, Clench - Lethe
L. e. fumosa L. appalachia
as in L, e. eurydice
cinnamon (6D6) == Rood's brown
greyish orange (5B5) = clay color
cocoa brown (6E6) := cinnamon
browna
sahara (6C5) = sayal browna
as in L. e. eurydice
as in L. e. eurydice
slight ( $ ) to moderate ( $ )
weak to strong
usually 4>3>2>1
frequently unpupilled
'high angled" in $ $ only
as in L. e. eurydice
as in L. e. eurydice
few setae on valve tip or sacculus
as in L. e. eurydice
unknown
unknown
permanent marshes
within prairie region
smoother, only slightly wavy
smoother, only slightly wavy
dark blonde (5D4) = buffy brown
topaz (5C5) = avellaneous
teak (6F5) = mummy brownb
teak (6F5) = mummy brownc
paler, not orange-tinted
lighter than ground, with violet
iridescence
moderate to strong
weak to strong
usually 1 and 4 largest
frequently unpupilled
frequently "high angled"
(both sexes)
less +sided ; short ;
narrower dorso-ventrally
inner lip smaller
some setae distally and on sacculus
dorsally flattened
horns only
irregular, large, pale
swamp forest, shrub swamp, forest-
edge ecotones
"high angled": ratio of length of forewing (base to apex) : outer margin < 1.5.




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9 4 Psyche [March
(without diapause) elsewhere as well. So far as is known, eurydice is single-brooded everywhere. Larvae of both species turn from yellow green to straw yellow when in diapause, and are capable of changing color in either direction overnight.
A usually small emergence of fresh eurydice occurs in some localities in New York, New Jersey and Pennsylvania in the first half of August, four to five weeks after the principal emergence. Males of this late "brood" are frequently of the dark form noted above. It is very unlikely that these butterflies are descendants of those which emerged a month earlier. There may be a genetic basis for the emergence times; a bimodal emergence of Hyalophora cecropia (L.) (Saturniidae) was recently reported by Sternburg and Wald- bauer (1969) with no genetic data. We do not believe the late eurydice are identical with funzosa, but the slight possibility exists that they represent another sibling species, unrecognizable in the adult except by its flight period and a statistical color difference. We have not obtained ova from these insects. Food Plants. - DOS Passes
(1969) speculates that a food plant
difference between L. eurydice and L. appalachia is likely. How-
ever, our observations suggest that both are sedge-feeders and that neither is species- or group-specific within Carex. Female appalachia occur near sedges in shrub swamp or forest habitats where observa- tion is difficult. One oviposition was seen in the field, on Carcx lacustris Willd. (Cyperaceae) at Texas Hollow, Schuyler Co., N.Y. Other sedges commonly associated with this species in New York, all of which were completely acceptable in the laboratory, are Carex gracillima Schwein., C. lanuginosa Michx., and Scirpus georgianus Harp. Wild hosts of L. e. eurydice in central New York include C. lacustris, C. stricta Lam., C. rostrata Stokes, and C. trichocarpa Michx. All of these sedges were fully acceptable to both species, as are some dozen other species tested (mostly undeter- mined). We reared both species from egg to adult on Carex torta Boott. Neither species would accept any of the following grasses (Gramineae) : Festuca ovina L. ; Elynzus ri-s L. ; Brachyelytrum erectum (Schreb.) Beauv. ; Muhlenbergia schreberi Gmel. ; Agrostis alba L. ; Phalaris arundinacea L. ; Leersia oryzoides (L.) Sw. ; Echi- nochloa crus-galZi (L.) Beauv. (B. erectum and P. arundinacea are wild food plants of Lethe portlandia anthedon A. H. Clark, and M. schreberi is acceptable in the laboratory; Shapiro and Card;, 1970.)




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19701
Cardi, Shapiro, Clench - Lethe
95
Fig. 21. Distribution of Let/zc eurvdicc euryd'ice (solid circles) and Lethe eurydice fmosa (solid triangles).
Add/å Behavior. -The most striking difference between L. e. eurydice and L. ajtpalachia, and the one leading to the discovery of their sympatry, is their differential habitat selection (Shapiro and Card;, 1970). At the McLean Bogs Reserve, Tom~kins Co., New York, these two species are frequently found flying within a few feet of each other, but do not mix. The preference of L. apwlachia for shaded habitats often results in its association with L. jt. anthedon upland or L. p. portiandia on the Coastal Plain. We have found L. c. eurydice only in relatively open sedge marshes 01- rarely, in drier meadows: it never enters dense shrub swamp or woods. ^Ve. have seen L. eurydice and L, p. antl~edon in co$ul~ in their usual habitats, once each (3 p.m. and 3 :30 p.m., respectively). DISCUSSION
Although the term "sibling species" has been in the literature for nearly thirty years and the concept is even older, it still seems necessary to point out that excessive dependence on morphological



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96 Psyche [March
Fig. 22.
Distribution of Lethe appalachia.
differences can hinder the recognition of such biologically interesting species as those of the Lethe eurydice group. Despite abundant mu- seum evidence of sympatry, these species went unrecognized for twenty years after Chermock (1947) was unable to find genitalic differences between them.
As usually happens with sibling species, recognition on biological grounds has led to discovery of morphological characters hitherto overlooked. These, however, are of a magnitude which would not be considered diagnostic of species in most groups of Lepidoptera. In fact, the genitalia seem to be among the most conservative characters in Lethe. Chermock found only very minor genitalic differences between L. portlandia and L. creola Skinner in the other American species group, and circumstantial evidence suggests that portlandia itself is really a pair of (largely allopatric) sibling species. Many Asiatic Lethe we have examined also show only slight differences among themselves and from their close American relatives. We consider it likely that what we are calling Lethe eurydice fumosa may also prove specifically distinct when its biology



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19701 Card& Shapiro, Clench -Lethe 97
- particularly the early stages - becomes better known. Similar
cases recently uncovered in the Lepidoptera include the tortricid moths Archips argyrospiLus and A. mortuanus, which differ only in sex attractant and in some characters of the last-instar larva (Roelofs and Comeau, 1969), and the papilionid butterflies Papilio zelicaon and P. gothica, said to differ consistently only in host-plant specificity but to behave as species in genetic tests (Remington, 1968). The Holomelina aurantiaca complex (Arctiidae) , often thought to consist of two species, actually includes at least ten, exceedingly similar in genitalic morphology, color and pattern, but differing in chromo- some number ( Cardt!, unpublished).
The seemingly inevitable problem with sympatric sibling pairs such as Lethe eurydice and appalachia is to account evolutionarily for the
"elegant" manner in which they coexist. The view that reproductive isolating mechanisms and ecological differences evolve in response to deleterious hybridization and competition in secondary sympatry (Brown and Wilson, 1956) is now very widely accepted. It was recently challenged by Ehrlich and Raven (1969), who proposed that isolating mechanisms usually develop during the genetic differentiation of allopati-ic populations under different selective regimes. This is in effect a reformulation of the view of most nineteenth- and early twentienth-century evolutionists. At- tempting to explain the ecological relationship of a given set of sibling species requires consideration of the following points: I. The apparent absence of ecological interaction (e.g., competition) or gene flow between presently sympatric populations does not rule out such events in the past, nor for competition, in the future. Furthermore, intermittent large-scale gene flow between normally allopatric populations, associated with fluctuations in population sizes, has probably been an important component of speciation ( Brown, 1957). Such fluctuations could also result in episodes of competition between otherwise non-competing species. 2. Biogeographical evidence may offer important clues to episodes of prior sympatry or allopatry in the evolution of species differences ( cf. IIengel, I 964).
3. In the absence of evidence for character displacement, it cannot be assumed that biological differences which appear to prevent com- petition evolved in response to the adverse effects of competition. The ecological differences among the American species of Lethe can be resolved into two parts: that involving the eurydice group alone and that concerning the eurydice and portlandia groups. The



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98 Psyche [March
species eurydice, appalachia, and portlandia (in the broad sense, including anthedon) divide neatly into "non-competing" pairs : the two sedge feeders (eurydice and appalachia) differ in habitat; the two woodland species (appalachia and portlandia) differ in larval food plant. (Similarly, in sexual behavior, enrydice and appalachia are essentially non-territorial ; portlandia is strongly territorial.) Lethe and the genera closely related to it are hypothesized to have originated in southeast Asia (Miller, 1968), a region with many forest-dwelling, grass- (mostly bamboo-) feeding representa- tives of both the eurydice and portlandia groups. It seems reason- able that the ancestors of both these groups migrated to North America via the Bering land bridge in the Arcto-Tertiary forest, and were forced southward by the events of the Pleistocene. With the vast majority of the many Asian Lethe feeding on grasses, the evolution of sedge feeding in North America by the ancestor of the eurydice group is a tempting hypothesis. Evolution of this trait by proto-eurydice independent of competition with proto-portlandia or by character displacement in syrnpatry with it are both possibilities. T. Shirbzu (pers. comm.) informs us that Lethe 71zargInaZis Motschulsky, which seems to be a member of the eurydice group, feeds on non-bamboo grasses and on sedges in Japan as does Kirinia epaminondas Staudinger formerly placed in Lethe. Ninguta ("Lethe") schrenckii Menetries is an obligate sedge feeder- On the other hand, the speciation of eurydice and appalarhia may have occurred when one of the Pleistocene glaciations isolated some populations of proto-wrydice in prairie to the west of populations in the eastern Austral forests. Virtually the entire range of mydice wa? glaciated, and the distribution is therefore of recent origin. The same can be said for the northern portions of the ranges of a~alachia and portlandia, but the southern portions are characteristic of many organisms which presun~ably survived the Wisconsin (and earlier glaciations) in the southeast. The lack of recorded relict populations of eurydice south of Pennsylvania in the Appalachians, if not due to inadequate collecting, suggests that the species did not have a Wis- consin refugium in the forested Austral Zone of the southeast; its habitat preference and developmental rate support this interpretation. (We do know of species of Hesperiidae, e.g. Euphyes binzacula, with ranges and biologies substantially similar to L. eurydice, which have relict populations in the southeast; Shapiro, 1970b.) Its most prob- able refugium, then, was in glacial Transition Zone somewhere west of the Appalachians. The existence of L. e. fumosa also supports



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19701 We, Shupiro, Clench -Lethe 99
a prior western distribution for eurydicle, but does not help in dating it. We have no grounds for estimating evolutionary rates in this group; all that can be said now with some confidence is that eurydice and appalachiu were more likely allopatric than sympatric in the Wisconsin (and appalachia and portlandia more likely sym- patric) .
The critical evidence concerns character displacement. We have found no morphological character displacement in sympatric vs. allopatric populations of Lethe eurydice and L. appalachia. One of us (Clench) believes he has observed behavioral character dis- placement between them in Pennsylvania, within the area of general sympatry; in certain localities where only one species occurs, it ap- pears that the habitat selection is not so rigorous as elsewhere. This needs additional study and quantification. Another geographic area also bears close investigation in this connection. Specimens of eurydice from southeastern New York (Orange, Rockland, and Westchester Counties) are somewhat anomalous, tending to vary in color and pattern (but not genitalia) toward appalachia. We have seen very few specimens from outside this small area which we would hesitate to classify to species by color and pattern. The area is completely surrounded by normal, sympatric, well-differentiated populations of both. It is thus critical to determine the ecology of these anomalous insects. Should appalachia be rare or absent, and twrydice occupying its niche at least in part, one would have a powerful argument for character displacement as the origin of the habitat difference. (There
is a chance of natural hybridization due to man's extensive disturb- ance of Lethe habitats in southeastern New York.) There are, then, two basic questions: Did the behavioral and food plant differences between the American eurydice and portlandia groups evolve independently, or largely as a result of competition ? Did the sharp habitat selection between eurydice and appalachia in close sympatry evolve in isolation, or was it intensified by behavioral character displacement ?
On the first point, any evolutionary scenario will require much more comparative data on the Asiatic species than is readily avail- able. Only a comprehensive revision, identifying the closest relatives of the American species and comparing their biologies, will allow a convincing argument.
On the second point, field studies in areas of allopatry will be critical. It should be noted that while we suspect the eurydice - appalachia habitat difference may have evolved to prevent competition



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100 Psyche [March
for larval food, it also may function, and have evolved, as a repro- ductive isolating mechanism.
Greater knowledge of mating behavior
in this group (Shapiro and Cardi, 1970)) as well as more informa- tion on the southeastern New York populations, may be able to dis- tinguish the correct hypothesis.
Our current state of knowledge does not allow us to choose between independent evolution and character displacement in ac- counting for the differences between the American eurydice and portlandia groups. But character displacement is an attractive hypothesis for the eurydice - appalachia habitat selection difference. ACKNOWLEDGMENTS
We wish to thank William D. Field of the U.S. National Museum and Dr. Alexander B. Klots of the American Museum of Natural History for their advice and cooperation; Dr. Michael Emsley for the loan of specimens of Lethe eurydice funzosa from the Philadelphia Academy of Natural Sciences; N. D. Riley of the British Museum (Natural History) and Dr. T. Nyholm of the Naturhistoriska Riksmuseum (Stockholm) for aiding in the search for the early types; Dr. Lee D. Miller for information on Euptychia spp.; Lucien Harris, Jr. and Frank Rutkowski for field notes and unpublished locality data; Edward L. Rittershausen for specimens from south- eastern New York; Dr. Ta.kashi Shiizu, Kyushu University, Japan, for data on Asiatic Lethe; and Dr. Peter A. Hyypio for deter- mining species of sedges. Dr. John G. Franclemont of Cornell University provided invaluable assistance in the taxonomic research. Mrs. Adrienne R. Shapiro assisted in color determinations and in the preparation of the manuscript. Dr. John Burns made several helpful suggestions.
This research received partial support from National Science Foundation Grant GB 7757 (Environmental Biology) and from the Grace Griswold Memorial Illustrations Fund, Department of Entomology and Limnology, Cornell University. To both we extend our thanks.
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