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K. W. Cooper.
A Southern California Boreus, B. notoperates n.sp. I. Comparative Morphology and Systematics. (Mecoptera: Boreidae).
Psyche 79(4):269-283, 1972.

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PSYCHE
Vol. 79 December, 1972 No. 4
A SOUTHERN CALIFORNIAN BOREUS,
B. NOTOPERATES N. SP.
I. COMPARATIVE MORPHOLOGY AND
SYSTEMATICS ( MECOPTERA : BOREIDAE ) *
BY KENNETH W. COOPER
University of California, Riverside
INTRODUCTION
AH but one of the 26 or so of the species of Boreus now known from Eurasia and North America (listed in Svensson 1972) are found chiefly within the bounds of north temperate or boreal forests where winters are cold. The northern margin of their distribution is roughly the oå¡ mean annual isotherm, in marked contrast to the more gentle climatic requiren~ents of the vast majority of Mecoptera. Though distributional limits are now probably poorly known, the southmost regions from which Boreus have been reported have mean annual temperatures generally less than 12.Så¡ (55OF), and in all cases there is a winter period of snow cover. Because the adults are charming insects, bizarre and easily recognized but seen alive by but few, and are most often collected on the surface of snow, active and mating at temperatures below freezing, it is not surprising that a fairly large, markedly anecdotal literature has developed about them. Despite frequent notes and longer narratives (nowhere fully re- viewed), there are few extensive accounts of their biology. Indeed Strubing's (1950) admirable study of Boreus hyemalis (L.) is the only substantial account available of the life history of a boreid. My own experience with Boreus extends over many years. As others before me, I have always associated Boreus with cold, snow, fairly high annual precipitation (20 inches or more), and north tem- perate forest.
It was with near disbelief that I found two female pupae of Boreus, on a very hot, dry autumn afternoon in an Upper Sonoran region of chaparral and yellow pine, in arid southern Cali- fornia. Not unexpectedly, this most southern of all Boreus proves to *Manuscript received by the editor November 28,1972 269




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2 70 Psyche
[December
be new, and in a surprising number of ways it is the most special- ized of all Boreus. In this account I describe Boreus noto-berates n. sp., discuss its taxonomic relationships and specializations, offer reasons for disregarding Euboreus Lestage into which the new spe- cies would otherwise fall, and record the latitudinal dispersion of Boreus in the Holarctic. An account of the environment and discus- sion of host and habitat mosses, as well as other biological n,otes, will be the subjects of a second article.
Boreus notoperates Cooper, n.sp.
Diagnosis
Boreuy notoperates n. sp. (figs. 1-2) may be separated from all other Boreus now known by the following combination of characters: male and
female dark, with light wings ; antennae I 9-segmented, frons foveolate, no median ocellus, occipital foramen not divided by a sclerotized corporatentorium, hypostomal bridge very long; prono- turn with 3
transverse folds; male: 2.7 mm long; forewing not spined on anterior margin; hindwing spined on posterior margin, without ventral brush; without tergal apophyses, 9th tergum with a glabrous, undivided, parallel-sided notch for styli ; hypandrium emarginate; internal, submedian tooth of style simple, acute; female: 3.9 mm long; ovipositor short, cerci separated at tip, 10th tergum prolonged as a spined, upturned blade on each side of cerci; gona- pophyses strongly spined laterally and ventrally. Description
Coloration. Eyes plum-colored to brownish black ; head, nota, procoxae and abdomen shining black with a bronze or greenish glint. Rostrum on sides, apically, and below, dark reddish brown. Flanks of thorax, meso- and metacoxae piceous or black, yet appearing ashen due to fine pubescence. Antennae and palps nearly black. Legs yel- lowish-brown to piceous, tarsi darker apically. Modified wings light brownish-yellow (sooty brown in one male) to yellow, those of male piceous apically, of female piceous at basal attachment. Gonocoxites and styli black, tips of styli flavescent. Ovipositor black; cerci black, flavescent or pallid laterally near tips. Pubescence. Of male, gray, except bristles which are yellow to brown; moderately sparse on abdomen, as long or longer than basal width of metatarsus; denser and a third or less as long on thoracic pleura. Occiput glabrous, vertex and frons with sparse hairs as long as width of scape; on each side a triangular patch of fine silvery



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Figs. 1-7.
Boreus notoperates n. sp. All specimens in fluid, under mod- erate pressure from above ; camera lucida representations. - Fig. 1. Male, aedeagus everted. - Fig. 2. Female. - Fig. 3. Epandrium and proctiger, dorsal aspect. - Fig. 4. Gonocoxites, dististyles and elongated median plaque that overlies aedeagus, dorsal aspect. - Fig. 5. Hypandrium, dotted line across notch denotes limit of hyaline membrane present in teneral male, ventral aspect, apex above.-Fig. 6. Ninth and succeeding terga of ovi- positor, dorsal aspect.-Fig. 7. Gonapophyses, ventral aspect, apex below.



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272 Psyche [December
pubescence in a depression running from ocellus to margin of antenna1 condyle; patch of 5-10 hairs at clypeal base, but no rostra1 subocular patch, only scattered hairs; few, fine hairs on stipes. Forewings with coarse, scattered, yellowish hairs.
Long white hairs on anterior and
posterior faces of pro- and meso-coxa, and anterior face of metacoxa. Abdominal pubescence somewhat longer, denser ventrally and to- ward sides; tergum I glabrous medially, as are t4 to t7 medially and, increasingly so, laterally.
Of female: much as in male, except pubescence appears yellowish, and everywhere somewhat shorter; patch at base of clypeus may be absent; but few long hairs on posterior surfaces of pro- and meso- coxae. Abdominal pubescence denser, posterior margins of terga polished, nearly glabrous, as are most of terga 8 and 9. Head. Rostrum (from lower margin of eye to apex) ca. 2.1 times length of eye in male, 2.6 times in female; longer than pro- tibia in female, slightly shorter in male. Eye slightly longer than broad, narrowed below. Antenna with 19 segments, condyle oppo- site lower margin of eye; scape much wider than long, pedicel swol- len in apical third and shorter (by one-fifth) than the combined lengths of segments 3 and 4. Ocelli small, median ocellus lacking. Frons and sides of vertex coarsely foveolate. Occipital foramen single, not divided to an alaforamen and neuroforamen by a sclero- tized corporatentorium. Hypostomal bridge (between occipital fora- men and cardo) very long, longer than length of eye in female, somewhat less in male.
Thorax.
Of male: pronotum polished, with occasional transverse wrinkles; strong anterior and posterior transverse grooves divide the pronotum into anterior, median and posterior transverse folds of widths roughly as 2 : 3 : I ; on each side: two or three long (and several shorter) bristles on anterior lobe, three or more shorter bristles on median fold, and two to four long bristles set in trans- verse, more or less confluent depressions along hind margin of pos- terior fold in front of wing base. Outer margin of forewing nearly straight in dorsal aspect, inner margin slightly concave behind basal third; hind margin with ca. 15 (range: 13-18) strong marginal spines below which are directed obliquely outwards; only vestiges of spines on front margin; hooked apex terminates in a pair of strong apical bristles. Hind wing shorter, glabrous, with ca. 16 (range 13- 20) stout spines below, no ventral brush of fine pubescence; apex hooked, spatulate at tip. A dark spine at apex above each femoral anterior condyle.




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19721 Cooper --- B'orens 273
Of female: pronotum similar, but widths of pronotal folds rough- ly 3 : 4 : I ; bristles of posterior fold in shallow depressions. Fore- wing pad nearly twice as long as broad, without marginal spines, completely covering atrophied hindwing. Femoral spines as in male. Genital Segments.
Of male (figs. 3-5) : eighth tergum and ster- num, and gt and 9s not fused at their respective pleural junctions; ninth tergum deeply infolded for reception of styles, the notch-like pocket with nearly parallel sides, glabrous within and not divided; a loose cluster of short spines ( 12-1 5å ) externally on each lateral border of notch.
Dististyles with a row of 15å short spines along apical one-third of inner margin; submedian internal tooth acute, not keel-like, relatively short; a membranous element lies within an ovate fenestra (or stylocavernula), broadest basally, on inner sur- face of each dististyle just distal to the submedian tooth. Hypan- drium broad, strongly emarginate apically (emargination may be partially closed by a hyaline membrane which is lost or worn away in older individuals).
Of female (figs. 6-7) : ovipositor short, from base of tergum 10 to apices of cerci about two-thirds length of rostrum (from lower margin of eye to tip).
Tenth tergum prolonged on each side as a pair of upwardly curved divergent blades bearing 4-6 strong spines above, tips of blades reach distal third of cerci ; cerci separated apical- ly; gonapophyses strongly spined laterally in apical halves. Reproductive System.
Of male (fig. 8) : each testis consists of four lobes or "sperm tubes", each ensheathed in a rich russet-brown tunic, right testis slightly anterior to left, the apices of the two testes reaching only slightly cephalad of swollen main portion of vas de- ferens
(that is, the "accessory gland") ; calyx and efferent vas of each testis short, thick; vas deferens tapering markedly to juncture of accessory gland; accessory gland with 4 lobes, externally poorly delimited ; lateral ejaculatory ducts barely demarcated ; median ejacu- latory duct strong, very muscular, U-shaped ; fleshy aedeagus trilobed when everted,
Spermatmoan (fig. 8) 87-96 p long: head 22-26 p long; tail with an "undulatory membrane". Of female (fig. 9) : Six ovarioles per ovary, each ovariole with 7-1 I panoistic egg chambers conjoined apically to ovarian ligament which unites with that of other ovary; ovariolar pedicels irregularly conjoined to common ducts which merge to form a calyx; lateral oviducts short; common oviduct nearly as long as an ovary; sperma- theca reniform, enclosing 24 sperm receptacles, 12 per side; sperma- thecal duct very short (less than short axis of spermatheca) arising



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Psyche [December




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19721 Cooper - B'orens 275
from hilus of spermatheca, entering common oviduct at its caudal third; a pair of elongated, tubular accessory glands jo,in oviduct by a very short common duct immediately posterior to entrance of spermathecal duct. M,ature egg (fig. 9) buffy-white, from 0.53 X 31 mm to 0.60 X 30 mm, slightly narrowed at one end; chorion smooth.
Measurements (in millimeters; in each case ordered thus : mean of measurements, [range of measurements], number of examples measured.) : Males, body length (in 70% isopropyl alcohol) - 2.7 r2.6-2.91 5 ; body length (dry) - 1.9 [1.5-2.51 g ; antennal length - 1.66
f I .4-I .g]
13 ; rostra1 length (eye to tip) - 0.7 f0.6-0.81 13 ; forewing length - I .o [ag-I .I] 13. Females, body length (in alcohol) - 3.9 r3.4-4.21 6 ; body length (dry) - 3.0 r2.7-3.61 5 ; antennal length - I .7 r1.5-1-91 10; ros- tral length - 0.9 [0.8-I .o] 10 ; forewing length - 0.3 [0.27-0.341 10; ovipositor length (base of tergum 10 to tip) - 0.6 l0.56-0.641 10.
Holotype. Male: 2.5 cm long (dry mount), collected 8 January 1972 ; Allotype, Female : 3.6 mm long (dry), collected 27 Decem- ber 1970; I 5 male and 12 female paratypes, collected 31 December 1971 to 22 January 1972. All specimens from Coldwater Canyon, ca 4400 ft altitude, below the town of Mountain Center (at 33' 42' N/ I 16O 44' W), Mt. San Jacinto, Riverside Co., California; collected by Geoffrey, Tera, Ruth and K. W. Cooper. Types and 2 paratypes are deposited in the Museum of Compara- tive Zoology at Harvard University where types of the majority of North American species are located. A pair of paratypes has been placed in each of the following colllections: U. S. National Mu- seum, California Academy of Sciences, Snow Entomological Museum (University of Kansas), and the University of California at River- side. Dissections and remaining specimens are in my collection. Figs. 8-1 1. Bow, reproductive system. All figures in dorsal aspect ; camera lucida representations. - Fig. 8. Male, B. notoperates n. sp., 4 lobes per testis;
right testis lies anterior to left, tilt conceals 2 of the 4 lobes; to right, mature spermatozoan.-Fig. 9. Female, B. notoperates, 6 ovarioles per ovary; to left, spermathecal vesicles and connections to as- sembly-duct ; to right, mature (laid) egg - the chorion is smooth. - Fig. 10. Male, B. nivoriundus Fitch, 1 lobe per testis, right testis anterior to left; above, testes from right side-note that a short efferent duct from each testis enters a common (fused) calyx; testes are mottled with an orange-brown pigmentation.-Fig. 11. Female, B. nivoriundus, 7 to 10 ovarioles per ovary; there are 8 ovarioles in the figured right ovary; to right, mature (laid) egg-the chorion is microscopically vermiculate.



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276 Psyche [December
Etynlology. The name notoperates derives from the Greek: notos, masc., south, and perates, masc., wanderer, alluding to the remark- able extension by this species of the range of Boreus to the south. Males of Boreus notoperates will classify as B. isolatus Carpenter when Carpenter's keys ( 1935, 1936) for North American Boreus are used. However, they strikingly differ from that species by their considerably longer rostrum ( ca 2. I
X eye-length ; "scarcely longer
than the eye" in B. isolates). Females of the new species will be excluded at couplet 7
(revised key, Carpenter 1936) because the ovipositor of B. notoperates is but two-thirds the length of the ros- trum.
The only known North American form with which B. notoper- ates might be confused is the closely related B. brevicaudus Byers ( 1961 ) . These two are markedly alike by having a reduced number of antennal segments (most Boreus have 21 or more), condyles of antennae opposite lower margins of eyes, no median ocellus, a long hypostomal bridge, no e~andrial "hood", and the 10th tergum of the female is abruptly narrowed on each side to form a pair of spined terminal blades. However they are readily distinguished in both sexes, for B. notoperates has 19-segmented antennae ( 18 in B. brevicaudus) , the frons more conspicuously foveate, a more heavily and richly spined area to each side of the epandrial notch, a very much shorter, inconspic~~ous submedian tooth of the dististyle, an emarginate hypandrium (entire in B. brevicaudus) , a relatively longer forewing in the female (L/W = 1.7 X ; 1.3 X in B. hicaudus) which lacks subapical spines, a relatively longer ovi- positer (0.67 X rostrum versus 0.5 X ) with more coarsely and extensively spined gonapophyses and apical blades, and the apical blades are divergent, Though color differences are often of little weight, the males of these two species have the color patterns of their genital segments and dististyles reversed; what is dark in the one is light in the other, possibly continuing a distinction important at a time that their progenitors were sympatric. At least four of the twelve recognized Eurasiatic forms share with B. notoperates (and B. brcvicmidus) the following set of characters: a reduced number of antennal segments
(21 or less), a relatively
short ovipositer, a male having the forewing narrow at base and no (or nearly no) apophyses on abdominal terga 2 and 3, namely B. chadzhi-gireji Pliginsky ( I 9 I 5 ) and 11. vlasovi Martynova ( 1954),



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19721 Cooper - Borens 277
each with 19 antennal segments, 13. mmi Pliginsky ( 1915) (20 segs.) , and B. bey-bienkoi Tarbinsky ( I 962) (2 I segs.) . To these
may perhaps be added B. aktijmi Pliginsky (1915), described from a single female said by Pliginsky to have 19-segmented antennae and to be very like his B. chadzhi-girejiL. Nevertheless large differences separate each of these species from B. notoperfltes and B. brevicau- dus. None of the Eurasiatic species have the apex of tergum-10 abruptly modified as a blade on each side; B. chadzhi-gireji and B. vlasovi have the 9th tergum of the male fused laterally with its ster- nite (Martynova, 1954) (a free suture is present in both America,n forms) 2; and males of B. n w i and B. bcy-h'ienkoi have apical brushes of fine pubescence on the under surfaces of the hind wings (absent in the two American forms).
It is likely that all 5 of the
Eurasiatic species have short hypostomal bridges, like most Bloreus but unlike B. notopert;tes and B. brevicaudits; regrettably this is definitely known to be so only for B. chadzhi-gireji (see Pliginsky, 1915, fig. 9).
STATUS OF EUBOREUS
I have not used the generic name Euboreus (genotype Boreus nivoriundus Fitch) which Lestage ( I 940-41 ) proposed for all boreids of which the males lack transverse apophyses on abdominal terga 2 and 3. Lestage would restrict Boreus (genotype Boreus hyemalif 'According to Martynova (1954), Pliginsky's types of B. navasi and B. aktijari cannot be found, but his specimens of B. chadzhi-gireji were studied by her. She states that the tergal apophyses are almost (pochti) absent in the males of B. chadzhi-gireji. Pliginsky (1915) did not de- scribe the male of this species, but merely observed, in his commentary in German (there is an obvious misprint in the corresponding text in Rus- sian), that it differs from that of his B. navasi only by having 19 antennal segments; but the male of B. navasi is pointedly stated by him (p. 365) to be without apophyses on abdominal terga 2 and 3. 'Mickoleit (1971b) has briefly discussed the fusion of the 8th abdominal tergum and sternum in male Boreus. The condition of the 8th (and 9th) abdominal segments in male Boreus, however, is more complicated than he knew.
Symbolizing such fusion with +, we have for the palearctic spe- cies:
Boreus orientalis (8+, 9+), B. chadzhi-gireji (0, +), B. semenovi (0, +), B. vlasovi (0, +), B. hyemails (+, O), and B. westwoodi (+, 0) (see Martynova 1954) ; for the nearctic species: B, borealis (+, O), B. californicus (+, O), B. coloradensis (+, O), B. brevicaudus (0, O), B. brumalis (0, O), B. nivoriundus (0, O), and B. notoperates (0, 0). It seems that such fusions have little to tell of the larger affinities of Boreus that Mickoleit discusses, yet it seems clear that these fusion-patterns must feature importantly in any world-wide taxonomic revision of Boreus as now constituted.




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278 Psyche [December
(L.) ) to include only those species having such apophyses, an as- semblage of species geographically restricted to western Eurasia which he believes to represent a derivative, more highly evolved, endemic palearctic lineage. So far as now known, Lestage's Boreus would include only four or five species : B. hyemalis (L.) , B. west- woodi Hagen, B. lokayi Klapalik, B. kratochwili Mayer, and B. chadzhi-gireji to judge from Martynova's ( 1954) remarks. If rec- ognized, Euboreus would encompass all, or nearly all, of the re- mainder, some 15 North American species and at least 5 of the Eurasiatic forms: B. navasi, B. semenovi, B. orientalis Martynova, B. v/asowi, and B. bey-bienkoi (no males are known for either B. aktijari or B. sjoestedti Navis, 1926). In Lestage's view, the spe- cies falling into Euboreus represent "un type palioendin~ique prim- itif", an important notion (were it true) which features prominently in his evolutionary explanation of boreid zoogeography. One test of the degree to which Lestage's two genera are useful concepts is to enquire whether other characters, not ordinarily em- ployed
in taxonomies, independently suggest that these aggregates do reflect important phyletic distinctions even though main, exter- nally visible characters traditionally used in diagnosis are shared by certain members in each of Lestage's two genera. Certainly it would seem to be so if forms placed in Euboreus share as a group important internal and cytological dissimilarities from the members of Les- tage's Boreus.
There is not much information to draw upon, but what there is seems conclusive. The reproductive systems of Boreus hyemalis (Steiner 1937 ; Potter I 938) and "Euboreus" brumalis (Cooper I 940) are strikingly dissimilar in lobulation oaf testis, testicular calyx, and spermatheca, as well as in their sex chromosomal comple- ments. I have found that B. hyemali,~ has an XO (male) - XX (fe- male) sex determination. "Euboreus" brumalis, however, has XiX2Y - males and X,XiX2Xa females (Cooper 1951). Nothing is known of the chromosomes of "E." notoperates, nor have others de- scribed the chromosomes of additional species. But the genital sys- tems of "E." notoperates (figs. 8-9)) though peculiar by possession of a very short vas deferens, one less lobe per testis, and a distinc- tively organized spermatheca which has a duct of negligible length, are not as unlike those of "E." brumalis as they are those of B. hye- malis. Indeed the chief distinctions shown by "E." notoperates may be viewed as larger departures along the same paths as those by which "E." brumalis differs from B. hyenzalis, and thus wholly con- sistent with Lestage's views.




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19721 Cooper - Bioreus 279
However, I find that the genotypic species of Eiiboreus, namely B. nivoriunh, has XO (male) - XX (female) sex determination and genitalia that are extremely similar to those of B. hyemalis*, thus the testes are single lobed and have fused calyces (fig. IO), and there is an extremely elongated spermathecal duct in the female (fig. I I) that is "coiled like a watch spring", just as Potter (1938) described it for B. hyemalis. Interestingly, XO-XX male hetero- gamety is the general mode of sex determination for the Mecoptera so far studied (Panorpa, Ullerich 1961 ; Bittacus, Matthey I 950; Chlorista, Bush 1967; and Merofie, Cooper, ~npublished)~, and is most probably primitive; primitive also is the very elongated sperma- thecal duct, a condition which Potter (1938) places among the five "most striking of the typically Mecopterous characters of the order". There are, therefore, no grounds at present for viewing Lestage's Euboreus as a useful taxonomic set, or even for attributing to its members as a group the emphasis which Lestage gave them as more primitive forms. Quite to the contrary, "E." notoperates indeed seems to be the least primitive among all the Boreus known. Its "advanced" characters include the long hypostomal bridge, loss of the median ocellus and the corporatentorial bridge4, loss of the brush of fine pubescence on the hind wing, the unparalleled reduction of the submedian tooth-complex of the dististyles, the peculiar apicolat- era1 blades of the 10th tergum of the female, and the distinctive spermatheca with its very short duct - all exceptional departures from the usual conditions in the Boreidae. Except for the marked reduction of the submedian complex of the dististyles, B. brevicaudus shares all of the external peculiarities with B. notoperates, and quite likely it possesses most or all of the remaining anatomical departures as well. These two Boreus, then, are the most specialized boreids known. Conversely, members of Lestage's "Boreus" have a quite generalized morphology, and should the tergal apophyses of their males prove to be vestiges of notal appendages homologous to those of Notiothaumids, Panorpids, and Panorpodids ( Crampton I 93 I ; Mickoleit 197 I a), they are evidently the most "primitive" forms known (but see ftnt. 2). At best they comprise a species-group within Boreus as now defined, namely the hyemalis-group. 3Also see Atchley and Jackson (1970).
*Boreus nivoriundus (Otanes 1922), B. hyemalis (Slais 1947), B. uni- color and B. californicus (Hepburn 1969), and1 as I have found for B. brumalis, have 3 ocelli and a divided occipital foramen. At present the
Boreidae is the only family of Mecoptera within which the number of ocelli varies, and in which the occipital foramen occurs in both states: divided and undivided (see Hepburn's fine study).



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Psyche [December
EXTENT OF THE LATITUDINAL RANGE OF BOREUS Boreus not'operates now provides the southmost record (33.y0N ) of all known species of Boreus. Nevertheless, in eastern North America, B. nivoriundus and B. brumalis extend nearly as far south. Both range from Jackson, New Hampshire (Dohanian, 1915) and Ellsworth, Maine to Tennessee, in the Great Smoky Mountains at 4000 ft (Carpenter 193 I, 1939; Cole and Gillespie 1g50), namely from 44.S0N to a latitude perhaps as low as 35.4ON. In the far West B. californicus Hine has a comparable range: from Kaslo, British Columbia (Carpenter I 93 1 ) to Hobart Mills, Nevada Co., California (new record), or from about 49.g0N to about 39.4ON. In the West, only B. notoperates has been found south of 3g0N. The total latitudinal range of Boreus in North America is therefore from the vicinity of McCarthy and Kennicott (ca 61.5ON in Alaska, B. intermedius Carpenter and B. yracilk Carpenter; Carpenter 1935, 1936) to Mt. San Jacinto (^^.'/ON)
in California, a total range
of nearly 28O of latitude, or about 1900 miles. Were Lestage's ( 1940, p. I 6 ; I 941, p. I 19) listing of Persia a valid record for the range of B. lokayi Klapilek, it would perhaps provide the most southern outpost of Boreus in Eurasia. It is an error evidently originating from Enderlein's (1910) rendering of the type locality5, which had been cited simply as Sedmihrad by Klapilek ( I 901 ) . Now "Sedmihrad" is 'Czech for Siebenburgen (Transylvania, now in Rumania), and KIaphIek ( I 903) leaves no doubt for he says he has many specimens of B. lokayi that were col- lected by Lokay "in Buczecz in Siebenburgen". Nevertheless it is an odd coincidence that Boreus has been found just across the northern border of Iran, providing the southmost rec- ord in Eurasia. It is that for B. vlasovi from Ashkhabad, Turk- men S. S. R. (ca 37.g0N) (Martynova 1954). The northmost record is that of B. westwoodi from the island of Kil'din (69.3ON)) slightly NE of Murmansk, U.S.S.R. (Tarbinsky 1962). Although Boreus ranges nearly 8' farther to the north, it has a known lati- tudinal range in Eurasia not quite 4' greater (about 250 miles) '". . Boreus lokayi Klap.
1903 (Persien, Sedmihrad) . ." thus, Enderlein (1910, p. 394). Perhaps Enderlein refers to the Muntii Persani of the Sedmihrad that, near Brasov, run north from the southern Carpathians of which the Bucsecs (Buczecz, M. Bucegi) is a member. If so, it may represent a valid, otherwise unrecorded locality of capture, in addition to the type locality. The most recent record for B. lokayi appears to be that of Miller and Povolny (1950): High Tatra Mountains, about 1600 m, Czechoslovakia.




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than that in North America - a datum of considerable ecologic and zoogeographic interest.
It is pleasant to express my gratitude to those who have so gen- erously helped me. Dr. Hildegard Strfibing and Prof. Dr. Erich Wolf, of the Freien Universitiit Berlin, many years ago made spe- cial collections and cytological preparations of Boreus hyemalis for me. Dr. Hugh B. Leech, of the California Academy of Sciences, loaned me specimens of Boms borealis Banks. Prof. George Byers,
of the University of Kansas, loaned specimens of his Boreus brevi- caudus and B. coloradensis, and commented helpfully on a draft of the description of the externals of the new species. Prof. A. Foro- stenko, University of California at Riverside, and my son Geoffrey translated many of the Russian passages, and Prof. Louis Pedrotti, also of UCR, translated Czechoslovakian accounts by Lokay ( I 908, 1912) and Obenberger ( 1928) (all in Casopis) which dispel1 any residual doubt concerning Lokay's Sedmihrad collections. LITERATURE CITED
ATCHLEY, W. R. AND R. C. JACKSON
1970. Cytological observations on spermatogenesis in four species of Mecoptera. Canad. Jour. Genet. Cytol. 12 : 264-272. BUSH, G. L.
1967. The comparative cytology of the Choristidae and Nannochor- istidae (Mecoptera) . Amer. Philos. SOC., Yrbk. 1966 : 326-328. BYERS, G. W.
1961, An unusual new species of Boreus (Mecoptera: Boreidae) from Oregon. Jour. Kansas Ent. Soc. 34: 73-78. CARPENTER, F. M.
1931. Revision of the nearctic Mecoptera. Bull. Mus. Comp. Zosl., Harvard College. 72 : 205-277.
1935. New nearctic Mecoptera, with notes on other species. Psyche 42: 105-122.
1936. Descriptions and records of nearctic Mecoptera. Psyche 43: 56- 64.
1939. Records and notes of nearctic Mecoptera and Raphidoidea. Bull. Brooklyn Ent. Soc. 34: 162-166.
COLE, A. C. AND D. S. GILLESPIE.
1950. Mecoptera records from Tennessee. Jour. Tenn. Acad. Sci. 25 : 84-85.
COOPER, K. W.
1940. The genital anatomy and mating behavior of Boreus brumalis Fitch (Mecoptera). Amer. Midland Nat. 23 : 354-367. 1951. Compound sex chromosomes with anaphasic precocity in the male mecopteran, Boreus brumalis Fitch. Jour. Morph. 89: 37-58.



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Psyche
[December
CRAMPTON, G. C.
1931. The genitalia and terminal structures of the male of the archaic mecopteron, Notiothauma Reedi, compared with related Holome- tabola from the standpoint of phylogeny. Psyche 38: 1-21. DOHANIAN, S. M.
1915. Notes on the external anatomy of Boreus brumalis Fitch. Psyche 22 : 120-123.
ENDERLEIN, G.
1910. ~ber die Phylogenie und Klassifikation der Mecopteren unter Beriicksichtigung der fossilen Formen. 2001. Anz. 35: 385-399. HEPBURN, H. R.
1969. The skeleto-muscular system of Mecoptera: the head. Univ. Kansas Sci. Bull. 48 : 721-765.
KLAPALEK, F.
1901. 0 novych a malo znamych druzich palaearktickych Neuropteroid. Rozpravy Ceska Akad. Ved. a Umeni, Prague. lO(21): 1-19. 1903. Ueber neue and wenig bekannte Arten der palaarktischen Neuro- pteroiden. Bull. Internat. Resume Trav. Pres. Sci., Math., Natur. Ceska Akad. Ved. a Umeni, Prague. 7: 1-14. LESTAGE, J. A.
1940. Pour I'histoire des Boreus (St6gopt&res-Micopt6res). Ann. Soc. Roy. 2001. Belgique. 71: 1-22; 1941. Deuxieme partie. ibid. 72: 5-29; 1941. Troisieme partie. ibid. 72: 105-125. MARTYNOVA, 0. M.
1954. Skorpionnitsy (Mecoptera) fauny SSSR. I. Trudy 2001. Inst. Akad. Nauk. SSSR 15 : 54-66.
MATTHEY, R.
1950. La formule chromosomique et le type de digametie chez Bit- tacus italicus Mull. (Mecoptera). Schweiz. Gesellsch. Vererb. Forsch. Arch. Julius Klaus-Stiftung. 25: 607-611. MICKOLEIT, G.
1971a. Zur phylogenetischen und funktionellen Bedeutung der sogen- annten Notalorgane der Mecoptera (Insecta, Mecoptera). Zeit- schr. Morphol. Tiere. 69: 1-8.
1971b. Das Exoskelet von Notiothauma reedi MacLachlan, ein Beitrag zur Morphologie und Phylogenie der Mecoptera (Insecta). Zeitschr. Morphol. Tiere. 69 : 318-362.
MILLER, F. AND D. POVOLNY
1950. Boreus lokayi Kip. (Panorpata, Boreidae) v. Csr. Entomol. listy. (Folia Ent.) 13 : 91-96.
NAVAS, L.
1926. Entomologische Ergebnisse der schwedischen Kamtchatka-Ex- pedition 1920-1922. 6. Neuroptera et Mecoptera. Ark. f. Zool. 18B (2) : 1-4.
~TANES, F. Q.
1922.
Head and mouth-parts of Mecoptera. Ann. Ent. Soc. Amer. 15: 310-327.
PLIGINSKY, V.
1915 (1914). Novye vidy roda Boreus Latr. iz' Kryma (Neuroptera, Panorpidae). Russk. Entom. Obozr., Petrograd. 14: 363-367.



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19721 Cooper - B'oreus 283
POTTER, E.
1938. The internal anatomy of the order Mecoptera. Trans. Roy. Ent. Soc. London. 87: 467-501.
SLAIS, J.
1947. Anatomic du Boreus hiemalis L. (Panorpata). Tete par raport ?i l'organ digestif. Vestnik Csl. 2001. Spolecn. 11 : 271-296. STEINER, P.
1937. Beitrag zur Fortpflanzungsbiologie und Morphologie des Geni- talapparates von Boreus hiemah L. Zeitschr. Morph. Okol. Tiere. 32: 276-288.
STRUBING, H.
1950. Beitrage zur Biologic von Boreus hiemalis L. 2001. Beitr. (N.F.) 1: 51-110.
SVENSSON, S. A.
1972. Boreus Latreille, 1825 (Mecoptera). A synopsis of described species. Studies on some winter-active insects. I. Ent. Scand. 3: 26-32.
TARBINSKY, S. P.
1962. 0 nakhozhdenii lednichnika-Boreus sp. n. (Mecoptera, Boreidae) v Predgor'yakh Kirgizskogo Ala-Too. Sbornik Ent. Rabot 1: 131-136.
ULLERICH, F. H.
1961. Achiasmatische Spermatogenese bei der Skorpionsfliege Panorpa (Mecoptera) . Chromosoma 12 : 215-232.




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