Cambridge Entomological Club, 1874
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N. Banks.
Some Characters in the Perlidæ.
Psyche 54(4):266-291, 1947.

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SOME CHARACTERS IN THE PERLIDB*
BY NATHAN BANKS
Holliston, Mass.
Many years ago when I started to classify our Perlidae I neglected to study the head, antennae, and legs. In re- cent years others have emphasized the importance of genitalia and nymphal structure. Species are physiolog- ical units, genera are structural units. Reproduction is one of the most important physiological processes, so that the structure of the genital parts is of great value in species. These vary from species to species so that only in a general way can they be used for genera. If the genus is to be something different from the species, it must rest on characters which are not commonly used for species. Species reflect adaptation to environment, the recent and temporary life. Genera should rest on long- inherited structures, of little or no use to the adult, but which reflect heredity; characters common to several species.
The wings of Pteronarcys and Isoperia have many dif- ferences, but for each genus the wing is efficient ; the great number of crossveins in the former genus is not neces- sary, they represent inheritance, not use, and so become of value in the classification of genera or higher groups. Of the several structures that I have observed, the anal lobe of the fore wings1 and a vein (anal brace), arising- from the under side of the anal cell, and crossing the anal lobe seem to me to represent the course of specialization. In Pteronarcys the anal lobe is very large and the anal brace (vein that crosses it) complete and strong. In * Published with a grant from the Museum of Comparative Zoology at Harvard College.
This anal lobe is rarely shown in figures of wings. Figure I in Needham
and Claassen shows the anal area of fore wing but does not indicate the line of fold which separates the anal lobe from the rest of the wing. Many figures indicate the anal brace but often only partially or incorrectly. In the Selys part one, Klapalek on page 6 shows line of fold, in part 2, page 9, he shows the anal brace. N. & 0. on plates 13 and 16 show the anal brace in Perla and Acronewria.
266




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19471 Banks-Characters k the Perlidis 267 Figures 1 to 8 I have indicated the gradual modification until it vanishes in Peltoperla. In Pteronarcys the anal brace extends obliquely backward (recurrent). Other general with an oblique recurrent anal brace are Acro- meuria, Togoperla, Banksiana, Neoperla, Perlesta, Ptero- narcella, Harrisiola,2 Perlinella.
In Isogenus the anal brace arises closer to the base of the anal cell and extends transversely to the margin: other genera in this group are Perlod&, Clioperla, &- droverla. Those in which the anal lobe is very small and folded under and the brace vestigial are ~so~erla, Para- perla, Diploperla, some species placed in Clioperla (I am not sure of genotype). In Atoperla there is a very short basal part of anal brace arising before base of anal cell, often difficult to determine. In many cases the outer part of anal brace is weaker than basal part. In some species of Isoperla the lobe and brace are very small and weak or absent, in few distinct. In five genera the lobe and anal brace are absent : Peltoperia, Alloperia, Chloroperla, Kathroperla, and Hastaperla. In Perlesta, and some- times other forms, the anal brace is bent at the line of fold.
ANTENNB.
The antennae in Perlidae are rarely mentioned except to note the color. Walker, in the descriptions of two species (decolorata and decisa) says antennae "very minutely pubescent " and "brown, minutely pubescent. " The antennae of Pteronarcys and Acromuria have joints near the base very short and broad, some, at least immovable; in Togoperia immarginata it is similar. In Perlodes the joints near base are usually longer, but still broader than long; also in Isogenus and Hydroperla. In many of the species placed by Needham and Claassen in Perla and Clioperla the third joint is plainly longer than the fourth, the latter and several following joints being much broader than long. In this type of antennae the joints are not narrowed at base until much further out 2 Harrisiola (type Peria flavescens Walsh). I we this name for the spe-
cies (except type) that were put by Klapalek in Neophasganophora; the type species being a Perla.




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268 Psyche
[Dec.
where the joints are much longer than broad. In Ptero-
narcella the joints become longer than broad much sooner. In all of these the hairs on the antennae are very small and fine, well described as "minutely pubescent. " Often they are erect, but in some slope toward tip of joint. In Togoperla the hairs, though longer, are closely ap- pressed; in Perlinella, though short, are also appressed. In the genotype of Isoperla (bilkeata) the joints near base are as long as broad, the hairs are very short, but near the tip or beyond the middle is a longer erect hair, which I shall call a sense hair. In other species of Iso- peda it is the same, but a few (ebria) that have some- times been placed in Isoperlu do not have these sense- hairs, or at least not well-developed. In Alloperla, in Perlesta, it is the same as Isoperla. In Neoperla the hairs are longer but the sense-hairs are very prominent, also Diploperla, Paraperla, and Katkroperla, as well as Chloropeda and Hastaperla have them more or less distinct.
In Isogenus and Perlodes there are no sense-hairs or else so small as to be unnoticed, in Hydroperla some spe- cies show very tiny erect hairs toward the tip of the joint, but in others they are very small or absent, in some Diploperla they are also short, but usually distinct and near tip of joint, in Clioperlu (sirnilis, slossom~) they are readily seen. In some Acrmeuria (xanfhenes, cali- fornica) the sense-hairs are fairly distinct, beyond the short joints, in others they are obscure or absent. In the Perlince distinct, but not as prominent as in Neoperla. HEAD.
In shape the head ranges from twice as broad as long (Peltoperla) to twice as long as broad (Paraperla) ; and behind the eyes the head may extend more than twice the diameter of the eye (Kathroperla) or there may be scarcely any space at all (some Hydroperla). The head may be considered as of three areas : occipital area, the part behind eyes and ocelli; the interocular area, and the face or area in front of anterior ocellus. The occipital area is often very short in middle, where it is



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19471 BanksÌÔCharacter in the Perlida 269 separated from the interocular area by a curved line or groove, the occipital line, the ends of which curve for- ward. Just behind the eyes the head usually narrows toward the pronoturn, but in a few forms (Perlodes, Atoperla) the side at first bulges outward before it starts to narrow. In the middle there is frequently a line or groove from the occipital line to the rear of head; this median groove is present in Perlodes, Isogenus, Clioperla, some Isoperla, also in P. nona and obscura. It is short or hidden in Perla (Togo per la), Acroneuria, Harrisiola, Perlinella, Perlesta, Atoperla, Alloperia, but is present in P. luctuosa, P. modesta and P. phalerata. It is present in some very small species as P. ebria. In ~~d~o~eria it is not present, at least in sight, but sometimes a small groove on the hidden part of head or as a very fine line. The occipital line is usually quite distinct in the primi- tive forms; in the more specialized it may show behind the hind ocelli, or be entirely obliterated. In Perlodes it slopes each side and ends in front of eyes ; in Perliptzanes it slopes less and runs into lower part of eye; in Perla and Acroneuria it slopes each side just outside of boss and is stopped by a ridge. In Calliperla it slopes each side a little concavely to a point near eye where it con- nects with a line sloping below inwardly, the outer part of the M-mark. In Harrisiola and Atoperla it makes a full semicircle, the ends running into the boss; in P. modestus it makes a broader curve, also ending in the boss each side, but not a full semicircle. In Isogenus it is lower and broad, stopping near boss.
In ~erl&ella the
sides disappear a little beyond boss.
In Perlesta it con-
nects top of ocelli, beyond vestigial.
In Isoperla it some-
times shows sloping from behind the hind ocelli toward the eye, but more often it is not visible. The interocular area has the three (or two) ocelli. They form a triangle as long or longer than broad in Perla, Acroneuria, Harrisiola, and some Perlodes; in Iso- genus, Pictetia, many Clioperla and Isoperia the triangle is broader than long or at least as broad. The hind ocelli are usually situated at about the middle of the eye-space; in Perlinella and KatJiroperZa a little



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270 Psyche [DW.
behind eyes, in Alloperia further back than usual, and sometimes behind the eyes. In Isogenus, Hpdroperh, Clioperlo, and some related species the hind ocelli look more or less laterally from the head of a broad groove, there being a ridge on their inner edge as high as the ocelli; in Isoperh the hind ocelli look more upward; in Acroneuria, Alloperia, Harrisiola the hind ocelli project above the surface of the head. The distance of the hind ocelli from the eyes is, at most, only a character of species. In nearly ail Perlids there is a small swollen spot a little in front of each hind ocellus, and often further out. This is the lateral tubercle or boss. It is more or less char- acteristic in shape and position for each species. Some- times hardly further than the diameter of the ocellus, again it may be five to ten times as far away. In a few cases it is circular, but usually longer than the ocellus, and one end may be enlarged, or narrowed at outer end. hi front of the anterior oeellus is the face. There is usually a transverse groove or a depressed spot in front of the anterior ocellns. In front of this there is on each side a smooth area, or raised area, usually elongate and sloping outward; frequently they are pyri- form. These are the middle part of what Keedham calls the M-mark; however it is better called the V-mark, since the outer sides are faint or absent in most species. Some- times the two are joined together above in a broad curve. They never reach the front margin of head, and near each lateral corner may be a dark rounded spot. The shape and divergence of the V-mark are very good characters for species and species-groups.
But sometimes widely
separated genera may have the V-mark of very similar shape. I have figured a number of these, and I think they should be considered in descriptive work. In some species the head is without marks, or only dark around the ocelli, lint in many there is a pattern of dark marks, which has long been recognized of specific value. There is, of course, variation in the development of a pat- tern. When the same pattern occurs in two or more spe- cies, there are differences in structure; often in shape of the ocellar triangle or on the lower face.



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1947 I Banks-Characters in the Perlida 271 The head of Perlodes, when viewed from above, shows the side behind each eye somewhat swollen before it be- onins to slope inward. Behind the eye of Isogenus the side is, at first straight, then slopes inward ; in Hydroperla it is less distinct, since the head is shorter behind the eyes, but in ebria and gravitans( ?) the side is plainly promi- nent before sloping. The last two species, when viewed from below, show a somewhat globose area behind the eye. In many species the side behind eye is convex, but it begins to slope at the eye. Perlinella and Calliperla show the side somewhat swollen before the slope. FEMORA.
The femora have on their lower edge a row of short, fine hairs or the hairs may have above them a row of stouter bristles, or the hairs may be more numerous and spreading up on the lower part of the femur, and with bristles interspersed or in a row. Those with the row of short fine hairs are the genera Perlodes (and subgen- era), Isogenus, Hydroperla, Clioperla and Isoperia, the other genera have bristles besides the hairs. In the Perla and Chloroperla sections there are always bristles on the femora, either in a row or among the hairs. In shape the femora may be very long and slender, with nearly parallel sides, but in many of the genera with bristles among the hairs (Perlesta, Neoperla, Calliper- linae) the femora are shorter, broader, and the upper edge convex.
WINGS.
Needham and Claassen in the table to species of Perla use the position of the cubito-anal cross-vein to divide the genus in two sections, one in which this cross-vein is placed its length beyond the end of anal cell, and the other where the cross-vein is at end of cell or only a trifle be- yond. I11 general it would be better to divide whether the cross-vein is at or before end of anal cell, or plainly beyond. The amount beyond varies a bit in some species. It is plainly beyond in Isogewus, Hydroperla, Perlodes, Perlinella. In Pteronarcys this vein is much beyond end of cell, so we may consider this a primitive character.



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272 Psyche [Dec.
However in Isoperla both ways occur commonly, even sometimes in the same specimen.
One point in venation is very interesting, it is the fork- ing of the cubital vein in fore wings; in all but two sub- families the cubitus forks in such a way that the anterior branch diverges at least a little, but plainly, from the main stem while the lower branch either runs out nearly as a continuation of the stem or also diverges from the stem, so if stem were continued it would divide the fork. In two subfamilies, the Isoperlinge and the Chloroperlinae the upper branch runs out as a continuation of the stem and the lower branch definitely diverges from the stem. These two subfamilies, although having much general resemblance, differ so much in details of structure that this similarity in forking of cubitus may be the result of similar needs for strengthening the flight structure. I11 the primitive forms there are more costal cross- veins, and more branches to the radial sector than in more specialized forms, but the result is not constant, but highly variable. One point is of interest, the origin of the radial sector is usually a little beyond the end of the first anal vein, if one moves further out so does the other, Perlinella having both points further out than in Perlesta. How- ever in Perla immargkta., in Perlinella, sometimes in Acroneuria the radial sector arises before end of anal vein, and in Hydroperla and Calliperla luctuosa often opposite. The position of the cord, is usually at begin- ning of outer third, of wing; in Perlesta, Calliperla, some Acroneuria it is before end of second third, while in Per- linella, and in some Clioperla it is beyond end of second third.
In the more primitive genera the subcosta extends out to the cord or near it, and in a few, Acroneuria, Perlodes, Perla, it may extend beyond the cord. In Isogenus, Calli- perla, Harrisiola, Peltoperla, Atoperia, and KatJiroperla it reaches the cord, or nearly so. In Alloperia, Para- perla, Isoperla the subcosta ends well before the cord. Sometimes it may fade out or be very indistinct as in Nunoperla and Perlesta (some species).
The number of veins in the anal area of the hind wings



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19471 Banks-Characters kn the Perlida 273 is normally five, Paraperla has but four, Alloperia three, and Hastaperla has no anal area. In most genera the second and fifth are branched, in some Acroneuria and Perla with several branches.
In the hind wings the number of cubital cross-veins, if more than one, has been used, it varies, but is somewhat useful. The number of branches of the second and last anal veins, is also helpful, but rarely dependable. The length of the union of radial sector and medius has also been used, but too variable for a generic character. PALPI.
The palpi in Pteronarcys, Perla, Acroneuria are short and inconspicuous, the last joint of the maxillary palpi very small, the third somewhat swollen at tip ; in Isogems, the fourth joint is longer, and in some species (nona, ebria, etc.) the palpi are much longer and more slender, the fourth joint prominent and the third joint not swollen at tip, while in Alloperia the last joint is very short, scarcely noticeable.
SETB.
The setae, or tails, in the primitive forms (Perla, Acro- meuria, Peltoperla) are short and in Perla and Acroneuria much thickened at the base and taper to the slender tip. The first few joints in Perla are simple annulations, not real joints; in Acroneuria they are usually more like a joint, plainly separated by a notch on the side ; in Pelto- peria some species are similar to Perla, in others more distinct. Also in Perlodes, and Hydroperlo, some species have short setae, the basal joints scarcely separated. But in most Perlidae the set= are long, often as long as ab- domen, and with many joints much longer than broad, and with long erect bristles near tip of each joint. RESULTS.
On considering these characters in the various genera one sees that there are two primary trends or groups of primitive Perlidae, one the Acroneuria group distin- guished by the oblique anal brace arising from near third anal vein, the other the Isogenus group with an anal brace arising from the base or near base of anal cell and ex-



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274 Psyche [Dec.
tended transversely across anal area and anal lobe when present. In the specialization that occurs in both lines, there comes reduction in venation, reduction in anal lobe, and many modifications that will be of use in the par- ticular group.
Both groups have a fringe of hair on lower edge of the femora, in Perlodes and Isogenus very fine and evenly short, in Acroneuria more dense and with some stiffer and longer bristles. Both groups have the basal antenna1 joints very short, but in each line they become longer in more recent genera. In both lines the anal brace grad- ually weakens and disappears. In the Acroneuria line there is more tendency to retain and develop the bristles ; but bristles appear in some of the fairly primitive genera of the Isogenus line, such as Calliperla and Diploperla; but in the more specialized forms (Isoperla) the bristles disappear, while in the most specialized group (Chloro- ~erlinae) of the Acroneuria line the bristles are most highly developed.
With the numerous similarities, as well as the differ- ences, of structure in the group, it appears to me that in the Plecoptera there are, as Newman in 1853 stated, but three families, Pteronarcidae, Perlidas, and Nemouridae, and that under the Perlidae are eleven subfamilies, which are tabulated below.
Two courses are open in making a key to the sub- families. In both Isoperla and Alloperia the forking of the cubitus in fore-wings is such that the upper branch of cubitus goes out straight as a continuation of the stem, while the lower branch (cubital fork) plainly diverges from the stem and upper branch of cubitus. In all other Perlidae the forking of cubitus is so that the upper branch diverges from the stem, plainly at least a little. But the Isoperlinas and the Chloroperlinae (Alloperia) differ in so many ways, and, as I believe, the Isoperlinse are an off- shoot of the Isogeninae that I think it better to take out the Chloroperlinae much before the Isoperlinae. Another matter permits of two solutions. Whether to
put together all those forms which have femoral bristles, or to dispose of them according to the forking of the



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19471 Banks-Characters in the Perlida 275 cubitus; I have chosen the latter course. Bristles are
ancestral, and the occurrence of ancestral characters in descendants is not unusual.
I had started this paper in order to do identification, especially of New England Perlidae, and had hoped to rely chiefly on body characters, but find (as others have) that there is much in the wing, particularly the basal portion that is sufficiently constant for synoptic purposes. I do
not underrate male genitalia and am pleased to find that in general my results agree with the plan made by Richer in 1943.
1. Head longer than broad, eyes situate more than twice their diameter from hind margin; in the forking of cubitus the upper branch is bent a little from the stem; the third anal vein arises from the cell; no anal lobe nor anal brace ; antennse with a few joints near base short; a carina or line connecting eyes and passing just behind hind ocelli.
Kathroperlinse
If head is longer tha,n broad the eyes are not more than one and one-half diameter from the hind ............................................................................................................... margin 2 2. Small forms less than an inch; no anal lobe nor brace; anal area of hind wings with not more than four longitudinal veins ; distinct erect bristles in a row on each side of femoral groove; occipital line absent or indistinct; antennae rather long, not thickened at base, the joints monilif orm ; usually in the fore wings the third anal vein branches from ............
the second beyond the anal cell
Chloroperlinse
Anal area of hind wings with more than four longi- tudinal veins ; in the fore wings the third anal vein leaves the second at or before end of anal cell ...... 3 3. But two ocelli ................................................................................................ 4 With three ocelli .......................................................................................... 6 4. Head about twice as broad as long, partly under the overhanging pronotum, latter very broad, rounded behind, straight or slightly concave in front; legs



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276 Psyche [DW.
short; setae very short, but basal joints distinct; costal cross-veins numerous ...................... Peltoperlinae Head usually longer; pronotum usually a little con- ........................ vex in front; setae not especially short 5
5. Ocelli hardly three diameters apart; anal brace dis- tinct, extending obliquely back; femora with bristles among the hairs, no cross-veins between first and second anal veins .............................. Neoperlinae Ocelli fully four diameters apart ; anal brace faint or absent, usually a few cross-veins between first and second anal veins ............ Atoperia in Perlestinae 6. In fore-wings there is an obliquely recurrent anal brace, often arising near base of third anal vein. Third antennal joint not or little longer than the fourth joint ; cubito-anal cross-vein usually at or before end of anal cell; lower edge of femora with ....................................
fine hairs and some longer bristles
7
In fore wings the anal brace, if present, arises from nearer the base of the anal cell, and extends trans- versely across anal area and anal lobe (if present) ; often anal brace faint or absent. Third antennal joint much longer than broad, and sometimes nar- rowed in middle ; cubito-anal cros s-vein frequently beyond end of anal cell, and first anal vein may be bent at that point. The lower edge of femora has a row of fine, usually short hairs, but sometimes with longer bristles present ...................................................... 9 7. Setae fairly long, not noticeably thick near base, the joints soon as long as broad, the fourth not several times as broad as long, often with long erect bristles at tip of joints; wings with few, if any, cross-veins beyond the cord; sometimes a few cross-veins in anal area; in male the genitalia are not visible from above .......................................... Perlestinae Setae fairly short, thickened at base, the first few joints usually three or four times as broad as long, and not notched at base; usually a number of joints of antennae toward base much broader than long ...................................................................................................... 8 8. Radial sector beyond cord usually forks three times,



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Banks-Characters k the Perlidce 277
and the forks slope behind the sector, so sector fairly parallel to radius ; usually some cross-veins beyond the cord; last joint of maxillary palpi usually one-half of fourth joint. In male no proc- ess or extension of fifth dorsal segment, but on ninth ventral segment is a raised median disc; ............
subcosta often ends beyond cord
Acroneurinse
Radial sector usually with but two forks and the first parts into an anterior as well as posterior branch; no series of cross-veins beyond cord, subcosta often ends before or at cord; last joint of maxillary palpi usually not one-half of fourth joint. In male the fifth segment is usually extended, and the genital prongs wholly visible from above, no median disc on the ninth ventral segment . Perline
9. Both wings with cross-veins beyond the cord forming several irregular cells; the portion of fore wing beyond cord rarely a third of wing-length; the cubito-anal cross-vein is well beyond the end of anal cell; ocellar triangle usually as long as broad ; anal lobe distinct and quite long .................. Perlodinse Wings rarely with cross-veins beyond the cord, and not forming several cells ; portion of fore wing beyond cord usually fully one-third of wing- ............................................................................................................ length 10
10. In the forking of the cubitus in the fore wing the upper branch bends a little upward at the forking, while the lower branch also bends down a little, rarely in a curve or else does not bend at all. Radial sector often forks two or more times. Often with more than one cubital cross-vein in hind wings ; cubito-anal vein sometimes broken ............ 11 In the forking of the cubitus, it is the upper branch which continues the straight course, and the lower one diverges usually in a curve.
Radial sector
rarely forks more than once, and usually but one cubital cross-vein in the hind wings. The fourth joint of antennae is usually as long as broad, and the sense-hairs distinct; median groove rarely present ; cubito-anal vein normally not broken. Isoperlinae




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11. Femora with bristles below among the shorter hairs; anal lobe very small .... .................................. ..... Calliperlinas Femora with only the fringe of short hairs below. Isogeninae
REVIEW OF SUBFAMILIES AND GENERA
Subfamily Perlodinas
Hanson has arranged the species of this subfamily on sternal structure, and, I believe, is continuing his work, so I have not utilized sternal structure in this paper, although I have previously (1938) based a genus, Hes- peroperia, on such characters, and in fact have called attention to the value of ventral structure many years ago (1900).
The only point I would make is that I think his Diety- operygella washkgtoniana belongs to the Isogeninae, close to or the same as Hydroperla; and add a new genus, dif- fering from Perlodes in the presence of bristles on the femora.
Perliphanes gen. nov.
Type : Dietyogenus ( ?) phaleratus Smith. Head plainly broader than long ; ocellar triangle a little broader than long, hind ocelli rather small, looking later- ally, nearer to each other than to eyes; median groove reaching occipital line ; palpi rather short, third joint thick, longer than fourth, last joint one-half of fourth ; pronoturn twice as broad as long, sides parallel, as also front and rear sides; setae short, several joints beyond first broader than long. "Wings with some cross-veins connecting branches of radial sector, thus forming cells much like Perlodes. Upper branch of cubitus bends at the forking; cubito-anal vein at end of anal cell, first anal vein not here angled; anal lobe does not reach lower end of anal cell; a transverse brace across anal area from near base of anal cell. The femora have bristles among the shorter hairs.
Subfamily Calliperlinae
Cubito-anal vein at or near end of anal cell; the first



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19471 Banks-Characters in the Perlidae 279 anal vein not angled or scarcely so near base; femora have distinct bristles among the shorter hairs. 1. Pronotum in front fully twice as broad as long; head extending back of eyes for the length of an eye; ocellar triangle twice as broad as long; last joint of maxillary palpi very short; radial sector forked twice . . . . . Calliperia
Pronotum not so broad, head not extended so far back, ocellar triangle rarely twice as broad as long ............ 2 2. Maxillary palpi long and slender, the joints not broad- ened at tip ; radial sector usually forks twice. Diplo perla
Maxillary palpi shorter, one or two joints plainly thickened near tip ; radial sector of ten forked but once .................................................................................................. Occiperla Calliperla gen. nov.
Type : Perla luctuosa Bks.
The ocellar triangle is twice as broad as long; hind ocelli nearer to eyes than to each other, head curving back of eyes more than length of eye. Ocellar line angu- late behind, the median groove very short. Palpi very long and slender. Femora with distinct longer bristles besides the short fringe. Pronotum more than twice as broad as long. In fore wing the subcosta ends at or near cord, radial sector usually forks twice, cubito-anal cross- vein at end of anal cell, first anal vein not angled; in hind wing several cubital cross-veins. From base of anal cell in fore wings there is short, transverse anal brace, but no anal lobe ; antennae with several joints beyond the third much broader than long, with only minute erect hairs. Diploperla Prison
This genus is based on D. (Perla) bilobata N. & 0. This is a rather slender species in which the male shows a lobe on two ventral segments (D. duplicata is similar). The other forms are rather more robust, and the male has but one ventral lobe.
But for the pesent I will keep
them together; they are modesta, verticalis, nona, tincta, sorpta and probably misnoma, alamda, astivalis, and f ulva.




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280 Psyche [Dec.
Occiperla gen. nov.
Type : Isoperia pinta Frison.
Head broad, behind eyes it at once slopes inward; median groove well developed; ocelli small, lateral boss about as far from hind ocellus as from base of antenna; femora with bristles among the shorter hairs ; maxillary palpi moderately short, second and third joints rather thick, fifth long and slender; antennse with moderately long joints, short hair, and minute sense hair near end of joint ; pronotum much broader than long ; cubito-anal cross-vein at end of anal cell; cubitus forks so both branches are at an angle with the base ; anal brace arising a little before base of anal cell, transverse, no distinct anal lobe.
I have Occiperla phta from Huntington, B. Col., and Cowichan Lake, Vancouver; there are at least two other species in the West, both with the area in front of an- terior ocellus wholly yellowish, one with a pale longi- tudinal streak in the ocellar area, the other with ocellar area covered by a triangular dark spot.
Subfamily Isogeninse
1. The anal lobe fairly large and long, ending beyond the lower end of anal cell; in hind wings a series of cubital cross-veins ; moderately large specimens . 2 The anal lobe very small or absent, and if present it ends before the lower end of anal cell; usually smaller species .......................................................................................... 3 2. Ocellar area about as broad as long, or nearly so ; head extending some distance behind eyes and plainly a little swollen before sloping inward ; median groove nearly or quite reaching occipital line .......... Isogenus Ocellar area but little if any broader than long; head very short behind eyes, scarcely space for more than the inward slope; median groove not reaching the ..................................................................... occipital line Hydroperla.
3. Hind ocelli far from hind border of head, not under a ridge, nor a shallow groove reaching laterally, but projecting a bit above surface, anterior ocellus almost as near each eye as to anterior margin of



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1947 I Banks-Characters in the PerIid6~ 281 head; fourth antenna1 joint fully as long as broad. Megahelus
Hind ocelli not so far from hind border of head and looking out over a broad shallow groove toward eye or antennae; anterior ocellus much farther from either eye than from anterior border of head ............ 4 4. Ocellar area about twice as broad as long; median groove complete and distinct; wings moderately broad ................................................................................................... Pictetia Ocellar area plainly not twice as broad as long ; median groove sometimes complete, often not, wings not so broad ............................................................ Clioperla This genus, proposed by Needham and Claassen, is based on the Isogenus clio Newman.
Through the kind-
ness of Mr. Kimmins of the British Museum I have learned of the generic characters which places the genus in the above table. The type is from Georgia, N. & C. identified specimens from North Carolina and Indiana. I have not been able to see these specimens; Frison saw them and said3 that the specimens from Raleigh, N. C., are the same ashis confusa. Specimens sent, as para- types of confusa, by Frison to the M.C.Z. are very differ- ent from clio. In the M.C.Z. are two specimens from Woodworth Lake, Pulton Co., N. Y., which agree with color description I made of the type in 1912. The species is very close, perhaps the same, as that later described by Pictet as Perla maculata. Specimens that I have iden- tified as maculata are from North Carolina and the White Mt's., N. H.;. these are smaller than clio and the tip of abdomen - - yellow, and the subgenital plate less strongly rounded.
Pictetia gen. nov.
Type : Peda expansa Bks.
The anal lobe is very small and short, the brace scarcely distinct ; cubito-anal vein beyond tip of anal cell ; palpi long and slender; back of eyes the head slopes inward. 3 Bull. 111. Nat. Hist. Survey 22: 331.
1942.




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282 Psyche [Dec.
Isogenus Newm.
This remains as used in Needham & Claassen. Hydroperlo, Frison
This includes besides the species mentioned by Frison, varians, subvarians, hastata, guerini.
Megatielus Klap.
This is based on Isoperia bellona Bks.; it has much resemblance to typical Isoperla, but the cubitus forks as in Isogenus. There are apparently one or two more species.
Subfamily Isoperlinse
1. Hind ocelli almost twice as near eyes as to each other, ocellar triangle fully twice as broad as long; pro- notum plainly more than twice as broad as long; spines under tibiae stout and long; small black spe- cies ................................................................................................... Nanoperia Hind ocelli not so near to eyes, and ocellar triangle not so broad ................................................................................................... 2 2. Femora with distinct longer bristles among the shorter hair; cubito-anal cross-vein at end of the anal cell ............................................................................................. Perliola Femora with only the fringe of short, even hairs ...... 3 3. Hind ocelli looking laterally out over a broad, shallow groove toward eye or antenna; inner edge of ocelli scarcely above the middle surface; mostly more or less marked with brown; sometimes with extra cubi- tal cross-veins in hind wing .................................... WalsMola Hind ocelli looking more upward, inner edge usually a little above surface between ocelli: no shallow @roove, at most a small, subtriangularpit just out- &.
side of ocellus; mostly pale species; rarely with extra cross-veins in cubital area of hind wings. Isoperla
Isoperla Bks.
This I reduce to the forms similar to the genotype (bilineata) in having the cubitus forked so that the upper branch is continuous and the lower branch arising from



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it at an angle or a curve; and further as in the above table. This is closely similar to the condition in Allo- perla.
Nanoperla gen. nov.
Type : CMoroperla minuta Bks.
This was placed in Isoperla by Needham and Claassen and by Prison.
It differs from Isoperla by the more
widely separated posterior ocelli, fully twice as far apart as each from eye; by the lack of an angle at anterior cor- ners of the pronotum; in the hind wing the median cell is fully twice as long as its pedicel; the legs and setae are shorter; the medius of fore wing is often stopped by the first median cross-vein, and does not extend basally close to the radius as is common in Perlidae, this is similar to Hastaperla and Chloroperla; in the anal area the third anal vein usually branches from the second at the end of anal cell, sometimes a little beyond, rarely before it. Its dark fore wings with the costal area yellowish also sepa- rates it from Isoperla.
In both sexes the genitalia are
different from the normal Isoperla.
-
Nanoperla minuta occurs in the mid-western States, Ohio, Michigan, Illinois.
It may be, as Prison suggests,
that it is the CMoroperIa m a Walsh.
His description
is short, and the size is a little large for a dried specimen ; his unique type was from Rock Island, Illinois; Prison says that minuta occurs in the central and eastern part of Illinois.
Walshiola gen. nov.
Type : Perlinella signata Bks.
In Isoperlinae, the upper branch of cubitus continues the line of the base; cubito-anal at tip of anal cell or a little beyond ; radial sector forked once or twice ; the head back of eyes at once curves inward; median groove usually reaches the occipital line, which, beyond each ocellus fades; hind ocelli looking out laterally over a broad, shallow groove; ocellar triangle a little broader than long; a rather wide space between arms of the V-mark; fourth antenna1 joint about or fully as long as broad ; palpi long and slender, fourth joint rather longer than third.




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284 Psyche [DW.
Contains also montana Bks. and marlyma N. & 0. Perliola gen. nov.
Type : Isoperlo, 5-punctata Bks.
In this the cubitus forks as in typical Isoperia, but the femora are usually thicker (front pair) and with more numerous hairs and among them longer bristles ; the ocelli are small, the lateral boss about as large and almost as far from hind ocellus as from the base of antenna; median groove present and usually visible. The maxil- lary palpi are short, the second and third joints at least rather thick; the joints of antennae though much longer than broad are not as long as in Isoperia, and the hair is usually shorter. There are several species in the West- ern States, one with much resemblance in subgenital plate to Isoperia sobria Hag.
Subfamily Perles tinae
The three genera in our fauna are separable as follows. 1. A series of costal cross-veins (five to ten or more) before end of subcosta: head broader behind than long; no cross-veins connecting anal veins beyond cell; three ocelli; first anal ends before origin of -
radial sector ................................................................................. Perlesta First anal vein usually ends about as far out as origin of radial sector; rarely more t,han one or two costal cross-veins before end of subcosta; head sometimes as long as broad; often one or mo.re cross-veins in -
anal field ............................................................................................................ 2 2. With but two distinct ocelli; rarely with more than one or two cross-veins between first and second anal veins; front femora about equal to width of pro- notum ................................................................................................ Atoperla With three ocelli; usually three or more cross-veins connecting first and second anals ; front femora not ..............................
as long as width of pronotum
Perlinella
Subfamily Neoperlinffi
There is but one genus, Neoperla, in the U. S., others with many species occur in tropical areas and in East Asia.




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19471 Banks-Characters i~ the PerZidce 285 Subfamily Acroneurinae
The numerous species of this subfamily are placed in Acroneuria, Claassenia, Eccoptura, Beloneuria, and Hes- peroperla.
Subfamily Perlinsa
The few genera, Perla, Togoperla, Banksiama, Har- risiola, have been separated by Klapalek and others; it is necessary to distinguish the female of Harrisiola from Perla (Togoperla). In Harrisiola (both sexes) the oc- cipital line ends on the outer end of the lateral boss, in Perla the line does not bend forward so far, but runs toward the eye, well behind the boss. Perla has a tiny last joint to maxillary palpi.
Subfamily Kathroperlinse
We have in America but the typical genus which occurs in British Colombia and adjacent areas. There are other genera in Formosa and China.
Subfamily Chloroperlinse
The four genera, Paraperla, Alloperia, Chloroperla, and Hastaperla, were tabulated by Prison in 1942. Subfamily Peltoperlinas
So far all the forms have been kept in the one genus -
Peltoperla, but P. Cora will doubtless become at least a subgenus.




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Psyche
Fig. 1.
Fig. 2.
Fig. 3.
Fig. 4.
Fig. 5.
Fig. 6.
Fig. 7.
Fig. 8.
Fig. 9.
Fig. 10.
Fig. 11.
Fig. 12.
Fig. 13.
Fig. 14.
Fig. 15.
Fig. 16.
Fig. 17.
Fig. 18.
Fig. 19.
Fig. 20.
Fig. 21.
Pteronarcys spinosa, anal part of fore wing. Harrisiola flavescens, anal part of fore wing. Clioperla ebria, anal part of fore wing. Perlodes signata, anal part of fore wing. Peltoperla maria, anal part of fore wing. Acroneuria lycorias, anal part of fore wing. Isogenus frontalis, anal part of fore wing. Walshiola montana, anal part of fore wing. Kathroperla perdita, anal part of fore wing. Diploporla, anal part of fore wing.
Clioperla similis, basal part of fore wing. Banksiana lcansensis, anal part of fore wing. Pictetia expansa, anal part of fore wing. Hydroperla subvarians, basal part of fore wing. Isoperla bilineata, anal part of fore wing. Isoperla transmarina, anal part of fore wing. Diploperla bilobata, basal part of fore wing. Perliola 5-punctata, forking of cubitus. Hastaperla brevis, anal part of fore wing. Togoperla immarginata, basal part of fore wing. Atoperia fumipennis, basal part of fore wing.



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29471 Banks-Characters in the Perlidce 287 PSYCHE, 1947
VOL. 54, PLATE 20




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Psyche
Fig. 22.
Fig. 23.
Fig. 24.
Fig. 25.
Fig. 26.
Fig. 27.
Fig. 28.
Fig. 29.
Fig. 30.
Fig. 31.
Fig. 32.
Fig. 33.
Fig. 34.
Fig. 35.
Fig. 36.
Fig. 37.
Fig. 38.
Fig. 39.
Fig. 40.
Fig. 41.
Fig. 42.
Fig. 43.
Fig. 44.
Fig. 45.
Fig. 46.
Fig. 47.
Fig. 48.
Fig. 49.
Perliphanes, femur.
Calliperla, femur.
Diploperla bilobata, femur.
Togoperla immarginata, femur.
Occiperla sp., femur.
Hastaperla brevis, femur.
Alloperla imbecilla, femur and tibia.
Isoperla, femur.
Hydroperla, femur.
Isogenus, basal portion of antenna.
Perlesta, basal portion of antenna.
Isoperla bilineata, basal part of antenna. Hastaperla, basal part of antenna.
Allopcrla, basal part of antenna.
Isoperla? mohri?, basal part of antenna. Diploperla modesta, basal part of antenna. Walshiola montana, basal part of antenna. Megahelus bellona, basal part of antenna. Neoperla clymene, basal part of antenna. Perlodes signata, basal part of antenna. Acroneuria, basal part of antenna.
Clioperla maculata, basal part of antenna. Calliperla luctuosa, basal part of antenna. Perla and Togoperla, basal part of antenna. Isogenus frontalis, maxillary palpus.
Diploperia bilobata, maxillary palpus.
Perliola 5-pzinctata, maxillary palpus.
Clioperla ebria, maxillary palpus.




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1947 I Basics-CJzuracters in the PerlidfE 289 PSYCHE, 1947
VOL. 54, PLATE 21




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Psyche
Fig. 50.
Fig. 51.
Fig. 52.
Fig. 53.
Fig. 54.
Fig. 55.
Fig. 56.
Fig. 57.
Fig. 58.
Fig. 59.
Fig. 60. ,
Fig. 61.
Fig. 62.
Fig. 63.
Fig. 64.
Fig. 65.
Fig. 66.
Fig. 67.
Fig. 68.
Fig. 69.
Fig. 70.
Fig. 71.
Fig. 72.
Fig. 73.
'Fig. 74.
Fig. 75.
Fig. 76.
Fig. 77.
Fig. 78.
Fig. 79.
Fig. 80.
Togoperla immarginata, maxillary palpus. Peltoperla maria, maxillary palpus.
Isoperla bilineata, maxillary palpus.
Harrisiola fiavescens, maxillary palpus. Alloperla coloradensis, maxillary palpus. Perlesta, maxillary palpus.
Kathroperla, head.
Harrisiola, ocellar area.
Perliola 5-punctata, ocellar area.
Hydroperla hastata, ocellar area.
Isogenus f rontalis, ocellar area.
Perla capitata, ocellar area.
Walshiola marlynia, (left) Diploperla modesta, (right) eye and side of head.
Togoperla immarginata, ocellar area.
Calliperla luctuosa, ocellar area.
Perlodes (left), Isogenus (right), eye and side of head. Togoperla ifnmarginata, lower face.
Acroneuria californica, ocellus and boss. Diploperla misnoma?, ocellus and boss.
Atoperla ephyre, ocellus and boss.
Neoperla clymene, ocellus and boss.
Megahelus bellona, ocellus and boss.
Clioperla maculata, ocellus and boss.
Walshiola signata, ocellar area.
Togoperla immarginata, ocellus and boss. Walshiola montana, ocellus and boss.
Peltoperla arcuata, ocellus and boss.
Alloperla borealis, ocellus and boss.
Clioperla ebria (left), Perlinella drymo (right), eye and side of head.
Eccoptura xanthenes, ocellar area.
Acroneuria carolinensis, ocellar area.




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19471 Banks-Characters in the PerlidfE 291 PSYCHE, 1947
VOL. 54, PLATE 22




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Volume 54 table of contents